|j Edited by Mies GEOFFROY fPaul MAUR/ES UY-JACQUEMIN
jVhistoire natur:
MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE
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Source :
Bibl lotheque Cent rale Mus6u
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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
© Editions du Museum national d’Histoire naturelle, Paris, 1996
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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
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 :
8
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
9
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 :
10
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 :
12
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
14
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
15
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.
16
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 :
in
INTRODUCTION
17
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
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.
22
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
23
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 :
26
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
27
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
28
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
29
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 :
30
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
31
"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 :
32
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
33
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.
34
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.
36
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.
Japan: Honshu.
REVISION OF THE EAST ASIAN GENUS MILLIPEDE ANA ULACIULUS
37
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.
38
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.
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
39
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.
40
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.
3
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
41
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
42
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.
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REVISION OF THE EAST ASIAN GENUS MILLIPEDE ANAULACIULUS
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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.
46
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
47
|
Sampling Site |
Taxon fr— on -op^.0^ |
•pKImont |
Goi-1 96 10C |
6Pgd 94 10C |
Hole Frequenc Pk 94 10C |
es pg 100 |
i o: |
1 00 |
Mpi |
no |
||||
|
Leukerbad |
m montivaga |
4 |
1.0 |
1.0C |
1.0C |
1.00 |
||||||||
|
Gemml |
m monUvaga |
1 6 |
0.16 |
0.8 |
1.0C |
1.0C |
1.00 |
0.37 |
||||||
|
llligenaip |
m. monUvaga |
13 |
1.0C |
1.0C |
1.0C |
0.96 |
0 96 |
0 04 |
||||||
|
Sanetsch |
m. montivaga |
12 |
1.0C |
0.04 |
0.9t |
1.0C |
1.00 |
1 00 |
||||||
|
Lauenen |
m. montivaga |
13 |
1.0C |
0.9£ |
1.0C |
1.00 |
1 00 |
|||||||
|
Tour de Famolon |
m montivaga |
7 |
1.0C |
1.0C |
1.0C |
1 .00 |
||||||||
|
Morgins |
m montivaga |
4 |
1.0C |
1.0C |
1.0C |
1.00 |
1 .00 |
|||||||
|
Vouvry |
m. montivaga |
1 |
1.0C |
1.0C |
1.0C |
1 00 |
||||||||
|
Rochers de Naye |
m. montivaga |
1 |
1.0C |
1.0C |
l.OC |
1.00 |
||||||||
|
1C |
Lossy |
m. montivaga |
1 |
1.0C |
1.0C |
1.0C |
- |
|||||||
|
1 |
Le Cachoi |
m. montivaga |
1 |
1.0C |
1.0C |
1.0C |
1.00 |
|||||||
|
1 |
La Brdvtne |
m. montivaga |
19 |
0.05 |
0.9£ |
i.6c |
1.0C |
1 .00 |
||||||
|
1C |
Mauborgel |
m. montivaga |
64 |
0.10 |
0.9C |
1.0C |
1.0C |
1.00 |
0.89 |
0 1? |
||||
|
1 |
Si. Georges |
m. montivaga |
39 |
0.19 |
0.8 |
1.0C |
1.0C |
1 .00 |
||||||
|
1 |
Grande Chanreuse |
m. montivaga |
10 |
1.00 |
1.0C |
1.0C |
1.00 |
1 00 |
||||||
|
1C |
Levier |
m. montivaga |
1 1 |
1.00 |
1.0C |
1.0C |
1.00 |
0 96 |
||||||
|
1 |
Oeschaux |
m. hessei |
4 |
1.00 |
1.0C |
1.0C |
1.00 |
1 .00 |
||||||
|
16 |
M6didre |
m hessei |
8 |
1.00 |
1.0C |
1 .00 |
1.00 |
0.90 |
0.10 |
|||||
|
1 |
Belverne |
m. hessei |
7 |
1.00 |
1.0C |
1 .00 |
1.00 |
0 21 |
0 79 |
|||||
|
2C |
Beaune |
m. hessei |
1 |
1.00 |
1.0C |
1.00 |
1.00 |
1 .00 |
||||||
|
2 |
Ancey |
montivaga -group ' |
1 |
1.00 |
1.00 |
1.00 |
1.00 |
1 00 |
||||||
|
22 |
Lantenay |
montivaga -group ' |
1 |
1.00 |
1.00 |
1.00 |
1.00 |
1 00 |
||||||
|
23 |
Vernot |
m. hessei |
10 |
1.00 |
1.00 |
1.00 |
1.00 |
1 00 |
||||||
|
24 |
Coudedoux |
alemannlca |
46 |
1.00 |
1.00 |
0.81 |
0.19 |
1.00 |
||||||
|
25 |
Noir Bo is |
alemannlca |
28 |
1.00 |
1.00 |
0.86 |
0.14 |
1.00 |
1 00 |
|||||
|
26 |
Le Breuil |
cervlna, alemannica" |
3 |
1.00 |
0.83 |
0.17 |
0.83 |
0.17 |
1.00 |
|||||
|
27 |
Si. Ursanne |
cervlna. alemannica " |
31 |
1.00 |
0.38 |
0.63 |
0.96 |
0.04 |
0.98 |
|||||
|
2& |
Bonlol |
alemannica |
1 |
1.00 |
1.00 |
1.-00 |
1.00 |
1 00 |
||||||
|
29 |
Boncourl |
alemannica ? |
4 |
1.00 |
1.00 |
0.50 |
0.50 |
1.00 |
1 .00 |
|||||
|
30 |
Masevaux |
alemannlca |
5 |
1.00 |
1.00 |
1.00 |
1.00 |
|||||||
|
31 |
Le Haul-du-Them |
alemannlca |
2 |
1.00 |
1.00 |
1.00 |
1 .00 |
|||||||
|
32 |
Linihal |
alemannlca |
8 |
1.00 |
1.00 |
1.00 |
1.00 |
1.00 |
||||||
|
33 |
Kenzlngen |
cervna * alemannica ? |
5 |
1.00 |
1.00 |
1.00 |
1 .00 |
|||||||
|
34 |
Hornberg |
cervlna, verhoelll " |
3 |
1.00 |
1.00 |
1.00 |
1.00 |
1 00 |
||||||
|
35 |
Marbach |
cervlna-group" |
2 |
1.00 |
1.00 |
1.00 |
1.00 |
1 .00 |
||||||
|
36 |
alemannlca |
25 |
1.00 |
1.00 |
0.88 |
0.10 |
1.00 |
|||||||
|
37 |
Otlwangen |
serrata |
23 |
0.91 |
0.07 |
1.00 |
0.39 |
0.61 |
1.00 |
0.98 |
||||
|
38 |
Inzlingen |
serrata |
10 |
1.00 |
1.00 |
0.20 |
0.80 |
0.85 |
0.15 |
1 00 |
||||
|
39 |
Hasel |
wehrana |
26 |
1.00 |
1.00 |
1.00 |
0.98 |
0.02 |
1 .00 |
|||||
|
40 |
Todlmoos |
wehrana |
20 |
1.00 |
1.00 |
0.95 |
0.05 |
0.95 |
1 00 |
|||||
|
4 1 |
Menzenschwand |
wehrana |
3 |
1.00 |
1.00 |
1.00 |
1.00 |
|||||||
|
42 |
Laulenburg |
verhoelll |
5 |
1.00 |
1.00 |
1.00 |
1.00 |
0 37 |
0 63 |
|||||
|
43 |
Tlefen6iein |
verhoelll |
5 |
1.00 |
1.00 |
1.00 |
0.10 |
0.90 |
1 .00 |
|||||
|
44 |
Sulz |
cervlna-group ' |
7 |
1.00 |
0.29 |
0.71 |
1.00 |
0.86 |
0.14 |
0 93 |
||||
|
45 |
Schuplari |
cervlna-group ’ |
4 |
1.00 |
0.50 |
0.50 |
1.00 |
1.00 |
0.50 |
0.50 |
||||
|
46 |
Homburg |
cervlna |
1 6 |
1.00 |
0.32 |
0.68 |
1.00 |
0.97 |
0.03 |
1 00 |
||||
|
47 |
Kussnach |
cervina |
4 |
1.00 |
0.13 |
0.88 |
1.00 |
1.00 |
0.50 |
|||||
|
48 |
Altglashuiten |
cervlna + verhoelll ? |
4 |
1.00 |
1.00 |
1.00 |
1.00 |
1 00 |
||||||
|
49 |
Gutachbrucke |
verhoelll, wehrana " |
4 |
1.00 |
1.00 |
0.50 |
0.50 |
1.00 |
0.75 |
|||||
|
50 |
Hu6mersee |
cervlna |
1 1 |
1.00 |
0.70 |
0.30 |
1.00 |
1.00 |
l .00 |
|||||
|
51 |
Dissenhofen |
cervina |
2 |
1.00 |
1.00 |
1.00 |
1.00 |
|||||||
|
52 |
Hemishofen |
cervlna |
1 4 |
1.00 |
1.00 |
1.00 |
1.00 |
1 00 |
||||||
|
53 |
Siaad |
cervlna. alemannica * ' |
8 |
1.00 |
0.13 |
0.88 |
1.00 |
1.00 |
1.00 |
|||||
|
54 |
Baar |
cervlna |
36 |
1.00 |
0.95 |
0.05 |
0.99 |
1.00 |
0.02 |
0 98 |
||||
|
55 |
Oberguisch |
cervlna ? |
2 |
1.00 |
1.00 |
0.75 |
0.25 |
1.00 |
1 .00 |
|||||
|
56 |
Trub |
cervlna |
7 |
1.00 |
0.86 |
0.14 |
1.00 |
1.00 |
||||||
|
57 |
Schelten |
cervlna |
4 |
1.00 |
1.00 |
1.00 |
1.00 |
|||||||
|
58 |
Berllncourl |
cervlna |
20 |
1.00 |
1.00 |
1.00 |
1.00 |
0 61 |
0 39 |
|||||
|
59 |
Gorges de Coud |
cervlna |
3 |
1.00 |
1.00 |
0.33 |
0.67 |
1.00 |
1 .00 |
|||||
|
60 |
Combe Biosse |
ervlna |
5 |
1.00 |
1.00 |
1.00 |
1.00 |
1 .00 |
||||||
|
6i |
Perluls |
ervlna |
20 |
1.00 |
1.00 |
1.00 |
1.00 |
1 .00 |
||||||
|
62 |
PrAvoux |
ervlna |
52 |
0.99 |
0.01 |
0.97 |
0.03 |
0.76 |
0.21 |
1.00 |
0.65 |
0.35 |
||
|
63 |
Yalsainte |
'ervina-group' |
1 |
1.00 |
1.00 |
1.00 |
1.00 |
1.00 |
||||||
|
64 |
Schwarzenmalt |
ervina |
3 |
1.00 |
1.00 |
0.33 |
0.67 |
1.00 |
0.18 |
0.82 |
||||
|
65 |
aun |
ervina-group' |
3 |
1.00 |
0.83 |
0.17 |
1.00 |
1.00 |
1 00 |
|||||
|
66 |
<andersteg |
ervlna |
20 |
0.95 |
0.05 |
0.95 |
0.05 |
0.58 |
0.43 |
1.00 |
0.08 |
0.92 |
||
|
67 |
Zwelsimmen |
ervlna / montivaga |
20 |
0.48 |
0.53 |
0.15 |
0.85 |
0.25 |
0.75 |
1.00 |
0.28 |
0.42 |
0.30 |
|
|
68 |
3oltigen |
ervina / montivaga |
25 |
0.78 |
0.22 |
0.75 |
0.25 |
0.71 |
0.30 |
1.00 |
0.15 |
0.85 |
||
|
69 |
3eseux |
ervlna / montivaga |
27 |
0.43 |
0.57 |
0.71 |
0.29 |
0.06 |
0.94 |
1.00 |
1 .00 |
|||
|
70 |
Rochelort |
ervlna / montivaga |
3 |
0.57 |
0.33 |
0.83 |
0.17 |
|||||||
|
71 |
»4auvaise Combe |
ervina / montivaga |
12 |
0.67 |
0.33 |
0.46 |
0.54 |
0.13 |
0.88 |
1.00 |
||||
|
72 |
a Chaux-du-Milieu |
ervina / montivaga |
39 |
0.67 |
0.33 |
0.60 |
0.40 |
0.18 |
0.82 |
1.00 |
||||
|
73 |
erridres |
ervina / montivaQa |
38 |
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
48
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
49
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.
50
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
51
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.
54
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
55
- 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) ;
56
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
57
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 :
58
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
59
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 :
60
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.
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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
63
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
64
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
65
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.
68
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
69
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.
70
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
71
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.
72
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.
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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
75
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.)
76
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
77
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.
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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
83
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)
84
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
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= 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
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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
87
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
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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