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|j Edited by
Mies GEOFFROY
fPaul MAUR/ES
UY-JACQUEMIN

jVhistoire natur:

MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE

Redacteur en chef ( Editor-in-Chief) : Jean-Lou JUSTINE
Redacteurs (Editors) : Jean-Marie BETSCH, Philippe BOUCHET, Christian ERARD & Jean-Lou JUSTINE
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Source :

Bibl lotheque Cent rale Mus6u

3 3001 00125732 7

Source : MNHN. Paris

^0 Cy,

Source : MNHN, Paris

Cover illustration:

VnllTZi? I00"*0’0 Sauss“re' 186°- 3 widesPread polydesmid millipede in the field, French Guiana: Diplopoda,
rolydesmida, Paradoxosomatidea (photograph by Michel BOULARD).

Illustration de couverture :

%^a^hD,ZZl)TpSlTUre^ I8P60' fplopode polydesmide ubiquiste photographic sur le vif en Guyane
rrangaise . Diplopoda, Polydesmida, Paradoxosomatidea (photographie de Michel BOULARD).

Acta Myriapodologica

Source : MNHN, Paris

ISBN : 2-85653-502-X
ISSN : 1243-4442

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MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE

TOME 169

ZOO LOG IE

Acta Myriapodologica

edited by

Jean-Jacques GEOFFROY*, Jean-Paul Mauries"
& Monique NGUYEN DUY- JACQUEMIN*"

* Mus6um national d’Histoire naturelle
Laboratoire d’Ecologie generate
4, avenue du Petit Chateau
F-91800 Brunoy

** Museum national d'Histoire naturelle
Laboratoire de Zoologie, Arthropodes
61, rue Buffon
F-75231 Paris Cedex 05

*** Museum national d’Histoire naturelle
Laboratoire de Zoologie, Arthropodes
61, rue Buffon
F-75231 Paris Cedex 05

EDITIONS
DU MUSEUM
PARIS

1996

Source : MNHN, Paris

CONTENTS/SOMMAIRE

Pages

Introduction . , . \ 3

Jean-Jacques GEOFFROY

List of participants and contributors . 1 9

Allocution d’ouverturc . 2 1

Jean-Marie DEMANGE

HISTORICAL MYRIAPODOLOGY

Myriapodology before and after Martin Lister's «Journey to Paris in the Year

1698» . 25

Stephen P. HOPKIN

ADVANCES IN SYSTEMATICS AND BIODIVERSITY

An approach to the revision of the East Asian millipede genus Anaulaciulus . 3 5

Zoltan KORSOS

The taxa of Rhymogona (Diplopoda: Craspedosomatidae): a ring species. Part one:
genetic analysis of the population structure . 4 5

Adolf SCHOLL & Ariane PEDROLI-CHRISTEN

Rhymogona (Diplopoda, Craspedosomatidae), un genre monospecifique. Deuxieme
partie : revision basee sur les resultats morphologiques, genetiques et faunistiques 5 3

Ariane PEDROLI-CHRISTEN & Adolf SCHOLL

Mastigophorophyllon (Verhoeff,1897) et Karpatophyllon Jawlowsky, 1928, genres
des Carpates (Chordeumatida, Diplopoda) . 6 1

Traian CEUCA

Sur la remarquable conformation des apophyses genitales males chez un
polydesmide neotropical . 67

Ionel TABACARU

Records of paradoxosomatid millipedes of India . 7 3

Kubra BANO

Systematics and biogeography of Ctenophilus Cook, 1898. A genus of centipedes
with disjunct distribution (Geophilomorpha, Schendylidae) . 7 9

Luis A. PEREIRA

Review and perspective of study on myriapodology of China . 8 1

Daqing WANG & Jean-Paul MAURIES

A taxonomic study of polydesmoid millipedes (Diplopoda) based on their
mandibular structures . 101

Kiyoshi ISH1I & Hiroshi TAMURA

Source :

ACTA MYRIAPODOLOGICA

Systematique et biogeographie des diplopodes penicillatcs des Ties Canaries et du

Cap Vert . 1 ] 3

Monique NGUYEN DUY - JACQUEMIN

Une approche des Diplopoda Penicillata de I'Amerique du Nord . 127

Bruno CONDf*

About the taxomomy of Spanish Scolopendrellidae . 137

Maria Teresa DOMINGUEZ RODRIGUEZ

Some observations on the onychophoran fauna of Tasmania . 139

Hilke RUHBERG & Robert MESIBOV

COMMUNITY STUDIES AND BIOGEOGRAPHY

Millipedes as aids for the reconstruction of glacial refugia (Myriapoda: Diplopoda) 15 1

Jorg SPELDA

On the distribution and faunogenesis of Himalayan millipedes (Diplopoda):
Preliminary results . 163

Sergei I. GOLOV ATCH & Jochen MARTENS

Etude systematique et ecologique des mvriapodes dans le Parc National de Chrea

(Atlas blideen), Algerie . 175

Ourida ABROUS-KHERBOUCHE

Etude des communautes de myriapodes (Chilopoda et Diplopoda) des forets

prepyreneennes (Huesca, Espagne) . 187

Antoni SERRA, Maria Cristina VICENTE & Eduardo MATEOS

Study of centipedes communities of three habitats in the Province of Ciudad Real .. 205

Andres GARCIA RUIZ & Francisco J. SANTIBANEZ

Svnanthropisation of the Diplopoda fauna of Poland

Wojcieh B. JEDRYCZKOWSKI

Chilopoda of urban greens in Warsaw

Jolanta WYTWER

Centipedes of Poznan town (Poland)

Malgorzata LESNIEWSKA

Contribution a la connaissance des lithobiomorphes (Chilopoda) de la region
palestinienne .

Stefan NEGREA & Zachiu MATIC (t)

Check-list, distribution and habitat in Bulgarian centipedes

Georgi RIBAROV .

Geographical distribution of diplopods in Great Britain and Ireland; possible causal
factors . . .

Anthony D. BARBER & Richard E. JONES

243

Millipedes recorded in the Grand Duchy of Luxembourg

Richard Desmond KIME

Source : MNHN. Paris

ACTA MYRIAPODOLOGICA

Some patterns in the distribution and origin of the lithobiomorph centipede fauna
of the Russian Plain (Chilopoda: Lithobiomorpha) . 265

Nadezhda T. ZALESSKAJA & Sergei I. GOLOV ATCH

The French Millipede Survey: towards a comprehensive inventory and cartography
of the Diplopoda in France . 269

Jean-Jacques GEOFFROY

Faunistique des mille-pattes de Suisse (Diplopoda) . 28 1

Ariane PEDROL1 -CHRISTEN

SYSTEMATICS AND EVOLUTION: PHYLOGENETIC RELATIONSHIPS

On myriapod / insect interrelationships . 283

Otto KRAUS & Margarete KRAUS

Morphology and evolution of circulatory organs in the Tracheata . 291

Gunther PASS

Some problems in the systematics of the order Scolopendromorpha (Chilopoda) . 293

Arkady A. SCHILEYKO

Plesiomorphic and apomorphic characters states in the class Chilopoda . 299

Carol C. PRUNESCU

A preliminary study on phylogeny and biogeography of the family Paracortinidae
(Myriapoda: Callipodida): a cladistic analysis . 307

Daqing WANG

The penis as a phylogenetic character in the millipede familv Julidae . 313

Henrik ENGHOFF

REPRODUCTIVE AND DEVELOPMENTAL TRENDS IN DIPLOPODA AND CHILOPODA

Functional morphology and evolution in genitalia of Diplopoda - Helminthomorpha

. . 327

Andreas T ADLER

Sperm competition and the evolution of millipede genitalia . 331

Mandy BARNETT & Steven R. TELFORD

Preliminary data on the anatomy of the genital systems in C rat e r o s ti gm u s
tasmanianus (Craterostigmomorpha) and Esastigmatobius longitarsis (Henicopidae,
Lithobiomorpha) (Chilopoda) . 341

Carol C. PRUNESCU, Robert MESIBOV & Keizaburo SHINOHARA

On some structural abnormalities in Dignathodon microcephalum (Lucas, 1846) and
their possible significance . 347

Francisco J. SANT1BANEZ & Andres GARCIA RUIZ

Developmental trends in the post-embryonic development of lithobiomorph

centipedes . . 35 \

Alessandro MINELLI, Enrico NEGRISOLO & Giuseppe FUSCO

Source :

ACTA MYRIAPODOLOGICA

Etude de la reproduction et du developpement post-enibryonnaire de Lithobius
pilicornis Newport, 1844 (Chilopoda, Lithobiomorpha) . 359

Antoni SERRA & Maria Carme MIQUEL

Developpement post-embryonnaire et cycle biologique de Bothropolys elongatus

Newport dans I'Est Algerien . 365

Tarek DAAS, Noureddine BOUZERNA & Michel DESCAMPS

The segmentation of the head and anterior trunk of millipedes (Diplopoda) - A

reassessment . 37 1

Wolfgang DOHLE

On periodomorphosis, iteroparity and life-cycles in males and females of
Tachypodoiulus niger (Leach) (Myriapoda, Diplopoda, Julidae) in France, Germany
and Great-Britain . 373

Francois SAHLI

PHYSIOLOGY, ECOPH YSIOLOG Y, CELL BIOLOGY

cAMP influence on brain and germinal cells RNA syntheses in Lithobius forficatus

(L.) an autoradiographic study . 3 85

Michel DESCAMPS, Catherine JAMAULT-NAVARRO & Marie-Chantal FABRE

Cadmium kinetics in Lithobius forficatus (L). during experimental contamination
and decontamination . 39 1

Sylvie GERARD, Marie-Chantal FABRE & Michel DESCAMPS

Cytochemistry of the tergite epicuticle of Glomeris marginata (Villers) (Myriapoda,
Diplopoda): Preliminary experimental results . 395

Philippe COMPERE. Stephane DEFISE & Gerhard GOFFINET

Coxal organs of chilopoda: the exocrine glands in Lithobius forficatus . 403

Jorg ROSENBERG & Hartmut GREVEN

The phenoloxidase from the hemolymph of Diplopoda . 411

Willi E. R. XYLANDER

In vitro cellular immune reactions of hemocytes against bacteria and their
differential degradation in myriapods . 42 1

Lutz NEVERMANN & Willi E. R. XYLANDER

Evidence for antibacterial activity in haemolymph of Diplopoda: preliminary

results . 43 1

Grzegorz KANIA, Jan JAROSZ, Mariola ANDREJKO & Malgorzata STEFANIAK

Supernumerary malpighian tubules in chilopods . 43 7

Carol C. PRUNESCU & Paula PRUNESCU

I he structure and possible function of the spiracles of some Scolopendridae
(Chilopoda, Scolopendromorpha) . 44 |

John G. E. LEWIS, Trevor J. HILL & Gavin E. WAKLEY

Source : MNHN, Paris

ACTA MYRJAPODOLOGICA

1 I

Population metabolism of millipedes at two altitudinal zones in the Central Alps
(Tirol, Austria) . 45]

Erwin MEYER, Peter MARSONER & Elisabeth FISCHER

Variation de la teneur en eau en fonction de la taille corporelle dans une population

du diplopode Polyzonium germanicum . 461

Guy VANNIER & Jean-Frant^ois DAVID

1 he respiratory response to changing temperature in millipedes belonging to the

genus Glomeris Latreille, 1802 . 473

Vladimir SUSTR

Submersion tolerance of some diplopod species . 477

Klaus Peter ZULKA

Eversible vesicles in Myriapoda . 433

Frantisek WEYDA

POPULATION BIOLOGY, SOIL ECOLOGY AND BEHAVIOUR

Etude comparative des techniques d'echantillonnage des macroarthropodes
saprophages (Isopoda and Diplopoda) . 48 5

Etienne BRANQU ART & Charles CASPAR

Experimental behaviour of a tropical invertebrate: Epiperipatus biolleyi
(Onychophora: Peripatidae) . ’ 493

Julian MONGE-NAJERA, Zaidett BARRIENTOS & Franklin AGUILAR

Scolopendra morsitans Linnaeus, 1758: a characteristic prey of the African carpet
viper Echis ocellatus Stemmier, 1970 . 495

Pascal REVAULT

The life cycle of Cylindroiulus latestriatus (Curtis, 1845) . 501

Karin VOIGTLANDER

Life-cycle of the millipede Melogona voigti (Verhoeff, 1899) from a suburban
forest in South Bohemia . 509

Karel TAJOVSKY

Compared life-cycles and reproductive strategies in local populations of Rossiulus
kessleri (Lohmander) (Julidae, Diplopoda) from isolated habitats . 515

Bella R. STRIGANOVA

Survival strategy of the terricolous millipede Cutervodesmus adisi Golovatch
(Fuhrmannodesmidae, Polydesmida) in a blackwater inundation forest of Central
Amazonia (Brazil) in response to the flood pulse . 5 23

Joachim ADIS, Sergei I. GOLOVATCH & Susanne HAMANN

Cycles d'activite compares de populations de diplopodes edaphiques dans un
ecosysteme forestier tempere . 533

Jean-Jacques GEOFFROY & Marie-Louise CELERIER

Source :

ACTA MYRIAPODOLOGICA

Traces de 1'activite de diplopodes dans des sols et des sediments karstiques du Maroc
atlantique . . . 555

Colette JEANSON, Hsain EL AISSAOUI & Jean-Pierre ADOLPHE

Feeding rates and nutrient assimilation in the millipede Jonespeltis splendidus
(Diplopoda, Paradoxosoniatidae) . 561

Kubra BANO

Sexual selection in savanna millipedes: products, patterns and processes . 5 65

Steven R. TELFORD & John Mark DANGERF1ELD

Trophic preferences of three soil macroarthropods (preliminary study) . 57 7

Jorge P. CANCELA DA FONSECA & Leila MEZIANE

Ecology and behaviour of Xanthodesmus physkon (Attems 1898), an aggregating
paradoxosomatid from tropical West Africa . 5 85

Dieter MAHSBERG

Deplacements en masse dans le sud-est de la France chez Ommatoiulus sa bill os us
(Myriapoda, Diplopoda, Julidae) avec invasions d'habitations . 5 87

Francois SAHLI

COMMUNITIES IN ECOSYSTEMS

Distribution patterns and qualitative composition of the centipede fauna in forestal
habitats of mainland Greece . 5 99

Marzio ZAPPAROLI

On abundance, phenology and natural history of Symphyla from a mixedwater
inundation forest in Central Amazonia, Brazil . 607

Joachim ADIS, Jose Wellington DE MORAIS & Ulf SCHELLER

The ecology of savanna millipedes in Southern Africa . 6 1 7

John Mark DANGERFIELD & Steven R. TELFORD

The diplopod community of a mediterranean oak forest in Southern France:

ecological and evolutionary interest . 62 7

Jean-Fran^ois DAVID

Centipedes (Chilopoda) of some forest communities in Slovenia . 63 5

Ivan KOS

Changes in the millipede (Diplopoda) community during secondary succession from
a wheat field to a beech wood on limestone . 647

Stefan SCHEU

Centipedes from Italian agroecosystems and their possible value as pest control

a8ents . 657

Marzio ZAPPAROLI

AUTHOR INDEX / INDEX DES AUTEURS . 663

SYSTEMATIC INDEX / INDEX SYSTEMATIQUE . 665

Source

Introduction

Jean-Jacques GEOFFROY

CNRS, Museum National d'Histoire Naturelle, Laboratoire d’Ecologie Generate. F-91800 Brunoy, France

Some twenty-seven years ago, a group of zoologists and biologists working on Myriapoda
met in Paris (France) for the First time. 1968 was the birth year of international congresses of
myriapodology and time when the Centre International de Myriapodologie took form. The
creation of the CIM was the work of three people : J. M. DEMANGE (Paris), J. P. MAURIES
(Paris) and O. KRAUS (Hamburg). Four years later, at Manchester, U.K. (1972), J. G.
Blower joined the initial trinity as the fourth CIM Father, when organizing the Second
International Congress of Myriapodology. These four men are the musketeers of the CIM. Ever
since, a new congress has been organized through out the world every three years : Hamburg,
Germany, 1975 (O. Kraus), Gargagno, Italy, 1978 (M. CAMATINI), Radford, USA, 1981 (R.
L. HOFFMAN), Amsterdam, The Netherlands, 1984 (C. A. W. JEEKEL), Vittorio Veneto, Italy,
1987 (A. MlNELLI) and Innsbruck, Austria, 1990 (E. MEYER & K. THALER).

Some five years ago, according to previous formal sessions of the CIM, it was suggested
that France, as first host-country and location of the permanent secretariat, should host the Ninth
International Congress of Myriapodology in 1993. This date appeared to be a very significant
one, as it saw the 25th anniversary of the CIM (1968-1993) and the bicentenary of the Museum
National d'Histoire Naturelle de Paris (1793-1993). We fully agreed with this idea, and decided
to prepare a proposal wich was submitted to the plenary session of the CIM held in Innsbruck,
Austria in July 1990. Paris was obviously the most appropriate place in France, due to the
souvenir of famous zoologists and myriapodologists, the availibility of convenient facilities, the
assistance of laboratories in the Museum National d'Histoire Naturelle (MNHN), the Centre
National de la Recherche Scientifique (CNRS) and the Universite Pierre et Marie Curie (UPMC),
the touristic interest of the city and the surroundings of the wide Fontainebleau forest...

In accordance with the discussions at Innsbruck, we decided to fix the period of the
Congress to the end of July ; we also considered the possibility to organize some visits in
National Galleries, an exhibition devoted to the activity of scientists durind the French
Revolution, and a one-day excursion in various cultural and natural sites in the Fontainebleau
area. Besides, we kept in mind to leave the scope of the Congress widely open to several topics,
in order to contribute, by lectures and posters, to an up-to-date and more or less comprehensive
knowledge about the biology of Diplopoda, Pauropoda, Symphyla, Chilopoda and - as is
traditional - Onychophora. Our favorite creatures would appear as models for fundamental and
applied biology and the contents of this volume plan to summarize this reality in 8 chapters :

Geoffroy, J. J.. 1996. — Introduction. In: Geoffroy, J.-J., Mauri£s, J.-P. & Nguyen Duy - Jacquemin, M.,
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 13-17. Paris ISBN : 2-85653-502-X.

Source

JEAN-JACQUES GEOFFROY

Historical Myriapodology; Advances in Systematics and Biodiversity; Systematics and
Evolution: Phylogenetic Relationships; Community Studies and Biogeography; Reproductive
Developmental Trends: Physiology, Ecophysiology and Cell Biology; Population Biology, Soil
Ecology and Behaviour; Communities in Ecosystems.

Fig. I. — A logo for the CIM (Centre International de Myriapodologie : Secretariat Permanent. MNHN Paris, 61 rue
Buffon F-7523I Paris Cedex 05, France). Conceiving : Geoffroy. MAURlfes & Nguyen Duy - Jacquemin. Drawn
by Jacques Rebi£re (mnhn).

Following the decision of the Centre International de Myriapodologie, an organization
committee was soon established in France and through out the world.

Organizers: J. J. GEOFFROY (CNRS, MNHN, Brunoy), J. P. MAURIES (MNHN, Paris), M.
Nguyen Duy - Jacquemin (cnrs, mnhn, Paris), M. L. Celerier (upmc, Paris).

President of the Congress: J. M. DEMANGE (MNHN, Paris).

Organizing Committee: J. F. David (CNRS, MNHN, Brunoy), M. DESCAMPS (USTL,
Lille I), C. JAMAULT-NAVARRO (Universite, Amiens), F. SAHLI (MNHN, Paris).

International Scientific Committee: J. ADIS (Plon, Germany), C.S. CRAWFORD
(Albuquerque, USA), W. DOHLE (Berlin. Germany), W. DUNGER (Gorlitz, Germany), H.
ENGHOFF (Copenhagen, Denmark), S. I. GOLOVATCH (Moscow, Russia), W. B.
JEDRYCZKOWSKI (Warsawa, Poland), C. A. W. JEEKEL (Amsterdam, The Netherlands), P. M.
JOHNS (Christchurch, New-Zealand), O. KRAUS (Hamburg, Germany), J. G. E. LEWIS
(Taunton, Somerset, U.K.). B. MEIDELL (Bergen, Norway), A. MlNELLI (Padova, Italy), E.
Meyer (Innsbruck, Austria), H. RUHBERG (Hamburg, Germany), U. SCHELLER (Jarpas,
Sweden), W. A. SHEAR (Hampden-Sydney, USA), R. M. SHELLEY (Raleigh, USA), B. R.
STRIGANOVA (Moscow, Russia) and M. R. WARBURG (Haifa, Israel).

Source

INTRODUCTION

The Ninth International Congress of Myriapodology was held from 26-31, July 1993 at
the University Pierre et Marie Curie, Paris VI and at the Museum National d'Histoire Naturelle
de Pans. A total of 129 members from 37 countries contributed or attended the Congress
Sessions were conducted over a five-day period with a mid-excursion to the Fontainebleau
i orest, castle and locky sites. Numerous attendees visited the future Evolutionary Gallery and
Micro-Zoo at the National Museum. 3 3

Fic. 2. Intemalional participation to the 9th International Congress of Myriapodology (Paris. France. July. 1993).

There were 97 scientific contributions, by lectures or posters. 14 sessions topics
represented the different themes. Seventy-nine papers were accepted for publication, some of
* em as short-papers or abstracts. This volume is based mainly on communications delivered
during the 9th Congress but its main aim is to produce a recent up-to-date review of the biology
(s.L) of millipedes, centipedes, symphylids, pauropods, and onychophorans. It is meant for
students ol terrestrial arthropods and soil biology; as well as for researchers, biologists,
zoologists, working in fields such as phylogeny, systematics, ecology, cell biology and others.

JEAN-JACQUES GEOFFROY

ACKNOWLEDGEMENTS

We gratefully acknowledge the financial support and practical assistance to the Congress
provided by the different ministries, scientific institutions, societies and other bodies:

- Ministere des Affaires Etrangeres (DDCSTE)

- Ministere de l'Enseignement Superieur et de la Recherche (ACCES)

- Service le lTnformation et de la Communication (UPMC, Paris VI)

- UFR Sciences de la Vie (UPMC. Paris VI)

- Parc Zoologique de Paris, Menagerie du Jardin des Plantes (MNHN)

- Cellule de Prefiguration de la Galerie de l'Evolution (MNHN)

- Service des Relations Exterieures et Presse (MNHN)

- Service des Cultures (MNHN)

- Societe de Biogeographie

- Societe de Biospeoiogie

- Societe Frangaise d'Ecologie

- Calypso Log

- CAES du CNRS

- Office National des Forets (ONF, Centre de Fontainebleau)

- RATP

- Societe AGISSON

Special thanks to their efficient help to Esther CLEMENT, Corinne GENOT, Gilles
HORTAULT, Mark JUDSON. Chantal LARROCHE, Marie-Anne MONTANE, Dominique MORO,
Anne Roussel-Versini, Michele Bertoncini.

Many thanks to the CAES/CNRS for the exhibition on “French Scientists and the
Revolution-’, and to the SOCIETE AGISSON for manufacturing the Tee-shirts.

Congratulations and friendly thanks to Jacques REBIERE for both the pleasant and serious
drawings, and to Valerie CHANSIGAUD for computer assistance.

A special mention must be adressed to the Laboratoire de Biologie & Physiologie des
Organismes (UPMC. Prof. Y. TURQUIER), the Laboratoire d'Ecologie Generate (MNHN. Prof. P.
BLANDIN) and the Laboratoire de Zoologie/Arthropodes (MNHN. Prof. Y. COINEAU) who
provided logistic support during the Congress organization and during the busy exciting period
of preparing the present volume.

Myriapodology moves on! We sincerely hope that myriapodologists (.?./.) will meet again
numerous and in good spirits in Copenhagen in 1996. for a new fascinating rendez-vous with
myriapod biology.

Paris, July 1995
Jean -Jacques GEOFFROY

Source :

INTRODUCTION

Fig. 3. During the Ninth International Congress of Myriapodology, Paris, July. 1993 : I. F. Minelli: 2. F. Minelli;
3. S. Negrea; 4. C.-C. Prunescu; 5. T. Ceuca; 6. A. Serra; 7. J.-M. Demange; 8. J.-J. Geoffroy; 9. VI. C. Vicente;
10. R. Bouzerna; II. M.C. Miquel; 12. K. Tajovsky; 13. B. Striganova; 14. J. Wytwer; 15. E. Branquart;
16. M. Kos; 17. P. Johns; 18. O. Abrous-Kherbouche; 19. A. Schileyko; 20. M.-L. Celerier; 21. M. Warburg;

22. Z. Korsos; 23. G. Kania; 24. H. Read; 25. M. Kraus; 26. W. Dunger; 27. K. Voigtliindcr; 28. N. Bouzerna;

29. D. Mashberg; 30. J. Spelda; 31. M. Lesniewska; 32. Z. Korsos; 33. S.R. Telford; 34. M.P. Minelli;

35 VI. Barnett; 36. A. Minelli; 37. S.P. Hopkin; 38. G. Ribarov; 39. E. Krabbe; 40. VI. Nguyen Duy-Jacquemin;

41. J.-F. David; 42. R.D. Kime; 43. J.W. de Vlorais; 44. E. Christian; 45. B. Meidell; 46. F.J. Santibanez;
47. F. Weyda; 48. A. Pedroli-Christen; 49. V. Sustr; 50. G. Pass; 51. B. Condc; 52. H. Borueki; 53. H. Friind;
54. K. Ishii; 55. O. Kraus; 56. I. Kos; 57. U. Scheller; 58. J. Adis; 59. A. Mette; 60. H. Ruhberg; 61. E. Robson;
62. P Reveillet; 63. E. Meyer; 64. M. Di Giovanni; 65. M. Zapparoli; 66. L. Nevermann; 67. J.G.E. Lewis;
68. E.H. Eason; 69. S.l. Golovatch; 70. J.-P. Mauries; 71. J. Rosenberg; 72. B. Rosenberg; 73. W. Dohle;
74. K.P. Zulka; 75. A. Tadler; 76. R E. Jones; 77. H. Enghoff; 78. M. Descamps; 79. A.D. Barber;
80. P. Compere; 81. W. Jedryczkowski; 82. J.P. Cancela da Fonseca; 83. K. Jedryczkowski; 84. G. Andersson.

Source : MNHN, Paris

Albania
Qirjo M.

Algeria

Abrous-Kerbouche O.
Bouzerna N.

DaasT.

Argentina
Pereira L. A.

Australia
Mesibov R.

Austria
Christian E.

Fischer E.

Marsoner P.

Meyer E.

Pass G.

Tadler A.

Zulka K. P.

Belgium
Branquart E.
Compere P.

DefiseS.

Gaspar C.

GoffinetG.

KimeR. D.

BlELORUS

Tarasevich Y.

Botswana
DangerfieldJ. M.
Kaunda S.K.

Brazil

De Morais J. W.

Bulgaria

Ribarov G.

China
Wang D.

Costa Rica
Monge-Najera J.
Barrientos Z.

Aguilar F.

List of the Participants and Contributors

Cuba

Ruhberg H.

Perez- Asso A. R.

SCHEU S.

Czech Republic

Spelda J.

V OIGTLANDER K.

Sustr V.

Xylander W. E. R.

Tajovsky K.

Weyda F.

Hungary

Denmark

Enghoff H.

KORSOS Z.

India

Egypt

Bano K.

Pandey M. K.

Ghabbour S.

Tripathi S. P.

France

Israel

Adolphe J. P.

Warburg M."

Cancela Da Fonseca J. P.
Celerier M. L.

Italy

Cond£ B.

Di Giovanni M.

David J. F.

Minelli A.

Demange J. M.

Negrisolo E.

Descamps M.

Zapparoli M.

El Aissaoui H.

Fusco G.

Fabre M. C.

Geoffroy J. J.

Ivory Coast

Gerard S.

Bourdanne Kadebe D.

Jamault-Navarro C.
Jeanson C.

Japan

Mauries J. P.

ISHII K.

Meziane L.

Shinohara K.

Nguyen Duy - Jacquemin M.

Tamura H.

Revault P.

Reveillet P.

New-Zealand

Sahli F.

Johns P.

Vannier G.

Germany

Norway

MeidellB.

Adis J.

Borucki H.

Poland

DohleW.

Andrejko M.

DUNGER W.

Jarosz J.

Emmerling C.

Jedryczkowski W. B.

FrOnd H.

Kania G.

GrevenH.

Lesniewska M.

Ham ann S.

Stefaniak M.

Krabbe E.

Wytwer J.

Kraus M.

Kraus 0.

Romania

Mahsberg D.

Ceuca T.

Martens J.

Matic Z.

Nevermann L.

Negrea S.

Rosenberg J.

Prunescu C. C.

Prunescu P.

Tabacaru I.

Russia

Golov atch S. I.
Mikhaijova E. V.
SCHILEYKO A. A.

Striganova B. R.
Zalesskaja N. T.

Slovenia

Kos I.

South Africa
Barnett M.

TelfordS. R.

Spain

Dominguez -Rodriguez M. T.
Garcia Ruiz A.

Mateos E.

MiquelM. C.

SantibanezF. J.

Serra A.

Vicente M. C.

Sweden

AnderssonG.

SchellerU.

Switzerland
Pedroli-Christen A.
Scholl A.

Ukraine
Chornyi N. G.

United Kingdom
Barber A. D.

Eason E.

HillT.J.

Hopkin S. P.

Jones R. e.

Lewis J. G. E.

Read H.

Robson E.

Wakley G. E.

USA

Crawford C. S.

Edgar G. A.

Mulvey M.

Source : MNHN. Paris

Allocution d’ouverture

Jean- Marie DEMANGE

President du IXeme Congres International de Myriapodologie (MNHN - Zoologie, Arthropodes - Paris)

M. le President, M. le Directeur, mes chers Collegues, Mesdames, Messieurs.

Le plaisir que nous eprouvons, celui de se trouver tous reunis, nous le devons a ceux qui
nous ont aides, soit materiellement, soit financierement. Je tieos a remercier tout particulierement
tous ceux grace a qui nous sommes la aujourd'hui : tout d'abord l'Universite Pierre et Marie
Curie en la personne du Professeur J. LEMERLE, Vice President, son representant ; non
seulement pour son aide financiere mais aussi pour tous les moyens mis a notre disposition, ne
serait-ce que cet amphitheatre et tout le materiel y afferent. Je remercie egalement le Professeur J.
GUERDOUX, directeur de l'UFR Sciences de la Vie pour sa genereuse participation financiere.
C'est grace aux efforts du Professeur Y. TURQUIER notamment et de notre collegue Madame
Marie-Louise CELERIER que furent elaborees les affiches du Congres, le fascicule des resumes
des communications, les panneaux, etc. M. L. CELERIER a assure, en outre, la liaison entre
notre Laboratoire du Museum et l'Universite. Le Museum National d'Histoire Naturelle,
actuellement en pleine renovation (Grande Galerie de l’Evolution, Amphitheatres,
Bibliotheque...) a mis a notre disposition la logistique de deux de ses laboratoires et accueille
plusieurs evenements de notre congres. De plus, une aide financiere assez importante a ete
debloquee a notre profit pour que tout se passe bien. Je remercie Monsieur le Professeur Jacques
FABRIES, Directeur du Museum , pour cette genereuse contribution.

Vous avez bien voulu egalement accepter. Monsieur le Directeur, que notre reunion
amicale d'accueil se tienne a la Rotonde de la Menagerie du Jardin des Plantes, ce qui est
exceptionnel. A cette occasion, j'adresse au Professeur Jean-Jacques PETTER, Directeur de la
Menagerie du Jardin des Plantes et de la Conservation des especes animales (Zoo), ma plus vive
reconnaissance. La Rotonde est un monument magnifique, recemment restaure, construit de
1804 a 1812, en forme de Croix de la Legion d'Honneur dont l'ordre etait cree par Napoleon ler
deux ans auparavant. Elle abrita, pendant pres de vingt ans, la celebre girafe offerte au Roi
Charles X par Mehemet Ali, Pacha d'Egypte. Cette Rotonde etait au depart destinee a presenter
les "animaux feroces" mais ce furent plutot les “animaux paisibles” qui l'occuperent. Elle est
aujourd'hui le temple des arthropodes, des microarthropodes notamment. dont l'exposition et les
appareils d'observation uniques qui la composent ont ete crees par le Professeur Yves COINEAU

Demange. J.-M., 1996. — Allocution d'ouvcrture. In: Geoffroy. J.-J„ Mauri£s, J.-P. & Nguyen Duy -
JACQUEM1N. M.. (eds), Acta Myriapodologica. Mem. Mus. nain. Hist. nat.. 169 : 21-23. Paris ISBN : 2-85653-502-X.

JEAN-MARIF. DEMANGE

et experimentes au Parc Zoologique du Bois de Vincennes... les gros animaux et les tout petits...
“de la Puce a l'Elephanf ’...

Nos collegues japonais auront, sans doute, ete surpris et intrigues de trouver au milieu du
petit temple qui forme la partie centrale du batiment, une magnifique sculpture representant une
japonaise. Ne voyez aucun message dans la presence de cette statue ; elle est superbe, en albatre

et comme un temple a toujours sa statue . Cette statue a une histoire et si cette japonaise parait

un peu fantaisiste aux yeux de nos collegues, qu'ils voient en cette oeuvre l'inspiration d'un
artiste. C'est une figure allegorique representant le Japon. Tun des Pays participants de
l'Exposition Universelle de 1878. Elle est l'ceuvre d'un artiste frangais Eugene AIZELIN et ceci
explique cela. Je remercie done egalement. a un double titre, le Professeur Yves CoiNEAU : en
tant que responsable scientifique, createur de cette exposition et en tant que Directeur du
Laboratoire de Zoologie/Arthropodes du Museum ; il a bien voulu nous apporter, en outre, une
aide precieuse et efficace pour la bonne realisation de notre Congres. Notre collegue le
Professeur Patrick BLANDIN a largement pris en charge, au Laboratoire d’Ecologie Generate,
toutes sortes de frais divers tres lourds pour notre petite communaute. N'oublions pas, enfin, le
Ministere des Affaires etrangeres, qui apporta sa contribution par son bureau des Congres, en
debloquant une forte somme et le Ministere de la Recherche qui y ajouta sa contribution
financiere. Le CAES du CNRS. quant a lui, nous a confie les panneaux de son exposition traitant
“des Savants et la Revolution ” qui sont presentes dans le hall. Tout cela fait done que nous
sommes la aujourd'hui, mais l'argent n'est pas tout. N'oublions pas mes collegues du Comite
d'Organisation qui se sont depenses sans compter, je dirais meme devoues pour aplanir toutes
les difficultes et Dieu sait qu'il y en eut ! et organiser de main de maitre cette rencontre.

Notre infatigable et dynamique collegue Jean-Jacques GEOFFROY fut un moteur tres
efficace epaule en cela par Monique NGUYEN DUY - JACQUEMIN et Jean-Paul MAURIES.
Beaucoup d'autres nous ont aide et beaucoup nous aiderons encore pendant nos reunions, je
veux parler de toutes les personnes de 1'Universite et du Museum, de tous les niveaux. Merci,
merci encore a tous et pour tout.

Je voudrais dire combien je suis sensible a l'honneur qui m'est fait d'avoir ete nomme a la
Presidence de ce Congres. Je veux voir dans cette distinction un temoignage de sympathie,
d'amitie et j’en suis d'autant plus touche. Ce signe de l'amitie n'est il pas exprime d'ailleurs
parfaitement dans le logo qui marque notre Congres? C'est meme le signe de bunion qui a
toujours ete celui du CIM (Centre International de Myriapodologie), que Ton peut reconnaitre
dans ce couple de Mille-Pattes enlaces qui reproduisent la tour Eiffel. Il y a 25 ans, nous nous
rencontrions a Paris pour la premiere fois dans le cadre du ler Congres international de
Myriapodologie. C'est en son sein que fut cree le CIM , par trois d'entre nous: le Professeur Otto
Kraus, ici present, alors a Francfort, Jean-Paul MAURIES du Laboratoire des Arthropodes et
moi-meme. En ce quart de siecle d'existence, plusieurs de nos collegues, presents a Paris en
1968, nous ont quittes : Mrs. Nell Bevel CAUSEY, Ulrich HAACKER qui anima si parfaitement et
si brillamment nos seances en se proposant spontanement "traducteur simultane" par sa
connaissance des langues et sa valeur scientifique, le Professeur Robert JOLY, des Universites
de Lille et d'Amiens, specialiste de l'endocrinologie de Lithobius forficatus et des phenomenes
de mue, le Dr. Karl STRASSF.R fun de nos doyens avec le Reverend Cannon BRADE-BIRKS et
enfin, plus recemment, le Professeur Max VACHON, ancien directeur du Laboratoire de Zoologie
des Arthropodes. J'ai, par ailleurs, voulu evoquer la memoire de tous les myriapodologistes
disparus, dans une sorte de preface au recueil des resumes des communications car je souhaite
qu'ils soient presents a nos cotes. Ne les oublions pas en un jour comme celui-ci. [A l'heure ou
paraissent ces lignes, un autre de nos fideles nous a quitte, en 1994 : Colin Peter FAIRHURST],
Les uns nous quittent mais d'autres nous rejoignent et nous voyons aujourd'hui beaucoup
dentre eux, de la nouvelle generation de chercheurs, qui viennent enrichir notre communaute.

Parti de Paris, notre mouvement revient a Paris apres avoir parcouru le Monde;
successivement Manchester, Hambourg. Gargnano, Radford, Amsterdam, Vittorio Veneto et

Source :

ALLOCUTION D'OUVERTURE

Innsbruck entin, en 1990. Depuis 1968, le nombre des Pays participants ne cesse d'augmenter:
de 13 a Paris, il y en eut 25 a Innsbruck et aujourd'hui 37. Je suis heureux de souhaiter
particulierement la bienvenue a ces nouveaux representants.

Cette annee 1993 revet une solennite toute particuliere, non seulcment parce que c'est un
retour aux sources mais surtout par deux symboles : - le bicentenaire de la creation de notre
grande et venerable Maison qui de Jardin du Roi devient Museum d'Histoire Naturelle par decret
de la Convention du 10 juin 1793 ; - la venue officielle au Museum, a cette meme date, de Jean
Baptiste Pierre Antoine de Monet, chevalier de LAMARCK, l'un des trois artisans de la creation
de notre Etablissement avec BUFFON et DAUBENTON. Ce dernier symbole revet, pour nous
chercheurs du Laboratoire de Zoologie/Arthropodes, une importance particuliere en ce que notre
laboratoirc est l'un des descendants directs de la chaire confiee a Lamarck. Elle fut divisee, en
1 830, en deux chaires : “Crustaces et Insectes" confiee a Pierre Andre LATREILLE et “ Annelides ,
Mollusques et Zoophytes ” confiee a Henri Ducrotay de BLAINVILLE. Une troisieme chaire en fut
detachee en 1917: " Vers et Crustaces", de laquelle est issue, plus recemment (1960) celle de
"Zoologie/Arthropodes" . Le Laboratoire de “ Zoologie/Arthropodes ” conserve les collections
d'origine de crustaces, arachnides, myriapodes, augmentees de collections celebres dont celles
de Henri Wilfrid BROLEMANN et Henri RlBAUT, pour ne parler que des myriapodes. En ce qui
concerne Henri-Wilfrid BROLEMANN, qu’il me soit permis d'evoquer brievement sa memoire
car il est incontestablement le pere de la Myriapodologie en France. Ne le 10 juillet 1860. il est
decede le 31 juillet 1933. Appartenant a une famille de grands industriels et de banquiers,
banquier lui-meme, il s'interesse tres tot aux myriapodes et finit par abandonner son metier
d'origine pour se consacrer totalement a l’etude des “Mille-pattes”. Specialiste de reputation
internationale incontestee, il publie 160 travaux plus particulierement consacres aux diplopodes.
C'etait un homme de grande culture, un erudit... Il est l'auteur d'une theorie evolutive, pour les
myriapodes, basee sur un “principe de contraction” ou il considere une evolution du groupe vers
une reduction du nombre des segments par arret de developpement. Ce principe de contraction
s'oppose a un “principe d'elongation” soutenu, a l'inverse, par Karl Wilhelm VERHOEFF,
specialiste allemand. Mais le temps des fondateurs est revolu, du point de vue de notre speciality,
la myriapodologie, nous nous retrouvons aujourd'hui tres nombreux et Ton a pu apprecier, au
cours de nos reunions successives la montee de la jeune generation. On peut done envisager
l’avenir avec confiance et souligner que la systematique est toujours a l'ordre du jour, bien
vivante et dynamique a une epoque ou le concept de biodiversite s'impose. Le programme de ces
journees est charge, tres charge, abordant tous les sujets de la molecule a la pure systematique ;
mettons nous vite au travail et pour cela je declare ouvert le 9eme Congres International de
Myriapodologie de Paris.

Source : MNHN, Paris

Myriapodology before and after Martin Lister’s
« Journey to Paris in the Year 1698»

Stephen P. HOPK1N

School of Animal and Microbial Sciences, University of Reading, P.O. Box 228, Reading, RG6 2AJ, U. K

ABSTRACT

The most famous publication of Martin Lister (1638-1712) was his account of his «Journey to Paris in the year
J698». The book is well-known for its detailed descriptions of everyday life in France at the end of the 17th century. Of
interest to myriapodologists, however, are the striking illustrations by Father Charles Plumier of two myriapods from
Brazil, a millipede lulus Americanus and a centipede “ Scolopendra Americana" . Indeed, myriapods have featured
prominently in zoological literature since the time of Aristotle 384-322 BC. The development of myriapodology has
mirrored the scientific revolution since the Renaissance. This paper gives an overview of the passage from folklore and
whimsy, through the seminal observations of Leeuwenhoek, the “compendia’* of 18th century zoologists including
Linnaeus, culminating with the flowering of scientific myriapodology in the 19th century.

RESUME

La myriapodologie avant et apres le «Voyage a Paris en Pan I698» dc Martin Lister.

La plus celebre publication de Martin Lister (1638-1712) fut sa relation de son « Voyage a Paris en Van I698». Le
livre est surtout connu pour sa description detaillee de la vie quotidienne des fran^ais a la fin du I7eme sieele. II presente
cependant un interet pour les myriapodologistes, a travers les illustrations saisissantes, dues au Pere Charles Plumier, de
deux myriapodes du Bresil. un diplopode. “ lulus Americanus" et un chilopode . "Scolopendra Americana". Les myriapodes
onl vraiment 6lc eminemment represents dans la literature zoologique depuis I'epoque d’Aristote (384-322 BC). Le
developpement ulterieur de la myriapodologie a reflete la revolution scientifique qui s’est operee depuis la Renaissance.
Ce travail se propose de passer en revue cette evolution qui, depuis le folklore et la fantaisie. h travers les observations de
Leeuwenhoek, grace aux precis et traites des zoologistes du 18eme sieele - parmi lesquels figure Linn£ -, a abouti au
developpement considerable de la myriapodologie qui a fleuri au 19eme sieele.

INTRODUCTION

"It is a noble employment to rescue from oblivion
those who deserve to be remembered'’

Pliny the Younger, Letters V.

Centipedes and millipedes are among the most prominent of terrestrial invertebrates. It
should not surprise us to find numerous references to myriapods throughout the literature of the
past. However, the modern approach to research emphasises topicality. Work rapidly becomes
“out of date". Few scientists have the time to study the books and papers of their predecessors
from previous decades, let alone earlier centuries.

HOPKIN, S. P.. 1996. — Myriapodology before and after Martin Lister’s « Journey to Paris in the Year I698». In:
Geoffroy, J.-J., MAURIES, J.-P. & Nguyen Duy - Jacquemin. M.. (eds), Acta Myriapodologica. Mem. Mas. natn. Hist,
nat.. 169 : 25-34. Paris ISBN : 2-85653-502-X.

Source :

STEPHEN P. HOPKIN

During the latter stages of research for " Biology of Millipedes” (HOPKIN & Read, 1992),
I began to uncover references to myriapods dating back as far as the 15th century. These
discoveries were made too late to include in our book. However, since then I have tracked down
more than 50 references to centipedes and millipedes in pre-19th century literature, many
illustrated with exquisite woodcuts, engravings and drawings, some in colour.

In this article, I shall give an overview of the development of myriapodology from the time
of Aristotle (384-322 BC) to the mid- 19th century. Before Martin LlSTER’s journey to Paris in
1698, most observations on myriapods were apocryphal, or related to medicines. In the late 17th
century, and 18th century, the diversity of invertebrate life began to be appreciated. Numerous
“compendia” were published, the most important of which was the 10th edition of the Sy sterna
Naturae of LINNAEUS (1758) which formed the basis of modem nomenclature.

The 19th century saw the application of scientific method to the study of centipedes and
millipedes and eventually symphylids and pauropods, although these two groups are not covered
here. This was the “Golden Age" of myriapodology. The beauty and accuracy of publications by
VON STEIN (1841). WAGNER (1841), NEWPORT (1843), SWAN (1864), and the magnificent
coloured plates of KOCH (1863). are testimony of the high standards that can be achieved from
long and careful observation with simple equipment. These workers laid the foundations of
modern myriapodology and we shall forever be in their debt.

THE DAWN OF MYRIAPODOLOGY

The earliest student of zoology whose work has survived was ARISTOTLE (384-322 BC).
Several references to myriapods can be found in translations of his work (e.g. THOMPSON,
1910). In one section on “insects”, millipedes and centipedes are recognised as different
organisms - “some insects are wingless such as the lulus and the centipede”. Elsewhere, the
distinction between the “Sea Scolopendra” polychaete worm and “Land Scolopendra” is made,
the source of much confusion in later centuries. The comment is made that if a Scolopendra is
cut in half, the two pieces move off in opposite directions!

PLINY the Elder (AD 23-79) brought together earlier bodies of scientific knowledge,
most notably in his 37-volume Naturalis Historia (FORD. 1992). Translations of Pliny’s work
(e.g. HOLLAND, 1601) include several references to “multipedes”. However, there is confusion
as to whether these are centipedes, millipedes or woodlice (terrestrial Isopoda). A description of
a cure for “biting of the cheeselips or many feet worms called multipedes” could refer to either.

There is one other pre-Renaissance reference to myriapods in the form of a small woodcut
of a “Skolopendra” (Fig. 1) made by a Byzantine artist in AD 512 to illustrate the Greek Herbal
compiled in the first century AD. by DIOSCORIDES (GUNTHER, 1934). “Skolopendra” are
included due to their supposed medicinal properties. However here, as on numerous other
occasions, it is impossible to decide whether centipedes, or marine polychaete worms, are being
discussed.

Fig. 1. — Illustration by a Byzantine artist in AD 512 to illustrate the Greek Herbal of Dioscoridf.s (from Gunther,
1934).

Source :

MYRIAPODOLOGY BEFORE AND AFTER MARTIN LISTER

The real dawn of zoology after the "dark” period of the Middle Ages is connected with the
name of an Englishman, Edward WOTTON, born at Oxford in 1492, who practised as a
physician in London and died in 1555. WOTTON’s De Different iis Animalium (1552) moved
away from the mythological creatures of earlier works and towards more factual descriptions.

The earliest unambiguous illustrations of marine polychaete worms appeared in the Libri
de Piscibus Marinis of RONDELETIUS (1507-1566) published in 1554. The woodcuts of “Sea
Scolopendra” included in this important book reappear many times in later centuries and are
frequently mis-identified as centipedes.

The first definite illustration of a centipede occurs in the herbal of MATTHIOLUS (1500-
1577) published in 1569. The good sale of his smaller herbal in 1554 with small woodcuts
caused MATTHIOLUS to prepare a luxurious edition. Ferdinand I whose physician ordinary
Matlhiolus was made a large contribution towards the cost. The fine woodcuts were done by
Giorgio LlBF.RALE and Wolf MEIERPECK and the blocks were first printed in the German edition
printed at Prague in 1563, and then sent to Venice. The illustrations of "Sea Scolopendra” were
copied and cited as such from RONDELETIUS (1554). However, there is an original woodcut of a
"Scolopendra'’ which is a true centipede (Fig. 2). The animal is clearly drawn from a specimen
rather than from memory.

In Lib Secundum Diofcoridis. ; 5 r

s'coi.oprs’Du a.

Fig. 2. — A "Scolopendra" from MATTHIOLUS (1569).

Aldrovandi (1522-1605) in De Animalibus Insectis (1638) distinguishes between
millipedes "lulus'', centipedes “ Scolopendra ter rest ris” and polychaete worms “ Scolopendra
marina ', but unfortunately, the accompanying illustrations of what are clearly Lithobius. have 1 1
or 14 pairs of legs instead of the correct 15 pairs. Indeed, the presence of the correct number of
legs on a myriapod as in the illustration of a Brazilian centipede in PlSO (1658), is a good guide
to the scientific accuracy of the artist.

The writings of WOTTON (1552), and Conrad CjESNER (1516-1565) in his huge five-
volume Historia Animalium (1551. 1558, 1587, 1617), were summarised and illustrated by
Thomas MOUFET sometimes MOUFFET, MUFFET or MOFFETT (1553-1604). MOUFET, a
contemporary of Shakespeare, studied medicine at Cambridge and Basel, and practised at
Ipswich and London. His Insectorum sive Minimorum Animalium Theatrum (1634) contains
several woodcuts of recognisable myriapods including a Lithobius with the correct number of
legs (15 pairs), and a rather fine millipede on the title page (Fig. 3). Some editions contain an
Appendix of four plates which are rarely seen. On one of these is a copy of the woodcut of the
" Scolopendra ” of MATTHIOLUS (1569) which has "lost” a pair of legs during the copying!

An English version of the work of WOTTON, GESNER & MOUFET was published by
Edward TOPSEL (1572-1628) in his History of Four-footed Beasts and Serpents (1658).
fOPSEL’s book contains several pages of delightful prose “concerning the Scolopendrae and

STEPHEN P. HOPKIN

Juli”. The Juli “the English after me will call them Gally-worms” - from the resemblance of the
numerous legs to oars on a ship - are treated separately from the Scolopendrae, although
polychaete worms are included with the latter judging from the accompanying illustrations. Both
Scolopendrae and Juli are included with the “Cheeselips” (woodlice) as the “Many-feet”, a
persistent theme (see e.g. KlRCHER. 1678; SlBBALD, 1684; BRADLEY, 1721; Hill, 1752;
SEBA, 1735). Topics mentioned include swarming, metachronal waves of the legs, and the use
of myriapods as medicines, particularly for removal of unwanted hair! There arc also references
on the use of “many feet" as diuretics, a common theme in early medicinal texts (e.g. BOYLE,
1744; JAMES. 1743-1745). Some authors have even reported “multipeda” being excreted with
the urine (Pare, 1634; Aldrovandi. 1638).

INSECTORVJVS

SIVE

Minimorum Annnaliuru
THE ATR VAT

Ohm ab

Edoabdo Wottoso,

CONRADO GllNIlO.

Thomaq-vr Pernio
i'icbojtuiii :

Taiulcm

Tho. Movnri LotkUfUiiiopcrifiimpcilmfj; nuxunu coru in/utuni,
auvtum,pe:li:£tum ;

Et ad vivumcxpicHis Icombusfupri quingcnimSlufliJiiini.

LoBdirdex OflicinJ typognp!uci7w«.C*/«. i 6

Fig. 3. — The litle page of MouFET (1634).

I he following passages from TOPSEL (1658) describe the effects of centipede bites in
vivid detail.

“This Scolopender being provoked bites so sharply that Ludovicus Armarus who gave me one brought out of
Alrica could scarce endure him to bite his hand, though he had a good glove on, and a double linen cloth; for he struck his
forked mouth deep into the cloth, and hung on a long time, and would hardly be shaken off'

"When the land Scolopender hath bitten, the place is all black and blue, putrifies and swells, and looks like to the
dregs of red wine, and is ulcerated with the first bite”

The Historiae Naturalis of JONSTONUS (1657a) and the English translation (1657b) are
examples of the pitfalls of plagiarism although to be fair to JONSTONUS, he does cite the sources
of his illustrations. Much of the text is based on earlier authors and many of the illustrations are
copied from the work of ALDROVANDI, GESNER, MOUFET & TOPSEL. In addition to repeating
the mistakes of earlier authors the 22 and 48 legged Lithobius of ALDROVANDI are reproduced.

Source :

M Y R I A PODOLOG Y BEFORE AND AFTER MARTIN LISTER

JONSTONUS introduced errors during the copying. Some creatures have “lost” or “gained" legs.
The small illustration of a woodlouse, for example, has seven pairs of legs in MOUFET &
TOPSEL, but has gained two extra pairs in JONSTONUS’s book. This “eighteen-legged
woodlouse” still turns up from time to time, most recently in an advertisement for Robinson’s
Barley Water in the U.K. as part of a series on ancient remedies.

1 he mid to late 1 7th century was a period of transition. Work of supreme quality was
published at the same time as anecdotal evidence for outdated concepts such as spontaneous
generation. The illustrations in KlRCHF.R (1678) appear to suggest the development of a
centipede from a putrifying horsetail plant Equisetum (Fig. 4). However, the invention of the
microscope enabled Robert HOOKE (1635-1702), Jan Swammerdam (1637-1680) and
Anthony van LEEUWENHOEK (1632-1723) to publish some of the most important and original
zoological observations ever made.

I. Xjlophjton ex r amnia Lilurni in Mnjro A niter is.

1 1. Ex purref.fli f:/ca an! junc: cju/e

Fig. 4. — “Spontaneous generation" of
animals from plants (from Kircher,
1678). including a centipede (III)
developped from a putrified
horsetai 1 ( Equisetum).

HOOKE’s Micrographia (1665) does not concern us here as this classic work contains no
reference to myriapods. The Historic i Insectorum Generalis of SWAMMERDAM (1669), English
translation (1758) again contains no illustrations of myriapods. However, in one passage,
SWAMMERDAM does make some brief observations on myriapods remarking that he is in
possession of "a Scolopendra of the largest kind which is even a span long and was sent to me
from the East Indies”. It is to LEEUWENHOEK'S Werken (1684-1718) that we must turn for the
first observations on myriapods displaying true application of scientific method.

LEEUWENHOEK discovered the aperture in the poison claws of centipedes (Fig. 5). In his
"Letter 104" sent to the Royal Society on 17th October 1687 from Delft (English translation,
1964), Leeuwenhoek wrote the following:

Source :

STEPHEN P. HOPKIN

"I have often heard people speak about the poisonous nips or Bites, by a certain vermin, which is called
Thousand-legs in the East Indies; this vermin as I was told comes to walk on the naked body of sleeping Persons, and as
this vermin is very cold. People often become restless when they feel these animals. But if People would lie quietly
without moving themselves, the same would not cause People any injury; but owing to this movement, they nip, with
the pincers that they have in front of their head into People's bodies; and although there is no effusion of blood
following this, and only a small red or blue spot remains where this vermin has nipped into the body, there nevertheless
follows an intolerable pain and swelling, which is greater and lasts longer in one Person than in another. To still this
pain there is, they say, no more effective remedy than to kill these Centipedes alive in the olive oil. and to rub this oil
into the affected part. Last year 1 instructed the workmen in this city, who receive the goods from the East Indies, to
bring me a live centipede, with the intention to discover, if possible, the reason for these harmful bites of the centipede.
They thereupon brought me a Centipede the length of a little finger while some others arc quite two fingers long and
more. I look hold of this Centipede by one of the two pincers, with a small pair of pliers; and on bringing the pincer
before the microscope. I saw that the pincers or nippers were continuously being moved towards and away from each
other, to nip or grasp something; in which movement I observed at the same time that each of these pincers was provided
with a tiny hole, which hole had a small groove or gutter, which was made in such a way as to bring the fluid that came
oozing out of this hole to the extreme end of this sharp, sting-like pointed part with which the pincer is fitted.

From these observations, I came to suppose that the Centipede, by nipping with his pincers into People's skin used so
much violence that he damaged some blood- and other vessels, and tore them apart, and that, at the same time, he injected
the aforesaid fluid into the skin. And I furthermore supposed that this fluid was mixed with an injurious sharp Salt: and
that it was not the damage done by the nipping that caused the great pain; but only the suffering inflicted by the noxious
fluid.

I had intended to continue my observations this year, and to this end I had instructed the Workmen to catch the
Centipedes. But they have not observed any. although several were seen on board ship during unloading of the goods,
and were killed there."

Fig. 5. — A plate from Leeuwenhoek's
Werken (1684-1718) sent to the
Royal Society on 17th October
1687. “Fig. 10" shows the poison
claw of a centipede ("Fig. 11")
which has one of its anterior-most
legs missing. The other
illustrations are of the stings of
nettle ( Urtica ).

The observation of LEEUWENHOEK on poison claws were referred to more than a century
later by SMELLIE (1790-1799) in his discussion of the effects of centipede bites.

Source : MNHN , Paris

M YRIA PODOLOG Y BEFORE AND AFTER MARTIN LISTER

"The poisonous weapons of ihe Scolopendra, or centipes, are somewhal different from those of the spider. Its bite
is so painful, especially in the East Indies, as we are informed by Bontius, that it makes the patient almost mad. When
the claws of its forceps are examined by a microscope, on the upper side of each of them, near the point, a small aperture
appears, through which the venom is conveyed to the wound. Of the East India centipedes. Leeuwenhoek had one sent to
him alive; and he found that by pressing the claw, a small drop of liquor issued out of this aperture".

LEEUWENHOEK was clearly a man ahead of his time.

MARTIN LISTER'S JOURNEY TO PARIS

Martin LISTER (1638-1712) was an English naturalist who published important books on
spiders and snails; for a recent biography of LISTER, see PARKER & HARLEY ( 1992). In 1698,
Lister was sent by King William III as a medical attendant to William BENTINCK. Earl of
Portland, on a diplomatic mission to Paris. He recorded his experiences in the one book he
published in English, A Journey to Paris in the year 1698.

LISTER’S account of his visit to Paris proved very popular and ran to three editions in his
own lifetime. It contains much of historical interest and, in particular, its information about
scientific, medical and other technical matters as well as its description of the city itself, and the
17th century way of life, are invaluable in their detail.

Included in the book are six folding plates. Two of these are among the most striking
illustrations of myriapods ever published. Plate 5 (Fig. 6) shows a large millipede “ lulus
Americanus ” and Plate 6 (Fig. 7) a centipede “ Scolopendra Americana', both drawn by Father
Charles PLUMIER. The centipede was in PLUMIER’s collection and was “a foot and a half long,
and proportionally broad". LISTER describes seeing the millipede in the collection of Monsieur
TOURNEFORT.

Fig. 6. — Left: "lulus Americanus " drawn by Father Charles PLUMIER (from LISTER, 1699).

Fig. 7. — Right: "Scolopendra americana" drawn by Father Charles Plumier (from Lister. 1699).

Source :

STEPHEN P. HOPKIN

- He showed me a very great Julus from Brazil, at least six inches long, and two about, round like a cord, very smooth
and shining, of a kind of copper or brazen colour: the feel infinite. like a double fringe on each side: this he had from F.
PlumiER, who afterwards gave me a design of it drawn by the life and in its proper colours".

For someone used to British myriapods, the sight of these spectacular creatures clearly had
a lasting impression on Martin LISTER.

COLLECTION AND CLASSIFICATION

The 18th and early 19th centuries were periods when the huge diversity of animal life
began to be appreciated and comprehensively described. The lavish texts of HILL (1752),
SCHAEFFER (1766), BARBUT ( 178 1 ), DONOVAN (1792-1807), GEOFFROY (1799), SHAW
(1800-1826), CUVIER (1838-1849). OKEN ( 1833-1842) and BERNARD et al. (1842/1843) all
contain illustrations of myriapods, many in colour. Huge collections of specimens were built up.
The wealthy Dutchman Albert SEBA (1665-1736) assembled the richest collection of natural
history objects of his time. His private museum contained several centipedes and millipedes
which are described and illustrated in the catalogue SEBA (1734-1765). SEBA’s specimens were
purchased by Peter the Great and moved to St. Petersburg.

The most important development of the 1 8th century was the system of classification
introduced by Carolus LINNAEUS (1707-1778) in the first edition of his Sy sterna Naturae
(1735). The tenth edition (1758) ranks as one of the most important zoological book ever
published.

In the first edition of Systema Naturae (1735), LINNAEUS recognises five classes of
animals. Class 5, the Insecta, is split into four groups namely Coleoptera, Angioptera,
Hemiptera and Aptera. The Aptera contains eight “Genera” which are separated mainly on the
basis of the number of legs. Woodlice Genus Oniscus, “Pedes 14” are distinguished from the
myriapods which are all in the Genus Scolopendria “Pedes 20” or more. Three “species” are
described, Scolopendria terrestris , Scolopendria marina (polychaete worm), and Julus.

Linnaeus’s introductory notes “Observationes in Regnum Animale”. Observations on the
Animal Kingdom were translated into English by ENGEL-LEDF.BOER & ENGEL in the facsimile
edition of 1964. Point 8 “Scintillas Scolopendrae ” is translated as “the luminescence of
Scolopendria marina a Nereide”. However, it seems much more likely that LINNAEUS is
referring to terrestrial species in which luminescence has been repeatedly observed (BARBUT,
1781: DONOVAN, 1792-1807; SHAW, 1800-1826).

The classification of myriapods is more detailed in the tenth edition of Systema Naturae,
with the “Insecta” comprising seven groups, the last of which “Aptera” contains 14 “Genera”
numbers 230-243. The centipedes (Genus 242 Scolopendra - nine species) are separated from
the millipedes (Genus 243 Julus - seven species), although nereid polychaetes are still included
as Scolopendra marina. The names of Scolopendra electrica from elektron, “a shining substance,
amber or an alloy of gold and silver” (EMMET, 1991) and Scolopendra phosphorea clearly refer
to properties of luminescence. Several of LlNNAEUS’s names are, of course, still in use today.

THE "GOLDEN AGE" OF MYRIAPODOLOGY

By the early 19th century, myriapods began to be recognised as a group distinct from
insects. The catalogue of British insects published by STEPHENS (1829) does not include
centipedes or millipedes. The Nomenclator Zoologicus of AGASSIZ ( 1 842- 1 846) contains many
genus and family names that are familiar to us today. The Myriapodum were divided into two
groups: Chilognatha, the millipedes comprising the families Glomeridae, Julidae, Polydesmidae,
Polyxenidae. Polyzonidae and Siphonophoridae and Chilopoda , the centipedes comprising the
families Cermatidae, Lithobiidae, Scolopendridae and Geophilidae.

The internal anatomy of millipedes and centipedes began to be studied in detail from the
mid 19th century onwards. The standard of draughtmanship of the plates in books by VON

Source :

MYRIAPODOLOGY BEFORE AND AFTER MARTIN LISTER

Stein (1841), Wagner (1841), Swan (1864), and the paper by NEWPORT (1843) has not
been bettered since. However, the peak of myriapodological illustration must surely be Die
Myriapoden by Carl Ludwig KOCH (1778-1857) published in 1863. This book contains
descriptions of more than 200 species of centipedes and millipedes, each of which is figured in
colour plates of breathtaking beauty. These paintings must rank among the most exquisite ever
produced and are a fitting tribute to the efforts of earlier myriapodologists. KOCH’s Die
Myriapoden released from relative obscurity what are surely among the most interesting of the
least-studied animals. Even NEWPORT (1841) bemoaned the preoccupation of naturalists with
insects to the detriment of other arthropods.

ACKNOWLEDGEMENTS

I am very grateful to Mike Bott, David Knott and Dermot O’Rourke of The University of Reading Library for
their help during preparation of this article. I would also like to acknowledge Cole (1944) and Ford (1992) as the
sources for much of the background material concerning the authors of the cited texts.

REFERENCES

Note : These references are to editions I have consulted in the Cole Library of Early Medicine and Zoology at Reading
University.

AGASSIZ, L., 1842-1846. — Nomenclator zoologicus, continens nomina systematica generum animalium turn viventium
quam fossilium . Solothurn, Jent et Gassmann.

Aldrovandi, U., 1638. — De Animalibus Insectis. Bologna. Clement Ferrohi.

Barbut, J., 1781. Les Genres des Insectes de Linne. London, Jacques Dixwell, J. Sewell.

Bernard. P., Couailhac, L., Lemaout, G. & Lemaout, E., 1842-1843. — Le Jardin des Plantes. Description complete,
historique et pittoresque du Museum d'histoire naturelle, de la menagerie, des serres, des galeries de mineralogie el
d’anatomie de la vallee suisse. Paris, L. Curmer.

BOYLE, R., 1744. — The Works of the Honourable Robert Boyle. London, A. Millar.

Bradley, R., 1721. — A Philosophical Account of the Works of Nature. London, W. Meare.

COLE, F. J., 1944. — A history of comparative anatomy from Aristotle to the eighteenth century. London, Macmillan &
Co.

Cuvier, G., 1838-1849. — Le Regne Animal distribue d'apres son Organisation. Paris, Victor Masson, 3rd "Disciples"
edition.

Donovan, E., 1792-1807. — The Natural History of British Insects. London, F. & C. Rivington.

Emmet, A. M., 1991. — The Scientific Names of the British Lepidoptera. Colchester, Harley Books.

FORD, B. J., 1992. — Images of Science: A History of Scientific Illustration. London, The British Library.

Geoffroy, E. L., 1799. — Histoire abregee des Insectes. Paris, Calixte-Volland, Remont.

GESNER, C., 1551, 1558, 1587, 1617. — Historiae Animalium Lib. I-V. Zurich & Frankfort, Apud Christ.
Froschoverum.

Gunther, R. T., 1934. — The Greek Herbal of Dioscorides. Oxford, Oxford University Press.

Hill, J., 1752. — A General Natural History. Vol. Ill Animals. London. Thomas Osborne.

Hooke. R., 1665. — Micrographia. London, J. Martyn & J. Allestry.

HOLLAND, P., 1601. — The Historie of the World, commonly called The Natural Historie of C. Plinius Secundus.

Translated into English by Philemon Holland, Doctor in Physicke. London, Adam Islip.

Hopkin, S. P. & Read, H. J., 1992. — Biology of Millipedes. Oxford, Oxford University Press, 233 pp.

James, R., 1743-1745. — A Medical Dictionary. London, T. Osborne.

JONSTONUS, J., 1657a. — Historiae Naturalis. De Insectis Libri III. Amsterdam, Apud Joannem Jacobi Fil. Schipper.
JONSTONUS. J., 1657b. — A History of the Wonderful Things of Nature. London, John Streater.

Kircher, A., 1678. — Mundus Subterraneus. Amsterdam. Jansson a Waesberge & Son.

Koch, C. L., 1863. — Die Myriapoden. Getreu nach der Natur abgebildet und beschrieben. Halle, H.W. Schmidt.
Leeuwenhoek, A. V., 1684-1718. — Werken. Leiden & Delft, Cornelius Boutesteyn, xxx pp.

Leeuwenhoek, A. V., 1964. — The Collected Letters of Antoni van Leeuwenhoek. Vol. 7. Amsterdam, Swets &
Zeillinger Ltd.

STEPHEN P. HOPKIN

Linnaeus, C., 1735. — Systema Naturae. [Facsimile of the First Edition. With an Introduction and a first English
translation of the "Observationes”. By M. S. J. Engel-Ledeboer and H. Engel. Vol. 8 Dutch Classics on the History
of Science. Nieuwkoop. B. de Graaf, 1964].

Linnaeus, C.. 1758. — Systema Naturae. Regnum Animate. Editio decima 1758. Cura Societatis Zoologicae Germanicae.

Engelmann facsimile reprint of 1894. Tomus I. Animals. Leipzig, Wilhelm Engelmann.

LISTER, M., 1699. — A Journey to Paris in the year 1698. London, Jacob Tonson.The Second Edition.

MATTHIOLUS, P. A.. 1569. — Commentarii in sex libros Pedacii Dioscorides Anazarbei de Medica materia. Venice,
Valgrisi.

MOUFET, T., 1634. — Insectorum sive Minimorum Animalium Theatrum. London. Thom. Cotes.

Newport, G., 1843. — On the structure, relations, and development of the nervous and circulatory systems, and on the
existence of a complete circulation of the blood in vessels, in Myriapoda and Macrourous Arachnida - First Series.
Philosophical Transactions of the Royal Society : 243-302.

Oken, L., 1833-1842. — Allgemeine Naturgeschichte fur alle Stande. Stuttgart, Hoffmann.

Pare, A., 1634. — The Workes of that famous Chirurgion Ambrose Parey. Translated out of Latine and compared with
the French by Th. Johnson. London, Th. Cotes & R. Young.

Parker, J. & Harley, B., 1992. — Martin Lister" s English Spiders 1678. Translated from the original Latin by M.
Davies & B. Harley. Colchester. Harley Books.

PlSO, G., 1658. — De Indiae Utriusque Re Naturali et Medica Libri Quatuordecim Quorum contenta pagina sequens
exhibet. Amsterdam, Ludovic & Daniel Elzevir.

RONDELETIUS, G., 1554. — Libri de Piscibus Marinis , in quibus verae Piscium effigies expressae sunt. Lyon, Apud
Matthiam Bonhomme.

Schaeffer. J. C., 1766. — Elementa Entomologica. Regensburg, Gedruckt mit Weissischen Schriften.

Seba, A., 1735. — Locupletissimi Rerum Naturalium Thesauri. Vol. II. Amsterdam, Janss-Waesberge, J. Wetstein & Gul.
Smith.

Shaw. G., 1800-1826. — General Zoology or Systematic Natural History. London, G. Kearsley.

Sibbald, R., 1684. — Scotia Illustrate. Edinburgh. Jacob Kniblo, Joshua Solingen, John Colmar.

SMELLIE, W„ 1790-1799. — The Philosophy of Natural History. Edinburgh. Charles Elliot.

STEPHENS, J. F.. 1829. — A Systematic Catalogue of British Insects. London. Baldwin & Cradock.

Swammerdam, J., 1669. — Historia Insectorum generalis, ofte Algemeene Verhandeling van de Bloedeloose Dierkens.
Utrecht, Meinard van Dreunen.

Swammerdam, J., 1758. — The Book of Nature; or the History of Insects. London, C.G. Seyffert.

Swan, J., 1864. — Illustrations of the comparative anatomy of the nervous system. 2nd Edn. London, Bradbury &
Evans.

Thompson, D. W. T., 1910. — The Works of Aristotle. Translated into English under the editorship of J. A. Smith & W.

D. Ross. Vol. IV. Oxford, Historia Animalium. Clarendon Press.

Topsel, E., 1658. — The History of Four-Footed Beasts and Serpents. London, G. Sawbridge.

Von Stein, F., 1841. — De Myriapodum partibus genitalibus, nova generationis theoria atque introduction systematica
adjectis. Berlin, Brandes & Klewert.

Wagner, R., 1841. — leones Zootomicae. Leipzig, L. Voss.

Wotton, E., 1552. — De differentiis Animalium Libri Decern. Paris, Apud Vascosanum.

Source : MNHN, Paris

An Approach to the Revision of the
East Asian Millipede Genus Anaulaciulus

Zoltan KORSOS

Department of Zoology, Hungarian Natural History Museum, Baross u. 13, H-1088 Budapest, Hungary

ABSTRACT

The millipede genus Anaulaciulus Pocock, 1895 (Julida: Julidae) comprises 44 species, distributed in Eastern Asia.
Based on fresh studies of numerous samples, type material and literature, a comprehensive overview of the genus is
proposed. A list of the presently known species is given, and a preliminary grouping is outlined on the basis of their
posterior gonopod structure. Other external and internal characters, such as penis, gonopod promerit and female vulva
structure, coloration, size, and shape of the preanal projection are discussed and evaluated. Two examples of
evolutionary gonopod transformation series are presented and illustrated.

RESUME

Essai de revision du genre est-asiatique Anaulaciulus .

Le genre Anaulaciulus Pocock, 1895 (Diplopoda, Julida, Julidae) comprend 44 especes reparties dans PEst asiatique.
La presente revision, basee sur Pexamen recent de nombreux exemplaires, permet de presenter une liste dans laquelle les
especes sont s^parees en groupes provisoirement bases sur la structure des gonopodes posterieurs. D’autres caract£res,
externes et internes, tels que le penis, le promerite des gonopodes, la structure des vulves, la coloration, la taille, et la
forme du telson sont discutes et evalues. Deux exemples de transformation evolutive des gonopodes sont pr^sentes et
illustres.

INTRODUCTION

The genus Anaulaciulus at present consists of 44 nominal species (with 4 proposed
subspecies) including 10 forms recently described from the Southern Himalaya region
(KORSOS, in press). Part of the original descriptions of the other species are rather old and not
properly detailed, type material of those is usually difficult to obtain. As the range of the genus
(see below) implies, there may certainly be a large number of yet undiscovered species.
However, considering the available material, a preliminary review of the genus seems not to be
premature.

The distribution of the species in the genus includes the temperate zone of Eastern Asia:
from Pakistan to the Russian Far East, through Nepal, northern India, Sikkim, Tibet,
northeastern China and Korea, including Hong-Kong and Taiwan. Numerous forms occur also
in Japan, south of Hokkaido (Honshu, Shikoku, Kyushu, Ryukyu Islands, Bonin Islands). To
the contrast of the other widespread Eastern Asian julid genus Nepalmatoiulus, Anaulaciulus

Kors6s, Z., 1996. — An approach to the revision of the East Asian millipede genus Anaulaciulus. In: Geoffroy,
J.-J., M AURlfes, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. nain. Hist, nat., 169 :
35-43, Paris ISBN : 2-85653-502-X.

ZOLTAN KORS6S

does not seem to penetrate into the tropical regions, it is confined to the temperate zone or high
altitudes.

The genus name itself was introduced by POCOCK in 1895. and subsequently generally
overlooked. The majority of the species belonging now to Anaulaciulus were originally
described in Fusiulus Attems, 1909, and only in 1966 did CAUSEY recognize the synonymy
with a redescription of the two POCOCK's species (paludicola and vallicola).

Anaulaciulus belongs in the tribe Brachyiulini, which can be characterized as follows:

Julidae (Brachyiulinae) without a free mesomerit on the posterior gonopod, with a well-
developed flagellum, and generally compressed gonopods in the antero-posterior direction.
About 24 genera are enlisted in this tribe, however, their relationship has not yet been
completely clarified.

The genus Anaulaciulus can be defined on the basis of some peculiarities externally as
well as in the gonopod conformation. The animals have no metazonal setae, no cheek lobes
expanded in the males. Male gonopod promerites are characteristically flattened, scale-like, a
rudiment of the telopodit is well visible. Posterior gonopods are rather simple, elongated, in situ
always protruding from beneath protecting promerites, and have several longitudinal, slightly
arched lamellae. The penis is long, bifurcate in every species; this character seems to be a unique
apomorphy for the genus in the entire millipede order Julida; even the closest relatives of the
genus in the tribe Brachyiulini have a completely different penis (Figs 1-3). The long, leaf-like
structure (differently developed in the different species) seems to be homologuous with the
apical membrane in the other species, and the opening of the seminal groove is situated most
probably caudally at the basis of the “leaves”.

The female vulval characters show also some peculiarities as compared to other members
of the tribe. They are slightly compressed in the antero-posterior direction (others are more-or-
less cylindrical), the well-separated operculum is always longer than bursa and apically provided
with two lateral cusps (often also a median one). The median cleft on bursa is deep, the
apodematic tube without secondary branches, the ampulla usually without an appendix.

A more detailed characterization of the genus is given elsewhere (KORSOS, in press).

There are very few works devoted to a summary or clarification of the internal
relationships of Anaulaciulus. An identification key is given to the species known at that time by
VERHOEFF, first in 1937 (for five species), then in 1941 (for 9 species, VERHOEFF, 1941a),
and by Takakuwa (1941, for 16 forms). They are all based mainly on minor gonopodal
character details, and not very useful, especially if one regards the different quality of the
descriptions and the possible morphological variations in the populations. ENGHOFF (1986) lists
28 nominal species and 6 subspecies with comments on their distribution. He establishes the
synonymies of A. ciliatus Shinohara, 1960 and F. trilobus quemoyensis Wang, 1963. Apart
from these, no attempt for the revision of the entire genus has been made.

REVIEW OF THE SPECIES

In the followings, a renewed alphabetical list of the presently known species in the genus
is given, together with a name history and distributional data of every species. Illustrations
wherever available are also referred to.

1. Anaulaciulus acaudatus Korsos, in press
Anaulaciulus acaudatus : Kors6s, in press (Figs 26-28)

2. Anaulaciulus acutus (Takakuwa, 1941)

Fusiulus acutus: Takakuwa. 1941 (Figs 2-3)

Anaulaciulus acutus : ENGHOFF, 1986

3. Anaulaciulus attemsii (Verhoeff, 1941)

Fusiulus attemsii: VERHOEFF, 1941a (FiGS 31-33)

Anaulaciulus attemsii : ENGHOFF, 1986

India: Sikkim.

Japan: Honshu.

REVISION OF THE EAST ASIAN GENUS MILLIPEDE ANA ULACIULUS

4. Anaulaciulus bilineatus Korsds, in press

Anaulaciulus bilineatus : Kors6s, in press (Figs 2-4, 6, 9, 11, 29-33)

5. Anaulaciulus bilobus (Takakuwa, 1941)

Fusiulus bilobus: Takakuwa, 1941 (Figs 10-11)

Anaulaciulus bilobus : ENGHOFF, 1986

6. Anaulaciulus capillatus (Takakuwa, 1941)

Fusiulus capillatus: Takakuwa, 1941 (Figs 12-13)

Anaulaciulus capillatus: ENGHOFF, 1986

7. Anaulaciulus cornutus (Takakuwa, 1941)

Fusiulus cornutus: Takakuwa, 1941 (FIGS 17-18)

Anaulaciulus cornutus: ENGHOFF, 1986

8. Anaulaciulus enghoffi Korsos, in press
Anaulaciulus enghoffi: KORSOS, in press (FlGS 34-41)

9. Anaulaciulus golovatchi Mikhajlova, 1982
Anaulaciulus golovatchi: Mikhajlova, 1982 (FlG. 2)

Anaulaciulus golovatchi: ENGHOFF, 1986

10. Anaulaciulus hirosaminus (Attems. 1909)

Fusiulus hirosaminus: Attems, 1909 (FlGS 76-78)

Anaulaciulus hirosaminus: ENGHOFF, 1986

11. Anaulaciulus inaequipes Enghoff, 1986
Anaulaciulus inaequipes: ENGHOFF, 1986 (FlGS 1-4)

Anaulaciulus inaequipes: Kors6s, in press (FlGS 20-25)

12. Anaulaciulus kashmirensis Korsos, in press
Anaulaciulus kashmirensis: Kors6s, in press (FlGS 42-47)

13. Anaulaciulus kiusiensis (Vcrhoeff, 1941)

Fusiulus kiusiensis: VERHOEFF, 1941a (FlGS 34-36)

Anaulaciulus kiusiensis: ENGHOFF, 1986

14. Anaulaciulus komatsui (Shinohara, 1957)

Fusiulus komatsui: Shinohara, 1957 in Takakuwa & Shinohara,
1957 (Fig. 2)

Anaulaciulus komatsui (Takakuwa & Shinohara, 1957): ENGHOFF,
1986

Nepal.

Japan: Kyushu.

Japan: Honshu.

Japan. Kyushu.

China: Kansu.

Russia: Far East, Maritime Province;
recently reported from North Korea as well
(Mikhajlova, 1993).

Japan: Hiro Sami.

Burma.

India: Kashmir.

Japan: Kyushu.

Japan: Honshu.

15. Anaulaciulus koreacolus Jedryczkowski. 1982
Anaulaciulus koreacolus: JEDRYCZKOWSKI, 1982 (FlGS 28-36)
Anaulaciulus koreacolus: ENGHOFF, 1986

16. Anaulaciulus koreanus (Verhoeff, 1937)

Fusiulus koreanus: VERHOEFF, 1937 (FlGS 4-8)

Fusiulus koreanus koreanus Verhoeff, 1937: Paik, 1976
Anaulaciulus koreanus: ENGHOFF, 1986
Anaulaciulus koreanus koreanus: LlM, 1988

16.1. Anaulaciulus koreanus boninensis (Verhoeff, 1939)
Fusiulus koreanus boninensis: VERHOEFF, 1939a (FlGS 16-17)
Anaulaciulus koreanus boninensis: Golov ATCH, 1980 (FlGS 1-2)
Anaulaciulus koreanus boninensis: ENGHOFF, 1986

16.2. Anaulaciulus koreanus tuberculatus (Takakuwa, 1941)
Fusiulus koreanus tuberculatus: Takakuwa. 1941 (FlG. 19)
Anaulaciulus koreanus tuberculatus: ENGHOFF, 1986

17. Anaulaciulus kuritai (Murakami, 1966)

Fusiulus kuritai: MURAKAMI, 1966 (FlG. 1)

Anaulaciulus kuritai: ENGHOFF, 1986

18. Anaulaciulus longus (Takakuwa, 1941)

Fusiulus longus: Takakuwa, 1941 (Figs 6-7)

Anaulaciulus longus: ENGHOFF, 1986

19. Anaulaciulus nepalensis Korsds, in press
Anaulaciulus nepalensis: Kors6s. in press (FlGS 1, 3, 7, 10, 48-52)

Korea: Sunchon and Hyangsan districts.

Korea: Hoko.

Japan: Bonin Islands, Ryukyu Islands;
Korea (Takakuwa, 1941; Paik, 1976; Lim,
1988; Golovatch, 1980).

Korea: Hoko (Paik, 1976; LlM, 1988).

Japan: Shikoku.

Japan: Akiyoshi: Korea (LlM, 1988).
Nepal.

ZOLTAN KORSOS

20. Anaulaciulus niger Korsos, in press

Anaulaciulus niger KORSOS, in press (FIGS 53-58) Nepal.

21. Anaulaciulus okinawaensis Shinohara, 1990

Anaulaciulus okinawaensis : Shinohara, 1990 (FlG. 1) Japan: Ryukyu Islands.

22. Anaulaciulus onychophora (Takakuwa. 1942)

Fusiulus onychophora : Takakuwa, 1942 (FlGS 1-2)

Anaulaciulus onychophora : Enghoff, 1986 Japan: Honshu.

23. Anaulaciulus otigonopus Zhang, 1993
Anaulaciulus otigonopus : Zhang, 1993 (FlGS 1-7)

Anaulaciulus otigonopus : Kors6s, 1994

Anaulaciulus otigonopus : Kors6s, in press China: Hunan Province, Changsa.

24. Anaulaciulus pakistanus Korsds, in press

Anaulaciulus pakistanus : KORSdS, in press (FlGS 59-60) Pakistan: Swat.

25. Anaulaciulus paludicola Pocock, 1895
Anaulaciulus paludicola : POCOCK, 1 895

Anaulaciulus paludicola: Causey, 1966 (FlGS 1-6) China: Wo-Lee Lake.

26. Anaulaciulus pinetorum (Attems, 1909)

Fusiulus pinetorum: ATTEMS, 1909 (FlGS 14-16, 69-75)

Fusiulus pinetorum: SHINOHARA, 1960 (FlG. 18)

Anaulaciulus pinetorum: ENGHOFF, 1986 Japan: Honshu.

26.1 Anaulaciulus pinetorum nivalis (Verhoeff, 1941)

Fusiulus pinetorum nivalis: VERHOEFF, 1941b (FlGS 8-10)

Fusiulus ciliatus: Shinohara, 1960 (Figs 14-17): Enghoff, 1986
Anaulaciulus pinetorum nivalis: ENGHOFF, 1986

27. Anaulaciulus quadratus (Takakuwa, 1941)

Fusiulus quadratus: Takakuwa, 1941 (Figs 14-16)

Anaulaciulus quadratus: Takano, 1978
Anaulaciulus quadratus: ENGHOFF, 1 986

28. Anaulaciulus riedeli Jedryczkowski. 1982
Anaulaciulus riedeli: JEDRYCZKOWSKI, 1982 (FlGS 19-27)

Anaulaciulus riedeli: ENGHOFF, 1986

29. Anaulaciulus ryugadensis Shinohara, 1990
Anaulaciulus ryugadensis: SHINOHARA, 1990 (FlG. 2)

30. Anaulaciulus simodanus (Takakuwa, 1941)

Fusiulus simodanus: Takakuwa, 1941 (Figs 8-9)

Anaulaciulus simodanus: ENGHOFF, 1986

31. Anaulaciulus simplex Verhoeff, 1936
Fusiulus simplex: Verhoeff, 1936
Anaulaciulus simplex: Shinohara, 1973
Anaulaciulus simplex: ENGHOFF, 1986

32. Anaulaciulus takakuwai (Verhoeff, 1941)

Fusiulus takakuwai: Verhoeff, 1941a (FIGS 37-38)

Anaulaciulus takakuwai: ENGHOFF. 1986
subspecies:

Anaulaciulus takakuwai coloratus (Verhoeff, 1941)

Fusiulus takakuwai coloratus: VERHOEFF, 1941a (Fig. 39)

Anaulaciulus takakuwai coloratus: ENGHOFF 1986

33. Anaulaciulus takanoi Shinohara, 1990
Anaulaciulus takanoi: Shinohara, 1990 (FlG. 3)

34. Anaulaciulus tibetanus Korsds, in press
Anaulaciulus tibetanus: KORS6S, in press (FlGS 61-63)

35. Anaulaciulus tigris Korsds, in press
Anaulaciulus tigris: Kors6s. in press (FlGS 5, 12, 64-69)

36. Anaulaciulus tonggosanensis Paik, 1976
Fusiulus longus Takakuwa, 1941: sensu Paik, 1963
Fusiulus tonggosanensis: Paik, 1976 (FlGS 1-1 1)

Japan: Honshu.

Korea: Hyangsan, Kyongsong and Puryong
districts.

Japan: Shikoku.

Japan: Honshu.

Japan: Honshu, in caves widely distributed
(Shinohara, 1973); Taiwan (Wang, 1963;
Shinohara, 1973).

Japan: Honshu.

Japan: Honshu, Niijima Island.

Japan: Honshu.

China: Tibet; India: Assam.

Pakistan: Swat.

Korea: Mt. Tonggo-san (LlM, 1988).

REVISION OF THE EAST ASIAN GENUS MILLIPEDE ANAULACIULUS

37. Anaulaciulus tonginus (Karsch, 1881)
lulus tonginus : Karsch, 1881
Anaulaciulus tonginus : ENGHOFF, 1986
Anaulaciulus tonginus : Kors6s, 1994 (FlGS 1-8)

Fusiulus trilobus khuuae Wang, 1963: Kors6s, 1994

38. Anaulaciulus topali Kors6s, in press
Anaulaciulus topali : KORS6S, in press (FlGS 70-75)

39. Anaulaciulus trapezoidus (Wang, 1955)

Fusiulus trapezoidus : Wang, 1955 (FlG. 3)

Anaulaciulus trapezoidus : ENGHOFF, 1986

40. Anaulaciulus trigonalis (Takakuwa, 1941)

Fusiulus trigonalis: Takakuwa, 1941 (FlGS 4-5)

Anaulaciulus trigonalis : ENGHOFF, 1986

41. Anaulaciulus trilobus (Wang, 1963)

Fusiulus trilobus quemoyensis : Wang, 1963
Anaulaciulus trilobus: ENGHOFF, 1986

42. Anaulaciulus vallicola (Pocock, 1895)
lulus vallicola: POCOCK, 1895 (FlG. 13)

Anaulaciulus vallicola: Causey, 1966 (FlG. 7)

43. Anaulaciulus yamashinai (Verhoeff, 1939)

Fusiulus yamashinai: VERHOEFF, 1939b (FlGS 1-3)

Fusiulus jamashinai Verhoeff, 1941a (Figs 40-42): Enghoff, 1986
Fusiulus insulariuni Verhoeff, 1941a: ENGHOFF, 1986

Fusiulus yamashinai: Takakuwa, 1941 (FlG. 1)

Anaulaciulus yamashinai: ENGHOFF, 1986 Japan: Ryukyu Islands.

44. Anaulaciulus yosidanus (Takakuwa, 1941)

Fusiulus yosidanus: Takakuwa, 1941 (FlGS 20-21)

Anaulaciulus yosidanus: Enghoff, 1986 Japan: Honshu.

INTRAGENERIC RELATIONSHIPS

The only internal classification of the genus appears in the division by VERHOEFF (1941b)
where he, on the occasion of a new subspecies, Fusiulus pinetorum nivalis, erected the
subgenus Parfusiulus for all the other members of the genus. The only species, pinetorum (with
the subspecies nivalis) remained in the subgenus Fusiulus s. str. in his sense. However, the
distuingishing character (i.e. two hairy fields on the mesal and lateral lamellae of the
opisthomerites) seems not to be warranted, especially in the light of a more careful study of the
gonopodal details in other species. As a result, virtually all species of the genus have more-or-
less hairs on their opisthomerit lamellae.

According to an examination of the shape of the telopodites of the posterior gonopods, the
following preliminary species-groups in the genus can be presented.

1. yamashinai- group (cf. Figs 8-11): acutus, bilobus, comutus, komatsui, onychophora,
pinetorum. quadratus, trigonalis and yamashinai

2. paludicola-group (cf. Figs 5-6): koreacolus, longus, paludicola, riedeli, simodanus and
tonggosanensis

3. koreanus- group: koreanus, okinawaensis, trapezoidus

4. h irosam in w.s-grou p : hirosaminus, kuritai

5. simplex- group (cf. Fig. 7): attemsii, simplex

6. tonginus- group: otigonopus, tonginus. trilobus

7. inaequipes- group: acaudatus, bilineatus, enghoffi, inaequipes, kashmirensis.
nepalensis, niger, pakistanus, tibetanus, tigris and topali

The species takakuwai can be considered as a bridge-species between the paludicola-
group and the koreanus-g roup (based purely on gonopod comparison).

Hong Kong; Taiwan.

India: Jammu and Kashmir.

Taiwan.

Japan: Kyushu, Kagoshima.
Taiwan: Quemoy Island.

China: Che Kiang, Da-Zeh valley.

ZOLTAN KORS6S

Six species could not be inserted in the groups above: capillatus, golovatchi (Fig. 4),
kiusiensis, ryugadensis, takanoi, yosidanus. In some cases their gonopods are so peculiar (e.g.,
in takanoi) that even their validity within the genus Anaulaciulus may be question-marked. (The
original description of this species does not deal with some important features like penis
structure, etc.).

One species, vallicola is known only by female, and although the type specimen has been
redescribed by CAUSEY (1966) and also seen by the author, nothing can be said about its
position in the genus.

Based on some fresh material, kindly loaned by Dr. H. ONO (National Science Museum,
Tokyo) some preliminary sketches are given to illustrate two main general pattern series. Figures
5 to 7 (samples from Korea and Japan) show the line of complete reduction of the
opisthomerites, from a “paludicola”- type gonopod to a simple “needle”. Anaulaciulus golovatchi
(Fig. 4. drawn from a paratype kindly loaned by Dr. S. 1. GOLOV ATCH, Moscow) may perhaps
also be inserted in this series.

Figs 1-3. — Penis, caudal view. - 1:
Anaulaciulus bilinealus Korsds, in
press from Nepal. - 2:

Megaphyllum unilineatum (C. L.
Koch, 1838) from Beograd,
Yugoslavia. - 3: Anaulaciulus
koreanus (Verhoeff, 1939) from
North Korea. Scale 0.5 mm.

Figs 4-7. — Left opisthomerit, frontal
view. - 4: Anaulaciulus golovatchi
Mikhajlova, 1982, paratype. - 5:
Specimen from North Korea, Mt.
Paekdu-san. - 6: Specimen from
Japan, Ryukyu Islands, Tokara. - 7:
Specimen from Japan, Ryukyu
Islands, Amami. Scale 0.5 mm.

The other line is more complicated but some characteristics can be observed. The lateral
lamellae of the opisthomerit appears as a “shoulder” (Fig. 8) and later, through a series of
intermediates (Figs 9-11), develops into a broad “wing” (Fig. 12) as it is seen in the inaequipes-
group. Although all this drawings are based on species originated from Japan, there is a striking
resemblance between the gonopods of the specimen from Honshu (Gifu) and those of the

Source :

REVISION OF THE EAST ASIAN GENUS MILLIPEDE ANAULACIULUS

the similar - maybe synonymous - species, otigonopus and trilobus) is believed to have a
somewhat peculiar position in the genus. Not only its intermediate penis and gonopod structure
(thick, antero-posteriorly not so flattened promerites; peculiar telopodits with a beginning of a
beak yet densely haired), but also its central geographical distribution (Hong-Kong, Taiwan
and maybe other parts of southeastern China) implies that it is close to the theoretical ancestor of

the whole genus.

Figs 8-12. Left opisthomerite, frontal view. - 8: Specimen from Japan, Honshu, Chojaga mori. - 9: Specimen from
Japan, Kyushu, Nagasaki. - 10: Specimen from Japan, Kyushu, Yaku-shima. -II: Specimen from Japan, Kyushu,
Kumamotol. - 12: Specimen from Japan, Honshu, Gifu. Scale 0.5 mm.

CONCLUSIONS

It is clear from the present observations, that the shape of the scale-like promerit is very
variable in the populations and that it is not a reliable character for distinguishing species This
was already introduced by Mikhailova (1982), and further discussed by Kors6s (in press).
Unfortunately, descriptions of former species, in some cases, have been exclusively based on
the shape of the promerit (e.g., acutus, bilobus & quadratus, all by Takakuwa, 1941). The
degree of the morphological variability of the opisthomerites is still to be defined, and a
clarification may well be resulted in a number of synonymies in the species-groups outlined
above.

Female (vulval) characters, as often neglected before, are also in urgent need to redescribe.
The species in the inaequipes- group (KORSOS, in press) show relatively consistent pattern in the
internal structure of bursa, usually having a simple or slightly curved apodematic tube and a
more-or-less sphaerical ampulla; whereas other species may have more complicated apodematic
tube ( golovatchi ), or an ampulla strongly elongated ( tonginus , riedeli , kiusiensis) or with a
distinct appendix ( koreanus ).

As it was shown by the analysis of the inaequipes- group, external characters have usually
an emphasized importance in distinguishing the different species. General body colouration

ZOLTAN KORS6S

(longitudinal stripes e.g., in bilineatus, bright yellow ground colour with dark brown blotches
ordered according to pro- and metazona: as in tigris and pakistanus) is more characteristic to
several species than the gonopod conformation, and may also be more useful in separating them.
Outside the inaequipes- group, one can also find similar feature: golovatchi, paludicola, tonginus
and yamashinai show three black, longitudinal stripes.

The shape of the epiproct may also help in distingushing the species, while members of
the inaequipes-group never have a preanal project turned upwards (usually it is short, straight,
or missing), the same character state, to a different degree, is not rare in the other continental and
in the Japanese species (e.g., in golovatchi, koreacolus , riedeli, ryugadensis, takanoi &
tonginus).

In some cases, maybe due to coexistence, significant size differences appear in closely
related species-pairs ( nepalensis-niger , pakistanus- tigris). This phenomenon is analyzed in more
detail elsewhere (KORSOS, in press).

Future investigations should aim at the more accurate characterization of the species, and,
with the accumulation of large material from the geographically remote areas, also from Japan!
the internal relationships of this highly diverse and complex genus will become possible to be
clarified.

ACKNOWLEDGEMENTS

I would like to thank Dr. Henrik Enghoff (Copenhagen) for his kind help and continuous encouragement during
my stay in the Zoologisk Museum, Copenhagen, where a part of this study was carried out. My participation at the 9th
International Congress of Myriapodology in Paris and the presentation of this paper was made possible by the support
ol the Phare Accord Mobility Project of the National Committee for Technical Development, Hungary (Project No. 126).

REFERENCES

Attems, C., 1909. — Die Myriopoden der Vega-Expedition. Ark. Zool. 5 : 1-84.

Causey, N. B.. 1966. — Redescriptions of two Chinese species of Anaulaciulus (Diplopoda, Julidae, Nemasomatidae), a
genus known also in Taiwan, Korea, and Japan. Proc. Louisiana Acad. Sci., 29 : 63-66.

Enghoff, H., 1986. Leg polymorphism in a julid millipede, Anaulaciulus inaequipes n. sp. With a list of congeneric
species (Diplopoda, Julida. Julidae). Steenstrupia, 12 : 117-125.

Golov atch, S. I„ 1980. — A contribution to the millipede fauna of Korea (Diplopoda). Folia em. hung.. 41 : 49-58.

Jeoryczkowski, W„ 1982. — New and rare millipedes (Diplopoda, Julida) from North Korea. Annls Zool PAN. 36 :

Karsch, F 1881. — Neue Juliden des Berliner Museums, als Prodromus einer Juliden-Monographie. Z. ges. Naturwiss.,
3** I-/?.

KorsOs, Z 1994. — Redescription of Anaulaciulus tonginus (Karsch. 1881) (Diplopoda. Julida, Julidae). Steenstrupia,
: 177-182.

KorsOs, Z., (in press). — Another Himalayan group of julid millipedes: Towards the clarification of the genus
Anaulaciulus Pocock, 1895) (Diplopoda: Julida). Senckenbergiana biol.

Lim, K. Y., 1988. — Taxonomical studies on Class Diplopoda from Korea. Mag. Rer. Natur. Thesis, Fac. Agric. Educ.
Inst. Educ., Univ. Yuanguan., 34 pp.

Mikhailova, E. V„ 1982. — New millipedes of the family Julidae (Diplopoda) from the Soviet Far East. Zool. Zhur.,
61 : 210-216.

M’KHajlova, E. V„ 1993. — The millipedes (Diplopoda) of Siberia and the Far East of Russia. Arthropoda Selecta. 2 :
3-36.

Murakami, Y„ 1966. — Postembryonic development of the common Myriapoda of Japan. XX11. Three new species of
millipeds. Zool. Mag.. 75 : 94-97.

P.MK K. Y 1963. — Survey of the myriapods of Mt. Sokkri, Chungcheung-pookdo, Korea. Theses Coll. Kyungpook
Univ., 7 : 33-42.

Paik K. Y„ 1976. — A new myriapod of the genus Fust ulus (Julidae: Diplopoda). Theses Coll. Grad. School Educ
Kyungpook Nat. Univ.. 6/7: 157-160.

Source :

REVISION OF THE EAST ASIAN GENUS MILLIPEDE ANAULACIULUS

Pocock, R. I., 1895. — Report upon the Chilopoda and Diplopoda obtained by P. W. Basset-Smith, Esq., Surgeon R.
N„ and J. J. Walker, Esq., R. N„ during the cruise in the Chinese Seas of H. M. S. Penguin', Commander W U
Moore commanding. Ann. Mag. Nat. Hist., Ser. 6., 15 : 346-372.

Shinohara, K„ 1960. — Three new species of Juloidea (Diplopoda) from Chichibu. Bull. Chichibu Mus. Nat. Hist.,

Shinohara, K., 1973. — The fauna of the lava caves around Mt. Fuiji-san XIII. Diplopoda and Chilopoda. Bull. Nat Sci
Mus., 16 : 217-251.

Shinohara, K., 1990.
42 : 21-25.

— Three new species of the genus Anaulaciulus (Diplopoda: Julidae) from Japan. Edaphologia

Takano, M.. 1978. — Comparative effects of BHC and Malathion against millipedes, Anaulaciulus quadratus
(Takakuwa). Acta Arachnol., 28 : 39-44.

Takakuwa, Y., 1941. — Die Fusiulus- Arten (Diplopoda). Trans. Sapporo Nat. Hist. Soc., 16 : 218-226.

Takakuwa, Y., 1942. — Einige neue Arten von Diplopoda aus Nippon. Zool. Mag., 54 : 237-239.

Takashima. H. & Shinohara, K., 1957. — Taxonomical and morphological study on myriapods collected in the
vicinity of Towada, Tohoku District, Japan. Misc. Rep. Yamashina Inst. Ornithol. Zool., 11 : 24-31.

Verhoeff, K. W., 1936. — Uber Diplopoden aus Japan, gesammelt von Herm Y. Takakuwa. Trans. Sapporo Nat. Hist
Soc., 14 : 148-172.

Verhoeff, K. W., 1937. — Zur Kenntnis ostasiatischer Diplopoden. II. Zool. Anz., 119 : 33-40.

Verhoeff, K. W.. 1939a. — Zur Kenntnis ostasiatischer Diplopoden. IV. Zool. Anz., 127 : 273-285.

Verhoeff, K. W.. 1939b. — Diplopoden von der Ryukyu-Insel Okinawa. Biogeographica. Trans, biogeogr. Soc. Japan,

Verhoeff, K. W. 1941a. — Asiatische Beitrage VI. Revue Fac. Sci. Univ. Istanbul , 6 : 31 1-318.

Verhoeff, K. W., 1941b. — Zur Kenntnis ostasiatischer Diplopoden. VI. Zool. Anz., 136 : 62-70.

Wang. Y. M., 1955. — Serica la: Records of myriapods on Formosa with description of new species. Quart. J Taiwan
Mus., 8 : 13-16. -

Wang, Y. M., 1963. — Serica lq: Millipedes and centipedes of Quemoy, Fukien Province, and Taiwan Island, Botel
Tobago (Lan Yu), Taiwan Province and of Singapore. Quart. J. Taiwan Mus., 16 : 89-96.

Zhang, C., 1993. — Small myriapoda in soil from China I. A new julidan species Anaulaciulus otigonopus (Julida:
Julidae). Acta Zootax. Sinica, 18 : 18-21.

Source : MNHN , Paris

The Taxa of Rhymogona
(Diplopoda: Craspedosomatidae): a Ring Species
Part One: Genetic Analysis of Population Structure

Adolf SCHOLL * & Ariane Pedroli-Christen **

* Department of Population Biology, Institute of Zoology
University of Berne, Baltzerstrasse 3
CH-3012 Berne, Switzerland
** CSCF, Musee d’Histoire Naturelle, Terreaux 14
CH-2000 Neuchatel, Switzerland

ABSTRACT

The genetic analysis of the population structure of Rhymogona is based on allozyme data from vertical starch gel
electrophoresis (14 enzyme loci surveyed) and includes all taxa (73 collecting sites). The genetic structure of
Rhymogona populations is not consistent with current taxonomy. We find five major groups of populations which are
arranged in a circular fashion around the Jura. Adjacent groups differ from each other in allele substitutions at five
polymorphic loci altogether and are connected by clinal variation. The extreme populations of this ring differ in allele
substitutions at four loci. They are found in Switzerland where they obviously came into secondary contact after the last
glaciation. They form narrow hybrid zones in the Jura and the Alps. Our data suggest that Rhymogona must be regarded
as a polytypic species if the biological species concept is applied.

RESUME

Rhymogona (Diplopoda, Craspedosomatidae), un genre monospecifique. Premiere partie :
analyse genetique de la structure des populations.

L’analyse genetique de la structure des populations de Rhymogona est bas6e sur les observations des allozymes par
electrophorese sur gel d'amidon vertical (14 loci enzymatiques) et porte sur tous les taxons de ce genre r6coltes dans 73
stations. La structure genetique des populations de Rhymogona ne coincide pas avec la taxinomie usuelle. Nous
constatons qu'il existe cinq principaux groupes de populations, celles-ci 6tant distributes de manitre circulate autour du
Jura. Les groupes adjacents se distinguent par des substitutions alleliques dans cinq loci polymorphiques et sont relids
par des variations clinales. Les populations se situant aux extremes se distinguent par des substitutions alltliques dans
quatre loci. Elies ont ttt recenstes en Suisse ou, de toute Evidence, elles semblent etre entrees secondairement en contact
aprts la dernitre glaciation. Elles forment d'ttroites zones hybrides dans le Jura et les Alpes. Sur la base de nos rtsultats,
Rhymogona doit etre considtrt comme espece polytypique si l’on veut tenir compte du concept biologique de l’esptce.

Scholl, A. & Pedroli-Christen, A., 1996. — The taxa of Rhymogona (Diplopoda: Craspedosomatidae): a ring
species. Part one: genetic analysis of population structure. In: Geoffroy, J.-J., Mauri£s, J.-P. & Nguyen Duy -
Jacquem tn, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 45-51. Paris ISBN : 2-85653-502-X.

ADOLPH SCHOLL & ARIANE PEDROLI-CHRISTEN

INTRODUCTION

Rhymogona is a small genus of the Diplopod family Craspedosomatidae which lives north
of the Swiss Alps and in adjacent parts of France and Germany. Seven nominal species are
recognized, most of these are known from one or a few localities only (PEDROLI-CHRISTEN &
Scholl, this volume). We have initially attempted to study the distribution in detail (PEDROLI-
CHRISTEN, 1990). Since species identification in this genus is based essentially on subtle
differences in morphology of genitalia and is often not unambiguous, we have asked if enzyme
electrophoretic data might be used as additional information for species identification. Our
samples cover the whole area of distribution of Rhymogona and all taxa described (73 collecting
sites).

MATERIAL AND METHODS

The collecting sites are shown in Fig. 1 and are listed in Table 1 along with the species diagnosis based on
morphological criteria (Pedroli-Christen & Scholl, this volume, for details of taxonomy and identification). The
specimens were stored at -80°C prior to electrophoresis. Electrophoretic studies (vertical starch gel electrophoresis) were
conducted using routine techniques of our laboratory (cf. Scholl et al. , 1990; Pedroli-Christen & Scholl, 1990). We
have scored 14 loci; Apk, Got- 1, Got-2, a-Gpd, Gpt, Hk. Idh. Mdh-1, Mdh-2, Mod, Mpi, 6Pgd, Pgi and Pk. Five loci were
polymorphic and indicated genetic differentiation among populations: Got-1. Mpi, 6Pgd, Pgi and Pk.

Mendelian inheritance of the electromorphs observed could not be assessed by breeding experiments but is
assumed by analogy (cf. Zimmermann & SCHOLL, 1993). Due to initial difficulties in resolving Mpi, this enzyme was not
scored in some populations (Table 1). The designation of alleles is based on electrophoretic mobilities (in mm) of the
electromorphs; R. montivaga from the Alps (sites 2-5 in Table 1 and Fig. 1) were used as reference (assigned index = 100
for the common allele at each locus). Coefficients of genetic identity (I) were calculated in pairwise comparisons of the
populations using the formula given by NEI (1972). These coefficients served as a matrix for average linkage cluster
analysis (UPGMA) (Nei, 1987).

RESULTS AND DISCUSSION

Table 1 shows the allele frequencies at five polymorphic loci. We have not listed seven
very rare alleles which were observed at one or the other locus in nine populations altogether.
These alleles do not contribute to the outline of the Rhymogona population structure presented.
Sample sizes were very low in some collecting sites. We have listed these sites in Table 1, but
samples with < 5 specimens were not included in further treatment of data because adequate
sample sizes are critical in genetic analysis of populations. Identification of individuals based on
morphology suggested that several samples contained more than one taxon (e.g. sites 27, 33, 53
in Table 1 and Fig. 1). The electrophoretic data, however, gave no evidence for the coexistence
of genetically separated gene pools at these sites or at any other site. With respect to the enzyme
phenotypes observed, we found no significant deviations from HARDY-WEINBERG
expectations. For calculations of allele frequencies and further treatment of data we have pooled
all specimens of a particular site.

Two alleles were found at the Got-1 locus, Got-1 ioo and Got-196 respectively. In most
populations, however, one or the other allele was fixed. Allele Got-1 1 00 was observed in most
R. montivaga populations (sites 1 - 1 1 in Table 1), montivaga populations from the western part
of the Swiss Jura (sites 12 - 14) were polymorphic, French R. montivaga populations (sites 15 -
16) instead had allele Got-196 fixed as all other populations and taxa except montivaga/cervina
hybrid populations (sites 67 - 73).

Two alleles were found at the 6-Pgd locus, 6-Pgdioo and 6-Pgd94 respectively. Allele 6-
Pgdioo was fixed in all populations and taxa except the Swiss R. cervina populations. The R.
cervina populations in the Swiss Jura (sites 55 - 62) had allele 6-Pgd94 fixed; R. cervina
populations along the Rhine (sites 44 - 47, 50 and 53), populations from the cervina/ alemannica
contact zone in the Swiss Jura (sites 26, 27) and populations from the montivaga/cervina hybrid
zone in the Swiss Jura and the Alps (sites 65 - 73) were polymorphic.

Source :

RHYMOGONA : GENETIC ANALYSIS OF POPULATION STRUCTURE

Sampling Site

Taxon

fr— on -op^.0^

•pKImont

Goi-1

96 10C

6Pgd

94 10C

Hole Frequenc
Pk

94 10C

100

i o:

1 00

Mpi

Leukerbad

m montivaga

1.0

1.0C

1.00

Gemml

m monUvaga

1 6

0.16

0.8

1.0C

1.00

0.37

llligenaip

m. monUvaga

1.0C

0.96

0 96

0 04

Sanetsch

m. montivaga

1.0C

0.04

0.9t

1.0C

1.00

1 00

Lauenen

m. montivaga

1.0C

0.9£

1.0C

1.00

1 00

Tour de Famolon

m montivaga

1.0C

1 .00

Morgins

m montivaga

1.0C

1.00

1 .00

Vouvry

m. montivaga

1.0C

1 00

Rochers de Naye

m. montivaga

1.0C

l.OC

1.00

Lossy

m. montivaga

1.0C

Le Cachoi

m. montivaga

1.0C

1.00

La Brdvtne

m. montivaga

0.05

0.9£

i.6c

1.0C

1 .00

Mauborgel

m. montivaga

0.10

0.9C

1.0C

1.00

0.89

0 1?

Si. Georges

m. montivaga

0.19

0.8

1.0C

1 .00

Grande Chanreuse

m. montivaga

1.00

1.0C

1.00

1 00

Levier

m. montivaga

1 1

1.00

1.0C

1.00

0 96

Oeschaux

m. hessei

1.00

1.0C

1.00

1 .00

M6didre

m hessei

1.00

1.0C

1 .00

1.00

0.90

0.10

Belverne

m. hessei

1.00

1.0C

1 .00

1.00

0 21

0 79

Beaune

m. hessei

1.00

1.0C

1.00

1 .00

Ancey

montivaga -group '

1.00

1 00

Lantenay

montivaga -group '

1.00

1 00

Vernot

m. hessei

1.00

1 00

Coudedoux

alemannlca

1.00

0.81

0.19

1.00

Noir Bo is

alemannlca

1.00

0.86

0.14

1.00

1 00

Le Breuil

cervlna, alemannica"

1.00

0.83

0.17

0.83

0.17

1.00

Si. Ursanne

cervlna. alemannica "

1.00

0.38

0.63

0.96

0.04

0.98

Bonlol

alemannica

1.00

1.-00

1.00

1 00

Boncourl

alemannica ?

1.00

0.50

1.00

1 .00

Masevaux

alemannlca

1.00

Le Haul-du-Them

alemannlca

1.00

1 .00

Linihal

alemannlca

1.00

Kenzlngen

cervna * alemannica ?

1.00

1 .00

Hornberg

cervlna, verhoelll "

1.00

1 00

Marbach

cervlna-group"

1.00

1 .00

alemannlca

1.00

0.88

0.10

1.00

Otlwangen

serrata

0.91

0.07

1.00

0.39

0.61

1.00

0.98

Inzlingen

serrata

1.00

0.20

0.80

0.85

0.15

1 00

Hasel

wehrana

1.00

0.98

0.02

1 .00

Todlmoos

wehrana

1.00

0.95

0.05

0.95

1 00

4 1

Menzenschwand

wehrana

1.00

Laulenburg

verhoelll

1.00

0 37

0 63

Tlefen6iein

verhoelll

1.00

0.10

0.90

1 .00

Sulz

cervlna-group '

1.00

0.29

0.71

1.00

0.86

0.14

0 93

Schuplari

cervlna-group ’

1.00

0.50

1.00

0.50

Homburg

cervlna

1 6

1.00

0.32

0.68

1.00

0.97

0.03

1 00

Kussnach

cervina

1.00

0.13

0.88

1.00

0.50

Altglashuiten

cervlna + verhoelll ?

1.00

1 00

Gutachbrucke

verhoelll, wehrana "

1.00

0.50

1.00

0.75

Hu6mersee

cervlna

1 1

1.00

0.70

0.30

1.00

l .00

Dissenhofen

cervina

1.00

Hemishofen

cervlna

1 4

1.00

1 00

Siaad

cervlna. alemannica * '

1.00

0.13

0.88

1.00

Baar

cervlna

1.00

0.95

0.05

0.99

1.00

0.02

0 98

Oberguisch

cervlna ?

1.00

0.75

0.25

1.00

1 .00

Trub

cervlna

1.00

0.86

0.14

1.00

Schelten

cervlna

1.00

Berllncourl

cervlna

1.00

0 61

0 39

Gorges de Coud

cervlna

1.00

0.33

0.67

1.00

1 .00

Combe Biosse

ervlna

1.00

1 .00

Perluls

ervlna

1.00

1 .00

PrAvoux

ervlna

0.99

0.01

0.97

0.03

0.76

0.21

1.00

0.65

0.35

Yalsainte

'ervina-group'

1.00

Schwarzenmalt

ervina

1.00

0.33

0.67

1.00

0.18

0.82

aun

ervina-group'

1.00

0.83

0.17

1.00

1 00

<andersteg

ervlna

0.95

0.05

0.95

0.05

0.58

0.43

1.00

0.08

0.92

Zwelsimmen

ervlna / montivaga

0.48

0.53

0.15

0.85

0.25

0.75

1.00

0.28

0.42

0.30

3oltigen

ervina / montivaga

0.78

0.22

0.75

0.25

0.71

0.30

1.00

0.15

0.85

3eseux

ervlna / montivaga

0.43

0.57

0.71

0.29

0.06

0.94

1.00

1 .00

Rochelort

ervlna / montivaga

0.57

0.33

0.83

0.17

»4auvaise Combe

ervina / montivaga

0.67

0.33

0.46

0.54

0.13

0.88

1.00

a Chaux-du-Milieu

ervina / montivaga

0.67

0.33

0.60

0.40

0.18

0.82

1.00

erridres

ervina / montivaQa

0.84

0.16

0.57

0.43

0.03

0.93

1.00

0.47

0.05

0.48

Table 1. — Allele frequencies at five polymorphic loci (rare alleles are not listed). * = females only; ** = identification
ambiguous; sites 67-73 = cervina/montivaga hybrid populations.

Source : MNHN, Paris

ADOLPH SCHOLL & AR1ANE PEDROLI-CHRISTEN

Two alleles. PklOO and Pk94 respectively, were found at the Pk locus. Allele Pkioo was
fixed in all R. montivaga populations. Polymorphism was observed in R. alemannica
populations from the Swiss Jura (sites 24 - 29), in populations from the southwestern Black
Forest region (sites 36 - 40) which were keyed out as alemannica , serrata and wehrana, and in
the montivaga /cervina hybrid zone.

a R.m. montivaga
o R.m. hessei
m R. cervina

b R. montivaga / cervina
hybrid zone

• R. alemannica

a R. cervina I alemannica
hybrids ?

a R. serrata

a R. verhoeffi

o R. wehrana

Fig. 1. — Sampling sites of electrophoretically analysed Rhymogona specimens (species diagnosis based on
morphological criteria).

At the Pgi locus most populations were monomorphic for allele PgiiOO. A second allele,
Pgii03, was observed in low frequencies or even fixed in six populations from the southern
Black Forest region, including the taxa serrata, wehrana , verhoeffi and cervina, as shown in
Table 1.

Due to initial difficulties in resolving Mpi, this enzyme was not scored in all populations.
Furthermore, many specimens, in particular those from R. cervina populations in Switzerland
and those from montivaga/cervina hybrid populations, failed to show Mpi activity. Possibly this
is due to the presence of a null allele. For calculation of Mpi allele frequencies we have assumed
that specimens with no Mpi activity are homozygous for a null allele.

Source : MNHN, Paris

RHYMOGONA : GENETIC ANALYSIS OF POPULATION STRUCTURE

In populations of R. montivaga and in montivaga/cervina hybrid populations we scored
the allele Mpiioo. Populations of other taxa were usually monomorphic for Mpil02, except two
populations from the French Jura (sites 18 and 19), both keyed out as R. m. hessei, which were
polymorphic. The allele frequencies observed in these two populations suggested clinal variation
towards populations from the Vosges.

Cluster analysis of coefficients of genetic identity (I) (populations from the
montivaga/cervina hybrid zone, sites 66 - 73, not included) resulted in several major groups of
populations with very high levels of genetic identity (I > 0.98). These groups are shown in
Fig. 2. Group A has the Swiss R. montivaga populations; group B has French populations
keyed out as R. m. montivaga and R. m. hessei respectively; group C has m. hessei populations;
group D has the northern Rhymogona populations and includes the taxa alemannica, cervina ,
and wehrana ; groups E and F have R. serrata and R. verhoeffi, respectively; group G has the
Swiss R. cervina populations and includes specimens from the type locality of R. aelleni (site
54). These groups usually differ, in the order as they are presented, by allele substitution at one
locus (Fig. 2). Group E which has the two R. serrata populations is exceptional because it is
polymorphic at the Pk locus and therefore has an intermediate position between groups C and D.
According to the allele frequencies observed (Table 1) group E is more close to group C with
respect to genetic identity.

□ R. m. montivaga
o R. m. hessei
■ R. cervina

n R. montivaga / cervina
hybrid zone

• R. alemannica

a R. cervina I alemannica
hybrids ?

& R. serrata

4 R. verhoeffi

o R. wehrana

allele 100

other alleles:

Got 96
Mpi 102
Pgi 103
6Pgd 94
PK 94

Fig. 2. — Genetic differentiation of Rhymogona populations.

ADOLPH SCHOLL & AR1ANE PEDROLI-CHRISTEN

li is important, however, to realize that the differentiation among these groups is not
abrupt. The allele substitutions observed between population groups change in a clinal fashion.
These clines appear to be shallow in some regions and steeper in other regions, as far as we can
see from a rather limited number of individuals and/or populations in some areas.

More generally, the electrophoretic data show that the genetic structure of Rhymogona
populations is not consistent with current taxonomy. This is most clearly evident from a
comparison of populations from the Black Forest region and from the Vosges (group D in
Fig. 2), which include the taxa aleinannica, cervina and wehrana according to morphology.
These populations are largely identical with respect to the alleles observed and to their
frequencies (Table 1). In contrast, R. cervina populations from Switzerland are different from R.
cervina populations in the Black Forest region. Furthermore, R. verhoeffi, which has the allele
Mpii03 substituted for Mpiioo, is clearly differentiated from the other taxa, however, Mpii03 is
also observed in low frequencies in other populations from nearby localities (sites 38, 39, 44
and 46 in Table 1 and Fig. 1). These specimens were keyed out as serrata, wehrana, and cervina
respectively. The electrophoretic data suggest gene flow among these taxa and an isolation-by-
distance model of genetic differentiation.

The more relevant information obtained from the electrophoretic survey are the
observations that the alleles and their frequencies change largely independently of morphological
characters and that they change in a clinal fashion within and among taxa. The groups of
populations are arranged in a more or less circular fashion around the Jura. Groups A and G,
which have obviously colonized this area after the last glaciation, come into secondary contact in
the Swiss Jura and in the Alps. These two groups differ by allele substitution in four loci, and
they form rather narrow hybrid zones in the Swiss Jura and the Alps, as we have shown
previously (PEDROLI-CHRISTEN & SCHOLL, 1990).

CONCLUSIONS

Rhymogona species, as in most other diplopods, were initially described using the
morphospecies concept. Other species concepts have been developed since (cf. HAFFNER,
1986). The biological species concept which defines species as "groups of interbreeding natural
populations that are reproductively isolated from other such groups" (Mayr, 1969) is now
chosen by the majority of zoologists (Mayr, 1963, 1970; HEWITT, 1990). As summarized in
Figure 2, our data show that Rhymogona consists of groups of genetically differentiated
populations. However, there is no evidence that these groups are reproductively isolated. In
contrast, our data show that there is gene flow between these groups. Our results therefore have
taxonomic consequences and suggest that Rhymogona must be regarded as a polytypic species.
All species presently recognized should be revised to subspecies of Rhymogona montivaga as
will be discussed (PEDROLI-CHRISTEN & SCHOLL, this volume).

ACKNO WLEDGEM ENTS

We are indepted to Mrs. V. Siegfried and Mrs. L. Frauchiger for technical assistance in the electrophoretic
studies and to S. Hunziker for computer graphics. Critical comments and suggestions of Dr. Henrik Enghoff and Dr.
Adam H. Porter and Dr. John R. Spence on an earlier version of this manuscript are gratefully acknowledged.

REFERENCES

Haffner, J., 1986. — Superspecies and species limits in vertebrates. Z. zool. Syst. Evolutionsforsch., 24 : 169-190.
Hewitt, G. M., 1990. — Divergence and speciation as viewed from an insect hybrid zone. Can. J. Zool., 68 : 1701-
1715.

Mayr. E., 1963. — Animal species and evolution. Cambridge, Massachusetts, Harvard University Press.

Mayr, E., 1969. — Principles of systematic zoology. New York, McGraw-Hill.

Mayr. E., 1970. — Populations, species and evolution. Cambridge, Massachusetts, Belknap Press of Harvard
University.

Nei, M., 1972. — Genetic distance between populations. Am. Nat. 106 : 283-292.

Source :

RHYMOGONA : GENETIC ANALYSIS OF POPULATION STRUCTURE

NEI, M., 1987. — Molecular evolutionary genetics. New-York. 512 pp.

Pedroli-Christen, A., 1990. — Field investigations on Rhymogona cervina Verhoeff and Rhymogona silvatica
Rothenbiihler (Diplopoda): Morphology, distribution and hybridization. In : A. Minelli, Proc. 7th int. Congr.
Myriapodology. Leiden, E. J. Brill : 27-43.

Pedroli-Christen, A. & Scholl, A., 1990. — Ecological and genetic studies on parapatric Rhymogona silvatica Roth,
and R. cervina Verh. (Diplopoda: Craspedosomatidae) with special reference to hybrid populations in a zone of
contact. Rev. suisse Zool., 97. 349-359.

Scholl, A., Obrecht, E. & Owen, R. E., 1990. — The genetic relationship between Bombus moderatus Cresson and the
Bombus lucorum auct. species complex (Hymenoptera: Apidae). Can. J. Zool.. 68 : 2264-2268.

Zimmermann. M. & SCHOLL, A.. 1993. — Specific status of Aquarius cinereus (Puton) and Aquarius najas (De Geer)
(Hemiptera: Gerridae) and the extent of hybridisation in the Mediterranean region. Ent. scand 24 : 197-210.

Source : MNHN, Paris

Rhymogona (Diplopoda, Craspedosomatidae), un genre
monospecifique. Deuxieme partie : Revision basee sur
les resultats morphologiques, genetiques et

faunistiques

Ariane PEDROLl-CHRISTEN * & Adolf SCHOLL **

* C.S.C.F., Musee d’Histoire Naturelle, Terreaux 14, CH-2000 Neuchatel, Suisse
** Departement Biologie des Populations, Institui de Zoologie, Universite de Berne
Baltzerstrasse 3, CH-3012 Berne, Suisse

RESUME

La revision du genre Rhymogona, qui comprend selon la bibliographic sept taxons nominaux, est.basee sur des
observations faunistiques et morphologiques (analyse des genitalia) ainsi que sur des resultats genetiques, presentes dans
la premiere partie de ce travail (Scholl & Pedroli-Christen, ce volume). Le taxon aelleni (Schubart, 1960) est mis en
synonymie avec cervina. Nous reconnaissons une espece polytypique. Rhymogona montivaga (Verhoeff, 1894) ; les
taxons cervina (Verhoeff, 1910), alemannica (Verhoeff, 1910), verhoeff i (Bigler, 1913) serrata (Bigler, 1913) et
wehrana (Verhoeff, 1910) sont considers comme des sous-especes.

ABSTRACT

Rhymogona (Diplopoda, Craspedosomatidae), a ring species. Second part: revision based upon
morphological, genetic and faunistic results.

The revision of the genus Rhymogona, which houses seven nominal species according to previous authors, is based
on faunistic and morphological studies (analysis of genitalia) and on genetic data, presented in part 1 of this work
(Scholl & Pedroli-Christen, this volume). The taxon aelleni (Schubart. 1960) is placed in synonymy with cervina
(Verhoeff, 1910). We recognize a polytypic species. Rhymogona montivaga (Verhoeff, 1894); the taxa cervina
(Verhoeff, 1910), alemannica (Verhoeff, 1910), verhoeffi (Bigler, 1913) serrata (Bigler, 1913) and wehrana (Verhoeff.
1910) are revised to subspecies.

INTRODUCTION

Plusieurs travaux concernant le genre Rhymogona ont fait l'objet de publications recentes.
PEDROLI-CHRISTEN (1990) resume, selon les donnees bibliographiques, la repartition
geographique des diverses especes alors recensees pour le genre et l'etat des connaissances
systematiques. Par ailleurs, des recherches faunistiques recentes permettent de mettre en
evidence des zones de contact entre deux taxons, montivaga (syn. silvatica voir PEDROLI-
CHRISTEN & SCHOLL, 1991) et cervina dans le Jura et les Prealpes suisses. L'analyse

Pedroli-Christen, A. & Scholl, A., 1996. — Rhymogona (Diplopoda, Craspedosomatidae), un genre
monospecifique. Deuxieme Partie : Revision basee sur les resultats morphologiques, genetiques et faunistiques. In:
Geoffroy, J.-J.. Mauris, J.-P. & Nguyen Duy - Jacquemin. M., (eds), Acta Myriapodologica. Mem . Mus. natn. Hist,
nat., 169 : 53-60. Paris ISBN : 2-85653-502-X.

AR1ANE PEDROLI-CHRISTEN & ADOLPH SCHOLL

morphologique des genitalia des individus males et femelles en provenance de ces zones mene a
l'hypothese de l’existence de phenomenes d'hybridation entre ces deux taxons.

Le recours a des analyses genetiques (electrophoreses enzymatiques) complementaires
permet de confirmer cette hypothese (PEDROLI-CHRISTEN & SCHOLL, 1990). Les resultats
obtenus relativisent le statut d'espece attribue a ces deux taxons. Afin de mieux comprendre la
systematique de Rhymogona, un elargissement de la recherche a l'ensemble du genre sur toute
son aire de repartition (Suisse, Nord des Alpes ; alentours de la Foret Noire en Allemagne ; en
France, de l'Alsace et des Vosges a la Savoie) a ete effectue. Les resultats obtenus sont presentes
en deux parties, fortement imbriquees : - la premiere partie (cf. Part 1, ce volume), fait etat des
analyses genetiques de structures des populations et mene a la notion de “ring species” - la
deuxieme partie presente les consequences systematiques des resultats sous forme d'une revision
du genre.

MATERIEL ET METHODES

Le materiel a et 6 recolte par chasse-^-vue sous des 6corces et bois morts au sol dans 250 sites r£partis sur
l’ensemble de l’aire de repartition du genre, surtout en septembre et octobre, de 1986 h 1991. 925 individus collectes en
73 stations ont fait l'objet d’analyses morphologiques et genetiques (Tableau 1 et Fig. 1 in Scholl & Pedroli-Christen,
Part I).

Le materiel recent a ete compare pour identification aux figures et descriptions des auteurs, mais aussi au materiel
original de VERHOEFF et de BIGLER.

Collection Verhoeff, Staatssammlung de Munich :

R. alemannica (Verhoeff. 1916. Staad/Rorschach; Verhoeff, 1935, Mindelsee) ; R. a. rotundatum (Verhoeff, 1916,
Badenweiler) ; R. cervina (Verhoeff, 1910, Pratteln, Schonberg bei Freiburg; Verhoeff, 1916, Sulz bei Laufenburg.
Immendingen) ; R. c. brevidentatum (Verhoeff, 1916, Tiengen) ; R. verhoeffi (Verhoeff, 1916. Rottweil am Neckar) ;
R. v. excavatum (Verhoeff. 1916, Andelsbachtal bei Klein-Laufenburg) ; R. wehrana genuinum (Verhoeff, 1910 Wehr,
VERHOEFF, 1936 Hollental) ; R. wehrana clavigerum (VERHOEFF, 1935, Schonau) ; R. wehrana calcivagum (VERHOEFF,
1910, Wehr) ; R. wehrana quadridentatum (VERHOEFF, 1935, Zell).

Collection BIGLER, Mus6e d'Histoire Naturelle de Bale :

R. alemannica (Vogesen, Sondernach, Nenzlingen, N'Lauchen, Tschapperli, Reinacherallmend) ; R. a. triarticulalum
(Bellackerkopf, Linthal, Sondernach) ; R. a. globosum (N'Lauchen, Hochfeld) ; R. a. alsaticum (Glasbachli) ; R. cervina
(Guldental, Oberdomach) ; R. verhoeffi (Gutach) ; R. serrata (Ottwangen, Hagenbach).

HISTORIQUE

- VERHOEFF (1894) decrit Atractosomci montivagum. Rochers de Naye (Vaud/CH) et
Daubensee (Valais/CH).

- COOK (1896) cree le genre Rhymogona qui reste oublie jusqu'au travail de HOFFMAN
(1980).

- VERHOEFF (1897) cree le genre Macheiriophoron, couramment utilise par la suite.

- ROTHENBUHLER (1899) decrit A. montivagum silvaticum (Villeneuve Vaud/CH).

- VERHOEFF (1910) decrit M. alemannicum, Hohentwil et Rufach en Alsace(F) ;
M. cervinum, Schonberg bei Freiburg (D) et Pratteln (Basel/CH) ; M. wehranum , Wehr (ouest)
dans le Wehratal (D). II attribue le statut d’espece a M. silvaticum decrit par ROTHENBUHLER
(1899).

- BIGLER (1913) decrit M. verhoeffi, Gutach en Foret Noire et M. serratum, Ottwangen et
Hagen au Dinkelberg (D) pres de Bale.

- SCHUB ART (1960) decrit M. aelleni, grotte de Baar (Zoug/CH).

Des sous-especes ou varietes ont ete decrites pour pratiquement toutes les especes :

- BROLEMANN (1935) M. silvaticum hessei, Prenois, Cote d'Or (F).

- BIGLER (1913) M. alemannicum genuinum, Jura Suisse, rive gauche de la Birse ;
M. alemannicum globosum, Niederlauchen en Alsace (F) ; M. alemannicum triarticulatum,
vallees dans le sud des Vosges (F).

- Verhoeff (1916) M. alemannicum rotundatum, Badenweiler (D).

- VERHOEFF (1916) M. cervinum brevidentatum, Tiengen und Thalmiihle (D).

RHYMOGONA, GENRE MONOSPECIFIQUE : REVISION SYSTEMATIQUE

- VERHOEFF (1910) M. wehranum calcivagum, Wehr (est) dans le Wehratal (D).

- VERHOEFF (1935) M. wehranum clavigerum , Schonau dans le Wiesetal (D) ; M. w.
quadridentatum, Zell dans le Wiesetal (D).

- VERHOEFF (1916) M. verhoeffi excavatum, Andelsbachtal bei Klein-Laufenburg (D).

- PEDROLI-CHRISTEN & SCHOLL (1991) proposent de considerer R. silvatica comme
synonyme de R. montivaga sur la base des analyses morphologiques et genetiques.

MORPHOLOGIE COMPAREE DES GENITALIA FEMELLES

Pour tous les taxons sus-mentionnes deux types de structures de vulves des femelles sont
reconnaissables, soit celui de R. montivaga hessei (RAVOUX, 1942) et R. montivaga (PEDROLI-
CHRISTEN, 1990) chez qui la valve externe est courte et la valve interne longue, soit celui de
R. alemannica (VERHOEFF, 1913) et R. cervina (PEDROLI-CHRISTEN, 1990), chez qui la valve
externe est longue et la valve interne courte.

Les taxons R. montivaga, R. wehrana et R. serrata ont une morphologie vulvaire du
premier type et sont difficilement discemables les uns des autres. R. alemannica, R. cervina et
R. verhoeffi presentent le deuxieme type morphologique et ne peuvent pas etre distingues les uns
des autres. Ceci est d'autant plus valable pour leurs sous-especes respectives.

MORPHOLOGIE COMPAREE DES GONOPODES
Taxons montivaga , wehrana et serrata (Fig. 1 A, B, C)

La determination de ces trois taxons, geographiquement bien separes, ne presente en
principe pas trop de difficultes. Cependant, relevons la variability existant a l'interieur meme de
chaque espece morphologique et ceci essentiellement pour la structure des cheirites et des
paragonopodes (PEDROLI-CHRISTEN, 1990; PEDROLI-CHRISTEN & SCHOLL, 1991 pour
R. montivaga).

Le fait que VERHOEFF ait decrit autant de formes appartenant au taxon wehrana va dans le
meme sens. La forme calcivagum, qui ne presente plus le crochet caracteristique a la base du
cheirite, ressemble alors tres fortement a R. montivaga. Rappelons ici qu'a l'inverse, plusieurs
males en provenance des zones hybrides localisees dans le Jura Suisse presentent, contrairement
aux deux formes parentales montivaga et cervina, un cheirite tres semblable a celui de
R. wehrana (Fig. 1, B et Fig. 8 in : PEDROLI-CHRISTEN, 1990).

Les differences morphologiques entre montivaga et montivaga hessei sont tres faibles et se
situent au niveau de l'extremite de la come rostrale qui est aplatie et accompagnee dune lamelle
chez la sous-espece.

Taxons cervina, aelleni, alemannica et verhoeffi
a) cervina (Fig. 1 E, F)

Ce taxon se distingue, selon VERHOEFF (1910 et 1916) :

- au syncolpocoxite: par une longue corne rostrale falciforme (raccourcie chez
brevidentatum) a peine plus courte que la lame en faucille ; par une courbe de la lame en faucille
simple et ne presentant pas de dent triangulaire, tout au plus une ou deux petites pointes ;

- au cheirite : par le prolongement basal du cheirite en general resserre vers le haut et vers
le bas ainsi qu'une pointe dressee bien developpee.

L'observation des males en provenance de 16 populations (cf. Table 1, Part 1) amene a
nuancer certains de ces points :

- la longueur de la come rostrale est variable selon les populations et, souvent, a l'interieur
meme d'une population (la distance entre la pointe de la lame en faucille et la pointe de la come
rostrale varie entre 0 (les deux pointes se juxtaposent) et 0,13mm (ce que Ton observe pour
R. alemannica) ;

ARI ANE PEDROLI-CHRISTEN & ADOLPH SCHOLL.

- si certaines populations presentent une lame en faucille a courbe simple (ex. stations 33,
48. 46 ou 50, Part 1) beaucoup d'autres developpent une dent bien marquee (Fig. 1 E, G ; Fig.
6B in : PEDROLI-CHRISTEN, 1990). Les deux variantes ont ete observees, par exemple, dans
des populations du Jura (stations 61, 62) ou a Baar (54).

Fig. 1. — Cheirites et colpocoxites de : A : R. m. montivaga, Mauborget (CH) 1990 (dessin : J. Spelda) ; B : R. m.
wehrana , Hasel (D) 1991 (dessin : J. Spelda) ; C : R. m. serrata, Inzlingen (D) 1990 (dessin : J. Spelda); D ; R.
m. verhoeffi , Hornberg, Gutachtal (D) 1991 (dessin: J. Spelda) ; E: R. m. cervina , Pratteln (CH) 1910.
Zoologische Staatssammlung Miinchen ; F : R. m. cervina , Kussnach (D) 1990 (dessin : J. Spelda) ; G : R. m.
alemannica , Staad (CH) 1916. Zoologische Staatssammlung Miinchen ; H : R. m. alemannica , Badenweiler 1990
(D) (dessin : J. Spelda).

FlG. 1. — Cheirites and colpocoxites of: A: R. m. montivaga, Mauborget (CH) 1990 ( drawning : J. Spelda); B: R. m.
wehrana. Hasel (D) 1991 (drawning: J. Spelda); C: R. m. serrata, Inzlingen (D) 1990 (drawning: J . Spelda); D: R.
m. verhoeffi, Hornberg, Gutachtal (D) 1991 (drawning J. Spelda); E: R. m. cervina, Pratteln (CH) 1910,
Zoologische Staatssammlung Miinchen; F: R. m. cervina. Kussnach (D) 1990 (drawning: ./. Spelda); G: R. m.
alemannica, Staad (CH) 1916, Zoologische Staatssammlung Miinchen; H: R. m. alemannica, Badenweiler 1990
(D) (drawning: J. Spelda).

Source : MNHN, Paris

RHYMOGONA, GENRE MONOSPECIFIQUE : REVISION SYSTEMATIQUE

b) aelleni

Ce taxon a ete decrit par SCHUBART sur la base d'un seul male en provenance d'une grotte
pres de Baar. Aucune autre station n'est connue. Nous avons recolte dans les environs
immediats de cette grotte (station 54) 17 males et 10 femelles. La morphologie de ces individus,
de meme que les figures de aelleni dessinees par SCHUBART, entrent dans les variations
observees pour cervina. Par ailleurs, les individus de Baar ne presentent aucune difference
genetique avec les cervina en provenance de Suisse (Table 1, Part 1).

Nous proposons done de considerer aelleni comme synonyme de cervina.

c) alemannica (Fig. 1 G, H)

Ce taxon se caracterise, selon VERHOEFF (1910 & 1916) par :

- une come rostrale courte au syncolpocoxite (comme chez serrata)

- la courbe de la lame a faucille divisee en deux par une dent triangulaire

- le haut et le bas du prolongement basal du cheirite en general non resserre et sa pointe
dressee et courte.

La differenciation des varietes decrites pour alemannica ( globosum , triarticulatum) se base
sur des criteres variables, tels les telopodites des paragonopodes. La variabilite de ces pieces a
deja ete soulignee anterieurement pour montivaga par exemple (PEDROLI-CHRISTEN, 1990). Si
les populations en provenance de France, d'Ajoie et de Badenweiler ne posent pas de probleme
de determination, d'autres populations, ou certains individus parmi elles, sont difficilement
distinguables de cervina (26, 27, 53 ; Table 1, Part 1), un ou-plusieurs caracteres se rapprochant
ou se confondant avec les caracteres de cervina.

d) verhoeffi (Fig. 1 D)

Du point de vue morphologique, ce taxon occupe une position intermediate entre cervina
et wehrana. Selon VERHOEFF (1916), la corne rostrale du syncolpocoxite egale ou depasse la
longueur de la lame en faucille et est droite ou recourbee vers le haut. Sur la courbure apicale du
cheirite il peut y avoir une petite dent. La base du prolongement basal presente vers l'arriere une
gibbosite. Ce taxon a ete identifie sans probleme dans deux localites (station 42, 43, Table 1,
Part 1), ailleurs (stations 34 et 49, Table 1, Part 1), une separation nette par rapport a cervina et
wehrana est plus difficile.

D'une maniere generate, on constate done pour tous les taxons une variabilite
morphologique relativement importante, pouvant engendrer des difficultes de diagnostic car la
limite entre deux taxons morphologiques est, dans certains cas, floue. Les populations
problematiques sont souvent situees dans les regions ou les resultats enzymatiques montrent des
transitions entre groupes de populations genetiquement differencies.

DISCUSSION

Les nombreux taxons du genre Rhymogona ont ete decrits au debut de ce siecle au moment
ou la myriapodologie connaissait un grand essor dans la region. La systematique etait alors
exclusivement basee sur le concept typologique de l'espece. Selon l'ampleur des variations
observees, de nouvelles sous-especes ou especes (morphologiques) etaient alors decrites a la
moindre difference. Si la classification typologique est un outil de travail facilement utilisable
dans la pratique et applique, selon MAYR (1967), au debut de toutes recherches scientifiques
d'un groupe, elle n'est aujourd'hui plus d'actualite. Les resultats de cette etude des populations
appellent a suivre le concept biologique de l'espece, definie comme un groupe de populations se
reproduisant entre elles, mais qui est toutefois reproductivement isole d'autres groupes de
populations (MAYR, 1967). En fonction des resultats obtenus, particulierement l'arrangement
geographique des populations genetiquement differenciees, nous proposons de considerer
Rhymogona comme un genre monospecifique et l'unique espece du genre comme une espece
polytypique. Les differentes “especes morphologiques’' doivent alors etre traitees comme des
sous-especes :

ARIANE PEDROLI -CHRISTEN & ADOLPH SCHOLL

Rhymogona Cook, 1896
Macheiriophoron Verhoeff, 1897

Genus Rhymogona Cook, 1896

Espece-lype : Atractosoma montivaga Verhoeff 1894
Espece-type : Atractosoma montivaga Verhoeff 1894

Rhymogona montivaga (Verhoeff, 1894)

Rhymogona montivaga montivaga (Verhoeff, 1894)

1894 Atractosoma montivagum Verhoeff
1899 Macheiriophoron montivagum silvaticum Rolhenbiihler
1910 Macheiriophoron silvaticum Verhoeff
1990 Rhymogona silvatica Pedroli-Christen

1990 Rhymogona montivaga Pedroli-Christen et Scholl

1991 Rhymogona montivaga Pedroli-Christen et Scholl
1993 Rhymogona montivaga Pedroli-Christen

R. montivaga hessei (Brolemann, 1935)

1935 Macheiriophoron montivagum hessei Brolemann
1942 Macheiriophoron montivagum hessei Ravoux
1959 Macheiriophoron montivagum hessei Demange

R. montivaga cervina (Verhoeff, 1910)

1910 Macheiriophoron cervinum Verhoeff
1913 Macheiriophoron cervinum Bigler

1915 Macheiriophoron cervinum Verhoeff

1916 Macheiriophoron cervinum Verhoeff

1916 Macheiriophoron cervinum var. brevidentatum Verhoeff
1934 Macheiriophoron cervinum Schubart
1936 Macheiriophoron cervinum Verhoeff
1960 Macheiriophoron aelleni Schubart (syn. nov.)

1990 Rhymogona cervina Pedroli-Christen

1991 Rhymogona cervina Pedroli-Christen et Scholl
1 99 1 Rhymogona cervina Spelda

1993 Rhymogona cervina Pedroli-Christen

R. montivaga alemannica (Verhoeff, 19 1 0)

1910 Macheiriophoron alemannicum Verhoeff
1913 Macheiriophoron alemannicum Bigler
1913 Macheiriophoron alemannicum var. globosum Bigler
1913 Macheiriophoron alemannicum var. triarticulatum Bigler
1913 Macheiriophoron alemannicum Verhoeff
1916 Macheiriophoron alemannicum genuinum Verhoeff
1916 Macheiriophoron alemannicum rotundatum Verhoeff
1916 Macheiriophoron alemannicum var. triarticulatum Verhoeff

1 934 Macheiriophoron alemannicum Schubart

1935 Macheiriophoron alemannicum Verhoeff
1983 Macheiriophoron alemannicum Kobel-Voss
1991 Rhymogona alemannica Spelda

1993 Rhymogona alemannica Pedroli-Christen

RHYMOGONA , GENRE MONOSPECIFIQUE : REVISION SYSTEMATIQUE

R. montivaga verhoeffi (Bigler 1913)

1913 Macheiriophoron verhoeffi Bigler
1916 Macheiriophoron verhoeffi genuinum Verhoeff
1916 Macheiriophoron verhoeffi excavatum Verhoeff
1991 Rhymogona verhoeffi Spelda

R. montivaga serrata (Bigler, 1913)

1913 Macheiriophoron serration Bigler
1 99 1 Rhymogona serrata Spelda

R. montivaga wehrana (Verhoeff, 1910)

1910 Macheiriophoron wehranum Verhoeff
1916 Macheiriophoron wehranum genuinum Verhoeff
1916 Macheiriophoron wehranum calcivagum Verhoeff
1935 Macheiriophoron wehranum genuinum Verhoeff
1935 Macheiriophoron wehranum quadridentatum Verhoeff

1935 Macheiriophoron wehranum clavigerum Verhoeff

1936 Macheiriophoron wehranum Verhoeff
1991 Rhymogona wehrana Spelda

REMERCIEMENTS

Nous tenons a remercier vivement Jorg Spelda pour sa collaboration sur le terrain et pour les dessins mis & notre
disposition, ainsi que le Dr. Henrik Enghoff et Yves Gonseth pour la lecture critique du manuscrit. Nos rcmerciements
vont egalement au Dr. H. Fechter, Zoologische Staatssammlung Miinchen et au Dr. M. BRANCUCCI, Naturhistorisches
Museum Basel, pour le pret de materiel de collection. L'Academie Suisse des Sciences, qui a accorde une allocation pour
les recherches sur le terrain en Allemagne et en France, m£rite aussi notre gratitude.

REFERENCES

Bigler. W., 1913. — Die Diplopoden von Basel und Umgebung. Rev. suisse Zool., 21 : 675-793.

Brolemann, H., 1935. — Faune de France 29. Myriapodes Diplopodes (Chilognathes I). Paris. P. Lechevalier, 1-369.
COOK, O. F., 1896. — II. On recent diplopod names. Brandlia : 5-8.

Demange, J.-M., 1959. — Myriapodes des cavites de la Cote d'Or, de la Saone-et-Loire et du Jura. Sous le Plancher , 2 :
32-35.

Hoffman. R. L., 1980. — Classification of the Diplopoda. Genfcve, Museum d’Histoire naturelle, (1979), 237 pp.
Kobel-Voss, A., 1983. — Zur Isopoden- und Diplopodenfauna des Naturschutzgebietes "Mindelsee". [In : Der Mindelsee
bei Radolfzell.] Natur- u. Landschaftschutzgebiete Bad.-Wiirti., 11 : 531-538.

Mayr, E., 1967. — Artbegriff und Evolution (dtsch. Obersetzung von Animal, Species and Evolution). Hamburg,
Berlin. 617 pp.

PEDROL1-CHRISTEN, A., 1990. — Field investigations on Rhymogona cervina Verhoeff and Rhymogona silvatica
Rothenbuhler (Diplopoda): Morphology, distribution and hybridisation. In : A. MlNELLI, Proc. 7th ini. Congr.
Myriapodology , Leiden, E. J. Brill : 27-43.

Pedroli-Christen, A., 1993. — Faunistique des Mille-pattes de Suisse (Diplopoda) / Faunistik der Tausendfussler der
Schweiz (Diplopoda). Neuchatel. Centre Suisse de Cartographic de la faune. Doc. faun, helv., 14, 1-248.
Pedroli-Christen, A. & Scholl, A., 1990. — Ecological and genetic studies on parapatric Rhymogona silvatica (Roth.)
and R. cervina (Verh.) (Diplopoda: Craspedosomatidae) with special reference to hybrid populations in a zone of
contact. Rev. suisse Zool., 97 : 349-359.

Pedroli-Christen, A. & Scholl, A., 1991. — Systematique et taxonomic du genre Rhymogona (Diplopoda:
Craspedosomatidae): Rhymogona silvatica (Rothenbuhler. 1899) synonyme de Rhymogona montivaga
(Verhoeff, 1894); resultats morphologiques et genetiques. Rev. suisse Zool., 98 : 83-92.

Ravoux, P., 1942. — Description de la femelle de Macheiriophoron silvaticum hessei. Arch. Zool. exp. & gen., 82 :
91-99.

Rothenbuhler, H., 1899. — Ein Beitrag zur Kenntnis der Diplopodenfauna der Schweiz I. Rev. suisse Zool., 6 : 1 99-
271.

Source :

ARI ANE PEDROLI -CHRISTEN & ADOLPH SCHOLL

Schubart, O., 1934. — Tausendfiissler Oder Myriapoda I. Diplopoda. In : F. Dahl, Tierw. Deutschl. 28 Jena, G.
Fischer, 318 pp.

Schubart, O., 1960. — Uber einige Hohlen-Diplopoden der Schweiz und Frankreichs. Rev. suisse Zool. 67 : 561-
588.

SPELDA, J., 1991. — Zur Faunistik und Systematik der Tausendfiissler (Myriapoda) Sudwestdeutschlands. Jh. Ges. Natur.
Want.. 146: 211-232.

Verhoeff, K. W., 1894. — Beitrage zur Diplopodenfauna der Schweiz. Berl. entom. Z.,39 : 281-296.

Verhoeff, K. W., 1897. — Ubersicht der mir genauer bekannten Europai'schen Chordeumiden-Gattungen (Beitrage zur
Kenntnis palaarktischer Myriopoden 5). Arch, f Naturg., 63 : 129-138.

Verhoeff, K. W., 1910. — Uber Diplopoden. Deutsche Craspedosomiden. Sitz.ber. Ges. naturforsch. Freunde Berlin :
19-62.

Verhoeff, K. W.. 1913. — Die weiblichen Fortpflanzungswerkzeuge von Listocheiritium und Macheiriophoron. Zool.
Anz., 41 : 398-409.

Verhoeff, K. W., 1915. — Beitrage zur Kenntnis der Diplopoden von Wurttcmberg, Hohenzollen und Baden. Jh. Ver.
vaterl. Naturk. Wiirttemberg , 71 : 1-54

Verhoeff, K. W., 1916. — Beitrage zur Kenntnis dcr Gattung Macheiriophoron und Craspedosoma. Zool. Jb., 39 :
273-416.

Verhoeff, K. W., 1935. — Quer durch Schwarzwald und schweizerischen Jura. Verb, natunviss. Ver. Karlsruhe, 31 :
153-174.

VERHOEFF, K. W.. 1936. — Unsere Kenntnis von den Diplopoden des alemannischen Gaues. Ber. naturf. Ges. Freiburg,
35 : 162-195.

Source : MNHN ' Paris

Mastigophorophyllon (Verhoeff, 1897) et
Karp atophy lion Jawlowsky, 1928 : genres carpatiques

(Chordeumatida, Diplopoda)

Traian CEUCA

Universitatea din Cluj-Napoca, Facultatea de Biologie. Catedra de Zoologie, str. Clinicilor 5-7

RO-3400 Cluj-Napoca, Roumanie

RESUME

Le genre Mastigophorophyllon comprend en g£n6ral des formes de haute altitude vivant surtout dans les prairies
alpines, parfois & la lisi£re des forets de coniferes et plus rarement & celle des forets de feuillus. Sur les six especes
depourvues de rameau plumiforme sur la partie posterieure des gonopodes anterieurs, cinq sont repandues uniquement dans
les Carpates Meridionales ; ce sont : M. alpivagum , M. deubeli , M. transsilvanicum , M. carpaticum et M. banarescui. La
seule espece situee en dehors de l’aire carpatique est M. bohemicum , repartie en Boheme. Sur les dix formes comportant
un rameau plumiforme sur la partie posterieure des gonopodes anterieurs, huit sont cantonnees dans les Carpates du Nord
et les Carpates Orientales ; ce sont : M. penicilligerum, M. cir rife rum, M. jickelii, M. serrulatum , M. s. apiculatum ,
M. crinitum, M. c. huculicum , M. aberratum et M. saxonicum. Deux autres formes se rencontrent en Bulgarie (Monts
Balkans) : M. bulgaricum et M. b. pirinicum. Le genre Karpatophyllon renferme seulement quatre especes : K. polinskii ,
K. dacicum, K. carpaticum et K. banaticum. Elies sont repandues dans unc aire qui relie les Carpates du Nord-Est aux
Carpates Meridionales, par P intermediate des Monts Apuseni et des Monts Poiana Ruscai ; cette repartition circonscrit
I’ensemble du Plateau de Transylvanie. On peut affirmer que les genres Mastigophorophyllon et Karpatophyllon sont
bien lies, d’un point de vue geographique, a la Chaine carpatique.

ABSTRACT

Mastogophorophyllon (Verhoeff, 1897) and Karpatophyllon Jawlowsky, 1928, Carpathian
genera (Chordeumatida, Diplopoda)

The genus Mastigophorophyllon comprises of forms living at high altitudes, especially in prealpine areas, sometimes
on the edge of coniferous woods but seldom on the edge of deciduous woods. Among the six species showing no
“featherlike” branches on the posterior part of the anterior gonopods, five are found only in the meridional Carpathian
Mountains, these being: M. alpivagum , M. deubeli , M. transsilvanicum , M. carpaticum and M. banarescui. The only
species found outside of the Carpathian area is from Bohemia. Ten forms have a “featherlike” branch at the posterior part
of the anterior gonopods, eight of them being distributed in the Northern and Eastern Carpathians: M. penicilligerum ,
M. cir rife rum, M. jickelii , M. serrulatum, M. s. apiculatum, M. crinitum, M. c. huculicum, M. aberratum and
M. saxonicum. .Species belonging to the genus Karpatophyllon seem to prefer deciduous woods reaching upwards to the
lower limit of coniferous forests. Only four species belong to this genus. They are distributed in an area that links the
North-Eastern to the Southern Carpathians (Apuseni and Poina Ruscai Mounts). These species are: K. polinski,
K. dacicum, K. carpaticum and K. banaticum. We may say that the two genera Mastigophorophyllon and
Karpatophyllon are geographically connected to the Carpathian Mountains.

CEUCA, T., 1996. — Mastigophorophyllon (Verhoeff, 1897) et Karpatophyllon Jawlowsky, 1928. genres des
Carpates (Chordeumatida, Diplopoda). In: Geoffroy, J.-J., Mauri£s, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 61-65. Paris ISBN : 2-85653-502-X.

TR1AN CEUCA

INTRODUCTION

Les deux genres qui font l'objet de notre analyse constituent, avec quelques autres, la
famille des Mastigophorophyllidae, et se distinguent nettement, tant du point de vue
morphologique (notamment par 1'aspect des gonopodes) que du point de vue geographique, des
especes se repartissant en Europe centrale et orientale. Le genre Mastigophorophyllon comprend
des formes vivant generalement en haute altitude, surtout dans les prairies alpines, parfois a la
lisiere des forets de coniferes et plus rarement a celle des forets de feuillus. On les trouve dans la
litiere, au bord des sentiers ou des routes forestieres. Certaines especes se rencontrent de
preference, dans les prairies alpines, sous les touffes d'herbe ou autour des quelques rares
coniferes presents. Pendant les periodes de secheresse prolongee, elles recherchent l'humidite
que conservent les branches de genevrier etendues au ras du sol (cf. STOJALOWSKA, 1961 ;
Tabacaru, 1990 ; VERHOEFF, 1900).

RESULTATS

Dans un travail publie en 1976 sur le genre Mastigophorophyllon, j'ai montre que les deux
sections creees par VERHOEFF, sur la base de la presence ou de l'absence d'un rameau
plumiforme sur la partie posterieure des gonopodes anterieurs, peuvent avoir valeur de sous-
genres. Notons que cinq des six especes du sous-genre Mastigophorophyllon depourvues de ce
rameau sont uniquement distributes dans les Carpates meridionales (Alpes de Transylvanie) ; ce
sont :

Mastigophorophyllon (M.) alpivagum (Verhceff, 1897) des Monts de Cindrel.

Mastigophorophyllon (M.) deuheli Verhceff, 1898 des Monts de Bucegi.

Mastigophorophyllon (M.) transsilvanicwn Attems, 1900 des Monts de Bucegi.

Mastigophorophyllon (M.) carpaticum Ceuca, 1976 des Monts de Retezat (FIG. 1 A).

Mastigophorophyllon (M.) banarescui Ceuca, 1976 des Monts de Retezat.

Ces endemismes peuvent etre dus a la fragmentation de la chaine carpatique par des vallees
transversales, dont le resultat est l'isolement paleogeographique, sous forme d'llots, des zones
de haute altitude. Beaucoup d'autres diplopodes strictement localises, ainsi que des especes
endemiques fort diverses, doivent avoir d'ailleurs la meme origine.

La seule espece connue en dehors de cette aire carpatique, est M. (M.) hohemicum Attems,
habitant la Boheme (Attems, 1900) ; sa presence dans une region aussi eloignee est
difficilement explicable.

Une distribution tout aussi etroitement delimitee caracterise le second sous-genre,
P aramastigophorophyllon (pourvu de rameau plumiforme), dont huit des dix formes connues
sont cantonnees dans les Carpates du Nord et les Carpates Orientales. Ce sont :

Mastigophorophyllon (P.) penicilligerum Verhceff, 1899 des Monts Rodna.

Mastigophorophyllon (P.) cirriferum Verhceff, 1899 des Monts de Tatra.

Mastigophorophyllon (P.) jickelii Verhceff, 1900 de Borsec.

Mastigophorophyllon (P.) serrulatum Attems, 1926 des Monts Rarau.

Mastigophorophyllon (P.) s. apiculatum Jawlowski, 1935 des Carpates de l'Ukraine.

Mastigophorophyllon (P.) crinitum Attems, 1926 de Virghis.

Mastigophorophyllon (P.) c. huculicum Jawlowski, 1935 des Carpates de l'Ukraine.

Mastigophorophyllon ( P ) aberration Ceuca, 1985 des Monts de Rodna.

La position zoogeographique de M. (P.) bulgaricum Schubart, 1939, et de sa sous-espece
M. (P.) b. pirinicum est tres interessante (GULICKA, 1967). On trouve les deux formes dans les
Balkans, a la limite sud de la repartition du genre Mastigophorophyllon , alors qu'il n'y a aucun
representant du sous-genre Paramastigophorophyllon le long des Carpates meridionales.

La seule espece qui occupe une aire tres vaste, depassant largement l'aire carpatique, est
M. (P.) saxonicum Verhceff, 1916. Elle est frequente en Allemagne, ou elle a ete trouvee dans

Source :

CHORDEUMATIDA DES CARPATHES

de nombreuses localites (SCHUBART, 1934), certaines probablement de faible altitude ; elle a ete
egalement signalee en Lettonie, Estonie, Pologne, Slovaquie, dans l'ouest de l'Ukraine et en
Roumanie (dans les Carpates du nord du pays). Avec une aussi large repartition, il semble
naturel de constater, chez cette espece, une certaine variability de la morphologic des gonopodes,
d'autant plus marquee que les individus proviennent des confins orientaux et occidentaux de son
aire geographique, ce qui peut etre interpret^ comine une distribution le long d'un cline.

Fig. 1. — A : exemple de gonopode anterieur sans rameau plumiforme sur la face posterieure, M. (M.) carpaticum ; B :
gonopode anterieur avec rameau plumiforme (r.), M. (P.) serrulalum ; C : gonopode anterieur, en vue posterieure,
de K. dacicum. (d'apres ATTEMS, 1926 ; Ceuca, 1964. 1976).

FIG. I. — A: anterior gonopod without "feather-like" branch on the posterior side, M. (M.) carpaticum ; B: anterior
gonopod with "feather-like’ branch (r). M. (P.) serrulalum; C: anterior gonopod, posterior view, K. dacicum.

Dans le travail deja mentionne ci-dessus (CEUCA, 1976), j’ai soutenu que le genre
Mastigophorophyllon faisait defaut dans les Monts Apuseni situes en Transylvanie, a l’interieur
de l'arc carpatique. J'ai cependant identifie, a la demande d'un collegue, des restes de
diplopodes, plus ou moins digeres, trouves dans l'estomac d'un lezard (Lacerta vivipara ) capture
dans les Monts Apuseni. Parmi les debris figurait un septieme anneau comprenant des
gonopodes intacts pouvant appartenir a M. (P.) saxonicum. Des recherches ulterieures effectuees
dans les memes montagnes m'ont fourni des exemplaires captures dans trois autres stations. Les
particularites morphologiques de leurs gonopodes pourraient justifier la creation d'une sous-
espece nouvelle dont l'etude constituera le sujet d’un travail particulier. II faut egalement
remarquer que les stations en question se situent a la lisiere de forets situees a de plus basses
altitudes et constitutes d'un melange de coniferes et de feuillus.

En ce qui concerne le troisieme sous-genre, Metamastigophorophyllon, dont la seule
espece connue actuellement est M. (M.) giliarovi Lang, 1959 du Caucase (Krasnaia Poliana),
une revision detaillee de la morphologie externe et des gonopodes parait necessaire afin de
demontrer son appartenance au genre Mastigophorophyllon .

L'autre genre qui fait l'objet de notre attention, Karpatophyllon, a ete cree par
JAWLOWSKY en 1928 lors de la description de K. polinskii , espece decouverte en Ukraine

TR1AN CEUCA

(Podolie, Collines du Prut) et retrouvee. plus tard, dans trois stations des Carpates du nord de la
Roumanie (Monts de Rodna). Ce genre renferme a ce jour trois autres especes, reparties, elles
aussi, dans le perimetre carpatique :

Karp atophy lion polinskii Jawlowski, 1928, Ukraine et Monts Rodna.

Karpatophyllon dacicum Ceuca, 1964, des Monts Apuseni (Fig. 1C).

Karpatophyllon carpaticum Ceuca, 1985 des Monts du Lapus.

Karpatophyllon banaticum Ceuca. 1989 des Monts Poiana Ruscai.

De ce qui precede, il s'ensuit que la repartition geographique du genre Karpatophyllon
permet de relier les Carpates du Nord-Est aux Carpates meridionales par l'intermediaire des
Monts Apuseni et des Monts Poiana Ruscai, ces demiers flanquant vers le Nord-Ouest les Monts
de Retezat ; on voit done cette repartition circonscrire le Plateau de la Transylvanie, du cote Ouest
(Fig. 2).

Fig. 2. — Repartition des genres Mastigophorophyllon et Karpatophyllon. 1 : M. (M.) alpivagum ; 2 : M. (M.)
deubeli ; 3 : M. (M.) transsilvanicum ; 4 : M. (M.) bohemicum ; 5 : M. (M.) carpaticum ; 6 : M. (M.) banaticum ;
7 : M. (P.) penicilligerum ; 8 : M. (P.) cirriferum ; 9 : M. (P.) aberration ; 10 : M. (P.) jickelii ; \ \ : M. (P.)
saxonicum ; 12 : M. (P.) serrulatum ; 13 : 14. (P.) s. apiculatum ; 14 : M. (P.) crinitum ; 15 : M. (P.) c. huculicum ;
16 : M. (P.) bulgaricum ; 17 : M. (P.) b. pirinicum ; 18 : K. polinskii ; 1 9 : A*, dacicum ; 20 : K. carpaticum ; 21 :
K. banaticum.

FlG. 2. — Distribution of the genera Mastigophorophyllon and Karpatophyllon. I: M. (M.) alpivagum; 2: M. (M.)
deubeli; 3: M. (M.) transsilvanicum; 4: M. (M.) bohemicum; 5: M. (M.) carpaticum; 6: M. (M.) banaticum; 7; M.
(P.) penicilligerum; 8: M. (P.) cirriferum; 9: M. (P.) aberratum; 10: M. (P.) jickelii; II: M. (P .) saxonicum; 12:
M. (P.) serrulatum; 13: M. (P.) s. apiculatum; 14: M. (P.) crinitum; 15: M. (P.) c. huculicum; 16: M. (P.)
bulgaricum; 17: M. (P.) b. pirinicum; 18: K. polinskii; 19: K. dacicum; 20: K. carpaticum; 21: K. banaticum.

Les especes du genre paraissent preferer les forets de feuillus, s’elevant en altitude jusqu’a
la liinite inferieure des forets de coniferes.

Source :

CHORDEUMATIDA DES CARPATHES

CONCLUSION

On peut affirmer que les genres Mastigophorophyllon et Karpatophyllon sont lies, du point
de vue geographique, a la chaine carpatique d'ou ils sont issus, se sont diversifies et ont etendu
leur aire de repartition. II apparait egalement que la presence ou l'absence du rameau plumiforme
des gonopodes anterieurs chez Mastigophorophyllon n'a pas du jouer un role tres important.

REFERENCES

ATTEMS, C. 190a — Ueber der Farbung von Glomeris und Beschreibung neucr Oder weniggekanter Myriopoden. Arch.
Naturg., LXVI : 313-316.

Attems, C., 1926. — Uber palaarktischer Diplopoden. Arch. Naturg., Abt. A. H., 1-2 : 82-108.

CEUCA r 1959. Genurile Karpatophyllon si Stenophyllum in fauna dc Diplopodc a Romaniei. Stud. Univ. B. B. ser
Biol., XXXIV : 52-56.

Ceuca, T., 1964. — Citeva Diplopodc noi in fauna RPR. Stud. Univ. B. B. ser. Biol., XXXIX : 89-92.

CEUo A; T¥’J976 “ Genul Mastigophorophyllon Verh. 1897 (Diplopoda-Ascospermophora). Stud. Univ. B. B. ser.
Biol. , LI : 37-43.

Gulicka, J., 1967. — Neue und interessante Diplopoden aus Bulgarien. Annotat. Zool. Bot., 39 : 1-3.

Jawlowsky H 1928. — Karpatophyllum polinskii n. sbg. n. sp., Leptoiulus czarnohoricus n. sp. (Diplop.). Ann
Mus. Zool. Polonici, 7 : 102-106. K K

Schubart, O., 1934. — Tausend fussier Oder Myriapoda I. Diplopoda. In : F. Dahl. Tierw. Deutschl. 28 Jena G
Fischer. 1-318. ’

Stojalowska, W., 1961. — Krocionogi (Diplopoda) Polski. Warszawa. Poiska Akademia Nauk: 216 pp.

TAB,A--. ' • 1969 <l97°)- — Sur I'origine de la faune des Diplopodes des Carpates. Bull. Mus. nail. Hist. not.. 41 :

1 J7- I 4j.

Verhoeff. K. W.. 1900. — Beitrage zur Kenntnis palaarktischer Myriapoden. Arch. Naturg., LXVI : 368-369.

Source ; MNHN, Paris

Sur la remarquable conformation des apophyses
genitales males chez un polydesmide neotropical

Ionel TABACARU

Instilut de Speologie “Emile Racovitza* Str. Frumoasa Nr. 1 1, RO-781 14 Bucuresti , Roumanie

RESUME

Description dun genre nouveau de diplopodes, Venezuelodesmus n. g. (Trichopolydesmoidea. Fuhrmannodesmidae,
Venezuelodesmini n. trib.), reprSsentc par trois especes (V. orghidani n. sp., V. decui n. sp.. V. bordoni n. sp.) trouvees
au Venezuela, chez lesquelles les coxa de la deuxieme paire de panes sont modifiees en remarquables apophyses genitales
portant des telopodites reduits et surmontant un long organe musculeux evaginable.

ABSTRACT

On the noteworthy structure of male genital apophyses in a Neotropical polvdesmid millipede.

Description of a new millipede genus, Venezuelodesmus n. g. (Trichopolydesmoidea, Fuhrmannodesmidae,
Venezuelodesmini n. trib.) including three species (V. orghidani n. sp.. V decui n. sp.. V. bordoni n. sp.) found in
Venezuela and showing the coxae of the second pair of legs transformed into remarkable genital apophyses supporting
reduced telopodites and overlying a long musculous evaginable organ.

INTRODUCTION

I] est bien connu que, chez les males de diplopodes polydesmides, les canaux deferents
perforent dans leur longueur les coxae des pattes de la deuxieme paire et debouchent a l’exterieur
chacun par un gonopore situe a Tangle distal interne de la hanche. On utilise chez les diplopodes
le nom de penis mais je prefere utiliser dorenavant le nom d'apophyse genitale car, ainsi que 1'a
montre le biologiste framjais Albert VANDEL (1943) dans un cas parfaitement similaire, le nom
de penis est manifestement inexact : en effet, ces formations, constitutes par la partie terminale
des canaux deferents, avec les orifices genitaux a leur extremite, ne jouent jamais le role
d’organe d'intromission.

En tout cas, chez les polydesmides, il s'agit d'un simple entonnoir situe parfois sur une
proeminence et entoure souvent de quelques soies (Fig. 1A). Cependant, en examinant des
petites formes de polydesmides recoltees au Venezuela par le regrette professeur Traian
Orghidan, M. Carlos BORDON, de Caracas, et mon ami V. DECU, j'ai eu la surprise de trouver
trois especes nouvelles, appartenant a un genre nouveau, chez qui les coxae de la deuxieme paire
de pattes, porteuses de telopodites reduits, sont modifiees en de remarquables apophyses
genitales. J'ai donne a ce genre le nom de Venezuelodesmus n. g. et aux trois nouvelles especes

Tabacaru, I., 1996. — Sur la remarquable conformation des apophyses gdnitales males chez un polydesmide
neotropical. In: Geoffroy, J.-J., Mauri£s, J.-P. & Nguyen Duy - Jacquemin. M.. (eds). Acta Myriapodologica. Mem.
Mns. nain. Hist. not.. 169 : 67-72. Paris ISBN : 2-85653-502-X.

IONEL TAB ACARU

les noms respectifs de : V. orghidani n. sp., V. bordoni n. sp. et V. decui n. sp. (Figs. 1 &
2).

Fig. 1. — A. Banat ode smus jeanneli (Tabacaru. 1980), patte de la 2eme paire. B et C, Venezuelodesmus decui n. g., n.

sp. : B, seconde paire de pattes ; C, la 2e paire de pattes sur l'organe musculeux evaginable.

Fig. I. — A. Banatodesmus jeanneli ( Tabacaru , 1980), second pair of legs. B and C, Venezuelodesmus decui n. g., n. sp.
: B: P2; C, P2 and musculous evaginable organ.

Les trois especes du genre Venezuelodesmus n. g. sont caracterisees par les hanches ou
coxae, tres longues et robustes de la 2eme paire de pattes. Ces coxae sont accolees et forment
ensemble une languette legerement elargie distalement et pourvue, sur la face orale, dans sa
moitie proximate ainsi que le long de ses bords externes, de soies robustes dirigees vers l'apex.
A l'apex, il y a quatre soies longues et plus robustes, surtout les deux laterales. Les deux coxae
se terminent en crochets diriges oralement et les orifices genitaux se trouvent a la base de ces

Source : MNHN, Paris

APOPHYSES GENITALES MALES CHEZ UN POLYDESMIDE NEOTROPICAL

crochets. Sur chaque coxa, dans la moitie basale, face caudale, sont inseres les telopodites
reduits de la 2eme paire de pattes. Ces telopodites sont constitues de six articles courts, surtout le
dernier. L’avant-demier article (le tibia) porte une tres longue soie (Fig. IB & C).

Fig. 2. — A. Venezuelodesmus bordoni n. sp., tete et les deux premiers tergites. B el C. Venezuelodesmus decui n. sp. :

B, gonopode gauche, en vue exteme ; C, gonopode gauche, en vue caudale.

FiG. 2. — A. Venezuelodesmus bordoni n. sp.. head and two first tergites. B and C. Venezuelodesmus decui n. sp.: B. left
gonopod, external view; C. left gonopod. caudal view.

Le sternite de la deuxieme paire de pattes n'est pas soude directement a fare pleuro-tergal
du troisieme segment car la languette constitute par les deux coxae accolees surmonte un organe
allonge, tres musculeux, a paroi membraneuse, qui presente une partie basale cylindrique et une
partie distale en forme de tronc de cone. Cet organe musculeux appartient au troisieme segment
mais il s'evagine entre le bord du 2eme segment et le bord du 3eme segment. Le bord caudal de la
partie ventrale du 2eme segment est profondement echancre et les lobes encadrant l'echancrure
sont tres saillants. L'organe musculeux evaginable est dirige soit en avant, et dans ce cas les
apophyses genitales depassent la tete, soit en arriere, entre les pattes des segments suivants
(Fig. 2A).

VENEZUELODESMUS N. G.

Polydesmida de taille tres petite (5 - 5,5 mm) ; c? et 9, 20 segments. Coloration
completement blanche ; teguments granuleux. Partie orale du corps non retrecie.

IONEL T ABACARU

Tete globuleuse. Antennes relativement longues ; 6eme antennomere nettement plus long
que le 5eme. Labre tridente : mandibules prolongees vers la partie ventrale pax- une grande lamelle
ovale, crenelee du cote anterieur ; gnathochilarium avec les stipes pourvus dans leur moitie
basale de nombreuses soies robustes ; dans l'angle distal interne des lamelles linguales se
trouvent deux styles a deux ou trois pointes emoussees.

Collum moins large que la tete, relativement long, de forme trapezoidale, a angles arrondis
et bords lateraux convexes, a surface garnie de trois rangees de soies claviformes.

Tergites convexes ; carenes peu saillantes a bords lateraux regulierement arques, sans
denticulations ; surface des tergites avec trois rangees de soies claviformes. Limbe a dents
courtes et pointues.

Le deuxieme segment du <f est plus grand ; dans sa partie ventrale il est profondement
echancre du cote caudal et les lobes encadrant l'echancrure sont tres saillants ; pas de pore
pleurotergal.

Formule des pores : 5, 7, 9, 10, 12, 13, 15-19.

Pattes sans denticulations sur les bords internes des articles. Premiere paire a tarse pourvu,
sur le bord interne, d'un peigne de soies alignees plus fortes que les autres soies ; un groupe de
fortes soies se trouve aussi sur la face orale du femur.

Deuxieme paire a sternite non soude avec la partie ventrale du 3eme segment ; coxae
immenses, accolees en une languette qui surmonte un long organe musculeux evaginable entre le
2eme et le 3eme segment ; telopodites reduits.

Dans la partie ventrale du Seme segment, sur le sternite de la 5eme paire de pattes ou sur les
stemites des 4eme et 5eme, il y a des processus portant des denticulations ou de longues epines.

Gonopodes : coxoides grands, globuleux ; les telopodites diriges obliquement vers la ligne
mediane s'entre-croisent. La zone prefemorale, en bourrelet pileux, est elargie dans la partie
caudale ou penetre le crochet coxal. Le telopodite, profondement divise, comprend d'une part,
une branche tarsale, longue et grele, recourbee vers l'arriere, et d'autre part une branche tibiale
plus courte, arquee en faucille, a partie basale large ; de cette partie basale se detache la branche
seminale. La rainure seminale est bien visible et son trajet est direct (Fig. 2B & C).

Espece type du genre Venezuelodesmus n. g. : V. decui n. sp.

CLE DES TROIS ESPECES DE VENEZUELODESM US

1 (2) Gonopodes : coxoi'de pourvu d'une proeminence anguleuse au bord distal
posterieur ; la region prefemoro-femorale se prolonge en un lobe arrondi et aplati, connecte, par
une lame, avec le solenomerite. Dans la partie ventrale du 5eme segment, sur le sternite des 4eme
et 5eme paires de pattes, se trouve un grand processus en fer a cheval pourvu de denticulations

. . . V. bordoni n. sp.

Localite type : Parque National Rancho Grande (Henri Pittier) (Station 44 in Decu, Bordon & Linares, 1987), 16-19. XI.
1982, 1000-1400 m, litiere, 3 & cf , 2 99, leg. V. Decu, C. Bordon & T. Orghidan.

2(1) Gonopodes : coxoi'de arrondi, sans proeminence anguleuse ; pas de prolongement
femoral. Dans la partie ventrale du 5eme segment il y a des processus qui n'ont pas la forme d'un
fer a cheval . 3

3 (4) Gonopodes : les deux longues soies orales du coxoi'de sont inserees dans une piece
en forme de coupe ; solenomerite sans eperon ; branche tarsale uniformement arquee a son
extremite. Dans la partie ventrale du 5emc segment, sur les stemites des 4eme et 5emc paires de
pattes, se trouvent deux processus longitudinaux paralleles, pourvus de denticulations

. . V. decui n. sp.

Locality type : Cerro La Pastora, Capadare, Edo Falcon (Station 51 in Decu, Bordon & Linares, 1987), 13. XI. 1982,
L4iere, 4 c? c? . 6 99, leg. V. Decu & C. Bordon (dont 1 c? et 1 9 paratypes ddposds au M.N.H.N. de Paris sous le n° JC

Source :

APOPHYSES GENITALES MALES CHEZ UN POLYDESMIDE NEOTROPICAL

4 (3) Gonopodes : les deux longues soies orales du coxoide sont inserees sur la surface de
celui-ci ; solenomerite pourvu d un eperon pointu ; branche tarsale brusquement recourbee en
crochet a son extremite. Dans la partie ventrale du 5cme segment il y a un seul processus
transversal sur le sternite de la 5cme paire de pattes, , pourvu de nombreuses et longues epines
. V. orghidani n. sp.

Local ite type : Route vers la grotte Cucva del Tigre, Cerro la Passora, Edo Falcon, 12.X1.1982, 1 <? , leg. T. Orghidan.

SUR LA POSITION SYSTEMATIQUE DU GENRE VENEZUELODESMUS N. G.

Le nouveau genre Venezuelodesmus n. g. fait partie d’un groupe de genres neotropicaux
caracterises par un habitus de type Trichopolydesmus et des gonopodes dont le coxoide, tres
grand, enveloppe un telopodite condense et de dimensions reduites (type cryptodesmoi'de).

ATTEMS (1926, 1940) a range ces genres dans la famille des Vanhoeffeniidae Attems 1914
et on a longtemps considere comme valable cette opinion. Cependant, JEEKEL (1965) a montre
que ce nom de famille, en raison de son genre type, est synonyme de Sphaerotrichopidae
Attems, 1914 et aussi de Dalodesmidae Cook, 1896.

VERHOEFF (1910, 1926-1932, 1941, 1942) a range ces genres dans la famille des
Trichopolydesmidae Verhoeff, 1910 et cette position a ete adoptee par KRAUS (1957, 1959,
1960). par LOOMIS (1964) et par SHEAR (1973). Dans des travaux relatifs aux representants
europeens de la famille dcs Trichopolydesmidae (TABACARU, 1975, 1980), nous avons
considere cette famille dans le sens de VERHOEFF.

BROLEMANN (1916) a considere que ces genres appartenaient a la famille des
Cryptodesmidae Karsch, 1879 et les a classes a part dans la tribu des Fuhrmannodesmini
Brolemann 1916. Cette opinion semble etre soutenue par notre collegue Maurles (-1983) car il
parle de “Cryptodesmides trichopolydesmiformes”.

D'apres HOFFMAN (1980) ces genres appartiennent a la famille des Fuhrmannodesmidae
Brolemann, 1916, mais dans le cadre de la super-famille des Trichopolydesmoidea Verhoeff,
1910. Cette position a ete soutenue par SlMONSEN (1990) dans son etude cladistique des
Polydesmida. GOLOV ATCH (1986) a aussi accepte la famille des Fuhrmannodesmidae.

Dans un travail concernant des Fuhrmannodesmidae de la region d'Amazonie (Bresil),
GOLOVATCH (1992) a decrit une nouvelle espece qu'il a attribute au genre Cutervodesmus
Kraus, 1957 et qui semble presenter sur les P.2 une conformation similaire a celle que nous
avons trouvee chez Venezuelodesmus. Cependant, notre collegue GOLOVATCH ne dit rien du
long organe qui s'evagine entre les segments 2 et 3. En outre, les trois especes du genre
Venezuelodesmus n. g. different de l'espece decrite du Bresil par trois caracteres :

1 ) les mandibules prolongees par une grande lamelle crenelee,

2) des processus sur la partie ventrale du 5eme segment,

3) la presence d'une branche seminale sur les gonopodes.

Considerant l'ensemble des genres reunis dans la famille des Fuhrmannodesmidae
(BROLEMANN, 1916), il nous semble que cette immense famille, apparemment heterogene, est
mal definie et probablement polyphyletique. En tout cas, une revision de ces genres parait
necessaire ainsi que la description de nouveaux taxons, qui meneront sans doute a une nouvelle
definition des sous-familles et des tribus.

Tenant compte des remarquables caracteres du nouveau genre, Venezuelodesmus n. g.,
nous proposons pour celui-ci une tribu a part, la tribu Venezuelodesmini nov. trib.

REFERENCES

Attems, C. , 1926. — Myriopoda. In : W. KOkenthal & T. Krumbach, Handbuch der Zoologie, 4. Progoneata,
Chilopoda, Insecta , Berlin & Leipzig, W. de Gruyter & C° : 1-402.

Attems, C., 1940. — Myriapoda 3. Polydesmoidea III. In : F. E. Schulze. W. Kukenthai. & K. Heider, Das Tierreich,
70. Berlin & Leipzig. W. de Gruyter & C° : 1-577.

IONEL TABACARU

BrClemann. H. W., 1916. — Essai de classification des Polydesmicns (Myriapodes). Ann. Soc. Entom. France. 84 :
523-608.

Decu, V.. Bordon, C. & Linares O., 1987. — Las estaciones de America del Sur de donde ha sido colectado el material
zoologico que esta en presente en estudio en el Instituto de Espeleologia de Bucarest (Romania). Situacion del
material. In . Fauna hipogea y hemiedafica de Venezuela y oiros paises de America del Sur. I. Bucuresti, Ed. Acad. :
29-45.

Golovatch. S. 1., 1986. — Diplopoda from the Nepal Himalayas: Polydesmidae, Fuhrmannnodesmidae.
Senckenbergiana biol.. 66 : 345-369.

Golovatch, S. I., 1992. — Review of the Neotropical fauna of the millipede family Fuhrmannodesmidae, with the
description of four new species from near Manaus, Central Amazonia. Brazil (Diplopoda, Polydesmida).
Amazoniana, Kief 12 : 207-226.

Hoffman. R. L.. 1980. — Classification of the Diplopoda. Geneve, Museum d'Histoire Naturelle, (1979), 237 pp.
Jeekel, C. A. W., 1965. — The identity of Dalodesmus tectus Cook. 1896, and the status of the family names
Dalodesmidae Cook, 1896, Vanhoeffeniidae Attems, 1914 and Sphaerotrichopodidae Attems, 1914 (Diplopoda,
Polydesmida). Entom. Bericht., 25 : 236-239.

Kraus, O., 1957. — Myriapoden aus Peru, V. Senck. biol.. 38 : 95-1 14.

Kraus, O., 1959. — Myriapoden aus Peru, VII. Senck. biol.. 40 : 191-208.

Kraus, O., 1960. — Myriapoden aus Peru, IX. Senck. biol.. 41 : 241-264.

LOOMIS, H. F., 1964. — The Millipeds of Panama (Diplopoda). Fieldiana Zoology, 47 : 1-136.

Mauries, J. P.. 1983. — Le genre Galliocookia Ribaut, 1954. Deux especes nouvelles des grottes de l'Ardeche et du Gard
(Myriapoda, Diplopoda. Polydesmida). Bull. Soc. Hist, nat., Toulouse, 119: 103-110.

Shear, W. A., 1973. — Millipeds (Diplopoda) from Mexican and Guatemalan caves. Subterranean Fauna of Mexico,
Acad. Nazionalc Linceix 171 : 239-305.

SlMONSEN, A., 1990. — Phylogeny and biogeography of the Millipede Order Polydesmida, with special emphasis on the
Suborder Polydesmidea. Thesis, Bergen, Mus. Zool. Univ., 114 pp.

Tabacaru, I., 1975. — Napocodesmus florentzae n. sp. (Diplopoda. Polydesmida). Trav. Inst. Speol. E. Racovitza,
14 : 71-82.

Tabacaru, I.. 1980. — Trichopolydesmus (Banatodesmus) jeanneli n. sg., n. sp. (Diplopoda, Polydesmida). Trav. Inst.
Speol. E. Racovitza, 19 : 155-161.

vandel. A., 1943. — Essai sur forigine, revolution et la classification des Oniscoidea (Isopodes terrestres). Bull. biol.
Fr. Belg., Suppl. 30 . 1-136.

\ erhoeff. K. W., 1910. — 4. Uber Diplopoden 42. Aufsatz : Neue Polydesmiden aus Mitteleuropa und ihre Verwandten.
Zool. Anz.. 36 : 132-145.

Verhoeff, K. W., 1926-1932. — Diplopoda 1 & 2. In : H. G. Bronns Klassen und Ordnungen des Tierreichs, 5, Leipzig,
Akademische Verlagsgesellschaft : 1-2084.

VERHOEFF, K. W., 1941. — Hohlen-Diplopoden aus dem Trentino. Zeits. f. Karst, u. Holden. : 179-189.

Verhoeff. K. W., 1942. — Chilopoden und Diplopoden. hi : Beitrdge zur Fauna Perus I, Hamburg : 5-72.

Source :

Records of Paradoxosomatid Millipedes of India

Kubra Bano

Department of Zoology, University of Agricultural Sciences, G.K.V.K., Bangalore, 560065, India.

ABSTRACT

A review of the family Paradoxosomatidae along with a list of genera and species so far recorded from India has been
brought in this short paper.

RESUME

Ce travail presente une revue taxinomique de la famille Paradoxosomatidae, accompagnee d'une liste des genres el
especes actuellement repertories en Inde.

INTRODUCTION

The family Paradoxosomatidae was first proposed by Daday (1889) for the two genera of
the order Polydesmida, Trachydesmus and Paradoxosoma. COOK (1895) recognized the family
Paradoxosomatidae. In addition, he created a family Strongylosomatidae, a heterogenous group
that was later considered synonymous to Paradoxosomatidae (JEEKEL, 1968).

ATTEMS (1898), in his monograph of the order Polydesmida, rejected the name
Paradoxosomatidae, but recognized the family Polydesmidae in which he included the sub¬
family Strongylosominae, which included a number of genera along with Trachydesmus and
Paradoxosoma. Apart from this, he distinguished the sub-family Suliciferinae. Both the sub¬
families were quite heterogenous as are almost all of the genera that are now included and
referred to as Paradoxosomatidae. Subsequently, ATTEMS (1914), in his revised studies,
merged these two sub-families into a single family Strongylosomidae. He published his work as
a monograph in 1937 “ Das Tierreich " vol. 68. His work included a description of the genera and
species known up to 1937. This book acquired importance among the workers and became the
origin for all the subsequent studies on the order. Following this, a number of contributions
were made towards the revision, criticism and re-classification of the family Paradoxosomatidae
(Hoffman, 1953, 1961, 1963, 1964; JEEKEL, 1963 a, b).

HOFFMAN critically evaluated the classification of Ethiopian fauna and briefly reviewed the
genera. He also commented on the fauna of East Asia (HOFFMAN, 1961, 1963). JEEKEL (1963)
presented a survey of the Paradoxosomatidae of the Neo-tropical regions and his publications
dealt with the taxonomy of the Indo-Australian fauna. Further, he set right the anomaly in the
classification to a certain extent arranging the so far known genera and species of
Paradoxosomatidae according to their zoo-geographic regions (JEEKEL, 1968). He discussed

Bano, K., 1996. — Records of paradoxosomatid millipedes of India. In: Geoffroy, J.-J., Mauries, J.-P. &
Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 73-77. Paris ISBN : 2-

85653-502-X.

KUBRA BANO

and evaluated previously proposed classifications, bringing numerous changes in the generic
delimitation of the fauna of several regions, including India.

ATTEMS (1937) estimated about 490 species of millipedes of the family
Paradoxosonratidae whereas, in 1968, JEEKEL estimated 650 species. However, how many
more might have gone unsighted and remained obscure is not known. JEEKEL (1963a) stated
that "to the family Paradoxosomatidae are referable all genera included in the monograph on the
Strongylosomidae published by ATTEMS, 1937, with the exception of Aphelidesmus Brol.,
lulidesmus Silv., Antisoma Chamb., Fijiodesmus Chamb., Phyletodesmus Chamb.,
Semenellogon Chamb. and Strongylomorpha Silv.”. In the same paper JEEKEL reinstated the
name Paradoxosomatidae which, up to that time was mostly referred as Strongylosomidae or
Strongylosomatidae.

According to the present state of our knowledge, the family is the largest of the order
Polydesmida, which in turn is the largest order of the class Diplopoda.

The main character used for distinguishing Paradoxosomatidae is the presence of
unconnected gonopod coxae, which are not joined by membranous bridge as in the other
Polydesmid families. Coupled with this are the other typical features namely the unique presence
of a distinct post-femoral cingulum in the gonopods, mode of insertion of the coxal horn in the
gonopod coxa and the location of paired setae on the paraprocts.

Owing to the scantiness of the Indian faunistic studies, much remains to be done in the
way of revisionary studies of the described species. ATTEMS (1936) was the first to study and
describe some of the species belonging to this family under the name Strongylosomidae. Some
notes have been furnished concerning Anoplodesmus (JEEKEL, 1965), Chondromorpha
(JEEKEL, 1963a) and Sundanina (JEEKEL, 1953), but information on many genera is still
lacking. To strengthen the studies, the author has carried out this review work and has planned
to conduct a survey and studies on the family Paradoxosomatidae.

JEEKEL (1968) presented the diversity and distribution of oriental fauna and published a
consolidated list ol the fauna known till then, from which a list of the Indian paradoxosomatids
has been brought out here, for the reference of the Indian workers. The list includes millipedes
belonging to the four tribes, namely Sulciferini, Xanthodesmini, Sundanini and Polydrepanini.
Ol the tour, the first three belong to the sub-family Paradoxosomatinae and the fourth to
Alogolykinae.

CHECK-LIST OF PARADOXOSOMATID MILLIPEDES OF INDIA

List of species recorded by ATTEMS (1937) and reported by JEEKEL (1968)

1 . Orthomorpha ( Kalorthomorpha ) coonoorensis Carl, 1932

2. 0. (K). ursula Attems,1932

3. O. (K). dentata Carl, 1932

4. O. (K). almorensis Turk, 1947

5. Anoplodesmus tanjoricus (Pocock. 1892)

6. A . anthracinus Pocock, 1895

(Syn. J one spelt is splendidus Verhoeff, 1936)

7. A. insignis Attems. 1936

8. A. saussurii (Humbert, 1865)

9. A. indus (Chamberlin. 1920)

10. A. atopus (Chamberlin, 1920)

11. Chondromorpha severini Silvestri, 1897

12. C. severini var. robust i Attems, 1936

13. C. mammifera Attems, 1936

14. C. kelaarti (Humbert, 1865)

15. C. kelaarti sub. sp. valparaiensis (Carl, 1932)

Source :

RECORDS OF INDIAN PARADOXOSOMATID MILLIPEDES

16. C. kelaarti sub. sp. longipes (Verhoeff, 1936)

17. C. kaimura Turk, 1947

18. Paranedyopus subcylindricus Carl, 1932

19. Himantogonus rufocinctus (Carl, 1932) Comb. nov.

20. Streptogonopus phipsoni (Pocock. 1892)

(Syn. Strongylosoma contortipes (Attems, 1898)

21. S. nitens Attems, 1936

22. S. jerdani (Pocock, 1892)

23. Sundcmina nulla Attems, 1936

24. S. laevisulcata Carl, 1932

25. S. hirta Carl, 1932

26. S. contortipes (Schubart, 1935)

27. S, granulifera Attems, 1936

28. S. bimontana Carl, 1932

29. S. trifida Carl, 1941

30. S. pumila Attems, 1944

31. S. septentrionalis Turk, 1947

32. Dasypharkis rugulosa (Carl, 1932)

33. Polydrepanum tamilum Carl, 1932

34. P. implicatum Carl, 1941

35. Telodrepanum badaga Carl, 1932

3 6 . Grammorhabdus asperrimum Carl ,1932

37. Xiphidiogonus spinipleurus Carl, 1932

38. X. dravidus C arl, 1932

39. X. hendersoni Carl, 1932

40. Gyrodrepanum contortipes (Carl, 1932) Comb. nov.

41. Kaschmiriosoma contortipes (Schubart, 1935)

From the above list JEEKEL, in the same work, pointed out that the
"Orthomorpha" coonoorensis, "O". almorensis, "O". dendata, “ Polydrepanum ” implicatum,
“ Sundanina ” granulifera, “5”. trifida, “5”. hirta, “5”. simplex and "S". septentrionalis belonged
to unnamed genera, and stated that the allocation to definite genera could be done only after a
careful study of the pertinent material.

JEEKEL (1980) reexamined some of the Indian species of paradoxosomatids and proposed
two new genera, Parchondromorpha and Harpagomorpha of the tribe Suliciferini for the species
Orthomorpha coonoorensis (Carl, 1932) and Orthomorpha dentata (Carl, 1932) respectively. He
erected a nov. gen. for Sundanina laevisulcata (Carl, 1932) and Sundanina hirta (Carl, 1932):
the genus Antichirogonus. In the same work, he described the characteristics of the genera
Polydrepanum Carl, 1932 and Dasypharkis Attems, 1936 of the tribe Polydrepanini. He
discussed the status of the tribes Polydrepanini and Alogolykini. He also reported Desmoxytes
planata Pocock from the Andamans.

The following are the Indian paradoxosomatids reported by JEEKEL (1980):

1 . Paranedyopus rufocinctus (Carl, 1932)

2. Paranedyopus subcylindricus (Carl, 1932)

3. Paranedyopus simplex (Humbert, 1865) new comb.

4. Paranedyopus Ursula (Attems, 1936) new comb.

5. Parchondromorpha coonoorensis (Carl, 1932)

6. Harpagomorpha dentata (Carl, 1932)

7. Antichirogonus laevisulcatus (Carl, 1932)

8. Antichirogonus hirtus (Carl, 1932)

9. "Kronopolites" unicolor Attems, 1936

10. "Kronopolites” spiniger Attems, 1936

1 1. " Strongylosoma " montigena Carl, 1935

1 2. Dasypharkis Attems, 1936 (2 sp.)

KUBRA BANO

13. Gyrodrepantun Carl, 1932 (1 sp.)

14. Polydrepanum Carl, 1932 (2 sp.)

(Syn. Grammorhabdus Carl, 1932)

15. Telodrepanum Carl, 1932 (1 sp.)

16. Xiphidiogonus Carl, 1932 (3 sp.)

17. " Polydrepanum " implication Carl, 1941

1 8 . “ Sundanina ” granulifera Attems, 1 936

1 9 . “Sundanina “trifida Carl, 1 94 1

20. Desmoxyies planaia (Pocock, 1895)

(Syn. Prionopeltis planatus Pocock, 1895)

JEEKEL listed the genera that are under inverted commas above, as incertae sedis and stated
that these required reexamination of the gonopods for proper allocation to their genera.

GOLOV ATCH (1984) examined the millipedes collected from India by Dr. G. TOPAL of the
Hungarian Natural History Museum, Budapest, in 1967, and discovered some very important
specimens of paradoxosomatids. He distinguished 16 species belonging to 13 genera of this
family. Among these, 9 were found to be new to science. He erected 8 new genera and
synonymised one. His work presented the description and allocation of the new taxa established
by him.

The following is the list of paradoxosomatids which Dr. GOLOV ATCH listed from the
collection of Dr G. TOPAL.

A List of paradoxosomatid millipedes of India
(Reported by S. I. GOLOV ATCH, 1983, 1984)

1. Kaschmiriosoma contortipes Schubart, 1935

2. Chondromorpha mammifera Attems, 1936

3. Kronopeltis occidentalis Golovatch, 1983

4. Topalosoma setiferum sp. nov. Golovatch, 1984

5. Curiosoma bispinosum sp. nov. Golovatch, 1984

6. Polydrepanum horridum sp. nov. Golovatch, 1984

7 . Hindornorpha (= Sundanina ) granulifera (Attems, 1936)

8. Parchondromorpha indica sp. nov. Golovatch, 1984

9. Parchondromorpha similis sp. nov. Golovatch, 1984

10. A rmolites spiniger (Attems, 1936)

1 1. Laterogonopus simplex sp. nov. Golovatch, 1984

12. Substrongylosoma distinctum sp. nov. Golovatch, 1984

13. Substrongylosoma falcatum sp. nov. Golovatch, 1984

14. Himalomorpha montigena (Carl, 1935)

15. Paranedyopus cylindricus comb. nov. (Carl, 1935)

16. Paranedyopus elongissimus sp. nov. Golovatch, 1984

The above lists constitute a record of the Indian paradoxosomatid millipedes reported

so far.

ACKNOWLEDGEMENTS

The author thanks Dr. C. A. W. Jeekel, Amsterdam. Netherlands and Dr. S. 1. Golovatch of the Institute of
Evolutionary Morphology and Ecology of Animals, Russian Academy of Sciences. Moscow, for their help in providing
the literature on Paradoxosomatidae. She also thanks Dr. J.-J. Geoffroy of M.N.H.N.. Paris for his help in the
presentation of the results during the 9th International Congress of Myriapodology, Paris. France, July 1993.

REFERENCES

Attems, C„ 1898. — System der Polydesmiden 1. Teil. Denkschr. K. Akad. Wiss. Wien (Math. Naturwiss cl.),
LXX VII ; 221-482.

ATTEMS, C., 1914. — Die Indo-Australischen Myriopoden. Arch. Nat. Abt. AH , 80 : 1-398.

Source : MNHN, Paris

RECORDS OF INDIAN PARADOXOSOMATID MILLIPEDES

Attems, C., 1936. — Diplopoda of India. Mem. Ind. Mus ., 11.

ATTEMS, C., 1937. — Myriapoda 3. Polydesmoidea. I. Fam. Strongylosomidae. In : F. E. SCHULZE, W. KOkenthal & K.
Heider, Das Tierreich, 68, Berlin & Leipzig, W. de Gruyter & C° : 1-300.

Cook, O. F., 1895. — Introductory note on the families of Diplopoda. In: : O. F. Cook & G. N. Collins, The
Craspedosomatidae of North America. Ann. New- York Acad. Sci., 9 : 1-8.

Daday, E., 1889. — Myriopodie estranea Musaci nationalis Hungarici. Termeszetr. Fiiz ., 12 : 115-156.

Golovatch, S. I., 1983. — Two Paradoxosomatidae from the Kashmir, Himalayas (Diplopoda). Senckenhera Biol 63
: 297-302.

Golovatch, S. I., 1984. — Some new or less known Paradoxosomatidae (Diplopoda: Polydesmida) from India. Acta
Zoologica Hungarica , 30 : 327-352.

Hoffman, R. L., 1953. — Scolodesmus and related African millipede genera (Polydesmida : Strongylosomatidae). Proc.
Biol. Soc. Wash., 66 : 75-84.

Hoffman, R. L., 1961. — Two new Diplopod genera from Western China (Polydesmida : Strongylosomatidae). Ann
Mag. Nat. Hist., 13 : 533-543.

Hoffman, R. L., 1963. — A contribution to the knowledge of Asiatic Strongylosomoid Diplopoda (Polydesmida:
Strongylosomatidae). Ann. Mag. Nat. Hist., 13 : 577-593.

Hoffman, R. L., 1964. — Uber einege Ostafrikanishe Diplopoda Polydesmida der zoologischen Statsammlung
Miinchen. Opusc. Zool. Munchen.,19 : 1-10.

Jeekel, C. A. W., 1953. — Two new Strongylosomidae from Indochina (Diplopoda, Polydesmidae). Beaufortia , 2 : 1-8.
JEEKEL, C. A. W., 1963a. — Diplopoda of (1-5) slud. Fauna Surinam . 4 : 1-157.

Jeekel, C. A. W.. 1963b. — Paradoxosomatidae from Borneo (Diplopoda: Polydesmida). Tijdschr. Ent., 106 : 205-283.
Jeekel, C. A. W., 1965. — A revision of the Burmese Paradoxosomatidae (Diplopoda, Polydesmida) in the Museo
Civico di Storia Naturale at Genova (Part I). Tijdschr. Ent., 108 : 95-144.

Jeekel, C. A. W., 1968. — On the classification and geographical distribution of the family Paradoxosomatidae
( Diplopoda - Polydesmida). Amsterdam, 162 pp.

Jeekel, C. A. W.. 1980. — On some little known Paradoxosomatidae from India and Ceylon, with the description of four
new genera (Diplopoda: Polydesmida). Beaufortia, 30 : 163-178.

Source : MNHN. Paris

Systematics and Biogeography of Ctenophilus Cook,
1898. A Genus of Centipedes with Disjunct
Distribution (Geophilomorpha, Schendylidae)

Luis A. Pereira

Museo de La Plata. Paseo del Bosque s/n, 1900-La Plata, Argentina

ABSTRACT

Among all known genera ot Schendylidae Ctenophilus Cook. 1898 is the only one characterized by having the
pleurites of the second maxillae fused with the posterior border of the coxosternum (apomorphic state of the character).
In all the remaining genera of the family the pleurites are not fused (plesiomorphic state of the character).

This genus has a wide distribution in Africa, with twelve species known to date. It is also present (but much less
widespread) in the Neotropical Region with one species in the Caribbean area.

A historical summary is provided for the genus, as well as observations on the taxonomic significance of various
characters heretofore utilized to distinguish genera of Schendylids.

Ctenophilus amieti (Demange. 1963), C. chevalieri (Brolemann & Ribaut. 1911), C. corticeus (Demange, 1968). C.
edentulus (Porat. 1894), C. magnus (Demange, 1963), C. nesiotes (Chamberlin. 1918), C. nitidus (Brolemann. 1926),
C. oligopodus (Demange, 1963) and C. pratensis (Demange, 1963) arc redescribed and figured from type material and/or
additional specimens and a map showing the geographical distribution of all species of the genus is included.

It is not known enough about the genus Ctenophilus and its nearest relatives to be able to confidently suggest an
explanation of the amphiatlantic pattern of distribution (which is common to some other genera of geophilomorphs
such as Schendylurus . Pectiniunguis , etc.). Plate tectonic events are considered being very evident the convinience to
develop a cladistical analysis within the Schendylids together with a biogeographical study.

It is also considered the case of the halophilous geophilomorphs. The scattered and often wide-ranging distribution of
these centipedes has been commented upon several times, specially by Cloudslky-Thompson (1948), Crabill (1960)
and Kr van (1983). Such species are very probably dispersed by rafting across very large distances, although in a very
unpredictable way. Crabill (1960) even suggested that this way of dispersal might explain trans-Atlantic disjunction
between South America and Africa. More data are obviously required and individual cases must be investigated in depth
belore we can assess the actual extent of this phenomenon and its possible occurence within Ctenophilus.

RESUME

Systematique et biogeographie de Ctenophilus Cook, 1898 ; un genre de chilopodes a aire
disjointe (Geophilomorpha, Schendylidae).

Ce travail propose une revision de fensemble du genre Ctenophilus Cook, largement repandu d’une part en Afriquc (12
especes), d’autre part dans la zone neotropicale (1 especc dans l'aire Caraibe). La revision de la systematique et de la
classification des especes composant le genre conduit a une discussion relative aux modalites de sa dispersion en deux
aires actuellement disjointes et eloignees.

Pereira, L. A., 1996. — Systematics and biogeography of Ctenophilus Cook, 1898. A genus of centipedes with
disjunct distribution (Geophilomorpha, Schendylidae). In: Geoffroy, J.-J.. Mauries, J.-P. & Nguyen Duy - Jacquemin,
M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist. not.. 169 : 79. Paris ISBN : 2-85653-502-X.

Source : MNHN, Paris

Review and Perspective of Study on Myriapodology of

China

DaqingWANG * & Jean-Paul MAURIES **

* Department of Invertebrates, Institute of Zoology
Chinese Academy of Sciences, Beijing 100080
** Museum National d’Histoire Naturelle, Laboratoire de Zoologie/Arthropodes
61, rue Buffon, F-75231 Paris, France

ABSTRACT

This contribution reviews the history and the present state of research in Myriapodology in China. It introduces all
Chinese researchers and their work in the field. Considering the present state of knowledge of Myriapoda, perspectives
and some suggestions are presented for future studies in this field in China.

RESUME

Bilan et perspectives des recherches myriapodologiques en Chine.

Ce travail passe cn revue le developpcment historique et l'etat actuel des recherches myriapodologiques en Chine. II fait
etat des travaux de tous les chercheurs chinois dans ce domaine. Un certain nombre dc perspectives sont degagees et des
suggestions sont proposees en vue de futurs travaux sur ce sujet en Chine.

INTRODUCTION

The features of Chinese zoogeography and geology are unusual and diverse.
Zoogeographically, China covers two zones: the orient and the palearctic. Physiographically,
China occupies 6.5% of the land surface of the world. The varied features make China abundant
in diversity of animal species. However, the present situation of study on Myriapoda of China
does not match in possibility provided by the fauna.

The starting point of the modern period of myriapology in China began in the late 1940s.
when the study of Taiwan diplopods commences. Studies on the Chinese mainland only began
in the late 1970s, since when a relatively prosperous period of myriapod study started. For many
years, only a small fraction of the actual China myriapod fauna and the work of Chinese
myriapodologists were known. An important reason for this is the language barrier because
many papers published by Chinese scientists were only accompanied by a brief abstract in
English. Hence the present paper is interned to introduce the current situation of Chinese
myriapodology in five sections: historical review and perspective, literature survey, collecting
localities, checklist of taxa, geographical and physiographical notes.

Wang, D. & Mauri£s. J.-P.. 1996. — Review and perspective of study on myriapodology of China. In:
Gkoffroy , J. J., Mauries. J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist,
tun.. 169 : 81-99. Paris ISBN : 2-85653-502-X.

DAQING WANG & JEAN-PAUL MAURlfeS

GEOGRAPHICAL, PHYSIOGRAPHICAL AND GEOLOGICAL NOTES

The present nation of China occupies an area of about 9.6 millions km2, of which lies on
the east of Euro-asian continent and the western coast of the Pacific Ocean. China has an ancient
and complex geology. In terms of Plate Tectonics, China basically belongs to the Eurasian-plate,
connecting with the Indo-plate in south and jointing the Pacific-plate of the Philippine-plate in the
East. The geological history of China is the result of the interactions of the three plates
mentioned above.

The Sino-Indo orogenic movement was the key factor in the formation of the Chinese
region in the early Mesozoic. From that time, the outline of the region was fundamentally
formed. From Yianshan orogenic movement to the early Tertiary, the land surface of China and
the rest of the world has been relatively stable. The surface became lower and flatter because of
chronic erosion and weathering, and the climate became warmer.

The Himalayan orogenic movement was directly responsible for the formation of the
modern physiographical environment of China during the Cenozoic. The great orogenic
movement comprised two events: the first occured from the late Oligocene to the Middle
Miocene; the second continued from the late Pliocene to the early Pleistocene, this being the
more sporadic orogen.

The second orogenic event was the most significant factor in the formation of the modern
physiographical variations of China. Under the force of the Himalayan orogenic movement, the
physiographical environment of the Euro-asian continent greatly changed: the ancient
Mediterranean sea disappeared: the Euro-asian continent jointed together; the great Tibetan
plateau emerged and became the worlds crest. Due to these events, the climate of China
consequently changed.

The elevation of the Tibet plateau, which blocked the moist winds from the ocean, resulted
in the formation of an arid physiographical environment in western China. To a large extent,
other regions succeeded the tropical or subtropical environment which formed before the
Quaternary.

HISTORICAL REVIEW AND PERSPECTIVE

Myriapods have been collected and recorded in China for more than 2000 years because
the Chinese use some species, notably centipedes, in medicine. According to traditional Chinese
medicine, large centipedes, such Scolopendra , can treat diseases, such carbuncles, scabies and
the sting of some insects. This is based on the traditional medicine theory that one poison can be
overcome by another. Hence, for a long time, the collection and study of Chinese myriapods has
focused on the medicinal use of centipedes, particularly large species. Up to now, people in
countries of southern China have a habit that treat stings of insects by using the alcohol in which
centipedes have been immersed.

The Chinese Encyclopedia of material medicine, named “Ben Cao Gang Mu”, edited in
1596, listed this as a kind of animal medicine and described its medicinal effects in detail. As a
result, some medical experts have analyzed centipede toxins using biochemical techniques.
However, the scientific study of the systematics of Chinese myriapods did not start until this
century.

The first such study dealing with Chinese myriapods dates from the late 1940s, when
Professor WANG Youxie (Yu-Hsi) commenced the study of diplopods of Taiwan. This was the
first time that Chinese myriapodologists had studied Chinese myriapods by themselves. In his
early study career, Wang sent collections of millipedes and centipedes to LOHMANDER to
identify. Later, much work was accomplished by himself towards the description and
identification of specimens from Taiwan and it adjacent islands. Most of his papers were
published in the Quaterly Journal of the Taiwan Museum. After the late 1960s his name

Source :

M YRIAPODOLOGY OF CHINA

disappeared from the literature. One of us (W. D.) asked several entomologists from Taiwan
about him, but to no avail. In 1950, the Chinese archaeologists JlA Lanpuo and LlU Xianting
discovered several fossil myriapods in Choukoutien, Beijing, and their paper was published in
the Bulletin of the Geological Society of China. This was the first time that the fossil myriapods
were reported in China.

Since the 1970s, Professor ZHANG Chong-Zhou has begun the systematic study of
mainland Chinese myriapods, including Diplopoda, Chilopoda, Symphyla and Pauropoda.
From that time, myriapodology was just known in China as a systematics owing to Professor
Zhang's outstanding work.

From 1976 to 1979, Professor ZHANG was mainly engaged in the studies of medicinal
centipedes, including their ecological habitats, individual development and breeding. In 1977,
ZHANG reported a new species of spirostreptoid collected by Li Zhi-Yin from Yunnan. In 1978,
he described myriapods collected by the same collector from Xisha islands. In 1980, he reported
a preliminary study on the Symphyla of China, based on material collected by CHEN Zhong-Pin
from Jinhua, Zhejiang.

From 1981 to 1983, ZHANG & Li described five new species belonging to five different
groups of millipedes, described the new family Bilingulidae in 1981, and redescribed
Scolopendra mazhii, collected by Li Zhi-Yin from Tibet, in 1983. The next year, ZHANG Chong-
Zhou published a new xystodesmoid, taken by Mao Jerong from Zhejiang province and, in the
same year, Li Zhi-Yin published a summary of centipede species of medicinal use. From 1985 to
1990, ZHANG Chong-Zhou reported three new species taken by Li from southwestern China,
and described a new genus and species of harpagophoroid collected by ZHANG Nai-Gang from
Yunnan in 1990.

In 1988 an important paper by ZHANG & CHEN Zhong-Pin appeared on Pauropoda from
Zhemiang province, listing eight species, four of which were described as new. This paper also
gave the first checklist of Pauropods in China. CHEN Jian-Xiu & MENG Weng-Xin described a
new cambalopsoid in 1991. but have not published since. In 1992. ZHANG & WANG Daqing
reported six subtropical soil species in a resource survey on centipedes for medicinal use from
Wuling mountain. In 1993, WANG Daqinq reported six species of diplopods, three of them as
new, based on material from Fujian province.

By the end of 1993, more than 300 species taxa of myriapods had been described or
reported from China in the papers listed in the references. Obviously, we probably know only
5% of the actual and real number of taxa that occur in China, perhaps even less...

PERSPECTIVES AND SUGGESTIONS

From the summary given above, we may get two kinds of impressions: one is that there
are only a few researchers who are engaged in the study of myriapods in China; the other is that
the research mainly focuses on the description of new species, and lacks systematic and in-depth
studies. One of the reasons for this is that the study of myriapodology in China started late.
Another reason is the large size of China. Finally the study of myriapodology currently belongs
to the range of basic science, so that research funds are difficult to obtain, because the result of
study cannot quickly bring profit, especially in the reality of China today. Perhaps the latter is
the reason why few young specialists are interested in this field. Although this situation results
in a vicious circle, it does not necessarily mean that systematics will disappear. On the contrary,
we believe that systematics will prospere again, with the appearance of cladistics as an example.
And we believe that this could be soon, because there are, after all, many old and young
researchers willingly engaged in the study of myriapodology. However, it is true that we face a
very serious challenge, especially in China.

In order to overcome such a situation in China, perhaps it might be practicable to
systematically study other uses for medicinal myriapods, as food for instance (centipedes can be)

DAQING WANG & JEAN-PAUL MAURIES

or by extracting the pure toxin from their bodies to treat disease in order to obtain money for less
applied studies. If successful!, it would be a beneficial circle.

China is a large region including two zoogeographical zones. The study of Chinese
myriapodology needs a large number of biological researchers to join in, including foreign
myriapodologists. Therefore, it is important to develop all kinds international co-operation and
communication. The study of Chinese myriapod fauna will contribute to the myriapod fauna of
the world.

At present in China, the systematic and detailed study of some taxa such as the orders
Julida. Polydesmida and Scolopendromorpha, should carried out first. The next stage would be
to cover these orders in the two zoogeographical zones. The third will be devoted the study of
zoogeography in the two zones. These are just suggestions. It is earnestly hoped that this paper
will provide a usefull appeal to present and future researchers of Chinese Myriapoda, who will
be able to join us and to build their own greater and finer edifices.

PROVISIONAL CHECKLIST OF MYRIAPOD SPECIES OF CHINA

The following preliminary list is mainly compiled from the papers published in China, the partial from abroad.
Because the literature is so scattered and many are new reports, especially some published in Chinese, authors and dates
of publications for species have been included.

Class DIPLOPODA

Subclass Penicillata

Order Polyxenida

Fam. Polyxenidae

Polyxenus hangzoensis Ishii & Liang, 1990
Eudigraphis taiwaniensis Ishii, 1990
Eudigraphis sinensis Ishii & Liang, 1990
Fam. Lophoproctidae

Lophoiurus okinawai (Nguyen Duy - Jacquemin & Conde, 1982) - Ishii, 1990

Subclass Pentazonia

Order Sphaerotheria

Fam. Sphaeropoeidae

Chinosphaera majorina Zhang & Li, 1982
Chinosphaera maculosa Attems, 1935
Chinosphaera multidenta Wang & Zhang, 1 993
Zephronia (?) profuga Attems, 1936
Zephronia (?) hainana Gressitt, 1941

Order Glomerida

Fam. Glomeridae

Hyleoglomeris sinensis (Brolemann. 1896)

Hyleoglomeris emarginata Golovatch. 1981
Pentazonia incertae sedis

" Glomeris ” bicolor (Wood. 1865)

Subclass Colobognatha

Order Platydesmida

Fam. Andrognathidae

Sinocybe cooki Loomis, 1942
Symphyopleurium hozawai (Chamb. & Wang, 1953)

Order Siphonophorida

Fam. Siphonophoridae

Siphonophora sp. (Wang, unpublished)

Subclass Helminthomorpha

Supraorder Iuliformia

Order Fossil “lulus” peii Chioa & Liu, 1 95 1 : 24

Order Spirobolida

Fam. Spirobolidae

Spirobolus bungii Brandt, 1833

= Spirobolus exquisitus Karsch, 1881

Source : MNHN , Paris

MYRIAPODOLOGY OF CHINA

= Spirobolus joannesi Brolemann, 1896 - Wang, 1955, 1958
Spirobolus walkeri Pocock, 1895
Spirobolus cincinnalis Wang & Zhang, 1993
Spirobolus grahami Keeton, 1960
Spirobolus formosae Keeton, 1960
Spirobolus umbobrochus Keeton. 1960
Trigoniulus niger Takakuwa,1940 - Wang, 1955
Trigoniulus takahasii Takakuwa, 1940
Trigoniulus segmentatus Takakuwa, 1940 - Wang, 1955, 1964
Trigoniulus tertius Takakuwa, 1940 - Wang, 1958
Spirostrophus lanyusis Wang, 1955
Spirobolellus latakuwai Wang, 1961

Order Spirostreptida

Fam. Harpagophoridae

Gonoplectus astutus Attems, 1936
Junceustreptus reirorsus Hoffman, 1980
Junceustreptus browningi Demange, 1961
Junceustreptus prominulus Demange, 1961
Junceustreptus brevispinus Zhang, 1985
Uriunceustreptus afemorispinus Zhang & Chang, 1990
Agariogonopus acrotrifoliatus Zhang (in press)

Order Cambalida

Fam. Pericambalidae

Bilingulus sinicus Zhang & Li. 1981 _

Parabilingulus aramulus Zhang & Li. 1981
Fam. Cambalidae

Glyphiulus anophthalmus (Loksa, 1960)

Glyphiulus balaszi (Loksa, 1960)

Glyphiulus granulatus Gervais, 1847

= ? Glyphiulus vulgatus Zhang & Li. 1982

= ? Glyphiulus tuberculatus (Verhoeff, 1936) - Chamberlin & Wang. 1953,
Wang, 1955, 1957
Glyphiulus formosci (Pocock, 1895)

Glyphiulus pu Icher (Loksa, 1960)

Glyphiulus recticullus Zhang & Li, 1982
Glyphiulus multicarinus Zhang & Li, 1982
Glyphiulus adeloglyphus Zhang & Li. 1982
Glyphiulus quadrohamatus Chen & Meng. 1991

Order Julida

Fam. Nemasomatidae

Orinisobates gracilis (Verhoeff. 1933) - Enghoff, 1985
Sinostemmiulus simplicior Chamberlin &Wang, 1953 - Hoffman, 1966
Fam. Mongoliulidae

Skleroprotopus confucius Attems, 1901
Skleroprotopus laticoxalis Takakuwa, 1942
Skleroprotopus serratus Takakuwa & Takashima. 1949
Skleroprotopus membranipedalis Zhang. 1985
Fam. Paraiulidae

Karteroiulus niger Attems. 1909 - Enghoff, 1987

Fam. Julidae

Amblyiulus sp. Takakuwa & Takashima, 1949
Anaulaciulus paludicola (Pocock. 1895) - Causey, 1966
Anaulaciulus simplex (Verhoeff, 1936) - Wang. 1964
Anaulaciulus tonginus (Karsch, 1881)

Anaulaciulus trapezoidus (Wang, 1955. 58. 63)

Anaulaciulus trilobus (Wang, 1963)

= Anaulaciulus trilobus quemoyensis (Wang, 1963)

Anaulaciulus trilobus khuuae ( Wang, 1963)

Anaulaciulus vallicola (Pocock. 1895) - Causey, 1966
Nepalmatoiulus tibetanus Enghoff, 1987
Nepalmatoiulus rhaphimeritus Enghoff, 1987

DAQING WANG & JEAN-PAUL MAURIES

Nepalmatoiulus brachymeritus Enghoff, 1987
Nepalmatoiulus polyakis Enghoff. 1987
Nepalmatoiulus fraterdraconis Enghoff. 1987
Nepalmatoiulus eulobos Enghoff. 1987
Nepalmatoiulus yunnanensis Enghoff, 1987

Supraorder Coelochaeta

Order Callipodida

Fam. Caspiopetalidae

BoUmania sp. Golovatch, 1981
Fam. Sinocallipodidae

Sinocallipus simplicipodus Zhang. 1993
Fam. Paracortinidac

Paracortina voluta Wang & Zhang. 1993
Paracortina leptoclada Wang & Zhang. 1993
Paracortina (Ahum) carinata (Wang & Zhang. 1993)

Paracortina (Altum) serrata (Wang & Zhang, 1993)

Paracortina ( Relictus ) stimula (Wang & Zhang, 1993)

Paracortina (Relictus) thallina (Wang & Zhang, 1993)

Paracortina (Altum) viriosa (Wang & Zhang, 1993)

Order Craspedosomatida (=Chordeumatida auct.)

Fam. Diplomaragnidae

Syntelopodeuma gracilipes Verhoeff. 1941- Wang, 1958
Diplomaragna formosanum (Verhoeff. 1936) - Shear, 1990
Fam. Speophilosomatidae

Speophilosoma sp. Wang, 1958
incertae sedis

G.sp. Verhoeff, 1933 - Chamberlin & Wang, 1953

Superorder Merocheta
Order Polydesmida

Suborder Paradoxosomaiidea

Fam. Paradoxosomatidae
Subfam. Alogolykinae)

Tribe Alogolykini

Yuennanina ceratogaster Altems, 1 936
Yuennanina aceratogaster Zhang & Li, 1977
Yuennanina petalolobodes Chang & Zhang, 1989
Tribe Polydrepanini

Orophosoma hingstoni (Carl, 1935) - Jeekel, 1980
Orophosoma simulans (Carl. 1935) - Jeekel, 1980
Subfam. Paradoxosomatinae
Tribe Tectoporini

Helicorthomorpha holstii (Pocock, 1895) - Wang, 1955, Jeekel, 1980,
Golovatch, 1981

= Chinosoma hodites Chamberlin. 1923
= Kochliopus trivittatus Verhoeff, 1933
Helicorthomorpha ocellata (Pocock, 1895) - Jeekel, 1980
= Helicorthomorpha uncinata (Attems, 1937)
Helicorthomorpha orthogona (Sil vestri , 1898) - Jeekel, 1980
= Helicorthomorpha kosingai (Wang, 1958)

Tribe Sulciferini

Orthomorpha coarctata Saussure, 1860 - Wang, 1956, 1957
Oxidus gracilis C.L.Koch, 1847 - Pocock, 1895, Wang, 1955,

Wang & Zhang, 1993
Hedinomorpha hummeli Verhoeff, 1933
Hedinomorpha hummeli svenhedini Verhoeff, 1933
Hedinomorpha biramipedicula Zhang & Tang, 1985
Kronopolites swinhoei (Pocock, 1895) - Hoffman, 1963

= Kronopolites svenhedini Verhoeff, 1933 - Zhang & Li, 1978
= Kronopolites formosanus (Verhoeff, 1939)

= Kronopolites ralphi Wang, 1957
Kronopolites acuminatus biagrilectus Hoffman, 1963

MYRJAPODOLOGY OF CHINA

Mandarinopus gracilipes Verhoeff, 1933
Polylobosoma roseipes (Pocock, 1895) - Jeekel, 1980
= Orthomorpha penicillata Attems, 1 93 1
Sichotanus mandschuricus Golovatch, 1978
Sigipinius grahami Hoffman, 1961

"Orthomorpha" (unnamed genus!) nordenskjoeldi Attems, 1909 -
Wang, 1955, 1964

"Orthomorpha" (unnamed genus) corticina Attems, 1 936
Tribe Chamberlinini

Chamberlinius pekuensis (Karsch, 1881) - Wang, 1955 , Golovatch, 1981
= Oxidus corcifera Verhoeff, 1931 - Wang, 1957
= Orthomorpha affinis Verhoeff, 1936 - Takashima. 1939
Chamberlinius haulienensis Wang, 1956 - Hoffman, 1973
Chamberlinius shengmui Wang, 1957 - Hoffman. 1973
Chamberlinius picrofasciatus (Gressitt. 1941) - Hoffman. 1973
Tribe Hylomini

Desmoxytes planata (Pocock, 1895) = D. rastrituberus (Zhang, 1986)
Desmoxytes draco (Cook & Loomis, 1924)

Desmoxytes piceofasciata (Gressitt, 1941)

Desmoxytes longispina (Loksa, 1960)

Desmoxytes cornuta (Zhang & Li, 1982)

Desmoxytes minutubercula (Zhang, 1986)

Tribe Tonkinosomatini

Aponedyopus montanus Verhoeff. 1939 - Takakuwa, 1942 , Wang, 1964
Aponedyopus reesi (Wang, 1957)

Aponedyopus jeanae (Wang, 1957)

Aponedyopus maculatus Takakuwa, 1942
Szechuanella tenebra Hoffman. 1961
Tribe Nedyopodini :

Nedyopus pat riot icus (Attems, 1898) - Wang, 1955, 1964
Varyomorpha hsientienensis Wang, 1957
Varyomorpha pectinata Wang, 1957
Paradoxosomatidae incertae sedis

Orthomorpha bisulcata Pocock, 1895 - Wang, 1957
Orthomorpha flavomarginata Gressitt, 1941
Gonebelus sinensis Attems, 1936
Strongylosoma nadari Brolemann. 1896
Orthomorpha endeusa Attems. 1 898

Orthomorpha circulars Takakuwa in Takakuwa & Takashima, 1949
Suborder Polydcsmidea

Superfam. Polydesmoidca
Fam. Polydesmidae

Polydesmus liber Golovatch, 1991

Pacidesmus sinensis (Golovatch & Hoffman, 1989) - Golovatch. 1991
= Polydesmus hamatus Loksa, 1 960
Epanerchodus potanini Golovatch, 1991
Epanerchodus shirinensis (Chamberlin & Wang, 1953)

Epanerchodus stylotarseus Chen & Zhang, 1 990
Epanerchodus sphaerisetosus Zhang & Chen, 1983
Epanerchodus eurycomutus (Zhang, 1992)

Epanerchodus takakuwai Verhoeff, 1931 - Wang, 1958
Epanerchodus orientalis Attems, 1901 - Wang, 1956, 1964
Fam. Doratodesmidae

Eutrichodesmus arcicollaris Zhang & Wang, 1993
Crenatidorsus grandifoliatus Zhang & Wang, 1993
Pocillidorsus dorsiangulatus Zhang & Wang, 1993
Parapauroplus mono dent us Zhang & Wang, 1993
Fam. Haplodesmidae

Prosopodesmus jacobsoni Silvestri, 1910, Wang, 1964
Fam. Cryptodesmidae

Niponia nodulosa Verhoeff, 1931 - Wang, 1955, 1964

Source : MNHN. Paris

DAQ1NG WANG & JEAN-PAUL MAURIES

Niponia simplexus (Wang, 1957)

Superfam. Stylodesmoidea
Fam. Pyrgodesmidae

Cryptocorypha spinicoronatus Zhang & Li, 1981
Delurodesmus orienfalis Si 1 vestri . 1948
Thelodesmus armatus Miyoshi, 1951 - Wang. 1958
Suborder Chelodesmidca

Superfam. Xystodesmoidea
Fam. Xystodesmidae

Tribe Orophini

Kiulinga jeekeli Hoffman. 1956
Kiulinga lobosa Zhang & Mao, 1984
Pamelaphe lacustris (Pocock, 1895) - Hoffman, 1964
Tribe Harpaphini

Riukiaria taiwanalis (Takakuwa, 1942)

Riukiaria uraensis (Wang. 1956)

Riukiaria holstii (Pocock. 1895) - Wang. 1964
Riukiaria neptuna (Pocock, 1895) - Wang, 1964
Riukiaria variata (Pocock, 1895) - Wang, 1964
Riukiaria capaca Wang & Zhang, 1993
Riukiaria ochraceus (Gressitt, 1941)

Riukiaria taiwanus (Takakuwa. 1942) - Chamberlin & Wang, 1953
Rhysodesmus (?) cohaesivus Wang, 1957
Rhysodesmus (?) contiguus Wang, 1957
Pachydesmus (?) attemsi Wang, 1960
Polydesmida incertae sedis

Polydesmus moorei Pocock. 1895
Polydesmus paludicola Pocock, 1895

Subclass Epimorpha

Order Geophilomorpha

Fam. Himantariidae

Class CHILOPODA

Stigmaiogasier japonica Takakuwa, 1935
Fam. Schendylidae

Subfam. Schendylinae

Escaryus latzeli Sseliwanoff, 1881 - Attems, 1927

Escaryus japonicus Attems. 1927 - Takakuwa & Takashima, 1949, Wang, 1957
Escaryus sachalinus Takakuwa. 1935 - Takakuwa & Takashima, 1949
Subfam. Ballophilinae

Ballophilus liber Chamberlin. 1952
Thalthybius boiehoboensis Wang, 1955
Fam. Oryidae

Orphnaeus brevilabiatus Newport, 1845 - Pocock, 1895 , Wang, 1955
Fam. Geophilidae

Subfam. Geophilinae

Geophilus infossulatus Attems, 1901
Pleurogeophilus takakuwai Verhoeff, 1934
Subfam. Dignathodontinae

Paraplanes svenhedini Verhoeff, 1933

Scolioplanes transsilvanicum (Verhoeff, 1928) - Wang, 1959
Scolioplanes maritimus japonicus (Verhoeff, 1935) - Wang, 1959
Subfam. Pachymerinae

Pachymerium ferrugineum C.L. Koch, 1847 - Takakuwa, 1938, Wang, 1956,
Takakuwa & Takashima, 1949

Pachymerium atticum Verhoeff, 1901 - Takakuwa & Takashima, 1949
Fam. Mecistocephalidae

Subfam. Mecistocephalinae

Formosocephalus longichilatus Takakuwa, 1937
M ec is tocephalus rubriceps Wood. 1862 - Wang, 1956, 1959
Mecistocephalus mikado Attems, 1928 - Takakuwa, 1938, Wang, 1956

Source : MNHN, Paris

MYRIAPODOLOGY OF CHINA

Me c istocephal us brevisternalis Takakuwa, 1934

Mecistocephalus fenestratus Verhocff, 1934

M ecist ocepha l us lakakuwai Verhoeff, 1934

Mecistocephalus ongi Takakuwa, 1934

Mecistocephalus multidentatus Takakuwa, 1936

Mecistocephalus japonicus Meinert, 1870 - Wang. 1963

Mecistocephalus nannocornis Chamberlin. 1920 - Wang, 1957

Mecistocephalus diversisternus Silvestri. 1919 - Wang. 1957

Mecistocephalus punctifrons Newport. 1845 - Wang, 1963

Mecistocephalus smithi Pocock, 1895 - Chamberlin & Wang. 1952 - Wang. 1955

Mecistocephalus mirandus Pocock, 1 895

Mecistocephalus insularis (Lucas, 1863) - Attems, 1929. Wang. 1956, 1959
Mecistocephalus insulomontanus Gressitt, 1941
Mecistocephalus monticolens Chamberlin, 1920 - Wang. 1956
Nodocephalus dooi Takakuwa, 1940 - Wang, 1959
Nodocephalus edentulus Attems, 1910 - Wang, 1956. 1963
Nodocephalus pauroporus Takakuwa, 1936
Taiwanella striata Takakuwa in Takakuwa & Takashima, 1949
Taiwanella sculptulatus Takakuwa, 1936
Taiwanella yanagiharai Takakuwa, 1936
Tygarrup javanicus Attems, 1907 - Chamberlin & Wang, 1952
Sublam. Arrupinae

Prolamnonyx holstii (Pocock, 1895) - Takakuwa & Takashima, 1949
= Mecistocephalus indecorus Attems,- 1901
Prolamnonyx sauteri Silvestri. 1919
Order Scolopendromorpha

Fam. Scolopendridae

Subfam. Scolopendrinae

Scolopendra calcarata Porat, 1876
Scolopendra cingulata Latreille, 1829 - Haase, 1887
Scolopendra mazbii Gravely, 1912 - Zhang & Li, 1983
Scolopendra morsitans L. - Pocock, 1895, Wang, 1955, 1956
Scolopendra mutilans L. Koch 1878 - Pocock. 1895, Brblemann, 1896,
Takakuwa. 1938. Wang, 1955

Scolopendra multidens Newport, 1845 - Haase, 1887, Wang, 1955, 1956
Scolopendra rapax Gervais. 1 847
Scolopendra rugosa Meinert. 1886

Scolopendra subspinipes Leach, 1817 - Pocock, 1895 , Wang. 1955 & auct...

= Scolopendra septemspinosa Brandt. 1841- Newport, 1845
Scolopendra subspinipes dehaani Brandt, 1840 - Pocock, 1895,

Wang. 1955 , 1956

Scolopendra subspinipes japonica L. Koch 1878 - Pocock, 1895, Wang, 1955
Trachycormocephalus koreanus Verhoff, 1934 - Takakuwa, 1938
Subfam. Otostigminae

Otostigmus aculeatus Haase 1887 - Pocock. 1895, Wang, 1955. 1956
Otostigmus insularis (Haase, 1887) - Wang, 1959
Otostigmus malayanus (Chamberlin, 1922) - Wang, 1959
Otostigmus scaber ( =carinatus ) Porat, 1 876

Pocock, 1895, Brolemann,1896, Chamberlin & Wang, 1952,

Wang, 1955. 1956

Otostigmus politus Karsch, 1881 - Attems, 1901

Otostigmus politus mandschurius Verhoeff, 1942

Otostigmus politus pigmentatus Attems, 1930 - Wang, 1955

Otostigmus striatus Takakuwa, 1 940

Otostigmus striatus porteri Dobroruka, 1960

Otostigmus multispinosus Takakuwa, 1937

Otostigmus astenus (Kohlrausch, 1881) - Wang, 1955

Otostigmus frigidus Verhoeff. 1942

Otostigmus frigidus lakakuwai Verhoeff, 1942

Rhysida mandchurica Miyoshi, 1939

Rhysida nuda nuda Newport. 1845 - Wang, 1959

DAQ1NG WANG & JEAN-PAUL MAURIES

Rhysida tiuda brevicomuia Wang, 1951 - Wang, 1957
Rhysida nuda immarginata (Porat, 1876) - Wang, 1955, 1957
Rhysida longipes (Newport, 1845) - Wang. 1956

Rhysida longipes brevicornis Takakuwa, 1934 - Takakuwa.1938 et Wang, 1957
Rhysida yanagiharai Takakuwa. 1935
Rhysida lilhobioides (Newport, 1845)

Fam. Cryptopsidae

Subfam. Cryptopsinae

Cryptops nigropictus Takakuwa, 1936 - Takakuwa, 1938, Wang, 1956
Cryptops japonicus Takakuwa, 1934 - Chamberlin & Wang. 1952,

Takakuwa, 1938

Mimops orientalis Kraepelin, 1903
Subfam. Scolopocryptopsinae

Scolopocryptops brolemanni Kraepelin, 1903

Otocryptops rubiginosa L.Koch, 1878 = O. confucii Karsch. 1884

Otocryptops sexspinosus Say. 1821 - Pocock, 1895 , Attems, 1930

Subclass Epimorpha

Order Lithobiomorpha

Fam. Lithobiidae

Subfam. Lithobiinae

Arebius chengsiensis Chamberlin & Wang, 1952
Areebius bidens Takakuwa, 1941 - Wang, 1952
Chinobius chekianus Chamberlin & Wang, 1952

Chinobius chekianus lumeopes Chamberlin & Wang, 1952 - Wang, 1955, 1956..
Chinobius svenhedini (Verhoeff, 1933)

Chinobius sac hal inns Verhoeff. 1937 - Wang, 1956. 1959
Chinobius (?) pachypedatus Takakuwa, 1938 - Wang, 1954
Lilhobius hummeli (Verhoeff, 1933)

Lithobius bidivisa Takakuwa, 1939 - Wang, 1963

Lilhobius kiayiensis Wang. 1959

Lithobius ongi Takakuwa, 1941 - Wang, 1959

Lithobius trichopus Takakuwa. 1939 - Wang, 1955, 1959

Lithobius tetrophthalmus Loksa, 1960

Lithobius aeruginosus mongolicus Attems, 1901

Lithobius decessus Attems, 1901

Lithobius jangsleanus Verhoeff, 1942 - Loksa, 1965

Lithobius kansuanus Verhoeff, 1933

Lithobius mongolicus Verhoeff, 1933

Lithobius erratus Attems, 1938

= Chinobius (?) sulcipes Attems, 1934 - Wang. 1959
Lithobius bogdoulensis Loksa, 1965
Lithobius anornatus Loksa, 1965
Lithobius mongolellus Loksa, 1965
Lithobius mongolomedius Loksa, 1965

Lithobius sulcifemoralis Takakuwa in Takakuwa & Takashima, 1949
Lithobius gantoensis Takakuwa in Takakuwa & Takashima, 1949
Lithobius irregularis Takakuwa in Takakuwa & Takashima, 1949
Lilhobius rufus Muralevitch. 1929 - Loksa, 1965
Monotarsobius crassipes L. Koch, 1862 - Wang, 1963
Monotarsobius crassipes holstii (Pocock, 1895) - Wang, 1959
Monotarsobius rhysus (Attems, 1934) - Chamberlin & Wang, 1952
Monotarsobius argaeensis (Attems, 1905) - Chamberlin & Wang, 1952
Monotarsobius obtusus Takakuwa, 1941 - Wang, 1955, 1956
Monotarsobius ramulosus Takakuwa, 1941 - Wang, 1955, 1956
Monotarsobius alticus Loksa, 1965
Monotarsobius crassus Loksa, 1965
Monotarsobius kaszabi Loksa, 1965
Subfam. Ethopetolidae (=Polybothridae)

Bothropolys asperatus L.Koch, 1878 - Chamberlin & Wang, 1952,

Takakuwa, 1938, Wang, 1956
= Lithobius lethidis Karsch, 1880

M YRI APODOLOGY OF CHINA

= Lithobius asperatus L. Koch, 1878 - Pocock, 1895
= Lithobius rugosus Meinert, 1872 - Attems, 1901
Bothropolys crassidentatus Takakuwa in Takakuwa & Takashima, 1949
Bothropolys imaharensis (Verhoeff, 1937)

Takakuwa, 1938, Chamberlin & Wang, 1952, Wang, 1959
Bothropolys richthofeni Verhoeff, 1938 - Takakuwa & Takashima, 1949
Bothropolys shansiensis Takakuwa in Takakuwa & Takashima, 1949
Fam. Henicopidae

Alaskobius takakuwai Chamberlin & Wang, 1952
Esastigmatobius longicornis (Takakuwa, 1936) - Wang, 1959
Esastigmatobius longitarsis Verhoeff, 1934 - Wang, 1959
Hedinobius hummeli Verhoeff. 1933
Lamyctes gracilipes Takakuwa, 1941 - Wang, 1957
Fam. Pterygotergidae

Pterygotergum svenhedini Verhoeff, 1933

Order Scutigeromorpha

Fam. Scutigeridae

Scutigera coleoptrata L. - Wang, 1959
Scutigera sinuata Haase, 1887
Scutigera complanata Haase, 1887
Scutigera hispida Haase, 1887 - Attems, 1901
Thereuopoda clunifera Wood, 1 862

= Scutigera longicornis clunifera (Wood, 1862) - Pocock, 1895,
Chamberlin & Wang, 1952 - Wang, 1955, 1956
= Scutigera sinensis Meinert, 1 886
Thereuopoda nivicomes Verhoeff, 1942
Thereuonema tuberculata Wood, 1862 - Pocock, 1895
Thereuonema variata Miyoshi. 1939

Thereuonema mandschuria Verhoeff, 1936 - Chamberlin & Wang, 1952 ,
Takakuwa, 1938

Thereuonema dilatationis Verhoeff, 1936 - Takakuwa & Takashima, 1949
Thereuonema hilgendorfi Verhoeff. 1905 - Chamberlin & Wang, 1952,
Takakuwa, 1938

Thereuonema viridescens Verhoeff, 1937 - Chamberlin & Wang, 1952
Class PAUROPODA

Ectomorphes

Fam. Pauropodidae

Allopauropus ovalapendicis Zhang & Chen, 1988
Allopauropus pilosisphaerus Zhang & Chen. 1988
Pauropus bifurcus Zhang & Chen, 1988
Pauropus longirarnus Zhang & Chen, 1988
Fam. Polypauropidae

Fagepauropus hesperius Remy, 1951 - Chalupsky, 1972

Endomorphes

Fam. Eurypauropodidae

Subfam. Eurypauropodinae

Eurypauropus sp. Zhang & Chen, 1988
Subfam. Sphaeropauropinae

Sphaeropauropus sp. Zhang & Chen, 1988

Class SYMPHYLA
Fam. Geophilellidae

Geophilella pyrenaica Ribaul, 1913 - Takashima, 1939
Fam. Scutigerellidae

Scutigerella immaculata (Newport, 1845) - Wang. 1957

Source : MNHN, Paris

DAQING WANG & JEAN-PAUL MAURJES

COLLECTING LOCALITIES

Figure 1 shows approximatively the locations of sites in China where myriapods have
been collected, during the modern period. In some cases, one single symbol represents several
nearby localities. It is obvious that most of the sampling efforts have been concentrared in the
south of China. The map unfortunately does not show the intensity of collecting; how many
species are known from a given locality? This information cannot be provided until all of the
groups of the specimens in the Institute of Zoology of Academia Sinica have been worked out in
detail. The collection was mainly taken by Li Zhi-Yin and WANG Daqing from 1969 to 1992 in
China.

Fig. I. — Localities in China at which myriapods have been collected. Some closely adjacent sites are represented by
single symbols. Due to lack of available material, the geographic coordonates are not given for the following
localities. Undoubtedly, some omissions have occured because a few places could not be located exactly,
particularly when the localities are small villages: Diao Luo Mountain. Hainan Island - Fuzhou, (vicinity of the
city), Fujan province - Yiang shou, a region of the city Gui Lin, Guangxi province - Tian cun, Guangxi province -
Mengman, Yunnan province - Luxi, Yunnan province - Jinhua, Zhejiang province - Hong Kang - Jilong, Taiwan -
Zhangjakou, Hebei province - Beijing, (vicinity of the city) - Tanmo Mountain, Zhejiang province - Daishan,
Zhejiang province - Lanzhou (vicinity), Gansu province - Tai Bai Mountain, Shannxi province - Kunming,
Yunnan province - Guanlin (in the caves), Gueizhou province - Hangzhou. Zhejiang province - Chayu, Tibet -
Shenyang (vicinity), Liaoning province - Muotuo. Tibet - Chang Bai Mountain, Jilin province - Ningbo,
Zhejiang province - Putuo. Zhejiang province - Changsha (vicinity), Hunan province - Yulushan, a hill of
Changsha vicinity - Mengla, Yunnan province - Hekou. Yunnan province - Xichou, Yunnan province - Mengzi.
Yunnan province - Zhongdian, Yunnan province - Deqin, Tibet - Yiajiang, Sichuan province - Batang, Sichuan
province - Taigu, Shanxi province - Zhousha Islands, Zhejiang province - Jiangle, Ml. longxi, Fujian province -
Taibei, Taiwan - Taizhong, Taiwan - Pingdong, Taiwan - Zhangjiajie, National Forest Park, Hunan province-
Sangzhi, Hunan province - Huhehot, Inner Mongolia - Tianjin( vicinity ), Hebei province.

Source :

MYRIAPODOLOGY OF CHINA

REFERENCES

The aim of the following list of references is to compile all the papers that have been published concerning
Chinese myriapodology, including those in wich were described taxa previously found out of China (*) and some ones of
general interest (**). Any omissions which may be discovered will be added in future works.

Attems, C, 1898. — System der Polydesmiden I. Denks. Math.-naturwiss. Kais. Akad. Wiss., Wien : 221-482.

Attems, C., 1901. — Myriopoden. In : Dritte Asiatische Forschungreise des Grafen Eugen Zichy, Bd. 2, Horvath : 275-
310.

*Attems, C, 1901. — Neue Polydesmiden des Hamburger-Museums. Milt. Naturhist. Mus., 18 : 85-105

* Attems, C, 1904. — Central-und hoch-asiatische Myriopoden gesammelt im Jahre 1900 von Dr von Almassy und Dr.

von Stummer. Zool. Jahrb., Syst., 20: 113-130

* Attems, C., 1905. — Myriopoden in Ergebnisse einer naturwissenschaftlichen Reise zum Erschias-Dagh (Kleinasien).

Ann. k.k. Naiurh. Hof museums, 20 : 1-5

* Attems, C., 1907. — Japanische Myriopoden gesammelt von Direktor Dr K. Kraepelin im Jahre 1903. Mitt. Naiurh.

Mus., 24: 77-142.

* Attems, C., 1909. — Die Myriopoden der Vega-Expedition. Arkiv f Zool., 5 : 1 -84.

ATTEMS, C., 1927. — 2. Neue Chilopoden. Zool. Anz .. 72 : 291-305.

* Attems, C., 1928. — Eine neue Gattung und eine neue Art der Mecistocephalidae (Chilopoden). Zool. Anz.. 75 : 1 15-

120.

A ITEMS, C., 1929. — Myriapoda 1. Geophilomorpha. In : F. E. SCHULZE & W. KOKENTHAL, Das Tierreich. 52. Berlin &
Leipzig, W. De Gruyter & C° : 1-388.

Attems, C., 1930. — Myriapoda 2. Scolopendromorpha In : F. E. Schulze & W. KOkenthal, Das Tierreich , 54. Berlin
& Leipzig, W. De Gruyter & C° : 1-308.

Attems, C, 1931. — Die Familie Leptodesmidae und andere Polydesmiden. Zoologica, 30 : 1-150.

ATTEMS, C., 1934. — Einige neue Geophiliden und Lithobiiden des Hamburger Museums. Zool. Anz., 107 : 310-317.
ATTEMS, C., 1935. — Myriopoden von Sumatra. Arch. Hydrobiol., suppl. 14: 114-142.

Attems. C.. 1936. — Diplopoda of India. Mem. Indian Mus., 11,4: 133-323.

Attems, C., 1937. — Myriapoda 3. Polydesmoidea. I. Fam. Strongylosomidae. In : F. E. Schulze, W. KOkenthal & K.

Heider. Das Tierreich, 68, Berlin & Leipzig, W. de Gruyter & C° : 1-300.

ATTEMS, C., 1938. — Die von Dr. G. Dawydoff in franzosisch Indochina gesammelten Myriapoden. Mem. Mus. natn.
Hist, nat., 6 : 187-353.

ATTEMS, C., 1953. — Myriapoden von Indochina Expedition von C. Dawydoff (1938-1939). Mem. Mus. natn. Hist,
nat.. A5 : 133-230.

*Brandt. J. F., 1840. — Observations sur les cspeces qui composent le genre Scolopendra suivies de caracteres des
espfcces qui se trouvent dans le Museum Zoologique de 1'Academie des Sciences de Saint-Petersbourg et quelques coups
d’ceil sur leur distribution geographique. Bull. Sci. Acad. Saint-Petersbourg, 1.

Brolemann, H. W., 1896. — Sur quelques Myriapodes de Chine. Mem. Soc. zool. France. : 349-362.

Causey, N. B., 1966. — Redcscription of two Chinese species of Anaulaciulus (Diplopoda. Julida, Nemasomatidae) a
genus known also in Taiwan, Korea and Japan. Proc. Louis. Acad. Sci., 29 : 63-66.

CHALUPSKY, J., 1972. — A new find of the genus Fagepauropus (Pauropoda). Vestnik Cesk. Spolecn. Zool., 2 : 89-92.
^Chamberlin. R. V., 1914. — Notes on Chilopods from the East Indies. Entom. News Philadelphia, 25 : 385-392.
*Chamberlin, R. V., 1920. — On the Chilopods of the family Mecistocephalidae. Canad. Entom., 52 : 184-189
Chamberlin. R. V., 1923. — Two Diplopod immigrant taken at Honolulu. Proc. Biol. Soc. Washington, 36 : 165-168.
Chamberlin, R. V., 1952. — Geophiloid Chilopods of the Hawaian and other Oceanic islands of the Pacific. Gr. Basin
Nat., 13 : 75-85

Chamberlin, R. V. & Wang, Y. M., 1952. — Some records and descriptions of Chilopods from Japan and other oriental
areas. Proc. Biol. Soc. Wash., 65: 177-186.

Chamberlin, R. V. & Wang, Y. M., 1953. — Records of Millipeds from Japan and other oriental areas, with description
of new genera and species. Amer. Mus. Novitates, 1621 : 1-13.

Chang, N. G. & Zhang, C. Z., 1989. — A contribution to knowledge of the genus Yunnanina and a new species in
Yunnan province (Diplopoda Paradoxosomatidae). Ada Zootaxonomica Sinica, 14 : 415-419.

Chen. J. X. & Zhang, C. Z., 1990. — A cave-dwelling new species of the Diplopod genus Epanerchodus from Guizhou
province. Acta Zootaxonomica Sinica, 15 : 407-409.

DAQING WANG & JEAN-PAUL MAURIES

Chen, J. X. & Meng, W. X., 1991. — A new species of the genus Glyphiulus (Diplopoda, Spirostreptida,
Cambalopsidae). Acta Zootcixonomica Sinica, 16 : 394-395.

Chia , L. P. & LlU, H. T., 1951. — Fossil Myriapods from Choukoutien. Bull Geoi Soc. China , 30 : 23-27.

COOK, O. F. & LOOMIS, H. F., 1924. — A new family of spined Millipeds from Central China. J. Wash. Acad. Sci., 14 :
103-108.

Demange, J. M., 1962. — Mat6riaux pour servir & une revision des Harpagophoridae (Myriapodes-Diplopodes). Mem.
Mus. natn. Hist, nat., ser. A, Zool. , 24, (1961) : 1-274.

Dobroruka, L. J., 1960. — A new formosan Centipede: Otostigmus striatus porteri n.sp. Ann. Mag. Nat. Hist., 13 :
61-62.

Dobroruka, L. J., 1960. — Ueber einc kleine Chilopoden-Ausbeute aus der Mongolei. Acta Arachnologica , 17 : 15-
18.

Enghoff, H., 1985. — The millipede family Nemasomatidae. With the description of a new genus and a revision of
Orinisobates (Diplopoda: Julida). Ent. Scand., 16 : 27-67 .

Enghoff. H.. 1987. — Karteroiulus niger Attems, 1909, in China (Diplopoda, Julida, Paraiulidae). Entom. Monthly
Mag..\23\ 207-208.

Enghoff, H., 1987. — Revision of Nepalmatoiulus Mauries 1983, a southeast Asiatic genus of Millipedes (Diplopoda:

Julida: Julidae). Courier Forsch.-Inst. Senckenberg, 93 : 241-331.

Golovatch, S. I., 1978. — Some east-asiatic Millipedes (Diplopoda) in collection of the Zoological Institut of the
Academy of Sciences of USSR. Rev. Entom. , USSR, 3 : 677-681.

Golovatch, S. I., 1981. — Some East-asiatic millipedes (Diplopoda) in the collection of the Institute of Zoology of the
Polish Academy of Sciences. Ann. Zool. 36 : 161-168.

Golovatch, S. I., 1991. — The millipede family Polydesmidae in southern Asia, with notes on phylogeny (Diplopoda:
Polydesmidea). Steenstrupia, 17 : 141-159

Golovatch, S. I. & Hoffman R. L., 1989. — Identity of Polydesmus hamatus Brandt, 1841, a Malagasy milliped
(Diplopoda, Polydesmida. Dalodesmidae). Tropical Zool.. 2 : 159-164.

Gervais, P., In WalC'KENAER, 1847. — Histoire Naturelle des insectes apteres T.4. Myriapodes. Nouvelles suites a
Buffon , Paris, Librairie Encyclopedique : 1-333.

Gravely, F. H., 1912. — Two new species of Scolopendridae. Rec. Indian Mus., 7 : 415-417.

Gressitt, J. L., 1941. — New Myriopods from Formosa and Hainan Islands. Ann. Mag. Nat. Hist., 11 : 55-61.

Haase, E., 1887. — Die indo-australische Myriopoden. I. Chilopoden. Abh. Ber. K. Zool. Anthrop. Mus. Dresden,
1886-1887 : 1-118.

Hoffman. R. L., 1956. — Studies on some oriental xystodesmine millipeds. Proc. Ent. Soc. Wash., 58 : 95-104.
Hoffman. R. L., 1961. — Two new Diplopod genera from western China (Polydesmida, Strongylosomidae). Ann. Mag.
Nat. Hist., I960, ser. 13, 3 : 533-543.

Hoffman, R. L., 1963. — A contribution to the knowledge of asiatic Strongylosomoid Diplopoda (Polydesmida,
Strongylosomidae). Ann. Mag. Nat. Hist., 1962, ser. 13,5 : 577-593.

Hoffman, R. L., 1964. — A new subfamily of Xystodesmid Millipeds from North America and China (Polydesmida).
Trans. Amer. Ent. Soc., 90 : 301-311.

Hoffman, R. L., 1966. — The ordinal position of the generic name Sinostemmiulus Chamberlin & Wang, 1953. Proc.
Ent. Soc. Wash., 68 : 322-325.

Hoffman, R. L., 1973. — Descriptions and allocation of new or poorly known genera and species of Paradoxosomatidae
from south-eastern Asia (Diplopoda: Polydesmida). J. Nat. Hist., 1 : 361-389.

**Hoffman, R. L., 1980. — Classification of the Diplopoda. Genfcve, Museum d'Histoire Naturelle (1979) : 238 pp.
Hoffman, R. L., 1980. — Studies on Spirostreptoid millipeds. XV. On some new or poorly known Harpagophorid
genera and species from China and the East Indies. J. Nat. Hist., 14 : 589-596.

Ishii, K., 1990. — Penicillate Diplopods (Diplopoda: Polyxenida) from Taiwan. Edaphologia, 42 : 1-20.

Ishii, K. & Liang L., 1990. — Two new species of Penicillate Diplopods of the family Polyxenidae from China. Cand.
Entom., 122: 1239-1246.

JEEKEL C. A. W., 1968. — On the classification and geographic distribution of the family Paradoxosomatidae
( Diplopoda , Polydesmida). Amsterdam: 1-162.

**Jeekel C. A. W., 1971. — Nomenclator generum et familiarum Diplopodorum. Monogr. Nederl. Entom. Ver.
Amsterdam, 5 : I-XII, 1-412.

Jeekel C. A. W., 1980. — The generic allocation of some little-known Paradoxosomatidae from South-East Asia
(Diplopoda, Polydesmida). Rev. Suisse Zool.. 87 : 651-670.

Karsch, F., 1880. — Ein neuer Lithobius. Zeitschr. f Ges. Naturwiss., 53 : 848.

Source : MNHN, Paris

MYRIAPODOLOGY OF CHINA

Karsch, F., 1881. — Zur Studium der Myriapoda Polydesmida I. Diagnosen neuer Polydesmiden des Berliner Museum.
Archiv f Nat urges., 47 : 36-43.

Karsch, F., 1881. — Neue Juliden des Berliner Museums, als Prodromus eincr Juliden-Monographie. Zeitschr. f Ges.
Naturwiss ., 54 : 1-79.

Karsch, F., 1884. — Classis Myriapoda. In : Ueber einigc neue und minder bekannte Arthropoden des Bremer
Museums. Abh.Ver. Bremen, 9 : 65-67.

Keeton, W. T., 1960. — A taxonomic study of the milliped family Spirobolidae (Diplopoda, Spirobolida) Mem Amer
Entom. Soc., 17 : 1-146.

*Koch, C. L., 1847. — System der Myriapoden III. Regensburg.

*K0CH. L., 1862. — Die Myriapodengatlung Lithobius. Niirnberg : 1-92.

Koch, L., 1878. — Japanesische Arachniden und Myriapoden. Verb. d. k. k. Zool. Bot. Ges. Wien, 1877 27 • 735-
798.

*Kohlrausch, E., 1881. — Gattungen und Arten der Scolopendriden. Arch. F. Naturges., 47 : 50-132.

Kraepelin, K., 1903. — Revision der Scolopendriden. Mitt. Naturh. Mus. Hamburg, 20 : 1-276.

*Leach, W. E., 1817. — A tabular view of the external characters of four classes of animals, which Linne arranged under
insecta; with the distribution of the genera composing three of the Classes into Orders, and description of several
new genera and species. Trans. Linn. Soc. London, 11, (1814) : 306-400.

Li, Z. Y. & Zhang, C. Z., 1980. — The individual development ol Scolopendra subspinipes mutilans (Scolopendrida,
Chilopoda). Chinese J. Zool., 15 : 17-19.

Li, Z. Y. & Zhang, C. Z., 1982. — Experimental studies on the increase in the breeding rate of Scolopendra subspinipes
mutilans by use of the artificial nest. Chinese J. Zool., 17 : 27-30.

Li, Z. Y., 1984. — The species of medicinal centipeds and the selection oftheir breeding. Chinese J. Zool., 19 : 12-15.
Lin, P. S. & DOUY, L. H., 1988. — The toxicological experiment of toxin taken from Scolopendra mutilans. Chinese J.
Zool. , 23 : 20-21.

Loksa, I., 1960. — Einige neue Diplopoden und Chilopoden- Arten aus Chinesischen Hohlen. Acta Zool. Hungar., 6 :
135-148.

Loksa, I., 1965. — Zoologische Ergebnisse der Forschungen von Dr Z. Kaszab in der Mongolei. 21. Chilopoda. Opusc.
Zool., 5: 199-215.

Loksa, I.. 1978. — Chilopoden aus der Mongolei (Arthropoda: Tracheata, Chilopoda). Ann. Hist. Nat. Mus. Nat. Hung.,
70 : 111-120.

LOOMIS, H. F., 1942. — Sinocybe, a new genus of Colobognath millipeds from China. J. Wash. Acad. Sci., 32 : 270-
273.

*Lucas, H., In Maillard, 1863. — Myriapodes. In : Note sur l'lle de La Reunion.

Matic, Z., 1973. — Revision du genre Chinobius Verhoeff, avec description de trois especes nouvelles (Chilopoda,
Lithobiida). Ann. Zool. Pologne, 30 : 33-47.

*Meinert, F., 1872. — Myriapoda Musaei Hauniensis. Naturh. Tidsskr. : 281-344.

MEINERT, F., 1886. — Myriapoda Musaei Hauniensis. Vid. Medd. f. d. naturh. Foren. i Kjobenhavn, (1884) : 100-156.

Meinert, F., 1886. — Myriapoda Musaei Cantabrigensis. Part I. Chilopoda. Proc. Amer. Phil. Soc.. (1885). 23 : 161-
233.

MlYOSHI. Y., 1939. — Chilopoda. In : Report of the first Scientific Expedition to Manchoukuo under the leadership of
Shigeyasu Tokunaga, June-Oct. 1933, 5, I, 14, n°77 : 1-26.

* Miyoshi, Y., 1951. — Beitrage zur Kenntnis japanischer Myriopoden. 1. Aufsatz: Ueber eine Gattung von
Leptodesmidae. Zool. Mag. Tokyo, 60 : 149-150.

*Muralevitch, V. S., 1929. — Scutigeridae et Lithobiidae de la faune du Caucase. Mem. Soc. Amis Sci. Nat. Moscou .
Zool. : 1-120.

Newport, G., 1845. — XXVI11. Monograph of the class Myriapoda. order Chilopoda; with observations on the general
arrangement of the Articulata. Trans. Linn. Soc. London, (1844), 19 : 349-438.

*NGUYEN Duy - Jacquemin. M. & Cond£, B., 1982. — Lophoproctides insulaires de l'Ocean Pacifique (Diplopodes:

Penicillates). Bull. Mus. nail. Hist. nat. . Paris. 4 : 95-118.

POCOCK, R. L., 1895. — Report upon the Chilopoda and Diplopoda obtained by P. W. Bassett-Smith and J. J. Walker
during the cruise in the Chinese Seas of H. M. S. Penguin. Ann. Mag. Nat. Hist., 6 : 346-372.

Pocock. R. L., 1895. — The Myriapoda of Burma. Pt. IV. Report upon the Polydesmoidea collected by S. L. Fea, M. E.
W. Oates and others. Viaggio di Leonardo Fea in Birmania e regione vicine. Ann. Mus. civ. Stor. nat. Genova, ser.
2a, 14, 34 : 787-834.

PORAT, C. O., 1876. — Om nagra exotiska Myriopoder. Sven. Vet. Akad. Handlingar, 4 : 1-48.

DAQ1NG WANG & JEAN-PAUL MAURIES

*Remy, P.. 1951. — Un nouveau type de Pauropode: Fagepauropus hesperius n. g., n. sp. du Sud-marocain. Bull. Mus.
natl. Hist. not.. Paris, 2 , 23 : 208-210.

*Ribaut, H., 1913. — Un nouveau genre de la Classe des Symphyles (Myriopodes). Bull. Soc. Hist. nat. Toulouse , 14 :
77-84.

*S.AUSSURE, H. de, 1860. — Essai d'une Faune des Myriapodes du Mexique. Memoires pour servir d THistoire naturelle du
Mexique , des Antilles el des Etats-Unis. Geneve : 1-135.

*Say, E., 1821 . — Description of the Myriapoda of the United States. J. Acad. Nat. Sci. Philad., 1° sen, 2 : 102-1 14.
*SELIVANOV, A., 1881. — Geophilidae Museja imperatorskoi. Mem. Acad. Imp. Sci. Saint-Peterbourg, 1.

Selivanov, A.. 1881. — Neue Lithobiiden aus Sibirien und Central Asien. Zool. Anz . . 4 : 15-17.

Shear, W. A., 1990. — On the Central and East Asian Milliped family Diplomaragnidae (Diplopoda, Chordeumida,
Diplomaragnoidea). Amer. Mus. Novitates. 2977 : 1-40.

*SlLVESTRl. F., 1898. — Alcuni nuovi Diplopodi della N. Guinea. Annali Mus. civ. Stor. nat. G. Doria, 39 : 441-449.
*SlLVESTRl, F.. 1910. — Descrizioni preliminari di nuovi generi di Diplopodi I. Polydesmoidea. Zool. Anz.. 35 : 357-
364.

Silvestri, F., 1919. — Contributions to the knowledge of the Chilopoda Geophilomorpha of India. Records Indian
Mus., 16 : 45-107.

Silvestri, F.. 1948. — Contribute alia conoscenza dei Diplopodi termitofili dell'America meridionale. Boll. Lab.
Entom. agraria Portia . 8 : 1-32.

*Takakuwa, Y., 1934a. — Sechs neue Arten der Chilopoden aus Japan. Trans. Sapporo Nat. Hist. Soc., 13 : 398-406.
Takakuwa, Y., 1934b. — Ueber drei neue Arten der Chilopoden aus Formosa und Marschallinseln. Trans. Nat. Hist.
Soc. Formosa, 24 , n°132 : 221-225.

*Takakuwa, Y„ 1935a. — Ueber neue Chilopoden aus Japan. Trans. Nat. Hist. Soc. Formosa. 25, n° 145 : 339-342.
*Takakuwa, Y., 1935b. — [Japanese Escaryus (Chilopoda)]. Dobuts. Zasshi, 47. n° 561: 407-41 1.

Takakuwa, Y., 1936a. — Eine neue interessante Mecistocephalus- Art aus Formosa. Trans. Nat. Hist. Soc. Formosa,
26 . n° 152 : 215-216.

Takakuwa, Y.. 1936b. — Ueber eine neue Gattung und zwei neue Arten von Mecistocephalidae aus Formosa. Trans. Nat.
Hist. Soc. Formosa. 26 , n° 159 : 435-439

Takakuwa, Y., 1936c. — Ueber eine neue Subspecies der Ethmostigmus aus Marshallinscln und eine neue spezies der
Nodocephalus aus Mandschurei. Trans. Nat. Hist. Soc. Formosa, 26, n° 151 : 105-106.

*Takakuwa, Y., 1937 . — Otostigmus-Anen aus Japan. Trans. Nat. Hist. Soc. Formosa, 21, n° 164 : 221-225.
Takakuwa. Y., 1938. — Verzeichnis der Japanischen und Mandschurcischcn Chilopodenarten. Annot. Zool. jap., 17 :
353-359.

*Takakuwa, Y., 1939. — Uber Japanische Lithobius- Arten. Trans. Sapporo Nat. Hist. Soc., 16 : 28-37.

*TAKAKUWA, Y., 1940a. — Eine neue Nodocephalus Arten (Chilopoda) aus Korea. Trans. Nat. Hist. Soc. Formosa, 30,
n° 197-198 : 41-42.

Takakuwa, Y., 1940b . — Eine neue Art von Otostigmus (Chilopoda) aus Formosa. Trans. Nat. Hist. Soc. Formosa, 30,
n° 200-201 : 209-210.

Takakuwa, Y., 1940c . — Ueber zwei neue Trigoniulus- Arten (Diplopoda) aus Taiwan. Trans. Nat. Hist. Soc. Formosa,
30, n° 200-201 : 211-215.

Takakuwa, Y., 1940d. — Weitere Trigoniulus- Arten aus Japan (Diplopoda). Ann. Zool. jap., 19 : 283-288.
Takakuwa, Y.. 1941. — Ueber einige japanische Lithobiiden. Trans. Nat. Hist. Soc. Formosa, 31, n° 213 : 292-297.
Takakuwa, Y., 1942a. — Ueber weitere japanische Rhysodesmus- Arten. Trans. Nat. Hist. Soc. Formosa. 32, n° 224:
197-203.

Takakuwa, Y., 1942b. — Die Myriapoden von Formosa, Philippinien, u.s.w. Trans. Nat. Hist. Soc. Formosa, 32, n°
231 : 359-367.

Takakuwa, Y., 1942c . — 3 n. sp. of Diplopoda from Japan. Zool. Mag. Tokyo, 54 : 237-239. (in japanese).
*Takakuwa, Y., 1942d. — Zur Kenntnis der japanischen Diplopoden. Annot. Zool. Jap., 21 : 39-47.

Takakuwa, Y., 1943. — Die drei neue Diplopoden aus Taiwan und West-nippon. Trans. Nat. Hist. Soc. Formosa, 33,
n° 242-243 : 603-606.

Takakuwa, Y. & Takashima, H., 1949. — Myriapods collected in Shansi, North China. Acta Arachnologica, 9,
n° 3 14 : 51-69.

Takashima, H., 1939. — Diplopoda. In : Report of the first Scientific Expedition to Manchoukuo under the leadership
of Shigeyasu Tokunaga, June-Oct. 1933, 5 , 1 , 14, n° 78 : 1-11.

Source MNHN, Pari$

MYRIAPODOLOGY OF CHINA

TaKASHIMA, H., 1939 — Symphyla. In : Report of the first Scientific Expedition to Manchoukuo under the leadership
of Shigeyasu Tokunaga, June-Oct. 1933, 5 . I . 14. n° 79 : 1-5.

* VERHOEFF, K. W., 1901. — Zur vergleichendc Morphologic, Systcmatik und Geographic der Chilopoden. Nova Acta

Leopoldina, 77 : 369-465.

* Verhoeff. K. W., 1905. — Ueber Scutigeridcn. 6. Aufsatz. Variabil itat und Thereuonema- Arten. Tarsen mit

sprungweiser Abanderung. Zool. Anz 29: 353-371.

*Verhoeff, K. W.. 1914. — Ascospermophoren aus Japan. Zool. Anz. ,43 : 342-370.

*Verhoeff, K. W., 1928. — Geophilomorphen-Beitrage und eine Lithobius-Fom. Mitt. Zool. Mus. Berlin , 14 : 227-
286.

* V erhoeff, K. W., 1931. — Chiiognathen aus den Bergamasker Alpen und Nachbargebieten; aus iiber zwei neue
Gattungen der Polydesmoidea aus Spanien und Japan. Zool. Jahrb., Syst., 61 : 397-452.

Verhoeff, K. W.. 1933. — Schwedisch-chinesische wissenschaftlich Expedition nach den nordwesilichen Provinzen
Chinas, unter Leitung von Dr. Sven Hedin und Prof. Su Ping-chang. Ark. Zool., 26 . 1-41.

* VERHOEFF, K. W., 1934. — Beitrage zur Systcmatik und Geographic der Chilopoden. Zool. Jarbh., Syst.. 66 : 1-152.
Verhoeff, K. W., 1935. — Ueber Scolioplanes. Zool. Anz Ill: 10-23.

Verhoeff, K. W.. 1936a. — Zur Kenntnis der Glyphiuliden (Cambaloidea). Zool. Anz., 113 : 49-62.

VERHOEFF, K. W., 1936b. — Kritische Untersuchung asiatischer Scutigeriden. Zool. Anz.. 115 : 1-18.

Verhoeff, K. W., 1936c. — Ueber Diplopoden aus Japan gesammelt von Herrn Y. Takakuwa. Trans. Sapporo Nat. Hist.
Soc., 14: 148-172.

* Verhoeff, K. W., 1937a. — Chilopoden aus Malacca nach den Objekten des Raffles Museum in Singapore. Bull.

Raffles Mus. Singapore. 13 : 198-270.

* Verhoeff, K. W., 1937b. — Zur Kenntnis der Lithobiiden. Arch. NaturgPsch., 6 : 71-257.

* Verhoeff, K. W.. 1938. — Uber einigc ostasiatische Lithobiiden. Zool. Anz., 123 : 99-105.

Verhoeff, K. W., 1939a. — Zur Kenntnis ostasiatischer Diplopoden. IV. Zool. Anz., 127 : 273-285.

Verhoeff, K. W., 1939b. — Diplopoden von den Ryukiu-Insel Okinawa. Biogeographica, 3 : 118-122. '

Verhoeff, K. W., 1942a. — Chilopoden aus innerasiatischen Hochgebirgen. Zool. Anz.. 137 : 35-52.

Verhoeff. K. W., 1942b. — Otostigmen der Mandschurei. Zool. Anz., 138 : 181-188.

Wang, D. Q.. 1993. — Animals of Jangle Mountain: Diplopoda, Chilopoda. Ed. Huang Chuenmei, China Forestry
Publishing House : 855-862.

Wang, D. Q. & Zhang, C. Z.. 1993. — Diplopoda. Chilopoda. Animals of Longqi Mountains, 8 : 845-851.

Wang, D. Q. & Zhang. C. Z., 1993. — A new Family of Millipeds (Diplopoda, Callipodida) from southwestern China.
Mem. Peking Nat. Hist. Mus., 53 : 375-390.

*WANG, Y. M., 1951. — The Myriapoda of the Philippine Islands. Ed. Edwards (Ann Arbor, Michigan, USA) : 1-80.
Wang. Y. M., 1955a. — Serica la: Records of Myriapods on Formosa with descriptions of new species. Quart. J. Taiwan
Mus., 8 : 13-16.

Wang, Y. M., 1955b. — Serica lb: A preliminary report on Myriapoda and Arachnida of Lan You Islets (Botel Tobago),
China. Quart. J. Taiwan Mus., 8 : 195-201.

Wang, Y. M., 1956. — Serica le: Records of Myriapods on Formosa with description of new species (2). Quart. J.
Taiwan Mus., 9: 155-159.

Wang. Y. M.. 1957a. — Serica If : Records of Myriapods on Taiwan Islands (3) Pescadore Islets, Kao-Young, Pingtung.
Changhua and Taipei. Quart. J. Taiwan Mus.. 10 : 23-29.

Wang, Y. M.. 1957b. — Serica lg : Records of Myriapods on Taiwan Islands (4) Six new Polydesmids. Quart. J. Taiwan
Mus., 10 : 103-1 1 1.

Wang, Y. M.. 1957c. — Serica lh : Records of Myriapods on Taiwan Islands (5) With descriptions of three new species.
Quart. J. Taiwan Mus., 10 : 112-115.

Wang, Y. M.. 1958a. — Serica li : On Diplopoda from Taiwan with a new Strongylosomid. Quart. J. Taiwan Mus., 11 :
340-344.

Wang, Y. M., 1958b. — Records of Formosan Myriapods. In : Proc. 1 0th hit. Congr. Ent. Montreal, 1. 1956 : 881-
882.

Wang, Y. M., 1959. — Serica Ij : On Chilopoda from Taiwan with a new Lithobid. Quart. J. Taiwan Mus.. 12 : 195-
199.

Wang, Y. M.. 1960. — On Millipedes and Centipedes from Taiwan. China. In : Verb. XL intern. Kongr. Entom. Wien,
17-25 August I960 : 288-291.

DAQING WANG & JEAN-PAUL MAURIES

Wang, Y. M., 1961. — Serica Ik: Millipedes of Taiwan. A new species of family Spirobolidae. Quart. J. Taiwan Mus .,
14: 141-142.

Wang. Y. M., 1963a. — Serica Iq: Millipedes and Centipedes of Quemoy. Fukien province and Taiwan Island, Bold
Tobago (Lan Yu), Taiwan province and of Singapore. Quart . J. Taiwan Mus.. 16 : 89-96.

Wang, Y. M., 1963b. — The Millipedes and Centipedes of Taiwan, China. In : Proc. XVl° Congress Zooi Washington.
J : 211.

Wang, Y. M., 1964. — Serica lop: Wallacea and insular Fauna of Millipedes. Quart. J. Taiwan Mus.. 17 : 67-76.

Wang, Y. M. & Wang T. N. G.. 1965. — Distribution of orders Polydcsmida and Spirobolida in some oriental islands.
In : Proc. XHth hit. Congress Entom. London. 1964 : : 444-445.

Wood. H. C. 1863. — On the Chilopoda of North America, with a catalogue of all the specimens in the collection of the
Smithsonian Institution. J. Philad. Acad.. 5: 5-52.

Wood. H. C.. 1865. — New Polyzoniidae. Proc. Acad. Philadelphie, 17 : 172-173.

Zhang, C. Z., 1985a. — A new species of millipeds of the genus Junceustreptus in Southwestern China. Acta Zool.
Sinica, 10: 137-139.

Zhang, C. Z., 1985b. — A new species of millipeds of the genus Skleroproropus in Stone Buddha Cave. Fangshang
County. Beijing. In : Karst Geomorphology and Speleology. Beijing, Science Press : 154-156.

Zhang. C. Z.. 1986. — On the genus Pratinus and its two new species from China. Ada Zootaxon. Sinica , II : 253-
257.

Zhang. C. Z., 1987a. — A study on decocting methods for Scolopendra spp. Bull. Chin. Mater Med., (1986), 11 : 66 1 -
663.

Zhang, C. Z., 1987b. — Breeding biology of medicinal Centipeds. Sci. Press.

Zhang, C. Z.. 1993. — Diplopoda from Yunnan caves (II): Contribution to the study of a new cavernous taxon of the
Nematophoran Millipeds (Diplopoda. Ccelocheta, Callipodida). In : Proc. Xlth hit. Congress Speleoi, August 2-8
1993. : 128-130.

ZHANG, C. Z.. (in press). — An interesting new millipede. Agariogonopus acrotrifoliatus, gen. et sp. nov., from a
limestone district of Yunnan, China.

Zhang, C. Z. & Chang. N. G.. 1990. — A new genus and new species of the family Harpagophoridae from Yunnan,
China (Diplopoda, Spiroslreptoidea). Acta Zootaxonom. Sinica. 15 : 32-35.

Zhang, C. Z. & Chen. Z. P.. 1983. — A new species of the genus Epanerchodus (Diplopoda: Polydesmida) from Zhejiang
province. Acta Zhejiang Teacher's College (Nature Sci.), 6 : 87-89.

Zhang, C. Z. & Chen, Z. P., 1988. — A preliminary study on a new record of the class Pauropoda (Myriapoda) from
China. Sinozoologica. 6 : 39-54.

Zhang, C. Z. & Jin, Y. L., 1981. — A textual study on the medicinal Myriapoda in Ben Cao Gang Mu (the Chinese
Encyclopedia of material medicine) 1596. J. Chin, tradition. Medicine, 478 : 50-56.

Zhang, C. Z. & Li, Z. Y.. 1975. — On the difference between Spirobolus joannesi (Brolemann) and Kronopolites
swinboei (Pocock) used in medicine. Chinese J. Zooi, 4 : 29-30.

Zhang, C. Z. & Li. Z. Y., 1977a. — Eine neue Yunnanina- Art (Diplopoda) aus China. Acta Zooi Sinica, 23 : 357-359.

Zhang, C. Z. & Ll. Z. Y., 1977b. — A survey of the Centipede used in medicine ( Scolopendra subspinipes muhidens
Newport) from Duan County, Guangxi, Chinese J. Zooi. 1977, 2 : 33-34.

Zhang, C. Z. & Li. Z. Y., 1977c. — Preliminary studies on the centipede ( Scolopendra subspinipes muhidens Newport)
from Hubei province. Chinese J. Zooi, 1977, 1 : 34-36.

Zhang, C. Z. & Li, Z. Y., 1978a. — On medical Kronopolites svenhedini (Verhoeff) (Diplopoda, Paradoxosomatidae).
Chinese J. Zooi, 1978, 3 : 12-13.

Zhang, C. Z. & Li, Z. Y.. 1978b. — On some Myriapoda from Xisha Islands, Guangdong province. Chinese J. Zooi,
1978, 4: 1-3.

Zhang. C. Z. & Li, Z. Y., 1979a. — Studies on the habits and characteristics of Scolopendra subspinipes mutilans L.
Koch (Chilopoda). Chinese J. Zooi, 1979, 1 : 20-22.

Zhang, C. Z. & Ll, Z. Y., 1979b. — The spawn (observations on egg-laying) of Scolopendra mutilans L. Koch. Chinese
J. Zooi, 1979, 4 : 13-15.

Zhang. C. Z. & Ll, Z. Y., 1979c. — The habits of Scolopendra mutilans L. Koch. Chinese J. Zooi, 1979, 4 : 20-22.

Zhang, C. Z. & Ll, Z. Y., 1980a. — A preliminary analysis of structure and product of Scolopendra mutilans in Daishan
County. Chinese J. Zooi. 1980, 2 : 1-3.

Zhang, C. Z. & Ll, Z. Y., 1980b. — Textual research of medicinal Spirobolus. J. Chin, tradition. Medicine, 6 : 66-67.

Zhang, C. Z. & Li. Z. Y.. 1981a. — Ueber Bilingulidae fam. nov. (Diplopoda Spirostreptida) aus dem sudchinesischen
Karstgcbiet. Acta Zooi Sinica. 6 : 376-377.

Source : MNHN. Paris

MYRI APODOLOGY OF CHINA

Zhang, C. Z. & Li, Z. Y., 1 98 1 o. — fcine neuc Art vom Archandrodestnus (Cryptodesmidac, Diplopoda). Acta
Zootaxonom. Sinica, 6 ; 250-252

Zhang, C. Z. & Li, Z. Y., !982a. — Centrodesmus commas sp. nov. eine neue Diplopoden-Art (Paradoxosomatidae,
Polydesmida) aus dem Sud-China. Acta Zootaxonom. Sinica. 7 : 37-39.

Zhang, C. Z. & Li, Z. Y., 1982b. — Eine neue Art vom Chinosphaera (Spherotheriida, Diplopoda) aus China. Acta
Zootaxonom. Sinica ,7: 152-154.

Zhang, C. Z. & Li, Z. Y., 1982c. — Die Gattung Glyphiulus (Diplopoda, Cambalidea) von China. Sinozoologica , 2 :
85-93.

Zhang, C. Z. & Li. Z. Y., 1983. — The Scolopendra mazbii Gravely. 1912 of Xizang Autonomus region (Tibet). Chinese
J. Zool.. 1983, 5 : 50-51.

Zhang, C. Z. & Li. Z. Y., 1990. — Textual research of medicinal Polydesmoid. J. Chin, tradition. Medicine. 466 : 66-
67.

Zhang, C. Z., Li, Z. Y.& Mao, J. R.. 1980. — The preliminary study on Symphyla of China. Chinese J. Zool.. 1980,
2: 4-7.

Zhang, C. Z., Ll, Z. Y. el al. 1983. — The development of Scolopendra subspinipes mutilans L. Koch (Scolopendridae,
Chilopoda). Chinese J. Zool.. 1983, 4 : 17-19.

Zhang, C. Z., Li, Z. Y. & Mao, J. R., 1984. — A new species of the genus Kiulinga (Diplopoda. Xystodesmidae). Acta
Zootaxonomica Sinica, 9 : 135-137.

Zhang, C. Z. & Tang, H.G., 1985. — Eine neue Art der Gattung Hedinomorpha (Diplopoda. Paradoxosomatidae) aus
China. Sinozoologica, 3 : 35-38.

Zhang, C. Z. & Wang, D. Q.. 1993. — Diplopoda from Yunnan caves (I): A study on new genera and species of the
millipede family Doratodesmidae. In : Karst landscape & Cave tourisni. China Environ. Sci. Press : 205-220.

Zhang, C. Z., Zhang, F. X. & Wang. D. Q., 1992. — A survey on the resources of Centipeds for medicinal use in Wuling
Mountains. Chinese J. Zool., 21 : 8-11.

Source :

Source : MNHN. Paris

A Taxonomic Study of Polydesmoid Millipedes
(Diplopoda) Based on their Mandibular Structures

Kiyoshi ISH1I * and Hiroshi TAM UR A **

* Department of Biology. Dokkyo University School of Medicine. Mibu. Tochigi 321-02. Japan
** Department of Biology. Ibaraki University. Mito. Ibaraki 310. Japan

ABSTRACT

The mandibular structures of 8 families. 25 genera and 34 species of polydesmoid millipedes have been examined based
on the material collected from Japan and China. Detailed examinations revealed that the molar structure was especially
useful for the taxonomy of this order. The key to the family on the basis of molar structure is also given.

RESUME

Etude taxinomique des diplopodes polydesmoides basee sur leurs structures mandibulaires.

Les structures mandibulaires de 8 families. 25 genres et 34 especes de diplopodes polydesmides ont ete examinees a
partir du materiel r£colle au Japon et en Chine. Des observations detaillees ont montre que les structures “molaires”
etaient particulierement utiles pour 1'etude taxinomique de 1'ordre Polydesmida. Une cle des families. bas£e sur la structure
en "molaire”, est egalement proposee.

INTRODUCTION

The mandibular structures of millipedes have so far been treated as minor taxonomic
characters since MANTON (1964), while LAUTF.RBACH (1972) noted function of mandible in
millipedes. ENGHOFF (1979, 1981, 1985) and ISH1I (1988) discussed on the taxonomic
significance of mandibular structures in a few millipede orders. Recently, ISHII & TAMURA
(1992) compared mandibles of 9 orders, 22 families, 46 genera and 65 species of millipedes,
suggesting usefulness of the mandibles as a diagnostic character in general in millipedes.

The mandibular structures of polydesmoid millipedes were investigated by ENGHOFF
1979, 1985) for only three families and four genera: Polydesmidae, Brachydesmus and
Serradium ; Macrosternodesmidae, Macrosternodesmus and Oxydesmidae gen.

In this study, we have made an intensive examination on the mandibular structure of 8
families, 25 genera and 34 species (TABLE 1) collected from Japan and China to confirm its
extensive usefulness as a taxonomic character throughout Polydesmoid millipedes.

METHODS

Mandibles were removed from the mouthparts using an ophthalmologic knife and forceps under a stereoscopic
binocular microscope, and further made clear using a microbrush and fluid pressure. The mandibles were fixed with 80%

Ishii. K. & Tamura, H.. 1996. — A taxonomic study of polydesmoid millipedes (Diplopoda) based on their
mandibular structures. In: Geoffroy. J.-J.. Mauries, J.-P. & Nguyen Duy - Jacquemin, M.. (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 101-111. Paris ISBN : 2-85653-502-X.

102

KIYOSHI ISHII & HIROSHI TAMURA

alcohol, dehydrated in a graded alcohol series, put overnight into isoamyl, and then dried in a critical point drier using
liquid carbon dioxide. The dried mandibles were coated with gold and observed with a scanning electron microscope.

Table 1. — List of Polydesmoid millipedes species examined.

Fam. Paradoxosomatidae: Haplogonosoma implication, Chamberlinius cristatus, C. haulienensis, Oxidus
gracilis. Orihomorpha coarctata, Nedyopus tambanus, Cemrodesmus sp. (unpublished sp. collected from
China), undetermined gen. collected from China.

Fam. Xystodesmidae: Levizonus takakuwai. Parafontaria ishiii, P. laminaia armigera, Xystodesmus sp.
(unpublished sp.), Riukiaria sp. (unpublished sp.).

Fam. Polydesmidae: Polydesmus japonicus, Epanerchodus mammillatus, E. orientalis, E. sp.-l
(unpublished sp.), E. sp.-2 (unpublished sp.), Prionomatis sp. (unpublished sp.).

Fam. Opisotretidae: Corypholophus sp. (unpublished sp.).

Fam. Cryptodesmidae: Kiusiunum sekii . K. nodulosum. Niponia nodulosa.

Fam. Pyrgodesmidae: Ampelodesmus granulosus. Cryptocorypha japonica.

Fam. Doratodesmidae: Eucondylodesmus elegans. Kylindogaster nodulosa, Thelodesmus armatus, T . sp.
(unpublished sp.), Dimorphodesmus sp.-l (unpublished sp.), D. sp.-2 (unpublished sp.).D. sp.-3 (unpublished
sp.), undetermined gen. collected from China.

Fam. Haplodesmidae. Rhipidopeltis sp. (unpublished sp.).

GENERAL MORPHOLOGY OF THE MANDIBLE OF POLYDESMOID MILLIPEDES
General morphology of the mandible of millipedes has been mentioned by ATTEMS
(1937), MANTON (1964), ENGHOFF (1979), ISHII (1988), ISHII & TAMURA (1992), and
HOPKIN & Read (1992). The general morphology of mandibles of Polydesmoid millipedes are
shown in Figure 1. The mandible is composed of 4 parts: 1) the proximal plate, armed with
molar plate consisting of wide or narrow molar processes, molar comb between molar process,
molar hook, molar tuft at proximal end, fringe, and granulated zone; 2) the intermediate plate,
armed internally with intermediate lobe covered by scaly hairs; 3) comb-lobe, with six rows of
comb teeth; 4) internal tooth and external tooth. Both molar plates has a muller as showed in
Figures 1-4.

RESULTS AND DISCUSSION

The mandible of Polydesmoid millipedes is characterized by extremely stout external and
internal teeth and comb teeth arranged in six rows, and also by crescent or triangular molar plate
with wide or sawtooth roof-like molar process, molar tuft at proximal end and lamellate fringe or
branched fringe at apex.

The external tooth of mandible is simple in the families Paradoxosomatidae and
Xystodesmidae (Fig. 2). Other families such as Polydesmidae, Opisotretidae, Cryptodesmidae,
Doratodesmidae, Pyrgodesmidae and Haplodesmidae have one or two lateral denticles. The
families Opisotretidae, Haplodesmidae and Pyrgodesmidae have a node.

The internal tooth is marginally divided into four to seven leaflets. The number of leaflets
and depth of emargination are varied depending on genus and/or species. Therefore, we assume
that the structures of both external and internal teeth are important diagnostic characters for lower
laxa rather than the family.

The intermediate lobe is less developed except the family Pyrgodesmidae in comparison
with the other millipedes orders such as Penicillata, Spirostreptida, Chordeumatida and Julida.

The molar structure of mandible is most stable within diagnostic characters of the family of
polydesmoid millipedes. The molar structures are given below for the 8 families examined in this
study.

Source :

POLYDESMOID MILLIPEDES MANDIBULAR STRUCTURES

103

internal tooth

intermediate lobe

comb teeth

tooth

molar process

molar plate

fringe

proximal plate

Fig. I. — The mandibular structure of Parafonlaria laminala armigera. 1. right mandible (dorsal view); 2, molar plate
(internal view); 3, disposition of mandible, muller, pharynx and oesophagus (dorsal view); 4, muller.

Source : MNHN, Paris

104

KlYOSHI ISHII & HIROSHI TAMURA

Nedyopus Xysiodesmus Doratcxlcsmidae gen.

(Fam. Paradoxosoinalidae) (Fam. Xystodesmidae)

Polydesmus
(Fam. Polydesmidae)

Corypholophus
(Fam. Opisotretidae)

Rhipidopellis
(Fam. Haplodesmidae)

Niponia Cryptocorypha

(Fam. Cryptodesmidae) (Fam. Pyrgodesmidae)

Ampelodesmus
(Fam. Pyrgodesmidae)

Fig. 2. — The externa! and internal teeth of the eight families in polydesmoid millipedes.

Fam. Paradoxosomatidae : Molar plate having wide sawtooth roof-like or lean-to roof-like
molar process, lamellate fringe with sharp tine at apex without inner branches and thick molar
tufts at proximal end (Fig. 3).

Fam. Xystodesmidae : Molar plate having shape sawtooth roof-like molar processes,
lamellate fringe with slightly round tine at apex without inner branches and tiny molar tufts at
proximal end (Fig. 4).

Source : MNHN. Paris

POLYDESMOID MILLIPEDES MANDIBULAR STRUCTURES

105

Fit;. 3. — Molar plate of the family Paradoxosomatidae. 1. Haplogonosoma implication ; 2, Oxidus gracilis', 3, Nedyopus
lambanus ; 4. Orthomorpha coarctata .

Source : MNHN, Paris

106

KIYOSHI [SHU & HIROSHI TAMURA

Fig. 4. — Molar plate of the family Xystodesmidae. 1. Levizonus takakuwai: 2, Parafontaria laminata armigera ; 3,
Xysiodesmus sp.; 4. Riukiaria sp.

Fam. Polydesmidae : Molar plate having wide molar process, separated fringes with inner
branches and thick, long molar tufts at the proximal end (Fig. 5).

Source : MNHN, Paris

POLY DESMOID MILLIPEDES MANDIBULAR STRUCTURES

107

Fig. 5. — Molar plate of the family Polydesmidae. 1. Polydesmus japonicus ; 2. Epanerchodus mammillatus; 3,
Prionomatis sp.; 4, Epanerchodus sp.-L

Source : MNHN. Paris

108

KIYOSHI ISH1I & HIROSHI TAMURA

Fig. 6. — Molar plate of the families Cryptodesmidae and Pyrgodcsmidae. Cryptodesmidae: I. Kiusiunum nodulosum ; 2,
Niponia nodulosa. Pyrgodcsmidae: 3, Ampelodesmus granulosus ; 4, Cryptocorypha japonica.

Source : MNHN, Paris

POLY DESMOID MILLIPEDES MANDIBULAR STRUCTURES

109

Pig. 7. — Molar plate of the family Doratodesmidae. L Eucondylodesmus elegans : 2. Kylindogasler nodulosa ; 3.
Thelodesnuis sp.; 4. Dimorphodesnius sp.-l.

Source : MNHN, Paris

110

KIYOSHI ISHII & HIROSHI TAMURA

Fig. 8. — Molar plate of the families Opisotretidae and Haplodesmidac. Opisotretidae: 1, Corypholophus sp.
Haplodesmidae: 2. Rliipidopeltis sp.

Fam. Opisotretidae : Molar plate having remarkably wide molar process, basal molar
process with a stout denticle on ventral border, brushed fringes at apex with inner branches and
long, numerous molar tufts (Fig. 8: 1 ).

Fam. Cryptodesmidae : Molar plate having wide molar process, wing process without
granulated zone, branched fringes at apex with inner branches and slender molar tufts (Fig. 6: 1,
2).

Fam. Pyrgodesmidae : Molar plate having a molar hook, wide molar process, wing
process with granulated zone, branched fringes at apex and slender molar tufts (Fig. 6: 3, 4).

Fam. Doratodesmidae : Molar plate internally having molar combs between narrow molar
processes, branched fringes at apex with few inner branches and dense, molar tufts on a limited
portion at proximal end (Fig. 7).

Fam. Haplodesmidae : Molar plate abruptly narrower sub-basally with dense molar tufts,
branched fringes at apex with inner branches, and molar combs scarce or absent between molar
processes (Fig. 8: 2).

KEY TO THE FAMILY OF THE POLYDESMOID MILLIPEDES BASED ON THE MOLAR

Molar plate with lamellate fringe — .

Molar plate with separate fringe . —

Molar plate with thick, long molar tufts-
Molar plate with tiny, short molar tufts-
Molar plate with molar wing process —
Molar plate without molar wing process

- 2

. — 3

Paradoxosomat idae
Xystodesmidae

- 4

- 5

Source :

POLYDESMOID MILLIPEDES MANDIBULAR STRUCTURES

111

4. Molar plate with a molar hook- . . . . . — Pyrgodesmidae

Molar plate without molar hook - - — Cryptodesmidae

5. Molar combs present between molar processes - - - - 6

Molar combs absent between molar processes - - - 7

6. Molar combs abundant- - - Doratodesmidae

Molar combs scarce or absent - - Haplodesmidae

7. Fringe narrow at apex - - - Polydesmidae

Fringe brush-like at apex - - - - - Opisotretidae

The following eleven genera well resemble morphologically each other: Kiusiunum,
Niponia, Cryptocorypha, Ampelodesmus, Pseudocatapyrgodesmus, Eucondylodesmus ,
Kylindogaster, Dimorphodesmus, Thelodesmus, Rhipidopeltis and undetermined genus
collected from China. These ten genera except the genus Pseudocatapyrgodesmus not
investigated in this study were classified into four groups from intensive examinations on molar
structure, and this grouping agreed with HOFFMAN (1980) as follows: Cryptodesmidae,
Kiusiunum and Niponia ; Pyrgodesmidae, Cryptocorypha and Ampelodesmus: Doratodesmidae,
Eucondylodesmus, Kylindogaster, Dimorphodesmus, Thelodesmus and undetermined genus
collected from China: Haplodesmidae. Rhipidopeltis. On the other hand, the genera Prionomatis
and Epanerchodus of the family Polydesmidae is distinctly the same in molar structure (FIG. 5).
Therefore the genus Prionomatis should be better considered as a synonym of the genus
Epanerchodus.

As far as the present study is concerned, mandibular structures are fairly stable within
taxa and distinctly difterent between taxa, clearly suggesting usefulness as an important
diagnostic character in the taxonomy of the Polydesmoid millipedes.

REFERENCES

ATTEMS. C., 1937. — Myriapoda 3. Polydesmoidea. I. Fam. Strongylosomidae. In : F. E. SCHULZE, W. Kukenthal & K.
HE1DER, Das Tierreich, 68. Berlin & Leipzig. W. de Gruyier & C° : 1-300.

Enghoff, H., 1979. — Taxonomic significance of the mandibles in ihe millipede Order Julida. In ; M. Camatini.
Myriapod Biology, London, Academic Press : 27-38.

Enghoff, H.. 1981. — A cladislic analysis and classification of the millipede order Julida. Z. zool. Syst. Evol.-Forsch..
19 : 285-319.

Enghoff, H.. 1985. — Modified mouthparts in hydrophilous cave millipedes (Diplopoda). Bijdragen tot de Dierkunde ,
55 : 67-77.

Hoffman, R. L., 1980. — Classification of the Diplopoda. Geneve, Museum d’Histoire Naturelle, (1979), 238 pp.
Hopkin, S. P. & READ, H. J.. 1992. — The biology of millipedes. Oxford. Oxford University Press, 233pp.

I SHI l, K.. 1988. — On the significance of the mandible as a diagnostic character in the taxonomy of penicillate
diplopods (Diplopoda: Polyxenidae). Can. Entomol.. 120 : 955-963.

Ishii. K. & Tamura. H., 1992. — The mandibular structure as a diagnostic character in taxonomy of diplopods. Acta
Zool. Fenn ., 196 : 232-235.

Lauterbach, K. E., 1972. — Uber die sogenannte Ganzbein-Mandibel der Tracheaten, insbesondere der Myriapoda.
Zool. Anz . 188 : 145-154.

M ANTON, S. M., 1964. — Mandibular mechanisms and the evolution of arthropods. Philosophical Transactions of the
Royal Society • of London, Series B, 247 : 1-183.

Source : MNHN, Paris

Systematique et biogeographie des diplopodes
penicillates des lies Canaries et du Cap Vert

Monique NGUYEN DUY - JACQUEMIN

Museum national d'Histoire naturelle, Laboratoire de Zoologie/Arthropodes
61, rue Buffon, F-75231 Paris, France

RESUME

Quatrc especes de penicillates ont ete recoltees sur les ties Canaries enrois aux Ties du Cap Vert. Trois de ces sept
especes sent nouvelles et font I'objet d'une description detaillee. Deux sont des Canaries : Fuertoventura. La Palma.
Tenerife pour Polyxenus oromii n. sp. et Fuertoventura pour Mcicroxenus enghoffi n. sp. La troisieme, Anopsxenus
cahoverdus n. sp., est de Santiago, Tune des Ties du Cap Vert. Une forme incertaine de San Antao (Cap Vert) est nommee
cf. enghoffi. Les trois autres especes ont une repartition continentale ou insulaire plus ou moins etendue.

ABSTRACT

Systematic and biogeographical study of Diplopoda, Penieillata of Canary Islands and Cape
Verde Islands.

Four species of Penieillata were collected from the Canary Islands and three from the Cape Verde Islands. These include
three new species which are described in detail. Two of the new species are from the Canary Islands: Polyxenus oromii n.
sp., from Fuertoventura and Tenerife: and Mcicroxenus enghoffi n. sp., from Fuertoventura: while the third. Anopsxenus
caboverdus n. sp.. is from Santiago, Cape Verde Islands. A form of uncertain status, referred to as M. cf. enghoffi, is
recorded from San Antao (Cape Verde). The other three species collected have fairly widespread, continental or insular,
distributions.

INTRODUCTION

Quatre especes de penicillates seulement ont ete citees des lies de la Macaronesie :
Polyxenus lagurus (L.) aux Agores (BROLEMANN, 1896 ; CONDE, 1961), P. fasciculatus Say,
1921 a Madere (CONDE & NGUYEN DUY - JACQUEMIN, 1994), Lophoturus madecassus
(Marquet et Conde, 1950) et Anopsxenus indicus Conde et Jacquemin, 1963 aux lies du Cap
Vert (ENGHOFF. 1993). L'examen des collections qui m'ont ete confiees par les Dr. ENGHOFF,
OROMI et VICENTE me permet d'exposer ici les premieres donnees sur le peuplement des lies
Canaries par les penicillates et de completer celui des lies du Cap Vert.

Nguyen Duy - Jacquemin, M., 1996. — Systematique et biogeographie des diplopodes penicillates des lies
Canaries et du Cap Vert. hr. Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist. nai.. 169 : 113-126. Paris ISBN : 2-85653-502-X.

114

MONIQUE NGUYEN DUY - JACQUEMIN

FAMILLE DES POLYXENIDAE

Polyxenus fasciculatus Say, 1921

Si A I IONS. — Gran Canaria. W of Artenara, 1200 m, slopes with Adenocarpus etc. under
stones, n° 2707, 07.1.1990, H. ENGHOFF leg. : 1 male a 13 pp. (ad), 2 femelles a 13 pp. (ad.).

— Cruz de Tejeda. 1500 m (27 km SW Las Palmas, loc. Luftlinie), wahrscheinlich
Ziegenweide, sehr steinig, unter Steinen, Trocken bis wenig feucht, n° 4122, 24.IV. 1976, W.
HUTHER leg. : 1 male a 13 pp. (ad.), 1 femelle a 13 pp (ad.), 2 femelles a 12 pp., 1 male a 12
pp.- Vega de Acusa. 15.X1I.1987. R. RODRIGUEZ leg. : 1 male a 13 pp. (ad.). — Valle de
Agaete, 30.X.1989. R. RODRIGUEZ leg. : 1 femelle a 13 pp (ad.).

Tenerife. Montana de la Hoya, 6. XI. 1989. R. RODRIGUEZ leg. : 1 femelle a 13 pp. (ad.).

— Los Carboneros, 3. II. 90, P. OROMI leg. : 2 males a 13 pp. (ad.), 1 ind. a 10 pp. — Vueltas
Taganana, 19.1.1991. P. OROMI leg. : 2 males a 13 pp. (ad.), 3 femelles a 13 pp (ad.). — Agua
Garcia et Las Mercedes, IV.94. M. BAEZ leg. : 1 femelle a 13 pp. (ad.) et 1 femelle a 12 pp.

— Sieste Canadas, N.-E. du pare national de Las Canadas, dans la litiere de Spartocytisus
supranubius et Adenocarpus viscosus , 10.VI.95, P. OROMI leg. : 8 femelles a 13 pp. (ad.), 2
males a 13 pp. (ad.), 2 males a 12 pp., 3 males a 10 pp., 1 femelle a 10 pp., 2 ind. a 8 pp.

Gomera. Barranco Na Sa Guadaloupe, 550m, 23.XII.78, V. MONSERRAT leg. : 1 femelle
a 1 3 pp. (ad.).

Hierro. Montana de las Cuevas, 30.III. 1989, R. RODRIGUEZ leg. : 1 femelle a 10 pp.
El Fayal c. 4 km SSW Mirador de Jinama, 1350 m, dense Fayal-Brezal, u. bark of log,
2. II. 1989, A. et H. ENGHOFF leg. n° 242 : 2 males a 13 pp. (ad.), 1 male a 12 pp., 1 femelle a
12 pp., 1 ind. a 8 pp.

P. fasciculatus est tres repandu dans le Centre et le Sud-Est des Etats-Unis, ou il remplace
la forme umsexuee de P. lagurus (CONDE & NGUYEN DUY - JACQUEMIN, 1994), aux Bermudes
(CONDE, 1972) et a Madere (CONDE, 1961, sous le nom de P. lagurus, forme bisexuee). II est
ties voisin de la forme bisexuee de Polyxenus lagurus, tres commune en Europe, notamment sur
le pourtour de la Mediterranee, et presente aux A?ores ; il ne s’en distingue que par un nombre
plus grand de sensilles basiconiques sur le 6emc article antennaire et un nombre inferieur de
sensilles sur l'expansion laterale des palpes gnathochilariaux (NGUYEN DUY - JACQUEMIN,

Figs. 1-6. — Polyxenus oromii n. sp. : 9 paralype de La Palma, I = tele, face dorsale. 2 = palpe droil du gnathochilarium

3 = antenne gauche ; aulres S adulles de La Palma. 4 cl 5 = sensilles des articles VI el VII d'une antenne gauche. 6 =
griffe et 6pine tarsale d une pane IX gauche.

Fk,. 1-6. — Polyxenus oromii n. sp. : 9 La Palma paratype, 1 = head, dorsal side, 2 = right palp of the gnathochilarium,
3 = left antenna ; other adults 9 of La Palma, 4 and 5 = sensillae of left antennal articles VI and VII. 6 = claw and
tarsal spine of a left leg IX.

Figs. 7 4 9. — Polyxenus chalcidicus Conde el Nguyen, 1971 (d'apres les auieurs : p. 1254) : 7 el 8 = sensilles de P article
VI d'un cf a 12 pp et de Panicle VII du d holotype de Pile d'Eubee, 9 = Griffe de la patte III droite du d holotype.

Fla 7 “ 9~ Polyxenus chalcidicus Conde el Nguyen, 1971 I after the authors: p. 1254): 7 and 8 = sensillae of article VI
of a <f with 12 pairs of legs and article VII of the holotype <f from Eubea Island. 9 = claw of right leg III of
holotype <?.

Figs. 10 a 14. —Anopsxenus caboverdus n. sp. : 9 paralype de Santiago, 10 = antenne droite avec detail des sensilles des
articles VI et VII, 1 1 = vulve gauche ; 9 holotype de Santiago. 12 = palpe gauche du gnathochilarium, 13 et 14 =
soie du subcoxa et griffe de la patte IV gauche.

F,G- 10 d ,4; - Anopsxenus caboverdus n. sp.: paratype 9 of Santiago, 10 = right antenna with detailed sensillae of
articles VI et VII, 11 = left vulva; holotype 9 of Santiago. 12 = left palp of the gnathochilarium, 13 and 14 =
subcoxa seta and claw of the left leg IV.

Fig. 15. — Anopsxenus indicus : 9 adulte, griffe de la patte VI gauche.

Ftc. 15. — Anopsxenus indicus : adult 9 , claw of the left leg VI.

Source :

DIPLOPODES PENICILLATES DES ILES CANARIES ET DU CAP VERT

115

Abreviations. Sensilles : basiconique anterieur (a) ; cccloconique (c) : basiconique intermediate (/) ; basiconiquc
posterieur ( p ) ; setiforme anterieur ( s ).

Abbreviations. Sensillae: anterior basiconicum (a); cceloconicum (c); intermediate basiconicum (i); posterior
basiconieum ( p ); anterior setiforrn (s).

Source :

116

MONIQUE NGUYEN DUY - JACQUEM1N

Polyxenus oromii n. sp. (Figs. 1-6)

STATION. — La Palma. Teneguia Colada Costera. n° 5753, P. OROMI leg. : 2 males a 13
pp. (ad.), 7 femelles a 13 pp. (ad.), 1 femelle a 12 pp., 1 male a 10 pp.

Tenerife. Punta del Teno, n° 5760. 6-10. IV. 1988, P. OROMI leg. : 1 male a 12 pp.

Fuertoventura. Puerto Lajas, zone supralittorale, 13. IV. 1987, R. RODRIGUEZ leg. : un
male a 12 pp., une femelle a 12 pp.

Les specimens de La Palma et Tenerife ont tous ete recoltes sur des laves recentes, en
bordure de mer.

DESCRIPTION. — Un male (holotype) et une femelle (paratype) adultes de La Palma,
montes dans le medium II de Marc Andre ont servi pour la description.

Longueurs.- Corps (sans le pinceau caudal) : holotype = 1,50 mm ; paratype = 1,80 mm.
Pinceau caudal : holotype = 0,20 mm. 2eme tarse de la 1 3eme paire de pattes : holotype = 80 jim ;
paratype = 82 pm.

Tete.- Plages posterieures du vertex coalescentes sur la ligne mediane, comprenant deux
rangees de trichomes, Panterieure d'une vingtaine de ces phaneres, la posterieure de 10 a 13. En
arriere de ces rangees, quelques trichomes mediaux : 3, un anterieur et deux posterieurs,
disposes en triangle chez l'holotype et 4 sur une seule rangee chez le paratype. L'orientation et la
taille de ces trichomes sont indiquees sur la Fig. 1 .

Le 6eme article antennaire (Fig. 3), une fois 1/3 a une fois 1/2 plus long que large, porte 2
sensilles basiconiques epais, un sensille setiforme anterieur et un sensille cceloconique
posterieur; le basiconique (/) proche du setiforme (s) est plus long et a peine plus epais que le
posterieur (p) (Fig. 4). Le 7&me article porte toujours 2 sensilles basiconiques epais, separes par
un sensille setiforme a base plus distale, et suivis par un sensille cceloconique posterieur ; 3 a 5
sensilles basiconiques greles entourent le basiconique epais anterieur (Fig. 5).

6 stemmates subegaux : 5 tergaux et un sternal anterieur. Trichobothries typiques.

Labre couvert de petits tubercules acumines, ceux des rangees anterieures ne paraissant pas
plus volumineux que les autres ; 4+4 lamelles marginales.

Palpes du gnathochilarium avec 17 sensilles sur le mamelon ; bras lateraux environ deux
lois plus longs que le plus grand diametre du mamelon, portant 9 sensilles chez les deux sexes.

Tronc.- Collum avec 3 rangees de trichomes ; la rangee intermediate comprenant chez le
male 8 trichomes subspheriques, diriges vers I'avant. Tergites II a VIII avec 44 a 55 trichomes
sur deux rangees subrectilignes reunies lateralement par quelques phaneres formant l'ebauche
d'une rosette.

Soies biarticulees des pattes, reparties ainsi chez le cf holotype : 2 sur les subcoxas I et II
uniquement ; 1 sur tous les coxas et trochanters ; 1 sur les tibias I a XII. Epine du 2eme tarse tres
effilee, de longueur a peine inferieure a la griffe ; processus anterieur du telotarse tres fin et plus
court que la griffe, processus posterieur lamellaire assez etroit et denticule basal effile (Fig. 6).

Male. Subcoxas VIII et IX depourvus d'invaginations glandulaires. Penis sans zone
glabre.

Telson.- De meme type que celui de P. lagurus. Le groupe medio-dorsal avec environ 23
trichomes barbeles chez la femelle. Trichomes appendicules, pourvus de 3 ou 4 expansions sous
la crosse terminale.

AFFINITES. — La nouvelle espece P. oromii est proche de P. chalcidicus par les trichomes
de la tete et des tergites troncaux presentant la meme forme et la meme distribution, mais elle s'en
distingue par un ensemble de caracteres enumeres ci-dessous :

- sensilles des articles antennaires VI et VII : Particle VI ne porte que 2 sensilles
basiconiques, au lieu de 3, l'anterieur faisant defaut (Fig. 7) ; sur Particle VII, le nombre de
sensilles basiconiques greles est plus eleve (4 a 7 au lieu de 2 : Fig. 8) ;

Source :

DIPLOPODES PENICILLATES DES ILF.S CANARIES ET DU CAP VERT

117

- bras lateraux des palpes du gnathochilarium : 9 sensilles au lieu de 12. La femelle adulte
de Lahav (Israel), rapportee a P. chalcidicus avec quelques reserves, ne possede aussi que 9
sensilles (CONDE & NGUYEN DUY - JACQUEMIN, 1971);

- processus anterieur du telotarse beaucoup moins developpe ;

- males depourvus de glandes subcoxales qui tissent les fils signalisateurs des
spermatophores. Les males de P. chalcidicus sont deja depourvus de ces formations a la base
des pattes IX, tandis que les autres especes de Polyxenus en possedent aux pattes VIII et IX.

Le moindre nombre de sensilles aux palpes du gnathochilarium, constate chez P. oromii,
peut etre considere comme une persistance, chez cette espece, de la formule des stades juveniles
n. III et IV de P. chalcidicus porteurs de 9 sensilles ; de meme, l'absence du sensille
basiconique anterieur de ('article antennaire VI, commune aux deux premiers stades larvaires de
P. chalcidicus ; et celle enfin des glandes subcoxales VIII apparaissant ordinairement au stade
VI.

DISCUSSION. — Le genre Polyxenus compte a present 4 especes nominales possedant des
trichomes tergaux globuleux : P. lapidicola Silvestri, in BF.RLESE. 1903 ; P. macedonicus
Verhoeff, 1952 ; P. chalcidicus Conde et Nguyen Duy - Jacquemin 1971 et P. oromii n. sp.

Les types de P. lapidicola ont ete recoltes dans des fissures de rochers sur le rivage marin,
pres de Portici, dans une zone mouillee par mer agitee. L'espece a ete ainsi consideree comme
halophile (ou halobie), d'autant qu'elle a ete recherchee sans succes dans des biotopes secs, sur
les pentes du Vesuve en particulier (SILVESTRI, 1903). La description est tres incomplete,
comme le souligne VERHOEFF (1921) qui citera l'espece de Macedoine (Skoplje) (1941) et de
l ile d'Ischia (1952). CONDE (1950, 1953) signale lapidicola de Saint-Raphael. puis de Corse,
plus ou moins loin de la mer, mais jamais sur le rivage. Tout recemment enfin, ENGHOFF &
SCHEMBRI (1989) attribuent a lapidicola des specimens de Malte, recoltes dans la litiere, sous
d'epais buissons, sans justifier cette determination.

L'examen de 3 syntypes de P. macedonicus, de Skoplje, conserves dans la collection K.
Verhoeff, a Munich, a permis de decrire les groupes de sensilles des articles VI et VII de
l'antenne avec, pour consequence, l'attribution a P. macedonicus des specimens de France
meridionale el de Corse rapportes a P. lapidicola (CONDE & NGUYEN DUY - JACQUEMIN,
1971 : 1256).

P. chalcidicus est largement repandu en Grece continentale : Thessalie, Beotie, Attique
(collections du Museum d'Histoire naturelle de Geneve. CONDE det., inedit) ; la presence de 12
sensilles, a partir du stade a 8 pp., sur les expansions laterales des palpes du gnathochilarium est
ainsi confirmee, de meme que celle d'invaginations glandulaires sur les subcoxas VIII des
males, a partir du stade a 1 0 pp.

P. oromii, enfin. a ete recolte dans des biotopes littoraux, semblables a ceux qui ont livre
les types de P. lapidicola. Cette demiere espece demeurera enigmatique jusqu'a la revision des
types.

Macroxenus enghoffi n. sp. (Figs. 16-26)

STATION. — Fuertoventura. Cumbre Jandia, 14.11.1977, P. OROMI leg. : 4 males a 13
pp. (ad.), 3 femelles a 13 pp. (ad.), 2 males a 12 pp., 1 male a 10 pp.

DESCRIPTION. — 2 males adultes, 1 femelle adulte. le male a 10 pp., montes dans le
medium II de Marc Andre, constituent la serie typique.

Adultes.

Longueurs.- Corps (sans le pinceau caudal) : males = 3,30 mm (holotype) et 3 mm ;
femelle =4 mm. Pinceau caudal = 0,40 mm (holotype). Trichomes du vertex = 0,20 mm
(holotype). 2eme tarse de la 13eme paire de pattes : males = 172 pm (holotype) et 179 pm ;
femelle = 188 pm.

MONIQUE NGUYEN DUY - JACQUEMIN

I 18

Tete.- Plages posterieures du vertex allongees
transversalement et tres fortement obliques, plus de deux
fois et demi plus longues que leur ecartement. Elies
comprennent chacune 29 trichomes chez I'hololype et 24
chez la femelle formant une rangee anterieure de 13 a 18
trichomes rapproches les uns des autres et une rangee
posterieure de 8 a 10 trichomes plus espaces que les
precedents, le plus lateral etant nettement plus eloigne des
autres (Fig. 18) ; parfois une 3emc rangee tres courte de 2 a
4 trichomes s'ajoute entre les deux autres, vers la region
centrale de la tete.

Les longueurs relatives des articles antennaires sont
donnees par la Figure 17. Article VI deux fois plus long
que large portant 13 (holotype et femelle) a 16 sensilles
basiconiques subegaux, tres effiles a l'apex, inseres sur
une surface triangulaire dont une base suit la limite distale
de Particle ; a ceux-ci s'ajoutent un sensille cceloconique
posterieur (c) et un sensille setiforme anterieur. L'article
VII porte toujours deux sensilles basiconiques subegaux,
separes par un sensille setiforme et accompagnes d'un
sensille cceloconique posterieur.

Trichobothries subegales, a funicule grele. 8
stemmates : 6 dorsaux, un lateral et un ventral (Fig. 18).

Fig. 16. — Male adulte de Macroxenus enghoffi n. sp, habitus, face
dorsale.

Fig. 16. — Adult male of Macroxenus enghoffi n. sp, habitus, dorsal
view.

Marge anterieure du labre bordee de 15 (9) a 22 (holotype, Fig. 20) lamelles arrondies,
aussi hautes que larges, sauf aux extremites laterales. Face externe couverte de granules pourvus
d'une petite pointe apicale ; les plus marginaux, sur 1 a 3 rangs, sont plus volumineux, les
suivants diminuent progressivement de taille jusqua la moitie de la largeur du labre. 10
(holotype) a 13 soies greles bordent la limite clypeale, elles sont presque deux fois plus courtes
que la plus grande largeur du labre.

Figs. 17-26. Macroxenus enghoffi n. sp. : cf holotype de Fuertoventura, 17 = antenne gauche, face dorsale. avec detail

des sensilles des articles VI et VII, 18 = demi-tete gauche, face dorsale, 19 = palpe droit du gnathochilarium, 20 =
labre (une partie seulement des granules est dessinee), 21 = trichome hamule du telson, 22 = soie du tibia de la
patte I gauche, 23 = soie du subcoxa de la patte III gauche, 24 = griffe de la patte X droite ; 9 adulte n°2, 25 = tarse
de la patte I droite avec detail de lepine et de la griffe, 26 = trichome a crochets specifique des 9.

Fig. 17-26. — Macroxenus enghoffi n. sp.: holotype cf of Fuertoventura, 17 = left antenna, dorsal side, with detailed
sensillae of articles VI et VII, 18 = half left head, dorsal side, 19 = right palp of the gnathochilarium, 20 = labrum
(only a part of the granules is drawn), 2/ = trichome with hooks of the telson, 22 = tibial seta on left leg I, 23 =
subcoxal seta on left leg 111, 24 = claw of the right leg X; adult 9 n°2, 25 = tarsus of the right leg 1 with detailed
spine and claw, 26 = trichome 9 - specific hooks.

Figs. 27 a 30. — Macroxenus cf. enghoffi : ind. h 8pp de S. Antao, 27 et 28 = sensilles des articles VI et VII de l'antenne
gauche, 29 = palpe gauche du gnathochilarium, 30 = soie du subcoxa de la patte I gauche.

Fig. 27 a 30. - Macroxenus cf enghoffi ; ind. (8 pairs of legs) of S. Antao, 27 and 28 = sensillae of left antennal
articles VI et VII, 29 = left palp of the gnathochilarium, 30 = subcoxal seta of the left leg I.

Source :

DIPLOPODES PENICILLATES DES 'iLES CANARIES ET DU CAP VERT

119

Source : MNHN, Paris

120

MONIQUE NGUYEN DUY - JACQUEMIN

Palpes dll gnathochilarium a expansion laterale environ 4 fois plus longue que le plus
grand diametre du mamelon, portant 34 et 37 sensilles chez l'holotype (Fig. 19), 43 et 44
sensilles chez l'autre male et 15 sensilles chez une femelle'.Tous ces sensilles presentent une
pseudoarticulation a peu de distance de 1'apex. Le mamelon des males porte 1 8 a 20 sensilles et
celui de latemelle 20 et 21 sensilles, de longueurs inegales, mais pseudoarticules comme ceux
de l'expansion, a l'exception des 7 antero-internes courts.

Tronc.- Les plages laterales des tergites (collum et tergites IX-X exceples) sont 2 a 3 fois
plus courtes que leur ecartement. Elies portent de 43 a 65 trichomes (cf ), 33 a 49 (9) et sont
reunies entre elles par une rangee marginale posterieure sinueuse de 14 a 21 trichomes au
collum. 26 a 48 (c? ) et 23 a 35 (9) aux tergites II a VIII ; en avant de la rangee marginale 5 a 14
trichomes non alignes sont presents sur les tergites II a X de l'holotype, II a IX des 2 autres
specimens et 2 sont presents uniquement sur le collum de l’holotype. Tous ces trichomes sont
orientes vers l'arriere.

Soies des subcoxas (Fig. 23), coxas et trochanters a funicule fusiforme glabre ; il y en a
une seule au subcoxa I. deux ou trois aux subcoxas II a XIII des cf et II a XII de la 9 ; une seule
au bord distal de chaque coxa et trochanter. Deux soies (parfois 1 ou 3) de meme type, mais
beaucoup plus petites, se rencontrent sur la region moyenne des trochanters ; une seule au bord
distal des femurs des pattes II a XIII des o’ , II a XII de la 9 et au bord distal de chaque tibia
(Fig. 22). L'epine du 2eme article des tarses est prolongee par une pointe extremement fine qui
souvent n'apparait pas au microscope ; sa longueur est voisine ou legerement superieure (1,1 a
1,2 fois) a celle de la griffe de la patte correspondante (Fig. 25). Griffe courte et trapue, pourvue
de deux dents accessories (anterieure et posterieure) subegales, longues et effilees (Figs. 24,
25). Processus telotarsaux setiforme et lamellaire presents.

c? . Penis sans zone pyriforme glabre. Vastes invaginations glandulaires sur les subcoxas
VIII et IX.

Telson.- II appartient au type II, base sur le genre Macroxenus , et defini par CONDE
(1970) : de chaque cote, 10 a 13 grosses embases de trichomes barbeles (c), groupees en une
plage subcirculaire, occupent une echancrure de la marge anterieure du pinceau ; au bord antero-
interne de la plage, 1’embase de b se distingue des autres par sa paroi un peu plus mince et sa
forme plus allongee ; un seul trichome a en avant de chaque pinceau. Trichomes hamules du
pinceau caudal portant generalement 3 crosses (Fig. 21), parfois 4. En plus des trichomes
barbeles et hamules des males, les femelles possedent un type different de trichomes hamules
(Fig. 26). La hampe de ces trichomes est garnie, vers l'apex, d'une rangee rectiligne de crochets
a pointe orientee vers la base du poil, de taille decroissant rapidement jusqu'a devenir de petites
dents sur plus de la moitie de la longueur du phanere ; ces dents sont ensuite remplacees par des
dents d'orientation inverse, c'est a dire dirigees vers l'apex, comme c'est le cas pour tous les
trichomes. Ces poils caudaux, specifiques des femelles, paraissenl occuper une position
ventrale, mais leur emplacement est impossible a reconnaitre d'apres la structure de leur cupule
d insertion ne pouvant etre distinguee de celle des autres trichomes hamules. C'est la seconde
fois qu’un caractere sexuel secondaire concernant la structure d'un type de poil du pinceau
caudal est observe chez un penicillate. La premiere fois (CONDE & NGUYEN DUY - JACQUEMIN.
1990), il sagissait du male du polyxenide Unixenus aff. broelemanni (Conde & Jacquemin,
1963), qui est depourvu des trichomes hamules presents chez la femelle.

1 . Chez Macroxenus rubromarginatus , les palpes du gnathochilarium des males et des femelles portent le meme
nombre de sensilles (24-26) sur I expansion laterale, mais chez M. caingangensis, il y cn a 16 au plus chez les
femelles et 18 a 28 chez les males (CONDE & MASSOUD. 1974).

D1PLOPODES PENICILLATES DES ILES CANARIES ET DU CAP VERT

121

Immature.

c? a 10 pp.

Les articles antennaires VI portent 12 et 13 sensilles basiconiques greles ; les palpes du
gnathochilarium onl chacun 20 sensilles sur le mamelon et respectivement 17 et 19 sensilles sur
l'expansion laterale. Marge anterieure du labre bordee par 5+6 lamelles hyalines laterales. Une
invagination glandulaire sur le subcoxa VIII. Pas de bourgeons externes ; faisceaux transitoires
presents.

Plage subcirculaire de 6 et 7 trichomes c ; 3 trichomes a sur le telson.

AFFINITES. — Les deux especes de Macroxenus decrites jusqu'ici, M. rubromarginatus
(Lucas, 1846), d'Afrique septentrionale, et M. caingangensis (Schubart, 1944), du Bresil. sont
excessivement voisines l'une de l'autre (CONDE, 1971 : 633). Les sensilles basiconiques de
1' article VI sont disposes en une rangee transverse, rectiligne, de 5 et parfois 6 unites, avec un
sensille cceloconique situe avant les 2 basiconiques posterieurs, soit 3+ccel.+2 ou 4+ccel.+2.
Les nombreux sensilles de Particle VI de M. enghoffi, disposes en un groupe subtriangulaire,
evoquent davantage Macroxenodes bartschi (Chamberlin, 1922), redecrit assez recemment
(NGUYEN DUY - JACQUEMIN & CONDE, 1984).

Les especes a sensilles gnathochilariaux pseudoarticules, reparties entre les genres
Macroxenus Brolemann, 1917, Macroxenodes Silvestri, 1948 et Chilexenus Silvestri, 1948 ont
en commun un telson du type II qui est unique dans le groupe - de meme que les sensilles
pseudoarticules - et constitue un argument de parente indiscutable. Toutefois, la definition de ces
trois genres est peu satisfaisante et, dans l’attente d'une revision des types, l'attribution
generique des especes doit etre consideree comme provisoire.

Macroxenus cf. enghoffi ( Figs 27-30)

Station. — S. Antao. Around villa de Ribeira Grande, 4-7.XII. 1988, A. VAN HARTEN
leg. : 1 ind. a 8 pp.

Description.

Longueurs.- Corps (sans pinceau caudal) = 1,70mm. 2e tarse de la VUIe paire de pattes =
99 pm.

Tete.- Plages posterieures du vertex composees de 10 trichomes : 7 a la rangee anterieure,
3 a la rangee posterieure. 15 et 16 sensilles basiconiques greles (Fig. 27) sur le 6eme article
antennaire ; 20 et 21 sensilles sur les mamelons des palpes gnathochilariaux, 12 et 13 sensilles
sur les expansions laterales (Fig. 29) : la pseudo-articulation des sensilles est difficile a observer
a ce stade juvenile.

Telson.- Plages subcirculaires comprenant 4 grosses embases de trichomes c, et 6
trichomes a formant les groupes lateraux.

AFFINITES. — Voisin de M. enghoffi n. sp., par le nombre et la disposition des sensilles
basiconiques du 6cme article antennaire, il s'en distingue par la forme de ces phaneres qui sont
plus longs et plus greles, et par les soies des subcoxas (Fig. 30), des coxas et des trochanters
dont le funicule est pubescent et proportionnellement plus court. Les dents accessoires de la
griffe paraissent plus trapues que celles de M. enghoffi, mais un adulte de cette forme incertaine
serait necessaire pour la comparer a M. enghoffi .

Anopsxenus caboverdus n. sp. (Figs. 10-14)

(= Anopsxenus indicus in : ENGHOFF, 1993)

REMARQUE. — Lors d'une premiere identification, j'avais rapporte les specimens du Cap
Vert a une espece decrite de Bombay, Anopsxenus indicus Conde et Jacquemin, 1963, type et
seul representant du genre. Toutefois, un examen plus attentif m'a conduit a considerer qu'il
s’agit en fait d'une espece distincte qui est decrite ici.

122

MONIQUE NGUYEN DUY - JACQUEMIN

Station. — Santiago. S. Jorge dos Orgaos, n° 2228, VII. 1989, A. van Harten leg. : 2
femelles a 13 pp (ad.), 2 ind. a 6 pp., 1 ind. a 5 pp., 1 ind. a 3 pp.

Description. — Les adultes sont designes respectivement comme holotype et paratype.
La tete dissequee de l'holotype, montee dans l'Euparal, preparation n° 1363, est conservee au
Musee de Zoologie de Copenhague ; son corps est monte dans le Medium II de Marc Andre,
ainsi que le paratype et les juveniles.

Adultes.

Longueurs.- Corps du paratype (sans le pinceau caudal) = 2,30 mm. Pinceau caudal =
0,25 mm. 2£me tarse de la 13eme paire de pattes = 150 (holotype) et 143 (im.

Teguments.- Aucune trace de pigment n'est decelable.

Tete.- Plages posterieures du vertex allongees, plus de trois fois plus longues que leur
ecartement, comprenant chacune une rangee anterieure de 1 8 trichomes et une posterieure de 4
(paratype). Les antennes sont conformes a la definition du genre (3 et 2 sensilles basiconiques
respectivement en VI et VII, mais le cceloconique manque en VI et il est dedouble en VII (Fig.
10), ce qui est typique des antennes regenerees (une antenne a 7 articles du paratype est en cours
de regeneration) (NGUYEN DUY - JACQUEMIN, 1972).

Trichobothries, labre et gnathochilarium (Fig. 12) comme chez A. indicus. 19 et 20
sensilles sur les mamelons du gnathochilarium (paratype).

Tronc.- Plages laterales du collum avec 66 et 70 trichomes (holotype) ; aux tergites
suivants, les plages comprennent de 35 a 48 trichomes et sont unies par une rangee marginale
ininterrompue de 40 a 48 trichomes (holotype).

Pilosite des pattes identique a celle de A. indicus, mais les vulves portent de 13 a 16 soies,
arquees pour la plupart, et differentes en cela de celles portees par les subcoxas, coxas et
trochanters (Fig. 1 1). Rapport tarse/griffe = 10 aux pattes XII et XIII, 9 a la patte I de la femelle
paratype. Les denticules situes a la base de la griffe sont bien developpes, le posterieur de
longueur egale au 1/3 de la griffe qui possede une faible dent stemale (Fig. 14).

Telson.- Groupes lateraux avec 9 et 10 trichomes a.

Immatures : Stades I, III et IV.

Le 6eme article antennaire du stade I (3 pp.) n’a que 2 sensilles basiconiques et un sensille
ccEloconique posterieur ; le 3eme sensille basiconique, anterieur aux 2 autres, n'apparatt qu’aux
stade II ou III ; il est present en effet chez l'individu a 5 pp.

7 sensilles sur les expansions des palpes gnathochilariaux et 20 a 22 sensilles sur les
mamelons. Nombre de trichomes des plages posterieures de la tete augmentant avec la croissance
: 6+2 au stade I, 8-9+2 au stade III, 9-10+2 au stade IV ; il en est de meme pour les trichomes a
du tergite telsonien : 5+5 (stade I), 7+7 (stade II), 8+9 (stade III). Les trichomes C2 et b du
telson sont presents chez les larves III et IV ; ce mode d'acquisition est probablement identique a
celui observe chez Monographis tamoyoensis et Pauropsxenus vilhenae ou ces trichomes
n'apparaissent qu'au stade III (NGUYEN DUY - JACQUEMIN, 1973).

AFFINITES. — Anopsxenus Conde et Jacquemin, 1963, a pour type le seul polyxenide
aveugle et pigmente connu. Un pigment brun qui occupe des plages de forme definie sur la tete,
le tronc et les pattes, a subsiste apres un sejour de 3 annees en alcool. Anopsxenus indicus,
recolte a Bombay par P. Remy en 1959, est represente par une femelle a 12 pp., choisie comme
lectotype en raison d'un meilleur etat de conservation, et une femelle paralectotype a 13 pp.
(ad.).

La nouvelle espece est totalement depourvue de pigmentation et il n'y a aucune preuve que
le sejour en alcool ou l'exposition a la lumiere aient ete responsables de la destruction d'un
pigment. Les griffes courtes (rapport tarse/griffe = 10 vs 6,4 chez A. indicus ) et trapues, a
volumineux denticules basilaires et a petite dent stemale, sont tres differentes des griffes longues
et greles de A. indicus , presentant de minuscules denticules a leur base (Figs. 14, 15). Les
nombreux phaneres des vulves manquent chez indicus. Enfin, le 6cme article antennaire est plus
allonge chez caboverdus que chez indicus : rapport L/l = 1,40-1,60 et 1,70- 1,80 vs 1,13-1,28

Source :

DrPLOPODES PENICILLATES DES ILES CANARIES ET DU CAP VERT

123

pour le lectotype de indicus, on notera cependant qu'il s’agit d'antennes regenerees pour
caboverdus.

Ces differences nous paraissent justifier le statut specifique accorde a present aux
specimens du Cap Vert.

Tableau 1. — Liste des espkces dc Macaronesie.
Table /. — Check- list of Mcicaronesian species.

Families

Esp&ces

CANARIES

CAP VERT

MADERE

AZORES

Polyxenus fasciculatus

Gran Canaria
Tenerife
Gomera
Hierro

Madeira

Selvagem Grande

Polyxenus lagurus

(forme bisexu£e)

Sao Miguel
Pico

POLYXENIDAE

Polyxenus oromii n. sp.

Fuertoventura
La Palma
Tenerife

Macroxenus enghoffi n. sp.

Fuertoventura

Macroxenus cf. enghoffi

S. Antao

Anopsxenus caboverdus n. sp.

S. Tiago

LOPHOPROCTIDAE

Lophoproctinus inferus maurus

Fuertoventura
Gran Canaria
Tenerife

Lophoturus madecassus

S.Tiago

FAMILLE DES LOPHOPROCTIDAE

Lophoproctinus inferus maurus Marquet & Conde, 1950.

STATIONS. — Fuertoventura. Jandia : Barranco del Ciervo, Morro de Cavedero N of
Morro Jable, grassy, stony W slope, 700 m, Astericus etc., under stones n°2660 : 6 femelles a
13 pp. (ad.), 1 femelle a 12 pp., 1 femelle a 10 pp., 1 individu a 8 pp. — Localite precedente,
grassy ridge and N & E slopes, 700 m, Astericus , etc., under stones, n° 2631, 4.1.1990 : 2
femelles a 13 pp. (ad.). — Gran Canaria . Roque Bentayga, SW slope, 1100 m. Euphorbia
obtus, Kleinia, under deeply imbedded stone, n° 2632, 1.1.1990 : 1 male a 13 pp. (ad.). Tous
recoltes par M. BAEZ, H. ENGHOFF. — Tenerife. Barranco de Las Cuevas, Teno Alto,
5.XI.1989 : 1 femelle a 13 pp. (ad.), R. RODRIGUEZ leg.

REPARTITION. — Cette sous-espece, decrite d'Algerie (Dar-el-Oued) par MARQUET &
CONDE (1950), se distingue de la forme typique, connue d'ltalie aux environs de Portici
(SELVES TRI, 1903), par sa chetotaxie cephalique et la presence d'une dent dans la concavite de la
griffe. Elle a ete retrouvee en Algerie (Blida), au Maroc occidental (Safi, Sidi Kacem,
Marrakech) et oriental (Oudja), et en Tunisie (Le Kef) (CONDE, 1954).

Source :

124

MONIQUE NGUYEN DUY - JACQUEMIN

Lophoturus madecassus Marquet & Conde, 1950.

Cette espece, signalee de Santiago (lie du Cap Vert) par ENGHOFF (1993), presente une
tres vaste repartition circum-tropicale : decrite de Madagascar (Tulear), elle a ete signalee
d'Afrique (Hoggar, Cote d' Ivoire), des Antilles (Jamai'que), de Floride (Dry Tortugas) et d'lles
du Pacifique Sud (Archipel des Tonga, Atoll Suvorov).

BIOGEOGRAPHIE

Deux especes a large repartition geographique sont presentes aux Canaries : Polyxenus
fasciculatus et Lophoproctinus inferus maurus (Fig. 31).

Fig. 31. — Repartition locale et mondiale des especes de penicillates des ties Canaries et du Cap Vert.
Fig. 31. — Local and world distribution of Penicillata species of Canary Islands and Cape Verde Islands.

Source : MNHN, Paris'

DIPLOPODES PENICILLATES DES ILES CANARIES ET DU CAP VERT

125

P. fasciculatus est 1 'espece la plus repandue aux Canaries, peuplant 13 stations : 2 a
Hierro, 1 a Gomera, 6 a Tenerife et 4 a Gran Canaria. El le occupe la portion meridionale de
l'Amerique du Nord (Centre et Sud-Est dcs Etats-Unis), les Bermudes et Madere (Ribero Seco,
Pico do Gato, Selvagein Grande), alors que la forme bisexuee de P. lagurus est implantee en
Eurasie et aux Azores (CONDE & NGUYEN DUY - JACQUEMIN, 1994) ou elle a ete identifiee a
Sao Miguel par CONDE (1961) et plus recemment, par moi-meme, a Pico, sur des rochers de
laves a Costa Cachorro (leg. OROMI 1987). On notera que si l'Archipel des Azores est plus
eloigne de l'Europe que les lies Canaries et que Madere, il est en revanche plus septentrional. La
limite geographique entre ces deux especes n’est pas precisee, mais on ne les a jamais vu
cohabiter. La forme parthenogenetique de P. lagurus est presente dans le Nord de l'Amerique et
de l'Eurasie ; elle parait mieux adaptee aux climats continentaux (CONDE & NGUYEN DUY -
JACQUEMIN. 1994) et la temperature moyenne pourrait etre aussi le facteur limitant l'extension,
vers le Nord, de P. fasciculatus ( ce qui est net aux Etats-Unis), comme celle de P. lagurus
bisexue en Eurasie.

Lophoproctinus inferus maurus a ete recolte sur 3 des lies Canaries : Fuertoventura, Gran
Canaria et Tenerife ; elle est connue sur le continent le plus proche, 1'Afrique du Nord (Maroc,
Algerie, Tunisie), situee a 90 kin de Fuertoventura.

Autre exemple d’espece a large repartition geographique, Lophoproctus madecassus, le
seul penicillate a 1 1 paires de pattes, signale par ENGHOFF(1993) de Santiago (Cap Vert) est une
espece tropicale : Hoggar, Cote d'Ivoire, Madagascar. Floride, Jamaique, Pacifique Sud
(Tonga, Suvorov).

Les especes nouvelles trouvees aux Canaries peuvent-elles etre considerees comme des
endemiques? M. enghoffi est eloigne de M. rubromarginatus, d'Afrique septentrionale et de M.
caingangensis, du Bresil, par la disposition des sensilles du Vie article antennaire. Anopsxenus.
caboverdus , en revanche, est voisin de la seule autre espece du genre, A. indicus, de Bombay.
P olyxenus orotnii enfin est tres proche de P. chalcidicus , du Bassin de la Mediterranee (Grece,
Israel). Deux specimens a 10 pp. (male et femelle) de Karpathos (Pigadia, 12. IV. 52, H.
SCHMALFUSS leg.), determines par B. CONDE, sont rapportes sous reserve, a cette espece, dans
l'attente de specimens adultes.

REMERCIEMENTS

J’adresse mes plus vifs remcrciements a Monsieur le Professeur B. Cond£ pour ses conseils dans la redaction de ce
manuscrit et a Jacques Rebiere pour la realisation de Ticonographie.

REFERENCES

BrOlemann, H. W., 1896. — Myriapodes provenant des Campagnes scientifiques de YHirondelle et de la Princesse
Alice. Bull. Soc. zool. Fr. .21 : 198-204.

Cond£, B. , 1950. — Un diplopode nouveau pour la France. L'Entomologiste , 6 : 109-116.

Cond£, B. , 1953. — Diplopodes Penicillates de Corse. Bull. Soc. zool. Fr.. 78 : 33-35.

Cond£ B., 1954. — Diplopodes Penicillates d’Afrique septentrionale. Bull Mus. natl. Hist. nat.,2eme Ser.. 26 : 496-

500.

Cond£, B., 1961. — Diplopodes Penicillates des Azores et de Madere. Bol. Mus. municipl. Funchal 14 : 7-10.

Co.nd£, B.. 1970. — Essai sur 1'evolution des Diplopodes Penicillates. Bull. Mus. natl. Hist, nat., 2eme Ser., 41, suppl.
2 : 48-52.

Cond£, B.. 1971. — Diplopodes penicillates des nids bresiliens de Camponotus rufipes. Rev. Ecol. Biol. Sol. 8 : 631-
634.

Cond£, B.. 1972. — Presence aux Bermudes de Diplopodes Penicillates et d’Arachnides Palpigrades. Rev. Ecol. Biol.

501, 9: 127-129.

Conde. B. & Massoud, Z., 1974. — Diplopodes Penicillates du Bresil et de la Republique Argentine. Rev. Ecol. Biol.
Sol.. 11 : 223-232.

Cond£, B. & Nguyen Duy - Jacquemin, M., 1971. — Penicillates d’Israel rassembles par G. Levy. Bull. Mus. natl. Hist,
nat., s. D. 42 : 1251-1258, 1970.

Cond£ B. & Nguyen Duy - Jacquemin. M., 1990. — Decouverte d'un caractere sexuel secondaire nouveau chez le male
d'un Polyxenid6 (Myriapodes, Penicillates). Ber. nat.-med. Verein Innsbruck, suppl. 10 : 57-62.

126

MONIQUE NGUYEN DUY - JACQUEMIN

Cond£ B. & Nguyen Duy - Jacquemin, M., 1994. — Parthenogen£se et reproduction bisexu£e dans le complexe de
Polyxenus lagurus (L.). Biogeographica, 70 : 41-48.

ENGHOFF, H., 1993. — Cape Verdean millipedes (Diplopoda). Tropical. Zool.. 6 : 207-216.

Enghoff, H. & Schembri, J., 1989. — The Millipedes of the maltese islands (central mediterranean). Boll. Soc. ent.
iial., Genova , 120 : 164-173.

MARQUET M. L. & Conde. B.. 1950. — Contribution a la connaissance des Diplopodes Penicillates d'Afrique et de la
Region madecasse. Mem. Inst. sci. Madagascar , s6r. A. 4 : 113-134,

Nguyen Duy - Jacquemin, M., 1972. — Regeneration antennaire chez les larves et les adultes de Polyxenus lagurus
(Diplopode, Penicillate). C. R. Acad. Sc. Paris (D). 274 : 1323-1326.

Nguyen Duy - Jacquemin, M., 1973. — Contribution & la connaissance de l’anatomie cephalique, des formations
endocrines et du developpement postembryonnaire de Polyxenus lagurus (Diplopodes penicillates). These Doctoral
d'etat es-Sciences natureiles, UPMC, Paris VI, 148 pp.

Nguyen Duy - Jacquemin. M., 1976. — Etude de la variability des caracteres de deux espfcces du genre Polyxenus. P.
lagurus (L.) et P. fasciculatus Say (Diplopode Penicillate). basee sur les mensurations d'articles tarsaux. Bull. Mus.
natl. Hist. nat. Paris, 3 eme Ser..Zool. 249. 356 : 105-118.

Nguyen Duy - Jacquemin, M. & Conde, B., 1984. — Nouvelle description et statut de Polyxenus bartschi Chamberlin
(Diplopodes, Penicillates). Bull. Mus. natl. Hist, nat., Paris. 4eme Ser.. 6. A. 3: 721-728.

SlLVESTRi. F.. 1903. — In : BERLESE, Acari, Myriapoda et Scorpiones hucusque in Italia reperta. Fasc. 98, n° 4.

Verhoeff, K. W.. 1921. — Ueber Diplopoden der Riviera und einige alpenadische Chilathognathen. Arch. natg. Berlin,
87 A : 1-110.

Verhoeff, K. W., 1941. — Zur Kenntnis der Polyxenus- Arten. Zool. Am.. 133 : 259-264.

Verhoeff, K. W., 1952. — Weitere Beitrage zur Kenntnis der Isopoden- und Diplopodenfauna von Ischia und Capri.
Bonn. Zool. Beitr., 3 : 125-150.

Source :

Une approche des Diplopoda Penicillata de l'Amerique

du Nord

Bruno CONDE

Musee dc Zoologie, 34 rue Sainte-Catherine, F-54000 Nancy. France

RESUME

Neuf esp£ces ou sous-esp£ces nominales de Diplopoda Penicillaia ont etc repertoriees en Amerique, au nord du Mexique.
et attributes, a une exception pres, au genre holarctique Polyxenus. Trois d’entre elles sont actuellement inclassables.
mais les six autres ont fait I’objet d'une revision. Quatre especes appartiennent de fait au genre Polyxenus ( anacapensis ,
fasciculatus, lagurus, pugetensis ), une au genre subtropical Macroxenodes ( bartschi ) et une au genre Lophoturus
( madecassus ). Nous ajoutons ici une seconde espece (cf. aequatus) a ce dernier genre. Un neotype de P. fasciculaius , la
premiere espece americaine decrite, est designe afin d’eviter toute confusion avec les formes bisexute ou
parthtnogenetique de P. lagurus.

ABSTRACT

An approach to the Diplopoda Penicillata from North America.

Nine nominal species or subspecies of Diplopoda Penicillata have been recorded from America, north of Mexico, and
assigned , with one exception, to the holarctic genus Polyxenus. Three of them are unclassifiable for the time, but the
six others were revised. Four of them belong in fact to Polyxenus ( anacapensis . fasciculatus, lagurus, pugetensis ). one to
the subtropical genus Macroxenodes ( bartschi ) and one to the wide ranging genus Lophoturus ( madecassus ). We add here
a second species (cf. aequatus) of the last genus. A neotype of P. fasciculatus , the first described american species, is
designed to avoid confusions with the bisexual or parthenogenetic stocks of P. lagurus.

INTRODUCTION

Neuf formes nominales de Penicillata ont ete decrites ou citees d' Amerique septentrionale
au nord de Mexico. Ce sont. dans l'ordre chronologique :

1- Polyxenus fasciculatus Say. 1821. Southern States

2- Polyxenes (sic) fasciculatus var. pallidas Ryder, 1878. Maryland

3- Polyxenus pugetensis Kincaid. 1898. Westerns Washington

4- Polyxenus bartschi Chamberlin, 1922. Florida (Tortugas)

5- Polyxenus lagurus (L.). Nova Scotia

6- Polyxenus fasciculatus var. victoriensis Pierce, 1940. Texas (Victoria)

7- Polyxenus anacapensis Pierce, 1940. California (Anacapa Is.)

8- Polyxenus tuberculatus Pierce. 1940. Texas (Sabinal)

9- Lophoturus madecassus (Marquet et Conde, 1950). Florida (Loggerhead Key)

Conde. B., 1996. — Une approche des Diplopoda Penicillaia de l'Amerique du Nord. In: Geoffroy. J.J..
Mauries, J.-P. & Nguyen Duy - Jacquemin. M., (eds). Acia Myriapodologica. Mein. Mus. natn. Hist. nat.. 169 : 127-
135. Paris ISBN : 2-85653-502-X.

128

BRUNO CONDE

Nous y ajoutons :

10- Lophoturus cf. aequatus (Loomis, 1936). Florida (Key Largo).

Polyxenus bartschi a ete attribue a Macroxenodes (NGUYEN DUY - JACQUEMIN & CONDE,
1984) et les caracteres des trichomes telsoniens ecartent aussi fasciculatus victoriensis et
tuberculatus du genre Polyxenus, sans que Ton puisse leur assigner une position generique
convenable en l'absence d'un nouvel examen du materiel typique. En revanche, il est probable
que fasciculatus pallidus soit un Polyxenus authentique.

ENUMERATION

Polyxenus fasciculatus Say, 1821 (Fig. 1A, B. C)

La description originale, citee par PIERCE (1940), ne permet pas de reconnaitre l'espece,
aucun type n'est connu et la mention “Inhabits the Southern States” rendait fort incertaine la
recherche de topotypes. II existe heureusement une biographie de SAY, consultee pour nous par
le Dr. Richard L. HOFFMAN, qui precise que SAY n'a effectue qu'une seule expedition dans les
Etats du Sud avant 1821. de Philadelphie a Savannah (Georgia) et jusqu'au Nord-Est de la
Floride, en suivant les ties cotieres, ou il a recolte de nouvelles especes d’lnsectes. HOFFMAN
(in litt. 10.02.65) conclut : “the locus typicus of P. fasciculatus Say is the costal part of Georgia,
between Savannah and Jacksonville”.

Neotype.

Devant la necessite de designer un neotype, en raison d'une situation confuse entre cette
espece et les lignees bisexuee ou parthenogenetique de Polyxenus lagurus, le Dr. HOFFMAN
nous a communique un male adulte (13 pp.) etiquete : “Georgia : Glynn County : St. Simon's
Island, Brunswick, 19 june 1977 R. L. HOFFMAN leg.”, depose au Laboratoire de Zoologie,
Arthropodes, du Museum national d’Histoire naturelle de Paris.

Longueurs.- Corps = 2,06 mm (extension moyenne) ; pinceaux telsoniens = 0,54 mm
(trichomes en crosses) et 0,68 mm (trichomes barbeles) ; ta I = 89,7 pm, ta XIII = 105 pm, ta
XHI/ta I = 1,17.

Tete.- Plages posterieures du vertex 3 fois plus longues que leur ecartement (92-95/30)
comprenant une rangee anterieure de 13 et 15 trichomes diriges vers l'avant et une rangee
posterieure de 8 diriges vers l'arriere ; une paire de trichomes parasagittaux en arriere des plages.
Calice de la trichobothrie la plus interne beaucoup plus petit que les autres. 6 stemmates
subegaux.

6eme article antennaire environ 1 fois 1/10 plus long que large (L/l = 1,1 1) (Fig. 1A). A
l’antenne gauche, 8 sensilles basiconiques, dont un epais, entre un sensille setiforme a base
renflee anterieur et un sensille coeloconique posterieur ; le basiconique epais, plus court que ses
voisins, est situe dans la moitie anterieure du groupe et entoure de 7 basiconiques greles1.
L'antenne droite, atypique, est depourvue du basiconique epais. Article VII avec 4 basiconiques
(2 greles, 2 epais) et un coeloconique (Fig. IB. C).

Marge anterieure du labre pourvue de lamelles hyalines imbriquees, non denombrables
avec certitude chez ce specimen fortement eclairci. Face externe couverte de granules a courte
pointe apicale, ceux des premieres rangees marginales plus gros que les suivants.

Palpes du gnathochilarium avec 14 sensilles a gauche et 12 a droite sur l'expansion
laterale2, et 17 sur le mamelon.

1 Le nombre total de sensilles basiconiques greles varie de 7 a 17 selon les indi vidus et les populations (Nguyen Duy -
Jacquemin, 1976 : 113, Tableau 4).

2 De 10 a 15 dans les proportions suivantes : 1. 14, 139, 9. 4. 1 (Nguyen Duy - Jacquemin, 1976 : 115).

Source :

DIPLOPODES PENICILLATES DE L'AMERIQUE DU NORD

129

Fig. 1. — Polyxenus fasciculatus Say. A : articles VI et VII de I'antenne gauche du neotype, face tergale. B : sensilles de
Particle VI droit dun male adulte de St Bernard Pa. Louisiane. C : sensilles de Particle VI gauche dune femelle
adulte de Baton Rouge, Louisiane.- Polyxenus pugetensis Kincaid, femelle adulte de Oak Creek, Oregon. D .
articles VI et VII de I'antenne gauche,' face tergale. 1-11 = sensilles basiconiques greles ; b-b2 = sensilles
basiconiques epais ; c = sensille coeloconique ; s = sensille setiforme a base renflee.

FlG. I. — Polyxenus fasciculatus Say. A: left antennal articles VI and VII of the neotype, tergal side. 8: sensilla of the
right article VI in an adult male from St Bernard Pa, Louisiana. C: sensilla of the left article VI of an adult female
from Baton Rouge, Louisiana. Polyxenus pugetensis Kincaid, adult female from Oak Creek, Oregon. D: left
antennal article VI and VII, tergal side. 1-1 1 = thin hasiconic sensilla ; b-b2 = thick basiconic sensilla; c =
coeloconic sensilla; s = setiform sensilla with a bulbous base.

Tronc.- Chetotaxie tergale du type de P. lagurus , les trichomes des deux rangees
marginales etant toutefois disposes moins regulierement.

Epine du 2eme article du tarse et griffes comme chez P. lagurus. Invaginations glandulaires
sur les subcoxas VIII et IX.

130

BRUNO CONDE

Telson.- Trichomes des plages subtriangulaires medio-dorsales : 22 et 21. Trichomes
principaux des pinceaux termines en une crosse appendiculee.

Repartition.- J'ai determine l'espece des Etats suivants : Illinois, District of Columbia,
North Carolina, Tennessee, Florida, Alabama, Arkansas, Mississippi, Louisiana et Texas, outre
le neotype de Georgia. Au total : 175 specimens repartis entre 21 males et 44 femelles a 13 pp.
(ad.), 8 males et 24 femelles a 12 pp., 10 males et 8 femelles a 10 pp., 8 ind. a 10 pp. de sexe
non reconnu, 8 ind. a 8 pp., 9 ind. a 6 pp., 12 ind. a 5 pp., 10 ind. a 4 pp., 13 ind. a 3 pp.

La sex-ratio est de 0,51 sur 115 individus (39 males et 76 femelles). Toutes les
populations dont l'echantillonage est suffisant renferment des representants des deux sexes, a
l'exception de Boca Raton, Florida (13 femelles).

L'espece peuple aussi les Bermudes (St George's West, CONDE, 1972), Madere (CONDE,
1961, sous le nom de P. lagurus ) et les Canaries (CONDE & NGUYEN DUY - JACQUEMIN,
1993).

Polyxenus lagurus (L.), lignee parthenogenetique

J'ai determine l'espece des Etats suivants : Massachusetts, New Jersey, Michigan, Illinois,
Montana, Washington, Colorado, Arizona. Au total : 1 16 specimens : 55 femelles (32 a 13 pp.
(ad.), 15 a 12 pp., 8 a 10 pp.), 26 ind. a 8 pp., 17 ind. a 6 pp., 10 ind. a 5 pp., 6 ind. a 4 pp., 2
ind. a 3 pp. Tous sont identiques aux specimens europeens de la lignee parthenogenetique, la
disposition et le nombre des sensilles basiconiques du 6eme article antennaire (4 a 6 greles, 5 le
plus souvent. NGUYEN DUY - JACQUEMIN, 1976 : 1 14) permettanl de les distinguer facilement
de P. fasciculatus.

On ne connait pas encore d'aires de contact entre P. lagurus et P. fasciculatus, comme cela
existe au Nord de l'Europe occidentale entre les lignees parthenogenetique et bisexuee de P.
lagurus. Les stations les plus proches des deux especes sont situees dans l'lllinois, a quelque
600 km de distance. La limite meridionale de P. lagurus coincide, de fa?on assez satisfaisante,
avec celle du climat continental defini par des amplitudes de temperature superieures a 20°C. II
n'est pas possible de decider actuellement si la presence de P. lagurus en Amerique du Nord est
la consequence d'une repartition holarctique ancienne ou si elle est due a une intervention de
l'Homme.

Polyxenus pugetensis Kincaid, 1898 (Fig. 2)

Selon son auteur, cette espece serait beaucoup plus proche de P. lagurus que de P.
fasciculatus. Le nom fait reference a la localite de Puget (ou au Puget Sund), a l’ouest de Seattle,
tandis qu'une indication plus vague figure dans la description originale : “Hab. : Westerns
Washington”. L'auteur precise qu'il n'a observe que des femelles, ce qui suggere une possible
confusion avec la lignee parthenogenetique de P. lagurus qui est presente au moins dans l'Est de
I' Etat (Spokane).

Cotype.

Le Dr. F. RICHARDSON nous a communique une lame qui porte les indications suivantes :

" Polyxenus pugetensis Kincaid, cotype”, “Thomas Burke Memorial-Washington State
Museum, Seattle, Washington. Catalogue n° 20344”,“Sex...-Date 1897-Local. University of
Washington Campus, Seattle, Washington”.

II s'agit d'une femelle a 13 pp. (ad.), montee dans le Baume du Canada. L'epaisseur de la
preparation et l'opacite du specimen rendent l'observation tres difficile, mais j'ai pu distinguer
neanmoins l'apex de 3 sensilles basiconiques seulement sur le 6eme article antennaire.

SPECIMENS COMPLEMENTAIRES. — Communiques par la regrettee Dr. Nell B. CAUSEY
et le Dr. R. L. HOFFMAN.

Washington. Mason Co., 1 mi E Lake Cushman Dam, Olympic Pen., 07.07.1959, L. M.
Smith : 1 male a 13 pp. (ad.). — Oregon. Benton Co. Moss sample (Berlese), 03.1962, L.

DIPLOPODES PENICILLATES DE L’AMERIQUE DU NORD

131

Abrahamsen : 1 femelle a 13 pp. (ad.).- Oak Creek, 6 mi NW Corvallis, Oak/Douglas fir litter,
29.04.1972, L. Russell : 11 males et 1 1 femelles a 13 pp. (ad.), 1 male et 1 femelle a 12 pp., 3
ind. a 8 pp., 4 ind. a 6 pp.

Adultes. Longueurs.- Corps = 3,00-3,60 mm ; pinceaux telsoniens = 0,48 mm (trichomes
en crosses) et 0,65 mm (trichomes barbeles). Ta I : males = 100-1 1 1 pm (x = 104,8 pm,
n = 10) ; femelles = 100-1 17,5 pm (x= 1 10,3 pm, n = 9). Ta XIII : males - 1 10,3-127,8 pm
(x = 1 19,9 pm, n = 10) ; femelles = 125,8-140,2 pm (x = 132,6 pm, n = 10). Ta Xlll/ta I:
males = 1,09-1,27 (x = 1,15 ; n = 9) ; femelles = 1,17-1,28 (x = 1,20 ;n =9).

Tete.- Plages posterieures du vertex
environ deux fois a deux fois et demi plus
longues que leur ecartement, comprenant
chacune 19 a 27 (21-24 le plus souvent)
phaneres sur 2 rangs rapproches ; 1+1
trichomes parasagittaux en arriere.

Les articles antennaires ressemblent a ceux
de P. lagurus ou de P. fasciculatus par leurs
longueurs relatives et leur forme. En revanche,
les sensilles basiconiques du 6eme article sont au
nombre de 3 seulement, le median un peu plus
epais et plus court que les autres ; en outre, un
sensille setiforme a base renflee et un
coeloconique sont presents, comme chez les
deux especes precedentes.

Fig. 2. — Polyxenus pugetensis Kincaid, femelle adulte :
plages pigmentaires de la tete et des trois premiers
tergites. Dessin de Claude Poivre.

Fig. 2. — Polyxenus pugetensis Kincaid, adult female:
pigmentary areas on the head ant the first three
tergites. Drawing by Claude POIVRE.

0.5 mm

Article VII avec 3 sensilles basiconiques greles, suivis de 2 plus epais et d'un
coeloconique (Fig. ID).

6 stemmates subegaux. Trichobothrie an tero- interne, a calice de dimensions reduites, qui a
echappe a KINCAID.

Labre avec 6+6 lamelles marginales imbriquees, les tubercules des 2-3 premiers rangs
beaucoup plus volumineux que les suivants.

Palpes du gnathochilarium portant 15 sensilles le plus souvent (n = 23), rarement 16 ou 17
(n = 2) et 13 ou 14 ( ? phaneres non vus, arraches ou reellement absents) ; 15 sensilles aux
palpes des deux individus a 12 pp., 12 et 9 chez les individus a 8 et 6 pp.

Tronc.- Les plages laterales du collum sont unies par deux rangees posterieures
ininterrompues de trichomes diriges vers l'arriere ; une courte rangee anterieure et une rangee
intermediaire sont interrompues en leur milieu.

Aux tergites suivants, les phaneres des deux rangees marginales, mais surtout ceux de la
rangee posterieure (diriges vers l’arriere) sont inseres suivant une ligne brisee, ce qui provoque
un dedoublement plus ou moins regulier des rangees et l'impression d’une troisieme rangee
intermediaire.

Telson.- Trichomes des plages subtriangulaires medio-dorsales : males 14+14 - 16+16 (15
le plus souvent) ; femelles 16+16 - 21+22 (16 le plus souvent).

132

BRUNO CONDE

Polyxenus anacapensis Pierce, 1940 (Fig. 3)

Decrite de Middle Anacapa Island (California), j'ai pu examiner des paratypes de cette
espece, communiques par le Dr. Charles L. HOGUE, et etablir qu'elle se distingue des autres
Polyxenus nord-americains par la presence de 5 stemmates (vs 6) et la disposition des sensilles
du 6cme article antennaire (Fig. 3B). L'espece est bisexuee : 2 males et trois femelles a 13 pp.
(ad.), remontes dans le medium II de Marc Andre, ont permis une etude detaillee.

100 pm 25 pm

Fig. 3. — Polyxenus anacapensis Pierce. A : portion droite de la capsule cephalique et antenne de la femelle adulte
paratype n°45 de Middle Anacapa Island. B : articles VI et VII de I'antenne droite d'un paratype adulte (sexe non
identifiable) de la preparation n°20, face tergale. Chiffres et lettres comme sur la Fig. 1.

FlG. 3. Polyxenus anacapensis Pierce. A: right pari of the head capsule and antenna of the adult female paratype n°45
from Middle Anacapa Island. B: right antennae VI & VII articles of an adult paratype (unidentifiable sex) of slide
n° 20, tergal side. Symbols as in FlG. I.

Source : MNHN , Paris

DIPLOPODES PENICILLATES DE L’AMERIQUE DU NORD

133

La presence de 5 stemmates (Fig. 3A) est partagee avec 3 especes : une du Japon (ISHII,
1983), une de Coree (ISHII & CHOI, 1988) et une autre de Chine (ISHII & LIANG, 1990) ;
neanmoins, certains details (4 a 6 sensilles basiconiques dont un epais au 6eme article de
l'antenne, palpes, mandibule) rapprochent davantage anacapensis du complexe de lagurus. Une
nouvelle description sera presentee ailleurs, mais dans cette attente le Tableau 1 resume les
principaux caracteres des quatre especes.

Tableau 1. — Principaux caract£res de 4 especes de Penicillaia.
Table I. — Main features of 4 species of Penicillata.

anacapensis

shinoharai

ko reanus

hangzhoensis

Longueur du corps (mm)

2,42 - 3,01

1,80 - 2

2,35 - 2,66

1,94

Tarse 2, XIII (pm) male

116,5 - 120

81,20

100

femelle

119 - 122,5

88,75

Basiconiques ant. VI

1+ 3-5

2 + 7-9

2 + 5-9

2 + 6

ant.VII

2 + 2-3

2 + 4

2 + 3

2 + 4

Lamelles du labre

7 + 7

5 + 5

Md. elements denticul£s

ca 15

—

1 1

Palpe : mamelon

exp. laterale

11 - 12

Glandes subcoxales

VIII - IX

VII-VIII-IX

VIII -IX

VIII - IX

Trichomes telson male

24-26

19-21

Trichomes telson femelle

21 - 26

20 - 24

—

Macroxenodes bartschi (Chamberlin, 1922), sub Polyxenus

Une nouvelle description d’apres des topotypes (3 males et 1 femelle a 13 pp. ad.) a ete
proposee et un neotype (male a 13 pp.) a ete designe (NGUYEN DUY - JACQUEMIN & CONDE,
1984) et depose au Laboratoire de Zoologie, Arthropodes, du Museum national d’Histoire
naturelle de Paris.

La localite type de l'unique specimen decrit par CHAMBERLIN “Tortugas, Florida” est tres
imprecise. De deux petites series de topotypes presumes, l'une, de Loggerhead Key, Dry
Tortugas, etait constitute de 6 Lophoproctidae (cf. infra) qui ne pouvaient correspondre a
l'espece recherchte ; l'autre, de Upper Snipe Keys, Lower Keys, comprenait 4 specimens d'un
Polyxenidae, le plus vraisemblablement identique a l'espece de CHAMBERLIN, que nous avons
attribues au genre Macroxenodes. A ce genre appartient aussi Polyxenus pcecilus Chamberlin
1923, dont nous avons propose une nouvelle description d'apres l’holotype de South Santa Inez
Island, dans le Golfe de Californie. Le seul critere differentiel incontestable est le nombre et la
disposition des sensilles basiconiques du Vie article antennaire (CONDE & NGUYEN DUY -
JACQUEMIN, 1987).

Lophoturus madecassus (Marquet et Conde, 1950), sub Alloproctus

Seule espece de Penicillata ne possedant que 1 1 paires de pattes au dernier stade, elle
presente une vaste repartition circumtropicale (Madagascar. Sahara, Cote d'Ivoire, Jamaique,
Floride, Pacifique central). Les specimens de Loggerhead Key avaient ete presumes etre des
topotypes de Polyxenus bartschi (NGUYEN DUY - JACQUEMIN & CONDE, 1984 : 722). La sex-
ratio s'etablit a 1/34. le seul male ayant ete recolte sur Nomukaiki (Archipel des Tonga).

Lophoturus cf . aequatus (Loomis, 1936), sub Lophoproctus

Les formes attributes au complexe aequatus - niveus ont en commun un labre a marge
anterieure entiere, sans languettes ou formations analogues, l'omementation de la surface exteme
rappelant un pavage irregulier sans epines cuticulaires ( niveus ) ou avec une seule rangee le long
du bord posterieur {aequatus). Les types de aequatus sont de Haiti, Petite Riviere de Artibonite ;

134

BRUNO CONDE

l'holotype (male a 13 pp.) et un paratype (male a 12 pp.) ont ete revus et compares a ceux de L.
niveus (Loomis, 1936), de Beata Island (CONDE & TERVER, 1965).

Florida. — Key Largo, John Pennekamp St. Pk., 21.10.84, M.A.Deyrup : 1 male et 1
femelle a 13 pp. (ad.), 2 femelles a 12 pp.

Le labre est conforme a celui des types de aequatus. Le
Vie article antennaire (Fig. 4A) est plus allonge (L/l = 1,90-
2 vs 1,63) et les sensilles sont de longueurs un peu inegales,
le posterieur (26,5) entre l'intermediaire (34) et l'anterieur
(23,5). Palpes avec 18-20 sensilles chez les femelles et
environ 40 chez les males. Le rapport 2eme tarse/griffe en
XII est beaucoup plus eleve (1 1,5 vs 8,3) et surtout la griffe
presente un denticule sternal tres net a toutes les pattes,
conime chez niveus (Fig. 4B).

La taille est un peu plus faible que celle du paratype de
aequatus (2eme tarse XII = 126, 128 vs 147 (im) et la pilosite
tres legerement moins fournie.

Fig. 4. — Lophoturus cf. aequatus Loomis, de Key Largo, Florida. A ::
article VI de 1'antenne gauche d'un male adulte. face tergale. B : tarse
XII gauche d'une femelle & 12 pp. a. i, p = sensilles basiconiques
anterieur, intermediate, posterieur ; c = sensille coeloconique.

Fig. 4. — Lophoturus cf. aequatus Loomis, from Key Largo, Florida. A: left
antennal article VI of an adult male, tergal side. B: left tarsus XII of a
12 leg-paired female, a, i, p = anterior, intermediate, posterior
basiconic sensilla; c = coeloconic sensillum.

Le Tableau 2 ci-dessous regroupe quelques valeurs comparatives.

Tableau 2. — Caracteres morphologiques compares de trois especes de Lophoturus.
Table 2. — Compared morphological features in three Lophoturus species.

Ant. VI, L/l

Ant. VI, sens.

ta XU

ta XIII

Dent griffe

L. niveus

indgaux

220 urn

252 um

L. cf. aequatus

1,90-2

inegaux

126 - 128 urn

135 - 146 Jim

L. aequatus

1,63

subegaux

147 Jim

174 Jim

CONDE & TERVER (1979 : 143) ont cite de Cuba (Jatibonico) des specimens proches des
types de aequatus (griffes sans dent, tarse XII : 146 pm). Des Petites Antilles et des Bahamas
(Saint-Eustache, New Providence), CONDE &TERVER (1965 : 134) ont pu etudier des
specimens pourvus d'une dent plus ou moins marquee aux griffes. Ceux de Saint-Eustache (2
femelles a 13 pp., 1 femelle a 12 pp.), assez grands (ta XIII : 202, 208 pm ; ta XII : 170 pm),
avec 22-23 sensilles sur les palpes. Ceux de New Providence (5 males, 7 femelles a 13 pp.)
sont, comme les types de niveus, les plus grands du complexe (ta XIII : 178-240 pm males,
222-256 pm femelles) avec 22-29 sensilles sur les palpes des femelles et 41-56 sur ceux des
males.

Les specimens de Floride montrent une combinaison de caracteres attribues les uns a L.
aequatus (labre, faibles dimensions), les autres a L. niveus (griffes, allongement du tarse), avec
aussi des elements intermediaires (Vie article de 1'antenne). En rapprochant ces specimens de L.
aequatus, plutot que de L. niveus, nous privilegions le critere du labre en considerant que
l'absence totale d'epines cuticulaires chez niveus est un caractere derive par rapport a la presence
de plusieurs rangees ou d'une seule, comme chez aequatus.

Source :

DIPLOPODES PENICILLATES DE L’ AMERIQUE DU NORD

135

REFERENCES

CONDE, B., 1961. — Diplopodes P6nicillates des Azores el de Madcrc. Bol. municip. Funchal, 14 : 7-10.

Cond£, B., 1972. — Presence aux Bermudes de Diplopodes Penicillates el d'Arachnides Palpigrades. Revue Ecol. Biol .
Sol, 9: 127-129.

Cond£, B. & Nguyen Duy - Jacquemin, M., 1987. — Le siaiui de Polyxenus ceylonicus Pocock el de Polyxenus poecilus
Chamberlin (Diplopodes Penicillaies). Revue Ecol. Biol. Sol, 24 : 99-107.

COND£, B. & Nguyen Duy - Jacquemin, M., 1993. — Parihenogenese el reproduclion bisexuee dans le complexe de
Polyxenus lagurus (L.). Biogeographica, 70 : 41-48.

Conde, B. & TERVER, D., 1965. — Les Penicillaies de Haiti decrits par H. F. Loomis. Studies on the Fauna of Curaqao and
other Caribbean Islands, 22 : 124-134.

Cond£, B. & Terver, D., 1979. — Missions Museum Antilles, Diplopodes Penicillates. Revue Ecol. Biol. Sol, 16 :
137-149.

Ishii, K.. 1983. — A new Species of Penicillata Diplopods of the Family Polyxenidae (Diplopoda : Penicillaia) from
Japan. Can. Em., 115 : 1355-1357.

Ishii, K. & Choi, S. S., 1988. — A new Species of ihe Genus Polyxenus (Diplopoda : Penicillata : Polyxenidae) from
Korea. Can. Em., 120 : 711-715.

Ishii, K. & Liang L., 1990. — Two new Species of Penicillate Diplopods of ihe Family Polyxenidae (Diplopoda :
Penicillaia) from China. Can. Em., 122 : 1239-1246.

NGUYEN Duy - JACQUEMIN, M., 1976. — Elude de la variability des caracleres de deux especes du genre Polyxenus, P.
lagurus (L.) el P. fasciculatus Say (Diplopode, Penicillale). Bull. Mus. natn. Hist, nat., Zool. 249, Seme Ser. 356 :
105-118.

Nguyen Duy - Jacquemin, M. & Cond£, B.,1984. — Nouvelle description el statui de Polyxenus bartschi Chamberlin
(Diplopodes Penicillaies). Bull. Mus. natn. Hist, nat., 4eme Ser., sec. A, 6 : 721-728.

Pierce, W. D., 1940. — A rare Myriapod from Anacapa Island compared with two Texas Species. Bull. South. Calif.
Acad . Sci.. 39 : 158-171.

Source : MNHN , Pans

About the Taxomomy of Spanish Scolopendrellidae

Maria Teresa DOMINGUEZ RODRIGUEZ

Centro de Ensenanza Superior San Pablo C.E.U. Carrctera Boadilla del Monte. Km. 5.300
E-28660 Boadilla del Monte, Madrid, Espagne

ABSTRACT

According to the usual morphological characters. Spanish specimens of the family Scolopendrellidae. genus
Scolopendrellopsis, subgenus Symphylellopsis . has been analysed. Those show some mixed characters; for example, it
has been found species that show the character "Tomosvary organs with long prolongations” that identified with
Scolopendrellopsis (Symphylellopsis) pauli n. sp.. joined to the character "12 tergal setae on the tergites without
posterior prolongations" that belongs to the species Scolopendrellopsis (Symphylellopsis) selgae Dominguez. This
suggests that the importance of the last character should be decreased.

It has been analysed and discussed the other more important characters that are usually employed in this group of
symphylids.

RESUME

A propos de la taxinomie des Scolopendrellidae d'Espagne (Symphyla).

Des specimens espagnols de la famille Scolopendrellidae. du genre Scolopendrellopsis, et du sous-genre
Symphylellopsis , ont ete etudies en accord avec les criteres morphologiques usuels. Ils montrent la presence de
caracteres hybrides : par exemple. on a rencontre des specimens presentant le caractere "Organes de Tomosvary pourvus
de longs prolongements", ce qui caracterise Scolopendrellopsis (Symphylellopsis) pauli n. sp.. associc au caractere “12
soies tergales sur les tergites depourvus de prolongations postcrieures" qui appartient a Pespece Scolopendrellopsis
(Symphylellopsis) selgae Dominguez. Ceci suggere que I* importance accordee & ce dernier critere doit etre reduite.
D’autres caracteres habituellement employes dans ce groupe de symphyles ont ete etudies et sont discutes.

Dominguez - Rodriguez, M. T.. 1996. — About the taxomomy of Spanish Scolopendrellidae. In: Geoffroy, J-
J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M.. (cds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat 169 :
137. Paris ISBN : 2-85653-502-X.

Source : MNHN, Paris

Some Observations on the Onychophoran Fauna of

Tasmania

Hilke RUHBERG * & Robert MESIBOV **

* Zoologischcs Institut und Zoologisches Museum der Universitat Hamburg
Martin-Lulher-King-Platz, 3 D- 20146 Hamburg, Germany
** P.O. BOX 700, Burnie, Tasmania, Australia 7320

ABSTRACT

At least nine species of Peripatopsidae (Onychophora) are native.io Tasmania. The four currently recognized
viviparous species, all endemic, have 15 pairs of legs and are restricted to northeastern or southwestern parts of the main
island: Tasmanipaius anophthalmus and T. barretti are only found in the North East, and two species of a new genus (as
yet undescribed) mainly occur in the South West's World Heritage Area (WHA). All other known species are oviparous,
have 14 pairs ot legs and were previously identified as Ooperipaiellus ins ignis, found in Victoria on the Australian
mainland. Egg-laying Tasmanian Onychophora are widely distributed and sometimes locally abundant. Taxonomic-
characters for oviparous species arc here reviewed and it is suggested that “0. insignis" in Tasmania is in fact a group of
endemic species.

RESUME

Observations sur la faune des onychophores de Tasmanie.

Neuf especes au moins de la famille Peripatopsidae (Onychophora) sont originaires de Tasmanie. Les quatre especes
vivipares connues, toutes endemiques, sont munies de 15 paires de pattes ; elles montrent une distribution restreinte :
Tasmanipaius anophthalmus et T. barretti oni ete exclusivement trouvees dans le Nord-Oucst et deux especes d’un nouveau
genre (inedit) existent surtout au South-West’s World Heritage Area (WHA). Toutes les autres especes connues sont
ovi pares et posscdent 14 paires de pattes. On les a regroupees jusqu’& present sous 1’espfece Ooperipatellus insignis,
trouvee a Victoria sur 1c continent australien. Les Onychophores ovipares de la Tasmanie presentent une large repartition
et abondent parfois en certains endroits. Ce travail propose une revision des caracteres taxinomiques et suggere que
“0. insignis ” represente en fait un groupe d'especes endemiques de Tasmanie.

INTRODUCTION

Onychophora frequently appear in phylogenetic discussions of the Arthropod
relationships, and in zoogeographic discussions of the Gondwanan element in fauna of the
Southern Hemisphere. Despite the scientific importance of the group onychophoran taxonomy,
especially at the species level, is far from satisfactory (RUHBERG, 1992). This is particularly true
for the Onychophora of Tasmania. Although the first record of a Tasmanian species was
published 100 years ago (SPENCER, 1895), very little collecting and no taxonomic studies were
carried out over the following 80 years. A “boom” in Tasmanian onychophoran research began

RUHBERG, H. & Mesibov, R., 1996. — Some observations on the Onychophoran fauna of Tasmania. In:
Geoffroy, J.-J., MAURifes, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist,
not., 169 : 139-150. Paris ISBN : 2-85653-502-X.

140

HILKE RUHBERG & ROBERT MESIBOV

with a visit to the island by Dr. V. van der LANDE in 1977. The present authors and their
colleagues have been studying the Tasmanian fauna since the mid- 1980’s, and considerable
progress has been made towards a comprehensive monograph (RUHBERG & MESIBOV, in
prep.).

TASMANIA - A SPECIAL ISLAND

Global climatic changes and other events have resulted in Tasmania being separated from
the Australian continent on several occasions during the past 2.5 million years. Australia itself is
an isolated remnant of Gondwana and thus is rich in ancient groups of its flora and fauna with
high percentage of endemisms (SMITH et al ., 1993). Tasmania experienced several highland
glaciations during the Pleistocene (DARLINGTON. 1969). The present isolation as an island is
believed to have stabilized some 6000 years ago. Cooling factors peculiar to this island together
with isolation periods resulted in the evolution of numerous taxa of plants and animals which are
now endemic to Tasmania, and has made this island an important repository and refuge for
archaic elements of great biological interest and significance. Amongst these are the
Onychophora. commonly referred to as “ Peripatus ”, "Velvet Worms” or "Living Fossils”.

HISTORY OF ONYCHOPHORAN RESEARCH IN TASMANIA

A "rather bleached specimen with fifteen pairs of legs” in the Macleay Museum in Sydney
was first noted by FLETCHER (1890) as demonstrating “the occurrence of ‘ Peripatus Leuckarti '
in Tasmania”. Unfortunately this specimen no longer exists in the museum's collections (D.S.
HORNING, pers. comm.. 1994), and nothing more is known of its morphology or provenance.

Three years later. Sir Baldwin SPENCER collected some 15 specimens of what he called
" Peripatus insignis ” at Dee Bridge in south central Tasmania (SPENCER, 1895). Peripatus
insignis was the name which had been given by DENDY (1890) to the second only known
oviparous onychophoran described from Macedon. Victoria on the Australian mainland. Both
the Victorian and Tasmanian specimens had 14 pairs of legs. However, SPENCER (1895)
mentioned differences in size between the mainland and the Tasmanian form. DENDY (1900)
erected the genus Ooperipatus to contain all oviparous Australian Onychophora, regardless of
whether they had 14 or 15 pairs of legs. In his famous monograph on the oviparous species of
Onychophora, he himself laid the foundation for future taxonomic confusion when he
synonymized the Victorian Ooperipatus insignis from Macedon and the Tasmanian “ insignis ”
from Dee Bridge (DENDY. 1902: 403, 408).

A note by BAEHR (1977) on Australian Onychophora included the description of a new
species with 14 pairs of legs, Ooperipatus decoratus, from Dip Falls in northwestern Tasmania.
The type material, thought until recently to be missing, has now been relocated and re-examined
by the senior author.

Following a collecting trip to western Tasmania in 1977, Dr. V. van der LANDE requested
additional material from Dr. J. HICKMAN of the University of Tasmania, as a form with 15 pairs
of legs among her specimens could obviously not be identified as Ooperipatus insignis. Her
request encouraged local zoologists to deliberately search for Onychophora. In 1983, Leigh
WlNSOR (unpublished report) noted the occurrence of a form with 15 pairs of legs, which he
called “Peripatoides leuckarti ”, near the Franklin River in the South-West (MALCOLM, 1987). A
second form with 15 pairs of legs from northeastern Tasmania, was found by the junior author
in 1984, and was later described as Tasmanipatus barretti together with a third species,
Tasmanipatus anophthalmus by RUHBERG et al. (1991). At the same time, in 1984, the senior
author was completing a revision of the Peripatopsidae of the world, and had available for study
only 18 preserved museum specimens from Tasmania, all of them in rather bad condition.
Accordingly the results were tentative and indicated a need for further work on fresh material.
RUHBERG (1985) retained the genus name Ooperipatus for its generotype Ooperipatus oviparus

THE ONYCHOPHORAN FAUNA OFTASMANIA

141

(a larger oviparous form with 15 pairs of legs from Victoria) and erected a new genus:
Ooperipatellus, to contain all remaining oviparous species with 14 pairs of legs. She considered
the meagre Tasmanian material in hand to be conspecific with two redescribed and renamed
species from the Australian mainland: Euperipatoides leuckarti (SAENGER, 1869), a viviparous
form from New South Wales, with 15 pairs of legs, and Ooperipatellus insignis (DENDY,
1890), from Victoria, with 14 pairs of legs.

An Australia-wide survey of Onychophora was begun in 1985 by Drs. N. N. Tait and D.
A. BRISCOE following their discovery of New South Wales forms with peculiar head organs
(Tait & Briscoe, 1990). In 1987, Tait & Briscoe collected Onychophora throughout
Tasmania, including remote portions of the World Heritage Area in the South-West. These
visits, aimed principally at securing material for allozyme electrophoretic investigations (Tait &
BRISCOE, in Smith et al. , 1993), stimulated the junior author to begin intensive field studies of
onychophoran conservation (e.g. MESIBOV, 1988, 1990. 1994). The senior author made the
first of three visits to Tasmania in 1989, and Dr. D. ROWELL has included recently collected
Tasmanian forms in his studies of chromosomal variation and chromosomal evolution within a
sample of Australian Peripatopsidae (ROWELL, unpubl. obs.; ROWELL et al., 1995).

As a result of all these recent activities there is now a rich supply of material available for
further taxonomic work on the Peripatopsidae of Tasmania enabling the earlier studies to be
reassessed. The first questions to be answered are:

(1) Are the Tasmanian species referred to Euperipatoides leuckarti and Ooperipatellus
insignis conspecific with their mainland counterparts?

and:

(2) Are there more species in Tasmania than at present described?

In what follows, we attempt to answer these questions, concentrating on the taxonomic
complexities of the oviparous forms. We begin, however, with a brief review of the viviparous
species.

VIVIPAROUS TASMANIAN ONYCHOPHORA

All known viviparous1 forms from Tasmania have 15 pairs of legs in both sexes. Their
distributions are remarkably restricted, with one group found only in the North-East and the
other in the South-West (FIG. la).

One of the northeastern species, Tasmanipatus anophthalmus Ruhberg et al.. 1991, is
white and “blind" (RUHBERG et al.. 1991; MESIBOV & RUHBERG, 1991). Its congener,
T. barretti Ruhberg et al.. 1991, has obvious eyes and is dorsally pinkish pigmented. The two
species occur parapatrically in forest habitats over ca. 1,000 sq. km. (RUHBERG et. al. 1991;
MESIBOV & Ruhberg. 1991). Histological investigations have revealed that an “inner” eye
occurs in both species, and that T. anophthalmus lacks the lens and the retina-pigment found in
this structure in T. harretti (RUHBERG et al, in prep.).

Viviparous forms from southwestern Tasmania were previously referred to Euperipatoides
leuckarti (Saenger. 1869), (RUHBERG, 1985). It now seems clear that the southwestern
viviparous Onychophora represent two allopatrically distributed species (FlG. la) in a new
genus, to be described in a forthcoming paper (RUHBERG, in prep.).

In summary, there is now strong evidence from morphological and phylogenetic studies
(RUHBERG in prep.; REID, 1995, in prep.), chromosome studies (ROWELL et al. 1995) and
allozyme investigations (Tait & BRISCOE in: SMITH et al. 1993), that none of the Tasmanian

1 All viviparous Peripalopsidae from Australia are new considered to be ovoviviparous (Campiglia & Walker, 1995;
Reid, pers. comm.).

142

HILKE RUHBERG & ROBERT MESIBOV

viviparous species are conspecific, or even congeneric, with mainland Australian forms. All
show clear-cut diagnostic features and are further characterized by their geographically restricted
distributions.

Fig. 1 a-b. — Distribution of (a) viviparous and of (b) oviparous Peripatopsidae in Tasmania.

OVIPAROUS TASMANIAN ONYCHOPHORA

Oviparous forms were previously identified with the Victorian Ooperipatellus insign is
(DENDY, 1890), (RUHBERG, 1985). Egg-layers are found throughout the main island of
Tasmania and on several offshore islands (Fig. lb). They occur in forest, woodland and scrub
habitats from sea level to at least 1,100 m and are sometimes locally abundant. Occurrences in
forest after clearfell logging, part-clearing or burning demonstrate that oviparous species, at least
in the short term, are remarkably tolerant of habitat disturbance (MESIBOV, unpubl. results).

In contrast to viviparous species in Tasmania, oviparous forms are superficially very
similar. All females have a prominent ovipositor and lay shelled eggs (Figs 4e, 3d), all males
have a nearly uniform distribution-pattern of crural papillae on leg-pairs 6-13, and both sexes
have 14 pairs of walking appendages. These characters are shared by Victorian and New
Zealand oviparous species within the genus Ooperipatellus which are also alike in having a 2n
chromosome number of 42. In contrast there is much variation within the viviparous forms.
Tasmanipatus- spp. have 2n = 34 or 36 and the yet undescribed viviparous southwestern genus
has 2n = 18, with interspecific variation in sex chromosomes (ROWELL et al., 1995).

Early taxonomic studies relied to a large extent upon colour and colour-pattern, as can be
seen in DENDY's impressive opus “On the oviparous species of Onychophora” (DENDY, 1902;

THEONYCHOPHORAN FAUNA OFTASMANIA

143

pi. 19, Figs 1-3). We have found that colour and colour-pattern per se are unreliable characters
in onychophoran systematics, and that other traits need to be examined (RUHBERG, 1992).

In an effort to improve the taxonomy of oviparous onychophorans, and to arrive at a well
grounded biospecies-concept, the present authors, in collaboration with specialists, are using
data on external and internal anatomy, histology and ultrastructure, allozyme electrophoresis,
behaviour and distribution patterns. Specimens for these investigations derive almost entirely
from our own field collections (mainly deposited in the Queen Victoria Museum and Art Gallery.
Launceston, Tasmania ), and include animals bred in the laboratory by H. R. in Hamburg. The
results of particular studies will be published in forthcoming papers. Here we review progress in
identifying species-diagnostic characters.

DIAGNOSTIC CHARACTER VARIATIONS IN OVIPAROUS ONYCHOPHORA

For the sake of completeness we begin with a revision of the genus Ooperipatellus.

Ooperipatellus s. str. Ruhberg, 1985.

Type species: Peripatus insignis Dendy, 1890 from Macedon, Victoria (to be redescribed
in REID, in prep.).

Distribution

Victoria, Tasmania, New Zealand.

Diagnosis

Ooperipatellus is a genus of Australasian oviparous peripatopsid Onychophora with 14 pairs
of legs. Females have a prominent ovipositor and lay shelled eggs, males have crural papillae on
leg pairs 6-13. Outer jaw blade without accessory tooth.

Differential diagnosis

Ooperipatellus, s. str., is distinguished from all other known oviparous peripatopsid
genera on the basis of a unique combination of characters (for comparison see RUHBERG, 1985).
It is distinguished from Ooperipatus, the “larger Victorian genus” which has 15 pairs of legs,
male crural papillae on leg pairs 2-14, and an accessory tooth on the outer jaw blade.

Ooperipatellus is separable from most other currently recognized, but as yet undescribed
oviparous mainland Australian species (REID, in prep.), in lacking characteristic head organs.
Further the latter forms show different patterns of male crural papillae.

Ooperipatellus nanus Ruhberg, 1985, a tiny form from southern New Zealand, with only
13 pairs of legs, which was previously tentatively assigned to this genus (RUHBERG, 1985: 131)
shows more unique characters in adults now than could be deduced from juveniles at hand in
1985. This species has to be transferred to a new genus (RUHBERG, in prep.).

Description

Oviparous peripatopsids. Leg number constant within species, last leg pair well
developed, with claws. Foot with 3 distal papillae, no basal papillae. Anal cone of variable
length (Fig. 2f, 4a-c). Genital pore in males of variable shape (Figs 4a-c), females with distinct
ovipositor of variable length (Fig. 4e).

Males with accessory glands coiled around each other.

Females with paired, flat ovaries, closely attached to the pericardial septum. Ovarial eggs
exogeneous and highly variable in size; rudiments of receptacula seminis only present in juvenile
females, lost in adults. Additional pouches lacking. Uterine eggs of varying developmental
stages.

144

IIILKE Rl'HBERG & ROBER T MES1BOV

Fig. 2 a-f. — SEMs of Tasmanian Onychophora: (a) Fourth foot with three distal papillae (dpp). nephropore (np) and
spinous pads (sp); (b) Dorsal skin with primary (arrowhead) and secondary dermal papillae (a-b. e: Oop&ipatellus
sp. from Black River); (c) Typical pattern of dermal papillae and plical folds in O. decorcitus ; (d) Yet undescribed
structure on dorsolateral skin in OoperipateUus sp. from Christmas Hill; (e) Antennal tip: second and third
annulus, each with one row of mechanoreceptors only (arrowhead); (f) Tasmanipatus barretti : Hind end of body
with a pronounced anal cone.

Source : MNHN, Paris

THEONYCHOPHORAN FAUNA OFTASMANIA

145

Fig. 3 a-f. — SEMs: (a-b) Different size and shape of male crural papillae in Ooperipatellus sp. from Black River: (a) I ! th
(left on FIG.) and 12th leg (right on FIG.); (b) 7th leg; (c) Crural papilla on 12th leg in O. decoratus\ (d-f) Chorion
of ripe eggs; (d) Egg from a yet undescribed oviparous species from Bellendcn Ker. N- Queensland; (e) Sculpture o!
egg-chorion in O. deco rat us ; (D egg-chorion in Ooperipatus oviparus from Victoria.

Source : MNHN , Paris

146

HILKE RUHBERG & ROBERT MESIBOV

Fig. 4 a-f. — SEMs: (a-c) Different shape and size of genital pores (arrowheads), crural papillae of last legs (asterisks)
and anal-slits (arrows) within male oviparous species; (a) Body's hind end in a sexual active O. decorciius ,
collected by R. M. during a “swarming night” in October 1992; (b) Ooperipcitellus sp. from Black River,
Tasmania; (c) Ooperipatus oviparus from Victoria; (d) Cruciform male genital pore in the viviparous
Tasmanipatus barretti ; (e) Ovipositor of O. decorums ; (f) Distinct head organ in a yet undescribed viviparous
species from the Tindcrry Mts., NSW (lateral view).

Source : MNHN , Paris

THEONYCHOPHORAN FAUNA OFTASMANIA

147

Species-diagnostic characters so far recognized in Ooperipatellus, are noted below:

External Anatomy; Internal Anatomy; Histology and Ultrastructure; Allozyme
Electrophoresis; Behaviour; Distribution; Other Characters.

a) External Anatomy

The number, size, shape and pigmentation of spinous pads of the foot; the position of the
nephropore on the 4th and 5th pair of legs; the number, size and arrangement of distal and basal
foot papillae (Fig. 2a); the structure and distribution of dorsal skin papillae (Figs 2b-c), the
number of dorsal plical folds (Fig. 2c), the presence or absence of as yet undescribed structures
on the latero-dorsal surface (Fig. 2d); the size of the primary dorsal papillae (Figs 2b-c), which
varies to give an overall “smooth” or “warty” appearance to the body; the number of rows of
mechanoreceptors on the second and third distal annulus of the antenna (Fig. 2e); the presence or
absence of a deep wrinkle at the antennal base (TAIT & BRISCOE, 1987); the size, shape and
position of the male crural papillae on leg-pairs 6-13 (Figs 3a-c, 4a-c), the degree of elongation
of the anal cone (Fig. 2f), the degree of reduction of the last legs; and the overall size, e.g. the
“stoutness” or the “slenderness” of the body (cf. BOUVIER, 1905; pi. I, Figs 4 & 6).

Coloration characters must be used with care. Even when very distinctive dorsal patterns
appear in a population (e.g. a “striped”, “chequered”, “diamond”, “chessboard”, “spotted”, or
“speckled” pattern) there can be substantial within-population variation (BROCKMANN, 1994;
Figs 7-8). Eye coloration is an unreliable character, it seems to change in fixative. Nevertheless,
antennae appear to have species-characteristic patterns of annular coloration which are already
visible in late embryos (BROCKMANN, 1994). The occurrence or absence of pigment and/or
colour patterns of the ventral body surface is characteristic as well and so is the pigmentation of
the hatchling in oviparous respectively of the newborn in viviparous species (RUHBERG, pers.
observations).

b) Internal Anatomy

Among the anatomical characters are mainly the peculiarities of the genital tracts in both
sexes. In Ooperipatellus, species can be distinguished by the following characters: the position
and shape of male crural glands (within the leg or free in the body cavity) and anal glands (e.g.
accessory glands coiled or uncoiled). The structure and development of the ovarial and uterine
eggs in the females vary in number and age.

c) Histology and Ultrastructure (SEM, TEM)

The sculpture of the chorion of the ripe uterine or freshly deposited egg (DENDY, 1902;
pi. 21; Figs 20-27, and this report. Figs 3d-f) is highly characteristic. Of further taxonomic
value are SEM-details of the integument, including the size, shape and number of scales on the
main dorsal papillae (Fig. 2b); features of the head, feet and genital region, and as yet
undescribed structures in certain species (Fig. 2d) which are currently under investigation with
regard to their function and taxonomic value (RUHBERG, in prep.).

First results from histological studies of O. decoratus (males, females and eggs) are
promising (BROCKMANN, 1994), in that this species shows clear histological differences when
compared with O. insignis, as described by DENDY (1902). Unfortunately, freshly killed
material is needed for such studies and histological characters may not be of use in identifying
museum specimens. The same caution applies to use of ultrastructural ditferences noted in
examination of fresh O. decoratus and O. viridimaculatus (DENDY, 1900; RUHBERG &
BROCKMANN, in prep.); among the most valuable TEM-criteria are details of the
spermatophores. In contrast, SEM-investigations are possible with old museum material as well
(RUHBERG, 1985, 1992).

d) Allozyme Electrophoresis

Characteristic allozyme patterns in Tasmanian oviparous forms have been used by TAIT &
BRISCOE (in: SMITH et al., 1993) as the basis for separating taxa in the “(9. insignis ” complex.
These “electrotaxa” are tentative, but the patterns suggest, that oviparous forms can have

148

HILKF. RUHBERG & ROBERT MESIBOV

restricted distributions within Tasmania, and that the Tasmanian forms are only distantly related
to oviparous Onychophora on the Australian mainland.

e) Behaviour

Ooperipatellus viridimaculatus from Shennandoah Saddle. New Zealand. South Island,
lies still when picked up and is remarkably sluggish in culture (RUHBERG, pers. obs.), while a
new Ooperipatellus -species from northwest Tasmania (RUHBERG & MESIBOV, in prep.) very
rapidly coils itself into a tight helix when disturbed. O. viridimaculatus is also unusual in
carryin° its antennae mostly bent backwards. A curious “swarming" of O. decoratus was noted
one night in October 1992, when hundreds of individuals were seen climbing trees about an
hour after sunset at a field site in northwest Tasmania (MESIBOV, unpubl. obs.). A random
sample from the swarm proved to be 90% males, with individuals showing widely opened
genital pores (FIG. 4a), and drops and threads of secretion clinging to crural gland openings
rRUHBERG. unpubl. obs.). It is not yet known whether any other Ooperipatellus species exhibit
swarming.

f) Distribution

Onychophoran populations are generally rather small and their distribution is disjunct on
both large and small scales. In several places egg-layers occur sympatrically with live-bearers.
Within Tasmanian Onychophora all possible distributional patterns do occur: broad sympatry,
naiTOw sympatry. parapatry and allopatry (MESIBOV. unpubl. obs.). A few species seem to be
better dispersers than others (e.g. T. barretti compared to T. anophthalmus ). Onychophora in
the State have been collected in almost all forest types: dry. wet and alpine. Although oviparous
Onychophora are almost ubiquitous in Tasmania (Fig. lb), it is clear from our preliminary
taxonomic work that individual species can be restricted to relatively small areas. It seems
unlikely that differing habitat preferences account for range limitation, and in all cases the
microclimate parameters are similar in the prefered shelters: under logs and stones, in leal litter
and soil crevices. Distribution limits may be “historically” determined or may be controlled by
interactions with other Onychophora, as is suspected to be the case for the parapatric
Tasmanipatus species of northeast Tasmania (MESIBOV & RUHBERG, 1991). Where range
boundaries are sharp, location may be used as a species-diagnostic character.

g) Other Characters

There is potential for using secretions as species-diagnostic biochemical characters.
Recently ELIOTT et al. (1993) have shown that crural gland secretion in males of the viviparous
Cephalofovea tomahmontis Ruhberg et al., 1988, acts as a chemoattractant for conspecific
females. Extensions of this study to oviparous species may reveal a range of biochemically
distinctive, pheromonal attractants. Onychophoran slime may also be taxonomically useful
(RENWRANTZ & RUHBERG, in prep.), although the slimes of the Ooperipatellus- species so far
studied appear to be nearly indistinguishable. Chromosome studies also have so far documented
the close relationship of Ooperipatellus- species from Tasmania. Victoria and New Zealand
(ROWELL, pers. comm.). Onychophora have the right properties (small, isolated populations)
for the study of chromosome driven speciation. As has been outlined before there is much
variation in the chromosome numbers within the Australian viviparous forms while all oviparous
representatives of Ooperipatellus investigated are alike in having a 2n = 42 chromosome-
number-pattern (ROWELL et al., 1995, in press). The latter is also the largest chromosome
number observed to date in Onychophora.

DISCUSSION

Returning to our earlier questions, we are now confident on the basis of morphological
and other studies that (1 ) neither Euperipatoides leuckarti nor Ooperipatellus insignis is present
in Tasmania, and (2) that Tasmania is home to four, not three, viviparous species and to at least
five, not one, oviparous species of Onychophora. The oviparous species have congeneric

Source :

THEONYCHOPHORAN FAUNA OF TASMANIA

149

relatives in Victoria and New Zealand and will be described in forthcoming papers (RUHBERG &
MESIBOV, in prep.).

For identification in the field it is convenient that viviparous Tasmanian species all have 15
pairs of legs, while oviparous Tasmanian species all have 14 pairs of legs. Identifications within
the taxonomically difficult oviparous group will depend on careful examination of many of the
subtle characters noted in this paper, a procedure made more difficult by the absence in the
Tasmanian forms of “complicated characters" sensu HENNIG, such as the head organs (Fig. 4f)
found in some oviparous and viviparous Onychophora on the Australian mainland (RUHBERG et
al. , 1988; Tait& Briscoe, 1990; Reid, in prep.).

Phylogenetic studies of oviparous forms will also be difficult, and a combined
morphological, genetic and ecological approach will be the key to understanding their radiation
within Australia. The effort will be worthwhile, as it will shed light on whether egg-laying or
(ovo-)viviparity is the primitive reproductive mode in the peripatopsid Onychophora. The co¬
existence of both reproductive modes in Tasmania is both a mystery and an opportunity for
understanding the respective advantages of the two strategies.

ACKNOWLEDGEMENTS

The authors are grateful for financial support from the following sources: (to H.R.) the Deutsche
Forschungsgemeinschaft; grants DFG Ru: 358/1-5 and 2-1; (to R.M.) the Plomley Foundation (through the Queen
Victoria Museum and Art Gallery. Launceston), and the Australian Heritage Commission (through the Tasmanian
Department of Lands, Parks and Wildlife). For the loan of museum specimens H.R. is indepted to Dr. T. Kingston,
Curator of Zoology at the QVMAG. Dr. R. Raven. Senior Curator of Chelicerata at the Queensland Museum. South
Brisbane, Dr. P. M. Johns, Canterbury University. Christchurch, N. Z.. Dr. D. Burckhardt, collections for the Museum
d'Histoire naturelle, Geneve, and Mr. P. P. Parillo, Division of Insects , Field Museum of Natural History, Chicago,
Illinois.

For skillful technical assistance at the Scanning Electron Microscope we would like to thank Mrs. R. Walter,
Zoology Dept., University of Hamburg. For unpublished notes, records and additional material both authors thank Drs.
D. Rowell and A. Reid (Canberra). N. N. Tait and D. A. Briscoe (Sydney) and V. v. d. Lande (Nottingham). Dr. M. H.
Walker (Leicester) kindly corrected H. Ruhberg’s original text version.

REFERENCES

Baehr, M., 1977. — Uber einige Onychophoren aus Australien und Tasmanien mil Beschreibung einer neuen Art und
Anmerkungen zur Stcllung von Ooperipcitus paradoxus Bouvier. 1915. Zool. Jb. Syst ., 104 . 9-19.

Bouvier, E. L., 1905. — Monographic des Onychophores. Annls. Sci. nat. (Zool.). 2 : 1-383.

Brockmann, C., 1994. — Zur Eidonomie, Anatomie und Entwicklungsbiologie von Ooperipatellus decoraius (Baehr,
1977) (Pcripatopsidae, Onychophora). Diplomarbeii des Fachbereiches Biologie der Universitat Hamburg
(unverdffentlicht) : 1-102.

Campiglia. S. S. & Walker. M. H., 1995. — Developing embryo and cycling changes in the uterus of Peripatus
(Macroperipatus) acacioi (Onychophora. Peripatidae). 7. Morph.. 224 : 179-198.

Darlington . P. J.. 1969. — Biogeography of the Southern End of the World. Harvard University Press (2nd Ed.) :1-
236.

Dendy. A., 1890. — Preliminary Account of a New Australian Peripatus. Viet. Naturalist . 6 : 173.

Dendy, A., 1900. — Preliminary Note on a proposed New Genus of Onychophora. Zool. Anz., 23 : 510.

Dendy, A., 1902. — On the Oviparous Species of Onychophora. Q. Jl. Micr. Sci., 45 : 363-413.

Eliott, S., Tait, N. N. & Briscoe, D. A.. 1993. — A pheromonal function for the crural glands of the Onychophoran
Cephalofovea tomahmontis (Onychophora: Pcripatopsidae). J. Zool.. Loud.. 231 : 1-9.

FLETCHER, J. J., 1890. — Additional Notes on Peripatus Leuckarti. Proc. Linn. Soc. NSW (2) V (Dec. 16) : 469-486.
Malcolm, H. E.. 1987. — Invertebrate Fauna of the Franklin River Area, Tasmania. Report on the Australian and New
Zealand Scientific Exploration Society (ANZSES) expedition 1983. The Tasmanian Naturalist., 91 : 5-6.

MESIBOV, R., 1988. — Tasmanian Onychophora. Unpttbl. Report for the Dept, of Lands, Parks and Wildlife. Tasmania :
1- 44.

MESIBOV, R., 1990. — Velvet Worms: A Special Case of Invertebrate Fauna Conservation. Tasforests July 1990 : 53-
56.

Mesibov, R.. 1994. — Faunal breaks in Tasmania and their significance for invertebrate conservation. Mem. Qld. Mus.,
36 : 133-136.

150

HILKE RUHBERG & ROBERT MESIBOV

MESlBOV. R. & Ruhberg, H.. 1991. — Ecology and conservation of Tasmanipatus barretti and T. anophthalmus ,
parapatric onychophorans (Onychophora: Peripatopsidae) from northeastern Tasmania. Pap. Proc. Roy. Soc.,
Tasmania . 125 : 11-16.

rEid, A. L.. 1995. — A systematic review of the Peripatopsidae (Onychophora) in Australia. Thesis, North Ride, N. 5.
W., Macquarie University (unpublished).

Rowell. D. M.. Higgins. A. V., Briscoe. D. A. & Tait. N. N., 1995. — The use of chromosomal data in the systematics
of viviparous onychophorans from Australia (Onychophora: Peripatopsidae). Zool. J. Linn. Soc.. 114 : 139-153.
Ruhberg. H.. 1985. — Die Peripatopsidae (Onychophora). Systematik, Okologie, Chorologie und phylogcnetische

Aspekte. Zoologica. 137 : 1-183. . , .

RUHBERG, H., 1992. — “ Peripatus " - an Approach towards a Modern Monograph. Per. nat.-med. Verein Innsbruck.

Suppl. 10 : 441-458.

Ruhberg. H., Tait, N. N.. Briscoe. D. A. & Storch, V.. 1988. — Cephalofovea tomahmontis n. gen., n. sp., an
Australian Peripatopsid (Onychophora) with a Unique Cephalic Pit. Zool. Anz., 221 : 117-133.

Ruhberg, H.. Mesibov. R.. Briscoe. D. A. & Tait. N. N., 1991. — Tasmanipatus barretti gen. nov., sp. nov. and
T. anophthalmus sp. nov.: two new and unusual onychophorans (Onychophora: Peripatopsidae) from northeastern
Tasmania. Pap. Proc. R. Soc. Tasm., 125 .11-16.

SAENGER, H.. 1869. — Peripatus capensis Sr. ct Peripatus Leuckartu n. sp. (cn russe). Trav. 2. Congr. nat. russ. Moscou
: 239-262. (Traduit par Bouvif.r, E. 1. 1901- A propos dun travail de H. Saenger sur les Peripales. Bull. Soc.

philom. Paris 3 : 9-36, pi. : 12-13.

Smith. S. J.. 1993. — Tasmanian Wilderness (Chapter 15) - In : S. J. SMITH & M. R. Banks, Tasmanian Wilderness -
World Heritage Values. Pap. Proc. Roy. Soc. Tasmania. Hobart : 129-143.

SPENCER, W. B.,1895. — Note on the presence of Peripatus insignis in Tasmania. Proc. R. Soc. Viet.. 7 : 31-32.

Tait. N. N. & BRISCOE, D. A.. 1987. — Onychophora in Tasmania. Unpublished Report on WHA Directed Research

Contract : 4 pp + Appendix 5pp. .

Tait., N. N. & Briscoe. D. A.. 1990. — Sexual head structures in the Onychophora: unique modifications for sperm

transfer. J. nat. Hist.. 24 : 1517-1527.

Tait N N & Briscoe. D. A., 1993. — Peripatus or Velvet Worms - a contribution to chapter 13. In : Threatened, rare
and localised plants and animals (by S. J. Smith & L. GlLFEDDER). In : S. J. SMITH & M. R. Banks, Tasmanian
Wilderness - World Heritage Values. Pap. Proc. Roy. Soc. Tasmania. Hobart : 136-138.

Winsor. L., 1983. — Onychophorans (Arthropoda: Onychophora) from the Franklin River Area, South Western
Tasmania. Unpubl. report on a collection made by the ANZSES Expedition. 1983, 4 pp.

Source ; MNHN, Paris

Millipedes as Aids for the Reconstruction of Glacial
Refugia (Myriapoda: Diplopoda)

Jbrg Spelda

University of Hohenheim, Institute for Zoology, Garbenstrasse 30, D-70593 Stuttgart, Germany

ABSTRACT

This paper shows that we can discover Quaternary refugia by studying the present distribution of millipedes. Four
preconditions are proposed that should be performed by a species used for the reconstruction of a refuge. It should have a
low tendency of outspreading, it should be an endemic species, it should be easy to catch and its statements should be
supported by other groups of organisms. Concerning two sites at the northern border of the Alps (in the vicinity of
Basel/Switzerland and Salzburg/Austria) it is shown that some species of chordeumatids fulfill these conditions. The
possibility of speciation after the Ice Age, the character of natural borders, the influence of extinction and men are
discussed.

RESUME

Utilisation dcs diplopodcs dans la reconstitution des refuges glaciaires.

Ce travail montre qu’ il est possible de dccouvrir des refuges quatemaires en Studiant la repartition actuelle des
diplopodes. Quatre conditions pr£alables sont proposees, qu'une espece doit remplir afin de pouvoir etre utile a la
reconstitution d’un refuge glaciaire. Elle doit presenter une faible tendance a la dispersion, etre endemique. etre facile a
capturcr et son statut doit etre appuye par d’autres groupes d'organismes. II est montre que. dans deux sites de la bordure
nord des Alpes (pres de Bale, en Suisse et pres de Salzbourg, en Autriche). certaines especes de chordeumatides reunissent
ces conditions. La possibility d’une speciation post-glaciaire, les caracttSristiques des frontieres naturellcs, I’ influence
des extinctions et celle de 1’homme sont discuses.

INTRODUCTION

The idea of using the present distribution of animals for the reconstruction of glacial
refugia has been born at the beginning of our century and is connected with the name of
HOLDHAUS (1954), who investigated large parts of the eastern Alps. This research caused a
dispute with JANETSCHEK ( 1956), the other one working on this subject. The main contradiction
between them was the existence of inneralpine refugia (as JANETSCHEK stressed) against
secondary immigration into those parts (HOLDHAUS' argumentation), however both principally
agreed in the existence of glacial refugia in the Alps. HOLDHAUS (1954) mainly investigated
beetles (especially wingless ground-beetles and weevils) while JANETSCHEK ( 1956) based his
argumentation on a larger number of taxa. But even earlier the great german myriapodologist
VERHOEFF (1917) recognized the importance of millipedes on this subject. He discovered the
endemic species in the southern Black Forest and during many excursions in the Alps he

Spelda. J., 1996. — Millipedes as aids for the reconstruction of glacial refugia (Myriapoda: Diplopoda). In:
Geoffroy. J.-J., Mauries. J.-P. & Nguyen Duy - Jacquemin. M.f (eds). Acta Myriapodologica. Mem. Mils. natn. Hist,
not.. 169 : 151-161. Paris ISBN : 2-85653-502-X.

152

JORG SPELDA

improved our knowledge on the distribution of these animals. A lot of his publications deal with
the zoogeography of millipedes and he always tried to give explanations of their distribution,
summarized in VERHOEFF (1938a).

MATERIAL AND METHODS

Beside the critical evaluation of the older (e.g. Attems, 1949. Bigler. 1913) and recent literature (c.g. Spelda,
1991; Pedroli-Christf.n. 1993), many collections have been made by the author since 1988 m southwestern Germany
and in the northern Alps.

FAUNIST1C RESULTS

The following records have been unpublished up to now. Making localisation easier for
subsequent workers the degrees of longitude and latitude are given for each locality. The sex of
the specimens is represented by the scheme (males/females). Collectors are only listed i! not
identical with the author.

Abbreviations: ri , c,

Xylvom = Xylophageuma vomrathi Verhoeff. 191 1; Haanor — Haasea norica (Veihoeri,
1913); Hapocu = Haplogona oculodistincta (Verhoelf, 1893); Synace = Syngonopodium aceris
Verhoeff, 1913; Pyrtit = Pyrgocyphosoma titianum (Verhoelf, 1910); Liscer = Listrocheiritium
cervinum Verhoeff, 1925; Lisnor = Listrocheiritium noricum Verhoeff, 1913; Lissep =
Listrocheiritium septentrionale Gulicka, 1965; Rhyale = Rhymogona alemannica (Verhoelf,
1910); Rhycer = Rhymogona cervina (Verhoeff, 1910); Rhyser = Rhymogona serrata (Bigler,
1912); Rhyver = Rhymogona verhoeff i (Bigler, 1913); Rhyweh = Rhymogona wehrana
(Verhoeff, 1910); Rhy = Rhymogona sp.

A = Austria; CH = Switzerland; D = Germany.

CH Aargau, 2 km ESE Sisseln (08°00'E, 47°33'N), 09.10.1991: Rhy (0/2); A: summit
of the mountain GaiBberg (13°06'E, 47°48'N), 17.10.1991: Haanor (2/5); Lisnor (3/4); A:

1 5 km SW Scharfling at lake Mondsee (13°23'E, 47°47'N), 17.10.1991: Hapocu (3/3 , first
record in the northern Alps !); Synace (3/2); Lisnor (4/3); A: highway station “Tauernalm
(13°25'E, 47°15'N), 18.10.1991: Liscer (2/1); A: 3 km SSW Rossatz (15°30'E, 48°22 N),
21.10.1991: Lissep (3/1 , first record for Austria)-, CH: Aargau, 1 km NW Sulz (08°05'E,
47°32'N), 27.10.1991: Rhy (0/6); CH: Aargau, 1.5 km SW Eiken (07°59'E, 47°3TN),
27.10.1991: Rhycer (2/1); D: 1 km N Hasel near Wehr (07°53'E, 47°39'N). 30.10.1991:
Xylvom (4/1); Rhyweh (9/15); Pyrtit (3/1); D: 1.5 km SO Inzlingen (07°42'E, 47°34'N),
30 10 1991: Rhyser (10/8); D: N Maulburg (07°46'E, 47°39'N), 30.10.1991: Rhyale (3/1); D:

1 km S Schallsingen (07°39'E. 47°45’N), 30.10.1991: Rhy (0/6); D: 1 km ESE Wittlingen near
Lorrach (07°40'E, 47°439N), 30.10.1991: Rhy (0/1); D: 1 km NE Neckarhausen (08°39 E,
48°24’N), 01.11.1991: Rhy (0/1); D: 1 km E Marbach near Villingen (08°29'E. 48°01'N),
01 11 199L Rhy (0/3); D: ObergieBhof, 4.5 km S Hornberg (08°13'E. 48°10'N), 01.11.1991:
Rhyver (1/2); D: NiedergieB, 3.5 km SSW Hornberg (08°12'E, 48° 1 l'N), 01.1 1.1991: Xylvom
(1/1)- Rhyver (2/1); D: T km NW Hirschsprung, Hollental near Freiburg (08°01'E, 47°56’N),
01 11 1991: Pyrtit (1/0); 1): 1 km SE Sulzburg (07°43'E, 47°50'N), 18.10.1992: Pyrtit (3/3);
Rhy (0/2); D: E of Lorrach-Brombach (07°42'E, 47°38'N): Rhyser (2/4): D: quarry 1 km E
Gerhausen (09°49'E, 48°23'N), pitfall trap, 17.09.-01.10.1992, leg. J. BOHMER: Rhycer (1/0);
D: Scheibenfelsen SE Hausern (08°10'E, 47°44'N), pitfall trap, 12.1988.-05.1989, leg. R.
MOLENDA: Rhyweh (1/0); D: Prag 6 km SSE Todtnau (07°57'E, 47°46'N), pitfall trap,
10.1991, leg. R. MOLENDA: Pyrtit (5/8): I): S Badenweiler (07°40'E, 47°47'N), 07.10.1986,
leg. A. PEDROLI-CHRISTEN: Pyrtit (0/1); I): S Hierbach (08°05'E, 47°40'N), 08.10.1986, leg.
A. PEDROLI-CHRISTEN: Pyrtit (0/1); D: 1 km SW Altglashiitten (08'05'E, 47°51’N),
30.09.1990, leg. A. PEDROLI-CHRISTEN: Pyrtit (1/0).

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MILLIPEDES AS AIDS FOR THE RECONSTRUCTION OF GLACIAL REFUG1A

153

DISCUSSION AND FURTHER RESULTS

A. Conditions for the use of a species as an aid for the reconstruction of glacial refugia

The use of the present distribution of organisms for the reconstruction of glacial refugia is
based on the assumption that it mainly is a consequence of the depauperation during the Ice Age.
The main argument for this is, that neither petrophilous (meaning restricted to rocky areas) nor
endemic animals occur in Scandinava (HOLDHAUS, 1954; VERHOEFF, 1938a), although we
have similar climatic conditions than in the Alps. As Scandinavia was totally covered with an ice-
shield during the glaciated periods (NORDAL, 1987) we have a very good explanation for this
fact.

To increase the suitability of the used organisms and consequently the validity of their
distribution for our purpose, the following conditions should be performed:

A. 1. Low tendency of spreading out

This is of course very important, because there might have been many species that have
survived at the refugia. But if they largely spread out after the climate had changed to better
conditions their vestiges got lost. Only the few species, that did stay nearby their former refugia,
will show us where they have been situated. In some cases the present distribution indicates a
north-alpine persistance (e.g. in the beetle Trechus glacialis ) but it cannot be proved where this
species has survived and whether it had a monocentric or polycentric refuge. Automatically this
means the more restricted a species is, the belter it is suitable for the reconstruction of glacial
refugia.

A. 2. Speciation

Endemic species will provide a stronger argument for a refuge than isolated populations of
a wider distributed species (but also see below under C. 3.). Without knowing of possible
vectors, we simply cannot decide whether there had been a polycentric refuge including several
nunataks or massifs de refuge or if the species has obtained parts of its present distribution in
more recent times.

Of course we have to ask the question why speciation should take place in the small,
isolated populations on their refugia. Referring to this, the theory of “sexual selection by female
choice” (EBERHARD, 1985), is of importance. Using its arguments we might suppose that in a
small area with a low amount of natural resources (food, hiding-places, etc.) there must be an
intensive struggle between the males about the females. These favour males on the base of their
genitalia, so that we have a strong selective pressure to surpass the competition. As the direction
of this evolutionary process is of random, different populations will go different ways. The
smaller a population is, the faster this process will go on.

At smaller refugial areas or at the border of larger ones, suitable places for the survival will
be intermitted by hostile areas, e.g. ice streams. As a result of this partition we will probably
find a complex of sibling species or subspecies at such places that furnish proof for this.

A. 3. Easy to catch

This condition is of practical value for the researcher and certainly depends on his
experience. But there are some species that live at inaccessible places or occur in so low numbers
that they will be found only by chance. We know of many species that have been described as
being endemic and have later been found elsewhere. Good examples are subterranean species,
especially the phreatic species that live in small crevices deep inside the rocks, where equal
climatic conditions occur. Most of them have been described as cave species, simply because
caves are the only places where men and phreatic species can meet.

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JORG SPELDA

A. 4. Confirmation by other organisms

If we do not assume that the outlasting was only a special case in one species or genus, we
might expect that endemism may also occur at other groups. This means, the more endemic
species from different groups (millipedes, beetles, earthworms, etc.) occur, the bettei a possible
refuge is verified. On the other hand there may occur isolated populations of non-endemic
species at this places too. Although we cannot exclude a secondary immigration (even with
human influence, see below) they may support our conclusions.

B. Biogeographic reference for the refugia near Basel and Salzburg

B. 1. The refugia in general . .

Many of the classical localities of former investigators have been visited and in most cases
it was possible to confirm the occurence. It has been proved, that the genera Haasea,
Xylophageuma, Syngonopodium, Pyrgocyphosoma, Rhymogona and Listrocheiritium perform
the condition of being relatively easy to find. It is most important to choose the right time. At the
middle to late October they will be quite common under bark and dead wood. Nevertheless there
are differences between the genera: Rhymogona and Pyrgocyphosoma will be found quite
regulary in October, with no or only minute dependence from the weather. In spite of BIGLER s
(1913) opinion P. titianum is quite common, but strongly related to high humidity, so that it is
mostly found near springs and small brooks. Xylophageuma vomrathi is much more difficult to
catch, possibly contrary to its sibling species X. zschokkei, that was found quite common by
PEDROLI-CHRISTEN and myself during our excursions in the Vosges Mountains and the Jura.
But although we both collected Pyrgocyphosoma and Rhymogona quite regulary, PEDROLI-
CHRISTEN has never found X. vomrathi. Its occurence seems to depend on cold weather with
temperatures just above the freezing-point.

Other species, like the possibly phreatic ones Alpityphlus seewaldi (only one record
known, STRASSER, 1967). Polydesmus rothi and P. xanthocrepis have very seldom been
collected and seem to be not suitable for a survey. Nevertheless, as long as we have no
contradictions for their endemism, we may use them as additional arguments lor the refugia.

At the investigated sites the endemic species have been found sympatric and often syntopic
under the same bark, in community with a rich fauna of other millipedes. During the
investigations, at two places (Hasel, Scharfling) the maximal amount of endemic Choideumatida
has been found syntopic. This supports the assumption of a glacial refuge and contradicts the
conception that the endemism results from competition with superior species.

Both refugia contain limestone areas. This might be of general importance for glacial
refugia, as calcareous areas mean warmer soil and the crevices provide places with moderate,
although cold climatic conditions, that may render a retreat during hard limes.

B. 2. The “Basel-refuge”

When comparing the maps (Figs 1-3) P. titianum shows the most closed distribution of its
records, so that it is regarded as the most suitable species tor reconstructing the “Basel-
refugium”. The three endemic Rhymogona- species indicate a partition to at least three different
sites. A possibly similar species-complex occurs in the snail genus Bythiospeum.

Another endemic animal in the southern Black Forest is the earthworm Lumbricus
badensis. Its distribution (KOBEL-LAMPARSKI & LAMPARSKI, 1989) is much the same as in
P. titianum. Other endemisms in this region are known from snails ( Bythinella badensis,
Bythiospeum sterkianum, SCHMID, 1979; SCHMID, 1989). In the snail genus Trichia 4 endemic
species arc known from northern Switzerland and adjacent Germany (T. caelata, 7. clandestina,
T. graminicola , T. biconica , KERNEY et. al, 1983), with T. caelata having a similar distribution
as Polydesmus rothi (Fig. 1). In the caves of the northern Jura the endemic cave-beetle Royerella
villardi matheyi occurs. Somewhat more southward in the Jura we will find other cave animals
like Trichaphaenops sollaudi (Coleoptera), Trichoniscoides pulchellus (Isopoda),

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155

Archiboreoiulus sollaudi and Boreoiulus simplex (both Diplopoda). As a support from wider
distributed species we have isolated populations of subalpine plants like Primula auricula
(MEUSEL el al., 1965-92) and beetles like Nebria gyllenhali (missing in the Vosges Mountains)
that occur in the Black Forest.

Fig. 1. — Distribution of Pyrgocyphosoma tilianum and
Polydesmus rothi in southwestern Germany and
adjacent Switzerland.

The lack of the endemic Black-Forest-species, as well as related species in Switzerland
(PEDROL1-CHRISTEN, 1993), at the alpine level contradicts a nunatak-refuge and supports a
massif de refuge at lower altitudes. When we decrease the formerly glaciated areas (HANTKE,
1978-1983), the calcareous mountains at the southern border of the Black Forest remain as most
probable refuge. This might be supported by the sympatric occurence of members of all three
endemic genera and the junction of the ranges of the three endemic Rhymogona- species. In
agreement with a map provided by HANTKE (1978-1983) we are even able to attach the ice-free
parts of the southern Black Forest as refugia to each of the endemic Rhymogona- species.

FiG. 2. — Distribution of Xylophageuma vomrathi and
Haasea flavescens in southwestern Germany and
adjacent Switzerland.

Germany

• Pyrgocyphosoma
titianum
a Polydesmus
rothi

Lake

Constance

\ Rhine

Switzerland

J • Xylophageuma
. vomrathi

c» Haasea
flavescens

Germany

Lake

Constance

Rhine

Switzerland

Source

156

JORG SPELDA

Germany

Danube

c R. alemannica
■ R. cervina
n R. serrata
b R. wehrana
a R. verhoeffi
□ R. sp.

Rhine

Constance

Switzerland

Austria

FlG. 3. — Distribution of Rhymogona species in southwestern Germany and adjacent Switzerland.

B. 3. The “Salzburg-refuge”

The “Salzburg-refuge” can be delimited by the occurence of Syngonopodium aceris,
Haasea norica and Listrocheiritium noricum (Figs 4-6). At present time we still have gaps in the
knowledge of the millipede fauna in northern Austria, so that not too many conclusions about the
exact extension of the refuge should be drawn. The results are in accordance with HOLDHAUS
(1954), and we can add the beetles Trechus wagneri and Otiorhynchus schaubergeri as further
endemisms of this region. Very interesting is the occurence of the also endemic cave beetle
Trichciphaenops angulipennis in the Dachstein-mountains, an area regarded as beeing glaciated
during the Wiirm. But as phreatic/cave species are difficult to record this species might have
survived at the unglaciated sites nearby. Its distribution resembles Syngonopodium cornutum.
As the eastern Alps have only been slightly glaciated (HOLDHAUS, 1954), the number ol
endemic species increases eastward, so that we might assume a series of refugia along the
northern Alps from Salzburg on. This is supported by the distribution of the genus
Listrocheiritium , showing a sequence of species there. The isolated record of L. nubium in the
mountains of the “Totes Gebirge” is very strange and may refer to L. noricum as a possible
misidentification. L. cervinum has a wider distribution inside the Alps, and the new record
(“Tauernalm”) fills the gap between the mountains of Hochstaufen, Grimming and
GroBglockner. Listrocheiritium- species show a large vertical distribution, reaching the alpine
level and have possibly survived on both, nunataks and massifs de refuge. The Austrian record
of L. septentrionale (Rossatz) is identical with “Buchental bei Spitz” given by A'lTEMS (1949) as
locality for the never described L. nibelungiacum. Examination of topotypic specimen and
ATTEMS’ types proved their identity, although L. septentrionale has been described from a site
more than 80 km northerly. This may show, that in contrast to VERHOEFF'S (1917) opinion
large rivers like the Danube provide no hindrance for Chordeumatida. As most parts of northern

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MILLIPEDES AS AIDS FOR THE RECONSTRUCTION OF GLACIAL REFUGIA

157

Austria are myriapodological badly investigated this species might be well distributed in the
intermediate area.

Fig. 4. — Distribution of Haasea norica and Polydesmus xanihocrepis in the northeastern Alps.

Fig. 5. — Distribution of Alpityphlus seewaldi and Syngonopodium species in the northeastern Alps.

Isolated populations of wider distributed species, that support the “Salzburg-refuge” are
found in many snails e.g. Acicula gracilis and Renea veneta (KERNEY et al. , 1983). In respect of
this, the isolated occurence of the chordeumatid Haplogona oculodistincta at Scharfling is very
interesting, as it could belong to a relict population. As this site provides a rich, pretentious

Source :

158

JORG SPELDA

millipede fauna, synanthropism might be excluded. The species is distributed in the southeastern
Alps up to Vienna and adjacent Balkan Peninsula.

C. Discussion of problems and counter-arguments

C. 1 . Speciation after the Ice Age ,

Someone may argue, that the endemic species have been developed alter the time ol
°laciation An argument" for this is the occurrence of endemic plants like the Papaver radicatum
complex, on which a discussion for ice-free refugia in Scandinavia was based on. NORDAL
(1987) showed, that we can explain the endemism by postglacial immigration and subsequent
speciation. This is certainly in larger accordance with the geological results there. The question
is can we transfer this to animals? The arguments against are, that we have no similar
endemisms of animals in Scandinavia, and that plant “speciation” can occur extremely rapid by a
single mutation with distinct phenotypic effect. As the presence of a mate is not absolutely
necessary (self fertilisation) a single specimen can be the ancestor of a whole changed population
beside a refuge. Animals, if not parthenogenetic, are liable to bisexual propagation, that will
suppress extraordinary mutations if they are not advantageous. In times ot spieading, when
there are less meetings of males and females sexual selection would be less effective
(EBERHARD, 1985).

C. 2. The character of natural borders

VERHOEFF (1917) was the first who discovered the importance of rivers as distributional
borders. An argument against them as absolute borders for millipedes might be the occurence of
species with generally small ranges on both sides of the Danube ( Listrocheiritium septentrionale ,

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MILLIPEDES AS AIDS FOR TIIF. RECONSTRUCTION OF GLACIAL REFUGIA

159

Haploporatia eremita, Leptoiulus marcomcinnius) and the Rhine ( Rhymogona cervina,
R. alemann ica, Orthochordeumella fulva, Polydesmus helveticus, “Helvetischer
Rheintaldurchbruch” of VERHOEFF, 1917).

With respect to EBERHARD's ( 1985) hypotheses I will propose another explanation. If two
species have been separated only by sexual selection they still use the same ecological niche. As
we know that two species cannot occupy the same niche at the same time, they must occur
allopatric or parapatric (vicariance). If we assume that their selective fitness is quite equal, the
native species is in favour against the invasive. As a result we will have relative stable borders at
places that will not be crossed so often like rivers and mountain ridges. This explains the
complex distribution of the genus Rhymogona in southwestern Germany much better, as the
distribution follows small rivers and brooks with species often changing between parallel ones
and - on the other hand - changing between larger rivers like the Rhine and the lower course of
the Wiese. The parapatric distribution of the related species Xylophageuma vomrathi and Haasea
flavescens (Fig. 2) migth be another case of parapatry by ecological competition.

C. 3. “Natural” extinction

Endemism can not only be explained by speciation e.g. during the ice-age, including
reduction of the areal and - up to now - only a minute reimmigration, but also by reduction of the
distribution caused by other (ecological) effects like competitors, enemies and diseases. The
occurence of such palaeoendemisms may simulate a glacial refuge, but normally in this case we
will have more than one isolated population of the same species (meaning places where the
hostile conditions are not effective) or they occur on special places, where other species cannot
survive (displacement to extreme sites). In spite of this, the places where the endemic millipedes
occur contain a very rich fauna so that we may regard displacement as less probable.

C. 4. Human influences

There might be the possibility that a species is delimited to a small area that cannot be left
naturally. But if transported by men, perhaps with the earth surrounding a plant, it may arrive a
place where spreading means no problem. This has been followed quite profound in North
America by the invasion of European species. Perhaps this is also a good explanation for the
colonization of the British Island by millipedes, as they might have been companions of early
settlers. On the other hand devastation by men took part in many of the areas populated by him,
e.g. by cutting down the forest. So we might also think of the change of areals by men in recent
and former days.

CONCLUSIONS

Comparison of the refugia presented here with the map of true cave-animals given by
HOLDHAUS (1954) shows, that the northernmost parts of their distribution cover with them.
Also a map provided by JANETSCHEK (1956) shows our refugia being included, but as
demonstrated here, they are more ensured and obviously of larger importance than most of the
other north-alpine “refugia” shown by him. So there is much evidence, that the southern part of
the Black Forest and the mountains of the Salzkammergut are the northernmost places where a
pretentious fauna have survived at least the last glaciation (Wurm) in middle Europe (excluding
ihe Carpathians). Although there might have been no larger trees, there is much evidence that
Salix- species have survived the last cold stage north of the Alps (BENNET el al, 1991). Their
litter might have served as food for the persistent diplopods.

An interesting aspect has been pointed out on forest trees by BENNET et al. (1991), saying
that remainig tree populations at mid-altitude sites in the mountainous areas of southern Europe
are most important for the long-term survival of species, as they cannot follow the rapid climatic
changes. As a result, the refugia are important at all times, both cold and warm stages. This
might also be true for animals with a low tendency of spreading out and may explain the
“petrophilism" just because of the fact that mountains show closed sites with different climatic

160

JORG SPELDA

conditions, that allow changes to suitable biotopes, so that a long-distance travelling to such
places is not necessary in cases of global climatic changes.

As no one can travel through time and test hypotheses concerning the past, we have to use
the “principle of parsimony”, that is also used in the discussion ot phylogenetic pathways,
meaning that we favour hypotheses that need less additional assumptions than others. 1 he
present distribution of organisms seems to be an important argument in cases where we have no
pollen evidence or macrofossils. With the exception of three phreatic/cave species (Alpityphlus
seewaldi . Polydesmus xanthocrepis and P. rothi) the endemic millipedes all belong to the
Chordeumatida. This shows, that the members of this order are the most important indicators tor

our purpose. , . . r

Up to now, no other group is known, showing so much endemism in the buropean

mountains as the millipedes (VERHOEFF, 1938b) and especially the chordeumatids. These
species - “Glazialresistente” of VERHOEFF (1917) - must be considered as beeing preglacial or at
least interglacial relicts.

ACKNOWLEDGMENTS

The author is greatly indebted to Professor Dr. h. c. Burkhard Frenzel (University of Stuttgart- Hohenheim,

Institut of Botany), Dr Sergei I. Golov atch (Russian Academy of Science, Moscow) and Pro!. Dr. Hinnch Rahmann

(University of Stuttgart -Hohenheim, Institut of Zoology) for giving valuable advices. I also wish to thank my

colleagues Dr. Jurgen BOhmer and Dietmar Rothmund for critical reading of the manuscript.

REFERENCES

Attems, C. 1949. — Die Myriopodenfauna der Ostalpcn. SB Ak. Wien., math.-naturw. Kl, /. 158 : 79-153.

Bennett. K. D.. Tzedakis, P. C. & Willis. K. J., 1991. — Quaternary refugia of north European trees. J. Biogeogr., 18
103-115.

BIGLER, W., 1913. — Die Diplopoden von Basel und Umgebung. Rev. suisse Zool , 21 : 675-793.

EBERHARD. W. G., 1985. — Sexual selection and animal genitalia. Cambridge & London. C.U.P., 255 pp.

Hantke. R.. 1978. — Eiszeitalter. Thun, Vol. 1, 467 pp.

Hantke. R., 1980. — Eiszeitalter. Thun, Vol. 2. 703 pp.

Hantke, R.. 1983. — Eiszeitalter. Thun, Vol. 3, 730 pp.

HOLDHAUS, K.. 1954. — Die Spuren der Eiszeit in der Tierwelt Europas. Abh. zool.-bol. Ges. Wien , 18 : 1-493.

JANETSCHEK. H., 1956. — Das Problem der inneralpincn Eiszeitubcrdauerung durch Tiere (Ein Beitrag zur Geschichte der
Nivalfauna). Osterr. tool. Zeitschrift, 6 : 421-506.

KERNEY. M. P., Cameron, R. A. D. & JUNGBLUTH, J. H., 1983. — Die Landschnecken Nord- und Mitteleuropas . Hamburg
& Berlin, Paul Parey, 384 pp.

Kobel-Lamparski. a. & Lamparski, F.. 1989. — Der Badische Regenwurm Lumbricus badensis und andere
bemerkenswerte Regenwiirmer aus dem Belchengebiet. In : Der Belchen. Natur- und Landschaftschutzgebiete Bad.-,
Wiirtt ., Karlsruhe, 13 : 891-905.

MEUSEL., H.. JAGER. E. J. & WEINERT. E.. 1965. — Vergleichende Cliorologie der zeniraleuropaischen Flora. Vol. I.
Jena, 582+258 pp.

MEUSEL.. H„ JAGER. E. J. & WEINERT. E.. 1978. — Vergleichende Chorologie der zeniraleuropaischen Flora. Vol. 2,
Jena, 418+162 pp.

MEUSEL.. H.. JAGER, E. J. & WEINERT, E.. 1992. — Vergleichende Chorologie der zeniraleuropaischen hlora. Vol. 3 ,
Jena, 333+256 pp.

Nordal, I., 1987. — Tabula rasa after all? Botanical evidence for ice-free refugia in Scandinavia reviewed. J. Biogeogr .,
14 : 377-388.

Pedroli-ChrisTEN, A., 1993. — Faunistique des millc-pattes de Suisse (Diplopoda) / Faunistik der I ausendliissler dei
Schweiz (Diplopoda). Neuchatel, Centre Suisse de Cartography de la Faune, Doc. faun, helv., 14. 248 pp.

SCHMID, G.. 1979. — Mollusken vom Grenzacher Horn. In : Der Buchswald bei Grenzach (Grenzacher Horn). Natur- und
Landschaftschutzgebiete Bad. -Wiirtt., Karlsruhe, 9 : 225-359.

SCHMID, G., 1989. — Schnecken und Muscheln vom Belchen. In : Der Belchen. Natur- und Landschaftschutzgebiete Bad.-
Wiirtt. , Karlsruhe. 13 : 891-905.

Source : MNHN. Paris

MILLIPEDES AS AIDS FOR THE RECONSTRUCTION OF GLACIAL REFUGIA

161

Spelda. J., 1991. — Zur Faunistik und Systematik der TausendfuBler (Myriapoda) Sudwestdeutschlands. Jh. Ges.
Naturkde. Wiirttemberg, 146 : 211-232.

Strasshr, K., 1967. — Ein Typhloiuline aus den nordlichen Kalkalpen (Diplopoda, Symphyognatha). Ber. nat.-med.
Ver. Innsbruck, 55 : 145-154.

Verhoeff, K. W.. 1917. — Zur Kenntnis der Zoogeographie Deutschlands zugleich iiber Diplopoden namentlich
Mitteldeutschlands und Beitrage fiir die biologische Beurteilung der Eiszeiten. Nova Acta. Abh. der Kaiserl. Leop.-
Carol. Deutschen Akademie der Naturforscher, 103 : 1-157.

VERHOEFF, K. W., 1938a. — Diplopoden der Germania zoogeographica im Lichte der Eiszeil. Zoogeographica, 3 . 494-
547.

Verhoeff, K. W., 1938b. — Ein halbes Jahrhundert Diplopodenforschung und ihre Bedeutung fur die Zoogeographie.
Zoogeographica , 3 : 548-588.

Source : MNHN, Paris

On the Distribution and Faunogenesis of Himalayan
Millipedes (Diplopoda): Preliminary Results1

Sergei I. GOLOV ATCH * & Jochen MARTENS **

* Institute for Problems of Ecology and Evolution, Russian Academy of Sciences
Leninsky prospekt 33, Moscow V-71, Russia
** Institut fur Zoologie, Johannes Gutenberg-Universitat
Saarstrasse 2 1 , D-55099 Mainz, Germany

ABSTRACT

The fauna of Diplopoda of the Himalayas (over 200 species, mostly endemic) is reviewed, with particular reference to
that of their central part. In spite of the preliminary state of knowledge, the patterns of vertical and geographic
distributions suggest the fauna to be eventually entirely Oriental and/or Indian in origin, and primarily associated with
forest tropical and/or subtropical communities. The so-called Palearctic influence in the relatively well-explored Central
Himalayas actually also originates in the present-day subtropical regions of Southeast and East Asia. The Himalayas
seem to have served as a pathway for repeated spreads of a uniform Turgai biota (with Diplopoda being an accompanying
group) which, chiefly during the early and middle Tertiary, advanced northwestward, following the receding southern
coast of the Tethys Sea. Naturally, during their relatively short orogenic history, the Himalayas also served as a major
center of secondary diversification for numerous groups, especially during the Plio-Pleistocene.

RESUME

Repartition et genese des faunes de Diplopodes de l’Himalaya : resultats preliminaires.

La faune des diplopodes des massifs himalayens (plus de 200 especes, la plupart endemiques), plus particulidrement de
leur partie centrale, est revisee. Les modalit£s de la repartition verticale et geographique suggerent pour cette faune une
origine entierement orientale et/ou indienne. primitivement associee k des peuplement forestiers tropicaux et/ou
subtropicaux. L’ influence dite palearctique provient, dans les regions relativement bien explores des chaines centrales
de f Himalaya, des aires subtropicales actuelles du Sud-Est et de 1’Est asiatique. Les massifs himalayens semblent avoir
servi de “bordure" & des extensions repetees d'un type d’ecosysteme uniforme de type “Turgai” (les diplopodes
apparaissant commc un groupe accompagnateur) qui, surtout durant le debut et le milieu de l’fcre tertiaire, a progresse vers
le Nord-Ouest a la suite du recul de la cote meridionale de la Tethys. Bien entendu, durant cette histoire orogenique
relativement courte, les massifs himalayens ont ete £galement un centre majeur de diversification secondaire pour de
nombreux groupes d’etres vivants, notamment au cours du Plio-Pleistocene.

1 Results of the Himalaya Expeditions of J. Martens, No. 201. — For No. 200 see: Bonner Zoologische Monographien,
39, 1995. — J. M. sponsored by Deutscher Akademischen Austauschdienst, Deutsche Forschungsgemeinschaft and
Feldbausch Foundation, Fachbereich Biologie, University Mainz.

Golovatch, S. I. & Martens, J., 1996. — On the distribution and faunogenesis of Himalayan millipedes
(Diplopoda): preliminary results, hi: Geoffroy, J.-J., M.AURifcS, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 163-174. Paris ISBN : 2-85653-502-X.

164

SERGEI I. GOLOV ATCH & JOCHEN MARTENS

INTRODUCTION

Being one of the globe's greatest mountainous lands supporting the highest peaks such as
Everest, Kanchenjunga, Manaslu, Annapurna, etc., the Himalayas occupy a vast area
encompassing the ranges lying between the rivers Indus and Brahmaputra and roughly delimited
by 74°E in the west and 95°E in the east (Fig. 1). The adjacent Karakorum and Kohistan-
Baluchistan regions in the west, and the Arakan-Chin-Yoma fold belt and the Magok belt in the
east are of the same orogenesis (MASCLE et al., 1990).

From a biologist's viewpoint, the Himalayas represent a highly important barrier between
the cold and arid uplands of Central Asia and largely tropical South and Southeast Asia,
reinforcing the contrast and, in spite of numerous local inversions, creating their own climate.
During the southwestern monsoon period, precipitation mainly occurs on the southern slopes,
being greatly reduced on the northern ones. However, this barrier function holds true only for
the central parts of the mountains, more or less within Kumaon, Nepal, Sikkim, and Bhutan. In
the western Himalayas, the aridity of Central Asia extends across the southern slopes, while in
the eastern parts rainstorms, though declining in amount and frequency, reach as far as
Southeast Tibet (TROLL, 1967).

This drastic climatic gradient within the Central Himalayas is of great importance,
influencing the distribution of various organisms. Although phyto- and zoogeographic regions
differ in certain details, both emphasize the role of the Himalayas as a contact region between
two great biogeographic realms, the Palearctic and Oriental, which meet and intermesh there in
various ways. All areas north of the Central Himalayas obviously belong to the Palearctic, as do
the highest parts of the inhabited southern flanks. The lower and lowest altitudes of the southern
slopes are largely attributable to the Oriental realm. However, the border between both regions is
generally neither striking nor abrupt, forming more (especially in the eastern Himalayas) or less
(in their central parts) vast transition areas, numerous inversions or anomalies. In other words,
the otherwise manifest rule “(sub)tropical organisms for (sub)tropical environments only” is

Source :

DISTRIBUTION AND FAUNOGENESIS OF HIMALAYAN MILLIPEDES

165

very often violated in the Himalayas, particularly in the central parts of this great mountainous
land and especially as regards animals (e.g. MARTENS, 1984, 1993). Even the altitudinal
zonation of Himalayan plant communities is rather conventional (DOBREMEZ, 1972) (Fig. 2).

nival stage

submval stage

scattered patches eternal snow

of vegetation " -

<li

upper

alpine level

thorn steppes

alpine meadows

lower

alpine level

heath with dwarf
Rhododendron
and jumpers

heath with dwarf
Rhododendron

o .

" 5

2 g |

f S i

2 c 8

— 52 2
8 1

N 3

at cl —
a §• 5 °
<x> $

a. o

E “

upper

subalpine level

birch forest

forest of tree
Rhododendron
and fir (Abies)

lower

subalpine level

fir forest

mountain region

Cedrus,
Cupressus
and Juniperus
forest

coniferous (Pmus.
Piceo) and deciduous
(Quercus) forest

hygrophilic Quercus
forest

hill region

forest of evergreen oaks and laurels

s S
s 1
s §

a. o

E E

| §
i/i j

upper

subtropical level

Olea forest

pine forest
(Pinas roxburghn)

subtropical deciduous
forest (Schimo.
Costonopsis )

lower

subtropical level

northwest IV

tropical zone

collme zone U

upper

tropical level

tropical forests, mainly Shoreo

lower

tropical level

dry

mesophilic

damp

west III

central. 11

east I

■6000

■5500

5000

4500

4000

3000

2600

2000

1500

1000

400
0

Fig. 2. — The vegetation belts and most important plant communities in the Nepal Himalayas. The Roman numerals at

the bottom indicate the floral regions of Nepal (modified, after Dobremez, 1972).

The present paper is the first attempt to trace the vertical distribution patterns of Himalayan
millipedes which is basic for reliable faunogenetic reconstructions. This study is mainly
restricted to the fauna of Nepal, the Himalayas' centralmost and particularly well-explored part,
but all available information is also incorporated on the faunas of adjacent areas and of some
ecologically similar soil/litter-dwelling animal groups for comparative purposes. At the start,
these results must be regarded as quite preliminary, for a considerable proportion (perhaps over
50%) of the existing collections of Diplopoda remains untreated. In addition to published
material, certain unpublished data are presented, chiefly derived by the senior author from the
long-term research project conducted by the junior author and his collaborators since 1969
(MARTENS, 1987a; GOLOV ATCH, 1990). In spite of its preliminary character, this paper seems
warranted to draw some general conclusions on the patterns of diplopod faunogenesis. The latter

166

SERGEI I. GOLOV ATCH & JOCHEN MARTENS

is the main objective, while the patterns of altitudinal distribution of Himalayan millipedes are an
important tool.

DIPLOPODA OF THE HIMALAYAS

Table 1 presents all available information on the millipede fauna of, and its distribution
within, the Himalayas, with over 200 species represented. However, some taxonomic remarks
are necessary. A few genera are between quotation marks, for they are obscure either as taxa or
as Himalayan elements (see HOFFMAN, 1980). In some cases, no information on elevations is
available, this being reflected by a question mark [(?)]. Introductions (N° 1 83-184) are extremely
rare and are referred to as synanthropic.

Table 1. — Geographic and vertical distribution of Himalayan diplopods.

Taxa

Country

Elevations ( m. )

Order Polyxenida

Family Polyxenidae

Genus Polyxenus Latreille, 1802-03

1. Polyxenus sp.

Kashmir

1585

Genus Monographis Attems, 1907

2. M. mirus (Turk, 1947)

Kumaon

1600

Genus Unixenus Jones, 1944

3. Unixenus sp.

Nepal

4550

Order Sphaerotheriida

Family Sphaeropoeidae

Genus Indosphaera Attems, 1936

4. Indosphaera curiosa Attems, 1936

Assam

Genus Kophosphaera Attems, 1 936

5. K. brevilamina Attems, 1936

North Bengal, Darjeeling Distr.

1700

6. K. devolvens Attems, 1936

Sikkim, Darjeeling Distr.

1700-2050

7. K. excavata (Butler, 1874)

Nepal, Assam

8. K. excavata mammifera Attems, 1936

Darjeeling Distr., Assam

9. K . politissima Attems, 1936

Darjeeling Distr.

1700

Genus “ Sphaeropoeus ” Brandt, 1833

10. 5. rnontanus Karsch, 1881

Himalayas

Genus “ Sphaerotherium” Brandt, 1833

11.5. maculatum Butler, 1874

Sikkim

12.5. politum Butler, 1874

Sikkim

Genus “ Zephronia " Gray, 1 832

13. Z. alticola Attems, 1936

Assam, Darjeeling Distr.

400-1700

14. Z. debilis Attems, 1936

Darjeeling Distr.

1700

15. Z. densipora Attems, 1936

Assam

16. Z. disparipora Attems, 1936

Assam

140

17. Z. hirta Attems, 1936

Darjeeling Distr.

1700

18. Z. hysophila Attems, 1936

Assam

19. Z. juvenis Attems, 1936

Assam

20. Z. laevissima Butler, 1874

Sikkim

21. Z. lignivora Attems, 1936

Assam

180-330

22. Z. manca Attems, 1936

Vietnam. Darjeeling Distr.

1000-1700

23. Z. nigrinota Butler, 1872

Darjeeling Distr.

2300-2700

24. Z. specularis Attems, 1936

Assam

25. Z. tigrinoides Attems, 1936

Darjeeling Distr.

170

26. Z. tumida Butler, 1882

Assam, Burma

27. “ Zephronia ” spp.

Nepal

250-500

Order Glomerida

Family Glomeridae

Genus Hyleoglomeris Verhoeff, 1910

28. H. crassipes Golovatch, 1987

Nepal

2450-2720

29.//. electa Silvestri, 1917

Darjeeling Distr.

500-1700

Source : MNHN: Paris

DISTRIBUTION AND FAUNOGENESIS OF HIMALAYAN MILLIPEDES

167

30. H. gorkhalis Golovatch, 1987

Nepal

1200

31. H. khumbua Golovatch, 1987

Nepal

3250-3300

32. H. modesta Silvestri, 1917

Assam

150

33. H. nagarjunga Golovatch, 1987

Nepal

1900-2100

34. H. tinjurana Golovatch, 1987

Nepal

2450

35. H. venus tula Silvestri, 1917

Assam

Order Siphonophorida

Family Siphonophoridae

Genus Pterozonium Attems, 1951

36. P. cingulatum (Attems, 1936)

Vietnam, Darjeeling Distr.

500-1700

37. P. coniceps (Attems, 1936)

Darjeeling Distr.

1700

38. P. lanx’oodi (Turk. 1947)

Kumaon

1600

Order Platydesmida

Family Andrognathidae

Genus Pseudodesmus Pocock. 1887

39. ? Pseudodesmus sp.

Nepal

<2000

Order Chordeumatida

Family Cleidogonidae

Genus Tianella Attems, 1904

40. T. ausobskyi Shear, 1987

Nepal

2500-3050

41. T. bobanga Shear, 1979

Nepal

2460-2500

42. T. daamsae Shear, 1987

Nepal

3600-3900

43. T. gitanga Shear, 1987

Nepal

2550

44. T. jaljalensis Mauries, 1988

Nepal

2350

45. T. kathmandua Maurids, 1988

Nepal

1700

46. T. lughla Shear, 1979

Nepal

2950-3300

47. T. mananga Shear, 1987

Nepal

2550

48. T. mangsingma Mauri&s, 1988

Nepal

2250

49. T. martensi Shear, 1979

Nepal

1150-2900

50. T. smetanai Mauries, 1988

Nepal

3250

51. Tianella sp.

Darjeeling Distr.

900-1400

Family Kashmireumatidae

Genus Kashmireuma Maurids, 1982

52. K. nepalensis Mauries, 1988

Nepal

3600-4100

53. K. nielseni Mauries, 1982

Kashmir

2600-3500

54. K. schawalleri Shear, 1987

Nepal

3450-3600

55. Kashmireuma sp.

Nepal

2500-3000

Family Megalotylidae

Genus Nepalella Shear, 1979

56. N. deharvengi Mauries, 1988

Nepal

2900-3500

57. N. gairiensis Mauries, 1988

Nepal

3000

58. N. gunsa Shear, 1987

Nepal

3600-3800

59. N. jaljalae Mauries, 1988

Nepal

2200

60. N. khumbua Shear, 1979

Nepal

3250-3300

61. N. phulcokia Mauries, 1988

Nepal

2250

62. N. ringmoensis Mauries. 1988

Nepal

2750-3000

63. N. taplejunga Shear, 1987

Nepal

3000-3300

64. N. thodunga Shear, 1979

Nepal

3200

65. N. tragsindola Mauries, 1988

Nepal

2450-3000

66. Nepalella sp.

Nepal

1900-4100

Order Julida

Family Julidae

Genus Anaulaciulus Pocock, 1895 (cf. KorsOs, 1996)

67. A. acaudatus Korsds, 1996 Sikkim

3990

68. A. bilineatus Kors6s, 1996

Nepal

3300-4300

69. A. kashmirensis Korsos, 1996

Kashmir

3100-3200

70. A. nepalensis Korsds, 1996

Nepal

2600-3400

71. A. niger Korsds, 1996

Nepal

2600-4500

72. A. tibetanus Korsds, 1996

China (E-Tibet), Assam

3700

73. A. topali Korsds, 1996

Kashmir

2300

Source : MNHN , Paris

168

SERGEI I. GOLOV ATCH & JOCHEN MARTENS

Genus Nepalmaioiuliis Mauries, 1983

74. N. appendiculatus Enghoff, 1 987

Kumaon

1900-2100

75. N. defiarvengi (Mauries, 1983)

Nepal

2550-3350

76. N. dhaulagiri Enghoff, 1987

Nepal

3000-3350

77. N. generalis Enghoff, 1987

Nepal

3400

78. N. hyalilobus Enghoff. 1987

Nepal

3600-3800

79. N. ivanloebli Enghoff, 1987

Nepal

2200-4800

80. N. juxtapositus Enghoff, 1987

Nepal

2800-3050

81. N. martensi Enghoff. 1987

Nepal

3250-3300

82. N. mauriesi Enghoff, 1987

Nepal

3600

83. N. nigrescens Enghoff. 1987

Bhutan

2300

84. N. pineti Enghoff, 1987

Nepal

2900

85. N. rugiflagrum Enghoff, 1987

Bhutan

3300

86. N. smetanai (Mauries, 1983)

Nepal

1900-2700

87. N. sympatricus Enghoff. 1987

Nepal

3000

88. N. uncus Enghoff, 1987

Nepal

2550

89. N. wuermlii Enghoff, 1987

Bhutan

1680-2600

90. N. zachonoides Enghoff, 1987

Nepal

2450-2600

Order Spiros trepti da

Family Harpagophoridae

Genus Gonoplectus Chamberlin, 1921

91. G. alius Demange, 1961

Assam

92. G. bhutanensis Demange, 1988

Bhutan

350-450

93. G. broelemanni Demange, 1961

Nepal

1800-2300

94. G. corniger (Attems, 1936)

Assam

95. G. gracilis (Attems, 1936)

Darjeeling Distr.

1200

96. G. hyatti Demange. 1961

Nepal

1200

97. G. lindbergi Demange, 1961

Darjeeling Distr., Bhutan

350

98. G. malayus (Carl, 1909)

Kumaon, Nepal, Bhutan

200-2500

99. G. probus (Attems, 1936)

Darjeeling Distr.

1000

100. G. remyi Demangc, 1961

Assam

101. G. sulcatus (Attems, 1936)

Darjeeling Distr.

2400

Order Cambalida

Family Cambalopsidae

Genus Podoglyphiulus Attems, 1909

102. P. elegans nepalensis Mauries, 1983

Nepal

<1000

Genus Trachyjulus Peters, 1 864

103. r. minius Silvestri, 1924

Assam

1200

104. T. wilsonae Mauries, 1983

Nepal

<1000

Order Spirobolida

Family Physobolidae

Genus Physobolus Attems, 1 936

105. P. olivaceus Attems, 1936

Darjeeling Distr.

1800.

Order Polydesmida

Family Cryptodesmidae

Genus Trichopeltis Pocock, 1894

106. T. watsoni Pocock. 1894

Darjeeling Distr., Assam,

Bhutan, West Bengal, Bangladesh

350-1000

Family Fuhrmannodesmidae

Genus Assamodesmus Manfredi, 1954

107. A. lindbergi Manfredi, 1954

Assam

Genus Hingstonia Carl, 1935

108. H. beaiae Golovatch, 1990

Nepal

2400-3500

109. H. dorjulana Golovatch, 1988

Bhutan

2450-3100

1 10. H. eremita Carl, 1935

Nepal

2000

III. H. falcata Golovatch, 1 986

Nepal

2650

112. H. fittkaui Golovatch, 1990

Nepal

3550-3650

113. H. gogonana Golovatch, 1988

Bhutan

3650-4000

114. H. pahakholana Golovatch, 1990

Nepal

2600-2800

115. H. pelelana Golovatch, 1988

Bhutan

3300-3400

116. H. perarmata Golovatch, 1986

Nepal

3150

DISTRIBUTION AND FAUNOGENESIS OF HIMALAYAN MILLIPEDES

117. H. serrata Golovatch, 1987

118. H. sympatrica Golovatch, 1990

119. H. variata Golovatch. 1987

120. Hingstonia sp.

Genus Magidesmus Golovatch, 1988

121. M. affinis Golovatch, 1988

122. M. bhutanensis Golovatch, 1988
Genus Sholaphilus Carl, 1932

123. S. asceticus Golovatch, 1986

124. S. dalai Golovatch, 1986

125. S. gompa Golovatch, 1990

126. S. lama Golovatch, 1986

127. S. martensi Golovatch, 1986

128. S. monachus Golovatch, 1990
Genus " Pseudosphaeroparia ” Carl. 1932

129. P. cavernicola Turk, 1945

Genus Topalodesmus Golovatch, 1988

130. T . communis Golovatch, 1988
Family Polydesmidae

Genus Bhutanodesmus Golovatch, 1988

131. D. velatus Golovatch, 1988
Genus Glenniea Turk, 1945

132. G. bhotiaensis Golovatch, 1988

133. G. indica Turk, 1945

134. G. minuscula Golovatch, 1988

135. G. perarmata Golovatch, 1988

136. G. martensi (Golovatch, 1987)

Genus Himalodesmus Golovatch, 1986

137. H. aiidax Golovatch, 1986

138. H. benefactor Golovatch, 1987

139. H. faustus Golovatch, 1987

140. H. parvus Golovatch, 1987

141. H. prosperus Golovatch, 1990

142. H. pulcher Golovatch, 1987

143. H. pygmaeus Golovatch. 1986

144. H. vigens Golovatch. 1987

Genus Typhlopygmaeosoma Turk, 1972

145. T. hazeltonae Turk, 1972

Genus Usbekodesmus Lohmander, 1932

147. U. buddhis Golovatch, 1986

148. U. occultus Golovatch. 1986

149. U. sacer Golovatch, 1987

150. U. theocraticus Golovatch, 1990

151. U. theosophicus Golovatch, 1986

152. Usbekodesmus sp.

Family Opisotretidae

Genus Martensodesmus Golovatch, 1987

153. M. bicuspidatus Golovatch, 1988

154. M. excornis Golovatch, 1988

155. M. himalayensis Golovatch, 1987

156. M. nagarjungicus Golovatch, 1987

157. M. sherpa Golovatch, 1987

158. Martensodesmus sp.

Family Paradoxosomatidae

Genus Armolites Golovatch, 1984

159. A. chulingensis Golovatch, 1994

160. A. communicans Golovatch. 1992

161. A. similis Golovatch. 1992

162. A. spiniger (Attems, 1936)

Genus Hirtodrepanurn Golovatch, 1994

163. H. latigonopum Golovatch, 1994

Nepal

3400-3600

Nepal

3550-3650

Nepal

2600-4500

Nepal

2200-3900

Bhutan

3300-3400

Bhutan

3100

Nepal

1300-1650

Nepal

2400

Nepal

2000-2100

Nepal

1800-2000

Nepal

1100-1850

Nepal

2050-2150

Kumaon

2800

Darjeeling Distr.

2000-2200

Bhutan

350-450

Bhutan

350-450

Kumaon

2800

Bhutan

1900-2300

Bhutan

1680

Nepal

1200

Nepal

2650

Nepal

2600-3400

Nepal

1000-1750

Nepal

2200

Nepal

2600-2800

Nepal

2450

Nepal

3300-3400

Nepal

2150-2250

Kumaon

1850

Nepal

3300-3400

Nepal

2300-2800

Nepal

3300-3400

Nepal

2600-2800

Nepal

3200

Nepal. Bhutan

3450-4250

Bhutan

1650-2000

Bhutan

2440

Nepal

1100-1300

Nepal

1900-2100

Nepal

1200

Nepal. Bhutan

1300-2150

Nepal

3000-3700

Nepal

2650

Nepal

2300-2700

Darjeeling Distr.

1000-2200

170

SERGEI I. GOLOV ATCH & JOCHEN MARTENS

Genus Kaschmiriosoma Schubart, 1 935

164. K. contortipes Schubart. 1935

Kashmir, N-Pakistan

2300-3300

165. K. nulla (Attems, 1936)

Himachal Pradesh

1000

166. K. pleuroptera (Attems. 1936)

Punjab (Pakistan)

2800

Genus Kronopolites Attems. 1914

167. K. occidemalis Golovatch. 1983

Kashmir

1500

Genus Martensosoma Golovatch. 1992

168. M. elegans Golovatch. 1992

Nepal

1350

169. M. foveatum Golovatch. 1992

Nepal

1800-2000

170. M. schawalleri Golovatch. 1992

Nepal

1000-2150

171. M. silvestre Golovatch, 1994

Nepal

2000-2600

172. M. splendens Golovatch. 1992

Nepal

1650-2150

173. M. unicolor ( Attems. 1936)

Assam. Darjeeling Distr.

1200-1700

Genus Nepalomorpha Golovatch, 1993

174. N. arunensis Golovatch, 1994

Nepal

1850-2150

175. N. hirsuta Golovatch, 1994

Nepal

3900-4100

176. N. kuznetsovi Golovatch, 1994

Nepal

3000

177. N. spinigera (Golovatch, 1992)

Nepal

600-1400

Genus Orophosoma Jeekel, 1980

178. O. fechteri Golovatch, 1990

Nepal

2330-3150

179. 0. hingstoni (Carl, 1935)

Tibet

3400

180. O. simulans (Carl. 1935)

Nepal, Tibet

3700

181. Orophosoma sp.

Nepal

1750-3450

Genus Orthomorpha Bollman, 1893

182. “O. ” almorensis Turk, 1947

Kumaon

1600

183. O. coarciata (Saussure, 1860)

Nepal (synanthr.)

600-650

Genus Oxidus Cook. 1911

184. O. gracilis (C. L. Koch, 1847)

Nepal (synanthr.)

570-1200

Genus Paranedyopus Carl, 1932

185. P. affinis Golovatch, 1990

Nepal

2475-2700

186. P. cylindricus (Carl, 1935)

Nepal, Darjeeling Distr.

1650-2850

187. P. elongissimus Golovatch, 1984

Darjeeling Distr.

1000

188. P. martensi Golovatch, 1990

Nepal

2250-3600

189. P. schawalleri Golovatch, 1990

Nepal

2050-2150

190. P. similis Golovatch, 1990

Nepal

2300-3000

191. Paranedyopus sp.

Nepal

2450-2900

Genus Par orthomorpha Golovatch, 1994

192. P. affinis Golovatch, 1994

Nepal

1400

193. P. granulosa Golovatch, 1994

Nepal

2000

194. P. intermedia Golovatch, 1994

Nepal

1000-1100

195. P. longiseta Golovatch, 1994

Nepal

1400-1600

196. P. nyakensis (Golovatch. 1992)

Nepal

2270-2450

197. P. philosophica Golovatch, 1994

Nepal

1650-2450

198. P. spectabilis Golovatch, 1994

Nepal

2650

199. P. tergalis Golovatch, 1994

Nepal

2650

200. P. tuberculata Golovatch, 1994

Nepal

3000-3300

Genus Substrongylosoma Golovatch, 1984

201. S. distinctum Golovatch, 1984

Darjeeling Distr.

1200-1500

202. S. falcatum Golovatch, 1984

Darjeeling Distr.

1400

203. S. montigena (Carl, 1935)

Darjeeling Distr.

1200-2300

204. S. schawalleri Golovatch, 1993

Nepal

1620-2000

Genus Topalosoma Golovatch, 1984

205. T. setiferum Golovatch, 1984

Darjeeling Distr.

900

Genus Touranella Attems, 1937

206. T himalayaensis Golovatch, 1994

Nepal

2300-2700

Family Pyrgodesmidae

207. Several genera and species

Nepal

450-1200

DISTRIBUTION AND FAUNOGENESIS OF HIMALAYAN MILLIPEDES

171

ZOOGEOGRAPHIC PATTERNS

The vast majority of Himalayan millipede species are local in distribution; there are few
relatively widespread species like Trichopeltis wcitsoni. Indeed, most Himalayan millipede
species are known from a single locality only, and many others appear to be restricted not only
in area, but also in altitude.

Conversely, most genera occur through a range of altitudes, as shown in Figure 3, but are
more or less restricted to forests, demonstrating sylvicoly. Table 1 and Figures 2-3 show that
the alpine zone of the Central Himalayas is only marginally populated by millipedes, whereas the
tropical and subtropical forest belts support the bulk of the fauna. This pattern conforms to
general knowledge that millipedes are primarily a class of forest floor-dwellers, which in
temperate regions of Eurasis seems trophecologically and historically associated with nemoral
(= broadleaved) forest communities (GOLOV ATCH, 1987, 1991a). In turn, this pattern provides
the basis for faunogenetic reconstructions based on phyto- and paleogeographic evidence.

Fig. 3. — Vertical distribution of some millipede genera in the Central Himalayas - genera Usbekodesmus, Hingstonia,

Sholaphilus , Orophosomci.

From the primary immigration routes of invertebrate and vertebrate faunal components in
the Central Himalayas, MARTENS (1984, 1993) distinguished Central Asian, West Asian
Himalayan, Tropical Indian, West Chinese Himalayan, and Indochinese Himalayan pathways
(Fig. 4). However, because of their preponderance in forests, only two major dispersal routes
are available to the Diplopoda, from Southeast Asia and the Indian subcontinent. Consequently,
the millipede fauna of the Central Himalayas is dominated by such tropical elements as the
families Sphaeropoeidae, Siphonophoridae, Andrognathidae, Harpagophoridae, Cambalopsidae,
Physobolidae, Cryptodesmidae, Opisotretidae, Fuhrmannodesmidae, Paradoxosomatidae,
Pyrgodesnudae, etc., which have Oriental and/or Indian affinities. In the relatively well-explored
Central Himalayas, most millipede species are restricted to tropical lowland forests such that it is
difficult to discriminate Southeast from Indian derivatives. Possibly only Sholaphilus,
Trichopeltis and certain Sphaeropoeidae hold eutropical Indian origins.

172

SERGEI I. GOLOV ATCH & JOCHEN MARTENS

West Asian

Fig. 4. — The main immigration routes of faunal components into the Nepal Himalayas.

Such genera as Hyleoglomeris, Tianella, Anaulaciulus, and Usbekodesmus are primarly
Palearctic and are restricted in the Central Himalayas to the uppermost forests, some even
spreading into alpine meadows above 4,000 m a.s.l., which contrasts with nearly 6,000 m for
certain other terrestrial invertebrates, e.g. spiders (MARTENS, 1993). The highest millipede
record in the Himalayas, and probably also in the world, is of Nepalmatoiulus ivanloebli
(ENGHOFF, 1987) encountered at 4,800 m a.s.l. Other millipedes demonstrate subtropical east
and southeast Asian elements, although there are occasional exceptions to the general rule,
“(sub)tropical creatures in (sub)tropical environments only”. Families that are 'more subtropical
than tropical include the Kashmireumatidae (with the oligotypic genus, Kashmireuma, in the
Himalayas and another monobasic genus in Vietnam), Megalotylidae (with the Oriental genus
Nepalella and a monobasic genus in the Russian Far East), Julidae (with Anaulaciulus and the
Oriental highly prolific genus Nepalmatoiulus), and certain Fuhrmannodesmidae and
Paradoxosomatidae [e.g., the endemic genera Hingstonia and Orophosoma (Fig. 3)].

The border between subtropical and purely tropical Himalayan components seems to be
vague (MARTENS, 1984, 1987b, 1993), as is that between forest (sub)zones (Fig. 2). Only a
few genera and even fewer tribes and families display clear vertical distribution patterns. In most
species there are only slight correlations with particular elevations. Some closely related genera
and species tend to occupy different altitudinal zones, probably because of niche segregation.
For example, Hingstonia and Sholaphilus (Fuhrmannodesmidae) tend to inhabit upper and
lower forests, respectively (Fig. 3), and this pattern is better demonstrated within speciose
genera. Usbekodesmus, for example, tends to be restricted to the upper forest belt, but a few
components are confined to low elevations, between 2,300 and 4,250 m a.s.l. (Table 1 , Fig. 3).

The same patterns have been reported for spiders, harvestmen, insects, birds, etc.
(Martens, 1984, 1987b, 1993), but the Diplopoda is distinguished in being almost strictly
sylvicolous and virtually entirely Oriental and/or Indian in origin.

The classical pattern of a prolonged cis-Himalayan band west of Brahmaputra, marking the
northwestemmost border of the Oriental realm, is highly characteristic of Oriental Diplopoda

DISTRIBUTION AND FAUNOGENESIS OF HIMALAYAN MILLIPEDES

173

(HOFFMAN & Burkhalter, 1978), and most Himalayan genera and tribes demonstrate this
pattern. An Oriental influence dominates in Kashmir, which is the classics, but also,
unexpectedly, to the north beyond the Indus Valley. Thus, judging from millipedes “...the less
elevated and more mild areas of modern North Pakistan seem to have retained particularly
ancient faunal elements as compared to the adjacent extremely high and severe Himalayas
nowadays supporting only relatively more advanced, younger forms” (GOLOV atch, 1991a:

FAUNOGENESIS

As the regional phyto- and paleogeography are well documented (WULFF, 1944; MEYEN,
1987), one can reasonably surmise that the Himalayas have served as a dispersal pathway for a
uniform Turgai biota (e.g. Quercus, Pyrus , Malus, and deciduous tree genera and associated
faunas) during the early and mid-Tertiary, which spread northwestward along the receding
southern coast of the Tethys Sea. A very considerable proportion of present-day European and
Mediterranean millipede genera, tribes, and families also seem to reflect repeated northwestward
dispersals from source areas in East and/or Southeast Asia, for example Hyleoglomeris and
possibly the Glomeridae as a whole, the tribes Brachyiulini and Leucogeorgiini (Julidae), the
tribe Paradoxosomatini and possibly all Paradoxosomatidae, and the genus Polydesmus and
possibly all Polydesmidae (GOLOV ATCH, 1987, 1991a, 1991b. 1993).

Beyond their effects on areas to the northwest, the Himalayas have also been a center of
secondary diversification since the Plio-Pleistocene, and much of the Central and West
Himalayas seem to have experienced a pronounced, secondary faunal impoverishment
(GOLOVATCH, 1991a), because all endemic millipedes display a relatively low taxonomic rank.
The development of local species swarms (among Tianella, Nepalella, Anaulaciulus,
Nepalmat&iulus, Gonoplectus, Hingstonia, Himalodesmus , Paranedyopus, etc.) through
allopatric speciation is a prominent characteristic of Himalayan Diplopoda irrespective of origin,
as has been observed among other soil/litter arthropods (MARTENS, 1987b, 1993). The few
anthropochorous introductions are very recent and have failed to alter the general zoogeographic
pattern of Himalayan Diplopoda.

Although preliminary, these reconstructions provide a basis for comparisons for future
faunistic and zoogeographic studies of the Himalayas. Compared to other terrestrial Arthropoda
(MARTENS, 1993), the salient aspects of Himalayan Diplopoda are pronounced sylvicoly and
Oriental and/or Indian origin, and their ostensible Palearctic influence also originates in present-
day subtropical regions of Southeast and East Asia.

REFERENCES

DOBRF.MEZ, J. F.. 1972. — Les grandes divisions phytogeographiques du Nepal et de I'Himalaya. Bull. Soc. boi. France,
119 : II 1-120.

Golovatch, S. I., 1987. — The alluaudi- group of Glomeris , another Macaronesian species swarm in millipedes
(Diplopoda: Glomeridae). Entomoi sc and. . 17 : 503-509.

Golovatch, S. I., 1990. — Diplopoda from the Nepal Himalayas. Several additional Polydesmidae and
Fuhrmannodesmidae (Polydesmida). Spixiana, 13 : 237-252.

Golovatch, S. I., 1991a. — On a small collection of millipedes (Diplopoda) from northern Pakistan and its
zoogeographic significance. Rev. suisse Zoo!.. 98 : 865-878.

Golovatch, S. I., 1991b. — The millipede family Polydesmidae in Southeast Asia, with notes on phylogeny
(Diplopoda: Polydesmida). Steenstrupia , 17 : 141-159.

Golovatch, S. 1., 1993. — On several new or poorly-known Oriental Paradoxosomatidae (Diplopoda. Polydesmida).
Arthropoda Selecta, 2 : 3-14.

Hoffman. R. L.. 1980. — Classification of the Diplopoda. Geneve. Mus. Hist, nat., (1979) 237 pp.

Hoffman, R. L. & Burkhalter, E. A.. 1978. — Studies on spirostreptoid millipeds XIV. A new species of Gonoplectus
from Thailand, with notes on the status and distribution of the genus (Spirostreptida: Harpagophoridae). J. nat.
Hist., 12 : 413-422.

174

SERGEI I. GOLOV ATCH &JOCHEN MARTENS

Kors6s, Z., 1996. — Another Himalayan group of julid millipedes: Towards the clarification of the genus Anaulaciulus
Pocock, 1895 (Diplopoda, Julida). Senckenberg. biol. (in press).

Martens. J., 1984. — Vertical distribution of Palaearctic and Oriental faunal components in the Nepal Himalayas.
Erdwissenschaftl. Forsch., 18 : 321-336.

Martens, J., 1987a. — Remarks on my Himalayan expeditions. Courier Forsch. -Inst. Senckenberg, 93 : 7-31.
Martens, J.. 1987b. — Beitriige zur Fauna, Faunengenese und Zoogeographie des Nepal-Himalaya. Arthropoda. Cour.
Forsch. -Inst. Senckenberg. 93 : 503 pp.

Martens. J.. 1993. — Bodenlebende Arthropoda im zentralen Himalaya: Bcstandsaufnahme, Wege zur Vielfalt und
okologische Nischen. Erdkundliches Wissen . 112 : 231-249.

MaSCLE, G., Delcaillau, B. & Herail, G.. 1990. — La formation de l'Himalaya. La Recherche, 217 : 30-39.

Meyen, S. V.. 1987. — Foundations of paleobotany. Moscow. “Nedra" Publrs, 403 pp. (In Russian].

Troll, C., 1967. — Die klimatische und vcgetationsgeographische Gliederung des Himalaya-Systems. Khumbu Himal,
1 -.353-388.

WULFF, E. V., 1944. — Historical geography of plants. The history of the world's floras. Moscow-Leningrad, Akad.
Nauk SSSR Publrs, 546 pp. (In Russian].

Source :

Etude systematique et ecologique des myriapodes dans
le Parc National de Chrea (Atlas blideen), Algerie

Ourida ABROUS-KHERBOUCHE

Universite des Sciences et de la Technologie Houari Boumedienne, I.S.N. Lab. d'ecologie animale

B.P. 32 El Alia Bab Ezzouar Alger, Algerie

RESUME

Les Myriapodes, notamment les diplopodes, jouent un role important dans la fragmentation de la litiere ct les
premieres etapcs du recyclage des mineraux dans le sol. Peu d etudes ont Ete consacrEes h ces groupes fonctionnels
d’arthropodes en Algerie. Pour cela, nous nous sommes proposes d’etudier les myriapodes (Diplopoda & Chilopoda) de
sept stations situees sur un gradient altitudinal dans le Parc National de Chrea (Atlas de Blida), plus particulierement sur
les djebels Chrea el Mouzaia, sEparEs par le protond ravin de l'oued Chiffa. Un premier resultat consiste en la liste
faunistique de toutes les especes rEcoltEes : 14 especes appartenant a 7 ordres differents sont repertoriees. L etude
autecologique des especes les plus abondanles permet de preciser leurs preferences biotiques el leur abondance relative au
sein de chacun des milieux qui sont largement decrits dans ce travail. L’etude comparative des peuplements montre que la
richesse specifique ainsi que 1'abondance relative augmentent generalement avec 1'altitude. L’une et l’autre sont plus
ElevEes sur le djebel ChrEa que sur le djebel Mouzaia. En outre, a 1'aide du coefficient ccenotique de Jaccard, nous
constatons l'existence de deux groupes distincts dans les stations d’Etude.

ABSTRACT

Systematic and ecological study of Myriapod communities in the Chrea National Park (Blida
Atlas, Alge ria).

Myriapoda - particularly Diplopoda - play an important role in litter-breakdown and soil mineralisation. Very few
studies have dealt with their ecological role in Algerian ecosystems. Hence it was decided to investigate millipede and
centipede populations in seven study sites ranging along an altitudinal gradient in the Chrea National Park (Blida Atlas),
precisely on Chrea djebel and Mouzaia djebel, which are divided by the deep canyon of oued Chiffa. The first result is a
check-list of all species collected at these sites: 14 species belonging to 7 different orders have been identified. The
ecological study of the most abundant species shows their biotic preferences and phenology. Following population
sampling, we can observe an increase in species richness and abundance related to altitude. Both are greater on Chrea
djebel than on Mouzaia djebel. Using the Jaccard coefficient, we recognize two distinct groups in the different study
sites.

INTRODUCTION

Dans toute etude ecologique, relative notamment a l’organisation des peuplements, la
systematique et la taxinomie sont de plus en plus indispensables ; sans elles, la comparaison des
biotopes et la connaissance precise de la structure des peuplements sont totalement impossibles
ou presentent un risque d’erreur d’ interpretation majeur.

Abrous - KHERBOUCHE, O., 1996. — Etude systematique et Ecologique des myriapodes dans le Parc National de
Chrea (Atlas blideen). Algerie. In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin. M.. (eds), Acta
Myriapodologica. Mem . Mus. natn. Hist . nat 169 : 175-186. Paris ISBN : 2-85653-502-X.

176

OURIDA ABROUS-KHERBOUCHE

Tres peu d'etudes a ce jour ont ete consacrees aux myriapodes dans les ecosytemes
naturels algeriens. Ces animaux y joucnt toutefois un role important, soit dans la fragmentation
de la litiere et le transfert ou le recyclage de la matiere minerale et organique (Diplopodes), soit
dans les processus de regulation des populations-proies de micro- ou de mesoarthropodes du sol
(Chilopodes).

C'est en 1840 qu'une premiere approche myriapodologique a ete entreprise par BRANDT
sur la region d' Alger, puis LUCAS (1846) a etudie une large partie de la faune myriapodologique
Nord Africaine.

Par la suite, BROLEMANN (1897, 1925, 1930, 1931) a approfondi les recherches
myriapodologiques du point de vue de leur taxinomie et de leur repartition au sein de diverses
entries ecologiques. En 1921, il signala 38 especes de diplopodes en Algerie, 23 en Tunisie, 4 au
Maroc et une espece en Lybie. Nous donnons dans ce travail la description des milieux d’etude
qui ont servi a une approche comparative de Porganisation des peuplements de chilopodes et de
diplopodes, dont les premiers resultats sont explodes et discutes.

MATERIEL ET METHODES

Dix pi£ges d' interception de type Barber ont «§te utilises dans chaque station pour capturer les myriapodes du Parc
National de Chrea. Ils ont ete releves mensuellemcnt durant la periode de Janvier 1989 a Septemhre 1990.

Du fait de problemes techniques, la periode de recolte n’a pas ete rigoureusement reguliere pour toutes les stations.
Pour les etudes comparatives, seule la periode de Juin 1989 a Juin 1990 a et 6 prise en consideration.

L’identification du materiel a etc rendue tres difficile a cause de 1'ancicnncte dc la litterature et par le fait que
plusieurs especes etaient inconnues ou de statut incertain. Nous avons benefici£ de l’aide du Dr. Serra, de I’Universite de
Barcelone (Espagne) et de J.-P. MAURiES,du Museum National d'Histoire Naturclle de Paris (France) respcctivement pour
la verification des especes de chilopodes et de diplopodes.

PRESENTATION DES MILIEUX

Le Parc National de Chrea est l'un des 9 pares nationaux que compte aujourd’hui
l'Algerie. D’une superficie de 26000 ha, il s'etend sur les versants Nord et Sud de 1' Atlas
Blideen, qui appartient a l'Atlas Tellien. Ce pare entoure le village de Chrea et englobe les djebel
s Guerroumen, Ferroukha et Mouzaia. Les deux premiers sont separes du dernier par le ravin de
l’oued Chiffa (Fig. 1).

D'apres HALIMI (1980), le climat du pare varie entre humide-doux sur les versants les plus
bas et humide-frais sur les versants plus eleves et les sommets. La formation vegetale la plus
caracteristique du Parc est la Cedraie, a partir de 1100 m sur le djebel Guerroumen. Le djebel
Ferroukha est couvert d'une foret degradee de Quercus ilex. En dessous de 1000-1 100 m, nous
rencontrons des forets de Finns halepensis, Quercus ilex ou Quercus faginea. Sur le djebel
Mouzaia, la Cedraie est absente. Elle est remplacee par des chenaies melangees de forets de Olea
europea. Pour etudier les myriapodes du Parc National de Chrea, sept stations situees sur un
gradient altitudinal ont ete choisies.

a) La station Roseaux

- Altitude et localisation : cette station est situee a 185 m d'altitude sur le djebel Mouzaia et
a quelques metres de l'oued Chiffa (Fig. 1 ).

- Exposition et etage bioclimatique : elle est a exposition Nord-Est et se localise dans
l'etage bioclimatique sub-humide a hiver legerement chaud (Fig. 2).

- Composition floristique et caracteristiques pedologiques : la station Roseaux se
caracterise par l'absence de strate arborescente. Elle presenle une strate arbustive assez dense
mais peu diversifiee. Phragmites australis et Laurus roseus sont les seules especes herbacees,
avec un recouvrement variant de 60 a 70%. Le sol de cette station est tapisse d'une litiere peu
epaisse, formee essentiellement d'herbes. Le pH est egal a 6,7 et l'humidite a 37%.

Source : MNHN, Paris

PEUPLEMENTS DE MYRIAPODES DE L' ATLAS BLIDEEN

177

b) La station Broussaille riveraine

Altitude et localisation : elle est
situee a 300 m sur le djebel Mouzaia,
le long de la route Nationale n°l, a cote
d'un petit affluent permanent de l'oued
Chiffa, a quelques dizaines de metres
de la vallee de l'oued Chiffa.

Exposition et etage bioclimatique :
la station Broussaille riveraine est a
exposition Nord-Est. Elle est localisee
dans l'etage bioclimatique sub-humide
et sous etage doux (Fig. 2).

Composition floristique et
caracteristiques pedologiques : la
vegetation est composee essentiellement
de trois strates :

1) La strate arborescente a un
recou vrement moyen de 40 a 45%, avec
Salix alba comme seul representant.

2) La strate arbustive comprend 9
especes et recouvre 30 a 40%. Ficus
carica, Olea europea et Rosa
semperviens sont les especes
abondantes.

3) La strate herbacee est tres
dense, constitute de plusieurs especes
et presente une litiere plus ou moins
compacte, composee d'herbes et de
quelques feuilles, les herbes atteignant
40 cm. Le sol de cette station presente
un pH = 6,7 et une humidite de 35%.

Fig. 1. — Localisation de 1’ Atlas blideen (A) et des stations (B).
Ro : Roseaux, Br : Broussaille riveraine, Su : Sub6raie. Pi :
Pinede, Ve : Verger, Ch : Chenaie, Ce : Cedraie.

FlG. 1. — Localization of the Blidean Atlas and of the study sites.
Ro: reeds. Br: Riverside brushwood, Su: Oak-wood (Q.
suber), Pi: Pine-wood, Ve: Orchard, Ch: Oak-wood (Q.
ilex), Ce: Cedar-wood.

c ) La station Pinede

Altitude et localisation : la Pinede
est situee a 400 m sur le djebel Chrea,
au Sud du village Fordjouna, le long de
l'oued Bouredou qui est un affluent de
l'oued Chiffa.

Exposition et etage bioclimatique : la station est exposee au Nord-Nord-Est ; elle est situee
dans l'etage bioclimatique sub-humide a hiver doux (Fig. 2).

Composition floristique et caracteristiques pedologiques : la vegetation est repartie sur trois
strates :

1) La strate arborescente : Pinus halepensis est l'espece dominante avec un recouvrement
de 30 a 40%. II peut atteindre une hauteur de 12 a 13 m.

2) La strate arbustive est dense et constitute de 9 especes. Pistachio lentiscus, l'espece
dominante, et Erica arbor ea. espece abondante, atteignent une hauteur de 12 m.

3) La strate herbacee, avec 1 1 especes, presente un recouvrement de 30 a 40%. Acanthus
maulus est l'espece la plus abondante. Le substrat est represente par un sol a pH = 6.6 ; il est
recouvert d’une litiere peu epaisse, formee essentiellement de feuilles de Pinus halepensis. La
presence de mousse est indicatrice d'une humidite du sol un peu plus forte, egale a 39%.

178

OURIDA ABROUS-KHERBOUCHE

Fig. 2. — Les etages bioclimatiques des stations etudids
dans 1’ Atlas blideen.

FiG. 2. — Bioclimatic stages of the study site in the Blidean
Allas.

d) La station Suberaie

Altitude et localisation : Elle est situee
a 450 m sur le djebel de Mouzaia, au Sud-
Ouest du lieu dit “Cafe maure” (Fig. 1).

Exposition et etage bioclimatique : la
Suberaie est a exposition Sud-Est et se
localise dans l'etage bioclimatique sub-
humide doux.

Composition floristique et
caracteristiques pedologiques :

1) la strate arborescente dominee par
Quercus suber , presente un recouvrement
inferieur a celui de la pinede.

2) la strate arbustive, avec ses sept
especes, presente un recouvrement de 70 a
80%. Pistachio lentiscus domine, Erica
arbor ea et Ole a europea sont abondants.

3) le recouvrement de la strate
herbacee est en moyenne de 50 a 60 %. Les
especes les plus frequentes sont Ranunculus
spicatus et diverses Gramines.

Durant la periode d’etude, cette station
presentait une vegetation seche. La litiere,
essentiellement composee de feuilles seches
de Quercus ilex, Pistachio lentiscus, de
branches et bourgeons morts ainsi que de
feuilles-epines de Calycotome spinosa,
presente une humidite du sol faible (20%)
par rapport aux autres stations. Le pH est en
revanche plus eleve que dans les stations
precedentes (7,1).

e) La station Verger

Altitude et localisation : cette station est situee a 1000 m sur le djebel Chrea a quelques
centaines de metres du village Fourdjouna.

Exposition et etage bioclimatique : elle est exposee au Sud-Sud-Ouest et se situe dans
l'etage bioclimatique humide et sous-etage frais.

Composition floristique et caracteristiques pedologiques : la vegetation de cette station
comporte trois strates :

1) la strate arborescente : le recouvrement faible varie entre 10 et 20%. Cerasus avium est
la seule espece abondante, sa hauteur ne depasse pas 4 m.

2) la strate arbustive comprend quelques especes rares. Erica arborea, frequente, ne
depasse pas 1,5 m. Le recouvrement est de 50 a 60%.

3) la strate herbacee : elle est dense avec beaucoup d'especes qui recouvrent 90 a 95%. Le
sol, a pH = 6,7 et humidite = 33%, est recouvert d'une litiere epaisse de 3 a 4 cm, composee de
feuilles de Cerasus avium, Pyrus communis, Malus mitis ainsi que d'herbes et de quelques
branches.

Source :

PEUPLEMENTS DF. MYRIAPODES DE L' ATLAS BLIDEEN

179

f) La station Chenaie

Altitude et localisation : elle se trouve a 1400 m sur le djebel Chrea (Mont Djamaa Draa) a
1400 m d'altitude, dans la foret de Beni Salah.

Exposition et etage bioclimatique : la Chenaie, a exposition Sud-Ouest, est situee dans
l'etage bioclimatique humide a hiver frais.

Composition floristique et caracteristiques pedologiques : la vegetation de la station est
composee des strates suivantes :

1 ) la strate arborescente : le recouvrement est de 20 a 30% avec Quercus ilex comme seul
representant.

2) la strate arbustive : avec trois especes recouvrant 90 a 95%. Quercus ilex est dominant.

3) la strate herbacee : le recouvrement est de 80%. Calycotome spinosa est l'espece
dominante. Le sol rocheux de cette station, constitue de schiste, est tapisse par une litiere de 3 a
4 cm d'epaisseur, formee essentiellement par des feuilles de chene-vert ( Quercus ilex) et
quelques feuilles de Cist us monspeliensis. Le pH est identique a celui de la station Verger (6,7)
mais l'humidite est plus faible (33%).

g) La station Cedraie

Altitude et localisation : elle est situee a 1600 m dans la foret de Beni Salah a 29 m du pic
dit “Koudia Sidi Abdelkader” (1629 m).

Exposition et etage bioclimatique : La station Cedraie est a exposition Sud-Ouest et est
situee dans l'etage bioclimatique humide et sous etage frais (Fig. 2).

Composition floristique et caracteristiques pedologiques : cette station presente une
formation arborescente assez dense, formee uniquement de cedres ( Cedrus atlantica), d’une
hauteur moyenne de 5 a 18 m. Nous y retrouvons les trois strates vegetales :

1) la strate arborescente : la hauteur et la densite de ces arbres ( Cedrus atlantica) empechent
les rayons du soleil d'atteindre le sol : les autres strates presentent un recouvrement moins
important.

2) la strate arbustive : Cedrus atlantica est la seule espece rare avec un recouvrement de 20
a 30%.

3) la strate herbacee : contient un nombre d'especes plus eleve que les autres stations ;
parmi les 17 especes relevees, Chrysantemum segetum, Graminea sp. et Ranunculus spicatus
sont les plus frequentes.

Le sol de cette station est recouvert d'une litiere de 2 a 3 cm et plus dans certains endroits.
Elle est composee essentiellement d'aiguilles de cedre, de branches et de nombreux rameaux
morts. Le sol a un pH neutre (6,7) et son humidite est egale a 33%.

RESULTATS ET DISCUSSION

Dans le Parc National de Chrea, nous avons recolte 14 especes appartenant a 7 ordres
differents (Tableau 1 ). Le nombre d'especes dans les ordres Geophilomorpha et Lithobiomorpha
et le nombre d'individus des especes de Julida (Iuliformia) et Polydesmida sont les plus eleves.

Le Tableau 2 montre que la richesse specifique des diplopodes est ties faible dans les deux
premieres stations (Roseaux, 185 m et Broussaille riveraine, 300 m d'altitude). En revanche, ils
sont plus abondants dans les stations de haute altitude (Verger, Chenaie et Cedraie).

Les Chilopodes se rencontrent dans la majorite des stations. Roseaux, Pinede et Suberaie
presentent toutefois un faible effectif.

Sur le diagramme (Fig. 3), nous remarquons l'absence de l'ordre Scutigeromorpha dans la
station Roseaux (185 m) situee a quelques metres de l'oued Chiffa, et qui semble etre un milieu
de favorable. L'ordre Scolopendromorpha n'est present que dans la Chenaie (1400 m) ; il semble
preferer les stations homogenes avec une litiere epaisse, formee de feuilles seches, de branches
et de bourgeons morts.

180

OURIDA ABROUS-KHERBOUCHE

Tableau 1. _ Nombrc d'especes et d'indi vidus captures, par ordre, dans le Parc National de Chr6a (Atlas blidSen).

Table 1. — Number of species and individuals collected per order in the Parc National de Chrea (Blidean Atlas).

Classes

Ordres

Nbre d'especes

Nbre d'individus

Julida

329

DIPLOPODA

Polydesmida

21 1

Glomerida

Geophilomorpha

CHILOPODA

Lithobiomorpha

Scolopendromorpha

Scutigeromorpha

Total

778

Tableau 2. — Liste des especes de diplopodes et chilopodes captures et leur abondance dans les stations etudi£es. Ro :
Roseaux, Br : Broussaille riveraine, Su : Suberaie. Pi : Pinede, Ve : Verger, Ch : Chenaie, Ce : CSdraie. Tot =
nombre total d’individus.

TABLE 2. — Check- list of millipedes and centipedes, relative abundance in the study sites. Ro: reeds, Br: Riverside
brushwood, Su: Oak-wood (Q. suberj, Pi: Pine-wood, Ve: Orchard, Ch: Oak-wood (Q. ilex! Ce: Cedar-wood. Tot =
total number of individuals.

Ordres

Especes

Tot

Julida

Ommatoiulus gauthieri

1 1

Phalloiulus distinct us

182

244

Glomerida

Glomeris conspersa

1 1

Polydesmus superus

Polydesmida

Archipolvdesmus sp.

201

208

Geophilus carpophagus

Geophilus sp.

Geophilomorpha

Schendvla sp.

Lithobius crassipes

Lithobius castaneus

Lithobiomorpha

Lithobius sp.

Lithobius forficatus

Scolopendromorpha

Scolopendra sp.

Scutigeromorpha

Scutigera coleoptrata

L'etude synecologique montre que la richesse specifique ainsi que le nombre d'individus
augmentent generalement avec l'altitude (Fig. 4). La Suberaie (450 m) presente le nombre
d'especes le plus faible. Ceci peut etre du a la faible humidite du sol (20,40) de cette station.

Le dendrogramme obtenu a l'aide du coefficient ccenotique de JACCARD revele l'existence
de deux groupes distincts de stations ayant une similarite de 12,50 % (Fig. 5).

La station Verger (1000 m) et la Chenaie (1400 m) presentent la similarite la plus elevee
(69,85%), suivies par la Cedraie (1600 m), avec laquelle elles forment un premier groupe. Ce
sont des stations relativement humides situees sur le meme djebel Chrea et a exposition sud-
ouest.

Le second groupe rassemble quatre stations : la Suberaie, la Pinede, la Broussaille
riveraine et la station Roseaux.

La Suberaie et la Pinede, situees respectivement a 450 m et 400 m d'altitude, sont des
stations sub-humides, homogenes, qui presentent la similarite la plus elevee (43,60%). Elles
offrent des conditions climatiques tres voisines et, par consequent, des microclimats similaires.

Source . MNHN, Paris

PEUPLEMENTS DE MYRIAPODES DE L’ ATLAS BLIDEEN

181

Juliformia

Glomerida

Polydesmida

Geophilomorpha

Lithobiomorpha

Scolopendromorpha

Scutigeromorpha

0 1 km

i - 1

Fig. 3. — Diagramme representant les proportions du nombre d’especes pour les differents ordres dans les stations
etudiees. Ro : Roseaux, Br : Broussaille riveraine, Su : Suberaie, Pi : Pinede, Ve : Verger, Ch : Chenaie, Ce :
CSdraie.

FlG. 3. — Relative importance of the specific richness for the different orders in the study sites. Ro: Reeds. Br: Riverside
brushwood. Su: Oak-wood (Q. suberj, Pi: Pine-wood, Ve: Orchard, Ch: Oak-wood ( Q. ilex A Ce: Cedar-wood.

2 especes
i - 1 50 individus

0 1 km

I - 1

Fig . 4. — Nombre d’especes et d’individus captures par station dans le Parc National de Chrea (Atlas blideen).
Fig. 4. — Number of species and individuals in each site in the Parc National de Chrea.

182

OURIDA ABROUS-KHERBOUCHE

100

Verger (1 000 m)

Chenaie (1 400 m)

Cedraie (1 600 m)

Suberaie (450 m)

Pinede (400 m;

Roseaux (185 m)

Broussaille riveraine (300 m)

A ces deux stations s'ajoute la station
Roseaux (185 m) et enfin, la Broussaille
riveraine (300 m) avec une similarity de
32,9 %.

L'etude autoecologique de 3 especes
dont l’abondance relative est importante
permet d'indiquer leurs preferences
ecologiques et leurs phenologies :

Ommatoiulus gauthieri (Brolemann,
1931 )

Cette espece a ete decrite du djebel
Bouzegza par BROLEMANN (1931). Elle
presente une tres grande variability
moiphologique. Dans 1' Atlas blideen, nous
l'avons recoltee dans presque tous les sites
d’etude. Elle est absente seulement dans les
stations Roseaux et Broussaille. Le plus
grand nombre d'individus (62) a ete recolte
dans la station Verger (1000 m). Cette
espece semble preferer une vegetation
dense avec une litiere composee de feuilles
et d'herbes.

Les males ont ete recoltes durant l'ete, l'automne et l'hiver. Ils montrent une activite
intense a la fin de l'automne et au debut de l'hiver (Fig. 6). Les femelles sont plus actives au
debut de l'automne et se rencontrent elles aussi, durant l'ete, l'automne et l'hiver.

Cylindroiulus (= Phalloiulus) distinctus (Lucas, 1846)

Cet iulide a ete signale dans les bois de Boulogne pres d' Alger par BROLEMANN en 1931 ;
a l'exception de la station Roseaux ( 1 85 m), elle se retrouve dans toutes les autres stations. Le
plus grand effectif (182 individus) a ete trouve dans la Cedraie, ce qui laisse supposer une
preference pour les hautes altitudes, avec une litiere composee essentiellement d'aiguilles de
Cedrus atlantica. Les individus de cette espece ont ete recoltes en toute saison (Fig. 7) mais, le
plus grand nombre a ete capture a la fin de l'automne et au debut de l'hiver.

Archipolydesmus sp.

Cette espece qui, d'apres J.-P. MAURIES (Museum N. H. N., Paris) est polymorphe,
nouvelle et appartient a un genre inedit en Algerie, a ete recoltee dans la Pinede, la Chenaie, la
Cedraie et la station Broussaille, elle est absente dans les autres sites. Toutefois, la ou elle est
presente, l'effectif demeure faible. C’est dans la Cedraie (1600 m) que Ton observe l’effectif le
plus eleve (201 individus). Ceci montre que cette espece prefere les hautes altitudes avec une
humidite atmospherique relativement grande. Comme le montre la Figure 8, les males et les
femelles de cette espece ont ete captures de l'automne jusqu'a la fin du printemps. Ils sont tres
actifs pendant l'automne et le debut de l'hiver.

Fig. 5. — Dendogramme de similitudes des stations Studies dans
FAtlas blidden.

FlG. 5. — Similarities between the study sites.

CONCLUSION

Cette etude preliminaire relative aux myriapodes du Parc National de Chrea apporte une
contribution a une meilleure connaissance de la faune algerienne, en reunissant les donnees
initiales d'une approche de l’organisation des peuplements de chilopodes et de diplopodes au
sein de plusieurs types d’ecosystemes nord-africains typiques.

Bien que demeurant faibles en general, le nombre d'especes ainsi que le nombre
d'individus augmentent avec l’altitude : la station culminante (Cedraie, 1600 m) presente la plus
grande richesse specifique et la plus grande abondance ( 10 especes et 440 individus captures).

Source :

PEUPLEMENTS DE MYRIAPODES DE L’ ATLAS BLIDEEN

183

' Pinede

"Chenaie

—

' . on . 1

' . 1

i i i i i i i i i i i

J FMAMJ J ASOND

J FMAMJ JAS

XXXXXJ JASONDJ FMAMJ XXX

"Suberaie

[ Cedraie

- —

~ . Hill,

* * « » * * » • n

1 . 11 fl ■

— i — i — l-i i i i i i i i

J FMAMJ JASONDJ FMAMJ JAS XXXXXJJASONOJFMAMJXXX

Fig. 6. — Abondance et activity relatives mensuelles de Ommatoiulus gauthieri dans les stations etudiees (x = periode
non echantillonnee).

FlG. 6. — Relative abundance and activity of Ommatoiulus gauthieri in the study sites (x = no sampling).

Source : MNHN. Paris

184

OURIDA ABROUS-KHERBOUCHE

Broussaille

— jO — 1 — 1 — 1 1 l-

XFMAMJ J ASONDJ FMAMJ J AS

Pinede

30
25
20
15
10
5
0

J FMAMJ J ASONDJ FMAMJ JAS

Verger

i i i i ■ i i i i—i — i — i — i— j — i — i — i—i — i — »—

XXXXXJ J ASONDJ FMAMJ JAS

J FMAMJ J ASOND J FMAMJ J AS

Fig. 7. — Abondance et activite relatives mensuelles de Phalloiulus distinctus dans les stations ctudiees (x = pSriode non
echantillonnee).

FlG. 7. — Relative abundance and activity of Phalloiulus distinctus in the study sites (x = no sampling).

Source : MNHN, Paris

PEUPLEMENTS DE MYRIAPODES DE L’ ATLAS BL1DEEN

185

6 □ $

~ Broussaille

Cedraie

- 50

- 40

—1 _ 1 _ 1 1 _ 1 _ 1 — 1 - 1 — 1 — 1 — 1 — 1 — Lxd - 1 - 1 - 1 - 1 - 1 - 1 - 1 - 1 Q

—J 1_ 1 — 1 — 1 — 1 — 1 — 1 — 1 — 1 — 1

i4i , , x ,

XXFMAMJ J ASONDJ FMAMJ JAS X X X X X X J J A S O N D J F M A M J X X X

70
60

50 r

40
30
20
10
0

Pinede

J I - 1 - 1 - L

J - L

Chenaie

J FMAMJ J ASONDJ FMAMJXXX X X X X X X J J A S O N D J F M A M J X X X

Fig. 8. — Abondance ct activite relatives mensueiles de Archipolydesmus sp. dans les stations etudiees (x = periode non
6chantillonnee).

FlG. 8. — Relative abundance and activity of Archipolydesmus sp. in the study sites (x = no sampling).

La comparaison des listes faunistiques des diverses stations fait apparaitre deux points :

1 ) la repartition plus ou moins uniforme de la plupart des genres et especes au sein de l'aire
concemee par 1’ etude.

2) l’apparente diversite des habitats dans lesquels une meme espece peut etre representee.

Les populations les plus abondantes montrent une periode d'activite pendant l'automne,

l'hiver et le printemps ; les males sont plus actifs en surface en hiver et les femelles au
printemps, les conditions climatiques qui regnent durant I’ete rendant difficiles l’acces aux
individus installes temporairement dans des sites refuges. L'altitude, associee a 1’ importance
relative de l’humidite, semble representer un facteur ecologique fondamental dans la repartition
des especes de la region etudiee. Au-dela de cette premiere approche consacree aux chilopodes et
aux diplopodes, d'autres etudes sont envisagees pour determiner la nature et l’importance des
divers facteurs expliquant 1’ organisation et les variations de leurs peuplements.

186

OURIDA ABROUS-KHERBOUCHE

REMERCIEMENTS

Je tiens & remercier vivement le Dr. R. Bosmans pour son aide dans la r^colte du materiel et ses conseils precieux.
Je remercie Sgalement le Dr. A. Serra pour la verification de 1’ identification des chilopodes et M. J.-P. MAURlfcs pour la
verification de 1* identification des diplopodes.

REFERENCES

Brandt, C.,1840. — Rapport sur les Oniscides et les Myriapodes dans la regence d'Alger. Revue Soc. cuvierienne
Paris , 3.

Brolemann, H. W., 1897. — Iulides d'Alg6rie. Ann. Sc. nat. , ser . 8, Zool., 4 : 253-276.

Brolemann, H. W.. 1921. — Liste des Myriapodes signals dans le nord de l'Afrique. Bull. Soc. Sc. nat. Maroc, 1 : 3-6.

Brolemann, H. W., 1925. — Races nouvelles de Schizophyllum alg£riens (Myriapodes Diplopodes). Bull. Soc. Hist,
nat. Afrique du Nord , 16 : 245-253.

Brolemann, H. W., 1930. — Myriapodes du Sahara central recueillis par L. G. Seurat au cours de la mission du Hoggar
(fevrier-avril 1928). Bull. Soc. Hist. nat. Alger, 1 : 6-8.

Brolemann, H. W.. 1931. — Myriapodes recueillis par Mr. le Dr. H. Gauthier en Alg6rie. Bull. Soc. Hist. nat. de
l'Afrique du Nord, 22 : 121-134.

Halimi, A., 1980. — Atlas blideen. Off. Pub. Univ. Alger : 1-523.

Lucas, L., 1846. — Notes sur quelques nouvelles espfcces d'inscctes (Myriapodes) du Nord de l'Afrique. Rev. zool. Sci.
de Cuvier , Paris, 9 : 283.

Source : MNHN, Paris

Etude des communautes de myriapodes
(Chilopoda et Diplopoda) des forets prepyreneennes

(Huesca, Espagne)

Antoni SERRA *, Maria Cristina VICENTE ** & Eduardo MATEOS *

* Departament de Biologia Animal. Facultat de Biologia, Universitat de Barcelona, Avda. Diagonal, 645

E-08028 Barcelona, Espagne

** Departament de Biologia Animal, Biologia Vegetal i Ecologia, Facultat de Ciencias, Universitat Autonoma de

Barcelona, E-08193 Bellaterra, Barcelona, Espagne

RESUME

Ce travail est consacre & lfetude des chilopodes et des diplopodes des milieux forestiers du massif de San Juan de la
Pena, qui appartient au sysfeme prepyreneen de la province de Huesca (Nord de 1’Espagne). Douze stations ont ete
etudiees : trois pinedes, deux sapinieres, une hetraie, une foret mixte, une chenaie, une foret de chenes verts, deux zones
de broussailles et une prairie paturee. On a utilise la methode des pfeges d’interception de type Barber. Six pfeges,
distants d'au moins 10 m, ont ete installes dans chaque station et releves chaque semaine, de fevrier 1977 a mars 1978.
Les aspects suivants ont ete pris en consideration : composition taxinomique des peuplements, densite relative et
aclivite des chilopodes et des diplopodes. caracterisation de chaque station en fonction de son peuplement en myriapodes
et preferences specifiques pour les differents habitats.

ABSTRACT

Centipede and millipede population study in prepyrenean forests (Huesca, Spain).

This work is devoted to the study of the centipedes and millipedes living in the forest habitats of the San Juan de la
Pena mountains, which belongs to the pre-Pyrenean system of the Huesca province (North of Spain). Twelve plots were
studied, corresponding to three pine groves, two fir woods, a beechwood, a mixed forest, an oak grove, a holm-oak
wood, two brushwoods and a pasture land. The sampling was performed by means of pitfall traps. Six traps, at least 10 m
apart, were placed in each plot. The trapping was done weekly from February 1977 to March 1978. The following aspects
were investigated : taxonomic composition of the myriapod populations, relative densities of active Chilopoda and
Diplopoda during the study period, characterization of each plot on the basis of its myriapod population and specific
preferences for the different habitats studied.

INTRODUCTION

Dans ce travail est exposee letude realisee sur un grand echantillonnage de myriapodes
(chilopodes et diplopodes) recoltes par des methodes indirectes sur differents biotopes du Massif
de San Juan de la Pena, qui est situe dans la zone prepyreneenne de la province de Huesca
(Espagne).

Serra, A., Vicente, M. C. & Mateos. E., 1996. — Etude des communautes de myriapodes (Chilopoda et
Diplopoda) des forets prepyreneennes (Huesca, Espagne). In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy -
Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. nain. Hist. nat.. 169 : 187-204. Paris ISBN : 2-85653-502-X.

188

ANTONI SERRA. MARIA CRISTINA VICENTE & EDUARDO MATEOS

Le programme de piegeage a ete structure et execute par le Dr. Cesar PEDROCHI-RENAULT
de l'lnstituto Pirenaico de Ecologia de Jaca. 52059 exemplaires de differents groupes
d'arthropodes ont ete captures. Le groupe des arachnides est, en nombre, le mieux represente,
tandis que celui des chilopodes presente le plus faible effectif de tout l'ensemble (PEDROCCHI-
RENAULT, 1985).

SITE D’ETUDE

Le massif de San Juan de la Pena fait partie des Chaines PrepynSneennes meridionals ; il se trouve situe dans la
province de Huesca, au sud-est de Jack. II est esscntiellement constitue, du point de vuc geologique, de conglomerats
deltaiques qui reposent sur les marges de la Depression Moyenne, et atteint, k son point culminant, le pic dc San
Salvador, Taltitude de 1546 m. Le climat de la zone est de type subm6diterraneen continental, les zones basale el
peripherique du massif se trouvant dans la subregion phytoclimatique IV- VI (Allu£, 1964). L'examen des diagrammes
climatiques correspondant aux differentes stations du contour du massif dc San Juan de la Pena permet de noter les
constantes suivantes (Pedrocchi-Renault, 1985) :

- Les precipitations annuelles oscillent entre 700 et 900 mm, avec un maximum printanier en mai (pres de
100 mm) et un minimum en juillet (pres de 35 mm). La periode seche est peu intense et de courte duree (mois de juillet
seulement).

- La temperature annuelle moyenne oscille autour de 10°C.

- La moyenne des temperatures minimales du mois le plus froid (janvier ou decembre) oscille entre -1°C et -2°C,
avec des minimales absolues de -13°C.

- La moyenne des temperatures maximales du mois le plus chaud (juillet ou aoul) oscille entre 26°C et 28°C, avec
des maximales absolues qui atteignent 36°C.

- Dans presque toutes les stations, la periode des gelees sures s’etend de decembre k mars; celle des gelees
probables est en avril, octobre et novembre. Les mois de mai k septembre peuvent etre consideres comme en dehors des
p6riodes de gelees.

- Dans le centre du massif, k une altitude de 1200 m, on observe une modification importante des conditions
climatiques. La periode seche estivale disparait et le mois de mai presente un caractere pcrhumide, avec des precipitations
superieures a 100 mm. Les temperatures maxi males dete ne sont pas si elevees que dans la zone basale et la periode des
gelees probables se prolongue jusqu'en mai. Des telles caracteristiques permettent d'inclure le massif dans la subregion
phytoclimatique VI, qui est definie par 1'absence de periodes seches et par une longue saison froide, avec une temperature
moyenne du mois le plus froid inferieure a 6°C.

LES STATIONS D'ETUDE

Le programme de piegeage a ete realist sur un total de douze stations situees, a l’exception de la chenaie a Quercus
fagi/tea, sur le massif de San Juan de la Pena.

Hetraie

Altitude : 1290 m ; Orientation : N-NE ; Pente : 33°. Sol ires profond, riche en carbonates, couvert par une epaisse
couche de litiere qui rend difficile le developpement de la strate muscinale. La strate hcrbac6e est tres pauvre et
discontinue. La strate arbustive est un peu dense, formee par des buis et quelques houx. Quelques sapins et pins sylvestres
accompagnent les hetres dominants.

Foret mixte

Altitude : 1105 m ; Orientation : N-NW ; Pente : 20°. Le sol est une rendzine tres humifere, avec un horizon
organique atteignant 20 cm d'epaisseur. La couche de litiere ne permet pas le developpement de la couche muscinale. La
strate herbacee est dominee par Hedera helix qui forme un tapis et meme, ga el la. des masses de v£g6tation. La strate
arbustive est assez importante en abondance et diversite specifiques. La strate arboree, avec un recouvrement de 100%,
comprend les essences Tilia platyphyllos, Frcixinus excelsior, Ulmus montana, Fagus sylvatica et Abies alba.

Pinede basse

Altitude : 962 m ; Orientation : N-NE ; Pente : 17°. Sol profond de tcrre brune, maintenu sur un terrain caillouteux
et couvert par une strate muscinale continue. Le buis, bien developpe et dense, constitue I'espece arbustive dominante,
avec aussi Clematis vitalba et quelques jeunes sapins. Pinus sylvestris forme une voute arboree presque continue.

Pinede moyenne

Altitude : 1120 m ; Orientation : NE ; Pente : 32°. Sol brun, calcaire, profond et couvert d un strate muscinale
dense et abondante. La couche herbacee est abondante si on la compare avec celle d'autres milieux forestiers. La strate

Source : MNHN. Paris

MYRIAPODES DES FORETS PRE-PYRENEENNES

189

arbustive, avec un epais manteau de buis, contieni aussi quelques exemplaires de Juniperus communis et de hetres. La foret
est constitute de Pinus. sylvestris assez murs, tres frequents sur le massif.

Pinede haute

Altitude : 1390 m ; Orientation : W ; Pcnte : 12°. Sol profond avec dcs carbonates, couvert de feuilles de hetre. Les
strates muscinale et herbacee sont pauvrcs. Le niveau arbustif est peu dense et il est domine par Buxus sempervirens et
Ilex aquifolium. La loret, eclaircie par la coupe de Pinus sylvestris, comprend des hetres et des sapins.

Sapiniere basse

Altitude : 1035 m ; Orientation : N ; Pente : 29°. Sol acide superficiellement, tres riche en matiere organique, se
melangeant en profondeur avec des cailloux et de 1’argile. La strate arbustive est compose par des buis et des hetres
jeunes. La strate arboree se compose de sapins, b 100%, sans aucune autre espece d’arbre.

Sapiniere haute

Altitude : 1415 m ; Orientation : N-NW ; Pente : 25°. Sur la roche-mere, constitute de conglomerats, le sol est
profond, riche en humus et de pH acide. Strate muscinale abondante, avec un recouvrement de 50%. Les strates herbacee et
arbustive sont trts pauvres. La strate arboree est typique dune foret en regeneration, avec abondance de Abies alba jeunes
mais peu vigoureux ou mourants, avec beaucoup de bois mort au sol (exemplaires morts et restes de coupe). De rares Pinus
sylvestris s'intercalent entre les sapins.

Chenaie d'ye uses

Altitude : 940 m ; Orientation : S-SE ; Pente : 27°. Sol de type xerorendzine, peu profond et riche en carbonate de
calcium. Meme si actuellement la pression humaine est nullc, jusqu'aux..annees 1940, le paturage et la coupe des arbres
ont diminue la taille de la chenaie qui se reduit actuellement a quelques massifs d’yeuses (chene vert). En sous-bois,
I'absence de lumiere ne permet pas I’installation d’autres especes, et une <§paisse couche de feuilles (10 a 15 cm) s’est
accumulee. Autour de ces massifs apparait une bordure arbustive fondamentalement constituee de Buxus sempervirens et
Juniperus. Dans les espaces ou verts apparaissent des plantes qui resistent a des secheresses prolongees et a des
oscillations thermiques importantes, tel les que Thymus vulgaris, Festuca greca indigesta et quelques Sedum. Cette station
est la plus mediterraneenne du massif.

Chenaie a Quercus faginca

Altitude : 775 m ; Orientation : N-NW ; Pente : 10°. C’est la seule station qui n'est pas situee sur le massif de San
Juan de la Pena, mais sur un de ses chainons. C'est une foret de regeneration de Quercus faginea avec une vegetation
caracteristique du type submediterraneen montagnard, ou Taction du paturage a ete tres intense dans le passe. Le sol est
une lerre brune calcaire, en bon etat de conservation.

Fruticee a Echinospartum horridum

Altitude : 1272 m ; Orientation : S-SE ; Pente : 14°. Cette station est etablie sur une zone qui a brule en 1919 et qui
a ete repeuplee avec Pinus sylvestris en 1965. Aprfcs Tincendie. lerosion a laisse un sol squelettique sur le conglomerat
qui affleure souvent. Les aptitudes colonisatrices de la fruticee & Echinospartum horridum lui ont permis d'occupcr ces
aires alterees par le feu, avec le maintien de la communaute anterieure h Tincendie. Les pins, peu vigoureux et ne
depassant pas un metre de haut, souffrent actuellement de la s£cheresse et aussi des attaques de chenilles processionnaires.

Fruticee d Genista scorpius

Altitude : 840 m : Orientation : S ; Pente : 33°. Cette station est situee dans une aire tres perturbee par Thomme ;
1’abandon du paturage a 6te la cause probable de Tinvasion de Genista scorpius , espece qui domine actuellement. Le sol,
arase, contieni peu de matiere organique en surface ou affleure souvent la structure argileuse de Thorizon mineral. Outre les
genets, la vegetation, caracteristique des zones arides a contrastes thermiques, est composee principalement de Thymus,
Lavandula et Festuca gr. indigesta .

Paturage

Altitude : 1 130 m ; Orientation : N-NE et N-W ; Pente : 10°. II s’agit de prairies qui ont appartenu h la foret et dont
Tequilibre depend uniquement de la pression du paturage ; elles ont et<5 colonisees rapidement par la vegetation marginale
forestiere ou par la frutic6ee. Le sol est profond (60 & 100 cm), de terre brune et avec une notable quantite de matiere
organique en surface. La vegetation est toujours herbacee, avec en general des graminees. La zone choisie pour
Techantillonnagc a perdu son equilibre h la suite du manque de paturage. Actuellement, la fruticee a Echinospartum
horridum s’accroil chaque ann6e. alors qu’ apparaissent de jeunes Pinus sylvestris.

190

ANTONI SERRA. MARIA CRISTINA VICENTE & EDUARDO MATEOS

MATERIEL ET METHODES

Le materiel etudiiS a etc collecte a I'aide de pieges de type Barber de 57 mm de diametre a l'ouverture ; une solution
saturee de chlorure de sodium dans l'eau avec un peu de detergent incolore a ete utilisee comme liquide de conservation.
Dans chacune des douze stations d'echantillonage, six pifeges ont ete installes a plus de dix metres de distance les uns des
autres. Le nombre de pieges a ete determine par un echantillonnage fait avant selon les criteres de Lamotte el at. (1969).
Le materiel capture <Stait rdcupere chaque semaine, si les conditions climatiques le permettaient. Le piegeage a die realise
pendant la periode de temps s’etendant de fevrier 1977 S mars 1978.

L'6chantillon global de niyriapodes est compost de 3760 exemplaires correspondant h 3670 diplopodes et 90
chilopodes. L'ensemble de ces deux groupes represente a peu prbs 7.5% du total des arthropodes recoltds (52059
exemplaires).

On ne veut pas clore ce paragraphe sans rappeler que I’interpretation des donnees obtenues avec cette methode de
capture par piege est d’une fiabilite limitde et qu'elle concerne essentiellement les taux d'activitd des individus
composants une fraction des populations, il faut done en tenir compte au moment des conclusions (ADIS, 1979; Ascaso,
1984).

REMARQUES FAUNISTIQUES

Le nombre d'especes de diplopodes repertoriees dans l'ensemble des douze biotopes est de
17. Une dizaine d’entre elles sont pyreneennes : Hirudisoma pyrenaeum Ribaut, 1908 -
Marquetiella lunatum (Ribaut, 1920) - Archipolydesmus osellai Ceuca, 1968 - Polydesmus
racovitzai Brblemann, 1910 - Blaniulus dollfusi Brblemann, 1894 - Haplopodoiulus spathifer
(Brblemann, 1897) - Leptoiulus umbratilis Ribaut, 1905 - Ommatoiulus robustus Ceuca, 1974 -
Loboglomeris haasi Attems,1927 et Protoglomeris vasconica (Brblemann, 1897). Le reste des
especes, Polydesmus coriaceus coriaceus Porat, 1879. Cylindroiulus caeruleocinctus (Wood,
1864), Ommatoiulus sabulosus (Linne, 1758), Tachypodoiulus niger (Leach. 1815) (= T.
albipes), Glomeris hexasticha intermedia Latzel, 1884 et Glomeris marginata (Villers, 1789),
sont des especes de distribution europeenne plus ou moins large. Avec ces especes, trois
exemplaires de Ceratosphys sp. ont aussi ete trouves mais ils n'ont pas pu etre identifies
specifiquement car il s'agissait d'une femelle et de deux males immatures.

A noter que les especes H. pyrenaeum, M. lunatum, P. racovitzai et L. umbratilis ont ete
trouvees pour la premiere fois dans la Peninsule Iberique. D'autre part, A. osellai est retrouvee
pour la premiere fois depuis sa description originale. Tous ces faits mettent en evidence qu'il
reste encore beaucoup de donnees faunistiques a decouvrir sur le versant sud des Pyrenees,
meme si cette zone est une des mieux connues de la Peninsule Iberique (MAURIES, 1975).

En ce qui concerne les chilopodes (cf. BROLEMANN, 1930 ; EASON, 1964), on remarque
que les individus appartenant aux 1 1 especes identifies au cours du piegeage sont beaucoup
moins nombreux que pour les diplopodes. Deux des especes, Strigamia acuminata (Leach,
1814) et Lithobius foificatus (Linne, 1758) se caracterisent par une repartition holarctique ; cinq
autres especes, Lithobius borealis Meinert, 1872, Lithobius calcaratus C. Koch, 1844, Lithobius
dubosequi Brblemann, 1896 (= L. microps), Lithobius piceus L. Koch, 1862 et Lithobius
tricuspis Meinert, 1872 presentent differents types de repartition en Europe ; les quatre autres
presentent d’ autres types de distribution geographique : Lithobius castaneus Newport, 1844 est
une espece circummediterraneenne ; Lithobius pilicornis Newport, 1844 est un element
atlantique ; Lithobius variegatus rubriceps (Newport, 1845) se trouve en Irlande, Grande-
Bretagne, Bretagne et dans la Peninsule Iberique (EASON & SERRA, 1986) alors que
Nesoporogaster hispanica Matic & Darabantu, 1969 se trouve dans les Pyrenees espagnoles.

COMPOSITION ET STRUCTURE DES PEUPLEMENTS

Les Figs 1 et 2 representent les abondances relatives des populations d’especes de
diplopodes et de chilopodes dans l’ensemble des stations. Parmi les diplopodes, l'abondance de
Glomeris marginata est remarquable, suivie par Cylindroiulus caeruleocinctus, Marquetiella
lunatum et Glomeris hexasticha intermedia, qui representent plus de 10% du total. En ce qui

Source :

MYRIAPODES DES FORETS PRE-PYRENEENNES

191

concerne les chilopodes, Lithobius borealis et Lithobius variegatus rubriceps ressortent du lot
par leur abondance par rapport aux autres especes representees.

Glomeris marginata 35,9%

Glomeris hexasticha 11,0%

Polydesmus coriaceus
1,9%

Marquetiella lunatum
13,7%

Hirudisoma pyrenaeum 1,1%

Reste 1 ,4%

Ommatoiulus robustus
7,8%

Tachypodoiulus albipes
1,9%

Protoglomeris vasconica 7,4%

Haplopodoiulus spathiferl,7%

Cylindroiulus caeruleocinctus 1 6,2%

Fig. 1. — Abondance relative des differentes especes de diplopodes capturees.

Fig. 1. — Relative abundance of the different species of millipedes in the pitfall traps.

Lithobius calcaratus 3,3%
Lithobius castaneus
Lithobius forficatus 1.1%

Lithobius piceus

Lithobius duboscqui
1,1%

Lithobius pilicornis 1 0%

Lithobius borealis 33,3%

Strigamia acuminata
8,9%

Nesoporogaster hispanica
Lithobius tricuspis 1 • 1 %
3,3%

Lithobius variegatus rubriceps 25,6%

Fig. 2. — Abondance relative des differentes especes de chilopodes capturees.

Fig. 2. — Relative abundance of the different species of centipedes in the pitfall traps.

Source ;

192

ANTONI SERRA. MARIA CRISTINA VICENTE & EDUARDO MATEOS

Si l'on analyse les donnees du Tableau 1, on observe que, meme si le nombre d'especes
est moindre (et plus marque pour le nombre d'individus). les valeurs de richesse specifique,
equitabilite et dominance du total des diplopodes sont semblables a celles obtenues pour le total
des chilopodes. Le trop petit nombre d’exemplaires de ces derniers ne nous a pas permis
d’aborder leur etude precise dans chacune des stations. En revanche, les diplopodes, plus
abondants, permettent de realiser un etude particuliere de chaque zone de piegeage. L'ensemble
des ecosystemes forestiers, hetraie, foret mixte, pinedes et sapimeres, presentent des valeurs
d'indice de diversite tres proches (de 2,44 a 2,14) et tres superieures aux valeurs des autres
stations ; parallelement les valeurs de dominance dune ou deux especes sont relativement faibles
et sensiblement inferieures a celles des zones non forestieres ou alterees.

Les incendies, l’exploitation du bois et le paturage constituent des impacts sur une serie de
stations du massif de San Juan de la Pena qui ont conduit a des alterations notables des
associations vegetales originales. Meme si elles ont actuellement cesse, les perturbations
provoquees sur les communautes d'arthropodes continuent a se manifester. Ce que semblent
prouver les faibles indices de diversite (de 1 .41 a 1,08) et les hautes valeurs de la dominance au
sein des populations des diplopodes.

Tableau 1. — Nombre d'especes pr^sentes (N. sp.). indice de diversite de Shannon (H), diversite maximum (Hmax),
equitabilite (E). dominance par une [D(i)] ou deux [D(i, j)] populations pour le total des diplopodes, le total des
chilopodes et pour les diplopodes de chacune des stations.

Table 1 . — Specific richness (N. sp.), Shannon diversity index (H), max. diversity (Hmax). equitability ( E ). dominance
by one species [D(i)I or two specific populations [D(i, j)] for the whole Diplopoda. the whole Chilopoda, and the
Diplopoda of each site.

N. sp.

Hmax

D(i)

D(i,j)

Diplopodes : Total

2,77

4,09

67,81

35,88

52,1 1

Hetraie

2,44

3,46

70,65

32,88

65,09

Foret mixte

2,36

3,32

71.06

36,96

65,76

Pinede basse

2,36

3,70

63.83

47,25

67,25

Pinede moyenne

2,14

3,17

67,67

43,73

73,48

Pinede haute

2,22

2,81

78,99

37,86

67,14

Sapiniere basse

2,23

3,32

67,05

35,31

59,60

Sapiniere haute

2,25

2,58

86,97

44,90

60,20

Chenaie d'yeuses

1,08

2,32

46,59

61,15

98,43

Chenaie a Quercus faginea

1,32

3,46

38,29

74,64

90,94

Fruticee a E. horridum

1,32

3,00

44,17

74,58

87,43

Fruticee a Genista scorpius

L17

1.58

74,10

61,67

95,00

Paturage

1,41

3,46

40,65

73,56

86,59

Chilopodes : Total

2,70

3,46

78,15

33,33

58.89

La Figure 3 montre le profil d’activite-densite de l'ensemble des chilopodes captures sur le
massif. En general, le nombre d'exemplaires captures par piege et par jour est tres faible, ce qui
peut etre du aux basses densites de population des chilopodes, surtout si on la compare avec les
consommateurs primaires que sont les diplopodes. Les resultats obtenus avec la methode
utilisant des pieges de type BARBER dans d’autres ecosystemes montrent egalement un faible
effectif de chilopodes (ASCASO, 1986 ; SERRA & ASCASO, 1990). Une autre cause probable de
ce phenomene peut etre que les differentes especes de chilopodes presentent une mobilite de
surface assez limitee ; ce dernier facteur pourrait avoir comme consequence que la methode de
capture soit peu appropriee pour revaluation qualitative et quantitative de leurs populations.

Le principal facteur climatique qui semble avoir influence le taux de capture des chilopodes
est la temperature. En decembre et janvier, le nombre d'exemplaires captures est minimum ou
nul, ceci correspond aux mois ou l'on enregistre les temperatures minimales plus basses de
l'annee. Pendant les mois les plus chauds, fin du printemps et ete, le nombre d'individus

Source : MNHN , Paris

MYRIAPODES DES FORETS PRE-PYRENEENNES

193

septembre Tou'tefoh §en gulidrement jusqu'a atteindre un maximum au mois de

— °?Cf°1S’ j 978’ on enre§lstre aussi des valeurs elevees En tout cas

heterogeneite des taux de capture associee a la possibility de variations microclimatiques dans
les di verses stations rend 1'interpretation des resultats particulierement delicate 4

(i/p/j) * 100

Fig. 3. Activit6-densit6 (individus captures par piege et par jour; pour I’ensemble des chilopodes.
Fig. 3. — Activity-density ( individuals per pitfall trap per day) for the whole Chilopoda.

ETUDE DES PEUPLEMENTS DE DIPLOPODES

Le nombre important de diplopodes captures nous a permis de realiser une etude de la
composition de leurs populations pour chacune des stations prospectees. Sur les Figures 4 a 27
sont representes d'un cote les valeurs moyennes annuelles, avec leur deviation standard des
exemplaires de chaque espece, captures par piege et par jour [(I/P/J)* 100] et de l'autre cote le
total des individus captures par piege et par jour des differentes especes pour chaque mois
pendant la penode d echantillonnage. La Figure 28 correspond a une analyse factorielle de
correspondances ou les points “espece" et les points “biotope” sont representes simultanement.
Les abreviations utilisees sur les figures sont detaillees comme suit:

GHE

Glomeris hexasticha intermedia

PAT

Paturaee

GMA

Glomeris marginata

CHY

Chenaie d’yeuses

LHA

Loboglomeris haasi

FRG

Fruticee a Genista scorpius

PVA

Protoglomeris vasconica

CHQ

Chenaie a Quercus faginea

AOS

Archipolydesmus osellai

PIB

Pinede basse

PCO

Polydesmus coriaceus coriaceus

PIM

Pinede moyenne

PRA

Polydesmus racovitzai

PIH

Pinede haute

CCA

Cylindroiulus caeruleocinctus

SAB

Sapiniere basse

HSP

Haplopodoiulus spathifer

SAH

Sapini&re haute

LUM

Leptoiulus umbratilis

HET

Hetraie

ORO

Ommatoiulus robustus

FMI

Foret mixte

OSA

Ommatoiulus sabulosus

FRE

Fruticee a Echinospartum horridum

TAL

Tachypodoiulus niger (=albipes)

BDO

Blaniulus dollfusi

CSP

Ceratosphys sp.

HPY

Hi nidi soma pyrenaeum

MLU

Marquetiella lunatum

194

ANTONI SERRA. MARIA CRISTINA VICENTE & EDUARDO MATEOS

Fig. 4.- Paturage

Fig. 5.- Paturage

Fig.6.- Fruticee (Gen.)

Fig. 7.- Fruticee (Gen.)

Fig. 8.- Chenaie d’yeuses

Fig. 9.- Chenaie d'yeuses

Figs. 4-9. — Valeurs moyennes annuelles, avec leur erreur standard, du taux de capture pour chaquc espfcce (Figs. 4, 6, 8)
et taux de capture mensuel des difterentes especes pendant la periode de piegeagc (Figs. 5, 7, 9) pour les stations
Paturage (4, 5), Fruticee k Genista (6, 7) et Chenaie d’yeuses (8, 9). Se reporter au texte pour les abreviations.

Figs. 4-9. — Annual mean trappability, with standard deviation, for each species (Fig. 4, 6, 8) and monthly trappability
for the different species during the trapping period (Fig. 5. 7, 9) respectively for Pasture (4,5), Genista fruit-tree
site (6, 7) and Oak Q. ilex (8, 9). See the text for abbreviations.

Source : MNHN, Paris

MYRIAPODES DES FORETS PRE-PYRENEENNES

195

Paturage

Dans cette station sont representees 11 especes. Cylindroiulus caeruleocinctus est la
population la plus abondante, ce qui concorde avec les preferences de cette espece pour les zones
c aires. Glomeris marginata & st la plus frequente ; elle se rencontre toute l'annee sauf en ianvier
et en levner. Le graphique de 1 activite annuelle presente deux valeurs maximales, l'une en avril
et 1 autre en octobre (cette derniere est le maximum annuel). A partir de ce point, il y a une
bi usque diminution pendant 1’hiver avec des valeurs nulles au mois de ianvier. L'activite
augmente pendant le printemps et diminue remarquablement pendant les mois d'ete, coi'ncidant
avec les temperatures elevees provoquees par l'insolation et la diminution des precipitations.

En ce qui concerne la phenologie des differentes especes, on observe que Cylindroiulus
caeruleocinctus presente son activite maximale en avril et en octobre, Glomeris hexasticha
intermedia au mois de juin et aussi en octobre, Glomeris marginata en octobre. Protoglomeris
vascomca en aout et Leptoiulus umbratilis en mai. Pour les autres populations, il est impossible
de conclure a cause de leurs frequence et abondance trop faibles (Figs 4 et 5)

Fruticee d Genista scorpius

11 s'agit de l’ecosysteme le plus pauvre en nombre d'especes presentes (3!), et aussi en
nombre dexemplaires captures, seulement 60. Glomeris marginata est le diplopode le plus
abondant et Cylindroiulus caeruleocinctus le plus frequent.

L'activite annuelle est irreguliere et caracterisee par l'absence de captures aux mois de
juillet, septembre et fevrier. Comme sur la station de 1'autre fruticee, la valeur maximale de
1 activite est situee en mars, le seul mois, avec le mois de mai, ou les trois especes apparaissent
simultanement. Glomeris marginata s'y trouve regulierement pendant le printemps, disparait
pendant quelques mois en ete et reapparait en automme. Au contraire, Cylindroiulus
caeruleocinctus , totalement absent dans les piegeages d'ete est present en automme et au debut
de 1 hiver (Figs. 6 et 7).

Chenaie d'yeuses

On remarque que si le nombre d'especes presentes est faible (5) sur cette station le
nombre total dexemplaires est eleve, ce qui est du surtout a l'abondance de Glomeris marginata
et a 1 importante frequence de Cylindroiulus caeruleocinctus.

L activite annuelle est caracterisee par des valeurs peu elevees enregistrees a la fin de
1 automme et en hiver, plus fortes en mars, quand les temperatures commencent a augmenter.
Ces valeurs se maintiennent de fa9on similaire jusqu'a en juillet, epoque de temperatures elevees
et de secheresse et, a partir de ces dates, elles diminuent jusqu'a etre nulles en septembre. En
octobre, avec 1 augmentation des pluies et les temperatures plus douces, on note un fort
accroissement de l’activite, c’est a ce moment qu’on observe le maximum annuel.

Cylindroiulus caeruleocinctus presente une population qui est stable au printemps, dont
1 activite dinunue considerablement en ete pour atteindre sa valeur maximale en octobre, comme
Glomeris marginata. Ce dernier semble mieux supporter la periode estivale, car il maintient et
meme augmente son activite au cours des mois d'ete (Figs. 8 et 9).

Chenaie a Quercus faginea

Malgre une remarquable richesse specifique sur cette station (1 1 especes), la majorite des
especes est tres peu representee, car les 3/4 des exemplaires recueillis sont des Glomeris
maiginata, 1 espece de loin la plus abondante, suivie par Glomeris hexasticha intermedia. Ces
deux especes sont aussi les plus frequentes, presentes dans les pieges pendant 9 des 12 mois
etudies.

Sur le graphique representant les variations annuelles on remarque que, apres le maximum
annuel de captures du mois d'octobre, l'activite diminue brutalement jusqu'a sa disparition

196

ANTONI SERRA, MARIA CRISTINA VICENTE & EDUARDO MATEOS

presque totale fin octobre et au debut de l'hiver. L'activite, avec des valeurs plus ou moins
fluctuantes, se maintient pendant le printemps et l'ete, et diminue de nouveau au cours du inois
de septembre (Figs. 10 et 11).

Sapiniere haute

On y trouve 6 especes. La population dominante est Marquetiella lunatum, avec 44
individus recoltes sur un total de 98. De plus. Marquetiella lunatum est l'espece la plus frequente
car elle apparait pendant toute l'annee sauf en aout et septembre. On observe que l'activite est
faible en fevrier et mars, commence a croitre en mars jusqu'au debut de l'ete et diminue
rapidement pour atteindre les valeurs minimales de septembre. Ce phenomene est suivie d’une
nouvelle augmentation jusqu'en janvier ou se situe le maximum annuel. Marquetiella lunatum
montre son maximum d'activite en automme et en hiver, comme les autres craspedosomatides.

On peut distinguer trois groupes de populations dans cet ecosysteme :

a) celles dont l’activite maximale est centree en fin de printemps et en ete comme
dans le cas de Glomeris marginata ;

b) cedes dont l'activite presente deux maxima, 1'un en ete (juillet) et 1 autre en
automme (octobre) : Protoglomeris vasconica, Glomeris hexasticha intermedia, Polydesmus
coriaceus coriaceus et Haplopodoiulus spathifer ;

c) cedes dont le maximum d'activite se situe en automme et en hiver : Marquetiella
lunatum. On peut penser que ces activites maximales coincident avec les penodes de
reproduction, mais on ne peut toutefois pas negliger certains phenomenes d'attraction qui
peuvent se produire avec le type de pieges utilises (Figs. 12 et 13).

Sapiniere basse

Dans cette station de moindre altitude (1035 m) que la precedente (1415 m), on observe
une plus grande richesse specifique, dix populations au total. La plus frequente est Glomeris
hexasticha intermedia , car elle est seulement absente en janvier et en fevrier, et la plus abondante
est Glomeris marginata qui represente presque le tiers du total des exemplaires.

L'activite dans la sapiniere basse est minimale pendant le mois de fevrier, augmente
progressivement au printemps jusqu'au debut de l'ete, ou se situe le maximum annuel. La
diminution des valeurs d’activite-densite pour les mois les plus chauds est beaucoup moins
accentuee que pour la sapiniere haute, ceci pouvant etre du a i'effet protecteur de l'abondante
strate arbustive de la sapiniere basse, valeurs que Ton retrouve au debut de l’automme. A partir
de ce moment les valeurs diminuent rapidement jusqu'a la fin de l'hiver, ou seul Marquetiella
lunatum apparait. L’ augmentation de l'activite en mars est due a deux causes : a) on y trouve 7
populations du peuplement et b) Glomeris hexasticha intermedia et Glomeris marginata
presented un nombre eleve (qui ne correspond pas aux valeurs maximales) d'exemplaires.

Concernant les populations, on trouve des especes ayant une activite maximale en ete
(juillet), telles que Protoglomeris vasconica et Haplopodoiulus spathifer, d’autres ayant une
activite maximale en fin de printemps et en ete (juillet), comme Glomeris marginata, des especes
a activite maximale en ete (juillet) et en automme (octobre), comme Glomeris hexasticha
intermedia et des especes a activite maximale en hiver, comme Marquetiella lunatum (Figs. 14 et
15).

Hetraie

La richesse specifique de la hetraie est remarquable, avec un total de 1 1 especes presentes.
Glomeris marginata et Marquetiella lunatum represented d'une fa5on generale les 2/3 du total
des exemplaires ; en outre, on observe que Marquetiella lunatum est le diplopode le plus
abondant et le plus frequent.

Source : MNHN , Paris

MYRIAPODES DES FORETS PRE-PYRENEENNES

197

Fig. 10.- Chenaie (Que.) Fig.11.- Chenaie (Que.)

Fig. 12.- Sapiniere haute

Fig. 13.- Sapiniere haute

MR AV MA JN JL AO SE OC NO DE JA FE

Fig-14.- Sapiniere basse Fig.15.- Sapiniere basse

Figs. 10-15.— Valours moyennes annuelles, avec leur erreur standard, du taux de capture pour chaque espece (Figs. 10, 12,
14) et taux de capture mcnsuel des differentes especes pendant la periode de piegeage (Figs. 11. 13, 15) pour les
stations Chenaie (10, 11), Sapiniere haute (12. 13) et Sapiniere basse (14, 15). Se reporter au texte pour les
abreviations.

Figs 10-15 .— Annual mean trappability, with standard deviation, for each species (Figs 10. 12. 14) and monthly
trappability for the different species during the trapping period (Figs 11. 13. 15) respectively for Oak wood (10.
II), High fir-wood (12, 13) and low fir-wood (14, 15). See the text for abbreviations.

Source : MNHN . Paris

198

ANTONI SERRA. MARIA CRISTINA VICENTE & EDUARDO MATEOS

Fig. 16.- Hetraie

Fig. 17.- Hetraie

MR AV MA JN JL AO SE OC NO DE JA FE

Fig. 18.- Pinede haute Fig. 19.- Pinede haute

Fig. 20.- Pinede moyenne Fig. 21.- Pinede moyenne

Figs. 16-21. — Valeurs moyennes annuelles, avec leur erreur standard, du taux de capture pour chaque espece (Figs. 16, 18,
20) et taux de capture mensuel des differentes especes pendant la periode de pi6geage (Figs. 17, 19, 21) pour les
stations Hetraie (16, 17), Pinede haute (18, 19) et Pinede moyenne (20, 21). Se reporter au texte pour les
abreviations.

FIGS 16-21. — Annual mean trappability, with standard deviation, for each species (Figs 16, 18, 20) and monthly
trappability for the different species during the trapping period (Figs 17, 19, 21) respectively for Beechwood
(16, 17), High pine-wood (18, 19) and medium pine-wood (20, 21). See the text for abbreviations.

Source : MNHN, Paris

MYRIAPODES DES FORETS PRE-PYRENEENNES

199

. c L’actjvite annuelle est caracterisee par l'existence de plusieurs augmentations altemees avec

fa DrtseTeT r^,01^™11"1 annuelDde pactivite-densite est atteint en debut de l'ete, du fait de
la presence de G lomeris marginata. Pendant quelques mois d'hiver il existe aussi des valeurs
elevees dues a la presence de Marquetiella lunatum.

,, l>bse,7c °lue Glomeris hexasticha presente deux valeurs maximales, Tune au printemps
et ! autre en automme (mai et octobre). Haplopodoiulus spathifer, Himdisoma pyrenaeum et
Protoglomeris vasconica presented seulement un maximum en aout et au comrrnre

S^F^'r^etTfr = ^ S6mble 8tr£ P‘US aCt‘f 3U C°UrS dU printemPs et au d^ut de

Pinede haute

Dans cette station, oil apparaissent 7 especes, on remarque 1'absence de Marquetiella
lunatum, malgre 1 altitude de la foret (1390 m) favorable a cette espece. L'orientation Ouest de
ce e parcelle et 1 absence de sous-bois provoquant un ensoleillement du sol assez intense
pouiraient expliquer cette absence. L'espece la plus abondante est Protoglomeris vasconica et les
plus frequente*. Glomeris marginata et Polydesmm coriaceus coriaceus. Tom Zen es
seulement en janvier, fevner et septembre.

L’activite annuelle presente deux maxima, fun au mois de juin et l'autre en octobre ce

3nrnr?rrre|PI0ndant ? ma^imum annuel. On observe, dans cet ecosysteme, un deplacement
du pic estival du mois de juillet ajuin, l'importante augmentation de l’activite de mai a juin la
diminution qu, se poursuit jusqu’en octobre et 1'absence totale d'activite en janvier et Ser
a f!hcCe de farquetl^lla •')■ Les temperatures elevees du sol durant les mois d'ete, dues

n, i b| ^ st0us'b0is' 'a dim|nution des pluies, peuvent expliquer l’intense activite en juin

fernJruf! T ^ qUe la §rande activde d’octobre est la consequence "des

temperatures douces et de 1 augmentation des pluies.

de nrimemn^H ^ ph6nologie’ on trouve des especes avec des maxima d'activite en fin

H'Pet d fb'Ut|dete et en au!°mme’ comme Glomeris marginata, des especes avec deux

™ f- C T' UI1 6n 6te Ct aUtr£ Cn automme (octobre), comme Glomeris hexasticha
intermedia et Protoglomeris vasconica et finalement on trouve des especes avec un seul

maximum d activite en fin de pnntemps-debut d ete comme Ommatoiulus robustus (Figs. 18 et
Pinede moyenne

Plus ahnannfLCnerlteI0reu 9 0nt dtd capturdes’ Parmi lesquelles Glomeris marginata est la

plus abondante, car die represente approximativement la moitie des exemplaires captures Elle

est aussi la plus frequente. En second lieu, on observe Glomeris hexasticha intermedia qui
presente des valeurs d activite-densite bien inferieures, mais a une abondance moyenne annuelle
ies supeneure aux autres especes ; en outre, elle est presente pendant les memes mois que
Glomeris marginata sauf en decembre. 4

Comme dans la majorite des stations, le mois de septembre est caracterise par un minimum
au nombre de captures suivi d'un notable accroissement en octobre, ou sont atteintes les valeurs
maximales annuelles. Cependant, on remarque dans cette pinede qu'en fin d'ete il y a un
accroissement important de l’abondance des diplopodes, qui presente un pic au mois d'aout. En
liiver, 1 activite decroit jusqu’a etre nulle en janvier et fevrier ; on remarque l'abondance de
Protoglomeris vasconica parmi les captures de decembre (Figs. 20 et 21).

Pinede basse

Cette station est celle qui presente la plus grande richesse specifique parmi toutes les zones

etudiees, avec un total de 13 especes. La plus abondante et aussi la plus frequente est Glomeris
marginata. ^

200

ANTONI SERRA. MARIA CRISTINA VICENTE & EDUARDO MATEOS

Fig. 22.- Pinede basse

Fig. 23.- Pinede basse

MR AV MA JN JL AO SE OC NO DE JA FE

Fig. 24.- Foret mixte Fig. 25.- Foret mixte

Fig. 26.- Fruticee (Ech.) Fig. 27.- Fruticee (Ech.)

Figs. 22-27. — Valeurs moyennes annuelles, avec Icur crreur standard, du taux de capture pour chaque espece (Figs. 22, 24,
26) et taux de capture mensuel des differentes esp£ces pendant la periodc de piegeage (Figs. 23, 25, 27) pour les
stations Pinede basse (22, 23), Foret mixte (24, 25) et Fruticee a Echinospartum (26, 27). Se reporter au texte
pour les abreviations.

FIGS 22-27. — Annual mean trappability, with standard deviation, for each species (Figs 22, 24, 26) and monthly
trappability for the different species during the trapping period (Figs 23, 25, 27) respectively for low pine-wood
(22, 23), Mixed forest (24, 25) and Echinospartum fruit-tree site (26, 27). See the text for abbreviations.

Source : MNHN; Paris

MYR1APODES DES FORETS PRE-PYRENEENNES

201

Si Ion observe le profil de 1’activite-densite, on constate que la valeur maximale
correspond, comme dans les autres pinedes, au mois d’octobre. Pendant le mois suivant les
hmtm it e,nre?lstrees sorJ,t encorf notablement elevees, mais a partir de decembre il y a une
nlnr?, dl™nuUon ,de 1 activite qui atteint son minimum en fdvrier. L'espece la plus active
pendant cette penode est Marquetiella lunatum. Au printemps et en ete, les captures sont
nombreuses surtout en aout. En revanche en septembre, meme si l'activite decroTt on n'atteint
jamais les valeurs minimales observees sur les autres stations (Figs. 22 et 23).

Foret mixte

d-ins ceuffnrS'ift MSOn ab,0ndafe relatif °lue Par sa frequence de capture, l'espece dominante
dans cette foret est Marquetiella lunatum. La deuxieme est Glomeris marginata , qui presente une

ninir'y611^ CI?ent Tindre- L’activite-densite montre des ressen/blances\ve^

h verni nc!ffetH Va eUfS maximales se Sltuent en automme et, apres une diminution
nlSpnt’ ^ T aV pnntemps, une nouvelle augmentation conduisant a un niveau qui se
maintient jusqu a la fin d ete. Au mois de septembre, comme dans beaucoup d'autres stations
on enregistre une notable diminution de l’activite. ’

„r^P?l°meriS hexasticha intermedia possede un maximum en juillet, Glomeris marginata

l fS mdX'T/ Cn mai.,et(octobre- Proto glomeris vasconica en aout, Haplopodoiulus
spatnijer en mai et Marquetiella lunatum en octobre et novembre (Figs. 24 et 25).

Fruticee a Echinospartum horridum

Cette station est remarquable par la grande abondance des Ommatoiulus robustus. La
hequence maximale est le fait de deux populations, Ommatoiulus robustus et Glomeris
marginata , qui, simultanement, sont absentes en janvier et en fevrier.

Le piofil de 1 activite-densite montre l'existence de deux maxima ; le premier correspond
aux mois de printemps et au debut de l'ete et le deuxieme a Pautomme. Curieusement le
maximum de 1 activite est atteint en mars a cause de la remarquable abondance de Ommatoiulus
robustus. Comme en d autres stations, pendant les mois d'ete l'activite decroit notablement et
atteint un minimum pendant le mois de septembre (Figs. 26 et 27).

La Figuie 28 represente 1'analyse factorielle de correspondances realisee a partir de la
matrice des valeurs totales d'indi vidus captures par piege et par jour des 17 populations de
diplopodes dans chacune des douze stations etudiees. Le premier axe, qui explique 39.5% de la
variance, separe la fruticee a Echinospartum horridum , qui correspond a une ancienne foret
lnccndiee comme nous l’avons mentionne anterieurement, du reste des stations. Le second axe
qui mteiprete 34,2% de la vanance, separe clairement, d’une part, les trois types dissociations
vegetales, le paturage, les vegetations arbustives de la chenaie d’yeuses et la fruticee a Genista
scorpius et, d autre part, les differentes types de forets. Parmi ces derniers, on separe legerement
du reste la chenaie a Quercus faginea qui presente une vegetation typique de 1 etage
su meditenaneen montagnard altere par le paturage ; ces caracteristiques la rapprochent des
conditions observees sur les deux associations arbustives.

Chacune de ces communautes vegetales est caracterisee par differentes especes de
diplopodes qui montrent des preferences claires pour chacune d'elles. Le paturage presente
comme especes caracteristiques Cylindroiulus caeruleocinctus et Leptoiulus umbratilis ; ces deux
especes se trouvent aussi dans les associations vegetales de type arbustif. Parmi ces dernieres, la
chenaie dyeuses abrite aussi Ommatoiulus sabulosus, espece exclusive de cette station La
fruticee a Echinospartum horridum est caracterisee par Ommatoiulus robustus. population la
plus abondante de la station, comparativement a tous les autres sites. Finalement, ce sont les
stations forestieres qui hebergent le plus grand nombre d'especes de diplopodes, en presentant
quelques specificites au niveau de leur abondance relative et de leur frequence. II est clair que la

202

ANTONI SERRA, MARIA CRISTINA VICENTE & EDUARDO MATEOS

majorite des diplopodes du massif de San Juan de la Pena montre une remarquable preference
pour les milieux forestiers.

1.8

O'"

C\[

CvJ

1.6

1.4

1.2

1 -

0.8 -

0.6 -

0.4

0.2

-0.2 -

-0.4 -

Paturage

cca ;

um PAT /

osa

Vegetation arbustive

CHY

\ FRG

gma

/ CHQ,ha

FRE

oro

Foret brulee

ghe PIB
csp

PIM

Forets

PIH SA?ai
pvaHET

p;° fmi

hsp bdo
.hpy

SAH pra
aos

I I I

-3 -2.6 -2.2 -1.8 -1.4 -1 -0.6 -0.2 0.2 0.6

axe 1 (39,5%)

Fig. 28. — Analyse factorielle de correspondances dans laquelle sont represenles les points espfcce et les points station.
FlG. 28. — Factorial analysis of correspondences showing species and sites data.

Source : MNHN. Paris

MYRJAPODES DES FORETS PRE-PYRENEENNES

203

CONCLUSION

type BmSvSit'8 0bKenUfS 1,dtude des sP&im="s captures par les pieges de
K L? nombre de diplopodes est sensiblement plus grand que celui des

qT le,S crrdes' prddatcurs- so“ moi"s atatatdaS

^it/deS^

profondfqm om^rcfpm^^'10"10^65- Pr°PreS a“X h°riz0ns organil)ues ou P'“

Les preferences pour des compartiments edaphiques plus ou moins profonds pourraienr
laisser penser que lut.hsation de pieges d' interception donne des results biaises e,^ ™us
estimes des populations de chilopodes. L'etude comparee des resultats obtenus avec 1'utilisation

so^des^daphiques^^ exemnL^dahai|lti I l°-nnase> pi®gCS de type BaRBER et biocenometres ou
dllin f exemple, dans le meme biotope, permettrait de confirmer cette hypothese
et surtout donnerait une estimation d une part des densites des populations de chaque esmece or
d autre part de la mobilite potentielle de chacune d'elles. q P

De 1 etude des communautes de diplopodes on conclut que les differentes esoeces

vegdtales prtTentes dp ?r6fences Pour les Efferents types ^associations
ege tales presences a San Juan de la Pena. Les milieux forestiers non alteres hetraies

Hm/me^S’ pinedes et forets mixtes> constituent des' ecosystemes ou les peuplements de
dipJopodes montrent une plus grande diversite. Sur un total de 17 especes de? diplopodes

teeS-,dan-S lej, captures, 13 ont une nette preference pour ce type de milieu, etant donne
s aleurs elevees d abondance relative et de frequence de capture obtenues Liberation des
milieux naturels suite a la coupe de bois, au paturage ou aux incendies a co^me ^:ons1quence

nli!rnUtr?°tab e dI. ladlversitd des communautes de diplopodes. L'abondance des
Ke f! nS'dC beaucouP ‘ i especes diminue, jusqu’a disparition. tandis que d'autres semblent

Les valK!??leKe>sede |erH10nS 61 aUgmen,tent remarquablement leur importance numerique.
s. uion Ce dC 'f, dominauce que l’on observe, pour une ou deux especes, sur les

stations destabilises, illustrent b.en ce phenomene. Dans ce sens il faut mentionner

station d!WDitu 6t Lc'p,oudl.ls umbratilis qui montrent une nette preference pour la

station de paturage et, a un degre moindre, pour la fruticee a Genista scorpius et la chenaie

dyeuses. Ommatoiulus sabulosus est exclusif de la chenaie d'yeuses et Ommatoiulus robustus
manifesto une a finite remarquable par la fruticee a Echinosplrtum horruZ De Is ilC
temr compte du fait que 1 effet de perturbation du milieu est durable. A l’exception des pa’tura^es
3 gtre S°riS a CCtte aClivitd Pendant ,a Pcriode de piegeagKene-mfrKe
et S S“Syeu“S PaiU dePl"S ^ ^ nombreuscs am*s d™s » Mcees

REFERENCES

Adis, J„ 1979. — Problems of intercepting arthropod sampling with pitfall traps. Zool Anz., 202 • 171-185

l'lnvesJtigLad!n6e4s ^ * *"***«■ Madrid, .nstituto Fores, a. de

AsCAS°, C„ 1984. Utilizacidn de trampas de ca.'da en dos comunidades forestales de la region mediteminea-
observaciones. In : Ac, as II Congreso Iberico de Entomologfa. Bohn. Soc. Pan. En,. Suppl 1 497-^05

AmoDodo's^d^ 7TT'V° de Pobl“s a partir de muestreos indirectos: aplicacion a comunidades de

Artropodos en dos bosques del Montseny. Tests Doctoral. Barcelona. Publ. Univ. Autonoma de Barcelona

LecheTali’el1: 'Tito*0' ~ ^ ^ ^ Mynapodes de France~ Chilopodes. [Faune Fr.. 25] Paris, P.

Eason, E. H„ 1964. — Centipedes of die British Isles. London. F. Warne & Co Ltd. 294 pp.

EAS,dem!v of Afc I986'kT °n ‘he gCOgraphlCal distribution of Lithobius variegatus Leach. 1814. and the

identity of Ltthobtus rubrtceps Newport, 1845. (Chilopoda. Lithobiomorpha). Journal nat. His,.. 20 : 23-29.

204

ANTONI SERRA. MARIA CRISTINA VICENTE & EDUARDO MATEOS

LAMOTTE. M„ G1LLON, D.. GILLON, Y. & RiCOU. G.. 1969. — L'fichantillonnage quantitatif des peuplements d'inverl6br6s
en milieux herbaccs. In : Problemes d'ecologie : I'echantillonnage des peuplements ammaux des milieux terrestres.
Paris, Masson : 7-54.

MAURlfcS J. P., 1975. — Diplopodes epiges et cavernicoles des Pyrenees espagnoles et des Monts Cantabnques. VIU.

Liste recapitulative, additions, corrections, conclusions. Bull. Soc. Hist. Nat. Toulouse , 111,1/2: 126-134
Pedrocchi - Renault, C., 1985. — Los artropodos epigeos del macizo de San Juan de la Pena (Jaca, Huesca). I.
Introduccion general a su estudio. Pirineos , 124 : 5-52.

Serra A. & AscaSO, C., 1990. — Analisis de la composicion faunfstica y variacion estacional de los Quilopodos de
ires hdbiiats del Montseny (Cataluna) capturados con trampas de cafda. In : A. Minelli. Proceedings 7th Intern.
Congr. Myriapodology. : 385-401.

Source : MNHN, Paris

Study of Centipede Communities of Three Habitats in

the Province of Ciudad Real

Andres Garcia Ruiz & Francisco Javier SANTIBANEZ

Dcpartamcnto de Biologia Animal I (Entomolog.a), Facultad de Ciencias Biologicas, Universidad Complutense

E-28040 Madrid, Espana

ABSTRACT

The present work is a study of centipede communities of three different habitats in the province of Ciudad Real. These
are a poplar grove a brushwood and rubbish heap. The specimens have been collected by direct sampling and by Berlese
tunnels. A check-list of the species collected is given. A frequency and abundance analysis have been carried out The
specific richness and diversity have been compared for the three different biocenoses.

RESUME

Etude des peuplements de chilopodes de trois types d’habitats dans la province de Ciudad Real
(Lspagne).

Le present travail ctudie les peuplements de chilopodes de trois types d'habitats de la province de Ciudad Real, en
bspagne. 11 s agit d une peupleraie, d'un fourre de broussailles el d'un monceau de detritus. Le materiel a ete recolte par
echantillonnage direct e. par extraction au Berlese. On donne la lisle des especes collectees. ainsi que leur frequence et

leur abondance La nchesse specifique et la diversite specifique ont ete estimees de maniere comparative au sein des trois
biocenoses etudi6es.

INTRODUCTION

Usually, when we study the soil fauna, we can observe the great variation that exists
between the habitat characteristics and the community of organisms that live there.

The centipedes, because of their affinity to dampness and their restricted tolerance to
variations of environment humidity, are easier to find in wet areas.

Consequently, it is rather strange to find populations in places which have a low humidity
during the main part of the year.

MATERIAL AND METHODS

En viron m ental D e scrip ti on :

The area where the present study has been carried out is the Campo de Calatrava. province of Ciudad Real, in the
Cornu mdad de Castilla-La Mancha. The afore mentioned Campo de Calatrava is placed in the Submeseta Sur. between the
Montes de Toledo and Sierra Morena. Of note are its old volcanic eruptive defiles caused by isostatics adjustments that
took place alter the Pontiense. Of note are the volcanic products called “Negrizales” or “Castillejos” as well as some
volcanic cones. The soil is dun or red dun of crusty limestone and also mediterranean red soil over siliceous materials

Garcia Ruiz, A. & Santibanez, F. J.. 1996. — Study of Centipede Communities of Three Habitats in the
Province ol Ciudad Real. In: Geoffroy. J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 205-208. Paris ISBN : 2-85653-502-X.

206

ANDRES GARCIA RUIZ & FRANCISCO JAVIER SANTIBANEZ

mainly dedicated to agriculture. The climate is mesomediterranean and the vegetation is typical of the corological Luso-
Extremadurense province.

Studied Habitats

The studied habitats are three areas in the Campo de Calatrava. The first is a poplar grove with an altitude of
700 m and coordinates UTM 30SVJ5608. characterized by an abundance of dead leaves. The second is a brushwood with
an altitude of 869 m and coordinates UTM 30SVH5199. characterized by rocks and scrubs of phytosociological
communities as Cisto clusii - Rosmarinetum and Herniario - Teucrietum pumilii. The third is a rubbish heap, with an
altitude of 800 m and coordinates UTM 30SVH4898, characterized by plenty of debris and vegetal nitrophile
communities.

Methodology:

The habitats were sampled on one day of the first week of every month during 1987.

Five places in each habitat were chosen and the sampling was made in an area of 3 square meters. Check-lists with the
number of specimens of each species were established and the frequency and abundance estimated. In order to compare the
structure of each habitat, the following indices were used:

a) Shannon-Wiener diversity (1949): H’ = - iPj In Pi

This index could be defined as a measure of the specific structure in an ecosystem, based in the number of species
present and their relative abundances.

b) Specific richness according to Margaleff (1980): R = (S-1)/In N, where P is the abundance of each species in
the habitat (Pi = N/N); S is the number of species and N the number of specimens.

RESULTS

Faunistic Composition

On the three studied areas eleven species have been found and can be assembled,
according to a biogeographic point of view and to the BROLEMANN (1930) criteria, in the
following way:

European species:

Mediterranen species:

Holartic species:

Paleartic species:

Betico-riferenian species:

Endemic species:

Atlantic species:

Cosmopolitan species:

Taxocenoses study

In Tables 1, 2 and 3 we can see the number of specimens belonging to the different
species. The total amount of specimens captured was 284 (49 in the poplar grove, 104 in the
brushwood and 131 in the rubbish heap).

In the poplar grove, the total number of centipedes is 49 individuals, belonging to 5
different species. The most abundant species is Pachymerium ferrugineum (40.81%), after that
comes Lithobius lusitanus (32.62%) and Scutigera coleoptrata (22.44%). In relation to the
frequencies we must point out the relative importance of Lithobius lusitanus (31.66%) and
Pachymerium ferrugineum (30.00%).

In the brushwood, the total number of centipedes is 104 individuals, belonging to 8
different species. The most abundant species is Scolopendra cingulata (29.80%), after that
comes Lithobius variegatus rubriceps (24.03%) and Pachymerium ferrugineum (12.50%). In
relation to the frequencies we must point out the relative importance of Scolopendra cingulata
(48.33%) and Lithobius variegatus rubriceps (38.33%).

Lithobius lusitanus Verhoeff, 1925.
Dignathodon microcephalum Lucas, 1846.
Scolopendra cingulata Latreille, 1 829.
Lithobius variegatus rubriceps Newport, 1845.
Pachymerium ferrugineum (C. Koch, 1835).
Schendyla nemorensis (C. Koch, 1837).
Geophilus carpophagus Leach, 1814.

Lithobius inermis L. Koch, 1856.

Cryptops hispanus Brolemann, 1920.
Lithobius pilicornis Newport, 1845.

Scutigera coleoptrata (Linne, 1758).

Source :

CENTIPEDE COMMUNITIES OF THREE HABITATS IN THE PROVINCE OF CIUDAD REAL

207

TA,LE s— — - —

M/H

0.16

0.32

L. pilicornis

L l us it an us
L. inermis

L. variegatus rubriceps
S. nemorensis
P. ferrugineum
D. microcephalum
G. carpophagus
C. hispanus
S. cingulata
S. coleoptrata

0.33

0.77

0.81

2.66

1 1

1 1.66
31.66

3.33

30.00

16.66

0.04

0.40

0.22

Table 2 Brushwood taxocenoses: number of males (M) and females (H) sampled, proportion among the sexes (M/H I
total amount of spec.mens (NT), frequency (F) and abundance (At for each species. *'

M/H

L. pilicornis

L. lusitanus

2.66

1 1

15.00

0.10

L. inermis

3.33

0.01

L. variegatus rubriceps

1 1

1.27

38.33

0.24

S. nemorensis

P. ferrugineum

1.16

13.33

0.12

D. microcephalum

G. carpophagus

0.57

1 1

16.66

0. 10

C. hispanus

3.33

0.01

S. cingulata

1.38

48.33

0.29

S. coleoptrata

0.28

1 1.66

0.08

— Rubbish heap laxoccnoses:

number ol males (M) and females (H) sampled, proportion among th

/r„, total amount ot specimens (NT), frequency (F) and abundance (A) for each species.

M/H

L pilicornis

L. lusitanus

1.71

28.33

0.14

L. inermis

L. variegatus rubriceps

0.72

48.33

0.23

S. nemorensis

P- ferrugineum

D. microcephalum

0.55

23.33

0.10

G. carpophagus

4.50

1 1

16.66

0.08

C. hispanus

1.33

10.00

0.05

S. cingulata

1.31

58.33

0.28

S. coleoptrata

0.33

18.33

0.09

Source

208

ANDRES GARCIA RUIZ & FRANCISCO JAVIER SANT1BANEZ

On the rubbish heap, the total number of centipedes is 131 individuals, belonging to 7
different species. The most abundant species is Scolopendra cingulata (28.24%), after that
comes Lithobius variegatus rubriceps (23.66%) and Lithobius lusitanus (14.50%).

In relation to the frequencies we must point out the relative importance of Scolopendra
cingulata (58.33%) and Lithobius variegatus rubriceps (48.33%).

When comparing the results obtained from the three habitats, we must point out that the
values for the specific diversity are very similar, being 1.73 for the rubbish heap, 1.68 lor the
brushwood and 1 .46 for the poplar grove. The values for the specific richness are: 1 .50 for the
brushwood, 1.23 for the rubbish heap and 1 .02 for the poplar grove. (Table 4) .

Table 4. — Number of species (Nsp). total number of specimens (N), specific richness and diversity in the three studied
habitats.

Nsp

Richness

Diversity

Poplar grove

1.02

1.46

Brushwood

104

1.50

1.68

Rubbish heap

131

1.23

1.73

CONCLUSION

The most abundant species in the rubbish heap and in the brushwood is Scolopendra
cingulata , but it seems to be absent in the poplar grove; this is in agreement with its obvious
preferences for rocky places.

The second most important species, Lithobius lusitanus, is found in the three studied
habitats, showing probably a better adaptability to the different conditions of the environment.

Two other species, Lithobius pilicomis and Schendyla nernorensis, have been found only
in the poplar grove and another one, Lithobius inermis only in the brushwood. They could be
considered as a characteristic for the habitat.

In the present work dealing mainly with a comparison of the three ecosystem types, we
have found very similar diversity values but different richness values. However, these results do
not show any highly significant differences.

REFERENCES

Brolemann, H. W., 1930. — Elements d'une Faune des Myriapodes de France. Chilopodes. [Faune Fr., 25 ]. Paris, P.
Lechevalier : 1-405.

Margalef, R., 1980. — Ecologia. Barcelona, Omega.

Source : MNHN, Paris

Synanthropisation of the Diplopoda Fauna of Poland

Wojciech B. JEDRYCZKOWSKI

Museum & Institute of Zoology PAS, P.O. Box 1007, ul. Wilcza 64, 00-679 Warszawa, Poland

ABSTRACT

More than 5,000 specimens of millipedes belonging to 40 species have been collected and analysed from
synanthropic sites in Poland. Six main zoogeographical elements were distinguished. The European element was
represented by 26 species. Four ecological elements were distinguished, from which synanthropic species were most
abundant in Warsaw (13 species).

RESUME

Anthropisation de la faune de diplopodes de Pologne.

Plus de 5000 specimens de diplopodes appartenant a 40 especes ont ete collectes dans des sites anthropises de Pologne
et Judies. Six composantes zoog£ographiques principales ont 6te distinguees. La composante europeenne est
rcpresentee par 26 especes. On a distingue par ailleurs 4 composantes ecologiques, pour lesquelles les especes
synanthropiques etaient plus abondantes & Varsovie (13 especes).

INTRODUCTION

Most faunistic and ecological publications are dedicated to national parks, protected areas
and other natural environments. Only a few deal with habitats that have been created or modified
by human activities.

Towns, as a typical example of a new habitat created by man for man, are interesting
places for studying processes of plant and animal colonisation, ecological adaptation and
behavioural changes among native fauna. Within towns the process of synanthropisation occurs,
which leads to the colonisation of urban areas by species with different ecological
specialisations. These species and groups of species are able to overcome the ecological barriers
and colonize the direct environment of man. In this way they enrich the ecosystem, forming at
the same time new values for people themselves. Factors controlling these processes and
influencing the development of fauna in “urbiccenoses”, should be recognized as of great
importance for making forecasts as to evaluate them in the phase of planning housing estates,
and to control them in the most convenient way for man and the ecosystem.

MATERIAL AND METHODS

During a 6 year period. 1974 - 1979, faunistic and ecological studies on the animals inhabiting towns in Poland
have been carried out at the Institute of Zoology PAS in Warsaw. These studies were aimed at answering a number of
theoretical and practical questions concerning the state and the role of the fauna in urban environment (Trojan, 1981).

Jedryczkowski, W. B., 1996. — Synanthropisation of the Diplopoda fauna of Poland. In: Geoffroy, J.-J.,
Mauri£s, J.-P. & Nguyen Duy - Jacquemin. M.. (eds), Acta Myriapodologica. Mem. Mus. tiatn. Hist, nat .. 169 : 209-
212. Paris ISBN : 2-85653-502-X.

210

WOJCIECH B. JEDRYCZKOWSK1

So far most of the studies on millipedes (Diplopoda) in Poland have been carried out in natural habitats.
Relatively few workers were interested in habitats transformed by human activates. The research project mentioned
above created opportunities to study the influence of synanthropic factors on the millipede fauna on a large scale.
Warsaw was treated as a model of synanthropic processes which can take place everywhere.

RESULTS

During the study period more than 5,000 specimens of millipedes belonging to 40 species
were collected and analysed from synanthropic sites in Poland. Six main zoogeographical
components were distinguished. The European component was represented by 26 species. Pour
ecological components were distinguished, from which synanthropic species were most
abundant in Warsaw (13 species).

Habitat types

The studied localities can be grouped into the following five main types of habitats
(NOWAKOWSKI, 1981) (Fig. 1).

Warszawa

Fig 1. — Location of the study plots in Warsaw. 1 - suburban areas; 2 - parks; 3 - housing estates; 4 - lawns; 5 -
greenhouses.

1. Suburban areas. An urban wood - a nature reserve - with a surface area of over 130 ha.
It is located on the Vistula river left bank, dominated by a lime-oak-hornbean forest ( Tilio -
Carpinetum), in places modified towards a secondary mixed forest. The woods are inhabited by

Source :

SYNANTHROPIC MILLIPEDE FAUNA IN POLAND

211

10 species of millipedes (Table 1). Six of them: Glomeris connexa, Heteroporatia bosniensis
Nemasoma varicome, Julus terrestris, Leptoiulus proximus and Ophiulus fallax were found
only in this kind of habitat. They are not able to enter the centre of town.

2. Parks. There are several old parks in Warsaw covered by a variety of greens, with
patches ot preserved forests and an area of up to 86 ha. They create good conditions for eleven
species of millipedes living there. Most of them belong to synanthropic species such as
Cylindroiulus caeruleocinctus and Kryphioiulus occultus which achieve their abundance Some
of the natural environment species can also live there.

3. Greens of housing estates. This type of habitat can hardly be characterized, since it
varies in size and floristic composition. The only factors they share are great intensity of
cultivation and very strong penetration by people and domestic animals. In such conditions only
four synanthropic species of millipedes can exist.

4. Lawns and courtyard greens. In general, these types of habitats do not exceed 250 m2
in surface area. The lawns, especially along streets, are heavily contaminated with salt and
subject to frequent drought conditions. The species composition is poor and consist of four
species only.

5 Greenhouses. These are the most artificial of all types of habitats which have been
created by human activity. They have high and constant temperature as well as humidity.
Diplopods dwelling there are typical synanthropic species which origin from tropics. In
Warsaw, seven species of millipedes have been recorded from greenhouses. Three of them
( Ophiodesmus albonanus, Oxidus gracilis and Cylindroiulus truncorum) live exclusively in this
type of habitat.

Table 1. Check-list of the Diplopoda occurring in Warsaw. Numbers of specimens collected.

Suburban

areas

Parks

Housing

estates

Lawns

Green¬

houses

Total

G. connexa

H. bosniensis

B. superus

P. inconstans

138

P. complanatus

0. albonanus

0. gracilis

B. guttulatus

B. tenuis

C. pal mat us

N. varicome

P. fuscus

N. venustus

C. frisius

1 10

C. truncorum

C. caeruleocinctus

242

302

K. occultus

449

451

J. terrestris

L . proximus

1 1

0. fallax

U. foetidus

110

P. ger manic um

Total

211

967

110

1368

212

WOJCIECH B. JEDRYCZKOWSK1

Species groups

Analysing the species composition of the millipedes fauna of synanthropic habitats, we can
conclude that the Diplopoda occurring in Warsaw belong mostly to the species showing high
ecological amplitudes and due to this they are able to inhabit several urban habitats
(JEDRYCZKOWSKI, 1982). From an ecological point of view, they can be divided in three

groups depending on the transformation of environment.

I . The first group consists of native species having a great ecological tolerance which are
able to enter into suburban greens, but are not able to live in the central part of town. It is
represented by G. connexa, H. bosniensis, P. complanatus, N. varicorne, P. fuscus, J.

terrestris, L. proximus and O. fallax. . .

2 The second group represents mainly synanthropic species, which originates mainly
from the south and west part of Europe, they dwell in parks and estate greens but occasionally
can inhabit some natural or seminatural habitats. As representatives of this group, P. inconstans,
C. latestriatus (=frisius ) and C. caeruleocinctus can be mentioned.

3. The third group is build up by species of tropical or unknown origin. Usually, they live
in a variety of greenhouses (where they can achieve a high numbers of individuals) sometimes
they can live out of buildings for a long period, especially when winters are mild.

DISCUSSION

The species diversity and the number of specimens occurring in towns depend on the age
and type of the inhabited greens. The highest specific richness occurs in old parks, with large
green patches, dense lawns with shrubs and where the litter is well preserved, providing shelter

for animals. . .

A high percentage of millipedes belong to the group of expansive species, which have
successfully colonized almost the whole Europe. The high ecological amplitude enables them to
adapt to changing habitat conditions. Synanthropic species, especially those belonging to the
genus Cylindroiulus, native of the Mediterranean, are a good example here. Urban pressure
firstly eliminates the species associated with forests and thickets. They still can live in the
suburbs but they do not colonize parks and housing estates.

REFERENCES

JEDRYCZKOWSKI, W„ 1982. — Millipedes (Diplopoda) of Warsaw and Mazovia. Memorabilia Zool. Warszawa, 36 : 253-
26 1 .

Nowakowski, E., 1981. — Physiographical Characteristics of Warsaw and the Mazovian Lowland. Memorabilia Zool.
Warszawa, 34 : 13-31.

Trojan. P., 1981. — Urban Fauna: Faunistic, Zoogeographical and Ecological Problems. Memorabilia Zool. Warszawa,
34 : 3-12.

Source

Chilopoda of Urban Greens in Warsaw

Jolanta WYTWER

Muzeum i Instytut Zoologii PAN ul. Wilcza 64, 00-679 Warszawa, Poland

ABSTRACT

A total of twelve Chilopoda species have been registered in three types of urban greens in Warsaw: seminatural wooded
areas, big parks and streetside lawns. The common core of Chilopoda communities in all three types of urban areens
contained six eurytopic species, with Lithobius microps as the most abundant. There were noticeable changes 'in the
dominance structure of the epigeic part of Chilopoda communities related to urban greens types. Lithobius microps
gradually replaced Lithobius foificatus, which is a very abundant species in wooded areas but relatively scarce in street

RESUME

Chilopodes des espaces verts urbains de Varsovie.

Les peuplements de chilopodes ont et6 etudies dans trois types d’espaces verts urbains k Varsovie : des boisements

61 deS pe,ouses silu6es pr6s des chaussees. Le materiel a ete recolte au cours de la periode
1 ->88-1990 a 1 aide d echantillons de sol et de pieges Barber. Au total, on a recolte douze especes de chilopodes dont le
plus grand nombre se trouve, en moyenne. dans les boisements (5,8), alors que la plus faible richesse specifique est
enregistree dans les pelouses (4,5). Dans les trois types d’espaces verts, les six especes communes apparaissent avec une
Constance d£passant 50% et constituent environ 2/3 de la composition taxonomique du peuplement. Ce sont : Lithobius
microps, Necrop h loeophag us flavus, Schendyla nemorensis , Geophilus electricus , Strigamia crassipes et Lithobius
jorjicatus. Dans les trois types d’espaces verts L microps predomine. L’analyse des structures dominantes des
peuplements de chilopodes a I’aide des indices de Morisita, de l’homogen6ite de domination et de rangs, a montre
I existence de changements dependant du degre d’anthropogenisation des espaces verts urbains. Dans le compartiment
epigeique, on a observe que 1’importance relative de L microps s’accroit au detriment de L. forficatus. qu il remplace
progressivement. ^ y

INTRODUCTION

Studies of Chilopoda conducted over the last several decades in many European cities, e.g.
in Copenhagen (ENGHOFF, 1973), Kiel (TlSCHLER, 1980), Goteborg (ANDERSSON, 1983) and
Rome (ZAPPAROLLI, 1992) indicate that urban Chilopoda communities are characterized by high
proportions of alien species and a significant degree of faunal diversification. Striking
differences in the abundance of Chilopoda are noted between individual study sites. Equally
unusual is the occurrence of many species as single specimens, a phenomenon which was
particularly conspicuous in a quantitative study carried out in Bonn Bad-Godesberg (FRUND,
1989, SCHULTE et al. 1989). We do not know yet, however, whether Chilopoda communities
inhabiting different types of urban greens belong to one or more faunal associations, in other

Wytwer, J., 1996. — Chilopoda of Urban Greens in Warsaw. In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen
Duy - JACQUEMIN, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, not., 169 : 213-220. Paris ISBN : 2-85653-
502-X.

214

JOLANTA WYTWER

words whether the differences in species composition and species abundance are accidental or
testify to their individual characters. The following paper analysing Chilopoda of urban greens
of Warsaw attempts to provide an answer.

MATERIAL AND METHODS

Sampling^ was carried oul in 13 sampling areas that represented 3 types of urban greens (Fig. 1):

1. wooded areas with ground cover growing spontaneously. Such areas are notsubject toregularhortcutura
nractices (e 2 digging over or raking). They are located along the edge of the erosion valley of the Vistula river that cuts
across the area of Warsaw. Areas W1 and W5, situated on the outskirts of Warsaw, represent the inden-o

forest in phytosociological terms. Areas W1-W4 are characterized by a high inclination angle. Area W5 is the only one
situated on the upper erosion terrace within the Lasek Bielahski reserve.

2. park lawns are subject to regular horticultural practices i.e. lawn trimming and litter raking. Areas P1-P4 are

located within larger park areas at least 50 meters away from a roadway.

3. street lawns (S1-S4) are situated in the immediate vicinity of busy arterial roads.

Sampling methods

Two sampling methods were used for collecting Chilopoda: f

1) Barber’s pitfall traps. 10 traps were placed in each area. The animals were collected once a fortnight irom

April *988 tt> ^ with ^ area of Q j m2? laken down lQ a deplh of aboul 25 cm. 3 or 5 samples were taken in May
and September 1990. 32 soil samples were taken altogether in each of the three types of urban greens. The specimens
were sorted by hand.

0 wooded areas : W1 . W2, W3, W4, W5
| parks : PI, P2, P3. P4

street lawns : SI , S2, S3, S4

Fig. 1. — Location of the urban greens in Warsaw.

Source : MNHN . Paris

CHILOPODA OF URBAN GREENS OF WARSAW

215

RESULTS

Species composition

12 species of centipedes were recorded from urban greens in Warsaw (Table 1) The
greatest numbers of species were found in wooded areas - 5.8 on average, compared to 5 0 in
park lawns and only 4.5 in street lawns. As far as the number of species is concerned the
Ch, opoda communities of urban greens of Warsaw are not basically different from forest
foil )p°f aDroI^Tnn71nieD0f Central EuroPe’ where 5-10 species are usually recorded (THIELE
1992) ALBERT’ 979' BECKER’ !982; Fr°ND, 1987; KACZMAREK, 1989; WYTWER, 1990

Table 1. — Species of Chilopoda in urban greens of Warsaw.

Species

wooded areas

W1 W2 W3 W4 W5

parks

PI P2 P3 P4

street lawns

SI S2 S3 S4

1 1
12

Lithobius forficatus (L.)

Lithobius melanops Newport

Lithobius mutabilis L. Koch

Lithobius crassipes L. Koch

Lithobius microps Meinert

Lamyctes fulvicornis Meinert
Necrophloeophagus flavus (De Geer)
Clinopodes linearis (C. Koch)

Geophilus electricus (L.)

Brachy geophilus truncorum (Berg. & Mein.)
Strigamia crassipes (C. Koch)

Schendyla nemorensis (C.Koch)

+ + + +

+ +

+ + + + -+

+ + + + +

+ + + +

+ + + + +

+ + +

+ +

+ + + +

+ + +

+ + + +

+ + +

+ +

+ + + +

Number of species

7 5 6 6 5

6 5 5 4

6 5 2 5

The following six species occurred in all the three types of urban greens and were
consequently considered to be common: Lithobius forficatus, Lithobius microps,
Necrophloeophagus flavus, Geophilus electricus, Strigamia crassipes and Schendyla
Thc? Similanty of the sPecies composition of such communities, expressed as the
60^™eS2)'STEINHAUS mdeX (MARCZEWSK1 & Steinhaus, 1958) oscillates closely around

Table 2. Similarity (in percent) of species composition in Chilopoda communities in urban greens of Warsaw
according to Marczewski-Steinhaus index (MS).

Type of urban greens

wooded areas x parks

54.5

parks x street lawns

60.0

wooded areas x street lawns

66.7

In order to determine the degree of fidelity of individual Chilopoda species to the urban
f J!een 1labltat' a constancy analysis was performed using TlSCHLER’s method (TlSCHLER,
!( 4;>' The analysis. revealed the presence of two groups of species (Fig. 2). The first group is
characterized by an index of constancy above 0.5 and is composed of the same six species listed
above as common to communities of all the three types of urban greens. The other °roup
consists of species with much lower values of constancy (below 0.2).

U can be assumed that the Chilopoda species occurring in all the three types of urban
greens and characterized by high constancy indices form the core of the Chilopoda communities

216

JOLANTA WYTVVER

of urban greens. The species from the other group are found occasionally in single sites and
function as accessory elements ol the community.

Lithobius microps

Necrophloeophagus flavus
Schendyla nemorensis
Geophilus electricus
Strigamia crassipes
Lithobius forficatus
Lithobius mutabilis
Clinopodes linearis
Lamyctes fulvicornis
Lithobius melanops
Brachy geophilus truncorum
Lithobius crassipes

0.1 0.2 0.3 0.4 0

constancy

FIG. 2. — Constancy of occurence of individual Chilopoda species in the urban greens of Warsaw.

The structure of dominance

The combining of the two methods of sampling helped to clarify and compare the
dominance structure of Chilopoda communities inhabiting the surface and ^“Per
(Fig. 3a, b). The similarity of dominance relationships are well reflected by M°R1srrAs index
MO (HORN, 1966) and the index of homogeneity - HD (RlEDL, 1963). When the centiped
fauna of the deeper layers of soil is analysed, the values of both indices are relatively high -
above 90 and 70% (Table 3). Therefore it can be assumed that the structure of this part of the
community is uniform in all types of urban greens in Warsaw.

Table 3 — Similarity (in percem) of dominance structure of Chilopoda communities. MO: index of similarity '^orisita
index); HD: index of homogeneity, based on the species dominance structure; HR: index of homogeneity, based

on the ranks of dominance structure.

Method

Type of urban greens

soil

wooded areas x parks

98.5

80.2

83.1

samples

parks x street lawns

96.0

73.8

74.0

wooded areas x street lawns

98.1

78.2

78.1

pitfall

wooded areas x parks

79.4

51.0

83.9

traps

parks x street lawns

75.2

42.0

41.2

wooded areas x street lawns

55.3

32.0

36.9

The similarity of the dominance structures of the epigean parts of Chilopoda communities
of different types of greens is expressed by much lower values of the both indices - below / )
and 51%. This is mostly caused by a decrease in the proportion of Lithobius forficatus in parks
and street lawns (Fig. 3b). This species is being gradually replaced by Lithobius microps.

Source : MNHN, Paris

CHILOPODA OF URBAN GREENS OF WARSAW

PARKS

1.1% 1

3./%

0.5% |

STREET LAWNS

16.7%

LEGEND:

Lithobius forficatus
Lithobius microps
Necrophloeophagus flavus
Geophilus electricus
Strigamia crassipes
Schendyla nemorensis
Lithobius crassipes
Clinopodes linearis
Brachygeophilus truncorum

Fig. 3a. — Percentage contribution of Chilopoda species in the soil sample material.

JOLANTA WYTWER

WOODED AREAS

3.8%

PARKS

3.6%

STREET LAWNS

3.2%

2.4%

0.8%

LEGEND:

Lithobius forficatus
Lithobius melanops
Lithobius mutabilis
Lithobius microps
Lamyctes fulvicornis
Necrophloeophagus flavus
Geophilus electricus
Strigamia crassipes
Schendyla nemorensis

Fig. 3b. — Percentage contribution of Chilopoda species in the pitfall trap material.

CHILOPODA OF URBAN GREENS OF WARSAW

219

j The structure of dominance of the epigean component of Chilopoda communities is

rnfono!!? ^aJOr Chvges- Th,S is not’ however’ a s'gn Of a change of the dominance model of
Chilopoda communities in various types of urban greens. The degree of overlap of the

dominance models has been determined by means of a homogeneity coefficient where
dominance models arranged according to the ranks (shares) of species provided a basis for the

Swbi?n?g t°JhlS simila,nty of dominance relations could be analysed irrespective of the
ect what species lakes succeeding position. The values of “ranks homogeneity coefficient” (HR)

JlabTe 3 I ’SSnr" 0f WOOdHd r as a"d parkS have a 1- do_ idd

(lab e 3). It differs however, from the dominance model of Chilopoda communities of the

m the epigean ^yer- Where the dominance Of Lithobius microps is very
nreTsure whuh e’ T' JC[urmg ,of Chilopoda communities under marked urbanizing

nrmf w . c'^rentIy takin§ place in urban greens, affects above all else the epigean
J S? f inhabiting deeper layers of soil are able to preserve structural relations in an
almost intact form, the only modifications being due to the exchange of accidental species.

DISCUSSION

Except for Strigamia crassipes, all the species which form the core of Chilopoda

whe!-cUcemmpfHUrbfan T kn°Wn !° be euryt0Pic and occur in most of the European cities
where cent pede fauna has been studied. The occurrence of S. crassipes in urbanized

Ihlnrt Qmen C°U d ^RVC beCn underest,mated so far since this species reaches its maximum of

ChlotnrH T!' Jr ' ^BARf er &k.KeaY’. 1988) and « therefore not included in faunal studies of

of So hte atSt S thlS SpeC" WCTe by pi,M1 trappm8 ” ***

snrear/hv nfh SP,6C1 ?? wbich have been found in Parks a"d street lawns are known to
spread by means of horticultural practices in artificial man-made environments as it is the case

r„:?r linearis and Brachygeophilus truncorum. On an other hand, wooded areas and

h kn , 10 LhC C?nt;re ,°f the t0Wn 3X6 often abundant in species that once inhabited the natural

habitats, such as Lithobius crassipes, Lithobius melanops and Lamyctes fulvicornis which are

fawnt m 3 synanthr°Plcuenv,ronment. However, they are not able to occur on the street

lawns because they belong to the epigean part of the community which is undergoing marked

smvSonirin^H1^118' n CXu1US1Ve,ly forest sPecies’ such as Lithobius mutabilis , has
°n'y in w°oded areas on the outskirts of the city (linden-oak-hombeam forest) as it is a

Low and (WYTSaT990)ment “ ChlI°P°da communides of this habitat type in the Mazovian

CONCLUSION

One type of centipede community can be considered to occur in urban greens of Warsaw
its core consists of 6 eurytopic species that do not avoid “synanthropised” environments. The
other species are distributed randomly among various sites where they function as accessory
species The influence of the urban environment on the structure of Chilopoda communities
manifests itself in marked transformation of the epigean part of the community.

ACKNOWLEDGEMENT

I am gratelul to prof, dr hab. P. Trojan for helpful advice and critical review of ihe manuscript

REFERENCES

Albkrt, A. M 1979. — Chilopoda as part of the predatory macroarthropod fauna in forests: abundance, life-cycle,
biomass and metabolism, In : M. Camatini, Myriapod Biology. London, Academic Press * 215-231

ohnineH ' inG;h^Sn” faun* in the ™nity °f G°teborS ~ a comparison between collecting results

obtained in the 1920s and the 1970s years. Acta ent. fenn. Helsinki , 42 : 9-14.

220

JOLANTA WYTWER

Barber, A. D. & Keay A. N., 1988. — Provisional atlas of the centipedes of the British Isles. Huntington, Biological

Records Centre. 127pp. .. n , 77 . 7A

BECKER J , 1982. — Hundertfussler (Chilopoda) des Bausenbergs und der osthche Eifel. Dechemana , 27 . 70-Xb.
Enghoff. H.. 1973. — Diplopoda und Chilopoda from suburban localities around Copenhagen. Vidensk. Meddr dansk

naturh. Foren., 136 : 43-48. . ,

FrOnd. H. C.. 1987. — Raumliche verteilung und Koexistenz der Chilopoden in einem Buchen-Altebestand.

Pedobiologia , 30 : 19-29. . , , .. , r

FrOnd. H. C., 1989. — Untersuchungen zur Biologie stadtischer Boden. 5. Epigaische Raubarthropoden. Vern. ues.

Horn, H. S.. 1966. — Measurement of “overlap” in comparative ecological studies. Am. Nat.. 100 : 410-424.
Kaczmarek, J.. 1989. — Pareczniki (Chilopoda) wybranego lasu gradowego Wielkopolski na przyka?adzie Rczerwatu

“Jakubowo". Fragm. faun.. 32 : 369-379. f A n n

Marczewski. E. & Steinhaus, H., 1958. — On a certain distance of sets and corresponding distance ol function. Loll.

Math.. 6 : 319-327. , , ^ , _ . c ,

Riedl, R., 1963. — Problemc und Methoden der Erforschung des litoralen Benthos. Verb. d. Dtsch. Loot.. .Suppl. 2(»

Schulte, W. et al. 1989. — Untersuchungen zur Bodendkologischen Bedeutung von Freiflachen ini Stadthereich.

Forschungsbericht des Bundesministerium fur Forschung und Technologie, Hamburg, 200pp.

Thiele, H. V. U., 1956. — Die Tiergesellschaften den Bodenstreu in den verschiedenen Waldtypen des Niederbergischen
Landes. Z. angew. Ent.. 39 : 316-369.

TlSCHLF.R, W.. 1949. — Grundziige der terrestrischen Tierdkologie. Berlin, F. Vieweg & Sohn. 220pp.

Tischler, W., 1980. — Asseln (Isopoda), Tausendfussler (Myriapoda) eines Stadtparks im Vergleich mit der IJmgebung
der Stadt: zum Problem der Urbanbiologie. Drosera, 80 : 41-52.

Wytwer. J., 1990. — Chilopoda of linden-oak-hombeam ( Tilio-Carpinetum ) and thermophilous oak forests (Potentillo

albae-Quercetum) of the Mazovian Lowland. Frag. faun.. 34 : 73-94.

Wytwer. J., 1992. — Chilopoda communities of the fresh pine forests of Poland. In : (E. Meyer, K. Thaler & W.

SCHEDL, Advances in Myriapodology.] Ber. nat.-med. Verein. Innsbruck, Suppl. 10 ; 205-211.

Zapparoli. M.. 1992. — Centipedes in urban environments: records from the city of Rome (Italy). In : [E. Meyer. K.
Thaler & W. SCHEDL, Advances in Myriapodology.] Ber. nat.-med. Verein. Innsbruck. Suppl. 10 : 231-236.

Source :

Centipedes of Poznan Town (Poland)

Malgorzata LESNIEWSKA

Zaklad Zoologii Ogolnej, Uniwersytct im. Adama Mickicwicza, ul. Fredry 10, 61 - 701 Poznan, Polska

ABSTRACT

,iiS!'a",,ltfIVe S(UdfieS °r lhe cemipedes 01 Poznan have becn carried out since 1988. The studies cover areas with
dillerent degrees of transformation by man, like parks, squares, cemeteries, allotment gardens, dumping grounds etc As

orPnl ° 'hese studies, the occurence of 18 centipede species has been-found, constituting 33% of the Chilopodan fauna

ord^r Kl1,0n8 H m“*Jreque?| species found ln Pozna"- Lilhobius forfica'us and Liihobius mi crops from the
„ 'Obtomorpha and Necrophloeophagus flavus and Schendyla nemorensis from the order Geophilomorpha. The
occurence ol Haploplulus subterraneus has been registered for the first time in Poland. The studies are to be continued.

RESUME

Chilopodes de la ville de Poznan (Pologne).

Les chilopodes de Poznan son. etudids depuis 1988. Les recherches sont effectuees sur des sites diversement
translormes par les activitds humaines, tels que pares, squares, cimetidres. espaccs verts, jardins, amas d ordures, etc.
Nous avons constate la presence de 18 especes. representant 33% de la faune des chilopodes de Pologne. Parmi les
especes les p us frequentes, on note : Liihobius forficatus et Lilhobius microps pour lordre Lithobiomorpha,
Nea ophloeophagus flavus et Schendyla nemorensis pour lordre Geophilomorpha. L'espece Haplopliilus subterraneus a
ete repertoriee pour la premiere fois en Pologne.

INTRODUCTION

, *n European myriapodological literature of the recent years, we can find some works
aesciiDing the urban centipede fauna including -among others- Copenhagen (ENGHOFF 1973)
Goteborg (ANDERSSON, 1983) or Rome (ZAPPAROLI, 1990a. b).

In the polish literature, there are no works of this type so far, although in recent years,
thice masters theses were prepared referring to centipedes of Poznan and one study devoted to
this group of animals is under preparation in Warsaw (WYTWER, this volume).

This work presents preliminary results of qualitative studies carried out since 1988 in
Poznan.

STUDY AREA

f , c°/nan 'S lhC largeSI t0Wn in wie|k°polska, founded in the 9th century. It is situated on Warta river at the altitude
ol 52- 54 m above sea level, covering 261.3 km2, with 589.7 thousands inhabitants. The climate is moderately
continental . I he annual rainfall is the lowest in Poland, below 500 mm. The annual isotherm is 8.5°C. The winters are

Lesniewska, M., 1996. — Centipedes of Poznan town (Poland). In. Geoffroy, J.-J., Mauries, J.-P. & Nguyen
Duy - JACQUEMIN, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 221-224. Paris ISBN : 2-85653-
502-X.

222

MALGORZATA LESNTEWSKA

mild (-2°C) and summers are warm (18°C). The growing season lasts 210-220 days. The relief is characterised by Baltic
glaciation. The soils include podzolic and brown soils. , .

Poznan is a city of International Trade Fairs, visited since 1921 by business men presenting their merchandise
from all over the world. This may exert an influence on the spreading of some plant and animal species.

MATERIAL AND METHODS

The following results refer to qualitative studies carried out in 1988. The material was collected by direct
sampling of specimens under stones, timber, stems, etc. and by the use of litter sieving.

Samples were taken in 33 localities (in parks, cemeteries, squares, dumping grounds, etc.)- These localities were
divided into 3 categories, taking into consideration primarily their degree of influence by the activity of man.

Category I. - areas completely transformed by man and his continuous interference (e.g. railways, embankments,
wild dumping grounds, roadsides).

Category II. - areas partially changed, covered with vegetation maintained by man (squares, cemeteries, allotment
gardens).

Category III. - areas subject to the least interference (mainly less cultivated parks and afforested areas on town

edges).

Each locality was inspected at least 5 times. The material was collected by up to three people. During 5 years
(1988 - 1992), 1628 individuals belonging to Chilopoda orders were collected. The results include also some materials
collected for a masters theses.

RESULTS

During the studies on the area of Poznan, the occurence of 18 centipede species have been
found, including 1 1 belonging to the Geophilomorpha, 1 to the Scolopendromorpha and 6 to the
Lithobiomorpha. For the first time, the two species Clinopodes linearis and Clinopodes flavidus
have been registered for the fauna of Wielkopolska, and the occurrence of Haplophilus
subterraneus has been found for the first time for the fauna of Poland. The most frequently
occurring species include two lithobiomorphs: Lithobius microps and Lithobius forficatus and
two geophilomorphs: Necrophloeophagus flavus and Schendyla nemorensis (Table I).

Table 1. — The occurence of centipedes in particular habitat types (number of localities).

LIST OF IDENTIFIED SPECIES

Category

III

Frequency

Haplophilus subterraneus (Shaw)

Schendyla nemorensis (C. L. Koch)

48%

Strigamia crassipes (C. L. Koch)

21%

Strigamia acuminata (Leach)

Pachymerium ferrugineum (C. L. Koch)

Clinopodes linearis (C. L. Koch)

12%

Clinopodes flavidus C. L. Koch

Geophilus elect ricus (Linn6)

30%

Geophilus proximus C. L. Koch

Necrophloeophagus flavus (De Geer)

48%

1 1

Brachy geophilus truncorum (Bergso & Meinert)

27%

Cryptops hortensis Leach

24%

Lithobius forficatus (Linne)

73%

Lithobius erythrocephalus C. L. Koch

18%

Lithobius melanops Newport

24%

Lithobius crassipes L. Koch

27%

Lithobius curtipes C. L. Koch

Lithobius microps Meinert

1 1

90%

Number of species

Remarks on the species (in decreasing frequency order )

1. Lithobius microps Meinert - European, eurytopic species, showing a tendency to occur
very frequently in man-made habitats (EASON, 1964; ENGHOFF, 1973; ANDERSSON, 1983;

Source . MNHN, Paris

CENTIPEDES OF POZNAN TOWN

223

Barber, 1985; Lewis, 1985; Zych, 1989). In Poznan, this is the most frequent and numerous
species in all categories of localities (Table I). 4 numerous

2. Lithobius forficatus (Linne) - W-Palaearctic, eurytopic species. In Poland this is the

freque^UIewrywihere.SentatlVe °f ^ (KaCZMAREK’ 1979 • 1980). In Poznan it is very

3 Necrophloeophagus flavus (De Geer) - Palaearctic, eurytopic species. In Poland it is

categories ^ reC°rdS fr0m P°Znan haVe been ColIected ln a11

-S woodland *“*“• '* *

_ GeoP,nl“s electricus (Linne) - European, eurytopic species, with a tendency to be more
numerous in uiban localities. According to KACZMAREK (1980), in Poland it is rare and not

(Table”) 1 P ° 11 Seems t0 occur in habltats partially and completely changed by man

6- Lithobius crassipes L. Koch - European, woodland species. In Poland, it is common in
natural and synanthropic localities. In Poznan the highest proportion of records has been
collected in areas of the category III (Table I).

7. Brachygeophilus truncorum (Bergso & Meinert) - European, eurytopic species. In
Roland, common in woodlands in the west part of the country (KACZMAREK, 1980). In
Poznan, it has been found in all habitats sampled.

- ii f^ithobius fnelanops Newport - Palaearctic, woodland species. In Poznan, it has been
collected in all categories of habitats but the largest proportion of records has been obtained from
woody areas.

?■ Cryptops hortensis Leach - Palaearctic, eurytopic species, in some regions
synanthropic. The records from Poznan have been collected especially from less cultivated parks
and woody areas (Table I). F

iO Strigamia crassipes (C. L. Koch) - Palaearctic, eurytopic species. In Poland, it occurs
in woodlands and synanthropic areas.

11. Lithobius erythrocephalus C. L. Koch - European, eurytopic species. In Polish

lowlands, apart from L. forficatus and L. mutabilis - the most common representative of the
genus. In urban localities, it is rare (KACZMAREK, 1980). In Poznan, it is mostly found in
afforested areas on town outskirts (Table I). 3

12. Clinopodes linearis (C. L. Koch) - European, eurytopic species, in Poland mainlv in

synanthropic areas. J

13. Lithobius curtipes C. L. Koch - European, woodland species. In Poland, it occurs in

uiban localities (KACZMAREK, 1980). The records from Poznan have been collected only from
wooded areas (Table I). J

14. Geophilus proximus C. L. Koch - European, woodland species. In Poznan, it has
been recorded in areas changed by man.

15 Clinopodes flavidus C. L. Koch - Palaearctic, woodland species. In Poland it is very
rare. In Poznan, it has been collected for the first time for Wielkopolska.

16. Pachymerium ferrugineum C. L. Koch - Holarctic, eurytopic species. In Poland (and
in roznan), it occurs outside forests, in warm, dry places.

1 7. Strigamia acuminata (Leach) - Holarctic, woodland species. In Poznan, the records are
rrom one old park.

18. Haplophilus subterraneus (Shaw) - an introduced species new for the fauna of Poland
I he records (3 specimens) are an old park (LESNIEWSKA & WOJCIECHOWSKI, 1992).
Therefore, one may say that, from a zoographic point of view, the centipedes of Poznan

represent the following elements;

- European - 8 (47%),

- Palaearctic - 7 (41%),

224

MALGORZATA LESNIEWSKA

- Holarctic -2 (12%).

On an other hand, due to ecological requirements, the following species categories can be
distinguished:

- eury topic -10 (59%),

- woodland -7 (41%).

(Haplophilus subterraneus has not been taken into account.)

It has been found that the fauna of Poznan is poorer by 15 species than the Wielkopolska
region where it is situated. The centipedes of Poznan represent 55% of the fauna of
Wielkopolska and 33% of the fauna of Poland. Quantitative studies are under investigation.

CONCLUSION

The present results should be regarded as preliminary ones because quantitative studies are
still under investigation. Nevertheless, the species composition and data referring to the
frequency of occurrence are similar to those obtained by other authors investigating on the
Chilopoda fauna of European towns (ENGHOFF, 1973; ANDERSSON, 1983; ZAPPAROLI, 1990
a, b).

REFERENCES

Andersson. G„ 1983. — The Chilopod fauna in the vicinity of Goteborg - a comparison between collecting results
obtained in the 1920s and the 1970s. Acta Entomol. Fenn., 42 : 9-14.

Barber, A. D„ 1985. — Distribution patterns in British Chilopoda. Bijdr. Dierk.. 55 : 16-24.

Eason, E. H„ 1964. — Centipedes of British Isles. London. F. Warne& C° Ltd, 294 pp.

ENGHOFF, H., 1973. — Diplopoda and Chilopoda from suburban localities around Copenhagen, Vidensk. Meddr dansk
nathur. Foren ., 136 : 43 - 48.

KACZMAREK, J., 1979. — Pareczniki ( Chilopoda ) Polski. Poznan, UAM.

Kaczmarek, J., 1980. — Katalog fauny Polski. Pareczniki. Czeceze XIV.

LESNIEWSKA, M. & Wojciechowski, J., 1992. — Haplophilus subterraneus (Shaw. 1794) (Chilopoda, Geophilomorpha)
- nowy dla fauny Polski przedstawiciel parecznikdw. Przeg. Zool. XXXVI, 1 - 4 : 133 - 136.

Lewis, J. G. E., 1985. — Centipedes enterning houses with particular reference to Geophilus carpophagus Leach. Em.
mon. Mag., 121 : 257-259.

Zapparoli, M., 1990a. — Centipedes in Urban Environments: Records from the City of Rome (Italy). Per. nat. - med.
Verein Innsbruck. Suppl 10 : 231 - 236.

Zapparoli, M., 1990b. — Chilopodi di ambienti urbani e suburbani della citta di Roma. Boll. Ass.Romana Entomol.,
44 : 1 - 12.

Zych. M., 1989. — Uwagi o wystepowaniu Lithobius microps Meinert (Chilopoda, Lithobiomorpha). Przegl. Zool.,
XXXIII : 332-335.

Source : MNHN, Paris

Contribution a la connaissance des lithobiomorphes
(Chilopoda) de la region palestinienne

Stefan NEGREA *& Zachiu Matic **<t)

* Institut de Speologie “E. Racovitza”, Bucarest, Romania
** Lab. Zoologie. Fac. Biol. Geol., Univ. Cluj-Napoca, Romania

RESUME

Se basant sur le materiel de lithobiomorphes Henicopidae et Lithobiidae rapporte d'Israel en 1990 par S. Negrea et
coll., les auteurs presentent dc nouvelles donnees systematiques, ecologiques et zoog6ographiques concernant les
especes apparlenant aux genres Lamyctes, Eupolybothrus et Monotarsobius. Une espece nouvelle : Monotarsobius
teldanensis n. sp. est decrite. Le travail s'ach^ve par des remarques sur la faune de lithobiomorphes de la region
palestinienne.

ABSTRACT

Contribution to the knowledge of the Lithobiomorpha (Chilopoda) in the Palestinian region.

Based on the material of fam. Henicopidae and Lithobiidae (Lithobiomorpha) collected in Israel during the year 1990
by S. Negrea and others, the authors present some new systematical, ecological and zoogeographical data concerning
the species belonging to the genera Lamyctes, Eupolybothrus and Monotarsobius. A new species. Monotarsobius
teldanensis n. sp. is described. The paper also contains general remarks on the fauna of Lithobiomorpha from the
Palestinian region.

INTRODUCTION

La region palestinienne peut etre delimitee par la cote mediterraneenne a l'Ouest. la
Peninsule Arabique a l'Est, le Mont Hermon au Nord et le golfe Elat-Aqabah au Sud. Les
chilopodes de cette region geographique sont encore insuffisamment connus. Les premieres
contributions ont ete publiees dans la periode 1893-1934. particulierement par ATTEMS, PORAT,
SlLVESTRl et Verhoeff, synthetisees ulterieurement par BODENHEIMER (1937). Selon ce
dernier auteur, cinq especes de lithobiomorphes ont ete signalees jusqu'en 1937 dans la region
palestinienne : Archilithobius carinatus Koch (=macrops Karsch), Lithobius parvicomis Porat,
Lithobius vosseleri Verhoeff, Monotarsobius barbipes Porat et Polybothrus fasciatus Newport.
La systematique actuelle ne reconnaTt parmi elles que quatre especes et une sous-espece : 1)
Lithobius carinatus L. Koch 1862 - (syn. Lithobius macrops Karsch, 1888); 2) Lithobius
parvicomis (Porat, 1893) ; 3 ) Hessebius barbipes (Porat, 1893) - (syn. Lithobius vosseleri
Verhoeff, 1901) ; 4) Eupolybothrus litoralis (L. Koch, 1867) - (syn. Lithobius fasciatus sensu
Porat, 1893 non Eupolybothrus fasciatus (Newport, 1844) et Polybothrus fasciatus graecus

Negrea. S. & Matic, Z., 1996. — Contribution a la connaissance des lithobiomorphes (Chilopoda) de la region
palestinienne. hr. Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin. M., (eds), Acta Myriapodologica. Mem.
Mus. natn. Hist, nat ., 169 : 225-233. Paris ISBN : 2-85653-502-X.

226

STEFAN NEGREA & ZACHIU MATIC

var . fasciatograecus Verhoeff, 1901. C'est ZAPPAROLI (1991) qui a apporte la plus recente et
importante contribution a la connaissance des chilopodes de cette aire de grand interet
biogeographique. II a ajoute les 5 especes supplemental suivantes : Harpolithobius halophilus
Verhoeff, 1941, Lithobius viriatus Sseliwanoff, 1879, Lithobius erythrocephalus C. L. Koch,
1847, Monotarsobius crassipes L. Koch, 1862et Monotarsobius bolognai Zapparoli, 1991.

Dans le cadre des programmes diriges a Jerusalem par le Professeur F. D. POR (cl.
POR, 1975 ; POR et al., 1995), l'un de nous (S. N) a recolte, en mai-juin 1990, des chilopodes
d'Israel et il’lui a ete confie pour etude le materiel conserve dans les collections zoologiques des
Universites de Tel Aviv et de Jerusalem, pour la redaction de sa contribution dans le volume de
la serie Fauna Palaestina. Parmi ce materiel, seuls les lithobiomorphes Henicopidae et
Lithobiidae appartenant aux genres Lamyctes , Eupolybothrus et Monotarsobius, ont ete etudies
a ce jour ; les resultats sont publies dans ce travail. Les genres Hessebius et Lithobius s. str. plus
riches en especes, feront l'objet d'autres publications (NEGREA & MATIC, 1995).

LES ESPECES IDENTIFIES

Les donnees pour chaque espece comprendront les synonymies, le materiel examine, la
description des especes ou des populations nouvelles, la redescription des especes
insuffisamment decrites, les notes morphologiques complementaires des especes insuffisamment
connues et les remarques eventuelles.

Les differentes collections sont designees par les abreviations suivantes : HUJ = Hebrew
University of Jerusalem, Zoological Museum; UTA = Tel-Aviv University, Zoological Museum.

Fam. HENICOPIDAE Pocock, 1901

iMmyctes coeculus (Brolemann, 1889) (Figs. 1-3)

Synonymies: Lithobius coeculus Brolemann, 1889 ; Lamyctes coeculus Attems, 1908;
Lamyctes coeculus Silvestri,1909.

' Materiel etudie: 4 92 maturus senior, plantation irriguee de peupliers pres du Kibbutz
Gonen (Vallee de Hula, Galilee superieure, alt. +67 m), sol alluvial couvert de feuilles mortes et
de debris de vegetaux, temperature du sol 28°C, 28.5.1990, leg. S. NEGREA et C.
DlMENTMAN, HUJ ; 1 9 maturus senior, plantation irriguee de pamplemousse pres du Kibbutz
En Gedi (Oasis En Gedi sur le bord ouest de la Mer Morte, alt. -370 m), sol melange, debris
vegetaux et fragments calcaires, temperature du sol 26°C, 2.6.1990, leg, S. NEGREA, HUJ.

Redescription. La description de BROLEMANN (1930) etant tres sommaire, nous
presentons les caracteres des cinq femelles examinees. Longueur des adultes : 4,3 - 5,2 mm.
Coloration jaune-pale, avec les extremites (capsule cephalique, forcipules et tergites 8-16)
intensement orangees. Corps a bords paralleles, elances. Teguments luisants, linement reticules ;
pilosite plus dense sur les tergites 13-16. Une paire de stigmates sur le premier segment. Tete
attenuee en avant, plus longue que large (Fig. 1 ) ; sillon frontal evident ; le bord caudal rectiligne.
Antennes tres courtes, pileuses, formees de 24+24 ou 24+18 articles courts, dont le dernier est
presque egal aux deux precedents, en forme de massue (Fig. 2). Pas d'ocelles. Organe de
Tomosvary indistinct. Bord rostral du coxosternum forcipulaire (Fig. 3) preeminent portant 2+2
dents plus ou moins noires de dimension moyenne et peu aiguisees ; 1 + 1 dents rudimentaires
exlernes, ecartees des autres, a l'aspect de courte epine, placees exactement dans la position des
porodontes. NEGREA (1989) a nomme ces denticules “pseudoporodontes” et non “porodontes”,
comme les designe ZALESSKAJA (1978). Des soies longues pres du bord rostral. Tergites a
angles arrondis ou droits ; bord caudal quasi droit ou un peu echancre pour les tergites 8, 10, 12,
14 et 16. Pores coxaux ronds et relativement grands, au nombre de 1 a 2 pour chaque hanche (1,
2, 2, 2 ou 2, 2, 2, 2). Pattes totalement depourvues d'epines mais avec un prolongement
acumine a l'extremite du tibia des PI a PI 1, sur la face anterieure. Pattes 1-12 avec tarse soude
au metatarse ; il est independant seulement des P13 aux PI 5. La patte 15 est plus longue que la

Source :

LITHOBIOMORPHES DE LA REGION PALESTINIENNE

227

patte 14, mais ne depasse pas la longueur de l'antenne. Griffe apicale des pattes 1-15 flanquee de
chaque cote d une griffe accessoire plus petite. Appendices genitaux des femelles armes de 2+2
eperons comques, relativement longs et pointus. La griffe est nettement delimitee a la base
courte, tres arquee et sans dentelures laterales. Pas d'epine dorsale sur les trois articles du
gonopode.

Remarques. Selon BROLEMANN (1930)
il s’agit d'une “espece exotique (Australie)
importee, decrite des serres de Lombardie et
egalement acclimatee dans celles du Museum
National d'Histoire Naturelle de Paris”. La
repartition de L. coeculus reste encore peu
connue et discontinue. ENGHOFF (1975)
indique Australie, Hawaii, Mexique, Tanzanie
et des serres d’Europe (Danemark, Finlande,

France, Italie et Suede). D’apres NEGREA
(1977), a Cuba celle-ci peut etre consideree
comme une forme edaphophile, les o’er et 99
etant plus frequemment rencontres dans le sol
des forets et sous les pierres plutot que dans la
litiere (100-750 m alt.) ; elle a ete trouvee avec
Lamycles fulvicornis de maniere
exceptionnelle. 11 s'agit d'un genre et d'une
espece nouveaux pour la region palestinienne,
rencontres par Fun des auteurs seulement
dans les plantations (done probablement
importes!) et representes uniquement par des
femelles (populations parthenogenetiques?).

012

0.06

Figs. 1-3. — Lamyctes coeculus (Brolemann), 9:1, Tete ;
2, Demiers articles de l'antenne ; 3, Bord rostral du
coxostemum forcipulaire (mesures en mm) (orig.
S. Negrea).

Figs 1-3. — Lamyctes coeculus ( Brolemann ), 9 : 1, Head;
2, Last antennal articles; 3, Rostral edge of the
forcipular coxosternum (in mm) (after S. NEGREA).

Fam. LITHOBIIDAE Newport 1844

Eupolybothrus (Eupolybothrus) litoralis (L. Koch, 1867) (Figs. 4-12)

Synonymies : Lithobius litoralis L. Koch, 1867 (nec Muralewitsch, 1906); ? Polybothrus
fasciatus Porat, 1993 (citee pour la localite Ain, Couffin); ? Lithobius (Polybothrus) fasciatus
graecus vai \ fasciatograecus Verhoeff, 1901 (citee par VERHOEFF, 1925, des environs de Jaffa
et du lac Tiberiade = Kinneret) ; Lithobius praecursor Attems, 1902 ( nec Lithobius ankarensis
praecursor Verhoeff, 1943, decrit de Beyrouth); Eupolybothrus litoralis Eason, 1970
(redescription dapres 1'holotype et les paratypes); Eupolybothrus litoralis Zapparoli, 1991 (citee
des localites : Mont Hermon et Irbid).

Materiel etudie : 1 <J maturus senior , Nahal Bezet (= Wadi Qarzkara, Galilee superieure),
8.4.1950, UTA; 1 d maturus senior , Zikhron Ya'aqov (Monts Carmel), 21.1.1946, UTA.

Description de la population palestinienne. Etant donne que EASON (1970) a publie une
redescription de cette espece en utilisant la serie type de la Grece et que la population
palestinienne n'a pas encore ete decrite, il parait utile de presenter les caracteres des deux males
examines. Longueur : 30-32 mm. Coloration jaune-marron ; pattes et antennes presque jaunes.
Tete (Fig. 4) plus large que longue. Antennes tres longues, depassant la moitie du corps,
formees de 46 a 53 articles plus ou moins allonges. Ocelles (Fig. 5) de 14 a 16, en quatre
rangees subrectilignes (1+4, 4, 4, 3 ; 1+4, 4, 4, 2). Organe de Tomosvary rond et tres petit,
situe tout pres des ocelles. Coxosternum forcipulaire court; bord rostral (Fig. 6) tres large,
rectiligne, a echancrure mediane petite, arme de 7+7 dents petites ; pas d'epines externes
(porodontes) spiniformes, mais des soies habituelles. Angles des tergites 1, 3, 5 arrondis ; ceux

228

STEFAN NEGREA & ZACHIU MATIC

des tergites 2, 4, 8, 10, 12, 14 presque droits avec le bord caudal plus ou moins echancre ; des
prolongements aux tergites 6 et 7 (courts, larges et emousses) et aux tergites 9, 11 et 13
(triangulaires, aigus) ; tergite terminal a angles faiblement arrondis et a bord caudal peu ou non
echancre, portant de nombreuses soies (Fig. 7). Pores coxaux (Fig. 8) nombreux (de 23 a 32),
ronds ou non, de dimensions variables, irregulierement disposes, dont l'un distal, relativement
isole et bien dessine, toujours plus grand que les autres.

2.5

FIGS. 4-12. — Eupolybothrus litoralis (L. Koch), <? : 4, Tete; 5, Ocellcs et organe de Tomosvary ; 6, Bord rostral du
coxosternum forcipulaire ; 7, Tergites ; 8, Hanche droite de la P14 ; 9, Grifles apicales de la P15 (droite). vue
dorsale ; 10, Femur de la P15 (gauche), vue dorsale-interne ; 11, Articulation tarso-m<$tatarsienne de la P14
(gauche), vue ventrale-inteme ; 12, Sternite genital et appendices genitaux du male (mesures en mm) (orig. S.
Negrea).

FIGS 4-12. — Eupolybothrus litoralis (L. Koch), (f : 4, Head; 5, Ocellae and Tomosvary organ; 6, Rostral edge of the
forcipular coxosternum; 7, Tergites; 8, Right coxa of the PI4; 9, Apical claws of the PI 5 (right), dorsal view;
10, Femur of the PI 5 (left), dorsal-internal view; 11, Tarso-metatarsial joint of the P14 (left), ventrale-internal
view; 12, Genital sternite and genital appendix of the male (in mm) (from S. NEGREA).

Les pattes 14 et 15 sont tres longues (specialement les P 15 qui depassent de beaucoup la
moitie du corps) et relativement greles. Pas d'epines coxolaterales. Griffe apicale secondaire
interne des P15 depassant le 1/3 de la principale (Fig. 9). Les pores glandulaires des PI 5 sont
concentres sur la face interne des femur, tibia, tarse et metatarse. La pilosite du metatarse des
PI 5 s'etend en longueur jusqu'aux 3/4 du diametre de cet article. La serie de soies (“seriate
setae”) du metatarse des PI 5 est absente. La fossette dorso-basale du femur des P 15 du male est
grande et profonde ; le sillon interne qui la prolonge atteint ou non le bord de la nodosite dorso-
distale interne ; cette nodosite, assez proeminente et toujours uniformement arrondie, porte une

Source :

LITHOBIOMORPHES DE LA REGION P ALESTIN IENNE

229

wfTl * °Vate re ? ?ent petlte comP°see cie pores glandulaires et de nombreuses soies
fines et tres courtes . le sillon externe est tres mince et long ; le femur presente de meme une

Sr>f> fr,°Xin;dle dC ?ngUeS soies <Flg- 10). Les soies et les epines de la face ventrale-inteme de
1 articulation tarso-metatarsienne des P14 sont representees Figure 11.

Spinulation des pattes (ventrale/dorsale)

P H tr P

• - - amp

2-13 - - amp

14 - m amp

15 - m amp

amp

am-

amp

am-

amp

am-

-m-

amp

a—

amp

a-p

amp

a- P

-p

amp

—

Appendices gemtaux d1 (Fig. 12) biarticules, longs, greles et pileux ; bord rostral du
stemitc genital a echancrure medtane petite et soies marginales nombreuses.

Remarques. Cette espece, nord-mediterraneenne et orientale, est connue avec certitude de
la Grece (y compos les lies de la Mer Egee - d'oii elle a ete decrite par L. KOCH), du sud-ouest
e la lurquie, de Crete, Syrie, Liban, Jordanie et Israel. Nous ajoutons ici pour Israel deux
stations nouvelles, 1 une en Galilee superieure (Nahal Bezet) et 1'autre au Mont Carmel (Zikhron
Yaaqov). - v

Le materiel d'Israel a ete compare a des exemplaires de 1'tle Kassos (Grece) empruntes par
Marzio ZAPPAROLI et a la redescription de cette espece faite par EASON (1970) /Fapres la serie
type des ties egeennes de Tinos et de Rhodes (Grece). Les differences minimes constatees
peuvent etre considerees comme des variations individuelles qui caracterisent les populations
palestimennes. Ces dilferences sont les suivantes : longueur du male “ maturus ” d'Israel • 30-35
mm (materiel de Kassos : 30-31 mm ; materiel de Tinos et de Rhodes : 28-38 mm) ; ocelles • 14-
16 en 4 rangees (12-14 en 3-4 rangees ; 16-18 en 4 rangees) ; bord rostral : 7+7 petites dents
( /+/ ou S+8 ; 8+8) ; pas de porodontes (idem; “lateral spines peg-like”) ; pores coxaux : 23-32
lrreguherement disposes (18-27 irregulierement disposes ; 30-55 en 4-5 rangees) ; un pore coxal
distal plus grand et isole (idem ; pas de mention) ; sillon externe du femur des PI 5 long et mince,
allant jusqu a 1 extremite apicale de cet article (idem; “finer, extending to the margin of the pore-
tree area ) , nodosite dorso-distale interne du femur des P15 toujours uniformement arrondie et
assez proeminente, portant une aire centrale ovale de pores et de soies fines (idem ; “is barely
swollen avec 1'aire centrale circulaire-ovale, sans soies!) ; epines de la P14 ■ DaT absente
(presente ; presente) ; epines de la P15 : VpF, VaT et DpT absentes (VpF et DpT absentes VpF
VaT et DpT presentes). r ’

Eupolybothrus litoralis est a rapprocher de E. fasciatus (Newport, 1844) connue avec
de Florence et de Naples (terra typica - voir EASON, 1970 et MlNELLI & ZAPPAROLI,
, -), mais certains caracteres separent les deux especes. II s'agit surtout de : “seriate setae” du
metatarse des PI 5 (E. litoralis : absente; E. fasciatus : presente), sillon externe du femur des
1 15 (present ; absent) ; touffe proximale du femur des PI 5 avec (des soies fine et longues ; des
soies grossieres et plus courtes) ; nodosite dorso-distale-interne du femur "des PI 5
(umloimement arrondie et relativement proeminente ; non-uniformement arrondie a cause de
1 aire centrale ovale et mamillaire et fortement proeminente).

Monotarsobius teldanensis n. sp. (Figs. 13-17)

Materiel etudie : 1 cf maturus senior (holotype), reserve naturelle du bassin superieur de la
riviere Tel Dan (Galilee superieure, alt. +190 m), sol calcaire de la foret couvert de feuilles
mortes, debris vegetaux, bois pourri et pierres, temperature du sol 19,5 °C, 28.5.1990. leg. S.
NEGREA, HUJ (lieu de conservation de F holotype).

Derivatio nominis ; du nom Tel Dan (terra typica).

230

STEFAN NEGREA & ZACH1U MAT1C

Description de l’holotype (tf) : Longueur : 11,2 mm. Coloration jaune-paille. Teguments
unis, brillants. Tete un peu plus large que longue, sans ponctuations distinctes, a bord caudal
presque rectiligne et a bourrelet etroit et sans sinuosites. Antennes tres courtes, de 20 articles,
dont le dernier est environ le double du precedent (Fig. 13). Ocelles : 1+3 " pratiquement de la
meme dimension, disposes en croix (Fig. 14). Organe de Tomosvary plus petit qu'un ocelle.
Coxosternum forcipulaire a bord rostral large, divise par une encoche profonde et arme de
dents robustes, relativement rapprochees ; 1+1 porodontes spiniformes epais et longs (Fig. 15).
Tous les tergites ont les angles arrondis et le bord caudal droit ou plus ou moins echancre , pas
de prolongements triangulaires posterieurs (Fig. 16). Pores coxaux : 3, 3, 3, 3 - petits et ronds.

FIGS. 13-17. — Monoiarsobius teldanensis n. sp., <f : 13, Derniers articles de l'antenne ; 14, Ocelles et organe de
Tomosvary ; 15, Bord rostral du coxosternum forcipulaire ; 16, Tergites ; 17, F6mur de la P.15 (gauche), vue
dorsale (mesures en mm) (orig. S. Negrea).

FIGS 13-17. — Monotarsobius teldanensis n. sp., <? : 13, Last antennal articles; 14, Ocellae and Tomosvary organ ; 15,
Rostral edge of the forcipular coxosternum; 16, Tergites; 17, Femur of the PI 5 (left), dorsal view (in nun) (from
S. Negrea).

Figs. 18-19. — Monotarsobius bolognai Zapparoli, 9:18. Appendice genital (gauche) ; 19, Ocelles et organe de
Tomosvary (mesures en mm) (orig. S. Negrea).

Figs 18-19. — Monotarsobius bolognai Zapparoli. 9 : 18. Genital appendix (left); 19, Ocellae and Tomosvary organ (in
mm) (from. S. NEGREA).

L'articulation tarso-metatarsienne des PI - P12 absente, celle de la P13 peu distincte et
celles des PI 4 et P15 distinctes et fonctionnelles. Les deux demieres pattes plus epaisses que les
autres, ayant leur face interne criblee de pores. Pas d'epines coxolaterales. Griffe apicale
secondaire des PI 5 absente. Le femur des P 15 a sillon dorsal peu profond (Fig. 17). Spinulation
des pattes (ventrale/dorsale) :

—

am-

-m-

-p

a-p

2-6

-m-

am-

-m-

-P

a-p

7-10

-mp

am-

a-p

1 1

-mp

amp

am-

amp

a-p

-mp

amp

am-

amp

-p

—

-mp

amp

am-

-mp

-p

—

amp

-mp

—

-mp

—

amp

-mp

—

-mp

—

Source : MNHN , Paris

LITHOBIOMORPHES DE LA REGION PALESTIN IENNE

231

apicale 6 premier sternite genital avec 5+6 soies ; le deuxieme arrondi et pourvu dune soie

.Difgnosf* 1 !’2 mmde longueur ; 20 articles antennaires, 1+3 ocelles disposes en croix ■
1+- dents au bord rostral du coxosternum ; 1 + 1 porodontes epais et longs ; 3 3 3 3 pores
coxaux ; epines coxolaterales absentes ; griffe de la PI 5 simple ; femur de la P15 du male avec
un sillon ; spinulation assez riche pour un Monotarsobius (voir au-dessus).

Diagnose comparative. Monotarsobius teldanensis n. sp. est connu uniquement de la
localite type. II se rapproche de M. schizus decrit par CHAMBERLIN en 1952 de Turquie
(uniquement dapres le male) mais les caracteres suivants distinguent les deux especes : nombre
des ocelles (1+3 au lieu de 1+3, 2) ; encoche du bord rostral (en forme de V et non de U) ■ forme
de porodonte (spiniforme, epais et long - et non spiniforme, court et grele) ; nombre des pores
coxaux (3, 3 3 3, au lieu de 2, 2, 2, 2) ; petit lobe distal du tibia des PI 5 absent chez M t. et
piesent chez M. 5.) ; spinulation (PI : 00021/00122 et non 00000/00001 ; P14 ■ 01320/00^00 et
non 01 32 1/002 00 ; PI 5 : 01320/10200 et non 01320/10210). Une comparison plus
approfondie entre M. teldanensis et M. schizus est toutefois difficile a etablir du fait que
Chamberlin a decrit incompletement son espece, sans aucune illustration.

Monotarsobius bolognai ZapparoliT 1991 (Figs. 18-19)

Materiel dtudie : 1 9 maturus senior , reserve naturelle Shemurat haMasreq (Monts de
Judee, alt. +600 m), sol calcaire de la foret couvert de feuilles mortes et de bois pourri
temperature du sol 16°C, 12.5.1990, leg. S. NEGREA, HUJ. '

Notes morphologiques : la femelle de 7.5 mm de longueur correspond a la diagnose et a la
description (spinulation incluse) de ZAPPAROLI (1991). On observe cependant quelques
ditterences : organe de Tomosvary de dimension intermediaire entre celle de l’ocelle posterieur et
celle de 1 unique ocelle anterieur (Fig. 18), articulation tarso-metatarsienne des PI - 12 absente
trace d articulation non fonctionnelle aux PI 2 et PI 3, gonopode a 4+4 eperons et a griffe etroite
et pointue, pourvue dune dentelure externe et d’une epine dorsale grele, deux epines dorsales
sur le second article du gonopode (Fig. 19).

Remarques : M. bolognai est connue seulement de trois stations de la region palestinienne.
relativement anthropisees et a substrat calcaire : Wadi Kafrein (versant oriental de la depression
e a Mer Morte), Wadi Tajiba et Jaar na Nabi (Jerusalem). Une nouvelle s'ajoute aux deux
premieres, Shemurat haMasreq, qui est une foret de Quercus callyprinos, Pistacia, Arbutus et
t'mus non irriguee, mstallee sur roche calcaire, pres de la localite Bet Meyr.

Monotarsobius crassipes L. Koch, 1862

Materiel etudie : 1 9 maturus senior (11 mm de longueur), 2 <f <? praematurus (6,5-
/,5mm) et 3 & <f immaturus (4, 5-5, 8 mm), plantation irriguee de peupliers pres de kibbutz
Gonen ( Vallee de Hula, Galilee superieure. alt. +67 m), sol alluvial couvert de feuilles mortes et
de debris vegetaux, temperature du sol 28°C, 28.5.1990. leg. S. NEGREA, A. NEGREA et I.
CAPUSE, HUJ ; 2 cf cf pseudomaturus (7, 5-8,2 mm), 1 cf praematurus (6,4 mm) et 1 o'
immaturus (5,3 mm), grotte Me’arat Sharakh (Vallee Nahal Sharakh, Galilee superieure, alt.
+300 m), sur le plancher calcaire avec sol argileux et dechets abandonnes par les visiteurs de la
grotte. 6.6.1990, leg. V. DECU et C. DlMENTMAN, HUJ.

Notes morphologiques : le o' et la seule 9 examines correspondent aux caracteristiques
signalees par EASON (1964) dans sa description et ses dessins.

Remarques : espece palearctique occidentale (Europe, Afrique du Nord, Proche et Moyen
Orient et Asie Centrale). Dans la region palestinienne, l’espece est connue du Liban (Becherre),
de la Jordanie (Petra) et d’ Israel (Allone Abba, Segev et Basmat Tivon) (ZAPPAROLI, 1991). A
ces stations, nous en ajoutons deux d'Israel : une grotte et une plantation de peupliers de la
Galilee.

232

STEFAN NEGREA &ZACHIU MATIC

CONCLUSIONS

L'etude du materiel ci-dessus revele la presence de cinq especes : Lamyctes coeculus
(Brolemann, 1889). genre et espece nouveaux pour la region palestinienne ; Monotarsobius
teldanensis, espece nouvelle pour la science ; Eupolybothrus litoralis (L. Koch, 1867),
Monotarsobius bolognai Zapparoli, 1991, et M. crassipes L. Koch 1862, ces trois detniers ont
deja ete citees de la zone d’etude mais recoltees dans des stations nouvelles. Ainsi. le nombre des
lithobiomorphes connus de la region palestinienne est passe de 9 a 1 1 especes. Du point de vue
zoogeograph ique, celles-ci peuvent etre rattachees a differentes categories. Le genre Lamyctes a
un seul representant, L. coeculus, element holotropical a repartition discontinue, qui fut
sommairement decrit par BROLEMANN (1889). II est redecrit d’apres des echantillons d’Israel,
peut-etre issus de femelles parthenogenetiques (?) importees. Le genre Eupolybothrus est
egalement represente par une seule espece, E. litoralis (L. Koch), qui est, d apres ZAPPAROLI
(1991), un element nord-mediterraneen et oriental. Le genre Hessebius a deux representants :
H. barbipes (Porat) qui est une espece est-mediterraneenne touranienne, tres frequente dans la
region palestinienne et H. halophilus Verhoeff dont la geonemie est nord-meditenaneenne
(ZAPPAROLI. 1991). Le genre Monotarsobius est represente par trois especes, toutes recoltees
par les auteurs en Israef : M. crassipes L. Koch, espece palearctique occidentale qui a ete
capturee dans une grotte et dans une plantation de peupliers (nombreux adultes et juveniles) ; M.
bolognai Zapparoli, endemique de la region palestinienne (cantonnee particulierement dans la
zone de la Mer Morte) et apparentee a des especes siberiennes (ZAPPAROLI, 1991) ; M.
teldanensis n. sp., probablement endemique dans cette region et apparentee a M. schizus
Chamberlin, connue de la Turquie. Enfin, le genre Lithobius s. str., qui sera etudie dans un
prochain travail, a, pour le moment, quatre representants : deux especes nord-mediterraneennes
orientales (L. carinatus L. Koch et L. parvicornis Porat), une sud-est europeo-anatolico-
caucasienne (L. viriatus Sseliwanoff) et une palearctique occidentale (L. erythrocephalus L.

Koch). „ . ,

La faune des Lithobiomorphes de la region palestinienne se revele ties mteressante et
remarquable par les especes endemiques et d'origine mediterraneenne qui constituent la majorite
de la faune. La diversite specifique n'est pas elevee. Les stations occupees par des chilopodes
corespondent aux biotopes naturels proteges dans des reserves naturelles (par exemple Tel Dan
et Shemurat haMasreq) ou a ceux des vallees boisees traversees par des rivieres (par exemple
Nahal Sharakh). Dans les biotopes anthropises (telles les plantations irriguees de Gonen et En
Gedi) la diversite est moindre, mais la densite des indi vidus plus grande.

REMERCIEMENTS

Nous adressons nos vifs remerciements a M. le Professeur F. D. Por (Jerusalem) pour son aimable invitation a
etudier sur place les chilopodes et pour I'aide qu'il nous a apportec en Israel, a M. le Docteur M. Zapparoli (Viterbo) pour
le materiel de comparison d 'Eupolybothrus litoralis et E. fasciatus qu’il nous a confie, ainsi que pour son aide
bibliographique et ses informations precieuses.

REFERENCES

Bodenheimer, F. S., 1937. — Prodomus faunae Palestinae. Mem. Inst. Egypte,33 : 233-234.

Brolemann, H. W., 1889. — Contributions & la faune myriapodologique mediterraneenne. Trois especes nouvelles.
Ann. Soc. Linn. Lyon : 5-16.

Brolemann, H. W., 1930. — Elements dune faune des Myriapodes de France. Chilopodes. I Faune de France , 25]. Paris,
P. Lechevalier. 1-405

Chamberlin, R. V., 1952. — On the Chilopoda of Turkey. Rev. Fac. Sci. Univ. Istanbul, ser. B. Sci. Nat., 17 : 183-
258.

Eason, E. H., 1964. — Centipedes of the British isles. London, F. Warne & Co, 294 pp.

Eason, E. H., 1970. — A redescription of the species of Eupolybothrus Verhoeff s. str. preserved in the British Museum
(Chilopoda, Lithobiomorpha). Bull. British Mus. London. 19 : 289-310.

Source : MNHN, Paris

LITHOBIOMORPHES DE LA REGION PALESTINIENNE

233

EN (SSK*: Sex ss* ,Br6lemann) 3 cosmopo,ilic parthenogenetic

M,NSwg *^P16OLI21NI-243992' ~~ Considerazioni faunistiche e zoogeografiche sui chilopodi delle alpi occidental!.

NK<MEc’ SUir laP69Sn4e dC Lamyc,es Wvicomis Meinert. 1868 (Chilopoda : Henicopidae) en Roumanic.

Negrea, S., 1977 Considerations ecologiques et biogeographiques sur les Chilopodes de Cuba. Res. Exped Biosp
Cubano-roumatnes a Cuba, 2 : 303-312. 1

NEGiRREcA\Srrt-,MA aC' Z,' “mpara,lve de la variabili‘e de trois populations de Hessebius barbipes (Porat,

1893) (Chilopoda : Lithobndae). Soil Fauna of Israel, 1 : 29-39. 1

Por, F. D.. 1975. — An Outline of the Zoogeography of the Levant. Zool. Scripta, 4 : 5-20.

Por F. D„ Decu V., Negrea, S. & Dimentman. C. 1995 — A survey of the edafic fauna of Israel. Results of a
Romaman-Israeli joint collecting programme (May-June, 1990). Soil Fauna of Israel, 1:1-17.

Porat, C. O., 1893 . — Myriapodes recoltes en Syrie par le Docteur Theodore Barrois. Rev. Biol. Nord France. 6 : 62-
Verhoeef. K. W., 1925. — Mediterranean Chilopoden und Notiz zur Periodomorphose der Juliden. Zool. Anz., 64 : 61-

oU.

ZAL2nKppA’ N* T ' 1978’ ~ 0prede,ile,i mn°g°nojek kostyanok SSSR (Chilopoda, Lithobiomorpha). Moscou, Nauka,
Zapparoli, M., 1991. Note su alcune specie di Chilopodi della regione palestinese. Fragm. Entomol. , 23 : 15-33.

Source : MNHN, Paris

Check-List, Distribution and Habitat in Bulgarian

Centipedes

Georgi Rl BAROV

Natural History Department of Regional Museum, 2 Dzaldeti Str.. BG-8600 Jambol. Bulgaria

ABSTRACT

mate d af'exam ined bvThe °Llt cen n p^de '«una “ based on bibliographic data, collections and recent

" 1 exammed °y lhc author. The present check-list includes 26 species of Geophilomorpha 5 species of
Scolopendromorpha, 67 species o I Lithobiomorpha and 1 species of Scutigeromorpha. Subspecies are^ot listed because
ot their uncertain status. The district and altitudinal distribution of the centipedes is shown in Table 1 42% of all the

°^a,kan^demJ?- M°Sl endemic ^ « known from Rhodipi
U//o) and btrandza Mts. (35 /o). Some species ( Thracophilus bulgaricus Verhoeff. T. beroni Matic & Darabantzu) show
the connections between the Bulgarian (respectively Balkan) fauna with that of Asia Minor.

RESUME

Les Chilopodes de Bulgarie : Iiste des especes, repartition et habitat.

Cet inventaire prehminaire de la faune des chilopodes de Bulgarie est base sur les donnees de la litterature sur les

r6ne,^“ an3,yS6 Par raUleUr* La ,islC deS esp^ces inclul 26 Geophilomorpha.
cause du caract^^mn ?nr.r.l H°T0rP f el ,IScul,2ieromorPha- Les sous-especes ne sont pas prises en consideration a
l P Cerla.,n de leur stalul- L a,re de repartition et la distribution verticale sont donnees par le Tableau

. 42 /o de toutes les especes repertories sont des endemiques de Bulgarie ou des Balkans. Le plus grand nombre d*especes

ThraZZllTf. ‘/0UVe v h* ^0nIf. Rhod°Pe (37%) el dans les Monts Strandza (35%?. Cenaines especes (Liles

de ■■ f™

INTRODUCTION

.. ,The foi™atlon of the present day Bulgarian centipede fauna can be generally divided into
three historical periods: Tertiary, Pleistocene and Postglacial (GRUEV, 1981). At the end of the
Miocene and at the beginning of the Pliocene, the tropical and subtropical climate in South
-ui ope and in the Balkan peninsula changed to more temperate conditions. At the same time
Bulgaria fell under the influence of the Central European fauna from the north and Asiatic fauna
from the north-east.

The second main period in the formation of the Bulgarian centipede fauna began during the
Pleistocene when North America, Europe and Siberia were covered with an ice belt. Under the
influence of this cold spell, the climate changed in the Balkan peninsula (including Bulgaria).

Ribarov. G., 1996. — Check-list, distribution
Mauri£s. J.-P. & Nguyen Duy - Jacquemin, M.. (eds),
241. Paris ISBN : 2-85653-502-X.

and habitat in bulgarian centipedes. In: Geoffroy, J.-J.,
Acta Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 235-

236

GEORGI RIBAROV

The glaciation enveloped the high Bulgarian Ryla and Pirin mountains. In the lower
southern mountains: Strandza, Sakar and the eastern parts of the Rhodopi Mts., the influence of
the cold spell was on a lesser scale. Many species of Chilopoda have found here more
favourable conditions.

TAXONOMIC OBSERVATIONS

Contributions to the study of the centipedes (Chilopoda) of Bulgaria have been made by
several authors (JURINICH. 1904; VERHOEFF, 1926-1928; FOLKMANOVA. 1936; MATIC, 1964-
1973b- MATIC & DARABANTU, 1968; MATIC & GOLEMANSKY, 1964-1967c; NEGREA, 1965.
1971; KACZMAREK. 1969a, 1975; RIBAROV, 1984-1992). In 1936, FOLKMANOVA recorded
the species Scolopendra morsitans (L.) from Strandza Mts., Southeastern Bulgaria. During the
period 1980-1984, the present author collected numerous specimens of the genus Scolopendra in
Southeastern Bulgaria and examined them. On the base of this and the examination of the
collections of Chilopoda in the Natural History Museum, Sofia, RIBAROV (1984) considered
that probably FOLKMANOVA (1936) has wrongly identified some specimens of S. cingulata
Latreille as S. morsitans and in fact the last species does not occur in Bulgaria. On the other
hand. RIBAROV (1989a) identified the subspecies S. cingulata thracia Verhoeff as a new
synonym of S. cingulata. The lithobiomorphs Eupolybothrus grossipes (C. L. Koch) and
Lithobius borisi (Verhoeff) were also removed from the Bulgarian faunal list (RIBAROV,
1989a). The first one as misidentified by JURINICH (1904) and the second as a new synonym of
Lithobius erythrocephalus ( C . L. Koch).

In the present paper, subspecies are not listed because of their uncertain status. There are
some cases of different subspecies of the same species which reported from the same locality.
Such are: Cryptops anomalans anomalans Newport and C. a. schassburgensis Verhoeff, C.
parisi parisi Brolemann and C. p. rhenanus Verhoeff. The taxonomic status of the subspecies
Pachymerium ferrugineum insulanum Verhoeff also needs an examination, lhe description of
the specimens of P. f insulanum Verhoeff, anounced by KACZMAREK (1969a) from the
Bulgarian Black Sea coast show significant differences (more various than those peculiar for the
subspecies level) with the nominate form P. f ferrugineum C. L. Koch recorded in the same
region (RIBAROV, 1984-1990).

ZOOGEOGRAPHICAL DISTRIBUTION AND HABITAT

To clarify the distribution of the centipedes and their habitat preferences, the Bulgarian
territory is divided here into 7 regions, on the basis of climatic-geographic principles: 1 - Black
Sea coast (BSC), 2 - North Bulgaria (NB), 3 - Thracean region (TR), 4 - Central mountains
(CM), 5 - Southwestern region (SW), 6 - Ryla-Rhodopi region (RR) and 7 - Strandza Mts.
(SM) (Fig. 1 ). The check-list and distribution are presented in Table 1.

The western, higher and more humid parts of the Rhodopi Mts. were included in RR. but
the eastern lower and more arid parts of the same Mts. remains in TR.

42% of all the centipedes established in the country are Bulgarian or Balkan endemics.
Comparing the regions, the largest percentage of endemics is found in the Strandza Mts. (35%)
and in Ryla-Rhodopi region (32%). Somewhat larger is the respective percentage in the Rhodopi
Mts. which fall into neighbouring regions - RR and TR (Fig.l). Some of the Bulgarian endemic
species: Geophilus balcanicus Kaczmarek, Lithobius electron Verhoeff, L. glaciei Verhoeff, L.
rylaicus Verhoeff, L. jurinici Matic & Golemansky, are recorded only from the high parts of the
mountains of the Ryla, Pirin, Stara Planina range and W-Rhodopi. Probably the above
mentioned species belong to isolated communities of an older Euro-Siberian faunistic complex.
On the other hand, in the East-Rhodopi, Sakar Mts. and Strandza Mts. (2/3 of the last extend
south into Turkey), many thermophilic endemic species have evolved and survived. Such are:
Henia angelovi Ribarov, Lithobius maculipes Folkmanova, L. tiasnatensis Matic, L. thracicus

Source :

A BULGARIAN CENTIPEDE SURVEY

237

mmai°a°K^^kL S°,emamk>i Ribar0''- L «™>zmicm Ribarov, Harpoli.hobius

F'G' ’■ ^Trenf Tal rC8TS i"B^!8fa- ' - Black Sca coasl <BSC>- 2 - North Bulgaria (NB). 3 - Thracean region
mountains (SM) (CM)> 5 ’ SouIhwestern region (SW), 6 - Ryla-Rhodopi region (RR). 7 - Strandtza

. m 27,%,°l f11Bulgarian centipedes belong to European and Central European faunistic
elements, but then percentage is not the same in the different regions. In North Bulgaria, these

Mu n2<Tf c thm Central MtS’ 35%- This Percentage is smaller in the Strandza

Wh p!° p Southwestern region (32%). The more thermophilic South European,

hi£ \/iEr,r0pean and, Fr/nl^u '?°Uth Eur°Pean faunal patterns established in Bulgaria are

rerorr Jri f f /o‘ The hlgh/St Percenta§e of Mediterranean faunistic elements were

recorded from the Black Sea coast (26%) and from the Southwestern region (20%). This is

connected with the climatic influence and with the location of the two above mentioned regions
in the path of Mediterranean migrants. The number of Mediterranean species occuring in the

„™dtZa ^tSi Aafd "a t,h,e„TI?raCean regl0n is also hi§h- In sP‘te of 'his, their percentage is
respectively 16 T and 14%, because of the numerous endemic species occurring in the same
i tgions.

The thermophilic endemic species Thracophilus bulgaricus Verhoeff and T. beroni Matic &
Darabantu seem to be good indicators of the connections between the Balkan and the Asia Minor
fauna in Bulgaria. The endemics T. cilicus Attems and T. pachypus Verhoeff are representatives
ol the genus Thracophilus Verhoeff in Asia Minor according to ZAPPAROLI (1990).

Some of the thermophilic species distributed south of the Central Bulgarian mountains
fHelei more humid biotopes rich in vegetation: Harpolilhobius folkmanovae Kaczmarek,
Lithobius beroni Negrea, Pleurolithobius jonicus (Silvestri). Other taxa such as Henia angelovi
Kibaiov and Lithobius peregrinus Latzel occur in more arid and open sites

238

GEORGI RI BAROV

Table 1. — Present knowledge of the centipede fauna (Chilopoda) of Bulgaria - Distribution according to the regions
(BSC-Black Sea coast, NB - North Bulgaria, TR - Thracean region, CM-Central Mts., SW - Southwestern region,
RR - Ryla-Rhodopi region, SM-Strandza Mts.) - Vertical distribution (V.D.) m a.s.I.; Zoogeographic distribution
(Z.D.): en - Endemic, ben - Balkan endemic, il - Illiric, me - Mediterranean, erne - East Mediterranean, se - South
European, see - South-East European, cse - Central-South European, e - European, ce - Central European, t-e-me -
Turano-Euro-Mediterranean, e-a - Euro-Asiatic, co - Cosmopolitan.

Distribution BSC NB

Species

GEOPH 1LOMORPH A
Bothriogaster signala Attems, 1926
Brachyschendyla varnensis Kaczmarek. 1968 +

Clinopodes flavidus C. L. Koch, 1847 + +

Dignathodon nticrocephalum (Lucas. 1846) +

Geophilus balcanicus Kaczmarek. 1972
G. electricus (L., 1758) +

G. flavus (De Geer, 1783) +

G. linearisC. L. Koch, 1835
G. proximus C. L. Koch, 1847
G. rhodopensis Kaczmarek, 1970

G. strictus Latzel, 1880
Henia angelovi Ribarov, 1987

H. bicarinata (Meinert, 1870)

H . illyrica (Meinert, 1870) + +

Himantarium gabrielis ( L., 1767) +

Pachymerium ferrugineum C. L. Koch, 1835 +

P. flavum Folkmanova, 1949 +

Schendyla delicatula Kaczmarek. 1 969 +

S. montana Attems, 1895
S. nemorensis (C. L. Koch, 1836)

5. walachica Verhoeff, 1900
Strigamia acuminata (Leach, 1815)

S. crassipes (C. L. Koch, 1835) + +

S. iranssilvanica (Verhoeff, 1928) +

Thracophilus beroni Matic & Darabantu, 1973

T. bulgaricus Verhoeff, 1926
SCOLOPENDROMORPHA

Cryptops anomalans Newport, 1844

C. croaticus Verhoeff, 1931

C. hortensis Leach, 1815

C. parisi Brolemann. 1920

Scolopendra cingulata Latreille, 1829
LITHOB IOMORPH A

Eupolybothrus andreevi Matic, 1964

E. fasciatus (Newport, 1845)

E. ochraceus (Folkmanova, 1936)

E. transsylvanicus (Latzel, 1882)

E. tridentinus (Fanzago, 1874)

E. valkanovi Kaczmarek, 1973

Harpolithobius anodus (Latzel, 1880) +

H. aseni Kaczmarek, 1975

V. D.

Z.D.

150

erne

100

0-2100

e-a

0- 700

1300-1400

100

50-1000

150-2500

250- 600

150-1800

400- 500

ben

150- 700

250- 600

0-1600

0-1450

0-1100

t-e-me

0- 700

100

500-1300

cse

550

600- 700

ben

150-1000

0-2400

0-1800

450

500- 800

see

0-1800

erne

400-2000

150-1850

0-1100

600

0-1100

100-2350

150-1800

see

150-1600

cse

600

500-1650

see

600

Source :

A BULGARIAN CENTIPEDE SURVEY

239

H. banaticus Matic, 1961

H. folkmanovae Kaczmarek, 1975

H. hemusi Kaczmarek. 1975

H. radui Matic, 1955

Lithobius aeruginosus L. Koch, 1862

L. agilis C. L. Koch, 1 847

L. audax Meinert, 1872

L. balcanicus Matic, 1973

L. beschkovi Matic & Golemansky, 1967

L. bifid us Matic, 1973

L. bulgaricus Verhoeff, 1925

L. burzenlandicus Verhoeff, 1931

L catascaphius Verhoeff, 1937

L. christovici Matic & Golemansky. 1964

L. eras sipes L. Koch, 1 862

L. curtipes C. L. Koch, 1847

L. dalmaticus Latzel, 1880

L. diampolisi Ribarov, 1987

L. dobrogicus Matic, 1962

L dubosequi Brolemann, 1896

L. electron Verhoeff, 1927

L. erythrocephalus C. L. Koch, 1 847

L. forficatus (L., 1758)

L glaciei Verhoeff, 1927
L. golemanskyi Ribarov, 1987
Ljurinici Matic & Golemansky, 1965
L. lakatnicensis Verhoeff, 1926
L lapidicola Meinert, 1872
L. latro Meinert, 1872
L. lucifugus L. Koch, 1 862
L. maculipes Folkmanova, 1936
L. microps Meinert, 1868
L. mutabilis L. Koch, 1862
L muticus C. L. Koch, 1847
L nigrifrons Latzel & Haase, 1880
L. nigripalpis L. Koch, 1867
L. oglednicus Ribarov, 1987
L. parietum Verhoeff, 1899
L. pelidnus Haase, 1880
L. peregrinus Latzel, 1880
L. piceus L. Koch, 1862
L. popovi Matic, 1973
L proximus Matic & Golemansky, 1967
L. pusillus Latzel, 1880
L. pustulatus Matic, 1964
L. ruschovensis Matic, 1967
L. rylaicus Verhoeff, 1937
L. strandzanicus Ribarov, 1987
L. beroni Negrea, 1965
L thracicus Matic & Golemansky, 1967
L. tiasnatensis Matic, 1973

1450

ben

50- 500

900

250

ben

300-1000

300

200- 300

see

700- 800

200- 400

150- 400

0-1800

ben

0- 700

700

400- 500

0-1100

400- 700

see

800- 900

ben

100- 400

200- 600

ben

700-1900

erne

2300

100-2750

0-2500

2200

150- 600

900-1900

250-1000

ben

700- 800

150- 600

0- 800

150- 400

600- 800

100-1400

100- 700

0-1800

150- 400

100-2500

see

300

150- 700

see

200-1800

900

600- 800

700-1400

150- 400

ben

800

2500

cse

150- 400

150-1300

200

300

240

GEORG I RI BAROV

L. totevi Kaczmarek, 1975

1000

L. trebinjanus Verhoeff. 1900

see

L. tricuspis Meinert, 1872

400- 500

L uniunguis Made & Golemansky, 1967

200- 800

L. viriatus Sselivanoff, 1879

150-2100

see

L. vizicae Ribarov, 1987

150- 400

L. zelazovae Kaczmarek, 1975

250-1500

Pleurolithobius jonicus (Silvestri, 1896)

«/T

erne

Lamyctes fulvicornis Meinert, 1868

150- 400

SCUTIGEROMORPHA

Scutigera coleoptrata (L., 1758)

+ 0- 800

CONCLUSION

The present knowledge of the centipede fauna of Bulgaria is obviously incomplete, for that
reason, we suggest the present account has to be considered as a preliminary one. For instance,
more data is necessary to complete the faunal list for North Bulgaria. Some species of Lithobius
found in the Rhodopi' Mts. are new for science and still in process of description. On an other
hand, the taxonomic status of some subspecies and also species is of questionable validity.

From a zoogeographical point of view, the Bulgarian centipede fauna is very rich because
of the geological history, the relief and the crossroad situation of the country. Here are
distributed: Endemic, Central and South European. Holomediterranean and East Mediterranean,
Euro-Asiatic and Balkan-Asia Minor zoogeographical faunistic elements.

The Bulgarian territory can be considered as a refuge for many species which survived the
glacial periods. Moreover, the process of specification took place here in some geographically
isolated populations. 42.4% of the species established in the country are Bulgarian or Balkan
endemics. For the thermophilic species, the most important refuge from the Tertiary times up to
now have been the mountains Strandza, Sakar and eastern parts of the Rhodopi Mts., for the
Euro-Siberian patterns similarly the mountains of the Ryla, Pirin and Stara Planina range.

A considerable influence on the formation of the modern centipede fauna in Bulgaria was
the migrations of the different faunistic elements. The most intensive pathways for the
Mediterranean migrants from south to north across the Bulgarian territory have been the Black
Sea coast and the valleys of the rivers Struma, Mesta, Toundztha, Maritsa and Arda.

The Central Bulgarian Mts. (Stara Planina range, Sredna Gora range, Vitosha) seemed to
have played a role as a natural northern boundary for the thermophilic Mediterranean and South
European centipedes such as Dignathodon microcephalum (Lucas), Eupolybothrus tridentinus
(Fanzago) and Pleurolithobius jonicus (Silvestri), and one of the last locations for the Central
European and Euro-Siberian elements, e. g. Lithobius mutabilis L. Koch which have migrated to
the south.

REFERENCES

FOLKMANOVA, B., 1936. — Ueber einige von Dr. Jaroslav Storkan in Bulgarien gesamraelte Chilopoden. Izv. tsarsk.
Prir. Inst., 9 : 92-97.

Gruev, B., 1981. — General biogeography. Plovdiv : 205-209 (in Bulgarian).

Jurinich, S., 1904. — Contribution h la faune des Myriapodes en Bulgarie. Sbor. Nar. Umotv., 20 : 1-44 (in Bulgarian).
Kaczmarek, J., 1969a. — Beitrage zur Kenntnis bulgarischer Chilopoda. Teil 1. Bull. Soc. Amis scien., Ser. D, 9 : 263-
277.

Kaczmarek. J., 1969b. — Beitrage zur Kenntnis bulgarischer Chilopoda. Teil 11. Bull. Soc. Amis scien., Ser. D\ 10 :
99-109.

Kaczmarek, J., 1970. — Beitrage zur Kenntnis bulgarischer Chilopoda. Teil III. Bull. Soc. Amis scien., Ser. D, 11 :
81-89.

Source : MNHN, Paris

A BULGARIAN CENTIPEDE SURVEY

241

Kaczmarek, J., 1972. — Beitrage zur Kcnntnis bulgarischer Chilopoda. Teil IV. Bull. Soc. Amis scien., Ser. D, 12/13
: 261-264.

Kaczmarek, J., 1973. — Beitrage zur Kenntnis bulgarischer Chilopoda. Teil V. Bull. Soc. Amis scien., Ser. D , 14
181-192

Kaczmarek, J., 1975. — Beitrage zur Kenntnis bulgarischer Chilopoda. Teil VI. Ann. Zool., 33 : 47-66.

Matic, Z., 1964. — Description dun nouveau Lithobiidae cavernicole de Bulgarie (Chilopoda). Ann. Spel. Romania
XIX : 507-510.

Matic, Z., 1967. — Contribution a la connaissance des Chilopodes cavcrnicoles de Bulgarie. Bull. Inst. Zool.
(Bulgarie), XXV : 17-21.

Matic, Z., 1973a. — Lithobius balcanicus n.sp. une nouvelle Lithobiidae (Chilopoda, Lithobiomorpha) de Bulgarie.
Bull. Inst. Zool. (Bulgarie) , XXXVIII : 249-251.

Matic, Z.. 1973b. — Nouvelles contributions a la connaissance des Chilopodes cavernicoles de Bulgarie. Bull. Inst.
Zool. (Bulgarie). XXXVIII : 253-263.

Matic, Z & Golemansky, V., 1964. — Contribution a la connaissance des Lithobiides (Chilopoda, Lithobiomorpha) en
Bulgarie. Ann. Univ. Sofia, LVII : 99-106.

M atic, Z & Golemansky, V., 1965. — Nouvelle contribution a la connaissance des Lithobiides (Chilopoda,
Lithobiomorpha) en Bulgarie. Ann. Univ. Sofia, 58 : 13-25.

Matic, Z & Golemansky, V., 1967a. — Recherches sur les especes et l'6cologie des Lithobiides (Chilopoda,
Lithobiomorpha) en Bulgarie. Bull. Inst. Zool. (Bulgarie), XXIV : 121-132.

Matic. Z & Golemansky. V., 1967b. — Le sous-genre Monotarsobius (Chilopoda. Lithobiomorpha) en Bulgarie. Bull.
Inst. Zool. (Bulgarie), XXIV : 39-50.

Matic, Z & Golemansky, V., 1967c. — Materiaux de la faune des Lithobiides (Chilopoda, Lithobiomorpha) en Bulgarie.
Bull. Inst. Zool. (Bulgarie), XXV : 17-21.

Matic, Z. & Darabantu, C., 1968. — Note critique sur quelques especes du genre Lithobius (Chilopoda. Lithobiidae).
Bull. hist. Zool. (Bulgarie), \\\ I : 103-117.

Negrea, S., 1965. — Contribution a 1 etude des Lithobiidae (Chilopoda) en Bulgarie. Fragm. Balcanica, V : 91-104.
Negrea, S., 1971. — Lithobius ( Monotarsobius ) taschevi Matic et Golemansky est un synonyme de Lithobius
(Monotarsobius) beroni Negrea (Chilopoda, Lithobiomorpha). Trav. Inst. Speol. E. Racovitza.X : 231-233.
Ribarov, G., 1984. — Esl-ce qu'on peut rencontrer en Bulgarie Scolopendra morsitans (L.)?. Bull. Musees Bulg. SE, VII
: 273-276 (in Bulgarian).

Ribarov, G.. 1985. — Species list, ecology, distribution and importance of centipedes Lithobiomorpha (Chilopoda)
from the mountains Strandza and Sakar. Stran. Sakar. Sbor., IV : 279-287 (in Bulgarian).

Ribarov, G., 1986a. — Contribution k la connaissance des Lithobiomorpha (Chilopoda) en Bulgarie Sud-est. Trav.
scien. Univ. Plovdiv. 24 : 155-160 (in Bulgarian).

Ribarov, G., 1987a. — Five new species Lithobiomorpha (Chilopoda) from Southeastern Bulgaria. Acta Zool. Bulg.,
34 : 45-52.

Ribarov, G., 1987b. — Henia angelovi sp. n. (Chilopoda, Geophilidae) - Eine neue Henia- Art aus Sudostbulearien. Acta
Zool., 35 : 86-89.

Ribarov, G., 1989a. — On some Chilopoda species (Myriapoda) published by S. Jurinich and K. Verhoeff. Hist. nat.
Bulg., 1 : 34-35.

Ribarov, G., 1989b. — A contribution to the study of centipedes (Chilopoda) from the Sredna Gora Mountains range.
Trav. scien. univ. Plovdiv, 21 : 165-184. (in Bulgarian).

Ribarov, G., 1990. — Chilopoda new to the science or to the Bulgarian fauna, hi : A. Minelli. Proc. 7th Int. Congr.
Myriapodology. Leiden, Brill : 429.

Ribarov, G., 1992. — The distribution of the centipedes of the genera Harpolithobius and Eupolybothrus in Bulgaria.

Ber. nat.-med. Ver. Innsbruck, suppl. 10 : 361-372.

Verhoeff, K. W., 1926. — Zwei neue Hohlen-Myriapoden aus Bulgarien. Zool. Anz ., 65 : 294-296.

Verhoeff, K. W.. 1927. — Zwei neue Geophilomorphen Gatungen aus Thracien und Mexico. Zool. Anz., 69 : 97-105.
VERHOEFF. K. W., 1928. — Uber Chilopoden aus Bulgarien gesammelt von Hemn Dr. I. Buresch. Bull. Entom. Druz., 4 :
115-124.

Zapparoli, M., 1990. — Distribution patterns and taxonomic problems of the centipede fauna in Anatolian peninsula.
In : A. Minelli, Proc. 7th hit. Congr. Myriapodology. Leiden, Brill : 51-59.

Source : MNHN, Paris

Geographical Distribution of Diplopods in Great
Britain and Ireland; Possible Causal Factors

Anthony D. Barber * & Richard E. Jones **

* Plymouth College of Further Education
Kings Road, Devonport, Plymouth PL1 5QG, U.K.
** 14 Post Office Road, Dersingham
Kings Lynn, Norfolk. PE31 6HP, U.K.

ABSTRACT

The locations and aspects of the habitats of Diplopoda have been recorded under the auspices of the British Myriapod
Survey Scheme since 1970 and records have been obtained from 1790 10 km squares of the British National Grid (Great
Britain and islands) and 419 squares of the Irish National Grid (Ireland), a total now in excess of 30,000 species/location
records. A provisional atlas was published in 1988 and a number of new records have increased our knowledge of the
distribution of millipedes in these islands. The present report examines the pattern of distribution of species and the
possible influence of climatic and other factors on the origins and distribution of the diplopod fauna. It is considered that
a high proportion of the British and Irish fauna is likely to have arrived following the loss of the land connections with
mainland Europe at the end of the last glaciation. Nevertheless there are considerable similarities with the fauna of
nearby countries.

RESUME

Repartition geographique des Diplopodes en Grande-Bretagne et en Irlande : les causes
possi bles.

Les localites et la nature des habitats des diplopodes des lies Britanniques ont ete rtpertoriees sous les auspices du
British Myriapod Survey, etablissant un bi lan depuis 1970 & parti r de 1790 carres dc 10 km de cott, correspondant au
carroyage national britannique UTM (Grande Bretagne et lies) et de 419 carres correspondant au carroyage national
irlandais UTM (Irlande). Le total des stations especes/localites depasse maintenant les 30 000. Un premier document a
ete publie en 1988 sous forme d’un atlas provisoire mais, depuis, un grand nombre de nouvelles donnees sont venues
accroitre notre connaissance sur 1’inventaire et la repartition des especes de diplopodes rtpertoriees dans les lies
Britanniques. Le present travail aborde les modes de repartition des especes, les influences possibles du climat ainsi que
d autres facteurs sur 1 origine et la distribution biogeographique de la faune des diplopodes. On considere qu'une grande
proportion de la faune britannique et irlandaise semble s’etre constitute a la suite de la perte de continuity des terres
emergees avec LEurope continentale a la fin de la derniere glaciation. II existe cependant de profondes similarites avec la
faune des pays voisins du nord de 1* Europe.

Barber. A. D. & Jones, R. E., 1996. — Geographical distribution of diplopods in Great Britain and Ireland;
possible causal factors. In: Geoffroy. J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, not., 169 ; 243-256. Paris ISBN : 2-85653-502-X.

244

ANTHONY D. BARBER & RICHARD E. JONES

INTRODUCTION

A distribution recording scheme for Diplopoda alongside one for Chilopoda was launched
in 1970 by the BRITISH MYRIAPOD GROUP using an itemised record card which has been
described elsewhere (BARBER & FAIRHURST, 1972). A new style card was introduced in 1985.
The present review concentrates on the distributional data that has been derived horn this.

The record card was designed for both professional and amateur usage and a very high
proportion of records were in fact obtained by non-professional but highly competent recorders
working in various parts of the British Isles. Identification was checked by a panel ol referees as
necessary so that the level of misidentification of specimens is likely to be insignificantly low.

By the nature of the scheme, the coverage was patchy.

1 . At least initially, recorders tended to collect the larger and more conspicuous species.

Smaller species and soil dwelling types will be under-recorded.

2. Some areas were recorded in great detail over many years (e.g. Yorkshire) whilst others
had only one or a few casual collections made there.

3. Millipedes are highly sensitive to microclimatic changes and often seasonal in their
occurrence so that a species may not be found on a particular occasion even though it is

common in the area. . ....

4. Different collecting procedures may yield quite different results. For instance, pittall

trapping generally collects" the larger active iuliforms, polydesmids and Chordeuma spp.;
finding Stygioglomeris crinita generally requires careful hand sieving of soil.

5. Immature specimens of some species e.g. Polydesmus spp. and Chordeuma spp.
cannot be determined with accuracy. These may be the only specimens of a species found in a
site.

There is a substantial element of chance in records being made. For instance BLOWER
(1985) wrote, “There remains no evidence that N. minutus (= venustus in the sense of
Schubart, 1934) has ever occurred in Britain, but there is a possibility that it may occur . It has
subsequently (as N. kochii ) been recorded on a number of occasions and is mapped from
twelve 10 km grid squares.

RESULTS OF THE SURVEY

More than 400 individual recorders participated in the scheme and collections made for
other purposes were also examined and a total of more than 30,000 species/site/data records aie
now held. This has allowed the plotting of distribution maps based on the British and Irish
national grids using the 10 km square as the unit of recording. A preliminary atlas using the then
available data was published in 1988 (BRITISH MYRIAPOD GROUP, 1988).

The present discussion is based on updated versions of these maps. Many more records
were made in certain areas compared with others, often with much greater detail. Such well
recorded areas include Kent, Surrey, Isle of Wight. Bedfordshire, parts of S. Wales, Yorkshire,
Lothians, parts of Devon, Norfolk, Suffolk. Figure 1 shows the 10 km squares from which one
or more records exist.

A summary of the regional distribution of species, including occurrence on outlying
islands is shown in Table 1 and examples of distribution patterns are shown in Figures 7-18. An
updated atlas of distribution will be published in due course, meanwhile records are held on
cards and on the database at the Environmental Information Centre, Monks Wood, Huntingdon.

Nomenclature is as in BLOWER (1985) except that a new species, Anthogona britannica , is
since described by GREGORY et al. (1994).

Source : MNHN, Paris

GEOGRAPHICAL DISTRIBUTION OF DIPLOPODS IN GREAT BRITAIN AND IRELAND

245

Table 1. — Distribution in various areas of the British Isles (X = presence in 1 or more 10 km grid square). Based on data
from the Millipede Survey Scheme. She = Shetland Islands, Ork = Orkney Islands, WIs = Western Isles, C&S =
Caithness & Sutherland (North Scotland), Sco = Scotland. Ire = Ireland. IOM = Isle of Man, L&Y = Lancashire &
Yorkshire, Sou = Southern England (South of line from Mersey - Wash, including SE and SW ), Wal = Wales, SWE
= South West England (Devon & Cornwall), KSS = Kent, Surrey, Sussex (extreme SE), CIs = Channel Islands
(Jersey, Guernsey.etc,). NB: Shetland, Orkney, Western Isles, Caithness & Sutherland, Isle of Man and Channel
Islands have relatively few records.

Species

She

Ork

WIs

C&S

Sco

Ire

IOM

L&Y

Sou

Wal

SWE

KSS

CIs

P. lagurus

G. marginata

S. crinita

A. gibbosa

T. lobata

P. germcmicwn

C. rawlinsii

N. polydesmoides

B. melanops

B. bagnalli/bradae

C. silvesire

C. proximum

M. gal lie a

M. sc ut ell are

T. 1 it t oralis

N. varicorne

P. fuscus

C. palmatus

N. kochii

B. guttulatus

A. pallidus

B. tenuis

0. sabulosus

T. niger

A. nitidus

C. londinensis

C. caeruleocinctus

C. vulnerarius

C. latestriatus

C. britannicus

C. punclatus

C. parisiorum

C. tr unco rum

E. armalus

J. scandinavius

0. pilosus

L . belgicus

L. kervillei

M. pratensis

B. pusillus

U. foetidus

A. britannicus

P. angustus

P. testaceus

P. inconstans

P. gallic us

P. denliculatus

B. superus

M. palicola

0. albonanus

S. italica

Source :

246

ANTHONY D. BARBER & RICHARD E. JONES

DISTRIBUTION PATTERNS IN BRITAIN

Geographical

A crude analysis for the commonest species is displayed in Table 2. For the puipose ot
this, Great Britain is divided into a series of regions based on the 100 km grid squares , (Fig. 2)
For each species the total 10 km squares for that species is taken as a percentage of the tota^ 10
km squares in that region for which records exist. This will underestimate smaller and more
difficult to find forms and will also reflect the relative intensity of collecting. Thus, for mst^ice
the northern part of Britain will have disproportionately more larger species recorded because
much collecting there so far has been of a casual and superficial nature. For this reason, the
values should be treated with great caution. Nevertheless they do show up the PI0IJ0unc^
relative scarcity of Ommatoiulus sahulosus and Julus scandinavius in he southeast and an
almost opposite pattern for Cylindroiulus caeruleocinctus which is relatively rarely found in the
southwest. The north-south pattern of some species is also shown.

Table 2 — Analysis of records for the commonest species for regions of the British Isles as a percentage of total 10 km
recorded square's for the region. NSco = Northern Scotland. SSNE = Southern Scotland & Northern England, M dE
= Midland England, East ^Eastern England, SEE = South East England. SWE = South West England, Wal - Wales,
GBT = Total Great Britain, Ire = Ireland.

Species

P. lagurus
G. marginata

N. polydesmoides
C. proximum

N. varicorne
P. fuscus

B. guttulatus

O. sabulosus
T. niger

C. caeruleocinctus
C. punctatus

C. britannicus
J. scandinavius

O. pilosus

P. angustus

P. inconstans
P. gallicus

P. denticulatus
B. superus
M. palicola
O. albonanus

NSco

SSNE

Y/L

MidE

East

0.0

36.8

53.5

56.0

46.8

9.2

27.4

55.1

46.0

51.7

0.0

0.5

0.0

1.3

0.5

3.9

7.3

25.7

24.6

18.9

27.5

36.8

58.3

41.9

57.9

2.4

9.4

21.9

38.2

24.4

32.7

29.9

34.8

23.9

32.3

22.7

91.4

90.4

75.7

80.1

4.8

5.3

4.9

18.4

17.9

57.0

63.9

85.6

83.5

73.6

5.3

13.5

17.6

22.3

7.0

23.7

28.8

49.7

32.0

34.8

24.6

25.7

50.3

47.9

37.8

31.4

38.2

75.9

61.5

64.7

6.8

8.7

12.3

10.7

11.4

0.0

1.4

5.3

27.5

18.9

2.9

3.1

1 1.2

11.3

23.9

8.2

16.3

32.1

31.1

43.3

SEE

SWE

Wal

GBT

Ire

9.6

0.7

67.3

59.7

54.1

46.3

23.9

55.6

38.1

45.2

40.1

32.5

9.3

8.3

42.6

6.0

0.2

30.9

27.6

15.5

18.9

1.7

47.8

54.6

55.8

47.6

18.1

34.6

25.4

27.7

23.0

1 1.3

5.9

28.2

30.4

26.9

4.8

67.3

55.8

68.9

66.1

28.9

33.7

5.0

2.7

11.1

0.2

79.5

76.2

63.5

73.1

45.1

19.0

9.4

20.2

14.6

10.0

6.8

54.1

47.3

33.4

6.9

39.5

35.9

60.8

39.3

31.0

59.0

47.0

58.1

53.8

27.0

15.6

3.3

2.7

9.3

6.7

32.7

13.8

21.6

15.1

24.1

16.6

13.3

29.1

12.7

2.2

54.0

42.0

27.0

30.9

10.0

8.4

0.7

7.1

0.7

Species with a southeasterly distribution include C. caeruleocinctus (above) which may be
increasing its range and Stosatea italica which is beginning to be found in a variety of areas in
England, Wales and Ireland having been found fairly widely in East Kent originally, again
presumably spreading, markedly synanthropic in many cases and found sporadically in rather
superficial habitats. Also Polydesmus testaceus known only from Kent with one older Cornish
record, Metaiulus pratensis, originally found in Kent and Sussex and recently found again in
Kent, presumably mostly soil dwelling, Polyzonium germanicum mostly from Kent ana
Cylindroiulus londinensis often found around the London area, commonly in synanthropic sites
but also recorded elsewhere in England.

Correspondingly, in the southwest Enantiulus armatus, so far found only in one area in
Devon, Chordeuma silvestre from Cornwall, Chordeuma proximum , widespread in southwest

Source : MNHN, Paris

GEOGRAPHICAL DISTRIBUTION OF DIPLOPODS IN GREAT BRITAIN AND IRELAND

247

England, in much of the rest of southern England and very common in South Wales. The two
Leptoiulus spp., L. belgicus and L. kervillei are typically southwestern but records across
southern Britain are known and L. belgicus has been found in Ireland. Several species have not
been found commonly in the southwest, if at all. These include Archiboreoiulus pallidus,
Craspedosoma rawlinsii, Cylindroiulus caeruleocinctus and C. londinensis. Brachychaeteuma
melanops is a distinctly southern species whilst the other two British Brachychaeteuma species
seem to have a more central/northem tendency in general.

There is a distinct group of species apparently rare or possibly absent in northern Scotland.
These include Glomeris marginata, Brachychaeteuma spp., Chordeuma spp., Cylindroiulus
caeruleocinctus, C. londinensis , possibly C. parisiorum, Blaniulus guttulatus, Choneiulus
palmatus and Polyxenus lagurus, although, especially given its distribution elsewhere in Europe,
the latter may simply have been overlooked, something that may also be true of some other
species listed. A number of species appear possibly to be absent from the Shetland Islands, etc.
(Table 1).

Other factors

Apart from the fact that we do not really know the exact regional distribution of species,
there are certain other factors which seem to affect where they are found. A frequently quoted
influence is the presence of calcareous soils; in fact the number of species in Britain which show
a clear calcicole tendency is quite small. Stygioglomeris crinita does appear to favour such soil;
it may in fact be very widespread but the difficulty in finding it makes it impossible at the present
time to be certain. Stosatea italica and possibly Polydesmus testaceus are also possible
calcicoles. The other likely species restricted in this way is Macrosternodesmus palicola.
Cylindroiulus caeruleocinctus, often found on calcareous soils, does not appear to be confined to
them.

There are also a considerable number of species which favour agricultural and/or
synanthropic sites. In the first category are possibly Archiboreoiulus pallidus, Brachydesmus
superus, and maybe Metaiulus pratensis. Of the synanthropes many are also found elsewhere
but they include B. guttulatus, Brachychaeteuma spp., B. superus (?), Choneiulus palmatus
(?), Cylindroiulus britannicus (?), C. londinensis, C. vulnerarius, C. truncorum, Nopoiulus
kochii (?), Ophiodesmus albonanus, Polydesmus angustus (?). Thalassisobates littoralis is a
purely littoral species whilst Cylindroiulus latest riatus is a common coastal species but also
found inland.

The very common Cylindroiulus punctatus is a distinct woodland species and is generally
only found in woods, close to them or on the site of former woodland. Possibly other species
show this tendency in a less pronounced form.

There are some species for which so few records exist that it is difficult to see a clear
pattern. These include Unciger foetidus (one Norfolk site), Anthogona britannica (one Devon
site, GREGORY et. al. , 1994, Trachysphaera lobata (Isle of Wight) and Adenomeris gibbosa
(Dublin). Correspondingly there are species which seem to occur in a wide variety of habitats
over a wide area. Oxidus gracilis and several other species are only known from glasshouses.

COMPARISON WITH OTHER AREAS IN EUROPE
Much of the British diplopod fauna is common with that of nearby areas of Western
Europe. For some species, the British Isles seem to be the centre or one of the centres of their
occurrence, a topic which is discussed by DOOGUE et al. (1993) with special reference to
Ireland. Table 3 shows British species and their occurrence elsewhere on the continental
mainland.

248

ANTHONY D. BARBER & RICHARD E. JONES

Table 3. — Distribution of species in various areas of Europe based on available information (X = presence out of
doors). Based on Doogue et al. (1993), Eason (1970), ENGHOFF (1974 & pers. comm.), Jeekel (1978), Kime
(1990, 1992 and pers. comm.), Lindroth (1957). Meidell (1972, 1979), Meidell & Solhy (1979), Palmen
(1949), Remy & Hoffmann (1959), etc. *U. foetidus and A. britannica are known from single localities only in E
and SW England respectively. Ice = Iceland. Fae = Faeroes, ShO = Shetland & Orkney, GBT = Great Britain, Ire =
Ireland, Nor = Norway. Den = Denmark. NNW = North West Netherlands, NSW = South Netherlands, BeL =
Belgium & Luxembourg, NFr = North France, Arne = Americas; var = various.

Species

Ice

Fae

ShO

GBT

Ire

Nor

Den

NNW

NSW

BeL

NFr

Amc

P. lagurus

G. marginata

S. crinita

A. gibbosa

T. lobata

P. germanicum

C. rawlinsii

N. polydesmoides

B. melanops

B. bagnalli/bradae

C. silvestre

C. proximum

M. gallica

M. scute llare

T. littoralis

N. varicorne

P. fuscus

C. palmatus

N. kochii

B. guttulatus

A. pallidus

B. tenuis

0. sabulosus

T. niger

A. nitidus

C. londinensis

C. caeruleocinctus

C. vulnerarius

C. latest riatus

C. britannicus

C. punctatus

C. parisiorum

C. truncorum

E. armatus

J. scandinavius

0 . pilosus

L. belgicus

L. kervillei

M. pratensis

B. pus ill us

U. foetidus

A. britannica

P. angustus

P. testaceus

P. inconstans

P. gallicus

P. denticulatus

B. superus

M. palicola

0. albonanus

S. italic a

Other species

var

Source : MNHN , Paris

GEOGRAPHICAL DISTRIBUTION OF DIPLOPODS IN GREAT BRITAIN AND IRELAND

249

Almost all species recorded from Norway occur in Britain with the exception of
Polydesmus complanatus (Britain has P. angustus ) and Leptoiulus proximus but a number of
biitisn species have not yet been found there. Eastern Fennoscandia similarly has a high
proportion of -British” species (PALMEN, 1949) but with other more eastern ones. Denmark
with 37 outdoor species, includes 9 not found as yet in Britain. The Netherlands has a somewhat
similar fauna to eastern Britain, several not found in Britain, with southeast Netherlands having
several species only found in southern Britain. Data for Belgium, Luxembourg and northern
h-ance shows most British forms although species such as Adenomeris gibbosa, Trachysphaera
Lobata, tnantiulus armatus and Metaiulus pratensis seem to come from further south. There are
about 16 species from the Belgium, Luxembourg area which do not occur in Britain as far as is
known as present.

Kjme (1990) has mapped a number of species in Europe and has demonstrated some
curious aspects ot this such as the disjunct distribution of Ophyiulus pilosus with a seeming <*ap
in its occurrence between Britain and Scandinavia/North Germany/Poland and Bavaria/Italy°He
demonstrates Nanogona polydesmoides as Britain/France, Chordeuma proximum similarly but
u , Stre (on|y ^ere known from Cornwall) with a much wider occurrence. It would be
helpful to have more data from northern France for comparative purposes.

POSSIBLE CAUSAL FACTORS IN DISTRIBUTION

Present Day Climate

The climate of the British Isles is usually described as Atlantic or Oceanic with
comparatively low temperatures in summer and comparatively high in winter compared with
nearby continental Europe. But within this description is considerable local variation including
the influence of the sea in coastal areas, altitude in so called “Highland Britain” (mostly western
and noithern areas) and the heat island ' effect of urban areas which may permit the survival of
species outside their normal range.

Mean annual rainfall varies from below 500 mm in part of Eastern England to more than
1 600 mm in areas of the west and up to more than 2400 mm in some mountainous areas, with a
general tendency for much of mid-southern, southwestern and northwestern England together
with Wales and Scotland to have 800 mm or more, as does Ireland (Fig. 3). Given the
sensitivity of myriapods to moisture one might seek some correlation with this but there is no
obvious one for most species except perhaps, in England and Wales, Chordeuma proximum.

Temperature is frequently a factor influencing animal distribution, affecting as it may do
survival for individuals, availability of food and breeding cycles. High summer (July) isotherms
tend to lun in an approximately east-west direction (Fig. 5) and there are a number of species
referred to which appear either to have a southerly distribution or are, as in the case for instance
of Glomeris marginata, absent from apparently suitable habitats in northern areas of Scotland.
Until more data is available on the effect of temperature on breeding cycles etc. in diplopods one
can only speculate on the causal factors here.

David (see this volume) describes how G. marginata in an oak forest in southern France
tends to move into the soil in winter, confirming previous work that suggests that it is in fact
cold intolerant. SUSTR (see this volume) describes how G. marginata has a much lower
metabolic rate than G. balcanica and G. hexasticha at low temperatures, which may mean that
in cold conditions it is unable to assimilate effectively and would therefore be unable to tolerate
these conditions for any length of time.

January mean isotherms (Fig. 4) tend to run in a more north-south direction with the
western areas relatively warmed by the influence of the sea whilst the eastern parts have a more

250

ANTHONY D. BARBER & RICHARD E. JONES

“continental” colder drier climate. There do not appear to be clear correlations with species here
but the more extreme, drier climate of the south-east, especially Kent, may favour, either directly
or by excluding competitors, certain species such as Polyzonium germanicum, Stosatea italica,
Polydesmus testaceus, and Metaiulus pratensis.

Past Climates and Land Bridges

Conditions during the Devensian (Wiirm) glaciation were such that much of the British
Isles were covered with ice sheets, reaching at their maximum the whole of the area north of
South Wales and North Yorkshire and the Norfolk coast. At the same time mainland Britain was
joined both to Ireland and to mainland Europe by land bridges. The area south of the ice sheets
was subject to periglacial conditions. Under such circumstances, the diplopod fauna was likely
to have been extremely sparse if not entirely absent in the present day area of the British Isles.
Sub-arctic conditions would have prevailed down to about 10,000 BP following the climatic
improvement of the Aller interstadial. A temperature with a July mean below +10 is quoted for
this Younger Dryas phase (NILSSON, 1982). This period was then followed by the transition to
Pre-Boreal and the re-establishment of forests. VAN DER HAMEN et al. (1971) describe
conditions as having reached mixed deciduous oak forest in the Netherlands by 8,000 BP.

Such conditions would clearly favour the spread of species north from climatic refuges.
However, rising sea levels led to the breaking through at the Dover Straits about 9,600 BP and
between East Anglia and the Netherlands in 8,600 BP (JONES, 1985). After this period the only
dispersal across the English Channel/North Sea would be by passive transport or human
activity. There is thus a period of about 400-1400 years during which species could re-invade
Britain directly. However, the effective separation of Ireland must be placed earlier and this has
been considered to account for the absence in that country of some species of animals (e.g.
certain amphibians and reptiles) present on mainland Britain.

A number of species of diplopod do occur in fairly northern locations in Europe. PALMEN
(1949) reported Proteroiulus fuscus north of latitude 66 degrees with Cylindroiulus latestriatus
and Ommatoiulus sabulosus north of 64 degrees and Polyxenus lagurus, Polydesmus
denticulatus and Polyzonium germanicum all north of 62 degrees in Eastern Fennoscandia.
MEIDELL (1972) included P. lagurus. P. fuscus, Polydesmus complanatus (which does not
occur in Britain), C. latestriatus and two species usually regarded as synanthropes in northern
Europe, Blaniulus guttulatus and Cylindroiulus londinensis as his most northerly recorded
species. EASON (1970) reports three species, Brachydesmus superus, Polydesmus coriaceus
and P. fuscus with possibly B. guttulatus from Iceland. These latter must have all presumably
arrived in some way from other parts of Europe but their existence in Iceland indicates a degree
of tolerance of local conditions there.

GOLOV ATCH (1992) in his survey of the Russian Plain describes P. fuscus from tundra,
P. fuscus, P. germanicum and two other species from northern taiga; these plus P. lagurus, O.
sabulosus and five other species from mid taiga, and amongst species from southern taiga, P.
denticulatus and N. varicome.

Fig. 1. — 10 km grid square distribution: total records for the British Isles.

Fig. 2 . — Regions used for Table 2 analysis.

Fig. 3. — British Isles: Mean Annual Rainfall in mm (based on Atkinson & Smithson in Chandler & Gregory, 1976).
Fig. 4. — British Isles: Mean January Temperatures 1941 - 70 (reduced to sea level) (after Tout in Chandler & Gregory,
1976).

Fig. 5. — British Isles: Mean July Temperatures 1941 - 70 (reduced to sea level) (after Tout in Chandler & Gregory,
1976).

Fig. 6. — British Isles: Fig of the Main Devensian ice advance (solid line) (after Sparks & West, in Evans, 1975).

Source :

GEOGRAPHICAL DISTRIBUTION OF DIPLOPODS IN GREAT BRITAIN AND IRELAND

251

Source : MNHN ' Paris

252

ANTHONY D. BARBER & RICHARD E. JONES

Fig. 7-18. — 10 km distribution Figs of selected species: 7. Polyzonium germanicum, 8. Glomeris marginata, 9.
Chordeuma proximum, 10. Proteroiulus fuscus, 11. Cylindroiulus caeruleocinctus, 12. C. latest riatus, 13. C.
londinensis, 14. C. punctatus, 15. Julus scandinavius, 16. Opliyiulus pilosus, 17. Leptoiulus kervillei ,
18. Enantiulus armatus.

Source : MNHN, Paris

GEOGRAPHICAL DISTRIBUTION OF DIPLOPODS IN GREAT BRITAIN AND IRELAND

253

254

ANTHONY D. BARBER & RICHARD E. JONES

Clearly there is therefore a possibility of some of the above species being able to tolerate
conditions in southern Britain during glacial times or at least to cross the land bridge belore it
broke down. Most of these species are widespread in both Britain and Ireland (although, for
some reason, P. germanicum is confined to the extreme southeast).

Once the land bridges had broken, then entry to the British Isles was possible only by
either passive transport by rafting or by being brought in accidentally by human activity. Rafting
is not easy to demonstrate although PALMEN (1949) refers to P. fuscus on driftwood in S.
Finland and suggest that such passive transport combined with parthenogenesis could account
for its occurrence on outlying islands.

Evidence for possible entry of animals via land bridges or survival during penglacial
conditions is provided by reptiles and amphibians, especially the latter which would be highly
vulnerable to salt water. These animals are of sufficient size that the likelihood of accidental
transport by human activity will be far less than that for soil invertebrates such as diplopods. Six
reptiles and the same number of amphibians are recorded from the British isles together with two
recent successful introductions, Rana esculenta and R. ridibunda. These latter, togethei with the
fact that other species have occurred in the past (HOLMAN, 1993), suggest that it is historical
factors that have determined the relative paucity of British species. Of the British forms, only 1
reptile and 3 amphibians occur in Ireland. One of these, Bufo calamita seems somewhat
anomalous and has been a field for some speculation (see, e.g. BEBEE, 1984). The lower
number of species fits in with the idea of an earlier isolation of Ireland. Of the species that do
occur these all occur in Scandinavia (ARNOLD & BURTON, 1978) with Lacerta vivipara and
Rana temporaria extending to the extreme north and Triturus vulgaris to mid Norway/Sweden.
Bufo sp„ R. temporaria , L. vivipara and the widespread British snake Natrix natrix are known
from Devensian deposits.

Given that the present herpetological fauna was in place by about 8,800 BP (HOLMAN,
1993), by which time separation would have been occurring, we could visualise that a relatively
small number of diplopod species, derived from adjacent areas of Europe, was already present.
These might have included Polyxenus lagurus , Proteroiulus fuscus, Nemasoma varicorne ,
Ommatoiulus sabulosus, Cylindroiulus latestriatus, Polydesmus denticulatus and others.

Other species would arrive either by rafting on tree trunks or other material or be brought
in accidentally with plant material or soil by human activity (see below).

Whatever the mode of their arrival, the improvement in climate towards the so-called
Climatic Optimum of about 7,000 - 5,000 BP, when mean temperatures were about 2-3 degrees
higher than at present, would have covered much of the country. Subsequent climatic changes
such as the climatic oscillation around 5,500 - 5,000 BP and the so called “Little Ice Age
AD 1550 - 1850 may well have brought about later contractions in range. The patchy distribution
one now sees for instance of species widespread in France such as Enantiulus armatus and the
Leptoiulus spp. could be vestiges of a once wider occurrence.

Entry by Rafting and Human Influence

It is difficult to give convincing evidence for rafting by organisms such as millipedes but it
has certainly been suggested for a variety of animal types for oceanic crossing (see for instance,
GARDNER, 1985 referring to geckos in the Seychelles and Mascarenes). Littoral or coastal
species (such as Cylindroiulus latestriatus, a very widespread island species) are most easily
transported in this way and parthenogenesis as in the case of Proteroiulus fuscus would assist.
However, a variety of species from woodland might be transported as a result of exceptional
conditions e.g. storm damage in coastal areas.

Human influence has undoubtedly assisted in the spread of some species of recent arrival
such as Cylindroiulus vulnerarius and human introduction ot invertebrates to islands is widely
quoted (see e.g. JONES & PRATLEY, 1987). Human activity undoubtedly plays a part in the

Source :

GEOGRAPHICAL DISTRIBUTION OF DIPLOPODS IN GREAT BRITAIN AND IRELAND

255

spread of soil animals. A recent example is the New Zealand planarian, Artioposthia triangulata
1992ltain (J' FREW’ pers- comm-) first recorded in Scotland in 1965 but with 416 records by

The occurrence of myriapods in isolated islands such as Iceland as well as the islands
around Britain or those of Denmark (ENGHOFF, 1974) are best explained in terms of transport
either by rafting or human influence. The very high proportion of British species in Ireland,
where land connection would have been lost soon after the disappearance of the ice, cannot be
convincingly explained by reference solely to land bridges but must involve transport across
water.

Lindroth (1957) has listed 17 species of diplopod known from Europe which occur in
the Americas, 16 of which (including Oxidus gracilis, a greenhouse form of tropical origin) are
found in Britain. Of the 18 species of millipede reported from Newfoundland 16 are regarded as
introduced forms lrom Europe with 37 of the 42 isopods and myriapods falling into this
category. Clearly such species have been introduced by human influence and presumably could
have reached the British Isles in the same way. LINDROTH gives five criteria for an introduced
species; it is difficult to apply these to the present situation except for very clearly recent arrivals
or species with clearly synanthropic habits such as Cylindroiulus vulnerarius or C. truncorum. If
glacial and penglacial conditions and subsequent breakdown of land bridges had left a number of
vacant niches then presumably these could be filled by incoming species with the appropriate
characteristics. It is difficult to conceive of common species of woodland and other habitats such
as Glomeris marginata and Tachypodoiulus niger as other than “native” species. In this case
then opportunities for crossing the land bridge in the wake of the ice must have been rather
greater than we have suggested.

Cylindroiulus londinensis, common in much of France, is an example of what may be an
"old introduction which has or is still spreading out from the London area, largely in
synanthropic or semi-synanthropic areas.

CONCLUSIONS

We would suggest that a fairly high proportion of British diplopods are likely to be forms
which may have arrived in the period after the breakdown of the land bridges between Britain
and Ireland and between mainland Europe and Britain. Although climatic and other conditions
are not identical with nearby areas the fauna is similar apart from some more eastern and
southern species, and the occurrence of species both on islands and in America confirms their
ability to cross water. There are likely to have been subsequent changes in distribution due to
climatic changes, the introduction of new species, habitat destruction and possibly to other as yet
imperfectly understood factors, such as those quoted by FORD (1982) for butterflies.

ACKNOWLEDGEMENTS

Clearly this scheme would not have been possible without all those too numerous to mention individually
contributed records, to past scheme organisers C. P. Fairhurst (responsible for the origin of the two myriapod recording
schemes) and D. T. Richardson (who also organised the highly detailed recording of the largest area. Yorkshire), to D.
Doogue in Ireland, J. G. Blower, author of the standard key and of many identifications, A. N. Keay for much help and
advice. R. D. Kime ol Brussels, H. Enghoff of Copenhagen and to P. T. Harding of the British Biological Records
Centre. We would also acknowledge C. M. Moiser for comments and references on reptiles and amphibians.

REFERENCES

Arnold, E. N. & BURTON, J. A., 1978. — A Field Guide to die Reptiles and Amphibians of Britain and Europe. London,
Collins. 242 pp.

Barber. A. D. & Fairhurst, C. P.. 1974. — A habitat and distribution recording scheme for Myriapoda and other
invertebrates. Symp. Zool. Soc. Lond., 32 : 611-619.

256

ANTHONY D. BARBER & RICHARD E. JONES

Bebee, T. J. C., 1984. — Possible Origins of Irish natterjack toads (Bufo calamita). Bril. J. Herpetology . 6 : 398-401.
Blower. J. G., 1985. — Millipedes (Synopses of the Br. Fauna NS. 35). London. E. J. Brill & W. Backhuys, 242 pp.
British Myriapod Group, 1988. — Preliminary Allas of the Millipedes of the British Isles. Huntingdon, NERC. 65 pp.
Chandler. T. J. & Gregory, S., 1976. — The Climate of the British Isles. London/New York, Longmans. 390 pp.
Doogue. D„ F.AIRHURST, C. P., Harding, P. T.& Jones, R. E., 1993. — A Review of Irish Millipedes {Diplopoda). In :

M. S. Costello, Biogeography of Ireland: past, present and future. Occ. Publ. Ir. biogeog. Soc. 2.

Eason, E. H., 1970. — The Chilopoda and Diplopoda of Iceland. Ent. sc and. , 1 : 47-54.

Enghoff, H., 1974. — Om tusindbenenes udbredelse i Danmark (Diplododa). Ent. Meddr.. 42 : 21-32.

Evans, J. G., 1975. — The Environment of Early Man in the British Isles. London. Unwin. 216 pp.

Ford, J. J.. 1982. — The Changing Climate: Responses of the Natural Flora and Fauna. London, Allen & Unwin.
190 pp.

Gardner. A. S., 1985. — Viability of the eggs of the day-gecko Phelsuma sundbergi in sea water. Brit. J. Herpetology ,
6 : 435-436.

Golovatch. S. I.. 1992. — Some patterns in the Distribution and Origin of the Millipede Fauna of the Russian Plain
(Diplopoda). Ber. nat.-med. Verein Innsbruck. 510 : 373-378.

Gregory. S. J., Jones. R. E. & Mauries, J. P.. 1994. — A new species of Millipede (Myriapoda, Diplopoda,
Chordeumatida) from the British Isles. ./. nat. Hist.. 28, 1993 : 47-52.

Holman, J. A., 1993. — British Quaternary herpetofaunas: a history of adaptations to Pleistocene disruptions.
Herpetological J.. 3 : 1-7.

Jeekel. C. A. W., 1978. — Voorlopige atlas van de verspreiding der Nederlandse Miljoenpoten (Diplopoda). Verslagen
technische Gegevens Inst, taxon, zool. Univ. Amsterdam, 15 : 1-69.

Jones. D. K. C.. 1985. — Shaping the Land: The Gcomorphological Background. In : S. R. J. Woodell, The English
Landscape. Past. Present, and Future. Oxford. Oxford University Press.

JONES. R. E. & PRATLEY, P., 1987. — Myriapods of the Isles of Scilly. Bull. Br. Myriapod Group., 4 : 7-15.

KlME, R. D., 1990. — Fauna Europaea Evertebrata: A Provisional Atlas of European Myriapods Part I. Luxembourg,
European Invertebrate Survey. 109 pp.

KlME, R. D.. 1992. — On Abundance of West European Millipedes (Diplopoda). Ber. nat.-med, Verein Innsbruck , 510 :
393-399.

LlNDROTH, C. H.. 1957. — The Faunal Connections between Europe and North America. New York/Stockholm,
Wiley/Almqvist & Wiskell, 326 pp.

Meidell, B. A.. 1972. — En faunistisk undorsokelse avyte Harangerfjordens myriapod fauna og en oversikt over norske
myriapoders taxonomiske og dyregografiske status. Thesis. Bergen, University of Bergen (Maps given in The
distribution of Norwegian myriapods as known to 1972 with a revised list of published papers devoted to the same
subject. Zool. Museum, University of Bergen.

Meidell, B. A., 1979. — Norvegian Myriapods: Some Zoogeographical Remarks. In : M. Camatini, M. Myriapod
Biology. London, Academic Press : 195-202.

Meidell. B. A. & Solhy, T., 1979. — Terrestrial Invertebrates of the Faroe Islands: VI Centipedes and Millipedes
(Chilopoda and Diplopoda). In : A. MlNELLl, Proceedings of the 7th International Congress of Myriapodology.
Leiden. Brill. : 413-427.

Nilsson. T, 1982. — The Pleistocene. Dordrecht/Boston/London. Reidel. 651 pp.

Palmen, E., 1949. — The Diplopoda of Eastern Fennoscandia. Ann. Zool. Soc. "Vanamon\ 13 : 1-54.

Remy, P. & Hoffmann, J., 1959. — Faune des Myriapodes du Grand-Duch6 de Luxembourg. Arch. Sect. Sci. Inst Gr.
Ducal Luxembourg, 26 : 199-236.

Van Der Hamen, T.. Wijmstra. T. A. & Zagwijn, W. H., 1971. — Floral Record of the Late Cenozoic in Europe. In :
K. K. TUREKION, The Late Cenozoic Glacial Ages, Newhaven, Yale University. Press. 606 pp.

Source : MNHN, Paris

Millipedes Recorded in the
Grand Duchy of Luxemburg

Richard Desmond KlME

Institut Royal des Sciences Naturelles de Belgique
Rue Vautier 29, B-1040 Brussels, Belgium

ABSTRACT

Knowledge of the occurence and distribution of millipedes in the Grand Duchy of Luxemburg is reviewed. To date 36
species have been recorded there. Regional variation is taken into account and the differences between the relatively
higher Oesling, a palaeozoic massif of Devonian age, and the lower-lying Gutland of Secondary age are emphasised.
Larger and longer-lived iteroparous species of millipedes are associated with open sites; the reasons for this are
discussed. Comments are made on the phenology of some species.

RESUME

Diplopodes repertories dans le Grand-Duche de Luxembourg.

Les peuplements de diplopodes edaphiques ont ete etudies dans plusieurs sites du Grand-Duche de Luxembourg. Une lisle
de 36 especes repertoriees dans le pays est donnee. La repartition des especes est liee aux diverses regions
luxembourgeoises et surtout h la nature des roches meres. Le resultat des echantillonnages fait apparaitre l'importance
relative des iulides dans les sites ouverts. Les raisons en sont discutees. De nouvelles donnees sur la phenologie de
certaines especes sont commentces.

INTRODUCTION

A list of millipedes from the Grand Duchy of Luxemburg was published by Joseph
HOFFMANN (REMY & Hoffmann, 1959) and consisted of 36 species. None of the material
collected at this time is to be found in the Luxemburg Natural History Museum. In 1982,
occasional collecting began again and from 1988 onwards the Museum has carried out an
intensive programme of pitfall trapping under the direction of Marc MEYER. Several invertebrate
taxa are being studied. This paper is intended to compare recent millipede records with the list of
species found by HOFFMANN (see Table 1) and takes into account regional variations which
remain to be analysed in more detail in the next few years. Some trends are so apparent that they
may be commented upon at this stage.

KlME, R. D., 1996. — Millipedes recorded in the Grand Duchy of Luxemburg. In: GEOFFROY. J.-J.. MAUR1&S, J.-P.
& Nguyen Duy - Jacquemin, M.. (eds), Acta Myriapodologica. Mem Mus . natn. Hist, nat 169 : 257-263. Paris ISBN ;
2-85653-502-X.

258

RICHARD DESMOND KIME

Table I. — Check-list of the 36 millipede species recorded in the Grand Duchy of Luxemburg during the 50’s, compared
with recent collections.

By HOFFMANN
Polyxenus l a gurus (Linne)

Glomeris conspersa C. L. Koch
Glomeris hexaslicha intermedia Lalzel
Glomeris marginata (Villers)

Blaniulus guttulatus (Fabricius)
Proteroiulus fuscus (Am Stein)
Choneiulus palmatus (Nemec)
Nopoiulus kochii (Gcrvais)
Archiboreoiulus pallidus Brade-Birks
Boreoiulus tenuis (Bigler)

Nemasoma varicorne C. L. Koch
Julus scandinavius Latzel
Leptoiulus simplex glacialis (Verhoeff)
Leptoiulus belgicus (Latzel)

Leptoiulus bertkaui (VerhoefD
Allajulus nitidus (VerhoefD
Cylindroiulus caeruleocinctus (Wood)
Cylindroiulus latest riatus (Curtis)
Cylindroiulus truncorum (Silvestri)
Cylindroiulus punctatus (Leach)
Brachyiulus pusillus (Leach)
Ommatoiulus rutilans (C. L. Koch)
Ommatoiulus sabulosus (Linne)
Tachypodoiulus niger (Leach)
Craspedosoma alemannicum Verhoeff
Craspedosoma simile Verhoeff
Melogona gallica (Latzel)

Chordeuma silvestre C. L. Koch
Mycogona germanicum (VerhoefD

Oxidus gracilis (C. L. Koch)

Stosatea italica (Latzel)

Brachydesmus superus Latzel
Polydesmus angustus Latzel
Polydesmus denticulatus C. L. Koch
Polydesmus inconstans Latzel
Polydesmus testaceus C. L. Koch

RECENTLY

Glomeris hexaslicha intermedia
Glomeris marginata

Archiboreoiulus pallidus

Nemasoma varicorne
Julus scandinavius
Leptoiulus simplex glacialis

Allajulus nitidus
Cylindroiulus caeruleocinctus

Cylindroiulus punctatus
Brachyiulus pusillus
Ommatoiulus rutilans
Ommatoiulus sabulosus
Tachypodoiulus niger
Craspedosoma rawlinsi Leach

Melogona gallica
Chordeuma sylvestre
Mycogona germanicum
Orthochordeumella pallida (Rothenbtihler)

Brachydesmus superus
Polydesmus angustus
Polydesmus denticulatus
Polydesmus inconstans
Polydesmus testaceus

COMPARISON OF HOFFMANN'S DATA AND RECENT DATA
In the last ten years 23 of HOFFMANN’S 36 listed species have been found again. With
regard to this it is important to remember that nearly all the recent collecting has been achieved by
pitfall trapping, and this probably accounts for the relatively low number of species found lately,
in particular the blaniulids, some of which hardly ever fall into pitfall traps because of their
hypogeal mode of life. Other species hardly ever or never taken in pitfall traps include Polyxenus
lagurus, recorded by HOFFMANN, and the polydesmid, Macrostemodesmus palicola , as well as
the glomerid, Stygioglomeris crinita , neither of which has been recorded at all from the Grand
Duchy, and both of which are likely to occur in the calcareous areas. A discussion about the
efficiency of pitfall traps in catching different species of millipedes is to be found in
BRANQUART et cil. (1995 and this volume). The other species found by HOFFMANN that have
not been recorded recently are on the whole synanthropic or at the extreme limit of their
geographical range. Leptoiulus bertkaui deserves special mention; HOFFMANN recorded it with
reservations: he did find males on one occasion but could not find the species again. As the site

Source : MNHN, Paris

MILLIPEDES OFTHE GRAND DUCHY OF LUXEMBURG

259

where he located it is in the drainage basin of the Rhine, it would not be particularly unexpected
to find it.

HOFFMANN recorded Craspedosoma rawlinsi as two species, Craspedosoma simile and
C. alemannicum, both described by VERHOEFF. This paper follows SPELDA (1991) in regarding
them as subspecies at best. The gonopods are variable and it seems certain that we are looking at
speciation in progress.

One species new to Luxemburg has been recorded. Seven adults of Orthochordeumella
pallida were found at Weicherdange in the North.

Thus, in the light of present taxonomy, the list of species recorded in the Grand Duchy still
stands at 36, though it will no doubt be added to in the future.

REGIONAL VARIATION IN SPECIES DISTRIBUTION

The northern part of the Grand Duchy is known as the Oesling, a dissected palaeozoic
massif of Devonian age continuous with the Ardenne in Belgium and across the valley of the
River Our which forms the frontier, with the Eifel in Germany. The plateau rises to about 550m
above sea level, and the valleys are quite deep. The parent rocks are chiefly metamorphic schists
and the soils tend to be mull-moder or moder brown earths, much of this area remaining
forested. There are some small patches of limestone in this region.

The larger southern part of the country is called the Gutland and geologically belongs to the
Secondary period. Lower-lying rocks shelve down to the Lorraine Plateau; these are Bunter
sandstones, Muschelkalk, Keuper sandstones and Keuper marls of the Triassic, and Liassic
sands and clays of the Jurassic period. While the Oesling tends to have oligotrophic acidic soils,
the Gutland has rather warmer sands and extensive calcareous soils forming mulls in forests. But
much of this land is cultivated and there are vines in the valley of the Moselle.

In the light of these observations, regional variations in species distributions are to be
expected. Table 2 shows species that were found to be present in some stations in four different
zones. The stations were sampled continuously from the end of the winter until the autumn by
means of barber traps, which were emptied regularly, roughly every three weeks. Species lists
for all the stations are likely to be incomplete at present, since only the one method of collecting
was employed.

Most julids are liable to fall into pitfall traps, however, and relatively very few did so in the
Oesling, which was dominated by polydesmids and glomerids in the traps, but which contains
large populations of chordeumatids, active in the winter when the traps were not operational.
Several species caught in the South were not caught in the Oesling; these include Cylindroiulus
caerideocinctus, C. punctatus, Ommatoiulus sabulosus, O. rut Hans, Melogona gallica and
Polydesmus testaceus. They are less common and some of them may be absent from the North,
especially O. rutilans, which is at its northern limit in Luxembourg, although it penetrates the
Ardenne massif in Belgium along the valley of the Meuse. The Gutland traps caught mainly
julids which attained maximum numbers in the meso-xerothermic calcareous grasslands of the
Keuper Marl.

Table 3 shows the numbers of julids caught in 22 of the sites where Barber traps were set
from March/April until October. The numbers as they stand are not reliable for all quantitative
calculations because the number of operational traps was not always the same on each site, some
of the traps were damaged and specimens either lost or not caught, and the captures are from
different years. They do nevertheless indicate some important qualitative trends and relative
numbers of species on any one site, as pitfall trapping studies use to do (see GEOFFROY &
CELERIER, this volume). The first 15 sites were sampled simultaneously by placing a series of
traps from 4 metres on one side of a hedge to 4 metres on the other side. Thus there were four
traps, one metre apart, at right angles to the hedge in the fields on either side, and there were

260

RICHARD DESMOND KIME

another four traps in the hedge itself. The traps in the hedge caught most of the millipedes. The
last seven sites in Table 3 were more or less open calcareous grassland

The results of the trapping lend further support to the view that the larger and longer-lived
iteroparous species such as the julids shown in Table 3 are abundant in open habitats, see e.g.
DUNGER& STEINMETZGER (1981), KIME (1992). , . ... ,,

The open Keuper sites were dry in the summertime and a lot ol large julids were caught in
them. O'NEILL (1969) subjected seven species of millipede to desiccation and found that they
differed significantly in their resistance to this stress; he attributed this to two factors, size and
cuticular structure. He concluded from observations in the forests of Illinois that tolerant species
are more numerous because of increased ability to disperse to new locations during unfavourable
periods. These tolerant species were the larger : the numerous larger julids in Luxembourg, some
of which are known to be thermophilous in summer, might well exemplify his argument.

Table 2. — Millipedes recorded in Barber traps in different regions of the Grand Duchy of Luxemburg. Cc = Cylindroiulus
caeruleocinctus ; Tn = Tachypodoiulus niger, ; Os = Ommatoiulus sabulosus\ Or = Ommatoiulus rutilans; An -
Allajulus nitidus ; Js = Julus scandinavius ; Cp= Cylindroiulus punctatus ; Bp = Brachyiulus pusillus ; Ls, Lsg -
Leptoiulus simplex glacialis\ Ap = Archiboreoiulus pallidus ; Mg = Melogona gallica\ Op = OrthochordeumeUa
pallida ; Cs = Chordeuma silvestre ; Csp = Craspedosoma sp.; Gm = Glomeris marginala\ Gh - Glomeris
hexasticha intermedia ; Pa = Polydesmus angustus; Pt = Polydesmus testaceus ; Pd = Polydesmus denticulatus ; Pi -
Polydesmus inconstans ; Bs = Brachydesmus superus.

SITE

MILLIPEDES RECORDED

Lsgj

Csp

OESLING:

DEVONIAN

Bauschelterbierg

Beim Weier

Wanterheck

Hartschlaegden

Gresbourg

Weicherdange

Sauerwisen

GUTLAND:

LIAS

Happfeldchen

Hanner Weller

Eiselsbierg

Aucheler

Ehlerange

Schuller

Mondercange

GUTLAND:

MUSCHELKALK

Rampelsbierg

Froumbierg

Tueschaker

Haerebierg

GUTLAND:

KEUPER

Rennpad

Doulen

Groebierg

Sonnebierg

Aarnest

Hunsdorf

Kleibierg

Dennebicrg

Source :

MILLIPEDES OF THE GRAND DUCHY OF LUXEMBURG

261

Table 3. — Numbers of julids caught in pitfall traps. Species as in table 2; TOT = total number of julids per site. For
species, see Table 2.

Year

Soil type

Habitat Site description

Numbers of julids

JOT

DEVONIAN

Surre, Bcim Weier

1989

Loam on schist

Hedge in grassland

Surre, Wanterheck

1989

Loam on schist

Hedge in grassland

Bavigne 1

1989

Loam on schist

Hedge in grassland

Bavignc, Gresler

1989

Loam on schist

Hedge in grassland

LIAS

Clemency 1

1989

Clay

Hedge in grassland

Garnich

1989

Clay

Grass/hedge/arable

Kahler

1989

Clay

Hedge in grassland

Clemency 2

1989

Calcareous clay

Hedge in arable land

181

■

187

Boursdorf

1989

Calcareous loam Grass/hedge/arable

240

255

Machtum

1989

Calcareous loam Hedge in arable land

Oberdonven

1989

Calcareous loam Hedge in grassland

* 4

123

129

Gostingen

1989

Calcareous loam Hedge in arable land

KEUPER

Junglinster,

Rennpad

Junglinster,

1989

Heavy clay

Hedge in grassland

Doulen

1989

Calc, colluvium

Hedge in grassland

168

189

366

Junglinster,

Groebierg

Bech,

1989

Leached loam

Hedge in grassland

110

143

Geyersknapp

Junglinster,

1989

Calcareous clay

Grassland with juniper

102

156

286

Weimericht

1989

Calcareous clay

Grassland with bushes

133

144

155

525

Godbrange,

Schleidelbierg

1989

Calcareous clay

Grassland

194

138

442

Altlinster,

Dennebierg

Reckange,

1990

Calcareous clay

Grassland

315

493

Billknapp

Walferdange,

1990

Calcareous clay

Pasture & set-aside

299

■

344

Sonnebierg

Oberanven,

1990

Calcareous clay

Grassland

278

116

1 1

487

Aarnescht

1991

Calcareous clay

Grassland/few pines

234

130

414

Crawford ( 1 979) noted that millipedes that live in deserts are usually large giving them
more resistance to loss of water. Furthermore, REMMERT (1981) studied body size of terrestrial
arthropods in relation to the abiotic parameters of their milieu, and concluded that for spiders and
winged insects at least, the average body size, and the numbers of animals were governed by the
relative humidity of the biotope.

The results from Luxemburg may reinforce these views strongly by statistically
demonstrating the same phenomenon in Diplopoda. With regard to particular species and the
environmental factors temperature and relative humidity, there is also accordance between these
results in Luxemburg and the detailed works of PERTTUNEN (1953), BARLOW (1957),
HAACKER (1968) and PEDROLI-CHRISTEN (1977, 1993) for Julus scandinavius, Ommatoiulus

262

RICHARD DESMOND KIME

sabulosus, O. rutilans and Tachypodoiulus niger in particular, which feature considerably in this
study. Equally indicative is the absence, or near absence, of other species for which the
parameters have been studied and which require more humid or stable environments to be found
in closed forests. The most abundant millipede in the traps, Cylindroiulus caeruleocinctus, is
described as hygrophile by HAACKER. Yet in Belgium. Luxemburg and Switzerland quantitative
results indicate that it is largely found in open habitats, with peak numbers occurring in the
spring. According to HAACKER, it may burrow in dry weather: a smaller peak is reported by
PEDROLI-CHRISTEN in the autumn. HAACKER and later BLOWER (1985) have also correlated the
presence of C. caeruleocinctus with a high pH value: the Luxemburg figures support this too.
However, C. caeruleocinctus occurs in open woods on acidic sandstone soils in S. E.England,
where the rainfall is relatively low, and where there are very few chordeumatids, and so it is
possible that this reflects its tolerance of dry environments, and that its abundance on calcareous
soils does the same. The same is true of Allajulus nitidus, which is common on calcic mulls, yet
occurs on sandstone. It is considered to be hygrophile too (HAACKER, 1968), and burrows
during the summer as well (GEOFFROY, 1981). The largest Western European julid,
Cylindroiulus londinensis , not found in Luxemburg, is another case in point.

The dominance of julids in periodically dry environments may be due to their resistance to
desiccation and/or their ability to burrow. The humid forests tend to be dominated by rapid¬
breeding semelparous species, reflecting their potential mathematical advantage in favourable
conditions.

PHENOLOGY

Since the traps were set from March or April until October, and caught mainly adults, the
numbers of chordeumatids recorded were disproportionately small. Adult Orthochordeumella
pallida were trapped at Weicherdange between the middle of March and the beginning of May,
1989, the last one obtained was in a map set on April 13. They occurred again in traps set from
September 16 until October 6. and from October 6 until October 27, when operations ceased. At
Clemency, 10 Melogona gallica (9 males, 1 female) were caught before April 25, 6 more males
between then and May 29, and a last male between May 29 and June 19. No more were obtained
before operations ceased on October 2. Results were similar in all the other sites where M.
gallica was found. There was a remarkable preponderance of males. Chordeuma silvestre was
also obtained in the spring on the Sonnebierg; adults of this species were captured by hand in
two forests on October 17, 1982. On the same day Craspedosoma rawlinsi/alemannicum was
similarly taken by hand in three forests. In the pitfalls traps, 26 segment specimens were caught
in August, 28 segment specimens in September, and 28 segment specimens were caught by hand
on October 12, 1991.

Looking at the polydesmids, adult Polydesmus angustus and P. testaceus were found in
every month of trapping. P. testaceus, common in the South, appears to have a spring peak at
least in a number of sites, but more results will be obtained and subject to analysis. P .
denticulatus shows spring and summer activity in the Oesling, and a marked burst of summer
activity in the South, where mainly wandering males were trapped during the warmest period of
the year in dry calcareous grassland sites with the highest summer temperatures in the country.
This was rather unexpected since P. denticulatus has a distinctly northern distribution in Europe
and has generally been associated with sites where the water table is close to or at the surface of
the soil, e.g. the polders (JEEKEL, 1978) and even in submerged sites (ZULKA, 1992). The
species is described as eurytopic by several authors; it is however a fairly small polydesmid.
There may be further support here for O’NEILL’S observation that stressed millipedes wander,
and we may be looking at migratory and/or sexual behaviour as well. In territory where there is a
mosaic of wooded and non-wooded country, it is a little difficult to separate the resident species

Source :

MILLIPEDES OF THE GRAND DUCHY OF LUXEMBURG

263

from the wanderers by using Barber traps. This is why there will be some Berlese-Tullgren
extractions made in this part of Luxemburg in the near future.

On the whole, julids and glomerids were found throughout the trapping period. There was
certainly an early spring peak for the abundant C. caeruleocinctus.

In recent years J. scandinavius has been taken in a large number of pitfall traps in
heathland and grassland; its numbers on the calcareous Keuper Marl are interesting: pitfall traps
in calcareous woodland in Belgium have not caught it, yet it is widespread on neutral and acidic
soils, both in woodland and open sites. PEDROLI-CHRISTEN (1993) reports its absence from
forests on calcareous rocks in Switzerland.

ACKNOWLEDGEMENTS

I give my thanks to Professor Norbert Stomp. Director of the National Museum of Natural History in Luxemburg,
and I owe a special debt of gratitude to Marc Meyer, who provided me with all the millipedes found in the pitfall trapping
schemes intended primarily to monitor beetle populations. Not only that. Dr. Meyer furnished information about the
sites, discussed the ecology of the regions with me and read through the manuscript.

REFERENCES

Barlow, C. A. , 1957. — A factorial analysis of distribution in three species of Diplopoda. Tijds. Ent ., 100 : 349-426.
Blower, J. G., 1985. — Millipedes ( Synopses of the Br. Fauna NS,35). London, E. J. Brill & W. Backhuys, 242 pp.
Branquart, E., Kime, R. D., Dufrene, M. & Wauthy. G., 1995. — Macroarthropod-habitat relationships in oak forests
in South Belgium. I. Environments and communities. Pedobiologia. 39 : 243-263.

Crawford, C. S., 1979. — Desert millipedes: a rationale for their distribution. In : M. Camatini, Myriapod Biology.
London, Acadademic Press : 171-181.

Dunger, W. & Steinmetzger, K., 1981. — Okologische Untersuchungen an Diplopoden einer Rasen-Wald-Catena im
Thuringer Kalkgebiet. Zool. Jh. Syst. 108 : 519-553.

Geoffroy, J. J., 1981. — Modalite de la coexistence de deux diplopodes, Cylindroiulus punctatus (Leach) el
Cylindroiulus nitidus (Verhoeff) dans un ecosysteme forestier du Bassin Parisien. Acta Oecol., Oecol. gener.. 2 :
227-243.

Haacker, U., 1968. — Deskriptive, experimented und vergleichende Untersuchungen zur Autokologie rhein-
mainischer Diplopoden. Oecologia , 1 : 87-129.

Jeekel, C. A. W., 1978. — Voorlopige atlas van de verspreiding der Nederlandse miljoenpoten (Diplopoda)
Amsterdam : 1-68.

Kime, R. D., 1992. — On Abundance of West-European Millipedes (Diplopoda). In: : [E. Meyer, K. Thaler & W.

SCHEDL, Advances in Myriapodology.) Ber. nat.-med. Verein Innsbruck, Suppl.10 : 393-399.

O’NEILL, R. V., 1969. — Comparative Desiccation Tolerance in Seven Species of Millipedes. Am. Midi Nat.. 82 : 1 82-
187.

Pedroli-Christen, A., 1977. — Etude des Diplopodes dans une tourbiere du Haut-Jura. Bull. Soc. Neuchatel Sci. nat.,
104 : 21-34.

PEDROLI-CHRISTEN, A., 1993. — Faunistique des Mille-pattes de Suisse ( Diplopoda ). Neuchatel, Centre Suisse de
Cartographic de la Faune : 1-167.

PERTTUNEN, V., 1953. — Reactions of Diplopods to the relative humidity of the air. Ann. Zool. Soc. " Vanamo ”, 16 : 1-
69.

REMMERT, H„ 1981. — Body Size of Terrestrial Arthropods and Biomass of their Populations in Relation to the Abiotic
Parameters of their Milieu. Oecologia . 50 : 12-13.

Remy, P. & Hoffman, J., 1959. — Faune des Myriapodes du Grand-Duche de Luxembourg. Archives de la Section des
Sciences de I'Institut Grand-Ducal . 26 : 199-236.

Spelda, J., 1991. — Zur Faunistik und Systematik der Tausendfussler (Myriapoda) Sudwestdeutschlands. Jh. Ges.
Naturkde. Wurttemburg , 146 : 211-232.

Zulka, K. P., 1992. — Myriapods from a central European rivers floodplain. In : [E. Meyer, K. Thaler & W. Schedl,
Advances in Myriapodology. ] Ber. nat.-med. Verein Innsbruck, suppl. 10 : 189.

Source : MNHN, Pahs

Some Patterns in the Distribution and Origin of the
Lithobiomorph Centipede Fauna of the Russian Plain
(Chilopoda: Lithobiomorpha)

Nadezhda T. Zalesskaja & Sergei I. GOLOV at CH

Institute for Problems of Ecology and Evolution,

Russian Academy of Sciences, Leninsky prospekt 33, 1 17071 Moscow (V-71), Russia

ABSTRACT ~

Based on the patterns of present-day landscape-zonal distribution, the lithobiomorph fauna of the Russian Plain (25
species or subspecies) appears to be ecologically and historically very strongly associated with a nemoral (=
broadleaved forest) type of vegetation. This allows the reconstruction of the group's regional faunogenesis, with the
fauna shown to be eventually fully migratory, derivative of the adjacent major (Carpathians/Moldova, Caucasus and/or
Crimea) and minor (Urals) nemoral refuges. This corresponds closely to the patterns reported for numerous other animal
groups.

RESUME

Repartition et origine de la faune de chilopodes lithobiomorphes de la plaine russe (Chilopoda :
Lithobiomorpha).

D’aprds les modalites actuelles de la repartition zonale des paysages. la faune de lithobiomorphes de la plaine russe,
compos£e de 25 esp£ces ou sous-especes, apparaTt fortement associee. historiquement et ecologiquement, a une
formation vegetale de type foret de feuillus. Ceci permet de reconstruire la genese regionale des groupes faunistiques,
lesquels peuvent avoir entierement migr£, derivant de refuges forestiers adjacents importants (Carpates/Moldavie.
Caucase et/ou Crimee) ou mineurs (Oural). Ces modalites se rapprochent beaucoup de celles decrites pour de nombreux
autres groupes d’animaux.

INTRODUCTION

The Russian Plain, a vast area covering most of the European part of the former Soviet
Union, has long been known as displaying a classical latitudinal nature zonation combined with
an increasing longitudinal continentality, presenting thereby a highly interesting and important
arena for biogeographical studies. The belts/zones of tundra, taiga, mixed coniferous-deciduous
forests, broadleaved forests, steppe, semidesert, and desert form a full and practically ideal
succession from north to south (e.g. MlLKOV, 1977). The problem of natural distributions in
Lithobiomorpha is still open to discussion (e.g. EASON, 1974,1992), primarily due to the
order's confused taxonomy (especially at the generic level) and our insufficient

Zalesskaja. N. T. & Golovatch, S. I., 1996. — Some patterns in the distribution and origin of the
lithobiomorph centipede fauna of the Russian Plain (Chilopoda; Lithobiomorpha). hr. GEOFFROY, J.-J., Mauries, J.-P.
& NGUYEN Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. nain. Hisi. nat., 169 : 265-268. Paris
ISBN : 2-85653-502-X.

266

NADEZHDA ZALESSKAYA & SERGEI I. GOLOV ATCH

collecting/identification efforts. Although the Russian Plain can boast to be perhaps the best
explorecTregion of the former USSR as regards the lithobiomorph fauna (ZALESSKAJA, 1978),
it still remains an area where lots of centipede records are dubious, and some are even new to the
regional list. Furthermore, general patterns of lithobiomorph chorology, let alone faunogenesis,
on° the Russian Plain have never been adequately discussed. This paper aims at filling in this gap
at least partially, with consideration of all 25 lithobiomorph species or subspecies cui iently
known to populate the region concerned. Much of new faunistic evidence derives from
numerous localities covering the entire nemoral (= broadleaved forest) biome as part of a broader
project on soil macrofauna communities of the Russian Plain (e.g. PENEV, 1992; ESJUNIN et al. ,
1993). It appears as Contribution Nr. 8 to the project entitled “Spatial variation in soil
macrofauna communities of East European oak forests in relation to environmental factors ,
conducted by L. D. PENEV and S. I. GOLOVATCH, sponsored by the USSR Academy of
Sciences, Moscow. This project has enabled us to be considerably more precise in the
distribution patterns and even enrich the list of Russian Plain lithobiomorphs, especially as
regards the fauna of oak, mainly Quercus robur L., forests.

Table 1 — Distribution of lithobiomorph species on the Russian Plain. Symbols: DP - general distribution pattern, C -
zonal-landscape distribution; H - Holarctic; P - Palearctic; E - (pan)-European; CE - central European; SE -
South(east) European; EM - East Mediterranean; Ca - Caucasian; Si - Siberian; End - endemic; Z - natural
vegetation zones; Tu - tundra; T - taiga; F - mixed broadleaved-coniferous forests; FS - forest-steppe; S - steppe;
Ml - mountainous lands south of the Russian Plain (Carpathians, Crimea and/or Caucasus).

Lithobiomorph species

Taxa

Lamyctes fulvicornis Meinert, 1868

T-S, Ml

Lithobius forficatus Linnaeus, 1758

F-Mt

Monotarsobius curtipes (C. Koch, 1847)

Tu-Mt

M. crassipes (L. Koch, 1862)

F-Mt

Lithobius erythrocephalus C. Koch. 1847

F-FS

L. melanops Newport, 1845

F-FS

L. tenebrosus Meinert, 1872 [= L nigrifrons Latzel & Haase, 1880)

T-FS

L. lucifugus L. Koch, 1862

F-Mt

L. piceus L. Koch, 1 862

FS-Mt

Monotarsobius aeruginosus (L. Koch, 1862)

FS-Mt

1 1

L. microps Meinert, 1868 [= M. dubosequi (Brolemann. 1896)]

Lithobius cvrtopus Latzel, 1880

L. borealis Meinert, 1868 [= L. lapidicola A. A. (non Meinert))

L. pelidnus Haase, 1880

h-Mt

L. validus Meinert. 1872

Monotarsobius microps (A. A. non Meinert)

M. sseliwanoffi (Garbowski, 1897)

FS-Mt

Lithobius mutabilis L. Koch, 1862

L. parietum Verhoeff, 1899

L. viriatus Sseliwanoff. 1880

FS-Mt

L. cronebergii Sseliwanoff, 1880

F, Mt

L. proximus Sseliwanoff, 1878

i-S

L. lusitanus tataricus Folkmanova & Dobroruka, 1960

End

Eupolybothrus verrucosus (Sseliwanoff, 1876)

End

Hessebius multicalcaratus Dobroruka, 1958

End

DISTRIBUTION OF LITHOBIOMORPHA ON THE RUSSIAN PLAIN
Table 1 shows the patterns of distribution of the Russian Plain lithobiomorph species,
some examples are also presented in maps (Figs 1-2). As one can see, the bulk of the fauna is
restricted to the zones of mixed broadleaved-coniferous forests and forest-steppe, while both to
the north (taiga and tundra) and south (steppe) the distribution becomes increasingly sporadic

Source :

CENTIPEDE FAUNA OF THE RUSSIAN PLAIN

267

and more closely associated with intrazonal, often anthropogenic, habitats. A similar pattern is
observed from west to east, with gradual impoverishment of the fauna from both Moldova and
the Carpathians toward the Urals. Thus, the Dniester seems to serve as the easternmost limit in
the distributions of Eupolybothrus verrucosus , Lithobius microps, Lithobius cyrtopus ,
L. viriatus, L. piceus , the Bug River “stops” also L. parietum , the Seversky Doniets also
Monotarsobius aeruginosus and L. microps as well as most of M. crassipes , the Don both
Lithobius erythrocephalus and perhaps also L. pelidnus (Figs 1-2). The Volga flow seems to
delimit the distributions of L. lucifugus , L. melcinops, and L. forficatus from the west, and
L. proximus from the east (Figs 1-2). The Urals appear to support the poorest fauna, i.e. the
extremely widespread, Holarctic Monotarsobius curtipes as well as the Siberian Lithobius
proximus.

Fig. 1. — Distribution of some Lithobius species on the Russian Plain and in adjacent mountainous lands: filled
diamond: L. cronebergii - filled square: L. melanops - open circle: L. parietum - open quadrangle: L. pelidnus -
filled triangle: L. piceus - filled circle: L. proximus. T: southern border of the taiga belt; F: southern border of the
nemoral forest belt; FS: southern border of the forest-steppe belt.

Fig. 2. — Distribution of some Lithobiomorpha on the Russian Plain and in adjacent mountainous lands: filled square:
Eupolybothrus verrucosus - open triangle: Hessebius multicalcaratus - open quadrangle: Lithobius cyrtopus - open
circle: L. lucifugus - open square: L. lusitanus tataricus - filled circle: L. tenebrosus - filled quadrangle: L. viriatus
- filled triangle: Monotarsobius sseliwanoffi.

268

NADEZHDA ZALESSKAYA & SERGEI I. GOLOV ATCH

FAUNOGENESIS

The above trends both in the preponderance of Russian Plain Lithobiomorpha to the
nemoral biome and in a west-east faunal impoverishment can be accounted for in terms of both
present-day ecological preferences of the group concerned and historical reasons.

As in the case of millipedes, the lithobiomorph fauna of the Russian Plain appears to be
eventually fully migratory in origin. The role of the adjacent major nemoral refuges is certainly a
leading one in the fauna's conservation and restoration during Pleistocene glaciations and
interglacials (including the Holocene), respectively. It is not by chance that nearly all
Lithobiomorpha populating the Russian Plain occur also in the Caucasus, Crimea and/or
Carpathians with the adjacent Moldova (see Table 1). This is particularly evident when both
Lithobius cronebergii and Monotarsobius sseliwanoffi are taken as examples (Figs 1-2) 1 he
former species had been believed to be confined to the Caucasus Major (ZALESSKAJA, 1978)
until it was discovered in the southern part of the Kaluga Area in 1991. Similaily,
M. sseliwanoffi occurs throughout the Caucasus and Crimea, with only a few records in the
lower Don and middle Volga flows involved. The role of a southern Ural refuge is clearly
subordinate, being probably best expressed only as regards Lithobius proximus. The present-
day distributions, but not necessarily origins, of Russian Plain endemic lithobiomorphs seem to
have been associated with spreading from the Carpathians/Moldova ( Eupolybothrus
verrucosus ), ?Urals (Lithobius lusitanus tataricus. a dubious form whose status requires a
revision), and Caucasus/Crimea (Hessebius multicalcar atus). Anthropochores must have attained
their vast distributions very recently, during the last few decades/centuries. This can be
suaCTested at least for Lithobius forficatus and Monotarsobius curtipes known to very often occur
in purely synanthropic habitats'. The above patterns correspond very closely to the faunogenetic
reconstructions recently conducted for Russian Plain soil/litter-dwelling spiders (Araneae)
(ESJUNIN et at., 1993), millipedes (Diplopoda) (GOLOVATCH, 1992), and earthworms
(Lumbricidae) (VASILEV, 1993). In other words, lithobiomorph centipedes join the numerous
other soil/litter macrofauna (and also some mammal, bird, insect, etc.) groups on the Russian
Plain proved to be both ecologically and historically very strongly associated with a nemoral type
of vegetation.

REFERENCES

Eason, E. H„ 1974. — The type specimens and identity of species described in the genus Lithobius by F. Mcinert. Zool.

J. Linn. Soc., 55 : 1-52. .

Eason, E. H., 1992. — On the taxonomy and geographical distribution of the Lithobiomorpha. Ber. nat.-med . Verein

Innsbruck, suppl. 10 : 1-9.

ESJUNIN, S. L.. GOLOVATCH, S. 1. & Penev L. D., 1993. — The fauna and zoogeography of spiders (Arachmda: Araneae)
inhabiting oak forests of the East European Plain. Ber. nal.-med. Verein Innsbruck. 80 : 179-249.

GOLOVATCH, S. I., 1992. — Some patterns in the distribution and origin of the millipede fauna of the Russian Plain
(Diplopoda). Ber. nat.-med. Verein Innsbruck, suppl. 10 : 373-383.

MlLKOV, F. N., 1977. — Nature Zones of the USSR. Moscow, "Mysl” Publ., 295 pp. (in Russian).

Penev, L. D., 1992. — Qualitative and quantitative spatial variation in soil wire-worm assemblages in relation to
climatic and habitat factors. Oikos. 63: 180-192. .

Vasilev, A. I., 1993. — Some particulars in the distribution and faunogenesis of earthworms in oak forests ot the
Russian Plain (Oligochaeta, Lumbricina. Lumbricidae). Doklady Ross. Akad. nauk, 332 (5) : 657-659 (in Russian).
ZALESSKAJA, N. T., 1978. — Identification Book of Lithobiomorph Centipedes of the USSR. Moscow, "Nauka Publ.,
212 pp. (in Russian).

Source :

The French Millipede Survey: Towards a
Comprehensive Inventory and Cartography of the

Diplopoda in France

Jean- Jacques GEOFF ROY

CNRS, Museum National d'Histoire Naturelle, Laboratoire d’Ecologie Generate
4, avenue du Petit Chateau, F-91 800 Brunoy, France

ABSTRACT

During recent years, field investigations have been carried out in the edaphic compartments of various ecosystem
types (forests, meadows, deep cave and high mountain biotopes, anthropogenic and suburban sites) providing new
zoogeograph ical data that have to be added to recent overviews dealing with the millipede check-fist and distribution in
France. 18 millipede species have been newly recorded from France during the period 1980-1995. This brings the total
number of species to 282 for all millipede taxa (most subspecies are not taken in account). Recent trends in
zoogeographical studies in Europe show that millipede biodiversity studies, monitoring, mapping, and the preservation
of special sites will be of interest for the future. Among the 8 millipede orders present in France, Chordeumatida /
Craspedosomatida is the dominant one, representing >40% of the specific and generic richness. This seem to be a result
of a wide range of origins related to Atlantic, north continental, alpine. Mediterranean and Pyrenean components, some
of them characterized by a high degree of endemism. The first cartographic exercise that has been initiated is a
provisional Allas based on the administrative boundaries of the 95 French “Departements". This atlas is up to date for
the orders Polyxenida, Glomerida, Polyzoniida, Plalydesmida and Callipodida; work is continuing on the orders
Chordeumatida, Polydesmida and Julida. Based on research carried out by EIS, maps of millipede distribution are being
drawn up, following the 10 km x 10 km UTM grid format. This step of the work is still in its early stages, because of the
need to verify much of the previous data. Future initiatives will be developed towards four main aims: (i) Permanent
updating of the check-list of species according to recent knowledge in millipede systematics and nomenclature, (ii)
Permanent updating of the “Provisional Departmental Atlas** of species, (iii) Progressive updating of the UTM
distribution maps, (iv) Proposals for precise recording of the zoogeographical distribution of species according to
cartographic methods developed by the Sendee du Patrimoine Naturel (1EGB. MNHN, Paris). A Fauna GalLICA Diplopoda
file is proposed.

RESUME

La faune des diplopodes de France : etapes vers un inventaire complet et un atlas de repartition
geographique des especes.

Les rZcentes recherches menees dans les compartiments Zdaphiques de divers ecosystemes portent a 282 le nombre
total d'especes de diplopodes rZpertoriees cn France. Elies se repartissent au sein de 98 genres. 19 d'entre elles ont ete
decrites ou inventorizes entre 1980 et 1995, ce qui traduit une augmentation de 6,7% de la richesse specifique connue au
cours de cette periode. Les especes de France appartiennent aux huit ordres europeens (sur les 16 ordres et sous-ordres qui
composent la classe Diplopoda) : Polyxenida. Glomerida, Polyzoniida, Platydesmida. Callipodida, Craspedosomatida,

GEOFFROY, J.-J., 1996. — The french millipede survey: towards a comprehensive inventory and cartography of
the Diplopoda in France. In: Geoffroy. J.-J., MAURlfeS, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 269-280. Paris ISBN : 2-85653-502-X.

270

JEAN-J ACQUE S GEOFFROY

Polydcsmida & Julida. La mise en forme des donnees les plus recentes confirme la large dominance de 1 ordre
Chordeumatida qui represente 40% de la richesse tant sp^cifique que gdneriquc. Cela s'explique par 1 origine des espies
representees par des composantes biogeographiques varies et caracterisees parfois par un ires fort cnd^misme. Base sur
le critere de presence-absence 5 I'interieur des limites administralives departementales, la realisation d un atlas est
destinee £ produire un document a partir duquel une cartographic en rapport avec les limites biogeographiques naturelles
sera facility. I! est a jour pour les ordrcs Polyxenida, Glomerida, Polyzoniida, Platydesmida. & Calhpod.da et en cours
d’amenagement pour les ordres Chordeumatida. Polydesmida & Julida. Une cartographic basee sur la grille des carres UTM
10 km x 10 km. faite en coordination avec les travaux europeens de 1’EIS parait, & terme, comme une contribution a
renrichissement de Fauna Europaka Evertebrata (banque de donnees faunistique europeenne). La precision apportee
par les carres de 10 km x 10 km parait largement suffisante pour une etude k 1'echelle nationale. Les bases de donnees
mises en oeuvre doivent etre precises mais demeurer claires et pratiques. 11 convienl done de ne considerer qu un nombre
restreint de types d’esp£ces ou d’£cosyst£mes de reference. On considerc dune part les espfcces a large repartition, d autre
part les especes liees soit aux influences mediterraneennes, atlantiqucs ou septentrionales, soit a des milieux particuliers
• cavernicoles, nivicoles ou halophiles. Conformement & une proposition du Conseil de l’Europe, la France est d^coupce
en 5 regions biogeographiques : atlantique. continentale. mediterraneenne, alpine & pyreneenne. En depit de son
caractere arbitraire. ce decoupage peut contribuer a letude de la biodiversite de fairness naturelles. De plus, nombre
d’especes chevauchant deux ou plusieurs zones, celles-ci pourront etre comparces a 1 aide d indices de similitude. Dans un
souci de simplification, l’appartenance des especes a divers environnements est limitee 5 types d ecosyslemes
ecosyst£mes de plaines. cotiers, d'altitude (hauts-plateaux & montagnes), insulaires, souterrains^ Fauna Gaix/CA
Diplopoda est une base de donnees destinee a gerer la repartition et la biodiversite des diplopodes de France. Elle est
elaboree dans 4£ Dimension sur systeme Macintosh et met en relation 14 fichiers dc donnees consumes autour du
catalogue des especes. Les fichiers principaux concernent les localites et les collectes. Le code-localite designe la
commune el un ensemble dc parametres prScisant sa situation et les caracteristiques dc I'environnement. Les coordonnees
geographiques autorisent un transfer! des donnees vers des systemes de cartographic automatique traites par divers
organismes. Un outil de cettc nature facilite, a terme, la mise ^ jour de I’invcntaire des especes en lonction de I avancee
des recherches en systemalique et en nomenclature, de meme que la mise en forme de l'atlas departemental . L essentiel des
informations provient a ce jour de donnees issues de collections de reference (MNHN, Pans) ou de bases bibliographiques
(Centre International de Myriapodologie, BiblioMac-Milpal). II convienl alors de mettre ces informations en relation
avec les capacities de traitement d'organismes tels que le Secretariat Faune-FIore du MNHN et de faciliter la constitution
d'un reseau d'observateurs-collecteurs capables. en liaison permanente avec des chercheurs specialistes, de multiplier les
points d’observation sur le territoire considere.

INTRODUCTION

The growing interest in biodiversity has prompted many authors to consider the question
of the number of species there are likely to be on Earth (see LEVEQUE, 1994; SOLBRIG, van
EMDEN & van OORDT, 1992; WILSON, 1993). Some papers describe steps towards an estimate
of the number of species either from a global assumption (MAY, 1992) or in connection with
particular taxa, area, environment or even locality (JUBERTHIE & DECU, 1994; SERRA-COBO et

al ., 1993). . r

Dealing with terrestrial invertebrates, there is obviously no group ol organisms tor which
the total number of species is known directly and certainly no group of arthropods tor which
even a majority of species are known or described. The inventory of living forms is still very tar
from complete, even in taxa usually considered as well-known. To this reality of which we are
reminded by BARB AULT (1994) has to be added the fact that such an inventory is still not
achieved in regions or countries usually considered as well-worked. Most arguments about
global biodiversity depend on a series of assumptions about the connection between local species
diversity and regional (or “national”) diversity (see CULVER & HOLSINGER, 1992; ERWIN,

1988). ... J .

Looking at this situation face to face, and so far as the specific richness and scarcity ol
species are good criteria for evaluating ecological systems (BLANDIN, 1989), we must
emphasize the importance of precise and comprehensive species inventories and taxonomical
diversity studies, wherever it is possible -notably in West-European areas- as preliminary
geographical tools for further analysis of biodiversity (MAURER, 1994). We describe below the
evolution of a FRENCH MILLIPEDE SURVEY, from the initial check-list of species to a computer
data-base to serve as a tool for future biodiversity studies on Diplopoda in France.

THE FRENCH MILLIPEDE SURVEY

271

RESULTS

Check-list of the millipede species

During last few years, field investigations have been carried out in the edaphic
compartments of various ecosystem types (forests, meadows, deep cave and high mountain
biotopes, anthropogenic and suburban sites, etc.), providing new zoogeographical data (see
GEOFFROY, 1981) to be added to recent reviews dealing with millipede check-lists and
distribution in France. Thus, 20 millipede species have been newly recorded on French territory
during the 1980-1995 period (Table 1). This allow us to conclude that the total number of
species is at present 282 for all millipede taxa (most of subspecies are not taken in account).
Some of these taxa have to be added to the previous check-list proposed by GEOFFROY (1990a).

Table 1. — Millepede species discovered or described in France during the 1980-1995 period. Data from Geoffroy,
1990b and Geoffroy & MauriEs, 1992. Original data from Geoffroy. Maurj£s, Klme & Pedroli-Christen.

Species

Distribution (“departement”)

Environment

Trachysphaera drescoi (Conde & Demange. 1961)

Brachychaeteuma bagnalli Verhceff, 1911
Crossosoma mauriesi Strasser, 1970
Crossosoma cavernicola (Manfredi. 1951)

Hispaniosoma racovitzai Ribaut, 1913
Opisthocheiron canayerensis Mauries & Geoffroy, 1982
Origmatogona kimeorum Mauries, 1990
Rhymogona cervina (Verhceff, 1910)

Rhymogona montivaga (Verhceff, 1894)

Vascosoma coiffaiii Mauries, 1966
Vascosoma coiffaiti falsaforma Mauries, 1990
Vascosoma duprei Mauries, 1990
Orthochordeumella leclerci Mauries, 1986
Galliocookia leclerci Mauries, 1983
Galliocookia balazuci Mauries, 1983
Mastigonodesmus lopezi Mauries, 1980
Mastigonodesmus fagniezi Mauries, 1982
Occitanocookia hirsuta Mauries, 1980
Archiboreoiulus pallidus (Brade-Birks, 1920)
Dolichoiulus tongiorgii Strasser, 1973 _

Pyrenees- Atlantiques

Caves

Rhone

Caves

Alpes-Maritimes

Soil & high mountain

AIpes-Maritimes

Caves

Ariege

Soil & high mountain

Gard

Caves

Dordogne

Soil

Doubs (new for France!)

Soil

Explanation in the text

Soil & Caves

Pyr6n6es- Atlantiques

Caves

Pyrenees- Atlantiques

Caves

Pyrenees- Atlantiques

Caves

Ardeche

Caves

Gard

Caves

Gard

Caves

Herault

Caves

Gard

Caves

Herault

Caves

Cher, Meurthe-et-Moselle, Ain

Caves

Alpes-Maritimes. Gard

Halophilous

Rhymogona cervina (Verhceff, 1910), after revision of the collection in MNHN, Paris, is
new for France (unpublished data, MAURIES & PEDROLI-CHRISTEN, pers. comm.). It is known
from Les Gras (Doubs), a locality closely related to the contact zone with Rhymogona montivaga
and the Swiss Jura.

Rhymogona montivaga (Verhceff, 1894) is new for the check-list of French millipedes,
because of its synonymy with Rhymogona silvatica (Rothenbiihler, 1899) and R. s. hessei
(Ravoux, 1935). It is distributed in Cote d'Or, Isere, Haute-Marne, Rhone, Haute-Saone,
Haute-Savoie.

A third nominal species, Rhymogona alemannica (Verhceff, 1910) is known from
Meurthe-et-Moselle and Haut-Rhin.

Richness and biodiversity of m illipede taxa

Recent zoogeographical trends in Europe show that invertebrate biodiversity studies,
monitoring and mapping, and preservation of special sites are more and more of great interest for
the future (GONSETH, 1993; MAURIN & GUILBOT, 1993; RASMONT, 1993). In connection with
the European Millipede Survey (E.I.S.), Kime (1990) published a first provisional atlas of

272

JEAN-JACQUES GEOFFROY

European myriapods, closely linked to the work done by the BRITISH MYRIAPOD GROUP
(1988; see Barber & Jones, this volume).

Among the 8 millipede orders present in France (GEOFFROY, 1992, 1993a),
Chordeumatida / Craspedosomatida is the most dominant one. It represents >40% of the specific
and generic richness (113 species). This appears to be a result of the wide range of origins
related to Atlantic, north continental, alpine, Mediterranean and Pyrenean components, some of
them characterised by a very high degree of endemism (Table 2). This result is quite different in
the British fauna, in which Chordeumatida (9 species) represent only 15.5% of the whole
Diplopoda (after BLOWER, 1985). On the contrary, in Switzerland, the importance of the order
Chordeumatida is relatively high, 33.9% for 43 species (after PEDROLI-CHRISTEN, 1993). In
Italy, where we can see a high specific richness of millipedes, it represents 3 1.5% (148 species)
of the Diplopoda (after STRASSER & MlNELLI, 1984).

Table 2. — Taxonomic diversity of the Diplopoda in France. See for comparison Geoffroy, 1989, 1990a, 1990b,

1993b; Geoffroy & Mauri£s, 1992.

Taxa (Orders & Sub-orders)

Species Richness

Polyxenida

1.8

Glomerida

10.6

Polyzoniida

1.1

Platydesmida

0.4

Callipodida

1.1

Chordeumatida/Craspedosomatida

113

40.1

Craspedosomatidea

33.0

Chordeumatidea

7.1

Polydesmida

12.7

Julida

32.2

Blaniulidea

10.6

Julidea

21.6

Class Diplopoda

282

100.0

The comparison of the species richness of the Diplopoda in France with other European
countries shows the possible mediterranean influence on these results related to some unnatural
administrative limits (Table 3). It would be of great interest to complete this comparative review
with more precise data from Scandinavia, Germany, Austria, Central Europe and Spain.

Table 3. — Compared specific richness of millipede fauna in different Western European countries.

Country

N species

Reference

Luxemburg (Gd Duchy)

Remy & Hoffmann, 1959; Kime, this volume

Denmark

Enghoff, 1974

Netherlands

Jeekel, 1978

U. K.

Blower, 1985

Switzerland

127

Pedroli-Christen. 1993

Germany

160

SCHUBART, 1934

France

282

This work

Italy

470

Strasser & MlNELLI, 1984

Source . MNHN, Paris

THE FRENCH MILLIPEDE SURVEY

273

Provisional departmental atlas

The first cartographic project that has started is a provisional Atlas using the administrative
limits of the 95 French “departements”, as initially used for the counties and vice-counties of the
United Kingdom, (BLOWER, 1985). Based on a “presence-absence” criterion, this atlas will
provide a basic document on biogeographic distribution. It is up to date for the orders
Polyxenida, Glomerida, Polyzoniida, Platydesmida and Callipodida (unpublished data); work on
the orders Chordeumatida/Craspedosomatida, Polydesmida and Julida is in progress
(GEOFFROY, 1989, 1993b).

UTM grids maps of French millipedes

In accordance with research by EIS members (KlME, 1990), distribution maps of
millipede species are in preparation, using the 10 km x 10 km UTM grid square. This step of the
work is still at an early stage, because of the necessity to examine old records from the literature
and museum collections and to correct their identification and bring nomenclature and
classification up-to-date. This will be a contribution to the EIS Fauna EUROPAEA
EVERTEBRATA data base (GEOFFROY, 1994a, 1994b).

Fauna Gallica Diplopoda: a data base

Fauna Gallica Diplopoda is a data base whose aim is the management and monitoring
of the distribution and biodiversity of millipedes in France. It is developped on “4th dimension”
data base for Macintosh computers which allows 14 data files to be organized around the
checklist of species. The main data files deal with localities and samples (Fig. 1). The main
structure of the data-base is adapted from a previous one used by the GlLlF Group for
Lepidoptera (Mothiron, 1993).

The locality file points out the commune, city-code, co-ordinates and several parameters
dealing with precise situation and environment. The geographic co-ordinates (grades and UTM
grid) allow data-transfer to automatic cartographic systems used in centralized institutes. It is the
first file to be completed. Then, linked to the previous one, the collect-file gives information
related on dates, collectors and lists of sampled species.

Such a data-base must be as precise and as complete as possible but, in order to be
understandable to new workers (colleagues, other specialists, ...), it must be quite simple and
clear. In order to avoid useless complexity, it seems advisable to select only a restricted number
of species categories and ecosystem types.

— Species categories

On one hand, we have to consider species with a wide-range distribution; on the other
hand, species closely related to either Mediterranean, Atlantic and northern influences, or special
environments such as the halophile category, coastal (MAURIES, 1982), high mountain
(GEOFFROY, 1981) and caves or any other deep subterranean biotope.

In regard to this, a provisional checklist of highly troglophitic and troglobitic French
millipedes has been proposed (GEOFFROY, 1994a). It is composed of 34 species : 5 Glomerida,
14 Chordeumatida, 6 Polydesmida and 9 Julida. Among these species, 7 taxa can be considered
as interesting ones for natural patrimony. They are of special interest either for biogeography or
evolutionary and paleontological history of lineages and territories. These species and their
habitat should be selected for conservation projects in order to maintain the biodiversity level of
these groups on a national and European scale. These taxa occur in high limestones sites in the
Alps (genus Broelemanneuma), the Pyrenees (genus Vascoblaniulus) and in the Cevennes
(“Causse de Canayere et de Bramabiau”). It should be possible, in the future, to be able to
distinguish other such interesting taxa among the French fauna.

274

JEAN-JACQUES GEOFFROY

At present, this selection contains the 7 following troglobitic species :

Broelemanneuma furcation Ribaut 1913 ai^pinf

Broelemanneuma gayi Demange 1968 ai p vp

Broelemanneuma gineti Ribaut 1954 i pimf

Broelemanneuma palmatum (Brblemann 1902) ^ ^

Broelemanneuma pectiniger (Brblemann 1 902) . KTr^ . K ,

Opisthocheiron canayerensis Maurtes & Geoffroy 1982 MED1 rERRANEAN

Vascoblaniulus cabidochei Mauries 1967 PYRENEAN

D6partements

Observateurs

NoDepartmt A

NomD^partmt A

CodeObs A

Ncm A

Pr6nom A

CodeStade

LlbStade

Especes

NoEspece

Nombre

Stade

Altitude

Biotope

Descripteurs

Code desc

Nom descnpteur

Coliectes

CodeLoc

Annee

Jour

JourOuNult

Observateur

Esp6ces

•

RefBiblo

Catalogue

Identi tiant

Genre

Espece

Descnpteur

SynGenre

Synespece

Sphere

ORDRE

Annee

Code Ordre

ListelDF

CodeEspece

Statut

DernAnnee

PeriodeAct

Commentaire

NumOrdre

Nb citations

CodeDouteux

TopBAN

TopCES

TopCOJ

TopETA

TopFON

TopMAN

TopRAM

TopRSM

CodeUibam

CodeMontig

Biotope

CodeBlotope

LibBiotope

Observations

CodelNSEE

NoOrdre

TopCarte

DateObs

Date2

Abscisse

Pr6cAbscisse

Ordonn^e

Pr^cOrdonnee

CodeEspece

Stade

Observateur

RefBiblo

Locaiit6s

CodeLoc

Libcommune

Lieudit

Altmin

Altmax

Altmoy

Grande Ville

DistGV

DifGV

Code commune

Abscisse

Ordonn6e

Biotope 1

Biotope2

Biotope3

Biocommentaire

Acces

Arbres

Arbustes

Herbac4es

Sol

Filler

UTM 10x10

Biblio

CodeRef A

Libels T

Commentaire T

Localit£slDF

CodeLoc

Secteur

NbDonnees

Av1970

Ap1970

Glomerida

Polydesmida

Julida

Code Regroup

LibelRegroup

Fig. 1. — Fauna Gallica Diplopoda : main structure of the database.

— Biogeographical and ecogeographical spheres

According to a proposal from the European Council, France could be divided into 5
biogeographic areas: 1: Atlantic, 2: Continental, 3: Mediterranean, 4: Alpine & 5: Pyrenean.
Despite the arbitrary nature of such a “biogeographic” division, it can facilitate the study of

Source : MNHN ' Paris

Ti IE FRENCH MILLIPEDE SURVEY

275

addition, the relationships between species and environment are taken into account in 5 chosen
large ecosystem types: 1: plains (s. 1.), 2: coasts, 3: high mountains and high table-lands, 4:
islands (Mediterranean, Atlantic and Channel Is.) & 5: caves (natural and artificial cave
environments). As often shown and discussed before, subterranean ecosystems are one of the
most important source of possible refuge for biodiversity (SERRA-COBO et al., 1993), in natural
caves or MSS compartments (GEOFFROY, 1984a, b; MAURIES & GEOFFROY, 1982) and
artificial quarries as well (GEOFFROY, 1991).

CONCLUSION AND FUTURE PROSPECTS

The present work shows the preliminary steps towards a long-term comprehensive
approach to the distribution and diversity of the millipede fauna of France. A large part of the
data originate from reference collections (MNHN, Paris), bibliographic data-bases (CIM-Paris,
BiblioMac-Milpat) and unpublished recent field collections. As the number of specialists is low
for taxa such as Diplopoda and other myriapods groups, it seems important now to consider the
possibility of establishing a more or less formal network of collectors in France. This would
augment the data-bank, that could be centralized and studied by researchers in the MNHN
laboratories at Paris and Brunoy.

Future developments can be summarised under four headings:

(i) Permanent updating of the check-list of -millipede species, according to recent
knowledge of millipede systematics and nomenclature.

(ii) Permanent updating of the “Provisional Departmental Atlas” of millipede species.

(iii) Progressive updating of the UTM grid cartography (10 km x 10 km).

(iv) Proposals for a precise zoogeographical distribution of species according to
cartographic methods developed by the Service du Patrimoine Naturel (SPN: MNHN, Paris).

The French Millipede Survey and Fauna Gallica Diplopoda are tools lor this
purpose.

annexe

Check-list of the French millipede species, from the data base Fauna Gallica Diplopoda (February 1996). In this list,
Geoglomeris subterranea Verhoeff. 1908. is considered as a senior synonym of Siygioglomeris cnniia
Brolemann, 1913 and Geoglomeris jurassica Verhoeff. 1918.

Lophoproctinus inferus

Silvestri

Lophoproctus jeanneli

(Brolemann)

Lophoproctus lucid us

Chalande

Polyxenus lagurus

(Linne)

Polyxen us ma cedon icus

Verhoeff

Adenonieris gibbosa

Mauries

Adenomeris hispida

Ribaut

Corsikonieris remyi

Verhoeff

Doderoa genuensis

Silvestri

Geoglomeris duboscqui

(Brolemann)

1 1

Geoglomeris granulosa

(Ribaut)

Geoglomeris provincial is

(Brolemann)

Geoglomeris subterranea

Verhoeff

Glomeridella kenillei

(Latzel)

Glomeris annulata

Brandt

Glomeris connexa

C.L.Koch

Glomeris conspersa

C.L.Koch

Glomeris guttata

Risso

Glomeris helve tica

Verhoeff

Glomeris humbertiana

Saussure

Glomeris intermedia

Latzel

Glomeris marginata

(Villers)

Glomeris pustulata

Latreille

Glomeris transalpina

C.L.Koch

Glomeris undulata

C.L Koch

Loboglomeris pyrenaicu

(Latzel)

Loboglomeris rugifera

Verhoeff

Onychoglonteris caslanea

(Risso)

Prologlomeris vasconica

(Brolemann)

1903

POLYXENIDA

Polyxenidea

1910

POLYXENIDA

Polyxenidea

1888

POLYXENIDA

Polyxenidea

1758

POLYXENIDA

Polyxenidea

1952

POLYXENIDA

Polyxenidea

1960

GLOMERIDA

Glomeridea

1909

GLOMERIDA

Glomendea

1943

GLOMERIDA

Glomeridea

1904

GLOMERIDA

Glomeridea

1913

GLOMERIDA

Glomeridea

1947

GLOMERIDA

Glomeridea

1913

GLOMERIDA

Glomeridea

1908

GLOMERIDA

Glomeridea

1894

GLOMERIDA

Glomeridea

1833

GLOMERIDA

Glomendea

1847

GLOMERIDA

Glomeridea

1847

GLOMERIDA

Glomeridea

1826

GLOMERIDA

Glomeridea

1894

GLOMERIDA

Glomeridea

1893

GLOMERIDA

Glomeridea

1884

GLOMERIDA

Glomeridea

1789

GLOMERIDA

Glomeridea

1804

GLOMERIDA

Glomeridea

1836

GLOMERIDA

Glomeridea

1844

GLOMERIDA

Glomeridea

1886

GLOMERIDA

Glomeridea

1906

GLOMERIDA

Glomeridea

1826

GLOMERIDA

Glomeridea

1897

GLOMERIDA

Glomeridea

JEAN-J ACQUES GEOFFROY

30 Spelaeoglomeris alpina

31 Spelaeoglomeris doderoi

32 Spelaeoglomeris jeanneli
3 3 Trachysphaera drescoi

34 Trachysphaera lohaia

35 Trachysphaera pyrenaica

36 Hirudisoma latum

37 Hirudisoma pyrenaeum

38 Polyzonium germanicum

39 Fiona tuherculata

40 Callipus corsicus

41 Callipus foetidissimus

42 Callipus sorrentinus

43 Anamastigona pulchella

44 Anthogona variegata

45 Brachychaereuma hagnalli

46 Brachychaeteuma bradae

47 Brachychaeteuma cadurcensis

48 Brachychaeteuma furcatum

49 Brachychaeteuma melanops

50 Brachychaeteuma peniculatum

51 Brachychaeteuma plumosum

52 Brachychaeteuma provinciate

53 Broelemanneuma furcatum

54 Broelemanneuma gayi

55 Broelemanneuma gineti

56 Broelemanneuma palmatum

57 Broelemanneuma pecliniger

58 Camptogona delamarei

59 Camptogona duboscqui

60 Ceratosphys amoena

61 Ceratosphys banyulsensis

62 Ceratosphys guttata

63 Ceratosphys nivium

64 Ceratosphys picta

65 Ceratosphys simoni

66 Ceratosphys vandeli

67 Chamaesoma broelemanni

68 Corsicosoma legeri

69 Cranogona dalensi

70 Cranogona delicata

7 1 Cranogona denticulata

72 Cranogona orientate

73 Cranogona pavida

7 4 Cranogona touyaensis
1 5 Cranogona uncinata

76 Cranogona vasconica

77 Craspedosoma alemannicum

7 8 Craspedosoma raw l ins ii

79 Craspedosoma taurinorum conforme

80 Crossosoma broelemanni

8 1 Crossosoma cavernicola
82 Crossosoma mauriesi

8 3 Crossosoma peyerimhoffi
84 Cyrnosoma beroni
8 5 Cyrnosoma coineaui

8 6 Cyrnosoma strasseri

87 Escualdosoma gourbaultae

88 Haasea flavescens

89 Helvetiosoma arvemum

90 Hispaniosoma racovitzai

9 1 Hypnosoma exornatum

92 Hypnosoma juberthieorum

93 Hypnosoma pallidum

94 lulogona tirolensis cisalpinum

95 Janetschekella valesiaca

96 Marquetiella auriculata

97 Marquetiella lunata

98 Marquetiella pyrenaica

99 Nanogona balazuci

1 00 Nanogona cebennica

1 0 1 Nanogona davidi

1 02 Nanogona digitata

1 03 Nanogona polydesmoides

1 04 Nanogona uncinata

105 Ochogona gallitarum

1 06 Opisthocheiron canayerensis

1 07 Opisthocheiron cornutum

1 08 Opisthocheiron elegans

1 09 Opisthocheiron fallax

1 1 0 Opisthocheiron lacazei
1 1 1 Opisthocheiron penicillatum
1 1 2 Origmatogona kimeorum
1 1 3 Pyreneosoma barbieri
1 I 4 Pyreneosoma bessoni
1 1 5 Pyreneosoma digitatum
1 1 6 Pyreneosoma ribauti

Brolemann

Silvestri

Brdlemann

<Cond£ & Demange)

(Ribaut)

Brandt

Silvestri

Verhoeff

(Savi)

Verhoeff

Silvestri

Ribaut

Verhoeff

Brolemann & Brade-Birks

Mauri&s

Ribaut

Brade-Birks & Brade-Birks

Ribaut

Demange

Ribaut

(Brolemann)

Mauries

(Brolemann)

Ribaut

Brolemann

Ribaut

Mauries

Ribaut & Verhoeff
(Brolemann)

Mauries

Del mas

Ribaut

Mauries

Ribaut

Verhoeff

Leach

Silvestri

Strasser

Manfredi

Strasser

(Brolemann)

Mauries

(Latzel)

(Ribaut & Brolemann)

Ribaut

Mauries

Ribaut

(Brolemann)

(Faes)

(Ribaut)

(Schubart)

(Ribaut)

(Demange)

(Ribaut)

(Leach)

(Ribaut)

(Brolemann)

Mauries & Geoffroy

Ribaut

Brolemann

Ribaut

Maurifes

Mauries

1913 GLOMER1DA
1 908 GLOMERIDA
1913 GLOMERIDA
1 96 1 GLOMERIDA
1954 GLOMERIDA

1 907 GLOMERIDA

1908 POLYZONIIDA
1908 POLYZONIIDA
1831 POLYZONIIDA
1898 PLATYDESMIDA
1943 CALLIPODIDA
1819 CALLIPODIDA

Glomeridea

Polyzoniidea

Platydesmidea

Callipodidea

Craspedosomatidea

Craspedoso mat i dea

Craspedosomatidea

1910 CALLIPODIDA
1898 CHORDEUMATIDA
1913 CHORDEUMATIDA

1911 CHORDEUMATIDA

1917 CHORDEUMATIDA

1967 CHORDEUMATIDA
1956 CHORDEUMATIDA

1918 CHORDEUMATIDA
1949 CHORDEUMATIDA
1947 CHORDEUMATIDA
1956 CHORDEUMATIDA
1913 CHORDEUMATIDA

1968 CHORDEUMATIDA
1954 CHORDEUMATIDA
1902 CHORDEUMATIDA

1 902 CHORDEUMATIDA

1969 CHORDEUMATIDA

1903 CHORDEUMATIDA
1920 CHORDEUMATIDA
1 9 26 CHORDEUMATIDA
1956 CHORDEUMATIDA
1927 CHORDEUMATIDA
1951 CHORDEUMATIDA
1920 CHORDEUMATIDA
1963 CHORDEUMATIDA
1913 CHORDEUMATIDA
1903 CHORDEUMATIDA
1965 CHORDEUMATIDA
1963 CHORDEUMATIDA
1925 CHORDEUMATIDA
1913 CHORDEUMATIDA
1951 CHORDEUMATIDA
1975 CHORDEUMATIDA
1951 CHORDEUMATIDA
1913 CHORDEUMATIDA
1910 CHORDEUMATIDA
1814 CHORDEUMATIDA
1 898 CHORDEUMATIDA
1975 CHORDEUMATIDA

1951 CHORDEUMATIDA

1970 CHORDEUMATIDA
1902 CHORDEUMATIDA
1 969 CHORDEUMATIDA
1 969 CHORDEUMATIDA
1969 CHORDEUMATIDA
1965 CHORDEUMATIDA
1882 CHORDEUMATIDA
1932 CHORDEUMATIDA
1913 CHORDEUMATIDA

1952 CHORDEUMATIDA
1968 CHORDEUMATIDA
1952 CHORDEUMATIDA
1930 CHORDEUMATIDA
1 902 CHORDEUMATIDA
1920 CHORDEUMATIDA
1920 CHORDEUMATIDA
1905 CHORDEUMATIDA

1958 CHORDEUMATIDA
1947 CHORDEUMATIDA
1967 CHORDEUMATIDA
1913 CHORDEUMATIDA
1814 CHORDEUMATIDA
1913 CHORDEUMATIDA
1900 CHORDEUMATIDA
1982 CHORDEUMATIDA
1922 CHORDEUMATIDA
1922 CHORDEUMATIDA
1922 CHORDEUMATIDA
1932 CHORDEUMATIDA
1913 CHORDEUMATIDA
1990 CHORDEUMATIDA
1970 CHORDEUMATIDA
1974 CHORDEUMATIDA

1959 CHORDEUMATIDA
1959 CHORDEUMATIDA

THE FRENCH MILLIPEDE SURVEY

277

1 1 7 Pyrgocyphosoma dalmazzense
1 1 8 Pyrgocyphosoma doriae
1 1 9 Rhymogona cervina
I 20 Rhymogona alemannica
1 2 I Rhymogona montivaga

122 Scutogona jeanneli

123 Semiosoma bardei
1 24 Semiosoma devil lei

125 Vandeleuma vasconicum

126 Vascosoma coiffailt

127 Vascosoma coiffaiti falsaforma

128 Vascosoma duprei

129 Xylophageuma zschokkei
1 30 Xysirosoma beatense

1 3 1 Xysirosoma cassagnaui

132 Xysirosoma caialnnicum

133 Xysirosoma muricum

I 34 Xysirosoma pyrenaicum
I 35 Xysirosoma lectosagum

1 36 Chordeuma consoranense

137 Chordeuma iluronense

138 Chordeuma inornatum

139 Chordeuma intermedium

1 40 Chordeuma monlanum
1 4 1 Chordeuma muticum

1 42 Chordeuma proximum
143 Chordeuma reflexum
1 44 Chordeuma silvesire

145 Chordeuma irifidum

146 Chordeuma uiriculosum
I 4 7 Chordeuma vasconicum

148 Melogona gallica

149 Melogona scuiellare

1 50 Mycogona germanica
1 5 1 Orthochordeumella fulva

152 Orthochordeumella leclerci

153 Orthochordeumella pallida

1 54 Orihochordeumella pyrenaica
155 Parachordeuma broelemanni
1 56 Oxidus gracilis

157 Stosalea italica

158 Archipolydesmus rihauli

159 Brachydesmus exiguus

1 60 Brachydesmus proximus
I 6 1 Brachydesmus super us
1 62 Devi Ilea tuberculata
163 Eumastigonodesmus boncii
1 64 Galliocookia balazuci

165 Galliocookia fagei

166 Galliocookia leclerci

167 Macrosiernodesmus palicola

168 Mastigonodesmus destefani
1 69 Mastigonodesmus fagniezi
170 Mastigonodesmus lopezi

I 7 1 Occiianocookia hirsuia
172 Ophiodesmus albonanus
I 7 3 Perapolydesmus progressus
174 Polydesmus anguslus
1 7 5 Polydesmus asihenesiaius

176 Polydesmus barbierii

177 Polydesmus coriaceus

178 Polydesmus corsicus

179 Polydesmus denticulalus
1 80 Polydesmus germanicus

I 8 1 Polydesmus helveticus
1 82 Polydesmus incisus
183 Polydesmus inconslans
1 84 Polydesmus mistrei
185 Polydesmus niveus
I 86 Polydesmus plicatus
187 Polydesmus racovilzai
1 88 Polydesmus raffardi
1 89 Polydesmus laranus
1 90 Polydesmus testaceus
I 9 I Polydesmus troglobius

192 Alpiobaies peyerimhoffi

193 Archiboreoiuius pallidus
1 94 Archiboreoiuius sollaudi
195 Boreoiulus dollfusi

1 96 Blaniulus guiiulaius

197 Blaniulus lichiensieini

198 Blaniulus lorifer

199 Blaniulus mayeti

200 Blaniulus orientalis

20 1 Blaniulus troglobius

202 Blaniulus troglodites

203 Blaniulus velalus

Verhoeff

1930

CHORDEUMATIDA

Craspedosomatidea

(Silvestri)

1898

CHORDEUMATIDA

Craspedosomatidea

(Verhoeff)

1910

CHORDEUMATIDA

Craspedosomatidea

(Verhoeff)

1910

CHORDEUMATIDA

Craspedosomatidea

(Verhoeff)

1894

CHORDEUMATIDA

Craspedosomatidea

Ribaut

1913

CHORDEUMATIDA

Craspedosomatidea

Ribaut

1913

CHORDEUMATIDA

Craspedosomatidea

(Brolemann)

1901

CHORDEUMATIDA

Craspedosomatidea

Mauries

1966

CHORDEUMATIDA

Craspedosomatidea

Maurifcs

1966

CHORDEUMATIDA

Craspedosomatidea

Mauries

1990

CHORDEUMATIDA

Craspedosomatidea

Mauries

1990

CHORDEUMATIDA

Craspedosomatidea

Bigler

1912

CHORDEUMATIDA

Craspedosomatidea

Ribaui

1927

CHORDEUMATIDA

Craspedosomati dea

Mauries

1965

CHORDEUMATIDA

Craspedosomatidea

Ribaut

1927

CHORDEUMATIDA

Craspedosomatidea

Ribaut

1927

CHORDEUMATIDA

Craspedosomatidea

Ribaut

1927

CHORDEUMATIDA

Craspedosomatidea

Ribaut

1927

CHORDEUMATIDA

Craspedosomatidea

Ribaut

1956

CHORDEUMATIDA

Chordeumatidca

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

Ribaut

1956

CHORDEUMATIDA

Chordeumatidea

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

Brolemann

1927

CHORDEUMATIDA

Chordeumatidea

C. Koch

1847

CHORDEUMATIDA

Chordeumatidea

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

Ribaut

1913

CHORDEUMATIDA

Chordeumatidea

(Latzel)

-1884

CHORDEUMATIDA

Chordeumatidea

(Ribaut)

1913

CHORDEUMATIDA

Chordeumatidea

(Verhoeff)

1892

CHORDEUMATIDA

Chordeumatidea

(Roihenbiihler)

1899

CHORDEUMATIDA

Chordeumatidea

Mauries

1979

CHORDEUMATIDA

Chordeumatidea

(Roihenbiihler)

1899

CHORDEUMATIDA

Chordeumatidea

Mauries

1965

CHORDEUMATIDA

Chordeumatidea

Ribaut

1912

CHORDEUMATIDA

Chordeumatidea

(C.L. Koch)

1847

POLYDESMIDA

Paradoxosomatidea

(Latzel)

1886

POLYDESMIDA

Paradoxosomati dea

(Brolemann)

1926

POLYDESMIDA

Polydesmidea

Brolemann

1894

POLYDESMIDA

Polydesmidea

Latzel

1889

POLYDESMIDA

Polydesmidea

Latzel

1884

POLYDESMIDA

Polydesmidea

Brolemann

1902

POLYDESMIDA

Polydesmidea

(Brolemann)

1908

POLYDESMIDA

Polydesmidea

Mauries

1983

POLYDESMIDA

Polydesmidea

Ribaut

1954

POLYDESMIDA

Polydesmidea

Mauries

1983

POLYDESMIDA

Polydesmidea

Brolemann

1908

POLYDESMIDA

Polydesmidea

Silvestri

1898

POLYDESMIDA

Polydesmidea

Mauries

1982

POLYDESMIDA

Polydesmidea

Mauries

1980

POLYDESMIDA

Polydesmidea

(Ribaut)

1948

POLYDESMIDA

Polydesmidea

(Latzel)

1895

POLYDESMIDA

Polydesmidea

(Brolemann)

1900

POLYDESMIDA

Polydesmidea

Latzel

1884

POLYDESMIDA

Polydesmidea

Pocock

1894

POLYDESMIDA

Polydesmidea

Latzel

1889

POLYDESMIDA

Polydesmidea

Porath

1870

POLYDESMIDA

Polydesmidea

Schubart

1931

POLYDESMIDA

Polydesmidea

C. Koch

1847

POLYDESMIDA

Polydesmidea

Verhoeff

1896

POLYDESMIDA

Polydesmidea

Verhoeff

1894

POLYDESMIDA

Polydesmidea

Brolemann

1921

POLYDESMIDA

Polydesmidea

Latzel

1884

POLYDESMIDA

Polydesmidea

Brolemann

1902

POLYDESMIDA

Polydesmidea

Brolemann

1900

POLYDESMIDA

Polydesmidea

Ceuca

1962

POLYDESMIDA

Polydesmidea

Brolemann

1910

POLYDESMIDA

Polydesmidea

Brolemann

1905

POLYDESMIDA

Polydesmidea

Verhoeff

1936

POLYDESMIDA

Polydesmidea

C.L. Koch

1847

POLYDESMIDA

Polydesmidea

Latzel

1889

POLYDESMIDA

Polydesmidea

(Brolemann)

1900

JULIDA

Blaniulidea

(Brade-Birks)

1920

JUL1DA

Blaniulidea

Brolemann

1921

JULIDA

Blaniulidea

Brolemann

1895

JULIDA

Blaniulidea

(Fabricius)

1798

JULIDA

Blaniulidea

Brolemann

1921

JULIDA

Blaniulidea

Brolemann

1921

JULIDA

Blaniulidea

(Brolemann)

1902

JULIDA

Blaniulidea

Brolemann

1921

JULIDA

Blaniulidea

(Latzel)

1886

JULIDA

Blaniulidea

Brolemann

1898

JULIDA

Blaniulidea

Ribaut

1954

JULIDA

Blaniulidea

Source : MNHN, Paris

278

JEAN-JACQUES C.EOFEROY

204 B lam ulus virei

205 Boreoiultis simplex

206 Boreoiulus tenuis

207 Choneiulus palmatus

208 Choneiulus subterraneus

209 Galliobates gracilis
2 1 0 Iberoiulus sarensis

2 1 I Mesobluntulus serrula

212 Munacobates monoecensis

213 Thalassisobates litloralis

214 Nemasoma varicorne

215 Nopoiulus kochii

216 Occituniulus rouchi

2 I 7 Proteroiulus broelemanni
2 I 8 Proteroiulus fuscus

219 Trichoblaniulus hirsutus

220 Trichoblaniulus lanuginosus

221 Vascoblamulus cabidocliei

222 Allajulus nitidus

223 Brachxiulus lusitanus
2 24 Brachviulus pusillus

225 Cxlindroiulus broti

226 Cxlindroiulus caeruleocinctus

227 Cxlindroiulus chalandei

228 Cxlindroiulus iluronensis

229 Cxlindroiulus latest rial us

230 Cxlindroiulus limitaneus

231 Cxlindroiulus londinensis

232 Cxlindroiulus parisiorum

233 Cxlindroiulus punctatus

234 Cxlindroiulus pxrenaicus

235 Cxlindroiulus Sagittarius
2 36 Cxlindroiulus schubarti

237 Cxlindroiulus segregatus

238 Cxlindroiulus spinosus

239 Cxlindroiulus verhoeffi

240 Cxlindroiulus vulnerarius

241 Daltchoiulus tongiorgii

242 Enantiulus armatus

243 Enantiulus nanus

244 Haplopodoiulus spathifer

245 Hypsoiulus alpivagus

246 Julus scandinavius

247 Leptoiulus arelatus

248 Leptoiulus belgicus

249 Leptoiulus bertkaui

250 Leptoiulus brevivelatus

251 Leptoiulus bruyanti

252 Leptoiulus demangei

253 Leptoiulus garumnicus

254 leptoiulus juvenilis

255 Leptoiulus ken’illei

256 Leptoiulus legeri

257 leptoiulus meridionals

258 Leptoiulus montivagus

259 Leptoiulus odieri

260 Leptoiulus piceus

261 Leptoiulus remxi

262 Leptoiulus simplex glacialis

263 Leptoiulus umbratilis

264 Leptoiulus uncinatus

265 Metaiulus pratensis

266 Ommatoiulus albolineatus

267 Ommatoiulus corsicus

268 Ommatoiulus haackert

269 Ommatoiulus ilhcis

270 Ommatoiulus imminutus

271 Ommatoiulus lienhardti

272 Ommatoiulus rutilans

273 Ommatoiulus sabulosus

274 Ophxiulus bastiensis

275 Ophxiulus chilopogon

276 Ophxiulus corsicus

277 Ophxiulus napolitanus

278 Ophxiulus pilosus

279 Ophxiulus renosensis

280 Pachyiulus varius

281 Tachypodoiulus niger

282 Typhloiulus sculterorum

Brolemann

(Bigler)

(Ncmec)

(Silvesiri)

Ribaut

Mauries

(Brolemann)

(Silvesiri)

C.L.Koch

(Gervais)

Brolemann
Lohmander
(Am Slein)

(Brolemann)

Ribaut

Mauries

(Verhoeff)

(Leach)

(Humbert)

(Wood)

(Ribaut)

Brolemann

(Curtis)

(Brolemann)

(Leach)

(Brolemann & Verhoeff)
(Leach)

Brolemann

Verhoeff

Brolemann

(Ribaut)

(Brolemann)

(Berlese)

(Strasser)

(Ribaut)

(Latzel)

(Brolemann)

(Verhoeff)

Latzel

Bigler

(Latzel)

(Verhoeff)

Bigler

Ribaut

Schubart

(Ribaut)

(Brolemann)

(Latzel)

(Brolemann)

(Risso)

Schubart

(Verhoeff)

(Ribaut)

Ribaut

Blower & Rolfe
(Lucas)

(Brolemann)

Mauries

(Brolemann)

(C.L Koch)

(Linn6)

Verhoeff

(Latzel)

Verhoeff

(Attems)

(Newport)

Mauries

(Fabricius)

(Leach)

(Brolemann)

1900

JUL1DA

1921

JULIDA

1913

JULIDA

1895

JULIDA

1903

JULIDA

1909

JULIDA

1970

JULIDA

1905

JULIDA

1905

JULIDA

1903

JULIDA

1847

JULIDA

1847

JULIDA

1923

JULIDA

1925

JULIDA

1857

JULIDA

1889

JULIDA

1947

JULIDA

1967

JULIDA

1891

JULIDA

1898

JULIDA

1815

JULIDA

1893

JULIDA

1864

JULIDA

1904

JULIDA

1912

JULIDA

1844

JULIDA

1905

JULIDA

1814

JULIDA

1896

JULIDA

1815

JULIDA

1897

JULIDA

1897

JULIDA

1943

JULIDA

1903

JULIDA

1904

JULIDA

1896

JULIDA

1888

JULIDA

1973

JULIDA

1909

JULIDA

1884

JULIDA

1897

JULIDA

1897

JULIDA

1884

JULIDA

1919

JULIDA

1884

JULIDA

1896

JULIDA

1919

JULIDA

1951

JULIDA

1962

JULIDA

1904

JULIDA

1908

JULIDA

1896

JULIDA

1897

JULIDA

1897

JULIDA

1884

JULIDA

1896

JULIDA

1826

JULIDA

1962

JULIDA

1894

JULIDA

1905

JULIDA

1951

JULIDA

1956

JULIDA

1845

JULIDA

1903

JULIDA

1969

JULIDA

1897

JULIDA

1926

JULIDA

1921

JULIDA

1847

JULIDA

1758

JULIDA

194 3

JULIDA

1884

JULIDA

1943

JULIDA

1903

JULIDA

1842

JULIDA

1969

JULIDA

1781

JULIDA

1815

JULIDA

1905

JULIDA

Blaniulidea

Julidea

THE FRENCH MILLIPEDE SURVEY

279

AC KNO WLEGEM ENTS

I am very grateful to my colleagues J. P. MauriES (Paris), R. D. Kime (Bruxelles) and A. Pedroli-Christen
(Neuchatel) for providing new localities and information. Many thanks to P. Mothiron for discussions and advice related
to the computer database. Comments of Prof. J. G. BLOWER and Dr. A. D. Barber on the English version of this
manuscript are gratefully acknowledged.

REFERENCES

Barbault. R.. 1994. — Des baleines, des bacteries et des hommes. Paris, Odile Jacob, Sciences, 327 pp.

Bi.andin, P.. 1989. — Sur la richesse spScifique et la rarete comme criteres devaluation de systemes ecologiques. In : F.
DH BEAUFORT & H. MAURIN, Utilisation des inventaires d'invertebres pour i identification et la surveillance d'espaces
de grand interet faunistique . Paris, SFF, MNHN : 71-80.

Blower, J. G., 1985. — Millipedes ( Synopses of the Br. Fauna NS, 35). London, E. J. Brill & W. Backhuys, 242 pp.

BRITISH Myriapod Group, 1988. — Preliminary Allas of the Millipedes of the British Isles. Huntingdon, NERC, 65 pp.

Culver, D. C. & Holsinger, J. R., 1992. — How many species of troglobites are there? NSS Bulletin , 54 : 79-80.

ENGHOFF, H., 1974. — Om tusindbenenes udbredelse i Danmark (Diplopoda). Ent. Meddr., 42 : 21-32.

Erwin, T. L., 1988. — The tropical forest canopy: the heart of biotic diversity. In : E. O. Wilson. Biodiversity.
Washington D. C., National Academic Press : 123-129.

Geoffroy. J. J., 1981. — Les myriapodes du Parc national des Ecrins. 1. Stations de recolte (1976-1979) et presentation
g£n£rale des peuplements de chilopodes et de diplopodes. Trav. Sci. Parc Natl. Ecrins, 1 : 97-123.

Geoffroy. J. J., 1984a. — Particularity du developpement 'post-embryonnaire du diplopode craspedosomide
cavemicolc Opisthocheiron canayerensis. Mem. Biospeol., 11 : 211-220.

Geoffroy, J. J.. 1984b. — Opisthocheiron canayerensis (Diplopoda : Craspedosomatida) : repartition de Fespece et
variations de la pigmentation. Mem. Biospeol ., 11 : 295-302.

Geoffroy, J. J., 1 989. — La faune des diplopodes de France. 2 : projet de cartographic des especes. In : F. de Beaufort
& H. MAURIN, Utilisation des inventaires d'invertebres pour I' identification et la surveillance d'espaces de grand
interet faunistique. Paris, SFF, MNHN : 270-271.

Geoffroy, J. J., 1990a. — La faune des diplopodes de France : un bilan des especes. In : A. Minelli, Proc. 7th Ini.
Congr. Myriapodology. Leiden, Brill : 345-359.

Geoffroy. J. J., 1990b. — Les diplopodes cavernicoles de France. Mem. Biospeol.. 17 : 3-11.

Geoffroy. J. J., 1991. — Les cavites artificielles et la repartition des diplopodes endog6s et souterrains : interet
biogeographique. Rev. Suisse Zool.. 98 : 93-106.

Geoffroy, J. J., 1992. — Cle d’identification des classes de myriapodes et des ordres de chilopodes frequents dans le sol
et ses annexes. Millepattia, 1 : 23-37.

Geoffroy, J. J., 1993a. — Cles d’identification des ordres de diplopodes frequents dans le sol. (Myriapoda ; Diplopoda).
Millepattia, 2 . 4-28.

GEOFFROY, J. J., 1993b. — L’inventaire et la cartographic des diplopodes de France. In: : J. LHONORE, H. Maurin,
R GuiLBOT & P. KEITH. Inventaire et cartographic des invertebres comme contribution a la geslion des milieux
naturels frangais. Coll. Patrimonies Naturels, 13. Paris, SSF / MNHN : 144-148.

GEOFFROY, J. J., 1994a. — Steps towards a comprehensive distribution of soil and cave-dwelling Diplopoda species. A
perspective on millipede biodiversity in France. In : G. Messana & S. FaiTI. XI International Symposium of
Biospeology, Firenze, Costello di Montegufoni, 28 August - 2 September 1994 (Italy), Abstracts Firenze.
Universita degli Studi di Firenze (65pp.).

Geoffroy, J. J., 1994b. — Fauna Gallica Diplopoda : Etapes pour une etude de la repartition et de la biodiversite des
diplopodes de France. In : B. CONDE. 98eme Journees de la Societe Zoologique de France, Nancy, 4-5-6 Juillet 1994,
Resume des communications Nancy, Universite Henri Poincan§ : 55.

Geoffroy, J. J., & Mauris, J. P.. 1992. — Les diplopodes edaphiques et souterrains de France : donnees recentes,
repartition des especes nouvellement decrites et peu connues. Mem. Biospeol., 19 : 127-133.

GONSETH, Y., 1993. — Le CSCF. un relai entre les recherches 6co-faunistiques et la pratique. In: : J. Lhonore, H.
Maurin, R. Guilbot & P. KEITH, Inventaire et cartographic des invertebres comme contribution a la gestion des
milieux naturels frangais. Coll. Patrimonies Naturels. 13. Paris, SSF / MNHN : 51-57.

JEEKEL, C. a. W.. 1978. — Voorlopigc atlas van de verspreiding der Nederlandse miljoenpoten (Diplopoda). Verslagen
en Technische Gegevens . 15. Amsterdam, Instituut voor Taxonomische Zoologie, 68 pp.

JUBERTHIE, C. & DECU, V., 1994. — Structure et diversile du domaine souterrain ; particularity des habitats et
adaptations des especes. In : C. Juberthie & V. DECU, Encyclopedia Biospeologica. Tome I. Moulis-Bucarest,
Soci6t6 de Biospeologie : 5-22.

Source :

280

JEAN-JACQUES GEOFFROY

Kime, R. D.. 1990. — A provisional alias of European myriapods. Part I. Luxembourg, European Invertebrate Survey,

1 09pp.

LEVEQUE, C. 1994. — Environnement et diyersite du vivant. Paris, CSI Pocket / ORSTOM, 127 pp.

Maurer. B. A.. 1994. — Geographical population analysis: tools for the analysis of biodiversity. Oxford, Blackwell
Scientific Publications, 130pp.

MAURlfcS J P 1982. — Dolichoiulus tongiorgii (Strasser), diplopode halophilc nouveau pour la faune de France.
Remarques sur la classification des Pachyiulini. Bull. Mus. nail. Hist, nat., Paris, (A). 4 : 433-444.

MAURifeS J p & GEOFFROY, J. J., 1982. — Decouverte. dans les Causses Majeurs, d'une remarquable espece cavemicole
du genre Opisthocheiron Ribaut, 1913 (Dipiopoda. Craspedosomida, Opisthocheindae). Bull . See. Hist. nat.
Toulouse, 1 18 : 131-140.

MAUR1N. H. & Guilbot, R., 1993. — La cartographic des invertebres et la gestion des milieux naturels. In: : J. LHONORE,
H Maurin, R. GUILBOT & P. KEITH. Inventaire et cartography des invertebres comme contribution a la gestion des
milieux naturels frangais. Coll. Patrimoines Naturels, 13. Paris, SSF / MNHN : 9-17.

May, R. M., 1992. — L'inventaire des esp&ces vivantes. Pour la Science, 182 : 30-36.

M othiron. P.. 1993. — Un exemple regional : l'inventaire des lepidoptfcres d Ile-de-France. In: : J. Lhonore,
H. Maurin, R. Guilbot & P. Keith, Inventaire et cartography des invertebres comme contribution a la gestion des
milieux naturels frangais. Coll. Patrimoines Naturels, 13. Paris, SSF / MNHN 103-105.

Pedroli-Christen, A., 1993. — Faunistique des mille-pattes de Suisse (Dipiopoda) / Fauntslik der Tausendf ussier der
Schweiz ( Dipiopoda ). Neuchatel. Centre Suisse de Cartographie de la Faune, Doc. faun, helv 14, 248 pp.

Rasmont. P.. 1993. — Methodologie et outillage de la cartographie <§cologique des invertebres. In\ : J. Lhonore.
H. Maurin, R. Guilbot & P. Keith, Inventaire el cartography des invertebres comme contribution a la gestion des
milieux naturels frangais. Coll. Patrimoines Naturels. 13. Paris, SSF / MNHN : 28-50.

Remy. P. & HOFFMAN, J.. 1959. — Faune des Myriapodes du Grand-Duche de Luxembourg. Archives de la Section des
Sciences de I’Institut Grand-Ducal , 26 : 199-236.

Serra-Cobo. J.. Barbault, R. & Estrada-Pena, A., 1993. - Le gouffre dc San Pedro de Los Griegos (Oliete, Teruel,
Espagne) : un refuge de biodiversite sans equivalent en Europe. Rev. Ecol. ( Terre Vie), 48 : 341-348.

Schubart, O., 1934. — Tausendfussler Oder Myriapoda. I: Dipiopoda. Tierwelt Deutchi, 28 : i-viii + 1-318.

Solbrig. O. T., van Emden, H. M. & van Oordt, P. G., (eds), 1992. — Biodiversity and Global Change. Paris, IUBS
Monographs , 8.

Strasser, C. & Minelli, A., 1984. — Elenco dei diplopodi d'ltalia. Lavori-Soc. Ven. Sc. nat.. 9 : 193-212.

Wilson. E. O, 1993. — La diversite de la vie. Paris, Odile Jacob, Sciences, 496 pp.

Source :

Faunistique des mille-pattes de Suisse (Diplopoda) -
Faunistik der Tausendfussler der Schweiz (Diplopoda)

Ariane PEDROLI-CHRISTEN

C.S.C.F., Musee d'Histoire Naturelle, Terreaux 14, CH-2000 Neuchatel, Switzerland

RESUME

Une synthese et un constat actuel et critique des connaissances relatives aux diplopodes de Suisse sont proposes dans
cet ouvrage (Pedroli-Christen, 1993), tant du point de vue systematique, chorologique qu’tScologique. Si 1'origine des
donndes permettant cette evaluation est lr£s disparate (donndes bibliographiques, collections, nkoltes recentes),
l'ampleur des observations permet cependant d’effecluer une analyse quantitative. Le catalogue complet des 14300
donndes est depose au Centre Suisse de Cartographic de la Faune, CSCF. La composition de la faune des diplopodes de
Suisse peut etre consid£ree actuellemenl comme relativement bien connue : 127 taxons identi fi6s sont repartis dans 6
ordres et 18 families. Plusieurs difficultes systematiques ont ete mises en evidence, dues au concept purement typologique
de l'espece, generalement en vigueur en myriapodologie. Elies sont egalement 1'expression de rimportance des facteurs
historiques sur ce groupe. Pour chaque espfcce, les resultats obtenus sont discutes et illustres selon un schema standard .
rapidc synonymie, references bibliographiques synth6tiques des meilleures descriptions et illustrations, references
bibliographiques helvetiques, aspects chorologiques, ecologiques et phenologiques, tableau synthetique du nombre
d'observations effectives et carte de repartition. Une etude qualitative de l'ensemble de la faune reprend plusieurs des
parametres retenus dans l'analyse par espece en considerant cette fois l'organisation des peuplements. Divers
groupements d'especes sont ainsi proposes selon leur repartition geographique. leur distribution altitudinale et les types
de milieux colonises.

ABSTRACT

Faunistics of the Swiss millipedes (Diplopoda).

An up-to-date and critical synthesis of the present knowledge of the Swiss Diplopoda is given in this book (Pedroli-
Christen, 1993) in the scope systematics, distribution and ecology. In spite of the lack of uniformity of the original
data (bibliographical, museum collections, recent collectings), the high number of observations enable a quantitative
analysis. The complete catalogue of captures is deposited at the Centre Suisse de Cartographie de la Faune, CSCF. The
Swiss Diplopoda fauna is now rather well known: 127 identified taxa belonging to 6 orders and 18 families. Several
systematical obstacles were encountered, mainly due to the purely typological concept of species used in
myriapodology. They also indicate the influence on this group of historical factors. For each species the results are
discussed and presented in a standard way: a rapid synonymy, bibliographical references of the best descriptions and
illustrations, bibliographical references concerning Switzerland, distribution, ecology and phenology, number of
observations and distribution map. A qualitative study of the whole fauna, considering populations, is based on the same
parameters. Various species-groups are defined after geographical distributions, altitudinal distributions and habitats
occupied.

Pedroli-Christen, A.. 1993. — Faunistique des mille-pattes de Suisse (Diplopoda) / Faunistik der Tausendfussler der
Schweiz (Diplopoda). Neuchatel, Centre Suisse de Cartographie de la Faune. Doc. faun, helv., 14, 248 pp.

Pedroli-Christen, A., 1996. — Faunistique des mille-pattes de Suisse (Diplopoda) - Faunistik der Tausendfussler
der Schweiz (Diplopoda). In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M.. (eds), Acta
Myriapodologica. Mem. Mus. nain. Hist, nat., 169 : 281. Paris ISBN : 2-85653-502-X.

Source : MNHN, Paris

On Myriapod / Insect Interrelationships

Otto Kraus & Margarete KRAUS

Zoologisches Institut und Zoologisches Museum, Universitat Hamburg, Martin-Luther-King-Platz 3,

D-20146 Hamburg. Germany

ABSTRACT

In the Tracheata (= Antennata), all non-insect taxa are traditionally classified as “Myriapoda". New insights suggest
that this may be mistaken. There is good reason to believe that the Chilopoda form the sister taxon ol all other
Tracheata. Further, a monophylelic unit formed by all progoneate taxa (Symphyla + Pauropoda + Diplopoda) is the most
probable sister taxon of the Insecta (= Hexapoda). Hence, Progoneata + Insecta also form a monophylum. Phis taxon
(sister taxon to the chilopods) is called Labiophora. The insects are maintained as a monophylelic unit. There is no
reason to separate the Collembola (as “Parainsecta") from the remaining "true" insects. - Available evidence suggests
that the basic phylogenetic branching events in the ’‘myriapods” and also in the insects into higher taxa happened very
early, presumably in Late Cambrian/ Early Silurian periods.

RESUME

Sur les interrelations entre myriapodes et insectes.

Chez les Tracheata ou Antennata, lous les non-insectes sont tradilionnellement considers comme “Myriapoda". De
nouvelles donndes suggerent que cela pourrait etre errone. II y a de bonnes raisons de penser que le groupe Chilopoda
constitue le taxon-frere de tous les autres Tracheata. D'autre part, Funite monophyletiquc formee par tous les Progoneata
(Symphyla, Pauropoda et Diplopoda) est le groupe-frere des insectes le plus probable. Desormais. Progoneata + Insecta
forment aussi un groupe monophyletique (taxon frerc des chilopodes) appele Labiophora. Les insectes sont maintenus
en tant qu‘ unite monophyletiquc, car il n’y a pas de raison d’en sdparer les collemboles sous le nom de Parainsecta. On
peut valablement penser que l’evenement instituant la base phylogenetique des myriapodes et aussi des insectes parmi
les autres grands taxons se produisit tres lot, probablcment dans la periode Cambrien supericur - Silurien inferieur.

INTRODUCTION

Most authors, and especially textbook authors, continue to maintain the traditional view
that myriapods form a taxon, i.e., a monophyletic unit. But various phylogeneticists feel that a
group called "Myriapoda” should be regarded as paraphyletic and therefore be abandoned.
Controversial discussions of the question which subtaxon of the so-called Myriapoda might be
most closly related to the Hexapoda (= Insecta) go back to the early days of POCOCK (1893) and
VERHOEFF (e.g.. 1910-1914). On the other hand, the concept of the Myriapoda as a taxon was
upheld by HENNIG (1969) and also by BOUDREAUX (1987).

There is no reason to question the monophyletic origin of the Tracheata (= Antennata) as a
whole, but this assumption should not be based on the presence ot tracheae as a character.
Convergent evolution of tracheal systems cannot be excluded and is, perhaps, even probable.

Kraus. O. & Kraus. M. 1996. — On Myriapod / Insect Interrelationships. In: Geoffroy. J.-J.. Mauri£s. J.-P
& Nguyen Duy - Jacquemin. M„ (eds). Acta Myriapodologica. Mem. Mus. natn. Hist. run.. 169 : 283-290. Paris ISBN
2-85653-502-X.

284

OTTO KRAUS & MARGARETE KRAUS

But there are other, more reliable characters available that should be regarded as autapomorphies
of the Tracheata (Fig. 1). For example, the second pair of antennae has been reduced, but its
metamere still forms part of the head capsule and is called the intercalary segment. Furthermore,
for reasons to be explained below, all tracheates lack a mandibular palpus. In the present paper,
we attempt a step-by-step reconstruction of early phylogenetic branching events within the
tracheates. In some instances, the fossil record permits estimation of the phylogenetic age of
various subtaxa - according to HENNlG’s terminus post quern non (see e.g., 1969).

MATERIAL AND METHODS

As usual, much of ihe relevant data is already available and can be derived from previously published papers.
Major problems were experienced with reference to the composition of the head capsule and the homology of
components of the mouthparts. Specimens preserved in alcohol or BoutN's fixative were dissected and studied by means
of light microscopy (Leitx interference contrast according to Smith), and also by scanning electron microscopy (SEM).
For liaht microscopy, chitinous parts were mounted on slides; for embedding, we used HOYER s mixture as this medium
has an advantageous light refraction index (for details: see Kraus, 1984). A camera lucida was used for all drawings.

RESULTS

In this sectio, we deal mainly with uncertainties concerning the composition of the
euarthropodean head capsule (see BOUDREAUX, 1987: 120, 121) and investigate various types
of mouthparts. The latter part of the investigation is concentrated on the homology of mandibles
and on the interpretation of components of the gnathochilarium in Diplopoda and Pauropoda (=
Dignatha).

Crustacea

Chilopoda

Progoneata

Symphyla Dignatha

Pauropoda Diplopoda

Pselaphogn Chilogn.

Nauplius eye
Mandibles gnathobasic

27.

tej

Tracheata

Labiophora

Entognatha

Diplura Ellipura

Protura Collemb.

Insecta

Ectognatha

Archaeogn. — Dicondylia

Zygeni. Pteryg

I |42

Terfe$tria!i*ation (lx?)

Stem lineage Mandlbulata

Fig. 1. — Phylogenetic relationships between higher taxa of the Tracheata, their outgroup (Crustacea) included. Arrows
indicate age of earliest hitherto known fossils of various groups. — LC, Lower Cambrium; LD, Lower Devonian;
MD, Middle Devonian; US, Upper Silurian.

Source :

ON MYRIAPOD / INSECT RELATIONSHIPS

285

Segments of the head capsule

It is now generally accepted that the euarthropodean head capsule includes an acron
followed by at least 5 (early fossils), in modern representatives by 6 metameres (LAUTERBACH,
1980a, b; WALOSSECK, 1993: 111). Various authors, however, have believed that the head
capsule of the Dignatha is made up of only by 5 metameres. The question arises of whether the
regular 6th segment in Pauropoda (TlEGS, 1947: 304) and in at least in Pselaphognatha
(Diplopoda) (ATTEMS, 1926: 109) was secondarily excluded from the head capsule or whether
it had not yet been fully included. A third alternative would be that it was and is included.

Homology of components of the gnathochilarium

The question of homology is directly concerned with the old problem as to whether the
gnathochilarium is made up primarily of the maxillae I or by both pairs of appendages, maxillae
I and II. VERHOEFF in particular (e.g., 1910-1914), argued that gnathochilaria included two
pairs of appendages. This is in contradiction to data derived from ontogenetic studies (DOHLE,
1964, 1980): in the ontogeny of Glomeris marginata, the mandibles are followed by only one
pair of prominent ornaments of appendages. Nonetheless, it remains quite uncertain whether
this can be regarded as proof for the assumption that the gnathochilarium does not include
elements derived from two pairs of appendages.

Pselaphognatha

A study of the gnathochilarium in Pselaphognatha seems to supply the key to solution of
the problem: there is no gnathochilarium at all in these diplopods! In Polyxenus, the mandibles
are followed by two (!) pairs of appendages (Fig. 2a, b). The posterior one shows a very broad
and partially bipartite basal plate. This piece bears a pair of appendages. They are equipped with
numerous sensillae; we interprete them as leg-like telepodites of the maxillae II. Further, the
reader is referred to the presence of traces of articulations between segments of these
appendages (Figs. 2a). In a somewhat lateral position, another pair of appendages is present in
front of these 2nd maxillae and posterior to the mandibles: these parts still show vestiges of
segments. We refer to the position of the duct of the “Putzdriise” (VERHOEFF’s term) and
interpret these parts as maxillae I. There is no reason to believe that they might be part of the
hypopharynx.

Pauropoda

In Pauropods, the head capsule also bears a posterior component which was designated
“intermaxillary plate” by TlEGS (1947: 182); this structure does not bear any appendages. In
agreement with the arrangement of mouthparts in Polyxenus, we interpret the subtriangular plate
as representing the maxillae II. In addition, distinct lateral and segmented mouthparts are also
present. It was TlEGS (1947), who clearly illustrated their position between the anterior
mandibles and the posterior “intermaxillary plate” (see his Fig. 2; also PI. 3 Fig. 33A). We have
studied brachypauropodids and especially Hexamerocerata and can confirm that the position of
these paired appendages is between the mandibles and maxillae II (Fig. 3). The obvious
interpretation is that they represent the maxillae I. We find it hard to understand how previous
authors could invent a pauropodean gnathochilarium (see, e.g., DOHLE 1980: 63, 91).

Chilognatha

Only in the Chilognatha is a true gnathochilarium present, forming the well-known
complex unit (VERHOEFF’s “Mundklappe”) with median and lateral components. In our view,
the lateral elements are homologous with the maxillae I and the median elements with the
maxillae II (HlLKEN & KRAUS, 1994; KRAUS & KRAUS, 1994). Apparently, this development

286

OTTO KRAUS & MARGARETE KRAUS

is correlated with the acquisition of new food niches, as the Chilognatha feed on larger food
particles than the Pselaphognatha. Accordingly, the “perfect gnathochilarium should be
understood in terms of constructional morphology.

FIG. 2a, b. — Polyxenus lagurus, mouthparts. a) SEM-pholo, b) drawing. — AR. articulations between segments of
telopodite, more or less reduced; BP, basal plate of maxillae II; MD. mandibles, distal part; MX /, maxillae I;
MX II. maxillae II; T. telepodites of second maxillae with sensillae.

Fig. 3. — Pauropoda. Hexamerocerata ( Millotauropus silvestrii Remy. 1953), lateral view of head capsule; AN. basis ol
antenna; MD. mandible made up by 3 segments; MX I. tip of maxillae 1; MX II, maxillae II.

Fig. 4. — Archaeognatha ( Trigoniophthalmus alternatus Silvestri, 1904), tclognathic mandible showing vestiges ol
original articulations ( AR ) between composing segments.

Being fully aware of the fact that this interpretation is in conflict with data derived from
ontogeny (DOHI.E, e.g., 1980), we argue (i) that there is no other imaginable interpretation ol
the structures present in postembryonic stages, and (ii) that it is not possible to state definitively

Source :

ON MYRIAPOD / INSECT RELATIONSHIPS

287

that features do not exist on the grounds that they have not yet seen or may even have remained
indiscernible during the course of ontogeny. Another, similar, situation concerning the so-called
thoracic segments was described by KRAUS (1990). It is now well established that “true”
diplosomites with only one pair of legs really exist among the diplopods (see ENGHOFF, 1993)!
Hence, one should ask the reverse question: how is it possible that details remain invisible
during the course of ontogeny when they are unquestionably present in postembryonic stages?

In conclusion, there seems to be no reason to doubt that the head capsule in Diplopoda and
Pauropoda includes 6 metameres - as in all other extant euarthropods.

Mandibles

In all mandibulates, the appendages of the fourth cephalic metamere have been
transformed into mandibles. The crustacean mandible is unquestionably gnathobasic
(LAUTERBACH, 1972, 1980a). In the ground pattern of the Crustacea, several distal segments of
this pair of appendages are accordingly represented by a palp.

On the other hand, there has been much dispute about whether the mandibles in the
Tracheata (= Antennata) are also gnathobasic (LAUTERBACH, 1972) or - as MANTON believed
(e.g., 1977) - telognathous. Attention is drawn to the various types of segmented mandibles in
Chilopoda, Symphyla and Diplopoda (see, e.g. MANTON, 1977); the well-known facts on these
can be supplemented. MANTON, who made extensive studies of the head capsule in a species of
the machilid genus Petrobius, did not realize that segment borders are also clearly visible in the
mandibles of the Archaeognatha (Fig. 4). The same is true in the Diplura, at least in
representatives of the genus Dinjapyx (see MARCUS, 1951 ). This finding is correlated with the
absence of a mandibular palp in all tracheate taxa: telognathous mandibles cannot bear a palp!

We therefore have to conclude that the mandibles in the Mandibulata are homologous as
far as they correspond to the appendages of the fourth metamere of the head capsule. Their
transformation into jaws happened independently, however: they are gnathobasic in the
Crustacea, whereas the appendages were suitably modified in the Tracheata as a whole.

DISCUSSION

Reconstruction of phylogenetic branching

There are convincing reasons for believing that the Tracheata are a monophyletic unit:

Combining our results with previously known details and referring to the hypothesis
expressed by the present cladogram (Fig. 1) we argue as set out below:

(1) Arthropodia are generally considered as a key character common to all arthropods.
This may be questionable. In addition, it is not absolutely certain whether the transition to
terrestrial life and the acquisition of uniramous walking legs (LAUTERBACH. 1980a: 147) took
place as early as in the stem lineage of the Tracheata as a whole. See character (24).

(2) As in the ground pattern of the Crustacea, the tracheate maxilla II was primarily leg¬
like (plesiomorphic condition, maintained in chilopods and also in various Pselaphognatha,
e.g., Polyxenus).

(3) Teleognathic mandibles are part of the ground pattern of the tracheates.

(4) Paired tarsal claws were regarded by HENNIG (1969: 89) as an autapomorphy of the
tracheates. This is highly questionable (see, e.g., diplopodean and ellipuran claws).

(5) An intercalary segment presents the third metamere of the head capsule — without
appendages.

(6) No digestive mitgut glands have been maintained in all tracheates. Instead, malpighian
tubules were developed. Nonetheless, the homology ot these organs needs clarilication. At

288

OTTO KRAUS & MARG ARETE KRAUS

present, the possibility that malpighian tubules evolved more than once, and hence may not be
homologous, cannot be excluded.

Chilopoda

(7) The assumed monophyletic origin of the Chilopoda is strongly supported by the
transformation of the appendages of the first postcephalic metamere into “maxillipeds”
(“Kieferfiisse”).

(8) Chilopods are functionally dignathous: their oral cavity is bordered ventrally by the
maxillae I; see character (2).

(9) - (10) Reduction of complex eyes to stemmata; loss of median eyes.

Labiophora

(1 1) The presence of coxal organs, including sty li, is assumed to be an autapomorphy of
the adelphotaxon to the chilopods: Labiophora. But DOHLE’s critical remarks (1980: 86) should
be considered.

(12) According to our inteipretation of the diplopodean gnathochilarium and of the
mouthparts in the Pauropoda, we conclude that in all subordinated taxa the oral cavity is
ventrally bordered by a plate formed by the maxillae II. Hence, all representatives of this major
taxon are functionally trignathous. The presence of special dorsal organs during the course of
ontogeny may constitute another autapomorphy of this group (for details: see DOHLE, 1980:
88).

Progoneata

(13) The anterior position of the genital opening forms a strong autapomorphy of the
taxon Progoneata. The opening is constantly located in front of the fourth pair of legs (but the
first pair may be reduced).

(14) All progoneates have trichobothria peculiar to this taxon and unknown in all other
terrestrial mandibulates. For details see DOHLE (1980: 72).

(15) In contrast to insects, there are no palpi on the maxillae I. It is assumed that the first
maxillae are telognathous in progoneates.

(16) Loss of median eyes.

(17) - (23) Symphyla have many autapomorphies. We will mention only a few: genital
opening unpaired; special position of a single pair of tracheal spiracles; complete reduction of
median and complex eyes; special structure of maxillae II, total loss of telopodites; spermathecae
formed by lateral pockets of the mouth cavity; presence of terminal spinning tubules.

(24) All Dignatha have their tracheal spiracles in a ventral position. Internally, they open
into tracheal pockets. They also serve as apodems. Such pockets are also present in pauropods
(see REMY, 1953: 37).

(25) Reduction of the first pair of postcephalic appendages. Only pauropods have
maintained vestiges: “exsertile vesicles” (see TlEGS, 1947: 182, 249).

(26) Presence of “penes” with openings of the vasal efferentia at the tip.

(27) - (30) Pauropoda have many autapomorphies, including specialized antennae;
exsertile vesicles [see (25)]; pseudoculus; maxillae II transformed into an unpaired triangular
plate.

Diplopoda

(31) Acquisition of diplopody.

(32) Antennae with four sensory cones on tip.

(33) , (35), (37) Complex eyes reduced to 5 isolated ommatidia; gnathochilarium with
separate maxillae I and specialized telopodites of maxillae II maintained; soft cuticle with
conspicuous groups of hairs.

Source : MNHN, Paris

ON MYRIAPOD/ INSECT RELATIONSHIPS

289

(34), (36), (38) Complex eyes reduced to stemmata; “complete” gnathochilarium;
calcification of cuticle; total loss of trichobothria.

Insecta

As far as insects are concerned, we will only mention the presence of a locomotory thorax
made up of the postcephalic metameres I to III (39), and the presence of 1 1 abdominal
metameres in the ground pattern (40). A detailed discussion of phylogenetic branching and
relationships within the Insecta (= Hexapoda) would not be appropriate here, the reader is
refered to the detailed arguments presented in HENNIG’s comprehensive work (1981).

PHYLOGENETIC AGE

The geological age of the earliest fossils presently known is indicated in FIG. 1 (arrows).
According to phylogenetic branching, the same age must be inferred to equivalent sister taxa. So
the presence of Crustacea as early as in Lower Cambrian times indirectly indicates that
representatives of the stem lineage of the Tracheata also existed at this period - irrespective of
the fossil record. The most important aspect within the Tracheata is the existence of Diplopoda
in deposits of Upper Silurian age. This indicates that previous branching events happened
earlier, presumably in Upper Cambrian / Lower Silurian times. It is therefore possible to predict
that chilopods are considerably older than the earliest known fossil ( Devonobius delta Shear,
1988).

AC KNO WLEDGEMENTS

Wc are indebted to Prof. Dr. W. Dunger (Gorlitz) and Prof. Dr. R. Willmann (Gottingen) for critical advice.
Dr. U. Scheller (Jarpas) and Prof. Dr. H. Sturm (Hildesheim) kindly helped by providing valuable materials. Karin
Meyer’s technical support is gratefully acknowledged.

REFERENCES

ATTEMS. K. W.. 1926. — Myriapoda. In : W. KOKENTHAL & K. KRUMBACH, Handbuch der Zoologie , 4. Progoneata.

Chilopoda, Insecta , Berlin und Leipzig, W. de Gruyter & C° : 1-402.

Boudreaux, H. B.. 1987. — Arthropod phytogeny, with special reference to insects. Florida, Malabar, 320 pp.

DOHLE, W., 1964. — Die Embryonalentwicklung von Glomeris marginata (Villers) im Vergleich zur Entwicklung
anderer Diplopoden. Zool. Jb. Anat.. 81 : 241-310.

Dohle, W., 1980. — Sind die Myriapoden eine monophyletische Gruppe? Eine Diskussion der

Verwandtschaftsbeziehungen der Antennaten. Abh. naturwiss. Ver. Hamburg, 23 : 45-104.

ENGHOFF, H., 1993. — Haplopodous diplopods: a new type of millipede body construction discovered in cambalopsid
juveniles (Diplopoda, Spirostreplida). Acta zool. Stockholm, 74 : 257-261.

Hennig, W., 1969. — Die Stammesgeschichte der Insekten . Frankfurt a. M.. Kramer, 436 pp.

Hennig, W., 1981. — Insect Phytogeny. New York, John Wiley & Sons, 514 pp.

Hilken, G. & Kraus. O.. 1994. — Struktur und Homologie der Komponenten des Gnathochilarium der Chilognatha
(Tracheata, Diplopoda). Verh. naturwiss. Ver. Hamburg , (NF), 34 . 33-50.

Kraus, O., 1984. — Hoyers Gemisch statt Polyvenyl-Lactophenol. Mikrokosmos. 73 : 54-55.

Kraus, O., 1990. — On the so-called thoracic segments in Diplopoda. In : A. MINELLI, Proc. 7th intern. Congr.

Myriapodology. Leiden, Brill : 63-68. , „ .

Kraus, O. & Kraus, M., 1994. — Phylogenetic System of the Tracheata (Mandibulata) : on ‘Myriapoda - Insecta
relationships, phylogenetic age and primary ecological niches. Verh. naturwiss. Ver. Hamburg, (NF), 34 . 5-31.
Lautbrbach. K. E., 1972. — Uber die sogenannte Ganzbein-Mandibel der Tracheata. insbesondere der Myriapoda.

Zool. Anz ., 188 : 145-154. , .

Lautbrbach, K. E., 1980a. — Schlusselereignisse in der Evolution des Grundplans der Mandibulata (Arthropoda). Aon.

naturwiss. Ver. Hamburg, 23 . 105-161. , .

Lauterbach, K. E., 1980b. — Schlusselereignisse in der Evolution des Grundplans der Arachnata (Arthropoda). Aon.

naturwiss. Ver. Hamburg, 23 . 163-327. D

Manton. S. M., 1977. — The Arthropoda. Habits, functional morphology and evolution. Oxford, Clarendon Press,

Marcus^H., 1951. — Observaciones morfologicas en Dinjapyx marcusi. Folia Jniv. Cochabamba , 5 : 83-106.
Pocock, R. I., 1893. — On the classification of the tracheate Arthropoda. Zool. Anz., 16 : 271-275.

290

OTTO KRAUS & M ARGARETE KRAUS

Remy. P. A.. 1953. — Description de nouveaux types de Pauropodes: “ Millotauropus'”ci "Rabaudauropus" Mem. Inst,
scient. Madagascar, A8 : 25-41.

Tiegs, O. W., 1947. — The development and affinities of the Pauropoda, based on a study of Pauropus sylvaticus. Quart.
J. microscop. Sci., 88 : 165-336.

Verhoeff. K. W., 1910-1914. — Die Diplopoden Deutschlands, zusammenfassend bearbeitet. Leipzig, Winter, 482 pp.
WaloSSEK, D., 1993. — The Upper Cambrian Rehbachiella and the phylogeny of Brachiopoda and Crustacea. Fossils &
Strata, 32 : 1-202.

Source : MNHN. Paris

Morphology and Evolution of Circulatory Organs in

the Tracheata

Gunther PASS

Institut fur Zoologie, Universitat Wien, Althanstrasse 14, A- 1090 Wien, Austria

ABSTRACT

A comprehensive description of the anatomy of the circulatory organs is given from all subtaxa of myriapods,
apterygots and some lower Pterygota. In the Chilopoda, a complex vessel system exists which obviously represents a
plcsiomorphic condition in many respects. According to the most common teaching this system has been widely
reduced during the evolution of the Tracheata and in the Hexapoda only the tubular dorsal heart remained. However, in
some ancestral insects blood vessels exist in addition which have been partly overlooked so far, but demand special
interest from the evolutionary and phylogenetic points of view. One specific trait is a vessel ring caudal to the brain
encompassing the gut and connecting the dorsal heart with a short ventral vessel. This structure is found in the
Chilopoda, Diplura, Archaeogonata and Zygentoma, but has never been reported in the Pterygota. Special reference is
given to the hemolymph supply of longer body appendages, especially the antennae. In general, antennal vessels exist
which are considered to be homologous within the Tracheata. In all subtaxa of the myriapods and in the Diplura they
originate as arteries from the dorsal vessel. In all other investigated insects they are separated from the latter. At their
proximal ends they form ampulla-like enlargements with valved ostia, which communicate with the hemolymph sinus in
front of the brain. The connection of the antennal vessels to the dorsal heart in myriapods and Diplura is considered a
plesiomorphic state which was apparently lost early in insect phylogeny. Space constraints due to constructional
changes in the cephalic capsule are discussed as possible reasons for this loss. In the Archaeogonata and Zygentoma. the
ampullae arc not pulsatile, and their function is only to funnel hemolymph into the antennal vessels. In higher insects,
the ampullae are true forcing pumps as a result of associated muscles (“antenna-hearts”). In different species these
muscles diverge with respect to their attachcment sites and act either as dilators or as compressors of the ampullae. A
derivation of the antenna-heart muscles from pharynx dilators is strongly indicated.

RESUME

Morphologie et evolution des organes circulatoires chez les Tracheata.

Une description complete de 1’anatomie des organes circulatoires est donnee pour les sous-groupes de myriapodes,
d’apterygotes et de quelques pt£rygotes inferieurs. Chez les chilopodes, il exisle un reseau complexe de vaisseaux qui
represente un etat plesiomorphe. Selon les interpretations les plus communement en vigueur, ce systeme a ete largement
reduit au cours de revolution des antennates et, chez les hexapodes, seul le vaisseau cardiaque dorsal tubulaire s’est
maintenu. Cependant, chez certains insecies primitifs, des vaisseaux sanguins existent egalement, ph6nomene qui
demande 5 etre reconsider^ d’un point de vue evolutionniste et phylogenetique. Un caractere spdcifique reside dans le
vaisseau caudal annulaire du cerveau qui entoure le tube digestif et relie le cceur dorsal & un court vaisseau ventral. Cette
structure se retrouve chez les Chilopoda, Diplura, Archeogonata et Zygentoma,. mais n’a jamais ete mise en Evidence
chez les Pterygota. II est particulierement fait reference h Tapprovisionnement en hemolymphe necessaire aux longs
appendices du corps, notamment aux antennes. Les vaisseaux antennaires sont consid6r6s comme homologues chez tous
les antennates. Dans tous les sous-groupes de myriapodes et chez les diploures, ils apparaissent comme des arteres issues

Pass, G„ 1996. — Morphology and evolution of circulatory organs in the Tracheata. In: Geoffroy, J.-J..
Mauries, J.-P. & Nguyen Duy - Jacquemin, M„ (eds), Acta Myriapodologica. Mem. Mus. natn. Hist. not.. 169 : 291 -
292. Paris ISBN : 2-85653-502-X.

292

GUNTHER PASS

du vaisscau dorsal alors que chez tous les autres insectes ctudies ils se scparent de ce dernier. 11s ferment a leur extremife
proximale des elargissements en forme d'ampoule. equipes dc valvules qui commumquent au

sinus de rhemolymphe. La connexion des vaisseaux antennaires avec 1c cceur dorsal chez es myriapodes et les d ploures
est^onsiderde'eomme un caractere plesiomorphe qui a apparemmen. disparu au cours deta pjjjjfaje **«*£«. Uj
contraintes spatiales dues aux changements survenus dans la construction de la capsule cdphalique sont discutees en tant
Sue cauies possibles de eette perte. Chez les Archeogona.a e. les Zygentoma. les structures en ampoule ne sont pas
pulsatiles et leur fenction consistc uniquement t> permettre a rhemolymphe de c.rculer jusqu aux
Chez les Insectes superieurs. elles jouent le role de pompes. sortes de occurs antenna.res resultant de l ^ociation de
muscles Selon les especes, ces muscles se distinguent par leurs points d attache, agissant soil comme dilatateurs, son
comme compresseurs .Vhypothfese d'une evolution de ces muscles du cceur antennaire a partir de dilatateurs du pharynx
est fortement suggerSe.

Some Problems in the Systematics of the Order
Scolopendromorpha (Chilopoda)

Arkady A. SCHILEYKO

Zoological Museum of the Moscow State University. Herzen Street 6, 103009 Moscow K-9, Russia

ABSTRACT

The class Chilopoda ought to be divided into Noto- and Pleurostigmophora in relation to its phylogeny. It is hard to
speak about poly- vs. oligomerization as a general pathway in the evolution of the Chilopoda as a whole, chiefly due to
an extremely early isolation of the Scutigeromorpha and a polymerous development in the Geophilomorpha. The family
Cryptopsidae (Scolopendromorpha) is an unnatural composite taxon because of its polyphyly. This is easily to explain
in terms of the theory of biological progress associated in all branches of scolopendromorphs.with a transition to a
hypogean mode of life.

RESUME

Quelques questions de systematique dans l’ordre Scolopendromorpha (Chilopoda).

La classe Chilopoda devrait etre divisee. d'apres sa phylogenie, en Noto- el Pleurostigmophora. 11 est difficile de
considerer le contraste “polymetamerisation - oligometamerisation" comme une voie generate de revolution de
P ensemble Chilopoda, principalement a cause de I'isolement extremement precoce des Scutigeromorpha et du
developpement “p^yn'ctamerique" des Geophilomorpha. La famille Cryptopsidae (Scolopendromorpha) apparait comme
un taxon composite non-naturel h cause de sa polyphylie. Ceci est assez facile a expliquer en theorie par Lassociation,
dans toutes les lignees de scolopendromorphes, d'une evolution biologique et d'une transition vers un mode de vie
hypoge.

INTRODUCTION

Chilopod evolution is a subject of active debate (e.g. MANTON, 1952; PRUNESCU, 1965;
SHINOHARA, 1970; DOHLE, 1988; SHEAR & BONAMO, 1988). Basically, some studies adhere
to oligomerization (= reduction in the number of body segments) as the major evolutionary trend
in the Chilopoda, while others document that in terms of polymerization. Systematically, the
class has been divided either into Noto- and Pleurostigmophora or Ana- and Epimorpha,
dependent on the pattern of allocation of the stigmata and the traits of postembryonic
development, respectively. (By the way, is such a character as the type of development (ana- vs.
epimorphosis) reliable taxonomically for dividing taxa of so high level?).

However, in addition to new evidence accumulated in the recent years, particularly the
discovery of a new extinct chilopod order (SHEAR & BONAMO, 1988), and a new cladistic
analysis (DOHLE, 1988), much remains to be clarified, either based on recent results or older
literature data.

SCHILEYKO, A. A., 1996. — Some problems in the systematics of the Order Scolopendromorpha (Chilopoda). In:
Geoferoy. J.-J., MauriBs, J.-P. & Nguyen Duy - Jacquemin, M.. (eds), Acta Myriapodologica. Mem. Mus. nain. Hist,
nat., 169 : 293-297. Paris ISBN : 2-85653-502-X.

294

ARKADY A. SCHILEYKO

The main impetus for presenting this preliminary paper lies in the deep interest we can
find, among Chilopoda, in the evolution and systematics of the order Scolopendromorpha (e.g.
SCHILEYKO, 1992; ZALESSKAJA & SCHILEYKO, 1992).

SYSTEM OF THE SCOLOPENDROMORPHA

At present, the system of ATTEMS (1930) of the centipede order Scolopendromorpha is
generally accepted, with such characters as the presence or absence of eyes serving as its basis.
The order is divided into two families: Scolopendridae (16 genera with eyes) and Cryptopsidae
(12 blind genera) (Fig. 1). However, about five years ago, when working with a collection of
Scolopocryptops ferrugineus (Brolemann, 1919), from Cuba, I found great similarity between
ScolopocryptopsPoral, 1876 and numerous Scolopendridae. In addition, I noted many
differences between Scolopocryptops and Cryptops Leach, 1815 (Fig.l). This provoked the
conclusion that Cryptopsidae is possibly a polyphyletic group. In other words, the main reason
for revising the system of this order is the apparent polyphyly of the family Cryptopsidae.

To my mind, the Attemsian system seems to reflect the order's eco-morphology rather than
phylogeny and fails to explain the allocation within a monophyletic family Cryptopsidae of such
quite different representatives as the genera Scolopocryptops, Dinocryptops Newport, 1844,
Plutonium Cavanna, 1881 or Cryptops (Fig. 1), regardless of the pathway centipede evolution
we accept (oligo- vs. polymerization).

1 have therefore analyzed all available material from the Zoological Museums of Moscow
and St-Petersburg. This amounted to about two thousand specimens from the following genera
(Fig. 1): Theatops Newport, 1845, Tonkinodentus Schileyko, 1992, Cryptops, Paracryptops
Pocock, 1891, Scolopocryptops, Dinocryptops, Newport ia (all Cryptopsidae), and Scolopendra
L., 1758, Cormocephalus Newport, 1844, Asanada Meinert, 1886, Otostigmus Porat, 1876,
Alipes Imhoff, 1845, Ethmostigmus Newport, 1845, Rhysida Newport, 1845 (all
Scolopendridae).

I have tried to evaluate the maximal number of characters, the main of which are the
following (Table 1):

(1) number of spiracles; (2) number of body segments; (3) structure of spiracles;
(4) presence of eyes; (5) presence of tooth plates of maxillipede coxostemite; (6) presence of
coxopleural pores; (7) presence of coxopleural process; (8) structure and ornament of last legs.

Plesiomorphy is coded by 0, apomorphy by 1 and serial tranformations by 2 to 4
(Table 1).

Table 1. — List of the characters with their evaluation as apomorphy or plesiomorphy.

Characters

Plesiomor phic

Apomorphic

1 . Number of spiracle pairs

19 (0)

11 (1), 10 (2),9 (3)

2. Number of body segments

23 (0)

21 (1)

3. Structure of spiracles

without flap (0)

with flap (1)

4. Eyes

presence (0)

absence (1)

5. Tooth plates

presence (0)

absence (1)

6. Coxopleural pores

presence (0)

absence (1)

7. Coxopleural process

presence (0)

absence ( 1 )

8. Structure of last legs

pincer-shaped ,
without spines (0)

normal-shaped,
with spines (1),
leaf-shaped (2),
with “saw” (3),
many-segmented tarsi (4)

However, I have not attempted a cladogram, because I have not seen representatives of all
genera. A cladogram, in this case, would be deficient. Besides, to my mind, the cladistic
methods are sometimes not objective, because the choice of characters, the evaluation of degree

Source :

SYSTEMATICS OF THE ORDER SCOLOPENDROMORPHA

295

of their expression and of their taxonomical importance is rather subjective (same as in the
“classic” methods). Because of all this, certainly the set of characters to be analyzed must be
extended.

SCOLOPENDROMORPHA

Scolopendrinae

Otostigminae

Cryptopinae

Theatopsinae

Scolopocryptopinae

Scolopendrini

Otostigmini

Cryptops

Theatops

Scolopocryptops

Scolopendra

Otostigmus

Paracryptops

Plutonium

Newport ia

Trachycormocephalus

Digitipes

Anethops

Tidops

Cormocephalus

Alipes

Mimops

Otocryptops

Arthrorabdus

Ethmostigmus

Kethops

Campilostigmus

Rhysida

Kartops

Rhoda

Allurops

Scolopendropsis

Arrhabdotini

Asanadini

Asanada

Pseudocryptops

Arrahabdotus

—

Fig. 1. — System of the Scolopendromorpha after Attems (1930).

A character matrix has been compiled (Table 2). Plutonium is a single genus, which I have
never personally seen, but I included it in the matrix, because of the great importance of this
form for phylogeny of the whole Scolopendromorpha. I analyzed the cardinal character of the
system of ATTEMS. To my mind, this character is highly adaptive and not reliable
taxonomically. There are numerous examples of eye losses in connection with the transition to a
hypogeal mode of life (edaphic and cavemicolous), e.g. in some Lithobiidae centipedes, Atyidae
shrimps, Trigonochlamydidae slugs, Characinidae fishes, etc.

Closely related forms with eyes are always present.

The system of ATTEMS is based on a single character. In this case, if one of that two
families is polyphyletic, all the system is not reliable. In my opinion, this matrix demonstrates
that Cryptopsidae, sensu ATTEMS, is not monophyletic (Table 2).

Table 2. — The matrix of the characters.

Genus

Characters

Cryptopsidae

Plutonium

Theatops

Tonkinodentus

Cryptops

Paracryptops

Scolopocryptops

Otocryptops

Newportia

Scolopendridae

Scolopendra

Cormocephalus

Asanada

Otostigmus

Alipes

Ethmostigmus

Rhysida

296

ARKADY A. SCHILEYKO

In addition, there is a very interesting question about the monotypical genus Plutonium,
which has 21 body segments with 19 pairs of spiracles, in other words the spiracles are
disposed on all body segments, except for the first and the last one (as in Geophilomorpha). As I
have already written, the Attemsian system fails to explain the allocation within the family
Cryptopsidae of some very morphologically different genera, and at first the allocation of
Plutonium zwierlainii Cavanna, 1881, regardless of the apo- or plesiomorph type of its
homonomity. Some of my ideas about the last problem are as follows.

The evolution of most groups of polymerous invertebrates, which left the soil environment
for surface habitats, is known to have undergone oligomerization (Arachnida, Insecta). In my
opinion, it is difficult to speak about poly- vs. oligomerization as a general pathway in the
evolution of the Chilopoda as a whole, primarily due to an extremely early isolation of the
Scutigeromoipha and a polymerous development in the Geophilomorpha. However, it is known
that the reduction of spiracles is associated with the development of anisotergy at first, and this
takes place in all orders of the Chilopoda which have moved to open habitats.

By the way, a second possible reason for this reduction is apparently connected with a
reduced transpiration rate through these structures devoid of epicuticule (KAUFMAN, 1959).
Water economy could have become more important during chilopod penetration into arid habitats
and regions. The dorso-medial spiracles of Scutigeromorpha are, possibly, the top of evolution
of this structure.

But representatives of Geophilomorpha have moved to the hypogean mode of life and have
a homonomous and polymerous body without well expressed anisosegmentation. Their
polymerization can be easily explained in terms of adaptation to active wormlike movements in a
more dense environment. But if the homonomity in Geophilomorpha is an apomorphy, I cannot
clearly imagine what their evolutionary pathway was, assuming so because their ancestor had an
anisosegmentation. But as an alternative, the Geophilomorpha could have had a homonomous
ancestor, and they retained homonomity.

Fig. 2. — The phylogenetic tree of the Chilopoda after Shear & Bonamo (1988).

Source

SYSTEMATICS OF THE ORDER SCOLOPENDROMORPHA

297

Moreover, the Scolopendromorpha and the Geophilomorpha are closely related groups
(PRUNESCU, 1965; DOHLE, 1988; SHEAR & BONAMO, 1988) (Fig. 2) and most probably have
a common ancestor. In this case, if body homonomity is a plesiomorphy in the
Scolopendromorpha, their ancestor would have had a homonomous body. However, amsotergy
would have been absolutely mandatory for groups of centipedes which colonized the soil
surface. Anisosegmenlation is gradually increasing in the following row: Scolopendromorpha -
Craterostigmomorpha - Lithobiomorpha - Scutigeromorpha. Apparently, this fact is due to an
increased velocity and manoeuvrability while moving and improving the transpiratory system.
All these are especially important for predators. I note that this succession is not an evolutionary
one, but it is only an attempt to a morpho-functional analysis of anisosegmentation.

CONCLUSIONS

1. So the family Cryptopidae ( sensu ATTEMS) is probably an unnatural composite taxon,
because of its polyphyly. This is easily explicable in terms of the theory of biological progress,
associated in all groups of scolopendromorphs with a transition to a hypogean mode of life.

2. If the homonomity is plesiomorphic in the Scolopendromorpha, Plutonium is a form
most closely related to their common ancestor. Hence, perhaps Plutonium deserves not only a
family of its own (SCHILEYKO, 1992), but even a suprafamily status, as an absolutely different
group.

ACKNOWLEDGMENTS

I am most grateful to the following persons who offered many useful suggestions and improvements to this
paper: Dr. A. A. Schileyko, Sr, Dr. S. 1. Golovatch, Dr M. V. Heptner (all Moscow), and Dr. Y. I. Starobogatov (St.
Petersburg). Valuable discussions have also been rendered by Dr. H. Enghoff (Copenhagen), Dr J. G. E. Lewis (Taunton),
Prof. W. Dohle (Berlin) and Prof C. Prunescu (Bucarest). In addition, I would like extend my deep appreciation to
Organising Committee of the 9th Inernational Congress of Myriapodology whose support has enabled me to participate
in the congress. A part of this work has been sponsored by the Soros Foundation.

REFERENCES

ATTEMS, G., 1930. — Myriapoda. 2. Scolopendromorpha. In : Das Tierreich. Berlin, Walter de Gruyter & Co, 307 pp.
DOHLE, W., 1988. — Myriapoda and the Ancestry of Insects. Manchester, Impact Print Services Ltd., 28 pp.

Kaufman, Z. S., 1959. — Morphology of spiracles of Geophilus proximus C. L. Koch (Chilopoda). Dokl. AN USSR.
129 : 698-701. (in Russian).

M anton, S. M., 1952. — The evolution of Arthropodan locomotory mechanisms. Part 2. General introduction to the
locomotory mechanisms of the Arthropoda. J. Linn. Soc. (Zool.). 42 : 93-167.

PRUNESCU. C. C., 1965. — Contribution a l'etude de 1'evolution des Chilopodes. Rev. Roum. Biol. (Zool.). 10 : 89-102.
Schileyko, A. A. Jr., 1992. — Scolopenders of Viet-Nam and some aspects of the system of Scolopendromorpha
(Chilopoda Epimorpha). Part 1. Arthropoda Selecta , 1 : 5-19.

Shear, W. A. & Bonamo, P. M.. 1988. — Devonobiomorpha, a new order of centipedes (Chilopoda) from the middle
Devonian of Gilboa, New York State, USA, and the phylogcny of Centiped Orders. Amer. Mus. Nov., 2977 ; 1-30.
Shinohara, K., 1970. — On the phylogeny of Chilopoda. Proc. Japan. Soc. Syst. Zool.. 65 : 35-42.

ZALESSKAJA, N. T. & Schileyko, A. A. Jr., 1992. — The scolopendromorph centipedes of USSR. Moscow , "Nauka"
Publ., 110 pp. (in Russian).

Source : MNHN, Paris

Plesiomorphic and Apomorphic Characters States in

the Class Chilopoda

Carol Constantin PRUNESCU

Institute of Biology, 296 Spl. Independentei, RO-79651 Bucarest, Romania

ABSTRACT

The plesiomorphic and apomorphic nature of characters used for a cladistic analysis in the class Chilopoda is taken
into account. The plesiomorphic or apomorphic status of the following features are proposed to be discussed here:
spiracles and types of respiratory systems, coxal/anal glands (organs), spines of the first article of the female gonopod,
male gonopods, testis, genital tract, supernumerary Malpighian tubules.

RESUME

Etats plesiomorphique et apomorphique des caracteres dans la classe Chilopoda.

La nature plesiomorphique ou apomorphique de chaque caracterc utilise pour une analyse cladistique des chilopodes est
prise en compte. On propose notamment de discuter ici le statut plesiomorphique ou apomorphique des organes suivants :
spiracles (stigma) et types de systeme respiratoire, glandes coxales/anales, gonopodes des femelles, gonopodes des
males, testicules, tractus genital, tubes de Malpighi supplementaires.

INTRODUCTION

In 1965 a preliminary phylogenetic tree of Chilopoda was published (PRUNESCU, 1965)
already made according to cladistic principles. For the reconstruction of the morphologic
characteristics of the primitive chilopods, the primitive morphologic features of representatives
of the orders Scutigeromorpha and Lithobiomorpha were selected. This was followed by a
synthesis of the research on the anatomy and evolution of the genital system in Chilopoda
(PRUNESCU, 1969a), as well as a discussion regarding the place of some atypical chilopods
regarding their systematics and evolution (PRUNESCU, 1969b). In 1985, W. DOHLE published a
cladistic analysis of the main chilopod groups and suggested a phylogenetic tree resembling that
published earlier (PRUNESCU, 1965). A series of morphological features are considered
plesiomorphic or apomorphic by DOHLE (1985), not as a result of a critical scientific analysis,
but to underline the main resemblances of chilopods ancestors with recent representatives of the
subclass Notostigmophora. In in a paper describing the fossil order Devonobiomorpha. SHEAR
& BONAMO (1988) dealt with cladistic analysis of several morphologic features treated by
DOHLE (1985). Taking into account that some features may have been wrong appreciated by
both authors, we considered necessary to reexamine them, as a contribution to the cladistic

PRUNESCU, C. C., 1996. — Plesiomorphic and apomorphic characters states in the class Chilopoda. In:
Geoffroy, J.-J., MAURlfcS. J.-P. & NGUYEN Duy - Jacquemin, M.. (eds), Acta Myriapodologica. Mem. Mus. natn . Hist,
nat., 169 : 299-306. Paris ISBN : 2-85653-502-X.

300

CAROL CONSTANTIN PRUNESCU

analysis of Chilopoda. A series of features, such as male gonopods, seminal vesicles,
supplementary Malpighian tubules were analysed here, for the first time.

CHILOPOD ENVIRONMENT

In our opinion, chilopods originate within the aquatic arthropods. The same opinion is
shared by other authors (KRAUS & KRAUS, 1994). The primitive chilopod environment could
have been wet or very wet. Most recent chilopods live within a wet environment created by
forest soil, deep cracks in rock and by caves. This option satisfies the vital needs of
Pleurostigmophora chilopods and offers the natural framework in which these Chilopoda
evolved and diversified from the Pleurostigmophora with 15 leg-bearing segments, unequal
tergal shields and anamorphic development, to elongated Pleurostigmophora with homonomous
secondary segmentation and epimorphic development. We consider the above mentioned wet
environment as plesiomorphic for Chilopoda.

The life environment of Notostigmophora which live and hunt in the open air, on rocky
walls and open beaches is an apomorphic one, conquered by an ancestral branch ol actual
Notostigmophora. detached directly from the primitive Pleurostigmophora and having the rank
of sister-group with the line from which the present Pleurostigmophora derived. The
Notostigmophora preserved most of the plesiomorphic features of the primitive chilopods, but
also adapted their metabolism and some of their organs to this different environment.

PAIRED LATERAL SPIRACLES

These structures serve a tracheal system through which gas exchange occurs at the level of
the cells of the whole organism. Pleurostigmophora kept a plesiomorphic circulatory system,
because their way of life did not imposed a decrease of the blood circulation.

The same type of plesiomorphic circulatory system was also maintained in
Notostigmophora. Since S. HAASE (1885), the presence of mid-dorsal spiracle has been
considered to have originated in displacement and subsequent median union of the two pleural
spiracles. However, tracheal lungs linked by this single median spiracle are paired and are in
accordance with the bilateral chilopod organization.

Radical modification of the breathing system in Notostigmophora chilopods results from
the gases exchange in “tracheal lungs” between the haemolymph which contains hemocyanin and
the oxygenated air, which goes through the tracheae, closed like in a glove finger. The changing
of the breathing system in the ancestors of the recent Notostigmophora can be related to the life
environment of these chilopods. Scutigera hunt almost all their life in the open air, namely in a
drier environment than the humid air of leaves and humus. In our opinion, the breathing system
in Notostigmophora allowed the inner humidity to be maintained through decreasing water
vapour loss at the gas exchange level, in the present apomorphic environment (Fig. 1). This is
why we consider that the bilateral spiracles as well as the tracheal breathing system in
Pleurostigmophora has plainly plesiomorphic features. Therefore these features can be
considered characteristic for the ancestor of the recent chilopods.

The presence of spiracles on leg-bearing segments with large tergites, their absence on leg¬
bearing segments with small tergites, and the absence of alternation which occur in the
successive large tergites on segments VII and VIII, where only the leg-bearing segment VIII has
spiracles, argues for the existence of these features in the ancestor of recent Pleurostigmophora.
The same strict distribution of the mid-dorsal spiracles is found again in Notostigmophora: the
spiracles are present on large tergites of leg-bearing segments inclusive on the unique tergite
covering the leg-bearing segments VII and VIII. This homology suggests the presenceof
alternation and the lack of alternation as well in the ancestor of Chilopoda (PRUNESCU, 1965).

The problem of lateral spiracles for tracheal breathing and mid-dorsal spiracles for tracheal-
lungs breathing must be treated separately from the problem of spiracle distribution on leg-

PLESIOMOR PH Y AND APOMORPHY IN THE CHILOPODA

301

bearing segments. According to the above argumentation, the tracheal breathing through pleural
spiracle is a plesiomorphic feature and the tracheal-lung breathing through mid-dorsal spiracles
an apomorphic one. The distribution of spiracles on leg-bearing segments with large tergites in
Scutigeromorpha, Lithobiomorpha, Craterostigmomorpha and Scolopendromorpha represents a
plesiomorphic feature and the distribution of spiracles on every leg-bearing segments in
Geophilomorpha or in the genus Plutonium (Scolopendromorpha) is an apomorphic one
(PRUNESCU, 1965).

COXAL - ANAL ORGANS

According to ROSENBERG (1982, 1983, 1989), coxal and anal glands are respectively
specialized organs for the uptake and release of water vapour and ions from and to the
environment. As mentioned above, the plesiomorphic environment of chilopods was a wet one,
similar to that now existing under leaves, in humus, etc. These organs are only useful in a wet
environment , in order to maintain the water and ion balance of the organism. If the pheromones
are eliminated together with the water vapours (LITTLEWOOD, 1983), the plesiomorphic state of
these glands is not changed. The lack of these organs in most of the species of Oryidae
(ATTEMS, 1929) adapted to dry environments supports the idea of the adaptational loss of these
organs in such species. The life style of Scutigera , which hunts on stone surfaces in the open
air, would not be possible, if the coxal-anal organ were maintained (Fig. 2). Therefore, we
consider the coxal-anal gland as a plesiomorphic feature and its absence in Scutigeromorpha as
an apomorphic feature. SHEAR & BONAMO (1988) seemed to agree this interpretation but finally
consider the absence of such organs as a plesiomorphic feature and their presence as an
apomorphic one.

Tracheal breathing

Chilopod

haemolymph

co2

wet environment

— = ► water vapour

- ► o2

tracheas

Chilopod

haemolymph
hemocyanin + C02
hemocyanin + 02

Tracheal lung

dry environment

indirect breathing

Chilopod

water

ions

wet environment

► water vapour

► ions

anal or coxal glands

Chilopod

water

ions

dry environment

integument

no anal nor coxal glands

Fig. 1. — Correlation of respiration type with life
environment in Chilopoda.

Fig. 2. — Presence and absence of coxal-anal glands
in connexion with the life environment of
Chilopoda.

In Lithobiomorpha, the coxal glands are located on the last 4 pairs of coxae. We consider
this distribution plesiomorphic.

302

CAROL CONSTANTIN PRUNESCU

In Scolopendromorpha and Geophilomorpha, coxal glands are distributed on the last pair
of coxae, sometimes on the margin of the respective sternite. This reduction of the coxal glands
can be considered an apomoiphic feature of the first degree.

The older name of “ano-genital capsule” (DOHLE, 1990) seems a misleading name for the
structure of the involved organ. Its presence in Craterostigmomorpha only is apomorphic
(SHEAR & BON AMO, 1988). In fact, in its two halves, this capsule contains several glands
homologous to the coxal glands of other chilopods (unpubl. observations). These glands,
leaving the coxae, located in an original organ. The ability of this organ to close firmly or open
widely may be linked with the need to con troll water vapour exchange with the environment. A
proper name would be the “capsule of coxal-anal glands”. This transformation can be considered
as an apomoiphic feature of the second degree. The lack of coxal glands in Scutigeromoipha can
be considered as an apomorphic feature of the third degree.

FEMALE GONOPODS

These organs are highly-modified ambulatory appendages (Fig. 3). As the articles of the
ambulatory appendages in Scutigeromoipha and Lithobiomorpha have a large spine (macrosetae)
at their distal ends, we consider that the female gonopod, which is characteristic to
Lithobiomorpha and has macrosetae, is plesiomorphic.

FEMALE GONOPOD

SCUTIGEROMORPHA

FIRST COXOSTERNITE ♦

GENITAL 2 -SEGMENTED GONOPOD

SEGMENT WITHOUT MACROSETAE

LITHOBIOMORPHA

(Anopsiidac-Henicopidae-

Lithobiidae)

/ J

GEOPHILOMORPHA

3 - SEGMENTED GONOPOD
WITH MACROSETAE

1/2 - SEGMENTED GONOPOD
IRUDIMENTARY)

Fig. 3. — Female gonopods. (Drawings reproduced from Attems. 1926). A: anal segment; AK: anal valves; Ec: coxite of
the last legs; Ev; sternite of the last leg-bearing segment; gon, Gon: gonopods; gon tel: telopodite of the
gonopods; gone: coxite of the gonopods; Gp: pleurite of the genital segment; gon tel: telopodite of the
gonopods; Gv: sternite of the genital segment; Pg. Pv: sternite of the pregenital segment; S 16 : sternite of the
16th segment; Ta: tergite of the anal segment; V|6: sternite of the 16th leg-bearing segment; vp: sternite of the
penultimate leg-bearing segment.

PLES IOMORPH Y AND APOMORPHY IN THE CH1LOPODA

303

The two female gonopods of Lithobiomorpha are firmly separated from one another and
formed of three distinct segments, while the female gonopods of Scutigeromorpha comprise two
articles, the first one being partly joined (Fig. 3). The female gonopod is a plesiomorphic feature
in Lithobiomorpha, an apomorphic one in Scutigeromorpha. Rudimentary female gonopods in
Geophilomorpha show a greater degree of apomorphism, of the second degree. The lack of
female gonopods in Craterostigmomorpha and Scolopendromorpha is equivalent to an
apomorphic feature of the third degree.

MALE GONOPODS

Although the authors of the previous cladistic analysis did not deal with this feature, we
think that a review of it is of phylogenetic interest.The most complete male gonopod exists in the
representatives of the Henicopidae (Lithobiomorpha). Within this group, the male gonopod
consists of 4 distinct segments. In comparison, the male gonopods of Lithobiidae are
rudimentary. The male gonopods of Geophilomorpha are formed of two well articulated
segments. The male gonopods in Scutigeromorpha are rudimentary but in two pairs (Fig. 4).

MALE GONOPOD

SCUTIGEROMORPHA

FIRST 1 • SEGMENTED

GENITAL GONOPOD

SEGMENT (RUDIMENTARY)

LITHOBIOMORPHA

Henicopidae

4 SEGMENTED
GONOPOD

Lithobiidae
1/2 ■ SEGMENTED
GONOPOD
(RUDIMENTARY)

2 - SEGMENTED
GONOPOD

SECOND 1 - SEGMENTED

GENITAL GONOPOD

SEGMENT (RUDIMENTARY)

Fig. 4. — Male gonopods. (Drawings reproduced from Attems, 1926). A. a: anal segment: AK: anal valves; Ec: coxite of
the last legs; Ev; sternite of the last leg-bearing segment; gi pregenital segment of the gonopodes; g 2:
rudimentary gonopods of the genital segment; gon. Gon: gonopods; Gp: pleurite of the genital segment; Gv:
sternite of the genital segment; p: penis; Pg, Pv: sternite of the pregcnital segment; Si6: sternite of the 16th
segment; Ta: tergite of the anal segment; Tg: tergite of the genital region; Vi6: sternite of the 16th leg-bearing
segment; vp: sternite of the penultimate leg-bearing segment; Z: sternite of the genital segment.

Taking into account their number, they represent a clear plesiomorphic feature of
Scutigeromorpha and are apomorphic for all the other chilopods. By the reduction in size and

304

CAROL CONSTANTIN PRUNESCU

number of articles, these gonopods are apomorphic. We therefore suggest the separation of
‘"gonopod pairs number” feature from the “gonopod articles number” feature. Thus, any future
cladistic analysis will be able to use both features.

The quantification of the “male gonopod articles number” feature could be made as
follows:

- Henicopidae: 4 segments = plesiomorphic feature.

- Lithobiidae: rudimentary gonopods = apomorphic feature of the second degree.

- Scutigeromorpha: rudimentary gonopods = apomorpic feature of the second degree.

- Geophilomorpha: biarticulated gonopods = apomorphic feature of the first degree.

- Craterostigmomorpha and Scolopendromorpha: gonopods absent = apomorphic feature
of the third degree.

TESTES AND TESTICULAR SYSTEM

In Scutigeromorpha, the male genital system consists of two testes, each formed of a
macrotestis and a microtestis (PRUNESCU. 1969c). This structure can also be observed during
the larval development (PRUNESCU, 1992b) and is clearly plesiomorphic (Fig. 5).

SCUTIGEROMORPHA LITHOBIOMORPHA CRATEROSTIGMOMORPHA SCOLOPENDROMORPHA GEOPHILOMORPHA

Fig. 5. — Ontogeny and phylogeny of the male genital system in Chilopoda. 1: macrotestis; 2: microtestis; 3; seminal
vesicles; 4: undifferenciated rudimentary testis.

In adults of the tribe Anopsobiini (Lithobiomorpha), one testis is undifferentiated and
exclusively populated with spermatogonia. The other functional one, consists of macrotestis and
microtestis, as in Scutigeromorpha (PRUNESCU & JOHNS, 1969; PRUNESCU, 1992a). This
testicular system is apomorphic in the first degree. Esastigmatobius, of the Henicopidae
(PRUNESCU, MESIBOV & SHINOHARA, this volume) and numerous genera of Lithobiidae
(PRUNESCU, 1964) have the testicular system formed of a single testis.

Source : MNHN, Paris

PLESIOMORPHY AND APOMORPHY INTHECHILOPODA

305

In Lithobius forficatus, during larval development, the unique testis results from the
joining of two male gonads (BlEGEL, 1922; ZERBIB, 1966). Hence, the single testis in
Lithobiidae and perhaps in Henicopidae s. str., is an apomorphic feature of the second degree.

During the larval development of Scolopendromorpha, the two embryonic testis merge to
form an unpaired median organ which, by subsequent lateral burgeonings, forms a large
deferens duct, to which are linked numerous, pseudometameric, testicular vesicles (HEYMONS,
1901). This is an apomorphic testicular system of the third degree.

Incomplete data for microscopic anatomy show that Craterostigmus tasmanianus testicular
system structure is similar to that of Scolopendromorpha (PRUNESCU, MES1BOV &
Shinohara, this volume).

The anatomic data of the testicular system in Geophilomorpha show the presence of two
lateral testicular vesicles, linked by a central deferents duct. Thus, Craterostigmomorpha,
Scolopendromorpha and Geophilomorpha have an apomorphic testicular system of the third
degree. According to the present state of knowledge, we cannot differentiate a distinct
apomorphic degree between the testicular system in Geophilomorpha with that of
Scolopendromorpha.

MALE GENITAL TRACT

The male genital tract is constituted by the genital organs between the testes and the genital
atrium. This anatomical structure follows a clear evolution simplification.

In Scutigeromorpha several organs play an important role in the deposition, maturation and
preservation of spermatozoa. We consider that this is a plesiomorphic feature. Of this very
complex and histologically varied anatomic system, only two elongated tubes are retained in
Lithobiomorpha. They are named seminal vesicles and have the same role as the similar organs
of Scutigeromorpha. This is an apomorphic situation of the first degree.

In Craterostigmomorpha, Scolopendromorpha and Geophilomorpha orders, the seminal
vesicles are absent, their function being taken over by the posterior half of the very expended
deferens duct. In this duct, the spermatozoa are deposited and the spermatophores formed
(JANGI, 1956). This can be considered as an apomorphic feature of the second degree.

SUPERNUMERARY MALPIGHIAN TUBULES

Only two Malpighian tubules were known to be present in all chilopod groups (LEWIS,
1981).

In Scutigera coleoptrata we found a supplementary pair of Malpighian tubules, which have
dorso- ventral insertion, at the level of the junction of the mid-gut with the hind-gut.

In Craterostigmus tasmanianus we found only one supplementary Malpighian tubule,
which has a medio-dorsal insertion at the same area of the intestine, and which is directed
towards the posterior region of the body (see PRUNESCU & PRUNESCU, this volume).

The presence of two supplementary Malpighian tubules in Scutigera coleoptrata suggests
they are plesiomorphic. As a rule, in Chilopoda, the evolution presents a tendancy to simplify
the features of different systems or organs. So, the absence of supernumerary Malpighian
tubules in Lithobiomorpha is an apomorphic feature of the second degree. The disappearance of
the ventral supernumerary tubule in Craterostigmus tasmanianus represents an apomorphic
feature of the third degree, while the presence of the dorsal supernumerary tubule represents a
plesiomorphic one. The absence of any supernumerary Malpighian tubule in
Scolopendromorpha and Geophilomorpha represents an apomorphic feature of the second
degree, distinct from the apomorphic feature of the second degree in Lithobiomorpha, which
was realized by its own evolutive line.

306

CAROL CONSTANTIN PRUNESCU

REFERENCES

ATTEMS, C. , 1926. — Myriopoda. In : W. KOKENTHAL & T. KRUMBACH, Handbuch der Zoologie, 4, Progoneata,
Chilopoda, Insecta, Berlin & Leipzig, W. de Gruyter & C° : 1-402.

Attems, C., 1929. — Myriapoda 1. Geophilomorpha. In : F. E. Schulze & W. KOkenthal, Das Tierreich , 52. Berlin &
Leipzig, W. De Gruyter & C° : 1-388.

Biegel, J., 1922. — Beitrage zur Morphologie und Entwicklung Geschichte des Herzens bei Lithobius forficatus (L.).
Rev. Suisse Zool., 29 : 444-480.

DOHLE, W., 1985. — Phylogenetic pathways in the Chilopoda. Bijdr. Dierk., 55 : 55-66.

Dohle, W., 1990. — Some observations on morphology and affinities of Craterostigmus tasmanianus (Chilopoda). In :
A. MlNELLl, Proc. 7th Intern. Congr. Myriapodology , Leiden. E. J. Brill : 69-79.

Haase, E., 1885. — Zur Morphologie der Chilopoden. Zool. Anz., 8, n° 216 : 693-696.

HEYMONS. R., 1901. — Entwicklungsgeschichte der Scolopender. Biblioth. Zoologica Chun., 33 : 82-196.

Jangi, B. S., 1956. — The reproductive system in the male of the centipede Scolopendra morsilans. Linn. Proc. Zool.
Soc. Lond., 127 : 145-159.

Kraus, O. & Kraus, M., 1994. — Phylogenetic System of the Tracheata (Mandibulata): on “Myriapoda” - Insecta
relationships, phylogenetic age and primary ecological niches. Verb, nalurwiss. Ver. Hamburg , (NF), 34 : 5-31.

Lewis, J. G. E., 1981. — The biology of Centipedes. Cambridge, Cambridge Univ. Press, 475 pp.

Littlewood, P. M. H., 1983. — Fine structure and function of the coxal glands of lithobiomorph centipedes: Lithobius
forficatus and L. crassipes (Chilopoda, Lithobiidae). J. Morphol. , 177 : 157-159.

PRUNESCU, C. C., 1964. — Anatomie microscopique du systeme genital male des Lithobiid6s. Rev. Roum. Biol. (Zool.),
9 : 101-104.

PRUNESCU, C. C., 1965. — Contribution a 1'etude de 1'evolution des Chilopodes. Rev. Roum. Biol. (Zool.), 10 : 89-
102.

PRUNESCU, C. C., 1969a. — Considerations sur 1'evolution du systeme genital des Chilopodes. Bull. Mus. natl. Hist,
nat., Paris, 41, suppl. 2 : 108-111.

PRUNESCU, C. C., 1969b. — Quelle est la place occupee par Cermatobius , Craterostigmus et Plutonium dans la
phylogenie des Chilopodes? Bull. Mus. natl. Hist. nat. Paris, 41, suppl. 2 : 112-115.

PRUNESCU, C. C., 1969c. — Le systeme genital male de S. coleoptrata, (Notostigmophora, Chilopoda). Rev. Roum.
Biol. (Zool.), 14 : 185-190.

PRUNESCU, C. C., 1992a. — The genital system in Dichelobius (Anopsobiidae. Lithobiomorpha, Chilopoda), Ber. nat.-
med Verein Innsbruck, suppl. 10 : 87-91.

PRUNESCU, C. C., 1992b. — The beginning of double spermatogenesis in Scutigera coleoptrata. Ber. nat-med Verein
Innsbruck. Suppl. 10 : 93-97.

PRUNESCU, C. C. & JOHNS, M., 1969. — An embryonic gonad in adult males of Anopsobius neozelandicus Silv.
(Chilopoda). Rev. Roum. Biol. (Zool.), 14 : 407-409.

Rosenberg, J., 1982. — Coxal organs in Geophilomorpha (Chilopoda), organization and fine structure of the
transporting epithelium. Zoomorphology, 100 : 107-120.

Rosenberg, J., 1983a. — Coxal organs of L. forficatus (Myriapoda, Chilopoda). Fine structural investigation with
special reference to the transport epithelium. Cell Tissue Res., 230 : 421-430.

Rosenberg, J., 1983b. — Coxal organs in Scolopendromorpha (Chilopoda). Topography, organization, fine structure
and signification in Centipedes. Zool. Jb. Anal., 110 : 383-393.

Rosenberg, J., 1989. — A key to the middle European Centipedes (Geophilomorpha) based on the coxal pores. Acts
Biol. Benrodis , 1988, 1 : 133-141.

Shear, W. A. & Bonamo P., 1988. — Devonobiomorpha, a new order of Centipeds (Chilopoda) from the middle
Devonian of Gilboa, New York State, USA, and phylogeny of centiped orders. Am. Museum Novitates, 2927 : 1-30.

Zerbib, C. W., 1966. — Etude descriptive et experimental de la differentiation de l'appareil genital du Myriapode
Chilopode Lithobius forficatus (L.). Bull. Soc. zool. France, 91 : 203-216.

Source : MNHN. Paris

A Preliminary Study on Phylogeny and Biogeography
of the Family Paracortinidae (Myriapoda: Callipodida):

a Cladistic Analysis

Daqing WANG

Department of Invertebrates, Institute of Zoology, Academia Sinica, Beijing 100080, P.R. China

ABSTRACT

The phylogeny and biogeography of a millipede family (Paracortinidae. fam. nov.) are preliminary examined by using
a cladistic analysis. One of the 35 most parsimonious trees (cladograms), which also has the lowest (best) F value, is
congruent with the scheme of evolution proposed by Wang & Zhang (1993) from systematical analysis. Yunnan
millipede R . stimulus is the sister group of all other extant Paracortinae. The other clade has the 'three Tibet species, A.
viriosum , A . serratum and A. carinatum as the sister group of three Sichuan species, P. voluta , P. leptoclcidci and P.
thallinus. The biogeography is inferred from the most parsimonious phylogenetic hypothesis of millipedes. The
ancestral millipede in Yunnan moved First northward and diverged into two stocks. Subsequently, one of them moved
eastward first and then vicariated into an eastern population and a western population. The another moved eastward and
then separated into a western population and an eastern population. Some geological events are discussed for their
possible effects in the formation of the present pattern of millipede distribution.

RESUME

Phylogenie et biogeographie de la famille Paracortinidae (Diplopoda : Callipodida) : analyse
cladistique preliminaire.

La phylogenie et la biogeographie d’une famille de diplopodes (Paracortinidae, fam. nov.) font I'objet d'une etude
prdliminaire a l’aide d’une analyse cladistique. L’un des 35 arbres les plus parcimonieux. qui presente la valeur de F la plus
basse (la meilleure?), est congruent avec le bilan evolutif propose par Wang & Zhang (1993) & partir d’une etude
systematique. Le diplopode du Yunnan R. stimulus est le groupe-fr£re de tous les autres Paracortinae. Trois espfcces du
Tibet, A. viriosum, A. serratum et A. carinatum constituent le groupe-frtre de trois especes du Sichuan, P. voluta, P.
leptoclada and R. thallinus . 11s sont monophyletiques et constituent une trichotomie avec l’espece du Yunnan, R.
stimulus. La biogeographie est deduite de l’hypothese de plus grande parcimonie pour les diplopodes. L'ancetre des
diplopodes du Yunnan s’est deplace tout d’abord vers le nord et s’est divis6 en deux stocks. Ult6rieurement, Fun d’entre
eux s’est d’abord deplace vers Test, puis a constitu6 une vicariance entre une population orientale et une population
occidentale. L’autre s’est alors d£place vers Test et s’est s£pare entre une population occidentale et une population
orientale. La possibility d’une influence d’6v£nements geologiques dans la formation des modalites actuelles de la
repartition des diplopodes est discut£e.

INTRODUCTION

Cladistic analysis is a systematic method that attempts to discover genealogical
(phylogenetic) relationships between taxa (HENN1G, 1966; WILEY, 1981). Since a detailed

Wang. D., 1996. — A preliminary study on phylogeny and biogeography of the family Paracortinidae
(Myriapoda: Callipodida): a cladistic analysis. In: Geoffroy, J.-J .. Mauri£s, J.-P. & Nguyen Duy - Jacquemin, M..
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist. not.. 169 : 307-311. Paris ISBN : 2-85653-502-X.

308

DAQING WANG

phylogenetic hypothesis for a group of organisms can and should serve as a basis for inferring
the biogeographic history (HENNIG, 1966; BRUNDIN, 1966; NELSON & PLATNICK, 1981;
HUMPHRIES & PARENTI, 1986), I have used this approach to study the phylogenetic
relationships and biogeography of some millipedes (Callipodida: Paracortinidae). This paper
preliminary reports some of these results, that will be precised in future works.

CHARACTER ANALYSIS

I chose the species of the genus Eurygyrus as the outgroup. From the analysis of
morphology, the species of the genus Eurygyrus are close to the family Paracortinidae in
affinities and the latter is rather ancient. This result can be derived from the descriptions of the
morphological and diagnostic characters of the seven species of paracortinids. The
morphological difference and diagnostic characters of the seven species were described in the
paper, including the following ten characters listed in Table 1 (Wang & ZHANG, 1993). Table 1
gives the coding of morphological characters proposed for the paracortinid species, and Table 2
shows the matrix of the character states in the seven extant species of paracortinid millipedes.
The branch and bound algorithm from the phylogenetic computer package PAUP 2.4.1, which
guarantees the finding of all the most parsimonious trees, was used in this preliminary analysis.

Table 1. — Coding of the morphological characters for the seven paracortinid species.

Character

States of character: (code)

1. ratio of 4th-6th segments antennae

0: 1:1

1: < 1:1

2. process of 7th pair of legs in male

0: none

1: two

3. body color

0: dark brown

1: slight yellow

4. size of crests on collum

0: small and short

1 : large and long

5. shape of collum edges

0: arch-shaped

1: parallel

6. shape of the median sclerite of hypoproct

0: rectangle

1 : square

7. shape of eyes

0: ladder-shaped

1: triangular

8. gonopods

0: protrude from body

1 : include in the body

9. concavity on front of head

0: deep

1: none

10. beak-shaped process in front head

0: none

1 : large one

Table 2. — Matrix of character states in the seven new species of paracortinid millipedes. (Outgroup = genus Eurygyrus).

species

coding of characters

4 5 6 7

R. stimulus

A. carinatum

A. viriosum

R. thallinus

P. voluta

A. serratum

P. leptoclada

OUTGROUP

Source :

PHYLOGENY AND BIOGEOGRAPHY OF THE FAMILY PARACORTINIDAE

309

PHYLOGENY

A large number of trees (cladograms) was obtained, 35 in total, all with a consistency
index of 0.636. The F value of these most parsimonious trees ranged from 0.155 to 0.61 I.
However, there is only one tree with the lowest (best) F value, shown in Figure 1. BROOKS et
al. (1986) suggested that the lower the F-ratio (= F value), the greater the degree of historical
constraint on the data. In this regard, the tree with the lowest F value (Fig. 1) should exhibit the
highest degree of congruence with the
hypothesis of paracortinid evolution.

The phylogenetic hypothesis indicates
that the Yunnan species, R. stimulus is the
sister group of all other extant paracortinids.

The species A. carinatum is the sister group of
A. viriosum and A. serratum. The species
R. thallinus is the sister group of P. leptoclada
and P. voluta. Three Tibet species,

A. viriosum, A. serratum and A. carinatum,
and the three Sichuan species, P. leptoclada,

P. voluta and R. thallinus are monophyletic
and constitute the sister group that forms a
trichotomy.

Based on the fact that the setal patterns of
Callipodida not only vary distinctly among different genera, but to a large extent correspond
closely to groupings made on the basis of gonopod structure, (HOFFMAN, 1972), I propose the
classification of the genera of the family Paracortinidae as follows:

g. Paracortina: Paracortina leptoclada, Paracortina voluta

g. Relictus: Relictus stimulus, Relictus thallinus

g. Altum: Ahum viriosum. Ahum carinatum, Ahum serratum

Comparing this classification with the hypothesis, the only difference between them is the
placement of the species R. thallinus. but the rest show the highest degree of congruence. This
point would strongly support the cladogram shown in Figure 1. As for the difference, it is an
interesting question and further study is needed.

stimulus thallinus voluta leptoclada serratum viriosum carinatum

Fig. .1. — Cladogram showing hypothesized

relationships among species of paracortinids.

BIOGEOGRAPHY

According to the principle of vicariance biogeography (NELSON & PLATNICK 1981;
HUMPHRIES & PaRENTI. 1986), a pattern of spatial distribution attained by the paracortinids can
be deduced from the phylogenetic hypothesis. The area summary cladogram in Figure 2
illustrates how the present pattern of paracortinid distribution was attained.

According to the progression rule of HENNIG (1966), it implies that the ancestral

paracortinid, residing in Yunnan, diverged into
two lineages. One gave rise to the modern
Zhongdian (Yunnan) species ( stimulus ,
Fig. 2) . and the other formed the species A
(Fig. 2), which is the ancestor of all other
extant paracortinid species. Species A migrated
northward and then vicariated into two
populations: the western population (B.
Fig. 2) whose descendants later occupied
Mongkang and Deqin (Tibet), and the eastern
population (C. Fig. 2), whose descendants
gave rise to all the paracortinids in Yiajang and

Sichuan Tibet Tibet

Yunnan Yunnan Sichuan Yunnan Tibet Yunnan Yunnan

stimulus thallinus voluta leptoclada serratum viriosum carinatum

Fig. 2. — Area summary cladogram of paracortinids with
ancestral species (A - E).

310

DAQING WANG

Batang (Sichuan). Species B migrated northward and then separated into the modern species
carinatum and a species E, the ancestor of extant two Tibet species serratum and viriosum.
Species C expanded eastward and diverged into the extant species thallinus and a species D,
which is the ancestor of the extant two Sichuan species, voluta and leptoclada.

Geological events that cause fragmentation of a continuous ancestral distribution are
considered the major reasons of distribution pattern formation (NELSON & PLATNICK, 1981).
Although not all vicariant events are identifiable at present, the following are known geological
events That could have produced the present pattern of paracortinid distribution.

Mong Kang and Deqin (Tibet) lie in a strip of land between two large rivers: the western
Lancang River and the eastern Jinsha River. Batang and Yajiang (Sichuan) face the Jinsha River
to the the west and the Yalong River to the the east. The three rivers flow rapidly so that it is not
possible for paracortinids to cross. In other words, this vicariance took place before the
emergence of the three large rivers.

The formation of the three rivers is the result of elevation of the Hengduan Mountains in
the later Tertiary (PHYSIOGRAPHY OF CHINA, 1985). The fossil members of the order
Callipodida indicate that callipodoids at least were widespread in Western Asia and North
America (HOFFMAN, 1969). That is to say that it is very possible for paracortinids to migrate
and diverge before the formation of the three rivers. The Tibetan species leptoclada and Sichuan
species carinatum are being in Zhongdian (Yunnan) strongly support this idea.

The early paracortinid ancestor of Yunnan (Zhongdian) migrated northward before the
Tertiary, then diverged into extant species stimulus and species A. Species A vicariated into
western and eastern populations: Tibet species B and Sichuan species. Because of the emergence
of the Langcang River and the Jinsha River, in the Tertiary, species B was separated, and then
diverged into the extant species thallinus and a species D. The latter is the ancestor of species
voluta and leptoclada . Species C was also separated in the later Tertiary because of the
formation of the Jinsha River and the Yalong River. Then it diverged into the extant species
carinatum and the species E, which is the ancestor of two species, serratum and viriosum.

CONCLUSION

The phylogenetic hypothesis of paracortinids (FlG. 1) presented in this work is the most
parsimonious scheme derived from the cladistic analysis. It is congruent with the scheme ol
evolution proposed by WANG & ZHANG (1993). The vicariance model proposed for paracortinid
biogeography is based on the adopted most parsimonious tree that shows congruence with the
extant pattern of spatial distribution attained by paracortinids. The model interprets that (1)
paracortinids arose in Zhongdian, Yunnan; (2) migrated northward before the Tertiary and then
diverged westward and eastward; (3) because of the formation of three large rivers in the
Tertiary, western and eastern populations were separated and then diverged into the extant
paracortinids.

Nevertheless, it should be noted that the biogeographic hypothesis presented in this paper
is to be considered as preliminary. More complete analysis will be proposed in future works.

ACKNOWLEDGMENTS

I wish to express my sincere thanks to Prof. HoJushey at the California State University (USA) for permitting me
to use his computer package PAUP 2.4.1 for cladistic analysis. And 1 am grateful to Ms. Wang Xiaowei and Mr. Wang
Jian at the Institute of Zoology, Academia Sinica, for their spending much time on the computer runs. In particular, I
thank Dr J.J. Geoffroy (editor) and unkown-names referees for their critical reading, comments and suggestions on this
preliminary paper.

Source

PHYLOGENY AND BIOGEOGRAPHY OF THE FAMILY PARACORTINIDAE

311

REFERENCES

Brooks, D. R., Grady, R. T. O. & Wiley. E. O., 1986. — A measure of the information content of phylogenetic trees,
and its use as an optimality criterion. Syst. Zool., 35 : 571-581.

Brundin, L., 1966. — Transantarctic relationships and their significance, as evidenced by chironomid midges. Kungl.
Svenska Vetenskap. Handl , 11. : 1-472.

Hennig, W., 1966. — Phylogenetic systematics. Urbana, Univ. Illinois Press. 263 pp.

HOFFMAN, R. L., 1969. — Myriapoda, exclusive of Insecta. In : Treatise on Paleontology. Part R, Arthropoda 4. 2 : 57 1 -
606.

Hoffman, R. L., 1972. — Studies on Anatolian callipoid Diplopoda. Mitt. Hamburg Zool. Mus. Inst., 69 : 81-108.
Humphries, C. J. & Parenti, L. R., 1986. — Cladistic biogeography. Oxford, Clarendon Press. 98 pp.

Nelson, G., & Platnick, N., 1981. — Systematics and Biogeography, cladistics and vicariance. New- York, Columbia
Univ. Press. 567 pp.

Physiography of China, 1985. — Edited by the Committee of “Physiography of China", Beijing, Academia Sinica. Sci.
Press : 13-49.

Wiley, E. O., 1981. — Phylogenetics : The theory and pratice of phylogenetic systematics. New-York, J. Wiley & Sons.
439 pp.

Wang, D. & Zhang, C. Z., — 1993. — A new family of millipeds (Diplopoda: Callipodida) from Southwestern China.
Peking Nat. Hist. Mus., Mem., 53 : 375-389.

Source : MNHN. Paris

The Penis as a Phylogenetic Character in the Millipede

Family Julidae

Henrik ENGHOFF

Zoologisk Museum, K0benhavns Universitetsparken 15, DK-2100 K0benhavn, Danmark

ABSTRACT

The double penis provides useful characters for analysing phylogenetic relationships within the family Julidae. In his
treatment of the Diplopoda in Bronx's Klassen und Ordnungen defTierreichs. Verhoeff (1926-32) noted the difference
between Pachyiulus and the other Julids examined. Study of numerous julid genera has confirmed this distinction: All
Pachyiulini have one type of penis - other julids (with a few, obviously secondary, exceptions) have another type. The
pachyiuline type is taken to be primitive, being more similar to penis types found in related families. The other type
thus constitutes a potential synapomorphy for all Julidae except Pachyiulini. The non-pachyiuline penis type shows
several further modifications which probably qualify as synapomorphies at lower hierarchical levels. Thus, all
Paectophyllini and Caly ptophy llini have an unusually stout and sclerotized penis, and all species of Anaulaciulus have
the terminal lobes of the penis particularly long.

RESUME

Le penis comme caractere phylogenetique dans la familie Julidae (Diplopoda).

Dans la familie Julidae, le double penis fournit dcs caracteres ires utiles pour Panalyse des relations phylogen6tiques.
Dans son traite des diplopodes dans le Bronn’s Klassen und Ordnungen des Tierreichs . Verhoeff (1926-32) notait deja la
difference entre Pachyiulus et les autres julides qu’il avait observes. L’etude de nombreux genres de julides a confirme
cette distinction : tous les Pachyiulini possedent un meme type de penis alors que les autres julides (sauf quelques
exceptions traduisant & V Evidence des modifications secondaires) pr6sentent un autre type. Le type de penis des
Pachyiulini est considere comme primitif (plesiomorphe) a cause de sa similitude avec le type de penis des families
phylogenetiquement voisines. L’autre type constilue une synapomorphie potentielle pour tous les Julidae sauf les
Pachyiulini. Plusieurs modifications du type de penis non-Pachyiulini constituent probablement des synapomorphies
etablies a des niveaux infericurs. Par exemple, tous les Paectophyllini et les Calyptophy Hi ni possedent un penis
exceptionnellement robuste et sclerifie. et toutes les especes du genre Anaulaciulus presentent des lobes peniens
terminaux particulierement longs.

INTRODUCTION

As in most other millipede groups, the taxonomy of the large Palearctic family Julidae
relies heavily on the gonopods. This is true both on species level and on higher levels. Recent
studies have demonstrated, however, that certain species in some julid genera cannot be
distinguished on gonopodal characters (see, e.g., ENGHOFF. 1987. 1992), and also that the
phylogenetic relationships of julidan families cannot be satisfactorily analysed by means of
gonopodal characters alone (ENGHOFF, 1981, 1991). At the intermediate level, the only recent

Enghoff, H., 1996. — The penis as a phylogenetic character in the millipede family Julidae. In: Geoffroy,
J.-J.. MAURIES. J.-P. & NGUYEN Duy - Jacquemin, M., (eds). Acta Myriapodologica. Mem. Mus. natn. Hist, nai ., 169
313-326. Paris ISBN : 2-85653-502-X.

314

HENRIK ENGHOFF

attempt at a phylogenetic analysis of the tribes within the Julidae is that of READ (1990) which
relies very much on gonopods.

The present study focuses on another part of the male sexual system, namely the penis,
and on the phylogenetic significance of the morphological variants found within this family.

In those millipedes which do have a penis, it is a single or double tube situated behind the
second pair of legs. The penis is used to load the proper copulatory organs, the gonopods, with
sperm (HAACKER & FUCHS, 1970).

Although the penis has received relatively little attention from diplopodologists, it is
noteworthy that the higher classification of the Diplopoda by COOK (1895) to some extent was
based on the nature of the penis (or rather: of the male gonopore, since some groups have no
penis proper). Many of the names that COOK gave to higher groups, and which refer to the penis
(see HOFFMAN, 1980: 44) survive in the current classification of millipedes (HOFFMAN, 1980):
Merocheta, Diplocheta, etc.

The julid penis is double in nature: there are two gonopores and the vasa deferentia remain
separate throughout the length of the penis. The latter is therefore sometimes referred to as a
double penis, or in the plural Latin form: penes. However, it is actually only its apical lobes
which are paired, the penial basis being externally undivided. This is also true of several other
julid families having a “double" penis, although in some the two “hemipenes” seem to be fully
separated. The julid penis is devoid of setae, whereas penial setae occur in several other julidan
families (Fig. 1). In most julids each apical lobe terminates in a hyaline “membranous tube”
which probably may be retracted into the more basal, more sclerotized part of the apical lobe.
The apical lobes are sometimes separated by a median lobe.

VERHOEFF (1926-32, p.687-689) described several important details of julid penis
structure. His most important conclusions were:

1) The penis of Pachyiulus is fundamentally different from that of the other genera he
examined: Julus, Megaphyllum (sub Brachyiulus), Unciger (sub Oncoiulus), Ommatoiulus (sub
Schizophyllum), and Leptoiulus.

2) There may be considerable intraspecific variability, as demonstrated by Unciger
foetidus.

3) Some genera seem to be characterized by particular penial features ( Onunatoiulus : hood¬
like median lobe; Leptoiulus'. penis slender, parallel-sided).

Several other authors have described the penis of various julid species but it was not until
1962 that another comprehensive treatment appeared, viz., in STRASSER’s monograph of the
erstwhile tribe Typhloiulini in which he presented outline drawings of the penis of 16
“typhloiuline” species (Fig. 32).

On this background, the aim of the present study is to explore the diversity of penis
structure within the Julidae, and to assess the utility of the penis as a phylogenetic character by
interpreting the differences found in a cladistic framework.

MATERIAL AND METHODS

More than a hundred species, representing fourty-four julid genera were examined, as well as representatives of all
other julidan families (see appendix). Euparal mounts were made of isolated penes of many species, but some species
were examined with the stereo microscope only. Some penes were prepared for scanning electron microscopy (SEM)
through dehydration in absolute alcohol, transfer to acetone, and air-drying. After being mounted and coated with gold,
the penes were examined with a Jeol SP840 scanning electron microscope. Drawing conventions: Although the paired
gonoducis can often be seen by transparency, they have only been drawn in a few species.

INTRASPECIFIC VARIABILITY

VERHOEFF (1913) described intraspecific variability in penis shape, and named four
varieties of Unciger foetidus, partly based on penial characters. The varieties appeared, at least in
part, to be allopatrically distributed. Also STRASSER (1962), studying the Typhloiulini,

Source :

THE PENIS AS A PHYLOGENETIC CHARACTER IN THE MILLIPEDE FAMILY JULIDAE

315

emphasized the individual
variability. ENGHOFF (1995) found
that in spite of modest individual
variability, penis shape may be
species-characteristic in the
Paectophyllini and Calyptophyllini.

The present, more sweeping
study suggests that at most moderate
intraspecific variability is in fact the
rule, and that penis characters
therefore may be of taxonomic-
phylogenetic value in the Julidae.

Fig. I. — Penis types in julidan families. The
cladogram is that of Enghoff (1991).
The columns to the right show
whether the penis is double (D) or
single (S), and whether penial setae
are present (+) or absent (-).

Famllly

PARAJULIDAE

MONGOLIULIDAE

.PAEROMOPODIDAE

OKEANOBATIDAE

BLANIULIDAE

ZOSTERACTINIDAE

GALLIOBAT1DAE

TELSONEMASOMAT1DAE

CHELOJULIDAE

PSEUDONEMASOMATIDAE

NEMASOMATIDAE

TRICHONEMASOMAT1DAE

RHOPALOIULIDAE

TRICHOBLANIULIDAE

JULIDAE

Double /
Single

Setae

♦/-

THE PENIS IN EACH JULID TRIBE

The tribes recognized by READ ( 1 990) have been used as the taxonomic framework of this
study, with a few modifications. The differences from READ (1990) are:

- Pteridoiulini are treated separately

- Catamicrophyllini and Symphyoiulini are included in Paectophyllini

- Calyptophyllini are considered

- Typhloiulini and Leptoiulini are included in Julini.

Neither this arrangement, nor the sequence of the tribes in the treatment reflects any
definitive ideas about julid interrelationships. See, however, the section “Phylogenetic
interpretation”.

Pachyiulini

According to VERHOEFF (1926-32), the penis of Pachyiulus differs from that of the other
julids in having the apical, separate lobes relatively much longer and lying parallel to each other
(see Fig. 2). In the other julids, the separate apical lobes were much shorter and directed
obliquely lateral.

Whereas the penis structure of “other julids” is much more diverse than envisaged by
VERHOEFF, there is a remarkable constancy within the tribe Pachyiulini. ENGHOFF (1992) found
that the penis in Dolichoiulus spp. is similar to that of Pachyiulus, and subsequent studies have
shown this to be true of numerous genera of the tribe. All Pachyiulini have a hyaline penis,
without any visible cuticular reinforcements. The two “hemipenes” are fused basally as in all
julids, and the apical lobes are long and are lying parallel to each other (Figs 2, 6, 7). Only in
Mesoiulus ciliciensis do the apical lobes diverge (STRASSER, 1975, confirmed by present
study). There are no differentiated membranous tubes at the orifices, and there is no median
lobe.

316

HENRIK ENGHOFF

Figs 2-5. — Scanning electron micrographs of penis in situ of 2: Pachyiulus flavipes , posterior view, 3: Cylindroiulus
caeruleocinctus , posterior view. 4: Ophyiulus pilosus , posterior view. 5: Ophyiulus pilosus , close-up of tip,
antero-distal view. Part of the second coxae is also shown in 2-4. Scales: 0.1 mm (2-4), 0.01 mm (5).

Source : MNHN, Paris

THE PENIS AS A PHYLOGENETIC CHARACTER IN THE MILLIPEDE FAMILY JULIDAE

317

Pteridoiulini

In Pteridoiulus aspidiorum (Fig. 8), the only species of this tribe, the body of the penis is
somewhat less hyaline than in the pachyiulines. The apical lobes are short and each ends in a
hyaline “membranous tube". The sclerotization of the penis is most evident in the narrow sinus
between the apical lobes. There is no median lobe.

Figs 6-8. — Penis of Pachyiulini
(6, 7) and Pteridoiulini (8).
6: Dolichoiulus vosseleri .
7: Amblyiulus barroisi , 8:
Pteridoiulus aspidiorum. -
Scales: 0.1 mm.

Brachyiulini

In the genus Brachyiulus and in the genus Megaphyllum , the penis is very short and stout.

Anaulaciulus inaequipes. - Scales: 0.1 mm (9, 12), 0.05 mm (10, 11)

318

HENRIK ENGHOFF

It is moderately sclerotized, and the lateral margins of the basal part only occasionally have a
constriction (M. Hercules, Fig. 9). The apical lobes are parallel and short (relatively long in
M adanense. Fig. 10); they lie close to each other in Megaphyllum (Figs 9 & 10) but are
separated in Brachyiulus apfelbecki (Fig. 1 1). The membranous tubes are broad and ± parallel-
sided. There is no median lobe. In M. adanense (Fig. 10), M. geniculatum , and M. brachyurum
the membranous tubes do not arise apically but subapically on the caudal surface of the apical
tubes - perhaps a synapomorphy for part of the large, catch-all genus Megaphyllum ?

A deviating and characteristic penis type is found in the genus Anaulaciulus. Here the basal
part of the well-sclerotized penis is slenderer and has concave lateral margins; the apical lobes are
divergent and are drawn out into long, finger-shaped projections, giving a donkey-headlike
outline to the penis (Fig. 12). KORSOS (1996, this volume) found this penis type in numerous
species of Anaulaciulus and suggested it to be an autapomorphy for the genus.

LeucogeOrgiini

This small tribe shows great variability in penis structure. Archileucogeorgia (Fig. 13) and
Heteroiulus (Fig. 14) have poorly sclerotized penes approaching the type found in Pachyiulini,
although the apical lobes are shorter. Chromatoiulus (Fig. 15) looks quite like the brachyiuline
Anaulaciulus, although the long apical lobes are parallel rather than diverging. Nepalmatoiulus
(Fi°. 16) is well-sclerotized like Chromatoiulus but instead of being drawn-out the short apical
lobes have long, slender well-differentiated membranous tubes. Neither genus has a median
lobe.

Figs 13-16. — Penis of Leucogeorgiini. 13: Archileucogeorgia sp., 14: Heteroiulus intermedius , 15: Chromatoiulus
podabrus , 16: Nepalmatoiulus bir manic us (with sperm ducts and spermatozoa shown). - Scales: 0.1 mm (13, 15,
16), 0.05 mm (14).

Oncoiulini

The penis of the only studied species, Unciger foetidus (Figs 17-21) looks quite like the
penis found in most Cylindroiulini (see below): well-sclerotized, slender, with concave lateral
margins, very short diverging apical lobes and well-differentiated membranous tubes. The
species is notable for intraspecific variability, especially as regards the presence/absence and
shape of a median lobe (VERHOEFF, 1913).

Source :

THE PENIS AS A PHYLOGENETIC CHARACTER IN THE MILLIPEDE FAMILY JULIDAE

319

Figs 1 7-21. — Penis of Unciger foetidus (Oncoiulini). 17: specimen from Italy, 18-21 (from Verhoeff, 1913):
specimens from Austria (18,19), Tatra Mts. (20), and Romania (21). - Scale (17): 0.1 mm.

Paectophyllini and Calyptophyllini

These two tribes, which are probably sister-groups (ENGHOFF, 1995) share a distinctive
penis type characterized by relatively extreme sclerotization. The basal part may be either
parallel-sided (Figs 25, 27), or with diverging (Fig. 24) or concave (Figs 22 - 23, 26) margins.

Figs 22-25. — Penis of Paectophyllini (22, 23) and Calyptophyllini (24, 25). 22: Macheirdiulus libicus, 23:
Catamicrophyllum mesorientale , 24: Calyptophyllum trapezolepis , 25: C. digitcitum. Sperm ducts shown in 22,
23, and 25. - Scales: 0.1 mm.

320

HENRIK ENGHOFF

The apical lobes are extremely short to apparently absent, with small membranous tubes. The
apical margin may be straight (Fig. 27). emarginate (Figs 24-26, the emargination mterpretable
as representing the sinus between the apical lobes, or as representing a bipartite median lobe) oi
convex (Fig. 23. interpretable as representing an undivided median lobe). The detailed penis
shape seems to be species-characteristic in several cases, although there is some individual
variability For instance, the penis of Catamicrophyllum caifanum may be parallel-sided as
shown in Figure 27, or the lateral margins may diverge slightly; the apical margin may be simple
as in Figure 27, or slightly concave. In Macheiroiulus libicus , the penis may have regularly
converging lateral margins, or may be parallel except basally; the apical margin may be entire, oi
shallowly trilobate as in Figure 22.

Figs 26-27. — Scanning electron micrographs of penis in situ of Paectophyllini. posterior view. Part of the second
coxae is also shown. 26: Paectophyllum escherichii , 27: Catamicrophyllum caifanum. - Scales: 0.1 mm.

Metaiulini

Metaiulus pratensis (Fig. 28), the only species of this tribe, has a penis which resembles
that found in Paectophyllini and Calyptophyllini in being strongly sclerotized. Its shape also
resembles that found in certain paectophyllines; in particular, the regularly convex apical margin,
without any indication of a separation of two apical lobes, is a trait which is otherwise seen only
in some Catamicrophyllum species (cf. Fig. 23).

Source . MNHN. Paris

THE PENIS AS A PHYLOGENETIC CHARACTER IN THE MILLIPEDE FAMILY JL'LIDAE

321

Figs 28-31. — Penis of Metaiulini (28) and Julini (29-31). 28: Metaiulus pratensis , 29: Ophyiulus major. 30: Leptoiuius
disparatus , 31: Julus scandinavius. - Scales: 0.1 mm.

Julini

In the Julini the penis has well-differentiated apical lobes and membranous tubes. The
apical lobes are usually very short (as in Fig. 4); those of Ophyiulus major (Fig. 29), are quite

unusual for the tribe. In most species the penis
is slender, although in a few, e.g., Leptoiulus
disparatus (Fig. 30), it is stout. The lateral
margins are often straight and parallel but may
also be converging or concave. STRASSER
(1962) studied the penis in several species of
Typhloiulini (part of Julini in the present
sense) and found that the penis shape was
often characteristic of genera/subgenera in this
group (see Fig. 32).

The genus Julus itself seems to be
characterized by a very constant penis shape
(Fig. 31). The lateral margins of the slender
penis are straight and converging, and the
short apical lobes and membranous tubes are
closely applied to each other, so that the apical
outline of the penis is distinctively angled.
This shape was seen in all Julus species
examined by me and was also recorded in
Julus terrestris L.. 1758, and Julus scanicus
Lohmander, 1925 by Lohmander (1925). Of
the other genera referred to Julini s.s. by
HOFFMAN (1980) I have examined
Haplopodoiulus where the penis is. however,
similar to that found in Ophyiulus etc. On the
other hand, the "typhloiulines” Serboiulus
lucifugus and Typhloiulus lohifer appear to
have penes like those in Julus (Fig 32, o, p).

Cylindroiulini

Most Cylindroiulini have a slender penis
with very short apical lobes and well-

FiG. 32. — Penis of various "typhloiulines” (Julini)
(from STRASSER, 1962). a: Buchneria sicula
Strasser, 1959, b: B. comma Verhoeff, 1941. c:
Trogloiulus mints Manfredi. 1931. d: T. boldorii
Manfrcdi, 1940, e: Typhloiulus serbani (Ceuca.
1956), f: T. tobias Berlese. 1886. g: T. maximus
(Verhoeff. 1929). h: T. ausugi Manfredi. 1953. i:
T. illyricus Verhoeff. 1929, j: 7. montellensis
Verhoeff, 1930. k: T. albanicus Allems. 1929. I:
T. bureschi Verhoeff, 1926. m: T. psilonotus
(Latzel, 1884). n: T. strictus (Latzel, 1882). o:
Serboiulus lucifugus Strasser. 1962. p:
Typhloiulus lobifer Attems, 1951.

322

HENRIK ENGHOFF

differentiated membranous tubes. The main difference from the Julini lies in the fact that the
openings are separated by a distinct apical margin which is usually emarginate (Figs 3, 33, 34).
Cylindroiulus ruber (Fig. 35) is somewhat deviating in being stouter. However, the penis of the
closely related C. bicolor (cf. READ. 1992) looks like that found in most other cylindroiulines.
Styrioiulus pelidnus (Fig 36) deviates in having the lateral margins converging and the
membranous tubes parallel close to each other.

FIGS 33-36. — Penis of Cylindroiulini. 33: Cylindroiulus broti. 34: C. laurisilvae , 35: C. ruber , 36: Styrioiulus
pelidnus. - Scales: 0.1 mm.

Schizophyllini

Whereas the penis of Tachypodoiulus looks quite like that found in most Cylindroiulini,
the examined species of Ommatoiulus differ in having a poorly sclerotized penis with a well-
developed. undivided median lobe. In O. rutilans (Fig. 37) and O. moreleti the median lobe is
remarkably well-developed: almost the same size as each of the well-differentiated apical lobes.
In other species (Figs 38, 39) the median lobe is more modest. The apical lobes may be large
and well-differentiated (Figs 37, 39) or virtually undifferentiated (Fig. 38).

Figs 37-39. — Penis of Schizophyllini. 37: Ommatoiulus rutilans. 38: O. kessleri, 39: O. navasi. - Scales: 0.1 mm.

Source :

THE PENIS AS A PHYLOGENETIC CHARACTER IN THE MILLIPEDE FAMILY JULIDAE

323

PHYLOGENETIC INTERPRETATION

In the light of the considerable variation in penis structure found both within the Julidae,
and among julidan families, it appears worthwhile to examine whether penial characters may be
useful for elucidating phylogenetic relationships. For the sake of clearness, the analysis is
arranged in three hierarchical levels: family, tribe, and genus.

Family-level considerations

According to ENGHOFF (1981, 1991) the Julidae occupy a very subordinate phylogenetic
position in the order Julida. The closest relatives of the family are, in descending order, three
small families: Trichoblaniulidae, Rhopaloiulidae, and Trichonemasomatidae. Together with the
Julidae, these families constitute the superfamily Juloidea, one of five superfamilies in the order.

The penis types found in the Julida can roughly be divided into four categories according
to whether they are double (with separate gonopores) or single, and to whether they have setae
or not (Fig 1). Using the Spirostreptida as an outgroup (the penis in the third juliformian order,
Spirobolida, is highly deviant and hardly comparable), one may conclude that a double, setose
penis is primitive within the Julida. All examined Spirostreptida have double penes, and
although both setose and naked penes occur in this order, it is regarded more likely that the
penial setae have been lost several times independently than that they have arisen several times
independently. (Furthermore, the preliminary observations on Spirostreptida suggest that there
may have been only one loss of penial setae).

In the Julida at least four losses appear to have
occurred (Fig. 1).

As shown in Figure 1 the Julidae agree
with other Juloidea in lacking penial setae, and
they agree with Trichoblaniulidae and
Rhopaloiulidae in having a double penis. Lack
of penial setae can be considered apomorphic
for Juloidea but is a weak character since
several non-Juloidea share the character. The
double nature of the julid penis is obviously
plesiomorphic.

The two closest relatives of Julidae,

Trichoblaniulidae and Rhopaloiulidae agree in
having the penis extremely short and without
differentiation into membranous tube and basal
part (Fig. 40). The longer penis of Julidae
could therefore be interpreted as an
autapomorphy of the family, but this
interpretation is counterindicated by the
generally longer penes found in
Trichonemasomatidae and non-iuloid Julida. „ .

The penis therefore does not provide any brolemann, 1923). The basal pans of the second

very useful phylogenetic information at family- |egs are also shown, as are their tracheal

level. apodemes.

Tribe-level considerations

In his classification of Diplopoda, HOFFMAN (1980) recognized three subfamilies of
Julidae but admitted that “this family may merit the distinction of being the most difficult family
of all diplopod groups to resolve”. At the present state of knowledge of julid intra-family

324

HENRIK ENGHOFF

phylogeny it therefore appears advisable to follow READ (1990) in only operating with one
suprageneric categorial level: the tribe.

In the preliminary cladogram of julid tribes given by READ (1990) there is a basal
trichotomy between Brachyiulini, Pachyiulini (inch Pteridoiulini) and other julids. The “other
julids” have a strong potential synapomorphy in the pro-mesomerital forceps of the gonopods,
whereas neither of the two basal subfamilies have any convincing apomorphies.

The present study has confirmed the idea of VERHOEFF (1926-32): that the penis of
Pachyiulini differs from that of other julids. Although the contrast is less striking than it
appeared to VERHOEFF. the Pachyiulini are still distinguished by having a hyaline penis with
relatively long, parallel apical lobes and no differentiated membranous tubes. Most other julids
have the penis more or less sclerotized. the apical lobes are mostly shorter and are mostly
directed obliquely lateral, and there are well-differentiated membranous tubes. Those non-
pachyiulines which resemble the Pachyiulini in one or more penis characters are comfortably
nested within groups with typical non-pachyiuline penis types. For instance, species of the
genus Ommatoiulus have hyaline penes, and some species even have very long apical lobes.
However. Ommatoiulus has convincing synapomorphies with Tachypodoiulus , the latter genus
having a typical non-pachyiuline penis. Ommatoiulus + Tachypodoiulus (= Schizophyllini) in
turn have synapomorphies with other non-pachyiulines (See READ, 1990: Fig. 16).

On the whole, the pachyiuline penis type more resembles that found in other julidan
families, although the long apical lobes in Pachyiulini do not at all resemble the very short ones
in Trichoblaniulidae and Rhopaloiulidae. In particular, the lack of differentiated membranous
tubes is a trait shared with the non-julids.

The Pachyiulini might therefore tentatively be placed as sister-group to all other julids,

which are united by the potential
synapomorphy: “non-pachyiuline” penis, with
differentiated membranous tubes. Pteridoiulini
would have to be included with the latter
group, the penis of Pteridoiulus being
obviously non-pachyiuline (see Fig. 41).

A second tribe-level relationship
supported by penial characters is the sister-
group relationship between Paectophyllini (=
Catamicrophyllini + Paectophyllini +
Symphyoiulini in HOFFMAN, 1980 and READ,
1990) and Calyptophyllini. (ENGHOFF, 1995).
Whether the resemblance between the penis
type found in these tribes and in Metaiulini has
any phylogenetic significance, remains to be
shown.

Genus-level considerations

Several julid genera have a consistent penis shape which in some cases may be regarded as
a generic autapomorphy. This is probably true of

-Anaulaciulus, in which the apical lobes are diverging and are drawn out into long, finger-
shaped projections (Fig. 12, see also KORSOS, this volume).

-Julus, in which the lateral margins of the slender penis are straight and converging, and
the short apical lobes and membranous tubes are closely applied to each other, so that the apical
outline of the penis is distinctively angled (Fig. 31).

- perhaps some “Typhloiulini” (STRASSER, 1962).

L_(1)'

(2)

Pachyiulini
Pteridoiulini
Brachyiulini
other julids

Fig. 41. — Tentative basal julid phylogeny. The non-
pachyiulini penis type is a potential
synapomorphy for non-pachyiuline julids (1).
The gonopodal pro-mesomerital forceps is a
potential synapomorphy for the "other julids”
(2).

Source :

THE PENIS AS A PHYLOGENETIC CHARACTER IN THE MILLIPEDE FAMILY JULIDAE

325

-Ommatoiulus, in which the penis is poorly sclerotized and has a well-developed,
undivided median lobe (Figs 37-39).

In some other cases, the potential significance of the penis lies at the subgeneric-species
group level, as in some Typhloiulini (STRASSER. 1962) and in Megaphyllum (see above).

The penial similarity between Tachypodoiulus and Cylindroiulus deserves special mention,
because this similarity would seem to support Hoffman’s (1980) reallocation of
Tachypodoiulus in the Cylindroiulini. It is, however, not clear whether the similarity is due to
Synapomorphy, symplesiomorphy or convergence, so the penial similarity cannot be regarded as
a serious challenge to the similarities (in part clear synapomorphies) between Tachvpodoiulus
and Ommatoiulus mentioned by READ (1990).

CONCLUDING REMARKS

Although the phylogenetic conclusions of the present study may seem to be of modest
extent, it is nonetheless obvious that future students of Julidae (and Juliformia in general) should
pay more attention to penial characters than has been commonplace so far. A better
understanding of the relationships between the numerous species of Julidae, many of which
abound in a wide range of habitats in Europe, temperate Asia and (introduced) other temperate
parts of the World, can only be achieved through consideration of all kinds of characters.
Gonopods are good, but they are not everything.

ACKNOWLEDGEMENTS

I am grateful to the late Bent W. Rasmussen for help with scanning microscopy, to numerous colleagues for
helping me to build up the large collection of julids in the Zoological Museum, Copenhagen, to J. Gruber
(Naturhistorisches Museum, Wien) for loan of Rhopaloiulus, and to Z. Kors6s (Budapest) for access to his unpublished
findings.

REFERENCES

Brolemann, H. W.. 1923. — Biospeologica XLVIII. Blaniulidae. Myriapodes (Premiere Serie). Arch. Zoo l. exp. gen.,
61 : 99-453, pi. i-xvi.

Cook, O. F., 1895. — Introductory note on the families of Diplopoda. [In : O. F. COOK & G. N. Collins. The
Craspedosomatidae of North America.] Ann. N.Y. Acad. Sci., 9:9-17.

ENGHOFF, H., 1981. — A cladistic analysis and classification of the millipede order Julida. Z. zool. Svst. Evolut-
forsch ., 19 : 285- 319.

Enghoff, H.. 1987. — Revision of Nepalmaioiulus Mauries 1983 - a southeast Asiatic genus of millipedes (Diplopoda:

Julida: Julidae). Courier Forsch.-lnst. Senckenberg . 93 : 241-331.

Enghoff. H., 1991. — A revised cladistic analysis and classification of the millipede order Julida. With establishment
of four new families and description of a new nemasomatoid genus from Japan. Z. zool. Syst. Evolut.-forsch.. 29 :
241-263.

Enghoff, H.. 1992. — Dolichoiulus - a mostly Macaronesian multitude of millipedes. With the description of a related
new genus from Tenerife, Canary Islands (Diplopoda. Julida. Julidae). Ent. scand., Suppl. 40 : 1-158.

Enghoff, H.. 1995. — A revision of the Paectophyllini and Caly ptophv II i ni : millipedes of the Middle East (Diplopoda.
Julida, Julidae). ./. nat. Hist.. 29 : 685-786.

HAACKER, U. & FUCHS. S., 1970. — Das Paarungsverhalten von Cylindroiulus punctatus Leach. Z. Tierpsvchol., 27 :
641-648.

Hoffman. R. L.. 1980 [1979). — Classification of the Diplopoda. Geneve. Museum d'Histoire naturelle, 237 pp.
Hoffman, R. L., 1961. — A new genus and subfamily of the diplopod family Nemasomatidae from the Pacific Northwest.
Proc. ent. Soc. Washington , 63 : 58-64.

Lohmander. H.. 1925. — Svcriges Diplopoder. Goteborgs K. Vetensk.- o. Vitterh-Samh. Handl. 4 Foljden, 30 : 1-1 15.
Read. H.. 1990. — The generic composition and relationships of the Cylindroiulini - a cladistic analysis (Diplopoda,
Julida: Julidae). Ent. scand.. 21 : 97-112.

Read, H., 1992. — The genus Cylindroiulus Verhoeff 1894 in the faunas of the Caucasus, Turkey and Iran. Senck. biol..
72 : 373-433.

326

HENRIK ENGHOFF

SHELLEY, R. M.. 1994. — Revision of the milliped family Paeromopodidae, and elevation of the Aprosphylosomatinae
to family status (Julida: Paeromopodoidea). Ent. scand., 25 : 169-214.

Strasser, K., 1962. — Die Typhloiulini (Diplopoda Symphyognatha). - Atii Mus. civ. Stor. nai. Trieste , 23 : 1-77.
STRASSER, K., 1975. — Uber einige Diplopoden aus der Tiirkei. - Rev. suisse Zool.. 82 : 585-597.

Verhoeff, K. W., 1913. — Zur Kenntnis von Haploporatia und Oncoiulus (uber Diplopoden 60. Aulsatz). - Abh. naturw.
Ges. ISIS Dresden .1 : 1-11.

VERHOEFF, K. W., 1926-1932. — Diplopoda 1 & 2. In : H. G. BRONNS Klassen und Ordnungen des Tierreichs, 5, Leipzig,
Akademische Verlagsgesellschaft : 1-2084.

APPENDIX: EXAMINED SPECIES

With the exception of Rhopaloiulus earner atanus, all examined material belongs to the Zoological Museum,
University of Copenhagen.

NB : The Paeromopodidae sensu ENGHOFF (1981, 1991) have recently been divided into
two families: Paeromopodidae and the monospecific Aprosphylosomatidae (SHELLEY. 1994)
which together constitute the superfamily Paeromopodoidea. The penis of Aprosphylosoma
darceneae Hoffman. 1961, is double and setose (HOFFMAN, 1961: Fig. 5), like that found in
Paeromopodidae sensu stricto.

Class DIPLOPODA

FAMILY JULIDAE:

Pachyiulini

Amblyiulus barroisi (Porai, 1893)

"A " creticus (Verhoeff. 1901)

Baskoiulus stammeri Verhoeff. 1938
Chersoiulus sphinx Strasser, 1962
Dolichoiulus vosseleri (Verhoeff. 1900)

(+ 37 further spp. (see ENGHOFF 1992)
Japanoiulus lobaius Verhoeff, 1937
Mesoiulus ciliciensis Strasser. 1975
Pachyiulus flavipes (C. L. Koch. 1847)
Parapachy tutus recessus Golovatch. 1979
Rhodopieila beroni (Strasser. 1966)

Syrioiulus cf andreevi Mauries. 1984
S continentalis (Attems, 1903)

Pteridoiulini

Pteridoiulus aspidiorum Verhoeff. 1913
Brachyiulini

Megaphyllurn adanense (Verhoeff, 1943)

M. bosniense (Verhoeff. 1897)

M brachyurum (Attems. 1899)

M. geniculatum (Lohmander, 1928)

M. Hercules (Verhoeff, 1901)

M. rossicum (Timotheev, 1897)

M. taygeti (Strasser, 1976)

M tenenbaumi (Jawlowski, 1931)

Brachyiulus apfelbecki Verhoeff. 1898
Anaulaciulus inaequipes Enghoff, 1986
A. tonginus (Karsch. 1881)

Leucogeorgiini

Heteroiulus iniermedius (Brolemann, 1892)
Archileucogeorgiu sp.

Chromatoiulus podabrus (Latzel. 1884)
Nepalmatoiulus birmanicus (Pocock. 1893)

Oncoiulini

Unciger foelidus (C. L. Koch, 1838)
Paectophyllini

Paectophyllum escherichii Verhoeff, 1898
Macheiroiulus compressicauda Verhoeff. 1901
M. libicus Manfredi. 1939
Symphyoiulus impartitus (Karsch, 1888)
Mesomeritius indivisus Enghoff. 1990
Catamicrophyllum caifanum Verb.. 1901
C. mesorientale Enghoff, 1995

Calyptophyllini

Calypiophyllum digitaium Enghoff. 1995
C trapezolepis Enghoff. 1995

Metaiulini

Metaiulus pratensis Blower & Rolfe. 1956
Julini s.l.

Julus scandinavius Latzel. 1884
J. colchicus Lohmander. 1936
J subalpinus Lohmander. 1936
J. ghiljarovi Gulicka, 1963
J jedryezkowskii Golovatch. 1981
Haplopodoiulus spathtfer (Brolemann. 1897)
Pachypodoiulus eurypus (Attems. 1895)
Hypsoiulus alpivagus (Verhoeff. 1897)
Ophyiulus pilosus (Newport. 1843)

O major Bigler. 1929
O chilopogon (Berlese. 1886)

O. osellai Strasser. 1970
O. germanicus Verhoeff. 1896
O. largionii Silvestri. 1898
Lepioiulus broelenumni (Verhoeff. 1895)

L cibdetlus (Chamberlin. 1921)

L macedonicus (Attems. 1927)

L proximus (Nemec. 1896)

L. disparalus Lohmander. 1936
L. helgicus (Latzel. 1884)

L. alenuinnicus (Verhoeff. 1894)

L. tanymorphus (Attems, 1900)

Xesloiulus laeiicollis (Porat, 1889)
Peltopodoiulus schesioperovi Lohmander. 1932
Chactoleptophyllum sp
Sihiriulus dentiger Gulicka. 1963
Pacifiiulus irrtbricaius Mikhaljova, 1982

Cylindroiulini

Allajulus spinosus (Ribaut, 1904)

Cylindroiulus broti (Humbert, 1893)

C. laurisilvae Enghoff. 1982
C. caeruleocinctus (Wood. 1864)

C ruber (Lignau. 1903)

C. bicolor Lohmander. 1932
C perforatus Verhoeff. 1905
C. lalzeli (Berlese. 1884)

C. propinquus (Porat. 1870)

C. punctatus (Leach. 1815)

Siyrioiulus pelidnus (Latzel, 1884)

Enantiulus dentigerus (Verhoeff. 1901)
Kryphioiulus occultus (C. L Koch. 1847)

Schizophyllini

Tachypodoiulus niger (Leach. 1815)
Ommaioiulus cingulatus (Attems. 1927)

O. kessleri (Lohmander, 1927)

O lapidarius (Lucas. 1846)

O moreleti (Lucas. I860)

O navasi (Brolemann, 1919)

O. nivalis (Schubart. 1959)

O. oxypygus (Brandt, 1840)

O. rutilans (C. L. Koch, 1847)

O. sabulosus (L., 1758)

FAMILY TRICHOBLAN1ULIDAE
Trichoblaniulus hirsulus (Brolemann. 1889)

FAMILY RHOPALOIULIDAE
Rhopaloiulus cameratanus Attems, 1927

FAMILY TRICHONEMASOMAT1DAE
Trichonemasoma peloponesius (Mauries. 1966)

FAMILY NEMASOMATIDAE
Nemasoma varicorne (C. L. Koch, 1847)
Orinisobates spp.

Basoncopus filiformis Enghoff. 1985

FAMILY PSEUDONEMASOMATIDAE
Pseudonemasoma femorotuberculata Engholl,

1991

FAMILY CHELO JULIDAE
Chelojulus sculpturatus Enghoff. 1982

FAMILY TEUSONEMASOMATIDAE
Telsoneniasoma microps Enghoff. 1979

FAMILY GALUOBATIDAE
(Gal ti obates gracilis (Ribaut. 1909).
see BROLEMANN 1923: Fig. 18)

FAMILY ZOSTERACTINIDAE
Ameractis chirogona Enghoff. 1982

FAMILY BLANIULIDAE

(see Brolemann 1923: Figs 39. 57)

FAMILY OKEANOBATIDAE
Okeanobates serratus Verhoeff. 1939
Yosidaiulus tuberculatus Takakuwa. 1940

FAMILY PAEROMOPODIDAE
Californiulus yosemitensis Chamberlin, 1941

FAMILY MONGOLIULIDAE
Skleroprotopus coreanus (Pocock, 1895)

FAMILY PARAJULIDAE
Aniulus sp.

Karteroiulus alaskanus (Cook. 1905)
Uroblaniulus sp

Source : MNHN, Paris

Functional Morphology and Evolution of the Genitalia
of Diplopoda - Helminthomorpha

Andreas TADLER

Institut fur Zoologie, Althanstr. 14, A-1090 Wien. Austria

ABSTRACT

Theories about the evolution of genitalia (lock and key, genitalia recognition, pleiotropy, sensory female choice,
mechanical mate choice) make different predictions about the mutual coadaptation between male and female genitalia. In
three species of Chordeumatida and four species of Julida different degrees of mutual mechanical coadaptation between
male and female genitalia have been found. This supports Eberhard’s “Mechanical Mate Choice Theory". The
“Pleiotropy Hypothesis” cannot explain the evolution of diplopod genitalia because pleiotropic effects are prevented by
heterochrony.

RESUME

Morphologie fonctionnelle et evolution des genitalia des Diplopodes Helminthomorphes.

Les theories relatives & revolution des genitalia (“cl6-serrure", reconnaissance des genitalia, plSiotropie, choix
sensoriel des femelles, choix mecanique de I'accouplement), font appel & differentes hypotheses predictives sur la
coadaptation des genitalia males et femelles. Chez trois esp£ces de chordeumatides et quatre especes de julides, differents
degr<§s de coadaptation mecanique cntre genitalia male et femelle ont 6t 6 definis. Ceci vient tout d’abord appuyer la
theorie de Eberhard du “choix mecanique de raccouplement”. L’hypothese "pleiotrope" ne peut pas expliquer
1’evolution des genitalia de diplopode car les effets pleiotropiques sont evites par 1’heterochronie.

INTRODUCTION

In many animal groups, the genitalia show an evolutionary pattern quite different from
other morpho-anatomic structures. The most important questions concern the “rapid and
divergent evolution” (EBERHARD, 1985) and the high degree of complexity of genitalia.

The male gonopods in the Helminthomorpha serve as a good example for the phenomena
of diversity and complexity. There were no really fundamental changes in the peripheral
phenotype of Helminthomorpha since the middle of the paleozoic (KRAUS, 1974), however, the
gonopods have developed completely different functional principles and “ Bauplans ” in each
order and family (VERHOEFF, 1928-32).

Theories, which have been formulated to answer the general questions about the evolution
of genitalia lead to predictions about morphological complexity of female genitalia and mutual
mechanical coadaptation between male and female structures (see EBERHARD, 1985 for
discussion).

Tadler, A., 1996. — Functional morphology and evolution of genitalia in Diplopoda - Helminthomorpha. In:
Geoffroy, J.-J., Mauri£s, J.-P. & Nguyen Duy - Jacquemin, M.. (eds), Acta Myriapodologica. Mem. Mus. natn. Hist,
nat ., 169 : 327-330. Paris ISBN : 2-85653-502-X.

328

ANREASTADLER

The “lock and key” (DUFOUR, 1844. review: SHAPIRO & PORTER. 1989) and the
“genitalic recognition" theories imply that sperm transfer between members of different species
should be restricted. If a mechanical lock and key mechanism works, there should be a more or
less tight mechanical fitting between male and female genitalia. In contrast, the genitalic
recognition theory holds that heterospecific sperm transfer is avoided by species-specific
stimulation. Therefore, the genitalia of females of different species should show differences in
their sensory and nervous structures, but not in their morphology.

MAYR's “pleiotropy theory” (1963) proposes that genitalia are less subject to the corrective
influences of natural selection and that changes in the structure of genitalia are caused by
pleiotropic effects. The theory predicts that taxonomically important structural components of
genitalia have no function. A tight mechanical correlation between male and female genitalia
should not exist (KRAUS, 1968).

“Male competition” or “sperm competition” means that males can diminish the mating
success of other males, for example, by displacing sperm from the receptacula of the female or
by plugging the females' genitalia, so that the next male cannot deposit sperm (PARKER. 1970;
SMITH, 1984). The theory predicts a rapid and divergent evolution of male genitalia, but no
similar pattern in female genitalia.

The most recent theory is the “female choice theory ” (EBERHARD, 1985). This theory
supposes that females choose between males of their own species on the basis of genitalic
structures. EBERHARD proposes two mechanisms, w-hereby female can discriminate between
males:

1 ) Females discriminate between male genitalia on the basis of sensory structures, for
example, mechanoreceptors. EBERHARD speaks about “internal courtship”. Complexity of male
genitalia arises because males evolve more and more efficient stimulatory organs. The theory
predicts a rapid and divergent evolution of only male genitalia; female genitalia should be
morphologically rather simple.

2) Females discriminate between male genitalia only by the mechanical fit. If genitalia
mechanically fit well, then the probability of successful sperm transfer is high. This mechanical
mate choice theory predicts that a morphological co-evolution between male and female organs
occurs (EBERHARD, 1985).

GENITALIA FITTING IN DIPLOPODA

Analysis of diplopods frozen instantly during copulation shows that male gonopods do not
represent simple casts of the female structures, but that there are different degrees of mutual
mechanical coadaptation between male and female genitalia.

In Nemasoma varicome (Julida, Nemasomatidae) the central area of the vulvae is modified
to fit with the male solenomerit. In Brachyiulus bagnalli (Julida, Brachyiulidae), and
Cylindroiulus boleti (Julidae, Cylindroiulinae) slits on the bursae of the female vulvae
correspond to projections on the male gonopods. The female opercula of Unciger foetidus and
Cylindroiulus boleti are modified to the different mechanical forces of the male pro-mesomerit
forceps (Haacker & FUCHS, 1970; Tadler, in press).

In three species of Chordeumatida different parts of the female vulvae are modified to fit
with male parts.

On the vulvae of Haploporatia eremita (Mastigophorophyllidae), the margin bulge is
enlarged. On the distal part of the anterior gonopods of the male, there are wing-like structures.
In copula, the wing-like structures of the male gonopods fit between the margin bulge and the
bursa of the female. In Mastigona bosniense (Mastigophorophyllidae), the basis of the vulvae is
modified, so that during copulation the bursa can be rotated for more than 270 degrees. The
sperm transferring distal part of the anterior gonopods is pressed by the basis against the
openings of the receptacula (TADLER, 1989).

Source :

FUNCTIONNAL MORPHOLOGY AND EVOLUTION OF GENITALIA OF DIPLOPODA

329

In Craspedosoma transsilvanicum (Craspedosomatidae) projections on the anterior
gonopods of the male (terminal projection and clasping projection of the cheirite) insert into
invaginations of the oviduct. During copulation the bursa of the female is pulled out from the
vulval sac, the openings of the receptacula are thereby pressed against the sperm transferring
parts of the male gonopods (brushes of the syncoxite) (TADLER, 1993).

DISCUSSION

The theories mentioned above are more or less mutually compatible. Genitalia could be
influenced therefore by different evolutionary patterns.

According to the present observations, the most important factor for the evolution of
genitalia of Helminthomorpha seems to be mechanical mate choice. Following MAYNARD
SMITH (1987) female choice exists when some behaviour or structure of females causes them to
mate more successfully with some males than with others. Therefore, even the simple
evolutionary adaptation of male genitalia to female genitalia can be regarded as caused by female
choice. It is important that the general theoretical models of female choice (FISHER, 1930;
LANDE, 1981; BORGIA, 1987; POMIANKOWSKY, 1988) show that female choice concerns not
only the evolution of male traits but also the evolution of female preferences. In our examples,
change in female preference also means changes in female genital morphology. Therefore, the
mechanical mate choice theory can explain the mutual mechanical adaptation of male and female
genitalia (EBERHARD, 1985). Mechanical and sensory female choice may work together in
Diplopoda. but unfortunately there is almost no information on the sensory structures of
diplopod vulvae. Sensory female choice must therefore be examined by neuro-morphological
and neuro-physiological studies.

The mechanical coadaptation between male and female genitalia may also be an indication
that a lock and key mechanism works, but, of course the hypothesis must be tested, especially
with regard to precopulatory isolation mechanisms.

A possible mechanism of sperm displacement has been found recently in a spirostreptid
(BARNETT, Telford & DE Villiers, 1991), and there are even older observations, which
suggest, that sperm competition exists in Diplopoda. For example the secretion caps (or
“Kappenspermatophoren”) described by VERHOEFF ( 1910) for the Chordeumatid Mycogona
germanica may in fact be mating plugs. Sperm competition may be an important factor for the
evolution of gonopods, however, it cannot explain the mechanical co-evolution between male
and female genitalia.

PLEIOTROPY HYPOTHESIS

The existence of mutual mechanical coadaptation between male and female genitalia
suggests that the pleiotropy hypothesis is less important.

For diplopods, one can turn the pleiotropy hypothesis around to arrive at a more plausible
story. The two major groups of Diplopoda-Helminthomorpha, the Colobognatha and the
Eugnatha, show great differences in the complexity and morphological diversity of gonopods.
Whereas the gonopods of colobognaths are rather uniform and similar to walking legs, the
gonopods of Eugnatha show a fantastic complexity and variety of forms (VERHOEFF, 1928-32).
This may have to do with the ontogeny of the gonopods. Walking legs and gonopods are
homologuous structures, but in Eugnatha, there is a heterochrony in the development of walking
legs and gonopods. The legs of the seventh trunk unit of immature males either disappear
entirely during post embryonic development or develop in to undifferentiated bumps (ENGHOFF.
1984).

It seems unlikely that in a metameric animal mutations would effect only a single segment.
If there is no heterochrony, pleiotropic effects between gonopods and walking legs should be
present. If, for example a mutation arises, which would have an advantageous effect for the

330

ANREASTADLER

gonopods, perhaps because it makes an additional projection on the tarsus, the same mutation
would be very disadvantageous for walking legs. The rapid and divergent evolution of complex
gonopods in Eugnatha is perhaps made possible, since heterochrony prevents pleiotropic effects
between the peripheral phenotype and the gonopods.

AC KNO WLEDGEMENT

I thank Peter ZULKA for many discussions and for converting the paper to a Macintosh Computer.

REFERENCES

Barnett, M., Telford, S. R. & de Villiers, C. J., 1991. — The genital morphology of the millipede Orthoporus
pyrhocephalus (Diplopoda Spirostreptidae) - a possible mechanism of sperm displacement. Proc. Electron
Microscopy Soc. S. A., 21 : 15-17.

Borgia, G., 1987. — A critical review of sexual selection models. In : J. W., Bradbury & M. B., Anderson, Sexual
selection, testing the alternatives. New York, Brisbane, Singapore, John Wiley & Sons, Chichester, 306 pp.
DUFOUR, L., 1844. — Anatomie generate des diptfcres. Ann. Sci. nat., 1 : 244-264.

Eberhard. W. G.. 1985. — Sexual selection and animal genitalia. Cambridge Massachusetts, Harvard University Press,
244 pp.

EnGhoff, H.. 1984. — Phylogeny of millipedes, a cladistic analysis. Z. zool . Syst u. Evolutionforsch., 22 : 8-26.
Fisher, R. A., 1930. — The genetical theory of natural selection. Oxford, Clarendon Press, 272 pp.

Haacker, U. & FUCHS S., 1970. Das Paarungsverhaltcn von Cylindroiulus punctatus Leach. Z. Tierpsych ., 27 : 641 -
648.

Kraus, O., 1968. — Isolationsmechanismen und Genitalstrukturen bei wirbellosen Tieren. Zool. Anz., 171 : 22-38.
Kraus, O., 1974. — On the morphology of Paleozoic diplopods. Symp. Zool. Soc. London , 32 : 13-22.

Lande, R., 1981. — Models of speciation by sexual selection on polygenetic trails. Proc. natn. Acad. Sci. U.S.A.. 78 :
3721-3725

Maynard Smith, J., 1987. — Sexual selection - A classification of models. In : J. W. Bradbury & M. B. Andersson,
Sexual Selection: Testing the alternatives. New York. Brisbane, Toronto, Singapore, John Wiley & Sons,
Chichester.

Mayr. E.. 1963. — Animal species and evolution. Cambridge, Massachusetts, Harvard University Press. 797 pp.
Parker, G. A., 1970. — Sperm competition and its evolutionary consequences in insects. Biol. Reviews ,45 : 525-
567.

Pomi ankowsky, A. N., 1988. — The evolution of female mate preferences for male genetic quality. Oxford. Surv. evol.
Biol., 5 : 136-184.

Shapiro, A. M. & Porter, A. H., 1989. — The lock and key hypothesis: evolutionary and biosystematic interpretion of
insect genitalia. Ann. Rev. Entomol., 34 : 231-245.

SMITH. R. L., 1984. — Sperm competition and the evolution of animal mating systems. Orlando, San Diego, New York,
Academic Press, 687 pp.

Tadler, A.. 1989. — Funktionsanatomie der Kopulationsorgane und Paarungsverhalten der Diplopoda-Chordeumatida
Craspedosoma transsilvanicum, Haploporatia eremila und Mastigona hosniense. Dissertation, Univ. Wien, 100 pp.
Tadler, A., 1993. — Genitalia fitting, mating behaviour and possible hybridisation in millipedes of the genus
Craspedosoma (Diplopoda, Chordeumatida, Craspedosomatidae). Acta Zool. (Stockholm) , 74 : 215-225.

Tadler, A., in press. — Functional morphology of genitalia of four species of julidan millipedes (Diplopoda,
Nemasomatidae, Julidae). Zool. J. Linn. Soc.

VERHOEFF, K. W.. 1910. — Juliden und Ascospermophora. Jh. Ver. vaterl. Naturkde. Wiirttemberg , 66 : 337-398.
VERHOEFF, K. W., 1928-32. — Diplopoda. 2. In : H. G. Bronn’s, Klassen u. Ordnungen des Tierreichs. Leipzig,
Akademische Verlagsgesellschaft : 1073-2084.

Sperm Competition and the Evolution of Millipede

Genitalia

Mandy BARNETT * & Steven R. TELFORD **

* Department of Zoology, University of Cape Town. Rondebosch 7700, South Africa
** Department of Zoology, University of Pretoria, Pretoria 0002. South Africa

ABSTRACT

Natural selection has presumably shaped much of genital morphology for the efficient transfer of sperm, but does not
account lor the evolution of seemingly bizarre male genitalic appendages. Gonopods of several species of spirostreptid
millipedes were examined using light and scanning electron microscopy, and the sequence of events representative of
their movement within the spermathecae demonstrated through the dissection of freeze-dried copula pairs and
simulations using scale models. Gonopods bear devices that may function in sperm displacement, including flagellae
with ridges and overlapping plates, scoops and regions of pitted spines. These are orientated correctly so as to facilitate
sperm removal and are accomodated within the spermathecae of the females. This morphological evidence, coupled with
spirostreptid physiology and behaviour, indicates that sperm competition may have played a major role in shaping
gonopod morphology.

RESUME

Transfert competitif du sperme et evolution des genitalia des diplopodes.

La selection naturelle a vraisemblablement beaucoup contribue a conformer la morphologie des genitalia pour un
transfert efficace du sperme, mais elle n’explique pas revolution morphologique en apparence bizarre des appendices
genitaux des males. Les gonopodes de plusieurs espfcces de diplopodes spirostreptides ont ete examines en microscopie
optique et en microscopie clectronique b balayage et la sequence des 6v£nements traduisant le mouvement des gonopodes
a l’intfrieur de la spermatheque a ct6 mi se en evidence par la dissection des pieces copulatrices et par des simulations h
partir de moderations. Les gonopodes portent des dispositifs varies qui interviennent dans le deplacement du sperme.
incluant flagelles avec aretes, lames se recouvrant, concavites et zones recouvertes d’epines enfoncees. Ils sont orientes
de manure & faciliter la reception du sperme et s’accordent la morphologie de la spermatheque des femelles. Cette
evidence morphologique, couplee avec la physiologie et le comportemenl, indiquc que la competition pour le transfert du
sperme a pu jouer un role majeur dans la conformation morphologique du gonopode.

INTRODUCTION

Gonopods are taxonomic characters of primary importance in many millipedes (HOPKIN &
Read, 1992) but, curiously, the selective processes responsible for the evolution of these
complex structures have not been considered. Selection for effective sperm transfer presumably
accounts for much of gonopod morphology, but, as in many taxa with complex genitalia
(EBERHARD, 1985), does not fully explain their dramatic diversity.

Barnett, M. & Telford, S. R.. 1996. — Sperm competition and the evolution of millipede genitalia. In:
Geoffroy, J.-J., Mauries. J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist .
nat ., 169 : 331-339. Paris ISBN : 2-85653-502-X.

332

MANDY BARNETT& STEVEN R. TELFORD

The most likely explanation for this genital complexity is sexual selection, conceived by
Darwin (1871). Sexual selection is believed to operate through intrasexual (usually male-male)
combat and intersexual (usually female) choice. In the context of intraspecific competition,
sexual selection is believed to favour devices and behaviours of males that would prevent
interference from other males before and during copulation (THORNHILL & ALCOCK, 1983). A
significant new dimension to sexual selection theory is the concept of competition between the
ejaculates of two or more different males for the fertilisation of ova (PARKER, 1970). Sperm
competition occurs between the time of insemination and fertilisation. Mechanisms of sperm
competition include the stratification, removal and dilution of ejaculates (BlRKHEAD & HUNTER,
1990). Because sperm competition is a powerful selective force in the evolution of reproductive
behaviour and genital morphology (PARKER. 1970; SMITH, 1984; BlRKHEAD, 1989) it may
simultaneously favour the evolution of devices that enhance an individuals ability to displace,
replace or dilute a rival gametes, and behaviours that resist preemption of ejaculates (PARKER,
1970; WAAGE, 1984, 1986a; and see SMITH. 1984).

The behaviour and genital morphology of spirostreptid millipedes can be interpreted in the
context of sperm competition. All the provisos for the evolution of sperm competition are
fulfilled in millipede mating systems: they are polygynandrous, store sperm and fertilisation is
delayed (TELFORD & DANGERFIELD. 1993a,b, c). Males protect their reproductive investment
in females by prolonging the duration of copulation; a behaviour that is best interpreted as a form
of mate guarding (TELFORD & DANGERFIELD, 1991, 1993c; BARNETT & TELFORD, 1994).

Here we focus on genitalic functional morphology and argue that gonopods are adaptive
devices designed to displace (via stratification or removal) rival ejaculates. In support of this
hypothesis, we present evidence to demonstrate that the gonopods reach the distal ends of the
spermathecae, bear the necessary devices with which to displace sperm and, in some species,
move within the spermathecae to effect sperm displacement.

MILLIPEDE GONOPODS

Millipede gonopods comprise three components: the sternite, the coxite and the telopodite,
the latter of which contains the sperm canal. The gonopods are normally drawn into the body of
the male so that only the distal ends of the coxites are visable. During copulation they are
protruded and sperm are transferred from the penes to the coxite from where they are released
into and stored in the spermathecae of the vulvae of the female (BARNES, 1986' KRABBE, 1982;
BLOWER. 1985).

telopodite
coxite
sperm canal
spined region

Fig. 1. — General plan of ihe LHS gonopod of (a) Harpagophoridae, (b) Spiroslreptidae and the RHS gonopod of (c)
Odontopygidae.

Source :

SPERM COMPETITION AND THE EVOLUTION OF Mil .LIPEDE GENITALIA

333

During copulation, the telopodite
is retracted and released, causing it to
perform a sequence of twists and turns
that depend on the configuration of the
telopodite arm and its association with
the coxite. The general association of
these components is family specific
(Fig. 1).

In the Harpagophoridae and the
Spirostreptidae, the telopodite is held
within the gonocoel, a fold formed by
the coxite. It originates at the base of
the gonocoel and rises to its opening
where it bends outwards, traversing the
top of the lateral margin of the coxite.
In the Harpagophoridae the telopodite
typically ends in a rigid comb-like
structure (ATTEMS, 1928. 1937). In the
Spirostreptidae the telopodite varies
from a single arm to one which
bifurcates medially (ATTEMS, 1928,
1937). In addition, the Spirostreptidae
typically bear a region of spines on the
distal oral coxite (ATTEMS. 1928,
1937). These vary in form from stout to
hair-like spines that may or may not be
situated in pits.

The Odontopygidae have

dramatically different gonopods. The
telopodite originates at the base of the
coxite but, because the coxite has no
gonocoel. is not held within it. Instead
it passes behind the coxite and bends
inwards. Telopodites are proportionally
larger than those of the
Harpagophoridae and the

Spirostreptidae and also bifurcate. The
sperm canal is held within the whip-like
arm.

GONOPODS AS DISPLACEMENT
DEVICES

In order to actuate displacement,
the gonopods need to bear

morphological devices with which to
manipulate rival sperm. In species
shown to displace sperm (Me VEY &
SMITTLE, 1984; SlVA-JOTHY, 1984,
1987; WAAGE, 1986a. b; MlCHIELS &
Dl-IONT, 1988; RUBENSTEIN, 1989;
Miller. 1991; von Helverson &

Fig. 2. — Scanning electron micrographs of gonopod features.
Orthopoms pyroceplialus distal telopodite scoop (a) and
region of spines (b); Allopoms sp. telopodite end (c) and
spines (d); AUoporus uncinaius telopodite end showing
medial scoop (e) and spines (0: Dorcitogonus sp.
telopodite end (g) and spines (h); Chaleponcus sp.
telopodite scoop (i) and distal end of sperm-canal bearing
arm (j); Chaleponcus limbatus sperm-canal bearing arm (k)
and overlapping plates at its distal end (I).

334

MANDY BARNETT & STEVEN R. TELFORD

VON HEL VERSON, 1991; Gage, 1992) the morphological devices that have evolved to facilitate
displacement include scoops (WAAGE, 1982), spines (WAAGE, 1986a, b) and flagellae with
overlapping barbs (WAAGE, 1984).

Scoop-like structures occur on the telopodites of several species of millipede belonging to
the families Odontopygidae and Spirostreptidae. These vary in form and in their position on the
telopodite. In Orthoporoides pyrocephalus, L. Koch the telopodite terminates in a spade-like
structure (Fig. 2a). In Chaleponcus sp. the telopodite bifurcates, and one arm (the one not
bearing the sperm canal) ends in a large rounded scoop (Fig. 2i). In addition, the sperm-canal
bearing arm of the telopodite bears a series of ridges at its distal end (Fig. 2j). The telopodite of
Chaleponcus limbatus also bifurcates with the sperm canal arm bearing a series of backwardly
overlapping plates (Fig. 2k, 1), and the other arm terminating in a less rounded scoop-like
structure.

In All op or us spp. and Doratogonus sp., where the telopodite also bifurcates, a trowel-like
scoop occurs half way up the sperm canal bearing arm (Fig. 2c, e, g). The telopodites of species
belonging to the family Harpagophoridae are more robust and terminate in rigid comb-like
structures.

Interestingly, spines are found in some species belonging to the family Spirostreptidae but
not in the other two families of Spirostreptid millipedes. The distal end of the oral region of the
coxite is the only place on the gonopod where spines occur. These spines vary in form from
stout pitted spines to long hair-like spines that are pitted in some species (Fig. 2b, h) and not in
others (Fig. 2d, f).

COMPATIBILITY OF GONOPOD SIZE AND SPERMATHECAL SHAPE

To manipulate the sperm of rivals, a male's genitalia need to be able to access the areas of
the female in which sperm are stored (PARKER, 1970; WALKER, 1980; KNOWLTEN &
GREENWELL, 1984; WAAGE, 1986a). In insects, spermathecae range from relatively simple
structures to complex convoluted organs, the latter of which may restrict access of the male
genitalia to the site of sperm storage (EBERHARD, 1985). If sperm competition occurs via
displacement, then spermathecal shape and size show strong correspondence (e.g damselflies:
WAAGE, 1984, 1986a; dragonflies: WAAGE, 1986a; MILLER, 1991; and see WALKER, 1980;
EBERHARD, 1985) to the size and shape of male genitalia (WAAGE, 1984, 1986a). This may
prove to be a generalisation that holds true for all invertebrates that displace sperm.

In millipedes, female gonopores open into paired vulvae (BLOWER, 1985; HOPKIN &
READ, 1992). These are opaque structures containing roughly oval chitinous structures which
form the inner chambers of the spermathecae. The spermathecae open distally into oviducts that
join to form a common oviduct running posteriorly to the ovaries (BARNETT, TELFORD & DE
VlLLIERS, 1993). Millipede spermathecae are relatively simple structures that are species specific
in both shape and size (Fig. 3).

To actuate effective manipulation (and placement) of sperm, selection should favour the
evolution of structures that can reach the areas of the spermathecae where the sperm are stored.
In millipedes the chitinous inner chamber of the spermatheca appears to be the main site of sperm
storage (unpublished data). For each species examined, the distal ends of the telopodites of
males can be accomodated within the spermathecae and can easily reach their distal ends. Thus,
the manipulation of sperm held within these regions of the female reproductive tract is possible.

Female sperm storage organs can be very complex structures with highly sophisticated
muscular control (e.g. ViLLAVASCO, 1975). Thus, females may be able to exert some control
over fertilisation events resulting in selection acting on males to overcome this control. This
conflict of interest can generate an escalating evolutionary spiral, or arms race ( sensu DAWKINS
& KREBS, 1979) between the sexes to gain control over copulatory events. The outcome of this
process of co-evolution would be concomitant genitalic adaptation and perhaps structural
complexity. The latter is true for millipede gonopods but not spermathecae.

Source :

SPERM COMPETITION AND THE EVOLUTION OF MILLIPEDE GENITALIA

335

Alloporus sp.

1mm

Alloporus uncinatus

Doratogonus sp.

Fig. 3. — Diagrammatic representations of corresponding gonopods and spermathecae. The oval shapes within the
spermathecae represent the chitinous inner chambers. (Orihoporus - Orthoporoides in the text; Poratophilus =
Zinop hora in the text)

Interestingly, the size of the genitalia is not related to the body sizes of the animals.
Chaleponcus sp. is one of the smallest species (mean mass(g) = 2.49, SD = 0.4, n = 45) and its
spermathecae are as large as those of Alloporus uncinatus (mean mass(g) = 9.31, SD = 5.99,
n = 295). Spermathecal size and shape in Chaleponcus sp. corresponds precisely with the large
scooped distal ends of the telopodites of the gonopods of conspecific males. It should also be
noted that present descriptions of spermathecal shape are based on external topography; internal
shape may be different. Also, gonopods and/or female musculature may expand and alter the
shape of spermathecae during copulation (see VlLLAVASCO, 1975;" SlVA-JOTHY, 1987;
Walker. 1980; Miller, 1987, 1991).

336

MANDY BARNETT & STEVEN R. TELFORD

GONOPOD ORIENTATION AND MOVEMENT WITHIN THE SPERMATHECAE
Movement of the gonopods is effected by retracting the proximal end of the telopodite and
is determined by both the point of emergence of the telopodite from the coxite and the shape of
the telopodite arm. Retraction-release sequences have been reconstructed for two of the species
examined here, and illustrate that describing the structure of the distal end of the telopodite is not
sufficient to fully explain its functional morphology. The shape of the telopodite arm. and
particularly the number and tightness of spirals that it describes are critical in determining its
movement path within the spermatheca and hence its sperm displacement action. For example, in
Orthopoides pyrocephalus, the telopodite emerges at the distal end of the coxite. When it is
retracted, it traverses the bridge formed by the margin at the fold of the coxite. The scoop at its
distal end twists within the spermathecae and is then brushed against the spined region on the
distal coxite. This movement may be a mechanism whereby sperm could be removed from the
spermatheca prior to insemination (Fig. 4).

Fig. 4. — Telopodite retraction-release cycle for the RHS gonopod of
Orthoporoides pyrocephalus. Proximal retraction of the
gonopod causes the distal end to rise (2); twist forward (3);
twist back (4); twist forward (5) and then flip round to make
contact with the spined region of the coxite. Release of the
telopodite results in the scoop brushing downwards against the
spines. Source: Modified from Barnett. Telford & DE Villiers
1993.

Fig. 5. — Telopodite retraction-release cycle
of Chaleponcus sp. Retraction causes
the scoop to twist about its own axis,
flipping forward (2); forward again
(3); then backwards (4) and backwards
again (1).

Source MNHN. Paris

SPERM COMPETITION AND THE EVOLUTION OF MILLIPEDE GENITALIA

337

In contrast, the telopodite of Chaleponcus sp. emerges from the posterior base of the
coxite and retraction yields a rotation of the scoop about its own axis. There are no spines
against which the scoop is brushed, and it is predicted that the scoop functions to reposition or
mix sperm within the spermathecae but not to remove it (Fig. 5).

This comparison serves to illustrate how two apparently similar structures can have
different functions due to the shape of the telopodite arm and its resultant plane of movement.
Thus, the evolution of sperm displacing devices in millipedes will not only be linked to the actual
displacing structures, namely the distal ends of the telopodites, but to the gonopods as a whole
because the mechanism of movement is dependent on the coxite, the shape of the telopodite arm,
and associated structural modifications. This is in contrast to the damselflies in which simple
horizontal movements during copulation make it possible to predict the mechanism of sperm
competition from the morphology of the terminal region of the penis (WAAGE, 1984, 1986a).

CONCLUSIONS

The evolution of complex genital morphology in millipedes can be explained and
understood in the context of sperm competition. Gonopods display the design features necessary
for efficient sperm displacement and their complexity is probably a product of sexual selection
via sperm competition. While sperm competition implies a focus on intermale competition, the
evolutionary perpective of females is also critical to understanding genitalic evolution
(KNOWLTON & GREENWELL, 1984). The spermathecae provide the arena for competitive
interactions and females may be capable of dictating the outcome of the competition (WALKER,
1980; Eberhard, 1985).

The relatively simple spermathecal structures of female millipedes contrast with male
gonopod complexity. This suggests that the manipulative capabilities of the gonopods dictate the
intensity of sperm competition and resultant patterns of sperm precedence.

Structural modifications of the terminal region of the telopodite and the shape of the
telopodite arm together may provide an accurate prediction of gonopod functional morphology
(see Table 1).

Table 1. — Between species comparisons of gonopod functional morphology, predicted mechanisms of sperm

competition and sperm precedence patterns.

GONOPOD FEATURES DISPLACEMENT PRECEDENCE

FAMILY SPECIES TELOPODITE COXITE MECHANISM PATTERN

SPIROSTREPTIDAE

0. pyrocephalus

distal scoop

stout spines

removal

last male

A. uncinatus

medial scoop

hair-like spines

stratification

last male

Alloporus sp.

medial scoop

hair-like spines

stratification

last male

Doratosonus sp.

medial scoop

hair-like spines

stratification

last male

ODONTOPYG1DAE

C. limbatus

distal scoop
& flagellum

removal

last male

Chaleponcus sp.

distal scoop
& flagellum

—

stratification

first male

HARPAGOPHORIDAE

Z. laminata

distal comb

—

removal

last male

Zinophora sp.

distal comb

—

removal

last male

Where displacement of rival ejaculates occurs then last-male sperm precedence is the
expected outcome of a multiple mating sequence (see Waage, 1986a; MILLER. 1991). Our data

338

MANDY BARNETT* STEVEN R. TELFORD

predict last-male precedence in all but one species ( Chaleponcus sp.. Table 1). We have shown
that sperm mixing or first-male precedence is likely to occur in this species (unpublished data).
This is because the scoop-like terminal region of the telopodite redistributes rather than removes
rival ejaculates; a consequence of the shape of the telopodite arm. These results suggest a
cautious approach to ascribing a precise function to a structure without a complete understanding
of its mode of action.

Descriptive studies of genital morphology are an essential first step towards understanding
the precise function of these complex structures. Knowledge of the mode of action of the
gonopods allows a priori predictions to be made about mechanisms of sperm displacement and
patterns of sperm precedence. This is an essential basis for beginning an iterative series of
experiments and manipulations designed to quantify mechanisms of sperm competition in
millipedes. Although numerous studies have quantified patterns of sperm precedence (see
SMITH. 1984), few have attempted the more challenging task of unravelling the underlying
mechanisms.

REFERENCES

A ITEMS, C., 1928. — The Myriapoda of South Africa. Ann. S. A. Mus., 26 : 353-354.

ATTEMS, C., 1937. — The myriapoda of Natal and Zululand. Natal Mus. Ann., 7 : 459-522.

Barnes. R. D., 1986. — Invertebrate Zoology. Philadelphia, Saunders College Publishing, 743 pp.

Barnett, M.. Telford. S. R. & De Villiers, C. J., 1993. — Sperm displacement in a millipede? - an investigation into
the genital morphology of the southern African Spirostreptid millipede Orthoporus pyrhocephalus. J. Zool. Lond.,
211 : 511-522.

Barnett, M. & Telford, S. R., 1994. — The timing of insemination and its implications for sperm competition in a
millipede with prolonged copulation. Anim. Behav.. 48 : 482-484.

Birkhead, T. R., 1989. — The intelligent sperm? A concise review of sperm competition. J. Zool. Lond., 218 : 347-
351.

Birkhead, T. R. & Hunter, F. M.. 1990. — Mechanisms of sperm competition. Trends Ecol. Evol ., 5 : 48-52.

Blower, J. G.. 1985. — Millipedes ( Synopses of the Br. Fauna NS. 35). London, E. J. Brill & W. Backhuys, 242 pp.

Darwin, C.. 1871. — The descent of man and selection in relation to sex. London, John Murray, 693 pp.

Dawkins. R. & Krebs. J. R., 1979. — Arms races between and within species. Proc. R. Soc. Lond., 205 489-51 1.

Eberhard, W. G., 1985. — Sexual selection and animal genitalia. Cambridge, Harvard, University Press, 231 pp.

Gage, M. J. G.. 1992. — Removal of rival sperm during copulation in a beetle. Tenebrio molitor. Anim. Behav., 44 :
587-589.

Hopkin, S. P. & Read. H. J.. 1992. — The biology of millipedes. Oxford. Oxford Univ. Press, 233 pp.

Knowlten, N. & Greenwell, S. R.. 1984. — Male sperm competition avoidance mechanisms: The influence of female
interests. In: R. L. SMITH, Sperm competition and the evolution of animal mating systems. New-York, Academic
Press : 62-83.

Krabbe, E., 1982. — Systematik der Spirostreptidae (Diplopoda: Spirostreptomorpha). Ablt. naturw. ver. Hamburg
(N.F.), 24 : 1-146.

McVey, M. E. & Smittle, B. J., 1984. — Sperm precedence in the dragonfly Erythemis simplicicollis. J. Insect
Physiol.. 30 : 619-628.

M ichiels, N. K. & Dhont, A. A., 1988. — Direct and indirect estimates of sperm precedence and displacement in the
dragonfly Sympetrum danae. Behav. Ecol. Sociobiol., 23 : 257-263.

Miller, P. L., 1987. — Sperm competition in Ischnura elegans. Odonatologica 16 : 201-207.

Miller. P. L.. 1991. — The structure and function of the genitalia in the Libellulidae (Odonata). Zool. J. Linn. Soc.,
102 : 43-73.

Parker, G. A., 1970. — Sperm competition and its evolutionary consequences in the insects. Biol. Rev., 45 : 525-
567.

Rubenstein. D. I., 1989. — Sperm competition in the water strider Gerris remigis. Anim. Behav., 38 : 631-636.

Siva-Jothy. M. T. 1984. — Sperm competition in the family Libellulidae (Anisoptera) with special reference to
Crocoihemis erythraea (Brulle) and Orthetrum cancellation (L.). Adv. Odonatoi, 2 : 195-207.

Siva-Jothy, M. T. 1987. — The structure and function of the female sperm storage organs in libcllulid dragonflies. J.
Insect Physiol., 33 : 559-567.

Source : MNHN, Paris

SPERM COMPETITION AND THE EVOLUTION OF MILLIPEDE GENITALIA

339

SMITH, R. L. 1984. — Sperm competition and the evolution of animal mating systems. New York, Academic Press,
687 pp.

Telford, S. R. & Danghrfield, J. M.. 1991. — Sex ratio manipulation and copulation duration in the tropical
millipede, Alloporus uncinatus : A test of the copulatory guarding hypothesis. Anim . Behav ., 40 : 984-986.
TELFORD, S. R. & Danghrfield, J. M., 1993a. — Mating tactics in the tropical millipede Alloporus uncinatus
(Diplopoda; Spirostreptidae). Behaviour, 124 : 45-50.

TELFORD, S. R. & Dangerfield, J. M., 1993b. — Mating behaviour and mate choice experiments in some tropical
millipedes (Diplopoda: Spirostreptidae). S. A. J. Zool., 28 : 155-160.

Telford, S. R. & Dangerfield, J. M., 1994. — Males control the duration of copulation in the tropical millipede,
Alloporus uncinatus. S. A. J. Zool., 29 : 266-268.

Thornhill, R. & ALCOCK, J., 1983. — The evolution of insect mating systems. Cambridge, Massachusetts, Harvard,
University Press, 547 pp.

Villavasco, E. J.. 1975. — Functions of the spermathecal muscle of the boll weevil, Anthonomus grandis. J. Insect.
Physiol., 21 : 1275-1278.

Von HELVERSON, D. & Von HELVERSON. O., 1991. — Premating sperm removal in the bushcricket Metaplastes ornatus.
Behav. Ecol. Sociohiol., 28 : 391-396.

Waage, J. K., 1982. — Sperm displacement by male Lestes vigilax (Zygoptera: Lestidae). Odonatologica, 11 :
201-209.

Waage, J. K., 1984. — Sperm competition and the evolution of odonate mating systems. In : R. L. Smith, Sperm
Competition and the Evolution of Animal Mating Systems. New- York, Academic Press : 251-288.

Waage, J. K., 1986a. — Evidence for widespread sperm displacement ability among Zygoptera (Odonata) and the means
for predicting its presence. Biol. J. Linn. Soc., 28 : 285-300.

Waage, J. K., 1986b. — Sperm displacement by two libellulid dragonflies with disparate copulation durations.
Odonatologica, 15 : 429-444.

Walker, W. F.. 1980. — Sperm utilisation strategies in nonsocial insects. Am. Nat., 115 : 780-799.

Source : MNHN. Paris

Preliminary Data on the Anatomy of the Genital
Systems in Craterostigmus tasmanianus
(Craterostigmomorpha) and Esastigmatobius longitarsis
(Henicopidae, Lithobiomorpha) (Chilopoda)

Carol C. PRUNESCU *, Robert MESIBOV **& Keizaburo

Shin ohara ***

* Institute ot Biology, 296 Spl. Independentei, RO-79651 Bucarest, Romania
** P.O. Box 431, Smithton, Tasmania 7330, Australia
*** 7-4, Wakamiya, 2-chome, Ichiara-si, J-Chiba 290, Japan

ABSTRACT

Microanatomical studies on the genital system of Esastigmatobius longitarsis and Craterostigmus tasmanianus
provide arguments to establish phyletic relationships between Henicopidae and Lithobiidae, and on the other hand
between Craterostigmomorpha, Lithobiomorpha and epimorphic chilopods. Microanatomical studies on adult males of
Craterostigmus tasmanianus indicate the presence of paired testes connected by the efferent canals to a single median
deferens canal. The male genital system of C. tasmanianus is similar to that of the orders Scolopendromorpha and
Geophilomorpha. The male genital system in Esastigmatobius longitarsis presents a single flagelliform median testis
and two seminal vesicles. The testis is continued by a deferens canal which opens in the zone of the confluence of the
two seminal vesicles. The genital tract is continued by two ejaculatory ducts, which open separately into the atrium. The
glandular system of the male genital tract is composed of dorsal accessory glands, ventral accessory glands and atrial
glands. In the single testis there is a unique type of spermatogenesis producing spermatocytes of large size. The female
genital system, similar to that ot Lithobiidae, is also described. The possible phyletic relationships of the family
Henicopidae s. st . and the tribe Anopsobiini are discussed.

RESUME

Donnees preliminaires sur I'anatomie du systeme genital male chez Craterostigmus tasmanianus
(Craterostigmomorpha) et des systemes genitaux male et femelle chez Esastigmatobius
longitarsis (Henicopidae, Lithobiomorpha) (Chilopoda).

Des etudes d'anatomie, en microscopic optique. des individus males de C. tasmanianus indiquent la presence de paires
de vesicules testiculaires reliecs par des canaux afferents a un canal deferent central, impair. Le systeme genital male de
Craterostigmus est similaire au systeme genital des ordres Scolopendromorpha et Geophilomorpha. Le systeme genital
male chez Esastigmatobius longitarsis presente un seul testicule median, flagelliforme et deux vesicules s£minales. Le
testicule est prolonge par un canal deferent allonge et contourn£ qui debouche dans la zone de confluence des vesicules
seminales. Le systeme glandulaire du tractus genital male est forme de glandes accessoires dorsales, accessoires ventrales
et atriales. Dans le testicule se d£roule un type de spermatogenese unique avec des spermatocytes de grande taille. On
decrit aussi le systeme genital femelle de cetle espece qui est semblable au systeme genital femelle des Lithobiidae. La
discussion finale concerne les relations phylogenetiques de la famille Henicopidae 5. str. et de la tribu Anopsobiini.

Prunescu, C. C., Mesibov, R. & Shinohara, K.. 1996. — Preliminary data on the anatomy of the genital
systems in Craterostigmus tasmanianus (Craterostigmomorpha) and Esastigmatobius longitarsis (Henicopidae,
Lithobiomorpha) (Chilopoda). In: Geoffroy. J.-J.. Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn . Hist, nat ., 169 : 341-346. Paris ISBN : 2-85653-502-X.

342

CAROL CONSTANTIN PRUNESCU, ROBERT MESIBOV & KEIZABURO SHINOHARA

INTRODUCTION

Genital systems are relatively well studied within the family Lithobiidae (cf. ATTF.MS,
1926; PRUNESCU, 1964. 1965a; RILLING, 1968 ; LEWIS, 1981).

Many years after the description of the microanatomy of the female genital system in
Craterostigmus tasmanianus was published (PRUNESCU, 1965b), we obtained new specimens
of this species. Difficulties in fixing and preserving of this material as well as a disadvantageous
sex ratio of this lot have not allowed us to make good quality dissections capable of clearing up
the problem of the precise number of pseudometameric testicular vesicles. Taking into account
that any positive data, even incomplete, dealing with the order Craterostigmomorpha are
opportune and can be useful in this period of cladistic phylogeny, we propose the preliminary
data presently at our disposal.

In the family Henicopidae, which presents an extra European distribution, the genital
system in general does not yet seem to have been studied. We present microanatomical data on
the genital system in Esastigmatobius longitarsis, as they appear from the study of the serial
cross-sections of the posterior half of the body, in male and female individuals.

MATERIAL AND METHODS

Several individuals of C. tasmanianus were collected from Goderich Road (N.W. Tasmania) by R. Mesibov at an
altitude of 580 m, on 12 September. 1991. They were fixed in 3% glutaraldehyde, in cacodylat buffer. pH 7.4, for 4 days.
The parts were then placed in 70% ethylic alcohol. After routine histological technique, paraffin-embedded material was
sectioned at 6 pm and coloured with hemalum-eosine. Some adult male and female individuals of E. longitarsis, collected
from Japan by K. Shinohara and fixed in 70% ethylic alcohol were studied using the same methods.

RESULTS

Genital system o/Craterostigmus tasmanianus

The testicular system of Craterostigmus tasmanianus consists of several testicular vesicles
placed one side and another of a deferens duct (Figs 1-4). Each testicular vesicle is an elongated
and sinuous formation. It communicates with the central deferens duct by two other afferens
ducts, one anterior and the other posterior.

The deferens duct is bifurcated, in the hind-gut area, into two ejaculatory ducts (Figs 5-6)
which descend to the ventral region of the body and open into the male genital atrium. The two
ducts of the dorsal accessory glands also open here. The two ducts of the ventral accessory
glands open into an unpaired ventral duct, which in turn communicates with the genital atrium
(Fig. 6). Both dorsal and ventral accessory glands are well-developed acinous glands.

Male genital system o/ Esastigmatobius longitarsis

The testis is unpaired, tubular and elongated (Fig. 7). Towards the posterior end, the
lumen of the testis narrows and takes on the aspect of a deferens duct (Fig. 8). In its caudal part,
the deferens duct holds numerous spermatogonia and even small spermatocytes. These
spermatocytes occur in different stages of cellular degeneration.

On the left and right sides of the testicle, there are elongated, tubular, seminal vesicles,
situated dorsally relative to the medium intestine and closed at their anterior end like a glove
finger. The two seminal vesicles join and immediately after their joining, in the medio-dorsal part
of the resulting formation, the deferens duct of the testis opens (Fig. 9).

The genital tractus is continued by two ejaculatory ducts (Fig. 10) which descend by the
posterior intestine. In their anterior part, the ejaculatory ducts are represented by two large,
dilated tubes with thick walls consisting of a secretory cylindric epithelium. This epithelium
synthesizes and secretes into the lumen a finely granulated eosinophil-rich secretion, mixed with
numerous basophil granules. As these ducts descend in the ventral and caudal part of the body,
their diameter grows smaller and the lumen narrows.

Source :

GENITAL SYSTEM IN SOMECHILOPODA

343

Figs 1-3. — Sections through the testicular vesicles at the deferens canal level. Note the efferens canals and the deferens
canal. xl20.

Fig. 4. — Testicular vesicle (detail). Spermatogonia and spermatocytes are seen during the division process or at the
beginning of the growth. x500.

Fig. 5. — Transverse section through the hind-gut region. The two ejaculatory canals (arrowed) can be seen between the
hind-gut and a ventral nerve ganglion. x!20.

Fig. 6. — Section through the male genital atrium. Ejaculatory canals (arrow). xl60.

The single pair of dorsal accessory glands and the single pair of ventral accessory glands
arc acinous tubular glands. The ducts of the ventral accessory glands join (anteriorly) to form an
unique duct which represents part of the genital atrium (Fig. 1 1). The ducts of the dorsal
accessory gland also join, posteriorly, to form a single duct. It is continued by a cylindrical

Source :

344

CAROL CONSTANTIN PRUNESCU. ROBERT MESIBOV & KEIZABURO SHINOHARA

structure which represents the central part of the genital atrium, also named the central duct of the
atrium (Fig. 1 1 ).

The two ejaculatory ducts
penetrate separately into a dorsal
structure of the genital atrium
(Fig. 12). The bottom of this
structure consists of a glandular
epithelium. The ejaculatory ducts
open in the atrium caudally, after
the central and the unique ducts of
the ventral accessory glands join
and open outwards. In the atrium,
an acinous atrial gland forms
caudally (Fig. 13).

Fig. 7. — Transverse section through the
single testes; it presents many
groups of spermatocytes and fascicles
of spermatozoa. xIOO. The thick
arrow shows the top of the slide.

Fig. 8. — Transverse section through
posterior extremity of seminal
vesicles. Within these vesicles can
be seen fascicles of spermatozoa and
a granular secretory material.
Between the seminal vesicles, many
profiles of the deferens duct are
visible. x90. The thick arrow shows
the top of the slide.

Fig. 9. — Transverse section at the level of
the joining of the two seminal
vesicles; the deferens duct can be seen
dorsally and the hind-gut ventrally.
x 90. The thick arrow shows the top
of the slide.

Fig. 10. — Transverse section at the level of
the ejaculatory ducts. x60.

Fig. 11. — Transverse section through the
ejaculatory ducts at the level of the
genital atrium. Single arrow indicates
the unique duct of the ventral
accessory glands: double arrow the
central duct of the atrium. x90.

Fig. 12. — Genital atrium at the level of the
opening of the ejaculatory ducts
(arrow). xI40.

Source : MNHN, Paris

GENITAL SYSTEM IN SOMECHILOPODA

345

Female genital system o/'Esastigmatobius longitarsis

The female genital system consists of an elongated tubular ovary which is above or beside
the mid-gut (Fig. 14). The caudal
part of the ovary is continued by
two oviducts which descend and
surround the posterior intestine. A
pair of large seminal receptacles
(Fig. 15) communicates with the
genital atrium by a narrow duct. It
is surrounded by a sheath of
circular muscles. The genital atrium
is a large structure which presents,
in its anterior part, a high epithelium
surrounded by numerous glandular
acini, the ducts of which open into
the atrium (Fig. 16). In cross-
section, the female genital atrium
presents a V-shape. Each oviduct
opens into the corresponding latero-
dorsal end (Fig. 16). The ducts of
ventral glands open separately
through the latero-ventral walls of
the atrium (Fig. 17). The ducts of
the seminal receptacles wind several
times in light spirals, penetrate the
dorsal wall of the atrium, and open
into the terminal part of the atrium
(Fig. 18). The dorsal accessory
glands open at about the same level,
through the latero-dorsal walls of
the atrium.

Fig. 13. — Acinous atrial gland. xl40.

Fig. 14. — Transverse section through the
ovary. xlOO.

Fig. 15. — Transverse section through the
seminal receptacles (arrow). Inside of
the receptacles are masses of
spermatozoa. x60.

Fig. 16. — Transverse section through the
anterior region of the genital atrium
at the level of the opening of the
oviducts. Arrow indicates oviduct, the
arrowheads the ducts of the seminal
receptacles, the double arrow the
ducts of the accessory dorsal glands.
x90.

Fig. 17. — Opening of the ducts of the
ventral accessory glands (arrow).
x90.

Fig. 18. — The terminal zone of the genital
atrium. Opening of the ducts of the
semi-receptacles. x200.

Source :

346

CAROL CONSTANTIN PRUNESCU. ROBERT MESIBOV & KF.IZABURO SHINOHARA

DISCUSSION

Our data on the testicular system of Craterostigmus tasmanianus do not allow us to specify
the number of the vesicular testicles. That seems to be equal to or larger than two pairs. The
testicular system in C. tasmanianus resembles that of epimoiphic chilopods. We have to mention
the missing of the seminal vesicles, characteristic for Lithobiomorpha. The fact that the larva of
C. tasmanianus has 12 leg-bearing segments at its eclosion (MANTON, 1965) shows that this line
detached from the main evolutionary line which linked anamorphic chilopods to epimorphic
ones. It is known that Craterostigmus females take care of their eggs (LEWIS, 1981) as do all
epimorphic chilopods. The presence of pseudometameric testes in a chilopod with 15 leg-bearing
segments and many resemblances in its outer morphology and way of life with the epimorphic
chilopods with an elongated body may appeared and evolved from ancestors with anamorphic
features (PRUNESCU. 1969a).

The male genital system of E. longitarsis resembles that of Lithobiidae. Unlike the
Lithobiidae, whose ejaculatory ducts join before opening in the atrium by an unique ejaculatory
duct, those of Henicopidae have the ejaculatory ducts opening separately into the genital atrium.
The presence of paired male genital tracts in some genera of Anopsobiini (Henicopidae)
(PRUNESCU & JOHNS, 1969: PRUNESCU, 1992a), indicates the phyletic complexity of
Henicopidae and supports the idea that they have retained numerous plesiomorphic features of
the genital system. The existence of a male gonopod of 4 articles in all lines of Henicopidae also
argues for the primitiveness of this group but. at the same time, covers its heterogeneity. The
presence in the deferens duct of the single testicle of small elements belonging to an abortive
spermatogenesis, suggests a closeness of this phenomenon to the microspermatogenesis in the
deferens duct-microtestes in Scutigera (FAHLANDER, 1938; PRUNESCU, 1969b. 1992b) and
Anopsobiidae (PRUNESCU & JOHNS, 1969). The female genital system presents less significant
morphological differences in comparison with that of in Lithobiidae.

ACKNOWLEDGMENTS

The iwo original papers which conslituted the present work have been revised and successfully unified by Dr J.-J.
Geoffroy (Editor). We deeply thank him for this benefic effort.

REFERENCES

Ant MS, C., 1926. — Chilopoda. In : W. KUKENTHAL, Handbuch der Zoologie. Progoneata. Chilopoda. Insecta. Berlin
und Leipzig, W. de Gruyter & Co. : 239-402.

FAHLANDER, K., 1938. — Beitrage zur Anatomie und systematischcn Eintcilung dcr Chilopoden. Zool. Beidr. Upps .,
17 : 1-148.

Lewis, J. G. E., 1981. — The biology of Centipedes. Cambridge, Cambridge Univ. Press, 475 pp.

Manton, M. S., 1965. — The evolution of arthropod locomotory mechanisms, 8. Zool J. Linn. Soc. . 46 : 251-483.
Prunescu, C. C., 1964. — Anatomic microscopique du systeme genital male des Lithobiides. Rev. Roum. Biol (Zool),
9: 101-104.

Prunescu, C. C., 1965a. — Contribution a I’etude anatomique et anatomo-microscopique du systeme gdnital femelle de
l ordre Lithobiomorpha. Rev. Roum. Biol (Zool), 10 : 11-16.

PRUNESCU, C. C., 1965b. — Les systemes genital el tracheal de Craterostigmus (Chilopoda). Rev. Roum. Biol (Zool),
10 : 309-312.

PRUNESCU, C. C., 1969a. — Quelle est la place occupce par Cermatobius, Craterostigmus et Plutonium dans la
phylogenie des Chilopodes? Bull. Mus. natl. Hist. nat. Paris, 41, suppl. 2 : 112-115.

Prunescu, C. C., 1969b. — Le systeme genital male de S. coleoptrata (Notostigmophora, Chilopoda). Rev. Roum.
Biol. (Zool.), 14 . 185-190.

PRUNESCU, C. C., 1992a. — The genital system in Dichelobius (Anopsobiidae, Lithobiomorpha, Chilopoda). Ber. nat.-
med . Verein Innsbruck, suppl. 10 . 87-91.

PRUNESCU, C. C., 1992b. — The beginning of double spermatogenesis in Scutigera coleoptrata. Ber. nat.- med . Verein
Innsbruck, suppl. 10 : 93-97.

Prunescu, C. C. & Johns, M., 1969. — An embryonic gonad in adult males of Anopsobius neozelandicus Silv.

(Chilopoda). Rev. Roum. Biol. (Zool.), 14 : 407-409.

Rll.UNG, G., 1968. — Lithobius forficalus. In : Grosses Zoologisch Praktikum, part. 13 b. Stuttgart. Fischer.

On Some Structural Abnormalities in Dignathodon
microcephalum (Lucas, 1846) and their Possible

Significance

Francisco J. Santibanez & Andres Garcia Ruiz

Departamento de Biologi'a Animal I (Entomologfa), Facultad de Ciencias Biologicas, Universidad Complutense

E-28040-Madrid, Spain

ABSTRACT

Some specimens of Dignathodon microcephalum (Lucas. 1846) with some structural abnormalities in the antennal
articles and the last pair of legs are described. There is no indication of damage or regeneration in Jhese specimens and we
presume that these are developmental abnormalities.

RESUME

Signification de quelques anomalies de structure chez Dignathodon microcephalum (Lucas,
1846).

On a ctudie des specimens de Dignathodon microcephalum prSsentant des anomalies de structure sur les articles
antennaires et la derniere paire de panes. L’absence de toute trace de dommage ou de regeneration chez les individus
observes am£ncnt a penser qu’il s’agit d'un developpement anormal.

INTRODUCTION

Among the large number of centipedes we have studied during the last few years we have
found some specimens with abnormal structures. MlNELLI & PASQUAL (1986) only found three
types of abnormal structures on centipedes: spiral segmentation, mutation of a structure into
another and branched appendix.

According to Lewis (1987) some anomalous structures in centipedes may not fit into
MlNELLI & PASQUAL's classification (1986) because in most of the cases the anomalous
structures are due to problems in the animal development or to structural regeneration after
damage.

DESCRIPTION

Abnormal size of left antenna

In a female of Dignathodon microcephalum collected at Moral de Calatrava (Ciudad Real)
on 6-1 V- 1 986 the antennae are of different sizes; both have all the antennal articles but the left
antenna is smaller than the right one because from the sixth to the penultimate article they are

Santibanez, F. J. & Garcia Ruiz, A., 1996. — On some structural abnormalities in Dignathodon microcephalum
(Lucas, 1846) and their possible significance. In: Geoffroy. J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M..
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 347-349. Paris ISBN : 2-85653-502-X.

348

FRANCISCO J. SANTIBANEZ & ANDRES GARCIA RUIZ

smaller than the corresponding articles of a normal antenna. The last article is of normal size
(Fig. 1).

Abnormal size of the last antennal article on the right antenna on a female

On a female of Dignathodon microcephalum collected in Talamanca del Jarama (Madrid),
on 30-111-1988 the last antennal article on the right antenna is three times larger than its left
equivalent (Fig. 2). This specimen shows no sign of damage and we think that the bigger size of
the right last antennal article is due to an abnormal development.

Fig. I. — Dignathodon microcephalum (Lucas, 1846). Head and antennals dorsal view.

Fig. 2. — Dignathodon microcephalum (Lucas, 1846). Head and antennals dorsal view.

Fig. 3. — Dignathodon microcephalum (Lucas, 1846). Last segment. Ventral view.

Fig. 4. — Dignathodon microcephalum (Lucas, 1846). Last segment. Ventral view.

Abnormal size of the right leg on the last pair of legs

On a female of Dignathodon microcephalum collected at Moral de Calatrava (Ciudad Real)
on 2-V-1987 the legs of the last pair were of different size. In both legs all articles are present
but the right leg is smaller than the left: from the fourth article on, the length of the articles is
lesser than the size of the corresponding (Fig. 3).

Source :

SOME S I RUCTURAL ABNORMALITIES IN D/GNATHODON MICROCEPHALUM

349

l described ^ similar case in a specimen of Tygarrup javanicus (Attems

U/ \\ U 7 eft eg was smaller than the n§ht one due to the different sizes of the articles
On the aforementioned specimen there is no sign of damage. We think that the smaller size
ol the ai tides on the right leg of the last pair of legs is due to abnormal development.

Abnormal development of the last four articles on the last left leg

-jo of Dignathodon microcephalum collected at Talamanca del Jarama (Madrid) on

: . , , 8 , , si four articles on the last left leg are not articulated instead there is a bigger

article that would fit with the fusion of the last four articles, because the size of the two le<>s is
the same (Fig. 4). 6

On the aforementioned specimen there is no sign of damage. We think that this fusion of
the last tour articles is due to abnormal development on the appendage.

In conclusion, the four cases of abnormal structures studied seem really due to abnormal
development of the articles on the respective appendage, because on none of the four specimens
is there sign of damage. Because the anomaly is always based on the legs it is reasonable to
think that this is due to their different development.

REFERENCES

Lbwis. J G. E„ 1987. — On some structural abnormalities in Lithobius and Cry, , tops (Chilopoda) and their possible
significance. Bull. Gr. Brit. Myriapod, 4 : 3-6.

LEWIS, J G. E., 1988. — Tygarrup javanicus (Attems) a Geophilomorph Centipede new to the British Isles. Bull Gr
Brit. Myriapod, 5 : 3-10.

M^ELUj A,. & Pasqual, C., 1986. — On some abnormal specimens of Centipedes. Lavori - Soc. Ven. Sc. Nat.. 11 :

Source ; MNHN, Paris

Developmental Trends in the Post-Embryonic
Development of Lithobiomorph Centipedes

Alessandro MlNELLI, Enrico NEGRISOLO & Giuseppe FUSCO

Dipartimento di Biologia, Universitadi Padova. 1-35121 Padova, Italy

ABSTRACT

The problem of comparing individual developmental stages of related species undergoing a different number of moults
is tentatively settled by taking as developmental time units both major developmental periods, the first being between
hatching and the transition from the last larval to the first postlarval stage, and the second between the first postlarval
stage and the first mature stage. On such a re-scaled developmental schedule, the developmental trajectories of several
characters in different species are studied, based on Andersson’s (1979) data. This analysis revealed extensive
heterochrony, as did further comparisons of metric and meristic characters in mature specimens belonging to 62 species
with adult length ranging from 6 to 35 mm. F ’

RESUME

Modalites du developpement post-embryonnaire des chilopodes lithobiomorphes.

*fin de resoudre le probleme de la comparaison entre les stades dc developpement d’especes presentant un nombre
ditterent de mues. les auteurs utihsent en tant qu'unites temporelles les deux 6tapes fondamentales du developpement -
d une part la phase allant de Peclosion jusqu’au passage des stades larvaires aux stades post-larvaires, d'autre pan la
phase allant du premier stade post-larvaire jusqu'a la maturity sexuelle. La comparaison des trajectoires ontogenetiques de
plusieurs caracteres chez differentes especes (donnees d'apres Andersson. 1979) revele plusieurs cas d’heterochronie.
l autres heterochronies sont mises en evidence par la comparaison de plusieurs caracteres metriques et meristiques entre
des exemplaires adultes appartenant k 62 espbees, pour lesquels la longueur varie de 6 & 35 mm.

INTRODUCTION

The overall uniformity of body structure within all recent Lithobiomorpha. and especially
within Lithobius s.L, where a great many species just seem to be “minor variations on a
common theme”, provides the scope for investigations of structural and morphogenetic
constraints in the evolution of form. However, current knowledge of post-embryonic
development of lithobiomorph centipedes is still in its descriptive phase; cf. especially the
detailed studies of ANDERSSON (1979, and literature cited therein). Many additional, even basic
aspects still need investigation, but we think that many insights, at least of a qualitative nature,
can be obtained from a careful consideration of the extant evidence.

M INELLI, A., NEGRISOLO, E. & Fusco. G., 1996. — Developmental trends in the post-embryonic development of
Lithobiomorph Cemipedes. In: Geoffroy, J.-J.. Mauris. J.-P. & Nguyen Duy - Jacquemin. M.. (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist. not.. 169 : 351-358. Paris ISBN : 2-85653-502-X.

352

ALESSANDRO MINELLI. ENRICO NEGRISOLO & GIUSEPPE FUSCO

MATERIALS AND METHODS

We have mostly relied on literature data, but lor the measures of individual podomeres in legs 1-15. we have
studied L microns Meinert (a female from Italy: Miogliola (AL)) and L. forficaius (Linnaeus) (a male from Italy: Bosco
della Mesola (FE)). Two (a, b) main data bases have been collected: (a) head length (HL), body length (BL), number of
ocelli (OC). number of antennomeres (AN), number of coxal pores (CP), number of coxosternal teeth (CT), number of
setae on the first genital sterniie of the male (SGM). id. of the female (SGF). in each post-embryonic stage of Luhobius
forficaius (Linnaeus. 1758). L. erythrocephalus C. L. Koch. 1847. L. melanops Newport, 1845, L. crassipes L Koch,
1862 L microps Meinert. 1868, L. calcaraius C. L. Koch. 1844. L. cunipes C. L. Koch. 1847 and L. tenebrosus
Meinert. 1872 (data compiled from Andersson. 1979); (b) BL. OC. AN, CP. CT. total number ol spines on the legs of
one side of the animal (ST) and plectrotaxy (distribution of the individual spines on all pairs of legs) in the adults of the
62 species listed in the Appendix (data compiled from Brolemann, 1930; Eason, 1964; Matic, 1966: Andersson,
1979).

As for the methods, the less trivial points are the re-scaling of ontogenetic stages onto a normalized x-axis as
justified in the next section and explained also in the legend to Figure 1. and the standardization of the values of BL. OC,
AN, CP, CT and ST before using them in multivariate analysis (principal components).

RESULTS

Comparability of ontogenetic stadia

When comparing the ontogenetic stadia of different, although closely related, arthropod
species, the first question to answer is which stage of species B, if any, is more correctly and
meaningfully comparable with a given stage of species A. Simply referring to the ordinal number
of the shtge (i.e., to the number of moults the animal has undergone) does not seem to be safe,
because of the different number of stages that A and B may possibly go through before getting
maturity, or during their whole life. We could even be faced with mtraspecific variation in the
number of post-cmbryonic stages. One of us (MINELLI, 1992) had already looked for a possible
application to centipedes of GRANDJEAN’s (1951) concept of stase, but could not find a lixed set
of structurally distinct stages, independent from the number of moults occurring between any
two of them.

We believe that only three points, along the post-embryonic development ot all
lithobiomorphs, can be more or less safely compared across species. These are the first larval
stage (L0 in ANDERSSON’ s terminology), the first post-larval stage, i.e. the first stage with the
full complement of segments and legs (PL1 ) A, and the first mature stage. The identification of
the onset of maturity is not easy to assess in objective terms. For the species whose
developmental schedules we have compared, we have simply accepted, for operational reasons,
the estimates of maturity as given by ANDERSSON (1979), although his assessments were
derived (but for L. forficatus) from comparative guesses rather than from objective, e.g.
histological, proofs.

On this basis, we have re-scaled the post-embryonic stages of all species onto a common
scale. Wc have dealt separately with the two intervals, the first between L0 and PL1, the second
between PL1 and the first (guessed) mature stage (Fig. 1). For the species under study, this
causes no relative changes between hatching and PL1, because the number of larval stages is
always the same, whereas the individual post-embryonic stages of the different species account
for different percentages of the normalized development. Some characters, e.g. head length,
follow similar ontogenetic trajectories in all species, but others are subjected to heterochrony, at
least for some species. Ontogenetic trajectories for ocelli start at diflerent points, but
subsequently evolve in comparable ways. In still other cases, as for the setae of lirst genital
stemite in both sexes, heterochrony is given by the different speed of comparable ontogenetic
changes that start at the same time. Again, as for the number of antennomeres, the first steps are
the same in all species under comparison, but their further increase goes on at different speed;
however, the different trajectories of antennomeres in L. calcaratus (or in L. forficatus ) and
L tenebrosus coincide again at the end. The passage from larval to post-larval stages seems to be
a major change only for those characters whose phenotypic expression requires the presence oi

Source : MNHN , Paris

TRENDS IN THE DEVELOPMENT OF LITHOBIOMORPH CENTIPEDES

353

fully developed segments XII and following (e.g., the number of setae on first genital stemite);
and that, not for all species.

setae of female 1
genital segment

0.00

0.20

0.40

■

0.60 0.80 1.00

16 setae of male 1st
genital segment
14

12 -
10 -
8
6
4
2

0.00 0.20 0.40 0.60 0.80

1.00

1.80

1.60

1.40

1.20

head length

forficatus

erythrocephalus

melanops

crassipes

microps

calcaratus

curtipes

tenebrosus

Fig. 1. — Ontogenetic trajectories for individual characters in selected Lithobius species. Data after Andersson (1979).
Horizontal axis represents normalized post-embryonic development, with first larval stage (LO) and first mature
stage (guessed), respectively, at the two ends of the scale; mid-point of the x axis corresponds to PL1. Individual
points along the trajectories refer to the individual post-embryonic stages, whose number before maturity is
different in the different species. Head length in mm.

Source

354

ALESSANDRO MI NELLI. ENRICO NEGRISOLO & GIUSEPPE FUSCO

Comparisons of mature representatives of species of different size

Further comparisons have involved a comparable (mature) stage of 62 species (see
Appendix) ranging in length from 6 to 35 mm. We have tried to identify the occurrence of
coherent trends of variation of different structural traits, as well as the independence of other
traits.

CN
*-
C
0)

o o

a
E
o
o

-1

-2

-3 -2 -1 0 1 2 3 4 5 6

component 1

. 9

: * 8

.6*6

"K"v- . e

14 14

-f - h

H - 1 - 1 - 1 - H

component 1

Fig. 2. — Scatter plot of the 62 Lithobius species listed in the Appendix onto the plane of principal components 1 and
2. the original variables being in (a) BL. OC. CP. CT and ST. in (b) BL. OC. CP. CT and AN. In (a), the percentual
contributions of the first two principal components to total variance are, respectively. 72.6 and 1 1.5; in (b), the
corresponding values are 64.0 and 18.0. In (a), the numbers within the plot identify the average number of CT;
squares without numbers are for the species with CT=4.

Source : MNHN , Paris

TRENDS IN THE DEVELOPMENT OF LITHOBIOMORPH CENTIPEDES

355

II we place the 62 species in the morphospace identified by the following 5 variables: BL.
OC, CP, CT, ST in a plot of principal components 1 and 2 (Fig. 2a), we see a very coherent
distribution, with points seemingly arranged in parallel rows, each of them corresponding to an
“isoodont line”, along which are aligned the species with the same number of coxostemal teeth.

The number of antennomeres does not behave as these five characters. Figure 2b shows
the disruptive effect of substituting AN for ST in the set of original variables. That means, that
the antennae are not subjected to the same constraints as the other four variables.

Thus, these interspecific comparisons provide further evidence for heterochrony, because
ol the different variation of individual characters with size at maturity.

large size Lithobius spp.

1 2 3 4 5 6 7 8 9 10 1112 13 14 15

body segment

small size Lithobius spp.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

body segment

L. forficatus

Lsbibb&BBBmL

!■!!■■!!!!!!!■!

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
body segment

2.5 •

L. microps

1 •
0.5 I

0 I*

nnnDDQQflflflflln

!!!!!!!!!!!!!■!

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
body segment

trochanter H praefemur

femur

tibia

tarsus I tarsus II

Fig. 3. — Plectrotaxy and leg length. Bar diagrams in the upper row give the mean number of spines per podomere (coxa
to tibia) per leg (I to XV) in the eight largest species listed in the Appendix (left) and in the seven smallest
species of the same list (right). Bar diagrams in the under row give the length (mm) of the individual podomeres
in each leg (I to XV) in one individual of a large species (left: Lithobius forficatus , male) and in one individual of
a small species (right: Lithobius microps, female).

Source

356

ALESSANDRO MINELL1. ENRICO NEGRISOLO & GIUSEPPE FUSCO

Size at maturity has also an effect over the segmental pattern of distribution of leg spines.
In the small species (Fig. 3. upper row, right), the average number of spines per leg increases
steeply from segment I to segment III, goes on with trifling differences until segment IX or X,
peaks at XI, then slowly decreases, whereas in the large species (Fig. 3, upper row, left) a
“near-saturation” is soon reached on leg II. The prefemur is most responsible for the changes in
the number of spines throughout the segments.

Changes in spinulation of the proximal podomeres (coxa and trochanter) of all species and
especially of the prefemur of small species parallel leg length changes along the body. However,
leg length profiles are not different in small vs. large species (Fig. 3, under row).

FDa

15
13
1 1
9
7
5
3
1

■ ■■ ■ ■

□ □

□ □ □
□ □□□

□ □ □

□ □

□□ □ □

□ □ □

□□□□□□

□□

□ □

D □
- 1— O-

□ □

- f-o

15 20

body length

□ □
□

- <-

PVm

U)
a)

-O-O-O-a-OCDD-C

15 20

body length

Fig. 4. — Range of segments where the anterior dorsal spine of the femur (FDa. above), and the median ventral spine of
the prefemur (PVm, below) occur, in typical representatives of each of the 62 species listed in the Appendix,
plotted against average body length (mm). Full squares mark the most anterior segment, empty squares the last
segment with the given spine.

Source :

TRENDS IN THE DEVELOPMENT OF LITHOBIOMORPH CENTIPEDES

357

1 ,uLkg S1Z16 and le§ sPinulalion could be both explained as controlled, over the whole trunk
length, by at least two factors acting from centers at both ends of the body.

., For those Positions where the number of pairs of legs with spines increases with the size
ol the animal at maturity, two mam patterns are observed, one centered in the fore trunk and
another centered in the hind trunk (Fig. 4).

DISCUSSION

While acknowledging that a definitive evaluation of evolutionary trends, including
heterochronies, necessitates a sound and detailed phylogenetic background, such as we cannol
yet provide for lithobnds, we believe that our preliminary comparisons of structural and
developmenta trends are already suggestive of an interesting interplay of strong constraints -
such as those keeping together, across a large number of species, body size, number of ocelli,
simulation and number of coxal pores - with the largely independent variation of other traits- at
the same time, a great deal of specific differences seem to be easily (we would dare to say-
inexpensively) obtamed by fine-tuning starts, speeds and end points of fundamentally identical

APPENDIX

Lnl°Llhe SPCdeS *h°seuadult 'rails wcre compared (cf. Fig. 2). Data on these species, which are generally listed here
/ uw cnameS USCd, by, lhC aulhors’ were comP'led from Andersson (1979), Brolemann (1930): Eason ( 1964). Matic
Figure 3SPCC,eS marked WUh (L) °r (S) are lhe large or smal1 sPecies considered for the analysis of plectrotaxy as in

Lithobius ( Monotarsobius ) aeruginosus L. Koch. 1862
L. agilis pannonicus Loksa, 1948
L. allotyphlus Silvestri, 1908
L. aulacopus Latzel, 1880,

L. (A/.) baloghi Loksa, 1947 (S)

L. (M.) biunguiculatus Loksa, 1947 (S)

L. borealis Meinert, 1872
L. bulgaricus Vcrhoeff, 1 925 (L)

L. (M.) burzenlandicus Vcrhoeff, 1931 (S)

L. calcaratus C. L. Koch, 1844
L. castaneus Newport, 1 844 (L)

L. cavernicola Fanzago, 1877
L. ( M .) crassipes L. Koch, 1862
L. crypt icola Ribaut, 1926
L. (M.)curtipes C. L. Koch. 1847
L. cyrtopus Latzel, 1880
L. (Thracolithobius) dacicus Matic. 1959
L. decapolitus Matic et al., 1962
L. (M.) dobrogicus Matic, 1962 (S)

L. (Dacolithobius) domogledicus Matic, 1961
L. (M.) duboscqui Brolemann, 1896

(=L. microps Meinert, 1868) (S)

L. (M. ) dudichi Loksa, 1947 (S)

L. erythrocephalus C. Koch, 1 847
L.fagniezi Ribaut, 1926
L. forficatus (Linnaeus, 1758) (L)

L. inermis L. Koch. 1856
L. ( Th .) inexpectatus Matic, 1962
L. lapidicola Meinert, 1868
L. latro Meinert, 1872
L. lucifugus L. Koch, 1862
L. luteus Loksa, 1947

L. matici matici Prunescu, 1 966 (L)

L. melanops Newport, 1 845

L. (M.) microps auct. nec Meinert. 1868

L. mutabilis L. Koch, 1 862

L. muticus C.L. Koch, 1847

L. nicoeensis (Brolemann. 1904)

L. nigrifrons Latzel, 1880
L. nodulipes Latzel. 1880
L. parietum Verhoeff, 1 899 (L)

L.pelidnus Haase, 1888
L. peregrinus Latzel, 1880
L. piceus L. Koch, 1862
L. pilicornis Newport. 1844 (L)

L. punctulatus vasconicus (Chalande. 1905) (L)

L. (M.) pustulatus Matic, 1964

L. ribauti Chalande, 1 907

L. {M.) sciticus Prunescu, 1965

L. silvivagus Verhoeff, 1925

L. speluncarum Fanzago, 1877

L. (M.) subterraneus Matic, 1962 (S)

L. tricuspis Meinert, 1872

L. troglodytes scutigeropsis Brolemann, 1930

L. typhlus Latzel. 1886

L. variegatus Leach, 1817

Harpolilhobiits anodus dentatus Matic, 1957

H. banaticus Matic, 1961 (L)

H. intermedius Matic, 1958
H. oltenicus Negrea, 1 962
H. radui Matic, 1955
H. triacanthos Matic, 1964
H. tridentatus Matic, 1962

358

ALESSANDRO MINELLI. ENRICO NEGRISOLO & GIUSEPPE FUSCO

ACKNOWLEDGEMENTS

This research was supported by grants of the Italian National Research Council (CNR) and the Italian Ministry of
University and Scientific and Technical Research (MURST) to A. Minelli.

REFERENCES

A ndersson G., 1979. — Taxonomical studies on the post-embryonic development in Lithobius , with a brief
comparison with Lamyctes (Chilopoda: Lithobiomorpha). Ph. Thesis, Gdteborg. Department of Zoology, Goteborg
Univ.

Brolemann, H. W.. 1930 — Elements d’une faune des Myriapodes de France : Chilopodes. Toulouse, Imprimerie
Toulousaine, 405 pp.

Eason, E. H.. 1964. — Centipedes of the British Isles. London. Frederick Wame & Co, 294 pp.

Grandjean, F., 1951. — Les relations chronologiques entre ontogenese et phylogenese d'aprks les petits caracteres
discontinus des Acariens. Bull. biol. France Belg., 85 : 269-292.

MatIC, Z.. 1966. — Fauna Republicii Socialiste Romania. C-lasa Chilopoda Subclasa Anamorpha., Vol VI, Fasc. I.
Bucuresti. Acad. Rep. Soc. Romania. 267 pp.

Minelli, A.. 1992. — Towards a new comparative morphology of myriapods. Ber. nat.-med. Verein Innsbruck, suppl.
10 : 37-46.

Source : MNHN, Paris

Etude de la reproduction et du developpement post-
embryonnaire de Lithobius pilicornis Newport, 1844
(Chilopoda, Lithobiomorpha)

Antoni SERRA & Maria Carme Ml QUEL

Department de Biologia Animal. Facultat de Biologia, Universitat de Barcelona, Avda. Diagonal. 645

E-08028 Barcelona, Espagne

RESUME

_Dans leeadrcde 1' etude du developpement post-embryonnaire de Lithobius pilicornis. un elevaee de couples d’adultes a

£ Dmelle!niam°r|t01re ^ f °b,en'r dC* Spdclmcns d'^e connu- Aucun des males n'a depose de spermatophore mais
es cmelles, teeondces avant la capture, ont pondu de nombreux ceufs. Ces experiences ont permis dobserver la pome

' ® la n,ue et. 1 alimentation des stades juveniles. Le grand nombre de larves de differents stades de developpement

ibtenu ainsi a permis d <5tudier la variability des caractbres morphologiques. Les criteres suivants ont ete utilises alin de
e mir Ies diflerents stades : taille, spinulation des pattes. nombre d*appendices ambulatoires, d’ocelles darticles
antennaires, de dents du coxostemum forcipulaire, de pores coxaux et d’appendices genitaux.

Lithobius pilicornis Newport, 1844

ABSTRACT

Reproduction and post-embryonic development of
(Chilopoda, Lithobiomorpha).

The main goal of this work was to study the post-embryonic development of Lithobius pilicornis. Laboratory rearing
was carried out in order to provide specimens of known age. No male of the several couples reared lav spermatophores
bu the females spawned numerous eggs (they were already fertilized when captured). This experiment allowed
observations on the egg-laying, hatching, moulting and feeding of the juvenile stages reared in laboratory The lame
number of larvae in different developmental stages obtained in this way allowed the study of the variability of their
morphological leatures. The lollowing characters were used in order to establish the different post-embryonic
deve opmental stages: size number of ambulatory appendages, ocelli, antennal articles, forcipule coxostemum teeth
coxal pores, spinulation and genital appendages.

INTRODUCTION

Le processus de developpement postembryonnaire des chilopodes lithobiomorphes
comporte 1 existence de differents stades dont les caracteres different souvent de ceux qu’on
trouve chez 1 adulte. Ce fait rend difficile l’identification specifique des exemplaires non adultes.

Les auteurs interesses, jusqu a ce jour, par letude et la description des differents stades du
developpement post-embryonnaire des especes appartenant a Lordre Lithobiomorpha sont en
nombre reduit. II faut toutefois citer des auteurs comme VERHOEFF (1905), BROLEMANN

. SkRKf; A- & M|QUEC M. C, 1996. — Etude de la reproduction et du developpement post-embryonnaire de
Lithobius pilicornis Newport, 1844 (Chilopoda, Lithobiomorpha). ln\ Geoffroy. J.-J., Mauries. J.-P. & Nguyen

Duy - Jacquemin. M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 359-364. Paris ISBN : 2-85653-
502-X.

360

ANTONI SERRA & MARIA CARME MIQUEL

(1930), MURAKAMI (1958, 1960), EASON (1964, 1970), SCHEFFEL (1969) et surtout les
nombreux travaux d'ANDERSSON (1976, 1978a, 1978b, 1979, 1980, 1981a, 1981b, 1982a,
1982b, 1983, 1984a, 1984b, 1990) sur la description et la caracterisation du developpement des
differentes especes de lithobiomorphes.

Dans ce travail, les caracteristiques morphologiques sont decrites pour les differents stades
larvaires de Lithobius pilicomis Newport, 1844. Afin d'obtenir un nombre eleve d'exemplaires
d'age connu, on a fait se reproduire cette espece en captivite, ce qui nous a permis d'observer et
remarquer maints aspects de sa biologie de reproduction.

MATERIEL ET METHODES

Les individus de Lithobius pilicomis utilises pour notre elude ont 6te captures sur la Serra de Roqueroles, La Pena,
Poblet (Tarragona), le 1.X.I990 et le 6.IV.1992. Au cours de la collecte effectuee en 1990 on a obtenu 80* et 199, qui ont
cte separes en 6 couples et 15 individus (2cf et 139) isoles chacun dans une boite en matiere plastique de 12 x 8 x 6 cm,
dont le fond a ete recouvert avec une couche de 4-5 mm de terre humide a surface completement lissee. Sur I’un des cot6s,
une lame couvre-objets a 616 legerement surelevee de fagon a permettre aux individus en Slevage de s’y abriter. Ces boites
ont ainsi ete installees dans une chambre h 20-25°C pendant 1'automne et I’hiver et h 22-26°C pendant le printemps et
lete. L'alimentation reguli&re des individus consistait en des larves de Tenebrio molitor, des petits grillons, des termites
et des drosophiles ; de 1'eau distillee etait rajoutde afin de maintenir la terre humide. On remarque que, dans deux cas
seulement, la femelle a devore le male et une seule fois le male a devoid la femelle. En juin 1991. a P exception d'un seul
male isole, tous les males etaient morts et il n'y avait done plus aucun couple reproducteur. En ce qui concerne les
femelles, seulement 8 sur 19 avaient survecu ; aucune observation en relation avec la reproduction n'avait etc notee.

Le produit de la collecte de 1992 a ete de 229. installees elles-aussi dans des conditions semblables. Toutefois, le
premier substrat avail ete change, car suspecte d' avoir ete la cause du taux elev6 de mortality. Le nouveau substrat utilise a
etc obtenu par tamisage des premieres couches du sol ou les individus ont ete recoltes, dans la Serra de Roqueroles.

Pendant la periode de mai a aout 1992, de nombreux oeufs furent comptabilisSs, issus des pontes effectives par 15
des 22 9 de 1992 et des 3 survivantes de 1990. II faut cependanl remarquer, concernant ces dernieres, qu'on n'avait detecte
aucun indice de l'existence d'une ponte en 1990 et en 1991. Etant donne qu'on n’a jamais observe de spermatophores, on
peut en conclure que toutes les femelles se trouvaient deja fecondees avant leur capture.

RESULTATS

Reproduction

Nos connaissances sur les differents aspects de la reproduction des especes du genre
Lithobius , telles que la formation des couples, remission des spermatophores, la fecondation, la
taille des pontes, 1'eclosion des oeufs, la survivance de chacun des stades larvaires, etc., sont
tres limitees. Peu de biologistes ont essaye de reproduire des especes de Lithobius en captivite ;
dans ce sens il faut remarquer le travail realise par DEMANGE (1956) sur Lithobius piceus
gracilitarsis, qui apporte des donnees interessantes sur la biologie de cette espece. ANDERSSON
(1978a, 1981b) expose aussi brievement la methodologie utilisee pour reussir la reproduction de
differentes especes au laboratoire afin d’obtenir des exemplaires permettant l'etude du
developpement post-embryonnaire.

Le Tableau 1 donne le nombre d'oeufs pour la ponte de chacune des femelles, le nombre
d’eclosions et le pourcentage de viabilite de celles-ci ainsi que les valeurs moyennes de ces
parametres.

L'enorme disparite de taille des pontes (entre 5 et 105 oeufs!) est difficilement explicable ;
il faut tenir compte du fait que des oeufs ont pu echapper a notre attention car Pootheque qui les
protege est une petite boule de materiel du substrat. Il faut egalement remarquer que la diversite
de taille des femelles indique que leur age etait sans doute different. Dans tous les cas et avec ces
conditions de reproduction en captivite, la valeur moyenne du pourcentage de viabilite des oeufs
(m = 77,53%) est remarquablement elevee. Elle permet de supposer que, dans certaines
populations sauvages elle pourrait etre plus importante, ce qui indique un taux de reproduction
assez eleve.

Source : MNHN, Paris

REPRODUCTION ET DEVELOPPEMENT POST-EMBRYONNAIRE DE LrTHOBIUS PILICORN1S

361

Tableau I. — Dates de capture des diftercntes femelles ayant pondu, nombre d'oeufs, periode de pome, nombre
d'eclosions et taux de survie a I'eclosion (%). m designe les valeurs moyennes.

Table I. Catch dates of females showing egg deposition, number of eggs, egg-laying period, hatching, survival
rate ( %). m = mean data.

Recoltc

Oeufs

Ponte (1992)

Eclosion

%Viabilit

1.X.1990

Juillet

96,67

1.X.1990

Mai-Juillet

70,83

1.X.1990

Mai

88,46

6. IV. 1992

Juin-Aout

28,89

6. IV. 1992

105

Juin-Aout

61.90

6.1V. 1992

105

Juin-Juillet

74,29

6. IV. 1992

Juillet

70,00

6. IV. 1992

Juillet

83,33

6. IV. 1992

Juin-Juillet

42,86

6. IV. 1992

Juin-Aout

65,85

1 1

6. IV. 1 992

Juillet-Aout

83,33

6. IV. 1992

Juin-Juillet

96,87

6. IV. 1992

Juin

100,00

6. IV. 1992

Juillet

71.43

6. IV. 1 992

Juin-Juillet

94,12

6.1 V. 1 992

Juin-Juillet

86,67

6. IV. 1992

Juillet

80.00

6. IV. 1992

Juin

100,00

m = 34, 1 1

m= 24,94

m= 77,53

En diverses occasions, nous avons pu observer la ponte des oeufs par certaines femelles,
ce qui nous a permis d'enregistrer ce processus sur une cassette-video. Le mecanisme observe
est tres semblable a celui decrit par DEMANGE (1956). La femelle soutient l'oeuf avec les valves
anales et les eperons gonopodiaux ; les gonopodes, tres turgescents, donnent a l’oeuf un
mouvement de rotation en lui evitant tout contact avec le sol. Les ongles apicaux des gonopodes
sont utilisees pour arracher les petites particules du sol et les coller a la surface de l'oeuf qui se
trouve totalement impregne dune secretion visqueuse jusqu'a la formation de l’ootheque en
forme de boule de boue. Cette derniere, spherique, est remarquablement differente de celle de
Lithobius piceus gracilitarsis, laquelle est de forme lenticulaire avec un gonflement au centre ; ce
fait indique que la forme de l'ootheque est specifique. La duree de la ponte varie entre une heure
et dcmie et deux heures et se termine quand l'ootheque est abandonnee au sol.

Nous avons isole dans de petites bottes munies du meme substrat les oeufs qui viennent
d'etre pondus, ce qui a permis d’observer et d’enregistrer I’eclosion, au bout de 24 a 32 jours.
Certains de ces oeufs ont ete delicatement deshabilles de l'ootheque protectrice, ce qui a permis,
par transparence, d’observer l'embryon, qui reste d’abord immobile et ne montre quelques
mouvements qu’au moment de la naissance.

L'eclosion debute lorsque se produit la cassure des deux couches qui forment l'oeuf
(Figs. 1 et 2), une couche externe plus epaisse et une autre interne plus fine, a travers une ligne
medio-equatoriale. La tete est la premiere partie du corps qui sort de l'oeuf ; les antennes sont
maintenues entre les forcipules de fagon a rester dirigees vers l’arriere, repliees sur la partie
ventrale du corps. Avec beaucoup d’efforts, la larve commence a extraire les premiers segments
du corps, et des que quelques-unes des premieres paires de pattes sont liberees, le petit Lithobius
les utilise pour liberer le reste du corps beaucoup plus rapidement. La nouvelle larve est tres
active des qu’elle se sent liberee; cependant. dans aucun cas elle ne se nourrit, comme nous
avons pu le constater. La duree du processus de la naissance : cassure de l'oeuf, jusqu'a
1'abandon de celui-ci par la larve, est de 1,5 a 2 heures.

Au bout de deux ou trois jours, la larve subit une premiere mue, qui a ete egalement
enregistree sur cassette-video. Le debut du rejet de l'ancienne cuticule commence avec sa rupture

362

ANTONI SERRA & MARIA CARME MIQUEL

au niveau du sillon frontal de la region antero-dorsale de la tete, immediatement en arriere de
celle-ci et ce sont les premiers articles des antennes qui apparaissent les premiers a 1 exteneur.
L'exuvie se replie progressivement jusqu’a la partie posterieure du corps et, simultanement, tout
le corps sort de celle-ci ; l'extraction des antennes est sans doute la partie plus longue et la plus
laborieuse du processus. A la fin de la mue. qui dure de 60 a 90 minutes, la nouvelle larve
abandonne sa vieille cuticule ; elle y reviendra plus tard pour la devorer. II taut signaler qu a
partir du moment ou la larve sort de sa vieille cuticule, elle passe beaucoup de temps a nettoyer
ses antennes avec les maxilles et en s'aidant des forcipules pour les soutenir ; on remarque
egalement que cette nouvelle larve est tres active et qu’elle se nourrit de petits invertebres tels que
des enchytreides et des collemboles.

Figs 1-2. — 1 : oeuf avec la ligne dc cassure medio-equatoriale. 2 : larve sonant de l'oeuf.
FlCS 1-2. — 1: egg with medio-equatorial broken line. 2: batching larva.

Le nombre de jours qui separe deux mues successives, c'est-a-dire la duree de chacun des
stades larvaires, est donnee ci-apres ; la terminologie utilisee est celle d'ANDERSSON (1976,

1978a) :

Oeuf (24-32j.)

L0 (2-3j.) - LI (4-8j.) - LII (6-8j.) - LIII (39-44j.) - LIV (28-32j.) - PL

Description des larves

A partir des larves obtenues en captivite et dont l'age est connu, on a pu etudier les
variations des principales caracteristiques morphologiques de chacun des stades. Elies sont
exposees dans le Tableau 2; les nombres entre parenthese qui suivent le nombre de paires de
pattes indique les paires de bourgeons pour chaque stade : les valeurs de la longueur du corps
d'une part, de la longueur et de la largeur de la tete d’ autre pan, sont exprimees en millimetres.

Tableau 2. — Principaux caracteres morphologiques des stades larvaires (L0 a LIV) et du premier stade post-larvaire (PL).
TABLE 2. — Main morphological features of the lar\fal stages (L0 to LIV) and the first post-larval stage (PL).

LII

LIII

LIV

Pattes

7(1)

8(2)

10(2)

12(3)

Tergites

1 1

Antennes

7+7

11 + 1 1

14+14

17+17

20+20

23-27

Ocelles

4-5

5-6-7

Cox. forcip.

2+2(3)

3+3

VmH

P.15

Long, corps

2,96-3,63

2,66-2,98

4,13-4,27

4,19-4,62

4,60-4,71

5,54

Long, tete

0,43-0,50

0,39-0,44

0,62-0,63

0.65-0,68

0.74-0,78

0,83

Larg. tete

0,58-0.68

0,50-0,52

0,62-0,65

0,67-0.74

0,74-0,76

0,82

Source :

REPRODUCTION ET DEVELOPPEMENT POST-EMBRYONNAIRE DE LlTHOBlUS PIUCORNIS

363

Dans le Tableau 3 la spinulation des pattes est exposee pour chacun des stades larvaires.
Pour chaque epine sont indiquees sa limite anterieure (premiere paire de pattes ou elle apparait) et
sa limite posterieure (derniere paire de pattes qui la presente), separees par un tiret. Dans le cas
ou l'une ou l’autre de ces limites varie selon les individus, les paires de pattes-limite (qu’elles
soient anterieure ou posterieure), sont separees par une virgule ; quand l'une de ces paires est
plus frequente, elle est indiquee en gras. Dans le premier stade post-larvaire, il y a une grande
variability pour les limites de la majorite des epines, la plus variable etant la DmP qui se trouve
en series continues (7-12, 8-12, 8-14, 7-15, 8-15, 10-15 ) ou qui apparait de fagon irreguliere
sur certaines pattes (6, 8, 10, 11, 12 ; 11, 12 ; 9, 10). On remarque aussi que l'epine VmH,
caractenstique de Lithobius pilicomis, apparait au stade post-larvaire (PL), le premier stade oil
l'°n trouve la paire de pattes 15, chez tous les exemplaires.

Tableau 3. — Spinulations des stades larvaires (LO a LIV) et du premier stade post-larvaire (PL).

Table 3- ~ Diagram of spinulation of the larval stages (LO to LIV) and the first post-larval stage (PL).

LII

LII1

LIV

DmP

8-10

6 ... 15

DpP

4.6,7.8-10

2,3,6-9,12.15

DaF

1-3.4. 5

1-7

1-8

1,4-8 -

2,5.6.7-8,10,11

1.2.3,5-10,1 1,12.13

DpF

7,8.9-10

2,3,4,5-12,14,15

DaT

1-7

1.5-7

1-8

1-9,10

1,2,3,5-7,8,9,10

1,2.4-10,1 1,12

DpT

6,7.8,9,-12,13,14

VmH

Vmtr

13,15

VaP

15/13-15

VmP

1,4,8-10

1,2-12

1-15

VpP

12.14-15

VaF

4,5,6.8,9-12,14,15

VmF

1,2-7

1-8

1.5-10

1-12

1-15

VmT

1-7

1-8

1-10

1-12

1-13,14,15

REFERENCES

ANDERSSON, G.. 1976. — Post-embryonic development of Lithobius forficatus (L.), tChilopoda: Lithobiidae) Em
sc and., 7 : 161-168.

ANDERSSON, G„ 1978a. — An Investigation of the Post-Embryonic Development of the Lithobiidae. Some Introductory
Aspects. Abh. Verh. naturwiss. Ver. Hamburg. (NF) , 21/22 : 63-71.

ANDERSSON, G., 1978b. — Post-embryonic development of Lithobius ervthrocephalus C. L. Koch (Chilopoda-
Lithobiidae). Ent. scand.. 9 : 241-246.

ANDERSSON, G.. 1979. — On the use of larval characters in the classification of Lilhobiomorph Centipedes (Chilopoda.
Lihobiomorpha). In : M. Camatini .Myriapod Biology. London, Academic Press : 73-81.

Andersson. G , 1980. — Post-embryonic development of Lithobius melanops Newport (Chilopoda: Lithobiidae) Em
scand.. 11:225-230.

Andersson, G„ 1981a. — Post-embryonic development and geographical variation in Sweden of Lithobius crassipes
L. Koch (Chilopoda: Lithobiidae). Em. scand.. 12 : 437-445.

Andersson, G., 1981b. — Taxonomical studies on the post-embryonic in Swedish Lithobiomorpha (Chilopoda). Ent.
scand., Suppl. 16 : 105-124.

Andersson, G., 1982a. — Post-embryonic development of Lithobius calcaratus C. L. Koch (Chilopoda: Lithobiidae)
Ent. scand., 13 : 435-440.

Andersson, G., 1982b. — Post-embryonic development of Lithobius microps Meinert (Chilopoda: Lithobiidae) Ent
scand., 13 : 89-95.

Andersson, G.. 1983. — Post-embryonic development of Lithobius curtipes C. L. Koch (Chilopoda: Lithobiidae) Ent
scand., 14 : 387-394.

Andersson, G.. 1984a. — Post-embryonic development of Lithobius tenebrosus fennoscandius Lohmander (Chilopoda-
Lithobiidae). Ent. scand., 15 : 1-7.

364

ANTONI SERRA & MARIA CARME MIQUEL

ANDERSSON. G„ 1984b. — Posi-embryonic developmenl of Lamyctes fulvicornis Meineri (Chilopoda: Hemcopidae).
Ent. scand., 15 : 9-14.

Andersson, G„ 1990. — About the duration of the different stadia in the post-embryonic development of some
Lithobiomorph species, hi : A. MlNELLI. Proceedings 7th Intern. Congr. Myriapodology. Leiden, Brill : 323-335.
Brolemann, H. W„ 1930. — Elements d'une Faune des Myriapodes de France. Chilopodes. [Faune Fr.. 25 ]. Pans, P.
Lechevalier : 1-405.

Demange. J. M., 1956. — Contribution a l'etude dc la biologie, en captivite, de Lithobius piceus gracilitarsis Brol.

(Myriapode-Chilopode). Bull. Mus. natl. Hist, nat., 2e ser., 28 : 388-393.

Eason, E. H., 1964. — Centipedes of the British Isles. London, F. Wame & Co Ltd, 294 pp.

Eason, E. H.. 1970. — The Chilopoda and Diplopoda of Iceland. Ent. Scand., 1 : 47-54.

Murakami, Y., 1958. — The life-history of Bothropolys asperatus (L. Koch). Zool. Mag. Tokyo , 67 : 217-223.
Murakami, Y., 1960. — Postembryonic development of the common Myriapoda of Japan. V: Lithobius pachypedatus
Takakuwa (Chilopoda. Lithobiidae). 3. Variation in the number of articles of antennae and coxal pores. Zool. Mag.
Tokyo. 69 : 167-170.

Scheffel, 1969. — Untersuchungen iiber die hormonale Regulation von Hautung und Anamorphose von Lithobius
forficatus L. (Myriapoda, Chilopoda). Zool. Jb. Physiol., 74 : 436-505.

Verhoeff, K. W., 1905. — Uber die Entvvicklungsstufen der Steinlaufer, Lithobiiden, und Beitrage zur Kenntms der
Chilopoden. Zool. Jb., Suppl. 8 : 195-298.

Source : MNHN, Paris

Developpement post-embryonnaire et cycle biologique
de Eupolybothrus elongatus (Newport) dans Test

algerien

Tarek DaaS *, Noureddine BOUZERNA * & Michel DESCAMPS **

* Institut des Sciences de la Nature, Laboratoire de Biologic Animale
Universite de Annaba, BP 12 Annaba, Algerie

** Ecophysiologie d’Invertebres du Sol, Laboratoire de Biologie Animale, Universite de Lille I

F-59655 Villeneuve d'Ascq Cedex, France
(to whom all correspondence must be sent)

RESUME

Le cycle de developpement du Chilopode Eupolybothrus elongatus a ete suivi dans la region d’Annaba, a l’Est de
1 Alg6rie. Les caracteristiques des differents stades ont ete d£crites (nombre de paires de panes, de segments antennaires.
longueur et, pour les stades epimorphes, masse moyenne). Labondance relative des differents stades a ete evaluee en
utilisant des pi£ges. Les observations sur le terrain, sur deux sites h couverture veg&ale differente (Sidi Amar : foret
d 'Eucalyptus ; Oued Zied : steppe de type mediterraneen), ont permis de reconnaitre deux periodes de ponte preferentielles,
d octobre a janvier, et, dans une moindre mesure, d'avril k juin-juillet. Le travail, mene en parallele sur les deux zones
delude, a permis de mettre en Evidence l'influence du milieu. Les resultats montrent I’importance des facteurs extemes sur
le taux de capture : les mois les plus secs sont caracterises par l'absence (ou la quasi-absence) d’adultes dans les pieges,
meme dans le biotope le moins aride.

ABSTRACT

Post-embryonic development and life-cycle of Eupolybothrus elongatus (Newport) in Eastern
Algeria.

The biological cycle ot the chilopod Eupolybothrus elongatus has been studied in Eastern Algeria, near Annaba. The
characteristics ot the different stages are described (number of leg pairs and of antennal segments, length of animal and,
for the epimorphic stages, mean mass). The relative abundance in the field of the different stages has been estimated by
using pitfall traps. The results concerning two sample sites with different kinds of vegetation (Sidi Amar: Eucalyptus
forest; Oued Zied: Mediterranean steppe) show two egg-laying periods, from October to January, and at a lesser extend,
from April to June-July. Analysis of the results from the two sample sites points out the influence of rainy periods: the
driest months are characterized by no adults (or so) in traps, even for the less arid biotope.

Daas, T., Bouzerna, N. & Descamps, M., 1996. — Developpement post-embryonnaire et cycle biologique de
Eupolybothrus elongatus (Newport) dans lest algerien. In: Geoffroy, J.-J., Mauries. J.-P. & NGUYEN DUY -
JACQUEMIN, M., (eds). Acta Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 365-370. Paris ISBN : 2-85653-502-X.

366

TAREK DAAS. NOUREDDINE BOUZERNA & MICHEL DESCAMPS

INTRODUCTION

Chez les chilopodes, la plupart des recherches en physiologie experimentale ont ete menees
chez Li thobius fotficatus (L.) (cf. les raises au point de SCHEFFEL, 1987 ; JOLY & DESCAMPS,
1988 : DESCAMPS, 1992). Seuls quelques travaux ont ete menes sur cl autres especes
(Scolopendra cingulata : JOLY, 1966 : L. crassipes : BENIOURI el al 1983).

En ce qui concerne le cycle biologique. la succession des stades et la croissance ont etc
bien etudiees chez differents Lithobius, en particular, pour citer les travaux les plus recents, par
ANDERSSON (1976-1984). Le genre “ Bothropolys " n'a ete etudie que par MURAKAMI (#.
asperatus , 1958). Le chilopode le plus commun dans la region d’Annaba est Eupolybothrus
eloneatus (Newport, 1849), ex-Bothropolys elongatus auct. (cf. EASON, 1972 ; MATIC, 1974).
Avant d'entreprendre une etude experimentale sur ce materiel, il nous est apparu necessane den
connaitre le cycle biologique dans son environnement mediterraneen.

MATERIEL ET METHODES

Let animaux recoltes dans divers biotopes aulour d'Annaba. soni maintenus dans des boites en plastique ou se trouve de
la terre humidifiee reconvene dun papier nitre imbibe d'eau. Les animaux son. nourris regul.erement avec des moustiques,
des mouches ou de petites araignees. La temperature d'elevage a vane en cours d annee entre 10 C (janvicr) et 26.5 C
(aout) refletant en cela les variations de temperature externc (de 10°C &'29,2CC).

Des pontes ont ete obtenues au laboratoire. qui nous ont permis d’6tudier les caractenstiques des premiers stades larvaires.
Les stades les plus ages ont ete suivis a partir d’animaux recoils dans la nature. Les resultats exprimes dans les tableaux
coiTespondent a un minimum de 4 observations.

Recoltes sur le terrain , ... , , i

Deux stations ont 6t€ etudiees : Sidi Amar (foret d' Eucalyptus) et Oued Zied (steppe de type mediterraneen). Les pifcges
d’ interception (pitfall trap), garnis d'eau formolde. sont releves au bout de 4 jours. Quatre prdlevements par mots ont ete
fails au cours de la periode d'etude (novembre 1990 - octobre 1991).

RESULTATS

Developpement post-embryonnaire

Nous avons pu reconnaitre 5 stades larvaires (L0 a L4 suivant la nomenclature de
ANDERSSON 1978), dont les durees de developpement sont consignees dans le Tableau 1 et les
caracteristiques dans le Tableau 2. A partir du stade PL4, les durees d'intermue deviennent tres
longues (45 jours et plus).

Tableau I. — Duree des stades. de la ponte a la larve 4. chez E. elongatus (d'apres des observations lattes entre octobre

1990 et mai 1991). . . ,

Table /. — Post-embryonic stadia of E. elongatus, duration in days, from egg to post-larval stage 4 ( after observations

made between October, 1990 and May, 1991).

Stades

Duree (en jours)

oeuf

21 a 25

Larve 0

13 * 22

Larve 1

Larve 2

Larve 3

10 a 13

Larve 4

Post-larve 1

13 a 17

Post-larve 2

Post-larve 3

30 h 36

Post-larve 4

Source :

DEVELOPPEMENT ET CYCLE BIOLOGIQUE D’UN L1THOBIOMORPHE ALGER FEN

367

Tableau 2. — Caractdristiques des stades larvaires et posi-Iarvaires chez E. elongatus
Table 2. — Morphological features of larval and post-larval stages of E, elongalus.

Stades

Nb.de paires
de panes

Nb. d'articles

antennaires

Longueur

(mm)

Masse (mg)

Larve 0

5 & 6

Larve 1

11 a 13

6,5

Larve 2

Larve 3

1 I

7,5

Larve 4

13 * 14

PL 1

34 h 36

9 & 1 1

4,1 a 6,3

PL 2

13 a 15

11,5 a 20

PL 3

38 a 39

16 & 18

23,5 a 40

PL 4

39 h 40

45 a 62

52 a 75

PL 5

21 & 23

75 a 85

80 a 115

PL 6

41 h 42

27 ii 30

93 a 140

120 a 165

> k PL 6

42 a 43

33 h 45

156 a 200

1 80 a 260

femelles

males

Cycle biologique

Les resultats des recoltes sont consignes dans les Tableaux 3 (Sidi Amar) et 4 (Oued Zied).
bn ce qui concerne Sidi Amar, nous pouvons constater un maximum de stades larvaires pieges
au cours du mois de decembre. Compte tenu de la duree de 1'incubation et des stades larvaires
obtenus au laboratoire, nous pouvons dire que les pontes ont lieu preferentiellement entre
octobre et janvier. Une autre periode de ponte, moins favorable semble-t-il d'apres le nombre de
captures, intervient entre avril et juin-juillet. Le maximum de capture des stades post-larvaires est
lui, atteint au mois de janvier. Notons un minimum dans les recoltes d'adultes (PL6 et au-
dessus) en aout et septembre.

Le site de Oued Zied est beaucoup plus pauvre en faune, mais on peut observer a peu pres
les memes phenomenes qu a Sidi Amar, avec cependant une periode d'absence de grands stades
post-larvaires durant la periode mai-aout (plus octobre).

DISCUSSION ET CONCLUSION

Le premier point que nous voudrions discuter conceme les caracteristiques du stade LO : la
duree de ce stade au laboratoire nous a semble anormalement longue. En effet, les differents
travaux faisant etat de la duree des premiers stades larvaires indiquent que ceux-ci n'excedent
jamais quelques jours (1 a 3 en general). Nos observations ont porte sur 9 individus, issus dune
ponte de 15 oeufs. Si les caracteristiques morphologiques permettent de separer nettement les
stades LO et LI, ll n'en reste pas moins etonnant qu'il y ait une aussi grande difference de duree
entre ces deux stades successifs. II est done possible que nous ayons observe, par une
malneureuse coincidence, des larves au developpement perturbe par une cause inconnue. La
reponse ne pourra bien sur etre apportee qu'en etudiant un plus grand nombre d'individus
provenant de femelles prelevees dans differents lieux de recolte.

En ce qui concernant les autres stades, nous sommes en presence du developpement
classique d un chilopode lithobiomorphe.

368

TAREK DAAS, NOUREDDINE BOUZERNA & MICHEL DESCAMPS

Tableau 3. — Recoltes sur le site de Sidi Amar.
Table 3. — Number of individuals collected at Sidi Amar.

Nov.

D<5c.

Jan.

Fev.

Mars

Avr.

Mai

Juin

Juil.

Aout

Sept.

Oct.

Larve 1

Larve 2

Larve 3

Larve 4

PL 1

PL 2

PL 3

1 1

PL 4 & 5

PL 6

1 1

> & PL 6

Total larves

1 1

Total PL U PL 5

1 1

Total matures (> PL 5)

1 1

Total general

Tableau 4. — R6coltes sur le site de Oued Zied.
TABLE 4. — Number of individuals collected at Oued Zied.

Nov.

D6c.

Jan.

Fev.

Mars

Avr.

Mai

Juin

Juil.

Aout

Sept.

Oct.

Larve 1

Larve 2

Larve 3

Larve 4

PL 1

PL 2

PL 3

PL 4 & 5

PL 6

> a PL 6

Total larves

Total PL U PL 5

Total matures (> & PL 5)

Total general

1 1

Source : MNHN, Paris

DEVELOPPEMF.NT ET CYCLE BIOLOGIQUE D'UN LITHOBIOMORPHE ALGERIEN

369

Le second point de discussion concerne la methodologie utilisee lors des recoltes : en effet,
le piegeage d' interception (pitfall trap) est peu propice a la recolte des stades jeunes (voir les
articles de Branquart & GASPAR et de GEOFFROY & CELERIER dans ce volume) ;
neanmoins, s’agissant de l’etude d’une seule et meme population specifique, et notre but etant la
recherche du deroulement d’un cycle et non l’estimation correcte dune densite de population, les
resultats restent significatifs.

Enfin, ce qui nous semble important au point de vue du cycle biologique, est pour le site
de Oued Zied, 1'absence ou la quasi absence d'individus dont les caracteristiques
morphologiques correspondraient a des stades posterieurs a PL6. Comme il est statistiquement
improbable que nous n'ayons recolte que de jeunes individus, il nous faut plutot conclure a une
population de taille moyenne plus faible, en rapport avec une nourriture disponible plus faible
(aridite du milieu).

Le rythme saisonnier, caracterise par 1'absence de grands froids, par des periodes de
grande chaleur (juillet et aout), par la secheresse, peut expliquer, avec un decalage dans le temps
que I’on pourrait qualifier d’“inertie du systeme”, 1'absence ou la diminution de l’activite-densite
des populations durant les periodes les plus chaudes et les plus arides. Inversement, c'est durant
les periodes les plus humides que Ton obtient les maximums de recolte.

Les populations de E. elongatus sont done fortement infeodees aux conditions climatiques,
et l'on peut d'ores et deja conclure que comme L. forficatus (DESCAMPS, 1971 ; HERBAUT,
1975), cet animal possede une physiologie qui est fortement sous la dependance des facteurs
extemes, en particulier de la temperature.

REFERENCES

ANDERSSON, G., 1976. — Posi-embryonic development of Lithobius forficatus (L.), (Chilopoda : Lithobiidae). Ent.
scand., 1 : 161-168.

ANDERSSON, G., 1978. — An investigation of the post-embryonic development of the Lithobiidae - Some introductory
aspects. Abh. Verb, naturwiss. Ver. Hamburg . 21/22 : 63-71.

ANDERSSON, G., 1981. — Post-embryonic development and geographical variation in Sweden of Lithobius crassipes L.
Koch (Chilopoda : Lithobiidae). Ent. scand. , 12 : 437-445.

ANDERSSON, G., 1982. — Post-embryonic development of Lithobius microps Meinert (Chilopoda : Lithobiidae). Ent.
scand., 13 : 89-95.

ANDERSSON, G., 1983. — Post-embryonic development of Lithobius curtipes C. L. Koch (Chilopoda : Lithobiidae). Ent.
scand., 14 : 387-394.

ANDERSSON, G., 1984. — Post-embryonic development of Lithobius tenebrosus fennoscandius Lohmander (Chilopoda :
Lithobiidae). Ent. scand., 15 : 1-7.

Beniouri, R., Descamps. M., Porcheron. P. & Joly, R.. 1983. — Correlations naturelles et experimentales entre
croissance spermatocytaire et taux d'ecdysteroi'des chez les Lithobiidae (Chilopoda). Rev. Can. Biol. Experiment..
42 : 183-189.

Branquart, E. & Gaspar, C., in press. — Comparative study of sampling techniques of saprophagous macroarthropods
(Diplopoda and lsopoda).

Descamps, M., 1971. — Le cycle spermatogenetique chez Lithobius forficatus (L.) (Myriapode, Chilopode). 11. Influence
des facteurs externes sur 1'evolution du testicule et des vesicules s£minales. Arch. Zool. exp. gen., 112 : 731-746.
Descamps, M., 1992. — Endocrine events during the life cycle of Lithobius forficatus (L.). Ber. nat.-med. Verein
Innsbruck , suppl 10 : 11-116.

Eason, E. A., 1972. — The type specimens and identity of the species described in the genus Lithobius by George
Newport in 1844, 1845 and 1849. Bull. Br. Mus. nat. Hist. (Zool.), 21 : 297-311.

HERBAUT, C., 1975. — Influence des facteurs externes sur le cycle ovogtSnetique chez Lithobius forficatus (L.)

(Myriapode Chilopode). Arch. Zool. exp. gen.. 116 : 293-302.

Joly, R., 1966. — Contribution a I'etude du cycle de mue et de son determinisme chez les Myriapodes Chilopodes. Bull.
Biol. Fr. Belg.,3 : 379-480.

Joly, R., Descamps, M., 1988. — Endocrinology of Myriapods. In : H. Laufer & R. G. H. Downer, Endocrinology of
Selected Invertebrate Types . New York, Alan R. Liss : 429-449.

370

TAREK DAAS, NOUREDDINE BOUZERNA & MICHEL DESCAMPS

Matic, Z., 1974. — Contribution h la connaissance du genre Bothropolys Wood, 1863 (Lithobiomorpha, Ethopolidae).
Ann. Zool., Warsawa, 31 : 329-341.

Murakami, Y., 1958. — The life-history of Bothropolys asperatus (L. Koch). Zool Mag. Tokyo , 67 : 217-223.
SCHEFFEL, H.. 1987. — Hautungsphysiologie der Chilopoden : Ergebnisse von Untersuchungen am Lithobius forficatus
(L.). Zool. Jb. Physiol., 91 : 257-282.

Source : MNHN, Paris

The Segmentation of the Head and Anterior Trunk of
Millipedes (Diplopoda) - A Reassessment

Wolfgang DOHLE

Institut fur Zoologie, Konigin-Luise-Str. 1-3, D-1000 Berlin, 33, Germany

ABSTRACT.

In recent years, the segmentation of the head and trunk of arthropods has gained new interest through the revelation
and characterization of segment polarity genes and pair rule genes. Are there indications of a general double-segment
organization in millipedes? A reinvestigation of germ band formation in the millipede Glomeris marginata was
performed with fluorescent dyes and with SEM. Most results which had been gained by traditional histological methods
have been confirmed. The intercalary segment remains without appendage buds. The mandibles become subdivided into 2
articles. The gnathochilarium is formed by the first maxillae and the triangular sternite of this segment. There is no
indication of appendage buds on the postmaxillary segment. The first 4 trunk segments are simple segments with one
pair of groove-like invaginations for the formation of the ganglia. Only in the subsequent segments (V+VI. VII+VIII)
the lateral and dorsal parts combine to form doubie-pleurites and double-tergites.

RESUME

La segmentation de la tete et de la partie anterieure du tronc chez les diplopodes - une
reevaluation.

Durant ces dernieres ann6es, 1'etude de la segmentation de la tete et du tronc des Arthropodes a connu un regain d’interet
grace a la decouverte et a la caracterisation de genes gouvernant la polarite de la segmentation et determinant
1 association des segments par paires. Y-a-t-il alors des informations relatives a une organisation generate d'une double
segmentation chez les diplopodes? Une reevaluation de la formation de la bandc germinale chez le diplopodc Glomeris
marginata a ete eftectuee dans ce but a 1 ' aide de coloration en fluorescence et en microscopie electronique a balayage
(MEB). La plupart des resultats qui avaient ete acquis grace aux techniques histologiques traditionnelles ont ete
confirmes. Le segment intercalate demeure depourvu de bourgeons. Les mandibules commencent a se subdiviser en deux
articles. Le gnathochilarium est forme par la premiere paire de maxilles et le sternite triangulate correspondant. Nous
n avons aucune confirmation concernant la presence de bourgeons appendiculaires sur le segment post-maxillaire. Les
quatre premiers segments du tronc sont simples et presentent une paire d’invaginations en forme de rainure qui conduisent
& la formation de ganglions. Ce n’est que dans les segments suivants (V+VI. VII+VIII) que les parties laterales et dorsales
sc combi nent pour former des doubie-pleurites et des double-tergites.

Dohle, W.. 1996. — The segmentation of the head and anterior trunk of millipedes (Diplopoda) - A reassessment.
In: Geoffroy, J.-J., Mauries. J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus . natn.
Hist, nat .. 169 : 371. Paris ISBN : 2-85653-502-X.

Source : MNHN, Paris

On Periodomorphosis, Iteroparity and Life-Cycles in
Males and Females of Tachypodoiulus niger (Leach)
(Myriapoda, Diplopoda, Julidae) in France, Germany

and Great-Britain

Frangois Sahli

Museum National d'Histoire Naturelle, Zoologie/Arthropodes, 61 ruc Buffon, F-75231 Paris Cedex 05
& Laboratoire souterrain du CNRS, F-09200 Moulis, France

ABSTRACT

In spite of appearances, the cycles and life-history of Tachypodoiulus niger have been hitherto poorly known. In the
same way as he did previously in Ommatoiulus sabulosus. the author describes and interprets the cycles of
Tachypodoiulus niger , particularly according to VERHOEFF's investigations in Germany, those of Fairhurst in Great-
Britain, and to the author’s 38 years of mostly unpublished data and experiments on this species in France, Germany and
Great-Britain, and in the light of recent knowledge.

RESUME

Sur la periodomorphose, I'iteroparite et les cycles de vie des males et des femelles de
Tachypodoiulus niger (Leach) ( Myriapoda , Diplopoda , Julidae ) en France, Allemagne et Grande-
Bretagne.

En depit des apparences, les cycles de Tachypodoiulus niger sont mal connus. Comme il Fa fail anterieurement pour
Ommatoiulus sabulosus , 1'auteur decrit et interprete les cycles de Tachypodoiulus niger. Pour ce faire, il s'appuie sur les
investigations de Verhoeff en Allemagne, de Fairhurst en Grande-Bretagne. ainsi que sur des donnees et experiences -
pour la plupart inediles - de 38 annees de recherches personnelles sur cette espece en France, en Allemagne et en Grande-
Bretagne. Les interpretations ont, de plus, ete faites & la lumiere de nos connaissances actuelles sur la periodomorphose.

INTRODUCTION

Our current knowledge of the cycles in T. niger mainly goes back to the data of
VERHOEFF (1915-1934: particularly 1928, 1932) and Sahli (1966). In collaboration with J. G.
BLOWER, Fairhurst (1968) translated the data of his predecessors into English, mentionned
those of HALKKA (1958) and added his own worthwhile observations.

[In reading SAHLI’s thesis (1966) the British authors did not understand that in Tachypodoiulus (and
Ommatoiulus) it is possible for a specialist - taking attention to details - to distinguish an intercalary from a
juvenile male (cf. SAHLI, 1966): only extremely rare cases (say in the region of 1: 500 intercalates) may
constitute exceptions].

Sahli. F., 1996. — On periodomorphosis, iteroparity and life-cycles in males and females of Tachypodoiulus
niger (Leach) (Myriapoda, Diplopoda, Julidae) in France. Germany and Great-Britain. / n: Geoffroy, J.-J.. Mauries,
J.-P. & Nguyen Duy - JACQUEMIN, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat .. 169 : 373-384. Paris
ISBN : 2-85653-502-X.

374

FRANCOIS SAHLI

New investigations were made by the present author from 1966 to 1992, in Burgundy, the
French Pyrenees (Ariege), French Alps, Germany and Great Britain. On the one side, these
researches and on the other the reinterpretation of VERHOEFF's data and my own prior to 1966,
showed that the cycles of T. niger were still poorly understood and much more complicated than
myriapdologists thought.

[Concerning cycles, periodomorphosis, adult to adult moults and combined strategies, HOPKIN & READ
(1992) partly overlook or misinterpret some recent results of the present author, concerning, among others,
Tachypodoiulus sp.. Ommatoiulus sp., Allajulus nitidus and Blaniulus guttulatus].

This paper aims to set the record straight regarding our knowledge of cycles in
Tachypodoiulus niger.

ADOPTED NOMENCLATURE

The definitions and abbreviations used here have been given in several previous papers (Sahli 1990a, b, 1991a,
b, c) to which the reader is referred.

Concerning the seriations, from reasons of symmetry, and taking into account the intercalary appearance
season, the same numbers as those adopted for O. sabulosus (i.e. seriation 1: intercalates si in autumn of the year x;
seriation 2: intercalates si in spring of the year x+1) will be used for T. niger (Table 3). We will add a seriation 3 and 3’
specific to Tachypodoiulus .

RESULTS

My own cultures, experimental investigations, field observations, along with those of
VERHOEFF (passim) led to the results recorded in Tables 1-8 and in Figures 1 and 2.

At low altitudes, adl production (maturation moults of juvenile to first adult males adl =
MMJ) can take place (see, among others. Table 7) in the following ways:

- (case a) at the end of the winter/spring of the year x (before mating and egg deposition),

- (case b) at the summer/autumn of the year x (i.e. several months before egg-laying in
spring of the year x+1, as it is the case in Allajulus nitidus). Nevertheless, mating can
additionally take place in autumn of the year x,

- both cases (a and b) in the same year.

In other words, depending on the environmental conditions, T. niger is able to use two
strategies of adl male production (either spring or autumn), or a combination of the two (spring
+ autumn).

Results concerning seasonal cycles in T. niger are recorded in Table 4. This gives the
“base cycle” which includes all fundamental possibilities. From the 8 basic possibilities (a to h)
one can derive all the possible ways, should one wish to go into details and individual cases.
From the base cycle all the possible patterns can be reconstituted.

Base cycles have been given for O. sabulosus in Burgundy (SAHLI, 1990a , Fig. 1) and
for Mediterranean populations (SAHLI, 1991a, b, Table 1). [in Sahli (1991, table l) a forgotten vertical
line should connect the adl (in case b. seriation 1 ”p) and the si (in case a, seriation lp) in summer. In the same table,
the two vertical unbroken lines must be regarded as two vertical braces].

INTERPRETATION AND DISCUSSION

As in O. sabulosus, short standing (SL) and long-standing (LL) intercalaries exist in
T. niger (SAHLI, 1990b).

Results taking into account the author’s experimental data and field observations, as well
as VERHOEFF's investigations are recorded in Tables 1 & 2. In the present state of our
knowledge, two cases can be distinguished in Germany and Burgundy up to 1000 m altitude:
one in lowlands and hills, another in the Allgau mountains (800 m) and in a rock shelter in the
Saarland, both in Germany.

As shown in Tables 1 & 2, in the sites under 1000 m, there are two typical possibilities, at
least in the present state of our knowledge:

Source :

PERIODOMORPHOSIS. ITEROPARITY AND LIFE-CYCLES IN A JULIDAE

375

Table I. — The two possible cases in T. niger, at an altitude under 1050 m in some German regions (particularly in
Saarland) and in Burgundy. SUM = summer.

Egg deposition

adl production

sch cf production (ad-schcf)

ad2 production (schcf-ad2)

- ad2 of SL origin

- ad2 of LL origin

SPRING

few

SUM-AUTUMN

and/or +

Case b

SPRING

SUM-AUTUMN

egg deposition

+(?)

typical adl poduction

typical schd* production

Table 2. — T. niger in Burgundy and some German regions. A: subdivision of adl. B: typical appearance of sch cf . In
case b\ adl mating is also possible -(first-) in autumn of year x.

case a : spring adl (reproduction = year x)

case b : summer/autumn adl (= reproduction year x)

case b’ : summer/autumn adl (= main reproduction year x+1 )

Spring

Sum/Autumn

Spring

Sum/Autumn

case a : schcf from adl of case a
case b : schcf from adl of case b

- case a = in Burgundy (at the numerous sites studied) and in epigean animals from
Germany (Saarland. Hunsriick, Eifel, Rheinland, Taunus and Hessen). In this case: (al) egg
laying takes place in spring; (a2) adl production in spring and/or autumn; (a3) typical - or
majority - schd" production in summer/autumn. Schd" are mainly (or only?) made up of LL and
R individuals. The predominance of LL in T. niger contrasts with that of SL in Mediterranean
O. sabulosus and probably in O. moreleti in Australia and southern Portugal (BAKER, 1978,
1984).

- case b = in Germany in the Allgau (750-1050 m), at least according to the culture results
of VERHOEFF (1934), and in a rock shelter at Wadern in Saarland. In this case : (bl) egg
laying(?) and adl male production take place at the end of summer/onset of autumn; (b2) schd1
are produced in spring. In VERHOEFF's cultures (from 1924 to 1928) only scho" SL (thus an a
succession) were observed. (The question of whether this was the result of the culture
conditions remains open). A complete absence of LL scho" in the Allgau mountains would be
astonishing. In Saar cultures (in a unheated basement - with open windows - of the Saarland
University) of individuals collected in the Wadern rock shelter, the following succession has
been observed: ad (collected in autumn x) - s (spring x+1) - ss (autumn x+1 ). A succession |3
(with R scho’) as been obtained instead of the a one (ad — s — ad) of VERHOEFF.

In case a, some schd" (ad- schd") may sometimes be (exceptionally?) produced in spring in
Burgundy, as well as in Saarland. If this occur, such scho" (which would be SL ones) could be
regarded as minority schd" in comparison with the whole scho" liable to be collected in spring.

Like his predecessors, FAIRHURST(1968, 1974) was unaware of the existence of LL
schd". He also did not know about the possibility for adults maturing in autumn to stay adults
without moulting until the following spring. He trusted the informations given by VERHOEFF

376

FRANCOIS SAHLI

(passim) and HALKKA ( 1958). For these reasons, it seems that FAIRHURST regarded - at least
implicitly - the spring postadult male production (e.g. the passage from males become SL
intercalaries 1 in autumn into ad2 males in the following spring) as common, even exclusive
(based on the last row of ocelli appeared in animals preserved in alcohol). In the same way
FAIRHURST regarded implicitly the ad-scho"moult as common in spring (schef being produced
in spring from adults which become adults in the previous autumn).

Table 3. — Numeration adopted for the seriations in Burgundy and some regions of Germany. Sedations 1 and 2
homologous to those I and 2 of T. niger - exist in O. sabulosus.

spring (year x) adl do' (low altitudes: Burgundy. Germany)

SERIATION 1: si intercalaries in summer/autumn (year x) (cf O. sabulosus) _

summer (year x) adl dd (e.g. in the Allgau mountains)
reproduction period = year x

SERIATION 2: si intercalaries in spring x+1 _ _ _

summer/autumn adl dd (year x) (in Germany plains)

typical reproduction period = year x+1

SERIATIONS 3 & 3': si intercalaries = at the end of spring or sum/autum year x+ 1 |

Table 4. - Basic cycle of T. niger in Burgundy. Great-Bri.am and some German regions.A.jartingfromadl,
becoming adults in spring (SP). B and C: starting from adl. becoming adults in summer (SUM) /autumn (Al l).
SER.: seriation. Unbroken line means a moult. Doited line means no moult.

CASE SER

SP x

SUM-AUT x

SP x+1

SUM-AUT x+1

1 SL

ad2

adl -

ad2

1 LLa

cl .

1 i i i\

adl -

\ -

- ss

r 1

adl

2 SL a

? i I ft

ad I - -

adl -

Si -

- ss

1 e 1

ad2

acJl .

- - ss

3’

adl .

?sl

In the present state of our knowledge in Burgundy and in Germany, SL successions never
appear as exclusive or even common. From numerous cultured animals in different conditions
and during several years, I never obtained spring intercalaries from epigeous adult males (say
adl or presumed ad2) collected in the previous autumn. I obtained spring intercalaries only from
adults collected in the rock shelter at Wadern in autumn (case b). Conversely, scho' collected in
autumn, as well as as scho” (particularly si ones) obtained in cultures in summer/autumn, never
gave postimaginal adults the following spring in cultures. Only autumn scho1 from the Wadern s

rock shelter gave spring postadult males (case b).

It is possible that spring SL (case b) exist in a more or less high numbers in Great-Britain.
Nevertheless, according to my own investigations made at Milldale (G.B.) in 1981 and 1483,
the cycle seems to include fundamentally schd" LL (case a) as in Burgundy and Saarland.
FAIRHURST's observations in Britain require confirmation: the case in Britain needs more

Source : MNHN, Paris

PERIODOMORPHOSiS. ITEROPARITY AND LIFE-CYCLES IN A JULtDAE

377

In^rnimn ^ ons particular|y using cultures. Freshly moulted spring schtf might be ss ones (schtf
in autumn of the year y giving ss in spring y+1?).

„ j c,1" b°th,pases,(a and b) °"e can admit the possibility of a double production of LL schtf
u's‘ 1 ’ the number of the SL sch o’ being variable. For instance, the SL may be
either few or absent (case a) or more frequent (case b) as in the Allgau mountains (if appearances

197?SfannadH hfeai bl^llgau resuIts’ VERHOEFF (1923, 1925) obtained SL in

adult on 15 9 1922) d ^ ^ 4' 922 Wh‘Ch tUmed im° 3 SL °n ’2‘ 6' 1922 and then int0 an

Fig. 1. Cycle ol males in T. niger in Germany and
Burgundy. The figure takes into account
possibilities of both LL and SL intercalates.
I: males which became adl in spring of year x
(Px). II: males which became adl in
summer/autumn x (Ex). In I and II it is
arbitrarily assumed that the number of spring
adl (adl = solid rectangles with an open circle;
ad 2 = entirely solid rectangles) in year x is
equal to the number of summer/autumn adl of
year x (under this assumption the solid
rectangles in PX and EX are of equal
dimensions). Ill and IV: new generation (of
year x+1). In III and IV it is arbitrarily
assumed that the number of spring adl (III) is
much higher than the number of

summer/autumn adl (IV) (under this
assumption the solid rectangles with an open
circle are of different dimensions). Schc? are
represented by open rectangles. Dashed lines:
without moult or transformation; solid lines:
transformation after a moult. - m: death, ss:
double intercalary form. Asterisk: schcf at the
end of the spring/beginning of summer (after
VERHOEFF, 1923).

m T'™1 that tW0 strateSies can be used by Mediterranean O. sabulosus: type 1 =
i 1MJ anhe end ot winter/onset of spring, i.e. a long time before egg-laying in summer/autumn;

r fnm Cnd of fP^g/^mmer of the year x, just before mating and egg deposition
(Sahli, 986, 1991a, b. c, 1992). Nevertheless, in O. sabulosus MMJ of type 1 or 2 (or both)
and egg-laying take place the same year. The respective strategies used by T. niger (at low
altitudes in Germany and in Burgundy) and O. sabulosus in the South-East of France are not
strictly superimposable: this is the reason why we speak of cases a and b in T. niger and tvpes 1
and 2 in O. sabulosus.

In the Departement des Hautes-Alpes (1000 - 1600 m) the production of ad. and schd"
ur!nSpC° CCted under snow bridges or stones) might take place as it does in the case a (SAHLI
1770 & unpubl.), at least in the present state of the investigations. In the Pyrenees ariegeoises,
the production of ad-scho' might correspond to case b. schef being numerous in autumn
according to preliminary observations. A thorough study, over numerous years, needs to be
carried out in the Hautes-Alpes, in the Pyrenees ariegeoises and in the Hautes-Pyrenees (in
which, on the slopes of the Pic de Campbielh, schefas well as ad males are frequent under
stones at the end of the summer).

Although simplified, Tables 5 & 6 allow the interpretation of virtually all the innumerable
field and culture data accumulated, be it in spring (year x+2 in Table 5) or in summer/autumn
(year x+1, Table 6).

378

FRANCOIS SAHL1

In sorin'3 (x+2) the schd" number will depend of the number of adl of year x+1, which
were prcdS in spring x+1. If one supposes, for instance tha, these adl were numerous, the
supposed situation will lead to numerous LL. shown in bold type in Table 5.

Year x+2 Will include on .he one hand new adl (year x+2). assumed (arb.irar.ly) to bepredomman.andonthe
taken into account, aut: autumn; n: new; sp: spring.

spx

aut x

schcf (SL)

sp x+1

aut x+1

sp x+2

some fast ad2

(x)

cO .

(x)

some fast adz -

— ss

(X)

adl

schcf (LL)

schcf (LL) - ad2

adl n

slow ad2 . (X)

new adl

(x+2)

“old” adl

(x+1)

some fast ad2

(x+1)

schcf (LL)

(X+1)

TABLH 6 _ T niger in Saarland and Burgundy. Starting from spring of year x+1 . The following autuntn.1 slow ad2 (bold,
' '■ re assUmed (arbitrarily) to be well represented. The number of new autumn (x+1) adl is variable compared w h
the number of^l (autumn year x+1), it can be higher (in this case adl would be in bold letters) or lower (si would

be in bold).

spring x+1

autumn x+1

some fast ad2

(X)

schcf (LL)

slow ad2

new autumn adl

(X)

(x+1)

new spring adl

s 1

(x+1)

As a general rule in spring and in case a, the ratio adl/schcf in spring x+2 (Table 5,

F'c l,(aTthed number of “new” adl (i.e. the number of “new” freshly MMJ) in year x+2, plus
the number of possible “old” adl from autumn x+1 (MMJ in autumn x+1) which have lemained
winter adl without moulting. These new adl can be high, middle or low in n^erp ,

(b) the number of LL si (which became si in autumn x+1). LLsl in turn depend on the
number of adl in spring x+1. A few SL si (? in Fig.l) might add to the LL si, as well as some

S llC The number of adl - new ones in spring x+2 and/or old ones from autumn x+1 (i.e. adl
which remain adults without moulting in spring x+2) - often outnumber, more or less, the

number of LLsl or the number of all schcf (LL si +? SL si + ss). ■

It is worth noting that in spring x+2 one does not compare males from the same ye<m
fact one compares adl and LL si which became, respectively, adults and intercalaries in two
different years.

Source : MNHN. Paris

PERIODOMORPHOSIS, ITEROPARITY AND LIFE-CYCLES IN A JULIDAE

379

Even adl (e.g. belonging to stadium 7RO) collected in spring x+2 can be either males
which became adl in spring x+2, or “old” adl which became 7RO adults the previous autumn:
succession 7RO autumn adl .... 7RO spring adl (Table 7).

Table. 7. — Development and cycles of males of T. niger in Saarland and Burgundy, represented in a highly simplified
way. Cases 1, 2, 3: starling from juvenile male (juv), with 6RO in summer/autumn (aut) x+1. - Cases 4, 5, 6:
starting from juv or adl, with 7RO in summer/autumn x+1. - Cases 7 to 12: males of the new generation (G) -
i.e. adl or juv in spring x+3 - are represented. Only the case of LL schcf (and not that of SL ones) has been
taken into account. Males from the generation of year x-1 have not been represented. First adults are underlined
at the time they become adl. Dashed line means a moult. Dotted line means no moult.

autumn x+1

spring x+2

autumn x+2

spring x+3

autumn x+3

6RO 7RO

6RO 7RO 8RO

6RO 7RO 8RO 9RO

7RO 8RO 9RO 10RO

8RO 9RO 10RO

juv . adl . si si . ad 2

juv . juv — . adl . adl . si

juv . juv . juv . adl . si

juv . adl . si . si . ad2

juv . juv . - . adl . adl . si

adl adl . si si ad2

1 1

(new G) adl . . s 1

juv . adl

juv juv

adl . si

juv . adl

adl . . si

In other words, in the case of LL, the intercalaries produced in summer/autumn of year
“y” will not be “useful” for the reproduction which takes place in spring of the following year
y+1. They will be able to reproduce - as ad2 - in spring of year y+2 (or in autumn of year y+1).

The above mentioned ad2 will add to the adl possibly produced in year y+2. When the
new (= freshly) adl are few, we will observe situations like those found in Burgundy (France)
near Chamboeuf and in the “Combe de Sainte Foi” (SahLI, 1989): in these sites the postadult
males (mostly ad2) were well represented in spring 1988.

In order to better unterstand the difference between LL and SL, let us add the following
comment: if we were in the presence of SL (instead of LL) produced in autumn of year y, the
males could mate the following spring y+1, after becoming postadult males (instead of staying
intercalaries).

Contrary to SAHLI’s 1967 statement, LL si may sometimes outnumber adl, in spring.
Such a case has been observed in the forest glade of Segrois (“Sommiere de Segrois” near
Chamboeuf, Cote-d'Or, France ), March 30-31, 1985. In this case, the number of adl produced
in spring 1985 was low (the winter 1984-1985 was exceptionally cold).

Summer/autumn (Table 6 and Fig. 1) is the main (or even exclusive) “season” of
schd1 production (e.g. ad. sch<+) in Saarland (epigean animals) and in Burgundy. At this time of
the year, the si (which have just become schcr) may outnumber the newly appeared adl under
two conditions (Table 6): (a) if the number of newly produced adl is low and (b) if the new
spring adl of year x+1 were well represented. Such a case was observed in Wadrill (15. 9.
1957).

380

FRANCOIS SAHLI

Table 8. — Simplified table of the cycles in males (A, C) and females (B) of T. niger in Saarland and Burgundy (only
some of all the possibilities are given). Among other things, one can see when adl males appear (9RO males
have not been represented). The possibility of 3RO larvae in autumn of the year x is not indicated. In A brackets
mean ad2 originating from SL. In C, generation x-2. ED = Egg Deposition, N.G. = New Generation. Dashed line
means a moult. Dotted line means no moult.

Autumn x

Spring x+1

Autumn x+1

Spring x+2

Autumn x+2

Spring x+3

4RO

5RO

6RO

7RO

cfJuv

cf adl--

7RO

adl

c?Juv

8RO

- adl

adl

8RO

schc?-*- .

\ _

adl .

9RO

. sc he?

schc?

8RO 9RO 10RO

schc?

. (ad2)

adl

9RO

. schc?

10RO

- . <ad2)

N.G.

6RO c? Juv
7RO <? Juv
7RO c?adl

6RO

7RO

. 8RO

7RO

. 8RO

. 9RO

N.G.

6RO

7RO

adl 7RO .

schc? 8RO .

7RO

adl .

schc? 8RO .

7RO

adl .

- ad2 9RO

8RO 9RO
si-* — ad2

^ — ss .

si .

. ad2 9RO

9RO

. ad2

. ss

(8RO)

At low altitude in Saarland (epigean animals) - and probably in different regions of
Germany, like Eifel, Taunus, Hunsruck, Hesse - and in Burgundy, summer and autumn
typically constitute the time at which schc? are produced in T. niger (case a). Consequently
schc? may be well represented in autumn (SAHLI, 1967). But, paradoxically, this is not always
the case - at least apparently. Thus the number of adl can prevail over that of si when, for
instance, the autumn adl production (year x+1. Table 6) was good. Such a case was observed at
Wadrill (30.9.1962). In other words, the number of new autumn adl can prevail over autumn
si of year x+1 (Table 6) when the production of spring adl of year x+1 was low.

Until recently, we had not understood what really happens in nature in T. niger (SAHLI
1966, 1967. 1970). Tables 5-8 and Figure 1 allow us to explain all previous field observations
and culture results.

In T. niger the predominance of LL Scho” over SL (case a) might correspond to a sexual
rest period (Table 9) in females (in the sense that females cannot be fertilized each year) - a rest
period of one or several years, during which females may elaborate new ripe eggs. In other
words, a female “indirect iteroparity” (SAHLI, this volume ) might be another raison d'etre of LL
schc? and repetitive ones, which are male forms able to withstand harsh conditions (SAHLI,
1991c). Nevertheless, another hypothesis can be put forward: a splitting up and spreading of 99
adl (and of egg depositions) over several years might be possible - such a strategy has been
called the “CAT strategy” in males (SAHLI, 1990b). The appearance of female adl (from a single
generation) during several years has been considered in mediterranean#. sabulosus (SAHLI,
1991b).

Source : MNHN. Paris

PERIODOMORPHOSIS, ITEROPARITY AND LIFE-CYCLES IN A JULIDAE

381

Table 9. — T. niger in Burgundy and Saarland. Indirect iteroparity (hypotetically admitted) in females, combined with
LL in males is represented. The year x+1 during which no egg deposition occurs, is labelled “sexual rest” period
and corresponds to the time needed for a new egg production; sum/aut: summer/autumn.

x+1

x+2

spring

sum/aut

spring

sum/aut

spring

cfcf adl .

si .

si LL .

ad2 .

ad2

99 first egg deposition

adl

"sexual rest" period

2nd egg deposition

BLOWER (1969) & coll. ( 1964, 1974, 1977) had the great merit to introduce the notion of
female iteroparity in myriapodology. Nevertheless this is only an hypothesis, and not a well
established fact. I have partly adopted Blower's idea, at least as a working hypothesis. SAHLI
(1993) subdivided it into (a) a direct iteroparity (egg deposition occurs in two consecutive years)
and (b) an indirect one (the first egg deposition is separated from the second by an interval of
over 2 or 3 years). The possibility of an indirect female iteroparity - with presence of LL schc?
and R ones - has been suggested (SAHLI, 1993) in the case of the Pyrenean cave blaniulid
Typhloblaniulus lorifer consoranensis , particularly because of the low temperatures which exist
in these caves. The same reasoning can be applied to T. niger, which lives rather in relatively
“cold” regions. The South limit in France seems to be the Departement de la Saone-et-Loire.
This species can be found further south in France at higher altitudes (particularly in the south¬
west, in the Pyrenees). In spite of periodomorphosis, T. niger is far less adapted to a hot and
dry climate (e.g. of the Mediterranean type) than O. sabulosus.

Fig. 2. — T, niger. A: at Wadrill (Saarland, Germany).
Al: stadium and specific sexual (ad or schcf)
frequencies of adult males (solid) and intercalaries
(open), respectively at each stadium, from stadia
7 to 12RO. [e.g.: in stadium 8RO: ratio 8RO adl
cT /all adult cf (184); or: 8RO schcf /all schcf
(92)). The frequencies are established from 184 ad
cf and92schcf. respectively , collected between
September 1957 and 1963 (inclusive). A2: total
frequencies of ad cf and schcf, established from
375 individuals (adcf + schcf) collected between
May 1957 and 1963 (inclusive) [e.g. ratio 7RO ad
cf/375 or ratio 8RO schcf /375. B: at Citeaux
(Burgundy, France). Bl: stadium and specific
sexual frequencies of ad cf (204) and schcf (1 15)
collected in 1963 and 1964. B2: total frequency
of adcf (204) and schcf (1 15), according to 319
individuals collected in 1963 and 1964.

Figure 2 shows the ratios of adults and schc? at two Burgundy sites. Notice, particularly at
Citeaux (a) the lower percentage of s2 intercalaries (typically stadium 10RO when adl appear at
7RO) compared with the percentage of si (stadia 8 and 9RO) and, correlatively, (b) the lower

382

FRANCOIS SAHLI

percentage of postadult ad2 males (a mixture of stadia 9RO males pro parte and possibly 10RO
males pro parte). Drawing a parallel between ad2 and adl is in fact a comparison between
animals which became adults at two different years; in this respect, one has to take into account
the comment made before and Tables 5 & 6. Moreover, stadia 9 and 10RO may comprise not
only ad2, but also adl (SAHLI, 1989). As for 12RO s3, they are very few: they imply some rare
ad4 at both sites.

CONCLUSION

Taking into account the author’s experimental results and observations, as well as those of
VERHOEFF, in the present state of our knowledge, the following statements can be made. In
good conditions and at low altitude in Burgundy and in Saarland, a majority of 7 and 8RO adl is
able to turn into si. The si are, in the majority, able to give either ad2 or ss. Then a fall seems to
appear: only a part of the ad2 are able to turn into s2 and afterwards into ad3 - the relative
importance of ad2 depends more or less on the temperature. A very low number of ad3 is able to
give s3 and then ad4 (the number of s3 and ad4 seems to increase with altitude and depends on
thermic conditions).

If one takes into account only the existence of LL all over the cycle and if the starting point
for adl takes place in spring of the year y, then males will become ad4 in autumn of year y+4. If
adl are 2 years old and possess 7RO, the ad4 might in theory be 6 years old, with 13RO; in the
"Cirque de Gavamie”, in the Pyrenees at 2500m alt. and over, the oldest adult male with a ring
formula of 72/2 (BROLEMANN, 1927) might belong to a stadium equivalent of 23RO (SAHLI,
1969).

Concerning T. niger , three comments will be added.

a) A possible dispersal - in time and space - variable for the male categories from one year
to the next one, has to be taken into account. Spring schd" which became schd" the previous
autumn y may have dispersed the year y in a given direction, while new adl which became
adults in spring y+1 may disperse in another direction year y+1. In other words the dispersion
areas in years y and y+1 might be different. So animal collections from a single locality may be
biased.

b) Intercalaries can sometimes be more or less grouped or “gathered” (cf. SAHLI, 1991a in
O. sabulosus). In Burgundy and Saarland T. niger , in spring and in case a, the activity of LL
scho" and R individuals may be lower - because they do not moult in spring and do not search
for females - than in new spring adult males. New spring adult c fcT are very active for two
reasons (a) in spring they have just moulted (P) they move in order to search for females for
spring mating (for an account of male activity in general, see SAHLI, this volume). FAIRHURST
(1968) stated that, in Britain, Tachypodoiulus spring sch <f - which, according to him, have just
moulted in spring (= SAHLI's case b?; thus they might be SL - or ss?) - would, in spite of the
moult, have a lower activity than spring adult males. FAIRHURST may be right. But field (or
laboratory) experiments should be carried out. With experimental animals in equal numbers, the
activity of fresh spring scho" (si -and not ss) should be compared with that of fresh spring
add" (adl) in England and in Tachypodoiulus ; if, under these conditions, there is a difference,
then it can be attributed to sexual activity.

c) Due to the lengh of time of LL or R sch d" states - thus to the long length of time
between the appearances of adld" and ad2d" - it is not impossible that a loss of individuals might
occur. If a loss really happens, it might be higher in LL and R schd" of T. niger than in SL of
Mediterranean O. sabulosus.

Source : MNHN, Paris

PERIODOMORPHOSIS, ITEROPARITY AND LIFE-CYCLES IN A JULIDAE

383

REFERENCES

Baker, G. H. , 1978. — The post-embryonic development and life history of the millipede, Ommatoiulus moreletii
(Diplopoda: Iulidae), introduced in south-eastern Australia. J. Zool. Lond 186 : 209-228.

Baker, G. H.. 1984. — Distribution, morphology and life history of the millipede Ommatoiulus moreletii (Diplopoda:
Iulidae) in Portugal and comparisons with Australian populations. Aust. J. Zool.. 32 : 811-822.

Blower, J. G., 1969. — Age structures of millipede populations in relation to activity and dispersion. In : The Soil
Ecosystem, Syst. Ass. Pubis., 8 : 209-216.

Blower, J. G. & Gabbutt, P. D., 1964. — Studies on the millipedes of a Devon oak wood. Proc. Zool. Soc Lond
143 : 143-176.

Blower, J. G. & Miller, P. F., 1974. — The life cycle and ecology of Ophyiulus pilosus (Newport) in Britain. Symp.
Zool. Soc. Lond., 32 : 503-525.

Blower, J. G. & Miller, P. F., 1977. — The life-history of the julid millipede Cylindroiulus nitidus in a Derbyshire
wood. J. Zool. Lond., 183 . 339-351.

Brolemann, H. W., 1927. — Myriapodes du cirque de Gavarnie et note sur les vulves de trois d'entre eux. Bull. Soc. Hist,
nat. Toulouse, 56 : 531-548.

Fairhurst, C. P., 1968. — Life cycles and activity patterns of schizophylline millipedes. Ph. D. Thesis, University of
Manchester, 407 pp.

Fairhurst, C. P., 1974. — The adaptive significance of variations in the life cycles of schizophylline millipedes.
Symp. zool. Soc. Lond , 32 : 575-585. -

Halkka, R.. 1958. — Life history of Schizophyllum sabulosum (L.) (Diplopoda, Iulidae). Ann. Zool. Soc. Fenn.
"Vanamo" , 19 : 1-72.

Hopkin, S. P. & Read, H., 1992. — The biology of millipedes. Oxford, Oxford Univ. Press, 233 pp.

Sahli, F ., 1966. — Contribution a letude de la periodomorphose et du systeme neurosecreteur des Diplopodes Iulides.
These de Doctorat £s-Sciences Naturelles, Universite de Dijon. Bernigaud & Privat, 226 pp.

Sahli, F., 1967. — Sur la frequence des males intercalaires du Diplopode Tachypodoiulus albipes (C. L. Koch). C. R.
Acad. Sc. Paris, ser. D., 264 : 618-620.

Sahli, F., 1969. — Contribution a I'etude du developpement post-embryonnaire des Diplopodes Iulides. Ann. Uni vers,
saraviensis, Berlin. Stuttgart, Gebr. Borntraeger, 154 pp.

Sahli, F., 1970. — Altitude et males intercalaires chez le Diplopode Tachypodoiulus albipes (C. L. Koch). C. R. Acad.
Sc. Paris, ser. D., 271 : 2175-2178.

Sahli, F., 1986. — Influence des orages sur certains deplacements d 'Ommatoiulus sabulosus (L.) (Myriapoda,
Diplopoda, Julida) dans les Alpes-Marilimes. Bull, sci . Bourg., 39 : 55-57.

Sahli, F., 1989. — The structure of two populations of Tachypodoiulus niger (Leach) (Myriapoda, Diplopoda. Julida) in
Burgundy and some remarks on male periodomorphosis. Rev. Ecol. Biol. Sol, 26 : 355-361.

Sahli, F., 1990a. — Recherches sur le cycle des males et la frequence des intercalaires $ Ommatoiulus sabulosus
(Myriapoda, Diplopoda, Julidae) en Bourgogne. Inversion experimentale du nombre des intercalaires et des adultes
en automne. Bull. sci. Bourgogne, 45 : 51-59.

Sahli. F., 1990b. — On post-adult moults in Julida (Myriapoda, Diplopoda). Why do periodomorphosis and
intercalaries occur in males? In : A. MlNELLl, Proc. 7th Intern. Congr. Myriapodology, Leiden. Brill : 135-156.
Sahli, F., 1991a. — Recherches sur le cycle des males d Ommatoiulus sabulosus (L.) (Myriapoda. Diplopoda. Julidae )
dans les Alpes-Maritimes et en Provence. Remarques sur les rapports entre la periodomorphose et l’altitude. Bull. sci.
Bourg., 44 : 33-39.

Sahli, F., 1991b. — Frequence de la periodomorphose chez le Diplopode Julide Ommatoiulus sabulosus (L.) dans les
Alpes-Maritimes et en Provence : II Passages males intercalaires - males adultes. Bull. sci. Bourgogne, 44 : 41-47.
Sahli. F., 1991c. — Augmentation et production experimentales de males intercalaires chez des Diplopodes Julida. Note
preliminaire. C. R. Acad. Sci. Paris, Ser. Ill, 313 : 59-63.

Sahli, F., 1992. — On male reproduction strategies in Ommatoiulus sabulosus (L.) in the Maritime Alps and Provence
(France) : juvenile to adult maturation moults. Ber. nat.-med. Verein, Innsbruck, suppl. 10 : 167-176.

Sahli, F., 1993. — Signification de la periodomorphose et des intercalaires males et femelles chez les Diplopodes
Blaniulides cavernicoles pyreneens. Mem. Biospeol., 20 : 209-215.

Verhoeff, K. W., 1915. — Zur Kenntniss einiger alpiner Chilognathen. Zool. Anz., 45 : 219-238.

Verhoeff, K. W., 1916. — Abhangigkeit der Diplopoden und besonders der Juliden-Schaltmannchen von ausseren
Einflussen. Z. wiss. Zool., 116 : 535-586.

Verhoeff, K. W., 1923. — Periodomorphose. Zool. Anz.. 56 : 233-238 & 241-254.

Verhoeff, K. W., 1925. — Mediterrane Chilopoden und Notiz zur Periodomorphose der Juliden. Zool. Anz., 64 : 63-80.

384

FRANCOIS SAHLI

VERHOEFF, K. W„ 1928. — Diplopoda 1 .In : H. G. Bronn's Klassen und Ordnungen des Tierreichs, 5, Leipzig,
Akademische Verlagsgesellschaft : 1-1072.

VERHOEFF, K. W, 1932. — Diplopoda 2. In : H. G. Bronn's Klassen und Ordnungen des Tierreichs, 5, Leipzig,
Akademische Verlagsgesellschaft : 1073-2084.

VERHOEFF, K. W., 1934. — Ueber Diplopoden der allgauer Alpen. deutschc Craspedosomidae und Penodomorphose.
Zool. Anz 108 : 27-40.

Source : MNHN, Paris

cAMP Influence on Brain and Germinal Cells RNA
Syntheses in Lithobius forficatus (L.): an
Autoradiographic Study

Michel DESCAMPS *, Catherine Jamault-Navarro **
& Marie-Chantal FABRE *

* Ecophysiologie d’lnvertebres du Sol, Laboratoire de Biologie Animale, Universite de Lille I, F-59655
Villeneuve d'Ascq CedeX, France (to whop* all correspondence must be sent)

** Laboratoire de Biologie Animale, UFR Sciences Exactes et Naturelles, Universite de Picardie, 33 rue Saint-

Leu, F-80039 Amiens Cedex, France

ABSTRACT

Mature L. forficatus were injected with 2 nmol of dibutyryl cAMP (sodium salt) [dBcAMP]. Animals of day 1, 2, 3 and
7 after injection were investigated. Autoradiographs were analysed either by cytophotometry (Leitz MPV) or by image
analysis (Biocom 2000 device). Germinal cells : in oocytes, the supply of dBcAMP led to a significative increase in
[3H]-uridine uptake during the first 3 days of the experimental series. Nevertheless, on day 7, control values were
obtained. In spermatocytes, two cases must be reported: if the testes were not in a phase of active spermatogonial
divisions (“phase de reconstitution”), an increase in RNA syntheses was observed. At the opposite of that measured for
oocytes, the maximum of label was observed on day 7. For testes with active mitoses, the uptake remained at a low level,
either in controls or in dBcAMP injected animals. Brain: analysis performed on various areas (middle and lateral parts of
the so-called pars intercerebralis, frontal lobes and cerebral glands) showed that dBcAMP has a stimulating effect on the
uptake of the tritiated precursor. The increase of RNA syntheses observed in germinal cells and in neurons shows that
cAMP is a good candidate to be the second messenger of the stimulating neuropeptide(s) issued from pars intercerebralis
neurosecretory cells. Our results have, in addition, shown that a refractory period occurs during the period of active
mitoses in the testis.

RESUME

Influence de cAMP sur la synthese de TARN du cerveau et des cellules germinates chez Lithobius
forficatus (L.): etude autoradiographique.

Apres injection de 2 nmol de dibutyryl cAMP (sel de sodium) (dBcAMP], des adultes matures de Lithobius forficatus ont
<$te etudies aux jours 1, 2, 3 et 7. Les autoradiographies analysees, soit par cytophotometrie, soit par un analyseur
d'images Biocom 2000, montrent que les ovocytes incorporent significativement plus d'uridine tritiee que les t6moins
durant les 3 jours qui suivent l’injection de dBcAMP. Des valeurs temoins sont cependant recuperecs au jour 7. En ce qui
concernc les spermatocytes, deux cas se presentent : si le testicule n’est pas en phase de reconstitution (periode de
divisions goniales), une augmention des syntheses d’ARN est observee. Aucune augmentation de synthese n’est observe
si de nombreuses divisions goniales sont en cours dans le testicule. Les analyses effectuees sur differentes zones du
cerveau (pars intercerebralis, lobes frontaux) ou sur la glande cerebralc, montrent un effet stimulates sur I'incorporation

Descamps, M., JAMAULT - Navarro, C. & FABRE , M.-C.. 1996. — cAMP influence on brain and germinal cells
RNA syntheses in Lithobius forficatus (L.): an autoradiographic study. In: GEOFFROY, J.-J., MAURlfcS, J.-P. & NGUYEN
Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. natn . Hist. nat.. 169 : 385-390. Paris ISBN : 2-85653-
502-X.

386

MICHEL DESCAMPS. CATHERINE JAMAULT-NA V ARRO & MARIE-CHANTAL FABRE

du precurscur tritie. L’augmentation des syntheses d’ARN observee aussi bien dans les cellules germinales que dans Ie
systeme nerveux montre que I'AMP cyclique est sans doute Ic second messager du (des) neuropeptidc(s) issu(s) des cellules
neuros6cretrices de la pars intercerebralis. Nos r£sultats momrent cn outre I’cxistcnce dune periode refractaire a Taction
de I'AMP cyclique.

INTRODUCTION

In Lithobius, the endocrine control of gametogenesis is the result of the balance between
stimulating factors - ecdysteroids and a hormone issued from pars intercerebralis neurosecretory
cells (pi NSC) - and a moderating one released from the cerebral glands (neurohemal cephalic
organs) (reviews: JOLY & DESCAMPS. 1988; DESCAMPS, 1992a). In addition, in females, it
has been shown that too high levels of ecdysteroids triggers the release of the moderating factor
(DESCAMPS, 1992b). It has been previously demonstrated that cAMP was present in testes and
that the level of this messenger was increased, except during meiosis, after electrical stimulation
of the pi NSC (DESCAMPS et at, 1986). So, it was of interest to proof the effects of cAMP on
germinal cells and on brain neurons metabolism, in order to compare the results to those
obtained after electrical stimulation or ecdysteroid injection.

MATERIAL AND METHODS

The experiments were conducted on mature Lithobius forficatus (L.) collected in northern France, during autumn, in order
to have animals showing a low rate of RNA syntheses, at least in male germinal cells.

Animals were injected with 2 nmol of dibulyryl cAMP (sodium salt) [= dBcAMP: purchased from Sigma] in solution in a
saline adapted to chilopods. The autoradiographic study was conducted after the injection of 185 KBq (= 5 jiCi) of [3HJ-
uridine (CEA. France; specific activity 166,5 TBq (= 45 Ci]/mMol) 48 hrs before fixation. The animals were fixed 1, 2.
3, and 7 days after the injection of the dBcAMP. Tissue sections, treated according to FlCQ (1961), were covered with
Kodak NTB3 emulsion. Kodak D19 was used as developer. The cytophotometric study of the labelling of germinal cells
was conducted either with a Leitz MPV cytophotometer, using a 500 }im2 diaphragm. Only growing spermatocytes (40 to
70 pm in diameter) or vitcllogenetic oocytes (stage 2B and beginning of stage 3 according to HERBAUT, 1972) were
investigated. Brain endocrine areas were studied with an image analysis device (Biocom 2000). In the graphs concerning
the Iattest results no error bars are shown, measurements being the results of area labelling expressed as a mean
percentage of label of the area (neuropil label is defined as an internal control and fixed to 1 ) and not as the result of cell
types measured individually.

RESULTS

Germinals cells

In oocytes, a significant increase of uptake of ['HJ-uridine was recorded in animals of day
1, 2, 3. In day 7 animals the label was comparable to that of the controls (Fig. 1).

In males, two cases were recorded. In animals with a testis showing growing
spermatocytes, an increase of uptake was present in all the experimental series (Fig. 2). In the
animals that undergo the renewal of their stock of spermatocytes (“periode de reconstitution”,
JOLY & DESCAMPS, 1969), at the opposite of that observed precedently, the level of label was
low in controls ( circa 30 units), and in addition, no effect of dBcAMP was found (label staying
between 20 and 25 units).

Brain

In all females, an increase uptake of the tritiated precursor was found in the endocrine
areas [median (Fig. 3) and lateral (Fig. 4) parts of the pi, frontal lobes (Fig. 5)]. Nevertheless,
the maxima recorded here were different according to the type of NSC studied.

The results were quite different for the cerebral glands. An increased uptake took place
during the first three days, a control value was measured from day 7 animals (Fig. 6).

Source : MNHN, Paris

BRAIN AND GERMINAL CELLS RNA SYNTHESES IN A LITHOBIOMORPH

387

time (days)

Fig. 1. — Cytophotometric measurements of labelling
over oocyte nuclei after 1, 2, 3, 7 days.
Means ± SD. Controls: open bars.

100-

2 3 7

time (days)

Fig. 2. — Cytophotometric measurements of labelling
over spermatocyte nuclei after 1, 2. 3, 7 days.
Means ± SD. Controls: open bars.

Median part of the pars intercerebralis area

Lateral parts of the pars intercere&ralls area

Figs. 3-6. — Mean label of brain endocrine areas of pi and of cerebral glands in females. Controls: open bars.

Source : MNHN, Paris

388

MICHEL DESCAMPS, CATHERINE JAMAULT-NAVARRO & M ARIE-CH ANTAL FABRE

In males, we can only report the results concerning animals not implied with the renewal
or the spermatocyte stock. As a consequence, the label was low over the brain and cerebral
gland, but not strong enough to enable measurements by the image analyser. In animal not
concerned with this refractory period, an increase in the uptake of [-'Hj-uridine was observed,
but tor shorter times than in females. For the median part of the pi, this increase lasted two days
Then, the values recorded were significantly under the control values (Fig. 7). Lateral parts of
the pi showed, compared to the controls, increases of uptake, but only during the first three
days; nevertheless, it must be noticed that these increases were of less great extend than those
recorded for the median part (Fig. 8). In frontal lobes labelling was increased only in the first
two days and then control values were recorded (Fig. 9). The most dramatic increases were
recorded tor the cerebral glands, particularly for animals of day 2 and day 3 In day 7 animals
values recorded were slightly under the control values (Fig. 10).

Frontal lobes

Cerebral glands

Figs. 7-10, - Mean label of brain endocrine areas and of cerebral glands in males. Comrols: open bars.

vuinllumuns and DISCUSSION
. A* a conclusion, we can say that cAMP stimulates the uptake of PHl-uridine and

3 NSCC or g“c=,fs°°d Cand,da,C t0 be 2nd “8“ of variL kind ‘of ceUs as fhose

Source : MNHN, Paris

BRAIN AND GERMINAL CELLS RNA SYNTHESES IN A LITHOBIOMORPH

389

Stimulation of uptake in oocytes is only transient: this result can be compared to findings
after injection of (at least) 0.4 pg of 20-hydroxyecdysone and was explained by the release of a
moderating factor, in order to limitate the level of the metabolism and to enable a normal and
regular vitellogenesis (DESCAMPS, 1992b). So, also in this case, the release of a moderating
factor is more likely involved in the regulatory process, explaining as a consequence the rather
short time of increased metabolism in oocytes.

At the opposite, stimulation of spermatocyte metabolism was observed during the whole
time of experiment, comparable to previous results (DESCAMPS, 1981, 1991). It appears that,
either there is no release of a moderating factor in males, the balance between the stimulating and
the moderating hormones being not regulated in the same manner in males and in females, or, as
another explanation, the spermatocytes are less sensitive to variations of hormonal levels. They
might be protected by the testis blood barrier, the lattest being regulated partly by 20-
hydroxyecdysone (BENIOURI, 1984).

Concerning the brain, the increase of tritiated uridine uptake in pi NSC requires more time
in females (at least 7 days) than in males (2 or 3 days). This fact can be related to the
reproductive physiology of animals: females are in their vitellogenetic phase whereas males are
entering in a period of minimal rate of metabolism (the so called winter rest period). Differences
between female and male brains were previously evidenced in transplantation experiments on the
influence of the pars intereerebralis on the gametogenetic cycle (DESCAMPS, 1974), and, at this
time, it was suggested that these differences took their origin in the course of gametogenetic
cycles. The present results are in full agreement with these statements.

In the frontal lobes, increased values were recorded during the whole time of experimental
series for both sexes, whereas in cerebral glands an increase was measured for only the first
three days. It is difficult to explain or to relate these findings to physiological events: release of
moderating factor by the cerebral gland do not imply increased syntheses, numerous secretory
granules being stored in the cells and in the axonal endings of the glands. In short, for the brain,
differences are only recorded according to the sex, and this was previously reported in
transplantation experiments (DESCAMPS, 1974).

cAMP is involved in various kind of processes. For example, in Crustacea it has been
found that the processes of protein synthesis are involved in previtellogenic oocytes (EASTMAN-
REKS & FlNGERMAN, 1984) and those which are necessary for eedysteroid production are
inhibited by MIH (molt inhibiting hormone) through cAMP (among other authors: MATTSON &
Spaziani, 1985; SEDLMEIER & Fenrich. 1993). At the opposite, in Insecta. the process of
eedysteroid synthesis is stimulated by PTTH (prothoracotropic hormone) through cAMP (among
others: SMITH eta!., 1984, 1993).

We have found that, in the brain, stimulating or moderating factors of NSC secreting show
both an increased uptake of ['HJ-uridine after dBcAMP supply. It is the question if the increase
of this uptake that is triggered in NSC frontal lobes is directly induced by the cyclic nucleotide or
if the activation of NSC frontal lobes is a consequence of the activation of pi NSC in order to
counteract their action in a regulatory process of metabolism? The answer to such a question
cannot be given before the localization of adenylate cyclase will be investigated.

REFERENCES

Beniouri, R., 1984. — Testis blood barrier control by 20-hydroxyecdysone in Lithobius forficatus (L.) (Myriapoda,
Chilopoda). Cytobios, 40 : 159-170.

Descamps. M., 1974. — Elude du controle cndocrinien du cycle spermatogenetique chez Lithobius forficatus (L.)

(Myriapode, Chilopode). Role de la pars intereerebralis. Gen. Comp. Endocrinol .. 25 : 346-357.

DESCAMPS, M., 1991. — Role of morphogenetic hormones in spermatogenesis in Myriapoda. In : A. P. Gupta,
Morphogenetic Hormones of Arthropods. Vol. I. part 3: Roles in histogenesis, organogenesis and morphogenesis.
New Brunswick & London. Rutgers University Press : 567-592.

390

MICHEL DESCAMPS. CATHERINE JAMAULT-NAVARRO & MARIE-CHANTAL FABRE

Descamps. M.. 1992a — Endocrine events during the life cycle of Lithobius forficatus (L.) (Myriapoda, Chilopoda). Ber.
nat.-med. Verein Innsbruck, suppl. 10 : 111-116.

Descamps, M., 1992b — Influence de la 20-hydroxyecdysone sur les syntheses d’ARN dans les ovocytes de Lithobius
forficatus L. (Myriapode, Chilopode). Bull. Soc. Zooi Fr 117 : 139-147.

Descamps, M., Cardon, C. & Leu. B., 1986. — Influence of brain electrical stimulation or 20-hydroxyecdysone
injection on the cAMP level in the testes of Lithobius forficatus (L.) (Myriapoda, Chilopoda). hi : M. Porchet, J. C
ANDRLfeS & A. Dhainaut, A dvances in Invertebrate Reproduction 4. Amsterdam, Elsevier Science Publishers : 505.

Eastman-Reks, S.. Fingerman. M., 1984. — Effects of neuroendocrine tissue and cyclic AMP on ovarian growth in vivo
and in vitro in the fiddler crab. Uca pugilator. Comp. Biochem. Physiol ., 79 A : 679-684.

FlCQ, A., 1961. — Contribution a I etude du metabolisme cellulaire au moyen de la technique autoradiographique.
Bruxelles, Inst. Inter. Univ. Sci. Nucl. Beige, Monographic n° 9 : 1-121.

Herbaut, C., 1972. — Etude cytochimique et ultrastructurale de l'ovogenese chez Lithobius forficatus (L.) (Myriapode,
Chilopode). Evolution des constituants cellulaires. Wilhelm Roux'Archiv , 170 : 115-134.

Joey, R. & Descamps, M.. 1969 — Evolution du testicule, des vesicules seminales et cycle spermatog£netique chez
Lithobius forficatus L. (Myriapode Chilopode). Arch. Zooi exp. gen. 110 ; 341-348.

Joly, R. & Descamps, M., 1988 — Endocrinology of Myriapods. In : H. Laufer et R.G.H. Downer, Endocrinology of
selected Invertebrate Types, New- York, Alan R. Liss, 429-449.

Mattson, M. P. & Spaziani, E., 1985. — Cyclic AMP mediates the negative regulation of Y-organ ecdysteroid
production. Mol. Cell. Endocrinol.. 42 : 185-189.

Sedlmeier, D. & Fenrich, R. 1993. — Regulation of ecdysteroid biosynthesis in crayfish Y-organ : I. Role of cyclic
nucleotides. J. exp. Zooi, 265 : 448-453.

Smith, W. A., Gilbert, L. I. & Boi.lenbacher, W. E., 1984. — The role of cyclic AMP in the regulation of ecdysone
synthesis. Mol. Cell. Endocrinol., 37 : 285-294.

Smith, W. A., Varghese, A. H. & Lou. K. J.. 1993. — Developmental changes in cyclic AMP-dependant protein kinase
associated with increased secretory activity of Manduca sexta prothoracic glands. Mol. Cell. Endocrinol ., 90 : 187-
195.

Source : MNHN, Paris

Cadmium Kinetics in Lithobius forficatus (L.) during
Experimental Contamination and Decontamination

Sylvie GERARD , Marie-Chantal FABRE & Michel DESCAMPS *

Ecophysiologie d'Invertebres du Sol, Laboratoire de Biologie Animalc, Universite de Lille I
F-59655 Villeneuve d'Ascq Cedex, France

* to whom all correspondence must be sent

ABSTRACT

Experiments were conducted on adult mature L. forficatus specimens collected in an are^ of Northern France
contaminated by various metals (Al, Cd, Cu, Pb). The mean level of cadmium in the soil reached about 60 ppm. Kinetics
of decontamination, performed in autumn experiments showed a decrease in Cd level in centipedes, from about 30 ppm at
the beginning of the experiment to about 12 ppm ten weeks later. Kinetics of contamination, starting after a period of
decontamination, showed at first a dramatic increase in Cd levels (up to a mean 80 ppm), followed by a decrease,
although the animals were regularly fed with cadmium contamined larvae. Difference between animals fed or not with
cadmium containing diet ranged from 18 ppm to only 8 ppm at the end of the experimental series. Experiments conducted
either in autumn or spring showed comparable evolution of level curves, if not the same values.

RESUME

La cinetique du cadmium chez Lithobius forficatus (L.) au cours d'une experience de
contamination et de decontamination.

Des experiences ont ete menees chez des adultes de Lithobius forficatus preleves dans le Nord de la France, dans une
zone contaminee par divers mctaux (Al, Cd, Cu, Pb). La cinetique de decontamination, suivie experimentalement durant
rautomne, montre une diminution des taux de 30 ppm au debut a environ 12 ppm dix semaines plus tard. Les cinctiques de
contamination, debutant apres une periode initiale de decontamination montrent dans un premier temps une
augmentation importante des taux atteignant 80 ppm. Une decroissance reguliere dcs taux esl ensuite observee, alors que
les animaux continuent a recevoir une nourrilure contaminee par le cadmium. A la fin de I’experience, la difference entre
les animaux nourris ou non avec de la nourriture cadmiee est d'cnviron 10 ppm. Aux valeurs pr£s. l'experimentation,
qu'elle soit automnale ou printanni^re. montre les memes phenomenes.

INTRODUCTION

Pollution by heavy metals is one of the main ecological problems of industrial areas, at
least for invertebrates (HOPKIN, 1989). In Northern France, numerous metal works were closed
during the last 20 years, and after demolition of furnaces and buildings, the resulting industrial
wastelands were rehabilited. Part of such a wasteland, located in Mortagne du Nord. previously
a zinc works, was used as an experimental area.

Gerard. S., Fabre, M.-C. & Descamps, M., 1996. — Cadmium kinetics in Lithobius forficatus (L.) during
experimental contamination and decontamination. In: Geoffroy. J.-J., MauriSs, J.-P. & Nguyen Duy - Jacquemin, M.,
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat ., 169 : 391-394. Paris ISBN : 2-85653-502-X.

392

SYLVIE GERARD. MARIE-CHANTAL FABRE & MICHEL DESCAMPS

It has been shown in a previous work (HOPKIN et al., 1985) that chilopods are one of the
invertebrate groups that is common in metal contaminated areas. Despite studies concerning the
rate of assimilation of heavy metals by Lithobius variegatus (HOPKIN et al., 1985; HOPKIN &
Martin, 1984), and some other soil arthropods (Janssen et al. 1991), no data exist about the
dynamics of accumulation and detoxication of metals in centipedes.

The present study is part of a program initiated by “Espace Naturel Regional du Nord Pas
de Calais” about the biology and evolution of industrial wastelands.

MATERIAL AND METHODS

Animals

Mature adults Lithobius forficatus (L.) were collected on the metalliferous grassland located in Mortagne du Nord.
For some experimental series, animals of the same subareas were collected separately, the level of cadmium varying
according to the place of sampling (reported as "A" (lawn], "B" [stub] and "C" [mixed poplar-willow-birch woodland] in
the corresponding graph).

In the lab, the centipedes were fed with Chironomus larvae bought at a fisher supplier and maintained either in tap
water or in water containing 20 mg/1 of CdCb- When used for feeding, the Cd mean level of the latter was 150 mg/kg.
Animals were starved for 3 days before analysis in order to ensure digestion and to not measure the gut content.

Each point of the decontamination or of the experimental contamination curve resulted from four to five animals
analysed individually.

Mineralization

Animals were dried at 80°C (12 hrs) and weighed. The samples, reduced to powder, poured into test tubes and added
with 1 ml HNO3 (Normapur) were kept at room temperature. Samples were then warmed up to 120°C and kept at this
temperature till half the acid mixture was evaporated. 1 ml of a mixture of HNO3 - H2SO4- HCIO2 (10v/2v/3v; Normapur
grade) was then added and warmed up to 180°C. When half the solution was evaporated, the resulting mixture was diluted
to 20 ml with deionized water.

The digests were analysed for cadmium by flame (Perkin Elmer 2380) or flameless (Varian AA 300) atomic
absorption spectrophotometry. Results are expressed hereafter in mg of cadmium per kg of dry mass.

RESULTS

The mean level in soil was about 40 mg/kg of Cd (ranging from 6 to 76 mg/kg), i.e more
than one hundred fold higher than in agricultural soils of Northern France (0.32 mg/kg, data
from “Chambre d' Agriculture du Nord”, October 1990). As a consequence, animals contained
more cadmium than those collected in unpolluted sites (for example, 4.2 mg/kg in another site
located in St-Amand, some kilometers southwest of Mortagne versus 10 to 30 mg/kg).

Animals fed with unpolluted diet showed a decrease of the level of Cd, as shown by the
slope of the linear regression (Fig. 1), evidence for the existence of a detoxication process.
Animals fed with Cd polluted diet (after a period of unpolluted feeding) showed a dramatic
increase of the Cd level during two weeks, and then decreased values were recorded, with a rate
of elimation far higher than that observed in controls (Fig. 2).

At the end of the experimental series, the values recorded in the two kinds of animals were
not so different (Fig. 2), controls ranging from 8 to 14 mg/kg and Cd fed animals about 20
mg/kg.

In experiments conducted during spring, only experimental contamination was performed,
after about two weeks of natural detoxication in the lab. The same shape of curve was observed,
as in autumn experiments: a dramatic increase followed by active decontamination (Fig 3). In
this particular case, the area of collection of animals was recorded, so we were able to
demonstrate the variations observed from one animal to another, according to the sample site
(Fig. 4).

Concerning the assimilation rate of Cd, we have indirect evidence for a quick elimination
or, at best, a non assimilation at the time of feeding. Indeed, based on the mean Cd level of the
diet, an animal ingested about 5.5 |ig Cd per week and the maximum level recorded a week after
the beginning of polluted feeding reached a mean 80 mg/kg of dry weigh, corresponding to
about 1.6 (ig of Cd per centipede, i.e. 1 |ig more than at the beginning of the experiment.

Source : MNHN, Paris

mg/kg (dry weight)

CADMIUM KINETICS DURING CONTAMINATION AND DECONTAMINATION

393

13 20

27 34 41

time (days)

48 55 62 69

Fig. 1. — Elimination of cadmium in L. forficatus
fed with unpolluted diet. Original data are
given as means with SEM. Solid line: linear
regression.

100-

50-

i i

. s .

0 6 13 20 27 34 41 48 55 62 69

time (days)

FlG. 2. — Accumulation and elimination of cadmium in
Lilhobius. Black squares: animals fed with
-- unpolluted diet (same animals as in Fig. 1); open
circles: animals fed with Cd polluted diet (starting
on day 15). Means ± SEM.

50-i

_ 4CH

4-»

-C
o>

J 30-

20-

10-

beginning of feeding with Cd polluted diet

20 27 34 41

time (days)

48 55 62 69

5 On

S 40-1
gi
o
2

>» 30-

C7>

^ 20-

10-

14 21 *28 35

time (days)

42 49

FlG. 3. — Accumulation and elimination of cadmium in
Lilhobius during spring experiments. Feeding
with Cd polluted diet starting on day 9. Means ±
SEM.

FlG. 4. — Same animals as in Fig. 3. but with individual
values according to the sample site (A. B. C; for
details, see the text).

DISCUSSION

The first point to discuss is the ability of Lilhobius to eliminate cadmium. Indeed, we have
shown a decrease of the Cd level when animals were supplied with clean diet. This rate of
elimination in our autumnal experimental series is about 20 ppm for ten weeks. It must be notice
that in spring experiments the level of Cd in animals was a bit lower. This fact may be related to
low amounts of metal ingested, consequence of the winter low rate of metabolism and poor
feeding.

394

SYLVIE GERARD. M ARIE-CH ANTAL FABRE & MICHEL DESCAMPS

The second point concerns the dynamics of accumulation: the animals reacted rather
quickly to Cd supply. Only two points of analysis show high values: during two weeks the
animals accumulate more Cd than they can eliminate. Then, they eliminate more Cd than they
assimilate. As the values recorded at the end of experimental series were quite comparable to that
measured at the time of collection, we can consider that the latter are equilibrium values for
animals when they are in a period of active physiology in metal polluted soil.

The assimilation rate of Cd shown by our results is a little bit higher (about 18%) than the
10% found by HOPKIN el cd. (1985) but we may keep in mind that the diet was not the same,
and that Chironomus larvae bring Cd both as compounds linked to the body and as soluble
CdCh to be found in the water layer surrounding the body.

ACKNOWLEDGMENTS

The present study was supported by a gram from "Espace Naturel Regional du Nord-Pas-dc-Calais” and “Fondation
de France" concerning the study of "Pelouse metallicole de Mortagne".

REFERENCES

HOPKIN, S. P., 1989. — Ecophysiology of metals in terrestrial invertebrates. London, Elsevier Applied Science.

Hopkin, S. P. & Martin. M. H., 1984. — Assimilation of zinc, cadmium, lead and copper by the centipede Lithobius
variegatus (Chilopoda). J. Appl. Ecol.. 21 : 535-546.

Hopkin, S. P., Watson, K., Martin, M. H. & Mould, M. L., 1985. — The assimilation of heavy metals by Lithobius
variegatus and Glomeris marginata (Chilopoda; Diplopoda). Bijdr. Dierk.. 55 : 88-94.

Janssen. M. P. M., Bruins, A.. DeVries. T. H. & Van Straalen, N. M.. 1991. — Comparison of cadmium kinetics in
four soil arthropod species. Arch. Environ. Contam. Toxicol.. 20 ; 305-312.

Source : MNHN, Paris

Cytochemistry of the Tergite Epicuticle of Glomeris
marginata (Villers) (Myriapoda, Diplopoda):
Preliminary Experimental Results

Philippe Compere , Stephane DEFISE & Gerhard GOFFINET

Universite de Liege, Laboratoire de Biologie generate et de Morphologic ultrastruciurale
Institul de Zoologie, 22 quai Ed. Van Benedcn. B-4020 Liege, Belgique.

ABSTRACT

The present study determines the ultrastructural location of chitin. proteins, and lipids in the eprcuticie of the diplopod
Glomeris marginata (Villers). The results lead to the conclusion that the cuticle includes two functionally different parts:
the upper part is involved in the permeability of the cuticle whilst the lower has mainly a mechanical role. The upper part
includes three epicuticular layers probably homologous to those described in insects: the cuticulin layer, the wax layer,
and the proteinaceous cement layer. The former seems to consist of a median leaflet of stabilized lipid polymers
sandwiched in two protein leaflets. This arrangement is assumed to be a primitive, general feature of the arthropod
cuticle, having been identified as the main waterproofing barrier in the cuticle of marine decapod crustaceans. The inner
epicuticle and the mineralised procuticle play a mechanical role. The inner epicuticle consists of a lipoprotein matrix
that surround rod-shaped protein elements and chitin-protein fibres which are probably of procuticular origin.
Structurally and functionally, it might be regarded as a structure convergent with that of decapod crustaceans, that plays a
part as a reinforcement to prevent the epicuticle splitting off from the mineralised exocuticle.

RESUME

Etude cytochimique de Fepicuticule des tergites de Glomeris marginata (Villers) (Myriapoda,
Diplopoda).

Lc present travail concerne la localisation ultrastruciurale de la chitine, des proteines et des lipides de fepicuticule du
diplopode Glomeris marginata (Villers). D’un point de vue fonctionnel. les resultats amenent a conclure que la cuticule
comprend deux parties difterentes : la partie superieure intervient dans la permeabilite de la cuticule tandis que la partie
inferieure joue un role essentiellement mecanique. La partie superieure comporte trois couches epicuticulaires de surface,
probablement homologues de celles dccrites chez les inscctes : la couche de cement proteinique, la couche de cire, et la
cuticuline. Cette demiere semble former un feuillet median de polym&res lipidiques stables pris en sandwich entre deux
feuillets proteiques. On admet que la cuticuline est un constituant primitif commun a toutes les cuticules d'arthropodes.
El le est reconnue comme la principale barriere impermeable de la cuticule des crustaces decapodes marins. Les couches
jouant un role mecanique sont fepicuticule interne et la procuticule mineralisee. L’epicuticule interne consiste en une
matrice lipoproteique entourant dcs elements proteiques en forme de batonnets el des fibres chitinoproteiques d origine
probablement procuticulaire. D un point de vue structural cl fonctionnel, cette constitution de fepicuticule interne peut
etre consideree comme une convergence avec celle rencontree chez les crustaces decapodes, en raison de son role de
renfort empechant fepicuticule de se detacher de fexocuticule mineralisee.

Compere. P.. DEFISE, S. & GOFFINET, G., 1996. — Cytochemistry of the tergite epicuticle of Glomeris marginata
(Villers) (Myriapoda, Diplopoda): preliminary experimental results. In: GEOFFROY, J.-J.. Mauries. J.-P. & Nguyen
Duy - JACQUEMIN, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist . nal ., 169 : 395-401. Paris ISBN : 2-85653-
502-X.

396

PHILIPPE COMPERE, STEPHANE DEFISE & GERHARD GOFFINET

INTRODUCTION

The arthropod cuticle is commonly regarded as an outer integumental structure acting as an
exchange surface in addition to providing mechanical protection. From an adaptive and
evolutionary point of view, the differentiation of several epicuticular layers with peculiar
structures and chemical composition has probably contributed to I he success of this group in
colonising a wide variety of environments.

As a result, the layers of epicuticle show a high degree of specialisation according to the
physiology of the integumental regions and to the habitat. For instance, it is well known that
cuticular lipids such as the surface waxes represent a barrier against water loss in terrestrial
arthropods such as insects and arachnids (HADLEY. 1981 ). In this respect, diplopods appear as
a very original and interesting group to study, since they are phylogenetically close to insects but
live in nearly the same wet microhabitats as terrestrial isopods and possess a mineralised cuticle
as it is the rule in numerous crustaceans. Little is known about the structure and especially the
chemical composition of the epicuticular layers.

Recent ultrastructural observations of ANSENNE el al. (1990), have shown that the
organisation of the cuticle of Glomeris marginata (Villers) fits the classical scheme known from
arthropods. In both the cuticle consists of a thin epicuticle overlying a thick, lamellate procuticle
which is subdivided into an exocuticle and an endocuticle. The cuticle is traversed by pore canals
and ducts of dermal glands. However, the epicuticle exhibits peculiar features whose
interpretation is critical (Fig. 1). The outermost epicuticular layers of diplopods with its
cuticulin, wax and cement layers are nearly identical in appearance to those of insects, but have
never been clearly identified. In addition, the structure of the inner epicuticle appears to be very
peculiar because microfibre-like elements that are arranged in a twisted plywood structure are
embedded in a matrix of medium electron density. Consequently, these structural peculiarities
raise important questions about the chemical nature of the epicuticle layers of diplopods in
relation to their roles in integument physiology and to their possible degree of homology or
analogy to corresponding structures in other arthropods.

The main purpose of this study was to determine the chemical nature of the epicuticle
components in the tergites of G. marginata, using cytochemical methods for the ultrastructural
demonstration of chitin, proteins, and lipids. The results are discussed with special reference to
the identity of the layers, their role in waterproofing or cuticle hardening, and their comparison
with the cuticles of terrestrial and aquatic arthropods.

MATERIAL AND METHODS

Individuals of Glomeris marginata (Villers) were collected on the University campus of the Sart Tilman, Liege. To
demonstrate chitin, ultrathin sections of glutaraldchydc-fixed. EDTA-decalcified and epoxy-resin-embedded tergites were
incubated for 45 min on drops of a WGA-BSA-gold complex (wheat germ agglutinin. Sigma) in 0.02 M Na-phosphate
buffer. pH 7.2 containing 0.5% BSA (HORISBERGER & Rosset, 1977).

Tannic acid in the fixative medium was used as an indicative reagent for proteins (Hayal, 1993). EDTA-
demineralised tergites were first fixed for 2 h at 20°C in a mixture of 1% tannic acid and 2.5% glutaraldehydc in 0.1 M Na-
phosphate buffer pH 7.4, followed by a 72-hours incubation in 1% tannic acid. Protein-bound tannic acid was then
revealed by "en bloc" uranyl acetate staining.

Two methods based on the reduction of OSO4 were used to demonstrate lipids. The first was to increase the
specificity of OsC>4 for unsaturated bonds in lipids under controlled experimental conditions (WlGGLESWORTH, 1981) and
blocking reactions. Prior to staining (1 h al 20°C in 1% Os04 in 0.1 M Na-phosphate buffer, pH 7.4) the glutaraldehyde-
fixed and EDTA-demineralised samples were treated for 4 h at 37°C in 1.25% N-ethylmaleimide-buffered solution for
blocking sulfhydryl groups (Gabe, 1968), then for 16 h at 20°C in nitrous acid (Lillie, 1954 in Gabe, 1968) or for 72 h
in 2.5% glutaraldehyde fixative solution for blocking primary amines and then for 1 h at 60°C in saturated bromine water
for blocking unsaturated bonds (Mukherji et al., 1960). Free and bound lipids were distinguished after extraction of free
lipids from glutaraldehyde-fixed material in a hot chloroform/melhanol mixture.

The second method was detection of hydrophobic substances. A highly unsaturated lipid-soluble marker,
niyrcene. was incorporated by partition in 50% ethanol, then revealed by reduction of OSO4 (Wigglesworth, 1981).
This treatment was performed after bromi nation.

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TF.RGITF. EPICUT1CULE OF GLOMERIS MARC, I NAT A (DIPLOPODA)

397

RESULTS

The WGA-BSA-gold complex which is used to demonstrate chitin labels the microfibres in
the procuticle and inner epicuticle (Fig. 2). In both layers, labelling depends on microfibre
orientation as determined by the twisted plywood arrangement of the microfibres. The gold
particles are distributed only along successive horizontal bands where the microfibres are seen in
oblique section and are absent where the fibres appear in longitudinal section.

The method used to demonstrate proteins, tannic acid treatment followed by “en bloc”
uranyl acetate staining, strongly enhances the electron density of the whole procuticle and of the
cement layer, the cuticulin layer, and the inner epicuticle (Figs 3 & 4). In contrast, the wax layer
remains completely electron-lucent. In the inner epicuticle, electron-dense rod-shaped elements
are prominent against the electron lucent material of the matrix. As observed after classical
staining (Fig. 1 ). these elements are oriented parallel to the microfibres.

The first method of lipid demonstration reveals unsaturated lipids in different epicuticular
layers and shows that they are insoluble in organic solvents (Figs 5 & 6). The lipids are mainly
located in the wax layer but also impregnate the cement layer and the upper leaflet of the cuticulin
layer, both of which appear as intensely electron-dense borders. The moderate electron density
of the lower leaflet of the cuticulin layer and inner epicuticle indicates that these layers are
relatively poor in such lipids. As a control, oxidation of double bonds by bromination prior to
0s04 staining prevents any osmiophilic reaction in the wax layer but merely reduces the contrast
in the other epicuticular layers (Fig. 7). This remaining osmiophily is probably due to the
presence of proteins. The presence of lipids in the epicuticular layers of G. marginata is
confirmed by the results of the second procedure for detecting hydrophobic substances.
Incorporation of myrcene after bromination and before 0s04 staining restores a high electron
density in the previously osmiophilic layers, i.e. the cement layer, the wax layer, and the upper
leaflet of the cuticulin layer (Figs 8 & 9). However, it only slightly enhances the contrast of the
lower leaflet of the cuticulin layer and the matrix material of the inner epicuticle, which seem to
consist of lipoproteins. In contrast, the rod-shaped protein elements and the chitin microfibres
remain electron-lucent.

DISCUSSION

The present cytochemical results combined with the previous ultrastructural observations
of ANSENNE el al. (1990) allow accurate identification of the structural components of Glomeris
marginata tergite epicuticle. Furthermore, considering the respective structures, chemical
compositions, and roles of its constituents, the cuticle of G. marginata is consistent with the
functional model recently proposed by COMPERE & GOFFINET (1992) for decapod crustaceans.
According to this model, the cuticle includes two functional parts: the upper part is responsible
of the integument permeability characteristics while the lower part contributes to the mechanical
resistance of the exoskeleton. In G. marginata, the upper part includes the three outer
epicuticular layers, i.e. the cement layer, the wax layer, and the cuticulin layer. The cement and
wax layers can be regarded as integument adaptations to a terrestrial mode of life and as
structures homologous to the corresponding layers of insect and arachnid cuticles (NEVILLE.
1975; FlLSHIE, 1976; HADLEY, 1986). This view is strongly supported by their structure,
location, chemical composition, and role in cuticular waterproofing in addition to the fact that the
cement layer of G. marginata is discharged on the cuticular surface by dermal gland ducts
(ANSENNE et al., 1990), similar as in insects.

Since it labels both protein- and lipid-positive, the cement layer appears as an outer
protection layer made of wax-impregnated proteins. Overlying the cuticulin layer, the wax layer
consists exclusively of stabilized lipid compounds, most of them unsaturated. Similar solvent-
resistant waxes have been reported for some insects (WlGGLESWORTH, 1985). for some
arachnids (HADLEY, 1981), and for the diplopod Orthoporus ornatus (Girard) (WALKER &

398

PHILIPPE COMPERE. STEPHANE DEFISF. & GERHARD GOFFINET

CRAWFORD. 1980). More recently, the presence of a strongly osmiophilic wax layer was also
described in Ophyiulus pilosus (Newport) (THOREZ et al., 1992). The higher resistance of
G. marginata to a dry environment compared to the poor water resistance of the isopod Oniscus
asellus (L.) (EDNEY, 1951), could be ascribed to the absence of this layer in O. asellus, which
only possesses free lipids inside the cuticulin layer instead of a distinct outer wax layer
(COMPERE, 1990).

Although the accurate identification of the cuticulin layer components is still lacking, the
present observations combined with those of ANSENNE et al. (1990) strongly suggest that the
structure of this layer is comparable to that described for other arthropods (insects: LOCKE,
1966; arachnids: FlLSHIE, 1976; crustaceans: COMPERE, 1988, 1990). The cuticulin layer of
G. marginata exhibits a membrane-like structure approximately 20 nm thick which seems to
consist of two protein leaflets, the upper one being impregnated with overlying waxes. These
leaflets are separated by an electron-lucent layer probably made of lipid polymers, as suggested
by the studies of WlGGLESWORTH (1985) and HACKMAN (1986) for insects and COMPERE &
GOFFINET (1992) for the crab Carcinus maenas (L.). Appearing as a general arthropodian
feature, the cuticulin layer can be regarded as a primitive structure, since it already constitutes the
main permeability barrier of the cuticle in marine crustaceans (COMPERE & GOFFINET, 1992).

The inner epicuticle and procuticle are layers of the lower part which probably contribute to
the mechanical properties of the cuticle. The inner epicuticle consists of a lipoprotein matrix
surrounding proteinaceous rod-shaped elements and chitin-protein microfibres that prolong the
helicoidal twisted plywood arrangement of the exocuticle fibres. This organisation appears very
peculiar, never having been reported before in any other arthropod cuticle. The presence of
chitin-protein fibres contradicts the first and classical definition of the insect epicuticle as a non-
chitinous layer (KUHNELT, 1928a,b). Considering that the inner epicuticle of the soft
intersegmental regions of G. marginata (COMPERE, unpublished results) is composed, as in
insect tergites (LOCKE, 1969; NEVILLE, 1975), of a homogeneous fibreless matrix, the thick
inner epicuticle of the mineralised tergites of G. marginata can be interpreted as a result of close
interpenetration between the epicuticular matrix material and the chitin-protein fibres of the
exocuticle. Functionally, it can be regarded as a structure convergent with the inner epicuticle of
decapod crustaceans, in addition to other features such as the pseudo-reticulate pattern of the
exocuticle and mineralisation of the procuticle (ANSENNE et al, 1990). As proposed by
COMPERE & GOFFINET (1992) for the crab C. maenas, the thick fibrous inner epicuticle bearing
roots on its lower side ensures mechanical reinforcement, preventing the upper layers and the
mineralised exocuticle from splitting off.

Fig. 1. — Transverse section through the tergile epicuticle of Glomeris marginata after classical tissue fixation and
uranyl acetate/lead citrate section staining, c, cuticulin layer; ct, cement layer; ie. inner epicuticle; rs, rod-shaped
elements; w, wax layer;

> , cross-sectioned chitin-protein microfibres; 0/^/0, microfibre orientation. Bar 5 pm.

Fig. 2. — Transverse section of the tergite epicuticle of Glomeris marginata after incubation with the WGA-BSA-gold
complex and stained in uranyl acetate, ex, exoculicle; ie, inner epicuticle; O/=/0, fibre orientation. Bar 0.5 pm.
Figs 3 & 4. — Detail of the upper epicuticular layers after “en bloc” uranyl acetate staining (Fig. 4) and after exposure to
tannic acid prior to “en bloc” uranyl acetate staining (Fig. 3). c, cuticulin layer; ct, cement layer; ie, inner
epicuticle; rs, rod-shaped elements; w. wax layer. Bars 250 nm.

Figs 5-9. — Vertical sections of the tergite epicuticle of Glomeris marginata after different cytochemical treatments, c,
cuticulin layer; ct. cement layer; ex, cxocuticlc; ie, inner epicuticle; rs, rod-shaped elements; w, wax layer.

Figs 5-6. After extraction of free lipids in a chloroform/methanol mixture, long fixation in buffered 2.5%
glutaraldehyde and OSO4 staining. Fig 5. Bar I pm. Fig. 6. Detail of the upper epicuticular layers. Bar 250 nm.
Fig. 7. Detail of the upper epicuticular layers after extraction of free lipids in a chloroform/methanol mixture,
incubation in bromine water and OSO4 staining. Bar 250 nm.

Figs 8-9. — After extraction of free lipids in a chloroform/methanol mixture, incubation in bromine water,
exposure to myreene and 0$04 staining. Fig. 8. Bar 1 pm. Fig. 9. Detail of the upper epicuticular layers. Bar 250
nm.

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rERGITE EPICUTICULE OF GLOMER1S MARGIN AT A < DIPLOPODA)

399

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400

PHILIPPE COMPERE. STEPHANE DEFISE & GERHARD GOFFINET

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TERGITE EP1CUTICULE OF GLOMERIS MARGINATA (DIPLOPODA)

401

CONCLUSION

In conclusion, the present study sheds some light upon the relationships between the
different cuticle layers in diplopods, their role in integumental physiology, and their ecological
significance. In addition, our evidence supports the dual-function model of the cuticle, recently
defined by COMPERE & GOFFINET (1992) for a marine decapod crustacean. This model also
agrees with previous observations made on the cuticle of insects and arachnids (for reviews, see:
NEVILLE, 1975; Hadley, 1981, 1984. 1986). we tentatively propose it as more general for
arthropod cuticular structure.

ACKOWNLEDGEMENTS

The authors are indebted to the Belgian Fund for Joint Basic Research for its financial support of this work (F.R.F.C..
convention N°2.4527.89). P.C. is the recipient of a grant front the National Fund for Scientific Research (F.N.R.S..
Belgium).

REFERENCES

Ansenne, A., Compere . P. & Goffinet. G.. 1990. — Ultrastruciural organization and chemical composition of the
mineralized cuticle of Glomeris marginata (Myriapoda. Diplopoda). In : A. Minelli. Proc. 7th Intern. Congr. of
Myriapodology ., Leiden, Brill : 125-134.

Compere, P.. 1988. — Mise en place de 1'epicuticule chez le crabe Carcinus maenas . Aspects recents de la Biologie des
Crustaces. Actes de Colloques IFREMER , 8 : 47-54.

Compere. P.. 1990. — Fine structure and elaboration of the epicuticule and pore canal system in tergite cuticle of the
land isopod Oniscus asellus during a molting cycle, hi : P. JuChault & J. P. Mocquard. Biology of terrestrial
Isopods III. Proc. 3rd hit. Symp. Biology terrestrial Isopods , Poitiers : 169-175.

Compere, P. & Goffinet, G.. 1992. — Organisation tridimensionnelle et cytochimie de 1'epicuticule et des systemes
canaliculaires des sclcrites du crabe Carcinus maenas (Crustace decapode). Mem. Soc. r. beige Ent.. 35 : 715-720.

Edney, E. B., 1951. — The evaporation of water from woolice and the millipede Glomeris. J. exp. Biol., 28 : 91-1 15.

FlLSHiE, B. K.. 1976. — The structure and deposition of the epicuticle of the adult female cattle tick ( Boophilus
microplus). In : H. R. Hepburn, The Insect Integument. Amsterdam, Elsevier : 193-206.

Gabe, M., 1968. — Techniques histologiques. Paris. Masson, 1113 pp.

Hackman, R. H., 1986. — The chemical nature of the outer epicuticle from Lucilia cuprina larvae. Insect Biochem .,
16 : 91 1-916.

Hadley. N. F., 1981. — Cuticular lipids of terrestrial plants and Arthropods: a comparison of their structure,
composition and waterproofing function. Biological Rev.. 56 : 23-47.

Hadley. N. F.. 1984. — Arthropoda : Cuticle : Ecological Signifiance. In : Bereiter-Hahn, A. G. Matoltsy & K. S.
Richards. Biology of the Integument, I. Invertebrates, Berlin, Springer Verlag : 685-693.

Hadley. N. F., 1986. — La cuticule des Arthropodes. Pour la Science, 107 : 64-72.

Hayat, M. A., 1993. -- Stains and cytochemical methods. Plenum Publ. Corp., New York, London, 445 pp.

Horisberger, M. & Rosset, J.. 1977. — Colloidal gold, a useful marker for transmission and scanning electron
microscopy. J. Histochem. Cytochem., 25 : 295-305.

Kuhnelt, W., 1928a. — Ein Beitrag zur Histochemic des Insektenskelettes. Zool. Anz.. 75 : 111-115.

KUHNELT, W., 1928b. — Studien iiber den mikrochemischen Nachweis des Chitins. Biol. Zeniralbl., 48 : 374-382.

Locke, M.. 1966. — The structure and formation of the cuticulin layer in the epicuticle of an insect, Calpodes ethlius
(Lepidoptera, Hesperiidae). J. Morph.. 118 : 416-494.

Locke, M., 1969. — The structure of epidermal cells during the development of the protein epicuticle and uptake of
molting fluid in an Insect. J. Morph., 127 : 7-40.

M ukherji, M., Df.b, C. & Sen, P. B.,1960. — Histochemical demonstration of unsaturated lipids by bromine silver
method. J. Histochem. Cytochem.. 8 : 189.

Neville, A. C., 1975. — Biology of the arthropod cuticle. In : W. S. Hoar. J. Jacobs. H. Lanc.er & M. Lindauer,
Zoophysiology and Ecology, Vol. 4/5. Berlin, Springer Verlag, 448 pp.

Thorez, A., Compere, P. & Goffinet. G., 1992. — Ultrastructure and mineral composition of the tergite cuticle ol the
iulid millipede Ophyiulus pilosus (Myriapoda, Diplopoda). Ber. nat.-med. Verein Innsbruck , suppl. 10 : 63-69.

Walker, L. J. & Crawford, C. S., 1980. — Integumental ultrastructure of the desert millipede, Orthoporus ornatus
(Girard) (Diplopoda : Spirostreptidae). hit. J. Insect Morphol. Embryol..9 : 231-249.

Wigglesworth, V. B„ 1981. — The distribution of lipid in the ceil structure: an improved method for the electron
microscope. Tissue Cell , 13 . 19-34.

Wigglesworth, V. B., 1985. — Sclerotin and lipid in the waterproofing of the insect cuticle. Tissue Cell, 17 : 227-
248.

Source : MNHN. Paris

Coxal Organs of Chilopoda: the Exocrine Glands in

Lithobius forficatus

Jdrg Rosenberg * & Hartmut Greven **

* Institut fur Tierphysiologie, Fakultiit fur Biologie, Ruhr-Universitat Bochum, D-44780 Bochum, Germany
** Institut fur Zoologie (Zoomorphologie und Zellbiologie) der Heinrich-Heine-Universitat-Diisseldorf
UniversitatsstraBe 1. D-40225 Dusseldorf, Germany

ABSTRACT

The exocrine glands within the coxal organs of Lithobius forficatus are described. Each gland consists ot secretory
cells, an additional cell and a canal cell, forming a cuticular ductule (class 3 gland according to NoiROT &
Quennedy, 1974). Secretory cells are rich in rER and PA-TCH-SP (periodic acid-thiocarbohydrazide-silver proteinate)-
positive secretory granules. The additional or intercalary cells possess numerous mitochondria and prominent infoldings
of the plasma membrane beneath the cuticle of the transport duct. PA-TCH-SP-positive secretory products are obviously
discharged along the cuticular ductule into the pore channel of the coxal organ, forming a mucous layer that covers the
specialized cuticle of the transport epithelium. Within its subcuticle, chloride can be localized cytochemically; its
accumulation in the mucous layer is not significant.

RESUME

Organes coxaux des Chilopodes : les glandes exocrines de Lithobius forficatus.

Les glandes exocrines des organes coxaux de Lithobius forficatus sont decrites. Chaque glande est composee de cellules
secretrices, d’une cellule additionnelle et d’une cellule-canal, constituant un canalicule cuticulaire [glande de classe 3
selon Noirot & QUENNEDY (1974)]. Les cellules secretrices sont riches en rER ct en PA-TCH-SP (“periodic acid-
thiocarbohydrazide-silver proteinate”) sous forme de granules de secretion. Les cellules additionnelles ou intercalaires
possedent de nombreuses mitochondries et des protuberances en doigts de gant de la membrane du plasma en dessous de la
cuticule du canalicule. Les produits secretes (PA-TCH-SP-positif) sont evacues le long du conduit cuticulaire jusqu’au pore
de Porgane coxal, formant une couche de mucus qui couvre la cuticule specialist de fepithtHium de transport. A
finterieur de sa sous-cuticule, les chlorures peuvent etre localises cytochimiquement ; leur accumulation dans la couche
de mucus n’est pas significative.

INTRODUCTION

Coxal organs of Chilopoda are complex and possibly multifunctional structures. They are
localized on the coxae of the last trunk segment (Geophilomorpha, Scolopendromorpha) or last
four trunk segments (Lithobiomorpha) and characterized by numerous pores, each leading into a
cuticle-lined pore channel surrounded by a columnar single-layered transport epithelium and -
arranged like a collar - junctional cells and several exocrine glands pouring out their secretory
products into the lumen of the pore channel. This secretion covers the specialized cuticle of the
transport epithelium (for review see ROSENBERG, 1985).

Rosenberg, J. & Greven, H„ 1996. — Coxal organs of Chilopoda: the exocrine glands in Lithobius forficatus.
In. Geoferoy. J.-J., Mauri6s. J.-P. & Nguyen Duy - Jacquemin, M„ (eds). Acta Myriapodologica. Mem. Mus. natn.
Hist. not.. 169 : 403-409. Paris ISBN : 2-85653-502-X.

404

JORG ROSENBERG & HARTMUT GREVEN

The general ultrastructure of the coxal organs in Chilopoda (ROSENBERG, 1982, 1983a, b,
1984. 1990) and also in experimental studies, particularly those on Lithobius forficatus, suggest
that they are involved in water vapour uptake from the environment (ROSENBERG, 1985;
ROSENBERG & BAJORAT, 1984). More recently, some evidence has been accumulated
suggesting that coxal organs of Lithobiomorpha release a sex-specific pheromone
(LlTTLEWOOD, 1988, 1991; LlTTLEWOOD & BLOWER, 1987). The authors speculate that sub-
epithelial blood cells beneath the coxal organs might synthesize this pheromone, which moves
across the epithelium of the coxal organ. These assumptions prompted us to examine the
“glandular system” of the coxal organs, especially as recent investigations have revealed
numerous small epidermal glands close to the coxal pores (ROSENBERG, 1994). On principle
such structures could act as pheromone glands.

The following note deals with the exocrine glands of the coxal organs in Lithobius
forficatus which were not described by LlTTLEWOOD (1983) in Lithobiomorpha.

MATERIALS AND METHODS

Coxae of adult Lithobius forficatus were fixed as described previously (Rosenberg 1983a).

For detecting “mucosubstances”, animals were fixed with 2.5% glutar dialdehyde and 2% formaldehyde (freshly prepared
from paraformaldehyde) in phosphate buffer without any postfixation. After graded ethanol dehydration, tissue was
embedded in LR White (London Resin Co.). Ultrathin sections were mounted on formvar-coated Ni grids. The periodic
acid-thiocarbohydrazidc-silvcr proteinate (PA-TCH-SP) reaction was performed as described by Neiss (1988). The
sections remained unstained.

For demonstration of chloride the coxae were fixed according to Wichard & Komnick (1973) with osmium
tetroxide and silver lactate and treated with nitric acid during dehydration.

Sections were examined in a Zeiss 109 T electron microscope.

RESULTS

The topography of the exocrine glands within the coxal organs of Lithobius forficatus has
been described elsewhere (ROSENBERG, 1983a, 1985). Each exocrine gland consists of three
types of cells: secretory cells with well-developed granular ER, an additional or intercalary cell,
forming microvilli-like projections surrounding a cuticular ductule, and a canal cell, whose
cuticular duct runs into the pore channel (Figs 1 & 5).

The secretory cells are spheroidal or spindle-shaped. Their plasma membrane is
moderately infolded. Cells are connected with adjacent additional cells by septate desmosomes.
The cytoplasm of most secretory cells is packed with stacks of granular endoplasmic reticulum,
which is filled with a fineley particulate substance (Fig. 3). In addition, these cells contain
numerous dictyosomes in different stages of development and numerous electron dense
secretory granules, which are membrane bounded and vary widely in size. Small mitochondria,
and some lysosomes and multivesicular bodies, are distributed randomly throughout the
cytoplasm. Each secretory cell has a large, lobate nucleus; its chromatin is mostly located
peripherally. The thin cuticular ductule of the secretory cell is continuous with the wall of the
transport duct of the additional cell.

PA-TCH-SP reaction reveals precipitations at the margins of secretory granules and in
some profiles, derived from endoplasmic reticulum (Fig. 6).

The additional duct forming cell is more elongated and its cytoplasm is lighter than that of
the secretory cells or adjacent cells of the transport epithelium. Large profiles of endoplasmic
reticulum and dictyosomes are absent, and the small nucleus is oval. The plasma membrane
surrounding the cuticular duct is folded forming long microvilli-like projections (Fig. 4).
Mitochondria are numerous along the apical infoldings, they are larger than in the secretory cell.
The wall of the cuticular duct is continuous with the cuticle of the small canal cell and that of the
epicuticle of the pore channel of the coxal organ (Fig. 5). As seen by SEM (Fig. 2) and TEM
(Fig. 1), secretion is deposited as a distinct mucous layer on the cuticle of the transport
epithelium.

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EXOCRINE GLANDS OF A LITHOBIIDAE

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Pig. I. — Section of two coxal pores of Lithobius forficaius. showing the mucous layer (ml) on the bottom of the pore
channel and several openings of epidermal glands (circles) around the coxal pores. Scale line, 20 pm.

Fig. 2. — Part of the coxal organ of Lithobius forficaius with the transport epithelium (te) and its specialized cuticle,
covered by the mucous layer (arrow), and the exocrine gland with secretory cells (sc) and an additional cell (ac)
with its cuticular duct (*). p pore channel. Scale line. 0.08 pm.

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406

JORG ROSENBERG & H ARTMUT GREVEN

Fig. 3. — Secretory cells ol the exocrine gland of the coxal organ with stacks of ER, dictyosomes, and secretory
products. Scale line, 0.08 pm.

Fig. 4. Additional cell ot the exocrine gland of the coxal organ. The cuticular ductule (*) is surrounded by infoldings of
the apical plasma membrane, m: mitochondrium. Scale line, 0.08 pm.

Fig. 5. — Additional cell (ac) and canal cell (cc) of the exocrine gland of the coxal organ. Their cuticular ductule (*) opens
into the pore channel (p) of the coxal organ. Scale line. 0.1 pm.

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EXOCRINE GLANDS OF A LITHOBUDAE

407

Within the accessory cells (intercalary and canal cell), treatment by PA-TCH-SP stains the
content of the transport duct, its cuticle, the material underlying the cuticle of the duct and the
space between the microvilli-like projections (Fig. 7). A positive reaction is also seen in the
mucous layer, covering the modified cuticle of the main epithelium (Fig. 8).

In coxal organs, fixed in the osmium-silver-lactate mixture, coarse precipitates are
localized predominantly in the subcuticle of the specialized cuticle, covering the transport
epithelium (Fig. 9). Fine precipitations seem to be scattered within the overlying endocuticle and
in the mucous layer (Fig. 10). No precipitates are found within the cells of the transport
epithelium, within the cuticle of the pore channel (Fig. 9), or within the cells of the exocrine
gland.

DISCUSSION

The exocrine glands described for the coxal organs of Lithobius forficatus can be
characterized as “class 3 glands” according to NOIROT & QUENNEDY (1974). In a simple case “a
cuticular ductule or canal penetrates the gland cell and the canal runs into a ductule or canal cell
which has secreted it” (p. 63). Within the exocrine glands of Lithobius forficatus, this
description is complicated by the presence of an additional or intercalary cell between the two
others. In Lithobius forficatus “class 3 glands” appear to be common; they are known to be
present adjacent to the telopodal glands and are associated with sensilla trichodea (KEIL, 1975)
as well as with the organs of Tomosvary (TlCHY, 1973). It is assumed that these glands,
although similar in organization, produce substances of different chemical composition and
significance.

Considering the ultrastructure of secretory cells, in particular the abundant rough
endoplasmic reticulum and the electron dense granules, there is reason to believe that secretory
products contain a considerable amount of proteins. However, well-developed dictyosomes and
the positive reaction after PA-TCH-SP are indicative of a carbohydrate component
(“mucosubstances”). PA-TCH-SP positive material has also been demonstrated within the
transport ductule and the mucous layer covering the specialized cuticle of the transport
epithelium.

Location of the exocrine glands suggests discharge of secretory products through transport
ductules into the pore channel of the coxal organ. As seen by SEM and TEM, the secretion
spreads over the modified cuticle of the transport epithelium (LlTTLEWOOD, 1983; ROSENBERG,
1983a) and fills up the bottom of the pore channel. Material that is electron-dense to varying
degrees, interpreted as the mucous layer, has been observed in all coxal organs hitherto
examined (ROSENBERG, 1985). It stains with PA-TCH-SP in Lithobius forficatus , but also in
Cryptops hortensis (Scolopendromorpha; ROSENBERG, 1983b). It was suggested that this
mucus consists of a hygroscopic material, which would gather water vapour from moist air
(ROSENBERG, 1983a, 1985; ROSENBERG & BAJORAT, 1984).

Apart from forming the transport ductule, the role of the additional cell is unclear. Enlarged
surfaces and abundant mitochondria in these cells suggest transporting ability, perhaps to modify
secretion products within the ductule.

Localization of chloride in the modified cuticle of the coxal organ has been regarded as an
indication of transepithelial solute transport as it is in the “chloride cells" of other transporting
systems (e.g. anal papillae, anal organs, ventral tube, coxal vesicles) in a variety of insects
(WICHARD & KOMNICK. 1973; KOMNICK, 1977; ElSENBEIS, 1 976; ElSENBEIS & WlCHARD,
1975). Accumulation in the mucous layer, however, seems not to be significant.

In general, the results presented here neither contradict the assumed uptake of water
vapour from the air by coxal organs nor the pheromone release. However, there is still no
definite proof for a hygroscopic capacity of the mucous layer; the epidermal glands close to the
coxal pores are the presumed site of pheromone production (ROSENBERG, 1994).

408

JORG ROSENBERG & HARTM UT GREVEN

Pig. 6. PA- TCH-SP-reaction in secretory cells (unstained sections): reaction products are visible at the margin of the
secretory granules and in profiles of the ER (arrow). Scale line, 0.04 pm.

Pig. 7. PA-TC H-SP-reaction in additional cell (unstained section): reaction products are visible within the cuticular
duct (*) and the space underlying the duct, and between the microvillar infoldings (arrows). Mitochondrium
(arrowhead). Scale line, 0.05 pm.

Fig. 8. PA-TCH-SP-reaction in mucous layer (unstained section): the reaction products are only visible within the
mucous layer (ml). C cuticle of the transport epithelium. Scale line, 0.03 pm.

Fig. 9. — Chloride-reaction in cuticle of the transport epithelium: a part of the coxal organ is shown with the transport
epithelium (te) and the cuticle (c) ot the pore channel (p). Reaction products (arrow) are only visible within the
subcuticle of the main epithelium. Scale line. 0.1 pm.

Pig. 10. Chloride-reaction in cuticle of the transport epithelium: coarse reaction products are localized within the
subcuticle (su); fine precipitations are scattered within the endocuticlc (en) and in the mucous layer (arrow) Scale
line, 0.03 pm.

Source : MNHN. Paris

EXOCRINE GLANDS OF A LITHOBIIDAE

409

ACKNOWLEDGMENTS

We would like to thank Mr. H. Schlierenkamp for careful technical assistance.

REFERENCES

Eisenbeis, G., 1976. — Zur Feinstruktur und Histochemie des Transportepithels abdominaler Koxalblasen der
Doppelschwanz-Art Campodea staphylinus (Diplura: Campodeidae). Ent. Germ., 3 : 185-201.

Eisenbeis. G. & WlCHARD, W., 1975. — Feinstruktureller und histochemischei Nachweis des Transportepithels am
Ventraltubus symphypleoner Collembolen (Insecta, Collembola). Z. Morph. Tie re, 81 : 103-110.

Keil, T., 1975. — Die Antennensinnes- und Hauldrusenorgane von Lithobius forficatus (L ). Eine licht- und
elektronenmikroskopische Uniersuchung. Dissertation. University of Berlin.

Komnick, H., 1977. — Chloride cells and chloride epithelia of aquatic insects, hit. Rev. Cytol.,49 : 285-327.

Littlewood, P. M. H., 1983. — Fine structure and formation of the coxal glands of Lithobiomorph centipedes:
Lithobius forficatus (L.) and Liihobius crassipes Koch (Chilopoda. Lithobiomorpha). J. Morphoi. Ill : 157-180.

LITTLEWOOD, P. M. H., 1988. — The chemosensory behaviour of Liihobius forficatus (Myriapoda: Chilopoda). 2.
Bioassay and chemistry of the coxal pheromone. J. Zool. . London . 215 : 523-535.

Littlewood, P. M. H., 1991. — Chilopod coxal organs: morphological considerations with reference to function. J.
Zool.. London . 223 : 379-393.

LITTLEWOOD, P. M. H. & Blower. J. G., 1987. — The chemosensory behaviour of Lithobius forficatus. 1. Evidence for a
pheromone released by the coxal organs (Myriapoda: Chilopoda). J. Zool.. London. 211 : 65-82.

NEISS, W. F., 1988. — Enhancement of the periodic acid-Schiff (PAS) and periodic acid-thiocarbohydrazide-silver
proteinate (PA-TCH-SP) reaction in LR White sections. Histochemistry. 88 : 603-612.

NoiROT, C. & Quennedey, A.. 1974. — Fine structure of insect epidermal glands. Annual Rev. Entomology, 19 : 61-80.

Rosenberg. J., 1982. — Coxal organs in Geophilomorpha (Chilopoda). Organization and fine structure of the
transporting epithelium. Zoomorphology. 100 : 107-120.

Rosenberg, J.. 1983a. — Coxal organs of Lithobius forficatus (Myriapoda, Chilopoda). Fine-structural investigation
with special reference to the transport epithelium. Cell Tiss. Res., 230 : 421-430.

ROSENBERG. J., 1983b. — Coxal organs in Scolopendromorpha (Chilopoda): Topography, organization, fine structure
and signification in centipedes. Zool. Jb. ( Anal .), 110 : 383-393.

ROSENBERG. J., 1984. — Ultrastructure of the anal organs in the larval stages of Lithobius forficatus (L.) (Chilopoda:
Lithobiomorpha). hit. J. Insect Morph. Embryol. , 13 : 29-35.

Rosenberg, J., 1985. — Untersuchungen zur feinstrukturellen Organisation und Funktion der Coxalorgane und
Analorgane bei Chilopoda. Bijdr. Dierk. .55 : 181-189.

Rosenberg, J., 1990. — Untersuchungen zur funktionellen Morphologie der Analorgane von Geophilidae
(Geophilomorpha). hi : A. MlNELLl, Proc. 7th. Intern. Congr. Myriapodology . Leiden, Brill : 115-123.

Rosenberg, J., 1994. — Fine structure of epidermal glands in vicinity to the coxal organs of Lithobius forficatus
(Chilopoda). Acta Biol. Benrodis. 6 : 37-47.

ROSENBERG. J. & Bajorat. K. H., 1984. — EinfluB der Coxalorgane von Liihobius forficatus (L.) (Chilopoda) auf die
Sorption von Wasserdampf. Zool. Jb. (Physiol.) , 88 : 337-344.

Tichy, H., 1973. — Untersuchungen liber die Feinstruktur des Tbmdsvaryschen Sinnesorgans von Liihobius forficatus
(L.) (Chilopoda) und zur Frage seiner Funktion. Zool. Jb. (Anal.) . 91 : 93-139.

WlCHARD, W. & Komnick, IT, 1973. — Fine structure and function of the abdominal chloride epithelia in caddisfly
larvae. Z. Zellforsch., 136 : 579-590.

Source : MNHN, Paris

The Phenoloxidase from the Hemolymph of Diplopoda

Willi E. R. XY LANDER

Institut fur Allgemeine und Spezielle Zoologie, Justus-Liebig-Universitat Giessen
Siephanstr. 24, D - 35390 Giessen, Germany

ABSTRACT

The phenoloxidases of the diplopods Chicobolus sp. and Rhapidostreptus virgator occur in the hemolymph as
proenzyme, prophenoloxidase (proPO). It can be activated in vitro by incubation with ethanol or methanol whereas
chymotrypsin only activates the proPO of Rhapidostreptus. Microbial substances have little effect on the proPO of both
species. Dopa showed to be a good substrate, dopamine and pyrogallol were less, tyrosine and others hardly converted at
all. Phenylthiourea is a potent inhibitor. The pH optimum was about pH7 in both species. In SDS-PAGE the proPO had a
molecular weight of about 230 kDa in both species and in Lithobius forficatus. The proPO is located in the grana of the
two types of granular hemocytes but not in the plasmatocytes. Microbial substances and other material (glass, sephadex.
latex beads) did not induce exocytosis and activation of the proPO but it is activated during wound clot formation (as
shown by mclanisation of the clot). Thus the activation mechanism of this defence system is somewhat different from
that of insects and crustaceans and may be involved in antimicrobial defence at wound margins.

RESUME

La phenoloxydase de l'hemolymphe des diplopodes.

La phenoloxydase des diplopodes Chicobolus sp. el Rhapidostreptus virgator existe dans 1 hemolymphe en tant que
proenzyme de la phenoloxydase (proPO). Elle peut etre activee in vitro par I'&hanol et le methanol, tandis que la
chymotrypsine active uniquement celle de Rhapidostreptus. Les substances microbiennes ont tres peu deffet sur les
proPO des deux especes. La dopa s'av^re etre un bon substrat. tandis que la dopamine et le pyrogallol le sont moins. En
revanche, la tyrosine et d'autres substances ne sont pour ainsi dire pas metabolisees. La phenylthiouree convient
parfaitement commc substance inhibitrice. La valeur optimale du pH est pH7. Dans le SDS-PAGE, la proPO des deux
esp&ces, de memc que celle de Lithobius forficatus , a un poids moleculaire d’environ 230 kDa. Elle est surtout localisee
dans les grana des deux types d'hemocytes granulaires. mais elle manque dans les plasmocytes. Des substances
microbiennes el d’autres mat£riaux (verre, grains de sephadex et de latex), ne provoquent pas 1 exocytose des grana
contenant la proPO mais sont actives pendant la cicatrisation (comme on peut le depister dans la melanisation).
Apparemment, les mecanismes d'activation de ce systfcme immunologique sont differents de ceux des insectes et des
crustacSs ; ils sont probablement impliques dans la defense antimicrobienne qui se developpe au bords des plaies.

INTRODUCTION

The phenoloxidase (PO) from the hemolymph of arthropods and the enzymes involved in
its activation play an important role in immune defence responses. In many arthropods, they are
stored in the hemocytes as inactive zymogen, the prophenoloxidase (proPO). Once set free by
exocytosis the enzyme becomes “sticky” and attaches to the surface of foreign particles where
the activation of the proPO and subsequent formation of melanin occurs (SODERHALL, 1%2).

XYLANDER. W. E. R., 1996. — The Phenoloxidase from the hemolymph of Diplopoda. In: Geoffroy, J.-J
M AURifes, J.-P. & Nguyen Duy - Jacquemin, M„ (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, not., 169 : 41 1
420. Paris ISBN : 2-85653-502-X.

412

WILLI E. R. XYLANDER

The immune defence reactions in which the hemolymph PO is involved comprise synthesis of
bacteriostatic and fungicide intermediate products of melanin formation, opsonization, melanin
deposition on the surfaces of metazoan and protozoan parasites, and wound closure (ASHIDA &
Yamazaki. 1990; Chadwick & ASTON, 1978; Gotz& Vey, 1974; Pye, 1974; RATCLIFFE et
al., 1984; Vey & GOTZ. 1975). This paper presents the results of investigations on the
hemolymph PO of two diplopods ( Rhapidostreptus virgator and Chicobolus sp.).

MATERIAL AND METHODS

Two diplopods (Chicobolus sp. and Rhapidostreptus virgator) were reared and hemolymph was obtained
following the description given by Xylander & Nevermann (1990), The methods of PO investigations in the in-vitro-
system (see Fig. 1). the calculation of PO-activity and the preparation of SDS-PAGE under non-reducing conditions were
described in detail by Xylander & Bogusch (1992). The procedure of demonstration of intracellular proPO in the
hemocytes was presented by Xylander & Nevermann (1993).

measurement of extinction
at *190 nm for
30 min or lh

Fig. I. — Procedure of in-vitro-measurement of PO activity.

RESULTS

In-vitro-activity

n ac rTt]e in'vitr°'actiyity °f the phenoloxidase (PO) without any activator is comparatively low
0.65 Lml-1 min-i] in Rhapidostreptus (XYLANDER & BOGUSCH, 1992) and 0.133 [ml-i min-il
in Chicobolus (Fig. 2). This means that the PO occurs in the hemolymph as inactive zymogen,
the piophenoloxidase (proPO). The level of PO-activity in the hemolymph depended oifthe
season; it was generally higher in summer (during the main period of activity) than in winter

Source : MNHN, Paris

PHENOLOX I DASE FROM THE HEMOLYMPH OF DIPLOPODA

413

File proPO can be activated by incubation of hemolymph with organic solvents (e. g.
ethanol, methanol, SDS) and proteases (a-chymotrypsin) for 10 min; maximum activity (the
highest activity measured) was generally higher in Rhapidostreptus than in Chicobolus,
however, the relative increase in activity after activation (activity without activation = 1) was
higher in Chicobolus (compare Figs 2 & 3). Ethanol is the best activator tested and raises the
PO-activity 20times (in Rhapidostreptus ) to 35times (in Chicobolus ; A = 7.42; see Figs 2 & 3).
Methanol is a less efficient activator than ethanol in both species, a-chymotrypsin is a good
activator in Rhapidostreptus but shows only little effect in Chicobolus (Figs 2 & 3).

Fig. 2. — Absolute in-vitro-activity of the
hemolymph of Rhapidostreptus
virgator and Chicobolus sp. without
and with application of various
potential activators.

moximum activity

Rhapidostreptus Chicobolus

without activ.

ethanol

Wilt methanol

(WVW

t.'-ii-J zymosan

bact. LPS

. - 1 chymotrypsin

Fig. 3. — Relative in-vitro-activity of the
hemolymph of Rhapidostreptus
virgator and Chicobolus sp. without
and with application of various
potential activators, (activity
without activator = 1). Note that the
relative activity increase in
Chicobolus is higher than in
Rhapidostreptus although the latter
has a higher absolute activity.

relotive activity

Rhapidostreptus Chicobolus

without activator

ethanol

! _ I chymotrypsin

zymosan

methonol

The activity after chymotrypsin activation increases with elongation of incubation time in
both species (Fig. 4); the highest activity is reached after 60 min of incubation. After activation
with ethanol the molecular weight (MW) of the PO-active band in SDS-PAGE does not differ
from unactivated hemolymph whereas after activation with a-chymotrypsin two different bands
occur with lower MW (XYLANDER & BOGUSCH, 1992), indicating that during ethanol
activation - in contrast to a-chymotrypsin - a conformational change of the proPO leads to its
activity rather than a protein cleavage.

414

WILLI E. R. XYLANDER

maximum activity

10 30

incubation time I min

Chicobolus

Rhapidostreptus

Fig. 4. — PO activity after preincubation of
hemolymph samples [min-i ml
hemolymph-i], from
Rhapidostreptus vir gator and
Chicobolus sp. with bovine o-
chymotrypsin for 1, 10, 30 and 60.
The activity of the PO increases in
both species with duration of
chymotrypsin incubation.

Chicobolus

Rhopidostreptus

FlG. 5. — Activity of ethanol and of o-
chymotrypsin activated and
unactivated hemolymph samples
[min-i ml hemolymph-i] from
Rhapidostreptus and Chicobolus
with or without potential PO-
inhibitors.

ethanol

ethanol + ED'A

chymotrypsin

ethanol ♦ EGTA

□

chymotrypsin + PTU
without activator

Fig. 6. — Activity of PO [min- 1 ml
hemolymph- 1] from the hemolymph
of Chicobolus at different pH.

Source : MNHN , Paris

PHENOLOXIDASE FROM TI IE HEMOLYMPH OF DIPLOPODA

415

7. — Polyacrylamid-gel (7.5%
acrylamid) of hemolymph (tH)
and hemocyte lysate (Hz) of
Li t ho bi us forficatus ( .'•).
Rhapidostreptus (:<h) and
Chicobolus (Ch), with
subsequent reactivation of
enzymes by incubation in a 2%
Triton-X 100 aquaeous solution
for 30 min. washing in buffer
and incubation in an 0.02 M
dopa solution in 0.01 M
cacodylate buffer (pH 7.0). At
the position of reactivated PO
dopachrom and melanin is
formed resulting in pinkish and
later brownish or black band.
Molecular standard (SDS-6H.
Sigma. Munich) run in the same
gel indicates the MW of the PO
(corresponding at about 230 kD
in all three species).

205 — -

1 16 ►

97 — ^

68-

46-

-Ch-

+«*

— PAGE of hemocyte lysate, total hemolymph and plasma of Rhapidostreptus (run under the same conditions and
with subsequent procedure described for Fig. 7). The left part of the gel was stained with Coomassie Brillant
Blue, the right treated as described for Fig. 7). No difference in MW is visible.

Source

416

WILLI E. R. XYLANDER

octivity

5 I

Fig. 9. — PO-activity of Chicobolus in the
hemocyte lysate (obtained after
moderate centrifugation at about 60-
100 g at 4°C for 10 min) and
hemolymph after ethanol
activation.

hemolymph

hemocyte lysate

Fig. 10. — Staining capabilities of
hemocyte monolayers (in % of each
hemocyte type) on glass slides of
Chicobolus after ethanol activation
and dopa incubation.

i n

o staining

weak reaction

strong reaction

Microbial activators (zymosan, murein, bacterial lipopolysaccharides) which are potent
inductors of PO-reaction in other arthropods have little effect neither on total hemolymph nor on
hemocyte lysate (Figs 2 & 3 and unpubl. results); therefore, another activation mechanism may
occur in Diplopoda than in Decapoda and Insecta.

Dopa is the substrate used best by the PO of EtOH-activated hemolymph from
Rhapidostreptus and Chicobolus as dopamin and pyrrogallol are less effectively used and
tyrosin, pyrocatechol and norephedrin hardly at all (see also XYLANDER & BOGUSCH, 1992).

The PO can be inhibited in the two diplopods and Lithobius by phenylthiourea and at least
in the diplopods by EDTA and EGTA (Fig. 5) indicating that the PO is of the tyrosinase-type
and Ca2+ dependend. The pH-optimum of the PO in both diplopods is pH 7.0, although the
activity is rather high between pH 6.0 and 8.0 (Fig. 6; see also XYLANDER & BOGUSCH, 1992
for results on Rhapidostreptus).

SDS-PAGE under non-reducing conditions

After SDS-PAGE under non-reducing conditions, reactivation of enzymes by incubation in
Triton-X 100 and incubation in dopa brownish to black bands occur at the position in the gel of
the PO and proPO, respectively. The proPO has corresponding molecular weights (MW) of

Source : MNHN, Paris

PHF.NOLOXIDASF. FROM THE HEMOLYMPH OF DIPLOPODA

417

about 230 kD in Lithobius, Rhapidostreptus and Chicobolus (Fig. 7). After incubation with
a-chymotrypsin the MW is reduced to about 200 and 180 kD whereas EtOFI has no such effect
(XYLANDER & BOGUSCH, 1992). PO-active bands have the same MW in hemocyte lysate, total
hemolymph and plasma of Rhapidostreptus (Fig. 8).

Localization of the proPO

After moderate centrifugation of hemolymph of Rhapidostreptus virgator at 4°C at about
60-100 g, resuspension and sonification of the hemocyte pellet the activity was mainly (about
80%) found in the hemocyte lysate whereas only little (about 20%) occurred in the plasma
(XYLANDER & BOGUSCH, in prep.); hemolymph samples from the same pool not centrifuged
had about the sum of plasma and hemocyte lysate activity. Investigations with Chicobolus with
the same procedure led to degranulation of hemocytes and artificially high activity in the plasma
(Fig. 9); if hemolymph was poured into the same amounts of ice-cold hemocyte stabilizing
buffer (after SODERHALL et al., 1979) about half of the activity occured in the hemocytes.

Investigations of glutaraldehyd-fixed hemocyte monolayers of Rhapidostreptus and
Chicobolus after activation with ethanol and subsequent dopa-overlay showed that PO-activity
occured mainly in the granular hemocytes (with different intensities which could be designed to
two types of granular hemocytes with different spreading capabilities; FIG. 10). Plasmatocytes
remained unstained whereas the prohemocytes showed high variability in their PO-reaction
(Figs 10 & 11). PTU in control hemocyte monolayers inhibits intracellular PQ reaction in both
species.

Exocytosis of the PO system in Rhapidostreptus cannot be initiated by addition of
solutions of various microbial cell wall components as it has been found in insects and
crustaceans (JOHANSSON & SODERHALL, 1985. 1989a, b, c).

A P £ B

*1 4 « " #

FiG. II. — Granular hemocytes and plasmatocytes of Chicobolus in a hemocyte monolayer. A. Phase contrast. B. Bright
lield. Only the stained granular hemocytes are visible under these conditions. P: plasmatocytes; G; Granular
hemocyte; U: Prohemocyte.

418

WILLI E. R. XYLANDER

DISCUSSION AND CONCLUSION

As in other arthropods the PO of diplopods occurs in the heinolymph as inactive proPO.
Activators found to be efficient in the diplopods also have been reported to activate the proPO of
insects and crustaceans (cf. ASHIDA & YAMAZAKI. 1990; GOTZ, 1988; JOHANSSON &
SODERHALL, 1989b). However, microbial cell wall components (murein, 13-1,3-glucans,
lipopolysaccharides from outer cell membrane of Gram-negative bacteria) which activate the
proPO in insects and crustaceans (ASHIDA et a!., 1983; ASHIDA & SODERHALL, 1984; ASHIDA
& Yoshida. 1988; Johansson & Soderhall, 1989b; Smith & Soderhall, 1983;
SODERHALL. & HALL. 1984; SODERHALL & UNESTAM. 1979; SODERHALL et al., 1988) and
have been considered to be the inducers of in vivo PO-reactions did not have any effect on
diplopods with the system used (XYLANDER. 1992; XYLANDER & BOGUSCH, 1992, this
paper). Regarding other capabilities (PTU-inhibition, pH-optimum. Ca2+ dependence,
preference for dopa as substrate) the proPO and PO of diplopods correspond to that of insects,
crustaceans and the few chilopods investigated.

The MW of the two diplopods and Lithobius ranges at about 230 kD. This is higher than
most data found in most insects and crustaceans the PO and proPO of which have a MW of 60-
80 kD; however, corresponding MW (higher than 200 kD) have also been reported for various
crustaceans and insects (NELLAIAPPAN et al., 1989; YAMAURA et al, 1980). ASPAN &
SODERHALL (1991) and GILLESPIE et al. (1991) reported that the proPO tends to aggregate to
form polymeres indicating that the bands found could be a distinctive polymere (tetramere?) of
the proPO (cf. ASPAN & SODERHALL, 1991). Own investigations using gel filtration
chromatography of hemolymph of Rhapidostreptus (data not shown), however, also led to MW
of more than 230 kD of the proPO.

Whether the proPO of arthropods occurs mainly free in the hemolymph or in the
hemocytes depends on the taxon investigated. In most insects and all crustaceans investigated,
the proPO is predominantly located in the hemocytes; it is discharged and activated after
infections (cf. JOHANSSON & SODERHALL, 1989b; ASHIDA & YAMAZAKI, 1990). In some
insects, however, the proPO is reported to occur free in the plasma (GOTZ et al., 1987. SAUL et
al, 1987). In Rhapidostreptus the majority of the proPO is also found intracellularly
(XYLANDER & BOGUSCH, in prep.), whereas in Chicobolus (probably as a preparation artifact
due to unsufficient stabilization of hemocytes in a crayfish saline) only about 50% are found in
the hemocyte lysate.

In all diplopods and chilopods investigated the granular hemocytes are the site of proPO
localization (Bowen, 1968; Krishnan & Ravindranath, 1973; Nevermann et al. 1991;
XYLANDER & NEVERMANN, 1993), whereas only few plasmatocytes show a faint staining after
dopa incubation; the spherulocytes of chilopods are PO-negative (NEVERMANN et al, 1991;
XYLANDER & NEVERMANN, 1993). In chilopods staining is rather slow in comparison to
diplopods (XYLANDER & NEVERMANN, 1993).

Although microbial cell wall components do not initiate exocytosis of the PO cascade as in
crustaceans and insects (cf. reviews in JOHANSSON & SODERHALL, 1989b; ASHIDA &
YAMAZAKI, 1990) rather strong melanization can be found at wound margins of diplopods. This
indicates that the PO is activated after an injury or infection and is involved in subsequent
immune response: one function of this enzyme may be to support rapid wound closure and to
kill bacteria and fungi before they enter the hemocoel and may lead to dangerous infections.
Furthermore, an intracellular activation has been found in hemocytic aggregates around foreign
material in vitro and in vivo (NEVERMANN, 1989; NEVERMANN & XYLANDER, this volume).
As in other arthropods where intracellular activation has been reported (VOLKMANN, 1991;
NAYAR et al, 1992) the “melanized hemocytes” in the capsule may constitute an efficient
barriere for metabolic waste from potential parasites and invaders on one hand and for nutrients
from the hemolymph necessary for their survival on the other hand. Thus melanized hemocytes

Source : MNHN, Paris

PHENOLOXIDASE FROM THE HEMOLYMPH OF DIPLOPODA

419

may support the function of a multilayered hemocyte capsule formed around parasites as a
typical cellular defence reaction of arthropods.

ACKNOWLEDGEMENTS.

I would like to thank O. BOGUSCH, H -U. Jahn. L. Nevermann. A. Hudel, and Prof. Dr. P. Gotz for their support.
Investigations and participation in the CIM congress were supported by the President of the Justus-Liebig-University
Giessen.

REFERENCES

Ashida. M., Ishizaki Y. & Iw ah an a H., 1983. — Activation of pro-phenoloxidase by bacterial cell walls or B-1,3
glucans in plasma of the silkworm. Bombyx mori. Biochem. Biophys. Res. Comm. 113 : 562-568.

Ashida, M. & SOderhaall. K., 1984. — The prophenoloxidase activating system in crayfish. Comp Biochem.
Physiol., 77 B : 21-26.

Ashida. M. & Yamazaki H. I.. 1990. — Biochemistry of the phenoloxidase system in insects: with special reference to
its activation. In : E. Ohnishi & H. Ishizaki. Moiling and metamorphosis. Tokyo. Japan Sci. Soc. Press, Berlin.
Springer-Verlag : 239-265.

Ashida, M. & Yoshida H., 1988. — Limited proteolysis of prophenoloxidase during activation by microbial products in
insect plasma and effect of phenoloxidase on electrophoretic mobilities of plasma proteins. Insect Biochem.. 18 :
1 1-19.

Aspan, A. & SOderhall. K., 1991. — Purification of prophenoloxidase from crayfish blood cells, and its activation by
an endogenous serine proteinase. Insect Biochem., 21 : 363-373.

Bowen, R. C., 1968. — Tyrosinase in millipede hemocytes. Trans. Amer. Microsc. Soc., HI : 390-392.

Chadwick. J. M. & Aston. W. P.. 1978. — An overview of insect immunity. In : Gershwin. M. E. & E. L. Cooper.
Animal models of comparative and developmental aspects of immunity and diseases. Oxford. Pergamon Press : 1-14.

Gillespie, J. P.. Bidochka, M. J. & Khachatourians. G. G., 1991. — Separation and characterization of grasshopper
hemolymph phenoloxidases by sodium dodecyl sulfate-polyacrylamide gel electrophoresis. Comp. Biochem.
Physiol., 98C : 351-358.

GOTZ. P.. 1988. — Immunreaktionen bei Wirbellosen, insbesondere Insekten. Verb. Dtsch. Zool. Ges.. 81 : 113-129.

Gotz, P., Enderlein, G. & ROETTGEN, I., 1987. — Immune reactions of Chironomus larvae (Insecta: Diptera) against
bacteria. J. Insect Physiol., 33 : 993-1004.

Gotz, P. & Vey, A.. 1974. — Humoral encapsulation in Diptera (Insecta) - defense reactions of Chironomus larvae
against fungi. Parasitology, 68 : 193-205.

Johansson, M. W. & SOderhall, K., 1985. — Exocytosis of the prophenoloxidase activating system from crayfish
haemocytes. J. Comp. Physiol., B156 ; 175-181.

Johansson, M. W. & SOderhall, K., 1989a. — A peptide containing the cell adhesion sequence RGD can mediate
degranulation and cell adhesion of crayfish granular haemocytes in vitro. Insect Biochem., 19 : 573-579.

Johansson, M. W. & SOderhall. K., 1989b. — Cellular immunity in crustaceans and the proPO system. Parasitology
Today, 5 : 171-176.

Johansson, M. W. & SOderhall, K.. 1989c. — A cell adhesion factor from crayfish haemocytes has degranulating
activity towards crayfish granular cells. Insect Biochem.. 19 : 183-190.

Krishnan, G. & Ravindranath. M. H.. 1973. — Phenoloxidase in the blood cells of millipedes. J. Insect Physiol.,
19 : 647-653.

Nayar, J. K., Mikarts, L. L., Knight, J. W. & Bradley. T. J.. 1992. — Characterization of the intracellular
melanization response in Anopheles quadrimaculatus against subperiodic Brugia malax i larvae. J. Parasitol.. 78 :
876-880.

Nellaiappan. K., Vinayagam, A. & Kalyani. R.. 1989. — Electrophoretic pattern of blood and cuticular phenoloxidase
of different crustaceans. Exp. Biol.. 48 . 177-179.

Nevermann, L., 1989. — Licht- und elektronenmikroskopische Untersuchungen der Haemocyten von Lithobius
forficatus und ihre Bedeutung fur zellulare Abwehrreaktionen. Diploma-thesis. University of Giessen, 69 pp.

Nevermann, L., Xylander, W. E. R. & Seifert, G.. 1991. — The hemocytes of the centipede Lithobius forficatus
(Chilopoda, Lithobiomorpha) - Light and electron microscopic studies using in-vitro techniques. Zoomorphology,
1 10 : 317-327.

Pye. A. E., 1974. — Microbial activation of prophenoloxidase from immune insect larvae. Nature, 251 : 610-613.

Ratcliffe, N. A., Leonard. C. & Rowley. A. F., 1984. — Prophenoloxidase activation: Nonself recognition and cell
cooperation in insect immunity. Science, 226 : 557-559.

420

WILLI E. R. XYLANDER

Saul, S. J., Bin, L. & Sugumaran. M., 1987. — The majority of prophenoloxidase in the hemolymph of Manduca sexta
is present in the plasma and not in the hemocytes. Dev . Comp. Immunol., 11 : 479-485.

Smith. V. & SOderhAll. K.. 1983. — B-1,3 Glucan activation of crustaceans hemocytes in vitro and in vivo. Biol.
Bull., 164 : 299-314.

SOderhAll, K.. 1982. — Prophenoloxidase activating system and melanization - a recognition mechanism of
arthropods? A review. Dev. Comp. Immunol.. 6 : 601-611.

SOderhAll. K. & HALL, L.. 1984. — Lipopolysaccharide-induced activation of prophenoloxidase activating system in
crayfish haemocyte lysate. Biochim. Biophys. Acta. 797 : 99-104.

SOderhAll. K.. HAll. L.. Unestam. T. & Nyhlen, L., 1979. — Attachment of phcnoloxidase to fungal cell walls in
arthropod immunity. J. Inv. Pathol., 34 : 285-294.

SOderhAll. K.. Rogener. W.. SOderhAll, I., Newton, R. P. & Ratcliffe, N. A., 1988. — The properties and
purification of a Blaberus craniifer plasma protein which enhances the activation of haemocyte prophenoloxidase by
a 8-1.3-glucan. Insect Biochem., 18 : 323-330.

SOderhAll. K. & Unestam. T., 1979. — Activation of serum prophenoloxidase in arthropod immunity. The specificity
of cell wall glucan activation and activation by purified fungal glycoproteins of crayfish phenoloxidase. Can. J.
Microbiol.. 25 : 406-414.

V ey, A. & GOtz. P.. 1975. — Humoral encapsulation in Diptera (Insecta) - comparative studies in vitro. Parasitology,
70 : 77-86.

VOLKMANN, A., 1991. — Localization of phenoloxidase in the midgut of Periplaneta americana parasitized by larvae of
Moniliformis moniliformis (Acanthocephala). Parasitol. Res., 77 : 616-621.

Xylander, W. E. R., 1992. — Immune defense reactions of Myriapoda - A brief presentation of recent results. [ /// : E.
MEYER. K. Thaler & W. SCHEDL , Advances in Myriapodology.] Ber. nat.-med. Verein Innsbruck, Suppl. 10 : 101-
1 10.

XYLANDER, W. E. R. & BOGUSCH, O., 1992. — Investigations on the phenoloxidase of Rhapidostreptus virgator
(Arthropoda. Diplopoda). Zool. Jb. Physiol., 96 : 309-321.

Xylander, W. E. R. & BOGUSCH. O., (in prep.) — The Prophenoloxidase of the millipede Rhapidostreptus virgator
(Diplopoda, Spirostreptidae) is mainly located in the granular hemocytes.

Xylander. W. E. R. & Nevermann, L.. 1990. — Antibacterial activity in the hemolymph of Myriapoda (Arthropoda). J.
Inv. Pathol., 56 . 206-214.

Xylander, W. E. R. & Nevermann, L.. 1993. — Phenoloxidase-active hemocytes in Lithobius forficatus, Scolopendra
cingulata (Chilopoda) and Chicobolus sp. (Diplopoda). Verb. Dtsch. Zool. Ges., 86 : 197.

Yamaura, I., YONEKURA, M.. Katsura, Y., Ishiguro. M. & Funatusu, M.. 1980. — Purification and some physico¬
chemical properties of phenoloxidase from the larvae of housefly. Agric. Biol. Client., 44 : 55.

Source : MNHN, Paris

In vitro Cellular Immune Reactions of Hemocytes
against Bacteria and their Differential Degradation in

Myriapods

Lutz NEVERMANN & Willi E. R. XY LANDER

Inslitut fur Allgemeine und Spezielle Zoologie, Justus-Liebig-Universitat Giessen
Stephanstr. 24, D-35390 Giessen. Germany

ABSTRACT

In vitro the hemocytes of various diplopods and chilopods are capable of phagocytosis and degradation of bacteria.
The sequence of this process depends on the myriapod species and the type of bacteria. In Rhapidostreptus virgator many
Micrococcus luteus have been phagocytosed but occur uneffected after 4 hours whereas E. coli shows indications of lysis.
Alter iO hours M. luteus also has been degradated. In Lithobius forficatus , nodules, aggregations of hemocytes
phagocytosing bacteria, are formed when hemocytes are added to a bacterial culture which is turned around continuously.
During this process, the hemocytes dcgranulate. M. luteus is lysed within 1 hour when phagocytosed or even located
close to a hemocyte cell. Enterobacter cloacae is enclosed extracellulary within the matrix of the nodule and subsequently
phagocytosed. E. coli was not aggregated within the nodule but hemocytes discharged vesicular content onto their
surface. They become phagocytosed but do not show indications of lysis after 1 hour. In Scolopendra cingulata
hemocytes also form nodules with bacteria in the continuously moved culture. These nodules contain a large extracellular
matrix in which the bacteria arc embedded. Only very few bacteria are phagocytosed within Ih of incubation and there are
no signs of lysis in M. luteus.

RESUME

Reactions immunitaires in vitro d'hemocytes contre des bacteries et leur degradation chez les
myriapodes (Diplopoda & Chilopoda).

In vitro, les hemocytes de differents diplopodes et chilopodes sont capables de phagocyter et de degrader des bacteries.
La sequence de ces processus depend des especes de myriapodes et de bacteries. Au bout de 4 heures. chez Rhapidostreptus
virgator, de nombreux Micrococcus luteus sont phagocytes mais ne subissent aucun dommage. landis que E. coli montre
deja des indices de lyse. Au bout de 20 heures, M. luteus cst, lui aussi, degrade. Chez Lithobius forficatus , de petits
nodules se forment. agregations d'hemocytes phagocytant des bacteries, si des hemocytes sont ajoutes ^ une culture
bacterienne continuellcment remuee. Durant ce processus, les hemocytes degranulent. M. luteus est lys6 en 1 heure. si les
bacteries sont phagocytes ou meme seulement se trouvent & proximite d'un hemocyte. Enterobacter cloacae se trouve
inclus de maniere extracellulaire dans la matrice du nodule, puis phagocyte. E. coli ne montre pas d'agregation en
nodules, mais les hemocytes d^chargcnt le contenu de certaines v6sicules sur la surface bacterienne. Elies seront
finalement phagocytes mais aucune trace de lyse n’est visible au bout de 1 heure. Chez Scolopendra cingulata , les
hemocytes forment des nodules avec les bacteries en culture. Ces nodules contiennent une volumincuse matrice
extracellulaire dans laquelle les bacteries sont incorporees. Au bout de 1 heure. seul un trfes petit nombre d'entre elles est
phagocyte ou lys6.

Nevermann, L. & XYLANDER, W. E. R.. 1996. — In vitro cellular immune reactions of hemocytes against bacteria
and their differential degradation in myriapods. In: Geoffroy. J.-J.. Mauries. J.-P. & Nguyen Duy - Jacqukmin, M.,
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist. not.. 169 : 421-430. Paris ISBN : 2-85653-502-X.

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LUTZ NEVERMANN & WILLI E. R. XYLANDER

INTRODUCTION

Two mechanisms are involved in defense of arthropods against infections by
microorganisms: the formation of antibacterial substances which in insects mainly act free in the
plasma, and the action of defense cells e. g. hemocytes which form nodules or phagocytose the
microbes. Lysozyme, an enzyme depolymerizing the cell wall of Gram-positive bacteria, is
permanently found in the hemolymph (DUNN, 1986; GOTZ, 1988) whereas most antibacterial
substances against Gram-positive bacteria are formed within a few hours after an external
stimulus, e. g. infections or injury (cf. BOMAN, 1986; GOTZ, 1988). Therefore, nodule
formation and phagocytosis are the initial defense mechanisms against microbes that help to
absorb infective bacteria until other mechanisms are available.

The mechanisms and the procedure of cellular defense against bacteria has been well
documented in insects, whereas little is known about such processes in “myriapods”. This
paper, therefore, describes nodule formation, phagocytosis, specificity of phagocytic hemocytes
to different bacteria and degradation of ingested bacteria in the diplopod Rhapidostreptus virgator
and the chilopods Lithobius forficatus and Scolopendra cingulata.

MATERIAL AND METHODS

Animals

Specimens of Scolopendra cingulata were collected in 1990 at different sites in northern Spain and maintained
solitary since then in large bcllaplast boxes with 2 cm of clean sand or soil covered with paper towels at room
temperature at a shaded place. Humidity of the substratum was controlled weekly and moistened with tab water if
necessary. Specimens were fed with Acheta domesticus, Tenebrio moliiov mainly larvae and Sarcophaga sp. mainly
imagines. Lithobius and Rhapidostreptus were obtained and reared as described by Xylander & Nevermann (1990).
Hemolymph was obtained as specified earlier (Xylander & Nevermann. 1990; Xylander & Bogusch. 1992) and
dropped directly into the culture medium.

Hemocyte preparations

For investigations with hemocytes of Rhapidostreptus hemolymph was added to a 1 cm2 piece of gelatine foil
alter swelling it in I-Ringer (1 1 g NaCI, 7 g KC1. 5.5 g CaCte, in 11 H2O) for 30 min in culture dishes. Hemocytes were
allowed to adhere to the substratum in I-Ringer for 50 min at room temperature. Then 10 ul of a bacterial suspension
were added. Hemocytes cultures with bacteria were slowly moved on a shaker for 1 h. 4 h and 20 h.

For investigations on chilopods 10 pi of suspended "washed" bacteria (see below) were added to 1 ml LAH (, Lithobius -
artificial-hemolymph according to Wenning,1989) in an Eppendorf cap and then 10 pi freshly collected hemolymph
was supplied. The Eppendorf caps were Fixed with cello tape to a spinbar and rotated vertically on a magnetic stirrer for
1 h at room temperature. Subsequently, the Eppendorf cap was centrifuged at lOOxg for about a minute, the
supernatant was removed and the pellet was processed for electron microscopy.

Preparation of bacterial cultures

The bacteria used in this study were obtained from Prof. Dr. P. GOTZ, Berlin (Micrococcus luteus, Enterobacter
cloacae 6-12 and Escherichia coli K12 D31). Bacteria were cultivated in nutrient broth (Merck Standard I) over night at
35 C in a water bath. Small amounts of this “over-night-culture" were added to fresh broth and raised until optical density
reached 0.65 at 565 nm.

For investigations on chilopods I ml of bacteria in culture were washed twice by centrifugation (room
temperature, 5 min at 1000 xg) and resuspension of the bacterial pellet to 1 ml with LAH. Finally, 10 pi of the
resuspension were added to the hemocyte culture.

Preparation for transmission electron microscopy

All samples were fixed in 2.5% glutaraldehyde. 2% paraformaldehyde in 0.1 M sodium cacodylate buffer,
pH 7.0 tor 2 h at 4-6°C, washed in buffer, postfixed in 2% OsOj, dehydrated through an acetone series and embedded in
araldite. Semithin and ultrathin sections were made on a Reichert OmU3 ultracut microtome, mounted on formvar coated
copper grids, stained with uranyl acetate and lead citrate and investigated with a Zeiss EM 9 A transmission electron
microscope (TEM).

RESULTS

Rhapidostreptus

Within 4 h after addition of bacteria the hemocytes of Rhapidostreptus have phagocytosed
both Gram-negative E. coli and Gram-positive Micrococcus luteus (Figs 1, 2, 5 & 6). The
bacteria are located in vacuoles mainly in that part of the hemocytes without contact to the

Source : MNHN, Paris

IN VITRO CELLULAR IMMUNE REACTIONS OF HEMOCYTES IN MYRIAPODS

423

substratum (Figs I & 5); the vacuoles have been found to be smaller around Micrococcus than
around E. coli (Figs 1 & 5). Most Micrococcus remain rather uneffected and still show their
typical electron dense internal structure (Figs 1 & 2) whereas the inside of E. coli occurs less
electron dense and flocculent indicating degradation (Figs 5 & 6). After 20 h dramatic changes
are lound. The cell wall of phagocytosed Micrococcus has been completely degradated and the
bacteria have been killed as noticed from an obvious decrease in their electron density (Figs 3 &
4). Plasmatocytes of Rhapidostreptus have disintegrated after 20 h leaving a translucent
vesicular cell debris whereas granulocytes look relatively unchanged. Bacteria which have not
been phagocytosed still exhibit their normal structure but show electron dense amorphous
material at their surface.

Lithobius forficatus

After 1 h in the “stirrer culture” plasmatocytes, granulocytes and some spherulocytes (for
description and classification of hemocytes, see NEVERMANN et al., 1991) have aggregated to
form nodules (Fig. 7). Plasmatocytes and granulocytes, however, no longer can be
differentiated, since they have lost most of their grana obviously by exocytosis into the culture
medium. All three species of bacteria are phagocytosed. Phagocytic vesicles enclosing bacteria
have been observed to fuse with lysosomes and become electron dense (Figs 8, 9, 1 1 & 16).

Micrococcus is lysed in electron translucent phagocytic vesicles (Fig. 14). Their cell wall
is thinned off and their plasm becomes more electron" translucent in TEM compared to living
bacteria (Figs 14 & 15). Even bacteria attached to the surface of hemocytes or lying in short
distance to hemocytes frequently undergo lysis. Bacteria, located in some distance are uneffected
(Fig. 15).

Enterobacter , in contrast to Micrococcus and E. coli, becomes aggregated in the
hemolymph culture and, therefore, clusters of bacteria are surrounded by hemocytes (Fig. 7).
Nevertheless, some of the bacteria are phagocytosed and lysed (Figs 7-9). In one single
phagocytic vesicle different stages of bacterial lysis may be found. Destroyed bacteria swell and
occur less electron dense (Figs 7 & 8).

E. coli are not aggregated but show a scattered distribution throughout the hemocyte
nodule. They become phagocytosed occasionally but no indications of lysis was found after 1 h
of culture neither in translucent nor in electron dense phagocytic vesicles (Figs 10-13). Prior to
phagocytosis fibrous material from the so-called structured vesicles (see NEVERMANN et al.,
1991) is discharged onto the surface of E. coli (Fig. 10). The same material may also be found
in phagocytic vesicles containing bacteria (Fig. 12).

Scolopendra cingulata

The hemocytes of S. cingulata aggregate to form nodules which contain an voluminous,
possibly fibrous, extracellular material (Fig. 17). Many bacteria (only Micrococcus was tested)
are embedded in this matrix and most of them seem to have no contact to the hemocytes.
Nevertheless, some bacteria are phagocytosed. As in Lithobius the phagocytic vesicles of
Scolopendra may have an electron dense matrix but - in contrast to Lithobius - Micrococcus does
not get lysed (Fig. 16). Presumably, in this species it takes more time than 1 h until visible signs
of lysis are found in TEM.

DISCUSSION

Phagocytosis has been described for insects in a number of papers but for other groups of
arthropods there is only very little information on phagocytosis and intracellular degradation of
microbes by hemocytes (e. g. JOHNSON, 1981; PALM, 1953; TYSON & JENKIN. 1973). The
TEM-micrographs of phagocytosis in arthropods, however, are very similar indicating that the

424

LUTZ NEVERMANN & WILLI E. R. XYLANDER

mechanisms must be quite alike. Even intracellular degradation of bacteria and the appearance ol
electron-dense phagocytic vesicles are much alike in insects (cf. ROWLEY & RATCLIFFE, 1976a
and our investigation on chilopods). .

Two initial mechanisms are involved in host cellular defense reaction against invading
bacteria: phagocytosis and nodule formation. Phagocytosis is supposed to be the main reaction
against low numbers of bacteria in arthropods (GOTZ, 1982; CHRISTENSEN & NAPPI, 1988).
But when there are high numbers of microbes entering the hemocoel nodule formation occurs
(RATCLIFFE & GaGEN, 1977). Our in vitro observations of Lithobius and Scolopendra indicate
that nodule formation and phagocytosis are related processes in the coagulum.

In the spiny lobster, Panulirus japonicus a factor is released from granular hemocytes after
stimulation with bacteria which provokes clotting of the other hemocyte types (AONO et al.,
1993). This implies that even a small number of microbes can elicit clotting. Nodule formation
may, therefore, not be restricted to reactions against higher numbers of bacteria but is a general
process occurring parallel to phagocytosis.

Lithobius hemocytes can degradate bacteria faster (in only 1 h) than Scolopendra and
Rhapidostreptus and other arthropods investigated. In an in vitro investigation with hemocytes
of Calliphora erythrocephala ROWLEY & RATCLIFFE (1976a) found cell wall damage and
swollen bacteria not before 2 h of incubation. Heat killed bacteria Bacillus cereus injected into
the hemocoel were entrapped into nodules but they did not show any signs of breakdown even
after 24 h (RATCLIFFE & GaGEN, 1977).

The reaction of hemocyte types to different bacteria varies. Granular cells of Galleria
meUoneUa make contact with E. coli in vitro, get stressed and degranulate (ROWLEY &
RATCLIFFE, 1976b). Eventually, they may phagocytose E. coli but most bacteria remain
attached to the outside of the cells. The plasmatocytes of this species remain unstressed and
phagocytose bacteria and the decaying granular cells. However, no killed bacteria have been
observed. In this investigation all hemocyte types were involved in phagocytosis and nodule
formation. In Rhapidostreptus granular hemocytes even seemed to be more stable and less
sensible to lysis than plasmatocytes.

F,GS 1-6. — Rhapidostreptus virgator : 1. Plasmatocyte (P) containing some not lysed Micrococcus luteus in phagocytic-
vesicles. 4 h of incubation. Scale bar: 2 pm. 2. Plasmatocyte with phagocytosed M. luteus . 4 h of incubation.
The diplococcal structure of the bacteria is visible (arrowhead). Scale bar: 1 pm. 3. Granulocyte (II) with
phagocytosed M. luteus which has become translucent due to lysis. 20 h of incubation. Scale bar: 0.2 pm.
4. Granulocyte (II) with phagocytosed lysed M. luteus , 20 h of incubation. Scale bar: 1 pm. 5. Granulocyte (I)
with phagocytosed and lysed E. coli . 4 h of incubation. The phagocytic vesicle surrounding the bacterium is
significantly larger than that around M. luteus. Scale bar: 1 pm. 6. Granulocyte (I) with lysed /:. coli in the
phagocytic vesicle (pv) showing a flocculent less electron dense content. 4 h of incubation. Scale bar: 1 pm.

Figs 7-9. — Lithobius forficatus: 7. Hemocytic nodule. Plasmatocytes (P) forming a primary capsule around agglutinated
Enterobacter cloacae (E). Arrowheads indicate bacteria just being phagocytosed. Scale bar: 5 pm. 8. Large
electron dense phagocytic vesicle (pv) containing E. cloacae in various stages of degradation. Scale bar: 1 pm.
9. Fusion of an electron dense lysosome with a phagocytic vesicle containing E. cloacae. Scale bar: 0.5 pm.

FIGS 10-13. — Lithobius forficatus: 10. E. coli during phagocytosis. Note fibrous material (*) discharged onto bacteria.
11. Electron dense phagocytic vesicle (pv) with five E. coli without clear indications of lysis. 12. Phagocytic
vesicle of a granular hemocyte with structured content (sv) and phagocytosed E. coli. The cell shows signs ol
disintegration. 13. Small translucent vesicles (arrow) gathering around phagocytosed E. coli. All scale bars on
this plate: 1 pm.

FiGS 14-15. — Lithobius forficatus & FIGS 16-17. — Scolopendra cingulata: 14. Phagocytosed and lysed Micrococcus
luteus (M). Note the thinned murein sacculus. The hemocytes have lost most ol their grana. Scale bar: 5 pm.
15. Few still electron dense M. luteus with an intact murein sacculus (arrow) at some distance to a hemocyte.
Bacteria located closer are more electron translucent on lysis. Arrowhead: semi-lysed diplococcus. Scale bar:
1 pm. 16. Four M. luteus in an electron dense phagocytic vesicle (pv) without signs of degradation. Scale bar:
1 pm. 17. Nodules of S. cingulata with an voluminous extracellular matrix (X) embedding the hemocytes and
M. luteus (M). Some bacteria are being phagocytosed (Mp). Scale bar: 1 pm.

Source : MNHN. Paris

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IN VITRO CELLULAR IMMUNE REACTIONS OF HEMOCYTES IN MYRIAPODS

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IN VITRO CELLULAR IMMUNE REACTIONS OF HEMOCYTES IN MYRIAPODS

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The time span until bacterial degradation becomes visible differs in the species investigated
and may be due to antibacterial substances which have earlier been demonstrated by bacterial-
agar dittusion tests in various diplopods and chilopods (VAN DER WALT et al. 1990-
XYLANDER & Nevermann, 1990). Antibacterial substances produced by hemocytes may be
involved in the intracellular and extracellular degradation of bacteria as observed. The thinnin° of
the cell wall of Micrococcus luteus indicates the action of lysozym which also has been shown to
hemolymph of various diplopods and chilopods (XYLANDER & NEVERMANN
1990). In the investigations by XYLANDER & NEVERMANN (1990), Scolopendra oraniensis and
Rhapidostreptus had less effect on living bacteria than e. g. Lithobius ; especially the lysozyme
effect on lyophilized Micrococcus luteus was low. Thus, the delay in cell wall lysis in
bcolopendra and Rhapidostreptus may be caused by lower titers of lysozym available in the
hemocytes. I his corresponds to recent immunocytochemical detection of varying amounts of
lysozyme in hemocytes of chilopods (Nevermann, unpublished).

In many insects investigated the fat body is the main site of synthesis of antibacterial
substances after infections; they are discharged into the hemolymph where they destroy bacteria
Hemocytes may also produce such substances (Trenczek. 1988) but their importance in insects
tor humoral antibacterial defense seems to be low. However, in other arthropods like
crustaceans (FENOUIL & ROCH, 1991; SMITH & CHRISHOLM, 1992), xiphosurans (MURAKAMI
et al, I 991; NAKAMURA et al. , 1988; TOH et al., 1991), diplopods (XYLANDER, unpublished)
and chdopods (Nevermann, unpublished), the antibacterial substances are. mainly located in
the hemocytes which most probably also are the site of their formation. Located in the hemocytes
the antibacterial substances may be strongly involved in killing and subsequent or simultaneous
lysis of bacteria. This could probably be the original function (and site of formation) of the
antibacterial substances in arthropods; in insects or a subgroup of this taxon the fat body seems
to have taken over this function from hemocytes. Thereby, the antibacterial defense may have
become subdivided in an initial cellular and a subsequent humoral phase.

However, further proteins may be involved in the cellular defense against bacteria.
Agglutination oi bacteria as found in Lithobius could be due to lectins which have been found in
the hemolymph plasma of different diplopods (JAHN & SEIFERT, 1992; JAHN & XYLANDER,
1991; XYLANDER, 1990, 1992) and may - as shown in e. g. insects (PENDLAND et al., 1988) -
be responsible for bacterial agglutination and opsonization facilitating phagocytosis.

ACKNOWLEDGEMENTS

We would like to thank Miss A. HUDEL for printing the micrographs and the President of the Justus-Liebig-
University, Giessen, who supported this investigation and our participation in the congress.

REFERENCES

Aono H., Oohara, I. & Mori, K.. 1993. — Cell type-specific roles in the hemocyte clotting system of the spiny
lobster, Panulirus japonicus. Comp. Biochem. Physiol., 105A : 11-15.

Boman, H. G., 1986. — Antibacterial immune proteins in insects. In : A. M. LaCKIE , Immune mechanisms in
invertebrate vectors. Symp. Zool. Soc., London . 56 : 45-58.

CHRISTENSEN, B. M. & Nappi, A. J., 1988. — Immune responses of arthropods. IS/ Atlas of Science. Animal and Plant
Sciences : 15-19.

Dunn, P. E., 1986. — Biochemical aspects of insect immunology. Ann. Rev. EntomoL.il : 321-339.

Fenouil, E. & ROCH. P. 1991. — Evidence and characterization of lysozyme in six species of freshwater crayfishes from
Astacidae and Cambaridae families. Comp. Biochem. Physiol. , B 99 : 43-49.

Gotz, P.. 1982. — Wie Insekten sich gegen Krankheitserreger und Parasiten vertcidigen. Sitzunesber. Ges. Naturforsch.
Freunde Berlin N. F.,22 . 33-48.

GOTZ, P., 1988. — Immunreaktionen bci Wirbellosen, insbesondere Insekten. Verb. Dtsch. Zool. Ges., 81 : 113-129.

Jahn, H. U. & SEIFERT, G.,1992. — Humorale Lektine in der Haemolymphe von Rhapidostreptus virgator. Verb. Dtsch
Zool. Ges.. 85 : 263.

430

LUTZ NEVERMANN & WILLI E. R. XYLANDER

Jahn, H. U. & XYLANDER. W. E. R., 1991. — Lektine bei Rhapidoslreptus - Beslandieile des Immunsystems? In : W.
Schoner & M. Kroger. 3 . Werkstattberichte aus der Experimentellen Biologic und Experimentellen Medizin.
Giessen. Uni versitat Giessen : 53.

JOHNSON. P. T., 1981. — Histopalhology of Aerococcus viridans var. homari infection Gaffkemia in the lobster.
Homarus americanus , and a comparison with histological reactions to a gram-negative species. Pseudomonas
perolens.J. Inv. Pathol., 38 : 127-148.

Murakami. T., Niwa. M.. Tokunaga, F.. Miyata, T & Iwanaga, S., 1991. — Direct virus inactivation of Tachyplesin I
and its isopeptides from horseshoe crab hemocytes. Chemotherapy, 37 : 327-334.

Nakamura. T.. Furunaka. H.. Miyata, T.. Tokunaga, F.. Muta. T., Iwanaga. S., Niwa. M.. Takao. T & Shimonishi.
Y..1988. — Tachyplesin. a class of antimicrobial peptides from the hemocytes of the horseshoe crab Tachypleus
tridentatus. J, Biol. Chem., 263 : 16709-16713.

Nevermann, L., XYLANDER. W. E. R. & Seifert. G.. 1991. — The hemocytes of the centipede Lithobius forficatus
Chilopoda. Lithobiomorpha: Light and electron microscopic studies using in-vitro techniques. Zoomorphology .
110 : 317-327.

Palm, N. B.. 1953. — The elimination of injected vital dyes from the blood in Myriapods. Arkiv Zool., 6 : 219-246.
Pendland. J. C., Heath, M. A. & Boucias. D. G.. 1988. — Function of a galactose-binding lectin from Spodoptera
exigua larval haemolymph: opsonization of blastospores from entomogenous hyphomycetes. J. Insect Physiol .,
34 : 533-540.

Ratcliffe, N. A. & Gagen, S. J.. 1977. — Studies on the in vivo cellular reactions of insects: An ultrastructural analysis
of nodule formation in Galleria mellonella. Tissue Cell . 9 : 73-85,.

Rowley. A. F. & Ratcliffe, N. A.. 1976a. — An ultrastructural study of the in vitro phagocytosis of Escherichia coli by
the hemocytes of Calliphora erythrocephala. J. Ultrastr. Res,. 55 : 193-202.

Rowley, A. F. & RATCLIFFE, N. A.. 1976b. — The granular cells of Galleria mellonella during clotting and phagocytic
reactions in vitro. Tissue Cell , 8 : 437-446.

SMITH, V. J. & Chrisholm, J. R. S., 1992. — Non-cellular immunity in crustaceans. Fish Shellfish Immunol., 2 : 1-31.
Toh, Y., Mizutani. A. .Tokunaga, F.. Muta, T. & Iwanaga, S.. 1991. — Morphology of the granular hemocytes of the
Japanese horseshoe crab Tachypleus tridentatus and immunocytochemical localization of clotting factors and
antimicrobial substances. Cell Tissue Res.. 266 : 137-147.

Trenczek, T.. 1988. — Injury and immunity in insects. Studies with Hyalophora cecropia fat body and hemocytes in
vivo and in vitro. In : F. SEHNAL, A. Zabza & D. L. DENLINGER. Endocrinological frontiers in physiological insect
ecology. Wroclaw, Wroclaw Technical University Press : 369-378.

Tyson, C. J. & Jenkin, C. R.,1973. — The importance of opsonic factors in the removal of bacteria from the circulation
of the crayfish Parachaeraps bicarcinus. Austral. J. Exp. Biol. Med. 5c*/., 51 : 609-615.

Van Der Walt, E.. McClain, E.. Puren. A. & Savage, N.. 1990. — Phylogeny of arthropod immunity. An inducible
humoral response in the Kalahari millipede. Triaenophorus triodus Attems. Naturwiss. ,77 : 189-190.

Wknning, A., 1989. — Transporteigenschaften der Malpighischcn GefaBe von Lithobius forficatus (L. ), Myriapoda.
Chilopoda. Verb. Dt. Zool. Ges.^ 82 : 215-216.

XYLANDER, W. E. R., 1990. — Immune defense reactions of myriapods - recent results and perspectives. In : “ 8th
International Congress of Myriapodology. Abstracts” . Veroffentl. d. Universitdt Innsbruck: 111 : 65.

Xylander. W. E. R., 1992. — Immune defense reactions of Myriapoda - A brief presentation of recent results. [/// :
Thaler, K., E. MEYER & W. SCHEDL, Advances in Myriapodology .1 Ber. d. naturw.-med. Verein Innsbruck, Suppl.
10 : 101-110.

XYLANDER, W. E. R. & BoGUSCH. O., 1992. — Investigations on the phcnoloxidase of Rhapidoslreptus virgator
(Arthropoda, Diplopoda). Zool. Jb. Physiol., 96 . 309-321.

Xylander, W. E. R. & Nevermann, L., 1990. — Antibacterial activity in the hemolymph of Myriapoda Arthropoda. J.
Inv. Pathol.. 56: 206-214.

Source : MNHN, Paris

Evidence for Antibacterial Activity
in Haemolymph of Diplopoda:
Preliminary Results

Grzegorz KANIA *, Jan JAROSZ **, Mariola ANDRE JKO **

& Malgorzata STEFANIAK**

* Department of Biology and Parasitology. Medical Academy. 20-080 Lublin. Poland
** Department of Insect Pathology. Marie Curie-Sklodowska University, 20-033 Lublin. Poland

ABSTRACT

In a screening bioassay, the antibacterial activity in haemolymph from eight species of millipedes ( Megaphyllum
projection kochi, Ommatoiulus sabulosus, Unciger foeiidus, Polydesmus complanatus. Glomeris connexa,
Strongylosoma pallipes auct., Leptoiulus proximus, Oxidus gracilis ) was compared with cell-free antibacterial immunity
of Galleria mellonella pupae. The only millipedes where lysozyme was not constantly detectable are M. projection kochi
and S. pallipes. In others, the low constitutive titre of lysozyme was unaffected by injections with Enterohacter cloacae
812 or nutrient broth. In Galleria, as for the majority of insects, such previously present antibacterial activity increased
markedly alter preinjection of the pupae with E. cloacae or sterile broth. In pupae of Galleria, the antibacterial activity of
cecropin-like type is induced by E. cloacae as well as by non-living material, the broth. A trace activity against
Escherichia coli D31 was present in untreated Unciger, but this litre did not increase after bacterial inoculation.
Haemolymph from M. projection kochi, P. complanatus and O. gracilis investigated 2 days after injections of either
broth or E. cloacae did not show any inducible antibacterial activity. Injections of E. cloacae into Ommatoiulus. but
broth into Glomeris. induce a measurable antibacterial activity against E. coli D3L

RESUME

Mise en evidence d'une activite antibacterienne dans rhemolymphe de Diplopodes.

Chez huit especes de diplopodes ( Megaphyllum projection kochi . Ommatoiulus sabulosus , Unciger foetidus,
Polydesmus complanatus, Glomeris connexa, Strongylosoma pallipes auct., Leptoiulus proximus, Oxidus gracilis ),
P activity antibacterienne de Ph6molymphe a ete compare & Pimmunite antibacterienne des pupcs de Galleria
mellonella. Les seuls diplopodes chez lesqucls les lyzozymes ne sonl pas constamment decelables sont M. projection
kochi et S. pallipes. Chez les autres especes. on ne note aucun effet a la suite d’injections de Enterobacter cloacae 612 ou
de milieu de culture nulritif. Chez Galleria , comme chez la plupart des insectes. P activity antibacterienne s’accroit
significativement apres injection de la pupe par E. cloacae ou un milieu sterile. Chez la pupe de Galleria, 1 ’ activite
antibacterienne de type cecropine est induite aussi bicn par E. cloacae que par un milieu non-vivant. Unc trace d’activite
anti -Escherichia coli D31 est presente chez Unciger non traite, mais ce taux ne s’accroit pas apres une inoculation
bacteriennc. L’hemolymphe de M. projection kochi, P. complanatus et O. gracilis, etudies deux jours apres injection soil
de milieu de culture, soit de E. cloacae, ne montre aucune activite antibacterienne d£celable. Des injections de E. cloacae
chez Ommatoiulus induisent une activite antibacterienne mesurablc a rencontre de E. coli D31 alors que cet effet est
produil par le milieu de culture chez Glomeris.

Kania, G., JAROSZ, J.. Andrejko. M. & Stefaniak, M., 1996. — Evidence for antibacterial activity in
haemolymph of Diplopoda : preliminary results. In: Geoffroy, J.-J.. Mauries. J.-P. & Nguyen Duy - Jacquemin, M..
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 431-435. Paris ISBN : 2-85653-502-X.

432

GRZEGORZ KANIA. JAN JAROSZ, MARIOLA ANDREJKO & M ALGORZATA STEFAN I AK

INTRODUCTION

Invertebrates can defend themselves against bacterial infections by both cellular (SALT,
1970) and humoral (CHADWICK, 1975) defence mechanisms. A potent humoral immune system
that can be induced by an infection with live non-pathogenic bacteria or injections of abiotic
foreign bodies is specially well characterized in pupae of Lepidoptera (BOMAN & HULTMARK,
1987) which respond to the infection with Enterobacter cloacae by the synthesis of several
classes of immune proteins. Humoral immunity in lepidopterans and other holometabolous
insects is due mainly to the antibacterial action of lysozyme (MOHRING & MF.SSNER, 1968) and
a new class of small basic polypeptides, the cecropins (BOMAN & HULTMARK, 1987).

The presence of lysozyme in normal and immunized arthropods has been reported in
phylogenetically distant species of invertebrates (GOTZ & TRENCZEK, 1991), including
myriapods (XYLANDER & NEVERMANN, 1990). Of inducible bactericidal immune proteins that
are de novo synthesized by several orders of Insecta, cecropins produced in Hyalophora
cecropia and other lepidopterans are the first antibacterial factors well defined biochemically.

In this paper we compared the antibacterial activity in haemolymph from eight species of
millipedes with the pupal Galleria mellonella immune system, using techniques for antibacterial
activity assays developed in the study of cell-free insect immune responses.

MATERIALS AND METHODS

Millipedes

Using a cup plate agar-diffusion assay technique, antibacterial activities of lysozyme and cecropins were detected
in native (non-immune) and immune haemolymph of diplopod species: Megaphyllum projection kochi (Verhoeff),
Ommatoiulus sabulosus (L.), Unciger foetidus (C. L. Koch). Polydesmus complanatus (L.). Glomeris connexa C. L.
Koch. Strongylosoma pallipes (auct.), Leptoiulus proximus (Nemec). Oxidus gracilis (C. L. Koch). All the millipedes
investigated live within woodland litter and soil as their natural habitat. In laboratory, the specimens caged in glass
vessels fed plant litter in varying stages of decomposition. Animals were maintained at 12° C until bleeding for
bioassays of antibacterial activities.

Induction of immune response and sampling of haemolymphs

For immunization, the millipedes were injected into the abdominal haemocoel with either live, log phase
Enterobacter cloacae (0.6 x 104 bacteria per specimen) or sterile nutrient broth (3.0 ml), an abiotic soluble foreign
molecule. By the same way. two day old pupae of Galleria mellonella (Lepidoptera. Pyralidae) taken out of their cocoons
were inoculated with immunizing bacteria or nutrient broth. Fully vigorous, unwounded pupae were served as a control
because of available already evidence on antibacterial immune proteins active in insect immunity (Jarosz, 1993).
Millipedes treated with foreign bodies were incubated for 48 hours at 12°C, but pupae of Galleria at 26°C. Haemolymph
from millipedes were obtained after incision of the intersegmental abdominal cuticle, using a sterile glass micropipette.
Only small volumes of blood may be collected from each individual, but trace amounts of haemolymph obtained from
most of the diplopod species herein investigated were quite sufficient to assay the antibacterial activities in a thin agar
layer with the wells of 0.7 mm in diameter.

Bioassays for antibacterial activities

Lysozyme activity (E C. 3.2.1.17; endo-IWl -4/-N-acetylmuramide glycanohydrolase) was determined in an
inhibition zone assay around the well, using freeze-dried Micrococcus luteus incorporated into an agar medium at a
concentration of 1.0 mg/ml. according to Mohrig & MESSNER (1968). The test for haemolymph lysozyme activity was
conducted in 0.066 M Sorensen buffer (pH 6.4) with 1.0% agarose and 70 mg/ml streptomycin sulfate to inhibit bacterial
contaminations.

Bactericidal activity of cecropins provoked in pupal haemolymph of Galleria was quantified as a diameter of the
lysis zone around the well in a thin agar layer inoculated with an overnight culture of Escherichia coli (about 0.3 x 105
log phase cells per ml), strain D31 sensitive to cecropin-like activity. Haemolymph samples loaded into 0.7 mm
diameter wells cut in the soft (0.7%) agar medium, were incubated at 28°C for 36 hours. Agar medium for assay ol
cecropin activity (but not for lysozyme) contained a trace of phenylthiourea to prevent melanization due to
phenoloxidase activity.

Source : MNHN , Paris

ANTIBACTERIAL ACTIVITY IN HAEMOLYMPH OF DIPLOPODA

433

RESULTS

The antibacterial activity in haemolymph of non-immunized and E. cloacae-, or broth-
injected millipedes was compared with the activity of lysozyme and that of cecropins induced in
Galleria pupae. Though considerable differences in haemolymph lysozyme activity of different
individuals and different diplopod species were noticed, the normally low lysozyme activity was
unaffected by E. cloacae 812 or broth. The normally low lysozyme titer present in untreated
O. sabulosus, U. foetidus and P. complanatus did not increased in specimens injected with
foreign bodies (Table 1). No differences in haemolymph activity were noted between immunized
and non-immunized diplopods. In our screening experiments, haemolymph lysozyme was not
present in untreated M. projection kochi and S. pallipes but a relatively high innate activity of
lysozyme was found in G. connexa and L. proximus. Injections of foreign materials into body
cavity of the millipede O. gracilis did not increase the hardly any detectable the constitutive
antibacterial activity of haemolymph lysozyme. In contrast, the normally high innate lysozyme
titer in Galleria pupae becomes elevated after bacterial infections (0.4 x 105 E cloacae ) so much
as after injections of broth.

Table 1. — Antibacterial activity of lysozyme in haemolymph of non-immunized millipedes and those immunized with
Enterobacter cloacae or sterile nutrient broth. Tr; trace activity; lysis zone diameter less than 1.0 mm ( diameter
of wells; 0.7 mm ). not examined because of difficulty in collecting the haemolymph sample.

Haemolymph lysozyme activity; lysis zone diameter (mm)

Diplopod species

Non-immunized

Specimens immunized with:

Enterobacter cloacae

Sterile nutrient broth

Megaphyllum projection kochi

Ommatoiulus sabulosus

1.5

Unciger foetidus

1.5

1.8

1.6

Polydesmus complanatus

1.7

1.8

Glomeris connexa

5.0

5.5

5.0

Strongylosoma pallipes auct.

Leptoiulus proximus

4.0

Oxidus gracilis

Galleria mellonella

8.0

10.5

10.0

In pupae of Galleria, cecropin antibacterial activity could normally be provoked by both
living E. cloacae and sterile broth. Injections of broth (3.0 ml per animal) into body cavity of G.
connexa can induce within 2 days a measurable antibacterial activity against E. coli D31
(Table 2). A similar but less pronounced antibacterial activity could be generated in
O. sabulosus treated with E. cloacae. Other millipedes injected with E. cloacae (or broth) did not
develop antibacterial activity against E. coli. Trace activity directed against E. coli was found in
immunized and non-immunized U. foetidus. Further investigations are, however, needed to
elucidate if the antibacterial activity appeared in immunized G. connexa and O. sabulosus could
tentatively be classified to an inducible activity of cecropin-like type. Antibacterial activity against
E. coli in the haemolymph of G. connexa and O. sabulosus might be detected only in some

434

GRZEGORZ KANIA. JAN JAROSZ, MARIOLA ANDREJKO & MALGORZATA STEFAN1AK

individuals. These inconsistent results may partly be caused by the difficulty in collecting
enough haemolymph from these millipedes. Furthermore, many specimens of millipedes treated
with E. cloacae died just one day after bacterial injections.

Table 2. — Efforts to induce the cecropin-iike antibacterial activity in haemolymph of Diplopoda by bacterial
infections with Enierobacter cloacae or inoculations of the millipedes with sterile nutrient broth. Tr; trace
activity, lysis zone diameter of E. coli D31 less than 1.0 mm (diameter of the well. 0.7 mm). not examined.

Diplopod species

Cecropin-like activity; lysis zone diameter of E. coli (mm)

Non-immunized Specimens immunized with:

Enierobacter cloacae Sterile nutrient broth

Megaphyllum projectum kochi

Ommatoiulus sabulosus

1.7

Unciger foetidus

Polydesmus complanatus

Glomeris connexa

4.5

Strongylosoma pallipes auct.

Leptoiulus proximus

Oxidus gracilis

Galleria mellonella

8.5

8.0

DISCUSSION

Despite of the progress in insect immunology, only scant evidence are still available about
antibacterial substances, both innate and inducible, conditioning humoral immunity in
myriapods. Intensive research activity of the last few years has led to the identification of new
and interesting groups of peptides with antibacterial activity. It seems that their occurrence is not
restricted to insects. XYLANDER & NEVERMANN (1990) have described at least two antibacterial
substances in haemolymph of Diplopoda; one substance is lysozyme but the other one is
different from lysozyme. Activity against living Micrococcus luteus increased after inoculation
with E. cloacae 612 in two diplopod species, Rhapidostreptus virgator and Chicobolus sp.
Independently of the bacterial strain used as an immunizing agent, growth of E. cloacae , but not
E. coli , was inhibited by haemolymph (XYLANDER & NEVERMANN, 1990). Thus far, the
activity against E. coli was detected only in haemolymph from the millipede Triaenostrepus
triodus (VAN DER WALT et al., 1990), and in haemolymph of immunized diplopods
Ommatoiulus sabulosus and Glomeris connexa (Table 2).

CONCLUSION

The low innate haemolymph lysozyme titer that increases drastically in insects invaded
with non-pathogenic bacteria (MOHRIG & MESSNER, 1968) and inducible antibacterial immune
proteins like cecropins that are synthesized in the fat body of several insect species (BOMAN &
HULTMARK, 1987), but rather absent in millipedes, could confirm the suggestions of
RAVINDRANATH (1973) and NEVERMANN & XYLANDER (1992 ), concerning the crucial role of
cellular immune responses in antibacterial defences of Diplopoda. The phenoloxidase system,
that is considered to be one of the main systems of immune defense in arthropods being
responsible for foreign recognition, killing of microbial invaders, encapsulation of parasites and

Source : MNHN , Paris

ANTIBACTERIAL ACTIVITY IN HAEMOLYMPH OF DIPLOPODA

435

wound healing helps the haemocytic reactions in diplopods since the melanization is found at
wound margins, in the haemocytic wound closure and in haemocytic capsules around foreign
bodies (XYLANDER & BOGUSCH, 1992).

REFERENCES

BOMAN. H. G. & Hultmark. D., 1987. — Ccll-frcc immunity in insects. Ann. Rev. Microbiol .. 41 : 103-126.
Chadwick. J. S., 1975. — Hemolymph changes with infection or induced immunity in insects and ticks, hi : K.

Maramorosh & R. E. Shope. Invertebrate Immunity, New York, Academic Press : 241-271.

Gotz. P. & Trenczek, T.. 1991. — Antibacterial proteins in insects other than Lepidoptera and Diptera and in some
other invertebrates. In : A. P. Gupta, Immunology of Insects and other Arthropods . London. CRC Press, Boca
Raton : 323-346.

Jarosz, J., 1993. — Induction kinetics of immune antibacterial proteins in pupae of Galleria mellonella and Pieris
brassicae. Comp. Biochem. Physiol., 106B : 415-421.

Mohrig, W. & MESSNER, B.. 1968. — Immunoreaktionen bei Insektcn. I. Lysozym als grundlegender antibakterieller
Factor im humoralen Abwehrmechanismus der Insekten. Biol. Zentralbi, 87 : 439-470.

Nevermann, L. & Xylander, W. E. R., 1992. — Cellular immune responses in centipedes and millipedes (Arthropoda,
Tracheata). In : XXV Annual Meeting of Society for Invertebrate Pathology, Heidelberg. Aug. 16 - 21 , Germany :

1 24.

Ravindranath. M. IF, 1973. — The hemocytes of a millipede, Thyropygus poseidon. J. Morphol., 141 : 257-268.
Salt, G., 1970. — The Cellular Defense Reactions of Insects. Cambridge, Cambridge University Press.

Van Der Walt, A.. McClain, A. E., Puren, A. & Savege, N., 1990. — Phylogeny of arthropod immunity. An inducible
humoral response in the Kalahari millipede, Triaenostreptus triodus (Attems). Naturwiss.. 77 : 189-190.

Xylander. W. E. R. & Nevermann, L., 1990. — Antibacterial activity in the hemolymph of Myriapoda (Arthropoda). J.
Invertebr. Pathol., 56: 206-214.

Xylander, w. E. R. & Bogusch, O., 1992. — Investigations on the phenoloxidase of Rhapidostreptus virgator
(Arthropoda, Diplopoda). Zool. Jb. Physiol. .96 : 309-321.

Source : MNHN, Paris

Supernumerary Malpighian Tubules in Chilopods

Carol Constantin PRUNESCU & Paula PRUNESCU

Institute of Biology, 296 Spl. Independentei, RO-79651 Bucharest, Romania

ABSTRACT

In Chilopoda, up to now, one pair of Malpighian tubules was described. These tubules are inserted on each side of the
gut at the junction of the mid and the hind-gut. In Scutigera coleoptrata an additional pair of Malpighian tubules is
present, differing from the main one by its smaller diameter and in being dorsally and ventrally inserted on the gut. at the
same level as the main pair, through small vesicles. In Craterostigmus tasmanianus , in addition to the main Malpighian
tubules, there is a third one, inserted dorsally. in the median plane, and orientated backwards. Animals belonging to
Lithobiidae, Henicopidae and Geophilomorpha show only the main pair. Additional Malpighian tubules of Scutigera and
Craterostigmus may represent plesiomorphic characters.

RESUME

Tubules de Malpighi surnumeraires chez les chilopodes.

Jusqu'fc present, on a decrit chez les chilopodes une seule paire de tubules de Malpighi. Les tubules sont inseres (un sur
la partie gauche, l’autre sur la partie droite de l'intestin) au niveau de la jonction de I'intestin moyen (glandulaire) avec
l'intestin posterieur. Chez Scutigera coleoptrata les deux tubules suppl6mentaires de Malpighi, avec un diametre plus
petit, sont inserts sur la partie dorsale el ventrale de l'intestin au memc niveau que la paire principale. Chez
Craterostigmus tasmanianus , un tubule de Malpighi supplemental, un troisieme, est insure dorsalement. dans le plan
median et oriente vers l'cxtremite posterieure du corps. Les recherches realises chez les Lithobiidae, Henicopidae el
Geophilomorpha ont demontr£ la presence d’une seule paire de tubules de Malpighi. La presence de tubules de Malpighi
surnumeraires chez Scutigera et Craterostigmus peut etre consideree comme un caractere plesiomorphe.

INTRODUCTION

All previous data concerning the anatomy of the Malpighian tubules indicates the existence
of only one pair of Malpighian tubules in chilopods (LEWIS. 1981). A study of the microscopic
anatomy permits the description of some supplementary Malpighian tubules in Scutigera
coleoptrata and Craterostigmus tasmanianus.

MATERIAL AND METHOD

Specimens of S. coleoptrata (males, females and larvae) were collected in Sicily (Italy) in 1969 and in Dobrogea
(Romania) between 1967 and 1992. Fixation was made in Bouin’s solution or 70% ethylic alcohol. A material
constituted by the caudal part of the body of two females of C. tasmanianus , embedded in paraffin, was offered to us by S.
M. Manton in 1964. This material was from Tasmania. Also from Tasmania were some females and males of C.
tasmanianus gathered by R. Mesibov in 1991. This material was fixed in formaldehyde-calcium (S. M. M anton's lot) or
in glutaraldchyde 2.5% in cacodylat buffer (R. Mesibov’s lot). The material was processed according normal histological
technique.

Prunescu, C. C. & PRUNESCU, P.. 1996. — Supernumerary malpighian tubules in chilopods. In: Geoffroy,
J.-J., MAURlfcS, J.-P. & NGUYEN Duy - Jacquemin. M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat ., 169 :
437-440. Paris ISBN : 2-85653-502-X.

438

CAROL CONSTANTIN PRUNESCU & PAULA PRUNESCU

RESULTS

In Scutigera coleoptrata , transverse serial sections, taken through the junction of the mid¬
gut and hind-gut, confirmed the opening of the two main Malpighian tubules in the horizontal
(bilateral) plane (Figs 1-2). Before the opening, each of the Malpighian tubules present an
ampulla which narrows at the level of the opening in the intestine. Approximately at the same
level, two other Malpighian tubules open into the intestine, but in a dorso-ventral plane. One of
these Malpighian tubules opens in the median-dorsal plane and the other in the median-ventral
plane (Figs 3-5). Before opening into the intestine, each of the dorso-ventral Malpighian tubule
presents its own, small ampulla. The dorso-ventral Malpighian tubules are extended and coiled
along the mid-gut, like the main pair of Malpighian tubules. We cannot make any statement
about the length of the dorso-ventral Malpighian tubules.

Fig. I. — Lateral Malpighian tubules opening into the intestine (S. coleoptrata), H-e. x80.

Fig. 2. — Detail of the opening of the lateral Malpighian tubules at the level of the junction of the mid and hind-gut
(5. coleoptrata) xl60.

Fig. 3. — Opening of the dorso-ventral Malpighian tubules into the gut (5. coleoptrata) x80.

Fig. 4. — Detail of Fig. 3. x200. The arrow shows the top of the slide.

Fig. 6. — Dorsal Malpighian tubule (arrow) (C. tasmanianus) x400. (see next page Fig. 6-9).

Fig. 7. — Dorsal Malpighian tubule drawn near the dorsal wall of the gut (arrow). — The lateral Malpighian tubules
ampullas open into the intestine (C. tasmanianus). x!20.

Fig. 8. — Dorsal Malpighian tubule opening into the dorso-median zone of the intestine (arrow) (C. tasmanianus) xl20.
Fig. 9. — Detail of the opening of the dorsal Malpighian tubule (arrow). (C. tasmanianus) x400.

Source : MNHN. Paris

SUPRANUMERARY MALPIGHIAN TUBULES IN CHILOPODS

439

Fig. 5. — Schematic representation of
the insertion of the Malpighian
tubules of Scutigera coleoptrdta.
A: dorsal view;

B: Transversal section;

1; lateral Malpighian tubule;

2: dorsal Malpighian tubule;

3: ventral Malpighian tubule;

4; hind-gut;

5: vas dorsalis;

6: ventral nerve ganglia.

In Craterostigmus tasmanianus, the laterally-inserted Malpighian tubules show an ampulla
of great size at their proximal end (Figs 7-8), by which they open into the intestine. At the same
level (the junction of the mid and the hind-gut) in the medio-dorsal plane, opens a short and
relatively thin Malpighian tubule (Figs 6-10), which has its own, small ampulla. This tubule
follows a sinuous line along the hind-gut. It is oriented towards the posterior extremity of the
body. The dorsal Malpighian tubule has the distal extremity blindening. The histological
structure of the dorsal Malpighian tubule is similar to that of the main lateral Malpighian tubules
(Fig. 6). The epithelium of the tubule is of cuboid shaped cells with brush borders.

Source

440

CAROL CONSTANTIN PRUNESCU & PAULA PRUNESCU

The dorso-ventral Malpighian tubules of S. coleoptrata show a similar histological
structure. Examination of the serial sections of the contact zone between the mid and hind-gut in
many species of Lithobiomorpha, Scolopendromorpha and Geophilomorpha confirmed the
presence of the main pair of bilaterally-inserted Malpighian tubules. We never found Malpighian
tubules with a dorsal or ventral insertion in any representatives of these orders.

1 --

Fig. 10. — Schematic representation of the
insertion of the Malpighian
tubules of C. tasmanianus.

A: dorsal view;

B: transverse section;

1: lateral Malpighian tubule;

2: dorsal Malpighian tubule;

3: hind-gut;

4: vas dorsalis;

5: ventral nerve ganglion.

DISCUSSION

This work does not present data concerning the function of the Malpighian tubules with a
dorso-ventral or ventral insertion. Sections through these Malpighian tubules show a similar
histological structure to those with a lateral insertion (PALM, 1953; BERTHEAU, 1971).

The presence of a pair of Malpighian tubules with dorso-ventral insertions in S. coleoptrata
suggests that this feature could represent a plesiomorphic character (PRUNESCU, this volume).

Since, during evolution, different systems and organs in Chilopoda have become simpler,
the existence of four Malpighian tubules, in an order which present numerous plesiomorphic
characters, may be considered as a plesiomorphic feature. The disappearance of the dorso-
ventral Malpighian tubules in the more evolved groups may be considered as an apomorphic
feature.

C. tasmanianus represents the archetype of the epimorphic Chilopoda, meaning that their
ancestor presented many of the subsequent features of epimorphic Chilopoda. The presence, in
this primitive type, of a Malpighian tubule homologous to the dorsal Malpighian tubule in
Scutigeromorpha, confirms the plesiomorphic nature of this supernumerary tubule.

REFERENCES

Bertheau, P., 1971. — Histologie comparee des tubes de Malpighi de quelques Chilopodes (Myriapodes). C. R. Acad.
Sci., Paris. 212 : 2913-2915.

Lewis. J. G. E., 1981. — The biology of Centipedes. Cambridge. Cambridge, Univ. Press. 475 pp.

Palm. N. B., 1953. — The elimination of injected vital dyes from the blood in Myriapods. Ark. Zool., Ser. 2. 6:219-

246.

Source :

The Structure and Possible Function of the Spiracles of
some Scolopendridae (Chilopoda, Scolopendromorpha)

John G. E. LEWIS *, Trevor J. HILL * & Gavin E. WAKLEY **

* Taunton School, Taunton, Somerset TA2 6AD, U. K.

** Department of Biological Sciences, Washington Singer Laboratories, University of Exeter.

Perry Road, Exeter EX4 4QG, U. K.

ABSTRACT

The results of a scanning electron microscope investigation into the structure of the spiracles of four species of
scolopcndrid centipedes are reported. Rhysida nuda togoensis (Kraepelin), Ethmostigmus trigonopodus (Leach).
Scolopendra morsitans L., Scolopendra valida Lucas, were studied. The spiracles serve to prevent debris entering the
tracheal system. The relatively simple spiracles of Rhysida and Ethmostigmus may function as a plastron in small
specimens. The more complex spiracles of Scolopendra spp. may function principally to prevent water loss, although it
is possible that the large sub-atrial cavities in S. morsitans may form a plastron. R. nuda and E. trigonopodus are absent
from arid habitats.

RESUME

Structure et fonction probable des spiracles de quelques Scolopendridae (Chilopoda,
Scolopendromorpha).

L’ ultrastructure des spiracles de Scolopendridae est etudiee chez Rhysida nuda togoensis (Kraepelin). Ethmostigmus
trigonopodus (Leach), Scolopendra morsitans L.. et Scolopendra valida Lucas. La fonction des spiracles est
essentiellement d’empecher Pentree de debris dans le systeme tracheen. Us sont simples chez Rhysida et Ethmostigmus ,
qui manquent dans les habitats aridcs. et fonctionnent comme plastron chez les petits individus. Plus complexes chez
Scolopendra sp., ils pourraient fonctionner comme preventifs du dessechement, bien qu'il soit possible que la grande
cavite sous-atriale de S. morsitans forme un plastron.

INTRODUCTION

The literature on the structure of centipede spiracles has been reviewed by VERHOEFF
(1941) and LEWIS (1981). Most genera of the order Scolopendromorpha have 21 leg-bearing
segments and in these spiracles are present on segments 3. 5. 8. 10. 12. 14. 16. 18 and 20 and
sometimes 7.

In genera with 23 leg-bearing segments spiracles are, in addition, present on segment 22.
The genus Plutonium is unusual in having spiracles on every leg-bearing segment except the first
and last.

The scolopendromorphs show a considerable variation in spiracle structure. In the family
Cryptopsidae the elliptical spiracle in Cryptops leads to the atrium which itself opens by a

Lewis, J. G. E.. Hill, T. J. & Wakley, G. E., 1996. — The siructure and possible function of the spiracles of some
Scolopendridae (Chilopoda. Scolopendromorpha). In: Geoptroy. J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M.,
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 441-449. Paris ISBN : 2-85653-502-X.

442

JOHN G. E. LEWIS. TREVOR J. HILL & GAVIN E. WAKLEY

crescent-shaped slit into a subatrial cavity (FULLER, 1960): both cavities are lined by trichomes.
In Otocryptops the atrium is funnel-shaped and there is no subatrial cavity (VERHOEFF, 1941).
The two subfamilies of the Scolopendridae are distinguished by the structure of their spiracles.
In the Otostigminae the spiracles are mostly rounded and without valves (Fig. 1 A) whereas the
Scolopendrinae have triangular spiracles with three-flapped valves (Fig. 3A).

MATERIALS AND METHODS

Four species have been investigated using the scanning electron microscope, namely: Rhysida nuda togoensis
Kraepelin, Eihmosligmus trigonopodus (Leach) and Scolopendra morsitans L., all from Nigeria, and Scolopendra valida
Lucas from Oman.

The material, which had been preserved in 70 per cent ethanol, was dehydrated in absolute ethanol for at least 24
hours and then air dried, sputter coated with gold and then examined in a Cambridge I00S scanning electron microscope.
Preparations for examination under the light microscope were mounted in Hoyer's mountant.

RESULTS

Subfamily Otostigminae

Rhysida nuda togoensis Kraepelin

The spiracles of Rhysida are approximately elliptical (Fig. 1A), the axis of the ellipse
sloping obliquely forwards and upwards on segment 3 but more or less vertical on the posterior
spiracles, the lower border less curved than the upper. The first spiracle, is almost twice the
length of the subsequent ones. The peritrema is scalloped. The atrial wall is thrown into a
number of vertical ridges and the floor into humps (Fig. IB. C). The atrial surface is covered by
complex trichomes which are of variable shape. Those immediately beneath the peritrema have
angular heads but most are elongated. The sides show a reticulate strutting so that they are
honey-combed with cavities (Fig. ID). The ridge-like trichomes are 1 1 pm high, 10-14 pm long
and 1.25-2.50 pm wide. The wide tracheae open between the humps of the atrial floor. Their
openings are surrounded by digitate trichomes whose surfaces are covered by a network of
ridges (Fig. IE). They are 60 pm long and are here termed guard hairs.

Ethmostigmus trigonopodus (Leach)

The general structure of the spiracle of Ethmostigmus ( =Heterostoma) was accurately
described by Haase (1884) and by VERHOEFF (1941). As in Rhysida the spiracles are
approximately elliptical but the first spiracle (Fig. 2A) is particularly large and the atrium saucer¬
shaped, its floor being only slightly below the level the surrounding stigmatopleurite (Fig. 5 A).
The subsequent spiracles become progressively more bowl-like and in small specimens resemble
those of Rhysida. The floor of the spiracle (Fig. 2B) is thrown into large humps or ridges
covered with trichomes. Those on top of the humps have scalloped heads 7-12 pm across (Fig.
2C). These were described as six-pointed stars by VERHOEFF. The sides and bases show
reticulate strutting (Fig. 2E). The trichomes become more elongated towards the base of the
humps so that they resemble those of Rhysida. The narrow trichomes are 9-1 1.4 pm long, 1.4-
1.8 pm wide and 10 pm high. The tracheae open at the bases of the humps, their openings
protected, as in Rhysida. by elongated guard hairs 64 pm long (Fig. 2D, E).

Subfam ily Scolopendrinae

Scolopendra morsitans Linnaeus

The structure of the spiracle is very similar to that described for 5. cingulata Latreille by
Haase (1884) and CHALANDE (1885). It is triangular, the apex being anterior (Fig. 3 A). The
peritrema is scalloped, most lobes having a short seta centrally.

Source :

STRUCTURE AND POSSIBLE FI 'NOTION OF THE SPIRACLES OF SOME SCOLOPENDRIDAE

443

Fig. I. — Rhysida nuda logoensis. Spiracle of segment 3. A. Surface view. B. Vertical section. C. Detail of wall of
atrium. P. peritrema. D. Atrial trichomes. E. Guard hairs of tracheal openings.

The atrium is divided into outer and inner cavities by a three flapped valve (Fig. 3B. 5B).
The inner atrial cavity is often termed the sub-atrial cavity. Beneath the peritrema the atrial wall
bears ridged columnar trichomes 8 pm high, these increase in length towards the valves (Fig.
3D). At the base of the outer atrial cavity there is a row of setose cones (CHALANDE's recumbent
plumes) pointing vertically towards the opening of the spiracle (Fig. 3B. C). On the spiracle of
segment three there are 8 on the posterior valve and 21-22 on the dorsal and ventral valves. They
are about 100 pm high and the longest setae or bristles are 41 pm long. They fill much of the

Source :

444

JOHN G. E. LEWIS. TREVOR J. I IILL & GAVIN E. WAKLEY

outer atrial cavity, having hut a narrow Y-shaped aperture between them. Beneath the row of
cones is a band of vertically ridged cuticle devoid of trichomes which forms a valve. The
subatrial cavity is extensive, with deeply folded walls covered with trichomes (Fig. 5B). These
are 4.6 pm high, their irregularly pitted heads measuring about 6 pm across. The sides show
reticulate strutting which is continued on the atrial floor between the trichomes (Fig. 3E). The
tracheae open into the floor and sides of the inner atrial cavity, their openings being surrounded
by guard hairs like those of Rhysida and Ethmostigmus (Fig. 3F). These are about 40 pm long.

Fig. 2. — Ethmostigmus trigonopodus. Spiracle of segment 3. A. Surface view. B. Vertical section (detail). C. Surface
view of atrial trichomes. D. Guard hairs of tracheal opening. E. Detail of guard hairs and trichomes.

Scolopendra valida Lucas

The spiracle of S. valida (Fig. 4A) resembles that of 5. morsitans in shape and its division
into an outer and inner atrial cavity. The wall of the outer atrial cavity bears trichomes similar to
those of S. morsitans (Fig. 4B. C). The setae are, however, not borne on cones but form a
dense strip along the top of each valve. The valves are composed of trichome free cuticle (Fig.
4D). The inner atrial cavity is not enlarged like that of 5. morsitans and lacks trichomes. The
tracheae open into the chamber, their openings being surrounded by guard hairs 60 pm long.

Fig. 3. — Scolopendra morsitans. Spiracle of segment 3. A. Surface view. B. Vertical longitudinal section. C. Setose
cone. D. Trichomes of outer atrial cavity. E. Trichomes of inner atrial cavity. F. Guard hairs.

Source :

STRUCTURE AND POSSIBLE FUNCTION OF THE SPIRACLES OF SOME SCOLOPENDRIDAE

445

Source : MNHN, Pahs

446

JOHN G. E. LEWIS. TREVOR J. I IILL & GAVIN E. WAKLEY

Fig. 4. — Seolopendra valida. Spiracle of segment 3. A. Surface view. B. Atrial trichomes and setae. C. Detail trichomes.
D. Vertical longitudinal section. V, valve.

Source : MNHN. Paris

STRUCTURE AND POSSIBLE FUNCTION OF THE SPIRACLES OF SOME SCOLOPF.N DRI DAE

447

DISCUSSION

Spiracles function to reduce water loss from the tracheal system under dry conditions and,
in arthropods that experience immersion in water, they frequently act as plastrons. The trichomes
may be involved in both these processes. Other functions that have been suggested for spiracular
trichomes are that they filter out dust (KAUFMANN, 1962) and prevent spiracular occlusion
during locomotion (CURRY, 1974). PUGH et al. (1991) suggested that the peritreme of
holothyrid mites might prevent suffocation during immersion or act as a water trap.

Spiracular function in Otostigminae

Water loss from the tracheal openings of Rhysida and Ethmostigmus may be impeded by
the spiracular guard hairs which will also prevent debris entering the tracheae. The crevices
between the humps of the atrial floor will also retain humid air. It is difficult to visualise a role
for the trichomes in this respect as gases diffusing in and out of the tracheae will pass over them
rather than between them. A more likely function is that they form a plastron, retaining a layer of
air when the centipede is immersed in water as may happen during the rainy season.

HINTON (1968) determined the basal limit of plastron efficiency for insects in terms of the
ratio between the area of the plastron and the wet body weight as 1.5 x 104 pm2.mg-i.
Assuming that the air-water interface is across the tops of the trichomes. a conservative estimate
for this ratio for a large specimen of R. nuda togoensis length 74 mm, mass 1080 mg is 3.75 x
103 pm2.mg-i : well below HlNTON's figure. For a small specimen body length 13 mm, mass
11 mg the figure is 1.5 x 104 pm2.mg-i, equal to HlNTON's minimum value. In an E.
trigonopodus length 88 mm. mass 2700 mg the ratio is 4.2 x 103 pm2.mg-i but in a small
specimen length 29.5 mm, mass 91 mg the value is 2.2 x 104 pm2.mg-L It would appear that in
both species small but not large specimens may be able to utilise plastron respiration. These
calculations assume that the interface is across the tops of the trichomes.

Although the relative area of a plastron decreases with increased mass of the organism,
tracheal volume will increase in proportion to increasing mass. The tracheae of large specimens
appear to be particularly voluminous and may function as air stores during immersion.

Spiracular function in Scolopendrinae

It is tempting to suggest that the greater complexity of the scolopendrine spiracle, with the
atrium divided horizontally by a three-flapped valve and the presence in Scolopendra spp. of
dense setae above the valves either borne on cones or not, is related to the need to restrict water
loss in dry conditions. PUGH et al. (1987) described structures similar to the setose cones from
the pcritrematic groove of the mite Phaulodinychus repleta (Berlese). They consist of
micropapillae arranged on Christmas tree-like structures and termed compound fimbriae. They
suggested that the compound fimbriae of P. repleta carried out a protective function preventing
the entry of foreign/harmful material into the tracheal system rather than supporting an air film.
The irregular and spiky compound fimbriae of Holothyrus coccinella (Wormersley) cannot
support an airfilm but would pierce the air water interface (PUGH et al. , 1991 ). The setae of the
setose cones of S. morsitans and the setae of S. valida clearly act as sieves and are often covered
with debris. The setae will clearly reduce diffusion. If their surface is not hydrophobe then
dipole-dipole interactions between water molecules and the protein and chitin molecules of the
cuticle will allow free diffusion of oxygen and carbon dioxide whilst impeding that of water. The
spiracles of the Scolopendra species are small and the area covered by trichomes in the atrial
cavities is low. The ratio between the area and body mass for a S. morsitans length 70 mm,
mass 1042 mg is 1.6 x 103 pm2.mg-i, lc. An order of magnitude below HlNTON's figure. The
figure for a S. valida length 1 10 mm, mass 4370 mg is even lower: 3.7 x 102 pm2.mg-i. S.
morsitans. though not S. valida, has large sub-atrial cavities lined with trichomes (Fig. 5B). If

448

JOHN G. E. LEWIS. TREVOR J. HILL & GAVIN E. WAKLEY

these were flooded with water it is possible that they would act as a plastron. Currently,
however, there are insufficient data to calculate the area involved.

Fig. 5. — A. Vertical longitudinal section of spiracle of segment 3 of Ethmostigmus trigonopodus. B. Vertical
transverse section of spiracle of segment 3 of Scolopendra morsitans. IAC, inner atrial cavity; OAC, outer atrial
cavity; P. peritrema; V, valve.

Ecology

Data on the distribution of scolopendrids in West Africa and Saudi Arabia shows that
members of the subfamily Scolopendrinae with their triangular spiracles occur in dry and humid
regions, whereas members of the subfamily Otostigminae are absent from drier habitats. Thus
Rhysida nuda togoensis and Ethmostigmus trigonopodus are virtually absent from the dry Sudan
and Sahel savanna regions of Nigeria (LEWIS, 1972) whereas the scolopendrines Asanada
socotrana Pocock and S. morsitans are widespread there (LEWIS, 1973 and unpublished data).

Rhysida and Ethmostigmus have not been recorded from Saudi Arabia but A. socotrana
and three species of Scolopendra ( canidens Newport, mirabilis (Porat) and valida Lucas) occur
there (LEWIS, 1986).

In the guinea savanna region of Northern Nigeria, R. nuda and E. trigonopodus are
virtually absent from surface habitats during the latter part of the dry season (mid-November to

Source : MNHN, Paris

STRUCTURE AND POSSIBLE FUNCTION OF THE SPIRACLES OF SOME SCOLOPENDRIDAE

449

April) but S. morsitans is surface active throughout the year being found under cow dung during
the dry season (LEWIS, 1969). The ecological data support the conclusions drawn about possible
spiracular functions on the basis of morphological observations.

ACKNOWLEDGMENTS

This work was carried out between 1990 and 1993 with a series of sixth form pupils from Taunton School. The
major participants were Susan Badley. Tom Basher, Tom Blandford, Nicola Irvin, Helen Jewell, Katie Newbold,
Philip Smith, Robert Tudor and Paul Yeung. It was supported by generous grants to J. G. E.Lewis from the Royal
Society and the Association for Science Education Research in Schools Committee which are gratefully acknowledged.
J.G.E.Lewis’s thanks are also due to Dr D. J. Stradling of Exeter University for continued advice and support which are
much appreciated. Professor J. A. Bryant and Dr M. R. Mcnair kindly allowed us to use the SEM facilities in the
Washington Singer Laboratories.

REFERENCES

Chalande, J., 1885. — Recherches anatomiques sur I’appareil respiratoire chez les chilopodes de France. Bull. Soc.
Hist. rial. Toulouse , 19 : 39-66.

Curry, A., 1974. — The spiracle structure and resistance to desiccation of centipedes. Symp. zool. Soc. Lond. ,32 :
365-382.

Fuller, H.. 1960. — Untersuchungen liber den Bau der Stigmen bei Chilopoden. Zool. Jb. (Anat.), 78 : 129-144.

Haase, E., 1884. — Das Respirationssystem der Symphylen und Chilopoden. Zool. Beitr.. 1 : 65-96.

HINTON. H. E.. 1968. — Spiracular gills. Adv. Insect Physiol.. 126 : 65-162.

Kaufman, Z. S., 1962. — The structure and development of stigmata in Lithobius forficatus L. (Chilopoda,
Lithobiidae). Ent. Obozr., 41 : 223-225. (In Russian with English summary).

Lewis, J. G. E., 1969 (70). — The biology of Scolopendra amazonica in Nigerian Guinea savannah. Bull. Mus. nail.
Hist, nat., Paris. 41, suppl. n°2 : 85-90.

LEWIS, J. G. E.. 1972. — The life histories and distribution of the centipedes Rhysida nuda togoensis and Ethmostigmus
trigonopodus (Scolopendromorpha, Scolopendridae) in Nigeria. J . Zool., Lond.. 197 : 399-414.

Lewis, J. G. E., 1973. — The taxonomy, distribution and ecology of centipedes of the genus Asanada
(Scolopendromorpha, Scolopendridae) in Nigeria. Zool. J. Linn. Soc.. 52 : 97-112.

Lewis, J. G. E., 1981. — The biology of centipedes. Cambridge, Cambridge University Press, 476 pp.

Lewis, J. G. E., 1986. — Centipedes of Saudi Arabia. Fauna of Saudi Arabia. 8 . 20-30.

Pugh, P. J. A., King, P. E. & Fordy, M. R., 1987. — Structural features associated with respiration in some intertidal
Uropodina (Acarina: Mesostigmata). J. Zool., Lond.. 211 : 107-120.

Pugh, P. J. A., Evans, G. O., King, P. E. & Fordy. M. R. & King, P. E., 1991. — The functional morphology of the
respiratory system of the Holothyrida (=Tetrastigmata) (Acari: Anactinotrichida). ./. Zool.. Lond., 225 : 153-172.
VERHOEFF, K. W., 1941. — Zur Kenntnis der Chilopodenstigmen. Z. Morph. Okol. Tiere.. 38 : 96-1 17.

Source : MNHN, Paris

Population Metabolism of Millipedes at
Two Altitudinal Zones in the Central Alps

(Tirol, Austria)

Erwin MEYER, Peter MARSONER & Elisabeth FISCHER

Institute of Zoology, University of Innsbruck
Technikerstr. 25, A-6020 Innsbruck, Austria

ABSTRACT

The respiratory metabolism of Enantiulus nanus (Latzel, 1884) from a mixed oak wood (670 m a.s.l.) and of 3
subalpine species, Leptoiulus saltuvagus (Verhoeff, 1898), Ochogona caroli (Rothenbuhler, 1900) and Haasea
fonticulorum (Verhoeff, 1910) from an Alnus viridis community (2000 m) was measured using a Gilson respirometer and
a Warburg respirometer with electronic manometers. Temperature (6-20°C) and mass-specific ( E . n .: 2-17 mgfw; L. s.\ 3-
75 mgfw; O. c.\ 1-10 mgfw; //./.: 2-12 mgfw) oxygen consumption values of these species are presented. The lowland
species has its greatest sensitivity to temperature changes between 10°C and 15°C, the subalpine species between 6°C
and 10°C with Qio-values between 3.1 and 7.9. Based on a mean E. nanus- biomass of 2.9 g fresh mass per m2 (= 73% of
the total millipede-biomass) in the oak wood, the population metabolism of this species equalled 1488 ml O: m-i and
year (= 30 KJ). In contrast a total millipede biomass of 2.2 g fresh mass per m2 in the subalpine site respires only 732 ml
O2 m-i and year (= 15 KJ). The most important single factor in determining the metabolism for field populations is its
population structure and biomass.

RESUME

Metabolisme de populations de diplopodes dans deux zones altitudinales des Alpes Centrales
(Tyrol, Autriche).

Le metabolisme respiratoire de Enantiulus nanus (Latzel, 1884) d’une foret mixte de chene (altitude 670 m) et de trois
especes subalpines, Leptoiulus saltuvagus (Verhoeff, 1898), Ochogona caroli (Rothenbuhler, 1900) el Haasea
fonticulorum (Verhoeff. 1910) d’un peuplement h Alnus viridis (altitude 2000 m) a ete mesure a I’aide d'un respirometre
de type Gilson et d un respirometre de type Warburg equipes de manometres 61ectroniques. On donne ici les temperatures
(6-20°C), les masses sp6cifiques fraiches (£. n. : 2-17 mg mf; L. s. : 3-75 mg mf; H. f : 2-12 mg mf) et les valeurs
respectives de consommation en oxygene. L’espece de plaine pr£sente une plus grande scnsibilite aux changements de
temperature entrc 10°C et 15°C, les especes alpines entre 6°C et 10°C, avec des valeurs de Q10 variant de 3,1 a 7.9. En se
basant sur une biomasse moyenne pour E. nanus egale a 2,9 grammes de matiere fraiche par metre carre (qui represente
73% de la biomasse totale des diplopodes) dans la chenaie, Ie metabolisme respiratoire annuel de cette population
equivaut a 1488 ml O2 m-i (= 30 KJ). En comparison, une biomasse totale de diplopodes de 2,2 g de mature fraiche par
m&tre carre ne repr£sente, dans les sites subalpins, qu’une respiration de 732 ml O2 m-i (= 15 KJ). Le facteur le plus
important dans le determinisme du metabolisme respiratoire des populations sur le terrain semble etre la structure du
peuplement et la biomasse.

Meyer, E., Marsoner, P. & Fischer, E., 1996. — Population metabolism of millipedes at two altitudinal zones
in the Central Alps (Tirol, Austria). In: Geoffroy, J.-J.. Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (cds), Acta
Myriapodologica. Mem. Mus. natn. Hist, nat ., 169 : 451-460. Paris ISBN : 2-85653-502-X

452

ERWIN MEYER. PETER MARSONER & ELISABETH FISCHER

INTRODUCTION

In a recent contribution PENTEADO el al. (1991) have summarized available data on oxygen
consumption in millipedes. Respiratory rates of more than 20 species living in temperate or
tropical regions, ranging in size from 10 to 4000 mg have been evaluated and discussed in
relation to size of individual and sex. Further variables such as temperature, life stages or
decreased oxygen tension have been investigated in numerous papers such as DWARAKANATH
(1971), Gromysz-Kalkowska (1970, 1973), Gromysz-Kalkowska & Stojalowska
(1966) or PENTEADO & HEBLING-BERALDO (1991). Data on oxygen consumption rates of
millipedes in context with the life-cycle and population structure are rare. WOOTEN &
CRAWFORD (1974) gave such an example by combining monthly measurements of the
respiration rate of a desert millipede with its behaviour in the field.

The aim of the present paper is to combine detailed laboratory investigations of the
respiratory rates of four alpine millipede species (FISCHER, 1985; MARSONER, 1992) with the
results from studies on the population structure, life-cycle and temperatures in their habitats
(Meyer, 1979, 1985; KOFLER & Meyer, 1992). In this way it is possible to evaluate the
relative importance of variables such as age structure, biomass, time of the year, temperature and
altitude affecting the population metabolism in the field.

MATERIAL AND METHODS

Specimens of Enantiulus nanus (Latzel) (2-17 mg live mass) were collected from an inneralpine mixed oak wood
(670 m a.s.l.) in the Inn-valley near Stams at four occasions between October 1990 and October 1991. The life-history,
abundance and production of this species have been studied in detail by Kofler & Meyer (1992). The annual mean
abundance of E. nanus is 859 inds m*2 and this species dominates the total millipede fauna in this oak wood. Males reach
maturity first as stadium VIII in the 3rd year, females also become adult with stage VIII and lay eggs after the third
overwintering. This iteroparous julid species shows a seven-year life-cycle. A mean overwintering biomass of 2.9 g
fresh mass m-2 produces 1.5 g fresh mass m-2 year-f The animals were kept in plastic boxes provided with tap water agar
to maintain a near 100% humidity and fed dead leaves. These cultures were maintained at field-like conditions with
fluctuating temperatures (night/day): March, April and October: 7.5°C/1 2.5°C, May, June and September:
12.5°C/17.5°C, July and August: 17.5°C/22.5°C. November: 2.5°C/7.5°C. Approximately 470 inds of E. nanus were used
in the assessment of respiratory metabolism between March and November 1991. Experimental temperatures were: 5°C
(November), 10°C (March, April, October). 15°C (May, June, September) and 20°C (July and August). Rates of oxygen
uptake of individual specimens were measured using a Warburg respirometer with electronic manometers connected to a
module box and a PC. During each experiment 13 respirometer flasks (Volume: 2. 2-2. 6 ml) contained animals and one
acted as a thermo-barometer-control. In each flask CO2 was absorbed by a 5% solution of NaOH (20 pi) pipetted into the
lower part of a two-piece flask. The animals were placed into the upper part of the flask which had a sintered floor to
allow gas exchange. A high humidity was maintained by inserting a damp and crumpled piece of filter paper. After
introducing the individuals into the flasks an half hour settling period was allowed to elapse before closing the
respirometer valves. Each experiment lasted at least 24 h. At the end of the experimental period the millipedes were
weighed individually. Oxygen consumption was calculated from a series of usually 144 measurements (10 min measuring
intervals) by excluding the first hour (6 measurements) after closing the valves.

Specimens of Leptoiulus saltuvagus (Verhoeff) (3-75 mg life mass), Ochogona caroli (RothenbLihler) ( 1 - 1 0 mg
life mass) and Haasea fonticulorum (Verhoeff) (2-12 mg life mass) were collected from an Alnetum viridis (2000 m a.s.l.)
in the Otztal Alps near Obergurgl at several occasions between August and October 1984. Previous investigations by
Meyer (1979, 1985) established for L. saltuvagus a mean density of 209 inds m-2 and a biomass 1.2 g fresh mass m-2.
The semelparous julid species reaches maturity in stage IX, X or XI after four or five years. At the same site the two
chordeumatid species have a mean density of 112 inds m-2 (O. caroli ) and 107 inds m-2 (//. fonticulorum ). This
corresponds with a mean biomass of 0.42 g fresh mass m-2 ( O . c.) and 0.63 g fresh mass m-2 ( H . /). A three-year life-
cycle is probable for both chordeumatid species.

The animals were kept in light-temperature chambers representing approximately field conditions (day/night-
temperature: I2°C/8°C, 12/12 h) in plastic boxes provided with tap water agar to maintain a near 100% humidity and fed
dead leaves. Approximately 80 individuals of L. saltuvagus and 40 individuals each of O. caroli and H. fonticulorum were
used in the assessment of respiratory metabolism. Between July 1984 and February 1985 rates of oxygen uptake of
individual specimens of the three species were measured using a refrigerated GR-14 Gilson differential respirometer.
Measurements were made at 6°C, 10°C, 15°C and 20°C. During each experiment 9-12 respirometer flasks (Volume: 25 ml)
contained animals and three acted as controls. In each flask CO2 was absorbed by 5N NaOH pipetted onto a roll of filter
paper held in the centre well. A perforated plastic cylinder placed over the central wall prevented the animals making
contact with the NaOH. A high humidity was maintained with moist filter paper on the floor of the chamber. Two damp

Source : MNHN. Paris

POPULATION METABOLISM OF MILLIPEDES IN CENTRAL ALPS

453

and crumpled I cm2 pieces of the same material provided shelter and the animals usually became quiescent within 1 h of
being placed in the vessels. Because of the respirometer response and precision, 3 individuals of the smallest size classes
were placed in each chamber.

Temperatures in the litter layer of the two sites were recorded with a Goerz Thermoscript (clockwork mechanism
and bimetallic probe).

RESULTS

The relationship between respiratory ’ rate and size of individual

Enantiulus nanus (Latzel, 1884)

As indicated in Table 1, the smallest specimens of E. nanus consume at least twice as
much oxygen per unit mass than the largest ones at all experimental temperatures between 5°C
and 20°C. The b-values according to the equation R = a Wb where R is the respiratory rate
expressed in |il oxygen ind-i are scattered between 0.012 (5°C, Nov.), 0.205 (10°C, Oct.),
0.236 (15°C, June), 0.306 (20°C, Aug.), 0.329 (15°C, May), 0.441 (15°C, Sept.) and 0.488
(20°C, July) depending on temperature and time of the year.

Table I. — Mean ( x±S.E.) rates of oxygen consumption per unit mass (pi mg-i h-i) over the size range of Enantiulus
nanus at 5°C, I0°C, I5°C and 20°C.

Size range (mg)

5°C

10°C

15°C

20°C

1.6 -

4.9

0.0910.01

0.1310.03

1 1

0.2110.03

0.2210.03

5.0-

6.9

0.0510.01

0.0810.01

0.1410.01

7.0 -

10.9

l 1

0.0410.01

0.0610.01

0.1 110.01

0.1210.01

11.0 -

16.6

0.0310.01

0.0510.01

0.0810.01

0.1010.01

Leptoiulus saltuvagus (Verhoeff, 1898)

Table 2 shows the respiratory rates of L. saltuvagus. Again juveniles consume
considerably more oxygen per unit mass than adults. The calculated b-values are between 0.67
(10°C), 0.68 (20°C), 0.82 (15°C) and 0.92 (6°C).

Table 2. — Mean(x+S.E.) rates of oxygen consumption per unit mass (pi mg-i h-l) over the size range of Leptoiulus
saltuvagus at 5°C. I0°C, I5°C and 20°C.

Size range (mg)

6°C

10°C

15°C

20°C

3.0 - 16.0

0.0810.01

0.1810.01

0.2110.03

0.3610.03

17.4 - 74.8

0.0710.01

0.1 110.01

0.1510.01

0.2110.04

Ochogona caroli (Rothenbiihler, 1900)

Mass specific oxygen consumption rates of O. caroli (Table 3) indicate higher values for
juveniles than for adults. Depending on temperature b-values range between 0.60 (6°C), 0.66
( 1 5°C), 0.8 1 ( 1 0°C) and 0.83 (20°C).

Table 3. — Mean ( x±S.E.) rates of oxygen consumption per unit mass (pi mg-i h-l) over the size range of Ochogona
caroli at 5°C, 10°C, 15°C and 20°C

Size range (mg)

6°C

10°C

1 5°C

n 20°C

1.2 - 3.2

0.1610.01

0.2410.03

0.3010.06

1 1 0.3210.02

5.1 - 10.3

0.0910.01

0.1710.02

0.1710.01

8 0.2710.04

454

ERWIN MEYER. PETER MARSONER & ELISABEPH FISCHER

Haasea fonticulorum (Verhoeff, 1910)

The rates of oxygen consumption per unit mass over the size range of H. fonticulorum
(Table 4) indicate unexpectedly high values for the adults. Even the calculated b-values ranging
between 0.94 (6°C), 1.41 (15° and 20°C) and 1.67 (10°C) do not show the expected relationship
between oxgen consumption and size of the individuals. However, the evident activity peak of
adult H. fonticulorum between September and November known from pitfall trapping (Meyer,
1979) may be responsible for the extraordinary respiration rates as the experiments were carried
out between 20th Sept, and 12th Dec.

Table 4. — Mean (x+S.E.) rates of oxygen consumption per unit mass (pi mg-i h-i) over the size range of Haasea
fonticulorum at 5°C. I0°C. 15°C and 20°C.

Size range (mg)

6°C

10°C

15°C

20°C

1.9 - 4.6

0.09±0.02

0. 1 7±0.05

0. 19±0.02

0.28±0.05

5.4 - 11.8

0.10±0.01

0.25±0.0 1

0.30±0.02

0.45±0.03

The relationship between respiratory rate and temperature

This relationship was investigated using specimens that had been reared at quasi field
temperatures throughout the year and measured at corresponding experimental temperatures of
5°C, 10°C, 15°C and 20°C. The results given in Tables 1-4 and Figures 2-5 indicate that oxygen
consumption rates of the four species investigated do not gradually increase with increasing
temperature. In E. nanus the steepest increase in oxygen comsumption takes place between 10°C
and 15°C which can be summarised as a Qio value of 3.08 (mean over all size classes). Between
15°C and 20°C the respiratory metabolism is nearly balanced (Qio = 1.21). In the three subalpine
species (L. saltuvagus, O. caroli and H. fonticulorum) the independence of their respiratory
metabolism of the temperature lies between 10°C and 15°C. In all cases the Qio values are not
significantly different from 1 (MANN-WHITNEY test). Between 6°C and 10°C the relationship
between respiratory rate and temperature is very close with Qio values of 7.9 ( L . saltuvagus),
3.2 ( O . caroli ) and 7.2 (H. fonticulorum).

Variation in respiratory rate with season

Measurements of the respiratory metabolism of E. nanus were carried out during the period
between April and November. The experimental temperatures corresponded to the rearing
temperatures and those to the temperature in the litter layer of its habitat. Hence in April and

October the experimental temperature was

|jl per mg per hour

Temperature Degree C

10°C, in May, June and September 15°C, in
July and August 20°C and in November 5°C.
In Figure 1, the respiratory rates at
corresponding experimental temperatures but
different months are compared.

Fig. 1. — Mean (x ± S.E.) respiratory rates of
Enantiulus nanus at different temperatures and
seasons. The letters within the columns indicate
the month in which the respiration experiments
at the given temperature were made. The figures
within the columns indicate the number of
animals used.

Source :

POPULATION METABOLISM OF MILLIPEDES IN CENTRAL ALPS

455

Relating results to the season (months), there is no significant difference in the respiratory
rate of E. nanus at 10°C and 15°C. Only in August at the given temperature of 20°C did the
animals respire at a significantly lower rate than in July. That corresponds with the life cycle of
E. nanus. Adults undergo their annual moulting phase in August and are therefore inactive.

Estimation of population metabolism

PHILLIPSON (1970) recommended a “best estimate” of respiratory metabolism as the mean
energy loss per unit mass per unit time calculated from laboratory measurements on all life stages
of a given species. Its calculation is independent of field population data, fluctuations in field
temperatures and generation time. During this investigation, oxygen consumption of all life
stages of four millipede species has been measured at temperatures of 5°C, 10°C, 15°C and 20°C.
The results are summarised in Tables 1-4. Estimates of population metabolism have been
obtained by multiplying the size- and temperature- specific respiratory rates with the time- and
age- specific biomass data from the field (Figs 2-5). It should be mentioned that the temperature
specific respiration rates were calculated on the basis of the Q!0 values and not on the assumption
that respiratory rate is an exponential function of temperature, as was demonstrated by HASSALL
(1983) for the isopod Philoscia muscorum.

20 D*gr»« C

ml 02 per square meter per day

1000

Litter/soil temperature

Fig. 2. — Top left: The relationship between oxygen consumption and temperature for the four age groups (AG) of E.
nanus . The size range of the different groups are given in Table 1. Bottom left: Mean monthly temperature in the
litter layer of the oak wood. Bottom right: Mean monthly biomass of juveniles (age group I. stage II- VIII) and
adults (age group II - IV, stage IX-XIV) of E. nanus in the oak wood (taken from Kofler & Meyer 1992). Top
right: Oxygen consumption by the whole population over the period March-December.

456

ERWIN MEYER, PETER MARSONER & ELISABETH FISCHER

Enantiulus nanus (Latzel, 1884) (Fig. 2)

The sum of estimates for each age group gives a total oxygen consumption of 1488 ml O2 m-i
year-i (= 30 KJ m-i) for the population of E. nanus in the oak wood. The graphical presentation
of the data provides an impression of the relationships between the field temperatures, the
dynamics of the biomass and the population metabolism during the vegetation period. Highest
total respiration rates are obtained between April and July. In spite of declining biomass due to
the disappearance of adults (but favoured by the rising temperature) the population metabolism
stays on the same level in the first half of the year. The second biomass peak during October
brings the population metabolism again nearly to the same level. The portion attributable to
juvenile metabolism is 59-88% in the total population and it is highest in July and August when
the new generation is appearing. Adults show their highest respiration rates in May and June
during the egg-laying period.

pi per mg per hour

16 20 D*gr*« C

ml 02 per square meter per day

Degree C mg fresh weight per square meter

Fig. 3. — Top left: The relationship between oxygen consumption and temperature for juveniles and adults of L.
saltuvagus. The size range of the two groups are given in Table 2. Bottom left: Mean monthly temperature in the
litter layer ot the Alnetum viridis at 2000 m a.s.l. Bottom right: Mean monthly biomass of juveniles (stage III-
VIII) and adults (stage IX-XI) of L. saltuvagus in the Alnetum viridis (taken from Meyer, 1985). Top right:
Oxygen consumption by the whole population over the period May-October.

Leptoiulus saltuvagus (Verhoeff, 1898) (Fig. 3)

For the population of L. saltuvagus living in alder litter at the timberline, an annual
respiratory metabolism of 528 ml O2 m-i (= 10.6 kJ m-i) was calculated. According to the
fluctuations in biomass, the oxygen consumption is highest in late summer and autumn. The

Source : MNHN, Paris

POPULATION METABOLISM OF MILLIPEDES IN CENTRAL ALPS

457

period with highest litter temperatures (July) does not become apparent in the population
metabolism because at that time the population structure undergoes the yearly change.
Overwintered adults disappear, overwintered juveniles are probably moulting and the “this-
year's” generation has not yet hatched. The portion attributable to juvenile metabolism is 53% in
the total population on average with highest values in spring and autumn.

Ochogona caroli (Rothenbuhler, 1900) and Haasea fonticulorum (Verhoeff, 1910)

(Figs 4 and 5)

Total population metabolism of these two Chordeumatida is low. (O. caroli: 99 ml O2 m-i
year- 1, (= 1.9 KJ m-i); H. fonticulorum: 105 ml O, m-i year-i (= 2.1 KJ m-ij. In
correspondence with the biomass, highest respiratory rates are obtained in September and
October when juveniles and adults occur in large numbers. Adults are short-lived and die in the
early spring. 78% (79%) of the total population respiration per year is attributable to the
juveniles.

/jl per mg per hour

16 20 Degree C

ml 02 per square meter per day

Population respiration

juveniles

-12.6. -1.7. -2a7. -13.8. -2.9. -14.9. -8.10.

Degree C

mg tresh weight per square meter

— Top left: The relationship between oxygen consumption and temperature for juveniles and adults of O. caroli.
The size range of the two groups are given in Table 3. Bottom left: Mean monthly temperature in the litter layer
of the A In e turn viridis at 2000 m a.s.l. Bottom right: Mean monthly biomass of juveniles (stage II-VI1I) and
adults (stage IX) of 0. caroli in th cAlnetum viridis (taken from Meyer, 1979). Top right: Oxygen consumption
by the whole population over the period May-October.

458

ERWIN MEYER. PETER MARSONER & ELISABETH FISCHER

fil per mo per hour

ml 02 per square meter per day

-12.8. - 17. -23.7. -13.8. - 2.9. -14.9. - 8.10.

-12.8. - 1.7. -23.7. -13.8. - 2.9. -14.9. - 8.10.

Litter/soil temperature

Fig. 5. — Top left: The relationship between oxygen consumption and temperature for juveniles and adults of H .
fonticulorum. The size ranges of the two groups are given in Table 4. Bottom left: Mean monthly temperature in
the litter layer of the Alnetum viridis at 2000 m a.s.l. Bottom right: Mean monthly biomass of juveniles (stage
II- VIII) and adults (stage IX) of H. fonticulorum in the Alnetum viridis (taken from MEYER 1979). Top right:
Oxygen consumption by the whole population over the period May-October.

DISCUSSION

The results presented above show that in all species there is a relationship between the
respiratory rate and size of individual with b-values ranging from 0.2 to 0.92 depending on
species and experimental temperature. Only in H. fonticulorum does the exponent exceed the
known range (0.1 - 1.2, as summarised by PENTEADO et al., 1991) reaching values of 1.41-
1.67. The measurements coincided with the usual high autumnal activity of this species in its
habitat.

The effects of temperature on millipede metabolism have been studied in several species by
Gromysz-Kalkowska & Stojalowska (1966) and Grom ysz-Kalkowska (1970, 1973,
1974). As in many other invertebrates (WlESER, 1973) there is no continuous increase in oxygen
consumption with increasing temperature. “Balanced” respiratory metabolism was found within
different temperature ranges depending on the species. Such temperature-insensitive phases
frequently occur around the mean temperature to be expected in the environment where the
species is active. Such a phenomenon could also be demonstrated during the present study. The
oak wood species E. nanus shows its “temperature-insensitive” phase between 15°C and 20°C
with a Qio of 1.21. This reaction is not fully explained by the temperature conditions in its

Source : MNHN, Paris

POPULATION METABOLISM OF MILLIPEDES IN CENTRAL ALPS

459

habitat, as the daily maxima certainly exceed 15°C for approximately 60 days between June and
August, but in no month does the mean temperature reach 15°C (Fig. 3). In the lower
temperature range between 10°C and 15°C the Qio is 3.08. This great sensitivity to temperature
changes allows the animal to speed up its metabolism as much as possible following hibernation
or coldness. Consistently the three subalpine species (L. saltuvagus , O. caroli and H.
fonticulorum) have their greatest sensitivity to temperature changes between 6°C and 10°C with
Qio values of 7.9, 3.2 and 7.2. In their habitat at the timberline wet weather is often
accompanied by coldness and snowfall even in the summer. Large and rapid temperature
changes often occur during the vegetation period. Between June and September the oscillations
of the daily mean temperatures in the litter layer of Alnus viridis are mostly between 5°C and
10°C (MEYER, 1990). A high sensitivity in this temperature range must be ecologically
significant and allows the millipedes to exploit warmer periods efficiently in the overall short
growth period. The temperature-insensitive phase of these species occurs between 10°C and
15°C with Qio values of 1.1, 1.5 and 1.2.

Figures 2-5 allow the comparison of the relative influences of the factors temperature, age
structure and biomass affecting the final estimates of the population metabolism. In all species
the seasonal changes in oxygen consumption of the population reflects changes in population
structure and biomass much more than changes in temperature. Similar observations by
HASSALL (1983) during investigations into the population metabolism of the isopod Philoscia
muscorum led to the suggestion that the accuracy with which the population structure and
biomass can be assessed is likely to be the most important single factor in determining the
metabolism for field populations.

REFERENCES

Dwarakanath, S. K., 1971. — The influence of body size and temperature upon the oxygen consumption in the
millipede Spirostreptus astenes (Pocock). Comp. Biochem. Physiol., 38A : 351-358.

Fischer, E., 1985. — Sauerstoffverbrauch von 3 subalpinen Diplopodenarten in Abhangigkeit von Temperatur und
Korpergewicht. Magisterarbeit, Univ. Innsbruck, 52 pp.

Gromysz-Kalkowska, K., 1970. — The influence of body weight, external temperature, season of the year, and fasting
on respiratory metabolism in Polydesmus complanatus L. (Diplopoda). Folia Biol. ( Krakow ) , 18 : 311-326.

Gromysz-Kalkowska, K., 1973. — Some exogenous and endogenous effects on rate of respiration in Strongylosoma
pallipes (Olivier) (Diplopoda) and behaviour in temperature gradient. Folia Biol. ( Krakow ) , 21 : 251-269.

Gromysz-Kalkowska, K.. 1974. — The effect of some exogenous factors and body weight on oxygen consumption in
Glomeris connexa C. L. Koch (Diplopoda). Folia Biol. ( Krakow ) , 22 : 37-49.

Gromysz-Kalkowska, K. & Stojalowska, W., 1966. — Respiratory metabolism in Onhomorpha gracilis C. L. Koch
(Diplopoda) as a function of temperature and body size. Folia Biol. ( Krakow ) . 14 : 379-389.

Hassall, M., 1983. — Population metabolism of the terrestrial isopod Philoscia muscorum in a dune grassland
ecosystem. Oikos . 41 : 17-26.

KOFLER, E. & MEYER, E., 1992. — Lebenszyklus, Abundanz und Produktion von Enamiulus nanus (Latzel) in einem
inneralpinen Eichenmischwald (Tirol, Osterreich). Ber. nai.- med. Verein Innsbruck, Suppl. 10 : 153-166.

Marsoner, P., 1992. — Atmungsstoffwechsel von Enamiulus nanus (Diplopoda, Julida). Magisterarbeit, Univ.
Innsbruck, 54 pp.

Meyer, E., 1979. — Life-cycles and ecology of High Alpine Nematophora. In : M. Camatini, Myriapod Biology.
London, Academic Press : 295-306.

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.

MEYER, E., 1990. — Altitude-related changes of life histories of Chordeumatida in the Central Alps (Tyrol, Austria). In :
A. MlNELLl, Proc. 7th Ini. Congr. Myriapodology. Leiden, Brill : 311-322.

Penteado, C. H. S., Hebling-Beraldo M. J. A. & Mendes E. G., 1991. — Oxygen consumption related to size and sex
in the tropical millipede Pseudonannolene tricolor (Diplopoda, Spirostreptida). Comp. Biochem. Physiol.. 98 A :
265-269.

Penteado, C. H. S. & Hebling-Beraldo, M. J. A., 1991. — Respiratory responses in a Brazilian millipede,
Pseudonannolene tricolor, to declining oxygen pressure. Physiol. Zool., 64 : 232-241.

460

ERWIN MEYER. PETER MARSONER & ELISABETH FISCHER

Phillipson, J., 1970. — The “best estimate" of respiratory metabolism: its applicability to field situations. Pol. Arch.
Hydrobiol. , 17 : 31-41.

Wieser. W.. 1973. — Temperature relations of ectotherms: a speculative review. In : W. Wieser, Effects of temperature
on ectothermic organisms. Berlin-New York. Springer-Vcrlag : 1-23.

Wooten. R. C. & Crawford. C. S.. 1974. — Respiratory metabolism of the desert millipede Orthoporus ornatus
(Girard) (Diplopoda). Oecologia (Berl.) . 17 : 179-186.

Source : MNHN, Paris

Variation de la teneur en eau en fonction de la taille
corporelle dans une population du diplopode
Polyzonium germanicum

Guy VANNIER & Jean-Franqois DAVID

CNRS, Laboratoire d'Ecologie Generate, Museum National d'Histoire Naturelle
4 avenue du Petit Chateau, F-91800 Brunoy, France

RESUME

Des echantillons saisonniers de Polyzonium germanicum, comprenant des individus de di-ff6rente taille, ont ete
prelev£s en foret d'Orleans. La teneur en eau, exprimee en pourcentage de la masse s£che, a ete mesuree chez chaque
individu. Quand la masse s&che est prise comme critere de taille, la variation de la teneur en eau en fonction de la taille
peut etre analysec a I'aide dun module de regression. La relation masse fraiche-masse seche est lineaire (Y = a + bX) et le
signe de l'ordonnee a l’origine (a) permet de prevoir le sens de variation de la teneur en eau en fonction de la masse s&che
— qui suit une relation hyperbolique si a * 0. Dans la population etudiee, a est positif, ce qui implique que la teneur en eau
decroit quand la masse seche augmente. Les influences de la saison et du sexe sont analysees de la meme fa9on. Quand le
nombre d'anneaux du corps est pris comme critere de taille, cela permet de preciser le stade de d£veloppement des
individus. Mais l'utilisation des modeles de regresssion se complique et il faut recourir aux comparaisons de moyennes
pour etudier les variations de la teneur en eau en fonction de la taille, de la saison et du sexe. Les deux methodes montrent
que la baisse de la teneur en eau quand la taille corporelle augmente est plus marquee chez les femelles que chez les males.
De plus, quels que soient la taille et le sexe des individus, leur teneur en eau varic saisonnierement, avec des valeurs plus
elev6es lete que l'hiver. Ces resultats sont discutes a la lumiere des connaissances sur lecologie et la biologic de
l'espece.

ABSTRACT

Water content in relation to body size in a population of the millipede Polyzonium
germanicum.

Polyzonium germanicum individuals of nearly all sizes were collected seasonnaly in the forest of Orleans (France).
Water content, expressed as a percentage of dry mass, was determined for each specimen. When body size is equated with
dry mass, the changes in water content with body size can be analysed using regression models. The relationship
between fresh mass and dry mass is linear (Y = a + bX), and the sign of the intercept (a) makes it possible to predict the
direction of variation in water content vs. dry mass — which is a hyperbolic relation if a * 0. In the population studied a
is positive, i.e. water content decreases as dry mass increases. The influence of season and sex is studied in the same way.
Body size can also be expressed as a number of rings, which makes it possible to distinguish the stages of development.
However, regression analyses are not so straightforward as above, because of curvilinear relationships between mass
components and number of body rings. In this case, classic comparisons of means are used to study the changes in water
content with body size, sex and season. Both methods show that the decrease in water content as body size increases is
more pronounced in females than in males. A seasonal effect on water content is apparent, animals of all stages being
more hydrated in warm and dry season than in cold and moist season. These results are discussed taking biological and
ecological features of the species into consideration.

Vannier. G. & David, J.-F., 1996. — Variation de la teneur en eau en fonction de la taille corporelle dans une
population du diplopode Polyzonium germanicum. In: GEOFFROY, J.-J., MAURlfcS, J.-P. & NGUYEN DUY - JACQUEMIN. M.,
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 461-471. Paris ISBN : 2-85653-502-X.

462

GUY VANNIER & JEAN -FRAN^OI S DAVID

INTRODUCTION

L'economie hydrique des Diplopodes a surtout ete etudiee sous Tangle de la resistance a la
dessiccation et des mesures de transpiration, dans le but d'etablir des relations entre la
distribution geographique des especes et les conditions climatiques de leur habitat (PERTTUNEN,
1953 ; BARLOW, 1957 ; Haacker. 1968 ; O'Neill, 1969 ; Crawford, 1972 ; Stewart &
Woodring, 1973 ; Baker, 1980 ; Wegensteiner, 1982 ; Meyer & Eisenbeis, 1985).
D’autres auteurs ont aborde le sujet a travers l'etude de l'ovogenese (CRAWFORD & WARBURG,
1982) ou l'analyse du milieu interieur en evaluant les reponses osmotiques des individus selon
leur degre de deshydratation et de rehydratation (e.g. WOODRING, 1974 ; RIDDLE et al. , 1976 ;
Riddle, 1985).

Nous avons aborde l'etude de la teneur en eau corporelle chez le diplopode Polyzonium
germanicum Brandt, 1831, dont nous connaissons le cycle biologique (David & COURET,
1985) et les preferences ecologiques (DAVID, 1990). Notre contribution differe des precedentes
en ce qu'elle utilise des relations statistiques nouvelles qui mettent en evidence la variation de
parametres caracterisant l'etat hydrique des individus a l'interieur d'un echantillon representatif
de tous les stades de developpement au sein d’une population. C'est ainsi que la relation du
premier degre qui lie la masse fraiche a la masse seche permet de connaitre le sens de variation de
l'hyperbole qui lie la teneur en eau corporelle a la masse seche consideree comme critere de taille
(VANNIER, 1975). Ces variations ont ete etudiees en fonction des saisons et en fonction du sexe.
D'autres relations ont ete formulees en prenant plus classiquement comme critere de taille le
nombre d'anneaux pediferes ; elles ont egalement permis de montrer un effet saisonnier des
variations hydriques corporelles.

MATERIEL ET METHODES

Quelques dizaines d'individus de P. germanicum ont et6 r£colt£s environ tous les deux mois au cours d'une periode
allant de mars 1991 k octobre 1992. Le site de pr&kvements est une litiere de pins sylvestres (Pinus silvestris ) dans le
massif d'Ingrannes de la Foret d'Orl&ms (120 km au sud de Paris) ; le site a 6l6 decrit en detail par David (1990) (site n°
10).

L’echantillonnage des animaux a ete effectue par une recherche manuelle directe k l'aide de pinces brucelles de
maniere k capturer l'6ventail le plus complet possible des classes de taille presentes dans la litidre. Les individus ainsi
captures ont ete ramenes sans tarder au laboratoire. Ils ont ete peses k l'aide d’une microelectrobalance (precision 1
microgramme) pour mesurer leur masse fraiche ; places ensuite dans une 6tuve (60°C) pendant 48 heures, puis transfers
dans un dessiccateur (0% H.R.) pour une duree de 15 jours et obtenir leur masse seche.

Le nombre d'anneaux pediferes et le sexe ont ete determines sur les carcasses seches apres un court passage dans
une solution de potasse k 10%.

Au cours de notre 6tude, nous avons traite un ensemble de 279 individus segmentes et 5 oeufs.

RESULTATS

Variation de la teneur en eau corporelle par rapport a la masse seche

1.- Rappel du principe de l'analyse mathematique

La masse fraiche (Y) est lie a la masse seche (X) par une relation lineaire :

Y = a + bX (1)

La pente (b) de la droite est toujours positive.

A l'aide de cette equation, on demontre que la relation entre la teneur en eau (W exprimee
par rapport a la masse seche) et la masse seche (X) est une fonction hyperbolique de la forme :

W = a/X + (b-1) (2)

Trois cas peuvent se presenter :

- lorsque la constante (a) de l'equation (1) est positive, la derivee de l'equation (2),

W' = -a/X2, est negative ;

- lorsque la constante (a) de l'equation (1) est negative, la derivee de l'hyperbole est
positive ;

Source :

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463

lorsque la constante (a) de l'equation (1) est nulle, l'hyperbole se confond avec une droite
parallele a l'axe des masses seches et tous les individus ont alors la meme teneur en eau ■

W = b- 1.

Ainsi done le signe de la constante (a) de la relation lineaire entre la masse fraiche et la
masse seche indique le sens de variation de la teneur en eau corporelle dans la population sans
meme avoir besoin de la calculer (VANNIER, 1975).

2. - Etude de l'ensemble des individus echantillonnes

La relation qui lie les masses fraiches (Y) aux masses seches (X) des 279 animaux s'ecrit ■

Y = 0,321 + 1,979 X avec R2 = 0,97

La constante de l'equation est positive a l'interieur de limites de confiance comprises entre
+ 0,454 et + 0,188, au seuil de 95% (Fig. la).

Dans ce grand echantillon qui comprend la plupart des stades de developpement de
P. germanicum, la moyenne des masses fraiches (± erreur standard) est de 5,039 mg ± 0,264,
celle des masses seches est de 2,384 mg ± 0,131.

La relation qui lie la teneur en eau (W exprimee en pourcentage) a la masse seche (X
exprimee en milligrammes) est done une hyperbole convexe (Fig. lb) :

W% = 32,1/X + 97,9

avec une derivee negative : W' = -32,1/X2 et une limite theorique de la teneur en eau
corporelle chez les plus grands individus egale a 97,9%. Cet exemple revelant que les jeunes
individus sont plus riches en eau que les individus adultes est le plus frequemment rencontre
dans la nature. Les deux autres cas de figure indiquant l'inverse ou montrant des valeurs stables
de la teneur en eau quel que soit la masse seche sont plus exceptionnels. La teneur en eau
moyenne des 279 individus est de 121,06% ± 2,05.

3. - Etude des variations saisonnieres sans distinction des sexes

La dispersion des points observee sur la Figure, lb nous a incites a rechercher son origine
dans un effet saisonnier. Nous avons analyse nos donnees en les regroupant en trois categories
correspondant aux animaux recoltes en hiver du 9 decembre au 19 mars (n = 126), ceux recoltes
a la fin du printemps et au debut de l'ete du 21 mai au 9 juillet (n = 104) et ceux recoltes au debut
de l'automne du 30 septembre au 9 octobre (n = 49).

Les relations lineaires entre masse fraiche (Y) et masse seche (X) correspondant a ces trois
situations s'ecrivent comme suit :

- Hiver : Yh = 0,183 + 1,867 X avec R2 = 0,995

- Ete : Ye = 0, 173 + 2,280 X avec R2 = 0,987

- Automne : Ya = 0,760 + 1,843 X avec R2 = 0,975

Les constantes (a) de ces trois equations lineaires sont toutes significativement positives au
seuil de 95% :

- Hiver : + 0,264 < a < + 0, 102

- Ete : + 0,326 < a < + 0,020

- Automne : + 1,037 < a < + 0,483

De cette premiere analyse, on peut deja relever que la pente de droite de l'ete est
significativement plus elevee que celle caracterisant les deux autres saisons (P < 0,001) ; que la
teneur en eau des plus jeunes individus est dans les trois cas de saison superieure a celle des plus
ages ; celle-ci tend vers des limites significativement differentes : 128% en ete, 87 et 84%
respectivement en hiver et en automne.

464

GUY VANNIER & JEAN -FRANCOIS DAVID

Fig. I. — Relations hydriques chez les 279 indi vidus £chantillonnes de mars 1991 h octobre 1992. a : Relation lineaire
entre la masse fraiche (Y) et la masse seche (X). b : Relation hyperbolique entre la teneur en eau corporelle (W, en
pourcentage de la masse s£che) et la masse seche (X).

Fig. I. — Water relationships for the whole sample (n = 279). a: Linear relation between fresh mass (Y) and dry mass
(X). b: Hyperbolic relation between relative water content (W expressed as a percentage of dry mass) and dry
mass (X).

Les hyperboles correspondant aux trois saisons sont representees sur la Figure 2. Leur
etagement demontre bien que les teneurs en eau des animaux d'ete sont systematiquement
superieures a celles des deux autres saisons. On remarque aussi qu'en automne, saison
intermediaire, les individus de petite taille possedent des teneurs en eau elevees proches de cedes
mesurees en ete ; au contraire les individus de plus grande taille tendent vers des valeurs faibles,
proches de cedes rencontrees en hiver.

Source : MNHN. Paris

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Fig. 2. — Variations saisonnieres tie la relation hyperbolique entre la teneur en eau corporelle (W, en pourcentage de la
masse s£che) el la masse seche (X) chez les indi vidus recoltes au cours de 1'ete (E : carres blancs), I’automne (A ;
carrcs pointes) et 1'hiver (H ; carres noirs).

FlG. 2. — Seasonal changes in the hyperbolic relation between water content ( W expressed as a percentage of dry mass)
and dry mass (X) (E = summer; white squares) (A = autumn; dotted squares) (H = winter; black squares).

4.- Etude des variations saisonnieres selon les sexes

a) Les femelles sont analysees avec les jeunes individus indifferencies en fonction des trois
periodes saisonnieres : hiver (n = 78), ete (n = 64), automne (n = 31). Les equations lineaires
entre masses fraiches (Y) et masses seches (X) correspondant aux trois situations climatiques
s'enoncent comme suit :

- Hiver : Yh = 0,137 + 1,867 X avec R2 = 0,996

- Ete : Ye = 0,203 + 2,276 X avec R2 = 0.987

- Automne : Ya = 0,980 + 1,81 1 X avec R2 = 0,974

Les constantes (a) de ces trois equations lineaires peuvent etre considerees comme
significativement positives au seuil de 95% en hiver et a l'automne :

- Hiver : + 0,244 < a < + 0,030

- Ete : + 0,41 1 < a < - 0.005

- Automne : + 1,395 < a < + 0,565

b) De la rneme maniere, les males et les jeunes individus indifferencies ont ete rassembles
dans chacune des trois periodes du cycle annuel que nous avons analysees : hiver (n = 69). ete
(n = 45), automne (n = 22). Les equations lineaires entre masse fraiche (Y) et masse seche (X)
correspondant aux trois saisons s'ecrivent comme suit :

- Hiver : YH = 0,048 + 1,968 X avec R2 = 0,990

- Ete : Ye = 0, 124 + 2,286 X avec R2 = 0,985

- Automne : Ya = 0,288 + 1,960 X avec R2 = 0,987

Les constantes (a) des equations lineaires ci-dessus sont comprises dans un intervalle de
valeurs significativement positives a l'automne, mais non significativement differentes de zero
1’hiver et 1'ete :

- Hiver : + 0,134 < a < - 0,038

- Ete : + 0,324 < a < - 0,076

- Automne : + 0,490 < a < + 0,086

La variation de la teneur en eau en fonction de la taille est done soit decroissante (a > 0),
soit a peu pres constante (a # 0), mais jamais croissante chez P. germanicum. La diminution de

466

GUY VANNIER & JEAN-FRANCOIS DAVID

la teneur en eau quand la taille augmente semble aussi plus significative chez les femelles que
chez les males.

Les equations lineaires montrent en outre que les limites minimales theoriques de la teneur
en eau sont variables selon les saisons. En hiver, cette limite est atteinte chez les femelles vers
86.7% et chez les males vers 96.8%. En ete, ellc est atteinte chez les femelles et les males vers
des valeurs plus elevees et tres proches l'une de l'autre, respectivement 127,6% et 128,6%. En
automne, les limites ne sont pas significativement differentes de celles calculees en hiver ; elles
sont de 81,1% chez les femelles et 96% chez les males.

II ressort nettement de cette analyse que les femelles comme les males sont plus riches en
eau pendant la periode estivale.

Variation de la teneur en eau par rapport au nombre d'anneaux pediferes

1.- Comparaison des valeurs moyennes de la teneur en eau en fonction du sexe et des saisons

Le Tableau 1 repartit l'ensemble de l'echantillon de 279 individus en classes de nombre
d'anneaux pediferes, correspondant, pour chaque sexe, a des etapes du developpement post-
embryonnaire, avec une reference concernant quelques oeufs pondus au debut de l'ete. Les
teneurs en eau moyennes de six classes de tailles biologiquement significatives ont ete calculees
pour comparer deux saisons opposees.

Tableau 1. — Valeurs moyennes de la teneur en eau corporelle (en % de la masse seche) en fonction du sexe et des saisons
au cours du cycle vital de P. germanicum.

Table I. — Average water content of P. germanicum (expressed as a % of dry mass ± standard error), in relation to number
of rings, sex and season (Hiver = winter; Ete = summer).

ANNEAUX SEXES PHASES STADE S TENEURS EN EAU

PEDIFERES (% ± E.S.)

HIVER

ETE

—

Oeufs

—

205 ±5

Sail

—

Immatures

Ill

103 ± 2

117 ± 15

13 & 15

Males

Immatures

128 ± 6

Femelles

Immatures

—

158 ± 17

17 a 25

Males

Adultes

V a VII

102 ±3

149 ±9

Femelles

Immatures

V a VI

108 ±6

120 ±8

28 * 34

Males

Adultes

VII a IX

98 ±2

159 + 30

Femelles

Adultes

VII a IX

91 ±2

172 ± 16

35 h 40

Males

Adultes

VIII ^ XI

103 ±4

130 ± 8

Femelles

Adultes

VIII a XI

93 ±2

141 ± 4

41 a 54

Males

Adultes

IX a XIII

103 ± 12

136 ±6

Femelles

Adultes

IX a XIII

81 ± 1

131 ±9

a) En hiver, le pourcentage d'eau des males est a peu pres constant quel que soit le stade de
developpement. 11 n'est pas significativement different de celui des immatures du stade III (8-1 1
anneaux). Chez les femelles, la teneur en eau decroit significativement quand la taille augmente et
de maniere accentuee a partir de la maturite (28-34 anneaux) pendant la formation des oocytes en
periode hivemale. On remarquera que les femelles immatures (17-25 anneaux) possedent une
teneur en eau moyenne qui ne differe pas significativement de celle des formes immatures du
stade III (8-11 anneaux).

Source :

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467

b) En ete, les teneurs en eau moyennes sont toujours plus elevees qu'en hiver, meme si les
tests de comparaison de STUDENT et de MANN-WHITNEY ne sont pas tous probants. On notera
que les erreurs standards dans les deux sexes sont plus elevees en ete qu'en hiver (dans huit cas
sur neuf) ; cette plus grande dispersion des valeurs de la teneur en eau corporelle pendant la
saison chaude peut avoir une signification biologique.

Fig. 3. — Relations entre la masse et le nombre d’anneaux pediferes chez les femelles et les jeunes ( P . germanicum)
recoltes au cours de l’6t6 (E ; carres blancs) et de 1’hiver (H ; carres noirs). a : Variations saisonnieres de la relation
du second degre entre la masse s£che (X) et le nombre d'anneaux pediferes (Z). b : Variations saisonnieres de la
relation du second degre entre la masse d'eau corporelle (M) et le nombre d'anneaux pediferes (Z).

FIG. 3. — Relationships between mass components and number of podous rings in females and juveniles (P.
germanicum) collected in two seasons (E = summer; white squares ) (H = winter; black squares), a: Seasonal
changes in the quadratic regression of dry mass (X) versus the number of rings (Z). b: Seasonal changes in the
quadratic regression of water mass (M) versus the number of rings (Z).

468

GUY VANNIER & JEAN-FRANQOIS DAVID

Fig. 4. — Relations entre masse et nombre d’anneaux pediferes chez les males et les jeunes {P. germanicum ) r6colt6s au
cours de 1 ete (E ; carres blancs) et de I'hiver (H ; carres noirs). a : Variations saisonnieres de la relation du second
degre entre la masse seche (X) et le nombre d'anneaux pediferes (Z). b : Variations saisonnieres de la relation du
second degre entre la masse d’eau corporelle (M) et le nombre d'anneaux pediferes (Z).

FlG. 4. — Relationships between weight components and number of podous rings in males and juveniles (P.
germanicum) collected in two seasons (E = summer; white squares) (H = winter; black squares), a: Seasonal
changes in the quadratic regression of dry mass (X) versus the number of rings (Z). b: Seasonal changes in the
quadratic regression of water mass (M) versus the number of rings (Z).

2- Etude complementaire des variations saisonnieres en fonction du sexe

Du point de vue dimensionnel, le nombre d'anneaux pediferes a l'avantage d'etre
apprehende independamment des mesures ponderales et permet done de tester si les variations
saisonnieres de la teneur en eau sont imputables soit aux changements de masse d'eau, soit aux
variations de masse seche.

Source : MNHN, Paris

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Chez les formes indifferenciees correspondant au stade III du developpement, la masse
seche est la meme dans les echantillons d'hiver et d'ete (0,21 mg) ; cependant la masse d'eau est
significativement plus elevee en ete qu'en hiver (respectivement 0,36 mg et 0,21 mg ; P < 0,01).

Chez les femelles et chez les males on compare les equations de regression soit de la masse
seche (X), soit de la masse d'eau (M) en fonction du nombre d'anneaux pediferes (Z) entre
l’hiver et lete. Le fait nouveau est que toutes les regressions sont des fonctions du second
degre.

Chez les jeunes et les femelles, les variations de la masse seche (X) en fonction du nombre
d'anneaux pediferes (Z) correspondent aux trinomes suivants (Fig. 3a) :

- Hiver : Yu = 2,002 - 0.286 Z + 0,01 1 Z2 avec R2 = 0,950

- Ete : Ye= 3,622 - 0,399 Z + 0,01 1 Z2 avec R2 = 0,806

A nombre d'anneaux pediferes egal, la masse seche tend a etre plus faible en ete qu’en
hiver, principalement chez les femelles adultes a partir du stade VII porteur de 28 anneaux et
plus.

Devolution de la masse d'eau (M) chez les femelles par rapport au meme critere de taille
(Z) suit le meme type d'equation (Fig. 3b) :

- Hiver : Mh = 1,317 - 0,195 Z + 0.008 Z2 avec R2 = 0.944

- Ete : Me = 4.499 - 0,501 Z + 0,015 Z2 avec R2 = 0,844

La masse d'eau corporelle tend a etre plus elevee en ete qu'en hiver. Cependant, un
chevauchement des deux courbes vers le 35eme anneau montre que les femelles de tailles
comprises entre 28 et 34 anneaux (stades VII a IX) ont une masse d'eau plus faible en ete qu’en
hiver ; chez les femelles plus grandes ou plus petites la tendance generate est respectee.

Chez les jeunes et les males, les equations du second degre liant la masse seche (X) au
nombre d'anneaux pediferes (Z) s'ecrivent comme suit (Fig. 4a) :

- Hiver : Xh = 0,479 - 0,058 Z + 0.003 Z2 avec R2 = 0,927

- Ete : Xe = 0,536 - 0,060 Z + 0,003 Z2 avec R2 = 0,887

A nombre d'anneaux pediferes egal, la masse seche est plus faible chez les individus de
l'ete, mais de maniere moins marquee que chez les femelles.

Les variations de la masse d'eau (M) chez les males en fonction du nombre d'anneaux
pediferes (Z) sont aussi representees sur la Figure 4b et les equations s'ecrivent :

- Hiver : Mh = 0,541 - 0.065 Z + 0,003 Z2 avec R2 = 0,965

- Ete : Me = 0,403 - 0,045 Z + 0,003 Z2 avec R2 = 0,844

La masse d'eau corporelle tend a etre plus elevee en ete qu'en hiver, mais de fa^on moins
accentuee que chez les femelles.

Dans les deux sexes, on a constate les memes tendances : a niveau de developpement egal,
les masses seches sont plus elevees en hiver qu'en ete, alors que c'est l'inverse pour les masses
d'eau. Ces deux observations ont pour consequence de reduire la valeur relative de la teneur en
eau en hiver et de l'augmenter en ete. Ce phenomene dependant des stades a moins d'amplitude
chez les males que chez les femelles.

DISCUSSION & CONCLUSION

Notre etude sur P. germanicum reprend des points traites par les auteurs qui ont aborde le
probleme de l'economie hydrique chez les diplopodes. De maniere non equivoque, nous savons
maintenant que les teneurs en eau corporelle varient selon les saisons et sont dependantes de
1'age et du sexe des individus selon des fonctions specifiques (hyperboles, trinomes).
Cependant, il n'est pas tres aise de reconnaTtre l'influence preponderante d'un facteur ou d'un
autre sur les variations de la teneur en eau corporelle dans une population aussi complexe que
celle de P. germanicum dont le cycle biologique se developpe sur plusieurs annees (David &
COURET, 1985).

470

GUY VANNIER & JEAN-FRAN'COIS DAVID

Parmi les facteurs intrinseques qui influent sur les variations de la teneur en eau des
individus. la croissance est determinante. Dans nos releves, nous avons montre que la teneur en
eau tend a diminuer quand la taille augmente. Cette diminution est surtout visible en hiver, quand
la dispersion des mesures hydriques est la plus etroite. En outre, la comparaison entre sexes
montre que cette decroissance concerne principalement les femelles. Deux interpretations peuvent
etre avancees :

1 . La masse seche des femelles a stade egal est plus elevee que celle des males, ce qui peut
se repercuter sur le calcul de la valeur relative de la teneur en eau.

2. Les tissus reproducteurs (coips gras, ovocytes) peuvent peser sur cette difference entre
sexes, car on constate que la teneur en eau diminue surtout chez les femelles adultes
reproductrices (Tableau 1).

BAKER ( 1 980) a egalement mis en evidence une diminution de la teneur en eau en fonction
de la masse seche chez le iulide Ommatoiulus moreleti dont les femelles sont plus riches en eau
que les males, contrairement a P. germanicum.

Les saisons marquent de leur empreinte les variations de la teneur en eau corporelle.
L'opposition ete-hiver, saison chaude-saison froide, est manifeste (Figs. 2-4, Tableau 1).
Plusieurs mecanismes lies a la phenologie du cycle vital de P. germanicum peuvent etre mis en
cause :

- La masse seche d'un individu de stade donne tend a etre plus faible en ete qu'en hiver. A
cela trois explications possibles : a) Le rejet de la mue en periode estivale ; mais dans nos releves
du debut de fete beaucoup d'individus n'avaient pas encore mue. b) Sachant que chez P.
germanicum le stock d’ovocytes se constitue en hiver et que la ponte a lieu a la fin du printemps
(COURET & David, 1985 ; David & Coijret, 1985), le corps des femelles se trouve done
allege au debut de l'ete. c) Chez les deux sexes, les masses de graisses ont aussi leur part
d'influence sur les fluctuations de la masse seche. Nous developperons cet aspect dans un
prochain article.

- La masse d'eau d'un individu a un stade donne tend a etre plus elevee en ete qu'en hiver.
Nous proposons l'interpretation suivante : au debut de l'ete, avant la periode d'exuviation, les
animaux augmenteraient leurs reserves hydriques, comme l'un de nous l'a observe chez quatre
especes de Collemboles (VANNIER. 1981). Nous avons effectivement observe que de nombreux
individus possedant des teneurs en eau elevees etaient proches de l’ecdysis. Les animaux du
stade IV sont particulierement representatifs de ce phenomene de reprise hydrique. On ne les
rencontre qu'au debut de l'ete et le plus souvent, ils viennent de muer. Leur teneur en eau
(Tableau 1) est beaucoup plus elevee (128% chez les males, 158% chez les femelles) que celle
des stades III d'hiver qui les ont precedes (103%).

Des variations saisonnieres de la teneur en eau corporelle ont ete signalees chez plusieurs
especes de diplopodes dans la litterature. Des valeurs maximales ont ete trouvees au printemps et
en ete chez Ommatoiulus moreleti par BAKER (1980), comme chez P. germanicum de la foret
d'Orleans. Selon cet auteur, cela pourrait s'expliquer par des modifications de structure dans la
population, comme la presence de gros individus a faible teneur en eau dans les echantillons
d'hiver. Cette explication ne convient pas pour P. germanicum dans notre etude. Chez les
Spirostreptidae, Archispirostreptus tumuliporus et Orthoporus ornatus, le maximum hydrique
est atteint en hiver pour la premiere espece qui vit en Israel et en ete pour la seconde vivant dans
le Sud des Etats-Unis. Ces observations ont ete rapportees par CRAWFORD et al. (1987) et
coincident avec la saison humide. Selon ces auteurs, l'accroissement du taux hydrique corporel
chez ces especes peuplant des zones arides serait du a l'ingestion de nourriture humide.

L'analyse que nous faisons de ces donnees bibliographiques peut se resumer en trois
points : a) Le maximum hydrique au printemps et en ete coincide avec la periode des mues chez
Ommatoiulus moreleti (BAKER, 1980). b) C'est egalement avec les mues et avant la sortie en
surface de Archispirostreptus tumuliporus que les teneurs en eau les plus fortes sont mesurees
(CRAWFORD et al., 1987). c) De meme, l'augmentation de la masse d'eau dans la cuticule et les

Source . MNHN, Paris

TENEUR EN EAU ETTAILLE CORPORELLE CHEZ UN DIPLOPODE POLYZONIIDA

471

tissus chez Orthoporus ornatus se produit en fin de saison seche, avant la mue, precedant la
periode d'activite en saison humide (Crawford, 1978).

Du point de vue biologique, il semble bien que P. germanicum reagisse de la meme
maniere, avec une augmentation significative de la teneur en eau corporelle dans les mois qui
encadrent la periode de mue. L'economie hydrique des diplopodes serait davantage regie par des
facteurs intrinseques (crises physiologiques) que par des facteurs extrinseques (climat).

REFERENCES

Baker, G. H., 1980. — The water and temperature relationships of Ommatoiulus moreletii (Diplopoda: lulidae) J Zool
(Loncl.), 190 : 97-108.

Barlow^C- A., 1957. — A factorial analysis of distribution in three species of diplopods. Tijdschr. Entomol., 100 :

CoURET, T. & David, J. F., 1985. — Recherche des stades de maturity sexuelle chez le Diplopode Polyzonium
germanicum Brandi, 1831 (Polyzoniida). Rev. Ecol. Biol. Sol, 22 : 247-258.

Crawford, C. S., 1972. — Water relations in a desert millipede Orthoporus ornatus (Girard) (Spirostreptidae). Comp.
Biochem. Physiol., 42(A) : 521-535.

Crawford, C. S., 1978. — Seasonal water balance in Orthoporus- ornatus, a desert millipede. Ecology, 59 : 996-1004.
Crawford, C. S.. Bercovitz, K. & Warburg, M. R., 1987. — Regional environments, life-history patterns, and
habitat use of Spirostrcptid millipedes in arid regions. Zool. J. Linn. Soc.. 89 : 63-88.

Crawford, C. S. & Warburg, M. R.. 1982. — Water balance and apparent oocyte resorption- in desert millipedes. J.
exp. Zool., 222 : 215-226.

David, J.-F., 1990. — Habitat dimensions of Diplopoda in a temperate forest on acid soil. Rev. Ecol. Biol Sol 21
95-112.

David, J. F. & COURET, T., 1985. — Le cycle biologique du Diplopode Polyzonium germanicum Brandt. 1831
(Polyzoniida). Rev. Ecol. Biol. Sol, 22 : 367-380.

HaackER, U., 1968. — Deskriptive, experimentelle und vergleichende Untersuchungen zur Autokologie rhein-
mainischer Diplopoden. Oecologia ( Berl .), 1 : 87-129.

Meyer, E. & Eisenbeis, G., 1985. — Water relations in millipedes from some Alpine habitat types (Central Alps, Tyrol)
(Diplopoda). Bijdr. Dierkd.,55 : 131-142.

O’Neill. R. V., 1969. — Comparative desiccation tolerance in seven species of millipedes. Am. Midi. Nat.. 82 : 182-
187.

Perttunen, V., 1953. — Reactions of diplopods to the relative humidity of the air. Ann. Zool. Soc. Zool. - Bot . Fenn.
Vanamo, 16 : 1-69.

Riddle, W. A., 1985. — Hemolymph osmoregulation in several Myriapods and Arachnids. Comp. Biochem. Physiol.,
80(A) : 313-323.

Riddle, W. A., Crawford, C. S. & Zeitone, A. M., 1976. — Patterns of hemolymph osmoregulation in three desert
arthropods. J. Comp. Physiol., 112(B) : 295-305.

Stewart, T. C. & Woodring, J. P., 1973. — Anatomical and physiological studies of water balance in the millipedes
Pachydesmus crassicutis (Polydesmida) and Orthoporus texicolens (Spirobolida). Comp. Biochem. Physiol.,
44(A) : 735-750.

Vannier, G., 1975. — Les trois cas de figure de la relation teneur en eau corporelle-poids sec chez un insecte Collembole
au cours du cycle annuel. C. R. Acad. Sci. Ser. D, 280 : 117-120.

Vannier, G., 1981. — Desequilibrc de la balance hydrique corporelle chez quatre especes d’insectes Collemboles apres
un jeune de courte durcc. Rev. Ecol. Biol. Sol, 18 : 29-38.

WEGENSTEINER, R.. 1982. — Zusammenhange zwischen dcr okologischen Potenz von Polyzonium germanicum Brandt
(Diplopoda, Colobognatha) und Standortparametern im Vorderen Rotmooz (Lunz, NO). Zool. Jahrb. Syst., 109 :
309-327.

Woodring. J. P., 1974. — Effects of rapid and slow dehydration on the hemolymph osmolarity and Na+-K +
concentration in the millipede Pachydesmus crassicutis. Comp. Biochem. Physiol., 49(A) : 115-119.

Source : MNHN, Paris

The Respiratory Response to Changing
Temperature in Millipedes belonging
to the Genus Glomeris Latreille, 1802

Vladimir SUSTR

Institute of Soil Biology, Academy of Sciences of the Czech Republic, 370 05 Ceske Budejovice, Czech Republic

ABSTRACT

The relationship between respiration rate and temperature of 3 millipede species: Glomeris marginata (Villers, 1789).
Glomeris hexasticha Brandt, 1833 and Glomeris balcanica (Verhoeff, 1906), with different geographical distribution was
measured and analysed. Simple linear relationship was found in G. marginata. A zone of the relative independence of
respiration rate on temperature in the temperature range 15-20°C was observed in G. hexasticha. The zone was less
apparent in G. balcanica. Specific respiration rates were higher in males than in females in all species. Possible
ecological significance of the difference in the respiration-temperature dependence was discussed.

RESUME

Reponse respiratoire aux changements de temperature chez les diplopodes du genre Glomeris
Latreille, 1802.

Les relations entre le m^tabolisme respiratoire et la temperature ont etc mesurees el analysces chez trois espfcces de
diplopodes prSsentant des aires de repartition differentes : Glomeris marginata (Villers, 1789), Glomeris hexasticha
Brandt, 1833 et Glomeris balcanica (Verhoeff. 1906). Une relation lineaire simple a ete mise en evidence chez
G. marginata. Chez G. hexasticha , le metabolisme respiratoire est relativement independant de la temperature, entre
15°C et 20°C, phenomene moins Evident chez G. balcanica . Pour toutes les especes, les taux respiratoires sont plus
elev6s chez les males que chez les femelles. La possibility d’une interpretation ecologique de cette difference dans la
relation respiration-temperature est discutee.

INTRODUCTION

Glomeris balcanica is a dominant species of the soil macrofauna in extreme biotopes on an
altitudinal gradient on Mt. Olympus and in a Quercus coccifera formation in Northern Greece.
The detailed description of its biology and ecology, including respiration activity, was given by
IATROU (1989) and lATROU & STAMOU (1989). Glomeris marginata is a common inhabitant of
litter layers in deciduous forests of western Europe, Glomeris hexasticha is a middle and east
European species. The biology and ecology of G. marginata and G. hexasticha was described by
DUNGER & STEINMETZGER (1981). More information about respiration-temperature
relationships have been published (e.g. PENTEADO & MENDES, 1981; GROMYSZ-KALKOWSKA
& TRACZ, 1983). However, different authors have used different experimental conditions, so

Sustr, V.. 1996. — The respiratory response to changing temperature in millipedes belonging to the genus
Glomeris Latreille, 1802. In: Geoffroy, J.-J.. Mauries, J.-P. & Nguyen Duy - Jacquemin, M.. (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 473-476. Paris ISBN : 2-85653-502-X.

474

VLADIMIR SUSTR

generalizations are difficult. The main aim of this study was to test differences between the
species in specific metabolic rate (SMR) and to assay the SMR - temperature relationship using
almost identical experimental procedure and the same method of measurement.

MATERIAL AND METHODS

The specimens of G. balcanica were collected in the northwest of Greece, some 20 km from Thessaloniki about 400 m
a.s.I. in June 1991 (leg. Tajovsky). and. after transport to our laboratory, kept for 3 weeks at 15°C in darkness. A
mixture of Quercus coccifera and Quercus robur litters was used as food. Specimens of G. marginata were collected from a
Quercus pubescens forest in Ruoms near Montdlimar (south of France) at an altitude 400 m a.s.I. in May 1991 (leg.
Frouz) and kept on Q. robur litter in similar conditions as G. balcanica. Individuals of G. hexasticha were collected in a
mixed oak forest near Netolice in South Bohemia at an altitude from 485 to 570 m a.s.I. in May 1991 (leg. Sustr &
Tajovsky). and kept on Quercus robur litter for 1- 3 weeks in conditions similar to those for G. balcanica. Oxygen
consumption was measured in a Warburg respirometer over a 5-hour period; 30 minutes were allowed for
thermostabilization. The respiration rate of every animal was measured successively at 5, 10, 15. and 20°C during 4 days
of experiment. Animals were reared in the laboratory culture (15°C. dark, with food) for approximately 19 hours between
the measurements. Seventeen individuals of G. hexasticha (7 males and 10 females), 15 specimens of G. balcanica (2
males and 13 females) and 16 individuals of G. marginata (7 males and 9 females), with body mass in the range of 0.030 -
0.219 g, 0.091 - 0.282 g, and 0.073 to 0.399 g, respectively, were used in the experiments. Qio values were calculated
according to the equation Qio = (ki/k2)10/(tl-t2), where ki and k2 are mean respiration rates at temperatures tl and t2
respectively.

RESULTS

Specific metabolic rate (SMR)-temperature curves are shown in Figure 1. anova indicated
a significant effect of temperature on SMR in all three species (F = 36.6, P < 0.01 in G.
marginata, F = 40.4. P < 0.01 in G. hexasticha and F = 49.0, P < 0.01 in G. balcanica). The
shapes of the curves were similar in both sexes of the same species (see Table 1). An almost
linear curve was observed in G. marginata. The increase of SMR with increasing temperature
was larger from 5 to 15°C than from 15 to 20°C in G. balcanica. A small range of relative
temperature independence (RRTI) was observed between 15 and 20°C in G. hexasticha (Fig. 1,
Table 1).

SMR values were higher in males than in females. Differences were significant in G.
hexasticha (F = 5.8. P < 0.02) and in G. balcanica (F = 8.5, P < 0.01). In G. balcanica ,
however, the comparison is disputable because of the limited number of males used in the
experiment.

The SMR of G. marginata was significantly lower than those of G. balcanica and G.
hexasticha at 5 and 15°C (F = 14.6, P < 0.01 and F = 1 1.1, P < 0.01 respectively).

The mean individual body mass was 0.090 g in G. hexasticha, 0.175 g in G. balcanica,
and 0.187 g in G. marginata. Changes in body mass were not significant during the experiment
(F < 0.01, P > 0.9) for any species.

Table 1. — Qio coefficients in three species of glomerid millipedes in the temperature range from 5°C to 20°C.

Species

5-10

5-15

Temperature range
10-15 5-20

10-20

15-20

Glomeris hexasticha

2.0

2.3

2.6

1.9

1.8

1.2

Glomeris hexasticha males

1.9

2.3

2.9

1.8

1.1

Glomeris hexasticha females

2.3

1.9

1.8

1.3

Glomeris balcanica

2.8

3.0

3.1

2.3

2.1

1.3

Glomeris balcanica males

4.4

4.2

4.0

3.0

2.5

1.5

Glomeris balcanica females

2.7

2.1

1.9

1.3

Glomeris marginata

4.3

3.0

2.1

2.5

2.0

1.8

Glomeris marginata males

4.0

3.1

2.3

2.5

2.0

1.8

Glomeris marginata females

5.3

3.2

2.0

2.7

1.9

Source : MNHN. Paris

RESPIRATORY RESPONSE TO CHANGING TEMPERATURE IN MILLIPEDES

475

FlG. I. — Mean SMR - temperature relationships in three species of millipedes. Errors bars: 95% confidence intervals,
filled triangle: Glomeris balcanica, filled square: Glomeris marginata, empty square: Glomeris hexasticha.

DISCUSSION

G. marginata , in agreement with its geographical distribution, would be best adapted to an
oceanic climate with relatively variable weather. The population used in this study lives in a
relatively exposed, warm and dry site. Adaptation to a microclimate with frequent temperature
fluctuations, as well as relatively low locomotion activity appears to have prevented G.
marginata from establishing an RRTI (that does or does not occur in a species). A wide range of
preferred temperatures ( 1 8°C to 26°C) was observed in G. marginata in laboratory by DUNGER
& STEINMETZGER (1981). The site of collection of G. hexasticha has a forest microclimate
without great temperature fluctuations. These conditions may have enabled the establishment of a
clearly expressed RRTI between 15°C and 20°C in the species. Its prefered temperature of 20°C
was reported in laboratory by DUNGER & STEINMETZGER (1981). The lack of RRTI should be
expected in G. balcanica because of high temperature fluctuations in the soil organic layer in Q.
coccifera formation. However, G. balcanica shows greater locomotion activity and irritability
than G. marginata and G. hexasticha. The difference between the standard and active metabolic
rate may be larger and the RRTI consequently more expressed. The combination of the above
mentioned factors contributes to the establishment of the RRTI, which is less apparent in
comparison to G. hexasticha. The placement of the RRTI (between 15°C and 20°C) corresponds

476

VLADIMIR SUSTR

to a temperature optimum of food consumption (IATROU & STAMOU, 1989) and with Qio values
obtained by IATROU (1989).

REFERENCES

Dungkr. W. & Steinmetzger, K.. 1981. — Okologische Untersuchungen an Diplopoden einer Rasen-Wald-Caiena im
Thuringer Kalkgebiet. Zool. Jb . Syst., 108 : 519-553.

GrOMYSZ-Kalkowska, K. & Tracz, H., 1983. — Effect of temperature, food kind and body weight on the oxygen
consumption by Proteroiulus fuscus (Am Stein) (Diplopoda. Blaniulidae). Ann. Warsaw. Agricult. Umv. - SGGW-AR,
For. a. Wood Technol. , 30 : 35-42.

IATROU. G. D., 1989. — Dynamics and activity of the diplopod Glomeris balcanica in the soil subsystem of an
evergreen-sclerophyllous formation in Mt. Hortiatis. Ph. D. Thesis, Aristotelian University of Thessaloniki (In
Greek), 216 pp.

Penteado, C. H. S. & Mendes, E: G., 1981. — Respiratory metabolism and tolerance in a tropical millipede,
Rhinocricus padbergi Verhoeff, 1938. Ill: the response to temperature variations. Rev. Brasil. Biol.. 41 : 499-509.

IATROU, G. D. & Stamou. G. P.. 1989. — Seasonal activity patterns of Glomeris balcanica (Diplopoda, Glomcridae) in
an evergreen-sclerophyllous formation in northern Greece. Rev. Ecol. Biol. Sol. 26 : 491-503.

Source : MNHN, Paris

Submersion Tolerance of some Diplopod Species

Klaus Peter ZULKA

Institute of Zoology, University of Vienna
Althanstr. 14, A- 1090 Vienna, Austria

ABSTRACT

Submersion tolerance of Polydesmus denticulatus was compared to that of other diplopod species. About ten specimens
of five species were placed individually in plastic tubes and flooded in an aquarium with aerated water (temperature
9±1°C). The highest median tolerance times were found in Polydesmus denticulatus , but one specimen of Brachyiulus
bagnalli even reached a higher maximum of 65 days submersion. The other species are considerably less tolerant. In the
four julid species median submersion tolerance times are significantly correlated with the surface/volume-ratio.
Polydesmus denticulatus , however, is an outlier, and shows comparably higher tolerance values. The possible
mechanisms of submersion tolerance are discussed.

RESUME

Tolerance a Timmersion chez quelques especes de diplopodes.

La tolerance & Timmersion de Polydesmus denticulatus a ete comparee a celle d’autres especes de diplopodes. Une
dizaine de specimens de cinq especes ont ete places individuellement dans des tubes de matiere plastique et immerg6s dans
un aquarium contenant de Teau a£ree (temperature 9±1°C). En moyenne, la plus longue duree de tolerance & ce milieu est
presentee par Polydesmus denticulatus. Cependant un invidu de Brachyiulus bagnalli a atteint la duree maximale de 65
jours d’ immersion. Les autres especes sont nettement moins tolerantes. Chez les quatre especes de julides. la duree de
tolerance a Timmersion est correlee de maniere significative avec le rapport surface/volume. Toutefois, Polydesmus
denticulatus reste un cas particular, montrant une tolerance & la submersion plus 61ev6e. Les mecanismes eventuels de
cette capacitc sont discutes.

INTRODUCTION

Diplopods are usually supposed to be weak tolerators if they become submerged by rain or
inundation (BLOWER, 1955; ElSENBEIS & WlCHARD, 1985). However, HOFFMAN (1978)
reports a diplopod from Papuan caves entering voluntarily the water, ADIS (1986) describes long
term submergence in an Amazonian diplopod, and in an investigation of floodplain soil animals
the widespread European diplopod Polydesmus denticulatus was shown to survive up to 75
days in oxygenated cold water (ZULKA, 1991, 1992).

The present experiment should decide if this is a special adaptation of Polydesmus
denticulatus to life in flood plains or if diplopods in general are able to withstand submersion
longer than previously expected.

Zulka, K. P., 1996. — Submersion tolerance of some diplopod species. In: Geoffroy, J.-J., Mauri£s, J.-P. &
Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodoloeica. Mem. Mus. natn. Hist, nat., 169 : 477-481. Paris ISBN : 2-
85653-502-X.

478

KLAUS PETER ZULKA

MATERIAL AND METHODS

Specimens of Polydesmus denticulatus C. L. Koch, Brachyiulus bagnalli (Brolemann), Ophyiulus pilosus
(Newport), Cylindroiulus boleti (C. L. Koch), and Leptoiulus proximus (Nemec) were collected by hand in May 1992 in
an alder forest near Marchegg, Lower Austria. All these species are widespread in Central Europe and inhabit a vast range
of habitats (Blower, 1985), but P. denticulatus is the only one of them living also in temporarily flooded areas and
surviving inundations submerged (ZULKA, 1991). Since the females of Ophyiulus pilosus and Leptoiulus proximus could
not be separated from each other and from the syntopic Julus scandinavius with certainty, only males of these two
species were used in the experiment. About ten specimens of each species were placed individually into plastic tubes of
2.2 cm diameter and 5.2 cm height that were closed with gauze. The tubes were flooded with aerated water of 9±1°C in a 12
litre-aquarium. Bubbles from an air stone connected to an aquarium pump maintained oxygen saturation in the water as
well as slow continuous circulation of the water body. Air bubbles in the tubes trapped beneath the gauze cover were
sucked off with a pipette. Oxygen saturation was measured with a WTW oxymeter at the beginning and near the end of the
experiment and ranged between 94% and 98%, i. e. around 10.4 mg/I. Typically the animals adhered at the gauze cover.
The tubes were checked daily and the animals were removed when their body became elongated (endosmosis) and they
were unable to walk anymore.

To estimate the surface-volume ratio of the species, the julids were modelled as cylinders with a surface
S=2rcr(r+h) and volume V=7tr2h, leading to a surface-volume ratio R=2(r+h)/rh (r: radius, h: height). However, in
Polydesmus denticulatus the cylinder model would have underestimated the surface-volume ratio because of the well-
developed paranoia. Thus, the calculation was based on a cross-section geometry as described in Figure 1 and leads to
S=27tr(r+h)+4rh, V=4r2h, R=(7t(r+h)+4rh)/2rh. For r and h the average values were taken from Schubart (1934).

The medians of the submersion times were compared by Kruskal-Wallis ANOVA with subsequent multiple a-
posteriori comparisons after Conover (1980). see also Bortz et al. (1989). p. 231. The average ranks for every species
were tested against:

AR(crii) = t(N - k. a / 2)

N.(N + 1)

emp

N-k

RESULTS

Polydesmus denticulatus showed the highest median submersion times of all five species

between P. denticulatus and B. bagnalli is not

(Fig. 2). However, the difference of the medians

Fig. 1. — Cross-section geometry of Polydesmus
denticulatus for the calculation of the
surface/volume ratio R=(rc(r+h)+4rh)/2rh.

significant (Table 1). The maximal tolerance
time in B. bagnalli was even higher (Fig. 2)
with one individual drowning only after 65
days. The other species are significantly less
tolerant. The maximal of survival time in
Ophyiulus pilosus and Leptoiulus proximus
are about two weeks. Females generally
showed a better performance (Fig. 3), but the
differences between sexes were not
significant.

In Figure 3 the median survival time is
plotted against the surface-volume ratio R. The
values are scattered around the regression line
1 and the correlation between survival time and
R is insignificant (r = 0.53). However, if one
excludes males and females of Polydesmus
denticulatus as obvious outliers, the correlation

becomes significant (regression line 2, r =
0.88, P<0.05, two-tailed). If one takes only males into consideration, the fit of the regression
line gets even better and the correlation becomes highly significant (regression line 3, r = 0.99,
P<0,01, two-tailed). This suggests, that in Julidae the survival times are to a high extent
determined by the surface/volume ratio: the smaller the species, the higher the submersion
tolerance.

Source : MNHN, Paris

SUBMERSION TOLERANCE OF SOME DIPLOPOD SPECIES

479

Table 1. — Multiple a-posteriori contrasts after Kruskal-Wallis-ANOVA between species (Conover, 1980). n. s
significant, * = significant (P<0.05).

Polydesmus

denticulatus

Brachyiulus

bagnalli

Cylindroiulus

boleti

Ophyiulus

pilosus

Leptoiulus

proximus

Polydesmus

denticulatus

n.s.

Brachyiulus

bagnalli

n.s.

Cylindroiulus

boleti

n.s.

Ophyiulus

pilosus

n.s.

Leptoiulus

proximus

n.s.

60 -

50 -

</>

2. 30
(0
TD

20 -

10 -

maximum (100%)
upper quartile (75%)

median (50%)

lower quartile (25%)

— minimum (0%)

Polydesmus

denticulatus

Brachyiulus

bagnalli

Ophyiulus

pilosus

Cylindroiulus

boleti

Leptoiulus

proximus

Fig. 2. — Survival times of five diplopod species submerged in water of 9±1°C and 94-98% oxygen saturation

not

Source : MNHN \ Pahs

480

KLAUS PETER ZULKA

DISCUSSION

In P. denticulatus flooding tolerances are highest. But there are small julids like B. bagnalli
with survival tolerances of the same magnitude that do not live in flood-prone habitats. So
submersion tolerance of adults cannot be the only factor to explain why P. denticulatus can live
in flooded places and others cannot. The unusual phenology of the species with reproduction in
summer (SCHUBART, 1934) or the wide oscillations in population density indicating a high
reproductive potential (BLOWER, 1970; ZULKA. 1991) may be additional preadaptations to life in
floodplains.

Fig 3 _ Scalier diagram of submersion tolerance limes against surface/volume ratio in 5 diplopod species 1:
regression line based on all species and sexes. 2: regression line without Polydesmus data, 3: regression line
without Polydesmus and female data.

In the other investigated julids flooding tolerances differ widely, depending mainly on the
size of the species. Since most of them withstand a few days in water, they should be able to
survive short time flooding caused by rain but unable to live in frequently inundated places.

Regarding the physiological mechanisms that allow the species a long survival under

water, three possibilities could be imagined; .

1. The species lives on its anaerobic pathways, and accumulates end products ot

glycolysis (see survey in CRAWFORD, 1978). . .

2. Oxygen supply by diffusion over the whole or over parts of the body cuticle is

sufficient. . . . c ,

3. There are surface structures that maintain an air cover acting as a plastion. buen

structures were found in the tropical millipede Gonographis adisi (MESSNER & ADIS, 1988).

In the present experiment the first possibility cannot be excluded, but it is rather unlikely,
since the tolerance times are long and Polydesmus denticulatus drowns soon when submerged in
unsaturated water (ZULKA, 1991).

Source : MNHN, Paris

SUBMERSION TOLERANCE OF SOME DIPLOPOD SPECIES

481

There are indications for the second possibility at least in julids, since tolerances are highly
correlated with surface/volume ratio.

No continuous air film around the body was observed except for the very first time when
the animals got under water. When the last air bubbles between the hind edges of the metazonite
and the ring duplicatures had already disappeared for a long time they still were active.
However, spiracle structures might act as an interface between water and tracheal air, preventing
the tracheae from being flooded and enabling plastron respiration. In this case, a similar
relationship between tolerances and surface/volume ratio should be expected, since the spiracle
area is correlated with the body surface.

Possibly the main oxygen source in all species is cutaneous diffusion, but in P.
denticulatus special features like spiracle plastron structures (MESSNER, in litt.) could enhance
the air supply under water. This would explain the higher tolerance values in this species
(Fig. 3).

From the present data a clear decision is not possible. A comparison of flooding tolerances
among Polydesmus species covering a broad range of size classes and living in very different
habitat types in the East Alps (TADLER & THALER, 1993) should further elucidate the problem.

ACKNOWLEDGMENTS

I am very indebted to R. L. Hoffman and A. Tadler for valuable discussions, to Z. Kors6s for taxonomic advice
regarding Brachyiulus, to G. Pass for comments on an earlier version of the manuscript, to G. REfMER and A. Tadler for
help during field work, and to G. Schaller for supervising the experiment.

REFERENCES

Adis, J., 1986. — An “aquatic" millipede from a Central Amazonian inundation forest. Oecologia. 68 : 347-349.
Blower, J. G., 1955. — Millipedes and centipedes as soil animals. In : D. K. McE. Kevan, Soil Zoology. Proc. Univ.

Nottingham Second Easter School Agricultural Science. 1953. London : 138-151.

Blower, J. G. 1970. — The millipedes of a Cheshire wood. 7. Zool. Loud., 160 : 455-496.

Blower, J. G., 1985. — Millipedes ( Synopses of the Br. Fauna NS. 35). London, E. J. Brill & W. Backhuys, 242 pp.
Bortz, J.. LlENERT. G. A. & BOEHNKE, K., 1989. — Verteilungsfreie Methoden in der Biostatistik. Berlin, Springer
Verlag, 939 pp.

CONOVER, W. J., 1980. — Practical nonparametric statistics. New York. Wiley.

Crawford, R. M. M., 1978. — Biochemical and ecological similarities in marsh plants and diving animals.
Naturwissenschaften, 65 : 194-201.

ElSENBElS, G. & Wichard, W., 1985. — Atlas zur Biologie der Bodenarthropoden. Stuttgart. New York. Gustav Fischer.
435 pp..

Hoffman, R. L., 1978. — Diplopoda from Papuan caves (Zoological results of the British speleological expedition to
Papua-New Guinea. 1975, 4). Ini. J. SpeleoL. 9 : 281-307.

MESSNER, B. & ADIS, J.. 1988. — Die Plastronstrukturen der bisher einzigen submers lebenden Diplopodenart
Gonographis adisi Hoffman 1985 (Pyrgodesmidae, Diplopoda). Zool. Jb. Anat 117 : 277-290.

Schubart, O., 1934. — TausendfuBler oder Myriapoda 1: Diplopoda. In : F. Dahl. Tierw. Deutschl. 28. Jena, G.
Fischer, 1-318.

Tadler, A. & Thaler, K., 1993. — Genitalmorphologie, Taxonomie und geographische Verbreitung ostalpiner
Polydesmida (Diplopoda: Helminlhomorpha). Zool. Jb. Syst., 120 : 71-128.

Zulka, K. P., 1991. — Uberflutung als okologischer Faktor: Verteilung. Phiinologie und Anpassungen der Diplopoda,
Lithobiomorpha und Isopoda in den FluBauen der March. Dissertation. Wien. Formal- und Naturwiss. Fakultat Univ.
65 pp.

Zulka, K. P., 1992. — Myriapods from a Central European river floodplain. I In : E. Meyer, K. Thaler & W. Schedl.
Advances in Myriapodology.) Ber. nat.-med. Verein Innsbruck, Suppl. 10 : 189.

Source : MNHN, Paris

Eversible Vesicles in Myriapoda

F r antis ek WE YD A

Dept, of Insect Morphology, Institute of Entomology, Branisovska 31,
370 05 Ceske Budejovice, Czech Republic

ABSTRACT

Land arthropods can employ several mechanisms of water uptake. In addition to drinking and water vapour absorption,
the atelocerate arthropods (= Tracheata, Antennata: myriapods and insects) frequently possess special organs for
capillary water uptake, the eversible vesicles (EVs), segmentally arranged paired sacs situated on the ventral side of the
trunk or its part. They are probably functionally analogous to EV-like structures of some Onychophora and Chelicerata.
EVs have been studied by various authors since the mid 19th century but their ecological and evolutionary roles are still
understood poorly. Comparative study of EVs is important for understanding of some acpects of phylogeny of
Onychophora & Arthropoda and their adaptations to terrestrial mode of life. In pauropods Silvestri (1902) and Tiegs
(1947) described two EV-like structures on the collum. Each of these organs consists of several large cells; no
ultrastructural and functionnal data are available. In diplopods EVs are known in a Carboniferous millipede Euphoberia
(Scudder, 1882) as well as in the recent groups (Verhoeff, 1903; Manton, 1958; Dohle, 1988). More than eighty
pairs of EVs are present on the limb bases in Brachycybe lecontii. Specialized cuticle and transporting epithelium are
typical for the absorbing part of an EV of that species. Basic experiments prove that the water absorption from a wet
substrate is possible. In chilopods no EVs are developed (see Dohle. 1988). In symphylans EVs are present in various
numbers (Haase, 1889; Tiegs, 1945; Ravoux, 1962; Dohle, 1988); they are usually located on bases of the legs of trunk
segments 3-10 but reduction of their number is a common feature. Specialized transporting epithelium is present in the
absorbing part of EV. and the water absorption proper has been proved by simple experiments (Tiegs. 1940).

RESUME

Vesicules reversibles chez les myriapodes.

Les arthropodes terrestres sont capables de mettre en ceuvre plusieurs mecanismes d’hydratation. Outre la capacite
d’ ingestion d’eau et d’absorption de vapeur, les arthropodes Atelocerata possedent des organes speciaux destines a
1’hydratation capillaire. les vesicules reversibles (VR), paires de sacs repartis suivant la segmentation dans la partie
ventrale du tronc. Elies ont vraisemblablement une fonction analogue a celle des structures de certains onychophores et
chelicerates. Les VR ont etc etudiees par divers auteurs depuis le milieu du 19e siecle mais leur role ecologique evolutif est
tres mal compris. Une Etude comparative de ces vesicules parait importante pour la comprehension de la phylogenie des
onychophores et des arthropodes et de leur adaptation au mode de vie terrestre. Chez les pauropodes on a decrit deux
structures semblables aux VR sur le collum. Chacun de ces organes comporte plusieurs grandes cellules ; aucune donnee
ultraslucturale ou fonctionnelle n'est disponible. Chez les diplopodes. on les connait tant chez Euphoberia , un diplopode
du Carboniftre, que dans les groupes rEcents. Plus de 80 paires de ces vesicules existent chez Brachycybe lecontii. Une
cuticule specialist et un epithelium de transport constituent les elements typiques de leur partie absorbante. Les
experiences prouvent que (’absorption d'eau a partir d’un substrat humide est possible. Aucune structure de ce genre
n’existe chez les chilopodes. Chez les symphyles, elles existent en nombres varies ; elles sont generalement situees a la
base des panes des segments du corps III & X mais la reduction de leur nombre est un caracttre frEquent. Un Epithelium de
transport specialist est present dans la partie absorbante de la vEsicuIe, et I’absorption d’eau a ete demontree par des
expEriences simples de Tiegs.

Weyda, F., 1996. — Eversible vesicles in Myriapoda. In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy -
Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat ., 169 : 483. Paris ISBN : 2-85653-502-X.

Source MNHN, Paris

Etude comparative des techniques d'echantillonnage
des macroarthropodes saprophages
(Isopoda & Diplopoda)

Etienne BRANQUART & Charles GAS PAR

Unite de Zoologie generate et appliquee, Faculte des Sciences agronomiques, B-5030 Gembloux, Belgique

RESUME

Des peuplements d’isopodes et de diplopodes ont 6te £chantillonnes a 1’aide de trois techniques au printemps 1992
dans trente sites forestiers du sud de la Belgique : pieges d’interception, echantillonnage par quadrats et extraction au
berlese, collecte manuelle dans les bois morts. Les resultats obtenus sont tout h fait differents car chacune des trois
methodes autorise une collecte preferentielle d'especes particulieres. Les avantages, limites et inconv6nients de ces
methodes d’echantillonnage sont mis en evidence et le concept d’activite est discute en rapport avec la biologie des
macroarthropodes saprophages. Des criteres en vue du choix d’une methode sont proposes en accord avec les buts que se
fixent les recherches entreprises.

ABSTRACT

Comparative study of the sampling methods for saprophagous macroarthopods (Isopoda and
Diplopoda).

Woodlouse and millipede populations have been sampled by three methods in Spring 1992 in thirty forest stands of
Southern Belgium: pitfall trapping, berlese extraction and dead wood hand-sorting. The results are quite different because
each of the three methods allows a preferential collect of particular species. The advantages, limitations and drawnbacks
of these sampling methods are underlined and the concept of activity is discussed compared to saprophagous
macroarthropod biology. Criteria for the choice of sampling method are proposed according to the study aims.

INTRODUCTION

II existe plusieurs techniques d'echantillonnage quantitatif ou semi-quantitatif des
populations de macroarthropodes saprophages (isopodes et diplopodes notamment) : (1°)
i’echantillonnage par la methode des quadrats suivi d’une extraction a l'aide d'appareils de type
Berlese-Tullgren ou de type Kempson ; (2°) le piegeage par pieges d'interception, encore appeles
pieges a fosse ou pieges Barber (= pitfall-traps), qui mesure l’activite ou l'abondance relative des
populations ; (3°) des techniques moins usitees de tri manuel ou de chasse a vue, essentiellement
qualitatives.

D'apres un survol de plus de 50 travaux ayant trait a des inventaires regionaux et a des
etudes biocenotiques, 72% des etudes reposent sur un echantillonnage par quadrat suivi
d’extraction selective. Ce type de methodologie semble done constituer une reference, meme s'il

Branquart, E. & Gaspar, C., 1996. — fitude comparative des techniques d'echantillonnage des
macroarthropodes saprophages (Isopoda & Diplopoda). In: Geoffroy, J.-J.. Mauri£s, J.-P. & Nguyen Duy -
Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. nain. Hist, nat., 169 : 485-492. Paris ISBN : 2-85653-502-X.

486

ETIENNE BRANQUART & CHARLES GASPAR

ne permet pas toujours d'obtenir des resultats exhaustifs et donne des estimations d'abondance
biaisees par le rendement parfois deficient des extractions (Van Der DRIFT, 1951 ; EDWARDS
& Fletcher, 1970).

Des techniques de piegeage sont mises en oeuvre dans 45 % des cas, seules ou en
complement a des extractions. Leur emploi relativement commode et peu onereux ainsi que
certaines difficultes inherentes a 1'echantillonnage par quadrat et extraction des populations
evoluant sur des sols ties superficiels (pelouses calcaires, par exemple) semblent justifier leur
utilisation (Davis & JONES, 1978 ; Van Etten & ROOS, 1984 ; SlMONSEN, 1985 ; KlME,
1992 ; Pedroli-Christen, 1993). Par ailleurs, on sait que les techniques de piegeage sont
couramment utilisees pour la capture des arthropodes predateurs epiges comme les arachnides
(MAELFAIT & Baert, 1975 ; UETZ & Unzicker. 1976) ou certains groupes de coleopteres
(THIELE, 1977 ; BRYAN & WRATTEN, 1984 ; DUFRENE, 1992). L'existence d’une relation entre
l'activite de ces arthropodes et leur densite effective constitue par ailleurs un vaste sujet de
discussion (e.a. LUFF. 1975 ; ADIS, 1979 ; BAARS, 1979a ; LOREAU, 1984 ; HALSALL &
WRATTEN, 1988 ; TOPPING & SUNDERLAND, 1992).

Concernant les macroarthropodes saprophages, les etudes detaillees permettant de
comparer les mesures d'activite resultant de piegeages avec les densites obtenues par quadrat et
extraction sont encore rares ou incompletement exploitees. Ce travail a pour but d'effectuer une
etude comparative de differentes techniques d'echantillonnage et de preciser a quoi correspond le
concept d'activite au niveau des populations d'isopodes et de diplopodes.

MATERIEL & METHODES

Unc etude destinee & caracteriser les peuplements de la macrofaune saprophage des sols forestiers du Sud de la
Belgique (entre Sambre et Meuse) est en cours de realisation. Des campagnes d’echantillonnage ont 6t6 menses
simultanement dans 30 sites forestiers representatifs de la diversite pedologique et phytosociologique de cette region.
L'6chantillonnage de la faune a ete realise durant le printcmps 1992, suivant 3 techniques differentes, a savoir des
quadrats suivis detractions au Berlese, des mesures d'activite superficielle ainsi qu'une fouille manuelle du bois mort.
Des echantillons de sol (litiere + sommet de 1'horizon humifere) ont ete preieves sur 6 quadrats de 625 cm2 dans chacun
des sites etudies et extraits durant 3 semaines. Le choix dune surface unitaire de 625 cm2 tient compte a la fois de la taille,
de la densite moyenne et du taux d'agregation des animaux etudies (Sutton, 1972; Blower, 1985; David, 1988). Trois
pieges d'activite ont ete installs par station. II s'agit de pots transparents en P.E.T. de 8 cm de diametre, contenant une
solution diluee de formol, additionnee de quelques gouttes de detergent inodore (TEEPOL) [Dufrene, 1988]. Quant au bois
mort. il a ete fouille minutieusement dans chaque station suivant un effort de chasse constant (2 x 1/2 h par site).

L'efficacite relative des differentes techniques d’echantillonnage vis-^-vis de chacune des 30 especes de
macroarthropodes rencontres durant cette etude (13 especes d'isopodes et 17 especes de diplopodes) a ete evaluee par le
calcul de 2 indices differents. Le premier, appeie indice d'efficacite relative, exprime la frequence d'occurrence de chaque
espece echantillonnee par 3 methodes differentes ; le piegeage, I'extraction et la recherche manuelle dans le bois mort. Le
calcul de cet indice a permis un groupement des especes suivant leur aptitude etre echantillonnees par chacune des 3
techniques. Les groupes proposes ont ete crees par la methode “k-means”, suivant 1'algorithme de groupement
agglomeratif propose par Hartigan & Wong (1979), disponible dans le logiciel SYSTAT.

Comme les valeurs attribu6es l'efllcacite relative des differentes techniques d'echantillonnage sont susccptibles
de varier suivant l'intensite avec laquelle ces dernieres ont ete mises en oeuvre, dies ont ete affinees par le calcul de
V indice d'efficacite du piegeage (ou indice d'activite) propose par Branquart et al. (1995). Celui-ci permet une
comparison de l'activite (resultats des piegeages) et de la densite (resultats des extractions) de la faune saprophage sur
une base quantitative. La valeur de cet indice a ete calcuie en considerant de manierc globale le "pcuplement” des 30
stations; pour chaque taxon, I’indice d'activite mesure le rapport de I'abondance relative estimee par extraction et par
piegeage:

le = 2 x [(nP/NP)/((nP/NP) + (nE/NE))] - 1

ou nP et nE symbolisent le nombre d'individus d'une espece echantillonnes respectivement par piegeage et par
quadrat/extraction alors que NP et NE correspondent au nombre total d’isopodes et de diplopodes echantillonnes par les
memes techniques.

Source : MNHN. Paris

TECHNIQUES D'ECHANTILLONNAGE DES MACROARTHROPODES SAPROPHAGES

487

RESULTATS ET DISCUSSION

Nous avons d'abord cherche a savoir s'il existait une technique qui permette un
echantillonnage exhaustif de la richesse specifique globale propre a un site, celle-ci etant
caracterisee, dans le cas present, par le nombre d'especes echantillonnees par au moins I’une des
trois techniques utilisees. Dans les differentes stations, le nombre de taxons captures par
piegeage, par quadrat-extraction et par chasse a vue dans le bois mort a ete compare au nombre
total d'especes presentes (Fig. 1). Pour chacune des 3 techniques, on trouve une correlation tres
elevee (P < 0.001) entre le nombre d'especes capturees par un seul type d'echantillonnage et le
nombre total d'especes presentes dans un site. La pente de chacune des 3 droites ajustees differe
significativement de 1 (b Piegeage — 0,85; b quadrai-extraclion = 0,65; b recherche manuelle = 0,44), ce
qui signifie qu'aucune technique ne permet de recenser tous les taxons presents dans un site
donne. On constate cependant que le piegeage se montre globalement plus efficace que les 2
autres techniques pour la recherche de la richesse specifique globale.

g 25

0 5 10 15 20 25

Nombre total d’especes

Fig. 1. — Comparison du nombre d'especes de macroarthropodes saprophages echantillonnees par piegeage (carres
noirs), par extraction (signes *'+”) et par recherche manuelle dans le bois mort (carrds blancs) avcc le nombre total
d’especes collect6es par les 3 techniques dans un meme site (diversity stationnelle).

FlG. 1. — Comparison of the number of saprophagous macroarthropods sampled by pitfall traps ( black squares), berlese
extraction (symbol" + ’) and hand-sorting of dead wood (white squares) with the total number of species found in
each stand".

Afin de savoir si certaines especes ne sont pas echantillonnees preferentiellement par l'une
ou 1'autre technique, les 2 indices d'efficacite definis plus haut ont ete calcules pour les
differentes especes. La projection des 30 taxons de macroarthropodes saprophages dans un
systeme de coordonnees triangulaires suivant les valeurs de leur indice d'efficacite relative pour
les 3 techniques d'echantillonnage (Fig. 2) montre que celles-ci sont fortement selectives.
Globalement, 4 groupes de taxons peuvent etre definis (Tableau 1) :

- le groupe 1, constitue de petites especes au mode de vie endoge, bien representees dans
les sols calcimorphes; celles-ci ne sont bien echantillonnees que par quadrat et extraction ;

- le groupe 2, constitue d'especes tres mobiles, souvent peu representatives des milieux
forestiers, essentiellement capturees par piegeage ;

- le groupe 3, compose de taxons a comportement corticole bien marque (efficacite relative
de la recherche dans le bois mort superieure a 35 %), generalement bien recoltes par piegeage et
beaucoup moins par quadrat-extraction ;

- le groupe 4, forme d'especes collectees preferentiellement par piegeage et par quadrat-
extraction, se rencontrant rarement dans le bois mort : il s'agit de taxons qui presentent une

488

ETIENNE BRANQUART & CHARLES GASPAR

importante activite de surface et
et l'horizon humifere.

vivent essentiellement dans la litiere et a l’interface entre celle-ci

Tableau I. - io„ des 4 grcupes !=«««“• ‘’r6Bren,iSl

t.,le t**V««°* <- ««■

APi

APu

PMt

Armadillidium pictum
Armadillidium pulchellum
Craspedosoma rawlinsii
Cylindroiulus punctatus
Nemasoma varicorne
Oniscus asellus

Porcellio monticola
Porcellio scaber
Polyde smus angustus
Tachypodoiulus niger

Brandt, 1833
(Zenker, 1798)
Leach. 1815
(Leach, 1815)

C. L. Koch, 1847
Linne, 1758
Lereboullet, 1853
(Latreille, 1804)
Latzel, 1884
(Leach, 1815)

Allaiulus nitidus
Chordeuma silvestre
Glomeris hexasticha
Glomeris margincita
Ligidium hypnorum
Leptoiulus kervillei
Leptoiulus simplex ssp.
Melogona gallicum

Orthochordeumella pallida
Polydesmus denticulatus
Philo sc ia muscorwn
Porcellium conspersum
Trichoniscus pusillus

(Verhoeff, 1891)

C. L. Koch, 1847
Brandt. 1833
(Villers, 1789)
(Cuvier, 1792)
(Brolemann, 1896)
(Verhoeff, 1894)
(Latzel, 1884)
(Rothenbuhler, 1899)
C. L. Koch, 1847
(Scopoli, 1763)
Koch, 1841
Brandt, 1833

pour l'extraction (31%) ou pour la recherche dans le hois mort (-7%).

FlG 2 — EfficacitS relative des 3
techniques d’echantillonnage
(piegeage, extraction et recherche
dans le bois mort) vis-a-vis de 30
espbces de macroarthropodes
saprophages. Groupement des
taxons suivant leur aptitude h etre
echantillonnes par ces diffdrcntes
techniques.

FlG. 2. — Relative efficience of the three
sampling methods (pitfall¬
trapping. funnel-extraction and
hand-collecting in the decaying
woods) towards 30 saprophagous
macroarthropod species. Grouping

of the taxa after their ability to be
sampled by these different methods.

La Figure 3 illustre la valeur de I'indice d'efficacite du piegeage f "

caWdfur' et 1769 par extraction. A nouveau.

Source : MNHN, Paris

TECHNIQUES DECHANTILLONNAGE DES MACROARTHROPODES SAPROPHAGES

489

les abondances cumulees des macroarthropodes saprophages offrent une image tres differente du
peuplement suivant la technique d'echantillonnage utilisee. Le peuplement global “Berlese”,
moins diversifie, est essentiellement domine par une petite espece d'Isopode, Trichoniscus
pus illus, qui represente plus de 50% des effectifs totaux. Y sont egalement bien representees les
petites especes endogees du groupe 1 ainsi que tous les stades juveniles des diplopodes. En
revanche, le peuplement global resultant des piegeages est beaucoup plus equilibre et diversifie.
II est domine par des individus adultes appartenant aux grandes especes mobiles des groupes 2 et
3. Plusieurs de ces especes presentent des pics d'activite ponctuels tres marques dans le temps,
probablement lies a l'activite sexuelle car les captures comprennent une forte proportion
d'individus males : c'est le notamment le cas de Craspedosoma rawlinsii, Ligidium hypnorum,
Polydesmus denticulatus et Tachypodoiulus niger.

TAXA

Fig. 3. — Valeurs de 1'indice d'efficacite du
piegeage pour differents taxons
(abscisse). La signification des
symboles est reprise au Tableau 1, h
('exception de JIu. JPo. JCh, JCr et
JGI qui correspondent
respectivement aux juveniles de
Iulides, che Polydesmides, de
Chordeumatides, de Craspedosoma-
tides et de Glomerides.

FlG. 3. — Trappability of the taxa. For
symbols, see Table I except for JIu,
JPo, JCh, JCr and JGI that
correspond respectively to
juveniles of Julida, Polydesmida,
Chordeumatidea, Craspedosoma -
[idea and Glomerida.

Rappelons que les resultats exposes ci-dessus ne se rapportent qu'a une seule saison
d’echantillonnage (le printemps) et sont a priori susceptibles de varier en fonction de la periode
de prelevement et du type de milieu etudie. Cependant, on notera que les valeurs de 1'indice
d'efficacite du piegeage correspondent tres bien avec celles qui ont ete calculees sur un laps de
temps plus important dans une etude anterieure effectuee dans la meme region (r = 0,925;
P < 0,001) (BRANQUART et al. , 1995). Les valeurs de cet indice, calculees durant les periodes
d’activite des animaux, semblent done correspondre a une caracteristique intrinseque des taxons
etudies.

Au vu de ces resultats, il apparait clairement que l'efficacite relative des differentes
techniques d'echantillonnage ou de capture depend de l'ecologie des taxons et du stade de
developpement des individus. Les abondances mesurees par quadrat-extraction donnent une
bonne image de la densite des especes dominantes vivant dans la litiere et dans les premiers
centimetres de l'horizon humifere. En outre, elles conduisent a une estimation fiable de
l'abondance de la plupart des stades de developpement, a l'exception des tous jeunes individus
(DAVID, 1988). Cependant, ce type d'echantillonnage est susceptible de donner des resultats
largement sous-estimes lorsqu'il est pratique en dehors des periodes d'activite des
macroarthropodes. Ainsi, durant les periodes critiques de secheresse ou de gel, bon nombre
d'especes s'enfoncent verticalement dans le sol ou se regroupent dans des micro-sites
particuliers, sous les pierres, dans le bois mort et les plages de mousse et echappent ainsi a
i'echantillonnage (SUTTON, 1972 ; GEOFFROY, 1981). De plus, il faut souligner que 1'indice de
dispersion variance/moyenne des arthropodes etudies est presque toujours superieur a 2. Les

490

ETIENNE BRANQUART & CHARLES GASPAR

dispersion variance/moyenne des arthropodes etudies est presque toujours superieur a 2. Les
estimations d'abondance s'accompagnent done souvent d'un coefficient de variation assez
important, meme lorsque le nombre d'echantillons unitaires augmente. Meme si on considere
habituellement que les extractions realisees avec des appareils de type Berlese-Tullgren donnent
des resultats reproductibles (GEOFFROY et al. , 1981), on devra toujours avoir a l'esprit les
differentes sources d'erreur associees a l'estimation des densites de macroarthropodes. Celles-ci
decoulent d'une part, de la distribution fortement agregative des organismes, de leurs migrations
saisonnieres, des rythmes nycthemeraux d'activite et, d'autre part, des caracteristiques
inherentes a la technique utilisee.

Les mesures d'activite se rapportent essentiellement a des individus adultes ou subadultes
qui se deplacent activement a la surface du sol. Ce type de capture permet en outre de recolter les
especes presentes en plus faible densite ou des taxons montrant une preference pour le bois
mort. Du fait de sa bonne efficacite relative pour la majorite des taxons et de son fonctionnement
en continu, le piegeage permet de realiser a peu de frais un inventaire qualitatif assez complet des
populations d’isopodes et de diplopodes presentes dans un site determine. Cet avantage a deja
ete souligne pour d'autres groupes taxonomiques comme les arachnides (UETZ & UNZICKER,
1976 ; Topping & Sunderland, 1992) et les carabides (Thiele, 1977). On notera cependant
qu'il arrive de recolter dans les pieges de type Barber des individus de passage, appartenant a
des formes tres mobiles peu caracteristiques du site etudie (julides, polydesmides,
armadillidiides). Cet effet est d'ailleurs susceptible de s'accentuer dans les habitats defavorables,
ou l'activite locomotrice des arthropodes augmente fortement (GRUM, 1971 ; BAARS, 1979b).
En outre, on a montre que l'efficacite relative du piegeage est eminemment variable d'un taxon a
l'autre : elle depend a la fois de l'“agilite” des individus (SUTTON, 1972), e'est-a-dire de leur
faculte a reperer et a eviter le piege, ainsi que de leur pouvoir de dispersion (BLOWER, 1969), en
relation avec la taille, la mobilite et le taux d'agregation au sein de la population. A ce propos, il
semble que les taxons vivant dans le bois mort, au moins durant une partie de leur cycle de
developpement (groupe 3), forcement repartis de maniere agregative, soient caracterises par un
pouvoir de dispersion important, comme en attestent les valeurs prises par l'indice d'efficacite du
piegeage. On notera encore que l'efficacite relative du piegeage est bien meilleure pour les
groupes predateurs, tels que les carabides, les chilopodes ou certains arachnides, que pour les
especes saprophages, relativement moins mobiles.

Au cours d'un cycle de piegeage annuel realise dans les forets feuillues du sud de la
Belgique, BRANQUART et al. (1995) ont montre que les macroarthropodes saprophages
presentent en general deux periodes marquees d'activite de surface, separees par deux phases de
repos. La premiere est induite par la secheresse estivale (eventuellement accompagnee de
phenomenes de diapause), la seconde est declenchee par le froid hivernal (cf. GEOFFROY &
CELERIER, ce volume). Ceci s'accorde assez bien avec les travaux de Barlow (1957) et de
CLOUDSLEY-THOMPSON (1988) qui ont montre que l'activite des isopodes et des diplopodes est
essentiellement regulee par les conditions climatiques. En particulier, l'activite trophique de ces
animaux est tres nettement regulee par la temperature, l'humidite du substrat et la photoperiode
(VAN DER DRIFT, 1975 ; DAVID, 1987 ; GEOFFROY etal., 1987 ; MOCQUARD et al., 1987). De
plus, le pic d'activite le plus important est generalement situe au printemps, et semble
correspondre a l'activite de reproduction des animaux (recherche de partenaires et de sites de
ponte) (BANERJEE, 1979). En consequence, durant les periodes d'activite autorisees par des
conditions climatiques favorables, les macroarthropodes saprophages partagent leur temps entre
leur activite sexuelle et leur activite trophique. On pourrait done formuler l’hypothese que le
niveau d'activite des isopodes et des diplopodes enregistre par piegeage est proportionnel au role
joue dans la decomposition de la matiere organique tout comme, chez les carabes, le concept de
“densite d'activite” est souvent relie a l'intensite de l'activite predatrice (THIELE, 1977).

On peut enfin se poser la question de savoir s'il existe une relation entre la densite et le
niveau d'activite des differents taxons. BRANQUART et al. (1995) ont montre qu'une relation de

Source :

TECHNIQUES DECHANTILLONNAGE DES MACROARTHROPODES SAPROPHAGES

491

ce type ne pouvait etre verifiee que pour les especes bien echantillonnees par quadrat-extraction
et par piegeage (groupe 4), a la condition que le piegeage soit realise, sinon au cours d'un cycle
annuel complet, du moins durant les phases majeures d'activite des animaux, en accord avec les
etudes de BAARS (1979a) et de LUFF (1982) pour les carabides.

CONCLUSION

Le choix d'une technique d'echantillonnage des populations de macroarthropodes
saprophages edaphiques doit toujours etre determine par le but de l’etude (Tableau 2). Dans tous
les cas, les mesures d’activite devront etre interpretees avec prudence, etant donne qu’elles
dependent a la fois de l'abondance des populations, du degre d'activite des individus qui les
composent ainsi que de leur l'agilite. Ainsi, on pourra comparer valablement le niveau d'activite
de plusieurs populations d’une meme espece, echantillonnees simultanement dans des conditions
aussi similaires que possible. Le piegeage pourra etre utilise avantageusement dans le cadre de la
realisation d'atlas ou de campagnes de surveillance faunistique et, en complement aux methodes
d'extraction, pour comparer des peuplements provenant de differents ecosystemes.
L'echantillonnage par quadrats suivi d'extraction s'appliquera davantage dans le cadre d'etudes
biocenotiques ou de dynamique de populations, car il permet de recolter l'ensemble des stades de
developpement. En milieu forestier, il devra neanmoins toujours etre complete par un
echantillonnage du bois mort par fouille manuelle ou par extraction afin d'obtenir une image
aussi complete que possible des peuplements etudies.

Tableau 2. — Choix d'une technique d'echantillonnage en fonction du type detude.

Table 2. — Choice for a sampling technique in relation to the study.

PIEGEAGE

QUADRAT-EXTRACTION

Inventaires regionaux

* *

Etudes de peuplements

* *

Etudes biocenotiques

* *

Dynamique de population

* *

Mesures d'activite

* *

REMERCIEMENTS

Ce travail a et£ subventionne par un financement I.R.S.l.A. Nous exprimons ici nos remerciements aux societes

anonymes Spa (groupe Spadel) et Chaudfontaine Monopole qui ont fourni gracieusement les bouteilles en P.E.T.

necessaires & la realisation des pi£ges d’interception.

REFERENCES

Adis, J., 1979. — Problems of interpreting arthropod sampling with pitfall traps. Zool. Anz 202 : 177-184.

Baars, M., A., 1979a. — Catches in pitfall traps in relation to mean densities of Carabid Beetles. Oecologia ( Berl .),
41 : 25 - 46.

Baars, M. A., 1979b. — Patterns of movement of radioactive Carabid Beetles. Oecologia {Berl.), 44 : 125-140.

Banerjee, B., 1979. — Diurnal and seasonal variations in the activity of the millipedes Cylindroiulus punctatus
(Leach), Tachypodoiulus niger (Leach) and Polydesmus angustus Latzel. Oikos, 18 : 141-144.

Barlow. C. A., 1957. — A factorial analysis of distribution in three species of Diplopods. Tijdschr. Entom ., 100
349-426.

Blower, J. G., 1969. — Age-structure of millipede populations in relation to activity and dispersion. In : J. G. Sheals,
The soil ecosystem, London : 209-205.

Blower, J. G., 1985. — Millipedes ( Synopses of the Br. Fauna NS. 35). London. E. J. Brill & W. Backhuys, 242 pp.

Branquart, E., Kime, R. D., Dufrene, M., Tavernier J. & Wauthy, G.. 1995. — Macroarthropod-habitat relationships
in oak forests in South Belgium. 1. Environment and communities. Pedobiologia, 3 9 : 243-263.

Bryan, K. M. & Wratten, S. D., 1984. — The response of polyphagous predators of prey spatial heterogeneity;
aggregation by carabid and staphylinid beetles to their cereal aphid prey. Ecological Entomology, 9 : 251-259.

492

ETIENNE BRANQUART & CHARLES GASPAR

Ci.OUDSLEY-ThompSON, J. L.. 1988. — Evolution and Adaptation of Terrestrial Arthropods . Berlin, Springer- Verlag,
141 pp.

David, J. F.. 1987. — Consommation annuelle dune litidre de chene par une population adulte du Diplopode
Cylindroiulus nitidus. Pedobiologia ,30 : 299-310.

David, J. F., 1988. — Les peuplements de Diplopodes dun massif forestier tempere sur sols acides. These Doctorat d’etat
es-Sciences Naturelles, Museum National d'Histoire Naturelle & University de Paris VI, 225 pp.

Davis. B. N. K. & Jones, P. E., 1978. — The ground arthropods of some chalk and limestone quarries in England.
Journal of Biogeography . 5 : 159-171.

Dufrene, M., 1988. — Description d’un piege a fosse original, efficace ct polyvalent. Bull. Annls Soc. r. beige Em.,
124 : 282-285.

Dufrene, M., 1992. — Biogeographie et Ecologie des Communautes de Carabidae en Wallonie. These Doct. Sciences,
Universite Catholique de Louvain.

Edwards, C. A. & Fletcher. K. E., 1970. — Assessment of terrestrial invertebrate populations. In : J. Philupson,
Methods of study in soil ecology, Paris, UNESCO : 57-66.

Geoffroy, J. J., 1981. — Etude d’un 6cosysteme forestier mixte. - V. Traits gcneraux du peuplement de Diplopodes
edaphiques. Rev. Ecol. Biol. Sol , 18 : 357-372.

Geoffroy, J. J.. Christophe, T., Molfetas, S. & Blandin, P., 1981. - Etude dun ecosysteme forestier mixte. - III :
Traits generaux du peuplement de macroarthropodes £daphiques. Rev. Ecol. Biol. Sol , 18 : 39-58.

Geoffroy, J. J., Celerier. M. L., Garay, I.. Rherissi, S. & Blandin, P., 1987. — Approche quantitative des fonctions
de transformation de la matiere organique par des macroarthropodes saprophages (Isopodes et Diplopodes) dans un
sol forestier a moder. Protocoles exp^rimentaux et premiers resultats. Rev. Ecol. Biol. Sol, 24 : 573-590.

Grum, L., 1971. — Spatial differentiation of the Carabus L. (Col., Carabidae) mobility. Ekol. Pol., 19 : 1-34.

Halsall, N. G. & Wratten. S. D.. 1988. — The efficiency of pitfall trapping for polyphagous predatory Carabidae.
Ecological Entomology, 13 : 293-299.

Hartigan. J. A. & WONG, M. A.. 1979. — A K-mcans clustering algorithm : Algorithm AS 136. Applied Statistics ,
28 : 126-130.

KlME, R. D., 1992. — On abundance of west-european millipedes. Ber. nat.-med. Verein Innsbruck, suppl. 10 : 393-
399

Loreau. M., 1984. — Population density and biomass of Carabidae (Coleoptcra) in a forest community. Pedobiologia ,
27 : 269-278.

Luff, M. L., 1975. — Some features influencing the efficiency of pitfall traps. Oecologia (Berl.), 19 : 345-357.

Luff, M. L.. 1982. — Population dynamics of Carabidae. Ann. Appl. Biol., 101 : 164-170.

Maelfait, J. P. & Baert. L., 1975. — Contribution to the knowledge of the Arachno- and Entomofauna of different
woodhabitats. - Part I : sampled habitats theoretical study of the pitfall method; survey of the captured taxa. Biol. Jb.
Dodonea , 43 : 179-196.

Mocquard, J. P., Juchault, P., Jambu, P. & Eustel, E.. 1987. — Essai devaluation du role des crustaces oniscoides
dans la transformation des litieres vegetales dans une foret feuillue de 1’ouest de la France. Rev. Ecol. Biol. Sol, 24 :
311-327.

Pedroli-Christen, a., 1993. — Faunistique des mille-pattes de Suisse (Diplopoda). Documenta faunistica hclvetiae 14,
Neuchatel, Centre suisse de cartographic de la faune, 167 pp. + annexes.

Simonsen, A., 1985. — Interspecific association among millipeds from western and south-eastern Norway. Fauna nor\>.
Ser. B, 32 : 42-44.

Sutton, S. L., 1972. — Invertebrates types. London, Woodlice. Ginn & Company Ltd, 144 pp.

THIELE, H. U.. 1977. — Carabid beetles in their environments. Berlin, Springer- Verlag, 369 pp.

Topping, C. J. & Sunderland, K. D., 1992. — Limitations to the use of pitfall traps in ecological studies examplificd
by a study of spiders in a field of winter wheat. J. Appl. Ecology. 29 : 485-491.

UETZ, G. W. & UNZICKER J. D., 1976. — Pitfall trapping in ecological studies of wandering spiders. J. arachnoi, 3 :
101-1 11.

Van Der Drift, J., 1951. — Analysis of the animal community in a beech forest floor. Tijdschr. Entomoi, 94 : 1-168.

Van Der Drift, J., 1975. — The significance of the millipede Glomeris marginata (Villers) for oak-litter decomposition
and an approach of its part in energy flow, hi : J. Vanek , Progress in soil zoology. The Hague : 293-298.

Van Etten, J. & ROOS, M., 1984. — De invertcbratenfauna van de Zuidlimburgse kalkgraslanden : Landpissebeden
(Crustacea : Isopoda : Oniscoidea), Natuurhist. Maandblad., 73 : 5 -12.

Experimental Behaviour of a Tropical Invertebrate:
Epiperipatus biolleyi (Onychophora: Peripatidae)

Julian Monge-Najera, Zaidett Barrientos & Flanklin Aguilar

Biologfa Tropical, Universidad de Costa Rica, Costa Rica

ABSTRACT

Several aspects of the basic behaviour of Epiperipatus biolleyi Bouvier have been studied experimentally in the
laboratory. The main preliminary results are presented in this short paper.

RESUME

Comportement experimental d’un invertebre tropical : Epiperipatus biolleyi (Onychophora :
Peripatidae).

Divers aspects du comportement de Epiperipatus biolleyi Bouvier ont ete etudies experimentalement en laboratoire.
Quelques resultats pr£liminaires sont prdsentes dans cette courte note.

INTRODUCTION

The limited knowledge on the behaviour of living onychophorans is based on casual
observations (RUHBERG, 1985). Quantitative experimental data are currently limited to feeding
behavior in one species (READ & HUGHES, 1987) and pheromonal function of crural glands in
another (ELLIOTT et al„ 1993). This short paper presents the results of controlled experiments
on the genera] behaviour of Epiperipatus biolleyi Bouvier collected in Coronado, Costa Rica.

PRELIMINARY RESULTS

In choice tests of natural substrates in the field, for unknown reasons bryophyte vegetation
and the soil associated with it were preferred to grass and its soil. In the field, E. biolleyi is
usually found in the moss-substrate interface and in burrows in the soil. Under experimental
conditions, they stayed mainly in the vegetation and rarely in the interface or within the soil. This
difference between the laboratory and the field may be due to the fact that Marchantia and grass
were used, instead of the moss which is also common in the area and provides hiding places for
the animals.

In the laboratory this species was unable to form burrows (N=20) suggesting that, in the
field, they need natural openings on the substrate during daytime. Their feeding and mating
grounds are possibly limited by this factor. Individuals did not show any fidelity to a particular
burrow when given the choice of four identical burrows and within 87 hours they switched
burrows almost three times. This suggests that they show an opportunistic behaviour, entering

Monge - Najera, J.. Barrientos, Z. & Aguilar, F., 1996. — Experimental behaviour of a tropical invertebrate:
Epiperipatus biolleyi (Onychophora: Peripatidae). In: Geoffroy, J.-J., MAURlfcs, J.-P. & Nguyen Duy - Jacquemin.
M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat .. 169 : 493-494. Paris ISBN : 2-85653-502-X.

494

JULIAN MONGB NAJERA. ZAIDETT BARRIENTOS & FLANKLIN AGUILAR

any burrow found nearby when the “resting” time approaches. Most animals enter burrows by
walking foward (N=31) but could also enter backwards. They show a tendency to “rest” facing
the burrow's entrance, possibly to speed reaction to enemies and climatic factors, as well as to
detect passing prey.

No aggressive behaviour for limited burrows was observed. Pairs of animals were seen to
rest with some body contact about half of the time, possibly to reduce desiccation and thermic
stress. The seven basic resting body postures identified which we called: Line, when the body is
straight, U shape, S shape, J shape (head) and J shape (tail), Roll and Ring, were displayed
with decreasing frequency by animals in burrows. The frequency of display differs between
animals on the surface and those in burrows and there is a slight tendency for the S position to
be more frequent for animals in the surface than in the burrows, in which there is a tendency for
the U position to occupy the second place in frequency. Onychophoran body posture has not
been studied previously, with the exception of reports about coiling or shortening in response to
dessication or touch (RUHBERG, 1985).

E. biolleyi hide from light of wave lenght between 470 and 600 nm. Perhaps
onychophorans lack the ability to detect light in the infrared and ultraviolet range (MONGE-
Najera, 1991).

On freshwater, these animals floated and became slightly turgid after the 25 mn. that the
test lasted (N=5). They remained in good health after the test, in contrast to those floated on sea
water (N=4), which died after 14-18 mn. (test suspended). This suggests that onychophorans
could survive contact with water while transported over freshwater (e.g. during floods), but that
dry places are required in natural rafts during possible dispersal across sea (see MONGE-
Najera, 1995).

E. biolleyi produces an adhesive lacking smell and colour. When fresh it tastes bitter to a
vertebrate predator (i.e. humans, N=9). This is in agreement with the idea that the adhesive
evolved originally for defense (MONGE-NAJERA, 1995). The bird Turdus grayi and the snake
Micrurus hemprichii feed on onychophorans in nature but this fact has hitherto remained
unknown to workers in this field because the reports appeared in herpethological and
ornithological publications.

In captivity and with a constant food supply, E. biolleyi survives for up to 150 days. A
marked retraction of the antennae, which become flaccid and curved downwards (sometimes
crossed in an X), and elimination of saliva, adhesive substance, faeces and sometimes immature
embryos are all indications that the animal is severely stressed. Faeces have small rounded
corpuscles which may contain excretory crystals or coccus type bacteria. The species avoids
daylight and even weak wind currents. It moults every 15 days (N=7). The mean speed of
locomotion away from a light source was 1.1-3 cm/s.

AKNOWLEDGEMENTS

We are grateful to William Lamar (University of Texas), Alejandro Sol6rzano and Esther Dominguez
(Universidad de Costa Rica), Wolfgang Bockeler (University of Kiel) and Hilke Ruhberg (University of Hamburg).

REFERENCES

Eliott, S., Tait, N. N. & Briscoe, D. A., 1993. — A pheromonal function for the crural glands of the onychophoran
Cephalofovea tomahmontis (Onychophora: Peripatopsidae). J. Zool., Lond., 231 : 1-9.

Monge-Najera, j., 1991. — An evolutionary interpretation of fertilization patterns in the Onychophora. Onych.
Newsl.y 3 : 2-4.

Monge-Najera, J.. 1995. — Phylogeny. biogeography and reproductive trends in the Onychophora. Zool. J. Linn.
Soc.y 114 : 21-60.

Read, V. M. S. J. & Hughes., R. N., 1987. — Feeding behaviour and prey choice in Macroperipatus torquatus
(Onychophora). Proc. R. Soc. Lond. B , 230 : 483-506.

Ruhberg, H., 1985. — Die Peripatopsidae (Onychophora). Systematik, Okologie, Chorologie und phylogenetische

Aspekte. Zoologica , 137 : 1-183.

Scolopendra morsitans Linnaeus, 1758:
a Characteristic Prey of the African Carpet Viper Echis

ocellatus Stemmier, 1970

Pascal Revault

ORSTOM BP 182 Ouagadougou 01, Burkina Faso

ABSTRACT

This short paper deals with the relationships between Echis ocellatus, the African Carpet Viper, which contributes
largely to human envenomation, and its chilopodan prey, Scolopendra morsitans.

RESUME

Scolopendra morsitans Linne, 1758 : une proie caracteristique de la vipere africaine Echis
ocellatus Stemmier, 1970.

En zone soudano-sahelienne, les vipSrides du genre Echis sont responsables d’une grande partie des envenimations
humaines (Habib. 1992). La predation jouant un role majeur dans la dynamique et l’organisation des peuplements de
reptiles tropicaux (Barbault, 1991), il est interessant de connaitre leur regime alimentaire. 47 Echis ocellatus ont ete
recoltes, dont 37 en octobre 1992 et 8 en avril 1993. dans quatre localites situees aulour de Ouagadougou (Burkina Faso) ;
iixtSs dans du formol a 10%, ils ont <§te disseques et conserves dans de Talcool a 60°. 8 individus avaient un tube digestif
vide. Chez les 39 autres, on trouva 34 scolopendres du genre Scolopendra (dont 9 S. morsitans) et 16 rongeurs (dont 3
Nannomys sp.). 19 viperes contenaient des scolopendres, contre 12 qui avaient consomme des rongeurs ; seules 3 d’entre
elles avaient consomme I un et l’autre. Par ailleurs, furent trouves dans les contenus intestinaux trois restes d’ophidiens
(dont un Psammophis), un crapaud ( Bufo sp.), une araignee, un scarabee. trois fourmis, trois arthropodes non identifies et
deux loholes de tamarinier (Tamarind us indica). II n'y avait pas de correlation entre la taille (ou le sexe) des Echis (35 cm
en moyenne) et le type de proie. La predation des rongeurs par les viperides est bien connue (Villiers, 1975). Elle
lavorise la presence de ces ophidiens aux abords des maisons et dans les champs. La consommation de reptiles, de
batraciens et d’insectes est egalement bien etablie. En revanche, I’importance de la capture de scolopendres etait
jusqu'ici, h notre connaissance, complement ignore, meme si Villiers signale la predation de myriapodes par des
reptiles fouisseurs des genres Typhlops et Leptotyphlops et que Warrell & Arnett (1976) ddcrivent une consommation
occasionnclle de myriapodes par Echis. La capture de S. morsitans , lucifuge et hygrophile, commune dans les
concessions, pourrait expliquer la frequence des rencontres homme/serpent la nuit, au crepuscule et a faube, dans et
autour des habitations, principalement au debut et a la fin de la saison des pluies. Le venin du genre Echis est
essentiellement hemorragipare, tr£s efficace pour la predation des petits rongeurs. On peut se demander aTors dans quelles
conditions s’est developpec, de maniere aussi importante, la predation des scolopendres. La description recente d'une
glande supralabiale h debouche externe dans le genre Echis (Ineich & Tellier, 1992), cas unique chez les serpents,
apporte peut-etre des elements de reponse. Le developpement de cette etude, sur plusieurs cycles pluviometriques, dans
d^autres regions d’Afrique soudano-sahelienne (Nigeria notamment), chez d'autres especes du genre Echis, serait riche
d enseignements pour la comprehension du fonctionnemcnt de la bioc^nose et des interactions entre fhomme et son
milieu.

Revault. P., 1996. — Scolopendra morsitans Linnaeus. 1758: a characteristic prey of the african carpet viper
Echis ocellatus Stemmier, 1970. In: Geoffroy. J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds). Acta
Myriapodologica. Mem. Mus. natn. Hist, nat., 169 : 495-499. Paris ISBN : 2-85653-502-X.

496

PASCAL REVAULT

INTRODUCTION

In the Sudan and the Sahel Savannah, the genus Echis (Reptilia, Viperidae) contributes
largely to bites of human and serious envenomations (HABIB, 1992; HUGHES, 1976; ROMAN,
1980). In the Ouagadougou area (Burkina Faso, Fig. 1), snake bites occui during the iainy
season (Fig. 2), mostly during the night (including dusk and dawn), sometimes in houses, and
principally at the beginning and the end of the season. Therefore it seems necessary to look tor
the factors that determine the encounter between man and snake. .

The rainfall pattern and the predation pressure play a leading role in the dynamics and
organization of tropical herpetofaunas (BARB AULT, 1991). We were curious to know more
about the diet of Echis ocellatus in Burkina Faso.

Fig. 1. — Study area in the Sudan Savannah: Ouagadougou (Burkina Faso). A: Sudan savannah, B: Sahelian steppe,

C: Tropical forest, D: Marsh areas; according to LACOSTE (1990).

MATERIAL AND METHODS

In October (end of the rainy season) and in April (beginning of the rainy season), a snake collection was carried

out, corresponding to the first and last bite peak (Fig. 2). t . .

Echis ocellatus are crepuscular vipers, living especially in the Sudan Savannah (Fig. 1). Peasants captured the
vipers from an area 100 km around Ouagadougou as they encountered them. The snakes were preserved in 10% lormalin
solution, after an injection with formalin (anus, tail). They were dissected in the 15 days after capture. The size, the sex
were noted and the digestive content analysed (from the cardia to the anus). The animals were preserved in 60 alcohol
(Laboratoire des Reptiles et Amphibiens MNHN, Paris, n° 3110-31 17).

Source : MNHN, Paris

SCOLOPENDRA MORSITANS : PREY FOR THE AFRICAN CARPET VIPER

497

RESULTS

47 Echis ocellcitus were captured (39 in October in three localities: Gonse, Sapone and

250 •

Fig. 2. — Correlation between snake bites and rainfall: Fig. 3. — Digestive contents of 39 Echis ocellaius :
number of patients monthly hospitalized at the H. taxonomic distribution of the 64 prey-types

N. of Ouagadougou (n = 83) during one year / Rain identified in the gut after dissection. Tamarind =

fall (665.5 mm in one year) from "Direction de la tamarind leaflets).

Meteorologie”.

Sapogo, and 8 in April in one locality: Kouloulou). The average length of 41 individuals, was
35 cm (2 individuals were 18 and 21 cm, the others were longer than 30 cm). The sex-ratio was
* 1 . Five animals had non mature eggs (2 in April and 3 in October).

There was no difference of size (or sex) between the vipers with no digestive content and
the others. Equally, there was no correlation between the type of prey and the size (or the sex) of
the Echis.

8 Echis had no gut contents. The gastro-intestinal contents of the 39 others are presented in
Figure 3. 34 centipedes, genus Scolopendra, were found. 9 were S. morsitans (J.-M.
DEMANGE det.). The average size, amongst 15 individuals, was 8 cm (from 5 to 12 cm). 21
snakes were concerned.

16 rodents were found. 3 belonging to the genus Nannomys (M. TRANIER det.). The
average size, found in 6 individuals, was 8 cm (from 6 to 12 cm). 15 snakes were concerned.

In three cases only, the snake had swallowed a rodent and a Scolopendra.

The other prey were: 3 snakes (1 Psammophis sp.), 2 tamarind leaflets, 1 Solifugae, 1
Coleoptera, 3 ants and the rest non identifiable arthropods.

Scolopendra is a common prey of vipers (Fig. 4) but does not appear to be inportant in rice
fields at Sapone in comparison with the other areas (Sudan Savannah).

DISCUSSION

The centipede Scolopendra morsitans is lucifugal and requires moisture. Snakes may enter
houses at night, dusk and dawn, looking for centipedes and this may explain the frequency of
man-snake encounters.

This study encompassed only one rainy season. Further data are required on the age,
predators and reproduction of Echis ocellatus.

BARB AULT (1991) doesn't describe such a characteristic diet for Echis ocellatus, in the
Sudan Savanah (Ivory-Coast) even if WARRELL & ARNETT (1976) describe that Echis sp. in
Nigeria occasionnally swallow Myriapoda. In the same region, VlLLIERS (1975) cites the

498

PASCAL REVAULT

predation of Myriapoda by the genus Typhlops and Leptotyphlops, both burrowing snakes. In
South Africa, Colubridae from the genus Asparallactus (BROADLEY & COK, 1993) are called
centipede eaters, described as having an effective venom, making their prey helpless. But to our
knowledge, the selective predation on centipedes amongst Viperidae, and especially on
Scolopendra, has not been remarked up to now. The venom of Echis is essentially
haemorrhagiparous, very effective for the predation of small rodents but not for centipedes. It
can be asked under what type of conditions did the predation of centipedes develop in such a
characteristic way. Is this particular diet found uniquely in the area of Ouagadougou, amongst
the genus Echis ? It is possible that the supra labial gland described by INEICH & TELLIER (1992)
may be important in this respect.

no digestive content

tamarind

batracian

ophidian

other arthropods

rodent

Scotopendra

■ Koukoulou 04/93:8

□ Total 10/92:39

□ Gons6: 8

□ Sapone: 5
IESapogo:26

Lewis (1970) has pointed out that
Scolopendra morsitans is atypical, amongst
the scolopendromorph centipedes that have
been studied, because it seems to be surface
active throughout the year. The life cycle is
completed in one year and young
individuals appear in March and again in
October (Lewis, 1974).

In order to develop this study in the
future, observations are required on several
seasonal cycles and in other regions of
Sudan and Sahel savanna. Other species of
Echis should be investigated in order to
enrich the comprehension of the biocenosis
and the interaction between man and his
environment.

Fig. 4. — Relative importance of prey-types captured by 47
Echis ocellatus in 4 snake sampling sites.

ACKNOWLEDGMENTS

I am much indebted to Dr. M. Goyffon for his assistance in collecting information and to Dr. I. Ineich for his
help to conserve and analyse the snake collection.

My thanks are also due to Prof J.-M. Demange for the identification of centipedes, and to Dr. J. C. Gautun for his
support related to collecting the snakes.

REFERENCES

Barbault, R., 1991. — Ecological constraints and community dynamics: linking community patterns to organismal
ecology. The case of tropical herpetofaunas. Acta (Ecologica, 12 : 139-163.

Broadley, D. G. & COK, E. V., 1993. — Snakes of Zimbabwe. Bundu Series, Zimbabwe, Longman, 152 pp.

Habib, A. G., 1992. — Tropical snake bite in Northern Nigeria. A clinical review. Nigerian Medical Practitioner , 23 :
3-8.

Hughes. B., 1976. — Notes on African Carpet Viper Echis carinatus, Echis leucogaster and Echis ocellatus (Viperidae,
Serpentes). Rev. Suisse Zool., 83 : 359-371.

Ineich, I. & Tellier, J. M., 1992. — Unc glande supralabiale & debouche externe chez le genre Echis (Reptilia,
Viperidae), cas unique chez les serpents. C. R. Acad. Sci. Paris, 315 : 49-53.

LaCOSTE, Y., 1990. — Atlas 2000. La France et le monde. Paris, Editions Nathan, 160 pp.

Source : MNHN. Paris

SCOLOPENDRA M0RS1TANS : PREY FOR THE AFRICAN CARPET VIPER

499

Lewis, J. G. E., 1970. — The biology of Scolopendra amazonica in Nigerian Guinea savannah. Bull. Mus. natl. Hist,
nat., 41, suppl. 2 : 85-90.

Lewis, J. G. E., 1974. — The ecology of Centipedes and Millipedes in Northern Nigeria. Symp. zool. Soc. Lond ., 32 :
423-431.

Roman, B., 1980. — Serpents de Haute Volta. Ouagadougou, Editions du CNRST, 129 pp.

VlLLlERS, A., 1975. — Les serpents de I'Ouest africain. Initiations et etudes africaines. N° II, 3^me edition. Dakar,
University de Dakar. IFAN, 195 pp.

Warrell, D. A. & Arnett, C., 1976. — The importance of bites by the saw scaled or carpet viper ( Echis carinatus ):
Epidemiological studies in Nigeria and a review of the world literature. Acta Trop., Basel , 33 : 307-341.

Source : MNHN , Paris

The Life Cycle of Cylindroiulus latestriatus

(Curtis, 1845)

Karin VOIGTLANDER

Staatliches Museum fur Naturkunde Gorlitz, PF 300 154, D-02806 Gorlitz, Germany

ABSTRACT

A population of Cylindroiulus latestriatus from Eastern Germany has been studied in 1983. On the basis of the
characterization of stadia, by the defence gland method, all possible variations of increments of ring numbers are given
and a scheme of developmental pathways is built up. By examining individual development and the resulting main
pathways a review of the regularities in the development of ring increment in the species C. latestriatus can be made. The
increment of body rings together with the increase of body length, volume, and biomass allows to clarify the regulatory
mechanisms in the development of julids. The age structure and life cycle of the East German population of C. latestriatus
is compared with those of other authors, made in different countries and habitats, and with different methods of stadial
determination. They are generally in agreement with only minor differences.

RESUME

Le cycle de vie de Cylindroiulus latestriatus (Curtis, 1845).

Le cycle de vie d'une population de Cylindroiulus latestriatus d’Allemagne orientale a ete etudiSe selon la methode de
comptage des glandes repugnatoires et comparee aux resultats obtenus par d'autres auteurs. Les divers modes d’acquisition
de nouveaux anneaux sont explores et l’hypothese d’un mecanisme regulateur de l’ontogenese des julides est testee sur le
materiel ctudie.

INTRODUCTION

The study of the post-embryonic development and the laws of anamorphosis in millipedes
has a high level of interest. Life histories offer phylogenetical information and help to clarify
relationships between millepede orders.

Despite there being more problems to overcome than in other groups (e.g. Chordeumatida,
Polydesmida) the order Julida is. with 41 species investigated for developmental pathways, one
of the most frequently studied orders. Difficulties in investigations arise from the overlap of ring
numbers of successive stadia. The total number of rings cannot be used to determine the stadia.
Maturity is reached in several different stadia and ages (see ENGHOFF, DOHLE & BLOWER,
1993). For this reason many more investigations on the life cycles of this order are necessary.

The species C. latestriatus has been studied very intensively by BLOWER & GaBBUTT
(1964) and by BlERNAUX (1972). Other authors (COTTON & MILLER, 1974; LANG, 1954) have
given additional remarks to the life history of this species. No detailed investigations were
known from Germany.

VoigtlAnder. K., 1996. — The life cycle of Cylindroiulus latestriatus (Curtis, 1845). In: Geoffroy, J.-J..
M AURifes, J.-P. & Nguyen Duy - Jacquemin, M„ (eds), Acta Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 501-
508. Paris ISBN : 2-85653-502-X.

Source : MNHN} Paris

502

KARIN VOIGTLANDER

This situation offers the rare possibility of comparing the results from different sites and
by various methods of stadial determination.

MATERIAL AND METHOD

About 100 specimens of C. latestriatus were collected from the litter layer under hazelnut bushes in a garden at a
very dry site in NE Germany (Premnitz near Brandenburg) in 1983. A more detailed description of the habitat is given by
VOIGTLANDER (1987).

Irrespective of stadia and age, the specimens were reared at room temperature in Petri dishes with a natural
substrate on a layer of moist filter paper to study the further development.

To identify the stadia of the freshly collected specimens I used the defence gland method (Halkka, 1958;
Brookes, 1963). Measurements of length and width as well as weighing were made on living specimens (for the methods
see VOIGTLANDER, 1987).

CHARACTERIZATION OF STADIA AND DEVELOPMENTAL PATHWAYS

The Table 1 shows the total numbers and increments of defence glands related to each
moult. The first and by far the biggest variation in increase of rings (or defence glands
respectively) occurs at the moult into the fourth stadium. From this there follows a nearly regular
decrease both in variation and in newly occuring ring numbers.

Table 1. — Total number and increment of defence glands in C. latestriatus.

increment of defence glands

stadium

individuals defence glands 8

7 6

4 3

100.0

III

6.4

3.2 12.9

51.6

25.8

10-14

3.3 46.7

43.3

6.7

15-19

40.5

42.9

16.7

19-25

4.4

66.7

24.4 6.7

24-29

4.4

28.9 66.7

Vffl

28-32

•L

10.0

88.0

2.0

31-34

6.9

65.5

27.6

32-36

12.5

87.5

33-38

100.0

34-39

100.0

XIII

38-40

100.0

XIV

Source : MNHN, Paris

THE LIFE CYCLE OF CYIJNDROIULUS LATESTRIATUS

503

On the average, the highest increase in rings is gained during the moult into the Vth
stadium. The following moultings show progressively fewer numbers of added rings, down to
1.0 from the Xlth stadium on.

The pathways of defence gland increments (Fig. 1 ) are originally determined by the “basic
groups”, according to PEITSALMI (1981), occuring at the IVth stadium. Basic groups with the
highest and lowest gland numbers become “normalized” during the following 3 moults.

Regarding the number of observed moulting events (indicated by the little numbers in
Fig. 1), it is clearly visible that 2 or 3 main pathways of development (thick lines) dominate.

Fig. 1 . — Observed defence glands and the main developmental pathways of C. latesiriatus.

In Figure 2 the post-embryonic development of C. latestriatus is shown as a result of
studies in an oak wood on sandy soil in England (BLOWER & GABUTT, 1964), in marram dunes
on the east coast of Scotland (COTTON & MILLER, 1974), in a field of carrots in Belgium
(BlERNAUX, 1972) and in a garden on sandy soil in Germany (this paper). The methods used
for estimations were the analysis of discontinuities of dimensions (BLOWER & GABBUTT,
1964), the observation of eye rows (COTTON & MILLER, 1974; BlERNAUX, 1972) and the
number of defence gland (BlERNAUX, 1972 and this paper).

504

KARIN VOIGTLANDER

401

</)

-U

VIII

VII

Cotton G
Miller (1974)

VIII ^

Biernaux (1972)

Blower G
Gabbutt(1964)

XIII

□ ■ □ U □

juveniles free-living premature adult only ^o

with yolk juveniles 66 and rfrfandcxj)

Fig. 2. — The post-embryonic development in C. latestriatus studied by different authors. The first stadium to contain
mature animals is the stadium VII. Results by VoigtlAnder deal with this paper.

Up to stadium IV there are few differences between the results. Later on variations
increase, for example specimens of stadium XI show a maximum of 41 rings according to
BLOWER & GabbuTT (1964), 42 rings according to BIERNAUX (1972) and 43 rings after the
results of the present study.

Males reach stadium IX only, irrespective of the indicated ring numbers according to all
authors. The highest stadium reached by females is stadium XIII, showing 43 rings (BLOWER &
GABBUTT, 1964) or 45 rings (this work). COTTON & MILLER (1974) estimate the highest
stadium being X with up to 42 podous rings. They argue that some females which were placed
in stadia IX and X in their investigation may belong to stadia XI or XII because it is difficult to
determine the addition of ocelli to the ocular field at higher stadia.

The results of the investigations of stadial development of C. latestriatus correspond with
the rules known for other Juliformia. The species belongs to the group of anamorphotic julids
with an indefinite number of body ring increments.

As in other related species, such as Cylindroiulus punctatus or Kryphioiulus occultus,
older stadia add one new body ring only at each moulting.

Source : MNHN, Paris

THE LIFE CYCLE OF CYLINDROIULUS LATESTRIATUS

505

C. latestriatus reaches stadium XIII as a maximum. The low number of stadia seems to be
typi-I for Cylindroiulinae (for example Enantiulus nanus XV, K. occultus XIV, C. punctatus

GROWTH INCREMENTS

Because of the relatively low number of specimens observed, these results are not
significant but they indicate the overall tendency.

males

females

Fig. 3. — Mean increase of length in C. latestriatus (p = premature, m = mature individuals).

volume

males

females

Fig. 4. — Mean increase of volume in C. latestriatus.

506

KARIN VOIGTLANDER

Each moult of C. latestriatus is correlated with a relatively constant increase of length (Fig.
3). The same is true for the volume except for the Xlllth stadium of females (Fig. 4). Judging
from the biomass (Fig. 5), more or less identical results can be seen, with the exception of some
flattened sections of the curves for gravid females according to the number of eggs. In contrast
the numbers of newly formed rings decrease at each moult.

mg <i

35-

Fig. 5. — Mean increase of biomass of C. latestriatus.

ENGHOFF, DOHLE & BLOWER (1993) have suggested 4 hypotheses to explain the range of
addition of rings. According to these, the addition can depend on:

1. the available energy ,

2. the randomness of splitting of the germinal field,

3. a combination of 1+2,

4. a predisposition to variable increments for whatever reason.

As to point 1, observations on C. latestriatus cannot be explained by different energy
supply if we suppose that available energy is not continously decreasing from stadium IV to the
end.

Furthermore the growth in biomass is not closely related to the numbers of added rings.

Regarding hypothesis 2 it should be considered that the first 3 stadia have an entirely
constant increment, and after that there is a nearly constant decrease of addition from the Vth
stage on. These events cannot be explained by randomness.

A predisposition of some kind may be assumed, although the cause has not yet been
identified.

biomass

without eggi

vi vii viii ix

xii stadia

EGG LAYING AND EARLIEST DEVELOPMENT

BLOWER & Gabbutt (1964) did not find any eggs neither in the field nor in culture. They
assume, that eggs are laid in small numbers and not aggregated together in a typical nest.

In this study one nest was found consisting of 6 eggs in a rearing vessel in the laboratory
at the beginning of May. The pupoids “hatched” after two weeks and developed after 4 days into
a typical stadium I with 3 pairs of legs. The duration of this stadium was 3 to 4 days. The
following stadium II is the first free living and feeding one. All observed young specimens
overwintered at stadium V.

Source :

THE LIFE CYCLE OF CYLINDROIULUS LA TESTRIA TUS

507

SEXUAL DIMORPHISM AND BEGINNING OF MATURITY

The sexes of C. latestriatus can be distinguished first at stadium V. Premature males
clearly show the absence of the leg pairs on the 7th body ring. Animals with appearance of
females need a further moult to clarify the real sex.

In all investigations made in C. latestriatus both sexes reached maturity in stadium VII or
rarely in VIII. In females the eggs, which can be seen through the body wall, indicate the
maturity.

Blower & Gabbutt (1964) have investigated precisely the gonopod development of
some species. They pointed out, that 2 to 3 gonopod stadia exist in C. latestriatus , which extend
over 5 developmental stadia of the males. The primary gonopod stage always coincides with the
fifth stage, the secondary and also the tertiary with the sixth. Mostly, maturity is reached in
stadium VII, rarely the tertiary gonopod stage extends to stadium VIII or IX (1 male).
Corresponding with their investigations only 20% premature males in stadium VIII were found
here, but no premature specimens in stadium IX.

Females reached the highest stadium of XII in their natural habitat (Table 2) and stadium
XIII in the laboratory (only ?). Males reach a maximum of stadium VIII (with the exception of
one male in stadium IX). This suggests that males die mostly after one or certainly after two
reproduction periods.

Table 2. — Moulting activity of C. latestriatus.

Month

Febr.

March April

May

June

July

Aug.

Sept.

Oct.

Nov.

Dec.

juvenile + premature

mature

Total

The temporal distribution of the moults is shown in Table 2. Juveniles, premature and
mature individuals are able to moult over the whole year, but with different maxima. Adults
mostly moult in autumn, juveniles and prematures in spring and summer. This agrees with other
characteristics of the species such as the high locomotory activity of adults in spring during the
search for the partners. A moult is the peak of the physiological activity and can only take place
after the end of a reproduction period. Young individuals can moult in spring too, because they
do not reproduce.

AGE STRUCTURE AND LIFE CYCLE

The age structure of the population is strongly connected with the life cycle of C.
latestriatus. The population consists of all stadia and age groups throughout the whole year with
exception of the youngest stadia, which can only be found after the egg laying period.

As a consequence of the different speeds of the individual development, individuals of
several generations can belong to the same stadium. This may firstly be observed in stadium VI
with two overlapping generations. In stadium VIII the overlap can include 3 generations. This
tact makes the interpretation of samples very difficult. This is the reason why a combination
between field observations and results of laboratory rearings was necessary.

The egg laying takes place in spring. The young individuals develop, according to the
results of all authors, into the stadia (III), mostly IV or V (and VI) in which they overwinter.
Individuals of stadium VI may belong to the first generation or to the second if they started from
overwintered stadium III. Provided that the maturity moult takes place in autumn

508

KARIN VOIGTLANDER

(VOIGTLANDER, 1987) it is concluded, that the first reproduction is possible at the 2nd or 3rd
year of life. This short time for development is very uncommon for julids.

Males die after one or at least two reproduction periods. They live for a maximum of 3 or 4
years, as a rule only 2-3 years. Females have on average a higher expectation of life usually 3-4
years, with a maximum of 7-8 years.

REFERENCES

Biernaux, J., 1972. — Chorologie et 6tude biologique comparee de deux families de Myriapodes - Diplopodes beiges:

les Blaniulidae et les Iulidae. Dissertation, Gembloux, Faculte des Sciences agronomiques de I'Etat, 193 pp.

Blower. J. G., & Gabbutt, P. D., 1964. — Studies on the millipedes of a Devon oak wood. Proc. zool. Soc. London,
143 : 143-176.

Brookes, C. H., 1963. — Some aspects of the life histories and ecology of Proteroiulus fuscus (Am Stein) and Isobates
varicornis (Koch) (Diplopoda) with information on other blaniulid millipedes. Thesis, Univ. Manchester.

Cotton, M. J.. & Miller, P. F., 1974. — A population of Cylindroiulus latestriatus (Curtis) on sand dunes. Symp. Zool.
Soc. Lond.. 32 : 589-602.

Enghoff, H., Dohle W. & Blower, J. G., 1993. — Anamorphosis in Millipedes (Diplopoda) - The present state of
knowledge with some developmental and phylogenetic considerations. Zool. J. Linnean Soc., 109: 103-234.
Halkka, R., 1958. — Life history of Schizophyllum sabulosum (L.) (Diplopoda, Julidae). Ann. Zool. Soc., Zool. bot.
Fenn. " Vanamo ”, 19 : 1-72.

Lang, J.. 1954. — Postembryonalentwicklung der Diplopoden. Vest. Cs. spol. zool. (Acta soc. zool. Bohemoslov.),
18 : 161-176.

Peitsalmi, M.. 1981. — Population structure and seasonal changes in activity of Proteroiulus fuscus (Am Stein)
(Diplopoda. Blaniulidae). Acta Zoologica Fennica, 161: 1-66.

VOIGTLANDER, K., 1987. — Untersuchungen zur Bionomie von Enantiulus nanus (Latzel, 1884) und Allajulus occullus C.
L. Koch, 1847 (Diplopoda. Julidae). Abh. Ber. Naturkundemus. Gorlitz, 60 : 1-116.

Source : MNHN, Paris

Life-Cycle of the Millipede Melogona voigti
(Verhoeff, 1899) from a Suburban Forest in South

Bohemia

Karel TAJ OV SKY

Institute of Soil Biology, Academy of Sciences of the Czech Republic
370 05 Ceske Budejovice, Czech Republic

ABSTRACT

A population of the millipede Melogona voigti in a suburban deciduous forest in South Bohemia has been studied by
soil sampling. The changes of density in the course of the year and the post-embryonic development have been
described. The biological cycle was annual for the greatest part of the population. The effects of climatic factors on the
life-cycle of Chordcumatida are discussed.

RESUME

Cycle de vie du diplopode Melogona voigti (Verhoeff, 1899) dans une foret suburbaine decidue
du sud de la Boheme.

Une population de Melogona voigti d’une foret decidue suburbaine du sud de la Boheme a ete etudiee par
echantillonnage du sol. Les variations de densite au cours de Tannee et le developpement post-embryonnaire sont
decrits. Le cycle est annuel pour la plus grande partie de la population. Les effets des facteurs climatiques sur le cycle des
chordeumatides sont discuss.

INTRODUCTION

Three species of the genus Melogona Cook, 1895 (= Microchordeuma Verhoeff, 1896) are
distributed in Central and North-West Europe. Post-embryonic development of M. scutellare
(Ribaut, 1913) was described by BLOWER (1978, 1979) and that of M. gallica (Latzel, 1884) by
David (1984). Some primary data for the third species M. voigti (Verhoeff, 1899) were
published by VERHOEFF (1913, 1928) and later supplemented by SCHUBART (1957) and by
DUNGER & STEINMETZGER (1981). M. scutellare, belonging to the subgenus Chordeumella
Verhoeff, 1897, has eight post-embryonic stadia and adults with 28 body segments. M. gallica
and M. voigti belong to the subgenus Melogona Cook, 1895, the adults of which have 30 body
segments and mature one stadium later.

During the faunistic research in South Bohemia, a suburban forest near Ceske Budejovice
was discovered to contain a population of Melogona voigti. To obtain data for completion of its
life-cycle, soil sampling was used.

Tajovsky, K., 1996. — Life-cycle of the millipede Melogona voigti (Verhoeff, 1899) from a suburban forest in
South Bohemia. In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica.
Mem. Mus. natn . Hist, nat 169 : 509-514. Paris ISBN : 2-85653-502-X.

510

KAREL TAJOVSKY

STUDY SITE AND METHODS

The population of Melogona voigii was studied in the suburban deciduous forest Stromovka in south-west
outskirts of Ceske Budejovice-cily in South Bohemia (Czech Republic). Stromovka (430 m a.s.L, average air
temperature 7.8°C, precipitation 620 mm. see Fig. 2) is an allochtonous sparse wood dominated by Populus nigra L. and
Alnus glulinosa (L.) Gaertn. The plant cover consists mostly of Urtica dioica L. and Filipendula ulmaria (L.) Max. The
site is characterized by brown soil type (gleic cambisol) with mull-moder to mull humus horizon. pH (H20) 4.4.

Ten soil samples (sampling area 1/16 m2, depth 5 cm) were taken at approximately fortnightly intervals from
February 24, 1992 until April 19, 1993. Millipedes were heat extracted from soil samples by modified Kempson
extraction apparatus (Kempson el al., 1963). For the evaluation of post-embryonic development of M. voigii, the
numbers of body segments, pairs of legs and ocelli were counted. Females were dissected and the presence or absence of
eggs investigated.

RESULTS

Population density and dynamics during the year

Sequential fortnightly sampling confirmed, that the millipede M. voigti is active during the
whole year. Mean annual density of the population under study was 60.6 ind.m2. During the
winter months up to March, adults only were present (Fig. 1). The first increase in density in
mid-April can be connected with the increasing activity of adults. Practically all successive peaks
of density were due to culminations of separate post-embryonic stadia: 11.5. - stadium III, 26.5.
- stadium IV, 13.7. - stadia V and VI, 24.8. - stadium VII, and 4.11. - stadium IX (new adults).
The marked depression in mid-June was probably due to the rainfall deficiency and the
consequent drying up of the litter and upper soil layer. The maximum number of adults was
observed in November. In winter months the density of millipedes decreased.

Mating and oviposition

Both, mating and oviposition appeared to take place early in spring, judging from the
appearance of stadium II as from the first half of April. In the field the mating was observed still
in April. Dissection of females showed the presence of eggs in the ovarium from November until
the end of April. Therefore it was not possible to be more precise about the time of oviposition
or the number of eggs laid.

Post-embryonic stadia

Numbers of body segments (podous and apodous) and numbers of pairs of legs for
individual stadia of M. voigti are given in Table 1. M. voigti has nine post-embryonic stadia.
Sexual differentiation is in the last three stadia. Numbers of ocelli and their arrangement in the
ocular field based on the material from South Bohemia are given in Figure 3.

Table 1. — Post-embryonic stadia of Melogona voigti. Pleurotergites - number of podous rings, apodous rings and
telson. Ad = adults, F = female. M =male.

Stadium:

III

VII

VIII

IX(Ad)

pleurotergites:

1 1

podous:

apodous:

leg pairs:

49(F)

”

45(M)

Source :

LIFE-CYCLE OF A MILLIPEDE OF A SUBURBAN FOREST IN SOUTH BOHEMIA

511

Fig. 1. — Density of dynamics of the millipede M. voigti showing the separate post-embryonic stadia.

FIG. 2. — Mean monthly temperature and monthly precipitations for Ceske Budejovice.

Post-embryonic development

The adults were present until the end of May (Figs 4 and 5). The first stadium was not
recorded by the method used. Individuals of stadium II were noted in mid-April. Stadium III
was observed from the end of April until the end of May with a maximum in the first half of
May. Stadium IV appeared as from the end of May in the highest density, then it was present
during the whole of June through to mid-July, and was also noted at the end of August. Stadium
V was present in soil samples from the end of June until the end of August, stadium VI from
mid-July up to September 21. Both stadia VII and VIII observed from the end of July and mid-
August, respectively, were present up to the beginning of October. At that time the first adults
(stadium IX) appeared. The absence of stadia VII and VIII in mid-October and following
samples confirmed the end of post-embryonic development.

KAREL TAJOVSKY

512

0.5 mm

In the period March 30 - July 13, in
addition to beside stadia II to VI, the
individuals of stadia VI. VII and VIII were
also present. This means that a smaller part
of the previous year's population did not
quite complete its development and
overwintered in the same locality as stadia
VI and/or VII and VIII. The successive
sampling did not quite elucidate, whether
this part of the population finished their
development during this second year.
During the next March and April 1993. the
younger overwintering stadia VI, VII and
VIII were not sampled.

M. voigti is therefore largely an
annual millipede, however a part of the
population can take more than one year to
complete its life-cycle.

Fig. 3. — Growth of the ocular field from stadium II to
maturity. Ocelli invariably present are cross-
hatched, ocelli not always present are open,
the rows of ocelli are marked.

DISCUSSION

Numbers of body segments and numbers of pairs of legs for individual stadia of M. voigti
correspond to the general pattern for the suborder Chordeumatidea (BLOWER. 1984) and are in
agreement with the data for M. gallica (DAVID, 1 984).

VERHOEFF (1913, 1928) described the occurrence of the adult millipedes of M. voigti
from October until May and of the juveniles in the remaining part of the year. SCHUBART (1957)
confirmed these data, and DUNGER & STEINMETZGER (1981) found the adults even in mid-
June. In contrast to VERHOEFFs' data (VERHOEFF, 1928), the appearances of individual stadia
III to VII were always later and the development up to stadium IX was shorter (Fig. 5). In this
way, post-embryonic development of M. voigti differs from that of the related West-European
species M. gallica (Fig. 5). A shorter period of development with a fast sequence of older stadia
V-VHI of the population under study may be the result of the colder continental climate.

The influence of climatic factors on life-cycles of Chordeumatida is known. In addition to
prolongation of the time to maturity, longevity and a general slowing down of the life-cycle with
the increase in altitude and decrease in temperature (MEYER, 1990), there is evidence of the
interruption of the life-cycle in a part of the population due to the unfavorable microclimatic
conditions (dryness, coldness) and of the life-cycle prolongation into the following year
(PEDROLI-CHRISTEN, 1978; David, 1989). The prolongation of life-cycle and the suggestion of

Source : MNHN \ Paris

LIFE-CYCLE OF A MILLIPEDE OF A SUBURBAN FOREST IN SOUTH BOHEMIA

513

a two year development for M. gallica were described for a French population (David, 1984)
and noted for a British population as well (BLOWER, 1984). David (1984) mentioned that this
phenomenon can be either regular, i.e. a certain smaller part of the population always hibernates
as a juvenile stadium (in our case stadia VI, VII and/or VIII) to finish the development during the
next year, or this phenomenon is evoked under certain conditions. On the other hand M.
scutellare , probably also due to the shorter post-embryonic development with eight stadia, is
only an annual species (Fig. 5).

% IX (ad)

n =112

n =4

n = 36

n =24

n =49

VII

VIII

n =78

n =129

n = 185

IX (ad) J"

i = 272

F ' M ' A ' M '

J ' J ' A' S ' O' N* D

1992

1993

“IX*

wmmm

— 1 - 1 - 1 -

M. voigti

□

WZZ]

WA _

□

r . i mr

□ «

VII

o □ me

zzi

VIII

r*“: r”~

: i.J

wzzzzm.

IX (ad)

— _ j.

WZZZZZM "

— 1 - 1 - 1 - 1 - 1 -

1; ju pr ■ - 1 - 1 -

M. scutellare

1 - 1 izz

•v 1

ZZ) 0

v zz

EZJ 1

VII

1 _ 1

VIII (ad)

□

j F M A M J

J A S O N D

FiG. 4. — Post-embryonic development of M. voigti during the year. Filled fields: generation of previous year 1991.

Fig. 5. — Comparison of the life-cycles of M. voigti, M. gallica (according to David, 1984), and M. scutellare
(according to Blower, 1979). Areas with dotted borders - generation of previous year, cross-hatched areas -
according to Verhoeff (1928).

514

KAREL TAJOVSKY

In 1991 a severe rainfall deficiency between August and October was observed (Fig. 2).
At this time stadia VI. VII and VIII of M. voigti were present, which were then found again the
next spring 1992. These stadia were probably minimally active and/or quite inactive in winter,
because only adults were sampled. No convincing evidence about the development of this
stadium VIII into stadium IX was given by subsequent sampling. Either the members of this
stadium mature together with the new generation of stadium VIII or, only a small part of them
mature which is difficult to find by sampling, or they do not mature at all. Consequently the
absence of juvenile stadia VI, VII and/or VIII in spring 1993 is associated with favorable
climatic conditions during the year 1992, when the whole population probably completed its
development.

REFERENCES

Blower, J. G., 1978. — Anamorphosis in the Nematophora. Abb. Verb, naturwiss. Ver. Hamburg, (NF) , 21/22 : 97-
103.

Blower. J. G.. 1979. — The millipede faunas of two British limestone woods. In : M. Camatini, Myriapod Biology.
London, Academic Press : 295-306.

Blower, J. G., 1984. — The British Chordeumatidae. Bull. Br. Myriapod Group, 2 : 8-23.

David, J. F., 1984. — Le cycle annuel du Diplopode Microchordeuma gallica (Latzel, 1884). Bull. Soc. zool. Fr., 109 :
61 - 70.

David, J. F.. 1989. — Le cycle biologique de Chamaesoma brolemanni Ribaut & Verhoeff, 1913 (Diplopoda,
Craspedosomatida) en foret d’Orldans (France). Bull. Mus. natl. Hist, nat., Paris, 4e ser., II, section A, 3 : 639-
647.

Dunger, W. & Steinmetzger, K., 1981. — Okologische Untersuchungen an Diplopoden einer Rasen-Wald-Catena im
Thuringen Kalkgebiet. Zool Jb. Syst., 108 : 519-553.

Kempson, D., Lloyd, M., Ghelardi, R., 1963. — A new extractor for woodland litter. Pedobiologia, 3 : 1-21.

Meyer, E.. 1990. — Aititute-related changes of life histories of Chordcumatida in the Central Alps (Tyrol, Austria). In :

A. Minelli, Proceedings 7th Intern. Congr. Myriapodology. Leiden, E. J. Brill : 311-322.

Pedroli-Christen, A., 1978. — Contribution a la connaissance du developpement post-embryonnaire de Craspedosoma
alemannicum Verhoeff et de Xylophageuma zschokkei Bigler (Diplopoda, Nematophora) dans une tourbiere du Haut-
Jura Suisse. Rev. suisse Zool., 85 : 673-679.

Schubart O., 1957. — Die Diplopoden der Mark Brandenburg. Eine Okologische Studie. Mill. Zool. Mus. Berlin, 33 :
3-94.

Verhoeff K. W.. 1913. — Erscheinungszeiten und Erscheinungsweisen der reifen Tausendfussler Mitteleuropas und zur
Kenntnis der Gattungen Orobainosoma und Oxydactylon. Verb. Zool.-Bot. Ges. Wien, 63 : 334-381.

VERHOEFF, K. W., 1928. — Diplopoda 1. In : H. G. Bronn’s Klassen und Ordnungen des Tierreichs, 5, Leipzig,
Akademische Verlagsgesellschaft : 1-1072.

Source : MNHN, Paris

Life Cycles and Reproductive Strategies in Local
Populations of Rossiulus kessleri (Lohmander)
(Julidae, Diplopoda) from Isolated Habitats

Bella R . STRIGANOVA

Institute of Ecology and Evolution, Russian Academy of Sciences. Moscow, Russia

ABSTRACT

Peculiarities of the postembryonic development and reproductive parameters in separate populations of Rossiulus
kessleri Lohmander, 1926 were studied in two remote isolated forest habitats in the dry steppes of South Russia.
Observations were carried out in a natural river-plain forest and shelter forest plantation differing in the edaphic and
hydrothermal conditions. Seasonal rhythms of the development, duration of different stages, age and stage of the first
reproduction in females, fecundity and natural mortality were considered in both populations. Patterns of the life-cycle of
Rossiulus kessleri in secondary dry anthropogenic habitats seem to determine the following features of the reproductive
strategy: delay in first reproduction and decrease of a total egg-production, which is compensated by low mortality rates
in adult stages. Population differences in Rossiulus kessleri from isolated habitats are discussed in terms of K-selection
and expansion capacity.

RESUME

Comparaison des cycles de vie et des strategies de reproduction de populations locales de
Rossiulus kessleri (Lohmander) (Julidae, Diplopoda) dans des habitats isoles.

Les modalit£s du developpement post-embryonnaire et de la reproduction de populations separees de Rossiulus kessleri
Lohmander, ont 6te etudi6es dans deux habitats forestiers isoles des steppes seches du sud de la Russie. Les observations
ont ete effectives dans une foret naturelle de plaine et dans une plantation foresti&re protegee differant par leurs
conditions Sdaphiques et hydrothermiques. Les rythmes saisonniers du developpement, la duree des differents stades.
Page et le stade de la maturation sexuelle des femelles, la f£condite et la mortalite naturelle ont ete pris en compte dans
les deux populations. Les modalitSs du cycle de vie de Rossiulus kessleri dans des habitats secs anthropisSs semblent
determiner les paramfctrcs des strategies de reproduction - retard de la premiere reproduction et diminution de taille des
pontes - qui sont compens£s par un faible taux de mortalite des adultes. Les differences entre populations d’habitats
isoles sont discutees en terme de selection-K et d’augmentation de la capacite biotique.

INTRODUCTION

Reproductive parameters and peculiarities of the postembryonic development have been
studied in a number of diplopod species. These indices indicate variation in separate species and
in separate populations of the same species from different geographical regions. For example,
inter-population differences in the number of sexual and epimorphic stages of males have been
recorded in Bacillozonium nodulosam , Narceus annularis , Spelaeoglomeris doderoi (SAHLI,

Striganova, B. R.. 1996. — Life cycles and reproductive strategies in local populations of Rossiulus kessleri
(Lohmander) (Julidae, Diplopoda) from isolated habitats. In: Geoffroy, J.-J., MauriSs, J.-P. & Nguyen Duy -
JacQUEMIN, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat .. 169 : 515-522. Paris ISBN : 2-85653-502-X.

516

BELLA R. STRIGANOVA

1974). In different populations of Schizophyllum sabulosum the number of asexual stages
during anamorphic development was found to vary from 3 to 5 (HALKKA, 1958; SAHLI, 1968,
1969). Interpopulation differences seem to be especially strongly expressed in species with
polyzonal ranges, inhabiting sites with various edaphic and hydrothermic conditions.

The aim of this work was to compare the life-cycles and the reproductive indices in local
isolated populations of Rossiulus kessleri from a river floodplain forest and shelter forest strips
in the dry steppe subzone of Southern Russia.

Rossiulus kessleri is abundant in the forest-steppe and steppe zones of the European part
of Russia. It usually predominates in diplopod communities of both natural and anthropogenic
habitats (STRIGANOVA, 1977). These diplopods are well adapted to the climatic conditions of a
steppe landscape with a hot and dry summer season. Rossiulus kessleri is often located in plots
with tree and bush cover. Numerous aggregations of this species occured in ravine and river
plain forests and in wind protecting forest plantations (PRISHUTOVA, 1985; SlZOVA, 1985;
STRIGANOVA, 1972; STRIGANOVA & PRISHUTOVA, 1990).

Extensive ploughed fields with a long-term monoculture of wheat are characteristic of this
region of Russia. Plots with the natural tree and bush cover preserved and artificial forest strips
represent isolated island habitats. The natural exchange between these forest islands seems to be
impossible for non-flying invertebrates. The perennial isolation of separate populations of
diplopods in island habitats suggests the possibility of the existence of different
microevolutionary trends in populations living in diverse ecological situations.

SITES AND METHODS

Materials for this study were collected in Rostov and Stavropol regions in two sites being 30-40 km apart:

1. River flood-plain forest of one of the Don tributaries two species dominated: Salix alba, Alnus glutinosa. Soil
was grey sandy river-plain soil. Litter layer was a constant, 7-10 cm depth, grass cover under a closed tree canopy.

2. 30-year old shelter forest strip in a watershed position. Predominate tree species: Fraxinus excelsior, Robinia
pseudoacacia, Populus nigra. Soil - typical chernozem of the loamy texture. Litter layer was 23 cm depth, completely
decomposed by June. Grasses covered about 70 % of the ground under the canopy.

The habitats differed significantly in the hydrothermic regime of the upper soil horizons. Soil humidity in the
river-plain forest was conststantly high, due to the high level of ground water. A high relative humidity at aboveground
level was maintained by a dense grass cover. Dry leaf litter dried out only in mid-summer during a drought.

The forest plantation was situated in the zone of an unstable moisture. The litter and soil dried out in summer and
the relative humidity of the upper horizon ranged between 5-12 %.

Quantitative samples of diplopods were taken twice per month in both plots from April to October. Diplopods
were collected by a hand-sorting from standard quadrats (25 x 25 cm, depth 10 cm).

The live body mass, sex and developmental stage were determined in all specimens sampled. Mature females were
dissected to record the presence of eggs.

Periods of the spring moulting, oviposition, presence of juveniles and start of the winter diapause were recorded.
The post-embryonic development (longevity of separate stages, growth of the body mass and mortality) was studied in
the laboratory.

Eggs obtained from females kept into laboratory jars were placed in Petri dishes. Both longitudinal and
transversal diameters of eggs were measured to compare egg sizes from different populations. Larvae that hatched from
eggs in the laboratory were reared in groups in jars with soil and leaf litter taken from their natural habitats.

RESULTS

The spring moult of Rossiulus kessleri in the river-plain was observed in late April.
Diplopods moulted in the soil and did not build moulting chambers. Oviposition was completed
in May, and in the mid-June pupoids were recorded in experimental clutches.

In the forest plantation the spring moulting was delayed by 10-15 days. Almost the
diplopods built moulting chambers what was observed in both the natural habitat and
laboratory). Females began to oviposit in late June, and young larvae appeared in July-August.
During dry years the oviposition was delayed until autumn.

Source :

COMPARED LIFE-CYCLES AND REPRODUCTIVE STRATEGIES OF MILLIPEDE POPULATIONS

517

The stadium of 1st oviposition
differed in separate populations: X - in
the river-plain and XI - in the forest
plantation. Relative numbers of egg-
laying females increased in
progressively older stadia (Fig. 1). The
percentage of egg-laying females
increased in stages X-XII in the river-
plain from 56 to 100%. In the forest
plantation the percentage of egg-laying
females was much lower in all
reproductive stages and increased from
19% to 81%.

Mature females showed
differences in body mass and fecundity
(Figs 2, 3). An increase in the fecundity
of older stadia has been recorded in
other diplopod species - Cylindroiulus
latestriatus, C. punctatus, Julus
scandinavius (BAKER, 1978; BLOWER.
1970). The relationship between body
mass and fecundity has been described
in Ommatoiulus moreleti, Julus
scandinavius and Glomeris marginata
(Heath, Bocock & Mountford,
1974). The multiple correlation between
the developmental stage, body mass and
fecundity has been calculated for both
populations of R. kessleri under
study(PRISHUTOVA & MlNORANSKY,
1984). The correlation coefficients
averaged 0.91 - for the river-plain and
0.80 - for the forest plantation. Figure 3
shows that the smaller females of the
stage XII from the river-plain have a
higher fecundity than bigger females of
the same stage from the forest
plantation.

Fig. I. — Relative number of reproductive females (%) in the older
stadia of the development. 1: river-plain forest (n=98); 2:
wind protecting forest plantation (n= 1 1 3).

600.0

500.0
O)

(/) 400.0

(/)

(0
E

"g 300.0
-Q

200.0

100.0

Fig. 2. — Body mass of mature females (mg) in the older stadia of
the development. The bars represent one standart error of
the mean. 1: river-plain forest (n=98); 2: wind protecting
forest plantation (n= 1 1 3).

-i - 1 - 1 - r - 1-

X » XI XI XN

stadia

Table 1. — Total volume (mkl) of the egg production in females of different stages from separate populations of R
kessleri.

Stadia X XI XU XIII XIV

River-plain forest 34.969 37.587 58.157

Forest plantation - 61.446 67.683 76.923 104.874

518

BELLA R. STRIGANOVA

Fig. 3. — Fecundity vs. body mass in mature females. x±SE. 1 & 2
as in Figure 1 .

Egg sizes had clear interpopulation differences: river-plain: longitudinal diameter (mm)
0.826 ± 0.03 and transversal diameter (mm) 0.658 ± 0.03; forest plantation: longitudinal
diameter (mm) 0.860 ± 0.03 and transversal diameter (mm) 0.714 ± 0.01 (P = 0.001)

(PRISHUTOVA, 1985). The mean egg
volumes were calculated approximating
their form to an ellipsoid. Individual
mean volume of eggs from the river-
plain averaged 0. 1 87 mkl and that from
the forest plantation 0.231 mkl. Table 1
shows the total volume of egg
production in females of different stages
from both populations.

The relationship between the mean
mass of reproductive females and the
total volume of their egg-production is
shown by Figure 4. Correlation
coefficients between these indices
averaged 0.99 and 0.94 for the
population from the river-plain and
forest plantation respectively. The
correlation for the whole range of female
biomass values was not found to be
significant.

Duration of individual stadia was
different for each population. Figure 5.
shows the rates of the development for
stadia II-VII in the laboratory. The total
duration of these stages averaged 1 15
days in the river-plain population and
100 days in the forest plantation one.
Significant differences (P<0.05) were
observed in stadium VII only, and these
were apparently related to differences in
sex differentiation. Sex differentiation
begins in stadium VII in the river-plain
population, and stadium VIII in the
forest plantation population.

Rates of post-embryonic
development under favourable
laboratory conditions were evidently
higher than in natural habitats, as
revealed by parallel field samples. But
the growth of body mass progressed
slowly. Mass of individual diplopods in
natural habitats was 1.5-2 times as high

as that in laboratory specimens of the same stadia.

In the river plain, R. kessleri reached V-VIII during the first summer, IX-X the next year,
and the 1st reproduction took place during the third year. Individuals of stadium XIV were
recorded in this population. The minimum duration of the life-span is 4-5 years.

A mass emergence of young larvae in the forest plantation was observed in August. All
larvae built moulting chambers near the soil surface. 10-15 individuals were found in one

Fig.

4. — Relationship between the mean body mass of
reproductive females and the total volume of their egg
production. 1 & 2 as in Figure 1 .

Source : MNHN , Paris

COMPARED LIFE-CYCLES AND REPRODUCTIVE STRATEGIES OF MILLIPEDE POPULATIONS

519

chamber. Before a winter diapause they
reached III-IV, in the second year - VIII-
IX and in the third - X-XI. The first
reproduction took place in the fourth
year of life. This population had 15
post-embryonic stages. The total life¬
span is 5-6 years - a year longer than
that in the river-plain. During
unfavourable years, the number of egg-
laying females remained relatively low
and did not exceed 40% of the total
amount laid by mature females. This
seems to be the reason why the life-span
of this population can be prolonged to 8
years.

The mortality dynamics were
studied under laboratory conditions
(Fig. 6). In the river-plain, maximum
mortality rates were recorded in adult F|G
stages, which reproduced at least once.

In the forest plantation a significant
increase in natural mortality was
observed during the sexual
differentiation. These features, together
with the differences of seasonal rhythms
affect the population structure.

Figure 7 presents the population
structure in both habitats during two
subsequent years as stadia recorded
from soil samples. The bulk of the
population in the river-plain was
represented by reproductive stadia. The 3?
relative importance of the stadium IX
increased in autumn because of the |
recruitment from the younger =

generation. In different seasons all stadia
were present in the population. The
seasonal changes in the age composition
in both years were similar.

In the forest plantation the pre-
reproductive stadium X predominated in
the first spring. Older mature were
absent. In autumn, stages XI-XII
predominated and single specimens of
older stadia appeared. The following fig.
spring all adult stadia were present, but
VIII-X together barely exceeded 10%.
Their mass increased slightly by the
following autumn. Replacement rates of
generations were delayed by a year in the fore:
plain population.

days

5. — Development rates of stadia II-VII in the laboratory for
individuals of the river-plain population (1) and of the
forest plantation population (2).

stadia

6. — Mortality dynamics studied under laboratory conditions
for individuals of the river-plain population (1) and of the
forest plantation population (2).

plantation, in comparison with those in the river-

520

BELLA R. STRIGANOVA

stadia

Fig. 7. — Population structure during two subsequent years: river-plain population (1); forest plantation population (2)

Source : MNHN , Paris

COMPARED LIFE-CYCLES AND REPRODUCTIVE STRATEGIES OF MILLIPEDE POPULATIONS

521

DISCUSSION

The analysis of the population structure, the reproduction tactics and the development rates
of R. kessleri in isolated habitats revealed clear interpopulation differences in the life-cycle. A
long summer drought in the soil of an artificial forest plantation promotes a long aestivation
period for all stadia. The timing and duration of the summer pause was found to depend on
weather conditions in distinct years. The juvenile were particularly susceptible to the
hydrothermic conditions. They ceased to feed at the elevated soil temperature even under high
moisture levels, while adults continued the normal feeding activity (STRIGANOVA, 1972).

Diplopods living in dried out habitats have to accumulate energy reserves for a summer
aestivation, which minimizes the energy quota for reproduction. Accumulation of energy
reserves results in the increase in body mass in all stadia, which was observed in diplopods
from the forest plantation, in comparison with those from the river-plain (PRISHUTOVA, 1985).
The delay in first reproduction can be also interpreted in terms of the energy allocation within the
population.

In addition, diplopods from the forest plantation have significant energy expenses for the
building of moulting chambers. This building activity was recorded in all stadia. The activity of
larvae of the first stadia seems to be provided by a nutrient supply in the eggs. The eggs in these
populations were bigger than those from the river-plain populations.

Numerous julid species use to present typical K-selection features such as perennial life-
cycle, iteroparity, fixed seasonal rhythms, large body mass etc. (cf. PlANKA, 1970). All these
features are characteristics of Rossiulus kessleri.

Rossiulus kessleri has 7 asexual stadia. This is the maximum number recorded in Julidae
(SAHLI, 1969, 1974). Interpopulation differences in the duration of separate larval stages are
insignificant, depending more on the longevity of inactive periods, than on those of an active
growth.

The number of sexual stadia shows the inter-population differences (2 - in the river plain
and 3 - in the forest plantation). The same was shown in separate populations of C. teutonicus
(SAHLI, 1969).

Discussion of the factors determining the time of first reproduction considers two
alternatives - genetic determination of an age or of a stadium of first reproduction (DAVID,
1992). The results here are consistent with the idea of the stadia as the main determinant. Under
unfavorable conditions, for example during heat deficiency, diplopods reach their genetically
fixed stadium of maturity at an older age, as described in high mountains (MEYER, 1985).

Interpopulation differences in the stadium of first reproduction can be considered as a
result of the micro-evolutionary processes. Populations of R. kessleri under study differed in
both stage and age of the first reproduction. Age can vary between individuals within the same
population, for example in the forest plantation only 1 8% of females reaching stadium XI
participated in reproduction. Hence, age appears to be a more resilient population feature,
depending on local conditions.

An acceleration of maturation is associated with a restriction of life-span, as shown, for
example, in Nemasoma varicorne (BROOKES, 1974). Populations of R. kessleri showed the
same relationship. The shorter life-span was characteristic of diplopods from the more
favourable natural habitat (river plain forest), which can be considered the primary biotope for
R. kessleri in steppes. The artificial forest plantations represented the secondary habitats
populated by these diplopods. The prolongation of the life-cycle and the delay in first
reproduction seem to be adaptations to unfavourable anthropogenic habitats.

The prolongation of the life-span to account for long inactive periods is characteristic of
many poikilotherms under extreme conditions. K-selection features allow diplopods to select
these tactics. The development of this evolutionary trend was recorded only in stable populations
capable of increasing their accumulated reproduction reserve with aging, and of continuing an

522

BELLA R. STRICANOVA

active growth between reproductive cycles (STEARNS, 1976). These traits are characteristic of
R. kessleri. This is why this species is so widely distributed in dry steppes where it occurs in
both natural and anthropogenic habitats. Life-cycle traits are obligatory population features, they
were recorded under both field and laboratory conditions and can be considered as phenotypical
features.

REFERENCES

Baker, G. H., 1978. — The post-embryonic development and life-history of the millipede Ommatoiulus morelellii
(Diplopoda, Julidae) introduced in south-eastern Australia. J. zool. London , 186 : 209-228.

Blower, J. G., 1970. — The millipedes of Cheshire wood. J. zool. London , 160 : 455-496.

Brookes, C. H., 1974. — The life-cycle of Proteroiulus fuscus (Am Stein) and Isobates varicornis (Koch) with notes of
the anamorphosis of Blaniulidae. Symp. Zool. Soc. London, 32 : 485-501.

David, J. F., 1992. — Some questions about the evolution of life-history traits in Diplopoda. Ber. nat-med. Verein
Innsbruck, Suppl. 10 : 143-152.

Halkka, R., 1958. — Life-history of Schizophyllum sabulosum (L.), Diplopoda. Julidae. Ann. Zool. Soc. Zoologicae
Botanicae Fenn ., 19. 1-72

Heath. G. W., Bocock, K. L. & Mountford. M. D., 1974. — The life-history of the millipede Glomeris marginata
(Villers) in North-West England. Symp. Zool. Soc. London. 32 : 433-461.

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.

Pianka, E. R., 1970. — On r- and K-selection. Am. Nat., 104. : 592-598.

Prishutova. Z. G.. 1985. — Ecology of diplopods in southern steppes (on example of Rossiulus kessleri Lohmander).
Ph D Thesis. Moscow, Moscow State Pedagogical Institute, 21 pp. (in Russian).

Prishutova, Z. G. & Minoransky, V. A., 1984. — About the reproduction of diplopods Sarmatiulus kessleri Lohm. in
Nizhny-Don region. Abstracts All-Union Congress the USSR Entomol. Soc., Kiev. 2:114 (in Russian).

Sahli, F., 1968. — Observations sur la biologie el la periodomorphose chez le Diplopode Schizophyllum sabulosum
(L.) en Allemagne. Bull. Sci. Bourgogne . 25 : 333-346.

Sahli, F., 1969. — Contribution a l’etude du developpement post-embryonnaire des Diplopodes Julides. Ann. Univ.
saraviensis Reihe math.-naturw. Fak..l : 1-154.

Sahli, F.. 1974. — Sur les periodes larvaires asexuee et sexuee male et sur Tapparition des males adultes chez les
diplopodes Chilognathes. Bull. Soc. zool. Fr., 99 : 295-305.

Sizova. M. G., 1985. — Population structure of diplopods Rossiulus kessleri Lohmander in different habitats of Rostov
region. Abstracts IX Intern Colloq. Soil Zool., Vilnius : 264.

Stearns, S. C., 1976. — Life-history tactics: a review of ideas. Quart. Rev. Biol., 51 : 3-47.

Striganova, B. R., 1972. — Effects of temperature on the feeding activity of Sarmatiulus kessleri (Diplopoda). Oikos.
23 : 197-199.

Striganova, B. R., 1977. — Adaptations of diplopods to the life in soils with different hydrothermic conditions. In :
M. S. Ghilarov, Adaptation of soil animals to environmental conditions. Moscow. Nauka : 151-166. (in Russian,
English Summary)

Striganova. B. R. & Prishutova, Z. G., 1990. — Food requirements of diplopods in the dry steppe subzone of the
USSR. Pedobiologia, 34: 37-41.

Source : MNHN , Paris

Survival Strategy of the Terricolous Millipede
Cutervodesmus adisi Golovatch
(Fuhrmannodesmidae, Polydesmida)
in a Blackwater Inundation Forest of Central Amazonia
(Brazil) in Response to the Flood Pulse

Joachim ADIS*, Sergei I. GOLOVATCH ** & Susanne HAMANN *

* Max-Planck-Institute for Limnology, Tropical Ecology Working Group. Postfach 165, D-24302 Plon,
Germany, in cooperation with National Institute for Amazonian Research (INPA), CJP. 478, 69011-970

Manaus/ AM, Brazil

** Institute of Evolutionary Morphology and Ecology of Animals, Russian Academy of Sciences, 33 Leninsky

prospect, 117071 Moscow V-71, Russia

ABSTRACT

Reaction of Cutervodesmus adisi Golovatch, 1992 to 5-7 months of flooding was studied in 1976/77 and 1983/84 in a
blackwater inundation forest near Manaus, Brazil. The study area was annually covered by several metres of floodwater,
due to the monomodal flood pulse of the Rio Negro. Juvenile migratory stages with 18 (and 17) segments spent the
inundation period on tree trunks. In 1976/77, they represented 42% of all Polydesmida caught in arboreal traps on 6 tree
trunks over a period of 18 months (n = 5661). After flooding, the migratory stages of C. adisi recolonized the forest
floor. They developed to adults which subsequently reproduced. Vertical distribution of 94% of all animals extracted from
0-14 cm soil depth was restricted to the upper 7 cm. Migratory stages of the offspring moved into tree trunks. Trunk
ascents began several weeks before forest inundation and after the rainy season had started. Trunk ascents and descents,
as well as the vertical distribution of C. adisi in the soil during the non-inundation period, are discussed with respect to
abiotic factors in the study area (precipitation, insolation, temperature and humidity of soil and air) as well as
macroclimatic influences (El Nino events). C. adisi is considered to be an endemic species of the blackwater inundation
forests in the Rio Negro Valley. Vertical migration of its juvenile stages represents an ethological adaptation to escape
annual long-term flooding, which was not found in Polydesmida of neighbouring non-flooded upland forests.

RESUME

Strategic de survie du diplopode terricole Cutervodesmus adisi Golovatch (Fuhrmannodesmidae,
Polydesmida) dans une foret inondable de l’Amazonie centrale en reponse a la frequence des
inondations.

Les reactions de Cutervodesmus adisi Golovatch. 1992 i \ une periode de 5 a 7 mois d'inondation ont ete etudiees en
1976/77 et 1983/84 dans une foret inondable d'eau noire pr£s de Manaus au Bresil. Le site d’6tude est annuellement
recouvert par plusieurs metres d'eau, cons£cutivemement au flux d’inondation monomodal du Rio Negro. Les stades
juveniles migrateurs & 18 (et 17) anneaux passent la periode d'inondation dans les troncs d’arbres. En 1976/77, ils

Adis, J., Golovatch, S. I. & Hamann, S., 1996. — Survival strategy of the terricolous millipede Cutervodesmus
adisi Golovatch (Fuhrmannodesmidae, Polydesmida) in a blackwater inundation forest of Central Amazonia (Brazil) in
response to the flood pulse. In: Geoffroy, J.-J., MAURifes, J.-P. & Nguyen Duy - Jacquemin. M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist . nat., 169 : 523-532. Paris ISBN : 2-85653-502-X.

524

JOACHIM ADIS. SERGEI. I. GOLOVATCH & SUSANNE HAMANN

represeniaient 42% de tous les Polydesmida captures dans les pieges arboricoles releves sur six troncs d'arbres durant 18
mois (n = 5661). Apres Finondation, les stades migrateurs de C. adisi ont recolonisS le sol foreslier. Us se sont
developpes en adultes qui se sont ensuite rcproduits. La repartition verticale de 94% de tous les indi vidus extraits d’une
epaisseur de sol de 14 cm se reduit aux 7 cm superieurs. Les stades migrateurs issus du recrutement annuel se d<§placent sur
les troncs d'arbres. L’ascension du tronc commence plusieurs semaines avant Finondation de la foret et apr£s le
demarrage de la saison des pluies. L’ascension et la descente le long des troncs, de meme que la repartition verticale de C.
adisi dans le sol en dehors de la periode d*inondation, sont discutees en rapport avec les facteurs abiotiques du site delude
(precipitation, insolation, temperature, humiditc de Fair et du sol) ainsi qu’avec les influences macroclimatiques
(6venements dus au El Nino). C. adisi est consider comme une espece endemique des forets inondables de la vallee du Rio
Negro. La migration verticale de ses stades juveniles reprdsente une adaptation ethologique lui permettant d’6chapper &
une inondation a long terme, comportemcnt qu’on ne retrouve pas chez les polydesmides des forets non-inondees plus
elevees.

INTRODUCTION

Terrestrial invertebrates in periodically flooded ecosystems require special “survival
strategies’’ (cf. ADIS, 1992a). In Central Amazonia, the monomodal “flood pulse” (JUNK et al.,
1989) of the Rio Negro and the Rio Solimoes- Amazon causes flooding of forests near rivers -
the so-called seasonal inundation forests (PRANCE, 1979) - and their adjacent shores by several
metres of floodwater for 5-7 months each year. Terrestrial invertebrates have adapted to this
ecosystem. The fauna comprises terricolous and arboricolous animals. Both groups include non¬
migrants and migrants. Migratory reaction of terricolous animals is horizontal (following the
high water line), vertical (temporal ascent to trunk or canopy) or includes a temporal flight to
upland forests. Non-migrants have active or dormant stages under water. The latter pass
inundation in naturally available retreats, in self-made retreats or as eggs. Non-migrant
arboricolous animals reproduce and live exclusively in the trunk and canopy region, whereas
migrants include life stages that live on the ground as well. Characteristics and examples of
species for each of these categories are given by ADIS (1992a, b).

In the vicinity of Manaus, millipedes alone display a good variety of responses to seasonal
floods. Thus, Gonographis adisi Hoffman, 1985 (Pyrgodesmidae, Polydesmida) appears to be
unique in being the only hitherto known millipede capable of surviving submersion for up to 1 1
months due to a hydrophobic secretion layer on the cuticula which enables plastron respiration
(HOFFMAN, 1985; ADIS, 1986; MESSNER & ADIS, 1988). On the contrary the even more
widespread synanthropic Muyudesmus obliteratus Kraus, 1960 (Pyrgodesmidae), another forest
floor-dweller, escapes flooding by moving to the non-inundated tree trunks and canopy areas
(like most other millipedes); its plastron is incomplete thus long-term submersion is fatal (ADIS,
1986; MESSNER & ADIS, 1988). Mestosoma hylaeicum Jeekel, 1963 (Paradoxosomatidae,
Polydesmida) generally displays the same pattern of behaviour, with all the phases of its life-
history neatly corresponding to local seasonality (ADIS, 1992c). Probably the same holds true
for Prostemmiulus adisi Mauries, 1984 (Stemmiulidae, Stemmiulida), perhaps the only one of
three congeners (the others being P. amazonicus Mauries, 1984. and P. wellingtoni Mauries,
1984) encountered in inundation forests that shows similar seasonal vertical migrations from the
forest floor to the trunk/canopy areas and back (MAURIES, 1984). The same can obviously be
said about Moojenodesmus pumilus Schubart, 1944, M. susannae Golovatch, 1992
(Fuhrmannodesmidae, Polydesmida) (GOLOVATCH, 1992a) and Onciurosoma adisi Golovatch,
1992 (Paradoxosomatidae) (GOLOVATCH, 1992b).

Another different survival strategy is demonstrated by Epinannolene arborea Hoffman, 1984
(Pseudonannolenidae, Spirostreptida), obviously a strict arboricole (HOFFMAN, 1984; ADIS,
1984) which remains in the upper trunk and canopy region unless forced down the trunk by
insolation/drought when it retreats under the bark and estivates there.

In this paper, adaptive reaction of the diplopod Cutervodesmus adisi Golovatch, 1992 from a
seasonal blackwater inundation forest in the Rio Negro valley to the annual flooding is

Source :

SURVIVAL STRATEGY OF A TERRICOLOUS MILLIPEDE IN A BLACKWATER INUNDATION FOREST 525

discussed. Its survival strategy is compared with that of Diplopoda already known to inhabit
inundation forests in the surroundings of Manaus.

STUDY AREA AND METHODS

Diplopoda were collected between 1975 and 1988 in the course of ecological studies on terrestrial invertebrates from
Central Amazonian floodplains, in particular the seasonal inundation forests (cf. Adis, 1981, 1984, 1992 a-c- Adis &
Schubart, 1984).

The study site was situated on the lower course of the Rio Taruma Mirim (03°02’S, 60°17'W). a tributary of the Rio Negro,
about 20 km upstream from Manaus. The seasonal blackwater inundation forest (for definition see Prance, 1979) was
situated on a slope and extended from the non-inundated dryland area (= upland or terra firme) with a constant decline (<
5%) to the bare sandy shoreline of the Rio Taruma Mirim (see profile in Beck, 1976). The central part of the study site
was covered annually by up to 3.35 m of floodwater between March/April and August/September. Further information on
the study site is given by Adis (1981, 1984, 1992a), Meyer (1991) and Worbes (1986).

The activity density of Diplopoda evaluated in this study was monitored on the forest floor with 8 ground photo-
eclectors- of Funke (= emergence traps) during the non-inundation period in 1976/77. Trunk ascents and descents were
detected at weekly or bi-weekly intervals with arboreal photo-eclectors (= funnel traps) on three tree trunks each of the
dominant tree species (cf. Table 1) between December 1975 and May 1977. The killing/preserving agent used in all traps
was aqueous picric acid solution (without detergent), which is known to be mostly neutral in terms of attraction or
repellence in temperate zones (Adis, 1979). All capture devices arc fully described by Adis (1981) and Funke (1971), who
also explain their mode ol utilization and function. Trunk ascents of Diplopoda and their activity on the ground were
additionally monitored with funnel traps on one tree and with 1-4 emergence traps on the forest floor during the non¬
inundation periods in 1982/83 and 1983/84, respectively.

Distribution of Diplopoda in the non-flooded soil was studied between September. 1981 and' February, 1982. Once a
month, six soil samples were taken at random along a transect with a split corer (= steel cylinder with lateral hinges;
diameter 21 cm, length 33 cm) which was driven into the soil by a mallet. Each sample of 14 cm depth was then
subdivided into four subsamples of 3.5 cm each. Animals were extracted from subsamples following a modified method of
Kempson (Adis, 1987).

The presence of Diplopoda in the Hooded soil was studied at the end of each inundation period in 1984-88. Twelve soil
samples were taken at 3-weekly intervals under water as described above. Each of the subsamples was kept moist for JO-
14 days on a grid inside a bucket, which was covered by a cotton screen (sealed up by a plastic snap ring) and contained
aqueous picric acid in the bottom. Animals were subsequently extracted with the modified Kempson apparatus.

The presence of Diplopoda in tree crowns was tested by fogging canopies with pyrethrum in the early dry season (July
1977, 1979), when the inundation forest was completely Hooded (cf. Adis et ai 1984; Erwin, 1983). Bromeliads. 5-25
m above ground, were also sampled and checked for terrestrial invertebrates in August, 1979 (forest not Hooded) and
June, 1981 (forest Hooded).

Seasonal inundation lorests in Central Amazonia are subject to a rainy season (December - May: average precipitation
1550 mm), and a "dry" season (June - November: average precipitation 550 mm, but each month has some rain events;
cf. Ribeiro & Adis, 1984). Vertical distribution of Diplopoda in relation to changing conditions of soil moisture
content, temperature and pH, was statistically evaluated with the linear correlation test (Cavalli-Sforza, 1972), using
the original field data. This method was also used to evaluate the activity of Diplopoda on the soil surface and tree trunks
in relation to insolation, precipitation, temperature and humidity of the air.

The taxonomic work for this paper was done by S. I. Golovatch (cf. Golov atch. 1992a, b), the evaluation of field data
by J. Adis and S. Hamann. Diplopoda sampled were classified as juveniles (7, 9, 12, 15, 17 and 18 pairs of legs),
subadults (19 pairs of legs) and adults (20 pairs of legs) according to Schubart (1934). Sex was determined in the adults.

RESULTS AND DISCUSSION

About 2600 specimens of Cutervodesmus adisi Golovatch, 1992 were collected in the
seasonal blackwater inundation forest under study. Of these, 98% could be grouped into
developmental stages. The majority were juveniles (97.1% of the total catch; n = 2420), 1.7%
were subadults (n = 43) and 1 .2% adults (n = 29).

C. adisi represented 60.4% (329 ind. m-2 month-i) of all Diplopoda (545 ind. m-2 month-i)
extracted from soil samples during the non-inundation period 1981/82. A total of 7.2 ± 4.0
millipedes per m2 (1.4 ind. m-2 month-i) were collected on the soil surface during the non¬
inundation period in 1976/77. Animals occurred solely from January to April (rainy season) in
ground photo-eclectors and were not detected during the preceding dry season. Out of these, C.
adisi represented 4.5 ±0.7 ind. m-2 (= 62.5 %; 0.9 ind. m-2 month- 1). The majority of the total
diplopods obtained in 1976/77 was sampled on tree trunks (cf. Table 5 in ADIS, 1981): out of

526

JOACHIM ADIS. SERGEI. I. GOLOV ATCH & SUSANNE HAMANN

the 13516 specimens caught, 17.5% (n = 2369) were represented by C. adisi. It was the
eudominant species of all Polydesmida collected (n = 5661) and represented 4 1 .9% of their total
catch. The percentage of trunk ascending individuals of C. adisi was higher (28.2% of the total
Diplopoda and 66.7% of all Polydesmida caught), when compared to trunk descending animals
(15.1% of the total Diplopoda and 36.1% of all Polydesmida sampled).

C. adisi reaches 6.0 mm in length (cf. GOLOV ATCH, 1992a). The species is considered
hemiedaphic, as 94% of all specimens extracted from soil samples in 1981/82 were found in the
top 7 cm (Fig. 1), independent of seasons. This becomes evident, when significant correlations
between the abundance of C. adisi and different soil conditions in the study area are carefully
analysed.

Fig. 1. — Distribution of
developmental stages of
Cutervodesmus adisi in the
soil at Rio Taruma Mirim.
Monthly samples taken
every 3.5 cm to a depth of
14 cm between September,
1981 and February, 1982
(non-inundation period);
total catch = 100%.

■■ juvenile subadult _ adult

During the dry season the decreasing abundance with greater soil depth was positively
correlated with the increasing soil moisture content (e.g. Nov. 19, 1981: p < 0.01, r = +0.9924
for the total catch and p < 0.05, r = +0.9798 for juveniles; n = 4, respectively). However,
during the rainy season, particularly after heavy rainfalls, the decreasing abundance was
negatively correlated with the now decreasing soil moisture content in greater soil depth (e.g.
Dec. 17, 1981: p < 0.001, r = -0.9995 for juveniles; n = 4). Similar changes were found with
regard to soil temperatures: an increase with greater soil depth after heavy rainfalls was
negatively correlated with the decreasing abundance of C. adisi (p < 0.05), whereas a
temperature decrease with greater soil depth during dryer periods was positively correlated with
the decreasing abundance of the species (p < 0.05).

Grain size and mineral composition of the soil seem to be especially important for the
vertical distribution of terricolous arthropods in seasonal inundation forests (cf. ADIS et al., this
volume). First analyses of soil data (ADIS & IRION, unpubl.) indicated, that the decreasing
abundance of C. adisi with greater soil depth during the dry and rainy seasons corresponded
with an increasing amount of grains > 1000 |im in lower soil layers (from 0.7% in 0-3.5 cm to
2.7% in 10.5-14 cm depth; p < 0.05) and with a decreasing amount of silt from 14.6% in the top
3.5 cm to 10.8% in 14 cm soil depth (p < 0.05).

During the rainy season the decreasing abundance of C. adisi was negatively correlated
with the increasing pH in lower soil layers as well (p < 0.05).

About 85% of the C. adisi population extracted from soil samples in 1981/82 was
represented by juveniles, 7% by subadults and 8% by adults (Fig. 2). The sex ratio of males and
females was 1:1.5 (n = 27). Adults and early larval stages (7, 9 & 12 segments) occurred solely
in the soil (Fig. 3) where reproduction must have taken place.

Source :

SURVIVAL STRATEGY OF A TERRJCOLOUS MILLIPEDE IN A BLACKWATER INUNDATION FOREST 527

Fig. 2. — Percentage of

developmental stages of C.
adisi caught in the soil (0-
14 cm depth). Monthly
samples taken between
September, 1981 and
February, 1982 (non¬
inundation period) at Rio
Taruma Mirim.

Fig. 3. — Temporal occurrence and
abundance of C. adisi
(ind./m2) in the soil (0-14
cm depth). Monthly
samples taken between
September, 1981 and
February, 1982 (non
-inundation period) at Rio
Taruma Mirim.

21.09.81 19.10.81 19.11.81 17.12.81 21.01.82 17.02.82

mM 7s

d] 178

f 98

CZD 18a

8 • number of segments

12s

19s (subad.)

15s

20s (ad.)

Advanced juvenile stages (the majority with 18 segments, some with 17 segments; cf.
Table 1) were found to pass the inundation period in the trunk region (Figs 4, 5) and to
recolonize the forest floor after the floodwater had receded. In 1981, some animals had already
moulted to adults and reproduced within the first five weeks after the forest floor had dried, as
juvenile stages of the offspring occurred from mid-September onwards (Fig. 3). Abundance of
C. adisi in the soil was highest during the dry season (October 19, 1981: 678 ind. m-2). The first
larval stage of the progeny (7 segments, 3 pairs of legs) was obtained at this time only and is
believed to be of short duration (cf. HOPKIN & READ, 1992). In 1981/82, 49% of all C. adisi
specimens extracted from soil samples comprised advanced juvenile stages with 18 segments
and to a lesser extent with 17 segments (Fig. 2). They represented “migratory stages” which
came to the soil surface at the beginning rainy season and started ascending tree trunks (Fig. 4).

Source :

528

JOACHIM ADIS, SERGEI. I. GOLOVATCH & SUSANNE HAMANN

In 1976. they were caught in ground photo-eclectors from January onwards, and activity density
became' greater with an increasing water saturation of the soil during subsequent weeks (p <
0 05, r = +0.521; n = 16; Fig. 4). Shortly before forest inundation, abundance of C. adisi in the
soil was lowest (131 ind. m-2; Fig. 3). The juveniles of non-migrating stages remaining in the
soil were forced into the trunk area by the inundation of the forest (Table 1). Similar behaviour
was found in two species of pseudoscorpions, and where tritonymphs represented the migratory
stage (ADIS & MAHNERT, 1985; ADIS et al ., 1988).

Table 1 — Number (ind.) and dominance (%) of developmental stages of Cutervodesmus adisi caught during trunk ascents
(BET) and trunk descents (BET) on three different tree species of Leguminosae in different years (capture periods:
BET from December to May in 1975/76. 1976/77. 1982/83 & 1983/84; BEi from July to October, 1976). No
specimens were captured on Mora paraensis DUCKE (Caesalpiniaceac) between July and October. 1976. N.I. = No
Investigation.

Trunk ascent (BET)

Stage

1975/1976

1976/1977

1982/1983

1983/1984

Total

Aldina latif olia

Benth var. latifolia I
(Fabaceae)

1.1

8.4

75.8

14.7

Total

100.0

Aldina latifolia

N.I.

2.7

Benth var. latifolia II

476

505

97.3

(Fabaceae)

Total

490

519

100.0

Peltogyne venosa

N.I.

1.2

Benth ss.

densiflora (Benth) M.

138

163

98.8

Silva

(Caesalpimaceae)

Total

139

165

100.0

Trunk descent

Stage

1976

SUM

Trunk

Total

(BEi)

ascent

descent

Aldina latifolia
Benth var.
latifolia

252

23.1

Stage

837

76.9

0.1

—

0.1

III (Fabaceae)

Total

1089

100.0

3.1

273

20.5

297

14.1

Aldina latifolia

8.7

740

95.0

1057

79.5

1797

85.2

Benth var.
latifolia

IV (Fabaceae)

220

91.3

1.8

0.7

Total

241

100.0

Total

779

100.0

1330

100.0

2109

100.0

In C. adisi the number of migratory juveniles caught in arboreal photo-eclectors was
higher during periods of less insolation (February-May 1977: p < 0.01, r = -0.9914, n = 4; cf.
Fig. 4). This was also reported for trunk ascending adults of Hanseniella arborea, a migrating
symphylan in seasonal mixed- and blackwater inundation forests (ADIS et al., this volume).
During rainy seasons which showed an approximately equal amount of precipitation between
months and a steady increase of flood waters in the study area, trunk ascents of C. adisi
occurred over a long period of time, e.g. from December to April in 1983/84. They were
relatively short in duration during rainy seasons which were marked by a greater amount of
rainfall in certain months and by fast rising flood waters, e.g. in February /March 1976 and in
April/May 1983. These two years were characterized by macroclimatic El Nino-Southern
Oscillation (ENSO) events (strong in 1982/83 and weaker in 1976/77), which were statistically
shown to cause a decrease in total precipitation during the rainy season in Central Amazonia and
a lower average water-level of the Rio Negro seven months after the event had begun (ADIS &
Latif, 1995; cf. ROPELEWSKI & HALPERT, 1987; PHILANDER, 1983; RICHEY et al., 1989).

Source : MNHN, Paris

SURVIVAL STRATEGY OF A TERRICOLOUS MILLIPEDE IN A BLACKWATER INUNDATION FOREST 529

From December 1982 until May 1983 (Fig. 5) the trunk ascent in C. adisi was significantly
correlated with the rising water gauge of the Rio Negro at Manaus (p< 0.05, r= +0.8380; n= 6).

C. adisi is believed to be nocturnal as it
was found to pass the flood period aggregated
under loose bark of trunks during the day. The
number of specimens caught on individual
trees of the same and of different tree species
varied considerably (cf. Table 1) and tree
trunks with smooth and/or thin bark may be
avoided (e.g. Mora paraensis). In 1977, trunk
descents were observed to occur within 1-2
weeks and at the time when the forest floor
was about to emerge from the receding
floodwater (cf. Fig. 4).

C. adisi was neither detected in tree
crowns (by means of canopy fogging) nor in
epiphytes (5-25 m above ground), nor was it
found in flooded soils which were taken from
under the water during forest inundation.
Adults and early juvenile stages occurred
solely on the forest floor. Based on the
characteristics outlined. C. adisi represents a
terricolous, migrating and univoltine species,
which is considered endemic of seasonal
blackwater inundation forests in the Rio Negro
valley. Vertical migration of advanced juvenile
stages represents an ethological adaptation to
escape annual long-term flooding, which was
not found in Polydesmida and other terrestrial
invertebrates of neighbouring non-flooded
upland forests (cf. ADIS, 1992a;
GOLOV ATCH, 1992a, b).

Fig. 4. — Activity density of Cutervodesmus adisi on the
forest floor (8 ground photo-eclectors (E);
ind./8m2), trunk descents (BEi)and trunk ascents
(BET); three arboreal photo-eclectors, respectively
between July, 1976 and May, 1977 at Rio Taruma
Mirfm. Soil moisture content is expressed as the
percentage portion of the soil humidity (vol. %) at
the maximal water capacity of the soil (= 34 vol.
%; cf. ADIS, 1981).

300

(hrs.)

(%)

Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May
;—3 insolation (hrs.) * soil moisture content (%)

BEt (ind.)

N - 633

0- -

300

600

trunk ascents

trunk descents

BE* (ind.)

N - 1333

Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May

E (ind. /8m2)

N - 36

activity density
-forest floor-

nail

Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May
1976 1977

18 a

17 •

20 a (adult!

Inundation period

GEO 19 a (aubad.)

non-inundation period

530

JOACHIM ADIS, SERGEI. I. GOLOV ATCH & SUSANNE HAMANN

600

400

200

Dec. Jan. Feb. Mar. Apr.

May

600

400

200

1976/77

Dec. Jan. Feb. Mar. Apr. May

* precipitation

water-level study area flooded

Feb. Mar. Apr.

• number of segments

May

* precipitation : water-level study area flooded

□ 17 s Hie. 8 • "umber of segments

600

1982/83

400

200

Dec.

Apr.

May

200

600

1983/84

400

Feb. Mar. Apr. May

precipitation

water-level study area flooded

100

■ '5 s 17 s HD 16 s ( _ 1 19 s

8 • number of segments

precipitation water-level study area flooded

(ZD 17 s HJ 18 s (ZD 19 s

s • number of segments

Fig. 5. Percentage of developmental stages of Cutervodesmus adisi ascending trunks caught per month in arboreal
photo-eclectors during four rainy seasons in relation to water-level of the Rio Negro and precipitation in the
Manaus area; total catch per season = 100 %. (See left page).

Source : MNHN, Paris

SURVIVAL STRATEGY OF A TERRICOLOUS MILLIPEDE IN A BLACKWATER INUNDATION FOREST 53 1

The flood pulse is regarded as the original determinant of the upward and downward
migrations of terrestrial invertebrates on tree trunks in the seasonal inundation forests of Central
Amazonia (ADIS, 1992a). However, it is still the primary control mechanism or ecofactor only
among certain species. Most of the invertebrates, like C. adisi, have apparently become sensitive
to secondary, mainly abiotic ecofactors, which are no longer directly related to the cycle of
flooding. The migration of animals from the ground to tree trunks (and their flight to upland
forests) is triggered mainly by the rainy season (December - May), which begins three to four
months before the flooding, and by the changes in the edaphic and climatic factor it causes.

Following GOLOV ATCH's (1987) proposed division of the Diplopoda into morphotypes
and ecomorphotypes (= life-forms), C. adisi represents a good climber: its body is fairly small
(up to 6 mm in length with females being larger (5.0-6.0 mm) than males (4.5-5. 3 mm)), the
paraterga are relatively small, the legs relatively strong and not too slender, with the somewhat
pretarsal podomeres and more slender and long tarsi each crowned with a well-developed claw.
Such a morphological pattern also fits a soil dweller (= edaphobiont), which is also the case. The
high number of migratory stages (which represented almost half the population in the soil), the
probable predominance of females on the forest floor and the synchronization of reproduction in
the soil with the non-inundation period compensates for the decline in population density during
flooding and assures the persistence of this species in a harsh environment.

ACKNOWLEDGEMENTS

We arc indebted to all our colleagues at the National Institute of Amazonian Research (INPA) in Manaus (Brazil) and at
the Max-Planck-Institute for Limnology in Plon (Germany) who, in the field or laboratory, contributed to this study,
especially Edilson De Araujo Silva, M. Sc. Elizabeth Franklin, M. Sc. Jose Wellington De Morals and Irmgard Adis.
This study was supported by a grant from the Max-Planck-Society for the second author. We wish to acknowledge the
valuable support received by PD Dr. W. J. Junk, head of the Tropical Ecology Working Group at the Max-Planck-
Institute for Limnology in Plon, Germany. Dr. Helen Read (Bucks, United Kingdom) and Dr. Ulf Scheller (Jarp&s,
Sweden) kindly corrected the English manuscript.

REFERENCES

Adis, J., 1979. — Problems of interpreting arthropod sampling with pitfall traps. Zooi Anz.. 202 : 177-184.

Adis, J., 1981. — Comparative ecological studies of the terrestrial arthropod fauna in Central Amazonian inundation
forests. Amazotiiana, 7 : 87-173.

Adis, J., 1984. — ’Seasonal Igapo’-forests of Central Amazonian blackwater rivers and their terrestrial arthropod fauna.
In: H. SlOLl, The Amazon. Limnology and landscape ecology of a might y tropical river and its basin. Monographiae
Biologicae, Dordrecht, W. Junk Publ. : 245-268.

Adis, J., 1986. — An “aquatic” millipede from a Central Amazonian inundation forest. Oecologia, 68 : 347-349.

Adis, J., 1987. - Extraction of arthropods from Neotropical soils with a modified Kempson apparatus. J. trop. Ecol. , 3 :
131-138.

Adis, J., 1992a. — Uberlebensstrategien terrestrischer Invertebraten in Uberschwemmungswaldern Zentralamazoniens.
Verb, naturwiss. Ver. Hamburg, 33 : 21-114.

Adis, J., 1992b. — How to survive six months in a flooded soil : Strategies in Chilopoda and Symphyla from Central
Amazonian floodplains. In: : J. ADIS & S. Tanaka, Symposium on life-history traits in tropical invertebrates.
[INTECOL, Yokohama. Japan 1990. Studies on Neotropical Fauna and Environment, 27]. Lisse, Swets &
Zeitlingcr : 117-129.

Adis, J., 1992c. — On the survival strategy of Mestosoma hylaeicum Jeekel (Paradoxosomatidae. Polydesmida,
Diplopoda), a millipede from Central Amazonian floodplains. Ber. nat.-med. Ver. Innsbruck. Suppl. 10 : 183-187.
Adis, J. & Latif, M., 1995. — Amazonian arthropods respond to El Nino. Biotropica : (in press).

Adis, J. & Mahnert, V., 1985. — On the natural history and ecology of Pseudoscorpiones (Arachnida) from an
Amazonian blackwater inundation forest. Amazoniana. 9 : 297-314.

Adis, J. & Schubart. H. O. R., 1984. — Ecological research on arthropods in Central Amazonian forest ecosystems
with recommendations for study procedures. In: J. H. COOLEY & F. B. GOLLEY, Trends in ecological research for the
1980s. (NATO Conference Series, Series 1 : Ecology] New York, London. Plenum Press : 111-144.

Adis, J., Lubin, Y. D. & Montgomery, G.G., 1984. — Arthropods from the canopy of inundated and terra firme forests
near Manaus, Brazil, with critical considerations on the pyrethrum-fogging technique. Studies on Neotropical Fauna
and Environment, 19 : 223-236.

532

JOACHIM ADIS. SERGEI. I. GOLOVATCH & SUSANNE HAMANN

Adis. J., Mahnert, V.. Morais, J. W. De & Rodrigues, J. M. G.. 1988. — Adaptation of an Amazonian pseudoscorpion
(Arachnida) from dryland forests to inundation forests. Ecology, 69 : 287-291.

Beck. L., 1976. — Zum Massenwechsel der Makro-Arthropodenfauna des Bodens in Uberschwemmungswaldern des
zentralen Amazonasgebietes. Amazoniana, 6 : 1-20.

Cavalli-Sforza, L., 1972. — Grundziige biologisch-medizinischer Stcitistik. Stuttgart. G. Fischer. 212 pp.

Erwin, T. L., 1983. — Beetles and other insects of tropical forest canopies at Manaus. Brazil, sampled by insecticidal
fogging. X. hi. S. L. Sutton, T. C. Whitmore & A. C. Chadwick, Tropical Rain Forest: Ecology and Management.
Oxford, Blackwell Scientific Publications : 59-75.

Funke. W.. 1971. — Food and energy turnover of leaf-eating insects and their influence on primary production.
Ecological Studies, 2 : 81-93.

GOLOVATCH. S. I., 1987. — On life-forms in millipedes (Diplopoda). In: B. R. Striganova. Soil fauna and soil fertility ;.
Moscow, Nauka : 210-213.

Golovatch, S. I., 1992a. — Review of the Neotropical fauna of the millipede family Fuhrmannodesmidae, with the
description of four new species from near Manaus, Central Amazonia, Brazil (Diplopoda, Polydesmida).
Amazoniana, 12 : 207-226.

Golovatch, S. I., 1992b. — Review of the Neotropical millipede genus Onciurosoma Silvestri, 1932, with the
description of three new species from near Manaus, Central Amazonia, Brazil (Diplopoda, Polydesmida,
Paradoxosomatidae). Amazoniana, 12 : 227-237.

Hoffman, R. L., 1984. — A new species of Epinannolene from the Amazon Basin, Brazil (Spirostreptida:
Pseudonannolenidae). Myriapodologica , 1 : 91-94.

Hoffman. R. L., 1985. — A new milliped of the genus Gonographis from an inundation forest near Manaus, Brazil
(Pyrgodesmidae). Amazoniana. 9 : 243-246.

Hopkin. S. P. & Read. H. J., 1992. — The biology of millipedes. Oxford. Oxford Univ. Press, 233 pp.

Junk, W. J.. Bayley, P. B. & Sparks. R. E.. 1989. — The flood pulse concept in river-floodplain systems. Can. Spec.
Publ. Fish. Aquatic. Sci., 106 : 110-127.

Mauries, J.-P.. 1984. — Les premiers Stemmiulides signales au Bresil : trois especes nouvelles de la region de Manaus,
dont une de la foret inondable ( Prostemmiulus adisi n. sp.)( Myriapoda : Diplopoda : Stemmiulida). Amazoniana, 8 :
375-387.

Messner, B. & Adis, J., 1988. — Die Plastronstrukturen der bisher einzigen submers lebenden Diplopodenart
Gonographis adisi Hoffman 1985 (Pyrgodesmidae, Diplopoda). Zool. Jb. Anat., 117 : 277-290.

Meyer, U., 1991. — Feinwurzelsysteme und Mykorrhizatypen als Anpassungsmechanismen in zentralamazonischen
Uberschwemmungswaldern - Igapo und Vdrzea. Univ. Hohenheim. FR Germany: Ph.D.-thesis, 223 pp.

Philander, S. G. H., 1983. — El Nino Southern Oscillation phenomena. Nature, 302 : 295-301.

Prance, G. T.. 1979. — Notes on the vegetation of Amazonia III. The terminology of Amazonian forest types subject to
inundation. Brittonia, 31 : 26-38.

Ribeiro, M. De N. G. & Adis. J., 1984. — Local rainfall variability - a potential bias for bioecological studies in the
Central Amazon. Acta Amazonica, 14 : 159-174.

Richey, J. E., Nobre, C. & Deser, C., 1989. — Amazon River discharge and climate variability: 1903-1985. Science,
246 : 101-103.

Ropelewski, C. F. & Halpert, M. S.. 1987. — Global and regional scale precipitation patterns associated with the El
Nino/Southern Oscillation. Mon. Wea. Rev., 115 : 1606-1626.

Schubart, O.. 1934. — Tausendfussler Oder Myriapoda I: Diplopoda. In: F. Dahl, Tierw. Deutschl. 28. Jena. G.
Fischer, 1-318.

Worbes, M., 1986. — Lebensbedingungen und Holzwachstum in zentralamazonischen Uberschwemmungswaldern.
Scripta Geobotanica, 17 : 1-112.

Source : MNHN. Paris

Cycles d'activite compares de populations
de diplopodes edaphiques dans un ecosysteme

forestier tempere

Jean- Jacques GEOFFROY * & Marie-Louise CELERIER **

* CNRS. Museum National d'Histoirc Naturelle, IEGB, Laboratoire d'Ecologie Generate
4, avenue du Petit Chateau F-91800 Brunoy, France
** Universite Pierre & Marie Curie, BoTte 6, UFR Sciences de la Vie, Bat. A
4, place Jussieu F-75252 Paris Cedex 05, France

RESUME

Le cycle d'activite de cinq cspeces du peuplement de diplopodes edaphiques d’un ecosysteme forestier du Bassin Parisien
(Station Biologique de Foljuif, Seine-et-Marne, France) a ele suivi & l'aide de pieges d'interception durant plusieurs
annees. Cette etude permet de determiner I’importance relative des fractions adultes et juveniles des populations dans les
couches superieures de litiere (couche L et couche F) au cours des diverses periodes caracterisant le cycle annuel. Outre la
mise en evidence des phases successives de colonisation des nouvelles couches de litiere par des populations
fonctionnellement compiementaires. cette approche, qui prend en compte les variations interannuelles, conduit a
seiectionner les categories d'individus qui interviennent de maniere fondamentale dans 1'organisation d'un groupe
fonctionnel de “macrodiplopodes saprophages”.

ABSTRACT

Compared Activity Cycles of Millipede Populations in a Temperate Woodland Ecosystem.

The activity cycle of five species mainly representative of the edaphic millipede community have been investigated in
a temperate woodland ecosystem belonging to the Fontainebleau Forest (Foljuif Biological Station, Seinc-et-Mame,
France). The research has been conducted using Barber Pitfall Traps (PT) during several successive years. This study
allows to estimate the relative importance of the adult and juvenile fractions of specific populations acting in the upper
litter layers of the Oak (Quercus petraea) and Hornbeam ( Carpinus betulus) forest (L layer and F layer). The situation has
been examined during the characteristic successive periods of the annual cycle. This investigation contributes to
distinguish obvious steps in the colonization of recently fallen leaves by millipede populations and individuals,
showing complementary ecological functions and migrating activity in the litter layers. Considering interannual
variations, it is possible to select individuals that play an important role in the composition and organization of a
“functionnal saprophagous macro-diplopod group”. The investigated species are two short-cycle ones : Melogona
gallica (Latzel) and Polydesmus angustus Latzel, and three long-lived ones: Glomeris mcirginata (Villers), Cylindroiulus
punctalus (Leach) and Allajulus nitidus (Verhoeff).

Geoffroy, J.-J. & Celerier, M.-L., 1996. — Cycles d'activitE compares de populations de diplopodes Edaphiques
dans un EcosystEme forestier tempErE. In: Geoffroy, J.-J., Mauri£s, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist. nat.% 169 : 533-554. Paris ISBN : 2-85653-502-X.

534

JEAN-JACQUES GEOFFROY & MARIE-LOUISE CELERJER

INTRODUCTION

De nombreux types de pieges, de conception et de forme variees, ont ete employes pour la
capture qualitative et semi-quantitative des invertebres (PARK, 1935 ; JOOSE & KAPTEIJN,
1968 ; LE BERRE, 1969 ; SOUTHWOOD, 1978 ; DRACH et al. , 1981 ; PANTIS et al., 1988).
Dans le cas des Macroarthropodes terrestres de foret temperee, on utilise le plus souvenl des
pieges d'interception constitues de pots enfonces dans le sol, derives de la technique mise au
point par BARBER (1931) pour 1'etude d'insectes cavernicoles (DRIFT, 1951 ; GIST &
CROSSLEY, 1973 ; LUFF, 1975). On fait largement usage des pieges d'interception dans 1'etude
d'animaux particulierement mobiles tels que les coleopteres (BRIGGS, 1961 ; MURDOCH,
1966; MADER & Muhlenberg, 1981 ; Franke & FRIEBE, 1983 ; Friebe, 1983 ;
LECORDIER & BENEST, 1986 ; Rusdea, 1992). De nombreux auteurs ont ete amenes a
interpreter les cycles d'activite de diplopodes a l'aide de cette technique bien adaptee a des
animaux se deplagant entre les interstices que forment les constituants organiques des litieres
forestieres et capables de migrations actives entre les differents compartiments edaphiques
(Tableau 1). De faqon generate, la technique des pots-pieges enfonces renseigne sur les periodes
d'activite des populations envisagees a l'echelle de la journee (PARK et al. , 1931 : PARK,
1935 ; Williams, 1958. 1959 ; Banerjee, 1967a ; Dondale et al.; 1972) ou au cours des
phases saisonnieres lors d'etudes portant au minimum sur un cycle annuel (GILBERT, 1956 ;
Greenslade, 1964 ; MURDOCH. 1966 ; Banerjee, 1967a, b).

Les etudes par piegeage ne component un aspect veritablement quantitatif que dans
quelques experiences particulieres (BANERJEE, 1970 ; GIST & CROSSLEY, 1973). Elies
cherchent a montrer l'existence de relations entre l'activite locomotrice des individus et la densite
des populations durant la periode de capture (MITCHELL, 1963 ; GREENSLADE, 1964 ;
BLOWER, 1970 ; ADIS, 1979 ; David & POUSSARDIN, 1983). Divers auteurs s'accordent a
placer ce type de relations dans le cadre du concept d'activite-densite (KACZMAREK, 1978) qui
traduirait la capacite moyenne des individus a effectuer des deplacements (BANERJEE, 1970 ;
DAVID, 1983 ; COURET, 1985 ; FRANKE et al., 1988). Si des donnees semi-quantitatives liees
a l'abondance des populations sont ainsi acquises, elles ne portent cependant que sur une
fraction de celles-ci, celle qui est capable de se deplacer horizontalement et entre les couches
constituant la litiere. Les larves de premiers stades, sedentaires et rarement representees dans les
couches superieures de litiere, de me me que les individus en cours de mue, echappent en grande
partie a ce type de mesure. En revanche, les formes subadultes et adultes y sont bien adaptees.

Pour cette raison, la technique presente un interet supplementaire, en contribuant a separer
deux groupes d'individus jouant des roles differents a l'interieur d'un ensemble
taxinomiquement homogene. De plus, associe a d'autres methodes de capture, le piegeage
d'interception permet de localiser dans le temps certains phenomenes lies a la biologie des
adultes reproducteurs (Blower, 1978 ; David & POUSSARDIN, 1983 ; David, 1984).

Sans perdre de vue les limites imposees par l'emploi des pieges d'interception dans les
programmes de recherche en ecologie des peuplements (TOPPING & SUNDERLAND, 1992), le
propos de ce travail est de decrire les cycles d'activite des principales especes de diplopodes d'un
sol forestier, cherchant a determiner, pour chaque saison, quelle fraction du peuplement
frequente majoritairement les couches les plus superficielles de la litiere (L+F). Le piegeage
permet en outre de situer dans le temps les phases de mobilite des adultes males et femelles liees
a des activites de reproduction, de consommation ou de migration (DRIFT, 1951 ; BARLOW,
1958 ; Blower, 1970 ; COTTON & Miller, 1974 ; David & POUSSARDIN, 1983). Enfin, a
condition d'etendre l'operation sur plusieurs annees, cette methode d'etude contribue a mettre en
evidence des fluctuations significatives dans l’importance relative des populations et, par ia-
meme, dans la structure du peuplement lie aux couches holorganiques du sol.

Cette etude se situe en amont d'un programme de recherche a long terme mene sur les
communautes edaphiques d'un ecosysteme forestier (BLANDIN et al., 1980, 1985 ; GEOFFROY

Source :

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPER!:

535

et al. 1981, 1987). Elle s'integre aux donnees quantitatives obtenues tant sur les diplopodes
(GEOFFROY, 1981a, 1985) que sur des groupes ecologiquement complementaires tels que les
isopodes oniscoides (MOLFETAS, 1982) ou les larves de dipteres (MOLLON, 1982, 1983) et aux
recherches menees sur la dynamique de la transformation des materiaux organiques lors des
phases de colonisation par la faune edaphique (Garay el al. 1986a, b). En outre, cette
presentation des cycles d'activite des diplopodes de la foret de Foljuif s’insere dans une etude
des variations pluriannuelles de l'abondance des populations (GEOFFROY, in prep.).

Tableau 1. — Selection des travaux relatifs a T activity des populations de diplopodes edaphiques,
menes a l’aide de pieges d’interception de type Barber (PT).

Table 1. — Some selected studies dealing with soil millipede population activity \
investigated by Barber Pitfall Traps (PT).

REFERENCES

ECOSYSTEM

IiXALITY

COUNTRY

Adis 1979

Beech forest

Soiling, Gottingen

FRG

Albert 1978

Mixed forests

Wuppertal

FRG

Baker 1979, 1985. 1988

Sclerophyllous Forests & Grasslands

Aust. South-West

Australia

Banerjee 1967a. b

Oak forest

Roy. Holloway College

Barlow 1957. 1958

Dunes and forests

Meijendel Dune, Leiden

Netherlands

Blower 1970

Sycamore / ash wood

Ernocroft, Cheshire

Cotton & Miller 1974

Sand dunes

Marram Dune.Tentsmuir

Couret 1985

Quercus petraea & Pinus sylvestris

Orleans Forest

France

David & Poussardin 1983

Quercus petraea & Pinus sylvestris

Orleans Forest

France

Dethier & Pedroli-Christen 1983

Alpine grassland

Mt La Schera. Grisons

Switzerland

Drift van der 1951

Beech forest

NP de Hooge Veluwe

Netherlands

Fairhurst 1979

Sand dune

Gibraltar Pt, Lincolnsh.

Gillon & Gillon 1976

Grassland & savannah

Darou (Kaolack)

Senegal

Haacker 1968

Different ecosystems

Rhein-Main-Gebiert

FRG

Janati-Idrissi 1988

Brachypodium grass plot

St-Gely-du-Fesc

France

Korsos 1991

Dolomitic grassland

Buda Mts

Hungary

Kurnik 1985

Subalpine grassland to nival zone

GroBglockner

Austria

Kurnik 1988

Xerothermic & agricultural sites

Albeins : South Tyrol

Austria

Kurnik & Thaler 1985

Mixed oak forest

Stams. North Tyrol

Austria

Loksa 1988

Mixed grassland & forest sites

Pilis Mountains

Hungarv

Meyer 1979

Mountain to alpine ecosystems

Otzal

Austria

Meyer 1980, 1985

Mountain to alpine ecosystems

Obergurgl

Austria

Pedroli-Christen 1981

Mountain & subalpine forests

Swiss Jura

Switzerland

Pedroli-Christen & Scholl 1990

Mountain & subalpine forests

Swiss Jura

Switzerland

Spelda 1993

Coniferous forests

Oberreichenbach

FRG

SZEKELYHIDY & LOKSA 1979

Oak forets (Q. petraeae - cerris)

‘Sikfokut Projekt"

Hungary

Vajda & Hornung 1991

Sandy grassland

Kiskuns£g natl. Park

Hungary

MILIEU D'ETUDE ET METHODES

La foret mixte de Foljuif (Massif de Fontainebleau. Seine-et-Mame. France) se situe a 80 km au Sud-Est de Paris.
Elle pfesente deux zones principales qui different par la composition specifiquc de leurs peuplements vegetaux. Les
essences dominantes sont le charme, Carpinus betulus L., et le chene sessile, Quercus petraea (Mattus.) Liebl. dans la
premiere zone, lc pin sylvestrc, Pinus sylvestris L., et le chene sessile. Q. petraea (Mattus.) Liebl. dans la seconde. Les
differences notables dans la chute des materiaux au sol induisent plusieurs types d* humus : un moder dans la zone 1 et un
dysmoder dans la zone 2 (Blandin et al ., 1980 ; Garay, 1980). Dans la station & moder. trois couches de litiere existent
tout au long de Fannie : couches L, F et H. au sein desquelles la decomposition des feuilles est achevee au bout de trois
ans (Garay et al ., 1986a). Le programme de pfegeage a 6te conduit dans la station a moder de la zone 1 sous chene ( Q .
petraea) et charme (C. betulus).

Chaque pfege est constitue d'un pot de matfere plastique de 130 mm de diametre et de 90 mm de profondeur. A sa
surface, un orifice de capture de 80 mm de diam&tre est amenage grace h une plaque rectangulaire dont les bords s'insinuent
au niveau de la couche H de la litiere. evitant une rupture brutale entre l'environnement immediat et 1’orifice. On assure
ainsi une position precise du bord superieur du pot par rapport au substrat (Greenslade, 1964).

536

JEAN-JACQUES GEOFFROY & M ARIE-LOUISE CELERIER

Les appareils ont 6t6 installs dans une placette protegee de la zone d’£tude. en rangees paralleles de cinq pieges. La
distance entre deux pots est de I m. Les batteries de cinq pieges, disposees en quinconce, sont equidistantes de 10 m.

La capture d’un specimen resulte avant tout de l'interception naturelle de I’animal. Pour cela, le piege doit demeurer neutre,
d£pourvu & la fois de caractcrc attractif ou repulsif (Greenslade & Greenslade, 1971). Les individus captures sont
preserves dans un liquide de fixation neutre qui ne doit pas affecter la capture des especes (Luff. 1968). Nous avons
employ^ un liquide peu volatile, qui ne gele pas et assure une bonne conservation des animaux : l'ethylene-glycol
(Szekelyhidy & Loksa, 1979). Afin d'^viter une dilution par l'eau de pluie (Fichter, 1941 ; Steiner et al. , 1963), chaque
pidge est equipe d’une plaque de protection. En d6pit de certains risques de modification de capture (Dunger &
Engelmann, 1978), ce dispositif 6vite le rayonnement solaire et surtout la chute des debris organiques h I'interieur des
pieges (Fig. 1).

Les pifcges ont fonctionne simultanement duranl quatre ans selon les modalites presentees par le Tableau 2.
Toutefois. 1 'arret temporaire du piegeage, de juillet & octobre 1975 explique l'absence de capture durant cette periode sur
les Figures 2 a 8.

11 est important de signaler ici que le cycle annuel 1976 a £te marque par des conditions climatiques tr£s particulieres,
notamment par une secheresse considerable qui s'est etendue d'avril a septembre el qui a eu des effets immediats sur les
abondances des populations edaphiques (Blandin et ai, 1980, 1985).

La duree moyenne separant deux releves est de 14 jours. Les animaux recup6r6s sont fixes dans I'alcool 70°. Le laux de
capture par piegeage (“Trappability" : Pantis et al. , 1988) resultant des 81 experiences realis6es est exprime en nombre
d’individus captures pour 20 pieges-trappes et pour 14 jours (Ind/20 PT/14 D).

Fig. 1. Un piege d’ interception en place dans le sol en position ouverte. La limitc de la capacite de capture correspond
aux couches L et F de la liliere.

FtG. 1. — A pitfall— trap in open— position in the soil of the temperate Foljuif Forest. The limit of catching efficiency
corresponds to L and F litter layers.

RESULTATS

Diplopoda

Le peuplement global de diplopodes de la foret de Foljuif est compose de 14 especes parmi
lesquelles 7 especes sont frequentes ou abondantes. Les cinq especes dominantes sont :

Source : MNHN , Paris

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPERE

537

- Glomeris marginata (Villers, 1789) [Glomerida, Glomeridea]

- Melogona gallica (Latzel, 1884) [ Chordeumatida, Craspedosomatidea]

- Polydesmus angustus (Latzel, 1884) (Polydesmida, Polydesmidea]

- Allajulus nitidus (Verhoeff, 1891) [Julida, Julidea]

- Cylindroiulus punctatus (Leach, 1815) [Julida, Julidea]

auxquelles il faut ajouter :

- Cylindroiulus caeruleocinctus (Wood, 1864) [Julida, Julidea]

- Ommatoiulus sabulosus (Linne, 1758) [Julida, Julidea],

Seules ces especes ont ete collectees dans les pieges d'interception. Globalement, le cycle
d'activite du peuplement presente deux pics. Le premier se situe au printemps, entre mai et juin,
le second apparait en automne, de fin-octobre a debut decembre. Dans l'ensemble, les captures
sont moins importantes durant 1'hiver et le milieu de l'ete. La meme tendance saisonniere se
repete durant les quatre cycles annuels successifs et traduit notamment la capacite de deplacement
des adultes. En effet, les juveniles sont toujours tres faiblement represents dans les pieges,
meme en periode de recmtement des jeunes au debut de l’ete (Fig. 2).

Un cycle d’activite presentant deux maxima est souvent observe chez les diplopodes
edaphiques (e.g. DUNGER & STEINMETZGER, 1981 ; PEITSALMI, 1981 ; MEYER, 1985) alors
qu'une succession de trois pics est plus rarement mise en evidence chez certaines populations
(VAJDA & HORNUNG, 1991).

Le rapport saisonnier entre les sexes est rarement disproportionne avec toutefois une legere
surabondance des males adultes, traduisant la grande mobilite de ces demiers durant les periodes
actives.

Le faible taux de capture observe en automne 1976 est manifestement du a la secheresse
qui a marque cette annee-la. Remarquons toutefois que l'effet ne s'en fait pas sentir a long terme
car le niveau d'activite observe en automne 1977 et en 1978 est sensiblement le meme que celui
qui caracterisait le cycle de 1975.

Les pics d’activite de printemps sont le plus souvent plus importants que ceux de
l'automne, exception faite du printemps de 1977, dont le maxima plus faible traduit l'effet de la
secheresse de 1976 sur l'ensemble des especes. On observe une reponse quasi-immediate de
l'activite des adultes de plusieurs especes aux conditions climatiques tres contrastees de cette
annee. Ainsi, le pic isole de juillet 1976 (4 especes) coincide exactement avec une augmentation
ponctuelle des precipitations a la meme date. (cf. BLANDIN et ai, 1980, Fig. 1). De plus, on
peut observer une richesse specifique maximum au printemps, 7 especes de mai a juin,
regulierement superieure a celle de l'automne ou Ton rencontre 4 especes au plus (Fig. 3). Ce
phenomene s'explique par l'activite manifestee au printemps par les especes les moins frequentes
0 Ommatoiulus sabulosus et Cylindroiulus caeruleocinctus) et par une absence relative de
Cylindroiulus punctatus des couches superieures de litiere en automne. Ce Julide effectue alors
une migration en direction des horizons plus profonds du sol mais se trouve encore en grande
partie dans les bois morts en decomposition ou il a passe la periode estivale. C’est dans ce milieu
tres particulier que sont deposees la plus grande partie des pontes, qu’a lieu l’eclosion et que se
deroule les premieres etapes du developpement post-embryonnaire (GEOFFROY, 1981b). Durant
l'automne, l'activite la plus importante est partagee par Glomeris marginata et Allajulus nitidus ,
ce dernier occupant a ce moment toutes les couches de la litiere. (GEOFFROY, 1981b, 1985). La
periode hivernale est caracterisee par un taux de capture moyen ou faible selon les annees mais
qui est associe a une richesse specifique elle-meme tres faible. Cette periode coincide presque
exclusivement avec l'activite de surface des adultes males et femelles du chordeumide Melogona
gallica .

538

JEAN-JACQUES GEOFFROY & MARIE- LOUISE CELERIER

Tableau 2. — Programme de pi^geage realisd dans la foret temper£e dc Foljuif (zone I) durant la phase initiale du
programme de recherche, du 07.02.75 au 19.12.78 (series 1 & 81). S = n° de serie ; PT = nombre de pieges ; D =
nombre de jours.

TABLE 2. — Pitfall trapping programme realized in the temperate Foljuif forest ( zone I) during the initial period of the
research programme, from 07.02.75 to 19.12.78 (series I to 81 ). S = n° series ; PT = number of pitfall-traps ; D
= number of days.

PER I ODE

07.02.75-21.02.75

21.02.75-21.03.75

21.03.75-04.04.75

04.04.75-18.04.75

19.04.75-17.05.75

17.05.75-29.05.75

29.05.75-12.06.75

12.06.75-04.07.75

15.10.75-29.10.75

29.10.75-13.1 1.75

13.1 1.75-28.1 1.75

1 1

28.1 1.75-12.12.75

12.12.75-29.12.75

29.12.75-10.01.75

10.01.75-23.01.76

23.01.76-06.02.76

06.02.76-20.02.76

20.02.76-05.03.76

05.03.76-19.03.76

19.03.76-02.04.76

02.04.76-15.04.76

15.04.76-30.04.76

30.04.76-15.05.76

15.05.76-28.05.76

28.05.76-1 1.06.76

11.06.76-25.06.76

25.06.76-08.07.76

08.07.76-22.07.76

22.07.76-19.08.76

19.08.76-17.09.76

17.09.76-28.09.76

1 1

28.09.76-19.10.76

19.10.76-28.10.76

28.10.76-1 1.1 1.76

11.11.76-25.11.76

25.11.76-09.12.76

09.12.76-23.12.76

23.12.76-07.01.77

07.01.77-21.01.77

21.01.77-04.02.77

04.02.77-18.02.77

PERIODE

18.02.77-04.03.77

04.03.77-18.03.77

18.03.77-01.04.77

01.04.77-21.04.77

21.04.77-18.05.77

18.05.77-15.06.77

15.06.77-01.07.77

01.07.77-18.07.77

18.07.77-28.07.77

28.07.77-05.09.77

05.09.77-20.09.77

20.09.77-04.10.77

04.10.77-18.10.77

18.10.77-02.1 1.77

02.1 1.77-16.1 1.77

16.1 1.77-02.12.77

02.12.77-14.12.77

14.12.77-23.12.77

23.12.77-06.01.78

06.01.78-20.01.78

20.01.78-03.02.78

03.02.78-17.02.78

17.02.78-17.03.78

17.03.78-31.03.78

31.03.78-1 1.04.78

1 1

1 1.04.78-24.04.78

24.04.78-09.05.78

09.05.78-25.05.78

25.05.78-06.06.78

06.06.78-20.06.78

20.06.78-04.07.78

04.07.78-19.07.78

19.07.78-05.09.78

05.09.78-19.09.78

19.09.78-03.10.78

03.10.78-19.10.78

19.10.78-31.10.78

31.10.78-14.1 1.78

14.1 1.78-06.12.78

06.12.78-19.12.78

Source :

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPERE

539

2. — Cycles d’activite de 1'ensemble du peuplement de diplopodes dans les couches superieures du sol, de fevrier
1975 a decembre 1978 (fractions male, femelle ct juvenile de la communaute).

2. — Activity cycles of the whole millipede community in the upper soil layers of the temperate Foljuif forest, from
February 1975 to December 1978 (male, female and juvenile parts of the community).

Source :

540

JEAN-JACQUES GEOFFROY & M ARIE-LOUISE CELERIER

20-

10-

iji f'm'aim'j ijia1 s'oIn'd

Fig. 3. — Cycles d’activit£ des sept especes dominantes de diplopodes dans les couches sup£rieures du sol, de fevrier
1975 a decembre 1978 (total des indi vidus captures dans les pieges).

FlG. 3. — Activity cycles of the seven mainly representative millipede species (total trapped individuals for each
species) in the upper soil layers of the temperate Foljuif forest, from February » 1975 to December 1978.

Source :

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPERE

541

Melogona gallica

Une activite de surface du chordeumide Melogona gallica (= Microchordeuma gallicum
auct.) est mise en evidence uniquement pendant 1'hiver, de la fin d'octobre au debut d'avril. Les
individus captures sont en grande majorite des adultes (stade IX) parmi lesquels on note une
dominance relative frequente des males. Les juveniles, rares dans les pieges, sont represents
par des individus subadultes appartenant aux stades VII et VIII. Tous les Chordeumatida ont une
duree de vie courte et un cycle biologique etabli sur 1 ou 2 ans (BLOWER, 1978, 1979 ;
PEDROLI-CHRISTEN, 1978 ; MEYER, 1979 ; David, 1984). A Foljuif, le cycle de M. gallica
est annuel et la periode d'activite des adultes est en totale correspondance avec leur maximum
d'abondance a la limite des couches F et H. En revanche, le maximum de densite observe a la fin
du printemps a partir de releves quantitatifs (GEOFFROY, 1985) correspond a une grande
quantite de juveniles qui ne montrent pas d'activite de deplacement dans les couches de surface.
L'absence totale de capture durant 1’hiver 76-77 traduit l'effet immediat de la secheresse de 1976
sur la population (Fig. 4).

Fig. 4. — Cycles d’activite de Melogona
gallica (Chordeumatida) dans les
couches superieures du sol, de
fevrier 1975 a decembre 1978.

20-

1977

Fig. 4. — Activity cycles of the species
Melogona gallica (Chordeumatida)
in the upper soil layers of the
temperate Foljuif forest, from
February 1975 to December 1978.

1978

542

JEAN-JACQUES GEOFFROY & MARIE-LOUISE CELERIER

Polydesmus angustus

P. angustus est une espece relativement peu abondante dans le site etudie, oil elle effectue
son cycle biologique soit en un an, soit en deux ans, selon les individus. Les periodes de pontes
sont tres etalees dans le temps, ce qui peut etre reflete par la longue periode d'activite de surface
des adultes (stade VIII), notamment des femelles. Le cycle de 1978 en est particulierement
caracteristique (fevrier a septembre), le maximum de mai-juin pouvant correspondre a la
maturation des femelles (cf. COURET, 1985). La fraction juvenile piegee est representee par des
subadultes appartenant aux stades VI et VII. On les capture essentiellement en automne
(septembre a novembre). Ils correspondent a une partie de la population hivemale qui deviendra
adulte au printemps suivant. La fraction immature de la population (stades II a V), presente dans
les couches profondes de la litiere et du sol en ete, n'est pas capturee par un systeme de piegeage
visant a selectionner les individus actifs en surface (BLOWER, 1970). Cela n'empeche ni
l'activite, ni la mobilite a plus grande profondeur (COURET, 1985).

Polydesmus angustus

20-

10-

Ind/20PT/14D

1975

1978

I N 1 D 1

1977

F I M 1 A 1 M I J I J

Fig. 5. — Cycles d'activite du Polydesmida
Polydesmus angustus dans les
couches superieures du sol, de
fevrier 1975 a decembrc 1978.

Fig. 5. — Activity cycles of the
Polydesmid species Polydesmus
angustus in the upper soil layers of
the temperate Foljuif forest, from
February 1975 to December 1978.

L’activite des adultes est surtout marquee par un pic de printemps pour les deux sexes avec
un maximum en mai-juin. En automne en revanche, un tres leger pic ne concerne, certaines
annees, que les males (octobre). Le maximum de printemps s’accorde bien avec les resultats
obtenus par divers auteurs (BANERJEE, 1967a ; HAACKER, 1968 ; BLOWER, 1970 ; COURET,
1985). Par ailleurs, on observe une importance relative plus grande des males au debut de la
periode d'activite (mars a mai). Cependant, ce phenomene ne dure pas et le reste de la periode
active montre un equilibre entre les sexes ou au contraire, une plus forte presence des femelles en
mai-juin (Fig. 5).

En depit de toute preuve formelle liant l'activite de surface des adultes aux periodes de
reproduction, on est tente de conserver cette interpretation pour la periode active de printemps

Source :

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPERE

543

chez P. angustus. Nous devons garder a l'esprit que les femelles matures de P. angustus sont
capables d'effectuer plusieurs ovipositions ± etalees dans le temps au cours de leur vie adulte
(Snider, 1981 ; David & CELERIER, 1993) et que les modalites de la copulation elles-memes
peuvent presenter de tres larges variations (VERHOEFF, 1928 ; HUSSON, 1937 ; SAHLI, 1969).
Ces caracteristiques de la biologie de I'espece, liees a une activite prolongee des adultes
(Blower, 1969 ; BANERJEE, 1973) sont confirmees par l'observation d'accouplement et de
pontes a Foljuif s'etendant de debut avril a la fin novembre. Le nombre d'oeufs trouves dans les
logettes de pontes est variable, allant de 160 a 658 (!) pour une meme logette et reflete la capacite
potentielle des femelles a realiser 1 a 4 pontes successives au cours de l'annee. La periode de
mobilite maximum est suivie d'une disparition progressive des adultes actifs que Ton peut lier
d’une part a la mortalite des males et des femelles apres la reproduction, d’autre part aux
modifications du climat a l'approche de l'hiver durant lequel les subadultes et adultes survivants
migrent dans les couches profondes de l’humus et dans les bois morts (GEOFFROY, 1985).

Fig. 6. — Cycles d’activite du Glomerida
Glomeris marginata dans les
couches sup£rieures du sol, de
f6vrier 1975 h decembre 1978.

FlG. 6. — Activity' cycles of the Glomerid
species Glomeris marginata in the
upper soil layers of the temperate
Foljuif forest, from February 1975
to December 1978.

^ Glomeris marginata
9

1975

Ind/20PT/14D

30 D

1977

544

JEAN-JACQUES GEOFFROY & MARIE-LOUISE CELERIER

Glomeris marginata

Espece a longue duree de vie (jusqu'a 14 ans selon HEATH et at., 1974), Glomeris
marginata presente une activite de surface plus ou moins importante tout le long de l'annee. Elle
est presque exclusivement le fait des stades adultes epimorphes males et femelles. Le rapport des
sexes est le plus souvent tres en faveur des males, notamment durant la periode d'activite
maximale qui s'etend de mars a juillet (Fig. 6). On observe durant cette periode une diminution
de 1'abondance relative des males pieges par rapport aux femelles qui dominent au debut de l'ete
et au debut de l'automne, ce qui s'accorde avec les observations de HEATH et al. (1974). Le pic
automnal est moins marque que celui du printemps. C'est a l'automne (septembre a novembre)
que la proportion de juveniles dans les pieges est la plus elevee. II s'agit d'individus appartenant
soit aux stades epimorphes immatures (VI, ...?), soit au dernier stade anamorphe, stade V
(BOCOCK etal., 1967).

La position des pics d'activite des adultes au cours du cycle annuel est en correspondance
avec celui des pics des densites de la fraction epimorphe de la population (BLOWER & GABBUT,
1964). Le pic de densite des immatures que Ton observe frequemment au printemps n'apparait
pas ici. Leur niveau d'activite et de deplacement se situe dans les couches inferieures de
l'humus. Seuls les derniers stades subadultes sont pieges.

Selon BOCOCK & HEATH (1967), les periodes d'inactivite de G. marginata dans les
couches de litiere correspondent au moment de l'annee ou la temperature de surface est basse
(<6°C) et au moment de la mue. Dans la foret de Foljuif, l'inactivite hivernale s'etend de
decembre a fevrier et la periode estivale de mue des adultes est ponctuelle (3 a 4 semaines en
juillet-aout le plus souvent). Cela nous conduit a une duree annuelle d’activite de surface de 275
jours (3/4 de l’annee) superieure a celle rapportee par CARREL (1984) pour les forets
d'Angleterre. Cette duree d'activite peut meme etre bien superieure dans le cas d'annees a hivers
doux.

Cylindroiulus punctatus

L'activite de surface de ce Julida est principalement marquee par un pic au printemps (de
mars a juin) et diminue progressivement durant l’ete. En revanche, durant l'automne, les
captures demeurent faibles ou nulles, l'activite reprend regulierement des la fin de l'hiver suivant
(Fig. 7). Les adultes males et femelles (stades VII a XII) representent la quasi-totalite des
individus pieges, les immatures sont constitues d’individus appartenant aux stades V et VI. Le
rapport entre les sexes est extremement variable d'une annee sur l'autre et difficilement
interpretable. Tres favorable aux males adultes en 1975 et 1976, le sex-ratio montre au contraire
une forte importance des femelles en 1977 et un etat proche de l'equilibre en 1978. Cela montre,
une fois de plus, combien l'interpretation des cycles d'activite est delicate et comme il est facile
de privilegier une hypothese (e.g. l'activite des males adultes lie a l'accouplement) au detriment
d'une autre en observant un seul cycle annuel. La diminution du niveau d'activite observee en
1976 et 1977 peut etre mise en relation avec les conditions climatiques caracterisant l'annee
1976. En 1978. le taux de capture saisonnier devient superieur a celui de 1975, ce qui peut
s'expliquer par un accroissement global de l'activite de la population au cours des annees. Le pic
d'activite unique au printemps semble bien etre une caracteristique constante de l'espece.
BANERJEE (1967a) constate un maximum d'activite en avril-mai suivi d'une decroissance
reguliere jusqu'en decembre alors que DRIFT (1951) signale une periode d'activite exclusivement
de debut-avril a la mi-mai.

L'activite de surface printanniere a lieu au moment ou la population de C. punctatus de la
foret de Foljuif presente une abondance relative importante par rapport aux autres diplopodes
(GEOFFROY, 1981a). Elle coincide avec un pic de densite observe a la meme periode dans les
couches superieures L+F, suivi d’une migration d'une grande partie de la population dans les
bois morts tombes au sol des le debut de l’ete (BANERJEE, 1967b ; GEOFFROY, 1985). Le pic

Source : MNHN , Paris

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPERE

545

de densite, observe a la fois chez les adultes et les juveniles au printemps, concorde avec une
importante activite de surface. En revanche, le pic de densite des adultes signale en automne
(GEOFFROY, 1981b) correspond a une periode de migration en direction des couches organo-
minerales du sol (H+Al), ce qui explique en partie l'absence de capture par les pieges a cette
saison. De meme, les fortes densites observees de juillet a novembre par BLOWER & GABBUT
(1964), qui sont en grande partie dues a des stades juveniles successes, ne semblent pas avoir
de repercussions sur une activite de surface des individus en dehors de la periode printanniere.
La relation etroite que presente cette espece avec les bois morts en decomposition lors des
periodes de mue et de reproduction joue un role capital dans l'interpretation de son cycle
d'activite de surface (GEOFFROY, 1981b).

Fig. 7. — Cycles d'activite du Julida
Cylindroiulus punctatus dans les
couches superieures du sol. de
fevrier 1975 a decembre 1978.

FlG. 7. — Activity cycles of the Julid
species Cylindroiulus punctatus in
the upper soil layers of the
temperate Foljuif forest, from
February 1975 to December 1978.

20-

10-

Ind/20PT/14D

30D

Allajulus nitidus

Contrairement a C. punctatus. Allajulus nitidus (appele Cylindroiulus nitidus dans les
travaux anterieurs de nombreux auteurs) presente manifestement deux pics d'activite de surface
au cours du cycle annuel : le premier au printemps, de mars ajuin, et le second en automne, de
septembre a novembre. Ce phenomene se repete regulierement chaque annee. L'ete et l'hiver
sont caracterises par un arret total de l'activite de surface. Les individus captures sont
essentiellement des adultes males (stades VII a IX) et femelles (stades VIII a XI). Les juveniles,
peu frequents et peu nombreux, sont representes par des subadultes des stades VI et VII.

On ne remarque aucun decalage evident entre les periodes d'activite des males et des
femelles, les deux sexes etant largement representes lors des deux periodes d'activite maximale.

546

JEAN-JACQUES GEOFFROY & MARIE-LOU1SE CELERIER

On note toutefois une preponderance des femelles durant le printemps 1976 et l’automne 1978.
Un cycle d'activite proche de celui-ci existe dans d'autres populations forestieres de A. nitidus
(e.g. David, 1982, en foret d'Orleans). Des differences portent cependant sur le decalage
temporel ou l'activite relative observes entre les sexes. L'activite des femelles adultes domine au
printemps et est precoce en automne en foret d'Orleans, alors que l'activite relative des males est
plus frequente et plus durable dans la population de Foljuif, a l'exception de l'automne 1978. II
convient toutefois de rappeler que 1’interpretation biologique de ces taux de captures est delicate
et que plusieurs causes sont a l'origine des variations saisonnieres inter-annuelles et inter¬
populations.

Fig. 8. — Cycles d’activite du Julida
Allajulus nitidus dans les couches
superieures du sol, dc fevrier 1975 &
decembre 1978.

Fig. 8. — Activity cycles of the Julid
species Allajulus nitidus in the
upper soil layers of the temperate
Foljuif forest, from February 1975
to December 1978.

A. nitidus est une espece tres abondante dans certains ecosystemes forestiers (BLOWER,
1979 ; Blower & Miller, 1977), le maximum d'activite coincide alors avec le maximum de
densite au printemps et en automne (GEOFFROY, 1981b ; David, 1982). A Foljuif, durant la
periode d'etude consideree ici, A. nitidus domine numeriquement le peuplement de diplopodes
de la litiere toute l’annee sauf une partie de 1'hiver (GEOFFROY, 1981a). On ne peut exclure

Source :

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTER TEMPERE

547

qu'une augmentation de l'activite soit en partie liee a la reproduction : a l'accouplement d’une
part, traduit par l’activite intense des males adultes, a la ponte d'autre part lors de l'activite
maximale et dominante des femelles au printemps. C’est Interpretation la plus couramment
admise par les auteurs etudiant l’activite de diplopodes julides (e.g. BANERJEE, 1967b ;
Cotton & Miller, 1974). Nous devons considerer aussi que les periodes de forte activite des
adultes sont liees aux conditions climatiques qui favorisent la dispersion de la population dans
1 'ensemble des couches de la litiere, notamment au debut de l'automne, oil elle coexiste
temporairement avec C. punctatus (GEOFFROY, 1985). Les variations inter-annuelles ne
concernent ni le rythme ni la nature du cycle saisonnier d'activite. En revanche, on constate une
diminution progressive du niveau d'activite, qui se traduit par un taux de capture de printemps
decroissant regulierement de 1975 a 1978. La diminution observee en automne 1976 et au
printemps 1977 s'interprete comme une reaction immediate aux conditions de secheresse de
1976. Cet effet n'est pas durable puisque l'activite reprend fortement a l'automne de 1977.
Pourtant, le cycle de 1978 est caracterise par un niveau d'activite plus faible, contrairement a
toutes les especes etudiees precedemment. Cela induit un changement dans l'organisation
fonctionnelle du peuplement que Ton doit considerer en comparant les variations relatives de
populations proches, notamment celles des autres Julida.

DISCUSSION

Considere dans son ensemble, le peuplement de diplopodes edaphiques de la foret de
Foljuif presente constamment une activite de surface dans la litiere. Le cycle annuel, d'allure
“bimodale”, montre un premier pic au printemps et un second pic a l'automne. Ceci est la
resultante de l'activite de la fraction adulte et subadulte de plusieurs populations dont
1'importance relative varie considerablement au cours du temps. Glomeris marginata et
Polydesmus angustus assurent, a des degres divers, un apport constant d'individus actifs tout au
long du cycle annuel. L'activite hivernale est essentiellement due au chordeumide Melogona
gallica. Les pics printaniers et automnaux sont accentues par l'activite conjuguee des julides,
surtout au printemps pour Cylindroiulus punctatus, et reflete en grande partie le cycle d'activite
de la population la plus abondante lors des premieres annees de la periode d'etude : Allajulus
nitidus.

Les traits generaux du cycle annuel d'activite ainsi defini se repetent regulierement chaque
annee. La reponse a court terme due aux effets du climat local dans l'ecosysteme ne s'etend pas
au-dela du cycle suivant. Ainsi, l'impact de la secheresse du printemps et de l'ete de 1976 se fait-
il sentir de l'automne 1976 a la fin du printemps 1977.

Au-dela de ce constat general de relatives stabilite et regularite, des modifications dans la
composition du peuplement apparaissent au cours du temps, sur lesquelles il y a lieu de
s'interroger. Elies concernent notamment 1'importance relative des deux especes de Julida les
plus frequemment capturees dans les pieges : A. nitidus et C. punctatus (FIG. 9). A. nitidus
domine clairement le peuplement de diplopodes en 1975 et il conserve une importance relative
considerable jusqu'en automne 1977. Mais l'observation des quatre cycles successifs montre
une diminution progressive de cette espece durant la phase de printemps, au profit de C.
punctatus qui finit par le remplacer en grande partie en 1978. A. nitidus reste dominant durant le
debut de l'automne, les grands indi vidus de C. punctatus etant alors encore distribues dans les
bois morts, ce qui explique le fort pourcentage que represente A. nitidus dans les demieres series
de piegeage (FIG. 10). Cette diminution considerable de A. nitidus pourrait n'etre en fait qu'un
phenomene temporaire du aux conditions locales autorisant les captures ou, partiellement, au
hasard des deplacements. Il semble que la modification constatee ait une signification plus
profonde et plus durable. Il ne s'agit pas settlement d'une diminution de l'activite mais egalement
de l'abondance relative, mesuree par la densite des individus dans les echantillonnages
quantitatifs. En outre, les etudes ulterieures confirment bien cette tendance.

548

JEAN-JACQUES GEOFFROY & MARIE-LOUISE CHEERIER

1975

Allajalus nitidus

1976

Fig. 9. — Cycles d’activit£ compares de
deux cspeccs de Julides, Allajulus
nitidus (An) et Cylindroi ul us
punctatus (Cp), dans les couches
superieures du sol, de fevrier 1975 a
decembre 1978 (nombre total
d’indi vidus pour chaque
population).

FlG. 9. — Compared activity cycles of two
Julid species, Allajulus nitidus (An)
and Cylindroiulus punctatus (Cp). in
the upper soil layers of the
temperate Foljuif forest, from
February 1975 to December 1978
(total number of individuals for each
specific population).

Durant le cycle annuel 1976, A. nitidus apparait nettement comme le diplopode le plus
abondant du peuplement, tant en densite qu'en biomasse (GEOFFROY, 1981a). II domine
largement C. punctatus dans les couches de litiere, ainsi que le montrent les modalites de la
coexistence de ces deux populations dans lecosysteme (GEOFFROY, 1981b). Au cours du cycle
annuel de 1982 au contraire, on observe un accroissement relatif et une dominance de C.
punctatus par rapport a A. nitidus et par rapport aux autres diplopodes durant toutes les phases
saisonnieres de l’annee (GEOFFROY, 1985).

En premiere analyse, les variations relatives observees entre les deux populations tout au
long de la periode de piegeage sont considerees comme un phenomene traduisant une reelle

Source : MNHN , Paris

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTER TEMPERE

549

modification de I'activite de l'une par rapport a l'autre et un remplacement progressif, dans les
couches L+F dc la litiere, de A. nitidus par C. punctatus (Fig. 10). La realite de ce phenomene
doit etre verifiee par l'analyse comparative des cycles d'abondance des deux especes au cours de
l’etude a long terme qui a suivi la periode de piegeage (BLANDIN el al., 1985) et au cours de
laquelle l'importance relative de C. punctatus semble se confirmer (GEOFFROY, in prep.). Cela
presente un interet majeur dans la dynamique de l'organisation et des modalites d'action d'un
groupe fonctionnel de macroarthropodes saprophages consommateurs et transformateurs des
couches superieures peu degradees de la litiere, au cours des deux premieres annees de
decomposition de celle-ci.

E3 Cp% Cp = Cylindroiulus punctatus
B An% An = Allajulus nitidus

Series (1 - 81)

Fig. 10. — Importance relative de deux especes de Julides, Allajulus nitidus (An) et Cylindroiulus punctatus (Cp). durant
la periode de piegeage (series 1 k 81), dans les couches superieures du sol de la foret de Foljuif.

FlG. 10. — Relative importance of two Julid millipedes, Allajulus nitidus (An) and Cylindroiulus punctatus (Cp), during
the trapping period (series 1 to 81) in the upper soil layers of the temperate Foljuif forest.

A l'echelle de 1'ensemble des compartiments de litiere et des horizons du sol, e'est la
totalite de la guilde des diplopodes saprophages qui entre en jeu, depuis les plus jeunes stades
jusqu'aux adultes a mues post-imaginales. Mais toutes les categories d'individus n'agissent pas
en meme temps et de la meme fagon a l'interieur d'un meme compartiment edaphique. La
selection d'une partie du peuplement lors de la capture par piegeage nous renseigne sur la nature
des individus capables d'agir au sein des couches les plus superficielles de la litiere. Elle
contribue a distinguer, parmi les diplopodes. deux groupes d'animaux montrant des modalites
d'activite et de fonction differentes dans les divers compartiments edaphiques de l'ecosysteme.
Seuls les individus les plus grands - et done les plus ages - semblent capables de se mouvoir
dans les couches les moins profondes de la litiere, et d'y realiser une fonction de consommation
et de transformation. Ainsi que Ton distingue couramment “micro-arthropodes” et “macro¬
arthropodes”, il conviendrait de distinguer des “micro-diplopodes” et des “macro-diplopodes”.
Ces deux groupes sont separes ici par une limite liee au taux de capture potentiel dans nos pieges

550

JEAN-J ACQUES GEOFFROY & MARIE-LOUISE CELERIER

d'interception qui selectionnent preferential lement les individus qui se deplacent activement en
surface. Une limite approximative est ainsi definie au sein de l'ensemble des diplopodes. Les
macro-diplopodes, frequents dans les pieges, representent un groupe fonctionnel particulier
(Fig. 11). Bien entendu, 1’organisation dun tel groupement d'individus doit etre confirmee par
d’autres voies d'observation et d'experimentation : repartition spatio-temporelle des individus et
modalites de la consommation des feuilles. En outre, la comparaison et l'integration de ces
mesures a celles obtenues chez les isopodes oniscoi'des devraient favoriser la comprehension
d'un “groupe fonctionnel de macroarthropodes saprophages” dans I'ecosysteme forestier etudie.

STADIA

~ i n i in

IV 1 V

VII

VIII

XII

Melogona

gallica

Polydesmus

angustus

Glomeris

marginata

Allajulus nitidus

Cylindroiulus

punctatus

Cylindroiulus

caeruleocinctus

Ommatoiulus

sabulosus

Fig. 11. — Repartition des differents stades de sept especes de diplopodes dans les couches supdrieurs de liti£re et de sol
de la foret temperee de Foljuif : individus captures par piegeage d’interception de fevrier 1975 & d£cembre 1978.
Ligne en gras : peuplement actif de macro-diplopodes ; ligne double : peuplement de micro-diplopodes.

F/C. 11. — Occurence of the different stadia of seven millipede species in the upper soil and litter layers of the temperate
Foljuif forest: individuals caught in Pitfall traps from February t 1975 to December 1978. Thick line: active macro-
diplopod community: double line : micro-diplopod community.

* : Frequent or abundant. 0 : Occasional or scarce.

REMERCIEMENTS

Ce travail a benelici£ dune aide du Ministere des Universes et du Ministere de l'Enseignement Superieur et de la
Recherche. Nous sommes reconnaissants & M. le Pr. P. Blandin (M.N.H.N., Brunoy) pour l'attention soutenue qu'il a
apportee a l'61aboration de cette publication. Nous remercions M. Loyau et Mme Raulo (E.N.S.. Station Biologique de
Foljuif) pour leur aide technique sur le terrain et au laboratoire.

REFERENCES

Adis, J.. 1979. — Problems of interpreting arthropod sampling with pitfall traps. Zool. Anz., 202 : 177-184.

Albert, M., 1978. — Bodenfallenfange von Diplopoden und Isopoden in Wuppertaler Waldern (MB 4708/09). Jahr.
Naturw. V. Wuppertal, 31 : 46-49.

Baker, G. H.. 1979. — The activity patterns of Ommatoiulus moreletti (Diplopoda: Julidae) in South Australia. J. Zool.
Lond ., 188 : 173-183.

Baker. G. H., 1985. — The distribution and abundance of the Portuguese millipede Ommatoiulus moreleti (Diplopoda:
Julidae) in Australia. Aust. J. Ecol., 10 : 249-259.

Baker, G. H., 1988. Patterns in the activity of the millipede Ommatoiulus moreleti (Diplopoda: Julidae). Aust.
Entomol. Mag., 15 : 127-139.

Banerjee, B., 1967a. — Diurnal and seasonal variations in the activity of the millipedes Cylindroiulus punctatus
(Leach), Tachypodoiulus niger (Leach) and Polydesmus angustus Latzel. Oikos, 18 : 141-144.

Banerjee, B., 1967b. — Seasonal changes in the distribution of the millipede Cylindroiulus punctatus (Leach) in
decaying logs and soil. J. Anim. Ecol., 36 : 171-177.

Source :

CYCLES D’ACTTVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPERE

551

Banerjee, B., 1970. — A mathematical model on sampling diplopods using pitfall traps. Oecologia , 4 : 102-105.
Banerjee, B., 1973. — The breeding biology of Polydesmus angustus Latzel (Diplopoda, Polydesmidae). Nork. ent.
Tidsskr. , 20 : 291-294.

Barber, H., 1931. — Traps for cave-inhabiting insects. J. Elisha Mitchell sci. Soc., 46 : 259-266.

Barlow, A., 1957. — A factorial analysis of distribution in three species of Diplopods. Tijdschr. Ent., 100 : 349-426.
Barlow, C. A., 1958. — Distribution and seasonal activity in three species of diplopods. Arch. Neerland. Zool., 13 :
108-133.

Blandin, P., Abbadie, L., Courault, S., Garay, I., & Geoffroy, J. J., 1980. — Etude d un ecosysteme forestier mixte.

I. Climat, structure de la vegetation et retomb£es de litiere. Rev. Ecol. Biol. Sol, 17 : 181-198.

Blandin, P.. Flogaitis, E., Arbeille, J., & Geoffroy., J. J., 1985. — Les variations interannuelles de
macroarthropodes edaphiques dans une foret temperee. Bull. Ecol., 16 : 273-283.

Blower, J. G., 1969. — Age structure of millipede populations in relation to activity and dispersion. In : J. G. Sheals,
The soil ecosystem.. London, Publ. Syst. Ass., 8 : 209-216.

Blower, J. G., 1970. — The millipedes of a Cheshire wood. J. Zool. Lond., 160 : 455-496.

Blower, J. G., 1978. — Anamorphosis in the Nematophora. Abh. Verb, naturwiss. Ver. Hamburg , 21/22 : 97-103.
Blower, J. G., 1979. — The millipede faunas of two British limestone woods. In : M. Camatini, Myriapod Biology.
London, Academic Press : 203-214.

Blower, J. G. & Gabbut., P. D., 1964. — Studies on the millipedes of a Devon oak wood. Proc. Zool. Soc. London ,
143 : 143-176.

Blower, J. G. & MILLER., P. F., 1977. — The life-history of the Julid millipede Cylindroiulus nitidus in a Derbyshire
wood. J. Zool. Lond., 183 : 339-351.

Bocock, K. L. & J. Heath., 1967. — Feeding activity of the millipede Glomeris marginata (Villers) in relation to its
vertical distribution in the soil. In : O. Graf & J. E. SATCHELL, Progress in Soil Biology. Amsterdam, North Holland
Publ. C° : 233-240.

Bocock, K. L.. M. D.. Mountford & Heath., J., 1967. — Estimation of annual production of a millipede population.

In : K. PETRUSEWICZ, Secondary productivity of terrestrial ecosystems. Warsawa : 727-739.

Briggs, J. B., 1961. — A comparison of pitfall trapping and soil sampling in assessing populations of two species of
ground beetles. Rep. E. Mailing Res. Sta., 1960 : 108-112.

Carrel, J. E., 1984. — Defensive secretion of the pill millipede Glomeris marginata. I. Fluid production and storage. J.
Chem. Ecol., 10 : 44-51.

COTTON, M. J. & Miller, P. F., 1974. — A population of Cylindroiulus latestriatus (Curtis) in sand dunes. Symp. Zool.
Soc. London . 32 : 589-602.

Couret, T., 1985. — Recherches sur la biologic et l'ecologie de deux populations de diplopodes en foret d Orleans :
Polyzonium germanicum Brandt, 1831 et Polydesmus angustus Latzel, 1884. These. Doctoral de 3e cycle, Ecologie,
Orleans, 144 pp.

David, J. F., 1982. — Variability dans l'espace et dans le temps des cycles de vie de deux populations de Cylindroiulus
nitidus (Verhoeff) (Julida). Rev. Ecol. Biol. Sol, 19 : 411-425.

David, J. F., 1983. — Contribution a Ietude des populations de Diplopodes en Foret d'Orleans, et de leur role dans la
degradation de la litiere. Th£se, Doctoral de 3e cycle, Ecologie, Orleans, 149 pp.

David, J. F., 1984. — Le cycle annuel du diplopode Microchordeuma gallicum (Latzel, 1884). Bull. Soc. zool. France,
109 : 61-70.

David, J. F. & C£l£rier., M. L., 1993. — Nombre d'ovipositions et fecondite chez le Diplopode Polydesmus angustus
Latzel (Polydesmida : Polydesmidae). C. R. Acad. Sci. Paris, 316 : 225-228.

David, J. F.& Poussardin, C., 1983. — ^interpretation du piegeage du Diplopode Microchordeuma gallicum (Latzel,
1884). Rev. Ecol. Biol. Sol , 20 : 579-587.

Dethier, M. & Pedroli-Christen., A.. 1983. — Diplopodes et Chilopodes dune pelouse alpine du Parc National Suisse.
Bull. Soc. Vaudoise Sci. nat., 364 : 373-379.

Drach, A.. BenestG. & Cancela da Fonseca, J. P.. 1981. — Analyse comparative de differents types de pieges basee
sur Ietude de deux peuplements de Carabiques (Col. Carabidae). Rev. Ecol. Biol. Sol, 18 : 91-114.

Drift, J. van DER, 1951. — Analysis of the animal community in a beech forest floor. Tijdschr. Ent.. 94 : 1-168.
Dunger, W. & Engelmann, H. D., 1978. — Testversuche mit immissiongeschiitzen Bodenfallen fur Mikroarthropoden.
Pedobiologia, 18 : 448-454.

Dunger, W. & Steinmetzger, K., 1981. — Okologischc Untersuchungen an Diplopoda einer Rasen-Wald-Catena im
Thiiringer Kalkgebiet. Zool. Jahrb. Abt. Syst. Okol. Geogr. Tiere, 108 : 519-533.

552

JEAN-JACQUES GEOFFROY & MARIE-LOUISE CELERIER

Fairhurst, C. P., 1979. — The distribution of Millipedes on a Lincolnshire sand dune system, In : M. Camatini,
Myriapod Biology. London, Academic Press : 241-250.

Fighter, E., 1941. — Apparatus for the comparison of soil surface arthropod populations. Ecology , 22 : 338-339.

Franke, U. & Friebe., B., 1983. — Erfassung der Makrofauna eines Buchenwaldbodens mittels Handauslcse und
Barbenfallen. Verh. Disch. Zool. Ges.. 1983 : 216.

Franke, U., Friebe B. & Beck, L., 1988. — Methodisches zur Ermittlung der Siedlungsdichte von Bodentiercn aus
Quadratproben und Barbenfallen. Pedobiologia. 32 : 253-264.

Friebe, B., 1983. — Zur Biologie eines Buchenwaldbodens 3. Die Kdferfauna. Carolinea . 41 : 45-80.

Garay, I., 1980. — Etude d'un ecosysteme forestier mixte. II. - Les sols. Rev. Ecol. Biol. Sol, 17 : 525-541.

Garay. I., Mollon A. & Flogaitis, E., 1986. — Etude dune litiere forestiere mixte a charme ( Carpinus betulus L.) et

chene ( Quercus sessiliflora Smith). II. - Succession des macroarthropodes au cours de la decomposition. Acta

(Ecologica, (Ecol. Gener., 7 : 263-288.

Garay. I.. Nazoa S. & Abbadie, L., 1986. — Etude dune litiere forestiere mixte k charme et k chene. I. - Decomposition
de la mature organique et dynamique des elements biogenes. Acta (Ecologica, (Ecol. Gener., 7 : 151-169.

Geoffroy, J. J.. 1981a. — Etude d'un ecosysteme forestier mixte. V. Traits generaux du peuplement de diplopodes
edaphiques. Rev. Ecol. Biol. Sol. 18 : 357-372.

Geoffroy, J. J., 1981b. — Modalites de la coexistence de deux diplopodes, Cylindroiulus punctatus (Leach) et
Cylindroiulus nitidus (Verhoeff), dans un ecosysteme forestier du Bassin Parisicn. Acta (Ecologica, (Ecol. Gener., 2 :
227-243.

Geoffroy, J. J., 1985. — Les diplopodes d’un ecosysteme forestier temper : repartition spatio-temporelle des
populations. Bijdragen tot de Dierkunde, 55 : 78-87.

Geoffroy, J. J., C£l£rier, M. L., Garay, I.. Rherissi S. & Blandin, P., 1987. — Approche quantitative des fonctions
de transformation de la matiere organique par des macroarthropodes saprophages (Isopoda et Diplopoda) dans un sol
forestier k moder : protocoles experimentaux et premiers resultats. Rev. Ecol. Biol. Sol , 24 : 573-590.

Geoffroy, J. J.. Christophe, T., Molfetas S. & Blandin, P.. 1981. — Etude d’un ecosysteme forestier mixte. III. Traits
generaux du peuplement de Macroarthropodes edaphiques. Rev. Ecol. Biol. Sol, 18 : 39-58.

Gilbert, O., 1956. — The natural histories of four species of Calathus (Col. Carabidae) living on sand dunes in
Anglesey, North Wales. Oikos, 7 : 22-47.

Gillon, Y. & Gillon, D.. 1976. — Comparaison par piegeage des populations de Diplopodes Iuliformes en zone de
vegetation naturelle et champ d’arachide. Cahiers ORSTOM, ser. Biol., XI : 121-127.

Gist, C. S. & CROSSLEY, D. A., 1973. — A method for quantifying pitfall trapping. Environ. Entomol. , 2 : 951-952.

Greenslade, P. & Greenslade, P. J. M., 1971. — The use of baits and preservatives in pitfall traps. J. Aust. Ent. Soc.,
10 : 253-260.

Greenslade, P. J. M., 1964. — Pitfall trapping as a method for studying populations of Carabidae. J. Anim. Ecol. ,33 :
301-310.

Haacker, U., 1968. — Descriptive, experimented und verglcichende Untersuchungen zur Autokologie Rhein-
mainischer Diplopoden. Oecologia , I : 87-129.

Heath, J., BOCOCK K. L. & MOUNTFORD, M. D., 1974. — The life-history of the millipede Glomeris marginata (Villers)
in North-West England. Symp. Zool. Soc. London , 32 : 433-462.

HUSSON, R.. 1937. — Reproduction non saisonniere des polydesmides (Dipl.) des galcries de mines. Bull. Soc. Sci.
Nancy .7: 195-198.

Janati-Idrissi, A., 1988. — Analyse du role des diplopodes en region mediterraneenne et influence de l'impact humain
sur leurs communautes. These de Doctorat. Ecologie. Montpellier III, 229 pp.

Joose, E. N. G. & Kapteijn, J. M., 1968. — Activity-Stimulating phenomena caused by Field-disturbance in the use of
pitfall-traps. Oecologia, 1 : 385-392.

Kaczmarek, W., 1978. — Die lokomotorische Akti vital der Bodenfauna als Parameter der trophischen Struktur und der
Sukzession von Waldokosystem. Pedobiologia, 18 : 434-441.

Korsos, Z., 1991. — Abundance and seasonal activity of millipedes in a dolomitic grassland community (Diplopoda).
Annls. hist.-nat. Mus. naln. Hung., 83 : 239-244.

Kurnik, I., 1985. — Uber Diplopoden aus Barberfallen im Bereich der Grossglockener - Hochalpenstrasse (Karnten,
Osterreich). Carinthia II, 175 : 271-283.

Kurnik, I., 1988. — Uber Diplopoden von Xerotherm und Kulturstandarten bei Albiens (Stidtirol). Ber. nat.-med. Verein
Innsbruck, 75 : 109-113.

Kurnik. I. & Thaler, K., 1985. — Weiterc Diplopoden-Fallenfange in Nordtirol (Osterreich) (Myriapoda : Diplopoda).
Ber. nat.-med. Verein Innsbruck, 72 : 145-154.

Source : MNHN. Paris

CYCLES D'ACTIVITE DE DIPLOPODES DANS UN ECOSYSTEME FORESTIER TEMPERE

553

LeBerre, J. R.. 1969. — Les methodes dc piegeage des invcrtebres. In : Lamotte, M. & Bourliere, F., Problemes
d'ecologie : I'echantillonnage des peuplements animaux des milieux terresires. Paris, Masson : 55-96.

Lecordier, C. & Benest., G., 1982. — Etude d'un ecosystEme forestier mixte. VI. Les carabiques (Col.). Rev. Ecol Biol
Sol , 19 : 89-104.

Loksa, 1., 1988. — Uber einige Athropoden-Gruppen aus dem Biosphare-Reservat des Pilis-Gebirgcs (Ungarn). 1. Die
Diplopoden, Chilopodcn, Weberknechte und Spinnen vom Szamar-Berg und aus der Umgebung der Loss-Wand von
Basaharc. Opusc. Zool. Budapest, XXIII : 159-176.

Luff, M. L., 1968. — Some effects of formalin on the numbers of Coleoptera caught in pitfall traps. Entomol. mon.
Mag., 104 : 115-116.

Luff, M. L., 1975. — Some features influencing the efficiency of pitfall traps. Oecologia, 19 : 345-357.

Mader, H. J. & MOhlenberg, M., 1981. — Artenzusammensetzung und Ressourcenangebot einer kleinflachigen
Habitatinsel, untersucht am Beispiel der Carabidenfauna. Pedobiologia, 21 : 46-59.

Meyer, E., 1979. — Life cycles and ecology of high alpine Nematophora. In : M. Camatini, Myriapod Biology.
London, Academic Press : 295-306.

Meyer, E., 1980. — Okologische Untersuchungen an Wirbellosen des zentralalpinen Hochgebirges (Obergurgl. Tirol) :
IV. Aktivitatsdichte, Abundanz und Biomasse der Makrofauna. Veroffentl. Univ. Innsbruck, 125 : 1-54.

Meyer, E., 1985. — Distribution, activity, life-history and standing crop of Julidae (Diplopoda, Myriapoda) in the
central High Alps (Tyrol, Austria). Holarci. Ecol, 8 : 141-150.

MITCHELL, B., 1963. — Ecology of two carabid beetles, Bembidion lampros and Trechus quadristriatus. J. Anim. Ecol.,
32 : 377-392.

Molfetas, S., 1982. — Etude d'un ecosysteme forestier mixte. VIII. - Les Isopodes. Rev. Ecol. Biol. Sol, 19 : 427-438.

Mollon, A., 1982. — Etude d'un ecosysteme forestier mixte. VII. - Composition et phenologie du peuplement des
DiptEres a larves edaphiques. Rev. Ecol. Biol. Sol, 19 : 289-306.

Mollon, a., 1983. — Etude d'un ecosysteme forestier mixte. IX. - Les Dipteres Edaphiques, structure temporelle du
peuplement larvaire. Rev. Ecol. Biol. Sol, 20 : 367-383.

Murdoch, W. W., 1966. — Aspects of the population dynamics of some marsh Carabidae. J. Anim. Ecol., 35 : 127-
156.

Pantis, J. D., Stamou G. P. & Sgardelis., S. S., 1988. — Activity patterns of surface ground fauna in Asphodel deserts
(Thessalia, Greece). Pedobiologia, 32 : 81-87.

Park, O., 1935. — Studies on nocturnal ecology. III. Recording apparatus and further analysis of activity rhythm.
Ecology, 16 : 152-163.

Park. O., Lockett J. A. & Myers., D. J.. 1931. — Studies on nocturnal ecology with special reference to climax forest.
Ecology, 12 : 709-727.

Pedroli-Christen, A., 1978. — Contribution a la connaissance du developpement post-embryonnaire de Craspedosoma
alemannicum Verhoeff et de Xylop/iageuma zschokkei Bigler (Diplopoda. Nematophora) dans une tourbiere du Haut-
Jura. Rev. Suisse Zool., 85 : 673-679.

Pedroli-Christen, A., 1981. — Etude des peuplements de diplopodes dans six associations forestieres du Jura et du
Plateau Suisse (Region Neuchateloise). Bull. Soc. Neuchdteloise Sci. nat., 104 : 89-106.

Pedroli-Christen, A. & A. Scholl., 1990. — Ecological and genetic studies on parapatric Rhymogona silvatica Roth,
and R. cervina Verh. (Diplopoda : Craspedosomatidae) with special reference to hybrid populations in a zone of
contact. Rev. Suisse Zool. 97 : 349-359.

PEITSALMI, M., 1981. — Population structure and seasonal changes in activity of Proteroiulus fuscus (Am Stein)
(Diplopoda, Blaniulidae). Acta Zool Fenn., 161 : 1-66.

Rusdea, E., 1992. — Stabilisierende Selektion bei Microphthalmen Hohlentieren : Untersuchungen zur Tages-
Zeitlichen Aktivitatsverteilung und Populationsdynamik von Lemostenus schreibersi (Kiister) (Carabidae). Mem.
Biospeol., 19: 1-110.

Sahli. F., 1969. — Sur la biologie sexuelle (accouplement, ponte, instincts sexuels) du Diplopode Polydesmus angustus
Latzel. Rev. Ecol Biol. Sol, 6 : 437-445.

Snider, R., 1981. — The reproductive biology of Polydesmus inconstans (Diplopoda: Polydesmidae) at constant
temperatures. Pedobiologia, 22 : 354-365.

SOUTH WOOD, T. R. E., 1978. — Ecological Methods, with particular reference to the study of Insect populations (2nd
ed.). London, Methuen & C°.

SPELDA, J.. 1993. — Hundert- und Tausendfusser aus Missen der Umgebung von Oberreichenbach, Lkr. Calw (Chilopoda,
Diplopoda). Beih. Veroff Naturschutz Landschaftspfelge Bad.-Wurtt., 73 : 399-402.

554

JEAN-JACQUES GEOFFROY & MARIE-LOUISE CELERIER

Steiner. P.. Wenzeland. F. & Baumert., D., 1963. — Zur Beeinllussung der Arthropodenfauna nordwest deutscher
Kartoffelfelder durch die Andwendung synthetischer Kontaktinsektizide. Mitt. biol. BundAnst. Ld.-u. Forstw.,
Berlin , 109 : 1-38.

Szekelyhidy, E. H. & Loksa, I., 1979. — Oniscoiden-, Diplopoden- und Chilopoden- Gemeinschaften im
Untersuchungsgebiet “Sikfdkut-Projekt” (Ungam). Opusc. Zool. Budapest, XVI : 151-174.

Topping, C. J. & Sunderland, K. D., 1992. — Limitations to the use of pitfall traps in ecological studies examplified
by a study of spiders in a field of winter wheat. J. Appl. Ecology. 29 : 485-491.

Vajda. Z. & HORNUNG, E., 1991. — Temporal and spatial pattern of a diplopod population (Megaphyllum unilineatum
(C.L. Koch) in a sandy grassland. Acta Biol. Szeged.. 37 : 75-81.

Verhoeff, K. W., 1928. — Diplopoda 1. In : H. G. Bronn’s Klassen und Ordnungen des Tierreichs. 5, Leipzig,
Akademische Verlagsgesellschaft : 1-1072.

Williams, G., 1958. — Mechanical time-sorting of pitfall captures. J. Anim. Ecol. . 27 : 27-35.

Williams, G., 1959. — The seasonal and diurnal activity of the fauna sampled by pitfall traps in different habitats. J.
Anim. Ecol.. 28 : 1-13.

Source : MNHN , Paris

Traces de l'activite de diplopodes dans des sols et des
sediments karstiques du Maroc Atlantique

Colette JEANSON *, Hsain El AlSSAOUl ** Jean-Pierre ADOLPHE **

* CNR S, Laboratoire d’Ecologie Generate, Museum National d'Histoire Naturelle
4, avenue du Petit Chateau, F- 91800 Brunoy, France

** Laboratoire de Geomicrobiologie, Universite Pierre & Marie Curie, 4, place Jussieu, F-75005 Paris, France

RESUME

Une 6tude pluridisciplinairc et multiscalaire conduit les auteurs & utiliser des donn£es de la pedozoologie pour
interpreter des observations stklimentologiques et geologiques. Les concretions carbonatees des sediments karstiques,
des agr£gats terreux de sols actuels et les dejections obtenues experimentalement par elevage de Glomeris marginata
(Diplopoda, Glomerida), presentent une tres nette analogie morphologique et physicochimique. Des relations sont
etablies entre les phenomenes de pedobioturbation, pateobioturbation et sedimentation.

ABSTRACT

Activity traces of millipedes in soils and karstic sediments in Atlantic Morocco.

The authors are carrying into effect a pluridisciplinary and multiscalar approach for using pedozoology data to
interpret sedimentological and geological observations. Carbonated concretions of karstic sediments, earthy
aggregates of Present soils, and experimental feacal pellets of Glomeris marginata (Diplopoda, Glomerida) show a very
clear morphological and physico-chemical analogy. Relationships are then identified between pedobioturbation,
paleobioturbation and sedimentation phenomena.

INTRODUCTION

Les formations carbonatees continentales du Haut-Atlas occidental marocain montrent un
developpement spatial important et presentent une grande diversite de facies : des revetements
pelliculaires sur les galets des oueds, des croutes en dalles compactes, des croutes zonaires
laminees, des croutes conglomeratiques, des edifices travertineux et des concretionnements
pedo-karstiques (ADOLPHE, 1981 ; HOURIMECHE, 1988). La demarche initiale de cette etude
consiste a circonscrire et a decrire sur le terrain le domaine des concretionnements pedo-
karstiques constitues par des assemblages de concretions carbonatees de quelques millimetres en
forme de “cocons". Cela dans le but de tenter de comprendre leur origine, les conditions de leur
mise en place et leurs relations avec les microorganismes de leur environnement (ADOLPHE et
al. , 1995).

Jeanson, C., El ATssaoui, H. & Adolphe, J.-P., 1996. — Traces de l'activite de diplopodes dans des sols et des
sediments karstiques du Maroc atlantique. In: Geoffroy, J.-J., MaURiSs, J.-P. & Nguyen Duy - Jacquemin, M.. (eds),
Acta Myriapodologica. Mem. Mus. natn. Hist, nat .. 169 ; 555-560. Paris ISBN : 2-85653-502-X.

556

COLETTE JEANSON, HSAIN EL AISSAOUI & JEAN-PIERRE ADOLPHE

A cette occasion, nous avons egalement mis en evidence le role de myriapodes diplopodes
dans l'elaboration de telles concretions. Ce travail concerne des echantillons provenant de deux
des sites de l'etude entreprise sur les concretionnements carbonates dans le Haut-Atlas.

MATERIEL ET METHODES

Localisation des sites : Imouzzer et Tali wine

Le site d'lmouzzer fait partic du domaine des Ida-ou-Tanane, zone la plus montagneuse de l’Atlas atlantique, au cceur
d'un anticlinal incisE par l’Oued Tidili qui entame les calcaires du Lias superieur jusqu'au contact avec les argiles du Lias
inferieur.

La coupe type du “gisement” a concretions carbonatees est situEe sur la rive gauche de 1’oued Tidili k I km de la
cascade d'lmouzzer. Sur les collines, un paysage de glacis-cones ou les cavites (lapiEs) de 1 h 5 cm de profondeur sont
remplies d'un sol rouge rEsiduel a concretions en "cocons” de couleur allant du rouge au noir. Au fond de la combe, 1'oued
entaille des terrasses quaternaires cultivees ou les sols limoneux actuels contiennent aussi des concretions en forme de
“cocons”. Cylindriques ou cylindroconiques, leur taille varie de 1 mm h 2 mm de longueur.

Le site de Taliwine sur le flanc sud du Haut-Atlas occidental presente un “gisement” moins important que le
prEcEdent. La coupe representative est levee pres de Bouyalgua a 1 km a l'ouest de Taliwine.

Au laboratoire. Analyse morphologique, t nine ralogi que el geomicrobiologique.

Les Echantillons entiers sont photographies puis examines & la loupe binoculaire. Des lames minces y sont taillees aprEs
induration puis examinees au microscope polarisant pour I'identification des materiaux organiques et des mineraux.

Le microscope electronique a balayage apporte un complement d’informations au niveau ultramicroscopique sur I'etat de
surface des concretions et sur les microorganismes qui s’y developpent.

RESULTATS

Sur le terrain

A Imouzzer, les concretions sont localisees dans la partie marno-calcaire des coupes selon
un systeme lenticulaire a repartition discontinue. La taille des lentilles varie de 1 cm d'epaisseur
et 10 cm de longueur a 40 cm d'epaisseur et 150 cm de longueur. Ces dernieres sont situees au
sommet des coupes et parfois superposees par groupe de deux ou trois. Les concretions y sont
regroupees en amas irregulierement consolides par une matrice egalement marnocalcaire.

La majorite des lentilles de concretions est de couleur blanche et grise ; les autres sont
colorees de jaune a rouge-brun par des oxydes de fer. Souvent, la base des lentilles est tapissee
d'une croute pelliculaire de quelques millimetres (croute zonaire).

A la partie sommitale des memes coupes, un banc de calcaire dur de 30 cm environ est
surmonte d'un sol residuel de couleur rouge qui remplit des cavites de dissolution (lapies). Ce
paleosol renferme aussi des concretions cylindriques, cylindroconiques, voire ovoides de
couleur rouge a noire.

A Taliwine, les lentilles de concretions sont localisees au sommet de la coupe dans une
formation mamo-schisteuse. Elies ont de 1 a 10 cm d'epaisseur et de 20 a 150 cm de longueur et
sont disposees de fagon discontinue sans aucun regroupement particulier. Les concretions ont la
meme forme que les precedentes et sont de couleur grise et blanche.

Au laboratoire

A la loupe binoculaire, on note une homogeneite morphologique des concretions et une
variabilite de leur couleur et de leur taille. La forme est cyiindroconique, ovoi'de ou
subspherique ; une extremite est arrondie avec une legere depression centrale, l'autre presente
souvent une petite protuberance subtriangulaire legerement incurvee (Fig. 1). Leur taille varie de
1 mm x 0,5 mm a 2 mm x 1 mm. Dans 1'ensemble, celles d'lmouzzer sont plus grandes que
cedes de Taliwine.

La couleur des concretions varie avec le milieu ou elles se trouvent. Ainsi, dans les sols
rouges remplissant les cavites de dissolution (lapies), elles sont rouges ou noires. Dans les sols
cultives des terrasses, elles sont grises ou noires. Dans les lentilles enclavees dans les formations

Source : MNHN . Paris

ACTIVITF. DE DIPLOPODES DANS LES SOLS ET SEDIMENTS KARSTIQUES MAROCAINS

557

carbonatees, elles sont en general blanches mais il existe aussi quelques lentilles beige-rose ou
rouges colorees par les oxydes de fer.

Fig. 1. — Concretion - dejection cylindroconique de Glomeris sp. (Myriapode. Diplopode) ; longueur 1 h 2 mm ; schema
de synthese dc la surface d'apr£s photos & la loupe binoculaire et au microcope & balayage (m.e.b.) : 1. extremite
anterieure h protuberance conique. 2. extremite posterieure a depression centrale. 3. rainures et cannelures entre
les deux extremes. 4. traces de mycelium. 5. taches de carbonate de calcium ou d'oxydes ferriques. 6. cassure.
7. debris v£g6taux. 8. turricules de nematodes, ik la surface. 9. id. dans une cavitd. 10. canalicules, sillons, pistes
de nematodes. 11. stries obliques de frottement. 12. cristaux de calcite, en rosette, en baguette, au m.e.b.
13. bacteries filamenteuses et en spherules, au m.e.b.

FlG. 1. — Cylindroconic fiscal pellet of Glomeris sp. ( Diplopoda ); length: 1 to 2 mm. Synthetic drawing after
photographs f Stereomicroscope and SEM). /. anterior end with cone-shaped protuberance. 2. posterior end with
central depression. 3. grooves and striations between the two ends. 4. mycelium traces. 5. stains of calcium
carbonate or ferric oxydes. 6. break. 7. vegetal fragments. 8. nematod excreta, on the surface. 9. id., in a cavity \
JO. small grooves and nematod trails. 11. rubbing oblique scores. 12. rosette and stick-shaped calcite crystal,
SEM. 13. filamentous and spheruleous bacteria, SEM.

558

COLETTE JEANSON, HSAIN EL A1SSAOUI & JEAN-PIERRE ADOLPHE

La surface des concretions est ornementee de traces, d'empreintes, de cavites, de
minuscules monticules, de fins debris vegetaux. Des reseaux de fins sillons et des canalicules
ramifies, parfois colores en blanc ou en noir, s'insinuent entre des corpuscules spheriques isoles
ou regroupes par trois ou quatre. Les reseaux sont discontinus et ne couvrent qu'une partie de la
surface de la concretion. Une minuscule cavite est parfois presente, proche de 1'extremite
arrondie de la concretion et de tres petits turricules en gamissent souvent le fond (Fig. 1 [9]).

La microtopographie de la surface des concretions est aussi marquee par des cannelures et
des rainures plus ou moms rectilignes qui rejoignent les deux extremites a intervalle regulier
(Fig. 1 [3]). Les debris vegetaux ou leurs empreintes, des fragments translucides de filaments
myceliens sont nettement visibles. II existe aussi des microcassures, fractures, fissures parfois
remplies d'une substance noire (Fig. 1[6]) et de fines cristallisations en baguette et en rosette.

Autant de traces des etapes successives de la construction et de la mise en place de la
concretion, de l'impact de l'environnement et de 1'evolution consecutive qu'il va falloir
interpreter dans le temps et l'espace.

Au microscope polarisant, l'examen des lames minces fait apparaitre trois sortes de
concretions : minerales, organiques, organominerales.

Les concretions minerales sont blanches, constitutes de micrite, de microsparite, de rares
cristaux de quartz, de quelques constituants noirs amorphes de matiere organique decomposee.
La matrice est une calcite et l'ensemble de la lame presente une structure compacte et homogene.

Les concretions brun-noir contiennent des debris organiques allonges, souvent disloques,
de couleur noire, jaune ou brun-rouge ou les tissus vegetaux sont encore visibles. Quelques
cristaux de quartz et de micrite y sont parfois associes. Ces concretions sont poreuses et
l'ensemble de la lame presente une structure alveolaire peu induree.

Les concretions grises ont une composition mixte organo-minerale. La micrite et la
microsparite sont associees a une matiere organique amorphe et sans structure visible ou a des
fantomes de debris organiques. Ces concretions ont une structure relativement compacte.

Les concretions brun-rouge appartiennent a l'une des categories ci-dessus. Elies ont une
matrice coloree par les oxydes de fer et sont beaucoup plus dures que les precedentes et resistent
mieux a la pression

Au microscope electronique a balayage, la surface des concretions presente: 1) des
corpuscules spheriques de 0,1 pm a 1 pm en chapelet, 2) des enchevetrements de filaments
myceliens, 3) des reseaux de sillons superficiels stries perpendiculairement a leur axe. Ces
divers microorganismes et ces microtraces sont fortement mineralises et associes a des aiguilles
de calcite.

INTERPRETATION

Les concretions-agregats-dejections

Les concretions de ces sols et sediments sont comparables par leur taille, leur forme et
quelques traits de leur micromorphologie externe, aux agregats terreux decrits dans des sols
bruns forestiers (JEANSON, 1981). Des agregats identiques sont aussi construits par des
myriapodes diplopodes, Glomeris marginata eleves dans des microcosmes en conditions
controlees selon la methode de pedozoologie experimental (JEANSON, 1968).

D'autres types d’ agregats, construits par vingt cinq especes d'invertebres terricoles et
vasicoles actuels, ont ete selectionnes. Ces modeles sont proposes pour, eventuellement, servir a
l'interpretation des phenomenes de biostructuration et bioturbation dans les paleosols etudies en
Geologie de la Prehistoire (JEANSON, 1987).

Dans les sols actuels, des agregats analogues sont elabores et constants par des Glomeris
et sont en realite des dejections deposees dans les premiers centimetres. Le materiel consomme y
est encore bien visible : de nombreux fragments vegetaux sont reconnaissables a leur structure

Source :

ACTIVITE DE DIPLOPODES DANS LES SOLS ET SEDIMENTS KARSTIQUES MAROCAINS

559

cellulaire et souvent associes a une matiere minerale limono-argileuse. Ces agregats-dejections
sont ainsi des indices : par leur forme, des animaux terricoles qui les ont construits ; par leur
constituants, du milieu d'ou ils proviennent ; par l'etat de leur surface, de revolution de leur
environnement.

La couleur et l'etat de la surface des concretions peuvent ainsi etre interprets a la lumiere
des donnees biopedologiques actuelles :

- les concretions blanches sont formees surtout de carbonate de calcium, les noires, de
matiere organique humifiee, les grises, d'un melange des deux ;

- les concretions rouges, brunes, beiges sont colorees par des oxydes de fer a des degres
de concentration variables. Cette diversite de couleurs est a mettre en relation avec l'importance
du deplacement des oxydes de fer et leur accumulation dans les concretions.

Dans l'etat actuel de nos investigations, la chronologie de la coloration serait la suivante :
des concretions-dejections elaborees a partir du sediment du sol sont de la couleur brunatre de ce
dernier, par la suite elles blanchissent progressivement en se mineralisant par le carbonate de
calcium sous l'influence des bacteries calcifiantes. Les concretions impregnees d'oxydes de fer
prennent une coloration brun-rouge, parfois marquee de taches blanches, indices d'une
carbonatation en cours.

L'interpretation des microtraces de la surface des concretions est biologique ou
physicochimique.

Des cannelures joignent les deux extremites ; elles sont plus marquees sur la partie conique
des concretions et dues a l’empreinte de la partie terminale du tube digestif des Glomeris. La
partie conique emise en dernier correspond ainsi a la partie anterieure de l’animal. Cette extremite
est parfois erodee ou absente ce qui donne a la concretion une forme subovoide, et pourrait
constituer un indice de deplacement par ruissellement.

Les sillons marques en creux et peu ramifies pourraient etre des traces laissees par des
nematodes. Une trentaine d'especes de ces vers bacteriophages et mycophages peuvent proliferer
dans les dejections actuelles de Glomeris hexasticha (TAJOVSKY et al., 1992). Ils pourraient
aussi etre a l'origine des petits turricules et de la logette situee a l'extremite arrondie.

Les sillons peu marques et plus ramifies pourraient etre des traces laissees par des
filaments myceliens. En effet, dans des dejections actuelles de Glomeris marginata, une
quinzaine d'especes de champignons se succedent au cours de leur decomposition (NICHOLSON
et al, 1966). Les petits lambeaux transparents et luisants qui subsistent par endroits sur les
concretions-dejections pourraient etre des vestiges des reseaux myceliens. Ils sont parfois
colores en noir par la matiere organique issue de leur decomposition ou de substances
organiques qui ont migre. Ils peuvent etre aussi colores en blanc par le carbonate de calcium tres
visible sur les concretions ferruginisees. Les corpuscules spheriques en chapelet sont des
bacteries ou des cyanobacteries. Cette microflore est souvent associee a une forte mineralisation
caracteristique des depots microkarstiques et pedologiques. Les cristaux calciques en baguettes
ou en rosettes, a la surface des concretions-dejections, sont d'origine bacterienne (ADOLPHE et
al, 1989).

Les lentilles et amas de concretions

Les concretions carbonatees s'accumulent en amas en forme de lentilles, dans des fentes
ou des cavites du reseau karstique souvent delimitees par de fins depots calcaires en lamelles
rubanees.

L’hypothese proposee pour interpreter la genese de ces formations geologiques externes
lait appel a des phenomenes pedologiques, biologiques et climatiques : un couvert vegetal
forestier et une litiere abondante ont pu favoriser le developpement des populations de
diplopodes. Des phases de destruction de la foret ont pu favoriser l'erosion des sols et le

560

COLETTE JEANSON, HSAIN EL AISSAOUI & JEAN-PIERRE ADOLPHE

ruissellement, l'entramement et l'accumulation des dejections de diplopodes dans des zones
preferentielles du reseau karstique.

Les dejections riches en matiere organique deja decomposee et associee au mucus digestif
des diplopodes constituent des micro-milieux propices au developpement des microorganismes.
La circulation de solutions bicarbonatees et ferreuses induit ensuite une carbonatogenese
bacterienne et une concentration d'oxydes ferriques. Les dejections, d'abord organiques,
deviennent organominerales puis se transforment en concretions minerales, carbonatees et/ou
ferriques.

CONCLUSION

Les traces de l'activite des diplopodes dans des sols actuels et des sols experimentaux sont
materialisees par des biostructures caracteristiques ; des agregats terreux construits par ces
arthropodes sont en realite des dejections. Celles des Glomeris actuels sont comparables aux
concretions carbonatees rassemblees dans des formations lenticulaires de sediments karstiques
du Haut-Atlas Atlantique.

La nature, la forme, la structure, le degre d'alteration et l'agencement spatial des materiaux
mineraux et organiques de ces concretions-agregats-dejections permettent de proposer une
chronologie des phenomenes qui sont a l'origine de leur apparition dans l'environnement.

Un champ d'investigation interdisciplinaire entre les Sciences de la Terre et les Sciences de
la Vie, la “Pedozoologie”, a ainsi fourni des bases pour l'interpretation de faits
sedimentologiques et geologiques. Cette demarche pourrait etre appliquee a d'autres formations
sedimentaires et se baser sur d’autres analogies avec des biostructures construites par divers
animaux terricoles.

REMERCIEMENTS

Nous remercions vivemenl Jacques Rebiere. dessinateur au Laboratoire de Zoologie-Arthropodes (Paris), pour

sa realisation du schema de synthese.

REFERENCES

Adolphe, J. P.. 1981. — Observations et experimentations geomicrobiologiques et physico-chimiques des
concretionnements carbonates continentauxs actuels et fossiles. These Sciences, UPMC, Mem. Sci. Terre , Paris VI,
339 pp.

Adolphe, J. P., Hourimeche, A., Loubiere, J. F., Paradas, J. & Soleilhavoup, F.. 1989. — Les formations
carbonatees continentales d'Afrique du Nord. Bull. Soc. geol. Fr., (8), V : 55-62.

Adolphe, J. P.. El Aissaoui, H., Hourimeche, A., Paradas, J.. Soleilhavoup, F. & Jeanson, C. 1995. — Contribution
a 1 etude des biostructures. Excmple marocain. In : Bassins sedimentaires africains. Actes du 4eme Colloque de
Geologie africaine. [118eme congrds national des societes historiques et scientifiques. Pau, 25-29 octobre 1993],
Paris, Editions du CTHS : 259-268.

HOURIMECHE, A., 1988. — Etude sedimentologique et geomicrobiologique des depots quaternaires de la region
d'Essaouira ( Maroc ). These Sciences, UPMC. Paris VI, 222 pp.

Jeanson, C., 1968. — Essai de Pedozoologie experimental. Morphologie dun sol artificiel structure par les
Lombricides. Mem. Mus. natn. Hist, nat.. A, 46 : 21 1-357.

Jeanson, C., 1981. — Structuration du sol par la faune terricole. Incidences sur les concentrations organominerales. In :
Migrations organominerales dans les sols temperes. Colloque de Nancy 1979. Nancy, Editions du CNRS : 114-123.

Jeanson, C., 1987. — Biostructures construites par la faune dans des sols et des sediments actuels. Leur utilisation en
Prehistoire. In : Geologie de la Prehistoire , Paris, Geopre . 725-735.

Nicholson, P. B„ Bocock, K. L. & Heal, O. W., 1966. — Studies on the decomposition of the faecal pellets of a
millipede ( Glomeris marginata (Villers)). J. Ecol., 54 : 755-766.

Tajovsky, K., Santruckova, H., Hanel, L.. Balik, V. & Lukesova, A., 1992. — Decomposition of faecal pellets of
the millipede Glomeris hexasticha (Dipopoda) in forest soils. Pedobiologia ,36 : 146-158.

Source : MNHN, Paris

Feeding Rates and Nutrient Assimilation in the
Millipede Jonespeltis splendidus
(Diplopoda, Paradoxosomatidae)

Kubra BANO

Department of Zoology, University of Agricultural Sciences, G.K.V.K., Bangalore, 560065 India

ABSTRACT

Laboratory studies have been conducted on the feeding and egestion of the millipede Jonespeltis splendidus. The
millipedes were found highly specific in their diet. Ingestion and egestion rates varied with the component of the diet as
well as with the sex of the individuals. Acceptability of the food depended on the moisture content and the material
softness. Palatability was based on the nitrogen levels of the food. Egestion rates were directly proportional to the
quantity ingested. Further, the percentage of assimilation was found to be higher with litter. The nutrient uptake by the
millipede was recorded at the rate of 62, 26 and 12% with respect to proteins, fats and carbohydrates. With the feeding
recorded rates the consumption by the millipede was estimated to be about 4-5 grams of dry litter per year per millipede.
As the sampled population density ranged near 200 millipedes per square metre, the rate of the litter breakdown would
approach 1 kg per year, per suare metre.

RESUME

Taux d'ingestion et dissimilation chez le diplopode Jonespeltis splendidus (Diplopoda,
Paradoxosomatidae).

Des etudes en laboratoire ont etc menees sur la consommation et la rejection chez le diplopode Jonespeltis splendidus
qui montre une relation hautement sp6cifique visi-vis de son regime alimentaire. Les taux d'ingestion et d'egestion
varient avec la composition de la nourriture et avec le sexe des individus. L'app6tance pour la nourriture depend du taux
d’humidite et de la souplesse du materiel, de meme que de son contenu en azote. Le taux de consommation correspond a la
quantite de materiel ingdre ; toutefois, le pourcentage dissimilation parait plus eleve dans le cas de consommation de
litidre. On a enregistre pour le diplopode un apport nutritif de 62, 26 et 12% en rapport avec les protdines, les lipides et
les glucides. En fonction des taux d'ingestion mesures, la consommation par le diplopode est estimee a environ 4-5
grammes de litidre sdche par an et par individu. La densite de la population etant estimde a environ 200 diplopodes par
mdtre carre, la degradation annuelle de la litiere s'etablit autour de 1 kg par metre carre.

INTRODUCTION

Jonespeltis splendidus (Verhoeff), a soil and litter dwelling millipede, is normally found in
large populations under dead and decaying organic matter, under humus layers on the forest
floors, in small number underneath pots in the gardens, and under stones and logs where
moisture is conserved. It is also found to occur in cow dung pits. Large populations are always
associated with availability of organic matter and high moisture levels. Dry litter drives the

Bano, K., 1996. — Feeding rates and nutrient assimilation in the millipede Jonespeltis splendidus (Diplopoda,
Paradoxosomatidae). In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, not., 169 : 561-564. Paris ISBN : 2-85653-502-X.

562

KUBRA BANO

population away. It feeds on the substratum on which it lives and deposits pad like faecal matter
on the surface that can be easily picked up as crust.

Reports are available on the food consumption and assimilation in other millipedes (see,
among others, BLOWER, 1974; KAYED, 1978; BRUGGL, 1992). As J. splendidus is known to
be a regulator and decomposer in its habitat, it was decided to record its feeding rates and
nutritive requirements which were correlated to assess the fertility levels of its surroundings.

MATERIAL AND METHODS

The millipedes were collected from gardens and fields in and around Bangalore and were held in vivaria on moist
garden soil (alfisol), with a layer of moist decomposing organic litter.

Before the beginning of the present experiment, the millipedes were fed on wet filter paper to clean their guts.
Determination of feeding and defecation rates: one kg of finely sieved soil was spread uniformly in a glass trough
305 mm in diameter and 150 mm deep. The soils were moistened to 50% level.

100 adult millipedes, which were fed on wet filter paper earlier, were allowed to feed on the soils for four hours.
After four hours of feeding the millipedes were picked and transferred to clean Petri dishes (150 mm diameter) for two
hours. The covered Petri dishes were lined with wet filter papers for maintaining the humidity. The faecal pellets that
were deposited in the Petri dishes were collected. The feeding and egestion rates were determined gravimetrically.

Similar experiments were conducted with Mango leaf litter. 1 kg of fine sand was uniformly laid in each glass
trough and moistened to 50% level. About 200 g of wet decomposing mango leaf litter was deposited over the sand bed.
The experiment then went on as mentioned earlier.

The determination of feeding rates was carried with known quantities of moistened soil and decomposing litter
alone (separately) with known number of millipedes in small plastic boxes. Every day the material was weighed after
picking the millipedes and separating the faeces. The difference in weight between the feed material before and after
feeding was recorded as the consumption. The moisture level was maintained by covering the containers with lids lined
with wet polystyrene foam.

The containers used for the experiment were maintained at 25 ± 2°C. The experiments were conducted on round the
clock basis. In darkness red light was used to record observations.

Protein, carbohydrates and fats were analysed bio-chemically by adopting standard techniques.

RESULTS

The recorded food intake and defecation of adult millipedes in relation to sex (Fig. 1),
when soil was used as, the medium, are given in Table 1. The rate of consumption of the males
was higher than that of the females. The intake of soil was higher than that of the decomposing
litter in both sexes. The millipedes feed on the soft tissue of the litter, avoiding the veins and the
midrib, which are woody in nature. The intake of food was dependent on the nutritive value and
also on the microbial fauna of the medium, which was also reported by BLOWER (1974).

Table 1. — Feeding and excretory rates (mg/day) of Jonespeliis splendidus (Mean result for 100 individuals).

Sex

Males

Females

Feed

Soil

Mango litter

Soil

Mango litter

Ingestion

31.84

13.2

51.54

1 2.2

Egestion

24.83

3.3

31.95

4.5

% Assimilation

The active period of feeding was found to be the morning hours. The feeding rates
recorded during these hours are given in Table 2.

Table 3 presents the proximate composition of the mango leaf litter and the excrements of
the male and female millipedes. The composition of the excrements did not differ with reference
to sex. The excrements contained more nitrogen than carbohydrates. The fat contents were
sigmlicantly higher in the male excrements. This suggests that the female assimilated more fats
than the males, perhaps as their energy requirements are more important during ovulation.

Source : MNHN, Pahs

FEEDING RATES AND NUTRIENT ASSIMILATION IN A MILLIPEDE

563

Table 2. — Feeding and excretory rates of Jonespeltis splendidus with soils in mg. per hour (forenoon)

(Mean result for 100 individuals).

Hour

Consumption

9.6

6.6

1 1.33

6.6

5.3

8.2

Excretion

2.3

1.0

2.5

1.0

1.9

0.9

Table 3. — Composition of the mango leaf litter and faecal

pellets of the millipede Jonespeltis splendidus.

Material

mg / 100 mg dry mass (X ± S.D. ; n=4)

Nitrogen

Soluble Carbohydrates

Total fats

Dry mango leaf

4.34

4.50

7.54

Faeces (Female)

2.95 ± 0.47

1.83 ± 0.38

2.72

Faeces (Male)

2.41 ± 0.81

1.77 ± 0.39

3.32

Table 4 demonstrates the daily caloric intake of adults. More ingestion was noted in both
sexes for proteins than for fats and carbohydrates.

Table 4. — Caloric intake of Jonespeltis splendidus (Estimated).

Food

1st day

3rd day

Male

Female

Male

Female

Soluble Carbohydrates

4.7871

4.7709

1.0701

1.0683

Protein

31.9737

30.9207

8.3626

7.6960

Total fats

13.3851

13.6359

3.2368

3.2733

O SOIL CONSUMPTION BV ADULT MALES
• SOIL CONSUMPTION BY ADULT FEMALES
A EXCRETORY OUTPUT BY ADULT MALES
A EXCRETORY OUTPUT BY ADULT FEMALES

Fig. 1. — Soil ingestion and soil egestion by adult males and females of Jonespeltis splendidus , in relation to sex.

564

KUBRA BANO

DISCUSSION

Earlier investigations have revealed that the millipede Jonespehis splendidus is a
saprophagous macroarthropod feeding selectively on decaying plant organic matter, and
preferring Mango leaf litter, when choice is offered (BANO & KRISHNAMOORTHY, 1981). The
feeding activity depends on the nitrogen content of the food source, which was evident when
different types of litter, cow dung and soils were offered. In the present experiment, smaller
intakes were recorded with litter, and larger intakes were associated with soils in which organic
nitrogen and carbon contents were low. If the surface layer was covered with organic matter, the
activity of the millipedes was restricted to the surface. When soils alone were offered the
millipedes exhibited burrowing activity, which is indicative of foraging behaviour for organic
matter. The consumption of soil was important in the absence of leaf litter. These features allow
the millipedes to get their nutritive requirements. There was no significant difference in the litter
intake between the sexes. The mango litter consumption was more or less equivalent in both the
sexes, but the soil consumption varied. The assimilation varied with soil and litter. The energy
requirements of the millipedes were obtained mostly from proteins (62%), followed by fats
(26%) and carbohydrates (1%).

The role of arthropods and particularly diplopods in soil-litter system has been described in
various ways (cf. CRAWFORD, 1992). They are considered as accelerators, regulators and
decomposers. While working on the millipede J. splendidus the author has elaborated similar
interpretation for the role of this millipede (Bano & KRISHNAMOORTHY, 1976, 1977; BANO,
1992). The feeding activity involved destruction of litter as these millipedes feed on the soft
tissues, leaving the veins and the fibrous portions to decay further. They contribute to the
transformation of the organic constituents, improving the humic part of the soils. Their direct
effect is the acceleration of the formation of humus, and the indirect effect is the incrementing of
the microflora through their faecal pellets. With the present data it could be calculated that a
millipede consumes 4 to 5 g of leaf litter (dry mass) in a period of six months, which could be
taken as the active period of the millipede's life cycle. The distribution of the species varies in
relation to the nature of the soils and organic matter. A maximum of 200 individuals/m2 was
reported earlier (BANO & KRISHNAMOORTHY, 1985). This distribution would destruct about
1 kg of dry leaf litter per year, thus suggesting their role as decomposers in their habitats.
Similar results were obtained by BLOWER (1974) with Ophyiulus pilosus regarding the mass of
leaf consumed per year.

REFERENCES

Bano, K., 1992. — The role of the millipede Jonespehis splendidus (VerhoefO in an ecosystem (Diplopoda,
Polydesmida, Paradoxosomatidae). Ber. nat.-med. Verein Innsbruck , Suppl. 10 : 319-327.

Bano, K„ Bagyaraj, D. J. & Krishnamoorthy. R. V., 1976. — Feeding activity of the millipede Jonespehis
splendidus Verhoeff and soil humification. Proc. Indian Acad. Sci. (Animal Sci.), 83B : 1-11.

Bano, K. & Krishnamoorthy, R. V., 1977. — Changes in the composition of soils due to defecation by the millipede
Jonespehis splendidus (VerhoefO- Mysore J. Agric. Sci.. 11 : 561-566.

Bano, K. & Krishnamoorthy, R. V., 1981. — Consummatory responses of the millipede Jonespehis splendidus
(VerhoefO in relation to soil organic matter. Proc. Indian Acad. Sci. ( Animal Sci.) , 90 : 631-640.

Bano. K. & Krishnamoorthy, R. V., 1985. — Reproductive strategy and life history of Jonespehis splendidus
(VerhoefO (Diplopoda: Polydesmida) with environmental synchronisation. J. Soil Biol. Ecol.,5 : 48-57.

Blower, J. G., 1974. — Food consumption and growth in a laboratory population of Ophyiulus pilosus (Newport).
Symp. Zool. Soc. Land., 32 : 527-551.

Bruggle, K., 1992. — Gut passage, respiratory rate and assimilation efficiency of three millipedes from a deciduous
wood in the Alps (Julidae, Diplopoda). Ber. nat.-med. Verein Innsbruck, Suppl. 10 : 319-326.

Crawford, C. S., 1992. — Millepedes as model detritivors. Ber. nat-med. Verein Innsbruck, Suppl. 10 : 277-288.

Kayed, A. N., 1978. — Consumption and assimilation of food by Ophyiulus pilosus (Newport). Abli. Verb. Naturwiss
Ver. Hamburg , 21/22 : 115-120.

Source

Sexual Selection in Savanna Millipedes
Products, Patterns and Processes

Steven R. TELFORD * & John Mark DANGERFIELD **

* Department of Zoology, University of Pretoria, Pretoria 0002, South Africa
* Department of Biology, University of Botswana, P. Bag 0022, Gaborone, Botswana

ABSTRACT

The polygynandrous mating systems of savanna millipedes are a dynamic combination of simple effective male
mating tactics, male behaviours that protect their reproductive investment in females and possibly female choice of high
quality males. Sexual selection has led to the evolution of dimorphic characters that aid in courtship and mate
acquisition. Differential development between the sexes at the onset of breeding activity leads to changes over the
breeding season in the OSR, and a prevalence of alternative male mating tactics. Selection via sperm competition may
be responsible for the evolution of diverse and complex male gonopods (Barnett & Telford, this volume) and
behaviours such as prolonging the duration of copulation. For a morphologically simple and conservative class of
invertebrates, millipedes offer great potential for the study of sexual selection.

RESUME

Selection sexuelle chez les Diplopodes de savane : resultats, modalites et processus.

Lcs modes polygynandres d’accouplement chez les diplopodes de savane constituent une combinaison dynamique des
tactiques d’accouplement simples et efficientes des males, des comportements des males protegeant leur investissement
reproducteur aupres des femelles et d’un choix possible des femelles pour des males de grande qualite. La selection
sexuelle a conduit a Involution de caracteres dimorphiques facilitant les parades et I’accouplement. Le developpement
ditferentiel entre les sexes a l’approche de la phase d’activite de reproduction conduit a des modifications de la sex-ratio
operationnelle durant la saison de reproduction et avantage les tactiques alternatives d’accouplement des males. La
selection par la competition pour le sperme peut etre responsable de revolution de gonopodes males complexes et
diversities (Barnett & Telford, ce volume), ainsi que de comportements tels que celui de prolonger la duree de
copulation. Pour une classe d'invertebres a morphologic simple et conservatrice, les diplopodes offrent un large
potentiel de recherche dans le domaine de la selection sexuelle.

INTRODUCTION

For most higher organisms, males produce a surplus of cheap gametes capable of
fertilising an infinite number of females, while females produce a finite number of expensive
gametes that require only a single mating for effective fertilisation (BATEMAN, 1948; WILLIAMS,
1966). Because of this disparity in the cost of gamete production, males display indiscriminate
mating tactics, mate frequently and compete intensely with rivals, while females often exercise
highly selective mate choice.

Telford, S. R. & Dangerfield, J. M., 1996. — Sexual selection in savanna millipedes: products, patterns and
processes, In: Geoffroy, J.-J., Mauri£s, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta Myriapodologica. Mem.
Mus. natn. Hist, nat .. 169 : 565-576. Paris ISBN : 2-85653-502-X.

566

STEVEN R. TELFORD & JOHN MARK DANGERF1ELD

This fundamental behavioural difference between the sexes is the source of a selective
process described by DARWIN (1871) as sexual selection. Sexual selection acts on both
behavioural and morphological traits that confer a mating advantage to individuals with the most
elaborate variant of the trait. Sexual selection acts in two ways: competition between males for
access to females (intrasexual selection); competition between males, usually via display, to be
chosen by females (intersexual selection). One or both forms of selection are assumed to be
responsible for the evolution of the bright and often elaborate plumage of male birds and the
horns and antlers of antelope and deer (Darwin. 1871; TRIVERS, 1985) as well as numerous
other examples of pronounced sexual dimorphism (e.g. ALEXANDER et ai, 1979; EBF.RHARD,
1985). Recently, PARKER (1970) described a form of indirect intrasexual selection, namely
sperm competition, in which the sperm of different males compete to fertilise ova from within
the female. This process of ejaculate competition has proved to be widespread in both vertebrates
(BIRKHEAD & HUNTER. 1990; BlRKHEAD, 1989; GOMENDIO & ROLDAN, 1993) and
invertebrates (PARKER. 1970; WALKER, 1980; THORNHILL & ALCOCK, 1983; WAAGE, 1986)
with polygynandrous mating systems.

Over the past ten years much research has focused on teasing apart the relative
contributions of these selective processes to the evolution of mating systems in general (EMLEN
& ORING, 1977; BORGIA, 1979), and more specifically to understanding what determines
variation in male and female mating success (BROWN, 1988; CLUTTON BROCK, 1988; GRAFEN,
1988).

The study of animal mating systems aims to describe who mates with who, how often,
and why. More specifically, researchers wish to identify the selective pressures responsible for
the evolution of sexual dimorphism and differences in behaviour (product), determine potential
morphological and behavioural correlates of variation in mating success (pattern), and identify
the tactics of mate acquisition and the degree of competition therein (process). In this paper we
summarise our work to date on the mating systems of southern African savanna millipedes using
the themes of product, pattern, and process. We highlight aspects of the work which deserves
further consideration and suggest future directions for the study of millipede mating systems.

PRODUCTS

Morphology

Southern African juliform millipedes display marked sexual dimorphism, with females
typically heavier and wider, but not longer than their conspecific male counterparts (TELFORD &
DANGERFIELD, 1990, 1993a, b). The ovaries of females are paired structures housed in a
common median ovitube that runs from ring 15 to the last podous ring (see BLOWER, 1985;
HOPKIN & READ, 1992). Our measures of clutch size for savanna millipedes range from 200-
800 ova over a size range of 1 .0-20.0 grams (n = 15 species, unpublished data).

A cylindrical body plan is a unifying feature of juliform millipedes. The volume of a
cylinder is calculated as: h.7cr2, where h is the height of the cylinder (body length) and r the
radius. If, for example, selection for increased fecundity resulted in a twofold increase in radius
the resultant increase in volume could only be matched through a fourfold increase in length. The
energetic costs associated with increasing body diameter are probably less than would be
associated with increasing body length. Presumably natural selection has acted on female body
volume in this way to accommodate the ovary and maximise the number of eggs it can contain.

Body volume is not the only sexually dimorphic character. Millipede legs are longer and
broader in males compared to females and have adhesive pads on the tarsal segments. The
evolution of this sex difference in morphology probably relates to courtship and copulation (see
HOPKIN & READ, 1992). Our observations have shown that males of all taxa (except members
of the genera Calostreptus and Chersastus) walk along the back of the female prior to engaging
in copulation. The longer broader legs of males with their adhesive pads presumably aid in this

Source :

SEXUAL SELECTION IN SAVANNA MILLIPEDES

567

manoeuvreing and may also function in stimulating sexual receptivity in females. Both natural
and sexual selection may have contributed to the evolution of sexual dimorphism in limb
structure. Data on sexual dimorphism in Alloporus uncinatus together with the probable sources
of selection are summarised in Table 1.

Table 1. — Sexual dimorphism in Alloporus uncinatus. Leg width measured in micrometer units. Significance levels:
***P<0.001, **P<0.01. N.S. = Natural Selection; S.S. = Sexual Selection. Source: Modified from Telford &
Dangerfield (1990).

Males

Females

t-test

Selective Pressure

Body mass (g)

8.0 ± 0.11

8.7 ± 0.13

-4.25***

110

N.S.

Body length (mm)

114.1 ± 0.65

110.7 ± 0.87

3.16**

110

N.SAS.S.

Body width (mm)

8.9 ± 0.06

9.5 ± 0.08

-6.41***

1 10

N.S.

Leg length (mm)

5.9 ± 0.10

4.9 ± 0.80

-7.83***

S.S.

Leg width (^im)

15.9 ± 0.16

13.4 ± 0.26

-8.18***

S.S.

In many sexually dimorphic species, males are typically the larger sex because sexual
selection favours larger males for their superior competitive ability in contests with rivals
(Thornhill & ALCOCK, 1983; TRIVERS, 1985). Body design in male millipedes may reflect
their greater mobility. Mobility seems to be an important determinant of mate acquisition (see
Behaviour section below) with both longer legs and longer body resulting in males being able to
move more quickly and efficiently than females. This idea remains to be tested.

Table 2. — Sex differences in ihe behaviour of three species of juliform millipede expressed as the percentage of the
total number of observations (values in brackets). Expected frequencies for between sex comparisons are based on the
overall sex ratio for the species and the total number of observations in the behaviour category. Source: modified from
Dangerfield, Milner & Matthews, (1992).

Walking

Feeding

Resting

Copulatin

Calostreptus carinatus

males (62)

64.5

1 1.3

9.7

14.5

females (223)

25.1

34.1

36.8

4.0

22.0***

8.6**

10.9***

8.4*

Chaleponcus digitatus

males (148)

66.2

23.0

10.1

0.7

females (212)

34.9

42.0

22.6

0.5

17.5***

9.7**

7.9**

0. 1 ns

Alloporus uncinatus

males (86)

69.8

18.6

3.5

8.1

females (158)

47.5

40.5

7.6

4.5

4.9*

7.9**

2.6ns

1 .3ns

568

STEVEN R. TELFORD & JOHN MARK DANGERFIELD

Behaviour

Savanna millipedes are surface active during the summer wet season and, depending on
habitat and rainfall patterns, may remain active for as long as six months (DANGERFIELD &
Telford, 1991; Dangerfield, Milner & Matthews, 1992; Telford & Dangerfield,
1993a). During this period adults, intercalary males and juveniles emerge from underground to
feed and reproduce. Significant differences between the sexes are apparent in the time invested in
the performance of these behaviours as well as in their general activity patterns (Table 2). For
example, in a population of Alloporus uncinatus from Mazowe, Zimbabwe (17°30’S, 30°57'E)
males were more active than females, who spent more time feeding in aggregates, and climbing
vegetation; apparently to avoid interference from the more mobile mate-seeking males
(Dangerfield & Telford, 1992; Telford & Dangerfield, 1993a). In a small patch of
Acacia savanna in Gaborone, Botswana (24°40'S, 25°52'E) there were significant differences
between the sexes of three species in time spent walking, feeding, resting and copulating
(Table 2, and see DANGERFIELD, MILNER & MATTHEWS, 1992).

Males tend to be the more mobile sex and invest most of their time searching the habitat for
females. Females invest more time in feeding, probably to repay their significant energetic
investment in ova.

PATTERNS

Mating patterns

Correlating variation in mating success with some measure of behaviour or morphology is
the standard protocol for detecting patterns of non-random mating (e.g. PARTRIDGE &
Halliday, 1984). Non-random mating is assumed to be the consequence of some process of
sexual competition in which individuals of superior competitive ability gain a mating advantage
over rival conspecifics.

In a study of mating patterns in the Mazowe population of Alloporus uncinatus over a
single breeding season (November 1988 - May 1989) mating was random with respect to body
mass (g): unmated males (mean ± 1 s.e.) = 8.16 ± 1.08, n = 868; mated males = 8.24 ± 1.05,
n = 295. MANN-WHITNEY U-test; U = -1.2, P>0.1, and see Fig. 2 in TELFORD &
DANGERFIELD (1993a). A lack of any significant correlations between various measures of the
body sizes (mass, length, width, head width) of males and females in copula pairs supported
this conclusion (see Fig. 1). However, this evidence for random mating is not conclusive as the
sampling protocol did not take into account the mating histories of individuals over the duration
of the breeding season. This highlights the problems of studying mating success in natural
populations of small, highly abundant, inconspicuous organisms. The focus of such studies
must be at the level of the individual and requires the use of effective, reliable marking
techniques.

The operational sex ratio

Another powerful predictor of local mate competition is the operational sex ratio (OSR),
which is the ratio of sexually receptive males to females (EMLEN & ORING, 1977). Under male
biased OSR conditions males are limited in their access to females and must compete for females
as a limiting resource. The reverse is true under female biased OSR conditions.

Savanna millipede communities are characterized by large-scale temporal variation, and sex
differences, in emergence and activity patterns (DANGERFIELD & TELFORD, 1991;
Dangerfield, Milner & MATTHEWS, 1992) with the nett effect of changing OSR conditions
over the breeding season.

Source :

SEXUAL SELECTION IN SAVANNA MILLIPEDES

569

2 - * - ‘ - *

2 6 10 14

female body mass (g)

Fig. 1. — The relationship between body mass (g) of males
and females in all mated pairs of Atloporus uncinatus
collected over a single breeding season. Arrow
indicates a pair in which the female received a fatal
injury. Source: Telford & Dangerfield, 1993a.

Fig. 2a. — Seasonal change in the operational sex ratio (•)
and the occurrence of triplet associations as a
proportion of the number of mated pairs (O) for the
Mazowe population of Alloporus uncinatus. Source:
modified from Telford & Dangerfield, 1993a.

The magnitude and timing of change
in the OSR varies between populations and
is also dependent on the duration of the
breeding season. For example, in the
Mazowe population of A. uncinatus the
OSR was female biased at the onset of
breeding. This bias gradually declined,
ending in a peak of male bias coincident
with the major peak in mating activity and
the appearance of an alternative male mating
tactic (Fig. 2a, and see TELFORD &
Dangerfield, 1993a). This process of
change in the OSR is a consequence of the
presence of large numbers of intercalary
males early in the season, and the gradual
decline in number of mature females who
presumably burrowed back into the soil to
lay egg clutches.

Botswana populations of A .
uncinatus, Calostreptus carinatus and
Chaleponcus digitatus showed similar
though less predictable change in the OSR
(Fig. 2b). In a millipede community at
Richards Bay, Natal (28°37'E, 32°17’S)
females of the dominant species Chersastus
sanguinipes were present constantly, but
males only appeared immediately after a
rainfall event, stayed for 2-3 days and then
disappeared. Copula pairs were only
observed during the times when males were
present (Rudi Van Aarde, pers. comm.).

In millipede breeding populations, the
co-occurrence of alternative male mating
tactics is a common observation under male
biased OSR conditions. Instead of always
searching for single females, males often
joined copula pairs to form triplet
associations. This observation will be
discussed further in the following section.

PROCESSES

Mating experiments

Separating the effects of male-male
competition from female choice remains one
of the major challenges to students of sexual
selection (see for example, KIRKPATRICK,
1982; ARNOLD, 1983; PARTRIDGE, 1983).
Previously we argued that our field data on
mating patterns were insufficient to refute

Source MNHN, Paris

570

STEVEN R. TELFORD & JOHN MARK DANGERF1ELD

the occurrence of non-random mating.

Fig. 2b. — Seasonal change in the OSR for Botswana populations of AUoporus uncinatus (•) Calostreptus carinatus (A)
and Chaleponcus digitatus (O).

Fig. 3. — The relationship between body mass (g) and
gonopod distal width for a sample of male AUoporus
uncinatus from Mazowe.

Here we summarise results from
controlled laboratory mating experiments
designed to test for size assortative and size
selective mating, and separate the effects of
inter and intra-sexual selection in generating
these mating patterns (TELFORD &
DANGERFIELD, 1993b). Sequential choice
experiments were conducted with AUoporus
uncinatus, Calostreptus carinatus,
Spinotarsus tenuis, Chaleponcus digitatus
and Chaleponcus limbatus. Individual male
and female mating histories for
A. uncinatus from Mazowe (Fig. 3) and the
two species of Chaleponcus revealed that,
in the absence of a choice situation, mating
was size selective (Table 3) and best
explained by female choice (TELFORD &
DANGERFIELD, 1993a, b). In the other
species and populations mating was
random.

In multiple choice mating experiments
a wide variety of outcomes were observed
(Table 4) including size assortative and

random mating, and a mating advantage for
both large and small males. In addition, between population comparisons of mating pattern were
not always consistent. Our results suggest that both processes of sexual competition operate in
millipede mating systems but do not always correspond for geographically separate populations
of the same species (Table 5). These results should viewed as a source of testable a priori
predictions about between-species and between-population differences in the competitive
processes that generated these observed mating patterns.

Source ;

SEXUAL SELECTION IN SAVANNA MILLIPEDES

571

Table 3. — Regression statistics for the relationship between frequency of acceptance and male body mass (g) for males
in sequential choice mating experiments. Source: Telford & Dangerfield (1993b).

Regression statistics

Species

Population

Sample size

intercept

slope

A. uncinatus

Mazowe

245

-6.10

0.77

0.503***

Hwange

-0.43

0.49

0.212ns

Calostreptus

Hwange

2.50

0.78

0.054ns

carinatus

Sengwa

1.93

1.48

0.095ns

Chcileponcus

V. Falls

-5.54

8.60

0.500**

limbatus

Marondera

-5.43

5.14

0.677***

Chaleponcus

Marondera

-8.77

8.77

0.583**

digitatus

Spinotarsus

Marondera

2.06

0.42

0.001ns

Table 4. Acceptance by females of first, second and third rank males as first and second mating partners in multiple
choice mating experiments and the proportion of females that accepted a second male (PA). Source: modified from
Telford & Dangerfield (1993b).

Species

Population

1st rank
males

2nd rank
males

3rd rank
males

A. uncinatus

Mazowe

1st

7.85*

2nd

2.00ns

Calostreptus

Hwange

1st

1 .40ns

carinatus

2nd

2.00ns

Sengwa

1st

5.60*

2nd

1 .00ns

Chaleponcus

V. Falls

1st

1.08ns

limbatus

2nd

Marondera

1st

6.19*

2nd

Chaleponcus

Marondera

1st

6.37*

digitatus

2nd

2.48ns

Spinotarsus

Marondera

1st

8.00**

tenuis

2nd

1 .00ns

Spinotarsus

Marondera

1st

9.39**

cuspidosus

2nd

Source : MNHN , Paris

572

STEVEN R. TELFORD & JOHN MARK DANGERF1ELD

MATING TACTICS

Intrasexual competition

Observations of courtship and copulation as well as mate seeking behaviour in both the
field and laboratory provide further insight into the tactics of mate acquisition and processes of
sexual competition employed by millipedes. Competition between males can be both direct and
indirect. The primary mating tactic appears to be a scramble for mates in which males seek
females through a random search of the habitat (TELFORD & DANGERFIELD, 1993a). When
conspecifics are encountered males attempt copulation and tend to be indiscriminate.
Interindividual differences in time spent searching and speed of movement may generate variance
in male mating success and is, therefore, a form of indirect competition. Under male biased OSR
conditions single males often join with copula pairs to form triplet associations (Fig. 2a, and see
TELFORD & DANGERFIELD, 1990, 1993a) and either wait for the pair to separate before
attempting to mate with the female (Spirostreptidae) or actively attempt to displace the copulating
male (Odontopygidae). Attempted displacement of copulating males is a common mating tactic in
invertebrate and anuran mating systems (THORNHILL & ALCOCK, 1983; LAMB, 1984; TELFORD
& VAN SICKLE, 1990) and is a form of direct male competition.

Why males switch from searching for single females to associating with copula pairs when
the OSR is male biased is an interesting unanswered question. We suggest that this change in
behaviour does not reflect inferior competitive ability; as is the case for males of many species
that perform alternative mating tactics (KREBS & DAVIES, 1987). A more parsimonious
explanation is that as the OSR becomes increasingly male biased, encounter frequency with
solitary females declines. Males then simply associate with the first female they meet regardless
of whether she is paired or alone. This behaviour is “tolerated” by copulating males because they
would have to release the female in order to repel the other male. One reason for this apparent
tolerance of copulating males is because the physical nature of copulation (TELFORD & WEBB, in
prep.) together with its long duration under male control (TELFORD & DANGERFIELD, 1993b, c)
creates a potential sexual conflict of interest in which the male stands to gain more than the
female.

Prolonged copulation is advantageous for males because it protects their reproductive
investment in females under conditions of intense intrasexual competition (TELFORD &
DANGERFIELD, 1991). The potential evolutionary benefits to females are less obvious (see
THORNHILL & ALCOCK, 1983). Females can incur physical damage during copulation (see
Fig. 1) which may be compounded through multiple mating (see for example, FOWLER &
PARTRIDGE, 1989). Therefore, if a male releases a female to repel a rival male she may become
unwilling to resume copulation. However, this still begs the question of why females remate
with the second male in triplet associations. While the evolutionary benefits a male enjoys from
multiple mating are obvious and well documented the same is not true for females (THORNHILL
& ALCOCK, 1983). An often argued potential benefit of multiple mating by females is the
increased heterozygosity in their offspring; this prediction still lacks empirical verification.

Copulation in spirostreptid millipedes occurs either in parallel ( Calostreptus ), head to head
(Rhodesiostreptus matabele), or with the male coiled around the female (TELFORD &
DANGERFIELD, 1990, 1993b). Interestingly, parallel copulators tend to mate for a shorter time
period than coiled copulators. Chaleponcus sp. has the shortest copulation duration for a coiled
copulator and single males of this species actively attempt to displace copulating males.
However, attempted displacement has never been observed in parallel copulators which also
typically perform short duration copulations. The significance of between species variation in
copulation duration remains unclear, and definately warrants further study. Data on copulation

Source :

SEXUAL SELECTION IN SAVANNA MILLIPEDES

573

position, duration and displacement, copulatory guarding, and triplet formation are summarised
in Table 6.

Table 5. Predictions of expected mating patterns and processes of sexual competition in the mating systems of nine
populations of spirostreptid millipede. Predictions are based on results obtained from sequential and multiple choice
mating experiments (tables 3 & 4). * Small male mating advantage, cf-o' Comp.= male-male competition.

Sequential Choice Multiple Choice

Species _ Population Pattern _ Process _ Pattern Process

A. uncinatus

Mazo we

Hwange

Size-

Selective

Random

9 Choice

Size-

Selective

Size-

Assortative

riuucss

9 Choice
c T-d* Comp.

9 Choice

Calostreptus

carinatus

Hwange

Random

Size-

Assortative

9 Choice

Sengwa

Random

Size-

Selective

9 Choice

Chaleponcus

limbatus

V. Falls

Size-

Selective

9 Choice

Random

Marondera

Size-

Selective

9 Choice

Size-

Selective

9 Choice
cf-d* Comp.

Chaleponcus

digitatus

Marondera

Size-

Selective

9 Choice

Size-

Selective

9 Choice
cT-d" Comp.

Spinotarsus

tenuis

Marondera

Random

Size-

Selective

9 Choice
cf-cf Comp.

Spinotarsus

cuspidosus

Marondera

Size-

Selective

9 Choice
d’-d’ Comp.

Prolonged copulation in invertebrates appears to have evolved as a form of mate guarding
in which males protect their reproductive investment in females by limiting their opportunity to
remate (see THORNHILL & ALCOCK, 1983). In Alloporus uncinatus, males alter the duration of
copulation according to predictions of the mate-guarding hypothesis (TELFORD &
DANGERFIELD, 1993 b, c). Sperm competition ( sensu PARKER, 1970) is the most likely process
responsible for selection favouring the evolution of this form of mate guarding by males. This
may also be true for other species of millipede with prolonged copulation (see Table 6).

Intersexual competition

In some invertebrates, primary or accessory genitalia scale positively with body size
(EBERHARD, 1985; unpublished data for eight species of spirostreptid millipedes). In libellulid
dragonflies, the hook-like structures at the end of the penis scale positively with body size and
both size and degree of symmetry correlate with sperm volumes removed (MILLER, 1991).
EBERHARD (1985) has suggested that females could use such a scaling relationship as a cue to
male body size and use the information to choose large males as mates. We have already
suggested that female choice may partially explain the mating advantage enjoyed by large males

574

STEVEN R. TELFORD & JOHN MARK DANGERFIELD

of several species in sequential and multiple choice mating experiments (Table 5). However,
invoicing genitalic scaling as the mechanism of female choice may be incorrect.

For example, although the gonopods of A. uncinatus scale positively with body size
(Fig. 3), the sizes of males and females in copula pairs from Mazowe do not correlate. If
females do choose males on this basis then we would predict a positive relationship between the
size of mating partners, or a correlation between mating success and body size in natural
populations. To more fully understand this potential mechanism of female choice requires a
systematic study of the ultrastructure of male and female genitalia (see BARNETT & TELFORD,
this volume) together with a study of mating patterns in populations of known individuals.

Table 6. — Copulation duration in minutes (mean ± 1 s.d.) for eight populations of spiroslreptid millipede (sample
sizes are given in brackets). Copula positions are given below species names. Source: modified from Telford &
Dangerfield (1993b), and unpublished data.

Family / Species

Population

Copulation

duration

Triplets

(Y/N)

Displacement

(Y/N)

Spirostreptidae

Alloporus uncinatus
(coiled)

Mazowe

122.7 ± 49.4
(35)

Hwange

205.8 ± 60.8
(25)

Calostreptus sp.

Hwange

60.3 ± 25.6(22)

(parallel)

Sengwa

33.8 ± 22.9
(25)

Odontopygidae

Chaleponcus sp. 1
(coiled)

Marondera

22.6 ± 17.9
(40)

Chaleponcus sp.3
(coiled)

V. Falls

80.2 ± 25.3
(20)

Marondera

85.6 ± 16.6
(19)

Chaleponcus digitatus
(coiled)

Marondera

66.0 ± 11.2
(28)

Spinotarsus tenuis
(coiled)

Marondera

92.3 ± 18.6
(19)

CONCLUSIONS

Millipedes are a conspicuous group of organisms, present in a wide variety of habitats and
typically occurring at high population densities. They adapt well to laboratory conditions and are
useful subjects for experimental manipulation.

Our study of the mating system of Alloporus uncinatus over a single breeding season
revealed the nature of male mating tactics, the impact of change in the OSR on male mating
tactics, and the role of stochasticity in generating the observed mating pattern. Our comparative
data from laboratory mating experiments suggests that both male competition and female choice
shape mating systems and that the relative roles of the two processes can differ between
populations of the same species. Between population plasticity in mating systems is believed to
be a consequence of the combined effects of environmental variability, differing population
densities and operational sex ratios. Together, these results and predictions can be used to
generate a priori hypotheses about plasticity in millipede mating systems, testable through
detailed longitudinal studies of natural populations.

Source :

SEXUAL SELECTION IN SAVANNA MILLIPEDES

575

Descriptive studies, for example between species variation in the duration of copulation,
are a necessary starting point for the investigation of the adaptive significance of any behaviour
pattern. We have shown experimentally, that in A. uncinatus prolonging the duration of
copulation is a form of mate guarding (TELFORD & DANGERFIELD, 1991) and our comparative
data allow us to make similar predictions for other species. Where copulation duration is not
prolonged the mate guarding explanation does not hold but nevertheless makes way for
alternative a priori predictions to be made. For example, if copulation duration is short it may be
because female remating frequency is low. Therefore, selection to guard females is relaxed and a
better tactic for males is to reduce time spent in copula and maximise mating frequency.

The data presented here confirm the polygynandrous nature of savanna millipedes which,
together with the capacity of females to store sperm and the functional role of gonopods in sperm
displacement (BARNETT & TELFORD, this volume), highlight the importance of sperm
competition in millipede mating systems. Quantifying sperm precedence patterns is essential to a
complete understanding of the relative contributions of different processes of sexual competition
to observable variation in male and female mating success.

REFERENCES

Alexander, R. D„ Hoogland, J. L„ Howard, R. D„ Noonan, K. M. & Sherman, P. W„ 1979. — Sexual dimorphism
and breeding systems in pinnipeds, ungulates, primates and humans. In : N. A. Chagnon & W. Irons. Evolutional
biology and human social behaviour: An anthropological perspective. North Scituate.

Arnold, S. J., 1983. — Sexual selection: The interface of theory and empiricism. In : P. Bateson, Mate Choice
Cambridge, Cambridge University Press : 67-108.

Barnett, M., Telford. S. R. & de Villiers, C. J., 1993. — Sperm displacement in a millipede? - an investigation into
the genital morphology of the southern African Spirostreptid millipede Orthoporus pyrhocephalus. J. Zool. Lond.,
3 311.

Bateman, A. J., 1948. — Intrasexual selection in Drosophila. Heredity. 2 : 349-368.

BlR35^AD T R 1989 _ ThC inlel,igenl sPerm? A conc'se review of sperm competition. J. Zool. Lond.. 218 : 347-

Birkhead, T. R. & Hunter, F. M., 1990. — Mechanisms of sperm competition. Trends Ecol. Evol. , 5 : 48-52.

Boake, C. R. B.. 1986. — A method for testing adaptive hypotheses of mate choice. Am. Nat., 127 : 654-666.

Borgia, G., 1979. — Sexual selection and the evolution of mating systems. In: N. S. Blum & M. A. Blum .Sexual
selection and reproductive competition in insects. New York, Academic Press.

Blower, J. G., 1985. — Millipedes. [In : D. M. Kermach & R. S. K. Barnes, Synopses of the British Fauna.. 351.
London, E. J. Brill. & W. Backhuys, 242 pp.

Brown, D., 1988. — Components of Lifetime Reproductive Success. In : T. H. Clutton Brock, Reproductive Success.
Chicago, University of Chicago Press : 439-453.

Clutton Brock, T. H., 1988. — Reproductive Success. In: T. H. CLUTTON Brock, Reproductive Success. Chicago,
University of Chicago Press : 472-485.

Dangerfield, J. M. & Telford, S. R.. 1991. — Seasonal activity patterns of Julid millipedes in Zimbabwe. J. Trop.
Ecol., 1 : 281-285.

Dangerfield, J. M. & Telford, S. R.. 1992. — Species diversity of Juliform millipedes: Between habitat comparisons
within the seasonal tropics. Pedobiologia ,36 : 321-329.

Dangerfield, J. M., Milner, A. E. & Matthews, R., 1992. — Seasonal activity patterns and behaviour of juliform
millipedes in south-eastern Botswana. J. Trop. Ecol.,S : 451-464.

Darwin, C., 1871. — The descent of man and selection in relation to sex. London, John Murray.

Eberhard, W. G., 1985. — Sexual selection and animal genitalia. Cambridge: Harvard University Press.

Emlen, S. T. & Oring, L. W., 1977. — Ecology, sexual selection and the evolution of mating systems. Science , 197 :
215-223.

Fowler, K. & Partridge, L., 1989. — A cost of mating in female fruitflies. Nature , 338 : 760-761.

Grafen, A., 1988. — On the uses of Data on Lifetime Reproductive Success, hi : T. H. Clutton Brock. Reproductive
Success.. Chicago, University of Chicago Press : 454-471.

Gomendio, M. & Roldan, E. R. S., 1993. — Mechanisms of sperm competition: Linking physiology and behavioural
ecology. Trends Ecol. Evol., 8 : 95-100.

576

STEVEN R. TELFORD & JOHN MARK DANGERFIELD

HOPKIN, S. P. & Read, H. J., 1992. — The biology of millipedes . Oxford, Oxford Univ. Press, 233 pp.

Kirkpatrick, M., 1982. — Sexual selection and the evolution of female choice. Evolution , 36 : 1-12.

Krebs, J. R. & Davies, N. B., 1987. — An introduction to behavioural ecology. Oxford, Blackwell Scientific
Publications.

Lamb. T., 1984. — Amplexus displacement in the southern toad Bufo terrestris. Copeia : 1023-1025.

Miller. P. L.. 1991. — The structure and function of the genitalia in the Libellulidae (Odonata). Zool. J. Linn. Soc.,
102 : 43-73.

Parker, G. A., 1970. — Sperm competition and its evolutionary consequences in the insects. Biol. Rev., 45 : 525-
567.

Partridge, L., 1983. — Non random mating and offspring fitness. In : P. Bateson, Mate Choice. Cambridge,
Cambridge University Press : 67-108.

Partridge, L. & Halliday, T. R.. 1984. — Mating patterns and mate choice. In : J. R. Krebs & N. B. Davies,
Behavioural ecology: An evolutionary approach. Oxford. Blackwell Scientific Publications : 222-250.

Telford, S. R. & Dangerfield. J. M.. 1990. — Sex in millipedes: laboratory experiments on sexual selection. J. Biol.
Ed., 24 : 233-238.

Telford, S. R. & Dangerfield, J. M., 1991. — Sex ratio manipulation and copulation duration in the tropical
millipede, Alloporus uncinatus: A test of the copulatory guarding hypothesis. Anim. Behav ., 40 : 984-986.

Telford, S. R. & Dangerfield, J. M., 1993a. — Mating tactics in the tropical millipede Alloporus uncinatus
(Diplopoda; Spirostreptidae). Behaviour, 124 : 45-57.

Telford. S. R. & Dangerfield, J. M., 1993b. — Mating behaviour and mate choice experiments in some tropical
millipedes (Diplopoda: Spirostreptidae). S. A. J. Zool.. 28 : 1 55.

Telford, S. R. & Dangerfield, J. M.. 1993c. — Males control the duration of copulation in the tropical millipede,
Alloporus uncinatus. S. A. J. Zool., 29 : 266.

Telford, S. R. & Van Sickle, J., 1989. — Sexual selection in an African toad (Bufo gutteralis ): The roles of
morphology, amplexus displacement and chorus participation. Behaviour, 110 : 62-75.

Thornhill, R. & Alcock, J. 1983. — The evolution of insect mating systems. Cambridge, Massachusetts, Harvard
University Press.

Trivers, R. L., 1972. — Parental investment and sexual selection. In : B. Campbell, Sexual Selection and the Descent
of Man. Chicago, Aldine : 136-179.

Waage, J. K.. 1986. — Evidence for widespread sperm displacement ability among Zygoptera (Odonata) and the means
for predicting its presence. Biol. J. Linn. Soc., 28 : 285-300.

Walker, W. F., 1980. — Sperm utilisation strategies in nonsocial insects. Am. Nat., 115 : 780-799.

Williams, G. C., 1966. — Adaptation and Natural Selection. Princeton, NJ, Princeton University Press.

Source : MNHN, Paris

Trophic Preferences of Three Soil Macroarthropods

(Preliminary Study)

Jorge P. CANCELA DA FONSECA & Leila MEZIANE

Analyse des Systemes Ecologiques, Ecologie du Sol, C.N.R.S.

Universite Paris 7, Laboratoire de Biologie vegetale et d'Ecologie forestiere, F-77300 Fontainebleau, France

ABSTRACT

The main objective of this study is to compare the trophic behaviour of two macroarthropod species belonging to two
different invertebrate groups - Diplopoda and Isopoda - but which are, apparently, morphologically similar and have a
similar defence, armadillo behaviour (roller species): Glomeris marginata (Villers) and Armadillidium vulgare (Latreille).
Oniscus asellus Linn6, a dinger species, was also taken into account. These species coexist in the soil of Fontainebleau
and Montmorency Forests. They feed on beech, oak and chestnut litter. Three classes of leaf litter were tested out: I.
Litter of the year; dark leaves, thick, with few or no rotting spots; II. Old litter; dark leaves, thick but thinner than in I.,
with light rotting spots; and, III. Old litter; bleached, thin leaves. A Student-t test of the data shows that the three
species have a similar trophic behaviour concerning the chestnut litter, but a different one concerning the litter of both
beech and oak.

RESUME

Preferences alimentaires de trois macroarthropodes edaphiques (etude preliminaire).

Le principal objectif de cette etude est de comparer le comportement trophique de deux esp£ces de macroarthropodes
appartenant & deux groupes differents d'invertebres - Diplopoda et Isopoda - mais qui sont, en apparence,
morphologiquement semblables et ont un comportement de defense similaire en se roulant en boule : Glomeris marginata
(Villers) et Armadillidium vulgare (Latreille). Oniscus asellus Linn6, isopode depourvu de capacite de volvation, a ete
aussi pris en compte. Ces especes coexistent dans les sols des forets de Fontainebleau et de Montmorency. Elies se
nourrissent de liti&re de hetre, de chene et de chataignier. On a teste trois classes de litiere : I. Litiere de 1'annee ; feuilles
sombres, epaisses, sans ou avec peu de taches de pourriture blanche ; 11. Litiere des ann6es pr6cedentes ; feuilles sombres,
6paisses, mais plus minces qu'en I., avec des taches claires de pourriture blanche ; et. III. Litiere des annees prScedentes ;
feuilles minces, blanchies. L'application aux donnSes du test-t de Student a montre que les individus de ces trois especes
d'arthropodes presentent un comportement trophique similaire en ce qui concerne la litiere de chataignier, mais diff6rent
vis-a-vis des liti£res de hetre et de chene.

INTRODUCTION

The main objective of this study was to compare the trophic behaviour of two soil
macroarthropod species belonging to two different invertebrate groups, Diplopoda and Isopoda,
but which are, apparently, morphologically similar and have a similar rolling defence,
“armadillo” behaviour: Glomeris marginata (Villers) and Armadillidium vulgare (Latreille). Few
studies compare these two macroarthropod species. In general, they compare either the two

Cancela da Fonseca, J. P. & Meziane, L., 1996. — Trophic preferences of three soil macroarthropods
(Preliminary study). In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M„ (eds), Acta Myriapodologica.
Mem. Mus. natn Hist . nat ., 169 : 577-584. Paris ISBN : 2-85653-502-X.

578

JORGE P. CANCELA DA FONSECA & LEILA MEZIANE

isopods A. vulgare and Oniscus asellus Linne, or the latter species with G. marginata (e.g.
HARTENSTEIN, 1964; NEUHAUSER & HARTENTSTEIN. 1978; HASSALL & RUSHTON, 1984;
Ineson & ANDERSON, 1985; SUTTON & HARDING, 1989). One of the few authors that
compared these two species of isopods with one species of Glomeris, not G. marginata but G.
connexa Koch, was DUNGER (1958). However, these three species coexist in the same
ecosystem we studied several years ago. a beech woodland: La Tillaie in Fontainebleau Forest
(MEZIANE, 1976; CANCELA DA FONSECA & MEZIANE, 1978). This is why O. asellus is also
taken into account.

Our aim was to study comparatively the ecological niches of two roller species coming
from two different arthropod classes (Crustacea and Diplopoda) and their differential roles on the
breakdown of forest litter. For this, one of the points was to detail the trophic preferences of
such species. This preliminary work presents some significant results related to litter
preferences, useful for the comprehension and development of future studies dealing with
comparative ecological importance of individuals and populations forming such “functionnal
macroarthropod groups”.

MATERIAL AND METHODS

The two roller species studied here are A. vulgare (AVU) and G. marginata (GMA), and the dinger species,
O. asellus (OAS).

All were present in both the Fontainebleau and Montmorency Forest ground floors. They feed on litter. Three
types of litter were given to them in our experiments: Beech litter (Fagus sylvatica Linne - FSY), Oak litter (Quercus
sessilijlora Salisbury - QSE) and Chestnut litter ( Castanea saliva Miller - CSA). Three classes of litter were used under the
experimental conditions : 1. Litter of the year, autumn 1976; dark leaves, thick, with few or no rotting spots; II. Old
litter of the years before 1976; dark leaves, thick but thinner than in I„ with light rotting spots; and. III. Old litter of the
years before 1976; bleached, thin leaves. Five replicates were done with a number of leaves variable according to the
number of leaves available in each class. The emphasis was put on the beech litter. The experiments were made at room
conditions of about 15-I7°C and 80% R.H. from May to November 1977. Each adult, after 48 hours with no food
(fasting), was put in contact with each type of litter for a period of 2 days.

The consumption rates (in %) were evaluated by assessing the area of the leaves eaten by each adult in relation to
the total leaf area available. The numerical results (Table 1, Fig. 1) were analysed by the Student-t test (Table 2).

RESULTS

First ol all, the three species, but mainly the millipede G. marginata and the woodlouse
O. asellus, have a clear preference to the chestnut litter (classes I, II and III), and a significant
avoidance, less for G. marginata, of the beech litter of the year (class I). However, the most
important antagonistic difference of the feeding behaviour between the millipede G. marginata
and the woodlouse A. vulgare concerns the almost complete avoidance of, respectively, the oak
and the beech litter of the year (class I). The same kind of opposition concerned their preferences
to the old, thick beech litter (class II). In relation to G. marginata, O. asellus has a trophic
behaviour similar to that of A. vulgare, except for the old, thick beech litter (class II).

Furthermore, A. vulgare seems to prefer the thick chestnut and oak litter to the beech one
(classes I and II), while G. marginata seems to prefer the beech litter to the oak litter of the year
(class I), the chestnut litter to the oak and the beech litter of the year (class I), and the old, thick
chestnut litter to the old, thick beech litter (class II). O. asellus prefers the chestnut and the oak
litter ot the year to the beech one (class I), and the whole old chestnut litters to those of beech
(classes II and III).

Comparing the frequency profiles of the data some “odd” data were excluded. This mainly
increased the significant differences already observed. However, some significant differences
appeared, but above all for the classes with rather few data.

Source : MNHN, Paris

TROPHIC PREFERENCES OF THREE SOIL MACROARTRHROPODS

579

Table 1. — Mean consumption rates (%) of litter classes by Glomeris marginata, Armadillidium vulgare and Oniscus
asellus. X = Corrected means.

Tree species

Litter

classes

No. of
leaves

marginata

x±sx

No. of
leaves

vulgare

X±sx

O. asellus

No. of

leaves x±sx

F. sylvatica

36. 7± 7.4

14. 5± 4.0

11.8+ 2.0

8.0± 1.9

46. 3± 5.8

29. 3± 4.7

43.61 4.4

IIx

24. 6± 3.9

45.81 4.3

III

27. 1 ± 1 5.5

47 . 8± 1 1 .8

53.6110.5

IIIx

13.4± 8.9

53.8±1 1 .7

60.3+ 9.1

Q. sessiliflora

7.3± 4.7

52.2±10.6

48.1+ 8.7

3.0± 2.1

58.2±10. 1

1 1

54.2±14.3

1 1

52. 2± 8.9

54.41 7.4

IIx

60. 1±14.7

62. 7± 8.6

57.2+ 7.3

III

42. 6± 13.8

5 1 .2±1 5.8

64.3123.1

IIIx

5 1 . 1±1 3.3

60.9±1 5.3

87.3+ 4.7

C. saliva

82. 1± 7.3

56. 6± 9.5

68.91 9.1

51. 8± 8.9

75.71 6.6

78. 1± 8.2

66. 9± 9.6

75.8+ 9.6

IIx

1 1

85. 2± 4.5

III

76. 2± 1 9. 1

71.1111.4

83.51 7.6

IIIx

95. 3± 2.0

88.91 6.1

DISCUSSION AND CONCLUSION

In the beech woodland of “La Tillaie” (Fontainebleau Forest) of the three macroarthropod
species, A. vulgare was dominant (72%) followed by O. asellus (15%) and G. marginata (13%)
(February 1972-January 1973; MEZIANE, 1976). Their coexistence in time and space, measured
in terms of “activity behaviour” by pitfall trap method, was more important for G. marginata
and A. vulgare (SCHOENER's index Rt=0.636 and Rs=0.746) than for G. marginata and O.
asellus (Rt=0.583 and Rs=0.604), while, for A. vulgare and O. asellus, it was somewhat higher
in time (Rt=0.652) and lower in space (Rs=0.563). Their spatial distribution was also different:
more random for G. marginata (negative binomial distribution parameter k=8.48), more
aggregative for O. asellus (k=0.62), and for A. vulgare in between (k=3.21). Though the
surface of the site studied was not very large, its central part was not covered by herbaceous
vegetation, only by beech litter, but their peripheric borders have a great number of
mesohabitats, like fallen beech trunks and branches, decayed logs, small grassy patches, several
holly bushes, and a small grassland glade. Thus, as the three species were present everywhere
and coexist in this site, they can easily overlap part of their ecological niches. Nevertheless, they
had some habitat preferences: G. marginata for the uncovered litter, O. asellus for the decayed
wood places, and A. vulgare for the grassy patches and the small grassland glade. It is well
known that G. marginata prefers woodland to grassland soils where it inhabits very often with
A vulgare, which prefers them, and that O. asellus prefers woodland soils and decayed wood
(WARBURG, 1968; WALLWORK, 1976; RUSHTON & HASSALL, 1983; HASSALL & RUSHTON,
1984; Sutton & Harding, 1989).

Source :

580

JORGE P. CANCELA DA FONSECA & LEILA MEZJANE

Table 2. — Significant trophic preferences between Glomeris marginata, Armadillidium vulgare and Oniscus asellus.
In brackets: a) Classes of litter; b) Corrected means.

Significance: n.s. = not significant; * = 0.05>P>0.01; ** 0.01>P>0.001; *** = PcO.OOl.

Species

Type of preferences

Consumption rates {%)

P<5%

G. marginata

.Over A. vulgare :

Thick BEECH litter of the year (I)

36.7 vs

14.5

(36.7 vs

8.0

***>

Thick, old BEECH litter (II)

46.3 vs

29.3

(46.3 vs

24.6

**,

.Over 0. asellus :

Thick BEECH litter of the year (I)

36.7 vs

11.8

* *

.Thick BEECH litter of the year (I)

36.7 vs

7.3

vs thick OAK litter of the year (I)

(36.7 vs

3.0

.Thin, old OAK litter (III)

42.6 vs

27.1

n.s.

vs thin, old BEECH litter (III)

(51.1 vs

13.4

.Thick CHESTNUT litter of the year (I)

82.1 vs

36.7

* *

vs thick BEECH litter of the year (I)

.Thick, old CHESTNUT litter (II)

78.1 vs

46.3

* *

vs thick, old BEECH litter (II)

(85.2 vs

46.3

***)

.Thin, old CHESTNUT litter (III)

76.2 vs

27.1

n.s.

vs thin, old BEECH litter (III)

(95.3 vs

13.4

***)

.Thick CHESTNUT litter of the year (I)

82.1 vs

7.3

***

vs thick OAK litter of the year (I)

(82.1 vs

3.0

***)

.Thin, old CHESTNUT litter (III)

76.2 vs

42.6

n.s.

vs thin, old OAK litter (III)

(95.3 vs

51.1

.Thick, old BEECH litter (II)

46.3 vs

27.1

n.s.

vs thin, old BEECH litter (III)

(46.3 vs

13.4

.Thick, old OAK litter (II)

54.2 vs

7.3

vs thick OAK litter of the year (I)

(60.1 vs

3.0

***)

.Thin, old OAK litter (III)

42.6 vs

7.3

vs thick OAK litter of the year (I)

(51.1 vs

3.0

**)

Species

Type of preferences

Consumption rates (%)

P<5%

A. vulgare

.Over G. marginata :

Thick OAK litter of the year (I)

52.2 vs 7.3

* * *

(58.2 vs 3.0

***)

Thin, old BEECH litter (III)

47.9 vs 27.1

n.s.

(53.8 vs 13.4

.Thick OAK litter of the year (I)

52.2 vs 14.5

***

vs thick BEECH litter of the year (I)

(58.2 vs 8.0

**»)

.Thick, old OAK litter (II)

52.2 vs 29.3

* *

vs thick, old BEECH litter (II)

(62.7 vs 24.6

***)

Source

TROPHIC PREFERENCES OF THREE SOIL M ACROARTRHROPODS

581

A. vulgare

.Thick CHESTNUT litter of the year (I)

56.6 vs

14.5

***

(continued)

vs thick BEECH litter of the year (I)

(51.8 vs

8.0

***,

.Thick, old CHESTNUT litter (11)

66.9 vs

29.3

* *

vs thick, old BEECH litter (II)

(66.9 vs

24.6

***)

.Thick, old BEECH litter (II)

29.3 vs

14.5

vs thick BEECH litter of the year (I)

(24.6 vs

8.0

**>

.Thin, old BEECH litter (III)

47.8 vs

14.5

* *

vs thick BEECH litter of the year (I)

(53.8 vs

8.0

*«*>

.Thin, old BEECH litter (III)

47.8 vs

29.3

n.s.

vs thick, old BEECH litter (II)

(53.8 vs

24.6

•*>

Species

Type of preferences

Consumption rates (%)

P<5%

0. asellus

.Over G. marginata :

Thick OAK litter of the year (I)

48.1 vs

7.3

* *

(48.1 vs

3.0

***)

Thin, old BEECH litter (III)

53.6 vs

27.1

n.s.

(60.3 vs

13.4

**)

.Over A. vulgare :

Thick, old BEECH litter (II)

43.6 vs

29.3

(45.8 vs

24.6

***)

.Thick OAK litter of the year (I)

48.1 vs

11.8

* * *

vs thick BEECH litter of the year (I)

.Thick CHESTNUT litter of the year (I)

68.9 vs

11.8

* * *

vs thick BEECH litter of the year (I)

(75.7 vs

11.8

***)

.Thick, old CHESTNUT litter (II)

75.8 vs

43.6

vs thick, old BEECH litter (II)

(75.8 vs

45.8

.Thin, old CHESTNUT litter (III)

83.5 vs

53.6

vs thin, old BEECH litter (III)

(88.9 vs

60.3

.Thick CHESTNUT litter of the year (I)

68.9 vs

48.1

n.s.

vs thick OAK litter of the year (I)

(75.7 vs

48.1

.Thick, old BEECH litter (II)

43.6 vs

11.8

***

vs thick BEECH litter of the year (I)

(45.8 vs

11.8

***)

.Thin, old BEECH litter (III)

53.6 vs

11.8

* * *

vs thick BEECH litter of the year (I)

(60.3 vs

11.8

***)

.Thin, old OAK litter (III)

64.3 vs

48.1

n.s.

vs thick OAK litter of the year (I)

(87.3 vs

48.1

Fontainebleau Forest is mainly a beech-oak forest while Montmorency Forest has also chestnut
woods. This is why our experimental trophic research concerned principally beech litter and
secondly oak and chestnut litter. Laboratory results showed significant trophic differences
between the three macroarthropod species in relation to the consumption rates of beech and oak
litter. No significant differences being observed in relation to the consumption rates of chestnut
litter. Thus, the pill millipede, G. marginata, avoids the thick oak litter of the year (which was
also observed by GEOFFROY et al., 1987) more intensely than the two woodlice, A. vulgar e and

Source

582

JORGE P. CANCELA DA FONSECA & LEILA MEZIANE

Fig.

80 -

CSE CSA

TREE SPECIES

— Consumption rates (%) of three classes of litter (I, II, III) by one pill-millipede (C. marginata) and two
woodlice (A. vulgare and O. asellus ) adult individuals fed on Fagus sylvalica (FSY), Quercus sessiliflora (QSE) and
Castanea saliva (CSA) leaves.

Source : MNHN , Paris

TROPHIC PREFERENCES OF THREE SOIL MACROARTRHROPODS

583

O. asellus , avoid the thick beech litter of the year. These two species have a similar trophic
behaviour against that of G. marginata behaviour which is also showed by their preferences to
the thin, old beech litter. However, some significant differences were observed between them,0.
asellus preferring the thick, old beech litter more than A. vulgare. Moreover, the three species
prefer the chestnut litter to the other types of litter. Though, this was also observed by
ANDERSON (1973), it seems no to be directly attribuable to the nitrogen contents, the C/N ratio
or even the polyphenol contents of the leaf litter. The same was pointed out by NEUHAUSER &
Hartenstein (1978) which indicates however that Fagus and Quercus litter are “scarcely
palatable” to A. vulgare and O. asellus. Nevertheless, BECK & BRESTOWSKY (1980) say that
O. asellus grew better on freshly fallen leaves of beech and oak than on overwintered ones
which contradicts DUNGER (1958) and PlEARCE (1989) observations. In our essays, they
preferred significantly the thick oak litter of the year to the thick beech litter of the year, contrary
to G. marginata. It seems, however, that the pill millipede is not very common on the beech
woods (WALLWORK, 1976), but more common in mixed beech-oak woodlands when oak
leaves form part of the litter (VAN DER Drift, 1951). In any case, all species preferred old litter
to freshly fallen one. The freshly fallen leaves have normally high polyphenol, like lignin, and
tannin contents which inhibit their feeding by the animals (MILLER & CAMERON, 1983;
RUSHTON & HASSALL, 1983; HASSALL & RUSHTON, 1984; GUNNARSSON, 1987;
MOCQUARD et al.9 1987; JAMBU et al., 1988).

Besides, it is well known that microorganisms are able to degrade the phenolic and tannin
compounds of the leaves, and by that way to render them more palatable to the animals
(DUNGER, 1958; HASSALL & RUSHTON, 1984; GUNNARSSON, 1987; BlGNELL, 1989). This
can justify the preferences for the old litter, but the preferences for the litter of the year need a
more detailed biochemical research.

REFERENCES

ANDERSON, J. M., 1973. — The breakdown and decomposition of sweet chestnut ( Castcinea saliva Mill.) and beech
( Fagus sylvatica L.) leaf litter in two deciduous woodland soils. II. Changes in carbon, hydrogen, nitrogen and
polyphenol content. Oecologia ( Bert. ), 12 : 275-288.

Beck. L. & BRESTOWSKY, E., 1980. — Auswahl und Verwertung verschiedener Fallaubarten durch Oniscus asellus
(Isopoda). Pedobiologia.lt) : 428-441.

Bignell, D. E., 1989. — Relative assimilation of i^C-labelled microbial tissues and i4C-plant Fibre ingested with leaf
litter by Glomeris marginata under experimental conditions. Soil Biol. Biochem.. 21 : 819-827.

Cancela da Fonseca, j. P. & MEZIANE L.. 1978. — Macroarthropodes : abondance relative et activity saisonni^re de
quelques groupes (Isopodes, Diplopodes, Chilopodes et Opilions). [hi : Lemee G., La hetraie naturelle de
Fontainebleau.] In : F. BourliEre & M. LAMOTTE, Problemes d'Ecologie : Structure et fonctionnement des
ecosystemes terrestres. Paris, Masson : 116-119.

Dunger W.. 1958. — Uber die Zersetzung der Laubstrcu durch die Boden-Makrofauna im Auenwald. Zool. Jb. (S\st.),
86: 139-180.

Geoffroy, J. J., CElerier, M. L., Garay, I., Rherissi, S. & Blandin, P., 1987. — Approche quantitative des fonctions
de transformation de la matiere organique par des Macroarthropodes saprophages (Isopodes et Diplopodes) dans un
sol forestier a moder. Protocoles experimentaux et premiers resultats. Rev. Ecol. Biol. Sol , 24 : 573-590.

Gunnarsonn, T., 1987. — Selective feeding on a maple leaf by Oniscus asellus (Isopoda). Pedobiologia . 30 : 161-165.

Hartenstein, R., 1964. — Feeding, digestion, glycogen and the digestive system in Oniscus asellus. J. Insect Physiol..
10 : 611-621.

Hassall, M. & Rushton, S. P., 1984. — Feeding behaviour of terrestrial Isopods in relation to plant defenses and
microbial activity. Symp. zool. Soc. London. 53 : 487-505.

Ineson, P. & Anderson, J. M., 1985. — Aerobically isolated bacteria associated with the gut and faeces of the litter
feeding macroarthropods Oniscus asellus and Glomeris marginata. Soil Biol. Biochem.. 17 : 843-849.

Jambu, P., Juchault, P. & Mocquard, J. P.. 1988. — Etude experimental de la contribution du crustace isopode
Oniscus asellus a la transformation des litieres forestieres sous chene sessile. Pedobiologia , 32 : 147-156.

Meziane, L., 1976. — Activite saisonnidre de quelques groupes de Macroarthropodes. Memoire de DEA d’Ecologie
animale, Universite Paris VI, 54 pp.

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JORGE P. CANCELA DA FONSECA & LEILA MEZIANE

Miller. R. H. & Cameron. G. N.. 1983. — Intraspecific variation of life parameters in the terrestrial Isopod,
Armadillidium vulgare. Oecologia (Berl.).S 7 : 216-226.

Mocquard. J. P., Juchault, P.. Jambu, P. & Fustec, E., 1987. — Essai devaluation du role des crustaces oniscoi'des
dans la transformation des litieres vegetales dans une foret feuillue de 1'ouest de la France. Rev. Ecol. Biol. Sol . 24 :
31 1-325.

Neuhauser, E. F. & Hartenstein. R.. 1978. — Phenolic content and palatability of leaves and wood to soil isopods and
diplopods. Pedobiologia. 18 : 99-109.

PlEARCE, T. G., 1989. — Acceptability of pteridophyte litters to Lumbricus lerresiris and Oniscus asellus, and
implications for the nature of ancient soils. Pedobiologia , 33 : 91-100.

Rushton, S. P. & Hassall. M., 1983. — Food and feeding rates of terrestrial isopod. Armadillidium vulgare (Latreille).
Oecologia (Berl.),Sl : 415-419.

Sutton, S. L. & Harding, P. T.. 1989. — Interpretation of the distribution of terrestrial Isopods in the British Isles.
Monitor. Zool. ital. (N.S.) Monogr.,4 : 43-61.

Van Der Drift. J., 1951. — Analysis of the animal community in a beech forest floor. Tijdschr. Em. , 94 : 1-168.
Wallwork, J. A.. 1976. — The distribution and diversity of soil fauna. London, Academic Press, 355 pp.

Warburg, M. R., 1968. — Behavioural adaptations of terrestrial isopods. Am. Zool., 8 : 545-559.

Source : MNHN , Paris

Ecology and Behaviour of Xanthodesmus physkon
(Attems, 1898), an Aggregating Paradoxosomatid from

Tropical West Africa

Dieter MAH SB ERG

Department of Animal Ecology & Tropical Biology (Zoology III), Biocenter
University Wurzburg, Am Hubland, D-97074 Wurzburg, Germany

ABSTRACT

Xanthodesmus physkon is widely distributed from Central Africa to Liberia. The ecological studies presented here were
conducted in the Comoe National Park/Ivory Coast, where a diverse millipede fauna is under research. In addition,
experiments on laboratory bred animals are reported. X. physkon is restricted to the galery forest and is ecologically
separated from the sympatric Habrodesmus duboscqui) a soil-dwelling paradoxosomatid occurring in the adjacent
savanna. Both species live in aggregations which are formed with the onset of the newborns' surface activity; adults live
solitarily. Aggregations of X. physkon comprise 150 to 700 individuals; after fusion, they may consist of 10,000
specimens. Disturbing or hurting of only one individual instantly leads to a dissolving of aggregations for a certain
time and corroborates the anti-predatory function of this behaviour. The millipedes avoid spots where a conspecific was
disturbed. Escape and avoiding behaviour are elicited by secretions of the defensive glands. The large number of X.
physkon-sclcriies in the food middens of Paltothyreus tarsatus points to this abundant stink ant as a possible predator
of these millipedes but attacks were rarely observed in the field. In the laboratory, stink ants mostly avoid contact with
live X. physkon. X. physkon are mainly found on the bark of trees where they exclusively feed on algae. Tree species
with a high density of algae are prefered. The activity of X. physkon is restricted to the the most humid periods of the
rainy season (April to August). Aggregations in the laboratory in Germany which were permanently kept under
favourable conditions (high soil humidity, 27°C, D:L = 12:12), showed an extraordinary activity pattern: surface
activity and moulting cycles among individuals were highly synchronized, with distinct peaks of activity in the
intermoult phases. One period of total inactivity of six months occurred. The activity patterns of separately kept
aggregations were nearly coincident, with a deviation of only a few days. This is a hint at an endogenous rhythm in the
activity of X. physkon which adjusts it to the seasonal climate of its tropical habitat.

RESUME

Ecologie et comportement d’un Paradoxosomatidae gregaire d’Afrique tropicale occidentale :
Xanthodesmus physkon (Attems, 1898).

X. physkon s’etend de l'Afrique centrale au Liberia. Des recherches 6cologiques ont ete menees dans le Parc National
de la Comoe (Cote d'Ivoire) ou la faune de Diplopodes est tr£s diversifiee, ainsi qu’en laboratoire. X. physkon se
cantonne dans la foret-galerie et se distingue ecologiquement d 'Habrodesmus duboscqui , un Paradoxosomatidae edaphique
sympatrique qui se rencontre dans la savane adjacente. Les deux especes prSsentent un mode de distribution agregatif dont
les amas se constituent lors du d6but d’activite de surface de la nouvelle generation, les adultes vivant en solitaire. X.
physkon s'agrege en amas composes de 150 a 700 individus; apr£s fusion, les agregats peuvent reunir 10000

Mahsberg, D., 1996. — Ecology and behaviour of Xanthodesmus physkon (Attems, 1898), an aggregating
paradoxosomatid from tropical West Africa. In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen Duy - Jacquemin, M., (eds),
Acta Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 585-586. Paris ISBN : 2-85653-502-X.

586

DIETER MAHSBERG

specimens. Le fait de perturber ou de blesser un seul d'entre eux provoque une dissolution des agregats et vient corroborer
la fonction aniiprtklation de ce comporlement. D’ailleurs, ces Diplopodcs dvitcnt les endroits ou un groupe conspecifique
a ete perturbe. De tels comportements de fuite ou d’evitement sont provoques par les s£cr£tions £mises par les glandes
defensives. Une grande quantity de sclerites de X. physkon rencontres dans les residus alimentaires de la fourmi-cadavre
Paltothyreus tarsatus montre que cette fourmi peut 6ventuellement jouer un role de predateur sur les populations de
Diplopodes. Toutefois, les attaques directes sont tres rarement observees sur le terrain. Au laboratoire, ces fourmis
evitent la plupart du temps tout contact avec les individus vivants de X. physkon. Ces derniers se rencontrent
principalement sous les dcorces, ou il se nourrissent exclusivement d'algues. On note une preference marquee pour les
essences qui accueillcnt une forte densitc d'algues. L’activite de X. physkon est limit^e aux p^riodes les plus humides de
la saison des pluies (avril h aout). Les agr£gations obtenues en laboratoire en Allemagne ont ete maintenues en
permanence dans des conditions favorables (humidite du sol elevee, 27°C , D:L = 12:12). Elies montrent des modalit6s
d’activite extraordinaires : l’activite de surface et les cycles de mue entre les individus sont hautement synchronises,
presentant des pics d’activit^ distincts au cours des phases d’intermue. Une periode d’inactivite totale s’6tend sur six
mois. L'activite d'agregats eleves separement est en coincidence presque totale, avec une marge de difference de quelques
jours seulement. Ceci am£ne a penser que le rythme endogene de l’activite de X. physkon s'ajuste bien aux phases
climatiqucs saisonnieres de son environnement tropical.

Deplacements en masse dans le sud-est de la France
chez Ommatoiulus sabulosus (Myriapoda, Diplopoda,
Julidae) avec invasions d'habitations

Frangois Sahli

Museum National d'Histoirc Naturelle, Zoologie/Arthropodes, 61, rue de Buffon, F-75231 Paris Cedex 05
& Laboratoire Souterrain du C.N.R.S., F-09200 Moulis, France

RESUME

Deux cas sonl decrits et analyses : 1'un concerne la Provence en 1987. 1’autre les Alpes-Maritimes cn 1988. Ces cas.
personnel lement observes in situ , ont trait a Ommatoiulus sabulosus. L'auteur cherche a repondre & di verses questions et a
interpreter le phenom&ne des rassemblements et des migrations en masse, en tenant compte des connaissances actuelles
sur la periodomorphose.

ABSTRACT

On mass migrations with dwelling invasions in Mediterranean Ommatoiulus sabulosus (L.)
(Myriapoda, Diplopoda, Julidae) in the south-east of France.

The two mass migrations (MM) observed at the Val de Sibourg (Provence) and at Peillon (Alpes-Maritimes) are the
result of a double overpopulation - viz. in spring and in a same place - (a) of countless “tall individuals” (most of them
being post-adults), with, in addition, (b) countless very young larvae, which originated from egg depositions during the
previous autumn. The presence of shelter sites (or relay shelters) seems to have play a role in Peillon. In South of France,
mass migrations with dwelling invasions can only occur if a village or a part of it (often a recent construction) is set up
in the garrigue or more or less near a large garrigue. The year of the mass migration, spring rains are a sine qua non
condition. In addition to spring rains of the year y, heavy rains during the previous autumn seem to have play an
important role in the case of Peillon. Finally, the influence of predators and parasites on (a) the population which got
adult in 1984 and on (b) the individuals born in 1984 (which gave adults in 1986 or 1987) seems to have been poor. Such
situations - combined with the adoption of "explosive” juvenile to adult maturation moult strategies several times
(instead of spreading “CAT” strategies) - allowed a first demographic explosion in 1984, followed by other explosions
in 1986, 1987 at Sibourg and 1987, 1988 at Peillon.

INTRODUCTION

Les deplacements en masse de diplopodes sont connus depuis la fin du siecle dernier. Ils
presentent deux cas extremes : (1) ceux effectues par un tres grand nombre d'individus (jusqu’a
des centaines de milliers), (2) les deplacements ne comportant qu'un petit nombre d'individus.
Les premiers seront appeles “migrations en masse" (MM). Les seconds, appeles migrations ou
deplacements tout court, seront designes par M. Dans les cas extremes, la limite entre les deux
types est nette. II existe neanmoins des cas ou la separation est floue et devient affaire

Sahli, F., 1996. — Deplacements en masse dans le sud-est de la France chez Ommatoiulus sabulosus (Myriapoda,
Diplopoda, Julidae) avec invasions d’habitations. In: Geoffroy, J.-J., MAURlfes, J.-P. & Nguyen Duy - Jacquemin, M..
(eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat ., 169 : 587-598. Paris ISBN : 2-85653-502-X.

588

FRANCOIS SAHL!

^appreciation subjective. Parmi les MM, on distingue celles comportant l'invasion d'habitations
humaines et cedes ne la comportant pas.

Chez O. sabulosus , on se doit de mentionner tout d'abord une migration en masse en
Alsace (France) en 1900 (VERHOEFF, 1900). Des milliers de O. sabulosus se sont deplaces sur
une voie ferree et sont parvenus a bloquer un train dont les roues patinerent. Dans les annees
1950. un deplacement de O. sabulosus a ete observe dans la region de Cologne en Allemagne
(SEIFERT, comm. pers.). En 1973, un autre deplacement important eut lieu en Sarre (Allemagne)
(HELB, 1975) et a fait la une des journaux : les sarrois du village d'Ensdorf (non loin de
Sarrelouis) - particulierement ceux du nouveau lotissement de l'epoque - furent parait-il
epouvantes de trouver des iules dans leurs assiettes, sur leurs tables de cuisine ou dans leurs lits,
ce qui nous semble avoir ete exagere par la presse. En Provence (France), le seul cas signale est
celui mentionne par DEMANGE (1960. 1963) dans les Alpes-de-Haute-Provence.

II faut insister sur le fait que toutes les invasions de O. sabulosus qui se sont produites en
France et en Allemagne dans le passe n'ont jamais ete observees directement par les auteurs qui
en ont parle dans leurs publications ; de plus, la biologie s. 1. et les cycles des O. sabulosus
etaient alors presqu'inconnus (cf. SAHLI 1990a, 1991a). Les observations elles-memes, sur le
terrain, ont ete faites par des temoins n'ayant pas forcement la rigueur et la competence
scientifiques voulues, qui ont assiste au phenomene ou qui en ont “souffert” ; d'ou des
possibility d'imprecisions, de deformations et d'exagerations des faits. (Dans un journal
allemand , en guise de O. sabulosus, un Spirobolide a ete represente en photographie !).

Des deplacements d'O. sabulosus ont ete signales par HALKKA (1958 ) en Finlande et par
FAIRHURST (1968, 1969) en Grande-Bretagne : peut-etre ne s'agit-il dans les deux cas que de M
et non de MM (?). Quoi qu'il en soit, il semble qu'il ne s'agisse pas du phenomene que nous
traitons ici. Des MM de Ommatoiulus moreleti , avec invasions d'habitations, ont ete signalees
dans le cas particulier de populations introduites par 1’homme en Australie (BAKER, 1978a, b, c,
1979, 1984), cas qui ne sera pas analyse ici.

Nous avons personnellement assiste, sur le terrain, a 3 MM : une en Yougoslavie en 1969
(SAHLI, 1984 ) concernant Pachyiulus fuscipes, une au Val de Sibourg pres de Lan<jon-de-
Provence (France) en 1987 et une a Peillon dans le Mentonnais (Alpes-Maritimes, France) en
1988. Nous n'envisagerons que les deux derniers cas qui concernent O. sabulosus.

MATERIEL

Les animaux etudies sont des Ommatoiulus sabulosus aimatopodus (in Demange, 1981), sabulosus punctatus et
apunctulatus selon Verhoeff (1921 a, b). L'ornementation et la coloration du tegument permettent de distinguer deux
"races" geographiques, reparties dans deux zones situees de part et d'autre du Var. En outre, les animaux localises a l'Ouest
du Var (zone : Cannes, Grasse, Antibes. Bouches-du-Rhone) sont nettement plus sveltes que ceux habitant & l’Est du Var
(zone : arriere-pays ni^ois et Mentonnais). D'autres types d'ornementation et de coloration different des pr6c6dents
existent dans 2 autres zones, dune part pres de Toulon (Var), d’autre part dans les Alpes-de-Haute-Provence (Colmars).

EXPOSE SOMMAIRE DES CAS

Le cas du Val de Sibourg

La MM etudiee a eu lieu a la mi-avril 1987 (essentiellement les 16, 17, 18 avril). Nos
investigations ont ete effectuees, in situ, au lotissement du Val de Sibourg, commune de Lan?on,
a quelques kilometres de Salon-de-Provence (Bouches-du-Rhone). Lanijon lui meme est reste
indemne tandis que dans le Val de Sibourg seul un quartier a ete fortement touche. La majeure
partie du lotissement, constitue de maisons individuelles, etait de construction recente (1-3 ans)
tandis qu'une partie etait encore en construction au moment des faits. D'autres invasions ont ete
signalees dans des villages voisins (Ventabren, La Barben) aux memes dates.

Les iules sont descendus des collines environnantes occupees par de la garrigue. Le
“Camp long” a constitue l'arrivee majeure Est. Les populations de O. sabulosus de cette colline
sont pratiquement encercles par un ensemble de canaux (canal du Verdon, canal EDF, canal de

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DEPLACEMENTS EN MASSES ET INVASIONS DE DIPLOPODES

589

Marseille) et par la ville d'Aix-en-Provence. Face au lotissement du Val de Sibourg, qui a ete
envahi, existe une issue terrestre Est et Nord-Est constituee par un pont. Une bonne partie des
animaux en migration ont emprunte ce pont ainsi que la partie de la colline ou le canal est
souterrain. D'autres ne l'ont pas franchi : ils ont ete arretes par la partie aerienne du canal
(constituant une barriere) et sont restes suspendus aux parois extemes et internes du canal. A
plus grande echelle (chaine de la Trevaresse, chaine des Cotes, chaine d'Eguilles, etc.) les iules
sont en fait encercles entre la Durance, le Canal de Marseille, l'Arc et la ville d'Aix-en-Provence.

Des milliers d'iules, provenant de la garrigue environnante (qui recouvrait le lotissement
avant sa construction), se sont deplaces dans les rues, sur les trottoirs, les murs de cloture et les
murs exterieurs des maisons (parfois, pour un petit nombre d’individus, sur les murs interieurs
de certaines habitations).

Le deplacement est survenu apres les fortes pluies de la premiere moitie d’avril.

Quatre generations participent au deplacement : trois generations de 1983, 1984 et 1985.
[les individus nes en automne 1984 (ages d’un an 1/2) etant majoritaires par rapport aux autres] ;
une generation de petites larves a 4 ou 5 rangees d'ocelles (RO) de l'annee 1986 (larves issues
des pontes de l'automne 1986), elles aussi, particulierement nombreuses. Apres l'intervention
des pompiers a l'aide de gaz toxiques, un amoncellement de jeunes larves, mortes ou a demi-
mortes, pouvaient s'observer dans les caniveaux sur une epaisseur de 10 a 15 cm. De nombreux
individus de toute taille accroches aux murs du canal furent epargnes par l'intervention des
pompiers qui craignirent de contaminer l'eau. Au cours de cette “invasion” de printemps 1987, il
convient de noter la participation de jeunes larves nees l'automne precedent (agees de 6 a 7
mois) : des larves aussi jeunes ne figuraient dans les MM precedemment analysees, ni chez
O. sabulosus (au printemps, en Alsace et en Allemagne), ni chez P.fuscipes (en automne en ex-
Yougoslavie).

Le cas de Peillon

L'invasion a commence debut avril 1988, un an environ apres celle de Sibourg. Deux
parties du village de Peillon ont ete envahies. On distingue d'une part, un foyer principal
N.N.W, concernant quelques maisons de construction plus ou moins recente localisees sur la
D121 a 1'entree du village, le cimetiere, la chapelle des Penitents blancs ; d'autre part un foyer
Est situe a l'extremite superieure du village englobant l'eglise et quelques maisons individuelles
(de construction ancienne) proches. Les deux foyers sont disjoints.

Caracteristiques sommaires :

Le deplacement eut lieu apres de fortes pluies. La partie Nord et Nord-Ouest du foyer
principal, boisee (essentiellement d'oliviers), comporte une couverture vegetale protegeant mieux
des pluies torrentielles que la garrigue peu dense des versants alentour. Le foyer de l'eglise,
quant a lui, est borde par un a-pic rocheux. inaccessible a l'Homme. recouvert d'une garrigue
dense d'ou proviennent les animaux.

Aux alentours des foyers, sur les hauteurs de la zone N et NW ainsi que sur les collines
environnantes, il n'y a que fort peu d'individus (les animaux ayant probablement quitte ces
collines pour se concentrer dans le foyer Nord-Ouest).

Trois generations sont presentes sur les sites d'invasions : deux des annees 1984 et 1985,
(avec une predominance des individus nes en 1984) et une constituee de petites larves (4-5RO)
issues des pontes de l'automne 1987.

Les grands individus (larves agees, adultes, post-adultes des generations 1984 et 1985)
migrent, en un premier temps, en avril. Les petites larves de 1987 - alors qu 'elles appartiennent
maintenant aux stades 5 et 6RO - apparaissent en grand nombre dans un second temps, en mai,
sur les murs des maisons, de l'eglise, de la chapelle et sur les tombes en marbre du cimetiere
(done dans les deux zones d'invasion). Contrairement au Val de Sibourg (ou le developpement
est plus rapide), il y a ici decalage dans le temps entre les deplacements sur les murs, d'une part,

590

FRANCOIS SAHLI

des generations 1984 et 1985 et d'autre part de la generation de 1987 (son developpement est en
retard de 1 a 2 mois sur celui de Sibourg).

INTERPRETATIONS ET DISCUSSION

A un endroit donne, on constate une concentration massive d'individus dont la mobilite est
incontestablement en rapport avec de fortes pluies. Tentons de repondre aux principales
questions qui se posent.

Origines de la concentration

a) On peut envisager une origine geographique soit proche (deplacements courts de 1'ordre
de plusieurs metres a plusieurs centaines de metres), soit lointaine (5 a 10 km ou plus?).

En ce qui conceme le Val de Sibourg et ses environs, rien n'empeche de penser qu'il y a
eu deplacement par rayonnement, a partir de la vaste garrigue, par 1 'ensemble (ou une partie) des
points de sortie possibles (ponts, passages aeriens correspondant aux passages souterrains des
canaux). Le Val de Sibourg representerait l'un de ces points d'echappement. Ce rayonnement a
pu s'effectuer, soit a partir du petit encerclement constitue par les canaux, soit a partir du grand
encerciement faisant intervenir la Durance.

En ce qui concerne Peillon, nous pensons que les animaux ayant migre en avril 1988
etaient deja sur place en hiver 1987-88 et/ou au printemps 1988. L'experience acquise montre
bien que, dans les Alpes-Maritimes, les pluies torrentielles automnales et printanieres entrainent
un abandon des sites, notamment a euphorbes, a romarin et a lavande, par les populations de
O. sabulosus. II est probable que les individus concentres au printemps 88, proviennent (au
moms pro parte) de deplacements effectues en automne 1987 lot s des accouplements et surtout
suite aux fortes pluies du 5 et 10 octobre 1987. Le meme raisonnement peut s'appliquer aux
O. sabulosus de la region de Sibourg. La migration aurait ainsi pu s'effectuer en plusieurs
temps.

b) La concentration n'a pas uniquement une origine geographique. On note, aussi bien au
Val de Sibourg qua Peillon, une predominance des individus nes en automne 1984, annee tout a
lait exceptionnelle par le nombre des pontes (abondance des femelles pondeuses et etendue des
sites de ponte) non seulement dans le sud-est de la France, mais aussi, paradoxalement, en
Bourgogne (Cote d'Or, Porron) et en Allemagne (Sarre) (SAHLI, inedit).

Le developpement post-embryonnaire se deroulant avec peu de pertes, comme ce fut le cas
cette annee de pontes abondantes (APA), il est suivi, deux ans plus tard a Sibourg et 3 ans plus
tard a Peillon. d’une annee d'abondance amplifiee des femelles adultes (AAA = annee
d abondance des adultes, femelles ou males) et des pontes (APA) ; ce qui s’est effectivement
passe.

Deplacements et activite

Outre la concentration et la surpopulation, les deplacements sont lies a l’activite de
1 espece, dont on distingue 7 types en Provence et Cote d'Azur : (a) une activite post-exuviale,
(b) une activite declenchee par des fortes pluies ou (c) par un temps lourd orageux (SAHLI,
1986c), (d) une activite sexuelle au moment de l'accouplement, (e) une activite propre aux
femelles fecondees cherchant un lieu de ponte, (f) une activite nocturne (FAIRHURST, 1968), (g)
une activite des femelles (en mai-juin) liee a la nutrition.

Dans la joumee, a certaines periodes de l'annee (par exemple en mai) les iules peuvent ne
pas utiliser leurs abris diumes et demeurer inactifs soit a Pair libre, comme en altitude (Rocca
Spaviera, Petra Cava : prairie avec herbe de 3 a 5 mm de haut, broutee par les moutons,
comportant a proximite des buissons de buis, de Juniperus, des touffes d'euphorbes et de
lavande), soit abrites (Rocca Spaviera, Petra Cava, ruines de Chateauneuf-de-Contes, Peillon,
Baousset, etc ).

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Les deplacements de Sibourg et de Peillon sont en rapport avec les causes d'activite (a) et
(b) : les individus sortaient de leur mue hivernale et il y a eu de fortes pluies coi'ncidant avec la
1’activite post-exuviale. A ces deux facteurs s'ajoute une surpopulation. Les deplacements a
distance eurent eventuellement lieu de nuit (type f) en plusieurs etapes. Des experiences inedites
et les observations faites sur une dizaine d'annees dans les Alpes-Maritimes demontrent
1 influence des precipitations violentes et soudaines avec un risque reel de noyade (cas observes
au Pas de l'Escous), les ouvertures metameriques des trachees etant localisees face ventrale
ainsi qu’une reaction de fuite (SAHLI, 1984).

Le decalage dans le temps entre les invasions de Provence (1987) et celles du Mentonnais (1988)

Ce decalage s’explique par une difference dans le rythme des pontes abondantes. Compte
tenu des circonstances environnementales, les pontes exceptionnellement abondantes de 1984
ont conduit 2 ans plus tard a Sibourg, 3 a Peillon, a une abondance d'adultes. On note des
differences dans le temps des mues de maturations juveniles-adultes (= MMJ), variables selon
les sites et les annees (SAHLI, 1992). Dans d'autres circonstances, les individus nes en 1984 a
Sibourg auraient, tres bien pu pondre massivement 3 ans apres leur naissance, au lieu de 2. De
meme, les individus de 1984 de Peillon auraient pu pondre 2 ans ou 4 ans apres, au lieu de 3. En
outre, au lieu de massives ou explosives, les pontes auraient pu etre plus ou moins equitablement
etalees sur 2, 3 et eventuellement 4 annees (strategic CAT , SAHLI, 1990a, b, 1991a, b).

La double surpopulation

L'analyse des MM de Sibourg et de Peillon se rapporte a trois points.

a) Concentration dans un meme endroit et au printemps des adultes, post-adultes,
intercalaires et des larves agees, appartenant a 2 ou 3 generations. Celle de 1984 est la plus
nombreuse, elle correspond aux descendants d'une annee de pontes abondantes. La
surpopulation des femelles est en fait due a celles ayant pondu l'automne precedent, celle des
males est due aux intercalaires (cas de Sibourg). La surpopulation est done essentiellement le fait
d'animaux post-imaginaux. Elle peut se schematiser ainsi, IN representant les adultes, post-
adultes, intercalaires et larves agees : IN 1983 + IN 1984 (APA) + IN 1985 = surpopulation d'IN
(SIN) au printemps dans un meme site (1987 a Sibourg, 1988 a Peillon ou les IN 1983 font
defaut).

b) Concentration de pontes puis de jeunes larves de 2 (voire 3) generations dans un meme
endroit et la meme annee. La majorite de ces pontes est le fait de males adl ou/et ad2
surabondants, provenant eux-memes d'une APA (generation de 1984). Ces pontes automnales
de 1’ annee x conduisent au printemps suivant (x+1) a une surpopulation de jeunes larves (SJL).

Dans le cas de Sibourg, (P = ponte d'une annee) les choses se passerent probablement
ainsi : “P 1 986 (des IN 1983) + PI 986 (des IN1984, APAde 1984) + ?P 1 986 (des IN 1 985) = APA 1986”.

Tout porte a croire qu il y a eu predominance des PI 986 (des IN 1984). II y a eu repetition
dans le temps de deux APA, l’une en 1984, l’autre en 1986 (Sibourg). Dans les 2 MM
observees, les pontes ont ete explosives deux fois de suite : en 1984 (APA a Sibourg et Peillon)
et en 1986 (APA a Sibourg) ou en 1987 (APA a Peillon).

L'experience montre que d’autres situations sont possibles lorsqu'il n'y a pas de MM.
D'une part, les sites de ponte de 2 generations peuvent etre differents dans l'espace (changement
de sites de ponte) ; d'autre part, les pontes d'une meme generation peuvent etre reparties
equitablement ou non dans le temps sur plusieurs annees : strategic de repartition “CAT”
(SAHLI, 1991 b) , l’oppose d'une strategic explosive.

c) Reunion dans un meme endroit et la meme annee d'une surabondance d’adultes, post-
adultes, intercalaires et larves agees (SIN), encore en vie dans le site, avec une surabondance de
jeunes larves (SJL). Avant la MM dans les aires d'origine on a une double surpopulation
SIN+SJL (a Sibourg, elle s'observe simultanement dans les aires post-migratoires tandis qu'a

592

FRANCOIS SAHLI

Peillon il y a un leger decalage dans le temps). Apres avoir pondu, les femelles adultes quittent
souvent le lieu de ponte (Col de Braus), alors que dans le cas des MM de Sibourg et de Peillon,
males et femelles post-imaginaux se trouvent dans le meme endroit que les pontes et les larves
qui en sont issues. A Sibourg, tant qu'il n'y a que des oeufs (automne 86), puis des larves a 2-
3(4)RO (automne 86, hiver 86-87), la densite dans les aires d'origine correspond simplement a
une seule surpopulation. Lorsqu'au printemps 87, apres l’hivernation, les larves parvenues aux
stades 4-5RO se mettent a se deplacer et a manger abondamment, il y a une double
surpopulation, traduite par une densite accrue, dont les effets se font sentir sur l'ensemble de la
population. Notons que les points b et c ne s'appliquent pas a la migration en masse automnale
de P.fuscipes en Yougoslavie.

A Sibourg, selon le point a, la ponte abondante de 1986 s’explique par la presence de
nombreux adultes cette annee-la (AAA 1986), provenant pour la plupart de la generation
exceptionnellement abondante de 1984. Mais, pour qu'il y ait beaucoup de femelles post-adultes
et d'intercalaires males au printemps 1987, il faut necessairement un maintien en vie qui conduit
les males et les femelles de la generation 1984, devenus adl en 1986, a des indi vidus post-
imaginaux en 1987.

Le maintien en vie, I'annee suivante, des femelles et des males apres I'accouplement et la ponte

a) Chez les femelles, il s’explique soit par l'existence dune iteroparite directe ou indirecte
(SAHLI, 1993 ) chez les femelles, les femelles adl ayant pondu en 1986 resteraient capables de
se reproduire une nouvelle fois en 1987 (iteroparite directe), en 1988 ou en 1989 (iteroparite
indirecte) ; soit, plus vraisemblabement, par la possibility d'un maintien en vie des femelles adl
de 6 mois a 10-12 mois apres la ponte, comme cela s'est produit dans les elevages. Dans cette
hypothese, la survie passagere serait suivie de mort, sans qu'il y ait eu iteroparite, pouvant
representer un vestige d'une strategic ancestrale (caractere genetique ancestral d'iteroparite
femelle). Cet hypothetique caractere ne s'exprimerait plus (ou peu ou rarement?) dans les
conditions de temperature et de pluviosite qui regnent, a faible altitude, dans le sud-est de la
France. L'iteroparite a d'autant moins de raisons de s'exprimer dans ces sites que le nombre
d'oeufs produits (et pondus en une fois) par femelle est important (de l'ordre de 500). Le
caractere ancestral pourrait aussi ne pas pouvoir s'exprimer dans le cas d'une forte surpopulation
comme c'est le cas ici. L'iteroparite chez Ommatoiulus pourrait traduire une adaptation au froid
de l'epoque glaciaire (SAHLI, 1990a, b : glacier du Mercantour il y a 20000 ans). Les choses
purent se passer ainsi : production et ponte d'un nombre relativement petit d'oeufs (tres inferieur
a l'actuel) avec repetition des pontes a des intervalles de plusieurs annees et des periodes de
repos entre 2 pontes comme cela a ete admis chez les femelles de Typhloblaniulus (SAHLI,
1993). L’iteroparite ancestrale presumee des femelle de O. sabulosus serait sous la dependance
d'un ou de plusieurs genes non lies au sexe et aurait pu se manifester autrefois aussi bien chez
les femelles que chez les males. Notons que le comportement de O. sabulosus (notamment des
femelles), apres la migration, donne l'impression d'un “suicide” collectif : inactivity, exposition
aux conditions environnementales, deshydratation et mort alors qu'ils pourraient se proteger et
se nourrir dans la garrigue toute proche.

b) Chez les males, le mecanisme de survie est la periodomorphose, phenomene a
determination genetique probable (SAHLI, passim ), propre a certains diplopodes et conduisant a
des males post-imaginaux. Les especes a males periodomorphiques ou a males post-imaginaux
iteropares sans regression (succession adulte-adulte = strategic y) sont des especes anciennes,
ayant probablement deja vecu a l'epoque glaciaire. Les especes a males periodomorphiques
presentent un grand avantage : elles sont, grace a la periodomorphose et aux intercalaires de
longue duree, [formes beaucoup plus resistantes a un environnement defavorable que les males
adultes (SAHLI, 1991c )] et peut-etre grace a l'iteroparite indirecte des femelles, adaptees aux
changements de climat, sans etre obligees d'avoir recours a une speciation, aussi bien dans un

DEPLACEMENTS EN MASSES ET INVASIONS DE DIPLOPODES

593

sens (froid, periode glaciaire) que dans 1 'autre (periodes interglaciaires). Cette adaptation permet
aussi une grande extension geographique actuelle Nord - Sud, des plaines basses aux sommets
montagneux de 3000 m. d'altitude, avec possibility d'occuper divers ecosystemes (forets,
prairies, garrigues, etc.). D'apres nos recherches experimentales, les intercalaires de
O. sabulosus paraissent mieux adaptes (plus resistants) au froid qu'a la chaleur (SAHLI, 1991c).

A Sibourg, le schema du developpement male retenu est le suivant : adl (automne 1986,
nes en 1984) - intercalaires (printemps 1987, males dedifferencies retrouvant une morphologie
quasi-juvenile) - ad2 (ete-automne 1987 apres une mue de maturation). Les males intercalaires
sont extremement nombreux au printemps 1987, le rapport intercalaires / tous les autres males
(juveniles de 7 a lORO + adl) est de 3/1 (SAHLI, 1990), le rapport intercalaires/femelles
(individus de 9 a 13RO) est de 1/1,55. Ceci confirme clairement trois regies - (rl) : dans de
bonnes conditions, presque tous les males adl peuvent se transformer en intercalaires (SAHLI,
1986 b) ; (r2) : les intercalaires si sont nombreux une annee x lorsque les adultes adl dont ils
sont issus etaient eux-memes nombreux l'annee x-1 (SAHLI 1986a). Le cas de Sibourg demontre
bien qu'apres de bonnes mues de maturations des juveniles en adl (MMJ) et lorsque tous (ou
presque tous) les individus sont rassembles en un point suite a une MM (au lieu d'etre disperses
dune fa$on irreguliere) la frequence des intercalaires si peut etre tres elevee ; (r3) : a faible
altitude, en Provence et dans les Alpes Maritimes, la grande majorite des males ad2 meurent a
1' automne de l'annee x (SAHLI, inedit), d’ou la rarete des s2.

A Peillon, en automne 1986 (secheresse), les males adl (nes en APA-1984) sont
nombreux, bien qu'il demeure une fraction de males juveniles tardifs non transformes en adl en
1986. Les femelles sont vraisemblablement restees majoritairement juveniles en 1986.

L'affirmation selon laquelle il n'y eut probablement que ires peu de pontes en 1986 a Peillon repose sur les raisons
suivantes : des larves 4-5RO au printemps 1987. des larves 7-8-9RO au printemps 1988 et des individus 9-lORO au
printemps 1989 etaient peu nombreux, voire rares ; or, ils auraient dte nombreux (au moins par moments) s'il y avait eu
des pontes abondantes en automne 1986 et ils n'auraient pu echapper a I'observation pendant plusieurs annees, sauf en
cas d'emigration massive sans retour ; de plus, ce que nous venons de dire de Peillon au sujet de la rarete des pontes en
1986 est vaiable pour d'autres stations du Mentonnais.

Au printemps 1987 et a Peillon, les intercalaires si devaient, en principe, etre nombreux a
certains endroits. En automne 1987 (pluies torrentielles depeuplant les sites d'observation par
emigration) les ad2 (issus des si) devaient, en principe, etre nombreux eux aussi a certains
endroits. Or, dans les endroits prospectes et la ou des animaux ont pu etre trouves, les si du
printemps 1987 et les ad2 de l'automne 1987 etaient peu nombreux. Nous ignorions qu'une MM
allait se produire en 1988 a l'entree du village et au cimetiere ; 1 'entree du village n'a done pas ete
prospectee au printemps et a l'automne 1987. Nous n'avons pas non plus pu trouver d'animaux
en automne 1987 aux environs de l'eglise (balayages trop frequents). La plupart des femelles
adl nees en 1984 sont devenues matures en 1987 (comme dans d'autres sites du Mentonnais).
Ce sont en majeure partie les meres des nombreuses larves nees en 1987. A l'automne 1987, la
situation s’interprete ainsi : (crl) croisements majoritaires ad2 males nes en 1984 x adl femelles
nees majoritairement en 1984, done decalage d'un an entre les premieres maturations males et
femelles ; (cr2) croisements minoritaires adl males tardifs nes en 1984 x adl femelles de 1984 ;
(cr3) croisements minoritaires de quelques adl males nes en 1985 x adl femelles de 1984 et
eventuellement de 1985. Des ad2 males d'automne 1987 de Peillon, seul un tout petit nombre a
survecu (r3) et s'est transforme, au printemps 1988, en intercalaires s2 qui, vu leur tres faible
nombre, n'ont peut-etre pas figure dans les prelevements des animaux en migration au printemps
1988. Le resultat est que lors de la migration de Peillon, contrairement a celle de Sibourg, les
intercalaires etaient relativement peu nombreux (le rapport intercalaires/femelles varie entre 1/1,7
et 1/2 selon les endroits).

Concemant les O. sabulosus mediterraneens de faible altitude, il en resulte une quatrieme
regie (r4) corollaire de la troisieme : les males adultes peuvent etre nombreux un automne x et les
intercalaires peu nombreux au printemps x+1 lorsque les adultes en question etaient des ad2.
Une ponte abondante a l'automne x, suivie d'abondantes larves au printemps x+1. peut masquer

594

FRANCOIS SAHLI

les geniteurs males adultes reels de 1'automne x lorsque ceux-ci sont des ad2. Dans ce cas, les
intercalaires peu nombreux recoltes au printemps x+1 sont pour la plupart des si et ne derivent
par consequent pratiquement pas des geniteurs ad2, morts en majorite. A Sibourg, au printemps

1987, il s'agissait, grosso modo, d'intercalaires 1 ages de 2 ans et demi. A Peillon, au printemps

1988, la plupart des ad2 etant morts, il n'y avait que quelques intercalaires si de la generation
1985 (ages de 2 ans et demi), plus quelques rares intercalaires pouvant etre des si tardifs, ages
de 3 ans et demi et correspondant peut-etre aux juveniles tardifs de 1986.

Le faible nombre des intercalaires si

Le nombre des intercalaires peut etre egalement plus ou moins faible dans le cas ou il n’y a
pas explosion d'adultes, c'est-a-dire lorsque la production des adultes n’est plus explosive et
concentree sur une annpe comme en 1986 a Sibourg mais repartie plus ou moins equitablement
sur deux a quatre ans. Dans ce cas, le nombre des larves d'une generation susceptibles de
devenir adultes est fractionne : certaines restent juveniles, d'autres deviennent adultes adl. Nous
pouvons admettre les rapports 50/50 ou 40/60 sur deux ans, 10/55/35 ou 10/40/40 sur 3 ans,
etc. La regie (r4), corollaire de la regie (rl) intervient alors : lorsque les adultes adl sont produits
en petite quantite, parce que leur production est plus ou moins largement fractionnee une annee
x, les intercalaires 1 de l'annee x+1 sont peu nombreux et leur nombre depend de l'importance
de la fraction d'adl de l'annee x.

Une faible quantite d'intercalaires une annee donnee dans un site donne peut done avoir au
moins deux raisons : (rail) l'existence de geniteurs ad2 ou (rai2) l'adoption d'une strategie de
fractionnement CAT. Il y a une troisieme raison (rai3) : l'existence d'une distribution non
homogene d'intercalaires (SAHLI, 1991a) qui fait que le nombre d'intercalaires recoltes a un
endroit donne ne traduit pas leur effectif reel dans la population. 11 existe enfin une quatrieme
raison (rai 4) : la delocalisation d'un site (par exemple du site de naissance) par emigration de la
quasi-totalite de la population dans un autre, suite a des pluies torrentielles de printemps ou/et
d'automne et sans qu'il y ait pour autant invasion d'habitations, ces dernieres faisant defaut.

Compte tenu des raisons 3 et 4, une MM presente l'avantage de rassembler tous les
individus (ou une bonne partie d'entre eux), intercalaires compris, au point de migration. Il en
resulte une cinquieme regie (r5) : il est necessaire de bien connaitre un site donne et les
antecedents biologiques des animaux qui y vivent (pontes, jeunes larves, apparition des stades
plus ages au cours des printemps et automnes successifs, maturation sexuelle des males (adl),
apparition d'intercalaires). Meme cela n'est pas suffisant puisque nous venons de voir qu'il peut
y avoir des evacuations massives dues aux antecedents environnementaux. Le nombre faible
d’intercalaires observe une annee x dans un site, lorsque les antecedents biologiques et
environnementaux du site sont mal connus, ne presente pas alors une grande signification.

Cette remarque, ainsi que celle ci-dessous, s'applique a nos travaux anterieurs a 1985 et
aux interpretations figurant implicitement ou explicitement dans ceux de nos predecesseurs :
VERHOEFF, BROLEMANN, HaLKKA, BLOWER, FAIRHURST (cf. 1974), BAKER, ENGHOFF...

Le nombre de RO est parfois difficile a etablir, notamment chez les differentes sortes de
males adultes et d'intercalaires ages (dimorphisme sexuel?). La determination de l’age se
complique du fait de la possibility de chevauchements dans un meme site (melange de stades
precoces et tardifs ayant le meme nombre de RO) ou dans plusieurs sites differents (rythmes
differents d'un site a l'autre) (SAHLI, 1992). De plus, en mai, a l'age d'l an et demi, une partie
des femelles est en avance d'une rangee ocellaire (et done d'un stade) sur les males de la meme
generation : dimorphisme sexuel couramment observe.

Source :

DEPLACEMENTS EN MASSES ET INVASIONS DEDIPLOPODES

595

La question de deux APA consecutives (preuve indirecte d'une iteroparite femelle importante)

Dans le sud-est de la France, nous n'avons jamais observe 2 vraies APA deux annees
consecutives. En revanche, il peut y avoir 2 AAA (pour les males : d'abord des adl puis, l'annee
suivante, des intercalaires suivis d'ad2).

Deux vraies APA , deux annees consecutives, seraient la preuve tangible d'une iteroparite
directe femelle, importante, dans laquelle presque toutes (=80 %) les femelles adl d'une annee
seraient capables de pondre une nouvelle fois l'annee suivante. Pareille preuve indirecte n'a
jamais pu etre apportee.

Admettons qu'il y ait fractionnement (strategic CAT) et que le rapport des adl males
produits 2 annees consecutives soit de 50/50 (ou 10/40/40 sur 3 annees consecutives). II peut
alors y avoir des adultes adl en plus ou moins grand nombre deux annees consecutives (50/50
ou 40/40). Si les femelles adl et les pontes sont fractionnees dans le temps dans les memes
proportions que celles admises ci-dessus pour les adl males, on est conduit a deux pseudo APA
(50/50 ou 40/40) et non a 2 vraies APA (80 et 80). Dans le cas de deux vraies APA
consecutives, il y aurait des pontes explosives deux annees de suite, done deux fois plus de
pontes par annee. Si la distinction entre pseudo et vraies APA est facile en theorie et en pratique
dans les cas extremes, elle peut cependant etre fort delicate lorsque les cas sont moms tranches.

Le role des intercalaires

Dans le cas de Sibourg, les nombreux intercalaires du printemps 1987, des SL pour la
plupart (composante « de la periodomorphose, SAHLI, 1990), ne furent tres probablement
d’aucune utilite et moururent sans avoir pu jouer de role : bien que devenus des ad2 en ete 1987,
les femelles (partenaires en puissance de ces ad2 males) ne purent probablement pas etre
fecondees une nouvelle fois en 1987.

A faible altitude dans le sud-est de la France et de nos jours, les roles des intercalaires chez
O. sabulosus peuvent etre les suivants :

(a) sous la forme de SL, ils foumissent des partenaires males (ad2) aux femelles adl dans
le cas d'une non-synchronisation massive des maturations males et femelles (decalage dans le
temps, cf. supra le cas de la MM de Peillon),

(b) sous forme de SL (ou de LL), ils fournissent des partenaires ad2 aux femelles adl dans
le cas ou les pontes sont fractionnees dans le temps (strategic CAT) : certaines femelles pondent
l'annee x, d'autres, de meme generation l'annee x+1, d'autres encore, de meme generation
l'annee x+2, etc.

Sur une vingtaine d'annees, il est possible que les femelles fassent usage, selon les
generations, les sites et l’environnement, tantot de la possibility (a), tantot de la (b), tantot de la
(c), voire peut-etre de combinaisons : (a) + un peu de (c) ou (b) + un peu de (c).

La presence d'abris

Lorsque surviennent de fortes pluies, les iules quittent les zones a vegetation peu dense et
basse (touffes d 'Euphorbia dans le Mentonnais) pour se refugier generalement dans des sites-
abris (sites-relais), ou la vegetation est abondante, dense et haute, composee d'arbres,
d'arbrisseaux ou de buissons protecteurs et non plus de touffes.

Halkka (1958) avait dej& signale de pareils deplacements chez O. sabulosus en Finlande : deplacements entre un
site bien abrite a vegetation dense et entre un site expose a vegetation clairsemee, lors de I'hivernation ou lors de pontes.
De pareils deplacements ont aussi ete notes par Fairhurst (1968) en Grande-Bretagne.

Lors des migrations en masse dans les villages de Sibourg et de Peillon, les specimens de
O. sabulosus sont, semble-il, partis (au printemps de la MM ) de sites-abris ou relais (garrigue
dense de Sibourg constitute notamment de chenes, zone boisee et garrigue dense de Peillon). Ce

596

FRANCOIS SAHLI

depart peut s'expliquer par une protection devenue insuffisante(?) ou, apres les pluies, par un
hygrotropisme negatif (reaction a l'egard d'une humidite trop elevee dans les sites-abris) et/ou en
grande partie sous la pression demographique (enorme densite d'individus sous chaque abri).

Ils grimpent alors sur les surfaces verticales qu'ils rencontrent sur leur trajet : murs
d'habitations humaines, d'eglises ou de chapelles, murs de canaux, rochers verticaux, parois de
tunnels autoroutiers (Sibourg), arbres ( cf Pachyiulus, SAHLI, 1984), oliviers notamment. Ils
peuvent aussi sejoumer sur des aires horizontales surelevees (tombes des cimetieres de Peillon et
de Gorbio).

La faible influence des predateurs, des competiteurs el des parasites

II peut y avoir des predateurs (coleoperes, araignees, scorpions, sangliers?, blaireaux,
renards?, scolopendres? presentes a Sibourg), des cohabitants qui ne sont pas reellement des
competiteurs (comme dans le Mentonnais Cylindroiulus limitaneus, Trichoblaniulus hirsutus,
Callipus foetidissimus ou Glomeris sp. dans certains sites : leur role est vraisemblablement
negligeable). Des parasites tels que les larves de certains dipteres peuvent jouer un role
important. Toujours est-il que : (a) le nombre des individus devenus adultes en 1984 a ete
considerable ; (b) les individus nes en automne 1984 de pontes extraordinairement importantes
sont, pour une bonne part, devenus adultes en 1986 ou 1987, l'influence des predateurs,
concurrents, et parasites ayant ete faible ou nulle.

On peut supposer que le nombre d'individus parvenus a 1'etat adulte (adl) a partir de
chaque ponte a ete exceptionnellement superieur en 1984 puis en 1986 (Sibourg) et en 1987
(Peillon) au nombre habituel (a partir de plusieurs centaines d'oeufs d'une ponte, une fraction
importante de larves meurt habituellement au cours du developpement).

La proximite d’une vaste garrigue

Dans le sud-est de la France il est logique de penser qu'une MM ne peut se produire que
lorsqu'une vaste garrigue permet l'hebergement, puis le deplacement, d'un nombre considerable
d'individus. De Salon-de-Provence a Aix-en-Provence, la garrigue s'etend sur des kilometres :
les sites a O. sabulosus sont etendus. On ne saurait imaginer un important deplacement a partir
d'un llot de garrigue ou d'une zone ne comportant que de petits fragments de garrigue, comme
c'est le cas dans nombre des stations du Mentonnais : ici, a part Peillon, peu de sites peuvent
constituer un terrain favorable a une MM avec invasion. A Peillon, la MM a ete de moindre
envergure qu'a Sibourg, sans doute parce que l'etendue du site a O. sabulosus y est moindre.

Une MM avec invasion d'habitations ne se produit que lorsque un village est implante dans
la garrigue, a proximite d'une garrigue assez vaste, ou sur l'itineraire d'un deplacement. Dans
l'etat actuel de nos connaissances, on ne saurait imaginer une MM dans de grandes
agglomerations comme celles de Nice, Monaco ou Menton. Dans le Mentonnais, les endroits les
plus exposes a une future MM avec invasion, outre Peillon, nous paraissent etre le hameau
d'Engarvin et les villages de Gorbio et de Sainte-Agnes. Dans ces deux derniers, une mini¬
invasion de murs d'habitations par O. sabulosus eut lieu non au printemps mais,
paradoxalement, en automne 1988 apres des pluies.

Pourquoi une MM n'a-t-elle pas eu lieu en 1985 ?

1984 ayant ete une annee exceptionnelle tant par l'abondance d'individus matures (AAA)
que par celle de pontes (APA, automne 1984), comment se fait-il qu'il n'y ait pas eu de MM, ni
en Provence (Sibourg), ni dans les Alpes-Maritimes (Peillon) l'annee suivante, au printemps
1985? Pourtant les conditions d’une double surpopulation SIN+SJL etaient remplies au
printemps 1985 comme au printemps 1987 a Sibourg.

Trois differences meritent d'etre signalees:

Source :

REPLACEMENTS EN MASSES ET INVASIONS DE D1PLOPODES

597

a) hiver 1984-1985 exceptionnellement froid, que les jeunes larves et les adultes, bien
adaptes, ont remarquablement supporte, meme en altitude (observations personnels);

b) secheresse inhabituelle du printemps 1985 et du printemps 1986 ;

c) surpopulation SIN (grands individus) du printemps 1985 sans doute moindre que celles
des printemps 1986 et 1987 a Sibourg : la composante IN de 1986 (puis de 1987) a
probablement ete amplifiee par rapport a celle de 1984 ; en effet chaque femelle adl de 1984 a
produit plusieurs centaines d'oeufs devenus, pro parte, a leur tour, des adl en 1986 (Sibourg),
augmentant ainsi le nombre des IN de 1986 (devenus des post-adultes en 1987) par rapport a
ceux de 1984 (cf supra).

En 1985, la secheresse a pu maintenir les populations de O. sabulosus dans leur habitat
(notamment les jeunes larves) empechant toute emigration. II n'y a eu en effet nude part de MM
avec invasion. Nous emettons l'hypothese que l'absence de pluie est responsable d'une absence
de MM meme en cas de surabondance des individus post-imaginaux et des jeunes larves. Peut-
etre aurait-il suffi de quelques pluies legeres au printemps 1985 pour declancher des MM a
certains endroits. Aussi admettrons nous que les pluies sont une condition determinante de MM.

CONCLUSION

La ou il y a eu des invasions, les habitations humaines sont typiquement construites dans
des sites a iules, souvent tres anciens : ainsi en Sarre, en 1973, en Yougoslavie en 1984, dans
les Alpes-de-Haute-Provence en 1989 (Cylindroiulus broti , d'apres SAHLi, inedit), dans divers
sites des Bouches-du-Rhone et a Peymenade dans les Alpes-Maritimes en 1963 (SAHLI, inedit).
C'est un peu comme si on avait bati sur une termitiere ou sur une fourmilliere, ou a proximite.
Vouloir gouter les joies de vivre en pleine nature, mais n’en supporter aucune consequense est
difficilement conciliable. Le corollaire de notre affirmation est qu'il existe des deplacements
massifs de O. sabulosus sans invasions, passant totalement inaper£us lorsque les habitations
font defaut. Nous en avons observe dans le Mentonnais a Rocca Spaviera et au Pas de l'Escous.
Le cas decrit par VERHOEFF en Alsace trouve sa place dans cette rubrique dans la mesure ou les
O. sabulosus ont simplement traverse une voie ferree.

Le fait de grimper sur les murs les parois d'un canal traduit, dans un premier temps, une
reaction de fuite ou un hygrotropisme negatif. Dans le cas d'une surpopulation, concernant les
femelles adl ayant pondu l'automne precedent, ce comportement conduit, dans un deuxieme
temps, a une mortalite considerable. Fixes sur des murs, les iules semblent ne pas se nourrir et
s’exposent a la deshydratation. Or, dans le Sud-Est, en avril et mai, l’alimentation est necessaire
a la production des ceufs chez les femelles, a la mue intercalaire 1- ad2 des males en vue de
l’accouplement.

Des essais d’ invasion experimental ont ete conduits a l'interieur et a l'exterieur (balcons)
de plusieurs immeubles du Mentonnais (O. sabulosus) ainsi que sur un balcon dans un
appartement a Dijon ( O . sabulosus et Tachypodoiulus niger) : les animaux, inoffensifs, sont
morts au bout de quelques jours.

Les invasions massives suivies de deplacements et d’une forte mortalite traduisent
vraisemblablement une regulation des populations. En effet, si tous les animaux adultes et
intercalaires qui migrent massivement au printemps se reproduisaient dans le meme site en
automne (ad2 males et femelles), les ressources alimentaires deviendraient vite insuffisantes. La
surpopulation doit logiquement conduire a coloniser des sites nouveaux ou moins peuples, a
condition que la chose soit possible compte tenu, par exemple, de l’anthropisation croissante des
milieux. A notre connaissance, il n'existe pas encore, en Europe, de sites envahis massivement
deux annees consecutives.

Il convient de dire que les invasions massives de iules constituent un risque periodique
dans tout le sud-est de la France, au mois d'avril et en ete, voire en debut d'automne. pour
certains equipements (autoroutes A8 et A9,...) et habitations.

598

FRANCOIS SAHLI

REFERENCES

Baker, G. H.. 1978a. — The distribution and dispersal of the introduced millipede Ommatoiulus moreletii (Diplopoda:
Julidae) in Australia. J. Zool. Lond ., 185 : 1-11.

Baker. G. H.. 1978b. — The post-embryonic development and life history of the millipede Ommatoiulus moreletii
(Diplopoda : Julidae) introduced in south-eastern Australia. J. Zool Lond., 186 : 209-228.

Baker. G. H.. 1978c. — The population dynamics of the millipede Ommatoiulus moreletii (Diplopoda: Julidae). J. Zool.
Loud., 186 : 229-242.

Baker, G. H.. 1979. — The activity patterns of Ommatoiulus moreletii (Diplopoda: Julidae) in South Australia. J. Zool.
Lond., 188 : 173-183.

Baker, G. H., 1984. — Distribution, morphology and life history of the millipede Ommatoiulus moreletii (Diplopoda:
Julidae) in Portugal and comparisons with Australian populations. Aust. J. Zool., 32 : 81 1-822.

Demange, J. M., 1960. — Sur un important rassemblement de Schizophyllum sabulosum (L.). Cah. Natural. Bull, nat.,
Paris, 16 : 89-91.

Demange, J. M.. 1963. — Myriapodes. In : Encyclopedic de la P16iade, Zoologie II. Les Arthropodes. Paris, Gallimard :
411-486.

Demange, J. M., 1981. — Les Millepattes. Myriapodes. Paris, Boubee, 284 pp.

Halkka, R.. 1958. — Life-history of Schizophyllum sabulosus (L.) (Diplopoda, lulidae). Ann. Zool. Soc. Zool. Bot.
Fenn. “ Vanarno ”, 19 : 1-72.

Helb. H. W., 1975. — Zum Massenauftreten des Schnurfussers Schizophyllum sabulosum im Saarland (Myriapoda:
Diplopoda). Entomologica, Stuttgart, Fischer Verlag.

Fairhurst, C. P., 1968. — Life cycles and activity patterns of schizophylline millipedes. Ph. D. Thesis, University of
Manchester, 407 pp.

Fairhurst, C. P., 1969. — Activity and wandering in Tachypodoiulus niger (Leach) and Schizophyllum sabulosum (L.).
Bull. Mus. nat. Hist. nat. Paris, 41, suppl. 2 : 61-66.

Fairhurst, C. P.. 1974. — The adaptive significance of variations in the life cycles of schizophylline millipedes.
Symp. zool. Soc. Lond , 32 : 575-585.

Sahli, F., 1984. — Quelques exemples de deplacements en masse ct de rassemblements chez les Myriapodes Diplopodes.
I09eme Congr. nat. Soc. savantes, Dijon, sc. II : 253-261.

Sahli, F., 1986a. — On some roles of periodomorphosis in Ommatoiulus sabulosus (L.) (Myriapoda, Diplopoda) in the
Maritime Alps. In : Porchet, Advances in Invertebrate Reproduction , 4 . Elsevier Sc. Publ. : 409-416.

Sahli, F., 1986b. — Frequence de la periodomorphose chez le Diplopode Julide Ommatoiulus sabulosus (L.) dans les
Alpes-Maritimes. I. Passages male adulte-male intercalate dans des elevages effectues in natura. C. R. Acad. Sc.
Paris, ser. Ill, 302 : 679-682.

Sahli, F., 1986c. — Influence des orages sur certains deplacements d 'Ommatoiulus sabulosus (L.) (Myriapoda,
Diplopoda, Julida) dans les Alpes-Maritimes. Bull sci. Bourgogne, 38 : 55-57.

Sahli, F., 1990a. — Recherches sur le cycle des males et la frequence des intercalates d' Ommatoiulus sabulosus
(Myriapoda, Diplopoda, Julidae) en Bourgogne. Inversion experimentale du nombre des intercalaires et des adultes en
aulomne. Bull. sci. Bourgogne, 45 : 5 1 -59.

Sahli, F., 1990b. — On post-adult moults in Julida (Myriapoda. Diplopoda). Why do periodomorphosis and
intercalaries occur in males? In : A. Minelli, Proc. 7th Intern. Congr. Myriapodology. Leiden, Brill : 135-156.

Sahli, F., 1991a. — Recherches sur le cycle des males d' Ommatoiulus sabulosus (L.) (Myriapoda, Diplopoda, Julidae)
dans les Alpes-Maritimes et en Provence. Remarques sur les rapports entre la periodomorphose et raltitude. Bull. sci.
Bourg.. 44 : 33-39.

Sahli, F., 1991b. — Frequence de la periodomorphose chez le Diplopode Julide Ommatoiulus sabulosus (L.) dans les
Alpes-Maritimes et en Provence : II. Passages males intercalaires - males adultes. Bull. sci. Bourg., 44 : 41-47.
Sahli, F.. 1991c. — Augmentation et production expcrimentales de males intercalaires chez des Diplopodes Julida. Note
preliminaire. C. R. Acad. Sci. Paris, Ser. HI, 313 : 59-63.

Sahli, F.. 1992. — On male reproduction strategies in Ommatoiulus sabulosus (L.) in the Maritime Alps and Provence
(France): juvenile to adult maturation moults. Ber. nat.-med. Verein, Innsbruck, suppl. 10 : 167-176.

Sahli, F., 1993. — Signification de la periodomorphose et des intercalaires males et femelles chez les Diplopodes
Blaniulides cavernicoles pyr6n6ens. Mem. Biospeol., 20 : 209-215.

Verhoeff, K. W., 1900. — Wandernde Doppelfussler Eisenbahnzuge hemmend. Zool. Anz., 23 : 465-473.

Vf.rhoeff, K. W., 1921a. — Chilognathen Studien. Arch. Naturgesch., 86A : 23-80.

Verhoeff, K. W., 1921b. — Ueber Diplopoden der Riviera und einige alpenlandische Chilognathen. Arch. Naturgesch.,

Source : MNHN, Paris

Distribution Patterns and Qualitative Composition of
the Centipede Fauna in Forestal Habitats of Mainland

Greece

Marzio ZAPPA RO LI

Dipartimento di Protezione delle Piante, Universita della Tuscia, 01 100 Viterbo, Italy

ABSTRACT

A preliminary overview of the centipede communities inhabiting the main forestal ecosystems of mainland Greece is
presented. The study is based on data collected during the last fifteen years. The literature records are thus included, after a
critical evaluation from both faunistic and taxonomic point of view. 54 species are considered. The following forms of
lorcst vegetation are considered: sclerophyllous forests ( Pistacia spp., Quercus coccifera) and their stages of
degradation, deciduous oakwoods ( Quercus spp.), firwoods (Abies cephalonica) and beech-fir forests (Fagus spp., Abies
sp. gr. alba). The centipede community of each formation is discussed from a faunistical, ecological and
zoogeographical point of view.

RESUME

Modalites de la repartition et composition specifique de la faune des chilopodes des milieux
forestiers de la Grece continentale.

Ce travail prcsente un premier bilan des peuplements de chilopodes occupant les principaux ecosystemes forestiers de
la Grfcce continentale. L’etude est basee sur les donnees rdunies durant ces quinze demieres annees. Les donnees de la
literature accessibles sont revues de manure critique a la fois du point de vue faunistique et du point de vue taxinomique et
incluses dans ce bilan. On a pris en consideration les formations forestieres suivantes : forets sclerophylles (Pistacia
spp., Quercus coccifera) et leurs slades de degradation, forets decidues de chenes (Quercus spp.), sapinieres (Abies
cephalonica) et hetraies-sapinieres ( Fagus spp., Abies sp. gr. alba). Le peuplement de chilopodes de chaque formation est
examine du point de vue faunistique, ecologique et zoogeographique.

INTRODUCTION

The main works published to date on the ecology of centipede communities in the
Mediterranean region deals only with some South-European areas, such as Spain (SERRA, 1978;
SERRA & ASCASO, 1990) and Italy (MlNELLI & lOVANE, 1987; ZAPPAROLI, 1992). No data
are available for North-African and East-Mediterranean countries. The aim of this paper is to
give a first qualitative picture of the centipede communities in the main forest habitats of
mainland Greece, in order to characterize these habitats and to allow for synecological
comparisons over the Mediterranean area.

Zapparoli, M.. 1996. — Distribution patterns and qualitative composition of the centipede fauna in forestal
habitats of mainland Greece. In: Geoffroy, J.-J., Mauri£s. J.-P. & Nguyen Duy - Jacquemin. M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist. nat.. 169 : 599-605. Paris ISBN : 2-85653-502-X.

600

MARZIO ZAPPAROLI

MATERIAL AND METHODS

In the present study the territory of Greece is considered as it is politically constituted today; the Egean Islands
and Crete are not analysed. The analysis is mainly based on data collected in the last fifteen years (Zapparoli, 1994), but
all available literature records, critically reviewed both from faunistic and taxonomic point of view, are also included.
About 280 sites are considered, 87 of which are known in their main vegetation features. 54 species of centipedes are
considered: i-e about the 85% of the species occurring in the studied area and 67% of the whole Greek centipede fauna.

The discussion follows an ideal ecological-altitudinal sequence, principally according to Debazac &
Navrommatis (1971), from the Mediterranean belt up to the higher mountain areas. The following forms of forest
vegetation, loosely named from their dominant plant species, are distinguished: sclerophyllous broadleaved forests with
Pistacia lentiscus, Quercus coccifera and Quercus ilex and their stages of degradation; deciduous thermophilous and
mesothermophilous oakwoods dominated by Quercus spp.; coniferous woods with Abies cephaloniccr, broadleaved woods
with Fagus spp. and Abies sp. gr. alba. The centipede community of each vegetation type is discussed from the
faunistical and zoogeographical point of view; both general and local features are pointed out; characteristic species are
also tentatively suggested.

In Table 1 the species are listed according to their presence/absence in the four habitats considered; the altitudinal
ranges are tentative; chorotypes have been established according to Vigna ei at. (1992). SS = evergreen sclerophyllous
formations: QQ = thermophilous and mesothermophilous broadleaved woods dominated by Quercus spp.; AC = woods
dominated by Abies cephalonica', FF = broadleaved woods dominated by Fagus spp. and Abies spp. gr. alba. Chorotypes
as follows: cae = Centroasiatic-european, eur = European, seu = S-European, sie = Sibiric-european, med = Mediterranean,
tern = Turanic-european-mediterranean, turn = Turanic-mediterranean, wmd = W-Mediterranean.

RESULTS

Evergreen mediterranean sclerophyllous forests

These forests are physionomically characterized mainly by evergreen shrubs of woods.
The most important species are Pistacia lentiscus, Juniperus phoenicea, Ceratonia siliqua, in the
thermorphilous sites, between 0-600 m a.s.l., Quercus coccifera and Phillyrea media, mostly in
more inland sites up to 1000-1200 m, while Q. ilex occurs in relatively mesophilous sites, from
the sea level up to 650-800 m. Open habitats, such as phrygana, mainly derived from human
activities (fire, grazing, cutting) in the Mediterranean forest, are related to the sclerophyllous
formations. These habitats are especially characterized by Cistus spp., Phlomis spp. and
Sarcopoterium spinosum. Sclerophyllous forests occur especially in Peloponnese and in
Thessaly, but are also present in Macedonia, in Central Greece and in Thrace.

The general features of the centipede communities in the sclerophyllous and related open
habitats of the whole mainland Greece are mostly characterized by very common widespread
Mediterranean species such as Scolopendra cingulata and Scutigera coleoptrata, both invasive,
the E-Mediterranean Pleurolithobius patriarchalis, and the S-European Eupolybothrus litoralis.
Besides, Bothriogaster signata, widespread from Turan to E-Mediterranean steppes,
Himantarium gabrielis, Mediterranean, and Stigmatogaster gracilis, S-European, inhabits
especially open habitats. However, all these species are euriecious and occur also in disturbed or
thermophilous broadleaved oakwoods and firwoods, up to 2000-2100 m a.s.l.

In the sclerophyllous formations of W-Greece, the above mentioned species are locally
accompanied by the E-Mediterranean Henia hirsuta, Nannophilus ariadnae, Scolopendra
dalmatica and the S-European Lithobius hauseri, all ranging between 0 and 900 m a.s.l.
H. hirsuta is endemic to Epirus; N. ariadnae is recorded from Kerkira, Crete and Karpathos;
S. dalmatica ranges from Greece to Istria along the Adriatic Sea coasts; L. hauseri is only
known from Kerkira. In the phrygana of the Ionian Islands Cryptops trisulcatus is also present,
mainly in thermophilous habitats of W-Mediterranean regions. Otherwise, this species has been
recorded in Greece only from the S-Sporades, no mainland records are to date known and it is
possible that the Ionian and Egean populations both result from introductions. In evergreen
formations of W- and S-Greece the Mediterranean Dignathodon microcephalum and H.
bicarinata have also been recorded. In E-Greece, characteristic Lithobiids occuring in
sclerophyllous open habitats of Attica and W-Peloponnese are L. carinatus, widespread from
Palestine to Greece, and L. nigripalpis, SE-European.

Source :

CENTIPEDE FAUNA IN FORESTAL HABITATS OF MAINLAND GREECE

601

I. Centipedes from forestal habitats of mainland Greece (* = present, - = absent): species are grouped according
to their ecological affinities, m = altitudinal range; C = chorotypes. Abbreviations & explanations in the text
(Material & methods).

Species/habitats

Stigmatogaster gracilis (Meinert)

0-500

seu

Dignathodon microcephalum (Lucas)

0->?

med

Henia bicarinaia (Meinert)

0-1000

med

H. hirsuta Verhoeff

0-300

emd

Nannophilus ariadnae Attems

0-300

emd

Cryptops trisulcaius Brolemann

0-350

med

Scolopendra dalmatica C. L. Koch

0-900

emd

Lithobius carinatus L. Koch

0-800

emd

L hauseri (Dobroruka)

0-500

seu

L agilis C. Koch

200-1200

cur

Pleurolithobius patriarchalis (Berlese)

100-1 100

emd

Bothriogaster signata (Kessler)

0-1700

turn

Henia devia C. L. Koch

0-1800

emd

Geophilus carpophagus Leach

50->?

eur

Pleurogeophilus mediterranens (Meinert)

0-2100

seu

Scolopendra cingulata Latreille

0-2350

med

Scutigera coleoptrata (Linn6)

0-1100

med

Eupolybothrus litoralis (L. Koch)

0-2200

seu

Lithobius brignolii (Made)

800-1800

seu

L. microps Meinert

200-1700

seu

L. nigripalpis L. Koch

0-2500

seu

Pleurolithobius orientis (Chamberlin)

200-1 100

emd

Lithobius muticus C. L. Koch

750-2200

ceu

Geophilus insculptus Attems

1900-2100

eur

Lithobius tenebrosus Meinert

1400-1500

eur

Geophilus linearis C. L. Koch

400-1900

eur

Lithobius crass ipes L. Koch

600-1500

eur

L. latro Meinert

600-2000

seu

L. microps sensu A A.

300-1200

eur

L. peregrinus Latzel

800-1500

seu

Strigamia engadina (VerhoefD

1 100-2100

ceu

Cryptops anomalans Newport

500-2000

eur

Eupolybothrus transsylvanicus (Latzel)

700-1850

seu

E. wemeri (Attems)

700-2300

sue

Lithobius lucifugus L. Koch

1200-2400

ceu

L. mutabilis L. Koch

1000-2000

ceu

L. schuleri Verhoeff

1200-2000

seu

Schendyla montana Attems

1200-1500

seu

Clinopodes trebevicensis (Verhoeff)

500-2200

seu

Strigamia transsilvanica (Verhoeff)

1 150-1250

seu

Lithobius beschkovi Matic & Golemansky

1400

seu

L. lakatnicensis Verhoeff

1 100

seu

Himantarium gabrielis (Linne)

0-1900

med

Henia athenarum Pocock

0-1300

emd

H. illyrica (Meinert)

0-2100

ceu

Clinopodes jlavidus C. L. Koch

0-2400

cae

Pachymerium ferrugineum (C. L. Koch)

0-2300

tern

Cryptops hortensis Leach

200-1450

eur

C. parisi Brolemann

400-2400

eur

Eupolybothrus caesar (Verhoeff)

300-2100

seu

Harpolithobius anodus (Latzel)

0-2100

seu

Lithobius erythrocephalus C. L. Koch

150-2900

eur

L. lapidicola Meinert

100-2400

eur

L. viriatus Sseliwanoff

200-2400

seu

N° species (54)

602

MARZIO ZAPPAROLI

33 species altogether have been recorded from Greek sclerophyllous woodlands, and 9 of
them seem to be exclusive of such habitats. The number of species in some Q. coccifera stands
of Epirus. Macedonia and Central Greece (Ioannina, Amphitea; Kavala, Palea Kavala; Fthiotida,
Purnari) range between 9-10. Under more thermophilous conditions, such as in Pistacia
lentiscus shrubs or in phrygana, the number is much lower, with no more than 4-5 species
(Evia, Nea Artaki).

Thermophilous and mesothermophilous broadleaved oakwoods

Greece oakwoods are mainly dominated by Quercus pubescens, Q. cerris, Q. frainetto ,
Q. petrea; other broadleaved species such as Castanea sativa and Ostrya carpinifolia may also be
associated with. These formations are mainly distributed in Thrace, Thessaly and Macedonia,
from low altitudes; the extend but marginally to Peloponese, where they are mainly restricted to
the mountain areas.

The centipede communities of these woods are not easy to characterize: these habitats are
heavily influenced by human activities which have affected both floristic composition and
geographic distribution. Besides, faunistic and ecological data are still few. A large number of
very euriecious species, widespread in European and in W-Paleartic regions, such as Clinopodes
flavidus, Pachymerium ferrugineum, Cryptops hortensis , Lithobius erythrocephalus ,
L. lapidicola, has been recorded from Greek oakwoods. Species mostly common in the adjacent
vegetation types, such as Pleurolithobius patriarchalis and L. nigripalpis, especially related to the
sclerophyllous formations, or Pleurogeophilus mediterraneus and L. brignolii, more linked to
the Fagus or Abies cephalonica woods, are also present. In oakwoods of Northern regions, we
can find European or S-European species such as Geophilus linearis, L. crassipes, L.
peregrinus, L. microps sensu auct., L. viriatus, also present in coniferous or in mesophilous
broadleaved forests. In the mesothermophilous oakwoods sites of Epirus occur the S-European
Eupolybothrus caesar, but it lives also in the beech-fir forest of the same area.

To date 23 species of centipedes have been recorded altogether from Greek oakwoods, but
this value it is probably underestimated. No exclusive species are known. The number of
centipede species recorded in some Quercus stands in N-Greece ranges from 6 (Kastoria,
Gavros) to 9 (Garakas, Xanthi).

Coniferous woods dominated by Cephalonian fir and mesophilous decidous broadleaved woods
dominated by beech and fir

Owing to their relative faunistic homogeneity, the centipede communities of the two most
important forestal habitats of mountain areas in Greece are discussed together.

Firwoods, dominated by the endemic Abies cephalonica, are characteristic of the
Peloponnese, Attica and Central Greece, mainly between 550-2000 m a.s.l., according to the
exposure. Pinus nigra is sometimes mixed with Cephalonian fir.

Fagus (F. sylvatica, F. orientalis) and Abies {A. spp. gr. alba) woods are mostly
distributed in the Northern regions, along the Pindus, in E-Thessalia and N-Macedonian along
the Ml. Rodopi. Beech and fir forests are also in Chalcidice and in Mt. Pangeo. Such formations
generally begin from 850-1000 m a.s.l., but exceptionally (Chalcidice) they may be present from
200 m a.s.l. In these forests, the presence of the beech progressively increases from South to
North, whereas that of Abies relatively decreases.

The main qualitative features of the centipede communities in Cephalonian firwoods and in
beech-firwoods is largely similar. 42 species altogether have been recorded and about the 55%
of these are recorded in both habitats. Many of the species recorded in such formations are
widespread all over Europe, such as Cryptops anomalans , in Central Europe, or Strigamia
engadina, Lithobius lucifugus and L. mutabilis or in SE-Europe, Eupolybothrus werneri and L.
schuleri. The S-European E. transsylvanicus also inhabits Fagus woods as well Southernmost

Source :

CENTIPEDE FAUNA IN FORESTAL HABITATS OF MAINLAND GREECE

603

-4. cephalonica stands but Northern populations are morphologically distinguishable from
Southernmost ones (Central Greece, Eubea, Peloponnese) (ZAPPAROLI, 1994). A relatively
larger stock of Mediterranean species characterizes the A. cephalonica forests, up to 2000 m. In
the Northernmost Fagus woods European especially S-European species, tend to predominate.
However, Mediterranean species, such as Himantarium gabrielis , Henia athenarum,
Scolopendra cingulata, are also present in Mt. Pangeo (Thracia) beechwoods, especially in the
open and lowest sites of the Southern slopes.

FIGS 1-4. — Distribution of Eupolybothrus species in Greece : E. litoralis (1), E. caesar (2), E. werneri (3),
E. transsylvanicus (4).

Lithobius tenebrosus, widespread in Europe, seems to be the main characteristic species
under Cephalonian firwoods, ranging between 1100-2300 m. Five species seem to be
characteristic of the beechwoods sites: Schendyla montana , Clinopodes trebevicensis , Strigamia

604

MARZIO ZAPPAROLi

transsilvanica, Lithobius lakatnicensis, all distributed in S-Europe and up to date recorded in
Greece only from Macedonia and Thracia, between 1 100-1250 m a.s.l., and L. beschkovi , to
date known from Macedonian. L. lakatnicensis is also known only from S-Bulgarian caves and
closely related species are present in European Turkey. In N-Greece beechwoods it is also
present L. erythrocephalus borisi, a race also known from SW-Bulgaria.

The number of species recorded in Abies cephalonica woods seems to be higher as
compared to the other forestal types: 36 species have been sampled altogether but only one
seems to be exclusive of this vegetation type. In Peloponnese (Mt. Killini, Mt. Taigetos) and in
Central Greece (Mt. Parnasus) firwoods stands 13-17 species have been recorded. In Greek
beechwoods, 29 species have been recorded altogether and 6 species seem to be exclusive. In
some Macedonian Fagus woods (N-Pindus, Mt. Falakro) 8-11 species have been recorded,
whereas 10 species are at least present in Mt. Olimpos beechwoods and 18 species have been
recorded in Katara Pass (Pindus) beechwoods.

CONCLUSION

Data on Chilopoda in forestal habitats of mainland Greece are still fragmentary and both
qualitative and quantitative information is needed to reach adequate knowledge, especially of
thermophilous and mesothermophilous oakwoods communities. Difficulties in drawing a
complete picture also rise from the still incomplete faunistical and taxonomical knowledge of the
Greek fauna. Besides, because of the heavy human impact that affects the S-Balkans forestal
landscape, especiallay at the lower altitudinal sites, it is sometimes difficult to assess the original
features of animal communities.

However, according to the preliminary data discussed in this contribution, the centipede
communities inhabiting in the forestal habitat of Greece seem to show a qualitative composition
characteristic and well related to the main bioclimatic and microclimatic conditions. Some
indicators of different conditions could be recognized. Among Lithobiidae, the four Greek
species of the S-European genus Eupolybothrus (Figs 1-4), well known both from taxonomic
and faunistic point of view (ZAPPAROLI, 1994), could be tentatively used. According to their
geographical distribution and habitat preferences, they can be listed according to a decreasing
thermophilic sequence: E. litoralis present in a wide range of thermophilous habitats in mainland
as well as in insular Greece from the sea level; E. caesar only inhabiting the thermophilous and
mesothermophilous sites of Epirus from 300 m a.s.l.; E. transsylvanicus and E. werneri in
mountain forestal habitats under A. cephalonica as well as Fagus, both from 700 m. The last
three species do not occur in the Southernmost xerothermic areas such as Attika and Egean
Islands.

The analysis of the chorotypes represented in the centipede fauna altogether recorded in
each forestal habitat gives further useful informations to describe these communities (see
Fig. 5).

The European s.l. species are the most important group in all forestal habitats of mainland
Greece, their relative importance ranging from about 48% in sclerophyllous forest up to 90% in
mesophilous Fagus woods. Species mainly ranging in S-Europe show, among the European
s.l., the highest percentage in all habitats investigated and are dominant also in the more
thermophilous biotopes (30%). The percentage of the Mediterranean s.l. species is high only in
the mainland sclerophyllous forests (42%) whilst it is very low in the other forestal habitats
(13% on an average). The E-Mediterranean group is the main among the Mediterranean s.l.
species in sclerophyllous forests (24%). W-Palearctic species are poorly represented in all
studied forestal habitats and they show similar percentages (9% on an average). A decrease of
the percentage of the Mediterranean s.l. species accompanied by a progressive increase of the
European s.l. species is clearly observed following the ecological sequence from the
sclerophyllous ecosystems to the mesophilous beechwoods.

Source :

CENTIPEDE FAUNA IN FORESTAL HABITATS OF MAINLAND GREECE

605

Fig. 5. — Chorotypes (%) represented in centipede communities of forestal habitats of mainland Greece. See Table 1 for
abbreviations.

ACKNOWLEDGEMENTS

This research of the Zoological Institutes of Roma Universities in the Near East (n° 173) has been supported by a
grant of the CNR.

REFERENCES

Debazac, E. F. & Mavrommatis. G., 1971. — Les grandes divisions ecologiques de la vegetation forestikre en Grece
continentale. Bull. Soc. hot. Fr.,118 : 429-452.

MlNELLl, A. & Iovane, E., 1987. — Habitat preferences and taxocenoses of Italian centipedes (Chilopoda). Boll. Mus.
civ. St. nat. Venezia. 31 (1986) : 7-34.

Serra, A., 1978. — Contribution al conocimiento de los quilopodos de algunas zonas aridas de la peninsula iberica. Los
Monegros (Aragon). Misc. Zool.. 4 : 31-42.

Serra, A. & Ascaso, C., 1990. — Analisis de la composicion faunistica y variacion estacional de los Quilopodos de
tres habitats del Montseny (Cataluha) capturados con trampas de cai'da. In : A. Minelli. Proc. 7th Int. Congr.
Myriapodology, Leiden, Brill : 385-401.

Vigna Taglianti, A., Audisio. P. A., Belfiore, C.. Biondi, M„ Bologna, M. A., Carpaneto, G. C., De Biase, A., De
Felici, S., Piattella, E., Racheli, T.. Zapparoli, M. & Zoia. S., 1992. — Riflessioni di gruppo sui corotipi
fondamentali della W-paleartica ed in particolare italiana. Biogeographia. 16 : 159-179.

Zapparoli, M., 1992. — I Chilopodi negli ambienti forestall italiani. Monti e Boschi. 5 : 1-12.

Zapparoli. M., 1994. — Note tassonomiche e faunistiche su Chilopodi della Grecia continentale ed insulare
(Chilopoda). Fragm. Entomol.. 26 : 11-66.

Source : MNHN, Paris

On Abundance, Phenology and Natural History of
Symphyla from a Mixedwater Inundation Forest in
Central Amazonia, Brazil

Joachim ADIS *, Jose Wellington de MORAIS ** & Ulf SCHELLER ***

* Max-Planck-Instituie for Limnology, Tropical Ecology. Working Group, Postfach 165,

D- 24302 Plon,. Germany

** Instituto Nacional de Pesquisas da Amazonia (INPA), C.P. 478, 69.01 1-970 Manaus/ AM, Brazil
*** Haggeboholm, Haggesled, S-53194 Jarpas, Sweden

ABSTRACT

Inundation forests in Central Amazonia are covered by several metres of floodwater for 5-7 months each year, due to
the monomodal flood pulse of rivers. Terrestrial invertebrates have adapted to this ecosystem by evolving several
survival strategies. The Symphyla of a mixedwater inundation forest near Manaus comprised solely terricolous species,
which represented non-migrants and migrants. Migratory reaction of Hanseniella arborea Scheller, 1979 was vertical'
with temporal ascent of adults into tree trunks. The non-migrating species Symphylella adisi Scheller, 1992 and
Ribautietla amazonica Scheller, 1984 had dormant stages under water. Subadults and advanced juvenile stages of R.
amazonica spent the inundation period in naturally available retreats (inside tree roots). During the non-inundation
period, abundance of this species was highest (52.6%) when compared to H. arborea (39.4%) and .S', adisi (7.9%) of the
total catch (n = 2139 specimens). Vertical distribution of all species in the soil (0-14 cm depth), their life cycle and the
vertical migration of H. arborea are discussed with respect to abiotic factors in the study area.

RESUME

Abondance, phenologie et histoire naturelle des symphyles d’une foret inondable en Amazonie
Centrale, BresiL

Les forets inondables dc P Amazonie Centrale sont couvertes par plusieurs metres d’eau durant 5 kl mois de l’annee, &
cause de la frequence d inondations monomodales des rivieres. Les invertebres terrestres se sont adaptes a ce type
particu lier d’ecosysteme en faisant evoluer plusieurs strategies de survie. Les symphyles d’une foret inondable d’eau
mixte pres de Manaus component uniquement des esp£ces terricoles, representees par des formes migratrices et non-
migratrices. La migration de Hanseniella arborea Scheller, 1979 s’effectue verticalement, avec un d6placement
temporaire d’adultes sur les troncs d’arbres. Les especcs non-migratrices Symphylella adisi Scheller, 1992 et Ribautietla
amazonica Scheller, 1984 presentent, sous I’eau, des 6tats de dormance. Les subadultes et les stades juveniles avances de
R. amazonica passent la periode d’inondation dans des retraites riaturelles accessibles. notamment le syst&mc racinaire
des arbres. En dehors de la periode d’inondation, 1’ abondance de ces especes est plus elevee (52,6%) que celles de H.
arborea (39.4%) et de S. adisi (7,9%) par rapport au total des captures (n = 2139 indi vidus). La repartition venicale de
loutes les especes dans le sol (0-14 cm de profondeur), leur cycle de vie et la migration verticale de H. arborea sont
discutes par rapport aux facteurs abiotiques du site d’etude.

Adis, J., de Morais, J. W. & Scheller, U., 1996. — On abundance, phenology and natural history of Symphyla
Irom a mixedwater inundation forest in Central Amazonia. Brazil. In: Geoffroy, J.-J., M.AURlkS, J.-P. & Nguyen Duy -
Jacquemin, M.. (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, nat ., 169 : 607-616. Paris ISBN : 2-85653-502-X.

608

JOACHIM ADIS, JOSE WELLINGTON DE MORAIS & ULF SCHELLER

INTRODUCTION

Inundation forests in Central Amazonia and their adjacent shores are covered by several
metres of floodwater for 5-7 months each year due to the monomodal periodic flood pulse of
rivers (cf. JUNK et a/., 1989). Terrestrial invertebrates have adapted to this ecosystem by
evolving several “survival strategies” (cf. ADIS, 1992a). The fauna comprises both terricolous
and arboricolous animals. Both groups include non-migrants and migrants. Migratory reaction
of terricolous animals is horizontal (following the high water line), vertical (temporal ascent to
trunk or canopy) or includes a temporal flight to upland forests. Non-migrants have active or
dormant stages under water. The latter pass inundation in naturally available retreats, in self-
made retreats or as eggs. Non-migrant arboricolous animals reproduce and live exclusively in
the trunk and canopy region, whereas migrants include life stages that live on the ground as
well. Characteristics and examples of species for each of these categories are given by ADIS
(1992a, b). In this paper, adaptive reaction of Symphyla from a mixed water inundation forest
near Manaus to the annual flooding is discussed. Abundance, phenology and the natural history
of three species are compared with data already known from symphylans inhabiting blackwater
inundation forests in the Rio Negro valley (cf. ADIS & SCHELLER, 1984; SCHELLER & ADIS,
1984).

STUDY AREA AND METHODS

The study site was at Lago Janauarf (03°20'S. 60°17'W), situated on a spit between the Rio Negro and the Rio
Solimoes about 10 km from Manaus, across the river. The region was influenced by blackwater of the Rio Negro during
low water-level and by whitewater of the Rio Solimoes during the high water period. The study site in this seasonal
mixed water inundation forest (cf. Prance, 1979) was Oat and had no direct connection with non-flooded dryland (=
upland) areas, which were several km distant (comp. Fig. 18 in Irmler, 1975). The soil consisted of clay, predominantly
montmorillonite, which represented alluvial deposits of the Rio Solimoes. A scanty litter layer was formed during the
non-inundation period (August/September - April/May). It was mostly carried out of the forest by the current of the
annual floodwaters and/or partially covered by sediments deposited during inundation. Further information on the study
site is given by Erwin (1983), Irmler (1975) and Adis & Righi (1989).

Symphylans evaluated for this study were collected between June, 1987 and May, 1988. In 1987, inundation lasted
until end of July. The study area was not flooded again until early May, 1988. On the forest floor, four ground photo-
eclectors (= emergence traps) provided data on the activity density of symphylans. Ground photo-eclectors are round,
tent-like capture devices which cover a basal area of 1 m2. They consist of lateral plastic walls, a roof of black cloth, a
transparent collecting box on top and a pitfall trap inside the apparatus. Trunk ascents and descents were detected at bi¬
weekly intervals with arboreal photo-eclectors (= funnel traps) on one tree trunk each throughout the collecting period.
The traps consisted of four connected capturing funnels of black cloth, each with a transparent collecting box at the
funnel mouth, and formed a closed ring around the trunk. The funnel opening faced either the forest floor (for trunk
ascending animals) or the canopy (for trunk descending animals). Traps for trunk ascents were mounted on Virola
surinamensis (ROL.) Warb. (Myristiaceae) and traps for trunk descents on Hevea spruceana MOLL. Arg. (Euphorbiaceae),
two dominant tree species in the study area. The killing/preserving agent used in all photo-eclectors on soil and trunk
was aqueous picric acid solution (without detergent), which is known to be mostly neutral in terms of attraction or
repellence in temperate zones (Adis, 1979). In one of the four ground photo-eclectors, however, an aqueous
formaldehyde solution (3%) was utilized. All capture devices are fully described by Adis (1981) and Funke (1971) who also
explain their mode of utilization and function. In addition, the distribution of symphylans in the soil was studied
between August, 1987 and May, 1988 (non-inundation period). Once a month (cf. Figs 4, 8, 12) six soil samples were
taken at random along a transect with a split corer (= steel cylinder with lateral hinges; diameter 21 cm, length 33 cm),
which was driven into the soil by a mallet. Each sample of 14 cm was then divided into four subsamples of 3.5 cm each.
Animals were extracted from subsamples following a modified method of KEMPSON (Adis, 1987). In addition,
symphylans were obtained from soil samples which were taken under water in August 1988 (end of flood period) as
described above and subsequently extracted by means of a flotation method via sugar water (for methodology see Adis el
a/., 1989).

Seasonal inundation forests in Central Amazonia are subject to a rainy season (December - May: average
precipitation 1550 mm), and a "dry” season (June - November: average precipitation 550 mm, but each month has some
rain events; cf. Ribeiro & Adis, 1984). Vertical distribution of Symphyla in relation to changing conditions of soil
moisture content, temperature and pH was statistically evaluated with the linear correlation test (Cavalli-Sforza,
1972), using the original field data. This method was also used to evaluate the activity of Symphyla on the soil surface
and tree trunks in relation to insolation, precipitation, temperature and humidity of the air. The taxonomic work for this
paper was done by U. Scheller (cf. Scheller, 1979, 1992; SCHELLER & Adis, 1984), the collection and the evaluation of

Source : MNHN. Paris

SYMPHYLA FROM A MIXEDWATER INUNDATION FOREST IN CENTRAL AMAZONIA

609

field data by J. Adis and J. W. De Morais. Symphylans sampled were classified as juveniles (8. 9 and 10 pairs of legs),
subadults (1 1 pairs of legs) and adults (12 pairs of legs). For subadults and adults sex was determined according to
Chardard (1947) and Scheller (1979).

RESULTS AND DISCUSSION

A total of 2386 Symphyla were collected in the seasonal mixedwater inundation forest
under study. Out of these, 97% could be classified to species and developmental stages. The
majority were juveniles (48% of the total catch; n = 1 105), 39% were adults (n = 899) and 13%
subadults (n = 312). They comprised three species: Ribautiella amazonica Scheller, 1984 and
Symphylella adisi Scheller, 1992 (Scolopendrellidae) were found solely in the soil (by soil
extraction). Hanseniella arborea Scheller, 1979 (Scutigerellidae) was caught on the soil surface
(in ground photo-eclectors) and also during trunk ascents and descents (in arboreal photo-
eclectors).

During the non-inundation period, 79%
of all symphylans obtained by soil extraction
from the 0-14 cm cores were caught in the
top 7 cm, irrespective of their developmental
stage (cf. Fig. 1). Abundance of R.
amazonica was highest (53%; 542 ind. m-2
month-i), when compared to H. arborea
(39%: 406 ind. m-2 month- 1) and S. adisi
(8% of the total catch; 82 ind. m-2 month-i).

The highest population density, of 1853 ind.
m-2, was recorded during the dry season (in
October, 1987) and the lowest, of 260 ind.
m-2, at the beginning of the non-inundation
period (in August, 1987). Average abundance
of Symphyla in the soil of the seasonal
mixedwater forest (1766 ind. m-2 month- 1)
was about nine times higher than the average
from a seasonal blackwater inundation forest
(208 ind. m-2 month- 1; ADIS & SCHELLER,

1984; SCHELLER & ADIS, 1984).

Data from this study provided conclusive information on the life cycle, habitat and ecology
of the three Symphyla species found.

Ribautiella amazonica Scheller (Scolopendrellidae)

This terricolous non-migrating and univoltine species reaches 2.1 mm in length. It is
known to pass inundation in the soil of seasonal blackwater inundation forests inside tree roots
in a dormant state (ADIS, 1992b). This behaviour was reconfirmed for R. amazonica from the
seasonal mixedwater forest, where subadults and the last juvenile stage (10 pairs of legs) were
obtained by flotation of soil samples which were taken under water during forest inundation.
Shortly after the floodwater had receded from the forest (in 1987 at the end of July), these
symphylans had moulted to subadults and adults (Fig. 4).

Soil extraction data, indicated that reproduction took place in the early part of the non¬
inundation period, as first juveniles of the offspring occurred from September onwards, four
weeks after the forest floor had dried. R. amazonica is considered hemiedaphic, as 76% of all
specimens extracted from soil samples in 1987/88 were found in the top 7 cm, irrespective of
their developmental stage (Fig. 2). This is in contrast to data obtained from the seasonal

R. amazonica ! _ H.arborea S.adisi

Fig. 1. — Distribution of Symphyla species in the soil.
Monthly samples taken every 3.5 cm to a depth of
14 cm between August, 1987 and April, 1988 (non¬
inundation period) in a seasonal mixedwater
inundation forest at Lago Janauarf; total catch =
100%.

610

JOACHIM ADIS. JOSE WELLINGTON DE MORAIS & ULF SCHELLER

blackwater forest, where the species was considered euedaphic, as 86% of all specimens
occurred below 7 cm soil depth (ADIS, 1992b).

Fig. 2. — Ribautietla amazonica Scheller (Scolopendrellidae): Distribution of developmental stages in the soil at Lago
Janauan'. Monthly samples taken every 3.5 cm to a depth of 14 cm between August, 1987 and April, 1988 (non-
inundation period); total catch = 100%.

Fig. 3. — Ribautiella amazonica Scheller (Scolopendrellidae): Percentage of developmental stages caught in the soil (0-
14 cm depth). Monthly samples taken between August, 1987 and April. 1988 (non-inundation period) at Lago
Janauan.

Fig. 4. — Ribautiella amazonica Scheller (Scolopendrellidae): Temporal occurrence and abundance of developmental
stages (ind./m2) in the soil (0-14 cm depth). Monthly samples taken between August. 1987 and April, 1988
(non-inundation period) at Lago Janauan,

Fig. 5. — Ribautiella amazonica Scheller (Scolopendrellidae): Temporal occurrence and abundance (ind./m2) of
specimens in the soil at Lago Janauarf. Monthly samples taken every 3.5 cm to a depth of 14 cm between
August. 1987 and April, 1988 (non-inundation period).

About 64% of the Ribautiella population in the seasonal mixedwater forest was
represented by juveniles (Fig. 3), 12% by subadults and 24% by adults. The sex ratio of males
and females was 1: 1.2 (n = 268). Population density was lowest two weeks prior to inundation
(April 29, 1988: 250 ind. m-2), when most of the symphylans must have entered tree roots to
pass flooding in a dormant state. Abundance was highest eight weeks after inundation had
ended (September 30, 1987: 1,477 ind. m-2), due to a high number of juveniles of the progeny in
the upper 7 cm of the soil (Figs 4, 5).

There was no clear correlation between the vertical distribution of R. amazonica in the soil
throughout the non-inundation period and the abiotic factors monitored in the study area.

Source MNHN, Paris

SYMPHYLA FROM A MIXEDWATER INUNDATION FOREST IN CENTRAL AMAZONIA

611

Symphylella adisi Scheller (Scolopendrellidae)

This is the smallest of the three species collected in the seasonal mixedwater inundation
forest reaching a maximum of 1.5 mm in length. It was only found in the soil but was recorded
from all soil layers sampled (Figs 6, 9). About 68% of its population was represented by
juveniles, 1 1% by subadults and 21% by adults (Fig. 7). The sex ratio of males and females
was 1: 1.1 (n = 35). The population density was lowest two weeks prior to forest inundation
(April 29, 1988: 5 ind. m-2) and highest during the early rainy season (February 2, 1988: 327
ind. m-2). There was no correlation found between the population density of S. adisi and
weather conditions in the area (dry season versus rainy season).

Ind./m2

350

300

250

200

150

100

3 8 87 1 9 30.9 29.10 112 30 12. 12.88 1.3. 30 3. 29 4 88

Fig. 6. — Symphylella adisi Scheller (Scolopendrellidae): Distribution of developmental stages in the soil at Lago
Janauarf. Monthly samples taken every 3.5 cm to a depth of 14 cm between August, 1987 and April, 1988 (non¬
inundation period); total catch = 100%.

Fig. 7. — Symphylella adisi Scheller (Scolopendrellidae): Percentage of developmental stages caught in the soil (0-14
cm depth). Monthly samples taken between August, 1987 and April, 1988 (non-inundation period) at Lago
Janauarf.

Fig. 8. — Symphylella adisi Scheller (Scolopendrellidae): Temporal occurrence and abundance of developmental stages
(ind./m2) in the soil (0-14 cm depth). Monthly samples taken between August, 1987 and April, 1988 (non¬
inundation period) at Lago Janauarf.

Fig. 9. — Symphylella adisi Scheller (Scolopendrellidae): Temporal occurrence and abundance (ind./m2) of specimens
in the soil at Lago Janauarf. Monthly samples taken every 3.5 cm to a depth of 14 cm between August, 1987 and
April, 1988 (non-inundation period).

It is presumed, that S. adisi passed forest inundation in a dormant state in the soil. Due to
its low abundance, it has not yet been located in soil samples taken under water during the flood
period and the type of its retreat (naturally available or self-made) is still unknown. Six weeks
after the forest floor had dried, adults and early juvenile stages of their offspring (8 and 9 pairs

612

JOACHIM ADIS. JOSE WELLINGTON DE MORAIS & ULF SCHELLER

of legs) were found in 3.5 - 14 cm soil depth (Figs 8, 9). Subadults occurred from end of
October onwards. The highest number of juvenile stages was observed during the early rainy
season (in February; Fig. 8) and main reproduction is assumed to occur at this time. The
abundance of subadults subsequently declined and advanced juvenile stages (9 and 10 pairs of
legs) as well as subadults dominated (Fig. 8). Ten weeks prior to inundation, only a few
specimens were found and those solely in the lowest of the soil layers sampled (10.5 - 14 cm;
Fig. 9). The main part of the population most probably had entered retreats at this time and, due
to the dormant state, could not be obtained by means of the soil extraction method.

Based on the characteristics outlined, S. adisi is considered a terricolous non-migrating
and univoltine species with dormant stages (presumably of subadults and advanced juveniles) in
retreats under water during annual inundation.

Hanseniella arborea Scheller (Scutigerellidae)

N - 843

0 - 3.5cm 3.5 - 7.0cm 7.0 - 10.5cm 10.5 - 14.0cm

juv. 8.9.10

ad.

60 N - 843
50

juv.8 juv.9

Ind. /nr
1200

1000

800

600

400

200

N ■ 4057

juv.8 03 juv.9 juv. 10 Wb s.ad.11 EM ad.

87. 1.9. 30.9 29.10. 1.12. 30.12 1.2.88 1.3. 30.3. 29.4.

0-3.5 cm 3.5-7 cm 7-10.5 cm wm 10.5-14 cm

Fig. 10. — Hanseniella arborea Scheller (Scutigerellidae): Distribution of developmental stages in the soil at Lago
Janauarf. Monthly samples taken every 3.5 cm to a depth of 14 cm between August, 1987 and April, 1988 (non¬
inundation period); total catch = 100%.

Fig. 11. — Hanseniella arborea Scheller (Scutigerellidae): Percentage of developmental stages caught in the soil (0-14
cm depth). Monthly samples taken between August, 1987 and April, 1988 (non-inundation period) at Lago
Janauarf.

Fig. 12. — Hanseniella arborea Scheller (Scutigerellidae): Temporal occurrence and abundance of developmental stages
(ind./m2) in the soil (0-14 cm depth). Monthly samples taken between August, 1987 and April, 1988 (non¬
inundation period) at Lago Janauarf.

Fig. 13. — Hanseniella arborea Scheller (Scutigerellidae): Temporal occurrence and abundance (ind./m2) of specimens in
the soil at Lago Janauarf. Monthly samples taken every 3.5 cm to a depth of 14 cm between August, 1987 and
April. 1988 (non-inundation period).

This terricolous, migrating and univoltine species reaches 2.7 mm in length. Adults are
known to pass inundation in the trunk/canopy area of seasonal blackwater inundation forests

Source : MNHN, Paris

SYMPHYLA FROM A MIXEDWATER INUNDATION FOREST IN CENTRAL AMAZONIA

613

(ADIS & SCHELLER, 1984). This behaviour was reconfirmed for H. arborea from the seasonal
mixedwater inundation forest (Fig. 14).

In the soil, about 51% of the population
caught during the non-inundation period was
represented by adults (the migrating stage), 31%
by juveniles and 18% by subadults. The sex ratio
of males and females was 1 : 1.5 (n = 429). H.
arborea is considered hemiedaphic, as 51% of all
specimens extracted from soil samples in
1987/88 were found in the top 3.5 cm and 90%
in 0-7 cm soil depth, irrespective of their
developmental stage (Figs 10, 13). Data differs
somewhat from results obtained in the seasonal
blackwater forest, where H. arborea was most
abundant in 3.5 - 7 cm soil depth (47% of the
total catch), and where only 12% of all
specimens were obtained from the top 3.5 cm
(ADIS & SCHELLER, 1984). First analysis of
grain size and mineral composition of soils from
mixed- and blackwater inundation forests (ADIS

6 IRION, unpubl.) indicated, that a lower
abundance of H. arborea in the soil layers
sampled corresponded with a greater presence of
grains < 2 |im, due to a higher amount of clay.
In the seasonal mixedwater forest under study,
decrease in population density of H. arborea
from the top 3.5 cm to 14 cm soil depth (cf.
Fig. 10) was correlated with an increase of
grains < 2 pm and of clay from 32% to 49% per
soil layer (p < 0. 10 for the total catch, p < 0.05
for monthly catches of Oct. 10 and Dec. 12,
1987 & March 1 and 30, 1988). In the seasonal
blackwater inundation forest, abundance of H.
arborea was highest where the amount of grains
< 2 ftm and of clay was the lowest (12% in 3.5 -

7 cm soil depth) as compared to the soil layers
below (25 - 31% in 7-14 cm depth), where
abundance was lower (cf. Fig. 3 in ADIS &
SCHELLER, 1984).

It is suggested, that H. arborea might
have difficulties inhabiting fine and clayey soil
layers due to its relatively large size.

Fig. 14. — Hanseniella arborea Scheller (Scutigerellidae):
Activity density of males and females on the forest
floor (4 ground photo-eclectors: (E); ind./m2), trunk
descents (BE i) and trunk ascents (BE T); one arboreal
photo-eclector, respectively) between July. 1987 and
June, 1988 at Lago Janauarf.

200

Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun.
: precipitation (mm) * insolation (hrs.)

BEt (ind.)

N ■ 72

o — -

trunk ascents

lili Eicj Uli lili 1

trunk descents

BEt (ind.)

N ■ 50

Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar Apr. May

E (ind./m2

N ■ 43

activity density
-forest floor-

Jul. Aug. Sep Oct Nov. Dec Jan Feb. Mar Apr May Jun
>987 ,988

ma,es females inundation period non~lnundatlon period

614

JOACHIM ADIS, JOSE WELLINGTON DE MORAIS & ULF SCHELLER

The lowest abundance of this species in soils from the mixedwater area was observed
during the beginning of the non-inundation period (August 3, 1987: 14 ind. m-2). H. arborea
was at this time represented by adults which had returned from the trunk/canopy area after
flooding and recolonized the upper 3.5 cm of the forest floor (Figs 12-14). Juvenile stages and
subadults of the progeny were detected only four weeks later (Fig. 12). Their abundance was
highest during the beginning of the rainy season (in December/January), shortly after the main
reproduction must have occurred. Adults dominated between February and April (Fig. 12) when
they came to the surface (where they were caught in ground photo-eclectors) and started
ascending tree trunks, about 12 weeks prior to forest inundation (Fig. 14). The predominance of
females assured the continuation of this species. During periods of less insolation, the number
of adults caught in arboreal photo-eclectors was lower (p < 0.05). No significant correlation
was found between catch numbers and the increasing water-level of the Rio Negro (cf. ADIS et
al, this volume). In the seasonal blackwater inundation forest, adults of H. arborea occurred
more frequently in ground photo-eclectors after heavy rainfalls, compared to drier periods (p <
0.05; cf. ADIS & SCHELLER, 1984).

Phenology data of H. arborea from this mixedwater study area supports the life cycle
proposed for this species based on results obtained by ADIS & SCHELLER (1984) from the
seasonal blackwater inundation forest.

CONCLUSIONS

Four terricolous species of Symphyla were found to inhabit the soils of different non-
flooded upland forests in Central Amazonia: Scolopendrellopsis tropicus , Symphylella adisi,
Ribautiella amazonica (Scolopendrellidae), and Hanseniella arborea (Scutigerellidae; cf.
SCHELLER, 1992).

Results of this paper show, that three of them succeeded in colonizing seasonally
inundated forests of the mixedwater and blackwater type in the Rio Negro valley. The survival
strategies which have evolved include a migratory state, represented by adults which pass
inundation in the trunk/canopy area (Scutigerellidae) and a dormant state, represented by
advanced juvenile and subadult stages which pass the flood period under water in retreats
(Scolopendrellidae). The concentration of dissolved oxygen in the water body near the soil is
known to be so low that flood resistant symphylans may have to switch from plastron and
cutaneous respiration to anaerobic respiration (ADIS & MESSNER, 1991; MESSNER & ADIS,
1988, 1992). The first and second larval stages (6 and 7 pairs of legs respectively) of the three
symphylan species may be inactive (TlEGS, 1945) or may only last a short time, as they were not
obtained by means of the soil extraction method. Only the migrating species H. arborea was
observed to respond to abiotic changes in its environment. Activity density on the soil surface
and vertical migration were somewhat triggered by these secondary ecofactors which are no
longer directly related to the cycle of flooding, and to which many terrestrial invertebrates of
inundation forests have become sensitive (cf. ADIS, 1992a). The flood pulse (JUNK et al., 1989)
is regarded as the original determinant of trunk ascents and descents. However, it was found to
act as the primary control mechanism or ecofactor among a few invertebrate species only (ADIS,
1992a).

Evaluation of capture data also indicated, that the three symphylan species changed from a
plurivoltine mode of life in upland forests (ADIS, MORAIS, RODRIGUES & SCHELLER, unpubl.)
to an univoltine life cycle in inundation forests. This was also observed in Pseudoscorpiones
and Meinertellidae (Archaeognatha) from the same study areas (ADIS et al., 1988; ADIS &
STURM, 1987). It remains to be investigated if the species of Symphyla in inundation forests
already differ ecologically and phenologically, as well as genetically, to such a great degree from
those in upland forests that they must be regarded as new species or subspecies (cf. WOLF &
ADIS, 1992).

Source :

SYMPHYLA FROM A MIXEDWATER INUNDATION FOREST IN CENTRAL AMAZONIA

615

ACKNOWLEDGEMENTS

anH p, b ld r 3 °ur colleagues at the National Institute of Amazonian Research (INPA) in Manaus (Brazil)

-rck-,nrute for L'mnol°gy m Plon (Germany) who, in the field or laboratory, contributed to this
r ’ fe*P“'al: y. S“e Hamann, Irmgard Adis, Edilson De Araujo Silva and M. Sc. Elizabeth Franklin Dr. Helen
Read (Bucks, United Kingdom) kindly corrected the English manuscript. This study was supported by a grant from the
German Academic Exchange Service (DAAD) lor the second author and from the Max-Planck-Society for the third author
We wish to acknowledge the valuable support received by PD Dr. W. J. Junk, head of the Tropical Ecology Working
Group at the Max-Planck-Institute for Limnology in Plon, Germany. ^

REFERENCES

Adis, J., 1979. Problems of interpreting arthropod sampling with pitfall traps. Zoo/. Anz., 202 • 177-184

“udi“ of *• “rres,,ial * c«rai a““» *—>•«»

ADIS, J., 1987. — Extraction of arthropods from Neotropical soils with a modified KEMPSON apparatus. J. Trop. Ecol.,

Adis J 1992a. - Uberlebensstrategien terrestrischer Invertebraten in Uberschwemmungswaldern Zentralamazoniens
Verh. natunviss. Ver. Hamburg, 33 : 21-114.

Adis. J., 1992b. — How to survive six months in a Hooded soil: Strategies in Chilopoda and Symphyla from Central

MNTFrntn nv°°dPiainS- ' 1 forln & S' TaNAKA’ Symposium on life-history traits in tropical invertebrates.
[INTECOL, Yokohama, Japan 1990. Studies on Neotropical Fauna and Environment, 27], Lisse, Swets &
Zeitlinger : 117-129

ADIS J„ Mahnert, V MORAIS, J. W. de & Rodrigues, J. M. G„ 1988. - Adaptation of an Amazonian pseudoscorpion
(Arachmda) from dryland forests to inundation forests. Ecology, 69 : 287-291.

Aims J. & Messner, B 1991. — Langzeit-Uberflutungsresistenz als Uberlebensstrategie bei terrestrischen
A^rtnropoden. - Beispiele aus zentralamazonischen Uberschwemmungsgebieten. Disch. ent. Z., N. F.. 38 : 211-

Adis, J. MESSNER, B. & Groth I., 1989. — Zur Uberflutungsresistenz und zum Spinnvermogen von Japygiden
(Diplura). Zool. Jb. Anat., 119: 371-382. °

Adis, J. & Righi, G., 1989. — Mass migration and life cycle adaptation - a survival strategy of terrestrial earthworms in
central Amazonian inundation forests. Amazoniana, 11 : 23-30.

Adis, J. & Scheller, U. ,1984. — On the natural history and ecology of Hanseniella arborea Schellcr, (Myriapoda,
ym phyla, Scutigerellidae), a migrating symphylan from an Amazonian black-water inundation forest
Pedobiologia, 27 : 35-41.

Adis, J & Sturm, H., 1987. — On the natural history and ecology of Meinertellidae (Archaeognatha, Insecta) from
dryland and inundation forests of Central Amazonia. Amazoniana, 10 : 197-218.

Cavalli-Sforza, L., 1972. — Grundzuge biologisch-medizinischer Statistik. Stuttgart. G. Fischer, 212 pp.

Chardard, R., 1947. — Nouvelles stations de Symphyles, distinction et rapport numerique des sexes. Bull. Mus Natl
Hist. nat. Paris, (2), 19: 177-184.

Erwin. T. L. 1983. — Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by insecticidal

m2' S‘ V,' ^UTT0N' r C- Whi™ORE & A. C. Chadwick, Tropical Rain Forest: Ecology and Management.
Oxlord, Blackwell Scientific Publications : 59-75.

Funke W., 1971. — Food and energy turnover of leaf-eating insects and their influence on primary production
Ecological Studies ,2: 81-93. v j v

Irmler, U., 1975. — Ecological studies of the aquatic soil invertebrates in three inundation forests of Central
Amazonia. Amazoniana, 5 : 337-409.

Jijnk, VV. J Bayley, P. B. & Sparks. R. E., 1989. — The flood pulse concept in river-floodplain systems. Can. Spec.
Publ. Fish. Aquat. Sci., 106 : I 10-127. r

Messner. B & Adis, J., 1988. — Die Plastronstrukturen der bisher einzigen submers lebenden Diplopodenart
uonographis adisi Hoffman 1985 (Pyrgodesmidae, Diplopoda). Zool. Jb. Anat.. 117 : 277-290.

Messner, B. & Adis, J., 1992. Die Plastronatmung bei aquatischen und flutresistenten terrestrischen Arthropoden
(Acari, Diplopoda und Insecta). Mitt. Dtsch. Ges. Allg. Angew. Ent., 8 : 325-327.

Prance, G. T., 1979. — Notes on the vegetation of Amazonia III. The terminology of Amazonian forest types subject
to inundation. Brittonia, 31 : 26-38.

616

JOACHIM ADIS, JOSE WELLINGTON DE MORAIS & ULF SCHELLER

RlBElRO, M. DE N. G. & Adis, J., 1984. — Local rainfall variability - a potential bias for bioecological studies in the
Central Amazon. Acta Amazonica, 14 : 159-174.

SCHELLER, U., 1979. — Hanseniella arborea n. sp., a migrating symphylan from an Amazonian blackwater inundation
forest (Myriapoda, Symphyla. Scutigerellidae). Acta Amazonica. 9 : 603-607.

Scheller. U., 1992. — A study of neotropical Symphyla (Myriapoda): list of species, keys to genera and description of
two new Amazonian species. Amazoniana, 12 : 169-180.

Scheller, U. & Adis. J., 1984. — A new species of Ribautiella (Myriapoda, Symphyla, Scolopendrellidae) trom an
Amazonian black-water inundation forest and notes on its natural history and ecology. Amazoniana, 8 : 299-310.

TiEGS. O. W., 1945. — The post-embryonic development of Hanseniella agilis (Symphyla). Quart. Jour. Micros. Sci.,
85 : 192-328.

Wolf, H. G. & Adis, J., 1992. — Genetic differentiation between populations of Neomachilellus scandens
(Meinertellidac, Archaeognatha, Insecta) inhabiting neighbouring forests in Central Amazonia. Verb, naturwiss.
Ver. Hamburg, 33 (NF) : 5-13.

Source : MNHN. Paris

The Ecology of Savanna Millipedes in Southern Africa

John Mark DANGERFIELD * & Steven R. TELFORD **

* Department of Biological Sciences, University of Botswana
PB 0022, Gaborone, Botswana
** Department of Zoology, University of Pretoria
Pretoria 0002, South Africa

ABSTRACT

Millipedes of the order Spirostreptida are abundant and diverse in the savanna habitats of southern Africa. Large body
size (2-20g live mass), mobility and considerable local abundance of up to 30 individuals per square meter are features of
several species. Life cycles and behaviours are strongly influenced by the seasonality of the climate whilst short term
changes in atmospheric conditions induce bouts of conspicuous surface activity during summer. The cylindrical body
plan is efficient for burrowing and enables individuals to find shelter in the soil during the dry winter. Energy efficient
mobility allows for opportunistic foraging tactics within a generalist feeding strategy. These tactics can result in the
development ol high density aggregations on suitable food sources. Few species appear to be exclusively litter feeders
although some species specialise on algae. Relatively low assimilation efficiencies require high ingestion rates and in
some habitats the turnover of detrital material by millipedes may have a significant influence on nutrient cycling.
Millipede species diversity, in particular relative abundance, varies between habitats and appears to be related to the
composition and vertical expression ot the vegetation. Millipede communities in savannas are likely to be input driven
but biotic interactions, including interspecific competition, may also be important.

RESUME

L’ecologie des diplopodes de savanne en Afrique du Sud.

Les diplopodes de l’ordre Spirostreptida sont abondants et diversifies dans les dcosystemes de savane d’Afrique du Sud.
La grande taille du corps (2 a 20 g de masse fraiche), la mobilite et fabondance considerable, d6passant localement 30
indi vidus par metre carre, caracterisent plusieurs especes. Le cycle biologique et le comportement paraissent fortement
influences par 1 aspect saisonnier du climat tandis que les changements ct court terme des conditions atmospheriques
induisent des reprises d activite manifestes en surface durant fete. La forme cylindrique du corps est efficace pour
I'enfouissement et permet aux individus de trouver des abris dans le sol durant 1’hiver sec. L’6nergie allouee k la mobilite
tacilite les tactiques opportunistes de recherche de la nourriture au sein d’une strategic alimentaire gSneraliste. Ces
tactiques peuvent provoquer fapparition d’une distribution fortement agregative avec de fortes densites sur les sites de
repartition des ressources alimentaires. Quelques especes apparaissent comme des consommatrices exclusives de litiere
bien que quelques-unes se special isent dans la consommation des algues. Les rapports dissimilation relativement bas
requierent des taux d’ ingestion 61ev6s et, dans certains milieux, le renouvellement des materiaux organiques fragments
par les diplopodes peut avoir une influence significative sur le cycle des Elements mineraux. La diversite specifique des
diplopodes, notamment fabondance relative des especes, varie selon les milieux et semble etre reliee a la composition et
a la repartition verticale de la vegetation. II semble que forganisation des peuplements de diplopodes des savanes soit
largement determince par les apports energ&iques mais les interactions biotiques, incluant notamment la competition
interspecifique, peuvent jouer aussi un role important dans felaboration de ces communautes d’arthropodes.

Dangerfield, J. M.,& Telford, S. R., 1996. — The ecology of savanna millipedes in Southern Africa. In:
Geoffroy. J.-J., Mauries, J.-P. & Nguyen Duy- Jacquemin, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist .
nat.. 169 : 617-625. Paris ISBN : 2-85653-502-X.

618

JOHN MARK DANGERFIELD & STEVEN R. TELFORD

INTRODUCTION

In southern Africa the savanna environment covers a large area north of the tropic of
Capricorn. A wide range of habitats occur characterized by a drought resistant vegetation mixture
in which C4 grasses potentially dominate the ground layer and woody plants vary in density
from widely scattered individuals to a closed canopy woodland broken occasionally by drainage
line grasslands (HUNTLEY, 1982). The functioning of these systems is driven primarily by the
availability of water which is supplied seasonally in summer rainfall (October-April), although
rainfall pattern and intensity are spatially and temporally unpredictable in volume and intensity
(VOSSEN, 1988). Also characteristic is a dry and cool winter season. The severity and frequency
of fires (TROLLOPE, 1982), soil nutrient status (SCHOLES, 1990) and a wide range of secondary
factors influence the vegetation structure which varies on a number of spatial scales from
100's km in response to rainfall gradients through 10's km in response to soil type and
geomorphology down to 1 0's m due to localised disturbance and nutrient enrichment by termites
(Campbell et al, 1988).

Although millipedes are abundant and diverse in these habitats (LAWRENCE, 1984;
DANGERFIELD & TELFORD, 1992) their ecology is poorly known. Savanna ecosystems present
ecological conditions for detritivores that place a premium on 1) avoidance of moisture stress,
particularly desiccation, 2) resource acquisition and 3) the ability to withstand unpredictable
stress including temporary food shortages.

Hence, in this paper we summarise our work on savanna millipedes in relation to the
evolutionary pressures created by this type of environment. We also consider the influence of
millipedes on the nutrient dynamics of savannas.

STUDY SITES

Much of the current study was carried out on three contrasting sites: 1) heterogeneous miombo woodland at Marondera,
Zimbabwe, dominated by the canopy trees Brachystegia spiciformis and Julbernardia globiflora described in detail by
Campbell et al (1988); 2) Acacia savanna in south-east Botswana dominated by Acacia erubescens , Dichrostachys
cinerea , and Euclea undulata (Dangerfield, Milner & Matthews, 1993) and 3) riparian savanna in south-east Botswana
dominated by Combretum erythrophyllum (Dangerfield & Milner, in press). A number of additional sites were studied
including managed areas derived from miombo woodland (arable fields, pasture and plantations) and natural savanna
types in Zimbabwe (riparian dominated by Celtis africana, Burkea africana woodland and Colophospermum mopane
woodland) and Botswana ( Baikiaea plurijunga woodland, C. mopane woodland and suburban habitats).

COMPOSITION AND ABUNDANCE OF THE MILLIPEDE FAUNA

Approximately 350 millipede species are thought to occur in southern Africa (LAWRENCE,
1984). The majority in savannas are juliform species of the order Spirostreptida, in particular
members of the Spirostreptidae, Odontopygidae and Harpagophoridae approximately in the ratio
of 2:1:1. Occasionally polydesmids, a group common in west Africa (LEWIS, 1971) and India
(Bhakat, 1989), can be found in the more moist habitats (>1000 mm rainfall) or synanthropic
sites.

Many of the southern African spirostreptids are characterised by large body size,
frequently in excess of 2.0 g fresh mass and in some species, such as Alloporus uncinatus
(Attems), up to 20.0 g. Although LAWRENCE (1984) considers that these animals have localised
distributions we have found several species over wide geographic areas. A. uncinatus is
common from Zululand to north-east Zimbabwe and across to southern Botswana, an area of
more than 10,000 km2, similarly separate populations of Calostreptus carinatus have been
recorded throughout this zone. However, along with the taxonomy (R. L. HOFFMAN, pers.
comm.), the biogeography of these species remains poorly known.

The abundance of millipedes in savannas varies greatly between the habitat types (Table 1)
and, together with other soil animals, many species appear to be sensitive to habitat change,

ECOLOGY OF SAVANNA MILLIPEDES

619

particularly when natural habitats are converted for human land use (Dangerfield 1990) In

• hnnH^nre •de"sities can reach 30 Ind- m'2> although accurate estimates of

abundance are difficult to obtain because some species remain cryptic even during the rainv

season, hence more than one sampling method is required for accurate estimates of abundance
and small-scale distribution appears to be highly aggregated. The range in abundance of <1 to 35
ind. m-2 is similar to that observed for other tropical species (BANERJEE, 1980; Bhakat, 1989)

Kr I s H N amo O R t*hV h f 9 8 5) ^ ^ reC°rded for JonesPeltis splendidus (Verhoeff) (Bano &

' ABLE in7 Ab7da"ce estimates (individuals m-2 ± I standard error) for millipede assemblages in southern African grass

"t nlrrlnr , TTmT S0UrCfS) 0 Danghrfield (1990); 2) Dangerfield, Milner & MATTHEWS (1993);
3) dangerfield & Milner (in press). 4) Dangerfield (unpublished data) and 5) Dangerfield & Telford (1993).

Habitat

Abundance

Sampling method

Miombo woodland

26 ±5

soil monolithsl

Acacia savanna

0.4 "

visual inspection of
permanent quadrats2

Riparian savanna

12 ± 3

litter quadrats (lm2)3

Teak woodland
( Baikiaea plurijunga)

2 ± 1

soil monoliths4

Riparian savanna
{Acacia erioloba)

25 ±5

soil monoIiths4

Mixed riparian

22 ±5

soil monoliths4

Mopane

( Colophospermum mopane)

33 ±5

soil monoliths4

Plantation
(Pinus patula)

0.3

visual inspection of
permanent quadrats^

SURFACE ACTIVITY

Most adult spirostreptids, odontopygids and harpagophorids show distinct patterns of
surface activity during the summer rainfall season from October to April (DANGERFIELD &
Telford, 1991; Dangerfield, Milner & Matthews, 1993). During such activity
individuals are conspicuous on the surface or can be located beneath leaf litter at the soil litter
interface. Frequently individuals climb trees, shrubs and tall grasses. This behaviour may be to
avoid waterlogged conditions, particularly in riparian habitats with clay rich soils; to access
additional food sources such as microbial populations; or to avoid mates (see TELFORD &
Dangerfield, this volume). A. uncinatus, Chaleponcus limbatus and C. digitatus can be added
to two other southern African pill millipedes, Sphaerotherium cinctellum and S. punctulatum,
that have been observed climbing trees (HaaCKER & FUCHS, 1972),

The onset of surface activity appears to be initiated by the second major rainfall event
(>20 mm) of the season as few animals are seen active after the first rainfall. Significant rainfall
may be necessary for moisture to percolate the desiccated upper soil horizons to depths of up to
30 cm, where the animals overwinter, and break aestivation. An alternative hypothesis is that

620

JOHN MARK DANGERFIELD & STEVEN R. TELFORD

delayed emergence may avoid late dry season fires which are a common feature of savannas.
Not all species emerge at the same time. In Acacia savanna Calostreptus carinatus , which feeds

primarily on algae, emerges four to five weeks

0 N D J F M

Fig. 1 . — Percentage of total observations (ordinate %) in
three categories; walking (O — O), feeding (□ —
□) and resting (• — •) for Alloporus uncinatus
(A), Chaleponcus digi talus (B) and Calostreptus
carinatus (C) during the 1991-92 rainfall season
in Acacia savanna, south-east Botswana.

after the first A. uncinatus and Chaleponcus
digitatus are recorded which suggests that food
availability may also influence emergence
pattern.

There are considerable species specific
differences in activity patterns within the
rainfall season, and between site differences in
overall millipede activity. In miombo
woodland surface activity, measured as
relative abundance, peaks in early December
and then declines steadily (DANGERFIELD &
TELFORD, 1991) whilst in wetter riparian
habitats the early season peak occurs but the
activity decline is less pronounced (TELFORD
& DANGERFIELD, 1993). In the semi-arid
Acacia savanna of south-east Botswana where
rainfall is more sporadic millipede activity is
pulsed, initially in response to each rainfall
event but during February and March
substantial rainfall events fail to produce
significant millipede activity (DANGERFIELD,

Milner & Matthews, 1993). Reddy &
VENKATAIAH (1990) also found positive
correlations between millipede abundance in
pitfall traps and rainfall and soil moisture in
both grass and tree savanna in India.

During surface activity individuals of all
species were observed walking, feeding,
resting, burrowing or copulating. The
frequency with which these activities were
observed varied between species and with sex
(DANGERFIELD, MILNER & MATTHEWS,
1993; DANGERFIELD & KAUNDA, 1994).
Seasonal variation in the proportion of
observations of these behaviours in Acacia
savanna is illustrated in Figure 1. Alloporus
uncinatus was most frequently observed
moving around the habitat and mobility
appeared to increase through the season at the
expense of feeding. In Chaleponcus digitatus
feeding and walking were equally important
and there was a higher frequency of
individuals resting. Calostreptus carinatus also
increased mobility during the season to the
extent that in February and March no
individuals were seen resting.

Source : MNHN, Paris

ECOLOGY OF SAVANNA MILLIPEDES

621

FEEDING BEHAVIOUR

In field observations of behaviour during surface activity up to 52% of all records were of
individuals feeding. A wide range of food sources were utilized including, tree leaf litter, seeds,
grass litter, Aloe grandidentata litter, soil, algae, faeces and dead invertebrates which confirms
for savanna species the wide dietary range seen in other diplopods (COLE, 1946; LEWIS, 197 L
Wooton & Crawford, 1975; Pobozsny, 1986 & review by Crawford, 1992). A feature
of these food types is that they represent a range of quality in terms of both energy and nutrient
content from relatively poor quality grass litter, known to retard growth and reproductive output
in woodlice (RUSHTON & HASS ALL, 1983), to the nutrient rich cotyledons of Acacia seeds.
Given such a range of foods energetic returns will vary with both the choice and quantity of
material ingested.

In a series of laboratory experiments a number of species were fed partially ground
Combretum erythrophyllum leaf litter and gravimetric estimates of ingestion and assimilation
recorded (DANGERFIELD & MILNER, 1993). Individuals ingested between 2.6 and 7.6% of their
dry body mass each day and assimilated around 10 to 25% of this material (Table 2), results that
are consistent with those recorded for other millipede species (see references in DANGERFIELD &
Milner, 1993). The discrepancy between ingestion and assimilation is considerable. Variation
in ingestion explains only 35 to 53% of variation in assimilation. The proportion of soil in the
diet influences assimilation rates in A. uncinatus (DANGERFIELD, 1993), whilst available
moisture, temperature, barometric pressure and food quality may also be important. There is
much to be gained from experiments designed to consider the fitness consequences of specialist
and catholic diets in detritivores.

Table 2. — Mean body size (mg dry mass) and estimates of mean (± i s.e.) ingestion rate (I, mg day-i), assimilation rate
(A, mg day-i) and assimilation efficiency (AE, %) for five species of savanna millipede fed ground Combretum
erythrophyllum litter. Data source: Dangerfield & Milner (1993).

Family/Species

mean body mass

Spirostreptidae
Alloporus uncinatus
Calostreptus carinatus

2.86 ±0.14

0.22 ± 0.03

75 ±7

17 ± 2

1.5 ± 2.9

4.3 ± 1.3

18 ± 3
26 ±6

Odontopygidae
Chaleponcus limbatus

C. digitalus

0.44 ± 0.04

0.88 ±0.10

30 ±7

34 ±5

7.7 ± 1.5

7.1 ± 4.6

14 ± 4
7 ± 9

Harpagophoridae
Zinophora sp.

0.98 ± 0.06

32 ±2

5.6 ± 0.8

16 ± 3

Inspection of the proportions of different foods utilized in Acacia savanna (Table 3)
suggest that there may be a strong selection of food items which may vary according to
frequency of food types and environmental conditions. In the 1990-91 rainfall season^A.
uncinatus was most frequently observed eating seeds of the canopy tree Acacia erubescens. As
mast-years are irregular in A. erubescens , this represents an opportunistic feeding tactic by A.
uncinatus which in 1991-92 reverted to a more mixed diet. Similarly C. carinatus specialised on
algae in 1991-92, when rainfall was 55% below the 25 year average, but also ate seeds and leaf
litter in 1990-91 (Table 3). This suggests that even the more selective feeders are not obligate
specialists.

During a study of the mating system in a population of A. uncinatus inhabiting riparian
savanna in Zimbabwe (TELFORD & DANGERFIELD, 1993) several aggregations of up to 42
individuals were observed. These aggregates were not associated with the mating system but
appeared to be part of the feeding tactics of immature individuals (DANGERFIELD & TELFORD,

622

JOHN MARK DANGERF1ELD & STEVEN R. TELFORD

1993). In a subsequent field experiment similar aggregates were generated by the addition of
high quality food, fruits of the highveld tree Uapaca kirkiana , to a pine plantation habitat with
limited herbaceous cover. After 48 hours, 15 to 20 millipedes, mostly A. uncinatus, were
feeding on the fruits in an area where background densities were 0.3 individuals m-2, hence the
addition of high quality food attracted individuals from an area of 140 m2 (DANGERFIELD &
TELFORD, 1993).

Table 3. — Percentage of total feeding observations on six different food types for three species of millipede in
ungrazed Acacia savanna in south-east Botswana in the 1990-91 and 1991-92 rainfall seasons, n is the total
number of feeding observations. Data sources: Dangerfield, Milner & Matthews (1993); Dangerfield &
Kaunda (1994).

Alloporus

uncinatus

Calostreptus

carinatus

Chaleponcus

limbatus

90/91

91/92

90/91

91/92

90/91

91/92

leaf litter

seeds

1 1

grass litter

1 1

Aloe grandidentata

1 1

algae/soil

faeces

131

107

1 17

In savannas the distribution of high quality food resources for detritivores is spatially and
temporally heterogeneous. High quality leaf litter often decays rapidly at the beginning of the
rainy season (M. J. SWIFT, pers. comm.) whilst faeces, fruits and fungal fruiting bodies are
spatially patchy and ephemeral. Such conditions would favour the evolution of opportunistic
feeding tactics in detritivores that would occasionally give the impression of considerable
specialisation (e.g. C. carinatus, Table 3). In turn, such a tactic requires cost effective mobility
and sophisticated sensory discrimination mechanisms.

CONSEQUENCES OF FEEDING ACTIVITY

Combinations of millipede abundance, activity, body mass, faecal pellet production rate
and pellet mass show that savanna millipedes produce 30 to 60 g m-2, of faecal material annually,
which may be up to 40% of the litter standing crop (DANGERFIELD & MILNER, in press).
Because of changes in particle size this conversion of leaf litter to faeces promotes moisture
retention (McBRAYER, 1973) and, given the importance of moisture to decomposition processes
(SWIFT, Heal & ANDERSON, 1979), faecal pellets are likely to decompose faster than
uningested leaf litter. As the greater proportion of leaf litter is ingested during the first two
months of the rainfall season (DANGERFIELD & TELFORD, 1991) the decomposition of litter is
enhanced at a time when the nutrient demand of the vegetation is high in response to the seasonal
growth of perennials and establishment of annual plants (MALAISSE, 1978). Millipedes may
thus be a factor in the evolved synchrony between nutrient release and uptake typical of natural
systems on poor soils where the bulk of the nutrients are stored in the vegetation and transient

Source :

ECOLOGY OF SAVANNA MILLIPEDES

623

soil organic matter. A reduction in millipede abundance due to land use changes (Lavelle &
PASHANASI, 1989; DANGERFIELD, 1990) would contribute to the disruption of this synchrony.

This effect on nutrient cycling combined with heterogeneous distribution of millipedes,
including feeding aggregations, suggest that millipede feeding activities may reinforce small-
scale patterns in the vegetation (e.g. BELSKY, 1983) and nutrient dynamics (CAMPBELL et al.,
1988) that already exist in savannas by enhancing plant growth in areas favoured by millipedes'
Such a process would be analogous to the effect of nutrient accumulation in termitaria (see
review by JONES, 1990). Experimental studies on millipede micro-distribution and correlations
with the distribution of root systems and soil nutrients would provide an important test of the
evolution of plant responses to the effects of soil fauna.

SPECIES DIVERSITY AND COMPETITION

Collections of surface active individuals from various savanna habitats contained between
one and five millipede species (DANGERFIELD & TELFORD, 1992). Repeated observations,
cryptozoan trap sampling (sensu COLE, 1946) together with hand sorting of soil samples
suggest that total millipede species richness can be as high as 10 in some habitats.

Managed habitats have fewer millipede species than the natural systems from which they
were derived, a pattern seen in other soil fauna groups (DANGERFIELD, 1990), and the rare
species in the natural systems appear to be those that are lost. Cosmopolitan species such as A.
uncinatus and several in the genus Chaleponcus appear to be less affected by habitat change,
although variation in abundance or behaviour between habitats have not been investigated in
detail. This combination of both sensitive and robust species, together with a significant yet
manageable number of species in a given habitat, makes millipedes a potentially useful taxa for
the monitoring of biodiversity.

Surface living detritivores are often considered to be largely regulated by abiotic factors
and rarely compete for food resources (ANDERSON, 1977; WARBURG, LlNSENMAIR &
BERKOVITZ, 1984). KlME & WAUTHY (1984) have shown that percentage clay and mean
annual temperature are good predictors of the numerical organisation of temperate millipede
assemblages. Two features of savanna millipedes suggest that biotic interactions may also be
important determinants of abundance and species composition. Firstly, the ability of several
species to show dietary specialisation and opportunism, a common process leading to niche
separation, and thereby avoiding competition. Secondly, the differences in relative abundance of
a species between habitats and the close approximation of species abundance plots to a geometric
series model (DANGERFIELD & TELFORD, 1992), which suggests that resource pre-emption and
competition may structure millipede assemblages. Tests of these hypotheses require long term
monitoring and experimental manipulations.

MILLIPEDES IN SAVANNA ENVIRONMENTS

In strongly seasonal environments resident organisms must evolve mechanisms to survive
regular periods of stress. In savannas spirostreptid millipedes avoid up to six months without
rainfall by burrowing into the soil. MANTON (1977) has established that the multi-legged
configuration of juliform millipedes is able to exert considerable forward force which is well
suited to burrowing. In this respect large body size is advantageous. The very large (40 g live
mass) Orthoporus spp. of the semi-arid savannas appear not to burrow but use the vent
structures of Macrotermes and Odontotermes termitaria to overwinter hence there may be an
upper limit to burrowing ability and size. The importance of burrowing may mean that the
distribution of the larger spirostreptids, as with temperate julids (KlME & WAUTHY, 1984) may
be restricted to light textured soils.

Large body size also allows considerable mobility. This is important for opportunistic
feeding tactics in heterogeneous environments or to access widely spaced shelter sites. If dry

624

JOHN MARK DANGERFIELD & STEVEN R. TELFORD

season fire has removed the litter layer emerging millipedes must either move to new areas or
utilize alternative food sources such as fresh shoots or ungulate dung, likely to be available as
the grass flushes in response to the nutrients released by fire. Mobility also allows for extensive
and rapid dispersal. We have observed individuals cover more than 30 m in one hour and
although no data are available, movement from natal habitats may be a key feature of several
species. There is some evidence that in managed environments fragments of natural habitat
contain higher densities of millipedes (DANGERFIELD, 1990). Such areas may act as refuges and
are favoured by mobile species.

Organisms that for the most part consume poor quality foods but have life history tactics
that require large adult body size are likely to be long lived, particularly if periods of active
foraging are restricted. We have kept adult specimens of A. uncinatus in laboratory culture for
30 months without significant mortality and suspect that most species live at least four years and
possibly up to 10 years. Although there is a lengthy list of potential predators (reduviid bugs,
suricates, ververids, large amphibians and hornbills) few feed exclusively on millipedes. The
apparent lack of intense adult mortality combined with high female fecundity of up to 600 eggs
female-1 (DANGERFIELD & TELFORD, unpublished data) suggest that, as with other
invertebrates, density-independent juvenile mortality is likely to be high.

The spirostreptid millipedes in southern African savannas offer valuable opportunities for
organism centred ecological studies. The present phase of inductive research has provided
information on the ecology of a little known group which invites comparisons with the juliform
species living in temperate habitats. Our analyses have also generated many hypotheses, both
theoretical and organism centred, that are readily testable in this system.

ACKNOWLEDGMENTS

We are very grateful to Peter Chibatamoto, Trevor Cooper, Samson Kaunda, Bowdin King, Charity Mackyii,
Tarombera Mwabvu. Busani Ndlela for field and laboratory assistance.

We thank the Department of Research and Specialist Services, Zimbabwe Ministry of Lands, Agriculture and Rural
Resettlement for permission to work at the Grasslands Research Station, Marondera and the Department ol National
Parks and Wildlife Management for a permit to collect animals from the national parks in Zimbabwe together with the
research boards of the University of Botswana and the University of Zimbabwe for financial support. We are also grateful
to SAREC for logistical support provided through the TSBF programme.

REFERENCES

Anderson, J. M., 1977. — The organisation of soil animal communities. Ecol. Bull. (Stockholm) , 25 : 15-23.
Banerjee, B.. 1980. — The population characteristics of the East African red-legged diplopod Metiche tanganyciense
Kraus (Diplopoda: Spirobolidae) Res. Popul. Ecol., 21 : 306-316.

Bano, K. & Krishnamoorthy, R. V.. 1985. — Reproductive strategy and life history of Jonespeltis splendidus
(Verhoeff) (Diplopoda: Polydesmida). J. Soil Biol. Ecol., 5 : 48-57.

Belsky, A. J., 1983. — Small-scale pattern in grassland communities in the Serengeti National Park, Tanzania.
Vegetatio ,55 : 141-151.

Bhakat, S. N., 1989. — The population ecology of Orthomorpha coarctata (Diplopoda: Polydesmidae). Pedohiologia,
33 : 49-59.

Campbell, B. M., Swift, M. J., Hatton, J. C. & Frost P. G. H., 1988. — Small-scale vegetation pattern and nutrient
cycling in miombo woodland. In: J. T. A. Verhoeven, G. W. HEIL & M. J. A. Merger, Vegetation structure in
relation to carbon and nutrient economy. The Hague, SPB Academic Publishing : 69-85.

Cole, L. C., 1946. — A study of the cryptozoa of an Illinois woodland. Ecol. Monogr. , 16 : 49-86.

Crawford, C. S., 1992. — Millipedes as model detritivores. Ber. nat.-med. Verein Innsbruck, suppl. 10 : 277-288.
Dangerfield, J. M., 1990. — Abundance, biomass and diversity of soil macrofauna in savanna woodland and associated
managed habitats. Pedobiologia, 34 : 141-151.

Dangerfield, J. M., 1993 — Ingestion of mineral soil/litter mixtures and faecal pellet production in the southern
African millipede Alloporus uncinatus (Attems). Pedobiologia ,37 : 159-166.

Dangerfield, J. M. & Kaunda, S. K., 1994 — Millipede behaviour in a savanna woodland habitat in south-east
Botswana. African J. Ecol., 32 : 337-342.

Source : MNHN, Paris

ECOLOGY OF SAVANNA MILLIPEDES

625

Danger FI eld, J. M. & Milner, A. E., 1993. — Ingestion and assimilation of leaf litter in some tropical millipedes. J.
Zool. , 229 : 683-693.

Dangerfield, J. M. & Milner, A. E., (in press) — Millipede faecal pellet production in selected natural and managed
habitats of southern Africa: Implications for litter dynamics. Biotropica .

Dangerfield, J. M. & Telford S. R., 1991. — Seasonal activity patterns of Julid millipedes in Zimbabwe. J. Trop.
Ecol. , 7 : 281-285.

Dangerfield, J. M. & Telford, S. R., 1992. — Species diversity of Julid millipedes: Between habitat comparisons
within the seasonal tropics. Pedobiologia ,36 : 321-329.

Dangerfield, J. M. & Telford S. R., 1993. — Aggregation in the tropical millipede Alloporus uncinatus (Diplopoda:
Spirostreptidae). J. Zool., 230.

Dangerfield, J. M., Milner, A. E. & Matthews, R.. 1993. — Seasonal activity patterns and behaviour of juliform
millipedes in south-eastern Botswana. J. Trop. Ecol., 8 : 451-464.

HaaCKER, U. & Fuchs, S., 1972. — Tree-climbing in pill-millipedes. Oecologia , 10 : 191-192.

Huntley. B. J., 1982. — Southern African Savannas. In : B. J. Huntley & B. H. Walker, The ecology of tropical
savannas. Berlin. Springer-Verlag : 101-119.

Jones, J. A., 1990. — Termites, soil fertility and carbon cycling in dry tropical Africa: A hypothesis. J. Trop. Ecol., 6 :
291-305.

Kime, R. D. & Wauthy, G., 1984. — Aspects of the relationship between millipedes, soil texture and temperature in
deciduous forests. Pedobiologia , 26 : 387-402.

LAVELLE, P. & Pashanasi, B., 1989. — Soil macrofauna and land management in Peruvian Amazonia (Yurimaguas,
Loreto). Pedobiologia ,33 : 283-291.

Lawrence, R. F., 1984. — The Centipedes and Millipedes of southern Africa. A Guide. Cape Town, A. A. Balkema,
148 pp.

Lewis, J. G. E.. 1971. — The life history and ecology of three paradoxosomatid millipedes (Diplopoda: Polydesmida) in
northern Nigeria. J. Zool.. 165 : 431-452.

Malaisse, F., 1978. — The miombo ecosystem. In: Tropical Forest Ecosystems. Unesco/UNEP/FAO Natural Resources
Research XIV . Paris, Unesco : 589-606.

MANTON, S. M., 1977. — The Arthropoda. Habits, Functionnal Morphology and Evolution. Oxford. Oxford University
Press, 527 pp.

McBrayer, J. F.. 1973. — Exploitation of deciduous leaf litter by Apheloria montana (Diplopoda: Eurydesmidae).
Pedobiologia , 13 : 90-98.

Pobozsny, M., 1986. — Uber Streuzersetzungsprozesse in Hainbuchen-Eichenwaldern unter Berucksichtigung der
Diplopoden. Opusc. Zool., Budapest., 22 : 77-84.

Reddy, V. M., Venkataiah, B.. 1990. — Seasonal abundance of soil-surface arthropods in relation to some
meteorological and edaphic variables of the grassland and tree-planted areas in a tropical semi-arid savanna. Int. J.
Biometeorol., 34 : 49-59.

Rushton, S. P. & Hassall, M., 1983. — The effects of food quality on the life history parameters of the terrestrial
isopod Armadillidium vulgare (Latreille). Oecologia . 57 : 257-261.

Scholes, R. J., 1990. — The influence of soil fertility on the ecology of southern African dry savannas. J. Biogeogr.,
17 : 415-419.

Swift, M. J., Heal O. W. & Anderson. J. M.. (Eds) 1979. — Decomposition in terrestrial ecosystems. Oxford,
Blackwell Scientific Publ.

Telford, S. R. & Dangerfield J. M., 1993. — Mating tactics in the tropical millipede Alloporus uncinatus (Diplopoda:
Spirostreptidae). Behaviour, 124 : 45-57.

Trollope, W. S. W., 1982. — Ecological effects of Fire in South African savannas. In: B. J. Huntley & B. H. Walker,
The ecology of tropical savannas. Berlin, Springer-Verlag : 292-306.

Vossen, P., 1988. — An agrometeorological contribution to quantitative and qualitative rainy season quality
monitoring in Botswana. Ph. D. Thesis, Stale University, Gent.

Warburg, M. R., Linsenmair. K. E. & Berkovitz, K.. 1984. — The effect of climate on the distribution and abundance
of isopods. Symp. Zool. Soc. London , 53 : 339-363.

Wooton, R. C. & Crawford, C. S., 1975. — Food, ingestion rates and assimilation in the desert millipede Orthoporus
ornatus (Girard) (Diplopoda). Oecologia , 20 : 231-236.

Source : MNHN, Paris

The Diplopod Community of a Mediterranean Oak
Forest in Southern France: Ecological and
Evolutionary Interest

Jean-Frangois David

CNRS, Centre d’Ecologie fonctionnelle et evolutive
1919, Route de Mende. B.P. 5051, F-34033 Montpellier cedex 01, France

ABSTRACT

Seven millipede species have been found in a holm oak forest ( Quercus ilex ) sampled for two years. Seasonal changes
in abundance are discussed in relation to the periods of recruitment and the changes in vertical distribution of species.
Details of the life cycle of Opisthocheiron elegans are given. The biomass of macrosaprophagous species (on average
10.5 g live mass/m2 tor Glomeris marginaia and Cylindroiulus caeruleocinctus) is the highest recorded in Europe, which
points to the ecological importance of millipedes in Mediterranean forests. The millipede populations from the site
studied differ intraspecifically from those found further north, as shown in Polytonium germanicurn. The significance of
this geographical variation is discussed.

RESUME

Le peuplement de diplopodes d'une foret de chene-vert dans le sud de la France.

Le peuplement de diplopodes d'une foret de chene vert ( Quercus ilex) est decrit apres deux annees d’6chantil!onnage. 11
comprend cinq especes regulieres et deux especes occasionnelles. Les variations saisonnieres d'abondance sont
examinees en liaison avec les pSriodes de recrutement et les variations de la distribution verticale des populations.
Quelques donnees sur le cycle biologique d' Opisthocheiron elegans sont presentees. La biomasse des especes
macrosaprophages presentes — en moyenne 10.5 g/m2 (masse fraiche) pour Glomeris marginaia a Cylindroiulus
caeruleocinctus — est la plus 61evee de toutes celles mesurees en Europe, ce qui souligne l'importance ecologique des
diplopodes dans certaines forets mediterraneennes. Les populations de diplopodes de la station etudiee se differencient
intraspecifiquement de celles trouvees plus au nord, comme le montre le cas de Polyzonium germanicurn. La signification
de cette variation geographique est discutee.

INTRODUCTION

The work of JANATI-lDRISSI (1988) on litter consumption in Mediterranean ecosystems
drew attention to the millipede community of the forest of Puechabon, Herault, on limestone
hills near- Montpellier. Several interesting aspects emerged: firstly, the abundance of Glomeris
and Cylindroiulus populations was very high; secondly, the presence of Polyzonium
germanicurn was recorded, which was surprising for a species associated with waterlogged
conditions near the centre of its west-European range (David, 1990).

David, J.-F., 1996. — The diplopod community of a mediterranean oak forest in Southern France: ecological and
evolutionary interest. In: Geoffroy, J.-J.. MauriSs, J.-P. & Nguyen Duy - Jacquemin, M., (eds), Acta
Myriapodologica. Mem. Mus. natn. Hist, nat ., 169 : 627-634. Paris ISBN : 2-85653-502-X.

628

JEAN-FRANgOIS DAVID

As information on millipede communities in French Mediterranean forests remained scanty
(Bigot & BODOT, 1973; Saulnier & ATHIAS-BINCHE, 1986; JANATI-IDRISSI, 1988), further
investigations were carried out at Puechabon for two years. Results have been obtained on three
points: (1) the seasonal changes in abundance have been described in terms of individuals and in
biomass (DAVID, 1995), which will be summarized in the present paper; (2) the extraordinary
abundance of macrosaprophagous species in the Mediterranean region has been confirmed;
(3) some traits distinguishing the local populations from others of the same species have been
underscored, as will be shown in P. germanicum.

SITE AND METHODS

The study site is located in the state forest of Puechabon. 25 km northwest of Montpellier, at an altitude ol
260 m. The climate, though rather humid with a cool winter, is typically Mediterranean. Rainfall is minimal in summer,
with a period of severe drought almost every year; the summer dryness index of EmberGER (1943) is < 7. A red tersiallitic
soil overlies karstic limestone; its clay content is high, as is the proportion of stones. The humus form is a mull, and
measurements from nearby sites have given pH ~ 7 and C/N = 16 (Floret et al. , 1989; Merzouki et a/., 1989). The lorest
is a holm oak coppice (Quercus ilex) with a typical understorcy of shrubs and herbs made up of several tens of species
(Braun-Blanquet, 1936; Dugrand, 1963).

Sampling was carried out in an area of about 3000 m2 surrounding a cleared plot with a CNRS meteorological
station. Eight samples, each including between 10 and 12 sampling units, were taken over two years: two in spring (30
April, 1 99T and 1 May. 1992), two in early summer (5 June, 1991 and 23 June, 1992), two in early autumn (7 October,
1991 and 1 October. 1992) and two in winter (3 January. 1992 and 29 January, 1993). Each sampling unit consisted of
two parts: (i) litter and soil down to about 2 cm deep, collected in a 25 x 25 cm quadrat (1/16 m2); (ii) soil between 2 and
10 cm deep, collected with a cylindrical corer 10 cm in diameter (=1/127 m2). The largest millipedes were sorted by hand
and the others extracted by Tullgren funnels in the laboratory. Species were determined according to Demange (1981).

In order to assess the fresh biomass of Chilognatha, as many individuals as possible were weighed alive after
cleaning on moist Filter paper, then their species, sex and stadium were determined. For dead animals, the mean live mass
corresponding to their species, sex and stadium was used.

The differences in abundance were compared statistically by means of parametric (Student’s t-test) and non-
parametric (Mann-Whitney U-test) methods. The differences were considered significant if both tests gave a probability
P< 0.05.

RESULTS AND DISCUSSION

The millipede species

A total of seven specifes were found at Puechabon. Four were present in all the samples:
Glomeris marginata (Villers), Cylindroiulus caeruleocinctus (Wood), Polyzonium germanicum
Brandt and Polyxenus lagurus\ Linne. Another, Opisthocheiron elegans Ribaut, was present in
seven out of eight samples. Two other species were found occasionally on the site: one specimen
of Ommatoiulus rutilans (C. L. Koch) was collected in the samples and several others by hand,
which made it possible to determine an adult male; two specimens of Polydesmus sp. were
collected by hand.

JANATI-IDRISSI (1988) reported the presence of Cylindroiulus londinensis (Leach),
Ommatoiulus sahulosus (Linne) and Blaniulus guttulatus (Fabr.), in addition to G. marginata
and P. germanicum. In fact, C. caeruleocinctus was obviously mistaken for C. londinensis , as
often happens in the literature, and there might also have been confusion of the two
Ommatoiulus species. As for the blaniulid, it was not found during the present study.

Population density

Overall, the average number of millipedes and its standard-error, stadium I not included,
was 667 ± 56 ind./m2, 556 ± 55 of which belonged to Chilognatha and 111 ± 1 6 to Penicillata.
The average biomass, for Chilognatha only, was 1 1 .0 ± 1.1 g/m2. From samples taken between

1 Southern France is inhabited by the sexual form (NGUYEN DUY - JACQUEMIN, 1973).

Source : MNHN. Paris

THE DIPLOPOD COMMUNITY OF A MEDITERRANEAN OAK FOREST

629

1984 and 1986 and sorted by hand, which yielded the largest individuals, JANATI-IDRISSI
(1988) had recorded a much lower number of Chilognatha (95 ind./m2 on average) but a barely
lower biomass (2.8 g dry mass! m2, i.e. between 8 and 9 g/m2).

The seasonal population densities of the five main species are given in Table 1. For each
season the data from the two years are pooled, as most differences between years were not
significant. There were significant seasonal changes in the population density of two species: (i)
the number of G. marginata was lower in spring than in early summer (P < 0.05) and early
autumn (P < 0.01 ); likewise, its biomass was lower in spring than in early autumn (P < 0.01 for
t-test; P < 0.05 for U-test); (ii) the number of P. lagurus was lower in early autumn than in early
summer (P < 0.01) and winter (P < 0.05). In all other species, the seasonal variations in
numbers of individuals and in biomass were not significant.

Table 1. — Seasonal abundance of the main millipede species at Puechabon (after David, 1995). The results are in

number and biomass / m2± standard-error, (s.u.: sampling units).

SPRING

(21 s.u.)

EARLY
SUMMER
(22 s.u.)

EARLY
AUTUMN
(22 s.u.)

WINTER

(24 s.u.)

MEAN

G. marginata

Ind./m2

100+ 16

216 ± 44

297 ± 47

228-± 47

212 ± 22

g/m2

4.0 ± 0.7

7.3 ± 1.6

11.5 ± 2.1

8.0 ± 2.2

7.8 ± 0.9

C. caeruleocinctus

Ind./m2

50 ± 11

124 ± 88

54 ± 1 1

38 ± 12

66 ±22

g/m2

3.3 ± 1.0

2.2 ± 1.3

3.1 ± 0.8

2.4 ± 0.9

2.7 ± 0.5

P. germanicum

Ind./m2

352 ± 124

311 ± 118

161 ±62

247 ± 63

267 ± 47

g/m2

0.4 ± 0.1

0.7 ± 0.2

0.4 ± 0.2

0.4 ± 0.1

0.5 ±0.1

O. elegans

Ind./m2

21 ± 14

16 ± 10

1 ± 1

6 ± 3

11 ±4

g/m2

P. lagurus

Ind./m2

119 ± 37

164 ±37

42 ± 13

121 ±30

111 ± 16

Total Diplopoda

lnd./m2

643 ± 132

831 ± 142

556 ± 76

640 ± 86

667 ± 56

Total Chilognatha

g/m2

7.8 ± 1.2

10.3 ± 2.4

15.0 ± 2.4

10.8 ± 2.5

11.0 ± 1.1

Aspects of the seasonal dynamics

Seasonal variations in population density are not easily explained, for they depend on both
actual changes in abundance {e.g. recruitment; mortality) and apparent changes ( e.g . burrowing
into the deep soil). Both are considered in the following discussion — keeping in mind that the
seasonal changes in the vertical distribution of species were measured in the daytime, and
without data regarding the height of summer. The five most abundant species all exhibit different
patterns.

G. marginata generally lives near to the surface, except in winter when it tends to go
deeper (Fig. 1). The vertical distribution seems the same as in Great Britain, on the opposite side
of its range (BOCOCK & HEATH, 1967). Recruitment occurs during warm months, as shown by
the high number of stadium II individuals (8 tergites; 8 leg pairs) in the samples from early
summer and early autumn. The species seems to withstand the summer drought easily, since
there is a continuous increase in abundance from spring to autumn, both in number and in
biomass (Table 1). On the other hand, G. marginata has difficulty in coping with winter cold:

630

JEAN-FRANCOIS DAVID

EARLY

SUMMER

EARLY

AUTUMN

G.marginata

C. caeruleocinc tus

P.germanicum

P.lag urus

O.elegans

while moving down in the soil, it undergoes a fall in abundance, which becomes minimal in
spring (Table 1).

C. caeruleocinctus appears to live a little
deeper than G. marginata in spring and autumn
but, like many julids in the temperate zone, it
burrows markedly during summer and winter
(Fig. 1). A batch of the smallest individuals
(stadium IV; 3 rows of ocelli) were found in
one sampling unit from the early summer of
1991, between 2 and 10 cm in depth, which
may be indicative of the recruitment period.
But if the young stay in the deep soil, that may
explain why significant seasonal changes in
density are difficult to detect.

P. germanicum is the most subterranean
species at Puechabon (Fig. 1). Contrary to
what could be assumed on the basis of
Table 1, recruitment does not start in winter,
but in summer; many brooding females were
observed in spring, and stadium I individuals
in early summer. But again, seasonal changes
in density are difficult to detect because the
whole cycle appears to occur deep in the soil;
moreover, the spatial distribution is highly
contagious (the variance to mean ratio reaches
a maximum in this species).

In contrast, P. lagurus appears to
remain in the litter and upper soil layer
throughout the year (Fig. 1). The species
seems sensitive to summer drought for its
population density is significantly reduced in
early autumn (Table 1). The subsequent
increase during winter is suggestive of
recruitment in the course of autumn, but the
samples were not frequent enough to follow
this fast-developing species.

The population density of O. elegans is
low and the proportions given in Figure 1 are
quite uncertain. Nevertheless, the species
appears to live in the soil in spring and early
summer, then to move upwards in autumn and
winter (Fig. 1). These vertical displacements
are concomitant with different stages in the life cycle, which looks annual at Puechabon
(Table 2). Adults emerge by early autumn at the surface; they breed in winter, as shown by the
appearance of the young of stadium II in January, still at the surface; then growth occurs deeper
in the soil during drier months.

Fig.

1. — Seasonal changes in the vertical distribution of
the main millipede species at Puechabon. The
results are expressed as percentages of the
population density (ind./m2) in two soil layers: (i)
litter and top-soil down to 2 cm (above the line);
(ii) soil between 2 and 10 cm deep (under the line).

Ecological importance of saprophagous species

It is interesting to compare the abundance of macrosaprophagous millipedes — those
which fragment litter for feeding, i.e., mainly Glomerida, Julida and Polydesmida — in forests

Source : MNHN, Paris

THE DIPLOPOD COMMUNITY OF A MEDITERRANEAN OAK FOREST

631

of different climatic regions of Europe. In the Atlantic zone, the density of these species is very
variable, but peak biomasses are usually below 4 g/m2. The highest figures of fresh biomass
recorded in Great Britain and northern France are, respectively, 3.9 g/m2 in autumn samples
from a mixed mull-like moder (BLOWER, 1979) and 2.3 g/m2in spring and autumn samples
trom an oak mull (David, 1989). The figure of 7.5 g/m2 in a Danish beech mull provided by
BORNEBUSCH (1930) may be an overestimate, owing to the approximate value of individual
biomasses. Similarly, the highest biomass figure recorded in more continental regions of
western Europe is an annual mean of 4 g/m2 for all Diplopoda, in a mixed oak wood in Austria
(Meyer etal., 1984).

Although data are more scarce in European regions of the Mediterranean, they are also
very variable. Nevertheless, peak biomasses are higher than in the Atlantic zone, notably in
climatic transition areas. IATROU (1989) has measured a density of 114 ind./m2 for
macrosaprophagous millipedes in a community of northern Greece where Glomeris balcanica is
dominant, which should correspond to a substantial biomass (above 7.1 g/m2, the figure for
G. balcanica alone). The biomass of macrosaprophagous species at Puechabon is higher than all
those mentioned above, on average 10.5 g/m2 for Glomerida and Julida.

Provided that the ingestion rate is of the same order of magnitude as in the conditions
prevailing in the Atlantic zone (about 10 g dry litter/g fresh mass/year — (Van DER Drift,
1975; David, 1987), millipedes probably play a very important role in Mediterranean forest
ecosystems. So it should be very interesting to pursue the ecological studies which have been
undertaken on litter consumption by millipedes in that region (BERTRAND etal ., 1987- JANATI-
Idrissi, 1988).

Table 2. — Seasonal changes in the stadial composition of O. elegans at Puechabon (ind./m2).
From stadium II in winter, individuals proceed to the adult stage in autumn,
through stadium V in spring and stadium VII in early summer.

(5/2)

III

(7/3)

(10/4)

STADIUM

(and number of rings)

V VI VII

(14/4) (18/4) (22/3)

(25/2)

(27/2)

WINTER

SPRING

EARLY SUMMER

EARLY AUTUMN

Particularity of local populations

From the taxonomic point of view, the millipede community of Puechabon does not differ
from typical atlantic communities, since all the species present can be found further north. This
similarity is misleading, however, for conspecific populations can be very different between the
two areas. For example, if the P. germanicum population from the forest of Puechabon is
compared with a population from the forest of Orleans, in the Centre of France, substantial
differences emerge as regards colour and body size: (1) individuals from Puechabon are of paler
yellowish colour than those from the forest of Orleans; (2) the difference in body size is striking,
which can be shown by comparing the fresh weight of individuals in relation to their number of
rings (Fig. 2); in both sexes, individuals from Puechabon are much smaller, the number of rings
being equal.

632

JEAN-FRANCOIS DAVID

Such intraspecific differences can be explained in two ways. Either there has been
selection for different genotypes in the two populations, which would be closely adapted to local
conditions, or there is phenotypic plasticity within the species for traits like colour and body
size, in which case individuals from the two populations maintained in the same conditions
would give the same phenotypes. Only cultures under controlled conditions could help to
distinguish the correct explanation.

Fig. 2. — Comparison of the curvilinear regressions between fresh mass (FW) and number of podous rings, in
P. germanicum populations from the forests of Orleans (black) and Puechabon (white). Individuals were collected
from October to May on both sites.

However, the special selective forces that act at the periphery of a species range, in
ecologically marginal areas, generally favour genetic differentiation of populations, regarding
many morphological, physiological, behavioural or demographic traits (MAYR, 1963). As all the

Source : MNHN, Paris

THE DIPLOPOD COMMUNITY OF A MEDITERRANEAN OAK FOREST

633

species present at Puechabon but P. lagurus are near the driest boundary of their geographical
range (MAURIES, 1964; MAURIES & GEOFFROY, 1982; KlME, 1990a, b), it is tempting to
suggest that such a genetic differentiation has occurred. This is a very likely scenario in the case
of P. germanicum , which is probably a small isolate at Puechabon given the species' rarity in the
Montpellier region.

Irrespective of the process actually involved, different factors can exert an influence on soil
animals in Mediterranean areas (Dl CASTRI, 1973). On one hand, the effects of climatic factors
such as summer drought can be felt in species remaining near the surface of the soil. On the
other hand, constraints associated with an endogeic way of life (e.g. low availability of
resources) may be the driving force in species which avoid the severity of climate by their
burrowing behaviour. As regards P. germanicum, its living in the soil in the Mediterranean
forest (Fig. 1) contrasts strongly with its high abundance in the litter in the forest of Orleans
(DAVID & COURET, 1985). Therefore, the differences in body size and colour between the two
populations — whether they are genetic adaptation or phenotypic characteristics due to
acclimatory or developmental responses — may result from their living in microhabitats differing
in depth rather than in climate.

ACKNOWLEDGEMENTS

I am grateful to Prof. J.-P. LUMARET (University of Montpellier III) for information on the study site; to J.-J.
Geoffroy and J.-P. Mauries (MNHN. Brunoy and Paris) for the identification of some specimens; to M. Grandjanny
and F. Romane (CEFE-CNRS. Montpellier) for providing local climatic data; and to H. Read for improvements in
English.

REFERENCES

Bertrand, M., Janati-Idrissi, A. & Lumaret. J.-P.. 1987. — Etude experimental des facteurs de variation de la
consommation de liliere de Quercus ilex L. et Q. pubescens Willd. par Glomeris marginata (V.) (Diplopoda,
Glomeridae). Rev. Ecol. Biol. Sol , 24 : 359-368.

Bigot, L. & Bodot, P.. 1973. — Contribution a l'&ude bioccenotique de la garrigue a Quercus coccifera. II. Composition
biotique du peuplement d'Invertebres. Vie & Milieu. Revue d'ecologie, 23 ; 229-249.

Blower. J. G., 1979. — The millipede fauna of two British limestone woods. In : M. Camatini, Myriapod Biology,
London, Academic Press : 203-214.

BOCOCK, K. L. & Heath, J., 1967. — Feeding activity of the millipede Glomeris marginata (Villers) in relation to its
vertical distribution in the soil. In : O. Graff & J. E. Satchell, Progress in soil biology , North Holland,
Amsterdam . 233-240.

Bornebusch, C. H., 1930. — The fauna of forest soil. Forstl. Forsogsvaes. Dan.. 11 : 1-256.

Braun-Blanquet, J., 1936. — La foret d'Yeuse languedocienne ( Quercion ilicis). Monographic phytosociologique.
Mem. Soc. Etud. Sci. nat. Nimes, 5 ; 1-147.

David, J. F., 1987. — Consommation annuelle d'une litidre de chene par une population adulte du Diplopode
Cylindroiulus nitidus. Pedobiologia, 30 : 299-310.

David, J. F., 1989. — Les peuplements de Diplopodes d'une foret temperee : variations spatiales et stabilite dans le
temps. Rev. Ecol. Biol. Sol, 26 : 75-90.

David, J. F., 1990. — Habitat dimensions of Diplopoda in a temperate forest on acid soil. Rev. Ecol. Biol. Sol , 27 ; 95-
112.

David, J. F., 1995 . — Seasonal abundance of millipedes in a Mediterranean oak forest (southern France). Isr. J. Zool.,
41 : 23-31.

David, J. F. & Couret, T., 1985. — Le cycle biologique du Diplopode Polyzonium germanicum Brandt, 1831
(Polyzoniida). Rev. Ecol. Biol. Sol, 22 : 367-380.

Demange, J. M., 1981. — Les mille-pattes. Myriapodes. Boubee. Paris ; 284 pp.

Dl Castri, F., 1973. — Soil animals in latitudinal and topographical gradients of Mediterranean ecosystems. In ; F. Dl
Castri & H. A. MOONEY, Mediterranean type ecosystems. Berlin. Springer-Verlag : 171-190.

Dugrand, R., 1963. — La garrigue montpellieraine. Bull. Soc. Languedoc. Geogr.. 34 ; 3-266.

Embergkr, L., 1943. — Les limites de l'aire de vegetation mediterraneenne en France. Bull. Soc. Hist. nat. Toulouse,
78 ; 159-180.

Floret, C., Galan, M. J.. LeFloch. E.. Rapp, M. & Romane, F.. 1989. — Organisation de la structure, de la biomasse
et de la mineralomasse dun taillis ouverl de chene vert ( Quercus ilex L.). Acta Oecol.. Oecol. Plant., 10 : 245-262.

634

JEAN-FRANgOIS DAVID

Iatrou. G. D.. 1989. — Dynamics and activity of the Diplopod Glomeris balcanica in the soil subsystem of an
evergreen sclerophyllous formation in Mt. Hortiatis. Thesis, Thessaloniki : 216 pp.

Janati-Idrissi, a., 1988. — Analyse du role dcs Diplopodes cn region mediterraneenne et influence de I'impact humain
sur leurs communautds. These, University Montpellier 3 : 229 pp.

KlME, R. D., 1990a. — Spatio-temporal distribution of European millipedes. In : A. MiNELLl, Proc. 7th. int. Congr.
Myriapodology. Leiden. Brill : 367-380.

Kime, R. D.. 1990b. — A provisional atlas of European Myriapods Part 1. Fauna Europaea Evertebrata I. Luxembourg,
E.I.S.: 109 pp.

Mauries, J. P.. 1964. — Notes sur les Diplopodes pyreneens. I. Les Cylindroiulus du sous-genre Bracheoiulus Verhoeff.
Bull. Soc. Hist. tiat. Toulouse , 99 : 444-449.

MAURikS, J. P. & Geoffroy, J. J., 1982. — Decouverte dans les Causses Majeurs d'une remarquable espece cavermcole du
genre Opisthocheiron Ribaut, 1913 (Diplopoda, Craspedosomida, Opisthocheiridae). Bull. Soc. Hist. nat. Toulouse ,
118 : 131-140.

Mayr, E., 1963. — Animal species and evolution. Cambridge, Harvard University Press, 797 pp.

Merzouki, A., Lossaint. P.. Rapp. M. & Billes, G.. 1989. — L’effet d’une coupe a blanc sur la mineralisation de I'azote
dun sol rouge mediterraneen. Rev. Ecol. Biol. Sol, 26 : 133-154.

Meyer, E., Schwarzenberger, I.. Stark. G. & Wechselberger, G.. 1984. — Bestand und jahreszeitliche Dynamik der
Bodenmakrofauna in einem inneralpinen Eichenmischwald. Pedobiologia, 27 : 115-132.

Nguyen Duy - Jacquemin, M., 1973. — Contribution h la connaissance de 1'anatomie c6phalique, des formations
endocrines et du developpement post-embryonnaire de Polyxenus lagurus (Diplopodes, Penicillates). Thfcse,
Universite Paris 6 : 148 pp.

Saulnier, L. & Athias-Binche, F.. 1986. — Modalites de la cicatrisation des ecosystemes mediterraneens aprys
incendie : cas de certains Arthropodes du sol. 2. Les Myriapodes edaphiques. Vie & Milieu, Revue d'ecologie, 36 :
191-204.

Van DER Drift. J., 1975. — The significance of the millipede Glomeris marginata (Villers) for oak-litter decomposition
and an approach of its part in energy flow. In : J. Vanek, Progress in soil zoology. The Hague, Junk : 293-298.

Source : MNHN, Paris

Centipedes (Chilopoda) of Some

in Slovenia

Forest Communities

Ivan KOS

Department of Biology, Biotechnical Faculty, Askerceva 2, 61000 Ljubljana, Slovenia

ABSTRACT

Centipede communities of seven forests in Slovenia were studied using the method of the quadrat sampling and
extraction by modified Tullgren funnels. Three different forestal ecosystems were investigated in the southern part of
Slovenia. This area is characterized by a small degree of pollution. On the three localities of Abieti - Fageium Dinaricum
36 species of centipedes were found, estimated density per square meter was between 108 (± 53) and 579 (± 124). 18
species were found in Luzulo albidae - Fagetum, estimated density varies between 354 (± 115) and 408 (± 94). In the
Asperulo - Carpinetum 18 species were found, estimated density was between 136 (± 34) and 218 (± 88). In the northern
part of Slovenia, around the town Velenje, three different forestal ecosystem types in five localities were investigated.
There is a great emission of different pollutants by steam power station in this region of Slovenia. The species diversity
was lower and the density was higher than in the non-polluted localities. In the Querco - Luzulo - Fageium 23 species
were identified and the estimated density ranged from 230 (± 137) to 655 (± 859), in the Bazzanio - Abietetum 16 species
were found, the density was estimated between 195 (± 122) and 640 (± 137). In the Vaccinia myrtilli - Pinetum 12
species were found, the density was estimated between 165 (± 66) and 345 (± 345).

RESUME

Chilopodes de quelques communautes forestieres de Slovenie.

Les peuplements de chilopodes de sept forets de Slovenie ont et£ echantillonnes par la methode des quadrats suivi dune
extraction selective & I’aide d’appareils de type Tullgren. Trois <§cosystemes forestiers ont ete etudies dans le sud de la
Slovenie, caracterisee par un faible degre de pollution. Dans trois sites ^ Abieti - Fagetum Dinaricum, 36 especes de
chilopodes ont ete recolies, la densite estimee par metre carre varie de 108 (± 53) a 579 (± 124). 18 especes ont 6te
troupes dans le Luzulo albidae - Fagetum ou la densite estimee varie de 354 (± 1 15) a 408 (± 94). Dans V Asperulo -
Carpinetum , 18 especes ont ei icoltees, dont la densite est estimee entre 136 (± 34) et 218 (± 88). Dans le nord de la
Slovenie, aux environs de Velenje, trois types d’ecosystemes forestiers ont 6te (Studies dans cinq locality. Cette region
est caracterisee par une forte Emission de polluants due h I’activite d’une centrale thermique. La richesse specifique est
plus faible et la densite plus elev<Se que dans les sites non-pollues. Dans le Querco - Luzulo - Fagetum, 23 espies ont ete
identifies, la densite estimee variant de 230 (± 137) a 655 (± 859). Dans le Bazzanio - Abietetum , on a trouve 16
especes, dont la densite estimee varie de 195 (± 122) & 640 (± 137). Enfin, dans le Vaccinio myrtilli - Pinetum, 12
especes ont ete nScoltees, leur densite estimee variant de 165 (± 66) a 345 (± 345).

INTRODUCTION

Very little is known about the communities of Centipedes (Chilopoda) from Slovenia; there
are only a few records of them also from the northern part of the Balkan Peninsula. Only MATIC

Kos, I., 1996. — Centipedes (Chilopoda) of some forest communities in Slovenia. In: Geoffroy, J.-J.,
MAURlfcs, J.-P. & NGUYEN Duy - JACQUEMlN, M., (eds), Acta Myriapodologica. Mem. Mus. natn. Hist, not., 169 : 635-
646. Paris ISBN : 2-85653-502-X.

636

IVAN KOS

(1966) and MATIC & TEODOREANU (1966) reported data about vegetation in localities. More
information is available about centipedes in certain plant communities in Europe (e.g. ALBERT,
1979 1982; DUNGER era/., 1972; FRUND, 1983, 1987; LOKSA, 1968, 1979; MEYER et al.,
1984; MlNELLI & lOVANE, 1987; POSER, 1988; ZAPPAROLI, 1992; WYTWER, 1992).
Comparing the region of our research with other places in Europe, we must emphasize that it has
some specialities because of its' geographical site. It is important that here there are numerous
mixed centipede species characteristic of different biogeographical areas (KOS, 1992).
Populations of these species confirm many endemic and special communities. It is very
important to research them and their basic characteristics for a better knowledge of the role of a
single group of animals in the environment. In Slovenia, seven forest communities were studied
at 1 1 different localities. The species composition, species diversity, density, and dominance
were determined. Only the results of the method of quadrat of soil and litter sampling are
represented here.

MATERIAL AND METHODS

The sampling sites

Centipede communities were studied in two regions in Slovenia. The lirst lies south of Ljubljana and is
characterized by a small degree of pollution and well sustained forests. In the three localities near Kocevje (Rog, 900 m
a.s.I., exposure SW) and Ribnica (Kot, 700 m a.s.l., exposure E; Mala gora, 850 m a.s.l., exposure SE) the sampling was
done in the Ahieti - Fagetum Dinaricum. The soil of these sites is shallow brown with underlying deep pockets in the
limestone. On the surface, many stones are present and there are also stumps and decaying tree trunks. The fourth locality
is near to Kocevje (Zeljne, 700 m a.s.l.. flattened). Here the forest community is Lamio orvule - Fagetum. On the surface,
there are few stones and the soil is similar to the previous ones. The forest of Asperulo - Carpinetum is near Ribnica
(hamlet Seljan, 700 m a.s.l., exposure SW), the soil is shallow brown and lies on the geological joint through the
limestone. Luzulo albidae - Fagetum is near Ribnica (Zrnovec, 600 m a.s.l.. exposure SW). The soil is brown, with a
thick layer of humus, the geological background is from Permian - Cretaceous silicate slate. The second research region
was in the north of Slovenia surrounding the town Vclcnje. It is known for a great emission of different pollutants by
steam powered electrical plant. Here we researched three plant communities. In the first two, two localities were selected
which differed in the degree of pollution. Thus, in the Querco • Luzulo - Fagetum the locality Veliki vrh (480 m a.s.l.,
exposure NE) the influence of the power station is greater than in Crnova (450 m a.s.l., exposure NW). In the Bazzanio -
Abietetum, the less polluted locality Topolsica (450 a.s.l., exposure NW) was researched and Lajse (450 m a.s.l.,
exposure NW), which is more polluted. In the locality of Zavodnje Vaccinio myrtilli - Pinetum grows.

Sampling

From the forest soil a predetermined number of sampling units were taken (size 25x25x10 cm or 20x20x10 cm).
The soil samples were taken randomly on the free surface between 8 and 12 a.m from areas, which are not covered with
stones, stumps and tree trunks. Centipedes were extracted slowly on the modificated Tullgren funnels, so that the
extraction lasted 18-21 days. The specimens were extracted in ethylene glycol and later transfered to 70% ethanol for the
species determination after Attems (1929, 1930). Eason (1982), Koren (1986), Matic (1966, 1972), Verhoeff (1937)
and others. The anamorph stages of Lithobiidae could not be determined and are labelled in the tables as L. sp. juv.
Species Si. n. sp. is a new species for science, species Si. non -microps is determined alter Verhoeff (1937) as
Lithonnanus microps Mein., but is another species (Eason, KoS, in prep.). The dates ol sampling and the number ol
sampling units are listed in the tables. The estimated density was made after Elliot (1977), Seber (1982) and Krebs
(1989), but the type of the distributions was not taken into account. The 95% confidence limits of the mean was
calculated as ± USE. The density is given for a m2 of free forest surface. The Shannon - Weaver diversity index was also
calculated.

RESULTS AND DISCUSSION

Species Composition

In the study of species composition we must take samples of suitable dimensions, about
20 sampling units in size 20x20x10 cm (KOS, 1988). Otherwise the species with small densities
and the species with very aggregative distribution would not be included in the samples. It is
also important to consider that some centipede species prefer some habitats (FRUND, 1983,
1987; KOS, 1988). Because of this, species which live in tree trunks, in stumps, and under
stones are not detected in their real number when using soil sampling methods. These must be
considered in comparing different localities and communities. The largest number of species of

Source : MNHN, Paris

CENTIPEDES OF FOREST COMMUNITIES IN SLOVENIA

637

centipedes was found in Abieti - Fagetum Dinaricum, where 23 species were present in two
localities and 22 species in one. This number is much larger than is reported from other parts of
Europe (ALBERT, 1982; DUNGER et at. , 1972; FRUND, 1983; LOKSA, 1968, 1979; MEYER et
at., 1984; WYTWER, 1992). The reason for this high number could be the biogeographical
situation of these communities. An open question is the influence of the limestone substrate in
the Karst, with its peculiar formations, on the number of species in these communities. Also, the
influence of pollutants on the number of species is yet to be studied. In the Asperulo -
Carpinetum and Luzulo albidae - Fagetum, 18 species were registered. Both localities are in
southern Slovenia. Around the town Velenje, most of the species were present in Querco -
Luzulo - Fagetum (16 in one locality and 15 in another). In Vaccinio myrtilli-Pinetum we found
12 species, and the smallest number of species was found in Bazzanio - Abietetum (11 and 10
species). The results from communities mentioned are similar to those known from the middle of
Europe (ALBERT, 1982; DUNGER et at., 1972; FRUND, 1983; LOKSA, 1968, 1979; MEYER et
at., 1984; WYTWER, 1992). The Shannon - Wieners diversity index shows the same situation;
the highest is in Abieti - Fagetum Dinaricum (between 2.17 and 2.60). In other communities its
value is lower (between 1.09 and 2.08).

Density

The density of centipedes was determined on the basis of sampling the “free” surface. The
results, which present the number of animals per squere meter of “free” surface are listed in the
Tables 1-10. The number is probably underestimated because of the sampling methods and
extraction. When sampling, we took only the upper 10 cm of soil, but some specimens also live
deeper. And at slow extraction, some specimens remain in the soil (ALBERT, 1982; FRUND,
1987). In comparing our density results to those of others we must take into account that they
probably do not show the density of animals in the surface of the forest, but in the “free”
surface, which is smaller. We suppose that other authors also estimated density in this way, but
this was not always emphasized in their reports.

There is no data avaliable about the density of centipedes in Abieti - Fagetum. The
estimated number of centipedes varied between 108 (± 53) in one locality (Kot, on 8.5.1987,
Table 2) and 579 (± 124) in the locality Rog (18.4.1990, Table 1). On the basis of our results
we can conclude, that the density of centipedes in coniferous forest is smaller than in decidious
forest. It would be interesting to study the influence of litter on the density of centipedes. Taking
into account the results of some authors who also researched centipedes in different communities
(ALBERT, 1982; DUNGER et at., 1972; FRUND, 1983, 1987; WYTWER, 1992), we see that in
spruce forests, the density of centipedes in studied communities in Slovenia is much bigger than
in other parts of Europe. Further investigations of spruce forests in this region will probably
give the explanation for such big differences in density. Similar as in spruce forests, also in
beech forests of studied communities, the density of centipedes was much higher than is
reported by some authors from the other parts of Europe (ALBERT, 1982; FRUND, 1983, 1987;
LOKSA, 1968; WYTWER, 1990).

On the basis of the shown results (Tables 1-10), we can see some specific interconnections
between different populations of centipedes in individual localities. These interconnections are
related to plant communities, but the reasons for these relations are probably not only due to the
plants themselves, but to the conditions in the locality as well, such as microclimate, habitat and
biogeography, which the plant community also defines.

The results confirmed our previous conclusions about the great number of specific
communities in the area of the northern Balkans (KOS, 1992), a characteristic of its
biogeographical situation, configuration and relative well sustained environment.

The specifity is presented in species composition and in the other characteristics of
communities: density, dominance, distribution of specimens, and probably reproductive
potentials.

638

IVAN KOS

Table 1. — Estimated density of centipedes per square metre of free forest surface in Abieii - Fagelum Dinaricum in the
locality Rog (near Kocevje, Slovenia) and Mala gora (near Ribnica, Slovenia). Estimated mean and 95%
confidence limits are given. 8 sampling units (25x25x10 cm) were taken in Mala gora, and 6 sampling units
(20x20x10 cm) in Rog. (Abbreviations, see Table 2).

species

18.4.1990

avg/m2 t*se

dom

avg/m2

11.1990

t*se

dom

19.5.1987

avg/m2 t*se

Mai

dom

gora

9.9.1987
avg/m2 t*se

dom

B. morn ana

12.5

12.8

2.2

55.0

45.5

11.5

8.0

9.5

3.8

24.0

31.8

6.0

C. abbreviate

12.5

28.5

2.2

10.0

24.8

2.1

0.0

C. linearis

0.0

12.0

19.9

5.7

3.2

4.6

0.8

C. sp.

8.3

12.0

1.4

0.0

C. irebevicensis

29.2

55.9

5.0

20.0

23.2

4.2

4.0

5.7

1.9

6.4

13.7

1.6

Ch. scheerpeltzi

0.0

4.0

5.7

1.9

0.0

D. carniolensis

0.0

5.0

12.4

1.0

9.9

9.2

4.8

0.0

G. insculptus

4.2

9.5

0.7

0.0

G. oligopus cf.

0.0

15.0

24.8

3.1

5.9

6.4

2.9

0.0

G. promontorii

0.0

8.0

9.3

2.0

G. sp.

4.2

9.5

0.7

10.0

24.8

2.1

0.0

G. flavus

0.0

1.6

3.4

0.4

S. carniolensis

0.0

10.0

24.8

2.1

0.0

1.6

3.4

0.4

S. nemorensis

0.0

5.0

12.4

1.0

0.0

St. acuminata

16.7

12.0

2.9

15.0

24.8

3.1

1.9

4.4

1.0

4.8

5.3

1.2

St. transsilvanica

8.3

19.0

1.4

15.0

24.8

3.1

9.9

9.2

4.8

14.4

15.7

3.6

H. gottscheensis

4.2

9.5

0.7

0.0

L. castaneus

0.0

3.2

4.6

0.8

L. dentatus

8.3

12.0

1.4

5.0

12.4

1.0

0.0

3.2

4.6

0.8

L forficatus

0.0

L. sp. juv.

137.5

43.6

23.7

75.0

19.6

15.6

40.2

28.2

19.0

144.0

79.7

36.0

L. lapidicola

0.0

1.6

3.4

0.4

L. latro

0.0

17.6

12.9

4.4

L. macilentus cf.

66.7

54.5

11.5

65.0

57.4

13.5

35.8

20.6

17.1

108.8

81.4

27.2

L. nodulipes

4.2

9.5

0.7

0.0

3.8

4.4

1.9

0.0

L. sp. pl2-3

0.0

5.0

12.4

1.0

1.9

4.4

1.0

1.6

3.4

0.4

Si. n.sp.

8.3

19.0

1.4

0.0

24.0

1 1.4

6.4

7.5

1.6

Si. non mi crops

58.3

28.2

10.1

30.0

36.1

6.3

13.9

18.2

6.7

11.2

11.4

2.8

Si. sp.

0.0

5.0

12.4

1.0

0.0

M. aeruginosus

0.0

1.9

4.4

1.0

0.0

Cry. croaticus

0.0

5.0

12.4

1.0

0.0

Cry. hortensis

108.3

50.3

18.7

85.0

84.5

17.7

20.0

14.6

9.5

22.4

18.5

0.0

Cry. par is i

58.3

38.0

10.1

45.0

45.5

9.4

12.0

17.4

5.7

16.0

12.5

0.0

Cry. rucneri cf.

29.2

27.2

5.0

0.0

total density

579.2

123.6

100.0

480.0

245.7

100.0

197.9

82.0

100.0

400.0

146.0

100.0

total species

Sha-W. div.

2.325

2.444

2.450

2.170

Source MNHN. Paris

CENTIPEDES OF FOREST COMMUNITIES IN SLOVENIA

639

Table 2. - Estimated density of centipedes per square metre of free forest surface in Abieii - Fagetum Dinaricum in the
“f i n?63' Ribmca). Estimated mean and 95% confidence limits are given. The sampling was done on
8 5.1 987 (12 sampling units), 21.7.1987 (16 sampling units), and on 6.11.1987 (14 sampling units). The size
of a samphng unit was 25x25x10 cm. Abbreviations - B: Brachyschendyla-, C: Clinopodes ; Ch: Chaetechelyne
or Chalandea D: Dicellophilus ; G: Geophtlu.-r, S: Schendyla, St: Strigamia, E: Eupolyboihrus, H: Harpolilhobius ;
L: Lithobius ; M: Monotcirsobius\ Si: Sigibius\ Cry: Cryptops.

species

avg/m2

8.5.1987

t*se

dom

21.7.1987
avg/m2 t*se

dom

avg/m2

5.1 1.1987

t*se

dom

B. montana

4.0

4.6

3.7

5.9

8.7

2.0

17.1

9.9

7.1

C. linearis

0.0

0.00

5.0

6.7

1.7

8.0

7.9

3.3

C. trebevicensis

0.0

4.6

13.1

4.8

0.0

Ch. scheerpehzi

0.0

5.5

6.2

2.8

0.0

D. carniolensis

9.3

9.2

8.6

4.2

3.4

1.1

4.6

3.4

1.9

G. carpophagus

0.0

3.8

2.1

0.3

0.0

G. pygmaeus

0.0

333

2.1

0.3

0.0

G. flavus

1.3

2.9

1.2

2.9

6.9

3.1

1.1

2.5

0.5

S. carniolensis

0.0

2.0

2.1

0.3

3.4

5.4

1.4

St. acuminata

1.3

2.9

1.2

2.5

5.3

2.0

3.4

3.9

1.4

St. transsilvanica

2.7

4.0

2.5

1.6

5.1

1.7

8.0

6.0

3.3

E. tridentinus

1.3

2.9

1.2

1.0

2.1

0.3

0.0

H. anodus

0.0

1.0

2.1

0.3

1.1

2.5

0.5

L. agilis

0.0

1.0

2.1

0.3

0.0

0.00

L. dentatus

5.3

6.6

4.9

1.9

4.3

0.7

0.0

L. sp. juv.

30.7

26.5

28.4

1 19.0

55.9

41.1

107.4

56.7

44.3

L. lapidicola

2.7

4.0

2.5

12.0

8.5

4.1

5.8

6.8

2.4

L. latro

2.7

4.0

2.5

5.9

6.9

2.0

5.8

2.4

L. macilentus cf.

5.3

5.0

4.9

32.0

16.8

1 1.0

24.0

24.2

9.9

L. nodulipes

0.0

3.0

3.4

1.1

0.0

0.00

L. sp.

1.3

2.9

1.2

0.0

M. aeruginosus

9.3

9.2

8.6

21.9

9.8

7.6

10.2

10.6

4.3

Si. n.sp.

1.3

2.9

1.2

5.0

6.7

1.7

0.0

Si. non microps

0.0

15.0

11.4

5.2

16.0

14.5

6.6

Cry. pari si

29.3

19.8

27.1

13.9

6.1

4.8

26.2

13.4

10.9

total density

108.0

52.7

100.0

289.9

91.7

100.0

242.2

95.6

100.0

total species

Sha-W. div.

2.090

2.600

2.320

Source : MNHN, Paris

640

IVAN KOS

Table 3. — Estimated density of centipedes per square metre of free forest surface in Lamio-ovulae-Fagetum in the
locality Zeljne (near Kocevje). Estimated mean and 95% confidence limits are given. 6 sampling units
(20x20x10 cm) were taken. (Abbreviations, see Table 2).

18.4.1990

6.1 1.1990

species

avg/m2

t*se

dom

avg/m2

t*se

dom

B. montcinci

33.3

28.2

10.1

62.5

52.6

11.2

C. cibbreviatus

8.3

19.0

2.5

16.7

24.1

3.0

D. camiolensis

0.0

12.5

12.8

2.3

S. camiolensis

0.0

8.3

19.0

1.5

Si. acuminata

0.0

8.3

12.0

1.5

St. transsilvanica

4.2

9.5

1.3

16.7

28.2

3.0

H. anodus

12.5

28.5

3.8

0.0

L. agilis

0.0

4.2

9.5

0.8

L. sp. juv.

91.7

74.6

27.9

145.8

78.5

26.2

L. macilentus cf.

75.0

76.6

22.8

62.5

43.6

1 1.2

L. melanops

0.0

8.3

12.0

1.5

L. pygmaeus

25.0

20.8

7.6

0.0

M. aeruginosus

50.0

48.9

15.2

179.2

124.5

32.2

M. sp.

0.0

4.2

9.5

0.8

Si. non microps

29.2

37.3

8.9

0.0

Cry. hortensis

4.2

9.5

1.3

0.0

Cry. parisi

12.5

19.5

3.8

37.5

35.3

6.7

total density

329.2

207.5

100.0

556.7

238.3

100.0

total species

Sha-W. div.

1.970

1.470

Source : MNHN, Paris

CENTIPEDES OF FOREST COMMUNITIES IN SLOVENIA

641

Table 4. — Estimated density of centipedes per square metre of free forest surface in Asperulo - Carpinetum in the
locality Seljan (near Ribnica). Estimated mean and 95% confidence limits are given. 8 sampling units
(25x25x10 cm) were taken. (Abbreviations, see Table 2).

species

avg/m2

19.5.1987

t*se

dom

avg/m2

9.9.1987

t*se

dom

B. montana

24.0

13.4

1 1.0

8.0

7.7

5.9

C. flavidus

5.9

9.2

2.8

0.0

C. linearis

4.0

5.7

1.8

1.6

3.4

1.2

C. trebevicensis

13.9

18.6

6.4

1.6

3.4

1.2

D. carniolensis

8.0

9.5

3.7

4.8

5.3

3.5

G. flavus

45.9

40.3

21.1

33.6

22.5

24.7

S. carniolensis

1.9

4.4

0.9

0.0

Si. crassipes

1.9

4.4

0.9

1.6

*3.4

1.2

St. transsilvanica

1.9

4.4

0.9

3.2

6.9

2.4

L. sp. juv.

41.9

25.0

19.3

32.0

26.5

23.5

L. lapidicola

1.9

4.4

0.9

0.0

L. nodulipes

0.0

1.6

3.4

1.2

L. macilentus cf.

16.0

13.3

7.3

20.8

15.3

L. tricuspis

1.9

4.4

0.9

0.0

M. aeruginosas

21.9

25.8

10.1

6.4

13.7

4.7

Si. n.sp.

4.0

5.7

1.8

3.2

4.6

2.4

Si. non microps

8.0

9.5

3.7

14.4

13.9

10.6

Cry. hortensis

0.0

1.6

3.4

1.2

Cry. par is i

20.0

25.6

9.2

1.6

3.4

1.2

total density

217.9

88.2

100.0

136.0

33.6

100.0

total species

Sha-W. div.

2.085

1.737

Source : MNHN, Paris

642

IVAN KOS

Table 5. — Estimated density of centipedes per square metre of free forest surface in Luzulo albidae - Fagetum in the
locality Zrnovec (near Ribnica). Estimated mean and 95% confidence limits are given. 6 sampling units
(25x25x10 cm) were taken. (Abbreviations, see Table 2).

avg/m2

19.5.1987

t*se

dom

avg/m2

1 1.9.1987

t*se

dom

B. montana

88.0

53.1

24.9

53.9

29.8

13.2

C. trebevicensis

53.9

41.2

15.2

61.9

34.4

15.2

C/i. illyriaca

0.0

1.9

4.4

0.5

D. carniolensis

9.9

6.4

2.8

9.9

6.4

2.4

G. oligopus

1.9

4.4

0.5

0.0

St. acuminata

0.0

8.0

9.5

2.0

St. transsilvanica

1.9

4.4

0.5

0.0

L. castaneus

1.9

4.4

0.5

0.0

L. forficatus

1.9

4.4

0.5

1.9

4.4

0.5

L. sp. juv.

33.9

30.2

9.6

65.9

31.7

16.2

L. lapidicola

4.0

5.7

1.1

17.9

10.4

4.4

L. latro

1.9

4.4

0.5

0.0

L. macilentus cf.

64.0

34.2

18.1

88.0

55.1

21.6

L. melanops

1.9

4.4

0.5

0.0

L. pygmaeus

1.9

4.4

0.5

0.0

Si. non microps

24.0

23.1

6.8

12.0

8.8

2.9

M. aeruginosus

33.9

22.6

9.6

53.9

25.8

13.2

Cry. hortensis

8.0

13.4

2.3

5.9

13.2

1.4

Cry. parisi

20.0

14.6

5.6

25.9

21.1

6.3

total density

354.0

1 15.3

100.0

408.0

94.3

100.0

total species

Sha-W. div.

2.055

2.030

Source : MNHN, Paris

CENTIPEDES OP FOREST COMMUNITIES IN SLOVENIA

643

Table 6. — Estimated density of centipedes per square metre of free forest surface in Vaccinia myrtilli- Pinetum in the

™ ™ ,^aV0lJnjc (nea,r VclenJe)- Estimated mean and 95% confidence limits are given. 6 sampling units
(20x20x10 cm) were taken. (Abbreviations, see Table 2).

species

avg/m2

22.6.1990

t*se

dom

avg/m2

23.10.1990

t*se

dom

avg/m2

21.3.1991

t*se

dom

B. montana

0.0

50.0

41.5

14.5

20.0

21.9

12.1

C. abbreviatus

5.0

11.7

1.8

0.0

C. irebevicensis

140.0

1 15.2

50.9

20.0

34.2

5.8

5.0

11.7

3.0

S. nemorensis

0.0

25.0

37.1

7.2

25.0

37.1

15.2

S. carniolensis

0.0

25.0

45.4

7.2

15.0

23.5

9.1

St. acuminata

5.0

1 1.7

1.8

0.0

L. macilentus

35.0

35.2

12.7

85.0

133.2

24.6

65.0

54.4

39.4

L. latro

0.0

20.0

1 1.7

5.8

5.0

1 1.7

3.0

L. tenebrosus

15.0

23.5

5.5

15.0

35.2

4.3

0.0

L. sp. juv.

20.0

21.9

7.3

7570

132.5

21.7

10.0

23.5

6.1

M. aeruginosus

25.0

18.5

9.1

20.0

34.2

5.8

10.0

23.5

6.1

Cry. hortensis

30.0

34.2

10.9

10.0

14.4

2.9

10.0

14.4

6.1

Cry. parisi

0.0

total density

275.0

143.64

100.0

345.0

344.7

100.0

165.0

65.8

100.0

total species

Sha-W. div.

1.402

1.978

1.613

Table 7. — Estimated density of centipedes per square metre of free forest surface in Bazzanio - Abietetum in the locality
Topolsica (near Velenje). Estimated mean and 95% confidence limits are given. 6 sampling units (20x20x10 cm)
were taken. (Abbreviations, see Table 2).

species

avg/m2

22.6.1990

t*se

dom

avg/m2

23.10.1990

t*se

dom

avg/m2

21.3.1991

t*se

dom

B. montana

0.0

5.0

1 1.7

1.6

15.0

23.5

7.7

G. insculptus

43.8

78.6

11.3

75.0

41.5

23.4

10.0

14.4

5.1

G. oligopus cf.

12.5

25.4

3.2

0.0

S. nemorensis

0.0

25.0

45.4

7.8

5.0

1 1.7

2.6

S. carniolensis

0.0

5.0

1 1.7

1.6

25.0

18.5

12.8

L. forficatus

0.0

L. sp. juv.

18.8

38.1

4.8

30.0

43.1

9.4

5.0

1 1.7

2.6

L. macilentus

18.8

24.3

4.8

25.0

32.1

7.8

20.0

46.9

10.3

M. aeruginosus

50.0

68.8

12.9

15.0

35.2

4.7

30.0

43.1

15.4

Cry. hortensis

137.5

44.0

35.5

140.0

73.2

43.8

85.0

75.5

43.6

Cry. parisi

0.2

12.7

1.6

0.0

total density

387.5

86.7

100.0

320.0

179.2

100.0

195.0

122.2

100.0

total species

Sha-W. div.

1.367

1.417

1 .606

Source :

644

IVAN KOS

Table 8. — Estimated density of centipedes per square metre of free forest surface in Bazzanio - Abietetum in the locality
Lajse (near Velenje). Estimated mean and 95% confidence limits are given. 6 sampling units (20x20x10 cm) were
taken. (Abbreviations, see Table 2).

22.6.1990

10.1990

17.6.1991

species

avg/m2

t*se

dom

avg/m2

t*se

dom

avg/m2

t*se

dom

C. flavidus

5.0

11.7

1.1

0.0

C. trebevicensis

75.0

61.5

15.8

50.0

26.2

7.8

95.0

78.4

17.6

D. carniolensis

0.0

5.0

1 1.7

0.8

0.0

S. carniolensis

0.0

75.0

37.1

11.7

0.0

S. nemorensis

0.0

60.0

68.4

9.4

0.0

St. acuminata

5.0

11.7

1.1

0.0

L. den tat us

5.0

1 1.7

1.1

0.0

L. sp. juv.

90.0

93.8

18.9

130.0

77.4

20.3

60.0

57.5

11.1

L. macilentus

70.0

65.3

14.7

105.0

98.8

16.4

65.0

104.9

12.0

M. aeruginosus

130.0

70.4

27.4

80.0

65.3

12.5

200.0

193.0

37.0

Cry . hortensis

85.0

77.8

17.9

120.0

56.9

18.8

1 10.0

92.9

20.4

Cry. parisi

10.0

14.4

2.1

15.0

14.4

2.3

10.0

30.8

1.8

total density

475.0

252.2

100.0

640.0

137.0

100.0

540.0

160.6

100.0

total species

Sha-W. div.

1.223

1.226

1.867

Table 9. — Estimated density of centipedes per square metre of free forest surface in Querco - Luzulo - Fagetum in the
locality Veliki vrh (near Velenje). Estimated mean and 95% confidence limits are given. 6 sampling units
(20x20x10 cm) were taken. (Abbreviations, see Table .2).

22.6.1990

10.1990

21.3.1991

17.6.1991

species

avg/m2

t*se

dom

avg/m2

t*se

dom

avg/m2

t*se

dom

avg/m2

t*se

dom

B. montana

0.0

10.0

14.4

2.9

10.0

14.4

3.9

0.0

C. abbreviatus

0.0

25.0

58.6

7.1

10.0

14.4

3.9

0.0

C. flavidus

5.0

1 1.7

2.2

0.0

C. linearis

0.0

5.0

1 1.7

1.4

5.0

11.7

2.0

5.0

11.7

1.0

C. trebevicensis

10.0

14.4

4.3

10.0

14.4

2.9

5.0

1 1.7

2.0

35.0

35.2

7.1

D. carniolensis

5.0

11.7

2.2

5.0

1 1.7

1.4

0.0

G. oligopus cf.

10.0

23.5

4.3

5.0

1 1.7

1.4

20.0

21.9

7.8

15.0

23.5

3.0

St. transsilvanica

0.0

5.0

1 1.7

1.4

0.0

E. tridentinus

0.0

5.0

1 1.7

1.4

0.0

L. dentatus

0.0

5.0

1 1.7

1.4

10.0

23.5

3.9

0.0

L. forficalus

0.0

5.0

1 1.7

1.4

0.0

L. sp. juv.

25.0

45.4

10.9

85.0

104.2

24.3

85.0

122.4

33.3

220.0

261.2

44.4

L. lapidicola

0.0

5.0

1 1.7

1.4

0.0

L. lusitanus

0.0

5.0

1 1.7

1.0

L. macilentus

165.0

1 13.7

71.7

160.0

1 18.2

45.7

100.0

92.7

39.2

195.0

1 13.4

39.4

M. aeruginosus

5.0

11.7

2.2

0.0

Cry. parisi

5.0

1 1.7

2.2

20.0

21.9

5.7

10.0

14.4

3.9

20.0

21.9

4.0

total density

230.0

136.8

100.0

350.0

194.5

100.0

255.0

241.4

100.0

495.0

344.1

100.0

total species

Sha-W. div.

0.831

1.569

1.348

1.000

Source : MNHN, Paris

CENTIPEDES OF FOREST COMMUNITIES IN SLOVENIA

645

'Fable 10. — Estimated density of centipedes per square metre of free forest surface in Querco - Luzulo - Fagetum in the
locality Crnova (near Velenje). Estimated mean and 95% confidence limits are given. 6 sampling units
(20x20x10 cm) were taken. (Abbreviations, see Table 2).

22.6.1990

21.3.1990

.3.199

7.6.1991

species

avg/m2

t*se

dom

avg/m2

t*se

dom

avg/m2

t*se

dom

avg/m2

t*se

dom

B. m on tana

0.0

40.0

52.4

14.0

25.0

32.8

3.8

0.0

C. abbreviate

0.0

5.0

6.6

1.8

5.0

6.6

0.8

0.0

C. trebevicensis

15.0

19.7

4.3

10.0

13.1

3.5

0.0

D. carniolensis

0.0

5.0

6.6

1.8

5.0

6.6

0.8

5.0

6.6

1.1

G. insculptus

75.0

98.3

21.7

30.0

39.3

10.5

30.0

39.3

4.6

205.0

268.8

43.6

G. flavus

0.0

5.0

6.6

1.1

S. nemo re ns is

0.0

10.0

13.1

1.5

0.0

St. acuminata

5.0

6.6

1.4

0.0

10.0

13.1

1.5

5.0

6.6

1.1

L. dentatus

0.0

L. forficatus

5.0

6.6

1.4

0.0

L. lusitanus

5.0

6.6

1.4

0.0

o.o-

0.0

10.0

13.1

1.5

20.0

26.2

4.3

L. macilentus

100.0

131.1

29.0

55.0

72.1

19.3

150.0

196.7

22.9

1 10.0

144.2

23.4

L. nodulipes

0.0

25.0

32.8

3.8

0.0

L. valid us

10.0

13.1

2.9

5.0

6.6

1.8

0.0

10.0

13.1

2.1

L. sp. juv.

70.0

91.8

20.3

45.0

59.0

15.8

265.0

347.5

40.5

65.0

85.2

13.8

L. sp.

5.0

6.6

1.4

0.0

10.0

13.1

1.5

0.0

H. sp.

0.0

5.0

6.6

0.8

0.0

Cry. hortensis

55.0

72.1

15.9

90.0

118.0

31.6

105.0

137.7

16.0

45.0

59.0

9.6

total density

345.0

452.4

00.0

285.0

373.7

100.0

655.0

858.9

100.0

470.0

616.3

100.0

total species

Sha-W. div.

1.615

1.638

1.864

1.345

REFERENCES

Albert, A. M., 1979. — Chilopoda as part of the predatory macroarthropod Fauna in forests: Abundance, Life-cycle,
Biomass, and Metabolism. In: M. Camatini, Myriapod Biology. London, Academic Press : 215-231.

Albert, A. M., 1982. — Species spectrum and dispersion patterns of Chilopods in Soiling habitats. Pedobiologia. 23 :
337-347.

Attems, C., 1929. — Myriapoda. I. Geophilomorpha. Das Tierreich. 52 : 1-388.

Attems, C., 1930. — Myriapoda II. Scolopendromorpha. Das Tierreich. 54 : 1-308.

Dunger. W., Engelmann, I. & Schneider R.. 1972. — Untersuchungen zur Langzeitwirkung von Industrie - Emissionen
auf Boden, Vegetation und Bodenfauna des Ncitzetales bei Ostitz/Oberlausitz. Abh. Bee Natur. Gorlitz . 47 : 1-40.
Eason, E. H., 1982. — A review of the North-West Lithobiomorpha with a revised key to their identification. Zool. J.
Linn. Soc ., 74 : 9-33.

Elliot, J. M., 1977. — Some methods for the Statistical Analysis of samples of Benthic Invertebrates. Freshwater
Biological Association Sci. Publication . 25 : 1-157.

Frund, H. C., 1983. — Untersuchungen zur Koexistenz verschiedener Chilopodenarten im Waldboden. Dissertation,
Wiirtzburg, 164 pp.

Frond, H. C., 1987. — Raumliche Verteilung und Koexistenz der Chilopoden in einem Buchen-Altbestand.
Pedobiologia. 30 : 19-29.

Koren, A., 1986. — Die Chilopoden-Fauna von Karnten und Osttirol, Teil 1, Geophilomorpha, Scolopendromorpha. -
Verlag des Naturwiss. Vereins fur Karnten, Klagenfurt. 87 pp.

Kos, I., 1988. — The problems of quality and quantity sampling of centipedes (Chilopoda). Ljubljana, University of
Ljubljana, Biotech. Faculty, Department of Biology, 85 pp. (in Slovene).

Source

646

IVAN KOS

Kos, I., 1992. — A Review of Taxonomy, Geographical Distribution and Ecology of the Centipedes of Yugoslavia
(Myriapoda, Chilopoda). ( In: E. Meyer, K. Thaler, W. Schedl , Advances in Myriapodology.] Ber. nat.-med. Verein
Innsbruck , suppl. 10 : 353-360.

Krebs, J. R., 1989. — Ecological methodology. New-York, Harper Collins Publishers, 654 pp.

Loksa, I.. 1968. — Quantitative Makrofauna-Untersuchungen in den Waldboden des Bukkgebirges (Ungarn). Ann. Univ.
Sci. Budapest, 9-10 : 265-289.

Loksa, I., 1979. — Quantitative Untersuchungen iiber die Makrofauna der Laubstreu in Zerreichen- und Hainsimen-
Eichen Bestanden des Biikkergebirges. Opusc. Zool. Budapest . 16: 87-96.

Matic, Z., 1966. — Clasa Chilopoda, Subclasa Anamorpha. Fauna Republicii socialiste Romania , 6 : 1-272.

MATIC, Z., 1972. — Clasa Chilopoda , Subclasa Epimorpha. Fauna Republicii socialiste Romania , 6 : 1-224.

Matic. Z. & Teodoreanu, M., 1966. — Contribution & la connaissance des Lithobiides (Chilopoda-Lithobiidae) de
Croatie. Biol. Glasnik , 19 : 19-26.

Meyer, E., Schwarzenberger. I.. Stark. G. & Wechselberger, G.. 1984. — Bestand und jahreszeitliche Dynamik der
Bodenmakrofauna in einem inneralpinen Eichenmischwald (Tirol, Osterreich). Pedobiologia, 27 : 115-132.

MINELLI, A. & Iovane, E., 1987. — Preferences and taxocenoses of Italian centipedes (Chilopoda). Boll. Mus. civ. St.
nat. Venezia . 37 : 7-34.

POSER, T., 1988. — Chilopoden als Pradatoren in einem Laubwald. Pedobiologia, 31 : 261-281.

SEBER. G. A. F., 1982. — The Estimation of Animal Abundance and related Parameters. London, Charles Griffin &
Company Ltd.. 654 pp.

VERHOEFF, K. W., 1937. — Chilopoden-Studien. Zur Kenntnis der Lithobiiden. Archiv fur Naturgeschichte, 6 : 171-257.

Wytwer. J., 1990. — Centipedes (Chilopoda) of linden - oak - hornbeam forests ( Tilio - Carpinetum) and the
thermophilous oak forests ( Potentillo albae - Quercetum) of Mazovian Lowland. Fragm. Faun., 32 :73-94.

Wytwer, J.. 1992. — Chilopoda Communities of the Fresh Pine Forests of Poland. [In : E. Meyer, K. Thaler, W.
Schedl, A dvances in Myriapodology.] Ber. nat.-med. Verein Innsbruck, suppl. 10: 205-211.

Zapparoli, M., 1992. — Preliminary Data on Centipede Communities of Quercetea ilicis and Fagetalia sylvaticae in
Central Italy. [In: E. MEYER, K. Thaler. W. Schedl. Advances in Myriapodology] Ber. nat.-med. Verein Innsbruck,
suppl. 10 : 197-204.

Source : MNHN. Paris

Changes in the Millipede (Diplopoda) Community
during Secondary Succession from a Wheat Field to a

Beechwood on Limestone

Stefan SCHEU

II. Zoologisches Institut, Abteilung Okologie, Berliner Str. 28. 37073 Gottingen. Germany

ABSTRACT

The diplopod communities of five sites, which were chosen to represent different stages af secondary succession from
a wheat field to a beechwood on limestone (wheat field, 4 year old fallow, 1 1 year old fallow, ca 50 year old fallow,
beechwood), were studied for 2 years. The sites were located in close proximity on a limestone plateau east of Gottingen
(southern Lower Saxonia, Germany). Diplopods were extracted by heat from soil cores four times a year. A total of 12
species were found: Allajulus nitidus , Cytindroiulus caeruleocincius, Lepioiulus belgicus, Unciger foeiidus,
lachypodoiulus mger. Ommatoiulus sabulosus, Blaniulus gullulalus , Mycogona germanicum, Glomeris marginaia.
Ulomens conspersa Stygioglomeris criniia , Polydesmus inconstant. Number of species, density and biomass increased
up to the 11 year old tallow stage. They were considerably lower in the 50 year old fallow (ash dominated wood) and
seemed to have increased again until the formation of the climax ecosystem of the beechwood. In contrast to succession
theory no continuous change in species composition occurred; rather, diplopods nourished at intermediate stages of
secondary succession. Changes in community structure are discussed and related to environmental factors. Canonical
correspondence analysts indicated the great importance of the amount of litter and of humidity for the species
composition of diplopods. The most important determining factor for low diversity, density and biomass of millipedes
at the 50 year old tallow is assumed to be the absence of a litter layer during summer.

RESUME

Variations dans un peuplement de diplopodes au cours d’une succession secondaire, d'un champ
de ble a une hetraie sur calcaire.

Les peuplements de diplopodes ont 6te etudies pendant deux ans dans cinq sites voisins, representatifs des different s
stades d une succession allant d’un champ de ble a une hetraie sur calcaire (champ de ble, jachere de 4 ans, jachere de 1 1
ans, Iriche de 50 ans, foret de hetres). Les stations sont siluees sur un plateau calcaire a Test de Gottingen (Allemaene).
Les diplopodes ont 616 obtenus par echantillonage de carottes de sol et extraction selective 4 fois par an. Douze especes
ont ete fee ol tees • Allajulus nitidus , Cylindroiulus caeruleocincius, Lepioiulus belgicus , Unciger foeiidus ,
Tachypodoiulus niger, Ommatoiulus sabulosus, Blaniulus guttulatus, Mycogona germanicum, Glomeris marginaia ,
Glomeris conspersa , Stygioglomeris criniia, Polydesmus inconstans. La richesse specifique. la densite et la biomasse
augmentent jusqu’au stade jachere de 1 1 ans. Elies sont nettement plus faibles dans la friche de 50 ans (Prene dominant) et
semblent de nouveau augmenter jusqu’a la formation forestfere “climacique” de la hetraie. A rencontre de la theorie des
successions, aucun changement continu n'apparait dans la composition specifique ; on observe plutot un developpement
florissant du peuplement de diplopodes durant les stades intermediates de la succession. Les variations de cette structure
sont discutees et mises en relation avec les facteurs environnementaux. Une analyse de correspondances canonique
montre la grande importance de 1 apport de litiere et de 1‘humidite dans la composition specifique des diplopodes. Le

Scheu, S., 1996. Changes in the millipede (Diplopoda) community during secondary succession from a wheat
held to a beechwood on limestone. In. Geoffroy, J.-J., Mauris, J.-P. & Nguyen Duy - Jacquemin. M., (eds), Acta
Mynapodologica. Mem. Mus . natn. Hist, nai., 169 : 647-656. Paris ISBN : 2-85653-502-X.

648

STEFAN SCHEU

facteur determinant les faibles diversity densite et biomasse de diplopodes dans la friche de 50 ans semble etre 1'absence
de liti&re durant V6l6.

INTRODUCTION

Diplopod communities of forest ecosystems (DUNGER, 1958; BLOWER, 1979;
GEOFFROY, 1981; MEYER et al., 1984; AXELSSON et al., 1984) and arable fields (HERBKE,
1962; PETERS, 1984; KLINGER, 1992) have been frequently investigated. In addition, diplopods
on restored mining soils found considerable attention (DUNGER, 1968; NEUMANN, 1971;
DUNGER. & VOIGTLANDER, 1990). In contrast, little is known about changes in diplopod
communities during secondary succession, e.g. after cessation of cultivation (TAJOVSKY,
1990).

Studies on secondary succession up to the climax are, for practical reasons, usually
performed at different sites representing different stages of secondary succession. In the present
study five sites were chosen to investigate changes in species composition, density and biomass
of diplopods during secondary succession from an arable field to the climax ecosystem of a
beechwood ( Fagus sylvatica L.). The sites were located on a limestone plateau east of Gottingen
(southern Lower Saxonia. Germany), most of them in close vicinity. The first site was an arable
field which had been planted with wheat. The second and third sites had been left uncultivated
for 4 and 1 1 years, respectively. The fourth site had been left abandened for ca 50 years and a
tree layer mainly of ash ( Fraxinus excelsior L.) had grown up at this site. The fifth site was a
beechwood which has been studied intensively (cf. SCHAEFER, 1991).

The present study forms part of a project which investigated changes in density and
biomass of soil animals during secondary succession in combination with functional aspects of
soil invertebrates for carbon turnover and nutrient cycling (SCHEU 1990a, b, 1992; SCHULZ,
1992; WOLTERS, in prep.). The aim of the present study was to get a closer understanding of
the factors responsible for changes in the millipede community during secondary succession.

MATERIALS AND METHODS

The sites

Five sites representing different successional stages from a wheat field to a beechwood were investigated. The
sites were located on a limestone plateau (360-420 m) east of Gottingen (Lower Saxonia, Germany). The wheat field was
planted with wheat during the investigations. In general, a rotation of wheat, barley and rape has been planted on that
field during the previous 20 years. Straw residues were burnt in autumn and the field worked with a disk cultivator. The
second site (first fallow) had been left uncultivated for 4 years until 1987. The flora consisted of a mixture of weeds and
grasses. The third site (second fallow) had been left uncultivated for 11 years and grasses dominated at this site. The
floral composition indicated a decrease in nitrogen supply until this stage (cf. SCHEU 1990b). The fourth site (third
fallow) had been left uncultivated for ca 50 years and was now an ash (F. excelsior ) dominated wood. Ash trees had
overgrown shrubs which were still present but in a stage of die back. The fifth site was a ca 130 year old beechwood
( F . sylvatica) which has been described in more detail in Schaefer (1991). More details on the other sites can be found
in SCHEU (1990a, 1992).

The climate in the study area is characterized by mild winters and humid summers. The annual mean temperature in
Gottingen is 8.7°C and the annual mean precipitation 613 mm. Variation among years is considerable and there might be
longer frost periods in winter and dry periods in summer; the former was the case in February 1987, whereas the latter
occurred in May and to a lesser extent also in August 1988 (Fig. 1).

The soil water content varied during the year with a maximum in late winter and spring (Fig. 2). Variation was
most pronounced in the litter layer and similar in the 0-3 and 3-6 cm soil depths. Generally, the water content of the soil
was similar in the arable field and the fallow sites and considerably lower than in the beechwood.

The amount of carbon and nitrogen (determined by an elemental analyser; Carlo Erba, Milano, Italy) in 0-3 and in
3-6 cm soil depth was similar in the arable field and the two younger fallow sites and considerably lower than in the ash
dominated wood and the beechwood (Fig. 3). The increase in the amounts of carbon and nitrogen in the two woodlands
was caused by an accumulation of humus in the upper soil layers of these ecosystems which was accompanied by a
decrease in soil bulk density. Mean annual amount of carbon and nitrogen in the litter layer was at a maximum in the
fallow sites but varied considerably during the year. Litter material al the first fallow consisted mainly of weed residues,
whereas grass leaf litter dominated at the second fallow. Litter materials estimated are presumably somewhat too high,
particularly for the 4 year old fallow, because the above ground vegetation was partly included in material taken as litter.

Source : MNHN, Paris

THE MILLIPEDE COMMUNITY OF A SECONDARY SUCCESSION

649

In the ash wood most of the litter consisted of small twigs and branches from overgrown shrubs, the leaf litter had
disappeared almost entirely by spring. In the beechwood a litter layer of beech leaves was present throughout the year

Faunal investigations

r S°tTeS ofj0 036 m2 were laken from lhe sludY sites and separated into litter layer, 0-3 and 3-6 cm soil depth in
the held. Eight randomly distributed samples were taken at 3 month intervals from October, 1986 to October 1988
except for the third tallow which was investigated from October, 1987 to October, 1988 only. Diplopods were extracted
by neat using a modified Kempson extractor (Kempson et al. , 1963. Schauermann, 1982) and determined to species
level. The biomass was estimated using regressions between the body diameter and ash free dry mass established by
Sprenobl (1986) for Mycogona germanicum (Verhoeff, 1897) and Allajulus nitidus (Verhoeff, 1891). Biomass of
Cylindroiulus caeruleocinctus (Wood, 1864), Leptoiulus belgicus (Latzel, 1844), Unciger foetidus (C. L. Koch 1838)
Tachypodoiulus niger (Leach, 1815). Ommatoiulus sabulosus (Linne, 1758), Blaniulus guttulatus (Fabricius, 1798) and
i7Qo d\5mA% inconslans Lalzel’ 1884 was estimated using the formula for A. nitidus. For Glomeris marginata (Villers,
1/89) and G conspersa (C. L. Koch, 1847) regressions between the width of the collum and ash free dry mass were used

(^prengel 1986). The regression for G. marginata was also used to calculate the biomass of S^gioglomeris crinita
broleman, 1913.

Statistical analysis

Canonical correspondence analysis (CCA, Ter Braak, 1988) was used to ordinate samples and to relate diplopod
data to environmental factors. Environmental factors (water content, carbon content, nitrogen content, C/N ratio,
amount of carbon, amount of nitrogen) were determined from the litter material, 0-3 and 3-6 cm soil depth of small soil
cores taken in close vicinity to the core for faunal extractions. Only four cores were taken at each sampling date at each
site for determination of environmental factors and therefore, only diplopods from the adjacent four cores were ordinated.
rhe water content in 3-6 cm soil depth was excluded from the analysis because of the collinearity with the water content
in 0-3 cm (ct. Fig. 2). The amount and content of carbon and nitrogen in 0-3 and 3-6 cm soil depth was excluded because
the correlation with species axes was poor (r < 0.3). Eigenvalues obtained by CCA ordination were compared with those
ot DCA (detrended correspondence analysis; cf. Ter Braak, 1988) ordination to estimate the relevance of the
environmental factors included in the analysis.

-10

1986 1987 1988

Fig. 1. — Monthly mean temperature and precipitation at the study sites in 1986-1988 (data were kindly provided by the

Institute for Soil Science, Gottingen).

RESULTS

Species composition

A total of 12 diplopod species were found. Six species of Julidae (A. nitidus ,
C. caeruleocinctus, L. belgicus, U. foetidus, T. niger, O. sabulosus) and three Glomeridae
(G. marginata , G. conspersa, S . crinita). The other three species belonged to three different

650

STEFAN SCHEU

families: B. guttulatus (Blaniulidae). M. germanicum (Chordeumatidae), P. inconstans
(Polydesmida e).

Wheat field ■ 4 year old fallow

► 1 1 year old fallow • ca 50 year old fallow
* Beechwood

□ Litter CD 0-3 cm £3 3-6 cm

Fig. 2. - Changes in gravimetric soil water content in
the litter layer (a>, 0-3 (b) and 3-6 cm soil depth
vc) at five sites representing different stages of
secondary succession during 1987 and i 988.

FlG. 3. — Amount of carbon (a) and nitrogen (b) in the
litter layer, 0-3 and 3-6 cm soil depth of the
study sites (means + SD of samples taken at 3
month intervals during 1987 and 1988; Field,
wheat field; Fal 1, 4 year old fallow; Fal 2, 11
year old fallow; Fal 3, ca 50 year old fallow; Bw,
Beechwood).

Species composition at the study sites was very different (Fig. 4). At the arable field five
species were found. However, A. nitidus and S. crinita were found only once (adult females in
both cases) and C. caeruleocinctus, in very small numbers (adult male, adult female and a
specimen of stage 3). It is doubtful therefore, if these species formed autochthonous populations
on the wheat field. Two species (P. inconstans and B. guttulatus) were found frequently and
different stages were present.

At the 4 year old fallow a total of 10 species was found (Fig. 4). Except for U. foetidus
juveniles were found, indicating that almost all of these species had reproduced on that site. At
the 1 1 year old fallow 1 1 species were present. From each of these species juveniles were found

Source . MNHN , Paris

THE MILLIPEDE COMMUNITY OF A SECONDARY SUCCESSION

651

indicating that these species formed autochthonous populations. In comparison to the 4 year old
lal ow a considerably higher ratio of specimens older than 5 years was present at the 1 1 year old
tallow, particularly for A. nitidus and C. caeruleocinctus. At the ca 50 year old fallow only five
species were present. Only one specimen of C. londinensis in stage 6 was found and it remains
unclear if this species was able to reproduce on this site. In the beechwood six species were
present adults and juveniles of each of them were found. Despite the similar number of species
in die beechwood and the ca 50 year old fallow, the species composition was very different

G. conspersa
G. marginata
U. foetidus
T. niger

O. sabu/osus
M. germanicum
L. be/gicus

C. nitidus
C. caenj/eocinctus
S. crinata

P. inconst ans
B. guttu/atus

Diplopoda

V 50 - \A

1 5 10 15 20 V 50 V 100 115 120

Years

Fig. 4. Changes in species composition ot diplopods during secondary succession from a wheat field to a beechwood
on limestone as indicated from the presence of diplopod species at five sites representing different stages of
secondary succession.

Density and biomass

Mean density and biomass of diplopods at the arable field were almost identical in 1987
and 1988 (Table 1). Ot the two dominating species, B. guttulatus contributed more to mean
annual density and biomass than P. inconstans (Fig. 5). Generally, density and biomass of
millipedes were high in spring and autumn and few animals were found in winter and summer.

Density and particularly biomass of diplopods at the 4 year old fallow was considerably
higher than at the arable field (Fig. 6). Biomass at the 4 year old fallow peaked in autumn and
only few specimens were found in winter (Table 1). A. nitidus dominated in density and
biomass (Fig. 5) indicating that this species was the most successful colonizer during early
secondary succession. Two other julid species (C. caeruleocinctus , L. belgicus ) also contributed
significantly to density and biomass at this site.

At the 1 1 year old fallow density and biomass of diplopods were generally similar to that at
the 4 year old fallow, however, mean annual density at the 4 year old fallow exceeded that at the
1 1 year old tallow in 1987 whereas mean annual biomass at the 1 1 year old fallow exceeded
considerably that at the 4 year old fallow in 1988 (Table 1). Similar to the 4 year old fallow. A.
nitidus dominated in density at the 1 1 year old fallow (Fig. 5). A. nitidus and C. caeruleocinctus
also contributed significantly to biomass in the latter, but the larger julid species O. sabulosus
dominated, despite its low numbers. As in the 4 year old fallow biomass of the millipedes
usually increased during the year and peaked in autumn.

652

STEFAN SCHEU

Table 1. — Changes in density and biomass of diplopods at five sites representing different stages of a secondary
succession from October 1986 to October 1988. ND = not determined.

Density (ind/m2)

Mean

10/86

1/87

4/87

7/87

10/87

1/88

4/88

7/88

10/88

1987

1988

Field

112

1 1

First fallow

137

343

126

179

156

102

Second fallow

270

165

130

182

102

109

Third fallow

242

BeechWood

147

116

Biomass (mg dry mass/m2)

Mean

Field

127

172

105

108

First fallow

1541

612

459

1455

976

320

1175

651

641

Second fallow

1102

450

606

294

744

1665

947

414

1956

524

1246

Third fallow

936

387

132

BeechWood

567

511

175

1560

109

108

557

434

584

302

Density and biomass of millipedes at the ca 50 year old fallow were exceptionally low and
similar to the field (Fig. 6). A. nitidus dominated in density and biomass. L. belgicus and B.
guttulatus had a similar density but L. belgicus contributed more to biomass.

In the beechwood the density and biomass of millipedes seems to have increased
considerably in comparison to the preceeding successional stage of the ca 50 year old fallow
(Fig. 6). However, in comparison to the two other fallow sites density and biomas of diplopods
were considerably lower in the beechwood. In the 4, 1 1 and ca 50 year old fallow sites A.
nitidus dominated in density and biomass as in the beechwood, but two other species, M.
germanicum and G. marginata, which were absent or rare at preceeding successional stages
contributed significantly to diplopod density and biomass in the beechwood.

Species environment correlation

The biplot of diplopod species composition and environmental variables (Fig. 7a) showed
a gradient from species which dominated in the wheat field (P. inconstans, B. guttulatus) to
species which occurred mainly in the beechwood (G. marginata, G. conspersa, U. foetidus , M.
germanicum). Species which were most abundant at the two younger fallow sites (C.
caeruleocinctus, O. sabulosus) were separated from those which dominated in the beechwood by
a moisture gradient and by a gradient in the amount of litter. As shown in Figure 3, the two
fallow sites had the highest amounts of litter, whereas the beechwood was the most humid site
(Fig. 2). The small difference in CCA eigenvalues (first axis 0.58, second axis 0.35) and DCA
eigenvalues (first axis 0.74, second axis 0.46) indicate that a substantial part of the variation in
species composition is represented by the environmental variables.

The CCA joint plot of samples showed four clusters (Fig. 7b). Parallel to the positions of
B. guttulatus and P. inconstans (Fig. 7a), the dominating species at the wheat field, samples
taken at this site were on the far right side of the diagram (first cluster). Positions of samples
from the beechwood (second cluster) and the 4 and 1 1 year old fallow (third cluster)
corresponded to the positions of the dominating species at these sites. Positions of samples from
the third fallow (fourth cluster) indicate that there was no continuous change in environmental
conditions during secondary succession in respect to the diplopod community. Environmental
factors representing much of the dipopod variation at this site were similar to those at the arable
field. Hence, in respect to the diplopod community, the arable field was most similar to the third
fallow, contrasting the floral dissimilarity of the two sites.

Source : MNHN, Paris

THE MILLIPEDE COMMUNITY OF A SECONDARY SUCCESSION

653

~Z! Cy/indroiu/us nitidus

Cylindroiulus caeru/eocinctus
Leptoiu/us be/gicus
(ZD Jnciger foetidus
i~i Tachypodoiu/us niger
IB Ommatoiulus sabu/osus

B/aniu/us guttulatus
Mycogona germanicum
iv?3 G/omeris conspersa
V72 G/omeris marginata
ES Stygiog/omeris crinata
CSJ Po/ydesmus inconstans

Fig. 5. — Dominance structure in density (a) and
biomass (b) of diplopods at five sites
representing different stages of secondary
succession (legend see Fig. 3).

Fig. 6. — Density (a) and biomass (b) of diplopods at
five sites representing different stages of
secondary succession (means of 1987 and 1988
except for the third fallow which was
investigated in 1988 only; legend see Fig. 3).

DISCUSSION

Succession refers to a continuous change in species composition, with time, following a
peiturbation. To explain the changes in species composition three hypotheses were presented
(CONNELL & Slatyer, 1977): facilitation, tolerance and inhibition. The facilitation hypothesis
assumes that species of early successional stages improve environmental conditions for
following species (ODUM, 1969). The tolerance model assumes that species of later successional
stages are more specialized; the changes in species composition being independent of the
preceeding species. The inhibition model assumes that species which have colonized a distinct
successional stage inhibit colonization by other species until their death. Most of these theories
were developed in studies on plant communities or sessile marine animals, little attention has
been paid to terrestrial animal communities.

The floral composition of the five sites investigated in the present study indicates a
continuous change in plant species composition during secondary succession from the arable
field to the beechwood (SCHEU, 1990b). In contrast to plants, the diplopod communities at the
live sites indicate that no continuous change in millipede species occurs when arable fields are
left uncultivated until formation of the climax ecosystem of a beechwood. Instead, diplopod
species flourished at the 1 1 year old fallow and several species which occurred at this site were
absent at the ca 50 year old fallow (S. crinata , M. germanicum , U. foetidus, G. marginata) but
occurred again in the beechwood. Hence, the data indicate that several diplopod species become

654

STEFAN SCHEU

extinct during secondary succession in an ash dominated stage and occur again in the climax
ecosystem of the beechwood. TAJOVSKY (1990) also found diplopods to thrive in early stages
of secondary succession. For a better understanding of changes in the diplopod community
during secondary succession gradients in environmental factors should be considered.

The most extreme habitat for millipedes was the field. Cultivation presumably caused an
extreme disturbance of the habitat and straw burning reduced the available food substrate. As a
consequence, only species with a short life cycle (e.g. P. inconstans) or with a high reproductive

potential (r-strategists; e.g. P. inconstans, B.
guttulatus) occurred at this site. Cessation of
cultivation enabled a variety of diplopod
species to colonize the abandoned fields. At
the 4 year old fallow at least nine species
formed autochthonous populations; at the 1 1
year old fallow 1 1 reproducing species were
present.

One of the most important factors
responsible for the thriving of diplopods at the
4 and 1 1 year old fallow sites was presumably
the presence of a litter layer throughout the
year. Litter material may have served as food
substrate, but the presence of a litter layer
might have also buffered temperature and
moisture extremes, e.g. frost during winter
and drought in summer. As indicated by CCA
ordination, the occurrence of species typical of
the 4 and 11 year old fallow (e.g. C.
caeruleocinctus, O. sabulosus) was related to
high amounts of litter at these sites. A second
factor which might have been important in
structuring the diplopod communities at these
sites was humidity. CCA showed that the
occurrence of millipede species typical of the 4
and 1 1 year old fallow was related to low
moisture levels in litter and soil. Obviously,
these species tolerate the low moisture levels
which occur at these sites particularly during
summer. C. caeruleocinctus, O. sabulosus
and T. niger as well were considered to prefer
warm dry habitats (e.g. HAACKER, 1968;
Thiele, 1968; Dunger & Steinmetzger,
1981) indicating their tolerance for low
humidity. Generally, the species composition
and dominance structure of the diplopod
community at the 1 1 year old fallow was
similar to that of other fallow sites investigated
by STRUWE-KUSENBERG (1981). In comparison to sites of similar age, which had been restored
or developed on raw soils by primary succession (DUNGER, 1968; NEUMANN, 1971), the
diplopod commmunity at the 1 1 year old fallow resembled those of restored afforested sites.

In comparison with the younger fallow sites, and the beechwood, the ash dominated wood
(ca 50 year old fallow) was an extreme habitat for millipedes. Only few species were present and
the density and biomass was low, resembling that of the wheat field. Low diversity, density and

Fig. 7. — Canonical correspondence analysis biplot with
environmental variables [amount of carbon in the
litter layer (C lit), C/N ratio of the litter (CN lit) and
gravimetric water content in the litter layer (H20
lit) and in 0-3 cm soil depth (H20 0-3)] represented
by arrows; (a) ordination of the composition of the
diplopod community from five sites representing
different stages of secondary succession with
centroids of the Five sites; (b) ordination of samples
taken at the five sites (data from 1988; for details
see text).

Source : MNHN. Paris

THE MILLIPEDE COMMUNITY OF A SECONDARY SUCCESSION

655

biomass of diplopods m_the ash dominated wood was unexpected because it has been shown
c eat ltter !s,a Preferred food substrate by diplopods (DUNGER, 1958, 1962-

™ I974) Howeyer, ash leaves are also known to decompose quickly. Ash leaf litter in
the ca 50 year old fallow had already disappeared almost entirely by May. The litter layer during
summer, until shedding of leaves in autumn, consisted mainly of small branches and twigs from
the overgrown shrub layer. The woody materials were presumably of low nutritional value for
diplopods and did not form a litter layer suitable for buffering extremes in temperature and

The beechwood was characterized by the absence of species typical of the open fallow
sites and the arable field and by the occurrence of G. conspersa. In addition, G. mareinata and
M. germamcum were most abundant at this site. CCA indicated that the occurrence of these
species was related to humid conditions. It has been frequently found that these species
preferentially colonize wooded habitats, especially G. conspersa which is known to occur almost
exclusively in moist woodlands (BEYER, 1964; THIELE, 1968; BROCKSIEPER. 1976). The
colonization of more humid ecosystems by these species presumably is related to a low tolerance
or dry conditions during summer.

The content of organic matter in the upper soil layers at the five study sites indicate that
S, option was accompamed by a strong increase in soil humus. It was hypothezised by
NCHEU ( 1 792) that this increase is of considerable impoitance for the changes in the lumbricid
community during secondary succession. In contrast, results of the CCA in the present study
indicate that the soil humus content is of minor importance in determining the structure of
diplopod communities. Rather, the analysis emphasized the importance of the litter layer,
moo Fhe hl0™* of diPloPods in the beechwood during the investigated period of 1987 and
1988 averaged 443 mg dry mass/m2. Variation between years in density [coefficient of variation
(C v) =1 1 %] was considerably lower than that in biomass (CV = 45%). The biomass at the 1 1
year old fallow also varied considerably between 1987 and 1988 (CV = 58%), whereas the
density was almost identical (CV = 1%). There was no unidirectional change in density or
biomass of the diplopod communities at the study sites from 1987 to 1988. The mean annual
biomass of diplopods at the 4 year old fallow and the wheat field remained at a very constant
level. At the 1 1 year old fallow it increased considerably (+138%), whereas in the beechwood it
decreased strongly (-48%). The summer in 1988 was exceptionally dry, particularly in May but
also in August (Fig. 1) Biomass data suggest that these unusually dry conditions did not have
detrimental effects on the diplopod communities of the younger fallow sites and the arable field
but did on that of the beechwood (density at the 4 year old fallow might also have been affected).
Presumably, the diplopod community of the beechwood is more susceptible to drought, despite
the buffering capacity of the canopy of trees and the presence of a litter layer throughout the
year. In a six year investigation SPRENGEL (1986) also concluded that dry summers adversely
affected density and biomass of millipedes at the beechwood, particularly in autumn.

In comparison to other woodlands, the density and biomass of diplopods at the
beechwood was in an approximatly central position (DUNGER, 1958; BLOWER 1979-
Geoffroy, 1981; Meyer et al., 1984; AXELSSON et al., 1984). Diplopods are considered to
be important agents for litter processing and nutrient cycling in forest ecosystems. The
considerably higher biomass of diplopods at the younger fallow sites indicate however that the
effect of diplopods in these ecosystems may even exceed their effect in woodlands.

REFERENCES

AXELSSON, B., Lohm, U. & Persson, T., 1984. — Enchytracids, lumbricids and soil arthropods in a northern deciduous
woodland: A quantitative study. Holarctic Ecol., 7 : 91-103.

Beyer. R., 1964. — Faunistisch-okologische Untersuchungen an Landisopoden. Zool. Jb. Syst., 91 : 341-402.

Blower. J. G., 1979. — The millipede fauna of two British limestone woods. In: M. Camatini .Myriapod Biology
London. Academic Press : 203-214.

656

STEFAN SCHEU

Brocksieper, J., 1976. — Isopoden und Diplopoden des Naiurparks Siebengebirge. Decheniana, 129 : 76-84.

Connell, J. H. & Slatyer, R. O., 1977. — Mechanisms of succession in natural communities and their role in
community stability and organization. Am. Nat.. Ill : 1119-1144.

Dunger. W., 1958. — Uber die Veranderung des Fallaubes im Darm von Bodentieren. Z. Pflanzenernaehr. Dueng.
Bodenkd. , 82 : 174-193.

DUNGER, W.. 1962. — Nahrungswahl bei Bodenarthropoden in produktionsbiologischer Sicht. Verb. XI. Ini. Entomol.
Kongr ., Wien 1960 : 169-173.

Dunger, W., 1968. — Die Entwicklung der Bodenfauna auf rekultivierten Kippen und Halden des Braunkohletagebaus.
Abhandl. Ber. Naturkundemuseum Gorlitz, 43 : 1-256.

Dunger. W. & Steinmetzger, K., 1981. — Okologische Untcrsuchungen an Diplopoden einer Rasen-Wald-Catena im
Thiiringer Kalkgebiet. Zool. Jb. Syst., 108 : 519-553.

Dunger, W., Voigtlander, K., 1990. — Succession of Myriapoda in primary colonization of reclaimed land. In: A.
MlNELLl, Proceedings 7th Intern. Congr. Myriapodology. Leiden, Brill : 219-227.

Geoffroy, J. J., 1981. — Etude d’un Scosysteme forestier mixte V. Traits generaux du peuplement de diplopodes
edaphiques. Rev. Ecol. Biol. Sol. 18 : 357-372.

Haacker, U., 1968. — Deskriptive, vergleichende und experimentelle Untcrsuchungen zur Autokologie rhein-
mainischer Diplopoden. Oecologia. 1 : 87-129.

Herbke, G., 1962. — Untersuchungen uber das Vorkommen von TausendfuBern in landwirtschaftlich genutzten Boden
des Dauerdiingungsversuches auf Dikopfshof. Monogr. Angew. Entomol ., 18 : 13-42.

Kempson, D.. Lloyd, M. & Ghelardi, R.. 1963. — A new extractor for woodland litter. Pedobiologia , 3: 1-21.

Klinger, K.. 1992. — Diplopods and chilopods of conventional and alternative (biodynamic) fields in Hesse (FRG).
Ber. nat.-med. Verein Innsbruck. Suppl. 10 : 243-250.

Meyer, E., Schwarzenberger, J.. Stark, G. & Wechselberger, G., 1984. — Bestand und jahreszeitliche Dynamik der
Bodenfauna in einem Eichenmischwald (Tirol. Osterreich). Pedobiologia, 27 : 115-132.

Neumann, U., 1971. — Die Sukzession der Bodenfauna (Carabidae, Coleoptera; Diplopoda und Isopoda) in den forstlich
rekultivierten Gebieten des Rheinischen Braunkohlereviers. Pedobiologia. 11 : 193-226.

Odum. E. P.. 1969. — The strategy of ecosystem development. Science , 164 : 262-270.

Peters, D.. 1984. — Faunistische und okologische Untersuchungen der Lumbriciden, Diplopoden und Chilopoden auf
verschieden bewirtschafteten Flachen der Niederrheinischen Tiefebene. Ph D Thesis, Bonn, Germany.

Sakwa. W. N., 1974. — A consideration of the chemical basis of food preference in millipedes. Symp. Zool. Soc.
London, 32 : 329-346.

Schauermann. J., 1982. — Verbesserte Extraktion der terrestrischen Bodenfauna im Vielfachgerat modifiziert nach
Kempson und Macfadyen. Arbeitsberichte SFB. 135 (1) : 39-45.

Schaefer, M.. 1991. — The animal community: diversity and resources. In: E. Rohrig & B. Ulrich, Temperate
deciduous forests. Ecosystems of the world 7. Amsterdam, Elsevier : 51-120.

Scheu, S., 1990a. — Changes in microbial nutrient status during secondary succession and its modification by
earthworms. Oecologia, 84: 351-358.

Scheu, S., 1990b. — Die saprophage Makrofauna (Diplopoda, Isopoda und Lumbricidae) in Lebensraumen auf
Kalkgestein: Sukzession und Stoffumsatz. Berichte des Forschungszentrums Waldokosysteme. A 57 : 1-302.

Scheu, S., 1992. — Changes in the lumbricid coenosis during secondary succession from a wheat field to a becchwood
on limestone. Soil Biol. Biochem ., 12 : 1641-1646.

SPRENGEL, T., 1986. — Die DoppelfiiBer (Diplopoda) eines Kalkbuchenwaldes und ihre Funktion beim Abbau dcr
Laubstreu. Ph D Thesis, Gottingen, Germany.

Schulz, E., 1992. — Die Milbenfauna (Acari: Mesostigmata und Cryptostigmata) in Lebensraumen auf Kalkgestein:
Populationsokologie, Sukzession und Beziehungen zum Lebensraum. Berichte des Forschungszentrums
Waldokosysteme, A79 : 1-245.

Struwe-Kusenberg, R., 1981. — Sukzession und trophische Struktur der Bodenfauna von Brachflachen. Pedobiologia,
21 : 132-141.

Tajovsky, K., 1990. — Diplopoda in a secondary soil succession row. In: A. MlNELLl, Proceedings 7th Intern. Congr.
Myriapodology. Leiden, Brill : 229-234.

Ter Braak, C. J. F., 1988. — Canoco - A Fortran programm for canonical community analysis by [partial] [detrended]
[canonical] correspondence analysis, principle component analysis and redundancy analysis. Technical report. TNO
Inst., The Netherlands, Wageningen.

Thiele, H. U., 1968. — Die Diplopoden des Rhcinlandes. Decheniana, 120 : 343-366.

Source : MNHN , Paris

Centipedes from Italian Agroecosystems and their
Possible Value as Pest Control Agents

Marzio Zapparoli

Dipartimento di Protezione delle Piantc, Universita della Tuscia, 01 100 Viterbo, Italy

Abo 1 KAL 1

The centipede communities of agroecosytems (vineyard, hazel-grove, olive-grove, chestnut-grove) and woodland
stands ( Castanea saliva coppiced wood, Quercus cerris woods) in Central Italy were studied in 1984-1991 by pitfall-
trapping tor one year in each site. Nineteen species were collected. In the agroecosystems tt.e number of specie's ranges
trom 2 (vineyard) to 7 (hazel-grove) whereas in oak woods 7 to 10 species were collected. LUhobius lap, dicola Meinert
and L. forficatus (Linne) are the most common species in the agroecosystems. An attempt to assess the possible value of
this group as pest control agent is discussed.

RESUME

Les chilopodes d ’agroecosysten.es d’ltalie et leur valeur possible en tant qu’agents controlant
les ravageurs.

Dcs recherches ont ete menees sur des peuplements de chilopodes dans des agntecosystemes (vignes, bosquets de
no. sellers, ol.vera.es, chata.gneraies) et des sites forestiers (taillis de chataigniers, Castanea saliva , forets de chenes
Quercus cerris) du centre de Pltalic. Les etudes ont etc realisees de 1984 * 1991 & I’aide de pieges d’interception durant un
an sur chaque site. 19 especes ont ete recoltees. Dans les agroecosystemes. le nombre des especes varie de 2 (vignes) a 7
(noisetiers alors que dans les forets de chenes, on trouve de 7 * 10 especes. LUhobius lapidicola Meinert et LUhobius
jorjicatus (Linne) sont les especes les plus communes dans les agroecosystemes. Une tentative d’accorder h ces groupes
une valeur d agent de controle des ravageurs est discutee.

INTRODUCTION

The centipede communities of Italian agroecosystems are little known and the few data
available are mostly synthetized in MlNELLI & IOVANE (1987). Other informations are in
Daccordi & Zanetti (1987) on NE Italy vineyard, PAOLETTI (1988) on NE Italy maize
monocultures, and ZAPPAROLI & TREMATERRA (1993) on N Italy apple orchads. Data
concerning chilopods as a component of the soil community are in PAOLETTI (1980, 1988) on
NE Italy maize agroecosystems, JONA LASINIO & ZAPPAROLI (1993) and TESTA & ZAPPAROLI
(1994), on olive-grove and on hazelnut-grove in Central Italy.

The aim of this paper is (i) to describe the centipede communities in some agroecosystems
of Central Italy, (ii) to verify the influence of the agricultural and sylvicultural activities on the
qualitative composition of such communities and (iii) to assess the possible role of these
polyphagous predators as pest control agent in integrated control programs.

Zapparoli. M., 1996. — Centipedes from italian agroecosystems and their possible value as pest control agents.
In: Geopfroy. J.-J., Mauries. J.-P. & Nguyen Duy - J.ACQUEMIN. M„ (eds). Acta Myriapodologica. Mem. Mas. natn
Hist, not., 169 : 657-662. Paris ISBN : 2-85653-502-X.

658

MARZIO ZAPPAROL1

SAMPLING SITES AND METHODS

Sampling sites

Nine sites have been sampled in different localities of the Viterbo province, Latium, Central Italy: sites 1-7 are
agroecosystems, representative of the predominant cultivations of the area, sites 8-9 are natural or seminatural stands,
representative of the local forest vegetation potentially occurring at the same altitudinal level. The main characteristics
of each site are summarized below :

1) Bassano in Teverina, 300 m a.s.l., a 15 year old vineyard, 5 ha, E exposure, bordering on open cultivated
fields. Agronomic practices: chemical manures in March, May; fungicide treatments all over the cultural cycle,
cultivations in spring and summer, no insecticide and herbicide treatment.

2) Caprarola, Mt. Venere, 500 m a.s.l., a 25-30 year old hazel-grove, 4 ha, SE exposure, bordering on
broadleaved mixed woods and hazel-groves (see also Testa & Zapparoli, 1994). Agronomic practices: chemical manures
in February, cultivations in April, May, August, insecticide treatments (Endosulfan 100 g/1000 1 H20) in May, June,
July, herbicide treatments (Glyphosate) in July, September.

3) Soriano al Cimino, St. Eutizio, 350 m a.s.l., a 20 year old hazel-grove, 3 ha, SE exposition, bordering on
hazel-groves (see also Testa & Zapparoli, 1994). Agronomic practices: chemical manures in February, cultivations in
April, May, June. July, August, no insecticide and herbicide treatments.

4) Canino, 120 m a.s.l., a 35-40 year old olive-grove, 4 ha. W exposure, bordering on open cultivated and grazed
areas (see also Jona Lasinio & Zapparoli, 1993). No agronomic practices.

5) Soriano al Cimino, St. Eutizio, 350 m a.s.l., chestnut-grove, 1 ha, W exposure, bordering on chestnut-groves
and hazel-groves. Agronomic practices: cultivations in May. September, no insecticide and herbicide treatments.

6) Canepina, 650 m a.s.l., chestnut-grove, 4 ha, SE exposure, bordering on chestnut-groves. Agronomic
practices: cultivations in May and September, no insecticide and herbicide treatements.

7) Caprarola. Poggio Nibbio, 550 m a.s.l.. Castanea sativa coppiced wood, SW exposure.

8) Caprarola, Mt. Venere, 560 m a.s.l., Quercus cerris wood with Ostrya carpinifolia and Acer obtusatum , E
exposure.

9) Caprarola, Poggio Nibbio, 580 m a.s.l., Q. cerris coppiced wood, SW exposure.

The area is marked by Mediterranean temperate climate, with a mean annual rainfall ranging from 700 to 1400
mm, max in October-November. min in July, mean annual temperature = 14.8 °C, max in July, min in December, aridity
period in July/August. The Mediterranean characteristics of the climate in site 4 (Canino, olive-grove) are stronger than
the other sites, according to its geographic position closer to the Thyrrenian sea coast. The geological substratum of the
whole area is of volcanic origin (Pleistocenic), soils are clayey (Testa & Zapparoli, 1994).

Methods

Sampling has been carried out between 1984-1991 by pitfall trapping (see Jona Lasinio & Zapparoli, 1993). Six
traps with a 4% solution of formaldehyde in vinegar are positioned in each site for one year. Traps have been located at
about 10 m one from the other along one or two rows, generally at the bases of trees, and emptied monthly. Because of
limitations in the use of pitfall traps, especially for Scolopendromorpha and Geophilomorpha (Zapparoli, 1992), the
results will be discussed mainly qualitatively. The faunistic similarity among the sampling sites has been calculated
using the Jaccard (1908) index; the hierarchic classification of the sites was undertaken using the average linkage
clustering method (UPGMA).

RESULTS

Nineteen species of centipedes were collected in the sampling sites with a total of 148
specimens (Table 1). More in detail, 14 species were collected in agroecosystems (sites 1-6), 12
species in the Quercus cerris and coppiced Castanea sativa stands (sites 7-9). In the
agroecosystems the number of species ranges from 2 (site 1, vineyard) to 7 (site 3, hazel-grove),
whereas in oak woods 7 (site 9) to 9 (site 8) species were recorded.

A clustering of the nine sampling sites according to the similarities of their centipede fauna
(cophenetic correlation index = 0.82) is shown in Figure 1. Cutting the dendrogram at low
similarity level (S = 0.20) results in three clusters.

The first cluster groups vineyard and hazel-grove agroecosystems. Euryecious species are
generally predominant in these habitats. L. lapidicola and L. forficatus are the most common
centipedes. These species were present in the sampled woodlands as well, but the number of
individuals recorded there was lower. Woodland species such as Cryptops anomalous, L.
acuminatus, L. calcaratus, L. castaneus and L. tylopus were represented in hazel-grove as well
but infrequent.

Source : MNHN. Paris

CENTIPEDES IN TAL1AN AGROECOSYSTEMS

659

Table I. — Centipedes collected in sites sampled (number of ind.). I = Bassano in Teverina. vineyard; 2 = Mt Venere
hazel-grove; 3 = St. Eutizio, hazel-grove; 4 = Canino, olive grove; 5 = St. Eutizo. chestnut-grove; 6 = Canepina!
chestnut-grove; 7 - Poggio Nibbio, Castanea saliva coppiced wood; 8 = Mt. Venere, Quercus cerris wood; 9 =
Poggio Nibbio, Q. cerris wood. H = habitat preference ; e = euryecious species, w = woodland species. C =
chorotype : eur = European, sie = Sibiric-european, med = Mediterranean, tern = Turanic-european-mediterranean,
wmd = W-Mediterranean.

species / sites

Himantarium gabrielis (Linn6)

med

Henia vesuviana (Newport)

seu

Schendyla nemorensis (C. L. Koch)

cur

Geophilus flavus (De Geer)

sie

G. linearis C. L. Koch

sie

Pachymerium ferrugineum (C. L. Koch)

tern

Strigamia crassipes (C. L. Koch)

eur

Cryptops anomalans Newport

eur

C. hortensis Leach

C. parisi Brolemann

eur

Eupolybothrus fasciatus (Newport)

seu

E. nudicomis (Gervais)

wmd

Lithobius acuminatus Brolemann

eur

L cal carat us C. L. Koch

eur

L. castaneus Newport

* 2

seu

L. forficatus (Linne)

eur

L. lapidicola Meinert

eur

L. romanus Meinert

seu

L tylopus Latzel

seu

tot species (19)

tot specimens (148)

1 1

euriecious spp. %

100

42.8

33.3

14.3

33.4

12.5

36.8

woodland spp. %

57.2

66.6

100

85.7

66.6

87.5

63.2

The second cluster groups together chestnut-groves and wood-land sites. The centipede
community recorded in these sites is mostly represented by common woodland species of the
broadleaved Central Italy woods, such as Henia vesuviana, Cryptops parisi, Eupolybothrus
fasciatus, Lithobius castaneus and L. tylopus. Other woodland species, such as Strigamia
crassipes and E. nudicomis, are present only in the two Q. cerris sampled sites. Anthropophilus
or euryecious species, such as Geophilus flavus, Pachymerium ferrugineum and L. forficatus,
are also present in the sampled Q. cerris and

coppiced C. sativa woods.

The third cluster is only represented by
the olive-grove site. The centipede community
is mainly represented here by woodland
species, of which L. tylopus is the most
abundant.

The centipede communities of the
sampled agroecosystems seem to be well
related to the whole centipede community of
the submediterranean-submontane belt of
Central Italy, at least in the general features.

Fig. 1. — Dendrogram of hierarchical classification of the
nine sampling sites. See Table 1 for abbreviations.

0.0 0.5 1.0

660

MARZIO ZAPPAROL1

The anthropic impact on the pre-existent communities seems to be low. especially in
chestnut-groves and hazel-groves where some woodland species are represented and a faunistic
exchange between the adjacent forest habitats is probably occurring. The lower number of
species generally recorded in the agroecosystems, as compared with woodlands, is probably
related to the occurence of agronomic pratices, especially of the cultivations. The high frequency
of these may induce changes of some soil microconditions, such as humidity degree and litter
structure, and changes in community structure may be induced (TESTA & ZAPPAROLI, 1994).

It is of some interest to verify the potential value of centipedes in the control of
phytophagous insects partially developing in soil, such as Coleoptera, Diptera and Lepidoptera,
in agroecosystems where the stucture of vegetation is similar to that of the forest habitats
(ZAPPAROLI & TREMATERRA, 1993). This potential function has already been pointed out for
other major polyphagous predator Arthropoda inhabiting open European agroecosystems
(cereals, arable field, etc.), such as Aranea, Coleoptera Carabidae and Staphylinidae (BRIGNOLI,
1983; Sunderland & Chambers, 1983). Besides, among Chilopoda especially Lithobiidae
have been pointed out as active soil-inhabiting predators in many European and Mediterranean
agroecosystems (SUNDERLAND. 1975; CAUSSE, 1976; TEMERAK. 1983).

Among the agroecosystems of the study area, centipedes are generally well represented in
species and numbers especially in hazel- and chestnut-groves which represent the most important
local cultivations. In hazel-groves, key pest in Central Italy is Balaninus nucum L. (Coleoptera
Curculionidae), this species completes one generation every 2-5 years and spends part of the
biological cycle in soil as mature larval stadium, pupa or adult as well, from June-July to the end
of spring, 15-50 cm deep (PUCCI, 1992).

According to the results of the sampling discussed above, the activity peak observed in the
centipede communities of the study area matches well with the phenology of the mature larvae
and pupae of B. nucum (Table 2). Moreover, in hazel-grove centipede community, the most
important species from the quantitative point of view is L. forficatus, which seems to be active
all over the year with a peak in July-September (Table 2). This species is well known as a
pioneer in artificial and disturbed European habitats and it also frequently occurs in Italian
agroecosystems (MlNELLI & IOVANE, 1987; PAOLETTI, 1988; ZAPPAROLI & TREMATERRA,
1993). L. forficatus usually preys on a large spectrum of small soft-bodied invertebrates
species, mainly arthropods and annelids (LEWIS, 1981) it has been already recorded as an
important predator of pupae of Rhagoletis pomonella Walsh (Diptera Tephirtidae), a key pest in
the Canadian apple ochards (MONTE ITH, 1975, 1976).

Table 2. — Number of individuals of centipedes (in brakets L. forficatus) collected monthly in the sites sampled.
1 = Bassano in Teverina. vineyard: 2 = Mt. Venere. hazel-grove; 3 = St. Eulizio, hazel-grove; 4 = Canino. olive
grove; 5 = St. Eutizo, chestnut-grove; 6 = Canepina, chestnut-grove; 7 = Poggio Nibbio, Castanea saliva
coppiced wood; 8 = Mt. Venere, Quercus cerris wood; 9 = Poggio Nibbio, Q. cerris wood.

sites/months

J-A

2(1)

2(2)

1(1)

l(-)

3(-)

4(2)

6(6)

5(3)

2(1)

i(-)

K-)

7(3)

4(4)

Id)

5(2)

9(5)

2(2)

2(1)

2(-)

3(-)

K-)

2(-)

8(-)

5(-)

l(-)

2(-)

U-)

2(1)

3(-)

2(-)

l(-)

2(-)

l(-)

3(-)

2(-)

l(-)

2(-)

Id)

2(-)

3(2)

Kl)

K-)

2(-)

l(-)

3(-)

H-)

l(-)

2(1)

2(-)

Ul)

2(-)

2(1)

K-)

8<-)

l(-)

1 (-)

l(-)

Source :

CENTIPEDES IN ITALIAN AGROECOSYSTEMS

661

CONCLUSION

Tentatively, we hypothesize that L.forficatus can play an important role among predators
in the biocontrol of B. nucum in Central Italy hazel-groves.

More evidence is necessary however, to support the possible role of Chilopoda as pest
control in agroecosystems. As SUNDERLAND & CHAMBERS (1983) suggest, many problems
will have to be solved in order to assess the potential value of such predators: field and
laboratory trials on which species are preyed on and the ability to prey are needed together with
informations on the predators behaviour, ecology and density during the years in different sites
and under different farming conditions.

On the other hand, as stated by WATERHOUSE (1969), the individual number of centipedes
in the field may be too low to give an adequate control of pest. As pointed out by several
European studies (see SUNDERLAND & CHAMBERS, 1982; THOMAS et at. , 1992), the activity of
these predators should therefore be useful in integrated pest control programs in cooperation
with other polyphagous Arthropoda, specialist parasites and predators, under more favourable
farming conditions such as reduced inputs of pesticides, reduced rates of cultivations and
increased and conserved structural diversity within the agroecosystems.

ACKNOWLEDGEMENTS

This research has been partly supported by grant of the Italian Ministero della Ricerca Scientifica e Tecnologica.

REFERENCES

Brignoli, P. M., 1983. I ragni quali predatori di insetti: il loro potenziale ruolo negli agroecosistemi (Araneae) In
Alii XIII Congr. Naz. II. Em.. Sestriere. Torino : 591-597.

Causse, R., 1976. — Etude de la localisation et de la mortalite hivernalc des larves de Laspeyresia pomonella L.
(Lepidoptera, Tortricidae) en vergers modernes de pommiers de la basse Vallee du Rhone. Ann. ZooL Ecol. Anim.. 8 :
83- 101.

Daccordi, M. & Zanetti, a., 1987. — Catture con trappole a caduta in un vigneto nella Provincia di Verona. Quad. Az.
Agraria Sper. Villafranca, 3 : 1-44.

Jaccard, P., 1908. — Nouvelles recherches sur la distribution florale. Bull. Soc. Vaud. Sci. Nat., 44 : 223-270.

Jona Lasinio, P. & Zapparoli, M., (1993). — First data on the soil Arthropod community in an olive grove in Central
Italy. In : M. PAOLETTI, W. FoiSSNER & D. COLEMAN, Soil Biota . Nutrient Cycling and Farming Systems. Proc. Ini.
nTi 2\gr0eC0l°gy and conservation issues in temperate and tropical regions", Chelsea, USA, Lewis Publisher .

Lewis, J. G. E., 1981. — The biology of centipedes. Cambridge, Cambridge University Press, 476 pp.

MlNBLLl, A. & Iovane, E., 1987. — Habitat preferences and taxocenoses of Italian centipedes (Chilopoda). Boll. Mus.
civ. St. nat. Venezia , 37 : 7-34.

Monteith, L. G.. 1975. — Laboratory feeding studies on potential predators of the apple maggot Rhagoletis pomonella
(Diptera Tephritidae) in Ontario. Proc. entom. Soc. Ontario , 106 : 28-33.

Monteith, L. G., 1976. — Field studies on potential predators of the apple maggot Rhagoletis pomonella (Diptera
Tephritidae) in Ontario. Proc. entom. Soc. Ontario , 107 : 23-30.

PAOLETTI, M. G., 1980. — L'agroecosistema a mais nella pianura veneta con raffronti all’ambiente planiziale forestale.
Alcune valutazioni qualitative e quantitative sugli invertebrati del suolo. In: Atti I Congr. naz. s. It. E. : 359-369.

PAOLETTI, M. G., 1988. — Soil invertebrates in cultived and uncultivated soils in Northeastern Italy. Redia, 71 : 501-
563 .

PUCCI, C., 1992. Studies on population dynamics of Balaninus nucum L. (Col. Curculionidae) noxoius to the hazel
( Corylus avellana L.) in Northern Latium (Central Italy). J. Appl. Ent., 114 : 5-16.

Sunderland, K. D., 1975. — The diet of some predatory arthropods in cereal crops. J. Appl. Ecol., 12 : 507-515.
Sunderland, K. D. & Chambers. R. J., 1983. — Invertebrate polyphagous predators as pest control agents: some
criteria methods. In : R. Cavalloro. Aphid antagonists, Proc. Meeting ZC Experts ’ Group, Rotterdam, Balkema :
100-108.

Iemerak, S. A., 1983. — A preliminary survey on the soil-inhabiting predaceous arthropods associated with pupae of
the large sucarcane borer. Sesamia cretica in sorghum field. Bull. Soc. ent. Egypte. 62 : 251-255.

662

MARZIO ZAPPAROLI

Testa. G. & Zapparoli. M., 1994. — First data on soil Arthropod communities of hazel groves in Central Italy. [In:

Proc. Ill ISHS hit. Congress Hazelnut, Alba, Italy]. Acta Horticulturae , 351 : 631-637.

Thomas, M. B.. Wratten, S. D. & Sotherton, N. W.. 1992. — Creation of “island” habitats in farmland to manipulate
populations of beneficial arthropods: predator densities and species composition. J. Appl. Ecology, 29 : 524-531.
Waterhouse, J. S., 1969. — An evaluation of a new predaceous centipede Lamyctes sp., on the garden symphylan
Scutigerella immaculata. Can. Entom., 101 : 1081-1083.

Zapparoli M., 1992. — Preliminary data on centipede communities of Quercetea ilicis and Fagetalia sylvaticae in
Central Italy. Ber. nat. -tried. Verein Innsbruck, suppl. 10 : 197-204.

Zapparoli. M. & Trematerra, P.. 1993. — Chilopodi nei meleti della Valtellina. Boll. Zool. agr. Bachic. (II), 25 : 41-
48.

Source : MNHN, Paris

Author Index / Index des auteurs

Abrous-Kherbouche, Ourida . 175

Adis, Joachim . 523, 607

Adolphe, Jean-Pierre . 555

Aguilar, Franklin . 493

Andrejko, Mariola . 431

Bano. Kubra . 73, 561

Barber, Antony D . 243

Barnett, Mandy . 331

Barrientos, Zaidett . 493

Bouzerna, Noureddine . 365

Branquart, Etienne . 485

Cancela da Fonseca, Jorge P . 577

C£l£rier, Marie-Louise . 533

Ceuca, Traian . 61

Compare, Philippe . 395

CondG, Bruno . 127

Daas, Tarek . 365

Dangerfield, John Mark . 565, 617

David, Jean-Frangois . 461, 627

DeMorais, Jos6 Wellington . 607

DEFIZE, St6phane . 395

Demange, Jean-Marie . 21

Descamps, Michel . 365, 385, 391

Dohle, Wolfgang . 371

Dominguez Rodriguez, Maria Teresa . 137

ElAissaoui, Hsai'n . 555

Enghoff, Henrik . 313

Fabre, Marie-Chantal . 385. 391

Fischer, Elisabeth . 451

Fusco, Giuseppe . 351

Garcia Ruiz, Andres . 205, 347

Gaspar, Charles . 485

Geoffroy, Jean-Jacques . 13, 269, 533

Gerard, Sylvie . 391

GOFFINET, Gerhard . 395

Golovatch, Sergei Illich . 163, 265, 523

Greven, Hartmut . 403

Hamann, Susanne . 523

Hill, Trevor J . 441

Hopkin, Stephen P . 25

Ishii, Kiyoshi . 101

Jamault-Navarro, Catherine . 385

Jarosz, Jan . 431

Jeanson, Colette . 555

Jedryczkowski, Wojcieh B . 209

Jones, Richard E . 243

Kania, Grzegorz . 431

Kime, Richard Desmond . 257

K ors6s, Zolt&n . 35

Kos, Ivan . 635

Kraus, Margarete . 283

Kraus, Otto . 283

Lesniewska, Malgorzata . 22 1

Lewis, John G. E . 441

Mahsberg, Dieter . 585

Marsoner, Peter . 451

Martens, Jochen . 1 63

Mauri£s, Jean-Paul . 81

Mateos, Eduardo . 1 87

Matic, Zachiu . 225

Mesibov, Robert . 139, 341

Meyer, Erwin . 451

Meziane, Leila . 577

Mi nelli, Alessandro . 351

Miquel, Maria Carme . 359

Monge-Najera. Julian . 493

Negrea, Stefan . 225

NEGRISOLO, Enrico . 351

Nevermann, Lutz . 421

Nguyen Duy - Jacquemin, Monique . 1 13

Pass, Gunther . : . 291

Pedroli-Christen, Ariane . 45, 53, 281

Pereira, Luis Alberto . 79

Prunescu, Carol Constantin . 299, 341. 437

Prunescu, Paula . 437

Revault, Pascal . 495

Ribarov, Goergi . 235

Rosenberg. Jorg . 403

Ruhberg, Hilke . 1 39

Sahli, Francois . 373, 587

Santibanez, Francisco J . 205, 347

Scheller. Ulf . 607

Scheu, Stefan . 647

SCHILEYKO, Arkady A . 293

Scholl, Adolf . 45. 53

Serra, Antoni . 187, 359

Shinohara, Keizaburo . 341

Spelda, Jorg . 151

Stefaniak, Malgorzata . 431

Striganova. Bella R . 515

Sustr, Vladimir . 473

Tabacaru, Ionel . 67

Tadler, Andreas . 327

Tajovsky, Karel . 509

Tamura, Hiroshi . 101

Telford, Steven R . 331, 565, 617

Vannier, Guy . 461

Vicente, Maria Cristina . 187

Voigtlander. Karin . 501

Wakley. Gavin E . 441

Wang, Daqing . 81, 307

Weyda, Frantizek . 483

Wytwf.r, Jolanta . 213

Xylander, Willi E. R . 411, 421

Zalesskaya. Nadezhda . 265

Zapparoli, Marzio . 599, 657

Zulka. Klaus Peter . 477

Source

Source : MNHN, Paris

SYSTEMATIC INDEX

Adenomeris gibbosa 245, 247. 248, 249, 275
Adenomeris hispida 275
Agariogonopus acrotrifoliatus 85
Alaskobius takakuwai 91
Alipes 294, 295

All aj ulus nitidus 245, 248, 258, 260, 262. 278, 374.
488, 533, 537, 545, 546, 547, 548, 549. 550, 647,
649, 650, 651, 652
Allajulus spinosus 326
Allopauropus ovalapendicis 91
Allopauropus pilosisphaerus 91
Alloporus 333, 334, 337

Alloporus uncinatus 333, 335, 337, 567, 568, 569, 570,
571, 573, 574, 575, 618, 619, 620, 621. 622, 623,
624

Alloproctus 133
Alogolykinae 74, 86
Alogolykini 75, 86
Alpiobates peyerimhoffi 277
Alpityphlus seewaldi , 154, 157
Ahum 309

Altum carinatum 307, 308, 309, 310
Altum serratum 307, 308, 309, 310
Altum viriosum 307. 308, 309, 310
Amblyiulus 85
Amblyiulus barroisi 326
Amblyiulus creticus 326
Ameractis chirogon 326
Ampelodesmus 1 1 1
Ampelodesmus granulosus 102, 108
Anamastigona pulchella 276

Anaulaciulus 35, 36, 42, 167, 172, 173, 313, 318, 324

Anaulaciulus acaudatus 36, 39, 167

Anaulaciulus acutus 36, 39, 41

Anaulaciulus attemsii 36, 39

Anaulaciulus bilineatus 37, 39. 40. 42, 167

Anaulaciulus bilobus 37, 39, 41

Anaulaciulus capillatus 37. 40

Anaulaciulus ciliatus 36

Anaulaciulus cornutus 37, 39

Anaulaciulus enghoffi 37. 39

Anaulaciulus golovatchi 37. 40, 41, 42

Anaulaciulus hirosaminus 37, 39

Anaulaciulus inaequipes 37, 39, 40, 41, 42, 317, 326

Anaulaciulus kashmirensis 37, 39, 167

Anaulaciulus kiusiensis 37, 40, 41

Anaulaciulus komatsui 37, 39

Anaulaciulus koreacolus 37, 39, 42

Anaulaciulus koreanus 37. 39. 40. 41

Anaulaciulus koreanus boninensis 37

Anaulaciulus koreanus tuberculalus 37

Anaulaciulus kurilai 37, 39

Anaulaciulus long us 37, 39

Anaulaciulus nepalensis 37, 38, 39, 42, 167

Anaulaciulus niger 38, 39, 42. 167

Anaulaciulus okinawaensis 38, 39
Anaulaciulus onychophora 38, 39
Anaulaciulus otigonopus 38, 39, 41
Anaulaciulus pakistanus 38, 39, 42
Anaulaciulus paludicola 36, 38, 39, 40, 42, 85
Anaulaciulus pinetorum 38, 39
Anaulaciulus pinetorum nivalis 38
Anaulaciulus quadratus 38, 39, 41
Anaulaciulus riedeli 38. 39, 41, 42
Anaulaciulus ryugadensis 38, 40, 42
Anaulaciulus simodanus 38, 39
Anaulaciulus simplex 38, 39, 85
Anaulaciulus takakuwai 38, 39
Anaulaciulus takakuwai coloratus 38
Anaulaciulus takanoi 38, 40. 42
Anaulaciulus tibetanus 38, 39, 167
Anaulaciulus tigris 38. 39, 42
Anaulaciulus tonggosanensis 38, 39
Anaulaciulus tonginus 39, 41, 42, 85, 326
Anaulaciulus topali 39, 167
Anaulaciulus trapezoidus 39, 85
Anaulaciulus trigonalis 39
Anaulaciulus trilobus 39. 41, 85
Anaulaciulus trilobus khuuae 85
Anaulaciulus trilobus quemoyensis 85
Anaulaciulus vcHlicola 36, 39, 40. 85
Anaulaciulus yamashinai 39, 42
Anaulaciulus yosidanus 39, 40
Andrognathidae 84, 167, 171
Aniulus 326
Anoplodesmus 74
Anoplodesmus anthracinus 74
Anoplodesmus at opus 74
Anoplodesmus indus 74
Anoplodesmus insignis 74
Anoplodesmus saussurii 74
Anoplodesmus tanjoricus 74
Anopsobiidae 346
Anopsobiini 304. 341
Anopsxenus 122

Anopsxenus caboverdus 113, 114, 121, 122, 123. 125
Anopsxenus indicus 113, 114, 121. 122, 123, 125
Anthogona britannica 244, 245, 247, 248,

Anthogona variegata 276
Antichirogonus 75
Antichi rogonus hirtus 75
Antichirogonus laevisulcatus 75
Antisoma 74
Aphelidesmus 74
Aponedyopus jeanae 87
Aponedyopus maculatus 87
Aponedyopus montanus 87
Aponedyopus reesi 87
Aprosphylosoma darceneae 326
Aprosphylosomaiidae 326

Archiboreoiulus pallidus 245, 247, 248. 258, 260, 271,
277

666

SYSTEMATIC INDEX

ArChiboreoiulus sollaudi 211

Archileucogeorgia 318, 326

Archilithobius carinatus 225

Archipolydesmus 180, 182. 185

Archipolydesmus osellai 190. 193

Archipolydesmus ribauti 277

Archispirostreptus tumuliporus 470

Arebius bidens 90

Arebius chengsiensis 90

Armolites 169

Armolites chulingensis 169

Armolites comm unicans 169

Armolites similis 169

Armolites spiniger 76, 169

Arrupinae 89

Asanada 294, 295

Asanada socotrana 448

Assamodesmus 168

Assamodesmus lindbergi 168

Atractosoma montivaga 53, 54, 55, 57, 58

Atractosoma montivaga hessei 55

Atractosoma montivagum silvaticum 54

Bacillozonium nodulosum 515
Ballophi linae 88
Ballophilus liber 88
Banatodesmus jeanneli 68
Baskoiulus stammeri 326
Basoncopus filiformis 326
Bhutanodesmus 169
Bhutanodesmus velatus 169
Bilingulidae 83
Bilingulus sinicus 85
Blaniulidae 326. 650
Blaniulidea 272, 277, 278
Blaniulus dollfusi 190, 193

Blaniulus guttulatus 211, 245, 246, 247, 248, 250. 258,
277, 374, 628. 647, 649, 650, 651, 652. 654
Blaniulus lichlensteini 277
Blaniulus lorifer 277
Blaniulus mayeti 277
Blaniulus orientalis 277
Blaniulus troglobius 277
Blaniulus troglodites 277
Blaniulus velatus 277
Blaniulus virei 278
Bollmania 86
Boreoiulus dollfusi 277
Boreoiulus simplex 278
Boreoiulus tenuis 211, 245, 248, 258, 278
Bothriogaster signata 238, 600, 601
Bothropolys 366
Bothropolys asperatus 90
Bothropolys crassidentatus 91
Bothropolys elongatus 366
Bothropolys imaharensis 91
Bothropolys richthofeni 91
Bothropolys shansiensis 9 1

Brachychaeteuma 247

Brachychaeteuma bagnalli 245, 248, 271, 276
Brachychaeteuma bradae 245, 248, 276
Brachychaeteuma cadurcensis 276
Brachychaeteuma f urea turn 276
Brachychaeteuma melanops 245, 247, 248, 276
Brachychaeteuma peniculatum 276
Brachychaeteuma plumosum 276
Brachychaeteuma provincial 276
Brachycybe lecontii 483
Bracliydesmus 101
Brachydesmus exiguus 277
Bracliydesmus proximus 277

Brachydesmus superus 211, 245, 246, 247, 248, 250,
258, 260, 277

Brachygeophilus truncorum 215, 219, 222, 223
Brachyiulidae 328
Brachyiulinae 36

Brachyiulini 36, 173, 317, 324, 326
Brachyiulus 314, 317, 481
Brachyiulus apfelbecki 317, 326
Brachyiulus bagnalli 328, 477, 478, 479, 480
Brachyiulus lusitanus 278
Brachyiulus pusillus 245, 248, 258. 260, 278
Brachyschendyla 639

Brachyschendyla montana 638, 639, 640, 641, 642, 643,
644, 645

Brachyschendyla varnensis 238
Broelemanneuma 273
Broelemanneuma furcation 274, 276
Broelemanneuma gayi 21 A , 276
Broelemanneuma gineti 21 A, 276
Broelemanneuma palmatum 21 A, 276
Broelemanneuma pectiniger 274, 276
Buchneria cornuta 321
Buchneria sicula 321

Californiulus yosemitensis 326
Callipodidea 276, 307
Call ip us corsicus 276
Callipus foetidissimus 276, 596
Callipus sorrentinus 276
Calostreptus 566, 572, 574

Calostreptus carinatus 567, 569, 570, 571, 573, 618,
620, 621, 622

Calyptophyllini 313, 315, 319, 320, 324, 326
Calyptophyllum digilatum 319, 326
Calyptophyllum trapezolepis 319, 326
Cambalidae 85
Cambalopsidae 168, 171
Camptogona delamarei 276
Camptogona dubosequi 276
Caspiopetalidae 86
Catamicrophyllini 315, 324
Catamicrophyllum 320
Catamicrophyllum caifanum 320, 326
Catamicrophyllum mesorientale 319, 326
Centrodesmus 102

SYSTEMATIC INDEX

667

Cephalofovea tomahmontis 148
Ceratosphys 190, 193
Ceratosphys amoena 276
Ceratosphys banyulsensis 276
Ceratosphys guttata 276
Ceratosphys nivium 276
Ceratosphys picta 276
Ceratosphys simoni 276
Ceratosphys vandeli 276
Cermatidae 32
Cermatobiidae 300, 301
Chaetechelyne 639
Chaetechelyne illyriaca 642
Chaetechelyne scheerpeltzi 638, 639
Chaetoleptophyllum 326
Chalandea 639

Chaleponcus 333, 335, 337, 338, 570, 572, 574, 623
Chaleponcus digitatus 567, 569, 570, 571, 573. 574
619, 620, 621

Chaleponcus limbatus 333, 334, 337. 570 571 573
619, 621, 622

Chamaesoma broelemanrii 276
Chamberlinini 87
Chamberlinius cristatus 102
Chamberlinius haulienensis 87, 102
Chamberlinius pekuensis 87
Chamberlinius picrofasciatus 87
Chamberlinius shengmui 87
Chelodesmidea 88
Chelojulidae 326
Chelojulus sculpturatus 326
Chersastus 566
Chersastus sanguinipes 569
Chersoiulus sphinx 326

Chicobolus 411, 412, 413, 414, 415, 416. 417, 418, 434

Chilexenus 121

Chinobius chekianus 90

Chinobius chekianus tumeopes 90

Chinobius pachypedatus 90

Chinobius sachalinus 90

Chinobius sulcipes 90

Chinobius svenhedini 90

Chinosoma hodites 86

Chino sphaera maculosa 84

Chinosphaera majorina 84

Chinosphaera mullidenta 84

Chondromorpha 74

Chondromorpha kaimura 75

Chondromorpha kelaarti 75

Chondromorpha kelaarti longipes 75

Chondromorpha kelaarti valparaiensis 74

Chondromorpha mammifera 74, 76

Chondromorpha severini 74

Chondromorpha severini vae robusti 74

Choneiulus palmatus 211, 245, 247, 248. 258, 278

Choneiulus subterraneus 278

Chordeuma 244, 245

Chordeuma consoranense 277

Chordeuma iluronense 277

Chordeuma inornatum 277

Chordeuma intermedium 277

Chordeuma montanum 277
Chordeuma muticum 277

Chordeuma proximum 245, 246, 248, 249, 252, 277
Chordeuma reflexum 277

Chordeuma silvestre 245, 246, 248, 258, 260, 262 277
488

Chordeuma trifidum 277
Chordeuma utriculosum 277
Chordeuma vasconicum 277
Chordeumatidae 650
Chordeumatidea 272 277 , 489
Chordeumella 509
Chromatoiulus 318
Chromatoiulus podabrus 318, 326
Cleidogonidae 167
Clinopodes 639

Clinopodes abbreviate 638, 639, 640, 642, 643 644
645

Clinopodes flavidus 222, 223, 238, 601, 602, 641, 644
Clinopodes linearis 215, 219, 222, 223, 638, 639, 641
644

Clinopodes trebevicensis 601, 603. 638, 639. 641 642
643. 644, 645
Cormocephalus 294, 295 '

Corsicosoma legeri 276
Corsikomeris remyi 275
Corypholophus 102, 110
Cranogona dalensi 276
Cranogona delicata 276
Cranogona denticulata 276
Cranogona orientale 276
Cranogona pavida 276
Cranogona touyaensis 276
Cranogona uncinata 276
Cranogona vasconica 276
Craspedosoma 262

Craspedosoma alemannicum 258, 259, 262, 276
Craspedosoma rawlinsii 245, 247, 248. 259, 262, 276,
488, 489

Craspedosoma simile 258, 259
Craspedosoma taurinorum conforme 276
Craspedosomatidae 45

Craspedosomatidea 272, 276, 277, 489, 537
Craterostigmus 346, 437

Craterostigmus tasmanianus 305,341, 342, 346,
437.438. 439, 440
Crenatidorsus grandifoliatus 87
Crossosoma broelemanni 276
Crossosoma cavernicola 271, 276
Crossosoma mauriesi 271, 276
Crossosoma peyerimhofft 276
Cryptocorypha 1 1 1
Cryptocorypha japonica 102, 108
Cryptocorypha spinicoronatus 88
Cryptodesmidae 71, 87, 102, 108, 110, 111, 168. 171
Cryptops 294, 295, 441, 639
Cryptops anomalans 238, 601, 602, 658. 659
Cryptops anomalans anomalans 236
Cryptops anomalans schassburgensis 236
Cryptops croaticus 238, 638
Cryptops hispanus 206. 207

668

SYSTEMATIC INDEX

Crypiops hortensis 222, 223, 238, 601, 602, 638, 640,
641, 642, 643, 644, 645, 659
Cryptops japonicus 90
Crypiops nigropictus 90

Cryptops parisi 236. 601, 638, 639, 640, 641, 642, 643,
644, 659

Cryptops parisi parisi 236

Cryptops parisi rhenanus 236

Cryptops rucneri 638

Cryptops trisulcatus 600. 601

Crypiopsidae 90, 293. 294, 295, 296, 297

Cryptopsinae 90

Ctenophilus 79

Ctenophilus amieti 79

Ctenophilus chevalieri 79

Ctenophilus corticeus 79

Ctenophilus edentulus 79

Ctenophilus magnus 79

Ctenophilus nesiotes 79

Ctenophilus nitidus 79

Ctenophilus oligopodus 79

Ctenophilus pratensis 79

Curiosoma bispinosum 76

Cutervodesmus 71

Cutervodesmus adisi 523. 524, 525, 526, 527, 528. 529.
530

Cylindroiulinae 328
Cylindroiulini 318, 321, 322, 325, 326
Cylindroiulus 212, 325, 627
Cylindroiulus bicolor 322, 326
Cylindroiulus boleti 328, 478, 479
Cylindroiulus britannicus 245, 246. 247. 248
Cylindroiulus broti 278, 322, 326, 597
Cylindroiulus caeruleocinctus 190. 193, 195, 201. 203,
211. 212. 245, 246. 247, 248, 252, 258, 259, 260,
262, 263, 278, 326. 537, 550. 627, 628, 629, 630,
647, 649, 651. 652, 654
Cylindroiulus chalandei 278
Cylindroiulus distinctus 182
Cylindroiulus frisius 211, 212
Cylindroiulus iluronensis 278

Cylindroiulus latestriatus 212, 245, 247, 248, 250, 252,

254, 258, 278, 501, 502, 503. 504, 505, 506, 507,
517

Cylindroiulus latzeli 326

Cylindroiulus laurisilvae 322, 326

Cylindroiulus limitaneus 278, 596

Cylindroiulus londinensis 245, 246, 247. 248, 250. 252,

255, 262, 278, 628, 651
Cylindroiulus nitidus 545
Cylindroiulus parisiorum 245, 247, 248. 278
Cylindroiulus perforatus 326
Cylindroiulus propinquus 326

Cylindroiulus punctatus 245, 246, 247, 248, 258. 259,
260. 278, 326, 488, 504, 505. 517, 533, 537, 544,
545. 547, 548, 549, 550
Cylindroiulus pyrenaicus 278
Cylindroiulus ruber 322 326
Cylindroiulus Sagittarius 278
Cylindroiulus schubarti 278
Cylindroiulus segregatus 278

Cylindroiulus spinosus 278

Cylindroiulus truncorum 211, 245, 247, 248, 255, 257,
258

Cylindroiulus verhoeffi 278

Cylindroiulus vulnerarius 245, 247, 248, 254, 255, 278
Cyrnosoma beroni 276
Cyrnosoma coineaui 276
Cyrnosoma slrasseri 276

Dalodesmidae 71
Dasypharkis 75
Dasypharkis rugulosa 75
Delurodesmus orientalis 88
Desmoxytes cornuta 87
Desmoxytes draco 87
Desmoxytes longispina 87
Desmoxytes minutubercula 87
Desmoxytes piceofasciata 87
Desmoxytes planata 75, 76, 87
Desmoxytes rastrituberus 87
Devillea tuberculata 277
Devonobius delta 289
Dicellophilus 639

Dicellophilus carniolensis 638, 639, 640, 641, 642, 644,
645

Dignathodon microcephalum 206, 207, 238. 240, 347,
348, 349. 600, 601
Dignathodontinae 88
Dimorphodesmus 102, 109, 111
Dinocryptops 294
Diplomaragna formosanum 86
Diplomaragnidae 86
Doderoa gen ue ns is 275
Dolichoiulus 271. 315
Dolichoiulus tongiorgii 271, 278
Dolichoiulus vosseleri 326
Doratodesmidae 87, 102, 109, 110, 111
Doratogonus 333, 334, 337

Enantiulus armatus 245, 246, 248, 249, 252, 254, 278
Enantiulus dentigerus 326

Enantiulus nanus 278, 451. 452, 453, 454, 455, 456,
458, 505

Epanerchodus 107, 111
Epanerchodus eurycornutus 87
Epanerchodus mammillatus 102, 107
Epanerchodus orientalis 87, 102
Epanerchodus potanini 87
Epanerchodus shirinensis 87
Epanerchodus sphaerisetosus 87
Epanerchodus stylotarseus 87
Epanerchodus takakuwai 87
Epinannolene arborea 524
Epiperipatus biolleyi 493, 494
Esastigmatobius 304

Source :

SYSTEMATIC INDEX

669

Esastigmatobius longicornis 91

Esastigmatobius longitarsis 91, 341. 342, 346

Esc ary us japonic us 88

Escaryus latzeli 88

Escaryus sachalinus 88

Escualdosoma gourbaultae 276

Ethmostigmus 294, 295, 441, 442, 444, 447, 448

Ethmostigmus trigonopodus 441, 442, 444, 447, 448

Ethopetolidae 90

Eucondylodesmus 1 1 1

Eucondylodesmus elegans 102, 109

Eudigraphis sinensis 84

Eudigraphis taiwaniensis 84

Eumastigonodesmus boncii 277

Euperipatoides leuckarti 141, 148

Euphoberia 483

Eupolybothrus 225, 226, 232, 603, 604. 639
Eupolybothrus andreevi 238
Eupolybothrus caesar 601, 602, 603, 604
Eupolybothrus elongatus 365, 366, 367, 369
Eupolybothrus fasciatus 225, 226, 229. 238, 659
Eupolybothrus gr os sipes 236

Eupolybothrus litoralis 225, 227, 228, 229, 232, 600,
601, 603, 604

Eupolybothrus nudicornis 659
Eupolybothrus ochraceus 238

Eupolybothrus transsylvanicus 238. 601. 602, 603, 604

Eupolybothrus tridentinus 238. 240, 639, 644

Eupolybothrus valkanovi 238

Eupolybothrus verrucosus 266. 267, 268

Eupolybothrus werneri 601, 602, 603

Eury gyrus 308

Eu ry paii ropodidae 91

Eurypauropodinae 91

Eurypauropus 91

Eutrichodesmus arcicollaris 87

Fagepauropus hesperius 91

Fijiodesmus 74

Fioria tuberculata 276

Formosocephalus longichilatus 88

Fuhrmannodesmidae 67. 71, 168. 171. 172, 524

Fuhrmannodesmini 71

Fusiulus 36, 39

Fusiulus acutus 36

Fusiulus at terns ii 36

Fusiulus bilobus 37

Fusiulus capillatus 37

Fusiulus ciliatus 38

Fusiulus cornutus 37

Fusiulus hirosaminus 37

Fusiulus insularumi 39

Fusiulus kiusiensis 37

Fusiulus komatsui 37

Fusiulus koreanus 37

Fusiulus koreanus boninensis 37

Fusiulus koreanus koreanus 37

Fusiulus koreanus tuberculatus 37
Fusiulus kuritai 37
Fusiulus longus 37, 38
Fusiulus onychophora 38
Fusiulus pinetorum 38
Fusiulus pinetorum nivalis 38, 39
Fusiulus quad rat us 38
Fusiulus simodanus 38
Fusiulus simplex 38
Fusiulus lakakuwai 38
Fusiulus lakakuwai coloratus 38
Fusiulus tonggosanensis 38
Fusiulus trapezoidus 39
Fusiulus trigonalis 39
Fusiulus trilobus khuuae 39
Fusiulus trilobus quemoyensis 36
Fusiulus yamashinai 39
Fusiulus yoshidanus 39

Galliobates gracilis 278, 326

Galliobatidae 326

Galliocookia balazuci 271, 277

Galliocookia fagei 277

Galliocookia leclerci 271, 277

Geoglomeris duboscqui 275

Geoglomeris granulosa , 275

Geoglomeris provincial is, 275

Geoglomeris subterranea , 275

Geophilella pyrenaica 91

Geophilellidae 91

Geophilidae 32, 88

Geophilinae 88

Geophilus 180, 639

Geophilus balcanicus 236 238

Geophilus carpophagus 180, 206. 207, 601. 639

Geophilus elect ricus 213, 215. 222, 223, 238

Geophilus flavus 238. 638. 639, 641, 645. 659

Geophilus infossulatus 88

Geophilus insculptus 601. 638, 643, 645

Geophilus linearis 238, 601, 602, 659

Geophilus oligopus 638, 642, 643, 644

Geophilus promontorii 638

Geophilus proximus 222. 223, 238

Geophilus pygmaeus 639

Geophilus rhodopensis 238

Geophilus strict us 238

Glenniea 169

Glenniea bhotiaensis 169

Glenniea indica 169

Glenniea martensi 169

Glenniea minuscula 1 69

Glenniea perarmata 169

Glomeridae 32, 84, 166. 173

Glomeridea 275, 276, 537

Glomeridella kervillei 275

Glomeris 431, 473, 557, 559. 560. 578, 596

Glomeris annulata 275

Source : MNHN, Paris

670

SYSTEMATIC INDEX

Glomeris balcanica 249, 473, 474, 475, 631
Glomeris bicolor 84

Glomeris connexa 21 1, 212, 275, 431, 432, 433, 434,
578

Glomeris conspersa 180, 258, 275, 647. 649, 652, 655
Glomeris guttata 275
Glomeris Helvetica 275

Glomeris hexasticha 199, 249, 473, 474, 475. 488, 559
Glomeris hexasticha intermedia 190, 191, 193, 195, 196,
199, 201, 258, 260
Glomeris Humbert iana 275
Glomeris intermedia 275

Glomeris marginata 190, 193, 195, 196. 199. 201. 245.
246. 247. 248, 249, 252, 255, 258, 260, 275, 285,

371, 395, 396, 397, 398, 473, 474, 475, 488, 517,

533, 537. 543, 544, 547, 550, 555. 558, 559, 577,

578, 579. 580, 581, 582. 583, 627, 628. 629, 647,

649. 652. 653, 655

Glomeris pustulata 275
Glomeris transalpina 275
Glomeris undulata 275
Glyphiulus adeloglyphus 85
Glyphiulus anophthalmus 85
Glyphiulus balaszi 85
Glyphiulus formosa 85
Glyphiulus granulatus 85
Glyphiulus multicarinus 85
Glyphiulus pulcher 85
Glyphiulus quadrohamatus 85
Glyphiulus recticullus 85
Glyphiulus tuberculatus 85
Glyphiulus vulgatus 85
Gonebelus sinensis 85
Gonographis adisi 480, 524
Gonoplectus 168, 173
Gonoplectus alius 168
Gonoplectus astutus 85
Gonoplectus bhutanensis 168
Gonoplectus broelemanni 168
Gonoplectus corniger 168
Gonoplectus gracilis 168
Gonoplectus hyatti 168
Gonoplectus lindbergi 168
Gonoplectus malayus 168
Gonoplectus probus 168
Gonoplectus remyi 168
Gonoplectus sulcatus 168
Grammorhabdus 76
Grammorhabdus asperrimum 75
Gyrodrepanum 75
Gyrodrepanum contortipes 75

Haasea 154

Haasea flavescens 155, 159, 276

Haasea fonticulorum 451, 452, 454, 457. 458, 459

Haasea norica 1 52, 1 56

Habrodesmus duboscqui 585

Hanseniella arborea 528, 607, 609, 612, 613, 614

Haplodesmidae 87. 102. 110, 111
Haplogona oculodistincta 152, 157
Haplogonosoma implicatum 102, 105
Haplophilus subterraneus 221, 222, 223. 224
Haplopodoiulus 321

Haplopodoiulus spathifer 190, 193, 196, 199, 201, 278,
326

Haploporatia eremita 1 59, 328

Harpagomorpha 75

Harpagomorpha dentata 75

Harpagophoridae 85, 168, 171, 332, 333, 337, 621

Harpaphini 88

Harpolithobius 639

Harpolithobius anodus 238, 601, 639, 640
Harpolithobius anodus dentatus 357
Harpolithobius aseni 238
Harpolithobius banaticus 239, 357
Harpolithobius folkmanovae 237, 239
Harpolithobius gottscheensis 638
Harpolithobius halophilus 226
Harpolithobius hemusi 239
Harpolithobius intermedius 357
Harpolithobius oltenicus 357
Harpolithobius radui 239, 357
Harpolithobius triacanthos 357
Harpolithobius tridentatus 357
Hedinobius hummeli 91
Hedinomorpha biramipedicula 86
Hedinomorpha hummeli 86
Hedinomorpha hummeli svenhedini 86
Helicorthomorpha hoist ii 86
Helicorthomorpha kosingai 86
Helicorthomorpha ocellata 86
Helicorthomorpha ortho gona 86
Helicorthomorpha uncinata 86
Helvetiosoma arvernum 276
Henia angelovi 236, 237, 238
Henia alhenarum 601. 603
Henia bicarinata 238, 600, 601
Henia devia 601
Henia hirsuta 600, 601
Henia illyrica 238, 601
Henia vesuviana 659

Henicopidae 91, 226, 303, 304, 305, 341, 346

Henicops 300

Hessebius 226. 232

Hessebius barbipes 225

Hessebius multicalcaratus 266, 267, 268

Heteroiulus 318

Heteroiulus intermedius 318, 326
Heteroporatia bosniensis 211. 212
Heterostoma 442
Hexamerocerata 285, 286
Himalodesmus 169, 173
Himalodesmus audax 169
Himalodesmus benefactor 169
Himalodesmus faustus 169
Himalodesmus parvus 169
Himalodesmus prosperus 169
Himalodesmus pulcher 169
Himalodesmus pygmaeus 169

SYSTEMATIC INDEX

671

Himalodesmus vigens 169
Himalomorpha monligena 76
Himantariidae 88

Himantarium gabrielis 238, 600, 601, 603, 659

Himantogonus rufocinctus 75

Hindomorpha granulifera 76

Hingstonia 168, 169, 171. 172, 173

Hingstonici beatae 168

Hingstonia dorjulana 168

Hingstonia eremita 168

Hingstonia falcata 168

Hingstonia fittkaui 168

Hingstonia gogonana 168

Hingstonia pahakholana 168

Hingstonia pelelana 168

Hingstonia perarmata 168

Hingstonia serrata 169

Hingstonia sympatrica 169

Hingstonia variata 169

Hirtodrepanum 169

Hirtodrepanum latigonopum 169

Hirudisoma latum 276

Hirudisoma pyrenaewn 190, 193. 199, 276

Hispaniosoma racovitzai 271, 276

Hyleoglomeris 166, 172, 173

Hyleoglomeris crassipes 166

Hyleoglomeris electa 166

Hyleoglomeris emargitiaia 84

Hyleoglomeris gorkhalis 167

Hyleoglomeris khumbua 167

Hyleoglomeris modesta 167

Hyleoglomeris nagarjunga 167

Hyleoglomeris sinensis 84

Hyleoglomeris tinjurana 167

Hyleoglomeris venustula 167

Hylomini 87

Hypnosoma exor natum 276
Hypnosoma juberthieorum 276
Hypnosoma pallidum 276
Hypsoiulus alpivagus 278, 326

Iberoiulus sarensis 278
Indosphaera 166
Indosphaera curiosa 1 66
lulidesmus 74

lulogona lirolensis cisalpinum 276
lulus 26, 27

lulus Americanus 25, 31
lulus tonginus 39
lulus peii 84
lulus vallicola 39

Janetschekella valesiaca 276
Japanoiulus lobatus 326

Jonespeltis splendidus 74, 561, 562, 563, 564, 619
Julidae 32, 35, 36, 85, 167, 173. 313, 314. 323, 325,
326, 328, 373, 649
Julidea 272, 278, 537
Julini 315, 321, 326
Juloidea 323
Julus 32, 314, 321, 324
Julus colchicus 326
Julus ghiljarovi 326
Julus jedryczkowskii 326

Julus scandinavius 245, 246, 248, 252. 258. 260, 261,
263, 278, 326, 488, 517
Julus scanicus 321
Julus subalpinus 326
Julus terrestris 211,212
Junceustreptus brevispinus 85
Junceuslreptus browning i 85
Junceustreptus prominulus 85

Karpatqphyllon 61, 63, 64, 65
Karpatophyllon banaticum T3 1 , 64
Karpatophyllon carpaticum 61, 64
Karpatophyllon dacicum 61, 63. 64
Karpatophyllon polinskii 61. 63, 64
Karteroiulus alaskanus 326
Karteroiulus niger 85
Kaschmiriosoma 170
Kaschmiriosoma contortipes 75, 76, 170
Kaschmiriosoma nulla 170
Kaschmiriosoma pleuroptera 170
Kashmireuma 167, 172
Kashmireuma nepalensis 167
Kashmireuma nielseni 167
Kashmireuma schawalleri 167
Kashmireumatidae 167, 172
Kiulinga jeekeli 88
Kiulinga lobosa 88
Kiusiunum 1 1 1

K i us i unum nodulosum 102, 108
Kiusiunum sekii 102
Kochliopus trivittatus 86
Kophosphaera 166
Kophosphaera brevilamina 166
Kophosphaera devolve ns 166
Kophosphaera excavata 166
Kophosphaera excavata mammifera 166
Kophosphaera politissima 166
Kronopeltis occidentalis 76
Kronopolites 170

Kronopoliles acuminatus biagrilectus 86
Kronopolites formosanus 86
Kronopolites occidentalis 170
Kronopoliles ralphi 86
Kronopolites spiniger 75
Kronopolites svenhedini 86
Kronopolites swinhoei 86
Kronopolites unicolor 75
Kryphioiulus occultus 211. 326. 504, 505

672

SYSTEMATIC INDEX

Kylindogaster 1 1 1
Kylindogaster nodulosa 102, 109

Lamyctes 225, 226, 232

Lamyctes coeculus 226. 227, 232

Lamyctes fulvicornis 215, 219, 227, 240, 266

Lamyctes gracilipes 91

Later ogonopus simplex 76

Leptoiulini 315

Leptoiulus 247, 254, 314

Leptoiulus alemannicus 326

Leptoiulus arelatus 278

Leptoiulus beigicus 245, 247. 248, 258. 278, 326, 647,
649. 651, 652

Leptoiulus bertkaui 258. 278
Leptoiulus brevivelatus 278
Leptoiulus broelemanni 326
Leptoiulus bruyanti 278
Leptoiulus cibdellus 326
Leptoiulus demange i 278
Leptoiulus disparatus 321. 326
Leptoiulus garumnicus 278
Leptoiulus juvenilis 278

Leptoiulus kervillei 245, 247. 248, 252, 278. 488

Leptoiulus legeri 278

Leptoiulus macedonicus 326

Leptoiulus marcomannius 159

Leptoiulus meridionalis 278

Leptoiulus montivagus 278

Leptoiulus odieri 278

Leptoiulus piceus 278

Leptoiulus proximus 211. 212, 249, 326, 431, 432, 433,
434, 478, 479
Leptoiulus remyi 278

Leptoiulus saltuvagus 451, 452, 453. 454, 456, 459

Leptoiulus simplex 488

Leptoiulus simplex glacial is 258, 260. 278

Leptoiulus tanymorphus 326

Leptoiulus umbratilis 190, 193, 195, 201, 203, 278

Leptoiulus uncinatus 278

Leucogeorgiini 173, 318, 326

Levizonus takakuwai 102, 106

Listrocheiritium 154, 156, 158

Listrocheiritium cervinum 152, 156

Listrocheiritium nibelungiacum 1 56

Listrocheiritium noricum 152, 156

Listrocheiritium septenlrionale 152, 156, 158

Lithobiidae 32. 90, 227, 295, 303, 304, 341, 346. 660

Lithobiinae 90

Lithobius 27, 28, 180, 226, 232, 240, 267, 351, 353,
354, 360 366, 386, 393, 381, 416, 418, 422, 423,
424, 429, 639

Lithobius acuminatus 658, 659
Lithobius aeruginosus 239
Lithobius aeruginosus mongolicus 90
Lithobius agilis 239, 601, 639. 640
Lithobius agilis pannonicus 357

Lithobius allotyphlus 357
Lithobius ankarensis praecursor 227
Lithobius anornatus 90
Lithobius asperatus 91
Lithobius audax 239
Lithobius aulacopus 357
Lithobius balcanicus 239
Lithobius beroni 237, 239
Lithobius beschkovi 239, 601, 604
Lithobius bidivisa 90
Lithobius bifidus 239
Lithobius bogdoulensis 90
Lithobius borealis 190, 191. 266, 357
Lithobius borisi 236
Lithobius brignolii 601, 602
Lithobius bulgaricus 239, 357
Lithobius burzenlandicus 239
Lithobius calcaratus 190, 352, 357, 658, 659
Lithobius carinatus 225, 232, 601
Lithobius castaneus 180, 190, 357, 638. 642, 658. 659
Lithobius catascaphius 239
Lithobius cavernicola 357
Lithobius christovici 239
Lithobius coeculus 226, 227, 232
Lithobius crassipes 180, 215, 219, 222, 223, 239, 352,
366, 601, 602

Lithobius cronebergii 266, 267, 268
Lithobius crypticola 357
Lithobius curtipes 222. 223, 239, 352
Lithobius cyrtopus 266, 267, 357
Lithobius dalmaticus 239
Lithobius decapolitus 357
Lithobius decessus 90
Lithobius dentatus 638, 639, 644, 645
Lithobius diampolisi 239
Lithobius dobrogicus 239
Lithobius duboscqui 190, 239, 266
Lithobius electron 236, 239
Lithobius erratus 90

Lithobius erythrocephalus 222, 223, 226, 232, 236, 239,
266, 267, 352, 357, 601, 602
Lithobius ery throcephalus borisi 604
Lithobius fagniezi 357
Lithobius fascial us 225

Lithobius forficatus 22. 180, 190, 213, 215, 216, 221,

222, 223, 239, 266, 267, 268, 305, 352, 355, 357,

366, 369, 385, 391, 392, 393, 403, 404, 405, 407.

411, 415, 421. 422, 424, 638, 642, 643, 644, 645.

657, 658, 659, 660, 661

Lithobius gantoensis 90
Lithobius glaciei 236, 239
Lithobius golemanskyi 237, 239
Lithobius hauseri 600, 601
Lithobius hummeli 90
Lithobius inermis 206, 207, 208, 357
Lithobius irregularis 90
Lithobius jangsteanus 90
Lithobius jurinici 236, 239
Lithobius kansuanus 90
Lithobius kiayiensis 90
Lithobius lakatnicensis 239, 601, 604

Source : MNHN. Paris

SYSTEMATIC INDEX

673

Lithobius lapidicola 239, 266. 357, 601, 602, 638. 639,
641, 642, 644, 657 658, 659
Lithobius latro 239, 357, 601, 638, 639, 642, 643
Lithobius litoralis 227

Lithobius lucifugus 239, 266, 267, 357, 601, 602
Lithobius lusitanus 206, 207, 208, 644, 645
Lithobius lusitanus tataricus 266, 267. 268
Lithobius luteus 357

Lithobius macilentus 638, 639, 640, 641, 642. 643, 644,
645

Lithobius macrops 225
Lithobius maculipes 236, 239
Lithobius matici matici 357

Lithobius melanops 215. 219. 222, 223, 266. 267, 352,
357, 640, 642

Lithobius microps 190. 213, 215, 216, 219, 221, 222,
239, 266, 267, 268, 352 357, 601. 602
Lithobius mongolellus 90
Lithobius mongolicus 90
Lithobius mongolomedius 90

Lithobius mutabilis 215, 219, 239, 240, 266, 357, 601,
602

Lithobius muticus 239, 357. 601
Lithobius nicoeensis 357
Lithobius nigrifrons 239, 357
Lithobius nigripalpis 239, 601, 602
Lithobius tiodulipes 357. 638, 639, 641, 645
Lithobius oglednicus 239
Lithobius ongi 90

Lithobius parietum 239, 266, 267, 357

Lithobius parvicornis 225, 232

Lithobius pelidnus 239, 266, 267, 357

Lithobius peregrinus 237, 239, 357, 601, 602

Lithobius piceus 190, 239, 266. 267, 357

Lithobius piceus gracilitarsis 360, 361

Lithobius pilicornis 190, 206, 207. 208, 357, 359. 360

Lithobius popovi 239

Lithobius praecursor 221

Lithobius proximus 239, 266, 267, 268

Lithobius punctulatus vasconicus 357

Lithobius pusillus 239

Lithobius pustulatus 239

Lithobius pygmaeus 640, 642

Lithobius ribauti 357

Lithobius romanus 659

Lithobius rufus 90

Lithobius rugosus 91

Lithobius ruschovensis 239

Lithobius rylaicus 236, 239

Lithobius schuleri 601, 602

Lithobius silvivagus 357

Lithobius speluncarum 357

Lithobius strandzanicus 237, 239

Lithobius sulcifemoralis 90

Lithobius tenebrosus 266, 267. 352, 601, 603, 643

Lithobius tethidis 90

Lithobius letrophthalmus 90

Lithobius thracicus 236, 239

Lithobius tiasnatensis 236, 239

Lithobius totevi 240

Lithobius trebinjanus 240

Lithobius trichopus 90
Lithobius tricuspis 190, 240, 357, 641
Lithobius troglodytes scutigeropsis 357
Lithobius tylopus 658, 659
Lithobius typhlus 357
Lithobius uni unguis 240
Lithobius validus 266, 645
Lithobius variegatus 357

Lithobius variegatus rubriceps 190, 191, 206, 207, 208

Lithobius viriatus 226, 232, 240, 266, 267, 601, 602

Lithobius vizicae 240

Lithobius vosseleri 225

Lithobius zelazovae 240

Lithobius (Dacolithobius) domogledicus 357

Lithobius (Monotarsobius) aeruginosus 357

Lithobius (Monotarsobius) baloghi 357

Lithobius (Monotarsobius) biunguiculatus 357

Lithobius ( Monotarsobius ) burzenlandicus 357

Lithobius (Monotarsobius) crassipes 357

Lithobius (Monotarsobius) curtipes 357

Lithobius (Monotarsobius) dobrogicus 357

Lithobius (Monotarsobius) duboscqui 357

Lithobius (Monotarsobius) dudichi 357

Lithobius (Monotarsobius) microps 357

Lithobius (Monotarsobius) pustulatus 357

Lithobius (Monotarsobius) sciticus 357

Lithobius (Monotarsobius) subterraneus 357

Lithobius ( Polybothrus) fasciatus graecus var.

fasciatograecus 227
Lithobius (Thracolithobius) dacicus 357
Lithobius ( Thracolithobius ) inexpectatus 357
Loboglomeris haasi 190. 193
Loboglomeris pyrenaica 275
Loboglomeris rugifera 275
Lophoproctidae 84, 123
Lophoproctinus inferus 275
Lophoproctinus inferus fnaurus 123, 124, 125
Lophoproctus 133
Lophoproctus jeanneli 275
Lophoproctus lucidus 275
Lophoproctus madecassus 125
Lophoturus 127, 128, 134
Lophoturus aequatus 127. 128, 133, 134
Lophoturus madecassus 113, 123, 124, 127, 133
Lophoturus niveus 133, 134
Lophoturus okinawai 84

Macheiriophoron 54, 58
Macheiriophoron aelleni 53, 54, 55, 57, 58
Macheiriophoron alemannicum 53, 54, 55, 57, 58
Macheiriophoron alemannicum genuinum 54, 58
Macheiriophoron alemannicum globosum 54. 57, 58
Macheiriophoron alemannicum rotundatum 54, 58
Macheiriophoron alemannicum triarticulatum 54, 58
Macheiriophoron cervinum 53, 54, 55. 57, 58
Macheiriophoron cervinum brevidenlatum 54, 55, 58
Macheiriophoron montivagum 58
Macheiriophoron montivagum silvaticum 58

674

SYSTEMATIC INDEX

Macheiriophoron serratum 54, 59
Macheiriophoron silvaticum 54, 58
Macheiriophoron silvaticum hessei 54
Macheiriophoron verhoeffi 54, 59
Macheiriophoron verhoeffi excavation 55, 59
Macheiriophoron verhoeffi genuinum 59
Macheiriophoron wehranum 54, 59
Macheiriophoron wehranum brevidentatum 55
Macheiriophoron wehranum calcivagum 55, 59
Macheiriophoron wehranum clavigerum 55. 59
Macheiriophoron wehranum genuinum 59
Macheiriophoron wehranum quadridentatum 59
Macheiroiulus compressicauda 326
Macheiroiulus libicus 319, 320, 326
Macrosternodesmidae 101
Macrosternodesmus 1 0 1

Macrosternodesmus palicola 245, 246, 247. 248. 258,
277, 488

Macroxenodes 121, 127. 128, 133

Macroxenodes bartschi 121. 133

Macroxenus 120, 121

Macroxenus caingangensis 121, 125

Macroxenus enghoffi 113, 117. 118, 121, 123, 125

Macroxenus rubromarginatus 121, 125

Magidesmus 169

Magidesmus affinis 169

Magidesmus bhutanensis 169

Mandarinopus gracilipes 87

Marquetiella auriculata 276

Marquetiella lunata 276

Marquetiella lunation 190, 193, 196, 199. 201

Marquetiella pyrenaica 276

Martensodesmus 169

Martensodesmus bicuspidatus 169

Martensodesmus excornis 169

Martensodesmus himalayensis 169

Martensodesmus nagarjungicus 1 69

Martensodesmus sherpa 169

Martensosoma 170

Martensosoma elegans 170

Martensosoma foveatum 170

Martensosoma schawalleri 170

Martensosoma silvestre 170

Martensosoma splendens 170

Martensosoma unicolor 170

Mastigona bosniense 328

Mastigonodesmus destefani 277

Mastigonodesmus fagniezi 271, 277

Mastigonodesmus lopezi 271, 277

Mastigophorophyllidae 62, 328

Mast igophorophy lion 61, 62, 63, 64, 65

M astigophor op hy l Ion aberration 61, 62, 64

Mastigophorophyllon alpivagum 61, 62, 64

Mast igophorophy lion banarescui 61, 62

Mastigophorophyllon banaticum 64

Mastigophorophyllon bohemicum 61. 62, 64

Mastigophorophyllon bulgaricum 61, 62, 64

Mastigophorophyllon bulgaricum pirinicum 61, 62, 64

Mastigophorophyllon carpaticum 61, 62, 63, 64

Mastigophorophyllon cirriferum 61. 62, 64

Mastigophorophyllon crinitum 61, 62, 64

Mastigophorophyllon crinitum huculicum 61, 62, 64

Mastigophorophyllon deubeli 61, 62, 64

Mastigophorophyllon giliarovi 63

Mastigophorophyllon jickelii 61, 62, 64

Mastigophorophyllon penicilligerum 61. 62, 64

Mastigophorophyllon saxonicum 61, 62, 63, 64

Mastigophorophyllon serrulatum 61, 62, 63, 64

Mastigophorophyllon serrulatum apiculatum 61, 62, 64

Mastigophorophyllon transsilvanicum 61, 62. 64

Mecistocephalidae 78

Mecistocephalinae 88

Mecistocephalus brevisternalis 89

Mecistocephalus diversisternus 89

Mecistocephalus fenestratus 89

Mecistocephalus indecorus 89

Mecistocephalus insularis 89

Mecistocephalus insulomontanus 89

Mecistocephalus japonicus 89

Mecistocephalus mikado 88

Mecistocephalus mirandus 89

Mecistocephalus monticolens 89

Mecistocephalus multidentatus 89

Mecistocephalus nannocornis 89

Mecistocephalus ongi 89

Mecistocephalus punctifrons 89

Mecistocephalus rubriceps 88

Mecistocephalus smithi 89

Mecistocephalus takakuwai 89

Megalotylidae 167, 172

Megaphyllum 314, 317, 318, 325

Megaphyllum adanense 317, 326

Megaphyllum bosniense 326

Megaphyllum brachyurum 318, 326

Megaphyllum geniculatum 318, 326

Megaphyllum hercules 317, 326

Megaphyllum projection kochi 431, 432, 433, 434

Megaphyllum rossicum 326

Megaphyllum taygeti 326

Megaphyllum tenenbaumi 326

Megaphyllum unilineatum 40

Melogona gallica 245, 248, 258, 259, 260, 262, 277,
488, 509, 512, 513, 533, 537, 541, 550
Melogona scutellare 245, 248, 277, 509, 513
Melogona voigti 509, 510, 51 1, 512, 513, 514
Mesoblaniulus serrula 278
Mesoiulus ciliciensis 315, 326
Mesomeritius indivisus 326
Mestosoma hylaeicum 524
Metaiulini 320, 324, 326

Metaiulus pratensis 245, 246, 247, 248, 249, 250, 278,
320, 326

Metamastigophorophyllon 61, 63

Microchordeuma gallicum 509, 512, 513, 541

Millotauropus silvestrii 286

Mi mops orientalis 90

Monacobates monoecensis 278

Mongoliulidae 85, 326

Monogr aphis 166

Monographis mirus 166

Monographis tamoyoensis 122

Monotarsobius 225, 226. 231, 232, 639

SYSTEMATIC INDEX

675

Monotarsobius aeruginosus 266, 267, 638, 639. 640.

641, 642, 643, 644
Monotarsobius alticus 90
Monotarsobius argaeensis 90
Monotarsobius barbipes 225
Monotarsobius bolognai 226, 231, 232
Monotarsobius crassipes 90, 226, 231, 232, 266, 267
Monotarsobius crassipes holstii 90
Monotarsobius crassus 90
Monotarsobius curlipes 266, 267, 268
Monotarsobius kaszabi 90
Monotarsobius interops 266
Monotarsobius obtusus 90
Monotarsobius ramulosus 90
Monotarsobius rhysus 90
Monotarsobius schizus 231
Monotarsobius sseliwanoffi 266, 267. 268
Monotarsobius teldanensis 225. 229, 231, 232
Moojenodesmus purnilus 524
Moojenodesmus susannae 524
Muyudesmus obliteratus 524
Mycogona germanica 277 . 329

Mycogona germanicum 258, 647, 649, 650, 652. 653,

655

Nannophilus ariadnae 600, 601
Nanogona balazuci 276
Nanogona cebennica 276
Nanogona davidi 276
Nanogona digitata 276

Nanogona polydesmoides 245, 246. 248, 249, 276
Nanogona uncinata 276
Narceus annularis 5 1 5

Necrophloeophagus flavus 213, 215, 221, 222, 223
Nedyopodini 87
Nedyopus patriolicus 87
Nedyopus tambanus 102, 105

Nemasoma varicorne 210, 211, 212, 245, 246, 248, 250.

258, 278, 326, 328, 488, 521
Nemasomatidae 85, 326, 328
Nepalella 167, 172, 173
Nepalella deharvengi 167, 168
Nepalella gairiensis 167
Nepalella gunsa 1 67
Nepalella jaljalae 167
Nepalella khumbua 167
Nepalella phulcokia 167
Nepalella ringmoensis 167
Nepalella taplejunga 167
Nepalella thodunga 167
Nepalella tragsindola 167
Nepalmatoiulus 35, 168. 172, 173, 318, 326
Nepalmatoiulus appendiculatus 1 68
Nepalmatoiulus binnanicus 318, 326
Nepalmatoiulus brachymeritus 86
Nepalmatoiulus deharvengi 168
Nepalmatoiulus dhaulagiri 168
Nepalmatoiulus eulobos 86

Nepalmatoiulus fraterdraconis 86
Nepalmatoiulus generalis 168
Nepalmatoiulus hyalilobus 168
Nepalmatoiulus ivanloebli 168, 172
Nepalmatoiulus juxtapositus 168
Nepalmatoiulus martensi 168
Nepalmatoiulus mauriesi 168
Nepalmatoiulus nigrescens 168
Nepalmatoiulus pineti 168
Nepalmatoiulus polyakis 86
Nepalmatoiulus rhaphimerilus 85
Nepalmatoiulus rugiflagrum 168
Nepalmatoiulus smetanai 168
Nepalmatoiulus sympatricus 168
Nepalmatoiulus tibetanus 85
Nepalmatoiulus uncus 168
Nepalmatoiulus wuermlii 168
Nepalmatoiulus yunnanensis 86
Nepalmatoiulus zachonoides 168
Nepalomorpha 170
Nepalomorpha arunensis 170
Nepalomorpha hirsuta 170
Nepalomorpha kuznetsovi 170
Nepalomorpha spinigera 170
Nesoporogaster hispanica 190
Newportia 294. 295
N ip o nia 1 1 1

Niponia nodulosa 87. 102, 108
Niponia simplexus 88
Nodocephalus dooi 89
Nodocephalus edentulus 89
Nodocephalus pauroporus 89
Nopoiulus kochii 244, 245. 247, 248, 278
Nopoiulus minutus 244
Nopoiulus venustus 211, 244

Occitaniulus rouchi 278

Occitanocookia hirsuta 271, 111

Ochogona caroli 451, 452, 453, 454, 457, 459

Ochogona gallitarum 276

Odontopygidae 332, 333, 337. 572, 574. 621

Okeanobates serratus 326

Okeanobatidae 326

Ommatoiulus 314, 322. 324, 325, 373, 374, 592

Ommatoiulus albolinealus 278

Ommatoiulus apunctulatus 588

Ommatoiulus cingulatus 326

Ommatoiulus corsicus 27 8

Ommatoiulus gaulhieri 180, 182

Ommatoiulus haackeri 278

Ommatoiulus illicis 278

Ommatoiulus imminutus 278

Ommatoiulus kessleri 322, 326

Ommatoiulus lapidarius 326

Ommatoiulus lienhardti 278

Ommatoiulus moreleti 322, 326, 375, 470. 517, 588

Ommatoiulus navasi 322. 326

Ommatoiulus nivalis 326

676

SYSTEMATIC INDEX

Ommatoiulus osellai 326
Ommatoiulus oxypygus 326
Ommatoiulus robustus 190. 193, 199, 201, 203
Ommatoiulus rutilans 258, 259, 260. 262, 278. 322. 326,
628

Ommatoiulus sabulosus 190, 193, 201, 245, 246, 248.

250, 254, 258. 259. 260, 261. 278. 326. 373, 374,

375. 376. 377. 380. 381, 382. 431. 432, 433, 434.

537, 550. 587. 588. 589, 590. 592, 593. 595, 596,

597, 628, 647, 649, 651, 652, 654

Ommatoiulus sabulosus aimatopodus 588
Ommatoiulus sabulosus punctatus 576
Onciurosoma adisi 524
Oncoiulini 318, 319, 326
Oncoiulus 314

Onychoglomeris castanea 275

Onychophora 39.139, 141, 142, 143, 148. 149, 483,
493, 494

Ooperipatellus 141. 142, 143, 144. 145. 146, 147, 148
Ooperipatellus insigms 139, 141. 142, 147, 148
Ooperipatellus nanus 143
Ooperipatellus viridimaculatus 147. 148
Ooperipatus 140, 143

Ooperipatus decoratus 140, 144, 145, 146, 147, 148

Ooperipatus insignis 140

Ooperipatus oviparus 140, 145, 146

Ophiodesmus albonanus 211, 245, 246, 247, 248, 277

Ophyiulus 321

Ophyiulus bastiensis 278

Ophyiulus chilopogon 278, 326

Ophyiulus corsicus 278

Ophyiulus fallax 210, 21 1, 212

Ophyiulus germanicus 326

Ophyiulus major 321. 326

Ophyiulus napolitanus 278

Ophyiulus pilosus 245, 246, 248, 249, 252, 278, 326.

398, 478, 479, 564
Ophyiulus renosensis 278
Ophyiulus targionii 326
Opisotretidae 102, 110, 111, 169, 171
Opisthocheiron canayerensis 271, 274. 276
Opisthocheiron cornutum 276
Opisthocheiron elegans 276, 627, 628, 629, 630
Opisthocheiron fallax 276
Opisthocheiron lacazei 276
Opisthocheiron penicillatum 276
Origmatogona kimeorum 271, 276
Orinisobates gracilis 85
Orinisobates 326
Orophini 88

Orophosoma 170, 171, 172

Orophosoma fechteri 1 70

Orophosoma hingstoni 86, 170

Orophosoma simulans 86, 170

Orphnaeus brevilabiatus 88

Orthochordeumella fulva 159, 277

Orthochordeumella leclerci 27 1 , 277

Orthochordeumella pallida 258, 259, 260, 262, 277, 488

Orthochordeumella pyrenaica 211

Orthomorpha 75, 170

Orthomorpha afftnis 87

Orthomorpha almorensis 75, 170

Orthomorpha bisulcata 87

Orthomorpha circularis 87

Orthomorpha coarctata 86, 102, 105. 170

Orthomorpha coonoorensis 75

Orthomorpha corticina 87

Orthomorpha dentata 75

Orthomorpha endeusa 87

Orthomorpha flavomarginata 87

Orthomorpha nordenskjoeldi 87

Orthomorpha penicillata 87

Orthomorpha ( Kalorthomorpha) almorensis 74

Orthomorpha ( Kalorthomorpha ) coonoorensis 74

Orthomorpha ( Kalorthomorpha) dentata 74

Orthomorpha (Kalorthomorpha) Ursula 74

Orthoporoides 333, 334

Orthoporoides pyrocepltalus 334, 336, 337

Orthoporus 623

Orthoporus ornatus 397. 470. 471
Orthoporus pyrocepltalus 333
Oryidae 88. 3*01
Otocry ptops 295, 442
Otocryptops rubiginosa 90
Otocryptops sexspinosus 90
Otostigminae 89, 442. 447
Otosligmus 294, 295
Otostigmus aculeatus 89
Otosligmus astenus 89
Otostigmus frigidus 89
Otostigmus insular is 89
Otostigmus malayanus 89
Otosligmus multispinosus 89
Otostigmus politus 89
Otostigmus politus mandschurius 89
Otostigmus politus pigmentatus 89
Otostigmus scaber 89
Otostigmus striatus 89
Oxidus 170
Oxidus corcifera 87

Oxidus gracilis 86, 102, 105, 170, 211, 247, 255, 258,
277, 431, 432, 433. 434
Oxydesmidae 101

Pachydesmus attemsi 88
Pachyiulini 313, 315, 324, 326
Pachyiulus 313, 314. 315. 596
Pachyiulus flavipes 326
Pachyiulus fuscipes 588, 589, 592
Pachyiulus varius 278
Pachymerinae 88
Pachymerium atticum 88

Pachymerium ferrugineum 88, 206, 207, 222, 223, 238,
601, 602, 659

Pachymerium ferrugineum insulanum 236
Pachymerium flavum 238
Pachypodoiulus eurypus 326
Pacidesmus sinensis 87
Pacifiiulus imbricatus 326

SYSTEMATIC INDEX

677

Paectophyllini 313, 315, 319, 320, 324, 326
Paectophyllum escherichii 320, 326
Paeromopodidae 326
Paeromopodoidea 326
Pamelaphe lacustris 88
Parabilingulus aramulus 85
Parachordeuma broelemanni 277
Paracortina (Ahum) carinata 86
Paracortina (Allum) serrata 86
Paracortina (Ahum) viriosa 86
Paracortina ( Relictus ) stimula 86
Paracortina ( Relictus ) thallina 86
Paracortina 309

Paracortina leptoclada 86, 307, 308, 309, 310
Paracortina voluta 86, 307, 308, 309. 310
Paracortinidae 86, 307, 308
Par aery ptops 294, 295
Par adoxo soma 73

Paradoxosomatidae 73. 74. 86, 87, 102, 104, 105. 110,
169, 171, 173. 524
Paradoxosomatidea 86, 277
Paradoxosomaiinae 74. 86
Paradoxosomatini 173
Paradoxosomatoidea 86
Parafontaria ishiii 102

Parafontaria laminata armigera 102, 103. 106
Paraiulidae 85, 326
Paramastigophorophyllon 62
Paranedyopus 170, 173
Paranedyopus affinis 170
Paranedyopus cylindricus 76, 170
Paranedyopus elongissimus 76, 170
Paranedyopus martensi 1 70
Paranedyopus rufocinctus 75
Paranedyopus schawalleri 170
Paranedyopus similis 170
Paranedyopus simplex 75
Paranedyopus subcylindricus 75
Paranedyopus Ursula 75
Parapachyiulus recessus 326
Parapauroplus monodentus 87
Paraplanes svenhedini 88
Parchondromorpha 75
Parchondromorpha coonoorensis 75
Parchondromorpha indica 76
Parchondromorpha similis 76
Parfusiulus 39
Parorthomorpha 1 70
Parorthomorpha affinis 170
Parorthomorpha granulosa 1 70
Parorthomorpha intermedia 1 70
Parorthomorpha longiseta 170
Parorthomorpha nyakensis 1 70
Parorthomorpha philosophica 170
Parorthomorpha spectabilis 170
Parorthomorpha tergalis 170
Parorthomorpha tuberculala 170
Pauropodidae 9 1
Pauropsxenus vilhenae 122
Pauropus bifurcus 9 1
Pauropus longiramus 9 1

Pectiniunguis 79

Pellopodoiulus scliestoperovi 326
Penicillata 84, 113, 127
Perapolydesmus progressus 277
Pericambalidae 85
Pcripatidae 493, 494
Peripaioides leuckarti 140
Peripatopsidae 139. 140, 141, 142
Peripatus 140

Peripatus insignis 140, 143
Phalloiulus dislinctus 180
Phyletodesmus 74
Physobolidae 168. 171
Phy sobol us 168
Physobolus olivaceus 168
Platydesmidea 276

Pleurogeophilus mediterraneus 601, 602
Pleurogeophilus takakuwai 88
Pleurolithobius jonicus 237, 240
Pleurolithobius orientis 601
Pleurolithobius patriarchalis 600, 601, 602
Plutonium 294, 295, 296, 297, 301. 441
Plutonium zwierlainii 296
Pocillidorsus dorsiangulatns 87
Podoglyphiulus 168
Podoglyphiulus elegans nepalensis 168
Polybothridae 90

Poly hot hr us fasciatus 225, 226, 227
Polydesmidae 32, 73, 87, 101, 102, 106, 117, 111, 169,
173, 650

Polydesmidea 87, 277, 537
Polydesmoidea 87
Polydesmus 173. 244. 480, 481

Polydesmus august us 245, 246. 247, 248, 249. 258, 260,
262, 277. 488,533, 537, 542, 543, 547, 550
Polydesmus asthenestatus 277
Polydesmus barbierii 211

Polydesmus complanatus 211, 212, 249, 250, 431, 432.
433, 434

Polydesmus coriaceus 250, 277

Polydesmus coriaceus coriaceus 190, 193, 196. 199

Polydesmus cor sic us 211

Polydesmus denticulatus 245, 246, 248. 250, 254, 258,
260, 262. 277 . 477, 478, 479, 480, 481, 488, 489
Polydesmus gallicus 245, 246. 248
Polydesmus german icus 211
Polydesmus hamatus 87, 277
Polydesmus helveticus 159, 277
Polydesmus incisus 211

Polydesmus inconstans 211, 212, 245, 246, 248, 258.

260. 277, 647, 649, 650. 651. 652, 654
Polydesmus japonicus 102, 107
Polydesmus liber 87
Polydesmus mistrei 211
Polydesmus moorei 88
Polydesmus niveus 211
Polydesmus paludicola 88
Polydesmus plicatus 211
Polydesmus racovitzai 190, 193, 277
Polydesmus raffardi 211
Polydesmus rothi 154, 160

Source :

678

SYSTEMATIC INDEX

Polydesmus superus 280
Polydesmus laranus 277

Polydesmus testaceus 245, 246, 247, 248, 250. 258, 259,
260, 262, 277
Polydesmus troglobius 277
Polydesmus xanthocrepis 154, 157, 160
Polydrepanini 74, 75. 86
Polydrepanum 75, 76
Polydrepcmum horridum 76
Polydrepanum implied turn 75, 76
Polydrepanum tamilum 75
Polylobosoma roseipes 87
Polypauropidae 91
Polyxenes (sic!) 127
Polyxenida 84, 166, 275
Polyxcnidae 32, 84. 114, 123, 166
Polyxenidea 275

Polyxenus 117, 127, 128. 132, 133. 166, 285, 287

Polyxenus anacapensis 127, 132, 133

Polyxenus bartschi 127. 128. 133

Polyxenus chalcidicus 114, 116. 117, 125

Polyxenus fasciculalus 113, 114, 123, 124. 125, 127,

128, 129, 130, 131
Polyxenus fasciculalus pallidus 128
Polyxenus fasciculatus vicloriensis 128
Polyxenus hangzoensis 84, 133
Polyxenus koreanus 133

Polyxenus lagurus 113, 114, 116, 123, 124, 125, 127,

129, 130, 131, 245, 246, 247. 248. 250, 258, 275,
286, 628, 629, 630

Polyxenus lapidicola 1 1 7

Polyxenus macedonicus 117, 275

Polyxenus oromii 113, 114, 116, 117, 123, 125

Polyxenus precilus 133

Polyxenus pugetensis 127, 129, 130, 131

Polyxenus shinoharai 133

Polyxenus luberculatus 127, 128

Polyzonidae 32

Polyzoniidea 276

Polyzonium germanicum 211, 245, 246,248, 250. 252,
254, 276, 461. 462, 463. 465, 466, 469. 470, 471,
627, 628, 629, 631, 632, 633
Prionomalis 102, 107, 111
Prionopeltis planatus 76
Prolamnonyx holstii 89
Prolamnonyx sauleri 89
Prosopodesmus jacobsoni 87
Prostemmiulus adisi 524
Prostemmiulus amazonicus 524
Prostemmiulus wellingtoni 524
Proieroiulus broelemanni 278

Proteroiulus fuscus 21 1, 212, 245. 246, 248, 250. 252,
254, 258, 278

Protoglomeris vasconica 190, 193, 195, 196, 199, 201,
275

Pselaphognatha 285, 286, 287
Pseudocatapyrgodesmus 1 1 1
Pseudodesmus 167
Pseudonannolenidae 524
Pseudonemasoma femorotuberculata 326
Pseudonemasomatidae 326

Pseudosphaeroparia 1 69
Pseudosphaeroparia cavernicola 1 69
Pteridoiulini 315, 317, 324, 326
Pteridoiulus 324

Pteridoiulus aspidiorum 317, 326

Pterozonium 167

Pterozonium cingulatum 167

Pterozonium coniceps 167

Pterozonium lanvoodi 167

Pterygotergidac 91

Pterygotergum svenhedini 91

Pyreneosoma barbie ri 276

Pyreneosoma bessoni 276

Pyreneosoma dig ita turn 276

Pyreneosoma ribauli 276

Pyrgocyphosoma 154

Pyrgocyphosoma dalmazzense 277

Pyrgocyphosoma doriae 277

Pyrgocyphosoma titianum 152, 154

Pyrgodesmidae 88, 102, 108, 110, 111, 170, 171, 524

Relictus 309

Relictus stimulus 307, 308, 309, 310
Relictus thallinus 307, 308, 309, 310
Rhapidostreptus 411, 413, 415, 416, 417, 418, 422,

423, 424, 429

Rhapidostreptus virgator 411, 412, 414, 417, 421, 422,

424, 434

Rhipidopeltis 102, 110, 111
Rhodesiostreptus matabele 572
Rhodopiella beroni 326
Rhopaloiulidae 323, 324, 326
Rhopaloiulus cameratanus 326

Rhymogona 45, 46, 48, 49, 50, 53, 54. 57, 58, 152,
154, 155, 156, 159
Rhymogona aelleni 49, 53, 55, 57
Rhymogona alemannica 46. 48, 49, 50. 53. 54, 55, 57,
152, 271, 277

Rhymogona alemannica alsaticum 54
Rhymogona alemannica globosum 54. 57
Rhymogona alemannica rot undatum 54
Rhymogona alemannica triarticulatum 54 .57
Rhymogona cervina 46, 47, 48, 49, 50, 53, 54, 55, 58,
152, 159, 271, 277
Rhymogona cervina brevidentatum 54
Rhymogona montivaga 46, 47, 48, 49, 50, 53, 55, 57,
58, 271, 277

Rhymogona montivaga alemannica 56, 58

Rhymogona montivaga cervina 56, 58

Rhymogona montivaga hessei 49, 55, 58

Rhymogona montivaga montivaga 49, 56, 58

Rhymogona montivaga serrata 56, 59

Rhymogona montivaga verhoeffi 56, 59

Rhymogona montivaga wehrana 56, 59

Rhymogona serrata 48, 49, 50, 53, 54, 55, 57, 59, 152

Rhymogona silvatica 53, 55. 58, 271

Rhymogona silvatica hessei 271

Rhymogona verhoeffi 48,49, 50, 53. 54. 55, 57, 59, 152

Source

SYSTEMATIC INDEX

679

Rhymogona verhoeffi excavation 54

Rhymogona wehrana 48, 49, 50, 53, 55, 57. 59. 152

Rhymogona wehrana calcivagum 54

Rhymogona wehrana clavigerum 54

Rhymogona wehrana genuinum 54

Rhymogona wehrana quadridentatum 54

Rhysida 294, 295. 441, 442, 444, 447, 448

Rhysida lithobioides 90

Rhysida longipes 90

Rhysida longipes brevicornis 90

Rhysida mandchurica 89

Rhysida nuda brevicornuta 90

Rhysida nuda immarginata 90

Rhysida nuda nuda 89

Rhysida nuda togoensis 441. 442, 443, 447, 448
Rhysida yanagiharai 90
Rhysodesmus cohaesivus 88
Rhysodesmus contiguus 88
Ribautiella 610

Ribautiella amazonica 607, 609, 610, 614

Riukiaria 102, 106

Riukiaria capaca 88

Riukiaria holstii 88

Riukiaria neptuna 88

Riukiaria ochraceus 88

Riukiaria taiwanalis 88

Riukiaria taiwanus 88

Riukiaria uraensis 88

Riukiaria variata 88

Rossiulus kessleri 515, 516, 517. 518 521

Schendyla 180, 639

Schendyla carniolensis 638, 639, 640, 641, 643, 644
Schendyla delicatula 238
Schendyla montana 238, 601, 603
Schendyla nemorensis 206, 207, 208, 213, 215, 221,
222, 223, 238. 638, 643, 644, 645, 659
Schendyla walachica 238
Schendylidae 79, 88
Schendylinae 88
Schendylurus 79
Schizophyllini 322, 324, 326
Schizophyllum 314
Schizophyllum sabulosum 516
Scolioplanes maritimus japonicus 88
Scolioplanes transsilvanicum 88
Scolopendra 26, 27. 29, 31. 32, 82, 180, 236, 294, 295,
423, 424, 441, 447, 495, 497, 498
Scolopendra Americana (sic!) 25, 31
Scolopendra calcarata 89
Scolopendra canidens 448

Scolopendra cingulata 89, 206, 207, 208, 236, 238, 366,
421, 422, 423, 429. 442, 600, 601. 603
Scolopendra cingulata thracia 236
Scolopendra dalmatica 600, 601
Scolopendra electrica 32
Scolopendra marina 27, 32
Scolopendra mazbii 83, 89

Scolopendra mirabilis 448

Scolopendra morsitans 89, 236. 441, 442. 444, 447,

448, 495, 497, 498
Scolopendra multidens 89
Scolopendra mutilans 89
Scolopendra oraniensis 429
Scolopendra phosphor ea 32
Scolopendra rapax 89
Scolopendra rugosa 89
Scolopendra seplemspinosa 89
Scolopendra subspinipes 89
Scolopendra subspinipes dehaani 89
Scolopendra subspinipes japonica 89
Scolopendra terrestris 27

Scolopendra valida 441. 442, 444, 446. 447, 448
Scolopendrellidae 137. 609. 610. 611, 614
Scolopendrellopsis 1 37

Scolopendrellopsis (Symphylellopsis) pauli 137

Scolopendrellopsis (Symphylellopsis) selgae 137

Scolopendrellopsis tropicus 614

Scolopendria marina 32

Scolopendria terrestris 32

Seolopendridae 32. 89, 294, 441

Scolopendrinae 89, 442, 447

Scolopocryptops 294, 295

Scolopocryptops brolemanni 90

Scolopocryptops ferrugineus 294

Scolopocryptopsinae 90

Scutigera 240. 300, 301,346. 437

Scutigera coleoptrata 91, 180. 206, 207, 240, 305, 437.

438, 439, 440, 600, 601
Scutigera complanata 9 1
Scutigera hispida 9 1
Scutigera longicornis clunifera 91
Scutigera sinensis 91
Scutigera sinuata 9 1
Scutigerella immaculata 9 1
Scutigerellidae 91, 612. 614
Scutigeridae 91
Scutogona jeanneli 277
Semenellogon 74
Semiosoma bardei 277
Semiosoma devillei 211
Serboiulus lucifugus 32 1
Serradium 101
Sholaphilus 169, 171, 172
Sholaphilus asceticus 169
Sholaphilus dalai 169
Sholaphilus gompa 169
Sholaphilus lama 169
Sholaphilus martensi 169
Sholaphilus monachus 169
Sibiriulus dentiger 326
Sichotanus mandschuricus 87
Sigibius 639
Sigipinius grahami 87
Sinocallipodidae 86
Sinocallipus simplicipodus 86
Sinocybe cooki 84
Sinostemmiulus simplicior 85
Siphonophora 84

680

SYSTEMATIC INDEX

Siphonophoridae 32, 84, 167. 171
Skleroprotopus confucius 85
Skleroprotopus coreanus 326
Skleroprotopus laticoxalis 85
Skleroprotopus membranipedalis 85
Skleroprotopus serratus 85
Skolopendra (sic!) 26
Spelaeoglomeris alpina 276
Spelaeoglomeris doderoi 276, 504
Spelaeoglomeris jeanneli 276
Speophilosoma 86
Speophilosomatidac 86
Sphaeropauropinae 86
Sphaeropauropus 9 1
Sphaeropoeidae 84. 166. 171
Sphaeropoeus 166
Sphaeropoeus montanus 166
Sphaerotherium 1 66
Sphaerotheriuni cinctellum 619
Sphaerotherium maculatum 166
Sphaerotherium politum 166
Sphaerotherium punctulatum 619
Sphaerotrichopidae 71
Spinotarsus 571

Spinotarsus cuspidosus 571. 573

Spinotarsus tenuis 570, 571. 573, 574

Spirobolellus tatakuwai 85

Spirobolidae 84

Spirobolus bungii 84

Spirobolus cincinnalis 85

Spirobolus exquisitus 84

Spirobolus formosae 85

Spirobolus graham i 85

Spirobolus joannesi 84

Spirobolus umbobrochus 85

Spirobolus walkeri 85

Spirostreptidae 322, 333. 334, 337, 621

Spirostrophus lanyusis 85

Stcmmiulidae 524

Stigmatogaster gracilis 600, 601

Stigmalogaster japonica 88

Stosatea italica 245, 246. 247, 248, 250, 258, 277

Streptogonopus phipsoni 75

Strigamia 639

Strigamia acuminata 190, 222. 223, 238, 638, 639, 640.
642, 643, 644, 645

Strigamia crassipes 213, 215, 219, 222, 223, 238, 641,
659

Strigamia engadina 600. 602

Strigamia transsilvanica 238. 601, 603, 638, 639, 640,
641, 642. 644
Strongylomorpha 74
Strongylosoma contort ipes 75
Slrongylosoma montigena 75
Strongylosoma nadari 87
Strongylosoma pallipes 431, 432, 433, 434
Sirongylosomatidae 73, 74
Strongylosomidae 73, 74
Strongylosominae 73

Stygioglomeris crinita 244, 245, 247, 248. 258, 275,
488, 647, 649, 650. 653

Stylodesmoidea 88

Styrioiulus pelidnus 322, 326

Substrongylosoma 1 70

Substrongylosoma distinctum 76, 170

Substrongylosoma falcatum 76, 170

Substrongylosoma montigena 170

Substrongylosoma schawalleri 1 70

Sulciferini 74, 86

Suliciferinac 73

Sundanina 74, 75

Sundanina bimontana 75

Sundanina contortipes 75

Sundanina granulifera 75, 76

Sundanina hirta 75

Sundanina jerdani 75

Sundanina laevisulcata 75

Sundanina nitens 75

Sundanina nulla 75

Sundanina pumila 75

Sundanina septentrionalis 75

Sundanina simplex 75

Sundanina trifida 75, 76

Sundanini 74

Symphylella adisi 607, 609, 611, 614
Symphylellopsis 137
Symphyoiulini 315, 324
Symphyoiulus impartitus 326
Symphyopleurium hozawai 84
Syngonopodium 154, 157
Syngonopodium aceris 152, 156
Syngonopodium cornutum 156
Syntelopodeuma gracilipes 86
Syrioiulus cf. andreevi 326
Syrioiulus continentalis 326
Szechuanella tenebra 87

Tachypodoiulus 322, 324, 325, 373, 374, 382
Tachypodoiulus albipes 190, 193, 199
Tachypodoiulus niger 190. 193, 199, 245, 246, 248
255. 258, 260, 260. 278, 326, 373, 374. 375, 376
378, 379, 380, 381, 382, 488, 489, 597, 647, 649
654

Taiwanella sculptulatus 89
Taiwanella striata 89
Taiwanella yanagiharai 89
Tasmanipatus 142, 148

Tasmanipatus anophthalmus 139. 140, 141, 148

Tasmanipatus barretti 139, 140, 141, 144, 146, 148

Tectoporini 86

Telodrepanum 76

Telodrepanum badaga 75

Telsonemasoma microps 326

Telsonemasomatidae 326

Thalassisobates litloralis 245, 247, 248, 278

Thalthvbius boteltoboensis 88

Theatops 294, 295

Thelodesmus 109, 111

Thelodesmus armatus 88, 102

SYSTEMATIC INDEX

681

Thereuonema dilatationis 9 1
Thereuonema hilgendorfi 9 1
Thereuonema mandschuria 9 1
Thereuonema luberculata 9 1
Thereuonema variata 9 1
Thereuonema viridescens 91
Thereuopoda clunifera 91
Thereuopoda nivicomes 91
Thracophilus 237

Thracophilus heroni 235, 237, 238

Thracophilus bulgaricus 235, 237, 238

Thracophilus cilicus 237

Thracophilus pachvpus 237

Tianella 167, 172,' 173

Tianella ausobskyi 167

Tianella bobanga 167

Tianella daamsae 1 67

Tianella gitanga 167

Tianella jaljalensis 167

Tianella kathmandua 167

Tianella lughla 167

Tianella mananga 167

Tianella mangsingma 167

Tianella martensi 167

Tianella smeianai 167

Tonkinodentus 294, 295

Tonkinosomatini 87

Topalodesmus 169

Topalodesmus communis 169

Topalosoma 170

Topalosoma setiferum 76, 170

Touranella 170

Touranella himalayaensis 1 70

Trachycormocephalus ko reanus 89

Trachydesmus 73

Trachyjulus 168

Trachyjulus mimus 168

Trachyjulus wilsonae 168

Trachysphaera drescoi 271 276

Trachysphaera lobata 245, 247, 248, 249, 276

Trachysphaera pyrenaica 276

Trichoblaniulidae 323,324, 326

Trichoblaniulus hirsutus 278, 326, 596

Trichoblaniulus lanuginosus 278

Trichonemasoma peloponesius 326

Trichonemasomatidae 323, 326

Trichopeltis 168, 171

Trichopeltis waisoni 168. 171

Trichopolydesmidac 71

Trichopolydesmoidea 67, 71

Trichopolydesmus 71

Trigoniophthalmus alternatus 286

Trigoniulus niger 85

Trigoniulus segmentatus 85

Trigoniulus takahasii 85

Trigoniulus lertius 85

Trogloiulus mirus 321

Tygarrup javanicus 89

Typhloblaniulus 592

Typhloblaniulus lorifer consoranensis 381
Typhloiulini 314, 315, 321, 325

Typhloiulus albanicus 321
Typhloiulus ausugi 321
Typhloiulus boldorii 321
Typhloiulus bureschi 321
Typhloiulus illyricus 321
Typhloiulus lobifer 321
Typhloiulus maximus 321
Typhloiulus montellensis 321
Typhloiulus psilonolus 321
Typhloiulus sculterorum 278
Typhloiulus serbani 321
Typhloiulus strictus 321
Typhlopygmaeosoma 169
Typhlopygmaeosoma hazehonae 169

Unciger 3 1 4

Unciger foetidus 211. 245. 247. 248, 314. 318, 319,
326. 431. 432, 433, 434, 647, 649. 650, 652, 653
Unixenus 166

Unixenus aff. broelemanni 1 20
Uriunceustreptus afemorispinus 85
Uriunceustreptus retrorsus 85
Uroblaniulus 326
Usbekodesmus 169, 171, 172
Usbekodesmus buddhis 169
Usbekodesmus occultus 169
Usbekodesmus sacer 169
Usbekodesmus theocraiicus 169
Usbekodesmus iheosophicus 169

Vandeleuma vasconicum 277
Vanhoeffeniidae 71
Varyomorpha hsientienensis 87
Varyomorpha pectinata 87
Vascoblaniulus 273
Vascoblaniulus cabidochei 274. 278
Vascosoma coiffaiti 271, 277
Vascosoma coijfaiti falsaforma 277
Vascosoma duprei 271, 277
Venezuelodesmini 67, 71
Venezuelodesmus 67, 68, 69, 70, 71
Venezuelodesmus bordoni 67, 68, 69, 70
Venezuelodesmus decui 67, 68, 69, 70
Venezuelodesmus orghidani 67, 68, 71

Xanihodesmini 74
Xanthodesmus physkon 585, 586
Xestoiulus laeiicollis 326
Xiphidiogonus 76
Xiphidiogonus dravidus 75
Xiphidiogonus hendersoni 75
Xiphidiogonus spinipleurus 75

682

SYSTEMATIC INDEX

Xylophageuma 154

Xylophageuma vomrathi 152, 154, 155, 159

Xylophageuma zschokkei 154. 277

Xystodesmidae 88. 102, 104, 106. 110

Xystodesmoidea 88

Xystodesmus 102, 106

Xystrosoma beatense 277

Xystrosoma cassagnaui 277

Xystrosoma catalonicum 277

Xystrosoma muricum 277

Xystrosoma pyrenaicum 211

Xystrosoma tectosagum 277

Yosidaiulus tuberculatus 326
Yuennanina aceratogaster 86
Yuennanina ceratogaster 86
Yuennanina petalolobodes 86

Zephronia 166
Zephronia alticola 166
Zephronia debilis 166
Zephronia densipora 166
Zephronia disparipora 166
Zephronia hainana 84
Zephronia hirta 166
Zephronia hysophila 166
Zephronia juvenis 166
Zephronia laevissima 166
Zephronia lignivora 166
Zephronia manca 166
Zephronia nigrinota 166
Zephronia profuga 84
Zephronia specularis 166
Zephronia tigrinoides 166
Zephronia tumida 166
Zinophora 337, 621
Zinop hora laminata 337
Zostcractinidae 326

BIBL DU
MUStUM
PARIS

★

SourcbSMNHN. Paris

Remerciements aux rapporteurs I acknowledgements to referees

La Redaction tient k remercier Ics experts exterieurs au Museum national d'Histoire naturelle dont les noms suivent, d'avoir bien voulu contribuer, avec les
rapporteurs de rEtablissement, a revaluation des manuscrits (1988-1996) :

The Editorial Board acknowledges with thanks the following referees who, with Museum referees, have reviewed papers submitted to the Memories du
Museum (1988-1996):

ADKISON D

Macon

U. S. A.

KIELAN-JAWOROWSKA Z.

Oslo

Norvfcge

AFZELIUS Bjdrn

Stockholm

Su&de

KILBURN R

Pietermaritzburg

Afrique du Sud

AKESSON Benil

Goteborg

Su£de

KNIGHT-JONES Phyllis

Swansea

Grande-Bretagne

AMIARD Jean-Claude

Nantes

France

KNIGHT-JONES Wyn

Swansea

Grande-Bretagne

ANDRES H.

Hambourg

Allemagne

KOHN A

Seattle

USA

BABA K

Kumamoto

Japon

KRANTZ G. W

Corvallis

U. S. A.

BACHELET Guy

Arcachon

France

KUDENOV Jerry D.

Alaska

U S. A.

BAUD C.A.

Gen£ve

Suisse

LAGARDfcRF.J -P.

La Rochelle

France

BELLAN Gerard

Marseille

France

LANA Paulo Da Cunha

Parana

Bresil

BEN-ELIAHU Nechama

Jerusalem

Israel

LA U BIER Lucien

Paris

France

BERGGREN M

Fiskebackskil

Sudde

LAVERDE-CASTILLO J. J. A

Bogota

Colombie

BERNET-ROLLANDE M. C

Puteaux

France

LETENDRE L.

Courbevoie

France

BERNOT L.

Anthony

France

LEG AY J. M.

Villeurbanne

France

BHAUD Michel

Banyuls-sur-Mer

France

LEVIN Lisa A.

La Jolla

U. S. A.

BLAKE James A.

Woods-Hole

U. S. A.

MACKIE Andrew

Cardiff

Grande-Bretagne

BOSS K.

Harvard

U. S. A.

MACPHERSON E.

Barcelona

Espagne

BOURDON R

Roscoff

France

MANNING R

Washington

U. S A

BOURLlfcRE F.

Paris

France

MARSHALL B.

Wellington

Nouvelle-Z61ande

BOUROULLEC J.

Pau

France

MAUCHLINEJ.

Oban

Grande-Bretagne

BRESSON F

Paris

France

MAURER Don

Long Beach

U. S. A.

BROSSET A

Paris

France

MAXWELL P

Waimate

Nouvelle-Zelande

BURKE Robert D

Victoria

Canada

MC ALPINE J. F

Ottawa

Canada

BUTLER P. M.

Surrey

Grande Bretagne

MCKENNA M

New York

U. S. A.

BUTMAN Cheryl Ann

Woods-Hole

U. S. A.

MCLAUGHLIN P

Washington

U. S. A.

CALDE D

Toronto

Canada

MEISTRICH Marvin L.

Houston

U. S A

CARR1CK Frank

Brisbane

Australie

METTAM Chris

Cardiff

Grande-Bretagne

CASTELLI Alberto

Modena

Italic

MUIR Alexander Ian

Londres

Grande-Bretagne

CHACE F. A

Washington

U. S. A.

NAGEL P

Saarbriicken

Allemagne

CHAREST P.

Quebec

Canada

NEWMAN W A

San Diego

U. S. A

CLARK P.

Londres

Grande-Bretagne

NOEL R.

Pau

France

COAN E.

Palo Alto

U. S. A.

OLIVA Rafael

Barcelona

Espagne

COMBES C.

Perpignan

France

OLIVE Peter James William

Tyne

Grande-Bretagne

CORNELIUS P.

Londres

Grande-Bretagne

PATERSON Gordon L. J.

Londres

Grande-Bretagne

CORNUDELLA Lluis

Barcelona

Espagne

PATTERSON C

Loiidres

Grande-Bretagne

CUZIN-ROUDY J

Villefranche-sur-Mer

France

PAXTON Hannelore

North Ryde

Australie

DAVIE P.

Brisbane

Australie

PEREZ FARE ANTE I

Washington

U. S. A.

DE BROYER C

Bruxelles

Belgique

PERKINS Thomas H.

Saint Petersburg

U. S. A

DESBRUYfcRES Daniel

Brest

France

PERTHUISOT J P

Nantes

France

DHAINAUT Andre

Villeneuve d'Ascq

France

PETERSEN Mary E

Copenhague

Danemark

DORRESTEIJN Adriaan

Mayence

Allemagne

PET1IBONL Manan H.

Washington

U. S. A.

DREUX P.

Paris

France

PEYROT-CLAUSADE M

Marseille

France

DUCHENE Jean-Claude

Banyuls-sur-Mer

France

PLEUEL Fredrik

Stockholm

Su£de

DUPUIS Y

Chatenay Malabry

France

POCCIA Dominic L.

Amherst

US. A.

EIBYE-JACOBSEN Danny

Copenhague

Danemark

POCKLINGTON Patricia

Halifax

Canada

ELDREDGE L. L.

Hawaii

U. S. A.

PONTIER J.

Villeurbanne

France

FAIN A.

Bruxelles

Belgique

POOR G.

Victoria

Australie

FAUCHALD Kristian

Washington

U. S. A.

PUIG H.

Paris

France

FISCHER Albrecht

Mayence

Allemagne

PURSCHKE Gunter

Osnabruck

Allemagne

FITZHUGH Kirk

Los Angeles

U. S. A.

PUTHZ V

Schlitz

Allemagne

FLEURY Anne

Orsay

France

RAIKOVA Olga

Saint-Petersbourg

Russie

FLORET J. J.

Paris

France

RAMIL F.

Vigo

Espagne

FOREY P. L.

Londres

Grande-Bretagne

REISH Donald J

Long Beach

U. S. A.

FOURNIER Judith

Ottawa

Canada

RICHER DE FORGES B

Noumea

Nouvelle-Cal£donie

FRANCOIS Y.

Paris

France

RIEMAN F

Bremerhaven

Allemagne

FRANSEN C.

Leiden

Hoilande

ROUSE Greg

Washington

U. S. A.

GAGNk R

Washington

U. S. A.

SAN MARTIN Guillermo

Madrid

Espagne

GAMBI M. Cristina

Napoli

Italie

SARDA Rafael

Blanes

Espagne

GEHU J. M.

Bailleul

France

SAVAGE D.E.

Berkeley

U. S. A.

GENTIL Frank

Roscoff

France

SCHMID M.

Paris

France

GEORGE David

Londres

Grande-Bretagne

SCHROEDER Paul

Pullmann

U. S. A.

GIANGRANDE Adriana

Lecce

Italie

SCOTT A C.

Surrey

Grande-Bretagne

GIBBS Peter E.

Plymouth

Grande-Bretagne

SIBUET Myriam

Brest

France

GILLET Patrick

Angers

France

SIGVALDADOTTIR Elin

Stockholm

Suede

GLASBY Chris

Canberra

Australie

SIMON Joseph L.

Tampa

U. S. A.

GL1-MAREC Michel

Brest

France

SPIRIDONOV V.

Moscou

Russie

GOERKE Helmut

Bremerhaven

Allemagne

STORK N. E.

Londres

Grande-Bretagne

GOODAY A. J.

Surrey

Grande-Bretagne

TAKEDA M.

Tokyo

Japon

GRASSHOFF M

Frankfurt

Allemagne

TAN C. G. S.

Singapore

GRASSLE Frederick

New Brunswick

Canada

TAYLOR P. D.

Londres

Grande-Bretagne

GRASSLE Judith

New Brunswick

Canada

THURSTON M H.

Surrey

Grande-Bretagne

GRUET Yves

Nantes

France

TOULMOND Andre

Paris

France

GUGLIELMO L.

Messina

Italie

TRICART J.

Strasbourg

France

GUILLAUMETJ. L.

Caen

France

TUDGE Christopher

Brisbane

Australie

HAMLEY Timothy

Brisbane

Australie

VACELETJ.

Marseille

France

HARDEGE Jorg Detelf

Oldenburg

Allemagne

VAN AMERON H. W. J

Krefeld

Allemagne

HAYWARD P J.

Swansea

Grande-Bretagne

VAN SOEST R. W M

Amsterdam

Hoilande

HEALY John

Brisbane

Australie

VOKES E

New Orleans

U. S. A.

HENSLEY D. A.

Puerto Rico

U. S. A.

VOVELLEJean

Paris

France

HILBIG Brigitte

Massachusetts

U. S. A.

WAGELE J. W.

Bielefeld

Allemagne

HODGSON Alan

Grahamstown

Afrique du Sud

WAREN A

Stockholm

Su6de

HOLTE Boerge

Tromsoe

Norv&ge

WARREN Lynda

Cardiff

Grande-Bretagne

HOLTHUIS L. B

Leiden

Hoilande

WATSON J.

Essendon

Australie

HOOPER J N. A.

Brisbane

Australie

WATSON Nikki

Armidale

Australie

HOVE Harry Ten

Amsterdam

Pays-Bas

WESTHEIDE Wilfried

Osnabruck

Allemagne

HUTCHINGS Patricia

Sydney

Australie

WILLIAMS A.

Washington

USA

JENKINS Parish

Cambridge

USA

WILSON Robin

Victoria

Australie

JOUIN-TOULMOND Claude

Paris

France

WITTMANN K.

Vienne

Autriche

KASINSKY Harold E.

Vancouver

Canada

ZEVINAG B

Moscou

Russie

KENDALL Michael

KENSLEY B

Plymouth

Washington

Grande-Bretagne

U. S. A.

ZIBROWIUS Helmut

Marseille

France

Source : MNHN, Paris

ACHEVE D* I M PRIMER
EN JUIN 1996

SLR LES PRESSES
DE

l’imprimerie F. PAILLART
A ABBEVILLE

Date de distribution : 28 juin 1996.
Depot legal : Juin 1996.

N° d'impression : 9745.

Source : MNHN, Paris

DERNIERS TITRES PARUS

RECENTLY PUBLISHED MEMOIRS

A partir de 1993 (Tome 155), les Memoires du Museum sont publies sans indication de serie.
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Prices in French Francs, postage not included.

This volume represents a fundamental contribution to our knowledge of myriapod and
onychophoran biology. It is divided into eight sections, dealing with the following topics:
Historical Myriapodology; Advances in Systematics and Biodiversity; Systematics, Evolution
and Phylogenetic Relationships; Community Studies and Biogeography; Reproductive
Developmental Trends; Physiology, Ecophysiology and Cell Biology; Population Biology. Soil
Ecology and Behaviour; Communities in Ecosystems.

The text includes 79 contributions from 107 authors and should prove essential reading for
all students and researchers of the biology, ecology and systematics of Diplopoda, Chilopoda,
Symphyla, Pauropoda and Onychophora.

Jcan-Jacques GEOFFROY is a researcher of the Centre national de la Recherche
scicntifique. He works at the Museum national d’Histoire naturelle. General Ecology (Brunoy,
France), on population ecology in forest soils, biogeography and biodiversity, mainly on the
arthropod groups Diplopoda and Chilopoda.

Jean-Paul MAURIES is Maitre de Conferences at the Museum national d’Histoire naturelle,
Zoology/Arthropods (Paris, France). A Diplopoda taxonomist, he has mainly published papers
on holarctic and pantropical forms, with special reference to, problems of speciation and
biogeography of millipedes, connected with their habitats in islands, caves and mountains.

Monique NGUYEN DUY - JACQUEMIN is a researcher of the Centre national de la
Recherche scientifique. She works at the Museum national d’Histoire naturelle,
Zoology/Arthropods (Paris, France), on development, sensory organ ultrastructure and
taxonomy of Diplopoda. She is a specialist of the millipede group Penicillata.

I "

EDITIONS
DU MUSEUM
57, RUE CUVIER
75005 PARIS

ISBN 2-85653-502-X
ISSN 1243-4442

550 FF TTC (France)
538,69 FF HT (Elranger)

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