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REVUE SUISSE
DE
ZOOLOGIE
REVUE SUISSE
ZOOLOGIE
ANNALES
DE LA
SOCIÉTÉ SUISSE DE ZOOLOGIE
ET DU
MUSEUM D'HISTOIRE NATURELLE
DE LA VILLE DE GENEVE
FI
GENÈVE
1996
ISSN 0035-418X
TABLE DES MATIÈRES
TOME 103 — 1996
Fascicule 1
DESQUEYROUX-FAUNDEZ, Ruth & Rob W.M. VAN SOEST. A review of Iophonidae,
Myxillidae and Tedaniidae occurring in the South East Pacific (Porifera:
Roecilosclenida)) wn eats ars RE ateneo sia enito dele
Monney, Jean-Claude, Lucia LUISELLI & Massimo CAPULA. Taille et mélanisme chez
Vipera aspis dans les Préalpes suisses et en Italie centrale et comparaison avec
différentes populations alpines de Vipera berus..........................
CoNDE, Bruno. Diploures Campodeides de la Pestera de la Movile (Movile Cave),
Dobroudjaméendonale(Roumante) see ei era CE ee
ANGELINI, Fernando. A New Species and records of the genus Pseudoagathidium
Angelini from Uganda (Coleoptera, Leiodidae, Agathidiini).................
ASSING, Volker & Paul WUNDERLE. A Revision of the Madeiran species of the genus
Geostiba Thomson, 1858 (Coleoptera: Staphylinidae). .....................
GOLOVATCH, Sergei I. The millipede family Paradoxosomatidae on Borneo, with
contributions to the faunas of some other islands of the Sunda area (Diplopoda,
BONES MIA) apy senses tu arts ane var RL ce A RA:
Pace, Roberto. Aleocharinae della Sottoregione Africana Orientale al Museo di
Ginevra (Coleoptera, Staphylinidae) Parte HI (conclusione). ................
MAHUNKA, Sandor. Oribatids from Sarawak I. (Acari: Oribatida). New and interesting
nitestrominelGenevaMuseumniEX XVII Mp EEE MR PR UC
PERREAU, Michel. Contribution à la connaissance des Cholevidae du Japon et de
TAgaiti((CO EROI ne RA RS EL e re
Fascicule 2
FRISCH, Johannes. Revision westmediterraner Scopaeus - Arten (Coleoptera, Staphy-
linidae: Paederinae) und Beschreibung einer neuen Art aus Südspanien und
Marokko. 2. Beitrag zur Kenntnis der Gattung Scopaeus Frichson. ...........
LOURENCO, Wilson R. Microcharmus hauseri, nouvelle espece de Scorpion de Mada-
SAS Calg (SCORPLONE Sess Wil CARRI TT
GORBUNOV, Oleg G. & Yutaka ARITA. New and little-known Oriental Melittia Hübner
(Lepidoptera, Sesiidae), from the collection of Muséum d’histoire naturelle,
Genève
GEIGER, Daniel. Haliotids in the Red Sea, with neotype designation for Haliotis uni-
lateralis Lamarck, 1822 (Gastropoda: Prosobranchia). . ......:.............
3-79
81-100
101-114
115-118
119-150
283-297
301-318
319-322
323-338
339-354
VI TABLE DES MATIERES
PAGÉS, Jean. Un Evalljapygidae (Diplura) canadien: Evalljapyx scaundersi n. sp. -
DicellunatanG enayensid Cx ERE RECETTORE
ConDE, Bruno & Camille BARETH. Une évaluation de Stygiocampa, sous-genre
troglomorphe de Plusiocampa (Diplura Campodeidae), avec la description d’une
mouvellerespecerde;Senbiecronientale san TRENO TSE
BOURGAT, Robert, Cyril ROURE & Sim-Dozou KuLo. Nouvelles données sur les Tré-
matodes d’Amphibiens d’ Afrique Occidentale. Description d’ Haematoloechus
CVO UAE TY, IOs eco NS en euere see eee D 0 Du ne Blais co 0
Pace, Roberto. Aleocharinae della Colombia e dell’Ecuador: Parte I (Coleoptera,
Stapliylimi daca nn at lake a cn a Sean eine Se Wee aie eee
STEINER, Werner A. Distribution of entomopathogenic nematodes in the Swiss Alps. . .
RIEDEL, Alexander & Klaus SCHONITZER. Revision of the soil-weevil genus Guineobius
Osella from New Guinea (Coleoptera, Curculionoidea, Entiminae, Celeuthetini).
Cuccoporo, Giulio. Revision of the Japanese rove-beetles of the genus Megarthrus
(Coleoptera, Staphylinidae, Proteininae). . ...............................
NAYROLLES, Pierre. Contribution to the knowledge of European Bourletiellidae
(Collembola, Symphypleona). III. Description of some species of Heteros-
minthunussandYDeuterosminthunus. «cls ar Da ee Se
BARBALAT, Sylvie. Influence de l’exploitation forestière sur trois familles de colé-
opteres liés au bois dans les Gorges de l’Areuse. .........................
Fascicule 3
KNOFLACH, Barbara. Three new species of Carniella from Thailand (Araneae, Theri-
AO) EN e oe a e i ee NER _
KODADA, Jän. Spalacosostea, an anomalous new terrestrial dryopid from South East
STANCO COP{ErAADEY OPTIO I
BAEHR, Martin. A peculiar new species of Anomotarus (Nototarus) from New Guinea
(Coleoptera; Carabidae; Eebiinae). EI RE
MESSNER, Benjamin, Joachim Apis & Klaus Peter ZULKA. Stigmale Plastronstrukturen,
die einigen Diplopoden-Arten eine submerse Lebensweise in kaltem und in
file bendemaWasseRermo SCIE oe eee
DALENS, Henri, André ROUSSET & Didier FOURNIER. Les formes épigées du genre
Oritoniscus (Crustacea, Isopoda, Oniscidea). I. Le complexe Oritoniscus flavus. .
PACE, Roberto. Aleocharinae della Colombia e dell'Ecuador: Parte II (Coleoptera,
Sta play limi eee eens, epics esi IONE
RIGHI, Gilberto. Some Venezuelan Oligochaeta Glossoscolecidae and Octochaetidae. . .
SALGADO Costas, José Maria. Un nouveau Apoduvalius du bassin de la rivière Nalon.
Stunes Espagne (Coleoptera, Trechidae)........2. er Eee eee
SCHAWALLER, Wolfgang. Spinolyprops pakistanicus sp. n. (Coleoptera Tenebrionidae),
an Oriental element in the fauna of northern Pakistan. ......................
BARTSCH, Ilse. Agaunopsis (Acari, Halacaridae) of the Sevastopol arcea; supplemen-
tanyanotesionitaxonomyzandfecolosya 4 ee eee eee
ROUGEMONT, G.M. DE. Stiliderus and Stilicoderus: New data and new species (Cole-
OpieramStaphylinidaetRaedennae) PAPER PERRET cia are
Pages
355-367
369-381
383-394
395-437
439-452
453-473
475-524
525-551
553-564
567-579
581-605
607-611
613-622
623-641
643-676
677-684
685-690
691-695
697-712
713-736
TABLE DES MATIERES
UHMANN, Gerhard. Indo-australische Anthicidae (Coleoptera) in Naturhistorischen
MUSCUMINGCNE RE IR NT
MENDES, Luis F. Further data on the Nicolettidae (Zygentoma), with description of a
THEMES DEC LES OMIMIAURUU SEE IRE ORI TTT
PERRET, Jean-Luc. Une nouvelle espèce du genre Ptychadena (Anura, Ranidae) du
BOHME, Wolfgang & Andreas SCHMITZ. A new lygosomine skink (Lacertilia: Scinci-
(ACMZANLASP IS) TOMICA MELOO NETTE TI
Fascicule 4
ZOOLOGIA ET BOTANICA 96, Fribourg, 21-23 February 1996 (Annual Conference of the
SWISSPZO0 0 21CA NS OC) RE RE RR E
SCHLÜSSEL, André & Jean-Paul THEURILLAT. Synusial structure of heathlands at the
subalpine/alpine ecocline in Valais (Switzerland). .........................
BAUR, Bruno, Jasmin JOSHI, Bernhard SCHMID, Ambros HANGGI, Daniel BORCARD,
Josef STARY, Ariane PEDROLI-CHRISTEN, G. Heinrich THOMMEN, Henryk LUKA,
Hans-Peter RUSTERHOLZ, Peter OGGIER, Stephan LEDERGERBER & Andreas
ERHARDT. Variation in species richness of plants and diverse groups of inver-
tebrates in three calcareous grasslands of the Swiss Jura mountains............
GRAF, Markus, Alexander Imanuel WANDELER & Peter LUps. Die räumliche Habitat-
nutzung einer Dachspopulation (Meles meles L.) im schweizerischen Mittelland.
LANG, Claude & Olivier REYMOND. Le zoobenthos comme indicateur des perturbations
d’origine humaine dans deux lacs de montagne. ..........................
GÖHNER, Manuela & Wolfgang PFEIFFER. Über die Verbreitung der Mauthner Axone
bei Fischen und Amphibien und ihren Zusammenhang mit der Schreckreaktion
demOStawOphy Sigund eA tates. sci O Rene
MERZ, Bernhard. Die Asteiidae (Diptera) der Schweiz. . .........................
DURETTE-DESSET, Marie-Claude & Claude VAUCHER. Molostrongylus acanthocolpos
gen. n., Sp. n., (Nematoda, Trichostrongylina, Molineoidea) parasite de Molos-
sops temmincki (Chiroptera, Molossidae) au Paraguay. .....................
BURCKHARDT, Daniel. On some Ancistria spp. from the Natural History Museum,
Vienna @oleopteranbassandzidae) ar an. ee
LOBL, Yvan & Serguei A. KURBATOV. A review of the Japanese Tychobythinus and
Bythoxenites (Coleoptera, Staphylinidae, Pselaphinae)......................
PERREAU, Michel. Nouveaux Cholevinae d’ Asie (Coleoptera Leiodidae)............
VII
Pages
737-748
749-756
757-766
767-774
777-794
795-800
801-833
835-850
851-858
859-891
893-904
905-913
915-918
919-938
939-949
| 77 AY, RA Ay | ; eo è a
DA x i à
Urne WIR REDET TMS EN
Donne dr | ES |
a OS EAN SEE h “ee |.
le U ge et ich Le
INDEX DES AUTEURS
par
ORDRE ALPHABÉTIQUE
ANGELINI, Fernando. A New Species and records of the genus Pseudoagathidium
Angelini from Uganda (Coleoptera, Leiodidae, Agathidnni). ................
ASSING, Volker & Paul WUNDERLE. A Revision of the Madeiran species of the genus
Geostiba Thomson, 1858 (Coleoptera: Staphylinidae). .....................
BAEHR, Martin. A peculiar new species of Anomotarus (Nototarus) from New Guinea
(Col opera bDIdaeMEebinae) RAP ERA N
BARBALAT, Sylvie. Influence de l’exploitation forestière sur trois familles de colé-
optères lies au bois dans les Gorges de l’Areuse.........................
BARTSCH, Ilse. Agaunopsis (Acari, Halacaridae) of the Sevastopol arcea; supplemen-
LARVANO (E SIONMAXONOMYIINAIECOLO VER ER ER
BAUR, Bruno, Jasmin JosHI, Bernhard SCHMID, Ambros HANGGI, Daniel BORCARD,
Josef STARY, Ariane PEDROLI-CHRISTEN, G. Heinrich THOMMEN, Henryk LUKA,
Hans-Peter RUSTERHOLZ, Peter OGGIER, Stephan LEDERGERBER & Andreas
ERHARDT. Variation in species richness of plants and diverse groups of inver-
tebrates in three calcareous grasslands of the Swiss Jura mountains............
BOHME, Wolfgang & Andreas SCHMITZ. A new lygosomine skink (Lacertilia: Scinci-
C'ACSPLANASPI) ATO MICAME TOO NIE ARTO ONT OTO
BOURGAT, Robert, Cyril ROURE & Sim-Dozou KuLo. Nouvelles données sur les Tré-
matodes d’Amphibiens d’ Afrique Occidentale. Description d’Haematoloechus
CUBE CRIES Aer EHEN PRE LILIAN RENI IE
BURCKHARDT, Daniel. On some Ancistria spp. from the Natural History Museum,
Wriennat(Coleopterasbassandndae) fem. see tices tat tg ete reece IN
CONDÉ, Bruno. Diploures Campodéidés de la Pestera de la Movile (Movile Cave),
Pobroudjasmertdionalel(Roumanie) ERRE see ace SER rere eee
CONDÉ, Bruno & Camille BARETH. Une évaluation de Stygiocampa, sous-genre
troglomorphe de Plusiocampa (Diplura Campodeidae), avec la description d’une
nouyellesespecezderSenbiesorientaleg ee PRE ROOT
Cuccoporo, Giulio. Revision of the Japanese rove-beetles of the genus Megarthrus
(Coleoptera Staphylinidae peroteminac) se. PRO eee O
DALENS, Henri, André Rousset & Didier FOURNIER. Les formes épigées du genre
Oritoniscus (Crustacea, Isopoda, Oniscidea). I. Le complexe Oritoniscus flavus. .
DESQUEYROUX-FAUNDEZ, Ruth & Rob W.M. VAN SOEST. A review of Iophonidae,
Myxillidae and Tedaniidae occurring in the South East Pacific (Porifera:
ROCCHOSCICH AA REANO
DURETTE-DESSET, Marie-Claude & Claude VAUCHER. Molostrongylus acanthocolpos
gen. n., sp. n., (Nematoda, Trichostrongylina, Molineoidea) parasite de Molos-
sops temmincki (Chiroptera, Molossidae) au Paraguay. .....................
Pages
115-118
119-150
601-611
553-564
697-712
801-833
767-774
383-394
915-918
101-114
369-381
475-524
623-641
3-79
905-913
X INDEX DES AUTEURS
FRISCH, Johannes. Revision westmediterraner Scopaeus - Arten (Coleoptera, Staphy-
linidae: Paederinae) und Beschreibung einer neuen Art aus Südspanien und
Marokko. 2. Beitrag zur Kenntnis der Gattung Scopaeus Erichson............
GEIGER, Daniel. Haliotids in the Red Sea, with neotype designation for Haliotis uni-
lateralis Lamarck, 1822 (Gastropoda: Prosobranchia). ....................
GÖHNER, Manuela & Wolfgang PFEIFFER. Uber die Verbreitung der Mauthner Axone
bei Fischen und Amphibien und ihren Zusammenhang mit der Schreckreaktion
TÉROSTATODY SAUNA ANUTA AE EEE oa ae ee a cane E
GOLOVATCH, Sergei I. The millipede family Paradoxosomatidae on Borneo, with
contributions to the faunas of some other islands of the Sunda area (Diplopoda,
Rolydesmida) nn cic es nee inne OM eal ole ee a eee
GORBUNOV, Oleg G. & Yutaka ARITA. New and little-known Oriental Melittia Hübner
(Lepidoptera, Sesiidae), from the collection of Muséum d’histoire naturelle,
GENÈVE RE le NE Mir II RIT O II GOTI
GRAF, Markus, Alexander Imanuel WANDELER & Peter LUps. Die räumliche Habitat-
nutzung einer Dachspopulation (Meles meles L.) im schweizerischen Mittelland.
KNOFLACH, Barbara. Three new species of Carniella from Thailand (Araneae, Theri-
III TREIA ET HERE AR ATO
KODADA, Jan. Spalacosostea, an anomalous new terrestrial dryopid from South East
Asian(Coleopteras Dry opidae) ras mia ISS
LANG, Claude & Olivier REYMOND. Le zoobenthos comme indicateur des perturbations
d’origine humaine dans deux lacs de montagne. ..........................
LÔBL, Yvan & Serguei A. KURBATOV. A review of the Japanese Tychobythinus and
Bythoxenites (Coleoptera, Staphylinidae, Pselaphinae). .....................
LOURENÇO, Wilson R. Microcharmus hauseri, nouvelle espèce de Scorpion de Mada-
SASCAM(SCONDIONES BULNITAE) EEE OE
MAHUNKA, Sandor. Oribatids from Sarawak I. (Acari: Oribatida). New and interesting
mutessinom: the, Geneva IMUuseumIXXVIII SR cee cpa eee
MENDES, Luis F. Further data on the Nicolettidae (Zygentoma), with description of a
NAW HOSES tom MEMOS, Geo RO
MERZ, Bernhard. Die Asteiidae (Diptera) der Schweiz. ..........1...............
MESSNER, Benjamin, Joachim Apis & Klaus Peter ZULKA. Stigmale Plastronstrukturen,
die einigen Diplopoden-Arten eine submerse Lebensweise in kaltem und in
fliebendemiWassenennogliChen 2... OLO OI
MONNEY, Jean-Claude, Lucia LUISELLI & Massimo CAPULA. Taille et mélanisme chez
Vipera aspis dans les Préalpes suisses et en Italie centrale et comparaison avec
differentes;populations alpines de: Vipera berus. .. Ri. Arlo see
NAYROLLES, Pierre. Contribution to the knowledge of European Bourletiellidae
(Collembola, Symphypleona). III. Description of some species of Heteros-
minthunussand Deuterosminthurus TIE AO EEE
PACE, Roberto. Aleocharinae della Sottoregione Africana Orientale al Museo di
Ginevra (Coleoptera, Staphylinidae) Parte III (conclusione). ................
PACE, Roberto. Aleocharinae della Colombia e dell’ Ecuador: Parte I (Coleoptera,
Staphylinidae) PART E HSS 48 VANNES ME A RARE
PACE, Roberto. Aleocharinae della Colombia e dell’ Ecuador: Parte II (Coleoptera,
SÉAPHYINITAC) et. RR LEO EEE
PAGES, Jean. Un Evalljapygidae (Diplura) canadien: Evalljapyx scaundersi n. sp. -
DicellujatanG enavensiaexexd ENTREE CETO
Pages
301-318
339-354
859-891
151-193
323-338
835-850
567-579
581-605
851-858
919-938
319-322
259-282
749-756
893-904
613-622
81-100
525-551
195-258
395-437
643-676
355-367
INDEX DES AUTEURS
PERREAU, Michel. Contribution à la connaissance des Cholevidae du Japon et de
RAA (COle Opera) ics see cess RESOR SIRIO
PERREAU, Michel. Nouveaux Cholevinae d’ Asie (Coleoptera Leiodidae)............
PERRET, Jean-Luc. Une nouvelle espèce du genre Prychadena (Anura, Ranidae) du
RIEDEL, Alexander & Klaus SCHÖNITZER. Revision of the soil-weevil genus Guineobius
Osella from New Guinea (Coleoptera, Curculionoidea, Entiminae, Celeuthetini).
RIGHI, Gilberto. Some Venezuelan Oligochaeta Glossoscolecidae and Octochaetidae. .
ROUGEMONT, G.M. DE. Stiliderus and Stilicoderus: New data and new species (Cole-
Opteraystaphydinidaes Pacdenin ac) sare a
SALGADO CosTAs, José Maria. Un nouveau Apoduvalius du bassin de la rivière Nalon.
Asturies-Espasne (Coleoptera Trechidae)e.n. 25. - 2.252 aon acess eee aes
SCHAWALLER, Wolfgang. Spinolyprops pakistanichs sp. n. (Coleoptera), an Oriental
element@inäthesfaunago@northernVBakistaney ER
SCHLÜSSEL, Andre & Jean-Paul THEURILLAT. Synusial structure of heathlands at the
subalpine/alpine ecocline in Valais (Switzerland). .........................
STEINER, Werner A. Distribution of entomopathogenic nematodes in the Swiss Alps. .
UHMANN, Gerhard. Indo-australische Anthicidae (Coleoptera) in Naturhistorischen
Mine tt Ce Ty à à TRO O UE aca
ZOOLOGIA ET BOTANICA 96, Fribourg, 21-23 February 1996 (Annual Conference of the
SWISs$7001l021ealSocie vB oH ou oN'e
XI
Pages
283-297
939-949
757-766
453-473
677-684
713-736
685-690
691-695
795-800
439-452
737-748
777-794
7
72
REVUE SUISSE DE ZOOLOGIE
REVUE SUISSE DE ZOOLOGIE
TOME 103 — FASCICULE 1
Publication subventionnée par l'Académie suisse des Sciences naturelles
et la Société suisse de Zoologie
VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genève
FRANÇOIS BAUD
Conservateur au Muséum d'histoire naturelle de Genève
DANIEL BURCKHARDT
Chargé de recherche au Muséum d'histoire naturelle de Genève
Comité de lecture
Le président de la Société Suisse de Zoologie
Le directeur du Muséum de Genève: Volker MAHNERT — Systématique des
vertébrés — Muséum de Genève
Le président du comité: Ivan LOBL — Systématique des Insectes — Muséum de
Genève
Patrick GUÉRIN — Physiologie et éthologie des arthropodes — Institut de Zoologie,
Neuchâtel
Willy MATTHEY — Ecologie, entomologie — Institut de Zoologie, Neuchâtel
Claude MERMOD — Ethologie et écologie des vertébrés — Université de Neuchâtel
Paul SCHMID-HEMPEL — Ecoéthologie, biologie des populations — Institut f.
Terrestrische Ökologie, ETH Zürich, Schlieren
Steve STEARNS — Biologie de l'évolution — Institut f. Zoologie, Basel
Beat TSCHANZ — Ethologie des Vertebres — Zoologisches Institut, Bern
Claude VAUCHER — Systématique des Invertébrés — Muséum de Genève
La préférence sera donnée aux travaux concernant les domaines suivants: Biogéographie,
systématique, écologie, éthologie, morphologie, et anatomie comparée, physiologie.
Administration
MUSÉUM D'HISTOIRE NATURELLE
1211 GENÈVE 6
PRIX DE L'ABONNEMENT:
SUISSE Fr. 225.— UNION POSTALE Fr. 230. —
(en francs suisses)
Les demandes d'abonnement doivent être adressées
à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse
ANNALES
de la
SOCIÉTÉ SUISSE DE ZOOLOGIE
et du
MUSEUM D'HISTOIRE NATURELLE
de la Ville de Genéve
tome 103
fascicule 1
1996
REVUE SUISSE DE ZOOLOGIE
g -
EI GENEVE MARS 1996 ISSN 0035 - 418X
REVUE SUISSE DE ZOOLOGIE
TOME 103 — FASCICULE 1
Publication subventionnée par l'Académie suisse des Sciences naturelles
et la Société suisse de Zoologie
VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genève
FRANÇOIS BAUD
Conservateur au Muséum d'histoire naturelle de Genève
DANIEL BURCKHARDT
Chargé de recherche au Muséum d'histoire naturelle de Genève
Comité de lecture
Le président de la Société Suisse de Zoologie
Le directeur du Muséum de Genève: Volker MAHNERT — Systématique des
vertébrés — Muséum de Genève
Le président du comité: Ivan LOBL — Systématique des Insectes — Muséum de
Genève
Patrick GUÉRIN — Physiologie et éthologie des arthropodes — Institut de Zoologie,
Neuchâtel
Willy MATTHEY — Ecologie, entomologie — Institut de Zoologie, Neuchâtel
Claude MERMOD — Ethologie et écologie des vertébrés — Université de Neuchâtel
Paul ScHMID-HEMPEL — Ecoéthologie, biologie des populations — Institut f.
Terrestrische Ökologie, ETH Zürich, Schlieren
Steve STEARNS — Biologie de l'évolution — Institut f. Zoologie, Basel
Beat TSCHANZ — Ethologie des Vertebres — Zoologisches Institut, Bern
Claude VAUCHER — Systématique des Invertébrés — Muséum de Genève
La préférence sera donnée aux travaux concernant les domaines suivants: Biogéographie,
systématique, écologie, éthologie, morphologie, et anatomie comparée, physiologie.
Administration
MUSEUM D'HISTOIRE NATURELLE
1211 GENÈVE 6
PRIX DE L'ABONNEMENT:
SUISSE Fr. 225.— UNION POSTALE Fr. 230.—
| (en francs suisses)
Les demandes d'abonnement doivent être adressées
| à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse
REVUE SUISSE DE ZOOLOGIE, 103 (1) : 3-79; mars 1996
A review of Iophonidae, Myxillidae and Tedaniidae
occurring in the South East Pacific (Porifera: Poecilosclerida)
Ruth DESQUEYROUX-FAUNDEZ* & Rob W. M. VAN SOEST**
*Muséum d’histoire naturelle, P.O. Box 6434, ch-1211 Geneva, Switzerland.
**Institute for Systematics and Population Biology (Zoölogisch Museum),
University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, the Netherlands.
A review of Iophonidae, Myxillidae and Tedaniidae occurring in the
South East Pacific (Porifera: Poecilosclerida).- Based on collections
from the coasts of Chile, Peru, and the Galapagos Islands, the SE. Pacific
species of Jophon, Myxilla and Tedania are revised. Descriptions and
illustrations are provided for eleven previously known species (including
thirteen new synonymy proposals). Six new species are described: /ophon
chilense, I. timidum, I. tubiforme, Myxilla (E.) dracula, M. (B.) asymme-
trica, and Tedania (T.) galapagensis. A key for the identification of the
species is included. The results of this study provided the basis for a
revised classification of related genera and a new family assignment of
Tophon, Myxilla and Tedania. In accordance with suggestions in the recent
literature /ophon is assigned to the revived family Iophonidae Burton, 1929
of the suborder Microcionina Hajdu er al., 1994. Myxilla and Tedania are
assigned to different families of the suborder Myxillina, viz. a restricted
Myxillidae Topsent, 1928 and Tedaniidae Ridley & Dendy, 1886 both of
the suborder Myxillina Hajdu er al., 1994. It is demonstrated that the
Myxilla-like genera Burtonanchora de Laubenfels, 1936, Ectyomyxilla
Lundbeck, 1909 and Stelodoryx Topsent, 1904, are based on characters
without phylogenetic significance. It is proposed to retain these as sub-
generic units within Myxilla. On similar grounds, Trachytedania Ridley,
1881 and Tedaniopsis Dendy, 1924, are considered subgenera of Tedania.
Diagnosis and generic composition of the families are discussed.
Key-words: Iophonidae - Myxillidae - Tedaniidae - SE. Pacific - Taxonomy
INTRODUCTION
Numerous species of Poecilosclerida are reported from the coasts of the South
East Pacific (RIDLEY 1881; RIDLEY & DENDY 1886, 1887; WILSON 1904; THIELE 1905;
DE LAUBENFELS 1939; DESQUEYROUX 1972, 1976, DESQUEYROUX-FAUNDEZ, 1990;
Manuscript accepted 04.05.1995.
4 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Haspu & DESQUEYROUX-FAUNDEZ 1994) and adjacent areas in the South West
Atlantic (BURTON 1932, 1940; SARÀ 1978). Among the poecilosclerids, particularly
the genera Jophon, Myxilla and Tedania are well-represented. However, type speci-
mens of newly described species from the area and specimens assigned to them sub-
sequently have never been revised, so the fauna is not well-known. Through her
contacts with many Chilean institutions, the senior author has assembled a com-
prehensive collection of sponges from most areas of the extensive coastline of Chile;
additional samples from Chile, Peru and the Galapagos Islands were obtained from
the collections of the former Smithsonian Oceanographic Sorting Centre. In addition
to this, the type and other specimens of previously described species were borrowed
from other institutions, and this combined comprehensive collection has been revised
by us. The present study is one of an ongoing series of studies of the Poecilosclerid
sponges of the South East Pacific made by the senior author (cf. HAIDU &
DESQUEYROUX-FAUNDEZ 1994),
The extensiveness of the available material enabled us to make a contribution to
the improvement of the classification of the Poecilosclerida initiated recently by HAJDU
et al. (1994). The large number of species of the genera Jophon, Myxilla and Tedania
described in the study area and adjacent areas induced us to concentrate the present
study on these genera. A further reason is that they occupy key positions in the preli-
minary changes in the Poecilosclerida classifications proposed by HAJDU et al. (1994).
Assignment of species to genera and families of the myxillid Poecilosclerida is
problematic because of the lack of consensus among recent authors over their content
and relationships. VAN SOEST (1984) employed a wide diagnosis of Myxillidae based
on the presence of diactinal (tylote or strongylote) ectosomal spicules and a reticulate
choanosomal skeleton. Thus diverging genera such as /ophon, Myxilla, Acarnus,
Tedania, and Lissodendoryx were included in a single family Myxillidae. BERGQUIST
(1978) and BERGQUIST & FROMONT (1988) assigned Tedania to a separate family
Tedaniidae based on the absence of chelas and the possession of onychaetes; they
emphasized an isotropic choanosomal skeleton as a synapomorphy for a more res-
tricted Myxillidae. Many authors, e.g. BOURY-ESNAULT & VAN BEVEREN (1982)
followed in part BERGQUIST (1978).
In their preliminary review of Poecilosclerid characters, HAJDU er al. (1994)
proposed an extensive reorganization of the genera formerly considered to be
"myxillids". They observed a striking consistency in the presence or absence of toxas
and the chela morphology: species and genera with arcuate and anchorate chelas
(together named "tridentate" chelas) never have toxas, (with some notable exceptions,
e. g. spp. of Dendrocia); these are confined to sponges possessing palmate chelas.
This was in line with a separation between palmate and "tridentate" coelosphaerid
sponges made earlier by LÉVI & LÉVI (1983). These authors proposed to recognize
two families of fistular Poecilosclerida, Coelosphaeridae with arcuate chelas and
Cornulidae with palmate chelas. LÉVI & Levi (1983) did not mention that most of
their Cornulidae had microspined tylote or strongylote apices. HAJDU er al. (1994)
suggested to include other genera with palmate chelas into the Cornulidae, and to de-
S-E PACIFIC SPECIES OF POECILOSCLERIDA 5
emphasize the fistular growth form. The genera concerned are Acarnus (cf. VAN
SOEST et al., 1991 for a revision), Megaciella and also /ophon, all three with micro-
spined tylotes. The latter genus, however, is a less clear-cut case because of the lack
of toxas, the possession of anisochelas and the peculiar autapomorphous bipocilla.
Moreover, if Iophon is included in the Cornulidae in the widened sense, the family
name has to change to Iophonidae Burton, 1929. VAN SOEST et al. (1994) revised the
Cornulidae s.s. and concluded these are a closely related assemblage probably related
to lophon and Acarnus; however, they did not include specimens of /ophon in their
studies. The present extensive collections contain ample material of this genus
allowing a comparison with the results of Van Soest ef al. (1994).
A further proposal of HAJDU er al. (1994) is a separation of myxillids pos-
sessing anchorate chelas (e.g. Myxilla) and those possessing arcuate chelas (e.g.
Lissodendoryx). The substantial number of Myxilla s.l. species found in the study area
is used to narrow down the differences between the proposed Myxillidae s.s. and an
extended Coelosphaeridae. The generic contents of the Myxillidae s.s. is discussed on
a preliminary basis. The Coelosphaeridae s.l. (including Lissodendoryx) will be
treated in a forthcoming paper.
HAJDU et al. (1994) expressed uncertainty over the distinctness of a family
Tedaniidae. The substantial number of Tedania species in the study area as well as a
large number of borrowed type slides available to us enabled us to revise this group.
The revisions presented below are intended to give a firmer basis to discus-
sions of poecilosclerid classification. In the absence of similar revisions of myxillids
from other areas, it is not possible yet to present a definitive new system.
MATERIAL AND METHODS
Specimens from the Chilean Coast between Iquique and Seno Almirantazgo
(20°-54°S) were collected during surveys realised from 1966 to 1992. Collector
references, are indicated in the text as: collecting number (abbreviations are explained
below), locality, geographical coordinates, date (day, month, year) and depth in m.
CHI.: Samples from the Instituto de Oceanologia Universidad de Valparaiso,
Chile;
Co.93: Samples from Facultad de Pesquerias y Oceanografia Universidad
Austral de Chile, Puerto Montt;
Ga: Samples from SEPBOP program;
HE: Samples from Hero expedition 1969-1972 from the National Science
Foundation and Departamentos de Biologia Marina y Oceanograffa y de Zoologia de
la Universidad de Concepcion, Chile;.
MNHNC: Samples from the Museo nacional de Historia natural de Chile,
Santiago;
MONT.: Samples from the Universidad de Chile, Valparaiso, Estacion de
Biologia Marina de Montemar;
6 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Por., E: Samples from Departamento de Ciencias Ecolögicas, Universidad de
Chile, Santiago;
VALD.: Samples from the Universidad Austral de Chile, Valdivia;
A second lot of specimens was obtained from the former Smithsonian Oceanographic
Sorting Centre: several specimens from Chile, Peru and Galapagos were collected
during the 1966 SE Pacific Biological Oceanography Program (SEPBOP) on board of
R.V. "Anton Bruun"; other specimens were collected by individual collectors. Repre-
sentative sets of specimens of this collection are deposited in the United States Natio-
nal Museum of Natural History at Washington, including holotypes of new species, in
the Zoölogisch Museum Amsterdam, and the Muséum d'histoire naturelle de Genève.
For comparison a comprehensive set of type fragments and type slides of
lophon, Myxilla and Tedania species and related genera were borrowed from a
number of institutions.
Abbreviations for institutions used in the text are:
BMNH: Natural History Museum, London;
CNM: Canadian National Museum;
IOUV: Instituto de Oceanologia Universidad de Valparaiso:
MCSN: Museo Civico di Storia Naturale “Giacomo Doria”, Genova;
MHNG: Museum d'histoire naturelle, Geneva;
MNHN: Museum National d’ Histoire Naturelle, Paris;
MNHNC: Museo nacional de historia natural de Chile, Santiago;
MZSF: Museo Zoologico de “La Specola”, Firenze (Italy);
MZUC: Museo zoologia Universidad Concepcion, Chile;
UCH: Departamento de Ciencias Ecolögicas, Universidad de Chile,
Santiago.
USC/AHF University of Southern California; Allan Hancock Foundation
USNM: National Museum of Natural History, Smithsonian Institution,
Washington; formerly United States National Museum;
ZMA: Zoölogisch Museum, Amsterdam;
ZMB: Museum für Naturkunde der Humboldt-Universität, Berlin;
ZMK: Zoologisk Museum Kobenhavn;
ZMH: Zoologisches Museum Hamburg
Skeletal slides and dissociated spicule mounts were made following Rützler
(1978). The SEM study was made using a Zeiss Digital Scan Microscope dsm 940,
with accelerating voltage of 20 KV and magnification up to 10,000 times.
Scales for specimens represent 0.5 cm. Scales for sem pictures are indicated
with each spicule. Measurements of spicules on Tables refer to minimum-mean
maximum in um; in text they refer to minimum-maximum, in um.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 7
SYSTEMATIC PART
Order: Poecilosclerida; Suborder: Microcionina Hajdu, van Soest & Hooper,
1994
Family: Iophonidae Burton (1929, as section Iophoneae)
Diagnosis: Microcionina with spined tylotes as ectosomal megascleres.
Tophon Gray, 1867
Synonyms: Alebion Gray (1867), Menyllus Gray (1867), Ingallia Gray (1867), Pocillon
Topsent (1891), Jophonopsis Dendy (1924), Burtonella De Laubenfels (1928) and /ophonota
De Laubenfels (1936).
Type species: Halichondria scandens Bowerbank, 1866 (by original desi-
gnation).
Diagnosis: Massive, branching or encrusting Iophonidae with ectosomal ske-
leton of intercrossing or scattered tylotes with spined heads, choanosomal skeleton
consisting of an isodictyal reticulation of smooth or spined styles, arranged singly or
in two’s and three’s, which may or may not be echinated by acanthostyles. Micro-
scleres include bipocilla and palmate anisochelas; toxas are absent (based on VAN
SOEST et al. 1994).
Remarks: The suborder assignment is based on the presence of palmate chelas
(as opposed to the "tridentate" chelas of the suborder Myxillina). The anisochelas are
shared with Melonchela, Acanthorhabdus (see below) and outside the family with the
mycalid genus Mycale s.l. The apparent absence of toxas is unusual for the family,
shared only with Acanthorhabdus. The genus is traditionally assigned to Myxillidae
(now in suborder Myxillina), and indeed the skeletal structure of many species is
reminiscent of Myxilla. However, Myxillidae in the restricted sense of HAJDU er al.
(1994) possess anchorate chelas and sigmas (cf. below). The "renieroid" or isotropic
skeletal structure is found in many Poecilosclerida, e.g. Jophon, Acarnus, Myxilla,
Lissodendoryx, Antho, Clathria, Ectyoplasia and Plocamionida. These are not all
closely related, and accordingly we assume this character to have been developed
several times in the evolution of the Poecilosclerida. A further argument for including
lophon in the Myxillidae would be the fact that ectosomal megascleres in Myxilla
often are tylote-like and quite often have some spines on their apices. Again, this
feature is not restricted to Myxilla and /ophon, but occurs in several other genera, e.g.
Tedania and Ectyodoryx. In all these cases the tylotes are not exactly similar to the
uniformly shaped /ophon tylotes, by being either anisotornotes (Myxilla, Ectyodoryx)
or very lightly spined (Tedania (Tedania)). It is assumed again that some parallel
evolution in the ectosomal megascleres has taken place (see a more extensive
discussion in HAJDU ef al., 1994).
It can be argued that the genus name /ophon is threatened by Menyllus Gray,
1867. Gray described the latter in the same publication, but one page earlier (page
533) than Jophon (page 534). Menyllus has not been used regularly (in fact there are
only half a dozen records of species as Menyllus in the past 50 years), and the name
was considered as a nomen oblitum under the rules of a previous edition of the
8 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
International Code of Zoological Nomenclature. However, since this rule has been
abandoned, we can only refer to Article 24 which covers the actions of the "first
revisor’. From among the various names available for this group of sponges, RIDLEY
(1881) chose Alebion (for his species A. proximum). RIDLEY (1881) may be
considered as the first revisor in the sense of Article 24, and his choice supersedes any
page priority considerations. Unfortunately, Alebion (also used by VOSMAER, 1882)
was found to be preoccupied, and subsequently RIDLEY & DENDY (1886) chose
Tophon as the name for this group. This is here explained as according to the principle
of Article 24, despite the recommendation 24A which urges the first revisor to take
what is described first. Gray's diagnosis of Jophon and Menyllus were sufficiently
different to explain, why RIDLEY & DENDY (1886) did not consider both synonymous,
as it turned out to be later. In conclusion, /ophon seems to be a valid name, with
Menyllus as a junior synonym.
Tophon species fall into two groups, those with and those without a special
category of fully spined echinating acanthostyles, and these have been given separate
generic status by DENDY (1924), with Jophon reserved for species with acanthostyles
(as in the type species /. hyndmani (Bowerbank, 1858), senior synonym of Halichon-
dria scandens Bowerbank, 1866) and /ophonopsis for species without them (as in the
type species /. nigricans (Bowerbank, 1858)). This distinction has not been accepted
by most authors, because in other Poecilosclerid groups the same presence or absence
is found (e.g. Clathria versus /sociella for example), and accordingly we consider
both as synonyms. Several authors, e.g. BAKUS (1966), went even further and
accepted presence or absence of acanthostyles not even as a distinction between spe-
cies. As will be shown below, this is partly true: in Jophon proximum the recognition
of smaller fully spined acanthostyles is often difficult to make. Bakus (l.c.) synony-
mized a long list of /ophon species, including /. proximum, under a I. pattersoni
(Bowerbank, 1866), now recognized as a junior synonym of /. nigricans (Bowerbank,
1858). However, in the sympatric Eastern Atlantic /. hyndmani and I. nigricans the
acanthostyle presence coincides with other morphological features, so in these it is a
species character.
lophon species are characteristic for cold water faunas (cf. VAN SOEST, 1994),
as they are common in the Northern Atlantic, North Pacific and Arctic, as well as in
the Southern Ocean, including New Zealand, South America and Antarctica. In
tropical and subtropical waters few species have been found, and then almost exclu-
sively in deeper waters.
Iophon proximum (Ridley, 1881) (Figs. 1-12)
Alebion proximum RIDLEY, 1881:119, pl. x, fig. 8.
? Iophon proximum var. reticularis HENTSCHEL, 1914: 89.
Tophon proximum BURTON, 1932:296, pl. 57, figs. 1-13, figs. 21-24, [ in part]; 1934: 25;
1978: 49; BOURY-ESNAULT & VAN BEVEREN 1982: 89, pl. 15, fig. 59, text figs. 25a-h; GENZANO
et al., 1991: 67, lam. 7, fig. B; CUARTAS, 1992: 78, text fig. 24;
lophon pattersoni sensu RIDLEY & DENDY, 1887: 117 & sensu THIELE, 1905 (in part)
[not /. pattersoni (Bowerbank, 1866) = 1. nigricans (Bowerbank, 1858)]
S-E PACIFIC SPECIES OF POECILOSCLERIDA 9
MATERIAL STUDIED: BMNH1879:12:27:5, holotype of /ophon proximum (Ridley, 1881),
Strait of Magellan, Sandy Point, substrate: one valva of Pecten, Coll. R. Coppinger “HMS”
Alert, 12-15 m; ZMH S2313, holotype of /. proximum reticularis Hentschel, 1914, Gauss Stn.,
66°02’S 89°38’W, Deutschen Südpolar-Expedition 1901-1903, 50-200 m.; zMB 3299, /ophon
pattersoni sensu Thiele, 1905, Calbuco, Punta Arenas.
Material studied for comparison: BMNH 1887:5:2:116, holotype of /ophon chelifer
Ridley & Dendy, 1886, Cape of Good Hope, 35°04'S 18°37'E, Challenger Coll., 12.1873, 274
m; cnm 1900-0266, specimen of /. chelifer sensu LAMBE, 1893, British Columbia, Discovery
Passage, 23.06.1885, 20-25 m
A. Encrusting form, on Zygochlamys patagonica:
MHNG 18781, 19207-19212: 18790-18792, HE 22, 23, 33, 34, 36, 58, 59, Bahia Inutil,
53°38’S 72°46’ W, 10-19.09.1972, 32-59 m; MHNG 18782, 18871, 19214: HE 20d, 52, 81c, Seno
de Otway, 53°00’S 71°30°W, 17.09.1972, 3-5 m; MHNG 18791: HE 62, Punta Guale, 53°04’S
73°03’W, 24.09.1972, 70 m, on Chaetopterus sp.; MHNG 18792: VALD. 6.la, Quintupeu,
42°10’S 72°24’W, 15.08.1972, 15 m.
B. Irregular form:
MHNG 18793, 19215, 19216: Mont. 4, 6, 20; San Antonio, 32°42’S 71°23’W,
09.07.1990, 5-10 m; MHNG 18794, 19217: MonT. 22, 27, El Quisco, 32°24’S 71°42’W,
09.07.1990, 10-15 m; MHNG 18795: VALD. 60.11, Estero Castro, 42°30’°S 73°46’W, 03.1987, 10
m; MHNG 18796, 19218: VALD. 27.1, 45, Quintupeu, 42°10°S 72°24’W, 26.04.1971, 10-15 m;
MENG 18797, 19218, 10339: PULL.1, E.85.2, Por.5, Pullinque, 39°33’S 72°11’W, 03. 1983, 5-7
m; MHNG 18798: CO.93.40, Pelluco, 41°30S 72°54’ W, 20.12.1992, 15 m.
C. Massive form:
MHNG 18799: HE. 2, Isla Madre de Dios, 50°15’S 75°05’W, 07.10.1972, 5-25 m; MHNG
18800, 19221: Mont. 2, 10a, San Antonio, 32°42’S 71°22’W, 09.07.1990, 5-10 m; MHNG 18801:
CO.93.6, Bahia Hueihue, 41°54’S 73°32’ W, 09.10.1972, 5-7 m; MHNG 18802: Co.93.19, Pelluco,
42°30°S 72°54’W, 12.12.1992, 15 m; MHNG 18803, 19222: VALD. 4, 37a, Golfo de Quetalmahue,
41°52°S 73°52’W, 05-06.01.1972, 10-15 m; MHNG 18804, 19223: VALD. 8.1, 50.2, Islote Pelù,
41°38°S 73°43°W,18.03.1972, 15 m; MHNG 18805: VALD. 15.3, Quintupeu, 42°10°S 72°24’W,
16.07.1972, 15 m; MHNG 18806: VALD. 31, Bahia Linao, 41°57’S 73°33’W, 28.06.1971, 10-15 m;
MHNG 18807: VALD. 58, Estero Quitralco, 45°43’S 73°25’ W, 07.07.1972, 10 m.
DESCRIPTION
Several specimens (figs. 1-6) and fragments. We differentiate three colour/
growth forms of these species: form A, encrusting on Zygochlamys patagonica or tubes
of polychaetes, with a finely corrugated and punctate surface, colour in alcohol gray,
size: 41-63 x 33-48 x 3-7 mm; form B, irregular often digitate, with finely corrugated
surface, colour in alcohol blackish, size: 30-74 x 31-76 x 22-48 mm; and form C,
massive, with oscules in a row, with a finely conulose surface, colour in alcohol brown,
size: 60-72 x 14-47 x 22-62 mm. Some specimens are agglutinating chitinous tubes or
calcareous fragments. No detachable surface membrane is visible. Small (0.5-1 mm)
oscules occur on conules or are scattered on the surface.
Consistency: Crumbly and friable, very fragile or firm, elastic.
Colour: Orange to yellowish alive, light beige, brownish to blackish in alcohol.
The different colours in alcohol may be attributable to artificial discolouring after
collection.
Skeleton: Ectosomal skeleton: bouquets of partly perpendicular, partly tangen-
tial bundles of tylotes, and abundant microscleres.
10 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
TABLE |
Spicule micrometries of Jophon proximum (Ridley, 1881). I. proximum (Ridley, 1881) bmnh
1879:12:27:5 holotype; /. proximum reticularis Hentschel, 1914 zMHs 2313 holotype; /. chelifer
sensu LAMBE, 1893 CNM
19001266 specimen from Elk Bay;
remeasured. /. proximum form A, specimens from Chile.
I. proximum
(Ridley, 1881)
BMNH 1879:12:27:5
Sandy Point
Remeasured
I. proximum reticularis
Hentschel, 1914
Holotype ZMH S2313
66°02°S 89°38°W
Remeasured
Acanthostyles 1
Acanthostyles 2
158 x 9.5
not observed
120-138-148
x 6-7-8
78-101-117
x 5-6
104-152
not observed
146-160
x 9-10
134x6
I. chelifer sensu LAMBE, 1893
CNM1900-0266
British Columbia
Remeasured
I. proximum. Form A
He 22
Bahia Inutil
He 23
Bahia Inutil
He 33
Bahia Inutil
He 34
Bahia Inutil x 6
He 36
Bahia Inutil
He 58
Bahia Inutil
He 59
Bahia Inutil
He 20d
Seno de Otway
262-327
x 13-19
259-292-316
x8
106-119-125
x6
83 x5
112-123-138
x 5-6-8
100 x 6
141-153-163
x 6-8-10
96 x 8
141-155-176
x 3-5
91-98 x 5-6
150-162-173
x 6-10
105 x 6
154-164-176
6-9-10
96-98-102
x 5
141-155-166
x 6-10
96 x 5
125-135-166
x 5-6-10
99 x 6
Tylotes
1585029
140-147-160
x 4
120-160
not observed
147-163-176
x 6
220-280
x 6-8
243-261-283
x 8
109-128-138
x 4-5
122-134-144
x 3-5-6
170-186-208
x 6
138-163-179
144-179-195
x 6
144-164-182
x 5-6
147-163-186
x 3-5-6
138-146-157
x 4
Anisochelas
25
23-24-27
16-18-20
18-24
16-21-26
29-36
29-36
32-34-38
13-17-19
11-14-18
13-14-18
19-22
10-14-19
13-18-22
22-23-26
16-20
13-15-18
from. literature and
Bipocilia
10.5
Absent
Absent
13-18
14-16-18
11-12-13
10-12-13
6-10-13
11-12-16
Absent
6-10-13
13
Absent
6-8-10
S-E PACIFIC SPECIES OF POECILOSCLERIDA 11
He 52 134-148-160 131-148-166 16-18-22 10-13-16
Seno de otway x 5-7-10 x 3-4-5
DAI
He 8lc 221-241-253 218-234-253 26-28-32 6-7-10
Seno de Otway x 10-10-13 x 6-8
102-120 x 6
He 62 150-162-176 152-184-203 12-14-16 8
Punta Guale x 6-8-10 x8
90-97-105
XD
Vald. 6.1a 170-193-272 157-168-179 13-19-22 6-9-10
Quintupeu x 6-8-10 x 5-5-6
130 x 6
Choanosomal skeleton: isodictyal to subisodictyal unispicular reticulation of
acanthostyles 1. Acanthostyles 2 (echinating acanthostyles) occur at the nodes; they may
be very rare to moderately abundant. Bipocilla and anisochelas are also variable in
abundance.
Spicules: (Table 1-3) Megascleres: Acanthostyles 1 (fig. 7) straight or slightly
bent, covered by numerous small spines, more abundant at the base, diminishing
regularly towards the smooth apex, 106-272 x 5-16 um.
Acanthostyles 2 (fig. 8) echinating, straight and entirely spined, 83-157 x 5-6
um.
Tylotes (fig. 9) straight, smooth except for the spines on the tyles, 109-252 x 3-
10 um
Microscleres: Spurred anisochelas, there are no separated categories of aniso-
chelas, differing just by the size, 10-35 um (fig. 10, 11) Their presence is variable:
abundant, rare, and sometimes absent (?). They have the same morphology: a straight
shaft, basal part shorter with spur-like structure, and alas diverging widely from shaft.
Bipocilla (fig. 12) compact, cup-shaped, with spines at the swollen extremities,
6-16 um. Their presence is variable: abundant, rare, or extremely rare.
Ecology: Occurring on calcareous, chitinous, cultured oysters or stone sub-
strates, 5-32 m.
Distribution: Chilean coast: 20°S-42°S, Falkland Islands, Kerguelen (?).
Remarks: This species stands out among all Jophon species of the studied area
in possessing a second category of small echinating acanthostyles (although they are
not always easy to find, and possibly not always differentiated). This character is
shared with /.chelifer Ridley & Dendy (1886) which may account for BURTON's
(1932) synonymy assignment. We distinguished three colour/growth forms in the
material at hand, but there is no matching character in the skeletal structure. All three
forms have essentially similar skeletons notwithstanding a wide individual variation
in spicule sizes and sometimes in the differentiation of two distinct acanthostyle
categories. Future in situ studies with live specimens might reveal other more subtle
differences between the three. Previous authors, starting with RIDLEY & DENDY (1887
as /. pattersoni) tended to include a wide range of specimens and species from all
12 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
TABLE 2
Spicule micrometries of Jophon proximum form B, specimens from Chile.
I. proximum. Form B Acanthostyles 1 Tylotes Anisochelas Bipocilla
Acanthostyles 2
Mont. 4 141-153-163 147-168-198 19-21-22 13-14-16
San Antonio x 6-8-10 x 6
Absent
Mont. 6 131-148-163 163-172-182 13-14-16 10-12-13
San Antonio x 6-8-10 x 6
Absent
Mont. 20 176-197-227 189-203-218 13-23-35 8-9-11
San Antonio x 6-11-13 x 6-10
Absent
Mont. 22 182-207-230 198-213-227 13-23-35 6-8-10
El Quisco x 6-11-13 x6
Absent
Mont. 27 147-160-189 145-167-182 16-17-19 10-12-13
El Quisco x 10 x6
Absent
Vald. 60.1 195-208-224 160-182-198 16-19-24 10-12-13
Estero castro x6 3
Absent
Vald. 27.1 157-174-186 170-185-208 16-27-32 10-12-13
Quintupeu x 10-12 x 6
Absent
Vald. 45 150-172-186 157-174-186 16-23-32 6-9-10
Quintupeu x 6-9-10 x 6
Absent
Pull. 1 176-186-198 157-172-182 13-15-16 10-12-13
Pullinque x 8-10 x 3-5-6
Absent
E.85.2 141-169-192 147-167-186 13-15-16 10-12-13
Pullinque x 5-6 x 3-4-5
Absent
Por. 5 150-163-173 147-164-176 13-15-16 10-12-13
Pullinque x 6 x 3-5
Absent
Co.93.40 179-198-222 191-204-218 20-26-35 8-10-12
Pelluco x 12-13 x 8
Absent
over the world under this species name. We were unable to examine every single
assignment, but nevertheless made an attempt to evaluate this alleged cosmopolitan
distribution. The correct and doubtful assignments are listed above in the synonymy.
Doubts remain over the identity of specimens assigned to /. proximum by BURTON
(1938, material not examined), DESQUEYROUX (1972, 1976, material presently
unavailable for reexamination), the Brazil specimen of BOURY-ESNAULT (1973) and
the Kerguelen specimens of HENTSCHEL (1914) and BOURY-ESNAULT & VAN BEVEREN
S-E PACIFIC SPECIES OF POECILOSCLERIDA 13
TABLE 3
Spicule micrometries of Jophon proximum form C, specimens from Chile.
Bipocilla
I. proximum. Form C Acanthostyles 1 Tylotes Anisochelas
Acanthostyles 2
He 2 128-140-152 148-164-182 16-19-22 10-14-16
I. Madre de Dios x 6-7-12 x 4-5-8
Absent
Mont. 2 144-156-166 157-170-179 13-18-22 6-9-10
San Antonio x 6-8-10 x 6
Absent
Mont. 10a 144-160-173 157-171-182 12-14-16 10-11-13
San Antonio x 6-8-10 x 5-6
Absent
Co 93.6 144-169-192 150-171-182 16-19-22 10-12-13
Bahia Heihue x 5-7-10 x 3-5-6
Absent
Co 93.19 166-180-192 138-175-195 16-22-26 10-12-13
Pelluco x 5-6 x 3-5-6
110 x 10
Vald. 4 144-160-173 147-162-173 19-22-26 10-12-13
Golfo Quetalmahue x 5-6 x 3-5
Absent
Vald. 8.1 186-201-218 179-199-227 14-16-22 10-12-13
Islote Pelt x 10-12-16 x 6-6-10
93-106
x 5-6
Vald. 50.2 186-202-218 198-210-221 13-18-22 8-10-12
Islote Pelu x 13-14-16 x 6-8-10
Absent
Vald. 15.3 160-173-189 170-196-202 19-22-26 10-12-13
Quintupeu x 10-11-13 x 5-6
90-106
x 5-6
Vald. 31 179-177-211 173-185-202 16-22-26 10-10-11
Bahia Linao x 5-8-10 x 5-6
Absent
Vald. 37a 154-166-182 154-166-179 16-19-22 10-11-13
Golfo Quetalmahue x 3-6 x 3-6
Absent
Vald. 58 182-201-227 195-208-230 16-22-31 6-8-10
Estero Quitralco x 10-11-13 x 6-8
109-157 x 6
(1982). For the Kerguelen material the name /. reticularis is available if it would turn
out to be a separate species.
In our opinion, the following specimens and species were associated incor-
rectly with /. proximum:
Iophon pattersoni (Bowerbank,1866) is originally described from the British
Isles. It is undoubtedly a junior synonym of /. nigricans (Bowerbank, 1858). This is a
14 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Fics 1-12
lophon proximum (Ridley, 1881). 1, 4, form A, encrusting specimen on Zygochlamys
patagonica, from Bahia Inütil and enlarged view of the surface. 2, 5, form B, irregular,
specimen from Pullinque and enlarged view of the surface. 3, 6, form C, massive specimen
from Isla Madre de Dios and enlarged view of the surface. 7, acanthostyles 1, straight or
slightly bent. 8, acanthostyles 2, or echinating styles. 9, Tylotes, straight, smooth with spined
tyles. 10, spurred anisochela of big size. 11, spurred anisochela of small size. 12, compact, cup-
shaped bipocilla. Scales: fig. 7 = 20um; fig. 8 = 10um; fig. 9 = Sum; fig. 10-12 = 2um.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 15
very common species in Western Europe. It differs from /. proximum in having no
category of smaller echinating acanthostyles, which are also considerable thinner.
THIELE's (1905) specimens labelled I. pattersoni are a mixture of /. proximum and
chilense n. sp. (cf. below). /. pattersoni sensu Bakus (1966) probably belongs to the
same, so far not further identified, species as /.chelifer sensu LAMBE, 1893 (cf.
below).
I. chelifer Ridley & Dendy, 1886 is originally described from South Africa. It
was assigned to /. proximum by BURTON (1932). We examined the type specimen. It
is similar to /. proximum but differs clearly in the "cut-off" shape of the tylote heads
which are also much lighter spined (only on the flattened apex) and in the clearly
longer and thicker (360-420 x 16-20 um) and more heavily spined structural acan-
thostyles. /. chelifer sensu LAMBE,1893 from British Columbia, that we examined,
differs from /. proximum in having two clearly separated size categories of bipocilla
(not mentioned by LAMBE), the tylotes are also lightly spined on the apices only and
the structural acanthostyles are more heavily spined and thicker. It is likely that
Lambe's material belongs to one of the Arctic Jophon species (cf. KOLTUN, 1959), and
that Bakus' material of /. pattersoni belongs to that same species. /. chelifer sensu
Thiele, 1905 belongs to the new species described below; it does not have the small
acanthostyles. /. chelifer ostiamagna Wilson, 1904, assigned to /. proximum by
Burton (1932) is a separate species of Jophon. I. chelifer californiana De Laubenfels,
1932 belongs to /. lamella Wilson, 1904 (cf. below). New Zealand specimens
assigned to /. proximum by BERGQUIST & FROMONT (1988) are unlikely to belong to
the present species; for them the name /. semispinosus Bergquist (1961) is available.
The description of /. proximum sensu URIZ (1987, 1989) from South West Africa
reminds strongly of /. chelifer, while her /. chelifer differs from /. chelifer proper in
having two distinct bipocilla sizes; the latter material is likely an undescribed species.
I. lamella, I. lamella indivisus, and I. indentatus, all described by WILSON
(1904) were assigned to /. proximum by BURTON, 1932, but are here considered a
separate species. The main differences are the shape of the bipocilla and the absence
of small acanthostyles.
Iophon lamella Wilson, 1904 (figs. 13-18)
lophon lamella WILSON, 1904:146, pl. 20, figs. 3, 7-9, 12, 13, pl. 24, figs. 2-4.
lophon indentatus WILSON, 1904: 151, pl. 19, fig. 6, pl. 20, figs. 1, 5, 6, pl. 23, fig. 4;
Hophon indentatus WILSON sensu DICKINSON, 1945: 16, pl. 21, figs. 41, 42, pl. 22, fig. 43.
Burtonella melanokhemia DE LAUBENFELS, 1928: 361, fig. 1.
lophon chelifer var. californiana DE LAUBENFELS, 1932: 82, fig. 47.
MATERIAL STUDIED: USNM 8277, holotype of Jophon lamella Wilson, 1904, Albatross
Stn. 3405, 00°57°S 89°38’W, 95 m ; USNM 8279, syntype of Jophon indentatus Wilson, 1904,
Albatross Stn. 3405, 00°57°S 89°38’W, 95 m; USNM 21401, holotype of Jophon chelifer var.
californiana De Laubenfels, 1932, S of San Pedro, California, 27.12.1916, 48 m; USNM 21369,
holotype of Burtonella melanokhemia De Laubenfels, 1928, California; USNM 39352, ZMA
10977, MHNG 18808 Ga.IIT.28, fragment, SEPBOP "Anton Bruun" Exped. Cruise 18B 791C,
Galapagos Islands, Santa Cruz Island, 00°26'S 90°20'W, 21.09.1966, 95 m.
16 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
TABLE 4
Spicule micrometries of: Jophon lamella Wilson, 1904. J. lamella Wilson, 1904 USNM 8277
holotype; /. indentatus Wilson, 1904 USNM 8279 syntype; /. chelifer var. californiana de Lau-
benfels, 1932 USNM 21401 holotype; Burtonella melanokhemia de Laubenfels, 1928 USNM
21369 holotype; from literature and remeasured. Specimen from Galapagos of /. lamella
Wilson, 1904.
Acanthostyles 1 Tylotes
Tophon lamella
Anisochelas 1 Bipocilla
Wilson, 1904 Anisochelas 2
Holotype USNM 8277 210-220 220-240 28 12-16
00°57’S 89°38°W x 12-16 x 7-8 14
Remeasured 210-222-234 218-223-235 10-14 8-13-16
x 12-13-15 x 7-8 20-26
lophon indentatus
Wilson, 1904
Syntype USNM8279 220 x 14-16 220 x 8 14 8
00°57°S 89°38W
Remeasured 199-218-234 207-219-220 16 8
x 16 x 7-8
I. chelifer var.
californiana
de Laubenfels, 1932 265-290 240-250 not observed 15
Holotype USNM 21401 x 12-13 x 6-8 15-33
South San Pedro
Remeasured 234-248-260 200-223-241 27-31-35 12-17-20
312 x 7-8 12-15-23
Burtonella
melanokhemia
de Laubenfels, 1928
Holotype USNM 21369 210 x 13 180 x 10 12-40 as sigmas:
California 50
Remeasured 270-282-304 222-238-253 20-21-25 12-15-16
x 16-18-20 x 7-8 12-14-16
Galll. 28 Sepbop 214-229-250 202-226-240 22-29 10-14-16
18B791C x 10-16 x 6 13-16
00°26°S90°20’W.
DESCRIPTION
A mass of fragments of small, irregular lamellate shapes; the largest is 5 x 6
cm and 5-15 mm thick (fig.13). The surface is differentiated in a more or less
undulated and punctate side and an opposite side with grooves covered by a thin,
easıly removed membrane.
Consistency: Firm but fragile.
Colour: Yellow or beige-brown in alcohol.
Skeleton. Ectosomal skeleton: a palisade of scattered tylotes.
Choanosomal skeleton: tight-meshed isotropical reticulation of spined styles
and tylotes. Microscleres abundant.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 167
Fics 13-18
lophon lamella Wilson, 1904. 13, lamellate specimen from Galapagos. 14, acanthostyle 1
heavily spined. 15, smooth tylotes, swollen apices, minutely spined. 16, 17, spurred anisochela
1 and 2, both with straight shaft. 18, strongly asymmetrical trilobate bipocilla with finger like
appendages. Scales: fig. 14 = 10um; fig. 15, 16 = Sum; fig. 17, 18 = 2um.
Spicules (Table 4): Megascleres: Acanthostyles I (fig. 14) heavily spined all
over the surface 214-250 x 10-16 um. No acanthostyles 2. Tylotes (fig. 15) with a
straight and smooth shaft and swollen apices, minutely but heavily spined, 202-240 x
6 um.
Microscleres: Spurred anisochelas 1 (fig. 16) 22-29 um and 2 (fig. 17) 13-16
um, both with the same morphology, straight shaft and the alas diverging from the
shaft. Bipocilla (fig. 18) strongly curved, and with one of the extremities expanded
and trilobate and the other with several finger like appendages, 10-14 um. Both types
of microscleres are abundant.
Ecology: Substrate, pebbles, 90-140 m.
Distribution: Galapagos Islands, West coast of Central America (DICKINSON,
1945) Pacific coast of Southern California, Baja California.
Remarks: This species differs from /. proximum in the lamellate shape and the
yellow-brown colour, in the possession of two distinct categories of anisochelas, and
in the much more openly curved trilobate bipocilla. These characters are shared with
Burtonella melanokhemia De Laubenfels and /. chelifer californiana De Laubenfels
18 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
and accordingly these are considered junior synonyms. Wilson's specimens of /.
lamella, I. lamella indivisus and I. indentatus differ somewhat in shape, but their
skeletons, spicule categories and bipocilla shape are essentially similar, so these are
united under the present species. DICKINSON's (1945) description of /. indentatus from
California differs substantially from that of /. lamella and I. indentatus, especially in
spicule size. Perhaps it belongs to the same species as the Lambe and Bakus speci-
mens, but study of Dickinson material is necessary for that conclusion.
Despite its lamellate habit, /. chelifer ostiamagna Wilson (1904) is clearly
separate from /.lamella in having much larger and lighter spined styles, only a single
category of anisochelas and two categories of bipocilla.
Iophon chilense n. sp. (figs. 19-26)
Tophon chelifer sensu THIELE, 1905: 445, figs 63a-d [non: /. chelifer Ridley & Dendy, 1886]
Iophon pattersoni sensu THIELE, 1905: 445 (in part) [non: /. pattersoni (Bowerbank, 1866)= 1.
nigricans (Bowerbank, 1864)]
MATERIAL STUDIED: HOLOTYPE: MHNG 18810, He 108, Somerset Canal, 47°58’S 74°35°W,
02.10.1972, 260 m.
Paratype: MHNG 18809, HE 81 Seno de Otway, 53° 00°S 71° 30°W, 17.09.1972, 250 m.
Material studied for comparison: ZMB 3300, Thiele's specimen of /ophon chelifer sensu
THIELE, 1905, Calbuco; ZMB 3299, Thiele's specimen of /ophon pattersoni sensu THIELE, 1905,
Calbuco.
DESCRIPTION
Sponge irregularly encrusting on calcareous substrates and tubes of poly-
chaetes, or massive, irregular (figs. 19, 20). Size: 35-46 x 25-39 mm. Surface mem-
brane thin, not easily removable, punctate.
Consistency: Crumbly, soft, very fragile.
Colour: Yellowish to orange alive, light to dark brown in alcohol.
Skeleton: Ectosomal, a palisade of bundles of tylotes and abundant micro-
scleres.
Choanosomal: Loose subisodictyal to confused reticulation of acanthostyles 1,
isolated or in occasional bundles. Abundant anisochelas
Spicules: (Table 5) Megascleres: Acanthostyles 1 (fig. 21) slightly bent, most
are smooth except for spines at the base, some have the shaft abundantly microspined,
257-308 x 8-16 um. No acanthostyles 2.
Tylotes (fig. 22, 23) straight, with swollen microspined apices, 207-259 x 5-8
um.
Spurred anisochelas | (fig. 24), curved, 18-29 um, and 2 (fig. 25) 10-16 um.
Bipocilla (fig. 26) one of the ends trifoliate, the other with claw-like
appendages, 10-16 um.
Etymology: named after its geographic distribution.
Ecology: Occurring on calcareous, polychetes tubes, and shells, 250-260 m.
Distribution: Chilean coast, 47°S.- 53°S.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 19
TABLE 5
Spicule micrometries of Jophon chilense n. sp. I. chelifer sensu THIELE, 1905, zMB 3300, specimen
from Plate collection; from literature and remeasured. Specimens from Chile of /. chilense n. sp.
I. chilense n. sp. Acanthostyles | Tylotes Anisochelas 1 Bipocilla
Anisochelas 2
I. chelifer sensu
Thiele, 1905
ZMB3300 20x15 ZIO xa 16-28 15
Calbuco
Remeasured 208-235-253 186-199-208 26-27-29 13-14-16
x 13-15-16 x 6 13-17-22
He 81 259-274-284 202-224-235 26-27-29 13-14-16
Seno De Otway x 8-16 x 5-6 13-16
Holotype MNHG 18810 = 257-285-308 207-249-259 18-20-21 10-12-13
He 108 x 8-16 x 5-6-8 10-12-13
Remarks: This species differs from /. chelifer sensu RIDLEY & DENDY, 1886 in
the absence of small acanthostyles, the almost smooth relatively short structural
acanthostyles, and the much heavier spined tylote heads. From /. proximum and 1.
lamella it differs also in possessing almost smooth styles, furthermore its bipocilla are
clearly of a different shape as those of /. proximum and show teeth different from
those of I. lamella. There are no matching descriptions from the studied area.
Iophon timidum n.sp. (Figs 27-32)
MATERIAL STUDIED: HOLOTYPE: MHNG 18811: HE 12, Puerto Caracciolo, 50°28’S
Togli WA OO 1019725 25 mi:
PARATYPES: MHNG 18812: He 28; MHNG 19224: HE 56, same data as the holotype.
MATERIAL STUDIED FOR COMPARISON: MNHNDT 1976, specimen 421, slide of the syntype of
lophon radiatus Topsent, 1901, 71°19’S 87°37’ W, 450 m.
DESCRIPTION
Sponge massive (figs. 27, 28) spherical to oval, size 55-95 x 55-60 mm.
Surface corrugated, irregular and covered by a thin membrane, which is easily
removed. A few small oscules (1-2 mm) are irregularly distributed.
Consistency: Soft, extremely fragile, easily torn.
Colour: Beige to dark brown alive, same colour in alcohol.
Skeleton: Ectosomal skeleton: a palisade of tylotes, free or in bundles. Normal
anisochelas are abundant.
20 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Fics 19-26
lophon chilense n.sp. 19 [HOLOTYPE, MNHG 18810] irregular encrusting specimen from
Sommerset Canal. 20, enlarged view of the surface. 21, acanthostyles 1 slightly bent, smooth,
with microspined shaft. 22, 23, straight tylotes, swollen microspined apices. 24, 25, spurred
anisochela | and 2 with curved shaft. 26, bipocilla trifoliate, with claw-like appendages. Scales:
fig. 21 = 10um; fig. 22, 23 = Sum; fig. 24, 26 = 2pm.
Choanosomal skeleton: a loose and irregular reticulation of styles, which
occasionally form longitudinal tracts connected by single styles. Anisochelas and
pigment are abundant.
Spicules (Table 6): Megascleres: Smooth styles (fig. 29) with a few spines at
the base and slightly bent, 186-259 x 3-6 um. No echinating acanthostyles.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 21
TABLE 6
Spicule micrometries of Jophon timidum n. sp. I. radiatus Topsent, 1901 LBIMDT 1976 syntypes;
from literature and remeasured. Specimens from Chile of /. timidum n. sp.
Tylotes
Anisochelas
I. timidum n. sp. Styles
Bipocilla
I. radiatus
Topsent, 1901
Syntypes LB/MDT 1976
specimen n° 306 SSUAO 300-350 17-60
71°14’S 89°14°W x 5-6 8-12
specimen n° 421 570-580 390 17/53-70
71919S 87°37 W x 20 x 8-9 8-16
Remeasured 502-552-607 340-368-413 57-61-73
specimen n° 421 x 16-19-24 x8 8-16
Anisochelas 1
Anisochelas 2
HOLOTYPE MNHG 18811
He 12 186-221-256 150-196-250 11-14
Puerto Caracciolo x 3-6 x 3-6 5-6
He 28 202-234-259 154-184-243 10-16
Puerto Caracciolo x 3-6 x 3-5 6-10
He 56 202-226-259 170-204-250 13-16
Puerto Caracciolo x 3-6 x 3-5 6-10
Tylotes (fig. 30) thin, long and fusiform, apices swollen and strongly spined
150-250 x 3-6 um.
Microscleres: Spurred anisochelas 1 (fig. 31) with a straight shaft, alas widely
diverging from the shaft, 10-16 um. Anisochelas 2 (fig. 32) with roundish concave
extremities and a short axis, rare, resembling bipocilla but provided with a distinct
spur, 5-10 um.
No bipocilla were found.
Etymology: the name refers to the shape of the anisochela 1, which looks as if
it is contracted with fear.
Ecology: Occuring on stones, sand, 25 m.
Distribution: Chilean coast, 50°S 75°W.
Remarks: The new species is based on the combination of (almost) smooth
styles and the second category of incurved anisochelas. Since all three specimens
were exactly similar in this respect, it is clear these are stable characters. The absence
of bipocilla is unusual, but reported occasionally from other Jophon species (cf.
KOLTUN, 1959) and specimens. (e. g. /. reticularis Hentschel, 1914 from Gauss Stn.,
that we examined). It is not a reliable character for species distinction.
DD RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
The smooth styles and "loose" anisotropic reticulation are shared with 7.
radiatus Topsent, 1901, originally described from the Antarctic region, but subse-
quently reported from along the Atlantic and Pacific coasts of South America up to
37°S. Our specimens have been compared with a type slide of /. radiatus and the two
were found to have some clear differences, the most important being the much larger
styles of /. radiatus, the much larger anisochelas (up to 60 um) (neither category of
anisochela is malformed) and the possession of 2 sizes of normal formed bipocillas.
Unfortunately, the specimens of /. radiatus recorded from the study area (DESQUEY-
ROUX, 1972) are not available for re-examination, so we cannot confirm the presence of
true /. radiatus in our area. However, the spicule sizes reported by Desqueyroux clearly
exceed those of the new species, so conspecificity with the present species is unlikely.
Iophon tubiforme n. sp. (Figs. 33-38)
MATERIAL STUDIED: HOLOTYPE: MHNG 18813: He 7, Isla Newton, Rada Shinglet,
51°51°S 73°42’ W, 01.10.1972, 5-10 m.
PARATYPES: MHNG 18814: HE 11, Isla Newton, Rada Shinglet, 51°51’S 73°42’W,
01.10.1972, 5-10 m.; MHNG 18815: He 20, Seno de Otway, 53°00’S 71°30’W, 17.09.1972, 35
m; MHNG 18816: HE 21, MHNG 19225: HE 26, Caleta Chica, Faro San Pedro, 47°47’S 74°54’ W,
01.10.1972, 30 m, 12 m; MHNG 18817, 18992: He 32, Co.93.31, Puerto Caracciolo, 50°26’S
75°09’W, 09.10.1972, 25 m; MHNG 18818: HE 86, Islas Grupo Porvenir, Canal Zenteno,
52°49’S 73°40’W, 11.10.1972, 30 m; MHNG 18819: HE 103, Punta Guale, 53°04’S 73°03’W,
24.09.1972, 70 m; MHNG 18820: VALD. 47, Laguna Cachana, 45°58’S 73°45’W, 28.08.10971,
10 m.
MATERIAL STUDIED FOR COMPARISON:
lophon unicornis Topsent,1907, LBIMDT 1665 type, specimen 798, Ile Anvers,
Antarctic, dragage, 06.01.1905, 25 m.
lophon spatulatus Kirkpatrick, 1907, SYNTYPES, BMNH 1908:2:5:144, 144a,b, 145, 145a,
Coulman Island, 1828 m, "Discovery" Coll.
DESCRIPTION
Several large specimens formed by up to 7 tubes, 81-87 mm high, of uniform
diameter, 18-31 mm, issuing from a common base: 87-111 mm (fig. 33, 34). Tubes
branch off into new tubes, 18-19 mm diameter and 32-62 mm high, all of the tubes
possess a terminal oscule of 6 to 14 mm in diameter. The oscular canal is as long as
the tube itself, tube wall is 3-10 mm large. Surface is irregular but smooth and
covered by a membrane overlying large subectosomal cavities.
Consistency: Soft and easily torn.
Colour: Brown alive, light to dark brown in alcohol.
Skeleton: Ectosomal skeleton: dense brushes consisting of bundles of tylotes
and free tylotes. Ectosomal brushes are pointed outwards, between them is a
subdermal lacunae system covered by the aspicular membrane.
Choanosomal skeleton: plumose, formed by longitudinal tracts of styles (2-4
across) connected by single styles. Numerous juvenile styles are present.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 23
TABLE 7
Spicule micrometries of Jophon tubiforme n. sp. I. unicornis Topsent, 1907 LBIMDT 1665, HOLO-
TYPE; from literature and remeasured. Specimens from Chile of /. tubiforme n. sp.
Styles
I. tubiforme n. sp. Tylotes Anisochelas Bipocilla
I. unicornis
Topsent, 1907
HOLOTYPE LBIMDT 1665 435-470 240-10 18-20 Not observed
Ile Anvers x 15
Remeasured 405-443-486 194-222-243 16-19-24 32
x 12-15-16 x8
HOLOTYPE MNHG 18813
He 7 195-207-218 186-202-221 17-19 8-12-16
Isla Newton x 8-10-11 x 6-8-10
He 11 150-173-192 141-157-170 8-10 6-10-13
Isla Newton x 5-6-10 x 3-5-6
He 20 170-186-195 141-158-166 12-15-16 8-10
Seno de Otway x 6-8 x 5-6
He 21 218-237-272 173-200-218 11-14-16 16-17-19
Caleta Chica x 6-7-10 x 3-6
He 26 170-184-202 134-158-173 13-16 10-16
Caleta Chica xe5=0 x 3-6
He 32 202-234-262 182-199-218 10-11-13 16-18-19
Puerto Caracciolo x 6-10-13 x 3-6
He 86 163-179-189 147-157-166 13-16 6-9-10
Canal Zenteno x 4-6 x3
He 103 218-227-262 211-221-243 10-11-13 13-14-16
Punta Guale x 6 x 3-6
Vald. 47 192-228-262 173-196-211 13-14-16 6-8-10
Laguna Cachana x 10-13 x 5-10
Spicules: (Table 7) Megascleres: mucronate, smooth, oxeote styles (fig. 35)
slightly bent, 150-272 x 5-13 um.
Tylotes (fig. 36) strongylote, fusiform, straight, with barely swollen, strongly
microspined apices 141-243 x 3-10 pm.
Microscleres: Spurred anisochelas in a single size category (fig. 37) common,
with curved shaft 8-19 um.
Bipocilla (fig. 38) uncommon, fragile, terminal apices diverging like small
fingers or teeth, 6-19 um.
Etymology: Refers to the morphology: a mass of tubes
Ecology: Stones, fine sand, mud, 10-70 m.
24 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Fics 27-38. lophon timidum n.sp. 27 [holotype, MNHG 18811] massive specimen from Puerto
Caracciolo. 28, enlarged view of the surface. 29, smooth styles with a few spines at the base. 30,
long, fusiforme tylotes with swollen apices, strongly spined. 31, spurred anisochela 1, with
straight shaft. 32, malformed anisochela 2, rounded concave extremities and short axis. Jophon
tubiforme n.sp. 33 [holotype, MNHG 18816] specimen from Caleta Chica. 34, enlaged view of the
surface. 35, mucronate smooth style. 36, smooth strongylote with fusiforme swollen microspined
apices. 37, spurred anisochela, with curved shaft. 38, bipocilla uncomon with finger like appen-
dages. Scales: fig. 29, 30 = Sum; fig. 31 = 2um; fig. 32 = Ium.; fig. 35, 36 = Sum; fig. 37, 38 =
2um.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 25
Distribution: Chilean coast, 45°S - 53°S.
Remarks: The new species shares with /. timidum the smooth structural
spicules and the loose skeletal reticulation; in this the two differ from all other Jophon
of the study area. Between them they differ clearly in the form of these spicules
(normal styles vs. oxeote/mucronate styles) and also in the form and spination of the
tylote heads.
There is a striking match between the new species and the Antarctic /. uni-
cornis Topsent (1907) in the form of the structural spicules (mucronate oxeotes). We
compared the new species with the description and a slide made of the holotype of /.
unicornis, and found several clear differences: /. unicornis is not tubiform, its
mucronate oxeotes are about twice as long and thick as those of the new species, and
its tylotes have clearly swollen apices. We also studied slides of the type of /.
spatulatus Kirkpatrick (1907) and can confirm that this is conspecific with /. uni-
cornis, aS suggested by BOURY-ESNAULT & VAN BEVEREN (1982). Since TOPSENT's
paper is dated July 1907 and KIRKPATRICK's September 1907, TOPSENT's name has
priority.
I. spatulatus has been recorded from the study area (Isla Huemules) by
DESQUEYROUX (1976) without description; since this material is not available for
study, we can only speculate, that it conformed to our new species.
It is likely that Boury-Esnault's record of /. unicornis from Kerguelen concerns
a separate new species, since its form is cup-shaped. The three mucronate-oxeote
styles bearing species, one Antarctic, one Southern South America, and one Ker-
guelen, probably form a monophyletic group within Jophon.
Iophon ostiamagna Wilson, 1904
lophon chelifer var. ostiamagna WILSON, 1904: 143, pl. 20, figs. 2, 4, 10, 11, pl. 24, fig. 1
MATERIAL STUDIED: HOLOTYPE: USNM 8280, Stn. 3384, 07°31'N 79°14'W, 08.03.1891,
880 m.
Not represented in our material, so only a diagnosis is given:
Lamellate growth form, 5-8 mm in thickness. Skeleton loosely reticulate.
Acanthostyles long and robust, lightly spined, 450 x 20 um. No echinating
acanthostyles. Tylotes with lightly spined heads, 325 x 10 um. Spurred anisochelas in
a single category, 12-20 um. Bipocilla in two categories, 1: 20 pm, 2: 12 pm.
Ecology: Substrate, sand, 880 m.
Distribution: 07°31'N 79°14'W.
Remarks: It is similar to the sympatric /. lamella, but differs in having much
larger and more lightly spined acanthostyles, a single category of anisochelas and two
categories of bipocilla.
Iophon radiatus sensu DESQUEYROUX, 1972
‘Aophon radiatus TOPSENT, 1901: 21, pl. 3, fig. 13.
lophon radiatus DESQUEYROUX, 1972: 23, figs. 70-75.
MATERIAL STUDIED: LBIMDT. 1976, microscopic slide of Topsent's type, specimen 421.
26 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Not represented in our material, so only a diagnosis of Desqueyroux's material,
which is from the study area, is given here:
Massive, with a loosely reticulated skeleton. Long smooth styles, 460-520 x 16
um. No echinating acanthostyles. Tylotes with distinctly spined heads, 280-350 x 9
um. Spurred anisochelas in two widely different size categories, 1: 70um, 2: 14 um.
Bipocilla 6-16 um.
Ecology: Substrate, stones, 600 m.
Distribution: Chilean coast, 20°S.
Remarks: Topsent's material from Antarctica was similar in spicule sizes, and
since the species has also been recorded from the Falkland Islands (BURTON, 1934)
and South Georgia (BURTON, 1940), it may be a wide-spread species.
REVIEW OF THE FAMILY IOPHONIDAE
Shapes in this microcionine family are varied (fistular, encrusting, lobate,
branching, vase-shaped). The spiculation includes those typical for the suborder:
smooth or spined styles, palmate chelas and toxas (although these are absent in
Tophon and Acanthorhabdus); sigmas are absent. Skeletal architecture in non-fistular
forms is variously anisotropic, plumose, or isotropic.
This new concept of a revived family was first introduced by HAJDU ef al.
(1994) and elaborated by VAN SOEST et al. (1994). The following fistular genera
conform to it: Cornulum (with junior synonym Coelosphaerella) Paracornulum
(with junior synonym Cornulacantha) Zyzzya (with junior synonym Damirina)
Acheliderma (with junior synonyms Astylinifer and Fusifer) Damiria, Melonchela,
Anisotylacanthaea and Xytopsene. These genera were previously united in the sub-
family Cornulinae LEvI & LEVI (1983). In addition, four non-fistular genera are con-
sidered to belong in this family, viz. lophon, Acarnus, Megaciella and Acan-
thorhabdus.
In VAN SOEST et al. (1994) the fistular members are described and discussed.
These sponges are not represented so far in the SE Pacific, so for more information on
these one is referred to that paper. The remaining genera are here briefly discussed.
Acarnus Gray, 1867
Synonyms: Fonteia Gray, 1867 and Trefortia Deszö, 1880.
Type species: Acarnus innominatus Gray, 1867: 544 (by monotypy).
Diagnosis (from VAN SOEST et al., 1994): Massive, branching or encrusting
Iophonidae with ectosomal skeleton of scattered tylotes with spined heads,
choanosomal skeleton consisting of an isodictyal, isotropic or anisotropic, reticulation
of styles and cladotylotes, arranged singly or in tracts of several spicules, which may
or may not be echinated by acanthostyles and/or cladotylotes. In thinly encrusting
specimens, the choanosomal skeleton is hymedesmioid, i.e. with styles or cladotylotes
erect on the substrate. The cladotylotes occur normally in two categories, but
S-E PACIFIC SPECIES OF POECILOSCLERIDA DE]
exceptionally in three or a single one. Microscleres include up to three categories of
toxas, and chelas.
Remarks: Acarnus has been revised and monographed recently by VAN SOEST
et al. (1991) (with additional data in HOOPER & LEVI, 1993). A species of that genus
occurring in the region is Acarnus peruanus Van Soest et al., 1991; for a description
one is referred to that paper. An additional specimen of that species is here recorded
from the Galapagos Islands (ZMA data collections).
Genus Megaciella Hallmann, 1920
Type species: Amphilectus pilosus Ridley & Dendy, 1886 (by original
designation).
Diagnosis (from VAN SOEST et al., 1994): Branching or encrusting Iophonidae
with ectosomal brushes of tylotes with spined heads, with a reticulate or confused
choanosomal skeleton consisting of smooth styles with spined or smooth heads,
arranged singly or in two’s or three’s. Thinly encrusting specimens have the styles
erect on the substrate. Microscleres are palmate isochelas and toxas.
Remarks: The type species lacks echinating acanthostyles, but similar species
with acanthostyles exist. There are no species answering to this diagnosis of Mega-
ciella recorded from the SE Pacific. The Californian species Myxichela microtoxa De
Laubenfels, 1935 is a clear Megaciella and “may be nearest representative” of that
genus in the area. i
Genus Acanthorhabdus Burton, 1929
Type species: Acanthorhabdus fragilis Burton, 1929 (by monotypy).
Diagnosis (from VAN SOEST et al., 1994): Iophonidae with modified styles in
the form of anisoxeas often provided with a mucron arranged in a coarse but ill-
defined plumoreticulation. Ectosomal megascleres are acanthorhabds with heavy
spines and microspined tylote apices. Microscleres are spurred palmate anisochelas.
Remarks: BURTON (1929) placed this rather enigmatic monotypical genus in
his section Mycaleae on account of its anisochelae, but the presence of the special
ectosomal megascleres makes it thoroughly untypical of this group. VAN SOEST ef al.
(1994) proposed to assign this species to Iophonidae, because the spurred anisochelae
are shared with /ophon and the acanthorhabds have spined tylote apices found in all
members of the Iophonidae. Although the occurrence of the species is technically
outside the study area, we decided to examine the holotype (BMNH 1926:10:26:189),
to supply more data on this intriguing species: the skeleton consists of massive
plumose columns of smooth oxea-like megascleres, carrying a surface crust of
acanthose spicules. The smooth oxeotes of 550 by 35 um are decidedly asymmetrical,
and often a distinct mucron (not unlike /ophon unicornis) is present at one end;
occasionally there is a single spine at one end. The acanthorhabds of 400 by 25 um
are fusiform, heavily spined, with spines curved towards the middle of the spicules,
28 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
clearly demonstrating their diactinal nature. The apices of the acanthorhabds are both
provided with small spines not unlike the heads of the tylotes of /ophon; it is assumed
- also by their position in the sponge skeleton - that these spicules are homologous to
ectosomal tylotes. The spurred anisochelas of 25-30 um length are very similar to
those of various Jophon species. No further material of this species has become
available.
KEY TO THE SPECIES OF Jophon OCCURING IN THE SE PACIFIC COAST
1 With two size categories of choanosomal styles............... I. proximum
— Sinsleisize category of choanosomal styles CL 2.2 2
2 Siyles’spined, bipocilla'strongly asymmetrical... =.=. CO EEE 3
- Styles smooth bipocilla almost symmatrical or absent. .................. 4
3 Spines on styles regularly distributed. Bipocilla trilobate. Galapagos Islands
RÉ RARA ent DE EEE ENTE DEL PARU EL RU SR ER I. lamella
- Spines on styles irregularly distributed Bipocilla trifoliate. Chilean coast.
WIRE RE EEE deb ue RER eee I. chilense n.sp.
- Bipocilla absent. The small category of anisochelas is malformed
CRE PR I IT I e CEE PLEASE ai) I. timidum n.sp.
- Bipocillaspresent almost symmetrieal® 9 ee zum I. tubiforme n.sp.
Suborder: Myxillina Hajdu, van Soest & Hooper, 1994
Family: Myxillidae Topsent, 1928
Diagnosis: Myxillina with smooth, mucronate or microspined tornote or tylote
ectosomal megascleres arranged as bouquets, with smooth or spined monactinal or
diactinal choanosomal megascleres arranged in isotropic, anisotropic or plumose reti-
culation. Microscleres include anchorate isochelas and/or derivates thereof (spatulate,
unguiferate or birotulate chelas), and smooth sigmas.
Myxilla Schmidt, 1862
Synonyms: Burtonanchora De Laubenfels, 1936; Crellomyxilla Dendy, 1924; Dendoryx
Gray, 1867; Emplocus Gray, 1867; Ectyomyxilla Lundbeck, 1909; Hastatus Vosmaer, 1880;
Stegxella Bowerbank, 1874; Stelodoryx Topsent, 1904; Styloptilon Cabioch, 1968; Tereus
Gray, 1867).
Type species: Halichondria rosacea Lieberkühn, 1859.
Diagnosis: Myxillidae with ectosomal spicules which are variably mucronate,
strongylotylote or tornote, frequently with one or a few prominent spines on the
apices. The choanosomal spicules are stylote, usually styles, with or without
spination. As is usual for the family, the microscleres are anchorate chelas -spatulate
and/or unguiferate- and sigmas, both types frequently in two categories.
Remarks: The suborder assignment is based on the "tridentate" nature of the
chelas. The family assignment is based on the combination of tornotes, anchorate
S-E PACIFIC SPECIES OF POECILOSCLERIDA 29
chelas and true sigmas (other myxilline genera with anchorate chelas do not have
tornotes and proper sigmas, and are united in the family Crambeidae new family
name).
The skeleton of the type species is a tight meshed isotropic reticulation of
strongly spined styles, but other species may have more loosely constructed skeletons
and smooth styles.
The possession of unguiferate chelas instead of or in addition to the spatulate
anchorates occurs in several Myxilla-like sponges as well as in the unrelated genus
Monanchora (cf. VAN SOEST, 1990), but is not considered evidence for a mono-
phyletic group. However, for convenience sake, we propose to distinguish a sub-
generic unit for Myxilla species with these chelas. The name Stelodoryx Topsent,
1904 (type S. procera Topsent, 1904) is available for these sponges.
The possession of smooth instead of spined styles also is not coinciding with
other features and accordingly is not considered evidence for a monophyletic group.
However, for convenience sake, we propose to distinguish a subgeneric unit for
Myxilla species with smooth styles. For these, the name Burtonanchora De Lauben-
fels, 1936 (type Myxilla crucifera Wilson, 1925) is available.
A separate genus Ectyomyxilla Lundbeck, 1909 with echinating strongly spi-
ned styles is likewise not recognized as a monophyletic group, because the presence
of these spicules does not coincide with other morphological features. However, it is
useful as a subgeneric unit. Technically, the name Ectyomyxilla is a nomen nudum
because LUNDBECK (1909) did not mention any species when erecting this genus.
However, since the name was proposed before 1930, the subsequent use of the name
by HENTSCHEL (1914) for E. kerguelensis HENTSCHEL 1914, satisfies Article 13 (1) of
the ICZN code (1985) as subsequent type species designation.
The remaining genera listed as junior synonyms above are synonyms of
Myxilla (Myxilla).
This wide diagnosis of Myxilla encompasses species from all areas of the
world.
Subgenus Myxilla Schmidt, 1862
Diagnosis: Myxilla with thight meshed isotropic skeleton and exclusively
spatulate anchorate chelae; without a special category of echinating acanthostyles.
Myxilla (Myxilla) mexicensis Dickinson, 1945 (Figs. 39-53)
Myxilla mexicensis DICKINSON, 1945: 18, pl. 24, figs. 47, 48, pl. 25, figs. 49, 50.
MATERIAL STUDIED: AHF 9, holotype of Myxilla mexicensis Dickinson, 1945, Stn.
513.36, off San Francisco Island, Gulf of California, 60 m.
USNM 37917, MHNG 18958: Ga.II.3, Stn. 17, Galapagos Islands, Kicker Rock, coll.
W.D.Hope, 02.1978, 23 m; USNM 37918, MHNG 18959: Ga.II.8, Galapagos Islands, Kicker
Rock; USNM 37919, MHNG 18960, ZMA 10978, USNM 39360: Ga.II.4, Stn. 8, Galapagos Islands,
Punta Vicente Roca, Isla Isabela, 02.1978, 18 m; USNM 39360, MHNG 18961: Ga.IV.33, Cruise
18B 794E, Galapagos Islands, Santiago Island, 00°12'S 90°51'W, 23.09.1966, 34 m ; ZMA
10979, Cruise 18B 791C, Galapagos Islands, Santa Cruz Island, 00°26'S 90°20'W, 21.09.1966,
95 m.
30 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
DESCRIPTION
Five specimens (figs 39-42): 45-80-100-120 x 30-60-100-70 mm and 3
fragments: 20-50-80 x 10-20-50 mm, high and diameter. Sponge tubiform, with the
basal part slightly enlarged and a central canal from 50 to 80 mm diameter in the
different specimens and fragments studied. Surface strongly tuberculate and irregular
(somewhat resembling Xestospongia) and covered by a membrane. The interior of the
sponges shows large open spaces and canals. Pores irregularly scattered in surface
depressions; oscules scattered over the inner wall of the tube.
Consistency: hard and fragile, crumbly.
Colour: Orange-brownish in formalin, brownish to yellow-white in alcohol.
Skeleton: Ectosomal skeleton a perpendicular compact palisade of tylotornotes
and free microscleres.
Choanosomal skeleton: an tight meshed isotropic reticulation of strongly
spined styles, abundant microscleres, especially sigmas.
Spicules (Table 8): Megascleres: strongly spined styles (figs 43, [44 holotype])
slightly bent and with strong spines, scarcely distributed but more abundant at the
base, 163-208 x 3-10 um.
TABLE 8
Spicule micrometries of M. (Myxilla) mexicensis Dickinson, 1945. M. mexicensis Dickinson,
1945 AHF 9 holotype; from literature and remeasured. Specimens from Galapagos of M. (M.)
mexicensis.
Tylotornotes
M. (Myxilla) Spined styles Isochelas 1 Sigmas |
mexicensis Isochelas 2 Sigmas 2
Dickinson, 1945
HOLOTYPE AHF.9 not indicated not indicated not observed 32
San Francisco Island 10-12 14
Remeasured 176-208-226 144-166-176 23-28-31 20-28-35
x 6-8-10 x4 12-18-20 12-14-15
GA.IL.3 163-178-195 144-159-173 22-27-29 22-28-32
Kicker Rock. Stn. 17 x 3-6-10 x3 10-13-16 11-15-19
USNM 37917
GA.II.4 173-184-195 144-160-173 22-25-29 22-26-32
Kicker Rock x 6-7-10 x 3-4-5 11-13-18 10-11-13
USNM 37919
Ga.IV.33 182-199-214 154-165-179 22-29-32 19-25-29
18B794E x 10-11-13 x 3-4-6 10-13-19 11-14-16
Santiago Island
Tylotornotes (figs.45, [46 holotype]) straight, regular diameter and with barely
swollen "cut-off" spined apices, about four to five spines at each end, 141-273 x 3-5
um.
Microscleres: Anchorate spatuliferous isochelas 1 (figs.47, [49 holotype]) 22-
29 um, with a shaft only slightly curved, with three free alas of the same length, the
innermost longer than the others and attached to the shaft. Anchorate spatuliferous
S-E PACIFIC SPECIES OF POECILOSCLERIDA 31
Fics 39-46
Myxilla (Myxilla) mexicensis Dickinson, 1945. 39, specimen from Galapagos Islands. 40,
enlarged view of the surface. 41 [HOLOTYPE, AHF 9] from Gulf of California. 42, enlarged view
of the surface. 43, slightly bent styles, strongly spined, spines scarcely distributed. 44, style
from holotype. 45, tylotornotes straight, regular diameter and with barely swollen "cut-off"
spined apices about four to five spines at each end. 46, tylotornotes from holotype. Scales: figs
43, 44, 46 = 5 um.
isochelas 2 (figs 48, [50 holotype]) 10-19 um, shaft slightly bent, three teeth are
visible in frontal view, the innermost longer than the others and attached to the shaft.
Since these chelas are morphologically identical, it is not entirely certain that the size
categories are genuine.
32 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Fics 47-53
Myxilla (Myxilla) mexicensis Dickinson, 1945. 47 [49 holotype] anchorate spatuliferous iso-
chela 1 with a shaft only slightly curved, with three free alas of the same lenght, the innermost
longer than the others and attached to the shaft. 48 [50 holotype] anchorate spatuliferous
isochela 2, shaft slightly bent, three teeth visible in frontal view, the innermost longer than the
others and attached to the shaft. 51, sigmas 1 and 2. 52, sigma I. 53, sigma 2 from holotype.
Scales: fig. 47 = 5 um; figs 48-53 = 2 um.
Sigmas (fig. 51): Sigma 1, 19-35 um (fig. 52) and sigmas 2: 11-19 um (fig. 53
[holotype]) with one of the apices strongly bent and the other largely open, both
abruptly pointed.
Ecology: Presumably fixed to hard bottom, no visible attaching points, 18-60 m.
Distribution: Galapagos, Southern California.
Remarks: The specimens were compared with a fragment of DICKINSON's type
and found to be essentially similar, as is apparent from figs 39-42. DICKINSON (1945)
reported only a single category of isochelas of 10-12 um, but we found the same size
range as in our specimens (cf. figs 47-50). Although the growth form of Dickinson's
specimen was not tube-shaped, it was a much smaller specimen. One of our
specimens also was smaller and it showed only a widened oscule in stead forming a
definite tube. We assume the growth form to be a tube only in mature specimens.
In spicule form, size and skeletal arrangement this species is closely related to
the type species of Myxilla, viz. M. rosacea (Lieberkühn, 1859), and to other
Northern Hemisphere species as M. fimbriata (Bowerbank, 1866) and M. incrustans
(Johnston, 1842).
S-E PACIFIC SPECIES OF POECILOSCLERIDA 33
Myxilla (Myxilla) mollis Ridley & Dendy, 1886 (Figs. 54-65)
Myxilla mollis RIDLEY & DENDY, 1886: 471; 1887: 133, pl. xxvii, figs. 4, 4a-b; Burton, 1932:
309, pl. lv, figs1-4; 1934: 28; 1940: 107.
Myxilla spongiosa RIDLEY & DENDY, 1886: 471, 1887: 134, pl. xxvii, figs. 3, 3a-f.
MATERIAL STUDIED: BMNH 1887:5:2:112, holotype of Myxilla mollis Ridley & Dendy,
1886, Challenger Exped. Stn.311, off sw coast of Patagonia , 52°45’S 73°46’W, 11.01.1876 ,
448 m; BMNH 1887:5:2:131, Stn. 308, 05.01.1876, 50°08°S74°41’W, 320 m; BMNH 1887:5:2:93,
holotype of Myxilla spongiosa Ridley & Dendy, 1886, Challenger Stn. 320, off mouth of the
Rio de la Plata, 37°17°S 53°52’W, 14.02.1876, 1097 m.
MHNG 18962: HE 8, HE 16, Boca Seno Baker, 47°49°S 74°38’W, 01.10.1972, 300 m;
MHNG 18963: HE 48, Isla Topar, 50°06°S 74°41’W, 16.01.1972, 150 m.
DESCRIPTION
Several specimens (figs. 54, 55) and fragments with basal holdfast, 41-97 x 30-
80 x 10-23 mm. Sponge massive. Surface irregularly tuberculate or minutely conu-
lose, covered by a thin membrane; where this is damaged, an irregular system of
small subdermal cavities is revealed. Several small oscules 2-4 mm diameter,
scattered on the surface, in one specimen, some of them are at the end of a small tube.
Consistency: Soft and compressive, elastic.
Colour: Yellowish-orange to grey alive; light to dark brown in alcohol.
Skeleton: Ectosomal skeleton: tangentially arranged anisotylotes in a loose
palisade. Abundantly distributed microscleres, producing a continuous crust.
Choanosomal skeleton: a loose reticulation of smooth styles, occasionally
arranged side by side (4-5). No continuous tracts. Microscleres abundantly scattered
all over the choanosome, but especially around the aquiferous canals.
Spicules (Table 9): Slightly spined styles to subtylostyles (figs. 56 [60 holo-
type]) smooth, swollen base with a few spines, slightly curved, 414-526 x 10-13 um.
Anisotylotes (figs. 57 [61 holotype]) straight, apices swollen unequally, one
end being more elongated than the other, with small spines on both extremities 221-
280 x 7-10 um.
Microscleres: Spatuliferous anchorate isochelas 1 (fig. 58), three to five rela-
tively long teeth, only slightly curved, 34-49 um. Spatuliferous anchorate isochelas 2
of similar shape (figs. 59 [62 holotype]) 16-22 um .
Sigmas 1, 34-79 and 2 20-32 um (figs. 63-65) common, C - or S - shape.
Ecology: Stones and calcareous substrate, 150-1097 m.
Distribution: West and east coast of South America: 47°S-52°S, 37°S;
Falkland Islands; South Georgia.
Remarks: Our material was compared to the type specimen of Ridley & Dendy,
and found to be quite similar, with the characteristic almost smooth styles, except at the
slightly spined base and two categories of chelas and sigmas. The smooth styles are
shared with M. (Burtonanchora) asymmetrica n.sp. (cf. below), but that species has and
asymmetric chelas, and with M. discoveryi but that species has unguiferate chelas.
Records of this species by BURTON (1932, 1934, 1940) presumably are correct,
extending the distribution to the east coast of South America, and to the Falkland
Islands and South Georgia.
34 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
TABLE 9
Spicule micrometries of M. (Myxilla) mollis Ridley & Dendy, 1886. M. mollis Ridley & Dendy
BMNH 1887:5:2.112 holotype; Myxilla spongiosa Ridley & Dendy, 1886 BMNH 1887:5:2:93
holotype; from literature and remeasured. Specimens from Chile of M. (M.) mollis.
M. (Myxilla) mollis Styles Anysotylotes Isochelas 1 Sıgmas |
R. & D., 1886 Isochelas 2 Sigmas 2
M. mollis R. & D. 1886 420 x 10 220 x 6 40 63
BMNH1887:5:2:112 not observed
sw. Patagonia
Remeasured 421-457-486 227-259-283 32-34-40 40-45-49
x8 x 4-6-8 24-26-28 16-22-28
M. spongiosa R. & D. 1886
BMNH1887:5:2:93 700 x 20 400 x 10 50 45-63
Off Rio de la Plata not observed not
observed
Remeasured 539-586-617 250-280-300 47-49-55 45-65-79
x 15-17-20 x 8-9-10 20-21-23 20-26-31
He 8 428-468-526 222-247-257 36-43-46 46-54-79
Boca Seno Baker x 10-11-13 x 8-10 17-19-22 20-23-24
He 16 414-477-513 221-236-249 34-43-49 40-51-58
Boca Seno Baker x 10-13 eH 16-19-22 21-25-32
He 48 428-471-503 250-267-280 41-48 34-49-58
Isla Topar x 10-12-13 x 16-18-21 21-24-26
By comparing microscopical slides of the types of M. mollis and M. spongiosa
Ridley & Dendy, 1886 from the area off Argentina (37°S), we could ascertain that
this is likely to be a junior synomym of M. mollis (the latter has page priority). Like
M. mollis occasional spines occur on the styles of the type specimen. The only diffe-
rence is the somewhat larger megasclere sizes of M. spongiosa. Hentschel's (1914)
record of this species from Antarctica is probably incorrect as the styles are reported
as spined. Hentschel's material has been elevated to a separate species M. hentscheli
(in subgenus Ectyomyxilla) by BURTON (1929).
Myxilla (Myxilla) caliciformis Sara, 1978
Myxilla caliciformis SARA, 1978: 46, figs. 28, 29.
MATERIAL STUDIED: EAI-76, holotype of Myxilla caliciformis Sara, 1978, Puerto Roca,
Isla Dos Estados, 11.01.74, 2-3 m.
Not represented in our material so only a diagnosis will be given:
Cup-shaped sponge.Anisotropic skeleton superimposed on the isotropic
meshes as usuals for the genus. Acanthostyles small, strongly spined, 120 x 6 um.
Tornotes with heavily spined apices, 120 by 3 um. Spatuliferous anchorate chelas in
two size categories: 18 and 12 um. Sigmas 30 um.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 35
Fics 54-65
Myxilla (Myxilla) mollis Ridley & Dendy, 1886. 54, massive specimen from Isla Topar. 55,
enlarged view of the irregular tuberculate surface. 56 [60. holotype, BMNH 1887:5:2:1 12]
slightly spined styles to subtylostyles. 57 [61, holotype] spined swollen apex from anisotylotes.
58, spatuliferous anchorate 1. 59, spatuliferous anchorate isochela 2 [62 holotype]. 63. sigmas l
and 2. 64 [65 holotype], sigma 2. Scales: figs. 56, 63 = 10 um; figs. 57, 59, 62, = 2 um; figs. 58.
60, 61, 64, 65 = 5 um.
36 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Distribution: Tierra del Fuego.
Remarks: Through its cup-shaped growth form this species stands out among
the Myxilla species of the study area. Judged from the spicules and skeletal structure
its closest relative is M. (M.) mexicensis.
Subgenus Burtonanchora de Laubenfels, 1936
Diagnosis: Myxilla with smooth instead of spined styles.
Myxilla (Burtonanchora) asymmetrica n. sp. (Figs. 66-73)
MATERIAL STUDIED: USNM 39335, HOLOTYPE: MHNG 18831, ZMA 10980: Ga.44 SEPBOP
"Anton Bruun" Exped., Cruise 18B 754, Peru, 07°49'S 80°38'W, 06.09.1966, 605-735 m.
DESCRIPTION
Numerous specimens (figs. 66 [67 Holotype]) of 20-40 x 10-25 mm, and
numerous fragments of 10-20 mm, of an encrusting to conical to massive sponge.
Although the substrate is no longer preserved with the sponge, impression marks on
the undersides of the specimens indicate that this species probably encrusts gastropod
molluscs. Surface rugose and irregular, several small oscules, 1.5-2 mm diameter are
visible.
Consistency: Soft and elastic, very fragile.
Colour: Greyish to brownish in alcohol.
Skeleton: Ectosomal skeleton: tangentially arranged compact palisade of
tylotes and microscleres abundantly distributed.
Choanosomal skeleton: a loose reticulation of smooth styles, irregular aqui-
ferous spaces rounded by isochelas.
Spicules (Table 10): Megascleres: Smooth styles (figs. 68, 69) irregularly bent
and of uniform diameter, 478-571 x 16 um.
Smooth isotylotornotes (fig. 70): 219-283 x 5 um.
Microscleres: Asymmetrical spatuliferous anchorate isochelas (figs 71, 72) that
at first glance appear arcuate, but the curved shaft is sheathed in continuous fimbriae
TABLE 10
Spicule micrometries of M.(Burtonanchora) asymmetrica n. sp. USNM holotype from Peru.
M. (Burtonanchora) Styles Isotylotornotes Isochelas Sigmas
asymmetrica n. sp.
Holotype USNM 39335
Ga.44 478-515-551 219-245-283 20-26-32 49-65
18B754 x 16 XÒ)
07°49’S 80°38’W
S-E PACIFIC SPECIES OF POECILOSCLERIDA 37
FIGs 66-73
Myxilla (Burtonanchora) asymmetrica n.sp. 66 [holotype, USNM 39335] massive specimen from
Galapagos. 67, enlarged view of the surface. 68, 69, smooth styles irregularly bent. 70, smooth
isotylotornote. 71, 72, asymmetrical spatuliferous anchorate isochela. 73, sigma widely
open. Scales: fig. 68 = 50 um; fig. 69 = 10 um; figs. 70, 72, 73 = Sum; fig. 71 = 2 um.
next to the three alae. The asymmetrical aspect is found in the length of the lateral
alas of which one is always considerably shorter than the central one. This modi-
fication occurs always in opposite position. Many of the alae have frayed or
denticulated rims. Size 20-32 um.
Sigmas (fig. 73): widely curved and abruptly pointed, abundant, 49-65 um.
Ecology: Probably encrusting gastropods, 605-735 m depth.
Distribution: Peru, 07°49’S 80°38’ W.
38 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Remarks: The asymmetrical chelas are unique for the genus. There are no
matching descriptions in the literature. In other families, also species with asymme-
trical chelas have been described: Esperiopsis glomeris Topsent, 1904 and E. pul-
chella Boury-Esnault & Van Beveren, 1982 (family Mycalidae) and Phorbas dayi
(Levi, 1963 as Anchinoe) (family Anchinoidae). These represent convergent deve-
lopments.
Subgenus Eetyomyxilla Lundbeck, 1909
Diagnosis: Myxilla with echinating strongly spined styles.
Myxilla (Ectyomyxilla) chilense Thiele, 1905 (figs 74-81)
Myxilla chilensis THIELE, 1905: 443, figs. 22, 62 a-e; BURTON 1932: 311, pl. liv, fig. 10; 1934
[in part]: 28; 1940: 108; BOURY-ESNAULT & VAN BEVEREN 1982, as Crellomyxilla [in
part?]: 84, pl. xiv, fig. 53, fig. 24 a-e.
MATERIAL STUDIED: ZMB 3298, holotype of Myxilla chilensis Thiele, 1905, Calbuco,
41°46°S 73°08’W. MHNG 18965-18967: VALD. 44.1, Quintupeu, 42°10°S 72°24’W, 23.07.
1971, 12 m; VALD. 53.7, Islote Caleta Lobato, 45°5’S 74°47’W, 17.07.1972, 15 m; Co 93-51,
Pelluco, 41°30°S 72°54’W, 20.12.1990, 15 m.
DESCRIPTION
One specimen (figs. 74, 75), 50 x 40 x30 mm, and two fragments of 31-48 x
30-39 x 29-39 mm. Sponge largely massive. Surface uneven, hispid, minutely corru-
gated or conulose and covered by a thick, transparent surface membrane. Oscules
numerous, scattered, 1.5-3 mm diameter.
Consistency: compressible.
Colour: bright yellow, yellowish-orange alıve, pink to whitish in alcohol.
Skeleton: The ectosomal skeleton is a variable thick, dense palisade of tornotes
and acanthostyles 2 positioned at oblique angles, forming part of the detachable sur-
face membrane. The choanosomal skeleton is a dense and irregular to subisodictyal
network of spined styles 1 and abundant microscleres. Numerous aquiferous canals
are distributed all over the choanosome.
Spicules: (Table 11) Megascleres: Strongly spined acanthostyles 1 (fig. 76) al-
most straight, abundantly distributed spines, which are longer at the basal end, sharp
apex, 176-229 x 8-13 um. Spined acanthostyles 2 (fig. 77) slightly curved, numerous
small spines, sharp apex, 83-144 x 3-6 um.
Tornotes (fig. 78) smooth, straight, sharp, abrupt apices, 157-211 x 4-10 um.
Microscleres: Spatuliferous anchorate isochelas 1 (fig. 79) curved shaft, com-
mon 18-45 um. Spatuliferous anchorate isochelas 2 (fig. 80) curved shaft, common,
10-16 um.
Sigmas, C- or S - shaped (fig. 81) common, tapering abruptly 32-64 um.
Ecology: Pebbles, calcareous and bryozoa fragments, 12-15 m.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 39
TABLE 11
Spicule micrometries of M. (Ectyomyxilla) chilense Thiele, 1905. M. chilensis Thiele, 1905 zmB
3298, HOLOTYPE; from literature and remeasured. Specimens from Chile of M. (E.) chilensis.
M. (Ectyomyxilla) Acanthostyles 1 Tornotes Isochelas 1 Sigmas
chilense Acanthostyles 2 Isochelas 2
Thiele, 1905 :
Holotype ZMB 3298 170-200 170-190 20-35 45-47
Calbuco x 10-12 x 6-7 12-15
not observed
Remeasured 182-195-208 163-181-202 19-26-29 45-47-51
x 6-8-10 x 6 11-14-16
102-122-134
x 6
Vald. 44-1 205-216-229 192-201-211 18-20-29 35-54-64
Quintupeu x 8 x 4-6 10-14-16
83-119-144
x 3-6
Vald. 53-7 176-191-202 157-181-298 38-40-45 32-43-48
I. Caleta Lobato x 8-10-11 x 5-6 10-14-16
112-125-138
x 3-6
Co.93-51 176-198-218 166-184-195 19-22-29 38-40-45
Pelluco x 10-11-13 x 4-7-10 10-14-16
128-132-144
x 6
Distribution: Chilean coast : 41°S-45°S; Falkland Islands; Kerguelen ?
Remarks: BOURY-ESNAULT & VAN BEVEREN (1982) recorded "Crellomyxilla"
chilensis and "C."kerguelensis sympatrically from the Kerguelen Islands. Apparently,
the two species may be distinguished on the different size of the sigmas, 45-58 and
12-32 um. Other authors (e.g. BURTON 1934) considered the two as conspecific.
Crellomyxilla Dendy (1924) with type species C. intermedia Dendy, 1924 from
northern New Zealand, is a junior synonym of Ectyomyxilla. Dendy interpreted the
acanthostyles 2, which are mixed in among the ectosomal tornotes, as evidence for
crellid affinity. This character, however, is also found in M. kerguelensis, the type of
Ectyomyxilla, so synonymy is clear. If M. chilensis, M. kerguelensis, and M.
intermedia are distinct species, they may be separated as a monophyletic subgenus
from other other Myxilla species possessing acanthostyles 2.
Myxilla (Ectyomyxilla) dracula n. sp. (Figs. 82-90)
HOLOTYPE: USNM 39351, MHNG 18968, ZMA 10981: Ga.IIT.36, SEPBOP "Anton Bruun"
Cruise 16 627 A, Peru, 05°01’S 81°25'W, 06.1966, 200 m.
40 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Fics 74-81
Myxilla (Ectyomyxilla) chilense Thiele, 1905. 74, massive specimen from Quintupeu. 75,
enlarged view of the surface. 76, acanthostyle 1, strongly spined, 77, acanthostyle 2, slightly
curved. 78, smooth oxotornote. 79, 80, spatuliferous anchorate isochela 1 and 2. 81, sigma,
abruptely pointed, C-shaped. Scales: figs. 76, 77 = 20um; figs. 78, 79, 81 = Sum; fig. 80 = 2pm.
DESCRIPTION
Small, thinly encrusting layer on a mass of hydroid chitinous stolons (figs. 82,
83). Surface irregular and covered by a spiculous membrane. No oscules are visible.
Consistency: Very fragile, but elastic.
Colour: Grayish in alcohol.
Skeleton: Ectosomal skeleton: a palisade of densely packed bundles of tylo-
tornotes and abundant free tylotornotes. Microscleres are abundant too.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 4]
Choanosomal skeleton: A loose and confused reticulation of strongly to
moderately spined acanthostyles 1 and 2 and extremely abundant microscleres. The
sponge skeleton surrounds the mass of hydroid stolons.
Spicules (Table 12): Megascleres: Acanthostyles 1 (fig. 84) almost straight,
with only a few strong spines, 308-397 x 8 um. Acanthostyles 2 (fig. 85) slightly bent,
spines abundant, 140-160 x 2 um.
TABLE 12
Spicule micrometries of M. (Ectyomxilla) dracula n. sp. MNHG 18968, holotype from Galapagos.
M. (Ectyomyxilla) Acanthostyles 1 Anisotylotornotes Isochelas 1 Sigmas 1
dracula n. sp. Acanthostyles 2 Isochelas 2 Sigmas 2
HOLOTYPE USNM 39351 2
Ga.III.36 308-355-397 170-182-194 40-45-49 32-42-51
16627A x 8 x4 12-13-16 18-21-24
05°01°S 81°25°W 140-160
x2
Anisotylotornotes (fig. 86) with a few small spines at the apices, 170-194 x 4
um.
Microscleres: Spatuliferous anchorate isochelas 1 (fig. 87) straight shaft, 40-49
um. Peculiarly small anchorate isochelas 2 (fig. 88) strongly curved and with long
narrow alas sharply bent inwards and almost meeting in the centre, 12-16 pm.
Sigmas 1 and 2 (figs. 89, 90) largely open and abruptely pointed, 32-51 and
19-24 um.
Etymology: the name refers to the long thin teeth of the chela 2, which evoke
associations of dragon's teeth.
Ecology: 200 m depth.
Distribution: Peru, 05°01’S 81°25'W.
Remarks: The characteristic curved chelas 2 make this new species distinct
from any known species of the genus. Apart from these peculiar chelas, the new
species shows similarities with Myxilla massa Ridley & Dendy, 1886 (cf. below),
described from Chile. This species has only a single category of sigmas and the
acanthostyles 1 are fully spined.
Myxilla (Ectyomyxilla) massa Ridley & Dendy, 1886
Myxilla mariana var. massa RIDLEY & DENDY, 1886: 473; 1887: 138, pl. xxvii, figs. 6,
6a-f.
MATERIAL STUDIED: BMNH 1887:5:2:95, HOLOTYPE of M. mariana var. massa Ridley &
Dendy, 1886, Challenger Expedition, Station 311, off the south west coast of Patagonia, 52°45'S
73°46'W, 01.1876, 448 m; BMNH 1887:5:2:108, holotype of M. massa Ridley & Dendy, 1886,
Challenger Expedition, Station 142, off Marion Island, 35°04'S 18°37'W, 12.1873, 274 m.
49 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
FIGs 82-90
Myxilla (Ectyomyxilla) dracula n.sp. 82 [holotype USNM 39351] encrusting specimen from
Galapagos. 83, enlarged view of the surface. 84, acanthostyle 1 almost straight with only few
strong spines. 85, acanthostyle 2 slightly bent abundantly spined. 86, anisotylotornotes with a
few small spines at the apices. 87, spatuliferous anchorate isochela 1. 88, peculiarly incurved
small isochela 2. 89, 90, sigmas | and 2, largely open, abruptely pointed. Scales: fig. 84 =
10um; figs 85-87 = Sum; figs. 88, 90 = 2um.
Not represented in our material, so only a diagnosis is given:
Massive sponge. Loosely reticulated skeleton. Acanthostyles 1 fully spined,
400-420 by 10 um. Acanthostyles 2 fully spined, 140-150 by 6 um. Smooth tornotes
250-280 by 4.5 um. Spatuliferous anchorate isochelas 1 and 2, 30-37 and 18 um.
Sigmas 45-56 um.
Distribution: Chilean Patagonia, 52°S.
Remarks: As discussed above, M. (E.) dracula n.sp. is similar to M.(E.)massa
but has strongly incurved isochelas and sparingly spined styles. M. massa was
S-E PACIFIC SPECIES OF POECILOSCLERIDA 43
synonymized with M. mariana Ridley & Dendy, 1886 from Marion Island (Indian
Ocean) by KOLTUN (1964). We examined a slide made from the type specimen and
found that a separate acanthostyles 2 category is not well differentiated. Also, the
tornotes have rounded smooth oval heads instead of the mucronate ones of M. massa.
We assume that these differences are evidence for specific distinctness.
?Myxilla (Ectyomyxilla) tenuissima (Thiele, 1905)
Hymedesmia tenuissima THIELE, 1905: 454, figs. 70a-f.
Ectyomyxilla tenuissima (Thiele, 1905); LÉVI, 1963: 36, fig. 39; DESQUEYROUX, 1976: 102.
No material examined, so only tentative remarks can be made:
Thinly encrusting sponge. Hymedesmioid skeleton or regular reticulation.
Acanthostyles 1 180-290 by 10-15 um. Acanthostyles 2 100-150 by 7-10 um. Tor-
notes oxea-like, with microspined ends. Anchorate chelas 10-30 um. Sigmas: 45-48
um.
Distribution: Calbuco, Chilean coast, 41-42°S; South Africa, 31°14'S 16°36'E,
12-272 m.
Remarks: On paper this description comes close to M. chilensis, with as the
only tangible difference the microspined condition of the tornotes. Type specimen is
not extant at ZMB, for the time being this species must be considered as incertae sedis.
Subgenus Stelodoryx Topsent, 1904
Diagnosis: Myxilla with unguiferate instead of or in addition to spatulate
anchorate chelae.
Myxilla (Stelodoryx) cribrigera Ridley & Dendy, 1886 (Figs. 91-98)
Myxilla cribrigera RIDLEY & DENDY, 1886: 472, 1887: 135, pl. xxvii, figs. 7, 7a,b.
Stelodoryx discoveryi BURTON, 1932: 316, fig. 28.
?Myxilla verrucosa BURTON, 1932: 312, text fig. 27; DESQUEYROUX, 1972: figs. 81-86; 1976:
101.
MATERIAL STUDIED: BMNH 1887:5:2:138, HOLOTYPE of Myxilla cribrigera Ridley &
Dendy, 1886, Challenger Expedition, Stn. 306A, off the south-west coast of Patagonia, 48°27°S
74°30°W, 02.01.1876, 631 m; BMNH 1928:2:15:426, holotype of Stelodoryx discoveryi Burton,
1932, St. WS 88, Falkland Islands, 54°07’S 58°16’W, 06.04.1927, 96-127 m; MHNG 18969-
18974: He 178, Seno de Otway, 53°00’S 71°30’W, 17.09.1972, 30 m; MNHNC 65, 50°43’S
76°08’ W, 10.03.1978, 248 m; Vald. 66a, Punta Roja, 43°21’S 73°44’W, 24.05.1971, 25 m; Co
93-27, Quintupeu, 42°10’S 72°24’ W, 25.03.1972, 25 m; Co 93-47, Pelluco, 41°30’S 72°54’ W,
10.02.1992, 20 m; GA HI 21, sEPBOP "Anton Bruun" Exped., 18B791C, Galapagos Islands,
00°37’S 90°51’ W, 08.05.1966, 78 m.
DESCRIPTION
Several fragments and one specimen (figs. 91, 92) 21-27 x 17-19 and 32 x 23
mm. Sponge cylindrical, but the basal part is missing. Surface even, microhispid,
44 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
covered by a thin transparent membrane, in a continuous layer firmly attached.
Terminal oscule, 3-7 mm diameter, from which diverge 4-5 aquiferous canals.
Numerous small oscula, 1.5-3 mm diameter, are scattered on the surface.
Consistency: Soft and fragile, elastic.
Colour: Varying form light brown to dark-brown, in alcohol.
Skeleton: Ectosomal skeleton: arranged as palisade of bundles of tylotornotes
and a continuous cover of spatuliferous anchorate isochelas.
Choanosomal skeleton: isodictyal reticulation of polyspicular tracts of styles,
ending by subectosomal bundles of styles passing through the surface (causing a
hispid surface). Longitudinal tracts connected by transverse bundles of styles (3-4) or
by single styles. Microscleres abundant.
Spicules (Table 13): Megascleres: styles (fig. 93) smooth, slightly curved 345-
632 x 8-24 um.
TABLE 13
Spicule micrometries of M. (Stelodoryx) cribrigera Ridley & Dendy, 1886. Myxilla cribrigera
Ridley & Dendy, 1886, BMNH 1887:5:2:138, holotype; Stelodoryx discoveryi Burton, 1932
BMNH 1928:2:15:426, holotype; from literature and remeasured. Specimens from Chile of M.
Myxilla (Stelodoryx)
Styles
(S.) cribrigera.
Isochelas 1
Tylotornotes
cribrigera Isochelas 2
R. & D., 1886
M. cribrigera R. & D. 1886
BMNH 1887:5:2:138 650 x 25 300 x 8 80
off sw coast Patagonia
Remeasured 502-576-607 243-279-316 57-63-79
x 16-23-24 x 8 32-39-43
Stelodoryx discoveryi
Burton, 1932
BMNH 1928:2:15:426 22556 165 x 5 35-45
Falkland Islands not observed
Remeasured 324-351-373 227-238-251 36-47-57
x8 x 8 40-46-49
He 178 389-460-494 202-235-250 48-49-58
Seno de Otway x 12-18-20 x 6-7-8 48-53-62
MNHNC 65 446-499-527 243-265-284 61-63-67
50°43°S76°08° W x 12-22-24 x 6-7-8 48-51-57
Vald. 66a 345-405-470 214-233-307 49-57-64
Punta Roja x 12-15-16 x 6-8 48-60
Co 93-27 454-476-494 182-193-198 51-60-67
Quintupeu x 6 x 6-8 35-42-48
Co 93.47 348-352-356 179-220-251 48-54-57
Pelluco x 8-13-16 x 6-8 37-47-51
Ga.IIT.21 567-608-632 227-254-283 81-86-89
18B791C x 16-22-24 x 8 74-78-86
002375 90°51 W
S-E PACIFIC SPECIES OF POECILOSCLERIDA 45
Fics 91-98
Myxilla (Stelodoryx) cribrigera Ridley & Dendy, 1886. 91, fragment of a cylindrical specimen.
92, enlarged view of the surface. 93, smooth slightly curved styles. 94, tylotornotes smooth
with equally swollen microspined cut-off apices. 95, 96, spatuliferous anchorate isochela 1. 97,
unguiferous anchorate isochela 2. 98, terminal enlarged region of the apex. Scales: figs. 93, 97
= 10um; figs. 94, 95, 98 = Sum; fig. 96 = 2um.
Tylotornotes (fig. 94) smooth, straight, with equal, slightly swollen micro-
spined apices which have a characteristic "cut-off" shape, 179-307 x 6-8 um.
Microscleres: Spatuliferous anchorate isochelas 1 (figs. 95, 96) with slightly
curved shaft and 5 to 9 spatuliferous teeth, parallel to each other, very common, for-
ming a continuous layer on surface, 48-89 um. Unguiferous anchorate isochelas 2
(figs. 97, 98) less numerous, present only in the choanosome, shaft almost straight ,
with 5 teeth, the innermost alas visible only as slightly enlarged region of the shaft ,
35-86 um.
46 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Ecology: no data about substrate, 20-248 m.
Distribution: Galapagos, Chilean coast, 41°S - 53° S ; Falkland Islands.
Remarks: The conspecificity of M. cribrigera and M. discoveryi was es-
tablished in a comparison of the slides made from both holotypes.
The unguiferous chelas are also described for Myxilla verrucosa Burton, 1932,
which was described simultaneously with M. discoveryi both from the Falkland
Islands. Differences are found in the spicule sizes (M. verrucosa: styles 240 x 8-18
um, tornotes 160 x 7 um, chelas 35 um). DESQUEYROUX (1972) reported M. verrucosa
from Golfo Corcovado: styles 250-400 x 8-21 um, tornotes 170-240 x 7-8 um,
isochelas 33-66. There is not indication about the morphology of anchorate chelas.
DESQUEYROUX (1976) reported M. verrucosa from San Pedro, Chile, but gave no des-
cription of her material. It is here left undecided whether M. verrucosa is a separate
species.
Stelodoryx was erected on the possession of unguiferous chelas in combination
with a stalked growth form. This combination is not found in the present species, but
we assume that a stalked habit is an adaptation to special environmental conditions. In
view of the widespread occurrence in many different groups of sponges, it is likely
that this character has no supraspecific significance.
REVIEW OF THE FAMILY MYXILLIDAE
Remarks: The above given diagnosis of the family Myxillidae excludes
traditional Myxillid genera, such as Lissodendoryx and Ectyodoryx, which have
arcuate chelas. These are united with the fistular genera Coelosphaera and Forcepia,
which also have arcuate isochelas, into an emended family Coelosphaeridae. For a
discussion and further arguments on this new assemblage cf. HAJDu et al. (1994). The
major synapomorphy distinguishing the Myxillidae from the Coelosphaeridae is the
anchorate condition of the chelas; other features are the more frequent spination of the
ectosomal tornotes which are also predominantly oxeote or strongylote rather than
tylote.
Genera possessing birotulate chelas (Amphiasterella Dendy, 1924, Hymetro-
chota Topsent, 1904 (with synonym Hymenotrocha Burton, 1930), /otrochota (with
synonyms Hiattrochota, lotrochostyla and lotrochopsamma, all of DE LAUBENFELS,
1950, 1954, cf. VAN Soest, 1987), Plocamiancora Topsent, 1927, and Rotuloplo-
camia Lévi, 1952) have traditionally been spread over various families of the Poeci-
losclerida (Desmacididae, Hymedesmiidae, Coelosphaeridae, Myxillidae and Micro-
cionidae), see for example discussion in VAN SOEST (1987) . However, the idea that
birotulates have been developed only once and thus that all birotuliferous genera are
monophyletic (suggested in HAJDU er al, 1994) has considerable merit. The poly-
dentate condition of the birotulate chelas reminds of the polydentate-unguiferous
chelas of several Myxillid genera. Also in one genus, /otroata, both birotulate and
anchorate chelas are found. However, birotulates in a clearly Antho-like sponge as
Rotuloplocamia octoradiata Lévi, 1952, almost birotulate-like palmate chelas of
S-E PACIFIC SPECIES OF POECILOSCLERIDA 47
Plocamiancora denticulata Topsent, 1928, or clearly anchorate-derived condition of
the /otroata spinosa (Lundbeck, 1905) (see SEM photo in VAN SOEST, 1987, plate I fig.
6), are indications that convergent development of the birotulate condition may be
equally likely. Since no birotuliferous genera are represented in the se Pacific, we
refrain from commenting upon their family assignment (excepting /otroata which is a
clear myxillid); we merely list them for future treatment.
The following genera are thought to belong to the new concept of Myxillidae:
Genus Desmacidon Bowerbank, 1864
Type species: Spongia fruticosa Montagu, 1818 (topotypical specimens exa-
mined).
Diagnosis: In the ectosome smooth oxeote spicules are arranged in bouquets.
Tracts of the same smooth oxeote spicules form an isodictyal reticulation in the choa-
nosome. Microscleres are spatuliferous anchorate isochelas and sigmas very similar in
shape to those of Myxilla.
Remarks: The megascleres are assumed to be tornotes, which have replaced
the usual styles in the choanosomal tracts. It is quite possible that D. fruticosum is a
Myxilla which has lost its styles, and synonymy of these genera may be warranted. In
the past, the genus has been in frequent use for a wide variety of sponges, most of
which are not closely related to D. fruticosum.
Desmapsamma Burton, 1934
Type species: Fibulia anchorata Carter, 1882 (topotypical specimens exa-
mined).
Diagnosis: Ectosomal coat of sand grains, partly replacing the ectosomal bou-
quets of oxeas. Choanosomal skeleton a paucispicular isotropic reticulation of oxeas
similar to those of the ectosome. Microscleres anchorate chelas and sigmas.
Remarks: Use of Desmapsamma as a subgeneric unit has merit in order to
manage the many arenaceous species known to live in the southern oceans (e. g.
WIEDENMAYER, 1989). This sponge is similar to D. fruticosum in most respects, but
oxeas are thinner. Chelas and sigmas smaller. Synonymy of Desmapsamma and
Desmacidon seems inescapable.
Ectyonancora Lévi, 1963
Type species: E. flabellata Lévi, 1963. (HOLOTYPE LBIMDCL 548, 549, exami-
ned)
Diagnosis: Smooth hastate/mucronate tornotes and an isotropic reticulation of
thick acanthostrongyles echinated by thick acanthostyles. Chelas are robust, spatuli-
ferous. No sigmas.
48 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Remarks: The generic distinctness rests on the acanthostrongyles being dif-
ferent from the often blunt-ending spined styles of Myxilla. The use of this genus
name so far is limited to South African waters (including Namibia and Kerguelen).
Hymenancora Lundbeck, 1910
Type species: Leptosia umbellifera Topsent, 1904 (not examined).
Diagnosis: Smooth strongyles as ectosomal spicules and hymedesmioid
architecture, i.e. with acanthostyles erect on the substrate. The anchorate isochelas are
unguiferous with 7-9 teeth.
Remarks: The genus is customarily assigned to Hymedesmiidae because of its
thinly encrusting habit and skeletal architecture. Four species have been described
under this genus name, all from cold deep waters.
Iotroata De Laubenfels, 1936 (junior synonym: lotaota De Laubenfels, 1936).
Type species: /otrochota acanthostylifer Stephens, 1916 (not examined).
Diagnosis: Smooth ectosomal tylotes. Isotropic reticulation of smooth or
acanthose styles, occasionally oxeas. Chelas include both anchorates and birotulates.
No sigmas.
Remarks: The birotulates are clearly derived from anchorates, as neighbouring
teeth show evidence of being the product of subdivision of a former ala. Several
North Atlantic species originally described under /otrochota belong to this genus (cf.
VAN SOEST 1987); at least one, /. spinosa (Lundbeck, 1905) (examined for this study),
has only birotulates, no anchorates. It is confined to cold and deep water. Possibly, the
genus needs to be united with Hymetrochota Topsent, 1904 (cf. below).
Melonanchora Carter, 1874
Type species: M. elliptica Carter, 1874 (topotypical specimen examined).
Diagnosis: Fistular growth form with a paper-like thin ectosome. Ectosomal
smooth tylotes, choanosomal spicules are smooth styles. Microscleres melonchelas
and anchorate chelas.
Remarks: The fistular growth form and ectosomal smooth tylotes make this
genus similar to many Coelosphaeridae. However, the melonchelas are clearly deri-
ved from anchorate chelas, as the youngest growth stages of these are indeed perfect
anchorates. Confined to cold and deep water.
Styloptilon Cabioch, 1968
Type species: Styloptilon ancoratum Cabioch, 1968 (examined).
This species is similar in spiculation to Myxilla but has a plumose rather than
an isotropic skeleton. In view of the variability of the skeletal structure a separate
S-E PACIFIC SPECIES OF POECILOSCLERIDA 49
genus Styloptilon for species with plumose skeletons is not accepted as valid, but if
there are more species like its type, then this can serve as a subgeneric unit.
Onychomyxilla Topsent, 1927
Type species: Dendoryx pectinata Topsent, 1892 (not examined).
Diagnosis: The ectosomal spicules are smooth tylotes; the choanosomal skele-
ton is an isotropic reticulation of acanthostyles echinated by smaller acanthostyles.
The chelas are unguiferous verging towards birotulates, and there are no sigmas.
Remarks: The genus sofar is monotypical and very close to Myxilla (Ectyo-
myxilla) differing in the unguiferous condition of the chelas and the lack of sigmas. It
is also close to Myxilla (Stelodoryx) differing in the possession of echinating
acanthostyles. Finally, the peculiar condition of the chelas indicates that it may be
close to /otroata but differs from it in the possession of echinating acanthostyles.
Plocamissa Burton, 1935
Type species: Plocamia igzo De Laubenfels, 1930 (not examined).
Diagnosis: The ectosomal spicules are smooth tornotes; the choanosomal ske-
leton is "plocamiid", i.e. an isotropic basal reticulation of single acanthostrongyles
echinated by long microspined styles. The chelas are spatuliferous anchorates.
Remarks: The family assignment rests on the presumed anchorate condition of
the chelas, but it is otherwise similar to microcionids (genus Antho). The genus is
monotypical, known only from the west coast of North America.
Pseudomyxilla Koltun, 1955 (with junior synonym Stelotrochota Bakus, 1966).
Type species: P. vitiazi Koltun, 1955 (not examined).
Diagnosis: Ectosomal tornotes and the choanosomal styles similar to those of
the type of Myxilla; the chelas are unguiferous and there are no sigmas.
Remarks: The genus is very close to Myxilla (Stelodoryx) and should probably
be considered synonymous. Species are described from Antarctic or Subantarctic
waters, and from the west coast of North America.
CONCLUSIONS
Next to several well-defined genera (Myxilla, Melonanchora, lotroata) there is
a large number of smaller genera with dubious synapomorphies which need to be
closely compared:
-Desmacidon + Desmapsamma: like Myxilla but lacking proper choanosomal
megascleres different from the ectosomal ones.
50 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
-Hymenancora + Onychomyxilla + Pseudomyxilla : like Myxilla (Stelodoryx),
but with echinating acanthostyles.
- Ectyonancora + Plocamissa: like Antho (Microcionidae) but with spatulate
anchorate chelas
The following genera with anchorate chelas are excluded from the Myxillidae,
and assigned to a separate family Crambeidae:
Crambe Vosmaer, 1887: no ectosomal tornotes.
Discorhabdella Dendy, 1924: ectosomal subtylostyles
Leptosiopsis Topsent, 1927: polytylote ectosomal styles
Monanchora Carter, 1883: ectosomal subtylostyles
Psammochela Dendy, 1896: no ectosomal tornotes.
KEY TO THE SUBGENERA AND SPECIES OF Myxilla OCCURING ON THE SE. PACIFIC COAST
1 Singsle/catesory; of smooth'styles... 2... 2... iS EE 2
- Two categories of styles: Larger spined or smooth styles. Smaller fully
spinedistvlessret anzio i cre DAL ae Subgenus Ectyomyxilla . . . . 5
2 Styles slightly to strongly spined stytes, 2 sizes of spatuliferous
anchoraterisochelasas. rar state cose ican ho Subgenus Myxilla . ...3
- Smoothstylesius asthenia ST el Dae Raa PNR 7
3 Spines on styles concentrated at the apices. Shaft largely smooth.......... 4
— Stronelyfspmedistylestall'overestee rene e M. (Myxilla) caliciformis
4 Sigma | small, from 20 to 35, sigma 2 from 10 to 20. . M. (Myxilla) mexicensis
- Sigmas | big, from 30 to 80, sigmas 2 from 20 to 30 ..... M. (Myxilla) mollis
5 With mucronate tornotes, two categories of spatuliferous anchorate
isochelas and one size of sigmas ........... M. (Ectyomyxilla) massa . . . . 6
- With tylotornotes, one category of normal and one of peculiarly in-
curved spatuliferous anchorate isochelas and two sizes of sigmas
CA SERPS EN ee ee den le M. (Ectyomyxilla) dracula
6 SigmasylongerthanS OMe cee zu: M. (Ectyomyxilla) chilensis
- Sismas:shorter than SO NME EC oe eee ce M. (Ectyomyxilla) tenuissima :
7 A single category of asymmetrical spatuliferous anchorate chelas and a
SIGS CACO] Or SITES 6 5 BS Ml CE clo sas M. (Burtonanchora) asymmetrica
- A single category of spatuliferous anchorate chelas, and one category of
unguiferate anchorate chelas, without sigmas . . . . . M. (Stelodoryx) cribrigera
Tedaniidae Ridley & Dendy, 1886
Diagnosis: Myxillina without microscleres other than onychaetes. Skeletal
architecture variously isotropic, anisotropic, or plumose.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 5]
Tedania Gray, 1867
Synonyms: Trachytedania Ridley, 1884, Tedaniopsis Dendy, 1924, Paratedania Bur-
ton, 1929, Oxytedania Sara, 1978).
Type species Reniera digitata Schmidt, 1862 (junior synonym of Halichondria
anhelans Lieberkühn, 1859).
Diagnosis: Tedaniidae possessing morphologically distinct ectosomal and
choanosomal megascleres. The type species has microspined tylotes as ectosomal
spicules and smooth styles as choanosomal spicules.
Remarks: Tedania anhelans (Lieberkühn, 1859), originally described from the
Mediterranean-Atlantic area, is a soft irregularly lobate sponge with rather variable
colour (brownish, greenish, bluish) and smooth surface. Ectosomal spicules are tylotes
of 140-300 by 2-7 um with well developed microspined heads, arranged in bouquets
and scattered singly in the peripheral region. Choanosomal spicules are smooth styles
170-280 by 5-11 um arranged in a loose reticulation with square meshes. Microscleres
are thin rugose asymmetrical onychaetes in the size range between 40 and 200 um.
Closely similar species have been reported from almost all subtropical and
tropical seas of the world. It is proposed here, in continuation of proposals made the
previous genera, to acknowledge the existence of a complex of probably closely
related species by erecting a nominal subgenus Tedania for them. Species assigned to
Tedania s.s.: T. anhelans (Lieberkühn, 1859) (Mediterranean-Atlantic), T. ignis
(Duchassaing & Michelotti, 1864) (Caribbean-Brazil), Tedania tepitootehenuaensis
Desqueyroux-Fatindez, 1990 (Isla de Pascua), and about 40 further species described
from all tropical and temperate seas of the world.
The genus Trachytedania Ridley, 1881 (with type species T. spinata Ridley,
1881) was differentiated from Tedania proper by the alleged spination of the choano-
somal styles. Re-examination of Ridley's type slides revealed that only very few
styles with spines are present; few authors have accepted the occasional spines on the
styles as sufficient ground for upholding a separate genus. There is, however, a
further difference with species of the subgenus Tedania: the ectosomal spicules differ
rather strongly from T. anhelans in being mucronate-oxeote, without spination on the
apices. The skeletal structure is very loose and no distinct meshes are recognized. It is
proposed here to retain Trachytedania as a subgenus of Tedania.
Species assigned to this subgenus are: Tedania spinata (Ridley, 1881) (with
Junior synonym Tedania murdochi Topsent, 1904) (Magellan), Tedania patagonica
(Ridley & Dendy, 1886) (with junior synonym Tedania inflata Sara, 1978) (Magellan),
Tedania gurjanovae Koltun, 1958 (Arctic), Tedania microrhaphidiophora Burton, 1935
(North Pacific and Arctic), Tedania mucosa Thiele, 1905 (with junior synonyms 7.
fuegiensis Thiele, 1905, T. pectinicola Thiele, 1905, and T. excavata Thiele, 1905)
(Magellan, SE Pacific), and possibly T. toxicalis De Laubenfels, 1930 (cf. below).
The genus Tedaniopsis Dendy, 1924 (with type species T. turbinata Dendy,
1924) was erected because of the replacement of choanosomal styles by strongy-
lotylotes. T. turbinata is an elaborate stalked species from New Zealand, related to T.
massa in spicule sizes, form and skeletal structure. The structural spicules are thick
52 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
large strongylotylotes, differing from the often strongylote spicules of T. massa, but
they are similar in size. Ectosomal spicules are thinner strongylotylotes with mucro-
nate apices, often on one side provided with two rather than a single spine. The
onychaetes 1 are long and thick, and have a clearly developed tyle. No further species
with exactly similar spiculation have been described. However, the large sizes of the
spicules and the irregular skeletal architecture unite the present species with several
others, and accordingly it is proposed to recognize a distinct subgenus for them.
Species assigned are: Tedania massa Ridley & Dendy, 1886 (with junior syno-
nyms T. actiniformis Ridley & Dendy, 1886 and Tedania tantula (Kirkpartick, 1907
as Oceanapia)) (Antarctic, Magellan) Tedania charcoti Topsent, 1917 (Antarctic,
Magellan, se Pacific) Tedania infundibuliformis Ridley & Dendy, 1886 (se Pacific)
Tedania tenuicapitata Ridley, 1881 (Magellan, SE Pacific) Tedania turbinata
(Dendy, 1924) (New Zealand) Tedania oxeata Topsent, 1917 (Antarctic) Tedania
vanhoeffeni Hentschel, 1914 (Antarctic) Tedania phacellina Topsent, 1928 (deep
water North Atlantic).
The genus Paratedania Burton, 1929 (with type species Oceanapia tantula
Kirkpatrick, 1907, which is probably a junior synonym of Tedania massa Ridley &
Dendy, 1886) was erected because of a densely packed layer of tangential mega-
scleres, as opposed to the bouquet-type arrangement of all other Tedaniidae. Ocea-
napia tantula (persistently named tarantula by BURTON, 1929, 1932) is a lobate to
erect sponge with smooth anisostrongyles as structural megascleres and tylotornotes
with an occasional mucron at one of the apices. The tangential arrangement at the
surface, however, was probably entirely induced by the periostracum of mollusc
shells on which the specimens were found to be attached, and thus must be considered
artifactual. BURTON (1932) withdrew this genus himself, and suggested that the type
was a junior synonym of Tedania massa. Examination of type material of both re-
vealed some differences in the shape of tornote endings (hastate in T. massa), but in
view of the variability of this character, conspecificity of both is certainly possible.
Both are also similar to Tedania turbinata, and accordingly Paratedania is considered
a junior synonym of the subgenus Tedaniopsis.
The genus Oxytedania Sara, 1978 (with type species O. bifaria Sara, 1978)
was erected for a species with the spicule combination of typical Tedania in addition
to a reticulate skeleton of oxeas. The type specimen available to us for study did not
contain any of the Tedaniid spicules described by Sara. The skeletal architecture was
typically that of the Haplosclerid family Niphatidae with multispicular tracts of oxeas
in a tight reticulation. It is assumed that the genus is based on a contamination of
Tedania spinata spicules in a niphatid specimen. Indications for this conclusion is not
only our failure to find any Tedania - like spicules, but also the fact that Sara not only
pictures the oxeas as structural megascleres, but also styles of quite dissimilar size
and form as a second category of structural megascleres. Two such different structural
megasclere categories together in a single sponge are unlikely to occur in a tedaniid.
Accordingly this genus is pronounced unrecognizable c.q. a junior synonym of
Tedania.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 53
Subgenus Tedania Gray, 1867:
Diagnosis: Tedania possessing smooth, relatively small styles, occasionally
strongylote styles, as structural megascleres and microspined tylotes as ectosomal
megascleres.
Tedania (Tedania) galapagensis n.sp. (Figs. 99-104)
MATERIAL STUDIED: HOLOTYPE USNM 39355, MHNG 18975 GA III 5 SEPBOP "Anton
Bruun" Cruise. 18B stat. 795D, Galapagos, 0°S 90°W, 78 m.
MATERIAL STUDIED FOR COMPARISON: USNM 21492, HOLOTYPE of Tedania toxicalis, de
Laubenfels, 1930, Point Pinos, Pacific Grove, California, 07.1925, intertidal; USNM 21490,
HOLOTYPE of 7. topsenti, De Laubenfels, 1930, Pescadero Point, near Carmel, California, 07.
1926, intertidal.
DESCRIPTION
Several small fragments (figs. 99, 100) sizes 7-12-35 x 10-15 x 2-3 mm.
Surface smooth, covered by a thin membrane which is not detachable. Several small
oscules 0.5-1 mm diameter, irregularly scattered. Small aquiferous cannals are visible
under the membrane.
Consistency : fragile, very soft.
Colour: whitish to pinkish in alcohol.
Skeleton: Ectosomal: palisade of bundles of tylotes partially included in the
membrane, and abundant onychaetes.
Choanosomal: tight meshed reticulation of short longitudinal tracts of smooth
styles connected by tylotes and abundant free onychaetes.
Spicules (Table 14): Megascleres: Thin, smooth styles (fig. 101) slightly
curved, 192-246 x 6-7 um.
TABLE 14
Spicule micrometries of Tedania (Tedania) galapagensis n. sp. T. toxicalis de Laubenfels, 1932.
USNM 21492 holotype; from literature and remeasured. Specimen from Galapagos of T. (T.)
galapagensis n. sp.
T. (Tedania) Styles Tylotes Onychaetes 1
galapagensis n.sp. Onychaetes 2
T. toxicalis
de Laubenfels, 1932
Holotype USNM 21492 100-200 200 150
Point Pinos 2] x 8-14 not observed
Holotype USNM 39355 192-226-246 179-198-234 173-188-205
Ga.III.5-18B795D x 2
00°S90°W x 6 x3 61-78-93
x 0.5-1
54 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Smooth tylotes (fig. 102) with oval microspined apices, regular diameter, 179-
234 x 3-4 um.
Microscleres: Onychaetes 1 (fig. 103) 173-205 x 2 um and 2 (fig. 104) 61-93
um x 0.5-1 um, strongly spined.
Ecology: 78 m.
Distribution: Galapagos Islands.
Remarks: T. toxicalis De Laubenfels, 1930 from California on paper sounds
close to this species, but re-examination of the type revealed some clear differences.
The choanosomal skeleton is a loose plumoreticulation with single or two spicules
interconnecting longitudinal tracts of 5-7 spicules thick; the styles have a swollen
head and frequently have one or two spines. The ectosomal spicules are smooth
tylotornotes with elongate heads. In fact, these characters make it likely that 7.
toxicalis belongs in the subgenus Trachytedania.
Other Tedania (Tedania) species in the E Pacific are T. tepitootehenuaensis
Desqueyroux-Fatindez, 1990, which has predominantly styles as choanosomal
megascleres, and T."nigrescens” sensu GREEN & GOMEZ (1986) which has onychaetes
not exceeding 100 um and thinner tylotes: 153-173 x 3-4.5 um. T. nigrescens
(Schmidt, 1862) is a junior synonym of the Mediterranean-Atlantic T. anhelans.
Tedania topsenti De Laubenfels, 1930, is a second species from California. It
differs quite substantially from T. toxicalis in having much more robust megascleres:
smooth styles averaging 360 by 14 um and smooth fusiform tylotes averaging 330 by
9 um and. Onychaetes are rare, possibly absent. The species could indeed be a
Kirkpatrickia as DE LAUBENFELS suggested himself.
Tedania (Tedania) tepitootehenuaensis Desqueyroux-Faundez, 1990
Tedania tepitootehenuaensis DESQUEYROUX-FAUNDEZ, 1990: 383, figs. 43-48.
MATERIAL STUDIED: MHNG 972229, HOLOTYPE of T. tepitootehenuaensis, Hotu Iti,
10.1972, 0 m.
Not represented in the present material, so only a brief diagnosis is given: is a
species from Easter Island with small thin styles, 240-272 x 4-9 um, straight tylotes,
oval microspined apices. Onychaetes are of two sizes: 160-285 x 2-3 um and 48-76 x
0.6 um. It differs from 7. mucosa from the Chilean coast, in having smooth, thin
styles and straight tornotes with oval heads instead of mucronate. It is probably an
endemic species.
Subgenus Tedaniopsis:
Diagnosis: Tedania with relatively long thick, smooth styles, occasionally
modified to anisostrongyles or anisoxeas, as structural megascleres. Ectosomal mega-
scleres are mucronate or tylostrongylote tornotes occasionally with one or more
vestigial spines.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 55
Tedania (Tedaniopsis) charcoti Topsent, 1907 (Figs. 105-110)
Tedania charcoti TOPSENT, 1907: 69; 1908: 30, pl. I, fig. 3, pl. IH, fig. 3, pl. V, fig. 6; Capon er
200019933263).
Tedania tenuicapitata sensu RIDLEY & DENDY, 1887: 52, pl. XI, fig. 5 [non: T. tenuicapitata
Ridley, 1881].
Tedania armata SARA, 1978: 51, figs. 30, 31.
MATERIAL STUDIED: LBIMDT 679, HOLOTYPE of Tedania charcoti Topsent, 1907, Ile
Wandel, Port Charcot, Stn. 346, 65°04'S 64°00'W, 30.03.1904, 40 m; MHNG 18845: He 3 Seno
de Otway, 53°00°S71°30’W, 16.09.1972, 260 m; MHNG 18848: He 114 Caleta Hale,
47°57°S74°39’W, 01.10.1972, 40-50 m, MHNG 10689: CH1.9, Zapallar, 32°33’S7143’W, 02.08
1985, 300 m; usnm 39353, MHNG 18854, ZMA 10982: Ga.III.3, SEPBOP "Anton Bruun” Exped.,
18A 697, Chile,35°27’S 73°01’W, 1966, 290-450 m.
MATERIAL STUDIED FOR COMPARISON: BMNH 1887:5:2:260, specimen of Tedania tenui-
capitata, Challenger Expedition, Stn. 311, off the south west coast of Patagonia, 52°45'S
73°46'W, 11.01.1876, 448 m; Ant 3, PARATYPE of Tedania armata Sarà, 1978, Spedizione
Antartica Italiana, 1882, Staz. VI, Tierra del Fuego, leg. D. Vinciguerra, .
DESCRIPTION
Sponge irregularly massive to lamellate (figs. 105, 106) size 27-30-60 x 19-40-
50 mm. Surface conulose, mammilate or minutely tuberculate. Oscules numerous, 1-5
mm diameter. Surface uneven and punctate, with a thin membrane.
Consistency: Soft, elastic, fragile.
Colour: Grayish to brownish in alcohol.
Skeleton: Ectosomal: A palisade of tornotes, free and in bundles. Abundant
free onychaetes
Choanosomal: Loose, irregular and confused reticulation of longitudinal tracts
of styles, and abundant free onychaetes.
Spicules (Table 15): Megascleres: Long, slightly bent, smooth styles (fig.107),
361-445 x 8-16 um.
Smooth mucronate straight tornotes (fig. 108), regular diameter, 202-300 x 4-6
um.
Microscleres: Onychaetes 1 (fig. 109) 112-392 x 2 um and 2 (fig. 110) 35-97x
0.5 um., straight, with numerous strong spines.
Ecology: Substrat, stones, 40-500 m.
Distribution: Chilean coast, 32°S - 53° S; east of the Strait of Magellan, 51°S
65°W, as T. tenuicapitata sensu RIDLEY & DENDY, 1887, Tierra del Fuego; 54°S
59°W, as T. armata Sarà, 1978; Wandel Island, Port Charcot, 65°S 64°W as 7.
charcoti, Topsent, 1907.
Remarks: This species is close to T. tenuicapitata, but differs clearly in having
much shorter onychaetes 1; the tornotes are also different, having a more pronounced
pointed end at one or both ends. The Challenger specimen assigned to tenuicapitata
conforms to the type of T. charcoti and our specimens.
56 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
TABLE 15
Spicule micrometries of Tedania (Tedaniopsis) charcoti Topsent. T. charcoti Topsent, 1907,
HOLOTYPE LBIMDT 679; T. armata Sara, 1978 Ant 3, PARATYPE; from literature and remeasured.
Specimens from Chile of 7 (T.) charcoti.
T.(Tedaniopsis)
Styles Tornotes Onychaetes 1
charcoti Topsent, 1907 Onychaetes 2
T. charcoti
Topsent, 1907
I Wandel. Port Charcot
HOLOTYPE LBIMDT 679 420-450 305-340 250-265
7 18} x 10 90-120
Remeasured 410-463-549 314-325-353 235-264-365
x 10-12 x 8-10-12 98-116-127
Sara, 1978
T. del Fuego
PARATYPE ANT.3 300-350 240 x 2-3 200-280
x 6-8 150-180
Remeasured 308-336-373 219-243-267 162-174-194
x8 x 4 97-126-146
He 3 361-374-401 202-233-271 112-148-176
Seno de Otway x x 6 48-59-70
He 114 298-339-355 208-237-259 160-177-204
Caleta Hale x 8-9-10 x6 35-44-54
CHI-9 377-403-426 262-274-291 147-271-393
Zapallar x 9-11-13 x 5-6 38-60-96
GA III 3a USNM 39353 381-415-445 251-269-300 154-206-292
18A697 x 12-14-16 x4 65-83-97
22 TES TBE OI
Tedania (Tedaniopsis) tenuicapitata Ridley, 1881 (Figs. 111-116)
Tedania tenuicapitata RIDLEY, 1881: 124, pl. xi, fig. 1.
[non Tedania tenuicapitata sensu RIDLEY & DENDY, 1887: 52, pl. xi, fig. 5, = T. charcoti
Topsent, 1907].
MATERIAL STUDIED: BMNH:1879:12:27:12, HOLOTYPE of Tedania tenuicapitata Ridley,
1881, Trinidad Channel, 50°00°S 75°00°W, near Madre de Dios Island, off SW Patagonia, 54.8
m. MHNG 18976, Chi. 22, 52°42’S 75°23’W, 30 m.
MATERIAL STUDIED FOR COMPARISON: LBIMDT 679, HOLOTYPE of Tedania charcoti
Topsent, 1967, Ile Wandel, Port Charcot, Stn. 346, 65°04'S 64°00'W, 30.03.1904, 40 m; BMNH
1887:5:2:260, specimen de Tedania tenuicapitata sensu Ridley & Dendy, 1887, Challenger
Exped. Stn. 311, off the south west coast of Patagonia, 52°45'S 73°46'W, 21.01.1876, 448 m.
Description: One specimen 27 x 14 x 10 mm (figs 111, 112) and a fragment of
a massive sponge. Surface smooth, covered by a thin membrane. Small oscules (1.5-2
mm diameter) irregularly scattered.
Consistency: Very soft and fragile.
S-E PACIFIC SPECIES OF POECILOSCLERIDA ST
Fics 99-110. Tedania (Tedania) galapagensis n.sp. 99, holotype, USNM 39355, fragment of
specimen. 100, enlarged view of the surface. 101, smooth styles slightly curved. 102, smooth
tylotes with oval microspined apices. 103, 104, onychaetes 1 and 2 strongly spined. Scales:
figs. 101-103 = Sum; fig. 104 = lum. Tedania (Tedaniopsis) charcoti Topsent, 1907. 105, mas-
sive specimen from Caleta Hale. 106, enlarged view of the surface. 107, long smooth slightly
bent styles, 108, smooth mucronate tornotes. 109, 110, onychaetes 1 and 2. Scales: figs. 107,
108 = Sum; fig. 109 = 2um; fig. 110 = lum.
58 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Colour: Whitish to light beige in alcohol.
Skeleton: Ectosomal, dense palisade of smooth tornotes and onychaetes.
Choanosomal: Loose and irregular to slightly isotropic reticulation of styles
and onychaetes.
Spicules (Table 16) Megascleres: long, smooth styles (fig.113), slightly bent,
437-543 x 8-16 um.
TABLE 16
Spicule micrometries of: Tedania (Tedaniopsis) tenuicapitata Ridley, 1881, HOLOTYPE BMNH
1879:12:27:12; from literature and remeasured. Specimen from Chile of T. (T.) tenuicapitata.
T. (Tedaniopsis) Styles Tomotes Onychaetes 1
tenuicapitata Onychaetes 2
Ridley, 1881
T. tenuicapitata
Ridley, 1881 380 x 12.7 279 x 6 316
BMNH 1879:12:27:12
Trinidad Channel
Remeasured 296-340-387 185-221-270 132-263-327
x 12-13 x4 52-64-75
CHI-22 437-500-543 308-349-389 121-178-267
52°42’S 74°23’W x 8-14-16 x8 xD
54-76-96
x Il
Smooth mucronate tornotes (fig. 114) with strongly pointed apices, 308-389 x
8 um.
Microscleres: Onychaetes | (fig. 115) 121-267 x 2 um, and 2 (fig. 116) 54-96
x 0.5um, strongly spined.
Ecology: Pebbles and stones, 30 m.
Distribution: Chilean coast, 50°S - 52°S.
Remarks: As discussed above, the nearest relative is T. charcoti, which has
smaller onychaetes 1, T. tenuicapitata sensu Ridley & Dendy, 1887 conforms to that
species. A further closely related species is T. infundibuliformis Ridley & Dendy,
1886 which is cup-shaped, but similar in spiculation.
Tedania (Tedaniopsis) infundibuliformis Ridley & Dendy, 1886
Tedania infundibuliformis RIDLEY & DENDY, 1886: 335; 1887: 54, pl. xi, fig. 1, pl. xxix, figs. 2,
2a.
MATERIAL STUDIED: BMNH 1887:5:2:151, HOLOTYPE of Tedania infundibuliformis Ridley
& Dendy, 1886, Challenger Expedition, Stn. 311, off the south west coast of Patagonia, 52°45'S
73°46'W, 21.01.1876, 448 m.
Not represented in our material so only a brief diagnosis can be given:
S-E PACIFIC SPECIES OF POECILOSCLERIDA 59
Fics 111-116
Tedania (Tedaniopsis) tenuicapitata Ridley, 1881. 111, massive specimen from 52°S 75°W.
112, enlarged view of the surface. 113, smooth style, strongly pointed apex. 114, smooth
mucronate tornote. 115, 116, onychaetes 1 and 2, strongly spined. Scales: figs. 113, 114 = Sum;
figs. 115, 116 = 2um.
Cup-shaped. Skeleton a loose reticulation. Long smooth styles averaging 500
by 12 um. Short, smooth tylotes with a few very faint spines on the heads, 280 by 6
um. Onychaete 1, 360 um, onychaetes 2, 60-70 um.
Distribution: Chilean coast 52°S.
Subgenus Trachytedania Ridley, 1881
Diagnosis: Tedania of which the structural spicules are relatively small styles
at least some of which show (a few) spines. The ectosomal megascleres are mucro-
nate or oxeote tornotes.
Tedania (Trachytedania) mucosa Thiele, 1905(Figs 117-128)
Tedania mucosa THIELE, 1905: 430, figs. 50a-c.; BURTON, 1934: 27.
Tedania excavata THIELE, 1905: 431, figs. Sla-c.
Tedania pectinicola THIELE, 1905,: 432, figs. 52a-d.
Tedania fuegiensis THIELE, 1905: 433, figs. 53a-d.
60 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
MATERIAL STUDIED: ZMB 3284, HOLOTYPE of Tedania mucosa Thiele, 1905, Calbuco,
41°46'S 73°08'W; ZMB 3285, HOLOTYPE of T. excavata Thiele, 1905, Calbuco; ZMB 3286,
HOLOTYPE of T. pectinicola Thiele, 1905, Calbuco; ZMB 3287, HOLOTYPE of T. fuegiensis Thiele,
1905, Cabo del Espiritu Santo, 52°58'S 68°36'W, Tierra del Fuego; MHNG 18857, 18977: He 25,
179, Seno de Otway, 53°00’S 71°30’W, 17.09.1972, 30 m; MHNG 18858, 10690: He 61, 127,
Caleta Hale, 47°57’S 74°37’W, 01.10.1972, 40-50 m; MHNG 18860: He 189, Bahia Scotchwell,
55°28’S 68°07’W, 10.1972, 50-95 m; MHNG 18861: VALD. 10.1, Canal Quinchao, 42°25’S
72°35’W; MHNG 18859, 18876: VALD. 46.1, 53.2, Palena 43°45’S 73°00’W, 10.01.1972, 10-15
m; MHNG 18862: VALD.53.5a, Islote Caleta Lobato, 45°05’S74°47’W, 10-15 m; MHNG 18863,
18873, 18983, 18993: Mont. 8, 9, 10, 30, San Antonio, 32°48’S 71°23’W, 10-15 m; MHNG
18864, 18823: Co.8, VALD. 14.1, Quetalmahue, 41°32’S 73°52’W; MHNG 18999, 18865, 19228,
19229: Co.93.38, 48, 49, 53, Pelluco, 41°30°S 72°54’W; MHNG 18866, 19230, 19231: MNHNC
25, 36, 57, 43°15’S 74°32’W, 266 m; MHNG 10691: Por.2, Pullinque, 39°33’S 72°11’W, 7 m,
DESCRIPTION
Several specimens and fragments. Sponge polymorphe, massive, (figs. 117,
118) irregularly roundish and lobate to lamellate and tubiforme 77-112 x 55-84 x 28-
89 mm, or repent and sending out cylindrical processes or stolon-like branches (figs.
119, 120), 40-55 x 37-59 x 9-17 mm. Surface uneven, tuberculate and conulose.
Minutely and entirely punctate and with numerous aquiferous canals visible at the
surface. Oscules at the summit of the conules, irregularly scattered on the surface, or
arranged along the edge of branches.
Consistency: Hard, slightly compressible.
Colour: Brown to reddish alive, colouring the alcohol when fixed and
producing great amounts of mucus.
Skeleton: Ectosomal skeleton: a perpendicular palisade of densely arranged
mucronate tornotes. Onychaetes are numerous.
Choanosomal skeleton: Loose or dense reticulation of tracts of styles.
Onychaetes abundant, free or in bundles.
Spicules (Table 17): Megascleres: Smooth styles (figs. 121, [122 holotype])
slightly bent, with sharp to roundish apex, 163-320 x 5-16 um.
Mucronate tornotes (figs. [123 holotype] 124, 125) regular diameter, 128-267x
3-6 um.
Microscleres: Onychaetes 1, 99-235 x 2 um and 2, 26-99 x 1 um, (figs. [126
holotype] 127, [128 holotype]) some of them present a tyle near the base.
Ecology: On stones, 7-266 m depth.
Distribution: Chilean coast, 32°S - 55°S; Argentinan Atlantic coast up mouth
of Rio de La Plata (37°S), BURTON, 1934.
Remarks: THIELE (1905) described four species based on minor differences
which were found to be partly incorrect: T. excavata was stated to have a digitate
growth form and to possess only a single onychaete category. T. pectinicola would be
encrusting and have a single onychaete category with distinct tyles ("tylonychaetes").
T. fuegiensis would have styles with occasional spines. We found a large variation in
growth forms among specimens all of which in their skeletal characteristics were
indistinguishable from the type of T. mucosa. All type specimens of the four Thiele
species had two categories of onychaetes, and minor differences in spicule sizes
S-E PACIFIC SPECIES OF POECILOSCLERIDA 61
117 118
119
Fics 117-128. Tedania (Trachytedania) mucosa Thiele, 1905. 117, 118, lamellate to massive
specimen, from 43°S Chile, and enlarged view of the surface. 119, 120, repent specimen, and
enlarged view of the surface. 121, smooth styles, slightly bent. 122, ZMB 3284, holotype of 7.
mucosa Thiele, 1905 from Calbuco, smooth styles. 123, zmB 3286 holotype, of T. pectinicola
Thiele, 1905 from Calbuco, mucronate smooth tornotes 124, Chilean specimen from Calbuco,
mucronate smooth tornote. 125, zMB 3284, mucronate smooth tornotes. 126, ZMB 3286, onychaete
1. 127, Chilean specimen from Calbuco, onychaete 1. 128, zmB 3284, onychaetes 2.
Scales: fig. 121 = 10pm; figs. 122-127 = Sum; fig. 128 = 2um.
62 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
TABLE 17
Spicule micrometries of Tedania (Trachytedania) mucosa Thiele, 1905. Tedania mucosa
Thiele, 1905 ZMB 3284 holotype; T. fuegiensis Thiele, 1905 ZMB 3287 holotype; T. pectinicola
Thiele, 1905 ZMB 3286 holotype; 7. excavata Thiele, 1905 zmB 3285 holotype; from literature
and remeasured. Specimens from Chile of T. (T.) mucosa.
Onychaetes 1
T. (Trachytedania) Styles Tornotes
mucosa Onychaetes 2
Thiele, 1905
T. mucosa
HOLOTYPE ZMB 3284 280-310 190-210 190-210
Calbuco x 14 x 6 not observed
Remeasured 304-364-349 208-213-220 204-219-233
x 12-13-16 x 5-6-7 40-53-64
T. fuegiensis
Thiele, 1905
HOLOTYPE ZMB 3287 200-225 160-200 120
T. del Fuego x8 y6 not observed
Remeasured 186-208-224 176-179-198 112-133-163
x 6-7-10 x 3-5-6 Ko)
74-94-10
x 0.5-1
T. pectinicola
Thiele, 1905
HOLOTYPE ZMB 3286 240 x 8 160 x 5 140
Calbuco not observed
Remeasured 205-225-246 147-158-173 102-119-134
x 6 x3 xe
45-73-93
x |
T. excavata
HOLOTYPE ZMB 3285 240 x 13-15 150 x 6 190
Calbuco not observed
Remeasured 208-253-275 157-178-192 122-155-186
x 10-12-13 x 6 xd
58-78-99
x |
He 25 251-370-413 194-246-267 105-156-235
Seno De Otway x8 x4 x 2
49-74-97
x |
He 179 275-299-320 166-207-234 157-194-230
Seno De Otway x 10-12-13 x 3-6 x 2)
35-54-83
x |
He 61 202-232-246 163-178-192 106-149-189
Caleta Hale x 6-9-13 x 3-5-6 x 2
26-65-99
x |
He 127
Caleta Hale
Vald. 46.1a
Palena
Vald. 53.2
Palena
He 189
BahiaScotchwell
Vald. 10.1
Canal Quinchao
Vald. 53.5a
I. Caleta Lobato
Mont. 8
San Antonio
Mont. 9
San Antonio
Mont. 10
San Antonio
Co. 8
Quetalmahue
Co. 93-48
Pelluco
Co. 93.49
Pelluco
234-244-272
x 6-9-13
202-247-278
x 13-14-16
163-237-269
x 6-13-16
199-218-234
x 6-10-12
x 6-7-10
228-244-262
x 9-10-12
170-189-205
x 5-6
237-254-270
x 10
220-237-256
qx 6-8-10
237-256-282
x 6-8-10
S-E PACIFIC SPECIES OF POECILOSCLERIDA
160-174-180
x 5-6
163-178-192
x 3-6
150-170-186
x 3-6
152-167-180
x 4-5
186-201-224
x 3-4-6
176-193-208
x 3-5
180-189-198
x 5-6
186-206-221
x6
128-148-160
x3
179-199-214
x 6
178-209-251
x 6
182-215-246
x 6
63
99-143-176
x 2
35-44-51
x |
112-145-182
x 2
106-160-205
RE
32-59-99
x |
109-154-183
2)
31-42-74
x |
102-144-179
xen,
29-52-86
x |
121-171-192
x2
154-161-166
x2
32-43-61
x |
144-175-198
x2
48-51-58
x |
102-113-144
x)
125-155-176
XD
43-51-70
el
138-155-176
160-170-176
Kee)
35-44-64
x |
64 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
Co. 93.53 269-306-326 202-207-211 208-218-224
Pelluco x 10 x 6 xD
38-48-64
x |
MNHNC.25 218-237-254 163-173-208 134-163-192
43°15°S 74932’ W x 8-13-16 x 5-6 xD
35-42-47
x |
MNHNC.36 224-241-256 157-171-208 80-115-179
43°15°S 74°32°W x 10-12-13 x 5-6 2
35-40-45
x |
MNHNC.57 210-240-256 166-173-182 86-122-144
43°15’S 74°32°W x 10-12-16 x 6 x 2
35-40-45
x il
Por. 2 201-229-246 179-192-208 99-131-166
Pullinque x 5-6 33 x2
35-44-51
x |
reported by Thiele could not be substantiated in remeasured series of spicules in slides
made from the four types. Occasional spines were found in several specimens,
including the type of T. mucosa. It is clear to us that all four are part of the same
rather variable species.
T. fuegiensis sensu SARA, 1978 is conspecific with T. spinata.
T. mucosa is similar to T. spinata (Ridley, 1881), but it has clearly thicker,
more robust styles, and the skeleton is denser and more confused.
Tedania (Trachytedania) spinata (Ridley, 1881) (Figs. 129-134)
Trachytedania spinata RIDLEY, 1881:122. pl. x, fig. 10.
Tedania murdochi TOPSENT, 1913: 629, pl. v, fig.
Tedania corticata SARA, 1978: 56, figs.34-35
Tedania fuegiensis sensu Sara, 1978 [non: Tedania fuegiensis Thiele, 1905: 50 = T. mucosa]
Tedania laminariae SARA, 1978: 54, figs. 32, 33.
o
Pe
Is
MATERIAL STUDIED: BMNH:1879:12:27:9, HOLOTYPE of Tedania (Trachytedania) spinata
Ridley, 1881, Portland Bay, 50°15°S 74°44’W, Chili (opposite the chief island of Madre de
Dios Archipelago) 18.8 m; LBIMDT 1591, HOLOTYPE of Tedania murdochi Topsent, 1913, Stn.
118, Stanley, Falkland Islands, 01.02.1904, 20 m; SAI 7, PARATYPE of Tedania corticata Sara,
1978 Spedizione Antartica Italiana, 1882 (locality and date not precised), SAI 1, 8, Tedania
fuegiensis sensu Sara, 1978, Spedizione Antartica Italiana, 1882, (locality and date not
precised); C-75, PARATYPE of Tedania laminariae Sara, 1978, Exped. 7, AMF MARES, GRST SAI,
Bahia Golondrina, Ushuaia, 54°49’S 68°16’W, 16.01.1974, on the beach, Sara coll.; MHNG
18877, 19232-19234: HE 15, 43, 63, 69, Puerto Caracciolo , 50°26’S 75°09'W, 09.10.1972, 25
m; MHNG 18878, 19235, 19236: HE 30, 40, 50, Isla Madre de Dios, Caleta Henry, 50°00’S
75°19’W, 6-28 m; MHNG 18825: HE 39, Puerto Eden , 49°10'S 74°23'W, 05.10.1972, 20 m;
MHNG 18826: VALD. 1, Punta Caucacura, 43°08'S 73°30'W, 16.01.1972, 20 m; MHNG 18827:
S-E PACIFIC SPECIES OF POECILOSCLERIDA 65
VALD. 5, Estero Quinchao, 42°32'S 73°26'W, 15.01.1972, 10 m; MHNG 18832: VALD. 22.1,
Compu , 42°52'S 73°33'W, 14.12.1971, 15 m; MHNG18836: Vald. 32, Linao, 41°57'S 73°33'W,
27.06.1971, 15 m; MHNG 18837: VALD. 43, Quintupeu, 42°10'S 72°24'W, 23.07.1971, 10 m:
MHNG 18864: VALD. 51, Corral, 39°52'S 73°25'W, 29.03.1981, 10 m; MHNG 18855: Co.9, Bahia
de Ancud, 41°50'S 73°47'W, 11.1983, 10 m; MHNG 18868: Co. 93.34, Pelluco, 41°30'S
72°54'W, 09.12.1992, 15 m; MHNG 18869: MNHNC 28c, Golfo Corcovado, Off Morro Yeli,
43°31'S 73°03'W, 30.05.1966, 69 m, MHNG 10692: Por. 14, Quetalmahue, 41°50'S 73°52'W,
04.1984, 10-15 m.
Fics 129-134
Tedania (Trachytedania) spinata (Ridley, 1881) 129, massive specimen from Puerto Eden. 130,
enlarged view of the surface. 131, small smooth styles. 132, smooth mucronate tornotes. 133,
134, onychaetes 1 and 2, strongly spined. Scales: figs. 131-133 = Sum: fig. 134 = Ium.
DESCRIPTION
Massive sponge (figs. 129, 130): 82-115 x 30-84 mm. Surface smooth, covered
by a translucent membrane, which is darker than the choanosome. Oscules, 2-3 mm
diameter, irregularly scattered.
Consistency: Compact, soft, compressible.
Colour: Surface dark gray, internally brownish to grayish.
Skeleton: Surface, dense perpendicular palisade of tornotes reinforced by the
terminal part of styles and abundant onychaetes.
66 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
TABLE 18
Spicule micrometries of: Tedania (Trachytedania) spinata (Ridley, 1881). Trachytedania
spinata Ridley 1881 BMNH 1879:12:27:9 holotype; T. murdochi Topsent, 1913 LBIMDT 1591
holotype; T. corticata Sara, 1978 SAI-7, paratype; T. fuegiensis sensu Sara, 1978 SAI-8, speci-
men from Tierra del Fuego; T. laminaria Sara, 1978 c-75 paratype from Ushuaia; from litera-
ture and remeasured. Specimens from Chile of T. (T. ) spinata.
134-163-192
MNHNC.25 218-237-254 163-173-208
T. (Trachytedania) Styles Tornotes Onychaetes |
spinata (Ridley, 1881) Onychaetes 2
Trachytedania spinata
Ridley, 1881
BMNH 1879:12:27:9 165 x 6 177-187 x 4 SO
Portland bay
Remeasured 148-159-163 148-167-191 117-131-150
x 4-5 x 3-4 x)
T. murdochi
Topsent, 1913
Holotype LBIMDT 1591 225-250 200-240 40-175
Stanley x 7-10 x 5-6
Remeasured 171-216-240 191-208-230 109-140-168
x 8-10 x4 x2
47-78
0.5
T. corticata
Sara, 1978
Paratype SAI-7 200-240 200-240 100-240
x 5-7 x 2-5
Remeasured 218-229-257 200-222-242 136-181-210
x?
T. fuegiensis
sensu SARÀ, 1978 200-240 160-200 100-200
SAI-8 x 6-10 x 4-6
T. laminariae
Sara, 1978
Paratype C-75 150-200-260 180-200 80-200
Ushuaia x 2-4-6 x 1-3-4 x 0.5-1
Remeasured 190-209-240 180-193-200 115-122-150
XS-0-7 x 4-6 x 0.5-1
38-48-60
x 0.5-1
He 15 217-240-256 172-185-230 115-162-201
Puerto Caracciolo x 3-7-10 x 3-6 x À
29-61-90
xl
He 43 192-214-237 122-167-214 106-148-214
Puerto Caracciolo x 6-8-10 x 3-5-6 xD
48-62-73
He 63 195-236-262 202-222-234 106-127-147
Puerto Caracciolo x 3-6 xs) 2
38-63-99
x Il
He 69
Puerto Caracciolo
He 30
Isla Madre de Dios
He 40
Isla Madre de Dios
He 50
Isla Madre de Dios
He 39
Puerto Edén
Vald. 1
Punta Caucacura
Vald. 5
Estero Quinchao
Vald. 22-1
Compu
Vald. 32
Linao
Vald. 43
Quintupeu
Vald. 51
Corral
Co.9
Bahia de Ancud
Co.93-34
Pelluco
214-239-266
x 6-7-10
161-219-253
x 3-5-6
189-214-227
x 6-7-10
211-248-277
x 6-8-9
230-251-275
x 6-7-10
240-246-278
x (0-7 10
224-242-256
x 6-8-10
230-251-307
x 6-7-10
S-E PACIFIC SPECIES OF POECILOSCLERIDA
154-180-221
x 3-6
179-203-230
x 3-5
192-206-283
x 3-4-5
186-199-214
x 3-6
153-165-170
X
191-209-242
x4
150-163-179
x 3-4-5
198-210-230
x 3-4-6
166-193-221
x3
198-215-230
x3
170-184-201
x 3-5-6
186-201-214
3-4
67
106-144-186
XD,
32-62-99
x |
112-131-157
Xen,
Absent
128-199-154
x)
Absent
189-209-214
KEL,
Absent
128-155-195
Xo,
Absent
999-119-144
x2
32-43-73
x 0.5
152-154-171
x 2
35-49-81
xl
93-115-157
xD
38-50-89
x |
144-157-166
XD,
Absent
128-174-202
x 2
35-52-93
x |
147-171-189
X À
32-57-99
x |
93-134-160
x2
32-43-54
al
112-159-192
x 2
Absent
68 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
MNHNC 28c 214-227-234 160-167-170 96-124-144
Golfo Corcovado x 6-9-10 x3 51-55-70
xe
Por.14 208-243-284 182-201-214 128-163-228
Quetalmahue x 6-8-10 x 3-5-6 x)
35-43-86
ll
Choanosomal: Loose, irregular to isotropic reticulation of longitudinal (3-4
spicules) and transversal (2-3 spicules) tracts of smooth styles. Abundant free and in
bundles onychaetes.
Spicules (Table 18): Megascleres: predominantly relatively small smooth
styles (fig. 131) with occasionally one or a few at the base spines, 161-284 x 3-10 um.
Smooth mucronate tornotes (fig. 132) abruptly pointed, 122-283 x 3-6 um.
Microscleres: Onychaetes 1 (fig. 133) 93-228 x 2 um, and 2 (fig. 134) 29-99 x
0.5 um length, straight, strongly spined.
Ecology: Stones pebbles, sand, 10-69 m.
Distribution: Chilean coast, 39°S - 50°S, Atlantic coast, 50°-54°S, Falkland
Islands.
Remarks: RIDLEY (1881) emphasized the presence of spines on the heads of the
styles when describing T. spinata, but re-examination of the type specimen by
BURTON (1932) and by us failed to reveal any but the faintest and rarest of spines in
Ridley's material. Such occasional spines are found in many species of Tedania and
not worthy of emphasis.
The type specimens of T. murdochi Topsent, 1904, T. corticata Sara, 1978 and
T. laminariae Sara, 1978 were re-examined and could not be separated from T.
spinata, on skeletal structure, spicule sizes and form, so their conspecificity is likely.
T. toxicalis De Laubenfels, 1930 (cf. above) from California is a close relative,
also possessing styles with vestigial spines. It differs from 7. spinata in having
tylotornotes in stead of mucronate tornotes; onychaetes 2 are distinctly longer in T.
toxicalis (100-130 um).
Tedania (Trachytedania) patagonica Ridley & Dendy, 1886
Trachytedania patagonica RIDLEY & DENDY, 1886:336; 1887: 57, pl.xxiii, figs. 6, 6a-c.
? Tedania biraphidora BOURY-ESNAULT, 1973: 281, fig. 36.
Tedania inflata SARÀ, 1978: 59, figs. 36, 37.
MATERIAL STUDIED: SAI 5, PARATYPE of Tedania inflata Sarà, 1978, Spedizione
Antartica Italiana, Patagonia.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 69
Not represented in our material, so only a brief diagnosis is given here:
Massive sponge, soft and fragile. Surface uneven and hispid. Skeleton as a
loose isodictyal network. Spicules: Styles fully spined all over, 350 x 13 pm.
Oxeotornotes 245 x 7 um. Onychaetes 100-200 um.
Distribution: Chilean coast, 50°S, Patagonia, Brazil, 24°S (?).
If a separate genus for Tedania With spined styles would have needed to be
erected, then this species would much more deserve to be in that genus than 7.
spinata. To date, this is the only known species of Tedania with fully spined styles.
KEY TO THE SUBGENERA AND SPECIES OF Tedania OF THE SE PACIFIC COAST
(N°)
W |
Long (from 300 to 700 um) thick smooth styles, tornotes of diverse
SI APESPE ERE E de ne de Subgenus Tedaniopsis .... 2
Short (from 150 to 300 um) to relatively short styles. ................... 3
iornotesstylotomoteser een T. (Tedaniopsis) infundibulifomis
Ihomnotes-mueronate 0r'0x0to1noteser an TOTI 4
Smooth styles, tylotornotes with microspined apices. Subgenus Tedania ...8
Smooth or spined styles. With oxeote tornotes or mucronate tornotes
CIRIE ay earn beens Wie eee a ten a Subgenus Trachytedania ....6
With oxotornotes, long onychaetes | (800 um) small onychaestes 2
(SO Imre Se T. (Tedaniopsis) massa
With mucronate tornotes, onychaetes | up 400 pm, onychaetes 2 up
OO I PES etc ses rea ace et RSR A a 5
Onychaetes 2 occasionally "tylonychaetes". Chile, S.W. Atlantic and
INTRO i en Lic > Tedania (Tedaniopsis) charcoti
Onychaetes 2 not "tylonychaetes".Chile (50-52°S)
SESIA ERE RIO IA A e T.(Tedaniopsis) tenuicapitata
Smooth styles with occasional spines, smooth mucronate tornotes. . . . . . U
Styles fully spined, smooth oxeote tornotes. . . . 7. (Trachytedania) patagonica
Thin styles (from 3 to 10 um}, Chilean, S.W. Atlantic, Falkland Islands
SON ER O rns II T. (Trachytedania) spinata
Thicker, robust styles from 6 to 16 um), Chilean, S.W. Atlantic sponge
i Cai ee en a US T. (Trachytedania) mucosa
Tylotes oval spined apices, from 190 to 250 um, Onychaetes 1 from
160 to 285 um, onychaetes 2 from 48 to 76 um. Easter Island sponge
toi ER T. (Tedania) tepitootehenuaensis
Tylotes from 197 to 234 um. Onychaetes 1 from 197 to 234 um, ony-
chaetes 2 from 60 to 90 um.Galapagos sponge..... T. (Tedania) galapagensis
70 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
REVIEW OF THE FAMILY TEDANIIDAE
Further genera assigned to the Tedaniidae, but not represented in the present
material are the following:
Genus Tedanione Wilson, 1894
Type species T. foetida Wilson, 1894 (topotypical specimen examined).
Diagnosis: Tedaniidae with smooth strongyles as the only megascleres.
Microscleres as usual onychaetes in single or two size classes.
Remarks: Tedanione foetida Wilson, 1894 (junior synonym Hemitedania baki
Van Soest, 1984) from the Caribbean is a thinly encrusting cryptic sponge, with a
loosely plumose skeleton. The major synapomorphy is the possession of a single type
of megasclere, viz. smooth strongyles. In this respect, the genus is similar to
Hemitedania, which, however, has sharply pointed oxeas as megascleres. The
onychaetes of foetida often have distinct tyles, a feature emphasized by VAN SOEST,
1984, but now thought to be of little significance because of widespread occurrence in
various unrelated Tedaniidae. Since at least a second species of Tedanione, T. wilsoni
Dendy, 1922 exists, the genus is considered valid.
Genus Hemitedania Hailmann, 1914
Synonym: Tedaniopsamma Burton, 1934
Type species Amorphina anonyma Carter, 1886 (type specimen examined).
Diagnosis: Tedaniidae with sharp pointed oxeas as the only megascleres.
Microscleres as usual onychaetes.
Remarks: Hemitedania anonyma (Carter, 1886) from South Australia is an
elaborate sponge containing a large quantity of sand grains. It is one of the "sand
sponges", i.e. sponges of various families of sponges which apparently replaced parts
or whole of their skeleton by sand grains (WIEDENMAYER, 1989), and the possibility
cannot be excluded that the choanosomal styles have become lost, while the ecto-
somal oxea-like tornotes have been retained. The genus is monotypical, although
Hemitedania baki Van Soest, 1984, was reported from the Caribbean and a Hemi-
tedania spec. from West Africa by VAN SOEST, 1993; both records, however, very
probably concern Tedanione foetida. In view of its deviating characters it is proposed
to keep the genus as valid and separate from Tedanione.
Tedaniopsamma Burton, 1934 (with type species Hircinia flabellopalmata
Carter, 1885: 113 (holotype figured in Von Lendenfeld, 1889 as Sigmatella) (not exa-
mined) is a probable synonym of Hemitedania. T. flabellopalmata is also a sand
sponge, and its tedaniid nature has not been established with certainty, because the spi-
cules, including the"raphides" have never been properly described or figured. If these
raphides are indeed onychaetes, then the species would fall to Hemitedania on account
of its curved 180 um long oxeas/strongyles (due to their thinness the exact nature has
not become clear) and sand cored fibres. A second species of Tedaniopsamma des-
cribed by VACELET & VASSEUR (1971), T. arenosa, appears to be a normal Tedania
(Tedania).
S-E PACIFIC SPECIES OF POECILOSCLERIDA gi
Three more genera have been associated with the family, viz. Kirkpatrickia
Topsent, 1912, Tedaniophorbas De Laubenfels, 1936 and Tedandoryx De Laubenfels,
1954. However, the type species of these genera do not possess onychaetes, and the
two latter have chelas. They belong to other families of the Poecilosclerida: Kirk-
patrickia goes to Anchinoidae because of its surface areoles; Tedaniophorbas is a
probable junior synonym of Megaciella (family lophonidae), and Tedandoryx either
is a Myxillidae, or possibly represents a commonplace Tedania (Tedania) conta-
minated with some arcuate isochelas.
To summarize, the family Tedaniidae comprises three valid genera, Tedania
(with subgenera Tedania, Trachytedania and Tedaniopsis), Tedanione, and Hemite-
dania.
DISCUSSION
SYSTEMATICS
The three genera treated here were previously considered members of a loosely
defined giant family Myxillidae. Tedania was more often than the other two genera,
separated into a family of its own (Tedaniidae) because of its lack of chelas and its
possession of onychaetes. For inexplicable reasons, /ophon did not receive the same
appreciation although in its peculiar bipocilla and its palmate anisochelas it had
strongly deviating characters, too. The proposal (HAJDU et al. 1994) to put these three
genera in three different families is here supported, because it increases the internal
consistency of the poecilosclerid families.
Judged on characters other than the anchorate chelas the family Crambeidae is
not an obvious sistergroup of Myxillidae. Relationships within the suborder Myxillina
still need to be further evaluated. It has been postulated by HAJDU er al. (1994) that
the arcuate chela is plesiomorphous, which means that families with arcuate chelas
(Coelosphaeridae, Crellidae, Anchinoidae, Hymedesmiidae) need additional defining
characters, and cannot be considered closely related by the possession of the arcuate
condition. The latter three families share similar surface structures, the pore sieves,
which indicates they might form a monophyletic group. Those three families as well
as Coelosphaeridae and Myxillidae share the possession of diactinal ectosomal mega-
scleres (as opposed to the monactinal ones in the family Crambeidae), which is
tentatively assumed to indicate that Crambeidae fall outside the five family group.
To test these ideas a preliminary character analysis of some major Poecilo-
sclerid families was performed.
The following families were studied: Mycalidae (outgroup), Microcionidae (to
seek support for the Microcionid nature of Iophonidae), Iophonidae, Myxillidae,
Tedaniidae, Coelosphaeridae sensu HAJDU et al. 1994 (to investigate whether they are
close to Myxillidae), Crambeidae, Hymedesmiidae, Crellidae and Anchinoidae (to see
whether arcuate and anchorate chela morphology is convergent with other morpho-
logical characters).
The following characters and states were distinguished:
72 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
1. Chela morphology (with states: absent, palmate, arcuate and anchorate)
2. Ectosomal megascleres (with states: absent, monactinal and diactinal)
3. Spination of ectosomal megascleres (with states: absent and present; outgroup
is scored as "?")
Echinating acanthostyles (with states: absent and present)
Choanosomal reticulation (with states: plumose, anisotropic and isodictyal)
Sigmas (with states: present and absent)
Toxas (with states: present and absent)
Pore sieves (with states: absent and present)
CORNE
The matrix of families and character states scores was analyzed with PAUP 3.1
(SWOFFORD, 1993), using the heuristics and unweighted characters option. The result
was a single tree of 18 steps length (consistency index 0.667, retention index 0.684,
rescaled consistency index 0.456). This tree is depicted in Fig. 135. Only characters 6
and 8 had a consistency of 1.00, indicating a lot of conflicting character distributions
and it is stressed here that this result is considered tentative.
Microcionidae
Iophonidae
Tedaniidae
Crambeidae
Myxillidae
Coelosphaeridae
Anchinoidae
Hymedesmiidae
Crellidae
Mycalidae
ÉIGAIES
Cladogram obtained by treating the matrix of families and character states in Table 19. Tree CI
= 0.667, RI = 0684, RCI = 0456.
S-E PACIFIC SPECIES OF POECILOSCLERIDA 73
Iophonidae is indeed the sister group of Microcionidae (but Raspailiidae was
not included in the analysis), Myxillidae and Coelosphaeridae are also sister groups,
clearly separated from the closely related family group Hymedesmiidae-Crellidae-
Anchinoidae. The position of Crambeidae and Tedaniidae on the branch of Micro-
cionidae-Iophonidae indicates the lack of support for a clear association of these
families with the "myxilline" group. There is only a single character shared with the
Iophonidae-Microcionidae group (absence of sigmas), but no character is shared
between Tedaniidae-Crambeidae and the "myxilline" group. Their position remains
problematical.
BIOGEOGRAPHY
The biogeography of Chilean sponges was treated extensively in previous
papers by one of us (DESQUEYROUX-FAUNDEZ & MoyanNO, 1989, DESQUEYROUX-
FAUNDEZ, 1994), as well as by older authors (e.g. EKMAN, 1953; BRIGGS, 1974). A
study of Magellanic-Antarctic sponge relationships was performed by SARÀ (1992).
From these studies two remaining questions are apparent:
- the existence of more than a single area along the coast of Chile
- the relationship and origin of the Chilean sponge fauna
Below, we will discuss to what extent our monographic treatments of the three
genera contribute to solving these questions.
The genus /ophon has a predominantly coldwater distribution, with only few
species recorded from the tropical or subtropical regions (VAN SOEST, 1994). In our
study it is reconfirmed that /ophon lamella and I. ostiamagna are genuine endemics of
the tropical-subtropical East Pacific. The other species reported here conform to the
general cold water distribution by being confined to the SE Pacific and (in the case of
I.proximum) to the area on both sides of South America. Endemism in this genus
seems to indicate the existence of two areas along the coasts of Chile, viz. 20°-42°S
(lophon proximum) and 43°-53°S (I.timidum and I.chilense). Since so many more
species of Jophon are found in the Southern Ocean and along the coasts of Antarctica,
and relatively few in the Arctic and boreal waters, it is likely that the genus originated
in the seas surrounding the southern continents.
The genus Myxilla is cosmopolitan but also has a predominance in cold water
(VAN SOEST, 1994). Since this is is a very large genus, with almost certainly some
distinct subgeneric groups, a further analysis will have to demonstrate whether all the
records from tropical regions are indeed Myxilla. We can here record at least one
species, M. (Myxilla) mexicensis, from the Galapagos Islands, with characters similar
to that of the type species. Another species from the Galapagos is M.(Stelodoryx)
cribrigera. Two further tropical records are M.(Myxilla) asymmetrica n. sp. and
Myxilla (Ectyomyxilla) dracula n. sp. from Peru, but there were collected in deep
water (200-600 m). The Myxilla distributions do not support a subdivision of a
northern and a southern Chilean area of endemism separated at about 42°S, most
species being confined to the tropics or to both sides of southern South America. One
species, M. (Stelodoryx) cribrigera occurs over the whole study area. The subgeneric
74 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
units of Myxilla used here are not likely to represent monophyletic groupings. Thus
their distributions cannot provide us with firm clues as to their origin. However,
Myxilla species in the study area (and also in neighbouring areas of the Southern
Ocean), with the exception of M. (M.) mexicensis have "loose" skeletons with a
tendency to form ill-defined plumoreticulate architecture. Styles often have only a
few spines. These characters may indicate supraspecific endemism in the southern
oceans. The Myxilla species of the Arctic-boreal regions are not well-studied, but an
obvious close relationship with southern ocean Myxilla is not apparent.
The genus Tedania is considered to have originated in the Antarctic-Antiboreal
regions (VAN SOEST, 1994), because most species have been reported from these parts
of the world. As indicated above, the tropical and subtropical regions contain Tedania
species with very similar characteristics, which by some authors are considered
members of a single cosmopolitan species (T. anhelans), or a complex of closely
related sibling species. From the Galapagos we describe here also a member of this
complex, Tedania (Tedania) galapagensis, while in a previous study of sponges of
Isla de Pascua (DESQUEYROUX-FAUNDEZ, 1990) a further species from the study area
was reported. The other Tedania species from the area belong to clearly different
subgeneric units, and their distributions are limited to both sides of southern South
America. Like the Myxilla distributions they confirm the existence of a Magellan area
on both sides of South America (EKMAN, 1953, BRIGGS, 1974), but do not present
evidence for a northern Chilean area of endemism. Some of the Arctic-boreal Tedania
belong to the subgeneric units occurring in the study area, but their number is limited.
In summary: endemism in the tropical region including the Galapagos fauna
and tropical continental faunas is clearly present in all three genera. The Chile-Peru
and Magellan regions are not clearly separated in the three genera, although /ophon
shows some evidence of the existence of these two separate areas. The close relation-
ship with faunas of Antarctica and other Southern Ocean regions is clear for all three.
The Arctic-boreal regions also show close relationships with the study area, but most
(sub) generic units appear poorer in species number. This perhaps indicates a southern
origin.
EKMAN (1953) and BRIGGS (1974) maintain that the Chilean fauna despite is
high degree of endemism is closely similar to that of the Pacific coast of North
America, with many species the same in both regions. VERMEU (1991) postulates a
Transequatorial interchange of north and south East Pacific faunas during the
Pliocene, following the gradual closure of the Panama seaway. This interchange was
mostly from north to south, although the subsequent Pleistocene interchange between
the regions supposedly was bothways. Before the Pliocene the tropical region was too
strong and currents too adverse for faunal interchange. We find little evidence to
support that 1) much interchange has ocurred, and 2) that the main stream was north-
south. This may have several causes, the most important probably being lack of
knowledge of the northern North Pacific fauna. Since the days of Lambe at the end of
last century no serious systematic work on sponges from the northern part of the area
has been done. Still, the large numbers of Tedania and Jophon in the south Pacific as
compared to those of the north are hardly likely to be an artifact. Also the morpho-
S-E PACIFIC SPECIES OF POECILOSCLERIDA 75
logical distinctness of Southern Ocean Tedania indicates that endemism in the south
is of a high taxonomic level and likely to be at least of Tertiary age. The few Arctic-
boreal Tedania may have invaded the north during Pleistocene times (following
VERMEIJ). Pliocene northern immigrants in the study area are not apparent.
Relationships with Antarctica are distinct in all three genera. Both in Jophon
and Tedania there are species described from the study area with peculiar morpho-
logical traits shared with Antarctic species (lophon chilense and I. unicornis, Tedania
charcoti, T. spinata. This indicates fairly recent interchange and subsequent spe-
ciation. During the Pleistocene, Antarctica and southern Chile were covered by a
thick icecap, possibly destroying the littoral fauna to considerable depths. Subsequent
reinvasion of these areas were inevitably from the north (VAN OPPEN et al., 1994), and
possibly the distribution of Tedania charcoti in Chile and Antarctica is a product of
that reinvasion.
ACKNOWLEDGMENTS
The following persons contributed to our study.
C. Valentine (BMNH); D. Kühlmann (ZMB); M. Dzwillo (ZMH); C.Lévi (MNHN);
C. Rützler, K. Smith and P. Norenburg (USNM); V. Raineri (MCSN: ); M. Borri
(MZSF); G. Bakus (usc); O. Tendal (ZMK); J. Fournier (CNM); J. Vermeulen (ZMA); C.
Osorio, A. San Martin (UCH); F. Jara, R. Schlatter (UACH); N. Bahamonde, L. Capurro,
A. Cornejo (MNHNC); J. Artigas, A. Gallardo, H. I. Moyano (MZUC) J. Redon (IOUV); J.
Wiiest; C. Ratton; I. Juriens (MHNG).
Reviewer greately contributed to our work.
76 RUTH DESQUEYROUX-FAUNDEZ & ROB W. M. VAN SOEST
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REVUE SUISSE DE ZOOLOGIE, 103 (1) : 81-100; mars 1996
Taille et mélanisme chez Vipera aspis dans les Préalpes suisses
et en Italie centrale et comparaison avec différentes
populations alpines de Vipera berus
Jean-Claude MONNEY*, Luca LUISELLI** & Massimo CAPULA**
*Université de Neuchatel, Institut de Zoologie, Rue Emile-Argand 11,
2007 Neuchatel, Suisse
**Universita di Roma “La Sapienza”, Dipartimento di Biologia Animale
e dell Uomo, Via Borelli 50, 00161 Roma, Italia
Body size and melanism in Vipera aspis in the Swiss Prealps and
Central Italy and comparison with different alpine populations of
Vipera berus. - In a contact zone of Vipera aspis and Vipera berus in the
Swiss Bernese Prealps, V.berus is smaller than V.aspis. V.berus seems to be
a dwarf mountainous form. The rate of melanism is higher in V.berus
(49%) than in V.aspis (34%), probably a consequence of the difference in
habitat use between the two species. Comparisons with other Swiss and
Italian populations of both V.aspis and V.berus suggest that melanism is
particulary advantageous in closed, forested areas. On the other hand, for
both species and in all the studied populations, the rate of melanism is
higher in females than in males, suggesting a higher advantage to be black
for the females. In Switzerland, as well as in Italy, melanistic V.berus tend
to be heavier and longer than cryptic ones, whereas in V.aspis the opposite
is the case. In both species, growth rates are higher in melanistic morphs
and we suggest that the larger body size of the cryptic V.aspis is related to
their higher survival rate and higher mean age.
Key-words: Vipera berus - Vipera aspis - Body size - Melanism - Ecology
INTRODUCTION
Le mélanisme chez Vipera aspis, fréquent localement et particulièrement en
montagne, a souvent été rapporté (FATIO 1872; SAINT GIRONS & FONS 1977; ARNOLD
& BURTON 1978, PILLET & GARD 1979; HOFER 1991; KRAMER & STEMMLER 1992)
mais il n’a fait l’objet que de rares études (NAULLEAU 1973; MONNEY 1990).
Manuscrit accepté le 29.05.1995.
82 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
Exceptionnel, voire inexistant dans la Chaîne jurassienne, il en est de même de
manière générale, au Nord de l’aire de répartition de l’espèce. Dans plusieurs régions
des Alpes et des Préalpes au contraire, ce caractère est fréquent et le taux de
mélanisme au sein d’une population est souvent voisin de 50%. Dans les Préalpes
fribourgeoises par exemple, à l'Ouest de la Suisse, NAULLEAU (1973) signale 54% de
vipères noires (n=79) et MONNEY (1990) 58% (n=21). Il en est de même dans les
Apennins, au centre et à l’Est de l’Italie, où le mélanisme est localement fréquent au
sein des populations montagnardes de V.aspis (BRUNO 1990).
La situation est bien différente pour V.berus. Le mélanisme chez cette espèce
n’est pas rare dans le Jura et au Nord de son aire de répartion, ainsi qu’en montagne.
Les hypothèses tentant d'expliquer le rôle écologique du mélanisme chez V.berus et
sa maintenance au sein d’une population sont nombreuses, mais bien des questions
restent encore ouvertes (ANDREN & NILSON 1981; FORSMAN & As 1987; MADSEN &
STILLE 1988; LUISELLI 1992, 1993; CAPULA & LUISELLI 1994; LUISELLI et al. 1994;
MONNEY et al. 1995). Afin de mieux cerner les facteurs impliqués dans ce polymor-
phisme, et comme le suggère LUISELLI (1993), il nous a semblé judicieux de comparer
la situation chez deux espèces proches philogénétiquement et vivant dans une même
station. Occupant la même niche écologique (SAINT GIRONS 1975), ces deux serpents
presentent-ils des caractéristiques morphologiques communes liées au mélanisme ?
Retrouve-t-on les mêmes tendances dans d’autres populations ? Cet article est la
première contribution détaillée qui tente de répondre à ces questions. D’autres publi-
cations par les mêmes auteurs aborderont le problème par le biais d’un modèle
théorique.
MATÉRIEL ET MÉTHODES
Nous comparons la taille et le mélanisme des vipères aspics provenant essen-
tellement de trois régions différentes: deux stations préalpines situées à l’Ouest de la
Suisse, l’Oberland bernois et les Préalpes fribourgeoises, et une station d'Italie cen-
trale (Abruzzes). En ce qui concerne la vipère péliade, les données proviennent de la
station de l’Oberland bernois qui abrite les deux espèces ainsi que d’autres stations
alpines (MONKEY er al. 1995). Les données climatiques sont tirées, pour les stations
suisses, de l’Atlas hydrologique de la Suisse (Kirchhofer et Sevruk, 1992) et de
l'Atlas de la Suisse (Imhof, 1965).
1. STATION DE L’OBERLAND BERNOIS
L’essentiel de nos données a été récolté sur une surface d’environ 40 ha située
à une altitude s’échelonnant de 1400 à 1600 m, dans l’étage subalpin inférieur. Cette
surface correspond à une zone de contact relativement étroite entre V.aspis et V.berus
qui fréquentent les mêmes types d’habitat mais dans des proportions différentes et
sans véritable cohabitation. Il s’agit d’un versant exposé au Sud-Est, comprenant deux
cônes de déjection torrentiels colonisés par une mosaïque d'associations végétales, des
TAILLE ET MELANISME CHEZ VIPERA ASPIS 83
pâturages et des prairies humides parsemés localement de murgiers, et une forêt
d’épicéas. Une troisième espèce de serpent vit dans cette station; il s’agit de Coronella
austriaca. La température annuelle moyenne est inférieure à 5°C et la hauteur annuelle
moyenne des précipitations est supérieure à 1600 mm.
De 1987 à 1993, nous avons marqué individuellement les vipères par coupe
d’écailles sous-caudales (BLANCHARD & FINSTER 1933). Les serpents ont été pesés a
l’aide d’un dynamomètre de type “PESOLA”. Pour mesurer leur longueur totale (LT)
et leur longueur museau-cloaque (LC), une fourchette modifiée (pointe médiane
coupée à sa base et remplacée par de la mousse) a été utilisée pour maintenir la tête de
la vipère plaquée au sol. Toutes ces manipulations ont été faites sur le terrain, le
serpent étant immédiatement relâché après ces mesures. Seule la taille des femelles
reproductrices et des mâles adultes est considérée. La longueur totale minimale de ces
derniers est celle des plus petits mâles observés en activité sexuelle, soit 40 cm pour
V.berus et 45 cm pour V.aspis. Le poids des vipères n’est relevé que si nous n’obser-
vons aucune trace d’activité trophique récente, soit essentiellement en fin de gestation
pour les femelles et au début des périodes d’accouplement pour les mâles. Si nous
n’avons considéré que les femelles reproductrices, c’est que le cycle sexuel des
femelles des deux espèces est pluriannuel, au plus triennal pour V.aspis et au plus
biennal pour V.berus (MONNEY 1995); de ce fait, les variations pondérales annuelles
chez ces vipères sont importantes et difficilement comparables. La récolte de nos
données s’échelonnant sur une période de 7 années, un même individu peut être
comptabilisé plusieurs fois, mais au plus une seule fois par année. L'état sexuel des
femelles a été déterminé par palpation des follicules ovariens ou des embryons. Les
cycles annuels d’activité de ces deux espèces, leur régime alimentaire ainsi que les
données sur le mélanisme de V.berus ont fait l’objet d’autres publications (MONNEY
1993, 1995; MONNEY et al. 1995).
2. STATION DES PRÉALPES FRIBOURGEOISES
Cette station est située à une altitude moyenne de 900 m et a été prospectée
intensivement en 1986 (MONNEY 1990, 1992). Elle est située dans l’étage montagnard
inférieur dominé par le hêtre (Carici- et Cardamino-Fagetum) et comprend, outre la
forêt, des pâturages parsemés de murgiers et de murs de pierres sèches, et de
nombreuses haies buissonnantes. En plus de la vipère aspic, dont les femelles se
reproduisent en moyenne tous les deux ans, on y trouve la coronelle lisse (Coronella
austriaca) et la couleuvre à collier (Natrix natrix). La température annuelle moyenne
est d’environ 8°C et la hauteur annuelle moyenne des précipitations est d’environ
1500 mm. Les données ont été récoltées de la même manière que dans l’Oberland
bernois.
3. STATION DE L'ITALIE CENTRALE
En 1993 et 1994, une troisième série de données concernant une population
mélanique de V.aspis a été récoltée suivant les mêmes méthodes en Italie centrale, à
une altitude de 800 m (Val Sagittario, Abruzzes). Le cycle sexuel des femelles est en
84 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
moyenne biennal. Pour cette station, le poids des femelles non reproductrices a
également été considéré. Ces vipères vivent en sympatrie avec quatre autres espèces
de serpents. Il s’agit de Coronella austriaca, de Natrix natrix, d’Elaphe longissima et
de Coluber viridiflavus, cette dernière étant fréquemment mélanique. La végétation
appartient à la ceinture samnitique de la région méditerranéenne (forêt de feuillus
dominée par le hêtre). La température annuelle moyenne est d’environ 10°C et la
hauteur annuelle moyenne des précipitations est d’environ 1150 mm. Les vipères sont
abondantes en bordure de la forêt mixte, et notamment à proximité des vieux murs de
pierres et des murgiers.
Les longueurs et poids moyens des vipères sont comparés à l’aide du test t de
Student. Le test du X? est utilisé pour comparer les taux de mélanisme. Les droites de
régression sont comparées selon la méthode de HALD (1967).
RÉSULTATS
1. COMPARAISON INTERSPÉCIFIQUE
Dans la station de l’Oberland bernois occupée par les deux espèces de vipères,
les adultes de V.aspis, mâles et femelles, sont significativement plus longs (LT et LC)
et plus lourds que les adultes de V.berus (poids des mâles: d.f.=217, t=8.283; poids
des femelles: d.f.=136, t=4.054; LT mâles: d.f.=237, t=13.250; LT femelles: d.f.=136,
t=6.436, p<0.0001 dans tous les cas). La différence est particulièrement marquée chez
les mâles. Les aspics mesurent environ 10 cm de plus et pèsent en moyenne 25 g de
plus (Tab. 1 et Fig. 1). La comparaison des droites de régression du logarithme de la
longueur du corps par le logarithme du poids montre que les mâles péliades sont
significativement plus trapus que les mâles aspics (Fig. 2). Les variances résiduelles
des deux droites ne sont pas significativement différentes (p=0.130), leurs pentes
valent 2.55 et 3.02 et sont significativement différente, p=0.018) (Fig. 2). Il n’y a par
contre pas de différence significative entre les femelles des deux espèces.
Le pourcentage de vipères noires est significativement plus grand chez V.berus
(49%, n=74) que chez V.aspis (34%, n=173) (X2=4.632, d.f.=1, p=0.031). Dans les
stations fribourgeoise et italienne au contraire, les taux de mélanisme chez V.aspis,
resp. 58% (n=36) et 56% (n=64) ne sont pas statistiquement différents de celui
observé chez V.berus dans l’Oberland (d.f.=1, X2=0.910 et 0.795, p=0.340 et 0.372
resp.), alors qu'ils diffèrent significativement de celui de V.aspis de l’Oberland
(d.f.=1, X2=7.405 et 9.540, p=0.006 et 0.002 resp.)
2. COMPARAISON INTRASPÉCIFIQUE
a. Vipera aspis
Dans l’Oberland bernois, la longueur totale des mâles et des femelles n’est pas
significativement différente (t=-0.332, d.f.=260, p=0.740) (Tab. 1). La longueur du
corps et le poids des femelles sont significativement supérieurs à ceux des mâles (t=-
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TAILLE ET MELANISME CHEZ VIPERA ASPIS
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86 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
140
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Poids (g)
80
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V.aspis, M V.aspis, F V.berus, M V.berus, F
FIG. |
Moyennes et écarts types du poids des vipères aspics et péliades dans une même station de
l’Oberland bernois. Mâles (M) et femelles (F) de V.aspis sont significativement plus lourds que
V.berus. On remarque d’autre part que les péliades mélaniques sont plus lourdes que les
normales et que c’est l’inverse chez V.aspis femelle.
2.893, d.f.=247, p=0.0042; t=-8.020, d.f.=241, p<0.0001 resp.). Les femelles “nor-
males” sont en moyenne significativement plus lourdes que les mélaniques, resp.
107.6 g et 94.1 g (t=-2.385, d.f.=85, p=0.0193). Leurs longueurs moyennes ne sont
par contre pas significativement differentes (pour LT et LC, t=1.6821, d.f.=85,
p=0.0961; t=1.654, d.f.=80, p=0.1020). Chez les mâles, les longueurs et poids moyens
ne different pas entre les deux phénotypes. Par contre, la comparaison des droites de
régression (logarithme du poids par logarithme de la longueur du corps) montre que
les mâles normaux sont plus lourds, pour une longueur donnée, que les mélaniques, la
distance entre les deux droites étant significativement différente de 0 (p=0.004). Elle
mesure sur Ÿ (X fixe) 0.0561 (Fig.3). Le taux de mélanisme des mâles (n=79) et des
femelles (n=94), respectivement 30 et 37%, n’est pas sigificativement différent
(X2=0.897, d.f.=1, p=0.343).
La taille respective de V.aspis des Préalpes fribourgeoises et de l’Italie centrale
ne diffère pas, pour les mâles et les femelles, de façon significative (pour LT: t=-
1.491, d.f.=46, p=0.142 et t=1.499, d.f.=49, p=0.140 resp.) (Tab. 2). Ces vipères,
males et femelles, sont par contre significativement plus grandes que celles de
l’Oberland (pour LT de la station fribourgeoise: t=-3.566, d.f.=184, p=0.0005 et t=-
2.643, d.f.=109, p=0.0094 resp. et pour LT de la station italienne: t=-2.184, d.f.=210,
p= 0.0301 et t=-5.030, d.f.=112, p<0.0001 resp.) (Fig. 4). Dans ces deux stations, le
taux de mélanisme ne diffère pas non plus de façon significative entre mâles et
TAILLE ET MÉLANISME CHEZ VIPERA ASPIS 87
5 + MVaspis
A M V.berus
4.6
u
Je
O
0.
= 4.2
=
38
3.4
36 i 5 5 4 4.1 4.2 4.3
Log Longueur totale
Log Poids = -7.674 + 2.967 * Log Longueur totale; R12 = .829 (M V.aspis)
Log Poids = -6.206 + 2.626 * Log Longueur totale; R*2 = .801 (M V.berus)
Fic. 2
Droites de régression du logarithme de la longueur en fonction du logarithme du poids de
Vipera aspis Male (M) et de V.berus Male. Les pentes des deux droites sont significativement
différentes (p=0.018).
hemellesi@x?—0:08/, Ad El p=0768 et X2=0.855; dif =i, p=0:355 resp) Ileest
cependant intéressant de constater que pour les trois populations, on observe la méme
tendance, soit un taux de mélanisme chez les femelles supérieur a celui des males,
resp. 37 et 30% dans l’Oberland, 60 et 54% dans les préalpes fribourgeoises, et 63 et
51% dans les Abruzzes. Cette méme tendance générale avait déja été relevée chez
V.berus dans les Alpes et dans une station de plaine (Monney et al., 1995).
Comme dans l’Oberland, la longueur totale des femelles dans les Abruzzes et
les Préalpes fribourgeoises ne diffère pas significativement de celle des mâles
(t=1.912, d.f.=62, p=0.0604 et t=-1.268, d.f.=33, p=0.2138, resp.). Il en est de même
pour le poids, les femelles étant significativement plus lourdes que les males (t=3.934,
d.f.=62, p=0.0002) et t=2.417, d.f.=33, p=0.0213 resp.).
88 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
O Maspis norm.
m Maspismél.
Log Poids
4
Log Longueur corps
Log Poids = -7.051 + 2.928 * Log Longueur corps; R"2 = .883 (M aspis norm.)
Log Poids = -8.386 + 3.258 * Log Longueur corps; R12 = .848 (M aspis mel.)
FIG. 3
Droites de régression du logarithme de la longueur en fonction du logarithme du poids des
mâles (M) mélaniques (mél.) et des mâles normaux (norm.) chez Vipera aspis de l’Oberland
bernois. La distance entre les deux droites est significativement différente de 0 (p=0.004).
La comparaison des longueurs et des poids moyens entre les deux phénotypes
mélanique et normal met en évidence la même tendance dans les Abruzzes que dans
l’Oberland. Les femelles normales ont une longueur moyenne identique à celle des
mélaniques, resp. 62.52 cm (n=10) et 62.68 cm, (n=17) (t=0.074, d.f.=25, p=0.941) et
tendent à être plus lourdes que celles-ci (resp. 143.70g et 132.41g), la différence
n'étant cependant pas significative (t=-0.853, d.f.=25, p=0.402). Les longueurs et
poids moyens des mâles ne diffèrent pas significativement entre les deux phénotypes.
Par contre, la comparaison des droites de régression (logarithme du poids par loga-
rithme de la longueur du corps) montre que, comme dans l’Oberland bernois, les mâles
normaux sont plus lourds, pour une longueur donnée, que les mélaniques (Fig. 5). La
distance entre les droites est significativement différente de 0 (p=0.025) et mesure sur
Y (X fixe) 0.0561. Les variances résiduelles des deux droites ne sont pas
TAILLE ET MÉLANISME CHEZ VIPERA ASPIS 89
TABLEAU 2
Longueurs et poids de Vipera aspis en Italie centrale (Abruzzes) et dans les Préalpes fribour-
geoises. Les étoiles indiquent les couples de valeurs qui diffèrent significativement (p<0.05).
Poids (g)
Longueur totale (cm)
N Moyenne Ecarttype Min.-Max. N Moyenne Ecart type Min.-Max.
V.aspis
Abruzzes
Mäles 371 SI 7.913 48.3-74.2 57 10222 35.643 53.0-189.0
Femelles DAI 62.6 5.414 53.2-72.1 27 136.6* 33.049 92.0-197.0
Mäles norm. 18 60.4 8.364 48.3-73.3 18 11572 43.521 53.0-189.0
Mäles mel. 19 5828075240 50.2742 19 hed 24.442 54.0-142.0
Fem. norm. 10226257776:.031°753.2-724 10 143.7 33.308 96.0-197.0
Fem. mel. 17 0272521907 539-7281 17 132.4 32.894 92.0-189.0
V.aspis
Prealpes frib.
Mäles Hl O WB) 48.8-73.0 11 122.2* 36.499 65.0-168.0
Femelles 24 60.2 6.086 49.0-70.0 8 167.9* 21.184 140.0-192.0
Mälesnorm. 5 61.1 9.619 48.8-73.0 5 118.7 46.154 65.0-168.0
Mäles mel. 6 69,2.26524°53:.0-72:0 6 1254 30.616 76.0-160.0
Fem. norm. 9 60.9 5.742 51.0-68.0 166.0 140.0-192.0
Fem. mél. 15 59.7 6.439 49.0-70.0 168.6 18.864 140.0-184.0
O N
significativement différentes (p=0.024), de même que leurs pentes qui valent 2.43 et
1.89 (p=0.107). Nous obtenons les mêmes résultats pour les femelles, relevant l’avan-
tage pondéral, pour une longueur donnée, des femelles normales (Fig.6) (Distance
entre les deux droites significativement différente de 0 (p=0.023) et mesurant sur Y
(X fixe) 0.0961, pentes non différentes et valant 2.362 et 2.628 ( p=0.561), variance
résiduelle non significativement différente (p=0.028) ).
En ce qui concerne la station fribourgeoise, le petit nombre de mâle ne permet
pas de comparaison statistique entre les phénotypes et il n’y a pas de différence
significative de longueur au sein des femelles.
b. Vipera berus
Les données concernant cette espèce sont présentées de façon plus détaillée
dans une autre publication (MONNEY et al. 1995) et nous ne ferons que les rappeler
brièvement pour la comparaison avec V.aspis vivant dans cette même station de
l’Oberland bernois. Les femelles de V.berus sont significativement plus longues et
plus lourdes que les mâles (t=-7.708, d.f.=113; t=-11.858; d.f.=112 resp., p<0.0001),
la différence étant beaucoup plus marquée que chez V.aspis. Contrairement à ce que
90 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
A Males aspics
© Femelles aspics
Longueur totale (cm)
Oberland Italie P.frib.
FIG. 4
Moyennes et écarts types de la longueur des vipères aspics mâles et femelles dans l’Oberland
bernois, dans les Abruzzes (Italie centrale) et dans les Préalpes fribourgeoises. Les vipères
aspics de l’Oberland, mâles et femelles, sont significativement plus petites que dans les deux
autres stations.
l’on observe chez V.aspis, ce ne sont pas les femelles normales mais les mélaniques
qui sont significativement plus lourdes (t=-2.209, d.f.=49, p=0.032). Comme pour les
femelles de V.aspis, il n’y a pas de différence de longueur entre les deux phénotypes
(pour LC, t=-1.353, d.f.=47, p=0.1826; pour LT, t=-1.739, d.f.=49, p=0.0884). En ce
qui concerne les mâles, la situation est également différente entre les deux espèces.
S’il n’y a pas de différence significative de poids moyen et de longueur moyenne chez
V.aspis, les mâles mélaniques de V.berus sont quant à eux significativement plus
longs et plus lourds que les normaux (pour LC: t=-2.185, d.f.=62, p=0.0327; pour LT,
t=-2.216, d.f.=62, p=0.0304; pour PDS: t=-2.159, d.f.=61, p=0.0348). Les droites de
régression entre les phénotypes (logarithme du poids par logarithme de la longueur du
corps) ne sont par contre pas significativement différentes, que ce soit leurs pentes
(p=0.608) ou la distance entre les droites (p=0.593).
Comme pour V.aspis, le taux de mélanisme des mâles (N=27) et des femelles
(n=47), respectivement 44 et 51%, n’est pas sigificativement différent (X2=0.301,
d.f.=1, p=0.5834).
TAILLE ET MELANISME CHEZ VIPERA ASPIS 9]
Log Poids
3.8 3.9 4 4.1 4.2 4.3 4.4
Log Longueur totale
Log Poids = -3.208 + 1.897 * Log Longueur totale; R/2 = .87 (mél.)
Log Poids = -5.286 + 2.432 * Log Longueur totale; R12 = .841 (norm.)
FIG. 5
Droites de régression du logarithme de la longueur en fonction du logarithme du poids des
mâles mélaniques (mél.) et des mâles normaux (norm.) chez Vipera aspis des Abruzzes (Italie
centrale). La distance entre les deux droites est significativement différente de 0 (p=0.025).
3. LE MÉLANISME CHEZ LES JEUNES VIPÈRES ET CHEZ LES ADULTES
A la naissance et dans toutes les stations étudiées, les vipéreaux des deux
espèces sont tous normalement colorés, ce qui confirme les observations d’autres
auteurs (NAULLEAU 1972, 1973, 1979; MONNEY 1990). Chez V.berus, il est souvent
possible de différencier les nouveau-nés qui deviendront noirs à leur couleur de fond
légèrement plus foncée que celle des autres, ce qui n’est pas le cas chez V.aspis. Chez
les deux espèces, le mélanisme apparaît progressivement au cours de la croissance,
par assombrissement de la couleur de fond des téguments. Durant leur première
année, nous avons observé, dans l’Oberland bernois, 8 jeunes vipères aspics méla-
niques qui mesuraient entre 23 et 29 cm (moyenne=25.9 cm). Toutes étaient très
foncées. Le dessin dorsal était encore visible et les yeux ainsi que les écailles labiales
92 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
Log Poids
4 4.1 4.2 4.3
Log Longueur totale
Log Poids = -4.818 + 2.363 * Log Longueur totale; R42 = .942 (norm.)
Log Poids = -6.008 + 2.628 * Log Longueur totale; R42 = .798 (mel.)
FIG. 6
Droites de régression du logarithme de la longueur en fonction du logarithme du poids des
femelles mélaniques (mél.) et des femelles normales (norm.) chez Vipera aspis des Abruzzes
(Italie centrale). La distance entre les deux droites est significativement différente de 0 (p=0.023).
étaient clairs. S’il est progressif, on peut relever cependant que l’assombrissement est
très marqué immédiatement après la mue. Une femelle de 29 cm par exemple
paraissait encore relativement claire le 19 juillet 1991. Suite à sa mue le 4 septembre
de la même année, elle devint presque entièrement noire mis à part ses yeux et ses
écailles labiales. Au cours de leur deuxième année, les vipères aspics deviennent
totalement noires, y compris les écailles labiales et l’iris de l’oeil. Certains individus,
essentiellement des femelles, demeurent rougeätres, coloration pouvant persister
jusqu’à l’âge adulte. Ce type de mélanisme est fréquent dans les Préalpes fribour-
geoises. Six jeunes vipères aspics de l’Oberland bernois, complètement noires, dans
leur deuxième année, mesuraient entre 29.5 et 35.5 cm (moyenne= 32.3 cm).
Si le mélanisme semble légèrement plus accentué et se manifeste un peu plus
rapidement chez les mâles que chez les femelles de vipères aspics, la différence est
TAILLE ET MELANISME CHEZ VIPERA ASPIS 93
encore plus nette chez la vipère péliade. Nous avons observé un mâle d’une année
exactement, complètement noir à l’exception de l’iris brun de ses yeux, mesurant 21
cm. Trois jeunes femelles mélaniques dans leur deuxième année, mesurant respec-
tivement 22, 23 et 27 cm, ainsi que deux autres dans leur troisième année, mesurant
30 et 33.5 cm, présentaient toutes une robe encore brun-foncée. Certaines femelles
mélaniques passent par un stade concolor brun-rouge cuivré lorsqu'elles mesurent
entre 40 et 45 cm. Cette coloration particulière a déjà été décrite par PIELOWSKI (1962)
et SCHIEMENZ (1987) et correspond probablement, dans certaine région, au terme
populaire de “vipere cuivrée”, “Kupferotter” en allemand.
Chez les vipères adultes, les différences interspécifiques liées au mélanisme
sont évidentes dans les stations suisses étudiées. La plus remarquable est la couleur de
l’iris qui est, à de rares exceptions près, complètement noire chez V.aspis. Dans
l’Oberland bernois, moins d’un pourcent des vipères aspics noires avait l'iris
rougeâtre ou blanchâtre. Au contraire, l’iris de l’oeil de V.berus est toujours bien
orangé chez les femelles et brun-foncé rougeätre chez les mâles. Chez la plupart des
femelles de cette espèce, les écailles labiales sont blanches ou jaunätres, et le corps est
parsemé de minuscules taches claires, souvent jaunätres. Ces deux dernières carac-
téristiques sont tout à fait exceptionnelles chez les mâles de V.berus et nous ne les
avons observées qu’à une seule reprise chez une femelle de V.aspis.
Dans les Abruzzes, les vipères aspics mélaniques, mâles et femelles, ont l’iris
bien rougeätre et souvent les écailles labiales claires, comme les femelles de V.berus
dans l’Oberland.
DISCUSSION
La comparaison de la taille des vipères adultes entre notre station de
l’Oberland bernois et une autre zone de contact, mais de plaine celle-là, en Loire-
Atlantique (SAINT GIRONS & NAULLEAU 1981), est intéressante. En effet, s’il n’y a pas
de différence significative de longueur et de poids pour V.aspis, V.berus est signifi-
cativement plus petite dans notre station. Les dimensions que nous observons chez
cette espèce correspondent à celles mentionnées par NEUMEYER (1984) dans les
Grisons. Il semble s’agir d’une forme naine de montagne (MONNEY et al. 1995),
beaucoup plus petite que les formes nordiques. Il est curieux de constater que dans les
Alpes orientales italiennes, à une altitude de 1100 mètres, les vipères péliades sont
également beaucoup plus grandes, les mâles excédant très rarement 63 cm et les
femelles 72 cm (CAPULA er al. 1992; MONNEY et al. 1995). Il s’agit là de tailles plutôt
exceptionelles, les péliades mâles des Alpes carniques mesurant pour la plupart entre
50 et 55 cm, les femelles entre 55 et 62 cm. Dans l’Oberland, les longueurs maxi-
males que nous avons enregistrées sont respectivement 55 et 58 cm pour les mâles et
les femelles. La plus petite taille des vipères aspics de l’Oberland bernois, comparée à
celle de l’Italie centrale et des Préalpes fribourgeoises, est peut-être due aux con-
ditions climatiques moins favorables de cette région, et notamment à l’enneigement
qui est beaucoup plus important. La durée de la période active s’en trouve raccourcie,
et par là même la croissance des vipères est ralentie. Le type de végétation reflète
94 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
parfaitement les différences de climat, les forêts de hêtres de nos stations des
Abruzzes et des Préalpes fribourgoises contrastant avec les forêts d’épicéas de
l’Oberland bernois. Il se pourrait également que des facteurs alimentaires jouent un
rôle. Dans notre station des Préalpes fribourgeoises par exemple, le lézard vivipare est
absent et le lézard agile plutôt rare. Dans ces conditions, les vipères adultes n’en
consomment pas ou exceptionnellement (MONNEY 1990). Ce n’est pas le cas dans
l’Oberland bernois où CAVIN (1993) a dénombré localement plus de 400 lézards
vivipares à l’hectare, ce qui contribue à en faire une proie habituelle des vipères
aspics, jeunes et adultes (MONNEY 1993). Cette abondance de petites proies pourrait
favoriser les vipères de petite taille. Il est également à relever qu’une taille réduite
peut être avantageuse pour la thermorégulation en zone de montagne.
Plusieurs auteurs ont tenté d'expliquer le taux de mélanisme élevé, souvent
voisin de 50%, au sein de populations de V.berus aussi bien alpines que de plaine. En
effet, si comme cela a été démontré par BLANCHARD & BLANCHARD (1940) chez
Thamnophis s.sirtalis le mélanisme chez les vipères est un caractère récessif ne
s'exprimant que chez les homozygotes, une forte proportion d’individus noirs dans
une population indique que cette coloration présente des avantages adaptatifs
importants qui permettent son maintien. Le principal avantage sélectif pour une vipère
d’être noire est lié à une thermorégulation plus efficace, avec toutes les conséquences
métaboliques que cela peut entraîner: croissance plus rapide, prolongation de la
période active, meilleure activité trophique (GIBSON & FALLS 1979; FORSMAN & Äs
1987). Cela pourrait donc également permettre aux serpents d’exploiter des milieux
plus frais et plus humides. Si Forsman et Äs ne trouvent pas de différence signi-
ficative entre les deux phénotypes, plusieurs auteurs ont pu montrer que les péliades
mélaniques sont plus grandes ou plus lourdes que les normales (ANDRÉN & NILSON
1981, 1983; MADSEN & STILLE 1988; LUISELLI 1992, 1993; CAPULA & LUISELLI 1994;
MONKEY et al. 1995). Andrén et Nilson pensent que le taux de mélanisme élevé au
sein d’une population résulte d’un équilibre entre les avantages que procure une
grande taille dans la reproduction (femelles plus fécondes et mâles plus puissants lors
des combats prénuptiaux) et la plus grande vulnérabilité des vipères mélaniques vis-à-
vis des prédateurs. Luiselli et ses collègues obtiennent des résultats similaires mais
pensent que le mélanisme présente des avantages sélectifs essentiellement chez les
femelles. En effet, la mortalité suite à la parturition semble inférieure chez les
femelles mélaniques (LUISELLI 1992) et ces femelles peuvent d’autre part se
reproduire plus régulièrement que les normales (CAPULA & LUISELLI, 1994). De plus,
ces auteurs constatent que le taux de mélanisme est significativement plus bas chez
les mâles que chez les femelles, respectivement 13% (n=46) et 35% n=37).
Nos résultats dans l’Oberland bernois vont dans le même sens que ceux
d’Andren & Nilson et de Luiselli et collaborateurs, les péliades mélaniques étant plus
lourdes et (ou) plus longues que les normales. Nous avons également mis en évidence
une fécondité supérieure chez les femelles mélaniques (MONNEY er al. 1995), mais
nous ne pouvons par contre pas évaluer de manière précise l’importance de la morta-
lité des femelles postparturientes. Quant au taux de mélanisme, il n’est pas signifi-
cativement différent entre mâles et femelles, mais la tendance semble être la même,
TAILLE ET MÉLANISME CHEZ VIPERA ASPIS 95
respectivement 44 et 51%. Nous retrouvons cette même tendance dans les trois
populations de vipères aspics étudiées, ce qui semble démontrer que cette coloration
est particulièrement avantageuse pour les femelles des deux espèces.
La niche écologique de V.aspis et V.berus dans les Alpes italiennes présente
d'importantes différences. Les types d’habitats utilisés ne sont pas les mêmes et le
régime alimentaire des jeunes est different (LUISELLI et al. 1995). Cela n’est pas le cas
dans la zone de contact de l’Oberland bernois où les deux espèces semblent occuper
une niche écologique similaire, comme c’est également le cas en Loire-Atlantique
(SAINT GIRONS 1975). Malgré cette similitude, V.aspis semble subir des pressions de
sélection très différentes de celles de V.berus. En effet, non seulement le taux de
mélanisme chez cette espèce est plus faible que chez V.berus, mais ce sont les indi-
vidus normalement colorés qui présentent un avantage pondéral. Le nombre d’em-
bryons et le poids des femelles étant positivement corrélés (NAULLEAU & SAINT
GIRONS 1981; MONNEY 1994), les femelles normales contribuent à contre-sélectionner
le mélanisme. On peut également soupçonner un taux de survie supérieur chez les
femelles normales postparturientes, comme cela a été démontré par Luiselli pour les
femelles mélaniques de V.berus. Si le maintien d’un taux de mélanisme élevé chez
V.berus dans notre station peut s'expliquer par un plus grand succès dans la
reproduction, il est plus difficile d'expliquer le maintien du mélanisme chez V.aspis.
Dans l’Oberland, V.aspis est à la limite de ses possibilités de survie et au delà de la
ligne de démarcation entre les deux espèces, elle ne pourrait vraisemblablement plus
se reproduire sans entrer en hivernage avec ses embryons dans les utérus (MONNEY
1995). Pourquoi donc le pourcentage de mélanisme, caractère qui semble pouvoir
augmenter les performances thermorégulatrices, est-il inférieur chez l’espèce la plus
sensible aux conditions thermiques ? MADSEN & STILLE (1988) suggèrent que la
disponibilité en proie du milieu peut avoir une influence sur le taux de mélanisme au
sein d’une population de vipères, une pénurie en proie ayant selon eux un impact
négatif plus important sur les grands individus. Cette hypothèse, qui contredit les
résultats d’ANDREN (1982), ne nous paraît guère plausible pour notre station. Nous
pensons que la pression des prédateurs, et donc indirectement les types d’habitats
occupés par les viperes, et dans une moindre mesure leurs conditions d'humidité, sont
des facteurs essentiels déterminant le taux de mélanisme au sein d’une population
porteuse de ce gene. Dans notre station de l’Oberland bernois, la ligne de démarcation
relativement étroite entre les deux espèces correspond au passage progressif d’un
milieu plutôt buissonnant et ouvert, vers un milieu plus fermé, constitué d’une forêt
fraîche, humide et clairièrée. La pression des prédateurs potentiels sur les vipères,
essentiellement les rapaces diurnes et les corvidés, bien représentés sur notre terrain,
est de toute évidence plus forte en milieu ouvert et non forestier, ce qui pourrait
contre-sélectionner le mélanisme. L’avantage pondéral des femelles aspics normales
ne serait pas dû à une croissance accélérée, plus rapide que chez les mélaniques, mais
bien au contraire à un taux de survie et à une moyenne d’äge plus élevée. Si l’on
compare la taille moyenne des plus petites femelles aspics reproductrices, soit la taille
des femelles lors de leur première reproduction, on constate que les normales sont
96 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
significativement plus petites que les mélaniques, leur longueur totale étant
respectivement 49.37+2.66 cm (n=8) et 51.50+0.80 cm (n=8) (t=2.157, d.f.=14,
p=0.049). Cette même tendance est encore plus marquée chez V.berus, les plus petites
femelles mesurant respectivement 46.81+1.53 cm (N=8) et 49.02+1.03 cm (n=8)
(t=3.379, d.f.=14, p=0.004). Si l’on admet que, pour une même espèce, l’âge de ces
femelles est identique, le taux de croissance semble plus rapide chez les individus
noirs des deux espèces. Si les mâles normaux de l’Oberland ne sont pas plus lourds en
moyenne que les mélaniques, ils sont significativement plus trapus. Cela pourrait
également refléter un âge plus avancé pour une longueur donnée, et donc un taux de
croissance inférieur à celui des mâles mélaniques, ou tout simplement une meilleure
condition physique que ces derniers. Il est intéressant de constater qu’en Italie
centrale, dans des conditions très différentes, la tendance semble être identique, mâles
et femelles de vipères aspics normalement colorés étant également plus trapus, ce qui
suggère un âge moyen plus avancé chez ces individus. Une croissance plus rapide des
vipères noires a déjà été décrite chez les mâles de V.berus (MADSEN & STILLE (1988)
et chez les femelles de cette espèce (LUISELLI et al. in prép.), alors que les résultats de
FORSMAN (1993) sembleraient contredire ces observations.
Si l’on compare les taux de mélanisme dans les trois stations étudiées avec
ceux observés dans d’autres populations, il en ressort clairement que le type d’habitat,
essentiellement la couverture forestière, joue un rôle primordial. En Italie centrale et
dans les Préalpes fribourgeoises, les vipères aspics sont primairement liées à des
hêtraies et les taux de mélanisme dépassent 50%. Il en est de même pour la station
préalpine étudiée par Naulleau, à l’étage subalpin dominé par l’épicéa. Au contraire,
plusieurs stations colonisées par la péliade dans les Préalpes fribourgeoises sont
complètement nues, dépourvues d’arbres voire de buissons, et le taux de mélanisme
est de l’ordre de 10% (9.5%, n=21 pour l’une d’entre elles, température moyenne et
hauteur des précipitations moyennes annuelles resp. 0-5°C et 2000-2400 mm). A
l’opposé, dans une vaste tourbière du Jura suisse (Température et hauteur des préci-
pitations moyennes annuelles resp. 0-5°C et 1600-2000 mm), milieu essentiellement
forestier, le taux de mélanisme chez la péliade est de 66% (n=18).
Dans la vallée de l’Oberland bernois que nous avons étudiée, nous connaissons
deux stations extrêmes situées au-dessus de la limite supérieure de la forêt, à plus de
1900 m d’altitude. Dans la première station, qui n’est colonisée que par V.aspis, 15%
des vipères sont mélaniques (n=13), et dans la seconde, qui n’abrite que V.berus, le
taux de mélanisme est de 17% (n=6). Bien que notre échantillon soit petit, il semble
indiquer une même tendance pour les deux espèces vivant dans un milieu ouvert.
A notre avis, et comme l’avait déjà relevé KJAERGAARD (1981) pour V.berus,
les populations de vipères aspics et péliades à forts pourcentages d’individus noirs
semblent être des populations primairement forestières qui sont liées à des forêts
plutôt fraîches et humides. L’étude très détaillée de Reinert (1984) sur l’occupation
spatiale de Crotalus horridus en Pennsylvanie semblerait à priori renforcer cette
hypothèse. L’auteur démontre une utilisation de l’habitat significativement différente
entre les deux phénotypes. Les crotales mélaniques “préfèrent “ la forêt claire de grands
TAILLE ET MELANISME CHEZ VIPERA ASPIS 97
arbres, avec du bois mort, les normalement colorés “préférant” la jeune forêt, avec une
importante couche de feuilles mortes. Nous ne pensons pas cependant qu’au sein d’une
même population de vipères, les individus noirs occupent préférentiellement les
habitats les plus fermés, les individus normaux les habitats les plus ouverts (LUISELLI et
al. 1994). La pression de sélection ne se manifeste pas sur le comportement individuel
des vipères, mais sur la population entière.
S'il est indéniable que les vipères mélaniques sont avantagées pour leur ther-
morégulation, nous n’avons cependant pas relevé, chez les femelles reproductrices, de
différences notables quant aux dates des parturitions entre les deux phénotypes. Cela
est très curieux, car la durée de la gestation est probablement le critère le plus sensible
d’une meilleure thermorégulation. Il est cependant possible que les différences liées à
l'emplacement (donc la durée de l’insolation), ou au comportement, soient suffi-
samment grandes pour masquer l’influence de la coloration (SAINT GIRONS comm.
pers.). En ce qui concerne la durée de la période active, les rythmes journaliers
d’activité ou les cas d’exposition à l’extérieur dans des conditions climatiques
extrêmes, l’influence de la coloration nous paraît négligeable. Ainsi, le mélanisme ne
semble pas avantager les femelles gravides en accélérant l’embryogénèse, alors qu'il
semble favoriser un taux de croissance accru avec tous les avantages qui y sont liés.
RÉSUMÉ ET CONCLUSION
La comparaison de la taille et du mélanisme chez V.aspis et V.berus provenant
de différentes régions nous amène aux conclusions suivantes:
Dans une zone de contact entre V.aspis et V.berus dans l’Oberland bernois,
V.berus est plus petite que V.aspis. Cela n’est pas le cas en Loire-Atlantique, à la
limite nord de l’aire de répartition de V.aspis.
V.berus de l’Oberland bernois semble être une forme naine de montagne. Sa
taille moyenne et sa taille maximale sont nettement inférieures à celles de V.berus
vivant dans les Alpes italiennes et les pays nordiques. Au contraire, la taille moyenne
relativement faible de V.aspis dans l’Oberland semble être uniquement due à des
facteurs climatiques et alimentaires, comme en témoignent ses tailles maximales
comparables à celles des autres populations étudiées.
La difference entre les taux de mélanisme chez V.aspis et chez V.berus dans
l’Oberland semble être en relation avec les types d’habitats fréquentés par ces deux
espèces, soit un milieu essentiellement forestier pour V.berus, et un milieu plus ouvert
et buissonnant pour V.aspis. Les conditions microclimatiques et la pression des pré-
dateurs ne sont pas les mêmes dans ces deux milieux.
Comme en témoigne la comparaison des taux de mélanisme au sein de diffé-
rentes populations de vipères, un milieu forestier à caractère humide semble favoriser
le mélanisme chez les deux espèces, à l’inverse d’un milieu ouvert.
Un taux de mélanisme plus élevé chez les femelles que chez les mâles est
observé dans toutes les stations étudiées; cela semble indiquer que le mélanisme est
particulièrement avantageux pour les femelles des deux espèces et (ou) que ces
dernières sont moins vulnérables à la prédation que les males mélaniques.
98 JEAN-CLAUDE MONNEY, LUCA LUISELLI & MASSIMO CAPULA
Chez les deux espèces, le mélanisme semble accélérer le taux de croissance
des vipères, mais ne semble pas avoir une influence notable sur la vitesse de l’em-
bryogénèse.
La tendance des vipères péliades mélaniques à être plus lourdes que les
normales, et celle des vipères aspics normales à être plus lourdes que les mélaniques
ne semblent pas être un cas particulier et se retrouvent dans des régions très diffé-
rentes.
L'avantage pondéral des vipères aspics normales pourrait être dû à leur âge
moyen supérieur, conséquence d’une croissance plus lente et d’un taux de prédation
inférieur, alors que l’avantage pondéral des vipères péliades mélaniques serait dû à un
taux de croissance plus rapide, mais à une moyenne d’äge identique à celle des
vipères normales.
Le mélanisme est un phénomène encore mal connu et ce caractère ne concerne
probablement pas que la seule coloration (DUGUY & SAINT GIRONS 1988), ce caractère
ne concerne probablement pas que la seule coloration, le génotype des individus
semble modifié de façon notable, comme en témoigne une variation accrue de l’écail-
lure chez les couleuvres mélaniques.
REMERCIEMENTS
Cette étude a pu être réalisée grâce à la collaboration de nombreuses personnes.
Qu’elles trouvent ici l'expression de notre profonde gratitude: Prof. Claude Mermod,
directeur de thèse du premier auteur (Université de Neuchâtel), Prof. Hubert Saint
Girons, pour sa critique du manuscrit (Université de Paris), Lionel Cavin, pour sa
collaboration sympathique sur le terrain, Dr Guy Naulleau, pour sa visite sur le terrain
et ses conseils judicieux (CEBAS, Chizé), Mme Jacqueline Moret, pour le traitement
statistique des données (Université de Neuchâtel), Mr François Reichenbach, pour la
mise à disposition de son chalet d’alpage. Cette étude n’aurait pu se réaliser sans les
autorisations de capture et de marquage décernées par le canton de Berne et sans le
soutien financier du Fonds de la Loterie du canton de Berne, ainsi que de la Ligue
bernoise pour la Protection de la Nature et de la Ligue Suisse pour la Protection de la
Nature. Nous les remercions vivement.
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REVUE SUISSE DE ZOOLOGIE, 103 (1) : 101-114; mars 1996
Diploures Campodéidés de la Pestera de la Movile (Movile Cave),
Dobroudja méridionale (Roumanie).
Bruno CONDÉ
Musée de Zoologie de l’Université et de la Ville de Nancy, 34, rue Sainte-Catherine,
F-54000 Nancy, France.
Diplura Campodeidae from the Pestera de la Movile (Movile Cave),
Southern Dobrogea (Rumania). — The Pestera de la Movile is carac-
terized by its thermomineral waters containing HyS. Most of the terrestrial
species discovered are troglobitic and new for science, a probable conse-
quence of a long isolation.Three species of Campodeidae are recorded
from this cave. While Plusiocampa isterina Condé ressembles endogean
species, two new species (Plusiocampa euxina n. sp. and Campodea
neuherzi n. sp.) present clearly troglomorphic adaptations. They are related
respectively to Plusiocampa evallonychia Silvestri and Campodea taurica
Silvestri known from a cave in the vicinity of Mount Taurus, Peninsula of
Crimea, just facing the southern Dobrogea across the Black Sea (Euxinus-
Pontus).
Key-words: Campodeidae — Taxonomy — Movile Cave — Rumania.
INTRODUCTION
Movile Cave est le nom anglo-saxon de la Pestera de la Movile, grotte de
Roumanie découverte en 1986, à 3 km à l’Ouest de la ville de Mangalia, sur la Mer
Noire, au Sud de la Dobroudja et à proximité de la frontière bulgare. Cette cavité qui
n’avait pas d’entrée naturelle s’ouvre maintenant par un puits creusé dans l’“Obanul
de la Movile” (un “oban” étant une vaste dépression a fond plat) qui a coupé par
hasard une galerie d’accès au niveau supérieur sec, long de 200 m; un niveau infe-
rieur, long de 40 m, est submergé par une eau thermo-minérale, sauf au niveau d’un
lac dont la surface s’ouvre sur l’étage supérieur et de cloches d’air creusées dans le
plafond; la température de l’eau, variant de 20,2° à 20,8°C assure à l’air une tempé-
rature de 20-21°C. Il s’agit donc d’une grotte chaude comparée aux autres cavités de
la même région dont la température est de 10-12°C.
A l'exception de rares épigés venus de l’extérieur par l’entrée artificielle, la
faune terrestre est représentée par des espèces troglobies, nouvelles pour la plupart, ce
Manuscrit accepté le 10.08.1995.
102 BRUNO CONDÉ
qui peut s’expliquer par un isolement de la surface se situant entre la fin du Miocène
et le début du Quaternaire, soit 5,5 à 3 MA (SARBU 1990).
Les Diploures figurent sans commentaire dans le tableau général de la faune
(Sarbu 1990: 194) et j'ai décrit les deux premiers spécimens, récoltés en septembre
1990, sous le nom de Plusiocampa isterina, en soulignant la ressemblance de cette
nouvelle espèce avec Plusiocampa dobati Condé qui peuple des grottes du Jura
souabe, situées sur le cours supérieur du Danube (CONDÉ 1993a).
Depuis, trois autres spécimens m'ont été soumis, deux appartenant à P.
isterina et le troisième représentant une espèce inédite qui, à l’inverse de la
précédente, présente un allongement considérable des appendices, caractère commun
aux Campodéidés dits troglomorphes. Les Campodéidés ont été récoltés à vue sur les
parois des cloches d’air ou sur la plage argileuse dans la salle du lac (Sarbu, in litt.
14.11.91).
Plus récemment, Sarbu (in litt. 01.08.92) a récolté sept nouveaux Campo-
déidés, mais cette fois à la base du puits artificiel d’accès, au voisinage d’appâts
(viande, fromage). Il s’agit d’une espèce de Campodea (Dicampa), appartenant à un
complexe déjà connu d’une grotte de Bulgarie (C. frenata Silvestri) et d’une grotte de
Crimée (C. taurica Silvestri), et présentant des indices d’une évolution cavernicole
(antennes, cerques). En avril et août 1993, Sarbu (in litt. 07.04.93 et 09.08.93) a
retrouvé deux représentants de cette dernière espèce, l’un à la base, l’autre à la partie
supérieure du puits artificiel.
Les holotypes de Plusiocampa euxina et de Campodea (D.) neuherzi, montés
dans le médium II de Marc André, ainsi que 2 paratypes (4 et £) en alcool de C. (D.)
neuherzi sont déposés au Muséum d’ Histoire naturelle de Genève.
DESCRIPTIONS
Plusiocampa isterina Condé, 1993
Station. — Movile Cave, 28.XII.90, S. Sarbu leg.: 1 9 juvénile de 3,50 mm, sans
cerques. — 10.1.91, S. Sarbu leg.: 1 © de 3,94 mm; cerques 2,01 mm; corps/cerques : 1,96.
Tete. — Toutes les antennes ont chacune 23 articles, comme chez les types.
Thorax. — Métanotum du plus grand spécimen avec un macrochete (/a,)
asymétrique, faible, sur la moitié droite (1+2), comme sur la moitié gauche de
l’holotype.
Abdomen. — La papille génitale de la femelle juvénile est très peu pileuse (2+2
phanères sur le volets et 3 sur le tubercule); celle du plus grand individu, au contraire,
possède le même nombre de phanères que le paratype (7+7 sur les volets et 21 sur le
tubercule). Cerques formés d’une base et de 6 articles (7 ou 6 chez les types).
Plusiocampa (s.str.) euxina n. sp.
Station. — Movile Cave, 28.XII.90, S. Sarbu leg.: holotype, & adulte (avec 1 © juvénile
de Plusiocampa isterina Condé).
DIPLOURES CAMPODÉIDÉS DE ROUMANIE 103
Longueurs. — Corps: 4,40 mm; cerque 10,57 mm; corps/cerque = 0,41; patte
metathoracique: 3 mm; corps/p.Ul = 1,46.
Téguments. — Cuticule sans ornementation, soies de revêtement finement
barbelées, certaines sur leurs 2/3 distaux au moins, très peu denses sur les tergites.
FIG. 1
Plusiocampa euxina n. sp., mâle holotype. Organe cupuliforme apical de l’antenne gauche en
coupe optique.
Tête. — Antennes de 45 et 44 articles; le sensille du Ile article subcylindrique,
mince et court (19 um), est postéro-sternal (entre d et e). Le rapport longueur/largeur
de l’article apical, mesuré de l’apex à la limite proximale de la portion pileuse, est
égal à 2,15 et le rapport longueur de l’article/profondeur de l’organe cupuliforme est
égal à 3,43. Ce dernier renferme 8 et 10 sensilles de forme massive, à branches
épaisses, de lecture difficile. Sensille du palpe maxillaire un peu plus épais et plus
court que celui de l’antenne (14 um). Sensille latéro-externe du palpe labial
légèrement conique, de même longueur que le sensille antennaire, mais beaucoup plus
épais; les deux poils de garde, distants l’un de l’autre, sont situés de part et d’autre du
sensille, vers le plan sagittal; le palpe porte en outre une rangée antérieure de 9
phanères ordinaires et un champ de plus de 130 phanères gustatifs.
104 BRUNO CONDÉ
Fic. 2
Plusiocampa euxina n. sp., male holotype. Portion frontale de la capsule céphalique. a, i, p =
macrochètes antérieur, intermédiaire et postérieur de la marge antennaire; x = macrochète de la
suture en Y.
Processus frontal peu saillant, avec un long macrochete glabre antérieur, les 7
autres phanères ressemblant aux soies de revêtement. Phanères de la marge antennaire
un peu barbelés sur leur région distale; longueurs relatives: a = 39,5, i = 60, p = 42.
Phanères x semblables aux intermédiaires (62)
Thorax. — Répartition des macrochètes:
ma la Ip
Aas I 1+1 1+1 2+2
Toul 1+1 2+2 2+2
1005900 1+1 1+1 2+2
Tous sont très robustes et finement barbelés sur leur moitié ou leurs deux tiers
distaux. En II, la, est beaucoup plus long et épais que la; et ma qui se ressemblent
(moyennes des longueurs: ma = 80, la, = 110, la; = 77). Les /p sont les plus longs de
tous les phanères, les /p, surpassant un peu les /p,, sauf en III (152/117, 162/151,
143/148). Soies de revêtement clairsemées, surtout sur le tergite I (ca 6,15 et 15 par
demi-tergite).
DIPLOURES CAMPODÉIDÉS DE ROUMANIE 105
FIG. 3
Plusiocampa euxina n. sp., mâle holotype. — A. Pro-, méso- et métanotum. — B. Patte
métathoracique droite à partir du trochanter, face postérieure. la = macrochète latéral antérieur:
ma = macrochete médial antérieur; ms = macrochète sternal; mt = macrochètes tergaux: p =
proximal, d = distal.
106 | BRUNO CONDÉ
Les pattes métathoraciques dépassent l’extrémité de l’abdomen du tiers distal
de leur tarse, les longueurs relatives des articles étant 16, 13, 54, 73, 53. On notera
que le fémur est beaucoup plus court que le tibia (fe/ti = 0,74), ce dernier étant
environ 12 fois plus long que large et le fémur 6 fois 3/4 seulement. Fémur avec 2
macrochètes tergaux très fortement différenciés, insérés de part et d’autre du milieu
du bord de l’article, le proximal un peu plus long que le distal (61,5 et 56,5). Le
macrochète sternal, beaucoup plus court (38,5), s’insère presque en face du tergal
proximal. Le tibia porte 3 macrochètes sternaux, 2 insérés sur le tiers médian et un au
milieu du tiers distal; ils sont équidistants à gauche, les proximaux étant au contraire
plus proches l’un de l’autre à droite. Tous les phanères du tarse sont barbelés, ceux
des deux rangées sternales beaucoup plus épais, avec une portion apicale glabre, de
même que les 3 longues soies subapicales.
FIG. +
Plusiocampa euxina n. sp., mâle holotype. — A. Griffe postérieure I droite. — B. Griffes II
gauches, face postérieure. — C. Griffe postérieure III gauche.
Griffes peu inégales (1,2-1,3) dont la longueur augmente progressivement de I
à II: antérieures = 56,5; 63,4; 68 um; postérieures = 63,4; 71,6; 76,2 um. Le corps et
les crêtes latérales sont ornementés, mais ces dernières sont peu développées et ne
forment pas de talon proéminent à la griffe postérieure. Processus télotarsaux plus
courts que les griffes, rectilignes et peu atténués à l’apex: le bord sternal présente
quelques barbules sur sa moitié proximale.
DIPLOURES CAMPODÉIDÉS DE ROUMANIE 107
Abdomen. — Répartition des macrochetes tergaux:
la post
Ab. I-III 0 1+1 (post; )
Ab. IV 1+1 4+4 (post) 3 à 5)
Ab. V 2+2 444 (post) 3 à 5)
Ab. VI-VO 2+2 4+5 (posto, asymétrique)
Ab. VIII 0 6+6
Ab. IX 0 9+9 (total du segment)
A tous les tergites, 2 soies marginales postérieures seulement entre les post].
Quand un post, est présent (asymétrique en VI et VII), il est plus grêle et plus court
(45) que ses voisins (57, 68). Valvule supra-anale avec 2 soies sagittales subapicales.
Sternite I avec 8+7 macrochètes très barbelés. Appendices subcylindriques à
large base, égale aux 2/5 environ de leur écartement, et régulièrement arrondis à
l’apex qui porte un champ d’une soixantaine de phanères a,. Pas de champ
glandulaire au sternite.
Sternite II à VI avec 6+6 macrochètes très différenciés. Styles dont les 3
phanères principaux (apical, subapical, moyen sternal) sont volumineux et densément
barbelés dès la base. Sternite VIIT avec 2+2 macrochètes; 13 soies à la rosette du
gonopore.
Le cerque droit est complet, formé d’une base et de 7 articles seulement, dont
les longueurs relatives sont les suivantes: 114 (base), 53, 68, 76, 88, 97, 108, 115. Ces
éléments sont extraordinairement longs: extrêmes 0,78 et 1,69 mm; X = 1,32 mm.
Sauf les courtes soies du verticille apical de chaque article, tous les phanères sont
semblables: longs (extrêmes 0,29 et 0,47 mm; X = 0,38 mm, n = 41), rectilignes et
très finement pubescents sur la plus grande partie de leur longueur.
Affinités. — La présence de deux macrochètes tergaux au fémur, sur laquelle
PACLT (1957: 46) a fondé son sous-genre Didymocampa (espèce type Plusiocampa
sinensis Silvestri, endogé de Chine méridionale) ne semble pas suffisante pour
justifier à elle seule le rapprochement des quatre espèces qui partagent maintenant ce
caractère. CONDÉ & SENDRA (1989: 617) l’ont déjà souligné en décrivant Plusiocampa
alhamae, du Sud de l’Espagne (Granada), qui s'apparente à des formes de la
Méditerranée occidentale, tandis que P. sinensis est une espèce à part, méritant sans
doute la coupure proposéel.
La nouvelle espèce est la plus voisine de Plusiocampa evallonychia Silvestri,
1949, de la grotte I-El-Koba (U-el-Choba dans Silvestri) dans la région du Mont
Taurus, en Crimée. L’une et l’autre présentent des caractères communs aux lignées
troglobies, mais plus accentués chez la nouvelle espèce: antennes de 44 et 45 articles
! Le premier Didymocampa troglobie, proche de sinensis, a été décrit de Chine
méridionale (Hupei): P.(D.) lipsae Condé (CONDÉ 1993b).
108 BRUNO CONDÉ
Fic. 5
Plusiocampa euxina n. sp., mâle holotype. Marge posterieure des urotergites IV, Vici Vile
DIPLOURES CAMPODÉIDÉS DE ROUMANIE 109
FIG. 6
Plusiocampa euxina n. sp., mâle holotype. — A. Urosternite I, diagramme, * = macrochète
asymétrique. — B. Moitié postérieure droite de l’urosternite I.
(vs 33); longueur totale/longueur de la patte métathoracique: 1,46 (vs 2,33); tibia plus
long que le fémur (ti/fe = 1,35 vs 1,33); tarse aussi long que le fémur (ta/fe = 1 vs
0,77). On ne peut pas comparer les caractères sexuels de l’urosternite I, ni ceux des
cerques, ces derniers remarquables chez la nouvelle espece, puisque le seul
représentant de P. evallonychia est une femelle incomplète. Mis à part quelques
détails de la chétotaxie, la forme des griffes et celle des processus télotarsaux écartent
euxina de evallonychia; chez cette dernière espèce, la griffe postérieure possède de
larges crêtes formant un talon qui recouvre l’extrémité du tarse, et les processus
télotarsaux sont asymétriques, le postérieur dépassant l’apex de la griffe
correspondante et l’antérieur étant un peu plus court que la moitié de la griffe.
Etymologie. — Du latin euxinus: du Pont-Euxin, ancien nom de la Mer Noire,
pour rappeler la situation de la localité type de la nouvelle espèce et ses affinités étroites
avec P. evallonychia, du Sud-Est de la presqu'île de Crimée, à quelque 400 km.
110 BRUNO CONDÉ
Campodea (Dicampa) neuherzi n. sp.
Station. — Movile Cave, à la base du puits artificiel d’accès, au voisinage d’appäts
(viande, fromage), S. Sarbu leg.: VII.92: 2 8,5 2; IV.93: 1 9. A la partie supérieure du puits
artificiel, S. Sarbu leg.: VIII.93: 1 larve.
Holotype: 2 de 6,65 mm. Paratypes: 2 d et 5 2.
C.PoeivRE
FIG. 7
Campodea (Dicampa) neuherzi n. sp., ¢ et © paratypes. Habitus montrant les plages blanches,
disposées symétriquement sur le corps, dues a la présence de cristaux d’urates qui réfléchissent
la lumière. Echelle = 2 mm. Détail d’une plage en bas à droite.
DIPLOURES CAMPODÉIDÉS DE ROUMANIE 111
Longueurs. — Corps. 3: 4,78 mm; 9: 6,55-7,60 mm (moyenne: 6,64 mm);
cerques: 9,10 (holotype) et 9,60 mm (2 n° 4); corps/cerques = 0,73 et 0,68; patte
métathoracique, depuis le trochanter: 2, 2,42, 2,48 mm; corps/p.IHI = 2,35 (d), 2,75-
3,06 (2).
Téguments. — Corps brun jaunätre avec des plages blanches disposées
symétriquement sur le corps, la tête, le dernier segment de l’abdomen et les
appendices exceptes. Cette ornementation exceptionelle a pour origine la réflexion de
la lumière, à travers la cuticule, sur des concentrations bien localisées de cristaux
d’urates dont l’opacité rend, par ailleurs, l'observation par transparence difficile.
Cuticule très finement granuleuse sur la face dorsale, cette structure particulièrement
visible sur la capsule céphalique. Soies de revêtement courtes, simples, et sensilles
sétiformes sensiblement de même longueur.
Tête. — Antennes des adultes de (34) 36-40 articles, sauf régénérats (r) souvent
identifiables par les dimensions des articles distaux, l’apical en particulier (-/35r,
36/34, 36/37, 26r/38, 39/40). Antennes de la larve: 30/30.
Le sensille du Ille article, subcylindrique, légèrement attenué aux deux
extrémités, est postéro-tergal (entre b et c). Le rapport longueur sur largeur de l’article
apical, mesuré de l’apex à la limite proximale de la portion pileuse, est égal à 1,86-
2,23 (X = 2,03; n = 4). L’organe cupuliforme apical est très peu profond et le rapport
longueur de l’article sur profondeur de l’organe cupuliforme varie de 7,68 à 12,5 (X =
9,73; n = 4), atteignant 18 à une antenne régénérée; il renferme 8 à 10 petits sensilles
de forme simple. Le sensille du palpe maxillaire et celui du palpe labial sont plus
courts et plus épais que celui de l’antenne; deux courts poils de garde sont insérés de
part et d’autre du sensille labial, mais à quelque distance; le palpe porte en outre une
rangée antérieure de 10 à 14 phanères ordinaires et de 149 (4 de 4,78 mm) à 234 (9
de 7,60 mm) phanères gustatifs.
Processus frontal peu saillant avec 3 macrochètes barbelés sur leur moitié ou
leurs 2/3 distaux. Phanères de la marge antennaire presque glabres, sauf l’antérieur
plus long et robuste que les autres (a = 69; i = 58,5; p = 54 chez l’holotype).
Thorax. — Formule typique du sous-genre Dicampa (3,2,1 par demi-tergite)
dans les rapports suivants:
Ip
ma/la Ip/ma >p/N e/la mall/malll
Woe Il 0,86-0,93 1,76-2 2,32-2,66
malt 0,60-0,78 = = == 1,36-1,63 0,94-1
Les macrochetes, comme les soies marginales, sont robustes et barbeles
presque dès la base. Au mésonotum, les ma sont environ les 2/3 des la et ces derniers
sont égaux aux 3/4 environ de la distance entre les embases ma-la.
Les pattes métathoraciques atteignent le bord postérieur du VIle urotergite, les
longueurs relatives des articles étant 33,5; 15,5; 84; 102; 69. On notera que le fémur
est beaucoup plus court que le tibia (fe/ti = 0,82), ce dernier étant 7,5 fois plus long
que large et le fémur un peu moins de 4 fois (3,8).
io BRUNO CONDE
Trochanters II et II (et parfois I) avec un très petit sensille grêle (13 um), très
faiblement claviforme. Tibia avec un macrochète sternal court et peu différencié,
inséré vers la moitié de la longueur de l’article, les plus souvent en decà du milieu.
Tarse de longueur intermédiaire entre fémur et tibia (ta/fe = 0,82); soies de la double
rangée sternale portant quelques barbules sur leur région moyenne, à l’exception de
celles de la paire subapicale qui sont beaucoup plus gréles et glabres. Griffes simples,
faiblement arquées et processus télotarsaux sétiformes atteignant l’extrémité des
griffes.
Abdomen. — Répartition des macrochètes tergaux: une seule paire de latéraux
postérieurs (/p) en VI et VII, 3 paires en VIII et 5+5 (total) en IX. Valvule supra-anale
avec 2+1+2 (4, 9), 2+2+2 ou 2+3+2 (2 dont l’holotype).
Sternite I avec 6+6 macrochètes. Chez la 2, les appendices sont cylindriques,
3,5 fois à près de 4 fois plus longs que larges (3,50-3,85), un peu atténués à
l'extrémité qui porte 26 phanères a, (9 de 7,60 mm). Chez le d, les appendices sont
subtriangulaires, l’angle le plus aigu vers le plan sagittal, et pourvus de 30-32 a, et de
46-52 a,. Un champ glandulaire étroit comptant 84 phanères sur 2 à 3 rangs sur la
marge postérieure.
Sternites II à VII avec 4+4 macrochètes et une paire de phanères, plus épais que
les soies de revêtement voisines (submacrochètes), de part et d’autre des styles. Soie
apicale des styles avec deux branches basilaires (la plus basale la plus courte) et 2-4
barbules sur la moitié distale; soie subapicale avec de faibles barbules sur la portion
moyenne; soie moyenne sternale fourchue, l’une des branches formant deux fourches
successives. Sternite VIII avec 1+1 macrochètes; papille génitale de la plus grande ©
avec 14 et 13 soies aux volets et 15, dont une médiane, sur le tubercule médian (5+5,
2+2, 1); les 10 soies proximales sont beaucoup plus longues et épaisses que les autres.
Le cerque droit de l’holotype est complet, formé d’une base et de 10 articles de
longueurs croissantes de la base à l’apex. Longueurs relatives: 173 (base), 70, 95,
106, 139, 152, 210, 220, 262, 271, 319 (extrêmes 0,31 et 1,43 mm). La base et les
trois derniers articles sont subdivisés plus au moins nettement en 3, 3, 7 et 8 articles
secondaires. Phanères de longueur modérée, les plus longs de la base atteignant 1,25
fois seulement la largeur de l’appendice; ils sont un peu barbelés sur leur portion
apicale, le nombre et les dimensions des barbules régressant progressivement, et les
phanères des articles terminaux sont tous glabres.
Affinités. — La nouvelle espèce est très voisine de C. (D.) frenata Silvestri,
1931 de la Javoreckata pestera (= caverna Jaworec de Silvestri), près de Lakatnik,
dans la défilé de l’Iskâr (Bulgarie) et de C. (D.)taurica Silvestri, 19492, de la même
grotte de la péninsule de Crimée que Plusiocampa evallonychia Silvestri, 1949.
Une comparaison des principaux caractères des trois espèces du sous-genre
Dicampa montre bien leur haut degré de ressemblance, quoique certains éléments de
2 A la fin de la description originale (SILVESTRI 1949: 29), un lapsus a fait écrire “C.
bulgarica Silv.”, une espèce qui n’existe pas, pour C. frenata. D'autre part, le nom de taurica a
été cité par PLIGINSKY (1927: 173), d’après une lettre de Silvestri de 1914, et la date de récolte
(06.06. 1913) précisée.
DIPLOURES CAMPODÉIDÉS DE ROUMANIE 113
comparaison fassent encore défaut. L’organe cupuliforme apical de l’antenne n’est
décrit que chez neuherzi et atteste une évolution souterraine de cette espèce, con-
firmée par l’allongement des cerques, inconnus chez faurica , mais beaucoup plus
courts chez frenata (0,72 et 1,25); les autres caractères n’ont pas de signification
quant à l’évolution souterraine.
C.(D.) frenata C.(D.) taurica C.(D.) neuherzi
Antennes 26-27 32-40 36-40
Organe cupuliforme % % 8-10
ma/la th.Il 0,533 1> 0,86-0,93
la VI-VII 1+1 0 0
Ip VI 1+1 0 1+1
corps/cerques 125 ? 0,68-0,73
Champ gld. st.I 3-4 rangs 4 rangs 2-3 rangs
Macro. ti. III 0 2 1
3 Mensurations sur les figures originales agrandies.
BIOGÉOGRAPHIE
La présence de deux paires d’espèces très voisines faurica-neuherzi et evallo-
nychia-euxina dans des grottes situées de part et d’autre de la Mer Noire est un bon
exemple de vicariance et, en ce sens, la faune des Diploures de Movile Cave est
exactement celle que l’on pouvait s'attendre à y rencontrer; le couple dobati-isterina
est d’affinités danubiennes, ce qui n’est pas davantage surprenant. La densité des
cristaux d’urates chez neuherzi est finalement la seule singularité rencontrée au cours
de cette étude, mais nous ignorons si frenata et taurica présentent quelque chose
d’analogue.
Etymologie. — Dédiée au Dr Heinz Neuherz, de Graz, qui a étudié les
Diploures et les Palpigrades des grottes d’Autriche.
REMERCIEMENTS
Madame E. Seraoui a assuré la mise au propre de l’abondante illustration de
l’une des nouvelles espèces et Monsieur Cl. Poivre a réalisé l’habitus de l’autre
espèce. Nous leur exprimons notre vive gratitude pour leur aide.
114 BRUNO CONDÉ
BIBLIOGRAPHIE
CONDÉ, B. 1993a. Une lignée danubienne du genre Plusiocampa (Diploures Campodéidés).
Revue suisse de Zoologie 100: 735-745.
CONDÉ, B. 1993b. Premiers Campodeidae cavernicoles de Chine, comme exemple de
l’évolution souterraine de la Famille (Diplura). Revue suisse de Zoologie 100: 823-828.
CONDE, B. & A. SENDRA 1989. Description du premier Campodéidé cavernicole du Sud de la
Péninsule ibérique (Diplura Campodeidae). Revue suisse de Zoologie 96: 611-617.
PACLT, J. 1957. Diplura. Genera Insectorum 212: 123 p.
PLIGINSKY, V. G. 1927. Contributions to the cave fauna of the Crimea. III. Russkoe Entomo-
logicheskoe Obozrenie 21: 171-180.
SARBU, S. 1990. The unusual fauna of a cave with thermomineral waters containing H,S, from
Southern Dobrogea, Romania. Mémoires de Biospéologie 17: 191-195.
SILVESTRI, F. 1949. Descrizione di due specie nuove cavernicole di Campodeidae (Insecta
Diplura) della regione del Monte Tauro. Bolletino del Laboratorio di Entomologia
Agraria “Filippo Silvestri “ 9: 27-31.
REVUE SUISSE DE ZOOLOGIE, 103 (1) : 115-118; mars 1996
A New Species and records of the genus Pseudoagathidium Angelini
from Uganda(Coleoptera, Leiodidae, Agathidiini)
Fernando ANGELINI
S.S.7 per Latiano, Km.0,500, I-72021 Francavilla Fontana (Brindisi) Italy
New species and records of Pseudoagathidium Angelini from Uganda. -
Pseudoagathidium ivani sp. n. is described and P. muehlei Angelini and P.
vignai Angelini & DeMarzo are recorded from Uganda.
Key-words: Coleoptera - Leiodidae - Pseudoagathidium - taxonomy -
Uganda.
INTRODUCTION
Afrotropical Agathidiini are under-represented in collections and appear rare
compared to those from Holarctic and Oriental realms. Thanks to a grant of the
“Académie suisse de sciences naturelles”, G. Cuccodoro, accompanied by D. Erne,
sampled humicolous arthropods in a number of forest remnants in Uganda. The study
of the collected Agathidiini revealed the presence of three species of Pseudo-
agathidium, one of which is new and described below. The material is deposed in the
Muséum d’histoire naturelle, Geneva (MHNG) and in the authors private collection
(AC).
Pseudoagathidium muehlei (Angelini, 1988)
Agathidium (Neoceble) muehlei ANGELINI, 1988: 126.
Material: Uganda, Mt. Elgon, Sipi, 1750 m, 31.V.1993, leg. Cuccodoro & Erne, 2 d
and 1 © in MHNG, 2 G in AC; Mt. Elgon, Sipi, 1650 m, 29.V.1993, leg. Cuccodoro & Ere, 2
? in MHNG; Kibale forest, 1250 m, 23.V.1993, leg. Cuccodoro & Erne, 1 d in MHNG.
Distribution: Rwanda and new for Uganda.
Manuscript accepted 20.03.1995.
116 FERNANDO ANGELINI
Pseudoagathidium vignai (Angelini & DeMarzo, 1984)
Agathidium (Neoceble) vignai ANGELINI & DE MARZO, 1984: 355; 1985: 76; ANGELINI & PECK,
1984: 420.
Material: Uganda, Mt.Elgon, Sipi, 1750 m, 1.VI.1993, leg. Cuccodoro & Erne, 1 9 in
MHNG, 1 6 in AC; Ruwenzori, above Kilembe, 2000 m, 4.V.1993, leg. Cuccodoro & Erne, 1
2 inMHNG.
Distribution: Zaire and new for Uganda.
Pseudoagathidium ivani sp. n. (Figs 1-7)
Length 2.55-2.85 mm (holotype d 2.6 mm). Dorsum of head and pronotum
black, pronotum dark reddish-brown; venter reddish-brown, mesosternum testaceous;
antennae testaceous with dark club; legs reddish-brown. Microreticulation absent,
only vague traces on whole dorsum; punctures small and superficial on whole
dorsum.
Head: Microreticulation absent, or only vague traces; punctures small, super-
ficial, spaced from each other by 2-10 times their own diameter. Widest at eyes (fig.
2); eyes protuberant; clypeus sharply excavated; clypeal line absent. 3rd antennal
segment 1.36 times as long as the 2nd and as long as than 4th + 5th together (fig. 1).
Pronotum: Microreticulation absent, only vague traces as on head; punctation
as that of head, spaced from each other by 5-10 times their own diameter. 1.44 times
as broad as head, moderately broader than long (W/L = 1.69) and moderately convex
(W/H = 1.58). Anterior margin very bent (fig. 3), lateral outline strictly bent (fig. 4).
Holotype: length 0.75 mm, width 1.27 mm, height 0.80 mm.
Elytra: Microreticulation absent, or only vague traces as on head; punctation as
that of head, but punctures spaced from each other by 2-15 times their own diameter.
Just a little broader than pronotum, moderately broader than long (W/L = 1.2) and
very convex (W/H = 1.07); lateral outline with sharp humeral angle, at the middle;
sutural striae weak and extending beyond the anterior 2/3 of elytra. Holotype: length
1.15 mm, width 1.38 mm, height 1.28 mm.
Metathoracic wings present. Meso- and metasternum: median carina weak
anteriorly, more evident posteriorly, lateral lines absent, femoral lines short, with a
tubercle between the metacoxae.
Legs: Tarsal formula d and 9 4-4-4.
Male copulatory organ as in figs 5-6.
Spermatheca as in fig. 7.
DISCUSSION
Pseudoagathidium ivani sp. n. is similar to P. vignai (Angelini & De Marzo),
(1984: 355, Zaire; 1985: 76) in absence of microreticulation on dorsum, punctation of
dorsum, form of the posterior tibiae; it differs only by lower length ratio 3rd/2nd
PSEUDOAGATHIDIUM FROM UGANDA 117
Fics 1-7
Pseudoagathidium ivani sp. n. : 1, antenna; 2, head; 3, pronotum in dorsal view; 4, pronotum
and elytra in lateral view; 5-6, aedeagus in lateral and ventral view; 7, spermatheca. Scale in
HeM2=02 mmn34=0Simm;ins"67=0 mm:
118 FERNANDO ANGELINI
antennal segments and the presence of vague traces of microreticulation on dorsum;
to distinguish the two species it is necessary to examine the aedeagi and sper-
mathecae.
Derivatio nominis: Dedicated in homage to Dr Ivan Löbl, Geneva.
HOLOTYPE d: Uganda, Mt. Elgon, Sipi, 1750 m, 31.V.1993, leg. Cuccodoro & Erne, in
MHNG.
PARATYPES: same data as holotype, 1 2 in MHNG, 1 d and 1 2 in AC.
ACKNOWLEDGEMENTS
I thank Dr Ivan Löbl, Geneva, for the loan of the material and my friend
Jonathan Cooter, Hereford, for help with the translation of the text into English.
REFERENCES
ANGELINI, F. 1988. Descrizione di Agathidium muehlei sp. n. del Rwanda e ulteriori dati sugli
Anisotomini dell'Africa centrale. Bolletino della Società entomologica italiana,
Genova, 120 (2): 125-128.
ANGELINI, F. & L. DE Marzo. 1984 (1985). Note sistematiche sugli Agathidium dell'Africa
centrale e descrizione di tre nuove specie. Fragmenta entomologica, Roma, 17 (2): 347-
358.
ANGELINI, F. & L. DE Marzo. 1985. Agathidium e Afroagathidium dell'Africa centrale. Ento-
mologica, Bari, 20: 69-80.
ANGELINI, F. & S.B. PECK. 1984. Agathidiini from South Africa (Coleoptera: Leiodidae: Leio-
dinae): a synopsis and description of a new genus and species. Annals of the Transwaal
Museum, Pretoria, 33 (28): 419-425.
REVUE SUISSE DE ZOOLOGIE, 103 (1) : 119-150; mars 1996
A Revision of the Madeiran species of the genus Geostiba
Thomson, 1858 (Coleoptera: Staphylinidae)
Volker ASSING ! & Paul WUNDERLE 2*
! Gabelsbergerstr. 2, D-30163 Hannover
> Hehnerstr. 15, D-41069 Mönchengladbach
A Revision of the Madeiran species of the genus Geostiba Thomson,
1858 (Coleoptera: Staphylinidae). - A revision of the types of all
Madeiran taxa of Geostiba Thomson as well as of further material
collected by the authors and from various collections yielded 15 species,
all of them endemic to the Madeiran archipelago. 11 species are described
for the first time: G. arieiroensis n. sp., G. ruivomontis n. sp., G. bica-
canaensis n. Sp., G. brancomontis n. sp., G. graminicola n. sp., G. vacci-
nicola n. sp., G. lauricola n. sp., G. caligicola n. sp., G. occulta n. sp., G.
endogea n. sp. and G. subterranea n. sp. Both G. israelsoni (Palm) and G.
waldeni (Palm) are considered junior synonyms of G. filiformis (Woll.),
and G. carli Palm is recognized as a synonym of G. lindrothi Franz. Sıpalia
leileri Palm is transferred to Atheta Thoms. Atheta juengeri Benick is
shown to be a junior synonym of Sipalia leileri Palm. The study includes
detailed descriptions of each species, illustrations of diagnostic characters,
faunistic and ecological data as well as a key allowing identification.
Key-words: Coleoptera - Staphylinidae - Geostiba - Madeira - Taxonomy.
INTRODUCTION
In his first account of Madeiran insects WOLLASTON (1854) described the genus
Xenomma with three species. Only the first of these (X. planifrons) has remained in the
genus, which today is placed in the Oxypodini. PALM (1981a) treated the other two
species (X. formicarum and X. filiformis) as members of Sipalia Muls. & Rey — a name
erroneously used and to be replaced with Geostiba Thomson (e.g. BLACKWELDER
(1952)) — and described four further species from the Madeiran archipelago: Sipalia
leileri, S. lindrothi, S. israelsoni and S. waldeni. In the same year FRANZ (1981) added
Geostiba lindrothi and G. portosantoi, and S. lindrothi Palm was renamed Geostiba
carli (PALM 1981b). Thus a total of 8 endemic species of Geostiba have so far been
described from Madeira and Porto Santo.
* Names in alphabetical order.
Manuscript accepted 31.03.1995.
120 VOLKER ASSING & PAUL WUNDERLE
During a joint excursion to Madeira proper and Porto Santo in spring 1993 we
collected mostly large series of various species of Geostiba. Their identification,
however, based on the original descriptions and the key in PALM (1981a) failed in
most cases. À subsequent examination of all the types revealed that not only did our
material contain a number of yet undescribed species, but some of the previously
described taxa had also been misinterpreted.
Material from the following museum collections was studied: The Natural
History Museum London (BMNH), the Muséum d'histoire naturelle, Genève
(MHNG), the Museum of Zoology and Entomology Lund (MZEL) and the Swedish
Museum of Natural History Stockholm (SMNH).
THE MADEIRAN SPECIES OF GEOSTIBA THOMSON
Our study of 812 specimens of Madeiran Geostiba both from our own and
from loaned material yielded 15 species, 11 of them are described as new. Three
names are placed in synonymy, and one species is excluded from the genus. Some
characters such as colour, punctation, microsculpture, body size and even body
proportions may be subject to considerable variability. The presence or absence of
functioning eyes, the shape of the genitalia and the sclerites of the abdominal segment
VII generally represent more reliable diagnostic criteria. As in Italian Geostiba (PACE
1977) and certain Oxypoda (ZERCHE, pers. comm.), the form of the apex of the
parameres and the parameral sensilla were found to be of high taxonomic value. For a
proper evaluation of these characters, particularly in the small subterraneous species,
the genitalia and the genital segments were mounted on slides.
In the descriptions, measurements of head width (HW), pronotal width (PW)
and length (PL), length of elytra at suture (EL) and the length from labrum to elytral
apex (SL) are indicated in um, the total length from labrum to hind margin of tergite
VII (TL) is given in mm. Measurements of body length, of course, strongly depend
on the mode of preparation. Whenever series of specimens were available, the range
of the measurements, the arıthmetic mean and standard deviation are also presented.
Paratypes of most of the new taxa described here as well as part of the material
of the remaining species collected by us are deposited in the Muséum d'histoire
naturelle, Geneve (MHNG).
Geostiba formicarum (Wollaston, 1854) (Figs la—k)
Xenomma formicarum WOLLASTON, 1854: 545; 1857: 172; 1865: 457.
Geostiba formicarum (Wollaston), FAUVEL 1897: 338.
Sipalia formicarum (Wollaston), JANSSON 1940: 16.
nec Sipalia formicarum (Wollaston), PALM, 1981a: 294.
Types: Lectotype: d, here designated and labelled accordingly, in coll. Wollaston.
(BMNH). Paralectotypes: 1 d, 1 2, in coll. Wollaston (BMNH).
The specimen here chosen as lectotype carries a round label ‘type’ and a hand-written
label ‘Xenomma formicarum Woll., type’. Since the former is a “curator label’ (HAMMOND,
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 121
Fic. la—k
Geostiba formicarum (Woll.) median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermatheca (d); right mandible (e); 4 tergite (f) and sternite (g) VII; © hind
margin of sternite VIII (h); antenna (i); labium (j); maxilla (k); pubescence partly omitted in
f-1. Scales: a-b, d-e, h-k: 0.1 mm; c: 0.05 mm; f-g: 0.25 mm.
122 VOLKER ASSING & PAUL WUNDERLE
pers. comm.) and therefore of no taxonomic significance and as Wollaston based his original
description on several specimens without fixing a type, the three specimens in the Wollaston
collection must be regarded as syntypes.
Further material studied: 18 8 8, 15 9 9: Madeira, Rabacal, 1050 m, 31.11.1993, leg.
& coll. Assing & Wunderle, MHNG; 1 4: Rabacal, 7.VIIL.1975, leg. Vit (coll. Assing).
DESCRIPTION
Measurements: HW: 380-460, 408+22; PW: 450-590, 527431; PL: 450-520,
485+20, EL: 250-320, 296+22; SL: 1100-1330, 241+72; TL: 3.0-3.6, 3.37+0.18.
Colour of body including the appendages reddish yellow to yellowish brown;
tergites III-VI increasingly darkened, tergites HI and IV mostly with slightly darker
antero-central area, most of the surface of tergites V and especially VI, except for the
hind margins, generally brown to blackish brown; tergite VII clearly lighter than
tergite VI, often brownish in anterior half, tergite VIII reddish yellow; colour of the
abdomen, on the whole, very variable.
Head clearly (0.76—0.80x) narrower than pronotum, with reticulate micro-
sculpture and therefore subdued shine, punctation mostly shallow and indistinct; eyes
present and functioning, their diameter equal to or slightly exceeding maximal width
of first antennal segment; antennae longer than head (measured from anterior margin
of labrum) and pronotum together (Fig. 11); mouthparts as in Figs le, j, k.
Pronotum usually slightly wider (0.97-1.15) than long, with distinct reticulate
microsculpture and often almost effaced punctation, caudadly directed pubescence
short and rather inconspicuous.
Elytra much shorter (0.55—0.67x) and slightly wider than pronotum, with
distinct granulose punctation, surface more shining than that of pronotum; alae reduced.
Abdomen increasing in width from segment III to V or VI, its maximal width
slightly exceeding that of elytra; surface somewhat shining, but with clear reticulate
microsculpture, and with rather sparse fine punctation.
d: hind margin of tergite VIII slightly concave, that of sternite VIII convex to
bluntly angled (Figs. 1f—g); median lobe and apex of paramere as in Figs. la-c.
2: posterior margin of sternite VIII weakly concave with a conspicuous row of
numerous bristles (Fig. 1h); spermatheca as in Fig. Id.
DISTRIBUTION AND BIONOMICS
WOLLASTON (1854) reports G. formicarum, an endemic of Madeira proper,
from the southern slope of Pico Arieiro and from the Lombo das Vacas in the north of
the island (1 and 2 specimens, respectively). We sieved numerous specimens from
soil and leaf litter in Laurus stands near Rabacal (1050m), where it has also been
collected by Lundblad (JANSSON 1940) and Vit. It has been recorded at the end of
March, in June and during the period from the end of July to the first half of August.
The presumed affinity to ants (name!), which WOLLASTON (1854) concludes from the
fact that he took his specimens “from beneath stones in the neighbourhood of ants'
nests”, appears very doubtful, particularly since G. formicarum does not possess any
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 123
of the external characters typical of other myrmecophilous Aleocharinae. Our own
observations do not confirm any such relationship either.
COMMENTS
PALM's (1981a) description and illustrations do not refer to this species, but to
the endemic Atheta (Mocyta) sanguinolenta (Wollaston, 1854), a light-coloured bra-
chypterous species which PALM apparently mistook for a Geostiba and whose types
the first author had the opportunity to study in the BMNH. Specimens of A. sangui-
nolenta in Palm's collection, which does not contain any true G. formicarum, are
labelled ‘Sipalia formicarum .
Geostiba filiformis (Wollaston, 1854) (Figs 2a-i)
Xenomma filiforme WOLLASTON, 1954: 545; 1857: 172; 1865: 457.
Geostiba filiformis (Wollaston) FAUVEL, 1897: 338.
nec(?) Sipalia filiformis (Wollaston), JANSSON 1940: 16.
Sipalia waldeni PALM, 1981: 298; syn. nov.
Sipalia israelsoni PALM, 1981: 298; syn. nov.
nec Sipalia filiformis (Wollaston), PALM, 1981: 297.
Types: Lectotype: 2, ‘Xenomma filiforme Woll., type’, here designated and labelled
accordingly, in coll. Wollaston (BMNH).
Paralectotype: 1 9, in coll. Wollaston (BMNH).
There were three syntypes (see remarks below G. formicarum) in Wollaston's collec-
tion, one of them heavily damaged. According to the original description two of the specimens
were collected on Porto Santo and one on Madeira proper.
Further material studied: 4 86, 6 99: Porto Santo, Pico do Castelo, 400 m,
1.1V.1993, leg. & coll. Assing & Wunderle; 28 4 4,31 2 9: Porto Santo, Pico Juliana, 400 m,
1.1V.1993, leg. & coll. Assing & Wunderle, MHNG; 1 6, 1 2: Porto Santo, Pico Branco, H.
Franz leg.; 1 2: Porto Santo, Pico Juliana, 3.11.1978, Waldén leg., Allotypus ‘Sipalia israel-
soni det. Thure Palm, coll. Palm (MZEL); 1 2: Madeiro, Faja da Pedro, 11.11.1978, Waldén
leg., Holotypus ‘Sipalia waldeni’ det. Thure Palm, coll. Palm (MZEL); 7 Ex.: Porto Santo, Pico
Branco, 10.1V.1970, leg. & coll. Franz.
DESCRIPTION
Measurements: HW: 345-365, 355+7; PW: 435-500, 474416; PL: 405-470,
434+19; EL: 280-350, 308416; SL: 1100-1230, 1164+44; TL: 3.0-3.5, 3.29+0.11.
Colour of body including the appendages yellow to reddish yellow; abdomen
dorsally darkened to various extents ranging from only slightly darker hue on central
area of tergites III to V and on anterior half of tergites VI and VII to almost com-
pletely blackish or darkbrown colour of tergites III to VII leaving only the hind
margins reddish yellow, anterior half of tergite VIII sometimes darkened, too.
Head with sides less rounded than in G. formicarum, clearly (0.72—0.79x)
narrower than pronotum, with reticulate microsculpture (similar to G. formicarum),
punctation usually distinct but often shallow; eyes present and functioning (ca. 6-8
ommatidia), their diameter (ca. 70 um) exceeding maximal width of first antennal
segment; antennae somewhat shorter than head and pronotum together (Fig. 21).
Pronotum a little wider (1.07-1.14) than long, with distinct reticulate micro-
sculpture; punctation variable, with transitions from deep and distinct to shallow and
fine; pubescence even shorter and less conspicuous than in G. formicarum.
124 VOLKER ASSING & PAUL WUNDERLE
FIG. 22-1
Geostiba filiformis (Woll.) median lobe in ventral and in lateral view (a, b); apex of paramere
(c); spermatheca (d); & tergite (e) and sternite (f) VIII; © sternite VIII (g-h); antenna (i);
pubescence partly omitted in fi. Scales: a-b, d-i: 0.1 mm; c: 0.05 mm.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 125
Elytra much shorter (0.68-0.75x) than pronotum, but relatively longer than in
G. formicarum; maximal width only indistinctly exceeding that of pronotum; with
usually distinct granulose punctation and with reticulate microsculpture; d d with a
characteristic longitudinal elevation on anterior half of both elytra near suture; alae
reduced.
Abdomen with maximal width slightly exceeding that of elytra; surface with
sparse fine punctation and somewhat shining, but with clear isodiametric to transverse
microreticulation.
d : tergites VII and VII with pronounced granulose punctures; hind margin of
tergite VIII slightly convex, that of sternite VIII bluntly angled (Figs. 2e-f); aedeagus
with conspicuous spines in internal sac, median lobe and apex of paramere as in Figs.
2a—<.
2: posterior margin of sternite VIII + convex, slightly concave or straight in
the middle, with a row of numerous bristles (Figs 2g—h); spermatheca as in Fig. 2d.
DISTRIBUTION AND BIONOMICS
G. filiformis appears to be the only Madeiran Geostiba occurring on both
Madeira proper, where it is apparently very rare, and Porto Santo. WOLLASTON (1854)
collected his Madeiran specimen above Funchal and the specimens from Porto Santo
“beneath stones ... on ... grassy mountain slopes” in April and “early spring”. The
records indicated by JANSSON (1940) obviously refer to a different species, since what
he considers to be G. filiformis is only 1.9-2.4 mm long. We sieved our specimens,
some of them still immature, together with larvae that apparently belong to the
species, on April | from deep and moist leaf litter and under logs of dead wood on the
northern slopes of Pico Juliana and Pico do Castelo (Porto Santo).
COMMENTS
In his account of Madeiran Geostiba PALM (1981a) clearly misiterprets G.
filiformis, although he states that he has seen the type. His description, illustrations
and records of G. filiformis as well as the corresponding material in his collection
refer to an as yet undescribed species dealt with below. In addition, he describes G.
israelsoni n. sp. from Porto Santo without commenting on the fact that WOLLASTON
(1854) reports G. filiformis from the same island in his original description. More-
over, PALM (1981a) describes another new species, G. waldeni n. sp., on the basis of
one 2 without antennae stating that G. waldeni is very similar to G. israelsoni, but
“ein wenig grösser, das [!] Hinterleib in grösserem Umfang verdunkelt und besonders
durch die Form der Spermatheca distinkt abweichend” (p. 298). Our examination of
the types revealed that G. israelsoni (Palm) is, without doubt, conspecific with G.
filiformis (Woll.). Similarly, with regard to all the characters available the (damaged)
type of G. waldeni is well within the range of variability of G. filiformis, and PALM's
differential diagnosis is based on two of the most variable characters as well as on
inadequate drawings of the spermathecae. Therefore, we regard both G. israelsoni
(Palm) and G. waldeni (Palm) as junior synonyms of G. filiformis (Woll.).
126 VOLKER ASSING & PAUL WUNDERLE
Geostiba arieiroensis spec. nov. (Figs. 3a-i)
Sipalia filiformis (Woll.), PALM, 1981: 297f.
Types: Holotype: 4 , Madeira, Pico Arieiro, 1600 m, 26.11.1993, leg. & coll. Assing.
Paratypes: 42 d d, 32 2 2: Madeira, Pico Arieiro, 1600 m, 26.1I1.1993, leg. & coll.
Assing & Wunderle; 81 4 4,88 2 2: Madeira, Pico Arieiro, 1600 m, 3.1V.1993, leg. & coll.
Assing & Wunderle, MHNG; 10 Ex.: Pico Arieiro, 1600 m, 7.11.1978, leg. & coll. Palm
(MZEL); 4 Ex.: Pico Arieiro, 23.1V.1978, leg. & coll. Palm (MZEL); 1 d: Pico Arieiro, Rib.
das Calas, 4.1V.1967, leg. & coll. Franz.
DESCRIPTION
Measurements: HW: 325-370, 354+12; PW: 390-425, 411+9; PL: 345-370,
362+8; EL: 225-245, 233+7; SL: 980-1090, 1024+47; TL: 2.5-2.8, 2.66+0.08.
Colour of body including the appendages yellowish with most of tergites V
and especially VI usually clearly darker, light brown to blackish, sometimes also
central area of tergites III and IV and anterior half of tergite VII slightly darkened.
Head narrower than pronotum (0.82-0.90x), but relatively wider than in the
two preceding species, with pronounced reticulate microsculpture and subdued shine
(similar to G. formicarum), punctation usually distinct, but often shallow; eyes pre-
sent and functioning (mostly 4 ommatidia), but small, their diameter (40-50 um)
shorter than maximal width of first antennal segment; antennae subequal in length to
or slightly shorter than head and pronotum together (Fig. 3e).
Pronotum a little wider (1.12-1.17) than long, its maximal width a short
distance behind the anterior angles; with reticulate microsculpture similar to that of
head; punctation usually very fine and indistinct; pubescence short and inconspi-
cuous; 6 d often with wide and shallow longitudinal impression on disk.
Elytra considerably shorter (0.60-0.67x) than pronotum; maximal width sub-
equal to that of pronotum; microsculpture much weaker than on head and pronotum,
therefore surface of shiny appearance; with distinct granulose punctation; alae
reduced.
Abdomen with maximal width slightly exceeding that of elytra; microreti-
culation and punctation as in G. filiformis.
d : hind margin of tergite VIII with pronounced concavity, posterior margin of
corresponding sternite rounded to obtusely pointed (Figs. 3f-g); median lobe with
conspicuous ventral process; median lobe and apex of paramere as in Figs. 3a—c.
9: hind margin of tergite VIII slightly concave, straight or weakly rounded;
sternite VIII posteriorly with shallow concavity and an interrupted row of long
bristles (Figs 3h-i); spermatheca as in Fig. 3d.
DISTRIBUTION AND BIONOMICS
So far this new species has only been recorded from the type locality (name!)
in early spring (February through April). We extracted our specimens from deep litter
layers and soil in stands of Erica sp. (arborea or scoparia) and Vaccinium padifolium
in both northern and southern expositions. On April 3, the ovaries of several 9 9
contained a mature egg.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 127
e
A)
NA bake h
KP Be N
SS Ae
lA i
AT he TT SI Ye
FIG. 3a-1.
Geostiba arieiroensis spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermatheca (d); antenna (e); d tergite (f) and sternite (g) VIII; sternite VIII
(h-1); pubescence partly omitted in e-i. Scales: a-b, d-i: 0.1 mm; c: 0.05 mm.
128 VOLKER ASSING & PAUL WUNDERLE
Geostiba ruivomontis spec. nov. (Figs. 4a-f)
Types: Holotype: d, Madeira, Ach. do Teixeira, 1350 m, 6.1V.1993, leg. & coll.
Assing.
Paratypes: 188 4,8 2 2: Madeira, Ach. do Teixeira, 1600 m, 6.1V.1993, leg. & coll.
Assing & Wunderle; 768 8,52 9 2: Madeira, Ach. do Teixeira, 1350 m, 6.IV.1993, leg. &
coll. Assing & Wunderle; MHNG.
DESCRIPTION
Measurements: HW: 320-365, 348+10; PW: 370-420, 400+13; PL: 320-365,
349+12; EL: 210-230, 225+8; SL: 920-1050, 986432; TL: 2.2—2.8, 2.53+0.15.
In external characters highly similar to the preceding species. It differs from G.
arieiroensis in its on average slightly smaller body size, in that tergite VII is usually
also darkened and of about the same colour as tergite V, in that the microreticulation
of head and pronotum is often less distinct and in that the dd lack the longitudinal
impression on the disk of the pronotum. The most reliable differences are, however,
found in the shape of the sclerites of the abdominal segment VIII and the genitalia.
d: hind margin of tergite VIII + straight or slightly concave (Fig. 4e), that of
sternite VIII strongly convex (Fig. 4f); median lobe with characteristic ventral process
(Figs. 4a—b), apex of paramere as in Fig. 4c.
2: posterior margin with shallow concavity, similar to G. arieiroensis; sper-
matheca as in Fig. 4d.
DISTRIBUTION AND BIONOMICS
At present, G. ruivomontis is only known from the northeastern slope of the
Pico Ruivo (name!) near the Achada do Teixeira. We collected it from debris and
moss in the shade of big rocks at 1600 m and from litter in a stand of old Erica sp. in
northern exposition at 1350 m.
Geostiba bicacanaensis spec. nov. (Figs. 5a—f)
Types: Holotype: d, Madeira, Bica da Cana, 1550 m, 29.11.1993, leg. & coll. Assing.
Paratypes: 62 8 d, 37 9 9: Madeira, Bica da Cana, 1550 m, 29.11.1993, leg. & coll.
Assing & Wunderle, MHNG.
DESCRIPTION
Measurements: HW: 315-350, 33649; PW: 370-400, 385+9; PL: 320-370,
344411; EL: 210-245, 223410; SL: 860-1020, 954+47; TL: 2.1-2.6, 2.34+0.13.
In external characters also highly similar to the two preceding species. G. bica-
canaensis is, on average, slightly smaller, its pronotum generally a little narrower;
length of eyes ca. 45-55 um, with 4-5 ommatidia; colour of tergites as in G. ruivo-
montis; microsculpture of head and pronotum as in G. arieiroensis; 3 d without
longitudinal impression on disk of pronotum. For a safe identification of this species
an examination of the genitalia, particularly of the males, is essential.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 129
Zs A x \ |
Molt \ Fr
|
Fic. da-f
Geostiha ruivomontis spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermatheca (d); d tergite (e) and sternite (f) VIII; pubescence partly omitted in
e-f. Scales: a-b, d-f: 0.1 mm; c: 0.05 mm.
d : hind margin of tergite VIII straight or very shallowly concave (Fig. 5e), the
corresponding sternite rounded posteriorly (Fig. 5f); median lobe without ventral
process (Figs. 5a—b), apex of paramere as in Fig. 5c.
9: posterior margin of sternite VIII slightly concave to almost straight, similar
to arieiroensis; spermatheca as in Fig. Sd.
DISTRIBUTION AND BIONOMICS
G. bicacanaensis has hitherto only been recorded from the type locality. It was
collected on the northern slope of Bica da Cana (name!) from deep litter in stands of
Erica sp. and Vaccinium padifolium. The gonads of several 2 2 contained a mature
egg.
130 VOLKER ASSING & PAUL WUNDERLE
ÉIGRS AE
Geostiba bicanaensis spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermatheca (d); d tergite (e) and sternite (f) VIII; pubescence partly omitted in
e-f. Scales: a—b, d-f: 0.1 mm; c: 0.05 mm.
Geostiba portosantoi Franz, 1981 (Figs 6a-h, 8a)
Geostiba portosantoi FRANZ, 1981: 329ff.
Types: Holotype: d, Porto Santo, Pico Juliana, leg. & coll. Franz.
Paratype: 1 ©, same data as holotype. leg. & coll. Franz.
FURTHER MATERIAL STUDIED: 22 dd, 15 9 9: Porto Santo, Pico Juliana, 400 m,
1.IV.1993, leg. & coll. Assing & Wunderle, MHNG.
DESCRIPTION
Measurements: HW: 250-275, 262+5; PW: 310-350, 334410; PL: 300-350,
B27 NOS EE 210245 23182 SE: 7710-9405 8632-462 lle 025259 alee
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 131
FIG. 6a-h.
Geostiba portosantoi Franz: median lobe in ventral and in lateral view (a, b); apex of paramere
(c); spermatheca (d); antenna (e); d tergite (f) and sternite (g) VIII; hind margin of @ sternite
VIII (h); pubescence partly omitted in e-h. Scales: a-b, d-h: 0.1 mm; c: 0.05 mm.
Colour of body including appendages and tergites more or less uniformly
yellowish to reddish yellow.
Head slender, suboval (Fig. 8a), clearly narrower than pronotum (0.74-0.81x)
and than in G. arieiroensis and allied species; with variable, but usually weaker
microreticulation and therefore more shine than in G. arieiroensis; punctation shallow
and sparse; length of antennae slightly exceeding that of head and pronotum together
(Fig. 6e); small eyes (maximal diameter ca. 40 um) present and functioning, with
usually 4 ommatidia. (Note that FRANZ (1981) states that this species is blind.)
Pronotum with lateral margins almost subparallel in the middle, about as wide
as long (1.0-1.05x), smaller than in the preceding species and somewhat narrower
than elytra; microsculpture and punctation similar to those of head; pubescence short,
sparse and inconspicuous.
Elytra considerably shorter (0.68—0.74x) than pronotum, microsculpture simi-
lar to, sometimes even more distinct than, that of head and pronotum; with granulose
punctation, which is, however, spaced more sparsely than in G. arieiroensis; alae
reduced.
132 VOLKER ASSING & PAUL WUNDERLE
Abdomen with maximal width slightly exceeding that of elytra; dorsal surface
with isodiametric microreticulation similar to that of elytra and with fine and sparse
punctation.
d: hind margin of tergite VIII + straight, that of the corresponding sternite
rounded (Figs. 6f-g); median lobe and apex of paramere as in Figs. 6a—c.
9: hind margin of tergite VIII weakly rounded, sternite VIII weakly concave
(Fig. 6h); spermatheca as in Fig. 6d.
DISTRIBUTION AND BIONOMICS
G. portosantoi is endemic to Porto Santo, where it has only been recorded
from the northern slope of Pico Juliana. FRANZ (1981) sieved the type material under
stones and dead trunks of Pinus sp. We collected our specimens from more or less
isolated spots with deep and still rather moist litter and soil. Since the distribution of
G. portosantoi is apparently restricted to a (forest) area of only a few hectares, where
— according to our observations — suitable conditions are scarce, this species must
be considered highly endangered. Small scale deforestation or even a fire might result
in its extinction.
Geostiba brancomontis spec. nov. (Figs. 7a-c, 8b)
Types: Holotype: d, Porto Santo, Pico Branco, Gipfelbereich, 13.1V.1968, leg. and
coll. H. Franz.
Paratypes: 1 4,2 9 2: same data as holotype, coll. Assing & Wunderle.
d
Fic. 7a-d
Geostiba brancomontis spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermatheca (d). Scales: a—b, d: 0.1 mm; c: 0.05 mm.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 133
DESCRIPTION
Measurements: HW: 260-275; PW: 310-335; PL 290-305; EL: 200-215; SL:
750-770.
Body entirely yellowish; in size and general appearance highly similar to the
following 4 species (see below), from which it is, however, externally distinguished
by the presence of functioning eyes (maximal diameter ca. 40 um) with 4 ommatidia.
G. brancomontis differs from G. portosantoi especially in the distinctly shorter head
(measured from anterior margin of clypeus to posterior margin of head), which is
about as wide as long in the former and clearly longer than wide (ca. 1.15x) in the
latter (Fig. 8b). Compared with G. portosantoi this species is also on the whole even
smaller, and the pronotum is on average narrower and shorter with its maximal width
a short distance behind the anterior angles and its sides more clearly converging
posteriorly.
d: hind margin of tergite VIII obtusely pointed, that of the corresponding
sternite convex, similar to portosantoi; internal sac with sclerotized spines (Fig.
7a-b), apex of paramere as in Fig. 7c.
9: hind margin of tergite VIII convex or very weakly obtuse, posterior margin
of sternite VIII with weak concavity centrally; spermatheca smaller than in porto-
santoi and of different shape (Fig. 7d).
DISTRIBUTION AND BIONOMICS
G. brancomontis is endemic to Porto Santo and apparently exclusively inhabits
the Pico Branco (name!). We have not seen the type locality during our visit to the
island. It would be interesting to examine if suitable biotopes and this species still
exist.
FIG. 8a—b
Habitus (forebody) of Geostiba portosantoi Franz (a) and G. brancomontis spec. nov. (b).
Scale: 0.5 mm.
134 VOLKER ASSING & PAUL WUNDERLE
Geostiba lindrothi Franz, 1981 (Figs. 9a—h)
Geostiba lindrothi FRANZ, 1981: 329.
Sipalia lindrothi PALM, 198 1a: 297.
Geostiba carli nom. nov. for Sipalia lindrothi PALM, 1981b: 447; syn. nov.
Types: Holotype: © , Madeira, Queimadas, 3.1V.1967, leg. & coll. Franz.
Paratype: 1 © : same data as holotype, leg. & coll. Franz.
Types of G. carli Palm: Holotype d, Madeira, Ribeira do Inferno (1150 m),
10.VIIL.1935, leg. & coll. Lundblad (SMNH). Paratypes: 11 Ex., same data as holotype; 1 Ex.,
Caramujo (1250 m), 6. — 14.VIII.1935, leg. & coll. Lundblad (SMNH); 1 9: Madeira,
Funchal, 500-1200 m, Terr. Luta, Febr. 1966, Palm (MZEL)
FURTHER MATERIAL STUDIED:
13 dd, 6 9 L: Madeira, Caramujo, 1300 m, 4.1V.1993, leg. & coll. Assing &
Wunderle, MHNG.
COMMENTS
An examination of the type material of Geostiba lindrothi Franz and of G.
carli Palm revealed that both taxa are conspecific. (It should be noted that the types of
G. carli deposited in the SMNH carry the handwritten label ‘Sipalia lundbladi Palm’,
apparently an in litteris name; the specimens are without doubt identical with those
mentioned in the original description of G. carli or Sipalia lindrothi, respectively.)
Both Geostiba lindrothi Franz and Sipalia lindrothi Palm were described in 1981 in
the same journal. Since PALM (1981b), as the first revising author, suggested the new
ri
OA 9
Sd dra
Es
| ki 2 ‘i i
di Mi ae
: TNT
Geostiba lindrothi Franz: median lobe in ventral and in lateral view (a, b); apex of paramere
(c); spermathecae (d); antenna (e); 4 tergite (f) and sternite (g) VIII; hind margin of 2 sternite
VII (h); pubescence partly omitted in e-h. Scales a-b, d-h: 0.1 mm; c: 0.05 mm.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 135
name Geostiba carli for his Sipalia lindrothi shortly afterwards and thus implicitly
recognized the priority of G. lindrothi Franz, his G. carli must be considered a
synonym of the former.
DESCRIPTION
Since this and the following four species of blind small subterraneous Geostiba
can only be distinguished on the basis of the sclerites of abdominal segment VII and
the genitalia, only the first will be fully described. The descriptions of the remainder
will focus on differential characters.
Measurements: HW: 250-275, 259+6; PW: 285-310, 301+7; PL: 275-305,
327+10; EL: 180-215, 194+7; SL: 710-830, 777£29; TL: 1.9-2.4, 2.18+0.16.
Colour of body including appendages entirely yellowish with only the antero-
central area of tergites III-VI occasionally with a slightly darker hue.
Body even smaller and, on the whole, slenderer than in G. portosantoi.
Head narrower than pronotum (0.83-0.88x), but relatively wider than in the G.
portosantoi. Surface somewhat shiny, but with distinct, yet variable microreticulation
and mostly rather fine, sparse punctation; eyes rudimentary and without ommatidia,
their maximal diameter ca. 27-30 um; antennae about as long as head and pronotum
together (Fig. 9e).
Pronotum small, as wide as or slightly wider than long (1.0-1.1x) with
maximal width in anterior half; microreticulation similar to that of head, punctation
very fine and sometimes indistinct; pubescence short and inconspicuous.
Elytra much shorter (0.63-0.70x) than and about as wide as pronotum;
microsculpture variable, reticulate, often superficial; punctation granulose and sparse;
alae reduced.
Abdomen with maximal width slightly exceeding that of elytra; dorsal surface
with isodiametric microreticulation similar to that of elytra and with fine and sparse
punctation.
3: hind margin of tergite VIII straight to weakly rounded (Fig. 9f), that of
sternite VIII obtusely pointed (Fig. 9g); median lobe as in Figs. 9a—b; apex of
paramere with one long and three short setae (Fig. 9c).
2: hind margin of tergite VIII weakly convex, sternite VIII posteriorly with
weak concavity (Fig. 9h); spermatheca as in Fig. 9d.
DISTRIBUTION AND BIONOMICS
So far G. lindrothi is only known from the localities indicated above. FRANZ
(1981) sieved the types from Laurus trunks. We extracted our specimens from soil in
a stand of old Erica sp. in northern exposition. Two of the females had mature eggs in
their abdomen.
Geostiba graminicola spec. nov. (Figs. 10a-h)
Types: Holotype: 4, Madeira, Pico Arieiro, 1750 m, 26.11.1993, leg. & coll. Wunderle.
Paratypes: 24 6d, 22 9 9: Madeira, Pico Arieiro, 1750 m, 26.III.1993, leg. & coll.
Assing & Wunderle, MHNG.
136 VOLKER ASSING & PAUL WUNDERLE
DESCRIPTION
Measurements: HW: 270-290, 276+5; PW: 300-350, 320+8; PL: 285-320,
302+8; EL: 190-210, 197+6; SL: 740-910, 809+45; TL: 2.0-2.6, 2.25+0.15.
In colour and external morphology highly similar to G. lindrothi; the antennae
are less slender with segments 4-6 clearly more transverse than in the preceding
species (Fig. 10e); rudiments of eyes variable in size, often even smaller than in the
preceding species (maximal diameter 20-27 um).
d : hind margin of tergite and sternite VIII similar to G. lindrothi (Figs. 10f-g);
median lobe as in Figs. 10a—b; apex of paramere with two long and two short setae
(Fig. 10c)
?: hind margin of tergite VIII + straight, that of sternite with pronounced
concavity (Fig. 10h); spermatheca as in Fig. 10d.
DISTRIBUTION AND BIONOMICS
G. graminicola has only been recorded from the type locality, where it was
collected in the shade of a big rock. A few specimens were found under stones, the
majority, however, was sieved from soil and the roots of the grass vegetation (name!).
The abdomen of one of the 2 2 contained a mature egg.
Fic. 10a-h.
Geostiba graminicola spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermatheca (d); antenna (e): ¢ tergite (f) and sternite (g) VIII; hind margin of 9
sternite VIII (h); pubescence partly omitted in e-h. Scales: a—b, d—h: 0.1 mm; c: 0.05 mm.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 137
Geostiba vaccinicola spec. nov. (Figs. 1 la—h)
Types: Holotype: 6, Madeira, Pico Arieiro, 1600 m, 3.1V.1993, leg. & coll. Wunderle.
Paratypes: 5 d d, 6 £ 2: Madeira, Pico Arieiro, 1600 m, 3.1V.1993, leg. & coll. Assing
& Wunderle, MHNG; 2 d d, 1 9: Madeira, Pico Arieiro, 1600 m, 26.III.1993, leg. & coll.
Assing & Wunderle; 2 4 Sd: Madeira, Pico Arieiro, Fonte Vermelha, 16.1V.1970, leg. & coll.
Franz.
DESCRIPTION
Measurements: HW: 265-290; PW: 300-335; PL: 285-330; EL: 190-210; SL:
770-880; TL: 2.1-2.5.
Colour and external morphology as in G. lindrothi.
d: hind margin of tergite and sternite VIII similar to G. lindrothi (Figs. 11f-g);
median lobe slenderer in ventral view and larger (Figs. 11a-b); apex of paramere with
one long and three short setae (Fig. 11c).
9: hind margin of sternite VIII similar to G. lindrothi (Fig. 11h); spermatheca
stouter and larger than in G. lindrothi (Fig. 11d).
Fic. 11a-h.
Geostiba vaccinicola spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermathecae (d); antenna (e); d tergite (f) and sternite (g) VIII; hind margin of
2 sternite VIII (h); pubescence partly omitted in e-h. Scales a-b, d-h: 0.1 mm; c: 0.05 mm.
138 VOLKER ASSING & PAUL WUNDERLE
DISTRIBUTION AND BIONOMICS
The type specimens of this subterraneous blind species were sieved from deep
litter and extracted from soil in stands of Vaccinium padifolium (name!) and Erica sp.
in northern exposition below the Pico Arieiro.
Geostiba lauricola spec. nov. (Figs. 12a-d)
Types: Holotype: & , Madeira, Rib. da Janela, 800 m, 31.11.1993, leg. & coll. Wunderle.
Paratypes: 5 d d, 2 9 9: Madeira, Rib. da Janela, 800 m, 31.11.1993, leg. & coll.
Assing & Wunderle, MHNG.
DESCRIPTION
Measurements: HW: 260-280; PW: 300-325; PL: 285-310; EL: 200-215; SL:
770-850; TL: 2.0-2.3.
Colour and external morphology as in G. lindrothi.
d: hind margin of tergite and sternite VIII as in G. lindrothi; median lobe
slenderer (Figs 12a-b); apex of paramere long and slender, with three short and one
long setae, the latter situated nearer to base than in G. lindrothi (Fig. 12c).
e,
Fic. 12a-d.
Geostiba lauricola Spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermathecae (d): Scales: a-b, d: 0.1 mm; c: 0.05 mm.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 139
2: hind margin of sternite VIII as in G. lindrothi, spermatheca slightly larger
and stouter than in G. lindrothi (Fig. 12d).
DISTRIBUTION AND BIONOMICS
The type specimens were extracted from soil and deep litter layers in a Laurus
stand (name!) on the ascent from Ribeira da Janela to Fanal in the northwest of
Madeira.
Geostiba caligicola spec. nov. (Figs 13a—h)
Types: Holotype: ¢, Madeira, Ach. do Teixeira, 1600 m, 6.1V.1993, leg. & coll.
Wunderle.
Paratypes: 3 dd, 4 9 9: Madeira, Ach. do Teixeira, 1600 m, 6.1V.1993, leg. & coll.
Assing & Wunderle, MHNG.
DESCRIPTION
Measurements: HW: 255-265; PW: 285-295; PL: 270-290; EL: 170-185; SL:
120-775:
Fic. 13a—h.
Geostiba caligicola spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); spermathecae (d); antenna (e); d tergite (f) and sternite (g) VIII; hind margin of
? sternite VIII (h); pubescence partly omitted in e-h. Scales: a—b, d-h: 0.1 mm; c: 0.05 mm.
140 VOLKER ASSING & PAUL WUNDERLE
Colour and general external morphology as in G. lindrothi; antennae similar to
G. graminicola, but shorter with the first three antennomeres less elongate (Fig. 13e).
d: hind margin of tergite and sternite VIII as in G. lindrothi (Figs 13f-g);
median lobe in lateral view apically slightly slenderer than in G. graminicola (Figs
13a-b); apex of paramere with setal pattern similar to G. graminicola, but the apical
long seta usually longer than in that species (Fig. 13c).
9: hind margin of sternite VIII as in G. lindrothi (Fig. 13h); spermatheca as in
Fig. 13d.
DISTRIBUTION AND BIONOMICS
The type series was sieved below the Pico Ruivo near Achada do Teixeira
from moss and litter in the shadow (name!) of rocks.
Geostiba occulta spec. nov. (Figs. 14a-i)
Types: Holotype: 4, Madeira, Rib. da Janela, 800 m, 31.11.1993, leg. & coll. Assing.
Paratypes: 2 6d, 8 2 2: Madeira, Rib. da Janela, 800 m, 31.11.1993, leg. & coll.
Assing & Wunderle, MHNG.
DESCRIPTION
Measurements: HW 405-430; PW 480-530; PL: 480-530; EL: 350-380; SL:
1330-1404; TL: 3.4-3.9.
G. occulta is the largest of all blind species of Madeiran Geostiba.
Colour of body including appendages entirely yellowish or reddish yellow with
only the antero-central area of tergites HI-VI sometimes with a slightly darker hue.
Head narrower than pronotum (0.81-0.84x); surface with distinct micro-
reticulation and mostly rather fine, sparse punctation; eyes reduced to minute rudi-
ments, without ommatidia; antennae slender, longer than head and pronotum together
(Fig. 14e).
Pronotum as wide as long (0.99-1.03x) with maximal width in anterior half,
microreticulation similar to that of head, punctation even finer than on head; pubes-
cence short and inconspicuous.
Elytra much shorter (0.69-0.75x) and a little wider than pronotum;
microsculpture reticulate, weaker than on head and pronotum, surface therefore more
shiny; punctation granulose and fine; alae reduced.
Abdomen with maximal width clearly exceeding that of elytra; dorsal surface
with distinct isodiametric microreticulation and sparse and fine punctation.
d: hind margin of tergite VIII deeply concave; sternite VIII rounded pos-
teriorly (Figs. 14f-g); median lobe and paramere as in Figs. 14a-c.
9: hind margins of tergite and sternite VIII shallowly concave centrally (Figs.
14h-1); spermatheca as in Fig. 14d.
DISTRIBUTION AND BIONOMICS
The type specimens were found under the same conditions and in the same
locality as G. lauricola.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 141
rem
aL
Fic. 14a-i
Geostiba occulta spec. nov.: median lobe in ventral and in lateral view (a, b); apex of paramere
(c); spermatheca (d); antenna (e); d tergite (f) and sternite (g) VIII; © tergite and sternite VIII
(h-i); pubescence partly omitted in ei. Scales: a-b, d, i: 0.1 mm; e-h: 0.2 mm; c: 0.05 mm.
Geostiba endogea spec. nov. (Figs. 15a-i)
Types: Holotype: d, Madeira, Caramujo, 1220 m, 29.IIT.1993, leg. & coll. Wunderle.
Paratypes: 3 dd, 4 2 2: Madeira, Caramujo, 1220 m, 29.III.1993, leg. & coll. Assing &
Wunderle, MHNG; 1 2: Madeira, Rosario, 800 m, 5.1V.1993, leg. & coll. Assing & Wunderle.
142 VOLKER ASSING & PAUL WUNDERLE
DESCRIPTION
Measurements: HW: 345-365, 353+7; PW: 405-455, 429+13; PL: 375-425,
404+13; EL: 290-325, 310+10; SL: 1090-1220, 1159+44; TL: 3.0-3.3.
In general appearance G. endogea resembles G. occulta, but it is censiderably
smaller than the latter; on the other hand it clearly exceeds the species of the G.
lindrothi group in body size.
Colour of body including appendages entirely yellowish or reddish yellow
with only the antero-central area of tergites III-VI sometimes with a slightly darker
hue.
Head narrower than pronotum (0.80-0.85x); surface somewhat shiny, with
apparent, yet superficial microreticulation and very fine, sparse punctation; eyes
reduced to minute rudiments; antennae less slender than in G. occulta, a little shorter
than head and pronotum together (Fig. 15e).
Pronotum slightly wider than long (1.05—1.10x) with maximal width near the
middle; microreticulation and punctation similar to that of head; pubescence short and
inconspicuous.
Elytra clearly shorter (0.74-0.81x) and a little wider than pronotum; micro-
reticulation rather superficial, with larger meshes than on head and pronotum;
punctation granulose and fine; alae reduced.
Abdomen with maximal width clearly exceeding that of elytra; dorsal surface
with distinct isodiametric microreticulation similar to that of elytra and sparse and
fine punctation.
d: hind margin of tergite VIII + straight to slightly convex, that of sternite
VIII convex (Figs. 15f-g); median lobe and paramere as in Figs. 15a-c.
9: hind margin of tergite VIII slightly convex, that of sternite VIII with
distinct concavity, centrally with rather many (7-9) long hairs clearly exceeding the
row of bristles in length (Figs. 15h-i); spermatheca as in Fig. 15d.
DISTRIBUTION AND BIONOMICS
At the localities indicated above the type material was sieved and extracted
from soil and deep litter layers, under moss and tree trunks in stands of deciduous
trees with northern exposition.
Geostiba subterranea spec. nov. (Figs. 16a-f)
Types: Holotype: 4, Madeira, Ribeiro Bonito, leg. & coll. H. Franz.
Paratype: 1 d: Madeira, Ribeiro Bonito, leg. H. Franz, coll. Wunderle.
DESCRIPTION
Measurements of the two specimens: HW: 362, 378; PW: 453, 498; PL: 438;
BESS Sie 21651223;
In general appearance rather similar to G. endogea.
Colour of body including appendages entirely yellowish or reddish yellow with
only the antero-central area of tergites III-VI sometimes with a slightly darker hue.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 143
us LI
fe
rm m ft ee
i
Fic. 15a-i
Geostiba endogea spec. nov.: median lobe in ventral and in lateral view (a, b); apex of paramere
(ce); spermatheca (d); antenna (e); d tergite (f) and sternite (g) VIII; © tergite and sternite VIII
(hi); pubescence partly omitted in e-i. Scales: a-b, d, i: 0.1 mm; e-h: 0.2 mm; c: 0.05 mm.
144 VOLKER ASSING & PAUL WUNDERLE
Head narrower than pronotum (0.75-0.80x), with distinct microreticulation,
punctation sparse and fine; eyes reduced to minute rudiments; antennae slenderer than
in G. endogea With segments 5-10 weakly transverse, longer than head and pronotum
together (Fig. 16d).
Pronotum a little wider than long (1.05-1.15x) with maximal width in anterior
half; microreticulation similar to that of head, punctation fine and sparse; pubescence
short and inconspicuous.
Elytra much shorter (0.7x) and a little wider than pronotum; microsculpture
less distinct than on head and pronotum; punctation granulose and fine; alae reduced.
Abdomen with maximal width clearly exceeding that of elytra; dorsal surface
with distinct microreticulation and fine and sparse punctation.
d: hind margin of tergite VIII weakly convex, sternite VIII rounded pos-
teriorly (Figs. 16e-f); median lobe and paramere as in Figs. 16a-c.
9: unknown.
DISTRIBUTION AND BIONOMICS
At present this species is only known from the type locality. Unfortunately, the
labels lacked any information as regards the ecological circumstances.
SPECIES EXCLUDED
Atheta leileri (PALM, 1981) comb. nov.
Sipalia leileri PALM, 1981a: 294ff.
Atheta (Parameotica) juengeri BENICK 1984: 166ff. syn. nov.
Types: Holotype; d, Madeira, Pico Arieiro, 23.1V.1975, Holotypus, Sipalia leileri
Palm, det. Thure Palm (MZEL).
Paratypes: 2 2 2: same data as holotype (MZEL).
FURTHER MATERIAL STUDIED: 21 dd, 18 22: Madeira, Pico Arieiro, 1600 m,
26.111.1993, leg. & coll. Assing & Wunderle; 8 dd, 13 9 9: Bica da Cana, 1550-1600 m,
29 111.1993, leg. & coll. Assing & Wunderle; 8 dd, 4 2 2: Madeira, Pico Arieiro, 1600 m,
3.1V.1993, leg. & coll. Assing & Wunderle.
PALM (1981a) erroneously believed this species to be a Sipalia or Geostiba,
respectively, apparently because of the general appearance, the small eyes and the
rather short elytra. However, it does not possess any of the further typical characters
of Geostiba. The construction of the genitalia is completely different. Furthermore,
the pubescence of the pronotum is not directed caudad, but + laterally on the sides;
along midline it is directed cephalad in the anterior half and caudad in the posterior
half (type III in BENICK & LOHSE (1974)). Having examined various characters
including the mouthparts and the genitalia we conclude that S. leileri Palm has to be
transferred to Atheta Thoms. For a detailed diagnosis and illustrations of the genitalia
(including the internal sac of the aedeagus) see BENICK (1984), who described the
same species again as Atheta juengeri, now a junior synonym of S. leileri Palm. J.
Vogel, Gürlitz, who studied the types of A. juengeri Benick, kindly confirmed that
they were indeed conspecific with our material of leileri (pers. comm.).
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 145
Pe
Fic. 16a-f
Geostiba subterranea spec. nov.: median lobe in ventral and in lateral view (a, b); apex of
paramere (c); antenna (d); d tergite (e) and sternite (f) VIII; pubescence partly omitted in d-f.
Scales: a—b: 0.1 mm; c: 0.05 mm; d-f: 0.2 mm.
DISTRIBUTION AND BIONOMICS
A. leileri (Palm) has repeatedly been collected at or near the type locality (Pico
Arieiro) at an altitude of ca. 1600 m (BENICK 1984; PALM 1981a; our own records, see
above). We sieved numerous specimens from litter in stands of Vaccinium and Erica
both in northern and southern exposition, also at Bica da Cana at 1600 m. One d
taken on 29.III.1993 was teneral
146
VOLKER ASSING & PAUL WUNDERLE
KEY TO THE MADEIRAN SPECIES OF Geostiba Thomson
D
Eyes functioning, with 4 ormore ommatidia "EN RE PEER ER 2
Eyes reduced to minute rudiments, without ommatidia; inhabitants of
Madétrapropert. 0420 Wr. TR RIRES 8
Larger species, 3.0-3.6 mm, PW >430 um, PL >400 um: eyes with 6 or
more ommatidia. d : apex of paramere with two or three long setae........ 9
Smaller species, 1.8-2.8 mm, PW >430 um, PL >380 um; eyes with
4-Sfomimatidia:») Muri, (eb EEE RIENTRO 4
Shape of head almost circular, HW >370 um, abdomen distincio
widened posteriorly, pubescence of head and pronotum longer and
more conspicuous, antennae slenderer with segments 6-10 only weakly
transverse (Fig. 11), elytra relatively shorter.
d: without longitudinal elevation on elytra, pronotum without im-
pression, punctation of tergites VII and VIII fine, median lobe and apex
of paramere as in Figs. la—c.
9: hind margin of sternite VIII concave (Fig. Ih), spermatheca as in
Fig. Id.
iad Sina DO DE tees cic ee a II G. formicarum (Woll.)
Head slenderer, HW <370 um, abdomen only slightly widened pos-
teriorly, pubescence of head and pronotum short, less dense and
inconspicous, antennae shorter with segments 5-10 distinctly transverse
(Fig. 21), elytra relatively longer.
d: with longitudinal elevation on elytra near suture, pronotum with
longitudinal impression on disk, tergites VII and VIII with pronounced
granulose punctation, median lobe and apex of paramere as in Figs.
2a-c.
© : hind margin of sternite VIII less concave (Fig. 2h), spermatheca as
in Fig. 2d.
Madeita,proper. a NIRO: AMORE E er ee G. filiformis (Woll.)
Body colour + uniformly yellowish to reddish yellow, head narrower
with HW <290 um; d : apex of paramere with one long and three shorter
setae? Species of Porto Santo. . . ...... 2... = u te eee 5
Body + bicoloured, yellowish with at least tergites V and VI somewhat
darkened, head wider with HW >310 um: d: apex of paramere with
four short setae. For a safe identification of these species an exami-
nation of the aedeagus and the ¢ tergite VIII is essential. Species of
Madeira proper. - ........... bic 2... ee TETTE 6
Slightly smaller species; nal shorter, about as long as wide (Fig. 8b),
pronotum on the average shorter and narrower.
d : median lobe smaller, with spines in internal sac (Figs. 7a-b); apex
of paramere as in Fig. 7c.
2: spermatheca smaller and of different shape (Fig 7d). Pico Branco
(BOrlO Santo). os Se RR G. brancomontis spec. nov.
10.
IN
2.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 147
Somewhat larger species; head longer (ca. 1.15x) than wide (Fig. 8a),
pronotum on the average longer and wider.
d: median lobe larger, without spines in internal sac (Figs. 6a—b); apex
of paramere as in Fig. 6c.
2: spermatheca larger (Fig. 6d).
Biconulianag(Porto?Santo)as re E G. portosantoi Franz
6 : median lobe with pronounced ventral process. ...................... 7
3d: median lobe without ventral process (Fig. Sb); tergite VIII weakly
concave posteriorly (Fig. Se); pronotum without impression.
Seas petmathecaas: mR 9 RR er G. bicacanaensis spec. nov.
Abdomen mostly with only tergites V and VI darkened.
d: pronotum often with longitudinal impression on disk; tergite VIII
with pronounced concavity posteriorly (Fig. 3f); ventral process of
median lobe bent (Fig. 3b), apex of paramere as in Fig. 3c.
CÉSpenmathecarasun Elo ad van E G. arieiroensis spec. nov.
Abdomen mostly with tergite VII as dark as tergites V and VI.
d: pronotum without longitudinal impression on disk; tergite VIII
weakly concave posteriorly (Fig. 4e); ventral process of median lobe +
straight (Fig. 4b), apex of paramere as in Fig. 4c.
Deispermatheeaasan Bis. Adnan. G. ruivomontis spec. nov.
PATS CIES CCIE See 3:0 3:9: mm, Mr ER NS Lin OL 9
SmalesSpecies 18022 GENE Sr nest er: Il
Body length 3.0-3.3 mm; HW <390 um, PL <460 um; body appen-
HASESIS NON ER APR ROLLE Ie e e 10
Larger species, 3.4-3.9 mm; HW >400 um, PL >490 um; legs and
antennae (Fig. 14e) long and slender.
d: hind margin of tergite VIII deeply concave (Fig. 14f), median lobe
and apex of paramere as in Figs. 14a-c.
ÉHSpermathecaæas initie Ad nn ca ec ak G. occulta spec. nov.
Antennae shorter with segments 4—10 transverse (Fig. 15e).
3: median lobe smaller (Figs. 15a-b); apex of paramere as in Fig. 15c.
Sraspemmatneca asim Kies sd... G. endogea spec. nov.
Antennae slenderer with segments 5—10 weakly transverse (Fig. 16d).
3: median lobe larger and of different shape (Figs. 16a—b); apex of
paramere as in Figs. 16c.
AUTONET n a AEs Bye ee Le RS G. subterranea spec. nov.
d : apex of paramere with one long and three very short setae (see e.g.
Eller, DOS RS OT SRE RER ER 12
2: apex of paramere with two long and two very short setae (e.g. Fig.
[LOTTE ee D SR 14
3: median lobe larger, ca. 0.3 mm long (Fig. 11a-b); apex of paramere
as in Fig. 11c.
ÉSpDermathe cafas n bie Md een G. vaccinicola spec. nov.
Jamedianiobe;smallerca 025 nm NR 15)
148 VOLKER ASSING & PAUL WUNDERLE
13. d: median lobe slenderer (Fig. 12a); apex of paramere more elongate,
long seta situated nearer to base (Fig. 12c).
9: spermatheca slightly larger and stouter (Fig. 12d). . G. lauricola spec. nov.
- d: median lobe wider in ventral view (Fig. 9a); apex of paramere less
elongate, long seta more distant from base (Fig. 9c).
2: spermatheca slightly smaller (Fig. 9d)............... G. lindrothi Franz
14. Antennae longer with the first three antennomeres more elongate (Fig.
10e).
d: median lobe in lateral view as in Fig. 10b; apical long seta usually
shorter (Fig. 10c).
2: posterior margin of sternite VIII with pronounced angular concavity
(Fig. 10h); spermatheca as in Fig. 10d............ G. graminicola spec. nov.
_ Antennae shorter (Fig. 13e).
d : median lobe in lateral view of different shape (Fig. 13b); apical long
seta usually longer (Fig. 13c).
9 : posterior margin of sternite VIII only shallowly concave (Fig. 13h);
spenmatheca' asile lE E G. caligicola spec. nov.
DISCUSSION
At present, 15 endemic species of Geostiba are known to occur on the Madeiran
islands. Thus speciation through adaptive radiation appears to have developed far more
endemics in this than in any other staphylinid genus on the archipelago. Of the better
known taxa only Stenus Latr., Othius Steph. (both 5 endemics) and Mycetoporus
Mannh. (3 endemics) include an appreciable number of species whose distribution is
restricted to Madeira (ASSING & WUNDERLE, 1994, 1995; PALM 1980). The genus
Atheta Thoms., particularly the subgenus Mocyta, can be expected to comprise a
considerable number of endemics, too, but in the absence of a recent revision precise
figures are impossible.
Furthermore, the number of Madeiran species of Geostiba considerably
exceeds that of other Atlantic Islands. Several species have been described from the
Canary Islands (FRANZ 1981; PALM 1975, 1976; WOLLASTON 1864), but according to
our own studies only two of them are true Geostiba: G. muscicola (Woll.) and G.
lanzarotensis (Palm). One species is reported for the Azores, G. melanocephala
(Crotch) (BERNHAUER 1940; CROTCH 1867), whose generic identity was confirmed
through an examination of two syntypes in coll. Wollaston (BMNH).
It may seem surprising that in spite of a fairly long tradition of entomofaunistic
activity on the Madeiran islands, so few species of Geostiba were described before;
many studies list no or, at the most; two species of the genus (e.g. BERNHAUER 1940;
ERBER 1990; ERBER & HINTERSEHER 1988, 1990; JANSSON 1940; LIKOVSKY 1963),
probably because the methods used were inadequate for these more or less
subterranean insects. Considering that during a two-week excursion it was possible to
collect not only the Geostiba species known at that time, but in addition an even
greater number of new taxa, most of them from one locality only, it seems certain that
a number of species remain to be discovered in the future.
MADEIRAN SPECIES OF THE GENUS GEOSTIBA 149
Regarding their morphological and ecological properties the Madeiran
Geostiba can be subdivided into 5 species groups. Three of them possess functioning
eyes: the G. formicarum group (G. formicarum, G. filiformis), the G. arieiroensis
group (G. arieiroensis, G. ruivomontis, G. bicacanaensis) and the G. portosantoi
group (G. portosantoi, G. brancomontis). The remaining two groups are blind and
subterraneous: the G. lindrothi group (containing the minute species G. lindrothi, G.
graminicola, G. vaccinicola, G. lauricola, G. caligicola) and the G. occulta group
(with the larger G. occulta, G. endogea, G. subterranea). There are, at first glance,
considerable differences between these groups regarding characters such as body size,
development of eyes, etc., particularly when measured against the relative uniformity
of continental Geostiba. However, having examined numerous external characters as
well as the mouthparts and the genitalia (also in comparison with the type species G.
circellaris (Grav.)), we are convinced that the species dealt with in this study are all
true members of Geostiba. After all, morphological (and ecological) diversity within
groups of closely related species is a well-known island phenomenon, exemplified not
only in Darwin's finches, but also many other animal taxa.
On the other hand the species of the G. lindrothi group are of remarkable
similarity. Nevertheless, we recognize them as different species for several reasons.
Above all, constant differences in the genitalia and other sex-related characters suggest
reproductive isolation. Moreover, most species of Geostiba in general cannot be safely
identified on the basis of external characters alone (cf. PACE 1977). A low morpho-
logical diversity should particularly be expected in the species of the G. lindrothi group,
all of them living subterraneously and representing similar life forms; selective pressure
in such habitats will eventually result in similar morphological adaptations, especially
various reductions (of pigmentation, eyes, size, etc.), which unfortunately also reduce
the number of available diagnostic features for the taxonomist.
ACKNOWLEDGEMENTS
We would like to express our sincere thanks to Dr. Roy Danielsson (MZEL),
Dr. Tord Nyholm and Dr. Per Lindskog (SMNH), and to Prof. Herbert Franz,
Mödling, for loans of type material. We are furthermore indebted to Mr Peter M.
Hammond and Ms Emma De Boise (BMNH), who arranged the loan of Wollaston's
types and who made it possible for the first author to study Wollaston's collection of
Madeiran Staphylinidae in the BMNH. Finally, we would like to thank Dr. Ivan Löbl
(MHNG) for his valuable advice and critical comments on the manuscript.
REFERENCES
ASSING, V. & P. WUNDERLE, 1994. The fifth endemic Stenus from Madeira: Stenus (Tesnus)
ruivomontis spec. nov. (Coleoptera, Staphylinidae). Bocagiana, Museu Municipal do
Funchal No. 175: 1-4.
ASSING, V. & P. WUNDERLE, 1995. A revision of the Madeiran species of the genus Othius
Stephens (Coleoptera: Staphylinidae). Boletim do Museu Municipal do Funchal 45
(1993): 53-65.
150 VOLKER ASSING & PAUL WUNDERLE
BENICK, G., 1984. Eine neue Afheta: A. (Parameotica) juengeri sp. n. von Madeira (Col.,
Staph.). Entomologische Blätter für Biologie und Systematik der Käfer 80: 166-168.
BENICK, G. & G.A. LOHSE 1974. Tribus 14 (Callicerini). /n: Die Käfer Mitteleuropas, Bd. 5,
Staphylinidae II (Hypocyphtinae und Aleocharinae, Pselaphidae), (Freude, H., Harde,
K.W. & G.A. Lohse) Krefeld: 72-220.
BERNHAUER M., 1940. Staphyliniden von den Azoren und Madeira. Commentationes Biolo-
gicae 8(2): 1-10.
BLACKWELDER, R.E., 1952. The generic names of the beetle family Staphylinidae with an essay
on genotypy. Smithsonian Institution, United States National Museum Bulletin 200:
1-483
CROTCH, G.R., 1867. On the Coleoptera of the Azores. Proceedings of the Zoological Society
London 1867: 359-391.
ERBER, D., 1990. New and little known Coleoptera from Madeira. Results of excursions to
Madeira in the years 1986-1990. Boletim do Museo Municipal do Funchal 42:
147-181.
ERBER, D. & W. HINTERSEHER, 1988. Contribution to the knowledge of the Madeira beetles.
Boletim do Museo Municipal do Funchal 40: 139-214.
ERBER, D. & W. HINTERSEHER, 1990. Additional notes to the knowledge of the Madeira beetles.
Boletim do Museo Municipal do Funchal 42: 141-146.
FAUVEL, A., 1897. Catalogue des Staphylinides de Barbarie et des Iles Acores, Madères,
Salvages et Canaries. Revue d’Entomologie 16: 237-371.
FRANZ, H. 1981. Neue blinde, subterrane Coleopteren von den Makaronesischen Inseln (Cole-
optera: Staphylinidae et Curculionidae). Entomologica scandinavica Supplementum 15:
328-332.
JANSSON, A., 1940. Die Arthropodenfauna von Madeira nach den Ergebnissen der Reise von
Prof. Dr. O. Lundblad Juli-August 1935. XXIX. Coleoptera. Arkiv för Zoologi B. 32 A.
No 24: 1-64. Stockholm.
LIKOVSKY, Z., 1963. A contribution to the study of beetles in the Madeira Islands. Results of
expeditions in 1957 and 1959. V. Staphylinidae II. (Unterfamilie Aleocharinae). Com-
mentationes Biologicae 25: 44-50.
PACE, R., 1977. Studio sul genere Geostiba Thomson, con descrizione di nuove specie italiane
(Coleoptera, Staphylinidae). Fragmenta Entomologica (Roma) 13: 183-229.
PALM, T., 1975. Zur Kenntnis der Käferfauna der Kanarischen Inseln. 4-7. 5. Die Gattung
Sipalia Muls. et Rey (Col. Staphylinidae). Entomologisk Tidskrift 95: 37-50.
PALM, T., 1976. Zur Kenntnis der Käferfauna der Kanarischen Inseln 15-16. 15. Sipalia franzi
n. sp. von Teneriffa (Coleoptera: Staphylinidae). Entomologica scandinavica 7: 37-50.
PALM, T., 1980. The beetle fauna of Madeira 2. — The genus Mycetoporus Mannh. (Coleoptera:
Staphylinidae). Entomologica scandinavica 11: 393-398.
PALM, T., 1981a. Zur Kenntnis der Käferfauna von Madeira 3. Die Gattung Sipalia Muls. &
Rey (Coleoptera: Staphylinidae). Entomologica scandinavica Supplementum 15: 293-
298.
PALM, T., 1981b. Geostiba carli n. nov. for Sipalia lindrothi Palm, 1981 (Coleoptera: Staphy-
linidae). Entomologica scandinavica 12: 447.
WOLLASTON, T.V., 1854. Insecta Maderensia, being an Account of the Insects of the Islands of
the Madeiran Group. London: 634 pp.
WOLLASTON, T.V., 1857. Catalogue of the Coleopterous Insects of Madeira in the collection of
the British Museum. London: 234 pp.
WOLLASTON, T.V., 1865. Coleoptera Atlantidum, being an enumeration of the Coleopterous
Insects of the Madeiras, Salvages, and the Canaries. London: 526 pp. + Appendix.
REVUE SUISSE DE ZOOLOGIE, 103 (1) : 151-193; mars 1996
The millipede family Paradoxosomatidae on Borneo,
with contributions to the faunas of some other islands
of the Sunda area (Diplopoda, Polydesmida)
Sergei I. GOLOVATCH
Institute for Problems of Ecology and Evolution, Russian Academy of Sciences,
Leninsky prospekt 33, 117071 Moscow (V-71), Russia.
The millipede family Paradoxosomatidae on Borneo, with contri-
butions to the faunas of some other islands of the Sunda area (Diplo-
poda, Polydesmida). — Several fresh collections of paradoxosomatid
millipedes from the Indo-Australian archipelago, mainly Borneo, comprise
14 species and two genera new to science: Euphyodesmus reductus sp. n.,
E. perarmatus sp. n., E. lobatus sp. n., E. lateridens sp. n., Borneochiropus
spinitergus Sp. n. (both genera allocated within the chiefly Australian tribe
Antichiropodini), Orangutana setosa gen. n., sp. n. (Orthomorphini),
Arthrogonopus proletarius sp. n., Opisthodolichopus bifidus sp. n., Mar-
garitosoma borneorum sp. n., Dajakina longipes sp. n., Kalimantanina
crucifera sp. n., K. bruneica sp. n., all from Borneo, as well as Gigan-
tomorpha mutilata sp. n., from Bali, and Margaritosoma singaporense Sp.
n., from Singapore. Notes on the variation range of Gigantomorpha imma-
nis Jeekel, 1963, are given. Tectoporus filarius (Attems, 1932), has been
rediscovered on Java, Opistodolichopus scandens Hoffman, 1973, in Singa-
pore, and Tectoporus pygmaeus (Pocock, 1894), hitherto reported solely
from Java, has been recorded in Bali for the first time. Keys to the tribes
and genera of Paradoxosomatidae encountered on Borneo as well as to
species of the genera Arthrogonopus Jeekel, 1963, Dajakina Jeekel, 1963,
Euphyodesmus Attems, 1931, Gigantomorpha Jeekel, 1963, Margari-
tosoma Jeekel, 1979, Kalimantanina Jeekel, 1963, and Opisthodolichopus
Verhoeff, 1941, have been compiled. A review of the Bornean
paradoxosomatid fauna is presented coupled with brief zoogeographical
comments.
Key-words: Paradexosomatidae - fauna - taxonomy - keys - Borneo -
Sunda Archipelago.
Manuscript accepted 15.09.1995.
152 SERGE! I. GOLOVATCH
INTRODUCTION
The millipede fauna of Southeast Asia, both continental and insular, is strongly
dominated by species of the large family Paradoxosomatidae. Yet the fauna of
Borneo, one of the biggest islands in Australasia, remains highly insufficiently
known. Only very few papers have been specially or mainly devoted to Bornean
paradoxosomatids, all by JEEKEL (1963, 1983a, 1983b), with the result that a total of
30 acknowledged species and nine genera have hitherto been reported from that great
island. I guess that actually several times as many species await discovery on Borneo.
The present paper focuses on part of a considerable millipede collection from
Borneo managed recently during several trips by Dr. Daniel Burckhardt, Dr. Bernd
Hauser and Dr. Charles Lienhard, all from the Muséum d'Histoire naturelle in Geneva
(MHNG). This contribution is restricted to the Paradoxosomatidae, hence providing
an excellent opportunity to review and key the entire Bornean fauna of that family as
well as to briefly discuss some zoogeographical issues. In addition, several sporadic
samples taken by the above collectors on a few other islands of the Sunda area on
their way to and back from Borneo have been incorporated in this study as well.
Besides, one more Bornean specimen representing a new species and recently
collected by Dr. Ott6 Merkl, of the Hungarian Natural History Museum in Budapest
(HNMB), has been added here too, bringing the paradoxosomatid fauna of Borneo to
a total of 42 species and 12 genera.
Only a few duplicates have been retained by the author for a subsequent
deposition in the collection of the Zoological Museum of the Moscow State Uni-
versity (ZMUM), whereas the bulk of material has been returned to MHNG or
HNMB, as indicated hereinafter.
TAXONOMIC PART.
Tectoporus filarius (Attems, 1932) (Figs 1-5)
Material: 1 4,2 2 (MHNG), 1 d, 1 2 (ZMUM), Indonesia, Java: Cibodas, Lithocar-
pus-Castanopsis forest along ravine above Botanical Gardens, 1350-1480 m, beaten from
vegetation, 25.XI.1987 (Sar-87/16). — 1 3, 1 2 (MHNG), same locality, lower part of Bota-
nical Gardens, ca. 1250-1300 m, beaten from vegetation, 27.X1.1987 (Sar-87/28), all leg. C..
Lienhard.
Remarks: The above material corresponds virtually fully to the original
description (ATTEMS 1932) and the only subsequent record (CHAMBERLIN 1945), as
seen from Figs 1-5. The only minor discrepancy concerns the presence both of very
poorly developed tubercles supporting the caudal row of tergal setae and a short axial
line behind the transverse sulcus (Fig. 1). However, the above variation in metatergal
sculpture can fairly well be considered as individual or populational, for the gonopod
structure is identical.
This species has hitherto been known only from West Java (see review: JEEKEL
1951), and the above material actually derives from the locus typicus: Cibodas (=
Tjibodas).
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 153
Tectoporus filarius (Attems, 1932), d: 1 — somite 10, dorsal view; 2 — sternal lamina between d
coxae 4, caudal view; 3 — left gonopod, mesal, lateral, and dorsal views, resp. — Scales in mm.
Tectoporus pygmaeus (Pocock, 1894) (Figs 6-14)
Material: 1 3 (NHMG), Indonesia, Bali: Bedugul, Botanical Gardens, 1230-1260 m,
forest and especially in hothouses, 30.IX.1987 (Sar-87/35); leg. B. Hauser. — 2 4,2 2 (NHMG),
2 d (ZMUM), same locality, forested part, 1230-1260 m, beaten from vegetation, 30.X1.1987
(Sar-87/36); leg. C. Lienhard.
Remarks: Like the preceding species, T. pygmaeus has hitherto been known
only from West Java (see reviews: JEEKEL 1951, 1968, 1979a), being new to the fauna
of Bali. Like the previous case, in spite of minor variations in some peripheral and
even gonopodal characters (POCOCK 1894; ATTEMS 1903, 1937; CHAMBERLIN 1945),
the new illustrations (Figs 6-14) leave no doubt that the above new material belongs
to T. pygmaeus. Moreover, introduction cannot be ruled out, since at least part of
material could have derived from hothouses.
In addition to T. pygmaeus, five paradoxosomatid species have been encoun-
tered on Bali, viz. both anthropochorous Orthomorpha coarctata (de Saussure 1860),
and Chondromorpha xanthotricha (Attems 1898) (JEEKEL 1983b), as well as the
probably endemic Orthomorpha baliorum Golovatch, 1994, Margaritosoma baliorum
Golovatch, 1994 (GOLOVATCH 1994b), and Gigantomorpha mutilata sp. n. (see below).
Euphyodesmus reductus sp. n. (Figs 15-18)
Material: HOLOTYPE d (MHNG), East Malaysia, Borneo, Sabah, West Coast
Residency, Mt Kinabalu, “Bukit Ular Trail”, 1790 m, sifted leaf litter and rotten wood in
Lithocarpus-Castanopsis forest, extracted with Winkler-Moczarski eclector, 28.IV.1982 (Sab-
82/15); leg. B. Hauser.
154 SERGEI I. GOLOVATCH
Fics 6-14
Tectoporus pygmaeus (Pocock, 1894), d : 6 — anterior body portion, lateral view; 7-8 — somite
10, lateral and dorsal views, resp.; 9 — sternal lamina between d coxae 4, caudal view; 10 — leg
3; 11-12 — right gonopod, mesal and lateral views, resp.; 13-14 — distal part of right gonopod,
ventromesal and subfrontal views, resp. — Scales in mm.
Etymology: The species epithet refers to the drastically reduced paraterga and
gonopod telopodite distally of the prefemoral portion.
Diagnosis: Differs from congeners by the especially poorly developed para-
terga and gonopod tibiotarsus as well as by the particularly short antennae and legs.
DESCRIPTION
Length ca. 8 mm, maximal width 0.75 mm on midbody somites and 0.8 mm on
head. Coloration uniform pale brownish, anterior third of body a little brighter.
Antennae only distally, but not apically, brownish. Legs, ventrum and a few basal
antennomeres yellowish-whitish.
Head a bit broader than subequally wide collum and somites 5-17. Postcollar
constriction evident on subequally narrowest somites 2-4. Body distinctly moniliform,
rather rapidly tapering caudad on somites 18-20. Antennae short, quite clavate, in situ
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 155
Fics 15-18
Euphyodesmus reductus sp. n., & holotype: 15 — somite 10, lateral view; 16 — left gonopod,
mesal view; 17-18 — tip of left gonopod, subdorsal and ventromesal views, resp. — Scales in mm.
reaching to end of somite 2. Paraterga almost entirely missing, very weakly displayed
as small and low ridges on somite 2, hardly traceable due to outlining sulci from
above and below on somites 3-4, onward as trace sulci only from above on somites
5-6, further caudad as poor swellings (Fig. 15). Ozopores lying very low. Surface
shining, smooth. Somites deeply constricted, suture dividing pro- and metazona deep,
strongly striate longitudinally. Transverse metatergal sulcus starting from somite 5,
rather deep, thin and lineiform, missing on somite 19. Tergal setae rather short, a few
on collum, 2+2 setae on somite 2, 1+1 presulcus ones on further somites but telson
(Fig. 15). Pleurosternal keels absent. Epiproct moderately long, somewhat flattened
dorsoventrally, in lateral view almost straight, from above subtriangular, rather
broadly truncate. Subanal scale subtrapeziform, with a pair of indistinct, setiferous,
paramedian knobs at a rather wide caudal margin.
Sterna virtually bare, strongly shining, with a big, linguiform, setose, lamina
directed forward between coxae 4. Legs short, rather stout, with neither adenostyles
nor tarsal brushes, prefemora and femora with dense, strong, setae adpressed distad.
Gonopods (Figs 16-18) suberect. Coxite rather long, ventrally setose. Telo-
podite with evidence of torsion, prefemoral part unusually prominent, perhaps the
most prominent not only among congeners, but also among all Paradoxosomatidae.
Seminal groove running mostly laterally, shifting laterad already parabasally. Distal
part of tibiotarsus with a simple, mesal, distofemoral tooth and a bifid lateral lobe
with proximalmost tooth supporting seminal groove.
156 SERGEI I. GOLOVATCH
Euphyodesmus perarmatus sp. n. (Figs 19-24)
Material: HOLOTYPE 4 (MHNG), East Malaysia, Borneo, Sarawak: SE of Serian, Sabal
(= Forest Research Station), 190 m, beaten from vegetation, 10.XII.1987 (Sar-87/72); leg. C.
Lienhard. — Paratypes: 1 2, 1 juv. (MHNG), same data, together with holotype. 1 2 (MHNG),
same locality, 180 m, under bark, 10.XII.1987 (Sar-87/71); leg. B. Hauser.
Diagnosis: Differs from congeners by the especially strongly developed
gonopod tibiotarsus.
DESCRIPTION
Length ca. 21 mm, maximal width 1.1 and 1.5 (6) and 1.1-1.2 and 1.5-1.6
mm (2) on midbody pro- and metasomites, respectively. Coloration rather uniform
dark chocolate-brown to blackish, ventrum and a few distal podomeres only a little
paler, only paraterga, tips of antennae and a few proximal podomeres whitish.
Head considerably broader than narrowest collum, but a bit narrower than
somite 5. Body distinctly moniliform, gradually broadening toward somite 5, parallel-
sided on somites 5-17, onward rather gently but rapidly tapering. Antennae very long,
slender (Fig. 19), in situ surpassing somite 5 (3) or 4 (d). Paraterga rather poorly
developed, set at about 1/3-1/4 midbody height, directed somewhat obliquely dorso-
caudad, only occasionally (on pore-bearing somites) level to dorsum, usually well
below dorsum, starting already from collum, as narrow (poreless somites) or some-
what broader (pore-bearing somites) ridges, from above invariably narrowly rimmed
laterally. Caudal corner of paraterga roundly subrectangular on collum, narrowly
rounded on somite 2, onward invariably beak-shaped and spiniform (Fig. 20), reach-
ing to slightly surpassing rear tergal contour, strongly surpassing the contour only on
somites 17-18, almost vestigial on 19th. Lateral rim of paraterga with one or two
minute incisions on pore-bearing and poreless somites, gradually coming (almost) to
naught toward midbody somites. Ozopores dorsolateral, inside an oblique groove.
Surface shining, generally smooth, uniform delicately shagreened, only postsulcus
halves of metaterga slightly rugulose and metazona 2—5 below paraterga coarsely
shagreened. Somites deeply constricted, suture dividing pro- and metazona deep,
smooth. Transverse metatergal sulcus starting already from collum, rather deep, thin,
arching and lineiform (Fig. 20), only traceable on somite 19. Tergal setae medium-
sized, mostly missing, a few on collum, onward a pattern of 2+2 setae in presulcus
row, of which only 1+1 median remain. Pleurosternal keels absent, as a vestigial
oblique ridge on somite 2 and poor swellings on somites 3-4. Epiproct moderately
long, somewhat flattened dorsoventrally, in lateral view almost straight, from above
conical, narrowly truncate, with two minute apical papillae and preapical incisions
removed very far from tip. Subanal scale subtriangular, with a pair of rather indistinct,
setiferous, paramedian knobs at a rather widely rounded caudal margin.
Sterna almost bare, with distinct cross-impressions and a big, linguiform,
setose, slightly emarginate lamina directed anteroventrally between coxae 4 (Fig. 21).
Legs very long, slender, growing considerably longer toward caudal body end, with
neither adenostyles nor tarsal brushes, relatively sparsely setose even ventrally,
distinctly longer and slightly incrassate in d as compared to 2.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 157
RE SP
PI Fe
REF ESS +. 21
parto AG
LE
1.0
Fics 19-24
Euphyodesmus perarmatus sp. n., 6 paratype: 19 — anterior body portion, lateral view; 20 —
somite 10, dorsal view; 21 — sternal lamina between do coxae 4, caudal view; 22-24 - left
gonopod, mesal, lateral, and ventral views, resp. — Scales in mm.
Gonopods (Figs 22-24) relatively complex. Coxite rather short and stout,
distally setose. Telopodite with evidence of torsion, prefemoral part not particularly
prominent, femorite stout and subcylindrical. Seminal groove running mostly
laterally, shifting laterad already parabasally. Tibiotarsus unusually big and complex,
subcyathiform, almost sheathing solenomerite, latter with a strong parabasal
distofemoral tooth.
158 SERGEI I. GOLOVATCH
Euphyodesmus lobatus sp. n. (Figs 25-30)
Material: HOLOTYPE d (MHNG), East Malaysia, Borneo, Sarawak: Kuching-Serian
road, near Kampong Kuap (18 km from Kuching), secondary forest, 30 m, beaten from
vegetation, 13.XII.1987 (Sar-87/88); leg. C. Lienhard.
Diagnosis: Differs from congeners by the especially strongly reduced gonopod
tibiotarsus.
DESCRIPTION
Length ca. 18 mm, maximal width 1.0 and 1.25 (6) on midbody pro- and
metasomites, respectively. Coloration dark brown to blackish, pattern like in E.
perarmatus.
Head and body (Figs 25-28) like in E. perarmatus, except for the following
characters. Paraterga set somewhat higher, at about 1/4-1/5 midbody height, with
caudal spike mostly slightly elevated above or at least level to dorsum on posterior
body half. Incisions on metaterga a little more distinct, especially on anterior body
half, gradually coming to naught until somite 17. Metatergal sulcus poorly traceable
already on somite 18, virtually missing on somite 19. Pleurosternal carinae present as
a small but distinct tooth only on somite 2, vestigial on somite 3. Epiproct also
narrowly truncate, but with a pair of big apical papillae divided by a very narrow
notch. Subanal scale semicircular, more strongly rounded at caudal margin.
Gonopods (Figs 29-30) relatively simple. Coxite short and stout, distally
setose. Prefemoral portion very big, femorite very strongly reduced, with a distodorsal
lobe at base of a relatively slender and apically unciform solenomerite, tibiotarsus
virtually totally missing.
Euphyodesmus lateridens sp. n. (Figs 31-36)
Material: HOLOTYPE d (MHNG), East Malaysia, Borneo, Sarawak: Kuching-Matang
road, Mt Gunung Serapi, forest along road toward TV station, 700 m, beaten from vegetation,
9.XII.1987 (Sar-87/67); leg. C. Lienhard. — Paratypes: 1 d, 2 juv. (19 segm.) (MHNG), 1 &
(ZMUM), same data, together with holotype. — 1 d (MHNG), Sarawak: Bau, near cave “Gua
Puang” near Kampong Pelaman Sekiang on a steep slope of Gunung Jambusan (“Pang Cave”
in Wilford, 1964), 10-30 m, beaten from vegetation, 4.XII.1987 (Sar-87/51); leg. C. Lienhard.
Diagnosis: Differs from congeners both by characteristic distolateral tooth of
the gonopod and sternal lamina between the d coxae 4.
DESCRIPTION
Length ca. 20-21 mm, maximal width 1.1-1.2 and 1.6-1.7 mm on midbody
pro- and metasomites respectively. Coloration dark brown to blackish, pattern like in
E. perarmatus.
Head and body (Fig. 31) like in E. perarmatus, except for the following
characters. Paraterga set relatively high, like in E. lobatus, mostly below or level to
dorsum, only seldom spikes surpassing dorsum, slightly surpassing rear tergal contour
already from somite 5, only from somite 4 pointed, before more or less narrowly
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 159
Fics 25-36
Euphyodesmus lobatus sp. n., & holotype (25-30), and Euphyodesmus lateridens sp. n., à
paratype (31-36): 25 — anterior body portion, lateral view; 26, 27 & 31 — somite 10, lateral,
dorsal, and dorsal views, resp.; 28 & 32 — sternal lamina between d coxae 4, caudal view; 29-30
— right gonopod, mesal and lateral views, resp.; 33-36 — left gonopod, mesal, lateral, dorsal, and
ventral views, resp. — Scales in mm
160 SERGEI I. GOLOVATCH
rounded, especially well-protruding on somites 16-18. Metatergal sulcus rather well-
developed also on somite 18, virtually missing on somite 19. Pleurosternal carinae
present only as small but distinct teeth on somites 2-3. Epiproct also rather narrowly
truncate, apical papillae indistinct, preapical incisions very strongly removed from tip.
Subanal scale as a very narrowly truncate trapeze or triangle. Lamina between d
coxae 4 particularly big and strongly notched in the middle (Fig. 32).
Gonopods (Figs 33-36) relatively complex. Coxite short and stout, distally
setose. Prefemoral portion very prominent, femorite strongly reduced, with a
characteristic, distolateral, scapuliform, serrate tooth at base of both a big, spiniform
tibiotarsus and a strong, distally unciform solenomerite.
Remarks: The genus Euphyodesmus Attems, 1931, has hitherto been known to
comprise only three species, all confined to Borneo: E. gracilis Attems, 1931 (the
type-species), E. postfemoralis Jeekel, 1963, and E. granulifer Jeekel, 1963 (JEEKEL
1963). Discovery in Sabah and Sarawak of four additional congeners reported here at
once seems to evidence that Euphyodesmus is perhaps far more species-rich, with
further new members still to be found.
The gonopod traits of all four new congeners are clearly those characteristic of
Euphyodesmus, while certain amendments in the concept of the genus, such as the
degree of development and caudal projection of the paraterga, presence of tarsal
brushes, length of antennae and legs, etc. (JEEKEL 1963), concern only some relatively
unimportant peripheral features.
Formally, Euphyodesmus has hitherto been referred to the tribe Tectoporini,
yet with certain reservations (JEEKEL 1963, 1968). With new evidence at hand, in
particular in view of the discovery of the new monobasic genus Borneochiropus, that
allocation appears false and actually refers to the tribe Antichiropodini (see just
below).
All currently known Euphyodesmus species can be separated using the
following key:
1(2) Metaterga with two transverse rows of knobs, presulcus one setiferous,
and postsulcus one bare. Gonopod suberect, tibiotarsus as a big, simple
lamina a little shorter than a bifurcate solenomerite............ E. granulifer
2(1) Metaterga without knobs, either hairless or with a single presulcus row
of setae. Gonopod either strongly enlarged or strongly reduced distally,
often unciform not suberect; tibiotarsus either strongly reduced, often
denti- or spiniform, or strongly folded and complex in shape.
3(4) Antennae relatively short, in situ reaching only the end of somite 2
dorsally. Paraterga strongly reduced, virtually missing on somites 7-19
(Fig. 15). Legs relatively short and stout (Fig. 15). Gonopod prefemoral
part hypertrophied out of all proportions, both tibiotarsus and soleno-
merite simple, dentiform, strongly reduced (Figs 6-18).......... E. reductus
4(3) Antennae and legs considerably longer. Paraterga always present,
mostly spiniform caudally. Gonopod prefemur never so strongly hyper-
trophied, both tobiotarsus and solenomerite never so strongly reduced.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 161
5(6) Paraterga relatively underdeveloped (Figs 19-20), usually well below
dorsum, set at 1/3-1/4 midbody height, only their caudal spikes some-
times scarcely reaching the level of dorsum. Transverse sulcus still
traceable on metasomite 19. Gonopod tibiotarsus hypertrophied, sole-
nopnore-like;(Elos22 DIN SH Eee ace E. perarmatus
6(5) Paraterga better developed, set mostly at 1/4-1/5 midbody height,
almost to fully level to dorsum. No transverse sulcus on metasomite 19.
Gonopod tibiotarsus never solenophore-like, nor sheathing solenomerite.
7(10) Suture dividing pro- and metazona striate, although quite indistinct.
Gonopod tibiotarsus strongly folded, complex in outline.
8(9) Head only a little wider than collum. Solenomerite about as long as
Gbietarsus Simplysacuminate attendre ee tn ee a ee E. gracilis
9(8) Head much wider than collum. Solenomerite distinctly longer than
ub1otarsusguneitermragend. EE E. postfemoralis
10(7) Suture dividing pro- and metazona virtually smooth. Gonopod tibiotar-
sus neither folded nor complex in outline.
11(12) Size lesser: width 1.25 mm. Sternal lamina between d coxae 4 without
notch (Fig. 28). Gonopod femorite without processes, tibiotarsus espe-
cially strongly reduced, retained only as a low lobe (Figs 29-30)... . E. lobatus
12(11) Size bigger: width 1.6 mm. Sternal lamina between d coxae 4 notched
apically (Fig. 32). Gonopod femorite with a distolateral tooth, tibio-
tarsussloncandispiniionm(Figs/3 3-30) 52st aba ae ene fae E. lateridens
Borneochiropus gen. n.
Type-species: Borneochiropus spinitergus sp. n.
Medium-sized Antichiropodini with very modestly developed paraterga.
Adenostyles absent. Sternal lamina between d coxae 4 present.
Gonopods remarkable by a medium-sized coxite, a hypertrophied prefemur, a
shortened and definitely tortiled femorite, and a very long and ribbon-shaped soleno-
merite, without trace of a solenophore, but with an outgrowth near base of soleno-
merite. Seminal groove running mostly laterally before reaching solenomerite.
Remarks: The tribe Antichiropodini has hitherto been considered as restricted
to Australia and Tasmania, with only a single possible member encountered in New
Guinea (JEEKEL 1968, 1979b). Therefore, the discovery of a representative on Borneo
is highly interesting and important zoogeographically (see below).
Within the tribe, Borneochiropus seems quite disjunct. Indeed, the absence of
adenostyles is almost unique, this character being shared only with the monobasic
genus Mjoebergodesmus Verhoeff, 1924, from Australia. However, the most striking
deviations lie in gonopod structure. What has no parallel not only among Antichiro-
podini, but perhaps amongst all Paradoxosomatidae but Euphyodesmus is the pre-
femoral part of Borneochiropus which is hypertrophied, and the femorite is relatively
strongly reduced and unusually strongly shifted dorsad in relation to the prefemur.
162 SERGEI I. GOLOVATCH
The relatively long solenomerite, presence of only a single medio-distofemoral
process at the base of the solenomerite, the seminal groove running first along a
torsate femorite and then between the distofemoral process and the base of the
solenomerite rather strongly resemble those observed in Haplochiropus Attems, 1944.
Originally, it was proposed as a subgenus of the Southwest Australian genus Anti-
chiropus Attems, 1911, for a single species, A. pustulosus Attems, 1944, from New
Guinea, but later its tribal position was questioned by JEEKEL (1968). However, in
view of the existence of a number of true antichiropodines on Borneo, there seems to
be no reason to further suspect that ATTEMS (1944) could have mistaken an anti-
chiropodine for an eustrongylosomatine. In other words, Haplochiropus is promoted
herewith to full generic status, being to some extent intermediate between Australian
Antichiropodini and Borneochiropus not only morphologically, but also geogra-
phically.
Yet Borneochiropus displays the most clear-cut relationships with Euphyo-
desmus, thus corroborating with zoogeographical evidence as well. This largely
concerns the shared strongly reduced and spiniform paraterga, the strongly hyper-
trophied gonoprefemorite and the thick solenomerite very often devoid of any sup-
port/trace of a solenophore. However, the course of the seminal groove in Borneo-
chiropus is singular in passing onto the solenomerite not on the ventral, as is the case
in Euphyodesmus, but on the dorsal side of the distalmost part of the gonofemorite.
This certainly makes both stand as “good” genera. In other words, Euphyodesmus and
Borneochiropus must be regarded as especially closely related genera of the tribe
Antichiropodini.
Borneochiropus spinitergus sp. n. (Figs 37-41)
Material: HOLOTYPE d (MHNG), East Malaysia, Borneo, Sabah, Sandakan Residency,
Sepilok, forest near pond, degraded lowland rain forest, beaten from vegetation, 26.1V.1982
(8240); leg. D. Burckhardt. — Paratypes: 1 d, 1 2 (MHNG), same locality, together with holo-
type. — 1 © juv. (19 segm.) (MHNG), same locality, 26.1V.1982 (8239); leg. D. Burckhardt.
DESCRIPTION
Length of adults ca. 19-20 mm, width of midbody pro- and metazona 0.8 and
1.0-1.1 (3 8), and 1.0 and 1.2 mm (@), respectively. Coloration of adults uniform
pale yellowish-brown to castaneous-brown, that of juvenile entirely pallid; in adults
only paraterga, distal half of antennomere 6, proximal half of antennomere 7, tip of
antenna, coxae, prefemora, and most of femora whitish.
Head considerably broader than collum and even midbody somites, postcollar
constriction rather evident due to narrowest somites 3-4. Body strongly moniliform,
parallel-sided on somites 5-17, onward gently and quite regularly tapering. Antennae
very long, slender, in situ reaching beyond somite 5. Paraterga very modestly
developed, poorly cariniform on collum and somites 2-4, onward mostly spiniform,
somewhat better developed, spike-like, directed largely more dorsally than caudally,
almost reaching to rear tergal contour on pore-bearing somites, gradually declining
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 163
Fics 37-41
Borneochiropus spinitergus gen. n., Sp. n., d paratype: 37 — somite 10, dorsal view; 38 —
sternal lamina between dé coxae 4, ventrocaudal view; 39-41 — left gonopod, mesal, lateral, and
ventral views, resp. — Scales in mm.
almost to naught on somites 16-19. Paraterga generally set very high, on pore-bearing
somites largely above dorsum and almost devoid of rim laterally (Fig. 37), on poreless
ones invariably below dorsum, rather strongly rimmed laterally and weil removed
from rear tergal contour. Surface poorly shining, entirely smooth and very delicately
shagreened. Somites strongly constricted, suture dividing pro- and metazona quite
deep, somewhat more roughly shagreened but not striate. Transverse metatergal
sulcus starting already from collum (d ) or somite 5 (2), very modestly deep, thin and
lineiform, almost wanting on somite 18, totally missing on somite 19. Tergal setae
medium-sized, a few on collum and 1+1 on anterior half of somites 2-3, onward
obliterated. Pleurosternal keels vestigial and present only on somite 2. Epiproct very
short, weakly flattened dorsoventrally, narrowly truncate at tip. Subanal scale broadly
and roundly subtrapeziform, usual.
Lamina between d coxae 4 setose, distinctly bifid, higher than coxa, directed
ventrad and slightly forward (Fig. 38). Sterna generally more shining, sparsely setose,
without modifications. Legs extremely long, several times as long as body diameter,
slender, in 9 a little shorter, d tarsal brushes only on pairs 1-2, onward setation
thinning out together with leg elongation toward telson, without adenostyles.
Gonopods (Figs 39-41) very particular. Coxite moderately long, distoventrally
modestly setose. Prefemur unusually strongly developed, setose as usual. Femorite
164 SERGEI I. GOLOVATCH
shortened, with evidence of torsion, strongly shifted dorsad and knee-shaped near
middle, with a curved digitiform outgrowth (vestigial tibiotarsus?) near border
between femorite and solenomerite. No trace of tibiotarsus. Solenomerite very long,
somewhat coiled, rather ribbon-shaped, apically with a spike terminating seminal
groove. Latter running mostly laterally before getting onto solenomerite.
Orangutana gen. n.
Type-species: Orangutana setosa sp. n.
Relatively small Orthomorphini with very modestly developed paraterga.
Adenostyles absent. Sternal lamina present only between 4 coxae 4.
Gonopods remarkable by the following combination: coxite long; femorite
very long and slender, lacking a sulcus demarcating a postfemoral part; tibiotarsus (=
solenophore) hypertrophied due to both lamina lateralis and a distally strongly
notched lamina medialis, both laminae sheathing most of a relatively short and
flagelliform solenomerite; free solenomerite beginning considerably proximally of,
not about level to, sulcus demarcating tibiotarsus. Seminal groove entirely mesal.
REMARKS: In the entire Sunda area, the chiefly Oriental tribe Orthomorphini
has hitherto been known to be represented by the genera Orthomorpha Bollman, 1893
(Malay Peninsula, Java as well as Farther India, Bali and Lombok), Gigantomorpha
Jeekel, 1963 (Borneo, Sulawesi, Bali), Dajakina Jeekel, 1963 (Borneo, Sumatra),
Nesorthomorpha Jeekel, 1980 (Java), Diglossosternum Jeekel, 1980 (Java), and
Luzonomorpha Hoffman, 1973 (Philippines). Of these, only Orthomorpha displays a
gonopod tibiotarsus simple and virtually undivided at tip, so Orangutana appears to
be more closely related in this respect to the remaining set. No gonopostfemoral
sulcus is marked only in Dajakina and Diglossosternum, so Orangutana approaches
here both of them. However, the distinctly hypertrophied gonopod solenophore and
the beginning of a free solenomerite considerably more proximally of the demarcation
sulcus between the femorite and tibiotarsus seem unique and make Orangutana easily
distinguished from all other known Orthomorphini (see also review: GOLOVATCH
1994a).
Orangutana setosa sp. n. (Figs 42-45)
Material: HOLOTYPE d (MHNG), East Malaysia, Borneo, Sabah: Sandakan Residency,
Sepilok, forest near pond, leaf litter in degraded lowland rain forest (extraction with Berlese-
Tullgren funnel), 10.V.1982 (Sab-82/41); leg. B. Hauser. — Paratypes: 1 d, 1 2 juv. (19 segm.)
(MHNG), same locality, together with holotype. — 1 d, 1 £ juv. (19 segm.) (ZMUM), same
locality, sifted leaf litter and rotten wood, Winkler-Moczarski extraction, 23.1V.1982 (Sab-
82/7); leg. B. Hauser.
Etymology: The generic and specific epithets refer to the cohabitation with the
famous ape Orang-Utan and to the densely setose head and metaterga, respectively.
DESCRIPTION
Length of adults ca. 11-12 mm, width of midbody pro- and metazona 0.95 and
1.0 mm, respectively. Coloration of adults uniform pale yellowish-brownish to pale
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 165
Fics 42-45
Orangutana setosa gen. n., Sp. n., d paratype: 42 — somite 10, lateral view; 43 — left gonopod,
mesal view; 44-45 — gonopod tip, sublateral and subventral views, resp. — Scales in mm.
brown, juvenile entirely pallid; in adults tip of antenna, legs and ventrum whitish-
yellow.
Head broadest, a bit broader than midbody somites; collum and somites 2-4
subequal in width, narrowest, but postcollar constriction poorly developed. Body
parallel-sided on somites 5-18, onward tapering mostly on telson. Head very densely
setose. Antennae rather short, slightly clavate, in situ reaching only beyond somite 2.
Paraterga poorly developed, as a thin lappet on collum, always spiniform on
subsequent somites, somewhat thicker on pore-bearing somites, set rather low, about
1/3-1/4 midbody height (Fig. 42), always within rear contour on poreless somites,
(almost) within contour even on pore-bearing somites in front of somite 12 (Fig. 42)
increasingly strongly surpassing rear tergal contour and beak-shaped until somite 15,
onward gradually reduced. Surface of prozona dull, very delicately shagreened; sterna
shining; metaterga shining dorsally, microgranulate only below paraterga, faintly
rugulose in front of metatergal sulcus, more strongly behind it. Somites relatively
poorly constricted, suture dividing pro- and metazona rather shallow, poorly striate
longitudinally. Transverse metatergal sulcus starting from somite 5, shallow, not very
thin, lineiform, almost reaching to base of paraterga, missing on somite 19. Traces of
166 SERGEI I. GOLOVATCH
an axial line on postsulcus parts of metaterga. Tergal setae rather long, very dense on
collum, onward considerably less dense (also many broken off), but little by little
growing more dense caudally, somite 19 already beset with setae almost like head;
setation pattern usually as 1-2 irregular rows in front of and 2-3 rows behind sulcus,
caudalmost row being especially dense and complete (Fig. 42). Pleurosternal keels
entirely missing. Epiproct medium-sized, a little flattened dorsoventrally, with pre-
apical incisions well removed from a narrowly rounded tip. Subanal scale trapezi-
form, with a usual paramedian pair of indistinct setiferous knobs near rear margin.
Lamina between d coxae 4 setose, broad, about as high as coxa, directed
ventrad and slightly forward, distinctly bimodal. Sterna sparsely setose, with vestigial
rounded knobs near coxae from d somite 8 onward. Legs relatively short, slightly
longer and incrassate in dd as compared to juvenile, with neither adenostyles nor
tarsal brushes.
Gonopods (Figs 43-45) very high, generally slender, only terminally falcate
caudally. Femorite with neither traces of torsion nor postfemoral demarcation.
Solenophore very strongly developed, about 1/3 femorite in length, set off from
femorite by a distinct demarcation sulcus, curved caudad, sheathing a relatively
moderately long solenomerite almost all along its length; both lamina medialis and
lamina lateralis equally well-developed, former terminally deeply notched. Seminal
groove entirely mesal, free solenomerite starting well before demarcation site between
femorite and solenophore.
Dajakina longipes sp. n. (Figs 46-53)
Material: HOLOTYPE ¢ (MHNG), East Malaysia, Borneo, Sabah: Sandakan Residency,
Sepilok, “Kabili-Sepilok Forest Reserve”, forest near “Orang-Utan Rehabilitation Station”, 30
m, lowland rain forest, 22.1V.1982 (Sab-82/2); leg. B. Hauser. — Paratypes: 1 d (MHNG), 1 d
(ZMUM), same locality, together with holotype. — 2 4 4 (MHNG), same locality, beaten from
vegetation, 12.11.1983 (Pal-83/7); leg. C. Lienhard.
Etymology: The species epithet refers to the extremely long legs.
Diagnosis: Differs from congeners by the presence of a sternal lamina between
the 8 coxae 4 combined with the absence of gonofemoral outgrowths, as well as by
the shape of the gonopod tibiotarsus.
DESCRIPTION
Length ca. 29-32 mm, width of midbody pro- and metazona 1.9-2.0 and 2 5-
2.6 mm, respectively. Holotype ca. 30 mm long, 2.0 and 2.5 mm wide on midbody
pro- and metazona, respectively. Coloration blackish-brown dorsally to somewhat
below paraterga, rather sharply contrasting with whitish-yellow ventrum, legs, and
lower halves of sides. Antennae brown to dark brown, basal half of antennomere 7
invariably, tip of antennomere 6 sometimes, very dark brown to blackish. Clypeus and
labrum whitish to pale brown.
Head, collum, somites 3-4 subequal in width, each a bit broader than narrowest
somite 2, but a little more narrow than somite 5. Body modestly moniliform, parallel-
sided on somites 5-17, onward very gently and regularly tapering. Antennae very
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 167
Fics 46-53
Dajakina longipes sp. n., à é paratypes: 46-47 — somite 10, lateral view; 48 — sternal lamina
between d coxae 4, caudal view; 49-53 — gonopods, mesal, lateral, ventral, mesal, and lateral
views, resp. — Scales in mim.
long, slender, very slightly clavate, in situ reaching beyond somite 4. Paraterga very
modestly developed, especially thin on collum and somite 2, considerably thicker on
pore-bearing somites, set rather low, 1/2-1/3 midbody height (Figs 46-47), always
delimited from above and, especially on poreless somites, from below, caudally
largely poorly dentiform, a little surpassing rear tergal contour only on somites 2, 18,
and 19. Surface poorly shining, generally smooth, rugulose only behind metatergal
sulcus. Somites rather strongly constricted, suture dividing pro- and metazona quite
168 SERGEI I. GOLOVATCH
deep and strongly striate longitudinally. Transverse metatergal sulcus starting from
somite 5, moderately deep, thin and lineiform, missing on somite 19. Tergal setae
short, a few on collum and at least 1+1 on anterior half of somite 2. Pleurosternal
keels present as rather strong, caudally pointed teeth surpassing rear contour on
somites 2-4, smaller denticles almost within contour on somites 5-7, scarcely tra-
ceable on somite 8, onward virtually missing. Epiproct medium-sized, somewhat
flattened dorsoventrally, with preapical incisions extremely close to a roundly trun-
cate tip, in dorsal view almost parallel-sided throughout, in lateral view almost
straight. Subanal scale semi-circular, with a usual paramedian pair of indistinct
setiferous knobs near rear margin.
Lamina between d coxae 4 setose, linguiform, about as high as coxa, directed
ventrad and slightly forward (Fig. 48). Sterna generally shining, scarcely setose,
without modifications. Legs very long (Fig. 47), slender, with neither adenostyles nor
tarsal brushes; very moderate setation thinning out together with leg elongation
toward telson
Gonopods (Figs 49-53) very slender. Femorite often with traces of dorsal
wrinkles parabasally, without postfemoral demarcation. Solenophore about as long
and slender as femorite, slightly curved caudad, sheathing a moderately long soleno-
merite almost all along its length. Tip of tibiotarsus slightly varying in shape.
Remarks: The genus Dajakina Jeekel, 1963, has hitherto been known to com-
prise four species: D. oculata Jeekel, 1963 (the type-species), D. rotundata (Attems,
1931), D. hastata Jeekel, 1983b, all from Borneo, and D. inermis (Silvestri, 1895),
from Sumatra (JEEKEL 1963, 1979a, 1983b). The new species is interesting in having
the gonofemorite often slightly rugose in the area supporting a fringe in D. oculata
and a strong process in D. hastata.
All currently known Dajakina species can be separated using the following key:
1(4) Paraterga of pore-bearing somites abbreviated, eye-shaped in lateral
view.
2(3) Size lesser: width 2.4 mm (d). Antennomere 6 unmodified, subcylin-
drical, without inflation. Pleural areas unmodified. Sternal lamina
between d coxae 4 present, conical. Gonopod femorite with a long,
lanceolate, dorso=parabasal process: 2. eee re D. hastata
3(2) Size bigger: width 4.0 mm (4). Antennomere 6 characteristically in-
flated. Pleural areas of d somites 8-18 each with a caudal emargina-
tion. No sternal process on d somite 5. Gonopod femorite with only a
spikemmstead sof panabasall process... sane eee D. oculata
4(1) Paraterga of pore-bearing somites as usual ridges, demarcated by a
furrow at least from above.
5(6) Paratergal caudal corner strongly spiniform on several caudalmost so-
mites. Transverse metatergal sulcus starting from somite 2. Sumatra. D. inermis
6(5) Paratergal caudal corner at most poorly dentiform. Transverse meta-
tergal sulcus starting from somite 5. Borneo.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 169
7(8) Paraterga rounded caudally, pore-bearing ones demarcated by a furrow
only from above. No sternal lamina between d coxae 4. Tarsal brushes
ontartewzantenionlegpairsiof er rer D. rotundata
8(7) Paraterga poorly dentiform caudally; pore-bearing ones demarcated by
a furrow both from above and below (Figs 46-47). Sternal lamina
between d coxae 4 present (Fig. 48). Tarsal brushes wanting... . . D. longipes
Gigantomorpha immanis Jeekel, 1963 (Figs 54-62)
Material: 2 33 (MHNG), East Malaysia, Borneo, Sabah, Sandakan Residency,
Sepilok, “Kabili-Sepilok Forest Reserve”, forest near “Orang-Utan Rehabilitation Station”,
lowland rain forest, 24.11.1983 (Pal-83/50); leg.B. Hauser. — 1 2 (MHNG), same locality,
3.V.1982 (Sab-82/26).- 1 2 (MHNG), Sepilok, forest near pond, degraded lowland rain forest,
4.V.1982 (Sab-82/28). - 2 dd (MHNG), 1 4 (ZMUM), Sepilok, “Kabili-Sepilok Forest
Reserve”, path toward mangrove, before “Ridge”, lowiand rain forest, 6.V.1982 (Sab-82/30). —
1 3 (MHNG), Interior Residency, Kimanis road, 8.5 mi from Keningau, path from
“Checkpoint” to above village “Temperate Vegetation Project”, 950-1150 m, 13.V.1982 (Sab-
82/47). — 1 4 (MHNG), same locality, 8.5 mi from Keningau, Checkpoint, near barracks, under
stones, 950 m, 14.V.1982 (Sab-82/52); all leg. B. Hauser. — 1 & (broken into several pieces)
(MHNG), Borneo, Brunei, Temburong Distr., “Peradayan Forest Reserve” (= “Bukit Patoi”),
14.5 km from Bangar (= 2.5 km from Labu), primary mixed dipterocarp forest, 10-300 m,
24.X1.1988 (Bru-88/39); leg. C. Lienhard.
Remarks: The genus Gigantomorpha Jeekel, 1963, has hitherto been known to
comprise nine species: G. immanis Jeekel, 1963 (the type-species), G. aterrima
Jeekel, 1963, G. cornalata Jeekel, 1963, G. fasciata (Attems, 1898), G. pallida
Jeekel, 1963, G. spinescens Jeekel, 1963, G. transmontana Jeekel, 1983b, G. tricho-
pleura Jeekel, 1963, all from Borneo, and G. socialis (Carl, 1912), from Sulawesi
(JEEKEL 1963, 1983b). A tenth congener, G. mutilata sp. n., has just been encountered
on Bali (see below). Of them, G. immanis seems to be quite widespread, being
encountered at least in Northwest and East Borneo. The above material extends
considerably the known range of this species northwesterly.
Already JEEKEL (1963) in his original description of G. immanis noted its
pronounced variability concerning size, shape of the paraterga and epiproct, leg
pubescence, etc. Based on such variations, he even referred some specimens to G.
immanis With qualifications, writing that “a much more abundant material is needed
to decide whether these characters are of really subspecific value or just represent
population varieties” (op. cit., p. 247).
The above new evidence allows to significantly refine the variation range of G.
immanis. The only Brunei d is 3.95 and 6.8 mm in width on midbody pro- and
metazona, respectively, while the Sabah specimens at hand are only 48-55 mm in
length, width of midbody pro- and metazona 3.9-4.2 and 6.7-7.0 (4 4), and 4.2-4.8
and 6.8-7.0 mm (9 9), respectively, this being on the average somewhat less than
reported by JEEKEL (1963). The 4 legs are ventrally densely setose, especially due to
prefemora, femora and tarsi, but the degree of gradual thinning out of setation toward
the caudal body end varies, sometimes concerning not only somites 15-18, but even
13-18. Sometimes the background coloration can be quite pale brown, but the pattern
170 SERGEI I. GOLOVATCH
Fics 54-62
Gigantomorpha immanis Jeekel, 1963, ¢ 4 from near Keningau Checkpoint (54, 56, 58-60)
and Kabili-Sepilok (55, 57, 61-62): 54-55 — somite 10, lateral view; 56-57 — somite 10, dorsal
view; 58 — telson, ventral view; 59-61 — gonopods, mesal, lateral, and mesal views, resp.; 62 —
distal part of gonopod, mesal view. — Scales in mm.
mostly persists, discarding the poor development to virtual absence of a pale
prosomital spot in the Brunei and some Sabah samples. What seems even more
important is that not only the outline of some paraterga somewhat varies in dorsal
view (cp. Figs 56 & 57), but the degree of elevation over the dorsal level ranges from
somewhat below (Fig. 54) to a little above it (Fig. 55) even on midbody somites of
Sabah specimens alone. The paraterga start surpassing the rear tergal contour on the
somites 11 or 12-19, sometimes being very narrowly rounded, more often pointed
throughout. The tubercles near the caudal margin of metaterga are often not very flat,
as small round knobs (cp. Figs 56-57), the pattern of tuberculation usually being quite
irregular, 3-4+3-4, on more caudal metaterga up to 5+5. The metatergal sulcus is
always lineiform, mostly quite shallow, only more seldom relatively deep. The
terminal knobs on the epiproct can be somewhat bigger or smaller (Fig. 58). Finally,
even the gonopods vary in outline a bit (Figs 59-62), although the tip is very.
characteristic.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 171
WANG & TANG (1965) reported Orthomorpha hydrobiologica Atems, 1930,
from Singapore. A bit later WANG (1967b) recorded there also Gigantomorpha
tenuipes (Atems), a Javan species long known to actually belong to the genus Ortho-
morpha (e.g., JEEKEL 1963). Why the latter species was transferred in Gigantomorpha
by Wang, remains totally obscure, the most probable reason why JEEKEL (1968)
ignored it altogether. What seems more important is that an endemic Singapore
Orthomorpha has since emerged. This is O. murphyi Hoffman, 1973, which might
well correspond to Wang's record(s) above.
In addition, WANG (1967a) reported Gigantomorpha fasciata (Attems) from
Brunei and simultaneously transferred three Philippine species of Paradoxosomatidae
to Gigantomorpha, again giving no explanation to the latter reallocations. It seems
noteworthy that all those three forms from the Philippines have since been placed in
the orthomorphine genus Luzonomorpha Hoffman, 1973. Such obvious mistakes of
WANG's, some more of which will be mentioned further below, coupled with a very
poor quality of his descriptions question most if not all his diplopod taxonomy (see
also below). Concerning WANG's (1967a) record of a Gigantomorpha in Brunei, I
believe he could have actually dealt with G. immanis.
Gigantomorpha mutilata sp. n. (Figs 63-70)
Material: HOLOTYPE d (MHNG), Indonesia, Bali: Bedugul, slope above Lake Bartan,
1130-1180 m, beaten from vegetation, 29.XI.1987 (Sar-87/34); leg. C. Lienhard. — Paratypes: 1
? (MHNG), same data, together with holotype. - 2 22 (MHNG), 1 2 (ZMUM), same
locality, Botanical Gardens, forested part, 1230-1260 m, chiefly beaten from vegetation,
30.X11.1987 (Sar-87/36); leg. C. Lienhard.
Diagnosis: Differs from congeners largely by the least body size and conspi-
cuous gonopod structure.
DESCRIPTION
Body length ca. 20-22 mm, midbody pro- and metazonites 2.0 and 3.0 (¢), 2.6
and 3.2 or 2.0 and 2.8-2.9 mm (2 ©), respectively. Coloration generally dark rusty
(purplish) brown to chocolate brown; paraterga, lower parts of sides, epiproct and legs
brown to pale brown; sterna yellow-brown, tip of antenna whitish.
Head subequal in width to collum and a bit narrower than somite 2, latter a bit
narrower than subequally wide somites 3-4. Body parallel-sided on somites 5-16,
onward gently and gradually tapering. Antennae not very long, rather slender, in situ
extending only beyond somite 3 (4) or 2 to midway of 3rd (@) dorsally. Paraterga
very strongly developed (Fig. 63), set high (at about 1/4 midbody height), yet not
reaching to level of dorsum (Fig. 64), in dorsal view broadly rimmed, in lateral view
almost equally narrow on pore-bearing and poreless somites, caudally mostly beak-
shaped, pointed and surpassing rear tergal contour (especially well so in ¢), narrowly
rounded only on collum (4) or collum and somite 2 (®), considerably less strongly
protruding only on somite 19. Ozopores big, lateral, inside a shallow groove. Surface
poorly shining, only sterna shining, prozonites delicately shagreened, metaterga
(72, SERGEI I. GOLOVATCH
Fics 63-70
Gigantomorpha mutilata sp. n., è holotype: 63-64 — somite 10, lateral and dorsal views, resp.;
65 — epiproct, dorsal view; 66 — sternal lamina between d coxae 4, caudal view; 67-68 — right
gonopod, mesal and lateral views, resp.; 69-70 — distal part of right gonopod, ventral and
subfrontal views, resp. — Scales in mm.
generally smooth, metazonites slightly rugulose only near base of paraterga and near
caudal edge, often almost smooth, sometimes (d ) caudal edge with traces of incisions
(Fig. 63); surface below paraterga slightly rugulose and delicately granulate, only
collum with traces of an axial line. Both dorsal and lateral surfaces of metazonites
micropiliferous, micropilosity being mostly fully or considerably obliterate. Somites
only slightly constricted, suture dividing pro- and metazonites shallow, without stria-
tion. Transverse metatergal sulcus starting from somite 5, rather shallow, thin and
lineiform, reaching to base of paraterga, missing on somite 19. Tergal setae medium-
sized, mostly broken off, a few on collum, pattern 2+2 in a presulcus row. Pleuro-
sternal carinae as little caudal teeth only on somites 2-4, onward wanting. Epiproct
long, flattened dorsoventrally, in lateral view almost straight, from above conical,
preapical incisions little, well removed from tip, apical papillae big (Fig. 65). Subanal
scale high and trapeziform, with a pair of well-developed, setiferous, paramedian
knobs at caudal margin.
Sterna densely setose, without modifications, between d coxae 4 with a
moderately big, roundly subtrapeziform, setose lamina directed mainly ventrally (Fig.
66). Legs not very long, slender, slightly longer and more strongly incrassate in à,
|
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 173
without adenostyles, femora normal, tarsal brushes present until 4 legs of somite 11,
onward setation thinning out yet very dense, especially so ventrally and in 4.
Gonopods (Figs 67-70) high and rather complex. Coxite rather short, slender,
cylindrical, distoventrally setose. Prefemoral part medium-sized, densely setose as
usual. Femorite rather long, folded laterally and, especially, mesally; distofemoral
portion demarcated by a well-developed postfemoral sulcus laterally and ventrally;
apical part with a ventral lobule with solenomerite starting at its base. Both tibiotarsus
and solenomerite long and slender, slightly curved, subequal in length. Apical part of
tibiotarsus (= solenophore) without distinct branching, with a few minute lobules and
a little ventro-preapical spinule.
Remarks: As noted before, the bulk of Gigantomorpha constituents is Bornean
(JEEKEL 1963, 1983b). Interestingly, the above new species extends the generic range
to the Lesser Sunda. The relatively small size and a little disjunct gonopod
conformation of G. mutilata, especially the particularly strongly folded femorite,
parallel its peripheral distribution, yet failing to broaden the concept of the genus
concerned.
A key to Bornean Gigantomorpha already exists (JEEKEL 1963) and still holds
valid, except that G. fasciata sensu JEEKEL (1963) has since been shown to represent
an independent species, G. transmontana (JEEKEL 1983b). Superficially, another key
based on such limited materials as above might seem superfluous, yet just below I
provide one, for it allows to incorporate all currently known congeners and to refine
some diagnostic features:
1(2) Lesser: width 2.8-3.2 mm (d, ©). Paraterga especially strongly bor-
dered (Fig. 63). Sternites usual, without cones near coxae. Gonopods as
INS OD ENTER e der, rca het Ma. AON Maa nena mec O G. mutilata
2(1) Bigger: width of adults at least 5 mm. Paraterga less strongly bordered.
Sternites modified, usually with a cone near coxa. Gonopods different.
Borneo or Sulawesi.
3(6) Lateral sides of collum rounded, caudally not angular. Paraterga 2
turned somewhat ventrad, distinctly below level of paraterga 3. Even 3
femora straight.
4(5) Lesser: width 5.0-5.4 mm (d, ©). A pattern of 4(5)+4(5) very small
tubercles near caudal margin of metaterga. Paraterga set relatively low
(at about midheight on d somite 4), below them almost all somites with
an irregular row of setae along caudal margin. Pleurosternal keels
extendinsibackito’somitesl 7 Emm BI ERREUR G. trichopleura
5(4) Bigger: width 6.3-8.0 mm (d, ?). Usually 3+3 bigger tubercles near
caudal margin of metaterga. Paraterga set distinctly higher (at about 1/3
height on d somite 4), without setae below them. Pleurosternal keels
exiendinsibackonly.tosomite 140192 2... 22 an G. spinescens
6(3) Lateral sides of collum angular caudally. Paraterga 2 turned somewhat
dorsad, nearly level to paraterga 3. d femora distinctly arched.
174 SERGEI I. GOLOVATCH
5(8) Coloration in ethanol very dark brown to blackish, with paraterga and a
rather narrow continuous middorsal band pale brownish. Gonopod
femorite rather strongly constricted about halfway.
6(7) Pleurosternal keels traceable back to somite 14 or 15. 4 legs more
strongly incrassate, all podomeres very densely setose ventrally, form-
ing real brushes of very dense and short setae at least on femora, the
brushes gradually thinning out caudally and disappearing only in last
PAS SSE ee OM GE NR M RENE De NR G. fasciata
7(6) Pleurosternal keels traceable back to somite 10 only, onward missing.
é legs less strongly incrassate, ventral setation on podomeres forming
Mintuallysnostiueybrushes sss). TOSI. RME RE G. transmontana
8(5) Somites without a continuous middorsal pale band. Gonopod femorite
devoid of a notable midway constriction.
9(10) Coloration in ethanol uniform blackish, only paraterga near caudal
corner brown. d proximal podomeres beset with short curved setae,
femora less;stronglysarched ye EC us... E G. aterrima
10(9) Coloration of dorsum in ethanol either somewhat paler, sometimes with
a pale central spot on prosomites, or dark with contrastingly yellow
parts, then at least paratergal caudal corners always yellowish. d legs
beset with long erect setae, femora more strongly arched.
11(14) Pleurosternal keels traceable until somite 16 or 17.
12(13) Coloration of dorsum rather uniform pale brownish-gray. Paraterga
more or less distinctly projecting caudally beyond rear tergal contour
from: somite Iiorl2 onward: East Bomeo..... 222... Er G. pallida
13(12) Coloration black-brown with caudal corners of paraterga yellow. Para-
terga extending caudally beyond rear tergal contour only from somite
[Ssonwardy South Sulawesi... RE G. socialis
14(11) Pleurosternal keels traceable back at best to somite 14 or 15.
15(16) Paraterga largely strongly elevated above dorsum. Pleurosternal keels
traceable back to somite 14 or 15. 4 legs more slender, proximal
podomeres without ventral brushes of setae. .................. G cornalata
16(15) Paraterga more or less level to dorsum (Figs 54-55). Pleurosternal
keels traceable back only to somite 9 or 10. 4 legs more strongly
incrassate, with ventral brushes on all podomeres (Figs 54-55). . . . G. immanis
Kalimantanina crucifera sp. n. (Figs 71-75)
Material: HOLOTYPE 4 (MHNG), East Malaysia, Borneo, Sabah, Sandakan Residency,
Sepilok, forest near pond, degraded lowland rain forest, beaten from vegetation, 8.V.1982
(82150), leg. D. Burckhardt. — Paratype: 2 (MHNG), same locality, 26.1V.1982 (8240); leg. D.
Burckhardt.
Etymology: The species epithet refers to the peculiar colour pattern on each
somite resembling a cross.
Diagnosis: Differs from congeners by the cruciform colour pattern combined
with the particularly strongly reduced tooth at base of the solenomerite.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 175
DESCRIPTION
Length ca. 26 (6) and 31 mm (9%), width 2.0 (6) and 2.6 mm (9). Back-
ground coloration pale whitish with a peculiar pattern on dorsum consisting of a dark
brown, rather narrow, axial stripe crossed on each metatergum by a similar transverse
band outlining sulcus, crossing areas and protergal line being often superimposed
over a more or less big, pale brown spot; a brown, almost continuous, lateral stripe on
each side level to paraterga and covering caudal half of paraterga and mostly
extending onto prozona. Antennae pallid, only distally darker, especially due to very
dark brown distal half of antennomere 6 and basal half of antennomere 7. Head pale
brown eventually throughout. Collum margined narrowly brown.
Head a bit broader than subequal collum and somite 2, but a bit narrower than
somites 5-18. Postcollar constriction evident on subequally narrowest somites 3-4.
Body distinctly moniliform, rapidly tapering caudad on somites 19-20. Antennae
long, slightly clavate, in situ reaching to end of somite 4 in d, missing (right) or too
short (left) in £ (perhaps regenerated). Paraterga modestly developed, somewhat
thicker on pore-bearing somites, set rather low, mostly at 1/2-1/3 midbody height,
demarcated by a sulcus from above and below, caudally invariably gently rounded
and lying mostly within rear tergal contour (Fig. 71), only on somite 19 just a bit
surpassing it. Ozopores lying near caudal corner of paraterga in an indistinct groove
(Fig. 71). Surface shining, generally smooth, slightly rugulose only below paraterga
and behind sulcus. Somites strongly constricted, suture dividing pro- and metazona
deep, strongly striate longitudinally. Transverse metatergal sulcus starting from
somite 5, moderately deep, thin and lineiform, missing on somite 19. Tergal setae
minute, a few on collum, pattern of 2+2 setae on somite 2, of 1+1 at least on somites
3 and 19, mostly missing. Pleurosternal keels present only until somite 7, onward
totally missing, better developed in à , displayed as small spines either surpassing rear
contour (d) or not (9). Epiproct long, distinctly flattened dorsoventrally, in lateral
view almost straight, from above coniform, rather narrowly truncated, with preapical
incisions quite distinctly removed from tip. Subanal scale roundly subtriangular, with
a pair of indistinct, setiferous, paramedian knobs at a rather narrow caudal margin.
Sterna scarcely setose, strongly shining, with a big, linguiform, setose lamina
between d coxae 4 (Fig. 72). Legs long, similarly slender in both sexes, rather
sparsely setose ventrally, without adenostyles.
Gonopods (Figs 73-75) subfalcate, typical for the genus. Base of free soleno-
merite with a very little, simple, mesal tooth. Tip of tibiotarsus characteristically
shaped.
Kalimantanina bruneica sp. n. (Figs 76-81)
Material: HOLOTYPE d (MHNG), Borneo, Brunei, Belait Distr., “Andulau Forest
Reserve”, 3.5 km S of Sungaim Liang (ca. 39.5 km from Labi), mixed dipterocarp forest, “K-7”
beaten from vegetation, 19.X1.1988 (Bru-88/22); leg. C. Lienhard. — Paratypes: 1 d (MHNG),
Belait Distr., near Sungai Mau, ca. 14 km along road from Sungai Liang to Labi (1.5 mi along
secondary road toward Badas), ca. 50 m, beaten from vegetation, 16.X1.1988 (Bru-88/9). — 1 d
(ZMUM), Belait Distr., “Badas Forest Reserve”, ca. 10 km along secondary road branching 32
176 SERGEI I. GOLOVATCH
km of Kuala Belait toward S, 10 m, “Kerangas” forest (= tropical heath forest) mostly of
Agathis dammara (Lambert) L.G.Rich. (Araucariaceae), under bark, 23.X1.1988 (Bru-88/36);
all leg. C. Lienhard.
Other material tentatively attributed to this species: 1 2 (MHNG), Borneo, Brunei,
Tutong Distr., little market on road coming from Muara, 77 km from Kuala Belait, ca. 40 m,
beaten from vegetation, 18.X1.1988 (Bru-88/17); leg. C. Lienhard.
Etymology: The species epithet refers to the terra typica.
Diagnosis: Differs from congeners by the generally pallid coloration combined
with the peculiar shape both of the tooth at base of the solenomerite and the gonopod
tibiotarsus.
DESCRIPTION
Length of dd ca. 25-26, width of midbody pro- and metazona 1.7-1.8 and
2.0-2.1 mm, respectively. Coloration entirely pallid to pale yellowish-brown, devoid
of a pattern, only distal half of antennomere 6 and most of antennomere 7 sometimes
dark brown to blackish.
Head a bit broader than collum, subequal to somites 2 and 4, a little broader than
narrowest somite 3 but a bit narrower than somites 5-18, onward body rapidly tapering.
Antennae relatively short, slightly clavate, in situ scarcely reaching to end of somite 3
in d. Paraterga modestly developed, much like in K. crucifera, somewhat thicker on
pore-bearing somites, set rather low, largely at 1/2-1/3 midbody height, caudally
invariably gently rounded and lying mostly within rear tergal contour (Fig. 76), only on
somite 19 just a bit surpassing it and more strongly dentiform; on somite 2 anterior
corner with a minute sharp denticle and caudal corner strongly rounded and somewhat
surpassing rear tergal contour. Ozopores lateral, lying near caudal corner of paraterga in
an indistinct groove (Fig. 76). Surface shining, generally smooth throughout, slightly
microgranulate only below paraterga 2-4. Somites strongly constricted, suture dividing
pro- and metazona deep, strongly ribbed longitudinally. Transverse metatergal sulcus
starting from somite 5, deep, thin and lineiform, missing on somites 18-19. Tergal setae
minute, mostly missing, a few on collum, pattern untraceable. Pleurosternal keels
preserved only as poorly developed teeth lying within rear contour, present only on
somites 2-4 and 6, virtually absent from somites 5, 7 and so on. Epiproct long,
distinctly flattened dorsoventrally, in lateral view almost straight, from above coniform,
rather broadly truncate, with preapical incisions very poorly developed and placed close
to tip (Fig. 77). Subanal scale high, subtrapeziform, with a pair of indistinct, setiferous,
paramedian knobs at a rather narrow caudal margin.
Sterna scarcely setose, strongly shining, with a big, trapeziform, setose lamina
between d coxae 4 (Fig. 78). Legs long, similarly slender in both sexes, rather
sparsely setose ventrally, without adenostyles.
Gonopods (Figs 79-81) subfalcate, typical for the genus. Base of free
solenomerite with a little, very slightly curved, serrate, mesal tooth. Tip of tibiotarsus
characteristically shaped.
Remarks: The genus Kalimantanina Jeekel, 1963, has hitherto been known to
comprise five species, all confined to Borneo: K. ruficeps Jekel, 1963 (the type-
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 177
Fics 71-81
Kalimantanina crucifera sp. n., 4 holotype (71-75), and Kalimantanina bruneica sp. n., &
holotype (76-81): 71 & 76 — somite 10, lateral and dorsal views, resp.; 72 & 78 — sternal lamina
between d coxae 4, caudal view; 73-75, 79-81 — gonopods, mesal, lateral, ventral, mesal,
sublateral, and subdorsal views, resp.; 77 — epiproct, dorsal view; — Scales in mm: 0.35 (76-77),
0.25 (78-80), 0.12 (81).
species), K. albonigra Jeekel, 1963, K. decolorata Jeekel, 1963, K. hirtitarsus Jeekel,
1963, and K. ocellata Jeekel 1963 (JEEKEL 1963). Both K. crucifera and K. bruneica
are quite typical congeners, failing to broaden the concept of the genus concerned.
178 SERGEI I. GOLOVATCH
All currently known Kalimantanina species can be separated using the
following key:
1(6) Colour pattern present, distinct on body somites.
2(3) Body lesser: width 2.0 (4) to 2.6 mm (9). Only paraterga 19 as a small
denticle projecting caudally beyond rear tergal contour. Tarsal brushes
missing. Transverse metatergal sulcus wanting only on somite 19. Pro-
cess at base of solenomerite particularly small, dentiform (Figs 73, 75).
Re pt I LINA K. crucifera
3(2) Body bigger, width closer to 3-3.5 mm (d). Caudal denticles on para-
terga 18-19 lying within rear tergal contour. Tarsal brushes present.
Transverse metatergal sulcus wanting on somites 18-19. Gonofemoral
process at base of solenomerite bigger.
4(5) White area dorsally of paraterga only half as wide as paraterga. Gono-
femoraliprocessilanceolate FERRO K. ocellata
5(4) White area dorsally of paraterga about as wide as paraterga. Gono-
FeImoORalDROCESSSSPINILOLNE EEE RO K. albonigra
6(1) Coloration more or less strongly uniform, pallid to very dark, some-
times only head and anal valves of a contrasting colour.
7(8) Body lesser: width 2.0-2.1 mm (6). Tarsal brushes missing. Pleuro-
sternal keels only on somites 2-4 and 6. Gonofemoral process little,
Sematen(Fres 79-8] RR OR K. bruneica
8(7) Body bigger: width at least 2.5 mm (d). Tarsal brushes present on most
d legs. Pleurosternal keels on somites 2-7, sometimes vestigial also on
somite 8. Gonofemoral process bigger.
9(10) Background coloration almost entirely pallid. Sternal lamina between
dé coxae 4 distinctly bilobed. Paraterga 18-19 caudally dentiform and
slightly surpassing rear tergal contour. Gonofemoral process arising
welliproximally of base of solenomenite:.. 4-54 sera K. decolorata
10(9) Background coloration dark to very dark, blackish, with head and anal
valves contrastingly reddish-brown. Sternal lamina between 4 coxae 4
at most scarcely emarginate. Caudal corner of paraterga 18-19 lying
within rear tergal contour. Gonofemoral process arising more distally,
just at base of solenomerite.
11(12) Lesser: width 2.5-2.6 (6) and 3.2-3.4 mm (2). Tarsal brushes absent
from d legs of posterior body half. Gonofemoral process arising from
Solenomerite sos near Bes Ara Oe) ee ee eee K. ruficeps
12(11) Bigger: width 3.1 mm (4). Tarsal brushes absent only from two last d
legpairs. Gonofemoral process arising just proximally of base of sole-
NOmenitesti. s patio NME RER AR OR K. hirtitarsus
Opisthodolichopus scandens Hoffman, 1973 (Figs 82-85)
Material: 1 8,2 2 (MHNG), Singapore: Bukit Timah Nature Reserve, from Summit
Hut to Simpang Hut along North View Path, Jungle Fall Path and Fern Valley Contour Path,
90-140 m, beaten from vegetation, 17.XII.1987 (Sar-87/97): leg. C. Lienhard.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 179
Remarks: The above strictly topotypical material fully corresponds to the nice
original description (HOFFMAN 1973), as can be seen from the new illustrations (Figs
82-85). This species has hitherto been known only from Singapore and is believed to
represent a true arboricole.
In addition to a new congener described just below, the genus Opisthodo-
lichopus Verhoeff, 1941, embodies four acknowledged species: O. javanicus (Attems,
1903), from Java, O. nigricornis (Pocock, 1894) and O. thienemanni (Attems, 1930)
(the type-species), both from Sumatra, as well as O. scandens Hoffman, 1973, from
Singapore (JEEKEL 1968; HOFFMAN 1973). Therefore, below is the first record of an
Opisthodolichopus species in Borneo. The new illustrations of O. scandens are
provided here for comparative purposes.
Opisthodolichopus bifidus sp. n. (Figs 86-89)
Material: HOLOTYPE d (MHNG), Malaysia, Borneo, Sarawak: Bau, near Cave “Gua
Puang” near Kampong Pelaman Sekiang on a steep slope of Gunung Jambusan (“Pang Cave”
in Wilford, 1964), 10-30 m, beaten from vegetation, 4.XII.1987 (Sar-87/51); leg. C. Lienhard.
— Paratype: d (MHNG), Sarawak: Serian District, Penrissen Road 12 miles from Kuching,
Semongok Wildlife Rehabilitation Centre: Nursery Centre of the Forest Department, 20-50 m,
8.X11.1987 (Sar-87/61); leg. B. Hauser.
Diagnosis: Differs from congeners by the distinctly and deeply bifid gonopod
tibiotarsus as well as by the considerably shorter gonofemorite.
DESCRIPTION
Length ca. 23-26 mm, maximal width 1.9-1.95 and 2.1-2.2 mm on midbody
pro- and metazonites, respectively. Coloration contrasting, being blackish dorsally
down to level of paraterga and yellow-whitish below; head, almost entire antennae,
epiproct and tips of tarsi pale brownish to brown; tip of antenna whitish, most of
antennomere 7 blackish.
Head a bit broader than somite 5 and subequal in width to somite 7, much
broader than subequally wide collum and somite 2 and, especially, narrowest somites
3-4. Body parallel-sided on pore-bearing somites (from 7th until 17th), caudally
tapering. Antennae relatively long and slender, in situ extending beyond somite 4
dorsally. Paraterga very poorly developed, invariably low (at about body midheight),
as a narrow ridge within rear contour only on somite 2, as very poorly developed
carinae on somites 3-4, onward as bulges (pore-bearing) or thin (poreless), hardly
traceable, vestigial ridges outlined only dorsally due to a sulcus, gradually reducing in
size on somites 17-19, caudally always rounded and lying well within rear tergal
contour (Fig. 86). Ozopores big, lateral. Surface delicately shagreened almost
throughout, shining, polished only on metaterga, slightly rugulose only near fore
margin of collum and more coarsely shagreened below paraterga 2-4. Only collum
with some traces of an axial line. Somites not very deeply constricted, suture dividing
pro- and metazona relatively shallow, strongly longitudinally striate dorsally and
dorsolaterally. Transverse metatergal sulcus starting as barely traceable from somite
180 SERGEI I. GOLOVATCH
FIGs 82-89
Opisthodolichopus scandens Hoffman, 1973, & (82-85), and Opisthodolichopus bifidus sp. n., &
paratype (86-89): 82 & 86 — somite 10, dorsal view; 83 & 87 — sternal lamina between d coxae 4,
caudal view; 84-89 — left gonopod, mesal, lateral, mesal, and lateral views, resp. — Scales in mm.
5, onward more deep, thin, lineiform, rather short, far from reaching to bases of
paraterga, missing on somites 18-19. Tergal setae short, mostly broken off, a few on
collum and 1+1 on somite 2, setation pattern perhaps 1+1 or 2+2 in a presulcus row.
Pleurosternal carinae as a lappet on somite 2, an oblique low ridge on somites 3-4, an
incomplete ridge hardly traceable only caudally on somites 5-6, onward totally
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 181
missing, never extending beyond rear contour. Epiproct long, flattened dorsoventrally,
in lateral view almost straight, from above rather broadly truncate, distal half parallel-
sided, preapical incisions distinct and placed close to tip, apical papillae indistinct.
Subanal scale semicircular, with a pair of setiferous, paramedian knobs at a convex
caudal margin.
Sterna moderately densely setose, without modifications, with a prominent,
linguiform, setose, apically slightly emarginate lamina directed mainly ventrally
between coxae 4 (Fig. 87). Legs long, slender, gradually growing in length toward
caudal body end, perhaps a little incrassate, with neither adenostyles nor tarsal
brushes, setation generally sparse to modest, mainly ventral, somewhat more dense
only on coxa, prefemur and tarsus, last two especially long and more strongly
enlarged.
Gonopods (Figs 88-89) simple, elongate. Coxite rather long, distoventrally
setose. Prefemoral part little, densely setose as usual. Femorite relatively short,
subcylindrical, folded mesally near base. Both tibiotarsus and solenomerite long and
slender, subequal in length, former deeply bifid.
Remarks: The deeply bifid gonopod tibiotarsus coupled with the relatively
short gonofemorite make the above new species disjunct among congeners. Such a
pattern of somewhat more strongly isolated congeners confined to Borneo has already
been observed in some other paradoxosomatid genera, e.g. Gigantomorpha, Dajakina
(JEEKEL 1963).
Superficially, judged from a strongly bifid gonopod tibiotarsus, O. bifidus
somewhat resembles Antheromorpha Jeekel, 1968, Dajakina as well as some other
Oriental genera of the tribe Orthomorphini. However, O. bifidus differs by the
completely mesal position of the solenomerite and entirely lateral tibiotarsus charac-
teristic of a sundaninine.
All currently known Opisthodolichopus species can be separated using the
following key:
1(2) Larger: width 3.3 mm. Pleurosternal keels traceable only on somites 2-4
5 6 oO LE TEIN È IR Scie ee el en MITE D III RIDIRE O. nigricornis
2(1) Smaller: width at most ca. 2.5 mm. Pleurosternal keels traceable at least
until somite 6
3(4) Live coloration carmine-red, in ethanol bleached white with brownish
middorsal areas. Pleurosternal keels until 4 somite 7. Gonopods as in
OSB SI re ee Bene O. scandens
4(3) Coloration in ethanol always contrasting, being dark blackish-brown
dorsally and pinkish to white on sides and ventrally. Pleurosternal keels
either until somite 6 or until somite 8. Gonopods different, especially tibiotarsus.
5(6) Pleurosternal keels extending back to d somite 8. Gonopod tibiotarsus
sienderslaminate JAVA. IAA MOLO: O. javanicus
6(5) Pleurosternal keels extending back only to d somite 6. Gonopod tibio-
tarsus much more stout, either strongly bifid or broadly laminate.
182 SERGEI I. GOLOVATCH
7(8) Sides and ventrum in ethanol pinkish. Gonopod tibiotarsus broadly
AMIN A (CES UAAR ie RAM OS IS ATE RE O. thienemanni
8(7) Sides and ventrum in ethanol whitish. Gonopod tibiotarsus deeply bifid.
BOTTONE VANNES AMEN ale MASE, A US, SEG See O. bifidus
Arthrogonopus proletarius sp. n. (Figs 90-93)
Material: HOLOTYPE d (NHMB), Indonesia, Borneo, Kalimantan Barat: Gunung
Palung National Park, Caban Panti Research Site (1°13'S, 110°7'E), primary lowland rainforest,
under bark, 20. VII.1993; leg. O. Merkl.
Diagnosis: Differs from congeners by the caudally dentate paraterga combined
with the relatively well-developed pleurosternal carinae (present as knobs even on
somite 8) as well as the pointed, non-branching and long gonopod tibiotarsus.
DESCRIPTION
Length ca. 38 mm, maximal width 2.5 and 3.0 mm on midbody pro- and meta-
zonites, respectively. Coloration generally blackish-brown; a wide, moniliform, yello-
wish, axial stripe on dorsum extending to epiproct (especially wide and bright on
prozonites and rear half of metazonites, more brownish on frontal half of meta-
zonites); antenna black, only tip whitish; head, paraterga, suture dividing pro- and
metazonites as well as ventrum and a few distal podomeres (mostly postfemur, tibia
and tarsus, more closely to telson also femur) pale brownish; proximal podomeres
whitish-yellow; tibiae and tarsi of last two legpairs considerably infuscate as com-
pared to those of preceding legs.
Head a bit narrower than subequally wide somites 5-16, but broader than
collum and, especially, narrowest somites 3-4. Body gently and gradually tapering
caudad on somites 18-20. Antennae very long and slender, in situ extending beyond
somite 5 dorsally. Paraterga very poorly developed, invariably low, as a narrow ridge
extending caudally beyond rear contour only on somite 2, onward as thick (pore-
bearing) or thin (poreless) ridges outlined only dorsally due to a sulcus, gradually
reduced in size on somites 17-19, caudally almost beak-shaped, very narrowly
rounded from somite 5 (Fig. 90). Ozopores big, lateral. Surface delicately shagreened
throughout, generally dull, poorly shining only on metaterga, rugulose only below
paraterga, some metaterga with traces of an axial line. Somites deeply constricted,
suture dividing pro- and metazona deep, strongly striate longitudinally. Transverse
metatergal sulcus stalting from somite 5, rather deep, thin and lineiform, at bottom
vaguely beaded, missing on somites 18-19. Tergal setae short, mostly broken off, a
few on collum, pattern perhaps 2+2 in a presulcus row. Pleurosternal carinae as strong
caudal teeth extending beyond rear contour on somites 2-6, as a moderately strong
tooth lying within the contour on somite 7, as a little knob on somite 8, onward
virtually wanting. Epiproct long, flattened dorsoventrally, in lateral view almost
straight, from above rather broadly truncate, distal half parallel-sided, preapical
incisions distinct and not greatly removed from tip. Subanal scale subtrapeziform,
with a pair of setiferous, paramedian knobs at a very poorly convex caudal margin.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 183
Fics 90-93
Arthrogonopus proletarius sp. n., d holotype: 90 — somite 10, dorsal view; 91 — sternal lamina
between d coxae 4, caudal view; 92-93 — right gonopod, mesal and lateral views, resp. — Scales
inmm.
Sterna poorly setose, without modifications, with a big, trapeziform, setose
lamina directed mainly ventrally between coxae 4 (Fig. 91). Legs long, slender, with
neither adenostyles nor tarsal brushes, setation sparse, mainly ventral, somewhat more
dense only on coxa, prefemur and tarsus.
Gonopods (Figs 92-93) subfalcate. Coxite rather long, ventrally setose. Pre-
femoral part small, densely setose as usual. Femorite rather long, folded laterally and,
especially, mesally; postfemoral portion set off due to well-developed lateral sulcus,
with a couple of little knobs distoventrally. Both tibiotarsus and solenomerite long
and slender, subequal in length, pointed.
Remarks: The genus Arthrogonopus Jeekel, 1963, has hitherto been considered
as being endemic to Borneo, with five species involved: A. edentulus Jeekel, 1963
(the type-species), A. denticulatus Jeekel, 1963, A. borneonus (Attems 1931), A.
bifidus Jeekel, 1963, and A. vicinus Jeekel, 1983b (JEEKEL 1963, 1983b). The above
new species is a typical congener, failing to alter the generic diagnosis.
The record of A. borneonus (as Orthomorpha borneona Attems, 1930, sic!) in
Singapore by WANG & TANG (1965) is dubious and most probably erroneous, reason
why JEEKEL (1968) as usual ignored it altogether (see also above and below).
184 SERGEI I. GOLOVATCH
Eventually, WAnG & TANG (1965) could well have dealt with Opisthodolichopus
scandens Hoffman, 1973, the only species of the tribe Sundaninini subsequently
reported from Singapore (see also above).
All currently known Arthrogonopus species can be separated using the
following key:
1(2) Coloration of two last legpairs not differing from that of other legs.
Gonopod tibiotarsus with a big, laminate, midway, distolateral process.
ee ei eee TP i en ai ae at) A o on oc À. bifidus
2(1) Last two legpairs contrastingly infuscate at least distally of middle of
tibiae, more proximally yellowish-brown, remaining legs brown only in
distal halves of tarsi. Gonopod tibiotarsus devoid of a considerable process.
3(6) Even 6 poriferous paraterga largely fully and more or less strongly
rounded caudally, at most subangular, never beak-shaped even on pos-
teriormost somites.
4(5) Pleurosternal keels present on somites 2-8. Gonopod femorite and
postfemoral part each with a distoventral knob............... A. borneonus
5(4) Pleurosternal keels only on somites 2-6(7). Gonopod femorite, not post-
femur withoutdistoventraliknob RE ee eee A. edentulus
6(3) Atleast a few posteriormost paraterga dentiform caudally (Fig. 90).
7(8) Paraterga relatively poorly developed, growing subangular only toward
posterior somites, distinctly angular only on somites 17-18. Pleuro-
sternal keels traceable only until 4 somite 6. Gonopod tibiotarsus
always dévoidiora distal. tooth; acuminate te =... ea eee A. vicinus
8(9) Paraterga subdentiform caudally at least from somite 15 onward. Pleu-
rosternal keels present also also on d somite 7. Gonopod tibiotarsus
sometimes rounded apically and bearing a distal tooth.
9(10) Paraterga strongly angular, almost dentiform from somite 5 onward (Fig.
90). Pleurosternal keels traceable until d somite 8. Gonopod tibiotarsus
acuminate at end, devoid of a distal tooth (Figs 92—93)......... A. proletarius
10(9) Paraterga produced into minute denticles caudally on somites of posterior
body half. Pleurosternal keels traceable only until d somite 7. Gonopod
tibiotarsus with a rounded apical lappet and a distal tooth....... A. denticulatus
Margaritosoma borneorum sp. n. (Figs 94-101)
Material: HOLOTYPE ¢ (MHNG), East Malaysia, Borneo, Sabah, Interior Residency,
Kimanis road, 16 mi from Keningau, heliport, 1400 m, mossy forest, beaten from vegetation,
14.11.1983 (Pal-83/14); leg. C. Lienhard. — Paratypes: 1 2 (MHNG), same locality, together
with holotype. — 1 4 (MHNG), Sabah, Interior Residency, Kimanis road, 10 mi from
Keningau, 1170 m, secondary forest, beaten from vegetation, 13.V.1982 (82187). — 1 2
(MHNG), Sabah, Interior Residency, Kimanis road, 16 mi from Keningau, heliport, 1380-1400
m, mossy forest, beaten from vegetation, 12.V.1982 (82178); all leg. D. Burckhardt. — 1 4,1 2
(MHNG), 1 é (ZMUM), Borneo, Brunei, Belait Distr., “Badas Forest Reserve”, ca. 10 km
along secondary road branching 32 km of Kuala Belait toward S, 10 m, “Kerangas” forest (=
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 185
tropical heath forest) mostly of Agathis damnara (Lambert) L.G. Rich. (Araucariaceae), under
bark, 23.X1.1988 (Bru-88/36). — 2 22 (MHNG), Borneo, Brunei, Belait Distr., “Andulau
Forest Reserve”, 3.5 km S of Sungaim Liang (ca. 39.5 km from Labi), mixed dipterocarp forest,
“K-7”, beaten from vegetation, 19.X1.1988 (Bru-88/22). — 1 2 (MHNG), Borneo, Brunei, Belait
Distr., “Labi Hills Forest Reserve”, “Teraja”, 42 km S of Sungai Liang (= 12 km S of Labi),
near “Rumah Panjang” (= Longhouse of Kampong Teraja), mixed dipterocarp forest, 40 m,
beaten from vegetation, 22.X1.1988 (Bru-88/33); all leg. C. Lienhard.
Etymology: The species epithet emphasizes the first congener encountered on
Borneo.
Diagnosis: Differs from congeners both by the especially poorly developed
paraterga and by the shape of the gonopod tibiotarsus.
DESCRIPTION
Length ca. 11-12 (6) and 14-15 mm (@), maximal width of midbody somites
0.85-0.95 (3d) and 1.0-1.05 mm (2). Coloration uniform red- to chocolate-brown,
body caudally somewhat more pale; only tip of antennae, suture between pro- and
metazona, and legs whitish to yellowish.
Head a little broader than subequally wide collum and midbody somites, post-
collar constriction slight on subequally narrowest somites 2-3. Body distinctly moni-
liform, parallel-sided on somites 5-16, onward very gently and gradually tapering.
Antennae moderately long, slightly clavate, in situ reaching to end (4) or midlength (9)
of somite 4. Collum with a lateral rim. Paraterga very weakly displayed as small and low
ridges slightly delimited from above and below on somites 2-4 (Fig. 94), onward entirely
missing. Ozopores lying very low, surmounting small swellings (Fig. 95). Surface
shining, smooth. Somites strongly constricted, suture dividing pro- and metazona deep,
extremely strongly striate longitudinally. Metatergal sulcus virtually missing, only
sometimes poorly traceable on somites 2-3 (2). Axial line wanting. Tergal setae rather
short to medium-sized, a few on collum, onward a pattern of 2-3+2-3 and sometimes
also 2-3+2-3 setae discarding a peculiar caudal seta level to ozopore on each side (Fig.
95). Setae of rear row often incomplete to virtually missing (especially so in Brunei
samples). Pleurosternal carinae strong and dentiform on somites 2—3, weak on somite 4,
onward obsolete. Epiproct medium-sized, somewhat flattened dorsoventrally, narrowly
truncate at apex, lateral incisions strongly removed from tip. Subanal scale trapeziform,
with a pair of rather distinct, setiferous, paramedian knobs at caudal corners.
Sterna usual, without modifications, sparsely to modestly setose, strongly
shining, with a big, transverse, setose, deeply notched lamina directed ventrad be-
tween d coxae 4 (Fig. 96). Legs rather long, somewhat incrassate and more long in d
as compared to 9, with neither adenostyles nor tarsal brushes.
Gonopods (Figs 97-101) rather stout. Coxite not too long, distoventrally
setose. Telopodite without evidence of torsion, prefemoral part rather prominent,
femorite with a mesal groove and a more (Sabah) or less (Brunei) strongly rounded,
dorsal, midway lobe. Lamina medialis of tibiotarsus unciform apically, with a small
simple tooth well removed from a more or less slender tip. Lamina lateralis little,
ribbon-shaped, more or less strongly bent. Distolateral process of tibiotarsus strong,
simple, more or less ribbon-shaped, bent subapically. Seminal groove entirely mesal.
186 SERGEI I. GOLOVATCH
Margaritosoma singaporense sp. n. (Figs 102-105)
Material: HOLOTYPE d (MHNG), Singapore, “Island Country Club”, between Lower
Peirce Reservoir and Windsor Park Estate along Upper Thomson Road, remains of primary
forest surrounded by secondary forest, ca. 60 m, beaten from vegetation, 12.X1.1988 (Bru-
88/2); leg. C. Lienhard.
Etymology: The species epithet refers to the terra typica.
Diagnosis: Differs from congeners both by the relatively poorly developed,
distinctly incised paraterga and by the peculiar shape of the gonopod tibiotarsus.
DESCRIPTION
Habitus much like in M. borneorum, but differing as follows. Length ca. 11-12
mm, maximal width 0.3 mm on midbody somites. Coloration uniform pale brown,
anterior third of body a little brighter, only legs and suture dividing pro- and metazona
whitish, and antennomere 7 a little infuscate.
Head subequal in width to somites 5-15, a bit broader than subequally wide
collum and somites 2-4. Body distinctly moniliform, very gradually and gently
tapering caudad on somites 16-20. Paraterga almost entirely missing, very weakly
displayed as small and low ridges on somite 2, hardly traceable due to outlining sulci
from above and below only on somites 3-4, onward virtually missing, preserved only
as poor swellings. Paraterga 2-3 with three, paraterga 4 with two distinct, lateral,
setiferous incisions. Ozopores very low, surmounting the swellings. Transverse
metatergal sulcus entirely missing. Tergal setae relatively long, ca. 1.5 times as long
as in M. borneorum; pattern much like in M. borneorum as well, a few on collum,
onward 3-4+3-4 and 4+4 setae in two rows, discarding 1-2 peculiar setae behind
ozopore until somite 7. Pleurosternal keels vestigial on somites 2-4, onward absent.
Epiproct rather long, from above almost parallel-sided, broadly subtruncate, with
preapical incisions well-developed and rather strongly removed from tip. Subanal
scale low, broadly trapeziform, with a pair of indistinct, setiferous, paramedian knobs
at a wide and straight caudal margin.
A big, subtrapeziform, setose lamina directed obliquely anteventrad between
3 coxae 4 (Fig. 102).
Gonopods (Figs 103-105) less strongly curved, midway dorsal lobe on femo-
rite almost wanting. Lamina medialis with a prominent rounded lobe at basal 1/3,
distolateral process of tibiotarsus big and scapuliform.
Remarks: WANG (1967b) once reported Tectoporus filum (Silvestri) from
Singapore, a species originally very poorly described from Sumatra and presently
known to be the generotype of Margaritosoma Jeekel, 1979, only since its relatively
recent redescription (JEEKEL 1979a). Why that species had been reallocated in Tecto-
porus by WANG & TANG (1965), remains totally obscure, the most likely reason why
JEEKEL (1968, 1979a) simply ignored both record and transfer altogether. What seems
more important, however, is that WANG's discovery could have well belonged in fact
to M. singaporense. Hence the entire situation strongly reminds of WANG's records of
Orthomorpha and/or Gigantomorpha in Singapore (see above).
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 187
Ay
N Ve
DS 99
Fics 94-105
Margaritosoma borneorum sp. n. (94-101), 8 6 paratypes from Sabah (94-98) and Brunei (99-
101), and Margaritosoma singaporense sp. n. (102-105), 4 holotype: 94 — anterior body
portion, lateral view; 95 — somite 10, lateral view; 96 & 102 — sternal lamina between d coxae
4, caudal view (96 drawn not to scale); 97-101, 103-105 — gonopods, mesal, lateral, mesal,
lateral, mesal, lateral, ventral, and ventral views, resp. — Scales in mm.
The genus Margaritosoma Jeekel, 1979a, has hitherto been known to comprise
four species: M. filum (Silvestri, 1895) (the type-species), from Sumatra, M. nanum
(Silvestri, 1895), from Enggano, M. sequens (Chamberlin, 1945), from Java, and M.
baliorum Golovatch, 1994b, from Bali (JEEKEL 1979a; GOLOVATCH 1994b). Besides,
JEEKEL (1979a) mentioned one more, yet undescribed congener from Padang,
Sumatra, suggesting the genus be much more prolific. The above two new species
from Borneo and Singapore confirm that statement to some extent, both are typical
members virtually failing to broaden the concept of the genus concerned.
188 SERGEI I. GOLOVATCH
All currently known Margaritosoma species can be separated using the
following key:
1(2) Transverse sulcus on metaterga present, usual. Paraterga absent, with
striae in their stead. Distolateral process of gonopod tibiotarsus
especially slender, a little longer than solenophore. Java......... M. sequens
2(1) Transverse sulcus on metaterga virtually absent. Somites 5-15 without
any traces of paraterga. Distolateral process of gonopod tibiotarsus
shorter than solenophore.
3(4) At least most d sterna with a little cone near coxa, lamina between d
coxae 4 fully bipartite. Gonopod tibiotarsus relatively little, only ca. 1/3
asslongrasitemornte Bali "#7. AE M. baliorum
4(3) Sterna without modifications, lamina between d coxae 4 always as a
single piece, usually lingui- to subtrapeziform. Gonopod tibiotarsus
always bigger and longer, closer to 1/2 as long as femorite.
5(6) Tergal setae especially numerous, largely 3+3 and 8+8 in two trans-
verse rows. Suture dividing pro- and metasomites not especially strong-
ly ribbed. Distolateral process of gonopod tibiotarsus oblong blade-like.
ST TANOME Peres ec ee een ty N A ie RIO ON M. nanum
6(5) Tergal setae less numerous. Suture dividing pro- and metasomites very
strongly ribbed (Fig. 95). Distolateral process of gonopod tibiotarsus
either as a very broad lobe or ribbon-shaped.
7(8) Head considerably broader than collum. Tergal setae largely 4(5)+4(5)
and 6+6 in two transverse rows. Sternal lamina between d coxae 4
high, linguiform. Tarsal brushes present until two last 5 legpairs.
SUV TOE es ek N ee A A LR EEE M. filum
8(7) Head only a little broader than collum. Tergal setae less numerous. Sternal
lamina neither too high nor linguiform. Tarsal brushes virtually absent.
9(10) Paraterga 2-4 quite strongly incised. Tergal setae largely 3(4)+3(4) and
4(5)+4(5) in two transverse rows. Sternal lamina between d coxae 4
subtrapeziform (Fig. 102). Distolateral process of gonopod tibiotarsus
scapulitonmi (1951035105) SINSApore PECE M. singaporense
10(9) Paraterga 2-4 only with very vague incisions at lateral margin. Tergal
setae usually either as 2(3)+2(3) and 3(4)+3(4) in two rows (Fig. 95) or
rear row up to (almost) entirely reduced. Sternal lamina between d
coxae 4 strongly emarginate apically (Fig. 96). Distolateral process of
gonopod tibiotarsus ribbon-shaped (Figs 97-101). Borneo... . . M. borneorum
KEY TO PARADOXOSOMATID TRIBES AND GENERA ENCOUNTERED ON BORNEO, WITH A
REVIEW OF THE BORNEAN FAUNA:
1(2) Body strongly polydesmoid; paraterga very prominent, devoid of lateral
calluses; ozopores located below paraterga. Males yet unknown
RR Dp ah N Sel, AE A APR ae ARS eee Tribe Centrodesmini
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 189
Genus Pleuroporodesmus Jeekel, 1983a. Only a single species from East
Borneo is known: P. serrulatus Jeekel, 1983a.
2(1) Body not so strongly polydesmoid; when paraterga prominent, lateral
calluses always strongly developed; ozopores always on paratergal
calluses. Males known.
3(6) Gonopod prefemorite hypertrophied; solenomerite not flagelliform,
very thick and rigid, completely or mostly free from solenophore-like
SEUCIUTO Salita EINE DB EEE RER PARENT er Tribe Antichiropodini
4(5) Seminal groove passing onto solenomerite on dorsal side of distalmost
remontes (es dd) PEER en RME nee Genus Borneochiropus n.
A single species from East Borneo.
5(4) Seminal groove passing onto solenomerite on ventral side of distalmost
femorite (Figs 16, 24,30, 34). ........... Genus Euphyodesmus Attems, 1931
Seven species, all from West or East Borneo (see key above).
6(3) Gonopod prefemorite never hypertophied, largely usual; solenomerite
always flagelliform, slender, mostly closely attached to or sheathed by
a well-developed solenophore (= tibiotarsus).
7(8) Gonopod tibiotarsus with a big additional lateral lobe or process (Figs
97-101). Body devoid of paraterga from somite 5 on (Fig. 95)
RE SE ve Genus Margaritosoma Jeekel, 1979a
A single and obviously disjunct genus of the tribe Sulciferini yet encountered
on Borneo, the above being only the generic diagnosis. Only one species has been
recorded in West and Fast Borneo, several others are widely scattered in the Sunda
area (see key above).
8(7) Gonopod tibiotarsus without such outgrowths. Body always with para-
terga also on somites 5-19.
9(16) Solenomerite fully mesal, attached to but not sheathed by a completely
lateral solenophore. Paraterga always poorly developed. . . . Tribe Sundaninini
10(13) Gonopod femorite devoid of a marked distal process, at best only poor
wrinkles in its place.
11(12) Distal part of gonofemorite with a prominent lateral sulcus marking a
postfemoral portion, latter with wrinkles distoventrally (Figs 92-93)
Do oo GR E SETS ER ACI a SI Genus Arthrogonopus Jeekel, 1963
Six species, all from Borneo (see key above).
12(11) Neither sulcus demarcating a gonopostfemoral portion nor traces of a
distofemoral outgrowth (Figs 88-89). Genus Opisthodolichopus Verhoeff, 1941
A single species yet encountered on Borneo (see key above).
13(10) Gonopod femorite with a more or less prominent process at or near
base of solenomerite.
14(15) Gonopod suberect, tibiotarsus shorter than femorite, curved slightly
CURO: Bey cg eee apc ae ER da Genus Borneonina Jeekel, 1963
A single species from West Borneo: B. retrorsa Jeekel, 1963.
15(14) Gonopod subfalcate, especially so due to a long and ventrally curved
solenophore (Figs 73-75, 79-81)........ Genus Kalimantanina Jeekel, 1963
190 SERGEI I. GOLOVATCH
Seven species, all for East and West Borneo (see key above).
16(9) Solenomerite strongly to almost fully sheathed by solenophore, latter
consisting of a lamina medialis and a lamina lateralis. Paraterga often
very well-developed, subhorizontal.................. Tribe Orthomorphini
17(20) Paraterga very well-developed, mostly subhorizontal, close to level to
dorsum.
18(19) Paraterga with moderately strong lateral calluses (Figs 56-57). Gono-
pod femorite with 1-2 distinct, dorsal, parabasal folds; tip of tibiotarsus
with a laminate or spiniform outgrowth (Figs. 59-62)
Sd O Bp PLOT LA O A E de ee Ce Genus Gigantomorpha Jeekel, 1963
Ten species, of which eight occur on Borneo (see key above).
19(18) Gonopod femorite without dorso-parabasal folds; tip of tibiotarsus as
2 SME APPESA cr. nn Genus Orthomorpha Bollman, 1893
Two species have hitherto been formally reported from Borneo: O. beaumontii
(Le Guillou, 1841), and O. coarctata (de Saussure, 1860). The former taxon is a big
form deriving from an unidentified Bornean locality and known only from a single ©
specimen (see redescription: JEEKEL 1963), whereas the latter congener is much
smaller, representing a pantropical anthropochore.
20(17) Paraterga poorly developed, sometimes next to obsolete.
21(22) Gonopod tibiotarsus relatively slender, solenomerite beginning very
near to border between femorite and tibiotarsus (Figs 49-53). Gono-
femorite sometimes with a dorso-parabasal process or its traces
ONE VARE i URE ARORA. CRATÙ Genus Dajakina Jeekel, 1963
Five species, of which four occur on Borneo (see key above).
22(21) Gonopod tibiotarsus hypertrophied, solenomerite beginning consider-
ably proximad of border between femorite and tibiotarsus (Figs 49-53).
Gonofemorite always without even traces of a dorso-parabasal process
OME AE À E ci. SIAT A RAI SOM GENS PE SE Genus Orangutana n.
A single species from East Borneo.
In addition, the Bornean paradoxosomatid fauma contains a couple of nomina
dubia. The first of these represents a species only possibly deriving from Borneo
(and/or Sulawesi?): Strongylosoma kuekenthali Attems, 1897. While it is clear now
that the taxon has nothing to do with a true Strongylosoma Brandt, 1833, its identity
remains totally obscure. All previous reallocations in Orthomorpha or Oxidus Cook,
1911 (ATTEMS 1937; JEEKEL 1963, 1968 — to his credit, the latter author always did
so with the necessary qualifications) are bound to be useless as well, since the generic
position of S. kuekenthali can be clarified only upon re-examination of type material.
The same can be said about yet one more Bornean paradoxosomatid, Strongy-
losoma nodulosum Attems, 1897. This species has been described from a single 9
specimen deriving from a closer unknown locality, thus discouraging any revisionary
attempt. Only a study of new abundant materials of Bornean Paradoxosomatidae may
reveal samples best matching the holotype of S. nodulosum, thus finally offering a
solution to the riddle. Yet one must be realistic that eventually this may never happen.
PARADOXOSOMATIDAE FROM SOUTHEAST ASIA 191
ZOOGEOGRAPHICAL NOTES
Having already been revised once in the scope of the entire Bornean fauna
(JEEKEL 1963), the Paradoxosomatidae definitely still remains a highly insufficiently
known millipede family in the entire Australasian area. Eventually, the same concerns
all Diplopoda in general of almost any region of the world, especially tropical. The
current estimates of the real species diversity of the class range between 50,000 to
80,000, of which only some 11-12% seem to have hitherto been described
GOLOVATCH et al. 1995). In other words, with 42 acknowledged species from as many
as 12 genera presently recorded in Borneo, perhaps several times as many para-
doxosomatids can be expected to be ultimately found there.
Of course, the above statement concerns first of all the species level. Already
now a number of genera are quite to highly species-rich on Borneo, e.g., Giganto-
morpha, Kalimantanina, Euphyodesmus, Arthrogonopus. All of such examples but
the first seem to represent true Bornean endemics, while Gigantomorpha can be
regarded subendemic, with the bulk of its constituents being confined to Borneo. A
further increase in the number of species to be yet revealed can be anticipated to be
largely due to such prolific genera. On the other hand, judged from the present
contribution, the number of genera to be discovered on Borneo is likely to at least
double, and not only due to some more pantropical/widespread anthropochores like
Chondromorpha xanthotricha (Attems, 1898), Oxidus gracilis (C. L. Koch, 1847),
Desmoxytes planata (Pocock, 1895), Anoplodesmus saussurei (Humbert, 1965) or
Helicorthomorpha orthogona (Silvestri, 1898).
Yet an outline of Bornean zoogeography as based on the Paradoxosomatidae,
highly preliminary as it is, seems justified. Already JEEKEL (1963) noted that Borneo
supports “a paradoxosomatid fauna which, although having general southeast Asiatic
features, is characterized by a large percentage of endemic genera” (op. cit., p. 205-
206). Discarding a few obvious anthropochores, virtually 100% Bornean paradoxo-
somatid species are endemic there. Zoogeographically, more instructive appear both
generic and tribal levels.
If the quantity of indigenous Bornean species or, to a lesser degree, genera is
sure to increase very considerably along with future explorations, the number of tribes
is more likely to remain more stable and representative. Altogether, five tribes are
currently known to occur on Borneo, with the following constituent genera: Anti-
chiropodini (Euphyodesmus, Borneochiropus), Orthomorphini (Orthomorpha, Gigan-
tomorpha, Dajakina, Orangutana), Sundaninini (Arthrogonopus, Opisthodolichopus,
Kalimantanina, Borneonina), Centrodesmini (Pleuroporodesmus), and Sulciferini
(Margaritosoma). As briefly outlined earlier, the Antichiropodini is a chiefly Aus-
tralian tribe, with only three genera encountered west of West Australia, viz. the
Papuan Haplochiropus and the above Bornean Euphyodesmus and Borneochiropus.
The remaining tribes are certainly Asian/Oriental in origin, with the Centrodesmini
and the Sundaninini clearly centering in the Sunda area. The obviously disjunct
sulciferine Margaritosoma, the orthomorphines Gigantomorpha and the weberi-group
of Orthomorpha, the sundaninine Opisthodolichopus join the examples of endemic
192 SERGEI I. GOLOVATCH
Sunda groups, yet not necessarily strictly Bornean. Hence, together with the truly
endemic Bornean elements, first of all probably from the largely Sunda tribe Sunda-
ninini, the paradoxosomatid fauna of that great island is indeed strongly dominated by
Oriental elements.
What seems more surprising is a relatively pronounced influence of the
Australian realm expressed in the obviously endemic Bornean antichiropodines.
Moreover, at least one of these, Euphyodesmus, appears a quite species-rich genus.
The gap between Borneo and New Guinea, in particular Sulawesi, still harbouring
virtually no paradoxosomatids Australian/Papuan in origin, e.g., representing the
tribes Antichiropodini and/or Eustrongylosomatini, seems to be an artifact, reflecting
the yet highly insufficiently explored faunas of the islands/archipelagos lying in
between rather than a complete absence of Australian/Papuan influence in the Sunda
area east of Borneo. Only further investigations can refute or confirm that. Until then,
the above remains mere speculations.
Similar observations of a mixed nature of the millipede fauna of Australasia,
with strong dominance of the Oriental realm up to New Guinea in the east, have been
reviewed and confirmed recently by SIMONSEN (1990, 1992). Yet there seems to be no
reason whatever to explain the modern distribution patterns of the millipedes of the
Sunda area in terms of continental drift and Gondwana fragmentation since the
Permo-Triassic (criticism of the so-called “mobilistic biogeography” see in: ESKOV &
GOLOVATCH 1986). All currently known evidence of the biogeography of Australasian
Diplopoda seems to point that the gigantic archipelago in question, both Borneo and
New Guinea included, definitely belongs to the Oriental Region, alghough the
reciprocal influence of the Australian realm is more or less strongly traceable there as
well (cp. GRESSITT 1961). As regards the Paradoxosomatidae, the abovementioned
relatively pronounced share of Australian/Papuan components on Borneo suggests an
even greater proportion of such forms further to the east. The little that remains to do
to prove or disprove this opinion, is to adequately describe at least the Sulawesi fauna
which, unfortunately, is still almost unknown.
ACKNOWLEDGEMENTS
I feel greatly obliged both to Dr. Volker Mahnert and Dr. Bernd Hauser, who
helped me during my brief visit and work at the MHNG in November 1993 made
possible through a grant from the Geneva Museum. Both Dr. Hauser (MHNG) and
Dr. Zoltan Korsés (NHMB) kindly sent me material under their care for study. This
work has been supported in part also through a grant of the Russian Academy of
Sciences, Biological Diversity Project, H11.
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Eskov, K. Y. & S. I. GOLOVATCH 1986. On the origin of trans-Pacific disjunctions. Zoologische
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GOLOVATCH, S. I. 1994a. Diplopoda from Nepal Himalayas. Further Paradoxosomatidae (Poly-
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GOLOVATCH, S. I., R. L. HOFFMAN, J. ADIS & J. W. DE MORAIS 1995. Identification plate for the
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REVUE SUISSE DE ZOOLOGIE, 103 (1) : 195-258; mars 1996
Aleocharinae della Sottoregione Africana Orientale al
Museo di Ginevra (Coleoptera, Staphylinidae)
Parte III (conclusione)
Roberto PACE
Via Vittorio Veneto, 13. 37032 Monteforte d'Alpone (Verona), Italia.
Aleocharinae from Eastern African Region in the Geneva Museum
(Coleoptera, Staphylinidae). Part III (conclusion). — In this paper, based
on the study of 70 species of the tribes Thamiaraeini, Myrmedoniini,
Oxypodini, Hoplandriini and Aleocharini, 60 species are described as new.
Illustrations for eight holotypes or lectotypes of species described by
Bernhauer are given. Five new combinations are recorded.
Key-words: Coleoptera - Staphylinidae - Aleocharinae - Taxonomy - East
Africa.
INTRODUZIONE
Con il presente lavoro si conclude l'esame e la pubblicazione delle Aleo-
charinae dell'Africa orientale conservate al Museo di Storia Naturale di Ginevra. Le
due precedenti parti sono state pubblicate nel 1994 e nel 1995 su questa stessa “Revue
suisse de Zoologie”. Nel presente lavoro sono incluse specie appartenenti, oltre che
alle tribù Thamiaraeini, Oxypodini, Hoplandriini e Aleocharini, anche specie appar-
tenenti alla tribù Myrmedoniini, che include uno sterminato numero di specie africane
tra loro più o meno affini. Purtroppo gli autori del passato (Fauvel, Bernhauer,
Scheerpeltz, Cameron) e anche recenti (Last, Tottenham), che si sono occupati anche
della tassonomia dei Myrmedoniini hanno trascurato l'esame accurato dell'edeago e
della spermateca. Hanno proposto chiavi di determinazione imperfette e fuorvianti.
Pertanto i lavori di questi autori sono di scarso valore e di minima utilità pratica per la
determinazione. Ho pertanto esaminato l'edeago e la spermateca di numerosi tipi della
tribù Myrmedoniini, come quelli di altre tribù, qui in parte illustrati. Ma l'esame di
tutti i tipi esistenti, appartenenti alla tribù Myrmedoniini va oltre le possibilità di un
solo individuo.
127° Contributo alla conoscenza delle Aleocharinae.
Manoscritto accettato il 23.12.1994.
196 ROBERTO PACE
Dato lo stato di incertezza attuale sullo studio sistematico di molti generi della
sottofamiglia Aleocharinae dell'Africa orientdle, e dati i limiti dell'attuale fase di
ricerca sul campo, al termine dell'esame delle specie africane di Aleocharinae del
Museo di Ginevra, non è possibile formulare varie considerazioni, nè proporre ipotesi
sul popolamento di questi minuscoli Coleotteri.
Gli holotypi si conservano nel Museo di Storia Naturale di Ginevra (MG), nel
Museo Zoologico “La Specola” dell'Università di Firenze e al D. E. I di Eberswalde.
Paratypi si conservano nel Museo di Storia Naturale di Ginevra e in collezione
dell'autore.
ELENCO SISTEMATICO DEI GENERI E DELLE SPECIE
THAMIARAEINI
Tyloplatyola microphthalma sp. n. (Figg. 1-8)
Holotypus d, Rwanda, Rangiro, 1800 m, 25.11.1973, (Werner leg., MG).
Paratypi: 2 d d, stessa provenienza, ma 6.VIII.1973; 8 es., Rwanda, Kayove, 2100 m,
12.VIII.1973, (Werner leg.); 1 2, Rwanda, Gisovu, 18.IV.1973, (Werner leg.).
Descrizione. Lungh. 2,0 mm. Corpo lucido e giallo-rossiccio; antenne giallo-
rossicce con i due antennomri basali gialli; zampe gialle. Sul capo la punteggiatura è
distinta e la reticolazione è assente. La reticolazione della superficie del pronoto e
delle elitre è distinta. Tubercoletti salienti coprono il pronoto e le elitre. Edeago figg.
6-7, spermateca fig. 8.
Comparazioni. Per la forma dell'edeago; la nuova specie appare sistemati-
camente affine a 7. orientalis Pace, 1986, dell'Africa orientale, ma quest'ultima specie
ha elitre molto più lunghe del pronoto e occhi lunghi quanto le tempie, mentre la
nuova specie ha elitre più corte del pronoto e occhi molto ridotti. Inoltre l'edeago
della nuova specie è più sviluppato e ha flagello del sacco interno più corto.
Tyloplatyola pseudomicrophthalma sp. n. (Figg. 9-11)
Holotypus 4 , Rwanda, Rangiro, 1800 m, 6.VIII.1973 (Werner leg., MG).
Descrizione. Lungh. 2,0 mm. Corpo lucido e giallo-rossiccio, comprese
antenne e zampe. La reticolazione del capo e del pronoto è molto superficiale, quella
delle elitre & svanita. La punteggiatura del capo & superficiale. I tubercoletti che
coprono la superficie del pronoto sono molto superficiali, quelli sulle elitre sono
molto salienti. Edeao figg. 10-11.
Comparazioni. La nuova specie è affine a T. microphthalma sp. n. sopra des-
critta. Se ne distingue per il pronoto meno trasverso e coperto di tubercoletti molto
svaniti (tubercoletti salienti in microphthalma) e per l'edeago più sviluppato, ampia-
mente ricurvo e angolato al lato ventrale (strettamente ricurvo e non angolato in
microphthalma).
ALEOCHARINAE DELL’ AFRICA ORIENTALE 197
01 mm
01 mm
FIGG. 1-8
Habitus, edeago in visione laterale e ventrale e spermateca. 1-4: Tyloplatyola microphthalma
Sp. n., paratypi di Kayove; 5-8: Tyloplatyola microphthalma sp. n., holotypus & di Rangiro,
paratypus 9 di Gisovu.
198 ROBERTO PACE
Tyloplatyola elgonensis sp. n. (Figg. 12-15)
Holotypus d, Kenya, Mt. Elgon, 2700 m, Nat. Park, 15.X1.1974, Mahnert & Perret
leg., MG).
Paratypi: 21 es., stessa provenienza, ma anche 2700-2800 m e 14.X1.1974.
Descrizione. Lungh. 2,1 mm. Corpo lucido e giallo-rossiccio con elitre ros-
sicce; antenne giallo-brune con 1 tre antennomeri basali gialli; zampe giallo-rossicce.
Il corpo è privo di reticolazione. Il capo presenta tubercoletti fini e salienti. Il pronoto
e le elitre sono coperti di tubercoletti salienti. Edeago figg. 13-14, spermateca fig. 15.
Comparazioni. Specie comparabile con T. microphthalma sp. n. e T. pseudo-
microphthalma sp. n. sopra descritte, per avere elitre molto più corte del pronoto e
forma simile dell'edeago. Ma la nuova specie presenta habitus più tozzo, occhi meno
ridotti, assenza di scultura squamosa dell'addome, edeago molto più sviluppato e parte
prossimale della spermateca molto più lunga.
Tyloplatyola coeca sp. n. (Figg. 16-19)
Holotypus d, Rwanda, Kayove, 2100 m, 29.1V.1973, (Werner leg., MG).
Descrizione. Lungh. 1,45 mm. Corpo lucido e giallo rossiccio con i due
antennomeri basali gialli; zampe gialle. La reticolazione del capo è svanita, quella del
pronoto nettissima, quasi vigorosa e quella delle elitre è distinta. Il capo e il pronoto
non presentano distinta punteggiatura. Le elitre sono coperte di tubercoletti molto
salienti. Edeago figg. 17-18.
Comparazioni. Nell'ambito del genere Tyloplatyola Pace, 1984, la nuova
specie si presenta isolata a motivo dell'anoftalmia ed elitre poco più corte del pronoto,
con addome allargato all'indietro e non appena ristretto all'indietro, come in altre
specie del genere. La ligula intera, i palpi labiali di due articoli, la formula tarsale
4-5-5 e il primo tarsomero posteriore lungo come i due seguenti tarsomeri compresi
insieme, sono caratteri riscontrati nella nuova specie e tipici del genere Tyloplatyola.
Tyloplatyola ruandensis sp. n. (Figg. 19-22)
Holotypus 4, Rwanda, Kayowe, 2100 m, 29.1V.1973, (Wernera leg., MG).
Paratypus: 1 2, stessa provenienza. |
Descrizione. Lungh. 2,2 mm. Corpo lucido e bruno-rossiccio con margini
laterali del pronoto e addome giallo-rossicci; antenne bruno-rossicce con i due anten-
nomeri basali gialli; zampe giallo-rossicce. La reticolazione del capo è estremamente
svanita, quella del pronoto è distinta e quella delle elitre superficiale. La pun-
teggiatura del capo è svanita. Il pronoto presenta tubercoletti distinti, le elitre hanno
tubercoletti superficiali. Edeago figg. 20-21, spermateca fig. 22.
Comparazioni. La nuova specie, in base alla forma della spermateca, sembra
tassonomicamente vicina a 7. simplex Pace, 1984, del Gabon, ma la spermateca della |
nuova specie è di 1/3 più sviluppata e ha parte prossimale proporzionalmente più
allungata. Anche l'habitus mostra caratteri differenziali: addome fortemente ristretto
all'indietro nella nuova specie e quasi a lati paralleli in simplex; elitre molto
199
ALEOCHARINAE DELL’ AFRICA ORIENTALE
01 mm
10 11
01 mm
he
15
Fico. 9-15
Tyloplatyola pseudo-
Habitus, edeago in visione laterale e ventrale e spermateca. 9-11:
microphthalma Sp. n.; 12-15: Tyloplatyola elgonensis sp. n.
200 ROBERTO PACE
sviluppate nella nuova specie ed elitre appena più lunghe del pronoto in simplex.
L'edeago di simplex presenta un lungo flagello del sacco interno, assente nel sacco
interno dell'edeago della nuova specie.
Tyloplatyola kenyensis sp. n. (Figg. 23-24)
Holotypus d, Kenya, Narok, Loita Hills, sous Morijo, 2050 m, 5.X1.1977, (Mahnert &
Perret leg., MG).
Descrizione. Lungh. 2,2 mm. Corpo lucido e giallo-rossiccio con meta pos-
teriore delle elitre bruna (per la presenza di ali brune sottostanti le elitre traslucide);
antenne giallo-rossicce con i due antennomeri basali gialli; zampe gialle. La retico-
lazione del capo è assente, quella del pronoto è molto svanita e quella delle elitre
superficiale. I tubercoletti che coprono a superficie del capo sono distinti, quelli del
pronoto e delle elitre sono molto salienti. Edeago figg. 24-25.
Comparazioni. Per la forma dell'habitus e dell'edeago, la nuova specie è simile
a T. watrousi Pace, 1984, del Gabon. Tuttavia la nuova specie ha occhi meno
sviluppati rispetto a watrousi e non presenta caratteri sessuali secondari sulla zona
suturale delle elitre del maschio e sul 5° urotergo libero del maschio, come in
watrousi. Il flagello del sacco interno dell'edeago della nuova specie, all'interno del
bulbo basale dell'edeago stesso, è avvolto in 6-7 spire nella nuova specie, mentre in
watrousi è avvolto in una sola spira.
Tyloplatyola magna sp. n. (Figg. 26-27)
Holotypus 9, Rwanda, Rangiro, 1800 m, 1.1976, (Werner leg., MG).
Descrizione. Lungh. 2,3 mm. Corpo lucido e giallo-rossiccio con capo e metà
posteriore delle elitre bruno-rossicci; antenne bruno-rossicce con 1 tre antennomeri
basali giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo e del pronoto è
svanita, quella delle elitre assente. I tubercoletti che coprono la superficie del capo
sono molto superficiali, quelli del pronoto sono svaniti e quelli delle elitre sono
distinti. Spermateca fig. 27.
Comparazioni. Specie affine e intermedia tra 7. simplex Pace, 1984, del Gabon,
e T. ruandensis sp. n., sopra descritta. Infatti è affine a queste due specie, per la forma
simile della spermateca, che ha grandezza vicina a quella della spermateca di ruan-
densis, ma la parte prossimale della spermateca della nuova specie è chiaramente più
corta di quella di ruandensis. La taglia della nuova specie è più sviluppata e l'addome è
appena ristretto all'indietro nella nuova specie e fortemente ristretto in ruandensis.
Anche 7. simplex ha addome appena ristretto all'indietro come nella nuova specie, ma
la sua taglia è nettamente minore (1,8 mm invece di 2,3 mm come nella nuova specie) e
la spermateca di simplex è anch'essa di dimensione molto minore.
Polyteinia andreinii Bernhauer, 1933 (Figg. 28-33)
Polyteinia andreinii BERNHAUER, 1933: 59
Materiale esaminato: Lectotypus d, Eritrea, Cheren, 28.XI1.1902, Dr A. Andreini, con
Thermes bellicosus, Polyteinia gridellii (cancellato), andreinii BRNH, n. gen. et sp. (Museo
Zoologico de “La Specola”, Firenze, N° 7786).
ALEOCHARINAE DELL’AFRICA ORIENTALE
01 mm
212
Ficc. 16-22
Habitus, edeago in visione laterale e ventrale e spermateca. 16-18: Tyloplatyola coeca sp. n.:
19-22: Tyloplatyola ruandensis sp. n.
202 ROBERTO PACE
Nota. Questo genere e la specie non sono stati inclusi da Seevers (1957) nel
suo volume “A Monograph on the Termitophilous Staphylinidae”. Questa specie
termitofila è qui per la prima volta illustrata. Sembra che il genere Polyteinia
Bernhauer, 1933, non abbia sinonimi.
MYRMEDONIINI
Catarractodes methnerianus (Bernhauer, 1915) (Figg. 34-39)
Catarractes methnerianus BERNHAUER, 1915: 182
Catarractodes methnerianus: STRAND, 1928: 2
Materiale esaminato: Holotypus 3, D. Ostafrica, Pangani Fälle, 1.VI.1910, Methner,
Catarractes methnerianus BRNH, Typus un. (Mus. Chicago).
La specie, tipo del genere, è qui per la prima volta illustrata.
Catarractodes fulgens (Bernhauer, 1934), comb. n. (Figg. 40-42)
Pachorhopala fulgens BERNHAUER, 1934: 244
Materiale esaminato: Lectotypus 4, Lubero, 23.VHI.1932, Burgeon, Pacorrhopala
(sic!) fulgens BRNH (Mus. Chicago).
Specie qui illustrata per la prima volta.
Catarractodes somalus sp. n. (Figg . 43-45)
Holotypus d, Somalia, Sar Uanle, litorale, 25.11.1973, (S.B.S., Mus. Zool. “La Spe-
cola” Firenze).
Descrizione. Lungh. 2,7 mm. L'esemplare è molto immaturo. Corpo lucido e
giallo, comprese le antenne e le zampe, solo gli uriti liberi 3°, 4° e base del 5° sono
rossicci. Solo sulle elitre è visibile una reticolazione svanita: sul resto del corpo la
reticolazione è assente. La punteggiatura del capo e del pronoto è molto superficiale,
quella delle elitre è svanita. Il 6° urotergo libero del maschio ha una carena mediana
smussata: ai suoi lati stanno fossette e tubercoletti arrotondati. Edeago figg. 44-45.
Comparazioni. La nuova specie presenta habitus simile a quello di C. brun-
neipennis (Bernhauer, 1929), dell'Africa orientale, ma gli antennomeri 4° a 10° della
nuova specie sono molto trasversi (più lunghi che larghi in brunneipennis). Non è
noto il maschio di brunneipennis.
Catarractodes embuensis sp. n. (Figg. 46-48)
Holotypus &, Kenya, Embu, 10 Km 0 Ishiara MI., 900 m, 14.X.1977, (Mahnert &
Perret leg., MG).
Descrizione. Lungh 2,5 mm. Corpo lucido e bruno con estremità addominale
rossiccia; antenne brune con i tre antennomeri basali giallo-rossicci; zampe rossicce.
Tutto il corpo non porta traccia di reticolazione. La punteggiatura del capo è molto
svanita sulla metà anteriore e distinta sulla posteriore, quella del pronoto è ben
visibile. Tubercoletti salienti coprono le elitre. Edeago figg. 47-48.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 203
FIGG. 23-27
Habitus, edeago in visione laterale e ventrale e spermateca. 23-25: Tyloplatyola kenyensis sp. n.;
26-27: Tyloplatyola magna sp. n.
ROBERTO PACE
204
01mm
Olmm
FIGG. 28-33
Habitus, edeago in visione laterale e ventrale, labio con palpo labiale, maxilla con palpo
mascellare e mento. 2-33: Polyteinia andreinii Bernhauer, lectotypus.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 205
FiGG. 34-39
Habitus, edeago in visione laterale e ventrale, maxilla con palpo mascellare, labio con palpo
labiale e mento. 34-39: Catarractodes methnerianus (Bernhauer), holotypus.
206 ROBERTO PACE
Comparazioni. Dato che la nuova specie presenta antennomeri 4° a 10°
fortemente trasversi, essa è comparabile con Catarractodes serranoi (Pace, 1988),
comb. n., (olim Catarractes serranoi Pace, 1988: 27) e Catarractodes minimus (Pace,
1988), comb. n. (olim Catarractes minimus Pace, 1988: 29), entrambe specie della
Guinea-Bissau, che presentano pure antennomeri 4° a 10° fortemente trasversi. Ma la
nuova specie ha corpo di taglia minore, con addome meno ristretto all'indietro ed
edeago più robusto, rispetto a serranoi e corpo di taglia maggiore ed edeago più
robusto rispetto a minimus.
Catarractodes obscurus sp. n. (Figg. 49-50)
Holotypus 9, Kenya, Hulmes Bridge, Nanyuki, 1700 m, 22.X1.1974, (Mahnert &
Perret leg., MG).
Paratypus: 1 ©, Rwanda, Kayove, 2100 m, 29.1V.1973, (Werner leg.).
Descrizione. Lungh. 4,0 mm. Corpo lucido e bruno; antenne brune con i tre
antennomeri basali giallo-rossicci; zampe giallo-rossicce con femori bruno-rossicci.
La punteggiatura del capo è distinta, ma assente sul disco. La netta punteggiatura del
pronoto non raggiunge il margine posteriore del pronoto stesso. Le elitre sono coperte
di punteggiatura simile a quella del pronoto. Sul corpo non vi è traccia di micro-
scultura reticolare. Spermateca fig. 50.
Comparazioni. La grande taglia del corpo, le antenne sottili, con antennomeri
4° a 10° più lunghi che larghi e gli occhi molto sviluppati, sono i più evidenti caratteri
che differenziano la nuova specie dalle restanti specie note. Anche C. methnerianus
(Bernhauer, 1915) (Figg. 34-39) e C. fu/gens (Bernhauer, 1934) presentano antenno-
meri 4° a 10° più lunghi che larghi, ma non sottili come quelli della nuova specie.
Catarractodes striolatus sp. n. (Figg. 51-52)
Holotypus £, Rhodésie, Umtali, I1.1969, (R. Mussard leg., MG).
Descrizione. Lungh. 2,6 mm. Corpo lucido e giallo-rossiccio con capo e 3/4
basali degli uroteghi liberi 3°, 4°, 5° e 6° bruno-rossicci; antenne bruno-ossicce con i
tre antennomeri basali e l'apice dell'11° giallo-rossicci; zampe giallo-rossicce. Il corpo
è privo di microscultura reticolare. La punteggiatura del capo è distinta. Il pronoto e le
elitre sono coperti di tubercoletti salienti. Gli uroterghi liberi 3°, 4° e 5° mostrano
nette strie longitudinali, assenti sulla linea mediana. Spermateca fig. 51.
Comparazioni. La nuova specie, per la presenza di striole longitudinali sugli
uroterghi liberi 3°, 4° e 5°, è unica nel genere. In base alla forma della spermateca,
appare affine a C. obscurus sp. n., sopra descritta, ma la spermateca della nuova
specie ha dimensione molto più ridotta, rispetto quella di obscurus.
Catarractodes hybridus sp. n. (Figg. 53-54)
Holotypus 9, Kenya, env. Endebess, pr. Kitale, 14.XI.1974, (Mahnert & Perret leg.,
MG).
ALEOCHARINAE DELL’ AFRICA ORIENTALE 207
Ficc. 40-45
Habitus, edeago in visione laterale e ventrale. 40-42: Catarractodes fulgens (Bernhauer), lecto-
typus; 43-45: Catarractodes somalus sp. n.
208 ROBERTO PACE
Descrizione. Lungh. 4,4 mm. Corpo lucido e bruno con pronoto giallo-
rossiccio e con uroterghi liberi 1° e 2° rossieci; antenne brune con i tre antennomeri
basali bruno-rossicci; zampe rossicce. La punteggiatura del capo & superficiale e
assente sul disco e ai lati, quella del pronoto é svanita e assente sulla meta posteriore.
Tubercoletti ben salienti coprono le elitre. Spermateca fig. 54.
Comparazioni. In base alla forma della spermateca, in parte avvolta a spirale,
la nuova specie è comparabile con C. brunneipennis (Bernhauer, 1929), dell'Africa
orientale. Pure essa ha la spermateca avvolta a spirale, ma il tubulo che compone la
spermateca stessa di brunneipennis, ha calibro pressoché uguale dal bulbo distale al
prossimale, mentre nella nuova specie il calibro del tubulo della spermateca è sottile
presso il bulbo distale e progressivamente più ampio verso il bulbo prossimale. Inoltre
gli occhi della nuova specie sono meno sviluppati che in brunneipennis.
Pachorhopala (s. str.) usambarae Bernhauer, 1929 (Figg. 55-58)
Pachorhopala usambarae BERNHAUER, 1929: 241
Lectotypus d, “Ost Usambara, D.O.Afr., 12-15, leg. Methner, Oxystigma Oxystigma
usambarae BRNH., Pachorhopala usambarae BRNH. Tupus” (Mus. Chicago).
14 e2 99, Kenya, Miss. Kisumu, NE Kisumu, 1650 m, 10.XI.1974 (Mahnert &
Perret leg.).
La specie è qui illustrata per la prima volta. La spermateca non appartiene a
materiale tipico, ma a una femmina riconosciuta perché associata a un maschio avente
edeago a habitus identici a quelli del lectotypus.
Pachorhopala (s. str.) rufonitens Pace, 1985
Pachorhopala rufonitens PACE, 1985: 153
3 es., Kenya, Mt. Kenya, S Met St. Lodge, 3250 m, 23.XI.1974, (Mahnert & Perret
leg)!
Specie finora nota solo del M. Aberdare.
Pachorhopala (s. str.) africana Bernhauer, 1915
Ocalea (Pachorhopala) africana BERNHAUER, 1915: 186
Pachorhopala africana: Scheerpeltz, 1934: 1677
4 99, Kenya, Embu, Kirimiri For., 13.X.1977, (Mahnert & Perret, leg.); 8 es., Kenya,
Embu, Irangi For., 2000 m, 11.X.1977, (Mahnert & Perret, leg.); 1 2, Kenya, Embu, 20 Km O
Ishiara, 900 m, 14.X.1977, (Mahnert & Perret, leg.); 1 d e 1 9, Kenya, Taita Hills, env. Wun-
danyi, 1400 m, 2.XII.1974, (Mahnert & Perret, leg.); 6 es., Kenya, Kiambu, près de Limuru, 2300 |
m, 2.X1.1977, (Mahnert & Perret, leg.); 1 es., Kenya, Thomson's Falls, 2350 m, 20.X1.1974,
(Mahnert & Perret, leg.); 1 es., Kenya, Mau For., Mau Saummit-Kedowa, 7.X1.1974, (Mahnert &
Perret, leg.); 3 ? ?, Rhodésie, Umtali, 11.1969, (R. Mussard leg.); 3 dd e 1 2, Kenya, Lac |
Nakuru; Parc Nat., 6.X1.1974, (Mahnert & Perret, leg.).
Specie finora nota di una sola località dell'Africa orientale: Usambara.
Pachorhopala (s. str.) kayovensis sp. n. (Figg. 59-61)
Holotypus d, Rwanda, Kayove, 2100 m, 12.VII.1973, (Werner leg., MG).
Paratypus: 1 d, stessa provenienza.
pei
ALEOCHARINAE DELL’ AFRICA ORIENTALE 209
FIGG. 46-52
Habitus, edeago in visione laterale e ventrale e spermateca. 46-48: Catarractodes embuensis
sp. n.; 49-50: Catarractodes obscurus sp. n.; 51-52: Catarractodes striolatus sp. n.
210 ROBERTO PACE
Descrizione. Lungh. 4,9 mm. Corpo lucidissimo e giallo-rossiccio con metà
posteriore delle elitre (tranne 1 lati esterni) e uriti liberi 1° (tranne la metà posteriore),
2° (tranne la metà posteriore e 1 lati), 3° e 4° (tranne 1 lati) bruni; antenne e zampe
giallo-rossicce. La punteggiatura del capo € molto svanita e assente sulla linea
mediana, quella del pronoto e delle elitre è distinta. Sul corpo non vi è traccia di
microscultura reticolare. Edeago figg. 60-61.
Comparazioni. Per l'habitus, la nuova specie sembra simile a P. africana
Bernhauer, 1915, dell'Africa orientale, tuttavia la punteggiatura del pronoto e delle
elitre della nuova specie è distinta, mentre è assai svanita in africana. Ma è l'edeago
che presenta caratteri differenziali più marcati: edeago profondamente incavato al lato
ventrale nella nuova specie, non incavato in africana; piastre del sacco interno
dell'edeago con stiletto distale nella nuova specie e con due punte distali in africana;
edeago, in visione ventrale, a lati debolmente convergenti verso l'apice nella nuova
specie, fortemente in africana.
Pachorhopala (s. str.) tambachensis sp. n. (Figg. 62-64)
Holotypus d, Kenya, Tambach, E Eldoret, 2000 m, 17.X1.1974, (Mahnert & Perret
leg., MG).
Descrizione. Lungh. 4,1 mm. Corpo lucido e bruno con capo bruno-rossiccio e
pronoto giallo-rossiccio; antenne e zampe giallo-rossicce. La punteggiatura del capo e
delle elitre è distinta, quella del pronoto è svanita. Non esiste microscultura reticolare
sul corpo. Edeago figg. 63-64.
Comparazioni. In base alla forma dell'edeago, la nuova specie presenta affinità
con P. kayovensis sp. n. sopra descritta. Tuttavia quest'organo nella nuova specie è
meno sviluppato e ha l'armatura del sacco interno a stiletto largo e tronco all'apice e
non sottile e aguzzo all'apice come in kayovensis. Inoltre il 3° antennomero della
nuova specie è lungo quanto largo, mentre in kayovensis e nettamente trasverso e la
punteggiatura del pronoto è superficiale nella nuova specie, ben distinta in kayovensis.
Pachorhopala (s. str.) laevigata sp. n. (Figg. 65-68)
Holotypus d, Kenya, Miss. Kaimosi, NE Kisumu, 1650 m, 10.X1.1974, (Mahnert &
Perret leg., MG).
Paratypi: 1 6 e 2 9 9, stessa provenienza.
Descrizione. Lungh. 3,0 mm. Corpo lucido e bruno-rossiccio con base ed
estremità addominali giallo-rossicci; antenne brune con l'antennomero basale bruno-
rossiccio; zampe giallo-rossicce. Edeago figg. 66-67, spermateca fig. 68.
Comparazioni. Questa nuova specie, in base alla forma della spermateca,
potrebbe essere affine a P. usambarae Bernhauer, 1929, (figg. 55-58), dell'Africa
orientale, ma l'habitus è nettamente differente: l'addome di usambarae è nettamente
ristretto all'indietro, mentre nella nuova specie lo è appena. La spermateca della nuova
specie presenta 5-6 spire e non 3 come in usambarae e l' edeago della nuova specie è
più ridotto e privo di flagello esteso dal bulbo basale, all'esterno dell'orifizio apicale
dell'edeago stesso, come in usambarae.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 211
FiGG. 53-57
Habitus, spermateca ed edeago in visione laterale e ventrale. 53-54: Catarractodes hybridus sp. n.;
55-57: Pachorhopala (s. str.) usambarae Bernhauer, lectotypus.
212 ROBERTO PACE
Imm
Fico. 58-62
Spermateca, habitus ed edeago in visione laterale e ventrale. 58: Pachorhopala (s. str.) usam-
barae Bernhauer; 59-61: Pachorhopala (s. str.) kayovensis sp. n.; 62: Pachorhopala (s. str.)
tambachensis sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 213
Imm
01 mm
68 ff
FIGG. 63-68
Edeago in visione laterale e ventrale, habitus e spermateca. 63-64: Pachorhopala (s. str.)
tambachensis sp. n., 65-68: Pachorhopala (s. str.) laevigata sp. n.
214 ROBERTO PACE
Pachorhopala (s. str.) confluens sp. n. (Figg. 69-70)
Holotypus 9, Rhodésie, Umtali, II.1969, (R. Mussard leg., MG).
Paratypus: | 9, stessa provenienza.
Descrizione. Lungh. 2,9 mm. Corpo lucidissimo e bruno-rossiccio con base ed
estremità addominale giallo-rossicce; antenne bruno-rossicce con antennomero basale
giallo-rossiccio; zampe giallo-rossicce. La punteggiatura del capo e del pronoto è
svanita, quella delle elitre e dell'addome è netta. Il capo presenta una reticolazione
svanita: sul resto del corpo non è visibile reticolazione. Spermateca fig. 70.
Comparazioni. La nuova specie è strettamente affine a P. laevigata sp. n. sopra
descritta, a motivo della forma della spermateca. Se ne differenzia per avere il pronoto
più trasverso e le spire della spermateca nettamente più strette, composte di tubulo di
calibro chiaramente più sottile di quello di /aevigata.
Pachorhopala (s. str.) ruandensis sp. n. (Figg. 71-72)
Holotypus 9, Rwanda, Kayove, 2100 m, 12.VIII.1973, (Werner leg., MG).
Paratypi: 2 9 9, stessa provenienza.
Descrizione. Lungh. 3,8 mm. Corpo lucidissimo e bruno-rossiccio con base ed
estremità addominali giallo-rossicce; antenne brune con i due antennomeri basali
giallo-rossicci; zampe giallo-rossicce. La punteggiatura del capo è svanita, quella del
pronoto è molto superficiale e quella delle elitre e dell'addome è netta. Solo il capo
presenta una reticolazione, sebbene superficiale: il resto del corpo non è reticolato.
Spermateca fig. 72.
Comparazioni. La struttura della spermateca della nuova specie è simile a
quella di P. africana Bernhauer, 1915. Tuttavia il bulbo prossimale della spermateca
della nuova specie è nettamente ipertrofico, mentre è indistinto in africana. Inoltre
l'habitus della nuova specie è nettamente snello rispetto quello tozzo di africana, dato
che capo e pronoto della nuova specie sono molto più stretti delle elitre (poco più
stretti in africana). Il 3° antennomero della nuova specie è lungo quanto largo, mentre
è nettamente trasverso in africana.
Pachorhopala (s. str.) ishiarensis sp. n. (Figg. 73-76)
Holotypus d, Kenya, Embu, 10 Km O Ishiara MI., 900 m, 14.X.1977, (Mahnert &
Perret leg., MG).
Paratypus: 1 ®, stessa provenienza.
Descrizione. Lungh. 3,7 mm. Corpo lucidissimo e giallo-rossiccio con elitre
con uriti liberi 3°, 4° e 5° bruno-rossicci; antenne brune con i due antennomeri basali
giallo-rossicci e il 3° rossiccio; zampe giallo-rossicce. La punteggiatura del capo e del
pronoto è molto svanita, quella delle elitre è netta. Non vi è traccia di reticolazione sul
corpo. Edeago figg. 74-75, spermateca fig. 76.
Comparazioni. Specie affine a P. africana Bernhauer, 1915, sia per l'habitus,
che per la forma dell'edeago e della spermateca. Tuttavia la nuova specie presenta
elitre nettamente punteggiate (elitre coperte di tubercoletti poco salienti in africana) e
ALEOCHARINAE DELL’ AFRICA ORIENTALE 215
FıGG. 69-76
Habitus, spermateca ed edeago in visione laterale e ventrale. 69-70: Pachorhopala (s. str.)
confluens sp. n.; 71-71: Pachorhopala (s. str.) ruandensis sp. n.; 73-76: Pachorhopala (s. str.)
ishiarensis sp. n.
216 ROBERTO PACE
chiaramente molto più larghe del pronoto. L'edeago della nuova specie ha minore
sviluppo e presenta un pezzo copulatore del sacco interno terminante a stiletto sottile
(e non tronco con due punte come in africana). La spermateca della nuova specie ha
minore sviluppo in lunghezza rispetto quella di africana.
Pachorhopala (s. str.) maculiventris sp. n. (Figg. 77-78)
Holotypus 2, Kenya, Embu, Irangi Forest, 2000 m, 11.X.1977, (Mahnert & Perret leg.,
MG).
Descrizione. Lungh. 4,8 mm. Corpo lucidissimo e giallo-rossiccio con capo,
elitre tranne la base e i lati esterni e una macchia centrale sugli uroterghi liberi 3° e 4°,
bruni; antenne nere con 1 due antennomeri basali giallo-rossicci e l'11° bruno; zampe
giallo-rossicce. La punteggiatura del capo è svanita e assente sulla linea mediana,
quella del pronoto è ombelicata e netta, assente sulla linea mediana e lungo il margine
posteriore e davanti agli angoli posteriori. La punteggiatura delle elitre è netta. Sul
corpo non vi è traccia di reticolazione. Spermateca fig. 78.
Comparazioni. La nuova specie, per la struttura della spermateca, è avvici-
nabile tassonomicamente a P. africana Bernhauer, 1915, ma è nettamente distinta sia
per l'habitus più snello dovuto al capo e al pronoto molto più stretti rispetto alla
larghezza delle elitre (capo e pronoto poco più stretti delle elitre in africana), che per
la spermateca che ha sviluppo maggiore. La macchia bruna sull'addome e la netta
punteggiatura del pronoto e delle elitre della nuova specie sono ulteriori caratteri
differenziali rispetto a P. africana che presenta capo e pronoto coperti di punteggia-
tura superficiale e che è priva di macchia addominale.
Pachorhopala (s. str.) embuicola sp. n. (Figg. 79-81)
Holotypus 4, Kenya, Embu, pr. Ishiara, 900 m, 14.X.1977, (Mahnert & Perret leg.,
MG).
Paratypus: 1 dé, Rhodésie, Embu, II.1969, (R. Mussard leg.).
Descrizione. Lungh. 4,0 mm. Corpo lucidissimo e rossiccio con metà pos-
teriore delle elitre e urite libero 4° bruno-rossicci e 1° e 2° giallo-rossicci; antenne e
zampe giallo-rossicce. La punteggiatura del capo è svanita e assente sulla linea
mediana, quella del pronoto è netta e addensata sulla metà posteriore, ma non ai lati,
nè lungo il margine posteriore. La punteggiatura delle elitre è netta sulla metà
anteriore, svanita sulla posteriore. Non è presente reticolazione sul corpo. Edeago
figg. 80-81.
Comparazioni. Il capo e il pronoto molto più stretti delle elitre, permettono di
associare tassonomicamente la nuova specie non tanto a P. africana Bernhauer, 1915,
quanto a P. maculiventris sp. n. sopra descritta. Tuttavia la nuova specie è differente
da maculiventris poiché presenta solo la metà posteriore del pronoto profondamente
punteggiata, come la metà basale delle elitre (pronoto ed elitre uniformemente
punteggiate in maculiventris).
ALEOCHARINAE DELL’ AFRICA ORIENTALE DAG
Figc. 77-81
Habitus, spermateca ed edeago in visione laterale e ventrale. 77-78: Pachorhopala (s. str.)
maculiventris sp. n.; 79-81: Pachorhopala (s. str.) embuicola sp. n.
218 ROBERTO PACE
Pachorhopala (Troposipalia) spectabilis (Bernhauer, 1934), comb. n. (Figg. 97-100)
Troposipalia spectabilis BERNHAUER, 1934: 242
Materiale esaminato: Lectotypus 3, Ruwenzori, 4200 m, VII.1932, Burgeon, Tropo-
sipalia spectabilis BRNH., Typus, (Mus. Chicago); paralectotypus 9, stessa provenienza (Mus.
Chicago).
Specie illustrata per la prima volta.
Pachorhopala (Troposipalia) spinifera sp. n. (Figg. 82-85)
Holotypus 6, Rwanda, Kayove, 2100 m, 29.1V.1973, (Werner leg., MG).
Paratypi: 1 6 e 1 9, stessa provenienza; 2 d d, Rwanda, Rangiro, 1800 m, 26.VIIL.1975,
(Werner leg.); 1 ¢, Rwanda, Kamiranzovu, 1900 m, marais, 1.1976, (Werner leg.); 2 dd,
Rwanda, Gisovu, 18.1V.1973, (P. Werner leg.).
Descrizione. Lungh. 3,8 mm. Corpo lucido e giallo-rossiccio con uriti liberi 3°
e 4° bruno-rossicci; antenne bruno rossicce con 1 due antennomeri basali giallo-
rossicci; zampe giallo-rossicce. Sul corpo è assente la reticolazione. La punteggiatura
del capo e del pronoto è superficiale e assente sulla linea mediana, quella delle elitre è
netta sulla metà basale e svanita sulla posteriore. Edeago figg. 83-84, spermateca fig.
85.
Comparazioni. La nuova specie, per alcuni caratteri dell'edeago e dell'habitus,
potrebbe essere affine a P. rufonitens Pace, 1985, del M. Aberdare e del M. Kenya,
ma l'edeago della nuova specie possiede delle spine ventrali, assenti in rufonitens e il
pronoto di rufonitens è più lungo che largo. La spermateca della nuova specie ha la
parte prossimale semplice, mentre quella corrispondente in rufonitens è un groviglio
serpentiforme.
Pachorhopala (Troposipalia) elgonensis sp. n. (Figg. 86-87)
Holotypus © , Kenya, Mt. Elgon, 2800 m, 14.XI. 1974, Manhert & Perret leg., MG).
Descrizione. Lungh. 4,2 mm. Corpo lucido e giallo-rossiccio, comprese
antenne e zampe. La punteggiatura del capo è svanita e assente sulla linea mediana,
quella del pronoto è distinta e pure assente sulla linea mediana, quella delle elitre è
netta. Assente è la reticolazione della superficie del corpo. Spermateca fig. 87.
Comparazioni. A motivo della grande taglia corporea, per l'habitus simile e per
la struttura della spermateca, è possibile che la nuova specie sia tassonomicamente
vicina a P. spectabilis Bernhauer, 1934 (figg. 97-100), del Ruwenzori, tuttavia la
taglia corporea della nuova specie è minore (4,2 mm, invece di 6,1 mm), le elitre sono
nettamente punteggiate nella nuova specie (coperte di tubercoletti salienti in spec-
tabilis) e la spermateca ha sviluppo molto minore nella nuova specie.
Pachorhopala (Troposipalia) burgeoni (Bernhauer, 1934), comb. n. (Figg. 88-90)
Troposipalia burgeoni BERNHAUER, 1934: 243
Materiale esaminato: Lectotypus dé, Ruwenzori, 4200 m, VII.1932, Burgeon,
Troposipalia burgeoni BRNH., Typus, (Mus. Chicago).
La specie è qui per la prima volta illustrata.
219
ALEOCHARINAE DELL’ AFRICA ORIENTALE
FIGG. 82-87
Habitus, edeago in visione laterale e ventrale e spermateca. 82-85: Pachorhopala (Tropo-
sipalia) spinifera sp. n.; 86-87: Pachorhopala (Troposipalia) elgonensis sp. n.
220 ROBERTO PACE
Pachorhopala (Troposipalia) ruwenzorii Bernhauer, 1934, comb. n. (Figg. 91-92)
Troposipalia ruwenzorii BERNHAUER, 1934: 243
Materiale esaminato: Lectotypus 9, Ruwenzori, 4200 m, VII.1932, Burgeon L., Tropo-
sipalia ruwenzorii BRNH., Typus, (Mus. Chicago).
Specie illustrata per la prima volta.
Pachorhopala (Troposipalia) ruwenzorensis sp. n. (Figg. 94-97)
Holotypus d, Ruwenzori, Kitandara Hut, 13.200 ft., 1.1984, (L. Lengeler leg., MG).
Paratypus: 1 ©, stessa provenienza.
Descrizione. Lungh. 6,3 mm. Corpo lucidissimo e interamente giallo-rossiccio,
comprese antenne e zampe. La punteggiatura del capo è distinta, quella del pronoto
molto svanita. Tubercoletti salienti coprono le elitre e l'addome. Su tutto il corpo non
vi è traccia di reticolazione. Edeago figg. 94-95, spermateca fig. 96.
Comparazioni. La nuova specie è distinta dalla simpatrica P. ruwenzorii
Bernhauer, 1940, per avere la spermateca più sviluppata (figg. 92 e 96), il pronoto un
po' più trasverso e il 4° antennomero lungo quanto largo (nettamente trasverso in
ruwenzoril). Di P. ruwenzorii non è noto il maschio.
Diplopeurus nanyukensis sp. n. (Figg. 101-102)
Holotypus ©, Kenya, Hulmes Bridge, Nanyuki, 1700 m, 22.X1.1974, (Mahnert &
Perret leg., MG).
Descrizione. Lungh. 4,4 mm. Corpo lucido e bruno-rossiccio con margine
posteriore degli uroterghi liberi 1°, 2° e 3° giallo-rossicci; antenne rossicce, con i due
antennomeri basali giallo-rossicci; zampe giallo-rossicce. Il capo presenta una bozza
tra le antenne e punteggiatura netta, profonda e diradata in avanti. Il pronoto mostra
tubercoletti molto salienti e un largo solco laterale, interrotto posteriormente da una
debole plica. Tubercoletti netti e salienti coprono le elitre. Spermateca fig. 102.
Comparazioni. La nuova specie è distinta da D. excavatus Bernhauer, 1915,
dell'Africa orientale, perché i tubercoletti che coprono il pronoto sono molto più fitti
nella nuova specie che in excavatus e perché le elitre della nuova specie sono coperte
di tubercoletti molto salienti e non di punteggiatura robusta come in excavatus. Inoltre
la base degli uroterghi liberi 3°, 4° e 5° della nuova specie è punteggiata, mentre in
excavatus è priva di punteggiatura.
Drusilla fissata (Bernhauer, 1915) (Figg. 103-106)
Astilbus fissatus BERNHAUER, 1915: 159
Drusilla fissata: PACE, 1986: 109
Materiale esaminato: Lectotypus ©, Africa or., Moschi, FL. Rau, Astilbus fissatus
BRNH., Typus, (Mus. Budapest); paralectotypi: 1 d e 1 ®, Pangani, 1905, Katona, (Mus.
Budapest).
20 es., Kenya, Tana River, Lac Shakababo, près de Ngao, 28.X.1977, (Mahnert &
Perret leg.); 1 es., Kenya, Hulmes Bridge Nanyuki, 1700 m, (Mahnert & Perret leg.).
ALEOCHARINAE DELL’ AFRICA ORIENTALE 221
SI
a
=
01 mm
Er = TR
grigi Myer NE
3,1 VE
ale ai
ro, LER
van tr
IERI -
Imm
92
FıGG. 88-93
Habitus, edeago in visione laterale e ventrale e spermateca. 88-90: Pachorhopala (Troposi-
palia) burgeoni Bernhauer, lectotypus; 91-92: Pachorhopala (Troposipalia) ruwenzorii
Bernhauer, lectotypus; 93: Pachorhopala (Troposipalia) ruwenzorensis sp. n.
222 ROBERTO PACE
Ficc. 94-100
Edeago in visione laterale e ventrale, habitus e spermateca. 94-96: Pachorhopala (Troposi-
palia) ruwenzorensis sp. n.; 97-100: Pachorhopala (Troposipalia) spectabilis Bernhauer, lecto-
typus d e paralectotypus 9.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 223
Fico. 101-106
Habitus, spermateca ed edeago in visione laterale e ventrale. 101-102: Diplopleurus nanyu-
kensis sp. n.; 103-106: Drusilla fissata (Bernhauer), lectotypus ©.
224 ROBERTO PACE
Specie finora nota di due località dell'Africa orientale: Moschi e Pangani;
illustrata per la prima volta.
Drusilla kisumuensis sp. n. (Figg. 107-109)
Holotypus 4, Kenya, Miss. Kaimosi, NE Kisumu, 1650 m, 11.XT.1974, (Mahnert &
Perret leg., MG).
Descrizione. Lungh. 4,1 mm. Corpo lucidissimo e bruno-rossiccio con addome
rossiccio e margini posteriori degli uroterghi giallo-rossicci; antenne giallo-rossicce;
zampe gialle con estremità distale dei femori bruna. La punteggiatura del capo è quasi
indistinta. Il pronoto presenta un'ampia e profonda concavità discale nel fondo della
quale sta un solco mediano; è coperto di tubercoletti svaniti ed è privo di retico-
lazione. Le elitre presentano delle rughe nell'area periscutellare e punteggiatura netta
e profonda, assente agli angoli posteriori esterni dove sta qualche fine tubercoletto.
Edeago figg. 108-109.
Comparazioni. La nuova specie è chiaramente distinta da D. fissata (Bernhauer,
1915), (figg. 103-106), dell'Africa orientale, sia per molti caratteri esoscheletrici, che
dell'edeago. Il capo e il pronoto sono privi di distinta punteggiatura nella nuova specie,
mentre in fissata il capo e il pronoto sono robustamente punteggiati. L'edeago della
nuova specie ha “crista apicalis” assai poco sviluppata e piastre chitinose del sacco
interno non dentellate dorsalmente, mentre nell'edeago di fissata la “crista apicalis” è
molto sviluppata e la piastra chitinosa corrispondente è dentellata al lato dorsale.
Drusilla mahnerti sp. n. (Figg. 110-113)
Holotypus d, Kenya, Shimba Hills, Makadara For., 30.X1.1974, (Mahnert & Perret
leg., MG).
Descrizione. Lungh. 4,3 mm. Corpo lucidissimo e rossiccio con addome
giallo-rossiccio e con lati esterni e angoli posteriori interni delle elitre bruni; antenne
rossicce con i tre antennomeri basali giallo-rossicci; zampe giallo-rossicce con tarsi
rossicci. La punteggiatura ombelicata del capo è distinta e assente su una larga fascia
mediana. Il pronoto presenta in avanti e ai lati punteggiatura molto svanita, assente
lungo la linea mediana; nell'area posteriore mediana del pronoto vi è una distinta e
profonda punteggiatura situata intorno una larga fossetta posteriore. Il 5° urotergo
libero del maschio mostra due carene arcuate e forte punteggiatura, tranne che nel
terzo basale. Edeago figg. 111-112, 6° urotergo libero del maschio fig. 113.
Comparazioni. La nuova specie esternamente è ben distinta da D. fissata
(Bernhauer, 1915) (figg. 103-106) dell'Africa orientale, per avere rada punteggiatura
e radi tubercoletti sull'avancorpo (punteggiatura robusta e fittissima in fissata), per le
due carene e la punteggiatura fitta sul 5° urotergo libero del maschio (assenti in
fissata) e per l'edeago non appuntito, in visione ventrale (appuntito in fissata).
Drusilla umtaliensis sp. n. (Figg. 114-116)
Holotypus 4, Rhodésie, Umtali, I1.1969, Melsetter, 1700 m, (R. Mussard leg., MG).
ALEOCHARINAE DELL’ AFRICA ORIENTALE 225
FIGG. 107-113
Habitus, edeago in visione laterale e ventrale e 6° urotergo libero del maschio. 107-109:
Drusilla kisumuensis sp. n.; 110-113: Drusilla mahnerti sp. n.
226 ROBERTO PACE
Descrizione. Lungh. 4,0 mm. Corpo lucido e rossiccio con gli uriti liberi 3° e
4° bruni, tranne 1l loro margine posteriore che & giallo-rossiccio, e il 5° interamente
bruno; antenne e zampe giallo-rossicce. La punteggiatura del capo è robustissima e
assente per una larga fascia mediana, quella del pronoto è profonda e robusta.
Punteggiatura fitta e netta con tubercoletti salienti copre le elitre. Edeago figg.
115-116.
Comparazioni. I caratteri differenziali più evidenti della nuova specie rispetto
a D. fissata (Bernhauer, 1915) (figg. 103-106), sono una larga fascia mediana non
punteggiata sul capo (assente in fissata) , il 5° e 6° urotergo libero del maschio
fortemente punteggiati (non punteggiati in fissata) ed edeago meno sviluppato e privo
di “crista apicalis”, (edeago più sviluppato e con “crista apicalis” grande in fissata).
Drusilla zimbabwensis sp. n. (Figg. 117-119)
Holotypus d, Rhodésie, Umtali, I1.1969, (R. Mussard leg., MG).
Paratypus: | d, stessa provenienza.
Descrizione. Lungh. 8,3 mm. Corpo lucido e rossiccio con capo e uriti liberi 4°
e 5° bruno-rossicci; antenne rossicce con i tre antennomeri basali giallo-rossicci;
zampe giallo-rossicce. La punteggiatura del capo e del pronoto è netta e assente sulla
linea mediana. Due fossette profonde solcano il pronoto. Una bozza smussata sta sulla
metà posteriore del 2° urotergo libero del maschio. Edeago figg. 118-119.
Comparazioni. La nuova specie è distinta da D. fissata (Bernhauer, 1915)
(figg. 103-106), per la presenza di due solchi sul pronoto e per l'edeago, in visione
ventrale, a lati convergenti verso l'apice (lati paralleli nella porzione submediana in
fissata).
Zyras (Androdonia) masai sp. n. (Figg. 120-122)
Holotypus d, Kenya, Nairobi, 3.X1.1974, (Mahnert & Perret leg., MG).
Descrizione. Lungh. 11,0 mm. Avancorpo debolmente opaco, addome lucido.
Corpo rossiccio con capo e addome bruni: sono rossicci i lati dell'addome, la base
delle spine e il 2° urotergo libero; antenne bruno-rossicce con i tre antennomeri basali
e l'11° giallo-rossicci; zampe giallo-rossicce. Il capo, il pronoto e le elitre presentano
reticolazione fine e netta e punteggiatura ben definita. Il 1° urotergo libero mostra una
rugosità mediana coperta di distinta reticolazione. Edeago figg. 121-122.
Comparazioni. Specie tassonomicamente avvicinabile a Z. bequaerti Bern-
hauer, 1928, dello Zaire. Ne è distinta per le elitre più lunghe del pronoto (più corte
del pronoto in bequaerti) e per le appendici del 1° urotergo libero del maschio, per
nulla allargate verso l'apice, in visione laterale (fig. 122 bis) come in bequaerti.
Zyras (Camonia) somaliensis Bernhauer, 1927
Zyras (Camonia) somaliensis BERNHAUER, 1927: 184
1 del 9, Somalia, Sar Uanle, 29.V.1973, (S.B.S. leg., Mus. “La Specola”, Firenze).
Specie finora nota solo della Somalia.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 227
116
Fico. 114-119
Habitus ed edeago in visione laterale e ventrale. 114-116: Drusilla umtaliensis sp. n.; 117-119
Drusilla zimbabwensis sp. n.
228 ROBERTO PACE
Zyras (Camonia) ngaoensis sp. n. (Figg. 123-125)
Holotypus d, Kenya, Tana River, Lac Shakababo, près Ngao, 28.X.1977, (Mahnert &
Perret leg., MG).
Descrizione. Lungh. 7,1 mm. Disco del capo opaco, resto del corpo lucido e
giallo rossiccio con capo e uriti liberi 3°, 4° e 5° bruno-rossicci e angoli posteriori
esterni delle elitre sfumati di bruno; antenne bruno-rossicce con i tre antennomeri
basali e l'11° giallo-rossicci; zampe giallo-rossicce. Il capo presenta reticolazione
netta solo sul disco e il resto della superficie senza reticolazione. La punteggiatura del
capo è distinta, quella del pronoto, delle elitre e dell'addome è netta. La reticolazione
sul pronoto è svanita, quella sulle elitre è molto svanita e quella dell'addome è
distinta. Edeago figg. 124-125.
Comparazioni. Specie tassonomicamente avvicinabile a Z. afer Gestro, 1892,
della Somalia. La nuova specie ne è distinta per avere il disco del capo opaco e non
depresso (capo largamente depresso e fortemente reticolato in afer) e per i prolun-
gamenti laterali del 1° urotergo libero del maschio più lunghi e ricurvi (corti in afer).
Zyras (Camonia) kilifensis sp. n. (Figg. 126-127)
Holotypus 9, Kenya, Kilifi Jilore, 29.X.1977, (Mahnert & Perret leg., MG).
Descrizione. Lungh. 7,8 mm. Avancorpo opaco, addome lucido. Corpo giallo-
rossiccio con capo, metà posteriore delle elitre e 4° urite libero bruno-rossicci;
antenne rossicce con i due antennomeri basali giallo-rossicci; zampe giallo-rossicce. Il
capo, il pronoto e le elitre presentano reticolazione netta a maglie regolari e
punteggiatura quasi indistinta. La reticolazione dell'addome è a maglie poligonali
irregolari nette. Spermateca fig. 127.
Comparazioni. Specie tassonomicamente vicina a Z. disputandus Last, 1963,
della Rodesia, ma distinta per avere il pronoto non sinuato lateralmente e privo di fine
solco mediano e per le elitre trasverse, se considerate insieme, (lunghe quanto larghe
in disputandus).
Zyras (s. str.) tambachensis sp. n. (Figg. 128-130)
Holotypus d, Kenya, Tambach, E Eldoret, 2000 m, 17.X1.1974, (Mahnert & Perret
leg., MG).
Descrizione. Lungh. 5,8 mm. Corpo lucido e giallo-rossiccio con capo bruno-
rossiccio ed elitre di un giallo sporco; antenne brune con i due antennomeri basali
bruno-rossicci; zampe giallo-rossicce. La punteggiatura del capo è quasi indistinta,
quella del pronoto è distinta e quella delle elitre è netta. E' assente la reticolazione su
tutto il corpo. Edeago figg. 129-130.
Comparazioni. Specie distinta da Z. abessinus (Bernhauer, 1915) dell'Abis-
sinia, per avere la punteggiatura del capo quasi indistinta (capo chiaramente pun-
teggiato in abessinus).
ALEOCHARINAE DELL’ AFRICA ORIENTALE 229
F1GG. 120-125
Habitus, edeago in visione laterale e ventrale e spina del 1° urotergo libero del maschio, in
visione laterale. 120-122 bis: Zyras (Androdonia) masai sp. n.; 123-125: Zyras (Camonia)
ngaoensis Sp. n.
230 ROBERTO PACE
127
Ficc. 126-130
Habitus, spermateca ed edeago in visione laterale e ventrale. 126-127: Zyras (Camonia) kili-
fensis sp. n.; 128-130: Zyras (s. str.) tambachensis sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 231
Zyras (s. str.) nakuruensis sp. n. (Figg. 131-132)
Holotypus 2, Kenya, Nakuru, Lac Naivasha, près de Mundui Estate, 1950 m, 8.X1.1977,
(Mahnert & Perret leg., MG).
Descrizione. Lungh. 6,0 mm. Corpo lucidissimo e giallo-rossiccio con capo,
elitre tranne la base e uriti liberi 4° e 5° bruni; antenne brune con i due antennomeri
basali e l'11° bruno-rossicci; zampe giallo-rossicce. La punteggiatura del capo è
svanita e assente sulla linea mediana, quella del pronoto e delle elitre è netta, diradata
sulla metà posteriore delle elitre. Spermateca fig. 132.
Comparazioni. Specie chiaramente distinta tassonomicamente da Z. dudu
Eichelbaum, 1913, dell'Africa orientale, per avere la spermateca a matassa nettamente
più fitta e per il capo e il pronoto sparsamente punteggiati (fittamente e profonda-
mente punteggiati in dudu).
Zyras (s. str.) mutarensis sp. n. (Figg. 133-136)
Holotypus d, Rhodésie, Umtali, 11.1969, (R. Mussard leg., MG).
Paratypus: 1 ®, stessa provenienza.
Descrizione. Lungh. 3,9 mm. Corpo lucidissimo e bruno-rossiccio con pro-
noto, base delle elitre e base ed estremità dell'addome giallo-rossicci; antenne brune
con i tre antennomeri basali giallo-rossicci; zampe gialle. La punteggiatura del capo e
del pronoto è molto svanita, quella delle elitre è netta. Reticolazione del corpo
assente. Edeago figg. 134-135, spermateca fig. 136.
Comparazioni. Specie ben distinta da Z. dudu Eichelbaum, 1913, dell'Africa
orientale, per avere il pronoto molto superficialmente punteggiato, l'addome ristretto
all'indietro (non ristretto in dudu) e per la fitta matassa della spermateca (matassa
della spermateca rada in dudu).
Zyras (Pachydonia) seditiosus sp. n. (Figg. 137-139)
Holotypus é, Kenya, Mt. Aberdares, Parc National, 2300 m, 25.X1.1974, (Mahnert &
Perret leg., MG).
Paratypus: | d, stessa provenienza.
Descrizione. Lungh. 5,9 mm. Corpo lucido e bruno-rossiccio con pronoto e
uriti liberi 1°, 2° e 6° rossicci; antenne brune con i due antennomeri basali e la base
del 3°giallo-rossicci e l'11° bruno-rossiccio; zampe giallo-rossicce. Il capo presenta
una bozza frontale tra le antenne e punteggiatura distinta, assente sulla linea mediana.
La punteggiatura del pronoto è distinta, fine ai lati e assente lungo il margine pos-
teriore, quella delle elitre è netta. Edeago figg. 138-139.
Comparazioni. Specie distinta da Z. dubius Bernhauer, 1928, dello Zaire, per
avere sul pronoto un solco posteriore mediano corto invece di una profonda fossetta,
come in dubius e per la punteggiatura del pronoto stesso distinta (e non svanita sulla
linea mediana come in dubius).
Zyras (Euryalonia) peranomalus sp. n. (Figg. 142-143)
Holotypus 9, Kenya, Mt. Aberdares, pres Nat. Park, 2300 m, 25.X1.1974, (Mahnert &
Perret leg., MG).
232 ROBERTO PACE
Descrizione. Lungh. 5,7 mm. Corpo debolmente opaco, un po' depresso e
bruno-rossiccio con base delle elitre e addome, tranne gli uroterghi liberi 4° e 5°,
giallo-rossicci; antenne brune con i due antennomeri basali giallo-rossicci; zampe
giallo-rossicce. L'avancorpo è coperto di reticolazione netta. La punteggiatura del
capo e del pronoto è svanita e fine, quella delle elitre è netta. Gii antennomeri sono
lateralmente lievemente compressi. Spermateca fig. 143.
Comparazioni. Specie ben distinta da Z. braunsi Bernhauer, 1928, del Sud-
africa per avere il pronoto più trasverso e per il differente colore delle elitre: elitre
bruno-rossicce con base giallo-rossiccia nella nuova specie, nero pece in braunsi.
Zyras (Tropidonia) rangirensis sp. n. (Figg. 144-145)
Holotypus 9, Rwanda, Rangiro, 1800 m, 4.IX.1976, (Werner leg., MG).
Descrizione. Lungh. 6,4 mm. Corpo lucido e rossiccio con capo e uroterghi
tranne il loro margine posteriore, bruni; antenne rossicce con i due antennomeri basali
giallo-rossicci; zampe giallo-rossicce. Gli antennomeri non sono compressi lateral-
mente. La punteggiatura del capo è fine e distinta, ma assente sulla linea mediana,
quella del pronoto è molto superficiale. Tubercoletti molto salienti stanno sulle elitre e
tubercoletti molto svaniti sull'addome. Il pronoto è lateralmente profondamente
depresso. Spermateca fig. 145.
Comparazioni. La nuova specie si distingue da Z. glaberrimus Bernhauer,
1926, dello Zaire, per avere il capo ristretto dietro gli occhi (capo un po' allargato
dietro gli occhi in glaberrimus), per gli antennomeri 4° e 5° più lunghi che larghi
(lunghi quanto larghi in glaberrimus) e per le elitre molto più lunghe del pronoto e
debolmente allargate all'indietro (poco più lunghe del pronoto e fortemente allargate
all'indietro in glaberrimus).
Zyras (Pycnodonia) microarmatus sp. n. (Figg. 140-141 e 146-149)
Holotypus d, Kenya, Nairobi, 4.XI.1974, (Mahnert & Perret leg., MG).
Paratypi: 1 d e 2 2%, stessa provenienza; 14 es., Namibia, East Caprivi: Katima
Mulilo, lux, 17' 29"S/24' 17"E, 3.-8.111.1992, (Uhlig Leg., Mus. Berlin).
Descrizione. Lungh. 5,4 mm. Corpo lucido e bruno-rossiccio; antenne bruno-
rossicce con i due antennomeri basali e l'11° giallo-rossicci; zampe giallo-rossicce. La
punteggiatura del capo è estremamente svanita, quella delle elitre è profonda nella
regione periscutellare: sul resto della superficie delle elitre ci sono tubercoletti salienti
e radi. Il pronoto è coperto di tubercoletti fini e radi. Gli antennomeri non sono
lateralmente compressi. Edeago figg. 148-149, spermateca fig. 141 e 146.
Comparazioni. Specie distinta da Z. lioglutoides Bernhauer, 1915, dell'Africa
orientale, per avere la punteggiatura del capo e del pronoto estremamente svanita
(punteggiatura robusta in lioglutoides), per le tempie lunghe quanto gli occhi (tempie
più lunghe degli occhi in lioglutoides), per gli antennomeri 6° e 7° più lunghi che
larghi (chiaramente trasversi in lioglutoides) e per le elitre robustamente punteggiate
solo nella zona periscutellare (fittamente punteggiate per intero in lioglutoides).
ALEOCHARINAE DELL’ AFRICA ORIENTALE 233
FIGG. 131-136
Habitus, spermateca ed edeago in visione laterale e ventrale. 131-132: Zyras (s. str.)
nakuruensis Sp. n.; 133-136: Zyras (s. str.) mutarensis Sp. n.
234 ROBERTO PACE
Imm
FIGG. 137-141
Habitus, edeago in visione laterale e ventrale e spermateca. 137-139: Zyras (Pachydonia)
seditiosus sp. n.; 140-141: Zyras (Pycnodonia) microarmatus sp. n.
ALEOCHARINAE DELL” AFRICA ORIENTALE 235
F1GG. 142-149
Habitus, spermateca ed edeago in visione laterale e ventrale. 142-143: Zyras (Euryalonia) pera-
nomalus sp. n.; 144-145: Zyras (Tropidonia) rangirensis sp. n.; 146-14: Zyras (Pycnodonia)
microarmatus sp. N.
236 ROBERTO PACE
Zyras (Pycnodonia) irangicola sp. n. (Figg. 150-152)
Holotypus d, Kenya, Embu, Irangi For., 2100 m, 12.X.1977, (Mahnert & Perret leg.,
MG).
Descrizione. Lungh. 4,8 mm. Corpo lucido e rossiccio con capo bruno; antenne
bruno-rossicce con 1 tre antennomeri basali giallo-rossicci; zampe giallo-rossicce. Il
capo presenta punteggiatura distinta e assente sulla linea mediana, reticolazione
distinta e una concavita discale. Il pronoto mostra punteggiatura superficiale e assente
sulla linea mediana e reticolazione distinta, tranne che sulla linea mediana che perciò
appare lucida. La punteggiatura e la reticolazione delle elitre sono come quelle del
pronoto. Edeago figg. 151-152.
Comparazioni. La nuova specie è distinta da Z. lioglutoides Bernhauer, 1915,
dell'Africa orientale, per la presenza di una concavità discale del capo del maschio
(assente in lioglutoides) e per avere sul 6° urotergo libero del maschio, un tubercolo
mediano e non due affilate protuberanze come in lioglutoides.
Zyras (Pycnodonia) mahnerti sp. n. (Figg. 153-155)
Holotypus 4, Kenya, Mt. Aberdares, près Nat. Park, 2300 m, 25.X1.1974, (Mahnert &
Perret leg., MG).
Paratypus: 1 d, stessa provenienza.
Descrizione. Lungh. 7,9 mm. Corpo lucido e rossiccio con 4° urite libero
bruno-rossiccio; antenne bruno-rossicce con i due antennomeri basali giallo-rossicci;
zampe giallo-rossicce. La punteggiatura è distinta sul capo e netta sul resto del corpo.
Sul capo la punteggiatura è assente su una larga fascia longitudinale mediana, sul
pronoto è assente sulla linea mediana, su due aree mediane anteriori e lungo il
margine posteriore del pronoto stesso. Edeago figg. 153-154.
Comparazioni. Z. mahnerti sp. n. è specie distinta da Z. lioglutoides Bern-
hauer, 1915, dell'Africa orientale, per la punteggiatura del capo non robusta come
quella di lioglutoides, per le tempie lunghe quanto gli occhi (e non più lunghe degli
occhi come in lioglutoides) e per i caratteri sessuali secondari sul 5° urotergo libero
del maschio: due tubercoli assenti nella nuova specie, presenti in lioglutoides.
Zyras (Pycnodonia) kisumuensis sp. n. (Figg. 156-157)
Holotypus 2, Kenya, Miss. Kaimosi, NE Kisumu, 1650 m, 10.X1.1974, (Mahnert &
Perret leg., MG).
Descrizione. Lungh. 6,8 mm. Corpo lucido e bruno-rossiccio con capo e 4°
urite libero bruni; antenne rossicce con i due antennomeri basali giallo-rossicci;
zampe giallo-rossicce. L'avancorpo è privo di microscultura reticolare, l'addome
invece è distintamente reticolato. La punteggiatura del capo, del pronoto e delle elitre
è netta: essa è assente sulla linea mediana del capo. Un fine solco mediano posteriore
è presentato dal pronoto. Gli antennomeri non sono lateralmente compressi.
Spermateca fig. 157.
ALEOCHARINAE DELL’ AFRICA ORIENTALE
N
WW
N
Fico. 150-155
Habitus ed edeago in visione laterale e ventrale. 150-152: Zyras (Pycnodonia) irangicola sp. n.;
153-155: Zyras (Pycnodonia) mahnerti sp. n.
238 ROBERTO PACE
Comparazioni. Il pronoto nettamente più stretto delle elitre & tra l'altro un
carattere differenziale sufficiente a distiguere la nuova specie da Z. lioglutoides
Bernhauer, 1915 dell'Africa orientale che ha il pronoto poco più stretto delle elitre.
Zyras (Parophthalmonia) narokensis sp. n. (Figg. 158-159)
Holotypus ©, Kenya, Narok, près de Narok, 2000 m, 3.X1.1977, (Mahnert & Perret
leg., MG).
Paratypus: 1 ©, stessa provenienza.
Descrizione. Lungh. 6,9 mm. Avancorpo lievemente opaco, addome lucido.
Corpo nero-bruno con estremità addominale posteriore bruno-rossiccia; antenne
bruno-rossicce con i due antennomeri basali e l'11° giallo-rossicci; zampe giallo-
rossicce. La reticolazione del capo, del pronoto e dell'addome è netta, quella delle
elitre è nettissima. La punteggiatura sull'avancorpo è netta, quella dell'addome è
superficiale. Sul pronoto la punteggiatura è assente sulla linea mediana e al lato
esterno delle due deboli fossette longitudinali mediane. Gli antennomeri sono
lateralmente compressi. Spermateca fig. 158.
Comparazioni. La nuova specie è ben distinta da Z. holtzi Bernhauer, 1928,
dell'Africa orientale, per avere il pronoto densamente punteggiato, tranne su due aree
discali (sparsamente punteggiato in holtzi) e per le elitre lunghe quasi quanto il
pronoto (e non 1/4 più lunghe del pronoto come in holfzi).
Zyras (Ctenodonia) luorum sp. n. (Figg. 160-162)
Holotypus d, Kenya, Mt. Elgon, 2400 m, For. Res., 4.X1.1974, (Mahnert & Perret leg.,
MG).
Descrizione. Lungh. 8,7 mm. Pronoto ed elitre debolmente opachi, resto del
corpo lucido. Corpo rossiccio compresi 1 lati del 2° urite libero e con capo e uriti liberi
2°, 3°, 4° e 5° bruni; antenne rossicce con i tre antennomeri basali e apice dell'11°
giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo e dell'addome è
distinta, quella del pronoto e delle elitre è vigorosa. Tutto il corpo presenta distinta
punteggiatura fine. Le antenne sono compresse lateralmente e il pronoto presenta una
lieve depressione laterale obliqua. Edeago figg. 161-162.
Comparazioni. Specie ben distinta da Z. magnificus Bernhauer, 1911, del
Kilimangiaro, per avere gli antennomeri 8° a 10° più lunghi che larghi, per il pronoto
lungo quanto largo (e non trasverso come in magnificus) e per le elitre più lunghe del
pronoto (e non lunghe quanto il pronoto come in magnificus).
Etimologia. La nuova specie prende nome dai Luo, antico gruppo etnico del
Kenya.
Zyras (Ctenodonia) perreti sp. n. (Figg. 163-164)
Holotypus 2, Kenya, Mt. Aberdares, près Nat. Park, 2300 m, 25.X1.1974, (Mahnert &
Perret leg., MG).
Paratypus: 1 2, stessa provenienza.
ALEOCHARINAE DELL’ AFRICA ORIENTALE
Ne pa ‘ Ve ce
(TE
ORA
qe
ES
FısG. 156-159
Habitus e spermateca. 156-157: Zyras (Pycnodonia) kisumuensis sp. n.;
(Parophthalmonia) narokensis sp. n.
Prada VER
{SURV ANA pate} 9} AB :
Ve TS
HEIN URI
239
Imm
158-159: Zyras
240 ROBERTO PACE
Descrizione. Lungh. 8,4 mm. Corpo lucido e rossiccio con capo, elitre e uriti
liberi 4° e 5° bruno-rossicci; antenne bruno-rossicce con 1 tre antennomeri basali
giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo é distinta, quella sul
resto del corpo è netta. La punteggiatura del capo e delle elitre è netta, quella del
pronoto è superficiale. Spermateca fig. 163.
Comparazioni. La nuova specie è distinta da Z. magnificus Bernhauer, 1911,
del Kilimangiaro, per il differente colore dell'addome (interamente nero in magnificus
e non rossiccio con uriti liberi 4° e 5° bruno-rossicci come nella nuova specie).
Trichodonia ruandorum sp. n. (Figg. 165-166)
Holotypus 9, Rwanda, Kayove, 2100 m, 29.VIII.1973, (Werner leg., MG).
Descrizione. Lungh. 7,1 mm. Corpo lucido e giallo-rossiccio con capo, elitre e
4° urite libero bruni; antenne brune con apice dell'11° antennomero rossiccio; zampe
bruno-rossicce con tarsi rossicci. Assente è la reticolazione su tutto il corpo. La
punteggiatura del capo è assente, quella dell'addome è distinta. Tubercoletti salienti
coprono la superficie del pronoto e delle elitre. Il pronoto ha una larga depressione
laterale. Gli antennomeri non sono compressi lateralmente. Spermateca fig. 166.
Comparazioni. Specie distinta dalle varie specie del genere, per l'assenza di
lunghe setole laterali del pronoto e delle elitre.
Porus nakuruensis sp. n. (Figg. 167-169)
Holotypus d, Kenya, Nakuru Mau Escarp., près Enangiperi, 2700 m, 6.X1.1977,
(Mahnert & Perret leg., MG).
Descrizione. Lungh. 6,8 mm. Avancorpo opaco, addome lucido. Corpo nero-
bruno; antenne nero-brune con 2° antennomero, base del 3° e l'11° rossicci; zampe
bruno-rossicce. I tubercoletti della superficie del capo, del pronoto e delle elitre sono
evidenti e allungati: sono assenti sulla linea mediana del capo e del pronoto. La
reticolazione della superficie dell'avancorpo è netta, quella dell'addome è distinta.
Edeago figg. 168-169.
Comparazioni. Specie distinta da P. cupulifer Fauvel, 1899, del Senegal, per
avere la fronte priva di impressione, per l'assenza di spine sul 2° urotergo libero del
maschio e per l'assenza di asperità trasversa sinuosa sul 6° urotergo libero del
maschio.
Porus ruandensis sp. n. (Figg. 170-172)
Holotypus d, Rwanda, Rangiro, 1800 m, 20.VIII.1976, (Werner leg., MG).
Descrizione. Lungh. 7, 6 mm. Corpo lucido e bruno con base ed estremità
dell'addome bruno-rossicce; antenne brune con i due antennomeri basali rossicci;
zampe giallo-rossicce. La punteggiatura del capo è ombelicata, netta e assente sulla
linea mediana, quella del pronoto è evidente e presente solo su due file dorsali e due
fasce laterali. La punteggiatura delle elitre è netta e assente ai lati esterni. L'addome è
coperto di punteggiatura profondissima. Edeago figg. 171-172.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 241
Imm
0)mm
163
F1GG. 160-164
Habitus, edeago in visione laterale e ventrale e spermateca. 160-162: Zyras (Ctenodonia)
luorum sp. n.; 163-164: Zyras (Ctenodonia) perreti sp. n.
242 ROBERTO PACE
Imm
Fico. 165-169
Habitus, spermateca ed edeago in visione laterale e ventrale. 165-166: Trichodonia ruandorum
sp. n.; 167-169: Porus nakuruensis sp. n.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 243
Comparazioni. Specie ben distinta da P. natalensis Fauvel, 1899, del Natal,
per avere il capo lucido (e non opaco e reticolato come quello di natalensis), per
l'assenza di spine sul 2° urotergo libero del maschio e di carena mediana longitudinale
sul 6° urotergo libero del maschio.
Porus umtaliensis sp. n. (Figg. 173-176)
Holotypus dé, Rhodésie, Umtali, 11.1969, (R. Mussard leg., MG).
Paratypi: 3 © 9, stessa provenienza.
Descrizione. Lungh. 3,6 mm. Corpo lucido e bruno-rossiccio; antenne bruno-
rossicce con i due antennomeri basali giallo-rossicci; zampe giallo-rossicce. La
punteggiatura del capo è netta e assente sulla linea mediana e nell'area occipitale,
quella del pronoto è lievemente svanita e assente lungo la linea mediana, quella delle
elitre è netta e quella dell'addome svanita. Non vi è traccia di microscultura reticolare.
Edeago figg. 174-175, spermateca fig. 176.
Comparazioni. A motivo della piccola taglia, la nuova specie è facilmente
distinguibile sia da P. cupulifer Fauvel, 1899, del Senegal, sia da P. natalensis Fauvel,
1899, che da P. longicollis Fauvel, 1899, del Sudafrica, che hanno corpo lungo quasi
il doppio.
Aenictonia kenyensis sp. n. (Figg. 177-179)
Holotypus d, Kenya, Mt. Aberdares, près Nat. Park, 2300 m, 25.X1.1974, (Mahnert &
Perret leg., MG).
Descrizione. Lungh. 6,1 mm. Corpo debolmente lucido e bruno con pronoto,
base degli uriti liberi 2°, 3° e 4° e latı dell'addome, rossicci; antenne rossicce con 1 tre
antennomeri basali giallo-rossicci; zampe gialle. La reticolazione del capo, del
pronoto e dell'addome è distinta, quella delle elitre è netta. I tubercoletti che coprono
il capo e il pronoto sono distinti, quelli delle elitre ben salienti. Edeago figg. 178-179.
Comparazioni. Specie distinta dalle altre note, per la presenza di un'accentuata
gibbosità ventrale dell'edeago, assente in tutte le specie note. La specie nuova è
tassonomicamente vicina ad A. carinata Kistner, 1968, dello Zaire, per i caratteri
esoscheletrici, ma l'edeago, oltre ad avere la citata sinuosità ventrale, in visione
laterale, è privo di elevata sinuosità preapicale dell'edeago, presente invece in
carinata.
OXYPODINI
Amarochara loitensis sp. n. (Figg. 180-183)
Holotypus 3, Kenya, Narok, Loita Hills, sous Morijo, 2050 m, 5.X1.1977, (Mahnert &
Perret leg., MG).
Paratypus: 1 ?, stessa provenienza.
Descrizione. Lungh. 2,7 mm. Corpo lucido e bruno-rossiccio con elitre e base
degli uroterghi brune; antenne brune; zampe giallo-rossicce. La punteggiatura del
ROBERTO PACE
244
Ficc. 170-176
Habitus, edeago in visione laterale e ventrale e spermateca. 170-172: Porus ruandensis sp. n.;
173-176 Porus umtaliensis sp. n.
__==
ALEOCHARINAE DELL’ AFRICA ORIENTALE 245
Ficg. 177-183
Habitus, edeago in visione laterale e ventrale e spermateca. 177-179: Aenictonia kenyensis sp.
n.; 180-183: Amarochara loitensis sp. n.
246 ROBERTO PACE
capo e del pronoto é fine e distinta. Le elitre presentano tubercoletti assai svaniti,
mentre l'addome li ha distinti. La superficie dei corpo non è reticolata. Edeago figg.
180-181, spermateca fig. 182.
Comparazioni. Specie distinta da A. sparsa Tottenham, 1957, del Ruanda, per
avere le elitre non fortemente punteggiate e per avere il bulbo distale della spermateca
molto più sviluppato e la parte prossimale della spermateca stessa, a forma di corno e
non sottile e a lati paralleli come in sparsa.
Amarochara nakuruensis sp. n. (Fig. 184-185)
Holotypus 9, Kenya, Lac Naivasha, Nakuru, près de Mundui Estate, 1950 m,
8.X1.1977, (Mahnert & Perret leg., MG).
Descrizione. Lungh. 2,5 mm. Corpo lucido e rossiccio con uriti liberi 4° e 5°
bruno-rossicci; antenne brune con i due antennomeri basali giallo-rossicci; zampe
giallo-rossicce. La punteggiatura del capo e del pronoto è fine e svanita. Tubercoletti
salienti coprono le elitre. Non esiste reticolazione sul corpo. Spermateca fig. 185.
Comparazioni. Specie esternamente simile ad A. sparsa Tottenham, 1957, del
Ruanda, ma la spermateca ha forma chiaramente differente nella nuova specie: la
parte prossimale è bisinuata e di calibro maggiore presso il bulbo distale e di calibro
minore verso il bulbo prossimale che è indistinto, mentre la parte prossimale della
spermateca di sparsa ha calibro uniforme ed esile e non è sinuata.
Apimela mahnerti sp n. (Figg. 213-215)
Holotypus d, Kenya, Narok, 5.X.1977, Loita Hills, 2050 m, (Mahnert & Perret leg.,
MG).
i Paratypi: 1 ©, stessa provenienza, ma 4.X.1977; 2 dd e 2 2 2, Kenya, Mt. Aberdares,
Parc National, 2300 m, 25.X1.1974, (Mahnert & Perret leg.); 2 es. Kenya, Taita Hills, env.
Wundanyi, 1350 m, 2.XII.1974, (Mahnert & Perret leg.); 3 es., Kenya, Embu, Irangi Forest st.,
2000 m, 11.X.1977, (Mahnert & Perret leg.); 1 ¢, Kenya, Mt. Elgon, 2700 m, 15.X1.1974,
(Mahnert & Perret leg.); 1 2, Kenya, Kiambu, près de Limuru, 2300 m, 2.X1.1977, (Mahnert &
Perret leg.); 1 ¢, Rhodésie, Umtali, Melsetter, 1700 m, 11.1969, (R. Mussard leg.); 1 d e 2
22, Rhodésie, Umtali, II.1969, (R. Mussard leg.); 5 es., Rwanda, Rangiro, 1800 m,
20.1X.1976 e 8.111.1972, (Werner leg.); 550 es., Rwanda, Kayove, 15-29.V.1973, (Werner leg.).
Descrizione. Lungh. 2,3 mm. Corpo lucido e nero-bruno con pronoto e 6° urite
libero bruni; antenne nero-brune con i due antennomeri basali bruni; zampe di un
giallo sporco con femori posteriori giallo-bruni. La reticolazione del capo e
dell'addome è svanita, quella del pronoto e delle elitre è svanita. I tubercoletti che
coprono la superficie del capo sono svaniti. La punteggiatura del pronoto è finissima e
poco evidente. Edeago figg. 211-212, spermateca Figg. 213.
Comparazioni. La taglia della nuova specie è maggiore di quella di A.
subparallela (Bernhauer, 1938), dello Zaire, l'edeago della nuova specie è tozzo,
mentre in subparallela è snello e la spermateca della nuova specie ha matassa molto
sviluppata, mentre in subparallela la matassa è poco sviluppata.
ALEOCHARINAE DELL’ AFRICA ORIENTALE 247
FIGG. 184-189
Habitus, spermateca ed edeago in visione laterale e ventrale. 184-185: Amarochara naku-
ruensis Sp. n.; 186-89: Apimela luorum Sp. n.
248 ROBERTO PACE
Apimela heteroclita sp. n. (Figg. 216-219)
Holotypus /, Kenya, Nairobi, 2.X1.1974, Mahnert & Perret leg., MG).
Paratypi: 10 es., stessa provenienza, ma anche 4.XI.1974.
Descrizione. Lungh. 2,2 mm, Corpo lucido e bruno con pronoto, i tre uriti
basali e l'estremità addominale bruno-rossicci; antenne brune con i due antennomeri
basali bruno-rossicci; zampe gialle. La reticolazione del capo e dell'addome è svanita,
quella del pronoto e delle elitre è distinta. La punteggiatura del capo è indistinta. Il
pronoto presenta tubercoletti svaniti, le elitre mostrano tubercoletti poco salienti.
Edeago fig. 217-218, spermateca fig. 219.
Comparazioni. La nuova specie è chiaramente distinta da A. subparallela
(Bernhauer, 1939a, dello Zaire, per la taglia maggiore, per avere l'edeago tozzo
(snello in subparallela) e la spermateca molto più sviluppata e molto più complessa e
con un numero di spire più ampie.
Apimela luorum sp. n. (Figg. 186-189)
Holotypus 4, Kenya, MT. Elgon, 2400 m, For. Res, 14.X1.1974, (Mahnert & Perret
leg., MG).
Paratypi: 5 es., stessa provenienza; 4 es., Kenya, Embu, Irangi Forest, 2100 m,
12.X.1977, (Mahnert & Perret leg.).
Descrizione. Lungh. 2,6 mm. Corpo lucido e rossiccio sporco con margine
posteriore degli uroterghi ed estremità addominale giallo-rossicci; antenne brune con
l'antennomero basale giallo-rossiccio; zampe giallo-rossicce. La punteggiatura del
capo e del pronoto è svanita. Tubercoletti distinti coprono la superficie delle elitre e
dell'addome. Sul corpo non vi è traccia di reticolazione. Edeago figg. 187-188,
spermateca fig. 189.
Comparazioni. Poiché la nuova specie presenta elitre più corte del pronoto, è
nettamente distinta dalle specie finora note, che presentano elitre più lunghe del
pronoto. Anche la forma molto caratteristica dell'edeago e della spermateca della
nuova specie non trova riscontri in altre specie, anche in quelle attere qui di seguito
descritte.
Etimologia. La nuova specie prende nome dai Luo, gruppo etnico nilotico del
sud del M. Elgon, località tipica.
Apimela rangirensis sp. n. (Figg. 190-193)
Holotypus d, Rwanda, Rangiro, 1800 m, 6.VIII.1973, (Werner leg., MG).
Paratypi: 73 es., stessa provenienza; ma anche 25.111.1973 e 14.IX.1976.
Descrizione. Lungh. 2,1 mm. Corpo lucido e giallo-rossiccio con capo, elitre e
4° urite libero bruno-rossicci; antenne brune con l'antennomero basale giallo-
rossiccio; zampe gialle. La punteggiatura del capo è fine, fitta e distinta, quella del
pronoto è svanita. Le elitre presentano tubercoletti distinti; l'addome ha tubercoletti
superficiali. Edeago figg. 191-192, spermateca fig. 193.
Comparazioni. Anche questa nuova specie, come A. luorum sp. n. sopra
descritta, mostra elitre più corte del pronoto, perciò è ben distinta da A. subparallela
ALEOCHARINAE DELL’ AFRICA ORIENTALE 249
0) mm
Ficc. 190-195
Habitus, edeago in visione laterale e ventrale e spermateca. 190-193: Apimela rangirensis sp. n.;
194-195: Apimela kirimirensis sp. n.
250 ROBERTO PACE
(Bernhauer, 1938), dello Zaire, e da altre specie che presentano elitre più lunghe del
pronoto. In base alla forma dell'edeago e della spermateca, la nuova specie si
distingue chiaramente dalle specie con elitre più corte del pronoto qui descritte.
Apimela kirimirensis sp. n. (Figg. 194-195)
Holotypus 2, Kenya, Embu, Kirimiri Forest, O de Runyenye, 1550 m, 13.X.1977,
(Mahnert & Perret leg., MG).
Descrizione. Lungh. 2,1 mm. Corpo lucido e giallo-rossiccio con elitre bruno-
rossicce; antenne brune con 1 due antennomeri basali giallo-rossicci; zampe gialle. Il
capo e il pronoto presentano punteggiatura fitta e distinta. I tubercoletti che coprono
le elitre sono svaniti, quelli dell'addome sono distinti. Solo l'addome mostra una
distinta reticolazione: sul resto del corpo la reticolazione è assente. Spermateca fig.
IQS),
Comparazioni. Poiché la nuova specie ha elitre lunghe quanto il pronoto, è ben
distinta da A. subparallela (Bernhauer, 1938), dello Zaire, che ha elitre più lunghe del
pronoto. Inoltre la spermateca della nuova specie ha sviluppo molto maggiore e ha
bulbo prossimale ipertrofico, mentre la spermateca di subparallela è poco sviluppata,
con bulbo prossimale non ipertrofico.
Apimela perreti sp. n. (Figg. 196-197)
Holotypus ©, Kenya, Narok, Morijo Loita Hills, 2300 m, 4.1.1977, (Mahnert & Perret
leg., MG).
Paratypi: 4 9 ©, Kenya, Embu, Irangi Forest, 2100 m, 12 X.1977, (Mahnert & Perret
leg.).
Descrizione. Lungh. 2,3 mm. Corpo lucido e bruno con base ed estremità
addominali bruno-rossicci; antenne brune; zampe bruno-rossicce. L'avancorpo è
coperto di punteggiatura svanita, posta su un fondo non reticolato. L'addome presenta
reticolazione a maglie trasverse e svanite. Spermateca fig. 197.
Comparazioni. La taglia corporea maggiore e le elitre molto larghe rispetto al
pronoto, sono i caratteri più evidenti che permettono di distinguere a prima vista la
nuova specie da A. subparallela (Bernhauer, 1938), dello Zaire. Inoltre la spermateca
della nuova specie è molto più sviluppata di quella di subparallela, con matassa molto
più fitta.
Apimela zerchei sp. n. (Figg. 198-199)
Holotypus ©, Usambara (Coll. Kraatz, D.E.I.).
Descrizione. Lungh. 2, 5 mm. Corpo lucido e bruno-rossiccio; antenne bruno-
rossicce con 11° antennomero giallo-bruno; zampe giallo-rossicce. Tutto il corpo è
coperto di punteggiatura fine e fitta, posta su un fondo non reticolato. Spermateca fig.
199 (purtroppo amputata).
Comparazioni. La nuova specie ha elitre più lunghe del pronoto, ma non così
lunghe come quelle di A. subparallela (Bernhauer, 1938), dello Zaire. Inoltre il bulbo
ALEOCHARINAE DELL’ AFRICA ORIENTALE 251
Ficc. 196-203
Habitus, spermateca ed edeago in visione laterale e ventrale. 196-197: Apimela perreti sp. n.;
198-199: Apimela zerchei sp. n.; 200-203: Apimela kayovensis sp. n.
252 ROBERTO PACE
prossimale della spermateca della nuova specie € molto più sviluppato di quello
corrispondente della spermateca di subparallela.
Etimologia. La nuova specie è dedicata al Dr. Lothar Zerche, noto studioso di
Staphylinidae del D.E.I. di Eberswalde, che me l'ha affidata in esame.
Apimela kayovensis sp. n. (Figg. 200-203)
Holotypus d, Rwanda, Kayove, 2100 m, 15.V.1973, (Werner leg., MG).
Paratypi: 10 es., stessa provenienza; 1 2, Rwanda, Kamiranzoyu, 1900 m, marais,
1.1976, (Werner leg.).
Descrizione. Lungh. 1,8 mm. Corpo lucido e rossiccio con capo, elitre e 4°
urite libero, bruni; antenne brune con i due antennomeri basali giallo-rossicci; zampe
gialle. La punteggiatura del capo e del pronoto è fine e fitta. Le elitre sono coperte di
tubercoletti svaniti come quelli dell'addome. Sul corpo non vi è traccia di reti-
colazione. Edeago figg. 201-202, spermateca fig. 203.
Comparazioni. In base alla forma della spermateca e dell'edeago, la nuova
specie è sistematicamente vicina ad A. rangirensis sp. n., sopra descritta, tuttavia la
nuova specie presenta elitre molto più lunghe del pronoto, mentre in rangirensis le
elitre sono più corte del pronoto. La taglia della nuova specie è minore di quella di
rangirensis e l'edeago della nuova specie è meno profondamente ricurvo al lato
ventrale.
HOPLANDRIINI
Tinotus minutus Bernhauer, 1915
Tinotus minutus BERNHAUER, 1915: 158
1 ©, Kenya, Lac Naivasha, 5 Km Fischeman's Camp, 5.X1.1974, (Mahnert & Perret leg.).
Specie nota solo dell'Africa orientale.
Tinotus natalensis Pace, 1986 (Figg. 204-205)
Tinotus natalensis PACE, 1986: 107
1 2, Kenya, Nairobi, 2.X1.1974, (Mahnert & Perret leg.); 3 2 2, Rhodésie, Umtali,
11.1969, (R. Mussard leg.).
ALEOCHARINI
Aleochara (s. str.) kiambuensis sp. n. (Figg. 206-208)
Holotypus d, Kenya, Kiambu, près de Limuru, 2300 m, 2.X1.1977, (Mahnert & Perret
leg., MG).
Paratypus: 1 d, stessa provenienza.
Descrizione. Lungh. 4,8 mm. Corpo lucido e nero, con elitre giallo-brune
aventi i lati esterni bruni; antenne nere con i due antennomeri basali rossicci; zampe
ALEOCHARINAE DELL’ AFRICA ORIENTALE 253
FIGG. 204-208
Habitus, spermateca ed edeago in visione laterale e ventrale. 204-205: Tinotus natalensis Pace;
206-208: Aleochara (s. str.) kiambuensis sp. n.
254 ROBERTO PACE
FiGG. 209-214
Habitus, spermateca e 6° urotergo libero del maschio. 209-210: Aleochara (Xenochara) kivu
Lik.; 211-214: Apimela mahnerti sp.
ALEOCHARINAE DELL’AFRICA ORIENTALE 255
til
POD
by tyra
OL OR
FiGG. 215-219
Habitus, edeago in visione laterale e ventrale e spermateca. 215: Apimela mahnerti Sp. n.;
216-219: Apimela heteroclita sp. n.
256 ROBERTO PACE
bruno-rossicce. La punteggiatura dell'avancorpo è distinta. La punteggiatura sui due
uroterghi basali è separata dal margine posteriore da una fascia senza punteggiatura.
Edeago figg. 206-207.
Comparazioni. Specie con habitus più tozzo di quello di A. arrowi Bernhauer,
1940, dell'Uganda, con elitre più corte del pronoto (elitre lunghe quanto il pronoto in
arrowi). In visione ventrale, l'edeao della nuova specie, all'apice termina a punta
molto ogivale, mentre l'edeago di arrowi termina a punta molto aguzza. Inoltre il
sacco interno dell'edeago della nuova specie, possiede un lungo stiletto, mentre il
corrispondente pezzo in arrow! è corto € ricurvo
Aleochara (Xenochara) kivu Likovski, 1983 (Figg. 209-210)
Aleochara (Polychara) plagiata CAMERON, 1932: 42
Aleochara kivu LIKOVSKI, 1983: 16
Aleochara (Xenochara) kivu: KLIMASZEWSKI, 1993: 74
i 2, Tanzania, Arusha, XI.1972, (E.C. Supper leg., MG).
Specie nota del Sudafrica, Zaire, Rwanda e Kenya. Nuova per la Tanzania.
Aleochara (Xenochara) puberula Klug, 1833
Aleochara puberula KLUG, 1833: 139
1 ©, Kenya, Embu, pr. Ishiara, 900 m, 14.X.1977, (Mahnert & Perret leg.).
Specie cosmopolita diffusa in Europa, Africa e Nordamerica.
RINGRAZIAMENTI
Un sentito ringraziameto é rivolto al Dr. I. Löbl et al Dr. C. Besuchet del
Museo di Storia Naturale di Ginevra, che mi hanno concesso di studiare il materiale
oggetto del presente lavoro. Per il prestito di tipi e di materiale di confronto ringrazio
anche il Dr. L. Bartolozzi del Museo Zoolegico de “La Specola” dell'Università di
Firenze, il Dr. L. Zerche del D.E.I. di Eberswalde, il Dr. M. Uhlig del Museo
Zoologico dell'Università Humboldt di Berlino, il Dr. L. Baert dell'Institut Royal des
Sciences Naturelles di Bruxelles, il Dr. A.F. Newton del “Field Museum of Natural
History” di Chicago.
RESUME
Ce travail énumère 70 espèces dont 60 sont décrites comme nouvelles. Elles |
apartiennent aux tribus Thamiaraeini, Myrmedoniini, Oxypodini, Hoplandriini et |
Aleocharini. Huit holotypes ou lectotypes d'espèces décrites par Bernhauer, sont |
illustrés. Cinq nouvelles combinaisons sont effectuées.
ALEOCHARINAE DELL’ AFRICA ORIENTALE DST
BIBLIOGRAFIA
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Ergebnis deutsche Zentral-Afrika-Expedition 1907-1908: 469-486.
BERNHAUER, M. 1915. Zur Staphylinidenfauna des tropischen Afrika. Annales historico-
naturales Musei nationalis hungarici 13: 95-189.
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114-168.
FAUVEL, A. 1899. Genres et species de Staphylinidae nouveaux d’Afrique. Revue d’Entomo-
logie: 1-44.
GESTRO, R. 1892. Di alcuni Coleotteri raccolti nel paese dei Somali dall'Ing. L. Brichetti
Robecchi. Annali del Museo civico di Storia naturali di Genova 12: 747-790.
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II. The genera Aenictonia and Anommatochara-Their Relationship and Behavior.
Annals of the Entomological Society of America 61: 971-986.
KLIMASZEWSKI, J. & JANSEN, R.E. 1993. Systematics, Biology and Distribution of Aleochara
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(Coleoptera: Staphylinidae). Annals of Transvaal Museum 36: 53-107.
LAST, H.R. 1963. A Revision of the African Species of Zyras Stephens (Coleoptera, Sta-
phylinidae), subgenus Camonia Bernhauer. Revue de Zoologie et Botanique africaines
67: 251-299.
LIKOVSKY, Z. 1983. Bemerkungen über Aleochara-Arten der Afrikanischen Region (Cole-
optera, Staphylinidae). Annotationes Zoologiae et Botanicae 152: 1-18.
MULSANT, M.E. & REY, C. 1874. Tribu des Brévipennes: Famille des Aléochariens: Septième
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rico-naturales Musei nationalis hungarici 78: 83-143.
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Sociedad portuguese de Entomologia 99: 1-34.
258 ROBERTO PACE
PACE, R. 1994. Aleocharinae della Sottoregione Africana Orientale al Museo di Ginevra
(Coleoptera, Staphylinidae). Parte I. Revue suisse de Zoologie 100: 117-193.
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1501-1881, Berlin.
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Fieldiana Zoology 40: 334 pp., Chicago.
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Entomologische Nachrichten 2: 2-3.
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Aleocharinae (part.). Annales du Musée royal du Congo belge 58: 73-135.
REVUE SUISSE DE ZOOLOGIE, 103 (1) : 259-282; mars 1996
Oribatids from Sarawak I. (Acari: Oribatida).
New and interesting mites from the Geneva Museum LXX VIII
Sandor MAHUNKA
Zoological Department, Hungarian Natural History Museum, Baross utca 13,
H-1088 Budapest, Hungary.
Oribatids from Sarawak I (Acari: Oribatida). New and interesting mites
from the Geneva Museum LXXVIII. — Seventeen Oribatid species are
listed from Sarawak; ten of them are new to science and 2 also represent
new genera: Bakobodes gen. n. (Carabodidae ) and Sarawakiella gen. n.
(Galumnidae).
Key-words: Acari - Oribatida - Taxonomy - New species, new genera -
Sarawak.
INTRODUCTION
The present paper is the first contribution to knowledge of the Oribatid mites
collected by Dr. Bernd Hauser, Head of the Arthropod Department of the Museum
d’Histoire naturelle, Geneva, during his 1987 expedition to Sarawak, organized
together with Dr. Charles Lienhard, Research Officer at the same Department, as a
part of the program of systematic exploration of the Microarthropoda of the rain
forests of South-East Asia and surrounding areas.
The main goals and aims of this research program on Oribatids of South-East
Asia were summarized in my recent paper on Brunei (MAHUNKA 1995). These first
results on the Oribatids of Sarawak confirm the great importance of Borneo for a
sound interpretation of the historical zoogeography of South-East Asia.
The elaboration of this rich material requires prolonged studies and the results
will therefore be published in several parts. I discuss here the occurrence of 17
species, most of them belonging to the groups Ptyctima and Carabodida; 10 species
are new to science, 2 of which also represent new genera: Bakobodes gen. n.
(Carabodidae) and Sarawakiella gen. n. (Galumnidae).
In the descriptions I generally apply the terminology used in several publications
by NORTON (e.g. 1982) and BEHAN-PELLETIER (e.g. 1984) based on Grandjean’s work.
! Partly sponsored by the Hungarian National Scientific Research Fund (OTKA T-
016729).
Manuscrit accepted 20.04.1995.
260 SÄNDOR MAHUNKA
The pilosity of the parts of the body and of the legs is expressed in formulae. The
sequence of the anogenital formula is: number of genital, aggenital, anal and adanal
setae. Within the setal formula of the palp and the legs, the solenidia of a given
segment are marked with the symbol +. The measurements given correspond to
extremes observed in the present material; length is measured from the rostral apex to
the furthermost opposite point of the body, width refers to maximum body width (in
the case of movable pteromorphae to maximum width without pteromorphae).
LIST OF LOCALITIES
Sar-87/60 : MALAYSIA: Sarawak: Serian District, Penrissen Road 12 miles de Kuching,
“Semongok Wildlife Rehabilitation Centre: Nursery Centre of the Forest Department”,
prélèvement de sol dans les angles formés par les contreforts de grands arbres, 50 m;
8.XII.1987; leg. B.H. (B)?
Sar-87/64 : MALAYSIA: Sarawak: route Kuching-Matang, mont Gunung Serapi, prélèvement
du sol dans la forêt le long de la route vers la station TV, 670 m; 9.XII.1987; leg. B. H.
(B)?
Sar-87/66 : MALAYSIA: Sarawak: route Kuching-Matang, mont Gunung Serapi, prélèvement
du se dans la forêt le long de la route vers la station TV, 320 m; 9.XII.1987; leg. B. H.
(B)
Sar-87/76 : MALAYSIA: Sarawak: Bako National Park, Jalan Lintang, prélèvement de sol dans
les angles formés par les contreforts de Austrobuxus nitidus Miq. [= Longetia mala-
yana (Benth.) P. & H.] (Euphorbiaceae), 30 m; 11.XII.1987; leg. B. H. (B)3
LIST OF SPECIES
Hypochthoniidae Berlese, 1910
Malacoangelia remigera Berlese, 1913
Locality: Sar-87/64: 3 specimens.
Distribution: circumtropical.
Phthiracaridae Perty, 1841
Hoplophorella cucullata (Ewing, 1909)
Locality: Sar-87/64: 2 specimens.
Distribution: approximately circumtropical.
Kakophthiracarus enigmaticus sp.n.
Locality: Sar-87/60.
Hoplophthiracarus (Plonaphacarus) aculeatus Mahunka, 1995.
Localities: Sar-87/60: 1 specimen; Sar-87/64: 1 specimen;
Sar-87/76: 2 specimens.
Distribution: Brunei (known from the type locality only).
Notophthiracarus lienhardi sp. n.
Locality: Sar-87/60.
Synichotritiidae Walker, 1965
Sabahtritia sarawak sp. n.
Locality: Sar-87/64.
2 (B) = extraction par appareil BERLESE à Kuching (Sarawak).
3 (B) = extraction par appareil BERLESE à Genève.
= —
ORIBATIDS FROM SARAWAK I 261
Temburongiidae Mahunka, 1990
Temburongia patoi Mahunka, 1990
Locality: Sar-87/76: 2 specimens.
Distribution: Brunei (known from the type locality only).
Eremaeozetidae Balogh, 1972
Eremaeozetes maculosus Mahunka, 1995
Locality: Sar-87/64: 2 specimens.
Distribution: Brunei (known from the type locality only).
Carabodidae C. L. Koch, 1837
Bakobodes orangutan gen. n., sp. n.
Localities: Sar-87/64; Sar 87/76.
Berndobodes hauseri sp. n.
Localities: Sar-87/60; Sar-87/76.
Congocepheus orientalis Mahunka, 1987
Locality: Sar-87/76: 2 specimens.
Distribution: Sabah (known from the type locality only).
Gymnobodes semengok sp. n.
Localities: Sar-87/60; Sar-87/76.
Hardybodes penicillatus Mahunka, 1995
Locality: Sar-87/76: 2 specimens.
Distribution: Brunei (known from the type locality only).
Pasocepheus bako sp. n.
Locality: Sar-87/76.
Yoshiobodes humidus sp. n.
Locality: Sar-87/66.
Peloppiidae Balogh, 1943
Austroceratoppia serapi Sp. n.
Localities: Sar-87/64; Sar-87/66.
Galumnidae Jacot, 1925
Sarawakiella longipilosa gen. n., sp. n.
Localities: Sar-87/60; Sar-87/64.
DESCRIPTIONS
Kakophthiracarus enigmaticus sp. n. (Figs 1-5)
Material examined: Holotypus: Sar-87/60, 23 paratypes from the same
sample. Holotypus and 15 paratypes deposited in the MHNG4 and 8 paratypes (1416-PO-1991)
in the HNHMS.
Measurements. — Length of aspis: 178-252 um, length of notogaster:
312-485 um, height of notogaster: 223-322 um.
Integument: The whole body surface covered by a thick cerotegument
layer, extremely thick in the deeper hollows of the notogaster.
4 MHNG = deposited in the Muséum d’Histoire naturelle, Genève.
5 HNHM = deposited in the Hungarian Natural History Museum, Budapest, with identi-
fication number of the specimens in the Collection of Arachnida.
262 SANDOR MAHUNKA
Fics 1-3
Kakophthiracarus enigmaticus sp. n. -1: body in lateral view, 2: aspis in lateral view, 3: aspis in
frontal view.
Aspis: Quadrangular in lateral view (Fig. 2), crown-shaped in frontal view
(Fig.3). A very strong median crista present, parallel with it a pair of smaller and
shorter cristae observable on each side. Rostral part medially excavated, rostral setae
arising frontally. Lateral crista absent, sinus line observable. Bothridial squama
modified, not bending over the bothridium (Fig. 2). Aspidial surface mostly alveolate,
but smooth in its lateral and basal parts and also some rugae visible basally. Prodorsal
setae thin, simple.
ci
ORIBATIDS FROM SARAWAK I 263
zio oh
Fics 4-5
Kakophthiracarus enigmaticus sp. n. - 4: notogaster in dorsal view, 5: anogenital region.
Sensillus very long, setiform, distinctly spinose unilaterally.
Notogaster: With large and wide anterior hood, behind it a deep hollow
(Fig. 1) present. Surface with many tubercles besides two pairs of larger hollows, and
three dorsal crests; all conspicuous in dorsal view (Fig. 4). Fifteen pairs of curved,
thin and simple notogastral setae; the alveoli of the vestigial f setae, a glandular
opening and two pairs of lyrifissures ia, im also visible.
Anogenital region: The position of the genital setae is typical for
the genus, setae gg-g¢ arising much nearer to the inner margin of the genital plates
than setae g5-g,. (Exceptionally 10 pairs of setae were recognisable). Among the
anoadanal setae two pairs arising strictly on the inner margin (Fig. 5), but the other
three (ad,- ad3) appearing also not far from it.
Legs: Leg chaetotaxy of the “complete type”, with the setal formulae:
I: 1-4-4+2-5+1-17+3-1
IV: 2-1-1-2+1-10-1
Seta d on femur I hooked. A small spine in front of w, of tarsus I conspicuous. Setae
d on tibia IV coupled with the solenidium.
264 SÄNDOR MAHUNKA
Remarks: On the basis of the form and position of the setae in the anoa-
danal region and of the notogastral structure the new species can be placed into the
recently established genus Kakophthiracarus Mahunka, 1992. The new species is
readily distinguished from the related species by the three median longitudinal crests
on the notogaster, absent in its congeners.
Notophthiracarus lienhardi sp. n. (Figs 6-8)
Material examined: Holotypus: Sar-87/60, 1 paratype from the same
sample. Holotypus deposited in the MHNG and paratype (1417-PO-1991) in the HNHM.
Measurements.- Length of aspis: 280-297 um, length of notogaster:
619-660 um, height of notogaster: 355-379 um.
Aspis: Median crista very high and wide in dorsal view (Fig. 8), rounded.
Lateral carina absent, sinus line conspicuous. Surface mostly alveolate, its lateral part
polygonate, and basally some rugae also visible. Bothridial squama well developed.
Rostral setae setiform, simple, arising on the frontal surface and bent downwards.
Lamellar and interlamellar setae straight, erect, blunt at tip and clearly spiculate or
spinose. Sensillus slightly dilated distally, this part spiculate and spinose asymme-
trically.
Notogaster: Anterior part of notogaster with a hood, slightly dilated
laterally and continuing in a broad median band posteriorly (Fig. 6). Surface of this
band smooth, all other surfaces ornamented by very deep and well framed alveoli.
Seventeen pairs of straight and erect, apically blunt and fairly spiculate or spinose
notogastral setae present. I was able to find only two pairs of lyrifissures ia, im.
Anogenital region (Fig. 7): Nine pairs of genital setae arising in one
axial row, distance between gg and gs greater than the distance between g, and gy.
All five pairs of anoadanal setae originating near the median margin but an, and an,
clearly separated from the adanal setae. This marginal region framed laterally by a
longitudinal crest.
Legs: All claws with two ventral teeth. Chaetotaxy of legs of the “complete
type” with the setal formulae:
I: 1-4-2+2-5+1-16+3-1
IN 2 FEIN
Seta d of femur I hooked, arising far from anterior margin. Solenidium ©, coupled
with a small seta on tarsus I, solenidium ¢, on tibia IV also coupled with seta d. All
solenidia of legs conspicuously long and curved distally.
Remarks: Despite the setae of the anoadanal plates originating in a typical
position, this species is easily assignable to the genus Notophthiracarus Ramsay,
1966. It is well characterised by the smooth, longitudinal median field of the noto-
gaster. On this basis it is related to N. orientalis (Mahunka, 1985) and N. planus
(Mahunka, 1985) (see MAHUNKA 1995), but it is easily distinguished from both and
from all other Notophthiracarus species by the characteristic ornamentation (very
deep alveoli) and by the number (17 pairs) of notogastral setae.
ORIBATIDS FROM SARAWAK I 265
FIGS 6-8
Notophthiracarus lienhardi sp. n. - 6: body in lateral view, 7: anogenital region, 8: body in
dorsal view.
266 SANDOR MAHUNKA
Sabahtritia sarawak sp. n. (Figs 9-13)
Material examined: Holotypus: Sar-87/64, 1 paratype from the same sample.
Holotypus deposited in the MHNG and paratype (1418-PO-1991): in the HNHM.
Measurements.- Length of aspis: 117-121 um, length of notogaster:
219-226 um, height of notogaster: 110-119 um.
Aspis: Wide, low. Lateral carina fine, thin, fused with the lateral rim.
Posterior median and laterocentral apodemes long. Surface foveolate medially,
smooth anteriorly and laterally, and also some large but shallow areolae present ba-
sally (Fig. 12). Sensillus bifurcate (Fig. 10).
Notogaster (Fig. 9): Surface heavily foveolate. Fourteen pairs of simple
notogastral setae present. I was able to find only two pairs of lyrifissusres (ia, im).
Anogenital region (Fig. 13): The whole surface fairly foveolate.
Suture kag very strong. Seven pairs of genito-aggenital setae arising before them, all
minute, except the posteromedian one. Six pairs (3+3) of setae in anoadanal position,
setae an, distinguishable from the others, each arising on a small tubercle, directed
backwards and slightly dilated basally. All other setae simple. Lyrifissures iad origi-
nating on posterior part of anoadanal plates.
Gnathosoma: Palp (Fig. 11) 4-segmented with the setal formula: 0-2-1-
Ik
Legs: All claws large with strong teeth on their ventral surface. Solenidia
®|, ©, and ¢, on leg I, è, on legs IT and III with coupled minute setae, but seta d on
legs IV long and not coupled with the solenidium. Genu IV without seta or sole-
nidium.
Remarks: The new species is unique in the genus Sabahtritia Mahunka,
1987, owing to the simple notogastral setae and the furcate sensillus. This latter
feature is also unique in the whole superfamily Euphthiracaroidea Jacot, 1930.
Bakobodes gen. n.
Diagnosis: Family Carabodidae. Prodorsum with strong transversal ele-
vation, notogaster gradually convex, dorsosejugal region normal, but the notogaster
partly covers the prodorsum. Lamellae with well developed cuspis, lamellar setae
arising between them. Interlamellar setae originating on the elevation, on the lamellar
surface. Ten pairs of phylliform notogastral setae, one pair in humeral position.
Mentum with anterior tectum. Coxisternal region large, with wide median field.
Epimeral setal formula: 3-1-3-3. Anogenital region much smaller than coxisternal
one, with strong costulae or tubercles. Anogenital setal formula: 4-1-2-3.
Type species: Bakobodes orangutan sp. n.
Remarks: Among the Carabodidae genera characterised by ten notogastral
setae only Meriocepheus Aoki, 1973 has a highly elevated prodorsum. The new taxon
is distinguished from it by the absence of the very high notogastral elevation and by
the simple but characteristic dorsosejugal region.
ORIBATIDS FROM SARAWAK I 267
Fics 9-13
Sabahtritia sarawak sp. n. - 9: body in lateral view, 10: basal part of aspis in lateral view, 11:
palp, 12: aspis in dorsal view, 13: anogenital region.
268 SÄNDOR MAHUNKA
Fics 14-17
Bakobodes orangutan gen. n., sp. n. - 14: body in dorsal view, 15: body in ventral view, 16:
rostral region in frontal view, 17: body in lateral view.
ORIBATIDS FROM SARAWAK I 269
Fics 18-21
Berndobodes hauseri sp. n. - 18: body in dorsal view, 19: body in ventral view, 20: leg I, 21:
body in lateral view.
270 SANDOR MAHUNKA
Bakobodes orangutan sp. n. (Figs 14-17)
Material examined: Holotypus: Sar-87/64, 11 paratypes from the same
sample; 8 paratypes: Sar-87/76. Holotypus and 12 paratypes deposited in the MHNG and 7
paratypes (1420-PO-1991) in the HNHM.
Measurements. — Length of body: 207-243 um, width of body: 107-
134 um.
Integument: Whole surface covered by a thick cerotegument layer.
Prodorsum: Lamellae thick, connected by a translamella anteriorly (Fig.
14). Lamellar and interlamellar setae phylliform, both groups arising on the lamellar
surface (Fig. 16). Interlamellar region divided by a longitudinal crest anteriorly and a
semicircular hollow basally. Sensillus directed outwards, dilated, characteristicaly
bent downwards. Tutorium well developed, without cusp.
Notogaster: Ten pairs of characteristically arched, phylliform noto-
gastral setae present. Surface ornamented by large tubercles or pustules, ordered in a
polygonal formation (Fig. 14).
Lateral part of podosoma: Pedotectum | narrow, pedotecta 2-3
small but sharp laterally, discidium well developed (Fig. 17).
Gnathosoma: Mentum fairly foveolate, with anterior tectum.
Coxisternal region: Well framed laterally by crests. All epimeral
setae minute or observable only by their alveoli.
Anogenital region: Ventral plates with rough structure. Anal
opening framed by a strong crest posteriorly, some other crests and tubercles also
observable (Fig. 15).
Remarks: The new species is readily characterized by the arched tectum
of the mentum.
Berndobodes hauseri sp. n. (Figs 18-21)
Material examined: Holotypus: Sar- 87/76, 19 paratypes from the same
sample; 6 paratypes: Sar-87/60. Holotypus and 16 paratypes deposited in the MHNG and 9
paratypes (1421-PO-1991) in the HNHM.
Measurements.- Length of body: 252-382 um, width of body: 139-
203 um.
Integument: Body surface covered by an irregularly thick and disin-
tegrating waxy layer comprising also “broken fragments”. Cuticular surface mostly
sculptured.
Prodorsum: Rostrum convex, rostral setae thick, arising on small
tubercles, curved. Lamellae narrow, decurrent marginally, their surface rugose.
Lamellar setae dilated, arising on dorsal surface of lamellae, far from their cusps.
Interlamellar surface smooth. Basal part of prodorsum strongly concave on both sides;
phylliform interlamellar setae very large, arising on additional tubercles of prodorsum
(Fig. 18). Bothridium strongly protruding laterally, sensillus long, arched upwards,
narrow, only dilate distally. Tutorium strong, with a sharp apex (Fig. 21).
271
ORIBATIDS FROM SARAWAK I
Fics 22-24
Gymnobodes semengok sp. n. - 22: body in dorsal view, 23: body in ventral view, 24: body in
lateral view.
DIO SANDOR MAHUNKA
Notogaster: Well developed humeral projections present. Dorsosejugal
suture straight, anterior margin of notogaster with large pustules, their number in-
creasing laterally. Notogastral surface smooth. Fifteen pairs of dilated notogastral
setae present (Fig. 18), two pairs in humeral and four pairs in posteromargainal
positions, smaller than the others. Two pairs near the anterior margin long, narrow
phylliform. without cilia or spicules, four pairs medially, much larger than the others,
spatulate or cuneiform, their surface spiculate.
Lateral part of podosoma: Pedotectum | narrow, pedotecta 2-3
and discidium small. Sejugal region finely granulate.
Coxisternal region: Epimeral borders frame the epimeral fields,
enclosing medially comparatively large hollows situated behind each other. Epimeral
surface ornamented by irregular alveoli and spots. Epimeral setal formula: 3-1-3-3
(Fig. 19). Setae 3c originating far from pedotecta 2-3.
Anogenital region: Ventral plate framed by a pair of longitudinal
ribs continuing with some round tubercles along anal aperture. Around the latter a
semicircular thickening observable, adanal setae arising on it. Anogenital setal
formula: 4-1-2-3. Genital and aggenital setae simple, genital setae conspicuously
long. Anal plates with a thickening along their inner margin, bearing setae an].
Adanal setae spatulate.
Legs: Claws of all legs strong. Femur of legs IT and IV with strong blade-
like formation ventrally ending in a robust distal spur. Unguinal setae (u) of all legs
short, spiniform. Solenidia @, of tibia I very long, characteristically directed forwards
and decumbent on tarsus I (Fig. 20). Setae d on tibiae I and II coupled, with sole-
nidium >. Setal formulae of legs:
I: 1-4-3+1-4+2-15+2-1
II: 1-4-3+1-4+1-12+2-1
1271222112 10
Remarks: Until now this genus was considered as monotypic.The type-
species, B. spathuliferé Mahunka, 1986, was described from Sabah. The new species
stands near to it, but differs by the shape of the notogastral setae and the sculpture of
the body (e.g. the longitudinal crests and tubercles along the anal aperture).
Gymnobodes semengok sp. n. (Figs 22-24)
Material examined: Holotypus: Sar-87/76, 1 paratype: Sar 87/60. Holotypus
deposited in the MHNG and paratype (1422-PO-1991) in the HNHM.
Measurements. — Length of body: 263 um, width of body: 129 um.
6 On the occasion of the first diagnosis of the genus (MAHUNKA 1986) I erroneously
named the type species spathulifer and not spiculifer as initially planned. Based on the same
type material the species was later described in detail under the name spiculifer Mahunka,
1988, which has to be considered as a junior objective synonym of spathulifer Mahunka, 1986.
ORIBATIDS FROM SARAWAK I 273
Fics 25-28
Pasocepheus bako sp. n. - 25: body in dorsal view, 26: body in ventral view, 27: prodorsum in
frontal view, 28: body in lateral view.
274 SANDOR MAHUNKA
Prodorsum: Rostrum weakly convex in dorsal view, its surface alveo-
late. Lamellae thin, with clearly visible apex. Intermellar surface covered by large
pustules (Fig. 22). All three dorsal pairs of prodorsal setae long, thin, flagelliform,
distinctly ciliate. Sensillus short, its head funnel-shaped. Tutorium absent, only a short
crest in its place.
Notogaster: Whole surface pustulate, similar to that between the
lamellae. Ten pairs of long, thin, densely ciliate notogastral setae.
Lateral part of podosoma: Pedotectum 1 narrow, pedotecta 2-3
very small. The surface mostly alveolate (Fig. 24), only smaller fields appear to be
smooth.
Coxisternal region: Apodemes and epimeral borders long, almost
composing a close network. Surface irregularly alveolate. Epimeral setal formula: 1-
1-3-3, all setae conspicuous.
Anogenital region: Major part of this surface pustulate (like the
notogaster), only smaller fields alveolate behind legs IV laterally. All setae of this
region long, thin, ciliate (Fig. 23). (I was not able to find the lyrifissures iad).
Remarks: On the basis of the free cuspis on lamellae, the weak tutorium,
the epimeral setal formula and the absence of aggenital setae the new species is
assignable to the genus Gymnobodes Balogh, 1965 (MAHUNKA 1986). Both of the
heretofore known species were described from Africa and the new species is readily
separated from them by the pustulate surface and the thin and well ciliate dorsal and
ventral setae. This latter feature is unique in the entire family Carabodidae C. L.
Koch, 1837.
The species is named after the “Semongok Wildlife Rehabilitation Centre” for
orang-utangs.
Pasocepheus bako sp. n. (Figs 25-28)
Material examined: Holotypus: Sar 87/76, 21 paratypes from the same
sample. Holotypus and 14 paratypes deposited in the MHNG and 7 paratypes (1423-PO-1991)
in the HNHM.
Measurements. —Length of body: 312-346 um, width of body: 188-
267 um.
Integument: A finely granulate cerotegument layer covering the whole
surface, except the tarsi and tibiae.
Prodorsum: Basal part of prodorsum - with the lamellae - distinctly
protruding and arched abruptly to the rostrum (Fig. 28). Between them a characteristic
U-shaped formation is observable, but less conspicous than in the type-species. In the
rostral part of prodorsum, in the interlamellar region (Fig. 27), and also between
lamellae and tutorium, some large alveoli are present; the rest of the surface wrinkled
or smooth, like the basal surface medially. All prodorsal setae small, the rostral setae
dilated, phylliform, lamellar setae thin, pilose. Interlamellar setae also phylliform,
bent inwards. Sensillus long, strongly bent backwards (Fig. 27), nearly falciform.
Tutorium well developed, but without cusp.
ORIBATIDS FROM SARAWAK I 275
Fics 29-32
Yoshiobodes humidus sp. n. - 29: body in dorsal view, 30: body in ventral view, 31: sensillus,
32: rostral region in frontal view.
Notogaster: Median part of notogaster strongly protruding, but gra-
dually declining to dorsosejugal suture. The sculpture of this region very similar to
that of Pasocepheus eremaeozetoides Mahunka, 1995. Position and shape of setae (15
pairs) also very similar to those in that species (Fig. 25).
276 SÂNDOR MAHUNKA
Figs 33-35
Austroceratoppia serapi Sp. n. - 33: body in dorsal view, 34: body in ventral view, 35:
podosoma in lateral view.
ORIBATIDS FROM SARAWAK I 277
Lateral part of podosoma: Pedotectum 1 and discidium well
developed, pedotecta 2-3 comparatively small. Anterior margin of pedotectum 1
ornamented by foveolae.
Coxisternal region: Framed laterally by a heavy crest. Median part
slightly hollowed.
Anogenital region: Nearly entire surface ornamented with longer
or shorter crests (Fig. 26). setae of ventral regions small, adanal setae clearly phylli-
form.
Legs: Femora of legs alveolate, all other joints smooth.
Remarks: In most of its features the new species exhibits a great simi-
larity with P. eremaeozetoides Mahunka, 1995. But the two species are readily distin-
guished from each other by the form of the sensillus (straight and plumose in P. ere-
maeozetoides) and by the prodorsal protuberances (smaller and shorter in P. eremae-
ozetoides).
Yoshiobodes humidus sp. n. (Figs 29-32)
Material examined: Holotypus: Sar-87/66, 1 paratype from the same sample.
Holotypus deposited in the MHNG and paratype (1424-PO-1991) in the HNHM.
Measurements. —Length: 306-342 um, width: 183-209 um.
Integument: Surface covered by a cerotegument layer, consisting of
granules ordered in different polygonal sculptures.
Prodorsum: Rostrum rounded. Lamellae typical for the genus, their dis-
tal end — in front of lamellar setae — strongly narrowed anteriorly, with rostral setae
arising on this part (Fig. 32). Lamellar setae much longer than rostral ones, setae in
phylliform, the largest of all. Interlamellar region distinctly, lamellar surface irre-
gularly, foveolate. Basal part of prodorsum smooth, two lateral hollows, framed by
cristae, present. Dorsosejugal region narrow, but deeply excavated, full of secretion.
Sensillus (Fig. 31) dilated distally and bent backwards.
Notogaster: Cerotegumental granules composing or framing longi-
tudinal fields anteriorly and rounded fields medially; sculpture typical for the genus.
Fourteen pairs of approximately spatulate notogastral setae present, the medial ones
widened distally, with straight distal margin (Fig. 29) in median region, the lateral
ones smaller, narrowed distally, spoon-shaped. Two pairs of setae originating in
humeral position.
Ventral regions: (Fig. 30): Mentum and coxisternal region strongly
foveolate and alveolate. Sternal region wide, two rounded hollows observable medially.
All epimeral setae minute, epimeral setal formula: 3-1-3-3. Anogenital region pustulate,
and also some longitudinal rugae visible along genital plates and laterally; anogenital
setal formula: 4-1-2-3. Aggenital setae minute, adanal setae spatulate (Fig. 30).
Remarks: The new species stands nearest to Yoshiobodes aokii Mahunka,
1987, but it is well distinguished from the latter by the shape of the notogastral setae
(distally narrowed, phylliform in Y. aokii) and by the structure of the prodorsum.
278 SÄNDOR MAHUNKA
Fics 36-38
Austroceratoppia serapi sp. n. - 36: leg I, 37: leg IV, 38: trochanter and femur leg II.
ORIBATIDS FROM SARAWAK I 279
Austroceratoppia serapi sp. n. (Figs 33-38)
Material examined: Holotypus: Sar-87/66, 6 paratypes from the same
sample; 5 paratypes: Sar-87/64. Holotypus and 7 paratypes deposited in the MHNG and 4
paratypes (1419-PO-1991) in the HNHM.
Measurements: — Length of body: 426-481 um, width of body: 267-
292 um.
Prodorsum: Rostrum wide, with very sharp, small apıces medially and
laterally (Fig. 33). Rostral setae thick, spiniform, arising on anterior margin, near lateral
corner. Lamellae long, with long free cusps, but not reaching to rostral margin (Fig.
35). Their cusps with small, but well observable lateral apex. Lamellar setae thick, as
the rostral setae, and arısing from the cusps. A transversal band present between the
lamellae resembling a translamella, but not connected with the lamellae. Interlamellar
setae long, slightly thicker than setiform sensillus. Exobothridial setae reduced,
represented only by their alveoli. Tutorium short, weakly developed, without cusps.
Notogaster: Notogastral setae chatacteristically reduced, only two pairs
of robust, thick setae present. Five pairs of setae represented by their alveoli, each of
which is a porose area, a pair of (median) alveoli without porose area (Fig. 33)
observable.
Lateral part of podosoma: Very strong and long genal teeth
directed forwards, reaching over rostral margin. Pedotectum | with a sharp dorsal
spur, pedotecta 2-3 small.
Coxisternal region: Apodemes and epimeral borders weakly
developed, short, not forming a transversal band. All epimeral setae characteristically
directed forwards, some of them (e. g. /b, 3b) robust, setae 3c and 4c short and thin.
All fairly ciliate.
Anogenital region: A thin, strongly arched transversal line decur-
rent from genital aperture to acetabulum of legs IV, a secondary line partly parallel
with the former one also present (Fig. 34). Circumpedal carina absent. All setae of
this region directed forwards, mostly setiform, but adanal setae spiniform or bacilli-
form, blunt at tip and much thicker than the rest. Lyrifissures iad in apoanal position.
Legs: All legs tridactylous. Setae (p) on tarsus I thick, straight, on tarsus H-
IV very thin, arising near claws, and similar to them. Trochanter of leg III with very
strong seta (Fig. 38). Setal formulae of legs:
I: 1-5-3+1-4+2-20+2-3 (Fig. 36)
IV: 1-2-3-3+1-12-3 (Fig. 37)
Remarks: The new species is readily distinguished from all related
species by the equal length of the adanal setae.
The new species is named after the mountain Serapi.
Sarawakiella gen. n.
Diagnosis: Family Galumnidae. Cuticula finely punctate, without heavy
sculpture. Rostrum sharply pointed. Lamellar and sublamellar lines well developed,
280 SÄNDOR MAHUNKA
slightly diverging from each other. Lamellar setae arising on median surface, between
lines L. Rostral setae arising laterally between lines L and S. Dorsosejegal suture
absent. Ten pairs of long flagellate notogastral setae and four pairs of not typical areae
porosae present. They are characteristic for the new genus. Epimeral setal formula: 1-
0-1-2. Adanal setae reduced or minute. Lyrifissures iad in adanal position. Area
porosa postanalis present. Chelicera of normal type. Legs tridactylous. Famulus
originating far from solenidia, setae ft’ short and almost coupled with solenidium ©.
Solenidium , arising laterally and slightly posteriorly from ).
Type species: Sarawakiella longipilosa sp. n.
Remarks: On the basis of the shape and position of the octotactic organs
the new taxon is related to Pilizetes Sellnick, 1937. However, the type-species of the
latter genus is characterised by a rounded rostrum, a strong notogastral sculpture, and
rigid notogastral and adanal setae.
Sarawakiella longipilosa sp. n. (Figs 39-44)
Material examined: Holotypus: Sar-87/60, 7 paratypes from the same
sample; 2 paratypes: Sar-87/64. Holotypus and 6 paratypes deposited in the MHNG and 3
paratypes (1425-PO-1991) in the HNHM.
Measurements. — Length: 252-302 um, width of body: 208-238 um.
Prodorsum: Rostrum with sharply pointed apex (Fig. 41). Lamellar and
rostral setae simple, setiform, long. Rostral setae arising between the lines L and S
(Fig. 42). Interlamellar setae very long, flagellate, their basal part darker than the rest.
Sensillus long directed backwards, its head comparatively small, smooth, asymmetric,
with a long spine on its distal end. Areae porosae dorsosejugales, small, elliptic.
Notogaster: Ten pairs of long, flagellate setae (like the interlamellar
ones), 4 pairs of small, hardly recognizable areae porosae, more resembling sacculi,
each with a long tubular formation in the cuticule, but the unframed pori well cons-
picuous on the surface (Fig. 39). All lyrifissures very narrow, inconspicuous, ih and
ips standing near each other transversally, im located near areae porosae Aj.
Ventral region (Fig. 43): Epimeral setal formula: 1-0-1-2, setae 4c
arising on discidium. Anoadanal region with anterior three pairs of genital setae
visible, all others minute or observable only by their alveoli.
Legs: All legs tridactylous. Setal formulae of legs:
I: 0-4-3+1-4+2-20+2-3 (Fig. 44)
II: 0-4-3+1-4+1-15+2-3
III: 1-2-1+1-3+1-15-3
IV: 1-2-2-3+1-12-3
The position of the solenidial group of tarsus I shown in Fig. 40.
Remarks: On the basis of the very characteristic dorsal chaetotaxy the |
new species is far removed from all known species of the family Galumnidae.
ORIBATIDS FROM SARAWAK I 281
Fics 39-42
Sarawakiella longipilosa gen. n., sp. n. - 39: body in dorsal view, 40: basal part of tarsus I, 41:
lateral part of prodorsum, 42: podosoma in lateral view.
282 SÄNDOR MAHUNKA
Fics 43-44
Sarawakiella longipilosa gen. n., sp. n. - 43: body in ventral view, 44: leg I.
REFERENCES
AOKI, J. 1973. Oribatid mites from Iriomote-jima, the southernmost island of Japan (1).
Memoirs of the National Science Museum Tokyo 6: 85-101.
BEHAN-PELLETIER, V.M. 1984. Ceratozetes (Acari: Ceratozetidae) of Canada and Alaska.
Canadian Entomologist 116: 1449-1517.
MAHUNKA, S. 1985. Neue und interessante Milben aus dem Genfer Museum LIV. Oribatids
from South India I (Acari:Oribatida). Revue Suisse de Zoologie 92: 367-383.
MAHUNKA, S. 1986. A survey of the family Carabodidae C. L. Koch, 1836 (Acari: Oribatida).
Acta Zoologica Hungarica 32: 73-135.
MAHUNKA, S. 1987. Neue une interessante Milben aus dem Genfer Museum LX. Oribatids from
Sabah (East Malaysia) II. (Acari: Oribatida). Revue Suisse de Zoologie 94: 765-817.
MAHUNKA, S. 1988. New and interesting mites from the Geneva Museum LXI. Oribatids from
Sabah(East Malaysia) III (Acari: Oribatida). Revue Suisse de Zoologie 95: 817-888.
MAHUNKA, S. 1990. A survey of the superfamily Euphthiracaroidea Jacot, 1930 (Acari: Oriba-
tida). Folia Entomologica Hungarica 51: 37-80.
MAHUNKA, S. 1992. Notes and remarks on Oribatid taxa (Acari), Il. Annales Historico-
Naturales Musei Nationalis Hungarici 84: 161-183.
MAHUNKA, S. 1995. Oribatids from Brunei I. (Acari: Oribatida) — New and interesting mites
from the Geneva Museum LXXV. Revue Suisse de Zoologie 102: 913-942.
NORTON, R.A. 1982. Arborichthonius n. gen., an unusual enarthronote soil mite (Acarina: Ori-
batei) from Ontario. Proceedings of the Entomological Society of Washington 84: 85-96.
SELLNICK, M. 1937. Eine neue Milbe aus Ostafrika (Acar. Oribat.). Zoologischer Anzeiger 117:
130-132.
REVUE SUISSE DE ZOOLOGIE, 103 (1) : 283-297; mars 1996
Contribution à la connaissance des Cholevidae du Japon
et de Taiwan (Coleoptera)
Michel PERREAU
Laboratoire de Magnétisme des surfaces, Université Paris 7
2, place Jussieu, F-75251 Paris cedex 05, France.
Contribution to the knowledge of the Cholevidae of Japan and Taiwan
(Coleoptera). - This paper contains the description of 8 new species of
Cholevidae from Japan and Taiwan: Ptomaphagus smetanai n. sp., Ptoma-
phaginus ishizuchiensis n. sp., Ptomaphaginus nipponensis n. sp., Ptoma-
phaginus pingtungensis, n. sp., Anemadus nipponensis n. sp., Anemadus
taiwanus n. sp., Anemadiola smetanai n. sp., Nargus taiwanensis, n. Sp.
New distributional data are given for the following species: Ptomaphagus
amamianus Nakane, Ptomaphaginus takaosanus Nakane, Ptomaphaginus
sauteri (Portevin), Micronemadus pusillimus Kraatz, Nemadus japanus
Coiffait et Ueno, Anemadiola itotateoi Hayashi, Sciodrepoides fumatus
Spence, Mesocatops japonicus Jeannel, et Catops miensis formosensis
Hayashi.
Key-words: Coleoptera - Cholevidae - Japan - Taiwan - new species.
INTRODUCTION
Les travaux consacrés à la faune des Cholevidae extrême orientale, particuliè-
rement Japonaise et taiwanaise, se sont multipliés ces derniers temps (SZYMCZA-
KOWSKI 1962, 1963, 1965; NAKANE 1963, 1982, HAYASHI 1969, 1985, 1986, 1987,
1988, 1990; Mryama 1985; NISHIKAWA 1983, 1986; Perreau 1992), et ont permis
d’apprécier son intérêt biogéographique et phylogénétique. Ils ont en effet mis en
évidence trois genres endémiques, dont deux pour le Japon: Cholevodes Portevin, et
Apterocatops Miyama, et un commun au Japon et à Taiwan: Anemadiola Szymcza-
kowski. D’autres lignées très diverses se rencontrent dans ces îles, combinant des
groupes paléarctiques (Catops Paykull, Ptomaphagus Illiger) et tropicaux (Ptoma-
phaginus, Micronemadus).
Le présent article contribue à l’inventaire de cette faune en ajoutant 8 espèces
nouvelles: Promaphagus smetanai n. sp., Ptomaphaginus ishizuchiensis n. sp., Ptoma-
phaginus nipponensis n. sp., Ptomaphaginus pingtungensis n. sp., Anemadus taiwanus
Manuscrit accepté le 20.02.1995.
284 MICHEL PERREAU
n. sp., Anemadus nipponensis n. sp., Anemadiola smetanai n. sp., et Nargus taiwa-
nensis n. sp., et en précisant certains détails sur la distribution géographique d’espèces
encore mal connues. Certaines espèces voient donc leurs aires de répartition s’étendre.
C’est ainsi le cas de Ptomaphagus amamianus Nakane, 1963, connu jusqu’à présent
du Japon, et retrouvé maintenant à Taiwan. De même, Ptomaphaginus takaosanus
Nakane, 1982, Ptomaphaginus sauteri (Portevin, 1914), Nemadus japanus Coiffait et
Ueno, 1955, Micronemadus pusillimus Kraatz, 1877, Anemadiola itotateoi Hayashi,
1990, Sciodrepoides fumatus Spence, 1815, Mesocatops japonicus Jeannel, 1936, et
Catops miensis formosensis Hayashi, 1988, ont été retrouvées dans différentes
localités japonaises et taiwanaises.
Parmi les espèces nouvelles, certaines ne font que prolonger la liste déjà
longue de groupes habituellement rencontrés en l’Asie, c’est le cas des 3 espèces de
Ptomaphaginus. En revanche, d’autres s’avèrent particulièrement intéressantes.
L’Anemadus nipponensis n. sp. est en effet la première espèce du genre Anemadus
récoltée au Japon où se rencontre par contre le genre monospécifique apparenté
Cholevodes. Ce dernier était jusqu'à présent le seul représentant des Anemadini dans
ces deux îles, puisqu’Anemadiola Szymczakowski, considéré lors de sa description
comme un Anemadini serait en fait, d’après GIACHINO & VAILATI (1993), apparenté
aux Nemadini. De même, l’Anemadus taiwanus n. sp. est la première espèce d’ Ane-
madini taiwanaise. Cette espèce, bien caractérisée, ne s’apparente a aucune autre
lignée d’Anemadus (GIACHINO & VAILATI, 1993).
Dans un autre genre également, la faune taiwanaise exhibe son originalité.
Ainsi, le Nargus taiwanensis, n. sp. est l’espèce la plus orientale connue dans le genre
Nargus, et présente des caractères qui l’éloignent de toutes les autres espèces. Ce
genre, essentiellement rencontré dans les régions méditerranéennes et en Asie
mineure, ne comprenait jusqu'à présent, que deux représentants orientaux: N. beatus
Szymezakowski, et N. besucheti Perreau, qui ne dépassent pas vers l'est la zone
septentrionale de la péninsule indienne.
La richesse de la faune chinoise continentale, qui commence à être inventoriée
en détail, et qui présente d'évidentes parentés avec celle de Taiwan et du Japon,
permet de présumer pour ces îles, d'autres intéressantes découvertes à l'avenir.
Abréviations utilisées : MHNG : Muséum d'Histoire naturelle de Genève ;
MNHNP : Muséum National d'Histoire Naturelle de Paris; CMP : collection Michel
Perreau ; CJF : collection Jürgen Frank.
TAXONOMIE
Ptomaphagus amamianus Nakane
Taiwan: Fenchihu, 1400 m, 25-V-1977, Klapperich leg. (5 ex. CJF, 2 ex. CMP, 1 ex.
MHNG)
Cette localité est la première citation de cette espèce dans l'île de Taiwan.
Les exemplaires sont en tous points semblables à ceux du Japon.
CHOLEVIDAE DU JAPON ET DE TAIWAN 285
Ptomaphagus smetanai n. sp. (Figs 1-5)
HOLOTYPE d: Taiwan, Hualien Hsien, Taroko, N. P. Duodyatunshan, 2660 m, 8/13-V-
1990 A. Smetana (MHNG).
Description: longueur: 4 mm. Espèce aptère. Corps très convexe. Coloration
générale brun foncé, les antennes et les tarses plus clairs. Tous le dessus du corps sauf
la téte recouvert d’une fine pubescence, et d’une striolation transversale fine et peu
espacée.
Téte à ponctuation fine, alignée en strioles transversales superficielles au voisi-
nage de la carène occipitale, et évoluant vers une ponctuation grosse et désordonnée
vers l’avant du front. Yeux petits relativement à la taille du corps: la largeur d’un oeil
en vision dorsale représente environ un dixième de la largeur totale de la tête au
niveau des yeux. Les trois premiers articles antennaires plus longs que larges, tous les
suivants transverses. Longueurs des articles antennaires en microns: 220, 155, 95,
115,95, 155, 70, 140, 130, 190 (précision: 10 microns).
Pronotum 1,7 fois plus large que long, légèrement rétréci à la base, la plus
grande largeur est située à une distance de la base égale au dixième de sa longueur.
Les angles postérieurs peu saillants en arrière.
Elytres 1,2 fois plus longs que larges, leur plus grande largeur au quart
antérieur. Striolation de même espacement longitudinal que celle du pronotum.
Tibias intermédiaires très arqués. Les tibias postérieurs sans courbure ni
épaississement particulier. Tarses antérieurs fortement dilatés.
Segment abdominal IX très sclérifié, les latérotergites en forme de coquille très
convexes, et le spiculum gastrale courbé en vision latérale, dépassant de plus de la
moitié de sa longueur le bord antérieur du segment (figs 3 , 4 et 5).
Edéage robuste long, et arqué, l’apex détourné vers la droite à l’apex (figs 1 et
2). Quelques soies fines sur les côtés, juste avant l’apex. Paramères libres (non
fusionnés avec le lobe médian) sur les trois quart de leur longueur.
Femelle inconnue.
Cette espèce est étroitement apparentée au P. yasutoshii Nishikawa. Elle s’en
distingue par la forme de l’édéage dont la bifurcation apicale vers la gauche est moins
prononcée et la forme du corps plus trapue, moins allongée.
Ptomaphaginus takaosanus Nakane (Figs 13, 18, 19)
Japon : Nagano, J. E. Kongen N. Park, Shiga, 1500 m, I. Löbl (12 ex. MHNG, et 3 ex.
CMP). Japon : Gunma, ss/Usui pass, 850 m, I. Löbl (3 ex. MHNG, et 1 ex. CMP).
Le mâle de cette espèce présente un enfoncement du ventrite VIII bordé de
petits tubercules (fig. 19). De tels ornements sur les derniers ventrites visibles ont été
observés récemment sur plusieurs espèces et s’avèrent de bons caractères spécifiques
chez les Ptomaphaginus. Ils sont sans doute plus fréquents qu’il n'apparaît à la lecture
de la littérature, et existent probablement chez des espèces connues où ils n’ont pas
été décrits.
286 MICHEL PERREAU
——
Fics 1 à 5
Ptomaphagus smetanai n. sp. 1: édeage face dorsale. 2: édeage face latérale. 3: segment IX face
dorsale. 4: spiculum gastrale face latérale. 5: segment IX face latérale droite. L’échelle
représente 0,5 mm.
On retrouve une conformation semblable chez les mâles de P. franki Perreau,
et P. trautneri Perreau. La première de ces deux espèces se rencontre à Taiwan, la
seconde vient de Thaïlande. Le segment XI du mâle est représenté sur la figure 18, et
la spermathèque sur la figure 13.
Ptomaphaginus ishizuchiensis n. sp. (Figs 6, 9, 12, 15)
HOLOTYPE d : Japon: Ehime, via Mt. Ishizuchi, 1000 m, 14-8-1980, I. Löbl (MHNG).
Paratypes de même provenance: 2 2 2 (MHNG, et CMP).
Description: longueur: 1,9 mm. Coloration brune, l'avant du pronotum et la
tête plus foncés, les antennes et les tarses antérieurs uniformément brun clair. Tout le
corps recouvert d’une fine pubescence couchée dorée.
Tête couverte d’une ponctuation fine alignée transversalement, mais non
confluante en strioles transversales nettes. Tout au plus quelques traits transversaux
éparses et très superficiels relient certains points entre eux. Antennes courtes à massue
compacte.
nee
CHOLEVIDAE DU JAPON ET DE TAIWAN 287
Pronotum à côtés rétrécis de la base à l’apex, la plus grande largeur à la base,
recouvert de strioles transversales.
Elytres couvertes de strioles transversales plus espacées que celles du
pronotum. Côtés presque parallèles sur la moitié basale.
Tarses antérieurs dilatés, aussi larges que l'apex des tibias.
Segment IX représenté sur la figure 15.
Edéage allongé et régulièrement arqué, arrondi à l’apex (figs 6 et 9). Trois
soies du côté droit, quatre du côté gauche, toutes insérées sur la face inférieure et
dirigées vers le bas.
Femelle différente extérieurement par l'absence de dilatation des tarses anté-
rieurs. La spermathèque est représentée sur la figure 12.
Cette espèce est apparentée aux deux suivantes mais s’en distingue facilement
par l’édéage dont l’apex comporte trois soies latérales et ventrales, et une expansion
dorsale (figs 6 et 9).
Ptomaphaginus nipponensis n. sp. (Figs 7, 10, 16)
HOLOTYPE d : Japon: Ehime, via Mt. Ishizuchi, 1000 m, 14-8-1980, I. Löbl (MHNG).
Description: longueur 2 mm. Coloration brun clair uniforme (même les yeux),
deux taches latérales symétriques pigmentées sur les côtés du clypéus. La coloration
claire est probablement due à l'immaturité du seul exemplaire connu. Pubescence
uniforme courte et couchée. Strioles des élytres et du pronotum de même espacement.
Pronotum et élytres à plus grande largeur à la base, régulièrement rétrécis de la
base vers l’apex.
Tarses antérieurs dilatés, mais moins larges que l’apex des tibias.
Ventrite VIII avec une fossette médiane bordée de fortes soies. La confor-
mation de cette fossette ressemble à celle du Promaphaginus takaosanus, mais elle est
moins étendue chez P. nipponensis n. sp, et n'est pas exactement circulaire, légère-
ment plus allongée dans l'axe du corps.
Segment IX représenté sur la figure 16. Le spiculum gastrale est effilé mais ne
dépasse pas vers l’avant le bord antérieur du segment.
Edéage allongé et régulièrement élargi de la base vers l'apex, et terminé en
palette plate et arrondie (figs 7 et 10). Stylet interne épais.
Femelle inconnue.
Probablement apparentée au P. takaosanus Nakane et au P. ishizuchiensis n.
sp., cette espèce s’en distingue facilement par l’édéage nettement plus élancé dans sa
partie basale.
Ptomaphaginus pingtungensis n. sp. (Figs 8, 11, 14, 17)
Holotype d: Taiwan: Pingtung Hsien, Pietawushan, trail at 2000 m, 23-V-1991, A.
Smetana (MHNG). Paratypes : même provenance, 6 ex.; même provenance, 1500 m, 1-V-1992,
1 ex.; Taiwan: Pingtung Hsien, Pietawushan, Kuai-Ku Hut, 2325 m, 21-V-199i, A. Smetana,
10 ex.; même provenance, 2125-2130 m, 27-30/IV/1992, 21 ex. (MHNG, MNHNP, et CMP).
288 MICHEL PERREAU
Description: longueur: 2,2 mm. Ailé. Coloration générale uniformément
brune, la massue antennaire un peu plus foncée. Tout le corps recouvert d’une fine
pubescence couchée dorée. Les insertions des soies constituant cette pubescence
s’alignent le long des strioles.
Tête à ponctuation fine, les points plus ou moins alignés en strioles très
superficielles. Antennes au cinquième article plus allongé que le quatrième.
Pronotum ample, plus large que les élytres, et englobant ces derniers, les
angles postérieurs légèrement saillants en arrière, 1,5 fois plus large que long. Les
côtés presque parallèles sur la moitié postérieure de leur longueur, juste un peu
rétrécis à la base.
Elytres courts, 1,8 fois plus longs que le pronotum. La plus grande largeur
voisine de la base, régulièrement arqués. L’intervalle entre deux strioles transversales
de même taille que celui du pronotum.
Tarses antérieurs largement dilatés, à peu près aussi larges que l’apex des
tibias.
Segment abdominal IX représenté sur la figure 17. Spiculum gastrale long et
fin, ne dépassant pas vers l’avant le bord antérieur du segment.
Edéage ovale, peu arqué, peu rétréci à l’extrémité (figs 8 et 11).
Femelle semblable à l’exception de la dilatation des tarses antérieurs. La
spermathèque est représentée sur la figure 14.
Cette espèce se reconnaît facilement par sa formule antennaire particulière. Le
cinquième antennomère est plus allongé que le quatrième. Chez les Ptomaphaginae,
les antennomères sont généralement de longueur régulièrement décroissante de la
base vers l’apex des antennes, entre le troisième article et le sixième.
Ptomaphaginus sauteri (Portevin)
Taiwan: Taichung Hsien, Wufeng 60 m, 24-IV/2-V-9192, A. Smetana, 6 ex. (MHNG
et CMP).
Le ventrite VIII du mâle présente une fossette large et peu profonde.
HAYASHI (1990) a représenté l’édéage d’une espèce de Ptomaphaginus taiwa-
naise qu'il attribue au Promaphaginus sauteri et le compare au dessin de l’édéage
donné par JEANNEL (1936). La forme générale est manifestement différente, et 1l est
douteux que ces exemplaires soient réellement des P. sauteri. D’après le dessin,
l’exemplaire représenté se rapporte plus vraisemblablement au P. pingtungensis n. Sp.
On peut d’ailleurs se demander si les variations importantes, observées par cet auteur
dans les longueurs relatives des antennomères entre les trois exemplaires mentionnés
dans son article, ne sont pas dues à un mélange de spécimens appartenant aux deux
espèces.
Fics 6 à 11: édéages faces ventrales et latérales. 6 et 9 : Promaphaginus ishizuchiensis n. sp., 7 |
et 10: Ptomaphaginus nipponensis n. sp., 8 et 11: Ptomaphaginus pingtungensis n. sp. Fics 12 à |
14: spermathèques. 12: Promaphaginus ishizuchiensis n. sp., 13: Ptomaphaginus takaosanus |
Nakane, 14: Ptomaphaginus pingtungensis n. sp.
290 MICHEL PERREAU
Ptomaphaginus franki Perreau
Taiwan : Kaohsiung Hsien, Tengchih, 1610 m, 24-IV-1980, A. Smetana, 1 d; Pingtung
Hsien, Peitawushan, Kuai-ku Hut, 2130m, 27-IV-1992, A. Smetana, 1 d (MHNG).
Micronemadus pusillimus (Kraatz)
Japon: Gunma, ss/Usui pass, 250 metres, 24-7-1980, I. Lobl, 2 ex. (MHNG).
Taiwan: Pingtung Hsien Peitawushan, Kuai-Ku Hut,, 2325 m, 21-5-1991, A. Smetana,
1 ex.; Nantou Hsien, Shanlicai, 1650 m, 19-5-1991, A. Smetana, 2 ex (MHNG).
Nemadus japanus Coiffait & Ueno
Un d: Japon: Ehime, via Mt. Ishizuchi, 1000 m, 14-8-1980, I. Löbl (MHNG).
Anemadiola itotateoi Hayashi (figs 20, 22)
Taiwan: Taoyuan Hsien, Takuanshan for. 17-IV-1990, 1660 m, A. Smetana, 1 d et 1 £
(MHNG).
L’édéage est représenté sur les figures 20 et 22 par comparaison avec l’espèce
suivante. :
Anemadiola smetanai n. sp. (figs 21, 23, 27)
HOLOTYPE d: Taiwan, Nantou Hsien, Houhuanshan, 3175 m, 15-V-1990, A. Smetana
(MHNG). Paratypes: 1 4, même origine; 1 femelle, Taiwan: Kaohschiung Hsien, Kuanshan
trail, above Kaunshanchi Riv. 2550 m, 21-IV-92 A. Smetana; 4 ? ®, Taiwan, Nantou Hsien,
Houhuanshan, Kuenyang, 3050 m, 27/29-IV-90, A. Smetana (MHNG et CMP).
Description: longueur: 2,2 mm. Corps brun clair recouvert d’une pubescence
assez longue et partiellement dressée.
Tête fortement ponctuée, à suture clypéo-frontale bien marquée et plus
pigmentée que la surface céphalique.
Pronotum à ponctuation rugueuse, non striolé transversalement, les côtés
sinués près des angles postérieurs, sans fossette latérale. Les angles postérieurs droits,
et la base nettement rebordée.
Elytres striolés transversalement, la pubescence alignées sur les strioles. Stries
élytrales longitudinales finement ponctuées. Ailes postérieures réduites.
Tibias antérieurs étroits à la base, brusquement élargis dans le premier tiers.
Tarses antérieurs fortement dilatés, aussi larges que l’apex des tibias. Tarses inter-
médiaires avec les deux premiers articles dilatés (le premier nettement plus dilaté que
le second).
Edéage large à la base, brusquement rétréci latéralement au milieu, effilé et
recourbé vers le bas à l’apex (figs 21 et 23), moins effilé toutefois que chez l’Ane-
madiola itotateoi Hayashi.
CHOLEVIDAE DU JAPON ET DE TAIWAN 291
TO
18
Fics 15 à 18: segment IX, face dorsale. 15: Ptomaphaginus ishizuchiensis n. sp., 16: Ptoma-
phaginus nipponensis n. sp., 17: Ptomaphaginus pingtungensis n. sp., 18: Ptomaphaginus
takaosanus Nakane. Fics 19: Ptomaphaginus takaosanus Nakane, ventrite VIII du mâle.
L’échelle représente 0,25 mm pour les figures 15 a 18, et 0,3 mm pour la figure 19.
Femelle sans dilatation des tarses antérieurs et intermédiaires, avec un
spiculum ventrale plus long que large, arrondi a l’apex (fig. 27).
L’Anemadiola itotateoi et l’Anemadiola smetanai n. sp sont étroitement
apparentées, elles présentent une forme similaire d’édéage effilée et recourbée vers le
bas à l’extrémité, et un spiculum ventrale plus long que large, mais arrondi à l’apex.
Anemadiola smetanai n. sp. diffère de Anemadiola itotateoi par la sinuosité plus
faible des angles postérieurs du pronotum, l’absence de gros points enfoncés le long
des stries longitudinales élytrales, et l’absence de fovéole sur le pronotum.
La caractéristique la plus importante de ce genre est la morphologie de
l’édéage, très homogène chez les trois espèces connues. La seconde moitié du lobe
292 MICHEL PERREAU
médian est comprimée latéralement, recourbée vers le bas, non coplanaire avec les
paramères, et l’apex légèrement dilaté en palette. Le sac interne est tapissé de courtes
écailles hyalines sur toute sa surface et de phanères constituées de grosses épines
pigmentées. Les différences portent sur la courbure de l’apex du lobe médian, très
prononcée chez smetanai, un peu moins chez inordinata, et encore moins chez
itotateoi, et sur le rétrécissement de l’apex du lobe médian, régulier chez smetanai,
plus brusque chez inordinata, et itotateoi.
La morphologie de l’édéage évoque certaines espèces de Nemadini, toutefois
les deux espèces taiwanaises présentent tous les caractères des Anemadini : lame
basale du tegmen courte, suture clypéo-frontale présente, non réduite à une ligne à
pigmentation plus prononcée, spiculum ventrale arrondi et non pas en forme d’épine,
et enfin, dilatation des deux premiers tarsomères intermédiaires chez les mâles.
Ce genre reste encore à découvrir en Chine continentale où il est très
certainement présent.
Le tableau de détermination suivant permet d'identifier facilement les trois
espèces actuellement connues d’Anemadiola.
TABLEAU DE DETERMINATION DES ESPECES DU GENRE Anemadiola
Il Forme large et convexe, côtés du pronotum non sinués devant les
angles postérieurs. Stries élytrales avec des gros points enfoncés
RE A O N ae inordinata Szymczakowski
- Forme plus allongée, côtés du pronotum sinués devant les angles postérieurs. 2
De Sinuosité des côtés du pronotum forte. Stries élytrales avec des gros
points enfoncés. Pronotum avec deux fovéoles situées au quart de la
lanseunde:schaque COESIONE itotateoi Hayashi
- Sinuosité des côtés du pronotum plus discrète. Stries élytrales sans gros
points enfoncés. Pronotum sans fovéole. .................. smetanai n. sp.
Anemadus taiwanus n. sp. (figs 24, 28)
HOLOTYPE d: Taiwan, Chial Hsien, Alishan, sister ponds, 2180 m, le 26-IV-1990, A.
Smetana (MHNG). Paratype: 1 ®, Ilan Hsien, Chyr Duan, 1050 m, 18-IV-1990, A. Smetana
(MHNG).
Description: longueur 2,8 mm. Corps large et convexe, recouvert d’une longue
pubescence dorée partiellement dressée. Coloration brun-foncé, les trois premiers
articles antennaires et les épaules éclaircis.
Tête à ponctuation forte et rugueuse. La suture clypéo-frontale bien marquée.
Palpes maxillaires à second article très dilaté, le dernier court et conique.
Pronotum transverse et rétréci à la base, couvert d’une ponctuation rugueuse
forte, non striolé transversalement. Angles postérieurs marqués, obtus.
Elytres avec des stries assez fortement ponctuées sur le tiers antérieur, la
ponctuation s’atténuant régulièrement vers l’arrière. Forme convexe, la largeur
CHOLEVIDAE DU JAPON ET DE TAIWAN 293
21 22 23
24
Fics 20 à 26: édéages, faces ventrales et latérales. 20 et 22: Anemadiola itotateoi Hayashi, 21 et
23: Anemadiola smetanai n. sp., 24: Anemadus taiwanus n. sp., 25 et 26: Nargus taiwanensis n.
sp. L’échelle représente 0,25 mm pour les figures 20 à 24, et 0,5 mm pour les figs 25 et 26.
294 MICHEL PERREAU
s’atténuant très rapidement et régulièrement à partir du tiers antérieur où se trouve la
plus grande largeur. Les angles postérieurs arrondis séparément à l’apex.
Tarses antérieurs modérément dilatés, nettement moins larges que l’apex des
tibias. Dilatation des deux premiers tarsomeres intermédiaires pratiquement imper-
ceptible.
Lobe médian régulièrement arqué de la lame basale jusqu'à l'apex. L'apex avec
une fine échancrure (fig. 24). Les paramères plus longs que le lobe médian, con-
vergents à l'apex, présentant trois soies parallèles subapicales, insérées sur la face
inférieure. Sac interne uniformément tapissé d’écailles fines et courtes, et équipé de
deux phanères latérales courtes.
Femelle sans dilatation des tarses antérieurs, spiculum ventrale (fig. 28) large
et court, arrondi, comme dans tout le genre Anemadus. Coloration uniformément plus
claire.
L'espèce se distingue des autres Anemadus par sa petite taille, l'absence de
dilatation des deux premiers articles mésotarsaux, l'apex de l'édéage échancré, et la
structure simple du sac interne, dépourvu de toute dent sclérifiée (mais pas des habi-
tuelles phanères). Il s'agit sans doute d’une structure très primitive de sac interne chez
les Anemadinae. Cette espèce ne prend pas clairement place dans un des groupes déjà
définis dans le genre Anemadus (GIACHINO & VAILATI 1993).
Anemadus nipponensis n. sp. (fig. 31)
HOLOTYPE ©: Japon: Nara, 27/31-7-1980, I. Löbl. (MHNG)
Cette espèce est proche de Anemadus asperatus, et sa description est donnée
par comparaison.
Description: Longueur 3,6 mm, coloration générale brun clair. Tout le corps
est recouvert d'une fine pubescence dorée couchée.
Tête avec une suture clypéo-frontale bien marquée et peu arquée, presque
rectiligne (fig. 31), et une ponctuation rugueuse entremêlée d'une microréticulation.
Pronotum 1,7 fois plus large que long, et 1,3 fois plus large que les élytres,
avec une ponctuation rugueuse, semblable à celle de À. asperatus et A. kuluensis.
Elytres 1,5 fois plus longs que larges ensemble, la ponctuation des stries forte
ainsi que la striolation transversale, et semblable à celle de A. asperatus. Par contre
les élytres sont plus convexes que A. asperatus.
L'Anemadus nipponensis n. sp. appartient au groupe "asperatus" (GIACHINO &
VAILATI 1993). C'est d'ailleurs l'espèce la plus proche de A. asperatus. Elle s'en
distingue par les élytres plus convexes et par la suture clypéo-frontale presque
rectiligne et non arquée (figs 30 et 31).
L'importance biogéographique de la découverte de cette espèce ainsi que de la
suivante a déjà été discutée dans l’introduction.
Nargus taiwanensis n. sp.
HOLOTYPE d: Taiwan: Pingtung, Hsien, Peitawushan, above Kuai-Ku Hut, 2750 m, 29-
IV-1992, A. Smetana (MHNG). Paratype: 1 femelle même provenance (MHNG).
CHOLEVIDAE DU JAPON ET DE TAIWAN 295
27 28
29)
30 31
|
Fics 27 à 29: femelles, ventrite VII et spiculum ventrale. 27: Anemadiola smetanai n. sp. 28:
Anemadus taiwanus n. sp. 29: Nargus taiwanensis n. sp. Fics 30 et 31 sutures clypéo-frontales.
30: Anemadus asperatus Champion, 31: Anemadus nipponensis n. sp. L’échelle représente 0,3
mm pour les figures 27 et 28, 0,5 mm pour les figures 29 a 31.
Description: Longueur: 3,6 mm. Espèce aptère, de forme générale convexe,
large et courte et de coloration brun clair uniforme.
Tête fortement microréticulée, d’aspect mat, la ponctuation extrêmement fine,
indiscernable au milieu de la microréticulation.
Pronotum ample dépassant la largeur des élytres, à angles postérieurs arrondis.
Les téguments présentent une microréticulation profonde, aussi forte que celle de la
296 MICHEL PERREAU
tête, et donnant aussi un aspect mat à la surface pronotale. Ponctuation fine mais bien
visible au milieu de la réticulation.
Elytres très convexes, moins profondément microréticulés que le pronotum,
l’aspect général brillant, et la ponctuation nettement plus grosse. Les angles apicaux
des deux élytres sont allongés en deux très fortes dents se recouvrant l’une l’autre.
Tibias antérieurs avec deux éperons externes visibles et quelques épines irré-
gulières. Tarses antérieurs fortement dilatés, un peu plus larges que l’apex des tibias.
Soies apicales des tibias inégales.
Edéage non symétrique, les parties dorsales et ventrales du lobe médian
réduites à deux longues lames bifurquées à l’apex. Ces deux lames se croisent a
l’apex, la première se déplaçant vers la gauche, et la seconde vers la droite (figs 25 et
26). Paraméres d’épaisseur constante, aussi longs que le lobe médian, avec deux soies
apicales de longueur inégales, l’une longue, dirigée vers l’apex du lobe médian,
l’autre plus interne et plus courte. Le sac interne est muni de courtes dents isolées, de
phanères, et à la base, d’une très forte dent.
Femelle semblable au mâle avec les élytres également dentés, mais sans
dilatation des tarses. Ventrite VIII et spiculum ventrale représentés sur la figure 29.
REMARQUE: l’absence de rangées de soies égales à l’apex des tibias, et la
conformation de l’édéage suggèrent l’appartenance au sous-genre Demochrus, mais
cette espèce présente des caractères tout à fait particuliers. Sa taille, nettement plus
grande que celle des autres espèces, l’absence de la seconde paire d’aile, et la
conformation complexe de l’édéage l’isolent très nettement au sein du genre. A terme,
un nouveau sous-genre, voire un nouveau genre pourrait être introduit, surtout si
d’autres espèces présentant une semblable évolution de l’organe copulateur venaient à
être découvertes dans cette région.
Cette évolution de l’organe copulateur chez les Cholevini rappelle celle ren-
contrée chez les Catopini du genre endémique japonais Apterocatops Nakane. Une
meilleure compréhension de la signification fonctionnelle de telles modifications
structurales permettra peut-être de préciser l’origine de cette évolution parallèle dans
les deux tribus, et se produisant dans la même région.
Sciodrepoides fumatus (Spence)
Japon: Tochigi, Nikko Nat. Park Senjogahara, 1400 m, 15-7-1980, I. Löbl, 3 ex.
(MHNG).
Espèce répandue dans toute l'Europe et une grande partie de l'Asie.
Mesocatops japonicus (Jeannel)
Japon: Gunma, 4 km sw Tsumagoi, 1050 m, 18-7-1980, I. Löbl, 1 ex. (MHNG).
Catops miensis formosensis Hayashi
Taiwan: Hualien Hsien, Taroko, N. P. Nanhushi Hut, 2220 m,
3; Taichung Hsien, Hsuehshan, above Shan-Liu-Gieu Hut, 3150 m, 8-
(MHNG).
8-V-1990, A. Smetana, 1
V-1991, A. Smetana, 1 d
CHOLEVIDAE DU JAPON ET DE TAIWAN 297
Contrairement à la description de HAYASHI (1988), les paramères portent deux
soies et non une seule, mais les deux soies sont de longueurs inégales, l’une est un
peu plus courte que la moitié de la plus longue.
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of the Faculty of Sciences, Kagoshima University (Earth Sciences and Biology), 15 :
101-111.
NISHIKAWA M., 1986. A new Catopid beetle (Coleoptera, Catopidae) from Japan. Special
Bulletin of the japanese Society of Coleopterology, 2: 107-110.
NISHIKAWA M., 1993. Occurrence of a New Apterous Species of Ptomaphagus (Coleoptera,
Cholevidae) in Taiwan. Elytra, Tokyo, 21 : 123-128.
PERREAU M., 1988. Les Cholevidae himalayens du Muséum d'Histoire naturelle de Genève.
Revue suisse de Zoologie, 95 : 1005-1018.
PERREAU M., 1992. Nouvelles espèces de Ptomaphaginus de Thaïlande et de Taiwan (Cole-
optera, Cholevidae). Bulletin de la Société entomologique de France, 97: 343-347.
PORTEVIN G., 1914. Silphidae. Supplementa Entomologica, 3: 5-8.
SZYMCZAKOWSKI W., 1962. Remarques sur quelques Catopinae du Japon. Niponius, 1: 1-7.
SZYMCZAKOWSKI W., 1963. Contribution à la connaissance des Anemadinae (Coleoptera
Catopidae). Polskie Pismo Entomologiczne, 33: 101-117.
SZYMCZAKOWSKI W., 1965. Zur Systematik und Verbreitung einiger Catopidae (Coleoptera) der
paläarktischen und orientalischen region. Polskie Pismo Entomologiczne, 35: 521-533.
ERRATUM
W.R. LOURENÇO 1995: Chaerilus sabinae, nouvelle espèce de Scorpion anophthalme des grottes
de Matampa en Inde (Scorpiones, Chaerilidae). - Revue suisse de Zoologie 102(3): 847-
850.
Suite à la publication de cette description, notre collègue Louis Deharveng
(Laboratoire d’Ecologie Terrestre/Zoologie, UMR9964 du CNRS, Toulouse) nous a
signalé une grave erreur dans la localité de ce scorpion anophthalme, due à une mau-
vaise transcription de l’étiquette originale par le prédécesseur de l’auteur.
La localité-type de cette espèce est la suivante: Indonésie, île de Sulawesi, pro-
vince de Sulawesi Selatan, Grotte “Gua Atas” (en Indonésien: grotte du haut), dans le
petit massif karstique de Matampa, près de Pangkajene, dans la région de Maros, coor-
données approximatives: 119° 38° E, 4° 48° 44” S; lg. L. Deharveng, 12.VIII.1985 (no.
INDO 76).
BIBLIOGRAPHIE concernant cette grotte et la campagne spéléologique:
BESSON, J.-P. 1986. Catalogue Indonésie. /n: Expédition Thai-Maros 85. Rapport spéléologique et
scientifique: 63-67. Association pyrénéenne de spéléologie, Toulouse.
DEHARVENG, L. & LECLERC, P. 1989. Recherches sur les faunes cavernicoles d’Asie du sud-est.
Mémoires de Biospéléologie 16: 91-110. (p. 95).
LECLERC, P. 1986. Arachnides. In: Expédition Thai-Maros 85. Rapport spéléologique et scienti-
fique: 181-185 (Scorpions: 184-185). Association pyrénéenne de spéléologie, Toulouse.
RIGAL, D. & A. BEDOS 1986. Le Karst de Matampa. /n: Expédition Thai-Maros 85. Rapport spé-
léologique et scientifique: 96-97. Association pyrénéenne de spéléologie, Toulouse.
idqiune
WARIS, SE
REVUE SUISSE DE ZOOLOGIE
Tome 103 — Fascicule 1
DESQUEYROUX-FAUNDEZ, Ruth & Rob W.M. VAN SOEST. A review of
Iophonidae, Myxillidae and Tedaniidae occurring in the South East
Pacific (Porifera: Poecilosclerida)........ PUCES a ae EN A Mi
MonNEy, Jean-Claude, Lucia LUISELLI & Massimo CAPULA. Taille et méla-
nisme chez Vipera aspis dans les Préalpes suisses et en Italie centrale
et comparaison avec différentes populations alpines de Vipera berus.
CONDE, Bruno. Diploures Campodéidés de la Pestera de la Movile (Movile
Cave), Dobroudja méridionale (Roumanie). ....................
ANGELINI, Fernando. A New Species and records of the genus Pseudo-
agathidium Angelini from Uganda (Coleoptera, Leiodidae, Agathi-
CU) a a ee na,
AssınG, Volker & Paul WUNDERLE. A Revision of the Madeiran species of
the genus Geostiba Thomson, 1858 (Coleoptera: Staphylinidae). . ..
GOLOVATCH, Sergei I. The millipede family Paradoxosomatidae on Borneo,
with contributions to the faunas of some other islands of the Sunda area
WBplopodasBolydesmida)ı.z, 2... u... een. See
PACE, Roberto. Aleocharinae della Sottoregione Africana Orientale al Mu-
seo di Ginevra (Coleoptera, Staphylinidae) Parte III (conclusione). . . .
MAHUNKA, Sändor. Oribatids from Sarawak I. (Acari: Oribatida). New and
interesting mites from the Geneva Museum LXXVII. ............
PERREAU, Michel. Contribution à la connaissance des Cholevidae du Japon
ciadeglialwani(Coleoptera)- ol RAR RR na
Pages
3-79
81-100
101-114
115-118
119-150
151-193
195-258
259-282
283-297
REVUE SUISSE DE ZOOLOGIE
Volume 103 — Number 1
DESQUEYROUX-FAUNDEZ, Ruth & Rob W.M. VAN SOEST. A review of
Iophonidae, Myxillidae and Tedaniidae occurring in the South East
Bacıei(BoriteragBoeciloselenda). Re
MONNEY, Jean-Claude, Lucia LUISELLI & Massimo CAPULA. Body size and
melanism in Vipera aspis in the Swiss Prealps and Central Italy and
comparison with different alpine populations of Vipera berus......
ConDE, Bruno. Diplura Campodeidae from the Pestera de la Movile
(Movile Cave), Southern Dobrogea (Rumania).................
ANGELINI, Fernando. A New Species and records of the genus Pseudo-
agathidium Angelini from Uganda (Coleoptera, Leiodidae, Agathi-
CR o BR Re od cc co cc
AssınG, Volker & Paul WUNDERLE. A Revision of the Madeiran species of
the genus Geostiba Thomson, 1858 (Coleoptera: Staphylinidae). . . .
GOLOVATCH, Sergei I. The millipede family Paradoxosomatidae on Borneo,
with contributions to the faunas of some other islands of the Sunda area
@iplopoda;Polydesmida) i... 6. sac ache © SEITEN eee
PACE, Roberto. Aleocharinae from Eastern African Region in the Geneva
Museum (Coleoptera, Staphylinidae). Part IH (conclusion). . ......
MAHUNKA, Sändor. Oribatids from Sarawak I. (Acari: Oribatida). New and
interesting mites from the Geneva Museum LXXVIIL ...........
PERREAU, Michel. Contribution to the knowledge of the Cholevidae of
Japan and Marwan) (Coleoptera): Nr RIE
Indexed in CURRENT CONTENTS
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259
283
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REVUE SUISSE DE ZOOLOGIE, 103 (2): 301-318; juin 1996
Revision westmediterraner Scopaeus - Arten
(Coleoptera, Staphylinidae: Paederinae) und Beschreibung
einer neuen Art aus Südspanien und Marokko
2. Beitrag zur Kenntnis der Gattung Scopaeus Erichson
Johannes FRISCH
Zoologisches Institut der Justus-Liebig-Universität Gießen,
Stephanstr. 24, D-35390 Gießen.
Revision of Western Mediterranean Scopaeus-species with description
of a new species from Southern Spain and Maroc. - Revision of eight
species of the genus Scopaeus Erichson, 1840 (Coleoptera, Staphylinidae:
Paederinae). New characters for identification of females are introduced.
Taxonomic results: Scopaeus anxius Mulsant & Rey, 1861 (Scopaeus
pourtoyi Coiffait, 1960; S. balazuci Coiffait, 1968; S. revestensis Coiffait,
1968: syn. n.) is revalided and redescribed. S. kerdousensis Coiffait, 1973
syn. n. = S. fageli Coiffait, 1960. S. kovaci sp. n. (Southern Spain, Northern
Maroc) is compared with S. longicollis Fauvel, 1874 and S. proculus
Normand, 1934. Lectotypes are designated for S. anxius and S. longicollis.
Key-words: Coleoptera - Staphylinidae - Paederinae - Scopaeus - new
species - new synonymies.
EINLEITUNG
Die Gattung Scopaeus Erichson, 1840 ist in der westlichen Paläarktis (sensu
CoIFFAIT 1984) gegenwärtig mit 88 validen Arten und Unterarten sowie 119
verfügbaren Namen vertreten. COIFFAIT (1984) führt 77 Arten an; FRISCH (1994) und
GUSAROV (1991, 1992, 1994) erhöhten die Zahl auf jetzt 88 Spezies.
Diese Arbeit ist der erste Teil einer Revision westpaläarktischer Scopaeus-
Arten, deren Notwendigkeit besonders durch das Studium der Coiffait'schen Typen
deutlich wurde; ein Großteil der Arten Coiffait's ist in die Synonymie zu stellen.
Coiffait beschrieb seine Arten ohne Typenstudium; auch bewertete er individuelle
oder geringfügige geographische Variationen viel zu hoch.
Die Gliederung der Gattung in Subgenera (CoIFFAIT 1960, 1968, 1981, 1984)
wird nicht übernommen, da sie nach den Kriterien der phylogenetischen Systematik
(HENNIG 1982) nicht haltbar ist. Eine phylogenetische Analyse der Artgruppen ist
gegenwärtig nicht möglich.
: Manuscript accepted 05.07.1995.
302 JOHANNES FRISCH
Klassifikation und Determination der © © fast aller Scopaeus - Arten bereiten
noch immer große Schwierigkeiten. In dieser Arbeit wird versucht, die 2 2 anhand
der Terminalsklerite und der artspezifisch gestalteten, sklerotisierten Spermatheka zu
charakterisieren. Die Skleritteile der Terminalsegmente 9 und 10 sind nicht immer
von hohem diagnostischem Wert, aber dennoch geeignet, die Determination der 2 £
zu erleichtern; ihre Terminologie richtet sich nach UHLIG (1989). Als Mittelfeld des
Tergits 10 und der lateralen Sternalteile 9 (Valve) wird das beborstete, innerhalb der
breiten Skleritränder gelegene Zentralfeld bezeichnet. Auf den Zeichnungen der
weiblichen Genitalsklerite ist die gesamte Beborstung dargestellt; ausgebrochene
Borsten sind an den Insertionspunkten erkennbar. Die feine Grundbeborstung der
Abdominalsternite 8 der 9 d ist nicht dargestellt.
Die Spermatheka der Scopaeus - Arten scheint in vielen Artgruppen eine
Klassifikation der 9 2 zu ermöglichen. Sie ist bei allen untersuchten Arten nach
einem Grundschema (vgl. Abb. 11, 12) gebaut. Die Kapsel setzt sich aus zwei distad
erweiterten, astförmigen Gliedern zusammen, die scheinbar gelenkartig verbunden
und in Seitenansicht zueinander gewinkelt sind; ihre Enden sind gegeneinander
gebogen. Ein sklerotisierter Ductus mündet in eines der Kapselglieder; ein
membranöser Ductus scheint der Gelenkstelle beider Kapselglieder zu entspringen.
Die meist unter 0,1 mm messenden Spermatheken sollten im Durchlichtmikroskop bei
einer Vergrößerung von mindestens 400x untersucht werden.
MABE UND ABKÜRZUNGEN: Die Längenangaben in den Diagnosen sind wie
folgt gemessen: Vorderkörperlänge vom Vorderrand der Mandibeln bis zum Hinter-
rand der Elytren. Kopflänge vom Vorderrand des Clypeus bis zum Hinterrand des
Kopfes. Augen- und Schläfenlänge sind von der Seite gemessen. Bei den 2 9 -Termi-
nalskleriten beziehen sich Maße oder Indizes auf das Mittelfeld innerhalb der breiten
Skleritränder. Proportionsmaße in Millimetern. NL = Nahtlänge, SL = Schulterlänge,
B = Breite der Elytren, PL = Länge des Pronotums.
REVISION DER ARTEN
Scopaeus anxius Mulsant & Rey, 1861 (Abb. 1-12)
Scopaeus (Polyodontus) anxius Mulsant & Rey, 1861. — BERNHAUER & SCHUBERT 1910. —
GANGLBAUER 1895. — GEMMINGER & HAROLD 1868. — MULSANT & REY 1861b, 1877.
Scopaeus (s. str.) pourtoyi Coiffait, 1960 syn. n. — COIFFAIT 1968, 1984.
Scopaeus (s. str.) balazuci Coiffait, 1968 syn. n. — COIFFAIT 1984.
Scopaeus (s. str.) revestensis Coiffait, 1968 syn. n. — COIFFAIT 1984.
UNTERSUCHTES TYPENMATERIAL: Zwei Syntypen aus der Sammlung Rey (MGL), die an
dieser Stelle als Lectotypus (4) und Paralectotypus (2) designiert werden. Beide Exemplare, die
auf sehr dünnen Drahtnadeln montiert waren, wurden genitalpräpariert und auf Kartonplättchen
geklebt. Rey'sche Stücke tragen neben einem Genuszeichen meist noch ein verschlüsseltes
Fundortetikett in Form eines kleinen farbigen Kartonplättchens. Die Syntypen tragen ein
schwarzes Etikett: 'Provence' und stammen laut Diagnose aus der Umgebung von Hyères.
Scopaeus pourtoyi syn. n.: Holotypus d (MHNP): Frankreich, Pyrenäen-Ostrand, env.
Banyuls, 05.06.1954, leg. Pourtoy.
WESTMEDITERRANE SCOPAEUS - ARTEN 303
Scopaeus balazuci syn. n.: Holotypus d (MHNP): Frankreich, Ardèche, Frigolet env.
Saint-Paul-Le Jeune, 25.08.1959, leg. Balazuc.
Scopaeus revestensis syn. n.: Holotypus d, Paratypus 2 (MHNP): Frankreich, Alpes
de Provence, Revest des Brousses, 08.1962.
WEITERES MATERIAL: 2 d d (Coll. Frisch): Spanien, Gueja Sierra, Ufer des Genil, 1200
mNN, leg. P. Wunderle.
Länge: 3,1-3,4 mm; Vorderkörper 1,9 mm. Die folgenden Proportionsmaße
beziehen sich auf den Lectotypus.
Ausgefärbt pechschwarz, doch scheinen häufig heller gefärbte (immature?)
Exemplare aufzutreten. Diese sind nur auf den vier vorderen Abdominaltergiten ge-
schwärzt. Vorderkörper einheitlich pechschwarz, Hinterrand der Elytren sehr schmal
aufgehellt. Labrum braun, Antennenwurzel aufgehellt. Mandibeln, Maxillarpalpen,
Antennen und Beine hellbraun. Femora stärker gebräunt, vorletztes Glied der
Maxillarpalpen geschwärzt. Abdomen pechschwarz, Abdominaltergit 7 nach hinten
schwach aufgehellt, Abdominaltergit 8 deutlich heller, zum Hinterrand gelbbraun.
Punktierung überall deutlich. Punktabstände am Kopf besonders auf den
Schläfen kleiner als die Punktdurchmesser, auf Frons und Clypeus diesen weitgehend
entsprechend. Punktierung des Pronotums obsolet und deutlich feiner als die des
Kopfes, die Punktabstände entsprechen dem Durchmesser. Punktfreie Mittellinie des
Pronotums schmal, im vorderen Drittel etwa von der Breite der Vorderschienenbasis.
Elytren grob und doppelt so stark wie der Kopf punktiert, Punktabstände kleiner als
die Durchmesser. Punktur zum Hinterrand der Elytren zunehmend obsolet, aber nicht
feiner oder weitläufiger. Abdomen sehr grob, auf den Abdominaltergiten 3-6 etwas
stärker und dichter als auf den Elytren punktiert, Punktur erst auf Tergit 7 feiner und
etwas weitläufiger.
Mikroskulptur obsolet, Gesamteindruck glänzend. Chagrinierung des Kopfes
nur auf den Schläfen deutlicher, auf Frons, Clypeus und der Mittellinie geschwunden.
Pronotum und Elytren glänzend, Chagrinierung kaum erkennbar. Abdominaltergite
3-6 deutlich chagriniert, doch ebenfalls glänzend. Mikroskulptur der Abdominal-
tergite 7 und 8 zunehmend obsolet.
Kopf etwas länger als breit (0,51 : 0,46, Index 1,11). Größte Kopfbreite über
den hinter den Augen schwach erweiterten Schläfen, diese gut doppelt so lang wie die
Augen (0,14 : 0,29, Index 0,48), von ihrer Mitte unter deutlichem Winkel zu den breit
verrundeten Hinterecken verengt. Hinterrand schwach konkav. Seitenzähne des
Labrums halb so lang wie mittlere Zähne.
Pronotum länger als breit (0,52 : 0,41, Index 1,24), unwesentlich länger (Index
0,98) und schmaler (Index 1,12) als der Kopf. Punktfreie Mittellinie im vorderen
Drittel stark, im hinteren Viertel schwächer gefurcht.
Elytren ein Viertel länger als breit (NL 0,46, SL 0,57, B 0,46, SL : B 1,24),
Naht kürzer (NL : PL 1,13), Schultern etwas länger (SL : PL 0,91) als das Pronotum.
Holopter. Abdominaltergit 7 mit Palisadensaum.
Antennen 0,9 mm lang. Antennomere 1-5 länger als breit, 6 und 7 so lang wie
breit, 8-10 geringfügig breiter als lang. Antennomere 1-11 (L : B): 2,7; 1,4; 1,5; 1,2;
PAO 0028-108078 ve
304
JOHANNES FRISCH
WESTMEDITERRANE SCOPAEUS - ARTEN 305
Beine ohne besondere Merkmale. Schienen schlank, ohne Erweiterungen. Vor-
dertarsen beider Geschlechter stark erweitert.
MANNCHEN: Sternit 8 (Abb. 7) caudal auf ein Viertel der Länge dreieckig
eingeschnitten, Spitzenwinkel des Ausschnittes schwach verrundet.
Aedoeagus (Abb. 1-6): 0,65-0,7 mm, Phallobasis mit ausgeprägten, ventral
mit kurzen Borsten besetzten Laterallobi. Apikallobi relativ kurz und gedrungen,
lateral gerundet erweitert und hier mit Sinnesgruben, die zum Teil äußerst kurze
Börstchen tragen, besetzt. Ventralrand der Apikallobi bogenförmig gerundet und
proximad gegen die Phallobasis vorgezogen. Länge der Apikallobi variabel. Dorsallo-
bus apikal eingeschnitten und in zwei ventrad gebogene Dorne ausgezogen. Tiefe
dieses Ausschnitts wie auch die Form der Dorne variabel, die breiter und kürzer, aber
auch schlanker und länger gestaltet sein können. Ein langer, unpaarer Dorn des
Endophallus, der zwischen den Dornen des Dorsallobus verläuft, erreicht fast den
Apex der Apikallobi. Ein kurzer, in Ventralansicht apıkal ausgerandeter Ventraldorn
ragt zwischen die Laterallobi.
WEIBCHEN: Laterale Tergalteile 9 (Abb. 8) in eine lange, schwach dorsad
gebogene Spitze ausgezogen, ohne dorsale Nebenspitze. Tergit 10 (Abb. 10) parallel,
etwa doppelt so lang wie breit (Index 2,11). Valven (Abb. 9) schlank, über sieben mal
so lang wie breit (Index 7,31), stark beborstet, die drei Hauptborsten kräftig.
Spermatheka (Abb. 11, 12): 0,1 mm, Kapselglieder distad stark beilförmig
erweitert. Das Glied, in das der sklerotisierte Ductus mündet, ist besonders breit. In
Seitenansicht ist das den Ducti entfernte Glied stark verlängert und gebogen.
Verbreitung: Scopaeus anxius, bislang nur aus Südfrankreich bekannt, ist im
Rhonetal nördlich bis in die Region Ardeche belegt und liegt nun auch aus Süd-
spanien (Andalusien) vor. Die Art ist vermutlich über ganz Spanien verbreitet.
VERWANDTSCHAFTSBEZIEHUNGEN: Scopaeus anxius ist mit keiner bekannten Art
in direkte Beziehung zu bringen, doch scheint auf Grund der Ausbildung der
Apikallobi und der Apikaldorne des Aedoeagus eine engere Verwandtschaft mit
Scopaeus gladifer Binaghi, 1935 zu bestehen.
Scopaeus anxius scheint farbvariabel zu sein. Die Rey'schen Typen sind fast
einheitlich hellbraun gefärbt; auch der Holotypus des Scopaeus balazuci syn. n. ist
relativ hell. Ausgefärbte Stücke sind aber von tief schwarzer Farbe. Vermutlich sind
die hellen Exemplare immatur wie auch der Typus des Scopaeus revestensis syn. n.,
was zu der irreführenden Zeichnung des verformten Aedoeagus mit stark gespreizten
Apikallobi (COIFFAIT 1968) führte. Die Gestalt der Apikallobi, der Apikaldorne und
die Tiefe des apikalen Einschnitts des Dorsallobus sind ebenfalls variabel, was aber
keine spezifische Gliederung rechtfertigt.
ABB. 1-6
Scopaeus anxius: Aedoeagus, lateral: 1) S. pourtoyi syn. n., Holotypus, 2) Lectotypus, 3)
Spanien, Gueja Sierra, dorsal: 4) Spanien, Gueja Sierra, 5) S. pourtoyi syn. n., Holotypus,
ventral: 6) S. pourtoyi syn. n., Holotypus.
306 JOHANNES FRISCH
ABB. 7-12
Scopaeus anxius, 3 -Lectotypus: 7) Sternit 8. 9 -Paralectotypus: 8) laterales Tergalteil 9, 9)
Valve, 10) Tergit 10, 11) Spermatheka, Aufsicht, 12) Spermatheka, Seitenansicht, Maßstab-
einheit 0,1 mm.
Scopaeus kovaci sp. n. (Abb. 13-15, 20, 22, 24, 26, 29)
TYPENMATERIAL: Holotypus d (MHNG): Spanien, Caseres, Terte, 24.08.1969, leg. A.
Senglet. 29 Paratypen: 1 d, 1 2 (MHNG, Coll. Frisch): Spanien, Cordoba, Villaviciosa,
29.06.1969, leg. A. Senglet. 2 22 (MHNG): Spanien, Cordoba, Palma del Rio, 26.06.1969,
leg. A. Senglet. 3 88,5 22 (MHNG, Coll. Frisch): Spanien, Huelva, Zalamea, 06.07.1969,
leg. A. Senglet. 1 d (MHNG): Spanien, Badajoz, Monesterio, 19.06.1969, leg. A. Senglet. 1 d,
1 2 (NMW): Spanien, Algesiras. 6 & 4,8 2 2 (DEI, HUB): Marokko, Tanger, leg. Rolph.
Länge: 2,86-3,2 mm; Vorderkörper 1,57-1,79 mm. Die Proportionsmaße
beziehen sich auf den Holotypus.
Vorderkörper matur dunkelbraun, Exemplare aus Marokko durchweg etwas
heller. Kopf und Pronotum einheitlich dunkelbraun. Elytren auf den Schultern ge-
schwärzt, zum Hinterrand heller und von der Farbe des Pronotums. Labrum braun,
Antennenbasis aufgehellt. Mandibeln, Maxillarpalpen, Antennen und Beine hell gelb-
braun. Abdomen schwarz, Tergit 7 aufgehellt, Tergit 8 hell braungelb.
Körper überall kräftig punktiert. Punktur des Kopfes stark und gedrängt,
Punktabstände kleiner als Durchmesser, auf Frons und Clypeus weitläufiger, die
Punktabstände entsprechen dort dem Durchmesser. Pronotum etwas obsolet und
WESTMEDITERRANE SCOPAEUS - ARTEN 307
wesentlich feiner als der Kopf punktiert, die Punktabstände entsprechen dem Durch-
messer. Pronotum ohne punktfreie Mittellinie. Elytren fast doppelt so stark, doch
nicht so gedrängt wie der Kopf punktiert, Punktur zum Hinterrand etwas feiner und
weitläufiger. Abdominaltergite 3-6 so grob wie die Elytren, doch stark gedrängt und
etwas tiefer eingestochen punktiert. Tergite 7 und 8 zunehmend feiner, weitläufiger
und nicht gedrängt punktiert.
Mikroskulptur obsolet. Glänzender Gesamteindruck, der nur durch die dichte
Punktierung besonders des Kopfes geschwächt ist. Mikroskulptur des Kopfes
erkennbar, doch in der dichten Punktur undeutlich, Frons und Clypeus völlig
glänzend. Pronotum, Elytren und Abdomen stark glänzend, Chagrinierung obsolet,
doch erkennbar.
Kopf etwas länger als breit (0,46 : 0,41, Index 1,12). Schläfen nicht erweitert,
knapp 1,4 mal so lang wie die Augen (0,16 : 0,22, Index 0,73) und bereits vom ersten
Drittel an schwach zu den sehr breit gerundeten Hinterecken verengt. Hinterrand
gerade, Seitenzähne des Labrums halb so lang wie Mittelzähne.
Pronotum schlank, mittleres Drittel parallel. Länger als breit (0,48 : 0,38 mm,
Index 1,26), etwas länger (Index 0,96) und schmaler (Index 1,08) als der Kopf.
Punktfreie Mittellinie im vorderen Drittel deutlicher, im hinteren Viertel schwach
gefurcht.
Elytren ein Fünftel länger als breit (NL 0,47, SL 0,61, B 0,51, SL: B 1,2) und
länger als das Pronotum. Die Nahtlänge entspricht der. Länge des Pronotums
weitgehend (PL : NL 1,02), Schulterlänge um ein Viertel länger als das Pronotum (PL
:SL 0,79). Holopter. Abdominaltergit 7 mit Palisadensaum.
Antennen 0,82 mm lang. Antennomere 1-4 länger als breit, 5-7 sehr schwach
quer, 8-10 knapp ein Drittel breiter als lang. Antennomere 1-11 (Le B)a2:5 05:13;
IS 40808410 *8=07107 05/5 1:5.
Beine schlank, Mittelschienen zur Spitze erkennbar erweitert (L : B = 5,56).
Vordertarsen beider Geschlechter stark verbreitert.
MANNCHEN: Abdominalsternit 8 (vgl. Abb. 27) im caudalen Viertel mit brei-
tem dreieckigem Ausschnitt, dessen Spitze völlig verrundet ist. Mittellinie des
Sternits durch intensivere Färbung und verdichtete, feine Grundbeborstung hervor-
gehoben.
Aedoeagus (Abb. 13-15): 0,6 mm, Laterallobi sehr umfangreich, die Basis der
Apikallobi weit überragend, ventral mit zahlreichen, feinen Borsten besetzt.
Apikallobi in Dorsalansicht parallel, ventral stark proximad verjüngt und mit
vereinzelten Sinnesgruben. Dorsallobus apikal breit ausgerandet und in zwei apikad
orientierte, etwas aufeinander zugebogene, dünne Dorne ausgezogen. Der Apex des
Dorsallobus trägt zwei weitere, ventrad gerichtete, kurze Dornen. Ein dünner, langer
Dorn des Endophallus verläuft apikad und endet zwischen diesen Dornen. Phallobasis
zwischen den Laterallobi mit breitem, in Ventralansicht ausgerandetem Fortsatz.
WEIBCHEN: Laterale Tergalteile 9 (Abb. 20) apikal spitz ausgezogen, dorsale
Nebenspitze gut entwickelt. Tergit 10 (Abb. 22) breit (Index 1,79), Seiten gleich-
mäßig schwach gerundet. Valve (Abb. 24) relativ breit gebaut (Index 5,36).
308 JOHANNES FRISCH
ABB. 13-18
Scopaeus kovaci sp. n., Holotypus: Aedoeagus 13) dorsal, 14) lateral, 15) ventral. S. longicollis,
Portugal, Vila Real: Aedoeagus 16) dorsal, 17) lateral, 18) ventral.
WESTMEDITERRANE SCOPAEUS - ARTEN 309
Spermatheka (Abb. 26): 0,07 mm, mit annähernd rechtwinklig orientierten,
apikal nur schwach gebogenen Kapselgliedern. Sklerotisierter Ductus robust, lang
ausgezogen, am Ende verdickt und in einen membranösen Ductus verlängert.
DERIVATIO NOMINIS: Dr Damir Kovac (FSF) sei mit der Wahl des Namens für
die Unterstützung dieser Arbeit herzlich gedankt.
VERBREITUNG: Scopaeus kovaci Sp. n. ist aus dem südlichen Spanien, nördlich
bis Badajoz, und von der Nordspitze Marokkos (Tanger) bekannt. Scopaeus kovaci
sp. n. wird vermutlich nach Norden durch Scopaeus longicollis und in Nordafrika
durch Scopaeus proculus ersetzt.
VERWANDTSCHAFTSBEZIEHUNGEN: Scopaeus kovacı sp. n. ist als vermutlich
vikariierendes Adelphotaxon des Scopaeus longicollis aufzufassen. Folgende Synapo-
morphien werden angenommen:
— Laterallobi des Aedoeagus flügelförmig vergrößert und ventral mit feinen
Borsten besetzt;
— Apikallobi apikad erweitert, proximad verjiingt, ventral abgeflacht;
— Dorsallobus apikad nicht verjüngt, breit abgestutzt und in zwei apikad
gerichtete Dornen ausgezogen, zusätzlich mit zwei weiteren, kurzen, ventrad
orientierten Dornen.
Eine engere Verwandtschaft ist auch zu dem nordafrikanischen Scopaeus pro-
culus erkennbar, der sich ebenfalls durch auffallend stark entwickelte, fein beborstete
Laterallobi des Aedoeagus auszeichnet, in anderen Merkmalen aber deutlich ab-
weicht.
DIFFERENTIALDIAGNOSE: Scopaeus kovaci Sp. n. soll mit Scopaeus longicollis
und Scopaeus proculus verglichen werden.
Scopaeus longicollis Fauvel, 1874 (Abb. 16-18, 19, 21, 23, 25, 28)
BINAGHI 1935. — COIFFAIT 1952, 1984. — GANGLBAUER 1895. — PORTA 1926. — PORTEVIN 1929. —
REITTER 1909.
UNTERSUCHTES TYPENMATERIAL: Zwei 9 9 -Syntypen, die der Beschreibung Fauvel's
zugrundeliegen. Der Lectotypus wurde umpräpariert und genitalisiert. Lectotypus 9 (ISNB):
Frankreich, Limoges. Paralectotypus © (ISNB): Frankreich, Carcassonne.
WEITERES MATERIAL: 2 dd (Coll. Frisch): Portugal, Vila Real, Cortico/Montalegre,
30.08.1969, leg. A. Senglet. 3 Expl. (MHNG): Spanien, Segovia, La Granja, 29.08.1971, leg. A.
Senglet. 3 d 4,4 2 2 (MHNG): Schweiz, Vaud, La Rippe, 31.03.1966, 04.04.1969, 04.04.1976,
leg. A. Comellini. 1 d, 1 2 (MHNG): Schweiz, Vaud, Bonmont, Chéserex, 22.07.1991,
21.06.1989, leg. Cl. Besuchet. 3 2 2 (MHNG): Schweiz, Valais, Erde-Aven, 03.07.1988, 830
mNN, leg. Cl. Besuchet. 1 d (MHNG): Frankreich, Provence, Hyères, 12.1959, leg. Ochs. 1 d
(MHNG): Frankreich, Provence, Foret Don, 06.08.1954, leg. Ochs. 11 44,3 22 (MHNG):
Frankreich, Alpes Maritimes, 1951-1958, leg. Ochs. 1 9 (Coll. Renner, Bielefeld): Deutschland,
Traisen/Nahe, 1989, leg. Renner.
Scopaeus longicollis ist von Scopaeus kovaci sp. n. durch etwas schlankeren
Körperbau, besonders den schlanken, fast parallelen Kopf und durch Unterschiede im
Bau des Aedoeagus und der Spermatheka deutlich unterschieden.
310 JOHANNES FRISCH
ABB. 19-27
Scopaeus kovaci sp. n., 9 -Paratypus, Spanien, Cordoba: 20) laterales Tergalteil 9, 22) Tergit 10,
24) Valve, 26) Spermatheka. S. longicollis, ?-Lectotypus: 19) laterales Tergalteil 9, 21) Tergit
10, 23) Valve, 25) Spermatheka. S. proculus, 8 -"Cotype": 27) Sternit 8, Maßstabeinheit 0,1 mm.
Länge: 3,2 mm; Vorderkörper 1,75 mm. Den Meßwerten liegt ein d aus
Portugal zugrunde.
Durchschnittlich etwas dunkler gefärbt. Elytren weniger stark aufgehellt,
Antennen häufig etwas gebräunt.
Punktierung und Mikroskulptur ohne Differenzialmerkmale.
Kopf etwas schlanker (0,48 : 0,41, Index 1,17). Schläfen schwach erweitert,
über die Hälfte ihrer Länge gerade, erst hinter ihrer Mitte zu den schwächer
gerundeten Hinterecken verengt. Kopf mit schmaler punktfreier Mittellinie.
Pronotum etwas schlanker (0,48 : 0,37, Index 1,3), hinter den Vorderecken
stärker gerundet. Kopf und Pronotum von gleicher Länge.
Elytren mit den gleichen Längen und Proportionen, auch im Verhältnis zum
Pronotum.
WESTMEDITERRANE SCOPAEUS - ARTEN 311
Abdomen ohne Differenzialmerkmale, Tergit 7 mit Palisadensaum.
Antennen 0,89 mm, mit schlankeren Antennomeren. Antennomere 2-4 etwas
länger als breit, 4-10 quadratisch. Antennomere 1-11 (L : B): 2,5; 1,2; 1,2; 1,2; 1,0;
OM POMLOMLO AS).
Beine schlanker, Mittelschienen schwächer erweitert (L : B = 5,58).
MANNCHEN: Abdominalsternit 7 (vgl. Abb. 27) wie bei Scopaeus kovaci sp. n.
gestaltet.
Aedoeagus (Abb. 16-18): 0,6 mm, von gleicher Organisation, doch breiter
gebaut. Laterallobi deutlicher flügelartig vergrößert, die Apikallobi in Seitenansicht
weitgehend verdeckend. Seiten der hyalin erscheinenden, schwach sklerotisierten
Apikallobi stark erweitert, nicht parallel, ihr dorsaler Innenrand stark gewinkelt.
Dorsallamelle apikal mit zwei kurzen, gegeneinander gebogenen Dornen und einem
Medianfortsatz, der einen apikalen, häutigen Lobus stützt und Scopaeus kovaci sp. n.
fehlt. Der apikad gerichtete Dorn des Endophallus erscheint in Ventralansicht sehr
schmal und hakenförmig gebogen. Ventraler Fortsatz zwischen den Laterallobi stark
ausgerandet, breiter als lang.
WEIBCHEN: Laterale Tergalteile 9 (Abb. 19) in einen breiteren Apex ausge-
zogen, dorsale Nebenspitze kleiner. Tergit 10 (Abb. 21) etwas breiter (Index 1,73).
Spermatheka (Abb. 25) von gleicher Größe, durch abgesetzte, parallel
stehende Endteile beider Kapselglieder unterschieden.
VERBREITUNG: Literaturangaben zufolge ist Scopaeus longicollis über das
südliche Mitteleuropa, Südwesteuropa und Nordwestafrika verbreitet. BERNHAUER &
SCHUBERT (1910) führen Südfrankreich, Spanien, Deutschland, Italien, Marokko,
Algerien und Tunesien an. Die Meldung aus Südkärnten durch Scheerpeltz (in HORION
1965) ist unbelegt und abzulehnen. HORION (1965) bezweifelt die unbelegte Angabe aus
Deutschland (EPPELSHEIM 1891), die auch REITTER (1909) übernahm. Ein sicherer
Nachweis aus Deutschland liegt von der Nahe (Traisen, 1989) vor. BINAGHI (1935)
meldet Scopaeus longicollis aus Algerien, Tunesien und Marokko, führt aber auch
Stücke aus Tanger (leg. Rolph, Coll. Peyerimhoff) an, die sicher der gleichen Serie
entstammen wie Paratypen des Scopaeus kovaci sp. n., die sich in den Sammlungen des
NMW, DEI und der HUB befinden. Alte Angaben aus Nordafrika werden schon von
COIFFAIT (1984) bezweifelt; sie sind auf Scopaeus kovaci sp. n. oder auf Scopaeus
proculus zu beziehen; historische Stücke des Scopaeus kovaci sp. n. waren vielfach als
Scopaeus longicollis determiniert. Scopaeus longicollis ist nur aus Mittel- und
Südfrankreich, Deutschland westlich des Rheins, dem westlichen Norditalien, der
Schweiz, Spanien ohne den Süden und dem nördlichen Portugal bekannt und wird nach
Süden vermutlich durch den vikariierenden Scopaeus kovaci sp. n. abgelöst.
Scopaeus proculus Normand, 1934 (Abb. 27, 30, 31-37)
COIFFAIT 1968, 1984.
Scopaeus (Polyodontus) cordifer Binaghi, 1935. — COIFFAIT 1984.
UNTERSUCHTES TYPENMATERIAL: Die Art wurde nach einem d beschrieben. Ein als
Cotype etikettiertes, vom Locus typicus stammendes d lag zur Untersuchung vor; ob es sich
312 JOHANNES FRISCH
um den Holotypus handelt, ist nicht zu klären, da die Sammlung Normand (Tunis) gegenwärtig
nicht zugänglich ist. "Cotype" ¢ (MHNG): Tunesien, Ain-Draham, leg. Normand.
Scopaeus cordifer: Nach vier Stücken beschrieben; drei Typen lagen vor. Zwei ver-
mutlich fälschlich als Cotypen etikettierte 2 2 befinden sich in der Sammlung Scheerpeltz
(NMW). Holotypus dé (MCG): Algerien, Mouzaia, 09.04.1905, leg. Peyerimhoff. Paratypen 2
dd (MCG): Tunesien, Ain-Draham, leg. Normand. 2 2 2 (NMW): Tunesien, Ain-Draham,
leg. Normand.
WEITERES MATERIAL: 1 ¢ (MHNG): Tunesien, Ain-Draham, 05.1937, leg. Normand.
Scopaeus proculus ist von Scopaeus kovaci sp. n. durch hellere Färbung,
bedeutendere Größe, breiteren Kopf, auffallend kurze, stark punktierte Elytren und
wesentlich breitere Mittelschienen zu trennen. Der schlankere Aedoeagus ist be-
sonders durch die Gestalt des Dorsallobus verschieden.
Länge: 3,36-3,4 mm, Vorderkörper 1,86-1,89 mm. Weitere Maße beziehen
sich auf die Normand'sche "Cotype".
Heller gefärbt. Vorderkörper rotbraun, Hinterhälfte des Pronotums und Elytren
geschwärzt, Schultern schwarz, Hinterrand der Elytren schmal bräunlich aufgehellt.
Abdomen schwarz, Tergit 7 auf der Hinterhälfte gelbbraun aufgehellt, Tergit 8 ein-
heitlich gelbbraun. Mandibeln, Maxillarpalpen, Antennen und Beine hell gelbbraun.
Kopf deutlich, aber fein punktiert, die Punktabstände entsprechen dem Durch-
messer. Punktur im Chagrin etwas aufgelöst. Frons und Clypeus weitläufiger punk-
tiert. Pronotum etwas feiner, aber nicht weitläufiger als der Kopf punktiert. Elytren
wesentlich stärker als Kopf und Pronotum punktiert, Punktur tief eingestochen und
gedrängt, zum Hinterrand etwas feiner. Abdominaltergite 3-6 halb so stark wie die
Elytren, doch ebenso gedrängt punktiert. Punktur der Tergite 7 und 8 feiner, im
deutlichen Chagrin schwer erkennbar.
Chagrin des Kopfes deutlich, auf Frons, Clypeus und der Kopfmitte geschwun-
den, ruft zusammen mit der feinen Punktierung einen matten Eindruck hervor.
Pronotum und Elytren obsolet chagriniert, stark glänzend. Abdomen zwischen der
dichten Punktur deutlich mikroskulptiert.
Kopf etwas länger als breit (0,56 : 0,51, Index 1,1). Schläfen stark erweitert,
mehr als doppelt so lang wie die kleinen, schwach gewölbten Augen (0,14 : 0,31,
Index 0,45), ab der Hälfte ihrer Länge ohne Winkelbildung zu den breit verrundeten
Hinterecken verengt. Seitenzähne des Labrums halb so lang wie Mittelzähne.
Pronotum mit den gleichen Proportionen, weniger parallel, Seitenränder
gleichmäßiger gerundet, im Verhältnis zum Kopf etwas schmaler (Index 1,11). Mittel-
linie in ganzer Länge, besonders im vorderen Viertel, tief gefurcht.
Elytren im Verhältnis zum Pronotum wesentlich kürzer, kaum länger als breit
(NL 0,46, SL 0,59, B 0,52, SL : B 0,13) und etwa so lang wie das Pronotum (PL : SL
0,97). Nahtlänge wesentlich kürzer als das Pronotum (Index 1,21). Holopter.
Abdomen ohne Differenzialmerkmale, Abdominaltergit 7 mit Palisadensaum.
Antennen 1,0 mm lang. Antennomere 2-5 ein Fünftel länger als breit, 6 und 7
so lang wie breit, 8-10 quer. Glieder 7-10 rundlich, perlschnurartig. Antennomere
[ERI 3B))2226:71:271623162:3102217,0:51.030:8720:820.821E82
Beine kräftiger, besonders die Mittelschienen stärker erweitert (L : B = 4,29).
Vordertarsen beider Geschlechter stark erweitert.
WESTMEDITERRANE SCOPAEUS - ARTEN 313
Abb. 28-30
Habitus von 28) Scopaeus longicollis, & , Portugal, Vila Real, 29) S. kovaci sp. n., 4 -Paratypus,
30) S. proculus, 8-"Cotype", Maßstabeinheit 1 mm.
MÄNNCHEN: Abdominalsternit 8 (Abb. 27) wie bei Scopaeus kovaci sp. n. und
Scopaeus longicollis gebildet.
Aedoeagus (Abb. 31-33): größer (0,7 mm) und schmaler, Apikallobi länger
ausgezogen, Laterallobi aber ebenso stark entwickelt. Dorsallobus apikal verlängert
und spitz zulaufend, nicht abgestutzt und keine weiteren Dorne tragend. Phallobasis
ventral mit zwei kurzen Borsten.
WEIBCHEN: Laterale Tergalteile (Abb. 34) breiter zugespitzt, dorsale Neben-
spitze kürzer. Tergit 10 (Abb. 36) etwas schmaler (Index 1,93).
Spermatheka (Abb. 37) etwas größer (0,08 mm), Kapselglieder distad stark
erweitert, sklerotisierter Ductus länger.
VERBREITUNG: Scopaeus proculus ist im westlichen Nordafrika beheimatet und
bislang nur aus Algerien und Tunesien bekannt.
314 JOHANNES FRISCH
ABB. 31-33
Scopaeus proculus, "Cotype": Aedoeagus 31) dorsal, 32) lateral, 33) ventral.
Scopaeus fageli Coiffait, 1960 (Abb. 38-45)
Scopaeus (s. str.) fageli Coiffait, 1960. — COIFFAIT 1984.
Scopaeus (Hyposcopaeus) kerdousensis Coiffait, 1973, syn. n. — COIFFAIT 1984.
UNTERSUCHTES TYPENMATERIAL: Holotypus d (ISNB), Paratypen 1 d,1 2 (MHNP):
Algerien, Gorges d'El Kantara, Djebel Metlilli, 05.1954, leg. G. Fagel.
Scopaeus kerdousensis syn. n.: Holotypus d (MHNP): Marokko, Anti Atlas, Col de
Kerdous, 11.1971, leg. H. Coiffait.
Länge: 2,86-3,2 mm; Vorderkörper 1,75—1,9 mm. Weitere Maße beziehen sich
auf den d -Paratypus.
Vorderkörper rotbraun, Hinterhälfte des Pronotums schwach geschwärzt, Ely-
tren auf den Schultern stark geschwärzt, Seitenrand und hinteres Drittel deutlich
aufgehellt. Abdomen dunkelbraun, hinteres Drittel von Tergit 7 und Tergit 8 gelb-
braun aufgehellt. Mandibeln, Maxillarpalpen, Antennen und Beine hell gelbbraun.
Kopf fein und gedrängt punktiert, Punktabstände kleiner als Durchmesser.
Frons und Clypeus weitläufiger punktiert. Schmale, punktfreie Mittellinie von der
Breite zweier Punkte nur auf dem Clypeus erkennbar. Pronotum etwas feiner und
weitläufiger als der Kopf punktiert, Punktabstände so groß wie Durchmesser, mit
WESTMEDITERRANE SCOPAEUS - ARTEN 315
ABB. 34-37
S. proculus: (9 -"Cotype" von S. cordifer): 34) laterales Tergalteil 9, 35) Valve, 36) Tergit 10,
37) Spermatheka, Maßstabeinheit 0,1 mm.
schmaler, punktfreier Mittellinie. Elytren etwas stärker als der Kopf und zum
Hinterrand nicht feiner oder weitläufiger punktiert, die Punktabstände entsprechen
dem Durchmesser. Abdomen so stark und dicht wie die Elytren, auf den Tergiten 7
und 8 aber zunehmend feiner und weitläufiger punktiert.
Chagrinierung obsolet, Gesamteindruck glänzend. Kopf verloschen chagri-
niert, Frons und Clypeus ohne Mikroskulptur. Pronotum und Elytren glänzend.
Mikroskulptur des Abdomens obsolet.
Kopf etwas länger als breit (0,45 : 0,47, Index 1,09), Schläfen hinter den
Augen schwach erweitert und fast doppelt so lang wie die Augen (0,14 : 0,26, Index
0,54). Schläfen ab der Hälfte ihrer Länge unter deutlicher Richtungsänderung zu den
stark gerundeten Hinterecken verengt. Seitenzähne des Labrums mehr als halb so lang
wie die Mittelzähne.
Pronotum schlank (0,46 : 0,39, Index 1,18), hinter den Vorderecken gleich-
mäßig gerundet. Etwas kürzer (Index 1,11) und ein Fünftel schmaler als der Kopf
(Index 1,21). Punktfreie Mittellinie von der Breite der Vorderschienenbasis auf dem
hinteren Drittel und dem vorderen Viertel schwach gefurcht.
Elytren ein Fünftel länger als breit (NL 0,54, SL 0,64, B 0,54, SL : B 1,19), gut
ein Drittel länger als das Pronotum (PL : SL 0,72). Holopter. Abdominaltergit 7 mit
Palisadensaum.
Antennen 0,89 mm. Antennomere 1—4 länger als breit, Glied 3 fast eineinhalb
mal so lang wie breit. Antennomere 5-10 quadratisch. Antennomere 1-11 (L : B):
AA A2 OO Os 0006:
316 JOHANNES FRISCH
ABB. 38-45
Scopaeus fageli: Aedoeagus von S. kerdousensis syn. n.: 38) dorsal, 39) lateral, 40) ventral. 9 -
Paratypus: 41) laterales Tergalteil 9, 42) Tergit 10, 43) Valve, 44) Spermatheka, Seitenansicht,
45) Spermatheka, Aufsicht, Maßstabeinheit 0,1 mm.
WESTMEDITERRANE SCOPAEUS - ARTEN 317
Beine schlank, Schienen ohne Erweiterung, Vordertarsen beider Geschlechter
etwas verbreitert.
MANNCHEN: Abdominalsternit 8 caudal mit breiter, dreieckiger, an der Spitze
breit verrundeter Ausrandung.
Aedoeagus (Abb. 38-40): 0,48 mm, Laterallobi mit langen, kräftigen Borsten,
die sich ventro-mediad fortsetzen. Apikallobi kurz, in Seitenansicht stark ventrad
gerundet, ihre Enden apikad geknickt und in Dorsalansicht sehr spitz ausgezogen.
Dorsallobus breit, abgestutzt und in zwei apikad orientierte Lappen ausgezogen,
zwischen die ein langer, äußerst schlanker Dorn des Endophallus ragt. Ventralseite
der Apikallobi mit zwei medianen, apikad divergierenden, ventrad stark vorgezo-
genen Lobi, die in Seitenansicht gut zu erkennen sind und den ventralen Winkel der
Apikallobi ausfüllen.
WEIBCHEN: Terminalia spärlich beborstet. Dorsale Nebenspitze der lateralen
Tergalteile 9 (Abb. 41) fehlend. Tergit 10 (Abb. 42) breit (Index 1,73), Valve (Abb.
43) schlank (Index 7,1) mit drei kräftigen Hauptborsten.
Spermatheka (Abb. 44, 45): 0,08 mm, mit sehr ungleich gestalteten Kapsel-
gliedern. Das Glied, an dem der sklerotisierte Ductus inseriert, ist in Aufsicht beil-
förmig erweitert. In Seitenansicht ist das winklig abgesetzte Spitzenteil lang und
gerade ausgezogen. Anderes Kapselglied nicht erweitert, in Seitenansicht lang haken-
förmig gebogen.
VERBREITUNG: Scopaeus fageli ist bislang nur von den Typenfundorten in
Algerien und Marokko bekannt.
VERWANDTSCHAFTSBEZIEHUNGEN: Scopaeus fageli ist mit keiner bekannten Art
in Beziehung zu bringen; die Ausbildung des Aedoeagus ist gegenwärtig nicht homo-
logisierbar.
DANKSAGUNG
Folgenden Institutionen und Kollegen möchte ich meinen herzlichen Dank für
die rasche Ausleihe des benötigten Materials aussprechen:
Deutsches Entomologisches Institut, Eberswalde, Dr L. Zerche (DEI). Institut
Royal des sciences naturelles de Belgique, Brüssel, Mr. G. Haghebaert (ISNB).
Museum für Naturkunde der Humboldt-Universität zu Berlin, Dr M. Uhlig (HUB).
Muséum d'histoire naturelle, Genève, Dr I. Löbl (MHNG). Muséum national d'his-
toire naturelle, Paris, Dr N. Berti (MHNP). Musée Guimet d'histoire naturelle, Lyon,
Dr J. Clary (MGL). Museo Civico di Storia Naturale "Giacomo Doria", Genua, Dr R.
Poggi (MCG). Naturhistorisches Museum Wien, H. Schillhammer (NMW).
Dr Cl. Besuchet (MHNG) und Dr D. Kovac, Forschungsinstitut Senckenberg,
Frankfurt am Main (FSF) möchte ich für die Unterstützung dieser Arbeit herzlich
danken. Für inhaltliche Diskussionen möchte ich Dr I. Löbl (MHNG) und Dr V.
Puthz, Schlitz, meinen herzlichen Dank aussprechen.
318 JOHANNES FRISCH
LITERATUR
BERNHAUER, M. & K. SCHUBERT. 1910. Staphylinidae 1. Jn: Junk, W. & S. Schenkling (1910):
Coleopterorum Catalogus. — Berlin, 245-252.
BINAGHI, G. 1935. Studio sul genere Scopaeus Erichson (Coleopt. Staphylinidae). — Memorie
della Societa Entomologica Italiana 14: 84-115.
COIFFAIT, H. 1952. Notes sur les Staphylinides 2. — Revue Française d’Entomologie 19: 5-16.
COIFFAIT, H. 1960. Démembrement du genre Scopaeus et description de 4 nouvelles espèces. —
Revue Française d’Entomologie 27: 283-290.
COIFFAIT, H. 1968. Scopaeus nouveaux ou mal connus de la région paléarctique occidentale. —
Bulletin de la Société d'histoire naturelle de Toulouse 104: 405-426.
COIFFAIT, H. 1973. Staphylinides nouveaux ou mal connus du Maroc. — Bulletin de la Société
d'histoire naturelle de Maroc 53, 269-291.
COIFFAIT, H. 1981. Staphylinides nouveaux du Moyen Orient appartenant au musée hongrois
d'histoire naturelle. — Folia Entomologica Hungarica 42: 15-23.
COIFFAIT, H. 1984. Coléoptères Staphylinides de la région paléartique occidentale 5: sous
famille Paederinae 2, sous famille Euaesthetinae. — Publications de la Nouvelle Revue
d’Entomologie, Supplement a la Nouvelle Revue d’Entomologie T. 13 fasc. 4, Labo-
ratoire de Zoologie de l'Université Paul Sabatier, Toulouse: 148-208.
FRISCH, J. 1994. Neue Arten der Gattung Scopaeus Erichson aus Griechenland, Anatolien und
dem Iran (Coleoptera, Staphylinidae, Paederinae). 1. Beitrag zur Kenntnis der Gattung
Scopaeus Erichson. — Coleoptera 2: 1-46.
GANGLBAUER, L. 1895. Die Käfer von Mitteleuropa 2. Familienreihe Staphylinoidea. 1. Theil:
Staphylinidae, Pselaphidae. — Wien, 526-530.
GEMMINGER, M. & E. HAROLD. 1868. Catalogus Coleopterorum 2. Staphylinidae. — München,
618, 619.
FAUVEL, A. 1874. Faune gallo-rhénane 3, Suppl. — Caen, 311.
GusAROV, V. 1991. New and little known Palearctic Staphylinidae (Coleoptera) 3. — Vestnik
Leningradskogo Universiteta Biol. 4: 3-12, 121.
GUSAROV, V. 1992. New and little known Palearctic Staphylinidae (Coleoptera) 4. — Vestnik St.
Petersbg. Universiteta Ser. 3 Biol. 1: 11-25, 117.
GUSAROV, V. 1994. New and little-known palaearctic Paederinae (Coleoptera, Staphylinidae). —
Annales de la Société Entomologique de France 30(4): 431-446.
HENNIG, W. 1982. Phylogenetische Systematik. — Berlin-Hamburg, 246 pp.
HORION, A. 1965. Faunistik der mitteleuropäischen Käfer. Staphylinidae, 2. Teil: Paederinae bis
Staphylininae. — Uberlingen, 41.
MULSANT, E. & CL. REY. 1861a. Description de quelques Coléoptères nouveaux ou peu connus.
— Opuscules Entomologiques 12: 149-152.
MULSANT, E. & CL. REY. 1861b. Description de quelques Coléoptères nouveaux ou peu connus.
— Annales de la Société Linnéenne de Lyon 8: 133.
MULSANT, E. & CL. REY. 1877. Histoire naturelle des Coléoptères de France. Brévipennes:
Pédériens. — Annales de la Société Linnéenne de Lyon 24: 195-201.
NORMAND, H. 1934. Contribution au Catalogue des Coléoptères de Tunisie 4. — Bulletin de la
Société d'histoire naturelle de l'Afrique du Nord 25, 367, 368.
PORTA, A. 1926. Fauna Coleopterorum Italiana 2, Staphylinoidea. — Piacenza, 73.
PORTEVIN, G. 1929. Histoire Naturelle des Coléoptères de France 1. — Paris, 385.
REITTER, E. 1909. Fauna Germanica. Die Käfer des deutschen Reiches 2. — Stuttgart, 148.
UHLIG, M. 1989. Zur Morphologie der weiblichen Terminalia einiger Staphylinidenarten (Cole-
optera). — Verhandlungen des 11. SIEEC Gotha 1986: 227-237.
REVUE SUISSE DE ZOOLOGIE, 103 (2): 319-322; juin 1996
Microcharmus hauseri, nouvelle espèce de Scorpion de Madagascar
(Scorpiones, Buthidae)
Wilson R. LOURENÇO
Laboratoire de Zoologie (Arthropodes), M.N.H.N., 61, rue de Buffon
75005 Paris, France.
Microcharmus hauseri, new species of scorpion from Madagascar
(Scorpiones, Buthidae). - The study of some scorpions deposited in the
collections of the Geneve Natural History Museum, conduct to the
description of a new species of Buthidae scorpion belonging to the genus
Microcharmus. The new species is the second known for this genus and
both species are endemic to the island of Madagascar.
Key-words: Scorpion - Madagascar - New species.
INTRODUCTION
Au cours d'une étude récente sur la faune des Scorpions de l'île de Madagascar
(LOURENCO, 1995), j'ai pu découvrir et décrire trois genres nouveaux appartenant à la
famille des Buthidae. Parmi ces trois genres, Microcharmus Lourenço, n'était connu
jusqu'à présent qui par la seule espèce Microcharmus cloudsleythompsoni Lourengo.
La description de la nouvelle espèce permet en outre de confirmer la validité des
caractères génériques définissant le genre Microcharmus, lesquels peuvent être
résumés ainsi:
"Scorpions de petite taille. Sternum sous-pentagonal. Peignes de petite taille
avec un nombre de dents réduit (10 à 11); absence de fulcres. Trichobothriotaxie du
type A-a, orthobothriotaxique".
Microcharmus hauseri n. sp. (Figs. 1 à 5)
Holotype: d, Madagascar (Prov. Antsiranana [anciennement Diego-Suarez], Sous-préf.
Andoany [anciennement Hell-Ville]): Ile Nosy Be, réserve naturelle intégrale «Lokobe”, forét
primaire près Ampasindava, sentier à partir de l’ancienne savonnerie, prélèvement de sol dans
les angles formés par les contreforts d’un grand arbre en décomposition mais encore debout, 80
m; 30.X1.1989; leg. Bernd Hauser (extraction par appareil Berlese à Antananarivo) Mad-89/34.
Déposé au Muséum d'Histoire naturelle de Genève.
Manuscrit accepté le 25.08.1995.
320 WILSON R. LOURENÇO
Fics 1-2
Holotype mâle, vues dorsale et ventrale.
Etymologie: Le nom spécifique est attribué en hommage au Dr. Bernd Hauser, res-
ponsable des collections d'Arthropodes au Musée de Genève.
DESCRIPTION. Coloration générale à fond jaunâtre, avec des très nombreuses taches
brunätres sur le corps, les pattes et les pédipalpes. Mains des pinces et telson jaunatres
sans taches. Chélicères très pigmentées. Sternum, hanches, processus maxillaires et
sternites très tachetés; peignes et opercule-genital jaunâtre-pale.
MORPHOLOGIE. Prosoma: plaque prosomienne avec le bord antérieur légèrement
concave. Tubercule oculaire antérieur par rapport au centre de la plaque prosomienne;
yeux médians séparés par moins de un diamètre oculaire. Trois paires d'yeux latéraux.
Carènes du prosoma et du mesosoma moyennement marquées. Granulation de la
plaque prosomienne et des tergites du mesosoma moyennement marquée. La région
antérieure de la plaque prosomienne avec une granulation plus importante. Metasoma:
carènes bien marquées sur les quatre premiers anneaux; les dorsales avec un petit
granule distal spiniforme; les ventrales plus discrètement marquées. Vème anneau
arrondi avec des carènes moins marquées que chez les autres quatre anneaux.
Anneaux I et II avec 10 carènes; III et IV à 8 carènes; anneau V avec 5 carènes.
MICROCHARMUS HAUSERI, NOUVELLE ESPECE DE SCORPION DE MADAGASCAR 321
Fics 3-5
M. hauseri, holotype mâle. 3. Chélicère, vue dorsale. 4. Pédipalpe, vue dorsale, avec la dispo-
sition des trichobothries. 5. Telson, avec soies caractéristiques.
Vésicule lisse très allongée avec des longues soies. Aiguillon court, dépourvu d'épine
ventrale. Sternites à stigmates linéaires aplatis. Peignes petits avec un nombre de
dents assez faible, 11-11, peu courant chez les Buthidae, mais caractéristique de
diverses espèces de micro-Scorpions; absence de fulcres. Sternum sous-pentagonal,
caractère retrouvé chez certains Buthidae tels les espèces des genres Charmus et
Microtityus. Pédipalpes faiblement granulés avec des carènes moyennement
marquées.Présence de granules spiniformes sur la face interne du tibia. Tranchant des
doigts mobiles des mains des pédipalpes avec 7-7 séries de granulations légèrement
obliques. Chélicères avec la dentition caractéristique des Buthidae; les deux dents
basales sont néanmoins très petites, peu marquées et difficiles à distinguer (VACHON,
1963). Trichobothriotaxie du type A-a, orthobothriotaxique (Vachon, 1973, 1975).
Eperon tibial présent. Pas de paratypes.
MENSURATIONS (en mm) DE L'EXEMPLAIRE DÉCRIT. Plaque prosomienne: longueur, 1,8,
largeur, 1,0. Metasoma: anneau I longueur, 0,9, largeur, 1,0; anneau V longueur, 2,2,
largeur, 0,9, hauteur, 0,9. Telson: largeur, 0,7, hauteur, 0,6. Pédipalpe: fémur lon-
gueur, 1,5; tibia longueur, 1,7; pince, 2,5; doigt mobile, longueur, 1,7.
322 WILSON R. LOURENCO
Microcharmus hauseri peut être facilement distinguée de Microcharmus
cloudsleythompsoni, par la présence de nombreuse taches brunâtres sur le corps,
pattes et pédipalpes, lesquelles sont absentes chez M. cloudsleythompsoni, espèce
globalement jaunâtre, et aussi par la présence de très nombreuses soies sur la vésicule,
caractère non observé chez M. cloudsleythompsoni.
REMERCIEMENTS
Je suis très reconnaissant aux Drs B. Hauser et V. Mahnert du Museum
d'Histoire naturelle de Genève de m'avoir facilité l'étude du matériel.
RÉFÉRENCES
LOURENCO, W.R. 1995. Description de trois nouveaux genres et de quatre nouvelles espèces de
Scorpions Buthidae de Madgascar. Bulletin du Museum national d'Histoire naturelle,
Paris, 4e sér., 17(1-2): 95-106.
VACHON, M., 1963. De l'utilité, en systématique, d'une nomenclature des dents des chélicères
chez les Scorpions. Bulletin du Museum national d'Histoire naturelle, Paris, 2e sér.,
35(2): 161-166.
VACHON, M., 1973. Etude des caractères utilisés pour classer les familles et les genres de Scor-
pions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et
types de trichobothriotaxie chez les Scorpions. Bulletin du Museum national d’ Histoire
naturelle, Paris, 3è sér., n° 140, Zool., 104: 857-958.
Vachon, M., 1975. Sur l'utilisation de la trichobothriotaxie du bras des pédipalpes des Scor-
pions (Arachnides) dans le classement des genres de la famille des Buthidae Simon.
Compte Rendu des séances de l’Académie des Sciences, Paris, ser. D, 281: 1597-1599.
REVUE SUISSE DE ZOOLOGIE, 103 (2): 323-338; juin 1996
New and little-known Oriental Melittia Hübner
(Lepidoptera, Sesiidae),
from the collection of Muséum d'histoire naturelle, Genève
Oleg G. GORBUNOV! and Yutaka ARITA2
1 Institute for the Problems of Ecology and Evolution. Russian Academy of Sciences,
Leninsky prospekt 33, Moscow V-71, 117071 Russia*
2Zoological Laboratory, Faculty of Agriculture, Meijo University, Tempaku-ku,
Nagoya, 468 Japan.
New and little-known Oriental Melittia Hübner (Lepidoptera, Sesii-
dae), from the collection of Muséum d'histoire naturelle, Genève. - A
small collection of clearwing moth of the genus Melittia Hübner, 1819
from Vietnam and the Philippines, deposited in the Muséum d'histoire
naturelle, Genève, Switzerland, is revised. Two new species, M. romieuxi
sp. n. and M. luzonica sp. n., are described from Vietnam and the Philip-
pines, respectively. M. newara Moore, 1879 and M. eurytion (Westwood,
1848) are recorded for Vietnam for the first time and the latter species for
Palawan Id, Philippine Is. as well. A redescription of M. gorochovi
Gorbunov, 1988 is presented. Melittia eurytion ab. microfenestrata Strand,
[1916] is synonymised under Melittia formosana Matsumura, 1911.
Key-words: Sesiidae - Melittia - new species - taxonomy - Vietnam - the
Philippines.
INTRODUCTION
The genus Melittia Hübner, [1819] is, perhaps, the largest genus of Sesiidae. It
is distributed worldwide in tropical and subtropical areas and consists of more than a
hundred presently known species. Unfortunately, most of them have been, in the past
up to the beginning of this century, described from a single or a few specimens. Their
original descriptions are very laconic. The descriptions contain, as a rule, only a few
external characters. This considerably hampers the study of these beautiful clearwing
moths. In addition, very important data about the types, type localities, etc., are
* Visiting Scholar of Zoological Laboratory, Faculty of Agriculture, Meijo University,
Tempaku-ku, Nagoya, 468 Japan
Manuscript accepted 15.09.1995.
324 OLEG G. GORBUNOV & YUTAKA ARITA
scattered among numerous books and periodicals, which in many cases are rare and
nearly unavailable. The majority of the species Melittia are highly beautiful, brightly
coloured and superficially bumble-bee-like species. They are diurnal heliophile
moths, but sometimes can be collected by the light trap. À number of everywhere
known larvae of Melittia live inside the stalks of Cucurbitaceae, especially of the
genus Trichosanthes (cucumber, pumpkin), and they may be easily reared.
Material was studied from Muséum d'histoire naturelle, Genève (MHNG). —
The Natural History Museum, London, Great Britain (BMNH). — Museum für
Naturkunde, Zentralinstitut der Humboldt-Universität zu Berlin, Germany (ZMHB).
— collection O. Gorbunov, Moscow, Russia (CG).
The holotypes of new species described herein are deposited in MHNG,
Switzerland.
Melittia gorochovi Gorbunov (Figs 1, 2, 9)
Melittia gorochovi Gorbunov, 1988: 195, figs 3-/, 4-1. Type locality: Vietnam, Shonla
Prov., Shongma. Holotype male (CG).
= Melittia indica Butler, 1874 — Le Cerf, 1917: 182 (part.), Pl. 476, Fig. 3922
(misdetermination).
MATERIAL EXAMINED: | male (holotype), Vietnam, Shonla prov., Shongma vic.,
3-10.V.1986, A.V. Gorokhov leg. (genital preparation No. G0064) (CG); 1 male, Vietnam,
Sam Con a Vang Lom, 10.1V.1950, J. Romieux leg. (genital preparation No. GA-050)
(MHNG).
REDESCRIPTION. Male (holotype) (Fig. 1). Alar expanse 29.5 mm; body length
15.5 mm; forewing 13.0; antenna 7.0 mm.
Head: antenna dorsally black with violet sheen, with a few snow-white scales
dorso-internally and an elongate yellow spot dorso-externally at tip; ventrally light
brown; frons grey-brown with purplish sheen, with a narrow pale yellow strip later-
ally; labial palpus basally pale yellow, apically black with a few yellow scales, second
joint yellow with two, narrow, black stripes both ventro-internally and ventro-
externally; vertex mixed with black and yellow scales; pericephalic hairs dorsally
mixed with olive, black and a few white scales, laterally pale yellow to white.
Thorax: patagia dorsally mixed with yellow-olive, light brown and a few black
scales; tegula, meso- and metathorax dark brown with violet sheen, with admixture of
yellow-olive scales; thorax laterally mixed with grey-brown with violet sheen, pale
yellow and yellow-olive scales.
Legs: fore coxa pale yellow with admixture of individual black scales basally;
mid tibia internally black; externally at base and apically narrowly black; medially
yellow ventrally and light brown-yellow with a small white spot dorsally; mid tarsus
black with a small white spot both at base of first and second tarsomeres externally;
hind tibia externally narrowly black at base; black with violet sheen, with a small
white spot ventro-externally; dorso-externally light brown-yellow with a small snow-
white spot medially and with a small yellow spot at base of apical spurs; dorsally
yellow with a few light brown-orange scales at tip; spurs black; hind tarsus black with
greenish sheen, with a small pale yellow spot medio-externally.
ORIENTAL MELITTIA B25)
Abdomen: dorsally black with bluish sheen; tergites 1 and 2 densely covered
with dirty olive-green scales; tergites 3-7 each with a narrow, olive-green, distal
margin; ventrally entirely pale yellow; anal tuft small, black with a few yellow scales.
Forewing: basally densely covered with yellow-olive and light brownish
scales; costal margin and veins within external transparent area black with violet
sheen; Cu-stem, anal margin and discal spot black with violet sheen, with a few light
brownish scales; apical area black with violet sheen, with a few snow-white scales
with bluish hue; discal spot relatively broad with a long, narrow, cuneiform projection
proximally; transparent areas well-developed; external transparent area relatively
large, divided into 6 cells (cell between veins R4 and R5 minute), on level of vein M1
about 2.5 times as broad as discal spot and ca. 1.5 times as broad as apical area; cilia
dark brown with bronze-violet sheen.
Hindwing: transparent; anal area black with violet sheen, densely covered with
yellow-olive scales and hairs; veins, discal spot and outer margin black with violet
sheen; discal spot extremely narrow and nearly undeveloped; outer margin ca. thrice
narrower than cilia; cilia dark brown with bronze-violet sheen.
Male genitalia (genital preparation No. MHNG-2480, GA-050) (Fig. 9).
Tegumen-uncus complex narrow; uncus bilobed distally with a small drop-shaped plate
of strong pointed setae internally on each side; gnathos small, narrow, membranous with
slight sclerotization basally (Fig. 9a); valva (Fig. 9b) elongate-oval; distal field of setae
rather well separated from medial one; medial field narrow with short setae; pocket-
shaped crista small; ventral lobe relatively broad and long, somewhat exceeding distal
margin; saccus relatively broad, mace-shaped basally (Fig. 9c); aedeagus (Fig. 9d)
narrow, broadened basally, about as long as valva; vesica with numerous minute cornuti.
Female. Unknown.
Variability. The second specimen of this species from Vietnam (Fig. 2) vir-
tually has neither coloration nor size differences from the holotype, only the external
transparent area of the left forewing divided into 5 cell, so a minute cell between
veins R4-RS is absent.
Diagnosis. M. gorochovi seems to be closest to Melittia newara Moore, 1879,
but differs clearly from it by the shape of the external transparent area of the forewing
(broader in newara (Fig. 3)), by the somewhat different coloration of the hind leg tuft,
and by the shape of the male genitalia (gnathos somewhat less developed and
sclerotized, pocket-shaped crista less developed and ventral lobe less broad, but slightly
longer in newara). From M. binghami Niceville, 1900, gorochovi can be distinguished
by the somewhat smaller size (alar expanse 35.0 mm in binghami), by the coloration of
the hind leg tuft (more yellow in the species compared), and by the shape of the
external transparent area of the forewing (broader and cell between veins R4-RS larger
in binghami). From M. indica Butler, 1874 and M. siamica Walker, [1865], gorochovi
is clearly distinguishable by the shape of the external transparent area of the forewing
(smaller, without hyaline cell between veins R4-RS in these species compared) and by
the brighter coloration of the hind leg tuft (with less yellow scales in indica and
siamica).
326 OLEG G. GORBUNOV & YUTAKA ARITA
Bionomics. The host plant unknown. Moths have been netted in April and
May.
Habitat. Borders of tropical forest; river valleys; road sides.
Distribution. Hitherto known only from Vietnam and northeastern India
(Sikkim).
Melittia newara Moore (Figs 3, 10)
Melittia newara Moore, 1879: 10. Type locality: "Darjiling" [= NE India, West Bengal,
'Darjeeling]. Holotype male (ZMHB).
MATERIAL EXAMINED: | male, Vietnam, Pahia, Col Bien, 16.1V.1950, J. Romieux leg.
(genital preparation No. MHNG-2467, GA-052) (MHNG).
Remarks. This species will be revised by us (Gorbunov & Arita, 1995). The
specimen from Vietnam (Fig. 3) slightly smaller than holotype from Darjeeling: alar
expanse 32.0 mm; body length 15.8 mm; forewing 14.2 mm; antenna 7.0 mm. Also,
this specimen has no small hyaline cell between veins Cul and Cu2, the external
transparent area of the forewing of this specimen is divided into 6 cells, and the
proximal projection of the discal spot of the forewing is somewhat shorter (see loc.
cit., text figs 1-2). The male genitalia (Fig. 10) of this specimen has a somewhat
narrower ventral lobe of the valva only. This is first record of the species for Vietnam.
Melittia romieuxi sp. n. (Figs 4, 11)
MATERIAL EXAMINED: 1 male (holotype), Vietnam, Pahia, 19.V.1950, J. Romieux leg.
(genital preparation No. MHNG-2483, GA-053) (MHNG).
DESCRIPTION. Male (holotype) (Fig. 4). Alar expanse 29.0 mm; body length
13.8 mm; forewing 12.5 mm; antenna 6.7 mm.
Head: antenna dorsally black with violet sheen, ventrally light brown with a
small yellow spot subapically; frons grey-brown with purplish sheen, with a narrow
white stripe laterally; labial palpus white mixed with dark brown, with a narrow black
stripe extro-ventrally on second joint; vertex grey-brown mixed with pale yellow;
pericephalic hairs black dorsally and white laterally.
Fics 1-8
Melittia spp. 1. M. gorochovi Gorbunov, 1988, holotype, male (CG). Alar expanse 29.5 mm. 2.
M. gorochovi Gorbunov, 1988, male, Vietnam, Sam Con à Vang Lom, 10.1V.1950, J. Romieux
leg. (genital preparation No. GA-050) (MHNG). Alar expanse: 30.0 mm. 3. M. newara Moore,
1879, male, Vietnam, Pahia, Col Bien, 16.1V.1950, J. Romieux leg. (genital preparation No.
GA-052) (MHNG). Alar expanse: 32.0 mm. 4. M. romieuxi sp. n., holotype, male (MHNG).
Alar expanse: 29.0 mm. 5. M. nepcha Moore, 1879, male, Vietnam, Shonla prov., Shongma,
Ma River, 4-9.X.1987, N.L. Orlov leg. (CG). Alar expanse: 32.0 mm. 6. M. luzonica sp. n.,
holotype, male (MHNG). Alar expanse 28.0 mm. 7. M. eurytion (Westwood, 1848), male,
Vietnam, Sam Con à Vang Lom, 10.1V.1950, J. Romieux leg. (MHNG). Alar expanse: 29.0
mm. 8. M. eurytion (Westwood, 1848), female, Philippines, Palawan, Puerto Princesa, oct.
1925 (MHNG). Alar expanse: 29.5 mm.
327
ORIENTAL MELITTIA
328 OLEG G. GORBUNOV & YUTAKA ARITA
Thorax: patagia brown with green-bronze sheen, with admixture of olive-green
scales anteriorly and laterally; tegula, meso- and metathorax grey-brown with bronze-
purple sheen, covered with olive-green scales; thorax laterally grey with purplish
sheen, with a few white scales.
Legs: fore coxa black with bronzed sheen, with pale yellow to white stripe
baso-internally; mid tibia externally black with green-violet sheen, with a small white
with blue hue spot medially; internally grey with bronze-purple sheen, with a small
pale yellow to white spot apically; spurs black; hind legs broken off.
Abdomen: dorsally green-brown; tergites 1 and 2 covered with thin olive-green
scales; tergite 2 with a narrow, yellow to pale yellow, margin distally; distal margin of
third tergite laterally with a few yellow scales; ventrally grey mixed with whitish
scales; anal tuft small grey-brown with a few whitish scales.
Forewing: basally black with a few olive-green and grey scales; costal and
anal margins, Cu-stem, discal spot, veins within external transparent area and apical
area black with violet sheen; discal spot narrow with a small cuneiform broadening
proximally; apical area very narrow, developed only at tip of wing; transparent areas
well-developed; external transparent area extremely large, rounded apically, divided
into 7 cells (cell between veins Cul and Cu2 narrow), about 7 times as broad as discal
spot; cilia dark brown with bronze-violet sheen.
Hindwing: transparent; anal area dark brown to black with violet sheen, with a
few pale yellow scales and yellow hairs; veins and outer margin black with violet
sheen; discal spot extremely narrow; outer margin narrow, ca. twice as narrow as
cilia; cilia dark brown with bronze-violet sheen.
Male genitalia (holotype, genital preparation No. MHNG-2483, GA-053) (Fig.
11). Tegumen-uncus complex narrow; uncus bilobed distally with a small drop-
shaped plate of strong pointed setae internally on each side; gnathos small, narrow,
membranous, nearly without sclerotization (Fig. 11a); valva (Fig. 11b) elongate-oval;
distal field of setae rather well separated from medial one; medial field triangular,
with short setae; pocket-shaped crista small; ventral lobe broad and short, not
exceeding distal margin; saccus relatively broad, mace-shaped basally (Fig. 11c);
aedeagus (Fig. 11d) narrow, broadened basally, somewhat longer than valva; vesica
with a few minute cornuti.
Female. Unknown.
Variability. Unknown.
Diagnosis. Superficially, this new species appears closest to M. newara
Moore, 1879, but can be separated from it by the coloration of the vertex (dark brown
to black mixed with yellow in newara), abdomen dorsally (dark brown to black with
purplish sheen; tergite 2 covered with thin olive-green scales basally; tergites 2 and 3
each with a narrow yellow margin distally in newara), and anal area of the hindwing
(dark brown to black with a narrow yellow strip in the species compared), and by the
form of the discal spot of the forewing (relatively narrow with a narrow and long
projection proximally in newara). From other closely related species (M. gorochovi
Gorbunov, 1988, M. callosoma Hampson, 1910, M. indica Butler, 1874, M. kulluana
ORIENTAL MELITTIA 329
Moore, 1888, and M. proxima Le Cerf, 1917), romieuxi sp. nov. is distinguishable by
the larger external transparent area of the forewing (relatively narrow, divided into
5-6 cells in all these species compared (see Figs 1-2)) and by the shape of the discal
spot of the forewing (with a relatively long, narrow, cuneiform projection proximally
in the species compared). Besides that, romieuxi sp. nov. clearly differs from all these
congeners by the structure of the male genitalia, especially the shape of the valva
(compare with Figs 9-10).
Bionomics. The host plant unknown. Holotype specimen has been collected in
May.
Habitat. Unknown.
Distribution. Known only from the type locality.
Etymology. We name this species after J. Romieux, who collected that species
and many other Sesiidae in Vietnam which we have studied.
Melittia nepcha Moore (Fig. 5)
Melittia nepcha Moore, 1879: 10. Type locality: Darjiling [= northeastern India, West
Bengal, Darjeeling]. Holotype male (ZMHB).
= Melittia amboinensis vietnamica Gorbunov
Melittia amboinensis vietnamica Gorbunov, 1988: 195, Figs. 2, 4-2. Type locality:
North Vietnam, Shonla prov., Shongma. Holotype male (CG).
GORBUNOV & ARITA, 1995 (as a synonym of M. nepcha Moore, 1879).
MATERIAL EXAMINED: | male (holotype of M. amboinensis vietnamica Gorbunov,
1988), Vietnam, Shonla prov., Shongma, 3—10.V.1986, A.V. Gorokhov leg. (CG); 1 male,
Vietnam, Shonla prov., Shongma, Ma River, 4-9.X.1987, N.L. Orlov leg. (CG).
Remarks. This species has been revised by the authors (Gorbunov & Arita,
1995). Specimens from Vietnam slightly vary in individual size: alar expanse 31.5—
32.0 mm; body length 15.2-16.0 mm; forewing 13.5-14.0 mm; antenna 6.5-7.0 mm
only and have not differences from the holotype in coloration.
Melittia luzonica sp. nov. (Figs 6, 12)
= Melittia siamica Walker, [1864] — Diakonoff, [1968]: 233, Figs 722—723 (misdeter-
mination).
MATERIAL EXAMINED. | male (holotype), Philippines, Luzon Id., Binoessoc, (genital
preparation No. MHNG-2482, GA-078) (MHNG).
DESCRIPTION. Male (holotype) (Fig. 6). Alar expanse 28.0 mm; body length
14.0 mm; forewing 11.6 mm; antenna 6.2 mm.
Head: antenna dorsally dark brown to black with purple-violet sheen, with
individual snow-white scales at fore margin, ventrally yellow; frons grey-brown with
a very narrow white stripe laterally; labial palpus dark brown to black with a few
white scales ventrally; vertex dark brown mixed with black hair-like scales, with a
narrow, V-shaped, white stripe between ocelli posteriorly; pericephalic hairs black
dorsally and white laterally.
Thorax: patagia black with violet sheen, with a small pale yellow spot late-
rally; tegula and mesothorax dark brown with violet sheen; metathorax grey-brown
330 OLEG G. GORBUNOV & YUTAKA ARITA
with two tufts of hair-like scales dark grey mixed with dirty yellow; thorax laterally
dark grey with violet sheen, with a few pale yellow scales anteriorly.
Legs: fore coxa dark brown with greenish sheen, with a few pale yellow scales
basally; mid tibia externally dark brown to black with green-purple sheen, with
admixture of individual rusty scales, with a small, diffuse, snow-white with blue hue
spot at basal third, and with a small dirty yellow spot dorso-apically; mid tarsus
yellow to pale yellow, externally four apical tarsomeres black with greenish sheen;
hind tibia mixed with dirty yellow, light brown, dark brown and a few white scales,
with a small snow-white with blue hue spot at base of mid spurs externally; spurs dark
brown to black with violet sheen; hind tarsus black with green sheen, with a few
white and dirty yellow scales basally and with a small snow-white spot medio-
ventrally.
Abdomen: dorsally tergite 1 dark grey, remaining tergites black with violet
sheen; tergites 2-4 and 6-7 each with a narrow, distinct, pale yellow stripe distally;
ventrally dark brown with greenish sheen; sternite 3 with a sparse pale yellow to
white stripe distally; sternites 4-6 each with a small pale yellow to white spot laterally
on distal margin; anal tuft small, dark brown to black with greenish sheen, mixed with
yellowish scales distally.
Forewing: costal margin, Cu-stem, discal spot, veins within external trans-
parent area and apical area black with bronzed sheen; anal margin black with bronzed
sheen, with admixture of individual olive-brown scales; discal spot narrow with a
short pointed projection proximally; transparent areas well-developed; external
transparent area rounded distally, divided into 6 cells (including minute cell between
veins R4-RS), ca. 4.5 times broader than discal spot on level of vein M2; cilia dark
grey with bronzed sheen.
Hindwing: transparent; anal area covered with dark brown and olive-brown
scales; veins and outer margin narrowly black; discal spot undeveloped; outer margin
about thrice narrower than cilia; cilia dark grey with bronzed sheen.
Male genitalia (holotype, preparation No. MHNG-2482, GA-078) (Fig. 12).
Tegumen-uncus complex narrow; uncus bilobed distally with a small drop-shaped
plate of strong pointed setae internally on each side; gnathos rather large, broad,
membranous with well-sclerotized oval plate basally; pedunculus with a long seta
(Fig. 12a); valva (Fig. 12b) trapeziform; distal field of setae nearly not separated from
medial one; setae of medial field relatively short, slightly covering pocket-shaped
crista; ventral lobe relatively narrow, barely exceeding distal margin; saccus (Fig.
12c) narrow, gradually broadened basally; aedeagus (Fig. 12d) narrow, somewhat
shorter than valva; vesica with numerous minute cornuti.
Fics 9-10
Male genitalia of Melittia spp. 9. M. gorochovi Gorbunov, 1988 (genital preparation No. GA-
050). a. Tegumen-uncus complex. b. Valva. c. Saccus. d. Aedeagus. Scale bar: 0.5 mm. 10. M.
newara Moore, 1879 (genital preparation No. GA-052). a. Tegumen-uncus complex. b. Valva.
c. Saccus. d. Aedeagus. Scale bar: 0.5 mm.
331
ORIENTAL MELITTIA
332 OLEG G. GORBUNOV & YUTAKA ARITA
Female. As far as we can tell from the figures published by DIAKONOFF (1968,
Figs 722 and 723), the female of this new species has virtually no differences in
coloration from male, but perhaps is more robust than male.
Variability. Unknown.
Diagnosis. This new species belongs to the amboinensis species-group. It
seems to be the closest to M. nepcha Moore, 1879, but can be distinguished by the
somewhat different coloration of the hind leg luft (dark brown both basally and
apically, medially mixed with brown, light brown to yellow-brown and snow-white
scales; extro-ventrally with three small white spots between bases of tibia and mid
spurs, and somewhat distally both mid and apical spurs in nepcha) and, especially, by
the coloration of the abdomen (dorsally dark brown to black mixed with olive-green
scales on tergites 2, 3 and 5; all tergites with a narrow, olive-green, distal margin;
ventrally entirely yellow to pale yellow in nepcha). However, these two species are
easy separable by the male genitalia (setae of medial field of valva long, entirely
covering pocket-shaped crista; saccus mace-shaped basally in nepcha). From M.
amboinensis Felder, 1861, luzonica sp. nov. differs also by the coloration of the hind
leg (with more rusty-brown scales dorsally in amboinensis) and abdomen (without a
narrow, distinct, pale yellow stripe distally in amboinensis). From M. batchiana Le
Cerf, 1917 this new species is clearly distinguishable by the more developed
transparent areas of the forewing (external transparent area divided into 4 cells, on
level of vein M2 about 1.6 times broader than apical area in batchiana). From other
closely related species, such as M. dorsatiformis Hampson, 1891, M. distincta Le
Cerf, 1916, M. congruens Swinhoe, 1890, and presently taxonomically unclear taxa
described by F. Le Cerf (1916) (meeki, celebica, javana, doddi), luzonica sp. nov.
clearly differs by the coloration of the abdomen (dorsally without a narrow, distinct,
pale yellow, distal stripe; ventrally entirely pale yellow to yellow in all these taxa
compared).
Bionomics and habitat. Unknown.
Etymology. This new species is named after the island Luzon, Philippine Is.,
where this species was collected.
Melittia eurytion (Westwood, 1848) (Figs 7-8, 13-14)
Trochilium enrytion Westwood, 1848: 62, Pl. 30, Fig. 5. Type locality: "India Sylhet"
[= NE Bangladesh, Sylhet]. Lectotype female (BMNH) (designated by Spatenka, 1992).
Melittia eurytion (Westwood, 1848) — Diakonoff, [1968]: 233, Figs 726-727.
Fics. 11-12
Male genitalia of Melittia spp. 11. M. romieuxi sp. nov., holotype (genital preparation No. GA-
053). a. Tegumen-uncus complex. b. Valva. c. Saccus. d. Aedeagus. Scale bar: 0.5 mm. 12. M.
luzonica sp. nov., holotype (genital preparation No. GA-078). a. Tegumen-uncus complex. b.
Valva. c. Saccus. d. Aedeagus. Scale bar: 0.5 mm.
ORIENTAL MELITTIA 333
334 OLEG G. GORBUNOV & YUTAKA ARITA
MATERIAL EXAMINED. | male, Vietnam, Sam Con a Vang Lom, 10.1V.1950, J. Romieux
leg. (MHNG); 1 male, Vietnam, Col Noé, Lao Sang, 19.V.1950, J. Romieux leg. (genital
preparation No. GA-056) (MHNG); 1 female, Philippines, Manila, Palanoz, Apr. 28 (genital
preparation No. GA-084) (MHNG); 1 female, Philippines, Palawan, Puerto Princesa, oct. 1925
(MHNG).
DESCRIPTION. Male (Fig. 7). Alar expanse 29.0 mm; body length 14.5 mm;
forewing 12.5 mm; antennae broken off.
Head: antenna broken off; frons grey-brown with purple sheen, with a narrow
white stripe laterally; vertex grey-brown with purple sheen mixed with black, white
and yellow hair-like scales; labial palpus white basally; mid joint white with two
black longitudinal stripes ventrally at margins, apical joint black with a few pale
yellow scales; pericephalic hairs dorsally black mixed with yellow, laterally pale
yellow to white.
Thorax: patagia dorsally dark brown with green-purple sheen, anteriorly
covered with more thin olive-green scales, laterally pale yellow to white; tegula and
mesothorax dark brown to black with purple-gold sheen, covered with a few olive-
green scales anteriorly; metathorax somewhat paler with two tufts of white hair-like
scales laterally and a few white with blue hue scales medially; thorax laterally mixed
with grey and white scales.
Legs: fore coxa white to pale yellow with admixture of a few dark brown
scales; mid tibia dark brown with gold sheen, with a small white with blue hue spot
both medio-externally and ventrally; hind tibial tuft black with greenish sheen, mixed
with white dorsally and externally, with two large white with blue hue spots both
somewhat basally of mid spurs and between base of both pairs of spurs ventrally;
spurs black, external spurs of both pairs with white inside; hind tarsal tuft black with
greenish sheen, with a small white spot medio-externally.
Abdomen: dorsally black with green-violet sheen, with admixture of indivi-
dual, narrow, white scales; scales of distal margin of each tergite with bronzed sheen;
additionally, tergites 2, 4, 6 and 7 each with a few white with blue hue scales distally;
ventrally sternite 1+2 black with a few white scales, other sternites entirely white;
anal tuft small, black, tipped white.
Forewing: costal and anal margins, Cu-stem, discal spot, apical area and veins
within external transparent area dark brown to black with bronze-purple sheen;
basally at anal margin with admixture of individual olive-green and white with blue
hue scales; Cu-stem basally shortly and narrowly bordered with white scales; apical
area with individual snow white with blue hue scales; discal spot broad with a long
cuneiform projection proximally; transparent areas well-developed; external trans-
parent area divided into 5 cells, narrowed costally, about as broad as apical area and
discal spot on level of vein M2; cilia dark brown with bronzed sheen.
Hindwing: transparent; anal area black, but densely covered with light bluish
scales; veins, discal spot and outer margin narrowly black with bronze-purple sheen;
cilia dark brown with bronzed sheen.
Male genitalia (genital preparation No. MHNG-2468, GA-056) (Fig. 13).
Tegumen-uncus complex narrow; uncus bilobed distally with a small oval plate of
ORIENTAL MELITTIA 335
Fic.13
Male genitalia of Melittia eurytion (Westwood, 1848) (genital preparation No. GA-056). a.
Tegumen-uncus complex. b. Valva. c. Saccus. d. Aedeagus. Scale bar: 0.5 mm.
strong pointed setae internally on each side; gnathos rather large, broad, membranous
with broad but slightly sclerotized plate medially (Fig. 13a); valva (Fig. 13b)
trapeziform; distal field of setae rather well separated from medial one; setae of
medial field relatively long, not reaching pocket-shaped crista; ventral lobe relatively
narrow, barely exceeding distal margin; saccus (Fig. 13c) narrow, mace-shaped
basally; aedeagus (Fig. 13d) narrow, broadened basally, somewhat longer than valva;
vesica with numerous minute cornuti.
Female (Fig. 8). Differs little from the male as to coloration and size, though
slightly more robust.
Female genitalia (genital preparation No. MHNG-2481, GA-084) (Fig. 14).
Papilla anales slightly sclerotized, covered with short setae; 8th tergite relatively
broad with relatively short setae at distal margin and with two long setae at inner
margin ventrally; both apophyses nearly equal in length; apophyses anterior with a
336 OLEG G. GORBUNOV & YUTAKA ARITA
long, narrow appendix baso-ventrally; ostium bursae opening near posterior margin of
7th sternite, slightly funnel-shaped, narrow, ring-shaped well-sclerotized; antrum
narrow, membranous; ductus bursae narrow, relatively long, membranous; corpus
bursae nearly pear-shaped, membranous with numerous wrinkles, with signum rela-
tively large, narrowly pear-shaped, with numerous small, well-sclerotized thorns, with
ca. 10 transverse, rather well-sclerotized, dentate stripes anteriorly, shortly bifurcate
posteriorly (Fig. 14b).
Variability. This wide-spread species of the Oriental Region is extremely
variable in individual size (alar expanse 26.0-33.0 mm), coloration of the labial
palpus and legs (more or less white or pale yellow scales) and, especially, in the size
of the external transparent area of the forewing (divided into 4-5 cells and narrower
than apical area and discal spot on level of vein M2 about 1.1-2.0 times). Also, it
slightly varies in the male (shape of gnathos and valva) and female (signum of corpus
bursae) genitalia.
Diagnosis. It seems to be the closest to M. formosana Matsumura, 1911, M.
chalybescens Miskin, 1892, M. flaviventris Hampson, 1919, and M. volatilis Swinhoe,
1890. Superficially, from formosana, eurytion can be distinguished by the relatively
narrower discal spot of the forewing, but tney rather clearly differ by the male
genitalia (valva shortly oval; distal and medial fields of setae nearly not separated;
saccus narrower in the species compared). From chalybescens, eurytion is
distinguishable by the coloration of the anal area of the both wings (bright blue-green
in chalybescens) and by the male genitalia (valva broadly oval; saccus bilobed basally
in chalybescens). From flaviventris, this species can be easily separated by the shape
of the anterior and external transparent areas of the forewing (anterior transparent area
not divided into two longitudinal stripes by proximal projection of discal spot;
external transparent area divided into 5—6 cells, about 1.8 times as broad as apical
area on level of vein M2 in flaviventris). From volatilis, eurytion clearly differs by the
more well-developed transparent areas of the forewing (anterior transparent area very
short and narrow; external transparent area divided only into one small cell between
veins M3-Cul in volatilis).
Bionomics. Specific host plant unknown, but most likely a species of the
family Cucurbitaceae. Imagos collected in April-May.
Habitat. Unknown.
Distribution. This species has been reported from West China, Nepal, West
(Bombey), Northeast (Sikkim) and East India (Assam), Sri Lanka, Myanmar, Phi-
lippines (Luzon, Mindanao). We record it for Vietnam and Palawan Id. (Philippines)
for the first time. Additionally, E. STRAND [1916] described an aberration of M.
Fic. 14
Female genitalia of Melittia eurytion (Westwood, 1848) (genital preparation No. GA-084). a.
General view. b. Corpus bursae. Scale bar: 0.5 mm.
337
ORIENTAL MELITTIA
Di
i
338 OLEG G. GORBUNOV & YUTAKA ARITA
eurytion, microfenestrata Strand [1916] from Taiwan [Formosa]. At present we
believe that all citations of eurytion for Taiwan should be regarded to M. formosana
Matsumura, 1911 and indicate Melittia eurytion ab. microfenestrata Strand, [1916]
(comb. nov. et syn. nov.) as a new synonym of M. formosana Matsumura, 1911
herein.
ACKNOWLEDGEMENTS
We would like to express our cordial thanks to Dr D.B. Burckhardt, Muséum
d'histoire naturelle, Genève, Switzerland, and Dr S.I. Golovatch, Institute for the
Problems of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia
for the opportunity to study this highly interesting collection. We also thank Dr T.D.
Eichlin, California Department of Food and Agriculture, Plant Pest Diagnostics
Branch, Sacramento, California, USA for linguistic help.
REFERENCES
DIAKONOFF, A.N., 1968. Microlepidoptera of the Philippine Islands. Bull. US Nat. Mus. 257:
1-458.
GORBUNOV, O.G., 1988. A new contribution to the knowledge of clearwing moths (Lepidoptera,
Sesiidae) of Vietnam. /n: Medvedev, L.N. & B.R. Striganova (editors), The Fauna and
Ecology of Insects of Vietnam: 192-198, Figs. 1-4. Moscow: Nauka Publ. (in Russian).
GORBUNOV, O.G & Y. ARITA, 1995. A revision of Frederic Moore's clearwing moths types (Lepi-
doptera, Sesiidae), at Humboldt University, Berlin. Tinea, 14(3): 204-224, figs 1-24.
LE CERF, F., 1917. Contributions à l'étude des Aegeriidae: Description et iconographie d'es-
pèces et de formes nouvelles ou peu connues. Etud. Lépid. comp. 14: 137-388, Pls
475-481.
MOORE, F., 1879. In: Hewitson, W.C. & F. Moore. Description of New Indian Lepidopterous
Insects from the Collection of the Late Mr. W.S. Atkinson: Heterocera. [Part 1]: 5-88,
Pl. 2-8. London: Asiatic Society of Bengal, Calcutta.
SPATENKA, K., 1992. Contribution à la stabilisation de la taxinomie des Sésiides paléartiques
(Sesiidae, Lepidoptera). Alexanor, 17(8): 479-503.
STRAND, E., [1916]. Fam. Aegeriidae. In: H. Sauter's Formosa-Ausbeute: Noctuidae p. p.
(Agaristinae, Macrobrochis), Aganaidae, Saturniidae, Uranidae, Cossidae, Callidulidae
und Aegeriidae. Archiv fiir Naturgeschichte (Berlin), 81A(8): 45-49.
WESTWOOD, J.O., 1848. The Cabinet of Oriental Entomology. 88 pp., 42 Pls. London: W. Smith.
REVUE SUISSE DE ZOOLOGIE, 103 (2) : 339-354; juin 1996
Haliotids in the Red Sea, with neotype designation for Haliotis
unilateralis Lamarck, 1822 (Gastropoda: Prosobranchia)
Daniel GEIGER
University of Southern California, Department of Biological Sciences,
University Park, Los Angeles, CA 90089-0371, USA.
Haliotids in the Red Sea, with neotype designation for Haliotis uni-
lateralis Lamarck, 1822 (Gastropoda: Prosobranchia). — Two species
of Haliotis are known from the Red Sea: A. pustulata cruenta Reeve, 1846
and A. unilateralis Lamarck, 1822. A neotype is here designated for
Haliotis unilateralis because the specimen in the Lamarck collection of the
Muséum d’Histoire Naturelle de Genève (MHNG) is clearly not the
original specimen described by Lamarck, which is not traceable in the
MHNG collections. The neotype, a complete specimen, is deposited in the
MHNG. The designated type locality is Gulf of Aqaba: Sinai: Elat. The
shell is redescribed and radular and epipodial characters are compared to
those of similar species.
Key-words: Gastropoda - Prosobranchia - Haliotis unilateralis - neotype -
Red Sea
INTRODUCTION
Members of the family Haliotidae occur in all tropical and temperate oceans
(e.g. COX 1962; LINDBERG 1992), mostly in the shallow subtidal. Approximately 200
species level taxa have been described, 55 of which are considered valid species
(Geiger, in prep.). Although seven taxa have been mentioned from the Red Sea, most
authors record one or two species, usually identified as H. pustulata cruenta Reeve,
1846 and H. unilateralis Lamarck, 1822 (TALMADGE 1971; YARON 1983).
Comparison of animal characters as well as those of the shell has resulted in a
new assessment of the Red Sea species. The specimen of H. unilateralis held in the
Lamarck collection of the Muséum d’Histoire Naturelle de Genève proved not to be
the specimen described by Lamarck, and this led me to designate a neotype for H.
unilateralis.
Although I will use only the single genus Haliotis, the taxa used by other
authors are often mentioned. Some authors use several genera in the family
Haliotidae. I consider the use of these genera as unjustified for the following reasons.
Manuscript accepted 08.08.1995.
340 DANIEL GEIGER
1) In the descriptions of the 17 supraspecific taxa (ranked either as subgenera or
genera; see PICKERY 1991 for list and references) only the type species had been
assigned, occasionally with selected species. Of the 200 species level taxa only
approximately 65 have ever been assigned to any supraspecific taxon, and 16 of those
to more than one group (GEIGER, unpublished) demonstrating the problematic
supraspecific taxonomy of this group. The descriptions of these genus-group taxa are
entirely typological and differential diagnoses between them do not exist. 2) Only a
few studies have been using modern systematic methods to determine the relationship
between 17 (enzyme electrophoresis) and 22 (cDNA sequencing) abalone species
(BROWN 1993; LEE & VACQUIER, 1995). The results show that the supraspecific taxa
and the limited number of associated species are not in accordance with the groups
shown in the more modern approaches. However, the type species of Haliotis sensu
stricto, H. asinina Linné, 1758 (by designation: MONTFORT 1810), had not been
included in either study, making any sound taxonomic decisions impossible. 3) The
only inferable consensus in the literature as well as in the abalone community
(Workshop “Evolutionary Biology and Genetics of Abalone” during the Second
International Symposium on Abalone Biology, Fisheries & Culture, February 1994,
Hobart, Tasmania) is not to use any supraspecific taxa until these groupings are based
on data.
ABBREVIATIONS
HUJ Hebrew University Jerusalem
KBIN Royal Belgian Institute for Natural Sciences (Brussel)
LACM Los Angeles County Museum of Natural History
MHNG Muséum d'Histoire Naturelle de Geneve
MNHA Museum of Natural History Amsterdam
MNHL Museum of Natural History Leiden
NMC Nature Museum Coburg
NHML Natural History Museum London (formerly British Museum, Natural
History)
NMW National Museum of Wales, Cardiff
TAU Tel Aviv University
USNM United States National Museum of Natural History
SUPPOSED TYPE SPECIMEN
The text of the original description of LAMARCK (1822) is given here:
11. Haliotide unilatérale. Haliotis unilateralis.
H. testâ ovali, convexo-depressä, rudi, subverrocosä, albido-flavescente, maculis
fuscis pictà; labio elevato, anteriùs latere producto; spira prominulà, obtusa.
An Rumph. Mus. 1.40. fig. G? H?
Habite les mers de Timor et de la Nouvelle-Hollande. Mon cabinet. Bord droit
fort court; nacre peu brillante, Diam. longit., 16 lignes; transv., 11 et demie.
NEOTYPE OF HALIOTIS UNILATERALIS LAMARCK, 1822 34]
The supposed type specimen has been dealt with by MERMOD & BINDER (1963)
who previously had cast doubts about its authenticity. I strongly support their
suspicion, as it is clear from the description that LAMARCK (1822) described a shell
with different characteristics. The specimen is olive green and white in colour
whereas Lamarck described it as white-yellow. It has a rather thick shell for the genus
(Figure 1), but Lamarck indicated it as being thin. The only quantitative indication in
Lamarck’s description is the length and width that he assigned to the shell (36 x 26
mm), which does not correspond exactly to its dimensions (35.5 x 24 mm) (see also
MERMOD & BINDER 1963), but the discrepancy may also be due to the inaccurate
conversion from lines to millimeters. The MHNG has the copy of Lamarck’s
“Histoire des animaux sans vertèbres”, which had been annotated by Lamarck’s
daughter. The annotations consist of handwritten indications of number of specimens
in Lamarck’s collection. For H. unilateralis two specimens are noted, indicating that
some changes occurred during the history of the collection. I identify the supposed
type specimen as H. varia Linné, 1758 and it matches the potential syntypes of H.
varia held in the Linnean Society in London (GEIGER, in prep.). It is now evident that
the original type specimen of H. unilateralis has been replaced by another specimen,
and as no other specimen has been traced in Lamarck’s collection (MERMOD &
BINDER 1963), the original type specimen is assumed to have been lost.
THE IDENTITY of H. unilateralis
SYNONYMS / MISIDENTIFICATIONS:
H. unilateralis Lamarck, 1822: Vol. 6, part. 2, p. 217, no 11.— YARON 1983:
489-491.
“Sanhaliotis cf. pustulata (Reeve, 1846)”.— SHARABATI 1984: PI. 2, figs. 1,
la, 1b.
H. varia Linné, 1758. sensu DRIVAS & JAY 1988: 32, fig. 17, not LINNÉ, 1758.
“A. cf. ovina Gmelin, 1791“. — GEIGER 1991: 95-103, figs. la-d, 2a-d, 3a-d,
4a-d, x (lower specimen).
“A. sp.”.— SINGER 1993: 15.
Non H. unilateralis of KAICHER 1981 [H. varia Linné, 1758].
AUTHORS UNCERTAIN AS TO THE IDENTITY OF H. unilateralis:
WEINKAUFF (1883: 55).— SOWERBY (1887: 37).— PILSBRY (1890: 97). —
MERMOD & BINDER (1963: 148-149).
CHRONOLOGICAL TREATMENT:
REEVE (1846), in his extensive monograph of the genus, did not mention H.
unilateralis, although he discussed other Lamarckian species.
WEINKAUFF (1883) called H. unilateralis a missing species (“verschollene
Art”). His wording subtly alludes to the possible loss of the type specimen. His
342 DANIEL GEIGER
FIG. 1
Supposed type specimen of H. unilateralis held in the MHNG. 35.5 x 24 mm. The specimen is
identified as H. varia. Photographs by C. Ratton, MHNG.
illustration (pl. 6: 4, 5) cannot be identified with certainty, but is closer to H. pustulata
cruenta than H. unilateralis as defined here. He (p. 55-56, 66) synonymised H.
pustulata and H. cruenta with H. unilateralis, and identified H. concinna as a juvenile
H. unilateralis (p. 55, 75). WEINKAUFF (1883: 2) used informal groupings termed
NEOTYPE OF HALIOTIS UNILATERALIS LAMARCK, 1822 343
“Formenkreis” or form groups, because he did not want to use the then described
genera. He assigned H. unilateralis to the “Formenkreis” of H. varia, but did not
synonymise H. varia With H. unilateralis. However, he mentioned that AH. varia
occurs in the Red Sea.
SOWERBY (1887: 37) called H. unilateralis “not identified”, i.e. gave it the status
of a nomen dubium. He compared H. varia with H. concinna, and called the differences
between H. pustulata and H. varia “of questionable specific value” (p. 28).
PILSBRY (1890) corroborated the similarity between H. concinna and H. varia.
He also cited Weinkauff (1883) on the synonymy between H. concinna and H.
unilateralis although he had not seen any specimens of the latter. PILSBRY (1890: pl.
17: 97, 98) had based his somewhat cruder drawing on Weinkauff’s figure. He also
indicated H. varia from the Red Sea.
TALMADGE (1971) first noted the confusion regarding the identity of H.
unilateralis that culminated in PILSBRY (1890), but did not realize that WEINKAUFF
(1883) as well as SOWERBY (1887) gave H. unilateralis the status of a nomen dubium.
He assigned the mistake to Weinkauff “ ... who evidently failed to check on the type
specimens ...’(p. 83). As an alternative the type specimens might have already been
lost at this time, although only subtle supporting evidence can be found in the word-
ing of Weinkauff (see above). TALMADGE (1971) recognized A. unilateralis according
to the supposed type specimen in the MHNG, figured in MERMOD & BINDER (1963).
KAICHER (1981) gave the only modern illustration under the name of H.
unilateralis, which clearly shows a specimen of H. varia.
Apparently only YARON (1983) has used H. unilateralis as defined here (see
below: studied specimens), but he did not illustrate his work.
GEIGER (1991) mentioned a further species of abalone in the Red Sea, which
was found, from the inspection of type material, neither to be H. unilateralis nor H.
pustulata cruenta, and finally was identified as H. cf. ovina Gmelin, 1791. At that
time the status of the supposed type specimen of H. unilateralis in the MHNG had not
been clarified.
In summary, H. unilateralis was called a nomen dubium early on, was com-
pared to H. concinna, which is a synonym of H. varia, and as the latter name was
better known to the scientific community, H. unilateralis was taken as a synonym of
H. varia. This error was compounded by erroneous citations of locations for H. varia
including the Red Sea. In a further complication the supposed type specimen is a
specimen of H. varia.
In modern collections specimens of H. unilateralis as defined here are very
uncommon (TALMADGE 1971; this study). Equally, no old specimens of this species
could be found in the European collections consulted: HUJ, KBIN, MHNG, MNHA
and MNHL (R. PICKERY, pers. comm.), NHML, NMC (W. Korn, pers. comm.) and
NMW, supporting the possibility that the original type specimen had been lost not
long after Lamarck described the taxon. Hence, the biological species may have
become unknown to the scientific community again resulting in confusion about the
identity of Lamarck’s taxon.
344 DANIEL GEIGER
Is it possible to positively identify the species from LAMARCK’s (1822)
description? First, the origin of Lamarck’s material has to be discussed. Lamarck
indicated Timor and Australia (his Nouvelle-Hollande) for the origin of his material.
As he did not collect the specimens himself, an error in these localities might have
been easily, but inadvertently, committed. In a similar case, GRAY (1826) described
H. squamosa from Australia; only recently that species has been relocated in a
restricted area of southern Madagascar (STEWART 1984; pers. comm.; D. PISOR, pers.
comm.). Therefore, an error concerning the geographical provenance of the original
specimens of H. unilateralis should not present an a priori hindrance to the
identification of the species, and biological species from outside the Australian region
should not be excluded from possibly being Lamarck’s H. unilateralis. The taxon has
primarily been reported from the Red Sea (WEINKAUFF 1883; SOWERBY 1887; PILSBRY
1890; HALL & STANDEN 1907; TALMADGE 1971; MERGER & SCHUHMACHER 1974;
YARON 1983: including three additional references; SINGER 1993), but at least some of
these indications may not refer to H. unilateralis as defined here.
LAMARCK (1822) made reference to two figures of RUMPHIUS (1766), but as he
marked them with question marks, these two illustrations are of no use for the correct
identification of H. unilateralis. Unfortunately, Lamarck did not illustrate his work,
and we are left with a short description. However, he mentioned that the form of
species no 12 (H. rugosa) is somewhat similar to the preceding one, 1.e. H. uni-
lateralis (“Forme un peu rapprochée de celle de la précédente”: p. 217). The type
material of H. rugosa contains two separate species (three specimens). HERBERT
(1990) designated a lectotype, and identified the other specimens as H. pustulata.
These three specimens are highly similar in the outline of the shell; therefore, the fact
that Lamarck mixed two species under one name does not impose any problem in
respect of the indicated similarity of form between H. rugosa and H. unilateralis.
The description of H. unilateralis is fairly general, potentially fitting several
species. However, a set of three characters - thin, warty, white yellow - is found in
only two species: the little known H. barbouri Foster, 1946 from Brazil, and the
species known from the Red Sea area as H. unilateralis (YARON 1983; SINGER 1993).
Of the latter, the shape of a typical shell is very similar to that of the type specimens
of H. rugosa in the sense of Lamarck (see above for discussion of this taxon), for
which Lamarck indicated to have a similar shape of the shell to H. unilateralis. The
shell is very thin for the genus, mostly as thin as or even thinner than H. brazieri
Angas, 1869. The shell most often is found in red and orange tones with some
markings in a darker colour: brown and green. The indication of colour is slightly in
contrast with “white and yellow” and “dark spots” by Lamarck. He also describes the
shell as “dull” and having “little shiny nacre”, and, therefore, I agree with WEINKAUFF
(1883), that he described the species from a beach or slightly worn shell. It is well
established that such shells lose the bright colouration and tend to fade to a certain
extent, i.e. a fading from orange to yellow is easily conceivable. One of the most
prominent features of the shell is the elevation and folding near the columella, a
feature not clearly addressed by Lamarck.
NEOTYPE OF HALIOTIS UNILATERALIS LAMARCK, 1822 345
In a statistical analysis, the length and width of the shell indicated in LAMARCK
(1822) lie near the regression line of the same data for all the available specimens
from the Red Sea species, which supports its identification as H. unilateralis (graph
not shown).
I have shown above that the species from the Red Sea is quite likely the same
species on which LAMARCK (1822) had based his description of H. unilateralis. For
the sake of stability of nomenclature (ICZN Article 75(b)) I shall maintain the rather
well established name. However such a decision necessitates the designation of a
neotype, as the original type must be assumed to be lost and the identity of the species
is highly controversial (ICZN Article 75(b)(ii)). The alternative is to designate A.
unilateralis a nomen dubium and to describe the species from the Red Sea as a new
species. However, for the following reasons the designation of a neotype is preferred:
No new taxon is introduced; the already used association of the taxon with a
biological species is retained; and, in contrast to the designation of a neotype, the
declaration of H. unilateralis as a nomen dubium could eventually be overturned,
causing further disruption of taxonomic stability.
The complete specimen designated here as the neotype represents the
specimen that best fits the description of Lamarck, although it is not a perfect match
(cf. ICZN Article 75(d)(4); MAYR & ASHLOCK 1991).
DESIGNATION OF NEOTYPE of H. unilateralis Lamarck, 1822
NEOTYPE: Haliotis unilateralis Lamarck, 1822. MHNG no. 18020. The com-
plete specimen is stored together with Lamarck’s collection. It comprises the dry
shell, the animal in alcohol, and the mounted radula.
TYPE LOCALITY: Red Sea: Gulf of Aqaba: Sinai: Elat. 40 m depth, under
backward slope of reef. Collected by M. Fainzilber, D. Korkos and B. Singer.
DESCRIPTION: Shell of neotype 33.8 mm long and 23.15 mm broad (Figure 2).
Dorsal side bears three spiral rows of bumps, corresponding to depressions in the
nacre. Tremata raised considerably. Last four tremata fully open, fifth partially closed.
Dorsal surface only slightly convex and forms an angle of approximately 70° with
lateral margin of shell. Most distinct character is the plicate margin which itself bears
blunt lamellae. Between fold and columella three spiral ridges. Suture pronounced
and spire distinctly elevated. No scar of hypertrophied adductor muscle.
Colour (based on wet shell; in dry condition colour dulled by periostracum):
Overall bright reddish orange with some light green spots on dorsal side in posterior
part, and at very margin (aperture) near last respiratory hole formed. Several fine,
lighter spiral lines. Red margin intersected from columella up to fold by pairs of
vertical, white bands. Length of intersection between pair of bands 1-2 times as wide
as pair of bands themselves. Nacre uniform and shiny.
Radula: see figure 3 for terminology used. On rachidian tooth, basally, pos-
terior end of basis bears vertical groove and not horizontal one. On lateral tooth 1,
cutting edge has distinct ridge, bent to the posterior. Primary ridge forms an angle of
346 DANIEL GEIGER
Fic. 2
Neotype of Haliotis unilateralis. A) Dorsal. B) Apertural. C) Ventral. D) Anterior part of row
of tremata with folded margin on the left side. A-C: Bar = 10 mm, D: Bar = 5 mm.
approximately 45° to cutting edge. Primary ridge approximately as long as cutting edge.
Secondary ridge separates from main part of tooth in its top third, only slightly shorter
than primary ridge, and very pointed. Cusps of lateral teeth 3-5 have no denticles.
Cusps of inner and middle marginal teeth slightly asymmetrically denticulated, with
denticles on inner margin closer to tip than those of outer margin. Outermost marginal
teeth with far less denticles on outer margin and close to tip (Figure 4).
NEOTYPE OF HALIOTIS UNILATERALIS LAMARCK, 1822 347
denticles on
cutting edge angle primary smooth inner
“2a ha
=.) RN
Dyula aD ta eas
cusp
denticles
= shaft —>
à
_ ros
elongated base base—
Fic. 3
Illustration of the terminology for the teeth of the radula of abalone.
FIG. 4
SEM photographs of raduar teeth of H. unilateralis. A: Rachidian tooth and lateral tooth 1.
Scale bar = 100 um. B: Lateral teeth 3-5. Scale bar = 100 um. C: Inner marginal teeth. Scale bar
= 25 um. D: Middle and outer marginal teeth. Scale bar = 25 um.
Epipodium: Very elaborate, despite rather narrow for genus. Dorsal margin
bears hand-shaped leaflets, several times fingered. On inside of epipodium, palm
gives rise to smaller, less fingered, hand-shaped projections. Longer dorsal tentacles
sparse. Ventral margin instead bears numerous long tentacles and palms of hands
longer than those on dorsal margin, but with fewer secondary hand-shaped leaflets. In
348 DANIEL GEIGER
epipodial fold large tentacles at greater intervals, 1.e. at intervals of 3-5 hand-shaped
structure on dorsal margin. Hand-shaped structures missing on either side, when a
tentacle in epipodial fold.
INTRASPECIFIC VARIABILITY
STUDIED SPECIMENS: The specimens are arranged from their northern-most localities
southward with the collection in brackets and the number of specimens after the colon. Elat 40
m under backward slope of reef (Neotype MHNG no. 18020: 1); Elat, 3. 1994, at foot of reef,
dead, 20 m (Singer: 1); Elat, 30.8. 1988, 10-30 m (Geiger: 1); off Elat: 2 m, on shell, 1972
(Stewart: 6); Elat (Stewart: 1); Elat 30 m (Stewart: 1); 30 km South of Elat, 8. 1993, beach,
dead (Singer: 2); Lagoon of Dahab, 6. 1991, dead on sand (Singer: 1); Dahab, 8. 1993, amongst
corals, 2m, dead (Singer:1); Gulf of Aqaba, approx. 1991, dead, shallow water (Singer: 2);
Na’ama Bay, Sinai (Singer: 1); Gulf of Aqaba, approx. 1988 (Singer: 6); Sinai (Jones: 1); Sinai,
1 m under stone (Geiger: 1); Tiran, Jackson Reef, corals, 30-40 m (Singer: 1); Woody’s Wreck,
2.9.1983 (NHML: 1); 5 mile reef, 8.1.1982 (NHML: 1); Safaga, Egypt (Pickery: 1); Djibuiti:
Ille des 7 frères (Stewart: 1); Djibouti, Ouaramous Island, 6 m rock (Stewart: 1); Aldabra: Ile
Picard, Passé Dubois, 07.04. 1983, Channel margin 3-10 ft (USNM 836532: 1); Mozambique
Channel: Bassas Da India, 04.90 live-taken (Stewart: 1); Djibuiti: Ille des 7 frères (Stewart: 1);
---, 2.9.1982 (NHML: 1). The size of the shells varies between 12.75 and 33.8 mm for their
largest dimension.
The shells mentioned by YARON (1983), housed in the HUJ and the TAU, were
not available for study. However, according to the curator of the HUJ, H.K. Mienis
(pers. comm.), the shells are “identical with the material you received from Solly
Singer”.
Shell: The dorsal surface of the shell can be planar or convex as in H. varia. The
flat shells are so far only known from the Sinai coast of the Gulf of Aqaba. The shells
from this locality are also similar to H. brazieri. The tremata are moderately to highly
elevated, i.e. to a similar extent as in H. parva Linné, 1758 or more, but in none of the
shells seen they are raised as much as in H. brazieri. The shell is thinner or occasionally
as thin as H. brazieri or H. parva. Between the row of tremata and the columella the
folded margin is one of the most noticeable features. It is comparable to the same struc-
ture found in H. scalaris Leach, 1814 [= H. emmae Reeve, 1846], but shows a reduced
range in the extent it is formed. In the most pronounced state it consists of tightly
spaced, delicate but short lamellae, approximately up to 15 between two tremata. These
may become fewer and sturdier and possibly may form nodes on the folded margin.
Colour: The whole range of light rose to orange and red leading occasionally
even to brown tones are found. Some green specimens are known. Most frequently
fine white mottling covers the shell, with larger spots in white or a second colour
being rather rarely encountered. Only two specimens show radial flammae, a pattern
often seen in H. ovina. The nacre is rather of a light colouration for the genus.
Radula and epipodium: The radula (GEIGER, unpublished) and the epipodium
(OWEN et al. 1971; GEIGER, pers. obs.) of abalones shows characters which vary
between species. In the description, only the features different from other species
investigated so far are indicated. Two radulae were investigated with a SEM, and four
alcohol specimens were available. The characters indicated in the description of the
neotype adequately indicate the intraspecific variability.
NEOTYPE OF HALIOTIS UNILATERALIS LAMARCK, 1822 349
Habitat: From all present information the species occurs during day time under
stones and coral heads and in deeper water on the underside of backward slopes of
reefs. No night time observations have been made to date. The vertical range extends
from at least 40 m up to the very shallow sublittoral (1 m). Therefore, in the shallow
water it can be found together with the 7. pustulata cruenta.
Geographic distribution: H. unilateralis has been reported primarily in the Red
Sea. It has been regularly found from the Gulf of Aqaba side of the Sinai from Elat
down to Sharm el Sheik (Singer, pers. comm.; this study). In the southern Red Sea,
SHARABATI (1984) has found the species at the locations called “Woody’s Wreck” and
“5 mile reef”; I was unable to find the precise location of these places. An underwater
photograph of the species was taken at Hurghada (Egypt) by U. Wüest, but the
specimen had not been preserved. Additionally, several specimen from Djibouti, one
specimen from Aldabra (USNM), one from Msibati, Tanzania (K. Stewart, pers.
comm.), one from Mozambique, and at least two specimens from Pointe du sel, St.
Leu, Reunion (M. JAY, pers. comm.: see DRIVAS & JAY 1988) are known. YARON
(1983: 491) indicates without references its occurrence in East Africa. Despite the
more frequent findings of the species in the Red Sea area, the abundance of A.
unilateralis must still be termed infrequent to rare.
COMPARISONS
— H. brazieri: For illustrations see KAICHER (1981), ABBOTT & DANCE (1983:
20), WILSON (1993: pl. 5 fig. 1). The flat form of H. unilateralis from around Elat
somewhat resembles A. brazieri. However, H. brazieri is found only in southeast
Australia from southern Queensland to Jeffries Bay, New South Wales (WILSON
1993). The tremata of H. brazieri are more raised than in H. unilateralis, and the shell
is usually heavier, the spire is much more elevated, the colour pattern usually involves
red oblique flammae, and no spiral row of depressions in the nacre can be observed.
The soft parts of this species are unknown to me.
— H. ovina: For illustrations see SPRINGSTEEN & LEOBRERA (1986: pl. 2 fig. 3),
ABBOTT & DANCE (1983: 22), DHARMA (1988: pl. 1 fig. 4), WILSON (1993: pl. 3 fig.
5). Whereas H. ovina extends as far west as the Gulf of Siam, Andaman Sea
(TALMADGE 1974) and the Maldives (BAER 1989), H. unilateralis has only been found
as far east as Reunion (M. Jay, pers. comm.). The shell of H. ovina is always much
heavier, and the margin between the tremata and the columella does not form a fold as
in H. unilateralis. H. unilateralis can also be distinguished easily from H. ovina on
the basis of the epipodium. The latter species has warty triangles whose broad bases
form the dorsal and ventral margin, and the tips point towards the epipodial fold. The
dorsal and ventral triangles are shifted by half their widths, which causes the tips of
the triangles of one side to point into the gap between the two triangles on the
opposite side. This arrangement forms an undulating epipodial fold. The dorsal and
ventral margin bear many small projections, arranged in a single line. Dorsal to the
dorsal epipodial margin large tentacles are found.
350 DANIEL GEIGER
FIG. 5
Shell of H. pustulata cruenta in dorsal, apertural and ventral view. Sinai, Gulf of Aqaba. 34 x
21.5 mm. Collection Geiger.
— H. p. pustulata and H. pustulata cruenta: For illustrations see KAICHER
(1981), BARASH & DANIN (1992: fig. 9), GIANUZZI-SAVELLI et al. (1994: figs. 105 a-c),
this study (fig. 5). A. pustulata is the only species to occur sympatrically with H.
unilateralis in the Red Sea. Its distribution in the Indopacific extends from the Persian
Gulf (TALMADGE 1974) to northeastern South Africa (JACKS 1983). The two
subspecies can be distinguished from A. unilateralis by having a thicker shell, no
elevated tremata, no folded margin and they usually have a dark brown to sepia
colouration with off-white mottling (Figure 5). The relation between H. pustulata and
H. rugosa Lamarck, 1822 (non Reeve, 1846) is currently debated (cf. HERBERT 1990).
The epipodium has dorsally and ventrally closely packed fingered structures,
with thick fingers. These bundles are separated by spaces bare of any such structures.
NEOTYPE OF HALIOTIS UNILATERALIS LAMARCK, 1822 35]
From within these dense bundles a few longer tentacles emerge. Large, isolated
tentacles appear in the middle of the epipodium, which, however, are not surrounded
by hand-shaped projections. No clear epipodial fold can be identified.
Occasionally H. unilateralis and H. pustulata have been confused, as in the
compilation by VINE (1986). There Sanhaliotis pustulata was described as having
raised tremata, a feature of A. unilateralis and not of H. pustulata cruenta; the second
species discussed is S. varia, which most likely represents H. pustulata cruenta. Note
that VINE (1986: 126) used the genus Sanhaliotis in the description of shells, but
Haliotis in the list on page 172 including the species mentioned in the text.
— H. varia: For illustrations see KAICHER (1981), ABBOTT & DANCE (1983: 22),
SPRINGSTEEN & LEOBRERA (1986: pl. 2 fig. 2), DHARMA (1988: pl. 1 fig. 6), this study
(fig. 1). H. varia seems to have its western distributional limit around Sri Lanka
(TALMADGE 1974; NHML) and Cape Comorin, India (Geiger, unpublished); occa-
sional specimens are found on the Maldives (NHML). One specimen has allegedly
been collected from Muscat, Oman in 1838 (NHML). Therefore, the western distri-
butional limit of the species seems to be very similar to that of H. ovina.
The shell is much heavier, the columella much narrower, and no fold between
the row of tremata and the columella is present. Numerous, narrow, spiral cords are
found on the dorsal surface and show in the nacre; the cords often bear scales. In
some specimens oblique lamellae can be found. The colours are only rarely bright;
muddy green and brown prevails (Caution: do not confuse with H. dohrniana Dunker,
1863).
The epipodium is rather narrow for the genus and is composed of distinct trian-
gular units. Each unit bears a large tentacle on the margin which is bordered on either
side by finer projections, which are often fingered and form small hand shaped struc-
tures, which, however, are less elaborate than in A. unilateralis. In addition, the
secondary palmate structures of H. unilateralis are not found in H. varia. Towards the
distinct undulating epipodial fold, which is formed as in H. ovina, no further struc-
tures can be found; the epipodium is smooth there.
SPECIES OF Haliotis INTHE RED SEA
Seven taxa have been indicated from the Red Sea: H. ancile Reeve, 1846, H.
dringii Reeve 1846, H. pustulata cruenta Reeve, 1846, H. cf. ovina Gmelin, 1791, H.
scutulum Reeve, 1846, H. unilateralis Lamarck, 1822 and H. varia Linné, 1758
(ABBOTT & DANCE 1983; YARON 1983; SHARABATI 1984; GEIGER 1991). YARON
(1983) discusses in detail the various taxa reported from the Red Sea. H. scutulum is a
little known taxon. No specimen labeled with this name could be found in the NHML
(GEIGER, pers. obs. 1993). From the description “waved, here and there larger”, the
colouration “olive brown ... dotted and spotted with green”, and the irregular spiral
ridges with occasional thickenings to be seen in the figure 63 of REEVE (1846), I
tentatively identify H. scutulum as H. varia. \t is mentioned a few times in the lite-
rature from the Red Sea (cf. YARON, 1983), possibly stemming from a confusion of H.
352 DANIEL GEIGER
pustulata With H. varia. Some additional indications on the remaining taxa are worth
mention. H. ancile is noted by ABBOTT & DANCE (1983) to occur in the Gulf of Suez
and the north western Indian Ocean. The figured specimen looks rather like a colour
form of H. pustulata and bears only limited resemblance to the type of H. ancile
(GEIGER, pers. obs. in NHML), which actually is a juvenile H. rubra Leach, 1814 (K.
STEWART, pers. comm.). In his compilation, VINE (1986) includes H. dringii which is
reported so far only from the Australian region (WHITEHEAD 1990). Despite mention
of the seven taxa listed above, only H. pustulata cruenta and H. unilateralis are
known to occur in the Red Sea. The more common H. pustulata cruenta has most
likely been involved if only one species of abalone had been identified, particularly in
the non-taxonomic literature.
In conclusion, only the well known H. pustulata cruenta and H. unilateralis
are known from the Red Sea area. The confusion about the latter biological species
has ceased to exist due to the present designation of a neotype, which does not disrupt
the usage of the taxon, hence provides taxonomic stability.
ACKNOWLEDGMENTS
Benjamin Singer, Daniel Korkos and Mike Fainzilber donated the animal
(neotype), and Mark Jones, Roger Pickery, Katharine Stewart and Urs Wüest made
some of the material available. Kathie Way (NHML) and Alan Kabat (USNM) sent
material for study. Yves Finet and C. Vaucher (MHNG) kindly sent pictures of the
supposed type specimen. R. Guggenheim and the team at the Labor fiir Raster-
elektronenmikroskopie of the University of Basel helped with the initial observations
on radulae. R. Bils and Alicia Thompson (USC) introduced me to the SEM on the
USC campus. Yves Finet, James McLean (LACM), Henk K. Mienis (HUJ), Katharine
Stewart and two anonymous referees read the manuscript and made valuable
comments on it. I express my gratitude to all.
REFERENCES
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BAER, T. 1989: Listes des mollusques gastéropodes des Maldives. Bulletin de la Société
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DHARMA, B. 1988: Siput Dan Kerang Indonesia (Indonesian Shells). PT. Sarana Graha,
Jakarta. 111 pp.
Drivas, J. & M. Jay 1988: Coquillages de La Réunion et de l’île de Maurice. Delachaux et
Niestlé, Neuchatel, 159 pp.
NEOTYPE OF HALIOTIS UNILATERALIS LAMARCK, 1822 353
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performed between the years 1818 and 1822 by Captain Phillip P. King. Appendix B.
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LAMARCK, J. B. 1822: Histoire Naturelle des Animaux sans Vertèbres. T.6(2): 1-232.
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poda): inference from DNA sequences of sperm lysin. Marine Biology 124: 267-278.
LINDBERG, D. R. 1992: Evolution, distribution and systematics of Haliotidae. In: Abalone of the
World: Biology, Fisheries and Culture (S.A. Shepherd, M.J. Tegner & S.A. Guzman del
Pröo, eds.). Fishing News Books, Oxford: 3-18.
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MERGER, H & H. SCHUHMACHER 1974: Morphologie, Ökologie und Zonierung von Koral-
lenriffen bei Aqaba (Golf von Aqaba, Rotes Meer). Helgoländer wissenschaftliche
Meeresuntersuchungen 26: 1-129.
MERMOD, G. & E. BINDER 1963: Les types de la collection Lamarck au Muséum de Genève:
Mollusques vivants. Revue Suisse de Zoologie 70: 127-172.
MONTFORT, D. DE 1810. Conchyologie Systematique 2: 114-120.
Owen, B., J.H. MCLEAN & R.J. MEYER 1971: Hybridization in the eastern Pacific abalones
(Haliotis). Bulletin of the Los Angeles Museum of Natural History and Science No.9: 1-37.
PICKERY, R. 1991: Chronological list of the references to the original descripitions of recent
subgenera and species belonging to the family Haliotidae Gloria Maris 29: 105-118.
PILSBRY, H.A. 1890: Manual of Conchology; structural and systematic with illustrations of the
species. 12: 72-127. pls. 1, 3-24, 46-50.
REEVE, L. 1846: Monograph of the genus Haliotis. 22 pp., pls. 1-17.
RUMPHIUS, G.E. 1766: Ambonische Raritätenkammer. Wien. 200 pp., 49 pls.
SHARABATI, D. 1984: Red Sea Shells. KP/, London, 127 pp.
SINGER, B.S. 1993: Taxonomic confusion in Red Sea Haliotis. Levantina 77-79: 15.
SOWERBY, G.B. 1887: Thesaurus Conchyliorum. 5: 17-37, pls. 1-14.
SPRINGSTEEN, F.J. & LEOBRERA, F.M. 1986: Shell of the Philippines. Carfel Seashell Museum,
Manila. 377 pp.
STEWART, K. 1984: Notes on H. squamosa Gray, 1827. Shells and Sea Life 16(7): 92-95.
TALMADGE, R.R. 1971: Notes on Israeli Haliotids. Argamon 2: 81-85.
TALMADGE, R.R. 1974: Some notes on the distribution and Taxonomy of Haliotis. Veliger 16:
411-412.
VINE, P. 1986: Red Sea Invertebrates. /mmel Publishing, London, 224 pp.
354 DANIEL GEIGER
WEINKAUFF, H.C. 1883: Die Gattung Haliotis. In: Systematisches Conchylien-Cabinet von
Martini and Chemnitz. 2(6)B: 1-83, pls. 1-30.
WHITEHEAD, T. 1990: Haliotis astricta and Haliotis dringii. Australian Shell News 70: 3.
WILSON, B. 1993: Australian Marine Shells 1. Odyssey Publishing, Kallaro, 408 pp.
YARON, I. 1983: The Haliotidae of the Red Sea. Levantina 42: 486-493.
Note added in proof.
The shells shown by Pickery & Steppe (1995: Gloria Maris 34: pl. 5, fig. 7) as H. unilateralis
are H. pustulata cruenta. Four additional specimens of H. unilateralis were located in the
California Academy of Sciences in San Francisco (CASIZ). Zanzibar: Pinna (CASIZ 1594: 1):
Mauritius, leg. Boswell (CASIZ 1532: 3).
REVUE SUISSE DE ZOOLOGIE, 103 (2): 355-367; juin 1996
Un Evalljapygidae (Diplura) canadien: Evalljapyx saundersi n. sp. —
Dicellurata Genavensia XXI -.
Jean PAGES!
51, rue du Faubourg Saint-Martin, F-21121 Fontaine-lès-Dijon, France.
A Canadian Evalljapygidae (Diplura): Evalljapyx saundersi n. sp.
Dicellurata Genavensia XXI -. Evalljapyx saundersi n. sp. is described,
based on an adult & from Vancouver (B.C.). The E. sonoranus Silv. (=
hubbardi [Cook]) recorded from Vancouver Island by SAUNDERS (1946) is
certainly a missidentification, hubbardi beeing localized in Arizona and all
the other species of Evalljapyx are until now only known from the South-
Western States of USA, the Equator and some Caribbean Islands. The
author is convinced that SAUNDERS' specimen belong to the new species
which is close to E. ombris Smith from California. Four chetotaxic patterns
are proposed for the seven first abdominal segments.
Key-words: Diplura - Evalljapygidae - Canada - Taxonomy - New species.
INTRODUCTION
En 1911 SILVESTRI crée le genre Evalljapyx pour des Japyx caractérisés par
leurs phanères pourvus de barbules et leurs mandibules d'un type particulier. Depuis
cette date 33 espèces en ont été décrites; toutes sont localisées du Sud-Ouest des
Etats-Unis (Californie, Nevada, Arizona, Texas), a l'Equateur et aux Caraïbes:
SILVESTRI (1948b) crée la sous-famille des Evalljapyginae que j'ai élevée au rang de
famille (PAGES 1989).
Or, en 1946 L.G. SAUNDERS rapporte la capture "at Brentwood, at the head of
the Saanish Inlet on Vancouver Island" de 3 "Japygids" qui, d'après la documentation
disponible à l'époque, ne pouvaient être que des E. hubbardi (Cook, 1899) (= E.
sonoranus Silv., 1911), seule espèce alors connue ayant 30 articles antennaires.
En fait, E. hubbardi n'est signalé que de l'Arizona et sa présence au Canada est
pour le moins surprenante, mais ni SILVESTRI (1947), ni SMITH (1960a) ne relèvent
cette anomalie chorologique et REDDELL (1983) se contente d'écrire que "the record of
this species for Canada... is almost certainly based on a misidentification".
| Professeur émérite de l'Université de Bourgogne, Equipe d'Ecologie et Dynamique
des Populations, F-21100 Dijon.
Manuscrit accepté le 17.08.1995.
356 JEAN PAGÉS
Cependant la présence d'un Evalljapygidé dans cette partie du Canada ne peut
être totalement exclue puisque un cas identique est connu chez les Japygidés: Occas-
Japyx americanus (Mac Gillivray, 1893) a été décrit d'Olympia (Etat de Washington)
et signalé du "Canada occidentalis" par SILVESTRI (1948a); deux autres espèces de ce
genre sont localisées en Californie et une dans l'Arkansas.
Grâce à la capture d'un d d'Evalljapyx indubitable à Vancouver par le Dr I.
Löbl, il m'est possible de confirmer la présence d'Evalljapygidae au Canada, tout au
moins dans cette région très restreinte et très particulière. Il s'agit d'une espèce
nouvelle que je dédie à L.G. Saunders, car je crois que les spécimens qu'il a recoltés
appartiennent à cette n. sp. qui n'a apparemment que de lointains rapports avec E.
hubbardi si l'on ne tient compte que de la forme des cerques par exemple; le fait que
ses exemplaires aient 30 articles antennaires et E. saundersi 29 seulement n'est pas un
obstacle à ce rapprochement, les espèces gravitant autour d'E. hubbardi pouvant avoir
de 29 à 31 articles aux antennes (SMITH 1960).
Dans cette note outre la description de la nouvelle espèce, je propose des
modèles chétotaxiques pour les 7 premiers segments abdominaux, car je crois qu'une
étude précise et comparative de la chétotaxie pourrait aider à une meilleure compré-
hension des affinités des espèces de ce genre, comme l'écrit REDDELL (1983): "A
thorough review of the genus may indicate it should be split into two or more genera".
Je remercie Mme M. Krähenbühl d'avoir dactylographié ce manuscrit et Mlle
F. Marteau qui a reporté sur calque mes dessins originaux.
MODELES CHÉTOTAXIQUES
A ma connaissance aucun des auteurs ayant décrit des Evalljapyx ne s'est
préoccupé de définir de façon précise des modèles chétotaxiques que l'on pourrait
considérer comme typiques des différents sclérites pris en compte dans les des-
criptions. Silvestri (op. cit.), quant à lui, n'indique qu'un nombre de "setis sat longis",
nombre souvent précédé de "circa"; quant aux figures qu'il donne, presque unique-
ment des derniers tergites abdominaux (du 6ème au 10ème au mieux) et des uroster-
nites 1 et 2, leur faible dimension et le manque presque constant de points de repère
rendent leur utilisation trés difficile. Smith qui est sans aucun doute le spécialiste qui
a pu étudier le plus grand nombre d'Evalljapyx (plus de 2000!), n'a jamais publié un
dessin précis d'un sclérite quelconque.
En 3 occasions, Smith donne des schémas trés simplifiés et sans point de
repère: en 1959b celui de l'urosternite 1 d'E. helferi Smith et du quart postérieur
gauche de celui de la 2 et des d de 3 espèces voisines (raneyi Smith, facetus Smith,
diversipleura Silv.); en 1960b il publie des dessins très simplifiés du mésonotum et de
la moitié droite du tergite 7 de 3 nouvelles espèces (leechi, decorus et mckenziei);
enfin en 1961, lorsqu'il compare les urosternites 1 de la 9 et du "second instar" d'E.
facetus Smith.
Il est certain que chez les Japygoidea la chétotaxie ne joue pas un rôle aussi
fondamental que chez les Rhabdura (Campodeoidea et Projapygoidea), mais elle
UN EVALLJAPYGIDAE (DIPLURA) CANADIEN 357
Schémas des modèles chétotaxiques proposés pour les 7 premiers segments abdominaux des
Evalljapyx Silv.. —
A. — Tergite abdominal 1; B. — Tergites abdominaux 2 à 7; C. — Sternite 1, les organes sub-
coxaux ont été représentés très schématiquement et très arbitrairement quant aux nombres de
soies glandulaires et sensorielles, ces dernières sont barbelées chez toutes les espèces connues:
D. — Sternites 2 à 7.
Sur tous les schémas les macrochètes ne sont représentés que par le contour de leurs embases;
le L;, des sternites 2 à 7 a son embase colorée en noir pour indiquer que ce M est
supplémentaire par rapport à ceux du sternite 1. Le rond noir entre Lg et Lo figure le sm typique
des sternites 1 à 7.
358 JEAN PAGÉS
permet très souvent de distinguer et de regrouper avec plus de précision les espèces de
genres dits "difficiles" comme par exemple Metajapyx Silv., Catajapyx Silv., Indjapyx
Silv. ou même Parajapyx Silv.; elle peut aussi apporter un complément important à la
diagnose de quelques genres comme Holjapyx Silv., Abjapyx Silv., etc.
Les 33 espèces d'Evalljapyx actuellement décrites constituent en première
analyse, un ensemble hétérogène: il est bien difficile d'après les seules descriptions,
souvent trop peu précises ou incomplètes, de reconnaître les affinités des espèces
entre elles. La solution adoptée par SMITH de les réunir d'après le nombre d'articles
antennaires me paraît avant tout pratique, mais peu phylogénique, compte tenu de ce
que l'on peut constater chez les autres Japygoidea. Par exemple SMITH (1960b) décrit
3 espèces (leechi, decorus, mckenziei) qui s'écarteraient des autres Evalljapyx par
leurs antennes de 26 articles; 1l ne cite pas cubanus et inermis, décrits par SILVESTRI
(1929) de Cuba qui ont le même nombre d'articles antennaires; or ces deux espèces
présentent au lobe interne des maxilles des lames dont la première est entière, les 4
suivantes pectinees; ceci en contradiction avec la définition du genre par SILVESTRI
(1911) ou SMITH (1959b, 1960a) qui indiquent 5 lames pectinées à ce lobe maxillaire.
Quatre lames pectinées seulement se retrouvent dans la sous-famille des Pro-
valljapyginae, dont c'est un des caractères principaux, proposée par SMITH (1959a,
1962). Par ailleurs SMITH (1961) écrit que le "second instar" d'E. facetus montre 1
lame entière et 4 autres pectinées à ce lobe; si l'on tient compte d'une part de la sim-
plicité de l'armature des cerques de ce stade qui se rapproche, par exemple, de celle de
Nanojapyx coalingae Smith et, d'autre part, des caractéristiques du genre Eojapyx
Smith (1960c), on peut concevoir que la sous-famille des Provalljapyginae, appa-
remment hétérogène, est plus proche des Evalljapyx qu'il n'y paraît et qu'elle est à
mon avis moins primitive que ne l'affirme SMITH (1959a).
Je crois qu'une comparaison très précise des chétotaxies aurait fourni des
points de repères phylogéniques importants que les descriptions des auteurs ne
permettent pas de détecter actuellement. Le manque de matériel et de documents
iconographiques utilisables me fait renoncer à définir une chétotaxie typique de la
tête, du thorax et des 3 derniers segments abdominaux; par contre les renseignements
fournis par les dessins de Silvestri (op. cit.) et mes observations sur l'unique
exemplaire en ma possession me semblent permettre de proposer des modèles
chétotaxiques applicables aux 7 premières plaques tergales et sternales, indiquant la
position idéale des seuls M, ne pouvant déterminer à coup sûr parmi les autres pha-
nères ceux susceptibles d'être des sm? ou de simples soies. Ces modèles corres-
pondent à un stade moyen du développement postembryonnaire et devront être affinés
ou même corrigés par l'étude de nombreux spécimens de tous stades; en effet, les
Evalljapyx montrent, d'après Smith (op. cit.), une certaine variabilité de la chétotaxie
liée au sexe et au stade de développement et l'on sait que chez les Japygidés la taille et
la densité des phanères peuvent, selon les lignées, se simplifier ou au contraire se
compliquer au cours des mues successives.
2 On trouvera la liste des abréviations et la définition des rapports dans PAGES (1954) et
PAGÉS & SCHOWING (1958).
UN EVALLJAPYGIDAE (DIPLURA) CANADIEN 359
Fics 1-5
Evalljapyx saundersi n. sp., 4 holotype. — 1. Vertex, e = 392 pm. — 2. Pronotum, e = 392 um. —
3. Mésonotum, e = 517 um. — 4. Métanotum, e = 517 pm. — 5. PIII droite, face antérieure, e =
417 um.
Quelques explications sont nécessaires pour justifier le systeme de numéro-
tation des M. Je n'ai pas considéré la répartition de ces phanères suivant des rangées
transversales, comme le propose Smith (1959b, 1961), mais en tenant compte du fait
que ces plaques sont constituées de 3 parties distinctes; c'est ainsi que tergalement 2
apodèmes longitudinaux, plus ou moins marqués, isolent le tergite proprement dit des
2 paratergites et que sternalement les 2 apodèmes stylaires du coxosternite séparent
une aire centrale de 2 aires latérales.
L'analyse de ces modèles chétotaxiques m'amène à faire les commentaires
suivants:
1. Comme chez les Japygidés, la plaque tergale du premier urite (modèle A) a
une chétotaxie très simplifiée par rapport à celle des segments suivants; elle est aussi la
seule à présenter un prétergite (= précosta, PAGÉS 1989) pourvu de phanères dont
typiquement 1+1 M; le macrochète médian A paraît propre à saundersi, le seul autre
Evalljapyx à présenter des M médians serait, d'après la littérature, E. leechi Smith,
1960b.
360 JEAN PAGÉS
2. Contrairement aux Japygidés la seconde plaque tergale (modèle B) a une
chétotaxie typique identique à celle des 5 plaques suivantes, mais quelques exceptions
semblent exister pour autant que l'on puisse en juger d'après les descriptions (darling-
toni Silv., anombris Smith); la précosta, entière ou formée de 2 sclérites ovalaires, est
toujours dépourvue de phanères.
3. Pour les plaques sternales (modèles C et D), j'ai considéré que les pré-
sternites (= apotomes auct., PAGÉS, loc. cit.) étaient ornés typiquement de 6+6 M mais
ce nombre est très variable, comme chez saundersi par exemple, et un M médian
semble assez fréquent; chez les Japygidés, sauf exceptions rarissimes, ces présternites
sont nus.
4. L'aire centrale du coxosternite | est remarquable par l'absence de M entre la
série M4-M et les M précédant les organes subcoxaux arbitrairement fixés à 6+6 sur
le modèle C; ce territoire correspond à peu près exactement à celui occupé chez les
Japygidés par les macrochètes de la série C et presque toujours par des soies à embase
circulaire caractéristique.
5. Les aires latérales 1 à 7 présentent toutes la même chétotaxie si l'on fait
abstraction du Li, des coxosternites 2 à 7; on notera plus particulièrement les 4
phanères insérés entre chaque style et le bord externe de l'aire, au moins 2 sont tou-
jours des M (Lg et Lio) alors que le Lo peut être un sm, comme celui inséré entre Lg et
Lo qui est chez toutes les espèces un sm.
II. Evalljapyx saundersi n. sp.
Matériel étudié: Canada. British Columbia: Vancouver, Stanley Park, niveau
de la mer; 7.V11.88; forêt tempérée hygrophile. Tamisage de feuilles mortes sur sol marécageux
dans la partie centrale. I. Löbl leg.: holotype, d de 9,6 mm, déposé dans les collections du
Muséum d'Histoire naturelle de Genève.
TÊTE
Aussi large que longue.
Vertex: sans sclérotisations particulières entre les bases des antennes;
20+20 M longs, pourvus de 5-7 barbules nettes; on peut les répartir sur chaque moitié
du vertex de la façon suivante: 2 le long de la base de l'antenne, les 18 autres dis-
posées sur 5 rangées longitudinales de 3, 2, 4, 5 et 4 M en partant de la ligne médiane;
3+3 sm assez longs ou assez courts, dont | entre les 2 M basiantennaires, | entre les M
de la seconde rangée longitudinale et 1 entre les 2ème et 3ème M de la rangée la plus
latérale.
Antennes: de 29 articles; des soies à barbules très fines sur les articles 1 à
3 inclus, au-delà tous les phanères sont lisses: 2 verticilles de soies droites et de sen-
silles recourbées jusqu'au 13ème article, puis au moins 3 à partir du 14ème, le
proximal ne comportant normalement que des sensilles recourbées; on observe dans le
verticille proximal des articles 4-13 et dans le verticille intermédiaire des articles
14-29, des soies droites dirigées vers la base de l'antenne; aires pileuses indistinctes;
les 13 trichobothries typiques ne dépassant pas la longueur des soies droites des
articles qui les portent, a médiane comme chez les autres Evalljapyx; 6 sensilles
UN EVALLJAPYGIDAE (DIPLURA) CANADIEN 361
FIGs 6-7
Evalljapyx saundersi n. sp., 4 holotype. — 6. Tergites 7 à 10 et les cerques, e = 517 um. — 7.
Détail des marges internes des cerques, face tergale, e = 82 um.
362 JEAN PAGÉS
placoïdes disposés en 2 groupes linéaires de 3 formant un anneau pratiquement
continu comme c'est la règle chez les Evalljapygidés.
Pieces buccales: typiques du genre, les 5 lames du lobe interne des
maxilles pectinées, la première presque aussi longue que les 4 suivantes et pourvue
d'environ 8-10 denticulations nettes; palpes labiaux 2,5 fois aussi longs que larges à
la base, chacun pourvu de 9 soies dont les subapicales sont en moyenne 1,5 fois aussi
longues que le palpe qui les porte, la plus externe 1,2 fois plus longue que l'interne.
THORAX
Pronotum: 6+6 M pourvus de 5 à 7 barbules, les 1+1 antérieurs
submédians sont assez longs, leur longueur égale environ 1,25 fois l'intervalle sépa-
rant leurs embases; les sublatéraux subantérieurs sont près de 2 fois aussi longs que
les M submédians antérieurs ou postérieurs; 3+3 sm assez courts, dont 1+1 sub-
antérieurs et 2+2 subpostérieurs.
Méso- et ménatotum: Préscutum: 1+1 M assez longs à 5-7 barbules.
0-1+0-1 minuscules soies de part et d'autre des M. Scutum: 9+9 M à 5-7 barbules,
ceux situés près de la ligne médiane et de part et d'autre des apodèmes longitudinaux
assez courts; les plus longs sont les 1+1 sublatéraux subintermédiaires; 3-4+3-4 soies
simples (sm?) assez longues à très courtes.
Pattes: peu allongées, les PIIT atteignant à peine le milieu du second
urosternite; la face antérieure (= externe) du tibia et du tarse est pourvue d'un grand
nombre de M, de soies barbelées et de soies simples, souvent aussi longues que les M
les plus développés: on notera à la base du tarse, face dorsale, un phanère coudé
présent à toutes les pattes: face postérieure, à l'apex du fémur, on observe 3 phanères
dont 2 barbelés et un simple comme par exemple chez E. anombris Smith (SMITH
1960a, fig. 8); aux PIII le tarse égale un peu moins des 2/3 de la longueur du tibia; pas
de soies spiniformes sur la face sternale des tarses, mais 8-10 longues soies simples;
griffes subégales, l'interne légèrement plus courte que l'externe; elles mesurent un peu
plus du tiers de la longueur du tarse; unguiculus aigu, égalant un peu moins du tiers de
la longueur des griffes.
Fics 8-21
Evalljapyx saundersi n. sp., 4 holotype. — 8. Urosternite 1, les organes subcoxaux latéraux
évaginés en partie, e = 209 um. — 9. Soie sensorielle et soies glandulaires d'un organe subcoxal
latéral, e = 42 um. — 10. Urosternite 3, e = 488 um. — 11. Pleurites 2 gauches, e = 488 pm. — 12.
Pleurites 3 droits, e = 517 pm. — 13. Style 4 droit, face antérieure, e = 110 um. — 14. Style 1
droit, face latérale externe, e = 110 pm. — 15. Urosternite et paratergites 8, e = 517 um. — 16.
Sternopleurites 9, e = 517 pm. — 17. Urite 7, vue latérale gauche, e = 488 um. — 18. Urite 8, vue
latérale gauche de sa marge postérieure, e = 517 um. — 19. Urite 9, vue latérale gauche de sa
marge postérieure, e = 517 um. — 20. Papille génitale d , vue latérale gauche, e = 251 um. — 21.
Papille génitale 3, vue antérieure, tous les phanères ne sont pas représentés, e = 251 um.
363
364 JEAN PAGÉS
ABDOMEN
Tergites 1 a 6: conformes aux modèles chétotaxiques proposés; on
relève les quelques irrégularités suivantes: l'absence des Ls et des L¢ au tergite 6, 1 M
supplémentaire près du Z4 droit du tergite 3, 1 M supplémentaire entre les L; et Lo
gauches des tergites 4 et 5 et 1 M supplémentaire entre les Ly et L4, à gauche au
tergite 4 et à droite au 5ème.
Tergite 7: à chétotaxie typique, on notera qu'il présente une sorte de
carène fortement colorée allant en biais de l'angle antérieur, passant entre Lj et Lo et
se terminant pratiquement sur le bord latéral substernal à hauteur de Lo.
Tergite 8: environ | fois 1/5 aussi large que long; 8+8 M dont 1+1
médians antérieurs, 1+1 submédians postérieurs, 1+1 antérieurs sublatéraux, 1+1
latéraux postérieurs et 4+4 latéraux (antérieurs, intermédiaires, postérieurs); 1+1 sm
entre les 2 M les plus postérieurs.
Tergite 9: 2,7 fois aussi large que long; 3+3 M le long du bord postérieur
dont 2+2 latéraux.
Tergite 10: à bords latéraux rectilignes, légèrement convergents vers
l'arrière; 1 fois un tiers aussi long que large au niveau des M latéraux subantérieurs;
carènes bien marquées, fortement colorées, abrégées à leurs deux extrémités; 5+1+5
M avec 6-8 barbules, le médian court, les autres longs ou assez longs; 3+3 sm assez
longs ou longs, lisses, dont 1+1 latéraux subpostérieurs, 1+1 insérés au tiers antérieur
des carènes, 1+1 submédians subpostérieurs, les plus développés: 2+2 soies simples
courtes dont 1+1 latérales subintermédiaires et 1+1 près de l'extrémité postérieure des
carènes. Sur tous les tergites, de très nombreux microchètes, dont un tout petit nombre
a été représenté.
Angles latéraux postérieurs des tergites: ne présentent
rien de particulier; tout au plus, on notera la forme arrondie des angles des para-
tergites 7 et des tergites 8 et 9.
Acropyge: peu saillant, large, régulièrement arrondi.
Longuweurs relatives des segments 7 a 10-ajene compaq
rerai ici que les longueurs médianes des parties découvertes de ces segments, le
dixième étant, par définition, égal a 100: 91-83-25-100.
Sternite 1: Préscutum: 6+1+6 M, les plus latéraux les plus longs, les
5+1+5 autres assez courts. Scutum: 25+26 M dont les 23+23 typiques, assez courts
sur l'aire centrale, longs ou assez longs sur les aires latérales: 2+3 M supplémentaires,
dont | près des Lg gauches.
Organes suboxaux latéraux: ils sont très développés occupant
presque toute la largeur interstylaire comme c'est la règle à quelques exceptions près,
par exemple E. crassicauda Silv., E aguayoi Silv., tous deux de Cuba et peut-être E.
duricauda Ewing & Fox du Guatemala pour autant que les descriptions nous
permettent de juger; ils ne sont pas précédés d'une rangée de soies barbelées très
courtes; 4 à 5 rangées très touffues de soies glandulaires, très courtes, SG/st; = 0,15
en moyenne; elles sont abouchées à un canal glandulaire au moins 5 fois aussi long
que la soie dont il est issu et terminé par un réservoir piriforme muni d'un très court
UN EVALLJAPYGIDAE (DIPLURA) CANADIEN 365
goulot; 23 soies sensorielles très densément barbelées, environ 5 fois aussi longues
que les soies glandulaires SS/SG = 4,8-5,2, SS/st; = 0,75.
Organe glandulaire médian: inexistant.
Sternites 2 à 7: Préscutum: 7+7 M au sternite 2, 7+1+7 au 3ème, 6+1
+6 au 4ème, 6+6 aux 5ème et 6ème, 5+5 au 7ème. Scutum: les 25+25 M typiques; on
note chez notre exemplaire les anomalies suivantes: au sternite 2, absence du Ms
gauche et du M» droit, présence d'un M supplémentaire de chaque côté entre M, et Ms
ainsi qu'entre Mg et M,, à gauche; au sternite 3 présence d'un M supplémentaire à
droite de la ligne médiane entre Mg et M;,; absence du My droit au Sème sternite et du
L4 au 6ème: 1 M supplémentaire à gauche au 4ème entre M> et My et | à droite entre
Le et L7 au 5ème.
Sternite 3 d: il montre juste en arrière de la limite du préscutum une
structure équivalente aux fossettes glandulaires des Japygidés. On observe un oper-
cule constitué de 2 pièces triangulaires occupant environ la moitié de la largeur du
sternite, pourvues chacune d'une demi-douzaine de minuscules soies; elles recouvrent
une sorte de poche s'étendant jusque dans le segment précédent et recouverte de cen-
taines de soies barbelées très courtes; chez saundersi, elle est identique au "setose
sac” décrit par SMITH (1960b) chez son E. ombris ou à celle notée par SILVESTRI
(1947) chez son E. dispar.
Sternite 8:10+11 M disposés sur 3 rangées longitudinales de 5, 4 et 1
(postérieur) M, le M supplémentaire à droite est subantérieur entre les 2 rangées de 4
et 5 M; une carène peu élevée, mais très colorée court entre la rangée de 5 M et la
limite latérale du sclérite.
Paratergites 8 :6 M longs à 5-7 barbules.
Sternopleurites 9: contigus sur la ligne médiane; 4-5 M longs.
Sternite 10: 10+10 M normaux.
Vésicules exsertiles: typiques aux urosternites | à 7, relativement
petites occupant moins du 1/10ème de la largeur interstylaire, celles de l'urosternite 1
très difficiles à observer.
Styles: allongés, aigus, à cône secondaire très aigu, bien séparé du cône
principal; 2 pores énigmatiques, petits; 2 soies, la plus développée 3 fois plus longue
que la petite qui doit être une sensille sétiforme; s1/5t1 = 0,36, sı/s7 = 0,84, st1/st7 =
0,71, s1/st7 — 023:
Pleurites: SMITH attache une grande importance à la chétotaxie des 2
sclérites présents dans la membrane pleurale; il nomme le sclérite antérieur "pleurite"
et le postérieur "pleuron" (SMITH 1959b et 1960a), mais en 1960b, il parle de "pré-
pleurite et de pleurite", ce dernier terme étant synonyme du pleuron; j'adopterai ici
cette seconde nomenclature en rappelant que, pour moi, le "prépleurite" est préseg-
mentaire et que le "pleurite" est homologue à un sternopleurite (PAGES 1989). D'après
mes observations et les données que l'on peut trouver dans la littérature, il apparait
que la chétotaxie du pleurite présente presque toujours 2 M et 1 soie simple posté-
rieure, exception: par exemple macswaini Smith, hubbardi (O.F. Cook) (= sonoranus
Silv.); c'est le prépleurite qui montre le plus de variations; chez saundersi, on observe
366 JEAN PAGÉS
les chétotaxies suivantes des prépleurites 1 à 7: 1 sans phanère; 2 avec 4 M du côté
droit de l'urite et 3 M à gauche; 3 avec 3 M à droite et 2 à gauche; 4 avec 2 M à droite
et 3 à gauche: 5 avec 2 M à droite et 3 à gauche: 6 avec 2 M à droite comme à gauche;
7 avec 1 M des 2 côtés; il semble donc que tout au moins chez saundersi la chétotaxie
des sclérites pleuraux n'ait pas la valeur que lui attribue SMITH (1960a) même s'il
avertit que "several ... pleura should be examined to establish the true number of these
setae”; on observe entre les 2 M normaux du pleurite 7 une sorte de carène, peu éle-
vée, très colorée.
Papille génitale d: aucune représentation n'en a été faite et je n'en ai
pas fait mention dans ma note de 1989; SMITH (1959b) décrit celle de E. helferi, mais
peu compréhensible sans figuration; chez saundersi, cette papille comporte une base
assez large et élevée, pourvue de soies simples assez longues latéralement face
antérieure et médialement quelques soies courtes; l'orifice génital est situé au sommet
d'un tronc de cône, assez court, occupant environ le 1/4 de la largeur de la base,
pourvu de nombreuses soies disposées sur 2 verticilles, les proximales courtes, les
distales très courtes; pas d'appendices génitaux; ce type de papille d me semble
corroborer encore plus la séparation entre Evalljapygidés et Japygidés; aucun autre
type de Diploures ne présente une papille d de ce genre.
CERQUES
Nettement plus courts que la partie normalement découverte du tergite 10,
Leq/L10 déc = 0,85; environ 2-3 fois aussi longs que larges à la base.
Cerque droit: une dent peu saillante à sommet arrondi, nettement post-
médiane, rg = 2,5; la largeur au niveau de la dent égale environ les 3/4 de la largeur à
la base; marge prédentale rectiligne avec 3 tubercules basaux supérieurs arrondis et
largement espacés l'un de l'autre; la marge inférieure montre près de la base du cerque
3-4 saillies arrondies, puis 1 tubercule arrondi suivi de 3 autres équilatéraux aigus; le
reste de la marge présente avant la dent 6-7 ondulations peu saillantes; la marge
postdentale dessine un profond sinus pourvu de 6 denticules arrondis peu saillants.
Cerque gauche: une forte dent à sommet aigu dirigé vers la base du
cerque, nettement prémédiane, rz = 0,57; marge prédentale avec 2 tubercules arrondis
largement séparés; marge postdentale régulièrement concave avec 2 rangées de den-
ticules peu saillants; le proximal inférieur aigu, tous ceux qui lui font suite arrondis;
ceux de la rangée supérieure ne sont que de simples ondulations.
Chétotaxie: tous les phanères sont lisses, sauf, sur chaque cerque, un M
latéral subantérieur pourvu de 6-8 barbules.
AFFINITÉS
Cette espèce est bien caractérisée par ses antennes, ses sternites 1 et 3 et ses
cerques; elle se rapproche beaucoup de E. ombris Smith, espèce propre, elle aussi, à
des biotopes humides et est bien distincte de E. hubbardi (Cook) par ses cerques et la
poche piligère du sternite 3 du d.
UN EVALLJAPYGIDAE (DIPLURA) CANADIEN 367
BIBLIOGRAPHIE
ALLEN, R.T. 1988. A new species of Occasjapyx from the Interior Highlands (Insecta: Diplura:
Japygidae). Proceedings Arkansas Academy of Science 42: 22-23.
Cook, O.F. 1899. New Dicellura. Proceedings of the entomological Society of Washington 4:
222929:
MACGILLIVRAY, A.D. 1893. North American Thysanura.— II. The Canadian Entomologist 25:
173-174.
PAGES, J. 1954. Japyginae (Japygidae, Insecta Diplura) de la Yougoslavie et des régions limi-
trophes. Glasnik Prirodnjackog muzeja srpske zemlje (= Bulletin du Museum
d'Histoire naturelle du Pays serbe) 5-6: 235-264.
PAGES, J. 1989. Sclérites et appendices de l'abdomen des Diploures (Insecta Apterygota).
Archives des Sciences, Genève 42: 509-551.
PAGES, J. & J. SCHOWING. 1958 Diploures Japygidés du Kivu et de l'Urundi (Congo belge).
Revue de Zoologie et de Botanique africaines 57: 193-240.
REDDEL, J.R. 1983. A checklist and bibliography of the Japygoidea (Insecta: Diplura) of North
America, Central America, and the West Indies. Texas memorial Museum, Pearce-
Sellards Series 37: 41 pp.
SAUNDERS, L.G. 1946. A Canadian Japygid (Thysanura). The Canadian Entomologist 78: 95.
SILVESTRI, F. 1911. Materiali per lo studio dei Tisanuri, XII. Un novo genere e undici specie
nove di Japygidae dell'America settentrionale. Bolletino del Laboratorio di Zoologia
generale e agraria della R. Scuola superiore d’ Agricoltura in Portici 5: 72-87.
SILVESTRI, F. 1929. Contribuzione alla conoscenza degli Japygidae (Thysanura) di Cuba.
Bolletino del Laboratorio di Zoologia generale e agraria del R. Istituto superiore
agrario, Portici 22: 263-281.
SILVESTRI, F. 1947. On some Japygidae in the Museum of Comparative Zoölogy (Dicellura).
Psyche, a Journal of Entomology, Cambridge entomological Club 54: 209-229.
SILVESTRI, F. 1948a. Descrizioni di alcuni Japyginae (Insecta Diplura) del Nord America. Boll.
Bolletino del Laboratorio di Entomologia agraria di Portici 8: 118-136.
SILVESTRI, F. 1948b. Specie di Japygidae (Insecta Diplura) finora raccolti nel Messico. Boll.
Bolletino del Laboratorio di Entomologia agraria di Portici 8: 297-320.
SMITH, L.M. 1959a. The Japygidae of North America 1 — Provalljapyginae and Nanojapyx.
The Pan-Pacific Entomologist 35: 99-107.
SMITH, L.M. 1959b. Japygidae of North America, 4. New species of Evalljapyx with twenty-
four antennal segments (Diplura). Proceedings of the entomological Society of Wash-
ington 61: 267-274.
SMITH, L.M. 1960a. Japygidae of North America, 5. Species of Evalljapyx with 30+1 segments
in the antenna (Order Diplura). Annals of the entomological Society of America 53:
137-143.
SMITH, L.M. 1960b. Japygidae of North America, 6. New species of Evalljapyx with 26
antennal segments. Journal of the Kansas entomological Society 33: 1-6.
SMITH, L.M. 1960c. Japygidae of North America, 7. A new genus in the Provalljapyginae from
Missouri. Proceedings of the biological Society of Washington 73: 261-266.
SMITH, L.M. 1961. Japygidae of North America, 8. Postembryonic development of Parajapy-
ginae et Evalljapyginae (Insecta, Diplura). Annals of the entomological Society of
America 54: 437-441.
SMITH, L.M. 1962. Japygidae of South America, 2. The genus Provalljapyx (Insecta: Diplura)
The Pan-Pacif. Entomologist 38: 237-241.
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REVUE SUISSE DE ZOOLOGIE, 103 (2): 369-381; juin 1996
Une évaluation de Stygiocampa, sous-genre troglomorphe
de Plusiocampa (Diplura Campodeidae),
avec la description d'une nouvelle espece de Serbie orientale.
Bruno CONDE* & Camille BARETH**
* Musée de Zoologie de l'Université et de la Ville de Nancy,
34, rue Sainte-Catherine, F-54000 Nancy, France.
** Biologie des Insectes, Université de Nancy 1, BP 239,
F-54506 Vandœuvre-lès-Nancy Cedex, France.
An appraisal of Stygiocampa, a troglomorphic subgenus of Plusiocampa
(Diplura Campodeidae), with description of a new species from eastern
Serbia. — Plusiocampa (Stygiocampa) christiani n. sp. from two caves in
Eastern Serbia is described. The geographical distribution of the now five
known troglomorphic species of the subgenus Stygiocampa and the mor-
phological adaptative characters are discussed, shared by those species are:
a large size of the body (up to 9.5 mm), very long antennae by shape and
number (up to 59) of the articles, enlargement of the cupuliform organ at
the tip of antenna which houses up to 15 olfactive receptors, elongated legs
and claws with large lateral crests, low number of macrochaetae on the
greater number of tergits and the legs, multiplication of macrochaetae on
abdominal sternits I-VII (up to 252 whole), propensity for the loss of
glandular setae in males, very long cerci (2—3.5 times longer than the
body). A map of the distribution area is provided.
Key-words: Campodeidae - Taxonomy - Caves- Balkans.
INTRODUCTION
La découverte d'un Campodéidé inédit dans deux grottes de Serbie orientale,
visitées par le Dr E. Christian, en juillet 1984, nous incite a tenter une évaluation des
criteres morphologiques du sous-genre Stygiocampa Silvestri, 1934 qui rassemble des
Plusiocampa troglomorphes de grottes de la région balkanique et dont l'espèce type
Campodea nivea Joseph, 1882, fut la première espèce de Diploure troglobie décrite en
Europe et la deuxième au monde, après Campodea cookei Packard, 1871, en
Amérique du Nord.
Le sous-genre Stygiocampa Silvestri, 1934, espèce type Campodea nivea
Joseph, 1882, par monotypie, correspond au groupe IV du genre Plusiocampa (CONDE
Manuscrit accepté le 16.08.1995.
370 BRUNO CONDÉ & CAMILLE BARETH
1956: 127), l'unique espèce du groupe V (Plusiocampa denisi Condé) lui ayant été
adjointe par PACLT (1957: 45).
L'aire géographique s'étend, du Nord au Sud, de la Vénétie julienne au
Montenegro; vers l'Est, elle atteint la province de Zagreb près de Kamange au Nord-
Est, et la rive gauche de l'Iskar, dans le district de Sofia, au Sud-Est. Les stations de la
nouvelle espèce, près de Zlot, en Serbie orientale, sont à moins de 200 km au Nord-
Ouest des localités bulgares.
( 1. P. (S.) bureschi Silvestri
* 2.P.(S.) christiani n. sp.
v 3. P. (S.) dalmatica Condé
4. P. (S.) nivea Joseph
I]
AUTRICHE 4
os)
~ LEN) Sl . P. (S.) remyi Condé
DIR, De ÿ Sr Ke HONGRIE Solis (Dy) aay) i
EA u x 6. P. (S.) cf.remyi
I
ITALIE >, Ljubljana N
4°
SA Zagreb xy
Sibenik® v
«Split
Nevesinje &
I
ALBANIE / i
i {
\ -d-
200 km \ >
it 4 \ Boss
/
Echelle S 7 GRECE
re
2
FIG. 1
Répartition des espèces du sous-genre Stygiocampa.
Les matériaux de cette contribution, y compris les types, sont déposés au
Muséum d'Histoire naturelle de Genève, Département des Arthropodes et d'Entomo-
logie I, à l'exception des spécimens étudiés par Denis et Silvestri, et des P. (S.)
bureschi, ces derniers appartenant au National Natural History Museum de Sofia.
EVALUATION DE STYGIOCAMPA, SOUS-GENRE TROGLOMORPHE DE PLUSIOCAMPA 371
LISTE DES ESPÈCES
P. (S.) nivea (Joseph, 1882)
Décrit de la Lukova Jama (Slovénie), a été révisé par DENIS (1923), puis par
SILVESTRI (1934), sur un spécimen de la Koschanski griza Pesina (Slovénie), donné
par Joseph à Moniez, et illustré par SILVESTRI (1947) sur des spécimens de la Grotta
del Tricolore (Postojna). Cette espèce occupe un vaste territoire dans la province de
Trieste, en Slovénie et, en Croatie, dans les provinces de Lika-Krbava (entre Otocac
et Gospic) et de Zagreb (une grotte près de la frontière slovène) (CONDÉ 1959). Déjà
connue d'une bonne trentaine de cavités où elle est abondante (plus de 250 spécimens
nous ont été communiqués), elle est aussi présente dans la Sercetova Jama, VG 290,
dans la commune de Trieste, où F. Gasparo a récolté 3 d et 7 2 en 1982, 1985 et
1986, à ca 40 m de l'entrée, dans la zone obscure et humide, errant sur les concrétions,
sous les pierres et près d'appâts de viande.
P. (S.) remyi Condé, 1947
Est cantonné en Herzégovine, Dalmatie méridionale aux environs de Dubrov-
nik et Monténégro. Aux stations énumérées en 1959 (168, 170), il convient d'ajouter,
pour l'Herzégovine, les trois suivantes: Provalija, district de Nevesinje, près de la
route reliant cette localité à Gacko, C.L. Deeleman, 25.07.62: 1 2; Snjetnica Pesina,
Kifino Selo, P.R. & C.L. Deeleman, 23.07.63: 1 2; Vodena Jama (ou Pezina) à une
heure au N de Gacko, P.R. & C.L. Deeleman, 07.62: 2 ©. Il est abondant dans la
Vjetrenica, l'une des deux localites des syntypes (36 spécimens reçus), les dix autres
grottes ayant fourni 30 individus.
P. (S.) cf. remyi Condé
Dans la Vranjaca Pesina de Kotlenica, près de Split (Dalmatie), P.R. & CL.
Deeleman ont récolté, le 30.07.64, 3 spécimens (1 4, 2 2) d'une forme incertaine,
dans laquelle les femelles n'ont pu être distinguées de celles de P. (S.) remyi (une
seule antenne intacte de 50 articles). Le mâle, en revanche, possède un premier
urosternite dont les angles latéro-postérieurs, à peine saillants, n'atteignent que le 1/5
proximal de l'appendice. Ce dernier est subcylindrique, à peine 2 fois plus long que
large (140/65), sa longueur égale aux 4/5 environ de la distance le séparant de son
symétrique; le champ apical comporte plus de 80 phaneres sensoriels épais (a).
Marge postérieure du sternite avec un champ de phanères glandulaires (g,) disposés
sur 5 à 7 rangs: macrochètes au nombre de 24+24. L'aspect général du sternite est plus
proche de celui de la femelle de P. (S.) remyi que de celui du mâle de cette espèce,
compte non tenu des phanères glandulaires marginaux.
Papille mâle très pileuse, avec une rosette de 16 soies autour du gonopore.
L'hypothèse d'un individu présentant une anomalie individuelle ne peut être
écartée et de nouvelles récoltes dans cette cavité sont nécessaires. On remarquera
cependant que la grotte est située à plus de 100 km des localités de P. (S.) remyi.
BRUNO CONDÉ & CAMILLE BARETH
372
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FIG. 2
Urosternite I de mäles adultes (semi-diagrammes). A. Plusiocampa (Stygiocampa) cf. remyi de
la Vranjaca Pesina, Kotlenica, Dalmatie. — B. Plusiocampa (Stygiocampa) remyi Condé,
spécimen de la série typique de la Vjetrenica Pesina, Herzégovine, d'après CONDE 1947.
EVALUATION DE STYGIOCAMPA, SOUS-GENRE TROGLOMORPHE DE PLUSIOCAMPA 373
P. (S.) dalmatica Condé, 1959
Décrit sur une unique femelle en très médiocre état de conservation, récoltée
dans la Pesina Strazbenica, non loin de Sibenik, il a été retrouvé en Vénétie julienne
dans la Grotta Bac, commune de Trieste, par F. Gasparo en 1979. BARETH (1988: 48)
en a décrit une larve de 2,7 mm, apportant à cette occasion de nombreuses précisions
sur cette espèce encore mal connue, en particulier une antenne complète de 47 articles
avec un organe cupuliforme apical à nombreux sensilles très modifies!.
L'espèce paraît assez abondante dans cette grotte où F. Gasparo & C. Semorile
ont récolté 9 nouveaux spécimens les 25.08.84 (1 2), 09.08.87 (1 9), 26.02.89 (3 2)
et 05.03.89 (1 d, 3 ©). Elle est présente également dans la Grotta Pocala, VG 91,
commune de Duino-Aurisina, province de Trieste, où F. Gasparo a récolté 1 9 sur un
morceau de bois à ca 80 m de l'entrée, le 19.06.88.
En Dalmatie, l'espèce a été reprise dans la Manita Pecina, près de Paklenica,
au
NE de Zadar, par P.R. & C.L. Deeleman, 02.07.64: 1 4,2 9.
Tête. — Antenne de 41, 42, 44 (2 fois) et 46 (2 fois) articles, outre des
régénérats de 32 à 42 articles.
Thorax. — Un seul métanotum à chétotaxie asymétrique (1+0 ma) chez une
femelle de 5,30 mm de la Grotta Bac.
Abdomen. — Les mâles possèdent un champ glandulaire continu sur la marge
postérieure du sternite I. Chez le spécimen de 3,9 mm (Grotta Bac), les phanères gy,
au nombre d'environ 90, sont disposés sur 2-3 rangs; les appendices sont subcylin-
driques et portent 5 phanères apicaux a,. Chez le mâle de 5,20 mm (Manita Pesina),
le champ glandulaire est plus large, comportant 6-8 rangs de phaneres gy.
Cerques. — Les cerques d'une femelle de 5,5 mm de la Grotta Bac sont pré-
sents, l'un et l'autre amputés d'une portion distale. Le gauche, long de 14,55 mm,
comporte une très longue base (653) subdivisée en 13 articles secondaires qui, sauf le
premier (46), sont de longueurs croissantes (23-82), suivie de 14 articles primaires de
longueurs croissantes (87-322). Le droit, long de 13,27 mm, comprend une base un
peu plus courte (534), découpée en 12 articles secondaires dans des rapports compa-
rables à ceux de l'autre cerque (48 et 27-72) et 16 articles primaires de longueurs
croissantes (34-222). Les rapports corps/cerques (0,37-0,41) sont comparables à ceux
mentionnés chez P. (S.) remyi (0,33-0,40), compte tenu de l'absence d'un article au
moins aux cerques disponibles.
Les longs macrochètes, y compris ceux des articles distaux, sont très finement
barbelés sur leur moitié apicale au moins.
P. (S.) bureschi Silvestri, 1931 (syn.: P. (S.) rauseri Rusek, 1965)
Cette espèce est cantonnée dans les grottes de la rive gauche de I'Iskar, District
! Un lapsus! (BARETH 1988: 49) a fait mentionner "le développement réduit des crêtes
latérales des griffes", ce qui est en contradiction avec le texte de la diagnose originale (:166)
"griffes inégales à crêtes latéro-tergales très développées" et la fig. 2,D de BARETH.
374 BRUNO CONDÉ & CAMILLE BARETH
de Sofia (Bulgarie). La grotte nommée Gornata Pestera par SILVESTRI (= Raziska
Dupka), l'une des trois localités citées dans la description originale, est la même que
la Suchata Pestera (V.B. Guéorguiev, in litt.) d'où RUSEK (1965) a décrit son P. ($.)
rauseri, un synonyme subjectif récent de P. (S.) bureschi.
Le pro- et le mésonotum portent seuls des macrochètes dont la disposition a été
décrite, mais non figurée par SILVESTRI; CONDÉ (1956: 44) a cru que la description du
mésonotum se rapportait au demi-tergite et non au tergite tout entier. Cette erreur
d'interprétation (formule 4+4 au lieu de 2+2: ma, /p3) est évidemment responsable de
la description de P. (S.) rauseri, dans laquelle se trouve la première et seule repré-
sentation (fig. 18) des tergites thoraciques de P. (S.) bureschi.
Plusiocampa (Stygiocampa) christiani n. sp. (Figs 1, 3, 4)
Serbie orientale. — Lazarova Pe3ina, près de Zlot, non loin (ca 30 km) de la frontière
bulgare, E. Christian, 15.07.84: 1 2 (holotype). Vernjikica Pesina non loin de la grotte précé-
dente et dans le même massif, E. Christian, 15.07.84: 1 4 mature, 1 9 (paratypes).
Femelle holotype
Les nombres entre parenthèses se rapportent aux paratypes lorsqu'ils diffèrent
de l'holotype.
Longueurs. — Corps: 6,82 mm (5,95; 5,20); cerque: 15,50 mm; corps/cerque =
0,44; patte métathoracique: 3,48 mm; corps/p. III = 1,96.
Teguments. — Epicuticule sans ornementation. Soies de revêtement finement
barbelées sur leur moitié distale au moins.
Tête. — L'antenne gauche est un régénérat de 27 articles, la droite en compte
31(32, n=3). Sensille de l'article III postéro-sternal, subcylindrique et atténué à l'apex,
de même forme que celui du palpe maxillaire et du palpe labial, mais environ 1,5 fois
plus court qu'eux (18/27,5 um) et beaucoup plus grêle. Les articles suivants présentent
chacun un verticille apical de sensilles en gouge, assez grêles, dont le nombre dépasse
la vingtaine sur les articles de la région moyenne. L'article apical est environ 2,25 fois
aussi long que large et l'organe cupuliforme occupe environ 1/5 de la longueur totale; ce
dernier renferme au moins 12 sensilles à l'antenne droite et environ 18 au régénérat (14
chez le mâle) de forme inhabituelle: la collerette périphérique est épaisse et le bouton
central fendu jusqu'à la base ou presque; en coupe optique, le sensille apparaît digité.
Palpe labial droit avec 13 phanères banaux sur la portion antérieure et 128
phanères gustatifs sur la portion postérieure; 2 courts poils de garde de part et d'autre
du sensille latéro-externe.
Processus frontal large, mais peu saillant, portant 3 phanères barbelés plus
forts que leurs voisins et disposés en triangle (macrochètes), ainsi que 3+3 phanères
ordinaires, aucun ne présentant d'embase tuberculée.
Thorax. — Le pronotum porte seul des macrochètes indubitables, au nombre de
4+4 (ma, la, laz, lp3) dont les longueurs relatives, de ma à /p3, sont 68; 68,5; 81; 87.
12+11 soies marginales entre les /p, longues (55-76, Zp/N = 64,5), grêles et finement
barbelées ressemblant, à l'embase près, aux macrochètes voisins.
EVALUATION DE STYGIOCAMPA, SOUS-GENRE TROGLOMORPHE DE PLUSIOCAMPA 375
Plusiocampa (Stygiocampa) christiani n. sp., femelle holotype de la Lazarova Pesina et mâle
paratype de la Vernjikina Pesina. — A. Pronotum de la femelle. — B. Sensilles du mâle: Ille
article antennaire (à gauche), palpes maxillaire (au centre) et labial (à droite). — C. Sensilles de
l'organe cupuliforme du mâle. — D. Portion apicale d'un article de la région moyenne de
l'antenne de la femelle avec une rangée de sensilles en gouge et un court sensille subclaviforme.
— E. Portion apicale du tarse III gauche et télotarse du mâle, face antérieure. — F. Urosternite I
du mâle. — G. Style gauche du IVe urosternite du mâle.
376 BRUNO CONDÉ & CAMILLE BARETH
Mésonotum avec une paire de phanères, un peu plus longs (50) et plus épais
que leurs voisins, qui correspondent, par leur position et leur forme, à des subma-
crochètes latéraux antérieurs (s/a). Ces phanères sont indiscernables chez les deux
paratypes. Métanotum sans phanères différenciés en dehors des soies marginales.
Pattes grêles et allongées, l'extrémité des métathoraciques atteignant presque le
bord postérieur du dernier segment de l'abdomen en extension. Les articles, à partir du
trochanter, sont dans les rapports 18, 62, 76, 56. Pas de macrochète tergal ni sternal au
fémur et au tibia; les macrochètes antéro-marginaux du fémur, ainsi que les calcars
tibiaux, sont les seuls phanères différenciés. Pas de pseudo-articulation tarsienne;
toutes le soies, y compris les 3 longues soies subapicales dorso-latérales et celles de la
paire subapicale sternale, sont barbelées jusqu'à l'apex. Griffes très inégales (95/66), à
crêtes ornementées, le talon de la postérieure recouvrant l'apex du tarse. Processus
télotarsaux sétiformes et glabres.
Abdomen. — Une paire de macrochètes postérieurs (posts) en VII (absents chez
les paratypes). Tergites VII avec 3+3 (post3_5) et IX avec 7+7 (total), les macrochètes
de la paire la plus tergale plus faibles que les autres. Valvule supra-anale avec 2 soies
médianes et 2+2 latérales.
Sternites I à VII présentant une néotrichie comparable à celle de la plupart des
espèces du sous-genre (dalmatica, nivea, remyi), le sternite I étant ici environ deux
fois plus pileux que les suivants et le sexe n'intervenant pas (tableau I).
TABLEAU I
Répartition des macrochètes des urosternites chez les espèces présentant une néotrichie
(ou hypertrichie)
No des sternites I II III IV V VI VII Total
P.(S.) dalmatica? 95,5 mm 21+19 11+11 10+10 949 901011066157
d3,9mm 18+19 11+12 10+10 10+10 10+9 -10+10 10+10 159
P.(S.) christiani Sholo. 29425 == 12+11 12+11 12+12 12412 13+13 1963
d para. 21H21 12+12 13412" 12412 VIBRARE ZINIO
© para. 21+25 11+117 12413 24772 “12414 RSA
P.(S.) nivea* 15+15 14+14 14+14 14+14 14+14 14+14 14+14 198
P.(S.) remyi (syntypes)> d22+23 19+19 16+18 16+19 17+16 18+18 14+17 252
223+23 18+18 16+16 14+14 15+14 17+16 15+15 234
Appendices du sternite I subcylindriques, légèrement atténués a l'apex, un peu
moins de 2 fois aussi longs que larges (1,85), leur base presque 4,5 fois dans leur
écartement (4,40) et leur longueur 2,3 fois environ (2,36). 24 et 23 phanères apicaux
épais (a1).
2 Grotta Bac (spécimens bien lisibles).
3 On a admis que le sternite II, mal lisible, portait 11+11 macrochétes, soit le nombre
minimum observé chez l'espèce.
4 cf. SILVESTRI 1934.
5 Vjetrenica Pesina.
EVALUATION DE STYGIOCAMPA, SOUS-GENRE TROGLOMORPHE DE PLUSIOCAMPA 377
Sternite VIII: avec 4+4 (4+5, mâle paratype). Papille génitale très pileuse: cha-
que volet avec respectivement 24 et 21 soies courtes, et le tubercule avec un total de 43,
soit un groupe subapical de 12 soies courtes, en avant desquelles des phanères, de plus
en plus longs vers la base du mamelon, forment 4 rangées de 4, 8, 7 et 12 éléments.
Cerques. — Deux cerques, l'un complet (15,5 mm), l'autre ne comportant que 6
articles distaux (10 mm) flottaient dans le tube renfermant l'holotype. Le cerque
complet comprend une base (108), subdivisée en 3 articles secondaires (32, 33, 43), et
6 articles primaires dont la longueur augmente progressivement, les derniers étant
extraordinairement allongés. Les 6 articles du cerque fragmentaire ne présentent pas
cette particularité (il pourrait s'agir d'un appendice régénéré).
Tableau II
Longueurs relatives des articles et longueurs totales des cerques (ces dernières exprimées aussi
en millimètres) de l'holotype.
Base I Il III IV V VI Total
Cerque complet 1035765 93 ee 133m el 225 258 1051 = 15,5 mm
Fragment -- 90 101 110 124 124 130 679 = 10 mm
Rapport corps/cerques = 0,44
Le revêtement est constitué de phanères courts, ceux de la base étant sensible-
ment égaux aux 2/3 de la largeur de l'article mis à plat. Tous sont pourvus de barbules
assez longues et bien séparées les unes de autres sur leurs 2/3 distaux ou au moins leur
moitié distale. Les phanères formant un verticille à l'extrémité des articles primaires
sont densément barbelés presque depuis la base, ayant l'aspect d'arbuscules. Les macro-
chètes de l'article apical sont seulement fourchus avec souvent une barbule subapicale;
leur longueur est égale à la largeur de l'article ou la dépasse d'environ un tiers.
Male paratype. — Appendices du sternite I plus courts et plus épais que ceux de
la femelle, environ 1,5 fois aussi longs que larges (1,36-1,50), leur base environ 3 fois
dans leur écartement et leur longueur un peu plus de 2 fois (2,19). 28 et 26 phanères
apicaux épais (a). Papille génitale très pileuse (les soies toutes glabres), gonopore
entouré d'une rosette de 19 phanères.
Affinités. — L'espèce est remarquable par le faible nombre de macrochètes
tergaux: 5 ou 6 seulement pour l'ensemble des demi-tergites thoraciques et abdo-
minaux. Elle se rapproche en cela de P. (S.) bureschi qui en possède 9. En revanche,
elle présente une néotrichie des urosternites I à VII (190-194 macrochètes) qui
n'existe pas chez bureschi (voir tableau III).
Faisceaux de spermatozoïdes (Fig. 4)
P. (S.) nivea. — Le filament spiralé, seul élément bien visible, ressemble à
celui de P. (S.) remyi (BARETH 1968: 382).Il décrit 1,5 tour de spire environ pour une
longueur de 200 à 250 um. De forme elliptique, ses diamètres externe et interne sont
378 BRUNO CONDÉ & CAMILLE BARETH
SOIL B,C,D,E
FIG. 4
Plusiocampa (Stygiocampa) dalmatica Condé. A. Urosternite I du mâle de 3,9 mm de la Grotta
Bac. — B à E: faisceaux de spermatozoïdes, à plat et en coupe optique. — B. P. (S.) nivea. — C. P.
(S.) remyi. — D. P. (S.) bureschi. — E. P. (S.) christiani. Les portions pleines des filaments spiralés
sont représentées par un pointillé régulier; les touffes de spermatozoides sont figurées par un
tireté irrégulier sur les vues à plat et par un fin pointillé (sections) sur les coupes optiques.
EVALUATION DE STYGIOCAMPA, SOUS-GENRE TROGLOMORPHE DE PLUSIOCAMPA 379
respectivement 50/40 et 35/25 um. La tête renflée, d'un diamètre d'une douzaine de
um sur une longueur de 20 environ, est suivie d'un étranglement à la suite duquel le
filament s'élargit pour atteindre jusqu'à 10 um de diamètre, puis s'atténue ensuite
progressivement jusqu'à son extrémité. Un peu après l'étranglement, une cloison
transverse est visible, précédée de 2-4 petites cavités contigués et surmontée d'une
mince cloison perpendiculaire qui rejoint la paroi du filament au delà de
l'étranglement .Toute cette région antérieure semble creuse, tandis que le reste du
filament dont la densité optique est plus élevée apparaît plein. Les spermatozoïdes,
peu nombreux, ne sont pas enroulés autour du filament comme d'habitude: fixés à
l'extrémité acuminée du filament, ils s'écartent ensuite de ce dernier.
P.(S.) remyi. — Le filament spiralé décrit 1,5 tour de spire pour une longueur de
220 um environ. La tête renflée, d'un diamètre d'environ 8 um sur une longueur de
18-20, est suivie d'un étranglement à la suite duquel le filament s'élargit pour
atteindre 8-10 um, puis s'atténue progressivement jusqu'à son extrémité. Tout le
filament semble plein et les structures décrites chez nivea n'ont pu être décelées. Les
spermatozoïdes, quoique peu nombreux, sont enroulés autour du filament, les
diamètres externe et interne de l'ensemble du faisceau, de forme elliptique, étant
60-65/45-50 et 30-33/25-27 um.
P. (S.) bureschi. — Les faisceaux de spermatozoïdes sont circulaires, leurs
diamètres externe et interne sont respectivement d'une cinquantaine et de 25 um
environ, sur une épaisseur de 20 um. Les spermatozoïdes, enroulés autour du filament
spiralé, sont bien visibles. Le filament décrit 3,5 tours de spire environ pour une
longueur de 300-350 um. Il est très légèrement renflé à une extrémité, atteignant 5-6
um, et progressivement atténué à l'autre. La portion renflée et le premier tour de spire
sont creusés d'une lumière, le reste du filament étant plein.
P. (S.) christiani. — Les faisceaux de spermatozoïdes sont circulaires, leurs
diamètres externe et interne sont respectivement de 45 et 20 um, sur une épaisseur de
10-12 um. Le filament spiralé décrit 1,5 tour de spire environ pour une longueur de
140-150 um seulement. Il est plein, atténué aux deux extrémités.
Le filament spiralé présente un certain intérêt taxonomique et sans doute
phylogénétique (BARETH 1968: 383) et il semble exister une relation entre ses dimen-
sions et la densité des phanères différenciés (macrochètes). C'est ainsi que le plus long
connu (12-13 tours de spire pour une longueur de 1500-2000 um) appartient à
Campodea sensillifera Condé & Mathieu, 1958, espèce pourvue de macrochètes
nombreux et très robustes. La chétotaxie tergale de P. (S.) christiani, la plus réduite
parmi les représentants du sous-genre, s'accorde avec les faibles dimensions du
filament.
DISCUSSION
Les cinq espèces les plus sûrement apparentées, nivea, bureschi, remyi,
dalmatica, christiani, dans l'ordre chronologique, présentent des tendances évolutives
communes qui peuvent se résumer ainsi:
380 BRUNO CONDÉ & CAMILLE BARETH
1. Grande taille: 6,5 mm dalmatica, 7 mm christiani, 7,5 mm nivea, 8 mm
remyi, 9,5 mm bureschi.
2. Longues antennes par la forme de leurs articles et le nombre de ceux-ci: 32 et
36 chez christiani et nivea; 47, 54 et 59 chez dalmatica, bureschi et remyi
respectivement, ces nombres correspondant aux maximums connus pour chaque espèce.
3. Grandes dimensions de l'organe cupuliforme de l'article apical de l'antenne
qui renferme de 7 à une quinzaine de sensilles (décompte souvent difficile), de forme
plus ou moins complexe, opposés aux 4 sensilles simples de la plupart des formes
endogées.
4. Allongement des articles des pattes et développement, sauf chez bureschi,
des crêtes latérales de la griffe postérieure
5. Régression du nombre des macrochètes tergaux du thorax et de l'abdomen,
de dalmatica à christiani, les autres espèces présentant des degrés intermédiaires,
dans l'ordre nivea, remyi, bureschi (tableau TT).
Tableau III
Répartition des macrochètes sur les tergites thoraciques et abdominaux.
Th. I Th. I Th. HI Ab.V VI VI
mas lass in ma late los masslamı Ip T post. T
dalmatica 1 4 2 1(0) 2 AAO) Ma? 1614) 4 4 4 12
nivea u Lane - 1 2 - - 2 12 4 4 4 12
remyi IRA? - 1 2 - - 2. 12 BAT 3 9
buneschu u lat 7222 1 - l - - - 7 - = 2 2
christiani 1 2 1 - 10) - - - - 5(4) SRE 1(0) 1
6. Réduction du nombre ou absence des macrochètes des fémurs et tibias.
7. Néotrichie (hypertrichie) des sternites abdominaux I a VII, sauf bureschi.
8. Absence de champ glandulaire marginal au sternite I du mâle, sauf chez
remyi, cf. remyi et dalmatica.
9. Allongement des cerques: presque 2 fois la longueur du corps (bureschi),
un peu plus de 2 fois (christiani) ou 3 fois à 3 fois 1/2 (dalmatica, remyi, nivea).
REMERCIEMENTS
La mise au propre de la figure 2 est due au talent de Madame E. Seraoui. Nous
lui exprimons notre vive gratitude pour son aide.
BIBLIOGRAPHIE
BARETH, C. 1968. Biologie sexuelle et formations endocrines de Campodea remyi Denis
(Diploures Campodéidés). Revue d’Ecologie et de Biologie du Sol 5: 303-426.
BARETH, C. 1988. Campodes cavernicoles de la Vénétie orientale (Frioule, Vénétie julienne) et
de la Calabre (Insecta Apterygota Diplura). Atti e Memorie della Commissione Grotte
"Eugenio Boegan" 27: 45-55.
EVALUATION DE STYGIOCAMPA, SOUS-GENRE TROGLOMORPHE DE PLUSIOCAMPA 381
ConDE, B. 1947. Campodéidés nouveaux des grottes balkaniques. Notes biospéologiques 1:
17-32.
CONDE, B. 1956. Matériaux pour une Monographie des Diploures Campodéidés. Mémoires du
Museum national d'Histoire naturelle, Paris 12: 1-202.
CONDÉ, B. 1959. Un singulier Diploure cavernicole de Dalmatie. Fragmenta Balcanica Musei
Macedonici Scientarum Naturalium 2: 165-172.
DENIS, R. 1923. Notes sur les Apterygotes. — I. Sur quelques Aptérygotes décrits par M.
Moniez. Annales de la Societé entomologique de France 92: 209-236.
PACLT, J. 1957. Diplura. Genera Insectorum 212: 123 pp.
RUSEK J. 1965. Zur Kenntnis der Campodeidae (Diplura) Bulgariens. Acta entomologica
bohemoslovaca 62: 92-97.
SILVESTRI F. 1931. Contributo alla conoscenza dei Campodeidae (Thysanura) delle grotte della
Bulgaria. /zvestiya na Tsarskite Prirodonauchni Instituti v Sofiya 4: 97-107.
SILVESTRI, F. 1934. Tisanuri cavernicoli della regione di Postumia. Arti del I. Congresso
Speleologico Nazionale, Giugno 1933, Trieste: 179-181.
SILVESTRI, F. 1947. Illustrazione della Plusiocampa (Stygiocampa) nivea Joseph (Campodeidae,
Diplura) delle grotte di Postumia. Bollettino del Laboratorio di Entomologia Agraria di
Portici 8: 88-92.
ax ite lavano eh
2 not
MER] fx
REVUE SUISSE DE ZOOLOGIE, 103 (2): 383-394; juin 1996
Nouvelles données sur les Trématodes d'Amphibiens
d'Afrique Occidentale.
Description d'Haematoloechus aubriae n. sp.
Robert BOURGAT, Cyril ROURE, & Sim-Dozou KULO
Laboratoire de Biologie, Université, 52 avenue de Villeneuve,
F-66860 Perpignan cedex.
New data on the Trematodes of Amphibians from Western Africa.
Description of Haematoloechus aubriae n. sp. - This study reports new
hosts for three Trematodes: Xenopus tropicalis (Gray, 1864) for
Progonimodiscus doyeri (Ortlepp, 1926), Ptychadena pumilio (Boulenger,
1920) for Mesocoelium monodi Dollfus, 1929, and Bufo togoensis Ahl,
1924 for Maederia eburnense (Maeder, 1969). It specifies or extends the
distribution of these Helminths and that of Pleurogenoides tener (Looss,
1898). Phrynobatrachotrema eburnense Goodmann, 1986 is considered a
synonym of Maederia eburnense (Maeder, 1969). Haematoloechus aubriae
n. sp., lung parasite of Aubria subsigillata (Duméril, 1856) in Porto-Novo
(Bénin) is erected.
Key-words: Trematodes — Western Africa — Amphibians — Progoni-
modiscus — Mesocoelium — Maederia — Pleurogenoides — Haematoloechus.
INTRODUCTION
Cette étude est consacrée à une série de 26 Trématodes livrés par la dissection
de 128 Amphibiens appartenant a 6 espéces provenant de la République du Bénin, du
Niger, du Nord Togo, et collectés en 1977 et 1978.
Les parasites, fixés sur le terrain au Bouin alcoolique et ultérieurement colorés
au carmin boracique de Grenacher, sont montés in toto et dessinés à la chambre claire.
Les dimensions sont données en micrometres, sauf indications contraires.
La nomenclature des hôtes Ranidae est fondée sur la révision de Dußois (1992).
PARAMPHISTOMIDAE
Progonimodiscus doyeri (Ortlepp, 1926) (fig. 1)
Synonymie: Diplodiscus doyeri Ortlepp, 1926; Diplodiscus subclavatus Goeze,
sensu GROBBELAAR 1922; Progonimodiscus doyeri victoriani Vercammen-Grandjean,
1960.
Manuscrit accepté le 12.09.1995.
384 ROBERT BOURGAT, CYRIL ROURE & SIM-DOZOU KULO
VERCAMMEN-GRANDIEAN (1960) observe que plusieurs caractères du Diplo-
discus doyeri d'Ortlepp ne permettent pas de le maintenir dans le genre Diplodiscus
Diesing, 1836 et crée pour lui un nouveau genre de Diplodiscinae: Progonimodiscus.
Dans ce même travail Vercammen-Grandjean crée pour les Progonimodiscus du lac
Kivu une sous-espèce nouvelle: P. d. victoriani. PRITCHARD (1964), aprés examen de
l'holotype de P. d. victoriani estime que les caractères invoqués par Vercammen-
Grandjean sont compris dans les limites de variation individuelle de P. doyeri et met
la sous-espèce P. d. victoriani en synonymie avec P. doyeri, conclusion acceptée par
SEY (1983). Suivant ce raisonnement, notre matériel se rapporte à l'espèce P. doyeri
(Ortlepp, 1926).
Mentions anterieures: VERCAMMEN-GRANDJEAN (1960) précise que Diplo-
discus subclavatus, sensu GROBBELAAR (1922), ainsi que D. doyeri, sensu ORTLEPP
(1926), BRAVO-HOLLIS (1941), SKRJABIN (1949) et YAMAGUTI (1958), sont en fait des
Progonimodiscus. Depuis cette date, P. doyeri a été retrouvé par:
PRITCHARD (1964) chez Xenopus laevis (Daudin, 1802) en Sud Afrique,
BEVERLEY-BURTON (1963), qui le nomme encore Diplodiscus doyeri, chez X.
laevis, en Rhodésie,
GASSMANN (1975), chez Conraua crassipes (Buchholz & Peters, 1875) au
Cameroun,
FISCHTHAL (1977), chez Xenopus muelleri (Peters, 1844) en Tanzanie, et chez
X. laevis victorianus Ahl, 1924 et X. fraseri Boulenger, 1905 au Zaire.
Hôte, localité et occurrence: Xenopus tropicalis (Gray, 1864), Porto-Novo
(République du Bénin), (10/31).
Date de la récolte: 25 mai 1977.
Habitat: Intestin.
Materiel examine: 12 exemplaires montes in toto.
Dimensions: L. (mm) = 2,9 à 4,1 (moy. 3,6); I. (mm) = ESP (me);
V.O. = 429 à 561 (moy. 474); Ac. (mm) = 1,5 à 2 (moy. 1,7); V.O./Ac. = 0,24 à 0,36
(moy. 0,28); Oes. = 429 à 627 (moy. 551); Tes. = 229 à 308 (moy. 261); Ov. = 198 à
270 (moy. 233); oeuf = 103 x 60.
Discussion: X. tropicalis est un hôte nouveau pour P. doyeri et la découverte
de ce parasite en République du Bénin étend son aire de répartition à l'Afrique
occidentale.
MESOCOELIIDAE
Mesocoelium monodi Dollfus, 1929 (fig. 2)
Synonymie: La taxonomie des Mesocoelium d'Afrique a fait l'objet de plusieurs
analyses contradictoires: THOMAS (1965), MAEDER et al. (1969b), MAEDER (1973).
Ces hésitations s'expliquent par la diversité des hôtes de ce parasite et son cosmo-
politisme sans qu'il n'offre de nettes variations morpho-anatomiques. Actuellement, la
dénomination spécifique M. monodi parait admise: FISCHTHAL (1977), BOURGAT
(1979), PIKE (1979)...
385
TREMATODES D’AMPHIBIENS D’AFRIQUE OCCIDENTALE
Fic. 2: Mesocoelium monodi
Fic. 1: Progonimodiscus doyeri
Dollfus, 1929, face ventrale.
(Ortlepp, 1926), face ventrale.
386 ROBERT BOURGAT, CYRIL ROURE & SIM-DOZOU KULO
Mentions antérieures: Une liste des précédentes récoltes de M. monodi en
Afrique, est donnée par BOURGAT (1979). Depuis cette date, M. monodi est encore
signalé par:
PIKE (1979) chez Bufo regularis Reuss, 1834 au Soudan,
FASHYI (1986) qui a obtenu son dékystement expérimental au Nigeria.
Hôtes, localités et occurrence: Bufo regularis Reuss, 1834, Niamey (Sud
Niger), (1/4); Ptychadena pumilio (Boulenger, 1920), Dapaong (Nord Togo), (1/3).
Date de la récolte: 28 février 1978 (Niamey).
Habitat: Intestin.
Matériel examiné: 6 exemplaires montés in toto.
Dimensions: L. (mm) = 2,6 à 3,15 (moy. 2,9); I. (mm) = 0,68 à 0,79 (moy.
0,73); V.O. = 188 à 255 (moy. 236) x 240 à 274 (moy. 261); Ac = 146 à 225 (moy.
186) x 174 à 225 (moy. 199); V.O./Ac. = 1,2 à 1,4 (moy. 1,32); Tes. gauche = 186 x
261, Tes. droit = 211 x 275; Ov. = 158 à 188 (moy. 176) x 191 à 248 (moy. 217); oeuf
= 23 à 28 (moy. 26) x 29 à 37 (moy. 33).
Discussion: Ptychadena pumilio (Boulenger, 1920) est un hôte nouveau, et le
Niger une région inédite pour M. monodi, dont le caractère euryxène et l'ubiquité sont
ainsi confirmés.
LECITHODENDRIIDAE
Pleurogenoides tener (Looss, 1898) (fig. 3)
Synonymie: Distomum tacapense Sonsino, 1894, sensu Looss (1896); Pleuro-
genes tener Looss, 1898.
Mentions antérieures: Looss (1896) chez Chamaeleo sp. en Egypte,
ODHNER (1910) chez Chamaeleo basilicus en Egypte,
Macy (1964) chez Chalcides ocellatus en Egypte,
FISCHTHAL (1976, 1977) chez Hoplobatrachus occipitalis (Gunther, 1859),
Bufo regularis et Chamaeleo dilepis dilepis Leach, 1820 en Afrique Equatoriale,
BOURGAT & COMBES (1977) chez H. occipitalis au Sud Togo,
BOURGAT (1979), idem,
PIKE (1979) chez H. occipitalis au Soudan,
GROSCHAFT & MORAVEC (1985) chez Chamaeleo chamaeleon (Linnée, 1758)
en Egypte.
Hôtes, localités et occurrence: Hoplobatrachus occipitalis (Gunther, 1859),
Kante (Nord Togo). Sur 81 Grenouilles disséquées a Kante, une seule a livré 7 P.
tener, dont 6 ont été préparés.
Date de la récolte: 13 décembre 1977.
Habitat: Intestin.
Matériel examiné: 6 exemplaires montés in toto.
Dimensions: L. (mm) = 0,95 à 1,05 (moy. 0,99); I. (mm) = 0,76 à 0,80 (moy.
0,79); V.O. = 120 à 150 (moy. 140) x 169 a 191 (moy. 176); Ac. = 161 à 244 (moy.
185) x 157 à 268 (moy. 185); V.O./Ac. = 0.71 à 1,11 (moy. 0,975); Tes. = 109 à 200
TREMATODES D’AMPHIBIENS D’ AFRIQUE OCCIDENTALE 387
LINE
dy]
D
A e
Upp FASS
©:
NZ
I
Ho DI
SR
VI
PSE
DATI
UA né Ù WES
SS
SEG DS 2207,
LAY ye X
Ri A
FIG. 3
Pleurogenoides tener (Looss, 1898), face ventrale.
(moy. 158) x 203 à 341 (moy. 273); Ov. = 124 à 143 (moy. 135) x 131 à 184 (moy.
164); oeuf = 19 à 30 x 12 à 16 (moy. 25 x 15).
Discussion: La découverte de P. tener au Nord Togo élargit sa répartition aux
savanes sèches d'Afrique occidentale. On peut noter que dans cette région il est infé-
odé à H. occipitalis. Au Cameroun P. tener a été trouvé par FISCHTHAL (1977) chez B.
regularis, mais au Togo aucune des 112 dissections de ce crapaud ne l'a livré.
L'expérience de Macy (1964) qui a obtenu l'infestation de P. mascareniensis, montre
que la spécificité de P. tener n'est pas de nature physiologique, mais vraisembla-
blement écologique ou liee à l'éthologie des hôtes intermédiaires.
PLAGIORCHIDAE
Maederia eburnense (Maeder, 1969) (fig. 4)
Synonymie: Haplometroides eburnense Maeder, 1969; Plagitura eburnense
(Maeder, 1969) Gassmann, 1975; Phrynobatrachotrema eburnense (Maeder, 1969),
Goodmann, 1986.
388 ROBERT BOURGAT, CYRIL ROURE & SIM-DOZOU KULO
Aprés un nouvel examen des exemplaires étudiés par Gassmann, ainsi que du
matériel collecté ultérieurement au Togo, BOURGAT & COMBES (1979) concluent à
l'impossibilité de les ranger ni dans le genre Haplometroides ni Plagitura, et pro-
posent la création du nouveau genre Maederia.
En 1986, GOODMANN décrit des helminthes livrés par Rana fuscigula
angolensis (Bocage, 1866) et par Phrynobatrachus natalensis (Smith, 1849) en
Ouganda; en particulier, 1l en donne une description trés fine du système excréteur.
Goodmann identifie son matériel à celui découvert par MAEDER (1969) en Côte
d'Ivoire et par GASSMANN (1975) au Cameroun, mais 1l ne reconnait pas les
attributions génériques (Haplometroides, Plagitura) proposées successivement par
ces deux auteurs et crée pour l'espèce eburnense le genre Phrynobatrachotrema. Ce
raisonnement, correct, ne prend toutefois pas en compte le travail de BOURGAT &
ComBEs (1979) et la création du genre Maederia qui, par antériorité, reste prioritaire.
Le genre Phrynobatrachotrema tombe donc en synonymie avec Maederia.
Mentions antérieures: MAEDER (1969) chez Phrynobatrachus sp. et Ptycha-
dena longirostris (Peters, 1870) en Côte d'Ivoire,
MAEDER & al. (1969a) chez Bufo funereus funereus Bocage, 1866 et B. came-
runensis camerunensis Parker, 1936 au Gabon,
MAEDER & al. (1970) chez Hylarana albolabris (Hallowell, 1856) en Centr-
afrique,
MAEDER (1973) chez Phrynobatrachus alleni Parker, 1936, P. liberiensis
Barbour & Loveridge, 1927, P. plicatus (Günther, 1858) et Prychadena longirostris
en Côte d'Ivoire,
GASSMANN (1975) chez Bufo latifrons Boulenger, 1900, Hylarana sp., H. lepus
(Andersson, 1903), H. longipes (Perret, 1960), Cardioglossa gracilis Boulenger,
1900, Phrynobatrachus auritus Boulenger, 1900, P. batesii (Boulenger, 1906), P.
steindachneri Nieden, 1910, Pedropedetes cameronensis Reichnow, 1914, Astylos-
ternus sp., A. batesi (Boulenger, 1900), Trichobatrachus robustus Boulenger, 1900, et
Leptodactylodon ventrimarmoratus (Boulenger, 1904) au Cameroun,
BOURGAT & COMBES (1979) chez Phrynobatrachus accraensis et P. calcaratus
(Peters, 1863) au Sud Togo,
BOURGAT (1979), idem,
GOODMANN (1986) chez R. f. angolensis et Phrynobatrachus natalensis en
Ouganda.
Hote, localité et occurrence: Bufo togoensis Ahl, 1924, Tinkoro (Togo), (1/7).
Date de la récolte: 4 juillet 1978.
Habitat: Intestin. M. eburnense est associé à Mesocoelium monodi.
Matériel examiné: | exemplaire monté in toto.
Dimensions: L. (mm) = 2,3; I. (mm) = 0,8; V.©. =209'x 218 AC Pas
V'OYAc = 1,22: Oes. = 155; Tes. = 150 x 221; Ova= 195 x 30050eUR 2252288
Discussion: L'exemplaire étudié ici présente l'intérêt de révéler un hôte
nouveau, B. togoensis, et une localisation inédite, le Nord Togo.
Il correspond tout à fait, par ses dimensions et ses caractères morpho-ana-
tomiques (situation de l'ovaire et des testicules, longueur des caeca jusqu'au deux tiers
TREMATODES D’AMPHIBIENS D’ AFRIQUE OCCIDENTALE 389
du corps, distribution des vitellogènes, extension de l'utérus jusqu'à l'extrémité posté-
rieure du corps) aux principales descriptions antérieures (MAEDER et al. 1969,
GASSMANN 1975). Le dessin de GOODMANN (1986) montre que l'utérus n'atteint pas
l'extrémite postérieure du corps mais se limite au niveau du testicule droit; le texte
cependant, précise "uterus ... extending posteriorly almost to end..." ce qui permet
d'associer le matériel ougandais aux M. eburnense du reste de l'Afrique. L'aire de
repartition de cette espèce s'étend donc de la Côte d'Ivoire à l'Ouganda, c'est-à-dire
sur un vaste territoire paléo-tropical.
HAEMATOLOECHIDAE
Haematoloechus aubriae n. sp. (fig. 5)
Hôte: Aubria subsigillata (Dumeril, 1856). Les poumons du seul individu
disséqué hébergeaient l'Haematoloechus étudié ici; son intestin des Mesocoelium et
des Diplodiscus. Au Togo, A. subsigillata héberge Opisthorchis lomeensis Bourgat &
Combes, 1975, exemple rare de Trématode biliaire d'Amphibien en Afrique.
Localité: Porto-Novo (Sud Bénin). Dans cette même station, un Trématode
rare a été découvert: Gorgoderina beninensis Bourgat, Dossou & Gasc, 1976, parasite
vésical de l'Amphibien A. occipitalis.
Occurrence: 1/1.
Date de la récolte: 25 mai 1978.
Habitat: Poumons.
Matériel examiné: 1 exemplaire monté in toto.
Description: Corps allongé, terminé postérieurement par un mucron; tegument
lisse, relativement épais (3,5 um); ventouse orale sub-terminale, acétabulum absent.
Appareil digestif: Cavité buccale séparée du pharynx par un court prépharynx;
pharynx pyriforme; oesophage court; les caecums, aprés leur bifurcation, se dirigent
transversalement, puis se contournent une fois en hélice avant de s'allonger jusqu'à
proximité de l'extrémité postérieure.
Appareil reproducteur mâle: testicules globuleux, légèrement lobés, inter-
caecaux, situés aux 2/3 du corps, disposés en diagonale, le droit étant décalé vers
l'arrière; poche du cirre conforme à la structure habituellement rencontrée chez les
Haematoloechus. Vaisseaux efférents visibles jusqu'à leur confluent au niveau de
l'ovaire.
Appareil reproducteur femelle: ovaire intercaecal, latéral droit, arrondi; récep-
tacle séminal grand, appliqué contre le bord interne de l'ovaire; vitellogènes rassem-
blés en grappes, 10 à gauche, 7 à droite, alignées latéralement en position paracaecale
externe; abondantes jusqu'au niveau des testicules, on n'en compte qu'une seule dans
la zone post-testiculaire; les circonvolutions de l'utérus comblent l'espace intercaecal
Jusqu'à la limite postérieure des caecums; il n'y a pas de boucle extracaecale.
Dimensions: L. (mm) = 9:T. (mm) = 2,45; V.O. = 454x484: Ac. = absent:
pharynx = 225 x 206 ; Test. gauche = 605 x 681, Test. droit = 577 x 615 ; Ov. = 397;
ER 13008700, ocut 348225:
390 ROBERT BOURGAT, CYRIL ROURE & SIM-DOZOU KULO
RS
[es
PSA
Ars.
Fi > À
n SII ASE"
AA Oy &
IN ax 2
Fic. 4: Maederia eburnense (Maeder, 1969), Fic. 5: Haematoloechus aubriae n. sp., face
face ventrale. dorsale.
TREMATODES D’AMPHIBIENS D’ AFRIQUE OCCIDENTALE 391
Dicussion: Ce Ver est un Haematoloechus. La présence du mucron suggère
l'espèce H. micrurus Rees, 1964; mais, l'absence de boucle utérine extracaecale, la
forme des testicules, (globuleux et non pas allongés ovales ou pyriformes), l'absence
d'acétabulum, (bien que "poorly developped" selon REES, 1964, l'acétabulum existe
chez H. micrurus), la disposition des vitellogènes, très rares dans le tiers postérieur du
Ver, ne répondent pas à la diagnose de H. micrurus.
Depuis sa description, H. micrurus a été signalé chez H. occipitalis par:
FISCHTHAL & THOMAS (1968), au Ghana,
BAER (1972), et MAEDER (1973), en Côte d'Ivoire,
GASSMANN (1975), au Cameroun,
BOURGAT et al. (1976), au Bénin,
FISCHTHAL (1977), au Zaïre,
BOURGAT (1979), au Togo,
PIKE (1979), au Soudan.
Au Cameroun, ASSANJI (1990) a découvert H. micrurus chez Bufo regularis,
qui n'est pas l'hôte habituel. Au Nigeria, OLADIMEN ef al. (1988-1990), signalent des
Haematoloechus sp. chez H. occipitalis.
Malgré le nombre des collectes, leur dispersion géographique et la diversité
des hôtes, aucun de ces travaux ne fait état de variations morphologiques telles que les
présente notre matériel; leur déterminisme n'est donc vraisemblablement pas de nature
écomorphotique. Les caractères concernant les ventouses, la taille des oeufs, les
boucles utérines extracaecales, et la position des testicules ayant, selon KENNEDY
(1980), la plus grande valeur taxonomique, l'Haematoloechus d'Aubria subsigillata du
Bénin ne peut être identifié à l'espèce A. micrurus.
L'absence d'acétabulum et la présence du mucron le séparent des autres
espéces d'Haematoloechus d'Afrique centrale et occidentale; en outre, chacune de ces
espèces s'en différencie par plusieurs caractères:
H. ocellati Gassmann, 1975 — ovaire lobé, boucles utérines extracaecales pré-
sentes, vitellogènes en zone post-testiculaire, oeuf = 12 à 15,5 x 18 à 26;
H. combesi Batchvarov & Bourgat, 1974 — boucles utérines extracaecales
déterminant une forme "épaulée" tout à fait particulière, vitellogènes en zone post-
testiculaire, oeuf = 17 x 22;
H. darcheni Combes & Knoepffler, 1967 — acétabulum "délicat à découvrir"
mais présent, ovaire trés lobé, boucles utérines extracaecales présentes, vitellogènes
en zone post-testiculaire, oeuf = 14 à 17 x 18 à 22;
H. lobogonadus Meskal, 1970 — tégument spinulé, ovaire et testicules lobés,
boucles utérines extracaecales présentes, vitellogènes en zone post-testiculaire, oeuf =
1032176330222 25;
H. johnsoni Bourgat, 1977 — tégument faiblement spinulé, boucles utérines
extracaecales présentes, vitellogenes en zone post-testiculaire;
H. dollfusinus (Odening, 1958) — selon Dollfus (1950), ovaire lobé, circon-
volutions utérines débordant les caecums, oeuf = 17 à 18,3 x 25 à 28,3;
H. variegatus leonensis Williams & Coker, 1967 — boucles utérines extra-
caecales présentes, vitellogènes en zone post-testiculaire.
392 ROBERT BOURGAT, CYRIL ROURE & SIM-DOZOU KULO
Cet ensemble de raisons justifie la création d'une espèce nouvelle que nous
proposons de nommer Haematoloechus aubriae pour rappeler l'identité de l'hôte. Ses
principaux caractères sont: Corps terminé postérieurement par un mucron, tégument
lisse, acétabulum absent, prépharynx présent, testicules intercaecaux, globuleux,
disposés en diagonale, ovaire arrondi, vitellogènes rares dans le tiers postérieur du
corps, pas de boucle utérine extracaecale, oeufs ovales mesurant (en um) 22,5 x 34.
L'holotype est déposé au Muséum d'Histoire naturelle de Genève sous le n°
214 77 INVE.
CONCLUSION
Cette étude a pour objet une série d'Helminthes originaires du Niger et du
"corridor dahoméen", territoire étroit, étiré de l'océan au Sahel, et formé de la
République du Bénin et du Togo. A ce niveau la savanne, descendue jusqu'au littoral,
sépare la zone humide et forestière "éburnéo ghanéenne" occidentale, de l'orientale,
ininterrompue du Nigeria à l'Afrique équatoriale. Cette situation se traduit dans la
composition de la faune des Trématodes d'Amphibiens dont certaines espèces, selon
BOURGAT (1979), sont ubiquistes, alors que d'autres restent inféodées à l'une ou à
l'autre des trois zones.
Les résultats confortent bien le schéma général:
— l'ubiquité de M. eburnense et de M. monodi se confirme; l'aire du premier,
qui parasite un hôte nouveau, se prolonge jusqu'au Nord Togo, celle du second
jusqu'au Niger.
— P. doyeri, en parasitant un hôte nouveau pénètre dans le "corridor daho-
méen" où, toutefois, il reste localisé à sa frontière orientale.
— L'aire de P. tener monte jusqu'aux savannes sèches où il se concentre sur un
seul hôte trés inféodé aux pièces d'eau permanentes.
— Quant à H. aubriae, nouvelle espèce de Trématode endémique découverte
dans le "corridor dahoméen" aprés H. combesi, G. beninensis, et O. lomeensis, il
témoigne que les conditions particulières de "l'exception écologique bénino-togolaise"
favorisent l'existence de formes endémiques sans interdire les espèces venues des
blocs voisins.
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REVUE SUISSE DE ZOOLOGIE, 103 (2) 395-437; juin 1996
Aleocharinae della Colombia e dell'Ecuador: Parte I
(Coleoptera, Staphylinidae)
(131° Contributo alla conoscenza delle Aleocharinae)
Roberto PACE
Via Vittorio Veneto, 13. 37032 Monteforte d'Alpone (Verona), Italia.
Aleocharinae from Colombia and Ecuador: Part I (Coleoptera, Sta-
phylinidae). - This paper adds further new genera, new species and data
obtained in the course of many new research in Colombia and Ecuador
made by the Dr H.G. Müller, Dr L. Bartolozzi and Dr Prof. H. Franz and in
the past. Three new species of the tribe Myllaenini are described; four of
the tribe Hygronomiini; eight of the tribe Gyrophaenini; eighteen of the
tribe Diestotini; four of the tribe Homalotini and one of the tribe
Boliticharini. Hoplandria rubicula (Erichson) is transferred to the new
genus Cyllogerusa; Thamiaraea discors (Erichson) in transferred to the
new genus Trichidryas. The new genera Polytelusa and Caralepta are
described for two new species. Four new combinations are recorded.
Key-words: Coleoptera - Staphylinidae - Aleocharinae - Taxonomy -
Colombia - Ecuador.
INTRODUZIONE
E' noto che le foreste pluviali e le regioni montuose del Sudamerica ospitano
un gradissimo numero di specie animali e vegetali. Allorché il Dr Hans Georg Miiller
dell'Università di Giessen ha effettuato ricerche sui Pantopodi in Colombia, non ha
trascurato di raccogliere anche esemplari di Aleocharinae. Essi, affidatimi in esame
tramite il Dr Volker Puthz della "Limnologische Flusstation" di Schlitz, sono qui
pubblicati. Al termine dell'esame del materiale di Aleocharinae raccolto dal Dr.
Miiller, ho constatato anche per questa sottofamiglia di Stafilinidi una presenza molto
consistente di specie per la Colombia, per lo più non ancora descritte. A questo
materiale ho aggiunto altre specie della Colombia e del confinante Ecuador: un
gruppo appartiene a un vecchio fondo delle collezioni dello "Zoologisches Museum"
dell'Università Humboldt di Berlino, fondo affidatomi in studio dal Dr M. Uhlig di
detto Museo. Un altro lotto di Aleocharinae fa parte delle raccolte effettuate in
Ecuador dal Prof. H. Franz di Mödling, Vienna. Un ultimo lotto aggiunto è il frutto di
recenti ricerche effettuate dal Dr Luca Bartolozzi del Museo Zoologico "La Specola"
dell'Università di Firenze.
Manoscritto accettato il 25.08.1995.
396 ROBERTO PACE
Gli holotypi sono conservati nel "Museum d'Histoire naturelle" di Ginevra
(MG), nello "Zoologisches Museum" dell'Università Humboldt di Berlino (MB), in
collezione del Prof. Franz (CFR) e nel Museo Zoologico "La Specola" dell'Università
di Firenze. Alcuni paratypi sono conservati anche nel "Museo Zoologico della
Pontificia Università Catdlica del Ecuador" di Quito (MQ).
ELENCO DELLE SPECIE (escluse le nuove)
OLIGOTINI
Oligota (s. str.) parva Kraatz, 1862
Oligota parva Kraatz, 1862: 300
1 2, Ecuador, Manabi, dint. Puerto Cayo, 21.11.1993 (L. Bartolozzi & G. Onore leg.).
Specie a larga diffusione, dal Nordafrica attraverso le Canarie fino al
Nordamerica, Messico e Brasile. Nuova per l'Ecuador.
Oligota (s. str.) tricolor Biering, 1934
Oligota tricolor Biering, 1934: 116; Frank, 1972: 139
1 2, Kolumbien Villa Culebra bei Bonda, X.1985 (H.G. Müller leg.).
Specie finora nota solo di Cuba.
GYROPHAENINI
Gyrophaena (s. str.) quassa Sharp, 1876
Gyropnaena quassa Sharp, 1876: 76
6 es., Colombia, Muzo (Collez. Mus. Berlin).
Specie diffusa nella regione amazzonica.
Gyrophaena (Agaricochara) mahunkai Pace, 1991
Gyrophaena (Agaricochara) mahunkai Pace, 1991: 154
1 & e 3 2 2, Kolumbien, Villa Culebra bei Bonda, ca. 10 Km E Sta. Marta, XI-X1.1985,
Barberfallen, (Müller leg.).
Specie finora nota solo dell'Isola di Sta. Lucia.
DIESTOTINI
Plesiomalota (s. str.) hispidula (Fauvel, 1866)
Homalota hispidula Fauvel, 1866: 316
Atheta (s. str.) hispidula: Bernhauer & Scheerpeltz, 1926: 644
Plesiomalota (s. str.) hispidula: Pace, 1986: 425
3 es., Ecuador, Rio Hollin, 9-13.11.1993, (Bartolozzi leg.).
Specie finora nota solo del Cile.
ALEOCHARINAE DELLA COLOMBIA E DELL'ECUADOR 397
Diestota (Apheloglossa) curticollis (Erichson, 1840), comb. n.
Homalota curticollis Erichson, 1840: 118
Atheta (s. str.) curticollis: Bernhauer & Scheerpeltz, 1926: 641
3 2 2, Kolumbien, Bahia de Cinto, Tayrona Park, ca. 30 Km NE Sta. Marta, Fallaub,
IV.1985 (Müller leg.); 1 9, Kolumbien, Dept. Magdalena, Rio Frio, ca. 50 Km NE Sta. Marta,
21.VIIL.1985, (Müller leg.).
Specie già nota della Colombia. La nuova combinazione è stabilita dopo esame
della serie tipica di Aragua (Colombia) composta di sole femmine.
Diestota (Apheloglossa) flavipennis (Erichson, 1940), comb. n.
Homalota flavipennis Erichson, 1840: 118
Atheta (s. str.) flavipennis: Bernhauer & Scheerpeltz, 1926: 643
3 2 2, Kolumbien, Punta de Betin, Sta. Marta, IV.1986, (Müller leg.).
Specie finora nota solo dell'Isola di St. John (Antille). Combinazione nuova
stabilita in base all’esame degli esemplari della serie tipica di St. John.
Eudera didyma (Frichson, 1840), comb. n.
Homalota didyma Erichson, 1840: 116
Ophioglossa didyma: Bernhauer & Scheerpeltz, 1926: 570
Diestota (Apheloglossa) didyma: Pace, 1987: 188
2 2 2, Kolumbien, Punta de Betin, Sta. Marta, 12.1V.1986, (Müller leg.); 1 6, Kolumbien,
Villa Culebra bei Bonda, ca. 10 Km E Sta. Marta, 1.XII.1985, (Müller leg.).
Specie già nota della Colombia e di Guadalupa.
Questa specie va esclusa dal genere Ophioglossa perché non presenta il mento
profondamente incavato al margine anteriore e da Diestota per avere il collo stretto.
Eudera bisulcata (Erichson 1840), comb. n.
Homalota bisulcata Erichson, 1840: 117
Ophioglossa bisulcata: Bernhauer & Scheerpeltz, 1926: 569
2 364, Kolumbien, Sierra Nevada de Sta. Marta, bei San Pedro de la Sierra, 12.1V.1986,
(Miiller leg.).
Specie presente in Colombia e in Brasile.
Anche questa specie va esclusa dal genere Ophioglossa perché il margine ante-
riore del mento non è profondamente incavato.
BOLITOCHARINI
Thecturota schuberti (Pace, 1983), comb. n.
Leptischnusa schuberti Pace, 1983: 308
1 2, Kolumbien, Punta de Betin, 1-26.1V.1986, (Müller leg.).
Specie diffusa in Brasile, in Bolivia e in Cile. Nuova per la Colombia.
Raccolta al lume.
FALAGRIINI
Falagria (Myrmecocephalus) strieta Erichson, 1840
Falagria stricta Erichson, 1840: 51
398 ROBERTO PACE
1 8, Kolumbien, Nördl. Sierra Nevada de Sta. Marta, San Lorenzo, 1700 m,
25.V111.1985, (Müller leg.); 9 es., Kolumbien, Sierra Nevada de Sta. Marta, San Pedro de la
Sierra, 12.1V.1986, (Müller leg.).
Specie gia nota della Colombia e del Venezuela.
Meronera albicincta (Erichson, 1840)
Falagria albicincta Erichson, 1840: 56
Meronera albicincta: Bernhauer & Scheerpeltz, 1926: 580
1 2, Kolumbien, Sierra Nevada de Sta. Marta, San Pedro de la Sierra, 12.1V.1986,
(Müller leg.); 1 2, Kolumbien, Villa Culebra bei Bonda, ca. 10 Km E Sta. Marta, XI-X11.1985,
Barberfallen, (Müller leg.); 25 es., Kolumbien, Punta de Betin, Sta. Marta, IV.1986, Farb-
schalen, (Müller leg.); 1 4, Ecuador, Cotopaxi, S. Francisco de las Pampas, 1300-1500 m,
11.1993, (Bartolozzi leg.).
Specie diffusa in Colombia, Venezuela, Brasile, Cuba, Messico e Guadalupa.
Nuova per l'Ecuador.
ATHETINI
Atheta (Microdota) brasiliana Bernhauer, 1908
Atheta (Microdota) brasiliana Bernhauer, 1908: 358
1 2, Kolumbien, Bahia de Gairaca, Tayrona Park, ca. 20 Km NE Sta. Marta, 31.X.1985
(Miiller leg.); 11 es., Kolumbien, Villa Culebra bei Bonda, ca. 10 Km E Sta. Marta,
XI-XI1.1985, (Müller leg.).
Specie finora nota solo del Brasile.
Il confronto con la spermateca e l'edeago dei tipi rende indubbia l'attribuzione
specifica.
Atheta (Acrotona) parcior Bernhauer, 1927
Atheta (Acrotona) parcior Bernhauer, 1927: 257
1 d, Ecuador, Cotopaxi, S. Francisco de las Pampas, 1300-1500 m, II.1993, (Bartolozzi leg.).
Specie finora nota dell'Argentina, del Brasile e del Cile. Esemplari comparati
con l'holotypus d di Caspinchango.
Atheta (Datomicra) columbica Fauvel, 1901
Atheta columbica Fauvel, 1901: 88
22 es., Kolumbien, Villa Culebra bei Bonda, ca. 10 Km E Sta. Marta, XI-X11.1985, (Müller leg.;
175 es., Kolumbien, Punta de Betin, Sta. Marta, IV.1986, (Müller leg.); 1 d e 2 2 2, Kolumbien,
Bahia de Gairaca, Tayrona Park, ca. 20 Km NE Sta. Marta, 19.VI.1985, (Miiller leg.).
Specie finora rinvenuta solo in Colombia, comparata con 1 tipi.
Atheta (Datomicra) conformis (Erichson, 1840)
Homalota conformis Erichson, 1840: 108
Atheta (s. str.) conformis Bernnauer & Scheerpeltz, 1926: 640
3 dd e 3 99, Ecuador, Cotopaxi, S. Francisco de las Pampas, 1300-1500 m, 1I.1994,
(Bartolozzi leg.); 2 2 2, Kolumbien, Punta de Betin, Sta. Marta, IV.1986, (Müller leg.); 2 dd,
Kolumbien, Villa Culebra bei Bonda, 10 Km E Sta. Marta, XI-X11.1985, (Müller leg.).
Specie diffusa in Colombia, Venezuela, Isola di St. Thomas e Puertorico,
comparata con 1 tipi.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 399
Atheta (Datomicra) pseudogagatina Bernhauer, 1934
Atheta (s. str.) pseudogagatina Bernhauer, 1934: 504
1 do, Kolumbien, Punta de Betin, Sta. Marta, XI-X11.1985, (Müller leg.).
Specie diffusa in Argentina e Brasile. Nuova per la Colombia.
L'holotypus € una femmina.
Atheta (Dimetrota) bicallosa Bernhauer, 1920
Atheta bicallosa Bernhauer, 1920: 152
1 gd, Kolumbien, Nord-Westl. Sierra Nevada de Sta. Marta, 1000 m, 12.1V.1986, (Müller leg.); 1
3 e2 2 2, Kolumbien, Sierra Nevada de Sta. Marta nahe San Pedro, 12.1V.1986, (Müller leg.).
Specie montana diffusa solo in Colombia, comparata con 1 tipi.
Atheta (Pseudobessobia) columbina Bernhauer, 1920
Atheta (s. str.) columbina Bernhauer, 1920: 151
1 2, Kolumbien, Punta de Betin, Sta. Marta, 27.1V.1986, (Müller leg.).
Specie finora nota sul solo holotypus 9 della Colombia da me esaminato. La
forma della spermateca, senza difficolta, permette di attribuire la specie al sottogenere
Pseudobessobia Bernhauer, 1921, il cui genotipo weiseri Bernhauer ho esaminato.
Lamprostiba schwabei (Bernhauer, 1939)
Atheta (Philhygra) schwabei Bernhauer, 1939: 14
Lamprostiba schwabei: Pace, 1986: 444
1 ©, Ecuador, Cotopaxi, S. Francisco de las Pampas, 1300-1500 m, 11.1993, (Bartolozzi leg.).
Specie del Cile e del Perù. Nuova per l'Ecuador.
SCHISTOGENIINI
Neoplatyola hospes (Pace, 1990), comb. n.
Platyola hospes Pace, 1990: 70
3 5 5, Kolumbien, Punta de Betin, 1-10.V.1986, (Müller leg.).
Specie segnalata per il Brasile e la Bolivia. Nuova per la Colombia.
DESCRIZIONI
MYLLAENINI
Myllaena chibcha sp. n. (Figg. 1-4)
Tri. Holotypus d, Kolumbien, Punta de Betin, Sta. Marta, IV.1986, (Müller leg. MG).
Paratypi: 2 9 2, stessa provenienza; 21 es. Punta de Betin, 1-26.IV.1986, Lichtfalle
und Farbschalen, (Miiller leg.).
DESCRIZIONE. Lunghezza 2,1 mm. Avancorpo opaco, addome lucido. Corpo
bruno-rossiccio; addome bruno con apice rossiccio; antenne nere con antennomeri 1,
9, 10 e 11 giallo-rossicci; zampe giallo-rossicce. Una pubescenza fitta copre l'intero
corpo. Edeago figg. 2-3, spermateca fig. 4.
400 ROBERTO PACE
COMPARAZIONI. Per la forma della spermateca la nuova specie pud essere
avvicinata tassonomicamente a M. insipiens Casey, 1911, del Nordamerica, ma l'ede-
ago è meno profondamente ricurvo al lato ventrale e ha pezzi copulatori del sacco
interno molto robusti (esilissimi in insipiens). Inoltre la spermateca della nuova specie
presenta più complesse circonvoluzioni della parte prossimale rispetto quelle semplici
di insipiens.
ETIMOLOGIA. Dai Chibha, popolazione colombiana stanziata sugli altipiani
della Colombia al tempo della conquista spagnola.
Myllaena muelleri sp. n. (Figg. 5-8)
Tipi. Holotypus 4, Kolumbien, Punta de Betin, Sta. Marta, IV.1986, Farbschalen,
(Müller leg., MG).
Paratypus: 1 ©, stessa provenienza.
DESCRIZIONE. Lunghezza 2,0 mm. Corpo debolmente lucido e bruno-rossiccio;
capo bruno; margine posteriore degli uroterghi e quinto urotergo libero, tranne la base,
rossicci; antenne nere con antennomeri | e 11 rossicci; zampe giallo-rossicce. L'intero
corpo è coperto di fitta e aderente pubescenza. Edeago figg. 6-7, spermateca fig. 8.
COMPARAZIONI. Specie tassonomicamente avvicinabile a M. currax Notman,
1920 della Florida, in base alla forma della spermateca che ha la parte prossimale
flessa su sè stessa. Tuttavia la nuova specie presenta bulbo distale della spermateca
stessa molto più dilatato e il sacco interno dell'edeago presenta delle spine, assenti nel
sacco interno dell'edeago di currax.
ETIMOLOGIA. Specie dedicata al suo raccoglitore, il Dr Hans Georg Müller di
Giessen, studioso di "Pantopoda".
Myllaena puthzi sp. n. (Figg. 9-12)
Tripi. Holotypus & , N Kolumbien, Nordwestl. Sierra Nevada de Sta. Marta, ca. 1000 m,
nahe San Pedro de la Sierra, | 12.1V.1986, Fallaub, (Müller leg., MG).
Paratypi: 1 4 e 3 © 9, stessa provenienza; 3 © 2, Kolumbien, Nördl. Sierra Nevada de
Sta. Marta, nähe El Campero, ca. 1000 m, aus Fall-Laub, 29.1V.1986. (Müller leg.).
DESCRIZIONE. Lunghezza 2,3 mm. Corpo debolmente opaco e bruno; estremità
addominale bruno-rossiccia; antenne brune con antennomeri 1, 8, 9, 10 e 11 giallo-
rossicci; zampe giallo-rossicce. L'intero corpo è coperto di pubescenza fitta e corta.
Edeago figg. 10-11, spermateca fig. 12.
COMPARAZIONI. Anche questa specie, come la precedente, per la forma della
spermateca è tassonomicamente avvicinabile a M. currax Notman, 1920, della Florida.
Ma la nuova specie è microttera, non atta al volo e la parte prossimale della sua
spermateca è nettamente più corta di quella di currax e il bulbo distale della spermateca
stessa è molto più sviluppato. Anche l'edeago della nuova specie presenta caratteri
differenziali: tra l'altro la parte preapicale in visione laterale, è gibbosa ventralmente,
mentre in currax è piana.
ETIMOLOGIA. La specie è dedicata al Dr Volker Puthz di Schlitz, noto studioso di
Steninae. Ciò in segno di ringraziamento per avermi affidato in studio le Aleocharinae
della Colombia raccolte dal Dr Miiller e qui illustrate.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 401
FIGG. 1-8
Habitus, edeago in visione laterale e ventrale e spermateca. 1-4: Myllaena chibcha sp. n.; 5-8:
Myllaena muelleri sp. n.
402 ROBERTO PACE
HYGRONOMIINI
Gyronycha filiformis sp. n. (Figg. 13-17)
Tipo. Holotypus 2, Kolumbien, Punta de Betin, Sta. Marta, 1-26.IV.1986, Lichtfalle
und Farbschalen, (Müller leg., MG).
DESCRIZIONE. Lunghezza 1,6 mm. Corpo lucido e bruno-giallioccio, comprese
le antenne che pero hanno i due antennomeri basali gialli; zampe gialle. L'avancorpo è
coperto di punteggiatura fitta e superficiale; l'addome é coperto di tubercoletti fini. La
reticolazione è assente sull'avancorpo ed è estremamente svanita sull'addome.
Spermateca fig. 17.
COMPARAZIONI. La nuova specie è distinta da G. valens Casey, 1893, specie
nordamericana, per avere contemporaneamente capo più stretto del pronoto e antenne
corte. Nella specie nordamericana, che ha capo più stretto del pronoto, le antenne
sono molto lunghe.
Caloderella neotropica sp. n. (Figg. 18-21)
Tipo. Holotypus d, Colombia, Buga (MB).
DESCRIZIONE. Lunghezza 3,7 mm. Corpo lievemente opaco e rossiccio; quarto
urite libero bruno-rossiccio; antenne rossicce; zampe giallo-rossicce. Il capo e il
pronoto non hanno microscultura reticolare e sono coperti di punteggiatura fine, netta
e profonda. La reticolazione che copre le elitre è svanita, quella dell'addome è netta, a
maglie circolari. La punteggiatura delle elitre è ben distinta. Il quinto urotergo libero
ha pubescenza meno fitta di quella del primo e secondo visibili. Edeago figg. 19-21.
COMPARAZIONI. La nuova specie è distinta da C. argentina Bernhauer, 1912,
dell'Argentina, per la taglia maggiore: 3,7 mm invece di 2,2-2,8 mm e per avere il
pronoto lungo quanto largo (più lungo che largo in argentina).
Caloderella humboldti sp. n. (Figg. 22-26)
Tripi. Holotypus 4 , Colombia, Muzo, (MB).
Paratypus: 1 ®, stessa provenienza.
DESCRIZIONE. Lunghezza 3,7 mm. Corpo debolmente lucido e rossiccio scuro;
elitre bruno-rossicce; antenne rossicce con antennomero basale giallo; zampe giallo-
rossicce. Indistinta è la reticolazione dell'avancorpo, quella dell'addome, al contrario,
è netta. La punteggiatura dell'avancorpo è fitta e molto superficiale. Il quinto urotergo
libero del maschio presenta un gruppo di tubercoli nettamente salienti sulla metà: ai
lati i tubercoli sono finissimi. Edeago figg. 23-25, spermateca fig. 26.
COMPARAZIONI. Specie simile a C. fraterna Bernhauer, 1912, dell'Argentina,
ma differente, tra l'altro, per la taglia maggiore (3,7 mm, invece di 3,0 mm) e per il
quinto urotergo libero del maschio coperto di granuli meno fittamente distribuiti.
ETIMOLOGIA. Specie dedicata ad Alexander von Humboldt, geografo e natura-
lista tedesco, a ricordo del suo viaggio di studio tra il 1759 e il 1804 attraverso i paesi
dell'America meridionale.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 403
FIGG. 9-17
Habitus, edeago in visione laterale e ventrale, spermateca, labio con palpo labiale, maxilla con
palpo mascellare e mento. 9-12: Myllaena puthzi sp. n.; 13-17: Gyronycha filiformis sp. n.
404 ROBERTO PACE
Caloderella carinata sp. n. (Figg. 27-30)
Tipi. Holotypus d, Bogota (MB).
Paratypus: 1 ©, stessa provenienza.
DESCRIZIONE. Lunghezza 3,9 mm. Corpo un po' appiattito, lucido e giallo-
rossiccio; elitre giallicce; antenne e zampe giallo-rossicce. L'avancorpo è privo di
reticolazione, l'addome presenta reticolazione distinta. La punteggiatura del capo, del
pronoto e delle elitre è fine, fitta e netta. Il primo urotergo libero del maschio è
attraversato sulla linea mediana da una carena smussata e molto saliente all'indietro.
Un tubercolo basale mediano inciso longitudinalmente sta sul quinto urotergo libero
del maschio. Edeago figg. 28-29, spermateca fig. 30.
COMPARAZIONI. A motivo della presenza di caratteri sessuali secondari sul
primo e sul quinto degli uroterghi liberi del maschio, costituiti da carena mediana e da
tubercolo mediano basale, la nuova specie è ben distinta sia da C. argentina
Bernhauer, 1912, che da C. fraterna Bernhauer, 1912, entrambe dell'Argentina.
GYROPHAENINI
Brachida columbiensis sp. n. (Figg. 31-34)
Tipo. Holotypus d, Colombia, (Moritz leg., N° 5272 MB, cotypus di Myrmedonia
tumidula Erichson, 1840).
DESCRIZIONE. Lunghezza 2,2 mm. Corpo lucidissimo e giallo-rossiccio; elitre
brune con base rossiccia; addome giallo-rossiccio chiaro con uriti liberi terzo e quarto
di un rossiccio scuro; antenne bruno-rossicce con gli antennomeri basali 1° a 4° e 11°
gialli; zampe giallo-rossicce. Sul corpo non vi è traccia di reticolazione. La pun-
teggiatura del capo è indistinta, quella del pronoto e dell'addome è assai svanita.
Edeago figg. 32-34.
COMPARAZIONI. La nuova specie è ben distinta da B. hirta (Erichson, 1840), del
Brasile, per la taglia minore, per gli antennomeri 4° a 7° molto più lunghi che larghi
(appena più lunghi che larghi in hirta) e per la pubescenza del corpo corta (molto
lunga in hirta). Non è possibile il confronto della forma dell'edeago poiché
l'holotypus di hirta è una femmina. Non è il caso di fare comparazioni con B.
importuna (Erichson, 1840) della Colombia poiché essa va trasferita al genere
Orphnebius Motschulsky, 1858, come ho constatato esaminando 1 tipi, comb. n.
Brachida caldasensis sp. n. (Figg. 35-38)
Tri. Holotypus d, C. Vieja (MB).
Paratypi: 3 © ©, stessa provenienza; 2 © 2, Colombie, Cordill.re Centr., 1877, (Dr O.
Thième, MB).
DESCRIZIONE. Lunghezza 3,4 mm. Corpo lucidissimo e giallo-rossiccio; parte
posteriore delle elitre sfumata di bruno; antenne giallo-rossicce con gli antennomeri
1° a 4° e 11° gialli; zampe giallo-rossicce. Sul corpo non vi è traccia di reticolazione.
Assente è la punteggiatura sul capo e sul pronoto, quella delle elitre è fine e svanita.
Solo il quinto urotergo libero presenta una reticolazione svanita. Edeago figg. 36-37,
spermateca fig. 38.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 405
Figc. 18-25
Habitus ed edeago in visione laterale, ventrale (solo l'apice) e dorsale. 18-21: Caloderella
neotropica sp. n.; 22-25: Caloderella humboldti sp. n.
406 ROBERTO PACE
FIGG. 26-30
Spermateca, habitus ed edeago in visione laterale, ventrale. 26: Caloderella humboldti sp. n.;
27-30: Caloderella carinata sp. n.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 407
FiGG. 31-38
Habitus, edeago in visione laterale, ventrale e dorsale e spermateca. 31-34: Brachida colum-
biensis sp. n.; 35-38: Brachida caldasensis sp. n.
408 ROBERTO PACE
COMPARAZIONI. La nuova specie mostra qualche affinità con B. tumidula
(Erichson, 1840), della Colombia, a motivo della forma dell'edeago. Ma quello della
nuova specie, se visto ventralmente, è molto largo e ha l'apice incavato, mentre
l'edeago di tumidula è stretto e termina a punta aguzza. Inoltre l'addome di tumidula
ha pubescenza molto lunga (corta nella nuova specie).
ETIMOLOGIA. La nuova specie prende nome da Caldas, regione della Cordillera
Centrale della Colombia.
Gyrophaena (s. str.) spatulata sp. n. (Figg. 38bis—43)
Tipi. Holotypus 4, Ecuador, Cotopaxi, S. Francisco de Las Pampas, 1300-1500 m,
V.1993, (L. Bartolozzi leg., N° 9853, MF).
Paratypi: 5 es., stessa provenienza (N° 9854 MF, MQ).
DESCRIZIONE. Lunghezza 2,2 mm. Corpo lucidissimo. Sono neri il capo e le
elitre, tranne gli omeri che sono bruni; pronoto bruno con margini laterali e posteriore
giallo-rossicci; addome giallo-rossiccio, tranne il quarto urite libero che è bruno;
antenne gialle con estremità distale sfumata di bruno; zampe gialle. La punteggiatura
del capo e del pronoto è come da fig. 38. Le elitre presentano punteggiatura fine e
tubercoletti salienti solo presso la sutura. La reticolazione è assente sul capo e il
pronoto, quella delle elitre e dell'addome è molto svanita: il quarto urotergo libero ha
reticolazione distinta, il quinto reticolazione netta e quattro carene, nel maschio.
Edeago figg. 39-40, spermateca fig. 41, sesto urotergo libero del maschio fig. 42,
sesto urotergo libero della femmina fig. 43.
COMPARAZIONI. La nuova specie è tassonomicamente vicina a G. quassa Sharp,
1876 dell'Amazzonia, a motivo della presenza di un robusto pezzo copulatore del
sacco interno dell'edeago. Tuttavia, l'edeago della nuova specie presenta un ampio
lobo laterale apicale, assente in guassa e apice dell'edeago molto prolungato al lato
ventrale (corto in quassa).
Gyrophaena (s. str.) cotopaxiensis sp. n. (Figg. 44-49)
Tipi. Holotypus d, Ecuador, Cotopaxi, S. Francisco de Las Pampas, 1300-1500 m,
II.1993, (L. Bartolozzi leg., N° 9851, MF).
Paratypi: 7 es. stessa provenienza (N° 9852, MF e MQ).
DESCRIZIONE. Lunghezza 3,3 mm. Corpo lucido e nero; elitre nero-brune,
antenne brune con i quattro antennomeri basali gialli e i tre seguenti giallo-bruni;
zampe gialle con femori bruni. Il corpo è privo di reticolazione, tranne sul quinto
urotergo libero dove è estremamente svanita. La punteggiatura del capo e del pronoto
è come da fig. 46. Le elitre sono coperte di robusti tubercoli poco salienti: fra essi
stanno tubercoletti fini, assenti all'angolo posteriore esterno che è compresso. Sesto
urotergo libero del maschio fig. 44, sesto urotergo libero della femmina fig. 45,
edeago figg. 47-48, spermateca fig. 49.
COMPARAZIONI. Specie tassonomicamente vicina a G. quassa Sharp, 1876,
dell'Amazzonia, ma ben distinta per l'edeago molto più sviluppato, con parte apicale
fortemente asimmetrica (in visione ventrale), (simmetrica in quassa).
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 409
FıGG. 38 bis-45
Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio e
della femmina. 38 bis-43: Gyrophaena (s. str.) spatulata sp. n.; 44-45: Gyrophaena (s. str.)
cotopaxiensis Sp. n.
410 ROBERTO PACE
Gyrophaena (Agaricochara) martensis sp. n. (Figg. 50-53)
Tipo. Holotypus d, Kolumbien, Punta de Betin, Sta. Marta, IV.1986, Farbschalen,
(Müller leg., MG).
DESCRIZIONE. Lunghezza 1,3 mm. Corpo lucido e bruno-gialliccio; antenne
giallo-rossicce con undicesimo antennomero oscurato; zampe gialle. La reticolazione
del capo e del pronoto è svanita, quella delle elitre è netta. Il capo e il pronoto sono
coperti di punteggiatura superficiale. Le elitre mostrano tubercoletti poco salienti su
tutta le loro superficie. Edeago figg. 51-52, sesto urotergo libero del maschio fig. 53.
COMPARAZIONI. In base alla forma dell'edeago, la nuova specie è tassono-
micamente affine a G. anomala Notman, 1920, della Florida. Tuttavia l'apice dell'
edeago della nuova specie è piegato ad uncino verso il lato dorsale, mentre quello di
anomala è volto al lato ventrale. Inoltre il sesto urotergo libero del maschio di ano-
mala è privo di caratteri sessuali secondari al margine posteriore, mentre nella nuova
specie è presente una punta mediana, larga e arrotondata.
Gyrophaena (Agaricochara) perincisa sp. n. (Figg. 54-56)
Tipo. Holotypus £, Ecuador, Cotopaxi, S. Francisco de Las Pampas, 1300-1500 m,
11.1993, (L. Bartolozzi leg., N° 9848, MF).
DESCRIZIONE. Lunghezza 1,8 mm. Corpo lucido e nero; antenne nere con i tre
antennomeri basali gialli; zampe gialle. La reticolazione del capo e del pronoto è
estremamente svanita, quella delle elitre è distinta e quella dell'addome è netta. La
punteggiatura del capo e del pronoto è fine. Distinti tubercoletti coprono la superficie
delle elitre. Il quinto urotergo libero del maschio ha dei tubercoli laterali assai
allungati. Spermateca fig. 55, sesto urotergo libero della femmina fig. 56.
COMPARAZIONI. Non sono note specie che presentano una profondissima inci-
sura sul sesto urotergo libero della femmina, come si osserva nella nuova specie. Solo
nel maschio di G. hubbardi Seevers, 1951, dell'Illinois, si riscontra una profonda
incisura sul sesto urotergo libero, ma essa è assai larga.
Gyrophaena (Agaricochara) praelapsa sp. n. (Figg. 57-58)
Tipo. Holotypus 9, Ecuador, Cotopaxi, S. Francisco de Las Pampas, 1300-1500 m,
11.1993, (L. Bartolozzi leg., N° 9849, MF).
DESCRIZIONE. Lunghezza 1,6 mm. Corpo lucido. Capo e pronoto nero-bruni;
margini laterali e posteriore del pronoto bruno-rossicci; elitre e uriti liberi quarto e
quinto neri; uriti liberi primo, secondo e terzo ed estremità addominale bruni; antenne
gialle, appena sfumate di bruno verso l'estremità distale; zampe gialle. La retico-
lazione del capo è molto svanita, quella del resto del corpo è distinta. La pun-
teggiatura del capo è fine. Il resto della superficie del corpo presenta tubercoletti
distinti. Spermateca fig. 58.
COMPARAZIONI. In base ai caratteri esterni, come il corpo ben reticolato e gli
angoli posteriori del pronoto indistinti, la nuova specie potrebbe essere tassono-
micamente vicina a G. sonorae Seevers, 1951, dell'Arizona, di cui è noto il solo
holotypus maschio. La nuova specie si distingue tra l'altro da G. sonorae per il
differente colore del corpo e la maggiore taglia (1,6 mm, invece di 1,0 mm).
411
ALEOCHARINAE DELLA COLOMBIA E DELL’ ECUADOR
Imm
0) mm
49
FicG. 46-53
Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio
46-49: Gyrophaena (s. str.) cotopaxiensis sp. n.; 50-53: Gyrophaena (Agaricochara) martensis
sp. n.
412 ROBERTO PACE
55
E
E
=
01 mm
58
FIGG. 54-58
Habitus, spermateca e sesto urotergo libero della femmina. 54-56: Gyrophaena (Agaricochara)
perincisa Sp. n.; 57-58: Gyrophaena (Agaricochara) praelapsa sp. n.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 413
Brachychara bartolozzii sp. n. (Figg. 59-61)
Tipo. Holotypus d, Ecuador, Cotopaxi, S. Francisco de Las Pampas, 1300-1500 m,
11.1993, (L. Bartolozzi leg., N° 9850, MF).
DESCRIZIONE. Lunghezza 2,0 mm. Corpo lucido. Capo e pronoto giallo-
rossicci; elitre e addome bruno-rossicci; antenne nere fino al settimo antennomero, il
resto delle antenne è andato perduto nella fase di raccolta; zampe gialle. La retico-
lazione del capo e del pronoto è assente, quella delle elitre è distinta e quella dell'
addome è svanita. La punteggiatura del capo è netta, quella del pronoto e dell'addome
è distinta. Tubercoletti ben salienti coprono la superficie delle elitre. Il quinto
urotergo libero ha reticolazione distinta e uno spazio posteriore privo di pun-
teggiatura. Edeago fig. 60, sesto urotergo libero del maschio fig. 61.
COMPARAZIONI. Specie distinta sia da B. crassa Sharp, 1883, che da B.
brevicornis Sharp, 1883, entrambe del Guatemala, per avere punteggiatura distinta sul
pronoto e pubescenza non molto rada (pronoto non punteggiato in crassa e pu-
bescenza molto rada in bevicornis) e per la forma del margine posteriore del sesto
urotergo libero del maschio. In crassa il lobo mediano tra le due spine del margine
posteriore del sesto urotergo libero del maschio, è quasi acuminato (arcuato nella
nuova specie). Di brevicornis è noto il solo holotypus femmina.
ETIMOLOGIA. Specie dedicata al Dr Luca Bartolozzi del Museo Zoologico de
"La Specola" dell'Università di Firenze, che l'ha raccolta.
DIESTOTINI
Cyllogerusa gen. n. (Figg. 62-67)
Corpo lucidissimo e convesso; addome a lati paralleli, fig. 62; tempie
robustamente marginate; palpi labiali di 2 articoli; ligula molto lunga, intera e con due
papille apicali; paraglosse molto prominenti in avanti, fig. 67; palpi mascellari di 4
articoli, fig. 66; mento subtrapezoidale, con margine anteriore arcuato all'indietro, fig.
63: mesocoxe lievemente separate tra loro; processo mesosternale insinuato tra esse e
impresso a metà; formula tarsale 4-4-5; primo tarsomero posteriore molto lungo.
TYPUS GENERIS: Cyllogerusa rubicula (Erichson, 1840), ("olim" Hoplandria).
ETIMOLOGIA. Il nome del nuovo genere significa: "Colei che è curva e vecchia".
COMPARAZIONI. Finora la specie rubicola Erichson, 1840, della Colombia, era
attribuita al genere Hoplandria Kraatz, 1857. L'esame del tipo maschio, etichettato
"Colombia, Moritz, rubicula Er., 5496, Typus" (MB), mi ha permesso di escludere
l'attribuzione a Hoplandria. Infatti questo genere presenta formula tarsale 4-5-5,
palpi labiali di 4 articoli e ligula lunga quanto quella della nuova specie, ma divisa
all'apice che presenta lunghe setole. Inoltre due setole sono inserite lateralmente a
metà della ligula, assenti nella ligula del nuovo genere.
Il nuovo genere probabilmente si colloca tassonomicamente presso il genere
Diestota Mulsant & Rey, 1870, da cui differisce per la ligula intera e assai lunga.
Occorrerà rinvenire la femmina del nuovo genere: la forma della sua spermateca
permetterà una sicura attribuzione tribale.
414 ROBERTO PACE
NE
va hae
la
rll yy ye
I 4
Fico. 59-63
Habitus, edeago in visione laterale, sesto urotergo libero del maschio e mento. 59-61:
Brachychara bartolozzii sp. n.; 62-63: Cyllogerusa rubicula (Erichson) gen. n.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 415
Fico. 64-71
Edeago in visione laterale e ventrale, maxilla con palpo mascellare e labio con palpo labiale.
64-67: Cyllogerusa rubicula (Erichson) gen. n.; 68-71: Trichidryas discors (Erichson) gen. n.
416 ROBERTO PACE
Trichidryas gen. n. (Figg. 68-73)
_ Corpo simile al genere Parasilusa Bernhauer, 1908, del Brasile, cioè con
pronoto molto ristretto in avanti e con una fossetta mediana posteriore trasversa, ma
l'addome è più ristretto all'indietro che in Parasilusa. Tempie robustamente mar-
ginate; palpi labiali di 2 articoli molto lunghi; ligula divisa in due rami: all'apice di
ciscuno è inserita una lunga setola, fig. 72; palpi mascellari di 4 articoli, con lobo
interno assai stretto all'apice, privo di lunghi denti, fig. 71; mento largamente incavato
al margine anteriore, fig. 73; mesocoxe largamente separate tra loro poiché i processi
mesosternale e metasternale si insinuano tra esse con larghe appendici, tronca all'
apice quella mesosternale, arrotondata la metasternale; formula tarsale 4-4-5; primo
tarsomero posteriore lungo quanto i due tarsomeri seguenti compresi insieme.
TYPUS GENERIS: Trichidryas discors (Erichson, 1840), ("olim" Homalota
Mannerheim 1831, "postea" Thamiaraea Thomson, 1858).
ETIMOLOGIA. Il nome del nuovo genere significa "(Ramo) di quercia con
setole". Esso allude alla forma della ligula.
COMPARAZIONI. Questo nuovo genere è creato in seguito all'accurato esame del
tipo di discors Erichson, 1840, della Colombia, così etichettato: "Columbia, Caracas,
Moritz, discors Er., Typus" (MB).
Il nuovo genere Trichidryas è tassonomicamente vicino al genere Parasilusa
Bernhauer, 1908, diffuso in Brasile con una sola specie (iheringi Bernhauer, 1908), da
me esaminata. Infatti i palpi labiali sono allungati in entrambi i generi e i denti del
lobo interno delle maxille sono corti pure in entrambi. Ma il nuovo genere è netta-
mente distinto per la ligula lunga a base assai stretta e con due setole apicali, mentre
Parasilusa presenta ligula corta, a base larga, e i due rami apicali sono privi di setole.
Il mento di Parasilusa è molto stretto al lato anteriore, largo nel nuovo genere.
La formula tarsale 4-4-5 esclude la specie discors dal genere Thamiaraea che
presenta formula tarsale 4-5—5.
Polytelusa gen. n. (Figg. 74-78)
Corpo simile a quello del genere Trichidryas, sopra descritto. Tempie robus-
tamente marginate; palpi labiali di 2 articoli corti; ligula corta, divisa in due rami
cortissimi che hanno, ciascuno, una setola apicale, fig. 77; palpi mascellari di 4
articoli, simili a quelli di Trichidryas, ma il secondo articolo è più corto del terzo;
lobo interno all'apice stretto e con fini e distinte spine; mento simile a quello di Tri-
chidryas, ma le setole anteriori sono meno lunghe e le posteriori più lunghe; meso-
coxe largamente separate tra loro; formula tarsale 4-4-5; primo tarsomero posteriore
corto.
TYPUS GENERIS: Polytelusa colombica sp. n.
ETIMOLOGIA. Il nome del nuovo genere significa "Colei che è magnifica".
COMPARAZIONI. Il nuovo genere si pone tassonomicamente in posizione
intermedia tra Parasilusa e Trichidryas. Si distingue per i caratteri dati nella seguente
chiave
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 417
CHIAVE DEI GENERI Parasilusa Bernhauer, Trichidryas gen. n. e Polytelusa gen. n.
BE Bimotarsomeropostenore corto lisula Cona RR ern 2
— Primo tarsomero posteriore lungo quanto 1 due seguenti considerati
INSIEME I QUA ASS ARNO hs CAE Trichidryas gen. n.
2 — Palpi labiali lunghi: il secondo articolo molto più lungo del primo e
molto più stretto; mento più profondamente incavato al lato anteriore;
parte basale della ligula assai larga; 1 due lembi apicali della ligula sono
lunebmigeisenzaisetole apicale eno Parasilusa Bernhauer
— Palpi labiali corti: il secondo articolo più corto del primo e dilatato
all'apice; mento meno profondamente incavato al lato anteriore; parte
basale della ligula stretta; i due lembi apicali della ligula sono cor-
tissimi e con una setola apicale ciascuno. ............... Polytelusa gen. n.
Polytelusa colombica sp. n. (Figg. 74-78)
Tipi. Holotypus d, Kolumbien, Nördl. Sierra Nevada de Sta. Marta, San Lorenzo, 2200
m, 28.IV.1986, aus niedriger Vegetation am Wegrand, (Miiller leg., MG).
Paratypi: 1 d e 1 ®, stessa provenienza; 1 d, idem, ma Nordwestl. Sierra Nevada de
Sta. Marta, nahe San Pedro de la Sierra, 12.1V.1986, aus Fall-Laub, (Müller leg.).
DESCRIZIONE. Lunghezza 3,0 mm. Corpo lucido e bruno; pronoto giallo-ros-
siccio; antenne brune con i quattro antennomeri basali giallo-rossicci; zampe giallo-
rossicce. La reticolazione dell'avancorpo è assente, quella dell'addome è distinta, ma
sugli uroterghi liberi quarto e quinto è vigorosa. La punteggiatura del corpo è netta e
profonda. Edeago figg. 75-76, spermateca fig. 78.
Gansia colombiensis sp. n. (Figg. 79-80)
Tipo. Holotypus ©, Kolumbien, Nördl. Sierra Nevada de Sta. Marta, nähe El Campano,
ca. 1000 m, 20.1V.1986, aus Fall-Laub, (Müller leg., MG).
DESCRIZIONE. Lunghezza 2,9 mm. Corpo lucidissimo e giallo-rossiccio; capo
nero; elitre bruno-rossicce; antenne nere con i quattro antennomeri basali e l'undi-
cesimo giallo-rossicci; zampe rossicce. Su tutto il corpo non vi è traccia di reti-
colazione. La punteggiatura del capo è svanita, quella del pronoto è distinta. Il solco
mediano del pronoto è allargato all'indietro. Tubercoletti salienti a raspa stanno sulla
superficie delle elitre; sull'addome i tubercoletti sono superficiali. Spermateca fig. 80.
COMPARAZIONI. Specie distinta da G. varicornis (Sharp, 1876) dell'Amazzonia,
per avere il capo nero e largo quanto il pronoto (capo rossiccio e più largo del pronoto
in varicornis). La nuova specie è pure distinta da G. bicolor Sharp, 1883, del
Guatemala e da G. antennaria Bernhauer, 1920, del Brasile, per avere i penultimi
antennomeri nettamente trasversi, e da G. tibialis Sharp, 1883, pure del Guatemala,
per avere un profondo solco mediano del pronoto (largamente, ma indistintamente
impresso in tibialis).
Gansia ecuadorensis sp. n. (Figg. 81-82)
Tri. Holotypus 9, Ecuador, Playa Tanga b. Guayaquil, IV.1975, (H. Franz leg., CR).
Paratypus: 1 ®, stessa provenienza.
418 ROBERTO PACE
Fico. 72-78
Labio con palpo labiale, mento, habitus, edeago in visione laterale e ventrale e spermateca.
72-73: Trichidryas discors (Erichson) gen. n.; 74-78: Polytelusa colombica gen. n., sp. n.
419
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR
=
FIGG. 79-82
Habitus e spermateca. 79-80: Gansia colombiensis sp. n.; 81-82: Gansia ecuadorensis sp. n.
420 ROBERTO PACE
DESCRIZIONE. Lunghezza 3,2 mm. Corpo lucido. Capo rossiccio; pronoto
giallo-rossiccio; elitre bruno-rossicce; addome giallo con uriti liberi terzo e quarto
bruno-rossicci; antenne giallo-rossicce con gli antennomeri quinto a decimo bruni e
l'undicesimo giallo paglierino; zampe giallo-rossicce. L'intero corpo è senza retico-
lazione. La punteggiatura del capo è fine e svanita, quella del pronoto e delle elitre è
distinta. Spermateca fig. 82.
COMPARAZIONI. Specie simile a G. varicornis (Sharp, 1876), dell'Amazzonia,
ma distinta per gli angoli posteriori del pronoto non prominenti all'infuori, come in
varicornis, e per le zampe interamente giallo-rossicce (zampe medie e posteriori con
base delle tibie picea in varicornis).
Plesiomalota (s. str.) puthzi sp. n. (Figg 83-86)
Tipi. Holotypus & , Kolumbien, Nördl. Sierra Nevada de Sta. Marta, San Lorenzo, 2200
m, 20.1V.1986, aus Fall-Laub und Moos, (Müller leg., MG).
Paratypi: 7 es., stessa provenienza.
DESCRIZIONE. Lunghezza 2,4 mm. Corpo lucidissimo e nero; antenne nere con
l'antennomero basale bruno e l'undicesimo giallo-rossiccio; zampe rossicce con
femori bruni. La reticolazione del capo e dell'addome è svanita, quella del pronoto e
delle elitre è estremamente superficiale. La punteggiatura del capo è ombelicata, netta
e assente sulla fronte, quella del pronoto è indistinta e quella delle elitre è netta.
Edeago figg. 88-89, spermateca fig. 90.
COMPARAZIONI. Specie che in base alla forma dell'edeago si pone tassono-
micamente accanto a P. incaica Pace, 1986, del Perù. Se ne distingue per l'edeago più
sviluppato; con l'appendice ventrale triangolare e non stretta e lunga come in incaica.
ETIMOLOGIA. Specie dedicata al Dr Volker Puthz di Schlitz, noto studioso di
Steninae.
Plesiomalota (s. str.) mendax sp. n. (Figg. 87-90)
Tipi. Holotypus d, N. Kolumbien, nordwestl. Sierra Nevada de Sta. Marta, ca. 1000 m,
nahe San Pedro de la Sierra, 12.1V.1986, aus Fall-Laub, (Müller leg., MG).
Paratypi: 10 es., idem, nähe El Campano, ca. 1000 m, 20.1V.1986, aus Fall-Laub
(Miiller leg.).
DESCRIZIONE. Lunghezza 1,9 mm. Corpo lucido e rossiccio scuro; capo bruno
rossiccio; antenne brune con i due antennomeri basali e l'undicesimo giallo-rossicci;
zampe rossicce. La reticolazione del capo e dell'addome è distinta, quella del pronoto è
vigorosa e quella delle elitre è svanita. La punteggiatura del capo è ombelicata e netta,
quella del pronoto è assente, quella delle elitre è netta e irregolarmente distribuita.
Tubercoletti svaniti stanno sugli uroterghi. Edeago figg. 88-89, spermateca fig. 90.
COMPARAZIONI. La presenza di una lunga lamina tronca all'apice che prende
origine dalla "crista apicalis" dell'edeago, rende unica la nuova specie.
Plesiomalota (s. str.) muelleri sp. n. (Figg. 91-93)
Tipo. Holotypus 4, N. Kolumbien, nordwestl. Sierra Nevada de Sta. Marta, nahe San
Pedro de la Sierra, 12.1V.1986, aus Fall-Laub, (Müller leg. , MG).
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 421
FıGG. 83-90
Habitus, edeago in visione laterale e ventrale e spermateca. 83-85: Plesiomalota (s. str.) puthzi
sp. n.; 87-90: Plesiomalota (s. str.) mendax sp. n.
422 ROBERTO PACE
DESCRIZIONE. Lunghezza 1,9 mm. Corpo lucido e bruno-rossiccio; capo bruno;
base dell'addome giallo-rossiccia; antenne brune con i due antennomeri basali e
l'undicesimo giallo-rossicci; zampe gialle. La reticolazione del pronoto è vigorosa,
quella delle elitre netta e quella dell'addome svanita. La punteggiatura del capo è
ombelicata e distinta, quella del pronoto è assente e quella delle elitre è netta. Edeago
figg. 92-93.
COMPARAZIONI. Specie che in base alla forma dell'edeago sembra affine a P.
venezuelana Pace, 1985. Se ne distingue per l'edeago meno sviluppato, privo di una
vistosa piastra del sacco interno e non ampiamente ricurvo al lato ventrale, come in
venezuelana.
ETIMOLOGIA. Specie dedicata al suo raccoglitore, il Dr H.G. Miiller di Giessen,
studioso di "Pantopoda".
Plesiomalota (Heteromalota) laminata sp. n. (Figg. 94-97)
Tipi. Holotypus 4, Nord-Kolumbien, Dept. Magdalena, nòrdl. Sierra Nevada de Sta.
Marta, nähe San Lorenzo, ca. 1700 m, 25.VIII.1985, aus eigenrotten Blättern, (Müller leg.
MG).
Paratypi: 2 2 2, Kolumbien, Sierra Nevada de Sta. Marta, nähe San Lorenzo, 1700 m,
25.VIII.1985, aus Moos und Streu und Wegrand, (Müller leg.).
DESCRIZIONE. Lunghezza 2,1 mm. Corpo lucido. Capo, elitre e uriti liberi terzo
e quarto e base del quinto, bruni; pronoto rossiccio; base ed estremita addominale
giallo-rossicce; antenne nere con antennomero basale e undicesimo giallo-rossicci;
zampe gialle. La reticolazione del capo é distinta solo sulla fronte, quella del pronoto
è vigorosa, quella delle elitre è ben distinta e quella dell'addome è svanita. La
punteggiatura del capo è ombelicata e netta, quella del pronoto non è visibile e quella
delle elitre è profonda. Edeago figg. 95-96, spermateca fig. 97.
COMPARAZIONI. La lamina ad apice arrotondato, che prende origine dalla
"erista apicalis" dell'edeago, è carattere sufficiente a distinguere la nuova specie dalle
note.
Plesiomalota (Microusa) cara sp. n. (Figg. 98-101)
Tripi. Holotypus d, Ecuador, Playa Tanga, b. Guayaquil, IV.1975; (H. Franz leg., CFR).
Paratypi: 36 es., stessa provenienza.
Descrizione. Lunghezza 1,8 mm. Avancorpo debolmente opaco, addome
lucido. Corpo bruno-rossiccio; capo bruno; antenne brune con antennomero basale
rossiccio scuro; zampe giallo-rossicce. La reticolazione del pronoto è vigorosa:
ciascuna maglia appare come un tubercoletto, ciò dà un aspetto rugoso alla superficie.
La reticolazione delle elitre è svanita, quella dell'addome è pure svanita, ma a maglie
trasverse. La punteggiatura del capo è fitta e profonda. I tubercoletti che coprono le
elitre sono distinti, quelli dell'addome sono poco salienti. Edeago figg. 99-100,
spermateca fig. 101.
COMPARAZIONI. Il corpo della nuova specie è simile, ma di lunghezza
lievemente maggiore, rispetto quello di P. roedingeri (Bernhauer, 1951), del Perù. I
caratteri differenziali più evidenti sono nell'edeago e nella spermateca. L'edeago della
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 423
Fico. 91-97
Habitus, edeago in visione laterale e ventrale e spermateca. 91-93: Plesiomalota (s. str.)
muelleri sp. n.; 94-97: Plesiomalota (Heteromalota) laminata sp. n.
424 ROBERTO PACE
nuova specie è privo di una plica ventrale, presente nell'edeago di roedingeri e la
spermateca della nuova specie descrive un numero molto maggiore di spire (2-3 spire
in roedingeri).
ETIMOLOGIA. Dalla popolazione Cara, che verso il 980 d. C. conquistò Quito.
Plesiomalota (Chaetopiezusa) tsantsaniana sp. n. (Figg. 102-105)
Tipi. Holotypus d , Ecuador, Playa Tanga, b. Guayaquil, IV.1975, (H. Franz leg., CFR).
Paratypus: 1 ©, stessa provenienza.
DESCRIZIONE. Lunghezza 2,8 mm. Corpo un po' appiattito. Avancorpo opaco,
addome lucido. Corpo nero pece; elitre di un giallo sporco; uriti liberi sesto e settimo
bruno-rossicci; antenne brune con i due antennomeri basali e la base del terzo rossicci;
zampe bruno-rossicce. La reticolazione del capo è netta, quella del pronoto è quasi
vigorosa e quella delle elitre e dell'addome è distinta. La punteggiatura del capo è
ombelicata e contigua. Il pronoto presenta una larga depressione mediana. Le elitre non
hanno distinti tubercoletti sulla superficie. Edeago figg. 103-104, spermateca fig. 105.
COMPARAZIONI. La nuova specie è chiaramente distinta da P. venezuelanorum
Pace, 1985, del Venezuela. Infatti l'edeago della nuova specie è poco ricurvo al lato
ventrale e la spermateca è composta da un numero minore di spire.
ETIMOLOGIA. La nuova specie prende nome dal "tsantsa", la testa del nemico
tagliata e ridotta nelle dimensioni dagli Jivaro.
Plesiomalota (Chaetopiezusa) bartolozzii sp. n. (Figg. 106-109)
Tipi. Holotypus d, Ecuador, Napo Rio Hollin, 1200 m, 9-13.11.1993, (L. Bartolozzi
leg., N° 9855, MF).
Paratypus: 1 /, stessa provenienza.
DESCRIZIONI. Lunghezza 1,8 mm. Corpo un po' depresso e lucido. Capo ed
elitre bruni; pronoto ed addome giallo-brunicci; antenne bruno-rossicce con undi-
cesimo antennomero rossiccio; zampe gialle. La reticolazione del capo è estrema-
mente svanita, quella del pronoto e delle elitre è assente, quella dell'addome è svanita.
La punteggiatura del capo è netta e fitta. Il pronoto è coperto di tubercoletti fitti e fini.
Le elitre non presentano punteggiatura, nè tubercoletti. Gli uroterghi presentano
tubercoletti distinti sulla loro superficie. Edeago figg. 107-108, spermateca fig. 109.
COMPARAZIONI. L'edeago della nuova specie è poco ricurvo al lato ventrale e
ha l'apice lievemente incavato, in visione ventrale, perciò la nuova specie è chiara-
mente distinta da P. venezuelanorum Pace, 1985, come dalla nuova specie sopra des-
critta P. tsantsaniana.
ETIMOLOGIA. La nuova specie & dedicata al suo raccoglitore, il Dr Luca
Bartolozzi del Museo Zoologico de "La Specola" di Firenze.
Plesiomalota (Chaetopiezusa) mucronata sp. n. (Figg. 110-112)
Tipo. Holotypus d, Ecuador, Napo Rio Hollin, 1200 m, 9-13.11.1993, (L. Bartolozzi
leg., N° 9856, MF).
DESCRIZIONE. Lunghezza 1,9 mm. Capo e pronoto lievemente opachi, resto del
corpo lucido. Capo, pronoto e uriti liberi terzo, quarto e quinto bruni; elitre nere;
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 425
101
FicG. 98-105
Habitus, edeago in visione laterale e ventrale e spermateca. 98-101: Plesiomalota (Microusa)
cara Sp. n.; 102-105: Plesiomalota (Chaetopiezusa) tsantsaniana sp. n.
426 ROBERTO PACE
01 mm
Ficc. 106-112
Habitus, edeago in visione laterale e spermateca. 106-109: Plesiomalota (Chaetopiezusa)
bartolozzii sp. n.; 110-112: Plesiomalota (Chaetopiezusa) mucronata sp. n.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 427
antenne bruno-rossicce con l'antennomero basale giallo e l'undicesimo giallo-
rossiccio; zampe gialle. Il capo e il pronoto mostrano una reticolazione vigorosa e una
punteggiatura poco distinta. La reticolazione delle elitre è distinta, quella dell'addome
svanita. Le elitre sono coperte di tubercoletti nettamente salienti. Il quinto urosterno
libero, al margine posterior ha una formazione appuntita che ha suggerito il nome
della specie.
COMPARAZIONI. L'edeago di dimensioni ridotte, rispetto quello delle specie del
sottogenere e la presenza di una spina mediana posteriore del quinto urosterno libero
del maschio, sono caratteri che distinguono chiaramente la nuova specie dalle note.
Plesiomalota (Chaetopiezusa) bilobifera sp. n. (Figg. 113-117)
Tipi. Holotypus d, Ecuador, Playa Tanga, b. Guayaquil, IV.1975, (H. Franz leg., CFR).
Paratypi: 6 es., stessa provenienza.
DESCRIZIONE. Lunghezza 2,5 mm. Avancorpo debolmente lucido, addome
lucido; capo e quarto urite libero, bruni; pronoto, elitre e uriti liberi terzo e quinto
bruno-rossicci; uriti liberi primo e secondo ed estremità addominale giallo-rossicci;
antenne brune con i due antennomeri basali e l'undicesimo giallo-rossicci; zampe
giallo-rossicce. La reticolazione del capo, delle elitre e dell'addome è svanita, quella
del pronoto è netta. La punteggiatura del capo è ombelicata e netta, quella del pronoto
è indistinta. Superficiali tubercoletti coprono la superficie delle elitre. Gli uriti liberi
quarto e quinto mostrano una reticolazione a maglie molto trasverse e svanite. Edeago
figg. 114-116, spermateca fig. 117.
COMPARAZIONI. L'edeago più sviluppato, con un lobo a ciascun lato e la
spermateca composta di poche, ma ampie spire, sono caratteri che permettono di
distinguere facilmente la nuova specie da P. venezuelanorum Pace, 1985.
Plesiomalota (Leptosomaphya) cotopaxiensis sp. n. (Figg. 118-121)
Tipi. Holotypus dé, Ecuador, Cotopaxi, 3900-4800 m, (H. Franz leg., CFR).
Paratypi: 2 9 9, stessa provenienza.
DESCRIZIONE. Lunghezza 2,0 mm. Specie attera. Corpo lucido e giallo-ros-
siccio, comprese le zampe e le antenne: solo gli antennomeri basali 1° e 2° e l'un-
dicesimo sono gialli. La reticolazione del capo è netta solo sul disco, quella del
pronoto è nettissima, composta di maglie ampie, quella delle elitre è distinta e quella
dell'addome è svanita. Il capo è coperto di punteggiatura ombelicata distinta. Il pro-
noto presenta una largo solco mediano, più profondo all'indietro ed è privo di pun-
teggiatura o di tubercoletti. Tubercoletti salienti coprono la superfice delle elitre. Il
quinto urotergo libero del maschio mostra una reticolazione distinta. Edeago figg.
120-121, spermateca fig. 119.
COMPARAZIONI. L'edeago della nuova specie, rispetto quello di P. franzi Pace,
1983 del Venezuela, è molto più sviluppato e meno profondamente ricurvo al lato
ventrale. Specie attera, endemica del Cotopaxi.
428 ROBERTO PACE
FIGG. 113-121
Habitus, edeago in visione laterale e ventrale (fig. 116 apice dell'edeago in visione latero—
ventrale) e spermateca. 113-117: Plesiomalota (Chaetopiezusa) bilobifera sp. n.; 118-121:
Plesiomalota (Leptosomaphya) cotopaxiensis sp. n.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 429
Plesiomalota (Microusa) filaria sp. n. (Figg. 122-123)
Tipo. Holotypus 9, Ecuador, Playa Tanga, b. Guayaquil, IV.1975, (H. Franz leg., CFR).
DESCRIZIONE. Lungh. 1,3 mm. Avancorpo opaco, addome lucido. Capo bruno;
pronoto bruno-rossiccio; elitre brune con base bruno-rossiccia; addome rossiccio con
uriti liberi terzo, quarto e base del quinto bruni; antenne bruno-rossicce con 1 due
antennomeri basali giallo-rossicci; zampe giallo-rossicce. La punteggiatura del capo e
del pronoto é fittissima e netta. Le elitre sono prive di punteggiatura e di tubercoletti,
pero presentano una reticolazione distinta. Il capo ha un solco mediano e il pronoto un
largo appiattimento mediano della superficie. Spermateca fig. 123.
COMPARAZIONI. La taglia molto ridotta e gli occhi ben sviluppati, lunghi quanto
le tempie, insieme alla forma della spermateca che ha bulbo distale con un’ appendice
falciforme apicale, sono caratteri sufficienti a rendere unica la nuova specie rispetto
alle specie del Venezuela da me descritte nel 1983.
Plesiomalota (Microusa) riobambensis sp. n. (Figg. 124-127)
Tipi. Holotypus d, Ecuador, Umg. Rio Bamba, IV.1975, (H. Franz leg., CFR).
Paratypi: 8 es., stessa provenienza.
DESCRIZIONE. Lunghezza 1,7 mm. Corpo lucido e rossiccio; elitre bruno-ros-
sicce; antenne bruno-rossicce con 1 tre antennomeri basali e l'undicesimo giallo-
rossicci; zampe gialle. Il capo e il pronoto sono coperti di tubercoletti fra loro con-
tigui. Il pronoto mostra un appiattimento mediano posteriore. Le elitre sono coperte di
tubercoletti e di reticolazione superficiali. Edeago figg. 125-126, spermateca fig. 127.
COMPARAZIONI. La nuova specie ha habitus simile a quello di P. perpusilla
Pace, 1983, del Venezuela, ma l'edeago della nuova specie è profondamente ricurvo al
lato ventrale e non bisinuato al lato ventrale come in perpusilla.
Diestota (Apheloglossa) chibchana sp. n. (Figg. 128-131)
Tipi. Holotypus d, Kolumbien, Punta de Betin, 1-10.V.1986, Farbschalen, (Müller leg.,
MG).
Paratypi: 3 9 9, stessa provenienza, ma in data IV.1986 & 1.11.1986, Barberfallen,
(Miiller leg.).
DESCRIZIONE. Lunghezza 1,9 mm. Corpo lucidissimo. Capo, pronoto e base
dell'addome rossicci; elitre e resto dell'addome nero-bruni; antenne brune con i tre
antennomeri basali e l'undicesimo giallo-rossicci; zampe giallo-rossicce. La retico-
lazione del capo e dell'addome è molto svanita, quella del pronoto è superficiale e
quella delle elitre è netta. La punteggiatura del capo è appena svanita, quella del pro-
noto è distinta e quella delle elitre è superficiale. Edeago figg. 129-130, spermateca
fig. 131.
COMPARAZIONI. L'appendice spatuliforme ventrale dell'edeago permette di
distinguere la nuova specie da tutte le specie del genere, poiché esse non presentano
tale curiosa struttura,
ETIMOLOGIA. Dai Chibcha, antica popolazione colombiana.
430 ROBERTO PACE
FIGG. 122-131
Habitus, spermateca ed edeago in visione laterale e ventrale. 122-123: Plesiomalota (Microusa)
filaria sp. n.; 124-127: Plesiomalota (Microusa) riobambensis sp. n.; 128-131: Diestota
(Apheloglossa) chibchana sp. n.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 431
Diestota (Apheloglossa) ecuadorensis sp. n. (Figg. 132-135)
Tipo. Holotypus dé, Ecuador, Cotopaxi, 3900-4800 m, (H. Franz leg., CFR).
DESCRIZIONE. Lunghezza 2,0 mm. Corpo debolmente lucido e bruno-rossiccio;
addome rossiccio con quarto urite libero bruno; antenne brune con 1 due antennomeri
basali di un giallo sporco; zampe gialle. La reticolazione del capo e del pronoto è
netta, quella delle elitre è distinta. La punteggiatura del capo è fitta. Fitti tubercoletti
coprono la superficie del pronoto e delle elitre. Edeago figg. 133-134, sesto urotergo
libero del maschio fig. 135.
COMPARAZIONI. Specie affine a D. guadalupensis Pace, 1987, a motivo dei
simili caratteri del sesto urotergo libero del maschio. Ne è distinta per l'edeago privo
di ampia incavatura ventrale e non terminante a punta acuta, ma a punta arcuata.
Diestota (Apheloglossa) cotopaxiana sp. n. (Figg. 136-140)
Tipi. Holotypus d, Ecuador, Cotopaxi, S. Francisco de Las Pampas, 1300-1500 m,
11.1993, (L. Bartolozzi leg., N° 9832, MF).
Paratypus: 1 9, stessa provenienza.
DESCRIZIONE. Lunghezza 2,2 mm. Avancorpo debolmente opaco, addome
lucido. Capo, elitre e quarto urite libero neri; pronoto bruno con margini laterali
rossicci; addome rossiccio, tranne il quarto urite libero nero, come indicato sopra;
antenne brune con antennomero basale rossiccio e l'undicesimo giallo-rossiccio con
base rossiccia; zampe giallo-rossicce. L'avancorpo è coperto di reticolazione distinta.
L'addome è privo di reticolazione. La punteggiatura del capo e del pronoto è poco
distinta, confusa nella reticolazione. Tubercoletti distinti stanno sulla superficie delle
elitre. Sesto urotergo libero del maschio fig. 137, edeago figg. 158-159, spermateca
fig. 140.
COMPARAZIONI. Per i caratteri del sesto urotergo libero del maschio, la nuova
specie è affine a D. guadalupensis Pace, 1987. Se ne distingue, oltre che per l'undi-
cesimo antennomero del maschio molto lungo, per l'edeago bruscamente flesso al lato
ventrale, perciò tozzo rispetto quello di guadalupensis e molto più largo di quello di
guadalupensis, in visione ventrale.
HOMALOTINI
Euvira tayronensis sp. n. (Figg. 141-142)
Tipo. Holotypus 2, Kolumbien, Dept. Magdalena, Bahia Ide Gairaca, Tayrona Park, ca.
20 Km NE Sta. Marta, VI.1985, Barberfallen, (Miiller leg., MG).
DESCRIZIONE. Lunghezza 1,4 mm. Corpo lucido e giallo-bruniccio; antenne
brune con i tre antennomeri basali gialli; zampe gialle. La punteggiatura del capo è
composta da punti piuttosto grandi e un po' svaniti. La punteggiatura del pronoto è
svanita, composta di punti fini. Le elitre presentano tubercoletti e reticolazione
svaniti. Spermateca fig. 142.
COMPARAZIONI. Specie simile a E. fervidula (Erichson, 1840) della Colombia,
ma d'aspetto più gracile. In base al confronto della forma della spermateca della nuova
432 ROBERTO PACE
Fısc. 132-140
Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del maschio e spermateca.
132-135: Diestota (Apheloglossa) ecuadorensis sp. n.; 136-140: Diestota (Apheloglossa) coto-
paxiana Sp. n.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 433
specie con quella di una femmina della serie tipica di fervidula, è possibile rilevare
consistenti differenze: presenza di introflessione apicale del bulbo distale della sperma-
teca della nuova specie e assenza del medesimo carattere nel bulbo distale della
spermateca di fervidula; parte mediana della spermateca della nuova specie poco
sviluppata, mentre è molto sviluppata quella della spermateca di fervidula.
Non è il caso di fare comparazioni con E. atratula (Erichson, 1840), pure della
Colombia, perché essa, come ho osservato sui tipi, ha taglia vistosamente maggiore
(2,1 mm) e spermateca priva di introflessione apicale del bulbo distale e parte
mediana della spermateca stessa cortissima.
Placusa (s. str.) chibchana sp. n. (Figg. 143-144)
Tipo. Holotypus 2, Kolumbien, Punta de Betin, Sta. Marta, IV.1986, Farbschalen,
(Miiller leg., MG).
DESCRIZIONE. Lunghezza 2,2 mm. Corpo lucido. Capo e pronoto bruni; elitre
bruno-rossicce con lati esterni e zona periscutellare bruni; addome rossiccio con
quarto urite libero bruno a metà; antenne nere con antennomero basale giallo sporco;
zampe giallo-rossicce. L'avancorpo è coperto di tubercoletti distinti e di reticolazione
svanita. L'addome non presenta reticolazione. Spermateca fig. 144.
COMPARAZIONI. In base alla forma della spermateca e all'habitus, la nuova
specie sembra affine a P. basiventris Pace, 1987, della Guadalupa. I caratteri distintivi
sono: bulbo distale della spermateca oviforme nella nuova specie, mentre è piriforme
in basiventris; assenza di introflessione apicale del bulbo distale della spermateca
nella nuova specie, mentre in basiventris è presente.
ETIMOLOGIA. Dai Chibcha, antica popolazione colombiana.
Placusa (s. str.) guambina sp. n. (Figg. 145-146)
Tipo. Holotypus ©, Kolumbien, Punta de Betin, Sta. Marta, IV.1986, Farbschalen,
(Miiller leg., MG).
DESCRIZIONE. Lunghezza 1,8 mm. Corpo appiattito e bruno-gialliccio; antenne
brune con i due antennomeri basali giallo-bruni; zampe giallo-brune. Una retico-
lazione distinta sta solo sul capo: sul resto del corpo la reticolazione è assente.
L'avancorpo è coperto di tubercoletti salienti. Fini sono i tubercoletti che coprono la
superficie dell'addome. Spermateca fig. 146.
COMPARAZIONI. La nuova specie, in base alla forma della spermateca, appare
simile a P. basiventris Pace, 1984 della Guadalupa. La nuova specie è differente da
questa specie per avere la spermateca più esile, con bulbo distale globoso e non
piriforme come quello di basiventris.
ETIMOLOGIA. Dai Guambinos popolazione caraibica esistente prima della con-
quista spagnola.
Placusa (s. str.) oligotina sp. n. (Figg. 147-148)
Tipo. Holotypus 9, Kolumbien, Villa Culebra bei Bonda, ca. 10 Km E Sta. Marta,
1-11.X1.1985, Trockengebiet, Farbschalen, (Müller leg., MG).
434 ROBERTO PACE
01 mm
144
148
01 mm
Fico. 141-148
Habitus e spermateca. 141-142: Euvira tayronensis sp. n.; 143-144: Placusa (s. str.) chibchana
Sp. n.; 145-146: Placusa (s. str.) guambina sp. n.; 147-148: Placusa (s. str.) oligotina sp. n.
DESCRIZIONE. Lunghezza 1,5 mm. Corpo lucido e rossiccio; capo, elitre, quarto
urite libero e metà basale del quinto, bruni; antenne giallo-rossicce; zampe gialle. La
reticolazione del capo è netta, quella del pronoto e delle elitre è distinta e quella
dell'addome è svanita. Il capo presenta una fossetta frontale, il pronoto tubercoletti
estremamente svaniti e le elitre punteggiatura superficiale. Spermateca fig. 148.
ALEOCHARINAE DELLA COLOMBIA E DELL’ECUADOR 435
COMPARAZIONI. Specie che presenta una fossetta frontale e una spermateca con
bulbo distale ipertrofico e parte mediana della stessa spermateca, esilissima. Questi
caratteri non sono presenti in specie note.
BOLITOCHARINI
Caralepta gen. n. (Figg. 149-155)
Habitus simile alle specie attere di Leptusa Kraatz d'alta quota altimetrica, cioè
con occhi ed elitre assai ridotti. Tempie non marginate; palpi labiali di 3 articoli;
N)
FısG. 149-155
Habitus, edeago in visione laterale e ventrale, spermateca, labio con palpo labiale, maxilla con
palpo mascellare e mento. 149-155: Caralepta cotopaxicola gen. n., sp. n.
436 ROBERTO PACE
ligula divisa fino alla base in due lembi, fig. 153; palpi mascellari di 4 articoli; lobo
interno con spine lunghissime, fig. 154; mento subtrapezoidale, a margine anteriore
rettilineo, fig. 155; mesocoxe contigue; processo mesosternale ad apice acuto, privo di
carene; formula tarsale 4-4-5; primo tarsomero posteriore corto.
TYPUS GENERIS: Caralepta cotopaxicola sp. n.
ETIMOLOGIA. Il nome del nuovo genere significa "Esile dei Caras". Caras è un
regno preincaico dell'Ecuador.
COMPARAZIONI. Il nuovo genere si colloca accanto al genere Thecturota Casey,
1893. Si distingue essenzialmente per avere la ligula divisa fino alla base in due
lembi, mentre in Thecturota è triangolarmente incisa all'apice. La spermateca ha
struttura simile nei due generi, ma è nettamente più robusta in Caralepta gen. n.,
nonostante la taglia delle specie dei due generi sia simile.
Caralepta cotopaxicola sp. n. (Figg. 149-155)
Tri. Holotypus d, Ecuador, Cotopaxi, 3500-4800 m, (H. Franz leg., CFR).
Paratypus: 1 ©, stessa provenienza.
DESCRIZIONE. Lunghezza 1,8 mm. Specie attera. Corpo lucido e giallo-ros-
siccio; addome sfumato di bruno rossiccio nella parte mediana; antenne e zampe
gialle. La reticolazione del capo e del pronoto è netta, quella delle elitre è distinta e
quella dell'addome poligonale e svanita, netta solo sul quinto urotergo libero. La
punteggiatura sul capo e sul pronoto è assente. Tubercoletti assai salienti stanno sulla
superficie delle elitre. Il capo presenta una fossetta discale. Edeago figg. 150-151,
spermateca fig. 152.
RINGRAZIAMENTI
Per il materiale affidatomi in esame, ringrazio molto cordialmente il Dr V.
Puthz della "Limnologische Flusstation" di Schlitz, il Dr L. Bartolozzi del Museo
Zoologico de "La Specola" dell'Università di Firenze, il Prof. Dr H. Franz di Mödling
e il Dr M. Uhlig dell'Università Humboldt di Berlino. Per il prestito di tipi ringrazio i
direttori e i conservatori dei seguenti Istituti: D.E.I. di Eberswalde, Museo Zoologico
dell'Università Humboldt di Berlino, Institut Royal des Sciences Naturelles de
Belgique di Bruxelles, il "Naturhistorisches Museum" di Vienna e il British Museum
(Natural History) di Londra.
BIBLIOGRAFIA
BERNHAUER, M. 1908. Beitrag zur Staphylinidenfauna von Südamerika. Archiv für Natur-
geschichte 74: 283-372.
BERNHAUER, M. 1912. Zur Staphylinidenfauna von Südamerika. Entom. Ztg. Wien 31: 68-82.
BERNHAUER, M. 1920. Neue Aleocharini aus Südamerika. Archiv für Naturgeschichte 86:
141-170.
BERNHAUER, M. 1921. Zur Staphylinidenfauna Südamerika, insbesondere Argentiniens. Archiv
für Naturgeschichte 86: 170-183.
ALEOCHARINAE DELLA COLOMBIA E DELL’ ECUADOR 437
BERNHAUER, M. 1927. Zur Staphylinidenfauna Siidamerika, insbesondere Argentiniens. Archiv
für Natur geschichte 91: 229-264.
BERNHAUER, M. 1934. Neue Staphyliniden aus Argentiniens. Revista de Entomologia do Rio de
Janeiro 3: 501-517.
BERNHAUER, M. 1939. Neue Staphyliniden aus Chile (Coleoptera). Arbeiten über
norphologische und taxonomische Entomologie aus Berlin-Dahlem 6: 12-15.
BERNHAUER, M. 1951 (1941). Staphylinidae (Col.). /n: Beitrag Fn. Perous 1: 277-293.
BERNHAUER, M. & SCHEERPELTZ, O. 1926. Coleopterorum Catalogus 82. Staphylinidae. Berlin
6: 499-988.
BIERING, A. 1934. Novedades de los géneros Oligota Mann. y Euvira Sharp de la fauna
neotropica. Memorias de la Sociedad Cubana de Historia Natural 8: 113-123.
CASEY, T.L. 1893. Coleopterological Notices V. Annals of the New York Academy of Sciences
7: 281-606.
Casey, T.L. 1911. New American Species of Aleocharinae and Myllaeninae. Memoirs on the
Coleoptera 2: 1-245.
ERICHSON, W.F. 1840. Genera et Species Staphylinorum Insectorum Coleopterorum Familiae 2:
Berlin, 954 pp.
FAUVEL, A. 1901. Voyage de M. le Dr Ed. Bugnion au Venezuela, en Colombie et aux Antilles.
Revue d’Entomologie de Caen 20: 69-91.
FRANK, J.H. 1972. The genus Oligota Mannerheim in the Caribbean Region (Coleoptera,
Staphylinidae). The Coleopterists Bulletin 26: 125-146.
KRAATZ, G. 1857. Genera Aleocharinorum Illustrata. Linnaea Entomologica 11: 1-43.
KRAATZ, G. 1862. Synonymische Bemerkungen. Ueber Coleopteren. Berliner entomologische
Zeitschrift 6: 298-300.
MANNERHEIM, C.G. 1830. Précis d'un nouvel arrangement de la Famille des Brachélitres de
l'Ordre des Insectes Coléoptères. Memoires de l'Académie des Sciences de Saint-
Petersbourg 1: 415-501.
MOTSCHULSKY, T.V. 1858. Enumération des nouvelles espèces de Coléoptères rapportés de ses
voyages. Bull. Soc. Imp. Nat. Moscou 31: 204-264.
NOTMAN, H. 1920. Staphylinidae from Florida in the collection of the American Museum of
Natural History, with descriptions of new genera and species. Bulletin of the American
Museum of Natural History 42: 693-732.
PACE, R. 1983. Aleocharinae orofile del Venezuela raccolte dal Prof. Franz. I. Bolitocharini e
Callicerini Geostibae. Bolletino della Società entomologica italiana 115: 91-102.
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versità Humboldt di Berlino. Giornale italiano di Entomologia 1: 295-316.
PACE, R. 1985. Aleocharinae del Venezuela riportate dal Prof. Franz. Giornale italiana di
Entomologia 2: 371-392.
PACE, R. 1986. Aleocharinae del Perù (Coleoptera, Staphylinidae). Redia 69: 417-467.
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Folia Entomologica Hungarica 48: 187-205.
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Staphylinidae). Bolletino del Museo civico di Storia Naturale di Verona 15: 151-162.
SEEVERS, C.H. 1951. A Revision of the North American and European Staphylinid Beetles of
the Subtribe Gyrophaenae (Aleocharinae, Bolitocharini). Fieldiana Zoology 32: 660-
762.
SHARP, D. 1876. Contribution to an insectfauna of the Amazon Valley, Staphylinidae.
Transactions of the Entomological Society of London 1: 27-414.
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Vetenskapsakademiens förhandlinger 15: 27-40.
=
REVUE SUISSE DE ZOOLOGIE, 103 (2): 439-452; juin 1996
Distribution of entomopathogenic nematodes in the Swiss Alps
Werner A. STEINER
Swiss Federal Research Station for Fruit-Growing, Viticulture and Horticulture,
CH-8820 Wädenswil
Distribution of entomopathogenic nematodes in the Swiss Alps. - A
total of 473 soil samples taken from alpine regions in Switzerland was
analysed for the presence of rhabditid entomopathogenic nematodes. These
parasites were found in 27% of the samples at altitudes between 490 and
2530 m asl. The recovered nematode isolates fall into six species:
Steinernema affinis, S. feltiae, S. intermedia, S. kraussei, Steinernema sp.
(a species close to S. intermedia), and Heterorhabditis sp. (North-West
European Group). The distribution of these species is discussed in relation
to altitude, vegetation, orientation, soil characteristics (1.e. pH, sand
content, content of organic matter), and potential host species. Steinernema
kraussei was the most commonly encountered species, usually occurring in
acidic soils rich in organic matter. Steinernema feltiae, the second most
prevalent species, was rather confined to grassland habitats of the lower
Alps and the Swiss Plateau. Similarly, the other species were usually found
in grassland, but were too rare to draw conclusions about their ecological
requirements.
Key-words: distribution - environmental characteristics - cold-activity -
steinernematids - Steinernema kraussei - feltiae - Heterorhabditis.
INTRODUCTION
Nematodes of the genera Steinernema (Steinernematidae: Rhabditida) and
Heterorhabditis (Heterorhabditidae: Rhabditida) are obligate parasites of soil-dwell-
ing insects. These nematodes include in their life cycle free-living infective juveniles,
able to locate and infect suitable host species. Several strains of Steinernema and
Heterorhabditis are commercially sold as biological control agents of various insect
pests. However, currently available strains can only be used effectively above
temperatures of ca 12 °C (e.g. Wouts 1991). Problems with the poor cold-activity of
entomopathogenic nematodes are common to all countries of temperate zones. To
enhance biological control of outdoor insect pests at low temperatures, cold-active
nematodes are required. Such nematodes are likely to be found in the Swiss Alps,
since alpine animals are adapted to long and cold winters.
Manuscript accepted 25.07.1995.
440 WERNER A. STEINER
The faunistic data presented here refer to a field survey realised as part of
COST Action 812 ("Selection and evaluation of cold-active lines of insect-parasitic
nematodes for outdoor application"; see EHLERS 1994). Although the present survey
was designed primarily to isolate cold-active nematodes, the data available allow us to
investigate the distribution of entomopathogenic nematodes in the alpine regions of
Switzerland. Moreover, examination of the influence of sample characteristics on
nematode prevalence provides insight into the ecology of rhabditid entomopathogenic
nematodes.
MATERIAL AND METHODS
A total of 472 soil samples was taken in 1991 at different altitudes in the Swiss
Alps. Sampling locations were chosen in order to collect nematodes over a wide range
of the Swiss Alps and of some adjacent regions. Sampling in the lower and alpine
regions was performed between May and July, and between August and September,
respectively. One additional sample was obtained in May 1994 from a small survey in
the framework of COST 813 (JENNY 1994). Each sample consisted of 10 subsamples (to
a depth of 10 - 15 cm), taken at regular intervals along a transect of ca 50 m. Sub-
samples were pooled, mixed and a 1 kg portion of the pooled sample was retained to
check the presence of entomopathogenic nematodes. At sampling locations below the
timberline two samples were generally taken (i.e. one sample within the forest, one on
the outside), whereas above the timberline only one sample was taken per location.
Entomopathogenic nematodes were baited in the laboratory (at 18 °C) with the wax
moth (Galleria mellonella L.). For each sample, five late instar larvae, placed on the
bottom of a 1-litre plastic pot, were carefully covered with damp soil. After 5 days, the
insect larvae were recovered, and placed into plastic dishes on moist filter paper.
Nematodes emerging from the G. mellonella larvae of each sample (1.e. an isolate) were
used to establish laboratory cultures. All isolates were identified using morphological
criteria (POINAR 1990; MRACEK 1994), and 35 selected isolates also by restriction
fragment length patterns (RFLP of steinernematids by A. Reid, GB-St Albans, Herts;
RFLP of the heterorhabditid isolate by P. Smits, NL-Wageningen).
For each sampling location, the following quantitative (a) and qualitative (b)
characteristics were recorded: a) altitude, pH, content of organic matter (%C), and sand
content; b) region (based on main mountain ranges), orientation (two classes: S-W;
NW-SE), vegetation types (woodland samples with subclasses: deciduous, mixed,
coniferous with spruce, coniferous with larch; grassland samples: pasture or hay,
pasture in vineyard, flowery meadows: "others": home gardens, dwarf shrubland,
rosette plants). The pH and %C of soil samples was determined electrometrically in
distilled water (using a glass electrode) and by a modification of the Walkley Black
method, respectively. The sand content of soil samples was estimated by feel. The
weather in 1991 presented generally average conditions, but May was relatively cool
and the period between August and September relatively warm and dry
(SCHWEIZERISCHE METEOROLOGISCHE ANSTALT 1994).
ENTOMOPATHOGENIC NEMATODES 441
Ordination and classification of positive samples (entomopathogenic nema-
todes present) were performed according to similarity in vegetation (two classes:
woodland as opposed to grassland and "others") and environmental characteristics
(i.e. pH, altitude, sand content, %C, and orientation) using correspondence analysis
and complete linkage clustering on square root transformed data (program MULVA-
4; see WILDI & ORLOCI 1990), respectively. The Mann-Whitney U-test or the Kruskal-
Wallis-test were used to compare rank sums of continuous variables achieved in
nominal classes (samples or species). x? contingency table analyses were used to test
whether entomopathogenic nematodes occur more frequently in certain classes of
samples (1.e. vegetation, orientation, pH, altitude, %C, sand content) than in others.
For rare species, the vegetational specificity was analysed with respect to two
vegetation types only (i.e. woodland as opposed to grassland and "others"). All
comparisons were corrected for ties and used a 0.05 level of significance, unless
otherwise indicated. For multiple analyses on the same data set, the Bonferroni
correction was applied for critical probabilities (SACHS 1992).
RESULTS
The following species of entomopathogenic nematodes were identified (fig. 1):
Steinernema affinis (Bovien, 1937), S. feltiae (Filipjev, 1934), S. intermedia (Poinar,
1985), S. kraussei (Steiner, 1923; sensu MRACEK 1994), Steinernema sp., and
Heterorhabditis sp. (North-West European Group, sensu SMITS et al. 1991). The latter
two species were found only once. Steinernema kraussei was clearly the most
frequent species (52% or 67 isolates). The isolates denoted as Steinernema spp. could
not be identified to species, mainly due to contamination and subsequent loss of initial
cultures.
|_| Heterorhabditis sp.
PTL U
ae
Steinernema affinis
4 5
ASIA
HESS LS Seok
EE M Steinernema feltiae
Il Steinernema intermedia
| ] Steinernema kraussei
M Steinernema Sp.
&& Steinernema spp.
52%
Fic. 1
Species representation of rhabditid entomopathogenic nematodes (n =128) in the Alps and
lower Alps of Switzerland. Steinernema spp. denote unidentified isolates.
442 WERNER A. STEINER
The overall percentage of soil samples yielding entomopathogenic nematodes
was 27% (128 isolates in 473 samples). However, nematode frequencies differed
considerably among regions, varying between 0.5 and 53% (tab. 1). The variation in
prevalence most probably reflects regional differences in sample characteristics,
because altitude, pH, %C, and sand content were unevenly distributed over the
regions (Kruskal-Wallis-test, n = 444, p < 0.001). For example, there were no low-
land samples (< 1000 m) in the regions R6, R8, and R9 (tab. 1).
TAB. 1
Recovery rates of rhabditid entomopathogenic nematodes in various geographical regions (R1
to R9) of the Swiss Alps related to altitude (asl) of the sampling locations. Regions were
defined based on main mountain ranges (see fig. 2).
Regions Altitude No of
No of Recovery
Median [m] Range [m] isolates samples rate [%]
R1: Northern Jura Mountains 805 630-895 2 18 11
R2: Eastern lower Alps 14415 805-1370 4 52, 8
R3: Central to eastern lower Alps 850 470-1180 3 60 05
R4: Central lower Alps 1130 785-1800 8 78 10
RS: Central to eastern Alps 1270 490-2050 17 52 53
R6: Eastern Alps 1810 1095-2330 39 We 93
R7: Central to southern Alps 1770 720-2320 22 56 39
R8: Central to western Alps 2215 1367-3090 19 45 42
R9: Western Alps 1830 1142-2460 14 39 36
R1 - R9: All regions 1320 470-3090 128 473 27
The regions RI-R4 were invariably characterised by recovery rates clearly
below the overall average of 27% (tab. 1). Conversely, the high proportion of positive
samples in the five alpine regions (i.e. R5-R9) suggests that entomopathogenic
nematodes are very common in the mountain range. The geographical distribution of
the 103 isolates identified to the species level is illustrated in fig. 2. The predominant
S. kraussei is restricted to the alpine regions, whereas S. affinis, S. feltiae, and S.
intermedia are more widely distributed. The two species Steinernema sp. and Hete-
rorhabditis sp. were too rare to interpret their geographical distribution.
The majority of the isolates was recovered in soil samples taken at altitudes
between 1500 and 2100 m (fig. 3A). The most elevated recovery site (i.e. with S.
kraussei) was located at 2530 m. Visual examination of fig. 3A suggests that S.
kraussei is the most important species in the alpine environment (mean altitude: 1800
m, 1045-2530 m; median and range). The isolates of S. kraussei were unevenly
distributed with respect to altitude (x2 = 98, DF = 7, p < 0.0001). This was largely
due to their high prevalence in altitudinal classes between 1650 and 2550 m, and an
absence at altitudes below 1000 m. Steinernema kraussei occurred at more elevated
sites than both S. feltiae (U-test, p < 0.001), which predominated in the lower Alps
ENTOMOPATHOGENIC NEMATODES 443
Heterorhabditis sp.
Steinernema affinis
Steinernema feltiae
Steinernema intermedia
Steinernema kraussei ea a
+ X U & © *X
Steinernema sp. ar
Fic. 2
Distribution of rhabditid entomopathogenic nematodes (n = 103) isolated in the Swiss Alps, in
the lower Alps, and in the Jura Mountains. RI to R9 refer to the regions explained in tab. 1.
Note that isolates from nearby sampling locations are hardly distinguishable.
40 50 30
n Koy re
2 304 SII RO
n © =
© 10
2 S 30 2
‘o 20 > o à
o © 20 | pa
2 3 10 à
= 3 io :
3 c -20 |
È rit 0 i z 5 7 = — -30
600 900 1200 1500 1800 2100 2400 600 900 1200 1500 1800 2100 2400
Altitude (m) Altitude (m)
Fic. 3
A) Altitudinal distribution of alpine entomopathogenic nematodes identified at the species level
(n = 103). For graphical patterns see fig. 1. B) Altitudinal variation of the recovery rate of total
entomopathogenic nematodes (n = 128) and the zones of variation in air (light grey) and soil
temperature (dark grey). Temperature was measured in 1991 at seven climatological stations in
Switzerland (SCHWEIZERISCHE METEOROLOGISCHE ANSTALT, 1994). mi: Air temperature (top:
mean of the warmest month; middle: annual mean; bottom: mean of the coldest month); O:
Soil temperature at a depth of 5 cm (top, middle, and bottom: see above), values for locations
above 1800 m are extrapolated (dashed lines); LJ: recovery rate of entomopathogenic nema-
todes related to altitudinal classes.
444 WERNER A. STEINER
(1255 m, 580-2170 m), and S. intermedia (U-test). No difference was observed in the
altitudinal distribution between the other species. Steinernema intermedia (1440 m,
605-2205 m) and S. affinis (1552 m, 490-2080 m) appear to be uniformly distributed
within the zone between ca 600 and 2200 m. Heterorhabditis sp. and Steinernema sp.
were recovered from sites at 1130 and 1530 m, respectively.
The variation in total entomopathogenic nematodes (unidentified isolates
included) and temperature changes along the altitudinal gradient are shown in fig. 3B.
Values of soil temperature in the coldest months indicate that soil-dwelling animals
are well protected from temperatures below the freezing point, irrespective of altitude
and associated air temperatures. In the warm season, however, soil temperature
decreases at a rate of 1.5 °C per altitudinal class. Therefore, soil-dwelling animals are
subjected to similar winter temperatures, whereas during the other seasons less
elevated habitats are characterised by considerably higher temperatures than alpine
areas. Divergent altitudinal profiles of recovery rate and temperature suggest that the
presence of entomopathogenic nematodes depends on factors other than temperature.
The environmental altitudinal gradient represents a complex gradient of tem-
perature (fig. 3B) and other factors related to elevation (e.g. humidity, soil formation).
In the present study, for example, altitude was significantly correlated with pH (r, = -
0.51, p < 0.001), %C (rs = 0.21, p < 0.001), sand content (r, = 0.44, p < 0.001), and
vegetation (Kruskal-Wallis test, p < 0.001). Since the nematodes are expected to
respond simultaneously to various of these factors, the distribution of the nematodes
was analysed using multivariate methods.
Fig. 4 shows the environmental similarity of the positive samples (n = 125)
based on correspondence analysis and complete linkage clustering. Characteristics of
most samples change along a continuum with respect to both axes, and only a few
samples are divergent. The main gradient along the first axis represents the transition
from sandy soils ("s", to the left) to soils rich in organic matter ("C", to the right). The
second axis, accounting mainly for the distinction between S. kraussei and the other
LAI 11
species, represents a shift from low land samples with relatively high pH levels ("p",
top) towards acidic alpine samples ("a", bottom). Vegetation differentiates samples
along both axes. Grassland samples ("g", as opposed to woodland samples) are
associated with comparatively high pH values, a high sand content, and occur usually
at the less elevated sites. The orientation of the sampling locations ("N") is only
slightly associated with other sample characteristics. The proximity of "N" to "a",
however, indicates that at elevated sampling locations the NW-SE oriented slopes
have a slightly higher nematode prevalence than south-facing slopes.
The comparison of species occurrence with sample characteristics within G1 to
G5 (tab. 2) provides information about the relative position of each species in the
niche defined by the six variables shown in fig. 4. Group 1 (G1) is heterogeneous with
respect to entomopathogenic nematodes as it includes all species (except Steinernema
sp.). The dominating species is S. kraussei, representing 67% of the identified iso-
lates. Samples of Gl are characterised by a high organic matter content, and a low
sand content. Furthermore, this group includes the majority (1.e. 56%) of all the
positive samples taken in woodland soils. In G2, the predominant species is S. feltiae,
ENTOMOPATHOGENIC NEMATODES 445
AXIS 2
Fic. 4
Ordination and classification of positive samples (n = 128) by correspondence analysis and
complete linkage clustering, respectively. Samples are characterised by vegetation ("g": grassland
and "others"; as opposed to woodland) and environmental characteristics ("a": altitude; "C":
organic matter content; "p": pH; "s": sand content; "N": orientation NW-SE; as opposed to S-W).
The eigenvalues of the first two axes are 0.40 and 0.29, respectively. Steinernema spp: @; for
other species see fig. 2. Gl to GS: group identification. For clarity, the true boundary of G3 was
omitted. The S. feltiae isolate located between "s" and "N" is a member of G4.
accompanied by one isolate of S. intermedia and S. affinis, each. All samples of G2
were taken from grassland habitats at low altitudes and were characterised by
relatively high pH values. Steinernema feltiae has its main representation in G2 and is
thus considered the most adapted species to grasslands in the lower Alps. Note that
grassland samples had generally higher pH values (U-test), a higher sand content (U-
test, p < 0.001), and a lower amount of C% (U-test, p < 0.001) than woodland
samples.
446 WERNER A. STEINER
TAB. 2
Environmental characteristics of the samples belonging to the groups G1 to G5, that were
revealed by correspondence analysis and complete linkage clustering (see fig. 4).
Environmental characteristics Gl G2 G3 G4 G5
pH (median) 5.5 7.1 6.0 4.8 5.6
(range) (4.1-7.3) (5.4-7.8) (4.3-8.0) (4.3 - 5.8) (4.4 - 7.5)
Altitude (median) 1565 1100 1800 1917 1765
(range) (970 - 1980) (580 - 1670) (970 - 2270) (1610 - 2300) (890 - 2530)
Sand content (median) 1 1 1 2 3
(range) (022) (0-4) (0-3) (1-4) (ES)
Organic content (median) (8 5.8 5.8 TA 4.5
(range) (5.3 -30.9) (2.8-7.9) (2.3-13.0) (4.6 - 10.4) (1.4 - 7.7)
Vegetation (No. of samples) 13/18 13/0 2515 14/5 28/6
grassland / woodland
Orientation (No. of samples) 4/27 3/10 18/10 1/18 3/31
S-W/NW-SE
The samples of G3 include all the species except Heterorhabditis sp., with S.
kraussei the dominating species (71%). G3 is a transitional group with a majority of
grassland samples, characterised by average values for most characteristics except
orientation. About 68% of all the samples taken from south-facing slopes (S to W) are
included in G3. Note that the overall recovery rate of entomopathogenic nematodes
was not the same for the eight radii (x? = 24.1, DF = 7, p < 0.005), with a lower
prevalence than expected on slopes with a southern orientation. G4 consists of
samples yielding almost exclusively S. kraussei (94%), accompanied by one S.
feltiae isolate. Samples of this group were taken at high altitudes and are characterised
by low pH values, as well as by a relatively high sand content. The percentage of
woodland samples in G4 corresponds to their overall rate in this survey. G5 is
dominated by S. kraussei (67%) and S. intermedia (21%). The latter species has its
main representation (1.e. 58%) in this group, and its ecological requirements are
presumably similar to those encountered in samples of GS. Steinernema affinis and S.
feltiae are represented in G5 by three and one isolates, respectively. Samples were
taken mainly from grassland soils characterised by intermediate pH values, a high
sand content, and a low organic matter content.
A more profound insight into the ecological requirements of relatively frequent
species can be obtained by comparing positive with negative samples. For S.
kraussei, this comparison shows that its presence in soil samples is related to
orientation, soil characteristics, and vegetation. Steinernema kraussei was more
prevalent at NW-SE facing slopes than at sampling locations exposed to S-W (x2=
10.4, DF = 1, p < 0.005). Moreover, positive samples differed from negative ones by
ENTOMOPATHOGENIC NEMATODES 447
a higher content of both organic matter (U-test) and sand (U-test, p < 0.001 ), as well
as by lower pH values (U-test, p < 0.001). The mean pH level of samples yielding S.
kraussei was 5.4 + 1.1 (average + S.D.). Steinernema kraussei occurred in samples
characterised by significantly lower pH values than samples with S. feltiae (U-test),
S. intermedia (U-test), and S. affinis (U-test, p = 0.005). Furthermore, S. kraussei
seems to have a specificity for the some types of vegetation (x? = 26.1, DF = 7, p<
0.001). It was exceedingly frequent in alpine larch stands, but was absent in mixed
forests and home gardens, and occurred less than expected in deciduous as well as in
spruce forests. In pastures and dwarf shrubland, S. kraussei occurred according to
sampling effort.
Only one isolate of S. intermedia and S. feltiae, each, were recovered from
forest soils, although woodland samples were relatively frequent (n = 153). Both S.
feltiae and S. intermedia seem thus to prefer grassland habitats. Similarly, S. affinis,
Heterorhabditis sp., and Steinernema sp. were confined to grassland habitats. How-
ever, due to the low prevalence of the species other than S. kraussei, their association
with the grassland habitat is only significant when they are treated as a group (x? =
12.8, DF = 1, p = 0.001). The pH values for S. feltiae (6.4 + 1.2; average + S.D.), S.
intermedia (6.2 + 0.7), S. affinis (6.9 + 0.4), Heterorhabditis sp. (6.5), and Steiner-
nema sp. (7.8) suggest that these species avoid extreme soil pH conditions.
DISCUSSION
TOTAL ENTOMOPATHOGENIC NEMATODES
Rhabditid entomopathogenic nematodes have been isolated from six continents
and appear to be among the most ubiquitous insect pathogens (POINAR 1990). While
most steinernematids were isolated from regions characterised by temperate or cool
climates, heterorhabditids were mainly recovered from soils in warmer regions
(POINAR 1990). In Europe, heterorhabditid and/or steinernematid nematodes occur in
all the countries surveyed so far, however, considerable variation exists in the
prevalence of entomopathogenic nematodes (BOAG et al. 1992). The European sur-
veys demonstrate that these nematodes become scarce as one moves northwards.
While they occur in central Europe (i.e. Germany, Switzerland, and Czechoslovakia)
with a recovery rate of over 25%, corresponding values are less than 20% in Northern
Europe (i.e. Norway, Republic of Ireland, Finland, Northern Ireland, and Scotland; in
decreasing order). Only one survey in Sweden with 25% positive samples (BURMAN et
al. 1986) and another one in Great Britain with 49% (HOMININCK & BRISCOE 1990a)
disclaim this tendency.
The observed differences in nematode prevalence may reflect variation in the
availability of suitable host species (MRACEK & WEBSTER 1993). However, since most
surveys concerning entomopathogenic nematodes are primarily performed to collect
new isolates for screening programs, ecological information necessary to explain the
distribution of the species is often lacking. Accordingly, sampling of the present
survey was designed to isolate cold-active nematodes, and no attempt was made to
448 WERNER A. STEINER
identify the natural hosts of the species detected. Another source for the variation in
recovery rates between surveys are methodological differences. Since the presence of
entomopathogenic nematodes in the soil is difficult to approve when their density is
low, the sensitivity of the detection method can substantially influence the estimation
of the nematode prevalence. In conclusion, only nematode-positive samples give
relevant information on the distribution of the species, whereas negative samples do
not provide solid evidence for the absence of the nematodes at a given sampling
location. Therefore, analysis of the species’ ecological niches using the information of
nematode-positive samples (tab. 2, fig. 4) should yield more reliable results than the
comparison of positive with negative samples, which served to analyse the environ-
mental preferences of the species.
SPECIES LEVEL
Steinernema feltiae is recorded all over Europe (e.g. MRACEK & JENSER 1988;
EHLERS et al. 1991; STURHAN pers. comm.). In Northern Europe, S. feltiae is usually
the most prevalent species (e.g. BOAG ef al. 1992; GRIFFIN ef al. 1991; VAINIO ef al.
1994). In Central Europe, S. feltiae represents a subdominant species. For instance in
Switzerland, it was the second most prevalent species (fig. 1). The observed
predominance of S. feltiae in the lower Alps and the Jura Mountains confirms the
view that it is a temperate species (HOMININCK & BRISCOE 1990). Complying with its
main prevalence in soils with relatively high pH values (G2, tab. 2), S. feltiae was
found to associate with calcareous soils in Great Britain (HOMININCK & BRISCOE
1990), and in pH neutral soils in Hungary (MRACEK & JENSER 1988). Contradictory
findings exist on the vegetational specificity of S. feltiae. HOMININCK & BRISCOE
(1990) recovered this species from diverse habitats, while in both Germany (STURHAN
pers. comm.) and Scotland (BoAG er al. 1992) S. feltiae was most common in
pastures. Results of the present study suggest that S. feltiae is associated with
grassland habitats (G2, fig. 4). The indicated preference of S. feltiae for grassland
soils is presumably caused by the distribution of its natural hosts, which include
lepidopteran larvae (Noctuidae and Hepialidae) feeding on the roots of tussock grass
(POINAR 1990). Fungus gnats and bibionid flies represent other natural hosts of S.
feltiae (POINAR 1992), which could explain its higher efficiency in parasitising sciarid
larvae as compared to the other steinernematid species (STEINER unpublished).
The overall prevalence of S. intermedia was similar to that of S. feltiae
(fig.1). In Germany, S. intermedia is the most commonly encountered species, fol-
lowed by S. affinis and S. feltiae (STURHAN pers. comm.). Steinernema intermedia
was further discovered in Norway (HAUKELAND 1993) and in South Carolina (POINAR
1990). The analysis of its distribution (fig. 4, tab. 2) suggests that S. intermedia is a
relatively unspecialised species, avoiding extreme pH conditions and exhibiting a
slight preference for grassland habitats. In Germany, however, S. intermedia was
significantly more often found in forests than in other habitats (STURHAN pers.
comm.). Therefore, its main distribution in grassland habitats of the alpine region
represents not a general vegetational specificity.
ENTOMOPATHOGENIC NEMATODES 449
Steinernema affinis, Steinernema sp. and Heterorhabditis sp. were too rare
(fig. 1) to allow a detailed interpretation of their distribution in the Swiss Alps.
Steinernema affinis was recovered in various regions (fig. 2) at altitudes between 500
and 2100 m, and was restricted to grassland samples with pH values near neutrality.
In Germany, S. affinis was more common than in Switzerland, and occurred most
often in pastures and arable soils (STURHAN pers. comm.). This confirms the
importance of the grassland habitat for the presence of S. affinis. Other records of this
species are reported from Norway (HAUKELAND 1993), Denmark (POINAR 1990), the
Republic of Ireland (GRIFFIN ef al. 1991), and Great Britain (REID & HOMININCK
1993). The natural hosts of S. affinis include bibionid fly larvae, which this species
therefore shares with S. feltiae (POINAR 1988).
Steinernema sp. is presumably a new species (close to S. intermedia), charac-
terised by the same restriction fragment length patterns (RFLP) as noted for a species
recently isolated in Great Britain (A. REID pers. comm.). Its geographical distribution
is therefore unknown. Heterorhabditis sp. (i.e. North-West European Group) was
detected only once in a pasture at 1130 m asl. Heterorhabditid species of the North-
West European Group occur also in Denmark, Germany, and Poland (Smits et al.
1991). A previously unpublished record of the same species in Switzerland at ca 400
m asl (KLINGLER pers. comm.) suggests that excessive sampling in the Swiss Plateau
or in the lower Alps would reveal further isolates of Heterorhabditis. The poor
representation of Heterorhabditis in the Swiss Alps, along with the high frequency of
steinernematids, supports the hypothesis that heterorhabditid nematodes are endemic
to warmer climates while steinernematids prevail in temperate climates (e.g. GRIFFIN
et al. 1991; MRACEK & WEBSTER 1993).
Steinernema kraussei was the most commonly encountered species in the
Swiss Alps (figs. 1 and 3). Its predominance was unexpected. In Germany, it occurred
equally frequent as S. feltiae, but was less common than S. intermedia and S. affinis,
while in Great Britain, only 20% of all samples were S. kraussei and around 50% S.
feltiae (A. REID pers. comm.). Since over 25% of the S. kraussei isolates in Switzer-
land originated from sampling locations above 2000 m, this species must endure long
periods of low temperatures, and is thus supposed to be a cold active species. In fact,
laboratory experiments showed that S. kraussei was significantly more active and
infective at low temperatures than S. feltiae or the heterorhabditid isolate (STEINER
1996).
The alpine distribution of S. kraussei indicates the ability of infective juve-
niles to survive for long periods at low temperatures (BRIAND & WELCH 1963 cited in
Wours 1991). The permanent snow-cover in winter stabilises soil temperatures near
the freezing point. The isolating effect of snow is especially important in regions with
extreme minima in air temperatures. In the Engadine (R6 in fig. 2; upper part of the
valley of the river Inn), for example, cold air sinking at night to the bottom of the
valley leads to very low air temperatures, but soil temperatures are similar to those
measured at other climatological stations (fig. 3B; 1800 m). While at 600 m the
period with a permanent snow-cover lasts for only 20 days, it increases constantly as
one goes upwards, reaching over 250 days above 2400 m. The upper altitudinal limit
450 WERNER A. STEINER
of S. kraussei could thus be governed by the sum of the positive temperatures in the
warmer season, which also affects the duration of the vegetational period and the
abundance of associated insect hosts. Evidence is drawn from laboratory obser-
vations, showing that 6-8 °C represent the lower limit for the successful propagation
of most alpine isolates (STEINER unpublished). This suggests that reproduction is
confined to the short summers. The precise role of temperature for the distribution of
entomopathogenic nematodes is thus direct via a lower limit for reproduction, and
indirect via the vegetation, the availability of associated host species, and soil for-
mation (see below).
The higher prevalence of S. kraussei at NW-SE facing slopes as compared to
sampling locations exposed to S-W is considered to be a further effect of temperature
via soil characteristics. According to the negative relationship between temperature
and soil formation, the relatively cool sampling locations oriented to NW-SE were
more acidic (U-test, p< 0.001) and had a higher content of organic matter (U-test, p <
0.05) than sites oriented to S-W.
Surprisingly, though, S. kraussei appears to have a lower limit of its natural
distribution in Switzerland. Why does S. kraussei not colonise areas downslope, all
the more as it occurs in Germany (STURHAN pers. comm.) and southern Bohemia
(MRACEK 1994) at lower altitudes? Unfortunately, the identity of this species was
unclear (e.g. POINAR 1990; MRACEK 1994), and notes on the actual distribution of S.
kraussei are scarce. Taking into account that laboratory cultures of the alpine S.
kraussei isolates can be maintained at 25 °C (STEINER unpublished), factors other than
high temperature (e.g. presence of natural hosts) must be responsible for the observed
lower limit of this species. Steinernema kraussei was originally isolated from the web
spruce sawfly (Cephalcia abietis [L.]) in a German forest stand (MRACEK 1994). The
high prevalence of S. kraussei in the Swiss Alps, however, must rely on another host
species, because C. abietis is rare in Switzerland. Occurring exactly in the same
altitudinal range as S. kraussei in the present survey, Zeiraphera diniana (Gn.), a
major pest of larch trees, could represent the natural host of this nematode. Since
sampling in the present survey was performed only one year after the population
collapse of Z. diniana in 1990, we can speculate that the larch bud-moth is partly
responsible for the high recovery rate of S. kraussei in larch stands of the Engadine
(R6 in fig. 2). The comparatively low frequency of S. kraussei in spruce forest is
presumably also related to the availability of suitable host species, all the more soil
characteristics of larch stands were similar to those measured in spruce forests.
The analysis of the ecological niche of S. kraussei showed that pH and sand
content of the soil, the humus, orientation, as well as the vegetation cover influence
the distribution of this species. Soil acidity, possibly associated with a high content of
organic matter, is considered a key factor for the distribution of S. kraussei. Evidence
is drawn from a general increase in soil acidity as one moves upwards, paralleled by a
habitat shift of the nematode. In the plains, S. kraussei seems to prefer woodland
soils (e.g. STURHAN pers. comm.), whereas in the Alps, its frequency was equal in
grassland and forest habitats. Forest soils differed from grassland at sample sites
below 1500 m by a lower pH (U-test, p < 0.001), whereas above 1500 m, their acidity
ENTOMOPATHOGENIC NEMATODES 451
was similar. Conversely, other differences between the two vegetation types remained
constant irrespective of altitude. Likewise, woodland samples were characterised by a
higher content of organic matter (U-test, p < 0.001), and a lower sand content (U-test,
p < 0.05) than grassland samples.
CONCLUSIONS
The present survey showed that rhabditid entomopathogenic nematodes are
represented in the Swiss Alps by six species. Steinernema kraussei is clearly the
dominant species at high elevations and must be well adapted to alpine climate. It
occurs especially frequent in alpine larch stands and occupies an ecological niche well
separated from the other nematodes encountered. Steinernema kraussei tolerates soils
with low pH and rich in organic matter, and is living in both grassland and forest
ecosystems, whereas S. feltiae, S. intermedia, and S. affinis are most prevalent in
grassland ecosystems at relatively high pH values.
ACKNOWLEDGEMENT
This research was funded by the Federal Office for Education and Science
under grant number COST812 ("Selection and evaluation of cold-active lines of
insect-parasitic nematodes for outdoor application"). The author thanks to Dr. J.
Klingler (CH-Wädenswil) for initiating this COST action in Switzerland. Special
thanks are extended to Dr. A. Reid (GB-St Albans) and to Dr. P. Smits (NL-Wagen-
ingen) for identifying selected nematode isolates, to Dr. W. Heller and H. Schwager
(CH-Wädenswil) for analysing soil characteristics, as well as to Dr. W. Stahel for
preliminary discussions about statistical analyses. Finally, the author is grateful to Dr.
K. Tschudi-Rein for critically reviewing the manuscript.
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REVUE SUISSE DE ZOOLOGIE, 103 (2): 453-473; juin 1996
Revision of the soil-weevil genus Guineobius Osella
from New Guinea
(Coleoptera, Curculionoidea, Entiminae, Celeuthetini)*
Alexander RIEDEL & Klaus SCHÖNITZER
Zoologische Staatssammlung, Münchhausenstraße 21, D-81247 München, Germany.
Revision of the soil-weevil genus Guineobius Osella from New Guinea
(Coleoptera, Curculionoidea, Entiminae, Celeuthetini). - Four new
species of the entimine soil-weevil genus Guineobius Osella (G. niger
Riedel, G. minutus Riedel, G. depressus Riedel, G. baliemensis Riedel) are
described from the highlands of Irian Jaya. This is the first record of this
genus for the western part of New Guinea. The two formerly known
species of this genus from Papua New Guinea are redescribed. A key is
given to all species and the phylogenetic significance of characters is
discussed. The relative length of abdominal sternites proved to be highly
variable within the genus. The possession of modified scales in most
species is suggested as an apomorphic character for Guineobius Osella.
Notes are given on ecology and conservation.
Key-words: Coleoptera - Curculionoidea - New Guinea - Taxonomy.
INTRODUCTION
Knowledge of the soil-weevil fauna of New Guinea perfectly reflects the
recent exploration of this island in other scientific and cultural fields. No endogean
weevil was known from New Guinea when OSELLA (1983) described the genus
Guineobius based upon two species, one of them known from a unique specimen, the
other known from two specimens only. Since then there are no further records.
However, this is not surprising, since most of the highland areas became accessible
only 20-30 years ago and the few collecters who have visited this region since,
resorted mainly to standard collecting techniques such as beating and sweeping.
Sieving the ground litter, the usual way to obtain endogean weevils was usually
neglected.
On a series of his own expeditions to New Guinea beginning in 1990 the
senior author repeatedly visited one locality on the eastern slopes of the Baliem valley
* Results of the entomological explorations of A. Riedel in New Guinea in 1990-1993.
Manuscript accepted 02.04.1995.
454 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
(rian Jaya) which resulted in the discovery of three additional species there. One
more species was found in the Nalca region. These four new species are described
below and their phylogeny and ecology are discussed.
MEASUREMENTS
Since the eyes in Guineobius are reduced or completely absent, the length of
the rostrum was not measured from the anterior margin of the eyes as is usual in most
Curculionoidea, but from the transverse furrow at the base of the rostrum. The breadth
of the rostrum was measured between the pterygia. The width of the elytra was
measured across both elytra at their widest point.
CHARACTERS
When OSELLA (1983) described the genus he defined it mainly on the loss of
eyes, the ferruginous coloration, the strong punctation of the pronotum and the
morphology of the abdomen (more or less straight suture between sternites III and IV;
and sternites V and VI subequal in length). Since characters associated with the soil-
dwelling habit, namely loss of eyes and ferruginous coloration are prone to conver-
gence, we were looking for better evidence to prove the monophyly of the genus
Guineobius. This was especially important since we discovered striking differences of
the relative lengths of abdominal sternites, which made at first a polyphyly appear
possible.
Modified scales of asymmetrical radiate pectinate shape are interpreted as
apomorphy for Guineobius (Figs. 1-3). They are often found in dense patches or
rows, especially on the ventral surface and are usually encrusted with dirt. The
function of these scales is unknown. They are similar to some scales described by
PAULAY (1985) in the genus Miocalles of Cryptorhynchinae, but we have not found
such scales in any other genus of Celeuthetini. There are two species of Guineobius
(G. minutus sp. n., G. viduus Osella) which lack these scales but this can be explained
by secondary loss (see Phylogeny).
The variability of the abdomen (Figs 16—21) is astonishing. The two extreme
types would probably lead a taxonomist to describe two different genera if the
intermediate forms were unknown. It must be noted that in Guineobius the length
ratios between abdominal sternites are identical in both sexes. In other genera of
Celeuthetini such as Stereogastrus and Piezonotus some abdominal sternites are en-
larged in females only.
An interesting genital character of various genera of Celeuthetini which is also
present in at least two species of Guineobius is a partial eversion of the median part of
the endophallus (Fig. 22). This is not an incomplete re-inversion but obviously the
normal state since the walls of this part of the endophallus part are partially
sclerotized. In Guineobius this character is most conspicuous in G. niger, but it is
present in G. minutus as well. We have found it also in another probably undescribed
genus of Celeuthetini. Sharp (1918) mentions in a "Trigonops, or Heteroglymma sp.
n.?" from New Guinea a "structure which has all the appearance of being a tube"
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 455
inside the median lobe and we believe it is the same character described above. Since
the species mentioned above do not seem to be closely related in other respects the
character of the secondary tube seems to be an ancient one in the evolution of
Celeuthetini.
The length of the flagellum seems to be highly variable within the tribe
Celeuthetini. Within Guineobius two forms occur: one is short and simply curved, the
other long, thread-like and spiral. The long form is probably a character reversal in
Guineobius (see also Phylogeny). According to THOMPSON (1977) both short and long
flagella also occur in the genus Apirocalus.
Guineobius Osella
Type-species: Guineobius viduus Osella, 1983, by original designation.
REDESCRIPTION:
Length. Small, pronotum plus elytron: 1.59-2.78 mm.
Rostrum with apical declivity evenly, not abruptly sloping, subglabrous, with
several long setae; dorsally separated from head by more or less distinct transverse
furrow which curves laterally apicad towards antennal scrobes.
Head with apical portion merging with rostrum; eyes lacking or small (not
larger than last funicular article), smooth, without visible facets; if present eyes placed
on anterior portion of head in front of level of median part of transverse furrow but
ventral to forward curving lateral parts of transverse furrow (Figs 10, 13-15).
Antennae with scape passing anterior margin of pronotum in repose, setose,
and more or less granulate; funicle with article 1 about 2X as long as broad apically, 2
shorter than 1, 3-7 subequal in length, ca. as long as broad.
Elytra smooth; strial punctures deeply impressed; humeri and scutellum
lacking.
Metasternum short, shorter than diameter of mesocoxa.
Abdomen with intercoxal process broad, 1.5-1.8X wider than metacoxae.
Legs with ventral surface of tarsı with long curved suberect setae, claws stout.
Vestiture generally lacking except most punctures (especially on elytra)
bearing single seta and except modified scales, often condensed in patches on head or
on thoracic and abdominal venter.
KEY TO THE SPECIES OF Guineobius
1 Body large (pronotum plus elytron 2.4-2.8 mm), black; sternites V-VI
shon:fequalinien st Gi) PC ER RO AEREE niger Sp. n.
- Body small (pronotum plus elytron 1.6-2.3 mm), ferruginous; sternites
V—VI longer, subequal or unequal in length (Figs 17-21). ............... 2
2 Sternites V and VI subequal in length with sternite VI slightly longer;
suture between sternites III-IV entirely absent (Figs 17-18)... ............ 3)
456 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
- Sternites V and VI slightly to strongly unequal in length with fifth
sternite 2X—3X longer than sternite VI; suture between sternites III-IV
distmettatileastilaterally.(Eies 192 ee eee 4
3 Rostrum above with transverse basal band of scales, head smooth (Fig.
here tati adoro Berner deharvengei Osella
— Rostrum lacking transverse basal band of scales, head punctate
(EISEN) nici et oo OSE viduus Osella
4 Fifth sternite 2.0X longer than sternite VI (Fig. 19); body size small,
pronotumiplusfelytrontdi:6/imm* 2 e minutus Sp. n.
— Fifth sternite 2.8X—3.1X longer than sternite VI (Figs 20-21); body size
larger, pronotum plus elytron 1.9-2.3 mm... 2... 2 2.2 2.22 ReSSSRA 5
5 Base of rostrum ventrally with round patch of scales; body elongate,
nlattenedi(Eie: 8) head smooth (Fig 14) 2 2 eee depressus sp. n.
- Base of rostrum ventrally with glabrous pit; body not strikingly elon-
gate or flattened (Fig. 9): head punctate (Fig. DS). 2... baliemensis sp. n.
Guineobius niger Riedel, sp. n. (Figs 1-4, 10, 16, 23, 28, 33)
MATERIAL EXAMINED: Holotype: Irian Jaya, Jayawijaya-Province, Wamena, Jiwika;
1900-2000 m, 23.1X.1992, leg. A. Riedel. Paratypes: Irian Jaya, Jayawijaya-Province,
Wamena, Jiwika, ca. 1700-2300 m, 2.1X.1991, leg. A. Riedel (4); 1700-2000 m, 11.IX.1991,
leg. A. Riedel (2 and allotype); 1900-2050 m, 24.X.1993, leg. A. Riedel (15); as holotype (28,
l coated for SEM).
Holotype and allotype deposited in the Zoologische Staatssammlung (München).
Paratypes deposited in collections A. Riedel (München), G. Osella (L'Aquila), C.W. O'Brien
(Tallahassee) and Muséum d'histoire naturelle (Geneva).
DIAGNOSIS: Large, black; base of rostrum ventrally with round patch of scales,
center of metasternum densely squamose; sternites V—VI short, equal in length.
DESCRIPTION:
Holotype, male. Length. Pronotum plus elytron: 2.78 mm. Rostrum 1.36X
longer than broad, in dorsal view lateral margins roundly diverging to antennal
insertion, there rostrum much broader than at base; furrow separating dorsal part of
rostrum partly merging with antennal scrobes anteriorly, laterally strongly impressed,
posteriorly shallow and medially weakly impressed; declivital area in front of
antennal scrobes moderately long; parts of scrobes in front of antennal insertion short,
at antennal insertion curved inwards, converging obliquely in straight line, posteriorly
open; dorsum between scrobes narrowing posteriad, almost flat, but slightly elevated
medially in anterior portion, with long setae arranged in form of anteriorly open V;
dorsum in posterior 1/2 of rostrum narrow, narrower than length of article 2 of
funicle; behind with shallow, indistinct median furrow and punctate; sides of dorsum
steeply concavely sloping; in lateral view straight in basal 1/2, dorsal and ventral
margins parallel; at level of antennal insertion broader, curved ventrad; ventrally with
large round patch of modified scales; with setiferous punctures.
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 457.
Guineobius niger sp. n., modified scales (SEM). (1) ventral aspect of mesothorax, metathorax
and sternite III with squamose patches; (2) ventral aspect of head with squamose patch at base
of rostrum; (3) modified scales with dirt incrustations. Scale bars: (1-2) 0.2 mm; (3) 0.05 mm.
Head with basal part globose, smooth; vestigial eyes as large as last funicular
article, located dorsolaterally on ovate elevated punctate lobes reaching from rostrum
onto smooth part of head. Antennae. Funicle with article 1 conical; 2 elongate, 1.7X
longer than 3; club with basal half conical, apical half broadly rounded, apical 1/3
more densely setose.
Prothorax slightly (1.09X) longer than broad, side margins strongly rounded;
disc strongly punctate with large punctures, distance apart greater than their diameter;
sides with widely scattered, smaller punctures; basally constricted, constriction
densely squamose, medially strongly rounded, apically gradually narrowing dorsally.
Elytra elongate-ovate, 1.37X longer than broad, broadest at basal 1/4, then subparallel
458 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
to apical 1/4; deeply punctate, setae minute, punctures separated by length of their
diameter; with 10 regular striae; stria 8 shortened at base, beginning at level of
metacoxa. Prosternum. Coxae separated by 0.20X coxal diameter; densely squamose
behind coxae; with anterior apical constriction; with deep posterior impression
between coxae. Mesosternum with intercoxal process densely squamose, coxae
separated by 0.92X coxal diameter. Metasternum medially densely squamose,
laterally subglabrous with row of squamiferous punctures bordering mesocoxae.
Abdomen. Sternites II and IV subequal in length, suture between fine but
distinct, slightly sinuate medially; sternite III with deep, sinuate impression bordering
front-margin of intercoxal process, with row of squamiferous punctures continuing to
elytral margin; remainder of sternites III and IV smooth, shallowly punctate, medially
concave; sternites V and VI equal, narrow, together 1/2 as long as sternite IV; sternite
VII half-rotund, longer than sternites V and VI together, punctate. Legs. Tarsi with
tarsomere 3 long, ventrally deeply bilobed, tarsal articulation at posterior 1/2.
Coloration of cuticle black, with slightly bluish, oily shine.
Genitalia. Median lobe much shorter (0.47X) than apodemes, stout, in lateral
view weakly curved from base to apex, in dorsal view apex evenly rounded. Endo-
phallus with distinct partial median eversion, here membrane sclerotized in part;
apical half curved forward to base of median lobe, with minute spines; flagellum
0.54X as long as median lobe.
Allotype, female. Same as male except: Length. Pronotum plus elytron 2.78
mm. Prothorax with side margins slightly less strongly rounded. Abdomen with ster-
nites III and IV convex.
Genitalia. Spermatheca with distinct ramus; extended at insertion of sperma-
thecal duct; insertion of spermathecal duct and of spermathecal gland well-separated;
body apically bent inward and much thinner than at base.
Infraspecific variation: Length. Pronotum plus elytron 2.41—2.78 mm. Colo-
ration from yellowish brown in one teneral specimen to piceous and completely black,
with bluish shine in most specimens. Prothorax. Punctation varying from denser
(diameter of punctures larger than interspaces between punctures) and deeper to spar-
ser (diameter of punctures smaller than interspaces between punctures) and shallower.
Etymological Note: This epithet is based on the Latin adjective niger (black,
dark) and refers to the black coloration in mature specimens.
Guineobius deharvengei Osella 1983 (Figs:5; 141702999)
Guineobius deharvengei Osella, 1983: 115.
MATERIAL EXAMINED: Holotype, female, Papua New Guinea, Port Moresby, Rauna
Falls, 5.XI.1978, leg. Deharveng. Deposited in Muséum d'histoire naturelle (Geneva).
DrAGNosis: With transverse band of scales dorsally at base of rostrum; dorsum
of rostrum between antennal scrobes with sides converging posteriad in straight line;
head impunctate; sternite VI longer than sternite V, suture between sternites III and
IV entirely effaced.
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 459
Fics 4-9
Habitus, dorsal: (4) Guineobius niger sp. n., paratype, length of pronotum plus elytron 2.65
mm; (5) G. deharvengei Osella, holotype, length of pronotum plus elytron 1.82 mm; (6) G.
viduus Osella, holotype, length of pronotum plus elytron 1.80 mm; (7) G. minutus sp. n.,
holotype, length of pronotum plus elytron 1.59 mm; (8) G. depressus sp. n., holotype, length of
pronotum plus elytron 2.34 mm; (9) G. baliemensis sp. n., allotype, length of pronotum plus
elytron 1.94 mm.
REDESCRIPTION:
Holotype, male. Length. Pronotum plus elytron: 1.82 mm.
Rostrum as long as broad, lateral margins subparallel in dorsal view: trans-
verse furrow separating dorsal part of rostrum medially broadly deeply impressed,
laterally finer, ending below antennal scrobes, throughout partly hidden by ring of
460 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
modified scales; declivital area in front of antennal scrobe moderately short; antennal
scrobes S-shaped, in front of antennal insertion narrow, curved inward, posteriorly
broad, converging slightly sinuate in almost straight line, posteriorly open; dorsum
between scrobes narrowing with straight margins posteriad, behind separated by
1.67X length of funicular article 2, posteriad scrobes widening, but very short;
dorsum smooth except median furrow in posterior 1/2 and few punctures and setae
anteriorly; in lateral view straight, tapering apicad, ventral margin straight, dorsal
margin sloping down roundly; base dorsally with transverse dense band of modified
scales, keeping posteriad antennal scrobes, laterally tapering; in the holotype band
asymmetrical, reaching further ventrad on right side; ventrally subglabrous, with few
long setae; postmentum triangularly elevated.
Head globose, smooth, shining. Antennae. Funicle with article 2 short, 1.2X
longer article 3; club pointed apicad, evenly setose.
Prothorax almost (1.07X longer) as long as broad, side margins strongly roun-
ded; disc deeply punctate, diameter of punctures larger than their distance, punctures
with suberect setae; laterally with same sculpture; with squamose basal constriction.
Elytra elongate, 1.59X longer than broad, broadest about middle, evenly curved from
base to apex, setae moderately long; stria 8 shortened at base, beginning at level of
metacoxa. Prosternum glabrous, punctate with distinct preapical constriction; coxae
separated by 0.56X coxal diameter. Mesosternum with coxae separated by 1.67X
coxal diameter. Metasternum especially on sides deeply punctate, subglabrous, with
scattered setae.
Abdomen glabrous, sternites III-IV densely shallowly punctate, medially
shallowly depressed, suture between sternite III and IV completely effaced; sternite V
narrow, sternite VI broad, 1.42X longer sternite V; sternite VII 1.27X longer sternites
V and VI together, apically broadly rounded. Legs. Tarsi with tarsomere 3 short,
broad, deeply bilobed. Coloration of cuticle light ferruginous, head and prothorax
little darker.
Genitalia: Spermatheca crescent-shaped, extended at insertion of spermathecal
duct; insertion of spermathecal duct not clearly visible in allotype, but probably well
separated from insertion of duct; body tapering apicad, outer margin evenly curved.
Remarks: The most distinct character of this species, a transverse dorsal band
of modified scales at the base of the rostrum is not a complete ring as OSELLA des-
cribed it. Ventrally there were only dirt-accrustations faking easily scale-covering, but
they could easily be removed with warm water.
Etymological Note: This species was named in honour of Mr. Deharveng. If it
was formed from this modern name the spelling would have to be deharvengi. It is
spelled this way on the label of the holotype. However, in the original publication
OSELLA has used the name deharvengei. This has to be taken as the genitive of the
latinized name deharvengeus. Usually the latinization of a male personal name is
formed by adding the ending -ius, but the ending -eus is valid as well. Thus the name
deharvengei is a correct original spelling although a little unusual.
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 461
Fics 10-15
Head, dorsal (SEM). (10) G. niger sp. n., paratype; (11) G. deharvengei Osella, holotype; (12)
G. viduus Osella, holotype; (13) G. minutus sp. n., holotype; (14) G. depressus sp. n., holotype;
(15) G. baliemensis sp. n., allotype. Scale bars: 0.2 mm.
Guineobius viduus Osella 1983 (Eigs’6, 1255185295 33)
Guineobius viduus Osella, 1983: 114.
MATERIAL EXAMINED: Holotype, male, Papua New Guinea, Chiumbu, Chuave,
22.X1.1978, leg. Deharveng. Deposited in Muséum d'histoire naturelle (Geneva).
DIAGNosIs: Devoid of scales; head strongly punctate; sternite VI longer than
sternite V, suture between sternites III and IV entirely effaced.
462 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
REDESCRIPTION:
Length. Pronotum plus elytron: 1.80 mm. Rostrum 1.17X longer than broad, in
dorsal view lateral margins slightly evenly roundly diverging apicad; furrow
separating dorsal part of rostrum deeply impressed throughout, anteriorly ending
below antennal scrobes; declivital area in front of antennal scrobe moderately short;
antennal scrobes intermediate between S- and kidney-shaped form, posteriorly
opposing each other at blunt angle, separated by 1.25X length of funicular article 2;
dorsum anteriorly between antennal scrobes widening posteriad, then sinuately
narrowing posteriad, posteriorly antennal scrobes parallel-sided, moderately broad,
curved; dorsum in apical part medially elevated, elevation surrounded by long setae
arranged in form of anteriorly open V; in lateral view straight, ventral margin straight,
dorsal margin convex; laterally with setiferous punctures, sculpture confluent with
head; ventrally at base with scattered non-modified suberect scales.
Head flat, above densely deeply punctate; ventrally smooth, glabrous. An-
tennae. Funicle with article 2 short, 1.3X longer article 3, stalked, basal 1/2 narrow
and parallel-sided; club ovate, apically weakly pointed and denser setose.
Prothorax 1.15X longer than broad, side margins rounded; disc deeply
punctate, laterally diameter of punctures larger than their distance, medially their
distance larger and partly impunctate; on sides punctation denser; base without cons-
triction. Elytra elongate, 1.63X longer than broad, broadest about middle, evenly
curved from base to shortly before apex, then slightly sinuate; setae moderately long.
Prosternum glabrous, punctate, preapical constriction indistinct; coxae separated by
0.50X coxal diameter; striae 6 and 7 shortened at base, beginning at level of
metacoxa. Mesosternum with coxae separated by 1.21X coxal diameter. Metasternum
subglabrous, with few setae.
Abdomen glabrous, sternites III and IV laterally punctate, medially deeply
impressed and smooth; suture between sternites III and IV medially completely
effaced, laterally with trace of suture; sternites V and VI broad, subequal, sternite VI
1.1X longer sternite V; sternite VII 1.1X longer sternites V and VI together, apically
rounded. Legs. Tarsi with tarsomere 3 short, broad, deeply bilobed. Coloration of
cuticle light ferruginous, head and prothorax little darker.
Genitalia. Median lobe slightly shorter (0.91X) than apodemes, in lateral view
strongly curved in basal 1/4, in dorsal view apex produced into pointed apical lip.
Endo-phallus without visible partial median eversion, inside median lobe with minute
spines, without distinct sclerites; outside median lobe first bent apicad, then bent
basad resulting in Z-shaped form; flagellum 0.35X as long as median lobe.
Guineobius minutus Riedel, sp. n. (Figs 7, 13, 19, 30, 33)
MATERIAL EXAMINED: Holotype: Irian Jaya, Jayawijaya-Province, Diuremna near Nalca,
1900-2100 m, 9-11.IX.1992, leg. A. Riedel. Left hind leg missing.
Holotype deposited in the Zoologische Staatssammlung (Miinchen).
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 463
Fics 16-27
(16-21) ventral aspect of thorax and abdomen; (16) G. niger sp. n.; (17) G. deharvengei Osella;
(18) G. viduus Osella; (19) G. minutus sp. n.; (20) G. depressus sp. n.; (21) G. baliemensis sp. n.;
(22) G. niger sp. n., aedeagus, schematic longitudinal section; (23-26) spermatheca; (23) G. niger
sp. n., allotype; (24) G. deharvengei Osella; (25) G. depressus, holotype; (26) G. baliemensis n.
sp., allotype; (27) G. depressus sp. n., holotype, sternite VIII. Scale bars: (16-21) 0.5 mm;
(23-26) 0.1 mm; (27) 0.2 mm.
464 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
Fics 28-31
Male genitalia; aedeagus in dorsal (A) and lateral (B) aspect. (28) G. niger sp. n., holotype; (29)
G. viduus Osella, holotype; (30) G. minutus sp. n., holotype; (31) G. baliemensis sp. n.,
holotype; (31c) apical aspect. Scale bars: (28-31B) 0.2 mm; (31C) 0.1 mm.
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 465
DIAGNOSIS: Small; without modified scales; posterior half of rostral dorsum
narrow, ridge-shaped; deep punctures of elytra each with long suberect seta; sternite
VI 1/2 as long as sternite V.
DESCRIPTION:
Holotype, male. Length. Pronotum plus elytron: 1.59 mm. Rostrum 1.49X
longer than broad, in dorsal view lateral margins subparallel from base to apex;
furrow separating dorsal part of rostrum anteriorly ending where antennal scrobes
bent inwards, shallow but distinct throughout; dorsum of rostrum extending tongue-
like far behind on head; declivital area in front of antennal scrobes long; antennal
scrobes kidney-shaped in dorsal view, posterior 1/2 strongly bent inward, scrobes
opposing each other at blunt angle, separated by dorsum equal to length of funicular
article 2; dorsum between antennal scrobes subquadrate, flat, with relatively short
setae, irregularly arranged in form of anteriorly open V; dorsum in posterior half of
rostrum extremely narrow, ridge-shaped, but with distinct median furrow; sides of
dorsum smoothly sloping, glabrous, shining; round lobes stretching backwards onto
smooth head including together with posteriorly produced dorsum cuneiform process
of head bearing setae directed medially and tiny vestiges of eye in the anterior corner;
in lateral view parallel basally, tapering apicad; ventrally obliquely wrinkled, sub-
glabrous, with few erect setae; medially with small pit, but without modified scales.
Head smooth and shining. Antennae. Funicle with second article 1.6X longer
article 3; club ovate, apex weakly pointed, apex denser setose than base. Prothorax
elongate, side margins rounded, 1.13X longer than broad; except along mid-line
densely punctate with large punctures, their diameter larger than their distance, each
puncture bearing long suberect seta; with narrow glabrous basal constriction. Elytra
elongate, at base weakly rounded, broadest behind middle, in apical 1/5 strongly and
broadly rounded to apex; strial punctures large, deep, bearing long erect seta subequal
in length to diameter of puncture; stria 6 shortened at base, beginning at level of
metacoxa; stria 7 complete; stria 8 shortened, beginning behind level of metacoxa;
stria 10 pressed to elytral margin.
Prosternum with preapical constriction, glabrous, punctate, coxae separated by
0.29X coxal diameter. Mesosternum glabrous, punctate with deep large punctures,
coxae separated 0.73X coxal diameter. Metasternum glabrous, punctate.
Abdomen glabrous, punctate; front- and hind-margin of sternite III bordered
by a row of punctures; sternites III and IV subequal in length, suture between distinct
throughout, medially sinuate; sternites V and VI together shorter (0.76) than sternite
IV; sternite VI half as long as sternite V; sternite VII slightly longer (1.1X) than
sternite V and VI together, broadly rounded apically. Legs. Tarsi with tarsomere 3
short, subquadrate, apex ventrally sinuate, cavity for articulation of tarsomere 5
occupying greatest part. Coloration of cuticle ferruginous.
Genitalia. Median lobe much shorter (0.54X) than apodemes, dorsoventrally
flattened, in lateral view weakly curved from base to apex, in dorsal view apex
bluntly pointed. Endophallus with short median partial eversion, without distinct
466 ALEXANDER RIEDEL & KLAUS SCHÖNITZER
sclerites, without minute spines; apex curved apicad towards base of median lobe,
without distinct sclerites; flagellum 0.62X shorter median lobe.
Etymological Note: This epithet is based on the Latin adjective minutus (small,
tiny) and refers to the small size of this species.
Guineobius depressus Riedel sp. n. (Figs 8, 14, 20, 25, 27, 33)
MATERIAL EXAMINED: Holotype: Irian Jaya, Jayawijaya-Province, Wamena, Jiwika;
1900-2000 m, 23.1X.1992, leg. A. Riedel
Holotype deposited in the Zoologische Staatssammlung (München).
DIAGNOSIS: Body flattened; rostrum ventrally with round patch of modified
scales; sternite V much longer (3.07X) than sternite VI;
DESCRIPTION:
Holotype, female. Length. Pronotum plus elytron: 2.34 mm. Rostrum 1.43X
longer than broad; in dorsal view lateral margins subparallel from base to apex;
furrow separating dorsal part of rostrum anteriorly ending below antennal scrobes,
laterally well impressed, posteriorly less distinct; declivital area in front of antennal
scrobes short; antennal scrobes kidney-shaped in dorsal view, opposing each other in
almost straight line, scrobes there separated by dorsum broader (1.2X) length of
funicular article 2, posterolateral margin not sharply delimited; dorsum between
scrobes subquadrate, flat, with long irregularly arranged setae; dorsum in posterior
half of rostrum broad, broader than funicular article 2 long, with flat, broad median
sulcus, sides of dorsum steeply sloping; in lateral view rostrum curved, dorsal and
ventral margins parallel; laterally shining with setiferous punctures; behind vestige of
eye wrinkly sculptured; ventrally with round patch of modified scales.
Head smooth. Antennae. Funicle stout, second article 1.7X longer article 3,
articles 3—7 parallel-sided; club ovate, apex weakly pointed, evenly densely setose
except extreme base subglabrous.
Prothorax slightly longer (1.11X) than broad, side margins moderately rounded,
disc convex, strongly punctate with large punctures, their distance smaller than their
diameter; sides denser punctate than disc; basal part constricted, separating cylindrical,
ventrally and laterally densely squamose basal part narrowing dorsally, there medially
glabrous; Elytra elongate, 1.46X longer than broad, at shoulders evenly rounded, then
subparallel till rounded at apex; dorsoventrally depressed; strial punctures small, later-
ally punctures bearing short erect seta, medially glabrous; striae 7-8 shortened at base,
beginning at level of metacoxa. Prosternum. Coxae separated 0.63X coxal diameter;
with preapical constriction; behind constriction to hind margin densely covered with
modified scales. Mesosternum. Coxae separated by 1.1X coxal diameter; margin of
intercoxal process with modified scales. Metasternum. Center glabrous, shining, punc-
tate; sides and margin of mesocoxae punctate, punctures with sparse modified scales.
Abdomen. Sternites II and IV subequal in length, suture between distinct
throughout and strongly sinuate; sternite III with a row of deep, squamiferous punctures
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 467
bordering front margin of intercoxal process: rest of abdomen glabrous, shining,
sparsely shallowly punctate; sternite V and VI together almost (0.88X) as long as
sternite IV; fifth sternite 3.07X longer than sternite VI; sternite VII shorter (0.83) than
sternites V and VI together, broadly rounded apically. Legs. Tarsı with tarsomere 3
moderately long, narrow, apex deeply bilobed. Coloration of cuticle ferruginous.
Genitalia. Spermatheca robust, with weak ramus; extended at insertion of
spermathecal duct; insertion of spermathecal duct and spermathecal gland well-
separated; body weakly tapering apicad, apical 1/2 in right angle bent inward.
Etymological Note: This epithet is based on the participle of the Latin verb
deprimere (flatten) and refers to the dorsoventrally flattened body-shape.
Remarks: Unfortunately this species is known only from a unique female.
However it can be clearly separated from the other known species and the male
genitalia are not of such crucial importance in this genus as in some other groups of
Curculionoidea. Therefore we consider it beneficial to provide a name for this species
here as well.
Guineobius baliemensis Riedel, sp. n. (Ios 957219631733)
MATERIAL EXAMINED: Holotype: Irian Jaya, Jayawijaya-Province, Wamena, Jiwika;
1900-2050 m, Winkler-ecclector, 24.X.1993, leg. A. Riedel. Paratypes: as holotype (2); Irian
Jaya, Jayawijaya-Province, Wamena, Jiwika, 1900-2050 m, 24.X.1993, leg. A. Riedel
(allotype).
Holotype and allotype in collection A. Riedel (München), deposited on indeterminate
loan in the Zoologische Staatssammlung (München). Paratypes deposited in collection A.
Riedel (München).
DIAGNOSIS: Rostrum ventrally at base with glabrous pit; head punctate;
prosternum with coxae widely (0.6X their diameter) separate, deeply impressed, with
modified scales; sternite V much longer (2.8X) than sternite VI.
DESCRIPTION:
Holotype, male. Length. Pronotum plus elytron: 1.60 mm. Rostrum 1.30X
longer than broad, in dorsal view lateral margins subparallel from base to apex;
furrow separating dorsal part of rostrum anteriorly ending below antennal scrobes, its
anterior 1/2 broadly and well impressed, behind shallow and almost effaced; declivital
area in front of antennal scrobes long; scrobes evenly kidney-shaped in dorsal view,
well delimited, posteriorly separated by dorsum broader (1.4X) length of funicular
article 2; dorsum between antennal scrobes subquadrate, subglabrous with few setae;
dorsum in posterior half of rostrum broad, broader than funicular article 2 long; sides
of dorsum smoothly convexly sloping; in lateral view weakly tapering apicad, ventral
margin straight, dorsal margin weakly curved; laterally mat, sparsely shallowly
punctate, setose; with vestige of eye; lateral lobes weakly developed; ventrally behind
with deep median elongate glabrous pit, laterally with sparse modified scales.
Head punctate. Antennae. Funicle with article 2 elongate, subequal in length to
article 1, 1.8X longer article 3; articles 3-7 subequal, about as long as broad; club
ovate, apical 1/2 denser setose than base.
468 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
Prothorax slightly longer (1.13X) than broad, side margins weakly rounded,
densely punctate but medially distance between punctures longer than diameter; basal
constriction narrow, glabrous. Elytra ovate, 1.40X longer than broad; strial punctures
large, deep, on declivity shallow; with 10 regular striae; striae 7 and 8 shortened,
beginning at level of metacoxa; laterally punctures each with short erect seta, me-
dially glabrous. Prosternum. Coxae widely separated by 0.6X coxal diameter; area
between coxae cavernous, with scattered modified scales; without preapical cons-
triction. Mesosternum with intercoxal process sparsely squamose, coxae separated by
1.15X diameter, suture with metasternum distinct. Metasternum glabrous, punctate.
Abdomen glabrous, center sparsely, shallowly punctate; anterior margin of sternite III
with row of deeper punctures; sternites III and IV medially with broad impression;
sternites III-IV subequal in length, suture between medially obsolete, laterally weak;
sternites V—VI together equal in length to sternite IV; sternite V little shorter (0.75)
sternite IV, 2.8X longer than sternite VI; sternite VII shorter (0.85X) than sternite V
and VI together, broadly rounded apically. Legs. Tarsi with tarsomere 3 moderately
long, apex deeply bilobed. Coloration of cuticle ferruginous.
Genitalia. Median lobe much shorter (0.53X) than apodemes, in lateral view
strongly curved, moderately flattened dorsoventrally, in dorsal view apex with
Straight sides angulately pointed, behind orifical area internally with sclerites. Endo-
phallus hyaline, without visible partial eversion, without minute spines, straight;
flagellum long, thread-like, spiralized.
Allotype, female: same as holotype except: Length. Pronotum plus elytron:
1.94 mm. Rostrum 1.36X longer than broad; dorsum between antennal scrobes with
few short, maybe broken setae, medially with flat broad ridge; dorsum in posterior
half with median furrow. Prothorax densely punctate throughout, sometimes distance
between punctures smaller than diameter. Elytra with stria 8 shortened. Mesosternum
with coxae separated by 1.6X diameter.
Abdomen. Sternites III-V convex. Spermatheca without ramus, insertion of
spermathecal gland approximate to insertion of spermathecal duct; body hardly ta-
pering apicad, thinner at base than at apex; outer margin evenly curved except slightly
concave near base.
Infraspecific variation. Prothorax. Area between procoxae more or less deeply
impressed to cavernous.
Etymological Note: This epithet is based on the name of the Baliem-river-
valley where the type locality is located.
SYSTEMATIC POSITION OF Guineobius
The only classification of Celeuthetini available is the revision by MARSHALL
(1956). Although being of great value for taxonomic purposes it does not provide an
appropriate basis for phylogenetic considerations. MARSHALL did not give a proper
diagnosis of the tribe except differentiating it against Elytrurini. From his comments
and from our observations the Celeuthetini sensu MARSHALL are likely a mono-
phyletic group; the character of the transverse sulcus dorsally separating head from
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 469
rostrum would be the only autapomorphy available at present. However, it must be
noted that some African Peritelini have a similar structure, so additional studies are
required to confirm the significance of this character. Most genera of Celeuthetini
have their rostrum abruptly declivous at the apex, but some genera including Guineo-
bius have it eveniy sloping apicad. We consider the first character state synapo-
morphic, the second one symplesiomorphic. Within the paraphyletic assemblage of
genera having evenly sloping rostrum, the genera Apirocalus, Hellerrhinus and Koko-
danus possess conspicuous spines on their elytral declivity and are thus considered to
constitute another monophyletic group. Looking for a sister group of Guineobius we
would have to consider following other groups in the analysis: All species with abrupt
rostral declivity, the group of Apirocalus — Hellerrhinus — Kokodanus and each of the
remaining (monotypic) genera with evenly sloping rostrum, namely Acoptorrhynchus,
Oedirrhynchus, Idorrhynchus and Pachyrrhynchidius. Unfortunately the four latter
genera are known to us from the description only. No characters were found to hint on
the position of Guineobius among the above mentioned groups.
Another possible sister group of Guineobius was found in leaf litter in New
Guinea. Two species were collected that should be placed in a new genus close to
Guineobius, but for a formal description the material at hand is not sufficient.
The species have an apicad evenly sloping rostrum. Furthermore they possess
in relation to Guineobius, plesiomorphic character states such as the small prominent
eyes with few facettes and the squamose vestiture. The males have a long spiral fla-
gellum. However, it must be admitted that no synapomorphy with Guineobius could
be found except the mode of life in the leaf litter and the somewhat smaller eyes in
relation to other Celeuthetini.
PHYLOGENY
Unfortunately four of the six species of Guineobius are known from only one
sex which renders inclusion of genitalic characters into a cladistic analysis difficult.
However, the genitalia possess characters that could be useful for such a purpose
when additional material becomes available. By using the available characters a
preliminary classification was obtained.
The polarity for all characters was determined by outgroup comparison.
Generally the remainder of Celeuthetini was used as an outgroup. To decide the polarity
of characters 3 and 5 where both states occur in in the remainder of Celeuthetini it was
necessary to use two yet undescribed species. These are also dwelling in the leaf-litter
in New Guinea and probably represent a new genus of Celeuthetini which is
presumably closely related to Guineobius (See "systematic position").
In the following character list numbers in brackets indicate the state of
polarity; (0) is plesiomorphic, (1) is apomorphic and (2) more apomorphic.
1. Scales
subovate and appressed or elongate and erect, but always with entire margins (0)
— asymmetrical radiate pectinate (1)
entirely missing (2)
470 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
2. Body length
— large, pronotum plus elytron: 2.4-2.8 mm (0)
small, pronotum plus elytron: 1.6-2.3 mm (1)
3. Antennal scrobes
in dorsal view margins curved inwards, converging obliquely in straight line (0)
— in dorsal view margins kidney-shaped in dorsal view, opposing each other in an
almost straight line (1)
4. Presence of eyes
— vestigial, but distinct (0)
— absent or vestiges extremely small (1)
5. Shape of elytra
— broad at base and apex, medially weakly rounded (0)
— relatively narrow, elongate, evenly rounded from base to apex (1)
6. Abdomen
— sternites V-VI equal or subequal with sternite VI slightly longer (0)
— sternite V at least 2X longer than sternite VI (1)
— sternite V ca. 3X longer than sternite VI (2)
7. Suture between sternites III-IV
— distinct (0)
— completely effaced, or at best only with lateral traces (1)
8. Cuticular coloration
— black (0)
— red-brown (1)
For discussion of characters | and 6 see under "Characters".
G. niger possesses most of the primitive traits, but whether it is really the
sister-group to all remaining Guineobius appears somewhat doubtful. Characters 2, 4
and 8 connecting the two following clades are extremely prone to convergence as
shown by numerous groups of subterranean Coleoptera having lost eyes and mela-
nızation. However, unless convergence can be demonstrated in this case, characters 2,
4 and 8 must be interpreted as synapomorphies.
Character 6 was treated as an ordered transformation series with a gradual
elongation of sternite V. The state in G. minutus (intermediate length) is apomorphic
to the state with a short sternite V, and the state in G. depressus and G. baliemensis
with a long sternite V would be most apomorphic.
G. deharvengei and G. depressus are known from female specimens only. For
that reason male genital characters could not be included in the analysis. Still, it may
be noted that a long thread-like spiral flagellum occurs both in the outgroup-repre-
sentatives and in G. baliemensis. A short simply curved flagellum is found in the
remainder of Guineobius. According to our present hypothesis of Guineobius-rela-
tionships this character is a reversal of the derived state (short flagellum) to the
formerly primitive one (spiral flagellum).
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 471
TABLE Il
Character state matrix for the species of Guineobius.
Character 1 2 3 4 5 6 Uf 8
G. niger 1 Bom Oude 0 0 0 0 0
G. deharvengi 1 1 0 1 1 0 1 1
G. viduus 2 1 0 1 1 0 1 1
G. minutus 2 1 1 1 0 1 0 1
G. depressus 1 1 1 1 0 2 0 1
G. baliemensis 1 1 1 1 0 2 1 1
G. niger
1 G. deharvengi
i O
(1)— G. viduus
2, 4,8 5
(1)— G. minutus
3, 6° G. depressus
6?
LE — G. baliemensis
Fıc. 32
Hypothesized relative relationships of Guineobius-species according to manual reconstruction.
Numbers refer to table 1 and to character list in the text. Convergence in parenthesis.
ECOLOGY AND CONSERVATION
G. niger, G. depressus and G. baliemensis were collected by sieving ground-
litter in a montane forest disturbed by human activity, but especially influenced by
pigs from the nearby villages digging the ground in search for food. The forest was
situated on sandy ground and unusually dry, but moist pockets were found for sieving.
The area is located at an altitude of 1900-2050 m, samples taken at 2300 m did not
contain Guineobius. It was found that Guineobius-species stay motionless for a long
time, even in bright sunshine, and may easily be overlooked during hand-sampling.
Sifted material that was searched in bright sunshine and that was completely dried out
provided a surprising number of Guineobius when put into a Winkler-ecclector over-
night. G. minutus was collected in a slightly disturbed forest that was moister than the
472 ALEXANDER RIEDEL & KLAUS SCHÔNITZER
Fic. 33
Distribution of Guineobius in New Guinea. (BI) G. niger sp. n., G. depressus sp. n. and G. balie-
mensis Sp. n.; (®) G. minutus sp. n.; (%) G. viduus Osella; (A) G. deharvengei Osella.
previous locality, but not as wet as usual montane forests in New Guinea. In those
other extremely mossy and wet forests good results were never obtained for soil-
weevils. It may be that these failures are due to technical problems, since the wet
particles stick together and may not get into the sieve or it takes the beetles even
longer than in drier material before they start moving and can be seen and collected.
But it is possible that the extremely wet mossy forests most common in New Guinea
do not meet the ecological requirements of soil-weevils.
Except for the type locality nothing more is known of the distribution of the
species. However, we expect that their range is very limited (See also below). Based
on this point of view G. niger, G. depressus and G. baliemensis are highly endangered
species. The greatest part of the presumably unique weevil fauna of the valley floor
probably already has become extinct: In the western part of the highlands of New
Guinea the Baliem valley was unique in possessing extensive swamps and gallery
forests at an elevation of about 1700m. They have been cleared before arrival of the
first explorers (ARCHBOLD et al., 1942); probably these extensive clearings date back
as long as 7000 to 5200 years BP (HABERLE er al. 1991). Now increasing pressure for
fire-wood and land is being exerted on relict forests on the slopes of the valley
through the dense and growing population of the fertile valley. Influenced by the
favourable climate of the grand valley these relict forests are usually much drier than
forests of comparable altitude in other parts of the highlands. For this reason an
unique fauna and flora might have developed. To prove this assumption an effective
SOIL-WEEVIL GUINEOBIUS FROM NEW GUINEA 473
protection and exploration of these forests is necessary, otherwise they are likely to
disappear before we can get sufficient information on their biodiversity.
The record of three sympatric species occurring on one small locality is of
some significance. It shows that there must be considerable ecological differentiation
in this genus, allowing their coexistance. One difference between G. niger and G.
depressus is rather obvious from their morphology. The flat and ferruginous G.
depressus seems to be better adapted to a life in deeper litter layers where it would fit
between small gaps in the material, wheras G. niger with its larger, more convex and
black body would be expected to dwell on or a little below the surface of the litter. As
a consequence a much greater number of species than is known today can be expected
from New Guinea. Within a group of slow-moving, apterous insects a lot of "trivial
species" e.g. allopatric species, not very different biologically, with trivial local
adaptions (HOLLOWAY, 1990) can be expected in a country as mountainous as New
Guinea. Probably the same set of ecological niches could be found in each suitable
locality, resulting in a vast number of species.
ACKNOWLEDGEMENT
We would like to thank Dr I. Löbl (Geneva) for the loan of the holotype of G.
deharvengei and G. viduus. We also owe many thanks to Prof. Dr C.W. O'Brien
(Tallahassee), M. Balke (Berlin), Prof. Dr H. Bohn (Munich) and Dr J. Muona (Oulu)
for reviewing and improving the manuscript and to Prof. Dr O. Kraus (Hamburg) for
his help in interpreting the International Code of Zoological Nomenclature. Mrs. R.
Kühbandner (Munich) kindly redrew some of the illustrations.
EIDERATURE CIMED
ARCHBOLD, R., A.L. RAND & L.J. BRASS. 1942. Results of the Archbold expeditions. No. 41.
Summary of the 1938-1939 New Guinea Expedition. Bulletin of the American Museum
of Natural History 79, 197-288, pls. 1-35.
HABERLE, S.G., G.S. HOPE & Y. DEFRETES. 1991. Environmental change in the Baliem Valley,
montane Irian Jaya, Republic of Indonesia. Journal of Biogeography 18, 25-40.
HOLLOWAY, J.D. 1990. Sulawesi biogeography — discussion and summing up. Pp. 95-102 in
W.J. Knight & J.D. Holloway: Insects and the Rain Forests of South East Asia
(Wallacea). 343 pp. London.
MARSHALL, G.A.K. 1956. The Otiorrhynchine Curculionidae of the tribe Celeuthetini (Col.).
134 pp. London.
OsELLA, G. 1983. Nuove specie di Curculionidi delle Regioni Orientale ed Australiana (XXV.
Contributo all conoscenza della curculionidofauna endogea). Revue suisse de Zoologie
90: 111-126.
PAULAY, G. 1985. Adaptive radiation on an isolated oceanic island: the Cryptorhynchinae (Cur-
culionidae) of Rapa revisited. Biological Journal of the Linnean Society 26(2): 95-187.
SHARP, D. 1913. Studies in Rhynchophora. IV. A preliminary note on the male genitalia.
Transactions of the Royal Entomological Society of London 66: 209-222, pl. 9.
THOMPSON, R.T. 1977. A revision of the New Guinea weevil genus Apirocalus Pascoe
(Coleoptera: Curculionidae). Bulletin of the British Museum of Natural History
(Entomology) 36(5): 193-280.
To.
a Bere
uy rer
ed LE
su bese BEN
= CI
REVUE SUISSE DE ZOOLOGIE, 103 (2): 475-524; juin 1996
Revision of the Japanese rove-beetles of the genus Megarthrus
(Coleoptera, Staphylinidae, Proteininae)!
Giulio CUCCODORO
Muséum d'histoire naturelle, Case postale 6434, CH-1211 Genève 6, Switzerland.
Revision of the Japanese rove-beetles of the genus Megarthrus (Coleop-
tera, Staphylinidae, Proteininae). - The Japanese species of Megarthrus
are revised to include 16 species, five of which are described as new: M.
aino, M. constrictus, M. conspirator, M. incubifer and M. sawadai. Megar-
thrus denticollis (Beck) and M. impressicollis Eppelsheim are recorded for
the first time from Japan, and M. japonicus Sharp and M. montanus
Sawada from the Asian continent. Three names are relegated into syno-
nymy: M. heteropus Sawada = M. hemipterus (llliger); M. montanus sub-
angulatus Sawada = M. montanus Sawada; M. subparallelus Sawada = M.
parallelus Sharp). Lectotypes are designated for M. convexus Sharp, M.
corticalis Sharp, M. japonicus Sharp, M. parallelus Sharp and M. scriptus
Sharp. Descriptions are given and diagnostic characters are figured for all
species, except for the widely distributed M. denticollis (Beck) and M.
hemipterus (Illiger). A key to the Japanese species is provided, and
biogeographical relationships are discussed.
Key-words: Staphylinidae - Proteininae - Megarthrus - taxonomy - Japan.
INTRODUCTION
In traditional biogeography a lot of emphasis was put on the delimition of
realms, regions, areas etc. In general the boundaries of zoogeographical realms
defined by WALLACE (1876) are still used today. Two major interfaces, however, are
problematics as the natural boundaries greatly depend on the group studied: the
Nearctic / Neotropical and Palaearctic / Oriental boundaries, respectively. This is
particularly relevant for the fauna of Japan, which is usually considered Palaearctic.
Among other staphylinoid groups, the Batrisinae and Scaphisomatini of Japan
appear Oriental (LÔBL, 1981 & 1984; NOMURA, 1991), whereas Bryaxis Kugelann and
Micropeplus Latreille have more Palaearctic affinities (CAMPBELL, 1968; JEANNEL,
! This paper is part of the author’s Ph.D. thesis at the University of Geneva, funded by
the Swiss National Science Foundation (Project no 31-32331.91).
Manuscript accepted 20.03.1995.
476 GIULIO CUCCODORO
1950). STEEL (1966) considered the Proteinini a Holarctic taxon represented in Japan
by two genera and 16 species (HAYASHI, 1988; SAWADA, 1962) but detailed phylo-
genetic and biogeographical relationships remain unknown.
Within the Proteinini, Megarthrus is of particular interest as it is very diverse
in the Himalaya range (MHNG data) and penetrates into the Southern hemisphere as
far South as New Caledonia, New Guinea (CUCCODORO, 1995), South Africa
(CUCCODORO & LÔBL, 1995) and northern Argentina. Ten species and one subspecies
of Megarthrus have been recorded so far from Japan. The present revision recognises
16 species and briefly discusses their biogeographical relationships.
MATERIAL AND METHODS
The present study is based exclusively on adults (380 specimens). The material
was mostly collected with Berlese and Winkler-Moczarsky eclectors, or with carrion,
dung and Malaise traps. For detailed examination, specimens were dissected, cleared
in 0.1 N potassium hydroxide and mounted in Eukit or Canada balsam on acetate
slides. Drawings were made using a drawing tube. Detailed locality data are repro-
duced according to labels. The name of major administrative units are standardised
according to KANO et al. (1967), and those of G. Lewis’ collecting localities are taken
from PARK (1948). Unless specified otherwise, material mentioned in the text has
been examined. Diagnosis of M. denticollis and M. hemipterus are not given here, as
these widely distributed species will be redescribed elsewhere. Generally only the
distribution of species in Japan is recorded here in detail, but for three East Asian
species - M. conspirator, M. impressicollis, M. japonicus and M. montanus - also
some non Japanese records are included.
The term frons, as used in the present study refers to the area anterior of the U-
shaped impression, the vertex to the area behind. Patches of sensilla on antennomeres
6 to 10 were detected by examining slide preparations. Abdominal sternites and
tergites are counted from the first morphological segment. Measurements and ratios
are defined as follows: length of specimens = interval from middle of anterior
pronotal margin to inner apical angle of elytron; width of specimens = maximum
pronotal width; AL = antennal length / pronotal length; EL = elytral sutural length /
pronotal length; ET = elytral sutural length / shortest interval between sutural margin
and lateral edge of elytron in dorsal view; EW = shortest interval between sutural
margin and outer apical angle of elytron in dorsal view / shortest interval between
sutural margin and lateral edge of elytron in dorsal view; EY = interval between
posterior ocular margin and apex of frons in dorsal view / interval between anterior
and posterior ocular margins in dorsal view; GT = posterior width of gula / median
length of gula; GW = width of neck / posterior width of gula; HW = maximum
pronotal width / interval between posterior ocular margins in dorsal view; ML =
median metasternal length / median mesosternal length; MP = length of segment 4 of
maxillary palpus / length of segment 3 of maxillary palpus; PT = maximum pronotal
width / pronotal length; SP = maximum width of abdominal sternite 8 / width of the
MEGARTHRUS OF JAPAN 477
basal projection; TPF = interval between basal angle and tip of medioapical projection
of female abdominal tergite 8 / lateral length of medioapical projection of female
abdominal tergite 8. The absence of that projection is indicated as “abs”.
Material was examined from the following collections: ASPC = A. Smetana
private collection, Ottawa; BMNH = The Natural History Museum, London; BPBM =
Bishop Museum, Honolulu; CNCI = Canadian National Collection of Insects, Ottawa;
DEI = Deutsches Entomologisches Institut, Eberswalde; FMNH = Field Museum of
Natural History, Chicago; KSPC = K. Sawada private collection, Nishinomiya;
MHNG = Muséum d'histoire naturelle, Geneva; NHMW = Naturhistorisches Mu-
seum, Vienna; ZMHB = Museum für Naturkunde der Humboldt-Universität, Berlin.
NATURAL HISTORY AND ECOLOGY
Except for few a specimens of M. montanus, the Japanese members of Megar-
thrus possess fully developed wings. According to label data they have been found in
forests, plantations and swamps, on and under the bark of logs, in leaf litter, moss,
fungi and decaying vegetable matter.
TAXONOMY
Pending a phylogenetic analysis defining natural species groups, the species
are listed alphabetically.
KEY TO SPECIES
1 elwiragyellow-brownswithiblackespOts iyi. te home aeegs ce 2
Fly ira gun OL OTO USS a. iy omc tas A LE Tee lata 5
2 Male metatibia bearing peg-like setae; female valvifers with lateral
massınsadistan (Eis SA) eee e M. corticalis Sharp
— Male metatibia lacking peg-like setae; female valvifers with lateral
MASINSICONISUOUSIE IS Sad nenn M. scriptus Sharp
3) Temples, in dorsal view, flattened or weakly convex (Fig. 1h)............. 4
— Temples, in dorsal view, strongly convex, subangulate or angulate
CRE ES ET: RE NN. II 7
4 Highest point of the eyes, in frontal view, reaching level of vertex
Ape yen ELI STAI CERTO DIARI D, O M. hemipterus (Illiger)
Highest point of the eyes, in frontal view, not reaching level of vertex. . .... 5
5 Eighth abdominal sternite 3.5-5.0x as wide as its basal projection
Te Cl on deu TO M. denticollis (Beck)
Eighth abdominal sternite 2.0-3.0x as wide as its basal projection. . ........ 6
6 Male metatrochanter lacking peg-like setae; female abdominal tergite
10 bearing two semicircular sclerites (Fig. 27b).......... M. shibatai Sawada
= Male metatrochanter bearing peg-like setae; female abdominal tergite
Weyathont@selentestteselsb) ons RE M. japonicus Sharp
478 GIULIO CUCCODORO
1 Pareraledses of pronotum rounded... Lern. nr. 2 ee 8
— Bateralledges of pronotum angular or/dentieulafe. . ere 15
8 Anterior portion of prothoracic hypomeron with an oblique ridge.......... 9
— Anterior portion of prothoracic hypomeron without ridge............... 10
9 Elytra, in lateral view, with sutural area strongly arcuate....... M. aino sp. n.
- Elytra, in lateral view, with sutural area almost straight
aaa u ee Lapo ASI M. impressicollis Eppelsheim
10 Male with tip of aedeagus incised (Fig. 4g); female abdominal tergite
lO\withteenteriof'apical:cupula hyaline (Fig-52) RE 11
- Male with tip of aedeagus pointed (Fig. 6a); female abdominal tergite
lO >with center of apical eupula sclerotised (Fig 7). fee 12
11 Male with internal sac of aedeagus bearing laterobasal sclerites (Fig.
10b); female with lateral contours of apical portion of valvifers parallel-
Sic cam (sac INR Ar AN I RENE RE AE Se M. convexus Sharp
- Male with internal sac of aedeagus without laterobasal sclerites (Fig.
4a); female with lateral contours of apical portion of valvifers tapering
(IPS A) ae ENS DR LR LR we M. conformis Sawada
12 Male with internal sac of aedeagus bearing laterobasal sclerites (Fig.
8a); female abdominal tergite 10 with cupula bearing a pit (Fig. 9b)
DA SERRA a En ee SA ee ES CO NE M. constrictus Sp. n.
- Male with internal sac of aedeagus lacking laterobasal sclerites (Fig.
6a); female abdominal tergite 10 with cupula lacking a pit (Fig. 9b)
i ears ERE N ORLO ET ORO OO Re aerate EAS M. conspirator sp. n.
18 Anterior portion of prothoracic hypomeron with an oblique groove....... 14
— Anterior portion of prothoracic hypomeron without groove. ............. IS
14 Eronsstormins/anidge above Clypeus tetano we M. incubifer sp. n.
- Frons not forming a ridge above clypeus............. M. montanus Sawada
15 Male metatrochanter bearing peg-like setae; female genital segment as
JG) BE AD ARIA AIS RAR AITRRGIE e EE RE RR A M. parallelus Sharp
- Male metatrochanter lacking peg-like setae (female unknown)
cai: (st a AH AR IR O RN M. sawadai Sp. n.
Megarthrus aino sp. n. (Figs 1g, 2a-g, 3a-f)
TYPE MATERIAL. Holotype d : Japan, Honshu, Iwate pref., Kawai, Yoshibezawa, 600-
700 m, 16.viii.1991 (A. Smetana) #J57, MHNG. Paratypes (8): same data as holotype, 2d in
ASPC and 16 in MHNG; Japan, Honshu, Iwate pref., Sugo, Takizawa, Iwate Un. Exp. For.,
220 m, 11.viii.1991 (A. Smetana) #J44, 15 in ASPC; Kyoto pref., Seryô-Tôge, 13 Km N
Kyoto, 500-600 m, 6.viii.1980 (C. Besuchet) ex leaf litter, moss and old log in plantation of
Cryptomeria japonica, 12 in ASPC; same data, but (I. Löbl) ex leaf litter at foot of rocky cliff,
12 in ASPC and 29 in MHNG.
DISTRIBUTION. Japan: Honshu (Kinki and Tohuku distr.).
MEGARTHRUS OF JAPAN 479
DESCRIPTION. Length 1.1-1.2 mm; width 0.7-0.8 mm. Body and appendages
yellow-brown. Dorsal pubescence fairly uniform, sparser on elytron. Anteriomedian
portion of frons with setae orientated backward. Elytral setae curved and recumbent.
Metasternal setae longer than prosternal setae, becoming denser posteriomedianly.
Abdominal pubescence parallel, becoming longer near apical margins of sternites 4-7.
Punctation on anterior portion of hypomeron coarse, posteriomedial portion of
metasternum impunctate. Frons on level with or raised above level of vertex, forming
a ridge above clypeus; frontal ridge sharp, fine. Anterior frontal edge weakly convex
in middle, oblique laterally. Frontal impression shallow. Eye moderately convex, with
highest point above level of vertex; supra-ocular margin sinuate in dorsal view.
Temple and occipital ridge similar to that in Fig. li, occipital ridge indistinct in
middle and sinuate laterally. Submentum weakly convex. Antenna (Fig. 2c) without
patches of sensilla; scape not flattened; antennomeres 3-4 symmetrical; short and
dense pubescence present on antennomeres 8-11. Pronotum (Fig. 3f) strongly convex
in frontal view; mesal portion slightly arcuate in lateral view. Pronotal disc with deep
depression along apical portion of lateral edge; median groove very shallow, parallel-
sided. Hypomeral ridge present anteriorly, oblique. Median prosternal ridge fine and
straight, interrupted in middle; anterior prosternal margin bordered by a regular row
of conspicuous longitudinal ridges. Protrochanter lacking transverse ridge. Lateral
portion of prepectal ridge angulate, then bifid. Scutellum as in Fig. 1g. Elytron not or
weakly narrowed basally; base abruptly inclined, then overhanging. Humeral callus
low. Elytral disc without swellings, flat along lateral edge; lateral edge finely carinate,
weakly convex in dorsal view; sutural area slightly arcuate basally and strongly
arcuate apically in lateral view; apical margin convex near suture; inner apical angle
obtuse. Metasternum with femoral line arcuate in middle; median ridge absent.
Abdominal tergite 3 with anterior portion flat, then strongly transversely vaulted.
Sternites 2 and 3 with median processes as in Fig. 2g, process of sternite 3 straight.
Sternite 4 flat.
Ratios AE EL 17218) El 1.9: EW 1:2: BY 2:3-2-4-Gi 2: 1-2: 23.G Ws 8"
1.9; HW 1.4-1.5; ML 1.4-1.5; MP 2.0-2.2; PT 2.0-2.1; SP 3.5-4.5; TPF abs.
3. Frontoclypeal area unmodified. Protarsomere 1 lacking tenent setae.
Mesofemur longer than metafemur. Mesotibia about as long as metatibia. Metatarso-
mere | shorter than combined length of metatarsomeres 2-4. Legs unmodified. Apex
of abdominal tergite 8 as in Fig. 2d, e. Sternite 8 as in Fig. 2f. Sternite 10 without
subbasal protuberance. Aedeagus as in Fig. 2a, b.
2. Abdominal tergite 8 (Fig. 3c, d) without medioapical projection. Sternite 8
as in Fig. 3b. Genital segment as in Fig. 3a, e.
COMMENTS. Megarthrus aino, M. conformis, M. conspirator, M. constrictus,
M. convexus, M. corticalis and M. scriptus are the only Japanese species to possess
angulate temples. Among these, M. aino is characterised by the unicolorous elytra and
an oblique hypomeral ridge. It resembles M. minor Coiffait (1976) from Nepal, from
which it may be distinguished by the sexually unmodified mesotibia. See also
comments under M. impressicollis.
480 GIULIO CUCCODORO
Megarthrus conformis Sawada (Figs la, i, 4a-g, Sa-1)
Megarthrus conformis SAWADA, 1962: 11.
TYPE MATERIAL. Holotype 9: Japan, Honshu, Kyoto pref., Kyoto city, Kitashirakawa,
27.x11.1952 (S. Huéno), KSPC, not examined (probably lost, Sawada, pers. comm.).
Additional material (17). Japan, Honshu, Gunma pref., E Usui Pass, 850 m, 24.v11.1980
(I. Löbl) ex rotted wood and leaf litter on slope of ravine with stream in forest, 19 in MHNG;
Nagano pref., Jò-shin-etsu Kogen N.P., Shiga, 1500 m, 23.vii.1980 (I. Löbl) ex leaf litter and
mould on steep rocky slope of ravine with stream in forest, 16 and 49 in MHNG; Shikoku,
Ehime pref., Mt. Ishizuchi N. P., Mt. Ishizuchi, 13-14.v111.1980 (C. Besuchet) ex leaf litter and
grass in dwarf bamboos above forest, 26 and 49 in MHNG; Ehime pref., Mt. Ishizuchi N. P.,
Mt. Ishizuchi, 1350 m, 13.v111.1980 (I. Löbl) ex leaf litter at foot of old Fagus with bamboos,
39 in MHNG; Ehime pref., Saragamine, 30.xii.1954 (K. Morikawa) 1 4 in BMNH; Tokushima
pref., Gy6-ba, 4.viii.1967 (Y. T. Yoshida) 1 © in KSPC.
DISTRIBUTION. Japan: Honshu (Chubu and Kanto distr.) and Shikoku.
DESCRIPTION. Length 1.3-1.5 mm; width 0.9-1.0 mm. Body and appendages
predominantly yellow-brown with vertex, pronotum and scutellum usually darkened.
Dorsal pubescence fairly uniform, sparser on elytron and near lateral edge of pro-
notum. Anteriomedian portion of frons with setae orientated backward. Elytral setae
curved and recumbent. Metasternal setae longer than prosternal setae, fairly uniform.
Abdominal pubescence converging on tergites 4-6, uniform on sternites 4-7, but with
a pair of long subapical setae on each sternite. Punctation fine on anterior portion of
hypomeron and posteriomedial portion of metasternum. Frons raised above level of
vertex, forming a ridge above clypeus; frontal ridge sharp, fine in middle, cons-
picuous laterally. Anterior frontal edge subangulate in middle, oblique laterally. Fron-
tal impression shallow in middle, deep laterally. Eye moderately convex, raised above
level of vertex; supra-ocular margin sinuate in dorsal view. Temple and occipital
ridge as in Fig. li, occipital ridge indistinct in middle and sinuate laterally. Sub-
mentum weakly convex. Antenna (Fig. 5d) without patches of sensilla; scape flatte-
ned; antennomeres 3-4 symmetrical; short and dense pubescence present on anten-
nomeres 7-11. Pronotum (Fig. 5b) weakly convex in frontal view; mesal portion
almost straight in lateral view. Pronotal disc with shallow depression along apical
portion of lateral edge; median groove almost indistinct, parallel-sided. Hypomeral
ridge absent. Median prosternal ridge present anteriorly, fine and straight; anterior
prosternal margin not bordered by longitudinal ridges. Protrochanter lacking trans-
verse ridge. Lateral portion of prepectal ridge angulate, then bifid. Scutellum as in
Fig. la. Elytron weakly narrowed basally: base abruptly inclined, then overhanging.
Humeral callus low. Elytral disc without swellings, shallowly depressed, or flat along
lateral edge; lateral edge conspicuously carinate, weakly convex in dorsal view;
sutural area slightly arcuate basally and strongly arcuate apically in lateral view;
apical margin convex near suture; inner apical angle obtuse. Metasternum with
femoral line arcuate in middle; median ridge absent. Abdominal tergite 3 slightly
transversely vaulted. Sternites 2 and 3 with median processes as in Fig. 51, process of
sternite 3 straight. Sternite 4 flat.
Ratios: AB 1.8-1.9; EL 2:0; ET 2.0; EW 1-0-1015 EY 2.5287 Gil 21 22677
1.8-1.9; HW 1.6-1.8; ML 1.4-1.5; MP 1.9-2.0; PT 2.0-2.1; SP 5.5-7.0; TPF 5.0.
MEGARTHRUS OF JAPAN 481
d. Frontoclypeal area raised, horn-like. Protarsomere 1 lacking tenent setae.
Mesofemur (Fig. 4d) longer than metafemur (Fig. 4c). Mesotibia (Fig. 4f) longer than
metatibia (Fig. 4e). Metatarsomere 1 about as long as combined length of
metatarsomeres 2-4. Peg-like setae absent from protibia and metatrochanter (Fig. 4c);
arranged in a single row on mesotrochanter (Fig. 4d), mesotibia and metatibia. Apex
of abdominal tergite 8 as in Fig. 5h. Sternite 8 as in Fig. 4b. Sternite 10 without
subbasal protuberance. Aedeagus as in Fig. 4a, g.
2. Abdominal tergite 8 (Fig. 5c, f) with medioapical projection. Sternite 8 as
in Fig. 5e. Genital segment as in Fig. 5a, g.
COMMENTS. The definition of this species is based on a female from Gyö-ba
identified as M. conformis by K. Sawada. Its characters agree with the data given in
the original description.
Megarthrus conformis, M. conspirator, M. constrictus and M. convexus differ
from other species with distinctly angulate temples (see comments under M. aino) by
the absence of a hypomeral ridge. Within these species, M. conformis may be
distinguished by the incised aedeagal tip and internal sac lacking laterobasal sclerites.
The former character is shared only with-M. convexus, in which the aedeagal internal
sac bears laterobasal sclerites. See also comments under M. impressicollis.
Megarthrus conspirator sp. n. (Figs 6a-g, 7a-1)
TYPE MATERIAL. Holotype d: Japan, Honshu, Gunma pref., Nikko N. P., below Konsei
Pass, 1500-1600 m, 15.v11.1980 (I. Löbl) ex very wet leaf litter in a ravine, MHNG. Paratypes
(15): same data as holotype, 15 in MHNG; Japan, Honshu, Gunma pref., 4 Km SW Tsumagoi,
1050 m, 18.vii.1980 (I. Löbl) ex trunk with fungi near river, 16 in MHNG; Gunma pref.,
Konsei Pass, 1800-1900 m, 13.viii.1980 (P. M. Hammond) 29 in BMNH and 12 in MHNG;
Iwate pref., Kawai, Yoshibezawa, 600-700 m, 16.viii.1991 (A. Smetana) #J57, 18 in ASPC;
Shizuoka pref., Mt. Amagi, 4.v.1957 (K. Ishida) 16 in BMNH; Tochigi pref., Nikko N. P.,
Senjogahara, 1400 m, 15.vii.1980 (I. Löbl) ex leaf litter near stream, 15 in MHNG; Tochigi
pref., Nikko N. P., Konsei Pass, 1800 m, 15.vii.1980 (I. Lôbl) ex leaf litter under Rhodo-
dendron in very degraded forest on very steep slope, 19 in MHNG; Shikoku, Ehime pref., Mt.
Ishizuchi N. P., Mt. Ishizuchi, 1550 m, 13.viii.1980 (I. Löbl) ex moss rotted wood and leaf litter
under old Abies and Fagus, 18 in MHNG; Ehime pref., Mt. Ishizuchi N. P., Mt. Kamegamori,
1650 m, 15.viii.1980 (S. & J. Peck) ex Abies & moss litter, 15 and 29 in CNCI; Russia,
Sakhalin, Aniva distr., Zunayiskiy Khrebet, Mt. Ishekovsk, 600-1000 m, 13.vii.1993 (A. Pütz
& Wrase) 29 in DEI.
DISTRIBUTION. Japan: Honshu (Chubu, Kanto, and Tohuku distr.) and Shikoku;
Russia: Sakhalin.
DESCRIPTION. Similar to M. conformis from which it differs as follows:
Antenna as in Fig. 6c. Pronotum as in Fig. 7g. Abdominal sternites 2 and 3 with
median processes as in Fig. 71. Ratios: AL 1.8-2.0; EL 2.0-2.2; ET 2.2; TPF abs.
3. Frontoclypeal area raised, horn-like. Protarsomere 1 lacking tenent setae.
Mesofemur (Fig. 6f) longer than metafemur (Fig. 6g). Mesotibia (Fig. 6e) longer than
metatibia (Fig. 6d). Metatarsomere 1 about as long as combined length of
metatarsomeres 2-4. Peg-like setae absent from protibia and metatrochanter (Fig. 6f),
arranged in a single row on mesotrochanter (Fig. 6g), mesotibia and metatibia. Apex
482 GIULIO CUCCODORO
of abdominal tergite 8 as in Fig. 7d. Sternite 8 as in Fig. 7b. Sternite 10 without
subbasal protuberance. Aedeagus as in Fig. 6a, b.
9. Abdominal tergite 8 (Fig. 7c, e) without medioapical projection. Sternite 8
as in Fig. 7h. Genital segment as in Fig. 7a, f.
COMMENTS. Megarthrus conspirator is one of the four species with distinctly
angulate temples and lacking hypomeral ridge (see comments under M. conformis). It
may be distinguished by the pointed aedeagal tip and the internal sac without
laterobasal sclerites. The former character is shared with M. constrictus, in which the
internal sac bears laterobasal sclerites.
Megarthrus constrictus sp. n. (Figs 8a-h, 9a-h)
TYPE MATERIAL. Holotype ¢: Japan, Honshu, Nagano pref., Minami Alps, Kitazawa
Toge, 2000 m, 3.ix.1985 (S. T. Martin) BMNH. Paratypes (7): same data as holotype, 2d, 32
in BMNH and 16,19 in MHNG.
DISTRIBUTION. Japan: Honshu (Chubu distr.).
DESCRIPTION. Similar to M. conformis from which it differs as follows: An-
tenna as in Fig. 9c. Pronotum as in Fig. 9g. Elytral disc shallowly depressed along
lateral edge; lateral edge weakly convex, except for median portion straight in dorsal
view. Abdominal sternites 2 and 3 with median processes as in Fig. 9h. Ratios: EL
ESSE led: IPE abs.
d. Frontoclypeal area raised, horn-like. Protarsomere 1 lacking tenent setae.
Mesofemur (Fig. 8g) longer than metafemur (Fig. 8h). Mesotibia (Fig. se) longer than
metatibia (Fig. 8f). Metatarsomere | about as long as combined length of metatar-
someres 2-4. Peg-like setae absent from protibia and metatrochanter (Fig. 8h),
arranged in a single row on mesotrochanter (Fig. 8g), mesotibia and metatibia. Apex
of abdominal tergite 8 as in Fig. 8b. Sternite 8 as in Fig. 8d. Sternite 10 without
subbasal protuberance. Aedeagus as in Fig. 8a, c.
2. Abdominal tergite 8 (Fig. 9e, f) without medioapical projection. Sternite 8
as in Fig. 9d. Genital segment as in Fig. 9a, b.
COMMENTS. See comments under M. aino, M. conformis and M. conspirator.
Megarthrus convexus Sharp (Figs 10a-h, 11a-h)
Megarthrus convexus SHARP, 1874: 100.
M. convex Sharp; SAWADA, 1962: 11 (incorrect spelling).
TYPE MATERIAL. Lectotype d: "Japan, Simabara, 10.11.1870, G. Lewis" (Kyushu),
BMNH. Paralectotypes (3): same data as lectotype, 22 in BMNH; "Japan, Lewis / 210" 1d in
FMNH, by present designation.
Additional material (13). Japan, Honshu, Gunma pref., Nikko, 1430-1500 m, 11-
12.viii.1980 (L. Maranuma) 26 and 19 in BMNH; Iwate pref., Kawai, Zaimokuzawa, 720 m,
15.viii.1991 (A. Smetana) #J54, 1d in ASPC; Kyoto pref., Kyoto city, Arashiyama, 18-
20.viii. 1980 (C. Besuchet) ex leaf litter and rotted wood in small ravine, 26 and 1 2 in MHNG;
Kyoto pref., Mt. Hiei, Yaseyuen, 4.viii.1980 (C. Besuchet) ex leaf litter in ravine, 16 in
MHNG; Osaka pref., Sakai, Hachi ga ne, Hodo-ji, 1.xii.1985 (K. Harusawa) Berlese, 1d in
MEGARTHRUS OF JAPAN 483
DEI; Shizuoka pref., Ko-Myosan, 26.11.1937 (Y. Saito) 12 in BPBM; Kyushu, Nagasaki,
21.iv.1881 (G. Lewis) 22 in BMNH; Shikoku, Ehime pref., Matsuyama, 15.11.1953 (S.
Hisamatsu) 1 2 in BMNH.
DISTRIBUTION. Japan: Honshu (Chubu, Kanto, Kinki and Tohuku distr.),
Kyushu and Shikoku.
DESCRIPTION. Similar to M. conformis from which it differs as follows:
Antenna as in Fig. Ile. Pronotum as in Fig. 11g. Abdominal sternites 2 and 3 with
median processes as in Fig. 10e. Ratios: AL 1.9-2.1; EL 2.1-2.3; ET 2.1-2.4; ML 1.5-
POPE S10:
d. Frontoclypeal area raised, horn-like. Protarsomere 1 lacking tenent setae.
Mesofemur (Fig. 10g) longer than metafemur (Fig. 10h). Mesotibia (Fig. 10c) longer
than metatibia (Fig. 10f). Metatarsomere 1 about as long as combined length of
metatarsomeres 2-4. Peg-like setae absent from protibia and metatrochanter (Fig.
10h), arranged in a single row on mesotrochanter (Fig. 10g), mesotibia and metatibia.
Apex of abdominal tergite 8 as in Fig. 10d. Sternite 8 as in Fig. Illa. Sternite 10
without subbasal protuberance. Aedeagus as in Fig. 10a-b.
2. Abdominal tergite 8 (Fig. 11b, f) with medioapical projection. Sternite 8 as
in Fig. 1 1c. Genital segment as in Fig. 11d, h.
COMMENTS. See comments under M. aino and M. conformis.
Megarthrus corticalis Sharp (Figs 12a-i, 13a-i)
Megarthrus corticalis SHARP, 1889: 468.
TYPE MATERIAL. Lectotype d: "Japan, Oyayama, 12.111.1881, G. Lewis" (Honshu)
BMNH. Paralectotype (d ): same data as lectotype, BMNH, by present designation.
Additional material (8). Japan (G. Lewis) ex bark, 15 in BMNH; Kuma (Honshu,
Kumagaya?) 14.iii.1881 (G. Lewis) 12 in BMNH; same data, but 17.111.1881 (written
underneath the card bearing the specimen) and with a locality label “Nagasaki, 13.x1.1880-
21.iv.1881”, 16 in FMNH; Honshu, Kyoto distr., Kibune, 9.111.1951 (A. Nobuchi) 16 in
BMNH, Shizuoka pref., Mt. Amagi, 3.v.1957 (Y. Watanabe) 16 in MHNG; Shikoku, Omogo
kei. 15.vi.1956 (M. Miyatake) 12 in BMNH; Mt. Takanawa,, 19.v.1956 (M. Miyatake) 19 in
BMNH; Sakase, Omogo Iyo, 19.vi.1951 (M. Miyatake) 19 in MHNG.
DISTRIBUTION. Japan: Honshu (Chubu and Kinki distr.) and Shikoku.
DESCRIPTION. Length 1.5-1.6 mm; width 1.0-1.1 mm. Body and appendages
predominantly yellow-brown with vertex, pronotum, scutellum and apical margins of
abdominal tergites darkened; elytron with colour pattern as in Fig. 13d. Dorsal
pubescence fairly uniform, sparser on elytron. Anteriomedian portion of frons with
setae orientated backward. Elytral setae curved and recumbent. Metasternal setae
longer than prosternal setae, fairly uniform. Abdominal pubescence converging on
tergites 4-6, uniform on sternites 4-7, but with a pair of long subapical setae on each
sternite. Punctation coarse on anterior portion of hypomeron and posteriomedial
portion of metasternum. Frons raised above level of vertex, forming a ridge above
clypeus; frontal ridge fine, blunt in middle, sharp laterally. Anterior frontal edge
weakly convex in middle, oblique laterally. Frontal impression shallow in middle,
deep laterally. Eye strongly convex, raised above level of vertex; supra-ocular margin
484 GIULIO CUCCODORO
sinuate in dorsal view. Temple and occipital ridge similar to that in Fig. li, occipital
ridge indistinct in middle, sinuate laterally. Submentum weakly convex. Antenna
(Fig. 13c) without patches of sensilla; scape flattened; antennomeres 3-4 symmetrical;
short and dense pubescence present on antennomeres 6-11. Pronotum (Fig. 131)
strongly convex in frontal view; mesal portion almost straight in lateral view. Pronotal
disc with shallow depressions along apical portion of lateral edge and beside median
groove; median groove shallow, widened apically. Hypomeral ridge present
anteriorly, oblique. Median prosternal ridge conspicuous, straight; anterior prosternal
margin bordered by a regular row of conspicuous longitudinal ridges. Protrochanter
lacking transverse ridge. Lateral portion of prepectal ridge angulate, then bifid.
Scutellum as in Fig. la. Elytron weakly narrowed basally; base abruptly inclined, then
overhanging. Humeral callus low. Elytral disc with low swellings, shallowly de-
pressed along lateral edge; lateral edge conspicuously carinate, weakly convex in
dorsal view; sutural area straight basally and strongly arcuate apically in lateral view;
apical margin convex near suture; inner apical angle obtuse. Metasternum with
femoral line arcuate in middle; median ridge absent. Abdominal tergite 3 slightly
transversely vaulted. Sternites 2 and 3 with median processes as in Fig. 13g, process
of sternite 3 straight. Sternite 4 flat at base, then strongly transversely vaulted.
Ratios: AL 2:0;-EL,2.0-2.1; ET 2:0; EW 1:0-1.1,EY2 52/8 CR2225 GIy
1282 HW. 1.6-1.8; ME 1.4-1.5; MP 1.9-2.0; PT 2.0-2.1; SP 31533: TPEvabs:
d. Frontoclypeal area unmodified. Protarsomere 1 lacking tenent setae.
Mesofemur (Fig. 12f) longer than metafemur (Fig. 12g). Mesotibia (Fig. 12d) about
as long as metatibia (Fig. 12e). Metatarsomere | about as long as combined length of
metatarsomeres 2-4. Peg-like setae absent from protibia, arranged in a single row on
mesotrochanter (Fig. 12f), mesotibia, metatrochanter (Fig. 12g) and metatibia. Apex
of abdominal tergite 8 as in Fig. 12c, i. Sternite 8 as in Fig. 12h. Sternite 10 without
subbasal protuberance. Aedeagus as in Fig. 12a, b.
2. Abdominal tergite 8 (Fig. 13e, f) without medioapical projection. Sternite 8
as in Fig. 13h. Genital segment as in Fig. 13a, b.
COMMENTS. Megarthrus corticalis differs from its Japanese congeners, M.
scriptus excepted, by the bicolorous elytra. It may be distinguished easily from M.
scriptus by the symmetrical aedeagus and, in the female, by the lateral margins of the
valvifers widely separated. See also comments under M. aino.
Megarthrus denticollis (Beck)
Omalium denticolle Beck, 1817: 26.
Megarthrus denticollis; ERICHSON, 1840: 906.
TYPE MATERIAL. Unknown. According to the original description from Bavaria, Germany.
Additional material (1). Japan, Honshu, Shiga Pref. (Tshusuki) MHNG.
DISTRIBUTION. Europe; Japan: Honshu (Kinki distr.). New to Japan.
COMMENTS. This species has been over a century currently identified as M.
denticollis (e.g. Erichson, 1840; Ganglbauer, 1895; Lohse, 1964). It is the only
member of the genus occuring in central Europe which has the metatibiae flat and
MEGARTHRUS OF JAPAN 485
wide, and very stout metafemora in male. In Megarthrus denticollis, M. japonicus and
M. shibatai, the highest point of the eyes is below the level of the vertex. Megarthrus
denticollis differs from the other two species by the 8th abdominal sternite which is
3.6-5.0x as wide as its basal projection.
Megarthrus hemipterus (Illiger)
Silpha hemiptera ILLIGER, 1794: 597.
Megarthrus hemipterus; ERICHSON, 1839: 645.
Megarthrus heteropus SAWADA, 1962: 12 (syn. n.).
TYPE MATERIAL. Megarthrus hemipterus: not existing.
Megarthrus heteropus. Holotype, d: Japan, Honshu, Aomori Pref., Spa Tsuta,
19.viii.1959 (T. Shibata) KSPC.
Additional material (105). Japan, Simarima, 25-26.vii.1946 (T. Nakane) 12 in BMNH;
Hokkaido, Sapporo, Usubetsu, Muine Bridge, 600 m, 7.viii.1991 (A. Smetana) #J37, 46 and 39
in ASPC; Honshu, Bibi or Nikko (G. Lewis) 16 and 19 in BMNH; Aomori pref., Fukaura, Mt.
Shirakami, Oku-akaishikawa-rindo, 270-400 m, 14.viii.1991 (A. Smetana) #J53, 85 and 19 in
ASPC; Iwate pref., Kawai, Yoshibezawa, 1050 m, 12.viii.1991 (A. Smetana) #J49, 19 in ASPC;
Nagano pref., Minami Alps, Kitazawa Toge, 2000 m, 3.1x.1985 (S. T. Martin) 14 and 19 in
BMNH; Tochigi pref., Nikko N. P., Senjogahara, 1400 m, 15.v11.1980 (I. Löbl) ex leaf litter near
stream, 16 in MHNG; Wakayama pref, Arida, Shimizu T., Kamiyukawa, Nikko-jinja, 28.viii-
23.1x.1989 (K. Harusawa) ex rotten chicken trap, 25 in DEI; Yamagata pref., Tachiyazawa-
Village, 2.vii.1960 (Y. Watanabe) 1? in BMNH; Shikoku, Ehime pref., Mt. Ishizuchi N. P.,
Tsuchigoya, 1400 m, 11-18.viii.1980 (S. & J. Peck) ex carrion traps in Fagus-Abies forest, 43
and 42 in FMNH; same data, but ex dung cup traps in Fagus-Abies forest, 1d and 29 in
FMNH; same data, but ex malaise trap trough in Fagus-Abies forest, 16 and 19 in FMNH; same
data, but ex berlese, litter in Fagus-Abies forest, logs, stumps, moss and fungi, 19 in FMNH;
Ehime pref., Mt. Ishizuchi N. P., Omogo Valley, 700 m, 18-25.viii.1980 (S. Peck) 126 and 189
in CNCI; same data, but ex rotted watermelon, 3d and 29 in CNCI; same data, but (S. & J.
Peck) ex carrion traps in mixed warm temperate forest, 13¢ and 92, FMNH; same data, but ex
malaise trap trough in mixed warm temperate forest, 1% in FMNH; same data, but ex berlese,
litter under rotting water melon in warm temperate forest, 16 in FMNH; Ehime pref., Omogo-
kei, 23.vini.1953 (M. Miyatake) 16 in BMNH; Ehime pref., Mt. Ishizuchi N. P., Tsuchigoya,
1400 m, 11-18.viii.1980 (S. & J. Peck) ex carrion traps in Fagus-Abies forest, 36 and 29 in
CENCI
DISTRIBUTION. Europe; Russia to Far East, China; Japan: Hokkaido, Honshu
(Tohuku and Chubu distr.) and Shikoku.
COMMENTS. Megarthrus hemipterus is a common Palaearctic species, unique in
Europe by its colour pattern, mentionned in the original description. Thus, it may be
reliably indentified though the type material does not exist. It is distinguished from its
Japanese congeners by the eye which does not reach the level of the vertex, in
combination with the temples similar to that in Fig. 1h.
Megarthrus impressicollis Eppelsheim (Figs le, m, 14a-1)
Megarthrus impressicollis EPPELSHEIM, 1893: 67.
Type material. Holotype 2: "Ost-sibirien, Quellgebiet des Irkut. Leder 1891" NHMW.
Additional material (19). Japan, Honshu, Nagano pref., J6-shin-etsu Kogen N. P.,
Shiga, 1500 m, 23.vii.1980 (I. Löbl) ex leaf litter and mould on steep rocky slope of ravine with
stream in forest, MHNG.
486 GIULIO CUCCODORO
DISTRIBUTION. Russia: Transbaikal; Japan: Honshu (Chubu distr.). New to Japan.
DESCRIPTION. Length 1.4-1.5 mm; width 0.8-0.9 mm. Body and appendages
yellow-brown with vertex somewhat darkened. Dorsal pubescence fairly uniform, spar-
ser on frons, denser along median pronotal groove. Anteriomedian portion of frons with
setae orientated backward. Elytral setae straight and recumbent. Metasternal setae
longer than prosternal setae, becoming sparser posteriomedianly. Abdominal pubes-
cence somewhat converging on tergites 4-6, uniform on sternites 4-7, but with a pair of
long subapical setae on each sternite. Punctation fine on anterior portion of hypomeron
and posteriomedial portion of metasternum. Frons raised above level of vertex; forming
a blunt ridge above clypeus. Anterior frontal edge weakly convex in middle, oblique
laterally. Frontal impression indistinct in middle, deep laterally. Eye almost hemis-
pherical, reaching level of vertex; supra-ocular margin sinuate in dorsal view. Temple
as in Fig. Im. Occipital ridge indistinct. Submentum weakly convex. Antenna (Fig.
14a) without patches of sensilla; scape not flattened; antennomere 3 somewhat asym-
metrical, antennomere 4 symmetrical; short and dense pubescence present on antenno-
meres 7-11. Pronotum (Fig. 141) strongly convex in frontal view; mesal portion straight
in lateral view. Pronotal disc with shallow depression along apical portion of lateral
edge, and deep depression along median groove; median groove shallow, parallel-sided.
Hypomeral ridge present anteriorly, oblique. Median prosternal ridge fine, straight;
anterior prosternal margin bordered by a regular row of fine longitudinal ridges.
Protrochanter lacking transverse ridge. Lateral portion of prepectal ridge sinuate, then
bifid. Scutellum as in Fig. le. Elytron not narrowed basally; base gradually inclined,
then overhanging. Humeral callus low. Elytral disc with low swellings, flat along lateral
edge; lateral edge finely carinate, almost straight in dorsal view; sutural area almost
straight in lateral view; apical margin straight near suture; inner apical angle right-
angled. Metasternum with femoral line arcuate in middle; median ridge absent. Abdo-
minal tergite 3 slightly transversely vaulted. Sternites 2 and 3 with median processes as
in Fig. 14h, process of sternite 3 straight. Sternite 4 flat.
Ratios-AL 2.05 EL 2.07 ET 2.2; EW 1.3; EY 2.6, GT 2:2: (GW. 3ESSEWZWSTE
MEIIFISNIBIRATBITINESZSP2:ITTPFrabs:
2. Abdominal tergite 8 (Fig. 14e, f) without medioapical projection. Sternite 8
as in Fig. 14k. Genital segment as in Fig. 14b-d, g.
3. Unknown.
COMMENTS. Six Japanese species of Megarthrus, M. aino, M. conformis, M.
conspirator, M. constrictus, M. convexus and M. impressicollis, possess uniformly
yellow-brown elytra and convex lateral edges of pronotum. Megarthrus impressicollis
may be distinguished by the sutural area almost straight in lateral view. The shape of
the apical abdominal tergites in female is particularly notable.
Megarthrus incubifer sp. n. (Figs If, k, 15a-1, 16a-g)
TYPE MATERIAL. Holotype d: Japan (G. Lewis), BMNH. Paratypes (29): Japan,
Hokkaido, Mt. Daisetsu, 3.vii.1958 (M. Miyatake) MHNG; Shikoku, Ehime pref., Mt. Ishizuchi
N.P., Tsuchigoya, 1400 m, 11-18.viii.1980 (S. & J. Peck) ex malaise trap trough in Fagus-
Abies forest, FMNH.
MEGARTHRUS OF JAPAN 487
DISTRIBUTION. Japan: Hokkaido and Shikoku.
DESCRIPTION. Length 1.4-1.5 mm; width 1.0-1.1 mm. Body uniformly dark
brown with appendages paler. Dorsal pubescence fairly uniform, sparser on elytron,
denser near medioapical margin of abdominal tergite 7. Anteriomedian portion of
frons with setae orientated forward. Elytral setae curved and recumbent. Metasternal
setae as long as prosternal setae, fairly uniform. Abdominal pubescence converging
on tergites 4-5, uniform on sternites 4-7, but with a pair of long subapical setae on
each sternite. Punctation coarse on anterior portion of hypomeron and posteriomedial
portion of metasternum. Frons raised above level of vertex; forming a ridge above
clypeus; frontal ridge sharp, fine. Anterior frontal edge evenly convex. Frontal
impression deep. Eye almost hemispherical, with highest point above level of vertex;
supra-ocular margin sinuate in dorsal view. Temple as in Fig. Ik. Occipital ridge
indistinct. Submentum weakly convex. Antenna (Fig. 16a) without patches of sensilla;
scape not flattened; antennomeres 3-4 symmetrical; short and dense pubescence
present on antennomeres 5-11. Pronotum (Fig. 16e) strongly convex in frontal view;
mesal portion almost straight in lateral view. Pronotal disc with deep depressions
along apical portion of lateral edge, and beside median groove; median groove deep,
parallel-sided. Hypomeral ridge present anteriorly, oblique. Median prosternal ridge
conspicuous, Y-shaped; anterior prosternal margin bordered by a regular row of
conspicuous longitudinal ridges. Protrochanter lacking transverse ridge. Lateral
portion of prepectal ridge sinuate, then bifid. Scutellum as in Fig. 1f. Elytron not
narrowed basally; base abruptly inclined, then overhanging. Humeral callus low.
Elytral disc with low swellings, flat along lateral edge; lateral edge finely carinate,
weakly convex in dorsal view; sutural area slightly and evenly arcuate in lateral view;
apical margin straight near suture; inner apical angle obtuse. Metasternum with
femoral line arcuate in middle; median ridge absent. Abdominal tergite 3 slightly
transversely vaulted. Sternites 2 and 3 with median processes as in Fig. 15e, process
of sternite 3 straight. Sternite 4 flat at base, then stongly transversely vaulted.
Ratios Ate 2. 1-2 2B 6-7 ET I GAME EWi LIE? SB Ye? 25 Gily2.02
2.3; GW 1.9-2.0; HW 1.8-1.9; ML 1.5-1.6; MP 1.8-2.0; PT 2.0-2.1; SP 2.9-3.0; TPF
abs.
dé. Frontoclypeal area unmodified. Protarsomere 1 lacking tenent setae.
Mesofemur (Fig. 15g) longer than metafemur. Mesotibia (Fig. 15c) shorter than
metatibia (Fig. 15d). Metatarsomere 1 about as long as combined length of
metatarsomeres 2-4. Peg-like setae arranged in a single row on mesotrochanter (Fig.
15g), mesotibia and metatibia, absent from protibia and metatrochanter. Apex of
abdominal tergite 8 as in Fig. 15f, i. Sternite 8 as in Fig. 15h. Sternite 10 without
subbasal protuberance. Aedeagus as in Fig. 15a, b.
2. Abdominal tergite 8 (Fig. 16c, d) without medioapical projection. Sternite 8
as in Fig. 16b. Genital segment as in Fig. 16f, g.
COMMENTS. Megarthrus incubifer and M. montanus differ from other Japanese
members of the genus by their uniformly dark brown elytra and oblique hypomeral
ridge. The former species may be distinguished by the Y-shaped prosternal ridge.
488 GIULIO CUCCODORO
Megarthrus japonicus Sharp (Figs Ic, h, 17a-k, 18a-h)
Megarthrus japonicus SHARP, 1874: 99.
TYPE MATERIAL. Lectotype d : Japan (G. Lewis) BMNH. Paralectotypes (14): same data
as lectotype, 78,62, BMNH and 1 d, FMNH, by present designation.
Additional material (50). China, Fukien prov., Chungan, Bohea Hills, 16.x1.1940 (T. C.
Maa) 15 in BPBM; Japan (V. Quedenfeldt) # 59 037, 18 in MHNG and 26 in ZMHB;
(Hiller) 22 in ZMHB; (Shönfeldt) 19 in DEI; Yokkaichi, Meer (Sauter) 16 in FMNH;
Honshu, Gose pref., Mt. Kongo (= Watamaya pref., Mt. Kongodoji) 1100 m, 30.v11.1980 (C.
Besuchet) ex leaf litter in forest, 12 in MHNG; Hyogo pref., Kobe, 25.1.1933, 2d in BMNH;
Hyogo pref., Kobe, ii-v.1930, 12, BMNH; Hyogo pref., Mayasan, 15.x.1928 (J. Ea?
unreadable, Lewis) 1 2 in BMNH; Kanagawa pref. (Sauter) 29 in FMNH; Kyoto distr., Kyoto,
Midorogaike Pond, 7-8.viii.1980 (C. Besuchet) 19 in MHNG; Tokyo pref., Tokyo, 1925, 1d,
32 in BMNH and 19 in MHNG; Tokyo, iv.1913 (F. Muir) 22 in BPBM; Tokyo, 1925 (Koch)
1d in FMNH; Tokyo pref., Inaginaganuma, 1.iv.1959 (Y. Watanabe) 1 din BMNH; Kanagawa
pref., Yokohama (G. Lewis) 26, 59 in BMNH and 16, 12 in MHNG; Kyushu, Nagasaki
pref., 7.111.1881 (G. Lewis) 18 in BMNH; same data but 9.iii.1881, 12 in BMNH; same data
but 3.iv.1881, 36 in BMNH: same data but 13.xi.1880-21.iv.1881, 26 and 32 in BMNH;
Shikoku, Ehime pref., Matsuyama, 12.v.1951 (S. Hisamatsu) 16 in BMNH; same data, but
9.11.1947 (M. Miyatake) 1d and 19 in BMNH; Kochi pref., Kuroson, 28.iv.1956 (S. Hisa-
matsu) 24 in BMNH and 14 in MHNG; Russia, Primorskij Kraj, Vladivostok, 11.vii.1993 (A.
Pütz) 16 in DEI; Sakhalin, Aniva distr., Nuovo Alexandrovsk, 10 Km N Yusshno-Sakhalinsk,
Zuzuya river, 12.vii.1993 (A. Pütz & Wrase) 19 in DEI.
DISTRIBUTION. Far East Russia; China: Fukien prov.; Japan: Honshu (Kanto
and Kinki distr.), Kyushu and Shikoku. New to China and Russia.
DESCRIPTION. Length 1.3-1.4 mm; width 0.8-0.9 mm. Body predominantly dark
brown or blackish with pronotum, elytron and appendages paler, sutural margin of
elytron darkened; antennomeres 1-4 somewhat paler than antennomeres 5-11. Dorsal
pubescence fairly uniform, somewhat denser along median pronotal groove, near
humeral area of elytron and apical margin of abdominal tergite 7. Anteriomedian por-
tion of frons with setae orientated backward. Elytral setae curved and recumbent.
Metasternal setae shorter than prosternal setae, becoming sparser posteriomedianly.
Abdominal pubescence parallel, uniform on sternites 4-7, but with a pair of long
subapical setae on each sternite. Punctation fine on anterior portion of hypomeron,
posteriomedial portion of metasternum impunctate. Frons raised above level of
vertex, forming a blunt ridge above clypeus. Anterior frontal edge evenly convex.
Frontal impression shallow or indistinct. Eye strongly convex, with highest point
below level of vertex; supra-ocular margin sinuate in dorsal view. Temple similar to
that in Fig. 1h. Occipital ridge indistinct. Submentum almost flat. Antenna (Fig. 18c)
with patches of sensilla on antennomeres 6-10; scape not flattened; antennomere 3
somewhat asymmetrical, antennomere 4 strongly asymmetrical; short and dense
pubescence present on antennomeres 5-11. Pronotum (Fig. 18h) weakly convex in
frontal view; mesal portion almost straight in lateral view. Pronotal disc not
depressed; median groove shallow, parallel-sided. Hypomeral ridge absent. Median
prosternal ridge absent; anterior prosternal margin bordered by an irregular row of
fine longitudinal ridges. Protrochanter lacking transverse ridge. Lateral portion of
prepectal ridge straight, then bifid. Scutellum similar to that in Fig. lc. Elytron not
MEGARTHRUS OF JAPAN 489
narrowed basally; base gradually inclined, then vertical. Humeral callus low. Elytral
disc with low swellings, shallowly depressed along apical portion of lateral edge;
lateral edge finely carinate, straight in dorsal view; sutural area almost straight in
lateral view; apical margin somewhat sinuate near suture; inner apical angle right-
angled. Metasternum with femoral line arcuate in middle; median ridge present
posteriorly, fine and low. Abdominal tergite 3 slightly transversely vaulted. Sternites
2 and 3 with median processes as in Fig. 17h, process of sternite 3 straight. Sternite 4
flat at base, then slightly transversely vaulted.
Ratios: AL 1.8-2.0; EL 1.8-2.0; ET 1.8-2.0; EW 1.2: EY 2.3-2.5; GT 2.1-2.2;
GW 1.7-1.8; HW 1.6-1.7; ML 1.6-1.7; MP 1.5-1.7; PT 2.2; SP 2.0-3.0; TPF 7.0-8.0.
d . Frontoclypeal area unmodified. Frontal pubescence parallel. Protarsomere 1
with tenent setae. Metafemur (Fig. 17k) longer than mesofemur (Fig. 17g). Metatibia
(Fig. 17d) longer than mesotibia (Fig. 17c). Metatarsomere 1 about as long as
combined length of metatarsomeres 2-4. Peg-like setae absent from protibia, arranged
in a single row on mesotrochanter (Fig. 17g), grouped to from a field on mesotibia
and metatrochanter (Fig. 17k), arranged in a single row on metatibia, but near
metatibial apex grouped to form a field. Apex of abdominal tergite 8 as in Fig. 17e, f.
Sternite 8 as in Fig. 18g. Sternite 10 with small subbasal protuberance (Fig. 171).
Aedeagus as in Fig. 17a, b.
2. Frontal pubescence converging. Abdominal tergite 8 (Fig. 18c, e) with
medioapical projection. Sternite 8 as in Fig. 18f. Genital segment as in Fig. 18a, b.
COMMENTS. Megarthrus japonicus differs from other Japanese congeners, M.
shibatai excepted, by the eyes which do not reach the level of the vertex, in combi-
nation with the 8th abdominal sternite which is 2.0-3.0x as wide as its basal pro-
jection. It differs from M. shibatai by the male metatrochanter bearing peg-like setae
and, in female, by the hyaline 10th abdominal tergite. See also comments under M.
denticollis.
Megarthrus montanus Sawada (Figs Id, n, 19a-1, 20a-h)
Megarthrus montanus SAWADA, 1962: 14.
Megarthrus montanus subangulatus SAWADA, 1962: 14 (syn.n.).
TYPE MATERIAL. Megarthrus montanus. Holotype, d : Japan, Honshu, Nagano pref.,
Inago-Yu, 27.vii.1959 (T. Shibata) KSPC. Paratype, 2: same data as holotype, KSPC.
Megarthrus montanus subangulatus. Holotype, 2: Japan, Honshu, Kanagawa pref.,
Osaka, Katsuoji, 24.xii.1954 (K. Sawada) KSPC. Paratype, 2: Japan, Honshu, Kii, Mt. Kôjin,
1.v11.1956 (M. Hayashi) KSPC.
Additional material (10). China, Heilungkiang prov., Harbin, 16.vi.1966 (P. M.
Hammond) #1364, 19 in BMNH; Japan, Honshu, Gunma pref., below Usui Pass, 750 m,
.20.v11.1980 (I. Lobl) ex leaf litter in ravine, 19 in MHNG; Gunma pref., Nikko, Kozawa, 1000
m, 15.viii.1980 (P. M. Hammond) 19 in BMNH; Iwate pref., Kawai, Yoshibezawa, 1050 m,
12.v111.1991, #J49 (A. Smetana), 16 in ASPC; Russia, slopes of Parwaja Rjetschka, Valley N
Wladiwostok, 1918-1920 (H. Frieb) 2? in NHMW; Primorskiy Kray, Ussuriysky Zapovednik,
33 Km SE Ussuriysk <43°37' N; 132°18' E> 500 m, 13.vi.1993 (L. Zerche) 16, 12 in DEI and
16,19 in MHNG.
DISTRIBUTION. Northeastern China; Japan: Honshu (Chubu, Kanto and Tohuku
distr.); Far East Russia. New to Russia and China.
490 GIULIO CUCCODORO
DESCRIPTION. Length 1.5-1.6 mm; width 0.9-1.1 mm. Body uniformly dark
brown with sutural margin of elytron darkened, appendages paler; antennomeres 1-4
paler than antennomeres 5-11. Dorsal pubescence fairly uniform, sparser on elytron,
denser near humeral area. Anteriomedian portion of frons with setae orientated
backward. Elytral setae curved and recumbent. Metasternal setae as long as prosternal
setae, becoming denser posteriomedianly. Abdominal pubescence converging on
tergites 4-6, uniform on sternites 4-7. Punctation coarse on anterior portion of
hypomeron, fine on posteriomedial portion of metasternum. Frons raised above level
of vertex, evenly deflected toward clypeus. Anterior frontal edge weakly convex in
middle, oblique or sinuate laterally. Frontal impression shallow in middle, deep
laterally. Eye almost hemispherical, reaching level of vertex; supra-ocular margin
sinuate in dorsal view. Temple and occipital ridge as in Fig. In, occipital ridge
straight in middle and sinuate laterally. Submentum almost flat. Antenna (Fig. 20c)
without patches of sensilla; scape not flattened; antennomere 3 slightly asymmetrical,
4 symmetrical; short and dense pubescence present on antennomeres 5-11. Pronotum
(Fig. 20g) strongly convex in frontal view; mesal portion slightly arcuate in lateral
view. Pronotal disc with deep depression along apical portion of lateral edge, and
shallow depression beside median groove; median groove shallow, parallel-sided.
Hypomeral ridge present anteriorly, oblique. Median prosternal ridge fine, straight,
interrupted in middle; anterior prosternal margin bordered by a regular row of
conspicuous longitudinal ridges. Protrochanter lacking transverse ridge. Lateral
portion of prepectal ridge sinuate, then bifid. Scutellum as in Fig. Id. Elytron weakly
narrowed basally; base abruptly inclined, then overhanging. Humeral callus low.
Elytral disc with low swellings, flat along lateral edge; lateral edge finely carinate,
weakly convex, or straight, in dorsal view; sutural area straight basally and slightly
arcuate apically in lateral view; apical margin weakly convex near suture; inner apical
angle obtuse. Metasternum with femoral line arcuate in middle; median ridge absent.
Abdominal tergite 3 slightly transversely vaulted. Sternites 2 and 3 with median
processes as in Fig. 20h, process of sternite 3 trifid. Sternite 4 flat.
Ratios: AL 1.7-1.8; EL 1.7-1.9; ET 1.8-2.0; EW 1.1-1:2>EX 2428567252
2.6; GW 1.7-1.8; HW 1.8-2.0; ML 1.3-1.9; MP 1.4-1.5; PT 2.0; SP 1.8-2-3; TPF abs.
dé. Frontoclypeal area unmodified. Protarsomere 1 lacking tenent setae.
Mesofemur (Fig. 19b) longer than metafemur. Mesotibia (Fig. 19d) shorter than
metatibia. Metatarsomere | about as long as combined length of metatarsomeres 2-4.
Peg-like setae absent from mesotrochanter (Fig. 19b), metatrochanter (Fig. 19c) and
metatibia, grouped to form a field on mesotibia. Apex of abdominal tergite 8 as in
Fig. 19h, 1. Sternite 8 as in Fig. 19e. Sternite 10 (Fig. 19g) with large subbasal protu-
berance. Aedeagus as in Fig. 19a, f.
2. Abdominal tergite 8 (Fig. 20e, f) without medioapical projection. Sternite 8
as in Fig. 20d. Genital segment as in Fig. 20a, b.
COMMENTS. Megarthrus montanus is characterised by the male mesotibia
deeply notched and, in female, the V-shaped posterior margin of the 8th abdominal
sternite. See also comments under M. incubifer.
MEGARTHRUS OF JAPAN 491
The two females collected respectively by I. Löbl and P. M. Hammond possess
smaller eyes, shortened wings and metasternum.
Megarthrus parallelus Sharp (Figs 1b, 21a-1, 22a-h)
Megarthrus parallelus SHARP, 1874: 99.
Megarthrus subparallelus SAWADA, 1962: 12 (syn.n.).
TYPE MATERIAL. Megarthrus parallelus. Lectotype, 9: "Japan, G. Lewis", BMNH.
Paralectotype, 2: same data as lectotype, BMNH, by present designation.
Megarthrus subparallelus. Holotype, 3: Japan, Honshu, Osaka pref., Minoo, Katsuoji,
24.x11.1954 (K. Sawada) KSPC. Paratype, 9 : Japan, Honshu, Nara pref., Mt Kasuga, 15.1.1953
(G. Imadate) KSPC.
Additional material (8). Japan (G. Lewis) 29 in BMNH, same data, but #228 [labelled
as "Megarthrus parallelus var?"] 19 in FMNH; Kiga (G. Lewis) 1? in BMNH; Fukuoji, Aki,
12.iv.1955 (M. Miyatake) 16 in BMNH; Honshu, Hyogo pref., Kobe, Mayasam, 13.vi.30, 350
m, 12 in BMNH; Kyushu, Nagasaki pref., 23.111.1881 (G. Lewis) 12 in BMNH; Shikoku,
Shirainot Aki, 13.xi.1949 (M. Miyatake) 19 in BMNH.
DISTRIBUTION. Japan: Honshu (Kinki distr.), Kyushu and Shikoku.
DESCRIPTION. Length 1.2-1.4 mm; width 0.8-0.9 mm. Body uniformly dark
brown with sutural margin of elytron darkened, appendages somewhat paler. Dorsal
pubescence fairly uniform, sparser on elytron. Anteriomedian portion of frons with
setae orientated forward. Elytral setae curved and recumbent. Metasternal setae as
long as prosternal setae, becoming denser anteriorly. Abdominal pubescence con-
verging on tergite 4, uniform on sternites 4-7, but with a pair of long subapical setae
on each sternite. Punctation fine on anterior portion of hypomeron, coarse on pos-
teriomedial portion of metasternum. Frons raised above level of vertex, forming a
ridge above clypeus; frontal ridge sharp, conspicuous. Anterior frontal edge evenly
convex. Frontal impression deep. Eye almost hemispherical, highest point somewhat
above level of vertex; supra-ocular margin sinuate in dorsal view. Temple as in Fig.
Im. Occipital ridge indistinct. Submentum weakly convex. Antenna (Fig. 22c)
without patches of sensilla; scape not flattened; antennomeres 3-4 symmetrical; short
and dense pubescence present on antennomeres 7-11. Pronotum (Fig. 22d) strongly
convex in frontal view; mesal portion almost straight in lateral view. Pronotal disc
with shallow depression along apical portion of lateral edge; median groove shallow,
parallel-sided. Hypomeral ridge present anteriorly, oblique. Median prosternal ridge
conspicuous and straight anteriorly, indistinct posteriorly; anterior prosternal margin
bordered by a regular row of conspicuous longitudinal ridges. Protrochanter lacking
transverse ridge. Lateral portion of prepectal ridge sinuate, then bifid. Scutellum as in
Fig. 1b. Elytron not narrowed basally; base abruptly inclined, then overhanging.
Humeral callus low. Elytral disc with low swellings, flat along lateral edge; lateral
edge finely carinate, almost straight in dorsal view; sutural area slightly and evenly
arcuate in lateral view; apical margin straight near suture; inner apical angle obtuse.
Metasternum with femoral line arcuate in middle; median ridge present anteriorly,
fine and low. Abdominal tergite 3 slightly transversely vaulted. Sternites 2 and 3 with
median processes as in Fig. 21e, process of sternite 3 straight. Sternite 4 flat at base,
then slightly transversely vaulted.
492 GIULIO CUCCODORO
Ratios: AL 2.1-2.2; EL 1.9-2.0; ET 1.7-1.8; EW 1.0-1.1; EY 2.6-2.7; GT 2.2-
2.3; GW 1.9-2.0; HW 1.6-1.7; ML 1.7-2.2; MP 1.9-2.1; PT 2.0-2.1; SP 3.4-3.7; TPF
abs.
4. Frontoclypeal area unmodified. Protarsomere 1 lacking tenent setae. Meso-
femur (Fig. 21h) longer than metafemur (Fig. 21k). Mesotibia (Fig. 21d) longer than
metatibia (Fig. 21c). Metatarsomere 1 about 1.5x as long as combined length of meta-
tarsomeres 2-4. Peg-like setae absent from protibia, arranged in a single row on meso-
trochanter (Fig. 21h), grouped to form a field on mesotibia, metatrochanter (Fig. 21k)
and metatibia. Apex of abdominal tergite 8 as in Fig. 21f, i. Sternite 8 as in Fig. 22g.
Sternite 10 (Fig. 21g) with small subbasal protuberance. Aedeagus as in Fig. 21a, b.
®. Abdominal tergite 8 (Fig. 22f, h) without medioapical projection. Sternite 8
as in Fig. 22e. Genital segment as in Fig. 22a, b.
COMMENTS. Among the Japanese species of the genus, M. parallelus and M.
sawadai are characterised by the uniformly dark brown elytra and the lateral edges of
the pronotum forming six distinct denticles. Megarthrus parallelus differs from M.
sawadai by the aedeagal characters and the shape of the legs in male.
Megarthrus sawadai sp. n. (Fig. 23a-n)
TYPE MATERIAL. Holotype d : Japan, Honshu, Gunma pref., below Usui Pass, 750 m,
20.vii.1980 (I. Löbl) ex leaf litter in ravine, MHNG. Paratype, d : Japan, Honshu, Kanagawa
pref., Monomitouge, 18.1v.1959 (Y. Watanabe) BMNH.
DISTRIBUTION. Japan: Honshu (Kanto distr.).
DESCRIPTION. Similar to M. parallelus from which it may be distinguished as
follows: Length 1.2 mm; width 0.8 mm. Anterior frontal edge weakly convex in
middle and oblique laterally. Antenna as in Fig. 231. Pronotum as in Fig. 23g. Abdo-
minal sternites 2 and 3 with median processes as in Fig. 23n. Ratios: EL 1.9-2.0; GW
1.7-1.8; ML 1.5-1.6; SP 3.1-3.5.
dé. Frontoclypeal area unmodified. Protarsomeres 1 lacking tenent setae.
Mesofemur (Fig. 23k) longer than metafemur. Mesotibia (Fig. 23b) longer than
metatibia (Fig. 23c). Metatarsomeres | about as long as combined length of
metatarsomeres 2-4. Peg-like setae absent from protibia and mesotrochanter, arranged
in a single row on mesotrochanter (Fig. 23k) and metatibia, arranged in a double row
on mesotibia, but near mesotibial apex arranged in a single row. Apex of abdominal
tergite 8 as in Fig. 23h, m. Sternite 8 as in Fig. 23e. Sternite 10 (Fig. 231) with small
subbasal protuberance. Aedeagus as in Fig. 23a, d.
2. Unknown.
COMMENTS. See discusion under M. parallelus.
ETYMOLOGY. The species is named in honor of Mr Kohei Sawada, Nishi-
nomyia.
Megarthrus scriptus Sharp (Figs 24a-g, 24a-1)
Megarthrus scriptus SHARP, 1889: 469.
TYPE MATERIAL. Lectotype d : Japan, Honshu, Gunma pref., Chuzenji, 19.viii.1881 (G.
Lewis) BMNH, by present designation.
MEGARTHRUS OF JAPAN 493
Additional material (89). Japan, Honshu, Aomori pref., Fukaura, Mt. Shirakami, Oku-
akaishikawa-rindo, 270-400 m, 14.viii.1991 (A. Smetana) #J53, 24 in ASPC; Gifu pref., 8 Km
SE Osaka, 750 m, 1.vin.1980 (I. Löbl) on and under bark of rotted trunk in Cryptomeria
plantation, 45 and 4? in MHNG: Gunma pref., below Usui Pass, 700 m, 20.vii.1980 (I. Löbl) ex
grass in abandoned field, 34 and 62 in MHNG; Gunma pref., E Usui Pass, 850 m, 24.vii.1980
(I. Löbl) ex rotted wood and leaf litter, forest on slope, 46 and 39 in MHNG; Gunma pref., J6-
shin-etsu Kogen N. P., Shirane, 1500 m, 22.vii.1980 (I. Löbl) ex swamp, 16 and 29 in MHNG;
Gunma pref., Chuzenji, Nataisan, 18.viii.1965 (D. H., A. C. & A. H. Kistner) field # 1002, 24
and 32 in BPBM; Iwate pref., Iwaizumi, Hitsutori, 790 m, 11.viii.1991 (A. Smetana) #J45, 19
in ASPC; Iwate pref., Kawai, Yoshibezawa, 1050 m, 12.viii.1991 (A. Smetana) #J49, 79 in
ASPC; Kanagawa pref., Sagami, Ohtakitoge, Nishi-tanzawa, 12.vi.1939 (H. Araki) 15 in
BMNH; Nagano pref., Jö-shin-etsu Kogen N. P., Shiga, 1500 m, 23.v11.1980 (I. Löbl) under bark
of dead Abies, 12 in MHNG; Nikko pref., 3-21.vi.1880 (G. Lewis) 16 in BMNH; Shizuoka
pref., Mt.Amagi, 18.v.1957 (K. Ishida) 16 in BMNH; Tottori pref., Dai-sen-oki, 8.viii.1933 (H.
Araki) 26 in BMNH; Shikoku, Ehime pref., Mt. Ishizuchi N. P., Tsuchigoya, 1400 m, 11-
18.vii1.1980 (S. & J. Peck) berlese, litter, logs, stumps, moss & fungi in Fagus-Abies forest, 1d
in FMNH; same data, but ex malaise trap trough in Fagus-Abies forest, 19 in FMNH; Ehime
pref., Mt. Ishizuchi N. P., Mt. Kamegamori, Siraza Pass, 1500 m, 15.viii.1980 (S. Peck) ex Fagus
logs, 16 and 32 in CNCI; Ehime pref., Mt. Ishizuchi N. P., Omogo Valley, 700 m, 18-
25.vii.1980 (S. Peck) 29 in CNCI; same data, but ex fungi on log & moss in mixed warm
temperate forest, 2d and 19 in CNCI; same data, but (S. & J. Peck), litter under rotting
watermelon, berlese, fungi on mossy logs, and malaise trap, 66 and 99 in FMNH; Ehime pref.,
Mt. Ishizuchi N. P., Mt. Ishizuchi, 1350 m, 13.viii.1980 (I. Löbl) ex leaf litter at foot of old
Fagus with bamboos, 19 in MHNG; Ehime pref., Omogo-kei, 13.vi.1954 (S. Hisamatsu) 19 in
BMNH; Ehime pref., Omogo, Sakase, 19.vi.1955 (M. Miyatake) 12 in BMNH; Ehime pref.,
Omogo-kei, 21, 22, 23.viii.1953 (M. Miyatake) 2d, 12 in BMNH; same data, but 1 and
2.vili.1954, 39, 21.viii.1955, 19, 27.viii.1955, 36, and 15.vi.1956, 19 in BMNH; Ehime pref.,
Saragamine, 12.xii.1953 (T. Mohri) 19 in BMNH.
DISTRIBUTION. Japan: Honshu (Chubu, Kanto and Tohuku distr.) and Shikoku.
DESCRIPTION. Length 1.3-1.5 mm; width 0.8-1.0 mm. Body and appendages
predominantly yellow-brown with vertex and pronotum occasionnally darkened;
elytra with colour pattern as in Fig. 25c. Dorsal pubescence fairly uniform, sparser on
elytron. Anteriomedian portion of frons with setae orientated backward. Elytral setae
curved and recumbent. Metasternal setae as long as prosternal setae, becoming denser
posteriomedianly. Abdominal pubescence converging on tergites 4-6, uniform on
sternites 4-7, but with a pair of long subapical setae on each sternite. Punctation fine
on anterior portion of hypomeron, posteriomedial portion of metasternum impunctate.
Frons raised above level of vertex, forming a ridge above clypeus; frontal ridge sharp,
fine. Anterior frontal edge strongly convex in middle, oblique laterally. Frontal im-
pression shallow in middle, deep laterally. Eye strongly convex, reaching level of
vertex; supra-ocular margin sinuate in dorsal view. Temple and occipital ridge as in
Fig. li, occipital ridge indistinct in middle and sinuate laterally. Submentum weakly
convex. Antenna (Fig. 25d) without patches of sensilla; scape not flattened; antenno-
meres 3-4 symmetrical; short and dense pubescence present on antennomeres 6-11.
Pronotum (Fig. 25g) weakly convex in frontal view; mesal portion almost straight in
lateral view. Pronotal disc with shallow depressions along apical portion of lateral
edge and beside median groove; median groove shallow, widened apically. Hypo-
meral ridge present anteriorly, oblique. Median prosternal ridge conspicuous, straight;
anterior prosternal margin bordered by a regular row of conspicuous longitudinal
494 GIULIO CUCCODORO
ridges. Protrochanter lacking transverse ridge. Lateral portion of prepectal ridge an-
gulate, then bifid. Scutellum as in Fig. la. Elytron not narrowed basally; base abruptly
inclined, then overhanging. Humeral callus low. Elytral disc with low swellings, flat
along lateral edge; lateral edge conspicuously carinate, moderately convex in dorsal
view; sutural area slightly and evenly arcuate in lateral view; apical margin convex near
suture; inner apical angle obtuse. Metasternum with femoral line arcuate in middle;
median ridge absent. Abdominal tergite 3 slightly transversely vaulted. Sternites 2 and
3 with median processes as in Fig. 25h, process of sternite 3 straight. Sternite 4 flat.
Ratios: AL 1.8-2.0: EL 2.02.12 ET 2:1-2.2; EW 11-1 22EVY22233G a
2.1; GW 1.8; HW 1.6-1.8; ML 1.1-1.3; MP 1.6-1.8; PT 2.0; SP 2.5-3.0; TPF abs.
d. Frontoclypeal area unmodified. Protarsomere 1 lacking tenent setae. Mesofe-
mur (Fig. 24d) longer than metafemur. Mesotibia (Fig. 24c) about as long as metatibia.
Metatarsomere 1 about as long as combined length of metatarsomeres 2-4. Peg-like setae
absent from protibia, mesotibia and metatrochanter, arranged in a single row on meta-
tibia and mesotrochanter (Fig. 24d). Apex of abdominal tergite 8 as in Fig. 24e, g. Ster-
nite 8 as in Fig. 24f. Sternite 10 without basal protuberance. Aedeagus as in Fig. 24a, b.
2. Abdominal tergite 8 (Fig. 25f, 1) without medioapical projection. Sternite 8
as ın Fig. 25e. Genital segment as in Fig. 24a, b.
COMMENTS. See discussion under M. aino and M. corticalis.
Megarthrus shibatai Sawada (Figs 26a-m, 26a-h)
Megarthrus shibatai SAWADA, 1962: 13.
TYPE MATERIAL. Holotype d : Japan, Honshu, Nagano pref., Inago-yu, 1500-2000 m,
16.vii.1959 (T. Shibata) KSPC. Paratype (19): same data as holotype, but 24.vii.1959, KSPC.
Additional material (18). Japan, Honshu, Iwate pref., Kawai, Yoshibezawa, 1050 m,
12.viii.1991 (A. Smetana) #J49, MHNG.
DISTRIBUTION. Japan: Honshu (Chubu, and Tohuku distr.).
DESCRIPTION. Similar to M. japonicus from which it differs as follows: Dorsal
pubescence fairly uniform. Submentum strongly convex. Antenna as in Fig. 27c, d.
Pronotum as in Fig. 27e. Apical margin of elytron weakly convex near suture; inner
apical angle obtuse. Abdominal sternites 2 and 3 with median processes as in Fig.
26m. Ratios: AL 1.6-1.7; EL 1.5-1.6; ET 1.6-1.7; ML 1.5; PT 1.8; TPF 6.0-7.0.
3. Frontoclypeal area unmodified. Frontal pubescence parallel. Protarsomere 1
with tenent setae. Metafemur (Fig. 26f) somewhat longer than mesofemur (Fig. 26h).
Metatibia (Fig. 26b) longer than mesotibia (Fig. 26a). Metatarsomere | about as long
as combined length of metatarsomeres 2-4. Peg-like setae absent from protibia and
metatrochanter (Fig. 26f), arranged in a single row on mesotrochanter (Fig. 26h),
grouped to from a field on mesotibia, arranged in a single row on metatibia, but near
metatibial apex grouped to form a field. Apex of abdominal tergite 8 as in Fig. 26g, 1.
Sternite 8 as in Fig. 26k. Sternite 10 (Fig. 26d) with small subbasal protuberance.
Aedeagus as in Fig. 26c, d.
2. Frontal pubescence converging. Abdominal tergite 8 (Fig. 27f, g) with
medioapical projection. Sternite 8 as in Fig. 27g. Genital segment as in Fig. 27a, b.
COMMENTS. See discussion under M. japonicus.
MEGARTHRUS OF JAPAN 495
DISCUSSION
All available Japanese material of Megarthrus is from the main islands, totalling
16 species (Fig. 28, table 1). Most of the Japanese Megarthrus are from Honshu, Chubu
district (10 species). Altitudinal records indicate a preference of Megarthrus for mon-
tane biotopes ranging from 700-1600 m a. s. 1. Only 5 species were found between 200-
700 m, and 3 above 1600 m. The surprisingly high number of species from Shikoku
compared to that from Kyushu may be explained by gaps in the detailed knowledge of
the distribution. The related Omaliinae are represented in both islands by an equal
number of species (WATANABE 1990). In unrelated groups, such as Calliphoridae and
Sarcophagidae (KANO ef al., 1967; KANO & SHINONAGA 1968) these figures are
essentially the same. The relatively high proportion of species that also occur outside
Japan (37.5%) is notable. In the Omaliini which usually inhabit similar habitats (Löbl,
pers. comm.) only one of 17 Japanese species is also found in other countries (WATA-
NABE 1990). All of the non-endemic Japanese species of Megarthrus occur, outside
Japan, in Russia, temperate China and Europe. This suggests that Megarthrus repre-
sents a Palaearctic rather than an Oriental element in Japan.
ACKNOWLEDGMENTS
The present paper is a part of a joint project between the Muséum d’histoire
naturelle, Geneva and the Natural History Museum, London, funded by the Swiss
National Science Foundation (Project no 31-32331.91, awarded to I. Löbl, D. H.
Burckhardt, MHNG, and P. M. Hammond, BMNH). Their help is gratefully ack-
nowledged. The following colleagues have generously lent specimens: M. Jäch,
NHMW; S. E. Miller, BPBM; A. F. Newton Jr., FMNH; K. Sawada, Nishinomiya; A.
Smetana, CNCI; M. Uhlig, ZMHB, and L. Zerche, DEI.
TABLE |. Geographical distribution of the Japanese Megarthrus.
En | te
Tohuku
X
za |
|
M. incubifer
M. japonicus
[Tohuku |
ll
Lars]
BR
DE
Be
Ben
ses
ES
esi
BER
PUT EL ELT EPI DT
RELL
mL LL EE] EL EEL EE
e [UFFI
ELLE:
PULL HE ERE
PLETE TRE
ee.
496 GIULIO CUCCODORO
SY
a b c
~ 7
> 7 N Fa
2? x >
N: 7
d e f
/ VÀ
SS - IR 7 SE
SA
© DS DAN
g h / \ I 7. N
x SERA / \ /
N DA I |
I i [ I
\ 1 \ !
\ 1 \ l
\ \
Fic. 1
Scutellum: a-g; Temple and occipital ridge: h-n; schematic. Megarthrus aino: g; M. conformis:
a, 1; M. impressicollis: e, m; M. incubifer: f, k; M. japonicus: c, h; M. montanus: d, n; M.
parallelus: b.
MEGARTHRUS OF JAPAN 497
Fic. 2
Megarthrus aino; a, b: aedeagus, ventral (apical portion) and lateral; c: antenna; d, e: male,
apex of abdominal tergite 8, lateral and dorsal; f: male, abdominal sternite 8; g: median
processes of abdominal sternites 2-4 (left to right), schematic. Scale bars = 0.2 mm.
498 GIULIO CUCCODORO
Fic. 3
Megarthrus aino; a, e: female, genital segment, sternites (a) dorsal and tergites (e) ventral; b:
female, abdominal sternite 8; c, d: female, apex of abdominal tergite 8, dorsal and lateral; f:
pronotum. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 499
o o
= ° + oo o
°
eo Ò o ° 5 oo
9° 9° 0°
° 0 TMD Aa < al Be
DANSE i
sp KA
Le = BIS
4 i,
AN, 41 )
AK RQ 7 \
i wwe
ANS
c-f
a, b, g
Fic. 4
Megarthrus conformis, male; a, g: aedeagus, lateral and ventral (apical portion); b: abdominal
sternite 8: c: metafemur and metatrochanter; d: mesofemur and mesotrochanter; e: metatibia; f:
mesotibia. Scale bars = 0.2 mm.
500 GIULIO CUCCODORO
FIG. 5
Megarthrus conformis; a, g: female, genital segment, sternites (a) dorsal and tergites (g)
ventral; b: pronotum; c, f: female, apex of abdominal tergite 8, dorsal and lateral; d: antenna; e:
female, abdominal sternite 8; h: male, apex of abdominal tergite 8, dorsal; i: median processes
of abdominal sternites 2-4 (left to right), schematic. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 501
TGS
J, SX FAN 1
CHENE
1 RNA
are PTE
FIG. 6
Megarthrus conspirator; a, b: male, aedeagus, ventral and lateral; c: antenna; d: male,
metatibia; e: male, mesotibia; f: male, metafemur and metatrochanter; g: male, mesofemur and
mesotrochanter. Scale bars = 0.2 mm.
502 GIULIO CUCCODORO
g,! b,d
C,e, ne meta ere La a,f
FIG. 7
Megarthrus conspirator; a, f: female, genital segment, sternites (a) dorsal and tergites (f)
ventral; b, h: abdominal sternite 8 (b) male and (h) female; c, e: female, apex of abdominal
tergite 8, dorsal and lateral; d: male, apex of abdominal tergite 8, dorsal; g: pronotum, contour,
basal and lateral; i: median processes of abdominal sternites 2-3 (left to right), schematic. Scale
bars = 0.2 mm.
MEGARTHRUS OF JAPAN 503
FIG. 8
Megarthrus constrictus, male; a, c: aedeagus, lateral and ventral (apical portion); b: apex of
abdominal tergite 8, dorsal; d: abdominal sternite 8; e: mesotibia; f: metatibia; g: mesofemur
and mesotrochanter; h: metafemur and metatrochanter. Scale bars = 0.2 mm.
504 GIULIO CUCCODORO
Fic. 9
Megarthrus constrictus, a, b: female, genital segment, sternites (a) dorsal and tergites (b)
ventral; c: antenna; d: female, abdominal sternite 8; e, f: female, apex of abdominal tergite 8,
dorsal and lateral; g: pronotum; h: median processes of abdominal sternites 2-3 (left to right),
schematic. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 505
DAT USAI
DS A SA
ST
IN
N
=
7
Fic. 10
Megarthrus convexus; a, b: male, aedeagus, lateral and ventral (apical portion); c: male,
mesotibia; d: male, apex of abdominal tergite 8, dorsal; e: median processes of abdominal
sternites 2-3 (left to right), schematic; f: male, metatibia; g: male, mesofemur and meso-
trochanter; h: male, metafemur and metatrochanter. Scale bars = 0.2 mm.
506 GIULIO CUCCODORO
FIG. 11
Megarthrus convexus; a, c: abdominal sternite 8 (a) male and (c) female; b, f: female, apex of
abdominal tergite 8, lateral and dorsal; d, h: female, genital segment, sternites (d) dorsal and
tergites (h) ventral; g: pronotum. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 507
Fic. 12
Megarthrus corticalis, male; a, b: aedeagus, lateral and ventral (apical portion); c, i: apex of
abdominal tergite 8, lateral and dorsal; d: mesotibia; e: metatibia; f: mesofemur and meso-
trochanter; g: metafemur and metatrochanter; h: abdominal sternite 8. Scale bars = 0.2 mm.
508 GIULIO CUCCODORO
Fic. 13
Megarthrus corticalis; a, b: female, genital segment, sternites (a) dorsal and tergites (b) ventral;
c: antenna; d: right elytron; e, f; female, apex of abdominal tergite 8, lateral and dorsal; g:
median processes of abdominal sternites 2-4 (left to right), schematic; h: female, abdominal
sternite 8; 1: pronotum. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 509
Fic. 14
Megarthrus impressicollis; a; antenna; b-d, g: female, genital segment, sternites (b, c) dorsal
and lateral, and tergites (d, g) lateral and ventral; e, f: female, apex of abdominal tergite 8,
lateral and dorsal; h: median processes of abdominal sternites 2-4 (left to right), schematic; 1:
pronotum; k: female, abdominal sternite 8. Scale bars = 0.2 mm.
GIULIO CUCCODORO
510
SX
SS
/
2 Ha a ai |
| AA Ny
| Yl \
c,d,g
a,b,f,h,i
Fic. 15
Megarthrus incubifer; a, b: male, aedeagus, ventral (apical portion) and lateral; c: male
mesotibia; d: male, metatibia; e: median processes of abdominal sternites 2-4 (left to right),
schematic; f, i: male, apex of abdominal tergite 8, dorsal and lateral; g: male, mesofemur and
mesotrochanter; h: male, abdominal sternite 8. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 511
Fic. 16
Megarthrus incubifer; a: antenna; b: female, abdominal sternite 8; c, d: female, apex of
abdominal tergite 8, lateral and dorsal; e: pronotum; f, g: female, genital segment, tergites (f)
ventral and sternites (g) dorsal. Scale bars = 0.2 mm.
512 GIULIO CUCCODORO
Fic. 17
Megarthrus japonicus; a, b: male, aedeagus, lateral and ventral (apical portion); c: male,
mesotibia; d: male, metatibia; e, f: male, apex of abdominal tergite 8, dorsal and lateral; g:
median processes of abdominal sternites 2-4 (left to right), schematic; h: male, mesofemur and
mesotrochanter; i: male, abdominal sternite 10; k: male, metafemur and metatrochanter. Scale
bars = 0.2 mm.
MEGARTHRUS OF JAPAN 513
Fıc. 18
Megarthrus japonicus; a, b: female, genital segment, sternites (a) dorsal and tergites (b) ventral;
c: antenna; d, e: female, apex of abdominal tergite 8, lateral and dorsal; f, g: abdominal sternite
8 (f) female and (g) male; h: pronotum. Scale bars = 0.2 mm.
514 GIULIO CUCCODORO
Fic. 19
Megarthrus montanus, male; a, f: aedeagus, lateral and ventral (apical portion); b: mesofemur
and mesotrochanter; c: metatrochanter; d: mesotibia; e: abdominal sternite 8; g: abdominal
sternite 10; h, i: apex of abdominal tergite 8, dorsal and lateral. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 515
= c
h ————— a,b,d-f
Fic. 20
Megarthrus montanus; a, b: female, genital segment, sternites (a) dorsal and tergites (b) ventral;
c: antenna; d: female, abdominal sternite 8; e, f: female, apex of abdominal tergite 8, lateral and
dorsal; g: pronotum; h: median processes of abdominal sternites 2-4 (left to right), schematic.
Scale bars = 0.2 mm.
GIULIO CUCCODORO
516
>
UPL
ef
a
Uf
NS
NIS
Fic. 21
Megarthrus parallelus; a, b: male, aedeagus, ventral (apical portion) and lateral; c: male,
metatibia and metatarsomeres; d: male, mesotibia; e: median processes of abdominal sternites
2-4 (left to right), schematic; f, i: male, apex of abdominal tergite 8, dorsal and lateral; g: male,
abdominal sternite 10; h: mesofemur and mesotrochanter; k: metafemur and metatrochanter.
Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN Sl],
FIG. 22
Megarthrus parallelus; a, b: female, genital segment, sternites (a) dorsal and tergites (b)
ventral; c: antenna; d: pronotum; e, g: abdominal sternite 8 (e) female and (g) male; f, h:
female, apex of abdominal tergite 8, dorsal and lateral. Scale bars = 0.2 mm.
518 GIULIO CUCCODORO
FIG. 23
Megarthrus sawadai; a, d: male, aedeagus, ventral (apical portion) and lateral; b: male, meso-
tibia; c: male, metatibia; e: male, abdominal sternite 8; f: antenna; g: pronotum; h, m: male,
apex of abdominal tergite 8, dorsal and lateral; i: male, abdominal sternite 10; k: mesofemur
and mesotrochanter; n: median processes of abdominal sternites 2-4 (left to right), schematic.
Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 519
SI SÈ
FIG. 24
Megarthrus scriptus, male; a, b: aedeagus, ventral (apical portion) and lateral; c: mesotibia; d:
mesofemur and mesotrochanter; e, g: apex of abdominal tergite 8, dorsal and lateral; f:
abdominal sternite 8. Scale bars = 0.2 mm.
520 GIULIO CUCCODORO
Fic. 25
Megarthrus scriptus; a, b: female, genital segment, sternites (a) dorsal and tergites (b) ventral;
c: right elytron; d: antenna; e: female, abdominal sternite 8; f, i: female, apex of abdominal
tergite 8, dorsal and lateral; g: pronotum; h: median processes of abdominal sternites 2-4 (left to
right), schematic. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN 521
Fic. 26
Megarthrus shibatai; a: male, mesotibia; b: male, metatibia; c, e: male, aedeagus, ventral
(apical portion) and lateral; d: male, abdominal sternite 10; f: male, metafemur and metatro-
chanter; g, i: male, apex of abdominal tergite 8, dorsal and lateral; h: male, mesofemur and
mesotrochanter; k: male, abdominal sternite 8; m: median processes of abdominal sternites 2-4
(left to right), schematic. Scale bars = 0.2 mm.
522 GIULIO CUCCODORO
Fic. 27
Megarthrus shibatai; a, b: female, genital segment, sternites (a) dorsal and tergites (b) ventral;
c: antennomeres 5-9; d: antenna; e: pronotum; f, h: female, apex of abdominal tergite 8, dorsal
and lateral; g: female, abdominal sternite 8. Scale bars = 0.2 mm.
MEGARTHRUS OF JAPAN
Un
N
Qo
1 (0) Hokkaido
2 (0)
1 (0)
Shikoku
FIG. 28
Numerical relationships of 16 species of Megarthrus between the main Japanese islands and the
continent (bold numbers = total number of species in main islands and total number of species
shared between them and the continent; number in parenthesis = number of species unique to a
single island or uniquely shared).
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(Reuter, 1876), Deuterosminthurus bicinctus (Koch, 1840), D. flavus
(Gisin, 1946), D. pallipes (Bourlet, 1843), and D. sulphureus (Koch, 1840).
Two new species are described: Deuterosminthurus maculatus sp. n. and
Deuterosminthurus pleuracanthus sp. n. A discussion is given on the
morphology of the claw, which is used to distinguish genera in the family
Bourletiellidae. Appendicular chaetotaxic characters are relevant for
defining genera, but more difficulty applicable for species distinctions. On
the basis of such characters, apomorphies of Heterosminthurus and
Deuterosminthurus are emphasized. Setae of body and color pattern are
also used for species diagnoses, and a key to European species of
Deuterosminthurus is provided.
Key-words: Collembola - Symphypleona - Bourletiellidae - Heterosmin-
thurus - Deuterosminthurus - taxonomy - Europe.
INTRODUCTION
Ontogeny of the appendicular chaetotaxy in species of the genera Heteros-
minthurus and Deuterosminthurus is described according to a standard I proposed
(NAYROLLES 1993a, 1993b) and since then have used to study the family Bour-
letiellidae (NAYROLLES 1994, 1995). Chaetotaxy of the body is less known, excepted
for the trichobothrial pattern (RICHARDS 1968; BETSCH 1980; BETSCH & WALLER
1989) and the small abdomen. I recently discussed the setal nomenclature of the small
Manuscript accepted 29.08.1995.
526 PIERRE NAYROLLES
abdomen (NAYROLLES 1995). I deemed that the paper of BETSCH & WALLER (1994) is
fundamental for any study to come. My view relied on two arguments: these authors
observed all ontogenetic levels, and used a wide taxonomic range. Nevertheless, they
did not propose a coherent nomenclature, probably because they considered further
observations to be necessary. For a practical reason, I have adopted BRETFELD' s
(1990) nomenclature of the small abdomen, which is sufficient for the adult
chaetotaxy. In the present paper, only the adult female circumanal setae are described.
The female of Heterosminthurus and Deuterosminthurus has seven circumanal setae
on the upper anal flap (the impaired a0 and the paired al, a2, and a3), and five setae
on the lower anal flap (from dorsal to ventral side: avl', avl, av2, av3, av4, and av5 =
anal appendage). Two species redescribed here, Heterosminthurus insignis and
Deuterosminthurus bicinctus, have already been studied by BRETFELD (1990). I
criticized this work (NAYROLLES 19935), and explained the reasons for dismissing
BRETFELD' s nomenclature of appendicular setae.
I give a definition of each genus with emphasis on the most obvious apo-
morphies. Other characters will be discussed in a future paper. Nevertheless, an
account of the empodial morphology is here needed.
According to BETSCH (1980), the empodial appendage of Bourletiellidae
displays an array of regressive evolutions, so that the definition of genera is mainly
based on its morphology. Using a scanning electron microscope, BETSCH & MASSOUD
(1970) provided an accurate survey of this organ in Jeannenotia stachi Stach, 1956
(Sminthurididae). In the most typical form, the empodial appendage consists of a
central rib, three lamellae, a subapical filament, and an apical needle. In comparison
with the leg orientation, a lamella is external (thus opposite the claw), another is
internal and slightly posterior, the last is anterior with a very slightly internal position.
The central rib and a large area of lamellae are covered with granular cuticle. The
ridge of lamellae is smooth, excepted for the basal part of the outer lamella. Smooth
cuticle is also present on the tip of the empodial appendage, at the apex of the fila-
ment and needle. RICHARDS (1968) described the subapical filament as a projection of
the central rib. BETSCH (1980) refuted Richards’ view, showing that the subapical
filament fitted with the outer lamella extension. More precisely, I consider that the
ridge of the outer lamella extends beyond the central rib in the subapical filament, and
the ridges of the other two lamellae join together into the apical needle.
European genera of Bourletiellidae can be readily defined by the shape of the
empodial appendage. Fasciosminthurus Gisin, 1960 sensu Bretfeld, 1992 and the
related genus Cyprania Bretfeld, 1992 — which perhaps should not have been
isolated (NAYROLLES 1993b) — show an empodial appendage made up of a granular
central rib ending in the filament and needle and devoid of lamellae. Filament and
needle are thus the only traces of lamellae in these genera. In Bourletiella and Cas-
sagnaudiella, the lamellae, filament, and apical needle are reduced but distinct.
Lamellae in Deuterosminthurus and Heterosminthurus are present, and they extend in
a single apical spike. I was not able to state whether this apical point corresponds to
the filament, the apical needle, or both welded together. BETSCH' s (1974) description
of the pretarsus of Bourletiellitas imerinensis Betsch, 1974 and Vatomadiella pauliani
EUROPEAN BOURLETIELLIDAE 527
Betsch, 1974 shows that the apical spike likely amounts to the filament and apical
needle welded together. It may be the same in Deuterosminthurus and Hetero-
sminthurus. The use of scanning electron microscope would possibly permit us to
resolve this problem.
All the species were collected in southwestern France and northeastern Spain.
Abbreviations and setal symbol follow NAYROLLES (1993a, 19935).
The material is deposited in the following institutions: Laboratoire d'Ecologie
des Invertébrés Terrestres, Université Paul Sabatier, Toulouse, France (LEITT) —
Muséum d'histoire naturelle de Genève, Switzerland (MHNG) — Museum National
d'Histoire Naturelle, Paris, France (MNHN) — Institute of Systematics and Popu-
lation Biology (Zoologisch Museum) Universiteit van Amsterdam, the Netherlands
(ZMA).
GENUS Heterosminthurus Stach, 1955
BRETFELD (1990) has noted the absence of Jp on all tibiotarsi of H. insignis.
However, Ili is always more basal than Ja and Jp, and, in comparison with Ja, the
apical remaining seta on the inner side of H. insignis is approximately at the same
level regarding the longitudinal axis of legs. Therefore, I consider that [lt is absent
and the remaining seta is Jp.
Heterosminthurus shows a characteristic pretarsus. The claw is devoid of
tunica, pseudonychia and distinct teeth on the inner crest. The empodial lamellae of
forelegs are reduced as well as the central rib which extends in a long apical point as a
bent outward blade. On the mid and hindlegs, the central rib and lamellae are very
distinct, and the apical point is shorter than on forelegs and bent toward inner side.
The posterior pretarsal seta is absent as in all Bourletiellidae.
Heterosminthurus is characterized by the following apomorphies: empodium
of forelegs different from the empodium of mid and hindlegs, (TI.)Hi absent, and
setae (AT)a+2 and (AT)ai+2 present in adults.
Three species are studied: H. insignis, H. bilineatus, and H. claviger.
Heterosminthurus insignis (Reuter, 1876) (Figs 1-8; Table I)
MATERIAL EXAMINED - FRANCE, dép. Lozère, district Nasbinals, Souvérols Lake, alt.
1230 m; peat bog, edge of lake on the floating vegetation, with Menyanthes trifolia, Sphagnum
sp., Drosera rotundifolia, Comarum palustre, Ligularia sibirica, etc.; collected with a net on
Sphagnum sp. and D. rotundifolia (LZ1d). 21-VI-1986: many specimens; 5 juv. mounted. 11-
IX-1988: many specimens; 7 juv. and 7 ad. mounted (LEITT). — Dép. Gironde, district Car-
cans, between Carcans and Maubuisson, Hourtin-Carcans Lake, alt. 0 m; near the lake, with
Phragmites communis and Hydrocotyle vulgaris; collected with a net on Phragmites communis
(GI4). 23-X-1987: 3 juv., 2 mounted (LEITT). — Dep. Pyrénées-Atlantiques, district Lecum-
berry, between Estérençuby and Larrau, near Astakieta wood, alt. 1150 m; small peat bog with
Sphagnum sp., Eriophorum sp., Vaccinium myrtillus and Erica vagans; collected with a net on
unselected plants (PA8a). 10-V-1987: 21 specimens; 1 juv. and 4 ad. mounted (LEITT).
528 PIERRE NAYROLLES
Table I. Appendicular chaetotaxy of Heterosminthurus insignis
D:i0 T:i+1,a+1,e+1 Q:i-1, pe+1
Ai is a trichobothrium.
Other setae: Q: ae+2, a+2, ai+2,
pe+2, pe+3
Setae -1
Setae 0
Setae +1
B completely zoned. Section M euwhorled with 5 whorls. Right euwhorlation.
Subsegmentation formula: 1 + 6 + 1 = (A) + (M1, 5 + BA) + (BM+BB). Right subsegmentation.
Apical bulb: joined to the apex Subapical organ: small
Allpe is a blunt microchaeta Alle is shifted toward apex
Hppe 1-1-1-1-1/0.6. m(h”) =5.6.
Setae of BB: P: BBe, BBae, BBa, BBp, BBpe / all G setae on M are present.
In female: BA complete whorl; n80%(BM) = 15-23; n80%(B) = 28-36; n80%(G) = 93-101.
In male: BA complete whorl; n80%(BM) = 23-30; n80%(B) = 36-43; n80%(G) = 101-108.
BBae and BBa are cucumiform setae on large base.
STE) (ES) aa Er N
(BAG NET Bun Pistes Et n [nit un Pi setae Luuii sia
itt Bra er i ee]
RA A ni
TR Brea]
P: i1
Te |
T:a2,011,0i2 Q:ae T: a2, Oil, Oi2 Q: ae
ren |
delayed primordial seta: Q: ae3 T: ai2, pe2, Op
T: pe2, Op Q:ai2, pe4 (T)Q: a5 (oc=0.4) Q:pe4
©
®
—
T: a5, ai2, pe2
Q: ai3, ai4, pe4, Oi
Ili, IVp, Vp Ii, Hp, IVp, Vp
lE Pee)
T: 4ai1, Vai, Vpi, FSa, O2pe T: 4ai1, Vai, Vpi, FSa, O2pe T: 3ai, 3i, 4ai1, 411, Vai, Vpi,
Q: 3a, 3p, dat, 4pi1, 4p1, 4ai2, | Q: 3a, 3p, dat, 4pit, 4p1, 4ai2, FSa, O2pe
FSai, FSpi FSai, FSpi Q: 2a, 3a, Spi, 4a1, 4pil, 4ai2,
Ipi, Ip and Ipe are spatulate | Ipi, Ip and Ipe are spatulate FSalzisopi
setae. setae. Ip and lpe are spatulate setae.
Ja is a winged seta.
CEE E
1... [PES RE SEE
Gelcae]lcaleannenleri ) Other setae: P:2pe
-
| B | setae on Gpi from Ipi to Vpi are
p
BEI) | macrochaetae
|
|
Seas:
m >
Chaetotaxy: — Morphology: anterior lamella simple, outer and inner lamellae smooth,
fr
(=
mucro ending in a small point.
EUROPEAN BOURLETIELLIDAE 529
DESCRIPTION
Development. — Number of juv. st. = 3.
Size ad. — d: 0.7 - 0.85 mm; 9: 0.75 - 1.5 mm (female specimens collected
in mountain do not reach beyond 1.2 mm).
Color. — Background yellow, often with ochre shades. Dark pigment
sometimes present, particularly in the posterior area of great abd., the color being
brown or wine-red in some specimens. À very dark male was collected, it was wholly
black except between eyes, with pigment on legs and furcula. Antennae brown,
eyepatches black. The different color-forms are similar to those described by
BRETFELD (1988).
Great abd. — Back strongly concave. Trichobothria: A, B, and C in linear
pattern. Dorsal setae: mesochaetae.
Antennae (figs 6 & 7). — Seta Alle of ant. IV shifted toward apex. Antennal
ratios: in 2, ant. I : Il : WI: IV = 1 : 2.3 : 3.5 : 6.4 and ant. : ceph. diag. = 1.9; in d, ant.
1:0: U1:IV=1:2.8: 4.0: 7.6 and ant. : ceph. diag. = 2.5. Chaetotaxic variables, in
both sexes: m(/Sh*) = 0.94 ; in 2 : for G: m = 97.6 / min = 91 / max = 102 ; m(RSh) =
0.55 ; in 6: for G: m = 104.6 / min = 100 / max = 108 ; m(RSh) = 0.51.
Head. — Eyes: 8+8; eyepatch with two setae. 2+2 ov. org. in the back-ventral
area (see BRETFELD, 1990). Labral formula: 6/5-5-4. Cephalic setae: only mesochaetae
in 2; d with 2+2 macrochaetae in inner edge of eyepatches. Several frontal setae
small, but not broadened at the basis as in H. bilineatus and H. claviger.
Legs (figs 1-5). — Femur: cup present. Pretarsus characteristic of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula (fig. 8). — Dental setae of Gpi, from Ipi to Vpi, are macrochaetae.
Small abd. — Made up of abd. V + abd. VI. Trichobothria: D and E. Male
genital papilla with 16 setae. Size comparison of female circumanal setae: avl' = avl
< av3 < av2 = av4. An. app. spine-like, slightly bent. Ratio an. app. : mucro = 0.59.
Heterosminthurus bilineatus (Bourlet, 1842) (Figs 9-15; Table II)
MATERIAL EXAMINED — FRANCE, dép. Aveyron, district Bessuéjouls, Le Plateau de la
Justice, alt. 450 m; meadow belonging to the phytosociological association of Orchido-
Brometum with some small trees (Quercus pubescens, Pinus sylvestris, Juniperus communis)
and Bromus erectus, Hippocrepis comosa, Globularia vulgaris, Eryngium campestre, etc.;
collected with a net on unselected plants (AVIa). Several collections from March until
December 1987. In all 62 specimens; 4 juv., 3 2, and 4 ¢ mounted (LEITT). — Dép. Aveyron,
district La-Bastide-Solages, D33 road between Plaisance and Brasc, alt. 350 m; grass alongside
a chestnut grove; collected with a net (AV15b). 22-VIII-1987: 77 specimens; 2 juv. and 2 d
mounted (LEITT). — Dép. Lozère, district Nasbinals, Souvérols Lake, alt. 1230 m; peaty soil
near the lake, with Eriophorum polystachyon, Equisetum limosum, Orchis incarnata; collected
with a net on unselected plants (LZ1c). 21-VI-1986: 114 specimens; 2 d and 2 9 mounted. 19-
VII-1986: 8 specimens. 11-IX-1988: 194 specimens; 5 juv. and 1 2 mounted (LEITT). — Dep.
530 PIERRE NAYROLLES
Table Il. Appendicular chaetotaxy of Heterosminthurus bilineatus
Ai is a trichobothrium. In adult male: Ae, Aae, Aa, and e+1 are acanthoids; Be, Bae, Ba, Bai, and a+1
are small and slender.
Other setae: Q: ae+2, a+2, ai+2,
pe+2, pe+3
Setae 0 | | | | Only in adult male: Bae is an
Setae +1 | | | | | acanthoid; Ae, Aa, Ai, Ap, Be, Ba, Bai,
Bi, and Bp are small and slender; p-1 is shifted toward i inner side.
B completely zoned. Section M euwhorled with 5 whorls. Right euwhorlation.
Subsegmentation formula: 1 + 6 + 1 = (A) + (M1, 5 + BA) + (BM+BB). Right subsegmentation.
Apical bulb: joined to the apex Subapical organ: small
Allpe is a blunt microchaeta Alle is shifted toward apex
Setae zi BB PE BBe, BBae, BBa, BBp, BBpe / all G setae on M are present.
In female: BA complete whorl; n80%(BM) = 16-22; n80%(B) = 29-35; n80%(G) = 94-100.
In male: BA complete whorl; n80%(BM) = 29-36; n80%(B) = 42-49; n80%(G) = 107-114.
BBae and BBa are cucumiform setae on large base.
SO PSN aston RS | ea ein | ee N
AG ea PSS [ee]
i‘ D_ziiPe]ee WE EF
TR[I]
FE Ir De 5
delayed primordial seta: Q: ae3 T: ai2, pe2, Op T: a5, ai2, pe2
T:pe2, Op Q:ai2, pe4 (T)Q: a5 (oc = 0.6) Q: ai3, aid, pe4, Oi
RE SE A ES Ili, lp, Vp, Vp
T: 4ai1, Vai, Vpi, FSa, O2pe
Q: 3a, 3p, 4a1, 4pit, 4p1, 4ai2,
FSai, FSpi
lpi, Ip and Ipe are spatulate
setae.
Ja is a winged seta.
Jp is a very slender micro-
chaeta.
T: 4ai1, Vai, Vpi, FSa, O2pe : 3ai, 3i, 4ai1, 411, Vai, Vpi,
Q: 3a, 3p, 4a1, 4pi1, 4ai2, FSai, FSa, O2pe
FSpi Q: 2a, 3a, Spi, 4a1, 4pil, 4ai2,
(Q): 4p1 (oc = 0.7) FSai, FSpi
lpi, Ip and Ipe are spatulate | Ip and Ipe are spatulate setae.
setae.
Jp is a very slender micro-
chaeta.
As ne Me N
nio en uu «n spe peste WO
Ge [Gae| Ga | Gai | Gi | Other setae: P:2pe
P P P P
P
p
Whorls | P
| to VI P
WhorlB
Chaetotaxy: — Morphology: anterior lamella simple, outer and inner lamellae smooth,
mucro ending in a small point.
EUROPEAN BOURLETIELLIDAE 53]
Tarn, district Murat-sur-Vèbre, D162 road, Laouzas Lake, alt. 800 m; lakeside with Baldingera
arundinacea, Juncus silvaticus, Molinia cerulea; collected with a net on unselected plants
(TN3). 22-VIII-1987: 62 specimens; 3 5 mounted (LEITT). — Dép. Tarn, district Murat-sur-
Vebre, D162 road between Laouzas Lake and Murat-sur-Vèbre, about 3.5 km from Murat-sur-
Vèbre, alt. 820 m; near a small stream in a damp meadow, with Juncus sp., Glyceria sp., and
Molinia cerulea; collected with a net on unselected plants (TN4a). 22-VIII-1987: 77
specimens; 2 juv. mounted (LEITT).
DESCRIPTION
Development. — Number of juv. st. = 3.
Size ad. — d: 0.5 - 0.55 mm; 9: 0.6 - 0.9 mm.
Color. — Background light yellow. Back of great abd. with a pair of
longitudinal brown or orange stripes, generally quite diffuse, and joined behind.
Several small patches on the posterior flanks of great abd. forming a pair of short
stripes which do not reach halfway the body. A pair of posterior and lateral small
elongated flecks is sometimes present. Trichobothrial sockets brown. Back of small
abd. marbled with variable brown pigment. Head light with two small flecks behind
eyes in the same line of great abd. streaks. Eyepatches black, antennae brown, legs
very light yellow, and furcula white.
Great abd. — Back weakly concave. Trichobothria: A, B, and C in linear
pattern. Dorsal setae: mesochaetae.
Antennae (figs 10-15). — Seta Alle of ant. IV shifted toward apex. & with
several special-shaped setae on ant. IT and II, and (AT)p-1 shifted toward inner side, so
that it might almost be thought of as included in the generatrix Gpi. Antennal ratios: in
AE VE 125-2172 and ant..ceph. das —2 0; nse ante 1
:IV=1:2.6:4.5:7.5 and ant. : ceph. diag. = 2.6. Chaetotaxic variables, in both sexes:
mast) = 092 ne tonG: m = 9123. /-mint=90:/max-=10le;m(RSAh) = 0.53 20€ :
on Gen 1103) min= 103 / max = 117: m(RSh)=0.47.
Head. — Eyes: 8+8; eyepatch with two setae. 1+1 ov. org. in the back-ventral
area (it remains the ventral — anterior in arthropleon orientation — pair). Labral
formula: 6/5-5-4. Cephalic setae: only mesochaetae in 9 ; d with several transformed
setae. In dorsal part of the frons, near the sagittal plane, 2+2 acanthoids, the more
posterior pair being the longer. Inner edge of eyes with 1+1 macrochaetae; frons with
several small onion-shaped setae.
Legs (fig. 9). — Femur: cup present. (TI1,2)Jp is a very slender microchaeta.
Pretarsus characteristic of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula. — See chaetotaxic table.
Small abd. — Made up of abd. V + abd. VI. Trichobothria: D and E. Male
genital papilla with 12 setae. Female circumanal seta avl' far shorter than the others,
with the following size comparison: avl' < avl = av4 < av2 = av3. An. app. spine-
like, slightly bent. Ratio an. app. : mucro = 0.76.
53) PIERRE NAYROLLES
SO pm EE et
50 ym 6
Fics 1-7
Heterosminthurus insignis ad.; 1: foretibiotarsus, anterior view; 2: schematic representation of
fig.1, setae of Ge and Gi, and ov. org. are drawn, other setae are schematized as follows: a full
symbol for a seta on reader’s side (thus on anterior side), an empty symbol for a seta on
opposite of reader’s side, a circle for a primary seta, a triangle for a secondary seta; setae
belonging to a same whorl are linked together by a line which is continuous on the anterior side
and discontinuous on the posterior side; 3: schematic representation of fig. 4 (hindtibiotarsus),
EUROPEAN BOURLETIELLIDAE 533
Heterosminthurus claviger (Gisin, 1958) (Fig. 16)
MATERIAL EXAMINED — FRANCE, dép. Lot-et-Garonne, district Durance, track of Le
Brocq farm, alt. 120 m; alongside a wood of Pinus pinaster, with Calluna vulgaris and Ulex
europæus; collected with a net on unselected plants (LG2). 10-X-1989: many specimens; 8 juv.
and 9 ad. mounted (LEITT). — Dép. Landes, district Brocas, Marais de la Coumme, alt. 80 m;
near a wood, with Molinia cerulea, Calluna vulgaris and Agrostis sp.; collected with a net on
unselected plants (LA6b). 22-X-1987: 38 specimens; 2 d mounted. 10-X-1989: 43 specimens
(LEITT).
DESCRIPTION
The Ist st. has not been observed, and the primary appendicular chaetotaxy 1s
an interpretation based on comparisons with other instars of this species as well as the
primary chaetotaxy of the other two studied species of Heterosminthurus.
Concerning the appendicular chaetotaxic table, refer to H. bilineatus (Table Il),
with the following differences:
Antennae. Sexual dimorphism: (AD)Be is an acanthoid in d, shape of other
setae similar as in bilineatus, the acanthoids being slightly thicker — Occurrences of
H setae on M & B: for Heae 1-1-1-1-1/0.8, and for Hppe 1-1-1-1-0.8/0.3. m(h*) = 5.5
— Chaetotaxic variables in 2 : n80%(BM) = 18-25 ; n80%(B) = 31-38 ; n80%(G) =
96-103. In d : n80%(BM) = 30-41 ; n80%(B) = 43-54 ; n80%(G) = 108-119.
Legs. (FE2)a5 : ontogeny type T — (TI2)4p1 : ontogeny type Q.
Development. — Number of juv. st. = 3.
Size ad. — d: 0.55 - 0.6 mm; 9: 0.65 - 0.9 mm.
Color. — Same pattern as in H. bilineatus with a darker pigment (dark brown).
Great abd. — Back weakly concave in posterior area. Trichobothria: A, B, and
C in linear pattern. Dorsal setae: mesochaetae.
Antennae (fig. 16). — Seta Alle of ant. IV shifted toward apex. 4 with several
special-shaped setae on ant. II and II, and (AT)p-1 shifted toward inner side, so that it
might almost be thought of as included in generatrix Gpi. Antennal ratios: in 2, ant. I:
HSIN, — alee D 3.7/1 6.O,and ant ceph. diag. 2:0 ini. ant AIME HE:
2.3 : 4.2 : 6.7 and ant. : ceph. diag. = 2.7. Chaetotaxic variables, in both sexes: m(U/Sh*)
= (Olin 2s fonG: m= 98.2 /min=94)/ max — 103 mRSh) 053 410 Ge tonGam
— #67 mim 310 max =120EmRSAr) = 0145:
Head. — Eyes: 8+8; eyepatch with two setae. 1+1 ov. org. in the back-ventral
area (it remains the ventral pair). Labral formula: 6/5-5-4. Cephalic setae: only
mesochaetae in 9 ; d with several transformed setae. In dorsal part of the frons, near
same legend as in fig. 2; 4: hindtibiotarsus, anterior view; 5: apex of midleg, detail in posterior
view, the arrow points the seta Jp, Ipi has its socket topped by an integumentary bulge; 6: ant.
III, anterior view; 7: schematic representation of fig. 6, setae of Ge and Gi are drawn as those of
antennal HI organ (Xe, Xi) and its guard setae (Aai, Api, Ape), other setae are schematized as
follows: a full symbol for a seta on reader’s side (thus on anterior side), an empty symbol for a
seta on opposite of reader’s side, a circle for a primary seta, a triangle for a secondary seta:
secondary setae belonging to a same generatrix are linked together by a line which is
continuous on the anterior side and discontinuous on the posterior side.
534 PIERRE NAYROLLES
10
_50pm _ 0.11.,14,15,16
SME
Fics 8-16
Fig. 8: Heterosminthurus insignis ad., furcula, posterior view; on the right, schematic
representation as follows: setae of Ge and Gi drawn, a full symbol for a seta on reader’s side
(thus on posterior side), an empty symbol for a seta on opposite of reader’s side, a circle for a
primary seta, a triangle for a secondary seta; for the dental whorls I to IV and B, setae
belonging to a same whorl are linked together by a line which is continuous on the posterior
side and discontinuous on the anterior side. Figs 9-15: Heterosminthurus bilineatus 3; 9: apex
EUROPEAN BOURLETIELLIDAE 535
the sagittal plane, 2+2 acanthoids, the more posterior pair being thick, very long, and
widened at apex as fan-shaped. Inner edge of eyes with several fairly long setae; frons
with several small onion-shaped setae.
Legs. — Femur: cup present. (TI1,2)Jp is a very slender microchaeta. Pretarsus
characteristic of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
Retinaculum. — Rami bidentate from 2nd st. on. Chaetotaxy: on anterior lobe
two setae appear at 3rd st. and one in ad.
Furcula. — Same chaetotaxy as in bilineatus.
Small abd. — Made up of abd. V + abd. VI. Trichobothria: D and E. Male
genital papilla with 12 setae. Size comparison of female circumanal setae: avl' < avi
< av? = av3 = av4. An. app. spine-like, slightly bent. Ratio an. app. : mucro = 0.71.
DISCUSSION
On the criterion of presence vs absence of setae, the appendicular chaetotaxy is
virtually the same among the three species. Indeed, few setae variable in a species and
constant in others are the only differences. Such characters, perhaps variable between
populations, cannot be used to distinguish species. Setal shape provides some relevant
characters, e.g., the dental macrochaetae in insignis, and (TI1,2)Jp as a minute
microchaeta in bilineatus and claviger. Morphology of cephalic setae makes the
distinction between males of claviger and bilineatus easy. Juveniles and females can be
identified by the lateral chaetotaxic pattern of great abd. described by BRETFELD (1986).
Genus Deuterosminthurus (Borner, 1901) sensu Gisin, 1960
Deuterosminthurus, as other genera of Bourletiellidae, displays a particular
pretarsal morphology. The claw is without tunica and pseudonychia, and its inner
crest bears a small and sometimes hardly distinguishable tooth. The empodium is
ending in a single apical spike. The reduction of empodial lamellae in all legs is an
apomorphy of Deuterosminthurus. The absence of the femoral setae (FE3)ai3 and
(FE3)ai4 in all the species of Deuterosminthurus I have observed also fits with a
of midleg, detail in posterior view, the arrow points the microchaeta Jp, the sockets of Jp and
Ipi are hidden by an integumentary fold; 10: ant. III, anterior view; 11: schematic
representation of fig. 10, same legend as in fig. 7, the arrow shows the displacement of the seta
p-1; 12: apex of ant. IV (with the section A and the whorl MI), anterior view; 13: schematic
representation of fig. 12, setae of Ge and Gi drawn, other setae are schematized as follows: a
full symbol for a seta on reader’s side (thus on anterior side), an empty symbol for a seta on
opposite reader’s side, a square for a H seta, a triangle for the secondary seta (Alai), and a
circle for a primary G seta except a cross for AA and a star for Allpe; setae belonging to a same
whorl are linked together by a line which is continuous on the anterior side and discontinuous
on the posterior side; 14: ant. II, anterior view; 15: schematic representation of fig. 14, setae of
Ge and Gi are drawn, other setae are schematized as follows: a full symbol for a seta on
reader’s side (thus on anterior side), an empty symbol for a seta on opposite of reader’s side, a
circle for a primary seta, a triangle for a secondary seta. Fig. 16: Heterosminthurus claviger à,
ant. II, anterior view.
536 PIERRE NAYROLLES
putative evolved character. The same assumption can be made for the absence of
(DE)llae and (DE)IVa.
From observations on an extensive material, I state that the character "strong
vs weak tooth on the inner lamella of claw", which has been frequently mentioned in
literature, is worthless. This point and chaetotaxy involve to consider that the species,
subspecies, or forms described as pallipes, repandus, mixtus, feneysi, and beckeri (see
ELLIS 1974) merely amount to only one species: D. pallipes. All these forms chiefly
corresponded to differences in color patterns, but color is a variable character and a
large range of tinges can be observed. ELLIS (1974) clustered these forms in two
species, repandus and pallipes, and in 1978 he questioned the separation of both. The
name pallipes (Bourlet, 1843), coined for a dark form, is prior to repandus (Agren,
1903). The valid name is thus pallipes; even though the form pallipes, in its original
description, relates to a rare color-form (males and females dark blue-purple excepted
for mouth, furcula, and legs) observed for some specimens collected in Atlantic
meadows, and the name repandus is more frequently cited in literature than pallipes.
The different color-forms of pallipes do not present a defined pattern. On the
other hand, D. maculatus Sp. n., a close species to pallipes, has a constant color pattern
which consists of sharply outlined maculae always laid down in the same arrangement.
In some places, both species occurred together, and never it was found a specimen with
an intermediate color; such a fact involved to give each form a specific status. Hence, I
tried in vain to find a chaetotaxic trait which would have permitted to distinguish these
species. I observed that all specimens of pallipes collected from Département Aveyron
bear six long setae on the most posterior cross row of great abd. whilst all other species
have four, and, for a while, I believed that this setal number was a worthwhile
character. Nevertheless, I collected specimens of pallipes from Atlantic populations
with four setae, and in intermediate locations with four to six. Another character lies in
the thickness of male antennal acanthoids, but the difference is not very strong, and this
character does not provide a sharp distinction between both species.
Distinction between D. bicinctus and D. flavus raised another problem. Typical
two black spotted specimens and others uniformly light yellow occurred together in
several populations of bicinctus. This last form absolutely looked like flavus, so that
one could question the specific status of flavus. Nevertheless, I uncovered a chaetotaxic
difference on the fourth antennal segment which permits to maintain the distinction
between bicinctus and flavus. The number of setae of the intergeneratrices Heae and
Hppe, and limited to the section M, is seven in flavus and nine in bicinctus. For these
setae, I observed a compensation phenomenon* in flavus, so that a sharp distinction
* Variable setae may present relationships for their presence vs absence, i. e., statistically
speaking, they are correlated (NAYROLLES 1991a). For a set of correlated setae, histogram of the
variable “observed number of setae” is compared with the theoretical histogram related to the
assumption of independence between setae. In the phenomenon of compensation (NAYROLLES
1993c), the observed histogram is much narrower than the calculated one. Consequently,
characters, i. e. features near constant within and variable between species, should refer not to
individual setae but to sets of correlated setae. In this case, variables are setal numbers.
EUROPEAN BOURLETIELLIDAE 537
between the species cannot be performed by considering the setae one by one but by
calculating a setal number. Another difference lies in the instar of appearance of the
seta (FE2)a5, in third instar in flavus and in adult in bicinctus, and (AD)a+1, in adult in
flavus and in third instar in bicinctus.
ELLIS (1974) described Deuterosminthurus sulphureus mediterraneus, and
justified this subspecies arguing that the male did not bear the two posterior
acanthoids on the upper anal flap, so conspicuous in the typical form. I deem this
character is sufficient for raising the subspecies to specific rank: Deuterosminthurus
mediterraneus Ellis, 1974, stat. nov. Furthermore, the ecological analysis of epigeic
Symphypleona (NAYROLLES 19915) showed that D. sulphureus scarcely occurred in
Mediterranean areas; it was only collected in submediterranean zone, and in stations
where the drought was temperate by local conditions as waterside or undergrowth. On
the other hand, D. mediterraneus is likely a true Mediterranean species since ELLIS
(1974) collected it in Rhodos Island (Greece).
Deuterosminthurus pleuracanthus sp. n. (Figs 17-20; Table III)
MATERIAL EXAMINED - SPAIN, prov. Huesca, Puerto de Monrepos, alt. 1260 m; pasture
and rest area of cattle, with many graminae, Poa sp., Phleum sp., Alopecurus pratensis,
Dactylis glomerata, and also Achillea millefolium, Daucus carota; collected with a net on these
plants (HU24a). 21-VI-1990. Syntypes: 59 specimens; 13 juv., 3 ¢, and 5 9 mounted (8
specimens in alcohol in MHNG, 8 in MNHN, 8 in ZMA, other specimens in LEITT).
Other material.— Same station. 1-VIII-1987: 5 £ and 1 6; 1 9 and 1 d mounted. 30-
V-1990: 1 2 mounted (LEITT).
DESCRIPTION
Development. — Number of juv. st. = 3.
Size ad. — 3: 0.45 - 0.55 mm; 9: 0.6 - 0.85 mm.
Color. — Light yellow, slightly pigmented with orange in posterior area of
great abd. Head light with a small orange fleck on the frontal ocellus. Antennae light
brown-yellow; eyepatches black. Legs very light yellow; furcula white.
Great abd (figs 17 & 18). — Back concave in posterior area. Trichobothria: A,
B, and C in linear pattern. Dorsal setae different between sexes: female with normal
mesochaetae, and male with several swollen and spine-like setae (acanthoids).
Antennae. — Antennal ratios: in 9, ant. I : II: II: IV = 1 : 2.0: 3.1 : 5.5 and
AAC ph a TE into N ante IE IE We 1:72 058565 andantetcephe
diag. = 2.1. Chaetotaxic variables, in both sexes: m(/Sh*) = 0.66 ; in 2 : for G: m =
90.8 / min = 87 / max = 95 ; m(RSh) = 0.44; in & : for G: m = 102.3 / min = 98 / max =
106 ; m(RSh) = 0.39.
Head. — Eyes: 8+8; eyepatch with two setae. Cephalic setae: mesochaetae. No
ov. org. Labral formula: 6/5-5-4.
Legs. — Femur: cup present. Tibiotarsal generatrix Gp without secondary seta.
Pretarsus characteristic of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
538 PIERRE NAYROLLES
Table Ill. Appendicular chaetotaxy of Deuterosminthurus pleuracanthus sp. n.
(T)Q: a+1 (oc=0.3) Q:i-1, pe+1
Other setae: Q: ae+2, pe+2, pe+3
Setae -1
Setae 0
| Setae +1
B completely zoned. Section M euwhorled with 5 whorls. Right euwhorlation.
Subsegmentation formula: 1 + 6 + 1 = (A) + (M1,5 + BA) + (BM + BB). Right subsegmentation.
Apical bulb: joined to the apex
Allpe is a blunt microchaeta
Setae of BE: PF. BBe, BBae, BBa, BBp, BBpo / all G setae on Mare present.
In female: BA with BAi variable (oc = 0.8); n80%(BM) = 10-16; n80%(B) = 23-28; n80%(G) =
88-93. In male: BA complete whorl; n80%(BM) = 21-27; n80%(B) = 34-40; n80%(G) = 99-105.
BBae and BBa are cucumiform setae on large base.
pe
ree à
(oe _
TRI IL |
ae primordial seta: Q: ae3 | T:a5, ai2, pe2, —— = Q: pe4 © a5, ai2, A Q: pe4, Oi
T: pe2, Op Q:ai2, pe4
TI a Dein te Weve _— ‚IVp, Vp
T: 4ai1, Vai, Vpi, FSa, O2pe T: 4ai1, Vai, Vpi, FSa, O2pe : , 4i1, Vai, Vpi,
Q: 3a, 4a1, 4pit, 4ai2, FSai, | Q: 3a, dal, 4pil, 4ai2, FSai,
FSpi FSpi Q: 2a, 3a, Spi, 4a1, 4pi1, 4ai2,
i È È FSai, FSpi
Ja is a winged seta. Ipi, Ip and Ipe are spatulate
Ipi, Ip and Ipe are spatulate | setae. Ip and lpe are spatulate setae.
setae.
T: pe2, pe3
tH Ge [Gae] Ga [Ga Gi | Gpi] Gp [Gpe Other setae: P:2pe
Nile Leake PLP ee
Il | FEW] PRIRER RE
Whors [tll || P| PILP Ee
ROVINA IN| Ses ee a] [PP |Pl|
Date 2er] TE SS
7 | ] { | © I
WholB | Ti P |
HE Chaetotaxy: — un anterior lamella — outer and inner lamellae smooth.
EUROPEAN BOURLETIELLIDAE 539
Fics 17-20
Deuterosminthurus pleuracanthus sp. n.; 17: é, small abd. and lateral and dorsal area of great
abd., lateral view; 18: detail of a swollen seta of great abd.; 19: 2, small abd. in lateral view;
20: right an. app. in ventral view.
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula. — See chaetotaxic table.
Small abd. (figs 19-20). — Made up of abd. V + abd. VI. Trichobothria: D and
E. Male genital papilla with 12 setae. Female circumanal setae avl' and avi shorter
than the others. An. app. fairly long, flat, and with few serration at apex. Ratio an.
app. : mucro = 0.64.
DISCUSSION
The acanthoids of male great abd. permit to distinguish D. pleuracanthus from
all other species of Deuterosminthurus.
Etymology: The name pleuracanthus, from Greek pleuron = flank and acantha
= spine, refers to the acanthoids on the male great abd.
540 PIERRE NAYROLLES
Deuterosminthurus bicinctus (Koch, 1840)
MATERIAL EXAMINED - FRANCE, dép. Gers, district Cravencères, between Cravencères
and Manciet, alt. 150 m; undergrowth of Quercus robur; collected with a net (GS1). 5-V-1990:
189 specimens; 6 juv., 2 2, and 1 d mounted (20 specimens in alcohol in MHNG, 20 in
MNHN, 20 in ZMA, other specimens in LEITT). — Dép. Tarn, district Rouairoux, between
Lacabarède and Rouairoux, about 1-2 km from Lacabarède, alt. 440 m; grove with chestnut,
Quercus pubescens, and Acer pseudo-platanus, in undergrowth: Lonicera periclymenum,
Teucrium scorodonia, Pteridium aquilinum, Euphorbia silvatica, Lamium hybridum, Rubus sp.,
etc.; collected with a net on unselected plants (TN1). 22-VII-1987: 66 specimens; 4 d and 4 9
mounted (LEITT). — SPAIN, prov. Guipüzcoa, between Bergara and Zumarraga, near Puerto
de Descarga, alt. 550 m; damp grove with Alnus glutinosa, Fraxinus excelsior, Sorbus
torminalis, Corylus avellana, in undergrowth: Athyrium filix-femina, Geranium robertianum,
etc. (phytosociological association of Alno Ulmion); collected with a net on unselected plants
(PB1): 36 specimens; 5 juv. mounted (LEITT).
DESCRIPTION
Concerning the appendicular chaetotaxic table, refer to D. pleuracanthus
(Table III), with the following differences:
Antennae. (AD)a+1 ontogeny type T — Occurrences of H setae on M & B: for
Heae 1-1-1-1-0/0, and for Hppe 1-1-1-1-1/0.2. m(h*) = 4.6 — Chaetotaxic variables
in 2: n80%(BM) = 13-16 ; n80%(B) = 26-29 ; n80%(G) = 91-94. In 4 : n80%(BM) =
18-25 ; n80%(B) = 31-38 ; n80%(G) = 96-103. BA complete whorl in both sexes.
Legs. (FE2)a5 ontogeny type Q.
Furcula. (MA)pe2 ontogeny type T(Q), oc = 0.8.
Development. — Number of juv. st. = 3.
Size ad. — 6: 0.5 - 0.55 mm; 9: 0.5 - 0.75 mm.
Color. — Background yellow with, in the normal form, two, one before, one
behind, black flecks on great abd. The anterior spot is the larger. Head dotted with
two small spots behind eyes. In some cases, I found other colored specimens living
with this typical form in a same population. Some had many minute black spots in
place of the black flecks, these minute spots were orange-pink in others, the lightest
form was pigmented with orange, particularly in the rear of great abd., and without a
trace of the typical flecks. In all cases, antennae orange-brown, eyepatches black, legs
and furcula light.
Great abd. — Back concave in posterior area. Trichobothria: A, B, and C in
linear pattern. Dorsal setae: mesochaetae.
Antennae. — Antennal ratios: in 9, ant. 1: If: U1: 1V=1: 1.8: 3.0: 5.6 and
ant. : ceph. diag. = 1.6 ;in 6, ant. 1: I : MTV = 1 21.9: 3.2 = St6fandiantS-ficephe
diag. = 2.1. Chaetotaxic variables, in both sexes: m(/S4*) = 0.77 ; in 2: for G: m =
92.6 / min = 90 / max = 95; m(RSh) = 0.48; in à : for G: m = 99.3 / min = 95 / max =
106 ; m(RSh) = 0.45.
Head. — Eyes: 8+8; eyepatch with two setae. Cephalic setae: mesochaetae. No
ov. org. Labral formula: 6/5-5-4.
Legs. — Femur: cup present. Tibiotarsal generatrix Gp without secondary seta.
Pretarsus characteristic of the genus.
EUROPEAN BOURLETIELLIDAE 541
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula. — Same chaetotaxy as in pleuracanthus (excepted for a manubrial
variable seta).
Small abd. — Made up of abd. V + abd. VI. Trichobothria: D and E. Male
genital papilla with 12 setae. Female circumanal setae with about the same length.
An. app. fairly long, flat, and serrated. Ratio an. app. : mucro = 0.61.
Deuterosminthurus flavus (Gisin, 1946)
MATERIAL EXAMINED — SPAIN, prov. Vizcaya, between Elgoibar and Markina-Xemein,
alt. 350 m; undergrowth of Pinus pinaster, with Athyrium filix-femina, Holcus lanatus, etc.;
collected with a net on unselected plants (PB2). 9-VII-1987: 106 specimens; 7 juv. and 9 ad.
mounted (12 specimens in alcohol in MHNG, 12 in MNHN, 12 in ZMA, other specimens in
LEITT). — Prov. Vizcaya, between Ondanoa and Lekeitio, alt. 100 m; moor with Preridium
aquilinum, Brachypodium pinnatum, Agrostis setacea, Lithospermum prostratum, Andro-
sæmum officinale, Molinia cerulea, etc.; collected with a net on unselected plants (PB3). 9-
VN-1987: 34 specimens; 2 d mounted (LEITT). — Prov. Vizcaya, between Bermeo and
Bakio, near Cabo Machichaco, alt. 70 m; undergrowth of Pinus pinaster with Pteridium
aquilinum, Hedera helix, Viburnum tinus, Smilax aspera; collected with a net (PB19). 16-IX-
1987: 7 specimens; 1 & and 1 2 mounted (LEITT).
DESCRIPTION
Concerning the appendicular chaetotaxic table, refer to D. pleuracanthus
(Table III), with the following differences:
Antennae. (AD)a+1 ontogeny type Q — Occurrences of H setae on M & B: for
Heae 1-1-0.9-0.2-0/0, and for Hppe 1-1-1-0.8-0.2/0. m(h*) = 3.6 — Chaetotaxic
variables in 2: n80%(BM) = 11-15 ; n80%(B) = 24-27 ; n80%(G) = 89-92. In &:
n80%(BM) = 17-21 ; n80%(B) = 30-34 ; n80%(G) = 95-99. BA complete whorl in à,
with variable setae in 2.
Legs. (TIl)3p ontogeny type (Q), oc = 0.3 — (TI1)4pl ontogeny type (Q), oc
= 0.7.
Development. — Number of juv. st. = 3.
Size ad. — d : 0.45 - 0.5 mm; ©: 0.55 - 0.85 mm.
Color. — Background yellow with orange on the back of great abd. Head more
or less pigmented, with a small orange fleck on the frontal ocellus. Antennae light
brown-yellow; eyepatches black, legs very light, and furcula white.
Great abd. — Back concave in posterior area. Trichobothria: A, B, and C in
linear pattern. Dorsal setae: mesochaetae.
Antennae. — Antennal ratios: in 2, ant. 1: IL: Wl: IV=1: 1.9: 2.9 : 5.5 and
die Cepie Gide. — 126-810 ant IE TAN OS SACS andlanteacepine
diag. = 2.1. Chaetotaxic variables, in both sexes: m(/Sh*) = 0.59 ; in © : for G: m =
90.2 / min = 87 / max = 93 ; m(RSh) = 0.40; in & : for G: m = 97.0 / min = 94 / max =
100 ; m(RSh) = 0.38.
542 PIERRE NAYROLLES
Head. — Eyes: 8+8; eyepatch with two setae. Cephalic setae: mesochaetae. No
ov. org. Labral formula: 6/5-5-4.
Legs. — Femur: cup present. The generatrix Gp of forelegs often bears a seta
which is either 3p or 4p1. No seta on Gp of mid and hindlegs. Pretarsus characteristic
of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula. — Same chaetotaxy as in pleuracanthus.
Small abd. — Made up of abd. V + abd. VI. Trichobothria: D and E. Male
genital papilla with 12 setae. Female circumanal setae with about the same length.
An. app. fairly long, flat, pointed and serrated. Ratio an. app. : mucro = 0.52.
Deuterosminthurus sulphureus (Koch, 1840)
MATERIAL EXAMINED — FRANCE, dép. Aveyron, district Bessuéjouls, Le Plateau de la
Justice, alt. 450 m; meadow of the phytosociological association of Orchido-Brometum with
some small trees (Quercus pubescens, Pinus sylvestris, Juniperus communis) and Bromus
erectus, Hippocrepis comosa, Globularia vulgaris, Eryngium campestre, etc.; collected with a
net on unselected plants (AV Ia). Several collections from 17-IV-1987 until 13-VI-1987. In all
55 specimens; 13 juv. mounted (LEITT). — Dép. Aveyron, district Montrozier, Causse Comtal,
road to Gages, alt. 620 m; grass on limestone with Bromus erectus, Keleria valesiaca, Festuca
gr. ovina, Coronilla minima, Hippocrepis comosa, Lotus corniculatus, Anthyllis vulneraria,
Carduncellus mitissimus, Eryngium campestre, Teucrium chamadrys, etc.; collected with a net
on unselected plants (AV2a). Several collections from 17-IV-1987 to 30-VII-1987. Many
specimens; 6 ad. and many juv. mounted (LEITT). — Dép. Haute-Garonne, distric Toulouse,
campus of Université P. Sabatier, alt. 160 m; long grass under fruit trees; collected with a net.
21-V-1988: many specimens collected and mounted (LEITT). — Dép. Ariège, district Ornolac-
Ussat-les-Bains, Ariège river bank, alt. 500 m; long grass with many graminae and Rhinanthus
major, Anthyllis vulneraria, Orchis militaris, etc.; collected with a net on unselected plants
(AR2a). 22-V-1987: many specimens; | juv. and 4 ad. mounted (LEITT).
DESCRIPTION
Concerning the appendicular chaetotaxic table, refer to D. pleuracanthus
(Table III), with the following differences:
Antennae. | have noted differences between two studied populations, parti-
cularly for the number of setae in males. (AD)a+1 ontogeny type T — (AD)pe+1
ontogeny type (T)Q or Q depending on populations — Occurrences of H setae on M
& B: for Heae 1-1-1-1-0/0, and slightly variable for Hppe, 1-1-0.8-0.3-0/0 or 1-1-1-0-
0/0, depending on populations. In all cases, m(h*) = 3.5 — Chaetotaxic variables in
2: n80%(BM) = 10-15 ; n80%(B) = 23-28 ; n80%(G) = 88-93. 3 with differences
between populations: n80%(BM) = 20-25 / 24-30 ; n80%(B) = 33-38 / 37-43 ;
n80%(G) = 98-103 / 102-108. BA complete whorl in both sexes.
Legs. (TI1)3p ontogeny type Q — (TIl)4pl ontogeny type Q — (TI2)3p
ontogeny type (Q), oc = 0.7 — (TI2)4p1 ontogeny type (Q), oc = 0.2 — Setae of the
EUROPEAN BOURLETIELLIDAE 543
outer side of tibiotarsi, (TI.)IlIe, (TI.)ITlae, (TI.)IMpe, (TI.)IVe, (TI.)IVae, (TI.)IVpe,
(TI1,2)Ve, (TI1,2)Vae, (TIl,2)Vpe, (TI3)IHe, are macrochaetae, their length grows
during development.
Furcula. The primary seta (DE)IVpi is generally present, but may be variable
in some populations.
Development. — Number of juv. st. = 3.
Size ad. — G : 0.55 - 0.6 mm; 9: 0.7 - 1.0 mm.
Color. — Golden yellow with more orange in males. A small orange fleck on
the frontal ocellus. Eyepatches black, antennae light yellow, legs very pale yellow,
furcula white.
Great abd. — Back concave in posterior area. Trichobothria: A, B, and C in
linear pattern. Dorsal setae: mesochaetae.
Antennae. — Antennal ratios: in 9, ant. 1: I : IT : IV = 1 : 1.8: 2.9: 5.4 and
anesecphaldiag Slo cantate: NE IV =e k9 = 3.3) "6:4 and: ant: ceph:
diag. = 2.1. Chaetotaxic variables of d have not the same values between popu-
lations; absolute minimum and maximum as well as range of average are given. In
both sexes, m(/Sh*) = 0.58. In © : for G: m = 90 / min = 86 / max = 94 ; m(RSh) =
0.40. In &: for G: m = 100.5 - 104.9 / min = 95 / max = 112 ; m(RSh) = 0.34 - 0.36.
Head. — Eyes: 8+8; eyepatch with two setae. Cephalic setae: mesochaetae. No
ov. org. Labral formula: 6/5-5-4.
Legs. — Femur: cup present. Tibiotarsal generatrix Gp with two setae on fore-
legs, and often one on midlegs. Several long macrochaetae on tibiotarsal outer side.
Pretarsus characteristic of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula — (DE)IVpi variable in some populations.
Small abd. — Made up of abd. V + abd. VI. Trichobothria: D and E. Male
genital papilla with 12 setae. Female circumanal setae avl' and avl shorter than the
others. An. app. fairly long, flat, pointed and slightly serrated. Ratio an. app. : mucro
= 0.68.
Deuterosminthurus maculatus sp. n. (Figs 21-24; Table IV)
MATERIAL EXAMINED — FRANCE, dép. Aveyron, district Montrozier, Causse Comtal,
road to Gages, alt. 620 m; collected with a net on Juniperus communis (AV2d). 23-V-1987.
Syntypes: 7 juv. and 20 ad.; 5 juv., 3 2, and 2 d mounted (LEITT).
Other material— Same station. 17-IV-1987: 2 juv. and 1 ®. 30-VII-1987: 1 juv
(LEITT). — Same location; grass and small shrub at the edge of a grove of Quercus pubescens,
with Teucrium chamadris, Hippocrepis comosa, Spiræa hypericifolia, Helianthemum canum,
Ornithogalum umbellatum, Euphorbia dulcis, and Fumana procumbens; collected with a net on
unselected plants (AV2e). Several collections from 17-IV-1987 to 1-X-1987. In all 24
specimens; 4 juv., 2 2, and 1 d mounted (LEITT). — Dép Aveyron, district Bessuéjouls, Le
Plateau de la Justice, alt. 450 m; meadow belonging to the phytosociological association of
544 PIERRE NAYROLLES
Orchido-Brometum with some small trees (Quercus pubescens, Pinus sylvestris, Juniperus
communis), and Bromus erectus, Hippocrepis comosa, Globularia vulgaris, Eryngium
campestre, etc.; collected with a net on unselected plants (AV la). 17-IV-1987: 1 2 mounted.
23-V-1987: 1 juv. (LEITT). — Dep. Aude, district Valmigere, road D54 between Valmigere
and Arques, near Valmigère, alt. 680 m; moor with Preridium aquilinum; collected with a net
on this plant (AUl le). 8-VII-1987: 33 specimens; 4 juv., 1 2, and 3 d mounted (LEITT). —
Dep. Haute-Garonne, district Roquefort-sur-Garonne, field between the roads D62 and N117,
alt. 270 m; hedge with Salix capræa, Cornus sanguinea, Rubus sp., Rosa sp., Cytisus scoparius,
Quercus pubescens, Clematis vitalba, etc.; collected with a net on the hedge (HG5b). 16-VI-
1987: 116 specimens (12 specimens in alcohol in MHNG, 12 in MNHN, 12 in ZMA, other
specimens in LEITT).
DESCRIPTION
Development. — Number of juv. st. = 3.
Size ad. — 36: 0.5 - 0.6 mm; 2: 0.65 - 0.9 mm.
Color (fig. 21). — Background white or very light yellow in female, yellow in
male. Several brown patches arranged in a characteristic pattern. Spots of female
forming a pair of laterodorsal stripes along the great abd. A mediodorsal band more or
less pigmented with orange. The laterodorsal stripes are completely or partially
connected by two cross flecks, one is near the halfway back, the other situated in the
rear quarter. End of great abd. spotted with a transversally stretched out fleck which
does not reach the laterodorsal stripes. Two oblong patches in a very lateral position.
Expanse of brown pigment on small abd. varying between specimens. Expanse of
dark pigment is lesser in males, lateral brown stripes being narrower and vanished
forward about halfway the back. At this level, the cross fleck, present in females, is
absent or very faded in males. Head colored with yellow excepted for a light area
around the eyepatches. Eyepatches black. Color of antennae varying from yellow to
brown, lighter on the three first segments than on the fourth. Legs light yellow;
furcula white.
Great abd (fig. 24). — Back concave in posterior area. Trichobothria: A, B, and
C in linear pattern. Dorsal setae: mesochaetae. Posterior cross row with always 4 setae.
Antennae. — Several setae transformed in acanthoids on ant. I and II in adult
male Antennal ratios: in 2, ant. 1: IL: IMI ; IV =1 =: 1.8 : 2.6 3 5:1 and antcepht
diag. = 1.6; n.d, ant. LIL: IM; IV = 1: 1.8: 2.82 5.5 and ant. °Ceph dia PA!
Chaetotaxic variables, in both sexes: mUSh*) = 0.67 ; in 2: for G: m = 97.0 / min =
94 / max = 101 : m(RSh) = 0.41 : in 6 : for G: m= 104.2 / min = 07 max
m(RSh) = 0.38.
Head. — Eyes: 8+8; eyepatch with two setae. Cephalic setae: mesochaetae,
some facial setae being longer in male than in female. 1+1 ov. org. in the back-ventral
area (it remains the ventral pair). Labral formula: 6/5-5-4.
Legs. — Femur: cup present, as well as the seta (FE3)pel. Tibiotarsal
generatrix Gp with two setae on forelegs, and two on midlegs. Pretarsus characteristic
of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
EUROPEAN BOURLETIELLIDAE 545
Table IV. Appendicular chaetotaxy of Deuterosminthurus maculatus Sp. n.
Cori EE a Owe SEO See [i GT OE
Ai is a trichobothrium. Ae, Be, Ape, Bpe, e+1, and pe+1 are acanthoids in male; Ap is also an
acanthoid, but less stout than the others.
Other setae: Q: ae+2, pe+2, pe+3
Setae -1
Setae 0
Setae +1
B completely zoned. Section M euwhorled with 5 whorls. Right euwhorlation.
Tentato formula: 1+6+1 = (A) + (M1, 5 + BA) + (BM + BB). Right subsegmentation.
Apical bulb: joined to the apex Subapical organ: very small D: Alai
Allpe is a blunt microchaeta
Occurrences of H setae on M & B: for Heae 1-1-1-1-0/0, for Hipi 1-1-1-0-0/0, and for
Hppe 1-1-1-1-0/0. È
Setae of BB: P: BBe, BBae, BBa, BBp, BBpe / all G setae on M are present.
In female: BA: complete whorl; n80%(BM) = 17-22; n80%(B) = 30-35; n80%(G) = 95-100.
In male: BA: complete whorl; n80%(BM) = 27-31; n80%(B) = 40-44; n80%(G)= 105-109.
BBae and BBa are cucumiform setae on large base.
P: 1 seta
P: 1 seta
P:ae,i1,ms T: Oi1 P:ae,i1,ms T: a, ai2, Oil
delayed primordial seta: Q: a
T:pe2,Op Q:ai2, pe4
T: 4ai1, Vai, Vpi, FSa, O2pe T: 4ai1, Vai, Vpi, FSa, O2pe T: Sai, 3i, 4ai1, 411, Vai, Vpi,
Q: 3a, 3p, 4a1, 4pil, 4p1, 4ai2, | Q: 3a, 3p, 4al, 4pil, 4p1, 4ai2, FSa, O2pe
FSai, FSpi FSai, FSpi Q: 2a, 3a, Spi, 4a1, 4pil, 4ai2,
FSai, FSpi
Ipi, Ip and Ipe are spatulate | setae. Ip and lpe are spatulate setae.
setae.
Ja is a winged seta. Ipi, Ip and Ipe are spatulate
A aN
II Mi ‘ai a AAA |
Other setae: P:2pe
oc(IVpi) =
WhorlB
GE Chaetotaxy: — e anterior An — outer =“ inner lamellae smooth.
546 PIERRE NAYROLLES
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula. — (DE)IVpi often absent.
Small abd. (figs 22-24). — Made up of abd. V + abd. VI. Trichobothria: D and
E. Male genital papilla with 12-13 setae. Female circumanal setae with about the
same length. An. app. flat, pointed and serrated. Ratio an. app. : mucro = 0.49.
DISCUSSION
D. pallipes and D. maculatus sp. n. can be distinguished from the other species
of Deuterosminthurus by the seta (FE3)pel present from the Ist st. on, the male
acanthoids on ant. I and II (see ELLIS, 1974, fig. 7b p. 135), and 1+1 ov. org. behind
the head. The color pattern is very characteristic and invariable in maculatus, whereas
pallipes displays a wide range of colors without specific pattern. Two other characters
can be used: pallipes has 4 to 6 long setae on the last cross row of great abd., and
maculatus, as the other species of Deuterosminthurus I have observed, always bears 4
setae, and the antennal acanthoids of maculatus are thicker than in pallipes.
Etymology: The name maculatus from Latin macula = mark, spot.
Deuterosminthurus pallipes (Bourlet, 1843) (Fig. 25)
MATERIAL EXAMINED — FRANCE, dép. Aveyron, district Montrozier, Causse
Comtal, road to Gages, alt. 620 m; grass on limestone with Bromus erectus, Keleria
valesiaca, Festuca gr. ovina, Coronilla minima, Hippocrepis comosa, Lotus corni-
culatus, Anthyllis vulneraria, Carduncellus mitissimus, Eryngium campestre, Teu-
crium chamadrys, etc.; collected with a net on unselected plants (AV2a). Several
collections from 17-IV-1987 to 11-IX-1987. In all 17 juv. and 11 ad.; 6 ad. and all
Juv. mounted (LEITT). — Same location; doline with Molinia carulea, Avena praten-
sis, Eryngium campestre, Carduncellus mitissimus, Lathyrus pratensis, Vicia pere-
grina, and some specimens of Prunus spinosa; collected with a net on unselected
plants (station AV2b). Several collections from 17-IV-1987 to 1-X-1987. In all 123
specimens; 32 juv. and 9 ad. mounted (LEITT). — Dép. Aveyron, district Sébazac-
Concourès, Causse Comptal, road D581 between Lioujas and Concoures, alt. 600 m;
collected with a net on Spiræa hypericifolia (AV7d). 21-VI-1986: many specimens; 4
juv. mounted (LEITT). — Dép. Pyrénées-Orientales, district Lesquerde, road D619
between Saint-Paul-de-Fenouillet and Ansignan, about 3 km from Saint-Paul-de-
Fenouillet, alt. 350 m; scrub with Quercus ilex, Bupleurum fruticosum, Quercus
coccifera, etc.; collected with a net on unselected plants (PO15a). 24-VI-1987: 142
specimens; 8 juv. and 7 ad. mounted (LEITT).
DESCRIPTION
Concerning the appendicular chaetotaxic table, refer to D. maculatus (Table
IV), with the following differences:
EUROPEAN BOURLETIELLIDAE 547
Fics 21-25
Figs 21-24: Deuterosminthurus maculatus sp. n. 2; 21: habitus; 22: small abd. in lateral view;
23: right an. app. in ventral view; 24: small abd. and posterior area of great abd. in dorsal view,
arrows point setae of the most posterior row of great abd. Fig. 25: Deuterosminthurus pallipes
2, small abd. and posterior area of great abd. in dorsal view, arrows point setae of the most
posterior row of great abd.
548 PIERRE NAYROLLES
Antennae. Acanthoids of male are a little less stout than in maculatus, and
(AD)Ap cannot be termed an acanthoid. I have noted some differences between
populations. (AD)pe+1, depending on populations, appears or not at 3rd st. (oc
varying from 0 to 1) — Occurrences of H setae on M & B: for Heae 1-1-1-1-0/0, and
fluctuate for Hppe from 1-1-1-1-0/0 to 1-1-1-1-0.8/0. Consequently, m(h*) varies
from 4.0 to 4.4. — Chaetotaxic variables in 9: n80%(BM) = 12-15 / 14-17 ; n80%(B)
= 25-28 / 27-30 ; n80%(G) = 90-93 / 92-95. In d: n80%(BM) = 19-24 / 22-30 ;
n80%(B) = 32-37 / 35-43 ; n80%(G) = 97-102 / 100-108. BA complete whorl in both
sexes.
Furcula. The primary seta (DE)IVpi is seldom present, its occurrence varying
from 0 to 0.2.
Development. — Number of juv. st. = 3.
Size ad — 6.2049 059:mm: 2.0.6 -.0.75:mm:
Color. — Very variable between populations, from light yellow to very dark
purple. Males are generally more colored than females. Color depends on climatic
zones in which populations live. Males of the Mediterranean area vary from orange
flecked to entirely dark red. Females are generally yellow, with sometimes orange or
ochre spots. In Atlantic area, two different patterns are mixed, most of specimens are
yellow, some others, males and females, are dark purple (this form corresponds to the
typical color of pallipes). In mountains, specimens living on Rhododendron ferrugi-
neum are wine-red (only one yellow female was observed in a collection of more 100
specimens). In other areas, specimens are yellow (this form was described as repan-
dus). In Mediterranean populations, females are colored with orange or ochre shades
as in D. maculatus, but the pigment is far more diffuse than in maculatus. In addition,
color widely varies between specimens, and in these populations, entirely red males
are present, so that one cannot confuse the species. Head more or less colored,
depending on the expanse of pigment on the body; surround of eyepatches often
remains light as in maculatus. Eyepatches black, antennae with pigment varying from
yellow to brown, legs light yellow, furcula white.
Great abd. (fig. 25). — Back concave in posterior area. Trichobothria: A, B,
and C in linear pattern. Dorsal setae: mesochaetae. Posterior cross row with 4 to 6
setae.
Antennae. — Several setae transformed in acanthoids on ant. I and II in adult
male. Antennal ratios: in 2, ant. 1: IT : II: IV = 1 : 1.8: 2.7 : 5.2 and ant. : ceph. diag.
= 16; in Gant 1: WM: TI: IV =1: 1.7: 27: 54 and ant Zeephadiaers ae
Chaetotaxic variables with some differences between populations. Absolute minimum,
maximum, and range of average are given. In both sexes: m(/Sh*) = 0.67 - 0.73. In 9:
for G: m = 91.8 - 93.1 / min = 90 / max = 96 ; m(RSh) = 0.44 - 0.46 ; in à : for G: m=
99.9 - 104.3 / min = 97 / max = 109 ; m(RSh) = 0.40 - 0.41.
Head. — Eyes: 8+8; eyepatch with two setae. Cephalic setae: mesochaetae,
some facial setae being longer in male than in female. 1+1 ov. org. in the back-ventral
area (it remains the ventral pair). Labral formula: 6/5-5-4.
EUROPEAN BOURLETIELLIDAE 549
Legs. — Femur: cup present, as well as the seta (FE3)pel. Tibiotarsal
generatrix Gp with two setae on forelegs, and two on midlegs. Pretarsus characteristic
of the genus.
Ventral tube. — Sacs warty from 2st st. on. Chaetotaxy: apical flaps with one
pair of primary setae, corpus without seta.
Retinaculum. — At Ist st. rami tridentate, bidentate from 2nd. Chaetotaxy: on
anterior lobe two setae appear at 3rd st. and one in ad.
Furcula. — In most cases, (DE)IVpi absent.
Small abd. (fig. 25). — Made up of abd. V + abd. VI. Trichobothria: D and E.
Male genital papilla wıth 12-13 setae. Female circumanal setae with about the same
length. An. app. flat, pointed and serrated. Ratio an. app. : mucro = 0.52.
DISTINCTION OF THE EUROPEAN SPECIES OF Deuterosminthurus
A key to the European species of Deuterosminthurus is provided, that includes
the studied species as well as D. pandayi Ellis, 1974 and D. mediterraneus Ellis,
1974, stat. nov. I recall that D. cruciata Haybach, 1972 should be taken out from
Deuterosminthurus and placed in Cassagnaudiella (ELLIS 1975; NAYROLLES 1995).
D. quadranculata (Loksa & Bogojevis, 1970) was described from only one female.
The description was insufficient, and this species ıs not included in the key.
In their description of D. quadranculata, LOKSA & BOGOJEVIS (1970) noted:
"Die neue Art steht der Art repanda (Ägren, 1903) am nächsten. Unterscheidet sich
von dieser durch das Muster, durch die Zahl der zirkumanalen Borsten und durch die
Anordnung dieser." However, the figure of small abd. given by LOKSA & BOGOJEVI2
(ibid., fig. 63, p. 141) does not show any difference from pallipes. The claw of
quadranculata was described as untoothed; on the contrary, a tooth is generally
present on the inner side of the claw of pallipes. Nevertheless, this character, difficult
to observe and sometimes variable, has long been given an excessive importance, and
I consider it to be irrelevant for distinguishing species. The color does not make up a
good character any more. It ıs variable, without a defined pattern in pallipes; on the
other hand, guadranculata is not really different from certain colored specimens of
pallipes. Consequently, the distinction between quadranculata and pallipes may be
questioned. However, LOKSA & BOGOJEVI3 described a retinaculum with two setae,
and all Deuterosminthurus species I have observed have three setae in adult. Do the
two quadranculata’ s setae come from a mistake in observation, an aberrant
specimen, or an actual species character? Only new collections in the location in
which guadranculata was found (Deliblat, Yugoslavia) and new observations may
give an answer.
KEY TO EUROPEAN SPECIES OF Deuterosminthurus
| - three thick setae (Ila, IIIa, and Ilai) on anterior dental side .......... pandayi
-Sn Outi ck Selac ON MÉNTENSE RE APE A al. ogre nope ed ce oe cunt TRE ee cs 2
550 PIERRE NAYROLLES
DRE im acrochaetaetontonterisidetofitibiotarsi EE 2.200 eee 3
Mubiotarsiwithoutmacrochaetae® u. LEO +
3 - posterior margin of the male upper anal flap with 1+1 acanthoids . . sulphureus
—malesupper analstlap_ without acanthoid ES, 00 mediterraneus
4 - (FE3)pl present, 1+1 ov. org. behind head, several acanthoids on male
ants angel. RER nl Bl ie 5
- (FE3)pl absent, no ov. org. behind head, no acanthoid on male ant. I
andere EV ina ead oo, at Re Go 6 0 0 0 © 6
5 - brown spots forming a characteristic pattern, 4 long setae on the most
POStEM ONTO OMercab abd! = eee maculatus Sp. n.
- variable color without a defined pattern, 4 to 6 long setae on the most
posterior row Of PTE AT ADEME O. cua: MEME RTE RENNES pallipes
Gere malesgreatabd with acanthoidsen re ee ee pleuracanthus sp. n.
= no,acanthoid.onimale great abdi troia eee 7
7. - back of great abd. with often two black patches, ant. IV with 9 setae in
thessewHleae-t"Lipperor M section? re RIT bicinctus
- back of great abd. unstained, ant. IV with 7 setae in the set Heae +
Fippe OMNISECHON Ne RE flavus
For the studied species, we can distinguish, on the one hand, those (maculatus
and pallipes) displaying the seta (FE3)pel from the Ist instar on, acanthoids on ant. I
and II in males, and 1+1 ov. org. on the back-ventral area of head, and on the other
hand, those (sulphureus, pleuracanthus, bicinctus, and flavus) without the seta
(FE3)pel, nor male special shaped setae, nor cephalic ov. org. Concerning pandayi
and mediterraneus, we know from ELLIS (1974) that they have no male antennal
acanthoid, it will be then of interest to determine whether the seta (FE3)pel and ov.
org. behind the head are present or absent.
REFERENCES
BETSCH, J. M. 1974. Etude des Collemboles de Madagascar. II. - Principaux cadres génériques
des Symphypléones de l'étage montagnard. Bulletin du Muséum national d'Histoire
naturelle, Paris, 3° série, 219, Zoologie, 147: 529-569.
BETSCH, J. M. 1980. Eléments pour une monographie des Collemboles Symphypléones
(Hexapodes, Aptérygotes). Mémoires du Muséum national d'Histoire naturelle, Paris,
AIG 22 7
BETSCH, J. M. & Z. Massoup. 1970. Etude sur les Insectes Collemboles. I. Systématique,
ultrastructure externe et écologie du genre Jeannenotia Stach, 1956 (Symphypléones,
Sminthurididae n. comb.). Description de deux Collemboles nouveaux (Proisotoma et
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BETSCH, J. M. & A. WALLER. 1989. L'armement en trichobothries des Collemboles Symphy-
pléones. Recherche de schémas chétotaxiques, pp. 15-31. /n Proceedings of the Third
International Seminar on Apterygota (R. Dallai ed.). Universita di Siena, Italy.
EUROPEAN BOURLETIELLIDAE 551
BETSCH, J. M. & A. WALLER. 1994. Chaetotaxic nomenclature of the head, thorax, and abdo-
men in Symphypleona (Insecta, Collembola). Acta zoologica Fennica, 195: 5-12.
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secta, Collembola, Symphypleona). Abhandlungen und Berichte des Naturkunde-
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BRETFELD, G. 1990. Chaetotaxy of four Species of the Genera Heterosminthurus, Bourletiella,
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ELLIS, W. N. 1975. On Bourletiella (Cassagnaudiella) pruinosa (Tullberg, 1871) and its allies
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a ¥
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ME Eur Bar:
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DA: MINE
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“1984S
REVUE SUISSE DE ZOOLOGIE, 103 (2): 553-564; juin 1996
Influence de l’exploitation forestière sur trois familles
de coléoptères liés au bois dans les Gorges de I’ Areuse!
Sylvie BARBALAT
Institut de zoologie
Emile Argand 11
2007 Neuchatel, Suisse.
Influence of forest management on three wood-eating Beetles families
in the Areuse Gorges (Canton of Neuchatel, Switzerland). - The species
richness and abundance of selected wood-eating Beetles (Buprestidae,
Lucanidae and Cerambycidae) between managed and unmanaged forests
since 1945 have been compared. The following traps were used: window
traps, small Malaise traps, beer traps and yellow plates. Among the 46
collected species, 34 belonged to the Cerambycidae, 9 to the Buprestidae
and 3 to the Lucanidae. This beetle fauna is not richer in the unmanaged
forests and is favourably influenced by the presence of clearings. Species
characteristic of old forests are missing in the studied area.
Key-words: Cerambycidae - Buprestidae - Lucanidae - Forest ecology -
Swiss Jura.
INTRODUCTION
Depuis le début du siècle, on a pu constater une forte raréfaction d’espèces de
Coléoptères du bois autrefois fréquentes. La liste rouge allemande des espèces mena-
cées (GEISER 1984) signale à ce sujet 60% d’espèces en danger parmi ces Coléopteres.
On ne dispose de données, principalement muséographiques, que depuis la fin du
XIXe siècle, mais il n’est pas exclu que le déclin de certaines espèces ait commencé
bien avant. Certaines de ces espèces, qui avaient un impact économique non négli-
geable, comme les Cérambycides Saperda carcharias (L.) et Tetropium castaneum
(L.) ou le Bupreste Agrilus viridis (L.) (SCHAEFFER 1949; BRAUNS 1964; CHARRARAS
1972), ne causent pratiquement plus de dégâts aujourd’hui, du moins dans la zone
étudiée (P. JUNOD, com. pers). Les espèces potentiellement nuisibles à la sylviculture
ne représentent, de plus, qu'une faible minorité des Coléoptères du bois, les autres,
comme par exemple les Lucanides Lucanus cervus (L.) et Dorcus parallelipipedeus
(L.), jouent un rôle essentiel dans le recyclage du bois mort. La régression de cette
| Cet article fait partie de la thèse de l’auteur.
Manuscrit accepté le 30.05.1995.
554 SYLVIE BARBALAT
faune est généralement attribuée à la raréfaction des arbres vieux ou morts (Geiser,
1984), ainsi qu'aux enrésinements (SPEIGHT 1989). Afin de conserver une richesse
faunistique maximale dans nos forêts, divers auteurs préconisent une sylviculture plus
extensive avec, par exemple, l'abandon des plantations d'arbres étrangers à la station,
la conservation des vieux arbres ou le maintien de forêts claires. (GEISER 1980;
PAULUS 1980; LSPN, 1993).
Cependant, à notre connaissance, très peu de travaux ont été consacrés à
l'étude de l'entomofaune du bois en relation avec l'exploitation ou la structure de la
forêt. Citons toutefois le travail de SPRECHER-UEBERSAX (1989) qui s'est intéresssé à
l'influence de la quantité de bois mort sur les Coléoptères du bois.
Pour cette première année de recherche, nous nous sommes fixé deux buts:
- le premier consiste à estimer l’influence de l’exploitation forestière sur trois
familles de Coléoptères liés au bois. Pour ce faire, nous avons tenté, pour quelques
types de forêts, de trouver une parcelle exploitée à comparer avec une parcelle
inexploitee;
- le second est l’étude de l’influence de deux éléments structurels supposés
favorables à ces insectes: les clairieres et le bois mort.
Les Coléoptères du bois formant un groupe très vaste, seules trois familles ont
été retenues. Il s’agit d’une part des Cérambycides et des Buprestides, familles essen-
tellement xylophages dont les larves se nourrissent de bois mort relativement frais, et
d'autre part des Lucanides, famille saproxylophage dont les larves se nourrissent
plutôt de bois en décomposition. Etant donné notre problématique, il nous a paru
pertinent de choisir des familles au sujet desquelles il existe une bonne documen-
tation, aussi bien bibliographique que muséographique. Elle permet une interprétation
appropriée des résultats en nous renseignant sur la biologie des espèces et l'évolution
de leur statut en Suisse. De plus, vu la raréfaction de nombreux représentants de ces
familles, il paraissait important de mieux connaître les milieux qu'ils fréquentent afin
de proposer des mesures de protection les plus adéquates possibles.
DESCRIPTION DES MILIEUX
Les Gorges de l'Areuse, à l'entrée du Val de Travers (NE), présentent toute
une gradation de forêts allant de la thermophile chênaie buissonnante à la monta-
gnarde hêtraie à Asplenium. Le terme de “station” désignera ci-après l’intérieur du
périmètre où les pièges ont été posés, c’est-à-dire environ 2500 m?. La figure 1
indique l'emplacement des stations dans la zone d’étude.
Dans le canton de Neuchâtel, le type d'exploitation forestière le plus répandu
est celui de la forêt jardinée. Il vise à maintenir dans la forêt des arbres de tous âges.
Lors de coupes, seuls les arbres d'âge mûr sont abattus, ce qui maintient un couvert
arborescent relativement constant. Toutefois, afin de favoriser des essences de lu-
mière comme le chêne (Quercus sp.) ou le pin sylvestre (Pinus sylvestris), les fores-
tiers pratiquent en certains endroits des ouvertures appelées coupes d'abri. Dans notre
zone, elles se rencontrent dans une aire assez restreinte où la composition du
peuplement est hétérogène et la forêt claire car le sol y est souvent mince. Ce sont les
COLEOPTERES LIÉS AU BOIS 555
Rochefort
722m
Chambrelien
Gare de Chambrelien
630m
Fretereules
Légende: = nvière
—— routes
LINGE CE RO —— lignes de chemin de fer
0 1 Km È x
= lignes de crêtes
Fic. 1
Situation des stations étudiées
uniques clairières rencontrées dans la zone étudiée. Elles mesurent le plus souvent
entre 600 m? et un hectare. Aucune forêt vierge ne subsiste dans le canton. Toutefois,
l'exploitation de forêts situées sur des pentes très raides a été abandonnée à la fin de la
deuxième Guerre mondiale.
Le bois mort est présent sous forme de branches de diamètre variable entassées
après une coupe, de petits arbres abattus ou de petits arbres secs sur pied.
Contrairement aux apparences, 1l n’est pas si simple de trouver une parcelle de forêt
sans bois mort. En ces périodes de conjoncture morose, seul le “bois rentable” est
évacué. Les petits arbres secs ne gênant pas la croissance des autres ainsi que les tas
de branches sont laissés sur place. L'inventaire de ces derniers a permis d'évaluer la
quantité de bois mort.
A l’intérieur d’une forêt exploitée, en l’occurence une hétraie thermophile, on
souhaitait évaluer l’importance de deux éléments structurels supposés favorables aux
insectes xylophages: les clairieres et le bois mort. A l’intérieur de ce type de forêt, il
fallait donc trouver quatre stations aux caractéristiques suivantes:
556
SYLVIE BARBALAT
- présence d’une clairière et de bois mort;
- présence d’une clairière et absence de bois mort;
- absence de clairière et présence de bois mort;
- absence de clairière et de bois mort;
Le tableau 1 résume les principales caractéristiques des 10 stations choisies.
TABLEAU 1: description des stations.
alt. = altitude, exp. = exposition; recouvr. = recouvrement de la strate arborescente, bm = station avec
bois mort, cl. = station avec clairière, coupe = date de la dernière coupe.
station commune coordonnées alt. exp. pente forêt exploitation recouvr. coupe
1 Rochefort 55172018 720m SE 30% chénaie exploitée 40% 1990
2 Rochefort 55142017 720m SE 60% chênaie inexploitée 40% 1949
3 Rochefort 55192027 790m SE 20% hétraie exploitée 90% 1977
4 Rochefort 5518 2025 760m SE 40% hêtraie exploitée, bm 75% 1991
5 Rochefort 55202019 725m SE 20% hêtraie mixte exploitée, cl., bm 40% 1985
6 Rochefort 5519 2025 770m SE 15% hêtraie mixte exploitée, cl. 60% 1987
7 Rochefort 5513 2018 670m SE 90% hêtraie inexploitée 90% 1944
8 Boudry 5500 2006 890m NW 50% hétraie à sapins exploitée 75% 1991
9 Boudry 5498 2008 810m NW 30% hêtraie à sapins "inexploitee"* 80% 1974
10 55122011 825m inexploitée 60% 1938
Boudry NE 100% hétraie à sapins
* La station 9 sert de parcelle-témoin pour l'observation du dépérissement des forêts; aucune inter-
vention forestière n'y a été réalisée depuis 1985, pas même le débardage des chablis.
MATERIEL ET METHODES
Ce travail a pour but de comparer l'abondance en Coléoptères du bois et leur
richesse spécifique dans les différentes stations. Nous avons donc opté pour des
pièges pouvant être disposés équitablement partout et représenter un effort d'échan-
tillonnage identique pour toutes les stations.
Les pièges suivants ont été employés (BARBALAT, 1995): le piège-fenêtre (80
x 50 cm; 1 par station) (HARTMANN & SPRECHER 1990), la mini-Malaise (40 x 40 x 25
cm; 3 par station) et l’assiette colorée (diamètre: 20 cm / hauteur: 10 cm; 1 par
station) (BASSET 1985), ainsi que le piège à bière (diamètre: 8 cm / hauteur: 23 cm; 3
par station) (ALLEMAND & ABERLENC 1991); en parallèle, une tournée hebdomadaire
(quand la météo était favorable) de chasse à vue a été effectuée. Cette dernière
méthode nous a fourni des informations essentiellement faunistiques, puisque les
insectes capturés de cette façon se trouvaient le plus souvent en dehors des stations.
La saison de piégeage a duré de début mai à mi-septembre 1993. Les pièges ont été
relevés tous les 10 jours. Il n'a pas été possible de poser de piège-fenêtre dans la
station 10 trop escarpée.
Les résultats obtenus ont été traités par une analyse canonique des correspon-
dances (ci-après: ACC) (TER BRAAK 1986, 1988a) à l’aide du programme CANOCO
(TER BRAAK 19885), cela afin d’identifer les variables les plus pertinentes qui influ-
encent la distribution des espèces dans les stations étudiées. Cette technique permet
d'extraire d'une matrice de données la variance expliquée par une ou des variables
COLEOPTERES LIÉS AU BOIS 557
explicatives introduites a priori. De même qu'en régression multiple, ces variables
peuvent être sélectionnées selon une procédure pas à pas. Les trois variables expli-
catives suivantes ont été introduites dans l'analyse pour y faire l'objet d'une sélection
pas-à-pas: presence ou absence de clairiere, présence ou absence d'exploitation et
présence ou absence de bois mort.
RESULTATS
Le tableau 2 présente les 46 espèces récoltées dans la zone d’étude. La répar-
tition spécifique par taxon est la suivante: Cerambycidae: 34 espèces; Buprestidae: 9
espèces; Lucanidae: 3 espèces. Les résultat obtenus par les différents pièges sont dis-
cutés dans une publication séparée (BARBALAT, 1995).
Comme une sélection pas à pas l'a montré, seule la variable “presence ou
absence de clairière” explique une proportion significative de la variance des données
(19,3% de la variance; p=0,01). Le tableau 3 a été diagonalisé en fonction de la
position des stations et des espèces sur le premier axe canonique, qui représente cette
partie de la variance.
DISCUSSION
INFLUENCE DES CLAIRIERES
Le petit Cérambycide floricole Strangalia melanura représente à lui seul
respectivement 84% et 56% des captures dans les stations 5 et 6, les seules situées en
clairière. Si l’on fait abstraction de cette espèce, sujette à d’importantes fluctuations
d'effectifs, ces deux stations conservent en moyenne un nombre d’individus plus de
dix fois supérieur à celui des autres stations, ainsi qu’un nombre d’espèces plus de
trois fois supérieur.
L'arrivée de soleil au sol dans les clairieres y permet une activité de l’entomo-
faune beaucoup plus importante que dans une forêt fermée. De plus, les fleurs de la
clairière représentent une source de nourriture pour beaucoup d'espèces butineuses et
favorisent les rencontres de partenaires sexuels. Il est donc logique que nos pièges,
qui mesurent essentiellement l'activité des insectes, se soient révélés plus efficaces en
clairière. La quantité d'individus dans les pièges peut donc simplement refléter une
activité accrue en clairière qui n'est pas forcément liée à l'abondance des insectes dans
le milieu. Cela dit, il paraît raisonnable de considérer que, pour les raisons évoquées
plus haut, une clairière représente un milieu fortement attractif, notamment pour les
espèces floricoles. De plus, les Cérambycides, qui constituent l'essentiel de nos
captures, sont généralement thermophiles, aussi bien à l'état larvaire qu'imaginal. Ils
auront tendance à pondre dans des troncs ou des branches exposés au soleil, donc de
préférence en clairière ou en lisière (KLAUSNITZER & SANDERS 1981). A notre avis, la
forte activité constatée dans les clairieres est donc également liée à une plus grande
abondance d'individus.
Il est intéressant de constater l'importance des milieux semi-ouverts pour ces
insectes, pour la plupart réputés forestiers. Dans notre cas, seul le Cérambycide
558 SYLVIE BARBALAT
Leiopus nebulosus ne semble pas avoir besoin de quitter la forêt pour boucler son
cycle. Cependant, d'après S. BILY (com. pers.), cette espèce serait indifférente à l'état
de fermeture de son milieu et non pas strictement forestière comme le laisseraient
supposer nos résultats.
INFLUENCE DE L'EXPLOITATION FORESTIERE
En ce qui concerne l’abondance et la richesse spécifique des Coléoptères
étudiés dans les parcelles non exploitées (les stations 2, 7, 9 et 10), on constatera
qu’elles sont au mieux équivalentes (stations 2 et 9) a celle des parcelles exploitées,
ou nettement plus faibles (stations 7 et 10).
On supposait a priori qu’une forêt inexploitée, comportant plus d’arbres vieux
ou morts et davantage d’arbres tombés créant des trouées qu’une forêt exploitée,
serait plus favorable aux insectes du bois. Les parcelles considérées ont été exploitées
jusqu’en 1945. Depuis lors, la structure du peuplement n’a pas sensiblement changé.
Tout au plus y voit-on de nombreux petits arbres secs, ainsi qu'une canopée très
dense, étant donné qu'aucune éclaircie n'y a été effectuée depuis 50 ans et que les
arbres n'ont pas encore atteint le stade sénescent auquel leur feuillage s'éclaircit.
Puisque l’ensoleillement constitue un facteur essentiel à la présence d’une faune de
Coléoptères du bois variée et abondante, on considérera comme logique la faiblesse
des effectifs capturés dans les stations inexploitées.
INFLUENCE DU BOIS MORT
Dans le cas présent, la quantité de bois mort seule ne semble pas constituer un
facteur déterminant l’abondance de la faune étudiée. La comparaison des stations 3 et
4 en témoigne. Cette dernière, pourtant nettement mieux dotée en bois mort, présente
une faune qui n’est guère plus diversifiée que celle de la station 3. En revanche,
lorsque l’offre en bois mort est couplée avec un bon ensoleillement, elle semblerait
constituer un attrait non négligeable, dont il est cependant difficile ici d’estimer
précisément l’importance.
Cela dit, le bois mort des forêts étudiées consiste principalement en branches
mortes au sol et en petits arbres secs sur pied. Il s'agit donc de bois mort récent et
généralement de faible diamètre. Ce bois n'a sans doute pas la même valeur pour les
insectes saprophages et xylophages que de vieux arbres pourvus de nombreuses
branches mortes. Ces derniers représentent en effet pour la faune concernée un milieu
stable et diversifé à long terme.
FAUNISTIQUE ET ÉCOLOGIE
D'un point de vue faunistique, cette étude a permis d’ajouter une espèce à la
faune du canton de Neuchâtel, le Bupreste Agrilus olivicolor. Il se rencontre surtout
en plaine mais remonte les vallées alpines jusque vers 1500 mètres. Seule une localité
(Delémont) a été signalée dans le Jura jusqu’à présent.
On peut également se réjouir de la présence sur le terrain d’étude de trois
espèces de Cérambycides rares figurant sur la liste rouge des espèces menacées
„> —
COLEOPTERES LIÉS AU BOIS 559
TABLEAU 2: espèces capturées et aperçu écologique
(nomenclature selon FREUDE, HARDE & LOHSE 1992, 1994) *
Distribution: M-O = Moyen-Orient, holomédit. = holoméditerranéenne; Etage: col = collinéen, mon =
montagnard, sub = subalpin; Milieu: ffeu = forêt de feuillus, fcon = forêt de conifères, mix = forêt
mixte, tran = clairières et lisières, mou = milieu ouvert, prm = forêt primitive, riv = milieu riverain;
Plante-hôte: feuil. = diverses essences de feuillus, conif. = diverses essences de conifères; Station:
sans autre indication que le numéro de la station, espèce capturée au piège, (chv) = espèce capturée
dans la station par chasse à vue, hs = espèce attrapée par chasse à vue en dehors des stations.
Espèce Auteur Distribution Etage Milieu Plante-hôte Station
BUPRESTIDAE
Agrilus angustulus (111.,1803) eurosiberienne col ffeu/tran Quercus 5
Agrilus biguttatus (F.,1777) européenne, M-O col ffeu/tran Quercus 5
Agrilus laticornis (111.,1803) européenne, M-O col ffeu/tran Quercus 6
Agrilus olivicolor Kiesw.,1857 eurosibérienne col ffeu/tran Corylus,Carpinus 4
Agrilus sulcicollis Lacord.,1835 eurosibérienne col ffeu/tran Quercus 2,5,6
Agrilus viridis (L.,1758) eurosibérienne col-mon ffeu/tran feuil. 5
Anthaxia helvetica Stierl.,1868 oréale col-sub fcon/tran conif. 4,5,6,hs
Anthaxia quadripunctata (L.,1758) oréale mon-sub fcon/tran conif. 5,6,hs
Chrysobothris affinis (F.,1794) eurosiberienne col-mon ffeu/tran feuil. 5,hs
LUCANIDAE
Platycerus caprea (Geer,1774) Europe centrale mon-sub ffeu feuil. 8
Platycerus caraboides (L.,1758) Europe centrale col-mon ffeu feuil. 7,hs
Sinodendron cylindricum (L.,1758) eurosiberienne col-mon ffeu/prm feuil. 9
CERAMBYCIDAE
Alosterna tabacicolor (Geer,1775) paléarctique col-mon ffeu/tran feuil. 1,2,3,4,5,6,7,hs
Anaglyptus mysticus (L.,1758) holomédit. col-mon ffeu/tran feuil. 3,6,8,hs
Callidium aeneum (Geer,1775) boréo-alpine mon-sub fcon conif. 8
Clytus arietis (L.,1758) européenne, M-O col-mon ffeu feuil. 1,2,3,4,5,6,7,hs
Gaurotes virginea (L.,1758) boréo-alpine mon fcon/tran conif. 6,hs
Grammoptera ruficornis (F.,1781) européenne col ffeu/mou feuil. 2,3,hs
Grammoptera ustulata (Schall.,1783) européenne col ffeu/tran feuil. 6
Grammoptera abdominalis (Steph.1831) holomédit. col ffeu/tran Quercus,Castanea 6,hs
Judolia cerambyciformis (Schrk.,1781) holomédit. col-mon ffeu/mou feuil./conif. 1,4,5,6,hs
Leiopus nebulosus (L.,1758) europeenne col-mon ffeu/mix feuil. 3,7,8,9
Leptura dubia Scop.,1763 européenne mon-sub fcon/tran conif. 5,6
Leptura livida F.,1776 eurosibérienne col mou/tran terricole hs
Leptura maculicornis Geer,1775 boréo-alpine mon-sub mou/tran conif/feuil. hs
Leptura rubra L.,1758 paléarctique col-mon fcon/mou conif/feuil. 1,4,5,6,hs
Leptura sanguinolenta L.,1761 boréo-alpine mon-sub tran conif. 5,6,hs
Leptura sexguttata F.,1975 europeenne col-mon ffeu/tran Quercus 2,3,5
Molorchus minor (L.,1758) holarctique col-sub fcon/mix conif. 6,10,hs
Obrium brunneum (F.,1792) européenne, M-O mon-sub fcon/mix conif. 1,5,6,9,hs
Oxymirus cursor (L.,1758) boréo-alpine mon-sub fcon/mix conif/feuil. 6,9
Parmena balteus (L.,1767) méditerranéenne col ffeu/tran feuil. 10,hs
Phymatodes testaceus (L.,1758) européenne, M-O col-mon ffeu feuil. 3,hs
Plagionotus arcuatus (L.,1758) européenne, M-O col ffeu/mix Quercus 5
Pogonocherus hispidulus (Pill.Mitt.,1783) européenne col-mon ffeu/mix feuil. 259
Pogonocherus hispidus (L.,1758) européenne col-mon ffeu/mix feuil. 9 (chv)
Pogonocherus ovatus (Goeze,1777) européenne mon-sub fcon/mix conif. 5,6
Pyrrhidium sanguineum (L.,1758) holomédit. col ffeu Quercus 3 (chv),5
Rhagium bifasciatum ETS européenne col-mon fcon/mix conif./feuil. hs
Rhagium inquisitor (L.,1758) eurosibérienne col-mon fcon/mix conif./feuil. hs
Rhagium mordax (Geer, 1775) eurosibérienne col-mon ffeu/mix feuil./conif. 9,hs
Stenocorus meridianus (L.,1758) eurosibérienne col ffeu feuil. 1235
Strangalia maculata (Poda,1761) européenne, M-O col-mon ffeu/tran feuil. 5,6,hs
Strangalia melanura (L.,1758) eurosiberienne col-mon ffeu feuil./conif. 1,2,4,5,6,8.hs
Tetropium castaneum (L.,1758) paléarctique col-sub fcon conif. 8
Tetropium fuscum (F.,1787) col-sub fcon conif. 8
* Cf. note à la fin de la bibliographie.
eurosibériennne
560 SYLVIE BARBALAT
TABLEAU 3
Stations et espèces ordonnées selon leur position sur le premier axe canonique représentant la
variance expliquée par la variable “présence ou absence de clairière”.
T
Especes / Stations SEION) MST SSL ANS T2 st.9 SE EST SR
Parmena balteus 1 |
Agrilus angustulus 10
Agrilus bigutatus 6
Agrilus laticornis 1
Agrilus viridis 3 Espèces
Anthaxia quadripunctata 9 5
Chrysobothris affinis 9 |
Gaurotes virginea 1 |
Grammoptera abdominalis 1 |
Grammoptera ustulata | 3 de
Leptura dubia 5 2
Leptura sanguinolenta SZ 6
Molorchus minor l | 1
Plagionotus arcuatus lil clairières
Pogonocherus ovatus 1 1
Potosia cuprea 1
Pyrrhidium sanguineum :
Strangalia maculata 24 14
Trichius fasciatus 12 2
Anthaxia helvetica 11 7 I
Leptura rubra 31 18 l I
Agrilus sulcicollis 2 3 1
Judolia cerambyciformis 6 8 1 1
Serica brunnea l 1 l |
Strangalia melanura 973, - 156 9 2 4 3 |
Clytus arietis 16 4 1 1 2 1 1 | Espèces
Oxymirus cursor* l I | ubiquistes
Obrium brunneum 1 3 1 2 2 | plutôt
Anaglyptus mysticus 2 l 1 thermophiles
Alosterna tabacicolor FORD 8 6 28 18 1 6
Pogonocherus hispidulus 2 1 2
Leptura sexguttata 1 1 1
Stenocorus meridianus l l 2
Agrilus olivicolor 2
Grammoptera ruficornis Die ic RSS i
Callidium aeneum 1
Leiopus nebulosus 2 1 1 7 | Espèces de
Phymatodes testaceus* DI Bforets
Platycerus caprea l | fraîches et
Platycerus caraboides 1 | plutôt
Rhagium mordax I l sombres
Sinodendron cylindricum l
Tetropium castaneum 4
Tetropium fuscum 1 |
nombre total d'individus 3 1155 280 23 15 15 9 31 4 19 1554
nombre total d’espèces 3 25 22 TI 8 8 6 9 4 U 44
Remarques: Etant donné le très faible nombre d’especes capturees dans la station 10, sa position sur
l’axe I n’est pas interpretable. Les espèces munies d’une astérisque ont été trouvées dans des milieux
ne correspondant pas à ceux cités dans la littérature.
COLEOPTERES LIÉS AU BOIS 561
d'Allemagne (GEISER 1984) (une telle liste n’existe pas encore pour la Suisse). Il
s’agit de Leptura sexgutatta, (statut 2: très menacé) Pogonocherus ovatus, et
Tetropium fuscum (statut 3: menacé). En Suisse, ces espèces sont peu communes. La
consultation de la base de données du Centre Suisse de Cartographie de la Faune à
Neuchâtel permet d'estimer le statut de ces espèces en Suisse. Si les effectifs de
Leptura sexguttata Sont stables, ceux de Pogonocherus ovatus et Tetropium fuscum
sont en diminution.
Pour ces espèces et d’une manière générale, il est difficile de savoir si le peu
de données jurassiennes correspond à une distribution réelle de l’insecte ou à un
effort de prospection faible comparé à d’autres régions du pays.
D'autres espèces méritent aussi quelque intérêt. Le Lucanide Sinodendron
cylindricum présente selon KOCH (1992) une affinité marquée pour les vieilles forêts
de hêtres (Fagus sylvatica). L'espèce a été capturée dans la station 9 qui comporte
effectivement quelques vieux hêtres. Le Cérambycide Plagionotus arcuatus, sans être
une espèce relique des vieilles forêts, indique selon RIECKEN & BLAB (1989) des
peuplements de chênes d’âge respectable. Il en va de même pour le Bupreste Agrilus
biguttatus (S. BILY, com. pers.). Parmena balteus est un petit Cerambycide. Il est
assez commun dans le bassin lémanique. Cette espèce méditerranéenne est rare en
Suisse en dehors de cette région. On la signale cependant dans plusieurs localités du
pied du Jura entre Yverdon et Bienne.
CONCLUSION
Cette recherche a permis de mettre en évidence certaines préférences écolo-
giques des familles étudiées. Elle représente également un certain apport faunistique à
la connaissance d’une région peu prospectée. Elle infirme aussi, pour les insectes
concernés du moins, une idée souvent répandue: toute exploitation forestière serait
nuisible à l'entomofaune. On a pu le constater, c’est précisément dans les endroits
exploités et particulièrement dans les clairières ouvertes par les forestiers que la
richesse faunistique et l'abondance sont les plus importantes. Cette variété est due à la
clairière elle-même, ainsi qu'à la qualité du peuplement dans lequel elle se situe.
En visitant une forêt primitive (Bialowieza, Pologne), nous avons pu nous
rendre compte qu'elle présentait une canopée relativement clairsemee et comportait de
nombreuses clairières dues à de vieux arbres tombés. Il semble donc qu'à l'origine, les
forêts étaient beaucoup moins sombres qu'actuellement. Cela expliquerait la prédilec-
tion pour les clairières que nous avons constatée chez des insectes réputés forestiers.
La richesse faunistique de la région étudiée est assez bonne si on la compare à
d’autres inventaires de l’entomofaune forestière (HARTMANN & SPRECHER, 1990;
SCHERLER & al. 1989). Il faut cependant relever qu’on n’y a trouvé qu’un seul
individu d'une espèce (le Lucanide Sinodendron cylindricum) caractéristique des
forêts primitives. Ces dernières n’ont pratiquement jamais subi d'exploitation et
présentent de nombreux vieux arbres. Il est certes réjouissant qu'une forêt fortement
marquée par l’influence humaine conserve une faune entomologique variée, mais il
faut toutefois garder en mémoire qu'il s’agit d’especes, pour la plupart fréquentes,
562 SYLVIE BARBALAT
ayant pu s'adapter aux conditions dictées par la sylviculture. D’après les ouvrages
consultés (PALM 1959; RIECKEN & BLAB 1989 et HOLZSCHUH 1984), il existe une série
d’espèces de Coléoptères du bois que l’on peut considérer comme des reliques de la
forêt primitive. Citons parmi elles les Cérambycides Cerambyx cerdo, Plagionotus
detritus, Mesosa curculionoides et Trichoferus pallidus, les Lucanides Aesalus
scarabaeoides et Ceruchus chrysomelinus ou le Bupreste Eurythrea quercus. Ces
espèces sont pour la plupart rarissimes en Suisse. Cette rareté, qui ne semble
d’ailleurs pas récente, est vraisemblablement imputable à la disparition déjà ancienne
des forêts primitives en Suisse. Il est également probable que certaines espèces à
répartition plutôt méridionale telles Cerambyx cerdo ou Eurythrea quercus se
montrent plus sensibles à des modifications de leur milieu lorsqu'elles se trouvent
dans les régions marginales de leur aire de répartition. Etant donné la quasi absence
de vieux arbres dépérissants dans nos forêts, il semble assez peu probable que de
telles espèces aient quelque chance de les recoloniser. On a toutefois pu constater que,
dans les forêts prospectées, un certain nombre d’arbres ayant visiblement dépassé
l’âge où ils sont d'habitude abattus, sont laissés sur pied. Nous ne pouvons
qu’encourager cette tendance favorisant le maintien à long terme d’une entomofaune
diversifiée dans nos forêts.
RÉSUMÉ
Une comparaison de la richesse faunistique et de l’abondance des Coléoptères
liés au bois (Buprestidae, Lucanidae et Cerambycidae) entre des forêts exploitées
actuellement et des forêts inexploitées depuis 1945 a été réalisée. Les pièges suivants
ont été utilisés: le piège-fenêtre, la mini tente Malaise, l'assiette jaune et le piège a
bière. Des 46 espèces récoltées, 34 appartiennent aux Cérambycides, 9 aux
Buprestides et 3 aux Lucanides. Les forêts inexploitées ne présentent pas une faune
plus riche que les forêts exploitées. La faune considérée est très favorablement
influencée par les clairières. Il manque cependant dans les peuplements étudiés des
Coléoptères caractéristiques de vieilles forêts.
ZUSAMMENFASSUNG
Der faunistiche Reichtum wurde mit der Häufigkeit zwischen heutzutage
bewirtschafteten Wäldern und seit 1945 nicht mehr bewirtschafteten Wäldern
verglichen. Dabei wurden die folgenden Fangmethoden benützt: Fensterfalle, kleine
Malaisefalle, Gelbschale und Bierfalle. Unter den 46 gefangenen Arten gehören 34 zu
den Cerambyciden, 9 zu den Buprestiden und 3 zu den Lucaniden. Die Wälder, die
nicht mehr bewirtschaftet werden, weisen keine reichere Totholzkäferfauna
(Buprestidae, Lucanidae und Cerambycidae) als bewirtschaftete Wälder auf. Diese
Fauna findet man bevorzugt auf Lichtungen. Arten, die als Urwaldrelikte gelten
fehlen in den untersuchten Wäldern.
COLEOPTERES LIÉS AU BOIS 563
REMERCIEMENTS
Je tiens a exprimer ma vive reconnaissance au Professeur W. Matthey et au Dr
D. Borcard pour le suivi de ce travail et la relecture du manuscrit, ainsi qu’à MM. M.
Plachta et P. Junod, responsables des arrondissements forestiers concernés par cette
étude, pour leur collaboration. Je remercie également l'expert anonyme de ses
remarques constructives.
BIBLIOGRAPHIE
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mofaune des frondaisons: le piège attractif aérien. Bulletin de la Société Entomologique
Suisse 64: 293-305.
BARBALAT, S. (1995). Efficacité comparée de quelques méthodes de piégeage sur certains
Coléoptères saprophages ou xylophages et influence de l’anthophilie sur le résultat des
captures. Bulletin de la Société neuchâteloise des Sciences Naturelles. 118: 39-52.
BASSET, Y. 1985. Comparaison de quelques méthodes de piégeage de la faune dendrobie.
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CHARRARAS, C. 1972. Les insectes du peuplier, Librairie de la Faculté des sciences, Paris, 372 pp.
FREUDE, H., HARDE, K. W., LOHSE, G. A. 1992. Die Käfer Mitteleuropas Band 13, Goecke &
Evers, Krefeld, 375 pp.
FREUDE, H., HARDE, K. W., LOHSE, G. A. 1994. Die Käfer Mitteleuropas Band 14, Goecke &
Evers, Krefeld, 403 pp.
GEISER, R. 1980. Grundlagen und Massnahmen zum Schutz der einheimischen Käfer.
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1, Greven, 270 pp.
HARTMANN, K. & SPRECHER, E. 1990. Ein Beitrag zur Insektenfauna des Arlesheimer Waldes
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schenden Gesellschaft Baselland 36: 75-124.
HOLZSCHUH, C. 1984. Rote Liste der in Österreich gefährdeten Bockkäfer (Cerambycidae) und
Borkenkäfer (Scolytidae), pp. 127-129. In: Rote Liste gefährdeter Tiere Österreichs
(Gepp J. ed.). Wien, 243 pp.
Kocu, K. 1992. Die Käfer Mitteleuropas, Ökologie, Band 3., Goecke & Evers, Krefeld, 389 pp.
LSPN. 1993. Réserves forestières et protection de la nature. Contributions à la protection de la
nature en Suisse 14. Bäle, 76 pp.
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Opuscula entomologica, Supplementum XVI, Lund, 374 pp.
PAULUS, H. 1980. Einige Vorschläge für Hilfsprogramme unserer gefährdeten Käfer. Natur und
Landschaft 55: 28-32.
RIECKEN, U. & BLAB, J. 1989. Biotope der Tiere in Mitteleuropa - Naturschutz Aktuell 7.
Greven, 123 pp.
SCHAEFFER, L. 1949. Les Buprestidae de France, Miscellanea entomologica, Paris, 511 pp.
SCHERLER, P., SEKALY, V. & TOUMAYEFF, G. 1989. Coléoptères de la réserve du Bois de Chênes
dans la région de Ferreyres-Moiry. Bulletin romand d’entomologie 7 (1): 11-29.
SPEIGTH, M. 1989. Les invertébrés saproxyliques et leur protection, Collection Sauvergarde de
la nature 42, Strasbourg, 77 pp.
564 SYLVIE BARBALAT
SPRECHER-UEBERSAX, E. 1989. Die Bedeutung des Totholzanteils für die xylobionte Cole-
opterenfauna eines Buchenwaldes. Diplomarbeit, Basel, 143 pp.
TER BRAAK, C.J.F. 1986. Canonical correspondence analysis. A new eigenvector technique for
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Press, Amsterdam.
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vironment relationships. Vegetatio 75: 159-160.
NOTE
Nous n’avons pas pu tenir compte dans les tableaux 2 et 3 du travail de BENSE (1995),
“Longhorn Beetles: Illustrated key to the Cerambycidae and Vesperidae of Europe”, Margraf,
Weikerscheim, 512 pp., qui nous est parvenu en cours d’impression.
REVUE SUISSE DE ZOOLOGIE
Tome 103 — Fascicule 2
FRISCH, Johannes. Revision westmediterraner Scopaeus - Arten (Coleop-
tera, Staphylinidae: Paederinae) und Beschreibung einer neuen Art
aus Südspanien und Marokko. 2. Beitrag zur Kenntnis der Gattung
NCOPACUSARNICNSOM IR NP ICAO
LOURENÇO, WILSON R. Microcharmus hauseri, nouvelle espèce de Scor-
piondeMadasascan(Scorpiones Buthidae) e cr.
GORBUNOV, OLEG G. & Yutaka ARITA. New and little-known Oriental
Melittia Hübner (Lepidoptera, Sesiidae), from the collection of
Miurscumedshistomematurcile Geneve sss u tee EE
GEIGER, Daniel. Haliotids in the Red Sea, with neotype designation for
Haliotis unilateralis Lamarck, 1822 (Gastropoda: Prosobranchia). . .
PAGÉS, Jean. Un Evalljapygidae (Diplura) canadien: Evalljapyx saundersi
nu spDieellunatal@enayensia ONE Ee ER ao elo cae
CONDÉ, Bruno & Camille BARETH. Une évaluation de Stygiocampa, sous-
genre troglomorphe de Plusiocampa (Diplura Campodeidae), avec la
description d’une nouvelle espèce de Serbie orientale..............
BOURGAT, Robert, Cyril ROURE & Sim-Dozou KuLo. Nouvelles données sur
les Trématodes d’Amphibiens d’Afrique Occidentale. Description
dyklaematoloechustaubriaen- spam... OE
PACE, Roberto. Aleocharinae della Colombia e dell’Ecuador: Parte I (Cole-
opteraStapNy.limidae). reno cate
STEINER, Werner A. Distribution of entomopathogenic nematodes in the
SUIS SHA SE ON era ee
RIEDEL, Alexander & Klaus SCHÖNITZER. Revision of the soil-weevil genus
Guineobius Osella from New Guinea (Coleoptera, Curculionoidea,
Entiminaes@eleuthetind) sr. tree Ree cao cet as ae ae
Cuccoporo, Giulio. Revision of the Japanese rove-beetles of the genus
Megarthrus (Coleoptera, Staphylinidae, Proteininae)..............
NAYROLLES, Pierre. Contribution to the knowledge of European Bourle-
tiellidae (Collembola, Symphypleona). IH. Description of some spe-
cies of Heterosminthurus and Deuterosminthurus. ...............
BARBALAT, Sylvie. Influence de l’exploitation forestière sur trois familles
de coléopteres liés au bois dans les Gorges de l’Areuse............
301-318
319-322
323-338
339-354
355-367
369-381
383-394
395-437
439-452
453-473
475-524
525-551
553-564
REVUE SUISSE DE ZOOLOGIE
Volume 103 — Number 2
FRISCH, Johannes. Revision of Western Mediterranean Scopaeus-species
with description of a new species from Southern Spain and Maroc. . .
LOURENÇO, WILSON R. Microcharmus hauseri, new species of scorpion
iomeviadasascam(Scorpiones, Buthidae). TER
GORBUNOV, OLEG G. & Yutaka ARITA. New and little-known Oriental
Melittia Hübner (Lepidoptera, Sesiidae), from the collection of
Muséumidhistorematurelle Genève. Be ann js a eee
GEIGER, Daniel. Haliotids in the Red Sea, with neotype designation for
Haliotis unilateralis Lamarck, 1822 (Gastropoda: Prosobranchia). . .
PAGÉS, Jean. A Canadian Evallijapygidae (Diplura): Evalljapyx saundersi
MaspaDicelurata Genavensia LON =. tay. EEE
CONDE, Bruno & Camille BARETH. An appraisal of Stygiocampa, a troglo-
morphic subgenus of Plusiocampa (Diplura Campodeidae), with des-
cription Oa new species romieastern Serbia. o Sia ee
BOURGAT, Robert, Cyril ROURE & Sim-Dozou KuLo. New data on the
Trematodes of Amphibians from Western Africa. Description of
klaematoloechustaubniaem “sp? OO et I ee
PACE, Roberto. Aleocharinae from Colombia and Ecuador: Part I (Coleop-
terasStaphylinidae). vr mn. ee NE e
STEINER, Werner A. Distribution of entomopathogenic nematodes in the
SWISS END SR RER N N ee
RIEDEL, Alexander & Klaus SCHÖNITZER. Revision of the soil-weevil genus
Guineobius Osella from New Guinea (Coleoptera, Curculionoidea,
Entiminaes Celeuthe tin) 2.10. WAVE e RIO: SAL eh RARE
CuccoDORO, Giulio. Revision of the Japanese rove-beetles of the genus
Megarthrus (Coleoptera, Staphylinidae, Proteininae)..............
NAYROLLES, Pierre. Contribution to the knowledge of European Bourle-
tiellidae (Collembola, Symphypleona). IT. Description of some spe-
cies of Heterosminthurus and Deuterosminthurus. ...............
BARBALAT, Sylvie. Influence of forest management on three wood eating
Beetles families in the Areuse Gorges (Canton of Neuchätel, Switzer-
Indexed in CURRENT CONTENTS
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323
589
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369
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