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Full text of "Revue suisse de zoologie"

1 
ANNALES 



tome 105 
fascicule 3 
1998 



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de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

et du 

MUSÉUM D'HISTOIRE NATURELLE ^^ 

de la Ville de Genève ^^J 

O 

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LU 

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Jël GENEVE SEPTEMBRE 1998 ISSN 0035 -418 X HH 



REVUE SUISSE DE ZOOLOGIE 



TOME 105 — FASCICULE 3 

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 

CHARLES LIENHARD 
Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Président: Ivan Löbl — Muséum de Genève 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des Instituts de zoologie des universités 
suisses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, écologie, éthologie, morphologie et anatomie comparée, 
physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.geneva-city.ch:80/musinfo/mhng/publications/revues.htm 



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 

MUSÉUM D'HISTOIRE NATURELLE 

de la Ville de Genève 



tome 105 
fascicule 3 
1998 




o 

o 



]§[ 



GENEVE SEPTEMBRE 1998 ISSN 0035 - 418 X 



O 

o 

N 

LU 

û 

LU 

D 

LU 

> 

LU 

ce 



REVUE SUISSE DE ZOOLOGIE 



TOME 105 — FASCICULE 3 

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 

CHARLES LIENHARD 
Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Président: Ivan Löbl — Muséum de Genève 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des Instituts de zoologie des universités 

suisses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, écologie, éthologie, morphologie et anatomie comparée, 
physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.geneva-city.ch:80/musinfo/mhng/publications/revues.htm 



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-121 1 Genève 6, Suisse 



Revue suisse de Zoologie 105 (3): 465-485; septembre 1998 



Evaluation de l'entretien des prairies sèches du plateau occidental 
suisse par le biais de leurs peuplements arachnologiques 
(Arachnida: Araneae)* 

Stefano POZZI 1 , Yves GONSETH 2 & Ambros HÄNGGI 3 

1 Muséum d'histoire naturelle, Case postale 6434, CH-121 1 Genève 6. 

2 CSCF, Terreaux 14, CH-2000 Neuchâtel. 

3 Naturhistorisches Museum, Ausustinergasse 2, CH-4001 Basel. 



Evaluation of dry grassland management on the Swiss occidental 
plateau using spider communities (Arachnida: Araneae). - Dry grass- 
land is a seriously endangered habitat in Switzerland. Within the frame- 
work of the Swiss Dry Grassland Inventory, the question of whether 
current management techniques meet the protection requirements was 
asked. The current research, using spiders as representatives of many 
epigeic groups, was expected to show what influence the management has 
on the small animal fauna. For that reason, a valuation method was 
developed which is based on the assessment of all the captured species at 
any given site, and does not only consider a few indicator species. It takes 
into account habitat fidelity and the rarity of individual species. The study 
showed that very extensive use (mowing in autumn, if possible not in 
every year) is required for the conservation of the most valuable dry grass- 
land possible. Grazing, either by sheep or cattle, was discovered to be less 
favourable. Rotation of used and fallow sections of a site is recommended. 
In areas with different uses in consecutive years, it has been shown that 
spiders react quickly to changes in use and are therefore good bioindi- 
cators. Spiders should therefore be used in the future in case studies for the 
description of the actual condition or development of specific habitat types, 
together with other parameters such as vegetation. 

Key-words: Araneae - ecology - habitat quality evaluation - dry grassland 
- conservation management - Switzerland. 

INTRODUCTION 

Les prairies sèches constituent les milieux herbacés les plus riches en espèces 
végétales et animales de Suisse (Antognoli et al. 1995). Elles se caractérisent par des 
communautés végétales qui poussent sur des terrains pauvres en substances nutritives et 



*Cet article fait partie de la thèse du premier auteur à l'Université de Genève sur 
l'étude des araignées des prairies sèches en tant que bioindicateur de la qualité du milieu, projet 
753-VA-l 1 16/00 de l'Office fédéral de l'environnement, des forêts et du paysage (OFEFP). 

Manuscrit accepté le 19.02.1998. 



466 s - POZZI, Y. GONSETH & A. HÄNGGI 

qui souffrent d'un manque d'eau périodique. Ces écosystèmes sont semi-naturels, ils 
ont été créés et maintenus par les activités agricoles traditionnelles non mécanisées, 
essentiellement le pâturage et la fauche. 

En Suisse, la disparition progressive des prairies sèches a encouragé l'Office 
fédéral de l'environnement à actualiser l'inventaire de ces milieux. Dans cette logique, 
les responsables du projet ont reconnu la nécessité d'étudier l'impact de leur entretien 
sur les communautés animales épiédaphiques. Parmi-celles-ci, les araignées sont 
connues pour fournir un bon reflet de l'état de leur habitat (Clausen 1986; Maelfait et 
al. 1989; Bauchhenss 1990; Platen 1993; Gonseth & Mulhauser 1995; Pozzi 1996). 
Présent dans tous les biotopes terrestres, ce groupe d'arthropodes très abondants 
comprend beaucoup d'espèces dont les peuplements sont révélateurs de conditions 
écologiques précises (Hänggi et al. 1995; Schulz & Finch 1996). 

Grâce aux méthodes d'ordination canonique partielle, Pozzi & Borcard 
(soumis) ont montré que les peuplements d'araignées des prairies sèches sont influ- 
encés, d'une part, par leur environnement naturel et d'autre part, par leur entretien par 
l'homme. Gonseth (1985), LOrtscher et al. (1994), Antognoli et al. (1995), Baur 
et al. (1996) ont utilisé les araignées pour évaluer la qualité de prairies sèches et l'effet 
de différents modes d'exploitation. Ces travaux montrent que l'évolution de la faune 
arachnologique des prairies sèches est intimement liée à l'évolution de leur végétation 
et par conséquent au traitement qu'elles subissent, mais ne proposent pas de méthode 
précise pour évaluer la qualité d'une station. 

La présentation détaillée d'une méthode d'évaluation de la qualité des prairies 
sèches par le biais des araignées, basée sur les travaux de Hänggi (1987, 1990), est la 
principale originalité de notre travail dont les buts étaient les suivants: 

comparer les effets de différents types d'entretien des prairies sèches sur leur 

peuplement arachnologique; 

définir des mesures susceptibles d'améliorer leur qualité respective pour la 

faune; 

souligner l'intérêt d'utiliser la faune, et plus particulièrement les araignées, 
comme outil supplémentaire d'appréciation des mesures de gestions adoptées 
pour assurer leur conservation. 

MATÉRIEL ET MÉTHODES 

Stations 

En 1995 et 1996, 40 stations ont été étudiées dans les régions de basse altitude 
(355 à 800 m) du pied du Jura vaudois, dans le canton de Genève, et dans le pays de 
Gex en France. Ces stations sont décrites par Pozzi (1997). D'après la typologie des 
milieux de Suisse (Galland & Gonseth 1990), les prairies étudiées peuvent être 
attribuées aux unités de végétation suivantes: a) prairies très sèches, b) prairies sèches 
typiques, c) prairies sèches légèrement amendées. Elles se caractérisent toutes par la 
dominance du brome dressé (Bromus erectus). Afin de ne pas compliquer l'inter- 
prétation des résultats par des effets de lisière (Heublein 1983; Hänggi 1993a; 



ÉVALUATION DE L ENTRETIEN DES PRAIRIES SECHES 467 

Bedford & Uscher 1994), le choix des surfaces s'est porté principalement sur des 
terrains homogènes, aussi bien du point de vue de l'entretien que du point de vue du 
type de végétation. En ce qui concerne les modes d'entretien, les types suivants ont été 
sélectionnés: a) pâturage bovin (6 stations), b) pâturage ovin (6 stations), c) fauchage 
précoce (mi-juin) (8 stations), d) fauchage tardif (automnal) (7 stations), e) entretien 
irrégulier (4 stations), f) station abandonnée (9 stations). Il faut souligner que la plupart 
des entretiens sont extensifs. Si l'exploitation devient intensive (surpâturage, fauchage 
trop précoce, engraissement) les prairies sèches se transforment rapidement en prairies 
grasses (Antognoli étal. 1995). 

RÉCOLTES DES ARAIGNÉES 

La récolte des données arachnologiques a été effectuée avec des pièges au sol 
(Barber): gobelets blancs en polypropylene de 7 cm de haut et 7 cm de diamètre, 
remplis au tiers d'un liquide conservateur (formaldehyde 4%). Les pièges (trois par 
station) étaient disposés en triangle au centre de la parcelle. Ils ont été vidés chaque 
quinzaine, d'avril à novembre (protocole selon Hänggi 1989). Le matériel sera déposé 
au Muséum d'histoire naturelle de Genève et au Naturhistorisches Museum de Bale. 

Le piège «Barber» est un piège d'activité particulièrement efficace pour l'étude 
de la macrofaune la plus mobile du sol, de la litière ou de la strate herbacée. Il est 
inadapté à la capture des espèces sédentaires (araignées à toile par exemple) dont 
l'activité au sol est, par définition, très limitée ou pour les espèces dont l'essentiel de 
l'activité a lieu sur les buissons, sur les troncs, les branches ou dans la couronne des 
arbres. Les résultats qualitatifs et semi-quantitatifs obtenus par cette méthode de 
piégeage ne concernent donc qu'une partie seulement des peuplements des milieux 
étudiés et ne permettent pas d'obtenir un reflet fidèle de la diversité réelle d'un milieu à 
forte structure tridimentionnelle. Par contre, les données rassemblées avec cette 
méthode sont extrêmement efficaces pour effectuer une comparaison des diverses 
stations prairiales choisies dans le cadre de cette étude. 

Liste des abréviations 

Les abréviations suivantes sont utilisées dans l'évaluation des stations, les 
résultats et la comparaison des stations par type d'entretien: 

AB stations abandonnées 

CLpEU classes de pourcentage des espèces euryèces 

CLVM 1 et CLVM2 classes des indices VM 1 et VM2 

CLVST classes de valeur stationnelle globale (VST) 

CLVSTH classes de valeur stationnelle théorique (VSTH) 

El stations entretenues irrégulièrement 

F indice de fidélité des espèces 

FP stations fauchées précocement 

FT stations fauchées tardivement 

HVL milieu de haute valeur 

IVL milieu intéressant de valeur limitée 

NIND nombre d'individus 

NOS nombre d'observations en Suisse (versant nord des Alpes uniquement) 



468 S. POZZI, Y. GONSETH & A. HÄNGGI 



NSP nombre d'espèces 

PB stations pâturées par des bovins 

pEU pourcentage des espèces euryèces 

PO stations pâturées par des ovins 

R indice de rareté des espèces 

st. station 

SUM somme des indices VM1 et VM2 

SVP milieu sans valeur particulière 

THVL milieu de très haute valeur 

UGBN charge en bétail, nombre UGB pour 100 jours de pâture 

V valorisation: SVP, IVL, VAL, HVL et THVL 

VAL milieu de valeur 

VM indices VM1 et VM2 

VM1 indice de valeur stationnelle VM1 = CLpEU * CLVST 

VM2 indice de valeur stationnelle VM2 = CLpEU * CLVSTH 

VSP valeur spécifique VSP = R * F 

VST valeur stationnelle globale VST est la somme des VSP d'une station 



EVALUATION DES STATIONS 

Plusieurs méthodes ont été proposées pour l'évaluation d'un milieu du point de 
vue de la protection de la nature (littérature compilée dans Marti & Stutz 1993). Elles 
présentent différents objectifs: évaluation qualitative d'un site, évaluation des effets des 
mesures de gestion, évaluation à différents niveaux de perception de l'espace: paysage, 
milieu complexe ou station. En Allemagne, plusieurs auteurs ont proposé des méthodes 
pour des évaluations plutôt globales en considérant différents facteurs abiotiques et 
divers groupes biologiques (Plachter 1992, 1994; Holstein 1995; Beinlich et al. 
1995). 

La méthode utilisée ici se base sur les travaux de Hänggi (1987) et sur leurs 
développements ultérieurs (Hänggi 1990). Elle n'est pas basée sur l'ensemble de la 
faune mais est au contraire focalisée sur les araignées. Cette approche est fondée sur 
«l'hypothèse» qu'une valorisation «globale» n'est pas réaliste, car ce qui semble béné- 
fique pour un groupe (lépidoptères par exemple) ne l'est pas forcément pour un autre 
groupe (araignées par exemple) puisque leur biologie et leurs exigences écologiques 
sont totalement différentes. D'autre part, cette méthode prend en considération toutes 
les espèces «piégées» dans une station. Nous pensons en effet, que la valeur d'une 
station ne peut être définie seulement à partir de quelques espèces indicatrices: la 
présence de certaines espèces de faible valeur (espèces typiques des stations entretenues 
de manière très intensive) peut être un indice précoce de changement des conditions du 
milieu même si toutes les espèces indicatrices de bonne qualité sont encore présentes. 

Cette méthode, qui se fonde sur l'étude du peuplement arachnologique d'une 
station, tient compte de paramètres importants pour l'évaluation de sa qualité: la rareté 
des espèces capturées et leur fidélité au biotope (Fig. 1). Contrairement aux indices de 
diversité, qui ne tiennent compte que du nombre d'espèces différentes capturées dans 
une station et de la distribution d'abondance de ces espèces, cette méthode privilégie la 
spécificité des peuplements par rapport aux habitats présents. Ainsi, elle permet de ne 



ÉVALUATION DE L'ENTRETIEN DES PRAIRIES SECHES 469 

pas sous-estimer la qualité des biotopes très homogènes renfermant peu d'espèces mais 
dont les liens avec les conditions du milieu sont très étroits (espèces sténoèces) et 
parallèlement de ne pas surestimer les stations riches en espèces ubiquistes (euryèces). 
Par exemple, une roselière renferme peu d'espèces mais de valeur spécifique élevée 
tandis que certains milieux artificiels bien structurés ont un grand nombre d'espèces 
triviales. 

Cette méthode a été développée afin de déterminer la valeur de chaque station et 
de faciliter la comparaison des résultats obtenus. Pour cela, nous avons attribué deux 
indices distincts aux différentes espèces capturées. Ces indices sont les suivants: 

Rareté (R) 

La notion de rareté est intimement liée à la répartition des espèces. Cet indice, 
qui oscille entre 1 et 6, est basé sur les connaissances faunistiques actuelles provenant 
de nombreuses publications (Maurer & Hänggi 1990; Hänggi 1993b; Baur et al. 
1996; Pozzi 1996). Certaines modifications ont donc été apportées aux indices retenus 
par Hänggi (1990). Ces modifications ont en outre entraîné une révision des limites des 
classes préalablement établies. 

classes rareté NOS %N % NSP 



1 


espèce banale 


NOS > 30 


17.7 


9.6 


2 


espèce très commune 


30 > NOS > 20 


20.0 


12.6 


3 


espèce commune 


20 > NOS > 12 


15.8 


18.4 


4 


espèce peu commune 


12 > NOS > 6 


17.2 


21.8 


5 


espèce rare 


6 > NOS > 2 


16.3 


25.4 


6 


espèce très rare 


2>NOS 


13.0 


12.1 



NOS Nombre d'observations en Suisse (versant nord des Alpes uniquement) 

%N Pourcentage du nombre d'espèces recensées pour l'étude des prairies sèches 

%NSP Pourcentage du nombre d'espèce total d'après la littérature 

limites des classes x 2 + x avec x = 1, 2, 3, 4 et 5 



Fidélité (F) 

La fidélité d'une espèce à un habitat précis livre de précieuses indications sur 
ses exigences écologiques. Cet indice, qui oscille entre 1 et 6, est principalement basé 
sur les connaissances écologiques actuelles (Maurer & Hänggi 1990 et Hänggi et al. 
1995) et sur les connaissances que nous avons acquises pour les milieux du nord des 
Alpes en Suisse. Cette remarque sous-entend que la valeur de l'indice attribué à une 
espèce donnée ne peut être reprise sans autre pour l'évaluation de stations appartenant à 
une autre région. De manière générale, un indice de faible valeur est attribué à une 
espèce peu exigeante fréquentant différents milieux de structure peu précise; un indice 
de valeur élevée est attribué à une espèce liée à un milieu de structure précise. La 
signification de l'indice attribué à chaque espèce est la suivante: 



470 



S. POZZI, Y. GONSETH & A. HÄNGGI 



CRITERES 



TRIVIALITE 



RARETE 



STATUT 



Expression des 
différents critères 



pourcentage des 

espèces euryèces 

pEU 



Nb de stations connues 

au nord des alpes 

R 



Fidélité au biotope 
F 



Standardisation des 
critères 



Valeur spécifique 
VSP = R * F 



Valeur stationnelle globale 
VST = S VSP 



Valeur stationnelle 

théorique 
VSTH = VST/NSP 



Classes de % des espèces 

euryèces 

CLpEU 



Classes de valeur 

stationnelle globale 

CLVST 



Classes de valeur 

stationnelle théorique 

CLVSTH 



. j 



Estimation de la valeur 
de chaque station 



VM1 
CLpEU * CLVST 



VM2 
CLpEU * CLVSTH 



Classification 
CLVM1 



SUM = 


VM1 


t-VM2 


SVP 


SUM 


=<8 


IVL: 


SUM = 


=<18 


VAL: 


SUM 


=<32 


HVL: 


SUM 


=<50 


THVL 


:SUM 


=<72 



Classification 
CLVM2 



Fig. 1. Méthode d'évaluation des stations. SVP= sans valeur particulière; IVL= intéressant, de 
valeur limitée; VAL= de valeur; HVL= de haute valeur; THVL= de très haute valeur. 



ÉVALUATION DE L'ENTRETIEN DES PRAIRIES SECHES 47 ] 

1: espèce très peu exigeante et/ou tolérant les milieux artificiels - euryèce (sans 

exigence), ubiquiste - ex: forêts et milieux ouverts, secs et humides, zones très 

artificielles 
2: espèce peu exigeante, présente dans plusieurs milieux de structure différente 

mais absente des milieux très artificiels - euryèce - ex: forêts et milieux ouverts, 

secs et humides 
3: espèce peu exigeante, présente dans différents milieux de structure «particu- 
lière» - mésoèce, sans exigences particulières - ex: milieux ouverts, secs et 

humides 
4: expèce exigeante, liée à quelques types de milieux précis - mésoèce (avec 

exigences) - ex: milieux ouverts, secs 
5: espèce exigeante, liée à un type de milieu bien précis - sténoèce modérée - ex: 

prairies sèches 
6: espèce très exigeante, liée à des conditions bien particulières d'un type de milieu 

bien précis - sténoèce stricte - ex: prairies sèches ouvertes à rocaille (sur sol peu 

profond) 

Estimation de la valeur d'une station 

Le produit des deux indices retenus (R * F) donne la «valeur» de chaque espèce 
(VSP = valeur spécifique). La valeur stationnelle globale (VST) est calculée en effec- 
tuant la somme des valeurs des espèces qui y ont été observées. Cette valeur transitoire 
peut ensuite être reprise directement comme une mesure d'évaluation de la qualité 
d'une station. Toutefois, VST favorisera les milieux dont le nombre d'espèces est élevé 
(milieux à forte structure tridimensionnelle par exemple) au détriment des milieux plus 
homogènes. Ainsi, on a choisi une méthode qui permet de reprendre les deux aspects 
les plus importants pour une évaluation (diversité et spécificité) représentées par les 
deux indices VM1 et VM2 (Fig. 1). 

Comparaison des indices VM 1 et VM2 

Les indices VM1 et VM2 expriment des aspects différents de la qualité d'une 
station donnée. VM1 favorise les stations ayant une haute diversité faunique (milieux 
mosaïques, écotones); VM2 favorise plutôt les milieux abritant une faune particulière 
(milieux extrêmes: homogènes, très stables). Ces deux indices sont donc complé- 
mentaires (et non contradictoires). De manière générale si leur valeur respective est 
élevée, la qualité de la station concernée n'en est que plus évidente. En outre, les écarts 
importants qui les séparent parfois soulignent particulièrement bien la présence de 
milieux extrêmes dans une étude donnée (par exemple: une roselière aura VM2 très 
haut et VM1 bas; tandis qu'un milieu mosaïque aura VM2 bas et VM1 très haut). 

Afin d'intégrer ces deux aspects de notre méthode d'évaluation dans la hiérar- 
chisation définitive des stations, nous avons effectué la somme de ces deux indices 
(SUM = VM1+VM2) et réparti les résultats obtenus dans les catégories SVP, IVL, 
VAL, HVL et THVL en tenant compte des limites de classe présentées ci-dessous: 



472 S. POZZI, Y. GONSETH & A. HÄNGGI 

SVP: milieu sans valeur particulière SUM < 8 

IVL: milieu intéressant de valeur limitée SUM < 18 

VAL: milieu de valeur SUM < 32 

HVL: milieu de haute valeur SUM < 50 

THVL: milieu de très haute valeur SUM < 72 

limites des catégories: x 2 + x 2 avec x = 2, 3, 4, 5, 6 

Le recours aux variables suivantes permet d'obtenir les indices de valeur stationnelle 
VMletVM2: 

— pourcentage d'espèces euryèces (pEU) exprimé en classes de pourcentage 
d'espèces euryèces (CLpEU) 

valeur stationnelle globale (VST, soit somme de toutes les valeurs spécifiques), 
exprimée en classes de valeur stationnelle globale (CLVST) 

— valeur stationnelle théorique (VSTH, soit VST divisé par le nombre d'espèces 
(NSP)), exprimée en classes de valeur stationnelle théorique (CLVSTH). 

Hänggi (1990) répartissait les indices suivants dans 4 classes différentes. Compte tenu 
de la bonne qualité générale des milieux que nous avons choisis et de notre objectif 
final (proposition de mesures de gestion susceptibles d'améliorer la qualité de toutes les 
stations étudiées), il nous a toutefois semblé nécessaire de les répartir en 6 classes afin 
de mieux distinguer les stations de valeur. 



Classes de pourcentage des espèces euryèces (CLPEU) 

Le pourcentage des espèces euryèces (pEU) est une variable importante pour 
l'évaluation de la qualité d'une station (Fig. 1). Plus ce pourcentage est élevé, plus le 
milieu est banal, ou en d'autres termes, plus pEU est faible, plus le milieu est peuplé 
d'araignées exigeantes (Hänggi 1987). Sous le terme d'euryèces sont comprises les 
espèces appartenant à la classe de fidélité 1 . Les limites de classe sont les suivantes: 

1 : pEU > 30% 1 : 25% < pEU < 30% 3: 20% < pEU < 25% 

4: 15%<pEU<20% 5: 10% < pEU < 15% 6: pEU < 10% 



Classes de valeur stationnelle globale (CLVST) 

Les limites de classe de valeur stationnelle globale ont été déterminées sur la 
base d'une station théorique présentant un nombre moyen de 45 espèces (moyenne des 
stations étudiées) dont la valeur spécifique serait égale à 2.25 (F=R=1.5 VSP=2.25 
VST=101.25), à 4 (F=R=2 VSP=4 VST=180), à 6.25 (F=R=2.5 VSP=6.25 VST= 
281.25), à 9 (F=R=3 VSP=9 VST=405) et à 12.25 (F=R=3.5 VSP=12.25 VST=551.25) 

1: VST < 101.25 2: 101.25 < VST < 180 3: 180 < VST < 281.25 

4: 281.25 < VST < 405 5: 405 < VST < 551.25 6: VST > 551.25 



ÉVALUATION DE L'ENTRETIEN DES PRAIRIES SECHES 473 

Classes de valeur stationnelle théorique (CLVSTH) 

Les limites des classes de valeur stationnelle théorique proviennent des valeurs 
spécifiques théoriques (2.25 pour F=R=1.5; 4 pour F=R=2; 6.25 pour F=R=2.5; 9 pour 
F=R=3; 12.25 pour F=R=3.5) 

1 : VSTH < 2.25 2: 2.25 < VSTH < 4 3: 4 < VSTH < 6.25 

4: 6.25 < VSTH < 9 5: 9 < VSTH < 12.25 6: VSTH > 12.25 

Classes des indices VM1 et VM2 (CLVM1 et CLVM2) 

1:VM=1 2:1<VM<4 3:4<VM<9 

4:9<VM<16 5:16<VM<25 6: 25 < VM < 36 

limites des classes: x 2 avec x = 1, 2, 3, 4, 5, 6 

Le choix des facteurs d'estimation des milieux et les notes attribuées aux diffé- 
rentes espèces recensées ont été fixées pour cette étude précise. Ils sont donc 
susceptibles de subir certaines modifications en fonction de l'évolution de nos connais- 
sances faunistiques et écologiques. Toutefois, dans le cadre de ce travail, ces variations 
potentielles sont tamponnées par le grand nombre de sites inventoriés et d'espèces 
recensées par station. De plus, les araignées des milieux ouverts sont assez bien 
connues (Thaler 1985; Gonseth 1985; Delarze 1987; Bauchhenss 1990; Maurer & 
Hänggi 1990; Hänggi 1992; Lörtscher et al. 1994; Hänggi et al. 1995, 1996; Baur 
étal. 1996). 

RÉSULTATS 

Au total, 22057 individus adultes appartenant à 215 espèces ont été récoltés avec 
les pièges Barber. Les résultats bruts (nombre d'individus de chaque espèce par station) 
sont publiés ailleurs (Pozzi 1997). Pozzi & Hänggi (1998) présentent les principaux 
résultats taxonomiques et faunistiques. Les analyses présentées dans le Tab. 1 ont été 
faites à partir du statut de rareté et de fidélité de chaque espèce (Tab. 2). 

La Fig. 2 présente les résultats des évaluations des 40 stations en fonction des 
différents types d'entretien. Les modifications de la méthode Hänggi (1990) apportent 
une plus grande clarté dans la répartition des stations de valeur (partie droite de la 
figure) mais ne modifient pas fondamentalement leur classement respectif (hiérarchi- 
sation). Cette amélioration de la méthode permet une meilleure évaluation des sites et 
de leur entretien. 

Premièrement, nous constatons une proportion importante de stations de 
«valeur»: plus des 3/4 des stations étudiées appartiennent aux catégories VAL, HVL et 
THVL, seules quelques stations appartiennent à la catégorie SVP (Fig. 2). Cette 
répartition très inégale des stations dans les 5 catégories susmentionnées trouve son 
origine dans leur choix initial. Il n'a pas été réalisé au hasard, mais bien au contraire en 
sélectionnant essentiellement des prairies sèches c'est-à-dire des stations qui à priori 
étaient de valeur. 



474 



S. POZZI, Y. GONSETH & A. HÄNGGI 



Tab. 1. 


Tableau des indices des différentes stations 


regroupées par type d'entretien. 








st4 


stl3 


stl7 


st43 


% El 


stl 


stll 


stl5 


st27 


st33 


st34 


st36 


st45 


st49 


XAB 


NIND 




1137 


408 


533 


521 


650 


540 


562 


515 


672 


273 


272 


544 


537 


622 


504 


NSP 




44 


43 


48 


54 


47.3 


49 


45 


50 


52 


38 


51 


46 


43 


48 


46.9 


VST 




379 


411 


375 


432 


399 


524 


499 


529 


573 


402 


484 


529 


521 


506 


507 


VSTH 




8.6 


9.6 


7.8 


8 


8.5 


10.7 


11.1 


10.6 


11 


10.6 


9.5 


11.5 


12.1 


10.5 


10.9 


pEU 




27.3 


16.3 


18.8 


20.4 


20.7 


12.3 


13.3 


18 


7.7 


10.5 


15.7 


15.2 


11.6 


14.6 


13.2 


CLpEU 




2 


4 


4 


3 


3.3 


5 


5 


4 


6 


5 


4 


4 


5 


5 


4.8 


CL VST 




4 


5 


4 


5 


4.5 


5 


5 


5 


6 


4 


5 


5 


5 


5 


5 


CLVSTH 




4 


5 


4 


4 


4.3 


5 


5 


5 


5 


5 


5 


5 


5 


5 


5 


VM1 




8 


20 


16 


15 


14.8 


25 


25 


20 


36 


20 


20 


20 


25 


25 


24 


CLVM1 




3 


5 


4 


4 


4 


5 


5 


5 


6 


5 


5 


5 


5 


5 


5.1 


VM2 




8 


20 


16 


12 


14 


25 


25 


20 


30 


25 


20 


20 


25 


25 


23.9 


CLVM2 




3 


5 


4 


4 


4 


5 


5 


5 


6 


5 


5 


5 


5 


5 


5.1 


SUM 




16 


40 


32 


27 


28.8 


50 


50 


40 


66 


45 


40 


40 


50 


50 


47.9 


V 




IVL 


HVL 


VAL 


VAL 


VAL 


HVL 


HVL 


HVL 


THVL 


HVL 


HVL 


HVL 


HVL 


HVL 


HVL 




st2 


st3 


stl2 


st42 


st44 


st46 


st47 


KFT 




st8 


st24 


st25 


st26 


st32 


st48 


KPO 


NIND 


385 


1106 


483 


629 


376 


512 


772 


609 




371 


389 


1103 


604 


424 


321 


535 


NSP 


46 


51 


39 


54 


55 


53 


45 


49 




50 


50 


49 


41 


39 


58 


47.8 


VST 


613 


540 


352 


499 


508 


621 


436 


510 




464 


570 


453 


403 


207 


441 


423 


VSTH 


13.3 


10.6 


9 


9.2 


9.2 


11.7 


9.7 


10.4 




9.3 


11.4 


9.2 


9.8 


5.3 


7.6 


8.8 


pEU 


15.2 


15.7 


23.2 


22.2 


16.4 


15.1 


15.6 


17.6 




22 


22 


26.5 


19.5 


41 


24.1 


25.9 


CLpEU 


4 


4 


3 


3 


4 


4 


4 


3.7 




3 


3 


2 


4 


1 


3 


2.7 


CL VST 


6 


5 


4 


5 


5 


6 


5 


5.1 




5 


6 


5 


4 


3 


5 


4.7 


CLVSTH 


6 


5 


5 


5 


5 


5 


5 


5.1 




5 


5 


5 


5 


3 


4 


4.5 


VM1 


24 


20 


12 


15 


20 


24 


20 


19.3 




15 


18 


10 


16 


3 


15 


12.8 


CLVM1 


5 


5 


4 


4 


5 


5 


5 


4.7 




4 


5 


4 


4 


2 


4 


3.8 


VM2 


24 


20 


15 


15 


20 


20 


20 


19.1 




15 


15 


10 


20 


3 


12 


12.5 


CLVM2 


5 


5 


4 


4 


5 


5 


5 


4.7 




4 


4 


4 


5 


2 


4 


3.8 


SUM 


48 


40 


27 


30 


40 


44 


40 


38.4 




30 


33 


20 


36 


6 


27 


25.3 


V 


HVL 


HVL 


VAL 


VAL 


HVL 


HVL 


HVL 


HVL 




VAL 


HVL 


VAL 


HVL 


SVP 


VAL 


VAL 




stl4 


st30 


st35 


st38 


st39 


st40 


st41 


st51 


KFP 


st21 


st22 


st28 


st29 


st37 


st50 


5? PB 


NIND 


417 


500 


701 


588 


266 


323 


958 


589 


543 


250 


451 


476 


296 


846 


785 


517 


NSP 


38 


52 


48 


43 


38 


41 


38 


48 


43.3 


34 


38 


44 


32 


44 


40 


38.7 


VST 


429 


491 


349 


492 


248 


290 


245 


438 


373 


305 


244 


389 


247 


388 


354 


321 


VSTH 


11.3 


94 


7.3 


11.4 


6.5 


7.1 


6.5 


9.1 


8.6 


8.9 


6.4 


8.8 


7.7 


8.8 


8.9 


8.3 


pEU 


15.8 


26.9 


31.3 


20.9 


29 


31.7 


36.9 


18.8 


26.4 


17.7 


23.7 


15.9 


28.1 


31.8 


27.5 


24.1 


CLpEU 


4 


2 


1 


3 


2 


1 


1 


4 


2.3 


4 


3 


4 


2 


1 


2 


2.7 


CL VST 


5 


5 


4 


5 


3 


4 


3 


5 


4.3 


4 


3 


4 


3 


4 


4 


3.7 


CLVSTH 


5 


5 


4 


5 


4 


4 


4 


5 


4.5 


4 


4 


4 


4 


4 


4 


4 


VM1 


20 


10 


4 


15 


6 


4 


3 


20 


10.3 


16 


9 


16 


6 


4 


8 


9.8 


CLVM1 


5 


4 


2 


4 


3 


2 


2 


5 


3.4 


4 


3 


4 


3 


2 


3 


3.2 


VM2 


20 


10 


4 


15 


8 


4 


4 


20 


10.6 


16 


12 


16 


8 


4 


8 


10.7 


CLVM2 


5 


4 


2 


4 


3 


2 


2 


5 


3.4 


4 


4 


4 


3 


2 


3 


3.3 


SUM 


40 


20 


8 


30 


14 


8 


7 


40 


20.9 


32 


21 


32 


14 


8 


16 


20.5 


V 


HVL 


VAL 


SVP 


VAL 


rvL 


SVP 


SVP 


HVL 


VAL 


VAL 


VAL 


VAL 


IVL 


SVP 


IVL 


VAL 



NIND= nombre d'individus; NSP= nombre d'espèces; VST= valeur stationnelle globale; VSTH= 
valeur stationnelle théorique; pEU= pourcentage des espèces euryèces; CLpEU= classe de pEU; 
CLVST= classe de VST; CLVSTH= classe de VSTH; VMl/2= indice VM1/2; CLVMl/2= classe 
de VM1/2; SUM= VM1+VM2; V= valorisation; stl= station no.l; xAB= moyennes des stations 
abandonnées; xFT= moyennes des stations fauchées tardivement; xFP= moyennes des stations 
fauchées précocement; xEI= moyennes des stations entretenues irrégulièrement; xPO= moyennes 
des stations pâturées par ovins; xPB= moyennes des stations pâturées par bovins; SVP= sans 
valeur particulière; IVL= intéressant, de valeur limitée; VAL= de valeur; HVL= de haute valeur; 
THVL= de très haute valeur. 



EVALUATION DE L ENTRETIEN DES PRAIRIES SECHES 



475 


















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476 



S. POZZI, Y. GONSETH & A. HÄNGGI 



Tab. 2. Tableau du statut de fidélité et de rareté des différentes espèces capturées. Nomenclature 
et systématique selon Maurer & Hänggi (1990); F= indice de fidélité; R= indice de rareté. 



espèce 


F 


R 


espèce 


F 


R 


Atypus affinis 


5 


5 


Walckenaeria dysderoides 


2 


3 


Atypus piceus 


4 


3 


Walckenaeria furcillata 


3 


5 


Dysdera crocata 


3 


5 


Walckenaeria monoceros 


3 


6 


Dysdera erythrina 


2 


2 


Bathyphantes gracilis 


1 


1 


Harpactea hombergi 


2 


3 


Centromerita bicolor 


1 


2 


Harpactocratus drassoides 


3 


3 


Centromerita concinna 


3 


5 


Zodarion italicum 


3 


4 


Centromerus dilutus 


3 


4 


Pachygnatha degeeri 


1 


1 


Centromerus incilium 


2 


4 


Aculepeira ceropegia 


3 


2 


Centromerus serratus 


3 


3 


Araneus diadematus 


1 


2 


Centromerus sylvaticus 


1 


1 


Hypsosinga albovittata 


4 


5 


Diplostyla concolor 


1 


1 


Hypsosinga sanguinea 


4 


3 


Lepthyphantes arenicola 


3 


6 


Neoscona adianta 


3 


6 


Lepthyphantes keyserlingi 


4 


5 


Ero aphana 


2 


4 


Lepthyphantes leprosus 


1 


3 


Erofurcata 


3 


3 


Lepthyphantes mengei 


1 


2 


Acartauchenius scurrilis 


5 


5 


Lepthyphantes pallidus 


2 


1 


Araeoncus humilis 


1 


2 


Lepthyphantes tenuis 


1 


1 


Ceratinella brevis 


1 


1 


Meioneta mollis 


2 


3 . 


Ceratinella scabrosa 


3 


3 


Meioneta rurestris 


1 


1 


Cnephalocotes obscurus 


3 


2 


Meioneta saxatilis 


5 


4 


Dicymbium brevisetosum 


1 


4 


Meioneta simplicitarsis 


2 


5 


Diplocephalus latifrons 


3 


1 


Microlinyphia pusilla 


2 


2 


Eperigone trilobata 


3 


4 


Microneta viaria 


2 


1 


Erigone atra 


1 


1 


Neriene furtiva 


4 


6 


Erigone dentipalpis 


1 


1 


Porrhomma microphthalmum 


2 


4 


Erigonella hiemalis 


3 


5 


Sintula cornigera 


3 


4 


Erigonoplus globipes 


5 


5 


Stemonyphantes lineatus 


2 


3 


Gonatium rubens 


3 


5 


Theonina cornix 


3 


5 


Gongylidiellum latebricola 


3 


3 


Dipoena coracina 


5 


5 


Jacksonella falconeri 


2 


6 


Enoplognatha thoracica 


3 


2 


Metopobactrus prominulus 


3 


4 


Episinus truncatus 


6 


3 


Micrargus herbigradus 


2 


1 


Euryopis flavomaculata 


3 


3 


Micrargus subaequalis 


1 


2 


Euryopis laeta 


5 


6 


Minicia marginella 


3 


5 


Euryopis quinque guttata 


5 


5 


Mioxena blanda 


4 


5 


Neottiura bimaculata 


2 


2 


Monocephalus fuscipes 


2 


2 


Neottiura suaveolens 


5 


5 


Oedothorax apicatus 


1 


1 


Robertus lividus 


1 


1 


Ostearius melanopygius 


2 


5 


Robertus neglectus 


2 


3 


Panamomops sulcifrons 


4 


3 


Steatoda albomaculata 


4 


6 


Pelecopsis parallela 


1 


4 


Steatoda phalerata 


5 


3 


Pocadicnemis juncea 


3 


3 


Theridion Impressum 


3 


2 


Silometopus bonessi 


4 


6 


Theridion nigrovariegatum 


4 


5 


Tapinocyboides pygmaeus 


4 


4 


Alopecosa accentuata 


4 


4 


Tiso vagans 


1 


2 


Alopecosa cuneata 


3 


2 


Trichoncus saxicola 


4 


6 


Alopecosa fabrilis 


4 


5 


Trichopterna cito 


5 


5 


Alopecosa pulverulenta 


1 


1 


Typhochrestus digitatus 


4 


6 


Alopecosa striatipes 


4 


5 


Typhochrestus simoni 


4 


6 


Alopecosa trabalis 


5 


3 


Walckenaeria acuminata 


2 


2 


Arctosa figurata 


5 


5 


Walckenaeria antica 


2 


1 


Aulonia albimana 


2 


1 


Walckenaeria corniculans 


3 


2 


Pardosa agrestis 


1 


2 



ÉVALUATION DE L' ENTRETIEN DES PRAIRIES SECHES 



477 



espèce 


F 


R 


espèce 


F 


R 


Pardosa bifasciata 


5 


4 


Micaria fulgens 


5 


4 


Pardosa hortensis 


1 


1 


Micaria guttulata 


5 


5 


Pardosa saltans 


2 


1 


Micaria pulicaria 


2 


1 


Pardosa monticola 


4 


2 


Phaeocedus braccatus 


4 


5 


Pardosa nigriceps 


4 


6 


Poecilochroa variana 


5 


6 


Pardosa palustris 


1 


1 


Zelotes apricorum 


3 


2 


Pardosa proxima 


4 


4 


Zelotes atrocaeruleus 


4 


6 


Pardosa pullata 


2 


1 


Zelotes civicus 


6 


6 


Pardosa riparia 


4 


4 


Zelotes erebeus 


5 


5 


Pardosa vinata 


3 


6 


Zelotes exiguus 


5 


5 


Pirata hygrophilus 


3 


1 


Zelotes latreillei 


I 


2 


Pirata latitans 


2 


1 


Zelotes lutetianus 


3 


4 


Tricca lutetiana 


3 


2 


Zelotes pedestris 


3 


2 


Trochosa robusta 


6 


4 


Zelotes petrensis 


3 


2 


Trochosa ruricola 


1 


1 


Zelotes praeficus 


4 


2 


Trochosa terricola 


2 


1 


Zelotes pumilus 


4 


4 


Xerolycosa miniata 


4 


4 


Zelotes pusillus 


2 


2 


Pisaura mirabilis 


2 


1 


Zelotes villicus 


5 


5 


Oxyopes lineatus 


3 


6 


Zora nemoralis 


2 


3 


Cicurina cicur 


2 


1 


Zora parallela 


5 


6 


Coelotes inermis 


3 


2 


Zora silvestris 


3 


5 


Coelotes terrestris 


2 


1 


Zora spinimana 


2 


1 


Histopona torpida 


2 


1 


Philodromus rufus 


3 


2 


Mastigusa arietina 


3 


5 


Thanatus formicinus 


4 


3 


Tegeneria silvestris 


3 


1 


Thanatus atratus 


5 


6 


Antistea elegans 


5 


2 


Oxvptila atomaria 


3 


2 


Hahnia nava 


3 


3 


Oxyptila blackwalli 


4 


4 


Hahnia pusilla 


2 


2 


Oxyptila claveata 


5 


4 


Argenna subnigra 


5 


4 


Oxyptila pullata 


3 


6 


Dictyna arundinacea 


3 


4 


Oxyptila scabricula 


5 


4 


Titanoeca quadriguttata 


6 


4 


Oxyptila simplex 


3 


2 


Agroeca brunnea 


3 


2 


Oxyptila trux 


2 


2 


Agroeca cuprea 


4 


4 


Xysticus acerbus 


4 


4 


Agroeca proxima 


3 


6 


Xysticus audax 


2 


2 


Apostenus fiiscus 


3 


3 


Xysticus bifasciatus 


3 


2 


Phrurolithus festivus 


2 


1 


Xysticus cambridgei 


3 


5 


Phrurolithus minimus 


4 


3 


Xysticus cristatus 


1 


1 


Phrurolithus nigrinus 


6 


5 


Xysticus erraticus 


2 


2 


Scotina palliardi 


2 


5 


Xysticus kempeleni 


3 


4 


Cheiracanthium virescens 


4 


5 


Xysticus kochi 


1 


2 


Clubiona brevipes 


3 


3 


Xysticus lineatus 


4 


4 


Clubiona coerulescens 


3 


3 


Xysticus robustus 


4 


4 


Clubiona diversa 


3 


4 


B alius chalybeius 


2 


3 


Clubiona frutetorum 


3 


4 


Bianor aurocinctus 


4 


3 


Clubiona neglecta 


4 


3 


Euophrys aequipes 


2 


3 


Clubiona pseudoneglecta 


4 


6 


Euophrys aperta 


3 


6 


Clubiona terrestris 


2 


2 


Euophrys frontalis 


2 


1 


Callilepsis schuszteri 


6 


6 


Evarcha arcuata 


3 


1 


Drassodes cupreus 


2 


4 


Evarcha laetabunda 


4 


6 


Drassodes lapidosus 


2 


2 


Heliophanus cupreus 


3 


3 


Drassodes pubescens 


2 


2 


Heliophanus flavipes 


3 


2 


Gnaphosa lucifuga 


6 


4 


Myrmarachne formicaria 


5 


2 


Haplodrassus dalmatensis 


5 


6 


Pellenes tripunctatus 


5 


3 


Haplodrassus kulczynskii 


4 


5 


Phlegra fasciata 


3 


3 


Haplodrassus signifer 


1 


1 


Phlegra insignata 


6 


3 


Micaria albimana 


4 


6 


Synageles hilarulus 


4 


6 


Micaria formicaria 


4 


4 









478 s - POZZI, Y. GONSETH & A. HÄNGGI 

COMPARAISON DES STATIONS PAR TYPE D'ENTRETIEN 

Pâturage bovin 

Les stations pâturées par les bovins se sont révélées les moins riches en espèces, 
et celles dont les valeurs stationnelles théorique et globale étaient les plus basses 
(Tab. 1). Soulignons qu'en plaine la pâture par les bovins n'est pas recommandée pour 
l'entretien des prairies sèches: les races modernes étant très lourdes, leur activité 
produit un tassement du sol et une forte dégradation de la prairie (Antognoli et al. 
1995). Les surfaces pentues sont particulièrement touchées par cet effet (station 29 
par ex.). D'autre part, les prairies sèches ne semblent pas suffisamment riches en 
substances nutritives pour la production de lait. Ainsi, seul l'élevage de vaches mères 
(avec des veaux) ou de génisses entre en ligne de compte (Maertens et al. 1990), ce 
qui a été le cas dans les parcelles de cette étude. 

D'après nos résultats, le pâturage bovin pratiqué de manière extensive (moins de 
1 UGBN/ha) 1 peut maintenir des stations de valeurs (st. 21 et 28, voir Fig. 2) mais dès 
que le pâturage devient trop important (plus de 2 UGBN/ha) ou trop précoce dans 
l'année, leur qualité baisse (st. 29, st. 37 et st. 50). La charge en bétail est primordiale, 
une forte pâture entraîne: 1) l'enrichissement en azote du sol qui se traduit par 
l'évolution vers des prairies plus grasses (Arrhenatherion) et moins intéressantes; 2) la 
destruction de la diversité structurelle de la végétation. 

Soulignons que les variations de pente et de structure (présence de buissons, de 
rocaille) au sein d'un pâturage permettent souvent le maintien de secteurs intéressants 
subissant une pression moins forte du bétail. La valeur plus faible de la station 22 
(SUM=21), zone plate jouxtant un pâturage extensif de pente par rapport à celle de ce 
dernier (SUM=32), illustre parfaitement ce fait. 

Le pâturage par des bovins, tel qu'il est actuellement pratiqué, semble défa- 
vorable au maintien de la qualité faunistique des prairies sèches. Il serait toutefois 
nécessaire d'étudier les effets réels d'une pâture extensive tardive pour exclure définiti- 
vement ce type d'entretien des mesures de gestion des milieux ouverts de qualité. Les 
travaux de Gonseth (1994) soulignent en effet qu'une pâture tardive même relative- 
ment forte, peut être compatible avec le maintien d'une faune lépidoptérique intéres- 
sante dans des pâturages maigres. 

Pâturage ovin 

Comme le pâturage bovin, le pâturage ovin ne devrait pas se pratiquer de 
manière intensive dans une prairie sèche. La st. 32 (SUM=6) est l'exemple typique 
d'une prairie surpâturée: grande densité de moutons sur une longue période et de plus 
commençant précocement dans l'année (fin mars). La forte charge dévalorise la station, 
l'herbe devient rase rapidement et la diminution de la structure de la végétation a un 



1 La charge en bétail est exprimée en UGBN, nombre UGB pour 100 jours de pâture; elle 
est calculée à partir des équivalences suivantes pour notre étude: génisse > 2 ans: 4/5 UGB; 
génisse de 1 à 2 ans: 3/5 UGB: veau: 1/3 UGB. 



ÉVALUATION DE L'ENTRETIEN DES PRAIRIES SECHES 479 

impact négatif sur les araignées. L'évolution du site des Curtilles (st. 08 et 48) illustre 
une exploitation défavorable à la faune par son intensité trop grande et sa précocité dans 
l'année. L'idéal serait de pratiquer un pâturage rotatif extensif (recréer l'impact naturel 
d'herbivores sauvages) en laissant un petit nombre d'ovins pendant une courte période 
en fin de saison, ce qui garantirait le maintien de la diversité structurelle du milieu 
essentielle aux araignées (Gibson et al. 1992). La st. 26 (SUM=36) qui a été exploitée 
plus tardivement que la st. 25 (SUM=20) illustre ce principe. 

A propos des stations entretenues annuellement, on peut remarquer que la 
fauche touche toutes les plantes d'une prairie à la même hauteur et au même moment, 
tandis que le pacage agit de façon progressive et sélective (Maertens et al. 1990; 
Gonseth 1994). Kirby (1992) considère qu'un pacage extensif conduit à des milieux 
de vie richement structurés et très favorables aux invertébrés. 

Fauchage précoce 

Concernant le fauchage précoce (mi-juin) des prairies sèches, deux tendances 
sont à relever. 1) Pour des milieux exploités intensivement dans le passé (plusieurs 
fauches dans l'année, engraissement), on peut constater une amélioration de la qualité 
de la station (par exemple, la st. 39 est une prairie en phase d'extensification). 2) Pour 
des milieux exploités extensivement dans le passé (pas fauchés toutes les années ou 
fauchés plus tardivement), on constate une dégradation de la station. La prairie sèche de 
Gland est un bon exemple de ce cas de figure. En 1995, la st. 14 a une valeur identique 
à celle de la st. 15 (HVL) qui se situe dans le même site mais qui n'est pas entretenue. 
Puis en 1996, la st. 38 (équivalente de la st. 14) a perdu de la valeur suite à son 
fauchage précoce. A signaler également dans ce site que la fauche est très rase ce qui, 
dans les conditions présentes, ralentit la repousse et fait que certaines araignées ne 
trouvent plus la structure du milieu nécessaire à leur développement. 

En résumé, on peut remarquer que dans un premier temps les prairies intensives 
vont gagner de la valeur avec un fauchage mi-juin mais à long terme il faut retarder le 
fauchage de plus en plus et extensifier l'entretien si l'on veut obtenir ou maintenir des 
stations de haute valeur. Gonseth (1985) confirme d'ailleurs qu'un fauchage précoce 
effectué sur une longue période (15 ans) contribue à appauvrir la communauté 
arachnologique des prairies sèches. A relever finalement que le fauchage précoce est le 
seul entretien à regrouper trois stations sans valeur particulière. Il obtient même la plus 
basse moyenne finale (SUM) avec le pâturage bovin (Tab. 1). 

Fauchage tardif 

Pour l'ensemble des stations étudiées, le fauchage tardif est le meilleur entretien 
régulier (SUM la plus élevée pour une station entretenue). La st. 2 obtient même la plus 
haute valeur stationnelle théorique (13.3) et la st. 46 la plus haute valeur stationnelle 
globale (621). 

Dans certains cas, ce mode d'entretien pourrait être encore amélioré ponctuelle- 
ment par des entretiens tous les deux ans ou par un fauchage parcellaire en rotation. Le 



480 S. POZZI, Y. GONSETH & A. HÄNGGI 

site de La Rippe illustre ce fait: si les stations 12 et 13 se situent dans la même prairie, 
la première est fauchée tardivement toutes les années alors que la deuxième est fauchée 
tous les deux ans. Malgré leur proximité, la st. 13 est classée dans les stations de haute 
valeur (SUM=40) tandis que la st. 12 dans les stations de valeur (SUM=27). Par contre, 
en 1996, après l'entretien de l'ensemble du site, la st. 43 (équivalente de la st. 13) 
retrouve une valeur semblable à la st. 12 (ou st. 42). Ainsi, son peuplement arachnolo- 
gique est moins intéressant que celui de la deuxième année sans entretien. L'expérience 
montre qu'une modulation des pressions anthropiques par rotation des contraintes entre 
les divers secteurs d'une même aire protégée permettrait une meilleure conservation des 
communautés animales, en augmentant leur richesse et en améliorant leur spécificité au 
biotope par rapport au cas d'une pression anthropique continue. 

Globalement, la réponse de la faune aux effets de la fauche varie selon l'inten- 
sité de l'intervention, sa fréquence et le moment de l'année où elle se situe. Des coupes 
bien menées (selon la dynamique du milieu) à des moments adéquats de l'année 
(automne), semblent favorables à la conservation de la valeur arachnologique des 
stations. 

Entretien irrégulier 

Vu la variété des entretiens rassemblés dans cette catégorie et le faible nombre 
de stations étudiées, il est difficile de tirer des conclusions sur ce type d'entretien. Des 
tendances semblent toutefois se dessiner: les stations fauchées tardivement mais irrégu- 
lièrement (st. 13, 17 et st. 43) sont généralement des prairies de valeur voir de haute 
valeur; la st. 4, quant à elle, entretenue intensivement par le passé (pâturée et légère- 
ment engraissée), est intéressante mais de valeur limitée. Elle nécessiterait, dans un 
premier temps, un entretien régulier (par exemple une fauche mi-juin) sans fumure mais 
avec exportation de la matière végétale pour diminuer progressivement la richesse du 
sol et la densité de la végétation, puis, dans un second temps, un fauchage de plus en 
plus tard dans la saison. 

Stations abandonnées 

Les stations abandonnées obtiennent les meilleures valeurs stationnelles et 
recèlent l'unique station de très haute valeur: st. 27. Elles se caractérisent par la plus 
faible moyenne des pourcentages d'espèces euryèces (Tab. 1), autrement dit par la 
présence d'un grand nombre d'espèces exigeantes face aux conditions de leur habitat. 

Lorsqu'une prairie sèche est abandonnée, différents facteurs, importants pour les 
animaux et les plantes sont modifiés. La suppression de la fauche annuelle et l'accu- 
mulation de litière qui en résulte sont les plus importants (Antognoli et al. 1995). Elles 
créent en principe de nouvelles conditions de vie au niveau de la surface du sol. Pour 
les araignées, l'absence de fauche a pour conséquence que la structure de leur milieu de 
vie, formée par la végétation, est conservée, et que les conditions microclimatiques qui 
y sont liées restent plus ou moins constantes. 



EVALUATION DE L ENTRETIEN DES PRAIRIES SECHES 4g 1 

Dans cette étude, les stations abandonnées sont de deux types: des prairies très 
xériques à évolution lente et des prairies en voie d'embroussaillement prononcé. Le 
premier cas est typique des zones xériques qui devraient être entretenues le moins 
possible. Les stations du plateau du Moulin-de-Vert (st. 1 et 45) et de l' Allondon (st. 1 1 
et 49) illustrent bien cette situation. Pour ces milieux de grande taille, l'absence 
d'entretien leur convient bien parce que les conditions extrêmes (climat, sol) empêchent 
un changement microclimatique trop rapide (végétation herbacée dense, buissons). La 
tendance au feutrage, due au manque de fauche, n'entraîne pas la disparition des 
espèces les plus xerophiles, puisqu'il existe encore assez de zones ouvertes (terre, 
cailloux) dans la station. Deux autres stations xériques (st. 24 et 14) entretenues 
régulièrement pourraient être améliorées en intervenant plus extensivement. 

En ce qui concerne les zones moins sèches avec un sol plus profond (st. 15 et 
34), il est nécessaire de contenir régulièrement la progression des ligneux sinon la forêt 
s'installe, mais il ne semble pas nécessaire de faucher toutes les années. D'après nos 
résultats, le fauchage tardif pourrait être une bonne méthode pour maintenir des zones 
ouvertes de qualité. 

L'évaluation de la qualité de deux stations abandonnées (st. 27 et 33) par le biais 
de la méthode que nous avons choisie illustre bien la différence de signification des 
indices VM1 et VM2. L'extrême variabilité du couvert végétal de la station 27, où 
alternent des zones de dalles nues ou peu colonisées, de pelouses et de buissons, se 
traduit par sa grande diversité faunistique; dans ce cas, la valeur de VM1 est supérieure 
à celle de VM2. Par contre, la st. 33 est un milieu extrême (très homogène et de grande 
stabilité) caractérisé par la valeur de VM2 supérieure à celle de VM1. D'ailleurs dans 
les analyses multivariées (Pozzi & Borcard soumis), cette prairie sèche se distingue 
faunistiquement des autres stations. 

En résumé, la non-intervention à moyen terme permet de maintenir une structure 
diversifiée de la végétation favorable à de nombreuses espèces pour réaliser leur cycle 
de vie (Kirby 1992). 

CONCLUSIONS SUR L'ENTRETIEN 

Notre étude se base sur une assez grande variété de situations qui reflète l'état 
des prairies sèches de la région. Si quelques mesures générales peuvent être proposées 
pour leur entretien (faucher tardivement, laisser des zones non entretenues), chaque 
station nécessite une gestion particulière en fonction des variables biotiques, abiotiques 
(pente, surface) et historiques qui l'influencent. Tous ces facteurs doivent être consi- 
dérés pour définir un plan de gestion de ces milieux. Les propositions suivantes liées 
aux différents entretiens étudiés permettent d'augmenter la valeur d'une prairie sèche. 

• Les stations abandonnées abritent une faune arachnologique plus intéressante 

(haute valeur VM1 et VM2) que les milieux entretenus. Elles sont utilisées 
comme refuge par les espèces appréciant un habitat de transition non perturbé. 
Toutefois, pour éviter leur recolonisation par la forêt à plus ou moins long 
terme, il est recommandé d'entretenir ces parcelles afin de maintenir l'éco- 



482 s - POZZI, Y. GONSETH & A. HÄNGGI 

système prairie sèche et les espèces qui le caractérisent. L'abandon d'une station 
ne devrait intervenir que dans les zones à végétation stable (sur des sols super- 
ficiels avec des conditions très xériques). Si la mise en friche a des effets positifs 
pour les araignées, la perte d'une partie de la diversité botanique qui en découle 
peut également être préjudiciable pour la faune à long terme. 

• La fréquence de l'entretien peut varier de quelques années pour les stations les 
plus xériques (steppes, xérobromion) à une intervention annuelle pour les sites 
plus dynamiques et très productifs. 

• En ce qui concerne l'importance de l'entretien, il faut éviter toute intervention 
intensive et proscrire tout épandage direct d'engrais. Ainsi, on doit diminuer la 
charge en bétail pour augmenter la valeur d'un site. En plus, il semble primor- 
dial de retarder et de réduire la période de pâture. C'est pour ces raisons qu'un 
pâturage extensif et tournant sur différents sites selon les années pourrait être 
judicieusement pratiqué. 

• La fauche doit être si possible tardive (automnale), ce qui favorise le déve- 
loppement des espèces estivales et ménage des refuges pour les araignées dont la 
période d'activité est longue ou tardive. Selon nos résultats, une fauche précoce 
se traduit par une baisse sensible de la qualité arachnologique des stations. De 
plus, nous recommandons une fauche parcellaire et alternée (sur 2 à 3 ans) en 
divisant les surfaces pour laisser certaines zones non entretenues. Ceci permet, 
en plus du maintien de la flore caractéristique des prairies sèches, la conser- 
vation de la structure du milieu nécessaire à la survie de nombreuses espèces. 
Certains buissons peuvent être ainsi conservés. Enfin, il faut éviter de faucher 
trop ras, car une action aussi brutale a un effet catastrophique sur certains 
éléments de la faune. De plus, il ne faut pas oublier d'enlever la matière orga- 
nique fauchée pour ne pas engraisser le sol. 

L'ensemble de ces informations doit être considéré comme un apport à la 
compréhension générale de l'écosystème prairie sèche, mais aussi comme des réponses 
à des questions individuelles des différentes stations étudiées. 

De manière générale, il faut rappeler qu'une gestion orientée sur la persistance 
de l'ensemble des phases d'une dynamique végétale, favorise la diversité spécifique. Le 
principe de base est une rotation des interventions sur le site et la mise en place d'un 
système de corridors permettant une recolonisation des habitats et le maintien de la 
diversité. Cependant, il faut préciser que l'objectif de la gestion des prairies sèches ne 
doit pas être forcément la présence d'une diversité biologique maximale mais le 
maintien d'une structure du milieu qui favorise les espèces caractéristiques des 
habitats menacés. 

CONCLUSIONS GÉNÉRALES 

Parallèlement à l'actualisation de l'inventaire national des prairies sèches, il 
nous a semblé important d'étudier la faune arachnologique de ces milieux. Ce travail a 
permis de perfectionner une méthode d'évaluation de la qualité de stations semi- 



ÉVALUATION DE L ENTRETIEN DES PRAIRIES SECHES 483 

naturelles par les araignées et de discuter des différents types d'entretiens dans une 
optique de protection de la nature. 

La méthode, basée sur l'étude du peuplement arachnologique d'une station 
permet, par le calcul de deux indices différents (VM1 et VM2), de ne pas sous-estimer 
la qualité des biotopes homogènes possédant des araignées spécifiques aux conditions 
du milieu. Les modifications apportées aux analyses permettent une meilleure hiérar- 
chisation des différentes stations, notamment celles de haute valeur. Ce résultat permet 
de proposer un entretien favorable à la faune arachnologique pour l'ensemble des 
stations. 

Nos résultats prouvent que les araignées sont de bons outils d'évaluation de la 
qualité des prairies sèches. Par leur position élevée dans la chaîne alimentaire, elles sont 
capables de donner une information qualitative pour d'autres groupes faunistiques. 
L'étude de certains sites durant deux années consécutives a permis de mettre en 
évidence la finesse et la rapidité de réaction des peuplements arachnologiques aux 
modifications des mesures de gestion. Trois cas de figures ont pu être observés: 1) des 
stations dont l'entretien inadapté a provoqué la diminution immédiate de leur valeur 
arachnologique; 2) des stations dont la valeur est restée stable; 3) des stations dont la 
valeur a rapidement augmenté à la suite d'un entretien plus extensif. Ces résultats sont 
importants mais devront être confirmés par des études à plus long terme. A l'avenir, il 
semble donc important d'utiliser les araignées pour caractériser un milieu et son évo- 
lution (dynamique) en parallèle de la phytosociologie. Pour cela, il est nécessaire de 
poursuivre les études autécologiques fines sur la faune arachnologique, en particulier 
sur les espèces liées à des habitats précis. Cette démarche est indispensable pour l'éla- 
boration de méthodes de gestion permettant la conservation des populations menacées. 

Les problèmes relatifs à la protection des prairies sèches ne sont de loin pas tous 
résolus. La prochaine étape est de compléter les propositions pratiques de mesure de 
gestion et de mettre sur pied un programme de suivi des effets des mesures proposées. 
Ce dernier doit servir à vérifier le bien-fondé des buts de protection et à améliorer les 
mesures de gestion. Le succès de ces différents pas dépend de l'obtention de connais- 
sances scientifiques sur les prairies sèches qui doivent englober des aspects biotiques et 
abiotiques ainsi que leurs interactions. 

Par ce travail, nous espérons avoir contribué à stimuler la sauvegarde de ces 
écosystèmes qui représentent un milieu de vie primordial pour de nombreuses espèces 
tant végétales qu'animales. 

REMERCIEMENTS 

Cette étude n'aurait pu s'effectuer sans l'aide du Prof. V. Mahnert, directeur du 
Muséum d'histoire naturelle de Genève. Elle a été réalisée avec le soutien financier de 
l'Office fédéral de l'environnement, des forêts et du paysage (OFEFP, BUWAL), du 
Service de la Conservation de la Faune (Canton de Vaud), du Service des forêts, de la 
faune et de la protection de la nature (Canton de Genève), du Naturhistorisches 
Museum Basel et du Muséum d'histoire naturelle de Genève. 



484 S- POZZI, Y. GONSETH & A. HÄNGGI 



BIBLIOGRAPHIE 

Antognoli, C, Lörtscher, M., Guggisberg, F., Häfelfinger, S. & Stampfli, A. 1995. Prairies 

maigres tessinoises en mutation. Cahier de l 'Environnement No 246, Nature et paysage, 

BUWAL, Bern. 137 pp. 
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Revue suisse de Zoologie 105 (3): 487-492; septembre 1998 



A revision of the Habrocerinae of the world. Supplement II 
(Coleoptera: Staphylinidae) 



Volker ASSING 

Gabelsbergerstr. 2, D-30163 Hannover, Germany. 



A revision of the Habrocerinae of the world. Supplement II (Coleop- 
tera: Staphylinidae). - Further data on the distribution of several species 
of Habrocerus Erichson and Nomimocerus Coiffait & Saiz are presented. 
H. neglectus sp. n. from southern Thailand, the sister species of H. rouge- 
monti Pace, and Nomimocerus septentrionalis sp. n. from Chile, a close 
relative of N. longispinosus Assing & Wunderle, are described. Their 
primary and secondary sexual characters are figured. 

Key-words: Coleoptera - Staphylinidae - Habrocerinae - Thailand - Chile 
- taxonomy - new species. 



INTRODUCTION 

Before the present study, the two genera of Habrocerinae comprised 19 species 
worldwide, 14 species of Habrocerus Erichson and 5 species of Nomimocerus 
Coiffait & Saiz. Habrocerus is widely distributed in the Palaearctic, the Oriental, the 
Nearctic and the Neotropical region; the genus is absent from the Ethiopian region, 
and in the Australian region represented only by the probably introduced and wide- 
spread H. capillaricornis (Gravenhorst). Nomimocerus, in contrast, is confined to the 
Chilean subregion (Assing & Wunderle 1995, 1996). 

In the course of examining new Habrocerus material from Thailand it was 
discovered that what had previously been treated as H. rougemonti Pace in fact refers 
to two closely related species, one of them new to science and described below. An 
examination of previously unrevised and recently collected material of Nomimocerus, 
deposited in the collections of the University of Kansas, yielded a further species of 
Nomimocerus, so that the figure now stands at 21 species of Habrocerinae worldwide. 

In addition to the descriptions, new faunistic data, which have become 
available since the first supplement (Assing & Wunderle 1996), are presented. 



Manuscript accepted 21.04.1998 



488 VOLKER ASSING 

Material from the following institutions and private collections was studied: 

DEI Deutsches Entomologisches Institut, Eberswalde (L. Zerche) 

MHNG Muséum d'histoire naturelle, Genève (I. Lobi) 

UKNHM University of Kansas Natural History Museum, Lawrence (J. S. Ashe, 

R. W. Brooks) 
cAss private collection, V. Assing 

cWun private collection, P. Wunderle 

NEW SPECIES AND RECORDS OF HABROCERINAE 

Habrocerus capillaricornis (Gravenhorst) 

Canary Islands: 13, 49, Gran Canaria, N Teror, E Osorio, El Palmar, 600m, in shady 
barranco with fragments of Laurisilva, 20.XII.1997, leg. Assing & Wunderle (cWun). 

United States of America: 1 3 , 3 9 , North Carolina, Watauga Co., Boone, 12.111./ 10.V./ 
19.V.1973, leg. Ashe (UKNHM). 

H. capillaricornis is here for the first time recorded from Gran Canaria. In the 
Canarian archipelago, the species was previously known only from Tenerife, La 
Palma and La Gomera. Numerous additional specimens from Central and Southern 
Europe and from Algeria were examined; the records are not listed in detail, as the 
species is very common in these regions. 

Habrocerus pisidicus Korge 

Bosnia-Herzegovina: 3â , 19, Bjelasnica planina (MHNG). 

Bulgaria: 23, 29, Pirin, leg. Weirather (MHNG); 3 9, Ali Botusch, 'N-Seite' near 
Goleschowo, 1015m, 41°42'13N, 23°35'21E, Fagus wood, 15.VI.1997, leg. Behne (DEI); 13, 
Sredna Gora, S Koprivschtiza, 1110m, 42°35'12N 24°22'19E, Fagus wood, 29.VI.1997, leg. 
Zerche & Behne (cAss). 

Greece: 1 3, 3 9, Crete, Zakros, 'Tal der Toten', 28.III.1973, leg. Falscher & Meybohm 
(MHNG); 5ó\ 49, Ródhos, M. Kariona, 400m, 11. IV. 1977, leg. Besuchet (MHNG, cAss); 29, 
Ródhos, Petaloudes, 8.IV.1977, leg. Besuchet (MHNG). 

H. pisidicus is widespread in the southeastern Mediterranean and has been 
known to occur in the regions indicated above. 

Habrocerus ibericus Assing & Wunderle 

Spain: 23, 19, E. Asturias, Cibes, Sierra de la Serrantina, 800m, 16. V. 1997, leg. 
Starke (coll. Feldmann, cAss); 19, Albacete, leg. Comellini (MHNG); lo*, Prov. Albacete, 
Villaverde, 15.IV.1959, leg. Besuchet (MHNG); \3, Prov. Tarragona, Sierra de Moutsant (?), 
23.III.1959, leg. Besuchet (MHNG); 1 o\ 19, Prov. Lèrida, 2.VI.1965, leg. Comellini 
(MHNG); 2o\ 1 9, Prov. Teruel, Rubielos de Mora, 19.IX.1971, leg. Comellini (MHNG); 19, 
Prov. Murcia, Sierra de Espuha, 26.III.1959, leg. Besuchet (MHNG); 33, 19, Prov. Jaen, 
Sierra de Cazorla, 12.IV. 1959, leg. Besuchet (MHNG). 

Previously, only the type specimens of H. ibericus were known from the 
Iberian Peninsula, where the species seems to be both widely distributed and quite 
common. 



A REVISION OF THE HABROCERINAE OF THE WORLD 



Habrocerus neglectus sp. n. Figs 1-4 

Habrocerus rougemonti Pace: Assing & Wunderle 1995 (partim, Figs 9a-g, 1 lg) 

Holotype 6: THAILAND. NE Bangkok, Khao Yai Nat. Park, Khao Khieo, 1 150m, leg. 

Burckhardt & Lobi, 28.XI.85 (MHNG). 

Paratypes: MS , 19$, same data as holotype (MHNG, cAss, cWun); 3cT, Chiang Mai, 

Mae Nang Kaeo, 900m, 54 km NE Chiang Mai, leg. Burckhardt & Lobi, 3.XI.1985 (MHNG, 

cAss). 

Diagnosis: Closely related to and externally indistinguishable from H. rougemonti 

Pace, its sister species. 

â: tergum and sternum VII slightly broader than in H. rougemonti (Fig. 3); modified 

sternum VIII anteriorly distinctly broader, less convex and mostly without median 

carina, posteriorly with wider emargination and more distinctly sclerotized margins 

(Fig. 2); internal sac with shorter row of semitransparent triangles and anteriorly with 

ca. four weakly sclerotized coniform spines (Fig. 1); appendices of pleurites VIII 

weakly S-shaped (see Fig. 9a in Assing & Wunderle 1995). 

9 : tergum and sternum VIII broader and shorter than in H. rougemonti (Fig. 4). 

Comments: For further details regarding external and secondary sexual characters, the 
description and figures in Assing & Wunderle (1995) are referred to. Both the 
figures and the material listed from the surroundings of Bangkok refer to this species 
and not to H. rougemonti. 

Distribution: H. neglectus is known only from two localities, one in northern and 
one in southeastern Thailand. It appears doubtful that H. neglectus and H. rougemonti 
should have a para- or allopatric distribution; both species are macropterous, and the 
northern locality of H. neglectus (Mae Nang Kaeo) is only some 70 km away from 
Doi Suthep, where H. rougemonti was found. 

Habrocerus rougemonti Pace Figs 5 - 9 

Habrocerus rougemonti Pace: Assing & Wunderle 1995 (partim) 

Additional material examined: 

Thailand: 5o\ 1 9, mountains near Umphang, 1250 m, 10.11.1993, leg. Schwendinger (MHNG, 

cAss). 

Diagnosis: â: tergum and sternum VII slightly more oblong than in H. neglectus 
(Fig. 8); modified sternum VIII anteriorly more slender and more convex than in 
H. neglectus and with median carina, posteriorly with narrower emargination and 
relatively weakly sclerotized margins (Fig. 6); internal sac with longer row of semi- 
transparent triangles and anteriorly with a large blackish spine (Fig. 5); appendices of 
pleurites VIII weakly and evenly curved (Fig. 7). 

$ : terminalia similar to H. neglectus, but tergum and sternum VIII more oblong 
(Fig. 9). 

Distribution: H. rougemonti appears to have a more restricted distribution than 
previously stated. So far, the species has become known from three localities in 
northern and western Thailand. 



490 VOLKER ASSING 



Habrocerus schwarzi Horn 



Canada: 9c? , 49, Alberta, George Lake, 53°57'N, 1 14°06'W, ex mushrooms, 11./ 
22.VIII.1977, leg. Ashe (UKNHM, cAss); lc?, Alberta, Fort MacKay, aspen spruce, pitfall 
trap, 2.IX.1978, leg. Ryan & Hilchie (UKNHM). 

Previously only one record from Alberta had been known (Assing & 

WUNDERLE 1995). 

Habrocerus tropicus Wendeler 

1 9 , Brazil, Linha Facao, Santa Catarina, V.1954, leg. Plaumann (UKNHM). 
H. tropicus has become known only from Brazil. 

Nomimocerus longispinosus Assing & Wunderle 

Chile: 2 c?, Chiloé, Isla Chiloé, San Juan de Chadmo, 20.11.1997, leg. Cekalovic 
(UKNHM, cAss); 2 9,1 ex., Chiloé, Isla Chiloé, 5km SW Chonchi, 20.11.1997, leg. Cekalovic 
(UKNHM). 

Previously recorded only from Aisén province, Chile, N. longispinosus is now 
also known from Isla de Chiloé, Chiloé province. 

Nomimocerus peckorum Assing & Wunderle 

Chile: 4c?, 49, Osorno, 14 km E Termas de Puyehue, 40°40 , S, 71°14'W, 350 m, ex 
fungus. on live vine, 30.XI.1994, leg. Leschen & Carlton (UKNHM, cAss); 1 9, same data, but 
450 m, ex sifting leaf litter (UKNHM); lc?, 19, Valdivia, 10 km NW Choshuenco, 39°45'S, 
72°20'W, 250 m, sifting leaf litter, 15.XI.1994, leg. Leschen & Carlton (UKNHM, cAss); 19, 
Valdivia. 37 km SE Panguipuli. 39°45'S, 72°20'W, 300 m, 14.XI.1994, leg. Leschen & Carlton 
(UKNHM); lc?, Cautin, 8 km S Pucon, 1075 m, 39°21'S, 71°58'W, 23.XI.1994, leg. Leschen 
& Carlton (UKNHM); lc?, Cautin, 26.7 km E Pucon, 625 m, 39°39'S, 71°47'W, 24.XI.1994, 
leg. Leschen & Carlton (UKNHM); 1 c? , Cautin, Termas de Palguin, Salto Puma, 725 m, 
39°22'S, 71°50'W, 25.XI.1994, leg. Leschen & Carlton (cAss). 

/V. peckorum was originally described from Osorno and Llanquihue province, 
Chile, and is here recorded from two further provinces: Valdivia and Cautin. 

Nomimocerus septentrionalis sp. n. Figs 10 - 12 

Holotype c?: CHILE: Coquimbo, 6 km SW Hurtado. 1040 m, Puente Morrillos, 
30°16'S, 70°40'W, 28 Oct 1994, R. Leschen & C. Carlton, #021, ex: along stream (UKNHM). 

Paratypes: 5c?, 3 9, same data as holotype (UKNHM, cAss); 6c?, 49, CHILE: 
Coquimbo, 850 m, 5 km S. Carén, Pte. El Cuiyano, Rio Limari, 30°52'S, 70°45'W, #033, 
R. Leschen, C. Carlton, ex: leaf litter, 30.X.1994 (UKNHM, cAss); 2c?, 9 9 [29 teneral], 
CHILE: Coquimbo, 6 km W Hurtado, 1040 m, Puente Morrillos, 30°16'S, 70°40'W, 28 Oct 
1994, R. Leschen & C. Carlton. #022, ex: sifting litter (UKNHM, cAss). 

Figs 1-12: Habrocerus neglectus sp. n. (1 - 4) and H. rougemonti Pace (5 - 9): internal sac in 
squeezed preparation (1, 5); outlines of c? sternum VIII (2, 6); appendix of e? pleurite VIII (7); 
e? tergum VII (left) and sternum VII (right) (3, 8); outlines of 9 tergum VIII (left) and sternum 
VIII (right) (4. 9). Nomimocerus septentrionalis sp. n. (10-12): internal sac in squeezed 
preparation (10); c? tergum VII (11); outline of hind margin of sternum VII of two c? (12); 
setae and pubescence partly or completely omitted in 1 1 and 12. Scales: 0.25 mm. 



A REVISION OF THE HABROCERINAE OF THE WORLD 



491 






492 VOLKER ASSING 

Diagnosis: Total length: 3.8 - 4.8 mm; pronotai length: 0.68 - 0.76 mm; pronotai 
width: 1.03 - 1.15 mm; elytral length: 0.71 - 0.80 mm. In external morphology and 
colour highly similar to N. peckorum and N. longispinosus, but on average larger (see 
measurements); hind wings in all the type specimens reduced. 

6: posterior margin of sternum VII with concavity of variable depth and width 
(Fig. 12); hind margin of tergum VII more deeply incised centrally than in N. pecko- 
rum and N. longispinosus (Fig. 11); internal sac with internal structures similar to 
N. longispinosus, but spines of long series stouter and caudad more gradually 
decreasing in length (in N. longispinosus the transition is ± abrupt), the second (short) 
series longer, and the sclerotized terminal piece distinctly more massive and less 
slender than in that species (Fig. 10). 

Distribution and bionomics: The species was collected near Coquimbo, Chile 
(between 30° and 31° southern latitude), distinctly further north (name!) than the 
known areas of distribution of its congeners. The type specimens were taken in leaf 
litter and in the vicinity of a stream at altitudes between 850 and 1040 m at the end of 
October; two of them were teneral. 

ACKNOWLEDGEMENTS 

I am indebted to the colleagues indicated in the introduction for the loan of the 
material which this study is based on. 

REFERENCES 

Assing, V. & Wunderle, P. 1995: A revision of the species of the subfamily Habrocerinae 
(Coleoptera: Staphylinidae) of the world. Revue suisse de Zoologie 102: 307-359. 

Assing, V. & Wunderle, P. 1996: A revision of the species of the subfamily Habrocerinae of 
the world. Supplement I. (Coleoptera: Staphylinidae). Beiträge zur Entomologie 46: 
373-378. 



Revue suisse de Zoologie 105 (2): 493-497; septembre 1998 



Beschreibung von vier neuen Arten der Gattung Derarimus 
(Coleoptera, Anthicidae) aus Malaysia 



Gerhard UHMANN 

Tannenhofstrasse 10, D-92690 Pressath, Deutschland. 



Description of four new species of the genus Derarimus (Coleoptera, 
Anthicidae) from Malaysia. - Following species are described and 
illustrated: Derarimus ampliaticornis sp. n., Derarimus bicavatus sp. n., 
Derarimus compacticornis sp. n. and Derarimus sabahensis sp. n. 

Key-words: Coleoptera - Anthicidae - Tomoderini - Derarimus - new 
species - Malaysia. 



EINLEITUNG 

Eine weitere Bestimmungssendung, die mir Herr Dr Ivan Lobi vom Natur- 
historischen Museum in Genf übergab, enthält vier bisher nicht beschriebene Arten 
der Gattung Derarimus, die nachfolgend beschrieben werden. 

Die Gattung Derarimus wurde 1978 für eine neue Art, D. carinatus aus Indien 
von Bonadona beschrieben. Außerdem stellte Bonadona Tomoderus excisicollis 
Heberdey aus Java in die neue Gattung. 1986 versetzte Sarai Tomoderus clavipes 
Champion in die Gattung Derarimus. Zur Zeit sind (einschließlich der hier beschrie- 
benen) 51 Arten beschrieben. Außer D. clavipes (Champion) aus der Paläarktis 
(Japan) sind alle Arten in der Orientalis beheimatet, nämlich in Malaysia, Indonesien, 
Indien, Vietnam, Thailand und Taiwan. 

Eine Bestimmungstabelle der Arten ist bei Uhmann, 1994 zu finden, eine 
Ergänzung dazu bei Uhmann 1996. 

Herrn Dr I. Lobi danke ich sehr, daß er mir die Bearbeitung dieser Tiere 
ermöglichte sowie für die Überlassung einiger der Käfer für meine Sammlung. Herrn 
Dr Volker Mahnen, ebenfalls vom Naturhistorischen Museum in Genf danke ich sehr 
für die redaktionelle Beratung. 

Alle Holotypen der hier beschriebenen Arten befinden sich im Naturhis- 
torischen Museum in Genf. Alle Maße sind in mm angegeben. 



59. Beitrag zur Kenntnis der Anthicidae 
Manuskript angenommen 05.02. 1 998 



494 GERHARD UHMANN 

BESCHREIBUNGEN 

Derarimus ampliaticornis sp. n. Abb. 1-3 

E-Malaysia, Sarawak, Gn Penrissen, 1000 m, 23.5.1994, edge prim, montane for., # 9 a, 
1 Ex., leg. Lobi & Burckhardt. Holotypus. 

Länge 2,9, größte Breite 1,0. Kopf 0,6 lang, über die Augen gemessen 0,6 
breit. Halsschild 0,7 lang, 0,4 breit. Flügeldecken 1,5 lang, 1,0 gemeinsam breit. 

Färbung: Braun. Taster, Fühler und Beine gelbbraun. 

Kopf: Glänzend. Fein und verstreut punktiert. Behaarung braun, kräftig, ziem- 
lich kurz, gebogen, etwas abstehend, in verschiedene Richtungen weisend. Außerdem 
mit wenigen, nicht sehr langen, geraden Borsten, Fühler mit langer, kräftiger 
Behaarung. 

Halsschild: Glänzend. In der Mitte ziemlich kräftig, zwischen den Kerben sehr 
kräftig, sonst sehr fein punktiert. Behaarung braun, kräftig, etwas gebogen, etwas 
abstehend, größtenteils nach hinten gerichtet. Außerdem mit zahlreichen langen, fast 
geraden Borsten, die senkrecht abstehen. 

Flügeldecken: Glänzend. Sehr kräftig, aber flach punktiert. Die Zwischen- 
räume sind kleiner als die Punkte. Zur Spitze Zur Spitze werden die Punkte etwas 
feiner und etwas tiefer. Die Zwischenräume werden etwas größer. Behaarung braun, 
kräftig, ziemlich lang, etwas gebogen, etwas abstehend, nach hinten gerichtet. Außer- 
dem mit zahlreichen, nicht sehr langen, meist geraden, senkrecht abstehenden 
Borsten. 

Beine unauffällig behaart. 

Beziehungen: Dem Derarimus luteipes Uhmann aus W-Malaysia (Pahang) 
ähnlich, aber der Halsschild ist schlanker und neben den Kerben befinden sich feine 
Längskielchen. 

Derarimus bicavatus sp. n. Abb. 4 und 5 

E-Malaysia, Sarawak, confi. Sun Oyan and Mujong riv., E Kapit. 50 m, 18.5.1994, 
# 5 a, 1 Ex., leg Lobi & Burckhardt, Holotypus. 

Länge 5,2, größte Breite 2,1. Kopf 0,9 lang, über die Augen gemessen 1,0 
breit. Halsschild 1,5 lang, 1,0 breit. Flügeldecken 3,0 lang, 2,1 gemeinsam breit. 

Färbung: Kopf und Halsschild rotbraun. Flügeldecken etwas dunkler braun. 
Fühler, Taster und Beine gelbbraun. 

Kopf: Glänzend. Sehr fein und flach punktiert. Zwischenräume viel größer als 
die Punkte. Behaarung braun, kräftig, gebogen, etwas abstehend, in verschiedene 
Richtungen weisend. Fühlerbehaarung kräftig und abstehend. 

Halsschild: Glänzend. Sehr fein, in der Abschnürung kräftig punktiert. Be- 
haarung braun, kräftig, wenig gebogen, abstehend, in verschiedene Richtungen 
weisend. In der hinteren Hälfte, jederseits der Mitte mit einer flachen, aber deutlichen 
Grube, so daß in der Mitte und seitlich kielartige Erhebungen stehen bleiben. 

Flügeldecken: Glänzend. Ziemlich kräftig punktiert. Zwischenräume kleiner als 
die Punkte. Zur Spitze wird die Punktur nur etwas feiner, bleibt aber dicht. Behaarung 
braun, kräftig, etwas gebogen, halb abstehend, ziemlich dicht, nach hinten gerichtet. 



BESCHREIBUNG VON VIER NEUEN ARTEN DER GATTUNG DERARIMUS 



495 




Abb. 1-5 

Derarimus ampliaticornis sp. n. (1-3): 1. Habitus; 2. Halsschildprofil; 3. Aedeagusspitze, 
dorsal. - D. bicavatus sp. n. (4-5): 4. Habitus; 5. Halsschildprofil. 



Beine mit kräftiger, aber sehr kurzer Behaarung. 

Beziehungen: Dem Derarimus robusticornis Uhmann aus Sabah etwas ähn- 
lich, aber die Schläfen sind länger, der Halsschild ist schlanker. 



Derarimus compacticornis sp. n. 



Abb. 6 und 7 



E-Malaysia, Sarawak, Gn Penrissen, 1000 m, 23.5.1994, edge prim, montane for., # 9 a, 
3 Ex., leg. Lobi & Burckhardt, Holotypus, 2 Paratypen. E-Malaysia, Sarawak, Gn Matang, 
20 km W Kuching, 800 m, 13.5.1994, submontane forest, # 2 a, 1 Ex., leg Lobi & Burckhardt, 
Paratypus. E-Malaysia, Sarawak, confi. Sun Oyan and Mujong riv., E Kapit, 50 m, 18.5.1994. 
# 5 a, 1 Ex., leg. Lobi & Burckhardt, Paratypus. 



496 



GERHARD UHMANN 




Abb. 6-11 

Derarimus compacticornis sp. n. (6-7): 6. Habitus; 7. Halsschildprofil. - D. sabahensis sp. n. 
(8-1 1): 8. Habitus; 9. Halsschildprofil; 10. Aedeagusspitze, dorsal; 1 1. Aedeagusspitze, lateral. 



Länge 3,0, größte Breite 1,2. Kopf 0,6 lang, über die Augen gemessen 0,7 
breit. Halsschild 0.6 lang, 0,6 breit. Flügeldecken 1,8 lang, 1,2 gemeinsam breit. 

Färbung: Dunkelbraun. Fühler, Taster und Beine etwas heller braun. 

Kopf: Glänzend. Äußerst fein und verstreut punktiert. Behaarung dunkelbraun, 
sehr kräftig, etwas gebogen, halb abstehend, größtenteils nach hinten gerichtet, ziem- 
lich dicht. Fühler unauffällig behaart. 

Halsschild: Glänzend. Ziemlich kräftig, aber unterschiedlich und flach punktiert. 
In der Einschnürung sehr kräftig und dicht punktiert. Behaarung dunkelbraun, sehr 
kräftig, wenig gebogen, halb abstehend. Außerdem mit zahlreichen steifen Borsten. In 
der Einschnürung und vor der Einschnürung mit je einem feinen Längskiel. 

Flügeldecken: Glänzend. Fein und etwas unterschiedlich punktiert. Zwischen- 
räume meist etwas größer als die Punkte. Zur Spitze werden die Punkte kaum feiner. 
Die Zwischenräume werden kaum größer. Behaarung dunkelbraun, kräftig, ziemlich 
dicht, etwas gebogen, halb abstehend, nach hinten gerichtet. 



BESCHREIBUNG VON VIER NEUEN ARTEN DER GATTUNG DERARIMUS 497 

Beine unauffällig behaart. 

Beziehungen: Dem Derarimus fulvescens Uhmann aus Malaysia (Taiping) 
ähnlich, aber kleiner, Punktierung und Behaarung anders, Flügeldecken ohne lange 
Borsten. 

Derarimus sabahensis sp. n. Abb. 8-11 

Malaysia, Sabah. Babu. Pungpul Resort env., 24.6.-1.7.1996, 1 1 c vegetation debris and 
forest floor litter accumulated arround large trees near river, no collector, 7 Ex., Holotypus, 
6 Paratypen. 

Länge 3,0, größte Breite 1,3. Kopf 0,6 lang, über die Augen gemessen 0,7 
breit. Halsschild 0,8 lang, 0,7 breit. Flügeldecken 1,8 lang, 1,3 gemeinsam breit. 

Färbung: Dunkelbraun. Fühlerbasis uns Schenkel heller braun. Fühlerspitze, 
Taster, Schienen und Tarsen gelbbraun. 

Kopf: Glänzend. Punktur fein, flach, verstreut. Behaarung braun, kräftig 
gebogen, halb abstehend, in verschiedene Richtungen weisend. Fühler mit langer, 
kräftiger Behaarung. 

Halsschild: Glänzend. Fein und verstreut punktiert, in der Einschnürung 
runzelig. Behaarung braun, kräftig, lang, fast gerade, fast senkrecht abstehend. In der 
Mitte des Vorderrandes ausgehöhlt (nur bei den Männchen). 

Flügeldecken: Glänzend: Kräftig punktiert, Zwischenräume etwa so groß wie 
die Punkte. Zur Spitze werde die Punkte sehr viel feiner, die Zwischenräume sehr viel 
größer. Behaarung braun, kräftig, lang, steil und fast gerade abstehend. Neben der 
Naht, hinter dem Schildchen beiderseits etwas niedergedrückt. 

Beine unauffällig behaart. 

Beziehungen: Dem Derarimus ovipennis Uhmann aus W-Malaysia (Pahang) 
etwas ähnlich, aber mit deutlichen Schultern und größeren Augen. 

LITERATUR 

Bonadona, P. 1978. Les Tomoderini subendogés d'Afrique centrale et de l'Inde méridionale. 

Revue suisse de Zoologie 85 (3): 645-656. 
Sakai, M. 1986. Studies on Anthicidae of Japan, I. Transactions of the Shikoku Entomological 

Society 17 (4): 247-251. 
Uhmann, G. 1994. Südostasiatische Anthiciden aus dem Naturhistorischen Museum in Genf, 4. 

Revue suisse de Zoologie 101 (3): 655-676. 
Uhmann, G. 1996. Indo-australische Anthicidae im Naturhistorischen Museum in Genf. Revue 

suisse de Zoologie 103 (3): 737-748. 



Revue suisse de Zoologie 105 (3): 499-555; septembre 1998 



A taxonomic revision of the family Oncopodidae I. 
New genera and new species of Gnomulus Thorell 
(Opiliones, Laniatores) 

Jochen MARTENS 1 & Peter SCHWENDINGER 2 

1 Institut für Zoologie, Johannes Gutenberg-Universität Mainz, 
D-55099 Mainz, Germany. 

2 Institut für Zoologie und Limnologie, Universität Innsbruck, Technikerstr. 25, 
A-6020 Innsbruck, Austria. 



A taxonomic revision of the family Oncopodidae I. New genera and 
new species of Gnomulus Thorell (Opiliones, Laniatores). - The genera 
Oncopus Thorell, 1876 and Gnomulus Thorell, 1890 are rediagnosed; 
Pelitnus Thorell, 1891 is synonymised with Gnomulus because of inter- 
mediate forms and identical penis structure. Seventeen species described 
under Pelitnus are transferred to Gnomulus; generic placement in six of 
them (known only from females or juveniles) is provisional. Taxonomic 
characters of Oncopodidae and relationships within the Oncopodidae and 
with other families are discussed. Pelitnus thorelli Schwendinger, 1992 is a 
primary homonym; the species unduely described under this name is 
transferred to Gnomulus and renamed G baharu Schwendinger nom. n. 
Three new genera and thirteen new species of mostly small oncopodid 
opilionids are described. Palaeoncopus gen. n., with P. gunung sp. n., P. 
kerdil sp. n., P. katik sp. n. from Sumatra possess a short, distad-directed, 
non-expandable glans penis, which is considered plesiomorphic. Bianton- 
copus gen. n., with B. fuscus sp. n. from the Philippines, has a similar but 
expandable glans. This character is regarded as apomorphic. Seven new 
species are placed in Gnomulus, i. e. G crucifer sp. n., G maculatus sp. n., 
G coniceps sp. n., G. leyteensis sp. n., G laruticus sp. n., G. asli sp. n. and 
G hirsutus sp. n. Their penis morphology, with a short proximad-directed 
glans, is typical for most Oncopodidae (also present in Oncopus) and is 
considered derived from the Palaeoncopus-type. Caenoncopus gen. n., 
including C. tenuis sp. n., C. affinis sp. n. and C. cuspidatus (Schwen- 
dinger) comb. n. (transferred from Oncopus) from Sumatra, has a glans 
penis comprising a strongly elongated, proximad-directed stylus wrapped 
in a membraneous collar. This structure appears highly apomorphic and is 
possibly an extreme modification of the penis type in Gnomulus and 
Oncopus. Intermediate states of reduction of glans sclerites and enlarge- 
ment of stylus are present in G. crucifer sp. n. and G. maculatus sp. n.. 



Manuscript accepted 07.04. 1998 



500 J - MARTENS & P. SCHWENDINGER 



though rhey probably belong to a different evolutionary lineage. Four penis 
types are distinguished for the Oncopodidae; their evolution, generic 
significance and functional morphology are discussed. 

Key-words: Opiliones - Oncopodidae - Taxonomy - Penis morphology - 
Evolution - SE Asia. 



INTRODUCTION 

Since a revisionai study on the Oncopodidae of the Natural History Museum of 
Geneva (Schwendinger 1992), extensive and exceptionally rich new material has 
become available from the collections of various colleagues and ourselves. Several of 
the small specimens in this material are particularly interesting, as they look similar to 
the enigmatic Oncopus cuspidatus Schwendinger. which has no genital-morpholo- 
gical resemblance to other Oncopodidae or even to other opilionids previously 
known. After close examination, surprisingly they turned out to include not only close 
relatives of O. cuspidatus but also other forms with unexpected penis morphology. An 
account of this remarkable new material is given here; a thorough revision of the 
remaining taxa with new descriptions shall follow in subsequent papers. Although the 
newly available material contains plenty of exceptional forms, it nevertheless orig- 
inates from quite sporadic samplings and we believe that it only represents the tip of 
an iceberg. With more systematic sampling by sifting and soil extraction in all parts of 
Southeast Asia, a plethora of small and inconspicuous oncopodid taxa is expected to 
be discovered. 

Abbreviations and terms used in the text: CCD collection of C, Deeleman- 
Reinhold, Sparrenlaan; MAR collection of J. Martens, Mainz; MCSNG Museo Civico 
di Storia Naturale, Genova; MHNG Muséum d'histoire naturelle, Genève; SMF 
Senckenberg Museum. Frankfurt; ZMA Zoological Museum, University of Amster- 
dam; ZMT Zoological Museum, University of Turku. Body measurements refer to the 
length of the dorsal scutum (i.e. distance between anterior margin of carapace and 
posterior margin of abdominal part of dorsal scutum). Leg articles were measured on 
their dorsal side, from joint to joint. All measurements are given in mm. 



TAXONOMIC REMARKS 

Species concept 

We tried to find characters for distinctions and relationships between species in 
their penis morphology, but faced the problem of where to draw the boundaries in 
regard to the biospecies concept. Allopatric, morphologically similar and obviously 
closely related populations needed to be divided into "biospecies", without knowing 
whether or not reproductive isolation exists. As empirical data for such a grouping in 
Oncopodidae are non-existent (see Martens 1978 for Biantidae). we had to draw 
species boundaries in an arbitrar}- manner. Each seemingly allopatric population with 



NEW GENERA AND SPECIES OF ONCOPODIDAE 5Q] 



clear morphological distinctiveness is here regarded as a separate species. Due to the 
scattered and very localized nature of Oncopodidae collections (formerly caused by 
restricted accessability, today by habitat destruction), individual finds may show 
morphological distinctiveness, which does not correspond with species identity in the 
sense of the biospecies concept. In addition, geographical variation in Oncopodidae 
remains largely unclear. Therefore the species concept necessarily used here is close 
to the phylogenetical species concept, which clusters diagnosable populations into so- 
called "phylospecies" (Zink 1997). 

Traditional system and generic limits 

Thorell (1876) described the subfamily Oncopodinae (under the family 
Cosmetoidae Koch) and later (1890) upgraded them to family rank. Within this group 
he successively distinguished three genera: Oncopus (Thorell, 1876), Gnomulus 
(Thorell, 1890) and Pelitnus (Thorell, 1891), which remained valid until the present 
day. The distinction between these genera is essentially based on tarsal formula 
(Oncopus 1-1-1-1, Gnomulus and Pelitnus 2-2-3-3) and presence {Pelitnus) or 
absence {Gnomulus) of an oblique, strong, triangular eye tubercle (Thorell 1891: 93 
- "Oculi basi tuberculi transversi fortis trianguli impositi."). The latter character, 
however, shows all transitions from a plane interocular area to a rounded hump and an 
acutely pointed prorect or erect eye tubercle in both nominal genera. The interocular 
area is sexually dimorphic in some species, with a low eye tubercle present in 9 ? 
and absent in SS. This was shown for G. lannaianus (sub Pelitnus lannaianus, 
Schwendinger 1992) and also occurs in G. sumatranus (in preparation). Therefore 
the traditional distinction between Gnomulus and Pelitnus cannot be maintained. 
Penis morphology in both genera (first illustrated for Gnomulus by Loman 1903: fig. 
5f) is of the same type. A detailed account of this shall be given in our next paper. 

In contemporary taxonomy of arthropods ó* genitalia are regarded to be among 
the most informative characters explaining relationships between taxa. Within the last 
decades this view became generally accepted and brought forth significant changes to 
high level systematics of opilionids. On the base of the genital morphology of S S, 
the family Fissiphallidae was established recently (Martens 1988) and the traditional 
suborders Cyphophthalmi and Palpatores were united to one suborder Cyphopal- 
patores (Martens 1980, 1986). Applying the same criteria to the family Onco- 
podidae, we place Pelitnus in synonymy with Gnomulus, as was already suggested by 
Loman (1902: 183). Three new genera with penis morphology distinctly different 
from Gnomulus and Oncopus are established in the following. 

Definition of the family Oncopodidae 

Oncopodid harvestmen are readily distinguishable by external characters, but 
most of these seem to be plesiomorphic for opilionids and are therefore not appro- 
priate to define the family. This holds true for the following characters in particular: 



502 J - MARTENS & P. SCHWENDINGER 

extensive dorsal scutum (carapace and abdominal tergites fused); low number (1-3) of 
tarsal articles; restricted unidirectional articulation of the legs and pedipalps. The 
large, fused dorsal scutum clearly distinguishes the Oncopodidae from other 
Laniatores, but similar structures are also found in the Sironoidea and the Trogulidae, 
which are likewise slow-moving, soil-dwelling opilionids. All three taxa are addition- 
ally characterized by a low number of tarsal articles (up to 2 in Sironoidea S S , up to 
4 in Trogulidae and up to 3 in Oncopodidae). Obviously the state of both characters is 
correlated with the mode of life of these cryptic animals. In Sironoidea large dorsal 
scutum and few tarsalia are considered plesiomorphic, whereas in the taxonomically 
distant Trogulidae the same most likely represent apomorphic reversals. Though 
oncopodids already possess clearly derived genital characters (e.g. hemolymph- 
pressure penis), their large scutum and low number of tarsalia are difficult to evaluate. 
We assume that they represent plesiomorphies. 

Only the following three character states are presently regarded as autapo- 
morphic of the Oncopodidae, defining the family as a monophyletic group: 

1. Glans penis with lateral sclerites fused by an intermediate (median) plate 
(Figs 1, 134). In other families, symmetrical glans structures are not interconnected 
and can be moved independently from each other (Martens 1986). 

2. Ovipositor laterally compressed, not dorso-ventrally flattened or circular in 
cross-section (Martens et al. 1981). This trait holds true not only for Oncopus 
(Martens et al. 1981), but was also found in Gnomulus, Biantoncopus gen. n., 
Palaeoncopus gen. n. and Caenoncopus gen. n. In the smallest representatives, 
however, ovipositor cross-sections generally tend to be more roundish. 

3. Cuticular appendages (paired or unpaired) on the carapace and the first 
abdominal tergite, respectively, form a small "bridge". This character is found only in 
Oncopodidae (Silhavy 1960). 

Regarding the scarcity of derived characters identified by now, the taxonomic 
position of the family Oncopodidae is again open to question. By no means can we 
presently trace character states that may warrant the status of a superfamily or a 
similarly high-ranking taxon for the Oncopodidae alone, as was suggested earlier 
(Silhavy 1960, Martens et al. 1981). At the present state of knowledge no sister 
group can be identified for the Oncopodidae. 



DESCRIPTIONS 

Caenoncopus gen. n. 

Diagnosis: Distinguished by penis with dorso-ventrally depressed truncus, 
basally bilobed, subbasally not constricted; subapical glans large, composed of a 
strongly elongated, cylindrical stylus proximally enclosed in a membraneous collar; 
tip of stylus asymmetrical; glans folded proximad in resting position, lying in a 
shallow trench on the dorsal surface of the truncus. Body small without elevated 
scutal areas; interocular area developed as a round hump, not abruptly separated from 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



503 




Fig. 1 

General scheme of an oncopodid penis of the GnomuluslOncopus type; ventral (a) and lateral 
(b) view of distal part. - Apex of truncus (ap); two lateral sclerites (Is); median plate (mp); 
membraneous socket (ms); two membraneous tubes (mt); subapical setae (ss); stylus (st). 



low thoracic area; chelicerae small, weak, with a pronounced dorso-distal and a small 
ventro-median hump on proximal article, no modifications on cheliceral hand; 
pedipalps with small ventral processes on proximal femur and trochanter; ventral coxa 
II with anterio-proximal process; legs 3142, tarsal formula 1-1-2-2 or 1-1-3-3. Exter- 
nal sexual dimorphism in shape of palpal trochanter or absent. 

Etymology: Greek: kainos = new, young; oncos = swelling; podos = foot; male 
gender. The Latinized name (correctly spelled "Caenooncopus", one "o" omitted to 
make it more euphonious) refers to the highly derived genital morphology of this 
genus and to its relationship with Oncopus. 

Type species: Oncopus cuspidatus Schwendinger. 

Species account and distribution: Three apparently allopatric species, i.e. C. 
cuspidatus comb, n., C. tenuis sp. n., C. affinis sp. n., are known from Sumatra. The 
latter two species occur in close proximity and are also closest relatives. 



504 



J. MARTENS & P. SCHWENDINGER 




Fig. 2 

Distribution of Oncopodidae in the Malay Peninsula and Sumatra. Only localities of species 
treated in this paper are shown. - a) Maxwell Hill (Gnomulus laruticus sp. n.); b) Chenderiang 
(G asli sp. n.); c) road to Genting Highlands (G. hirsutus sp. n.); d) Templer Park (G. hirsutus 
sp. n.); e) Ulu Gombak (G hirsutus sp. n.); f) Ketambe {Caenoncopus cuspidatus, Palaeoncopus 
kerdil sp. n.); g) Bukit Lawang (C. cuspidatus); h) Bohorok [Langkat Reserve] (C. cuspidatus, 
P. katik sp. n.); i) road Brastagi - Sibolangit [Deli Serdang] (C. cuspidatus); k) Panti (C affinis 
sp. n.); 1) road Lubuksikaping - Panti (C affinis sp. n.); m) Palopo Nature Reserve (C. tenuis 
sp. n.); n) 5 km SE of Payakumbuh (C. tenuis sp. n.); o) Mt. Singgalang (P. gunung sp. n.). 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



505 




Fig. 3 

Distribution of Oncopodidae in the Philippines. Only localities of species treated in this paper 
are shown. - a) Sagada (Gnomulus maculatus sp. n.); b) Baguio, Cristal Cave (G coniceps sp. 
n.); c) Mt. Santo Thomas (G. crucifer sp. n.); d) Puerto Galera (G. maculatus sp. n.), doubtful 
record; e) Lake Danao, Leyte (Biantoncopus fuscus sp. n.); f) Visca N of Baybay (B. fuscus sp. 
n., G. leyteensis sp. n.). 



506 



J. MARTENS & P. SCHWENDINGER 




Fig. 4. Scanning electron micrographs showing external characters of Oncopodidae species. - a) 
Palaeoncopus gumtng sp. n., left 9 palpal trochanter and base of femur, prolateral view; b) 
Caenoncopus tenuis sp. n.. tarsus and distal part of metatarsus IV; c-f) C. cuspidatus 
(Schwendinger), tarsus and distal part of metatarsus of leg II (c) and leg IV (d) , claws and 
arolium on leg IV of juvenile (e-f). 



NEW GENERA AND SPECIES OF ONCOPODIDAE 5Q7 



Caenoncopus cuspidatus (Schwendinger) comb. n. Figs 4c-f, 5a-d, 6-8 

Oncopus cuspidatus Schwendinger, 1992: 190-192, figs. 67-80. Description of 3 . 

Types: SUMATRA, Northern Sumatra Province, Langkat, Bukit Lawang Nature 
Reserve, 180 m, S holotype (MHNG Sum-85/49). - Deli Serdang, north of Brastagi, 1400 m, 
3 paratype (MHNG Sum-85/47); both leg. B. Hauser, 8.-20.XI.1985. 

New material: SUMATRA, Aceh Province, Mt. Leuser National Park, Ketambe 
Research Station, 300-500 m, 23.-30.XI.1989, 4 3, 13 9; 800 m, 28.XI.1989, 1 9; all leg. D. 
Agosti, I. Lobi & D. Burckhardt (MCSNG, MHNG). - North Sumatra Province, Langkat, Bukit 
Lawang Nature Reserve, 1 1 .-12.X. 1990, 1 3, 1 9; leg. A. Riedel (MAR). - Bohorok. 
7.VIII.Ï982, Ì 3, 1, 9, 13.XI.1983, 1 9, 30.XII.1993, 2 9; all leg. C. Deeleman-Reinhold & P. 
Deeleman (CCD). - Langkat, Bukit Lawang Nature Reserve, Bohorok river, 5.VII.1984, 1 3 , 2 
9; leg. J. Robert (MHNG). - Forest 7 km north of Brastagi, 1500 m, 2.XII.1989, 1 3, 6 9; all 
leg. D. Agosti, I. Lobi & D. Burckhardt (MHNG). 

Diagnosis (extended): Relatively large species; scuta smooth, with charac- 
teristic pattern; palpal femur with ventro-basal process (Figs 7, 8); palpal trochanter 
sexually dimorphic, with distinct ventral process only in 9 9 (Fig. 7); tarsal formula 
1-1-2-2; genital operculum with anterior hump (Fig. 6). Truncus penis with rounded 
apex drawn into an obtuse angle, bearing two widely separated rows of subapical 
setae on each side; stylus very long, almost reaching base of truncus, wrapped in a 
membraneous collar almost throughout its entire length; apex of stylus free, 
corkscrew-shaped (Fig 5a-d, Schwendinger 1992: figs 74-80). 

Remark: The small proximal tarsalia on the posterior legs, partly overlapped 
by the terminal edge of the metatarsus (Fig. 4d), were not recognized in the original 
description. Therefore C. cuspidatus was placed in the genus Oncopus. However, this 
placement was provisional. The author was aware of the uniqueness of the species, 
but decided not to establish a new genus because of the sparse material (2 3) 
available (Schwendinger 1992: 197). 

Variation: Measurements range: 3: body length 3.65-4.51 (x = 4.09, SD = 

0.274), width 2.08-2.68 (x = 2.44, SD = 0.193), n = 10; 9 : body length 3.59-4.39 (x = 

4.05. SD = 0.238), width 2.07-2.85 (x = 2.40, SD = 0.187), n = 27. Body size 

gradually decreases in between the three populations. 

Population: mean 3 body length (SD) mean 9 body length(SD): 

Ketambe 4.37 (0.089), n = 4 4.24 (0.090), n =14 

Bukit Lawang and env. 4.02 (0.056), n = 4 3.98 (0.052), n = 7 

Brastagi and environs 3.67 (0.015), n = 2 3.68 (0.049), n = 6 

Distribution (Fig. 2): Known from the southern part of Aceh Province and 
from the northern part of North Sumatra Province. The specimens examined originate 
from three separate areas: 1. Ketambe; 2. Bukit Lawang Reserve, Bohorok, Langkat; 
3. Brastagi, Deli Serdang. 

Bionomics: The specimens were collected from the leaf litter of a lowland rain 
forest and a montane rain forest with various degrees of disturbance. 

Caenoncopus tenuis sp. n. Figs 4b, 9-17 

Material: SUMATRA, West Sumatra Province , 5 km southeast of Payakumbuh. 600 
m, 3 holotype (MHNG), 9<î,4 9 paratypes (MAR, MCSNG, MHNG), 20.-21. XI. 1989. - 
Palopo Nature Reserve, north of Bukittinggi, 900 m, 1 3 paratype (MHNG), 18.-20.XI. 1989. 
All specimens leg. D. Agosti, I. Lobi & D. Burckhardt. 



508 



J. MARTENS & P. SCHWENDINGER 




Fig. 5 

Scanning electron micrographs of penis of Caenoncopus cuspidatus (Schwendinger): total 
penis, dorsal (a) and lateral (b) view; apex of glans, fronto-dorsal (c) and dorso-distal view (d). 




Figs 6-8 

Caenoncopus cuspidatus (Schwendinger); 6 holotype (8), 9 (6, 7). - Genital operculum, 
latero-ventral view (6); left palp, retrolateral view (7, 8). - Scale lines 0.5 mm. 



Etymology: Latin: tenuis = small, thin, delicate. 

Diagnosis: Closely related to C. cuspidatus, distinguished by smaller size and 
less extensive dark patches on dorsal and ventral scuta; distinct ventral process on 
palpal trochanter also present in ââ; tarsal formula 1-1-3-3; genital operculum 
without anterior hump. Truncus penis with broadly truncate apex carrying only one 
short row of subapical lateral setae on each side; stylus shorter, not reaching base of 
truncus; membraneous collar enclosing only basal half of stylus; apex of stylus 
different in shape (Figs 9-14). 

Description: 6 (holotype). Coloration: Body mostly light amber, except for: 
dark reticulation on carapace, dark margin and transverse bands on abdominal part of 
dorsal scutum (Fig. 17a), widely separated lateral pairs of dark transverse patches on 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



509 




Figs 9-16 

Caenoncopus tenuis sp. n.; S holotype (9-12), 6 paratype (13-14), 9 paratype (15-16). - Penis, 
dorsal (9) and lateral view (10); apex of stylus, dorsal (11), lateral (12), ventral (13) and contra- 
lateral view (14); left palp, retrolateral view (15); left chelicera, retrolateral view (16). - Scale 
lines 0.05 mm (11-14), 0.5 mm (9-10, 15-16). 



ventral scutum (Fig. 17b). Genital operculum, process on coxae II and proximal zone 
of tibiae I, II and metatarsus II darkened. 

Carapace short, interocular area a low hump; dorsal and ventral scuta smooth, 
without furrows or elevations (Figs 17a, c). Distinct antero- and posterio-proximal 
processes on coxa II, two smaller sub-proximal and median processes on coxa I. 
Genital operculum relatively large (Fig. 17b). 

Chelicerae (Fig. 16) small; proximal article with distinct dorso-distal boss and 
ventro-median process; hand slender, without process. 

Palps (Fig. 15) with basally wide ventral process on proximal femur; other 
articles unarmed. 

Legs 3142, tarsal formula 1-1-3-3. 

Penis (Figs 9-14): truncus depressed, bearing needle-like subapical setae in one 
group on each side. Membraneous socket of glans penis distally round; tube-like 
stylus more than half as long as truncus, resting in a shallow trench on the dorsal 
surface of truncus. Basal half of stylus covered by membraneous collar; crescent- 
shaped apex standing at right angles to axis of stylus. 

9 . As the ô ; no external sexual dimorphism discernible. 



510 



J. MARTENS & P. SCHWENDINGER 






Fig. 17 
Caenoncopus tenuis sp. n.. S holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale 
line 1 mm. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 5 1 ] 

Measurements (o\ in brackets $): body 2.32 (2.55) long, 1.50 (1.56) wide; 
carapace region 0.61 (0.64) long, 0.88 (0.93) wide. - Palp and legs: 



Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 0.23 (0.27) 


0.38 (0.41) 


0.32 (0.34) 


0.24 (0.24) 


- - 


0.41 (0.44) 


1.58(1.70) 


Leg I 0.23 (0.23) 


0.69 (0.69) 


0.43 (0.46) 


0.35 (0.36) 


0.71 (0.72) 


0.06 (0.06) 


2.47 (2.52) 


Leg II 0.27(0.31) 


0.92 (0.94) 


0.56(0.61) 


0.61 (0.61) 


1.09(1.09) 


0.06 (0.06) 


3.51 (3.62) 


Leg III 0.23 (0.23) 


0.53 (0.54) 


0.43 (0.44) 


0.37 (0.37) 


0.72 (0.72) 


0.06 (0.07) 


2.34(2.37) 


Leg IV 0.24 (0.26) 


0.85 (0.87) 


0.56 (0.55) 


0.66 (0.69) 


0.98 (1.00) 


0.06 (0.07) 


3.35 (3.44) 



Variation: Measurements of body length/width range: â 2.26-2.52/1.38-1.58 
(n = 12); 9 2.46-2.55/1.50-1.57 (n = 4). Some SS (including the holotype) have a 
smaller ventro-basal process on the palpal femur than shown in Fig. 15. In several S S 
the carapace is dorsally less distinctly saddle-shaped than in the holotype (Fig. 17c). 

Bionomics: The specimens were collected by sifting leaf litter and soil in 
abandoned rubber and coffee plantations (SE of Payakumbuh) and in a steeply 
sloping secondary forest (Palopo Nature Reserve). 

Caenoncopus affinis sp. n. Figs 18-26 

Material: SUMATRA, West Sumatra Province. Panti, 250 m, S holotype (MHNG), 2 
6,2 9 paratypes (MAR, MHNG), 19.XI.1989; all leg. D. Agosti, I. Lobi & D. Burckhardt. - 
Road from Lubuksikaping to Panti, ca. 700 m, 26.X.1991, 1 ? paratype, leg. A. Riedel (MAR). 

Etymology: Latin: affinis = related, neighbouring. The species name points to a close 
relationship with C. tenuis sp. n. 

Diagnosis: Very close to C. tenuis sp. n. but distinguished by posterior carapace 
low; genital operculum with straight lateral margins; dark lateral patches on ventral 
scutum larger, interconnected posteriorly; ventral processes on palpal trochanter and 
femur more pronounced. Membraneous socket of glans penis more pointed distally; 
stylus shorter, less strongly sigmoid distally, apex different in shape (Figs 18-23). 

Description: S (holotype). Coloration: Body mostly light amber, except dark 
reticulation on' carapace, dark margin and transverse bands on abdominal part of 
dorsal scutum (Fig. 26a); lateral pairs of dark transverse patches on ventral scutum 
interconnected posteriorly (Fig. 26b). Genital operculum, process on coxae II, tibia II. 
proximal 2/3 of metatarsus II and proximal 1/2 of metatarsus I darkened. 

Carapace short, interocular area elevated above rest of carapace; abdominal 
parts of dorsal and ventral scuta^smooth, without furrows or elevations (Figs 26a, c). 
Ventral prosoma with distinct conical process on proximal coxa II, smaller knob- 
shaped processes on sub-proximal and median coxa I and on palpal coxa. Genital 
operculum with straight lateral margins, almost trapezoidal in shape (Fig. 26b). 

Chelicerae (Fig. 25) small; proximal article with dorso-distal boss and low 
ventro-median process; hand slender. 

Palps (Fig. 24) with distinct ventral process on proximal femur and on distal 
trochanter. 

Legs 3142, tarsal formula 1-1-3-3. 



512 



J. MARTENS & P. SCHWENDINGER 



Penis (Figs 18-23): truncus depressed with straight distal margin, bearing a 
subapical group of needle-like setae on each side. Membraneous socket of glans 
distally tapering; tube-like stylus less than half as long as truncus, slightly bent below 
apex, basal half covered by membraneous collar; apex of stylus crescent-shaped, 
perpendicular to axis. 

9 . As the <5; no external sexual dimorphism discernible. 

Measurements (o\ in brackets 2): body 3.04 (3.20) long, 1.94 (2.06) wide; 
carapace region 0.75 (0.75) long, 1.14 (1.14) wide. - Palp and legs: 



Palp 
Legi 

Leg II 
Leg III 
Les IV 



Tr 

0.37 (0.38) 
0.29(0.31) 
0.34 (0.38) 
0.31 (0.29) 
0.34 (0.39) 



Fe 



Pa 



0.53 (0.52) 0.44 (0.43) 
0.87(0.81) 0.60(0.58) 
1.16(1.08) 0.75(0.72) 
0.69 (0.66) 0.59 (0.58) 
1.07(1.01) 0.78(0.76) 



Ti 

0.35 (0.34) 
0.47 (0.46) 
0.78 (0.75) 
0.47 (0.47) 
0.85 (0.82) 



Mt 

0.92 (0.90) 
1.40(1.31) 
0.99 (0.95) 
1.34(1.28) 



Ta 



Total 



0.55(0.56) 2.24(2.23) 

0.05(0.06) 3.20(3.12) 

0.08(0.11) 4.51 (4.35) 

0.06(0.08) 3.11 (3.03) 

0.06(0.08) 4.44(4.34) 



Variation: Measurements of body length/width range: S 3.04-3.25/1.90-2.00 
(n = 3); 9 3.20-3.24/1.96-2.12 (n = 3). One 6 paratype with interocular tubercle 
lower than in Fig. 26c. 




Figs 18-25 

Caenoncopus affinis sp. n.; S holotype (18-21), â paratype (22-23), $ paratype (22-25). - 
Penis, dorsal (18) and lateral view (19); apex of stylus, dorsal (20, 22) and lateral view (21, 23); 
left palp, retrolateral view (24); left chelicera, retrolateral view (25). - Scale lines 0.05 mm (20- 
23), 0.5 mm (18-19, 24-25). 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



513 






Fig. 26 

Caenoncopus affinis sp. n., 6 holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale 
line 1 mm. 



514 J- MARTENS & P. SCHWENDINGER 

Bionomics: The specimens were collected by sifting vegetational debris in a 
lowland swamp forest (near Panti) and in a lowland rain forest (between 
Lubuksikaping and Panti). 

Palaeoncopus gen. n. 

Diagnosis: Truncus penis very slender, cylindrical, basally truncate and sub- 
basally constricted; short subapical glans distad-directed, composed of a short thin 
stylus surrounded by a pair of forceps-like lateral sclerites interconnected by a median 
plate; pair of membraneous tubes absent; glans not expandable. Body small; inter- 
ocular area a round hump, abruptly separated from thoracic area carrying a pair of 
humps. Anterior margin of abdominal part of dorsal scutum with a pair of wide lobes, 
more or less distinctly separated from carapace hind-margin by a membraneous zone. 
Scutal areas distinctly elevated, medially subdivided by a deep longitudinal furrow. 
Chelicerae small, weak, without modifications apart from dorso-distal hump on prox- 
imal article. Palpal femur with a distinct ventro-basal process; palpal trochanter with a 
large, multilobate ventral process and a conical prodorsal process. Legs 1324, tarsal 
formula 1-1-3-3. External sexual dimorphism unknown. 

Etymology: Greek: palaios = old; oncos = swelling; podos = foot; male gender. 
The Latinized name (correctly spelled "Palaeooncopus", one "o" omitted) refers to the 
primitive genital morphology of this genus and to its relationship with Oncopus. 

Type species: Palaeoncopus gunung sp. n. 

Species account and distribution: Including three species from Sumatra, i.e. 
P. gunung sp. n., P. kerdil sp. n., P. katik sp. n. 

Relationships: Palaeoncopus gen. n. represents the most basic group within 
the Oncopodidae. The three species are closely related to each other and are known 
from distinctly separated localities. Presently, they are all to be considered morpho- 
species. 

Palaeoncopus gunung sp. n. Figs 4a, 27-39 

Material: SUMATRA, West Sumatra Province, Bukittinggi, Mt. Singgalang, 2100- 
2600 m, 3 holotype (MHNG), 1 â, 4 9 paratypes (MAR, MHNG); 16.X.1990, leg. A. Riedel. 
Etymology: Malay and Indonesian: gunung (gunong) = mountain. Noun in apposition. 

Diagnosis: Resembling Caenoncopus affinis sp. n. in body size, proportions 
and tarsal formula but thoracic area and scutal areas distinctly elevated and sub- 
divided by a deep median furrow; no ventral process on proximal article of chelicera; 
palpal trochanter with a large, trilobed ventral process and a conical prodorsal pro- 
cess; anterio-proximal process on ventral coxa III present; legs 1324. Penis with short 
distad-directed glans bearing forceps-like lateral sclerites connected by a median plate 
(Figs 27-32). 

Description: â (holotype). Coloration: Body mostly light amber; dark reticu- 
lations on carapace and dark, medially broken transversal bands on dorsal (Fig. 39a) 
and ventral scutal areas (Fig. 39b), indistinct in ventral areas III-V. Limbs mostly dark 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



515 




Figs 27-38 

Palaeoncopus gunung sp. n.; 6 holotype (27-30, 34), 6 paratype (31-32, 35), 9 paratypes (33, 
36-38). - Penis, dorsal (27) and lateral view (28); apex of penis, dorsal (29, 31) and lateral view 
(30, 32), glans slightly expanded (29-30); left palp, retrolateral view (33); left palp, trochanter 
and femur, retrolateral view (34-37); left chelicera, retrolateral view (38). - Scale lines 0.5 mm 
(27-28, 33-38), 0.1 mm (29-32). 



516 



J. MARTENS & P. SCHWENDINGER 






Fig. 39 
Palaeoncopus gunung sp. n., 3 holotype, body, dorsal (a), ventral (b) and lateral view (c). 
Scale line 1 mm. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 5 \ 7 

brown, except for light orange cheliceral hand and tarsi, distal tibiae and metatarsi of 
legs and pedipalps. 

Carapace short, narrow depression separating hump on interocular area from 
pair of humps on thoracic area; dorsal and ventral scutal areas distinctly elevated, 
dorsal areas separated from each other by deep transverse furrows and by a 
longitudinal median furrow (Figs 39a, c). Conical anterio-proximal processes on 
ventral coxae II and III, knob-shaped central ones on median coxa I and palpal coxa. 
Genital operculum wide, subtriangular (Fig. 39b). 

Chelicerae (Fig. 38) small; proximal article with dorso-distal boss but without 
ventro-median process; hand slender. 

Palps (Fig. 33-37): femur with conical ventral process; trochanter with large 
trilobate ventral process and small conical dorsal process. 

Legs 1324, tarsal formula 1-1-3-3. 

Penis (Figs 27-32): truncus slender, its distal margin arched and medially 
invaginated, forming two distinct lobes; strong, curved subapical setae in two 
distinctly separated groups on each side of truncus. Glans pointing distad, rising from 
a long, basally narrow membraneous socket; lateral sclerites tapering, with furrows on 
lower face. Median plate long, triangular; stylus slender. 

2 . As the â; no external sexual dimorphism discernible. 

Measurements (o\ in brackets ?): body 2.87 (2.94) long, 1.90 (1.99) wide; 
carapace region 0.75 (0.69) long, 1.11 (1.10) wide. - Palp and legs: 

Tr Fe Pa Ti Mt Ta Total 

Palp 0.34(0.32)0.46(0.42)0.42(0.40)0.29(0.26)- - 0.53(0.50)2.04(1.90) 

Legi 0.31(0.29) 0.75(0.75) 0.49(0.50) 0.46(0.46) 0.85(0.85) 0.08(0.08) 2.94(2.93) 

Legll 0.38(0.34) 1.05(1.04)0.63(0.64) 0.76(0.75) 1.30(1.28) 0.12(0.11) 4.24(4.16) 

Leg III 0.29(0.29) 0.70(0.67) 0.50(0.53) 0.50(0.49) 0.99(0.98) 0.09(0.08) 3.07(3.04) 

Leg IV 0.32(0.31) 0.98(0.95) 0.67(0.69) 0.82(0.79) 1.39(1.36) 0.11(0.09)4.29(4.19) 

Variation: Measurements of body length/width range: S 2.87/1.79-1.90 
(n = 2); 9 2.94-2.97/1.94-2.03 (n = 4). Variation in the shape of the ventral process 
on palpal trochanter (Figs 33-37) does not show sexual dimorphism. 

Bionomics: The specimens were sifted from leaf litter in a montane forest. 

Palaeoncopus kerdil sp. n. Figs 40-46 

Material: SUMATRA, Aceh Province, Mt. Leuser National Park, Ketambe Research 
Station, 300-500 m, 23.-30.XI. 1989, S holotype and 9 paratype; leg. D. Agosti. I. Lobi & 
D. Burckhardt (MHNG). 

Etymology: Malay and Indonesian: kerdil = small, dwarfish. 

Diagnosis: Close to P. gunung sp. n., distinguished by smaller body size; tho- 
racic area of carapace more elevated; wider gap between carapace and abdominal part 
of dorsal scutum; dark pattern on ventral scutum different; ventral process on palpal 
trochanter bilobed. Penis with narrower, truncate apex; short glans more remote from 
apex; lateral sclerites more strongly bent and closer to each other; membraneous 
socket at base of glans shorter, ovoid (Figs 40-43). 



518 



J. MARTENS & P. SCHWENDINGER 




Figs 40-45 

Palaeoncopus kerdil sp. n.; S holotype (40-43), 9 paratype (44-45). - Penis, dorsal (40) and 
lateral view (41); apex of penis, dorsal (42) and lateral view (43); left chelicera, retrolateral 
view (44); left palp, retrolateral view (45). - Scale lines 0.5 mm (40-41, 44-45), 0.1 mm 
(42-43). 



Description: S (holotype). Coloration: Body light amber; dark reticulations on 
carapace, abdominal part of dorsal scutum with dark margin and medially broken 
transversal bands on elevated areas (Fig. 46a); dark transverse bands on ventral scutal 
areas medially indistinct, not broken (Fig. 46b). Proximal leg articles slightly 
darkened; pedipalps, chelicerae and distal leg metatarsi and tarsi light orange. 

Carapace short, hump-shaped interocular area low, separated from thoracic 
area (with pair of humps) by a narrow depression; dorsal scutal areas distinctly 
elevated (Fig. 46c), separated from each other and medially by deep furrows (Fig. 
46a, c). Ventral scutum with indistinctly elevated areas and lobes behind spiracles. 
Ventral coxae II and III with distinct anterio-proximal processes, coxa II also with 
posterio-proximal one, smaller knob-shaped processes on central coxa I and palpal 
coxa. Genital operculum widely subtriangular (Fig. 46b). 

Chelicerae (Fig. 44) small; proximal article with moderately wide dorso-distal 
boss, ventral process indistinct; hand slender. 

Palps (Fig. 45): femur with conical ventrobasal process; trochanter with large 
bilobed ventral and small conical dorsal process. 

Legs 1324, tarsal formula 1-1-3-3. 

Penis (Figs 40-43): truncus slender, apex narrow, distal margin slightly 
indented; subapical setae in two widely separated groups on each side. Glans short. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



519 






Fig. 46 

Palaeoncopus kerdil sp. n.. 6 holotype, body, dorsal (a), ventral (b) and lateral view (c). 
Scale line 1 mm. 



520 J - MARTENS & P. SCHWENDINGER 

not reaching apex of penis, rising from a fairly short, ovoid membraneous socket; 
lateral sclerites distinctly bent, tapering, with furrows on lower face; median plate 
rounded, its lateral margins hidden under lateral sclerites; stylus slender. 
9 . As the S ; no external sexual dimorphism evident. 

Measurements (<?, in brackets $): body 1.72 (1.88) long, 1.12 (1.19) wide; 
carapace region 0.46 (0.47) long, 0.63 (0.71) wide. - Palp and legs: 

Tr Fe Pa Ti Mt Ta Total 

Palp 0.18(0.21)0.24(0.24)0.24(0.24)0.14(0.14)- - 0.31(0.32)1.11(1.15) 

Legi 0.16(0.17) 0.44(0.46) 0.29(0.33) 0.24(0.24) 0.46(0.50) 0.06(0.07) 1.65(1.77) 

Leg II 0.20(0.21) 0.63(0.64) 0.40(0.41) 0.37(0.38) 0.78(0.77) 0.08(0.08) 2.46(2.49) 

Leg III 0.17(0.18) 0.39(0.40) 0.32(0.34) 0.26(0.27) 0.56(0.56) 0.05(0.07) 1.75(1.82) 

Leg IV 0.20(0.20) 0.56(0.58) 0.43(0.43) 0.43(0.46) 0.81(0.82) 0.07(0.07) 2.50(2.56) 

Bionomics: The animals were collected by sifting leaf litter in a lowland 
dipterocarp forest. 



Palaeoncopus katik sp. n. Figs 47-57 

Material: SUMATRA, North Sumatra Province, Langkat, Bukit Lawang Nature 
Reserve, Bohorok River, 5.VII.1984, 6 holotype and 3 paratype; leg. J. Robert (MHNG). 
Etymology: Malay and Indonesian: katik = small, stunted. 

Diagnosis: Very similar to P. kerdil sp. n. but larger, eye tubercle (in dorsal 
view) closer to anterior carapace margin, anterior borders of abdominal part of dorsal 
scutum less arched, legs 1342. Apex of truncus penis arched, bearing two indistinctly 
separated groups of subapical setae on each side; membraneous socket of glans 
constricted proximally, lateral sclerites of glans distally wider (Figs 47-52). 

Description: S (holotype). Body coloration as in P. kerdil sp. n. except for 
darker palpal femora. 

Carapace with domed interocular area close to anterior margin, separated from 
pair of humps on thoracic area by a narrow depression; wide membraneous zone 
between carapace and abdominal part of dorsal scutum (Figs 55-57); dorsal scutal 
areas elevated, separated by deep furrows; ventral scutal areas moderately elevated. 
Processes on ventral coxae I, II, III and palps. Genital operculum wide; area behind 
spiracles drawn into lobes. 

Chelicerae (Fig. 53) small, with dorso-distal boss on proximal article. 

Palps (Fig. 54) with conical ventrobasal process on femur and large bilobed 
ventral plus small conical dorsal process on trochanter. 

Legs 1342, tarsal formula 1-1-3-3. 

Penis (Figs 47-52): truncus slender, apex slightly widened, distal margin 
medially invaginated, forming two small lobes; subapical setae in two indistinctly 
separated groups on each side. Glans short; membraneous socket ovoid, proximally 
narrow; lateral sclerites strongly bent, distally wide, with furrows on lower face; 
median plate rounded, mostly hidden under lateral sclerites; stylus slender. 

?. Unknown. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



521 




Figs 47-57 

Palaeoncopus katik sp. n.; 6 holotype (47-50, 55-56), S paratype (51-54, 57). - Penis, dorsal 
(47) and lateral view (48); apex of penis, dorsal (49, 51) and lateral view (50, 52); left 
chelicera, retrolateral view (53); left palp, retrolateral view (54); anterior part of body, lateral 
(55) and dorsal view (56-57). - Scale lines 0.5 mm (47-48, 53-57), 0.1 mm (49-52). 



Measurements (a): body 1.85 long, 1.24 wide; carapace 0.50 long, 0.72 wide. 



- Palp < 


ind legs: 
















Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.20 


0.26 


0.25 


0.15 


- 


0.36 


1.22 


Legi 


0.17 


0.50 


0.37 


0.27 


0.59 


0.05 


1.95 


Leg II 


0.23 


0.75 


0.47 


0.46 


0.92 


0.07 


2.90 


Leg III 


0.20 


0.44 


0.37 


0.31 


0.64 


0.05 


2.01 


Leg IV 


0.21 


0.66 


0.47 


0.52 


0.92 


0.08 


2.86 



Variation: The â paratype has body length 1.86, width 1.24. 

Relationships: This species is very similar to P. kerdil sp. n. Minor but consist- 
ent differences in external and genital morphology appear adequate for a species dis- 
crimination. 



522 J - MARTENS & P. SCHWENDINGER 

Biantoncopus gen. n. 

Diagnosis: Penis with stout cylindrical truncus, basally truncate, subbasally 
constricted; subapical glans distad-directed, composed of a short thin stylus sur- 
rounded by a median plate and a pair of lateral sclerites with apices curved away from 
the truncus; large pair of membraneous tubes present; parts of glans expandable. Body 
small, scutal areas low; eye tubercle conical. Chelicerae small, without modifications 
apart from dorso-distal hump on proximal article. Palpal femur with distinct ventro- 
basal process; palpal trochanter with large ventral process, without dorsal process. 
Legs 1324, tarsal formula 2-2-3-3. External sexual dimorphism unknown. 

Etymology: The genus name refers to similarities in penis morphology with the 
genus Biantes (Biantidae). 

Type species: Biantoncopus fuscus sp. n. 

Species account and distribution: At present the genus includes only the type 
species from Leyte Island, Philippines. 

Relationships: Biantoncopus gen. n. distinctly stands apart from the other genera 
of Oncopodidae, but its relationships are not clear. Penis morphology is similar to that 
of Palaeoncopus gen. n., both genera possess a distad-directed glans. However, glans 
direction is the only diagnostic character shared with Palaeoncopus species and 
according to the interpretation given in this paper, it presents a symplesiomorphy. 
Additional genital characters (i.e. expandable glans with a pair of membraneous tubes: 
mt in Figs 62, 64-65 and 134c) actually show B. fuscus sp. n. to be quite different from 
Palaeoncopus gen. spp. n. In the expanded state the tip of the stylus far surmounts the 
tip of the truncus and both inflated membraneous tubes are bent downwards, pointing to 
the base of the truncus (Figs 64-65). Morphological and operational similarities to the 
penes of Biantes spp. (Biantidae; Martens 1978) are considerable. 

On the other hand the habitus of B. fuscus sp. n. generally corresponds well with 
Gnomulus species and they also share the tarsal formula 2-2-3-3. Therefore Bianton- 
copus gen. n. may also be interpreted as a Gnomulus with reversal in glans orientation 
or as a separate lineage with partly conservative and partly derived penis morphology 
(see Fig. 134). Additional related species are expected to be discovered in the future, 
which hopefully will elucidate the role this glans type plays in oncopodid evolution. 



Biantoncopus fuscus sp. n. Figs 58-70 

Material: PHILIPPINES, Leyte, Lake Danao, 500 m. S holotype (MHNG), 2 ö\ 4 9 
paratypes (MAR, MHNG), 19.II.-9.III.1991, leg. W. Schawaller & J. Martens. - Visca, N of 
Baybay, 200-500 m, 1 S paratype (MAR), 2.III.1991, leg. W. Schawaller & J. Martens. 

Etymology: Latin: fuscus = brown; the species name refers to the distinct brownish 
colour of the body. 

Diagnosis: Externally similar to Gnomulus maculatus sp. n., but colour pattern 
different, palpal processes larger and genital operculum wider. Penis stout, glans with 
distad-directed expandable pair of membraneous tubes and a protrudable stylus 
(Figs 58-65). 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



523 




Figs 58-69 

Biantoncopus fuscus n. sp.; S holotype (58-61, 68-69), S paratypes (62-65) [Lake Danao (62. 
64-65), N of Baybay (63)], 5 paratype (66-67). - Penis, dorsal (58) and lateral view (59). Apex 
of penis, dorsal (60, 62-64) and lateral view (61, 65), glans partly expanded (64-65): left palp, 
retrolateral view (66); left chelicera, retrolateral view (67); distal part of left leg II, retrolateral 
view (68); distal part of leg IV, prolateral view (69). - Scale lines 0.5 mm (58-59, 66-69). 0.1 
mm (60-65). 



524 



J. MARTENS & P. SCHWENDINGER 






Fig. 70 
Biantoncopus fuscus sp. n., S paratype (from the type locality), body, dorsal (a), ventral (b) and 
lateral view (c). - Scale line 1 mm. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 525 

Description: â (paratype). Coloration: Body amber, dark reticulations on: 
carapace, margin and elevations of scuta, legs (except light orange tarsi) and palps; 
proximal article of chelicerae dark, cheliceral hand cream; dark transversal bands on 
dorsal scutal elevations I-IV medially broken (Fig. 70a). 

Carapace with conical eye tubercle; a pair of truncate lobes forming bridge 
between abdominal part of dorsal scutum and carapace (Fig.70a, c); dorsal and ventral 
scutal areas only slightly elevated (Figs 70c). Ventral coxae II and III with conical 
anterio-proximal processes, ventral coxae I and palpal coxae with knob-shaped central 
processes; genital operculum wide, broadly rounded when closed (Fig. 70b), a pointed 
anterior process visible when opened. 

Chelicerae (Fig. 67) weak; proximal article with a conical dorso-distal process, 
no ventral one; hand slender. 

Palps (Fig. 66): ventral femur with distinct conical proximal process; 
trochanter with large bilobed ventral process, no dorsal process. 

Legs (Figs 68-69) 1324, tarsal formula 2-2-3-3. 

Penis (Figs 58-65): truncus stout, its distal margin more or less distinctly 
invaginated medially; subapical setae in one large group on each side. Glans short, not 
reaching apex, membraneous socket not sharply outlined from truncus; wide lateral 
sclerites bent dorsad (pointing away from apex), embracing median plate, short, 
slender stylus and large pair of membraneous tubes (corresponding to titillatores in 
Biantidae; Martens 1978). Expanded glans with membraneous tubes folded down- 
wards and stylus stretching forward, extending far beyond apex of penis (Figs 64-65). 

9 . As the S ; no external sexual dimoiphism discernible. 

Measurements (o% in brackets 9): body 2.48 (2.60) long, 1.69 (1.73) wide; 
carapace region 0.64 (0.64) long, 0.93 (0.92) wide. - Palp and legs: 





Tr 


Fe 


Pa Ti 


Mt 


Ta 


Total 


Palp 


0.23 (0.23) 


0.34 (0.32) 


0.32(0.32) 0.21 (0.18) 


- - 


0.43 (0.43) 


1.53(1.48) 


Legi 


0.23 (0.23) 


0.59 (0.56) 


0.40(0.37) 0.32(0.32) 


0.50 (0.49) 


0.37 (0.35) 


2.41 (2.32) 


Leg II 


0.27 (0.27) 


0.76(0.71) 


0.47(0.46) 0.44(0.44) 


0.76 (0.73) 


0.43 (0.41) 


3.13(3.02) 


Leg III 


0.24 (0.24) 


0.52 (0.50) 


0.40(0.38) 0.34(0.34) 


0.65 (0.64) 


0.28 (0.25) 


2.43 (2.35) 


Leg IV 


0.29 (0.26) 


0.72 (0.69) 


0.50(0.49) 0.53(0.53) 


0.93 (0.92) 


0.32(0.31) 


3.29 (3.20) 



Variation: Body length/width ranges: S 2.25-2.48/1.60-1.69 (n = 4), $ 2.37- 
2.60/1.59-1.73 (n = 4). Bridge teeth connecting abdominal part of dorsal scutum and 
carapace vary in shape from indistinct and rectangular to distinct and triangular or 
rounded. One S (from the type locality) has a thin cylindrical stylus and a narrowly 
rounded median plate with a smooth margin (Fig. 62). In all other â â the stylus is 
fairly broad and slightly depressed and the median plate is broadly rounded with a 
more or less strongly indented anterior margin (Figs 60-61, 63-65). 

Bionomics: The specimens were extracted from soil and leaf litter on plane and 
strongly sloping forest floor of a primary evergreen forest interspersed by clearings 
with cultivated bananas. 



526 J - MARTENS & P. SCHWENDINGER 



Gnomulus Thorell, 1890 

Gnomulus Thorell (1890: 378), type species originally designated but described later, 

G. sumatranus Thorell, 1891; Pocock (1897: 285)"; Roewer (1923: 60-61); 

SCHWENDINGER (1992: 197). 
Pelitnus Thorell (1891: 757), type species by original designation, P. armillatus Thorell, 1891; 

Pocock (1897: 285); Roewer (1923: 62); Sorensen (1932: 213); Schwendinger 

(1992: 197). NEW SYNONYMY. 

Diagnosis (extended): Penis with basally truncate, subbasally constricted 
cylindrical truncus. Subapical glans proximad-directed, stylus flanked by a pair of 
sclerites (both structures variable in shape and length) and a pair of membraneous 
tubes; median plate present or absent. Interocular area low or elevated into a domed or 
conical tubercle. Bridge between carapace and abdominal part of dorsal scutum 
distinct or absent. Dorsal scutal areas elevated or low. with a more or less distinct 
longitudinal median furrow. Chelicerae weakly or strongly developed, proximal 
article with a more or less distinct dorso-distal hump or tubercle, rarely with a ventro- 
median process; hand unarmed. Palpal femur usually with (rarely without) ventro- 
basal process, rarely with ventro-median one; palpal trochanter with or without 
ventral process, dorsal process absent. Legs 1324, 1342, 3124, 3142; tarsal formula 2- 
2-2-2 or 2-2-3-3. External sexual dimorphism in shape or hair cover of ventral scutal 
elevations, in shape of carapace or in size of chelicerae. 

Species account: Twenty nominal oncopodid species (thirteen previously 
described and seven new), in which penis morphology is known, are here listed under 
Gnomulus. These are: 

G. sumatranus Thorell, 1891 [Sumatra] - Loman 1903: fig. 5f; <3, $9 syntypes in 

MCSNG examined. 
G. segnipes (Loman, 1892) comb, n., transferred from Pelitnus [Sumatra; doubtful 

records from Java and Borneo] - Schwendinger 1992: figs 58-61 (possibly not 

conspecific); presumably conspecific â â from Sumatra in CCD, SMF, ZMA, 

ZMT examined. 
G. aborensis (Roewer, 1913), transferred from Pelitnus by Roewer (1923: 61-62) 

[NE-India] -2 â paratypes in SMF examined. 
G. laevis (Roewer, 1915) comb. n.. transferred from Pelitnus [Borneo] - Schwen- 
dinger 1992: figs 63-66. 
G. insularis (Roewer, 1927) comb. n.. transferred from Pelitnus [Malaysia, Penang 

Island] - 6 holotype in SMF examined (probably identical with G. rostratus - 

$ holotype in MCSNG examined). 
G. drescoi (Silhavy, 1962) comb, n., transferred from Pelitnus [Sumatra] - Suzuki 

1982: figs 13, 14. 
G. imadatei (Suzuki, 1969) comb, n., transferred from Pelitnus [Brunei] - Suzuki 

1969: figs 4a-e. 
G. hyatti (Martens, 1977) comb. n.. transferred from Pelitnus [Nepal] - Martens 

1977: figs 3, 4. 
G. goodnighti (Suzuki, 1977) comb, n., transferred from Pelitnus [Philippines, 

Mindanao] - Suzuki 1977: figs 2f-i. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 527 

G. lannaianus (Schwendinger, 1992) comb, n., transferred from Pelitnus [Thailand] - 

Schwendinger 1992: figs 8-13. 
G. baharu Schwendinger nom. n. (Malay and Indonesian: baharu = new), homo- 

nymously described under Pelitnus thorelli [Brunei] by Schwendinger 1992: 

figs 21-26. 
G. conigerus (Schwendinger, 1992) comb, n., transferred from Pelitnus [Sabah] - 

Schwendinger 1992: figs 36-41. 
G. sundaicus (Schwendinger, 1992) comb, n., transferred from Pelitnus [Sarawak] - 

Schwendinger 1992: figs 48-53. 
G. crucifer sp. n. - Philippines, Luzon. 
G. maculatus sp. n. - Philippines, Luzon and Mindoro (?). 
G. coniceps sp. n. - Philippines, Luzon. 
G leyteensis sp. n. - Philippines, Leyte. 
G laruticus sp. n. - Malaysia, Perak. 
G. asli sp. n. - Malaysia, Perak. 
G. hirsutus sp. n. - Malaysia, Selangor and Pahang. 

In the remaining eight species described under Gnomulus and Pelitnus penis 
morphology is yet unknown. Therefore they may belong to either Gnomulus or Biant- 
oncopus gen. n., which are indistinguishable by external characters. As Gnomulus is 
by far the more widespread of both genera, we provisionally place these species in 
Gnomulus. 

G. rostratus Thorell, 1890 [Malaysia, Penang Island] - see G. insularis comb. n. 

G. armillatus (Thorell, 1891) comb. n„ transferred from Pelitnus [Sumatra]. 

G. annulipes (Pocock, 1897) comb, n., transferred from Pelitnus [Sarawak]. 

G. pulvillatus (Pocock, 1903) comb, n., transferred from Pelitnus [Malaysia, 

Selangor]. 
G piliger (Pocock, 1903) comb, n., transferred from Pelitnus [Malaysia]. 
G. thorelli (S0rensen, 1932) comb, n., transferred from Pelitnus [Java]. 
G. palawanensis (Suzuki, 1982) comb, n., transferred from Pelitnus [Philippines, 

Palawan)]. 
G. minor Tsurusaki, 1990 [Philippines, Luzon]. 

Distribution: Himalayan Region (Nepal and northeastern India) and Southeast 
Asia (northern Thailand to Java and the Philippines). 



New species of gnomulus 

Gnomulus crucifer sp. n. Figs 71-81 

Material: PHILIPPINES, Luzon, Mt. Santo Thomas close to Baguio, ca. 1850 m, 
3 holotype, 14.1.1980; leg. L. Deharveng (MHNG). 

Etymology: Latin: crux = cross, ferre = to carry; the species name refers to the cross- 
shaped glans penis of this species. 



528 



J. MARTENS & P. SCHWENDINGER 




Figs 71-80 

Gnomulus crucifer sp. n., S holotype. - Penis, dorsal (71) and lateral view (72); apex of penis, 
dorsal (73) and lateral view (74); glans penis, dorso-lateral view (75); left chelicera, retrolateral 
view (76); left palp, retrolateral view (77); trochanter and femur of left palp, retrolateral view 
(78); distal part of left leg II. retrolateral view (79); distal part of left leg IV, prolateral view 
(80). - Scale lines 0.5 mm (71-72. 76-80), 0.1 mm (73-75). 



Diagnosis: Similar to G. minor Tsurusaki but with elevated interocular area, 
process on palpal trochanter different in shape, anterio-proximal process on ventral 
leg coxa II present; legs 1324, tarsal formula 2-2-2-2. Glans penis proximad-directed, 
bearing an enlarged stylus with cross-shaped apex; lateral sclerites lobate, median 
plate absent (Figs 71-75). 

Description: <$ (holotype). Coloration: body light amber, with dark reticu- 
lations on carapace, dark transversal bands on abdominal part of dorsal scutum and 
dark, medially broken transverse bands on ventral scutum (Figs 81a, b); genital oper- 
culum with indistinct dark reticulation; chelicerae and pedipalps cream, except 
slightly darker palpal femur; legs grey-brown, except slightly lighter trochanters and 
cream coxae, tarsi and distal metatarsi. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



529 






Fig. 81 

Gnomulus crucifer sp. n., ê holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale 
line 1 mm. 



530 L MARTENS & P. SCHWENDINGER 

Carapace with low, broadly rounded eye tubercle (Figs 81a, c); connection 
between carapace and abdominal part of dorsal scutum indistinct; dorsal and ventral 
scutal areas only slightly elevated (Fig. 81c). Ventral leg coxae II and III with conical 
anterio-proximal process, coxa II with rounded posterio-proximal one; ventral coxae I 
and palpal coxae with knob-shaped paramedian and small central processes, respect- 
ively; genital operculum wide (Fig. 81b). 

Chelicerae (Fig. 76) weak; proximal article with dorso-distal boss, no ventral 
process; hand slender. 

Palps (Fig. 77-78): ventral femur with low conical proximal process; 
trochanter with pronounced, somewhat trilobed ventral process, no dorsal process. 

Legs (Figs 79-80) 1324, tarsal formula 2-2-2-2. 

Penis (Figs 71-75): truncus fairly slender, apex narrow, with broadly rounded 
distal margin; subapical setae in one small group on each side. Glans with a long, 
proximally wide stylus, constricted below a cross-shaped apex with acutely pointed 
tip; lateral sclerites flat, distally rounded, lying above stylus; median plate absent; pair 
of membraneous tubes partly covered by stylus. 

2 . Unknown. 





Measurements 


(S): 


body 


3.12 long. 


1.78 wide; 


carapace 


0.88 long, 


1.02 wide 


-Palp 


and legs: 


















Lr 


Fe 




Pa 


Li 


Mt 


La 


Lotal 


Palp 


0.31 


0.43 




0.35 


0.26 


- 


0.49 


1.84 


Legi 


0.23 


0.70 




0.41 


0.40 


0.62 


0.43 


2.79 


Leg II 


0.31 


0.94 




0.50 


0.64 


1.01 


0.49 


3.89 


Leg III 


0.24 


0.63 




0.44 


0.43 


0.76 


0.30 


2.80 


Leg IV 


0.29 


0.93 




0.61 


0.73 


1.11 


0.34 


4.01 



Relationships: Habitus, shape of palps and chelicerae and geographical 
proximity suggest close relationship with G. minor ( S unknown), although the tarsal 
formula is different. The penis with an enlarged stylus and without a median plate, on 
the other hand, distantly resembles that of Caenoncopus gen. n. At the present state of 
knowledge, however, it appears that this partial congruence in penis morphology was 
caused by parallelism. 

Bionomics: The specimen was extracted from vegetational debris in a humid 
ravine of an evergreen hill forest. 

Gnomulus maculatus sp. n. Figs 82-89 

Material: PHILIPPINES, Luzon, Mountain Province, dolines NE of Sagada, 6 holo- 
type (MHNG), 21. XII. 1979; 1 S, 1 2 paratypes (MAR, MHNG), Sagada, near Latan Cave, 
6.1.1980; Sagada, near village and near Sogong Cave, 4., 5., 9.1.1980, 3 juv. (MAR).- Mindoro, 
Puerto Galera, near San Theodoro Waterfall, 1 9 paratype (MAR), 2.-4.I.1979 (probably 
mislabeled). All specimens leg. L. Deharveng. 

Etymology: Latin: maculatus = spotted; the species name refers to the conspicuous 
colour pattern on the dorsal scutum. 

Diagnosis: Closely related to G. goodnighti, distinguished by distinct colour 
pattern; eye tubercle lower; dorsal scutum broadly rounded behind, without para- 
median humps on posterior scutal areas, with two lobate bridge teeth on anterior 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



53; 




Figs 82-88 

Gnomulus maculatus sp. n.; S holotype (82-85), S paratype (86), 9 paratype (87-88). - Penis, 
dorsal (82) and lateral view (83); apex of penis, dorsal (84, 86) and lateral view (85); left palp, 
retrolateral view (87); left chelicera, retrolateral view (88). - Scale lines 0.5 mm (82-83, 87-88), 
0.1 mm (84-86). 

margin (none on posterior carapace); small anterio-proximal processes present on leg 
coxae III. Distal margin of truncus penis rounded; membraneous socket of glans wide; 
paired lateral sclerites of glans only little bent, terminally truncate; median plate long, 
tongue-shaped; stylus stout, with bifurcate apex (Figs 82-85). 

Description: 6 (holotype). Coloration: body light yellow, with dark rings 
around eyes and dark procurved band on posterior carapace; dorsal scutum very dark 
along margin and on scutal areas, except for conspicuous light yellow median zone on 
last 3 scutal areas (Fig. 89a, c); dark transversal bands on ventral scutum unbroken, 
except for the last one (Fig. 89b); genital operculum grey-yellow, tarsus of leg I 
cream. 

Carapace with conical eye tubercle; lobed bridge teeth between abdominal part 
of dorsal scutum and carapace; dorsal and ventral scutal areas moderately elevated 
(Fig. 89a, c). Ventral leg coxae II and III with small anterio-proximal processes; 
ventral coxae I and palpal coxae with knob-shaped median processes; genital oper- 
culum wide (Fig. 89b). 

Chelicerae (Fig. 88) weak; proximal article with dorso-distal boss, no ventral 
process; hand slender. 



532 



J. MARTENS & P. SCHWENDINGER 






Fig. 89 
Gnomulus maculants sp. n., S holotype, body, dorsal (a), ventral (b) and lateral view (c). 
Scale line 1 mm. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 533 

Palps (Fig. 87): ventral femur with low proximal process; trochanter with 
distad directed ventral process. 

Legs 1324, tarsal formula 2-2-3-3. 

Penis (Figs 82-86): truncus fairly slender, apex narrow, distal margin broadly 
arched, without median indentation; subapical setae in two indistinctly separated 
groups on each side. Glans rising from a very wide membraneous socket; lateral 
sclerites long, forceps-like, connected by a long, tongue-shaped median plate with 
finely serrated distal margin; stylus stout, its apex bifurcate, with tip (carrying 
opening of sperm duct) bending towards the truncus at right angles below a pointed 
distad directed process; pair of membraneous tubes mostly covered by median plate. 

? . As the 6; no external sexual dimorphism discernible. 

Measurements (o\ in brackets 9): body 2.80 (2.94) long, 1.90 (1.90) wide; 
carapace region 0.76 (0.69) long, 1.03 (1.04) wide. - Palp and legs: 

Tr Fe Pa Ti Mt Ta Total 

Palp 0.29(0.29) 0.42(0.44) 0.35(0.34) 0.24(0.24) - - 0.54(0.55) 1.84(1.86) 

Legi 0.24(0.26) 0.67(0.69) 0.42(0.41) 0.43(0.41) 0.64(0.66) 0.41(0.41) 2.81(2.84) 

Leg II 0.34(0.34) 0.91(0.90) 0.53(0.53) 0.63(0.62) 0.96(0.99) 0.47(0.48) 3.84(3.86) 

Leg III 0.27(0.27) 0.61(0.63) 0.44(0.44) 0.46(0.45) 0.81(0.80) 0.32(0.33) 2.91(2.92) 

Leg IV 0.32(0.32) 0.90(0.93) 0.59(0.60) 0.76(0.75) 1.14(1.17) 0.35(0.35) 4.06(4.12) 

Variation: Body length/width ranges: 6 2.80-2.84/1.87-1.90 (n = 2), 9 2.94- 
3.18/1.90-2.09 (n = 2). Bridge teeth between abdominal part of dorsal scutum and 
carapace vary in shape; they are generally smaller in the 9 9 examined, in one of 
them missing on the left side. The median plate of the glans penis is narrower in the 
S paratype (Fig. 86). The largest specimen ( 9 ) is darker than all others, i.e. with dark 
reticulation on: carapace, genital operculum, leg coxae and trochanters, palps (except 
tarsus) and proximal chelicerae; femora to metatarsi of legs grey-brown. 

Relationships: The enlarged, uniquely formed stylus of this species probably 
represents an earlier stage of glans modification, which is more strongly developed in 
G. crucifer sp. n. In other characters G. maculatus sp. n. most closely resembles 
G. goodnighti. 

Bionomics: The specimens were collected from humid leaf litter and moss in 
an evergreen hill forest by means of sifting and Berlèse-extraction. 

Distribution (Fig. 3): According to the collecting data the specimens are from 
two Philippine islands, Luzon and Mindoro. This would be an exceptionally wide 
distribution for an oncopodid species. The record of a single specimen from Mindoro is 
very probably due to a confusion of labels (supported by L. Deharveng, pers. comm.). 



Gnomulus coniceps sp. n. Figs 90-96 

Material: PHILIPPINES, Luzon, Baguio, near Cristal Caves, 1500 m, â holotype 
(MHNG), 12.1.1980, leg. L. Deharveng. 

Etymology: Latin: conus = cone; ceps (from caput) = head; noun in apposition. The 
name refers to the unusually large eye tubercle of this species. 



534 



J. MARTENS & P. SCHWENDINGER 




Figs 90-95 

Gnomulus coniceps sp. n., S holotype. - Penis, dorsal (90) and lateral view (91); apex of penis, 
dorsal (92) and lateral view (93); left palp, retrolateral view (94); left chelicera, retrolateral 
view (95). - Scale lines 0.5 mm (90-91, 94-95), 0.1 mm (92-93). 



Diagnosis: Closest to Gnomulus goodnighti, distinguished by basally wider 
eye tubercle and narrower genital operculum. Penis with broadly invaginated anterior 
margin of truncus; glans penis with strongly geniculate lateral sclerites connected to a 
narrow, pointed median plate (Figs 90-93). 

Description: S (holotype). Coloration: body amber, with dark reticulations on 
carapace (especially around eyes), very dark dorsal scutal areas with a light 
longitudinal median band between areas II-IV (Fig. 96a); ventral scutal areas with 
transversal bands becoming increasingly darker posteriorly (Fig. 96b). Chelicerae, 
pedipalps and legs light amber, except slightly darker metatarsi I-IV and patella and 
tibia IV; terminal tarsal segment darker than basal ones. 

Carapace short, with broadly conical eye tubercle occupying almost entire 
length of carapace (Fig. 96c); triangular pair of abdominal scutal processes forming 
bridge with pair of processes from carapace. Abdominal part of dorsal scutum high, 
with paramedian pairs of humps on scutal areas (largest on V-VII); posterior margin 
of dorsal scutum slightly pointed (Figs 96a, c); ventral scutum with only slightly 
elevated areas (Figs 96b, c). Distinct anterio- and posterio-proximal processes on 
ventral leg coxa II, a smaller one on coxa III; palpal coxae with large ventral pro- 
cesses; genital operculum longer than wide (Fig. 96b). 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



535 





Fig. 96 

Gnomulus coniceps sp. n., 6 holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale 
line 1 mm. 



536 J - MARTENS & P. SCHWENDINGER 

Chelicerae (Fig. 95) weak; proximal article with dorso-distal boss, no ventral 
process; hand slender. 

Palps (Fig. 94): ventral femur with indistinct proximal process; trochanter with 
distad-directed ventral process. 

Legs 1324, tarsal formula 2-2-3-3. 

Penis (Figs 90-93): truncus stout, apex wide, anterior margin broadly 
invaginated; subapical setae in one large group on each side. Glans medially con- 
stricted; lateral sclerites strongly geniculate, proximally divergent, distally 
convergent, tips ramified, touching each other; lateral sclerites connected by a short, 
spike-like median plate above slender stylus; pair of membraneous tubes clearly 
visible at constriction of glans. 

9 . Unknown. 

Measurements (S): body 3.37 long, 2.66 wide; carapace region 0.78 long, 
1.38 wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.42 


0.61 


0.51 


0.33 


- 


0.65 


2.52 


Legi 


0.33 


1.05 


0.56 


0.64 


1.03 


0.59 


4.20 


Leg II 


0.35 


1.35 


0.73 


1.02 


1.55 


0.65 


5.65 


Leg III 


0.32 


1.05 


0.61 


0.76 


1.35 


0.53 


4.62 



Leg IV 0.41 1.40 0.73 1.05 1.89 0.61 6.09 

Relationships: According to external morphology G. coniceps sp. n. is closest 
to G. goodnighti and (more distant) to G. maculatus sp. n., but its glans penis is quite 
different from both of them and shows no close resemblance with any species known 
so far. 

Bionomics: The specimen was collected from a ravine in an evergreen hill 
forest. 



Gnomulus leyteensis sp. n. Figs 97-104 

Material: PHILIPPINES, Leyte, Visca, north of Baybay, 200-500 m, 3 holotype 
(MHNG), 10.III. 1991. leg. J. Martens & W. Schawaller. 

Etymology: The specific epithet is taken from the name of the island where the species 
was collected. 

Diagnosis: Closest to G. goodnighti, distinguished by more rounded posterior 
margin of body, narrower genital operculum and basally wider ventral process on 
palpal trochanter. Penis apically narrower; glans more remote from distal margin; 
lateral sclerites less strongly bent, with ramified tip; median plate indistinct (Figs 97- 
101). 

Description: â (holotype). Coloration: body amber, with dark reticulations on 
carapace (especially around eyes); dorsal scutum with very dark margin and 
transversal bands on elevations, in scutal areas I-IV broken by a light longitudinal 
median band (Fig. 104a); ventral scutal areas with unbroken dark transversal bands 
(Fig. 104b). Chelicerae, pedipalps and legs light amber, except slightly lighter tarsi 
and slightly darker metatarsus II. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



537 




Figs 97-103 

Gnomulus leyteensis sp. n., S holotype. - Penis, dorsal (97) and lateral view (98); apex of 
penis, dorsal (99) and lateral view (100); glans with tips of lateral sclerites folded downwards 
(101); left palp, retrolateral view (102); left chelicera, retrolateral view (103). - Scale lines 0.5 
mm (97-98, 102-103), 0.1 mm (99-101). 



Carapace short, with conical eye tubercle set back from anterior margin; 
posterior part of carapace flat, triangular pair of paramedian processes forming bridge 
with corresponding processes on anterior margin of abdominal part of dorsal scutum. 
Dorsal scutum high, with paramedian pairs of indistinct humps on areas I-VII; 
posterior margin of dorsal scutum slightly pointed (Figs 104a, c); ventral scutum with 
only slightly elevated areas (Figs 104b, c). Ventral leg coxa II with distinct anterio- 
and posterio-proximal processes, coxa III with small anterio-proximal one; palpal 
coxa with large ventral process; genital operculum about as long as wide (Figs 104b). 

Chelicerae (Fig. 103) small, slender; proximal article with dorso-distal boss, no 
ventral process; hand slender. 

Palps (Fig. 102): ventral femur with indistinct proximal process; trochanter 
with distad-directed, basally wide ventral process. 

Legs 1324, tarsal formula 2-2-3-3. 

Penis (Figs 97-101): truncus with narrow apex and slightly invaginated distal 
margin; subapical setae in one large group on each side. Glans distinctly remote from 
apex of truncus; lateral sclerites bent towards each other and away from the truncus in 
distal half, its tips strongly ramified, touching each other; median plate very short, 
indistinct; stylus fairly stout, its tip bent towards the truncus. 



538 



J. MARTENS & P. SCHWENDINGER 






Fig. 104 
Gnomulus leyteensis sp. n., S holotype, body, dorsal (a), ventral (b) and lateral view (c). 
Scale line 1 mm. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



539 



wide. 



9 . Unknown. 

Measurements {8): body 3.27 long, 2.39 wide; carapace region 0.76 long, 
- Palp and legs: 



.24 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.32 


0.50 


0.43 


0.31 


- 


0.64 


2.20 


Legi 


0.27 


0.85 


0.53 


0.56 


0.87 


0.55 


3.63 


Leg II 


0.35 


1.14 


0.69 


0.88 


1.33 


0.59 


4.98 


Leg III 


0.31 


0.87 


0.58 


0.64 


1.11 


0.46 


3.97 


Leg IV 


0.41 


1.19 


0.72 


0.96 


1.59 


0.50 


5.37 



Relationships: According to penis morphology Gnomulus leyteensis sp. n. is 
closest to G. goodnighti. 

Bionomics: The specimen was sifted from leaf litter and humus on steep slopes 
of an evergreen primary forest. In small interspersed clearings bananas were grown. 



Gnomulus laruticus sp. 



Figs 105-113 
Hill (= Bukit Larut), 1320 m, 



Material: MALAYSIA, Perak, Taiping, Maxwell 
S holotype (MHNG), 26.1.1995, leg. P. Schwendinger. 

Etymology: The species name is taken from Bukit Larut, the Malayan name of the type 
locality. 




Figs 105-112 

Gnomulus laruticus sp. n., holotype. - Penis, dorsal (105) and lateral view (106); apex of penis, 
dorsal (107) and lateral view (108); left palp, retrolateral view (109); left chelicera, retrolateral 
view (110); distal part of left leg IV, prolateral view (111) and of leg II, retrolateral view (112). 
- Scale lines 0.5 mm (105-106, 109-1 12), 0.1 mm (107-108). 



540 



J. MARTENS & P. SCHWENDINGER 






Fig. 113 
Gnomulus laruticus sp. n., â holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale 
line 1 mm. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 541 

Diagnosis: Resembling G. crucifer sp. n. in body shape and tarsal formula but 
different in shape of eye tubercle, colour pattern and presence of a wide bridge 
between carapace and abdominal part of dorsal scutum; genital operculum smaller: 
process on anterio-lateral margin of leg coxae I present; ventral process on palpal 
trochanter smaller; low ventro-median hump on proximal article of chelicerae. Glans 
penis with slender stylus and pair of sigmoid, pointed lateral sclerites interconnected 
by a short, broadly rounded median plate (Figs 105-108). 

Description: S (holotype). Coloration: body mostly light amber. Dark reticu- 
lations on carapace and dark transversal bands on abdominal part of dorsal scutum 
(Fig. 113a); indistinct dark lateral and paramedian patches on ventral scutal areas 
(Fig. 113b). Femora to metatarsi of legs slightly darkened, metatarsus II and proximal 
half of tibia IV distinctly darkened. 

Carapace with low, broadly rounded eye tubercle; posterior part of carapace 
slightly elevated; a single short, wide lobe there forming bridge with corresponding 
lobe (shorter, wider) on anterior margin of abdominal part of dorsal scutum (Figs 
113a, c); dorsal and ventral scuta low, areas indistinctly elevated (Fig. 1 13c). Ventral 
leg coxa II with anterio- and posterio-proximal processes, coxa III with anterio- 
proximal one; coxa I with conical process on anterio-lateral margin; palpal coxa with 
ventral process; genital operculum subtriangular, wider than long (Figs. 1 13b). 

Chelicerae (Fig. 110) weak; proximal article with dorso-distal boss and 
indistinct ventral process; hand slender. 

Palps (Fig. 109): ventral femur with small proximal process; ventral trochanter 
with pronounced distad directed process. 

Legs 3124 (Figs 1 1 1-1 12), tarsal formula 2-2-2-2. 

Penis (Figs 105-108): truncus with narrow apex and broadly arched, medially 
indistinctly invaginated anterior margin; subapical setae in one group on each side. 
Glans with sigmoid lateral sclerites, their acutely pointed tips bent away from the 
truncus and away from each other. Median plate short, broadly rounded, with few tiny 
teeth on margin; pair of membraneous tubes completely covered by median plate; 
stylus slender. 

$ . Unknown. 

Measurements (S): body 2.48 long, 1.52 wide; carapace region 0.63 long, 0.94 
wide. - Palp and legs: 





Tr 


Fe 


Pa 


Ti 


Mt 


Ta 


Total 


Palp 


0.26 


0.31 


0.29 


0.19 


- 


0.40 


1.45 


Legi 


0.23 


0.66 


0.40 


0.35 


0.53 


0.37 


2.54 


Leg II 


0.27 


0.88 


0.54 


0.57 


0.79 


0.47 


3.52 


Leg III 


0.23 


0.56 


0.40 


0.38 


0.64 


0.27 


2.48 


Leg IV 


0.26 


0.86 


0.55 


0.63 


0.92 


0.31 


3.53 



Relationships: Congruences in penis morphology show closest relationship 
between G. laruticus sp. n., G. asli sp. n. and G. hirsutus sp. n. An unusual tarsal for- 
mula (2-2-2-2) links G. laruticus sp. n. with G. crucifer sp. n., but their distinctly 
diffent penes indicate a parallel retention of the juvenile tarsal number on posterior 
legs. 



542 



J. MARTENS & P. SCHWENDINGER 



Bionomics: The specimen was collected by sifting leaf litter from the floor of a 
lower montane rain forest. 




Figs 114-121 

Gnomulus asli sp. n.; 6 holotype (114-117), S paratypes (118-119), 9 paratype (120-121). - 
Penis, dorsal (114) and lateral view (115); apex of penis, dorsal (116, 118-1 19) and lateral view 
(117); left palp, retrolateral view (120); left chelicera, retrolateral view (121). - Scale lines 0.5 
mm (114-115, 120-121), 0.1 mm (116-119). 



Gnomulus asli sp. n. 



Figs 114-122 



Material: MALAYSIA, Perak, forest 5 km northeast of Chenderiang, 290-330 m, 
6 holotype (MHNG), 2 8, 1 9 paratypes (MAR, MHNG), 22.-3 1.1.1994; 330-400 m, 6 9 
paratypes (MAR. MHNG), 15.-22.I.1995; leg. P. Schwendinger. 

Etymology: Malay and Indonesian: asli = indigenous, original. The species is named 
after the Orang Asli, the indigenous people of Malaysia. The specimens were collected from a 
forest utilized by the inhabitants of an Orang Asli (Semai Senoi tribe) village. 

Diagnosis: Close to G. laruticus sp. n. but distinguished by different colour 
pattern, lower eye tubercle, two bridge teeth between carapace and abdominal part of 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



543 






Fig. 122 
Gnomulus asli sp. n., 3 holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale line 



544 J - MARTENS & P. SCHWENDINGER 

dorsal scutum, process on palpal trochanter basally rounded and tarsal formula 2-2-3- 
3. Glans penis distally wider; tips of lateral sclerites pointing towards each other (Figs 
114-119). 

Description: o* (holotype). Coloration: body mostly light amber, with dark 
reticulations on carapace and distinct dark transversal bands on abdominal part of 
dorsal scutum (Fig. 122a), indistinct ones on ventral scutum (Fig. 122b). Genital 
operculum and femora to metatarsi of legs dark amber, tarsi light yellow, ventral side 
of distitarsus I cream. 

Carapace with low, broadly rounded eye tubercle; pars thoracica slightly elev- 
ated; two tubercles on posterior margin forming bridge with pair of short, obliquely 
truncate teeth on anterior margin of abdominal part of dorsal scutum (Fig. 122a, c); 
dorsal and ventral scuta low, areas indistinctly elevated; ventral scutal areas covered 
by fine short hairs (Fig. 122c). Ventral leg coxa II with distinct anterio- and posterio- 
proximal processes, coxa III with anterio-proximal one; coxa I with conical process 
on anterio-lateral margin; palpal coxa with ventral process; genital operculum 
subtriangular, distinctly wider than long (Fig. 122b). 

Chelicerae (Fig. 121) small, slender; proximal article with dorso-distal boss, no 
ventral process; hand slender. 

Palps (Fig. 120): ventral femur with small proximal process; ventral trochanter 
with pronounced distad directed process. 

Legs 1324, tarsal formula 2-2-3-3. 

Penis (Figs 114-119): truncus slender, distal margin widely arched and 
indistinctly invaginated medially; subapical setae in one group on each side. Glans 
with sigmoid lateral sclerites, their pointed tips distinctly bent away from the truncus 
and towards each other. Median plate short, broadly rounded, with several denticles 
on margin; pair of membraneous tubes completely covered by median plate; stylus 
slender. 

9 . As the 6 , except for an indistinct hair cover on ventral scutum. 

Measurements (6, in brackets 2): body 2.34 (2.39) long, 1.59 (1.63) wide; 
carapace region 0.61 (0.62) long, 0.92 (0.91) wide. - Palp and legs: 

Tr Fe Pa Ti Mt Ta Total 

Palp 0.26(0.27)0.31(0.31)0.27(0.27)0.18(0.18) - - 0.38(0.37) 1.40(1.40) 

Leg I 0.24 (0.24) 0.59 (0.60) 0.38 (0.37) 0.35 (0.34) 0.50 (0.50) 0.40 (0.40) 2.46 (2.45) 

Leg II 0.29(0.29) 0.81(0.82) 0.47(0.48) 0.50(0.50) 0.79(0.79) 0.48(0.47) 3.34(3.35) 

Leg III 0.24(0.24) 0.56(0.56) 0.40(0.38) 0.37(0.37) 0.64(0.64) 0.29(0.29) 2.50(2.48) 

Leg IV 0.26 (0.27) 0.79 (0.79) 0.50 (0.50) 0.56 (0.56) 0.96 (0.96) 0.32 (0.32) 3.39 (3.40) 

Variation: Body length/width ranges: 6 2.29-2.34/1.56-1.59 (n = 3), 9 2.28- 
2.51/1.51-1.69 (n = 7). 

Relationships: Penis morphology shows that G. asli sp. n. and G. laruticus sp. 
n. are closest relatives, despite differences in the tarsal formula (2-2-3-3 versus 2-2-2- 
2) and in the shape of the carapace-abdomen bridge. 

Bionomics: The specimens were sifted from leaf litter and humus in a lowland 
rain forest. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 



545 




Figs 123-132 

Gnomulus hirsutus sp. n.; S holotype (123-126), 6 paratype (127-128, 131-132), 9 paratype 
(129-130). - Penis, dorsal (123) and lateral view (124); apex of penis, dorsal (125, 127) and 
lateral view (126, 128); left palp, retrolateral view (129); left chelicera, retrolateral view (130); 
left chelicera, proximal article, retrolateral view (131); anterior part of body, lateral view (132). 
- Scale lines 0.5 mm (123-124, 129-132), 0.1 mm (125-128). 



Gnomulus hirsutus sp. n. 



Figs 123-133 



Material: MALAYSIA, Selangor, Templer Park, 6 holotype and 1 9 paratype 
(MHNG), 1-3.XII.1990, leg. C. Deeleman-Reinhold. - Ulu Gombak, University of Malaya 
Field Centre, 200 m, 1 o\ 1 9 paratypes (MAR, MHNG), 26.IX.1991, leg. D. Agosti. - 
Pahang, on the road to Genting Highlands, 500-800 m, 1 9 paratype (MAR), 13.XII.1982, leg. 
H. Ono. 

Etymology: Latin: hirsutus = hairy; the specific epithet refers to the relatively hairy 
body of this species. 



546 



J. MARTENS & P. SCHWENDINGER 






Fig. 133 
Gnomulus hirsutus sp. n.. 3 holotype. body, dorsal (a), ventral (b) and lateral view (c). - Scale 
line 1 mm. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 547 

Diagnosis: Close to G. asli sp. n. but distinguished by larger body covered by 
fine hairs; eye tubercle higher, prorect; ventral scutal areas sexually dimorphic; 
ventral process on palpal trochanter basally narrower; proximal article of chelicerae 
with distinct dorso-median boss; glans penis with tips of lateral scleri tes further apart 
from each other, median plate without marginal teeth (Figs 123-128). 

Description: â (holotype). Coloration: mostly dark amber, with dark reticu- 
lation on carapace, dark margin and transversal bands on abdominal part of dorsal 
scutum (Fig. 133a); ventral scutum light orange, its transversal bands indistinct (Fig. 
133b); ventral prosoma light amber. Leg trochanters, chelicerae and pedipalps light 
yellow with dark spots (except on cheliceral hand and on palpal tarsus); leg tarsi light 
amber, ventral distitarsus I cream. 

Whole body quite densely covered with fine light hairs. Carapace with prorect, 
rounded eye tubercle; pars thoracica only little elevated; two pairs of tubercles 
forming bridge between carapace and abdominal part of dorsal scutum; dorsal and 
ventral scuta low (Figs 133a, c); ventral scutal areas moderately swollen and pale 
(Figs 133b. c). Ventral leg coxa II with distinct anterio- and posterio-proximal 
processes overlapping anterio-proximal process on coxa III; coxa I with central and 
anterio-lateral processes; palpal coxa with ventral process; genital operculum broadly 
rounded anteriorly, slightly wider than long (Fig. 133b). 

Chelicerae (Figs 130-131) weak; proximal article with distinct dorso-distal and 
smaller dorso-median boss, no ventral process; hand slender. 

Palps (Fig. 129): ventral femur with distinct proximal process; ventral 
trochanter with pronounced distad-directed bilobed process. 

Legs 1324, tarsal formula 2-2-3-3. 

Penis (Figs 123-128): truncus slender, its distal margin broadly arched, 
indistinctly invaginated medially; subapical setae in one group on each side. Glans 
with sigmoid lateral sclerites, their pointed tips slightly bent away from the truncus 
and towards each other. Median plate short, without teeth on margin; pair of 
membraneous tubes mostly covered by median plate; stylus slender. 

9 . As the 6 , except for less elevated, more strongly pigmented ventral scutal 
areas. 

Measurements (a, in brackets 2): body 4.02 (4.07) long, 2.54 (2.63) wide; 
carapace region 0.86 (0.83) long, 1.45 (1.41) wide. - Palp and legs: 

Tr Fe Pa Ti Mt Ta Total 

Palp 0.43(0.43)0.52(0.50)0.43(0.43)0.28(0.29)- - 0.70(0.66)2.36(2.31) 

Legi 0.43(0.41) 0.96(0.96) 0.59(0.59) 0.56(0.57)0.78(0.78) 0.61(0.61) 3.93(3.92) 
Leg II 0.55(0.53) 1.30(1.33) 0.79(0.81) 0.88(0.88) 1.24(1.25) 0.70(0.70) 5.46(5.50) 
Leg III 0.42(0.42) 0.93(0.93) 0.64(0.64) 0.61(0.59) 1.02(1.03) 0.47(0.47) 4.09(4.08) 
Leg IV 0.50(0.50) 1.36(1.34) 0.82(0.84) 0.95(0.98) 1.55(1.60) 0.55(0.53) 5.73(5.79) 

Variation: Body length/width ranges: 6 3.81-4.02/2.54-2.57 (n = 2), 9 4.02- 
4.14/2.63-2.78 (n = 3). The 9 from the type locality and the S from Ulu Gombak 
have a more distinct dark pattern and a light longitudinal median stripe between areas 
II and III on their dorsal scutum. The 9 from Ulu Gombak shows a faint pattern in 
the central region of its dorsal scutum. The o* paratype possesses an obliquely 



548 J - MARTENS & P. SCHWENDINGER 

truncate eye tubercle (as seen from laterally; Fig. 132), smaller bridge teeth on the 
anterior margin of its abdominal part of dorsal scutum and an indistinct dorso-median 
boss on the proximal article of its chelicerae (Fig. 131). 

Relationships: External and genital morphology show closest relationship 
between G. hirsutus sp. n. and G. asli sp. n.; G. laruticus sp. n. is more distant. 
Another seemingly related form (only 1 9 available) occurs on Maxwell Hill. 

Bionomics: The specimens were collected from leaf litter of primary and 
secondary evergreen forests. 

UNDETERMINED MATERIAL 

Nine other related forms are available only in $ $ . We are reluctant to describe 
new species from 5 9 , since in the absence of penis characters a generic placement is 
by no means certain, considering the close external resemblance of Palaeoncopus gen. 
n. with Caenoncopus gen. n. and Biantoncopus gen. n. with Gnomulus. Nevertheless 
these specimens are recorded here, together with their presumed relationships, in the 
hope that further collecting at their find localities will yield S specimens. 

1. 1 9, Sumatra, 7 km north of Brastagi, 1500 m, 2.XII.1989, leg. I. Lobi et al. 
(MHNG) — similar to C. cuspidatus (tarsi 1-1-2-2) but distinctly smaller; colour 
pattern different; interocular area not elevated; posterior margin of body more 
rounded; ventral process on palpal trochanter blunt. Collected from the same locality 
as C. cuspidatus, probably syntopic. 

2. 1 9, Sumatra, Bukittinggi, Gunung Merapi, 2000-2200 m, 18.X.1990, leg. 
A. Riedel (MAR) — similar to C. tenuis sp. n. but different in colour pattern, more 
slender legs and wider genital operculum. 

3. 7 9, Sumatra, Mt. Kerinci, 1750-1900 m, 11.-13.XI.1989, leg. I. Lobi et al. 
(MHNG) — close to C. ajfinis sp. n. but different in colour pattern; eye tubercle more 
rounded; genital operculum much larger; ventral process on palpal trochanter smaller. 

4. 2 9, Sumatra, Panti, 250 m, 19.XI.1989, leg. I. Lobi et al. (MHNG) — close to 
P. gunung sp. n. but distinctly smaller; eye tubercle and two humps on pars thoracica 
more elevated, the latter almost pointed. Apparently syntopic with C. ajfinis sp. n. 

5. 1 9, Sumatra, Mt. Kerinci, 1750-1850 m, 11.-12.XI.1989, leg. I. Lobi et al. 
(MHNG) — close to P. gunung sp. n. but much larger; pars thoracica lower, without 
pair of humps; ventral palpal trochanter with much larger, multilobed process; prox- 
imal article of chelicerae with pointed prodorsal process on distal corner. 

6. 1 9, Indonesia, Kalimantan, Kaharian, 2.-16.IX.1985, leg. C. Deeleman-Reinhold 
(MAR) — similar to C. cuspidatus (tarsi 1-1-2-2), but much larger; leg tarsi longer, 
more Oncopus-iike; ventral process on palpal trochanter very long, bifurcate; 
chelicerae robust, proximal article with pointed prodorsal process on distal corner, 
second article with ventral process (as in Oncopus spp.). Judging from external 
morphology, this specimen appears intermediate between Caenoncopus gen. n. and 
Oncopus. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 549 

7. 1 9, Philippines, Leyte, Visca north of Baybay, 200-500 m, 23.11.1991, leg. 
J. Martens & W. Schawaller (MAR) — similar to B. fuscus sp. n. (tarsi 2-2-3-3) but 
much larger; eye tubercle higher; ventral palpal trochanter and femur with conical 
processes; proximal article of chelicerae with erect dorso-median process. Syntopic 
with G. leyteensis sp. n. 

8. 1 9, Philippines, Luzon, Sagada, 9.1.1980, leg. L. Deharveng (MAR) — similar to 
B. fuscus sp. n. but distinctly larger; ventral process on palpal trochanter smaller. 
Syntopic with G. maculatus sp. n. 

9. 1 9, Malaysia, Perak, Taiping, Maxwell Hill, ca. 1200 m, 10.IV.1990, leg. A. 
Riedel (MAR) — similar to G. hirsutus sp. n. (tarsi 2-2-3-3) but smaller, less hairy 
and without distinct ventral process on proximal palpal femur. Distinct from 
G. laruticus sp. n., which occurs at the same locality. 

DISCUSSION 

Genital morphology 

The description of Caenoncopus cuspidatus (Schwendinger 1992) already 
showed that penis morphology in Oncopodidae is not as uniform as previously 
assumed and the new material uncovers even more complex conditions. At present, 
we can distinguish four major penis types. Three of them are characteristic for one 
genus each, the remaining one is shared by two genera. 

Hypothetical ancestor. Our interpretation of phylogenetic relationships within 
the Oncopodidae is rooted in an ancestral penis type (Fig. 134a). Archaic Onco- 
podidae presumably possessed a cylindrical truncus and a short distad-directed glans 
with membraneous, slightly movable tubes. From this hypothetical form the four 
extant penis types may have evolved. These are: 

Type 1. In the species of Palaeoncopus gen. n. the penis is structurally almost 
identical with the ancestral form (Fig. 134a), except for the lack of membraneous 
tubes (Fig. 134b). 

Type 2. In Biantoncopus gen. n. (only known species B. fuscus sp. n.) the 
penis is similar to type 1, but stylus and membraneous tubes can be expanded by 
means of hemolymph pressure (Fig. 134c). During expansion the stylus is protruded 
distally and the membraneous tubes are bent to the opposite direction towards the 
base of the truncus. Structurally and functionally, this construction is very similar to 
other "hemolymph-pressure" lanitorean families (Martens 1986). 

Type 3. The penes of Gnomulus and Oncopus possess the same elements as 
the hypothetical ancestor, but the whole glans is proximad-directed (Fig. 134d). This 
penis form was previously considered typical for the whole family (Martens 1976, 
1986); it is found in the majority of species. A few of them, however, are already 
derived in that they have acquired an enlarged, prolonged "atypical" stylus (Fig. 134e; 
e.g. G. maculatus sp. n., G. crucifer sp. n.) and/or reduced the median plate (e.g. 
G. leyteensis sp. n., G. crucifer sp. n.). 



550 



J. MARTENS & P. SCHWENDINGER 




Fig. 134 

Hypothetical evolution of the four penis types in Oncopodidae. - a) hypothetical ancestor with 
glans parts directed distad; b) type 1: Palaeoncopus gen. n., similar to state a), but 
membraneous tubes (see Fig. 1) lost; c) type 2: Biantoncopus gen. n. with parts of glans 
expandable; d) type 3: Gnomulus and Oncopus, glans bent proximad; e) Gnomulus crucifer sp. 
n., parts of the glans (especially stylus) noticeably elongated; f) type 4: Caenoncopus gen. n. 
with median plate and lateral sclerites reduced, stylus strongly elongated, embraced by 
proximal sheath. - This scheme does not reflect phylogenetic relationships, but shows trends in 
the evolution of penis types in Oncopodidae. Caenoncopus gen. n., for example, probably has 
not evolved directly from Oncopus or Gnomulus, but from a common ancestor with a penis as 
in d (see Discussion: Evolution). 



Type 4. The penes of Caenoncopus gen. n. species are structurally unique and 
quite different from the other types. The truncus is dorso-ventrally depressed, its base 
not constricted and ending in two lobes. The glans essentially comprises only a very 
long, thick proximad-directed glans with an asymmetrical apex. Proximally, the stylus 
is embraced by a sheath-like membraneous collar formed by an extension of the 
membraneous socket; lateral sclerites, membraneous tubes and median plate are 
absent (Fig. 134f). This hypertrophic stylus is only partly homologous to the styli of 
the other penis types described above. 



new genera and species of oncopodidae 55 1 

Evolution 

Our interpretation of taxonomic characters leads to the following possible 
evolutionary scenario. Primitive Oncopodidae were small (2-5 mm), sironoid-like 
opilionids, with large dorsal and ventral scuta, weak chelicerae and few leg tarsalia. 
The ancestral oncopodid penis was devoid of muscles and already characteristic for 
the derived group of "hemolymph-pressure" Laniatores. It had a short, distad-directed 
subapical glans with a free slender stylus surrounded by membraneous tubes and by 
lateral sclerites connected to a median plate (Fig. 134a). A similar penis morphology 
as presumed for early oncopodids is characteristic for two other families of "hemo- 
lymph-pressure" Laniatores, i.e. Gonyleptidae and Cosmetidae (Martens 1976, 
1986). We take them as outgroup for our interpretation of relationships within the 
Oncopodidae. In these opilionids the subdistal glans is movable by means of hemo- 
lymph pressure only to a limited extent. As the stylus is pointing straight forwards, 
there is no need for a greatly movable glans. The penes of Palaeoncopus gunung sp. 
n., P. kerdil sp. n. and P. katik sp. n. from Sumatra largely accord with this primitive 
penis form, but are apomorphic in the loss of their membraneous glans tubes 
(Fig. 134b). Biantoncopus fiiscus sp. n. from the Philippines also possesses a distad- 
directed glans penis (with membraneous tubes still present), but it has become 
expandable (Fig. 134c) in the same way as shown for the Biantidae (Martens 1978: 
figs 8a-e, 9a-d). The expanding stylus probably increases the insemination rate by 
enabling the S to ejaculate deeper into the ? ovipositor. 

From the primitive, generalized penis type with short distad-directed, non- 
expandable glans, the evolutionary trend in "hemolymph-pressure" Laniatores went 
towards enlargment and enhanced movability of the stylus. To keep the penis under the 
genital operculum and protect it from damage, it became necessary to either transpose 
the glans to a more proximal position on the truncus, or to retreat it deeply into the 
distal part of the truncus, or to fold it downwards. The latter option was realized in the 
majority of oncopodid species, the former ones in different lineages, i.e. in the 
Fissiphalliidae, Assamiidae, Biantidae and Podoctidae (Martens 1986, 1988). After the 
glans became newly adjusted to a position with the stylus pointing to the truncus basis 
(Fig. 134d), evolution in Oncopodidae apparently went in two directions. Caenoncopus 
gen. spp. n. from Sumatra remained small, but greatly modified their penis morphology. 
In these species the lateral sclerites, median plate and membraneous tubes of the glans 
were lost, whereas the stylus gained enormous proportions and became asymmetrical 
(Fig. 134f). The rest of the Oncopodidae retained the typical penis structure (with short 
proximad-directed glans) but diversified in external characters instead. Most of them 
increased size of body and chelicerae (most pronounced in Oncopus) and acquired 
different tarsal formulas (2-2-2-2, 2-2-3-3 in Gnomulus, 1-1-1-1 in Oncopus). The 
evolutionary trend for stylus enlargement and reduction of glans sclerites, parallel to 
that in Caenoncopus gen. n., is also evident in the Gnomulus-Oncopus lineage, as 
shown in G maculatus sp. n. (Figs 84-86) and G. crucifer sp. n. (Figs 73-75, 134e). 

Divergent evolutionary lineages resulted in four distinctly different penis 
types, but specific distinctions within the same type are often minute. Considering 



552 J - MARTENS & P. SCHWENDINGER 

also the rapidly increasing number of newly discovered species with restricted 
distribution, it appears that the Oncopodidae currently undergo an active process of 
speciation. Unlike most other Laniatores, specific distinctions in Oncopodidae are 
expressed in penis modifications rather than in external morphology. Consequently, 
describing new taxa from 9 9 or juveniles should be strictly avoided. 

Generic limits 

Our reassessment of the Oncopodidae is primarily based on penis morphology, 
which is clearly outlined in the genera newly established in this paper. Only in 
Gnomulus penis shapes are fairly variable, with some species (e.g. G crucifer sp. n.) 
being already as far derived from the usual type that they could be regarded as 
generically different. The penes of Gnomulus and Oncopus, on the other hand, are of 
the same type and no relevant distinctions could be found. Externally, however, 
Oncopus is clearly distinguished by its unique tarsal formula (1-1-1-1) and its robust 
body. Tarsal formulas, however, do not entirely correspond with genital characters 
used for generic distinctions. Different tarsal formulas exist in genera clearly defined 
by genital characters (1-1-2-2 and 1-1-3-3 in Caenoncopus gen. n.; 2-2-2-2 and 2-2-3- 
3 in Gnomulus) and the same tarsal formulas are present in different genera (1-1-3-3 
in Caenoncopus gen. n. and Palaeoncopus gen. n., 2-2-3-3 in Biantoncopus gen. n. 
and Gnomulus). Consequently, the only reliable traits available at present to delineate 
genera within the Oncopodidae are genital characters. 

The five oncopodid genera that we recognize here are founded on the 
following putative autapomorphies: 

Palaeoncopus gen. n. - glans penis without membraneous tubes; palpal trochanter 
with prodorsal process. 

Biantoncopus gen. n. - glans partly retreated into the distal part of the truncus, 
expandable by means of hemolymph pressure. 

Gnomulus Thorell - glans proximad-directed. No relevant distinctions for Pelitnus 
Thorell, therefore placed in synonymy with Gnomulus. 

Oncopus Thorell - glans proximad-directed (but this is possibly synapomorphic with 
Gnomulus). Another possible autapomorphy is found in the strong chelicerae 
with 1-2 ventral processes on the cheliceral hand. At present, however, we 
cannot recognize any clear autapomorphies, which distinguish Oncopus from 
Gnomulus. External morphology (especially tarsal formula 1-1-1-1) is very 
characteristic for Oncopus, but it is not clear whether this (except for the large 
body size) is plesiomorphic or apomorphic. 

Caenoncopus gen. n. - stylus strongly enlarged with membraneous collar and asym- 
metrical apex; other parts of glans lost. 

Functional morphology of the oncopodid penis 

The oncopodid penis operates by hemolymph pressure. Like in most other 
families of Laniatores, muscles and tendons are absent. Internal pressure of the body, 
transmitted to the penis via hemolymph, causes positioning of the glans in relation to 



NEW GENERA AND SPECIES OF ONCOPODIDAE 553 

the truncus. Structure and alignment of the subdistal glans was previously regarded as 
fairly uniform. Only penis type 3 (Fig. 134d) was known and just a small degree of 
upward movement (less than 180°) by the short proximad-directed glans was thought 
to be possible (Martens 1986). 

With the new material, greatly different conditions within the Oncopodidae are 
pointed out. Some of the new species exhibit penis forms with structural and 
functional characteristics as otherwise found only in different well-defined opilionid 
families (Fig. 134). 

A similar short, distad-directed glans (considered plesiomorphic for the 
"hemolymph-pressure" Laniatores) as in Palaeoncopus gen. n. is present in Cos- 
metidae and Gonyleptidae. Within the primitive penis types derived forms can be 
identified, in which hemolymph pressure protrudes the stylus forward to reach the 9 
receptacula seminis more efficiently during copulation. Such is found in Biantonco- 
pus fuscus sp. n., Assamiidae, Biantidae and Podoctidae. A short, proximad-directed 
glans, comparable to Gnomulus and Oncopus, is typical for many Phalangodidae, an 
obviously polyphyletic group. In separate evolutionary lineages the styli became 
increasingly enlarged. This occurs in the species of Caenoncopus gen. n. and, pro- 
bably parallel to it, also in few Gnomulus species. Comparable hypertrophies of the 
stylus are present also in the family Triaenonychidae, where this phenomenon appar- 
ently evolved independently several times in different Australian and South American 
clades. In this family (belonging to Laniatores with a muscle-tendon complex inside 
the penis), the distad-directed stylus seems to be only slightly movable. In the cases 
mentioned, the stylus became a strongly elongated part of the truncus, but it was 
never folded downwards (Hunt & Maury 1993). 

The different penis types require modifications in mating behaviour. In this 
study, we did not examine the ovipositors of Oncopodidae in detail, but random 
samples showed that they are unsegmented, generally short and only movable to a 
small extent, if at all (Martens et al. 1981, observations from one Oncopus species). In 
terms of inner skeletal anatomy, the oncopodid ovipositor is very similar to that of other 
Laniatores. 

In order to transfer sperm into the receptacula within the ovipositor, the â needs 
to adjust his stylus to a suitable position by means of hemolymph pressure. This is done 
by slightly protruding the stylus in penis type 1 (Fig. 134b), or by inflating the 
membraneous tubes and strongly protruding the stylus in penis type 2 (Fig. 134c), or by 
folding the entire glans upwards about 180° (while only slightly protruding the stylus) 
in type 3 (Fig. 134d, e). It is not quite clear what happens during copulation in penis 
type 4 (Fig. 134f). Obviously the penis first has to be pushed out of the genital orifice 
for almost its entire length until the long stylus (Figs 5a, b, 9-10, 18-19) lies free and 
can be folded upwards. In Caenoncopus cuspidatus, with its glans only little shorter 
than the truncus, probably the maximum length of a proximad-directed glans is reached. 
To unfold this enormous sub-organ of the penis, the animal presumably has to rise high 
on its "tiptoes" and lift up its body quite a distance from the substrate. What movement 
these S S actually perform during mating can only be learned from observations on 
living animals. 



554 J - MARTENS & P. SCHWENDINGER 

ACKNOWLEDGEMENTS 

We are grateful to Dr Bernd Hauser (MHNG) for his persistent incitement to 
take up this study and for the loan of specimens. Further material was kindly made 
available by the following colleagues: Dr Ben Brugge (ZMA), Dr Christa Deeleman- 
Reinhold (Sparrenlaan), Dr Louis Deharveng (Université Paul Sabatier, Toulouse), 
Dr Giuliano Doria (MCSNG), Dr Manfred Grasshoff (SMF), Dr Pekka Lehtinen 
(ZMT), Dr Ivan Lobi (MHNG), Dr Hirotsugu Ono (National Science Museum, 
Tokyo), Mr Alexander Riedel (Zoologische Staatssammlung, München), Dr 
Wolfgang Schawaller (Staatliches Museum für Naturkunde, Stuttgart). Dr Konrad 
Thaler provided literature; Mr Konrad Eller, Mr Willi Salvenmoser and Mr Karl 
Schatz (all University of Innsbruck) helped producing SEM-micrographs. Dr J. 
Margraf (formerly Visca, Leyte) made a research stay of J.M. in Leyte, Philippines 
very successful. Mrs Käthe Rehbinder (Mainz) drew all whole-animal figures. Dr 
Stephen Elliott (Chiang Mai University) helped to improve the English text. The 
German Academic Exchange Service (Deutscher Akademischer Austauschdienst) 
supported P.S. with a research grant to work in Mainz for one month. The Feldbausch 
Foundation at "Fachbereich Biologie" of Mainz University granted travel funds to 
J.M. Our sincere thanks are due to all friends, colleagues and institutions mentioned. 



REFERENCES 

Hunt, G. S. & Maury. E. A. 1993. Hypertrophy of male genitalia in South American and 
Australian Triaenonychidae (Arachnida: Opiliones: Laniatores). Memoirs of the 
Queensland Museum 33: 551-556. 

Loman, J. C. C. 1892. Opilioniden von Sumatra, Java und Flores. In: Weber, M. (ed.). 
Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien 3: 1-26. Leyden. 

Loman, J. C. C. 1902. Neue aussereuropäische Opilioniden. Zoologische Jahrbücher, Abteilung 
für Systematik 16:1 63-2 1 6. 

Loman, J. C. C. 1903. Vergleichend anatomische Untersuchungen an chilenischen und anderen 
Opilioniden. Zoologischer Jahresbericht, herausgegeben von der Zoologischen Station 
zu Neapel (Berlin). Supplement 6: 1 17-120. 

Martens, J. 1976. Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida: 
Opiliones). Entomologica Germanica 3: 51-68. 

Martens, J. 1977. Opiliones aus dem Nepal-Himalaya. III. Oncopodidae, Phalangodidae, Assa- 
miidae (Arachnida). Senckenbergiana biologica 57: 295-340. 

Martens, J. 1978. Opiliones aus dem Nepal Himalaya. IV. Biantidae (Arachnida). Senckenberg- 
iana biologica 58: 347-414. 

Martens, J. 1980. Versuch eines phylogenetischen Systems der Opiliones. In: Gruber J. (ed.). 
Verhandlungen, 8. Internationaler Arachnologen-Kongreß, 355-360. Egermann, Wien. 

Martens, J. 1986. Die Großgliederung der Opiliones und die Evolution der Ordnung (Arach- 
nida). In: Barrientos, J. A. (ed.). Actas 10 Congreso Internacional de Aracnologia, 
Jaca/Espana 1: 289-310. Instituto Pirenaico de Ecologia & Grupo de Aracnologia, Bar- 
celona. 

Martens, J. 1988. Fissiphalliidae, a new family of South American Laniatorean harvestmen 
(Arachnida: Opiliones). Zeitschrift für zoologische Systematik und Evolutionsforschung 
26: 144-127. 



NEW GENERA AND SPECIES OF ONCOPODIDAE 555 



Martens, J., Hoheisel, U. & Götze, M. 1981. Vergleichende Anatomie der Legeröhren der 

Opiliones als Beitrag zur Phylogenie der Ordnung (Arachnida). Zoologische Jahr- 
bücher, Abteilung für Anatomie 105: 13-76. 
PococK, R. I. 1897. Descriptions of some new Oriental Opiliones recently received by the 

British Museum. Annals and Magazine of Natural History, including Zoology, Botany 

and Geology 19: 283-292. 
Roewer, C.-F. 1923. Die Weberknechte der Erde. Systematische Bearbeitung der bisher 

bekannten Opiliones. Fischer, Jena, IV+1 1 16 pp. 
Schwendinger, P. J. 1992. New Oncopodidae (Opiliones, Laniatores) from Southeast Asia. 

Revue suisse de Zoologie 99: 177-199. 
Silhavy, V. 1960. Die Grundsätze der modernen Weberknechttaxonomie und Revision des 

bisherigen Systems der Opilioniden. Verhandlungen des XI internationalen Kongresses 

für Entomologie in Wien (17-25 August 1960) 1: 262-267. 
S0RENSEN. W. 1932. Descriptiones laniatorum (Arachnidorum Opilionum Subordinis) fecit 

William S0rensen opus posthumum recog'novit et edidit Kai L. Hendriksen. Mémoires 

de l'Académie Royale des Sciences et des Lettres de Danemark, Copenhague (Section 

des Sciences) (9) 3 (4): 199-422. 
Suzuki, S. 1969. On a collection of opilionids from Southeast Asia. Journal of Science of the 

Hiroshima University, Series B, Div. 1 (Zoology) 22: 11-77. 
Suzuki, S. 1977. Report on a collection of opilionids from the Philippines. Journal of Science 

of the Hiroshima University, Series B, Div. 1 (Zoology) 27: 1-120. 
Suzuki, S. 1982. Contributions to the taxonomy and zoogeography of the Opiliones of the 

Philippines, Bismarck and Solomon Islands. With an appendix on some related species 

from the Moluccas and Sumatra. Steenstrupia 8 (8): 181-225. 
Thorell, T. 1876. Descrizione di alcune specie di Opilioni dell'Archipelago Malese appar- 
tenenti al Museo Civico di Genova. Annali del Museo civico di Storia Naturale di 

Genova (Serie 1)9: 111-138. 
Thorell, T. 1890. Aracnidi di Pinang raccolti nel 1 889 dai signori L. Loria e L. Fea. Annali del 

Museo Civico di Storia Naturale di Genova (Serie 2) 10: 269-383. 
Thorell, T. 1891. Opilioni nuovi o poco cognosciuti dell'Archipelago Malese. Annali del 

Museo Civico di Storia Naturale di Genova (Serie 2) 10: 669-770, plates Vili + IX. 
Zink, R.M. 1997. Species concepts. Bulletin of the British Ornithologists' Club 1 17: 97-109. 



Revue suisse de Zoologie 105 (3): 557-568; septembre 1998 



Redescription of Betta pietà (Teleostei: Osphronemidae) 
and description of B. falx sp. n. from central Sumatra 



TAN Heok Hui 1 & Maurice KOTTELAT *> 2 

1 School of Biological Sciences, National University of Singapore, Kent Ridge, 
Singapore 119260, Republic of Singapore. 

2 Route de la Baroche 12, Case postale 57, CH-2952 Cornol, Switzerland. 



Redescription of Betta pietà (Teleostei: Osphronemidae) and descrip- 
tion of B. falx sp. n. from central Sumatra. - Betta pietà is redescribed 
on the basis of material from Java. Betta falx sp. n. is described from 
Jambi, central Sumatra. The new species differs from B. pietà by having 
fewer lateral scales (27 vs. 27-30), a narrower head profile and by lacking 
median caudal-fin extensions in mature males. 

Key-words: Betta - Osphronemidae - Sumatra - taxonomy - biodiversity. 



INTRODUCTION 

Betta trif asciata Bleeker, 1850, the type species of the genus Betta by mono- 
typy, was described from Ambarawa, Java (Bleeker 1850). However, Valenciennes 
(in Cuvier & Valenciennes 1846) had earlier described the same fish from 
Buitenzorg (now Bogor), Java, as Panchax pictum and, therefore B. trifasciata is a 
junior subjective synonym of B. pietà. The type material of B. pietà is lost and the 
type(s) of B. trifasciata is(are) apparently mixed up with other Betta species from 
several localities (RMNH 6370, 30 ex., 31.8-77.5 mm SL; Indonesia Archipelago; 
P. Bleeker, bought 1879) and are in a bad condition (pers. obs.). Roberts (1993: 72, 
fig. 39) published a watercolour copy of the original drawing by Kuhl & van Hasselt 
(labeled as Panchax pictum) on which Valenciennes had based his description. A 
similar looking species, previously identified as B. pietà, from Jambi, central 
Sumatra, is here described as a new species. A redescription of B. pietà is also 
presented. 

The B. pietà group was first established by Witte & Schmidt (1992) and later 
emended by Tan & Ng (in press). All members of the B. pietà group share the 
common features of darkly pigmented anal and caudal distal fin margins, iridescent 
opercle, and a relatively small adult size (up to 60 mm TL). The B. pietà group has 
the unique combination of characters: total anal-fin rays 21-26, dorsal-fin rays 8-10, 



Manuscript accepted 06.02.1998 



558 TAN HEOK HUI & MAURICE KOTTELAT 

subdorsal scales 5-6, lateral scales 27-30, and vertebrae 27-29. The species group 
currently contains four species: B. pietà, B. taeniata Regan, 1910, B. simplex Kottelat, 
1994, and B.falx sp.n. 

MATERIAL AND METHODS 

The species concept used here is the phylogenetic species concept (Cracraft 
1989; Warren 1992; see discussion in Kottelat 1997). For practical purposes, and 
especially as used in the present context, the phylogenetic species concept does not 
differ significantly from the evolutionary species concept; see Mayden & Wood 
(1995) for a discussion of the hierarchy of species concepts. Species recognition in 
the genus Betta is further discussed in Kottelat & Ng (1994: 65-67) who also 
comment on the use of colour marks as diagnostic characters and the limited or lack 
of use of morphometric characters. Systematic research may take different forms, and 
different types of publications may fulfill different purposes; our priorities in docu- 
menting biodiversity has been discussed in Kottelat (1995b, 1997). We call species 
group (abbreviated group) an assemblage of species sharing a set of diagnostic 
characters, which may or may not be a monophyletic lineage. In some cases, available 
data may support the monophyly of a given group, while for other groups such data 
are (still) missing or have not yet been re-evaluated. Our inability to demonstrate 
monophyly today does not automatically imply that a given group is not mono- 
phyletic, just that we do not know. Meanwhile, the recognition of groups, be they for 
convenience only, is justified by the necessity of handling subsets of the genus Betta, 
for example for comparing a species with those it seems related with and avoiding 
trivial and lengthy comparisons with completely and obviously unrelated species. 

Methods for counts and measurements and the terminology for elements of the 
colour pattern follow Tan & Ng (in press), modified from Witte & Schmidt (1992). 
Chin-bar is used to refer to the dark thin stripe running under the preorbitai stripe, 
from the lower margin of the eye forwards and downwards to the throat. The 'second 
central stripe' is a faint or distinct stripe, continuous or interrupted, usually starting 
behind pectoral-fin base, situated 1-2 scales ventral to central stripe anteriorly and 
joining it posteriorly. The dark, narrow, often slightly curved, concentric bars perpen- 
dicular to the rays on the interradial membranes of the dorsal and caudal fins are 
called dorsal (respectively caudal) transverse bars. These descriptive terms follow that 
of Tan & Kottelat (1998). Measurements are point to point on the left side of fish 
and were obtained with a pair of dial calipers (0.05 mm accuracy). 

Specimens examined are deposited in the Muséum d'histoire naturelle, Genève 
(MHNG); Muzium Zoologicum Bogoriense, Bogor (MZB); Nationaal Natuurhisto- 
risch Museum, Leiden (RMNH); Instituut voor Systematiek en Populatiebiologie, 
Universiteit van Amsterdam (ZMA); Zoological Reference Collection, National Uni- 
versity of Singapore (ZRC); and the collection of the second author in Cornol (CMK). 



BETTA P1CTA AND B. FALK 



559 




Fig. 1. Betta pietà, male, 30.9 mm SL, ZMA 121.675, Java: Cipanas; right side, reversed. 
Fig. 2. Betta faix, holotype, MZB 9308, male, 32.7 mm SL; Sumatra: Jambi: Sungai Alai. 
FlG. 3. Betta falx, male, ca. 25 mm SL; Sumatra: Jambi: Pijoan, not preserved. 



560 



TAN HEOK HUI & MAURICE KOTTELAT 






Fig. 4. Schematic drawing of caudal fin and posterior part of anal fin of: a, Betta pietà, MZB 
1325, 32.5 mm SL. male; b, B.falx, ZRC 40974, 32.5 mm SL, male. 

Fig. 5. Dorsal, lateral and ventral views of head of: a. Betta pietà, ZMA 121.589, 31.6 mm SL, 
male; b, B.falx, ZRC 40974, 33.3 mm SL, male. 



BETTA PI CT A AND B. FALX 56] 



DESCRIPTIONS 

Betta pietà (Valenciennes, in Cuvier & Valenciennes, 1846) Figs 1, 4a, 5a 

Panchax pictum Valenciennes, in Cuvier & Valenciennes, 1846: 385. 

Betta tri/asciato Bleeker, 1850: 12; Günther 1861: 388; Regan 1910: 781. 

Betta pietà: Bleeker 1879: 26 (part); Weber & de Beaufort 1922: 360 (part); Witte & 

Schmidt 1992: 324 (key); Kottelat et al. 1993: 163 (part), plate 76; Roberts 1993: 

38, fig. 39. 

Material examined. ZMA 102.149, 10 ex., 22.3-31.6 mm SL, ZRC 40973, 2 ex., 29.9-32.0 
mm SL; Indonesia: Java, ponds near Trogon (ca. 6°21'S 106°34'E, ca. 30 km northwest of 
Bogor); M. Weber, 1888. - ZMA 121.691, 52 ex., 13.3-32.8 mm SL; Indonesia: Java, 
Buitenzorg (Bogor); M. Weber, 1888. - MZB 1325, 37 ex., 13.8-32.2 mm SL, ZRC 42495, 5 
ex., 28.0-32.1 mm SL; Indonesia: Java, Bogor, Tjidjeruk, Sawahbera; Sukardi, 24 Aug. 1970. - 
RMNH 10447, 3 ex., 29.3-32.4 mm SL; Indonesia: Java, Buitenzorg (Bogor); M. Weber. - 
RMNH 15794, 1 ex.. 36.4 mm SL; Indonesia: Java, Buitenzorg (Bogor), Tjiblagoeng; A. 
Bushkiel, 1935. - ZMA 121.675, 30.9 mm SL, male; ZMA 121.589, 13 ex., 19.5-34.3 mm SL, 
ZRC 40972, 2 ex., 29.9-31.6 mm SL; Indonesia: Java, Tjipanas (Cipanas: ca. 6°43'S 107°2'E, 
ca. 30 km southeast of Bogor) near Sindanglaja; M. Weber, 1888. - ZMA 121.694, 38 ex., 14.9- 
31.2 mm SL; Indonesia: Java, Tjinjiroean, after Poentjak Gedeh, ca. 1600m (Gunung Gede: ca. 
25 km southeast of Bogor); Kerkhoven, 1921. - MHNG 2090-92, 3 ex., 23.6-30.0 mm SL; 
Indonesia: Java, Sukabumi (ca. 40 km southeast of Bogor); Walsh, July 1930. - ZMA 121.693, 
2 ex., 28.8 mm SL; Indonesia: Java, Bandoeng (Bandung); Huysmans. - ZMA 121.689, 1 ex., 
24.5 mm SL; Indonesia: Java, Bandoeng (Bandung), ca. 700m; I. Jacobson, 1934. - RMNH 
26940, 4 ex., 31.8-35.3 mm SL; Indonesia: Java, Bezoeki (Besoki) (ca. 7°16'S 109°E, ca. 1050 
km southeast of Bogor); J. Semmelink, 1864-65. - RMNH 13698-13699, 2 ex., 32.6-35.2 mm 
SL; Indonesia: Java, Ambarawa (ca. 7°8'S 110°20'E); J. Sybrandi, 1933. - RMNH 10740, 4 
ex., 24.9-34.2 mm SL; Indonesian Archipelago; from P. Bleeker' s collections. 

Diagnosis. Betta pietà is distinguished from the other members of the species group 
in having iridescent yellow-gold opercle scales in male (vs. bluish or greenish), and 
anal and caudal fins with a narrow bluish distal band in live male (vs. broad band; 
bluish in B. taeniata and B. simplex, reddish in B. falx). Other characters distinguish 
B. pietà are listed under Affinities. 

Description. General appearance is shown in Fig. 1; meristics and morphometries of 
the B. pietà group are listed in Table 1. Body relatively slender (body depth 21.5- 
25.5 % SL), head long (head length 31.1-36.0 % SL). Dorsal and anal fins may be 
slightly pointed in male, more rounded in female and juvenile; dorsal fin placed 
relatively far back (predorsal length 63.0-67.6 % SL), anal-fin base length almost half 
of SL (42.6-48.4 % SL). Caudal fin of male with median rays sligthly elongated, 
caudal fin of female and juvenile rounded. Pelvic fin falcate with second filamentous 
ray short (23.8-33.0 % SL). 

Coloration. Life coloration illustrated in Linke (1990) and Kottelat (1994). 
Body light brown, head and dorsal part of body darker brown. Head with distinct pre- 
and postorbital stripes, chin-bar and second postorbital stripe present. Body with 
central and second central stripes present, with faint spot on middle of caudal fin base. 
Male with iridescent yellow-gold opercle scales. Unpaired fins reddish. Anal fin and 
lower half of caudal fin with a pale blue subdistal band and broad dark blue distal 
band; dorsal fin with faint transverse bars. Pectoral fin hyaline, pelvic fin reddish with 
white filament. Female with whitish opercle scales. Unpaired fins yellowish brown 



562 TAN HEOK HUI & MAURICE KOTTELAT 

with very narrow white distal margin on dorsal and anal fins. Anal fin with a narrow 
subdistal blue band; dorsal and anal fins with faint transverse bars; caudal fin with 
transverse bars very faint or absent in some specimens only (based on photographs in 
Linke 1990). 

Preserved specimens light brownish or yellowish. Opercle pattern distinct in 
both sexes, both preorbitai and postorbital stripes distinct, chin-bar and second 
postorbital stripe also distinct. Body with distinct central and second central stripes, 
with spot on middle of caudal fin base. Male with dark reddish distal band on anal and 
caudal fins, female and juvenile without dark margin. Dorsal and anal fins of only 
female and juvenile with transverse bars. 

Distribution. Betta pietà is known only from the western two-thirds of Java. The 
easternmost record in Central Java is Ambarawa (Tuntang basin, draining to the 
north). All known localities are in basins draining to the north, except Sukabumi, 
Western Java, which is in the Mandiri basin draining to the south. Betta pietà 
apparently inhabits hilly areas where it has been observed in slow-flowing side-waters 
of hill creeks (see Linke 1990, for habitat description). It is not clear whether this 
really is its preferred habitat. It might originally have had a larger distribution range in 
Java and anthropogenic pressure and habitat modification induced from intensive 
agriculture might be responsible for its restricted distribution. 

Affinities. Betta pietà is distinguished from B. taeniata in having iridescent yellow- 
gold opercle scales in male (vs. bluish-green opercle scales); anal and caudal fins with 
narrow bluish distal band in live male (vs. broad distal band); reddish caudal fin (vs. 
brownish); golden iris of eye (vs. dark brown); fewer modal vertebrae (27, vs. 28); 
fewer anal rays (21-23, mode 22, vs. 23-26, mode 26); fewer modal subdorsal scales 
(5V2, vs. 6); fewer modal lateral scales (28, vs. 29); dorsal-fin origin modally above 
13th scale of lateral series (vs. 14-15); more modal predorsal scales (20, vs. 19); 
smaller body depth (21.5-25.5 % SL, vs. 25.3-30.1); smaller anal-fin base length 
(42.6-48.4 % SL, vs. 47.9-52.7); slightly greater postorbital length (47.3-53.4 % HL, 
vs. 45.2-49.2); and slightly smaller interorbital width (26.0-31.1 % HL, vs. 29.2-36.2). 

Betta pietà is distinguished from B. simplex in having iridescent yellow-gold 
opercle scales in male (vs. greenish-blue opercle scales); anal and caudal fins with 
narrow bluish distal band in live male (vs. broad distal band); fewer transverse scales 
(9-9V2, mode 9, vs. 9V2-1 1 V2, mode IL/2); dorsal fin origin modally above 13th scale 
of lateral series (vs. 14-15); greater total length (135.4-143.9 % SL, vs. 132.1-137.2); 
smaller predorsal length (63.0-67.6 % SL, vs. 66.7-69.8); smaller caudal peduncle 
depth (14.4-17.3 % SL, vs. 17.8-19.3); smaller body depth (21.5-25.5 % SL, vs. 29.3- 
32.2); and smaller anal-fin base length (42.6-48.4 % SL, vs. 48.3-50.0). 

Betta pietà is distinguished from B. falx in having the anal and caudal fins with 
a narrow bluish distal band in live male (vs. reddish and broad band; Fig. 4); iri- 
descent yellow-gold opercle scales in live male (vs. greenish blue opercle scales); 
faint dorsal transverse bars in preserved male (vs. distinct); faint or absent caudal 
transverse bars in female (vs. distinct); head broad, width more or less constant (vs 
narrower anteriorly, resulting in a more pointed appearance in dorsal view; compare 



BETTA PICTA AND B. FALX 563 

Fig. 5a and 5b); narrow preorbitai black stripe (vs. thick); more modal dorsal rays (9, 
vs. 8); fewer modal subdorsal scales (5 l /i, vs. 6); more modal lateral scales (28, vs. 
27); dorsal fin origin modally above 13th scale of lateral series (vs. 12); anal fin 
origin modally below 7th scale of lateral series (vs. 6); more modal predorsal scales 
(20, vs. 19); slightly greater predorsal length (63.0-67.6 % SL, vs. 60.0-65.7); and 
slightly greater anal-fin base length (42.6-48.4 % SL, vs. 46.5-50.3). 





B. pietà 


B. falx 


B. taeniata 


B. simplex 


sample size 


12 


12 


16 


4 


MERISTICS 










[range (mode)] 










vertebrae 


2 + 8+ 17-18 


2 + 8+ 17-18 


2 + 8+ 17-19 


2 + 8+ 18 




total: 27-28 (27) 


total: 27-28 


total: 27-29 (28) 


total: 28 


anal-fin rays 


11,19-21 


II, 19-21 


I-II, 22-24 


11.21-22 




total: 21-23 (22) 


total: 21-23 (22) 


total: 23-26 (26) 


total: 23-24 (24) 


dorsal-fin rays 


0-1, 7-9 


III, 7-8 


Oil, 8-9 


0-1, 8-9 




total: 8-9 (9) 


total: 8-10(8) 


total: 8-10 ( 9) 


total: 9 


caudal-fin rays 


ii, 5 + 6, ii 


ii, 5 + 6, i-ii 
(ii, 5 + 6, ii) 


ii, 5 + 6-7, i-ii 
(ii, 5 + 7, i) 


i, 6 + 7, 1 


pelvic-fin rays 


I, 1,4 


I. 1,4 


I, 1,4 


I, 1,4 


pectoral-fin rays 


12 


12 


11-12(12) 


11-12(12) 


subdorsal scales 


51/2-61/2(51/2) 


51/2-61/2(51/2 or 6) 


5-6(6) 


5 


transverse scales 


9-9'A (9) 


9-9 'A (91/2) 


9-91/2(91/2) 


91/2-111/2(111/2) 


lateral scales 


27-30 (28) 


27 


27V 2 -30(29) 


27-28 (28) 


lateral scale below 










dorsal-fin origin 


12-14(13) 


11-12(12) 


12-15(14) 


15 


lateral scale above 










anal-fin origin 


6-8(7) 


6-8 (6) 


5-7 (7) 


- 


predorsal scales 


19-21 (20) 


19-20(19) 


19-21 (19) 


20 (20) 


postdorsal scales 


9-11 (10) 


9-11 (10) 


9-11 (10) 


9-10(10) 


MORPHOMETRICS 










percentage of SL [rar 


ige] 








total length 


135.4-143.9 


135.8-142.2 


132.9-141.6 


132.1-137.2 


predorsal length 


63.0-67.6 


60.0-65.7 


62.4-67.7 


66.7-69.8 


postdorsal length 


19.5-25.5 


22.7-25.7 


19.5-24.8 


23.2-25.1 


caudal peduncle depth 


14.4-17.3 


15.8-18.3 


14.8-19.7 


17.8-19.3 


preanal length 


48.7-53.6 


48.7-52.3 


47.2-54.1 


51.1-52.7 


head length 


31.1-36.0 


33.1-37.9 


30.3-34.5 


34.6-37.2 


body depth 


21.5-25.5 


22.6-28.0 


25.3-30.1 


29.3-32.2 


pelvic-fin length 


23.8-33.0 


24.2-34.3 


23.5-37.3 


28.1-28.7 


anal-fin base length 


42.6-48.4 


46.5-50.3 


47.9-52.7 


48.3-50.0 


dorsal-fin base length 


10.2-13.0 


10.5-13.3 


10.8-15.1 


11.2-11.8 


percentage of HL [range] 








orbit diameter 


23.4-27.1 


24.2-28.1 


23.8-30.7 


- 


postorbital length 


47.3-53.4 


46.1-53.4 


45.2-49.2 


- 


interorbital width 


26.0-31.1 


24.7-30.5 


29.2-36.2 





Table 1 . Meristic and morphometric data of the Betta pietà species group 



564 TAN HEOK HUI & MAURICE KOTTELAT 

Remarks. Betta pietà is the only species of the genus naturally occurring on Java. All 
the preserved specimens we have examined seem to belong to a single species. We 
cannot exclude, however, that once live specimens from throughout the species' range 
become available, the status of these populations might have to be revised. 

Previous workers have included in B. pietà numerous populations from 
Sumatra (Bleeker 1879; Weber & de Beaufort 1912, 1922; Kottelat et al. 1993). 
These Sumatran populations are described below as B. falx. 



Betta falx sp. n. Figs 2, 3, 4b, 5b 

Betta pictum: Weber & de Beaufort 1912: 541. 

Betta pietà: Bleeker 1879: 26 (part); Weber & de Beaufort 1922: 360 (part); Kottelat et al. 
1993: 163 (part). 

Holotype. MZB 9308, 32.7 mm SL, male; Indonesia: Sumatra: Jambi Prov.: Sungai Alai, km 
19.5 on Muara Bungo - Muara Tebo road (bridge at Sungai Alai: 1°28 , 42.6"S 102°18'31.7"E); 
H. H. Ng & S. H. Tan. 22 June 1995. 

Paratypes. All from Indonesia: Sumatra: Jambi Prov.: ZRC 40974, 20 ex., 17.2-33.0 mm SL, 
ZMA 121.673, 6 ex., 21.8-31.0 mm SL, MZB 9307, 4 ex., 17.5-26.3 mm SL, RMNH 33087, 5 
ex., 19.7-26.7 mm SL; same data as holotype. — ZRC 42496. 10 ex., 17.0-34.9 mm SL, 
MHNG 2593.95, 5 ex.. 29.7-30.1 mm SL; same locality as holotype; H. H. Tan & H. H. Ng, 22 
Jul 1997. — ZRC 38571. 44 ex.. 13.8-32.4 mm SL, CMK 11119. 44 ex., 10.5-30.8 mm SL; 
Sungai Alai at km 28 on Muara Bungo - Muara Tebo road, between half hour downriver of 
bridge to ca. 1 hour upriver, including small tributaries and Danau Gresik; M. Kottelat & H. H. 
Tan, 30-31 May 1994. — ZRC 38254. 3 ex.. 31.7-36.1 mm SL; Sungai Alai; M. Kottelat & H. 
H. Tan, May 1994. 

Other material (non type). All from Indonesia: Sumatra: ZMA 121.590, 29 ex., 17.1-27.4 
mm SL, ZRC 40975, 2 ex., 25.2-25.6 mm SL; East Sumatra, Deli [Medan]; L. P. de Bussy - Le 
Cosquino, 1905-1920. — ZMA 121.692, 5 ex., 23.7-32.0 mm SL; Aceh: Sungai Gloegoer, 
beekje by Bohorot, Boven Langkat; L. P. de Bussy, Aug. 1917. — ZMA 121.688, 2 ex., 32.9- 
33.8 mm SL; Sumatra Utara: Serdang [Sungai Serdang: 3°42'N 98°52'E]; V. Dedem, 26 July 
1909. — ZRC 38497, 5 ex., 13.1-18.8 mm SL, CMK 1 1034, 5 ex.; Jambi: Danau Pinang, lake 
connected to Sungai Pijoan, 1 boat hour upstream of Pijoan (19 km W of Jambi on road to 
Muara Bungo); M. Kottelat & H. H. Tan, 28 May 1994. — ZRC 38506, 2 ex., 15.1-22.6 mm 
SL, CMK 11055, 2 ex.; Jambi: Sungai Pijoan just downriver of confluence with stream 
draining Danau Pinang; M. Kottelat & H. H. Tan, 28 May 1994. — ZRC 38590, 1 1 ex., 15.8- 
26.5 mm SL, CMK 1 1 136, 10 ex.; Jambi: Danau Kamining near Kampung Transos, ca. km 5 
southwards on road branching off road Muara Bungo - Muara Tebo at km 36; M. Kottelat & H. 
H. Tan, 31 May 1994. — CMK 1 1071, 1 ex.; Jambi, Sungai Keruh, ca. 2 km south of mainroad 
at km 23 on road Jambi - Muara Tembesi, tributary of Sungai Pijoan; M. Kottelat & H. H. Tan, 
28 May 1994. — ZRC 40976. 9 ex., 11.3-31.6 mm SL; Sungai Pijoan, 1°35'35.0"S 
103°27'07.2"E; H. H. Tan et al., 8 Jun. 1996. — ZRC 40977, 2 ex., 24.0-24.1 mm SL; Jambi, 
Pijoan, Danau Souak Padang, 1°36'34.4"S 103°26'55.1'"E; H. H. Tan et al.. 8 Jun. 1996. 

Diagnosis. Betta falx is distinguished from the other members of the B. pietà group in 
having anal and caudal fins with reddish distal band in live male (vs. blue), distinct 
dorsal transverse bars in male (vs. absent in B. taeniata and B. simplex, faint in 
B. pietà) and distinct caudal transverse bars in female (vs. absent in B. taeniata and 
B. simplex, faint or absent in B. pietà). Other characters distinguishing B. falx are 
listed under Affinities. 



BETTA PICT A AND B. FALX 565 

Description. General appearance as shown in Figs. 2-3; meristics and morphometries 
of the B. pietà group are listed in Table 1. Body relatively slender (body depth 22.6- 
28.0 % SL), head long (head length 33.1-37.9 % SL). Dorsal and anal fins may be 
slightly pointed in male, more rounded in female and juvenile; dorsal fin placed 
relatively far back (predorsal length 60.0-65.7 % SL), anal-fin base length almost half 
of SL (46.5-50.3 % SL). Caudal fin of male, female and juvenile rounded. Pelvic fin 
falcate with second filamentous ray short (24.2-34.3 % SL). 

Coloration. Life coloration illustrated in Fig. 3. Body light brown, head and 
dorsal part of body darker brown. Opercle pattern distinct in both male and female, 
both preorbitai and postorbital stripes distinct, chin-bar and second postorbital stripe 
also distinct. Body with distinct central and second central stripes, with spot on 
middle of caudal fin base. Male with greenish-blue iridescent opercle scales. Unpaired 
fins yellowish. Anal fin and lower half of caudal fin with pale reddish subdistal band 
and broad reddish-brown distal band with very thin white margin; dorsal fin with 
distinct transverse bars. Pectoral fin hyaline, pelvic fin hyaline with white filament. 
Female with yellowish opercle scales. Unpaired fins yellowish without narrow dark 
distal band on dorsal and anal fins. Dorsal, anal and caudal fins with distinct 
transverse bars. Juveniles (less than 20 mm SL) with distinct transverse bars on 
unpaired fins, distal dark margin on anal fin absent, usually with a rather marmorated 
pattern on body. Young males (less than 25 mm SL) with caudal transverse bars 
which disappear in adult males. 

Preserved specimens are brownish. Opercle pattern distinct in both male and 
female, both preorbitai and postorbital stripes distinct, chin-bar and second postorbital 
stripe also distinct. Body with distinct central and second central stripes, black spot on 
middle of caudal fin base. Male with dark reddish-brown distal margin on anal and 
caudal fins, female and juvenile without dark distal margin. Male with distinct dorsal 
transverse bars only; female and juvenile with distinct transverse bars on dorsal, anal 
and caudal fins. 

Distribution. Betta faix is known from the Langkat area and Medan in Sumatra 
Utara, and Jambi province in central Sumatra. Weber & de Beaufort (1912: 541; 
1922: 360, 362) recorded B. pietà from Palembang, Upper Langkat, Muara Kompeh 
(Jambi province) and Deli (now Medan). From these localities, they had access only 
to material from Upper Langkat (ZMA 121.692) and Deli (ZMA 121.590), which we 
have re-identified as B.falx. The other records were based on Bleeker (1879), but the 
material on which Bleeker based his records has now been mixed with Betta material 
of various species and localities and cannot be sorted (RMNH 6370, 30 ex.). 

Notes on biology. We have observed B. falx only in swamp forests in the Batang 
Hari basin, Jambi province. Specimens are typically found among submerged bank 
vegetation, in near stagnant waters, with pH 4.7-6.8. Syntopic osphronemids are 
Belontia hasseltii, Betta aff. fusca, Luciocephalus pulcher, Parosphromenus suma- 
tranus, Sphaerichthys osphromenoides, Trichogaster leerii, T. trichopterus and Tri- 
chopsis vinata. Betta falx adapts well to captivity, where it readily spawns with 
fortnightly intervals. It is a paternal oralbrooder (THH, pers. obs.). 



566 TAN HEOK HUI & MAURICE KOTTELAT 

Etymology. From the Latin falx, meaning scythe, alluding to the continuous curved 
shape of the broad anal and caudal distal margins of a male in display. A noun in 
apposition. 

Affinities. Betta falx is distinguished from B. taeniata in having iridescent greenish- 
blue opercle scales in male (vs. strongly coloured bluish-green opercle scales); anal 
and caudal fins with reddish distal band in live male (vs. blue); reddish fins (vs. 
brownish); distinct dorsal transverse bars in male and caudal transverse bars in female 
(vs. absence); fewer anal-fin rays (21-23. vs. 23-26); fewer modal dorsal-fin rays (8, 
vs. 9); fewer modal lateral scales (27, vs. 29); dorsal fin origin above 1 1- 12th scale of 
lateral series (vs. 12-15); anal-fin origin modally below 16th scale of lateral series (vs. 
7); and smaller interorbital width (24.7-30.5 % HL. vs. 29.2-36.2). 

Betta falx is distinguished from B. simplex in having anal and caudal fins with 
reddish distal band in live male (vs. blue); distinct dorsal transverse bars in male and 
caudal transverse bars in female (vs. absence); fewer anal-fin rays (21-23, vs. 23-24); 
fewer modal dorsal-fin rays (8, vs. 9); more modal subdorsal scales (5*/2-6, vs. 5); 
fewer transverse scales (9-9'/2. mode 9 l h, vs. 9 1 /2-ll i /2, mode IIV2); fewer modal 
lateral scales (27, vs. 28); dorsal-fin origin above 11- 12th scale of lateral series (vs. 
15); slightly greater total length (135.8-142.2 % SL, vs. 132.1-137.2); smaller 
predorsal length (60.0-65.7 % SL, vs. 66.7-69.8); greater postdorsal length (22.7-25.7 
% SL, vs. 23.2-25.1); and smaller body depth (22.6-28.0 % SL, vs. 29.3-32.2). 

Betta falx is distinguished from B. pietà in anal and caudal fins with broad 
reddish distal band in live male (vs. narrow and bluish; Fig. 4); iridescent greenish- 
blue opercle scales (vs. yellow-gold); preserved male with distinct dorsal transverse 
bars (vs. faint); female with distinct caudal transverse bars (vs. very faint or absent); 
head narrower anteriorly, resulting in a more pointed appearance in dorsal view (vs. 
head broad, width more or less constant; compare Figs. 5a and 5b); thick preorbitai 
black stripe (vs. narrow); fewer modal dorsal-fin rays (8 vs. 9); fewer modal lateral 
scales (27 vs. 28); dorsal fin origin above ll-12th scale of lateral series (vs. 12-14); 
anal fin origin modally below 6th scale of lateral series (vs. 7); fewer modal predorsal 
scales (19. vs. 20); and slightly greater anal-fin base length (46.5-50.3 % SL, vs. 42.6- 
48.4). 

Remarks. The Sumatra species identified by earlier workers (Bleeker, Weber & de 
Beaufort) as B. pietà is B. falx. Witte & Schmidt (1992: 324) recorded a 
"B. (edithae) sp. B" from Jambi. Our recent collections at numerous localities around 
Jambi did not yield any species resembling B. edithae; B.falx is the only species from 
this area which one might possibly consider as having some similarities with 
B. edithae. 

No species of the B. pietà group is presently known from Lampung, the 
southernmost province of Sumatra. Betta falx might occur there, but B. pietà could 
also be present. Some Javanese species also extend over a limited range in southern 
Sumatra (e.g., Nemacheilus fasciatus, see Kottelat 1984). 

Due to their close morphological similarity, B. pietà and B. falx are apparently 
closely related, their niche preference, however, is quite marked. Betta pietà is known 



BETTA PICTA AND B. FALX 567 

(so far) only from hill stream habitats, whereas B. falx is known (so far) only from 
lowland swamp forest habitats. As mentionned above, the possibility cannot be 
discounted that B. pietà occupies a secondary niche (a sort of refuge habitat) in the 
hill stream as a result from anthropogenic pressures. The quasi totality of the lowland 
and foothills of Java has lost its natural forest cover and has been converted into rice 
fields. In Sumatra and Borneo, lowland and foothill streams are the habitats with the 
most diverse fish communities; Kottelat (1995a: 422) noted that about half of the 
fish species recorded from Java at the beginning of the century have not been 
collected again in the last 40 years. Beside the possible effect of bias as crude as the 
lack of sampling effort, these extinction may simply reflect the disparition of forest 
and foot-hill streams. 



ACKNOWLEDGEMENTS 

We are very grateful to Gunawan 'Thomas' and Verawaty Kasim, for the gift 
of material and being such good hosts; Peter K. L. Ng, for his advice and constructive 
discussions; Sonia Muller, Claude Weber (MHNG), Daisy Wowor, Renny Hardiaty 
(MZB), Martin van Oijen (RMNH), Isaac Isbriicker (ZMA), Mrs. C. M. Yang and 
Kelvin K. P. Lim (ZRC), for access to and loan of specimens; Tommy H. T. Tan, 
S. H. Tan, Darren C. J. Yeo and H. H. Ng, for their help and companionship in the 
field; S. H. Tan for helping with the radiography of specimens; David M. Armitage 
for donating phototanks for THJTs use. This paper is part of the higher degree for 
THH whose work has been partially supported by research grants RP 950326 and 
RP960314 to Peter K. L. Ng from the National University of Singapore. This is 
contribution 6/97 of the Systematics and Ecology Laboratory, School of Biological 
Sciences, National University of Singapore. 



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Revue suisse de Zoologie 105 (3): 569-580; septembre 1998 



Importance of forest structures on four beetle families 

(Col.: Buprestidae, Cerambycidae, Lucanidae and phytophagous 

Scarabaeidae) in the Areuse Gorges (Neuchâtel, Switzerland) 1 



Sylvie BARBALAT 

Institut de Zoologie, Emile Argand 11, CH-2007 Neuchâtel, Switzerland. 



Importance of forest structures on four beetle families (Col.: Bupre- 
stidae, Cerambycidae, Lucanidae and phytophagous Scarabaeidae) in 
the Areuse Gorges (Neuchâtel, Switzerland). - The species richness and 
abundance of selected xylophagous (Buprestidae, Cerambycidae and 
Lucanidae) and rhizophagous or saprophagous beetles (phytophagous 
Scarabaeidae) were compared between various forest stands with different 
ecotone structures. Window traps and water traps were used to sample the 
beetles. Among the 65 captured species, 13 belonged to the Buprestidae, 
41 to the Cerambycidae, 8 to the phytophagous Scarabaeidae and 3 to the 
Lucanidae. Forest stand and ecotone type were found to have a significant 
influence on these beetle communities. In oak stands, typical species such 
as Plagionotus arcuatus, Anoplodera sexguttata or Anthaxia Salicis were 
found, while in beech forests Platycerus caprea was found as a charac- 
teristic species. Natural edges are characterised by grassland and shrub 
species such as Agapanthia violacea, Phytoecia cylindrica and Anthaxia 
nitidula. In artificial clearings, species living in old stumps such as 
Corymbia rubra, Prionus coriarius, Rhagium bifasciatum, or Anastran- 
galia sanguinolenta are common as well as species living in the small 
branches left after a cutting, the most common of which being Stenurella 
melanura. In order to conserve a high diversity of forest beetles, oaks 
should be favoured (it hosts 9 typical species in our study) and diversified 
structures such as natural edges and artificial clearings must be maintained 
or created. 

Key-words: Forest ecology - Buprestidae - Cerambycidae - Scarabaeidae 
Pleurosticti - Lucanidae - Swiss Jura - Bioindicators. 

INTRODUCTION 

Buprestids, cerambycids and lucanids have xylophagous larvae (Dajoz 1980). 
Depending on the species, the larvae can colonise living trees, dead wood or rotten 
stumps. On the other hand, phytophagous scarabaeids are rather rhizophagous or sapro- 
phagous as larvae and phytophagous as adults (Allenspach 1970). 



1 This paper is part of the author's PhD. 
Manuscript accepted 23.03.1998 



570 SYLVIE BARBALAT 

Xylophagous beetles are an important element of forest ecosystems. They 
actively participate in dead wood decomposition. The larval galleries facilitate wood 
colonisation by micro-organisms, which considerably increases their efficiency (Dajoz 
1980). 

Thanks to museum collections, it has been possible to elaborate the rarefaction 
of several, mainly spectacular, species since the end of the last century, although the 
forest surface in Switzerland has not decreased during that time. The decline of many 
species can be attributed to coniferous tree plantings at low altitude, mainly spruce 
(Picea abies), instead of broad-leaved tree forests, causing a considerable loss of 
habitats for lowland species (Speight 1989). Forest trees are cut down before reaching 
an age at which they become attractive for xylophagous fauna and isolated old trees 
have almost disappeared. Regression of traditional orchards and humid habitats has also 
caused the rarefaction of specialized species (Geiser 1984). For species colonizing 
more open biotopes, mainly among scarabaeids, agriculture intensification has also 
been a cause of decline (Allenspach 1970). 

Some authors have worked on the influence of forest structures as well as 
woodland type on beetle communities. In Poland, for instance, Gutowski (1986) 
compared the fauna of a virgin forest with the fauna of a managed one. The changes in 
communities of cambio- and xylophagous insects in different age classes of forest 
stands have also been studied (Starzyk 1977; Starzyk & Witkowski 1981; Gutowski 
1995). Starzyk (1976; 1979) and Gutowski (1985) have studied the cerambycid 
communities occurring in different forest associations. 

In Switzerland, the importance of dead wood quantity for xylophagous beetles 
was underscored by Hartmann & Sprecher (1990). Barbalat (1996) and Barbalat 
& Borcard (1997) have shown the importance of artificial clearings on xylophagous 
beetles in managed forests. 

The aim of this work is to study, among different edges and clearings, what 
structures are most favourable for this fauna, in order to be able to make proposals to 
promote a forest management respecting biodiversity as much as possible. If buprestids, 
cerambycids and lucanids are good indicators for forest biotopes, only a few species are 
adapted to forest edges. For this reason, we consider as well a family, the scarabaeids, 
linked to more open habitats. 

MATERIALS AND METHODS 

Study area 

Our study was carried out in the Areuse Gorges near Neuchâtel (Western 
Switzerland) on the first Jura slopes (Fig. 1). We chose twelve sites in three forest 
types: oak, beech and mixed stands. Mixed forests are usually constituted of broad- 
leaved trees such as beech, oak and maple (Acer pseudoplatanus) mixed with 
coniferous trees such as spruce, pine and fir (Abies alba). These mixed forests are 
usually located on thin calcareous soils where tree growth is weak. In these stands, 
foresters create artificial clearings ranging from 650 to 10.000 m 2 , in order to favour 
pines which grow quite well on shallow soils and require much light to grow. In these 



IMPORTANCE OF FOREST STRUCTURES ON BEETLES 



571 




Fig. 1. Study area. After Barbalat (1997) 



station 


locality 


coordonnâtes 


altitude 


expo 


slope 


forest type 


cover 


habitat type 


edge type 


1 


Corcelles 


556125 204250 


695 m 


S 


20% 


oak 


80% 


edge 


natural 


2 


Colombier, Chanet 


555600 202950 


555 m 




0% 


beech 


40% 


artificial clearing 


clean 


3 


Chambrelien, Plan du Bois 


553250 202700 


670 m 


SE 


40% 


mixed 


60% 


artificial clearing 


clean 


4 


Boudry, Chanet 


553000 201150 


550 m 




0% 


oak 


30% 


artificial clearing 


clean 


5 


Chambrelien, Chassagne 


552000 201900 


725 m 


SE 


20% 


mixed 


40% 


artificial clearing 


clean 


6 


Chambrelien, Chassagne 


551900 202250 


770 m 


SE 


15% 


mixed 


60% 


artificial clearing 


clean 


7 


Rochefort, Les Grattes 


552470 204070 


880 m 


SE 


80% 


mixed 


40% 


edge 


natural 


8 


Fretereules 


549050 201600 


860 m 


SE 


45% 


beech 


90% 


edge 


natural 


9 


Brot-Dessous 


547600 200950 


890 m 


S 


40% 


beech 


90% 


edge 


clean 


10 


Brot-Dessous 


547850 200900 


790 m 


S 


40% 


beech 


50% 


natural clearing 


natural 


11 


Chambrelien, Bois-Devant 


553500 202450 


620 m 


SE 


15% 


mixed 


70% 


artificial clearing 


clean 


12 


Rochefort, Chaumes 


551375 202625 


840 m 


SE 


60% 


beech 


70% 


artificial clearing 


clean 


expo = exposure; cover = tree cover. 



















Table 1. Site description. After Barbalat (1997) 



clearings all the trees are cut down except a few selected pines, which are left for their 
seeds. Similar clearings are also created in oak stands in order to favour these trees 
which also need much light for their growth. This type of clearing is not made in beech 
forests, because beech can grow in more shady conditions. All the sites have been 
chosen either in clearings or forest edges with South or South-East exposure (Table 1). 



572 



SYLVIE BARBALAT 



Species 


Author 


Distribution 


Level 


Habitat 


Main host plants 


Site 


BUPRESTIDAE 














Agrilus angustulus 


ail-, 1803) 


eurosiberian, NA 


col 


broadl/tran 


Que reus 


1,2,3,4,6,8,10,11 


Agrilus biguttatus 


(F., 1777) 


holomediterranean 


col 


broadl/tran 


Quercus 


1,2,3,4,5,10,11,12 


Agrilus cyanescens 


Ratzb.,1837 


european 


col -mon 


broadl/tran 


Lonicera 


10 


Agrilus laticornis 


(111. ,1803) 


european, ME 


col 


broadl/tran 


Quercus 


3,4,7 


Agrilus olivicolor 


Kiesw.,1857 


eurosiberian 


col 


broadl/tran 


Corylus, Carpinus 


2,5 


Agrilus sulcicollis 


Lacord.,1835 


eurosiberian 


col 


broadl/tran 


Quercus 


1,2,3,4,6,7,11 


Agrilus viridis 


a, 1758) 


eurosiberian 


col -mon 


broadl/tran 


broadl 


2,3,4,5,8,11,12 


Antliaxia helvetica 


Stierl.,1868 


mountain 


col -sub 


conif/tran 


conif 


1,2,3,4,5,6,7,8,9,10,11,12 


Antliaxia nitidula 


(L-,1758) 


holomediterranean 


col-mon 


tran/orch 


Prunus 


1,7 


Antliaxia quadripunctata 


(L.,1758) 


mountain 


mon-sub 


conif/tran 


conif 


1,2,3,5,6,7,11.12 


Anthaxia Salicis 


(F., 1777) 


holomediterranean 


col 


broadl/tran 


Quercus 


1,4 


Anthaxia similis 


Saund.,1871 


mountain 


mon-sub 


conif/tran 


conif 


3,6,11 


Chrysobothris affinis 


(F., 1794) 


eurosiberian, NA 


col-mon 


broadl/tran 


broadl 


1,2,3,4,8,9,11 


SCARAB AEIDAE 














Amphimallon atrum 


(Hbst.,1790) 


W. european 


col-mon 


open/ tran 


? 


7,8,9,10 


Cetonia aurata 


(L,1761) 


palearctic 


col -sub 


broadl/tran 


broadl 


3,4,7,8,11 


Hoplia argentea 


(Poda,1761) 


C. andS. european 


col-sub 


open/tran 


? 


4,5,8,9,10 


Omaloplia ruricola 


(F., 1775) 


C. european 


col-mon 


open 


Poaceae, div 


10 


Phyllopertlm hordeola 


(L.,1758) 


eurosiberian 


col-sub 


open/tran 


div 


1,3,4,6,7,8,9,10,12 


Rhizotrogas aestivùs 


(01,1789) 


ponto medi terranean 


col-mon 


open/ tran 


div 


1 


Serica brunnea 


(L.,1758) 


eurosiberian 


col-mon 


broadl/tran 


div 


1,2,4,5,7,8,9,10,11,12 


Trichius fasciatus 


(L.,1758) 


eurosiberian 


col-mon 


broadl/tran 


broadl 


1,2,3,4,5,6,7,8,10,11,12 


LUCANIDAE 














Platycerus caprea 


(Deg.,1774) 


C. european 


mon-sub 


broadl 


broadl 


4,7,8,9,10,12 


Platycerus caraboides 


(L-,1758) 


C. european 


col-mon 


broadl 


broadl 


1,2,4,5,12 


Sinodendron cxlindricum 


(L.,1758) 


eurosiberian 


col-mon 


broadl 


broadl 


7 


CERAMBYCIDAE 














Agapanthia villosoviridescens (Deg.,1775) 


eurosiberian 


col-mon 


open/ tran 


Asteraceae, Apiaceae 1 1 


Agapanthia violacea 


(F.,1775) 


pont omedi terranean 


col-mon 


open 


Dipsacaceae 


1 


Alosterna tabacicolor 


(Geer,1775) 


palearctic 


col-mon 


broadl 


broadl 


1,2,3,4,5,6,7,8,9,10,12 


Anaglyptus mysticus 


(L.,1758) 


european 


col-mon 


broadl/tran 


broadl 


3,5,6,12 


Anastrangalia dubia 


(Scop., 1763) 


mountain 


mon-sub 


conif/tran 


conif 


2,3,4,5,6,7,10,11,12 


Anastrangaiia sanguinolenta 


(L.,1761) 


C. and E. european 


mon-sub 


conif/tran 


conif 


1,3,5,6,7,10,11,12 


Anoplodera sexguttata 


(F.,1775) 


european 


col 


broadl/tran 


Quercus 


1,4.5 


Arhopalus rusticus 


(L.,1758) 


hol arc tic 


col-mon 


conif 


conif 


11 


Ceratnbyx scopolii 


Füssl.,1775 


european, NA 


col-mon 


broadl/orch 


broadl 


10 


Clytus arietis 


(L.,1758) 


european, ME 


col-mon 


broadl/tran 


broadl 


1,2,3,4,5,6,7.8,9,10,11,12 


Clytus lama 


Muls.,1847 


mountain 


mon-sub 


conif/tran 


conif 


2,6,7,11 


Cortodera femorata 


(F., 1787) 


C. andE. european 


col-sub 


conif 


Picea 


1,3,12 


Corymbia rubra 


(L.,1758) 


palearctic 


col-sub 


conif/tran 


conif/broadl 


1,2,3,4,5,6,7,10,11,12 


Dinoptera collaris 


(L.,1758) 


palearctic 


col-mon 


broadl/tran 


broadl 


6,7,8,10,12 


Gaurotes virginea 


(L.,1758) 


boreo-alpine 


mon-sub 


conif/tran 


conif 


3,5,6,12 


Grammoptera abdominalis 


(Steph.,1831) 


C. and S. european, ME col 


broadl/tran 


Quercus, Castanea 


7 


Gratrunoptera ruficornis 


(F., 1781) 


european, ME 


col 


broadl/tran 


broadl 


1,2,12 


Leiopus nebulosus 


(L.,1758) 


european 


col-mon 


broadl/mix 


broadl 


2,3,7,8,11,12 


Leptura nwculata 


(Poda,1761) 


european, ME 


col-mon 


broadl/tran 


broadl 


1,2,3,4,5,6,7,8,9,10,11,12 


Molorchus minor 


(L.,1758) 


palearctic 


mon-sub 


conif/mix 


conif 


7,9 


Obrium brunneum 


(F., 1792) 


european, ME 


mon-sub 


conif/mix 


conif 


3,5,6,7,9,11,12 


Oxymirus cursor 


(L.,1758) 


E. european, ME 


mon-sub 


conif/mix 


conif/broadl 


3,6 


Pachytodes cerambycifonnis 


(Schrk.,1781) 


C. european, ME 


col-mon 


broadl/tran 


broadl/coruf 


1,2,3,4,5,6,7,8,10,11 


Panne na balte us 


(L.,1767) 


W. mediterranean 


col 


broadl/tran 


broadl 


3,5,7 


Phymalodes testaceus 


(L.,1758) 


holarctic 


col-mon 


broadl 


broadl 


3,4,7,9 


Phytoecia cylindrica 


(L:;1758) 


palearctic 


col-mon 


open 


Apiaceae 


7,10 


Pidonia lurida 


(F.,1792) 


mountain 


mon-sub 


conif/mix 


Picea, Fagus 


10 


Plagionotus arcuatus 


(L-,1758) 


european, ME, NA 


col 


broadl/mix 


Quercus 


1,4,11 


Pogonocherus fasciculatus 


(Deg.,1775) 


palearctic 


col-sub 


conif/tran 


conif 


3 


Pogonocherus hispidulus 


(Pill.MitL.1783) 


european 


col-mon 


broadl/mix 


broadl 


5,6,12 


Prionus coriarius 


(L-,1758) 


palearctic 


col 


broadl/mix 


broadl/comf 


3,4 


Pseudovadonia livida 


(F., 1776) 


palearctic 


col 


open/tran 


soil 


3,5 


Pyrrhidium sanguineum 


(L.,1758) 


european, NA 


col 


broadl 


Quercus 


1,3,7,12 


Rhagium bifasciatum 


F.,1775 


european 


col-mon 


conif/mix 


conif/broadl 


5,6,7,12 


Rhagium inquisitor 


(L-,1758) 


holarctic 


col-mon 


conif/mix 


conif/broadl 


2,3,6,11 


RJiagium mordax 


(Deg., 1775) 


eurosiberian 


col-mon 


broadl/mix 


broadl/conif 


4,5,8,10,12 


Stenocorus meridianus 


(L-,1758) 


eurosiberian 


col 


broadl 


broadl 


5,6,8,9 


Stenostola dubia 


(Laich., 1784) 


C. and N. european 


col-mon 


broadl/tran 


broadl 


8,10,12 


Stenurella bifasciaia 


(Müll., 1776) 


palearctic 


col 


mix/tran 


broadl/conif 


1,6,7 


Stenurella melanura 


(L.,1758) 


palearctic 


col-mon 


broadl/mix 


broadl/conif 


1,2,3,4,5,6,7,8,9,10,11,12 


Tetropium castaneum 


(L.,1758) 


palearctic 


mon-sub 


conif 


conif 


3,7 


Distribution: ME = Middle East, NA = North Africa, C = central, E. = 


east, N. = north 


S. = south, W. 


= west; 




Habitat: broad! = broad-leaved forest, conif = coniferous forest, mix = 


mixed forest, tran = clearings and edges, open = open 


habitat, orch = orchard; 


Level: col = colline, mon = montane, sub ^subalpine; Main host plants 


conif = coniferous trees, broad 


= broad-leaved trees 


div = diverse plants. 



Table 2. Species list and ecological overview. Nomenclature after Lohse & Lucht (1992) and 
Bense(1995). 



IMPORTANCE OF FOREST STRUCTURES ON BEETLES 



573 



A. violacea 


St. 1 


St. 4 


St. 11 


St. 3 


St. 2 


St. 5 


St. 6 


St. 12 


St. 7 


St. 8 


St. 10 


St. 9 


total 




10 
























10 




R. aestivus 


12 
























12 


Oak 


A. Salicis 


7 


6 






















13 


A. sexguttata 


2 


10 








1 














13 




P. arcuatus 


7 


4 


2 




















13 


pref ering 


A. sulcicollis 


26 


49 


14 


22 


1 




5 




1 








118 




A. bigutattus 
A. anguslulus 


32 
35 


167 
21 


19 
16 


16 
8 


2 
3 


1 


2 


2 




1 


1 
1 




240 
87 


species 


C. qffinis 
G. ruficornis 


1 


6 


5 


4 


1 
2 






1 




3 




1 


21 
21 




18 










C. aurata 




14 


1 


2 










2 


3 






22 




A. villosoviridescens 






1 




















1 




A. rustic us 






1 




















1 




A. olivicolor 










3 


1 














4 




P. caraboides 


1 


1 






11 


4 




1 










18 




A. viridis 




1 


34 


2 


5 


1 




1 




2 






46 




C.femorata 


1 






1 








1 










3 




P. livida 








1 




1 














2 




S. hif asciata 


15 












1 




21 








37 




P. sanguineum 


1 






3 








1 


2 








7 




A. taticornis 




1 




1 










1 








3 




P. balteus 








1 




1 






1 








3 




0. cursor 








1 






1 












2 




A. nilidula 
P. coriarius 


1 
















6 








7 




1 




3 










4 




P.fasciculatus 








1 


















1 




R. inquisitor 
A. similis 






1 
2 


1 

2 


1 




1 
1 












4 
5 


Artificial 


A. quadripunctata 


3 




5 


9 


2 


1 


12 


30 


4 








66 


clearings 


A. dubia 




3 


12 


23 


5 


13 


10 


17 


4 




1 




88 




A. sanguinolenta 


5 




16 


2 




7 


79 


17 


6 




1 




133 


and 


C. rubra 


5 


52 


108 


48 


56 


38 


75 


25 


1 




3 




411 


coniferous 


P. cerambyciformis 


6 


3 


3 


7 


15 


5 


10 




1 


1 


1 




52 


S. melanura 


33 


222 


53 


180 


25 


673 


481 


184 


42 


9 


32 


21 


1955 


tree 


P. hispidulus 
A. mysticus 








1 




1 
1 


1 
1 


1 
1 










3 

4 


prefering 


G. virginea 
R. bifasciatum 








3 




1 
4 


5 
4 


3 
14 


3 








12 
25 


species 


C. lama 






1 




1 




2 




1 








5 




0. brunneum 






1 


1 




1 


1 


1 


2 






3 


10 




L. nebulosus 
T.fasciatus 






2 


1 


2 






1 


3 






1 


10 




15 


11 


15 


1 


5 


15 


3 


19 


4 


3 


5 




96 




L. maculata 


21 


13 


28 


17 


5 


38 


60 


51 


3 


12 


28 


11 


287 


ubiquitous 


C. arietis 


30 


33 


38 


61 


17 


25 


27 


59 


14 


64 


61 


10 


439 


species 


A. helvetica 


11 


15 


46 


31 


3 


43 


90 


30 


112 


7 


6 


3 


397 




A. tabacicolor 
T. castaneum 


9 


20 




27 


2 


25 


23 


13 


9 


13 


19 


17 


177 










1 










1 








2 




R. mordax 




4 








3 




7 




1 


1 




16 




G. abdominalis 


















2 








2 




P. testaceus 
S. brunnea 


1 


1 
4 


2 


2 


2 


2 




1 


1 






1 


5 




2 


18 


25 


6 


63 




H. argentea 




4 










6 






51 


1 


1 


63 


Fresh 


P. horticola 


1 


2 




1 






1 


4 


13 


105 


10 


2 


139 




S. cylindricum 


















1 








1 


forest 


S. meridianus 












1 


2 






2 




4 


9 




P. capre a 




1 












16 


12 


6 


19 


6 


60 




D. collar is 














1 


1 


3 


1 


11 




17 


and 


P. cylindrica 


















3 




1 




4 




M. minor 


















2 






1 


3 


altitude 


S. dubia 
















1 




2 


4 




7 




A. atrum 

A. cyanescens 


















2 


3 


6 

1 


15 


26 
1 


prefering 


C. scopolii 






















1 




1 




P. lurida 
0. ruricola 






















1 

1 




1 
1 


species 


total 


309 


669 


426 


485 


169 


907 


905 


503 


285 


307 


241 


103 


5309 





Table 3. Number of collected species in each station, diagonalized according to the biotopes 
they have been found in. 



574 sylvie barbalat 

Beetle sampling and analyses 

The study was conducted from the end of April until the beginning of September 
of 1994 and 1995. The following trapping methods were used: window traps and water 
traps (yellow and white) (Barbalat 1995). Two traps of each type were placed in each 
site at about 20 cm above ground level for water traps and 80 cm for window traps. 
They operated without interuption during the whole season. 

The data analysis was made by canonical correspondence analysis (ter Braak 
1986, 1988a). The program CANOCO (ter Braak 1988b) was used in order to 
determine the most relevant environmental variables influencing species distribution in 
the studied sites. With this method, it is possible to extract the variance explained by 
one or more environmental variables introduced a priori in the analysis. These variables 
can be chosen by a forward selection and their significance tested by a permutation test. 
The following environmental variables were introduced in the analysis and submitted to 
a forward selection: "stand type (oak, beech, mixed)", "proportion of broad-leaved/ 
coniferous trees", "clearing size (small, medium, large)", "altitude", "tree covering", 
"deadwood quantity", "ecotone type [natural edge (with bush stratum), clean edge 
(without bush stratum), artificial clearing]", "slope" and "young tree size (< lm, 1-2 m, 
>2m )" in the clearings. 

RESULTS 

A total of 65 species (Table 2) were recorded in our study area: 13 Buprestidae, 
41 Cerambycidae, 8 phytophagous Scarabaeidae and 3 Lucanidae. The total number of 
collected specimens was 5309 (Table 3). 

The forward selection showed that two environmental variables ("stand type" 
and "ecotone type") explain a significant part of the data variation. We tested the 
environmental variables on each season separately as well as on both seasons together. 
In the three cases, the same variables ("stand type" and "ecotone type") were found 
significant (p < 0.01 for both variables in the three cases). Axis 1 explains 16.8 % of 
the variance in 1994, 20.7 % in 1995 and 18.9 % when both years are cumulated. For 
the second axis, these values are 16.3 %, 17.7 % and 17.7 % respectively. For this part 
of the analysis, the rare species (less than 3 specimens) were excluded. 

The effect of the year of capture on our results has also been tested. It represents 
only 3.4 % of the variance (p = 0.74; NS). The beetle communities can therefore be 
considered as stable during our two trapping seasons. Table 3 shows the species 
richness and abundance. It was diagonalized according to the different biotopes. 

Fig. 2 shows the species and sites distribution on the first two canonical axes. 

DISCUSSION 

Biological interpretation 

Several of the tested variables were found collinear and our significant variables 
have to be considered as synthetic. The variable "stand type" is in fact correlated with 
the altitude, which implies not only a climatic and vegetation change but also diffe- 



IMPORTANCE OF FOREST STRUCTURES ON BEETLES 



575 











Axis 2 


St 1 

R. aestivus 
° A. violacea 




( 
-2.5 


2.5 


^ 


v__ 


-l.i 


J 














A. Salicis □ 

Oak stand • 








□ S. bifasciata 

• Natural edge 


p P. arcuatus 
d G. ruticornis 

A. sexguttata □ 








D A. nitidula 


St. 4 


St. 9 . 
Axis 1 


St 
St. 10 . 


8 ■ 
St. 7 


A. ami m 
g M. minor 
° P. cylindrica 

P. hordeola o 

S. brunnea o ° H. argentea 
S. dubia a op caprea p tes 
D. collans ° p sanguinei™ ° 
A. tabacicolor ° 


A. angustulus 
C. aurata „ D o A. btgutattus 
C. femorata° A sulclcolIls 

Ba. laticornis 
aceus 

o C. affinis 

a T. fasciatus 








Altitude -^ " " C. arietis L q 
S. meridianus A. helvetica □ 

•Beech stand SmeU 

A sang 
0. brunneum d 
L. nebulosus a 
P. balteus o 


maculata ^ tnordax 
nura ° ^ ceram by c ' rorm ' s 

o P conarius 
uinolenta ° P. caraboides 
a n C. rubra 
a A quadripuncata 








C. lama o 
R. bifasciarum ° 


a viridis .Artificial clearing 

s * A. dubia . St 1 1 

Mixed forest " St3 








G. 
St. 12. A n 


A. olivicolor 
'irginea a similis ■ St - 
lysticus « r inquisitor 
P. hispidulus 

St*6 st. 5 



Fie 2. Diagram showing species and sites distribution along the first two canonical axes, 
representing a linear combination of the synthetic variables "oak stand" and "artificial clearing". 
The variables "beech stand", "altitude" and "mixed forest" were added a posteriori as passive 
variables for heuristic purposes. 

rences in meadow management. The other significant variable "ecotone type" is also 
related to other variables such as the number of coniferous trees (mainly pine) in a 
stand, implying specialized forest management. 

On the plane formed by axes 1 and 2, it was possible to identify the three 
following groups: sites 7, 8, 9 and 10, sites 2, 3, 5, 11 and 12 and sites 1 and 4. 

On the left end of axis 1, the variance of which is mainly due to stand type, we 
find sites 8, 9 and 10 located at a higher altitude in pure beech forests. Site 7 is located 
in a mixed forest but at a similar altitude. The only species which can be considered as 
a beech forest indicator is the lucanid Platycerus caprea, which usually lives in old 
stumps in mountain mixed beech forests (Koch 1992). 



576 SYLVIE BARBALAT 

The other species found in these four sites seem more related to the edge itself 
than to the forest type. For instance, the larvae of the scarabaeids Serica brunnea and 
Phyllopertha horticola are rhizophagous on many plants and the adults are 
phyllophagous on diverse plants, including trees. Serica brunnea, Hoplia argentea and 
Amphimallon atrum are more common in hilly or mountainous regions (Horion 1958; 
Allenspach 1970). Their abundance in these sites could also be due to a more 
extensive agriculture, using less pesticides than in the lowlands. 

At the other end of the first axis, sites 1 and 4 are located in almost pure oak 
stands, which is clearly indicated by species developing mainly in oaks, such as the 
cerambycids Plagionotus arcuatus and Anoplodera sexguttata as well as the buprestid 
Anthaxia Salicis. 

Sites 2, 3, 5, 6, 11 and 12 are in the middle of the first axis and are either located 
in mixed forests (sites 3, 5, 6 and 11) or in beech stands with some coniferous trees 
(sites 2 and 12). 

Axis 2, the variance of which is principally due to the clearing type, shows an 
opposition between sites 1, 4, 7, 8, 9 and 10 on the top of the diagram and sites 2, 3, 5, 
6, 1 1 and 12 on the bottom. All the "top" sites except site 4 are located in edges while 
all the "bottom" sites are in artificial clearings. On the top of the diagram, beside 
Hoplia argentea, Amphimallon atrum, Serica brunnea and Phyllopertha horticola, we 
can quote as edge indicators the cerambycid Agapanthia violacea and Phytoecia 
cylindriea, the scarabaeid Rhizotrogus aestivus and the buprestid Anthaxia nitidula. 

The five scarabaeids live in open or in half open habitats. They are typical for 
edges without indicating if they are natural or clean. Agapanthia violacea and 
Phytoecia cylindriea cannot really be considered as typical edge species because their 
host plants can be found in other biotopes such as meadows or embankments. 
Nevertheless, in our case, we would tend to consider these two species as natural edge 
indicators. Actually meadows are usually mown quite early in the season, often before 
the beetle emergence. The use of fertilizers leads to the disappearence of certain typical 
oligotrophic and mesotrophic lawn plants which, among others, host Agapanthia 
violacea and Phytoecia cylindriea. In our study area, we should therefore consider that 
these two species can only live where the edge is wide enough to allow the maintenance 
of their host plants. 

The only species, which can be considered as a typical edge species is the 
buprestid Anthaxia nitidula, which lives in treelike Rosaceae and which is often found 
on Crataegus sp., Prunus spinosa or Rosa canina, which are typical edge shrubs. 

On the bottom of the diagram, we find several species associated with artificial 
clearings. Some of them, such as the cerambycids Corymbia rubra, Prionus coriarius, 
Rhagium bifasciatum or Anastrangalia sanguinolenta live in old stumps and are 
favoured by those left in the artificial clearings after cutting. 

Species living in small branches seem also favoured by the branch heaps left in 
artificial clearings after cutting. The little cerambycid Stenurella melanura seems to 
favour particularly these structures. This species is abundant everywhere but parti- 
cularly in artificial clearings. According to Horion (1974), Stenurella melanura lives in 
rotten branches on the ground. To a lesser extent, species living in little branches such 



IMPORTANCE OF FOREST STRUCTURES ON BEETLES 577 

as Pogonocherus hispidulus, Agrilus olivicolor, Anthaxia similis, Leiopus nebulosus 
and Obrium brunneum also seem to be favoured by artificial clearings. 

Relationships between beetle communities and forest management 

Taking a closer look at axis 2, we can notice that the opposition between 
"artificial clearing" and "natural edge" is linked to another opposition: mixed forests 
against pure forests. This can be explained by a management adapted to each stand 
type. Sites 1, 4, 8, 9 and 10 on the top of the diagram are located in pure oak or beech 
forests, while the other sites are in mixed forests with a certain amount of coniferous 
trees. This is shown on the bottom of the diagram, by a higher number of species, living 
mainly in coniferous trees such as the cerambycids Corymbia rubra, Anastrangalia 
dubia, A. sanguinolenta, Gaurotes virginea, Rhagium bifasciatum and R. inquisitor, as 
well as the buprestids Anthaxia similis, A. qudripunctata and A. helvetica. 

In our study area, mixed forests are mainly situated on thin calcareous soils 
which are unfavourable to the growth of most trees. Pine is the only species which 
grows quite well under these conditions. It requires much light for its growth and 
artificial clearings are necessary to favour it. This explains why artificial clearings are 
opened mainly in this type of forests. 

In our study area, pure beech forests are usually located above 800 m, on deeper 
and more fertile soils. As beech is a shade species, beech forest management does not 
require the opening of large artificial clearings. These forests are usually quite dark as 
beech canopy is very thick and their beetle fauna is generally poorer than in mixed or 
oak forests. 

On the contrary, oak forests usually grow below 700 m and young trees need a 
lot of light for their growth. The management of this type of forests implies the opening 
of artificial clearings. Unlike beech, oak hosts a large number of specific insect species, 
much more related to the tree itself than to the forest structure. This explains the 
position of our site 4, an artificial clearing in a pure oak stand. Its fauna is closer to that 
of site 1 , located at the edge of a pure oak forest, than to the other artificial clearings in 
mixed stands. 

In normally managed forests, most of the trees are cut before reaching an age 
where the foliage becomes scarcer. A forest of healthy trees is therefore very dark, a 
feature not favourable for the studied fauna which is thermophilous. To a certain extent, 
we can consider artificial clearings as a substitute to natural clearings caused in 
primeval forests by the fall of old trees, all the more so that in addition to the light and 
the heat they cause, they also provide suitable biotopes for larvae as long as stumps and 
branch heaps are left on the site. These stumps and branches are more attractive in 
sunny places, than in dark places deeper in the woods, since not only the adults but also 
the larvae are thermophilous. This suggests that a careful management respecting local 
conditions can enhance forest beetle diversity. 

Apparently, edge beetle assemblages are rather constituted of species living in 
herbaceous plants. They would be very sensitive to any edge and surrounding mo- 
dification, be it a reduction of the edge width or a change in the agricultural practices. 



578 SYLVIE BARBALAT 

For instance the replacement of a meadow by a field would change the microclimate, 
supress an important food source and probably increase chemical treatments. 

The results therefore suggest that the link existing between the type of forest and 
the kind of management is reflected by the beetle communities. 

CONCLUSION 

This study shows that the main factor for the presence of xylophagous beetles is 
the occurence of their host plants. This concerns chiefly 9 species which are strongly 
dependant on their host plant. In our case, the species in question are mainly linked to 
oak and would be very sensitive to a vegetation change. Therefore, this tree of high 
biological value, hosting many typical species, has to be maintained. It should also be 
favoured by adapted management. 

The second main factor found in our study is the ecotone structure. Species 
found in artificial clearings are not the same as those trapped in natural edges. 

In order to preserve in our forests a diversified beetle fauna including specia- 
lized species, it is important to keep the number of indigenous trees adapted to site 
conditions as high as possible, among them oak. In a mountainous country like Switzer- 
land, oak is not a very common tree since most of the lowlands are intensively 
cultivated. It is therefore important to favour this species where it is possible. At higher 
altitude, beech often constitutes monospecific stands. Tree diversity could be enhanced 
by favouring other species such as linden (Tilia sp.) or maple. Diversified structures 
must also be maintained. Even if artificial clearings have a favourable effect when 
stumps and branch heaps are left after the cutting, one must keep in mind that they 
cannot replace natural edges. These have to be maintained where they already exist and 
encouraged elsewhere in favourable sites. It has also to be recalled that artificial 
clearings favouring pine have a very favourable effect when young trees are small. 
Nevertheless, their area should remain limited because the rich mixed forest should not 
be replaced little by little by a pine monoculture. 

ACKNOWLEDGEMENTS 

I would like to express my deep gratitude to Prof. W. Matthey and Dr D. 
Borcard for guiding this study, to Dr J. Gutowski for reviewing this manuscript, to E. 
Mitchell for correcting the English text and to P. Junod and M. Plachta for colla- 
boration. 

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186 pp. 

Barbalat, S. 1995. Efficacité comparée de quelques méthodes de piégeage sur certains Colé- 
optères saprophages ou xylophages et influence de l'anthophilie sur le résultat des 
captures. Bulletin de la Société neuchâteloise des Sciences Naturelles 1 18: 39-52. 



IMPORTANCE OF FOREST STRUCTURES ON BEETLES 579 



Barbalat. S. 1996: Influence de l'exploitation forestière sur trois familles de Coléoptères liés au 

bois dans les Gorges de l'Areuse (Canton de Neuchâtel, Suisse). Revue suisse de 

Zoologie 103(2): 1-12. 
Barbalat, S. 1997. Faunistique de 47 Cérambycides (Col. Cerambycidae) capturés dans les 

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Naturelles 120:99-119. 
Barbalat, S. & Borcard, D. 1997. Distribution of four beetle families (Coleoptera Buprestidae. 

Cerambycidae, phytophagous Scarabaeidae and Lucanidae) in different forest ecotones in 

the Areuse Gorges (Neuchâtel, Switzerland). Ecologie 28 (3): 199-208. 
Bense. U. 1995. Longhorn beetles. An illustrated key to the Cerambycidae and Vesperidae of 

Europe. Margraf, Weikersheim, 512 pp. 
Dajoz, R. 1980. Ecologie des insectes forestiers. Bordas, Paris, 489 pp. 
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(eds). Rote Liste der gefährdeten Tiere und Pflanzen in der Bundesrepublik Deutschland. 

Naturschutz Aktuell 1 . Kilda. Greven, 270 pp. 
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forest site types in Bialowieza Primeval Forest. Parki Narodowe i Rezerwaty Przyrody 

6(1): 77-94. (in Polish) 
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unter Berücksichtigung der Holzbewohnenden Käfer. Tätigkeitsberichte der Naturfor- 
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Überlingen, 343 pp. 
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Silvestria, Series Silvestris 16: 131-152. (in Polish) 
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Agraria et Silvestria, Series Silvestris 17: 1 17-135. (in Polish) 
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525-533. 



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TER Braak, C. J. F. 1986. Canonical correspondence analysis. A new eigenvector technique for 
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Revue suisse de Zoologie 105 (3): 581-664; septembre 1998 



Fossil and extant species of Cylindromyrmex 
(Hymenoptera: Formicidae) 

Maria L. DE ANDRADE 

Zoologisches Institut der Universität, Rheinsprung 9, CH-405 1 Basel, Switzerland. 

Fossil and extant species of Cylindromyrmex (Hymenoptera: Formi- 
cidae). - The genus Cylindromyrmex is restricted to the Neotropics and 
comprises ten Recent species and two fossil ones from Dominican amber. 
Cylindromyrmex parallelus Santschi is a junior synonym of Cylindro- 
myrmex meinerti Forel. Cylindromyrmex whymperi (Cameron) is re- 
established as a good species. Cylindromyrmex escobari n. sp. is described 
from Colombia. A cladistic analysis allows grouping of the twelve known 
species into four clades: the striatus, the boliviae, the brevitarsus, and the 
longiceps clades. No Cylindromyrmex has been reported from the Recent 
fauna of Hispaniola yet. This genus existed on Hispaniola during the Early 
or Middle Tertiary times and its apparent absence from the extant fauna of 
the island, if confirmed, should be due to a more recent extinction. 

Key-words: Formicidae - neotropical ants - Cylindromyrmex - Dominican 
amber - fossil ants - Tertiary. 



INTRODUCTION 

The subfamily Cerapachyinae contains 3 tribes: Cerapachyini (three genera), 
Acanthostichini (one genus) and Cylindromyrmecini (one genus, Cylindromyrmex, 
revised in this work). Cylindromyrmex nests in cavities of rotten wood, under bark, in 
hollow twigs, and in termite galleries. They are said to be termite predators. Among 
cerapachyines, Wilson (1985) and Baroni Urbani (1995) reported the presence of the 
genus Cylindromyrmex in Dominican amber without further specifications, and de 
Andrade (in press) describes a new species of Acanthostichus from Dominican amber. 
Only two more species of cerapachyines are known in the fossil record, both assigned 
to the extinct genus Procerapachys from Baltic Amber. Their systematic position is not 
clear. In fact, Brown (1975) within Cerapachys, considered the two Baltic species to 
represent a distinctive species group on the basis of their large eyes and complete 
promesonotal suture. The eyes of these two species, as they have been figured by 
Wheeler (1915), appear comparable to those described for several Recent represen- 
tatives, and the promesonotal suture is drawn uninterrupted only for C. annosus and not 
for C. favo s us. The two Baltic species, as far as I know, have never been re-studied 



Manuscript accepted 29.05.1998 



582 MARIA L. DE ANDRADE 

since Wheeler's descriptions at the beginning of this century. Most contemporary 
species, however, show a completely fused mesosoma without traces of suture, but a 
well visible suture is present at least in the S. African C. wroughtoni. 

MATERIAL AND METHODS 

Two fossil specimens of Cylindromyrmex have been examined in two samples 
of amber from the Dominican Republic: 

Do-4 130-1 (Fig. 1) of the amber collection of the State Museum of Natural 
History, Stuttgart (Department of Phylogenetic Research). A light yellow sample 
containing only one winged gyne of Cylindromyrmex. The preservation condition of the 
specimen is good, though whitish layers surround the right side of the frontal carina, the 
mesosoma, the wings, the gaster, and part of the legs. 

MCZC (Fig. 2) of the collection of the Museum of Comparative Zoology, 
Harvard, U.S.A. A dark yellow sample containing a dipteron, few impurities, small air 
bubbles, remaining of insect wings and one winged gyne of Cylindromyrmex. The 
preservation condition of the specimen is good. 

The Recent species of Cylindromyrmex examined in this study are deposited in 
the following collections, given here with the relative coden as it will be used in the 
following text: 

BMNH The Natural History Museum, London, England. Courtesy of Barry Bolton. 

CPCC Centro de Pesquisa do Cacao. CEPLAC, Itabuna, Bahia, Brasil. Courtesy of Dr. 

Jacques H. C. Delabie. 
WEMC William and Emma MacKay, Texas, United States. Courtesy of Prof. William P. 

MacKay. 
DEIC Deustches Entomologisches Institut. Eberswalde, Deutschland. Courtesy of Dr. 

Stephan M. Blank. 
IAVH Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Villa de 

Leiva, Santafé de Bogota. Colombia. Courtesy Fernando Fernândez-C. 
IEGG Istituto Di Entomologia "Guido Grandi", Bologna, Italy. Courtesy of Prof. Egidio 

Mellini. 
LACM Natural History Museum of Los Angeles County, USA. Courtesy of Roy R. Snelling. 
MHNG Muséum d'Histoire Naturelle, Geneva, Switzerland. Courtesy of Dr. Ivan Lobi. 
MIZA Museo del Instituto de Zoologìa Agricola "Francisco Fernandez Yépes", Maracay, 

Venezuela. Courtesy of John Lattke and José L. Garcia 
MNHN Muséum National d'Histoire Naturelle, Paris. Courtesy of Dr. Janine Casevitz- 

Weulersse. 
MCSN Museo Civico di Storia Naturale "Giacomo Doria", Genoa, Italy. Courtesy of Dr. Valter 

Raineri. 
MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, 

USA. Courtesy of Stefan Cover. 
MPEG Museu Paraense Emilio Goeldi, Para, Brazil. Courtesy of Dr. Ana Y. Harada. 
MZSP Museu de Zoologia, Universidade de Sào Paulo, Brazil. Courtesy of Prof. Carlos 

Roberto Ferreira Brandào. 
NHMB Naturhistorisches Museum Basel, Switzerland. Courtesy of Dr. Michel Brancucci. 
NHMW Naturhistorisches Museum Wien, Austria. Courtesy of Dr. Stefan Schödl. 
USNM United States Department of Agriculture, Agricultural Research Service, Systematic 

Entomology Laboratory, c/o National Museum of Natural History, Washington, D. C, 

USA. Courtesy of Dr. Ted R. Schultz. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



583 




Fig. 1 
Dominican amber. Specimen Do-41 30. Profile (top); dorsal view (bottom). 



The measurements and Indices I used are those defined by Brown (1975); these 
and other measurements defined here are: 



TL 



HL 

HW 

EL 
SL 
SW 
WL 

PeL 



(Total Length): combined head length in full face view (mandibles included), Weber's 

length, petiole length (side view), and postpetiole and remaining gastral lengths (both in 

side view). 

(Head Length): with head in full frontal view, maximum measurable distance between 

the middle of the vertexal margin and the middle of the anterior border of the clypeus. 

(Head Width): maximum measurable head width behind the eyes in frontal view. 

(Eye Length): maximum eye length. 

(Scape Length): length of scape shaft, excluding the basal condyle. 

(Scape Width): maximum width of scape. 

(Weber's Length): diagonal length of mesosoma from the anterior prenotai slope to distal 

edge of the posterior border of the propodeum. 

(Petiolar Length): maximum measurable distance, in dorsal view, between the middle of 

the anterior petiolar margin to the middle of its posterior margin. 



584 



MARIA L. DE ANDRADE 




FlG. 2 
Dominican amber. Specimen from MCZC. Profile (top); head in dorsal view (bottom). 

PeW (Petiolar Width): maximum measurable width of the petiole in dorsal view. 

HFeL (Hind Femur Length): maximum measurable distance on the anterior face of the hind 

femur. 
HFeW (Hind Femur Width): maximum measurable width of the anterior face of the hind femur. 
HTiL (Hind Tibia Length): maximum measurable distance on the anterior face of the hind 

tibia. 
HTiW (Hind Tibia Width): maximum measurable distance on the anterior face of the hind tibia. 
HBaL (Hind Basitarsus Length): maximum measurable distance on the anterior face of the hind 

basitarsus excluding the spines. 
HBaW (Hind Basitarsus Width): maximum measurable distance on the anterior face of hind 

basitarsus. 
CI (Cephalic Index): HW/HLx 100 
SI (Scape Index): SW/SL x 100 
HFel (Hind Femur Index): HFeW/HFeL x 100 
HTil (Hind Tibia Index): HTiW/HTiL x 1 00 
HBal (Hind Basitarsal Index): HBaW/HBaL x 100 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 585 

A species-level cladistic analysis was performed using as outgroups two repre- 
sentatives of two closely related genera, Acanthostichus and Simopone. No straight- 
forward autapomorphies have been included in the data matrix. The search for the most 
parsimonious tree(s) was performed by PAUP 3.1.1 (Swofford 1993). The search was 
performed by means of the Branch-and-Bound algorithm (Hendy & Penny 1982). 

In order to assess a statistical degree of confidence to the results obtained in this 
way, 1,000 replicates of a bootstrap analysis as described by Felsenstein (1985) were 
performed by the algorithm equally implemented in PAUP 3.1.1. The graphic tracing of 
synapomorphies of Fig. 35 was obatined by MacClade 3.01 (M Addison & M Addison 
1992). 

All characters were assumed to be unordered and with equal weight. 



DESCRIPTIONS 

Cylindromyrmex Mayr, 1870 

Cylindromyrmex Mayr, 1870: 967. Type species Cylindromyrmex striatus Mayr, by monotypy. 

Holcoponera Cameron, 1891: 92. Type species Holcoponera whymperi Cameron, by monotypy. 
Junior homonym of Holcoponera Mayr, 1887. Synonymy with Cylindromyrmex pro- 
posed by Forel (1892). 

Cylindromyrmex, Wheeler 1924: 106 (subgenus ad Cylindromyrmex). Type species Cylindro- 
myrmex striatus Mayr, by original designation. Synonymy with Cylindromyrmex pro- 
posed by Brown (1975). 

Hypocylindromyrmex Wheeler, 1924: 106 (subgenus ad Cylindromyrmex). Type species Cylin- 
dromyrmex longiceps Roger, by original designation. Synonymy with Cylindromyrmex 
proposed by Brown (1973). 

Metacylindromyrmex Wheeler, 1924: 106 (subgenus ad Cylindromyrmex). Type species 
Cylindromyrmex godmani Forel, by original designation. Synonymy with Cylindro- 
myrmex proposed by Brown (1973). 

Diagnosis and Description of the genus Cylindromyrmex Mayr 

The workers of Cylindromyrmex and Acanthostichus according to Bolton 
(1994) can be separated from those of the other genera of Cerapachyinae {Sphincto- 
myrmex, Simopone and Cerapachys) for the head dorsum lacking a carina between the 
antennal socket and the lateral margin of the head. In the same key Cylindromyrmex is 
separated from Acanthosthichus by means of the following combination of characters: 
antennal scrobes present, mid and hind tibiae each with 2 pectinate spurs, presternite of 
abdominal segment III approximately at midheight of the first gastric segment, side of 
the head without longitudinal groove, and distinct eyes. To these characters I would 
add another one, easier to detect: all known Cylindromyrmex species have the dorsum 
of the head, of the mesosoma and of the petiole longitudinally striate while no des- 
cribed Acanthostichus shows traces of similar heavy striation on the same body parts 
(only a very light striation close to the antennal articulation, on the pleurae and on the 
petiolar sides can be present in a few species). The groove and the size of the eyes are 
likely to have a weak diagnostic value. All workers and gynes of Cylindromyrmex 
possess a longitudinal groove running posteriorly from the mandibular articulation, but 



586 MARIA L. DE ANDRADE 

the groove of Cylindromyrmex is placed more dorsally than in Acanthostichus. The 
Cylindromyrmex groove is somehow difficult to see because all species have a longitu- 
dinally striate head, while no Acanthostichus are striated. For what corncerns the size 
of the eyes, I have examined workers of C. longiceps with 16 ommatidia and the 
worker of A. texanus should have 10 ommatidia only (Mackay 1996). 

A detailed description of the males, gynes and workers of Cylindromyrmex can 
be found in Brown (1975). Here, I will complement only Brown's diagnoses: 

Worker. Monomorphic but variable in size. Head longer than broad, with 
slightly convex, subparallel or parallel sides. Clypeus short. Frontal carinae parallel or 
subparallel diverging posteriorly. Ocelli present or reduced to an impressed pit. 
Compound eyes placed on the middle or slightly behind the mid line of the head and 
with a variable number of ommatidia (16-500). Antennae 12-jointed. Funicular joints 8- 
10 with spine-like seta on the proximal border; last joint with similar spine-like seta 
but almost on its all surface (Fig. 3). Scapes reaching or slightly surpassing the anterior 
border of the eyes. Funiculi thickening from the base to the apex. Mandibles sub- 
triangular, dorsally flat or convex. Masticatory margin of the mandibles with 4-14 
irregular denticles or edentate. Apex of the mandibles with pointed apical tooth. Palpal 
formula 2,2 or 2,3. Mesosoma elongate, cylindric, with parallel sides and weakly 
convex dorsally. Promesonotal and propodeal sutures absent, simply marked by a pit or 
superficially impressed. Promesopleural suture superficially or deeply impressed. 
Meso-metapleural suture superficially impressed. Humeral angles round. Propodeum 
with basal and declivous faces distinct separated or not by a margin. Propodeal spiracle 
round or oval and placed at mid height in lateral view. Petiole subcylindric, as long as 
broad, longer than broad, or shorter than broad. Petiolar sides subparallel and often 
diverging posteriorly. Ventral petiolar process small or large, subtriangular, sub- 
truncate, or subround. Postpetiole (abdominal segment III or gastral segment I) broader 
than petiole, broader than long, and as broad as the first gastric segment (abdominal 
segment IV or gastral segment II). Postpetiolar sternite antero-medially without or with 
a variably marked triangular "lip". Pygidium obliquely or perpendicularly truncate; 
apex of pygidium with or without a notch. Sides of pygidium surrounded by a set of 
many irregularly distributed denticles with in 2-4 larger denticles above the sting, or 
with a row of denticles enlarging apically. Sting developed, curved upwards and with 
flat sides. Legs incrassate or slender. Femora with a concavity of variable deepness to 
receive the tibiae. All tibiae with a large, pectinate spur. Mid and hind tibiae with an 
additional, smaller, pectinate spur close to the large one. Basitarsi of the three pairs of 
legs of variable length and with 3-7 spine-like setae on the outer apical edge. First, 
second and third tarsomeres with similar spines. Fourth (apical) tarsomeres of variable 
length. Pretarsal claws thicker proximally than distally and with a small denticle or an 
angle on the proximal part. Head, mesosoma and petiole covered by longitudinal striae 
of variable thickness. Postpetiole smooth or striate. First, second and third gastric 
tergites smooth and variably reticulate-punctate or longitudinally striated. Remaining 
gastric tergites, sternites and pygidium smooth and/ or reticulate-punctate. Legs smooth 
to superficially punctate; some species with hind or hind and mid coxae longitudinally 
striated. Body with pointed hairs of different size and variably distributed, generally 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 587 

denser on the gaster. Colour dark ferrugineous to black. Legs concolour with or lighter 
than the body. Some species with yellowish tibiae. 

Gyne. Very similar to the worker but differing from it in the following 
characters. Size slightly or much larger than the worker. Ocelli and compound eyes 
larger. Wings as in Fig. 4. Fore wing with well marked veins and pterostigma. Rsf5 
connected with Rl. Mf2 and r-m medially interrupted. Mf4 and Cu Al variably pig- 
mented. Hind wings with R+Sc, M+CuA and A pigmented. Distal veins faintly 
pigmented, CuA and 1A more pigmented than Rs and M. In some species the wings 
have violaceous reflexes. Dorsum of the mesonotum with or without striae on the sides. 
Mesopleurae striate or not on the anterior part. Scutellum smooth or with variably 
impressed longitudinal striae. 

Male. Size variable, generally smaller, but in some species as large as, or larger 
than the gyne. Head shorter than broad, as long as broad, or longer than broad. Vertex 
convex. Frontal carinae developed but never completely hiding the antennal socket. 
Sides of the frontal carinae subparallel, or broad anteriorly and converging posteriorly, 
or strongly broad anteriorly and touching each other posteriorly. Antennae 13-seg- 
mented, varying from 1/3 to 1/2 of the maximum body length. Ocelli large. Compound 
eyes very large, slightly longer than 1/2 of the head length and largely on the anterior 
half of the head sides. Scapes very short. First funicular joint less than or about 1/2 of 
the length of the second one; second and last two apical joints thinner than joints 3-10. 
Mandibles slender, edentate except for a visible apical pointed tooth. Mesosoma robust. 
Pronotum with subparallel or diverging sides. Mesonotum and scutellum gently convex. 
Pair Mayrian furrows impressed or not. Parapsidal furrows variably impressed. Pro- 
podeum with the sides converging posteriorly. Basal face of the propodeum separate 
from the declivous one by a well marked carina. Petiole cylindric, as long as or longer 
than broad. Anterior face of the petiole truncate and separate from the dorsal one by a 
marked carina. Subpetiolar process variable in size, subtriangular or subtruncate. 
Postpetiole broader than the petiole. Postpetiolar sides diverging posteriorly or gently 
convex. First gastric segment broader than the postpetiole. Second gastric segment as 
broad as or slightly narrower the first segment, rarely broader than the first segment. 
Remaining gastric segments narrowing posteriorly. Legs long and slender. Head with 
deep punctures or piligerous foveae sometimes separated by irregular or regular striae. 
Mesosoma pro- and mesopleurae smooth and with punctures or piligerous foveae of 
variable size. Propodeum and metapleurae with thick, longitudinal rugosities, some- 
times irregular. Petiole and postpetiole smooth or with irregular, longitudinal rugosities, 
very superficial on the postpetiole. Gaster and legs smooth and variably punctate. Body 
with pointed hairs denser than in the female castes. Sometimes the posterior part of the 
head, pronotum, gaster and legs with dense pilosity of variable size. Wings as in Fig. 5, 
similar to the one of the gyne. Colour brown to black. Legs concolour with or lighter 
than the body. Some species with yellowish tibiae. 

List of the characters 

The characters listed below are considered as of possible phylogenetic signi- 
ficance: 



588 MARIA L. DE ANDRADE 



1. Worker. Eyes small to medium (10-200 ommatidia) (0), or large (more than 400 omma- 
tidia) (1). 

2. Worker. Petiolar dorsum with at most 14 striae (0), or with at least 16 striae (1). Species 
with smooth or foveolate petiole (character 7 state 0) were coded as "?". 

3. Worker and gyne. Frons at most slightly broader than 1/3 of the head width (0), or ca. 1/2 
or more of the head width ( 1 ). 

4. Worker and gyne. Base of the mandibles not angulate laterally (0), or angulate laterally 
(1). 

5. Worker and gyne. Occiput high (Fig. 17) (0), or low (Figs. 8, 31) (1). 

6. Hypostomal bridge narrow (Fig. 9) (0), or broad (Fig. 32) (1 ). 

7. Worker and gyne. Head, mesosoma and petiole smooth or foveolate but never striate 
(only traces of fine striation can be present close to the antennal insertions, on the pleurae 
and on the petiolar sides) (0), or head, mesosoma and petiole clearly striate (1). 

8. Worker and gyne. Dorsum of head, mesosoma and petiole with thick striae (Figs. 6, 8, 

12) (0), or with thin striae separated by large interspaces (Figs. 29, 31) (1), or with thin 
striae very close each other (Figs. 17, 20) (2). Species with smooth or foveolate body 
coded as "?". 

9. Worker and gyne. Ventral process of the petiole different shape but never triangular (0), 
or broad, triangular ( 1 ). 

10. Worker and gyne. Dorsum of the petiole with more than 7 long pointed hairs (0), or with 
at most 3 long, pointed hairs (1). 

1 1. Worker and gyne. All gastric tergites smooth or foveolate but never striate (0), or only 
first gastric tergite striate (1), or first and second gastric tergites striate (2). 

12. Worker and gyne. Dorsal face of hind coxae without a concavity close to the articulation 
with trochanter (0), or with a concavity close to the articulation with the trochanter (1). 

13. Worker and gyne. Mid tibiae with no or with one pectinate spur (0), or with two pectinate 
spurs (1). 

14. Worker and gyne. Fore basitarsi longer than mid basitarsi (0), or fore basitarsi at most as 
long as the mid basitarsi ( 1 ). 

15. Worker and gyne. Fore basitarsi shorter than hind basitarsi (0), or fore basitarsi as long as 
or longer than hind basitarsi ( 1 ). 

16. Worker and gyne. Mid basitarsi longer than 1/2 of the hind basitarsi (0), or mid basitarsi 
shorter than 1/2 of the hind basitarsi ( 1 ). 

17. Worker and gyne. Outer apical edge of the hind basitarsi with 0, or 3, or 5 spine-like 
setae (0), or hind basitarsi with 6-7 spine-like setae (1), or hind basitarsi with 4 spine-like 
setae (2). 

1 8. Worker and gyne. Apical tarsomeres of hind legs shorter than the sum of second and third 
tarsomeres (0), or apical tarsomeres of hind legs as long as or longer than the sum of 
second and third tarsomeres (1). 

19. Gyne. Compound eyes largely behind the mid line of the head (0), or on the mid line of 
the head (1). 

20. Gyne. Scutellum smooth, foveolate, or with very thin striae (0), or scutellum with very 
thick striae (1). 

21. Gyne. Hind femora Index < 48 (0), or > 50 ( 1). 

22. Male. Frontal carinae subparallel (0), broad anteriorly and narrower posteriorly (1), or 
strongly broad anteriorly and touching each other posteriorly (2). 

23. Male. Antero-median border of the clypeus convex (0), or straight (1). 

24. Male. Anterior face of femora densely covered by hairs (0), or with only few hairs (1). 

25. Male. Hypopygium smooth or finely denticulate between the distal apodemes (Figs. 7, 11, 

13) (0), or with a simple, umpair, median projection between the apodemes (Figs. 27, 34) 
(1), or with a bidentate median projection between the apodemes (Figs. 16, 23, 24) (2). 

26. Male. Hypopygium not strongly constricted distally (Figs. 23, 27, 34) (0), or strongly 
constricted distally (Figs. 7, 11, 13) (1). 

27. Male. Ventral and dorsal borders of the aedeagus straight or partially concave (Figs. 7, 
16, 27. 34) (0). or convex on their entire length (Figs. 11, 13) (1). 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



589 



Data matrix 





1 


2 | 3 


4 


5 


6 


7|8 


9 


10 


11 1 12 


13| 14| 15 


16|17|18|19 


20 


21 


22 


23|24| 25 


26 1 27 


C. boliviae 


? 


1 1 





1 





1 








1 


1 











1 


2 





C. brasiliensis 


1 


1 1 


1 


1 





1 





1 





1 1 


10 


1 











1 


C. brevitarsus 





1 1 











1 2 


0&1 





1&2 1 


1 


12 10 








1 


2 





C. darlingtoni 





1 1 











1 2 


1 





1 1 


1 


12 10 








9 


7 7 9 


7 7 


C. escobari 





1 1 











1 2 


1 





1 


1 


1 2 1 ? 


? 


? 


9 


7 7 7 


? 7 


C. godmani 


? 


1 1 





1 





1 0&1 








2 1 


1 


10 10 








2 


1 





C. longiceps 





1 





1 


1 


1 1 








2 1 


1 1 


11 





1 


9 


? ? ? 


7 7 


C. meinerti 





1 





1 


1 


1 1 








2 1 


1 1 


10 11 





1 


2 


1 





C. striatus 


1 


1 


1 


1 





1 





1 





1 1 


10 


1 








1 1 


1 1 


C. whymperi 


1 


1 


1 


1 





1 





1 





1 1 


10 


1 








1 1 


1 1 


t C. antillanus 


7 


1 





1 





1 1 








2 1 


1 1 


10 11 








9 


7 7 7 


? ? 


t C. electrinus 


7 


1 1 











1 2 


1 





1 1 


1 


2 








9 


7 ? ? 


7 7 


A. texanus 





? 











? 














1 

















S. annettae 


? 


? 











? 














1 








9 


? 7 ? 


7 ? 



Tab. 1 

Matrix with the presence (1) or absence (0) of the 27 characters described in text among the 
known species of Cylindromyrmex and two outgroups. 

Results of the cladistic analysis 

The data of Table 1 allows the construction of only one most parsimonious tree 
of length 41 (Fig. 35). The tree has a Consistency Index of 0.854 (Rescaled CI = 0.782), 
a Retention Index of 0.915, and a Homoplasy Index of 0.220. The Cylindromyrmex 
species appear grouped in 4 clades with the two fossils, antillanus and electrinus, in 
different clades. A bootstrap test (Fig. 36) of significance as described in the methods 
chapter reveals that only the striatus clade (brasiliensis, (striatus, whymperi)), its 
subclade (striatus, whymperi), part of the longiceps clade (antillanus, (longiceps, 
meinerti)), and its subclade (longiceps, meinerti) are represented at frequencies higher 
than the conventional statistical limits in 1,000 replicates. 

The Clades of Cylindromyrmex and their species 

The striatus clade 

This clade includes three species: brasiliensis, whymperi and striatus. They are 
characterized by the following synapomorphies: (1) eyes large, (2) base of the man- 
dibles of the worker and of the gyne laterally angulate, (3) dorsum of the postpetiole of 
the worker and of the gyne with three long, pointed hairs at most, (4) fore basitarsi as 
long as the mid basitarsi, (5) outer apical edge of the mid and hind basitarsi of the 
worker and of the gyne with 6 or 7 spine-like setae, (6) scutellum of the gyne with very 
thick striae, (7) male hypopygium strongly constricted distally. 



Cylindromyrmex brasiliensis Emery 



Figs 3, 6-7 



Cylindromyrmex striatus Mayr, Mayr 1887: 545. Worker and male (Santa Catarina), nee gyne 
from Lima = whymperi (nee Mayr 1 870). Misidentification. 



590 



MARIA L. DE ANDRADE 



Cylindromyrmex brasiliensis Emery, 1901: 53. Worker and male (Santa Catarina). Originai 
description. Type locality: Brazil. Type material: 3 syntype workers labelled: "S. 
Catharina, Schmalz, typus", in MCSN; 1 syntype worker labelled: "Brésil, Mayr, typus, 
Cylindromyrmex brasiliensis Em (striatus Mayr 1887)". in MCSN, examined. 

Cylindromyrmex brasiliensis Emery , Borgmeier 1937: 218. Gyne. 

Cylindromyrmex brasiliensis Emery, Jaffé 1993: fig. 51. Worker. 

Cylindromyrmex brasiliensis Emery, Fowler & Delabie 1995: 328. 

Diagnosis. The basalmost species of the striatus clade and differing from both 
other species, striatus and whymperi, by the legs dark orange or light brown instead of 
black with at least part of the tibiae yellowish. 

Worker (Fig. 6). Head about 1/5 longer than broad, with subparallel sides. 
Occiput low. Vertexal angles round. Frontal carinae about half broad as the maximum 
head width. Anterior third of the frontal carinae diverging backwards and reaching the 
middle of the eyes posteriorly. Dorsum of the frontal carinae with an impressed, short, 
median sulcus anteriorly. Frontal carinae not reaching the anterior border of the 
clypeus. Compound eyes large, slightly convex and behind the mid line of the head. 
Ocelli developed. Scapes surpassing the anterior border of the eyes. Proximal fifth of 
the scapes about 1/2 narrower than the remaining parts. Mandibles weakly convex 
dorsally. Mandibles laterally angulate at the base. Masticatory margin of the mandibles 
with a set of 5-6 irregular denticles followed by an apical tooth. 




00007983 



10 Lim 



REM-Labor 
Uni Basel 



Fig. 3 

C. brasiliensis Emery. Worker from Capào Bonito, Sào Paulo, Brazil. Apical funicular joint with 
spine-like setae. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



591 




Fig. 4 
C. boliviae Wheeler. Gyne from Rancho Grande, Aragua, Venezuela. Fore and hind wings. 




Fig. 5 
C. godmani Forel. Male from Turrialba, Costa Rica. Fore and hind wings. 

Mesosoma slightly convex dorsally and as long as or slightly longer than the 
head (mandibles included). Pronotum with parallel sides. Mesonotum narrower than 
pronotum. Propodeal sides gently convex. Basal face of the propodeum separate from 
the declivous one by a marked margin superficially interruped medially. 

Petiole sub-cylindrical, slightly longer than broad, anteriorly truncate and dor- 
sally convex. Ventral process of the petiole small and triangular. Postpetiole slightly 
broader than long. Postpetiolar sides diverging backwards. Postpetiolar sternite antero- 
medially with a salient triangular "lip" pointed backwards. Postpetiole in dorsal view 
antero-laterally angulate. Pygidium in side view obliquely truncate. Pygidium in dorsal 
view with the sides bearing irregularly distributed small denticles followed by a row of 
larger denticles converging towards a pair of small teeth over the sting. 



592 



MARIA L. DE ANDRADE 




00007975 



500 |jm 



REM-Labor 
Uni Basel 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



593 




Q.5mm 



Fig. 7 

C. brasiliensis Emery. Male from Santa Catarina, Brazil. Genital appendages: a) lateral view of 
left parameres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 



Fig. 6. C. brasiliensis Emery. Worker from Capào Bonito, Sâo Paulo, Brazil. Head in full dorsal 
view (top), body in full dorsal view (middle), body in profile (bottom). 



594 MARIA L. DE ANDRADE 

Legs. Femora and tibiae not inflated. Hind basitarsi long and about 1/5 shorter 
than the maximum length of the hind tibiae. Outer apical edge of the hind and of the 
mid basitarsi with 6 or 7 spine-like setae. 

Sculpture. Posterior third of the head dorsum and frontal carinae with longitu- 
dinal striae, thinner on the anterior half of the frontal carinae; some striae bifurcated. 
Posterior third of the head dorsum with additional, small, sparse, piligerous foveae. 
Anterior half of the head dorsum in front of the eyes with striae converging towards the 
scrobes, these striae thinner than those on the anterior part of the frontal carinae. 
Ventral part of the head smooth, with sparse piligerous foveae and longitudinal, slightly 
irregular striae on the antero-lateral half only. Mesosoma longitudinally striate and with 
sparse, superficial, piligerous foveae. Dorsum of the pronotum with 17-21 striae similar 
to those on the posterior part of the head dorsum. Prenotai striae prolonging onto the 
mesonotum and to the propodeum but thinner. Pleurae with thin, longitudinal striae and 
rare piligerous foveae, the striae absent on the lower propleurae. Lower mesopleurae 
smooth in small specimens. Piligerous foveae on the meso- and metapleurae denser in 
large specimens. Petiolar dorsum with 15-17 striae similar to those on the pronotum and 
with piligerous foveae. Petiolar sides minutely and superficially reticulate; their dorsal 
half with piligerous foveae, in some specimens the foveae separated by few, thin, longi- 
tudinal striae. Declivous face of the propodeum and anterior face of the petiole minu- 
tely and superficially reticulate. Dorsum of the postpetiole with ca. 33-35 striae as those 
on the mesonotum and propodeum and with few piligerous foveae, more impressed on 
the anterior third; postpetiolar sternite smooth and with sparse piligerous foveae. First, 
second, third and fourth gastric tergites and first gastric sternite smooth and with sparse 
punctuations. Remaining gastric segments superficially reticulate-punctate. Legs with 
very superficial, minute punctures. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) one long, erect to suberect on the external border of the scape, a pair 
between the frontal carinae and clypeus, sparse on the external border of the mandibles, 
one close to each prenotai angle, two or three on the postpetiole, sparse on the gaster, 
and denser on the pygidium; (2) shorter than type (1), variably distributed on the whole 
body; (3) shorter than type (2), suberect or subdecumbent on the head dorsum, 
subdecumbent on the mesosoma, on the petiole and on the postpetiole, decumbet on the 
ventral part of the head, on the gaster and on the legs. In addition, the hypostomal 
bridge surrounded by a row of hairs similar to those of type (1) but appressed and 
apically curved. Outer ventral border of the mandibles with hairs similar to those of the 
hypostomal bridge but shorter. 

Colour black and shining. Mandibles castaneous red. Antennae and tarsi ferru- 
gineous-brown. Legs dark orange or light brown. 

Measurements (in mm) and indices: TL 5.56-8.48; HL 1.16-1.64; HW 0.93-1.32; EL 
0.37-0.49; SL 0.51-0.73; SW 0.17-0.22; WL 1.40-2.20; PeL 0.55-0.96; PeW 0.51-0.84; HFeL 
0.67-1.02; HFeW 0.25-0.34; HTiL 0.65-1.04; HTiW 0.19-0.26; HBaL 0.53-0.83; HBaW 0.09- 
0.12; CI 78.1-80.5; SI 30.7-34.4; HFel 33.3-37.3; HTil 25.0-29.2; HBal 14.4-15.1. 

Gyne. Similar to the worker but differing from it in the following details: head 
with parallel sides; ocelli larger; mesosoma broader; parapsidal furrows impressed; 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 595 

anterior corners of the postpetiole more angulate; striae on the head dorsum less regu- 
lar; piligerous foveae on the vertexal angles and on the ventral part of the head denser 
and deeper; pronotum with longitudinal striae as in the worker; one specimen has the 
pronotai striae irregular and separated by piligerous foveae; mesonotum medially with 
irregular, short striae and few piligerous foveae; sides of mesonotum smooth; scutellum 
with sculpture similar to the one on the mesonotum; pro- and mesopleurae almost 
completely smooth; petiolar dorsum with 10 longitudinal, irregular striae. 

Measurements (in mm) and indices: TL 8.56; HL 1.44; HW 1.14; EL 0.45; SL 0.60; 
SW 0.19; WL 2.40; PeL 0.81; PeW 0.77; HFeL 0.77; HFeW 0.30; HTiL 0.75; HTiW 0.22; 
HBaL 0.61; HBaW 0.10; CI 79.2; SI 31.7; HFel 39.0; HTil 29.3; HBal 16.4. 

Male. Head as broad as long. Vertexal margin subtruncate. Ocelli protuberant. 
Compound eyes broadly convex and largely on the anterior half of the head. Borders of 
the frontal carinae raised and diverging backwards. Frons anteriorly superficially 
concave, medially convex and posteriorly sloping towards the impair ocellus. Clypeus 
declivous; its anterior border gently convex medially. Mandibles long, with edentate 
masticatory margins and a pointed apical tooth. Scapes about half longer than broad. 
Second and last two funicular joints thinner than joints 3-10. 

Mesosoma robust. Pronotum in dorsal view with subparallel sides. Mesonotum 
slightly convex. Parapsidal furrows impressed. Scutellum slightly higher than the 
mesonotum. Basal face of the propodeum separated from the declivous one by a 
marked carina. 

Petiole cylindric, its anterior face truncate and separated from the dorsal one by 
a marked carina. Ventral process of the petiole subtriangular. Postpetiole antero- 
laterally angulate, broadening backwards and much narrower than the first gastric 
tergite. 

Genitalia as in Fig. 7. 

Legs. Femora not inflated. Mid and hind metatarsi long. 

Wings as in Fig. 5. 

Sculpture. Head dorsum minutely punctate, with transversal striae around the 
ocelli and on the antennal scrobes, and with large foveae on the vertexal angles and on 
the ventral part of the head. Dorsum of the pronotum punctate and densely covered by 
foveae slightly larger than those on the head. Mesonotum smooth and with very sparse, 
small foveae. Scutellum with foveae larger than those on the pronotum. Basal face of 
the propodeum and petiole covered by slightly irregular foveae of different sizes, sepa- 
rated by longitudinal striae. Declivous face of the propodeum with longitudinal striae. 
Pro- and mesopleurae smooth and with short striae close to the posterior borders. 
Metapleurae with irregular, longitudinal striae. Postpetiole, first gastric segment and 
legs smooth and with sparse, superficial punctures. Remaining gastric segments punc- 
tate. 

Pilosity. Body covered by pointed hairs of three types: (1) long and suberect, 
dense on the head, mesosoma, sparse on the gaster and on the legs; (2) shorter than 
type (1) variably distributed on the body, dense on the gaster; (3) shorter than type (2), 
decumbent, sparse on the vertexal angles, dense on the legs. 

Colour. Black . Mandibles brown. Antennae and legs yellowish-orange. 



596 MARIA L. DE ANDRADE 



Measurements (in mm) and indices: TL 8.78; HL 1.16; HW 1.16; EL 0.59; SL 0.32; 
SW 0.17; WL 2.76; PeL 0.81; PeW 0.72; HFeL 1.00; HFeW 0.23; HTiL 0.88; HTiW 0.18; 
HBaL 0.77; HBaW 0.07; CI 100.0; SI 53.1; HFel 23.0; HTil 20.4; HBaI 10.4. 

Material examined. BRAZIL: no further locality, 1 worker (syntype), Mayr [MCSN]; 18 
workers, G. Mayr [MCZC, MHNG, MNHN, NHMB, NHMW, USNM]. Pernambuco: Caruaru, 
IV. 1972, 1 worker, M. Alvarenga [MZSP]. Bahia: Encruzilhada, 980 m, XI. 1974, 2 workers, 
Seabra & Alvarenga [MZSP]; Buerarema, 22.IX.1996, 1 worker, R. Blatrix [CPCC]. Rio DE 
Janeiro: Floresta da Tijuca, 11.1957, 14 workers, C. A. Campos Seabra [MZSP]; Represa do Rio 
Grande, 08.11.1961, 6 workers [MZSP]. SÄO Paulo: Piracicaba, Escola Superior de Agricultura 
"Luiz de Queiróz", 25.X.1974, 10 workers, 1 gyne, E. Berti Filho [MZSP, MPEG]; Agudos, 
VIII. 1958, 5 workers, R. Mueller [MZSP]; Barueri, 03VI.1967, 88 workers, K. Lenko [MZSP]; 
Botucatu, 13.X.1987, 1 worker, L. C. Forti & I. M. P. Rinaldi [MZSP]; Sào Paulo, Butantan 23- 
31.VII.1969, 13.VIII.1969, 31VII.1973, 5 workers, L. Travassos Filho [MZSP]; Ilha de Sâo 
Sebastiào, VII. 1987, 500 m, 13 workers, C. R. F. Brandào [MZSP]; Salesópolis, Estaçào 
Biològica de Boracéia, 3-5 V. 1996, 1 worker, C. R. F. Brandào et al. [MZSP]; Capào Bonito, 
14.XI.1990, 1 worker, M. L. de Andrade [LACM]. Parana: Londrina, VII,1987, 1 worker, 
M. E. M. Tomotake [MZSP]. Santa Catarina: no further locality, 3 workers (syntypes), 
Schmalz [MCSN]; no further locality, 8 workers, 1 male, G. Mayr [NHMW]; no further locality, 
1 worker [IEGG]; Gaspar, 123 workers, 1 gyne, M. Silva Fontes [MZSP]; Blumenau, 2 workers, 
Hetschko [NHMW]; same locality, 1 worker [NHMW]. Rio Grande do Sul: Pareci Novo, 
10.V.1927, 3 workers, Hansen [MCZC, MZSP]; same locality, 18.III.1926, 4 workers, P. Rambo 
SJ [IEGG, MZSP]. PARAGUAY: La Cordillera: San Bernardino, 1 gyne, Fiebrig [MHNG]. 

Discussion. The workers and the gynes of brasiliensis possess on the mesosoma, 
on the petiole and on the postpetiole, striae thinner and less regular than striatus and 
whymperi. These three species are very similar each other but the characters already 
given in the diagnosis should be sufficient to clearly allow separation of brasiliensis 
from the other two. 

Mayr (1887) reported specimens of "striatus" collected by Hetschko in Brazil 
in termite galleries. Borgmeier (1937) cited specimens of brasiliensis collected by 
P. Rambo from Pareci Novo (Brazil) in a branch of Erythroxylum obovatum (Erythro- 
xylaceae). 

Distribution. Brazil and Paraguay. 



Cylindromyrmex whymperi (Cameron) sp. rev. Figs 8-11 

Cylindromyrmex striatus Mayr, Mayr 1887: 546 (gyne, Lima), nee worker and male ^brasi- 
liensis). Nee Mayr 1870. Misidentification. 

Holcoponera whymperi Cameron, 1891: 92, fig. Worker. Original description. Type locality: 
Ecuador. Type material: holotype worker labelled: "Whymp. Supp. App. p. 92, Holco- 
ponera wympheri Cam. type", in BMNH, examined. First combination in Cylindro- 
myrmex by Forel 1892: 256. 

Cylindromyrmex striatus Mayr, Emery 1901: 53. Misidentification. 

Cylindromyrmex whymperi (Cameron), Wheeler 1910: 228, fig. 127 (worker). 

Cylindromyrmex striatus Mayr, Wheeler 1919: 266. Misidentification. 

Cylindromyrmex williamsi Wheeler, 1924a: 101, fig. 19. Worker and gyne. Original description. 
Type locality: Seymour Island. Type material: 1 worker labelled: "S. Seymour I., 
Galapagos, W. M. Wheeler", in MCZC, examined. Syn. nov. 

Cylindromyrmex striatus var. tibialis Stitz, 1932: 367. Worker. Original description. Type 
locality: Galapagos Islands. Type material: not available for the present study. Not 
synonym of striatus as proposed by Brown 1975: 82. Syn. nov. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 597 



Cylindromyrmex schmidti Menozzi, 1931: 191, fig. 3. Partim. Worker. Nee gyne (= meinerti). 

Original description. Type locality: La Caja, Costa Rica. Type material: 2 workers 

(syntypes) labelled: "La Caja: 8 kil. w. San José C. R., Heinr. Schmidt. TYPUS, 

Cylindromyrmex schmidti Typus, Menoz.", in IEGG, examined. Synonymia nova. 
Cylindromyrmex striatus Mayr, Brown 1975: different pages, Fig. 94. Partim (only material from 

Peru, Ecuador, Galapagos = whymperi). Misidentification. 
Cylindromyrmex striatus Mayr, Snelling & Hunt 1975. Partim (only material from Peru, 

Ecuador, Galapagos, Chile = whymperi), Figs 19-22 (= gyne, male and worker of 

whymperi). Misidentification. 
Cylindromyrmex striatus Mayr, Fowler & Delabie 1995. Partim (only material from Peru, 

Ecuador, Galapagos, Chile = whymperi). Misidentification. 
Cylindromyrmex whymperi (Cameron), Holldobler & Wilson 1990: 85, n. n. Fig. (worker). 
Cylindromyrmex, Bolton 1994: Figs 9 & 10, worker. 

Diagnosis. A Cylindromyrmex species belonging to the striatus clade, resulting 
as sister species of striatus, but differing from it in the worker and gyne by the thicker 
body striation, and by the posterior third of the head dorsum with 25 longitudinal striae 
at most instead of more than 34. 

Worker (Fig. 8). Head about 1/6 longer than broad, with slightly convex sides. 
Occiput low. Vertexal angles round. Frontal carinae about half broad as the maximum 
head width. Sides of the frontal carinae diverging posteriorly or gently convex 
medially. Dorsum of the frontal carinae with an impressed, short, median sulcus ante- 
riorly. Frontal carinae not reaching the anterior border of the clypeus. Compound eyes 
large, convex and slightly behind the mid line of the head. Ocelli developed. Scapes 
surpassing the anterior border of the eyes. Proximal fifth of the scape 1/2 narrower than 
the remaining parts. Mandibles flat dorsally and shorter than in brasiliensis. Mandibles 
laterally angulate at the base. Masticatory margin of the mandibles with a set of 4-5 
irregular denticles followed by an apical tooth. Hypostomal bridge narrow, with the 
antero-lateral margin concave (Fig. 9). 

Mesosoma gently convex dorsally and as long as or slightly longer than the head 
(mandibles included). Pronotum with parallel sides. Mesonotum narrower than pro- 
notum. Propodeal sides converging posteriorly. Basal face of the propodeum separated 
from the declivous one by a marked margin converging medially. 

Petiole sub-quadrate, with the sides gently diverging backwards. Anterior face 
of the petiole truncate and the dorsal one convex. Ventral process of the petiole trian- 
gular and slightly smaller than in brasiliensis. Postpetiole broader than long and with 
convex sides. Postpetiolar sternite antero-medially with a variably marked triangular 
"lip". Postpetiole in dorsal view antero-laterally angulate. Pygidium in side view obli- 
quely truncate. Pygidium in dorsal view with the sides bearing a row of denticles 
strongly converging to a pair of small teeth over the sting. 

Legs. Femora and tibiae not inflated. Hind metatarsi long and about 1/5 shorter 
than the maximum length of the hind tibiae. Outer apical edge of the hind and of the 
mid basitarsi with 6 or 7 spine-like setae. 

Sculpture. Posterior third of the head dorsum and frontal carinae with thick, 
longitudinal striae, thinner on the anterior half of the frontal carinae. Rare, small, 
piligerous foveae can be present behind the ocelli. Anterior half of the head dorsum 



598 



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Fig. 9 

C. whymperi (Cameron). Worker from Antofagasta, Chile. Anterior portion of the cephalic 
capsule and mandibles in ventral view to show the narrow hypostomal bridge (character 6 state 
0). Notice the concavity of the anterior margin of the hypostomal bridge, a character not verified 
in all species of the genus. 

with striae converging towards the scrobes, these striae thinner than those on the 
anterior half of the frontal carinae. Ventral part of the head with longitudinal striae 
laterally, smooth and superficially punctate medially. Mesosoma with 11-15 longitu- 
dinal striae similar or slightly thicker than those on the posterior third of the head 
dorsum. Lower pro- and metapleurae, and mesopleurae with thin longitudinal striae 
similar to those on the anterolateral part of the head dorsum. Upper pro- and meta- 
pleurae with striae as on the anterior part of the frontal carinae. Petiolar dorsum with 9- 
14 striae similar to those on the mesosoma. Petiolar sides minutely reticulate and with 
less regular and thinner striae than those on its dorsum. Declivous face of the propo- 
deum and anterior face of the petiole minutely and superficially reticulate. Dorsum of 
the postpetiole with ca. 19-25 striae as thick as or slightly thinner than those on the 
mesosoma. Postpetiolar sternite smooth or reticulate and with sparse piligerous foveae. 



Fig. 8. C. whymperi (Cameron). Worker from Ecuador. Head in full dorsal view (top), body in 
full dorsal view (middle), body in profile (bottom). 



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Fig. 1 1 

C. whymperi (Cameron). Male from Lima, Peru. Genital appendages: a) lateral view of left 
parameres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 



Fig. 10. C. whymperi (Cameron). Male from Lima, Peru. Head in full dorsal view (top), body in 
full dorsal view (middle), body in profile (bottom). 



602 MARIA L. DE ANDRADE 

First, second and third gastric tergites and first gastric sternite smooth and with sparse 
punctuations. Remaining gastric segments reticulate-punctate. Hind coxae with few, 
faint, longitudinal striae. Legs with very superficial, minute punctures. 

Pilosity similar to brasiliensis. 

Colour black and shining. Tibiae of three pairs of legs yellowish with the distal 
borders brown. Tarsomeres brown. 

Measurements (in mm) and indices: TL 5.02-7.40; HL 1.08-1.56; HW 0.90-1.28; EL 
0.31-0.45; SL 0.44-0.68; SW 0.15-0.19; WL1.12-1.88; PeL 0.44-0.76; PeW 0.48-0.80; HFeL 
0.61-0.93; HFeW 0.22-0.30; HTiL 0.60-0.97; HTiW 0.16-0.23; HMeL 0.46-0.76; HMeW 0.07- 
0.10; CI 78.9-83.3; SI 27.9-34.1; HFel 32.4-36.1; HTil 23.7-26.2; HBal 13.0-15.2. 

Gyne. Similar to the worker, from which it differs by the following peculiarities: 
vertex less concave medially; mesosoma broad medially; parapsidal furrows impressed; 
petiole slightly longer than broad; pronotum with 14-17 thick, longitudinal striae; 
mesonotum medially with 9-10 longitudinal striae, slightly thinner than those on the 
pronotum, those on the sides less regular and shorter than the median ones; scutellum 
with 7-8 striae as those on the pronotum; dorsum of the propodeum with 14 striae as 
those on the pronotum; petiolar dorsum with 9-10 longitudinal striae as those on the 
pronotum; postpetiolar dorsum with 22-30 longitudinal striae slightly thinner than those 
on the pronotum 

Wings as in fig. 4. 

Measurements (in mm) and indices: TL 7.58-8.16; HL 1.44-1.46; HW 1.16-1.20; EL 
0.45-0.46; SL 0.56-0.60; SW 0.16-0.18; WL2.36-2.40; PeL 0.73-0.76; PeW 0.69-0.72; HFeL 
0.82; HFeW 0.30-0.31; HTiL 0.75-0.78; HTiW 0.23-0.24; HBaL 0.64-0.66; HBaW 0.09-0.1 1; CI 
80.5-82.2; SI 29.2-34.1; HFel 36.6-37.8; HTil 23.9-26.7: HBal 13.3-15.5. 

Male (Fig. 10). Head as broad as long. Vertexal margin subtruncate or convex. 
Ocelli protuberant. Compound eyes broadly convex and largely on the anterior part of 
the head. Borders of the frontal carinae raised and diverging backwards. Frons ante- 
riorly slightly concave, medially convex and posteriorly sloping towards the impair 
ocellus. Clypeus declivous; its anterior border straight. Mandibles long with edentated 
masticatory margins and with a pointed apical tooth. Scapes thick and shorter than first 
and second funicular joints. Funicular joints as in brasiliensis. 

Mesosoma robust. Pronotum in dorsal view with subparallel sides. Mesonotum 
convex. Pair Mayrian carinae superficially impressed. Parapsidal furrows impressed. 
Scutellum slightly higher than the mesonotum. Basal face of the propodeum narrowing 
backwards and separated from the declivous one by a developed and well marked 
carina. Middle of the basal face of the propodeum sometimes with a longitudinal 
sulcus. 

Petiole sub-quadrate. Anterior face of the petiole truncate and separated from the 
dorsal one by a marked carina. Ventral process of the petiole subtriangular. Postpetiole 
broadening backwards and narrower than the first gastric tergite. 

Genitalia as in Fig. 11. 

Legs. Femora not inflated. Mid and hind basitarsi long. 

Wings as in Fig. 5. 

Sculpture. Head dorsum with striae converging from the posterior half of the 
compound eyes to the ocelli. Striae behind the ocelli slightly transversal. Striae between 



FOSSIL AND EXTANT SPECIES OF CYL1NDROMYRMEX 603 

the pair ocelii transversal and converging from the pair to the impair ocellus. Posterior 
half of the frontal carinae with traces of longitudinal striae. Anterior half of the frontal 
carinae with striae converging posteriorly. Head dorsum behind the clypeus with striae 
diverging to the compound eyes. Area close to the insertion of the scape sometimes 
smooth or with few traces of irregular striae. Ventral part of the head minutely punctate 
and with sparse, superficial piligerous foveae and thin striae; the striae slightly longi- 
tudinal on the middle of the head and perpendicular close to the eyes. Dorsum of the 
pronotum with irregular, transversal striae and few irregular foveae. Mesonotum and 
mesopleurae smooth, with sparse piligerous punctures Scutellum with variably impres- 
sed longitudinal striae. Basal face of the propodeum and metapleurae covered by thick 
longitudinal striae. Declivous face of the propodeum smooth. Propleurae with longitu- 
dinal striae as those on the scutellum. Petiole with longitudinal striae and few foveae, 
the striae sometimes very superficial or absent on the dorsum and marked on the sides. 
Postpetiole, first gastric segment and legs smooth and with sparse, superficial punc- 
tures. Remaining gastric segments superficially punctate. 

Pilosity. Body covered by pointed hairs of three types: (1) long, suberect, dense 
on the last gastric segments; (2) shorter and denser than the type (1); (3) shorter than the 
type (2), decumbent, dense on the coxae, on the anterior face of the fore and mid 
femora and on the ventral face of the tibiae. 

Colour. Black and shining. Anterior third of the head dorsum, mandibles, funi- 

culli and tibiae yellowish-orange to light brown, scapes, coxae, femora and tarsi darker. 

Measurements (in mm) and indices: TL 7.58-8.14; HL 1.04-1.12; HW 1.02-1.14; SL 
0.24-0.28; SW 0.15-0.16; WL 2.40-2.72; PeL 0.68-0.80; PeW 0.67; HFeL 0.91-0.98; HFeW 
0.18-0.20; HTiL 0.79-0.87; HTiW 0.17-0.18; HBaL 0.62-0.71; HBaW 0.07-0.08; CI 96.3-101.8; 
SI 57.1-62.5; HFel 19.8-21.3; HTil 20.6-21.5; HBal 11.3. 

Material examined. GUATEMALA: near Tikal, Peten, 24.III.1963, 2 workers, R. M. C. 
Williams [BMNH]. COSTA RICA: La Caja, 8 km W of San José, 2 workers (syntypes of 
schmidti), H. Schmidt [IEGG]; same locality, 1931, 1 gyne (wrongly labelled as syntype of 
schmidti) Schmidt [IEGG]; same locality, 1931, 2 workers, Schmidt [IEGG]; S. José, 2 workers 
(wrongly labelled as holotype and paratype of schmidti) [DEIC]; same locality, in house, sting 
people, 17. V. 1937, 1 worker, F. Quirós [USNM]; same locality, 1940, 1 worker, H. Schmidt 
[MZSP]; Guanacaste, Santa Rosa, 20.X.1996, 1 worker, F. Fernandez-C [IAVH]; Cartago Prov., 
Turrialba, Catie, 25.V.1995, 1 gyne, J. Rifkind [LACM]. GALAPAGOS ISLANDS: S. 
Seymour, 1 worker (syntype of williamsi), W. M. Wheeler [MCZC]; Academy Bay, Indefatigable 
Is. 11-22.1.1906, 1 worker (erroneous labelled as syntype of williamsi), F. X. Williams [MCZC]; 
Fernandina I., 3 km inland from coast on N side, 450 m, 25-27. III. 1970, 19 workers, R. 
Silberglied [MCZC, MZSP, WEMC]; Fernandina I., Punta Espinosa, 13.V.1983, 9 workers, Y. 
D. Lubin [LACM]; Isabela I., Caleta Tagus, 9.V.1983, 15 workers, Y. D. Lubin [LACM]. 
ECUADOR: no further locality, 1 worker (holotype of whymperi) [BMNH]. Hac de Tenguel, 
09.VI.1934, 15 workers, W. von Hagen [LACM, MCZC, USNM]. Los Rios: Babahoyo, 
III. 1938, 1 gyne, H. Hanson & W. H. W. Komp [MCZC]. Guayas: Guayaquil, X.1922, 1 worker, 
F. X. Williams [MCZC]; same locality, 1 gyne, C. T. Brues [MCZC]. PERU: no further locality, 
3 workers, 1 gyne, 2 males, E. A. Martinez [LACM, USNM]; Lismaco, 1 gyne, Radoszkowski 
[MCSN]; Valle Chanchamayo, 800 m, 5 workers, Weyrauch [USNM]. Piura: Sullana, Hda. 
Mallares, 24.VII.1957, 1 worker, 1 gyne, W. Markl [NHMB]. Lambayeque: Chiclayo, 4 
workers, Weyrauch [MCZC]; same locality, Hda. Pâtapo, in wood for construction, 2 workers, 1 
gyne [MZSP]. Lima: Lima, 2 gynes, Radoszkowski [MCSN, NHMW]; same locality, in wood, 4 
males, P. Aguiar [USNM]; same locality, 9.VII.1982, 8 males, J. M. Wilson [LACM]: Ancon, 
15.V.1913, 1 gyne, [LACM]. Cuzco: Cuzco, 1.1995, 2 workers, M. A. B. Smith [CPCC]. 



604 MARIA L. DE ANDRADE 

BOLIVIA: Beni: Trinidad, 1 worker, Lizer & Deletang [NHMB]. CHILE: Tarapaca: Arica, 
18°29' S 70°20' W, 40 m, 24.IX.1966, 1 worker, 1 gyne, M. E. Irwin [LACM]. Antofagasta: 
Antofagasta, 1988, 13 workers, J. Vidal [MZSP]. BRAZIL: Santa Catarina: Blumenau, 1 
male, G. Mayr [NHMW]. 

Discussion. Forel (1892) considered H. whymperi a species disctinct from all 
the other Cylindromyrmex. Few years later Emery (1901) proposed the synonymy of 
Cylindromyrmex whymperi (Cameron) with Cylindromyrmex striatus because the des- 
cription of whymperi fits well Peruvian gynes of what he thought to be "striatus". 
Wheeler (1910), without justifying his point of view, published a figure of a Cylindro- 
myrmex worker under the name whymperi. A few years later, Wheeler (1924) 
described williamsi as a new species from the Galapagos, supposed to be different from 
his "striatus" from Guayaquil (Ecuador) and from the worker of whymperi. The exa- 
mination of the type material of striatus, williamsi and whymperi reveals that whymperi 
and striatus are distinct species and williamsi is a junior synonymy of whymperi. 

Examination of the material labelled as "typus" of schmidti by Menozzi shows 
several contradictory points. The type locality of schmidti is La Caja (Costa Rica) and 
the "type" (worker?) should have been deposited in the Deutsches Entomologiches 
Museum and a "cotype" in his own collection. Two workers labelled "S. José, Costa 
Rica, Holotypus, Paratypus, Cylindromyrmex schmidti, Typus! Menoz., Menozzi 
deter." are preserved in the Deutsches Entomologiches Institut of Eberswalde. These 
workers are identical to whymperi. They are unlikely to be the holotype and paratype of 
schmidti because the locality name does not correspond to the one given by Menozzi 
(1931). Two workers and a gyne of schmidti labelled "La Caja: 8 kil. w. San José C. 
R., Heinr. Schmidt, TYPUS, are preserved in the Menozzi collection (IEGG). These 
workers are similar to whymperi and are likely to be the true syntypes of schmidti. The 
gyne does not fit the description and drawing of the gyne of schmidti by Menozzi 
(1931). Additional material in the IEGG contains two other gynes with labels similar to 
those of the "syntype" workers of schmidti and fit exactly the description of Menozzi 
(1931). These two gynes correspond to meinerti Forel. 

C. whymperi has a much broader distribution than striatus. A male in the 
NHMW labelled "Blumenau (Brazil), striatus" (handwriting of Mayr) is definitively not 
striatus. It is identical to all the other males of whymperi I examined in this study. I 
have some doubts about the autenticity of this locality record which is the only one 
from Brazil for this species. 

The species whymperi and striatus are very similar each other in both worker 
and gyne. Examination of the sculpture shows that the striae of whymperi are much 
thicker than those of striatus, especially on the head dorsum and postpetiole. The head 
of whymperi is shorter and with more convex sides than the one of striatus. There 
seems to be little variation in the thickness of the striae on the mesosoma and on the 
postpetiole of workers of whymperi. The specimens from Hac de Tenguel are those 
with thickest mesosomal and postpetiolar striation. Two workers, one from Bolivia 
(NHMB) and the other from Costa Rica (IAVH) have thinner striation on the post- 
petiole but still definitively thicker than that of striatus. Normally gynes of whymperi 
have thicker and less striae on the postpetiole than the gynes of striatus. Ten out of 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 605 

eleven gynes of whymperi have 22-24 striae on the postpetiole. Only a gyne from Costa 
Rica (LACM) has 30 striae on the postpetiole, approaching in this way the gynes of 
striatus with 30-34. Whymperi exhibits also some colour variation. Rare workers and 
gynes have the distal half of the tibiae dark brown. The subspecies striatus tibialis Stitz 
is based on specimens with a similar type of coloration. 

There are no elements to assert whether whymperi is introduced or indigenous in 
the Galapagos Islands. The most remarkable fact about its distribution is that, judging 
from the collection records, it seems to be common on the islands. Its success there, 
however, can be explained in both ways, i. e. by being native of the islands and by the 
lack of competitors after its introduction (see discussion chapter). 

Wheeler (1919) mentioned "striatus" from a house of Indefatigable Island 
(Galapagos Is.). Wheeler (1924, 1936) reports "williamsi" nesting in dead branches of 
the Celastraceous shrub Maytenus obovata whose dead parts contained flourishing 
colonies of Calotermes pacificus. The specimens collected on the Ferdinanda Is. by 
R. Silberglied were under the bark of Bursera graveolens (Burseraceae). 

Distribution. Guatemala, Costa Rica, Galapagos Island, Ecuador, Peru, Bolivia, 
Chile, and Brazil. 

Cylindromyrmex striatus Mayr Figs 12-13 

Cylindromyrmex striatus Mayr, 1870: 967. Gyne. Nee Mayr 1887 (worker and male = brasi- 
liensis). Original description. Type locality: Surinam. Type material: two gynes labelled: 
"Surinam, Coll. G. Mayr, striatus, G. Mayr, Type"; one gyne labeled: "Surinam, 
M. Haab, Collect. G. Mayr, striatus, G. Mayr, Type"; all in NHMW, examined. 

Cylindromyrmex striatus Mayr, Brown 1975. Partim (only material from Guyanas = striatus). 
Nee Cylindromyrmex williamsi Wheeler, Brown 1975: 82. Incorrect synonymy of 
striatus. Misidentification. 

Cylindromyrmex striatus Mayr, Snelling & Hunt 1975. Partim (only material from Surinam and 
French Guiana = striatus). Nee figs. 19-22 (= whymperi). 

Cylindromyrmex striatus Mayr, Overal & Bandeira 1985. 

Cylindromyrmex striatus Mayr, Fowler & Delabie 1995. Partim (only material from Manaus = 
striatus). 

Diagnosis. A Cylindromyrmex species belonging to the striatus clade, resulting 
as sister species of whymperi in my analysis, but differing from it by the thinner body 
striation, by the posterior third of the head dorsum with more than 34 striae instead of 
25 at most. 

Worker (previously undescribed) (Fig. 12). Head 1/5 longer than broad. Sides of 
the head subparallel. Occiput low. Vertexal angles round. Frontal carinae about half 
broad as the maximum head width. Sides of the frontal carinae anteriorly diverging and 
posteriorly gently convex. Dorsum of the frontal carinae with an impressed, short, 
median sulcus anteriorly. Frontal carinae not reaching the anterior border of the cly- 
peus. Compound eyes large, convex and slightly behind the mid line of the head. Ocelli 
well defined. Scapes reaching the anterior border of the eyes. Proximal fifth of the 
scapes 1/2 narrower than the remaining parts. Mandibles flat and short. Mandibles 
laterally angulate at the base. Masticatory margin of the mandibles with a set of 4 
irregular denticles followed by an apical tooth. 



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Fig. 13 

C. striatus Mayr. Male from French Guyana. Genital appendages: a) lateral view of left para- 
meres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 



Fig. 12. C. striatus Mayr. Worker from Para, Rio Curuâ-Una, Brazil. Head in full dorsal view 
(top), body in full dorsal view (middle), body in profile (bottom). 



608 MARIA L. DE ANDRADE 

Mesosoma gently convex dorsally and as long as or slightly longer than the head 
(mandibles included). Pronotum with subparallel sides. Mesonotum narrower than 
pronotum. Propodeal sides slightly convex. Basal face of the propodeum separated 
from the declivous one by a marked margin converging medially. 

Petiole slightly longer than broad, anteriorly truncate and dorsally convex. 
Petiolar sides gently diverging backwards. Ventral process of the petiole small and 
triangular. Postpetiole slightly more than 1/4 broader than long. Postpetiolar sides 
gently convex. Postpetiolar sternite antero-medially with a marked triangular "lip" 
pointing backwards. Postpetiole in dorsal view antero-laterally angulate. Pygidium as in 
whymperi. 

Legs. Femora and tibiae not inflated. Hind basitarsi long, ca. 1.4 shorter than the 
maximum length of the hind tibiae. Outer apical edge of the hind and of the mid 
basitarsi with 6 or 7 spine-like setae. 

Sculpture. Head mesosoma and postpetiole with thinner striation than those in 
whymperi. Posterior third of the head dorsum and frontal carinae with longitudinal 
striae, thinner on the anterior part of the frontal carinae. Sides of the head dorsum in 
front of the compound eyes with thinner striae than on the anterior part of the frontal 
carinae. Ventral part of the head antero-laterally with longitudinal striae; remaining 
parts of the ventral part of the head smooth and with minute, superficial piligerous 
punctures. Mesosoma with longitudinal striae thicker than those on the posterior half of 
the head dorsum. Pronotum with 20-22 striae. Propodeum with ca. 17 striae. Pleurae 
with thin longitudinal striae as those in front of the compound eyes, the striae thicker on 
the upper pro- and metapleurae. Petiolar dorsum with 12-14 striae thicker than those on 
the mesosoma. Petiolar sides minutely reticulate and with less regular and thinner striae 
than those on its dorsum. Declivous face of the propodeum and anterior face of the 
petiole minutely and superficially reticulate. Dorsum of the postpetiole with ca. 29-30 
striae similar to those on the mesosoma. Postpetiolar sternite smooth and with small, 
sparse piligerous foveae. First, second and third gastric tergites and first gastric sternite 
smooth and with sparse punctuations. Remaining gastric segments reticulate-punctate. 
Hind coxae with few, thin, longitudinal striae. Legs with very superficial, minute 
punctures. 

Pilosity as in whymperi and brasiliensis. 

Colour black and shining with lighter legs. Tibiae of three pairs of legs yellow- 
ish with the distal borders brown. 

Measurements (in mm) and indices: TL 6.18-6.56; HL 1.24-1.28; HW 0.99-1.04; EL 
0.35-0.36; SL 0.52-0.56; SW 0.15-0.16; WL 1.52-1.72; PW 0.72-0.75; PeL 0.65-0.75; PeW 0.59- 
0.65; HFeL 0.72-0.78; HFeW 0.25-0.28; HTiL 0.71-0.80; HTiW 0.19-0.21; HBaL 0.52; HBaW 
0.09; CI 79.8-81.2; SI 28.6-28.8; HFel 34.7-35.9; HTil 26.2-26.8; HBal 17.3. 

Gyne. Similar to the worker but differing from it in the following peculiarities: 
mesosoma broad medially; parapsidal furrows impressed; posterior part of the head 
dorsum with slightly thicker striae than on those the anterior part; pronotum with 22-24 
longitudinal striae slightly thicker than those on the posterior part of the head dorsum; 
mesonotum smooth to weakly striated medially, the striae very thin and incomplete; 
scutellum with 8-10 striae similar to those on the pronotum; dorsum of the propodeum 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 609 

with 18-20 striae similar to those on the pronotum; petiolar dorsum with 13-15 
longitudinal striae thicker than those on the pronotum; postpetiolar dorsum with 30-34 
longitudinal striae slightly thinner than those on the pronotum 

Wings as in fig. 4. 

Measurements (in mm) and indices: TL 7.50-7.72; HL 1.30-1.32; HW 1.00; EL 0.40- 
0.42; SL 0.50-0.51; SW 0.16; WL 2.08-2.12; PW 0.76-0.80; PeL 0.67-0.70; PeW 0.63; HFeL 
0.68; HFeW 0.27; HTiL 0.67-0.68; HTiW 0.21-0.22; HBaL 0.53; HBaW 0.09; CI 75.7-76.9; SI 
31.4-32.0; HFel 39.7; HTil 30.9-32.8; HBal 17.0. 

Male (previously undescribed). Very similar to the one of whymperi but dif- 
fering from it in the following details: head (eyes excluded) slightly longer than broad; 
vertexal angles less convex. Scutellum with the sides less converging and with the 
posterior border less truncate. 

Genitalia as in Fig. 13. 

Wings as in Fig. 5. 

Sculpture. Ventral part of the head smooth and with few, small piligerous 
foveae. Scutellum, propleurae, petiole, postpetiole, gaster and legs smooth and with 
minutele, superficial punctures, more impressed on the last gastric segments. Ventral 
border of the propleurae with two-three longitudinal striae. 

Colour. Head and mesosoma dark brown-black and shining. Gaster brown. Man- 
dibles, antennae and legs yellow to light brown. The specimen in question is imature. 

Measurements (in mm) and indices: TL 7.74; HL 1.05; HW 1.00; SL 0.26; SW 0.15; WL 
2.54; PeL 0.73; PeW 0.68; HFeL 0.95; HFeW 0.20; HTiL 0.81; HTiW 0.17; HBaL 0.62; HBaW 
0.08; CI 95.2; SI 57.7; HFel 21.0; HTil 21.0; HBal 12.9. 

Material examined. SURINAM: no further locality, 3 gynes (syntypes), G. Mayr 
[NHMW]. FRENCH GUYANA: no further locality, 1 gyne, 1 male [MNHN]. BRAZIL: 
Amazonas: Manaus, 15.11.1989, 1 worker, H. G. Fowler [CPCC]. Para: Rio Curuâ-Una, 
13.XII.1984, 1 worker, W. L. Overal [MZSP]. 

Discussion. C. striatus is a rarely collected species occuring only in the northern 
part of South America. The similarities between striatus and whymperi and the small 
number of striatus specimens available for study is one of the reasons for which 
whymperi has been considered a junior synonymy of striatus. The sole male of striatus 
I examined is also very similar to whymperi. Few differences are visible in their 
genitalia (Figs. 10 & 13). Striatus and whymperi are allopatric (Fig. 37). 

Overal & Bandeira (1985) found specimens of striatus in nests of Nasuti- 
termes sp. and N. surinamensis. 

Distribution. Surinam, French Guyana and Brazil. 

The boliviae clade 

C. boliviae is an isolate species representing a clade of its own. It differs from 
the species of the striatus clade by the following characters: (1) hind coxae with a 
concavity close to the articulation with trochanter, (2) male with frontal carinae broad 
anteriorly and narrower posteriorly, and (3) male hypopygium with a bidentate median 
projection between the apodemes. It differs from the species of the brevitarsus and 
longiceps clades by the following characters: (1) mid basitarsi longer than 1/2 of the 
hind basitarsi, and (2) apical tarsomeres of hind legs shorter than the sum of second and 
third tarsomeres. 



610 



MARIA L. DE ANDRADE 




00007996 



1 mm 



REM-Labor 
Uni Basel 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



611 










300 Lim 




1 mm 



00007960 



1 mm 



REM-Labor 
Uni Basel 



612 



MARIA L. DE ANDRADE 




0.5mm 



Fio. 16 

C. boliviae Wheeler. Male from Rancho Grande, Aragua, Venezuela. Genital appendages: a) 
lateral view of left parameres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 



Fig. 14. C. boliviae Wheeler. Gyne from Rancho Grande, Aragua, Venezuela. Head in full dorsal 
view (top), body in full dorsal view (middle), body in profile (bottom). 

Fig. 15. C. boliviae Wheeler. Male from Rancho Grande, Aragua, Venezuela. Head in full dorsal 
view (top), body in full dorsal view (middle), body in profile (bottom). 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX ß] 3 



Cylindromyrmex boliviae Wheeler Figs 4, 14-16 

Cylindromyrmex boliviae Wheeler, 1924: 104, fig. 20. Gyne. Original description. Type locality: 
Mapiri, Bolivia. Type material: 1 alate gyne (head missing) labelled: "Mapiri, Bolivia: 
Cotype; Gift of W. M. Wheeler; M.C.Z. Cotype, 20336", in MCZC, examined. 

Cylindromyrmex striatus Mayr, Emery 1901: 54 (male, Peru). Nee Mayr 1870. Misidentification. 

Diagnosis. A Cylindromyrmex species differing from all the others by the post- 
petiole smooth or at most with traces of superficial, short striae on the the posterior 
half. 

Gyne (Fig. 14). Head ca. 1/4 longer than broad, with parallel sides. Occiput 
slightly higher than in the species of the striatus clade. Frontal carinae ca. 1/3 narrower 
than the maximum head width. Sides of the frontal carinae diverging backwards and 
reaching at least the middle of the compound eyes posteriorly. Dorsum of the frontal 
carinae with an impressed, median sulcus anteriorly. Frontal carinae reaching the 
anterior border of the clypeus. Compound eyes large, gently convex and largely on the 
posterior half of the head. Ocelli well developed. Scapes reaching the anterior border of 
the compound eyes. Proximal third of the scapes ca. 1/2 narrower than the remaining 
parts. Mandibles massive and strongly convex dorsally. Masticatory margin of the 
mandibles each with a set of 10-12 irregular, minute denticles followed by an apical 
tooth. 

Mesosoma dorsally flat and slightly more than 1/3 longer than the head (man- 
dibles included). Pronotal dorsum with the sides superficially marginate. Propleurae 
concave. Mesopleurae gently convex. Propodeal sides converging posteriorly. Basal 
and declivous faces of the propodeum subequal in size and delimited by a superficial 
margin. 

Petiole ca. 1/5 longer than broad, anteriorly truncate and the dorsally gently 
convex. Petiolar sides diverging backwards. Ventral process of the petiole large, sub- 
round or obliquely truncate. Postpetiole subquadrate and slightly broader posteriorly. 
Postpetiole in dorsal view antero-laterally angulate. Postpetiolar sternite antero-me- 
dially only with superficial traces of a triangular "lip". Pygidium in side view sub- 
truncate. Pygidium in dorsal view with the sides bearing many irregularly distributed 
small denticles converging to 4-6 small teeth over the sting. 

Legs. Femora and tibiae not strongly inflated. Hind basitarsi 1/4 shorter than the 
maximum length of the hind tibiae. Outer apical edge of the hind and of the mid 
basitarsi with 5 spine-like setae. 

Wings as in Fig. 4. 

Sculpture. Head covered by longitudinal striae, thicker on the posterior half of 
the head dorsum. Striae close to the antennal scrobes thinner than those on the remai- 
ning parts of the anterior half of the head. Dorsum of the pronotum with about 18-21 
striae similar to those on the posterior part of the head dorsum. Center of the 
mesonotum with about 9-12 striae, thinner than those on the pronotum; remaining parts 
of the mesonotum and scutellum smooth, or sides of the mesonotum with thin, super- 
ficial, short striae. Dorsum of the propodeum with about 21-24 striae similar to those on 
the mesonotum. Propleurae, lower mesopleurae, metapleurae and sides of the petiole 
minutely and superficially reticulate-punctated and with longitudinal striae similar to 



614 MARIA L. DE ANDRADE 

those close to the atennal scrobes. Upper mesopleurae smooth. Petiolar dorsum with 
about 15-17 striae similar to those on the propodeum. Declivous face of the propodeum 
and anterior face of the petiole minutely reticulate-punctate. Postpetiolar dorsum 
smooth and sometimes with very thin, short, superficial striae on the center of the pos- 
terior half. Postpetiolar sternite and gaster smooth and with variably impressed punctu- 
ations, denser and larger on the the postpetiolar sternite. Last three gastric sternites and 
sides of their corresponding tergites minutely and superficially reticulate-punctated. 
Coxae not striated. Legs with very superficial, minute punctures. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) long, erect to suberect, rare on the head, on the mandibles, on the anterior 
border of the clypeus, on the mesosoma, on the pedicel, on the ventral process of the 
petiole and on the gaster, dense on the pygidium; (2) shorter than the type (1) rare and 
suberect on the whole body except on the sternites these hairs are sub- or decumbent; 
(3) shorter than the type (2), erect to suberect on the whole body except on the posterior 
half of the ventral part of the head, on the gaster and on the legs these hairs are sub- or 
decumbent. In addition, the hypostomal bridge surrounded by a row of hairs similar to 
those of type (1) but appressed and apically curved. 

Colour black and shining. Legs dark orange-brown with darker tarsi and black 
coxae. Imature specimens with mandibles, antennae, coxae and pygidium reddish- 
brown, last funicular joints orange. 

Measurements (in mm) and indices: TL 9.64-10.28: HL 1.60-1.64; HW 1.24-1.28; EL 
0.50-0.54; SL 0.65-0.67; SW 0.23-0.24; WL 2.72-2.76; PeL 0.90-1.00; PeW 0.80-0.81; HFeL 
0.88-1.02; HFeW 0.37-0.45; HTiL 0.80-0.92; HTiW 0.24-0.29; HBaL 0.60-0.65; HBaW 0.10- 
0.1 1; CI 77.5-78.0; SI 35.4-35.8; HFel 42.0-43.1; HTil 30.0-32.5; HBal 16.4-16.9. 

Male (Fig. 15). Head as broad as long. Ocelli protuberant. Compound eyes 
broadly convex and largely on the anterior half of the head. Frontal carinae high. 
Borders of the frontal carinae broad, convex on the anterior third and subparallel 
posteriorly. Frons anteriorly concave, medially gently convex and posteriorly sloping to 
the impair ocellus. Anterior border of the clypeus convex medially. Mandibles long; 
their masticatory margin edentated and with a pointed apical tooth. Scapes short and 
thick. Funicular joints stout; first joint about 1/2 shorter than the second one. Second 
and last two funicular joints thinner than joints 3-10. 

Mesosoma robust. Pronotum in dorsal view with diverging sides. Mesonotum 
convex. Parapsidal furrows superficially impressed. Scutellum subround and as high as 
the mesonotum. Basal face of the propodeum narrowing backwards and separated from 
the declivous one by a marked carina. Posterior border of the basal face of the pro- 
podeum with a short sulcus in the middle. 

Petiole slightly longer than broad, broader on the posterior half. Anterior face of 
the petiole truncate and separated from the dorsal one by a marked carina. Ventral 
process of the petiole subtriangular. Postpetiole with the sides gently convex and 
narrower than the first gastric tergite. 

Genitalia as in Fig. 16. 

Legs not inflated. Hind basitarsi about 1/4 shorter than the hind tibiae. Mid 
basitarsi slightly more than 1/2 of the lenght of the hind basitarsi. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 615 

Wings as in Fig. 5. 

Sculpture. Head dorsum minutely punctate and striated, the punctures more 
impressed on the anterior half, the striae thicker on the posterior half, slightly longi- 
tudinal and short on the frons, concentric and irregular close to the internal border of 
the eyes, and converging from the posterior border of the compound eyes to the pair 
ocelli. Vertexal angles and sides of the ventral part of the head with small, deep, pili- 
gerous foveae, larger on the vertexal angles. Middle of the ventral part of the head with 
thick tranversal striae. Pronotum smooth and with sparse piligerous foveae on the 
center; some specimens with additional irregular, transversal rugosities between the 
foveae. Mesonotum and scutellum smooth, with rare, small foveae. Basal face of the 
propodeum and petiole covered by thick, irregular, longitudinal striae, sometimes mis- 
sing on the center of the petiole. Declivous face of the propodeum punctate and with 
rare, very thin, transversal rugosities close to the borders. Pro- and mesopleurae 
smooth. Metapleurae striated as on the basal face of the propodeum. Postpetiole, first 
gastric segment and legs smooth and with superficial punctures, denser and deeper on 
the three last gastric segments. 

Pilosity. Body covered by pointed hairs of four types: (1) long, sparse and 
suberect; (2) shorter than the type (1), sparse and suberect, dense, decumbent on the 
gaster and on the femora; (3) shorter than the type (2) dense, decumbent on the vertexal 
angles, on the posterior half of the ventral part of the head, appressed on the mandibles 
on the scapes, on the first funicular joints, on the coxae, on the tarsi and tarsomeres; (4) 
very short, thick and dense on the 2-12 funicular joints. 

Colour. Black and shining. Mandibles, antennae and legs lighter. 

Measurements (in mm) and indices: TL 8.22-9.54; HL 1.16; HW 1.16; EL 0.60-0.62; SL 
0.25-0.27; SW 0.18-0.19; WL 2.64-2.74; PeL 0.76; PeW 0.70-0.74; HFeL 1.02-1.09; HFeW 
0.23-0.25; HTiL 0.89-0.92; HTiW 0.18-0.19: HBaL 0.71-0.74; HBaW 0.08; CI 100.0; SI 66.7- 
73.1; HFel 21.1-24.5; HTil 19.8-20.6; HBal 10.8-1 1.3. 

Material examined. COLOMBIA: Cundinamarca: Medina, Quebrada Ardita, 1475 m, 
28.11.1997, 1 gyne, F. Escobar [IAVH]. VENEZUELA: Aragua: Rancho Grande, 17.VII.1972, 

I gyne, N. A. Weber [MCZC]; same locality, 1100 m, IV.1987. 1 gyne, C. Bordon [MIZA]; 
same locality, 1200 m, 15.V & 22.VI.1987, 3 males, C. Bordon [MIZA]. PERU: Lima: 
Calanga, 1 male, Staudinger [MCSN]. BOLIVIA: Songo, 1 male [MCSN]. La Paz: Mapiri, 1 
gyne (holotype), Staudinger [MCZC]. 

Discussion. Boliviae is known only on the sexuals. The gyne of boliviae is easily 

distinguished from all other species by the characters already listed before and by the 

very broad frontal carinae reaching the internal border of the eyes. In body shape the 

gyne of boliviae resembles the one of godmani of the longiceps clade and the worker of 

escobari of the brevitarsus clade. Boliviae, godmani and escobari have broad frontal 

carinae and large, convex mandibles. Boliviae and godmani share also a broad and large 

ventral process of the petiole, and boliviae and escobari have mandibles with more than 

I I denticles, and no gastric striae. 

The male of boliviae can be distinguished from the other Cylindromyrmex males 
by the legs dark brown or black. Boliviae males, in addition, have the frontal carinae 
more similar to males of the brevitarsus clade than to males of the striatus or longiceps 
clades. 



6 1 6 MARIA L. DE ANDRADE 

Emery (1901) attributed with doubts two Peruvian males to "striatus". I found in 
the Emery collection only one of these males and it belongs to boliviae. 

The size of the eyes of workers was considered an important diagnostic 
character in Cylindromyrmex. The workers of the longiceps clade have small and flat 
eyes. The workers of the brevitarsus clade have the eyes as in the species of the 
longiceps clade or slightly larger. The workers of the striatus clade have relatively large 
eyes. There is no difference in the size of the eyes of gynes of species with workers 
with large or small eyes. 

Material available for the present study proves that the range of boliviae, 
previously known only from Bolivia and Venezuela, is much broader than what was 
previously supposed (Fig. 38). A Bolivian locality (Songo) has not been plotted on the 
map of Fig. 38 because I was unable to localte it. 

Distribution. Colombia, Venezuela, Peru and Bolivia. 

The brevitarsus clade 

This clade includes four species: escobari, electrinus, brevitarsus and darling- 
toni. They are characterized by the following synapomorphies: (1) occiput high, (2) 
ventral process of the petiole triangular, and (3) hind basitarsi wit 4 spine-like setae. 

Cylindromyrmex escobari n. sp. Fig. 17 

Cylindromyrmex brasiliensis Emery, Fernandez-C. & Escobar, 1997: 347. Worker. Nee Emery 

1901. Misidentification. 

Holotype: Worker labelled: "Colombia, Narino, Ricaurte La Planada, 1°17' N 78° 15' W, 
1800 m, interior bosque, bmh-PM, Col: F. Escobar", in IAVH. 

Derivano nominis. C. escobari is named after Federico Escobar, the collector of 
this species. 

Diagnosis. The basalmost species of the brevitarsus clade, differing from all the 
other species by the antero-median margin of the clypeus convex and by the absence of 
striae on the gaster. 

Worker (Fig. 17). Head ca. 1.5 times longer than broad, with subparallel sides. 
Occiput high. Vertexal angles convex. Frontal carinae more than half broad as the 
maximum head width. Anterior third of the frontal carinae diverging backwards and 
reaching at least the middle of the eyes posteriorly. Dorsum of the frontal carinae with 
an impressed, broad, median sulcus anteriorly. Frontal carinae not reaching the anterior 
border of the clypeus. Antero-median clypeal margin strongly convex. Compound eyes 
intermediate in size between the species of the longiceps and striatus clades, slightly 
flat and on the posterior half of the head. Ocelli represented by superficial impression 
only. Scapes stout and surpassing the anterior border of the eyes posteriorly. Proximal 
third of the scapes 1/2 narrower than the remaining parts. Mandibles strongly convex 
dorsally. Masticatory margin of the mandibles each with a set of 13-14 irregular den- 
ticles followed by an apical tooth. 

Fig. 17. C. escobari de Andrade. Worker from Ricaurte, La Planada, Narino, Colombia. Head in 
full dorsal view (top), body in full dorsal view (middle), body in profile (bottom). 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



617 




00007978 



1 mm 



REM-Labor 
Uni Basel 



6 1 8 MARIA L. DE ANDRADE 

Mesosoma gently convex dorsally and slightly less than 1/5 longer than the head 
(mandibles included). Pronotum with parallel sides. Promesonotal and propodeal 
sutures superficially impressed. Mesonotum slightly narrower than pronotum. Tegula 
superficially marked. Propodeum with the sides weakly convex. Basal face of the 
propodeum separated from the declivous one by a superficial margin. 

Petiole rectangular, anteriorly truncate and the dorsally convex. Petiolar sides 
diverging backwards. Ventral process of the petiole large and subtriangular. Postpetiole 
ca. 1.5 broader than long. Postpetiolar sides diverging posteriorly. Postpetiole in dorsal 
view antero-laterally angulate. Postpetiolar sternite antero-medially with a superficial 
triangular "lip" pointing backwards. Pygidium in side view obliquely truncate. Pygi- 
dium in dorsal view with the sides bearing many irregularly distributed small denticles 
converging to 4 small teeth over the sting. 

Legs. Coxae and tibiae slightly inflated. Mid basitarsi strongly broadening 
distally. Hind basitarsi about 1/3 shorter than the maximum length of the hind tibiae. 
Mid basitarsi 1/2 of the length of the hind basitarsi. Outer apical edge of the hind and of 
the mid basitarsi respectively with 4,5 spine-like setae. 

Sculpture. Head covered by thin, longitudinal striae, slightly thicker on the 
posterior third of the head dorsum and on the center of the ventral part of the head, 
absent on the posterior corners of the ventral part of the head. Mesosoma longitudinally 
striated. Pronotum and mesonotum with striae thicker than those on the propodeum. 
Pronotum with about 28-30 longitudinal striae similar to those on the posterior part of 
the head dorsum. Propodeum with about 30-35 longitudinal striae. Pleurae with very 
thin, superficial, longitudinal striae, less impressed on the propleurae. Petiolar dorsum 
with about 30-35 striae slightly thinner than those on the propodeum. Petiolar sides 
with very thin, superficial striae. Declivous face of the propodeum and anterior face of 
the petiole with superficial reticulation. Dorsum of the postpetiole with striae thinner 
than those on the petiolar dorsum. Remaining gastric tergites, sternites and legs smooth, 
with minute, superfical reticulation more impressed on the distal segments of the gaster. 
Ventral face of the hind coxae with thin, longitudinal striae. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) long, erect to suberect, sparse on the head, on the mesosoma, on the pedicel 
and on the gaster, dense on the pygidium; (2) shorter than the type (1), suberect and 
variably distributed on the whole body; (3) shorter than the type (2), sparse, suberect or 
subdecumbent on the whole body. In addition, the hypostomal bridge surrounded by a 
layer of hairs similar to the type (1) but appressed and apically curved. Outer ventral 
border of the mandibles with hairs similar to those of the hypostomal bridge but shorter. 

Colour black. Mandibles, antennae and coxae dark ferrugineous-brown. Legs 
yellow-orange to light brown with darker tarsi. 

Measurements (in mm) and indices: TL 8.16; HL 1.56; HW 1.28; SL 0.732 SW 0.25; 
WL 2.16; PeL 0.68; PeW 0.56; HFeL 1.00; HFeW 0.37; HTiL 0.85; HTiW 0.26; HBaL 0.50; 
HBaW 0.09; CI 82.0; SI 34.7; HFel 37.0; HTil 30.5; HBal 18.0. 

Material examined. COLOMBIA: Narino: Ricaurte, Reserva La Plananda, 1°17' N 
78°15'W, 1800 m. 1 worker (holotype), F. Escobar [IAVH]. 

Discussion. Escobari differs from the other species of the brevitarsus clade 
mainly by the absence of striae on the first gastric tergite. In particular, it differs from 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 6 1 9 

electrinus by the larger and more massive mandibles with 13-14 denticles instead of 
smaller and less massive and with 6-7 denticles. From darlingtoni and brevitarsus, 
escobari differs by the more elongate body. Escobari in general body shape resembles 
more darlingtoni than brevitarsus. A comparison of escobari and darlingtoni proves 
that they are very different each other. Escobari can be separated from darlingtoni by 
the strongly convex anterior border of the clypeus, by the frontal carinae not reaching 
the anterior clypeal border and by the more enlongate femora. 

Comparisons were made also between the worker of escobari and the gynes of 
boliviae and godmani, two species the workers of which are still unknown and occur- 
ring close to the area where escobari was collected. Escobari has concolour femora and 
tibiae (yellowish-orange to light brown) and godmani has black femora and yellow 
tibiae. Escobari differs from boliviae by the postpetiole striate instead of smooth or 
with very superficial, short striation restricted to the center of the posterior half, and by 
thinner striation. 

Fernandez- C. & Escobar (1997) reported this species from decayed wood. 

Distribution. Colombia. 

Cylindromyrmex electrinus sp. n. Figs 2 & 18 

Holotype: Winged gyne in an amber sample without reference number from the MCZC. 

Derivatio nominis. From the Latin electrinus (= made of amber). 

Diagnosis. A species appearing in an unresolved position together with brevi- 
tarsus and darlingtoni within the brevitarsus clade, but differing from both other 
species by the following combination of characters: basal face of the propodeum sepa- 
rated from the declivous one by a marked margin, by the coxae and femora black 
instead of dark brown, and by the mid basitarsi long and not broad distally. 

Gyne (Figs 2 & 18). Head ca. 1/4 longer than broad, with parallel sides. Occiput 
high. Vertexal angles convex. Frontal carinae about half broad as the maximum head 
width. Anterior third of the frontal carinae diverging backwards. Dorsum of the frontal 
carinae with an impressed, median sulcus anteriorly. Frontal carinae reaching the 
anterior border of the clypeus. Compound eyes large, slightly flat and mostly on the 
posterior half of the head. Ocelli well developed. Scapes reaching the anterior border of 
the eyes. Proximal third of the scapes about 1/2 narrower than the remaining parts. 
Mandibles convex dorsally. Masticatory margin of the mandibles with a set of 6-7 
irregular denticles followed by a pointed apical one. 

Mesosoma dorsally flat and ca. 1.3 longer than the length (mandibles included). 
Pronotum dorsally with the sides superficially marginate. Propleurae gently concave. 
Mesopleurae gently convex. Propodeum with the sides converging posteriorly. Basal 
and declivous faces of the propodeum subequal in size and delimited by a margin. 

Petiole ca. 1/4 longer than broad, snteriorly truncate and the dorsally convex. 
Ventral process of the petiole triangular. Postpetiole diverging backwards and broader 
posteriorly. Anterior corners of the postpetiole angulate. Postpetiole in dorsal view 
antero-laterally angulate. Postpetiolar sternite antero-medially with a salient, triangular 
"lip" pointing backwards. Pygidium obliquely truncate; its sides bearing many irregu- 
larly distributed small denticles converging to 4 small teeth over the sting. 



620 



MARIA L. DE ANDRADE 






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FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 621 

Legs. Femora slightly inflated. Mid basitarsi with parallel sides. Hind basitarsi 
slightly less than 1/3 shorter than the length of the hind tibiae. Mid basitarsi ca. 1/2 of 
the length of the hind basitarsi. Outer apical edge of the hind and of the mid basitarsi 
respectively with 4,5 spine-like setae. 

Wings as in Fig. 4. 

Sculpture. Head covered by thin, longitudinal striae, thicker on the posterior part 
of the dorsum, thinner close to the antennal scrobes, absent on the posterior third of the 
ventral part of the head. Posterior third of the ventral part of the head minutely punctate 
and smooth. Dorsum of the pronotum with at least 40 striae similar to those on the 
posterior part of the head dorsum. Mesonotum with at least 25 striae thinner than those 
on the pronotum. Scutellum covered by striae slightly thinner than those on the meso- 
notum. Dorsum of the propodeum covered by striae similar to those on the mesonotum. 
Pleurae and petiole with longitudinal striae similar to those on the scutellum, the striae 
more superficial on the upper mesopleurae and on the sides of the petiole. Petiolar 
dorsum with at least 25 striae. Postpetiole covered by at least 50 striae similar to those 
on the mesonotum. First gastric tergite with very thin, short striae on the center of the 
anterior half only. Remaining gastric tergites and sternites smooth and with variably 
impressed punctuations more impressed on the last segments. Legs with very super- 
ficial, minute punctures. Hind coxae with thin, longitudinal striae. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) long, erect to suberect, one on the external border of the scape, a pair 
between the frontal carinae and the clypeus, rare on the external border of the man- 
dibles, rare on the gaster, sparse on the pygidium; (2) shorter than the type (1) and 
sparsely distributed on the whole body; (3) shorter than the type (2), erect to suberect, 
sparse on the whole body. In addition, the hypostomal bridge surrounded by a layer of 
hairs similar to the type (1). 

Colour black. Tibiae of three pairs of legs partially yellowish and transparent or 
dark brown. Tarsi dark brown, tarsomeres lighter. 

Measurements (in mm) and indices: TL 7.36; HL 1.24; HW 0.94; EL 0.36; SL 0.46; SW 
0.16; WL 2.20; PeL 0.68; PeW 0.56; HFeL 0.75; HFeW 0.30; HTiL 0.64; HTiW 0.21; HBaL 
0.47; HBaW 0.08; CI 75.8; SI 36.9; HFel 40.0; HTil 32.8; HBal 17.0. 

Material examined. Dominican amber: 1 gyne (holotype) without reference number 

[Mczq. 

Discussion. C. electrinus, in the phylogeny proposed in this paper, appears close 
to the Recent brevitarsus and darlingtoni. These three species differs from the basal- 
most species of the clade, escobari, by the presence of striae on the first gastric tergite. 
Electrinus shares with brevitarsus the frontal carinae reaching the anterior border of the 
clypeus and the mandibles with 6-7 denticles, and with darlingtoni the striae on the first 
gastric tergite thin, short and restricted on the anterior part only. Electrinus is very 
similar to both brevitarsus and darlingtoni, but the characters listed in the diagnosis 
allow an easy separation of the fossil from both Recent species. 

Distribution. Dominican amber. 



622 MARIA L. DE ANDRADE 

Cylindromyrmex darlingtoni Wheeler Fig. 19 

Cylìndromyrmex darlingtoni Wheeler, 1937: 441. Worker and gyne. Original description. Type 
locality: Cuba. Type material: 2 workers and 2 gynes labelled: "Gran Piedra Rge. Ote, 
Cuba, 2-3000 ft., 30.VI.1936, P. J. Darlington, MCZ cotype", in MCZC, examined. 

Diagnosis. A species belonging to the brevitarsus clade and resulting in a un- 
resolved position together with brevitarsus and electrinus, but differing from both by 
the frontal carinae surpassing the anterior border of the clypeus instead of as long as the 
clypeus, and by the mandibles with 9-10 denticles instead of 6-8. Darlingtoni differs 
from electrinus by the mid and fore basitarsi shorter and broader distally instead of long 
and with parallel sides, and from brevitarsus, by the ventral face of the hind femora 
with only traces of longitudinal striae instead of markedly striate. 

Worker (Fig. 19). Head ca. 1.5 times longer than broad, with parallel sides. 
Occiput high. Vertexal angles convex. Frontal carinae about half broad as the maximum 
head width. Frontal carinae anteriorly diverging backwards and reaching at the middle 
of the eyes posteriorly. Dorsum of the frontal carinae with an impressed, broad, median 
sulcus anteriorly. Frontal carinae slightly longer than the anterior border of the clypeus. 
Compound eyes small, flat and behind the mid line of the head. Impar ocellus minute, 
pair ocelli reduced to a superficial pit. Scapes almost reaching the anterior border of the 
eyes. Proximal third of the scapes 1/2 narrower than the distal parts. Mandibles convex 
dorsally. Masticatory margin of the mandibles each with a series of 9-10 irregular 
denticles followed by an apical one. 

Mesosoma convex dorsally and as long as the head (mandibles included). Pro- 
notum with parallel sides. Promesonotal and propodeal sutures superficially impressed. 
Mesonotum narrower than pronotum. Propodeal sides gently convex and converging 
posteriorly. Basal face of the propodeum separated from the declivous one by a faint 
margin. 

Petiole subquadrate, slightly broader than long, anteriorly truncate and dorsally 
convex. Petiolar sides diverging backwards. Ventral process of the petiole large and 
subtriangular. Postpetiole ca. 1.3 broader than long. Postpetiolar sides gently diverging 
posteriorly. Postpetiole in dorsal view gently angulate antero-laterally. Postpetiolar 
sternite antero-medially with a superficial, triangular lip pointing backwards. Pygidium 
truncate; its sides bearing many irregularly distributed, small denticles converging to 4 
small teeth over the sting. 

Legs. Femora and tibiae slightly inflate. Fore and mid basitarsi strongly 
broadening distally. Hind basitarsi short, ca. 1/2 shorter than the maximum length of the 
hind tibiae. Outer apical edge of the hind and of the mid basitarsi respectively with 4,5 
spine-like setae. 

Sculpture. Head covered by thin, longitudinal striae, thicker on the posterior 
third of the head dorsum, absent on the angles of the ventral part of the head. Meso- 



Fig. 19. C. darlingtoni Wheeler. Worker from Gran Piedra, Range Oriente, Cuba. Head in full 
dorsal view (top), body in full dorsal view (middle), body in profile (bottom). 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



623 





00008089 



500 Mm 



REM-Labor 
Uni Basel 



624 MARIA L. DE ANDRADE 

soma longitudinally striated, some mesosomal striae bifurcated. Dorsum of the pro- 
notum with about 40 longitudinal striae similar to those on the posterior part of the head 
dorsum. Pronotal striae prolonging to the dorsum of the mesonotum and propodeum. 
Pleurae with thin, superficial, longitudinal striae, less impressed on the propleurae. 
Petiolar dorsum with about 30 striae similar to those on the pronotum. Petiolar sides 
with minute and superficial reticulation. Declivous face of the propodeum superficially 
and sparsely reticulate. Anterior face of the petiole smooth. Dorsum of the postpetiole 
densely covered by striae similar to those on the petiolar dorsum. Second gastric tergite 
with extremely thin, superficial, longitudinal striae on the center of the anterior half 
only. Postpetiolar sternite and remaing gastric segments smooth and with sparse punc- 
tures. Pygidium, border of the sternites, and legs superficially reticulate. Hind coxae 
with traces of longitudinal striae. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) long, erect to suberect, one on the external border of the scape, a pair 
between the frontal carinae nd clypeus, rare on the mandibles, on the mesosoma, on the 
pedicel, on the gaster and on the legs, sparse on the pygidium; (2) shorter than the type 
(1) and sparsely distributed on the whole body; (3) shorter than the type (2), suberect on 
the head dorsum and mesosoma, subdecumbent on the pedicel, decumbet on the ventral 
part of the head, on the gaster and on the legs. In addition, the hypostomal bridge 
surrounded by a layer of hairs similar to the type (1) but appressed and apically curved. 

Colour black. Mandibles and antennae browish red. Coxae and femora dark 

brown. Last funicular joints, tibiae and tarsomeres yellowish-orange, tarsi darker. 

Measurements (in mm) and indices: TL 6.52; HL 1.28; HW 1.08; EL 0.25; SL 0.54; SW 
0.20; WL 1.64; PeL 0.58; PeW 0.60; HFeL 0.73; HFeW 0.33; HTiL 0.63; HTiW 0.23; HBaL 
0.34; HBaW 0.08; CI 84.4; SI 37.0; HFel 45.2; HTil 36.5; HBal 23.5. 

Gyne. Very similar to the worker but differing from it in the following details: 
compound eyes very large, flat and mostly on the posterior part of the head; ocelli well 
defined; mesosoma broad medially; parapsidal furrows superficially impressed; petiole 
slightly longer than broad; anterior half of the mesonotum and of the scutellum with 
very thin, superficial striae; posterior half of the mesonotum and of the scutellum with 
few traces of short striae or simply smooth. 

Measurements (in mm) and indices: TL 8.16-8.44; HL 1.34-1.36; HW 1.08-1.10; EL 
0.46-0.47; SL 0.57; SW 0.21; WL 2.28-2.36; PeL 0.70-0.72; PeW 0.66; HFeL 0.76-0.77; HFeW 
0.35-0.36; HTiL 0.66; HTiW 0.24; HBaL 0.40; HBaW 0.08; CI 80.6-80.9; SI 36.8; HFel 45.4- 
46.0; HTil 36.4; HBal 20.0. 

Material examined. CUBA: Gran Piedra Range, Oriente, 2-3000 ft 30. VI. 1936, 2 wor- 
kers, 2 gynes (all syntypes), P. J. Darlington, [MCZC]. 

Discussion. C. darlingtoni is the northernmost species of the genus. It is known 
only from the type series from Cuba and it is likely to be endemic on the island. 
Differences between darlingtoni and its closest relatives, brevitarsus and electrinus, are 
listed in the diagnosis and in the discussion of these species. 

The type series of darlingtoni was collected in decayed wood. 

Distribution. Cuba. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 625 

Cylindromyrmex brevitarsus Santschi Figs 20-24 

Cylindromyrmex brevitarsus Santschi, 1925: 5. Worker. Original description. Type locality: 

Brazil. Type material: 1 worker labelled: "Brésil, Rio Negro, Reichensperger", in NHMB, 

examined. 

Diagnosis. A species belonging to the homonymous clade and resulting in an 
unresolved position with darlingtoni and electrinus, but differing from darlingtoni by 
the frontal carinae reaching the anterior border of the clypeus instead of longer than the 
clypeus, and by the mandibles with 6-8 denticles instead of 9-10; and from electrinus, 
by the mid and fore basitarsi shorter and broader distally instead of long, with parallel 
sides, and by the yellow-light brown femora instead of black. 

Worker (Fig. 20). Head ca. 1.6-1.7 times longer than broad, with subparallel 
sides. Occiput high. Vertexal angles convex. Frontal carinae about half broad as the 
maximum head width. Anterior half of the frontal carinae diverging backwards and 
reaching the middle of the eyes posteriorly. Dorsum of the frontal carinae with an 
impressed, broad, median sulcus anteriorly. Frontal carinae as long as the anterior 
border of the clypeus. Compound eyes small (minimum 30 and less than 150 omma- 
tidia), flat and on the posterior half of the head. Ocelli reduced to very superficial pits. 
Scapes reaching the anterior border of the eyes. Proximal third of the scapes 1/2 
narrower than the distal parts. Mandibles convex dorsally. Masticatory margin of the 
mandibles each with a series of 6-8 irregular denticles followed by an apical one. 

Mesosoma convex dorsally and slightly longer or shorter than the head (man- 
dibles included). Pronotum with parallel sides. Promesonotal and propodeal sutures less 
impressed than in darlingtoni. Mesonotum slightly narrower than pronotum. Propo- 
deum with the sides gently convex and converging posteriorly. Basal face of the pro- 
podeum separated from the declivous one by a faint margin. 

Petiole quadrate, slightly broader than long, anteriorly truncate and the dorsally 
convex. Ventral process of the petiole large and triangular. Postpetiole ca. 1.4 broader 
than long. Postpetiolar sides gently diverging posteriorly. Postpetiole in dorsal view 
slightly angulate antero-laterally. Postpetiolar sternite antero-medially with traces of a 
triangular "lip". Pygidium truncate; its sides bearing many irregularly distributed den- 
ticles converging to 4-6 small teeth over the sting. 

Legs. Femora and tibiae inflated. Fore and Mid basitarsi strongly broadening 
distally. Hind basitarsi short, ca. 1/2 shorter than the maximum length of the hind tibiae. 
Mid basitarsi about 1/2 shorter than the hind basitarsi. Outer apical edge of the hind and 
of the mid basitarsi respectively with 4,5 spine-like setae. 

Sculpture. Head covered by thin, longitudinal striae, thicker on the posterior 
third of the head dorsum and absent on the corners of the ventral part of the head. 
Mesosoma longitudinally striated. Dorsum of the pronotum with about 34-38 longitu- 
dinal striae similar to those on the posterior part of the head dorsum. Prenotai striae 
prolonging to the dorsum of the mesonotum and propodeum. Pleurae with thin, super- 
ficial, longitudinal striae, less impressed on the propleurae. Petiolar dorsum with about 
31-35 striae similar to those on the propodeum. Petiolar sides minutely and superfi- 
cially reticulate. Declivous face of the propodeum and anterior face of the petiole 



626 



MARIA L. DE ANDRADE 




00007955 



500 pin 



REM-Labor 
Uni Basel 



FOSSIL AND EXTANT SPECIES OF CYL1NDROMYRMEX 



627 




00007986 



1 mm 



REM-Labor 
Uni Basel 



628 



MARIA L. DE ANDRADE 




REM-Labor 
Uni Basel 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



629 




0-5mm 



Fig. 23 

C. brevitarsus Santschi. Male from Corcovado, Rio de Janeiro, Brazil. Genital appendages: a) 
lateral view of left parameres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 



Fig. 20. C. brevitarsus Santschi. Worker from Corcovado, Rio de Janeiro, Brazil. Head in full 
dorsal view (top), body in full dorsal view (middle), body in profile (bottom). 

Fig. 21. C. brevitarsus Santschi. Gyne from Serra do Mar, Nova Friburgo, Rio de Janeiro, Brazil. 
Head in full dorsal view (top), body in full dorsal view (middle), body in profile (bottom). 



Fig. 22. C. brevitarsus Santschi. Male from Corcovado, Rio de Janeiro, Brazil. Head in full 
dorsal view (top), body in full dorsal view (middle), body in profile (bottom). 



630 



MARIA L. DE ANDRADE 




0.5 mm 



Fig. 24 

C. brevitarsus Santschi. Male from Butantan, Sào Paulo, Brazil. Genital appendages: a) lateral 
view of left parameres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 



smooth or with similar sculpture as on the petiolar sides. Dorsum of the postpetiole 
densely covered by striae similar to those on the petiolar dorsum. Center of the first 
gastric tergite with thin, short, longitudinal striae, thinner than those on the postpetiole; 
mature specimens have the whole first gastric tergite covered by striae and the second 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 63 \ 

gastric tergite with striae on the center only. Postpetiolar sternite and remaing gastric 
segments smooth and with sparse punctures. Pygidium, border of the sternites, and legs 
superficially reticulate. Hind coxae with thin, longitudinal striate. Mid coxae with few, 
fainter striate than those on the hind coxae. 

Pilosity as in darlingtoni. 

Colour light or dark brown. Legs yellowish-orange to light brown with darker 
coxae and basitarsi. 

Measurements (in mm) and indices: TL 4.20-6.44; HL 0.86-1.20; HW 0.75-0.98; SL 
0.36-0.43; SW 0.16-0.19; WL 1.04-1.50; PeL 0.36-0.56; PeW 0.41-0.58; HFeL 0.49-0.64; HFeW 
0.26-0.30; HTiL 0.40-0.54; HTiW 0.16-0.19; HBaL 0.20-0.29; HBaW 0.06-0.08; CI 81.7-89.2; 
SI 42.5-45.0; HFel 45.3-46.9; HTil 35.2-40.5; HBal 26.9-31.8. 

Gyne (previously undescribed) (Fig. 21). Very similar to the worker but differ- 
ing from it in the following details: compound eyes very large, flat or gently convex 
and largely on the posterior part of the head; mandibles with 6-8 denticles; ocelli well 
defined; mesosoma broad medially; parapsidal furrows weakly impressed; petiole 
slightly longer than broad; scutellum with very thin, superficial striae, sometimes on the 
anterior half only; propodeal dorsum with striae thinner than those on the pronotum and 
on the mesonotum. Some gynes have short striae on the second gastric tergite. 

Colour dark brown or black. Antennae, mandibles and coxae dark ferrugineous 

or brown. Some specimens have the anterior half of the head dorsum dark ferrugineous. 

Legs dark yellowish-orange or light brown with darker coxae and tarsi. The gyne from 

Ecuador (LACM) and Serra Norte (MPEG) have the femora light brown with yellowish 

tibiae. 

Measurements (in mm) and indices: TL 5.88-9.40; HL 0.95-1.52; HW 0.80-1.28; EL 
0.35-0.46; SL 0.37-0.64; SW 0.16-0.24; WL 1.52-2.04; PeL 0.47-0.78; PeW 0.45-0.72; HFeL 
0.53-0.93; HFeW 0.25-0.40; HTiL 0.46-0.81; HTiW 0.18-0.27; HBaL 0.28-0.50; HBaW 0.06- 
0.09; CI 78.5-84.5; SI 37.5-43.2; HFel 42.5-47.9; HTil 32.4-39.1; HBal 17.4-22.6. 

Male (previously undescribed) (Fig. 22). Head slightly longer than broad. Ver- 
texal angles converging to the subtruncate vertexal margin. Ocelli protuberant. Com- 
pound eyes broadly convex and largely on the anterior part of the head. Borders of the 
frontal carinae broad, convex anteriorly, converging and subparallel posteriorly. Frons 
anteriorly concave, medially convex and posteriorly sloping towards the impair ocellus. 
Anterior border of the clypeus medially convex. Mandibles long; their masticatory 
margin edentated and with a pointed apical tooth. Scapes about 1/4 longer than broad. 
Funicular joints narrowing from the base to the apex. 

Mesosoma robust. Pronotum in dorsal with the sides diverging posteriorly. 
Mesonotum slightly convex. Mayrian carinae absent. Parapsidal furrows superficialy 
impressed. Basal face of the propodeum narrowing backwards and separated from the 
declivous one by a marked carina. 

Petiole slightly longer than broad; anteriorly truncate and dorsally convex. 
Petiolar sides broadening backwards. Ventral process of the petiole subtriangular. 
Postpetiole broadening backwards and narrower than the first gastric tergite. Postpetiole 
antero-laterally subround. 

First gastric segment broader than the postpetiole. Second gastric segment nar- 
rower or as broad asthe the first segment. Remaining gastric segments narrowing back- 
wards. 



632 MARIA L. DE ANDRADE 

Genitalia as in Fig. 23 (normal size males) and Fig. 24 (large size male). 

Legs. Femora not inflate. Mid and hind basitarsi long. 

Wings as in Fig. 5. 

Sculpture. Head dorsum minutely punctate and with thin striae, the punctures 
more impressed on the anterior half, the striae slightly longitudinal between the ocelli 
and on the frons, diverging from the eyes to the frontal carinae. Vertex and sides of the 
ventral part of the head smooth and with variably distributed small piligerous foveae. 
Middle of the ventral part of the head with short, tranversal striae. Pronotum smooth 
and with sparse piligerous foveae on the center; some specimens with additional 
irregular, transversal striae between the foveae. Mesonotum and scutellum smooth and 
with rare, small foveae. Basal face of the propodeum and petiole covered by thick, 
irregular, longitudinal striae, sometimes missing on the center of the petiole. Declivous 
face of the propodeum punctate and with rare, very thin, transversal rugosities close to 
the border. Pro- and mesopleurae smooth. Metapleurae striated as on the basal face of 
the propodeum, the striae thicker and less regular ventrally. Postpetiole, first gastric 
segment and legs smooth and with superficial punctures, denser and deeper on the three 
last gastric segments. 

Pilosity. Body covered by pointed hairs of three types: (1) long, sparse and 
suberect, denser on the last three gastric segments; (2) shorter than the type (1) and 
variably distributed on the body; (3) mixed and shorter than the type (2), dense on the 
vertexal angles, on the posterior half of the ventral part of the head, on the coxae and on 
the gaster. 

Colour. Black and shining. Some specimens with the anterior third of the head 
dorsum, mandibles and antennae orange-ferrugineous or brown. Legs orange-light 
brown with darker coxae and basitarsi. 

Measurements (in mm) and indices: TL 5.98-8.96; HL 0.85-1.18; HW 0.81-1.24; EL 
0.44-0.61; SL 0.19-028; SW 0.13-0.19; WL 1.80-2.84; PeL 0.48-0.84; PeW 0.43-0.76; HFeL 
0.68-1.06; HFeW 0.17-0.23; HTiL 0.59-0.85; HTiW 0.14-0.19; HBaL 0.44-0.69; HBaW 0.05- 
0.07; CI 89.6-105.1; SI 57.1-73.1; HFel 22.3-27.9; HTil 21.0-25.4; HBal 10.1-13.6. 

Material examined. VENEZUELA: Aragüa: Rancho Grande, 1100 m, 9.IV.1987, 1 
gyne [MIZA]; same locality, 1200 m, 22.VI.1987, 1 gyne, C. Bordon [MIZA]; same locality, 
1100 m, 28.X.1987, 1 gyne, C. Bordon & H. Romero [MIZA]. ECUADOR: Pichincha: 
Tinalandia, 16 km SE of Santo Domingo de los Colorados, VI. 1975, 1 male, S. Peck & J. Peck 
[MCZC]; Sucumbios, 0.5° S, 76.5° W, 290 m, Sacha Lodge, 22.II-4.III.1994, 1 gyne, 1 male, P. 
Hibbs [LACM]. PERU: Apurimac: no further locality, 14VIII.1962, 1 worker, M. Dourojeanni 
[USNM]. BRAZIL: Para: Serra Norte, Estaçào. Manganês, 5-9.IX.1983, 1 gyne, F. F. Ramos 
[MPEG]. Goias: Jataf, XII. 1972, 1 gyne. F. M. Oliveira [MZSP]. Bahia: Ilhéus, Reserva 
Botànica. CEPEC, 23-24.IV. 1987. 1 male [CPCC]. Mato Grosso: Sinop, 55°37' W, 12°31' S, 
X.1974, 1 male, M. Alvarenga [MZSP]. Rio de Janeiro: lina Grande, 16.X.1944, 2 workers, H. 
Sick [MCZC, MZSP]; Rio de Janeiro, Corcovado, 25.IX.1962, 4 workers, 21 gynes, 23 males, R. 
L. Araujo [MZSP]; Guanabara, Floresta da Tijuca, 1.1974, C. A. C. Seabra & M. Alvarenga 
[MZSP]; Nova Friburgo, Serra do Mar, Bacco farm, forest, 4769 ft., 1991, 2 gynes, K. P. Bland 
[BMNH]; Silva Jardim, Vili. 1974, 1 gyne, F. M. Oliveira [MZSP]. SÄO Paulo: Sào Paulo, 
Butantan, Horto Osvaldo Cruz, 17.1.1971, 1 male, L. Travassos Filho [MZSP]. Parana: Rio 
Negro, 1 worker (holotype). Reichensperger [NHMB]; same locality and collector, 1 gyne, 
[NHMB]; same locality, II. 1929, 1 male [NHMB]. 

Discussion. Brevitarsus is very similar to darlingtoni. Both species can be sepa- 
rated as stated in the diagnosis and, in addition, also by the hind coxae ventrally 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 633 

markedly striate in brevitarsus instead of superficially striate in darlingtoni. Wheeler 
(1937) gave the following characters to separate brevitarsus from darlingtoni: mandi- 
bular shape and dentition, body colour, and size of the eyes. Material available for the 
present study proves that the body colour and the size of eyes are too variable to be 
useful to separate brevitarsus from darlingtoni. The gynes of brevitarsus vary remar- 
kably in size (see measurements). The gynes from Corcovado (MZSP) and Jatai 
(MZSP) are small (TL: 5.88-5.96). Those from Aragua (MIZA), Rio Negro (NHMB), 
Silva Jardim (MZSP), and Sucumbios (LACM) are intermediate (TL: 7.44-7.64). Those 
from Nova Friburgo (BMNH) and from Floresta da Tijuca (MZSP) are the largest (TL: 
8.20-9.40). There seems to be no relevant morphological differences between small and 
intermediate size gynes, only the mandibles are shorter and less convex in the small 
ones. The large gynes differ from the others by the ventral process of the petiole more 
round. The large size gynes also have the mandibles convex and massive as in the 
medium ones. From the material available for the present study I find insufficient 
evidence to regard them as belonging to two (or three) different species. 

A Brazilian male from Butantan which I also consider brevitarsus has larger size 
8.96 instead of 5.98-8.04 (see measurements), more impressed sculpture and darker 
legs. A comparison of its genitalia with those of two males of "normal" size (Fig. 23) 
and (Fig. 24) shows no significant differences. 

Some workers and gynes of brevitarsus may also have the second gastric tergite 
striate. 

Distribution. Venezuela, Ecuador, Peru and Brazil. 

The longiceps clade 

This clade includes four species: godmani, antillanus, longiceps and meinerti. 
They are characterized by the following synapomorphies: (1) first and second gastric 
tergites striate, (2) male frontal carinae strongly broad anteriorly and touching each 
other posteriorly, (3) male hypopygium with a simple, impair, median projection 
between the apodemes. 

Cylindromyrmex godmani Forel Figs 25-27 

Cylindromyrmex godmani Forel, 1899: 4, pi. 1, fig. 2. Gyne. Original description. Type locality: 
Panama. Type material: 1 gyne labelled: "V. de Chiriqui, 2-3000 ft., Champion, 
Holotype, B. C. A. Hym. Cylindromyrmex godmani, Forel, Type", in BMNH, examined. 

Diagnosis. The basalmost species of the longiceps clade differing from all the 
others by the distance between the frontal carinae, about 2/3 of the head width instead 
of about 1/3, and by the superficial gastric striae. 

Gyne (Fig. 25). Head ca. 1/4 longer than broad. Sides of the head behind the 
eyes gently converging posteriorly and in front of the eyes slightly convex. Occiput 
low. Vertexal angles convex. Frontal carinae about 2/3 as broad as the maximum head 
width. Anteriorpart od the frontal carinae gently diverging posteriorly. Dorsum of the 
frontal carinae with an impressed median sulcus anteriorly. Frontal carinae in full face 
view with a deep incision antero-medially and as long as the anterior border of the 



634 



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FOSSIL AND EXTANT SPECIES OF CYUNDROMYRMEX 



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Fig. 26. C. godmani Forel. Male from Turrialba, Costa Rica. Head in full dorsal view (top), body 
in full dorsal view (middle), body in profile (bottom). 



636 



MARIA L. DE ANDRADE 




0-5mm 



Fig. 27 

C. godmani Forel. Male from Turrialba, Costa Rica. Genital appendages: a) lateral view of left 
parameres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 



clypeus. Compound eyes large, gently convex and largely on the posterior half of the 
head. Ocelli developed. Scapes reaching the anterior border of the eyes. Proximal third 
of the scapes ca. 1/2 narrower than the distal parts. Mandibles massive and strongly 
convex dorsally. Masticatory margin of the mandibles with 4-5 irregular denticles 
followed by an apical tooth. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 637 

Mesosoma dorsally flat and 1/4 longer than the head (mandibles included). Pro- 
notum dorsally with the sides superficially marginate.Propleurae concave. Mesopleurae 
gently convex. Propodeum with the sides gently converging posteriorly. Basal and 
declivous faces of the propodeum subequal in size and delimited by a superficial margin. 

Petiole ca. 1/3 longer than broad,anteriorly truncate and dorsally gently convex. 
Ventral process of the petiole subround. Postpetiole subquadrate and broader pos- 
teriorly. Postpetiole in dorsal view antero-laterally angulate. Pygidium in side view 
truncate and postero-laterally concave. Pygidium in full face view the sides with a 
series of small denticles converging to a pair of large, pointed teeth separted by a deep 
notch over the sting. 

Legs. Femora not inflated. Tibiae strongly inflate. Hind basitarsi ca. 1/3 shorter 
than the maximum length of the hind tibiae. 

Wings as in Fig. 4. 

Sculpture. Head covered by thick, longitudinal striae, thicker on the anterior half 
of the ventral part, thinner on the scrobes and on the postero-lateral half of the ventral 
part of the head. Head with additional thin striae between the thick ones. Dorsum of the 
pronotum with ca.20 thick striae similar to those on the head dorsum, some striae 
separated by thin, bifurcated ones. Mesonotum medially with thin striae, fainter 
posteriorly; remaining parts of the mesonotum and scutellum simply smooth. Dorsum 
of the propodeum covered with about 24 striae thinner than those on the pronotum. 
Pleurae covered by thin, longitudinal striae, more impressed on the metapleurae. 
Petiolar dorsum with ca. 20 striae similar to those on the propodeum. Declivous face of 
the propodeum, anterior face of the petiole minutely reticulate-punctate. Postpetiole 
smooth, irregularly, minutelly and superficially punctate and with longitudinal striae, 
slightly sparsed, very thin and more impressed posteriorly. Center of the first gastric 
tergite with similar sculpture on the postpetiole, the striae thinner, sparser and fainter. 
Second gastric tergite with similar sculpture on the first tergite, the striae fainter. 
Remaining gastric tergites, sternites and legs punctate, denser on the two last sternites. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) long, erect to suberect, sparse on the head, on the mandibles, on the 
anterior border of the clypeus, on the mesosoma, on the pedicel and on the gaster, dense 
on the pygidium; (2) shorter than the type (1) and sparsely distributed on the whole 
body, dense on the sternites; (3) shorter than the type (2), subdecumbent to decumbent, 
very sparse on the whole body, dense on the tergites. In addition, the hypostomal bridge 
surrounded by a layer of hairs similar to the type ( 1 ) but appressed and apically curved. 

Colour black and shining. Anterior half of the head, antennae, mandibles, 
femora and pygidium ferrugineous, tarsi lighter. Proximal half of the last four funicular 
joints orange to light brown. Tibiae yellow. 

Measurements (in mm) and indices: TL 14.38; HL 2.18; HW 1.54; EL 0.64; SL 0.82; 
SW 0.31; WL 4.04; PeL 1.16; PeW 1.00; HFeL 1.16; HFeW 0.49; HTiL 1.00; HTiW 0.39; HBaL 
0.71; HBaW 0.14; CI 70.6; SI 37.8; HFel 42.2; HTil 39.0; HBal 19.7. 

Male (tentative attribution) (Fig. 26). Head slightly longer than broad. Vertexal 

margin convex. Ocelli protuberant. Compound eyes broadly convex and largely on the 

anterior part of the head. Frontal carinae with raised borders and partially covering the 



638 MARIA L. DE ANDRADE 

antennal socket. Borders of the frontal carinae subparallel anteriorly, slightly convex 
medially, and strongly converging, almost touching each other posteriorly. Frons of the 
frontal carinae concave anteriorly, raised medially and declivous posteriorly. Anterior 
border of the clypeus gently convex medially. Mandibles long; their masticatory margin 
edentated and with a pointed apical tooth. Scapes half longer than broad. Funicular 
joints thick. 

Mesosoma robust. Pronotum in dorsal view with subparallel sides. Mesonotum 
slightly convex. Pair of Mayrian carinae impressed but not connected each other pos- 
teriorly. Parapsidal furrows impressed. Basal face of the propodeum narrowing back- 
wards and separated from the declivous one by a marked carina. 

Petiole about 1/4 longer than broad, anteriorly truncate and dorsally convex. 
Ventral process of the petiole subtriangular. Postpetiole broadening backwards and 
narrower than the first gastric tergite. Postpetiole in dorsal view antero-laterally angu- 
late. First gastric segment broader than the postpetiole. Second gastric segment broader 
than the first segment. Remaining gastric segments narrowing backwards. 

Genitalia as in Fig. 27. 

Legs. Femora not inflated. Mid and hind basitarsi long. 

Wings as in Fig. 5. 

Sculpture. Head dorsum minutely punctate and with longitudinal, slightly 
irregular striate, the punctures more impressed on the anterior half, the striae more 
impressed on the posterior half and behind the the clypeus. Vertexal angles with addi- 
tional small, deep, piligerous foveae, continuing to the sides of the ventral part of the 
head. Middle of the ventral part of the head with thick tranversal rugae and piligerous 
foveae. Pronotum densely covered by deep, small piligerous foveae separated by thin, 
transversal striae. Mesonotum smooth and with sparse, minute piligerous punctures. 
Scutellum smooth. Basal face of the propodeum covered by thin, longitudinal striae. 
Petiolar dorsum smooth, with rare, superficial, small piligerous foveae and with short, 
longitudinal rugosities on the anterior part. Petiolar sides minutely reticulate and with 
sculpture similar to those on the anterior part of its dorsum, but sometimes with larger 
foveae and longer rugosities. Declivous face of the propodeum minutely and super- 
ficially punctate and with rugosities converging to the center. Pro- and mesopleurae 
smooth, with variably impressed punctuations and with traces of longitudinal rugosities, 
more impressed on the posterior border of the mesopleurae. Metapleurae striated as on 
the basal face of the propodeum. Postpetiole, first gastric segment and legs smooth and 
with superficial punctures, denser and deeper on the remaining gastric segments. 

Pilosity. Body covered by pointed hairs of four types: (1) long, sparse and 
suberect, denser on the last three gastric segments; (2) shorter than the type (1) and 
variably distributed on the body, dense on the mesopleurae; (3) mixed and shorter than 
the type (2), dense on the vertexal angles, on the posterior half of the ventral part of the 
head, on the pronotal dorsum, on the coxae, on the ventral face of the femora and tibiae, 
and on the gaster; (4) short and thick on the funicular joints. 

Colour. Black and shining. Anterior third of the head dorsum, mandibles, 
antennae, tibiae and tarsi ferrugineous to dark brown, femora darker. Outer face of the 
mid and of the hind tibiae, and tarsomeres yellowish to light brown. 



FOSSIL AND EXTANT SPECIES OF CYL/NDROMYRMEX 639 

Measurements (in mm) and indices: TL 10.1-10.7; HL 1.28-1.36; HW 1.20-1.26; EL 
0.66-0.76; SL 0.30-0.33; SW 0.16-0.17; WL 3.04-3.44; PeL 0.80-0.96; PeW 0.60-0.72; HFeL 
1.04-1.17; HFeW 0.23-0.27; HTiL 0.90-1.02; HTiW 0.20-0.23; HBaL 0.74-0.79; HBaW 0.07; CI 
92.6-93.7; SI 51.5-53.3; HFel 22.1-23.1; HTil 22.2-22.5; HBal 8.8-9.4. 

Material examined. COSTA RICA: Turrialba, 15-18.VII.1965, 1 male, P. J. Spangler 
[USNM]. PANAMA: Volcan de Chiriqui, 1 gyne (holotype), Champion [BMNH]. ECUADOR: 
Pichincha: Tinalandia, 16 km SE of S. Domingo de los Colorados, VI. 1975, 1 male, S. Peck & J. 
Peck [MCZC]. 

Discussion. C. godmani is the largest species of the genus. It is a rare species 
previously known only on the holotype and on a gyne from Ecuador (Wheeler 1924) 
not available for the present study. A striking character shared by godmani, antillanus 
and meinerti is a notch on the apex of the pygidium, more impressed in godmani and 
antillanus. The function of the notch is probably to facilitate the movement of the sting. 

The isolate males described here as godmani are tentatively referred to this 
species for the following reasons: 1- frontal carinae and genitalia similar to those of 
meinerti; 2- tibiae partially yellowish brown (yellowish in the gyne); 3- they originate 
from the geographic range of godmani; 4- the males of the other species occuring in 
Central and northermost countries of South America, i. e. striatus, whymperi, boliviae, 
brevitarsus are already known, except escobari. I exclude the possibility that the two 
males referred here to godmani could be attributed to the Colombian escobari because 
this species belongs to another clade the male of which {brevitarsus) differs signifi- 
cantly from the those of the longiceps clade by the frontal carinae and genitalia. If the 
attribution of these two males to godmani is not correct, they should represent an un- 
described species. 

Distribution. Costa Rica, Panama and Ecuador. 

Cylindromyrmex antillanus n. sp. Figs. 1 & 28 

Holotype: Winged gyne in the amber sample Do-4 130-1 from the SMNS. 

Derivatio nominis. The name antillanus is a neologism indicating the pro- 
venance of this amber sample from the Antilles. 

Diagnosis. A species resulting as outgroup of longiceps and meinerti, and 
differing from both for the CI > 77 (instead of < 70) and HFel < 46 (instead of > 50). 

Gyne (Figs. 1 & 28). Head slightly less than 1/3 longer than broad. Occiput low. 
Vertexal angles convex. Frontal carinae about 1/3 broad as the maximum head width. 
Sides of the frontal carinae parallel and reaching at least the middle of the eyes 
posteriorly. Dorsum of the frontal carinae with an impressed median sulcus anteriorly. 
Frontal carinae as long as the anterior border of the clypeus. Antero-median border of 
the clypeus with a minute pair of denticles. Compound eyes large, flat and on the 
middle of the head. Impar ocellus developed. Scapes reaching the anterior border of the 
eyes. Proximal third of the scapes 1/2 narrower than the distal parts. Mandibles gently 
convex dorsally. Masticatory margin of the mandibles each with a set of 4 irregular 
denticles followed by an apical tooth. 

Mesosoma slightly convex dorsally. Pronotum with parallel sides. Parapsidal 
furrows superficially impressed. Propodeum with the sides gently convex and conver- 
ging posteriorly. Basal face of the propodeum separated fromthe declivous one by a 
thin margin. 



640 



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FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 641 

Petiole subquadrate, slightly broader than long, anteriorly truncate and dorsally 
gently convex. Petiolar sides diverging backwards. Ventral process of the petiole large 
and subround. Postpetiolar sides gently diverging posteriorly. Postpetiolar sternite 
antero-medially with a marked, triangular "lip" pointing backwards. Pygidium in side 
view truncate. Posterior half of the pygidium in full dorsal view with the sides bearing a 
row of small denticles converging to a deep notch. 

Legs. Femora and tibiae slightly inflated. Hind basitarsi short, slightly less than 
1/3 shorter than the maximum length of the tibiae. Outer apical edge of the hind and of 
the mid basitarsi respectively with 3,5 spine-like setae. 

Wings as in Fig. 4. 

Sculpture. Head covered by thin, longitudinal striae, slightly thicker on the pos- 
terior third of the head dorsum. Mesosoma longitudinally striated. Dorsum of the pro- 
notum with about. 30 longitudinal striae similar to those on the posterior part of the 
head dorsum; some pronotal striae bifurcated. Mesonotum and propodeum coverd by 
longitudinal striae thinner than those on the pronotum. Scutellum, pleurae, declivous 
face of the propodeum, petiolar sides and ventral face of mid and hind femora with very 
thin, longitudinal striae, thinner on the scutellum, propleurae, petiolar sides and mid 
femora. Petiolar dorsum with about 28 striae similar to those on the propodeum. 
Anterior face of the petiole smooth. Dorsum of the postpetiole densely covered by 
striae as those on the petiole. First gastric tergite with thin, superficial, longitudinal 
striae on the center only. Second gastric sculptured as the first tergite but the striae 
extremely thin. Postpetiolar sternite, remaing gastric segments and legs smooth and 
with sparse punctures. Hind coxae covered by thin, longitudinal striae; mid coxae with 
similar sculpture but sparser and fainter. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) long, erect to suberect, one on the external border of the scape, a pair 
between the frontal carinae and clypeus, rare on the mandibles, on the mesosoma, on 
the gaster and on the legs, sparse on the pygidium; (2) shorter than the type (1) and 
sparsely distributed on the whole body; (3) shorter than the type (2), suberect on the 
head dorsum and mesosoma, subdecumbent on the pedicel, decumbet on the ventral 
part of the head, on the gaster and on the legs. In addition, the hypostomal bridge 
surrounded by a layer of hairs similar to the type (1) but appressed and apically curved. 

Colour dark brown. Tibiae yellowish to light brown. 

Measurements (in mm) and indices: TL 6.36; HL 1.08; HW 0.84; EL 0.35; SL 0.32; 
SW0.15; WL1.68; PeL 0.524 PeW 0.56; HFeL 0.53; HFeW 0.24; HTiL 0.44; HTiWO.16; 
HBaL0.25; HBaW 0.07: CI 77.8; SI 46.9; HFeI45.3; HTil 36.4; HBal 28.0. 

Material examined. Dominican amber: 1 gyne (reference number Do-4 130) [SMNS]. 

Discussion. Antillanus, longiceps and meinerti share the narrow frontal carinae, 
the eyes on the middle of the sides of the head, the mesosoma 2/3 longer than high, and 
the petiole with a short anterior face. The workers of longiceps and meinerti possess 
reduced and flat eyes. It is likely that the unknown worker of antillanus also has similar 
eyes. 

Distribution. Dominican amber. 



642 MARIA L. DE ANDRADE 

Cylindromyrmex longiceps André Figs 29-30 

Cylindromyrmex longiceps André, 1892: 47. Worker. Original description. Type locality: Brazil. 

Type material: 1 worker labelled: "Brésil, Type, Museum Paris, Collection Ernest André, 

1914, longiceps André", in MNHN, examined. 
Cylindromyrmex longiceps André, Kempf 1968: 372. Gyne. 

Diagnosis. Longiceps is the sister species of meinerti and differs from it in the 
worker and gyne by the narrower frontal carinae not reaching the anterior border of the 
clypeus. 

Worker (Fig. 29). Head about 1/3 longer than broad and with paralllel sides. 
Occiput very low. Vertexal angles round and protruding backwards. Frontal carinae 
slightly less than 1/3 broad as the maximum head width. Anterior fourth of the frontal 
carinae diverging backwards and not reaching the anterior border of the eyes pos- 
teriorly. Dorsum of the frontal carinae with a median sulcus anteriorly. Frontal carinae 
shorter than the anterior border of the clypeus. Antero-median border of the clypeus 
superficially notched and bearing a minute denticle. Compound eyes very small (Fig. 
30), flat and behind the mid line of the head. Ocelli reduced to a superficial pit, some 
specimens with the impair ocellus more developed than the pair ones. Scapes stout and 
short. Anterior fourth of the scapes half narrower than the distal parts. Mandibles short 
and flat dorsally. Masticatory margin of the mandibles edentated and with a pointed 
apical tooth. 

Mesosoma weakly convex dorsally and about 1/5 shorter than the head (man- 
dibles included). Sides of the mesosoma slightly narrower in the mesonotum. Propo- 
deum with the sides gently convex and converging posteriorly. Declivous face of the 
propodeum ca. 1/3 of the length of the basal face. Basal face separated from the decli- 
vous one by a very superficial margin. 

Petiole subquadrate. Anterior face of the petiole very short and deeply concave; 
dorsal face of the petiole weakly convex. Ventral process of the petiole small and 
subround. Postpetiole broader than long. Postpetiolar sides gently diverging posteriorly. 
Postpetiolar sternite antero-medially with a triangular "lip" pointing backwards. Pygi- 
dium truncate; its border with a semicircle of small teeth of similar size. 

Legs. Femora and tibiae inflated. Hind basitarsi slightly less than 1/2 shorter 
than the maximum length of the tibiae. Outer apical edge of the hind and of the mid 
basitarsi respectively with 5,6 spine-like setae. 

Sculpture. Head dorsum covered by thin longitudinal striae, more superficial and 
thinner close to the antennal scrobes. Ventral part of the head with small, superficial, 
oval piligerous foveae and with longitudinal striae, fainter on the posterior half, absent 
on the middle and on the posterior angles. Mesosoma with longitudinal striae thicker on 
the pronotum. Pronotum with 22-25 striae thicker than those on the posterior half of the 
head dorsum. Pleurae and petiolar sides with longitudinal striae similar to those on to 
the atennal scrobes. Petiolar dorsum with 24-26 striae similar to those on the pro- 
podeum. Declivous face of the propodeum and anterior face of the petiole minutely 

Fig. 29. C. longiceps André. Worker from Rio de Janeiro. Brazil. Head in full dorsal view (top), 
body in full dorsal view (middle), body in profile (bottom). 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



643 




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Fig. 30 
C. longiceps André. Worker from Rio de Janeiro, Brazil. Compound eye. 



punctate. Dorsum of the postpetiole and of the first gastric tergite covered by striae 
thinner than those on the petiole. Second gastric tergite with thin and superficial striae 
on the center only. Remaining gastric tergites and sternites sparsely and minutely 
reticulate and densely punctate. Legs with very superficial, minute punctures. Hind 
coxae covered by thin, longitudinal striae; mid coxae with similar sculpture but sparser 
and fainter. 

Pilosity. Body with pointed hairs of at least three lengths and distributed as 
follows: (1) long, erect to suberect, one pair on the clypeus, one close to each pronotal 
angle, rare on the on the gaster, sparse on the pygidium; (2) shorter than the type (1) 
and sparsely distributed on the whole body; (3) shorter than the type (2), sparse and 
suberect on the head dorsum and on the mesosoma, sparse and subdecumbent on the 
pedicel, and on the first gastric tergite, decumbent but sparse on the ventral part of the 
head and on the legs, dense on the postpetiolar and on the remaing gastric sternites. In 
addition the hypostomal bridge surrounded by a layer of hairs similar to the type ( 1 ) but 
appressed and apically curved. 

Colour black. Mandibles and anterior third of the head dark ferrugineous. 
Scapes, first funicular joints and tarsi brown. Legs orange to light brown. 



FOSSIL AND EXTANT SPECIES OF CYL1NDROMYRMEX 645 

Measurements (in mm) and indices: TL 7.44-8.50; HL 1.68-1.92; HW 1.08-1.28; EL 
0.15-0.22; SL 0.47-0.52; SW 0.20-0.22; WL 1.68-1.96; PeL 0.57-0.70; PeW 0.70-0.83; HFeL 
0.69-0.78; HFeW 0.33-0.38; HTiL 0.65-0.76; HTiW 0.25-0.29; HBaL 0.36-0.38; HBaW 0.10- 
0.1 1; CI 64.3-66.7; SI 42.3-42.5; HFel 47.8-48.7; HTil 38.1-38.5; HBal 27.8-28.9. 

Gyne. Very similar to the worker but differing from it in the following details: 
compound eyes very large, flat and on the middle of the dorsolateral part of the head; 
ocelli well defined and marked; mesosoma broad medially; parapsidal furrows super- 
ficially impressed; petiole slightly longer than broad; pronotum with about 28 striae; 
mesonotum and scutellum with very superficial, short, thin striae; postpetiolar striae as 
thick as on the pronotum. 

Wings as in Fig. 4. 

Measurements (in mm) and indices: TL 9.94; HL 1.84; HW 1.18; EL 0.54; SL 0.49; 
SW0.21; WL 2.76; PeL 0.74; PeW 0.76; HFeL 0.73; HFeW 0.38; HTiL 0.75; HTiW 0.27; HBaL 
0.44; HBaW 0. 1 1 ; CI 64. 1 ; SI 42.8; HFel 52.0; HTil 36.0; HBal 25.0. 

Material examined. BRAZIL: no further locality, 1 worker (holotype), MNHN. SÄO 
Paulo: Sâo Paulo, 5.1.1974, 1 gyne, R. L. Araujo [MZSP]. Rio de Janeiro: Rio de Janeiro, 
25.VIII.1962, 28 workers, R. L. Araujo [MZSP, NHMB]. 

Discussion. Longiceps is the species of the genus with the highest number of 
autapomorphies. They are the following: hypostomal bridge Y-shaped, broad and semi- 
transparent; head very elongate; frontal carinae very narrow; mandibles edentate; ante- 
rior border of the clypeus medially notched and denticulate; ventral process of the 
petiole very short; pygidium with a semicircle of small teeth. 

The largest known series of longiceps was collected by Araujo (a brazilian 
termitologist). It is very likely that all these specimens were collected in termite nests. 

Distribution. Brazil. 

Cylindromyrmex meinerti Forel Figs 31-34 

Cylindromyrmex meinerti Forel, 1905: 155. Worker. Original description. Type locality: Vene- 
zuela. Type material: 4 workers, two of which labelled: "C. meinerti, type, Forel, Las 
Trincheras, Venezuela, Meinert, in altem Baume", in MHNG, MCZC and MCSN; 
examined. 

Cylindromyrmex schmidti Menozzi, 1931: 192, fig. 4. Partim. Gyne. Nee worker (= whymperi). 
Original description. Type locality: Costa Rica. Type material: 2 gynes labelled: "La 
Caja: 8 kil. w. San José, C. R., Heinr. Schmidt", in IEGG, examined. Synonymia nova. 

Cylindromyrmex parallelus Santschi, 1932: 410, fig. 19. Gyne. Original description. Type 
locality: Panama. Type material: 1 gyne labelled: "Panama, France Field, Bierig, VI-30, 
Cylindromyrmex parallelus Sant. type", in NHMB, examined. Synonymia nova. 

Cylindromyrmex parallelus Santschi, Wheeler 1937: 443. Misidentification. 

Cylindromyrmex parallelus Santschi, Brown 1975: 38. Figs. 1 17 & 130, male genitalia. Misiden- 
tification. 

Diagnosis. Meinerti is the sister species of longiceps and differs from it in the 
worker and gyne by the frontal carinae as long as the anterior border of the clypeus 
instead of shorter. 

Worker (Fig. 31). Head ca. 1/4 longer than broad and with paralllel sides. 
Occiput very low. Vertexal angles round. Frontal carinae at most slightly broader than 
1/3 as the maximum head width. Anterior third of the frontal carinae diverging, 
remaining parts parallel and reaching the eyes posteriorly. Dorsum of the frontal carinae 



646 



MARIA L. DE ANDRADE 




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Fig. 31. C. meinerti Forel. Worker from Panama. Head in full dorsal view (top), body in full 
dorsal view (middle), body in profile (bottom). 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



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Fig. 32 

C. meinerti Forel. Worker from Panama. Anterior portion of the cephalic capsule and mandibles 
in ventral view to show the broad hypostomal bridge (character 6 state 1 ). Notice the convexity of 
the anterior margin of the hypostomal bridge, a character not verified in all species of the genus. 



with a median sulcus anteriorly. Frontal carinae as long as the clypeus. Anterior border 
of the clypeus laterally convex, medially concave and bearing a pair of small denticles. 
Compound eyes very small, flat and on the mid of the dorsolateral part of the head. 
Ocelli reduced to superficial pits, more developed in large specimens. Scapes not 
reaching the anterior border of the eyes. Proximal fourth of the scapes 1/2 narrower than 
the distal parts. Mandibles flat. Masticatory margin of the mandibles with 4 irregular 
denticles followed by an apical tooth. Hypostomal bridge broad, with the antero-lateral 
margin convex (Fig. 32). 

Mesosoma gently convex dorsally and slightly shorter than the head (mandibles 
included). Mesosoma 2/3 longer than heigh. Sides of the mesosoma parallel. Propodeal 
sides gently convex. Declivous face of the propodeum ca. 1/2 of the length of the basal 
face. Basal face of the propodeum separated from the declivous one by a faint margin. 

Petiole subquadrate. Petiolar sides diverging backwards. Anterior face of the 
petiole very short and concave; dorsal face of the petiole slightly convex. Ventral pro- 
cess of the petiole very large and subround. Postpetiole broader than long. Postpetiolar 
sternite antero-medially with traces of a triangular "lip" pointing backwards. Pygidium 



648 



MARIA L. DE ANDRADE 





00007972 



1 mm 



REM-Labor 
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FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



649 




0.5mm 



Fig. 34 

C. meinerti Forel. Male from Barro Colorado Is., Panama. Genital appendages: a) lateral view of 
left parameres; b) hypopygium; c) left aedeagus in profile; d) sternite VIII. 

truncate; its sides with a row of small teeth converging to a pair of larger teeth sepa- 
rated by a variably impressed notch over the sting. 

Legs. Femora and tibiae inflated. Hind basitarsi ca. 1/2 shorter than the 
maximum length of the tibiae. Outer apical edge of the hind and of the mid basitarsi 
with 3 spine-like setae. 



Fig. 33. C. meinerti Forel. Male from Barro Colorado Is., Panama. Head in full dorsal view (top), 
body in full dorsal view (middle), body in profile (bottom). 



650 MARIA L. DE ANDRADE 

Sculpture. Head dorsum covered by thin longitudinal striae, fainter and thinner 
on the antennal scrobes. Anterior half of the ventral part of the head with longitudinal 
striae as thick as those on the posterior part of the head dorsum but sparser; posterior 
half of the ventral part of the head with striae similar to those on the antennal scrobes. 
Mesosoma with longitudinal striae similar to those on the posterior part of the head 
dorsum. Pronotum with 20-21 striae. Pleurae and petiolar sides with longitudinal striae 
similar to those on the atennal scrobes. Petiolar dorsum with 17-19 striae similar to 
those on the mesosoma. Declivous face of the propodeum and anterior face of the 
petiole minutely punctate. Dorsum of the postpetiole covered by striae thinner than 
those on the petiole. First gastric tergite covered by striae thinner than those on the 
postpetiole. Second gastric tergite with thin and very superficial striae on the center 
only. Remaining gastric tergites and sternites sparsely and minutely reticulate and 
densely punctate. Legs with very superficial, minute punctures. Hind coxae covered by 
longitudinal striae; mid coxae with similar sculpture but fainter and sparser. 

Pilosity. Body with pointed hairs of at least of three lengths and distributed as 
follows: (1) long, erect to suberect, one pair between the frontal carinae and clypeus, 
one close to on each pronotai angle, rare on the on the gaster, sparse on the pygidium; 
(2) shorter than the type (1) and sparsely distributed on the whole body; (3) shorter than 
the type (2), sparse and suberect on the head dorsum and on the mesosoma, sparse and 
subdecumbent on the pedicel, and on the first gastric tergite, decumbent but sparse on 
the ventral part of the head and on the legs, dense on the postpetiolar and on the 
remaing gastric sternites. In addition, the hypostomal bridge surrounded by a layer of 
hairs similar to the type (1) but appressed and apically curved. 

Colour black. Mandibles and anterior third of the head dark ferrugineous. 
Antennae, coxae, femora, tarsi and tarsomeres light brown. Tibiae yellowish. 

Measurements (in mm) and indices: TL 5.32-6.58: HL 1.20-1.28: HW 0.88-0.94; EL 
0.11-0.15: SL 0.40-0.41: SW 0.17; WL 1.32-1.50; PeL 0.44-0.51; PeW 0.52-0.62; HFeL 0.53- 
0.60; HFeW 0.26-0.30; HTiL 0.44-0.49; HTiW 0.19-0.21; HBaL 0.22-0.25; HBaW 0.07-0.08; CI 
72.0-73.4; SI 41.5-42.5; HFel 49.0-50.1; HTÜ 41.7-43.2; HBal 30.4-32.0. 

Gyne. Very similar to the worker but differing from it in the following details: 
compound eyes very large; ocelli well defined; impar ocellus higher than the posterior 
border of the compound eyes; mesosoma broad medially; parapsidal furrows super- 
ficially impressed; petiole as broad, as long; pronotum with about 22-27 striae as thick 
as in the worker; mesonotum with thinner striae than on the pronotum; some specimens 
with striae only on the middle of the mesonotum; scutellum smooth or with striae on 
the anterior half only; propodeal striae thinner than on the pronotum. 

Wings as in Fig. 4. 

Measurements (in mm) and indices: TL 7.56-8.60; HL 1.24-1.44; HW 0.86-1.00; EL 
0.40-0.41; SL 0.41-0.42; SW 0.18; WL 1.96-2.24; PeL 0.71-0.72; PeW 0.70-0.72; HFeL 
0.56; HFeW 0.28-0.32; HTiL 0.48-0.56; HTiW 0.20-0.23; HBaL 0.26-0.31; HBaW 0.09-0.10; 
CI 69.3-70.0; SI 42.8-43.9; HFel 50.0-50.8; HTil 41.1-41.8; HBal 32.2-34.6. 

Male (Fig. 33) (previously undescribed). Head longer than broad. Vertexal 
margin convex. Ocelli protuberant. Compound eyes broadly convex and largely on the 
anterior part of the head. Frontal carinae with raised borders and partially covering the 
antennal socket. Sides of the frontal carinae subparallel anteriorly, slightly convex 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 65 1 

medially, and strongly converging and almost touching each other posteriorly. Frons 
concave anteriorly, raised medially and declivous posteriorly. Anterior border of the 
clypeus gently convex medially. Mandibles long; their masticatory margin edentated 
and with a pointed apical tooth. Scapes slightly less 1/2 longer than broad. Funicular 
joints stouts. 

Mesosoma robust. Pronotum in dorsal view with subparallel sides. Mesonotum 
slightly convex. Scutellum at the same level as the mesonotum. Pair Mayrian and 
parapsidal furrows superficially marked. Impar Mayrian furrow absent. Basal face of 
the propodeum separated from the declivous one by a developed and well marked 
carina. 

Petiole subcylindric; anteriorly truncate and dorsally convex. Ventral process of 
the petiole small and subtriangular. Postpetiole broadening backwards and smaller than 
the first gastric tergite. 

Genitalia as in Fig. 34. 

Legs. Femora not inflated. Mid and hind basitarsi long. 

Wings as in Fig. 5. 

Sculpture. Head dorsum covered striae converging from the internal border of 
the eyes to the ocelli; striae behind the pair ocelli thinner, tranversal, irregular and 
mixed with small piligerous foveae. Ventral part of the head variably punctate and with 
small, piligerous foveae; some specimens with diverging striae on the anterior part 
only. Pronotum punctate and with transversal, irregular striae, sometimes mixed with 
irregular piligerous foveae. Mesonotum and scutellum smooth and with minute punc- 
tures, denser on the mesonotum. Basal face of the propodeum, metapleurae and petiole 
covered by longitudinal striae. Declivous face of the propodeum smooth; some speci- 
mens with tranversal striae on the middle of the posterior half only. Propleurae punctate 
and with traces of thin, longitudinal striae. Mesopleurae smooth, minutely punctate and 
with rugosities on the posterior border. Postpetiole, first gastric segment and legs 
smooth and with sparse, superficial punctures; some specimens with longitudinal, irre- 
gular rugosities on the postpetiole. Remaining gastric segments superficially reticulate- 
punctate; this sculpture more impressed posteriorly. 

Pilosity. Body covered by pointed hairs of thrre types: (1) long, sparse, subde- 
cumbent, denser on the gaster, rare on the head; (2) shorter than the type (1), sparse on 
the head and legs, dense on the mesosoma and gaster; (3) short and thick on the 
funicullus. 

Colour. Head, mesosoma and petiole black. Anterior third of the head dorsum, 
mandibles, antennae and legs yellowish to light brown. Postpetiole, first and second 
gastric segments dark brown, remaing gastric segments lighter. 

Measurements (in mm) and indices: TL 7.90-8.30; HL 1.08-1.16; HW 0.92-1.04: EL 
0.56-0.59; SL 0.22; SW 0.13; WL 2.48-2.64; PeL 0.70-0.72; PeW 0.64-0.68; HFeL 0.77-0.84; 
HFeW 0.20-0.24; HTiL 0.68-0.82; HTiW 0.18-0.19; HBaL 0.52-0.61; HBaW 0.07-0.08; CI 85.2- 
89.6; SI 59.1; HFel 26.0-28.6; HTil 23.2-26.5; HBal 13.1-13.5. 



652 



MARIA L. DE ANDRADE 



Material examined. COSTA RICA: no further locality, 1 gyne, F. Nevermann [MZSP]; 
no further locality, 1 gyne, 1920, P. Serre [MNHN]; La Caja, 8 km W of San José, 2 gynes 
(corresponding to the description and drawing of Menozzi, 1931), H. Schmidt [IEGG]; Santa 
Rosa, Natural Park, Guanacaste Province, May-August 1984, 300 m, 1 gyne, D. H. Janzen & I. 
Gauld [BMNH]; Hambrug Farm, Santa Clara Province, 23. IV. 1926, 1 worker, 2 gynes, 2 males, 
F. Nevermann [USNM]; same locality, Reventazon River, 1 worker, F. Nevermann [USNM]; 
same locality, IV. 1921, 4 gynes, 3 males, 1 pupa, F. Nevermann [USNM]. PANAMA: France 
Field, VI.1930, 1 gyne (holotype of parallelus), A. Bierig [NHMB]; Barro Colorado Is. Canal 
Zone, 9. VI. 1935, 1 gyne, 3 gynes pupae, A. Emerson [MCZC]; same locality, 4. V. 1935, 4 males, 
A. Emerson [MCZC, USNM]: same locality, 2 workers, in termite nest, L. Schneider [IAVH, 
MZSP]; same locality, 1 worker, in termite nest, [WEMC]; same locality, III.IV.1949, Zetek, 1 
male [USNM]. VENEZUELA: Zulia: El Tucuco 45 km SW of Machiques, 5-6.VI.1976, 1 male, 
A. S. Menke & D. Vincent [WEMC]; same locality, IV. 1984, 1 male, E. Inciarte & E. Rubio 
[MIZA]. Distrito Federal: Los Canales. 120 m. 23.III.1938, 1 male, G. Vivas-Berthier 
[WEMC]. Bolivar: Las Trincheras, in altem Baume. 4 workers (syntypes of in e inerti), Meinert 
[MHNG, MCZC, MCSN]. BRAZIL: Amazonas: Ilha de Curari, vârzea, 2.LX.1976, 1 gyne, J. 
Adis [LACM]. 




Fig. 35 

Unique most parsimonious phylogeny of the known species of Cylindromyrmex. Acanthostichus 
texanus and Simopone annettae have been included into the analysis for outgroup comparison. 
The frames include the character changes at each branch with their respective identification 
number as given in the text and the apomorphic state change. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



653 



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92 



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Fig. 36 

Same phylogeny as in Fig. 35 with the frequency of the clades resulting from 1,000 bootstrap 
replicates. Further explanations in text. 



654 MARIA L. DE ANDRADE 

Discussion. Meinerti, from my cladistic analisys, results as the sister species of 
longiceps although it resembles more antillanus in body shape. This is because longi- 
ceps and meinerti share synapomorphically the HFel > 50 and the broad hypostomal 
bridge (Fig. 32), the first of which is of doubtful phylogenetic importance. 

The composite nature of the type series of schmidti Menozzi has been already 
described under whymperi. 

Forel (1905) reported meinerti from an old tree. Some specimens of meinerti 
were collected in termite nests (see material examined). 

Distribution. Costa Rica, Panama, Venezuela, Brazil. 

DISCUSSION 

The nesting place and feeding habits of Cylindromyrmex are still fragmentarily 
known. Part of the specimens examined during this study bear some biological 
information (see the discussion under each Recent species). Wheeler (1936) listed 
brasiliensis and "williamsi" (=whymperi) as termite inquilines. Overal & Bandeira 
(1985) equally supposed that specimens of striatus collected in a termite nest should be 
termite inquilines. Their statement is partly contradicted by their report in the same 
paper of striatus workers attacking Nasutitermes surinamensis in laboratory. 

Tomotake & Caetano (1997) described the digestive tract of C. brasiliensis 
and compare it with the one of the same subfamily Acanthostichus serratulus. 
Significant differences have been found neither between the two genera nor between 
the two Cerapachyinae and other ants. 

Clark et al. (1982) reported williamsi (- whymperi) as "endemic" in the North 
arid zone of Santa Cruz (Galapagos Is.). They collected "williamsi" in two out of 429 
samples and mention that "williamsi" has escaped the competition with Wasmannia 
because of its adaptation to inhabit the arid zone. Lubin (1984) uses the term native for 
"striatus" (=whymperi) from the Galapagos. 

The distribution of the members of the striatus, boliviae, brevitarsus and longi- 
ceps clades are given respectively in Figs. 37-40. The figures are based only on the 
material examined during this study. Previous literature records I have been able to 
verify revealed often erroneous identifications. 

The three species of the striatus clade appear to be allopatric (Fig. 37). The 
unique male of whymperi from Blumenau in NHMW is likely to be wrongly labelled. 
The allopatry of brasiliensis and striatus had been already stressed by Fowler & 
Delabie(1995). 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



655 




it brasiliensis 
■ striatus 
▲ whymperi 



^&^ 



Fig. 37 
Distribution of the species of the striatus clade. 



656 



MARIA L. DE ANDRADE 




FlG. 38 

Distribution of the species of the boliviae clade. 



FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 



657 




Fig. 39 
Distribution of the species of the brevitarsus clade. 



658 



MARIA L. DE ANDRADE 



?-^^ 



V: . 



^) 



H» antillanus 
▲ godmani 
"^ longiceps 
■ meinerti 



■«8 



43P 



Fig. 40 
Distribution of the species of the longiceps clade. 



FOSSIL AND EXTANT SPECIES OF CYL1NDROMYRMEX 659 

IDENTIFICATION KEYS 

WORKERS 

The workers of antillanus, boliviae, electrinus and godmani are not included in 
this key because they are not yet known. 

1. Eyes large and convex (> 400 ommatidia). Ocelli present and well 

defined (Figs. 6, 8, 12) 2 

Eyes small or of medium size (> 16 and < 200 ommatidia), flat or 
slightly convex. Ocelli absent or represented by superficial pits (Figs 17, 
19,29) 3 

2. Legs dark orange to light brown. Dorsum of the petiole with thin, 

irregular striae (Fig. 6). Brazil, Paraguay brasiliensis 

Legs dark ferrugineous to black, with at least part of the tibiae yellowish. 
Dorsum of the petiole with thick, regular striae (Figs 8, 12) 4 

3. Posterior third of the head dorsum at most with 25 striae. Postpetiole 
with 19-25 longitudinal striae. Guatemala, Costa Rica, Galapagos Is., 

Ecuador, Peru, Bolivia, Chile, Brazil whymperi 

Posterior third of the head dorsum with more than 34 striae. Postpetiole 

with about 29-30 striae. Surinam, French Guyana, Brazil striaius 

4. Head length (mandibles excluded) ca. 1/3 longer than broad; frons, 
slightly more than 1/3 of the head width. Mandibles dorsally flat 

(Figs 29, 31) 5 

Head length (mandibles excluded) ca. 1/5 longer than broad; frons broad, 
slightly less than 1/2 or more than 1/2 of the head width. Mandibles 
dorsally convex (Figs. 17, 19, 20) 6 

5. Frontal carinae not reaching the anterior clypeal border. Pygidium with a 

semicircle of teeth of similar size. Brazil longiceps 

Frontal carinae reaching the anterior clypeal border. Pygidium with a 
semicircle of teeth with two larger ones over the sting. Costa Rica, 
Panama, Venezuela, Brazil meinerti 

6. Frontal carinae not reaching the anterior clypeal border. Clypeus strongly 
convex medially (Fig. 17). Gaster without striation. Mesosoma, petiole 

and legs elongate. Hind femora Index (HFel) = 37. Colombia escobari 

Frontal carinae reaching or surpassing the anterior clypeal border. Cly- 
peus not strongly convex medially (Fig. 19, 20). At least the first gastric 
tergite with thin striation on the anterior part. Mesosoma, petiole and legs 
stout. Hind femora Index (HFel) > 45 7 

7. Frontal carinae surpassing the anterior clypeal border. Mandibles with 9- 

10 denticles. Scape Index (SI) = 37. Cuba darlingtoni 

Frontal carinae reaching the anterior clypeal border. Mandibles with 6-7 
denticles. Scape Index (SI) > 42. Venezuela, Ecuador, Peru and Brazil 
brevitarsus 



660 MARIA L. DE ANDRADE 



GYNES 

The gyne of escobari is not included in this key because it is not yet known. 

1 . First gastric tergite smooth 2 

First gastric tergite sculptured 5 

2. Postpetiole smooth or with very thin, short, superficial striae on the pos- 
terior half. Frontal carinae very broad, reaching the internal border of the 
eyes (Fig. 14). Mandibles not angulate basally, convex dorsally and with 

10-12 denticles. Colombia, Venezuela, Peru and Bolivia boliviae 

Postpetiole entirely striate. Frontal carinae not reaching the internal 
border of the eyes. Mandibles angulate basally, slightly convex or flat 
dorsally, with maximum 7 denticles 3 

3. Legs dark yellowish-orange to light brown. Body striation more irre- 
gular. Brazil, Paraguay brasiliensis 

Legs dark ferrugineous to black with large part of the tibiae yellowish. 
Body striation regular 4 

4. Cephalic Index (CI) > 80. Posterior third of the head dorsum at most with 
25 striae. Guatemala, Costa Rica, Galapagos Island, Ecuador, Peru, 

Bolivia, Chile, Brazil whymperi 

Cephalic Index (CI) < 77. Posterior third of the head dorsum with more 

than 34 striae. Surinam, French Guyana, Brazil striatus 

5. Frons at most slightly more than 1/3 the head width. Eyes on the middle 

of the head sides 6 

Frons at least slightly less than 1/2 or more than 1/2 of the head. Eyes 
behind the middle of the head sides 8 

6. Frontal carinae not reaching the anterior clypeal border. Pygidium api- 
cally without a distinct pair of large teeth. Distal border of hind basitarsi 

with 5 spine-like setae. Brazil longiceps 

Frontal carinae reaching the anterior clypeal border. Pygidium apically 
with a distinct pair of large teeth. Distal border of hind basitarsi with 3 
spine-like setae 7 

7. Outer apical edge of the mid basitarsi with 3 spine-like setae on the outer 
face. Cephalic Index (CI) < 70. HFel > 50. Costa Rica, Panama, 

Venezuela, Brazil meinerii 

Outer apical edge of mid basitarsi with 5 spine-like setae on the outer 

face. Cephalic Index (CI) > 77. HFel < 46. Dominican amber antillanus 

8. Head and mesosoma covered by thick and thin striae. Pygidium apically 
with a distinct pair of large teeth separated by a deep notch. Size large > 

12.5 mm. Cephalic Index (CI) < 71. Costa Rica, Panama, Ecuador . . godmani 
Head and mesosoma covered by uniform thin striae. Pygidium apically 
with 4-6 large teeth not separated by a notch. Size small < 9.5 mm. 
Cephalic Index (CI) > 78 9 

9. Coxae and femora black. Mid basitarsi with parallel sides and more than 

half longer than the hind basitarsi. Dominican amber electrinus 



FOSSIL AND EXTANT SPECIES OF CYL1NDROMYRMEX 66 1 

Coxae, femora dark brown. Mid basitarsi broad apically and half as long 

as the hind basitarsi 10 

10. Frontal carinae not surpassing the anterior clypeal border. Mandibles 
with 6-8 small denticles. Femora yellowish to light brown. Venezuela, 

Ecuador, Peru and Brazil brevitarsus 

Frontal carinae surpassing the anterior clypeal border. Mandibles with 9-10 
small denticles. Femora dark brown. Cuba darlingtoni 

MALES 

The males of darlingtoni, escobari, electrinus, antillanus and longiceps are not 
considered because they are not yet known. Whymperi and striatus are not separated 
because the unique specimen of striatus available for the present study is immature and 
does not allow a sure recognition of diagnostic characters. 

1 . Frontal carinae strongly converging and almost touching each other pos- 
teriorly and broadly separated anteriorly (Figs 26, 33). Hypopygium with 
a simple, umpair, median projection between the apodemes (Figs 

27b, 34b) 2 

Frontal carinae not strongly converging posteriorly (Figs 10, 15, 22), if 
almost touching each other posteriorly (few males of brevitarsus) then 
never broadly separated anteriorly. Hypopygium smooth, or finely 
denticulate, or with a bidentate median projection between the apodemes 
(Figs. 7b, 1 lb, 23b) 3 

2. Total length (TL) > 9.5 cm. Mesosoma massive. Petiole with traces of 
striae only anteriorly. Postpetiole smooth. Costa Rica, Panama, Ecuador 

godmani 

Total length (TL) < 8.5 cm. Mesosoma elongate. Petiole entirely striate. 
Postpetiole superficially striate. Costa Rica, Panama, Venezuela, Brazil . 
meinerti 

3. Head and basal face of the propodeum with thick striae (Fig. 10), 
sometimes with thick foveae between the striae. Hypopygium smooth or 

finely denticulate between the distal apodemes 4 

Head and basal face of the propodeum with thinner striae (Figs 15, 22). 
Hypopygium with a simple, umpair, median projection between the apo- 
demes 5 

4. Anterior clypeal border slightly convex. Scape Index (SI) < 53.1. Hypo- 
pygium finely denticulate between the distal apodemes (Fig. 7b). Brazil 

and Paraguay brasiliensis 

Anterior clypeal border straight (Fig. 10). Scape Index (SI) > 57.1. 
Hypopygium smooth between the distal apodemes. (Figs lib, 13b). 
Guatemala, Costa Rica, Galapagos Island, Ecuador, Peru, Bolivia, Chile, 

Brazil whymperi 

Surinam, French Guyana, Brazil striatus 



662 MARIA L. DE ANDRADE 

5. Coxae black with the remaining parts of the legs yellow to light brown. 
Ventral border of the aedeagus with at least 42 denticles. Venezuela, 

Ecuador, Peru and Brazil brevitarsus 

Coxae dark or black, concolour with the remaining parts of the legs. 
Ventral border of the aedeagus with at most 32 denticles. Colombia, 
Venezuela, Peru and Bolivia boliviae 

ACKNOWLEDGEMENTS 

I would like to express my warmest thanks to Cesare Baroni Urbani for the 
suggestions and help through all the stages of the work. I am grateful to all the curators 
who send material for this study. In particular, many thanks are due to Richard 
Guggenheim for the facilities and to Sabine Wirtz for the photographs at the Centre of 
Scanning Electron Microscopy of the Basel University, to Carlos Roberto Ferreira 
Brandäo and to Bodo Hasso Dietz for sending me pictures of some questionable spe- 
cimens. 

Publication of the present paper has been made possible through a financial 
contribution of the Emilia Guggenheim-Schnurr Foundation of Basel. 

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Revue suisse de Zoologie 105 (3): 665-732; septembre 1998 



Aleocharinae della Cina: Parte III 
(Coleoptera, Staphylinidae) 

Roberto PACE 

Via Vittorio Veneto 13, 1-37032 Monteforte d'Alpone (Verona), Italia. 

Aleocharinae from China: Part HI (Coleoptera, Staphylinidae). - In 

this paper further 69 species are described as new to science. These new 
species belong to the tribe Athetini (part II). Two new synonymies are 
proposed. The main diagnostic caracters are illustrated. 

Key-words: Coleoptera - Staphylinidae - Aleocharinae - Taxonomy - 
China. 



INTRODUZIONE 

Con il presente lavoro continua la descrizione delle specie nuove delle Aleo- 
charinae della Cina raccolte dal Dr Ales Smetana e dal collega Guillaume de 
Rougemont di Londra. Le specie note o nuove per la Cina sono elencate nella prima 
parte di questa serie (Pace 1998a). Le specie descritte nel presente lavoro apparten- 
gono alla vasta tribù degli Athetini la cui trattazione sarà conclusa nella quarta parte 
di questa serie. 

Gli holotypi delle nuove specie sono conservati nelle collezioni del Museo di 
Storia Naturale di Ginevra, Svizzera (MHNG). 

ATHETINI (parte II) 

Atheta (Chaetida) elephanticola sp. n. Figg. 1-3 

Holotypus <5, China, Yunnan, Xishuangbanna, Sanchahe, elephant res., 24.1.1993, de 
Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e nero con elitre giallo-brune; 
antenne nere; zampe anteriori e medie giallo-brune, posteriori pure giallo-brune, ma 
con tarsi gialli. Sulla superficie corporea non vi è traccia di reticolazione. La 
punteggiatura del capo è distinta. I tubercoletti che coprono la superficie del pronoto e 
dell'addome sono salienti, quelli delle elitre sono svaniti. Il pronoto ha un debole e 
stretto solco mediano. 

Comparazioni. La nuova specie è simile ad A. drescheri Cameron, 1939, 
dell'India. Se ne distingue per il colore giallo-bruno delle elitre e l'estremità addo- 



(141° Contributo alla conoscenza delle Aleocharinae). 
Manoscritto accettato il 13.02.1998 



666 ROBERTO PACE 

minale nera e non rossiccia come in drescheri. L'estremità delle antenne dell' holo- 
typus â di drescheri è perduta, perciò non comparabile. L'edeago della nuova specie 
è un terzo più breve, nonostante ciò è più largo nella regione mediana, se visto 
ventralmente. Inoltre la lunga armatura genitale interna dell'edeago stesso corris- 
ponde a un'armatura genitale corta e sinuata nell'edeago di drescheri. 

Atheta (Philhygra) conferta sp. n. Figg. 7-9 

Holotypus ó\ China, Yunnan, Dali, 9.II.1993, de Rougemont leg. (MHNG). 
Paratypi: 2 6 e 1 9, stessa provenienza. 

Descrizione. Lunghezza 2,8 mm. Corpo lucido e nero-bruno; antenne brune 
con l'antennomero basale bruno-rossiccio; zampe bruno-rossicce. La reticolazione 
della superficie del capo e del pronoto è nettissima, quella delle elitre è estremamente 
svanita e quella dell'addome è distinta e a maglie trasverse. Il capo e il pronoto non 
presentano distinta punteggiatura. Tubercoletti fini ed estremamente svaniti coprono 
le elitre. Uroterghi con tubercoletti distinti. Edeago figg. 8-9, spermateca indistinta. 

Comparazioni. La nuova specie è affine ad A. homoeopyga Eppelsheim, 1893 
della Siberia orientale (Baikal), se si osserva l'edeago. Se ne distingue per l'apice 
dell'edeago largamente ricurvo al lato ventrale, mentre homoeopyga ha l'apice 
dell'edeago strettamente ricurvo. Lo stesso, in visione ventrale è stretto nella nuova 
specie e largo in homoeopyga. L'armatura genitale interna dell'edeago della nuova 
specie è composta da un fascio di robuste spine che nell'edeago di homoeopyga sono 
esili e in numero molto minore. Inoltre nell'edeago della nuova specie esiste una lama 
ricurva appartenente all'armatura genitale interna, assente nell'edeago di homoeo- 
pyga. 

Atheta (Philhygra) dalijiensis sp. n. Figg- 10-12 

Holotypus S, China, Gansu, Dalijia Shan, 46 Km W Linxia, 2980 m, 10.VII.1994, 
A. Smetana leg. (MHNG). 

Descrizione. Lunghezza 3,9 mm. Corpo lucido e nero pece con addome nero; 
antenne nero-brune; zampe rossicce. La reticolazione della superficie dell'avancorpo 
è netta, quella degli uroterghi libero 4°, 5° e 6° è estremamente trasversa e distinta. I 
tubercoletti che coprono la superficie del capo sono distinti e assenti sulla fascia 
mediana, quelli del pronoto e delle elitre sono svaniti. Edeago figg. 1 1-12. 

Comparazioni. Per la forma dell'edeago, la nuova specie è tassonomicamente 
vicina ad A. terminalis (Gravenhorst, 1806) della regione paleartica occidentale. La 
nuova specie se ne distingue per avere l'estremità dell'edeago non dentata, sottile e 
molto prolungata (estremità dell'edeago breve e larga in terminalis). 

Atheta (Philhygra) gansuicola sp. n. Figg- 13-15 

Holotypus â, China, Gansu, M. ts 25 Km E Xiahe, 3000 m, 5.VIII.1994, A. Smetana 
leg. (MHNG). 

Paratypi: 2 <3\ stessa provenienza. 



ALEOCHARINAE DELLA CINA 



667 




FlGG. 1-7 

Habitus, edeago in visione laterale e ventrale. 1-3: Atheta (Chaetida) elephanticola sp. n.; 4-6: 
Atheta (Philhygra) yokkaichiana Bernhauer; 7: Atheta (Philhygra) corifena sp. n. 



668 



ROBERTO PACE 




Figo. 8-12 
Edeago in visione laterale e ventrale e habitus. 8-9: Atheta (Philhygra) conferta sp. n.; 10-12: 
Atheta (Philhygra) dalijiensis sp. n. 



ALEOCHARINAE DELLA CINA 669 

Descrizione. Lunghezza 3,7 mm. Corpo lucido e nero; antenne brune; zampe 
giallo-brune. La reticolazione del capo è nettissima, quella del pronoto è vigorosa, 
quella delle elitre è netta e quella dell'addome è svanita e a maglie molto trasverse. Il 
capo presenta una debole e larga bozza tra le antenne e tubercoletti salienti e assenti 
sulla fascia mediana. I tubercoletti del pronoto sono salienti, quelli delle elitre svaniti. 
Edeago figg. 14-15. 

Comparazioni. La nuova specie, simile per il capo e il pronoto nettamente 
reticolati ad A. pseudoelongatula Bernhauer, 1907 del Giappone, ne è distinta per 
avere l' edeago sinuato al lato ventrale (arcuato in pseudoelongatula) e la parte termi- 
nale dello stesso organo larga e non stretta come in pseudoelongatula. 

Atheta (Philhygra) tianmushanensis sp. n. Figg- 16-19 

Holotypus â , China, Zhejiang Tianmushan, 29. IV. 1993, de Rougemont leg. (MHNG). 
Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 3,9 mm. Corpo lucido e bruno con elitre bruno- 
rossicce e addome nero; antenne brune; zampe rossicce con femori bruno-rossicci. La 
reticolazione del pronoto è netta e quella sul resto della superficie del corpo è distinta. 
I tubercoletti della superficie del capo sono svaniti, assenti sulla fascia mediana, quelli 
del pronoto sono netti e quelli delle elitre e dell'addome sono distinti. Spermateca fig. 
16, edeago figg. 17-18. 

Comparazioni. La nuova specie è ben distinta da A. palustris Kiesenwetter, 
1844, a diffusione paleartica, per il capo e il pronoto distintamente o nettamente reti- 
colati (assenza di reticolazione in palustris). L'edeago della nuova specie pur avendo 
profilo ventrale simile a quello dell'edeago di palustris, ha l'armatura genitale interna 
molto differente. Ad esempio le due armature genitali dentate dell'edeago di palustris 
sono assenti neh' edeago della nuova specie. 

Atheta (Philhygra) ideogrammifera sp. n. Figg 20-22 

Holotypus ó\ China, Shanxi, Wutaishan, 4-5.VI.1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 3,2 mm. Corpo lucido e nero-bruno con elitre giallo- 
brune e il margine posteriore dei due uriti basali bruno-rossiccio; antenne brune con i 
due antennomeri basali rossicci; zampe rossicce. La reticolazione della superficie del 
capo è quasi vigorosa, quella del pronoto è distinta, quella delle elitre è netta e quella 
dell'addome è a maglie molto trasverse e ondulate, distinte. La punteggiatura del capo 
è distinta e assente sulla fascia mediana. I tubercoletti che coprono il pronoto sono 
ben svaniti, quelli delle elitre sono salienti. Edeago figg. 21-22. 

Comparazioni. Il profilo ventrale dell'edeago della nuova specie è simile a 
quello dell'edeago di A. pseudoelongatula Bernhauer, 1907, del Giappone, ma la 
robusta armatura genitale interna dell'edeago della nuova specie è assente neh' edeago 
di pseudoelongatula. Inoltre l'edeago della nuova specie, in visione ventrale, è 
strettissimo, mentre è largo in pseudoelongatula. 



670 



ROBERTO PACE 




Figo. 13-19 
Habitus, edeago in visione laterale e ventrale e spermateca. 13-15: Atheta (Philhygra) gansui- 
cola sp. n.; 16-19: Atheta (Philhygra) tianmushanensis sp. n. 



ALEOCHARINAE DELLA CINA 



671 




FlGG. 20-25 

Habitus ed edeago in visione laterale e ventrale. 20-22: Atheta (Philhygra) ideogrammi/era 
sp. n.; 23-25: Atheta (Philhygra) vulnerans sp. n. 



672 ROBERTO PACE 

Etimologia. Il nome della nuova specie significa "Portatrice di ideogramma". 
Infatti l'armatura genitale dell'edeago in visione laterale ha la vaga forma di un 
ideogramma cinese. 

Atheta (Philhygra) vulnerans sp. n. Figg. 23-25 

Holotypus â , China, Yunnan, Ruili, ca. 700 m, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 3,1 mm. Corpo lucido e nero-bruno con elitre giallo- 
brune, con margine posteriore dei tre uroterghi basali giallo-brunicci e con gli uriti 
liberi 4° e la base del 5° neri; antenne brune con l'antennomero basale giallo-rossiccio 
e il successivo bruno rossiccio; zampe giallo-rossicce. La reticolazione dell'avan- 
corpo è netta, quella dell'addome è svanita e a maglie molto trasverse. La punteggia- 
tura del capo è estremamente svanita. Il pronoto è coperto di tubercoletti svaniti, le 
elitre di tubercoletti distinti. Edeago figg. 24-25. 

Comparazioni. In visione laterale, l'armatura genitale dell'edeago della nuova 
specie possiede due distinte lame ricurve che nell' edeago di A. pseudoelongatula 
Bernhauer, 1907, non sono presenti. 

Etimologia. Data la presenza di lame dell'armatura genitale interna dell'ede- 
ago, la nuova specie è nominata "colei che ferisce". 

Atheta (Psammostiba) apicipilae sp. n. Figg 26-28 

Holotypus <?, China, Shanxi, Wutaishan, 4-5.VI.1993, de Rougemont leg. (MHNG). 
Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 3,3 mm. Avancorpo debolmente lucido, addome 
lucido. Corpo nero-bruno con addome nero; antenne nero-brune; zampe rossicce con 
femori nero-bruni. Una netta reticolazione copre la superficie del corpo. Il capo ha il 
disco impresso e la punteggiatura distinta, assente sulla fascia mediana. I tubercoletti 
del pronoto sono finissimi e radi, quelli del pronoto sono netti. Edeago figg. 27-28, 
spermateca indistinta. 

Comparazioni. La nuova specie, in base alla forma dell'edeago è affine ad 
A. kamtschatica Brundin, 1943, della Siberia. Se ne distingue per avere l'edeago più 
ampiamente ricurvo al lato ventrale e con profilo sinuato (non sinuato in kamtscha- 
tica), per la parte apicale dell'edeago, in visione ventrale, più larga, con appendice 
membranosa a ciascun lato e per la presenta di setoline nell'armatura genitale interna 
dell'edeago (assenti in kamtschatica). 

Etimologia. La nuova specie prende nome dai peli dell'armatura genitale 
interna dell'edeago. Il nome significa "Peli apicali". 

Atheta (Coprothassa) iucunda sp. n. Figg- 29-33 

Holotypus o\ China, Yunnan, Kunming, I-II.1993, de Rougemont leg. (MHNG). 
Paratypi: 2 9 , stessa pronenienza. 

Descrizione. Lunghezza 3,2 mm. Corpo lucidissimo e nero con elitre nero- 
brune; antenne nere con i due antennomeri basali nero-bruni; zampe bruno-rossicce 



ALEOCHARINAE DELLA CINA 



673 




Figo. 26-32 

Habitus, edeago in visione laterale e ventrale e spermateca. 26-28: Atheta (Psammostiba) 
apicipilae sp. n.; 29-32: Atheta (Coprothassa) iucunda sp. n. 



674 ROBERTO PACE 

con tarsi rossicci. Sulla superficie corporea non vi è traccia di reticolazione, tranne 
che sul quinto urotergo libero dove è visibile una reticolazione estremamente svanita 
e a maglie molto trasverse. La punteggiatura del capo e del pronoto è svanita, quella 
dell'addome è netta. Tubercoletti svaniti coprono la superficie delle elitre. Edeago 
figg. 30-31, spermateca fig. 32, sesto urotergo libero del maschio fig. 33. 

Comparazioni. In base alla forma della spermateca la nuova specie sembra 
affine ad A. melanaria (Mannerheim, 1830), diffusa nella regione paleartica occi- 
dentale. Tuttavia la spermateca della nuova specie ha il bulbo distale molto più 
sviluppato e la parte prossimale dello stesso organo, a stretta spira e non con due 
spire, di cui una ampia come in melanaria. Inoltre l'edeago della nuova specie è 
profondamente arcuato al lato ventrale, mentre l'edeago di melanaria è debolmente 
arcuato. 

Atheta (Coprothassa) hsin sp. n. Figg- 34-35 

Holotypus 9, China, Sichuan, Langmui, 3500-3600 m, 13.VII.1994, A. Smetana leg. 
(MHNG). 

Descrizione. Lunghezza 2,3 mm. Corpo lucido e bruno; antenne brune con i 
due antennomeri basali bruno-rossicci; zampe giallo-brune. La reticolazione del 
pronoto è netta, quella sul resto del corpo è distinta, trasversa e ondulata sull'addome. 
I tubercoletti che coprono la superficie del corpo sono salienti. Spermateca fig. 34-35. 

Comparazioni. La nuova specie è ben distinta da A. melanaria (Mannerheim, 
1830), per avere il bulbo distale della spermateca molto sviluppato e la parte 
prossimale dello stesso organo a spirale schiacciata. 

Etimologia. La nuova specie prende nome dall'aggettivo cinese "hsin" che 
significa nuova. 

Atheta (Acrotona) setipyga sp. n. Figg- 36-38 

Holotypus S, China, Yunnan, Ruili, ca. 700 m, 3.II.1993, de Rougemont leg. (MHNG). 
Paratypus: 1 S , stessa provenienza. 

Descrizione. Lunghezza 1,8 mm. Corpo lucido e bruno con pronoto bruno- 
rossiccio e con i due uriti basali e l'apice dell'addome giallo-bruni; antenne brune con 
il primo antennomero basale giallo-rossiccio con una macchia apicale bruna; zampe 
giallo-rossicce. Il capo e il pronoto sono privi di reticolazione, quella delle elitre è 
molto svanita. L'intero corpo è coperto di tubercoletti fini e distinti, fitti sul- 
avancorpo e radi sull'addome. Edeago figg. 37-38. 

Comparazioni. Per la presenza di lunghe setole dell'addome e per la forma e 
dimensione dell' edeago, la nuova specie può essere tassonomicamente vicina ad 
A. borneana Cameron, 1943, del Borneo, ma la nuova specie ha occhi più lunghi delle 
tempie (lunghi quanto le tempie in borneana) e l'edeago in visione ventrale non è 
sinuato ai lati come in borneana e la robusta armatura genitale interna flessa ad 
angolo dell'edeago di borneana, nell'edeago della nuova specie è sostituita da 
un'ovale piastra con punta. 



ALEOCHARINAE DELLA CINA 



675 





Figo. 33-38 

Sesto urotergo libero del maschio, habitus, spermateca ed edeago in visione laterale e ventrale. 
33: Atheta (Corpothassa) iucunda sp. n.; 34-35: Atheta (Coprothassa) hsin sp. n.; 36-38: Atheta 
(Acrotona) setipyga sp. n. 



676 ROBERTO PACE 

Atheta (Acrotona) exsuperans sp. n. Figg. 39-42 

Holotypus o\ China, Beijing, B.N.U., flight interception trap, 10.VI-10.VII.1993, de 
Rougemont leg. (MHNG). 

Paratypi: 1 del 9, stessa provenienza; 1 o\ China, Shanxi, Wutaishan, 4-5.VI.1993, 
de Rougemont leg. 

Descrizione. Lunghezza 2,4 mm. Corpo lucido e bruno con elitre ed estremità 
addominale bruno-rossicci; antenne nere con antennomero basale bruno; zampe 
giallo-brune con tarsi gialli. La reticolazione è distinta sul capo e svanita sul resto del 
corpo. I tubercoletti dell 7 avancorpo sono salienti, quelli dell'addome sono estrema- 
mente svaniti. Edeago figg. 40-41, spermateca fig. 42. 

Comparazioni. La nuova specie è simile ad A. suspiciosa Motschulsky, 1859, 
diffusa dallo Sri Lanka al Nepal. Se ne distingue per avere l'armatura genitale interna 
dell'edeago più robusta e per la presenza di una lunga spina all'interno del bulbo 
basale dello stesso edeago, poco evidente nel bulbo basale dell'edeago di suspiciosa. 

Atheta (Acrotona) collusa sp. n. Figg. 43-46 

Holotypus 6, China, Beijing, Xiaolongmen, 1100-1500 m, 1. VII. 1993, de Rougemont 
leg. (MHNG). 

Paratypi: 23 es., stessa provenienza; 24 es.. China, Shanxi, Wutaishan; 4-5.VI.1993, de 
Rougemont leg.; 3 es.. China, Gansu, pass btw Hezuo-Amqog, 3300 m, 12. VII. 1994, 
A. Smetana leg. 

Descrizione. Lunghezza 2,8 mm. Corpo lucido e nero pece con elitre nero- 
brune; antenne nere; zampe bruno-rossicce. La reticolazione della superficie delle 
elitre è netta, quella sul resto del corpo è svanita, tranne che sul quinto urotergo libero 
dove è distinta. I tubercoletti delle elitre sono salienti, quelli sul resto della superficie 
corporea sono superficiali. Edeago figg. 44-45, spermateca fig. 46. 

Comparazioni. Per i caratteri dell'edeago, la nuova specie è simile ad 
A. ostentata Pace, 1991. Tuttavia l'armatura genitale falciforme interna dell'edeago è 
meno robusta e assai meno ricurva nella nuova specie, mentre l'apice dell'edeago 
della nuova specie è più largo. Il bulbo distale della spermateca è meno dilatato nella 
nuova specie. 

Atheta (Acrotona) adesiana sp. n. Figg. 47-49 

Holotypus 6, Hong Kong, N.T., IV. 1996, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,8 mm. Corpo lucido e bruno con capo e addome 
neri; antenne nere con antennomero basale giallo-rossiccio e i due successivi bruno- 
rossicci; zampe rossicce. Il capo e il pronoto mostrano una reticolazione svanita, le 
elitre e l'addome hanno reticolazione distinta, quella addominale a maglie molto 
trasverse. L'intero corpo è coperto di tubercoletti salienti. Edeago figg. 48-49. 

Comparazioni. La nuova specie per i caratteri dell'edeago è simile ad 
A. paedida (Kraatz, 1859), largamente diffusa in oriente, in Africa meridionale e nel 
Madagascar. Se ne distingue per avere il bulbo prossimale dell'edeago di maggiore 



ALEOCHARINAE DELLA CINA 



677 




Figo. 39-46 

Habitus, edeago in visione laterale e ventrale e spermateca. 39-42: Atheta (Aerofono) exsuper- 
nans sp. n.; 43-46: Atheta (Acrotona) collusa sp. n. 



678 ROBERTO PACE 

sviluppo rispetto la parte distale (in paedida la parte distale ha maggiore sviluppo 
della parte basale), per l'apice dell'edeago, in visione ventrale, largo e non sinuato ai 
lati preapicali come in paedida. 

Etimologia. La nuova specie è dedicata a Garry Ades, zoologo che ha raccolto 
Aleocharinae della Cina, pure oggetto della presente serie di lavori. 

Atheta (Acrotona) litura sp. n. Figg. 50-52 

Holotypus 6 , China, Zhejiang, Tianmushan, 29. IV. 1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,8 mm. Corpo lucido e bruno con elitre giallo-brune 
con lati esterni oscurati di bruno e con i due uriti basali e la metà distale del quinto 
libero bruno-rossicci; le antenne mancano; zampe giallo-rossicce. L'avancorpo è 
coperto di reticolazione superficiale, l'addome è privo di reticolazione. L'intero corpo 
è coperto di tubercoletti salienti. Edeago figg. 51-52. 

Comparazioni. La nuova specie, poiché ha occhi molto sviluppati, lunghe 
setole all'estremità addominale e l'edeago poco arcuato, mostra qualche affinità con 
A. subscabrosa Cameron, 1939, dell'India. Tuttavia la presenza di un'armatura geni- 
tale interna dell'edeago robusta e flessa differenzia la nuova specie da subscabrosa 
che non presenta tale armatura genitale. Inoltre le lunghissime setole sporgenti ai lati 
del corpo della nuova specie, sono ridotte in subscabrosa. 

Etimologia. Il nome della nuova specie significa "scarabocchio" a motivo 
della forma simile a uno scarabocchio dell'armatura genitale interna dell'edeago in 
visione laterale. 

Atheta (Acrotona) appulsina sp. n. Figg- 53-55 

Holotypus a , China, Zhejiang, Tianmushan, 29.IV. 1993, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 2,7 mm. Corpo lucido e nero-bruno con pronoto ed 
elitre brune; antenne nere con i due antennomeri basali bruno-rossicci; zampe 
rossicce. La reticolazione del capo e delle elitre è svanita, quella del pronoto è distinta 
e quella dell'addome è netta. I tubercoletti della superficie del capo sono svaniti, 
quelli del pronoto sono molto superficiali e quelli delle elitre e dell'addome sono 
distinti. Edeago figg. 54-55. 

Comparazioni. Per la forma dell'edeago, la nuova specie sembra affine ad 
A. subscabrosa Cameron, 1939, dell'India. Ma l'edeago è meno profondamente 
arcuato al lato ventrale e in visione ventrale ha lati sinuati e non arcuati come in 
subscabrosa. L'armatura genitale interna dell'edeago della nuova specie è molto più 
sviluppata, mentre l'armatura genitale bisinuata è più esile rispetto quella dell'edeago 
di subscabrosa. 

Atheta (Acrotona) appulsinoides sp. n. Figg- 56-58 

Holotypus ó\ China, Shaanxi, Nanwutai, 17.IX.1995, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 2,1 mm. Corpo lucido e nero pece con elitre brune; 

antenne nere; zampe giallo-brune. La reticolazione del corpo è molto superficiale. I 



ALEOCHARINAE DELLA CINA 



679 




Figo. 47-55 

Habitus, edeago in visione laterale e ventrale. 47-49: Atheta (Acrotona) adesiana sp. n.; 50-52: 
Atheta (Acrotona) litura sp. n.; 53-55: Atheta (Acrotona) appulsina sp. n. 



680 ROBERTO PACE 

tubercoletti del capo sono svaniti, quelli del pronoto e delle elitre sono nettamente 
salienti. Edeago figg. 57-58. 

Comparazioni. La nuova specie è affine ad A. subscabrosa Cameron, 1939, 
dell'India. Se ne distingue per l'edeago meno sviluppato con armatura genitale interna 
differente, figg. 54-55 e 57-58. 

Atheta (Acrotona) intercepta sp. n. Figg- 59-61 

Holotypus ó\ China, Beijing, B.N.U., flight interception trap, 10.VI.VII.1993, de 
Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,2 mm. Corpo lucido e nero-bruno con elitre giallo- 
brune ed estremità addominale bruna; antenne nere; zampe bruno-rossicce. La 
reticolazione del capo è svanita, quella del pronoto è distinta, quella delle elitre netta e 
quella dell'addome è estremamente svanita e a maglie molto trasverse. I tubercoletti 
della superficie del capo e dell'addome sono superficiali, quelli delle elitre e 
dell'addome sono salienti. Edeago figg. 60-61. 

Comparazioni. La nuova specie, in base alla forma dell' edeago, è forse affine 
ad A. scabrosa Cameron, 1939, dell'India, ma l'edeago della nuova specie è netta- 
mente più sviluppato, più profondamente arcuato al lato ventrale e con due armature 
genitali lamellari assenti nell'edeago di scabrosa. 

Atheta (Acrotona) filia sp. n. Figg. 62-64 

Holotypus 6, China, Yunnan, Xishuangbanna, Mengdien, 26.1.1993, de Rougemont 
leg. (MHNG). 

Descrizione. Lunghezza 1,9 mm. Corpo lucido e nero-bruno con elitre brune; 
antenne nere; zampe giallo-brune. Sull'avancorpo non vi è traccia di reticolazione, 
quella sull'addome è estremamente superficiale. I tubercoletti della superficie del 
pronoto sono svaniti, quelli sul resto della superficie del corpo sono salienti. Edeago 
figg. 63-64. 

Comparazioni. La nuova specie appartiene al gruppo di specie affini ad 
A. fletcheri Cameron, 1939, dell'India. L'armatura genitale finemente squamiforme si 
osserva anche in fletcheri, ma è molto più lunga e arcuata nell'edeago della nuova 
specie e retta in fletcheri. Inoltre l'edeago della nuova specie, al lato ventrale, 
presenta un angolo retto, assente nell'edeago di fletcheri e specie affini e la sua 
lunghezza al livello del bulbo basale, in visione ventrale, è doppia rispetto quella del 
bulbo basale dell' edeago ài fletcheri. 

Atheta (Acrotona) singularis sp. n. Figg. 73-75 

Holotypus S , China, Yunnan, Ruili, ca. 700 m, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 2,9 mm. Corpo debolmente lucido e bruno con elitre 
giallo-brune e margine posteriore dei tre uriti basali e apice addominale bruno- 
rossicci; antenne brune con antennomero basale giallo-bruno macchiato di bruno 
all'apice; zampe gialle. Il corpo è coperto di reticolazione superficiale e di tubercoletti 



ALEOCHARINAE DELLA CINA 



681 




Figo. 56-64 

Habitus ed edeago in visione laterale e ventrale. 56-58: Atheta (Acrotona) appulsinoides sp. n. 
59-61: Atheta (Acrotona) intercepta sp. n.; 62-64: Atheta (Acrotona) fiìia sp. n. 



682 



ROBERTO PACE 




FlGG. 65-71 

Habitus, edeago in visione laterale e ventrale e spermateca. 65-71: Atheta (Acrotona) inquinata 
Cameron. 



ALEOCHARINAE DELLA CINA 683 

fini e svaniti. La pubescenza dell'addome è fitta, simile a quella di molte specie di 
Myllaena. Edeago figg. 74-75. 

Comparazioni. La nuova specie è simile ad A. inquinata Cameron, 1939, 
dell'India e della Cina, dato che l' edeago è nettamente meno sviluppato, con armatura 
genitale interna robusta e lati dell'edeago, in visione ventrale, convergenti verso 
l'apice e non paralleli come in inquinata. Due armature genitali interne dell'edeago, 
in visione ventrale, a forma di piastre con appendice falciforme, non si osservano 
neu' edeago di fletcheri. 

Atheta (Acrotona) lingicola sp. n. Figg- 76-77 

Holotypus 5, China, Gansu, Dalijia Shan, 46 Km W Linxia, 2980 m, 10.VII.1994, 
A. Smetana leg. (MHNG). 

Descrizione. Lunghezza 3,4 mm. Corpo lucido e nero pece con il quarto urite 
libero e la base del quinto neri; antenne nere; zampe rossicce. L'avancorpo è privo di 
reticolazione, l'addome la mostra estremamente svanita e a maglie molto trasverse. 
Tubercoletti distinti o molto salienti coprono la superficie dell'avancorpo. Spermateca 
fig. 76. 

Comparazioni. La nuova specie è probabilmente tassonomicamente vicina ad 
A. inquinata Cameron, 1939, dell'India e della Cina. Se ne distingue per il bulbo 
distale della spermateca più sviluppato e con robusta introflessione apicale e per la 
parte prossimale della stessa spermateca non ampiamente sviluppata come in inqui- 
nata. 

Etimologia. Il nome della nuova specie significa "colei che abita le catene 
montuose". Infatti il sostantivo cinese "ling" significa catena montuosa. 

Atheta (Acrotona) biserrula sp. n. Figg 78-81 

Holotypus â, China, Beijing, Xishan, IX. 1992, de Rougemont leg. (MHNG). 

Paratypi: 14 es., stessa provenienza; 1 9, Beijing, B.N.U., flight interception trap, 
10.VI-10.VII.1993, de Rougemont leg.; 7 es., China, Hebei, Chengde, 3.X.1993, de Rougemont 
leg.; 8 es., Beijing, Yingtaogou, III. 1993, de Rougemont leg. 

Descrizione. Lunghezza 1,8 mm. Corpo ben convesso, lucido e bruno con 
pronoto bruno chiaro, elitre giallo-brune e margine posteriore dell' urotergo basale 
bruno rossiccio; antenne brune con l'antennomero basale giallo-rossiccio e il 
successivo rossiccio; zampe giallo-rossicce. L'intero corpo è coperto di tubercoletti 
distinti ed è privo di reticolazione. Edeago figg. 79-80, spermateca fig. 81. 

Comparazioni. Una specie che presenta una robustissima armatura genitale 
interna dell'edeago, come quella dell'edeago della nuova specie, è A. iperarmata 
Pace, 1987a, del Nepal. La nuova specie è nettamente distinta da essa per l'edeago di 
un terzo minore, con apice largo in visione ventrale (appuntito e diviso in iper- 
armata). Inoltre le due lamelle apicali a margine seghettato dell'armatura genitale 
interna dell'edeago della nuova specie, non sono presenti in iperarmata. La parte 
prossimale della spermateca della nuova specie è esile a confronto della corris- 
pondente parte della spermateca di iperarmata. 



684 



ROBERTO PACE 




Figo. 72-77 

Spermateca, habitus ed edeago in visione laterale e ventrale. 72: Atheta (Acrotona) inquinata 
Cameron; 73-75: Atheta (Acrotona) singularis sp. n.; 76-77: Atheta (Acrotona) ìingicola sp. n. 



ALEOCHARINAE DELLA CINA 685 

Etimologia. Il nome della nuova specie significa "con due seghette" per la 
presenza di due lamelle a margine seghettato all'estremità dell'armatura genitale 
interna dell'edeago. 

Atheta (Acrotona) villaekadooriensis sp. n. Figg. 82-85 

Holotypus 6, Hong Kong, N.T., Kadoorie Farm, VI. 1992, Malaise trap, G. Ades leg. 
(MHNG). 

Paratypi: 1 6 e 1 9 , stessa provenienza. 

Descrizione. Lunghezza 3,3 mm. Avancorpo debolmente lucido, addome 
lucido. Capo bruno, pronoto, elitre e i due uriti basali bruno-rossicci, restanti uriti 
neri; antenne nere con i due antennomeri basali rossicci e l'apice dell'undicesimo 
bruno; zampe giallo-rossicce. La reticolazione del capo e delle elitre è svanita, quella 
del pronoto è distinta e quella dell'addome è estremamente svanita o assente. La 
punteggiatura del capo e del pronoto è molto superficiale, quella delle elitre è distinta. 
I due uroterghi basali mostrano una pubescenza piuttosto fitta, i restanti uroterghi 
appaiono nudi, senza setole, tranne quelle al margine posteriore e sul quinto libero 
quelle più robuste. Spermateca fig. 83, edeago figg. 84-85. 

Comparazioni. In base alla forma della spermateca e per la presenza di una 
larga lamella dell'armatura genitale interna dell'edeago, la nuova specie, nonostante 
le apparenze esteriori del corpo, va attribuita al sottogenere assegnatole e proba- 
bilmente si colloca tassonomicamente presso A. fletcheri Cameron, 1939, dell'India. 
L' edeago della nuova specie è sinuato ventralmente presso l'apice: ciò non si osserva 
neh' edeago di fletcheri che inoltre non presenta l'angolo ventrale, evidente, al 
contrario, nella parte ventrale dell'edeago della nuova specie. Il bulbo distale della 
spermateca della nuova specie è molto più sviluppato di quello corrispondente della 
spermateca dì fletcheri. 

Etimologia. Il nome della nuova specie ricorda la "farm" dei fratelli 
Kadoorie, grandi filantropi di Hong Kong, fondatori del "Kadoorie Agricultural 
Research Centre" dell'Università di Hong Kong. In questo centro è stata raccolta la 
nuova specie il cui nome significa "dell'azienda agricola dei Kadoorie". 

Atheta (Acrotona) pseudofungi sp. n. Figg- 86-87 

Holotypus 9, China, Gansu, Dalijia Shan, 46 Km W Linxia, 2580 m, 10.VII.1994, 
A. Smetana leg. (MHNG). 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e bruno con capo e uriti liberi 
4° e 5° neri; antenne brune con i due antennomeri basali giallo-bruni; zampe gialle. 
La reticolazione della superficie del capo e del pronoto è svanita, quella delle elitre è 
distinta e quella dell'addome è confusa. I tubercoletti che coprono la superficie del 
corpo sono fini e superficiali. La reticolazione degli uriti liberi 4° e 5° è a maglie 
molto trasverse. Spermateca fig. 87. 

Comparazioni. Per i caratteri delle antenne e altri esoscheletrici, la nuova 
specie è determinabile come A. fungi (Gravenhorst, 1806), specie subcosmopolita 



686 



ROBERTO PACE 




Figo. 78-85 

Habitus, edeago in visione laterale e ventrale e spermateca. 78-81: Atheta (Acrotona) biserrula 
sp. n.; 82-85: Atheta (Acrotona) villaekadooriensis sp. n. 



ALEOCHARINAE DELLA CINA 687 

della regione paleartica, ma la spermateca della nuova specie ha bulbo distale 
nettamente più sviluppato e la parte mediana della stessa spermateca è robusta (esile 
m fungi). 

Atheta (Acrotona) parasuspiciosa sp. n. Figg. 88-89 

Holotypus 9, China, Gansu, M. ts 25 Km E Xiahe, 2805-2925 m, 3.VII.1994, A. 
Smetana leg. (MHNG). 

Paratypus: 1 5 , stessa provenienza. 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e nero-bruno, comprese le 
antenne; zampe giallo-brunicce. L'avancorpo è privo di reticolazione, l'addome è 
coperto di reticolazione a maglie poligonali irregolari svanite. Tubercoletti distinti 
coprono la superficie del corpo. Spermateca fig. 89. 

Comparazioni. Per la forma della spermateca, la nuova specie sembra affine 
ad A. suspiciosa Motschulsky, 1859, diffusa dallo Sri Lanka al Nepal. La spermateca 
della nuova specie è distinta da quella di suspiciosa per la taglia nettamente minore 
(circa un quarto minore), per il bulbo distale nettamente meno sviluppato e per 
l'estremità della parte prossimale della stessa spermateca, meno robusta e prolungata 
per un breve tratto (non prolungata la parte corrispondente in suspiciosa). Esterna- 
mente il capo di suspiciosa rispetto alla larghezza del pronoto è distintamente poco 
sviluppato, mentre nella nuova specie lo è di più, rispetto al pronoto. 

Atheta (Acrotona) yuzhongensis sp. n. Figg- 90-91 

Holotypus 9, China, Gansu, Yonghai, ca. 20 Km SW Yuzhong, 2700-2800 m, 
9.VIII.1994, A. Smetana leg. (MHNG). 

Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 2,4 mm. Corpo lucido e nero pece; antenne nere; 
zampe brune. L'avancorpo è privo di reticolazione, l'addome presenta una retico- 
lazione poligonale irregolare distinta. Tubercoletti distinti coprono la superficie del 
corpo. Spermateca fig. 90. 

Comparazioni. In base alla forma della spermateca, la nuova specie si 
collocherebbe in posizione intermedia tra A. parasuspiciosa sp. n. sopra descritta e 
A. suspiciosa Motschulscky, 1859. La nuova specie si distingue da parasuspiciosa per 
il bulbo distale della spermateca più sviluppato e per il prolungamento più accentuato 
della parte prossimale della stessa spermateca. E' distinta da suspiciosa per la 
spermateca evidentemente di taglia minore e per il prolungamento prossimale della 
stessa spermateca (assente in suspiciosa). 

Atheta (Acrotona) xiahensis sp. n. Figg. 92-95 

Holotypus 6, China, Gansu, M. ts 25 Km E Xiahe, 2805-2925 m, 3. Vili. 1994, 
A. Smetana leg. (MHNG). 

Paratypi: 20 es., stessa provenienza. 

Descrizione. Lunghezza 2,6 mm. Corpo lucido e nero pece; antenne nere; 
zampe bruno-rossicce. La reticolazione della superficie del capo e del pronoto è 



688 



ROBERTO PACE 




FlGG. 86-91 

Habitus e spermateca. 86-87: Atheta (Acrotona) pseudofiingi sp. n.; 
parasuspiciosa sp. n.; 90-91: Atheta (Acrotona) yuzhongensis sp. n. 



): Atheta (Acrotona) 



ALEOCHARINAE DELLA CINA 689 

distinta, quella delle elitre è assente e quella dell'addome è svanita. I tubercoletti che 
coprono il capo sono salienti e assenti sulla fascia mediana, quelli del pronoto sono 
poco salienti e quelli delle elitre sono distinti. Edeago figg. 93-94, spermateca fig. 95. 
Comparazioni. In base alla forma della spermateca e dell'edeago, la nuova 
specie sembra tassonomicamente affine ad A. scabrosa Cameron, 1939, dell'India. 
Tuttavia l'edeago della nuova specie è di taglia nettamente maggiore, con una 
rudimentale carena ventrale e lati lievemente sinuati, in visione ventrale, mentre 
l'edeago di scabrosa è privo di quasiasi carena ventrale e i suoi lati sono regolarmente 
convergenti verso l'apice, in visione ventrale. La spermateca della nuova specie ha 
bulbo distale subsferico, mentre quello di scabrosa è allungato. 

Atheta (Acrotona) mutans sp. n. Figg- 96-97 

Holotypus 9, China. Gansu, M. ts 25 Km E Xiahe, 2806-2925 m, 3.VIII.1994, 
A. Smetana leg. (MHNG). 

Descrizione. Lunghezza 2,3 mm. Corpo lucido e bruno con capo e uriti liberi 
4° e 5° neri; antenne rossicce con i tre antennomeri basali giallo-rossicci; zampe 
gialle. Assente è la reticolazione della superficie del capo e delle elitre, essa è estre- 
mamente svanita sul pronoto e a maglie trasverse, ondulate e distinte sull'addome. I 
tubercoletti della superficie del capo e delle elitre sono poco distinti, quelli del pro- 
noto sono ben salienti. Spermateca fig. 97. 

Comparazioni. La nuova specie appare affine sia ad A. xiahensis sp. n. sopra 
descritta, che ad A. scabrosa Cameron, 1939, dell'India. Si distingue da entrambe per 
la minuscola introflessione apicale del bulbo distale della spermateca che ha taglia 
minore rispetto quella delle due specie a confronto. Inoltre la parte mediana della 
spermateca della nuova specie mostra alcune maglie di scultura interna, carattere 
questo assente nelle altre due specie. 

Atheta (Acrotona) xishanensis sp. n. Figg- 98-102 

Holotypus <?, China, Beijing, Xishan, IX. 1992, de Rougemont leg. (MHNG). 

Paratypi: 1 tî e 1 9, stessa provenienza; 1 9, Beijing, Songshan, 15. V. 1993, de 
Rougemont leg.; 23 es., Beijing, B.N.U., at light, V-VI.1993, de Rougemont leg.; 1 e? e 1 9, 
Beijing, B.N.U. campus, from birds nesting box, 7.VII.1993, de Rougemont leg.; 1 c3\ Beijing, 
B.N.U., flight interception trap, 10.VI-10.VII.1993, de Rougemont leg.; 2 9, China, Henan, 
Luoyang, 10.IX.1994, de Rougemont leg.; 1 S , China, Jiangsu, Suzhoo, 3. IX. 1994, de Rouge- 
mont leg.; 13 es., China, Hebei Prov., Yongnian, 6.X.1995, Shuqiang Li leg. 

Descrizione. Lunghezza 2,4 mm. Corpo lucido e nero con pronoto giallo 
rossiccio, elitre giallo-brune e i tre uriti basali bruni con margine posteriore giallo 
rossiccio. La reticolazione è netta sul disco del capo e sulle elitre, svanita al di fuori 
del disco del capo e sul pronoto ed è assente sull'addome. I tubercoletti del capo e 
delle elitre sono svaniti, quelli del pronoto e dell'addome sono salienti, presenti anche 
nel fondo dei solchi trasversi basali dell'addome. Spermateca fig. 99, edeago figg. 
100-101, sesto urotergo libero del maschio fig. 102. 

Comparazioni. Per la presenza di un'armatura genitale interna a squame 
dell'edeago, la nuova specie si presenta simile ad A. birmana Pace, 1984, della 



690 



ROBERTO PACE 




Figo. 92-99 
Habitus, edeago in visione laterale e ventrale e spermateca. 92-95: Atheta (Acrotona) xiahensis 
sp. n.; 96-97: Atheta (Acrotona) mutans sp. n.; 98-99: Atheta (Acrotona) xishanensis sp. n. 



ALEOCHARINAE DELLA CINA 691 

Birmania e della Cina, e ad A. siamensis Pace, 1984, della Thailandia. Da entrambe la 
nuova specie è differente per avere tale armatura genitale a squame estremamente 
sottile e con squame minute (squame larghe e armatura genitale robusta nelle due 
specie a paragone). Inoltre la spermateca della nuova specie ha dimensioni ridotte 
rispetto quelle della spermateca delle due specie citate, con parte prossimale avvolta a 
spirale come in birmana, ma con profonda introflessione apicale del bulbo distale 
della stessa spermateca, introflessione assente nel bulbo distale della spermateca di 
birmana. 

Atheta (Acrotona) nanjingensis sp. n. Figg. 103-105 

Holotypus 3, China, Jiangsu, Nanjing, 17. Vili. 1994, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 1,8 mm. Avancorpo debolmente lucido, addome 
lucido. Corpo bruno con margini laterali del pronoto bruno-rossicci, elitre giallo- 
brune, i due uriti basali bruno-rossicci e l'estremità addominale rossiccia; antenne 
brune con i due antennomeri basali rossicci; zampe giallo-rossicce. La reticolazione 
della superficie del capo è confusa, quella del pronoto è svanita, quella delle elitre è 
distinta e quella dell'addome è assente. La superficie del capo ha un aspetto rugoso. 
Tubercoletti fini e fitti coprono il pronoto. La punteggiatura delle elitre è distinta. 
Edeagofigg. 104-105. 

Comparazioni. La nuova specie è affine ad A. birmana Pace, 1984, ma da essa 
distinta per avere l'edeago maggiore di quasi un terzo. Nonostante ciò le squame 
dell'armatura genitale interna dell'edeago della nuova specie sono minuscole rispetto 
quelle ampie di birmana. Inoltre, in visione ventrale, l'edeago della nuova specie è 
decisamente più largo sia al livello del bulbo basale che della parte apicale. 

Atheta (Acrotona) guandongensis sp. n. Figg. 106-107 

Holotypus 9, Hong Kong, XII. 1995-1. 1996, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e bruno con capo e quarto urite 
libero neri; antenne brune con i due antennomeri basali bruno-rossicci; zampe gialle. 
L'avancorpo è coperto di tubercoli robusti e molto salienti su un fondo non reticolato 
su capo e pronoto, con reticolazione svanita sulle elitre. I quattro uroterghi basali 
presentano una reticolazione estremamente trasversa e distinta, il quinto urotergo 
libero ha una reticolazione pure estremamente trasversa, ma svanita. I tubercoletti 
della superficie dell'addome sono salienti e fini. Spermateca fig. 107. 

Comparazioni. In base alla forma della spermateca, vi è forse qualche affinità 
tassonomica tra la nuova specie ed A. xishanensis sp. n. sopra descritta. I caratteri 
distintivi più evidenti sono che la nuova specie ha l'avancorpo coperto di robusti 
tubercoletti e ha il bulbo distale della spermateca largo, con larga introflessione 
apicale. In xishanensis il bulbo distale è stretto come l'introflessione apicale. 

Etimologia. La nuova specie prende nome dalla peninsola di Hong Kong che 
appartiene alla provincia cinese di Guandong. 



692 



ROBERTO PACE 





102 




104 




Figo. 100-105 

Edeago in visione laterale e ventrale, sesto urotergo libero del maschio e habitus. 100-102: 
Atheta (Acrotona) xishanensis sp. n.; 103-105: Atheta (Acrotona) nanjingensis sp. n. 



ALEOCHARINAE DELLA CINA 693 

Atheta (Acrotona) fraudolenta sp. n. Figg. 108-1 10 

Holotypus S, China, Beijing, Xishan, IX. 1992, de Rougemont leg. (MHNG). 
Paratypus: 1 S , stessa provenienza. 

Descrizione. Lunghezza 2,3 mm. Corpo lucido e bruno con pronoto ed elitre 
giallo-bruni, uriti liberi 3°, 4° e 5° neri; antenne brune con antennomero basale giallo 
e il successivo rossiccio; zampe gialle. Sul corpo non vi è traccia di reticolazione e i 
tubercoletti sono fini e distinti. Edeago figg. 109-1 10. 

Comparazioni. La forma dell 1 edeago dell nuova specie ricorda quella di 
A. vicaria (Kraatz, 1859), a larga diffusione orientale. L'edeago della nuova specie 
però non presenta un dente apicale, in visione laterale, né robusta armatura genitale 
interna, come in vicaria, né l'apice dell'edeago stesso, in visione ventrale, ad ogiva 
come in vicaria. 

Atheta (Acrotona) mimannuliventris sp. n. Figg- 1 1 1-1 14 

Holotypus 6, Hong Kong, Tai Po, III. 1996, de Rougemont leg. (MHNG). 
Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 1,9 mm. Corpo lucido e giallo-rossiccio con elitre 
giallo-brunicce e con urite libero 4° e base del 5° bruni; antenne bruno-rossicce con i 
due antennomeri basali gialli; zampe gialle. Solo sulla superficie delle elitre è pre- 
sente una reticolazione distinta, sul resto della superficie corporea è assente. La 
punteggiatura del capo e del pronoto è fitta e superficiale. I tubercoletti della super- 
ficie delle elitre sono molto superficiali, quelli dell'addome sono salienti. Spermateca 
fig. 112, edeago figg. 113-114. 

Comparazioni. Il colore del corpo della nuova specie è molto simile a quello 
di A. annuliventris (Kraatz, 1859) dello Sri Lanka, Taiwan, ecc. e può portare a 
un'errata determinazione senza l'esame dell'edeago o della spermateca. Questi organi 
sono charamente differenti da quelli di annuliventris. L'edeago della nuova specie è 
di taglia minuscola rispetto quella di annuliventris, senza una robusta armatura 
genitale crestata interna, come al contrario si osserva nell'edeago di annuliventris. La 
spermateca di annuliventris ha il bulbo distale con introflessione apicale a base stretta 
ed è distinto da una strozzatura che lo divide dal resto della spermateca. La 
spermateca della nuova specie, al contrario, ha base larga dell'introflessione apicale 
del bulbo distale che non è separato da una strozzatura dal resto del corpo della stessa 
spermateca. 

Atheta (Dimetrota) insinuata sp. n. Figg. 117-118 

Holotypus ?, China, Shanxi, Wutaishan, 4-5. VI. 1993, de Rougemont leg. (MHNG). 
Paratypus: 1 $ , stessa provenienza. 

Descrizione. Lunghezza 2,8 mm. Corpo lucido e nero, antenne comprese; 
zampe nere con tarsi rossicci. Una reticolazione netta sta sul disco del capo, sul 
pronoto e sulle elitre. La reticolazione dell'addome è a maglie trasverse distinte al di 
fuori dei solchi trasversi basali che mostrano una reticolazione non trasversa e netta. 



694 



ROBERTO PACE 




FlGG. 106-114 

Habitus, spermateca ed edeago in visione laterale e ventrale. 106-107: Atheta (Acrotona) 
guandongensis sp. n.; 108-110: Atheta (Acrotona) fraudolenta sp. n.; 1 11-114: Atheta (Acro- 
tona) mimannuliventris sp. n. 



ALEOCHARINAE DELLA CINA 695 

La reticolazione del capo al di fuori del disco è svanita. I tubercoletti che coprono 
l'avancorpo sono salienti, quelli dell'addome distinti. Spermateca fig. 118. 

Comparazioni. La nuova specie è distinta da A. introducta Pace, 1991, per 
l'undicesimo antennomero più lungo, per le elitre meno larghe e per la spermateca di 
dimensioni molto ridotte, con debole introflessione apicale del bulbo distale e con 
parte prossimale non sinuata come in introducta. 

Atheta (Dimetrota) sericoides sp. n. Figg. 1 19-120 

Holotypus 9, China, Beijing, Yingtaogou, III. 1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e bruno con elitre bruno- 
rossicce e quarto urite libero nero; antenne brune con i due antennomeri basali bruno- 
rossicci; zampe bruno-rossicce. La reticolazione è netta sul disco del capo che è 
impresso e sulle elitre, è distinta sul pronoto e sull'addome dove è inoltre trasversa. 
La punteggiatura del capo è superficiale e assente su una larga fascia longitudinale 
mediana. I tubercoletti della superficie del pronoto e dell'addome sono fini e poco 
salienti, quelli delle elitre sono salienti. Spermateca fig. 120. 

Comparazioni. La forma della spermateca della nuova specie è simile a quella 
di A. orphanella Cameron, 1944a, del Kashmir. Se ne distingue per la minuscola 
introflessione apicale del bulbo distale della stessa spermateca (introflessione apicale 
larghissima in orphanella) e per il bulbo prossimale sempre della spermateca appena 
più largo della parte mediana della stessa spermateca (bulbo prossimale delle 
spermateca di orphanella nettamente più largo). 

Atheta (Dimetrota) apicalis sp. n. Figg 121-124 

Holotypus 6, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 26.VII.1994, 
A. Smetana leg. (MHNG). 

Paratypi: 1 9 , stessa provenienza, ma above camp 3, 3050 m, 22.VII.1994, A. Smetana 
leg.; 2 $ , China, Shanxi, Nonwutai, de Rougemont leg. 

Descrizione. Lunghezza 3,6 mm. Corpo lucido e nero-bruno con elitre brune e 
addome nero; antenne nere con i tre antennomeri basali bruno-rossicci; zampe giallo- 
rossicce. L'avancorpo è coperto di reticolazione distinta, l'addome da reticolazione 
molto trasversa e molto superficiale. La punteggiatura del capo è svanita. Tubercoletti 
fini e distinti coprono il pronoto e gli uroterghi basali, sulle elitre sono svaniti. Edeago 
figg. 122-123, spermateca fig. 124. 

Comparazioni. Soprattutto in base alla forma della spermateca la nuova specie 
potrebbe essere tassonomicamente vicina ad A. kotgarhensis Cameron, 1939, 
dell'India. Ma l'edeago di kotgarhensis è molto più profondamente e ampiamente 
arcuato al lato ventrale e, in visione ventrale, l'apice dello stesso edeago è molto più 
largo, cioè largo quanto la metà della larghezza del bulbo basale dell' edeago stesso, 
mentre nella nuova specie l'apice è molto stretto. Inoltre l'introflessione apicale del 
bulbo distale della spermateca della nuova specie è profonda e a base molto larga in 
kotgarhensis, mentre nella nuova specie tale introflessione è breve e a base stretta. 



696 



ROBERTO PACE 




Figo. 115-121 

Habitus e spermateca. 1 15-116: Atheta (Acrotona) inornata (Kraatz); 1 17-118: Atheta 
(Dimetrota) insinuata sp. n.; 1 19-120: Atheta (Dimetrota) sericoïdes sp. n.; 121: Atheta 
(Dimetrota) apicalis sp. n. 



ALEOCHARINAE DELLA CINA 697 

Atheta (Dimetrota) paritetica sp. n. Figg. 125-127 

Holotypus <3\ China, Yunnan, Kunming, 1. II. 1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,2 mm. Corpo lucido e nero; antenne nere con i due 
antennomeri basali giallo-bruni; zampe anteriori gialle, medie gialle con femori 
giallo-bruni e posteriori con tarsi gialli e tibie e femori giallo-bruni. La reticolazione 
della superficie del capo è estremamente superficiale, quella del pronoto e delle elitre 
è svanita e quella dell'addome è netta, anche nel fondo dei solchi trasversi basali degli 
uroterghi. Il capo e il pronoto sono coperti di tubercoletti poco distinti, essi sulle elitre 
sono distinti. Il capo ha un debole e largo solco mediano posteriore. Edeago figg. 
126-127. 

Comparazioni. La nuova specie appare simile ad A. prodita Cameron, 1939, 
dell'India e del Nepal. Se ne distingue per il pronoto largo quanto il capo (pronoto più 
largo del capo in prodita) e per l'edeago a profilo ventrale angoloso (senza angolo in 
prodita). L'armatura genitale interna dell' edeago di prodita è meno sviluppata di 
quella della nuova specie. 

Atheta (Dimetrota) mixtior sp. n. Figg- 128-131 

Holotypus <?, China, Hebei, Beidaihe, 29. V. 1993, de Rougemont leg. (MHNG). 
Paratypi: 9 es., stessa provenienza. 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e bruno con gli uriti liberi 3°, 
4° e 5° neri; antenne nere con i tre antennomeri basali bruno-rossicci; zampe anteriori 
e medie bruno-rossicce con tarsi gialli, posteriori brune con tarsi gialli. La 
reticolazione della superficie del pronoto è netta, quella sul resto della superficie 
corporea è distinta. Le maglie di reticolazione degli uroterghi sono trasverse, ma non 
nel fondo dei solchi basali dove non sono trasverse. La punteggiatura del capo e delle 
elitre è svanita, quella del pronoto è quasi indistinta. Edeago figg. 129-130, sperma- 
teca fig. 131. 

Comparazioni. La nuova specie presenta occhi grandi, edeago poco svi- 
luppato, con dente apicale dorsale (in visione laterale) e apice (in visione ventrale) 
assai stretto. Simili caratteri si riscontrano nel corpo e nell'edeago di A. putridula 
(Kraatz, 1859) dello Sri Lanka. Tuttavia il margine posteriore del sesto urotergo 
libero del maschio di putridula è dentellato, mentre nella nuova specie è lineare. 
L'edeago di putridula ha bulbo basale enorme, privo di "crista apicalis" e l'apice 
dell'edeago stesso è tronco, mentre nell'edeago della nuova specie il bulbo basale è 
poco sviluppato e l'apice è appuntito. 

Atheta (Dimetrota) salamannai sp. n. Figg. 132-135 

Holotypus 6, China, Shaanxi, Nanwutai, 17.IX.1995, de Rougemont leg. (MHNG). 
Paratypi: 3 9 , stessa provenienza. 

Descrizione. Lunghezza 1,8 mm. Corpo lucido. Avancorpo bruno, addome 

rossiccio con uriti liberi 4° e 5° bruni; antenne brune con antennomero basale 

rossiccio; zampe gialle. La reticolazione della superficie del capo e dei tre uroterghi 



698 



ROBERTO PACE 




FlGG. 122-128 

Edeago in visione laterale e ventrale, spermateca e habitus. 122-124: Atheta (Dimetrota) 
apicalis sp. n.; 125-127: Atheta (Dimetrota) paritetica sp. n.; 128: Atheta (Dimetrota) mixtior 
sp. n. 



ALEOCHARINAE DELLA CINA 699 

basali è svanita, quella del pronoto è estremamente svanita, quella delle elitre è 
distinta e quella degli uroterghi liberi 4° e 5° è molto trasversa e molto superficiale. I 
tubercoletti che coprono la superficie del capo sono fitti e superficiali, quelli del 
pronoto e delle elitre sono fitti e salienti. Edeago figg. 133-134, spermateca fig. 135. 

Comparazioni. Per alcuni caratteri somatici esterni, della spermateca e 
dell'edeago, sembra che la nuova specie sia tassonomicamente vicina ad A. sublugens 
Cameron, 1939, dell'India. Essa presenta però occhi più lunghi delle tempie e non 
molto più corti delle tempie come in sublugens. L' edeago della nuova specie è di 
taglia minuscola rispetto a quella dell'edeago di sublugens, con armatura genitale 
interna composta di due lame ricurve corte e non strette e lunghissime quanto quelle 
dell'armatura genitale interna dell'edeago di sublugens. Il bulbo distale della sperma- 
teca della nuova specie, pur avendo simile introflessione apicale, non è flesso verso 
sinistra come quello della spermateca di sublugens. 

Etimologia. La nuova specie è dedicata al Prof. Giovanni Salamanna 
dell'Università di Genova che per tanti anni è stato ottimo direttore delle pubbli- 
cazioni della Società Entomologica Italiana. 

Atheta (Dimetrota) muta sp. n. Figg 136-140 

Holotypus 3, China Shaanxi, Nanwutai, 17.IX.1995, de Rougemont leg. (MHNG). 
Paratypi: 3 9 , stessa provenienza. 

Descrizione. Lunghezza 2,7 mm. Corpo lucido. Avancorpo bruno con margini 
laterali del pronoto e base delle elitre bruno-rossicci; addome giallo-rossiccio con gli 
uriti liberi 3°, 4° e 5° nero-bruni; antenne brune con i tre antennomeri basali giallo- 
rossicci; zampe gialle. La reticolazione della superficie del capo e dell'addome è 
distinta, quella del pronoto e delle elitre è netta. I tubercoletti che coprono la 
superficie dell'avancorpo sono svaniti, quelli dell'addome sono salienti. La retico- 
lazione degli uroterghi è a maglie molto trasverse. Edeago figg. 137-138, spermateca 
fig. 139, sesto urotergo libero del maschio fig. 140. 

Comparazioni. La nuova specie sembra avere qualche affinità tassonomica 
con A. sublugens Cameron, 1939, dell'India e Nepal, a motivo della presenza del 
bulbo distale della spermateca flesso al lato sinistro come quello di sublugens. Ma la 
parte prossimale della spermateca della nuova specie è poco sviluppata in lunghezza, 
mentre è lunghissima e avvolta in spire quella di sublugens. L' edeago della nuova 
specie ha profilo ventrale simile a quello di sublugens, ma l'armatura genitale interna 
non presenta due lunghe e strette lame come quelle presenti all'interno dell'edeago di 
sublugens. 

Atheta (Dimetrota) reelsi sp. n. Figg. 141-144 

Holotypus ó\ Hong Kong, N.T., IV. 1996, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,3 mm. Corpo lucido e nero; antenne brune con i 
due antennomeri basali bruno-rossicci; zampe gialle con femori bruno-rossicci. La 
reticolazione della superficie del capo, del pronoto e dei quattro uroterghi basali è 



700 



ROBERTO PACE 




Figo. 129-136 

Edeago in visione laterale e ventrale, spermateca e habitus. 1 29- 1 3 1 : Atheta (Dimetrota) 
mixtior sp. n.; 132-135: Atheta (Dimetrota) salamannai sp. n.; 136: Atheta (Dimetrota) muta 
sp. n. 



ALEOCHARINAE DELLA CINA 701 

distinta, quella del quinto urotergo libero è lievemente trasversa e svanita. I tuber- 
coletti della superficie dell'avancorpo sono fini e superficiali. Edeago figg. 142-143, 
sesto urotergo libero del maschio fig. 144. 

Comparazioni. In base alle dimensioni e alla forma dell' edeago, la nuova 
specie sembra simile ad A. subatomaria Cameron, 1939, dell'India. Tuttavia la taglia 
corporea di subatomaria è di 1,5 mm e gli occhi sono assai ridotti. L'edeago della 
nuova specie è meno profondamente ricurvo al lato ventrale e presenta un'armatura 
genitale interna distinta, mentre quella dell'edeago di subatomaria è poco distinta. 

Etimologia. La nuova specie è dedicata allo zoologo inglese Graham Reels 
che ha raccolto Aleocharinae in Cina. 

Atheta (Dimetrota) beijingensis sp. n. Figg. 145-148 

Holotypus 6, China. Beijing, Xiaolongmen, 1100-1500 m, 1. VII. 1993, de Rougemont 
leg. (MHNG). 

Paratypi: 11 es., stessa provenienza. 

Descrizione. Lunghezza 2,9 mm. Corpo lucidissimo e bruno con capo e uriti 
liberi 4° e 5° nero-bruni e con il margine posteriore dei tre uriti basali bruno-rossiccio; 
antenne brune; zampe giallo-rossicce. Sulla superficie del corpo non vi è presenza di 
reticolazione. La punteggiatura del capo è superficiale sul disco e netta ai lati, quella 
del pronoto è fine e più fitta sulla fascia mediana e quella delle elitre è netta. Tuber- 
coletti distinti coprono la superficie degli uroterghi. Edeago figg. 146-147, spermateca 
fig. 148. 

Comparazioni. La nuova specie ha l'armatura genitale interna dell'edeago 
priva di pezzi copulatori evidenti come quelli dell'edeago di A. congruens Cameron, 
1939, dell'India. La nuova specie inoltre possiede un'evidente "crista apicalis" 
dell'edeago, quasi assente in congruens, e la parte apicale dello stesso edeago è 
molto larga nella nuova specie e stretta quella dell'edeago di congruens. 

Atheta (Dimetrota) amplificata sp. n. Figg. 149-151 

Holotypus S, China, Yunnan, Ruili, ca. 700 m, 3 .11.1993, de Rougemont leg. (MHNG). 
Paratypi: 2 6, stessa provenienza. 

Descrizione. Lunghezza 3,0 mm. Corpo lucidissimo e nero con elitre nero- 
brune; antenne nere con i due antennomeri basali bruni; zampe gialle. Sulla superficie 
corporea non vi è presenza di reticolazione. La punteggiatura del capo è svanita ed è 
largamente assente sulla fascia mediana, quella del pronoto è superficiale, assente ai 
lati e agli angoli posteriori e addensata sulla linea mediana. La punteggiatura delle 
elitre è svanita. Tubercoletti salienti stanno sulla superficie degli uroterghi. Edeago 
figg. 150-151. 

Comparazioni. La nuova specie è distinta da A. kotgarhensis Cameron, 1939, 
dell'India, per gli antennomeri 4° a 6° più lunghi che larghi e non nettamente trasversi 
come in kotgarhensis, per l'edeago meno sviluppato e assai largo, se visto dal lato 
ventrale (edeago stretto visto dal lato ventrale in kotgarhensis). 



702 



ROBERTO PACE 




FiGG. 137-144 
Edeago in visione laterale e ventrale, spermateca, sesto urotergo libero del maschio e habitus. 
137-140: Atheta (Dimetrota) muta sp. n.; 141-144: Atheta (Dimetrota) reelsi sp. n. 



ALEOCHARINAE DELLA CINA 



703 




Figo. 145-152 

Habitus, edeago in visione laterale e ventrale e spermateca. 145-148: Atheta (Dimetrota) beijin- 
gensis sp. n.; 149-151: Atheta (Dimetrota) amplificata sp. n.; 152: Atheta (Dimetrota) khitana 
sp. n. 



704 ROBERTO PACE 

Atheta (Dimetrota) khitana sp. n. Figg. 152-154 

Holotypus 6 , China, Beijing, Xiaolongmen, 1 100-1500 m, 1. VIII. 1993, de Rougemont 
leg. (MHNG). 

Descrizione. Lunghezza 3,8 mm. Avancorpo debolmente lucido, addome 
lucido. Corpo nero-bruno con elitre giallo-brune; antenne nere; zampe anteriori giallo- 
rossicce, medie bruno-rossicce con tarsi gialli, posteriori giallo rossicce con femori 
bruno-rossicci. La reticolazione del capo e delle elitre è netta, quella del pronoto è 
quasi vigorosa e quella degli uroterghi a maglie molto trasverse ondulate distinte. I 
tubercoletti della superficie del capo e del pronoto sono distinti e quelli delle elitre 
sono salienti. Il pronoto ha un'impressione mediana posteriore. Edeago figg. 153-154. 

Comparazioni. La nuova specie è simile ad A. reminiscens Pace, 1988, del 
Nepal, a giudicare dalla forma dell'edeago. Se ne distingue perché l'edeago non è così 
ampiamente ricurvo al lato ventrale quanto l'edeago di reminiscens e l'armatura 
genitale interna dell'edeago stesso è poco distinta, mentre in reminiscens sono ben 
distinte due lamelle, in visione laterale. 

Etimologia. La nuova specie prende nome dai Khitani, invasori di Pekino 
nell'anno 936 d.C. 

Atheta (Dimetrota) effulta sp. n. Figg. 155-156 

Holotypus 9, China, Zhejiang, Tianmushan, 29. IV. 1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,9 mm. Corpo lucido con capo e uriti liberi 4° e 5° 
nero-bruni, pronoto bruno-rossiccio, elitre giallo-brune e i due uriti basali giallo- 
rossicci con area mediana bruno-rossiccia e il terzo libero bruno-rossiccio; antenne 
brune con i due antennomeri basali rossicci; zampe gialle. La reticolazione del capo è 
svanita, quella del pronoto e delle elitre è distinta e quella dell'addome è a maglie 
estremamente trasverse e svanite. La punteggiatura del capo è fine e distinta. I 
tubercoletti della superficie del capo sono salienti, quelli delle elitre sono molto 
svaniti e quelli degli uroterghi sono evidenti. Spermateca fig. 155-156. 

Comparazioni. La nuova specie ha il bulbo distale della spermateca volto al 
lato sinistro, pertanto è probabile la sua affinità con A. sublugens Cameron, 1939, 
dell'India. Ma l'introflessione apicale del bulbo distale della spermateca della nuova 
specie è ampia e profonda e non poco profonda come quella di sublugens. Inoltre il 
pronoto della nuova specie è poco trasverso (molto trasverso in sublugens), gli occhi 
sono lunghi quanto le tempie (e non più corti delle tempie come in sublugens) e il 
colore del corpo è del tutto differente nelle due specie. 

Etimologia. Il nome della specie significa "sostenuta". 

Atheta (Dimetrota) furtivoides sp. n. Figg. 157-160 

Holotypus 6, China, Yunnan, Dali, 9.II.1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,4 mm. Corpo lucido e nero-bruno con elitre e i due 

uriti basali bruni; antenne nero-brune con antennomero basale rossiccio; zampe 

rossicce. La reticolazione del capo è distinta, quella del pronoto è netta e quella delle 



ALEOCHARINAE DELLA CINA 



705 




Figo. 153-159 

Edeago in visione laterale e ventrale, habitus e spermateca. 153-154: Atheta (Dimetrota) 
khitana sp. n.; 155-156: Atheta (Dimetrota) effulta sp. n.; 157-159: Atheta (Dimetrota) furti - 
voides sp. n. 



706 ROBERTO PACE 

elitre e dell'addome è svanita, sull'addome a maglie estremamente trasverse. La 
punteggiatura del capo è superficiale. I tubercoletti della superficie del pronoto e 
dell'addome sono fini e poco distinti, quelli delle elitre sono svaniti. Edeago figg. 
158-159, sesto urotergo libero del maschio fig. 160. 

Comparazioni. La nuova specie è distinta da A. furtiva Cameron, 1939, 
dell'India e del Nepal, per l'edeago meno robusto, privo di bozza ventrale e netta- 
mente più stretto in visione ventrale. 

Atheta (Dimetrota) shanxiensis sp. n. Figg. 161-165 

Holotypus S, China, Shanxi, Wutaishan, 4-5. VI. 1993, de Rougemont leg. (MHNG). 
Paratypi: 2 e? e 1 9 , stessa provenienza. 

Descrizione. Lunghezza 4,5 mm. Corpo lucido e nero con elitre nero-brune; 
antenne nere; zampe bruno-rossicce con femori nero-bruni e tarsi rossicci. La 
reticolazione è distinta sul disco del capo che è impresso, quella del pronoto è ben 
visibile, quella delle elitre è netta, quella dei quattro uriti basali è a maglie appena 
trasverse e svanite, quella sul quinto urite libero è pure a maglie appena trasverse, ma 
distinte e quella posta alla base del quarto urite libero è netta. La punteggiatura del 
capo è distinta e assente sulla fascia mediana, quella del pronoto è svanita. Tuber- 
coletti superficiali coprono la superficie delle elitre e tubercoletti salienti coprono 
quella degli uroterghi. Edeago figg. 162-163, spermateca fig. 164, sesto urotergo 
libero del maschio fig. 165. 

Comparazioni. L'angolo ventrale dell' edeago della nuova specie è molto più 
ampio di quello corrispondente dell' edeago di A. kotgarhensis Cameron, 1939, 
dell'India, e la parte distale dell'edeago della nuova specie è ben distinta dal bulbo 
basale dell'edeago stesso, in visione ventrale, grazie alla presenza di sinuosità laterali, 
mentre nell'edeago di kotgarhensis il bulbo basale non è distinto dalla parte apicale. 
Inoltre l'introflessione apicale del bulbo distale della spermateca è profonda e a base 
larga in kotgarhensis e minuscola nella nuova specie. 

Atheta (Dimetrota) inflatescens sp. n. Figg 166-167 

Holotypus 9, China, Beijing, Xiaolongmen, 1100-1500 m, 9. VII. 1993, de Rougemont 
leg. (MHNG). 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e nero-bruno con elitre e 
margine posteriore dei due uriti basali bruni; antenne nere con i due antennomeri 
basali e la base del terzo bruno-rossicci; zampe rossicce. La reticolazione del capo e 
delle elitre è svanita, quella del pronoto e degli uroterghi è distinta: su questi ultimi è 
a maglie estremamente trasverse. La punteggiatura del capo è fine e poco distinta. I 
tubercoletti della superficie del pronoto e delle elitre sono poco salienti, quelli degli 
uroterghi sono distinti. Spermateca fig. 167. 

Comparazioni. Per la forma della spermateca, la nuova specie è tassonomi- 
camente vicina ad A. mitis Pace, 1988, del Nepal. La nuova specie è distinta da essa 
per le elitre chiaramente meno sviluppate in lunghezza e larghezza e per la spermateca 
di un terzo minore, con bulbo distale meno dilatato. 



ALEOCHARINAE DELLA CINA 



707 




Figo. 160-165 

Sesto urotergo libero del maschio, habitus, edeago in visione laterale e ventrale e spermateca. 
160: Atheta (Dimetrota) furtivoides sp. n.; 161-165: Atheta (Dimetrota) shanxiensis sp. n. 



708 ROBERTO PACE 

Atheta (Dimetrota) fengicola sp. n. Figg. 168-170 

Holotypus â, China, Sichuan, Langmusi, 3500-3600 m, 13.VII.1994, A. Smetana leg. 
(MHNG). 

Descrizione. Lunghezza 4,0 mm. Corpo lucido e bruno; antenne brune con i 
due antennomeri basali rossicci; zampe rossicce. La reticolazione del capo è distinta, 
quella del pronoto e delle elitre è svanita e quella degli uroterghi è a maglie molto 
trasverse e quasi non visibili tanto sono superficiali: su alcune aree della superficie di 
essi è del tutto assente. La punteggiatura del capo è distinta e assente sulla fascia 
mediana. La superficie del pronoto e delle elitre è coperta di tubercoletti distinti. 
Edeagofigg. 169-170. 

Comparazioni. L'edeago della nuova specie ha forma simile a quella dell'ede- 
ago di A. imbrium Pace, 1988, del Nepal. La nuova specie ha elitre meno larghe 
rispetto alla larghezza del pronoto, undicesimo antennomero nettamente più lungo di 
quello di imbrium, edeago con netto angolo ventrale (angolo smussato in imbrium) e 
parte apicale dell'edeago stesso più stretta di quella deiredeago di imbrium. 

Etimologia. Il nome della nuova specie significa "abitatrice dei picchi mon- 
tani". Infatti il sostantivo cinese "feng" significa picco montano. 

Atheta (Dimetrota) langmuicola sp. n. Figg. 171-172 

Holotypus 9, China. Sichuan, Langmui, 3500-3600 m, 13.VII.1994, A. Smetana leg. 
(MHNG). 

Descrizione. Lunghezza 2,6 mm. Corpo lucido e nero-bruno; antenne nere con 
antennomero basale bruno; zampe giallo-rossicce. La reticolazione del capo e del 
pronoto è netta, quella delle elitre è svanita e quella degli uroterghi a maglie molto 
trasverse e distinte. I tubercoletti della superficie del capo sono svaniti, quelli del resto 
della superficie corporea sono salienti. Spermateca fig. 172. 

Comparazioni. La spermateca della nuova specie ha struttura simile a quella 
della spermateca di A. andrewesiana Cameron, 1939, dell'India. Se ne distingue per il 
grande sviluppo in larghezza e profondità dell 1 introflessione apicale del bulbo distale 
della spermateca (introflessione stretta in andrewesiana) e per la maggiore taglia della 
stessa spermateca della nuova specie. Inoltre il pronoto della nuova specie mostra una 
netta reticolazione, mentre quello di andrewesiana ha reticolazione estremamente 
superficiale. 

Atheta (Dimetrota) subchiniphila sp. n. Figg. 173-176 

Holotypus 3. China, Sichuan, Langmusi, 3500-3600 m, 13.VII.1994, A. Smetana leg. 
(MHNG). 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e nero-bruno con elitre giallo- 
brune e addome nero; antenne brune con i due antennomeri basali giallo-bruni; zampe 
gialle. La reticolazione del capo è netta, quella del pronoto e dell'addome è distinta (a 
maglie lievemente trasverse sull'addome) e quella delle elitre è assente. I tubercoletti 
della superficie del capo e delle elitre sono distinti, quelli del pronoto sono svaniti e 



ALEOCHARINAE DELLA CINA 



709 




Figo. 166-172 

Habitus, spermateca ed edeago in visione laterale e ventrale. 166-167: Atheta (Dimetrota) 
inflatescens sp. n.; 168-170: Atheta (Dimetrota) fengicola sp. n.; 171-172: Atheta (Dimetrota) 
lengmuicola sp. n. 



710 ROBERTO PACE 

quelli degli uroterghi sono salienti. Edeago figg. 174-175, sesto urotergo libero del 
maschio fig. 176. 

Comparazioni. La nuova specie ha la forma dell'edeago simile a quella 
dell'edeago di A. kotgarhensis Cameron, 1939, dell'India. Se ne distingue per il 
minore sviluppo di quest'organo, per la minore profondità ventrale, per i lati della 
zona preapicale (in visione ventrale) paralleli e non convergenti verso l'apice 
largamente arrotondato dell'edeago di kotgarhensis. L'armatura genitale interna 
dell'edeago della nuova specie, in visione laterale, è divisa in due gruppi, mentre in 
kotgarhensis vi è un'ampia lamella sormontata da un pezzo copulatore sclerificato 
corto e stretto. La nuova specie ha occhi molto più corti delle tempie, mentre kotgar- 
hensis ha occhi lunghi quanto le tempie. 

Atheta (Dimetrota) kadooriana sp. n. Figg 177-179 

Holotypus 2 , Hong Kong, Kadoorie Farm, V.1996, de Rougemont leg. (MHNG). 
Paratypus: 1 9 , Hong Kong, Tai Po. VII. 1996, de Rougemont leg. 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e nero con pronoto, metà 
basale delle elitre, i due uroterghi basali e la base del terzo giallo-rossicci; antenne 
nere con i due antennomeri basali giallo-rossicci e il successivo bruno; zampe gialle. 
La reticolazione del capo e dei due uroterghi basali è molto svanita, quella del pronoto 
e dei restanti uroterghi è svanita e quella delle elitre è netta. Le maglie di reticolazione 
degli uroterghi 3°, 4° e 5° sono molto trasverse. I tubercoletti che coprono la 
superficie del capo sono fini e superficiali, quelli del pronoto sono fini e distinti, 
quelli delle elitre sono poco salienti e quelli degli uroterghi sono distinti. Spermateca 
figg. 178-179. 

Comparazioni. Per la forma della spermateca, la nuova specie è comparabile 
con A. subincisa Cameron, 1939, dell'India, che presenta la spermateca a bulbo 
distale molto sviluppato e sua parte prossimale a ridottissimo sviluppo, come nella 
spermateca della nuova specie che però ha minore sviluppo generale e profonda 
introflessione apicale del bulbo distale, assente in subincisa. Gli occhi di subincisa 
sono molto più corti delle tempie e non lunghi quanto le tempie come nella nuova 
specie. 

Etimologia. La nuova specie è dedicata ai fratelli Kadoorie, noti filantropi di 
Hong Kong nella cui "farm" sono state raccolte varie specie di Aleocharinae esa- 
minate nella presente serie di lavori. 

Atheta (Diometrota) pseudocollusa sp. n. Figg 180-183 

Holotypus 3, China, Gansu, pass btw Herzuo-Amqog, 3300 m, 12.VII.1994. 
A. Smetana leg. (MHNG). 

Paratypi: 9 9 , stessa provenienza; 1 1 es., China, Sichuan, Langmusi, 3500-3600 m, 
13.VII.1994, A. Smetana leg. 

Descrizione. Lunghezza 2,6 mm. Corpo lucido e nero; antenne nere con 
antennomero basale bruno; zampe bruno-rossicce. La reticolazione del capo, del 
pronoto e dell'addome è svanita, quella delle elitre è distinta. La reticolazione degli 



ALEOCHARINAE DELLA CINA 



711 




Figo. 173-180 

Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del maschio e spermateca. 
173-176: Atheta (Dimetrota) subchiniphila sp. n.; 177-179: Atheta (Dimetrota) kadooriana sp. 
n.; 180: Atheta (Dimetrota) pseudocollusa sp. n. 



7 J 2 ROBERTO PACE 

uroterghi è a maglie trasverse. I tubercoletti della superficie del capo e del pronoto 
sono distinti quelli della elitre sono salienti. Edeago figg. 181-182, spermateca 
fig. 183. 

Comparazioni. La spermateca della nuova specie è simile a quella di 
A. congruens Cameron, 1939, dell'India, però è più sviluppata, con parte prossimale 
più prolungata. L' edeago della nuova specie non è profondamente arcuato al lato 
ventrale quanto quello di congruens e l'apice dello stesso edeago, in visione ventrale, 
meno sottilmente appuntito rispetto all'apice di quello di congruens. Inoltre il pronoto 
di congruens è poco trasverso. 

Atheta (Dimetrota) variolosa sp. n. Figg- 184-188 

Holotypus 6, Hong Kong, Tai Po, III. 1996, de Rougemont leg. (MHNG). 
Paratypi: 13 es., stessa provenienza. 

Descrizione. Lunghezza 3,8 mm. Corpo lucido e nero pece con elitre brune; 
antenne nere con i tre antennomeri basali nero-bruni; zampe rossicce con femori 
bruni. La reticolazione del capo è svanita, quella del pronoto è distinta, quella delle 
elitre è netta e quella degli uroterghi è molto trasversa, superficiale e assente alla base 
di ciascun urotergo. La superficie dell'avancorpo è coperta di tubercoletti salienti, 
quella dell'addome di tubercoletti superficiali. Edeago figg. 185-186, sesto urotergo 
libero del maschio fig. 187, spermateca fig. 188. 

Comparazioni. In base alla forma della spermateca, la nuova specie sembra 
affine ad A. subincisa Cameron, 1939, dell'India. Tuttavia il bulbo distale della 
spermateca della nuova specie è meno sviluppato e la parte prossimale non è piegata 
strettamente per due volte come in subincisa. L'edeago della nuova specie ha 
maggiore taglia rispetto quello di subincisa, ha l'apice assai largo (in visione ventrale) 
e non inciso come quello di subincisa. Inoltre gli occhi della nuova specie sono 
appena più lunghi delle tempie, mentre quelli di subincisa sono molto ridotti, circa tre 
volte più brevi delle tempie. 

Etimologia. Il nome della nuova specie significa "butterata di vaiolo" a 
motivo della presenza di formazioni chitinose ventrali del bulbo basale dell'edeago, 
simili a cicatrici lasciate dal vaiolo sulla specie umana. 

Atheta (Dimetrota) cooteri sp. n. Figg. 189-193 

Holotypus d\ China. Zhejiang Prov., Anji County, ca. 480 m, Long Wang Shan, N.R., 
1 I.V. 1996, J. Cooter leg. (MHNG). 

Paratypi: 1 6 e 1 9 , stessa provenienza. 

Descrizione. Lunghezza 2,9 mm. Avancorpo debolmente lucido, addome 
lucido. Capo e pronoto bruni con riflessi bronzei, elitre brune, addome nero pece; 
antenne brune con i due antennomeri basali giallo-rossicci e il successivo rossiccio; 
zampe gialle. La reticolazione dell'avancorpo è netta, quella dell'addome è a maglie 
molto trasverse, distinte sui tre uroterghi basali e svanita sui restanti. I tubercoletti 
della superficie delle elitre sono superficiali, quelli sul resto della superficie corporea 



ALEOCHARINAE DELLA CINA 



713 




FiGG. 181-188 

Edeago in visione laterale e ventrale, spermateca, habitus e sesto urotergo libero del maschio. 
181-183: Atheta (Dimetrota) pseudocollusa sp. n.; 184-188: Atheta (Dimetrota) variolosa sp. n. 



714 ROBERTO PACE 

sono salienti. Edeago figg. 190-191, spermateca fig. 192, sesto urotergo libero del 
maschio fig. 193. 

Comparazioni. Per i caratteri dell' edeago e del margine posteriore del sesto 
urotergo libero del maschio, la nuova specie sembra tassonomicamente vicina ad 
A. swayambunathana Pace, 1991, del Nepal. Ma la parte apicale dell'edeago, in 
visione ventrale, nella nuova specie è nettamente più stretta, gli occhi della nuova 
specie sono più lunghi delle tempie e non molto ridotti come nella specie del Nepal 
che mostra inoltre antennomeri 4° a 6° più lunghi che larghi e non lunghi quanto 
larghi come nella nuova specie. 

Etimologia. La nuova specie è dedicata al suo raccoglitore Jonathan Cooter di 
Hereford (Gran Bretagna), noto studioso di Liodidae. 

Atheta (Dimetrota) inaroides sp. n. Figg 194-195 

Holotypus 9, China, Beijing, Yingtaogou, III. 1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e nero, antenne comprese; 
zampe brune. La reticolazione del capo è quasi vigorosa, quella del pronoto è nettis- 
sima, quella delle elitre netta e quella degli uroterghi lievemente trasversa e distinta. I 
tubercoletti della superficie della fronte sono svaniti, quelli sul resto del capo sono 
salienti come quelli del pronoto. Le elitre presentano tubercoletti distinti e l'addome li 
ha salienti. Spermateca fig. 195. 

Comparazioni. Per la forma della spermateca, la nuova specie sembra affine 
ad A. inari Sawada, 1977, del Giappone. Se ne distingue per avere il bulbo distale 
della spermateca più allungato e la parte prossimale della stessa spermateca, corta: 
essa non raggiunge la metà della parte mediana della stessa spermateca (parte 
prossimale che raggiunge la metà della parte mediana della spermateca in inari). 

Atheta (Datomicra) vacua sp. n. Figg. 196-198 

Holotypus â, China, Beijing, Xishan, IX. 1992, de Rougemont leg. (MHNG). 
Paratypus: 1 â, stessa provenienza. 

Descrizione. Lunghezza 1,5 mm. Corpo lucido e nero con estremità addo- 
minale bruna; antenne nere; zampe gialle con femori giallo-bruni. Su un fondo non 
reticolato i tubercoletti della superficie del capo e del pronoto sono fini e salienti, 
quelli delle elitre sono superficiali e quelli dell'addome sono distinti, più fitti sugli 
uroterghi basali. Il sesto urotergo libero è coperto di reticolazione distinta. Edeago 
figg. 197-198. 

Comparazioni. La nuova specie è distinta da A. sordiduloides Cameron, 1939, 
dell'India, oltre che per l'edeago di taglia nettamente minore, per l'assenza di distinta 
armatura genitale interna dell'edeago stesso. Inoltre la nuova specie ha elitre meno 
sviluppate e il margine posteriore del sesto urotergo libero dal maschio non sinuato 
come quello di sordiduloides. 



ALEOCHARINAE DELLA CINA 



715 




FiGG. 189-195 

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 
189-193: Atheta (Dimetrota) cooteri sp. n.; 194-195: Atheta (Dimetrota) inaroides sp. n. 



716 ROBERTO PACE 

Atheta (Datomicra) viduoides sp. n. Figg. 199-202 

Holotypus S, China, Yunnan, Xishuangbanna, Jinghong, 11.1993, de Rougemont leg. 
(MHNG). 

Paratypus: 1 ? , stessa provenienza. 

Descrizione. Lunghezza 1,8 mm. Corpo lucido e bruno con capo nero-bruno, 
con margine posteriore dei tre uroterghi basali ed estremità addominale bruno- 
rossicci; antenne nere con i due antennomeri basali giallo-bruni; zampe gialle. La 
reticolazione del capo e delle elitre è svanita, quella del pronoto e dell'addome è 
estremamente superficiale. Le maglie di reticolazione degli uroterghi sono molto 
trasverse. I tubercoletti del capo sono superficiali, quelli del pronoto e delle elitre 
distinti e quelli dell'addome sono salienti. Edeago figg. 200-201, spermateca fig. 202. 

Comparazioni. La nuova specie è simile ad A. vidua Pace, 1988, del Nepal, in 
base alla forma della spermateca. Tuttavia il bulbo distale della spermateca della 
nuova specie è più sviluppato, con introflessione apicale larga (stretta in vidua). 
Inoltre gli occhi di vidua sono più corti delle tempie e non più lunghi delle tempie 
come nella nuova specie. 

Atheta (Datomicra) antesericea sp. n. Figg- 203-207 

Holotypus S , China, Zhejiang, Tianmushan, 29.IX.1993, de Rougemont leg. (MHNG). 
Paratypi: Idei?, stessa provenienza. 

Descrizione. Lunghezza 4,1 mm. Avancorpo quasi opaco, addome lucido. 
Corpo bruno con elitre giallo-brune e uriti liberi 3°, 4° e 5° neri; antenne nere con i 
due antennomeri basali giallo-rossicci. La reticolazione dell'avancorpo è netta, quella 
dell'addome è a maglie estremamente trasverse e distinte. La punteggiatura del capo è 
assai superficiale, quella del pronoto è indistinta. Superficiali tubercoletti coprono le 
elitre, quelli dell'addome sono salienti. Sesto urotergo libero del maschio fig. 204, 
edeago figg. 205-206, spermateca fig. 207. 

Comparazioni. In base alla forma della spermateca, la nuova specie sembra 
simile ad A. formicetorum Bernhauer, 1907, del Giappone, ma il bulbo distale della 
stessa spermateca è meno sviluppato nella nuova specie e la parte prossimale, sempre 
della spermateca, è notevolmente prolungata nella nuova specie e brevissima in 
formicetorum. La parte apicale dell'edeago della nuova specie è strettissima, mentre 
in formicetorum è il doppio più larga. 

Atheta (Datomicra) permimetica sp. n. Figg. 208-212 

Holotypus 6, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 26.VII.1994, 
A. Smetana leg. (MHNG). 

Paratypi: 23 es., stessa provenienza; 1 cT, Gongga Shan, above camp 3, 3050 m, 
22. VII. 1 994, A. Smetana leg. 

Descrizione. Lunghezza 2,8 mm. Corpo lucido con pronoto debolmente 
lucido. Corpo bruno con capo bruno scuro e addome giallo-rossiccio con uriti liberi 
3°, 4° e 5° nero-bruni; antenne brune con i tre antennomeri basali giallo-rossicci; 



ALEOCHARINAE DELLA CINA 



717 




Figo. 196-204 

196-198: Atheta (Datomicra) vacua sp. n.; 199-202: Atheta (Datomicra) viduoides sp. n.; 203- 
204: Atheta (Datomicra) antesericea sp. n. 



718 ROBERTO PACE 

zampe gialle. La reticolazione del capo e del pronoto è nettissima, quella delle elitre è 
distinta e quella dell'addome è a maglie molto trasverse e molto superficiali. I 
tubercoletti della superficie del capo e del pronoto sono distinti, quelli delle elitre 
sono svaniti. Edeago figg. 209-210, spermateca fig. 211, sesto urotergo libero del 
maschio fig. 212. 

Comparazioni. La forma della spermateca potrebbe indicare che la nuova 
specie può essere tassonomicamente vicina ad A. formicetorum Bernhauer, 1907, del 
Giappone. La più evidente differenza nella spermateca è la presenza di una gibbosità 
alla base del bulbo distale, assente nella spermateca ài formicetorum. L'edeago, al 
lato ventrale, ha un distinto angolo smussato, esso in formicetorum appare come 
debolissimo e molto ottuso angolo. 

Atheta (Datomicra) bimacula sp. n. Figg 213-217 

Holotypus 9, China. Yunnan. Ruili. ca. 700 m, 3. 11.1993. de Rougemont leg. (MHNG). 
Paratypus: 1 6 . China. Yunnan. Xishuangbanna. Mengdien, 26.1.1993, de Rougemont 
leg. 

Descrizione. Lunghezza 3,2 mm. Corpo lucido. Capo nero, pronoto giallo- 
rossiccio, elitre giallo-brune, due uriti basali giallo-rossicci con macchia bruna sulla 
linea mediana, uriti liberi 3°, 4° e metà beasele del 5° bruni, estremità addominale 
giallo-rossiccia; antenne brune con i due antennomeri basali e la base del terzo giallo- 
rossicci; zampe rossicce. La reticolazione del capo è distinta, quella sul resto della 
superficie del corpo è netta. Sugli uroterghi è ben trasversa. I tubercoletti della 
superficie del capo sono svaniti sul disco e ben salienti sul resto del capo, quelli del 
pronoto, delle elitre e dell'addome sono altrettanto salienti. Edeago figg. 214-215, 
spermateca fig. 216, sesto urotergo libero del maschio fig. 217. 

Comparazioni. La nuova specie per la forma della spermateca sembra affine 
ad A. permimetica sp. n. sopra descritta, ma il colore del corpo è differente, come 
differenti sono l' edeago e il margine posteriore del sesto urotergo libero del maschio 
(figg. 209-212 e 214-217). 

Atheta (Datomicra) mimoformosa sp. n. Figg 218-219 

Holotypus 9. Hong Kong, Kadoorie Agricultural Research Centre, Malaise trap, 19- 
3 1 .V. 1996, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 1.6 mm. Corpo lucido e nero pece con elitre nero- 
brune; antenne nere con i due antennomeri basali nero-bruni; zampe gialle. Su tutto il 
corpo vi è una reticolazione svanita. La punteggiatura del capo è estremamente 
superficiale. 1 tubercoletti del pronoto e delle elitre sono indistinti, quelli dell'addome 
sono salienti. Spermateca fig. 219. 

Comparazioni. La struttura della spermateca della nuova specie è simile a 
quella della spermateca di A. formosa Cameron, 1939, dell'India, ma è esile e non 
robusta come quella ài formosa. Inoltre l'introflessione apicale del bulbo distale della 
spermateca della nuova specie è emisferica, mentre è conica m formosa. 



ALEOCHARINAE DELLA CINA 



719 




Figo. 205-212 

Edeago in visione laterale e ventrale, spermateca, habitus e sesto urotergo libero del maschio. 
205-207: Atheta (Datomicra) antesericea sp. n.; 208-212: Atheta (Datomicra) permimetica sp. n. 



720 



ROBERTO PACE 




Figo. 213-219 

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 
213-217: Atheta (Datomicra) bìmacula sp. n.; 218-219: Atheta (Datomicra) mimoformosa sp. n. 



ALEOCHARINAE DELLA CINA 



721 




227 



jr^i 



Figo. 220-229 

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 
220-224: Atheta (Datomicra) canescens (Sharp); 225-229: Atheta (Datomicra) dadopora 
Thomson. 



722 ROBERTO PACE 

Atheta (Datomicra) dalijiamontis sp. n. Figg. 230-231 

Holotypus 9, China, Gansu, Dalijia Shan, 46 Km W Linxia, 2980 m, 10.VII.1994, 
A. Smetana leg. (MHNG). 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e bruno con uriti liberi 4° e 5° 
neri; antenne giallo-brune con i tre antennomeri basali giallo-rossicci; zampe gialle. 
La reticolazione del capo è distinta, quella del pronoto e delle elitre è svanita e quella 
dell'addome è assente. La punteggiatura del capo è superficiale e assente sulla fascia 
mediana. Sul resto del corpo stanno dei tubercoletti superficiali. Spermateca fig. 231. 

Comparazioni. Per la forma della spermateca, la nuova specie sembra affine 
ad A. sordidula (Erichson, 1839), a diffusione paleartica. Se ne distingue per la taglia 
corporea maggiore, per gli antennomeri 4° e 5° più lunghi che larghi (trasversi in 
sordidula) e per il bulbo distale della spermateca meno sviluppato, con introflessione 
apicale stretta (larga in sordidula). 

Atheta (Datomicra) zhejiangensis sp. n. Figg 232-235 

Holotypus S , China, Zhejiang, Tianmushan, 29.IV. 1993, de Rougemont leg. (MHNG). 
Paratypi: 3 6 e 2 9, stessa provenienza. 

Descrizione. Lunghezza 2,4 mm. Corpo lucido e nero con elitre brune; 
antenne nere; zampe brune con tarsi gialli. La reticolazione del disco del capo è 
vigorosa, quella sul resto della superficie del capo è svanita. Il resto della superficie 
del corpo è coperta di reticolazione distinta, a maglie trasverse solo sugli uroterghi. La 
punteggiatura del capo è superficiale e assente sulla fascia mediana che è impressa. Il 
pronoto è coperto da fine punteggiatura. Tubercoletti distinti coprono la superficie 
delle elitre e tubercoletti svaniti quella dell'addome. Il quinto urotergo libero mostra 
una reticolazione molto trasversa. Edeago figg. 233-234, spermateca fig. 235. 

Comparazioni. Per la forma della spermateca, la nuova specie sembra affine 
ad A. nigra (Kraatz, 1858) a diffusione olartica, Ma il bulbo distale della spermateca 
della nuova specie è ricurvo e senza introflessione apicale (rettilineo e con profon- 
dissima introflessione apicale in nigra) e la parte prossimale della stessa spermateca 
della nuova specie è bisinuata, al contrario descrive un'ampia curva in nigra. 

L' edeago è profondamente arcuato al lato ventrale nella nuova specie e a 
profilo ventrale sinuoso in nigra. 

Atheta (Datomicra) subinopinata sp. n. Figg. 236-240 

Holotypus 9, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 26.VII.1994, 
A. Smetana leg. (MHNG). 

Paratypi: 35 es., stessa provenienza. 

Descrizione. Lunghezza 4,0 mm. Corpo lucido e bruno con addome giallo- 
rossiccio avente gli uriti liberi 3°, 4° e la base del 5° neri; antenne brune con i quattro 
antennomeri basali giallo-rossicci; zampe rossicce. La reticolazione del capo e del 
pronoto è netta, quella delle elitre è distinta e quella dell'addome è molto svanita e 
composta di maglie molto trasverse. I tubercoletti della superficie del capo sono poco 



ALEOCHARINAE DELLA CINA 



723 




Figo. 230-235 

Habitus, spermateca ed edeago in visione laterale e ventrale. 230-231: Atheta (Datomicra) 
dalijiamontis sp. n.; 232-235: Atheta (Datomicra) zhejiangensis sp. n. 



724 



ROBERTO PACE 




Figo. 236-240 

Habitus, sesto urotergo libero del maschio, edeago in visione laterale e ventrale e spermateca. 
236-240: Atheta (Datomicra) subinopinata sp. n. 



ALEOCHARINAE DELLA CINA 725 

distinti e assenti sulla fascia mediana, quelli del pronoto sono salienti, quelli delle 
elitre sono distinti e quelli dell'addome sono fini. Sesto urotergo libero del maschio 
fig. 237, edeago figg. 238-238, spermateca fig. 240. 

Comparazioni. Per la forma della spermateca, la nuova specie è tassonomi- 
camente vicina ad A. inopinata Pace, 1991, del Nepal. Se ne distingue per avere il 
bulbo distale della spermateca flesso e non rettilineo e lievemente sinuato come in 
inopinata. L' edeago della nuova specie non è ventralmente bisinuato e profondamente 
arcuato come in inopinata, né la regione preapicale dello stesso edeago (in visione 
ventrale) è larghissima con margine angoloso. Il margine posteriore del sesto urotergo 
libero del maschio di inopinata è rettilineo con un dente arrotondato a ciascun lato, 
mentre nella nuova specie è come da fig. 237. 

Atheta (Datomicra) parainopinata sp. n. Figg 241-245 

Holotypus S, China, Sichuan, Gongga Shan, above capo 3, 3050 m, 22.VII.1994, 
A. Smetanaleg. (MHNG). 

Paratypi: 32 es., stessa provenienza, ma 2800 m, 28.VII.1994, A. Smetana leg. 

Descrizione. Lunghezza 3,8 mm. Corpo lucido e bruno con addome nero- 
bruno avente il margine posteriore dei due uriti basali e l'estremità addominale 
rossicci; antenne brune con i sei antennomeri basali giallo-rossicci; zampe giallo- 
rossicce. Il capo e il pronoto presentano una reticolazione netta. Le elitre mostrano 
d'essere coperte di tubercoletti poco salienti. L'addome è privo di reticolazione. 
Edeago figg. 242-243, spermateca fig. 244, sesto urotergo libero del maschio fig. 245. 

Comparazioni. La nuova specie è affine ad A. inopinata Pace, 1991, del 
Nepal, ma più affine ad A. subinopinata sp. n. sopra descritta, se si osserva la forma 
della spermateca che ha il bulbo distale flesso. Tuttavia questo prende origine al 
livello delle docce interne della stessa spermateca e non dopo un tratto più o meno 
lungo come in subinopinata. Il confronto della forma dell 'edeago delle due specie 
(figg. 238-238 e 242-243) insieme a quello della forma del margine posteriore del 
sesto urotergo libero del maschio (figg. 237 e 245) conferma che si è in presenza di 
specie distinte. 

Atheta (Datomicra) xinlongensis sp. n. Figg 246-250 

Holotypus S, China, Gansu, Xinlong Shan, ca. 70 Km S Lanzhou, 2225-2380 m, 
7.VH.1994, A. Smetana leg. (MHNG). 

Paratypi: 13 es., stessa provenienza. 

Descrizione. Lunghezza 3,7 mm. Corpo lucido e nero-bruno con elitre brune e 
con margine posteriore dei tre uroterghi basali ed estremità addominale rossicci; 
antenne brune con i cinque antennomeri basali giallo-rossicci; zampe giallo-rossicce. 
La reticolazione dell'avancorpo è netta, quella dell'addome è estremamente svanita, 
composta di maglie estremamente trasverse. Tubercoletti distinti o salienti coprono la 
superficie dell'avancorpo. Edeago figg. 247-248, spermateca fig. 249, sesto urotergo 
libero del maschio fig. 250. 



726 



ROBERTO PACE 




Figo. 241-246 
Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 
241-245: Atheta (Datomicra) parainopìnata sp. n.; 246: Atheta (Datomicra) xinlongensis sp. n. 



ALEOCHARINAE DELLA CINA 



727 




Figo. 247-253 

Edeago in visione laterale e ventrale, spermateca, sesto urotergo libero del maschio e habitus. 
247-250: Atheta (Datomicra) xinlongensis sp. n.; 251-253: Atheta (Datomicra) régressa sp. n. 



728 ROBERTO PACE 

Comparazioni. Per la forma della spermateca, la nuova specie è affine ad A. 
parainopinata sp. n. sopra descritta. Ma l'introflessione apicale del bulbo distale della 
spermateca è breve nella nuova specie e lunga in parainopinata e il bulbo prossimale 
della stessa spermateca è più sviluppato nella nuova specie che in parainopinata. 
Tuttavia è la forma dell' edeago (figg. 242-243 e 247-248) e del margine posteriore 
del sesto urotergo libero del maschio (figg. 245 e 250) che permettono di distinguere 
nettamente le due specie. 

Atheta (Datomicra) régressa sp. n. Figg 251-255 

Holotypus S, China, Gansu, Xinlon Shan, ca. 70 Km S Lanzhou, 2225-2380 m, 7.VIII. 
1994, A. Smetana leg. (MHNG). 

Paratypus: 1 9, China, Sichuan, Gongga Shan, above camp 3, 3300-3350 m, 23. VII. 
1994, A. Smetana leg. 

Descrizione. Lunghezza 3,9 mm. Corpo lucido e bruno con elitre bruno- 
rossicce e con uriti liberi quarto e quinto neri; antenne brune con i tre antennomeri 
basali giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo e delle elitre è 
distinta, quella del pronoto netta e quella dell'addome è estremamente superficiale e a 
maglie molto trasverse. La punteggiatura del capo è svanita e assente sul disco che è 
appena impresso. Tubercoletti distinti coprono il pronoto e le elitre. Sesto urotergo 
libero del maschio fig. 252, spermateca fig. 253, edeago figg. 254-255. 

Comparazioni. La nuova specie è affine ad A. xinlongensis sp. n. sopra des- 
critta a motivo della forma della spermateca. Se ne distingue per l'angolo ventrale 
dell'edeago stretto e non ampio come quello di xinlongensis e per il differente profilo 
del margine posteriore del sesto urotergo libero del maschio (figg. 250 e 252). 

Atheta (Datomicra) graffa sp. n. Figg 256-260 

Holotypus cT, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 25.VII.1994, 
A. Smetana leg. (MHNG). 

Paratypi: 1 9, stessa provenienza; 1 S e 1 9, China, Gansu, Xinlong Shan, ca. 70 Km 
S Lanzhou, 2225-2380 m, 7.VIII.1994, A. Smetana leg. 

Descrizione. Lunghezza 3,4 mm. Corpo lucido e bruno con elitre bruno- 
rossicce e con uriti liberi 4° e 5° neri; antenne brune con i tre antennomeri basali 
giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo è netta, quella del 
pronoto è nettissima, quella delle elitre è svanita e quella dell'addome è molto 
superficiale, composta di maglie molto trasverse. La punteggiatura del capo è svanita. 
Tubercoletti fini e distinti coprono il pronoto, quelli delle elitre sono svaniti. Edeago 
figg. 257-258, spermateca fig. 259, sesto urotergo libero del maschio fig. 260. 

Comparazioni. La nuova specie è affine alle quattro precedenti immediata- 
mente descritte. Il carattere differenziale più evidente è la forma del margine pos- 
teriore del sesto urotergo libero del maschio: essa non si osserva in nessuna delle 
specie del gruppo note. 

Etimologia. La nuova specie prende nome dalla parentesi graffa il cui anda- 
mento si osserva al margine posteriore del sesto urotergo libero del maschio. 



ALEOCHARINAE DELLA CINA 



729 




Figo. 254-260 

Edeago in visione laterale e ventrale, habitus, spermateca e sesto urotergo libero del maschio. 
254-255: Atheta (Datomicra) régressa sp. n.; 256-260: Atheta (Datomicra) graffa sp. n. 



730 ROBERTO PACE 



ADDENDA 



All'elenco delle specie note o nuove per la Cina dato nella parte I della 
presente serie di lavori sulle Aleocharinae della Cina, sono da aggiungere le seguenti 
specie: 

Atheta (Philhygra) yokkaichiana Bernhauer, 1907 Figg. 4-6 

Atheta (Metaxya) yokkaichiana Bernhauer, 1907: 410 

Atheta (Philhygra) yokkaichiana: Sawada 1977: 177 
1 S, China, Zhejiang, Tianmushan, 29.IV. 1993, de Rougemont leg.; 18 es. China, Zhejiang, 
Hangzhou, 27.IV. 1993, de Rougemont. 

Specie giapponese ora nota anche in Cina. 

Atheta (Acrotona) inquinata Cameron, 1939 Figg. 65-72 

Atheta (Acrotona) inquinata Cameron, 1939: 407 
4 es., China. Beijing. Xiaolongmen. 1 100-1500 m, 1. VII. 1993, de Rougemont leg.; 1 e? e 1 9, 
China, Ganzu, Dalijia Shan, 46 Km W Linkia, 2980 m, 10.VII.1994, A. Smetana leg.; 1 6 e 
1 9 , China, Gongga Shan, above camp 3, 3050 m, 22. VII. 1994, A. Smetana leg. 

La specie era finora nota solo del Kashmir. 

Atheta (Datomicra) canescens (Sharp, 1869) Figg. 220-224 

Homalota canescens Sharp, 1869: 239 
Atheta (Datomicra) canescens: Palm 1968: 269 
19 es., China, Shanxi, Nanwutai, 17.IX.1995, de Rougemont leg. 

Specie diffusa dall'Europa centrale e settentrionale alla Siberia. Nuova per la 
Cina. 

Atheta (Datomicra) dadopora Thomson, 1867 Figg. 225-229 

Atheta dadopora Thomson, 1867: 282 
Atheta (Datomicra) dadopora: Lohse 1974: 189 
Atheta (Datostiha) poroshirica Sawada, 1978: 243. syn. n. 
Atheta (Datomicra) shuteae Pace, 1987: 420, syn. n. 
1 3 e 2 9 , China, Gansu, Pass btw Hezuo-Amqog, 3300 m, 12.VII.1994, A. Smetana leg. 

Specie diffusa dall'Europa alla Siberia, all'India settentrionale e al Giappone. 



RINGRAZIAMENTI 

Rivolgo i miei più sentiti ringraziamenti a coloro che mi hanno affidato in 
studio le Aleocharinae della Cina oggetto del presente lavoro e frutto di recenti 
raccolte: il collega Guillaume de Rougemont di Londra, il Dr Ales Smetana di 
Ottawa, Jonathan Cooter di Hereford (Gran Bretagna), Garry Ades, Graham Reels e il 
Dr Shuquang Li di Stuttgart (Germania). Per il prestito di tipi e di materiale di 
confronto rigrazio sentitamente il Dr A. F. Newton del "Field Museum of Natural 
History" di Chicago, il Dr P.M. Hammond del "Natural History Museum" di Londra, 
il Dr L. Baert delflnstitut Royal des Sciences Naturelles" di Bruxelles e il Dr L. 
Zerche del D.E.I, di Eberswalde. 



ALEOCHARINAE DELLA CINA 73 ] 



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logiques 8: 25-1 18. 

Pace, R. 1984. Aleocharinae della Thailandia e della Birmania riportate da G. de Rougemont. 
Bollettino del Museo civico di Storia naturale di Verona 1 1: 427-468. 

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d'Entomologie A: 117-131. 

Pace, R. 1987b. Staphylinidae dell'Himalaya Nepalese. Aleocharinae raccolte dal Prof. Dr 
J. Martens (Insecta: Coleoptera). Courier Forsch. -Inst. Senckenberg 93: 383-441. 

Pace, R. 1988. Aleocharinae riportate dall'Himalaya dal Prof. Franz. Parte IV. Nouvelle Revue 
d'Entomologie 5: 181-194. 

Pace, R. 1991. Aleocharinae nepalesi del Museo di Ginevra. Parte IV: Autaliini ad Athetini 
(Coleoptera, Staphylinidae). Revue suisse de Zoologie 98: 107-158. 

Pace, R. 1998a. Aleocharinae della Cina: Parte I. Revue suisse de Zoologie 105( 1 ): 139-220. 

Pace, R. 1998b. Aleocharinae della Cina: Parte II. Revue suisse de Zoologie 105(2): 395-463. 

Palm, T. 1968. Skalbaggar Coleoptera Kortvingar: Fam. Staphylinidae. In: Svensk Insektfauna 
9: 467 pp., Stockolm. 

Sawada, K. 1977. Studies on the genus Atheta Thomsom (sic!) and its allies III: Japanese 
Species described by the previous Authors. Contributions from the Biological 
Laboratory Kyoto University 25: 171 -248. 



732 ROBERTO PACE 

Sawada, K. 1978. Studies on the Genus Atheta Thomson and its Allies IV: Three New Species 
from Japan. Contributions from the Biological Laboratory Kyoto University 25: 241-248. 

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Thomson, C. G. 1867. Skandinaviens Coleoptera 9: 407 pp., Lund. 



REVUE SUISSE DE ZOOLOGIE 

Tome 105 — Fascicule 3 

Pages 

Pozzi. Stefano, Yves Gonseth & Ambros Hänggi. Evaluation de l'entre- 
tien des prairies sèches du plateau occidental suisse par le biais de 
leurs peuplements arachnologiques (Arachnida: Araneae) 465-485 

Assing, Volker. A revision of the Habrocerinae of the world. Supplement 

II (Coleoptera: Staphylinidae) 487-492 

Uhmann, Gerhard. Beschreibung von vier neuen Arten der Gattung 

Derarimus (Coleoptera, Anthicidae) aus Malaysia 493-497 

Martens, Jochen & Peter Schwendinger. A taxonomic revision of the 
family Oncopodidae I. New genera and new species of Gnomulus 
Thorell (Opiliones, Laniatores) 499-555 

Tan, Heok Hui & Maurice Kottelat. Redescription of Betta pietà 
(Teleostei: Osphronemidae) and description oïB.falx sp. n. from cen- 
tral Sumatra 557-568 

Barbalat, Sylvie. Importance of forest structures on four beetle families 
(Col.: Buprestidae, Cerambycidae, Lucanidae and phytophagous Sca- 
rabaeidae) in the Areuse Gorges (Neuchâtel, Switzerland) 569-580 

De Andrade, Maria L. Fossil and extant species of Cylindromyrmex 

(Hymenoptera: Formicidae) 581-664 

Pace, Roberto. Aleocharinae della Cina: Parte III (Coleoptera, Staphy- 
linidae) 665-732 



REVUE SUISSE DE ZOOLOGIE 

Volume 105 — Number 3 



Pages 



Pozzi, Stefano, Yves Gonseth & Ambros Hänggi. Evaluation of dry 
grassland management on the Swiss occidental plateau using spider 
communities (Arachnida: Araneae) 465 

Assing, Volker. A revision of the Habrocerinae of the world. Supplement 

II (Coleoptera: Staphylinidae) 487 

Uhmann, Gerhard. Description of four new species of the genus Dera- 

rimus (Coleoptera, Anthicidae) from Malaysia 493 

Martens, Jochen & Peter Schwendinger. A taxonomic revision of the 
family Oncopodidae I. New genera and new species of Gnomulus 
Thorell (Opiliones, Laniatores) 499 

Tan, Heok Hui & Maurice Kottelat. Redescription of Betta pietà 
(Teleostei: Osphronemidae) and description of B.falx sp. n. from cen- 
tral Sumatra 557 

Barbalat, Sylvie. Importance of forest structures on four beetle families 
(Col.: Buprestidae, Cerambycidae, Lucanidae and phytophagous Sca- 
rabaeidae) in the Areuse Gorges (Neuchâtel, Switzerland) 569 

De Andrade, Maria L. Fossil and extant species of Cylindromyrmex 

(Hymenoptera: Formicidae) 581 

Pace, Roberto. Aleocharinae from China: Part III (Coleoptera, Staphy- 
linidae) 665 



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Penard, E. 1888. Recherches sur le Ceratium macroceros . Thèse . Genève . 43 pp. 
Penard, E. 1889. Etudes sur quelques Héliozoaires d'eau douce. Archives de Biologie 9: 1-61. 
Mertens, R. & Wermuth, H. 1960. Die Amphibien und Reptilien Europas, Kramer , Frankfurt am Main , XI + 264 pp. 
Handley, C. O. Jr 1966. Checklist of the mammals of Panama, pp. 753-795. In: Wenzel, R. L. & Tipton, V. J. (eds). 

Ectoparasites of Panama. Field Museum of Natural History . Chicago , XII + 861 pp. 
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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 

CHARLES LIENHARD 
Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Président: Ivan Löbl — Muséum de Genève 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des Instituts de zoologie des universités 
suisses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, écologie, éthologie, morphologie et anatomie comparée, 
physiologie. 

Administration 

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1211 GENÈVE 6 

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Revue suisse de Zoologie 105 (4): 733-765; décembre 1998 



ZOOLOGIA ET BOTANICA '98 
Geneva, 18-20 February 1998 

(Joint meeting of the Swiss Society of Zoology and the Swiss Society of Botany) 
18 February 1998: Satellite Symposium: Ecology, Behavior and Evolution 
79-20 February 1998: Main Symposium: From Gene Genealogy to 
Organismal History 

Author Index 

Altwegg Res, Ringsby Thor-Harald & Saether Bernt-Erik 

Natal dispersal in a metapopulation of house sparrows {Passer domesticus): 

characteristics and fate of the dispersing individuals 
Andrews S., see Cuenoud Philippe et al. 
Avise John 

The history and purview of phylogeography 
Bairoch Amos 

How can protein sequence database be helpful for phylogenetic studies 
Ballabeni Pierluigi, Conconi Davide & Rahier Martine 

Oviposition preference and larval performance in a leaf beetle 
Ballard Harvey, see Nadot Sophie et ai. 
Balloux François, Goudet Jérôme & Perrin Nicolas 

Breeding system and genetic variance in the monogamous, semi-social shrew. 

Crocidura russula (Insectivora, Soricidae) 
Bartoli Pierre, see Jousson Olivier et al. 
Baur Bruno, see Locher Rolf et al. 
Bayer Clemens, see Savolainen Vincent et al. 
Bernasconi Giorgina & Keller Laurent 

Phenotype and individuai investment of cooperating ant queens 
Bernasconi Marco, Valsangiacomo Claudio, Piffaretti Jean-Claude & Ward 
Paul 

Phylogeny of Scathophagidae (Diptera, Calyptratae) based on mitochondrial 

DNA sequences 
Bersier Louis-Félix, Dixon Paul & Sugihara George 

The behaviour of the link density in food webs with increasing sampling 

effort: from scale invariance to scale dependence 
Blanckenhorn Wolf U., see Mühlhäuser Claudia et al. 
Bray Rodney A., see Hoberg Eric P. et al. 
Brünner Harald, see Lugon-Moulin Nicolas et al. 
Buner Francis 

Habitat utilization of overwintering Kestrels {Falco tinnunculus) in relation to 

perch availability, volve abundance and distribution 



734 ZOOLOGIA ET BOTANICA '98 

Castella Vincent, Goudet Jérôme & Perrin Nicolas 

Allozyme variance and habitat in the freshwater snail Physa acuta 
Chase Mark W., see Savolainen Vincent et al. 
Chautems Alain, see Perret Mathieu et al. 
Conconi Davide, see Ballabeni Pierluigi et al 
Creach Jean-Baptiste, see Nadot Sophie et al. 

Cuenoud Philippe, Manen Jean-François, Loizeau Pierre-André, Andrews S. & 
Spichiger Rodolphe 

Molecular Phylogeny and Biogeography of the Genus Ilex L. (Aquifoliaceae) 
Curcic Bozidar P. M. & Makarov Slobodan 

'Endemic differentiation of millipedes (Diplopoda, Myriapoda) in Serbia, 

Montenegro and Macedonia: taxonomic implications 

2 On geographic distribution and historical development of some cave-dwelling 

diplopods (Myriapoda) in Serbia, Montenegro and Macedonia (FYROM) 
Dajoz Isabelle, see Nadot Sophie et al. 
De Bruijn Anette, see Savolainen Vincent et al. 
De Vargas Colomban 1 , Zaninetti Louisette & Pawlowski Jan 

Cryptic diversity in the open Ocean 

'see also Pawlowski Jan et al. 
Devore-Scribante Ariane 

The Pseudoscorpions (Arachnida) of Switzerland: systematic, faunistic and 

biogeographical study 
Dietrich Barbara & Wehner R. 

Parapatric distribution of desert ants: Inter- and intraspecific variation of 

mitochondrial DNA of desert ants, Cataglyphis Foerster, 1950 
Dixon Paul, see Bersier Louis-Félix et al. 
Donoghue Michael J. 

Phylogenies, TreeBASE, and comparative biology 
Donze Gérard 

Do non-hosts help Swallowtail butterflies to find their host plants? 
Duboule Denis 

Regulation and function of vertebrate Hox genes during limb development and 

evolution 
Ducrest Anne-Lyse, see Roulin Alexandre et al. 
Dupanloup Isabelle, Schneider Stefan & Excoffier Laurent 

Inferring genetic impact of cultural and ecological barriers: a new approach 
Excoffier Laurent, see Dupanloup Isabelle et al. and Schneider Stefan et al. 
Fahrni José, see Pawlowski Jan et al. 
Faucci Anuschka 

Structure of a hydroid-seaweed community 
Fay Michael F., see Savolainen Vincent et al. 
Flook Paul K. 1 & Rowell C. H. F. 

Examining hypotheses of grasshopper evolution with molecular sequence data 

'see also Mizrahi-Meissonnier Liliana et al. 



ZOOLOGIA ET BOTANICA '98 735 



Forstner Michael R. J., see Pastorini Jennifer et al. 
Frey Daniel 

RAPDs reveal extensive genetic variability in the apomictic Erigeron annum 
Galland Nicole, Savolainen Vincent, Salamin Nicolas & Sanjovo Alexandre 

Phylogeographic relationships among the polyploid complex Ornithogalum 

umbellatum (Liliaceae), based on allozyme and RAPD data 
Gantenbein Benjamin & Scholl Adolf 

Allozymes show an unusually high genetic differentiation of Euscorpius ger- 

manus (Scorpiones: Chactidae) populations 
Goudet Jérôme, see Balloux François, Castella Vincent et al, Lugon-Moulin 

Nicolas et al. and Monsutti Alice et al 
Goudet Jerome, see Balloux François, Castella Vincent et al, Lugon-Moulin 

Nicolas et al and Monsutti Alice et al 
Goudet Jérôme & Savolainen Vincent 

Key innovations and rate of speciation: Statistical artefact or real phenomenon? 
Gouy Manolo, see Tourasse Nicolas et al. 
Greenwood Alex, see Pääbo Svante et al. 
Griffin Andrea 

Updating the path integrator through a visual fix 
Grossenbacher Kurt, see Lüscher Beatrice et al 
Guggenheim Richard, see Kaupp Andreas et al 
Güntert Marcel, see Lüscher Beatrice et al 

Hausser Jacques, see Lugon-Moulin Nicolas and Wust Saucy Anne-Gabrielle et al 
Heer Lorenz 

Rank-specific mate expenditure of males in the polygynandrous Alpine 

Accentor (Prunella collaris) 
Hellriegel Barbara, see Saxer Gerda et al 
Hoberg Eric P., Mariaux Jean, Jones Arlene & Bray Rodney A. 

Phylogeny of Eucestoda: morphological and molecular congruence 
Holzmann Maria, Piller Werner & Zaninetti Louisette 

Molecular, morphological and ecological evidence for species recognition in 

Ammonia (Foraminifera) 
Hoot Sara B., see Savolainen Vincent et al 
Hosken David 

Aspects of sperm competition in yellow dungflies 
Jokela Jukka, see Wullschleger Esther et al 
Jones Arlene, see Hoberg Eric P. et al. 
Jousson Olivier, Bartoli Pierre & Pawlowski Jan 

Use of the ITS rDNA for elucidation of the life cycles of Mesometridae (Tre- 

matoda, Digenea) 
Kaupp Andreas, Guggenheim Richard, Wirtz Sabine & Nagel Peter 

Eggs as a subject of phylogenetic research in Paussinae (Coleoptera: Cara- 

bidae) 



736 ZOOLOGIA ET BOTANICA '98 

Keller Laurent, see Bernasconi Giorgina et al. 
Krings Matthias, see Pääbo Svante et al. 
Locher Rolf & Baur Bruno 

Is the sperm pool in the hermaphroditic land snail A rianta arbustorum limited? 
Loizeau Pierre-André, see Cuenoud Philippe et al. 
Losada Freddy, see Vizoso Dita et al. 

LuGON-MouLiN Nicolas, Goudet Jérôme, Brünner Harald & Wyttenbach 
Andréas 

Microsatellites reveal the fine-scale structure of a hybrid zone in Sorex 

araneus (Insectivora, Soricidae) 
Lüscher Beatrice, Grossenbacher Kurt, Güntert Marcel & Scholl Adolf 

Genetic differentiation of the common toad (Bufo bufo) in the Alps 
Makarov Slobodan, see Curcic Bozidar P. M. et al. 1 ' 2 
Manen Jean-François, see Cuenoud Philippe et al. 
Mariaux Jean, see Hoberg Eric P. et al. 
Martin Robert D., see Pastorini Jennifer et al. 
Melnick Don J., see Pastorini Jennifer et al. 
Mizrahi-Meissonnier Liliana, Flook Paul K. & Rowell C. H. F. 

Comparison of the utility of different data partitions of the 28S rRNA for the 

reconstruction of caeliferan phylogeny 
Monsutti Alice, Perrin Nicolas, Naciri-Graven Yamama & Goudet Jérôme 

Selection and life-history responses to size-dependent prédation in Physa acuta 

(Gastropoda) 
Montoya-Burgos Juan-Ignacio & Pawlowski Jan 

Enlightening the history of Neotropical river systems by using loricariid 

catfish phylogeny 
Morton Cynthia M., see Savolainen Vincent et al. 
Mühlhäuser Claudia & Blanckenhorn Wolf U. 

Female choice for larger males in the dung fly Sepsis cynipsea - Fisher's 

runaway or good genes? 
Müller Sonia 

Genetic diversity and relationships in the neotropical catfish genus Ancistrus 

(Siluriformes, Loricariidae) as revealed by allozyme electrophoresis 
Naciri-Graven Yamama, see Monsutti Alice et al. 
Nadot Sophie, Creach Jean-Baptiste, Ballard Harvey & Dajoz Isabelle 

The evolution of pollen heteromorphism in Viola: a phylogenetic approach 
Nagel Peter, see Kaupp Andreas et al. 
Pääbo Svante, Krings Matthias, Poinar Hendrik & Greenwood Alex 

Mitochondrial DNA sequences as a means to deduce human history 
Pastorini Jennifer, Forstner Michael R. J., Martin Robert D. & Melnick Don J. 

Morphology and molecules in conflict: the phylogenetic relationship of Calli- 

mico within the Callitrichidae 
Patton James L., see Ruedi Manuel et al. 



ZOOLOGIA ET BOTANICA 98 737 

Pawlowski Jan 1 , De Vargas Colomban, Fahrni José & Zaninetti Louisette 

Calibrating ribosomal clocks in foraminifera 

^ee also De Vargas Colomban et al., Holzmann Maria et al., Jousson 

Olivier et al. and Montoya-Burgos Juan-Ignacio et al. 
Perret Mathieu, Savolainen Vincent, Chautems Alain & Spichiger Rodolphe 

Evolution of pollination systems in Sinningieae (neotropical Gesneriaceae): 

insights from molecular phylogenetics 
Perrin Nicolas, see Balloux François et al., Castella Vincent et al. and Monsutti 

Alice et al. 
Piffaretti Jean-Claude, see Bernasconi Marco et al. 
Piller Werner, see Holzmann Maria et al. 
PoiNAR Hendrik, see Pääbo Svante et al. 
Pozzi Stefano 

Evaluation of dry grassland management using spider communities 
Qiu Yin-Long, see Savolainen Vincent et al. 
Rahier Martine, see Ballabeni Pierluigi et al. 
Reyer Heinz-Ulrich, see Saxer Gerda et al. 
Richner Heinz, see Roulin Alexandre et al. 
Ringsby Thor-Harald, see Altwegg Res et al. 
Roulin Alexandre, Richner Heinz & Ducrest Anne-Lyse 

Genetic, environmental and condition-dependent effects on female and male 

ornamentation in the barn owl Tito alba 
Rowe Tiffany 

Identifying ambiguous landmarks through vector addition 

see also Flook Paul K. et al. and Mizrahi-Meissonnier Liliana et al. 
RuEDi Manuel, Smith Margaret F. & Patton James L. 

Molecular phylogeography: A glimpse to the past of a Pocket Gopher hybrid 

zone 
Saether Bernt-Erik, see Altwegg Res et al. 
Salamin Nicolas, see Galland Nicole et al. 
Sanjovo Alexandre, see Galland Nicole et al. 

Saucy Francis, see Schneiter Beat et al. and Wust Saucy Anne-Gabrielle et al. 
Savolainen Vincent, see Galland Nicole et al, Goudet Jérôme et al, Perret 

Mathieu et al and Spichiger Rodolphe et al 
Savolainen Vincent, Chase Mark W., Morton Cynthia M., Hoot Sara B., 
Soltis Douglas E., Bayer Clemens, Fay Michael F., De Bruijn Anette, Sullivan 
Stuart & Qiu Yin-Long 

Phylogenetics of flowering plants based upon a combined analysis of plastid 

atpB and rbcL gene sequences 
Saxer Gerda, Hellriegel Barbara & Reyer Heinz-Ulrich 

Population dynamics of lynx in relation to its prey - A comparison between 

Canada and Europe 
Schneider Stefan 1 & Excoffier Laurent 

Estimation of past demographic parameters using the mismatch distribution 

^ee also Dupanloup Isabelle et al 



738 ZOOLOGIA ET BOTANICA '98 

Schneiter Beat & Saucy Francis 

Juvenile dispersal in the vole Arvicola terrestris (Rodentia, Arvicolidae) 

during rainy nights 
Scholl Adolf, see Gantenbein Benjamin et al. and Lüscher Beatrice et al. 
Schuchert Peter 

Phylogeny and Classification of the Hydrozoa (Cnidaria) 
Smith Margaret F., see Ruedi Manuel et al. 
Soltis Douglas E., see Savolainen Vincent et al. 
Spichiger Rodolphe 1 & Savolainen Vincent 

Teaching botany in a molecular world 

'see also Cuenoud Philippe et al. and Perret Mathieu et al. 
Sugihara George, see Bersier Louis-Félix et al. 
Sullivan Stuart, see Savolainen Vincent et al. 
Swalla Billie J. 

Evolution and Development of the Chordate Body Plan 
Taberlet Pierre, see Wust Saucy Anne-Gabrielle et al. 
Tourasse Nicolas & Gouy Manolo 

Accounting for evolutionary rate variation among sequence sites consistently 

changes universal phylogenies deduced from rRNA and protein-coding genes 
Valsangiacomo Claudino, see Bernasconi Marco et al. 
Vizoso Dita & Losada Freddy 

Effect of the habitat complexity on the size distribution of marine invertebrates 
Wackers Felix 

Extrafloral nectar production as a herbivore-induced plant defense 
Ward Paul, see Bernasconi Marco et al. 
Wehner R., see Dietrich Barbara et al. 
WiRTZ Sabine, see Kaupp Andreas et al. 
Wüest Jean 

The evolution of the pheromone dispersing apparatus by some Hesperidae 

(Lepidoptera) 
Wullschleger Esther & Jokela Jukka 

Parasites as selective agents in two host sister taxa: Prevalences of trematode 

infection in molluscan intermediate hosts in dependence of habitat factors 
Wust Saucy Anne-Gabrielle, Hausser Jacques, Taberlet Pierre & Saucy 

Francis 

Phylogeography of the vole Arvicola terrestris as revealed by mtDNA: The 

role of historical factors? 
Wyttenbach Andréas, see Lugon-Moulin Nicolas et al. 
Zaninetti Louisette, see De Vargas Colomban et al, Holzmann Maria et al. and 

Pawlowski Jan et al. 
Zehnder Marc 

Molecular phylogenetic analysis of the tapeworm order Proteocephalidea 

based on mitochondrial 16S rDNA sequences 



ZOOLOGIA ET BOTANICA 98 739 



Abstracts 

(alphabetically by first author) 

Altwegg Res 1 , Ringsby Thor-Harald 2 & S aether Bernt-Erik 2 

1 Department of Zoology; University of Zurich; Winterthurerstrasse 190; CH-8057 Zürich; 

Switzerland 
- Department of Zoology; Norwegian University of Science and Technology; N-7034 Trond- 

heim; Norway 

Natal dispersal in a metapopulation of house sparrows {Passer domesticus); charac- 
teristics and fate of the dispersing individuals 

In this study, we examine dispersal between local populations in a metapopulation of house 
sparrows {Passer domesticus) on an archipelago in Northern Norway. The following questions 
were addressed: 1) Is dispersal random movement, or is it influenced by the spatial arrangement 
of the islands? Are there differences in dispersal tendency or dispersal distance between age 
classes and between the sexes? 2) What is the relationship between dispersal probability and 
variation in several traits that are important for life history in birds, and 3) How do dispersers 
perform compared to residents in terms of survival. The observed distribution of dispersal 
distances differed significantly from the expected distribution under the assumption of equal 
colonization and emigration rates across islands. Dispersal was more intense among less 
isolated islands. In accordance with the general trend in passerine birds, dispersal was almost 
exclusively performed by juveniles. There was no significant sex bias in dispersal tendency and 
dispersal distance. Among females, dispersers tended to be larger and had higher adult survival 
compared to residents. Among males, early hatched and low ranking individuals were more 
likely to disperse. They did not differ from resident males in adult survival rate but had lower 
survival than female dispersers. Dispersal seemed thus to be performed by viable and 
successful females, and by subordinate males. These results idicate that dispersal within the 
studied metapopulation is a phenomenon influenced by many factors which may act in different 
ways on different parts of the population. 

Avise John 

The University of Georgia; Genetics Department; Life Science Building; Athens, GA 30602- 
7223; USA 

The history and purview of phylogeography 

About 10 years have passed since the birth of phylogeography as a formal discipline. However, 
the field's gestation began in the mid- 1970' s with the introduction of mitochondrial DNA 
analyses to population genetics, and to the profound shift toward genealogical thought at the 
intraspecific level (now formalized as coalescent theory) that these methods prompted. Here, I 
will briefly trace the history and explosive growth of phylogeography, consider some of the 
conceptual ramifications of the field for conservation and population biology, and close with a 
few thoughts on future challenges for the discipline. 

Bairoch Amos 

Université de Genève; Département de Biochimie Médicale; Genève; Switzerland 

How can protein sequence database the helpful for phylogenetic studies 
No abstract available. 

Ballabeni Pierluigi, Conconi Davide & Rahier Martine 

Institut de Zoologie; Université de Neuchâtel; Rue Emile- Argand ll;CH-2007 Neuchâtel; 
Switzerland 



740 ZOOLOGIA ET BOTANICA 



Oviposition preference and larvai performance in a leaf beetle 
No abstract available. 

Balloux François, Goldet Jérôme & Perrin Nicolas 

Institut de Zoologie et d'Ecologie Animale; Université de Lausanne; CH-1015 Lausanne; 
Switzerland 

Breeding system and genetic variance in the monogramous, semi-social shrew, Croci- 
dura russula (Insectivora, Soricidae) 

The population-genetics consequences of monogamy and male philopatry (a rare breeding 
system in mammals) were investigated in the semi-social and anthropophilic shrew C. russula 
with the use of microsatellite markers. Genetic diversity was large (H=0.813) with significant 
differentiation among subpopulations (5-6%) over a small geographical scale (16 km). 
Subpopulations were themselves structured into smaller units («breeding groups») comprising 
an estimate of four breeding pairs each. Members of the same breeding groups displayed 
significant coancestries (9-10%), essentially due to strong male kinship: syntopic males were 
on average related at the half-sib level. Female dispersal among breeding groups was not 
complete (estimate 39%), and insufficient to prevent inbreeding. From our results, the breeding 
strategy of C. russula seems less efficient than classical mammalian systems (polygyny and 
male dispersal) in disentangling coancestry from inbreeding, but much more so in boosting the 
effective size of subpopulations, and thereby retaining genetic variance. 

Bernasconi Giorgina 1 3 & Keller Laurent 2 

1 Behavioral Ecology; University of Berne; Hinterkappelen 

- IZEA; University of Lausanne; Switzerland 

3 Present address: Environmental Sciences; University of Zurich: CH-8057 Zürich; Switzerland 

Phenotype and individual investment of cooperating ant queens 

Fire ant (Solenopsis invida) queens founding a colony with unrelated nestmates potentially 
face a trade-off. Increased individual investment enhances worker production, colony survival 
and growth. However, increased investment may reduce a queen's probability of surviving 
fights that invariably arise after worker eclosion. Indeed, previous studies showed that (i) 
queens lose less weight (a mesure of investment) when initiating colonies with cofoundresses 
than when alone and (ii) within associations the queen losing more weight is more likely to die. 
In this study, we tested whether queens adjust weight loss to social environment and fighting 
ability, and whether restraining weight loss directly increases survival prospects. Experimental 
manipulation of colonies showed that reduced investment by queens within associations is 
primarily a response to the presence of a nestmate and not simply a response to differences in 
per-queen brood care demands. Differences in head width were associated with relative and 
combined weight loss of cofoundresses, as well as queen survival. Manipulation of the queens' 
relative weight by feeding and exposure to contrasting social environment (queens kept alone 
or in groups) did not significantly affect survival. These results indicate that head width 
differences, or other correlated phenotypic attributes of fighting ability, influenced both 
investment strategies and survival probability of queens. That the queens with larger heads both 
invested less energy into brood rearing and were more likely to survive suggests that early 
foundress associations of ants may not be as peaceful as previously assumed. 

Bernasconi Marco 1 2 , Valsangiacomo Claudio 2 , Piffaretti Jean-Claude 2 & 
Ward Paul 1 

1 Zoologisches Museum der Universität; Abt. Ökologie; Winterthurerstrasse 190; CH-8057 
Zürich; Switzerland 

2 Istituto Cantonale Batteriosierologico: Via Ospedale 6; CH-6904 Lugano; Switzerland 



ZOOLOGIA ET BOTANICA 98 741 

Phylogeny of Scathophagidae (Diptera, Calyptratae) based on mitochondrial DNA 
sequences 

Scathophagids flies, with more than 250 species, are mainly confined to the Holarctic region. 
Individuals of most species feed on vertebrate dung or decomposing carcasses, performing the 
ecologically-important function of resource recycling. One species, Scathophaga stercoraria 
has been used extensively to investigate questions in animal ecology and evolution. However, 
the taxonomy and phylogeny of this family still remain unclear. This study represents the first 
molecular approach aimed at uncovering the phylogenetic relationships among species of this 
family. A portion of nearly 800 bp of the terminal region of the mitochondrial gene COI (the 
subunit I of the cytochrome oxidase gene) was sequenced in over 30 species covering a wide 
geographic area, as well as a diverse spectrum of ecological habitats. In general, molecular data 
are in agreement with previous morphological information. However, some peculiarities reveal 
discrepancies between the two approaches. Some examples which will be discussed are: (i) S. 
taeniopa and S. suilla, morphologically distinct species, are molecular indistinct taxa (ii) the 
subdivision of the Norellia genus in two different subgenera, as proposed by morphological 
characters, is not supported by molecular data. 

Bersier Louis-Félix 1 , Dixon Paul 2 & Sugihara George 2 

1 Institut de Zoologie; Rue Emile Argand, 1 1 ; CH-2007 Neuchâtel; Switzerland 

2 Scripps Institution of Oceanography; UCSD; La Jolla, CA 92093; USA 

The behaviour of the link density in food webs with increasing sampling effort: from 
scale invariance to scale dependence 

We examined the scaling behaviour of the link density property of food webs with varying 
levels of sampling effort used to document the webs. The results of two models as well as 
empirical evidence concur in showing that the link density tends toward scale invariance with 
low sampling effort, and that it becomes scale dependent with increasing resolution in docu- 
menting the webs. This is a reasonable explanation for the discrepancies found in the literature 
on this question, suggesting that, for this property, scale-invariant and scale dependent 
hypotheses may not be exclusive hypotheses, but rather elements of a continuum. This finding 
raises two significant issues. First, in cross-system comparisons, it points to the difficulty of 
disentangling between effects due to differences in sampling effort and to biological dif- 
ferences. Second, in giving an equal weight to all trophic relations, binary links tend to distort 
the image of a web with increasing sampling effort, where weaker and weaker links (in term of 
biomass transferred) are reported. Both issues highlight the need of considering links quanti- 
tatively in the description of food webs. 

Buner Francis 

Zoologisches Institut Basel; Vogelwarte Sempach; Switzerland 

Habitat utilization of overwintering Kestrels {Falco tinnunculus) in relation to perch 
availability, vole abundance and distribution 

In the last two decades Kestrel numbers declined in most European areas, presumably because 
of agricultural changes. To this day most studies about Kestrels were looking at its finding 
behaviour, very little is known about its winter behaviour and ecology. 

In winter 1996/97 I radio tracked 30 Kestrels in an area of about 20 km 2 in the Klettgau 
(Canton of Schaffhausen) from December till April. In soon became clear that the studies 
winter population consisting of territorial pairs, individual birds, winter guests, migrants and 
cold weather migrants, was more complicated and dynamic as believed so far. During a cold 
spell at the end of December, beginning January, about 50% of the Kestrels did leave the study 
area. The movements were mainly observed in an area which held most individuals in late 



742 ZOOLOGIA ET BOTANICA '98 



autumn. This about 5 km- big area called «Widen» is intensively used agricultural land, 
characterized by a relatively high abundance of «ökologische Ausgleichsflächen» and small 
mammal numbers. I could show that the winter distribution and movements of the Kestrels 
were not only due to the weather situation but also to the combined influence of habitat 
distribution and quality, perch availability as well as vole abundance and distribution. 
To support our overwintering Kestrels it seems to be essential to lay out further «ökologische 
Ausgleichsflächen» and bigger extensively cultivated ares in combination with erecting more 
perches. 

Castella Vincent, Goudet Jérôme & Perrin Nicolas 

Institut de Zoologie et d'Ecologie Animale; Bâtiment de Biologie; Université de Lausanne; 
CH-1015 Lausanne; Switzerland 

Allozyme variance and habitat in the freshwater snail Physa acuta 

The wide-spread freshwater Gastropod Physa acuta colonizes big lakes as well as small, 
temporary ponds. Both habitats show a pronounced difference in temporal and spatial structure. 
Populations living in these two distinct habitats might then be genetically differentiated. 
460 snails were collected from 12 different sites, six sites being of lake habitats (Léman and 
Neuchâtel) and six sites being of pond habitats. All individuals were genotyped using six 
polymorphic allozyme loci. 

Differentiation was high between ponds' populations. Despite their low dispersal abilities snails 
from the lake Léman were genetically weakly differentiated. All populations showed a strong 
heterozygotes deficiency which was independent of their environment. This suggests that this 
deficiency is due to selling and consanguineous matings. In spite of their recent colonization, 
the genetic variability of some pond's populations was as high as those from the lakes. The 
presence of two genetically different gene pools between these distinct habitats suggests the 
existence of two sibling species or the possible action of selection. 

Cuenoud Philippe 1 , Manen Jean-François 1 , Loizeau Pierre-André 1 , Andrews S. 2 

& Spichiger Rodolphe 1 

1 Conservatoire et Jardin botaniques; Chemin de l'Impératrice, 1; CH-1292 Chambésy; 

Switzerland 
- Royal Botanic Garden; Kew; Richmond; Surrey; UK 

Molecular Phylogeny and Biogeography of the Genus Ilex L. (Aquifoliaceae) 

The genus Ilex L. (Aquifoliaceae) contains about 600 species of trees and shrubs distributed 
worldwide with a maximum diversification in east Asia and south America. It is a quite ancient 
genus, with fossils known from 90 my ago. The phylogeny of Ilex has been investigated using 
chloroplastic DNA. The atpB/rbcL spacer has been sequenced for 110 species. For 41 of these 
species, the rbcL gene has been sequenced in addition to the spacer. The phylogenies inferred 
from this database are presented, and their implications for the taxonomy and biogeography of 
the genus are discussed. In general, this genus appears to be a difficult taxon to study. In 
particular, the taxonomical treatments published by classical authors show only limited 
correlation with the molecular information. The biogeographical interpretation of the molecular 
phylogenies is problematic too. These difficulties may arise from the very long history of the 
genus, in which many extinctions and subsequent radiations have probably blurred the patterns 
that could have existed. 

Curcic Bozidar P. M. & Makarov Slobodan 

Institute of Zoology; Faculty of Biology; University of Belgrade; Studentski Trg 16; 11000 
Belgrade; Yugoslavia 

Endemic differentiation of millipedes (Diplopoda, Myriapoda) in Serbia, Montenegro 
and Macedonia: taxonomic implications 



ZOOLOGIA ET BOTANICA '98 743 



The fauna of diplopods in Serbia, Montenegro and Macedonia is very rich and diverse. Out of 
136 species inhabiting the areas studied, 48 (or 35.29%) are endemic to these regions; further- 
more, of 48 genera, only 4 (or 8.33%) are endemic to the same areas, but none of the families. 
In Serbia, 70 diplopod species have been found, and 59 and 59 in Montenegro and Macedonia, 
respectively. There exist no endemic genera in Serbia, but Montenegro and Macedonia are 
inhabited by 1 and 3 endemic general respectively. 

Serbia, Macedonia and Montenegro have in common 33 species, 23 genera and 14 families of 
diplopods. Serbia and Montenegro are inhabited by 18 common species, 15 genera and 9 
families, while Serbia and Macedonia have in common 29 species, 21 genus and 13 families; 
and Montenegro and Macedonia are characterized by shared 16 species, 12 genera and 7 
families. There are neither endemic species nor genera shared either by Serbia and Montenegro 
or by Montenegro and Macedonia. Only 5 endemic species and 1 genus are endemic both to 
Serbia and Macedonia; these are: Albanoglomus ljubetensis Attems, Brachydesmus (Styio- 
brachydesmus) ljubetensis Attems, Xestoiulus (Oroiulns) macédoniens (Attems), Megaphylllum 
crassum (Attems) and Typhloiulus (Typhloiulus) albanicus Attems. The majority of endemic 
diplopods in the areas studies belong to the families Polydesmidae (16 species, or 33.33% of all 
endemic forms), and Julidae (14 species, or 29.17% of all endemic forms). 
The abundance of both higher and lower taxa of endemic diplopods in Serbia, Montenegro and 
Macedonia may be partly explained by the long-lasting tectonic movements by the folding and 
faulting processes, by the epeirogenetic movements and by the evolution of the karstic relief 
which have greatly influenced the distribution of the ancient diplopod fauna in the Balkan 
Peninsula. 

In conclusion, Serbia, Montenegro and Macedonia are inhabited by an extremely varied fauna 
of endemic diplopods pertaining to different phyletic lineages; these forms are of different 
origins and ages. Therefore the Balkan Peninsula represents one of the main global centres of 
endemic differentiation of the millipede fauna. 



Curcic Bozidar P. M. & Makarov Slobodan 

Institute of Zoology; Faculty of Biology; University of Belgrade; Studentski Trg 16; 11000 
Belgrade; Yugoslavia 

On geographic distribution and historical development of some cave-dwelling diplo- 
pods (Myriapoda) in Serbia, Montenegro and Macedonia (FYROM) 

The analysis of geographic distribution and taxonomic interrelationships of cave millipedes has 
clearly shown that Serbia, Montenegro and Macedonia are characterized by an extremely rich 
and diverse diplopod fauna inhabiting the underground habitats of the areas studied. Therefore, 
this study has shown that each of these regions is inhabited by 16, 8, and 3 cavernicolous 
species, classified into 9, 5, and 2 genera, and 7, 5, and 2 families, respectively. Each endemic 
species is restricted to one of the areas studied. A single genus is common both to Serbia and 
Montenegro, while no genera have been found either common to Serbia and Macedonia, or to 
Montenegro and Macedonia, respectively. Only one millipede family is common both to Serbia 
and Macedonia, as well as to Montenegro and Macedonia, respectively; additionally, two diplo- 
pod families are common both to Serbia and Montenegro. Of all cave species, 13 troglobitic 
forms inhabit Serbia (3 are either troglophiles or trogloxenes), 6 - Montenegro (2 are troglo- 
philes and trogloxenes), and 1 - Macedonia (2 trogloxene and troglophile species). 
The underground diplopods of Serbia, Montenegro, and Macedonia are probably the remains of 
some ancient hygrophilic forms. With the climatic changes (and the subsequent increase of 
aridity) and the origin and development of different niches, these forms evolved as cave 
inhabitants. However, the main reasons for the outstanding variety of cave diplopods in the 
Balkan Peninsula (including the areas studied) are: the presence of a varied diplopod fauna 
once inhabiting the Proto-Balkans, the evolution of the underground karstic relief, the favour- 
able climatic changes and the subsequent radiation of the diplopod taxa in numerous isolated 
habitats. 



744 ZOOLOGIA ET BOTANICA '98 



De Vargas Colombari 1 , Zaninetti Louisette 1 & Pawlowski Jan 1 - 2 

'Department de Zoologie et Biologie Animale; Université de Genève; CH- 1224 Chêne- 

Bougeries; Switzerland 
2 Muséum d'histoire naturelle; CP 6434; CH-121 1 Genève 6; Switzerland 

Cryptic diversity in the open Ocean 

To estimate the biodiversity of the open ocean and to understand how speciation can occur in 
such a hormogeneous environment mixed by water currents, is a new deal for oceanography. 
We have examined the genetic diversity in the foraminifer Orbulina universa, one of the most 
commonly encountered planktonic organism inhabiting the world Ocean between 50° N and 
50° S. This species appeared in the fossil record 16 Myr ago and is largely used as a strati- 
graphic and paleoclimatic index. By sequencing SSUrRNA genes of several specimens 
collected in 33 stations across the Atlantic, we have shown large and geographically localized 
genetic differences among O. universa. This variability reveals three cryptic allopatric species, 
adapted to different water masses. The molecular timing of their speciation is compared to the 
history of the Atlantic. 

Devore-Scribante Ariane 

Muséum d'histoire naturelle; CP 6434; CH-121 1 Genève 6; Switzerland 

The Pseudoscorpions (Arachnida) of Switzerland: systematic, faunistic and biogeo- 
graphical study 

Presently, 58 species of pseudoscorpions are recorded from Switzerland, 13 of which are 
recorded for the first time from this country. 
The aims of the present study are: 

- to provide a detailed description of male, female and the 3 postembryonic stages, of each 
species, 

- to examine the taxonomic status of each species and to give reliable keys for the Swiss fauna, 

- to complete to knowledge the distribution (maps) and the ecology of the treated species. 

The systematic study includes morphometrical measures and chaetotaxy (tables); the diagnostic 

characters are figured. 

The distributional data are based on the material of the collection of the Museum of Natural 

History, Geneva. 

Switzerland, due to its specific position in Europe enhances the presence of a large range of 

species with different distributions patterns: 

- the widely distributed and cosmopolitan-type 

- the Mediterranean-type (South of Alps) 

- the eastern-type (North-East) 

- the western-type (along the Jura mountain) 

- the alpine-type 

- the endemics and apparent endemics (cavernicolous and other species from specific loca- 
lities). 

Dietrich Barbara & Wehner R. 

Institute of Zoology; University of Zürich; Winterthurerstrasse 190; CH-8057 Zürich, Switzer- 
land 

Parapatric distribution of desert ants: Inter- and intraspecific variation of mitochon- 
drial DNA of desert ants, Cataglyphis Foerster, 1950 



ZOOLOGIA ET BOTANICA '98 745 



The ants of the genus Cataglyphis are a highly specialized species, which inhabit the food 
impoverished areas of the Old World deserts. Cataglyphis has evolved amazing navigational 
abilities to cope with this scanty environment. The three prominent species of the bicolor 
group, formerly described as one species {"bicolor"), show a strongly parapatric North-South 
distribution (Wehner, R. et al. 1994: Senckenbergia biologica 74: 163). 

Restriction fragment length polymorphism (RFLP) analysis of mitochondrial DNA (mt-DNA) 
was chosen as the technique to investigate the ants population structures on a molecular basis. 
In within species comparisons mt-DNA phylogenies have been proven useful in population 
studies as well as in the defining themselves. Many disadvantages of traditional RFLP analysis 
for analysing population variation using individual insects, such as the small amount of DNA, 
are eliminated by carrying out polymerase chain reactions (PCR) in advance. Two primer pairs 
wer used (CoI->CoII, N1NFA->CB3FC) to amplify the mt-DNA sequences. To cut them we 
used three restriction enzymes Dral, Sspl and Taql. 

Ants of eleven different C. bicolor populations (oasis- and steppe populations) were analysed. 
A comparison between populations from continuous and disruptive habitats showed that these 
two population series hardly differ. 

The results of genetic variation in different populations of C bicolor were compared with the 
results received by examining closely related species of the same genus (C. savignyi, C. 
viaticus, C. velox and C. rosenhauri). Especially the N1NFA->CB3FC sequence confirms that 
all species of the bicolor group are clearly distinct species. Methodically the results reveal 
which part of the mt-DNA of Cataglyphis is effective in resolving the phylogeny and popul- 
ation structures of the bicolor species group. 

DoNOGHUE Michael J. 

University of Harvard; Department of Organismic and Evolutionary Biology and Harvard 
University Herbaria; 22 Divinity Avenue; Cambridge, MA 02138; USA 

Phylogenies, TreeBASE, and comparative biology 

Many evolutionary studies require access to multiple phylogenetic trees. I will illustrate a 
variety of such studies, including analyses of ( 1 ) general patterns in homoplasy, (2) change in 
particular characters over composite phylogenies (supertrees), (3) the sensitivity of particular 
comparative results to alternative phylogenetic hypotheses, (4) gene trees with respect to 
species trees, and (5) species trees with respect to biogeography and conservation. Access to 
original phylogenetic data is needed for reanalysis with the same or with different methods, and 
for the combination of different sources of evidence. In view of such needs, the development of 
databases of phylogenetic knowledge should be a high priority, yet this endeavor has received 
little attention. I will briefly describe the TreeBASE database in order to explore a number of 
issues surrounding the design and use of a database of published trees and data matrices. 

Donze Gérard 

Institut de Zoologie; Rue Emile Argand 11; CH-2007 Neuchâtel; Switzerland 

Do non-hosts help Swallowtail butterflies to find their host plants? 

Most caterpillars will only grow on a limited range of plant species so that the biochemical 
tolerance of larvae and female oviposition must be in accordance even as they are distinct 
physiological processes. Host choice by females is crucial for the survival of first instar 
caterpillars which are themselves unable to select a host. When searching for host plants, in a 
pre-landing phase, females receive visual and olfactory inputs. After landing on a plant, the 
female can further assess the quality of the plant by mechanical and gustatory stimuli. The 
efficiency of the host-finding depends on the ability to quickly differentiate the non-host 
species from suitable host plants. 

I have studied the pre-landing perception of plants by butterflies and hypothesized that Papilio 
polyxenes (Papilionidae) females use their previous experiences (i.e., both acceptances and 



746 ZOOLOGIA ET BOTANICA '98 



rejections) in order to enhance their judgement. We tested this hypothesis using olfactory cues 
from a host plant, the carrot (Daucus carota), and from a non-host plant, the yarrow (Achillea 
millefolium). To simplify the model, I controlled visual stimuli (i.e.. shape and color) and 
gustatory stimulants with the help of model plants cut from sponge. The results showed that 
experienced females, which had previously had been in contact with both the host and the non- 
host plants, were better able to avoir landings on sponges odorified with no-host odour 
compared to females which had contact only with the host plant. 

As the sensory input is multi-modal, and since insects are able to learn, use of previous positive 
and negative experiences reduces the error of landings on plants unsuitable for their cater- 
pillars. 

Duboule Denis 

Dept. of Zoology and Animal Biology; Faculty of Sciences; University of Geneva; CH-1211 
Genève 4; Switzerland 

Regulation and function of vertebrate Hox genes during limb development and 
evolution 

The function of Hoxd genes during development and their potential role in the evolution of the 
vertebrate limb has been studied in two different systems. In mice, single, double as well as 
triple inactivations (cre-mediated deletions) have been produced to assess the function of these 
genes during development. The results demonstrate that Hoxd genes act in a coordinated and 
cooperative way, where quantitative interactions seem to be at least as important as qualitative 
ones. In a different approach, we analysed the development of fish appendicular skeletons. 
Pectoral fins are the appendicular structures which are, in fishes, homologous to the tetrapod 
forelimbs. We initiated a comparative study in lower vertebrates in the aim of understanding 
the ontogenetic and phylogenetic relationships between these two structures. We have 
undertaken a detailed analysis of the development of the endoskeletal component of the 
pectoral fins of the zebrafish (Danio rerio). The sequence of appearance of the various 
mesenchymal prechondrogenic condensations has been determined as well as the mitotic 
dynamics in the mesenchymal sheets. In parallel, we cloned and characterized the posterior 
halves of the fish HOXA and HOXD complex, i.e. the Hoxd-9 to d-13 genes as well as the 
Hoxa-9 to Hoxa-13 genes. These genes were shown, in mammals, to play an important function 
in limb pattern formation. The expression of these genes during pectoral fin development were 
analysed in details and compared to the expression of a potential morphogen molecule encoded 
by the fish gene hedgehog. The significance of these results will be discussed in the light of the 
current theories (mainly derived from paleothological data) which account for the fin to limb 
transitions. The fin to limb transition will be taken as an example to illustrate some aspects of 
the functional organization of the Hox gene family in vertebrates, in relation with the evolution 
of functions. In particular, the involvement of this genetic system both in the evolution of 
morphologies through heterochrony and in the maintenance of a vertebrate body plan will be 
discussed. 

Dupanloup Isabelle, Schneider Stefan & Excoffier Laurent 

Département d" Anthropologie et d'Ecologie; Université de Genève; Rue Gustave Revilliod 12; 
CH-1227 Carouge; Switzerland 

Inferring genetic impact of cultural and ecological barriers: a new approach 

Several techniques have been developed to detect the presence of genetic boundaries which can 
be defined as areas where the rate of change of gene frequencies is particularly high. These 
boundaries may correspond to either steep ecological gradients or regions of limited gene flow 
between populations. Sometimes these frontiers are even defined a priori based on non genetic 
factors like ecology, geography or culture. We present here a new method to infer the biolo- 
gical impact of these frontiers on gene flow between populations. 



ZOOLOGIA ET BOTANICA 98 747 



Because the processes acting on different portions of the frontier may be heterogeneous, we 
first divide the frontier into segments of arbitrary sizes and then evaluate the "permeability" of 
each segment independently. For each segment, we compute a statistic equivalent to the 
proportion of genetic distances between samples located on different sides of the barrier that 
are larger than the distances between samples located on the same side of the barrier taking into 
account the observed geographic distances between samples. The confidence intervals of this 
statistic are also evaluated. The program output is a graphical representation of the populations 
and of the frontier; the width of the segments of the frontier is drawn proportional to the 
statistic computed for this segment and to its statistical significance. 

Faucci Anuschka 

Zoologisches Institut; Universität Basel; CH-4051 Basel; Switzerland 

Structure of a hydroid-seaweed community 

Epiphytism is a strategy for opportunistic species to escape high competition in marine hard 
bottom communities. In the present study the seasonal and spatial distribution of epiphytic 
hydroids on three species of the widespread brown algae Cystoseira have been investigated on 
two sites of different water exposure at the rocky shore of Porto Cesareo (Ionian Sea/Italy). The 
addressed questions were: 

1) Is there seasonal pattern in the hydroid community on Cystoseira? 

2) Which factors influence the community structure: a) exposure to wave action? b) substrate 
differences? 

3) Do the more fragile athecate hydroids occur at different sites than thecate hydroids? 
Ordination by Canonical Correspondence Analysis (CCA) was used to summarize the variation 
in species composition and frequency related to site and date. The two sites of different 
exposure are clearly separated and show a seasonal cycle. The algal stems from the exposed 
site are shorter and less colonized than the one from the sheltered site. Exposure to wave action 
has a bigger influence on the community than the differences in algal substrate. Athecate 
hydroids prefer clearly sites with less wave action and higher structured algae, i.e. sites with 
less mechanical stress than thecate hydroids. 

Flook Paul K. & Rowell C. H. F. 

Zoologisches Institut; Universität Basel; Rheinsprung 9; CH-4051 Basel; Switzerland 

Examining hypotheses of grasshopper evolution with molecular sequence data 

We have used nuclear and mitochondrial DNA sequences to examine the systematics of the 
insect order Orthoptera. Phylogenies reconstructed from the molecular data have proved robust 
at different taxonomic levels and have enabled us to make inferences about various aspects of 
orthopteran evolution. An important finding of the work concerns the phylogeny of the super- 
family Acridoidea (true grasshoppers and relatives). Analysis of a large, representative taxo- 
nomic sample indicates that mtDNA sequences of several of the major acridoid lineages have 
diverged to an approximately equal degree. One interpretation of this pattern is that the major 
acridoid groups arose simultaneously during evolution. Alternatively, the observed similarity in 
branch lengths separating members of different lineages may reflect site saturation in the 
mtDNA sequences. We have examined the two possibilities by 
(i) comparing sequence divergence within acridoid lineages; 
(ii) comparing sequence divergence between other orthopteran groups; 
(iiij estimating rates of intra-specific variability; 
(iv) analysing patterns of among site rate variation, and; 
(v) analysing the assumption of substitution rate constancy. 

We conclude that the observed pattern of sequence divergence does reflect a sudden radiation 
of acridoid species. The analyses also indicate that the mtDNA sequences are evolving at an 



748 ZOOLOGIA ET BOTANICA '98 



approximately equal rate, allowing us to make inferences concerning the ages of the main 
acridoid groups. We discuss the implications of these findings for our understanding of ortho- 
pteran evolution and relate the results to existing systematic hypotheses. 

Frey Daniel 

Geobotanisches Institut ETH; Zollikerstrasse 107; CH-8008 Zürich; Switzerland 

RAPDs reveal extensive genetic variability in the apomictic Erigeron annum 

Erigeron annuus is considered an obligate apomictic taxa of the Asteraceae. It is native in 
eastern North America and has successfully spread all over the world, mainly the northern 
hemisphere. Four RAPD primers were used to assess genetic variability in about 800 
individuals from a total of 100 Western European sampling locations. Up to 35 distinct and 
reliable bands were found per primer, most of them polymorphic. Different approaches were 
used to assess intra- and interpopulation variability, such as an analysis of molecular variance 
(AMOVA) and spatial autocorrelation analysis. Preliminary results are presented. 

Galland Nicole, Savolainen Vincent, Salamin Nicolas & Sanjovo Alexandre 

Institut de Botanique systématique; Université de Lausanne; Bâtiment de Biologie; CH-1015 
Lausanne; Switzerland 

Phylogeographic relationships among the polyploid complex Ornithogalum umbella- 
tum (Liliaceae), based on allozyme and RAPD data 

O. umbellatum forms a polyploid complex in Mediterranean and temperate Europe. Several 
former "species" have been recognized as 3 infraspecific entities: morph 1 (2x), morph 2 (3x) 
and morph 3 (4x, 5x, 6x). A close taxon, O. algeriense (6x) was identified as a separate species 
in Northern Africa and Southern Spain. The study of allozyme variation allows to confirm the 
conspecific nature of all diploids; their genetic distances are highly correlated with geographic 
distances, suggesting a trend to geographic speciation. Allozyme patterns of polyploids suggest 
an autopolyploid origin for O. umbellatum, whereas O. algeriense displays a fixed heterozy- 
gosity typical for an amphiploid origin. RAPDs were used to look for clusterings within the 
Ornithogalum complex: their use turned out to be informative for the segretion of O. algeriense 
and the remaining umbellatum s. str. A comparison of the informativity of RAPDs versus 
allozymes is presented in this context. 

Gantenbein Benjamin & Scholl Adolf 

Institute of Zoology; Division of Population Biology; Baltzerstrasse 3; CH-3012 Bern; 

Switzerland 

Allozymes show an unusually high genetic differentiation of Euscorpius germanus 
(Scorpiones: Chactidae) populations 

We examined the genetic population structure of Euscorpius germanus (C. L. Koch, 1837) 
using horizontal starch gel enzyme electrophoresis (18 loci surveyed). We collected eight 
population samples from Switzerland (Valais, Ticino, Grison), 19 samples from northern Italy 
(Lombardia. Trentino, Alto Adige), and two samples from Austria (Carinthia). These samples 
include the subspecies E. g. germanus (C. L. Koch, 1837) and E. g. alpha di Caporiacco, 1950. 
Eight of the north Italian samples come from the Valle Brembana (Alpi Bergamasci), where 
Bonacina (1980) suggested hybridization between the two subspecies, based on asymmetric 
numbers of trichobothria on the pedipalp tibia and of the pectinal teeth. We included four 
samples of E. flavicaudis (de Geer, 1778), two samples of Buthus occitanus (Amoreux, 1789), 
and four samples of Mesobuthus gibbosus (Brulle, 1832) for comparison. The analysis revealed 
two highly differentiated population groups in E. germanus which are separated by the river 



ZOOLOGIA ET BOTANICA 98 749 



Etsch in northern Italy. These population groups are fixed for alternative alleles at eight out of 
18 gene loci and coincide roughly with the subspecies E. g. germanus and E. g. alpha. 
However, there is no evidence of introgressive hybridization from the allozyme data. The 
easternmost population from Austria (Carinthia, Karawanken) indicates a third population 
group that is separated from the former groups by allele substitution at nine out of 18 gene loci. 
In the phenogram (UPGMA cluster analysis using Nei' s (1972) genetic distance), the branching 
points of the E. germanus population groups are found at unusually high distances as compared 
to the outgroup taxa. 

Goudet Jérôme & Savolainen Vincent 

Institut de Zoologie et d'Ecologie Animale & Institut de Botanique Systématique et Géobo- 
tanique; Bâtiment Biologie; Université de Lausanne; CH-1015 Lausanne; Switzerland 

Key innovations and rate of speciation: statistical artefact or real phenomenon? 

Investigations have often shown that key innovations such as phytophagy in insects or nectar 
spurs in angiosperms have led to an increased rate of speciation. Recently, evidence for 
correlation in the rate of microevolution (e.g. number of mutations along the branches in a 
cladogram) and macroevolution (number of species in families connected to these branches) 
was found. Here we review the statistical methods use to put forward these claims and suggest 
a new test. Extended data in angiosperms show that, despite tending toward a positive 
correlation between rate of micro and macroevolution, this relation is severely influenced by 
the taxonomies employed, the phylogenetic sampling and in turn the accuracy of the 
phylogenies. 

Griffin Andrea 

Laboratoire d'Ethologie; Université de Genève; Route des Acacias 54; CH-1227 Carouge; 
Switzerland 

Updating the path integrator through a visual fix 

To survive and to reproduce it is essential for any sedentary animal to know its way around its 
environment. Inded it must be able to relocate reliable food sources, shelter and water, avoid 
predators, find mates and provision offspring. Mammals can navigate through path integration 
(dead reckoning) by updating their position on the basis of internal signals generated during 
locomotion, without using any external references. However, being open to cumulative errors, 
path integration remains functional over short excursions only, unless it is corrected by 
positional information from familiar landmarks. The hypothesis that reference to learned 
landmarks can update the path integrator was examined in golden hamsters {Mesocricetus 
auratus W.) during hoarding excursions occurring in darkness, within a large open arena. The 
subjects proceeded from their peripheral nest to a platform located at one of seven 
feedingssites. During food uptake, they were submitted to more than 10 full rotations. Self- 
generated positional information having been jammed, the animals no longer oriented towards 
the nest. By contrast, when the subjects were rotated at the food source and then briefly 
presented with the familiar visual environment, before the initiation of homing, they returned 
significantly towards the nest. However, their homing performance was less accurate than in 
control trials involving neither rotations nor the opportunity for a visual fix. Our results suggest 
that the visual fix provided the animals with directional, but not positional information. 

Heer Lorenz 

Institute of Zoology; University of Berne; Baltzerstrasse 3; CH-3012 Bern; Switzerland 

Rank-specific mate expenditure of males in the polygynandrous Alpine Accentor 
(Prunella collaris) 



750 ZOOLOGIA ET BOTANICA 



By grouping and cooperating for territory defense, male Alpine Accentor of one breeding unit 
get into conflict for the access to fertile females within their group. Dominant males (alpha, 
sometimes beta) invest most in direct competition (social hierarchy, mate guarding, frequent 
copulations), while subordinate males are mostly restricted therefrom and invest more in the 
indirect competition (song, sitting self-advertisely). Dominant males are present full-time in 
their territory guarding a fertile female. Subordinate males spend only part-time in the territory 
and are preponderant present therein during the time of the 'insemination window'. Accord- 
ingly, dominant males sire most of the young. 

Hoberg Eric P. 1 , Mariaux Jean 2 , Jones Arlene 3 & Bray Rodney A. 3 

1 Biosystematics and National Parasite Collection Unit; USDA; Agricultural Research Service; 
Beltsville, MD 20705; USA 

- Muséum d'histoire naturelle; CP 6434; CH-121 1 Genève 6; Switzerland 

3 Department of Zoology; The Natural History Museum; London SW7 5BD; UK 

Phylogeny of Eucestoda: morphological and molecular congruence 

Advances in our understanding of relationships among the 12 orders of the Eucestoda have 
been achieved based on independent approaches linked to comparative morphology and 
analysis of sequence data from 18s rDNA. Historically, the phylogeny for tapeworms has been 
problematic; numerous conflicting hypotheses have been presented over the past century. 
Recent studies indicate that resolution of relationships among the 1 2 orders appears near at 
hand. Maximum parsimony analysis of morphological and molecular databases yielded largely 
congruent trees supporting monophyly for the Eucestoda. Monozoic Caryophyllidea are the 
basal taxon; difossate forms such as the Pseudophyllidea are primitive; tetrafossate groups 
including the Tetraphyllidea, Proteocephalidea, Nippotaeniidea, Tetrabothriidea and Cyclo- 
phyllidea are highly derived. Hypotheses differed in the placement of the Trypanorhyncha and 
the Diphyllidea. These studies are a robust foundation for resolution of higher-level relation- 
ships among the tapeworms linked to 'total evidence' analyses. Through a top-down approach 
this has also supported phylogenetic studies among families of Cyclophyllidea, the most 
diverse and economically significant group of tapeworms in avian and mammalian hosts. 
Comparative data from morphology, ontogeny and ultrastructure are validated; a comple- 
mentary nature for morphological and molecular approaches is emphasized. Phylogenetic data 
bases for the eucestodes represent model systems for evolutionary biology, cospeciation ana- 
lysis and historical biogeography. 

Holzmann Maria 1 , Piller Werner 2 , Zaninetti Louisette 3 & Pawlowski Jan 4 

1 Institut für Paläontologie der Universität Wien; Austria 

- Institut für Geologie und Palaeontoloaie der Universität Graz; Austria 
J Département de Géologie; Université de Genève; Switzerland 

4 Département de Zoologie et Biologie animale; Université de Genève; Switzerland 

Molecular, morphological and ecological evidence for species recognition in 
Ammonia (Foraminifera) 

The genus Ammonia is one of the most common shallow water benthic foraminifers. Taxo- 
nomic identification of Ammonia species, however, is impeded by their great morphologic 
variability. In the present study, we combined molecular, morphological and ecological data to 
define two Ammonia species, termed Ammonia sp. 1 and Ammonia sp. 2. We obtained partial 
large subunit ribosomal DNA (LSU rDNA) sequences from 42 living specimens collected from 
the Mediterranean Sea, North Atlantic and South Pacific. DNA sequence analysis confirms that 
Ammonia sp. 1 and Ammonia sp. 2 form two genetically distinct species. Cluster analysis of 
their morphological characters shows that both species differ by the size of their tests and size 
of wall perforations. They may also be adapted to different environmental conditions as 
suggested by differences in their distribution in the Lagoon of Venice. 



ZOOLOGIA ET BOTANICA '98 75 ] 



Hosken David 

Zoologisches Museum; Universität Zürich-Irchel; Winterthurerstrasse 190; CH-8057 Zürich: 
Switzerland 

Aspects of sperm competition in yellow dungflies 

Sperm competition occurs when the ejaculates of different males compete to fertilize a given 
set of a females ova. How ejaculates compete is generally inferred from paternity data and 
mathematical modelling rather than by direct observation. Yellow dungflies are particularly 
well studied in this respect. Female dungflies store sperm in fixed volume sperm stores 
(spermathecae), and models of dungfly sperm competition suggest constant random sperm 
displacement with instantaneous mixing from the stores when females copulate more than once, 
with each subsequent male displacing about 80% of the previous stored sperm. However, 
histological evidence suggests some model assumptions are possibly incorrect, and examination 
of the large female accessory glands indicates functions not previously described. Furthermore, 
since males vary consistently in sperm length it is possible to use sperm length variation in the 
female sperm stores as a means of testing the current dungfly sperm competition model: if 
random displacement with instantaneous mixing occurs it is unlikely that the sperm from more 
than two males would be detected in the spermathecae. The variance in sperm length in 
samples from field captured females was compared to expectations based on variance dis- 
tributions generated using sperm length data from field captured males. It appears females 
contain sperm from more males than expected with 80% displacement. 

JoussoN Olivier, Bartoli Pierre & Pawlowski Jan 

Département de Zoologie et Biologie Animale; Université de Genève; CH-1224 Chêne- 
Bougeries; Switzerland 

Use of the ITS rDNA for elucidation of the life cycles of Mesometridae (Trematoda, 
Digenea) 

Identification of larval stages is crucial for elucidating the life cycles of various Digenea. 
However, in many digenean species, the larvae are morphologically indistinguishable and it is 
very difficult to establish the affiliation between the larval and adult stages by using the 
morphological criteria. The molecular methods, based on DNA sequencing or PCR-RFLP 
analysis, can offer a new tool for larval stage identification. In this study, the sequences of 
internal transcribed spacer of the ribosomal DNA (ITS rDNA) were used to identify the 
cercariae of three out of five species of the family Mesometridae {Centroderma spinosissima, 
Elstia stossichianum, and Wardula capitellata). These species differ among the others by the 
number of repeats in the ITS1 region. The phylogeny of Mesometridae was inferred from their 
ITS rDNA sequences. 

Kaupp Andreas 1 , Guggenheim Richard 2 , Wirtz Sabine 2 & Nagel Peter 1 

Universität; Institut für Ntur-, Landschafts- und Umweltschutz / Biogeographie; St. Johanns- 
Vorstadt 10; CH-4056 Basel; Switzerland 
2 Universität Basel; REM-Labor; Bernoullistrasse 32; CH-4056 Basel; Switzerland 

Eggs as a subject of phylogenentic research in Paussinae (Coleoptera: Carabidae) 

The ant nest bettles (Paussinae) are a monophyletic taxon of approximately 450 myrmeco- 
philous species. Together with Metriinae, Ozaeninae and Protopaussinae they form the 
monophyletic paussine complex within the ground beetles. 

Despite the conspicuous structural diversity of the Paussinae, the external characters of the 
adults are insufficient to reconstruct well-founded genealogical relationships of the main 
lineages. The monophyly of the individual tribes and subtribes, however, is proven by several 
synapomorphic character states each. Preliminary studies indicate that the chorion structure of 



752 ZOOLOGIA ET BOTANICA 



the eggs may represent a good character for the phylogenetic reconstruction of the Paussinae. 
We distinguished two different types of chorion: 

A) Chorion thin, without aerenchym, netlike. A similar chorion type 'A' exists in several sub- 
families of the Carabidae representing lineages without close relationship (examined taxa: 
Brachininae (Pheropsophus), Carabinae (Carabus), Paussinae: Paussini (Paussus)). These 
findings indicate that this type of chorion structure represents an ancestral (plesiomorphic) 
character state within the Paussinae. 

B) Chorion thick, with spongiformous intrachorionic aerenchym. So far, this particular struc- 
ture was only found in three tribes of Paussinae (examined taxa: Cerapterini (Cerapterus), 
Heteropaussini (Heteropaussus), Pentaplatarthrini (Pentaplatarthrus). The currently avail- 
able results indicate that this structure represents the first known shared-derived character 
state (synapomorphy) of these three taxa. 

All figured eggs were recovered by dissection of dried or alcohol-preserved specimens. 

Locher Rolf & Baur Bruno 

Department of Integrative Biology; Section of Conservation Biology; University of Basel; St. 
Johanns-Vorstadt 10; CH-4056 Basel; Switzerland 

Is the sperm pool in the hermaphroditic land snail Arianta arbustorum limited? 

Many species of gastropods have different forms of sperm storage which provide the potential 
for sperm competition. However direct evidence for sperm competition is so far lacking, except 
in the simultaneously hermaphroditic land snail Arianta arbustorum. In this species multiple 
mating, multiple paternity and differential male fertilization success has been demonstrated. 
Sperm number is an important feature in sperm competition. We conducted a mating 
experiment to examine how long individuals of A. arbustorum need to recover from sperm 
depletion after a copulation. The results show that a delay of 8 and more days between two 
matings is enough to replenish the sperm reserves entirely. Furthermore, egg laying between 
two matings had no effect on the number of sperm transferred in the second copulation 
regardless the intermating interval. These findings contribute to a better understanding of sperm 
competition and sex allocation in simultaneous hermaphrodites. 

Lugon-Moulin Nicolas, Goudet Jérôme, Brünner Harald, Wyttenbach 

Andréas & Hausser Jacques 

Institute of Zoology and Animal Ecology; Biology Building; University of Lausanne; CH-1015 
Lausanne-Dorigny; Switzerland 

Microsatellites reveal the fine-sacle structure of a hybrid zone in Sorex araneus 
(Insectivora, Soricidae) 

The common shrew {Sorex araneus) is subdivided into numerous chromosomes races and 
several interracial hybrid zones have been discovered. Microsatellites were used to unravel the 
fine-scale genetic structure of a hybrid zone between chromosome races Valais and Cordon 
located in the French Alps. A total of 269 individuals collected between 1992 and 1995 were 
typed for seven microsatellite loci. Several approaches were used to study genetic structuring. 
We introduce the exact G-test to microsatellite data. The exact G-test was recently shown to be 
a powerful test of differentiation for diploid populations. Gene flow is clearly reduced between 
these chromosome races and has been estimated at one migrant every two generations using 
/^-statistics and one migrant per generation using F-statistics. Hierarchical F- and /^-statistics 
are compared and their efficiency to detect inter- and intraracial patterns of divergence is 
discussed. Within-race genetic structuring is significant, but remains weak. F f display similar 
values on both sides of the hybrid zone, although no environmental barriers are found on the 
Cordon side, whereas the Valais side is divided by several mountain rivers. A modified version 
of the classical Multiple Correspondence Analysis (CRT-MCA) is carried out. This analysis 



ZOOLOGIA ET BOTANICA 98 753 



clearly shows the dichotomy between the two races. Further analyses are realized to assess the 
genetic background of karyotypic hybrids to compare it with the genetic background of pure 
parental forms. Our results indicate that these karyotypic hybrids represent the trace of an 
ancient hybridization event. 

Lüscher Beatrice 12 , Grossenbacher Kurt 2 , Güntert Marcel 2 & Scholl 
Adolf 1 

1 Institute of Zoology; University of Berne; Switzerland 

- Natural History Museum Berne; Bernastrasse 15; CH-3005 Bern; Switzerland 

Genetic differentiation of the common toad {Bufo bufo) in the Alps 

Bufo bufo occurs throughout Europe (except in northern Scandinavia) and in parts of northern 
Africa and western Asia. In the Swiss Alps, it is found up to 2200 m above sea level. Three 
subspecies are recognized. Two of them occur in our study area. Bufo b. spinosus is found in 
the Mediterranean region and B. b. bufo in the other parts of Europe, including the north of the 
Alps. We analyse the genetic differentiation of 30 populations of both subspecies in the Alpine 
region, using horizontal starch gel electrophoresis of ten enzymes from tadpoles. For 
comparison, we include three populations from more distant localities: Doupov (Bohemia, 
Czech Republic, B. b. bufo), Genova (Italy, B. b. spinosus), and Aries (Provence, France, B. b. 
spinosus), respectively. 

À very low level of genetic differentiation is found among the populations from the Alpine 
region, including the B. b. bufo population from Bohemia and the B. b. spinosus populations 
from southern Switzerland and Genova. All these populations form a compact cluster in the 
dendrogram (UPGMA, using Nei' s genetic distance in pairwise comparisons of populations as 
matrix for cluster construction). However, the Provence B. b. spinosus population is clearly 
differentiated from this cluster. 

Mizrahi-Meissonnier Liliana, Floor Paul K. & Rowell C. H. F. 

Zoologisches Institut; Universität Basel; Rheinsprung 9; CH-4051 Basel; Switzerland 

Comparison of the utility of different data partitions of the 28S rRNA for the 
reconstruction of caeliferan phylogeny 

We amplified and sequenced 2300 base pairs of the 28S riboromal RNA gene in 50 caeliferan 
species (Insecta: Orthoptera). The sequenced region includes variable regions Dl-D7b variable 
regions (based on the D. melanogaster model of Hancock, 1988), and 4 hypervariable regions. 
After calculating the secondary structure for the orthopteran 28S gene we partitioned the data 
according to pairing properties of different positions. We also partitioned data according to the 
pattern of sequence conservation and reconstructed phylogenies from different data sets. We 
then used objective criteria (e.g. consistency indices, nonparametric bootstrapping) to assess the 
utility of different regions of the 28S gene for phylogenetic reconstruction. Using the most 
informative character set we show that the 28S gene is useful for resolving phylogenetic 
relationships among the basal groups of the caeliferan Acridomorpha. 

Monsutti Alice, Perrin Nicolas, Naciri-Graven Yamama & Goudet Jerome 

Institut de Zoologie et d'Ecologie animale; Université de Lausanne; Bâtiment de Biologie; CH- 
1015 Lausanne; Switzerland 

Selection and life-history responses to size-dependent prédation in Physa acuta 
(Gastropoda) 

In order to investigate the evolutionary response of life-histories to a size-dependent prédation, 
we performed two long-term (20 months) laboratory experiments involving the freshwater snail 
Physa acuta. 



754 ZOOLOGIA ET BOTANICA 



1 . A selection experiment on juvenile length - simulating prédation - resulted in a significant 
response in the line selected for larger size, as well as correlated responses for growth rate, 
size at maturity, and asymptotic size. 

2. The second experiment involved controlled prédation by two flatworm species on artificial 
populations, and also showed a significant increase in juvenile size in the populations under 
prédation. 

3. These two results show that important life-history parameters of the species under study 
may evolve quite rapidly (12 generations) under size-dependent prédation. 

Montoya-Burgos Juan-Ignacio & Pawlowski Jan 

Muséum d'histoire naturelle; 1, route de Malagnou; CP 6434; CH-121 1 Genève 6; Switzerland 

Enlightening the history of Neotropical river systems by using loricariid catfish 
phylogeny 

The catfish family Loricariidae (Siluroidei) includes more than 600 species inhabiting almost 
all Neotropical freshwater systems. The analysis of loricariids phylogeny is used here to 
unravel some aspects of paleohydrogeography in Tropical South America. We have sequenced 
the ITS1 - 5.8S - ITS2 region for several representatives of eight closely related loricariid 
genera. Three lineages have been identified in our phylogenetic tree. Within each lineage, the 
combination of present geographical distribution of species and their phylogenetic relationships 
is discussed. The results are compared to current knowledge on past drainage patterns. 

Mühlhäuser Claudia & Blanckenhorn Wolf U. 

Zoological Museum; University Zürich-Irchel; Winterthurerstrasse 190; CH-8057 Zürich; 
Switzerland 

Female choice for larger males in the dung fly Sepsis cynipsea - Fisher's runaway or 
good genes? 

In Sepsis cynipsea females show a characteristic vigorous shaking behaviour to dislodge the 
male on her back trying to copulate. We investigated the mechanisms of sexual selection in the 
laboratory. In a mate choice experiment we showed that females preferred larger males, and 
that copulations with larger males ensued faster. 

Female choice was expressed in the duration of her shaking. The full sib heritability of head 
width (i.e. body size) was high, as expected. Female shaking duration was also heritable. The 
crucial genetic correlation between male size and female shaking (i.e. choice) was nil, however 
either due to low sample sizes, because it does not exist, or because of a genetic constraint 
suggested by the genetic correlation structure. We have evidence that large body size indicates 
good genes, as large males have a higher larval survival, large females enjoy a higher 
fecundity, and larger individuals of both sexes benefit from a longer adult life. The data support 
the good genes hypothesis as both preference and trait are heritable, and as there are fitness 
benefits other than mating success. Although the genetic correlation of trait and preference was 
not substantiated by our data, we cannot exclude that the Fisher process is also partly 
responsible for the evolution of this mating system. 

Muller Sonia 

Muséum d'histoire naturelle: CP 6434; CH-121 1 Genève 6; Switzerland 

Genetic diversity and relationships in the neotropical catfish genus Ancistrus (Siluri- 
formes, Loricariidae) as revealed by allozyme electrophoresis 

Genetic differentiation was examined among 46 samples of Ancistrus representing 15 putative 
species from the main cis-Andean river systems. Further samples of other Ancistrinae genera 



ZOOLOGIA ET BOTANICA '98 755 



were examined for outgroup comparison. Protein electrophoresis were performed on Poly- 
acrylamid gels using a Pharmacia Phastsystem; the gels were stained for 8 enzyme systems. All 
of the 1 1 presumptive loci analysed are polyallelic. Fixed allelic differences at 4 loci demons- 
trate the existence of two syntopic species which were not diagnosed by morphology. High 
variability is showed in populations of a largely sampled species from Amazonian and Para- 
guayan systems. The genetic distances are compared in regard to taxonomic distinctiveness. 
Hypothesis of phylogenetic relationships are given using phenetic and cladistic methods. 

Nadot Sophie 1 , Creach Jean-Baptiste 1 , Ballard Harvey 2 & Dajoz Isabelle 3 

1 Laboratoire Evolution et Systématique; Université Paris 11; France 

2 Department of Biology; Ohio University; USA 

3 Laboratoire d'Ecologie; Université Paris 6; France 

The evolution of pollen heteromorphism in Viola: a phylogenetic approach 

Pollen heteromorphism (the production within a plant of several pollen morphs that differ in 
aperture number) occurs in 30% of Angiosperm species. Variation in aperture number may be 
selected, because the aperture is the only point of the pollen wall where the pollen tube 
germinates. We have focused on the evolutionary significance of pollen heteromorphism in the 
genus Viola in which about 1/3 of the 500 recorded species are pollen heteromorphic. In several 
species of Viola, the different pollen morphs have different fitnesses: aperture number is 
positively correlated with germination speed, but negatively with life expectancy. We aim to 
study the distribution of pollen heteromorphism in Viola and to understand which selective 
pressures act on its maintenance, using a molecular phylogeny based on ITS sequence data. 
Taxonomic studies on Viola distinguish two main groups: Violets and Pansies, based on 
differences in flower morphology; Pansies are monophyletic, Violets are polyphyletic. Pollen 
heteromorphism has evolved independently at least 7 times in Viola. Variations in sporophytic 
ploidy level are linked to the apparition of pollen heteromorphism in all groups of Violets, not 
in Pansies. The proportion of pollen-heteromorphic species, variance in mean aperture number 
and range of variation of pollen morphs is significantly higher in Pansies than in all Violet 
groups. Together with predictions of the theory about the maintenance of pollen hetero- 
morphism, this suggests that selective pressures may render pollen heteromorphism adaptive in 
Pansies, not in Violets. Our data indicate that its maintenance and development are probably 
contingent upon several prerequisites such as fitness differences between the different pollen 
morphs, pollination conditions and traits of flower morphology, which differ between Pansies 
and Violets. 

Pääbo Svante, Krings Matthias, Poinar Hendrik & Greenwood Alex 

Zoological Institute; University of Munich; PO Box 202136; D-80021 Munich; Germany 

Mitochondrial DNA sequences as a means to deduce human history 

DNA sequences from archaeological and palaeontological finds could potentially contribute 
substantially to our understanding of human history. However, most human remains are devoid 
of endogenous sequences that can be amplified by PCR. In those cases, contamination by 
contemporary human DNA poses serious problems. In the small proportion of remains where 
endogenous DNA exists, chemical degradation and modification make the retrieval of ancient 
DNA sequences difficult. For example, large amounts of oxidative base damage is present in 
most samples and correlate with the inability to amplify sequences. In order to overcome some 
of these problems, we have used HPLC to analyze the state of racemization of amino acids in 
ancient remains. This allows large numbers of samples to be screened in order to identify those 
where the state of conservation of amino acids is suggests that DNA retrieval may be possible. 
By this approach, samples from the Neandertal type specimen were analyzed and a mito- 
chondrial DNA sequence was determined by sequencing clones from short overlapping PCR 
products. Our experience with this and other ancient remains will be reviewed. 



756 ZOOLOGIA ET BOTANICA '98 

Pastorini Jennifer, Forstner Michael R. J., Martin Robert D. & Melnick Don J. 

Anthropologisches Institut; Universität Zùrich-Irchel; Winterthurerstrasse 190;CH-8057 Zürich; 
Switzerland 

Morphology and molecules in conflict: the phylogenetic relationship of Callimico 
within the Callitrichidae 

The New World monkeys are divided into two main groups, Callitrichidae and Cebidae. Recent 
morphological phyletic studies generally place Callimico asthe most basal offshoot within the 
Callitrichidae. By contrast, genetic studies have consistently placed Callimico somewhere 
within the Callitrichidae, not basal to this clade. In addition, the detailed nodal relationships of 
the two tamarin genera, Saguinus and Leontopithecus, remain controversial. A DNA sequence 
data set from a subfragment of the mitochondrial ND4 gene and the tRNA Hls , tRNA Ser , and 
tRNA Leu genes was generated in an attempt to clarify the phylogenetic relationships within the 
Callitrichidae. We extracted DNA from hair or tissue samples from 8 species (Ateles geoffroyi, 
Callimico goeldii, Callithrix jacchus, Cebuella pygmaea, Cebus apella, Leontopithecus rosolia, 
L. chrysomelas, and Saguinus midas), amplified the fragment containing those genes by PCR 
and directly sequenced the template. Additional data from outgroup taxa were available from 
GenBank. The 887 bp sequences were analysed by maximum parsimony, neighbor-joining, and 
maximum likelihood methods. The main results are that the phylogenetic position of Callimico 
is resolved between the marmosets (Callithrix, Cebuella) and the tamarins (Saguinus, Leonto- 
pithecus), while Leontopithecus and Saguinus together form the most basal clade of the Calli- 
trichidae. Combined analyses of all previously published nuclear and mitochondrial gene 
sequences (5201 base pairs) confirm these results. As available molecular evidence indicates 
that Callimico is more closely related to the marmosets than to the tamarins, a reconsideration 
of the morphological evidence in the light of the consensus tree from DNA sequence analyses 
is warranted. 

Pawlowski Jan 1 2 , De Vargas Colomban 1 , Fahrni José 1 & Zaninetti Louisette 1 

1 Département de Zoologie et Biologie animale; Université de Genève; CH-1224 Chêne- 
Bougeries; Switzerland 

2 Muséum d'histoire naturelle; CP 6434; CH-121 1 Genève 6; Switzerland 

Calibrating ribosomal clocks in foraminifera 

Because of their well known fossil record, foraminifera offer unusual opportunity to provide 
temporal dimension to the molecular phylogeny and to study the tempo and mode of molecular 
evolution. In order to test the current palaeontological hypotheses on evolution of foraminifera, 
we have obtained partial SSU rDNA sequences of 50 benthic and 18 planktonic species. By 
comparing the number of substitutions with the divergence times inferred from the fossil record 
we evaluated absolute rates of rDNA evolution for several species. Our study reveals important 
differences in rates of molecular evolution between different groups of foraminifera, ranging 
from 4.0x10-9 subst./site/year in planktonic species to less than 0.1x10-9 subst./site/year in 
some benthic species. However, with few exceptions, the rates are relatively stable within two 
planktonic (Globigerinidae, Globorotaliidae) and one benthic (Soritidae) families, suggesting 
that the local ribosomal clocks may exist in foraminifera. 

Perret Mathieu, Sovalainen Vincent, Chautems Alain & Spichiger Rodolphe 

Conservatoire et Jardin botaniques de la Ville de Genève; Chemin de l'Impératrice, 1; CH-1292 
Chambésy; Switzerland 

Evolution of pollination systems in Sinningieae (neotropical Gesneriaceae); insights 

from molecular phylogenetics 

The study of long-term patterns and processes of organisms diversification is a major challenge 
in evolutionary biology and ecology. In this context, we intend to study the evolution of plant- 



ZOOLOGIA ET BOTANICA '98 757 



pollinator systems within Sinningieae (Gesneriaceae). This tribe, distributed throughout the 
Neotropics, includes herbaceous plant and shrubs which are pollinated by a wide range of 
animals like bees, butterflies, moths, hummingbirds and bats. The systematic, morphological 
diversity and geographical distribution of the Sinningieae are well studied, but little has been 
done to understand how ecological relationships with their pollinators originate and evolve. To 
study these mechanisms, we intend to map the pollination systems onto an accurate phylogeny 
and to trace the evolutionary events that have given rise to the modern distribution of ecological 
festures. Until now, half of the currently described species have been collected for which the 
trnL-trnF and rbcL-atpB chloroplast intergenic spacer has been sequenced. Our intention is to 
present probable evolutionary scenarios (multiple syndromes) that can be inferred from the 
molecular phylogeny, and discuss them in the light of the actual morphological and biogeo- 
graphical knowledge. 

Pozzi Stefano 

Muséum d'histoire naturelle; CP 6434; CH-121 1 Genève 6; Switzerland 

Evaluation of dry grassland management using spider communities 

In Switzerland, the inventory of dry grasslands, with the aim to protect these semi-natural 
habitats, has triggered interest in studying the effects of their management on their fauna. A 
special emphasis is put on spiders, since they are one of the most important groups of terrestrial 
predators, and, thanks to their diversity and abundance, they are known as good bioindicators of 
the ecological state of their habitat. 

In 1995 and 1996, we worked on 40 sites on the Swiss occidental plateau, collecting spiders by 
means of pitfall traps. The evaluation method used takes into account biotop fidelity and the 
rarity of the speices. This method privileges the specificity of the population in relation to its 
present habitat. Therefore, it allows us to not underestimate the quality of the biotops, which 
although very homogeneous, comprise very few species but for whom the ties with their 
surroundings are very close (stenoecious species) and at the same time not to over-estimate the 
stations rich with ubiquitous (euryoecious) species. 

This study allows us to discuss the different management strategies. For dry grasslands 
conservation, it seems important to upkeep them extensively. This operation must be done late 
in the year (autumn). In addition, we recommend to divide the surface of a dry grassland in 
order to alternate the upkeep. This way of management allows us to maintain an abandoned 
part indispensable for some species to carry out their life cycle. The continued study of certain 
stations snowed the capacity of spiders to reflect the effects of management. In future, it seems 
important to use spiders for research in habitat descriptions and evolutions together with 
phytosociological transects. 

RouLiN Alexandre, Richner Heinz & Ducrest Anne-Lyse 

Department of Zoology; University of Berne; Wohlenstrasse 50a; CH-3032 Hinterkappelen; 
Switzerland 

Genetic, environmental and condition-dependent effects on female and male orna- 
mentation in the barn owl Tito alba 

In birds, usually the males only are ornamented. Interestingly, in the barn owl Tito alba both 
female and males display sex-limited plumage traits. Males are commonly lighter coloured, and 
females spottier. A partial cross-fostering experiment tested the relative importance of shared 
genes and a shared environment for the resemblance of related birds. Siblings raised in different 
nests converged towards similar trait values, offspring resembled the true but not the foster 
parents, and plumage traits of unrelated nestlings sharing the same nest were not correlated. 
Results were not inflated by maternal effects detectable in the mother's phenotype, since mid- 
daugther to mother resemblance was not higher than mid-son to father resemblance. This 
suggest the plumage coloration and spottiness are largely genetically inherited traits, and that 



758 ZOOLOGIA ET BOTANICA '98 



the rearing environment has not a strong impag on the expression of these traits. The further 
investigate whether the two sex-limited traits are condition-dependent, brood sizes were mani- 
pulated. Enlargement or reduction of broods by two netlings resulted in lower and higher body 
mass of nestlings respectively. However, nestlings raised in enlarged or reduced broods did not 
show a significantly darker or lighter, or a more or less spotted plumage. No genotype by 
environment interaction was detected. In conclusion, additive genetic variance for plumage 
coloration and spottiness is maintained, and both the rearing environment and boyd condition do 
not account for a large proportion of the phenotypic variance in female and male ornamentations. 

Rowe Tiffany 

70, chemin de la Montagne; CH-1224 Chêne-Bougeries; Switzerland 

Identifying ambiguous landmarks through vector addition 

Path integration (PI) is the process by which an animal uses information about its own 
movements in order to keep track of its current position relative to the starting point of its trip. 
PI and stable spatial information (such as memorized landmarks) complement each other. In 
particular, PI can help the animal solve the problem of locating a goal specified by ambiguous 
landmarks. An experiment was devised to test this. 

Each subject (hamsters) inhabited a round, optically shielded arena with a peripheral nest. 
There were four identical cylinders in a square pattern around the arena centre. One of the 
cylinders (always in the same position relative to the nest) was baited with food, and the animal 
was trained to climb into it and hoard the food. This allowed the animal to establish a long-term 
memory of the location of the baited cylinder relative to the nest. In test trials, the animals was 
lured in darkness along the periphery of the arena, then the lights were switched on, and the 
animal could now see the array of cylinders under a new perspective. Only by keeping track of 
its own movements could the animal identify which, among the four visually indistinguishable 
cylinders, was the correct one. Results indicate that the animal can indeed combine PI to a 
memory of goal position in order to reach the goal from novel points. This is akin to saying that 
the animal can vectorially add its current position vector to the vector stored in long-term 
memory. 

Ruedi Manuel 1 , Smith Margaret F. 2 & Patton James L. 2 

1 Institut de Zoologie et d'Ecologie Animale; UNIL; CH-1015 Lausanne; Switzerland 

2 Museum of Vertebrate Zoology; University of California; Berkeley; CA 94720; USA 

Molecular phylogeography: A glimpse to the past of a Pocket Gopher hybrid zone 

Mitochondrial DNA (mtDNA) variation in the cytochrome b gene was determined for two 
divergent taxa of pocket gophers, Thomomys bottae actuosus and T. b. ruidosae. These two 
taxa hybridize in a narrow contact zone in New Mexico (USA), but introgression of nuclear 
markers such as allozymes or chromosomes does not extend much beyond the hybrid zone. We 
found that despite their distinctness, the two subspecies shared very similar mtDNA haplotype. 
By a comparison of phylogenetic histories derived from nuclear markers (allozymes) and from 
mtDNA haplotypes sampled in different populations of T. bottae from New Mexico, we show 
that apparent similarity is due to an introgression of T. b. ruidosae mtDNA into T. b. actuosus 
nuclear background. Evidence of introgression is not limited to the present-day contact zone 
between these two taxa, but extends at least 75 km away from it. Of several potential 
mechanisms which could lead to such a geographic pattern of variation, we argue that a 
combination of range shifts due to climatic fluctuations, and genetic drift are most likely. We 
also discuss why selection, which might have promoted the spread of one haplotype across the 
hybrid zone, is a less likely explanation. Horizontal gene transfers due to hybridization are not 
uncommon historical events among animals and plants. Although they can be identified with 
careful phylogenetic study using independent data sets, the potential for misinterpreting a gene 
tree as an organismal tree is great. 



ZOOLOGIA ET BOTANICA '98 759 



Savolainen Vincent 1 2 , Chase Mark W. 1 , Morton Cynthia M. 3 , Hoot Sara B. 4 , 
SoLTis Douglas E. 5 , Bayer Clemens 6 , Fay Michael F. 1 , De Bruijn Anette 1 , 
Sullivan Stuart 1 & Qiu Yin-Long 7 

1 Royal Botanic Gardens Kew; Richmond; UK 

2 Conservatoire & Jardin botaniques; Geneva & IBSG; University of Lausanne; Switzerland 

3 University of Reading; UK 

4 University of Wisconsin-Milwaukee; USA 

5 Washington State University; USA 

6 Institut für Allgemeine Botanik; D-22609 Hamburg; Germany 

7 Indiana University; Bloomington 47405; USA 

Phylogenetics of flowering plants based upon a combined analysis of plastid atpB and 
rbcL gene sequences 

In plant molecular systematics, since the broad scale rbcL analysis of Chase, Soltis, Olmstead 
et al, there has been much debates on the use of such large datasets. Recently, Hillis showed 
using simulations based on a new 18S dataset from Soltis et al., that recovering complex phylo- 
genies were surprisingly easier than previously thought. We present here a new phylogenetic 
analyses for angiosperms, based upon the atpB and rbcL chloroplast gene sequences for 358 
taxa representing all major lineages. The results obtained by combining these genes, the 
feasibility of such analyses as well as the support/resolution of these phylogenies are presented. 

Saxer Gerda, Hellriegel Barbara & Reyer Heinz-Ulrich 

Zoologisches Institut; Universität Zürich; Winterthurerstrasse 190; CH-8057 Zürich; Switzerland 

Population dynamics of lynx in relation to its prey - A comparison between Canada 
and Europe 

Periodic fluctuations of mammalian populations and their potential causes have received 
considerable attention. One of the main hypothesis concerning predator-prey dynamics predicts 
that only populations of specialist predators follow the population fluctuations of their prey. 
The lynx (Lynx lynx) populations in Canada and Europe are an ideal system to test this 
hypothesis. The Canadian lynx is a specialist predator and its main prey, the snowshoe hare (L. 
americanus), shows periodic population fluctuations. The European lynx is a generalist with a 
diverse diet including roe deer (C. capreolus), reindeer (Rangifer tarandus) and red deer 
(Cervus elaphus). Therefore the European lynx should not show any periodic or even cyclic 
population fluctuations relative to its prey. I tested this hypothesis by analysing the hunting 
statistics of lynx shot in Norway from 1846-1980 and population size estimates from the lynx 
in Bialowieza Primeval Forest in Poland (1958-1994) with time series analysis. No regular or 
even cyclic fluctuations of the European lynx populations could be detected, in either 
population. These results support the hypothesis that only populations of specialist predators 
follow the population fluctuations of their prey. 

Schneider Stefan & Excoffier Laurent 

Biometry and Genetics Laboratory; Dept. of Anthropology and Ecology; University of Geneva; 
Rue Gustave Revilliod, 12; CH-1227 Carouge; Switzerland 

Estimation of past demographic parameters using the mismatch distribution 

The distribution of pairwise differences between the sequences observed in a sample (the 
mismatch distribution) can be very informative about the recent demographic history of the 
population (Rogers and Harpending, 1992). The theoretical model developed by Li (1977) 
describing the expected mismatch distribution was based on the infinite sites model and 
assumed the homogeneity of mutation rates over the sequence. However in the control region 



760 ZOOLOGIA ET BOTANICA '98 



of human mitochondrial DNA, quite large amount of heterogeneity of mutation rates has been 
observed (Wakeley 1993). In this case, because most mutations occur at a small number of loci, 
the effect of reverse mutations cannot be neglected. To take into account this effect, we have 
extended a model introduced by Yang (1996), to provide the expected mismatch distribution 
under more realistic conditions. We thus explicitly take into account the finiteness of the 
sequence under consideration, the transition bias observed for mtDNA, and up to 4 different 
mutation rates in the sequence. We also provide a way to compute confidence intervals for the 
estimated parameters (population size before and after the expansion and the time elapsed since 
the expansion), as well as for the expected mismatch distribution. We apply this new methodo- 
logy to the case of human mtDNA. 

References: Wakeley, J. 1993. Substitution rate variation among sites in hypervariable region I of human 
mitochondrial DNA. J.Mol.Evol. 37:613-623.; Li, W. H. 1977. Distribution of nucleotide differences between 
two randomly chosen cistrons in a finite population. Genetics 85:331-337.; Rogers, A. R., and H. Harpending. 
1992. Population growth makes waves in the distribution of pairwise genetic differences. Mol. Biol. Evol. 
9:552-569.; Yang, Z. 1996. Statistical properties of a DNA sample under the finite-sites model, genetics 
144:1941-1950. 

Schneiter Beat & Saucy Francis 

Institute of Zoology; University of Fribourg; Pérolles; CH-1700 Fribourg; Switzerland 

Juvenile dispersal in the vole Arvicola terrestris (Rodentia, Arvicolidae) during rainy 
nights 

Abnormally high densities recorded in enclosed populations of small mammals suggest that 
dispersal may be an important component of their population dynamics. For practical reasons, 
dispersal is a difficult phenomenon to address because it is uneasy to distinguish dispersers 
from residents. 

Previous studies conducted using the classical below-ground trapping approach have shown 
that fossorial Arvicola terrestris move mostly over short distances, but high turnover of young 
individuals suggest that our understanding is still incomplete. Taking advantage of the fact that 
this vole usually lives in underground tunnels, we have attempted to catch individuals dis- 
persing above the ground in traps set along drift fences. Observations were conducted in two 
permanent grasslands where we fenced two 50 x 50 m trapping grids. Fences were also estab- 
lished along the edges of neighbouring forests. Two unfenced grids served as controls. From 
March to July 1997, the average density of the below-ground populations increased approxi- 
mately from 70 to about 250 ind/ha. A total of 734 captures and recaptures of 450 A. terrestris 
were recorded along the fences during 121 days of continuous trapping (March 22-July 20 
1997). Interestingly, 91% of the voles caught above the ground were either juveniles or 
subadults, as compared to only 44% in our below-ground samples. No bias in sex-ratio could be 
found among dispersers. More surprisingly, most captures occurred during few rainy nights 
indicating that fossorial A. terrestris disperse en masse above ground during rainy periods. 

Schuchert Peter 

University of Bern; Insel MEM-B814; CH-3014 Bern; Switzerland 

Phylogeny and Classification of the Hydrozoa (Cnidaria) 

Traditional hydrozoan systematics is characterized by a primary dichotomy separating the 
Siphonophora from all other members of the class. The lack of complex characters, the frequent 
homoplasy, and their enormous morphological and life-history variation make the hydrozoans 
not an easy taxon for a phylogenetic analysis. Some progress towards a more natural classific- 
ation has eben made in recent years by recognizing that the Hydrocorallina (hydrozoans with a 
calcareous skeleton) are polyphyletic. In the present work, a phylogenetic analysis and clas- 
sification is attempted. Even the few characters available evidently show that the traditional 
dichotomy Siphonophora versus Hydroidomedusae is no longer acceptable. While some of the 



ZOOLOGIA ET BOTANICA '98 761 



traditional orders like the Narcomedusae, Trachymedusae and the Thecata are recognized as 
monophyletic groups, the order Limnomedusae must be revised and the order Athecata-Antho- 
medusae must be abolished. The Siphonophora clearly belong to a clade named Gastrogonae 
containing also the former members of the Athecata as well as the Hydrocorallina. The 
following classification is proposed: 

HYDROZOA: NARCOMEDUSAE 

MANUBRIOGASTRAE (new name) 
TRACHYMEDUSAE 
HYDROIDEA 

LIMNOMEDUSAE 
STOLONATA (new name) 
THECATA 
GASTROGONAE (new name). 

Spichiger Rodolphe & Savolainen Vincent 

Conservatoire et Jardin botaniques de la Ville de Genève; CH-1292 Chambésy; Switzerland 

Teaching botany in a molecular world 

The application of molecular biology in botany has drastically changed our knowledge in 
systematics and evolution. The most recent systems of classification proposed by Takhtajan, 
Dahlgren, Thorne and Cronquist are questioned by molecular phylogenetics whereas these 
results are not yet fully accepted. However, molecular botany is now close to draw the picture 
of plant phylogeny since large datasets are currently analysed in several institutes. It is 
consequently a difficult period for the teaching of academic botany where modern results have 
to be integrated into the conventional classification. This poster is not a new classification of 
flowering plants since such work will be published later by the angiosperm phylogeny group. 
However we present a general picture of the angiosperms as based on our formal botanical 
course (using idiosyncratic terminology), according to affinities (mainly based on Chase et al.) 
and macroscopic features, and in comparison with the classifications of Cronquist and Thorne. 

Swalla Billie J. 

Pennsylvania State University; Department of Biology; 208 Mueller Laboratory; University 
Park, PA 16802; USA 

Evolution and Development of the Chordate Body Plan 

Metazoan phyla are classified as either protostomes or deuterostomes based on morphological, 
phylogenetic and developmental studies. Deuterostomes have radial cleavage patterns, 
development of the embryonic blastopore into the adult anus and coelomic formation by entero- 
coely. Within the deuterostomes, chordates have a distinct body plan which is thought to have 
evolved from an ancestral deuterostome similar to extant hemichordates or urochordates, but 
studies of chordate evolution within the deuterostome phlya are hampered by the poor fossil 
record left by the soft-bodied ancestors. The urochordate ascidians possess definite chordate 
characteristics as tadpole larvae including a tail containing a dorsal neural tube, notochord, and 
muscle cells flanking the notochord. The head contains the sensory organs, a brain and most of 
the endoderm, or gut. We have used two closely erlated ascdian species with dramatically 
different larval phenotypes to look for genes involved in the specification of the chordate body 
plan during development. One of these genes, manx, appears to be a transcription factor that is 
necessary very early in development for the specification of tissues in the larval tail. 
We have isolated several other maternal genes implicated in specifying the larval body plan, 
including p58 and Cymric. P58 appears to be urochordate specific and is also implicated in 
autonomous muscle development. Recent studies conducted in my laboratory are aimed towards 
establishing a robust phylogeny of the deuterostomes, with special emphasis on the urochordates 



762 ZOOLOGIA ET BOTANICA '98 



and hemichordates. We will use this phylogeny to infer which organisms may closely resemble 
the ancestral chordates in an effort to understand the evolution of chordates. We will study the 
expression of genes we believe are involved in specifying the chordate body plan in extant 
embryos and larvae in an effort to understand which developmentally regulated genes may have 
been co-opted in a non-chordate ancestor to elaborate the chordate phenotype. Current progress 
on these studies will be discussed in my lecture. 

Tourasse Nicolas & Gouy Manolo 

Laboratorie de Biométrie; Génétique et Biologie des populations; Université Lyon I; F-69622 
Villeurbanne cedex; France 

Accounting for evolutionary rate variation among sequence sites consistently changes 
universal phylogenies deduced from rRNA and protein-coding genes 

Identification of the primary lineages of life and of their evolutionary relationships is essential 
for understanding early cellular evolution, particularly the transition between prokaryotes and 
eukaryotes. Most molecular phylogenetic analyses of small subunit (SSU) and large subunit 
(LSU) ribosomal RNA sequences as well as analyses of isoleucyl-tRNA synthetase and of the 
largest subunit of RNA polymerase support the existence of three monophyletic domains, 
Archaea, Bacteria, and Eukarya. In contrast, analyses of elongation factors lalpha/Tu and 2/G 
and of the second largest subunit of RNA polymerase suggested that Archaea are paraphyletic 
and that Eukaryota are specifically related to a subset of Archaea. Crenarchaeota (previously 
eocytes) as advocated by Lake. We have re-analyzed this question using the large numbers of 
sequences now publicly available and recently developed methods of phylogenetic analysis. 
These methods differ mainly from previous ones in using more realistic models of molecular 
evolution which account for the extensive variation among sequence sites of the rate of 
substitution. We report here that this approach gives considerable support to the crenarchaeote - 
eukaryote relationship, both from rRNA and protein sequence data. 

Vizoso Dita & Losada Freddy 

Zoologisches Institut; Universität Basel; Rheinsprung 9; CH-4051 Basel; Switzerland 

Effect of the habitat complexity on the size distribution of marine invertebrates 

The spatial arrangement of the habitat, also known as spatial structure, may affect the body size 
distribution of the inhabitant organisms by constraining the available space. In spite of the 
generalized idea of the existence of such a relationship, methodological problems in the mea- 
surement of habitat complexity have constrained studies in this topic. In the present study, the 
size distribution of diverse and discrete communities of marine invertebrates was analyzed and 
correlated with the spatial complexity of the habitat. This complexity was represented by the 
fractal dimension of the sessile macroalgae that serve as habitat to the communities. The fractal 
dimension offers a direct measure of the ruggedness of a surface, thus allowing the quantitative 
characterization of the habitat complexity. The fractal dimension is scale-invariant. This feature 
allows the characterization of the habitat a different scales without bias in the measure, cover- 
ing all the range of organisms that may be affected and offering a "organismal" point of view. 
The size of the invertebrates present different distributions according to the fractal dimensions 
of the studied macroalgae. 

Wäckers Felix 

ETH-Z; Applied Entomology; Clausiusstrasse 25; CH-8092 Zürich; Switzerland 

Extrafloral nectar production as a herbivore-induced plant defense 

The effect of herbivory on the quantity of extrafloral nectar production, as well as its composition 
and distribution was studied in castor, cotton, and faba bean plants. In ricinus and cotton. 



ZOOLOGIA ET BOTANICA 98 763 



herbivore damage increased the total volume of secreted nectar by a factor 3 and 12 respectively. 
No significant increase due to herbivory could be measured in faba beans. Inductio of nectar 
production was mainly restricted to the damaged leaf. Systemic effects were found in adjacent 
younger leaves only. The increased nectar production could be detected within 16 hours follow- 
ing the onset of herbivore feeding. The induced increase in nectar production ceased within 48 
hours following herbivore removal. 



Wüest Jean 

Muséum d'histoire naturelle; CP 6434; CH-121 1 Genève 6; Switzerland 

The evolution of the pheromone dispersing apparatus in some Hesperidae (Lepi- 
doptera) 

Within the family Hesperidae, we studied the organization of the male pheromone dispersing 
apparatus, which is localized on the forewings, in 3 species, Thymelicus limola, Th. actaeon and 
Hesperia comma. The organization of the apparatus, as well as the morphology of the andro- 
conias, present a growing degree of complexification, perhaps representing the way in which the 
apparatus and androconias were elaborated during evolution of this group. The apparatus is 
formed simply by patches of androconias in Th. limola. In Th. actaeon, around the patches of 
androconias. the adjacent scales are slightly modified and orientated towards the androconial line. 
In H. comma, the patches of androconias are completely covered by the adjacent scales, which 
form a closed space above the androconias. In Hesperidae, the androconias are tubular scales 
containing the pheromone within the hollow medulla. These scales can break into pieces named 
osmophores, which are the dispersing mean of the pheromone. In Th. limola, the scales are 
tubular and non-breakable, but some rare scales present constrictions which can be hypothesized 
as precursors of the dehiscent zones present in the two other species. In Th. actaeon, all the 
androconias present dehiscent zones and break into osmophores, but they often remain unbroken. 
In H. comma, all the androconias are broken and the liberated osmophores are glued together into 
a net just under the roof made of the adjacent scales. 



Wullschleger Esther & Jokela Jukka 

Abteilung Experimentelle Ökologie; ETH Zürich; Clausiusstr. 25; CH-8092 Zürich; Switzerland 

Parasites as selective agents in two host sister taxa: Prevalences of trematode infection 
in molluscan intermediate hosts in dependence of habitat factors 

Adaptation to parasitism can be an important selective factor which determines distribution and 
relative abundances of closely related species. In this study the trematode communities 
infecting two closely related freshwater snails which differ in habitat choice and distribution, 
Lymnaea peregra and L. ovata, were examined in a field survey. 

Trematodes commonly lead to complete castration of their molluscan intermediate host, thus 
the selective pressure imposed by these parasites upon snail populations is strong. The parasite 
life cycle involves further hosts, therefore factors external to snail populations are important to 
its maintenance. 

Several habitat factors were tested for an impact upon parasite prevalences. The surrounding of 
a freshwater habitat, which might determine presence of end hosts, the substrate, which in- 
fluences habitat choice of the snails, and habitat permanence showed the largest effects. 
Habitats with high prevalences were disproportionately often inhabited by L. peregra, the host 
with significantly more infections in total. The prevailing cercarial types were not the same in 
both snail species, suggesting that the snails differ in susceptibility to the various parasite types. 
L. ovata was far more abundant in the lowlands, while L. peregra was more abundant at higher 
altitudes. Sympatric sites were rare, and intermediate snail forms which would suggest hybrid- 
ization were found extremely rarely. 



764 ZOOLOGIA ET BOTANICA '98 



Since parasite prevalences vary across habitat types, the snail species meet differential selection 
pressures for parasite resistance in the varying habitats, and separation through habitat special- 
ization may be enhanced by parasites. 

Wust Saucy Anne-Gabrielle 1 , Hausser Jacques 1 , Taberlet Pierre 2 & Saucy 
Francis 3 

1 Institute of Zoology and Animal Ecology; University of Lausanne; CH-1015 Lausanne; 

Switzerland 
-Laboratoire des Populations d'Altitude; CNRS EP55; University Joseph Fourier; F-3 8041 

Grenoble; France 
3 Institute of Zoology; University of Fribourg; Pérolles; CH-1700 Fribourg; Switzerland 

Phylogeography of the vole Arvicola terrestris as revealed by mtDNA: The role of 
historical factors? 

Arvicola terrestris is a large vole (family Arvicolidae) with a wide geographic distribution 
covering most Eurasia. It is a highly polymorphic species with more than 35 subspecies which 
can be grouped, according to their ecology, into aquatic and fossorial forms. Aquatic animals are 
large and live in wet lowlands, whereas fossorial ones are smaller and live in mountainous areas. 
Aquatic populations, transitionally colonising drier habitat, have often been described. Aquatic 
transitional populations show fossorial behaviour and adopt fossorial life-history traits. The 
origin of these forms which are differentiated by characters such as habitat, body size, colour, 
weight, population dynamics, home range size and mating behaviour is still controversial. 
In order to solve this problem, we studied allozyme and mitochondrial DNA polymorphism for 
aquatic and fossorial animals originating from populations from most parts of the geographic 
distribution of the species. Whereas morphology and habitat suggest some hybridisation between 
forms, the mitochondrial phylogeny, based on 150 sequences of 800bp of the cytochrome b gene, 
distinguish three main groups. Fossorial populations living in mountainous areas of Europe are 
monophyletic. Aquatic populations from the south of the Alps form a separate monophyletic 
clade which has probably emerged first from the aquatic ancestral pool of populations. Finally, 
the remaining populations of northern Europe, England and the east of Eurasia belong to the last 
most diversified clade. The fossorial clade seems to result from a single, more recent, historical 
differentiation coupled with some morphological and behavioural adaptive traits. Phylogeo- 
graphic hypotheses will be discussed with the aim to propose a scenario which could explain the 
actual distribution of populations of Arvicola terrestris in relation to climatic events of the early 
Pleistocene. 

Zehnder Marc 

Institut de Zoologie; Université de Neuchâtel; Rue Emile Argand, 11; CH-2007 Neuchâtel; 
Switzerland 

Molecular phylogenetic analysis of the tapeworm order Proteocephalidea based on 
mitochondrial 16S rDNA sequences 



Systematics of the tapeworm order Proteocephalidea (Eucestoda) whose members are obligatory 
parasites of the alimentary tract of fishes, amphibians and reptiles has until recently been 
addresses using morphological and life-cycle data. These approaches failed so far to yield 
satisfactory results: other sources of characters, DNA sequences in particular, are being explorer 
to better understand the evolution of this tapeworm order. 

I attempt here to infer phylogenetic relationships among a variety of proteocephalidean represent- 
atives using a fragment of the mitochondrial 16S rDNA molecule. A 478 bp sequence was 
obtained for 41 proteocephalidean cestodes as well as for two outgroup species (a tetraphyllid and 
a cyclophyllid). 415 sites were unambiguously aligned, 115 of which were phylogenetically 
informative. Both parsimony and distance method-analyses yielded similar results: 



ZOOLOGIA ET BOTANICA 98 765 



the two traditional families: Monticelliidae and Proteocephalidae are not recognised 
the monophyly of the type genus Proteocephalus is not supported, although a clade consisting 
of palearctic Proteocephalus species only is well defined, thus excluding species from the New 
World. Morphological fetures that distinguish the two groups of Proteocephalus need yet to be 
carified 

resolution of relationships among the remaining Proteocephalidea is poor. Paucity of synapo- 
morphies and high homoplasy indices (HI=0.702) leading to a large number of equally parsi- 
monious trees account for this result. 



Revue suisse de Zoologie 105 (4): 767-770; décembre 1998 




Louis de ROGUIN 

(1948- 1998) 



Un ami nous a quittés le 9 mai 1998 en laissant un grand vide au département 
de Mammalogie et Ornithologie du Muséum d'histoire naturelle de Genève. 

Né le 31 août 1948 à Lausanne, Louis de Roguin fit ses études de sciences 
naturelles à l'université de cette ville après avoir suivi une scolarité classique, latin- 
grec. Licencié es sciences en 1976 (zoologie, botanique, physiologie, biochimie) il 
travaillera plusieurs mois sous mandat pour la «Ligue suisse pour la protection de la 
nature» actuellement «Pro Natura». En janvier 1977, il entre comme collaborateur 
scientifique temporaire au Muséum de Genève et commence une thèse de doctorat 
sous la direction du Prof. W. Huber de l'université de Berne. En 1983 il défend sa 
thèse sur le sujet «Biologie et gestion d'une population de chamois du Jura vaudois» 
et obtient le grade de docteur es sciences de l'université de Lausanne. 

Il est nommé chargé de recherche au Département de Mammalogie et Ornitho- 
logie du Muséum de Genève la même année. 

Personnalité attachante, esprit ouvert, humaniste, Louis de Roguin était tou- 
jours disponible. Ses intérêts multiples l'ont amené à s'intéresser à toutes les branches 
des sciences dites naturelles: botanique, zoologie, écologie. 



768 louis de ROGUIN ( 1 948- 1 998) 

Zoologue de terrain, il a parcouru le Jura durant des semaines en toutes saisons 
à la recherche des chamois, sujets de sa thèse. Dormant souvent à la belle étoile, il 
savait nous transmettre ses émotions lorsqu'il voyait ces animaux, parfois à moins de 
deux mètres de son poste d'observation. 

Homme de laboratoire, il a étudié les micromammifères d'Europe, d'Asie et 
d'Amérique du Sud, et décrit une nouvelle espèce de campagnol d'Iran ainsi qu'une 
sous-espèce de chauve-souris de cette région. Il a également participé à l'étude de la 
faune des micromammifères de sites archéologiques du moyen âge en Rhône-Alpes et 
en Bourgogne (programme PIREN / CNRS). 

«Naturaliste» il maîtrisait la botanique et ses applications dans des domaines 
aussi divers que l'écologie des zones humides ou l'herboristerie. 

Historien, il participa à la publication d'un livre sur le «Voyage aux Antilles et 
au Mexique» de Henri de Saussure et travaillait encore sur le «journal» de cette 
personnalité genevois du 19 e siècle. 

Rédacteur de 1991 à 1997 de la publication scientifique des «Archives des 
Sciences» organe de la société de Physique et d'Histoire Naturelle de Genève, il 
venait de prendre la corédaction de la revue «Le Rhinolophe» du Centre de coordi- 
nation ouest pour l'étude et la protection des chauves-souris. 

Son travail scientifique s'est concrétisé par de nombreuses publications, mais 
hélas, il n'eut pas le temps de réaliser tous ses buts. Dans son bureau, plus d'une 
douzaine de travaux resteront inachevés... 

Louis de Roguin savait écouter sans interrompre et donner un avis sans 
l'imposer. Personne n'hésitait à venir à lui pour toutes sortes de renseignements, de 
conseils, parfois personnels. Nous rendons hommage à cet homme de parole, à ce 
collègue, ce camarade et surtout cet ami... Adieu Louis... 

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d'élevage dans les pays bourguignons et rhodano-alpins (1 er au XVII e siècle), pp. 127- 
134. In: Beck, C. & Delort, R. (éds). «Pour une histoire de l'environnement». Actes 
du programme scientifique et du colloque de mars 1991 sur l'histoire de 
l'environnement et des phénomènes naturels. CNRS Editions, Paris, 272 pp. 

Burnand, J.-D., Cherix, D., Moret, J.-L. & de Roguin, L. 1976. Le marais des Monneaux. La 
Forêt 29 (6): 174-175. 

Burnand. J.-D., Cherix, D., Moret, J.-L. & de Roguin, L. 1977. La végétation du marais des 
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Burnand, J.-D., Cherix, D., Moret, J.-L. & de Roguin, L. 1977. La faune du marais des 
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Burnand, J.-D., Cherix. D., Moret. J.-L. & de Roguin, L. 1978. La faune du marais des 
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Chaix, L., Olive, C, de Roguin, L., Sidi Maamar, H. & Studer, J. (éds) 1995. L'animal dans 
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de Roguin, L. 1983. Connu comme... le chamois blanc. Musées de Genève 235: 2-4. 

de Roguin, L. 1985. Qui sont les Marsupiaux? Musées de Genève 255: 2-6. 

de Roguin, L. 1986. Les Mammifères du Paraguay dans les collections du Muséum de Genève. 
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de Roguin, L. 1988. La cellule domestiquée. Musées de Genève 286: 14-16. 

de Roguin, L. 1988. Notes sur quelques mammifères du Baluchistan iranien. Revue suisse de 
Zoologie 95 (2): 595-606. 

de Roguin, L. 1988. Über einige Gewöllfunde des Uhus Bubo bubo L. aus Nordtirol 
(Österreich). Bericht des naturwissenschaftlich-medizinischen Vereins in Innsbruck 75: 
241-244. 

de Roguin, L. 1989. New historical data on the distribution of Black Rat Rattus rattus (L.) in 
Europe. (Abstract in: Report of 2nd Int. Meeting "Rodents and the Environment", Lyon 
1989). Mammalia 53 (3): 480. 

de Roguin, L. 1990. La microfaune ou la part du rat, p. 81. In: Catalogue de l'exposition du 
Musée des Beaux-Arts et d'Archéologie de Besançon «Se nourrir à Besançon au 
Moyen-Age». Musée Beaux-Arts et Archéologie, Besançon, 84 pp. 

de Roguin, L. 1990. Les micromammifères de Portout, pp. 54-56. In: Pernon, J. & C. (éds). 
«Les potiers de Portout, production, activités et cadre de vie d'un atelier au V e s. ap. 
J.-C. en Savoie». Revue d'Archéologie Narbonnaise, suppl. 20, Editions CNRS, Paris, 
220 pp. 

de Roguin. L. 1990. Mammifères et Oiseaux. In: «Sentier naturaliste. Vallon de la Roulavaz 
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niques, Genève, 89 pp. 

de Roguin, L. 1991. Données historiques nouvelles sur la présence du rat noir Rattus rattus 
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Chabaud, Paris, 362 pp. 

de Roguin, L. 1992. Les micromammifères, p. 135. In: Raynaud, F. (éd.). Documents d'Ar- 
chéologie en Rhône-Alpes No 6: «Le Château et la Seigneurerie du Vuache, Haute- 
Savoie 74». Service Régional d'Archéologie, Lyon, 146 pp. 

de Roguin, L. 1993. Annexe I. La microfaune mammalienne du Castlar de Durfort. Annexe II. 
La microfaune du site de Laval-Basse. In: «Formes et fonctions de l'habitat castrai en 
France méridionale: les apports de la bordure méridionale du Massif Central», pp. 145- 
149. Rapport triennal de synthèse, projet collectif 01 du programme H18 du C.S.R.A., 
208 pp. 

de Roguin, L. 1993. La microfaune commensale (contribution au chap. 2.6. La faune terrestre, 
par Claude Olive), p. 112. In: Colardelle, M. & Verdel, E. (éds). Documents 
d'Archéologie Française, no 40: «Les habitats du lac de Paladru (Isère) dans leur 
environnement. La formation d'un terroir au XI e siècle.» Ed. Maison des Sciences de 
l'Homme, Paris, 416 pp. 



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de Roguin, L. 1995. Le Rat noir Rattus rattus L., pp. 288-292. In: Hausser, J. (éd.). 

Mammifères de la Suisse. Birkhäuser, Basel. 
de Roguin, L. 1995. Le Rat surmulot Rattus norvegicus (Berk), pp. 283-287. In: Hausser, J. 

(éd.). Mammifères de la Suisse. Birkhäuser, Basel. 
de Roguin, L. & Studer, J. 1991. Le rat noir à l'âge du Bronze final. Revue de Paléobiologie, 

Genève 10(1): 79-83. 
de Roguin, L. & Weber, C. 1985. Stenoderma tolteca Saussure, 1860 (Mammalia, Chiroptera): 

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(S) 2466. Bulletin of zoological nomenclature 42 (3): 302-303. 
de Roguin, L. & Weber, C. (éds) 1993. Henri de Saussure: Voyage aux Antilles et au 

Mexique, 1854-1856. Editions Olizane, Genève, 513 pp. 
Justine, J.-L. & de Roguin, L. 1990. Capillaria murissylvatici (Nematoda, Capillariinae), 

parasite d'un rongeur du Baluchistan iranien. Bulletin du Muséum national d'Histoire 

naturelle, Paris, 4 e ser., sect. A. 12(1): 19-33. 
Moeschler, P., Studer, L, Bénier, C, Burckhardt, D., LObl, I., de Roguin, L. & Weber, C. 

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rasti (Saussure). Revue suisse de Zoologie 90 (3): 747-750. 



François J. Baud 



Revue suisse de Zoologie 105 (4): 771-775; décembre 1998 



Description of a new troglophilous species of the genus Maxchernes 
Feio, 1960 (Pseudoscorpiones, Chernetidae) from Brazil 
(Sao Paulo State) 

Volker MAHNERT 1 & Renata DE ANDRADE 2 

1 Muséum d'histoire naturelle, case postale 6434, CH-121 1 Genève 6, Switzerland 

2 Departamento de Zoologia, Instituto de Biociencias da USP, c.p. 1 1461, 
05422-970 Sao Paulo, Brazil. 



Description of a new troglophilous species of the genus Maxchernes 
Feio, 1960 (Pseudoscorpiones, Chernetidae) from Brazil (Sao Paulo 
State). - Maxchernes iporangae n. sp. is described and figured. It occurs 
mainly in the Caverna Alambari de Baixo, Iporanga, in guano of fruit- 
feeding bats. It is compared with the two other known species of the genus, 
M. birabeni Feio from Argentina and M. plaumanni Beier from Brazil. 
Some biological observations are given. 

Key-words: Pseudoscorpiones - Chernetidae - Brazil - caves - guano - 
taxonomy. 

INTRODUCTION 

The cave-dwelling fauna of Brazil is the richest one of South America, at least 
one animal species has been recorded in more than 280 caves out of 1537 explored 
ones (Pinto-da-Rocha 1995). But only one pseudoscorpion species, Pseudochthonius 
strinata, has been upto now mentioned from Brazilian caves (Beier 1969), although 
the presence of this group (in general or identified to family or genus level) had been 
currently recorded (e.g. Trajano & Gnaspini-Netto 1991; Pinto-da-Rocha 1995). 
A large collection of pseudoscorpions of approximately 100 caves (mainly collected 
by Eleonora Trajano, Ricardo Pinto-da-Rocha and Pedro Gnaspini-Netto) is under 
final study by the senior author, but it is necessary to publish this new species 
separately since the junior author has to submit her PhD thesis on the biology of this 
yet undescribed species (Andrade & Gnaspini 1998) till end of this year. 

Chernetidae belonging to other genera are common cave pseudoscorpions in 
Brazil. 

Acronyms used: 

MHNG Muséum d'histoire naturelle, Geneva, Switzerland 

MZUSP Museu de Zoologia Universidade de Sao Paulo, Brazil 



Manuscript accepted 22.09.1998 



772 v - MAHNERT & R. DE ANDRADE 



DESCRIPTION 

Maxchernes iporangae n. sp. Figs 1- 9 

Material: Sao Paulo state, Caverna Alambari de Baixo, Iporanga, in guano of fruit- 
feeding bats, lg. P. Gnaspini-Netto, 16.X.1988: 1 ? (holotype) 3d 1 9 2 tritonymphs (MZUSP 
10298); 2 protonymphs (bred in captivity: from female 65, 2.12.97, and female 56, 12.1.98) 
(paratypes); Gruta das Aguas Quentes, Iporanga, lg. P. Gnaspini-Netto, 2.V.1986: 19 (para- 
type) (MZUSP 10297) (2d 1 9 paratypes in MHNG). 

Description: Carapace and palps yellowish brown; carapace laterally coarsely 
granulate, central parts of pro- and mesozone smooth, posterior margin granulate, two 
distinct, granulate transverse furrows, the subbasal one clearly nearer to posterior 
margin than to medial furrow, no eyes or eye-spots; 1,0-1,1 times as long as broad, in 
middle broadest; setae distinctly davate, 4+1 nearly smooth preocular ones on anterior, 
7-10 on posterior margin; tergites (XI excepted) divided, scaly sculpture, setae davate, 
longer on last tergites, 5-6 setae on posterior margin, III-IX also with a lateral and 
medial anterior one, XI 8-10 (2 medial discal setae); apical lobe of palpal coxa with 3 
marginal and 1-2 discal setae, palpal coxa 19-22, coxa I 10-12, II 13-15, III 18-21, IV 
approx. 32; anterior genital operculum of male with 25-30 long setae (semicircular 
arrangement), of female (fig. 1) with 17-21 setae in central group, genital chamber of 
male with 2-3/2-3 smooth setae, spermatheca of female (fig. 2) with two short tubules 
and a few tiny cribillate plates on them. Sternites divided (excepted III, XI), scaly, setae 
on anterior sternites smooth, VII-XI also with davate setae, III 13-15, 3 suprastigmal 
setae, half-sternite IV 3-5, 1 suprastigmal seta, normally 5-6 setae on posterior margin 
of following ones, VII-X also with a medial anterior seta, XI 6-7 (2 lateral and 2 medial 
discal tactile setae, all relatively short). Chelicera (fig. 3): palm with 6-7 setae (2-3 
apically denticulate), fixed finger with 4 bigger and 3 small teeth, movable finger with 
tooth-like subapical lobe, galea of female with 6 branches, of male (fig. 4) nearly 
smooth, serrala exterior 16-17 blades, flagellum 4 setae (distal one dentate). Palps (figs 
5-7) distinctly granulate, setae short, clavodentate, lateral setae on femur and patella 
longer, dentate; trochanter with well developped dorsal hump, femur 2,8-2,9 times as 
long as broad, patella 2,4-2,5, club 1,74-1,80 times as long as broad, hand with pedicel 
1,7 times (male: 1,5-1,6 times) as long as broad and 1,18-1,24 times longer than finger, 
chela with pedicel 2,8-2,9 times (male: 2,6-2,7) as long as broad; fixed finger with 36- 
38 marginal teeth, 4-7 external and 3-4 internal accessory teeth, movable finger 38-41 
marginal teeth, 4-7 external and 2-4 internal accessory teeth. Trichobothria see fig. 7; 
no long smooth seta on dorsal side of finger, near ist. Claws of legs (figs 8, 9) smooth, 
longer than undivided arolia, subterminal seta on tarsus IV smooth, curved. Leg I: 
femur 1,4-1,6 times as long as broad, patella 2,7-3,0 times as long as broad and 1,46- 
1,51 times longer than femur, tibia 3,7-4,1 times, tarsus 5,1-6,1 times as long as broad; 
leg IV: lateral setae clavodentate, femur+patella 3,6-4,2 times, tibia 4,3-4,8 times, 
tarsus without tactile seta, 4,7-5,4 times as long as broad. 

Measurements in mm (3c?/3 9): Carapace 0.60-0.72/0.55-0.65; palps: femur 
0.57-0.64/0.20-0.24, patella 0.53-0.61/0.21-0.25, hand with pedicel 0.54-0.62/0.32- 
0.37, finger length 0.45-0.50, chelal length 0.92-1.04; leg I: femur 0.18-0.22/0.11- 



A NEW TROGLOPHILOUS SPECIES OF MAXCHERNES 



173 



0.16, patella 0.26-0.31/0.10, tibia 0.27-0.31/0.07-0.08, tarsus 0.28-0.32; leg IV: 
femur+patella 0.48-0.57/0.12-0.16, tibia 0.38-0.43/0.08-0.10, tarsus 0.31-0.36/0.06- 
0.07. 




Figs 1-9 

Maxchernes iporangae n. sp. 1: female genital operculum; 2: spermatheca; 3: chelicera of 
female; 4: galea of male; 5-6: pedipalp of female and chelal hand of male (6b: chaetotaxy and 
granulation partially omitted); 7: trichobothrial pattern; 8: leg I (female); 9: leg IV (female). 
Scale unit 0.1 mm. 



774 v - MAHNERT & R. DE ANDRADE 

Tritonymph: similar to adults; sternite II with 4 central setae; palps: femur 2,6 
times as long as broad (0.46mm/0.18mm), patella 2,1 times (0.42/0.20), chela with 
pedicel 3,1 times as long as broad (0.76/0.25). 

Protonymph: Carapace smooth in posterior half; 26 davate setae (4 on an- 
terior, 6 on posterior margin; a dentate preocular seta on each side); tergites I-X 6 
setae, divided, XI 4; stergites V-IX 6, X-XI 4 setae, on the last sternites the lateral 
setae clearly dentate. Chelicera: palm with 4 smooth setae, galea with 2 apical and 1 
subapical branch, serrula exterior 10 blades, flagellum 4 setae. Palps: femur 2,2 times 
as long as broad (0.2 lmm/0. 09mm), patella 1,8 times (0.19/0.11), chela with pedicel 
3,0 times as long as broad (0.40/0.13), finger shorter than hand without pedicel; fixed 
finger with 19. movable finger with 21 teeth. 

Discussion: We place this new species in the genus described by Feio (1960) 
from Argentina (Jujuy prov.), since it shares the following characters with the type 
species birabeni Feio, 1960: flagellum with 4 setae; trichobothrial pattern {est clearly 
proximad ist; st clearly nearer to t than to sb); long venomous duct; no tactile seta on 
tarsus IV, no tactile setae on tergite XI and absence of discal setae on tergites. It 
differs from both birabeni and plaumanni Beier, 1974 (from Nova Teutonia, Brazil) 
in bigger size and more slender pedipalps, mainly femur and patella. 

KEY TO THE SPECIES OF MAXCHERNES (ADULTS) 

1 Small species, length of femur at most 0.46mm; stout pedipalps (femur 

at most 2.4 times, patella 2.4 times longer than broad) 2 

Bigger species, length of femur 0.57-0.64mm; pedipalps more slender, 
femur 2.8-2.9 times, patella 2.3-2.4 times longer than broad. Brazil, Sao 
Paulo (caves) iporangae n. sp. 

2 Length of palpal femur 0.45-0. 46mm; anterior transverse furrow in 
middle of carapace; serrula exterior with 17 blades. Brazil, Santa 

Catarina plaumanni Beier 

Length of palpal femur 0.40-0. 42mm; anterior transverse furrow in 
basal third of carapace; serrula exterior with 15 blades. Argentina, 
Jujuy birabeni Feio 

BIOLOGY 

Maxchernes iporangae n. sp. is very abundant on frugivorous bat guano piles 
in Alambari de Baixo cave, individuals can frequently be seen wandering around. It 
should be pointed out that Brazilian cave pseudoscorpions, although rather common, 
have been never observed in large numbers. Only one or two individuals are normally 
observed in cave passages and/or on guano piles. The new species seems to be 
restricted to the guano piles of this particular cave, although one single specimen has 
been collected in a cave nearby (Aguas Quentes). Frugivorous bat guano piles of the 
same kind and extension, and including a very similar fauna, have been found in other 
caves from the same region. Nevertheless, M. iporangae n. sp. has not been recorded 
from these other caves so far, in spite of similar collection effort. 



A NEW TROGLOPHILOUS SPECIES OF MAXCHERNES 775 

Considering its biology, iporangae n. sp. seems to fit very well into the gua- 
nobite concept (sensu Gnaspini 1992) - a species restricted to guano deposits in caves. 
Its reproductive and behavioural biology is being studied in laboratory, and is the 
subject of a paper in preparation by the junior author. 

BIBLIOGRAPHY 

Andrade. R. de & Gnaspini, P. 1998. Study of the life history of the cave pseudoscorpion 

Maxchernes sp. (Chernetidae) in the laboratory. Abstracts, XIV International Congress 

ofArachnology, Chicago: 45. 
Beier. M. 1969. Ein wahrscheinlich troglobionter Pseudochthonius (Pseudoscorp.) aus Bra- 
silien. Revue suisse de Zoologie 76: 1-2. 
Beier, M. 1974. Brasilianische Pseudoscorpione aus dem Museum in Genf. Revue suisse de 

Zoologie 81: 899-909. 
Feio, J .L. A. 1960. Consideraciones sobre Chernetinae con la descripcion de Maxchernes 

birabeni genero y especie nuevos (Arachnida, Pseudoscorpiones). Neotropica 5(1959): 

71-82. 
Gnaspini, P. 1992. Bat guano ecosystems. A new classification and some considerations, with 

special reference to Neotropical data. Mémoires de Biospéologie 19: 135-138. 
PlNTO-DA-RocHA, R. 1995. Sinopse da fauna cavernicola do Brasil (1907-1994). Papeis 

Avulsos de Zoologia, Sao Paulo, 39 (6): 61-173. 
Traiano, E. & Gnaspini-Netto, P. 1991. Composiçao da fauna cavernicola brasileira. com 

urna analise preliminar da distribuçao dos taxons. Revista brasileira de zoologia 7(3): 

383-407. 



Revue suisse de Zoologie 105 (4): 777-787; décembre 1! 



New species and records of Masuria Cameron from Nepal 
(Coleoptera, Staphylinidae, Aleocharinae) 

Volker ASSING 

Gabelsbergerstr. 2, D-30163 Hannover, Germany. 



New species and records of Masuria Cameron from Nepal (Coleoptera: 
Staphylinidae: Aleocharinae). - Further data on the distribution of Masuria 
plumbea Cameron, M. picipes Cameron, and M. loebli Pace are presented. 
Three new species are described from Nepal: M. rugosepunctata sp. n., 
M. ancoriformis sp. n.. and M. longicornis sp. n. Primary and secondary 
sexual characters are figured. 

Key-words: Coleoptera - Staphylinidae - Aleocharinae - Masuria - Nepal - 
Himalaya - taxonomy - new species - distribution. 

INTRODUCTION 

The genus Masuria was described by Cameron (1928), according to whom the 
new taxon was near Pronomaea, but separated from that genus especially by the 
anteriorly less produced head and the different mouthparts with three-jointed labial 
palpi, transverse labrum and not bifid ligula. Later the same author attributed the genus 
to the tribe Masuriini (containing only Masuria) and keyed the six species known to 
him, all of them from northern India (Cameron 1939). Recently Pace (1989) revised 
the species of Masuria, describing five new species and referring the former genus 
Oncosomechusa Pace with three species to Masuria as a subgenus. In the revision a 
total of 13 species were recognized, ten in Masuria s. str. and three in Oncosomechusa. 
Almost all the species of Masuria have become known from the Himalayan region of 
northern India (seven species) and Nepal (eight species). The obviously widespread 
M. plumbea Cameron and M. picipes Cameron have been recorded from various 
localities both in India and in Nepal; most species, however, are known only from their 
respective type localities. Very recently, one further species of the subgenus Onco- 
somechusa was described from Gansu, northern China (Pace 1997). 

Though very diverse in external characters such as body shape, length of elytra, 
punctation, colour, microsculpture etc. (see figures in Pace 1989), the genus, parti- 
cularly Masuria s. str., is readily identified by the long labial palpi, whose two terminal 
joints in normal preparation distinctly protrude from below the labrum and especially 
by the morphology of the primary sexual characters. For details regarding the morpho- 
logy of the mouthparts and external characters see Figs 1 a - d and the descriptions by 



Manuscript accepted 23.06.1998 



778 VOLKER ASSING 

Cameron (1928, 1939). These descriptions, however, lack a reference to dimorphisms 
of the secondary sexual characters: tergum VIII is usually of slightly different shape; 
the hind margin of the 9 sternum VIII, apart from being often more strongly convex, is 
characterized by a row of rather long marginal setae and by the presence of 
micropubescence in the central area, whereas in the S sternum IX the posterior setae 
insert at some distance from the hind margin, and the micropubescence is absent. 

The aedeagus, though often remarkably different in size, and the spermatheca 
are somewhat uniform in shape among the species of Masuria s. str. On the other hand, 
the internal structures of the aedeagus, particularly the shape of the base of the fla- 
gellum are characteristic and therefore the most reliable characters for the identification 
of the species. 

Unidentified material of Staphylinidae mainly from the collections of the 
Muséum d'histoire naturelle, Genève (MHNG), but also from the Staatliches Museum 
für Tierkunde, Dresden (SMTD), contained numerous Nepalese specimens of Masuria. 
An examination of this material and a comparison with types of similar species yielded 
three new species. In view of the fact that most species of Masuria are known only 
from their type localities and that the material from almost every locality, where 
Masuria was collected, contained at least one new species it can be assumed that our 
knowledge of this genus, presently comprising 17 species, is far from complete and that 
numerous further species remain to be discovered. 



NEW SPECIES AND RECORDS OF MASURIA 

Masuria (s. str.) plumbea Cameron 

50 ex., Nepal, Rasuwa District. Langtang Khola Valley, 2.5 km E Syabru, 1720- 1730m, 
14.IV. 1985, leg. Smetana (MHNG. cAss). 

The species was previously known from several localities in Northern India 
(Chakrata, Mussoorie, Almora) and one locality in Nepal (Tal) (Pace 1988, 1989). 

Masuria (s. str.) picipes Cameron 

11 ex., Nepal, Rasuwa District, Langtang Khola Valley, 2.5 km E Syabru, 1720-1730m, 
14.IV. 1985, leg. Smetana (MHNG. cAss); 7 ex., Nepal, Annapurna mountains, Lamjung Himal, 
below Taunja Danda. 2350m. 6 V. 1996, leg. O. Jäger (SMTD, cAss). 

Like M. plumbea, M. picipes is apparently widespread and was previously 
known from several localities in Nepal and northern India (Pace 1989). The material 
was compared with two â paralectotypes. 



Masuria (s. str.) loebli Pace Figs 2 a - g 

38 ex., Nepal, Nuwakot District, between Ghopte and Thare Pati, 3200m, 23.IV. 1985, leg. 
Smetana (MHNG, cAss). 



NEW SPECIES AND RECORDS OF MASURIA 



779 




b 





d 



Figs 1 a - d: Masuria plumbea Cameron: labium (a); maxilla (b); mandibles (e); labrum (d). 
Scales: 0.2 mm. 



780 



VOLKER ASSING 






b 




/iM'i'M 




Figs 2 a - g: Masuria loebli Pace: aedeagus in lateral and in ventral view (a); flagellum of internal 
sac (b); spermatheca (c); posterior part of 3 tergum (d) and sternum Vili (e); posterior part of 9 
teraum (f) and sternum VIII (g). Scales: 0.2 mm. 



NEW SPECIES AND RECORDS OF MASURIA 781 

Comments and comparative notes: 

Using the key in Pace (1989), most specimens belonging to this species would 
key out with M. rufescens Cameron, because the pronotum is mostly of the same colour 
as the head and elytra; a brief diagnosis is presented below. M. loebli is distinguished 
from M. rufescens by a less densely punctured head and pronotum, different coloration 
(in P. rufescens the body is reddish with the elytra and abdominal tergum VII darker) 
and by the different shape of the internal structures of the aedeagus (cf. Fig. 25 in Pace 
1989). The specimens indicated above were compared with type material. 

Diagnosis: 

2.9 - 3.6 mm. Size and proportions similar to M. picipes. Colour somewhat 
variable; usual coloration: pronotum and elytra ± reddish or light brown; head at least 
slightly darker, reddish brown to dark brown; abdomen brown to dark brown with the 
tergal hind margins slightly and the apex distinctly lighter; legs and basal antennomeres 
ferrugineous; antennae distally at least slightly darkened. Whole body without distinct 
microsculpture and shining; punctation of forebody distinct, but not very dense, with 
the interstices at least on head and pronotum on average at least as wide as, usually 
wider than punctures; punctation of abdomen sparse and extremely fine. 

Head with eyes in dorsal view approximately as long as temples; antennae 
slender and long, with antennomeres I - III distinctly oblong and subequal in length, 
IV - X gradually decreasing in absolute and relative length, but at least VI still clearly 
oblong, and X weakly transverse. 

Pronotum with weakly pronounced, ± obtuse posterior angles, lateral margins in 
posterior half usually weakly concave. Elytra distinctly wider and at suture (from apex 
of scutellum to hind margin) usually slightly longer than pronotum; hind wings fully 
developed. 

6 : hind margin of tergum VIII weakly pointed posteriorly, that of sternum VIII 
evenly convex (Figs 2 d - e); aedeagus in ventral view broad, base of flagellum shaped 
like an axe (Figs 2 a - b). 

9 : hind margin of tergum VIII weakly pointed, that of sternum VIII more 
strongly convex than in â (Figs 2 f - g); spermatheca as in Fig. 2c. 

Distribution: 

The material indicated above was collected near the type locality in northern 
central Nepal. 

Masuria (s. str.) longicornis sp. n. Figs 3 a - g 

Holotype 6: NEPAL, Khandbari Distr., Induwa Khola Valley, 2050m, 16.IV.1984, Smetana & 

Lobi (MHNG). 

Paratypes: 1 â , 2 9 9 , same data as holotype (MHNG, cAss). 

Derivatio nominis: The name (lat.: with long antennae) refers to one of the characters 

distinguishing this species from the similar M. picipes. 



782 



VOLKER ASSING 




b 



h 





Figs 3 a - h: Masuria longicornis sp. n. (a - g) and M. picipes Cameron (h): aedeagus in lateral 
and in ventral view (a); flagellum of internal sac (b, h); spermatheca (c); posterior part of 
6 tergum (d) and sternum Vili (e); posterior part of 9 tergum (f) and sternum VIII (g). Scales: 
0.2 mm. 



NEW SPECIES AND RECORDS OF MASURIA 783 

Diagnosis: 

3.5 - 3.8 mm. Externally very similar to M. picipes, but of larger, more slender 
and lighter appearance. Colour of body blackish brown to black, with the hind margins 
of the abdominal terga and the margins of the pronotum slightly lighter; legs brown to 
dark brown with the tarsi yellowish; antennae dark brown with antennomere I and often 
parts of II and III lighter. Whole body without distinct microsculpture and shining: 
punctation of forebody distinct and moderately dense, interstices on average less wide 
than punctures; punctation of abdomen fine, much finer than in M. picipes. 

Head with eyes in dorsal view approximately as long as temples or slightly 
shorter; antennae of similar shape as in M. loebli, more slender and longer than in 
M. picipes; antennomere V distinctly oblong (in M. picipes indistinctly oblong or sub- 
quadrate), VI usually weakly oblong (in M. picipes subquadrate or weakly transverse), 
and X subquadrate to weakly transverse (in M. picipes distinctly transverse). 

Pronotum with ± obtuse posterior angles, lateral margins in posterior half 
usually concave. Elytra distinctly wider and at suture (from apex of scutellum to hind 
margin) slightly shorter than pronotum; hind wings fully developed. Legs longer than in 
M. picipes, mesotarsomeres I - IV distinctly oblong (in M. picipes at most weakly 
oblong), length of metatarsomeres I - V 0.5 - 0.6 mm (in M. picipes 0.3 - 0.4 mm). 

S : hind margin of tergum VIII weakly convex posteriorly, that of sternum VIII 
moderately convex (Figs 3 d - e); aedeagus in ventral view slender (Fig. 3a), base of 
flagellum as in Fig. 3b (for comparison with flagellum of M. picipes see Fig. 3h). 

9 : hind margins of tergum and sternum VIII as in Figs 3 f - g; spermatheca as in 
Fig. 3c. 

Comparative notes: 

Using Pace (1989), the species would key out with M. picipes and M. kali Pace. 
For distinction from the former see diagnosis; from the latter P. longicornis is dis- 
tinguished by larger eyes, a more slender pronotum, the much more slender and smaller 
aedeagus and the different shape of the base of the flagellum (cf. Figs 13 - 15 in Pace 
1989). 

Distribution: 

The species is known only from the type locality in eastern Nepal. 

Masuria (s. str.) rugosepunctata sp. n. Figs 4 a - g 

Holotype 6: NEPAL, Khandbari Distr., Induwa Khola Valley, 2050m, 16.IV.1984, Smetana & 

Lobi (MHNG). 

Paratypes: 15 ex., same data as holotype (MHNG, cAss). 

Derivatio nominis: The name (lat.: rugosely punctured) refers to the conspicuously dense, coarse 

and partly rugose punctation, a character only shared with the similar M. plumbea. 

Diagnosis: 

3.0 - 3.5 mm. Externally (size, proportions, punctation, colour) highly similar to 
M. plumbea Cameron, though on average of slightly lighter colour, with smaller eyes, 



784 



VOLKER ASSING 




Figs 4 a - h: Maswïa rugosepunctata sp. n. (a - g) and M. plumbea Cameron (h): aedeagus in 
lateral and in ventral view (a); flagellum of internal sac (b, h); spermatheca (c); posterior part of 
S tergum (d) and sternum Vili (e): posterior part of 9 tergum (f) and sternum VIII (g). Scales: 
0.2 mm. 



NEW SPECIES AND RECORDS OF MASURIA 785 

which in dorsal view are somewhat shorter than temples (in M. plumbea as long as or 
longer than temples), and with slightly less densely punctured abdomen. 

â: tergum and sternum VIII broader and shorter than in M. plumbea (Figs 4 d - 
e); aedeagus of similar shape and size as in M. plumbea, but base of flagellum of 
different shape (Figs 4 a - b); for comparison with flagellum of M. plumbea see Fig. 4h. 

9 : tergum and sternum VIII broader and shorter than in M. plumbea (Figs 4 f - 
g); spermatheca as in Fig. 4c. 

Comparative notes: 

M. rugosepunctata is distinguished from its congeners by the extremely dense 
and coarse punctation, a character which it only shares with M. plumbea; for separation 
from that species see diagnosis. 

Distribution: 

The species is known only from the type locality in eastern Nepal, where it was 
collected together with M. longicornis. 

Masuria (s. str.) ancoriformis sp. n. Figs 5 a - g 

Holotype S: NEPAL, Rasuwa Dis., Langtang Kh. Vail., 2.5km E Syabru, 1730m, 14.IV. 1985. 

A. Smetana (MHNG). 

Paratypes: \6 , 1 9, same data as holotype (o* paratype: 1720m) (MHNG, cAss). 

Derivatio nominis: The name (lat.: shaped like an anchor) refers to the characteristic anchor-like 

shape of the base of the flagellum, which distinguishes this species from the similar M. plumbea. 

Diagnosis: 

3.0 - 3.5 mm. Externally (size, proportions, colour) highly similar to M. plum- 
bea, but punctation of forebody and abdomen less dense, that of head and pronotum 
less coarse than in that species; surface of body therefore more shiny. In addition, eyes 
slightly smaller; pronotum more transverse (1.2 - 1.3x wider than long) than in average 
M. plumbea (usually ca. 1.1 - 1.2x wider than long), and with less distinctly concave 
lateral margins in posterior half. 

S: tergum VIII broader and shorter than in M. plumbea, its hind margin ± 
truncate (in M. plumbea strongly convex); hind margin of sternum VIII evenly convex 
(in M. plumbea almost pointed) (Figs 5 d - e); aedeagus of similar shape and size, but 
ventral process longer and base of flagellum of different shape (Figs 5 a - b). 

9 : tergum and sternum VIII broader and shorter than in M. plumbea; hind 
margin of tergum VIII weakly pointed (in M. plumbea distinctly pointed) (Figs 5 f - g); 
spermatheca as in Fig. 5c. 

Comparative notes: 

For distinction from M. plumbea (and also M. rugosepunctata) see diagnosis. 
From M. parva Cameron from northern India (type specimens examined), M. ancori- 
formis is separated by the slightly larger size, the longer antennae - with antennomere 
IV almost 2x wider than long and V distinctly oblong (in M. parva antennomere IV is 
weakly oblong and V subquadrate) -, the more finely punctured pronotum, the clearly 



786 



VOLKER ASSING 







Figs 5 a - g: Masuria ancoriformis sp. n.: aedeagus in lateral and in ventral view (a); flagellum of 
internal sac (b); spermatheca (c); posterior part of S tergum (d) and sternum Vili (e); posterior 
part of ? tergum (f) and sternum VIII (g). Scales: 0.2 mm. 



NEW SPECIES AND RECORDS OF MASURIA 787 

less dense punctation and pubescence of the elytra, the more distinct punctation of the 
abdomen, and especially by the much larger size of the aedeagus, the longer flagellum 
and the different shape of the flagellar base (cf. Figs 17-19 in Pace 1989). 

Distribution: 

The species is known only from the type locality in northern central Nepal, 
where it was collected together with M. plumbea. 

ACKNOWLEDGEMENTS 

I am indebted to Dr. Ivan Lobi (MHNG), Mr Olaf Jäger (SMTD) and Mr Martin 
Brendell (BMNH) for arranging the loan of the material which this study is based on. 

REFERENCES 

Cameron, M. 1928. Description of a new genus of Staphylinidae (Col.) from India. The Entomo- 
logist's Monthly Magazine 64: 51-52. 

Cameron, M. 1939. The fauna of British India including Ceylon and Burma. Coleoptera 
Staphylinidae. Vol. IV, Part I. London, 410 pp. 

Pace. R. 1988. Aleocharinae dell'Himalaya raccolte da Marc Tronquet e Georges Ledoux 
(Coleoptera Staphylinidae). Bolletino del Museo Civico di Storia Naturale, Verona 14 
(1987): 403-419. 

Pace, R. 1989. Aleocharinae nepalesi del Museo di Ginevra. Parte II. Revisione del genere 
Masuria Cameron (Coleoptera, Staphylinidae). Revue suisse de Zoologie 96: 713-727. 

Pace, R. 1997. Aleocharinae della Cina: Parte I (Coleoptera, Staphylinidae). Revue suisse de 
Zoologie 105: 139-220. 



Revue suisse de Zoologie 105 (4): 789-796; décembre 1998 



Redescription of Compsobuthus rugosulus (Pocock, 1900) 
(Scorpiones, Buthidae) based on specimens from Pakistan 

Wilson R. LOURENÇO* & Lionel MONOD** 
*Laboratoire de Zoologie (Arthropodes), Muséum national d'Histoire naturelle, 
61, rue de Buffon, F-75005 Paris, France, e-mail: arachne@mnhn.fr 
** Muséum d'histoire naturelle, route de Malagnou 1, case postale 6434, 
CH-121 1 Genève 6, Suisse. 

Redescription of Compsobuthus rugosulus (Pocock, 1900) (Scorpiones, 
Buthidae) based on specimens from Pakistan. - A redescription of 
Compsobuthus rugosulus (Pocock, 1900) is given on the basis of 15 spe- 
cimens from Pakistan (9 males and 6 females), 10 of which were collected 
in Hyderabad, the type locality, from where a lectotype is designated. 

Key-words: Scorpiones - Compsobuthus - taxonomy - Pakistan. 

INTRODUCTION 

Compsobuthus rugosulus, was first described by Pocock (1900), as a subspecies 
of Buthus acutecarinatus Simon, based on specimens collected both in Gwalior, India 
and Hyderabad, Sind, which today is in Pakistan. By the time of his publication the 
generic classification of Middle East and Oriental scorpions was still very poor. 
Consequently most of the buthid scorpions were associated to the very large and 
complex genus Buthus Leach. 

In his monographic work, dealing mainly with the scorpions of North Africa, 
Vachon (1949) revised the relationships of several species included in the genus 
Buthus, and proposed a number of different genera, including Compsobuthus for the 
species associated with Buthus acutecarinatus Simon. In the following years, other 
species have been included in the new genus. A more detailed synthesis was proposed 
by Vachon (1952). 

The exact composition of the genus Compsobuthus remains uncertain at present. 
Sissom (1990) proposed 12 species to be included in the genus, with a geographic 
distribution ranging from northern Africa to the Middle East and India. More recently 
Sissom (1994) revised the taxonomic positions of Compsobuthus acutecarinatus 
(Simon), Compsobuthus brevimanus (Werner), Compsobuthus werneri werneri 
(Binila), and described the new species Compsobuthus vachoni. He also discussed a 
supplementary unnamed subspecies of Compsobuthus werneri, thereby attesting to the 
problematical status of species and subspecies within this genus. 



Manuscript accepted 1 1.05.1998 



790 w - R - LOURENÇO & L. MONOD 

For Compsobuthus rugosulus the status of species is accepted by some authors, 
while others only regard it as a subspecies of Compsobuthus acutecarinatus. In our 
opinion Compsobuthus rugosulus is to be considered as a valid species, and we give 
in this paper a precise redescription. 

Compsobuthus rugosulus (Pocock, 1900) Figs 1 to 10 

Syntypes: 4 females (the original description mentions males and females), BMNH. 
1896.12.15.14-17. Gwalior (26.13 N 78.10 E), India and Hyderabad (25.22 N 68.22 E), Sind, 
now in Pakistan. One female syntype from Hyderabad (examined by the senior author) is here 
designated as lectotype. 

Buthus acutecarinatus rugosulus Pocock, 1900: 20; Takashima 1945: 76. 

Buthus acuterinatus rugulosus (sic Birula, 1905: 139; Kraepelin 1913: 127. 

Buthus (Buthus) acutecarinatus rugulosus (sic!): Birula 1910: 173; Birula 1917; 
Werner 1936: 204. 

Compsobuthus rugolosus (sic!): Vachon 1949: 99; Vachon 1952: 219. 

Compsobuthus rugulosus (sic!): Vachon 1966: 211; Habibi 1971 : 43; Farzanpay 
1988:37. 

Compsobuthus rugosulus: Levy et al. 1973: 114; Perez 1974: 23; Levy & Amitai 
1980:60. 

Compsobuthus acutecarinatus rugosulus: Tikader & Bastawade 1983: 169. 

Material studied (deposited in the Natural History Museum, Geneva): Pakistan: 
Hyderabad (type locality), 1/1962 (Anderson leg.): 6 males, 4 females. Kavahari, XII/1958 
(Anderson leg.): 1 male-juvenile, 1 female. Mirpur-Sakro, south of Karachi, 11/1962 (Anderson 
leg.): 2 males. Near Karachi, HI/1962 (Anderson leg.): 1 female. 

Redescription (based on male and female topotypes). 

Morphometric measurements in Table I. 

Coloration. Basically yellowish with some darker reddish brown areas on the 
tergite keels. Prosoma: carapace yellowish with reddish areas over the keels; eyes 
surrounded by black pigment. Mesosoma: yellowish with two longitudinal reddish 
areas and longitudinal reddish brown pigment over longitudinal keels. Metasoma: 
segments I to V yellowish. Vesicle yellowish; aculeus reddish. Venter yellowish. 
Chelicerae yellowish; fingers reddish. Pedipalps: globally yellowish; chela with some 
reddish yellow areas on the articulations. Legs yellowish. 

Morphology. Carapace moderately granular; anterior margin with a feeble 
concavity. Anterior and posterior ocular keels strong; furrows moderate to feeble. 
Median ocular tubercle slightly anterior to the center; median eyes separated by one 
ocular diameter. Three pairs of lateral eyes. Sternum triangular. Mesosoma: tergites 
with moderate to strong granulations. Median keel strong in all tergites. Two latero- 
longitudinal keels, which arise after the posterior ocular keel of carapace, very strong 
in all tergites. Tergite VII pentacarinate; all keels strong. Venter: genital operculum 
divided longitudinally. Pectine: pedinai tooth count 19-19; basal middle lamellae of 
each pecten not dilated. Sternites feebly granular with moderately elongated stigmata; 
two feeble longitudinal keels on each sternite; VII with four keels. Metasoma: seg- 
ments I to IV with 10 keels, crenulate. Segment V with 5 keels. Tegument moderate 
to strongly granular. Telson less granulated than segments, with a long and strongly 
curved aculeus. Subaculear tooth absent, with only a vestigial granule present. 



REDESCRIPTION OF COMPSOBUTHUS RUGOSULUS 



791 




Fig. 1 



Compsobuthus rugosulus, female from Hyderabad in dorsal view. 



792 



W. R. LOURENÇO & L. MONOD 




REDESCRIPTION OF COMPSOBUTHUS RUGOSULUS 



793 




« 



-°0 






Figs 2 to 6: Compsobuthus rugosulus, male from Hyderabad. 2 to 4: Trichobothrial pattern. 
Chela, dorso-external, ventral and internal aspects. 5 and 6: Tibia and femur, dorsal and 
externa] aspects. 

Figs 7 to 10: 7: Pectine. 8: Disposition of the granulations over the dentate margins of pedipalp- 
chela fingers. 9 and 10: Tarsus of leg IV, lateral and ventral aspects. 



794 



W. R. LOURENÇO & L. MONOD 



Cheliceral dentition characteristic of the family Buthidae (Vachon 1963); ventral 
aspect of both finger and manus with long but not very dense setae. Pedipalps: femur 
pentacarinate strongly crenulate; tibia with 7 keels; chelae with 7/8 keels strongly 
crenulate; all faces moderately granular. Movable fingers with 10/11 oblique rows of 
granules. Trichobothriotaxy; A-ß, orfhobofhriotaxy (Vachon 1973, 1975). Legs: tarsi 
with numerous fine setae ventrally. Tibial and pedal spurs present, moderate to strong 
on all legs. 

Females: bigger and more bulk than males (see Table I). General coloration 
similar to that of males, but slightly darker yellowish. Pectines smaller (see Table II 
for variability in the number of teeth). Basal middle lamellae not dilated. 



Table I 

Morphometric values (in mm) of male and female of Compsobuthus rugosulus 

from Hyderabad. 



Male 



Female 



Carapace: 

- length 

- anterior width 

- posterior width 



3,0 
1.9 

3,4 



4,3 
2,8 
4,8 



Metasomal segment I: 

- length 

- width 



1,9 

2,1 



2,4 
2,6 



Metasomal segment V: 

- length 

- width 

- depth 



3,2 
1,5 
1,5 



4,0 

2,2 
2,1 



Vesicle: 

- width 

- depth 



1.2 
1,1 



1,8 

1,5 



Pedipalp: 

- Femur length 

- Femur width 

- Tibia length 

- Tibia width 

- Chelae length 

- Chelae width 

- Chelae depth 



2.5 
0,8 
3,2 
1,3 
5.2 
1,4 
1.3 



3,7 
1,2 
4,1 
1,7 
7,2 
1,8 
1,7 



Movable finger: 
- length 



3,6 



5,2 



REDESCRIPTION OF COMPSOBUTHUS RUGOSULUS 795 

Table II 
Variability of the number of pectines teeth in males and females of Compsobuthus rugosulus 

Males Females 

Number of teeth 

14-15 1 

15-15 1 3 

16-16 1 

16-17 1 

17-16 1 

17-17 2 

17-18 1 

18-17 1 

18-18 2 

19-19 1 



ACKNOWLEDGEMENTS 

We are very grateful to M. Gaillard, Laboratoire de Zoologie Arthropodes, for 
preparing several illustrations, and especially to Prof. John L. Cloudsley-Thompson 
formerly of University College London, for reviewing the manuscript. The junior 
author wishes to thank the "Département des affaires culturelles de la Ville de 
Genève" for financial support. 

REFERENCES 

Birula, A. 1905. Beiträge zur Kenntniss der Skorpionenfauna Persiens. Bulletin de l'Académie 
Impériale des Sciences de St. Pétersbourg, 5 e série 23 (1-2): 1 19-148. 

Birula, A. 1910. Ueber Scorpio maurus Linné und seine Unterarten. Horae Societatis Entomo- 
logicae Rossicae 35: 1 15-192. 

Birula, A. 1917. Arachnoidea Arthrogastra Caucasica. Pars I. Scorpiones. Zapiski Kavkazs- 
kogo Muzeya (Mémoires du Musée du Caucase), Tiflis Imprimerie de la Chancellerie 
du Comité pour la Transcaucasie, sér. A, 5: 253 pp. (in Russian). English translation : 
Birula A., 1964. Anthrogastric Arachnids of Caucasia. 1. Scorpions. Israel Program 
for Scientific Translations, Jerusalem, 170 pp. 

Farzanpay, R. 1988. A catalogue of the scorpions occuring in Iran, up to January 1986. Revue 
Arachnologique 8 (2): 33-44. 

Habibi, T. 1971. Liste de Scorpions de l'Iran. Bulletin Faculty of Science, Tehran University 2 

(4): 42-47. 

Kraepelin, K. 1913. Neue Beiträge zur Systematik der Gliederspinnen. III. A. Bemerkungen 
zur Skorpionenfauna Indiens. B. Die Skorpione, Pedipalpen und Solifugen Deutsch- 
Ost- Afrikas. Mitteilungen aus dem Naturhistorischen Museum 30: 123-196. 

Levy, G. & Amitai, P. 1980. Fauna Palaestina. Arachnida I. Scorpiones. The Israel Academy 
of Sciences and Humanities, Jerusalem, 130 pp. 

Levy, G., Amitai, P. & Shulov, A. 1973. New scorpions from Israel, Jordan and Arabia. 
Zoological Journal of the Linnean Society 52 (2): 1 13-140. 



796 w - R - LOURENÇO & L. MONOD 



Perez, S. M 1974. Un inventario preliminar de los escorpiones de la region paleartica y claves 
para la identification de los generös de la region paleartica occidental. Universidad 
complutense de Madrid, Faculdad de ciencias, Departamento de zoologia, Catédra de 
artrópodos, 7: 1-45. 

Pocock, R. I. 1900. Arachnida. In: Blanford, W. T. (ed.). The Fauna of British India, in- 
cluding Ceylon and Burma. London, Taylor and Francis. 279 pp. 

Sissom, W. D. 1990. Systematics, Biogeography, and Paleontology. Pp. 64-160. In: Pons, 
G. A. (ed.). The Biology of Scorpions. Stanford University Press. 

Sissom, W. D. 1994. Description of new and poorly known scorpions of Yemen (Scorpiones: 
Buthidae, Diplocentridae, Scorpionidae). Fauna of Saudi Arabia 14: 3-39. 

Takashima, H. 1945. Scorpions of Eastern Asia. Acta Arachnologica 9 (3-4) : 68-106. 

Tikader, B. K. & Bastawade, D. B. 1983. The Fauna of India. Vol. 3. Scorpions (Scor- 
pionida: Arachnida). Zoological Survey of India, Calcutta, 671 pp. 

Vachon, M. 1949. Etudes sur les scorpions. Archives de l'Institut Pasteur d'Algérie 27 (1): 
66-100. 

Vachon, M. 1952. Etudes sur les scorpions. Publications de l'Institut Pasteur d'Algérie, Alger: 
482 pp. 

Vachon, M. 1963. De l'utilité, en systématique, d'une nomenclature des dents des chélicères 
chez les Scorpions. Bulletin du Muséum National d'Histoire Naturelle, Paris, 2 e sér. 35 
(2): 161-166. 

Vachon, M. 1966. Liste des scorpions connus en Egypte, Arabie, Israîl, Liban, Syrie, Jordanie, 
Turquie. Irak, Iran. Toxicon 4: 209-218. 

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 Muséum National d'Histoire 
Naturelle, Paris, 3 e 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. 
Comptes Rendus des Séances de l'Académie de Sciences, Paris, sér. D 281: 1597-1599. 

Werner, F. 1936. Reptilien und Gliedertiere aus Persien. Festschrift zum 60. Geburstage von 
Professor Dr. Embrik Strand, 2: 193-204. 



Revue suisse de Zoologie 105 (4): 797-812; décembre 1998 



The jumping plant-lice of Lebanon (Hemiptera: Psylloidea) 

NajlaZEIDAN-GÈZE 1 & Daniel BURCKHARDT 2 

1 Faculté des Sciences (2), Université Libanaise, BP 90656, Jdeidth El Maten, Lebanon. 

2 Naturhistorisches Museum, Auaustinergasse 2, CH-4001 Basel, Switzerland. 



The jumping plant-lice of Lebanon (Hemiptera: Psylloidea). - Based on 
collections made in the last two decades, 35 species are recorded from 
Lebanon. Only four species have been previously recorded from the 
country, two of which are misidentifications ("Psylla pyricola" and "Eu- 
phyllura olivine?', for Cacopsylla bidens and Euphyllura straminea respec- 
tively). The biogeographical composition of the Lebanese fauna is briefly 
discussed and compared to that of the neighbouring countries. 

Key-words: Psylloidea - Lebanon - Middle East - Biogeography. 

INTRODUCTION 

Jumping plant-lice or psylloids are a small group of plant sap-sucking 
sternorrhynchous insects. They are generally highly specific with respect to their 
larval food plants. Unlike the closely related aphids, psylloids are particularly diverse 
in the Southern hemisphere from where they probably originate. The fauna of the 
Palaearctic is highly derived and relatively well-studied. Klimaszewski (1973) lists 
505 species from there, and Gegechkori & Loginova (1990) record 521 species from 
the territory of the former USSR. 

The knowledge of the psylloid fauna of the Middle East is very uneven. 
Following numbers of species have been recorded: Egypt 13 (Samy 1973); Arabian 
Peninsula 52 (Burckhardt & Mifsud 1998); Iraq 3 (Loginova 1974; Ossiannilsson 
1992; Burckhardt & Lauterer 1997); Iran 89 (Burckhardt & Lauterer 1993); 
Turkey 93 (Burckhardt & Önucar 1993; Güclü & Burckhardt 1996); Jordan 33 
(Al-Khawaldeh et al. 1997); Syria 6 (Talhouk 1969; Mustafa 1991); Israel 66 
(Bodenheimer 1937; Halperin et al. 1982; Halperin 1986; Halperin et al. 1988; 
Burckhardt & Halperin 1991; Burckhardt & Lauterer 1997). 

Up to now, only four species were known from Lebanon. Talhouk (1969) 
recorded Homotoma ficus (L.), Psylla pyricola Foerster and Euphyllura olivina 
(Costa); subsequently, the last two species have been recognised as species complexes 
(Burckhardt & Hodkinson 1986; Loginova 1973). Halperin et al. (1982) listed 
Euphyllura straminea Loginova from Lebanon; this is the same species as Talhouk' s 
Euphyllura olivina. Lebanese material of Cacopsylla myrthi (Puton) was mentioned 
by Burckhardt (1989) but without detailing collecting data. 



Manuscript accepted 23.06.1998 



798 



N. ZEIDAN-GÈZE & D. BURCKHARDT 



The present paper aims to fill this gap and reports 35 species from Lebanon 
with collecting dates and localities. Information is added on known host plants and 
general distribution, and a key is provided for the identification of adults. 



Batroun 



Jounìé 
Beyrouth fc^ 




S aida 



10 20 30km 
' i i 



Fig. 1 



Simplified map of Lebanon. The names accord with "Carte du Liban" (Anonymous 1997). 



PSYLLOIDEA FROM LEBANON 799 

MATERIEL AND METHODS 

The specimens were collected by sweeping vegetation with a net, or with pan, 
light or sticky traps. The material is preserved in the collections of the Muséum 
d'Histoire Naturelle du Liban, Faculté des Sciences II de l'Université Libanaise 
(MHNL), the Naturhistorisches Museum Basel (NHMB), and the Muséum d'histoire 
naturelle, Genève (MHNG). Most of the studied material comes from following 
Lebanese regions ("casas"): Kesrouan, Maten, Batroun, Jbeil, Chouf, Aley, Saida and 
the North (Fig. 1). The geographical names are cited according to "Carte du Liban" 
(Anonymous 1997). 

The classification is that of White & Hodkinson (1985) with the changes 
proposed by Burckhardt (1987). The morphological terminology follows mostly 
Ossiannilsson (1992). The treated taxa are arranged in alphabetical order. 

KEY TO ADULTS 

1 Antennal flagellar segments flattened bearing long black setae. Male 
proctiger distinctly 2-segmented. On Ficus carica. 

Homotomidae: Homotoma ficus (L.) 

Antennal flagellar segments more or less cylindrical. Male proctiger 

1 -segmented, sometimes indistinctly subdivided 2 

2 Forewings with vein R+M+Cuj bifurcating into R and M+Cuj; if tri- 
furcating then anal break close to apex of vein Cu lb and metabasitarsus 
with 1 or 2 black spurs. Costal break and/or pterostigma often deve- 
loped. Psyllidae 3 

Forewings with vein R+M+Cuj trifurcating into R, M and Cuj or 
bifurcating into R+M and Cu^ anal break distant from apex of vein 
Cu ]b ; costal break and pterostigma always absent. Metabasitarsus 
without black spurs. Triozidae 24 

3 Metacoxae without meracanthus; trochanteral cavity with weakly scle- 

rotised tubercle. Rhinocolinae 4 

Metacoxae with horn-shaped meracanthus; trochanteral cavity without 
tubercle 5 

4 Forewings without expanded pattern. Male parameres simple, lanceo- 
late. Antenna length/head width ratio more than 1.5. Coronal suture 
fully developed. On Pistacia vera. 

Megagonoscena gallicola Burckhardt & Lauterer 

Forewings with dark, sometimes very faint pattern forming a zig-zag 
band along outer margin. Male parameres with posterior process. On 
Pistacia spp Agonoscena pistaciae Burckhardt & Lauterer 

5 Vertex longer than large. Liviinae: Livia 6 

Vertex larger than long 7 

6 Antennal segment 2 long pear-shaped. Forewings oval. On Juncus spp. 
Livia juncorum (Latreille) 



800 N - ZEIDAN-GÈZE & D. BURCKHARDT 

Antennal segment 2 short, cylindrical. Forewings oblong with sub- 
parallel fore and hind margins. Host plant unknown. 
Livia mediterranea Loginova 

7 Head with large anterior flattened lobes enclosing median ocellus 
which is, therefore, visible only in dorsal view. Euphyllurinae: Euphyllura. . 8 
Head different, either regularly rounded anteriorly, or with separated 
lobes or cones. Median ocellus visible in frontal and/or ventral view 9 

8 Pterostigma of forewings long, more than 3 times the distance between 
the apices of pterostigma and vein Rs; with transverse veins which are 
more or less well-developed. Parameres in profile short with flattened 
anterior lobe in apical two thirds. Apex of female proctiger pointed. On 

Olea europaea Euphyllura straminea Loginova 

Pterostigma short, shorter than twice the distance between the apices of 
pterostigma and vein Rs; usually without transverse veins. Parameres in 
profile long with subparallel fore and hind-margins. Apex of female 
proctiger truncate. On Phillyrea, Olea, Osmanthus spp. 
Euphyllura phillyreae Foerster 

9 Basal metatibial spine always absent. Apical metatibial spurs more or 
less evenly spaced, forming a crown, or grouped and then vertex 
flattened, rectangular with anterior lobes. Metabasitarsus with two 
black spurs. Posterior margin of male proctiger bearing wing-like pro- 
cesses. Aphalarinae 10 

Basal metatibial spine often developed. Apical metatibial spurs always 
grouped. Vertex trapezoidal; head with genal cones 12 

10 Proepimeron much larger than proepisternum; propleural suture dia- 
gonal. On Tamarix spp Colposcenia kiritshenkoi Loginova 

Proepimeron and proepisternum subequal; propleural suture vertical. 
Craspedolepta 11 

1 1 Antennae 10-segmented. On Leontodon autumnale. 

Craspedolepta sonchi (Foerster) 

Antennae 9-segmented. On Achillea, Anthémis, Pyrethrum spp. 
Craspedolepta pontica Dobreanu & Manolache 

12 Metabasitarsus without or with a single black spur 13 

Metabasitarsus with two black spurs 15 

13 Male proctiger with posterior wing-like lobes. Forewings with long cell 
m 1+ 2 and high cell cu la , vein Cu lb longer than Cuj. On Acacia, Albizia 

spp Acizziinae: Acizzia uncatoides (Ferris & Klyver) 

Male proctiger without posterior wing-like lobes. Forewings with shor- 
ter and lower M 1+2 and Cu ]a cells, vein Cu lb shorter than Cuj. 
Arytaininae p. p 14 

14 Genal cones about half vertex length. On Calicotome spp. 

Arytainilla cytisi (Puton) 

Genal cones longer than vertex length. On Spartium junceum. 
Livilla spectabilis (Flor) 



PS YLLOIDEA FROM LEBANON 80 1 

15 Forewings with long cell m, +2 and high cell cu ]a . Male parameres 
truncate apically. On Astragalus spp. 

Arytaininae: Cyamophila stoklosai Klimaszewski & Lodos 

Forewings with shorter and lower m 1+0 and cu la cells. Male parameres 
different. Psyllinae 16 

16 Forewing membrane yellow, bearing irregularly, densely spaced sur- 
face spinules. Psylla 17 

Forewing membrane colorless or with brown pattern; surface spinules 
sparse forming regular rhombi or squares, or reduced to narrow stripes 

in the middle of the cells. Cacopsylla 18 

17 Genal processes more than half vertex length. Antennae less than 2.0 

times head width. On Ostrya carpini/olia Psylla colorata Low 

Genal processes less than half vertex length. Antennae more than 2.0 
times head width. On Alnus spp Psylla foersteri Flor 

18 Pattern of forewings consisting of well-defined dark patches at the 
apices of pterostigma and veins Rs, M 1+2 , M 3+4 , Cu la and Cu lb , as well 
as a more or less interrupted band from apex of vein Rs to the middle of 

vein Cu la . On Cercis siliquastrum Cacopsylla pulchella (Low) 

Pattern of forewings different 19 

19 Forewings with brown band along outer margin. On Crataegus spp. 

Cacopsylla mariannae (Baeva) 

Forewings without brown band along outer margin 20 

20 Surface spinules present in all cells forming broad fields; apart from 
narrow stripes along the veins, covering the whole surface of cell c+sc; 
spinules present in basal part of cell rs proximal to bifurcation of vein 

R; fields in apical part not tapering towards wing margin 21 

Combination of characters absent 22 

21 Hindmargin of parameres, in lateral view, convex. Dorsal margin of 
female proctiger weakly convex. On Salix spp. 

Cacopsylla brunneipennis (Edwards) 

Hindmargin of parameres, in lateral view, concave. Dorsal margin of 
female proctiger weakly concave. On Salix spp. 
Cacopsylla subpropinqua (Loginova) 

22 Forewings with brown clavus. On Pyrus spp Cacopsylla bidens (Sulc) 

Forewings with clavus of the same colour as surrounding membrane 23 

23 Surface spinules forming broad fields in apical half of forewings. On 

Pyrus spp Cacopsylla pyrisuga (Foerster) 

Surface spinules forming narrow stripes in apical half of forewings. On 
Rhamnus spp Cacopsylla myrthi (Puton) 

24 Forewings with vein R+M+Cuj bifurcating in veins R+M and Cuj. 

Host plant unknown Eutrioza opima Loginova 

Forewings with vein R+M+Cuj strictly trifurcating into R, M and Cuj 25 

25 Metatibiae with 1+3 black apical spurs. Trioza p. p 26 

Metatibiae with 1+2 black apical spurs 27 



g02 N - ZEIDAN-GÈZE & D. BURCKHARDT 

26 Forewing hind margin with conspicuous dark spots in the middle of 
the cells. Parameres truncate apically. Female genitalia short, truncate 

apically. On Rhamnus alaternus Trioza marginepunctata Flor 

Forewings without conspicuous dark spots in the middle of the cells 
along the hind margin. Parameres with small forward directed hook 
apically. Female genitalia long, pointed apically. On Urtica spp. 
Trioza urticae (L.) 

27 Bifurcation of vein M of forewings distinctly distal to line from apices 

of veins Rs to Cu la 28 

Bifurcation of vein M of forewings on, or proximal to line from apices 

of veins Rs to Cu la 31 

28 Vein Rs of forewings sinuous to almost straight; vein M ]+2 more than 
twice the length of vein M 3+4 ; fore margin of forewings strongly 
arched, hind margin only weakly curved. Male proctiger with large 
posterior lobes. Female proctiger long. On Rubus spp. 

Phylloplecta trisignata (Low) 

Vein Rs of forewings concave; vein M| +0 less than twice as long as 
M 3+4 ; fore margin of forewings only slightly stronger curved than hind 
margin. Hind margin of male proctiger weakly produced. Female geni- 
talia short. Trioza p. p 29 

29 Forewings narrow, more than 2.7 times as long as wide, without surface 

spinules in apical half. On Laurus nobilis Trioza alacris Flor 

Forewings wide, less than 2.7 times as long as wide, with surface spinules. 30 

30 Surface spinules present in all cells of forewings, forming large fields. 

On deciduous Quercus spp Trioza remota Foerster 

Surface spinules largely reduced in apical half of forewings. On ever- 
green Quercus ilex Trioza ilicina (de Stefani) 

31 Antennal segments 4-7 light. Body coloration green or yellow. 
Posterior lobes of male proctiger large, semicircular and bearing long 
marginal setae. Female proctiger with pointed posterior process, with 
long dorsal setae ending almost at apex of proctiger. On Elaeagnus 

angusti/olia Trioza neglecta Loginova 

Antennal segments 4-7 dark brown or black. Body coloration in mature 
specimens with dark parts. Posterior lobes of male proctiger small, 
without long marginal setae. Female proctiger short, subacute or blunt 
apically, without very long dorsal setae. Bactericera 32 

32 Forewings bearing surface spinules. On Salix spp. 

Bactericera albiventris (Foerster) 

Forewings without surface spinules in apical half 33 

33 Genal cones longer than half vertex length. On Salix spp. 

Bactericera curvatinervis (Foerster) 

Genal cones shorter than half vertex length. Polyphagous. 
Bactericera nigricornis (Foerster) 



PSYLLOIDEA FROM LEBANON 803 

LIST OF SPECIES 

HOMOTOMIDAE 

Homotoma ficus (L.) 

Material examined. Jabal Moussa: Kesrouan, 15.VI.1997; Naas: Maten, 
14.VI.1997; Darh El Souan: Maten, 16.VII.1997; Meyrouba: Kesrouan, 24.IX.1994 
(MHNL, NHMB, MHNG). 

Distribution. Albania, Algeria, Armenia, Austria, Azerbaijan, Bulgaria, France, 
Georgia, Great Britain, Greece, Israel, Italy, Jordan, Lebanon, Morocco, Portugal, 
Russia (European part), Spain, Switzerland, Syria, Turkey, USA (California), former 
Yugoslavia (Al-Khawaldeh et al. 1997; Burckhardt 1989; Gegechkori & 
LoGiNOVA 1990). Fig is probably native to SW Asia from where it spread early to the 
Mediterranean; Egyptians were cultivating it 4000 BC (Mabberley 1990). It is likely 
that the host specific H. ficus has a similar history of distribution. 

Host plant. Ficus carica L. (Moraceae). 

Comments. The larvae can cause an irregular growth of the leaves, but the 
species has not been reported to be of economic importance (Burckhardt 1994). 

Psyllidae: Acizziinae 

Acizzia uncatoides (Ferris & Klyver) 

Material examined. Naas: Maten (MHNL). 

Distribution. Australia, introduced into Algeria, Chile, France, Israel, Italy, 
Mexico, New Zealand, Portugal, USA (California, Hawaii) (Burckhardt 1989). 

Host plants. Acacia, Albizia spp. (Fabaceae). 

Comments. A. uncatoides damages its hosts by honeydew secretion and by 
sucking on young tissue which produces necrosis of flower racemes and young twigs 
(Burckhardt 1994). 

Psyllidae: Arytaininae 
Arytainilla cytisi (Puton) 

Material examined. Kfar Debiane: Kesrouan, 7.X.1984; Laklouk: Kesrouan, 
10.VI.1986; Yahchouch: Kesrouan, 16.VI.1981 and 2.V.1983; Dahr El Souan: Maten, 
I.V. 1983 and 22.V.1983 (MHNL). 

Distribution. Algeria, France, Greece, Israel, Italy, Jordan, Spain, Turkey, 
former Yugoslavia (Al-Khawaldeh et al. 1997; Burckhardt 1989). 

Host plants. Calicotome spp. (Fabaceae). 

Cyamophila stoklosai Klimaszewski & Lodos 

Material examined. Kfar Debiane: Kesrouan, 8.VII.1985 (MHNL; NHMB; 
MHNG). 

Distribution. Turkey (Burckhardt & Önucar 1993). 
Host plants. Astragalus spp. (Fabaceae). 



g04 N. ZEIDAN-GÈZE & D. BURCKHARDT 

Livilia spectabilis (Flor) 

Material examined. Dahr El Souan: Maten, 26.1.1981 and 23. VII. 1986; 
Chabrouh: Kesrouan, 22.XI.1983; Ghazir: Kesrouan, 2.1.1982; Hyata: Kesrouan, 
27.VII.1995 and 2.VIII.1996; Kartaba: Kesrouan, 22.IV. 1985; Meyrouba: Kesrouan, 
24.IX.1994; Kfar Abida: Kesrouan, 23.VI.1995 (MHNL; NHMB; MHNG). 

Distribution. Algeria, France, Greece, Italy, Portugal, Spain, Switzerland, 
former Yugoslavia (Burckhardt 1989). 

Host plant. Spartium junceum L. (Fabaceae). 

Psyllidae: Aphalarinae 
Colposcenia kiritshenkoi Loginova 

Material examined. Janné: Jbeil, 30.X.1985 (MHNL; NHMB). 
Distribution. Armenia, Azerbaijan, Bulgaria, Georgia, Iran, Russia (European 
part), Turkey (Gegechkori & Loginova 1990). 
Host plants. Tamarix spp. (Tamaricaceae). 

Craspedolepta pontica Dobreanus & Manolache 

Material examined. Meyrouba: Kesrouan, 5.V.1985; Yahchouch: Kesrouan, 
16.VI.1981 (MHNL). 

Distribution. Armenia, Azerbaijan, former Czechoslovakia, Georgia, Greece, 
Israel, Kazakhstan, Kyrgyzstan, Romania, Russia (European part), Tajikistan, Turkey, 
Turkmenistan (Gegechkori & Loginova 1990). 

Host plants. Achillea, Anthémis, Pyrethrum spp. (Asteraceae). 

Craspedolepta sonchi Foerster 

Material examined. Bhamdoun: Aley, 21. IX. 1979 (MHNL). 

Distribution. Armenia, Central Europe, Denmark, Finland, Georgia, Great 
Britain, Norway, Russia (European part, Dagestan, Siberia, Maritime Territory), 
Sweden (Ossiannilsson 1992). 

Host plant. Leontodon autumnale L. (Asteraceae). 

Comments. Literature records my concern a complex of closely related species 
(Lauterer & Burckhardt, in prep.). 

Psyllidae: Euphyllurinae 
Euphyllura phillyreae Foerster 

Material examined. Nahr-Ibrahim: Kesrouan, 4.V.1985; Meyan: Jbeil, 23. IV. 
1983 (MHNL). 

Distribution. Algeria, Bulgaria, France, Georgia, Greece, Israel, Iran, Italy, 
Morocco, Russia (European part), Spain, Tunisia, Turkey, former Yugoslavia 
(Burckhardt 1989; Gegechkori & Loginova 1990). 

Host plants. Phillyreae, Olea, Osmanthus spp. (Oleaceae). 



PSYLLOIDEA FROM LEBANON 805 

Euphyllura straminea Loginova 

Material examined. Bchamoun: Chouf, 9.III.1981 (MHNL). 

Distribution. Cyprus, Iran, Iraq, Israel, Jordan, Lebanon, Turkey (Al- 
Khawaldeh et al. 1997). 

Host plant. Olea europaea L. (Oleaceae). 

Comments. The presence of E. olivina in Lebanon recorded by Talhouk 
(1969) is unlikely. E. olivina is Western, E. straminea Eastern Mediterranean in dis- 
tribution (Burckhardt 1994). 

Psyllidae: Liviinae 
Livia juncorum (Latreille) 

Material examined. Dahr El Souan: Maten (MHNL). 

Distribution. Widely distributed throughout the Palaearctic (Ossiannilsson 
1992). 

Host plants. Juncus spp. (Juncaceae). 

Livia mediterranea Loginova 

Material examined. Kfarkatra: Chouf, 22.11.1981 (MHNL). 
Distribution. Algeria, Bulgaria, Caucasus, Crimea, Israel, Italy, Spain (Conci 
et al. 1993). 

Host plant. Unknown 

Psyllidae: Psyllinae 

Cacopsylla bidens (Sulc) 

Material examined. Bhamdoun: Aley, 10. VIII. 1978 and 21. IX. 1979; Kfar- 
Katra: Chouf, 8.III.1981; Bhannés: Maten, 4.IV.1980 and 23.VII.1986; Dahr El 
Souan: Maten, 17.VII.1997; Hajar El Afrit: Kesrouan, 16. IX. 1994; Hrajel: 
Kesrouan, 27. VI.1994; Kfar Debiane: Kesrouan, 19.VII.1994 (MHNL, NHMB). 

Distribution. Armenia, Central Asia, Crimea, France, Greece, Iran, Israel, 
Italy, Jordan, Ukraine and introduced into Argentina and Chile (Al-Khawaldeh et al. 
1997). 

Host plants. Pyrus spp. (Rosaceae). 

Comments. The record of Psylla pyricola from Lebanon (Talhouk 1969) 
probably concerns Cacopsylla bidens (Burckhardt & Hodkinson 1986). 

Cacopsylla brunneipennis (Edwards) 

Material examined. Laklouk: Kesrouan, 10.VI.1986 and 22.VI.1986; Yah- 
chouch: Kesrouan, 16.VI.1981; Zaarour: Maten, 1. VI. 1997; Naas: Maten, 19. VI. 1997; 
Broumana: Maten, 14.VI.1997 (MHNL, NHMB, MHNG). 

Distribution. Armenia, Austria, Bulgaria, Czech Republic, Denmark, England, 
Finland, France, Georgia, Ireland, Italy, Kazakhstan, Norway, Poland, Romania, 



806 N - ZEIDAN-GÈZE & D. BURCKHARDT 

Russia (European part, Siberia), Scotland, Slovakia, Spain, Sweden, Switzerland, 
Ukraine, former Yugoslavia (Lauterer & Burckhardt 1997). 
Host plants. Salix spp. 

Cacopsylla mariannae (Baeva) 

Material examined. Naas: Maten, 14.VI.1997; Zaarour: Maten, 28.VI.1997; 
Meyrouba: Kesrouan, 24. IX. 1994; Hrajel: Kesrouan, 27.VI.1994; Kfar Debiane: 
Kesrouan, 26.V.1985 and 8.VII.1985; Hajar el Afrit: Kesrouan, 24.IX.1994 and 
16.IX. 1994; Achkout: Kesrouan, 17.IX.1994; Baskinta: Kesrouan, 22.IX.1994 
(MHNL, NHMB, MHNG). 

Distribution. Tadzhikistan, Turkey (Gegechkori & Loginova 1990; 
Burckhardt & Önucar 1993). 

Host plants. Crataegus spp. (Rosaceae). 

Cacopsylla myrthi (Puton) 

Material examined. Jaj: Batroun, IV-VI.1984; Dahr El Souan: Maten, 23. VII. 
1986; Naas: Maten, 14.VI.1997; Zaarour: Maten, 1. VI. 1997; Achkout: Kesrouan, 
17.IX.1994; Baskinta: Kesrouan, 19. IX.1994; Hajar El Afrit: Kesrouan, 17.IX.1994, 
24.IX.1994 and 16.IX.1994; Hrajel: Kesrouan, 27.VI.1994; Jabal Moussa: Kesrouan, 
15.VI.1997 and 21. VI. 1994; Kartaba: Kesrouan, 22.XI.1985; Meyan: Kesrouan, 
23.IV. 1983; Meyrouba: Kesrouan, 5.V.1985 and 24.IX.1994; Yahchouch: Kesrouan, 
16.VI.1981; Becharré: Nord, 20.V.1987; Tannourine: Batroun, 31.V.1997; Saida: 
Saida, 28.X.1978 (MHNL, NHMB, MHNG). 

Distribution. Algeria, Crimea, France, Greece, Israel, Italy, Jordan, Spain, 
Turkey (Al-Khawaldeh 1997). 

Host plants. Rhamnus spp. (Rhamnaceae). 

Cacopsylla pulchella (Low) 

Material examined. Jaj: Batroun 4.V.1984; Naas: Maten, 8.VI.1997 and 
18. VII. 1997; Hajar El Afrit: Kesrouan, 24.IX.1994; Jouret Bedrane: Kesrouan, 
2.V.1983; Jabal Moussa: Kesrouan, 15.VI.1986, 14.VI.1997 and 15.VI.1997; Tan- 
nourine: Nord, 30.V.1997 (MHNL, NHMB, MHNG). 

Distribution. Austria, France, Great Britain, Greece, Italy, Russia (European 
part), Switzerland, Turkey, former Yugoslavia (Gegechkori & Loginova 1990). 

Host plant. Cercis silìquastrum L. (Fabaceae). 

Comments. The assignment of C. pulchella to Cacopsylla needs to be revised 
(Burckhardt 1994). 

Cacopsylla pyrisuga (Foerster) 

Material examined. Hajar El Afrit: Kesrouan, 24.IX. 1994; Achkout: Kesrouan, 
17.IX.1994; Jabal Moussa: Kesrouan, 15.VI.1997 and 21.VI.1994; Jaj: Batroun, IV- 
VI.1984 (MHNL, NHMB, MHNG). 



PSYLLOIDEA FROM LEBANON 807 

Distribution. Throughout the Palaearctic (Conci et al. 1993). 
Host plants. Pyrus spp. (Rosaceae). 

Cacopsylla subpropinqua (Loginova) 

Material examined. Naas: Maten, 8.VI.1997; Broumana: Maten, 14.6.1997; 
Zaarour: Maten, 28.VI.1997 (MHNL, NHMB, MHNG). 

Distribution. Mongolia, Russia (Siberia, Far East) (Gegechkori & Loginova 
1990). 

Host plants. Salix spp. (Salicaceae). 

Cacopsylla sp. 

Material examined. Hrajel: Kesrouan, 27. VI. 1994; Hajar El Afrit: Kesrouan, 
16.IX. 1994 (MHNL). 

Comments. The material at hand is insufficient for identification. 

Psylla (Baeopelma) colorata Low 

Material examined. Achkout: Kesrouan, 17.IX.1994; Dahr El Souan: Maten, 
13.VIII.1985 (MHNL). 

Distribution. SE European (Conci et al. 1993). 
Host plant. Ostrya carpinifolia Scop. (Betulaceae). 

Psylla (Baeopelma) foersteri Flor 

Material examined. Naas: Maten, 8. X. 1997; Dahr El Souan: Maten, 16. VII. 
1997 (MHNL, NHMB, MHNG). 

Distribution. Throughout the Palaearctic (Klimaszewski 1973). 
Host plants. Alnus spp. (Betulaceae). 

Psyllidae: Rhinocolinae 

Agonoscena pistaciae Burckhardt & Lauterer 

Material examined. Bhamdoun: Aley, 16.X.1978; Ghazir: Kesrouan, 2.1.1982; 
(MHNL). 

Distribution. Iran, Israel, Jordan, Turkey (Al-Khawaldeeh 1997). 
Host plants. Pistacia spp. (Anacardiaceae). 

Megagonoscena gallicola Burckhardt & Lauterer 

Material examined. Kfar Katra: Chouf, 17.V.1981; Jabal Moussa: Kesrouan, 
15.VI.1986 and 15. VI. 1997; Naas: Maten, 14.VI.1997; Broummana: Maten, 
14.VI.1997 (MHNL, NHMB, MHNG). 

Distribution. Israel, Jordan (Al-Khawaldeh 1997). 

Host plant. Pistacia vera L. (Anacardiaceae). 



g08 n. zeidan-gèze & d. burckhardt 

Triozidae 

Bactericera albiventris (Foerster) 

Material examined. Zaarour: Maten, 28. VI. 1997; Kfar Debiane: Kesrouan, 
31. VII. 1985 (MHNL, NHMB, MHNG). 

Distribution. Austria, former Czechoslovakia, Demark, Finland, France, Ger- 
many, Great Britain, Greece, Hungary, Italy, Latvia, Norway, Poland, Russia (Euro- 
pean part, Siberia, Far East), Spain, Sweden, Switzerland, Turkey, Turkmenistan 
(Burckhardt & Lauterer 1997). 

Host plants. Salix spp. (Salicaceae). 

Bactericera curvatinervis (Foerster) 

Material examined. Broumana: Maten, 12. VI. 1997 and 14. VI. 1997; Naas: 
Maten, 8.VI.1997, 14.VI.1997 and 18.VII.1997; Zaarour: Maten, 28.VII 1997 
(MHNL, NHMB, MHNG). 

Distribution. Most of Europe, Caucasus, Russian Far East, Japan (Burckhardt 
& Lauterer 1997). 

Host plants. Salix spp. (Salicaceae). 

Bactericera nigricornis (Foerster) 

Material examined. Nahr Ibrahim: Kesrouan, 24.1.1981; Zaarour: Kesrouan, 
28. Vn. 1997 (MHNL). 

Distribution. West Palaearctic, Central Asia, Siberia, Mongolia (Burckhardt 
& Lauterer 1997). 

Host plants. Polyphagous on herbaceous plants. 

Eutrioza opima Loginova 

Material examined. Laklouk: Kesrouan, 22.VII.1986 (MHNL, NHMB). 
Distribution. Caucasus, France, Italy, Tunisia, Turkey, Ukraine (Conci et al. 
1996). 

Host plant. Unknown. 

Phylloplecta trisignata (Low) 

Material examined. Bhannés: Maten. 6.VIII.1985; Dahr El Souan: Maten, 
16.XI.1983; Faraya: Kesrouan, 16.VIII.1985 (MHNL) 

Distribution. Mediterranean without North Africa (Conci et al. 1996). 
Host plants. Rubus spp. (Rosaceae). 

Trioza alacris Flor 

Material examined. Nahr Ibrahim: Kesrouan, 24.1.1981 and 4.VI.1985 (MHNL). 

Distribution. Native to the Mediterranean, now occuring in the West Palae- 
arctic from Portugal, Southern Great Britain and Fennoscandia to the Caucasus, 
Turkey and the Crimea; introduced into North and South America (Conci et al. 1996). 

Host plant. Laurus nobilis L. (Lauraceae). 



PSYLLOIDEA FROM LEBANON g09 

Trioza ilicina (de Stefani) 

Material examined. Meyrouba: Kesrouan, 5.V.1985 (MHNL, NHMB, MHNG). 
Distribution. Algeria, France, Italy, Spain, Turkey (Conci et al. 1996). 
Host plant. Quercus ilex L. (Fagaceae). 

Trioza marginepunctata Flor 

Material examined. Jaj: Batroun, IV-VI.1984; Kartaba: Kesrouan, 22.XI.1985 
(MHNL). 

Distribution. Croatia, France, Israel, Italy, Spain (Conci et al. 1996). 
Host plant. Rhamnus alaternus L. (Rhamnaceae). 

Trioza neglecta Loginova 

Material examined. Broumana: Maten, 12.VI.1997 (MHNL, NHMB). 
Distribution. Armenia, Azerbaijan, Georgia, Iran, Rumania, Russia (European 
part), Turkey (Gegechkori & Loginova 1990). 

Host plant. Elaeagnus angustifolia L. (Elaeagnaceae). 

Trioza remota Foerster 

Material examined. Meyrouba: Kesrouan, 5.V.1985 (NHML, NHMB, MHNG). 
Distribution. Throughout the Palaearctic, though the records from the Far East 
and Central Asia need to be revised (Conci et al. 1996). 

Host plants. On deciduous Quercus spp. (Fagaceae). 

Trioza urticae (L.) 

Material examined. Janné: Kesrouan, 16. V. 1997; Yahchouch: Kesrouan, 
16.VI.1981 (MHNL, NHMB, MHNG). 

Distribution. Throughout the Palaearctic (Klimaszewski 1973). 
Host plants. Urtica spp. (Urticaceae). 

CONCLUSIONS 

To the four psylloid species previously recorded from Lebanon, 3 1 are added 
here. Compared to the 66 species known from neighbouring Israel, the 35 species 
listed here constitute probably less than half of the existing Lebanese psylloid species. 

Due to the incomplete knowledge, a biogeographical analysis of the Lebanese 
psylloid fauna is premature. Here a narrative summary of the current knowledge is 
provided (Table 1). Except for Cacopsylla sp., an unidentified species with unknown 
distribution, the known Lebanese species are generally widely distributed. Species 
endemic to Lebanon are not known so far. Mediterranean species (Table 1: columns 
ME and EM) make up 41.2 % of the identified species; eleven species are widely 
distributed over the Mediterranean Basin, whereas only 3 species are Eastern 
Mediterranean in distribution. Eighteen species (52.9 %) (Table 1: column PA) are 



810 N. ZEIDAN-GÈZE & D. BURCKHARDT 



Table 1. Distribution of Lebanese psylloids. PA = Palaearctic; ME = Mediterranean; EM = E 
Mediterranean; IN = introduced; IS = Israel (based on records by Bodenheimer 1937; 
Halperin et al. 1982; Halperin 1986; Halperin et al. 1988; Burckhardt & Halperin 1991; 
Burckhardt & Lauterer 1997); S Y = Syria (based on records by Talhouk 1969; Mustafa 
1991). 

Lebanese species PA ME EM IN IS SY 

Acizzia uncatoides (Ferris & Klyver) + + 

Agonoscena pistaciae Burckhardt & Lauterer + + + 1 

Arytainilla cytisi (Puton) + + 

Bactericera albiventris (Foerster) + + 

Bactericera curvatinervis (Foerster) + 

Bactericera nigricornis (Foerster) + 

Cacopsylla bidens (Sulc) + + +- 

Cacopsylla brunneipennis (Edwards) + 

Cacopsylla mariannae (Baeva) + 

Cacopsylla myrthi (Puton) + + 

Cacopsylla pulchella (Low) + + 

Cacopsylla py risuga (Foerster) + 

Cacopsylla subpropinqua (Loginova) + 

Cacopsylla sp. 

Colposcenia kiritshenkoi Loginova + 

Craspedolepta pontica Dobreanu & Manolache + + 

Craspedolepta sonchi (Foerster) + + 

Cyamophila stoklosai Klimaszewski & Lodos + 

Euphyllura phillyreae Foerster + + 

Euphyllura straminea Loginova + + + 3 

Eutrioza opima Loginova + 

Homotoma ficus (L.) + + + 

Livia juncorum (Latreille) + 

Livia mediterranea Loginova + + 

Livida spectabilis (Flor) + + 

Megagonoscena gallicola Burckhardt & Lauterer + + 

Phylloplecta frisi gnata (Low) + + 

Psylla (Baeopelma) colorata Low + 

Psylla (Baeopelma) joer steri Flor + 

Trioza alacris Flor + + 

Trioza ilicina (de Stefani) + 

Trioza marginepunctata Flor + + 

Trioza neglecta Loginova + 

Trioza remota Foerster + 

Trioza urticae (L.) + + 

1 Recorded as Agonoscena targionii (Lichtenstein) by Talhouk (1969). 

-Recorded as Psylla py ricola Foerster by Talhouk (1969), and as Cacopsyalla pyricola by 

OSSIANNILSSON (1992). 

3 Recorded as Euphyllura olivina (Costa) by Talhouk (1969). 

more or less widely distributed in the Palaearctic but are not restricted to the 
Mediterranean or Eastern Mediterranean Basin. Of interest is here the occurrence of 
Cacopsylla subpropinqua in Lebanon, previously known only from Mongolia, Siberia 
and the Russian Far East. A similar disjunct known distribution has Cacopsylla 
saligna (Loginova) in Spain and Kazakhstan (Lauterer & Burckhardt 1997). 



PSYLLOIDEA FROM LEBANON g ] j 

Two species, viz. Acizzia uncatoides and Trioza neglecta, have been intro- 
duced from Australia and East Asia respecitively. Homotoma ficus is treated here as 
Mediterranean element as it is established there for probably several thousands of 
years (see comments to Homotoma ficus). 

A comparison with the fauna from Israel (Table 1) indicates that the Lebanon 
constitutes the Southern limit of a series of species associated with deciduous trees 
and shrubs such as Alnus, Ostrya, Quercus, Salix spp. etc. (Bactericera albiventris, 
Cacopsylla brunneipennis, C. mariannae, C. pyrisuga, C. subpropinqua, Psylla colo- 
rata, P. foersteri and Trioza remota). The psylloid fauna from Syria is insufficiently 
studied. Apart from the six species mentioned by Talhouk (1969) and Mustafa 
(1991), there is one additional record from Syria which probably concerns Israel: 
Livilla syriaca (Low) (Hodkinson & Hollis 1987). The species was originally 
recorded from "Syria: Kaifa (Reitter) ? ' (Low 1882); this is a likely misspelling of 
Haifa in Israel. 

Apart from two species with unknown host relationships (Cacopsylla sp. and 
Eutrioza opima), a vast majority of the known Lebanese psylloids develops on woody 
plants: 26 spp. on woody plants, 7 spp. on herbaceous plants. At this stage of 
knowledge it is difficult to judge whether this is real or due to the applied collecting 
techniques. 

ACKNOWLEDGEMENTS 

We thank the AUPELF - UREF (exchange between French-speaking Univer- 
sities) for financial support for a study and co-ordination mission by NZG to 
Switzerland, and following institutions for various help and support: Université 
Libanaise, C.N.R.S Libanais, University of Geneva, Natural History Museum of 
Geneva. 

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Jordan. Zoology in the Middle East 15:71 -82. 

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33:286 pp. 

Burckhardt, D. 1987. Jumping plant lice (Homoptera: Psylloidea) of the temperate neo- 
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roidinae). Zoological Journal of the Linnean Society 89: 299-392. 

Burckhardt, D. 1989. Les Psylles (Insecta, Homoptera. Psylloidea) de l'Algérie. Archives des 
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Burckhardt, D. 1994. Psylloid pests ottemperate and subtropical crop and ornamental plants 
(Hemiptera, Psylloidea): a review. Trends in Agricultural Sciences, Entomology 2: 173- 
186. 

Burckhardt. D. & Halperin, J. 1991. Additions to the psyllid fauna of Israel (Homoptera: 
Psylloidea). Israel Journal of Entomology 25-26: 41-50. 



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Burckhardt, D. & Hodkinson, I. D. 1986. A revision of the west Palaearctic pear psyllids 

(Hemiptera: Psyllidae). Bulletin of entomological Research 76: 119-132. 
Burckhardt, D. & Lauterer, P. 1993. The jumping plant-lice of Iran (Homoptera, 

Psylloidea). Revue suisse de Zoologie 100: 829-898. 
Burckhardt, D. & Lauterer, P. 1997. A taxonomic reassessment of the triozid genus 

Bactericera (Hemiptera: Psylloidea). Journal of Natural History 31: 99-153. 
Burckhardt, D. & Mifsud, D. 1998. Psylloidea (Insecta: Hemiptera) of the Arabia Peninsula. 

Fauna of Arabia 17: in press. 
Burckhardt, D. & Önucar, A. 1993. A review of Turkish jumping plant lice (Homoptera, 

Psylloidea). Revue suisse de Zoologie 100: 547-574. 
Conci, C, Rapisarda, C. & Tamanini, L. 1993. Annotated catalogue of the Italian Psylloidea. 

First part (Insecta Homoptera). Atti dell'Accademia Roveretana degli Agiati, series 7, 

2B, 242 (1992): 33-136. 
Conci, C, Rapisarda, C. & Tamanini, L. 1996. Annotated catalogue of the Italian Psylloidea, 

Second part (Insecta Homoptera). Atti dell Accademia Roveretana degli Agiati, series 

7, 5B, 245 (1995): 5-207. 
Gegechkori, A. M. & Loginova, M. M. 1990. Psillidy SSSR. Akademiya Nauk Gruzinskoi 

SSR, Tbilisi. 164 pp. 
Güclü, S. & Burckhardt, D. 1996. New records of jumping plant-lice from Turkey 

(Hemiptera, Psylloidea). Entomofauna, Zeitschrift für Entomologie 17: 381-384. 
Halperin, J. 1986. An Introduced Psyllid Injurious io Acacia Trees. Phytoparasitica 14: 234- 

235. 
Halperin, L, Hodkinson, I. D. & Burckhardt, D. 1988. Six local psyllids (Homoptera: 

Psylloidea) new to the Israeli fauna. Phytoparasitica 16: 283-284. 
Halperin, J., Hodkinson, I. D. & Russell, L. M. 1982. A contribution to the knowledge of 

the psyllids of Israel (Homoptera: Psylloidea). Israel Journal of Entomology 16: 27-44. 
Klimaszewski, S. M. 1973. The jumping plant lice or psyllids of the Palaearctic. An annotated 

check-list. Annales Zoologici, Warszawa 30: 155-286. 
Lauterer, P. & Burckhardt, D. 1997. Central and West European willow feeding jumping 

plant-lice of the genus Cacopsylla (Hemiptera: Psylloidea). Entomological Problems 

28: 81-93. 
Loginova, M. M. 1973. Taxonomy of the tribe Euphyllurini (Psylloidea, Homoptera). Zoo* 

logicheskii Zhurnal 52: 858-869. (In Russian). 
Loginova, M. M. 1974. Jumping plant lice of the Tribe Stigmaphalarini Vondr. (Psylloidea, 

Aphalaridae) from arid regions of the Palaearctic. Entomologicheskoe Obozrenie 53: 

106-121. (In Russian). 
Mabberley, D. J. 1990. The plant-book. A portable dictionary of the higher plants. Cambridge 

University Press, reprinted with corrections, 707 pp. 
Mustafa, T. M. 1991. The Psyllids (Homoptera: Psylloidea) of Jordan. Iraqi Journal of 

Science 31:139-145. 
Ossiannilsson, F. 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. Fauna 

Entomologica Scandinavia 26: 1-345. 
Samy, O. 1973. Psyllids of Egypt. Bulletin of Entomological Society of Egypt 56: 437-480. 
Talhouk, A. M. 1969. Insects and mites injurious to crops in the Middle Eastern countriesi 

Monographien zur angewandten Entomologie. Beihefte zur Zeitschrift für angewandte 

Entomologie 21: 99-102. 
White, I. M. & Hodkinson, I. D. 1985. Nymphal taxonomy and systematics of the Psylloidea 

(Homoptera). Bulletin of the British Museum (Natural History), Entomology 50: 153- 

301. 



Revue suisse de Zoologie 105 (3): 813-822; décembre 1998 



Les organes producteurs de phéromones de quelques Hespérides 
(Lepidoptera, Hesperiidae, Hesperiinae) ' 

Jean WÜEST 

Muséum d'histoire naturelle, Case postale 6434, CH-121 1 Genève 6. 



The pheromone dispersing apparatus in some Hesperiinae (Lepidop- 
tera: Hesperiidae). - In Hesperiinae (Thymelicus lineola. Th. acteon and 
Hesperia comma), the organization of the pheromone dispersing apparatus, 
as well as the morphology of the androconia, present a growing degree of 
complexification, possibly reflecting the evolution within the group. The 
apparatus is simply formed of patches of androconia in Th. lineola. In 
Th. acteon, the scales adjacent to the patches of androconia are slightly 
modified and oriented towards the androconial line. In H. comma, the 
patches of androconia are completely covered by the adjacent scales. In 
Hesperiinae, the androconia are tubular scales containing pheromone 
within hollow medulla. These scales can break into pieces named osmo- 
phores, which are the dispersing means of the pheromone. In Th. lineola, 
the androconia are tubular and do not break, but some of them present 
constrictions, which can be hypothesized as a precursor state of the 
dehiscent zones present in the two other species. In Th. acteon, all andro- 
conia present dehiscent zones and can break into osmophores, but they 
often remain unbroken. In H. comma, all androconia are broken and the 
freed osmophores are glued together into a net just under the roof of the 
covering scales. 

Key-words: Lepidoptera - Hesperiidae - Androconia - Osmophores - 
Pheromone apparatus. 

INTRODUCTION 

Dans un précédent article (Wüest 1997), nous avons présenté les appareils à 
pheromone des mâles de quelques papillons de jour du genre Argynnis sensu lato. 
Nous avons poursuivi notre étude au moyen du microscope électronique à balayage 
sur quelques espèces de la famille des Hespérides, sous-famille des Hesperiinae, qui 
présentent un appareil tout à fait original et sans exemple comparable connu: ce sont 
des écailles androconiales contenant le principe attractif actif, qui se rompent et jouent 



Communication présentée sous forme de poster au Congrès Zoologia et Botanica 
(Genève, 18-20 février 1998) et à l'Assemblée annuelle de la Société entomologique suisse 
(Genève, 13-14 mars 1998). 

Manuscrit accepté le 29.06.1998 



g 14 JEANWÜEST 

le rôle de support "macroscopique" de l'odeur. Ces éléments se comportent comme 
une poudre et peuvent être dispersés par le mâle au cours de ses battements d'ailes. 
On peut en retrouver collés sur les antennes de la femelle (Sellier 1972) et il semble 
que la phéromone impliquée soit fort peu volatile et ait plutôt une action par contact. 
Ces écailles spécialisées avaient fait l'objet d'une étude de Reverdin (1916) qui les 
avait décrites en microscopie optique et avait déjà vu les différences entre Thymelicus 
limola et les autres espèces, celle-là présentant des écailles filiformes qui ne se 
rompent pas en segments alors que les autres espèces présentent des écailles en 
chapelets se rompant aisément. Les segments de ces écailles, en forme de petites 
saucisses, ont été appelés osmophores (Sellier 1971, 1972; Grasse 1975). Cepen- 
dant, la morphologie de l'appareil à phéromone (disposition des écailles, types d'é- 
cailles, etc.) ne semble jamais avoir été étudiée en détail, malgré quelques indications 
chez Sellier (1971), et nous voudrions en faire la description ici. Notre étude porte 
sur trois espèces qui illustrent bien la complexification de l'appareil et des écailles. 



MATERIEL ET METHODES 

Les trois espèces étudiées [Thymelicus limola (Ochsenheimer, 1808), Th. 
acteon (Rottemburg, 1775) et Hesperia comma (Linné, 1758)] proviennent du Valais 
(val d'Hérens, 1996). Les ailes ont été observées, après pulvérisation cathodique à 
l'or, dans un microscope électronique à balayage Zeiss 940A. 



L'APPAREIL A PHEROMONE 

Chez les mâles des Hespérides de couleur générale orange (représentant la 
sous-famille des Hesperiinae, genres Hesperia et Thymelicus présentés ici), on peut 
remarquer à la face supérieure des ailes antérieures une sorte de dilatation noire de la 
nervure cubitale, le "trait androconial". Il s'agit d'une zone spécialisée dans la pro- 
duction et la diffusion des attracteurs sexuels de ce groupe de Rhopalocères (Higgins 
&RILEY 1971). 

La partie essentielle de cet appareil est constituée d'une masse d'écaillés plus 
ou moins fortement modifiées, les androconies (Figs 1, 5 & 11). Elle est organisée en 
plusieurs massifs placés à la suite l'un de l'autre et suivant plus ou moins le tracé de 
la nervure cubitale, qui borde la cellule discoidale vers l'arrière. Si la coloration noire 
n'est pas présente à la face inférieure de l'aile, les massifs d'androconies s'y 
retrouvent chez Th. limola. Les tissus de l'aile sont fortement épaissis à cet endroit, 
comme le signalent Pivnick et al. (1992), renfermant probablement les cellules glan- 
dulaires à phéromones. 

A partir de ce schéma, des états de complexification peuvent être mis en 
évidence en comparant les appareils de diverses espèces d' Hespérides. 

Chez Thymelicus limola, on trouve l'appareil le plus simple (Fig. 1). Les 
écailles entourant et bordant les massifs d'androconies ne sont pas modifiées, ni dans 
leurs formes, ni dans leurs orientations. Elles ont alignées longitudinalement comme 



ORGANES À PHÉROMONES DES HESPÉRIDES 815 

la grande majorité des écailles de l'aile. Ce qui est visible sur l'aile, ce sont directe- 
ment les androconies. Il n'y a pas spécialisation d'une des faces de l'aile, les andro- 
conies se trouvant sur les deux faces. 

Chez Thymelicus acteon, une légère modification apparaît dans les écailles 
adjacentes à la zone des osmophores (Figs 5 & 6), qui deviennent plus larges et 
s'orientent perpendiculairement à la nervure médiane. Mais la zone des écailles à 
osmophores n'est pas couverte. Par contre, la face inférieure de l'aile ne présente 
aucun appareil androconial. 

Hesperia comma présente l'appareil à phéromone le plus complexe. Cet 
organe est constitué de deux parties, une proximale arrondie, qui s'étend entre les 
nervures cubitale et anale, et une plus distale et allongée, qui court parallèlement à la 
nervure cubitale bordant la cellule discoidale vers l'arrière (Figs 11 & 12). Les 
massifs d' osmophores, qui ont pourtant la même organisation que chez Thymelicus 
acteon, ne sont plus visibles de l'extérieur. Ils sont totalement recouverts par des 
écailles de couverture qui forment une sorte de toit au-dessus des osmophores (Figs 
12 & 13). Ces écailles sont très nettement plus larges que les écailles banales des 
ailes. Elles sont recourbées dans le sens de la longueur, ce qui assure une couverture 
efficace des osmophores, et orientées perpendiculairement à la nervure médiane. Du 
côté antérieur, une partie des écailles adjacentes, de morphologie semblable aux 
écailles banales des ailes, est orientée vers la base de l'aile (Fig. 12). Cette zone est de 
couleur noire et tranche avec la couleur orange de l'aile. Les écailles à osmophores 
sont mélangées à des écailles classiques, filiformes ou légèrement aplaties, à côtes 
rectilignes (Fig. 14). A la face inférieure de l'aile, une légère boursouflure des écailles 
permet de localiser la zone, sans que des androconies soient présentes. 



LES ANDROCONIES DE LA ZONE A PHEROMONE 

Chez Thymelicus lineola, les androconies ne sont pas modifiées en écailles 
porteuses d'osmophores. Les éléments diffuseurs de phéromone se présentent comme 
de simples écailles filiformes à côtes longitudinales (Fig. 2). Leur tige d'implantation 
est relativement longue; l'apex est lisse et résulte de la coalescence des côtes. Elles ne 
présentent pas d'amincissements de dehiscence et ne se rompent pas. La phéromone 
doit ici être diffusée sous la forme de vapeur. Pourtant, en examinant un grand 
nombre de ces écailles, on peut repérer sur certaines des zones où les côtes s'effacent, 
et où le cylindre de l'écaillé a tendance à se boursoufler, préfigurant une zone de 
dehiscence (Fig. 4). Mais les côtes restent toujours strictement longitudinales. 

Chez Thymelicus acteon et Hesperia comma, les écailles portant le principe 
attractif sont fortement modifiées et forment des segments appelés osmophores. 

Il s'agit d'écaillés très allongées, filiformes, présentant sur leur longueur un 
certain nombre de restrictions ou zones de dehiscence. Les segments ou osmophores 
se détachent et peuvent reformer un réseau (Figs 7, 8 & 13), leurs extrémités étant 
adhésives (particulièrement chez H. comma), peut-être du fait de la présence de la 
substance active sous forme non volatile qui remplit la cavité centrale (Grasse 1975). 



8 1 6 JEAN WÜEST 

Il est rare de rencontrer de telles écailles entières. En effet, elles sont destinées à se 
rompre au niveau de chaque zone de dehiscence, libérant ainsi de courts tronçons en 
forme de saucisses qui sont les éléments de transport de la phéromone. On peut 
cependant trouver parfois plusieurs segments encore attachés les uns aux autres 
(Fig. 8). La zone d'implantation de ces écailles montre qu'elles présentent dès la base 
la structure des segments osmophores (Figs 7 & 14). 

La forme des osmophores est assez constante d'une espèce à l'autre. Par 
contre, leur longueur est extrêmement variable d'un osmophore à l'autre et peut 
mesurer entre 10 um et 50 um et plus. Quelques petits détails de structure ou d'orne- 
mentation différencient les espèces étudiées ici, en particulier dans les zones de 
dehiscence. 

D'une manière générale, les osmophores présentent une partie renflée terminée 
à chaque extrémité par les zones de dehiscence. La partie renflée présente des côtes, 
avec des perforations dans les parties creuses entre les côtes, comme dans la plupart 
des écailles. Cependant, les côtes, au lieu d'être rectilignes, sont enroulées autour du 
cylindre de l'osmophore, avec un pas variable. Les zones de dehiscence, ou d'accro- 
chage d'un osmophore à son voisin, forment un petit renflement lisse, porteur de 
l'orifice de communication entre les osmophores. C'est à ce niveau surtout que se 
constatent les différences spécifiques. 

Les écailles à osmophores de Th. acteon ne semblent pas très fragiles, puis- 
qu'on trouve beaucoup d'entre elles portant encore des chapelets d'osmophores; 
d'autre part, les masses d'osmophores consistent en majeure partie en baguettes de 
plusieurs osmophores non séparés (Figs 7 & 8). Les massifs d'osmophores sont 
constitués de ces baguettes emmmêlées, sans qu'un réseau d'osmophores collés se 
mette en place comme chez H. comma. La zone d'écaillés à osmophores consiste 
presque exclusivement en ce type d'écaillés, avec quelques rares écailles étroites en 
général peu modifiées (Fig. 7). La base des écailles à osmophores présente une mor- 
phologie en tous points semblable à celle des osmophores (longueur du segment, 
largeur, ornementation) (Fig. 7). 

Chez Th. acteon, les segments ont environ 2,5 um de diamètre. Huit côtes 
garnissent la surface et font environ 2,5 fois le tour du segment pour un osmophore de 
15 um de long (Fig. 9). Les côtes ne présentent pas d'ornementation et les perfo- 
rations ont des dimensions variables (Fig. 10). Les extrémités de chaque segment 
présentent un renflement très bombé et lisse. L'ouverture du canal est placée latéra- 
lement (Fig. 9). La longueur des osmophores peut être très variable (Fig. 8). L'orien- 
tation des côtes peut également varier et parfois onduler le long des écailles (Fig. 8). 
On peut trouver des figures aberrantes d'osmophores présentant des chapelets de 
boules de jonction entre eux. 

Chez H. comma, les écailles à osmophores semblent très fragiles, puisque nous 
n'avons trouvé aucun osmophore en place (Fig. 14). Tous les osmophores sont 
regroupés à la surface de la zone, sous les écailles de couverture où ils forment des 
aggrégats collés en réseau (Fig. 13). On peut voir, entre les écailles peu modifiées qui 
parsèment la zone à androconies, les bases des écailles à osmophores restées en place 
(Fig. 14). Les osmophores ont toujours le même diamètre de 2,5 um. Les huit côtes 



ORGANES À PHÉROMONES DES HESPERIDES 817 

présentent un pas plus ou moins serré et peuvent faire 1,5 à 3,5 fois le tour d"un 
segment de 20 u m de long. Les extrémités du segment sont fusiformes et l'ouverture 
est latérale (Fig. 15). Par contre, les côtes présentent une ornementation transversale 
particulière sous la forme de petits bourrelets (Fig. 16). La base des écailles à 
osmophores est beaucoup plus courte que. les osmophores, mais présente la même 
ornementation superficielle. Les osmophores de cette espèce ont la faculté de se 
recoller à un autre segment et de former un réseau très intriqué qui remplit tout 
l'espace entre les écailles banales qui sont mélangées aux androconies et les écailles 
de couverture (Figs 13 & 14). Ce réseau d'osmophores collés a tendance à se localiser 
en surface de la zone à androconies, peut-être poussé vers le haut par les écailles 
banales (Fig. 13). 

DISCUSSION 

Parmi les trois espèces étudiées, on peut distinguer une gradation nette dans la 
complexité de l'appareil à phéromone. Chez Thymelicus lineola, la zone à andro- 
conies se présente simplement comme une zone à écailles modifiées, sans que les 
écailles adjacentes aient subi une quelconque transformation (forme ou orientation). 
De plus, les massifs d' androconies se trouvent chez cette seule espèce sur les deux 
faces de l'aile, indiquant un état non encore spécialisé. Chez T. acteon, par contre, on 
note que les écailles bordant la zone à osmophores sont plus larges et s'orientent 
perpendiculairement à la zone à osmophores, préfigurant l'apparition d'écaillés de 
recouvrement. Mais les osmophores restent bien visibles de l'extérieur. C'est chez 
Hesperia comma enfin, que l'appareil est le plus complexe. La zone à osmophores est 
entièrement cachée par des écailles de recouvrement, et la libération des osmophores 
doit être plus restreinte, dans des étapes bien précises de l'accouplement. Chez cette 
dernière espèce, la présence d'écaillés filiformes normales parmi les écailles à osmo- 
phores permettrait d'envisager que ces dernières par leurs mouvements réciproques 
facilitent la rupture des segments osmophores et les conduisent vers la surface de la 
zone à androconies. 

Ces étapes évolutives discutées ci-dessus rappellent ce que nous avions décrit 
chez les espèces du groupe des Argynnis Fab. 1807 (Wüest 1997). Chez Speyeria 
aglaja (Linné, 1758), la zone des androconies (qui ne libère pas d'osmophores) n'est 
pas recouverte par des écailles de recouvrement, contrairement à ce que nous pouvons 
trouver chez Argynnis paphia (Linné, 1758) sur la seconde nervure cubitale; cepen- 
dant la zone modifiée (contenant des écailles à phéromone) des autres nervures (mé- 
diane, cubitale 1 et anale) de A. paphia ne présente pas ce toit d'écaillés de recou- 
vrement. Nous devons être chez les Argynnes en présence d'un cas montrant les 
étapes de l'évolution de l'appareil à phéromone, entre deux espèces et aussi entre les 
nervures à l'intérieur d'une même espèce. 

Il semblerait donc que l'évolution ait tendance à protéger de tels appareils du 
milieu extérieur en recourant à plusieurs occasions (exemple de phénomènes de 
convergence entre des superfamilles différentes, quoique proches, les Hesperioidea et 
les Papilionoidea) à des écailles de couverture, aussi bien chez les Hespérides 



818 



JEAN WUEST 




Figs 1-4. Thymelicus limola. - 1. Zone à phéromone de l'aile antérieure (face supérieure). On 
distingue directement les massifs d'androconies. G = 40x. - 2. Détail de la surface des massifs 
d'androconies. Il n'y a pas d'osmophores. G = 600x. - 3. Zone à androconies en coupe, 
montrant quelques rares restrictions sur la longueur des écailles filiformes. G = 430x. - 
4. Androconie présentant une restriction et amorçant le processus de formation des osmo- 
phores. G = 6300x. 



Figs 5-10. Thymelicus acteoii. - 5. Zone à phéromone. G = 12x. - 6. Détail de la zone à 
phéromone. Les écailles adjacentes sont élargies et orientées perpendiculairement à la nervure. 
G = 60x. - 7. Zone à phéromone en coupe. Les osmophores restent en baguettes non détachées. 
G = 300x. - 8. Massifs d'osmophores. De nombreux segments restent attachés les uns aux 
autres. G = 1220x. - 9. Osmophore. G = 3100x. - 10. Détail de la partie centrale d'un osmo- 
phore. G= 18'500x. 



ORGANES A PHEROMONES DES HESPERIDES 



819 




820 



JEAN WUEST 




ORGANES À PHÉROMONES DES HESPERIDES 821 

(Hesperiinae, mais aussi Pyrginae, qui feront l'objet d'un prochain travail) que chez 
les Argynnes (Sellier 1971; Barth 1944). On peut penser que cela permettrait 
d'éviter un gaspillage des principes actifs, ou d'assurer leur diffusion seulement dans 
certaines phases du comportement de cour. Mais il est bien sûr illusoire de penser 
vérifier cette hypothèse en visualisant la position des écailles de recouvrement 
pendant les différentes étapes de l'accouplement, même si Barth (1944) a décrit les 
déplacements des écailles de l'appareil des Argynnes pendant le vol, permettant la 
libération des phéromones. Il serait intéressant de vérifier si on retrouve une telle 
architecture des appareils à phéromone (massifs d'androconies recouverts par des 
écailles de couverture) chez d'autres groupes de rhopalocères: chez les papillons de 
jour, les phéromones jouent un rôle dans les phases rapprochées de l'accouplement, 
après les informations visuelles, contrairement aux papillons nocturnes où les sémio- 
chimiques ont une action primaire à longue distance. Il serait aussi intéressant de 
chercher de tels exemples chez d'autres Lépidoptères diurnes n'appartenant pas aux 
Rhopalocères. 

Quant aux androconies elles-mêmes, nous avons pu mettre en évidence ici 
également une gradation entre les espèces étudiées, et cette gradation recouvre celle 
concernant la complexité de l'appareil lui-même. Thymelicus lineola représente un 
état peu différencié, puisque les androconies se présentent comme de simples écailles 
filiformes peu modifiées. Cependant, quelques-unes de ces écailles montrent une 
tendance à former des zones modifiées rappelant les zones de dehiscence des autres 
espèces, et Pivnick et al. (1992) ont montré que ces zones de dehiscence semblent se 
multiplier au cours de la vie imaginale, sans en expliquer le mécanisme. Thymelicus 
acteon, lui, présente deux types d'écaillés dans la zone à androconies, des écailles 
filiformes semblables à celles de Th. lineola, et de véritables écailles à osmophores. Il 
faut noter que les écailles filiformes peuvent présenter des côtes non plus longitu- 
dinales, mais inclinées, et rappeler ainsi les écailles à osmophores. Nous trouvons 
donc ici aussi des intermédiaires entre les deux types d'androconies. Enfin, Hesperia 
comma présente des osmophores un peu différents de ceux de 77t. acteon, avec des 
extrémités fusiformes. Les écailles normales parsemées dans la zone à osmophores 
sont filiformes ou légèrement aplaties, mais ne présentent pas d'intermédiaires avec 
les androconies à osmophores. 

Un autre critère concerne la fragilité des androconies à osmophores. Chez 
Th. lineola, les limites entre segments de type osmophore n'étant que partiellement 
indiquées, ces écailles ne se rompent pratiquement pas et restent entières dans la zone 
à androconies, quoique Pivnick et al. (1992) démontrent que les cassures, comme les 
zones de dehiscence, se multiplient avec l'âge de l'imago. Chez Th. acteon, les 



Figs 11-16. Hesperia comma. - 11. Zone à phéromone. Les massifs d'osmophores sont 
recouverts par des écailles de couverture. G = 9x. - 12. Détail montrant les écailles de 
couverture élargies et recourbées à l'apex. G = 23x. - 13. Coupe d'une chambre à osmophores. 
G = 60x. - 14. Détail d'une coupe de chambre à androconies montrant les écailles filiformes 
non modifiées et les écailles à osmophores toutes rompues. G = 435x. - 15. Osmophore. G = 
2400x. - 16. Détail d'un osmophore. G = 18'500x. 



822 JEAN WÜEST 

osmophores sont libérés par rupture, mais on peut trouver des androconies entières ou 
de longs chapelets d' osmophores intacts. Enfin, chez Hesperia comma, toutes les 
androconies ont libéré leurs osmophores qui, par leurs hautes propriétés collantes, ont 
reformé un réseau en surface des androconies, poussés peut-être par les écailles fili- 
formes présentes dans la zone, et peuvent se retrouver collés sur les antennes des 
femelles. 

Nous avons voulu rechercher si les espèces étudiées ici (Th. lineola et Th. 
acteon, H. comma) avaient fait l'objet de travaux sur la constitution chimique de leurs 
phéromones, ou sur les séquences de certains de leurs gènes, ce qui aurait permis de 
déduire des liens de parentés entre ces trois espèces, et de vérifier si notre hypothèse 
d'une complexification et donc d'une évolution, dont les étapes se seraient mainte- 
nues chez Th. lineola, Th. acteon et H. comma, était corroborée par les méthodes 
moléculaires. A notre connaissance, seuls Pivnick et al. (1992) ont effectué des élec- 
troantennogrammes sur Th. lineola, qui ont permis de prouver l'émission de phéro- 
mones chez les Hespériides. Nous avons en outre consulté des banques de données 
sur les phéromones et sur les séquences de DNA, ainsi que le Zoological Record 
(entre 1972 et 1997), mais sans parvenir à trouver aucun représentant de la famille des 
Hespérides. Nous envisageons dans un prochain travail d'orienter nos recherches dans 
cette direction. 

BIBLIOGRAPHIE 

Barth, R. 1944. Die männliche Duftorgane einiger Argynnis Arten. Zoologische Jahrbücher, 
Abteilung Anatomie und Ontogenie der Tiere 68: 331-362. 

Grasse. P. P. 1975. Phanères épidermiques. In: Grasse P. P. 1975. Traité de Zoologie, tome 
Vili, fase. Ill, pp. 48-67. Masson éd., Paris. 

Higgins, L. G. & Riley, N. D. 1971. Guide des Papillons d'Europe (Rhopalocères). Delachaux 
& Niestlé, 414 pp. 

Pivnick, K. A., Lavoie-Dornik, J. & McNeil, J. N. 1992. The role of the androconia in the 
mating behaviour of the European skipper, Thymelicus lineola, and evidence for a male 
pheromone. Physiological Entomology 17: 260-268. 

Reverdin, J.-L. 1916. Adopaea nova, mihi, species nov. Bulletin de la Société lépidoptéro- 
logique de Genève 3: 122-128. 

Sellier, R. 1971. Etude morphologique en microscopie électronique à balayage de quelques 
types d' androconies alaires chez les Lépidoptères diurnes. Compte Rendu hebdoma- 
daire des Séances de l'Académie des Sciences de Paris, Série D 273: 2550-2553. 

Sellier, R. 1972. Etude ultrastructurale en microscopie électronique à balayage et essai d'inter- 
prétation du mode de fonctionnement des poils androconiaux alaires chez les Hespe- 
riidae (Lépid. rhopal.). Compte Rendu hebdomadaire des Séances de l'Académie des 
Sciences de Paris, Série D 275: 2239-2242. 

Wüest, J. 1997. L'appareil à phéromone d' Argynnis paphia et de Mesoacidalia aglaja mâles 
(Lépidoptères Nymphalides) en microscopie électronique à balayage. Bulletin romand 
d'Entomologie 15: 47-56. 



Revue suisse de Zoologie 105 (4): 823-833; décembre 1998 



Nouvelles espèces asiatiques du genre Bryaxis et quelques données 
sur des espèces connues (Coleoptera: Staphylinidae: Pselaphinae) 

Sergei A. KURBATOV 1 & Ivan LÖBL 2 

1 Severodvinskaya 9-84, Moscou 129224, Russie. 

2 Muséum d'histoire naturelle, Case postale 6434, CH-121 1 Genève 6, Suisse. 



New Asian species of the genus Bryaxis, and data on previously known 
species (Coleoptera: Staphylinidae: Pselaphinae). - The following new 
species of Bryaxis from China are described: B. kimi sp. n. and B. namet- 
kini sp. n. from the prov. Sichuan, B. pletnevi sp. n. from the prov. Hubei, 
B. ruidus sp. n. and B. mendax sp. n. from the prov. Zhejiang, B. fictor sp. 
n. from the prov. Shaanxi. Bryaxis valentulus sp. n. is described from the 
Russian western Altai. An unidentified member of Bryaxis is recorded 
from Burma, and new records are given for B. sacrificus Kurbatov & Lobi 
and B. panda Kurbatov & Lobi. 

Key-words: Coleoptera - Staphylinidae - Pselaphinae - Bryaxis - China - 
Russia - Burma. 

INTRODUCTION 

Avec quelque 300 espèces recensées, dont de très nombreux endémiques 
locaux, Bryaxis Kugelann est certainement le genre de Pselaphinae paléarctiques 
ayant la plus forte diversité spécifique et écologique. La plupart des espèces de 
Bryaxis se rencontrent dans les régions circum-méditerranéennes et à l'extrême Est de 
la Russie, en Corée, au Japon et à Taiwan. Hormis B. bulbifer (Reichenbach) qui est 
répandu en Europe et en Sibérie, le genre est apparemment absent d'un territoire 
immense s'étirant de la Mer Caspienne au bassin du fleuve Amour et incluant toute la 
région himalayenne ainsi que les pays limitrophes. 

Actuellement, 12 espèces de Bryaxis ont été recensées en République Popu- 
laire de Chine (ci-dessous: Chine), dont une, B. kolîzei (Reitter), est largement 
répandue (Coulon & Li 1995; Kurbatov & Löbl 1995), et une autre, B. heilong- 
jiangensis (Li & Chen 1993), comporte dans l'illustration adjointe à sa description 
originale des caractères manifestement artefactuels. La faune des Bryaxis de Chine 
semble pauvre par rapport à certaines régions voisines où l'on en dénombre 13 
espèces de Corée (Löbl 1977; Nomura & Lee 1993), 19 à Taiwan (Löbl & 
Kurbatov 1996) et 34 au Japon (Löbl et al. 1998). 

Basée sur des récoltes récentes, cette étude comporte la description d'une 
espèce nouvelle de Bryaxis de l'Altaï russe et de six espèces nouvelles de Chine. 



Manuscrit accepté le 1 1.09.1998 



824 s - A - KURBATOV & I. LÖBL 



D'autre part, une espèce vraisemblablement nouvelle pour la science, mais non- 
décrite en l'absence des caractères essentiels du mâle, est signalée de Birmanie, et de 
nouvelles données sur la distribution de deux espèces de Bryaxis sont fournies. 

Abréviations utilisées: 

CSKM Collection privée de S. A. Kurbatov, Moscou 

MHNG Muséum d'histoire naturelle, Genève 

ZMAN Muséum Zoologique de la Division Sibérienne de l'Académie Russe de Sciences, 

Novosibirsk 
ZMUM Muséum Zoologique, Moscou 

DESCRIPTIONS 

Bryaxis kimi sp. n. Figs 1, 12 

Holotype 6 : Chine, sud de la province de Sichuan, environs de Xichang, 1700 m, 
débris végétaux, 26.VII.1996, leg. S. A. Kurbatov (ZMUM). Paratypes: mêmes données que 
Phototype, 2 3, 10 9 (ZMUM, MHNG, CSKM). 

Longueur: 1,45-1,55 mm; largeur maximale: 0,65-0,68 mm. Corps brun. 
Pubescence semi-érigée, plus longue sur les élytres et sur l'abdomen que sur l'avant- 
corps. Tête densément et presque entièrement ponctuée, les diamètres des points plus 
grands que les intervalles entre eux. Moitié basale de la dépression frontale et 
tubercules antennaires lisses. Lobe frontal large de 0,19-0,20 mm, très légèrement 
rétréci en arrière. Bord antérieur du front incliné en avant. Centre des fossettes du 
vertex situé en arrière du bord antérieur des yeux. Carène occipitale atteignant le bord 
postérieur de la dépression frontale. Yeux 6 nettement plus longs que les tempes en 
vue latérale, comportant 20-25 ommatidies; yeux 9 aussi longs que les tempes en vue 
latérale, comportant 10-12 ommatidies. Articles 2 et 3 des palpes maxillaires dépour- 
vus de tubercules. Articles antennaires: 3 plus long que large, 4-8 égaux, aussi longs 
que larges ou 7-8 à peine transverses, 9 aussi long et plus large que 8, 10 plus long et 
plus large que 9, 1 1 plus long que 8 à 10 réunis. Pronotum plus large que long, avec la 
même ponctuation que le vertex. Elytres à ponctuation plus grande et plus super- 
ficielle que celle du pronotum, diamètres des points le plus souvent plus grands que 
les intervalles entre eux. 

S. Articles 1 à 3 de l'antenne comme Fig. 12; scape long de 0,17 mm, 
pédicelle long de 0,06 mm. Protibias dépourvus d'échancrure, métatibias avec une 
petite dent apicale. Edéage (Fig. 1 ) long de 0,40 mm. 

9 . Scape cylindrique, 2 fois plus long que large, pédicelle ovale, plus long que 
large, plus long que l'article 3. 

Cette espèce est proche de B. emeicus Kurbatov & Lobi, 1995. Elle en offre 
l'ensemble des caractères, notamment la forme de l'édéage, très similaire à ceux de 
B. emeicus dont elle diffère nettement par le nombre et la forme des pièces sclérifiées 
du sac interne de l'édéage, en particulier par les branches du sclerite apical plus 
étroites et moins asymétriques. 

Etymologic L'espèce est nommée en l'honneur du Dr Boris B. Kim (Uni- 
versité de Moscou), dont l'aide a permis au premier auteur de réaliser un de ses 
voyages en Chine. 



NOUVELLES ESPECES ASIATIQUES DE BRYAX1S 



825 




Figs 1 à 4 

Bryaxis, édéages, vue dorsale: 1. B. kimi sp. n.; 2. B. nametkini sp. n.; 3. B. pletnevi sp. n. 
4. B. ruidus sp. n. Echelles = 0,1 mm. 



826 S. A. KURBATOV & I. LÖBL 



Bryaxis nametkini sp. n. Figs 2, 13 

Holotype 6: Chine, sud de la province de Sichuan, au sud de Xichang, Mt. Luoji, 
2300 m, débris végétaux, 17.VII.1996, leg. S. A. Kurbatov (MHNG). Paratypes: mêmes 
données que l'holotype, 4 <3, 7 9; mêmes données mais 18.VII.1996, 4 â, 5 9; mêmes 
données mais 2500 m. 23.VII.1996, 2(5,22; mêmes données mais 2500 M, 24.VII.1996, 1 6 
(ZMUM, MHNG, CSKM). 

Longueur: 1,45-1,55 mm; largeur maximale: 0,65-0,69 mm. Corps d'un brun 
rougeâtre à brun. Pubescence semi-érigée, plus longue sur les élytres et sur l'abdomen 
que sur l'avant-corps. Tête densément ponctuée, les diamètres des points plus grands 
que les intervalles entre eux. Moitié basale de la dépression frontale lisse, ponctuation 
sur les tubercules antennaires très fine et éparse. Lobe frontal comme chez B. kimi, 
large de 0,19-0,20 mm. Bord antérieur du front, fossettes du vertex et carène occi- 
pitale comme chez B. kimi. Yeux S plus longs que les tempes en vue latérale, 
comportant 20 ommatidies environ; yeux 9 plus courts que les tempes en vue laté- 
rale, comportant 5-10 ommatidies. Palpes maxillaires comme chez B. kimi. Articles 
antennaires: 3 plus long que large, 4-5 aussi longs que larges, 6-7 à peine transverses, 
8 de la même longueur, mais à peine plus large, 9 plus large et à peine plus long que 
8, 10 plus long et plus large que 9, 1 1 aussi long que 8-10 réunis. Pronotum plus large 
que long, ponctué comme le vertex. Ponctuation des élytres formée de points plus 
grands et plus superficiels que ceux du pronotum, les diamètres des points généra- 
lement plus grands que les intervalles entre eux. 

â . Articles 1 à 3 de l'antenne comme Fig. 13; scape long de 0,17 mm, pédi- 
celle long de 0,06 mm. Protibias et métatibias comme chez B. kimi. Edéage (Fig. 2) 
long de 0,32 mm. 

9 . Scape cylindrique, environ 2 fois plus long que large, pédicelle ovale, plus 
long que large, plus long que l'article 3. 

Espèce ressemblant à B. kimi par les caractères externes, mais en différant par 
la proportion des articles antennaires 6-11 et par la forme du scape chez le â. 
L'édéage diffère très nettement de celui de B. kimi et de B. emeicus par les paramères 
plus larges, ornés chacun d'une seule paire de soies, et par la forme de pièces 
sclérifiées symétriques du sac interne de l'édéage. 

Etymologic L'espèce est nommée en l'honneur du Dr Serguei N. Nametkin 
(Université de Moscou) qui a soutenu les missions du premier auteur effectuées en 
Chine. 

Bryaxis pletnevi sp. n. Figs 3, 14 

Holotype â: Chine, est de la province de Hubei, à 30 km au nord-est de Macheng, 
environ 500 m, débris végétaux, 25. V. 1995, leg. S. A. Kurbatov (ZMUM). Paratypes: mêmes 
données que l'holotype, 2^9 (ZMUM, MHNG). 

Longueur: 1,5 mm; largeur maximale: 0,64-0,70 mm. Corps brun. Pubescence 
comme chez les deux espèces précédentes. Ponctuation sur la tête et le pronotum 
encore plus dense, les intervalles entre les points beaucoup plus petits que les dia- 
mètres des points. Fond de la dépression frontale et les tubercules antennaires lisses. 
Yeux â bien plus grands que les tempes en vue latérale, comportant 20 ommatidies 
environ; yeux 9 aussi longs que les tempes en vue latérale, comportant 10-12 omma- 



NOUVELLES ESPECES ASIATIQUES DE BRYAXIS g27 

tidies. Autres caractères de la tête comme chez B. kimi et B. nametkini. Articles 
antennaires: 3 un peu plus long que large, 4-8 progressivement raccourcis, 4 aussi 
long que large, 8 nettement plus large que long, 9 aussi long que 8, mais nettement 
plus large, 10 encore plus large et un peu plus long, presque 2 fois plus large que 
long, 11 aussi long que 7-10 réunis. Pronotum plus large que long, ponctué aussi 
densément que le vertex. Ponctuation sur les élytres formée de points plus grands, 
plus superficiels et beaucoup plus épars que les points sur l' avant-corps, les diamètres 
des points en général un peu plus grands que les intervalles. 

S. Articles 1 à 3 de l'antenne comme Fig. 14; scape long de 0,165 mm, 
pédicelle long de 0,06 mm. Fémurs renflés; protibias, sur leur quart distal, ornés d'une 
échancrure nette, délimitée par une dent; métatibias munis d'une longue dent apicale. 
Edéage (Fig. 3) long de 0,39 mm. 

9 . Scape et pédicelle comme chez les espèces précédentes. 

Cette espèce ressemble à B. kimi par la forme du scape et du pédicelle 6 . Elle 
en diffère par la ponctuation de l' avant-corps plus dense et par l'édéage beaucoup 
plus étroit, à sac interne symétrique, orné de nombreux stylets et d'une plaque apicale. 

Etymologic L'espèce est dédiée à Vladimir A. Pletnev, entomologiste de 
Moscou et ami du premier auteur. 

Bryaxis ruidus sp. n. Figs 4, 15 

Holotype S: Chine, province de Zhejiang, Tienmushan, 2.IX.1994, leg. G. de Rouge- 
mont (MHNG). Paratypes: mêmes données que l'holotype, 1 <3, 2 9 (MHNG). 

Longueur: 1,5-1,55 mm; largeur maximale: 0,63-0,70 mm. Corps brun, élytres 
parfois plus clairs, brun rougeâtres. Pubescence comme chez les espèces précédentes. 
Tête comme chez B. pletnevi, mais dépourvue d'une carène occipitale. Yeux compor- 
tant 22-25 ommatidies chez le â et 8-10 chez la 9. Articles antennaires: 3 plus long 
que large, 4 et 5 aussi longs que larges, 6 à 8 à peine transverses, 9 à peine plus large 
et plus long que 8, légèrement transverse, 10 nettement plus large et à peine plus long 
que 9, 1 1 plus long que 8-10 réunis. Pronotum plus large que long, sa ponctuation très 
dense, presque aussi dense que celle du vertex; les diamètres des points beaucoup plus 
grands que les intervalles. Elytres avec la ponctuation plus éparse, les points plus 
grands et plus superficiels que sur le pronotum, à peine plus grands que les intervalles 
entre eux. 

S . Articles 1 à 3 de l'antenne comme Fig. 15; scape long de 0,13 mm, pédi- 
celle long de 0,10 mm. Pattes comme chez B. pletnevi, mais la dent apicale des 
métatibias plus courte. Edéage (Fig. 4) long de 0,37 mm. 

9 . Scape cylindrique, à peu près 1 ,5 fois plus long que large, pédicelle ovale, 
d'un tiers plus court que le scape. 

L'espèce ressemble à B. holciki Lobi, 1964 par l'absence de la carène occi- 
pitale, par la forte ponctuation sur la tête et sur le pronotum et par les caractères 
sexuels secondaires. Elle en diffère notamment par le sac interne de l'édéage plus 
complexe, muni de stylets très fins et de pièces sclérifiées pus larges, progressivement 
rétrécies vers l'apex. 



g28 S. A. KURBATOV & I. LÖBL 

Bryaxis mendax sp. n. Figs 5, 16 

Holotype S: Chine, province de Zhejiang, Tienmushan, 2.IX.1994, leg. G. de Rouge- 
mont (MHNG). Paratypes: mêmes données que l'holotype, 2 9 (MHNG). 

Longueur: 1,45-1,55 mm; largeur maximale: 0,64-0,67 mm. Uniformément 
brun, pubescence comme chez B. kimi. Tête comme chez B. ruidus. Yeux deux fois 
plus longs que les tempes en vue latérale et comportant 25 ommatidies environ chez 
le â; yeux 2 un peu plus longs que les tempes en vue latérale et comportant 10 
ommatidies environ. Articles antennaires: 3 un peu plus long que large, 4-8 à peine 
plus larges que longs, égaux, 9 plus large et plus long que 8, légèrement transverse, 10 
aussi long et plus large que 9, 11 plus long que 8-10 réunis. Pronotum à peine plus 
large que long, sa ponctuation nettement moins dense que celle du vertex. Elytres à 
ponctuation aussi dense que celle du pronotum, mais formée de points plus grands et 
plus superficiels; les diamètres des points en général un peu plus grands que les 
intervalles entre eux. 

S . Articles 1 à 3 de l'antenne comme Fig. 16; scape long de 0,10 mm, pédi- 
celle long de 0,09 mm. Protibias très légèrement échancrés sur leur quart distal, 
dépourvus de dent à la base de l'échancrure; métatibias munis d'une dent apicale 
légèrement pointue. Edéage (Fig. 5) long de 0,35 mm, paramères fortement courbés 
dorsalement à l'apex. 

9 . Scape cylindrique, à peine plus long que large, pédicelle allongé, un peu 
plus court que le scape. 

Cette espèce ressemble à B. ruidus, notamment par la conformation de la tête. 
Elle en diffère par le pronotum à ponctuation nettement moins dense et, surtout, par 
l'édéage. Les paramères sont échancrés subapicalement chez B. mendax, semblables à 
ceux de B. smetanai Lobi; les pièces sclérifiées du sac interne, constituées d'une paire 
de grandes dents latérales et d'une plaque centro-apicale, sont très caractéristiques 
chez ces espèces. 

Bryaxis fictor sp. n. Figs 6, 17 

Holotype 6 : Chine, province de Shaanxi, Nonwutai, à peu près 25 km sud de Xian, 
400-800 m, 17.IX.1995, débris végétaux, leg. G. de Rougemont (MHNG). 

Longueur: 1,5 mm; largeur maximale: 0,67 mm. Corps uniformément brun. 
Pubescence comme chez B. kimi. Tête densément ponctuée, la dépression frontale et 
les tubercules antennaires lisses. Lobe frontal long de 0,20 mm, à côtés subparallèles. 
Bord antérieur du front incliné en avant. Centre des fossettes du vertex situé en arrière 
du niveau du bord antérieur des yeux. Carène occipitale presque absente, à peine 
visible au niveau de la moitié postérieure des fossettes du tentorium. Yeux plus longs 
que les tempes en vue latérale, comportant 20 ommatidies environ. Articles 2 et 3 des 
palpes maxillaires dépourvus de tubercules. Articles antennaires: 3 plus long que 
large, 4-8 subégaux, à peine transverses, 9 plus long et à peine plus large que 8, 
légèrement transverse, 10 plus grand que 9, plus transverse, 11 aussi long que 7-10 
réunis. Pronotum légèrement plus large que long, ayant la même ponctuation que le 
vertex. Elytres à ponctuation un peu moins dense que celle du pronotum; les points 



NOUVELLES ESPECES ASIATIQUES DE BRYAXIS 



829 




Figs 5 à 1 1 

Bryaxis, édéages, vue dorsale, et métatibias S , vue latérale: 5. B. mendax sp. n., édéage; 
6. B. fictor sp. n., édéage; 7, 10 et 11. B. valentulus sp. n., édéage (7) et métatibias ( 10 et 11): 
B. femoratus (Aube), édéage (8) et métatibia (9). Echelles = 0.1 mm (Figs 5 à 8); = 0.2 mm 
(Figs 9 à 11). 



330 S. A. KURBATOV & I. LÖBL 

plus grands et plus superficiels que sur le pronotum; les diamètres des points plus 
grands que les intervalles entre eux. 

â . Articles 1 à 3 de l'antenne comme Fig. 17; scape long de 0,10 mm, pédi- 
celle long de 0,10 mm, large de 0,12 mm. Partie centrale du métasternum avec une 
ponctuation fine mais particulièrement dense. Profémurs légèrement mais nettement 
déprimés à la base du bord inférieur, sur une surface à peu près aussi grande que le 
protrochanter. Protibias légèrement échanrés sur leur quart distal; échancrure limitée 
par une dent. Métatibias munis d'une petite dent apicale. Tubercule gulaire grand et 
complexe, saillant en avant en vue latérale. Edéage (Fig. 6) long de 0,36 mm. 

Cette espèce présente des caractères uniques sur le métasternum, les pro- 
fémurs, les antennes et l'édéage. La ponctuation particulière du métasternum et les 
dépressions profémorales sont considérées comme étant liées au sexe â. Bryaxis 
fictor semble être isolé au sein du genre. Les lobes dorsaux sur les paramères 
rapproche la nouvelle espèce de B. wolongensis Kurbatov & Löbl du Sichuan ainsi 
que de la plupart des espèces de Taiwan, dont elle diffère nettement par le sac interne 
de l'édéage relativement simple, muni d'un sclerite circulaire basai joint au sclerite 
apical bifide. 

Bryaxis valentulus sp. n. Figs 7, 10, 11, 18 

Holotype 6: Russie, Altaï occidental, chaîne Ivanovsky, à 10 km au sud de Lenino- 
gorsk, forêt clairsemée (¥ Abies, 1400 m, 30.V.1996, leg. R. Dudko (ZMAN). Paratypes: mêmes 
données que l'holotype, 3 et, 1 2 (ZMAN. MHNG, CSKM). 

Longueur: 2,05-2,2 mm; largeur maximale: 0,77-0,85 mm. Corps brun rou- 
geâtre ou uniformément brun. Pubescence double, à soies mi-dressées assez denses et 
à soies plus longues, plus dressées et beaucoup moins nombreuses. Tête lisse, aux 
points épars et très petits. Lobe frontal très légèrement rétréci en arrière, large de 
0,19-0,20 mm. Bord antérieur du front incliné en avant. Centre des fossettes du vertex 
un peu postérieur au niveau du bord antérieur des yeux. Carène occipitale atteignant 
le bord postérieur de la dépression frontale. Yeux aussi longs que les tempes en vue 
latérale, comportant 12-16 ommatidies chez 6, un peu plus petits chez 9. Articles 2 
et 3 de palpes maxillaires dépourvus de tubercules. Articles antennaires: 3-8 de la 
même largeur, 3 plus long que large, 4-5 égaux, plus longs que larges, plus courts que 
3, 6-8 aussi longs que larges ou à peine plus longs que larges, plus courts que les 
précédents, 9 aussi long que large, plus grand que 8 (8 et 9 à peine transverse chez 
5), 10 plus large, à peine plus long que 9, légèrement transverse, 11 plus long que 8- 
10 réunis. Pronotum presque aussi long que large (longueur/largeur = 19/20), lisse. 
Ely très à ponctuation formée de grands points; les diamètres des points aussi grands 
que les intervalles entre eux. 

6 . Articles 1 à 3 comme Fig. 18; scape long de 0,20 mm, pédicelle long de 
0,085 mm. Métasternum orné d'une dépression triangulaire. Fémurs et tibias, surtout 
les antérieurs et postérieurs, renflés. Protibias ornés d'une échancrure profonde, 
limités proximalement par une grande dent aiguë; en arrière de l' échancrure, une 
petite saillie prolongée par une carène très fine. Métatibias soit pourvus d'une saillie 



NOUVELLES ESPECES ASIATIQUES DE BRYAXIS 



831 







Figs 12 à 19 

Bryaxis, articles 1 à 3 de l'antenne 6: 12. B. kimi sp. n.; 13. B. nametkini sp. n.; 14. B. pletnevi 
sp. n.; 15. B. ruidus sp. n.; 16. B. mendax sp. n.; 17. B. fictor sp. n.; 18. B. valentulus sp. n.; 19. 
B. femoratus (Aube). Echelles = 0.1 mm. 

anguleuse, soit ornés d'une dent au milieu de leur bord intérieur; dent apicale longue 
(Figs 10, 11). Edéage (Fig. 7) long de 0,67 mm; sac interne éventuellement inversé. 

9 . Scape cylindrique, presque deux fois plus long que large, pédicelle ovale, 
nettement plus long que large, presque 2 fois plus court que le scape. 



832 s - A - KURBATOV & I. LÖBL 

Cette espèce semble proche de B. femoratus (Aube) par ses caractères sexuels 
secondaires et par la structure de l'édéage. Chez B. femoratus, le scape et le pédicelle 
sont plus petits, le scape est seulement 1,5 fois plus long que large et orné d'un 
tubercule glandulaire tronqué (Fig. 19), l'échancrure des protibias n'est pas limitée 
apicalement par une saillie et les paramères de l'édéage sont moins arqués. La 
nouvelle espèce en diffère également par la taille du corps plus grande, la coloration 
plus claire (les téguments sont foncés chez B. femoratus, les élytres parfois noirâtres), 
les proportions des articles antennaires (chez B. femoratus les articles 4-5 sont aussi 
longs que larges, 6-8 légèrement transverses, 9-10 nettement transverses), la forme 
des métatibias chez le S (Figs 9 à 11) et par la forme des structures sclérifiées du sac 
interne de l'édéage que comporte une large dent apicolatérale, absente chez B. 
femoratus (Fig. 8). 

NOUVELLES LOCALITÉS 

Bryaxis sacrificus Kurbatov & Lobi, 1995 

Matériel: 4 â , 2 9. Chine, ouest de la province de Hubei, Shennongjia Nat. 
Res., 2000-2200 m, débris végétaux, 7.VI.1995, leg. S. A. Kurbatov (CSKM). 
Note. L'espèce n'était connue que de la province chinoise de Shaanxi. 

Bryaxis panda Kurbatov & Lobi, 1995 

Matériel: 1 S , Chine, ouest de la province de Hubei, Shennongjia Nat. Res., 
2000-2200 m, débris végétaux, 7.VI.1995, leg. S. A. Kurbatov (CSKM). 

Notes. Espèce décrite et connue seulement de Sichuan. 

Le S de Shennongjia Nat. Res. diffère des spécimens de la série-type par 
l'absence totale de tubercule sur le pédicelle. Ainsi, nous interprétons la présence ou 
l'absence du tubercule glandulaire chez B. panda comme un cas de variabilité infra- 
spécifique. 

Bryaxis sp. 

Matériel: 5 9, Birmanie, Shan province, Namhsan, 1600 m, litière, 19.02. 
1996, leg. S. A. Kurbatov (CSKM). 

L'absence de S ne permet pas de donner une description basée sur des carac- 
tères sûrs. Toutefois, la découverte d'une espèce de Bryaxis en Birmanie est impor- 
tante. Le genre est inconnu de la région himalayenne, du Tibet, de Yunnan et de 
Thaïlande. 

LITTÉRATURE 

Coulon, G. & Li. J. 1995. On some Pselaphinae from China (Coleoptera, Staphylinidae). 

Bulletin et Annales de la Société Royale belge d'Entomologie 131: 483-486. 
Kurbatov, S. A. & Löbl, I. 1995. Contribution to the knowledge of the East Asian Bryaxis 

(Coleoptera, Staphylinidae. Pselaphinae). Archives des Sciences (Genève) 48): 161-172. 
Li, J. & Chen. P. 1993. Studies on fauna and ecogeography of soil animals. Press of Northeast 

Normal University, 265 pp. 



NOUVELLES ESPECES ASIATIQUES DE BRYAXIS 833 



Löbl, I. 1964. Neue ostasiatische Arten der Gattung Bryaxis Kugelann (Col., Pselaphidae). 

Acta Societatis entomologicae cechosloveniae 61: 43-46. 
Löbl, I. 1977. Weitere Pselaphiden von der Koreanischen Volksdemokratischen Republik. 

Bulletin de l'Académie Polonaise des Sciences. Série des sciences biologiques Cl. V, 

25:235-241. 
Löbl, I. & Kurbatov, S. A. 1996. The Bryaxis of Taiwan (Coleoptera: Staphylinidae:* 

Pselaphinae). Bulletion of the National Museum of Natural Science (Taichung) 8: 1-22. 
Löbl, I., Kurbatov, S. A. & Nomura, S. 1998. On the Japanese species of Biyaxis (Cole- 
optera: Staphylinidae: Pselaphinae), with notes on allied genera and on endoskeletal 

polymorphy. Species Diversity (in print). 
Nomura, S. & Lee, C. E. 1993. A revision of the family Pselaphidae (Coleoptera) from South 

Korea. Esakia 33: 1-48. 



Revue suisse de Zoologie 105 (4): 835-837; décembre 1998 



A new species of Stetholiodes Fall, 1910 
(Coleoptera, Leiodidae, Agathidiini) from Taiwan 

Fernando ANGELINI* & Jonathan COOTER** 

* s.s. 7 per Latiano, km 0-500, 1-72021 Francavilla Fontana, (Brindisi), Italy. 
** 19 Mount Crescent, Hereford, HR1 1NQ, England 

A new species of Stetholiodes Fall, 1910 (Coleoptera, Leiodidae, Aga- 
thidiini) from Taiwan. - Stetholiodes magnifica n. sp. is described from 
Taiwan. A new generic record from Taiwan. 

Key-words: Coleoptera - Leiodidae - Stetholiodes - Taiwan. 



INTRODUCTION 

A hitherto undetected unique specimen of Stetholiodes, collected by Dr Ales 
Smetana during 1991, was found amongst undetermined Leiodidae in the collection 
of the Natural History Museum, Geneva. This species is described here, it represents 
the first known occurrence of the genus Stetholiodes in Taiwan. 

DESCRIPTION 

Stetholiodes magnifica n. sp. Figs 1-5 

Length 4.1 mm (holotype 6). Dorsum black, posterior margin of pronotum 
broadly lighter, raised lateral margin of elytra narrowly lighter. Each elytron with a 
testaceous 'C'-shaped macula, internally limited by stria 2, anterior lobe extending as 
far as the 6th stria and the posterior lobe just reaching the 8th stria (figs 1 & 2). 
Mesosternum dark reddish-brown, metasternum black; antennae testaceous with 
segments 7-10 and basal part of 11 dark; legs reddish-brown. Microreticulation clear 
on head, more superficial on pronotum and absent on elytra, puncturation distinct on 
head an pronotum; each elytron with 9 punctured striae. Suturai striae clear, well 
defined, confined to apical half of elytron. 

Head. Microreticulation clear, well impressed, surface somewhat dull; punctures 
clear, moderately large, separated from each other by 1-3 times their own diameter. 
Antero-lateral margins rounded, uniform. Clypeus rectilinear. Clypeal line absent. Eyes 
protuberant, head widest just behind the eyes, temples very short. Antennae with 3rd 
segment 1-7 times longer than 2nd and longer than 4th+5th together. 



Manuscript accepted 23.06.1998 



836 



F. ANGELINI & J. COOTER 



Pronotum. Microreticulation more superficial than on head; puncturation more 
superficial than on head and the punctures smaller, separated from each other by 2-5 
times their own diameter. 1-6 times as broad as head, moderately broader than long 
(width/length = 1-67) and moderately convex (width/height = 174). Anterior margin 
slightly curved, sides sharply curved anteriorly, lateral outline with anterior and 
posterior margins sub-parallel. Measurements of pronotum of holotype: length 1-15 
mm, width 192 mm and height 110 mm. 





Figs 1-5 

Stetholiodes magnifica n. sp. 6: body outline 1, dorsal and 2, lateral, scale line = 10 mm; 
3, aedeagus lateral, apex of aedeagus 4, ventral and 5, dorsal view, scale line = 0-5 mm. 



A NEW SPECIES OF STETHOLIODES 837 

Elytra. Microreticulation absent. The 9 striae composed of large clear punc- 
tures, separated from each other by 0-5-1 times their own diameter; punctures in 
interstices in the order of one-quarter, or less, the diameter of those of the striae, 
clearly impressed and separated by 0-5-2 times their own diameter. Distinctly broader 
than the pronotum and slightly longer than broad (width/length = 0-97) slightly 
convex (width/height = 2). Lateral outline with sharp humeral angle. Measurements of 
elytra of holotype: length 2T5 mm, width 2T0 mm and height 1-05 mm. 

Metathoracic wings present. 

Meso- and metasternum: with weak median carina, lateral lines complete, 
femoral lines absent. 

Legs: Tarsal formula of male 5-5-4, segments 1-3 of anterior tarsi somewhat 
dilated (female not known). 

Aedeagus as in figures 3, 4 & 5. 

Holotype: â , Taiwan, Taichung Hsien, Hseuhshan, above Shan-Lin-Gien Hut, 
3360 m, lO.v.1991, leg A. Smetana, in Geneva Natural History Museum collection. 

Discussion: Stetholiodes magnifica sp. n. is very similar to Stetholiodes turnai 
Angelini & Svec. It differs by having large conspicuous testaceous maculae on the 
elytra, larger size, the microreticulation of the head more clearly impressed, colouring 
of the antennae, in the greater length ratio of 3rd/2nd antennal segments, and in 
having a differently formed aedeagus. 

Distribution: Taiwan (Taichung Hsien). First record of Stetholiodes Fall from 
Taiwan. 

ACKNOWLEDGEMENTS 

The authors wish to express their thanks to Dr Ivan Lobi, Department of 
Entomology, Geneva Museum of Natural History, for the loan of this material. 



Revue suisse de Zoologie 105 (4): 839-877; décembre 1998 



New data on Oribatids (Acari: Oribatida) from St. Lucia (Antilles). 
(Acarologica Genavensia LXXXIX) 1 

Sândor MAHUNKA 

Zoological Department, Hungarian Natural History Museum, 

Baross utca 13, H-1088 Budapest, Hungary. 

New data on Oribatids (Acari: Oribatida) from St. Lucia (Antilles). 
(Acarologica Genavensia LXXXIX). - Elaboration of the material 
collected by T. Jaccoud and L. de Roguin in St. Lucia in 1989. 67 species 
are identified and listed, 16 of them are described as new to science and 
one of them also represents a new genus: P arac arino galumna gen. n. 
(Galumnidae). Licneremaeus antillensis Mahunka, 1985 = Licneremaeus 
discoidalis (Willmann, 1930): new synonymy. 

Key-words: Acari - Oribatida - Taxonomy - New taxa - St. Lucia 
(Antilles). 

INTRODUCTION 

The soil-mite fauna of the Caribbean territory including the Lesser and the 
Greater Antilles is fairly well-known. The richness and the zoogeographical signi- 
ficance of the area have been shown by several authors (e.g. Grandjean 1929, 1930; 
Willmann 1933, Balogh & Mahunka 1974). The oribatids of St. Lucia have already 
been discussed by the present author (Mahunka 1985). The study of additional 
material preserved in the Muséum d'histoire naturelle, Geneva proves our frag- 
mentary knowledge of this fauna. Part of this material is presented in this paper-. 

The material comprises 67 species of which 16 are new to science. For one of 
them a new genus is established. 25 known species are recorded for the first time for 
St. Lucia. Several endemic species, only known from the original descriptions, are 
recorded again together with many species already known from various regions of the 
Neotropics. In my previous paper (Mahunka 1985) 69 species are recorded for 
St. Lucia, now a total of 1 10 species are known from this island. 

The terminology and the taxonomic arrangement correspond to my previous 
papers (e.g. Mahunka 1994). 



New title for the series "Neue und interessante Milben aus dem Genfer Museum 
I-LX" and "New and interesting mites from the Geneva Museum LXI-LXXX". 

2 This research programme was partly sponsored by the Hungarian Scientific Research 
Fund (OTKA 16729). 

Manuscript accepted 16.10.1998 



840 SANDOR MAHUNKA 

LIST OF LOCALITIES 

STL-79/1: Saint Lucia (Castries): Hotel Halcyon Sands et environs immédiats (jardin, forêt), 
sur la presqu'île de Vigie; (st. 1), 14.VI.1979; leg. T. Jaccoud et L. de Roguin. 

STL-79/2: Saint Lucia (Gros Islet): Pied du Mont Layau, avant le village de Monchy, vallée de 
l'Espérance River, rivière presque à sec, dans forêt de Lauriers et Manguiers; (st.2) 
50 m; 15.VI.1979; leg. T. Jaccoud et L. de Roguin. 

STL-79/3: Saint Lucia (Dennery): Entre Barre de l'Isle Ridge et la Mabouva Valley, avant le 
col, près de Thomaso, E de Morne Panache, forêt claire, prélèvement sur souche 
d'arbres pourris; (st. 3.), 50 m; 16. VI. 1979; leg. T. Jaccoud et L. de Roguin. 

STL-79/4: Saint Lucia (Dennery/Dauphin): Pied du Piton Flore, au NW de Dernière Rivière, 
récolte de feuilles dans forêt dense, très en pente, avec falaises et abris sous roche; 
(st.4), 300 m; 17.VI.1979; leg. T. Jaccoud et L. de Roguin. 

STL-79/5: Saint Lucia (Dennery): Entre Barre de l'Isle Ridge et la Mabouva Valley, avant le 
col, près de Thomaso, E de Morne Panache, forêt claire, prélèvement sur souche 
d'arbres pourris; (st.3.), 50 m; 16. VI. 1979; leg. T. Jaccoud et L. de Roguin (B) 3 . 

STL-79/6: Saint Lucia (Dennery): Entre Barre de l'Isle Ridge et la Mabouva Valley, avant le 
col, près de Thomaso, E de Morne Panache, forêt claire, prélèvement sur souche 
d'arbres pourris; (st.3.), 50 m; 16. VI. 1979; leg. T. Jaccoud et L. de Roguin (B) 3 . 

STL-79/7: Saint Lucia (Dennery): Entre Barre de l'Isle Ridge et la Mabouva Valley, avant le 
col, près de Thomaso, E de Morne Panache, forêt claire, prélèvement sur souche 
d'arbres pourris, tamisage, Berlese à Genève; (st.3.), 50 m; 16.VI.1979; leg. T. Jaccoud 
et L. de Roguin (WB) 4 . 

STL-79/8: Saint Lucia (Anse la Raye): Vers Massecré, au NE de Anse la Raye, végétation 
arbustive, dense mais peu haute, nombreux épineux, sol caillouteux; (st.5), 100 m; 
17.VI.1979; leg. T. Jaccoud et L. de Roguin. 

STL-79/1 1: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt 
de pluie typique, très bien conservée (protégée), très humide prélèvement de sol; (st.6.), 
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B) 3 . 

STL-79/1 3: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt 
de pluie typique, très bien conservée (protégée), très humide prélèvement de sol; (st.6), 
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B) 3 . 

STL-79/15: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt 
de pluie typique, très bien conservée (protégée), très humide prélèvement de sol; (st.6), 
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B) 3 . 

STL-79/1 7: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt 
de pluie typique, très bien conservée (protégée), très humide, tamisage; (st.6), 200-350 
m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin. 

STL-79/1 8: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt 
de pluie typique, très bien conservée (protégée), très humide prélèvement de sol; (st.6.), 
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B) 3 . 

STL-79/1 9: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt 
de pluie typique, très bien conservée (protégée), très humide prélèvement de sol; (st.6), 
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B) 3 . 



3 (B): extraction par appareils Berlese à Genève. 

4 (WB): extraction par appareils Winkler-Moczarski sur place et par appareils Berlese à 



Genève. 



ORIBATIDS FROM ST. LUCIA 841 

LIST OF IDENTIFIED SPECIES 

Mesoplophoridae Ewing, 1917 

Mesoplophora hauseri Mahunka, 1 982 

Localities: STL-79/5: 8 specimens; STL-79/7: 2 specimens. 

Distribution: Costa Rica (known from the type locality only); first 

record for St. Lucia. 
Mesoplophora (Parplophora) subiilis Niedbata, 1981 

Localities: STL-79/12: 5 specimens; STL-79-18: 3 specimens. 

Distribution: Well distributed in the Neotropical and Oriental Region 

(see Niedbala 1985); first record for St. Lucia (Figs 1-3). 

Phthiracaridae Perty, 1841 

Hoplophorella lanceosetoides Mahunka, 1985 

Locality: STL-79/8: 9 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Hoplophorella scapellata (Aoki, 1965) 

Locality: STL-79/8: 2 specimens. 

Distribution: Perhaps Circumtropical; first record for St. Lucia. 

Lohmanniidae Berlese, 1916 

Meristacarus longisetosus Mahunka, 1978 

Locality: STL-79/1 : 3 specimens. 

Distribution: Dominican Republic (known from the type locality only); 

first record for St. Lucia. 
Torpacarus omittens Grandjean, 1950 

Locality: STL-79/8: 1 specimen. 

Distribution: Well distributed in Central- and South America; second 

record. 

Trhypochthoniidae Willmann, 1931 

Afronothrus incisivus neotropicus Balogh & Mahunka, 1974 

Locality: STL-79/8: 2 specimens. 

Distribution: Cuba (known from the type locality only); first record for 

St. Lucia. 
Allonothrus neotropicus Balogh & Mahunka, 1969 

Locality: STL-79/1 : 2 specimens. 

Distribution: Bolivia (known from the type locality only); first record 

for St. Lucia. 
Archegozetes magnus mediosetosus Mahunka, 1978 

Localities: STL-79/4: 4 specimens; STL-79/8: 25 specimens. 



g42 SANDOR MAHUNKA 



Distribution: Mauritius (known from the type locality only); first 
record for St. Lucia. 

Epilohmanniidae Oudemans, 1923 

Epilohmannia pallida americana Balogh & Mahunka, 1981 
Locality: STL-79/4: 1 specimen. 

Distribution: Paraguay (known from the type localities only); first 
record for St. Lucia. 

Nanhermanniidae Sellnick, 1928 

Cyrthermannia simplex Mahunka, 1985 

Locality: STL-79/5: 3 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 

Hermanniellidae Grandjean, 1934 

Sacculobates horologiorum Grandjean, 1962 

Localities: STL-79/1: 1 specimen; STL-79/8: 1 specimen. 
Distribution: Central America and Northern part of South America; 
second record. 

Liodidae Grandjean, 1954 

Teleioliodes zikani (Sellnick, 1930) 

Locality: STL-79/4: 3 specimens. 
Distribution: Brazil; first record for St. Lucia. 

Microtegeidae Balogh, 1972 

Microtegeus borhidii Balogh & Mahunka, 1974 

Locality: STL-79/1 9: 3 specimens. 

Distribution: Cuba (known from the type localities only); first record 

for St. Lucia. 
Microtegeus hauseri sp. n. 

Localities: STL-79/1; STL-79/1 7. 
Microtegeus lucianus sp. n. 

Localities: STL-79/2; STL-79/7; STL-79/1 7; STL-79/19. 

Eremaeozetidae Piffl, 1962 

Eremaeozetes lineatus Mahunka, 1985 

Locality: STL-79/8: 1 specimen. 

Distribution: St. Lucia (known from the type locality only); second 

record. 



ORIBATIDS FROM ST. LUCIA 843 

Eremaeozetes roguini sp. n. 

Locality: STL-79/7. 

Microzetidae Grandjean, 1936 

Berlesezetes auxiliaris Grandjean, 1936 

Localities: STL-79/2: 1 specimen; STL-79/4: 1 specimen; STL-79/5: 4 

specimens; STL-79/6: 5 specimens; STL-79/7: 7 specimens. 

Distribution: Circumtropical; second record for St. Lucia. 
Cosmozetes jaccoudi sp. n. 

Locality: STL-79/19. 

Eremulidae Grandjean, 1 965 

Eremulus nigrisetosus Hammer, 1958 

Locality: STL-79/8: 3 specimens. 

Distribution: South America; first record for St. Lucia. 

Damaeolidae Grandjean, 1965 

Fosseremus laciniatus (Berlese, 1905) 

Locality: STL-79/4: 2 specimens. 

Distribution: Cosmopolitan; first record for St. Lucia. 

Eremobelbidae Balogh, 1961 

Eremobelba piffli Mahunka, 1985 

Localities: STL-79/1: 2 specimens; STL-79/4: 2 specimens; STL-79/8: 

2 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 

Basilobelbidae Balogh, 1961 

Basilobelba insularis Mahunka 1985 

Localities: STL-79/4: 1 specimen; STL-79/8: 4 specimens. 
Distribution: St. Lucia (known from the type locality only); second 
record. 

Zetorchestidae Michael, 1898 

Zetorchestes schusteri Krisper, 1984 

Locality: STL-79/8: 3 specimens. 
Distribution: Brazil; first record for St. Lucia. 

Carabodidae C. L. Koch, 1837 

Kalloia simpliseta Mahunka, 1985 

Locality: STL-79/1 : 5 specimens. 

Distribution: St. Lucia (known the type locality only); second record. 



844 SANDOR MAHUNKA 

Klapperiches penicillus sp. n. 

Localities: STL-79/2; STL-79/12. 

Tectocepheidae Grandjean, 1954 

Tegeozetes tunicatus Berlese, 1913 

Locality: STL-79/4: 3 specimens. 

Distribution: Circumtropical; second record for St. Lucia. 

Dampfiellidae Balogh, 1961 

Beckiella vitiosa Mahunka, 1985 

Localities: STL-79/3: 1 specimen; STL-79/7: 2 specimens. 
Distribution: St. Lucia (known from the type locality only); second 
record. 

Cuneoppiidae Balogh, 1983 

Cuneoppia laticeps Balogh & Mahunka, 1969 
Locality: STL-79/4: 1 specimen. 

Distribution: Bolivia (known from the type locality only); first record 
for St. Lucia. 

Oppiidae Grandjean, 1951 

Aeroppia adjacens Mahunka, 1985 

Localities: STL-79/1 : 2 specimens; STL-79/4: 1 specimen; STL-79/8: 1 

specimen. 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Aeroppia hammerae Mahunka, 1985 

Localities: STL-79/1: 1 specimen; STL-79/3: 4 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Aeroppia asymmetrica Mahunka 1985 

Localities: STL-79/1: 5 specimens; STL-79/8: 3 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Amerioppia extrema Mahunka 1985 

Localities: STL-79/1: 3 specimens; STL-79/3: 1 specimen; STL-79/8: 

1 specimen; STL-79/1 3: 2 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Amerioppia paraguayensis Balogh & Mahunka, 1981 

Locality: STL-79/3: 5 specimens. 

Distribution: Paraguay, Brazil; first record for St. Lucia. 



ORIBATIDS FROM ST. LUCIA 845 

Dissorhina neotropicalis sp. n. 

Locality: STL-79/19. 
Gittella insularis sp. n. 

Locality: STL-79/17. 
Machuella ventrisetosa Hammer, 1 96 1 

Locality; STL-79/19: 2 specimens. 

Distribution: Neotropics and Oriental Region; first record for 

St. Lucia. 
Mohtzoppia subfallax sp. n. 

Locality: STL-79/17. 
Multioppia insularis Mahunka, 1985 

Locality: STL-79/19: 2 specimens. 

D i s t r i b u t i o n : St. Lucia (known from the type locality only); second 

record. 
Oppiella nova (Oudemans, 1902) 

Localities: STL-79/5: 2 specimens; STL-79/11: 8 specimens; STL- 

79/13: 1 specimen. 

Distribution: Cosmopolitan; second record for St. Lucia. 
Oxyoppia antillensis sp. n. 

Localities: STL-79/6; STL-79/13. 
Striatoppia tribuliformis Balogh & Mahunka, 1981 

Locality: STL-79/4: 2 specimens. 

Distribution: Paraguay (known from the type locality only); first 

record for St. Lucia. 

Suctobelbidae Jacot, 1938 

Suctobelbella baculifera Balogh & Mahunka, 1981 

Locality: STL-79/19: 2 specimens. 

Distribution: Paraguay (known from the type locality only); first 

record for St. Lucia. 
Suctobelbella complexa (Hammer, 1958) 

Locality: STL-79/19: 2 specimens. 

Distribution: Neotropics; first record for St. Lucia. 
Suctobelbella variosetosa (Hammer, 1961) 

Localities: STL-79/13: 3 specimens; STL-79/19: 2 specimens. 

Distribution: Circumtropical; second record for St. Lucia. 

Cymbaeremaeidae Sellnick, 1928 

Scapheremaeus longilamellatus Mahunka, 1 985 
Locality: STL-79/1: 1 specimen. 

Distribution: St. Lucia (known from the type locality only); second 
record. 



846 SANDOR MAHUNKA 

Licneremaeidae Grandjean, 1931 

Licneremaeus discoidalis (Willmann, 1930) 

= Licneremaeus antillensis Mahunka, 1985 nov. syn. 

Localities: STL-79/18: 1 specimen; STL-79/19: 2 specimens. 

Distribution: Guatemala; St. Lucia. 

Scutoverticidae Grandjean, 1954 

Arthrovertex hauseri Mahunka, 1985 

Localities: STL-79/2: 1 specimen; STL-79/3: 1 specimen; STL-79/19: 

2 specimens. 

Distribution: St. Lucia (known from the type locality only); second 
record. 

Parakalummidae Grandjean, 1936 

Parakalumma piton sp. n. 

Localities: STL-79/3; STL-79/6; STL-79/7; STL-79/12; STL-79/17. 

Mochlozetidae Grandjean, 1960 

Mochlozetes asculpturatus Mahunka, 1985 

Locality: STL-79/1 : 6 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Unguizetes similis sp. n. 
Localities: STL-79/1 ; STL-79/19. 

Oripodidae Jacot, 1925 

Benoibates minimus Mahunka, 1985 

Localities: STL-79/12: 1 specimen; STL-79/17: 2 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Oripoda lobata Mahunka, 1985 

Localities: STL-79/8: 2 specimens; STL-79/17: 3 specimens. 

Distribution: St. Lucia (known from the type locality only); second 

record. 

Haplozetidae Grandjean, 1936 

Nasobates mirabilis Balogh et Mahunka, 1969 

Locality: STL-79/2: 2 specimens. 

Distribution: South- and Middle-America; first record for St. Lucia. 
Peloribates antillensis (Mahunka, 1985) 

Localities: STL-79/4: 1 specimen; STL-79/7: 1 specimen; STL-79/19: 

3 specimens. 



ORIBATIDS FROM ST. LUCIA 847 

Distribution: St. Lucia (known from the type locality only); second 

record. 
Peloribates capucinus (Berlese, 1908) 

Locality: STL-79/18: 1 specimen. 

Distribution: Cosmopolitan; first record for St. Lucia. 
Rostrozetes carinatus Beck, 1962 

Localities: STL-79/1: 2 specimens; STL-79/3: 3 specimens; STL-79/13: 

1 specimen. 

Distribution: Neotropical; first record for St. Lucia. 
Rostrozetes ovulum (Berlese, 1908) 

Localities: STL-79/4: 2 specimens; STL-79/13: 1 specimen. 

Distribution: Circumtropical; first record for St. Lucia. 

Austrachipteriidae Luxton, 1985 

Lamellobates molecula (Berlese, 1916) 

Localities: STL-79/4: 2 specimens; STL-79/8: 6 specimens. 
Distribution: Circumtropical; first record for St. Lucia. 

Phenopelopidae Petrunkevitch, 1955 

Eupelops spongiosus sp. n. 

Locality: STL-79/8. 

Epactozetidae Grandjean, 1930 

Truncozetes rugosus sp. n. 

Locality: STL-79/1. 

Galumnidae Jacot, 1 925 

Galumna flabellifera Hammer, 1958 

Locality: STL-79/1: 15 specimens. 

Distribution: Circumtropical; second record for St. Lucia. 
Galumna hamij er Mahunka, 1985 

Localities: STL-79/5: 3 specimens; STL-79/8: 9 specimens. 

Distribution: Guadeloupe, Brazil; first record for St. Lucia. 
Paracarino galumna genavensium gen. n., sp. n. 

Localities: STL-79/1; STL-79/5; STL-79/8; STL-79/1 1; STL-79/1 5; 

STL-79/1 7. 
Pergalumna cucheae sp. n. 

Locality: STL-79/1. 
Pilogalumna antillensis sp. n. 

Locality: STL-79/8. 



848 



SANDOR MAHUNKA 




Figs 1-3 

Mesoplophora (Parplophora) subtilis Niedbala, 1981 - 1: body in lateral aspect, 2: anogenital 
region, 3: aspis in dorsal aspect. 



ORIBATIDS FROM ST. LUCIA 849 

DESCRIPTION OF THE NEW TAXA 

Microtegeus hauseri sp. n. Figs 4-5 

Material examined: Holotype: St. Lucia: STL-79/17; 1 paratype: STL-79/1. Holotype: 
MHNG 6 , 1 paratype (1556-PO-96): HNHM 7 . 

Measurements.- Length of body: 198-214 urn, width of body: 126-134 
urn. 

Prodorsum: Rostral apex conical. Lamellae wide, without sharp inner 
spur, but a well developed incisure present. Surface of prodorsum granulated, the 
granules only partly composing a polygonal reticulation. Rostral setae simple, 
straight, lamellar setae arising on the anterior margin of the lamellae, slightly dilated 
basally. Interlamellar setae short, bacilliform, a characteristic, strong thickening pre- 
sent at their insertion. Sensillus conspicuously long, directed laterally and backwards, 
its capitulum elongate, with strong spines on its surface (Fig. 4). 

Notogaster: Well granulated, the granules form a polygonal reticulation. 
Larger protruding tubercles absent, in lateral view surface appearing irregular. Ten 
pairs of short, straight, bacilliform or spiniform notogastral setae present. Two median 
setae arising very near to each other. 

Ventral regions (Fig. 5): Weakly sclerotised, apodemes and epimeral 
borders composing a disjoint reticulation. Sternal apodeme absent between apodemes 
2 and the sejugal ones. Discidium smaller than in the following species. Ventral plate 
irregularly granulated. One shorter, one long longitudinal and one transversal lath 
observable. Neither tubercles nor protuberances along the genital aperture. All setae 
in the anogenital region minute, simple, excepting the median anterior genital setae. 

Remarks: Refer to the remarks on the following species. 

Derivatio nominis: I dedicate the new species to my friend Dr Bernd 
Hauser, in recognition of his enormous efforts in managing research on soil- 
microarthropods during his activity as curator of the Department of Arthropods of the 
Geneva Museum from 1968 to 1998. 

Microtegeus lucianus sp. n. Figs 6-7 

Material examined: Holotype: St. Lucia: STL-79/7; 1 paratype: STL-79/2; 3 paratypes: 
STL-79/17; 1 paratype: STL-79/19. Holotype and 3 paratypes: MHNG, 2 paratypes (1555-PO- 
96): HNHM. 

Measurements.- Length of body: 198-235 urn, width of body: 105-139 
urn. 

Prodorsum: Rostrum conical. Lamellae very wide, with well developed 
incisure and spur on their anteromedian corner, with a transversal lath between them. 
Lamellar surface covered by smaller, interlamellar region with greater, pustules or 
granules, the latter form a polygonal sculpture. Rostral setae thin, arched inwards. 



6 MHNG: deposited in the Muséum d'histoire naturelle, Geneva. 

7 HNHM: deposited in the Hungarian Natural History Museum, Budapest, with an 
identification number of the specimen in the Collection of Arachnida. 



850 



SANDOR MAHUNKA 




Figs 4-5 
Microtegeus hauseri sp. n. - 4: body in dorsal aspect. 5: body in ventral aspect. 



lamellar setae arising on the rounded anterior margin of lamellae, they are basally 
dilated. Interlamellar setae short, slightly bacilliform. Sensillus capitate, its dorsal 
surface with blunt spines. 

Notogaster (Fig. 6): Three pairs of large projections (clearly visible 
under low magnification) on the notogaster. Its surface covered also by pustules and 
granules like the prodorsal ones and mostly also forming a kind of polygonal pattern. 
Ten pairs of spiniform. short notogastral setae present, their position typical for the 
family. 

Ventral regions (Fig. 7): Coxisternal region strongly sclerotised, all 
epimeral borders and the apodemes conspicuous. Epimeral setal formula: 3-1-3-3. 
All epimeral setae slightly spiniform. Anogenital setal formula: 5-1-2-2. 
Lyrifissures iad in adanal position. 

Legs: All legs monodactylous. 



ORIBATIDS FROM ST. LUCIA 



851 




-\- 7- 




Figs 6-7 
Microtegeus lucianus sp. n. - 6: body in dorsal aspect, 7: body in ventral aspect. 

Remarks: On the basis of the sculpture of the notogaster (resembling 
Eremobelba) these two new species are readily distinguished from all heretofore 
known species of the genus. M. hauseri is distinguished from M. lucianus by the 
unique form of the sensillus. 

Derivatio nominis: Named after St. Lucia. 



Eremaeozetes roguini sp. n. Figs 8-1 1 

Material examined: Holotype: St. Lucia: STL-79/7; 1 paratype: from the same sample. 
Holotype: MHNG, 1 paratype (1557-PO-96): HNHM. 

Measurements.- Length of body: 346-367 urn, width of body: 205-226 
urn. 

Prodorsum: Lamellae large, covering the whole prodorsal surface, 
except posteromedially. Cerotegument layer (as on the rest of the body) thick, 
composed of irregular rugae. None of the prodorsal setae visible in dorsal view, 
rostral and lamellar setae minute, interlamellar setae absent. Sensillus conspicuously 
long and narrow, fusiform, its margin waved, surface mostly with bacilliform spines 
(Fig. 8). 



852 



SANDOR MAHUNKA 







Figs 8-11 

Eremaeozetes roguini sp. n. - 8: body in dorsal aspect, 9: body in ventral aspect, 
10: posteromedian part of the coxisternal region, 1 1 : tarsus and tibia of leg. I. 



ORIBATIDS FROM ST. LUCIA 853 

Notogaster: Pteromorphae very large, ventrally reflexed to the ventral 
plate. Notogastral surface ornamented by a polygonate design, created by irregular 
foveolae. The distance among the foveolae comparatively large, so the border-lines 
mostly wide. Lacking separate median and or lateral part of the notogaster. Ten pairs 
of minute, straight and simple, spiniform notogastral setae present, two pairs among 
them arising medially. Lyrifissures im and the glandular opening also seen. 

Gnathosoma: Mentum strongly rugose having a characteristic 
transversal lath anteriorly. 

Ventral regions (Fig. 9): The whole surface well sclerotised, but the 
epimeral borders and the apodemes faint and only partly observable. A characteristic 
median thickening exists in front of the genital aperture (Fig. 10). Epimeral setal 
formula: 3-1-2-2, setae 3c and 4c absent or not visible. Surface of the genital plate 
also rugose, the anal plate foveolate. Along the genital opening a pair of strong 
longitudinal rugae visible. Anogenital setal formula: 6 - 1 - 2 - 3. All setae minute, 
excepting the inner pair of anterior genital setae. Both pairs of posterior adanal setae 
(aJj, ad 2 ) inserted on an arched lath. 

Legs: All legs with strong claws. The surface of the joints well rugose, the 
femur of leg III and IV widened, bearing a blade-like formation. Solenidia cpj and cp 2 
of leg I arising on a large process (Fig. 1 1), cp 2 ver y sma U- 

Remarks: On the basis of the characteristic polygonate sculpture of the 
notogaster, the new species stands very near to Eremaeozetes undulatus Mahunka, 
1985, also described from St. Lucia. The new species may be distinguished from 
Eremaeozetes undulatus by its much longer sensillus and the much smaller "cell" of 
the sculpture, and especially by the sculpture of the coxisternal region in front of the 
genital opening. 

Derivatio nominis: I dedicate the new species to the collector of this 
material, late Dr L. de Roguin (Muséum d'histoire naturelle, Geneva). 

Cosmozetes jaccoudi sp. n. Figs 12-14 

Material examined: Holotype: St. Lucia: STL-79/19; 27 paratypes: from the same 
sample. Holotype: MHNG and 18 paratypes: MHNG, 9 paratypes: (1558-PO-96): HNHM. 

Measurements.- Length of body: 223-251 urn, width of body: 166-186 
urn. 

Prodorsum: Rostral apex triangulate in dorsal aspect (Fig. 12), beak- 
shaped in lateral aspect (Fig. 14). Lamellae typical for the genus, connected by an 
arched translamella medially. Their lateral cusps very long and strong, median cusps 
resembling tubercles, hardly protruding; lamellar seta arising on it. Rostral setae 
slightly, lamellar seta strongly, thickened, the latter spiniform, directed outwards, 
crossing the outer cusps of lamellae. Interlamellar setae fine, short, located on the 
lamellar surface. One pair of nearly round structures present in the interlamellar 
region. Exobothridial setae conspicuously long, straight, arising on small tubercles. 
Sensillus directed outwards, its pedicel long, capitulum rounded with long spines, 
which are shorter proximally than distally. 



854 



SANDOR MAHUNKA 




Figs 12-14 

Cosmozetes jaccoudi sp. n. - 12: body in dorsal aspect, 13: body in ventral aspect, 

14: podosoma in lateral aspect. 



ORIBATIDS FROM ST. LUCIA 855 

Notogaster: Dorsosejugal suture indistinct. Pteromorphae small, ending 
in sharp, triangular lateral cusps. Notogastral margin slightly excavated laterally. 
Strong heterotrichy among the notogastral setae, c the shortest. All slightly ciliate. 

Lateral part ofpodosoma: Tutorium well developed, it is simple 
with a normal, spiniform apex, continued in a long lath, reaching the insertion of the 
rostral setae (Fig. 14). Pedotecta I distinctly rugose, pedotecta 2-3 conspicuously 
large, seta 3c arising on its surface. Discidium also well developed. Circumpedal 
carina not reaching to the lateral margin of the ventral plate. 

Gnathosoma: All setae and eupathidium of the palpal tarsus spiniform, 
nearly equal in length and arising marginally around the joint. Setae of palpal tibia 
much thinner than tarsal ones. 

Ventral regions (Fig. 13): Apodemes and epimeral borders typical for 
the family. Epimeral setal formula: 3-1-3-3. All setae conspicuously ciliate. 
Anogenital setal formula: 6-0-2-3. Anterior pair of genital setae much longer than 
the others, conspicuously ciliate. All the other setae in this region small and fine. 

Remarks: The heretofore known species of the genus Cosmozetes Balogh 
& Mahunka, 1969 are distributed in Bolivia, Brazil and Cuba. On the basis of the 
spiniform lamellar setae the new species stands nearest to C. striatissimus Balogh & 
Mahunka, 1969 (from Bolivia). However, in the new species the characteristic 
striation of the lamellae is absent and the outer lamellar cusp is much longer than the 
lamellar setae (equal in length in C. striatissimus) and no interlamellar structure is 
present in C. striatissimus. 

Derivatio nominis: I dedicate the new species to Mr T. Jaccoud 
(Muséum d'histoire naturelle), collector of this material. 

Klapperiches penicillus sp. n. Figs 15-18 

Material examined: Holotype: St. Lucia: STL-79/12; 4 paratypes: from the same 
sample; 2 paratypes: STL-79/2. Holotype and 3 paratypes: MHNG, 3 paratypes: (1559-PO-96): 
HNHM. 

Measurements.- Length of body: 341-372 urn, width of body: 155-180 
urn. 

Prodorsum: Rostrum widely rounded, lamellae narrow, running margi- 
nally. Rostral and lamellar setae thin, simple, interlamellar ones penicillate. Lamellar 
surface smooth, interlamellar region distinctly pustulate. Sensillus comparatively 
long, directed outwards, its capitulum calyciform, open laterally (Fig. 15). 

Notogaster: Whole surface pustulate. Dorsosejugal suture nearly 
straight. Ten pairs of small, penicillate notogastral setae present. 

Lateral part ofpodosoma: Tutorium present. Near to the rostral 
margin a characteristic, serrate minitectum present (Fig. 18), it is connected with the 
lamellar apex. Surface of the lateral part also ornamented by large foveolae. 
Pedoctecta I with a sharp tooth on its distal end (Fig. 17). 

Ventral regions (Fig. 16): Mentum and coxisternal region medially 
and laterally foveolate. Ventral plate pustulate medially and foveolate laterally. 



856 



SANDOR MAHUNKA 




Figs 15-18 

Klapperiches penicillus sp. n. - 15: body in dorsal aspect, 16: body in ventral aspect, 
17: pedotecta I, 18: podosoma in lateral aspect. 

Epimeral borders, excepting the posterior ones, conspicuous. Epimeral setae minute, 
epimeral setal formula: 1-1-3-3. Four pairs of genital, one pair of aggenital, two 
pairs of anal setae simple, the aggenital setae thickened, conspicuously ciliate, ad\ 
penicillate, like the notogastral setae. 

Remarks: On the basis of the position of the genital plates, the new 
species is readily classified with the hitherto monotypic genus Klapperiches 
Mahunka, 1978. It is distinguished from the type species (K. nigrisetosus Mahunka, 
1978) by the small and penicillate notogastral setae and the much longer sensillus. 



ORIBATIDS FROM ST. LUCIA 857 

Derivatio nominis: Named after the characteristic form of the 
notogastral setae. 

Dissorhina neotropicalis sp. n. Figs 19-21 

Material examined: Holotype: St. Lucia: STL-79/19; 6 paratypes: from the same 
sample. Holotype and 4 paratypes: MHNG, 2 paratypes: ( 1560-PO-96): HNHM. 

Measurements.- Length of body: 276-302 urn, width of body: 155-171 
urn. 

Prodorsum: Rostral apex separated by two incisions, as typical for the 
genus. Rostral setae arising on this small, triangular apex. Prodorsal costulae of 
typical design, composing a strong structure, but not reaching to the lamellar setae. 
Bothridium also well sclerotised, with two separate tubercles behind it. Sensillus 
directed outwards, with fusiform, well-dilated capitulum, 2-3 minute spicules visible 
on its distal end. 

Notogaster: Median part of the dorsosejugal region protruding 
anteriorly, straight medially (Fig. 19). Ten pairs of simple notogastral setae, c 2 short, 
setae h^ much longer than setae p 2 andp 3 . 

Lateral part of podosoma: Exobothridial surface smooth, 
granules visible only around the acetabula. A well sclerotised longitudinal lath, and 
the two tubercles behind the bothridium, mentioned above, observable. Exobothridial 
setae extremely strong (Fig. 21). 

Ventral regions (Fig. 20): Coxisternal plate well sclerotised. Epimeral 
borders linked by transversal crests resembling small bridges. Epimeral setae short, 
simple; setae 1c arising on a longitudinal crest, setae 3c the longest of all. Anogenital 
setae conspicuously short, minute (excepting the anteromedian pair of genital ones). 
Their number and position typical for this genus. 

Remarks: The new species stands very close to Dissorhina ornata 
(Oudemans, 1900). The two species are difficult to distinguish, because of the high 
variability of several characters. However the form of the sensillus is decisive in the 
separation of these two species. The sensillus of D. ornata is much longer than that of 
D. neotropicalis sp. n. and bacilliform, only gradually thickened distally (Fig. 22). In 
the new species the sensillus is fusiform and has a well-separated head. An other 
distinguishing feature could be the degree of sclerotization of the coxisternal region 
(stronger in D. neotropicalis). 

Derivatio nominis: This is the first Dissorhina species from the 
Neotropical region, all others are known from the Holarctic. 

Gittella insularis sp. n. Figs 23-26 

Material examined: Holotype: St. Lucia: STL-79/17; 3 paratypes: from the same 
sample. Holotype and 2 paratypes: MHNG, 1 paratype: (1561-PO-96): HNHM. 

Measurements.- Length of body: 576-642 urn, width of body: 255-297 
urn. 

Prodorsum: Rostral part elongated, rostrum rounded. Weak lamellar 
lines present, which reach over to the insertion point of the lamellar setae. Three pairs 



858 



SANDOR MAHUNKA 






Figs 19-22 

Dissorhina neotropicalis sp. n. - 19: body in dorsal aspect, 20: body in ventral aspect, 

21: podosoma in lateral aspect. 

Dissorhina ornata (Paoli, 1908). - 22: basal part of prodorsum in dorsal aspect. 



ORIBATIDS FROM ST. LUCIA 



859 




Figs 23-26 

Gittella insularis sp. n. - 23: body in dorsal aspect, 24: podosoma in lateral aspect, 25: body in 

ventral aspect, 26: leg I. 



860 SANDOR MAHUNKA 

of large spots and a pair of strong tubercles present in the interbothridial region (Fig. 
23). The latter directed posteriorly. Rostral setae thin and simple, arising near to the 
lateral margin. Lamellar setae conspicuously ciliated, rostral and interlamellar setae 
smooth. Sensillus slightly dilated, with 5-6 long flagellate branches. Exobothridial 
region granulate. 

Notogaster: Twelve pairs of long notogastral setae and the alveoli of 
setae c-> present. All notogastral setae with 4-6 comparatively long cilia. A pair of 
spots, near to the lyrifissures iad visible. 

Lateral part ofpodosoma: Pedotecta I narrow, setae lc arising on 
it. The tectum, below the acetabula, disconnected behind pedotecta I (Fig. 24). A 
weak polygonal sculpture anteriorly and some crests and granulated areas above the 
acetabula of leg II-III present. 

Ventral regions (Fig. 25): Some parts of the surface (e.g. along the 
apodemes) covered by cerotegument granules. Epimeral borders and apodemes 
conspicuous, op. 2 in straight transversal position, with a rounded thickening in front 
of it. Genital opening located comparatively posteriorly, epimera IV completely 
framing it. This surface has a polygonal sculpture. Epimeral setal formula: 3 - 1 - 3 - 
3. Some of these setae conspicuously ciliate. Anogenital setal formula: 5-1-2-3. 
Adanal setae with long cilia. Lyrifissures iad in inverse apoanal position. 

Legs: All joints of legs thick, somewhat narrowed at both ends. Especially 
thick is the femur of leg I (Fig. 26). Their setal formula typical for the family: 

I: 1-5-2+1 -4+2-20+2- 1 

Remarks: The new species fits well into the genus Gittella Hammer, 
1961. It is distinguished from the other species by the size of the prodorsal condyles, 
the length of the notogastral setae and the form of the sensillus. 

Derivatio nominis: It is named for its island locality. 

Moritzoppia subfallax sp. n. Figs 27-29 

Material examined: Holotype: St. Lucia: STL-79/17; 5 paratypes: from the same 
sample. Holotype and 3 paratypes: MHNG, 2 paratypes: (1562-PO-96): HNHM. 

Measurements.- Length of body: 266-317 urn, width of body: 145-171 
urn. 

Prodorsum: Rostrum rounded, ciliated, rostral setae arising on the 
prodorsal surface. Well developed, inverse Y-shaped costulae. A pair of other laths 
run forwards from the bothridia, they do not connect with the costula. Some small 
tubercles visible in front of them. One pair of V-shaped laths also present in the 
interbothridial region basally. Lamellar and interlamellar setae smooth, thinner and 
shorter than the rostral ones. Bothridia with basal tubercles. Sensillus asymmetrically 
expanded, with short spines unilaterally. 

Notogaster: Dorsosejugal suture nearly straight medially, carina and a 
pair of humeral apophyses conspicuous. Setae c 2 not shorter than the other nine pairs 
of notogastral setae (Fig. 27). 

Lateral part ofpodosoma: Well granulated, with some stronger 
laths (Fig. 29). Exobothridial setae hardly shorter than the interlamellar ones. 



ORIBATIDS FROM ST. LUCIA 



861 




Figs 27-30 

Moritzoppia subfallax sp. n. - 27: body in dorsal aspect, 28: body in ventral aspect, 

29: podosoma in lateral aspect. 

Lauroppia fallax (Paoli, 1908). - 30: prodorsum in dorsal aspect. 



862 SANDOR MAHUNKA 

Ventral regions: Coxisternal region well sclerotised, most of the 
apodemes and borders well visible. A characteristic, serrate, transversal crista present 
along the posterior border of the coxisternal region. Sejugal borders thick, with 
longitudinal, parallel lines. Epimeral setae simple, thin, setae lc arising laterally on 
the margin of pedotecta I (Fig. 28). Anogenital setal formula: 4-1-2-3. Setae ad\ 
in postanal, setae ad^ in preanal position. The latter inserted closer to the aggenital 
setae than to other adanal setae. 

Remarks: The habitus, primarily the prodorsal and notogastral structure 
closely resembles the type species of the genus Lauroppia Subias & Rodriguez, 1986 
(cf. Fig. 30: fallax Paoli, 1908). But on the basis of the number of genital setae and 
the well developed setae c 9 , the new species would be better be placed in the genus 
Moritzoppia Subias & Rodriguez, 1988 (Subias & Balogh 1989), however I consider 
this as a somewhat provisional solution. 

Derivatio nominis: It is named after the closely related species. 

Oxyoppia antillensis sp. n. Figs 31-33 

Material examined: Holotype: St. Lucia: STL-79/13; 7 paratypes: from the same 
sample; 2 paratypes: STL-79/6. Holotype and 6 paratypes: MHNG, 3 paratypes: (1563-PO-96): 
HNHM. 

Measurements.- Length of body: 193-21 1 p.m, width of body: 91-102 
urn. 

Prodorsum. Rostrum widely rounded, rostral setae arising laterally on 
small tubercles. Between them an arched line observable. Costulae reaching far out to 
the insertion of the interlamellar setae, a weak transcostula seen between their 
insertions. A pair of well-seclerotised basal tubercles passing into a longitudinal crest 
present in the interbothridial region, they frame two pairs of round spots and the 
interlamellar setae (Fig. 31). A pair of well-developed and arched lateral laths, 
directed toward the costula. Nearly the whole surface granulated. Sensillus large, 
asymmetrically dilated and unilaterally spinose, pectinate. 

Notogaster: Dorsosejugal suture arched medially, with a well-developed 
crista. Humeral processes normal, standing opposite to the posterior bothridial 
tubercles. Ten pairs of notogastral setae present, they are all slightly thickened, 
bacilliform and pilose, excepting setae c 1 . 

Lateral part of podosoma: Well sclerotised, the surface mostly 
granulated (Fig. 33). 

Ventral regions (Fig. 32): Epimeral borders conspicuous, epimeral 
fields separated by thick bands. Epimeral fields ornamented by a polygonal sculpture. 
A sharp, arched minitectum present along the posterior border, near the genital 
opening. Anogenital setal formula: 5 - 1 - 2 - 3. All setae short (excepting the anterior 
genital ones), setae ad^ in postanal, setae ad^ in preanal position. 

Remarks: The new species stands nearest to Oxyoppia pilosa Balogh & 
Mahunka, 1981 described from Paraguay but differs from it by the conspicuously 
long crista, the longer costulae, the position of the weak transcostula and by the 
strongly arched laths near to the posterior epimeral borders. 



ORIBATIDS FROM ST. LUCIA 



863 




Figs 31-33 

Oxyoppia antillensis sp. n. - 31: body in dorsal aspect, 32: body in ventral aspect. 

33: podosoma in lateral aspect. 

Derivatio nominis: This is the first Oxyoppia species known from 
the Antilles. 



Parakalumma piton sp. n. Figs 34-36 

Material examined: Holotype: St. Lucia: STL-79/7; 6 paratypes: from the same sample: 
4 paratypes: STL-79/3; 2 paratypes: STL-79/6; 2 paratypes: STL-79/12; 3 paratypes: STL- 
79/17. Holotype and 1 1 paratypes: MHNG, 6 paratypes: (1566-PO-96): HNHM. 

Measurements.- Length of body: 291-376 pm (female), 331-352 pm 

(male), width of body: 160-21 1 pm (female), 175-181 pm (male). 



864 



SANDOR MAHUNKA 





36 



Figs 34-37 

Parakalumma piton sp. n. - 34: body in dorsal aspect (9), 35: body in dorsal aspect (S), 

36: lamella and sensillus ( 9 ). 

Parakalumma foveolata Balogh & Mahunka, 1969. - 37: lamella and sensillus ($). 



ORIBATIDS FROM ST. LUCIA 865 

Prodorsum: (Female): Rostrum rounded, rostral setae simple. Lamellae 
with a small anterolateral incision and a spine (Fig. 34). Lamellar setae twice as long 
as the rostrals, interlamellar setae conspicuous. Sensillus comparatively short, its 
capitulum wide. (Male): Lamellae without anterolateral incision and spine, gradually 
narrowing anteriorly. Sensillus thinner and longer (Fig. 35) than in the female. 

Notogaster: In both sexes: Ten pairs of minute notogastral setae. Four 
pairs of sacculi present, So much larger than the others and their inner surface 
punctate (resembling a porose area). Pustulate sculpture in males (Fig. 35). 

Lateral part of podosoma: Tutorium very short, without cusp, a 
conspicuous line runs from acetabulum I to the insertion of rostral seta. Pedotecta I 
and II-III normally developed, discidium large. Circumpedal carinae conspicuous, 
reaching the lateral margin of ventral plate and running far anteriorly. 

Gnathosoma: Palpal femur with strong transversal ridges. Palp setal 
formula: 2 - 1 - 3 - 9+1. Eupathidium acm and the solenidium fused with each other. 

Ventral regions: Median part of the coxisternal region ornamented by 
weak foveolae, on the ventral plate some rounded, but even weaker foveolae present. 
Epimeral setal formula: 3 - 1 - 3 - 3. Anogenital setal formula: 4 - 1 - 2 - 3. Lyri- 
fissures iad in adanal position. All setae in the ventral regions short and simple. 

Legs: All legs monodactylous, with large claws. Seta ft" and e inserted 
behind solenidia coj and co 2 . cpj and cp 9 arising on a projection. A well developed 
blade-like formation on femora III and IV ventrally. 

Remarks: The new species stands very near Parakalumma foveolaîa 
Balogh & Mahunka, 1969. Re-examination of a paratype of this species shows that it 
can unambiguously be distinguished from the new species by the form of the lamellae 
(Fig. 37) in the female and the lacking of the peculiar pustulate sculpture of the 
notogaster (shown in Fig. 35) in the male. In both sexes of the new species the 
foveolate sculpture is much weaker than in P.foveolata. This species differs also from 
the new one by the form of the sensillus (much thinner in P. foveolata) and the length 
and ratio of the prodorsal setae (much longer in the female of the new species, 
Fig. 36). 

Derivatio nominis: named after the characteristic mountain of 
St. Lucia Island. 

Unguizetes similis sp. n. Figs 38-41 

Material examined: Holotype: St. Lucia: STL-79/19; 4 paratypes: from the same 
sample; 1 paratype: STL-79/1. Holotype and 3 paratypes: MHNG, 2 paratypes: (1564-PO-96): 
HNHM. 

Measurements.- Length of body: 560-691 urn, width of body: 477-544 
urn. 

Prodorsum: Rostrum nearly triangular in dorsal view. Lamellae with 
comparatively long and narrow cusps (Fig. 41), their outer apex sharply pointed. 
Rostral, lamellar and interlamellar setae simple, setiform, somewhat ciliated. Sensillus 
directed outwards and backwards, with a long pedicel and a small, finely ciliate 
capitulum. 



866 



SANDOR MAHUNKA 




Figs 38-42 

Unguizetes similis sp. n. - 38: body in dorsal aspect, 39: body in ventral aspect, 

40: podosoma in lateral aspect, 41: lamellar cusp. 

Unguizetes incertus Balogh & Mahunka. 1969. - 42: lamellar cusp. 



ORIBATIDS FROM ST. LUCIA 867 

Notogaster: Surface finely punctate. Dorsosejugal suture faint between 
the insertion of interlamellar setae. Four pairs of small and round areae porosae and 
ten pairs of setal alveoli present on the notogaster (Fig. 38). 

Lateral part ofpodosoma (Fig. 40): Tutorium well developed, with 
a short cusp, rostral seta inserted close to it. Exobothridial setae conspicuously long 
and thin. Pedotecta II-III and the discidium large. 

Ventral regions (Fig. 39): Apodemes short or absent (excepting the 
sejugal ones). Epimeral borders hardly discernible. All epimeral setae long, but their 
length variable. Anogenital setal formula: 6 - 1 - 2 - 3. The setae in this region short 
and simple. A band of areae porosae observable along the margin of the ventral plate, 
beginning at the connecting of the circumpedal carina. 

Legs: All legs tridactylous, haterodactyly present. Solenidia of tibia I 
inserted on a protuberance. Femora of legs II-IV wit a blade-like formation ventrally. 

Remarks : The new species stands nearest to Unguizeies incertus Balogh 
& Mahunka, 1969. It is distinguished from the latter by the absence of the 
characteristic, polygonal sculpture, by the much narrowed lamellae running near to 
each other (broadened in U. incertus) and by the form of the lamellar cusps (for U. 
incertus see in Fig. 42). In U. incertus the dorsosejugal suture is also faint medially, 
but not absent, as mentioned in the original description. 

Derivatio nominis: The species is similar to Unguizetes incertus 
(Balogh & Mahunka, 1969). 

Eupelops spongiosus sp. n. Figs 43-44 

Material examined: Holotype: St. Lucia: STL-79/8: MHNG. 

Measurements.- Length of body: 476 urn, width of body: 372 urn. 

Prodorsum: Its size typical for the genus. Interlamellar region U-shaped, 
wide. Lamellar surface finely foveolated. Tutorium also wide, well developed (Fig. 
43). Rostral and lamellar setae distinctly barbed, interlamellar setae very large, 
phylliform. Sensillus short, its capitulum fusiform. 

Notogaster: Cerotegument thick, excepting the well separated 
notogastral lenticulus and the lateral part of the pteromorphae. It seems to be foveo- 
late. Pteromorphae large, the hinge-like line conspicuous. Anterior notogastral tectum 
covering the basal part of prodorsum, its anterior margin waved. Ten pairs of mostly 
dilated notogastral setae present, setae Ip and h^ stand close to one another. The 
former much smaller than the latter (Fig. 43). Owing to the cerotegument sculpture I 
was unable to find all four pairs of areae porosae and the position of the lyriffisures. 

Ventral regions: Mentum ornamented by mostly longitudinal, 
anteriorly converging striations. Coxisternal and ventral surface covered by thick 
cerotegument. Its sculpture similar to that of the notogaster; therefore, most of the 
epimeral setae invisible (Fig. 44). 

Legs: All legs tridactylous, a strong heterodactyly present. 

Remarks: The new species is well characterised by the sculpture of the 
cerotegument layer. Within the species group which is characterised by the closely 



SANDOR MAHUNKA 




Figs 43-44 
Eupelops spongiosus sp. n. -43: body in dorsal aspect, 44: body in ventral aspect. 

adjacent setae Ip and /? 3 , a similar sculpture is known in E. foveolatus Engelbrecht, 
1975 described from South Africa. However, the sensillus of the latter species is 
much shorter and its head is truncate. 

Derivatio nominisi named after the characteristic sculpture of the 
notogaster. 



Truncozetes rugosus sp. n. 



Figs 45-46 



Material examined: Holotype: St. Lucia: STL-79/1; 3 paratypes: from the same sample. 
Holotype and 2 paratypes: MHNG, 1 paratype: (1565-PO-96): HNHM. 



ORIBATIDS FROM ST. LUCIA 



869 



^3 




Figs 45-46 
Truncozetes rugosus sp. n. - 45: body in dorsal aspect, 46: body in ventral aspect. 

Measurements.- Length of body: 233-244 urn, width of body: 171-176 
urn. 

Integument: Nearly the whole surface covered by cerotegument 
granules which, on some parts of body (e.g., dorsosejugal region), is more distinct 
than on other parts. Cuticle foveolate or punctate, mentum with irregular longitudinal 
rugae. 



870 SANDOR MAHUNKA 

Prodorsum: Rostrum elongated, rounded. Lamellae wide, obliquely 
truncate anteriorly. Translamella narrow, almost linear (Fig. 45). Lamellar and rostral 
setae thin setiform, interlamellar ones minute. Sensillus fusiform, its distal end elon- 
gate, this part much darker than the rest. Bothridium opened anteriorly. 

Notogaster: Surface well foveolate. One pair of sacculi (?) and ten pairs 
of short notogastral setae present. A weak but conspicuous elevation in posteromar- 
ginal position. 

Ventral regions (Fig. 46): Mentum rugose and foveolate. Epimeral 
region ornamented by smaller, ventral plate by larger, foveolae. The smaller foveolae 
sometimes confluent. Epimeral borders and some apodemes also conspicuous, bo 2 
forming a transversal band, the others compose a network in front of the genital 
opening. Epimeral setae hardly visible, minute. Six pairs of genital setae present. 

Legs: All legs tridactylous, a strong heterodactyly present. 

Remarks: The new species stands near to the type species of the genus, 
Truncozeles mucronatus Balogh & Mahunka, 1969. However it is clearly distingu- 
ishable by the absence of the longitudinal pattern on the coxisternal region and the 
much weaker and irregular sculpture on the mentum, which consists of regular longi- 
tudinal sulci in T. mucronatus. 

Derivatio nominis: This species is named after the sculpture of the 
mentum. 



Paracarinogalumna gen. n. 

Diagnosis: Family Galumnidae. Prodorsal surface with a well 
sclerotised, sharp, median keel, resembling a crest or carina. Lamellar lines protruding 
anteriorly and composing a keel-shaped formation like a costula and similar to the 
median keel. All three crests run to the rostral apex, which is characteristically 
triangular. Sublamellar lines strong, normally developed, well arched. Lamellar setae 
arising on lamellar keel, rostral setae very near to it. Dorsosejugal suture complete. 
Notogaster with 4 pairs of areae porosae, 10 pairs of setal alveoli (true setae absent). 
Median pori absent. Coxisternal region normal, pedotecta I sharply pointed. 
Anogenital setal formula: 6 - 1 - 2 - 3. Lyriffissures iad located far laterally from the 
anal aperture. Porose area postanalis absent. All legs tridactylous, heterodactyly weak. 
Solenidium co 2 and e stand behind coj. 

Type species: Paracarinogalumna genavensium sp. n. 

Remarks: The lamellar line is formed like a blade. This feature, along 
with the median keel, is known in the family Galumnidae {Carino galumna Engel- 
brecht, 1973). The new taxon stands close to this genus, although the blade-like 
formation is only partial and it is the lamellar line which continues in an arched, sharp 
blade not bearing the lamellar seta; furthermore, the sublamellar line is reduced, or at 
least partly missing, while the porose area postanalis is completely lacking. The genus 
Carino galumna has median pori and the lyrifissure iad is in the adanal position; on 
the other hand, in the new species, the median pori are missing, while lyrifissure iad 



ORIBATIDS FROM ST. LUCIA 871 

is far removed from anal aperture. Consequently, it is not considered here as a 
member of the South African genus C arino galumna. 

Derivatio nominis: The new genus is very near to Carino galumna 
Engelbrecht, 1973. 

Paracarinogalumna genavensium sp. n. Figs 47-51 

Material examined: Holotype: St. Lucia: STL-79/8; 9 paratypes: from the same sample; 
18 paratypes: STL-79/1; 1 paratype: STL-79/5; 4 paratypes: STL-79/ll;2 paratypes: STL- 
79/15; 1 paratype: STL-79/17. Holotype and 23 paratypes: MHNG, 12 paratypes: (1569-PO- 
96): HNHM. 

Measurements.- Length of body: 766-865 urn, width of body: 589-642 
urn. 

Prodorsum: Rostral apex sharply pointed. Lamellar keel running toward 
the rostral apex but not reaching it. The position of the rostral and lamellar setae as 
shown in Fig. 50. Interlamellar setae reduced, hardly visible or represented only by 
their alveoli. Sensillus lanceolate, its head with some spicules (Fig. 47). 

Notogaster: Dorsosejugal suture complete. Four pairs of porose areae 
present, Aa gradually widened to the lateral margin of notogaster. A 3 narrow, long. 
Median pori absent, lyrifissures im located near to porose area A j . 

Lateral part of podosoma: As shown in Fig. 50. Circumpedal 
carina reaching to the lateral margin of the ventral plate. 

Ventral regions (Fig. 48): Coxisternal region without spots or other 
sculpture. All setae minute, hardly discernible. Epimeral setal formula: 1 - - 2 - 2 . 
Only two pairs of genital setae arising on the anterior margin of the plates, the others 
inserted along longitudinal lines. Aggenital, anal and adanal setae also minute, setae 
ad^ arising in front of lyrifissures iad. 

Legs: All legs tridactylous, a weak heterodactyly present. End of lateral 
claws slightly dilated. The position of the solenidial group of leg I as shown in Fig. 
51. Genu IV conspicuously long, bearing two very long setae. 

Legs setal formulae: 

I: 1 -5-3+1 -4+2-20+2-3 
IV: 1 -2-2-3+1 - 12-3 

Remarks: See the remarks after the generic diagnosis. 

Derivatio nominis: In honour of the staff of the Geneva Museum 
and especially of T. Jaccoud and L. de Roguin. 

Pergalumna cucheae sp. n. Figs 52-55 

Material examined: Holotype: St. Lucia: STL-79/1; 17 paratypes: from the same 
sample. Holotype and 10 paratypes: MHNG, 7 paratypes: (1567-PO-96): HNHM. 

Measurements:- Length of body: 650-758 urn, width of body: 460-552 
urn. 

Prodorsum: Rostral part characteristically modified, rostral apex pro- 
truding in a wide nose forwards, with a weak but wide transversal band behind it. 
Both anterior prodorsal setae long, thin, nearly equal in length. Interlamellar setae 
represented only by their alveoli. Sensillus setiform, smooth, simple. 



872 



SANDOR MAHUNKA 




Figs 47-48 

Paracarinogalumna genavensium gen. n., sp. n. - 47: body in dorsal aspect, 48: body in ventral 

aspect. 



ORIBATIDS FROM ST. LUCIA 



873 




Figs 49-51 

Paracarinogalumna genavensium gen. n., sp. n. - 49: podosoma in lateral aspect, 50: lamellar 
line, 5 1 : solenidial group of leg I. 



Notogaster (Fig. 52): Dorsosejugal suture interrupted between the round 
sejugal porose area. Structure hy conspicuously large. Three pairs of areae porosae 
and 10 pairs of setal alveoli present, no essential difference among the areae porosae. 

Lateral part of podosoma: Lamellar and sublamellar line well 
developed, gradually arching toward the margin of the coxisternal region. They run 
parallel to each other. Anterior margin of mentum very thick (Fig. 55). Circumpedal 
carina reaching to the lateral margins of the ventral plate. 

Ventral regions (Fig. 54): Epimeral surface without characteristic 
sculpture. All epimeral setae simple and conspicuous, epimeral setal formula: 1-0-2 
- 3. Three pairs of genital setae inserted on the anterior margin of the genital plates. 
Aggenital, anal and adanal setae minute, hardly visible. Their position typical for the 
family. 



874 



SANDOR MAHUNKA 




Figs 52-55 

Pergalumna cucheae sp. n. - 52: body in dorsal aspect, 53: solenidial group of leg I, 54: body 
in ventral aspect. 55: podosoma in lateral aspect. 



ORIBATIDS FROM ST. LUCIA 875 

Legs: The solenidial group of leg I as shown in Fig. 53. 

Remarks: The new species is well characterised by the form of the 
rostrum. It slightly resembles Pergalumna elencata (Berlese, 1914), which was 
designated by Pérez-Inigo & B aggio (1994) as the type of the genus Pseudo- 
galumna. However, the rostrum of the new species is rounded and, without a keel. 
The three pairs of notogastral porose areas are also typical for the genus Pergalumna 
Grandjean, 1936. Therefore it fits well to the latter genus. 

Derivatio nominis: I dedicate the new species to Mrs T. Cuche 
(Geneva Museum) in recognition of her invaluable help during my stay in Geneva. 

Pilogalumna antillensis sp. n. Figs 56-59 

Material examined: Holotype: St. Lucia: STL-79/82; 7 paratypes: from the same 
sample. Holotype and 4 paratypes: MHNG, 3 paratypes: (1568-PO-96): HNHM. 

Measurements.- Length of body: 683-78 1 urn, width of body: 526-622 
urn. 

Prodorsum: Rostrum simple, conical. Lamellar lines absent, sublamellar 
lines (S) well developed, arched downwards and fused with the epimeral margin (Fig. 
59). Prodorsal setae short or reduced, rostral setae longer than the lamellar ones. Inter- 
lamellar setae represented only by their alveoli. Sensillus directed forwards, narrow 
lanceolate, with a few capitular spicules. 

Notogaster: Dorsosejugal suture complete, the thickening behind it (hy) 
round (Fig. 56). Notogaster smooth, pteromorphae with some radiate crests. Four 
pairs of comparatively small porose areae present. Aa and A 3 characteristically elon- 
gated, A j irregular. Ten pairs of minute notogastral setae or their alveoli present, at 
least setae c 2 , da and/?j clearly visible. 

Lateral part of podosoma: Pedotecta I small, pedotecta II-III 
normal, discidium rounded laterally. Circumpedal carina short, ending far from the 
lateral margin on the ventral plate. 

Ventral regions (Fig. 58): Epimeral surface hardly ornamented, only a 
few spots visible. Epimeral setal formula: 1 - - 2 - 1, all setae minute. Anogenital 
setal formula: 6 - 1 - 2 - 3, all setae also minute. Three pairs of setae concentrated on 
the anterior margin of genital plates. Lyrifissure iad in adanal position. 

Legs: All legs tridactylous, a weak heterodactyly observable. Both solenidia 
of tarsus I inserted near to each other, in a transversal line (Fig. 57). 

Remarks: The classification of this species is rather problematic, because 
the absence of the sublamellar lines is one of the main characteristics of the genus 
Pilogalumna Grandjean, 1956. They are well developed in this species! However, on 
the basis of the other characters (minute notogastral setae, position of e on tarsus I) it 
fits well into this group. Further studies are necessary to decide the final position of 
this species. Hitherto, the genus has not been recorded from the Neotropical region. 

Derivatio nominis: Named after the zoogeographical region, where 
the species was collected. 



876 



SANDOR MAHUNKA 




Figs 56-59 

Pilogalumna antillensis sp. n. - 56: body in dorsal aspect, 57: solenidial group of leg I, 
58: body in ventral aspect, 59: podosoma in lateral aspect. 



ORIBATIDS FROM ST. LUCIA 877 



ACKNOWLEDGEMENTS 

I would like to express my gratitude to Mr T. Jaccoud and to late Dr L. de 
Roguin (1948 - 1998), both collaborators of the Geneva Museum, for their assistance 
in 1979. In that year my wife and I planned and paid for a trip to St. Lucia, but we did 
not get a visa. At the last minute T. Jaccoud and L. de Roguin offered to undertake - 
at their own expense - this collecting trip instead of us and helped us in this way to 
save our money and postpone our trip for the following year (1980). This exemplary 
help between colleagues should be especially emphasized. 

I am also very grateful to Dr Malcolm Luxton (National Museum of Wales, 
Cardiff) for critical reading of the manuscript and for his valuable suggestions. 

REFERENCES 

Balogh, J. & Mahunka, S. 1974. A foundation of the Oribatid (Acari) fauna of Cuba. Acta 

Zoologica Academiae Scientarum Hungaricae 20: 1-25. 
Grandjean, F. 1929. Quelques nouveaux genres d'Oribatei du Venezuela et de la Martinique. 

Bulletin de la Société Zoologique de France 54: 400-423. 
Grandjean, F. 1930. Oribates nouveaux de la région Caraïbe. Bulletin de la Société Zoologique 

de France 55: 262-284. 
Mahunka, S. 1985. Mites (Acari) from St. Lucia (Antilles). 2. Oribatida. Acta Zoologica 

Hungarica 31: 119-178. 
Mahunka, S. 1994. Oribatids from Madagascar II. (Acari: Oribatida). (New and interesting 

mites from the Geneva Museum LXXIX.). Revue suisse de Zoologie 101 : 47-88. 
Niedbala, W. 1985. Essai critique sur Mesoplophora (Acari, Oribatida, Mesoplophoridae). 

Annales Zoologici 39: 93-1 17. 
Pérez-Inigo, C. & Baggio, D. 1994. Oribatides édaphiques du Brésil (Vili). Oribates de l'état 

de Sào Paulo (Cinquième partie). Acarologia 35: 181-198. 
Subîas, L. S. & Balogh, P. 1989. Identification keys to the genera of Oppiidae Grandjean, 

1951 (Acari: Oribatei). Acta Zoologica Hungarica 35: 355-412. 
Willmann, C. 1933. Zoologische Ergebnisse einer Reise nach Bonaire, Curaçao and Aruba im 

Jahre 1930. No. 10. Trimalaconothrus pilipes, eine neue Oribatide aus Westindien. 

Zoologische Jahrbücher. Abteilung für Systematik 64: 447-452. 



Revue suisse de Zoologie 105 (4): 879-899; décembre 1998 



Japygoidea (Diplura) du Sud-Est asiatique n° 8: Indonésie 
(Java, Bali), Singapour et Brunei 
- Dicellurata Genavensia XXIII - 

Jean PAGES 1 

51, rue du Faubourg Saint-Martin, F-21 121 Fontaine-lès-Dijon, France. 

Japygoidea (Diplura) from South-East Asia n° 8: Indonesia (Java, 
Bali), Singapore and Brunei - Dicellurata Genavensia XXIII. - In this 
note are studied 14 specimens of Parajapygidae collected in 1987 and 
1988. Parajapyx (Grassjapyx) sabahnus Pgs, formerly described from 
Sabah (Malaysia), exists in Singapore where it may have been imported. 
Three new species are recognized: P. (Parajapyx) hauseri n. sp., P. 
(Grassjapyx) temburong n. sp. from Brunei, and P. (Grassjapyx) reniformis 
n. sp. from Java and Bali. One specimen from Teraja (Brunei) shows a 
mixture of characters which hinders me to describe it as a n. sp. until 
further specimens will be collected in the same area. New nomenclature 
and definition of the different instars of a and 9 are proposed. The 
subgeneric position of Parajapyx hwashanensis Chou, Parajapyx yangi 
Chou and Parajapyx jinghongensis Xie & Yang is precised. The possibility 
for Parajapyx paucidentis Xie et al. to be an anomaly or a casual mutation 
of Parajapyx (Parajapyx) isabellae (Grassi) is discussed. 

Key-words: Diplura - Parajapygidae - Indonesia - Singapore - Brunei - 
Taxonomy - New species - Instars. 

INTRODUCTION 

Au cours de ses missions entomologiques dans le Sud-Est asiatique, l'équipe 
du Département des Arthropodes et d'Entomologie I du Muséum d'histoire naturelle 
de Genève (B. Hauser et C. Lienhard), a recueilli 14 représentants de la famille des 
Parajapygidae. Cette famille cosmopolite n'était connue de ces régions que par les 
Parajapyx (Grassjapyx) sepilok et P. (G.) sabahnus que j'ai décrit du Sabah 
(Malaysia) (Pages 1987). 

A ma connaissance, douze autres taxons peuvent être cités d'Asie et des îles du 
Pacifique. L'espèce cosmopolite, Parajapyx (P.) isabellae (Grassi) a été signalée de 
Chine, du Japon et des îles Hawaï (Silvestri 1928, Chou 1966, Zwaluwenburg 
1934, Zimmerman 1948). 



Professeur emèrite de l'Université de Bourgogne, Equipe d'Ecologie et Dynamique 
des Populations, F-21 100 Dijon. 

Manuscrit accepté le 01.07.1998 



880 JEAN PAGES 

L'énigmatique P. (G.) grassianus var. indica Silv. du Sikkim (Silvestri 1913) 
est l'unique Parajapygidé signalé de la région indienne. D'Australie, Womersley 
(1934. 1945) a fait connaître P. (P.) swani Worn. d'Australie occidentale et P. (G.) 
queenslandica Worn, du Queensland; enfin, des îles du Pacifique, on peut citer, outre 
l'ubiquiste Parajapyx (P.) isabellae (Grassi) des îles Hawaï, une 9 d'un Parajapyx 
(Grassjapx) sp. des îles Marianes du Nord (Nakamura 1994) et P. (G.) samoanus 
Silv. de l'archipel des Samoa (Silvestri 1930). 

De Chine, 5 espèces et une "var." ont été décrites: Parajapyx (P.) emeryanus 
Silv. et sa "var." centralis Silv. (Silvestri 1928. Chou 1966); Parajapyx hwasha- 
nensis Chou du Shensi et Parajapyx yangi Chou de Pékin (Chou 1966), ainsi que 
Parajapyx jinghongensis Xie & Yang du Yunnan (XiE & Yang 1990) ont été décrits 
sans indication de sous-genre; il me semble que les marges internes des cerques 
figurés justifient leur attribution au sous-genre Grassjapyx Pgs: Parajapyx (Grass- 
japyx) hwashanensis Chou. Parajapyx (Grassjapyx) yangi Chou, Parajapyx (Grass- 
japyx) jinghongensis Xie & Yang. 

Quant au Parajapyx paucidentis décrit récemment par Xie et al. (1990) du 
Zhejang. je ne crois pas que l'on puisse le considérer comme une espèce valable, mais 
vraisemblablement plutôt comme une aberration ou une mutation accidentelle ou non 
de P. (P.) isabellae (Grassi) dont il semble posséder les antennes, la chétotaxie 
générale et surtout les organes subcoxaux latéraux. En effet, pratiquement tous les 
stades de régression de l'armature de la marge interne des cerques dans le groupe 
isabellae ont été décrits. Le premier exemple est peut-être celui du Parajapyx (P.) 
isabellae, variant IV d'El Golea (Maroc) (Pages 1954) dont la d^ droite n'est plus 
qu'un minuscule denticule. le reste de la marge étant normal 2 . Le cas le moins aty- 
pique de dégradation de la marge interne est celui que j'ai signalé (1952b) chez un 
Parajapyx (P.) normandi Pgs. de Tunisie dont le cerque droit montre des dents mal 
formées, peu saillantes, mais encore reconnaissables. On peut citer ensuite l'exem- 
plaire de Parajapyx (P.) isabellae (Grassi) figuré par Silvestri (1948a) chez lequel 
on ne distingue plus que quelques minuscules denticules à peine visibles. Parajapyx 
(= Hemijapyx) unidentatus Ewing de l'Alabama ne montrerait plus que la d^, le reste 
des marges étant totalement lisse (Ewing 1941). P. paucidentis pourrait être considéré 
comme le stade ultime de la régression, les marges ne présentant plus qu'un minus- 
cule denticule près de l'apex des cerques ne correspondant à aucune des dents 
normales. 

Des régressions semblables ont été signalées par Silvestri (1948a) d'une part 
chez un autre Parajapygidé. P. (G.) grassianus Silv. d'Amérique du Nord, dont les 
cerques sont dépourvus de toutes dents, mais avec 2 minuscules denticules près de 
l'apex; c'est le même type d'anomalie que présente paucidentis, mais dans l'autre 
sous-genre; d'autre part chez un Japyx solifugus Hai. d'Italie dont la marge interne du 
cerque droit est complètement dépourvue d'armature. Il me semble inutile de nommer 



- La fig. 6 représentant cette anomalie a été attribuée par erreur à un variant IV d'In 

Salah. 



JAPYGOIDEA DU SUD-EST ASIATIQUE ggl 

de telles anomalies à moins que la même station ne fournisse, au fil des années, que 
de ces individus apparemment "hors normes". 

C'est pourquoi la synonymie Parajapyx paucidentis Xie et al. = Parajapyx 
(Parajapyx) isabellae (Grassi) ne peut être établie avec certitude actuellement. 

Dans cette note, je répartis les 14 spécimens de Parajapygidés récoltés en 1987 
à Java et Bali (Indonésie) et en 1988 à Singapour et au Brunei entre 4 espèces dont 3 
inédites: Parajapyx (P.) hauseri n. sp. et Parajapyx (G.) temburong n. sp. du Brunei, 
Parajapyx (G.) reniformis n. sp. de Java et Bali et Parajapyx (G.) sabahnus Pgs de 
Singapour; enfin un spécimen de Teraja au Brunei présente des caractères ambigus et 
trop peu nets qui ne me semblent pas permettre pour l'instant une détermination 
précise. 

La découverte chez Parajapyx (P.) hauseri n. sp. d'un nouveau stade du déve- 
loppement postembryonnaire des S , de celle du plus jeune stade 9 connu et chez 
Parajapyx (G.) reniformis n. sp. du stade 9 le plus âgé me font proposer une nouvelle 
nomenclature des stades pour les deux sexes. 

Tous les spécimens étudiés dans ce travail sont conservés dans les collections 
du Muséum d'histoire naturelle de Genève. 

DESCRIPTION DES ESPÈCES 

On trouvera la description des chétotaxies typiques, la définition des rapports 
et des abréviations utilisés dans Pages 1952a, 1952b et 1954. La nouvelle numé- 
rotation des stades du développement postembryonnaire des S et des $ est exposée 
ci-après. 

Parajapyx (Parajapyx) hauseri n. sp. Figs 1-18 

Matériel étudié 3 : Bru-88/24: Brunei (Brunei-Muara District): "Berakas Forest 
Reserve" au nord de Bandar Seri Begawan sur la route, à 19,5 km de Muara (= à 102,5 km de 
Muala Bêlait), forêt "Kerangas" (= "Tropical heath forest"), prélèvemenet de sol au pied de 
Casuarina nobilis Whitmore (Casuarinaceae), 30 m; 20.XI.1988; leg. B. Hauser (B-,); holo- 
type: S 4n de 2,14 mm; paratypes: 1 S 3n de 1,94 mm, 1 â 2n de 1,66 mm, 1 2 st. 1 de 1,55 
mm, 1 9 de 2,02 mm, 1 9 st.4 de 2,22 mm, 1 sexe? (urites 8 à 10 et les cerques absents). 

TÊTE 

Vertex: chétotaxie typique. 

Pli oral: sans la soie 4. 

Antennes: de 18 articles assez pileux, pas d'aires pileuses différenciées; 
sensilles recourbées présentes à partir du 5e article; 4 sensilles placoïdes en position 
typique sur l'article apical. 

Pièces buccales: typiques du genre. 



3 Basé sur les listes établies par le Dr B. Hauser pour les stations de récolte des 
missions de 1987 et 1988 en Insulinde. Les spécimens ont été capturés soit directement à vue, 
soit après traitement des prélèvements par entonnoir Berlese, soit à Bogor (Java) (Bj). à Bandar 
Seri Begawan (Brunei) (B 2 ), Hong Kong (B 3 ) ou à Genève (B 4 ). 



882 



JEAN PAGES 




Figs 1-6 

Parajapyx (Parajapyx) hauseri n. sp., S holotype. - 1. Vertex, e = 105 pm. - 2. Pronotum, e = 
105 [am. - 3. Mésonotum. e = 105 pm. - 4. Métanotum, e = 105 p.m. - 5. Tergite 1, e = 105 pm. 
- 6. Tergites 7 à 10 et les cerques, e = 105 pm. 



japygoidea du sud-est asiatique 883 

Thorax 
Face tergale 

P r o n o t u m : les 5+5M typiques, M 3 près de 2 fois aussi longs que les 
autres M qui sont subégaux; S\ nulles. 

Mésonotum: Préscutum: 1 + 1 M assez longs. Scutum: les 5+5 M 
typiques, les M 3 les plus longs, les M 5 les plus courts, les 5+5 s typiques assez 
courtes, 1 + 1 soies supplémentaires entre Sj et s 2 et 1 + 1 autres entre s 4 et s 5 . 

M é t a n o t u m : Préscutum: 2+2 M assez longs. Scutum: M, s et soies 
supplémentaires comme au mésonotum. 

Face sternale 

Par rapport à la chétotaxie de P. (P.) isabellae (Grassi) prise comme type 
(Pages 1952a), les seules variations rencontrées sont la présence d'une soie supplé- 
mentaire au présternite du mésosternum ou l'absence de la soie médiane typique à 
celui du métasternum. 

Pattes: peu pileuses; tarses avec 2+2 soies sternales dont les 2 distales sont 
aiguës, dépassant de peu l'apex de la griffe postérieure; celle-ci est à peine plus 
longue que l'antérieure; unguiculus minuscule, aigu. 

Abdomen 

T e r g i t e 1 : Préscutum: 2+2 soies assez longues et environ 5+5 microsoies. 
Scutum: les 5+5 M typiques, les M 4 les plus longs, 2 fois aussi développés que les M 5 
qui sont les plus courts de tous; les 6+6 s typiques assez courtes, sauf les s 5 qui sont 
longues; on note sur certains spécimens, sans que le sexe ou le stade de développe- 
ment soit en cause, l'absence d'une des s 3 et/ou la présence d'une soie supplémentaire 
entre M 3 et s 6 . 

Tergites 2 à 7 : Préscutum: 1 + 1 + 1 M typiques; le â holotype présente 
2+2 M sur le préscutum du tergite 6. Scutum: les 8+8 M typiques longs, les M 9 et M 3 
les plus développés; les 7+7 s peuvent être présentes, l'une d'entre elles peut 
cependant manquer d'un côté ou de l'autre comme cela semble être la norme chez les 
espèces du genre; la $ st.l qui ne montre qu'une seule et unique paire de soies sur la 
papille génitale, n'est pourvue sur ces tergites que des seules s 1? SyS-j, de la s 1 gauche 
et d'une soie supplémentaire près de la s 7 gauche; cette soie supplémentaire peut être 
présente chez les individus plus âgés, ainsi que deux autres entre Mj et s 4 ou M 7 et 5v. 

Tergite 8: un peu moins de 1,25 fois aussi large que long (1/L = 1,25 chez 
le â holotype, 1,22 chez la 9 st.l, 1,16 chez la $ st.4); 7+7 M subégaux longs, ceux 
de la moitié postérieure du segment légèrement plus courts que ceux de la moitié 
antérieure; 4+4 s (s\ nulles) chez les 6 et la 9 st.4, seules 5 3 et s 5 sont présentes chez 
la $ st.l. 

Tergite 9 : 1/L = 2,24 en moyenne (2,25 chez l'holotype, 2,07 chez la 2 
st.l, 2,48 chez la $ st.4); les 3+3 M typiques longs; aucune 5 chez le S holotype; 
chez les autres spécimens, seules les s^ sont présentes si l'on admet que cette paire de 
phanères est venue se placer entre M i et M 2 . 



884 



JEAN PAGES 




Figs 7-14. Parajapyx {Parajapyx) hauseri n. sp., a holotype. - 7. Urosternite 1, e = 105 pm. - 
8. Urosternite 3, e = 105 pm. - 9. Hypopyge, e = 84 pm. - 10. 9 st.l, tergites 7 à 10 et les 
cerques, e = 105 pm. - 11. 9 st. 3, tergites 7 à 10 et les cerques, e = 105 pm. - 12. idem, 
acropyge, e = 51 pm. - 13. idem, angle externe de l'organe subcoxal gauche et style, e = 42 pm. 
- 14. idem, hypopyge, e = 51 pm. 



JAPYGOIDEA DU SUD-EST ASIATIQUE 



885 




Figs 15-18 

Parajapyx (Parajapyx) hauseri n. sp., évolution de l'armature de la marge interne des cerques 
suivant le sexe et le stade de développement. - 15. S holotype (= 6 4n ), e = 60 um. - 16. 6 3n , 
e = 60 um. - 17. 6 2rr e = 51 um. - 18. 9 st. 2, e = 46 um. 



T e r g i t e 10 : L/l = 1,61 en moyenne (v. ex. = 1,54-1,74), 1,56 chez le $ 
holotype; 6+1+6 M (M ] nuls) longs, le M 4 droit manque chez la 9 st.4; 4+4 s (s 4 
nulles) courtes, la Sj droite est nulle chez la 9 st.l; le phanère situé entre M 7 et s 2 
peut être interprété comme un M 4 qui est typiquement inséré au-dessus de la ligne 
joignant M 7 et s 2 , ce qui le place entre M 2 et M 7 , ou bien comme une s 3 typiquement 
située presqu' exactement entre M 7 et s 2 ; sur certains individus, se phanère peut être 
considéré comme un M 4 à droite et comme une s 3 à gauche ou inversement. Toutes 
les combinaisons sont possibles entre 6+6 M et 4+4 s ou 5+5 M et 3+3 s. 

Longueurs relatives des segments 8àl0: 63-33-100 en 
moyenne; 6 holotype: 61-33-100; S 2n . 61-29-100; 9 st.l: 64-35-100; 9 st.4: 64- 29- 
100. 

Acropyge: triangulaire, simple, aussi long que large à la base; chez le 6 
holotype il est beaucoup plus large que long, à sommet bifide et très en avant de la 
ligne joignant les condyles d'articulation des cerques; il ne s'agit pas d'un artefact, cet 
individu ayant été préparé et monté exactement comme tous ceux de la série; ceci est 
à rapprocher de ce que Silvestri (1929) représente chez son P. (P.) dorianus ou moi- 
même chez P. (P.) genavensium (Pages 1978). 

S t e r n i t e 1 : Préscutum: chétotaxie typique, la soie médiane peut être 
absente. Scutum: les 10+10 M typiques assez longs, M 5 , M 6 et M 9 un peu moins longs 
que les autres; 1+2+1 s (c nulles); 8+8 soies disposées sur deux rangées en avant des 
organes subcoxaux latéraux; on peut observer jusqu'à 4 soies supplémentaires à droite 
ou à gauche de la ligne médiane: 1 entre M x et M 2 , 1 entre M 5 et b, 1 ou 2 entre M 9 et b. 



gg6 JEAN PAGES 

Organes subcoxaux latéraux: ils occupent chacun environ le 
1/3 de la largeur interstylaire et sont peu saillants; de 14 à 22 soies glandulaires 
disposées sur une seule rangée, en moyenne SG/st^ = 1 (v. ex. = 0,96-1,04); 4 à 7 
soies sensorielles régulièrement espacées, SS/st^ = 0,6 en moyenne (v. ex. = 0,50- 
0,64); SG/SS = 1,90 chez l'holotype, 1,61 en moyenne chez les autres exemplaires. 

La partie médiane postérieure du sternite porte les 1 + 1 minuscules soies 
typiques. 

Sternites 2à7: Préscutum: les 5+1+5 soies typiques assez longues. 
Scutum: les 12+12 M typiques assez longs, 4+3+4 s, la c peut faire défaut comme au 
sternite 1. 

Styles: typiques du genre, le cône secondaire aigu, égal aux 2/5 de la 
longueur du cône principal; aux sfj_ 3 la soie recourbée (s') est subégale à la soie droite 
(s); s { /st { = s\/st { = 0,47; s { lst 1 = 0,38; s-j/stj = 0,34; stjs^ = 0,83. 

Vésicules exsertiles: typiques du genre. 

Hypopyge: Chez le S 4n , il occupe les 8/10 de la largeur intercondylaire 
et on constate que comme pour l'acropyge, l'allongement vers l'arrière des bords 
latéraux du segment 10 portant les condyles d'articulation des cerques amène une 
déformation remarquable de l' hypopyge, semblable à celle décrite par Silvestri 
(1929) chez son dorianus; chez les autres spécimens l'hypopyge est saillant comme 
c'est la règle, occupant environ les 3/4 de la largeur intercondylaire, à sinus largement 
ouvert, à bords présentant des dents irrégulières. 

Papille génitale c? : j'ai défini dans ma note de 1975 sur Parajapyx 
(P.) botosaneanui, 4 stades du développement postembryonnaire des â caractérisés 
par le nombre de soies présentes sur la papille génitale: 6 , = 6 soies, â 1 = 12 soies, 
â 3 = 14 soies, c?4 = 16 soies. Or, parmi les 3 â reconnus ici, un a 12 soies, un autre 
14 et le troisième en présente 10, ce qui le rend intermédiaire entre mes 6 i et â 2 - Je 
crois qu'il faut donc maintenant définir 5 stades dans le développement des S et 
modifier par conséquent la nomenclature précédente. Si l'on part toujours d'un â j à 6 
soies (1+1 latérales, 1 + 1 submédianes antérieures et 1 + 1 submédianes postérieures), 
le nouveau â-, possédant 10 soies s'obtient par dédoublement des soies submédianes 
antérieures et postérieures, ce qui donne 1 + 1 soies sublatérales antérieures et 1+1 
soies sublatérales postérieures; pour le nouveau â 3 c'est le dédoublement des 1 + 1 
soies latérales qui donne bien 12 soies; quant aux 1 + 1 soies supplémentaires du â 4 , 
elles proviennent du dédoublement des 1 + 1 soies sublatérales antérieures, ce qui 
corrobore ce que j'écrivais pour botosaneanui: "les nouvelles soies apparaissent dans 
les angles latéraux de l'orifice génital, médialement par rapport aux soies latérales"; 
enfin le Ô 5 voit cette fois le dédoublement des 1 + 1 soies sublatérales postérieures. En 
résumé, la nouvelle succession des stades postembryonnaires des S de Parajapyx 
s'établit ainsi: 

d ln = 6 soies; S -> n = 10 soies; â 3n = 12 soies; â 4n = 14 soies; c? 5n = 16 soies. 
On a donc les équivalences suivantes entre la nouvelle nomenclature â xn et l'an- 
cienne â x : 

^ln =c? l' ^2n nouveau, â 3n =â 2 , <3 4n =<3 3 , <3 5n =â 4 . 



JAPYGOIDEA DU SUD-EST ASIATIQUE g87 

Papille génitale 9 : typique de la famille. En 1952a, grâce à 
l'abondant matériel en ma possession, mais ne comprenant que des individus pourvus 
de cerques indurés, j'ai défini 1 stade asexué st.I et 4 stades du développement post- 
embryonnaire des 9 , st.II à st.V caractérisés par le plus ou moins grand déve- 
loppement du canal antérieur de la spermathèque. Or en 1961, Smith découvre des 
pontes de P. (P.) isabellae et peut ainsi décrire les 2 premiers stades suivant 
l'éclosion. Il faut donc considérer que chez les Parajapygidés et les Japygidés existent 
3 stades asexués dont le troisième possède des cerques indurés; il s'ensuit que logi- 
quement, mon st.I doit être dénommé st. III par analogie avec celui des Japygidés; 
l'absence de trichobothries antennaires ne permet pas de le subdiviser en A ou B. La 
succession des stades chez les 9 doit être lui aussi modifié pour 2 raisons: tout 
d'abord, la 9 de 1,55 mm ne possède qu'une seule et unique soie sur chacune des 
valves supérieures, aucun autre phanère sur la papille et la spermathèque est indi- 
cernable; vu la taille de l'individu, je ne crois pas qu'il s'agisse d'une régression tem- 
poraire, mais du premier stade reconnaissable du développement postembryonnaire 
des 9 ; je le nomme st.I. La deuxième raison est l'extraordinaire développement de la 
spermathèque des 9 de P. (G.) reniformis n. sp. décrite plus loin (p. 892). En con- 
clusion, je propose la succession suivante des stades 9 (avec entre parenthèses l'an- 
cienne notation): st.I nouveau, st.2 (= st.II), st.3 (= st.III), st.4 (= st.IV), st.5 (= st.V) 
et st. 6 nouveau. 

Cerques 

Aussi longs en moyenne que les 4/5 de la partie normalement découverte du 
tergite 10, L /L 10d = 0,8 (v. ex. = 0,77-0,82); environ 2 fois 1/5 aussi longs que 
larges à la base; L/l = 2,14 (v. ex. = 2,05-2,27); leur largeur au niveau de la <i 3 égale 
en moyenne le 1/3 de leur longueur, le ld 3 /lcq = 0,58 chez le S holotype alors qu'il 
est en moyenne égal à 0,73 (v. ex. = 0,70-0,76) chez les paratypes. Un sinus peu 
profond entre d^ et d^ chez les paratypes, plus net chez l'holotype. 

Marges internes: celles du c? ?n et des 9 sont semblables; toutes les 
dents sont aiguës, à sommet dirigé vers la base du cerque; d 3 la plus forte, les autres 
plus petites subégales; d^ avec un très faible épaulement postérieur, obsolète chez le 
ó* 2n ; seule la d 3 montre un épaulement postérieur net; en faisant l'intervalle d^-d-, égal 
à 100 ceux entre les dents et entre û? 5 -apex du cerque sont en moyenne comme 100- 
96-62-59-170; il faut noter qu'il n'y a jamais une symétrie parfaite entre les 2 cerques 
d'un même individu, mais les écarts sont minimes; les plus grands se rencontrent chez 
le 6 2n : 100-105-63-63-168 au cerque droit, 100-90-60-60-173 au gauche. Les marges 
internes des 2 autres 6 sont bien différentes par la forme de la d x d'une part, et 
d'autre part, par le déplacement des d^ à d 5 les unes vers les autres et vers l'apex du 
cerque: chez le c? 3n la d x devient prépondérante, son sommet est saillant et arrondi, la 
d 2 s'aplatit et ne forme plus qu'une légère élevure mousse, peut saillante sur le sinus, 
les d 3 à d 5 sont identiques à celles des 9 , et les intervalles sont comme 100-93-37-33- 
96 au cerque droit et comme 100-81-26-29-90 au gauche. 

Chez le â 4n il y a une déformation remarquable des deux d^ qui s'allongent 
vers l'apex du cerque, le sommet reste arrondi; elles ont le même aspect que celle du 



888 JEAN PAGES 

seul cerque gauche de P. (P.) dorianus Silv.; les d 2 sont réduites à de minuscules tuber- 
cules arrondis, les d^ à d 5 sont assez semblables à celles des 9 , mais à sommet moins 
aigu; les intervalles sont comme: 100-74-23-23-54 au cerque droit et comme 100-72- 
21-23-62 au gauche, on voit bien le tassement vers l'apex des cerques des d^kd^ 

Plaques d'évaporation: les 9 et le â 2n ne montrent qu'une seule 
plaque d'évaporation assez grande située sur une ligne joignant M 3 à c et près de cette 
dernière; le â^ n en possède 1 à gauche et 2, plus ou moins jointives, à droite, quant au 
ó* 4n il en présente 3 à droite et 4 à gauche assez petites; chez ces 2 8 les plaques 
tendent à se déplacer d'une part vers la minuscule soie insérée au niveau de d 2 et, 
d'autre part, vers la marge interne du cerque. La présence d'une seule plaque d'éva- 
poration ne peut donc plus être considérée, à elle seule, comme indiquant un stade 
asexué; il faut tenir compte en outre de la présence d'une seule soie sur les préscutum 
des tergites 2 à 7 (Pages 1952a). 

C h é t o t a x i e : les 10 M typiques présents, M 6 courts, les autres, longs; 7 s 
courtes, les soies typiques d et e étant nulles. 

Affinités 

L'évolution de la marge interne des cerques des S rapprocherait cette espèce, 
en premier lieu de P. (P.) dorianus Silv. par les modifications de la d\, mais aussi le 
bonetianus Silv. (Silvestri 1948b), kocheri Pgs (Pages 1954), botosaneanui Pgs 
(Pages 1975) et genavensium Pgs (Pages 1978) par le rejet vers l'apex des cerques 
des û?3 à d 5 . Le nombre d'articles antennaires, l'absence de M ou de s sur certains 
tergites, en particulier le dixième et, sur les cerques, la présence d'une seule plaque 
d'évaporation chez les 9 et les S jeunes caractérisent cette espèce. 

Derivatio nominis 

Ce m'est un réel plaisir que de dédier cette remarquable espèce au Dr B. 
Hauser, Conservateur du Département des Arthropodes et d'Entomologie I au Mu- 
séum d'histoire naturelle de Genève. Grâce à ses nombreuses missions vouées avant 
tout à la récolte de la faune du sol de régions peu prospectées ou écologiquement sen- 
sibles, il a réuni, parmi d'autres, une très importante collection de Diploures dont 
d'innombrables Japygoidea, mes préférés. Invité il y a plus de vingt ans à venir étu- 
dier ce "trésor", j'ai pu apprécier la compétence, la grande gentillesse et la disponi- 
bilité totale de ce Collègue et ami. 

Parajapyx (Grassjapyx) temburong n. sp. Figs 19-26 

Matériel étudié: Bru-88/38: Brunei (Temburong District): "Peradayan Forest 
Reserve" (= "Bukit Patoi"), à 14,5 km de Bangar (= 2,5 km de Labu), forêt primaire ("Mixed 
dipterocarp forest"), prélèvement de sol dans les angles formés par les contreforts de grands 
arbres morts, 80 m; 24.XI.1988; leg. B. Hauser (B 3 ); holotype: 9 st. 4 de 2,35 mm. 

TÊTE 

Vertex: les 11 + 11 soies typiques assez courtes, aucune soies supplé- 
mentaires. 



JAPYGOIDEA DU SUD-EST ASIATIQUE 



889 




Figs 19-26 

Parajapyx (Grassjapyx) temburong n. sp., 9 st. 4 holotype. - 19. Vertex, e = 126 um. - 20. 
Mésonotum, e = 126 um. - 21. Tergite 1, e = 126 um. - 22. Tergites 7 à 10 et les cerques, e = 
126 um. - 23. Marges internes des cerques, e = 63 um. - 24. Urosternite 1, e = 105 um. - 25. 
Urosternite 3, e = 105 um. - 26. Hypopyge, e = 84 um. 

Pli oral: typique. 

Antennes: de 18 articles; chétotaxie typique du genre; 4 sensilles 
placoïdes sur l'article apical. 

Pièces buccales: typiques du genre. 



890 jean pages 

Thorax 
Face tergale 

P r o n o t u m : les 5+5 M typiques; 2+2 s, S\ nulles. 

Mésonotum: Préscutum: 1 + 1 M longs et 5-6+5-6 microsoies. Scutum: 
les 5+5 M typiques, M 2 et M 5 assez courts, les autres longs; 5+5 s typiques assez 
courtes; 2+2 soies supplémentaires dont 1+1 entre s^ et s 9 et 1 + 1 entre s 4 et s 5 . 

M é t a n o t u m : Préscutum: 2+2 M. Scutum: en tout point identique à celui 
du mésonotum. 

Face sternale 

C h é t o t a x i e : typique, identique à celle de P. (P.) isabellae, sauf pour le 
préscutum du métasternum qui ne porte que 4+4 soies, la médiane étant absente. 

Pattes: peu pileuses; 2+2 soies sternales aux tarses dont les subapicales 
sont courbées et aiguës dépassant la griffe postérieure; celle-ci 1 fois 1/3 aussi longue 
que T antérieure; unguiculus normal. 

Abdomen 

T e r g i t e 1 : Préscutum: 2+2 M assez longs. Scutum: les 5+5 M typiques, 
M 3 assez longs, les autres longs; les 6+6 s typiques, les 3+3 postérieures assez 
longues, les 3+3 médianes courtes; pas de soies supplémentaires. 

Tergites 2à7: Préscutum: les 1 + 1 M typiques assez courts. Scutum: les 
8+8 M typiques, Afj, M 3 et M 4 longs, les autres assez longs; s^ et 5 5 . 7 en paires 
typiques, s-, et s 3 présentes d'un seul côté de la ligne médiane, en général l'une d'elles 
à droite et l'autre à gauche; 1 + 1 soies supplémentaires entre s 6 et s 7 . 

Tergite 8: un peu moins de 1 fois 1/5 aussi large que long (1/L = 1,15); 
6+6 M longs (M 5 nuls); 4+4 s typiques (s nulles), s 3 aussi longues que les M, les 3+3 
autres courtes. 

Tergite 9:2 fois 1/5 aussi large que long; les 3+3 M typiques longs ou 
assez longs; ni s, ni soies supplémentaires. 

Tergite 1 : 1 fois 3/4 aussi long que large, à côtés parallèles; 6+6 M (Mj 
nuls), M 4 courts, les autres longs ou assez longs; 3+3 s fs 3 et s 4 nulles), les remarques 
faites chez l'espèce précédente à propos du choix à faire entre M 4 et s 3 s'appliquent 
parfaitement ici. 

Longueurs relatives des segments 8 à 10: 58-29-100. 

Acropyge: en triangle isocèle très allongé, 1 dent sur le côté gauche. 

S t e r n i t e 1 : Préscutum: 4+1+4 soies assez longues. Scutum: les 10+10 M 
typiques. M 3 et M 10 plus longs que les autres qui sont assez longs; 1+2 soies typiques 
assez courtes (c nulles); 1+1 soies supplémentaires entre M 5 et M 6 ; 7+7 soies assez 
courtes, unisériées en avant des organes subcoxaux latéraux. 

Organes subcoxaux latéraux: peu saillants, ils occupent environ 
le 1/3 de la largeur interstylaire; 16 soies glandulaires à droite, 15 à gauche, uni- 
sériées, toutes de même longueur, SG/st l = 0,77; 5 soies sensorielles régulièrement 
espacées, SS/st l = 0,46; SG/SS = 1,65. 



JAPYGOIDEA DU SUD-EST ASIATIQUE 891 

La partie médiane postérieure du sterilite porte les 1 + 1 minuscules soies 
typiques. 

Sternites 2 à 7 : Préscutum: 5+1+5 soies typiques longues, les 1+1 les 
plus externes les plus longues. Scutum: les 12+12 M typiques, M 3 et M 8 longs, les 
autres assez longs ou assez courts; 4+3+4 s typiques assez courtes ou courtes; une 
soie supplémentaire peut s'observer entre M 5 et M 6 . 

Styles: typiques du genre, le cône secondaire, aigu, aussi long que le 1/4 
ou le 1/3 du cône principal qui est plutôt étroit; la soie recourbée (s') des st^ est à 
peine plus courte que la soie droite (s): s^st^ = 0,38; s\/st l = 0,35; s l ls 1 = 1,3; st^/stj 
= 0,90; s^/stj = 0,34. 

Vésicules exsertiles: typiques du genre. 

Hypopyge: il occupe les 3/4 de la largeur intercondylaire, saillant, à sinus 
large et peu profond, orné de nombreuses dents aiguës. 

Papille génitale: typique d'une 9 st.4. 

Cerques 

Allongés, aussi longs que les 7/10 de la partie normalement découverte du 
tergite 10; un peu plus de 2 fois 1/3 aussi longs que larges à la base; leur largeur au 
niveau de la J 3 égale le 1/3 de la longueur d'un cerque et les 3/4 de sa largeur à la 
base. 

Marges internes: dents aiguës à sommet dirigé vers la base du cerque; 
dy et d 3 les plus fortes, les 3 autres subégales, petites; épaulements nuls; les intervalles 
entre les d et entre la d 5 et l'apex du cerque sont assez différentes d'un cerque à 
l'autre, à droite ils sont comme 100-120-78-53-240 et à gauche comme 100-83-67-61- 
211; cela peut indiquer que les d s'engrènent bien les unes dans les autres quand les 
cerques se referment; les fortes valeurs des intervalles d 5 -apex des cerques montrent 
que le crochet terminal du cerque est bien dégagé de la rangée des dents. 

Plaques d'évaporation: 2 petites plaques par cerques situées à 
l'intersection des lignes joignant M x à M 5 et M 4 à c. 

Chétotaxie: leslOM typiques, M 6 courts, les autres longs; 7 s typiques, 
d et e étant nulles, apparemment comme chez toutes les espèces de ces régions. 

Affinités 

Cette espèce est surtout caractérisée par ses organes subcoxaux latéraux et ses 
cerques, ainsi que par quelques détails de la chétotaxie de l'abdomen. Proche de 
P. (G.) sepilok Pages, les marges internes des cerques permettent de les différencier 
facilement. 

Derivatio nominis 

Le District de Temburong est situé dans la partie N de l'Etat de Brunei, 
séparée de la partie S, la plus importante, par un étroit territoire appartenant au Sabah. 



892 JEAN PAGES 

Parajapyx (Grassjapyx) reniformis n. sp. Figs 27-37 

Matériel étudié: Sar-87/37: Indonésie: Bali: Ubud, "Monkey Forest", dans le 
virage de la route traversant la forêt, sous des pierres de la pente, 200 m; 30. XI. 1987; leg. B. 
Hauser; holotype: 9 st. 6 de 2,22 mm. Sar-87/8: Indonésie: Java: Bogor, Jardin Botanique, 
prélèvement de sol dans les angles formés par les contreforts de grands arbres près du "Guest 
House", env. 250 m; 24.XI.1987; leg. B. Hauser (B,); paratype: î st.6. de 2, 20 mm. 

TÊTE 

Vertex et pli oral: chétotaxie typique. 

Antennes: typiques de 18 articles peu pileux, sans aires pileuses 
différenciées; 4 sensilles placoïdes en position typique sur l'article apical. 

Pièces bucales: typiques du genre. 

Thorax 
Face tergale 

Pronotum: 5+5 M typiques, M et M 5 assez courts, les autres longs; 3+3 s 
typiques courtes, s l à mi-distance de M 3 et M 4 , s~> et s$ rapprochées l'une de l'autre, 
seules les s 3 sont à leur place normale. 

Mésonotum: Préscutum: 1 + 1 M assez courts et 3+3 microsoies. Scutum: 
les 5+5 M typiques, M 3 les plus longs de tous, M 5 les plus courts, les autres subégaux, 
longs; les 5+5 5 typiques assez courtes; 1 + 1 soies supplémentaires entre les s^ et s 2 , 
celle de droite nulle chez la 9 de Java qui présente une soie supplémentaire à droite 
entre M-, et s-,.. 

Métanotum: Préscutum: 2+2 M assez longs et 6+6 microsoies. Scutum: 
les 5+5 M typiques, M 3 longs, M 5 courts, les autres subégaux et assez longs; la 9 de 
Bali possède les 5+5 s typiques et 3+3 soies supplémentaires entre les Sj et s 2 , M 2 et 
s 2 , s 4 et s 5 ; chez la 9 de Java, on note que si ces 2 dernières paires de soies 
supplémentaires sont présentes et les s nulles, il y a 2+2 soies supplémentaires, 1 + 1 
entre les M 5 et la seconde paire de soies supplémentaires de la 9 de Bali et 1 + 1 entre 
les s 4 et les soies supplémentaires postérieures; tous ces phanères sont assez courts. 

Face sternale 

Chétotaxie: identique à celle de P. (P.) isabellae, sauf pour les aires 
infracoxales des sternites des méso- et métasternum qui portent chacune 4 soies dont 
1 supplémentaire entre les 2 postérieures; en outre, on observe une petie soie 
supplémentaire dans la rangée antérieure de la plage médiane des mêmes sternites. 

Pattes: peu pileuses; 2+2 soies sternales aux tarses, les subapicales sont 
aiguës, atteignant au plus le milieu des griffes qui sont typiques; unguiculus petit, 
aigu. 

Abdomen 

Tergi te 1 : Préscutum: 2+2 M assez longs. Scutum: les 5+5 M typiques 
assez longs, les M 4 un peu plus développés que les autres, M 2 les plus courts; les 6+6 
s typiques assez courtes, la s 3 droite manque chez la 9 de Java; 1+1 soies supplé- 
mentaires entre A/ 4 et M 5 et une autre paire entre s 5 et s 6 . 



JAPYGOIDEA DU SUD-EST ASIATIQUE 



893 




Figs 27-37. Parajapyx {Grassjapyx) reniformis n. sp., 9 st. 6, holotype. - 27. Vertex, e = 126 
p.m. - 28. Mésonotum, e = 126 pm. - 29. Métanotum, e = 126 p.m. - 30. Tergite 1, e = 126 p.m. - 
31. Tergites 7 à 10 et les cerques, e = 126 pm. - 32. Marges internes des cerques, e = 63 pm. - 
33. Plaque d'évaporation du cerque gauche, e = 32 pm. - 34. Urosternite 1, e = 105 pm. - 
35. Urosternite 3, e = 105 pm. - 36. Canal de la spermathèque caractéristique du st. 6, e = 
238 pm. - 37. Hypopyge, e = 51 pm. 



894 JEAN PAGES 

Tergites 2 à 7 : Préscutum: les 1 + 1+1 M typiques. Scutum: les 8+8 M 
typiques, M 6 assez courts, les autres longs ou assez longs; les 7+7 s typiques présentes, 
assez courtes, s^ et s 6 les plus longues, une des s 3 manque le plus souvent chez la 9 de 
Java; 1+1 soies supplémentaires entre M 1 et M 8 et une autre paire entre s 6 et s 7 . 

T e r g i t e 8 : à peine plus large que long chez la 9 de Java, chez celle de 
Bali 1/L - 1,20; 6+6 M longs, M 3 les plus longs, M 5 nuls; 3+3 s (s 2 et 5 5 nulles), s 3 
longues, 5 } et s 4 courtes, s^ nulles chez la 9 de Java; 1 + 1 soies supplémentaires 
courtes entre M { et M 3 et 1+1 entre M l et M 7 . 

T e r g i t e 9 : entre 1,70 et 2 fois aussi large que long; les 3+3 M typiques 
longs; seules les $j sont présentes et longues. 

T e r g i t e 1 : en moyenne 1 fois 3/4 aussi long que large; 6+1+6 M (Mj 
nuls) longs; on constate la même ambiguïté entre M 4 et 5 3 que chez P. (G.) hauseri 
n. sp.; il y a au plus 4+4 s (s 4 nulles) ou 3+3 si s 3 est nulle elle aussi. 

Longueurs relatives des segments 8 à 1 : 65-35-100 en 
moyenne. 

Acropyge: saillant, triangulaire, un peu plus long que large à la base. 

S t e r n i t e 1 : Préscutum: 5+1+5 phanères longs à assez courts. Scutum: 
8+8 M, M 9 et Mjq nuls, M 5 assez courts, les autres longs, les M 8 les plus développés; 
1+2+1 5 courtes; 1 + 1 soies supplémentaires entre M 4 et M 8 et 1 + 1 autres entre M 5 et 
b; 6-7 + 6-7 soies en avant des organes subcoxaux latéraux. 

Organes subcoxaux: chacun occupe environ le 1/4 de la largeur 
interstylaire et est relativement saillant; 15 soies glandulaires unisériées et 5 soies 
sensorielles régulièrement espacées à chaque organe chez les 2 exemplaires; SG/s/j = 
0,86, SS/tfj = 0,60, SG/SS = 1,50 en moyenne. 

La partie médiane postérieure du sternite porte les 1 + 1 minuscules soies habi- 
tuelles. 

Sternites 2 à 7: Préscutum: 5+5 soies longues, la médiane est plutôt 
courte et est dédoublée chez la 9 de Bali. Scutum: les 12+12 M typiques longs; 4+2+4 
s typiques, les c étant nulles; à noter que le sternite 7 de la 9 de Bali n'a pas de Sy 

Styles: typiques du genre; la soie recourbée des st\ _ 3 est égale à la soie 
droite. s l /st l = 0,40; s\/st l = 0,40; s-Js^ = 0,32; Sjlst-, = 0,40; st^st-j = 0,80. 

Vésicules exsertiles: typiques du genre. 

Hypopyge: Il occupe le 3/4 de la largeur intercondylaire, saillant à sinus 
médian largement ouvert et à bords ornés de tubercules plus ou moins développés, 
ceux du fond du sinus aigus. 

Papille génitale 9 : Typique du genre; chez la 9 de Bali, le canal de 
la spermathèque est extraordinairement long formant un peloton lâche logé dans 
l'urite 6. Les circonvolutions de la spermathèque sont un peu moins complexes chez 
la 9 de Java. C'est la première fois que j'observe un tel développement de ce canal 
qui caractérise à mon avis un st.6 qui est pour l'instant le stade le plus "âgé" des 9 du 
genre Parajapyx (cf. p. 887). 

Cerques 

Leur longueur est égale à un peu moins des 3/4 de la partie normalement 
découverte du tergite 10, assez élancés, ils sont 2 fois (9 de Bali) à 2 fois 1/4 (9 de 



JAPYGOIDEA DU SUD-EST ASIATIQUE 895 

Java) aussi longs que larges à la base; leur largeur au niveau de la d^ égale un peu 
moins des 2/5 de leur longueur et les 4/5 de leur largeur à la base. 

Marges internes: les dents sont aiguës à sommet dirigé vers la base: 
Jj et 6?3 les plus fortes, les 3 autres subégales; pas d'épaulements antérieurs ou 
postérieurs nets; les intervalles entre les d et l'apex du cerque sont comme 100-88- 
77-63-154 en moyenne; on constate que la d^ est plus proche de l'apex du cerque à 
droite qu'à gauche: d 5 -apex = 142 au cerque droit des 2 spécimens, alors que pour le 
cerque gauche, cet intervalle est de 163 chez l'exemplaire javanais et de 170 chez le 
balinais. 

Plaque d'évaporation: d'un type inédit; on observe une surface 
reniforme lisse; au milieu de son bord concave s'ouvre un pore glandulaire qui est le 
débouché d'un canal à bords épaissis visible par transparence; la fig. 32 montre la 
petite masse de sécrétion, coagulée par le fixateur, issue de la (ou des) cellule(s) 
glandulaire(s) associée(s) à la plaque d'évaporation du cerque droit. 

Chétotaxie: Iesl0+10M typiques, M 2 , M 5 et M 6 assez courts ou courts, 
les autres longs; 5+5 s typiques, d et e nulles; à noter que je considère le phanère situé 
dans le triangle formé par M 5 , M 7 et h comme un M 6 uniquement parce qu'il est 
inséré en dessous de la ligne joignant M 5 et i et sur celle joignant /et i, la soie e se 
plaçant typiquement au dessus de la ligne M 5 -i et plus près de M 1 que de Mg. 

Affinités 

Par l'allure générale de ses cerques, P. (G.) reniformis n. sp. rappelle P. (G.) 
sepilok du Sabah. On séparera sans difficulté ces 2 espèces par des détails de la 
chétotaxie tergale, la forme et la disposition des dents des cerques et avant tout par la 
plaque d'évaporation qui est d'un type jusqu'à présent unique chez les Parajapygidés. 

Derivatio nominis 

Allusion à la forme très particulière de l'unique plaque d'évaporation des 
cerques. 

Parajapyx (Grassjapyx) sabahnus Pages Figs 38-44 

Matériel étudié: Bru-88/70: Singapour: Sentosa Island (partie orientale), forêt 
près de la "Earth Satellite Station", prélèvement de sol dans les angles formés par les 
contreforts d'un grand arbre, 70 m; 6. XII. 1988; leg. B. Hauser (B 4 ); 3 9 st.3 de 2,17 mm, 
2,34 mm et 2,54 mm. 

Ces exemplaires correspondent parfaitement à la description de cette espèce du 
Sabah (Pages 1987). La chétotaxie tergale, les rapports 1/L et L/l des segments 
abdominaux et des cerques entrent dans les limites de variations de l'espèce; les 
intervalles entre les dents sont en moyenne un peu plus faibles que chez les spécimens 
du Sabah: 100-90-65-63, au lieu de 100-87-70-68, mais ces nombres entrent aussi 
dans les limites de variations rencontrées au Sabah. Le fait le plus remarquable est 
que les exemplaires de Singapour ont les J 3 à d 5 et l'apex des cerques émoussés 
comme ceux de la 9 st.3 de 2,1 mm du Sabah. Comme je le laissais entendre en 1987, 



896 



JEAN PAGES 




Figs 38-44. Parajapyx {Grassjapyx) sabahnus Pages, 9 st. 3 de 2,54 mm. - 38. Tergite 1, e = 
1 12 jam. - 39. Tergites 7 à 10 et les cerques, e = 1 12 um. - 40. Marges internes des cerques, e = 
49 um. - 41. Urosternite 1, e = 1 12 um. - 42. Angle latéral de l'organe subcoxal gauche, e = 
42 um. - 43. Urosternite 3. e = 112 um. - 44. Hypopyge, e = 63 um. 



on ne peut donc pas expliquer cet aspect par une "usure" anormale des dents; le 
problème reste entier. 

Plaques d'évaporation: très petites, mais plus nettes que chez les 
exemplaires du Sabah; au nombre de 4-5 par cerque, elles sont inscrites dans un 
trapèze dont les sommets sont les Mj, M 3 , M 5 et la soie c comme au Sabah. 

C h é t o t a x i e : caractérisée par l'absence des soies d et e et la brièveté des 
M*. 



JAPYGOIDEA DU SUD-EST ASIATIQUE 



897 



Parajapyx (subgen. ?) sp.? 



Figs 45-47 



Matériel étudié: Bru-88/32: Brunei (Bêlait District): "Labi Hills Forest 
Reserve", "Teraja", à 42 km au sud de Sungai Liang (= 12 km au Sud de Labi), environs de 
"Rumah Panjang" (= Longhouse du Kampong Teraja), forêt primaire ("Mixed dipterocarp 
forest"), prélèvement de sol dans les angles formés par les contreforts d'un très grand arbre, 
40 m; 22.XI.1988; leg. B. Hauser (B 9 ); 1 9 st.3 de 1,80 mm. 




Figs 45-47 

Parajapyx (subgen. ?) sp.?, 9 st. 3. - 45. Tergites 7 à 10 et les cerques, e 
internes des cerques, e = 45 um. - 47. Hypopyge, e = 105 um. 



79 um. - 46. Marges 



g98 JEAN PAGES 

Cet exemplaire possède des cerques dont les caractéristiques ambiguës me font 
hésiter à le classer spécifiquement. 

Les cerques sont nettement plus trapus que chez les espèces précédentes et se 
rapprocheraient de ceux de P. (G.) sabahnus, mais la forme de la d 2 et la présence 
d'une seule plaque d' evaporation située sur une ligne joignant M 5 et c l'en écartent 
incontestablement. 

Ces 2 caractéristiques se retrouvent chez P. (P.) hauseri, mais si les 
chétotaxies sont semblables, l'absence d'un sinus entre d\ et dy même aussi peu 
marqué que chez la 9 st. 2 de P. hauseri, et l'aspect plus trapus des cerques rendent ce 
rapprochement difficile. 

Les intervalles entre les dents des cerques de cet exemplaire sont comme 100- 
107-73-80-213 à droite et comme 100-94-59-65-212 à gauche, valeurs voisines de 
celles rencontrées chez P. (G.) temburong, mais cette dernière espèce a des cerques 
beaucoup plus élancés et de petites plaques d' evaporation situées elles aussi, il est 
vrai, sur une ligne joignant M 5 et c, enfin les acropyges sont identiques, présentant 
une dent sur le côté gauche. Je ne crois ni utile, ni souhaitable de créer un taxon 
nouveau en l'absence d'autres exemplaires, en particulier de S provenant de la même 
station. 

REMERCIEMENTS 

Je tiens à remercier Mme M. Krähenbühl qui a dû dactylographier mon manus- 
crit et M. G. Roth qui a reproduit sur calque mes dessins originaux. 

BIBLIOGRAPHIE 

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115-119. 
Ewing, H. E. 1941. New North American genera and species of Apterygotan Insects of the 

Family Japygidae. Proceedings of the Entomological Society of Washington 43: 69-75. 
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History Research, Special Issue 1: 219. 
Pages, J. 1952a. Parajapyginae (Insecta, Entotrophi, Japygidae) de l'Angola. Publicaçôes 

Culturais da Companhia de Diamantes de Angola 13: 53-96. 
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suisse de Zoologie 94: 41-47. 
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9:51-62. 



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Zoologia Generale e Agraria della R. Scuola Superiore d'Agricoltura, Portici 22: 

49-80. 
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Superiore d'Agricoltura, Portici 23: 210-226. 
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Revue suisse de Zoologie 105 (4): 901-910; décembre 1998 



Redescription oï Diplomystes mesembrinus 
(Siluriformes, Diplomystidae) 

Maria de las Mercedes AZPELICUETA 1 & Atila Esteban GOSZTONYI 2 

1 Depto. Cientifico Zoologia de Vertebrados, Museo de La Plata, 1900 La Plata, 
Argentina 

2 Centro Nacional Patagónico, Conicet, Bvd. Brown s/n, 9120 Puerto Madryn, 
Argentina. 

Redescription of Diplomystes mesembrinus (Siluriformes, Diplomysti- 
dae). - New specimens of Diplomystes mesembrinus collected in the rio 
Chubut, Argentina, allow a thorough description of the species, hitherto 
known only by four specimens. Diplomystes mesembrinus may be distingu- 
ished from D. viedmensis and D. cuyanus by the possession of a low number 
of maxillary teeth, no more than 23 in adults. This character is shared by 
D. mesembrinus and the Chilean species D. chilensis, D. camposensis, and 
D. nahuelbutaensis. The suture between basibranchials 2 and 3 differentiates 
D. mesembrinus from the three Chilean species and D. viedmensis. Some 
ontogenetic changes in D. mesembrinus are commented upon. 

Key-words: Siluriformes - Diplomystidae - Diplomystes mesembrinus - 
Neotropical Region - Patagonia. 

INTRODUCTION 

Patagonia, in southern South America, is an extense region with scarce rivers. 
Most of the Patagonian rivers have headwaters in the Cordillera de los Andes, many of 
them at high altitude, and run from the West to the East, pouring into the Atlantic 
Ocean. In the rivers of Patagonia, fresh water fishes are scarce. Only species of the 
siluriform families Diplomystidae and Trichomycteridae are recorded within the 
ichthyofauna of the area. Six species are included into the family Diplomystidae; three 
of them are present in Chile and other three species in Argentina. The three East- 
Andean species Diplomystes cuyanus, D. viedmensis and D. mesembrinus respectively 
occur in the Colorado, Negro, and Chubut-Senguerr basins. Chubut-Senguerr basin is 
one of the most important Patagonian basins; it mainly occurs in the province of Chubut 
between 41° 30 and 46° S. From the mouth of the rio Senguerr, Ringuelet (1982) 
described Diplomystes mesembrinus based on two specimens. Although several records 
of the "bagre pintado" appeared in a chronicle of an expedition to the rio Chubut in 
1865-66 (Claraz 1988), only material type and two very small specimens of 
D. mesembrinus, captured by one of the authors in that river (Gosztonyi 1988), were 
known until five years ago, and the adults of D. mesembrinus were unknown. 



Manuscript accepted 08.05.1998 



902 AZPELICUETA & GOSZTONYI 

The objective of this paper is to redescribe D. mesembrinus using external and 
osteological characters. Remarks about disgnostic characters and ontogenetic changes 
of the species are added. 

MATERIAL AND METHODS 

Specimens of D. mesembrinus were cleared and counterstained following 
Taylor & Van Dyke (1985). Measurements are straight distances taken with calliper 
to nearest 0. 1 mm, on the left side of the body whenever possible. 

Cleared and stained specimens (C&S) of Diplomystes viedmensis Mac Donagh, 
1931, D. cuyanus Ringuelet, 1965, D. chilensis Molina, 1782, D. camposensis Arratia, 
1987, and D. nahuelbutaensis Arratia, 1987, have been examined. 

Material is deposited in Centro Nacional Patagónico (CENPAT), Muséum 
d'histoire naturelle de Genève (MHNG) and Museo de La Plata (MLP). 

REDESCRIPTION 

Diplomystes mesembrinus Ringuelet, 1982 Figs 1-8, table 1 

Diplomystes viedmensis mesembrinus Ringuelet, 1982: 350. 
Diplomystes mesembrinus; Azpelicueta 1994a: 13. 

Material examined. 66 specimens, provincia del Chubut, Argentina. Holotype: MLP 8452, 
168.0 mm, mouth of rio Senguerr. MLP uncat., 1 ex., 33.8 mm, rio Chubut inferior (C&S), col. 
A. Gosztonyi. MLP uncat.. 9 ex., 80.4-170.7 mm (1 ex., C&S), rio Chubut near Los Altares (43° 
51' 30" S-68° 28' W), 18-12-95. MLP uncat., 3 ex., 65.0-101.0 mm (2 ex. C&S), rio Chubut 
near km 261 provincial road 25, 19-12-95, col. A. Gosztonyi and R. Laylor. MLP uncat., 20 ex., 
63.0-135.3 mm (2 ex. C&S), rio Chubut en canadon Carbon /43° 49' 8" S- 67° 53' 8"), 18-4-97, 
col. A. Gosztonyi, L. Kuba, and R. Taylor. MLP uncat., 1 ex., 103.2 mm, rio Chubut en Los 
Altares, same date and collectors. MLP uncat., 8 ex., 99.0-159.5 mm, rio Chubut, near cerro 
Condor (43° 19' I"-69° 8' 2"), 19-4-97, same collectors. CENPAT uncat., 2 ex., 137.0-187.0 
mm (C&S), rio Chubut near Los Altares, 9-12-1993, col. R. Taylor. CENPAT uncat., 14 ex., 
82.2-215.0 mm, same locality, 18-12-95, col. A. Gosztonyi and R. Taylor. CENPAT uncat., 5 ex., 
69.0-122.0 mm, km 261 of the provincial road 25, 19-12-95, same collectors. MHNG 2593.66- 
67, 2 ex., 127.0-134.0 mm, rio Chubut. near cerro Condor, 19-4-97, col. A. Gosztonyi, L. Kuba, 
and R. Taylor. 

Diagnosis. Diplomystes mesembrinus is differentiated from D. cuyanus and 
D. viedmensis by the presence of few maxillary teeth, no more than 23 in adults, a 
character of uncertain polarity shared with the Chilean D. chilensis, D. cuyanus, and 
D. nahuelbutaensis. The suture between basilbranchials 2 and 3 distinguishes D. mes- 
embrinus from the three Chilean species and D. viedmensis. Diplomystes mesembrinus 
has the narrowest mouth (0.26-0.32 vs. 0.34-0.42 in head length of the other species). 

Redescription. The previously known descriptions of D. mesembrinus were 
based on the holotype (Ringuelet 1982; Azpelicueta 1994b). Completing those 
descriptions, only the differences found in the new material are mentioned below. 
Measurements are listed in table 1, as percentages of different lengths. 



REDESCRIPTION OF DIPLOMYSTES MESEMBRINUS 903 

Dorsal fin with first spine small, second spine well developed and 7 rays 
branched, as in all diplomystid species. Second spine with 6-8 small serrae in half or 
distal one third of posterior margin, serrae worn in adults; anterior margin of second 
spine smooth. Adipose-fin origin placed at same level of anal-fin origin or scarcely 
anteriorly (Fig. 1). Adipose depth in percentage of its length 16.67-30.0. 

Pectoral fin with one spine and 8-10 branched rays; pectoral-fin tip extended 
beyond pelvic-fin origin in juveniles (Fig. 2), not reaching such origin in adults. 
Pectoral spine with 18 posterior serrae in adults, highest number found in diplomystid 
species. 

Two pectoral axillary gland pores present; posterior one usually divided into two 
apertures. 

Pelvic-fin origin always, placed in anterior half of standard length. Pelvic-fin tip 
extended beyond origin of anal fin in youngs and some juveniles, but not in adults. 

Anal fin with 4-5 unbranched rays and 8-9 branched rays. Last simple ray or 
first branched ray longer. Dorsal and anal fins with fleshy bases. 

Lower caudal lobe scarcely longer and wider than upper one. Lateral line end 
usually curved dorsally, sometimes straight (five specimens). Dorsal procurrent caudal 
rays 16-17 and ventral procurrent rays 16-18. 

Dorsal head surface on supraoccipital smooth. Fleshy fold around posterior nare 
completely or partially covering the aperture. Mouth narrow; premaxillary tooth plate 
wide in relation to mouth width. Maxillary barbel reaching beyond pectoral-fin origin 
in small specimens, and near branchiostegal membrane in adults. 

Total number of vertebrae 42-44, including the Weberian apparatus and preural 
centrum 1+ural centrum 1. Anterior gill rakers on first arch as follows: 6-8 in epi- 
branchial, 1 in cartilage, 12-14 in ceratobranchial and hipobranchial. Pseudobranch 
bearing 14-15 filaments in adults. Eleven or twelve pairs of ribs present; first rib 
reduced and fused to 5th parapophysis. A well-developed rib joined to 5th. para- 
pophysis present in one specimen (140 mm SL). 

Coloration in life: Body grey, upper area of flanks dark, lower area light; body 
whitish ventrally. Small black dots on flanks, more abundant dorsally. Few dots on 
dorsal surface of head, more concentrated on snout. Small specimens grey, without dots 
(Fig. 2). 

Pectoral fins dark grey, pelvic fins light grey. Dark chromatophores only on 
dorsal surface of both paired fins. Dorsal and caudal fins with very small dots and distal 
margin dark; five specimens with caudal margin unpigmented. Larger specimens with 
dots scattered on anal fin, anal-fin base of small specimens whitish, and light grey 
distally. Adipose fin completely covered by very small dots. 

Papillae: Filamentous papillae, scarce in juveniles and numerous in adults, 
cover the entire body; also, papillae are present at the base of dorsal and pectoral fins, 
and spines. Papillae are short in juveniles and long in adults (1.5 mm). There are 
rounded papillae around mouth, posterior surface of maxilla, and branchiostegal area. 
Many of those papillae around mouth become prismatic during growth. The dorsal area 
of each papilla has eight taste-buds. Scarce rounded or filamentous papillae are always 
found in mouth roof. 



904 



AZPELICUETA & GOSZTONYI 




Fig. 1 
Diplomystes mesembrinus, rio Chubut near cerro Condor, 150.5 mm SL. 




Fig. 2 

Young specimen of Diplomystes mesembrinus, no Chubut near cerro Condor, 102.2 mm SL, 
without dots on body. 



REDESCRIPTION OF DIPLOMYSTES MESEMBRINUS 



905 



Large pit-organs in a line, parallel to lateral line, present along the entire flanks 
(Azpelicueta 1993). Also, large pit-organs occur in lines in the dorsal surface of head 
and in front of dorsal fin, on body (Azpelicueta 1994a; Arratia & Gayet 1995). 




Figs 3-6 

Diplomystes mesembrinus, 187 mm SL. 3: Right autopalatine, dorsal view, an incomplete 
fusion between the two anterior autopalatine processes; 4: Right maxilla, with conic and 
spatulate teeth, lateral view; 5: Right maxilla, ventral view, schematic illustration showing the 
arrangement of the maxillary teeth; 6: Right Suspensorium, mesial view, a small metapterygoid 
process present, metp: metapterygoid process. Scale bar: 1 mm. 



906 



AZPELICUETA & GOSZTONYI 




Figs 7-8 

Diplomystes mesembrinus, 215 mm SL, right branchial elements, dorsal view; 7: Suture 
between basibranchial 2 and 3; 8: Process in the anterior branch of the uncinate epibranchial 3 
with the ligament attached, bb 2: basibranchial 2; bb 3: basibranchial 3; epi: uncinate 
epibranchial 3; lig: ligament; pab: process in the anterior branch of the third epibranchial; phb: 
pharyngobranchial 4. Scale bar: 1 mm. 



Osteology: The sphenotic of small specimens is short and wide whereas the 
sphenotic of large specimens has a very long anterior process. The wide frontal of early 
ontogenetic stages becomes narrow posteriorly in adults and extremely narrow in very 
large specimens. One crest for muscle attachment crosses along dorsal surface of the 
extrascapula. 

The number of maxillary teeth does not exceed 23 in adults; teeth are arranged 
in three somewhat unordered rows anteriorly and two rows posteriorly (Figs 4, 5). 

Tooth plates develop under autopalatine on both sides in 18 specimens (N = 40); 
only on one side in seven specimens, and they are absent in the rest. Tooth plates have 
irregular number of teeth and different size; also, more than one tooth plates may occur 
under autopalatine, on each side. All specimens have two vomerine tooth plates. The 
autopalatine of small specimens and juveniles has two anterior processes that fuse 
during ontogeny, although they are not completely fused in some large specimens 
(Fig. 3). Ectoperygoid and entopterygoid always occur under the autopalatine posterior 
process which is long in adults. The metapterygoid process only develops in one 
specimen (Fig. 6). A large crest for insertion of levator arcus palatini is present on the 
hyomandibula. 



REDESCRIPTION OF DWLOMYSTES MESEMBR1NUS 



907 



Posterior margin of basibranchial 2 and anterior one of basibranchial 3 suture 
during ontogeny (Fig. 7); the beginnig of this trend is observed at 130 mm SL. A small 
process directed posteriorly, for attachment of a short and strong ligament that firmly 
joins the bone to pharyngobranchial 4, grows in the anterior branch of the uncinate third 
epibranchial (Fig. 8). 

In the Weberian apparatus, the reabsorption of the horizontal process of the 
intercalarium occurs in early stages of ontogeny. One supraneural develops between the 
supraoccipital and the neural arch of the complex vertebrae, not reaching the supra- 
occipital in large specimens. The dorsal margin of the claustrum does not contact the 
supraneural in the largest specimen (215 mm SL). 

There are three proximal radiais in the pectoral fin. The number of pectoral 
distal radiais is reduced from five to two during ontogeny. The pelvic bone has three 
anterior processes and one pelvic radial in all specimens; the youngest specimen 
examined has the medial process yet cartilaginous (Azpelicueta 1994b, fig. 14c). 

Table 1 

Measurements of 35 specimens of Diplomystes mesembrinus in percentage of indicated lengths. 
SL = 33.8-215.0 mm. 



Percentage of standard length 
Predorsal-fin length 
Preadipose fin-length 
Preventral-fin length 
Preanal-fin length 
Dorsal-fin base 
Adipose-fin base 
Anal-fin base 
Pelvic-fin length 
Pectoral-spine length 
Dorsal-spine length 
Greatest body depth 
Greatest head depth 
Head length 
Head width 
Mouth width 
Interorbital length 
Preorbitai length 

Percentage of head length 
Head width 
Mouth width 
Greatest head depth 
Interorbital length 
Preorbitai length 
Orbital length 
Maxillary length 
Premaxillary length 

Percentage of mouth width 
Premaxillary tooth plate 



37.8 


35.1 


39.2 


64.2 


57.8 


68.2 


48.6 


44.5 


51.0 


66.7 


59.9 


69.0 


13.7 


12.3 


16.4 


25.9 


21.4 


31.9 


14.0 


11.0 


17.3 


16.6 


13.6 


20.0 


18.4 


15.2 


21.5 


18.0 


11.8 


21.7 


20.0 


14.4 


23.3 


15.7 


13.0 


20.0 


26.6 


24.2 


29.0 


18.4 


16.2 


22.2 


7.7 


6.0 


7.7 


8.1 


6.8 


8.1 


11.2 


8.3 


13.1 


69.1 


61.6 


86.7 


20.0 


22.6 


35.9 


59.2 


50.0 


74.5 


30.5 


25.7 


35.8 


42.1 


33.3 


49.2 


16.4 


11.3 


29.8 


23.9 


17.7 


28.4 


22.7 


15.6 


27.3 



82.7 



65.8 



99.9 



908 AZPELICUETA & GOSZTONYI 

BIOLOGY 

Little is known about the biology of the family (Arratia 1983; Azpelicueta 
1994a). Examination of a few specimens shows that five stomach contents include adult 
Hymenoptera, large amount of terrestrial Coleoptera, and numerous specimens of the 
gasteropod Chilina sp. 

Males have testes with broad lobes, similar in anterior and posterior regions. As 
in other species of diplomystids, the females only have one gonad. At the beginning of 
the warm season (December), the females were not ripe. 

Large specimens have been collected in a slow, deep run on the southern side of 
a wide turn in a meandering section of Chubut river. The medium-sized specimens were 
collected in shallow waters, usually with faster current. According to the observations 
of Gozstonyi (1988) the small specimens were caught in a fast flowing narrow section 
of the river. 

DISCUSSION 

The family Diplomystidae is the only group of living catfishes retaining a 
dentate maxilla with long medial process and laminar lateral expansion. Maxillary teeth 
are placed along most of the oral surface of the bone and are arranged in somewhat 
unordered rows. A low number of maxillary teeth, not exceeding 23 in adults, have 
been considered by Arratia (1987) as a diagnostic character for the species living in 
the Western slope of the Andes. Nonetheless, the same number is present in adults of D. 
mesembrinus (Figs 4, 5). This number of maxillary teeth distinguishes D. mesembrinus 
from D. viedmensis and D. cuyanus which have a high number of teeth. At 215 mm SL, 
specimens of D. cuyanus and D. viedmensis have about 40 teeth. Such number changes 
during ontogeny and about 60 maxillary teeth, placed in five rows anteriorly, occur in 
the largest D. viedmensis (324 mm SL; Azpelicueta 1994b, fig. 16f). The teeth of 
D. mesembrinus are arranged in three unordered rows anteriorly and two rows poster- 
iorly. Diplomystes camposensis, D. nahuelbutaensis, and D. chilensis have two rows 
anteriorly and one row posteriorly (Arratia 1992), an arrangement that differentiates 
them from D. mesembrinus. 

The autopalatine of diplomystid species has two anterior processes in early 
stages of ontogeny. D. nahuelbutaensis and D. camposensis retain those processes in 
large specimens (Arratia 1987) whereas both processes fuse during grow in D. 
viedmensis, D. cuyanus, D. chilensis, and D. mesembrinus; nonetheless, some large 
specimens of D. mesembrinus have an incomplete fusion (Fig. 3). 

A metapterygoid process is found in large specimens of D. cuyanus and in 
different ontogenetic stages of D. viedmensis (Azpelicueta 1994b); such process bears 
a small ligament attached to parasphenoid. This process only occurs in one specimen of 
D. mesembrinus (Fig. 6). 

The presence of a suture between basilbranchial 2 and 3 differentiates 
D. mesembrinus from all diplomystid species, but D. cuyanus. The small process deve- 
loped in the anterior branch of the uncinate third epibranchial is a character shared by 
D. mesembrinus and D. cuyanus. 



REDESCRIPTION OF DIPLOMYSTES MESEMBRINUS 909 

Fink & Fink (1981, 1996) listed the absence of third supraneural as a Characi- 
physan character. Usually, one supraneural appears in the Weberian apparatus of all 
Diplomystes, although some specimens of D. chilensis and D. cuyanus have one 
ossified element posterior to the first ossification (Arratia 1987, fig. 9b; Azpelicueta 
1994b). Arratia (1992) mentioned one supraneural with two centers of ossification. In 
two juveniles of D. cuyanus, the suture between both elements is clearly distinguished 
but none of the adult specimens examined have two elements. The examination of 
larval and postlarval specimens of D. chilensis or D. cuyanus will confirm the origin of 
the second ossification which may represent other supraneural. The very small D. vied- 
mensis, D. nahuelbutaensis and D. mesembrinus examined have only one supraneural 
(30, 26, and 33 mm of SL respectively). 

During ontogeny, the cranium of the diplomystid species becomes narrow, 
specially at the epiphyseal bar level. The shape of some skull bones as sphenotic, 
frontal, and supraoccipital strikingly changes in large specimens of D. mesembrinus, as 
in the remaining species of Diplomystes. The presence of one crest for muscle 
attachment on the extraescapula of D. mesembrinus distinguishes this species from 
adult D. viedmensis. 

ACKNOWLEDGMENTS 

The authors thank Luisa Kuba and Roberto Taylor for their help during 
collecting field trips, Rodolfo Casamiquela for mentioning the chronicle of Claraz and 
comments about its expedition to the rio Chubut, and Gloria Arratia her comments on 
the manuscript. 

REFERENCES 

Arratia, G. 1983. Preferencia de habitat de peces siluriformes de aguas continentales de Chile 
(Fam. Diplomystidae y Trichomycteridae). Studies on Neotropical Fauna and Environ- 
ment 18: 217-237. 

Arratia, G. 1987. Description of the primitive family Diplomystidae (Siluriformes, Teleostei, 
Pisces): morphology, taxonomy and phylogenetic implications. Bonner zoologische 
Monographien 24: 1-120. 

Arratia, G. 1992. Development and variation of the Suspensorium of primitive catfishes 
(Teleostei, Ostariophysi) and their phylogenetic relationships. Bonner zoologische Mono- 
graphien 32: 1-148. 

Arratia, G. & Gayet, M. 1995. Sensory canals and related bones of Tertiary siluriform crania 
from Bolivia and North America and comparison with Recent forms. Journal of Verte- 
brate Paleontology 15: 482-505. 

Azpelicueta, M. de las M. 1993. Neuromastos superficiales en Diplomystes. Neotrópica 39: 
101-102. 

Azpelicueta, M. de las M. 1994a. Los diplomfstidos en Argentina, pp. 5-27. In: Castellanos, 

Z. A. de (ed.). Fauna de Agua Dulce de la Repüblica Argentina. PROFADU-CONICET. 

Estudio Sigma, Buenos Aires 40: 1-49. 
Azpelicueta, M. de las M. 1994b. Three East- Andean species of Diplomystes (Siluriformes, 

Diplomystidae). Ichthyological Exploration of Freshwater s 5: 223-240. 
Claraz, J. 1988. Diario de viaje de exploración al Chubut 1865-66. Ediciones Marimar, Buenos 

Aires. 191 pp. 



910 AZPELICUETA & GOSZTONYI 



Fink, S. V. & Fink, W. L. 1981. Interrelationships of ostariophysan fishes. Zoological Journal of 

the Linnean Society 72: 297-353. 
Fink, S. V. & Fink, W. L. 1996. Interrelationships of ostariophysan fishes (Teleostei), pp. 209- 

249. In: Stiassny, M. L., Parenti, L. R. & Johnson, G. D. (eds). Interrelationships of 

fishes. Academic Press, San Diego, xiii + 496 pp. 
Gostonyi, A. E. 1988. Peces del rio Chubut inferior, Argentina. Physis (Buenos Aires), Sección B 

46:41-50. 
Ringuelet, R. A. 1982. Una nueva subespecie de bagre patagónico Diplomystes viedmensis Mac 

Donagh, 1931 en el no Senguer (Chubut, Argentina). Limnobios 2: 349-351. 
Taylor, W. R. & Van Dyke, G. C. 1985. Revised procedures for staining and clearing small 

fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-1 19. 



Revue suisse de Zoologie, 105 (4): 91 1-982; décembre 1998 



Aleocharinae della Cina: Parte IV 
(Coleoptera, Staphylinidae) 



Roberto PACE 

Via Vittorio Veneto 13; 1-37032 Monteforte d'Alpone (Verona), Italia. 

Aleocharinae from China: Part IV (Coleoptera, Staphylinidae). - In 

this paper further 69 species are described as new to science. These new 
species belong to following tribes: Athetini (part III) (42 species), 
Thamiaraeini (4), Pygostenini (6) and Myrmedoniini (17). One subgenus 
and two genera are described as new, assigned to following tribes: 
Pygostenini {Mesomegaskela n. gen. and Cephaplakoxena n. gen.) and 
Thamiaraeini (Aidemonusa n. subgen. of M imoxypoda). The main diagnos- 
tic characters are illustrated. 

Key-words: Coleoptera - Staphylinidae - Aleocharinae - Taxonomy - 
China. 



INTRODUZIONE 

Nel presente lavoro viene esaurita la descrizione di nuove specie dell'este- 
sissima tribù Athetini ed è compresa la descrizione di nuove specie delle tribù 
Thamiaraeini, Pygostenini e Myrmedoniini. Anche queste descrizioni, come quelle dei 
tre precedenti lavori di questa serie (Pace 1998a, b, e), sono fatte su materiali di recente 
raccolta da parte dei colleghi studiosi di Staphylinidae Guillaume de Rougemont e 
Dr Ales Smetana di Ottawa. Ho incluso in questo lavoro due specie nuove che non 
appartengono propriamente al territorio cinese: una del Kazachistan, l'altra della 
Siberia. 

Gli holotypi contrassegnati con la sigla (MHNG) sono conservati nelle 
collezioni del Museo di Storia naturale di Ginevra. Un holotypus contrassegnato con la 
sigla (CASS) è conservato in collezione Volker Assing di Hannover. 

ATHETINI (parte III) 

Atheta (Phanerosphaena) retroarmata sp. n. Figg. 1-3 

Holotypus ó\ Hong Kong, Kadoorie Agricultural Research centre, flight interception 
trap, 19-31.V.1996, de Rougemont leg. (MHNG). 



(142° Contributo alla conoscenza delle Aleocharinae) 
Manoscritto accettato il 14.02.1998. 



912 ROBERTO PACE 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e bruno con estremità addo- 
minale giallo-rossiccia; Antenne brune con i due antennomeri basali e l'apice dell'un- 
dicesimo giallo-rossicci; zampe giallo-rossicce. La reticolazione della superficie del 
capo è estremamente superficiale, quella sul resto del corpo è assente. I tubercoletti 
che coprono la superficie dell'avancorpo sono svaniti, quelli dell'addome sono 
salienti e fini. Edeago figg. 2-3. 

Comparazioni. La nuova specie è chiaramente distinta da A. tronqueti Pace, 
1987a del Nepal e della Thailandia, sia per i caratteri esterni che per l' edeago. Gli 
enormi occhi della nuova specie contrastano con quelli ridotti di tronqueti. Inoltre 
l' edeago della nuova specie è strettamente e profondamente infossato al livello della 
"crista apicalis" e l'armatura genitale interna è costituita da due lamine falciformi. 
Questi caratteri, insieme a molti altri non elencati, non si riscontrano nell'edeago di 
tronqueti. 

Atheta (Microdota) iperintroflexa sp. n. Figg- 4-5 

Holotypus 9, China, Zhejiang, Tianmushan, 29.IV. 1993, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 2,0 mm. Corpo lucido e bruno con capo e uriti liberi 4° 
e 5° nero-bruni e con elitre giallo-brune; antenne brune con antennomero basale 
rossiccio e il successivo rossiccio; zampe gialle. La reticolazione del capo è super- 
ficiale, quella del pronoto, delle elitre e dei tre uroterghi basali è estremamente svanita, 
quella del quarto urotergo libero è ben trasversa come quella degli uroterghi anteriori 
ma è netta, quella del quinto e sesto urotergo libero è quasi vigorosa e a maglie 
isodiametriche. I tubercoletti della superficie del capo sono fittissimi e poco salienti, 
quelli del resto della superficie del corpo sono fini e salienti. Spermateca fig. 5. 

Comparazioni. In base alla struttura della spermateca, la nuova specie è 
comparabile con A. pseudocoprophila Cameron, 1950, di Selangor, ma la nuova 
specie ha taglia corporea maggiore (2,1 mm invece di 1,6 mm), il quarto antennomero 
più lungo che largo (e non trasverso come in pseudocoprophila) e per la spermateca 
che ha una profondissima introflessione apicale del bulbo distale (breve in pseudo- 
coprophila). 

Atheta (Microdota) tricoloroides sp. n. Figg- 6-10 

Holotypus 6 . China, Zhejiang, Hangzhou, 27.IV. 1993, de Rougemont leg. (MHNG). 
Paratypi; 8 es., stessa provenienza. 

Descrizione. Lunghezza 2,0 mm. Corpo lucido. Capo e uriti liberi 3° e 4° 
bruni, pronoto, i due uriti basali ed estremità addominale giallo-rossicci, elitre giallo- 
brune; antenne brune con i due antennomeri basali giallo-rossicci; zampe gialle. La 
reticolazione della superficie del capo e dell'addome è distinta, quella del pronoto è 
netta e quella delle elitre è svanita. Le maglie di reticolazione della superficie 
dell'addome sono poligonali irregolari. I tubercoletti della superficie dell'avancorpo 
sono superficiali, quelli dell'addome sono salienti. Edeago figg. 7-8, sesto urotergo 
libero del maschio fig. 9, spermateca fig. 10. 



ALEOCHARINAE DELLA CINA 



913 




FlGG. 1-6 

Habitus, edeago in visione laterale e ventrale e spermateca. 1-3: Atheta (Phanerosphaena) 
retroarmata sp. n.; 4-5: Atheta (Microdota) iperintroflexa sp. n.; 6: Atheta (Microdota) trico- 
loroides sp. n. 



914 ROBERTO PACE 

Comparazioni. La nuova specie è simile ad A. masuriensis Cameron, 1939, 
dell'India. Il solo carattere distintivo somatico esterno più evidente è la maggiore 
lunghezza dell'undicesimo antennomero della nuova specie, rispetto quello di masu- 
riensis. La spermateca ha forma e grandezza simili nelle due specie, tuttavia l'intro- 
flessione apicale del bulbo distale della stessa spermateca è minuscula nella nuova 
specie e profonda in masuriensis. E' nell'edeago che si notano le maggiori differenze 
morfologiche tra le due specie. Tra le molte è da segnalare che l'armatura genitale 
falciforme interna dell'edeago della nuova specie, non si riscontra nell'edeago di 
masuriensis. 

Atheta (Microdota) jiensis sp. n. Figg. 11-14 

Holotypus S. China, Beijing. Xiaolongmen. 1100-1500 m, 1. VII. 1993, de Rougemont 
leg. (MHNG). 

Paratypi: 9 es., stessa provenienza. 

Descrizione. Lunghezza 1,9 mm. Corpo lucido e bruno-rossiccio con capo 
bruno; antenne nere con antennomero basale rossiccio; zampe gialle. La reticolazione 
del disco del capo è netta, quella sul resto della superficie del capo e sulle elitre è 
svanita, quella del pronoto è quasi virorosa, quella dell'addome è distinta e a maglie 
poligonali irregolari. I tubercoletti della superficie del capo e del pronoto sono poco 
salienti e assenti sulla fascia mediana del capo, quelli della superficie delle elitre sono 
distinti. Edeago figg. 12-13, spermateca fig. 14. 

Comparazioni. La nuova specie è simile ad A. placita Cameron, 1939, 
dell'India, ma il colore del corpo è differente (corpo bruno-rossiccio con capo nero 
invece di corpo giallo-rossiccio con capo, elitre e fascia addominale bruni come in 
placita) e le elitre di placita hanno maggiore sviluppo in lunghezza e larghezza. Il 
bulbo basale dell'edeago della nuova specie è più sviluppato di quello di placita e 
l'armatura genitale interna dello stesso edeago della nuova specie è nettamente più 
robusta di quella di placita. La spermateca della nuova specie ha sviluppo maggiore 
di quello della spermateca di placita. 

Etimologia. La nuova specie prende nome dall'antico nome "Ji" di Pekino, 
risalente all'ottavo secolo a.C. 

Atheta (Microdota) yanensis sp. n. Figg- 15-18 

Holotypus ó\ China, Beijing, Xiaolongmen, 1100-1500 m, 1. VII. 1993, de Rougemont 
leg. (MHNG). 

Paratypi: 8 es., stessa provenienza. 

Descrizione. Lunghezza 1,6 mm. Corpo lucido e bruno; antenne brune con 
antennomero basale bruno-rossiccio; zampe gialle con femori bruni. La reticolazione 
sul disco del capo è netta, sul resto della superficie del capo e sul resto del corpo 
superficiale. I tubercoletti della superficie del capo, del pronoto e delle elitre sono 
salienti: sono assenti sulla fascia mediana sia del capo che del pronoto. I tubercoletti 
dell'addome sono svaniti. Edeago figg. 15-17, spermateca fig. 18. 



ALEOCHARINAE DELLA CINA 



915 




FlGG. 7-14 

Edeago in visione laterale e ventrale, sesto urotergo libero del maschio, spermateca e habitus. 
7-10: Atheta (Microdota) tricoloroides sp. n.; 1 1-14: Atheta (Microdota) jiensis sp. n. 



916 ROBERTO PACE 

Comparazioni. La nuova specie è affine ad A. sororcula Cameron, 1939, 
dell'India, a motivo della forma dell'edeago e della spermateca. Ha elitre appena più 
larghe del pronoto e non nettamente più larghe del pronoto come in sororcula. 
L'edeago della nuova specie ha minore sviluppo, la sua parte apicale, in visione 
laterale, meno ampia di quella dell'edeago di sororcula che ha struttura dell'armatura 
genitale interna dell'edeago più robusta e più ricca di pezzi copulatoli rispetto quella 
dell'edeago della nuova specie. Il bulbo distale della spermateca della nuova specie ha 
maggiore sviluppo della parte restante della stessa spermateca e ha profonda e robusta 
introflessione apicale, mentre in sororcula il bulbo distale, con introflessione apicale 
breve, ha minore sviluppo rispetto al resto della stessa spermateca. 

Etimologia. La nuova specie prende nome dallo Stato feudale Yan dell' ot- 
tavo-quinto secolo a.C. della regione di Pekino dove è stata raccolta. 

Atheta (Microdota) gonggaensis sp. n. Figg. 19-21 

Holotypus S, China, Sichuan, Gongga Shan, above camp 2, 2800 m. 25. VII. 1994, 
A. Smetana leg. (MHNG). 

Paratypi: 2 S , stessa provenienza. 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e bruno con capo e uriti liberi 
3°, 4° e 5° neri e con pronoto bruno-rossiccio; antenne brune con i tre antennomeri 
basali giallo-rossicci; zampe giallo-rossicce. La reticolazione è netta solo sul pronoto, 
sul resto del corpo è svanita. Quella dell'addome è a maglie poligonali irregolari. 
I tubercoletti che coprono la superficie dell'avancorpo sono poco distinti. Edeago 
figg. 20-21. 

Comparazioni. La nuova specie è affine ad A. placita Cameron, 1939, 
dell'India. Se ne distingue per avere la parte apicale ventrale sporgente del bulbo 
basale dell'edeago poco saliente (molto saliente in placita) e per l'assenza di una 
lamella copulatrice ricurva basale del sacco interno dell'edeago (presente al contrario 
in placita). 

Atheta (Microdota) ipercristata sp. n. Figg- 22-25 

Holotypus a. China, Gansu, Xinlong Shan, ca. 70 Km S Lanzhou, 2225-2380 m, 
7.VIII.1994, A. Smetana leg. (MHNG). 

Paratypus; 1 S , stessa provenienza. 

Descrizione. Lunghezza 2,2 mm. Corpo lucido e nero, antenne comprese; 
zampe giallo-brune. La reticolazione del capo è netta, quella del pronoto e dell'addome 
è distinta e quella delle elitre è svanita. Le maglie di reticolazione dell'addome sono 
appena trasverse. I tubercoletti della superficie del capo sono poco salienti, più fitti ai 
lati e assenti sulla fascia mediana, quelli del pronoto e delle elitre sono salienti. Edeago 
figg. 23-24, sesto urotergo libero del maschio fig. 25. 

Comparazioni. A mia conoscenza la nuova specie è unica nell'ambito del 
sottogenere, dato che presenta una "crista apicalis" a sviluppo abnorme. 

Atheta (Microdota) lanzhouensis sp. n. Figg. 26-29 

Holotypus 6, China, Gansu, 120 Km S Lanzhou, Guanghe Xian Mai Jia, 2300 m, 
8. VII. 1994, A. Smetana leg. (MHNG). 



ALEOCHARINAE DELLA CINA 



917 




Figo. 15-21 

Habitus, edeago in visione laterale e ventrale e spermateca. 15-18: Atheta (Microdota) yanensis 
sp. n.; 19-21: Atheta (Microdota) gonggaensis sp. n. 



918 ROBERTO PACE 

Descrizione. Lunghezza 1 ,8 mm. Corpo lucido e bruno scuro con uriti liberi 3°, 
4° e 5° neri; antenne nero-brune con i tre antennomeri basali giallo-bruni; zampe gialle. 
La reticolazione del capo è distinta, quella del pronoto è netta e quella delle elitre e 
dell'addome è svanita. L'avancorpo è coperto di tubercoletti poco distinti, l'addome di 
tubercoletti salienti. Edeago figg. 27-28, sesto urotergo libero del maschio fig. 29. 

Comparazioni. Per la forma dell 'edeago e per alcuni caratteri dell' esoscheletro, 
la nuova specie può essere tassonomicamente vicina ad A. tuberculata (Kraatz, 1859) 
dell'India. Infatti entrambe le specie presentano occhi ridotti ed edeago di dimensioni 
pure ridotte, con poco accentuata parte apicale ventrale sporgente dal bulbo basale. La 
nuova specie se ne differenzia per gli occhi ancor più ridotti, per le elitre appena più 
larghe del pronoto (e non molto più larghe del pronoto come in tuberculata) e per 
V edeago, in visione ventrale, stretto, con apice largamente arrotondato e non con apice 
appuntito come si osserva in tuberculata. 

Atheta (Microdota) philamicula sp. n. Figg- 30-31 

Holotypus 9 , China, Sichuan, Langmusi, 3500-3600 m, A. Smetana leg. (MHNG). 
Descrizione. Lunghezza 2,1 mm. Corpo lucido e nero, antenne comprese; 
zampe giallo-brune con tibie nere. La reticolazione del capo e del pronoto è netta, 
quella del pronoto è dell'addome è distinta. I tubercoletti della superficie del capo e del 
pronoto sono distinti, quelli delle elitre sono svaniti e quelli dell'addome sono salienti. 
Il capo presenta un debole solco discale. Spermateca fig. 31, con parte prossimale nera, 
fortemente chitinizzata. 

Comparazioni. La forma della parte prossimale della spermateca della nuova 
specie è pressoché identica a quella della spermateca di A. amicula (Stephens, 1832), 
diffusa nella regione paleartica occidentale. Ma questa parte della spermateca della 
nuova specie è nera, mentre è incolore in amicula e il bulbo distale della spermateca 
della nuova specie ha scultura interna a maglie ampie ed è privo di introflessione 
apicale, mentre in amicula le maglie sono finissime ed è presente un'introflessione 
apicale. Gli occhi della nuova specie sono più corti delle tempie, mentre in amicula 
sono lunghi quanto le tempie. Le elitre della nuova specie sono molto più larghe del 
pronoto, mentre in amicula sono poco più larghe del pronoto. 

Atheta (Microdota) permixta sp. n. Figg- 32-33 

Holotypus 9, China. Gansu. Yonghai, ca. 20 Km SW Yuzhong, 2700-2800 m, 
9.VIII.1994, A. Smetana leg. (MHNG). 

Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 2,6 mm. Corpo lucido e nero pece; antenne nere; 
zampe bruno-rossicce con femori bruni. La reticolazione del capo, del pronoto e 
dell'addome è svanita, quella delle elitre è distinta. Le maglie di reticolazione della 
superficie dell'addome sono tras verse, quelle del quinto urotergo libero sono appena 
trasverse e distinte. I tubercoletti che coprono la superficie dell'avancorpo sono molto 
superficiali. Spermateca fig. 33. 

Comparazioni. La posizione sistematica della nuova specie è problematica. La 
forma della spermateca è tipica e simile a quella di alcune specie del genere Aloconota 



ALEOCHARINAE DELLA CINA 



919 




Figo. 22-31 

Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del maschio e spermateca. 
22-25: Atheta (Microdota) ipercristata sp. n.; 26-29: Atheta (Microdota) lanzhouensis sp. n.; 
30-31: Atheta (Microdota) philamicula sp. n. 



920 ROBERTO PACE 

Thomson, 1858, ma la ligula non è propria del genere Aloconota, ma del genere Atheta. 
La forma della spermateca della nuova specie è simile a quella di A. nana (Kraatz, 
1859) dello Sri Lanka, ma le dimensioni di quest'organo sono maggiori e la parte 
prossimale è notevolmente più dilatata. 

Atheta (Microdota) kadooriorum sp. n. Figg. 34-38 

Holotypus 6, Hong Kong, N.T., Kadoorie Agricultural Research Centre, IX-X.1991, 
Malaise trap, Ades leg. (MHNG). 

Descrizione. Lunghezza 1,4 mm. Corpo lucido e nero-bruno con pronoto e i due 
uriti basali bruni e con elitre giallo-brune; antenne nero-brune con i due antennomeri 
basali bruno-rossicci; zampe gialle. La reticolazione del capo è superficiale, quella del 
pronoto e delle elitre è molto svanita e quella dell'addome è assente. La punteggiatura 
del capo è fitta e assai superficiale. I tubercoletti che coprono la superficie del pronoto e 
delle elitre sono fini e molto svaniti, quelli dell'addome sono salienti. Edeago figg. 34- 
35, spermateca fig. 38. 

Comparazioni. Poiché ha taglia corporea minuta ed edeago di ridotte dimen- 
sioni, la nuova specie potrebbe essere vicina ad A. nana (Kraatz, 1859), dello Sri 
Lanka. Se ne distingue per l'enorme sviluppo degli occhi (occhi molto più corti delle 
tempie in nana) e per l'edeago notevolmente ricurvo al lato ventrale (appena ricurvo in 
nana). 

Etimologia. La nuova specie è dedicata ai fratelli Kadoorie, noti filantropi di 
Hong Kong, nella cui tenuta agricola sono state raccolte varie specie di Aleocharinae 
esaminate per la presente serie di lavori sulle Aleocharinae cinesi. 

Atheta (Microdota) alternantoides sp. n. Figg- 39-40 

Holotypus 9, China, Yunnan, Dali, 9.II.1993, de Rougemont leg. (MHNG). 
Paratypi: 1 e? (edeago non rinvenuto dentro l'addome) e 1 9 , stessa provenienza. 

Descrizione. Lunghezza 2,2 mm. Corpo lucido. Capo e uriti liberi 3°, 4° e metà 
basale del 5° neri, pronoto, i due uriti basali, la metà distale del 5° urotergo libero e 
l'estremità addominale giallo-rossicci, elitre giallo-brune con base gialla; antenne brune 
con i due antennomeri basali e la metà basale del terzo giallo-rossicci; zampe gialle. La 
reticolazione è distinta sul disco del capo, sulle elitre e sull'addome, sul pronoto è quasi 
vigorosa, sulle tempie e sul sesto urotergo libero è svanita. I tubercoletti della superficie 
del capo e del pronoto sono poco distinti, quelli delle elitre e dell'addome sono salienti. 
Spermateca fig. 40. 

Comparazioni. Per il colore del corpo e per la forma della spermateca, la nuova 
specie potrebbe essere affine ad A. piacila Cameron, 1939, dell'India. Se ne distingue 
per la base delle elitre e l'estremità addominale gialle o giallo-rossicce (elitre e addome 
interamente bruni in placito). Inoltre la nuova specie ha la spermateca a dimensioni 
minori nonostante la taglia corporea sia maggiore (2,2 mm invece di 2,0 mm come in 
piacila), priva di introflessione apicale del bulbo distale (presente invece nel bulbo 
distale della spermateca di piacila) e per la parte prossimale della stessa spermateca, 
breve e non lungamente protratta come in piacila. 



ALEOCHARINAE DELLA CINA 



921 




Figo. 32-40 

Habitus, edeago in visione laterale e ventrale e spermateca. 32-33: Atheta (Microdota) permixta 
sp. n.; 34-38: Atheta (Microdota) kadooriorum sp. n.; 39-40: Atheta (Microdota) alternantoides 
sp. n. 



922 ROBERTO PACE 

Atheta (Microdota) chinamicula sp. n. Figg. 41-45 

Holotypus S, China, Zhejiang, Hangzhou, 27.IV. 1993, de Rougemont leg. (MHNG). 
Paratypi: 13 es., stessa provenienza. 

Descrizione. Lunghezza 1,7 mm. Corpo lucido e bruno con uriti liberi 3°, 4° e 
5° neri; antenne brune con i due antennomeri basali giallo-bruni; zampe gialle. La 
reticolazione del disco del capo è netta, composta di maglie isodiametriche ampie, 
quella del pronoto è vigorosa, quella delle elitre è svanita, quella dei quattro uroterghi 
basali è a maglie lievemente trasverse e distinte e quella del quinto urotergo libero è 
netta, composta di maglie appena ovali. Il capo presenta una superficie coperta di 
tubercoletti distinti posti ai lati e sulla regione occipitale: sono assenti sulla fascia 
mediana che dal disco all'occipite è impressa da un solco. I tubercoletti della superficie 
del pronoto e delle elitre sono superficiali, quelli dell'addome sono salienti. Edeago 
fìgg. 42-43, spermateca fig. 44, sesto urotergo libero del maschio fig. 45. 

Comparazioni. In base alla struttura della spermateca, la nuova specie è 
probabilmente affine ad A. contingens Cameron, 1939, dell'India. Infatti la parte distale 
di quest'organo è molto simile nelle due specie. Tuttavia la spermateca della nuova 
specie ha minore sviluppo in lunghezza. Inoltre le elitre della nuova specie sono distin- 
tamente più corte, con loro sutura lunga quanto il pronoto, mentre in contingens la 
sutura delle elitre è un quinto più lunga della lunghezza del pronoto. 

Atheta (Microdota) elytralis sp. n. Figg- 46-49 

Holotypus 6 , China, Zhejiang, Trianmushan, 29.IV. 1993, de Rougemont leg. (MHNG). 
Paratypi: 6 ó* e 2 9 , stessa provenienza. 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e bruno con elitre, base ed 
estremità addominale di un bruno chiaro; antenne brune con i due antennomeri basali 
giallo-bruni; zampe giallo-rossicce. La reticolazione dell'avancorpo è netta e a maglie 
isodiametriche ampie sul capo. La reticolazione dell'addome è distinta e lievemente 
trasversa sui quattro uriti basali, sul quinto libero è isodiametrica e pure distinta. La 
punteggiatura del capo è distinta e assente per una larga fascia longitudinale mediana. I 
tubercoletti che coprono il pronoto e le elitre sono poco distinti, quelli dell'addome 
sono molto salienti. Spermateca fig. 47, edeago figg. 48-49. 

Comparazioni. La nuova specie è distinta da A. subluctuosa Cameron, 1939, 
dell'India, per avere le elitre appena più lunghe e poco più lunghe e più larghe del 
pronoto come in subluctuosa. L'edeago della nuova specie, in visione ventrale, ha 
bulbo basale molto largo, mentre in subluctuosa tale bulbo basale è poco largo. Il bulbo 
distale della spermateca della nuova specie è privo di introflessione apicale, presente al 
contrario nel bulbo distale della spermateca di subluctuosa. 

Atheta (Microdota) nanior sp. n. Figg. 50-53 

Holotypus 6, China, Yunnan, Xishuangbanna, Sanchahe, elephant res., 24.1.1992, de 
Rougemont leg. (MHNG). 

Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 1,3 mm. Corpo lucido e nero-bruno con uriti liberi 4° 

e 5° neri; antenne nere; zampe giallo-brune. La reticolazione del capo e dell'addome è 



ALEOCHARINAE DELLA CINA 



923 




Figo. 41-49 

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 
41-45: Atheta (Microdota) chinamicula sp. n.; 46-49: Atheta (Microdota) elytralis sp. n. 



924 ROBERTO PACE 

estremamente svanita, quella sul resto della superficie del corpo è assente. L'intera 
superficie corporea è coperta di tubercoletti molto salienti e fitti. Edeago figg. 51-52, 
spermateca fig. 53. 

Comparazioni. Per la taglia corporea estremamente ridotta (1,3 mm), la nuova 
specie sembra A. inquinula (Gravenhorst, 1802), diffusa nella regione paleartica 
occidentale. Ma l'edeago e la spermateca sono nettamente differenti. L'edeago di 
inquinula non è profondamente arcuato al lato ventrale come quello della nuova specie 
e la spermateca di inquinula, breve e tozza con distinti bulbi distale e prossimale, si 
distingue nettamente dalla spermateca della nuova specie che è sottile, lunga e varia- 
mente sinuata. 

Atheta (Microdota) laminarum sp. n. Figg- 54-56 

Holotypus â. China, Yunnan, Xishuangbanna, Chayanhe F.P., 24.1.1993, de 
Rougemont leg. (MHNG). 

Paratypi: 5 S e 6 9 , stessa provenienza, ma Sanchahe, elephant res., 24.1.1992, de 
Rougemont leg. 

Descrizione. Lunghezza 1,8 mm. Corpo lucido e nero con elitre nero-brune; 
antenne nere; femori bruni, tibie e tarsi anteriori e medi giallo-bruni, i posteriori bruno 
chiari. La reticolazione del disco del capo è nettissima e a maglie isodiametriche ampie, 
quella del resto della superficie del capo è svanita come quella delle elitre e dei quattro 
uroterghi basali che l'hanno a maglie lievemente trasverse, quella del quinto urotergo 
libero è netta come quella del pronoto e a maglie isodiametriche. La punteggiatura del 
capo è indistinta. I tubercoletti della superficie del pronoto sono salienti, quelli delle 
elitre sono superficiali e quelli dell'addome sono molto salienti. Edeago figg. 55-56, la 
spermateca non è sufficientemente scarificata perciò indistinta. 

Comparazioni. La nuova specie è distinta da amiculoides Cameron, 1939, 
dell'India, per avere lunghissime setole isolate ai lati del corpo e delle meso-metatibie, 
assenti in amiculoides, per avere l'edeago meno sviluppato, in visione ventrale allargato 
nella regione preapicale. carattere questo assente nella regione preapicale dell' edeago di 
amiculoides. 

Atheta (Dicolyota) hongkongiphila sp. n. Figg- 57-61 

Holotypus 6. Hong Kong, XII. 1995-1. 1996, de Rougemont leg. (MHNG). 
Paratypi: 7 es., stessa provenienza. 

Descrizione. Lunghezza 1,8 mm. Corpo lucidissimo e nero, antenne comprese 
con i due antennomeri basali nero-bruni; zampe gialle. La reticolazione del capo e del 
pronoto è netta e a maglie molto ampie, quella delle elitre è a maglie ancor più ampie di 
quelle del pronoto e altrettanto nette, la reticolazione dell'addome è trasversa e svanita. 
La punteggiatura del capo e del pronoto è rada e fine, quella delle elitre è indistinta. 
Tubercoletti fini e radi stanno sulla superficie del pronoto. Edeago figg. 57-58, 
spermateca fig. 59, sesto urotergo libero del maschio fig. 61. 

Comparazioni. In base alla forma della spermateca, la nuova specie sembra 
affine ad A. scabriventris Cameron, 1939, del Tibet. Il bulbo basale della spermateca 



ALEOCHARINAE DELLA CINA 



925 




FiGG. 50-60 

Habitus, edeago in visione laterale e ventrale e spermateca. 50-53: Atheta (Microdota) nanior 
sp. n.; 54-56: Atheta (Microdota) laminarum sp. n.; 57-60: Atheta (Dicolyota) hongkongiphila 
sp. n. 



926 ROBERTO PACE 

della nuova specie è molto lungo, mentre quello di scabriventris è corto, quasi ovale. 
Inoltre la sutura delle elitre della nuova specie è lunga quanto la lunghezza del pronoto, 
mentre la sutura delle elitre di scabriventris è nettamente più lunga del pronoto, circa 
un quarto più lunga. Il pronoto della nuova specie è nettamente trasverso, mentre quello 
di scabriventris è appena trasverso. Non è noto il maschio di scabriventris. 

Atheta (Dicolyota) ponderata sp. n. Figg. 62-63 

Holotypus 9 , China, Yunnan, Ruili, ca. 700 m, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 1,9 mm. Corpo lucido e nero con estremità addominale 
bruno-scura; antenne nere; zampe brune con tibie anteriori e tarsi gialli. La retico- 
lazione del capo e del pronoto è netta, quella delle elitre è distinta e quella dell'addome 
è superficiale. I tubercoletti della superficie del capo sono molto salienti. La punteggia- 
tura del pronoto è svanita. Le elitre presentano una superficie d'aspetto rugoso per la 
presenza di tubercoletti fini, fitti e superficiali. I tubercoletti della superficie dell'ad- 
dome sono molto salienti. Il capo ha un solco mediano distinto. Il pronoto presenta una 
impressione mediana posteriore allungata. Spermateca fig. 63. 

Comparazioni. Il base alla forma della spermateca, la nuova specie, anche se 
evidentemente affine, è nettamente distinta da A. scabriventris Cameron, 1939 del 
Tibet. Infatti il bulbo prossimale della spermateca della nuova specie è esile come il 
resto della spermateca e non largamente ellittico come quello di scabriventris. 

Atheta (Diaprepota) ruiliensis sp. n. Figg- 64-69 

Holotypus ó\ China, Yunnan, Ruili, ca. 700 m, 3.II.1993, de Rougemont leg. (MHNG). 
Paratypus: 1 9 stessa provenienza. 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e giallo rossiccio con capo e 
uriti liberi 3°, 4° e base del 5° neri e con elitre di un giallo sporco con lati estermi e 
zona periscutellare bruni; antenne nere con i due antennomeri basali e la base del terzo 
giallo-rossicci; zampe gialle. La reticolezione del disco del capo è nettissima e svanita 
ai suoi lati, quella del pronoto è netta e quella delle elitre e dell'addome è distinta. La 
punteggiatura del capo è svanita e assente sulla fascia mediana, quella del pronoto è 
poco distinta sul disco e distinta ai lati. I tubercoletti che coprono la superficie delle 
elitre sono poco distinti, quelli dell'addome sono salienti. Edeago figg. 65-66, sesto 
urotego libero del maschio fig. 67, spermateca fig. 69. 

Comparazioni. La nuova specie è distinta da A. subnigritula Cameron, 1950, di 
Selangor, per le elitre di un giallo sporco con lati bruni e non uniformemente bruno 
rossicce, per il quarto antennomero trasverso e non più lungo che largo come in 
subnigritula e per le tempie sfuggenti, e non largamente arrotondate come in subnigri- 
tula. Di quest'ultima specie non è noto il maschio. 

Atheta (Bessobia) gentilior sp.. n. Figg- 70-74 

Holotypus 9, China, Yunnan, Kunming, 1. II. 1993, de Rougemont leg. (MHNG). 
Paratypi: 1 d, stessa provenienza; 1 ó\ China, Yunnan, Dali, 9.II.1993, de Rougemont 
leg. 



ALEOCHARINAE DELLA CINA 



927 




Figo. 61-67 

Sesto urotergo libero del maschio, habitus, spermateca ed edeago in visione laterale e ventrale. 
61: Atheta (Dicolyota) hongkongiphila sp. n.; 62-63: Atheta (Dicolyota) ponderata sp. n.; 64- 
67: Atheta (Diaprepota) ruiliensis sp. n. 



928 



ROBERTO PACE 




Figo. 68-73 



Habitus, spermateca ed edeago in visione laterale e ventrale. 68-69: Atheta (Diaprepota) 
ruiliensis sp. n.; 70-73: Atheta (Bessobia) gentilior sp. n. 



ALEOCHARINAE DELLA CINA 929 

Descrizione. Lunghezza 2,7 mm. Corpo lucido e bruno scuro con elitre brune e 
con addome nero; antenne nere; zampe bruno-rossicce con femori bruni. La reti- 
colazione dell'avancorpo è netta, quella dell'addome è molto svanita, composta di 
maglie estremamente trasverse. Il capo è coperto di tubercoletti superficiali, il pronoto 
di tubercoletti salienti e le elitre di tubercoletti svaniti. Edeago figg. 71-72, spermateca 
fig. 73, sesto urotergo libero del maschio fig. 74. 

Comparazioni. La nuova specie è simile, ma ben distinta da A. smetanaorum 
Pace, 1991, del Nepal, per avere l' edeago non ristretto nella regione mediana, in visione 
ventrale, per l'assenza di due prominenze semicircolari mediane al margine posteriore 
del sesto urotergo libero del maschio, per l'assenza di doppia carena mediana al quinto 
urotergo libero del maschio e per le elitre meno larghe rispetto al pronoto (più larghe in 
smetanaorum). La spermateca della nuova specie ha forma simile a quella di A. occulta 
(Erichson, 1839), diffusa dall'Europa, alla Siberia e al Giappone, ma quella della nuova 
specie è priva della profonda e larga introflessione apicale del bulbo distale della 
spermateca stessa. 

Atheta (Bessobia) wutaishanensis sp. n. Figg. 75-78 

Holotypus 6, China, Shanxi, Wutaishan, 4-5. VI. 1993, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 3,5 mm. Corpo lucido e nero, antenne comprese; 
zampe brune con femori neri. L'avancorpo è coperto di reticolazione nettissima, 
l'addome l'ha svanita e a maglie molto trasverse, ma nel fondo dei solchi trasversi 
basali è vigorosa. I tubercoletti della superficie del capo e del pronoto sono salienti, 
quelli delle elitre sono svaniti. Edeago figg. 76-77, sesto urotergo libero del maschio 
fig. 78. 

Comparazioni. Poiché la parte apicale dell'edeago, in visione ventrale, è note- 
volmente stretta, la nuova specie è chiaramente distinta da A. smetanaorum Pace, 1991, 
del Nepal che presenta tale parte apicale larga. Inoltre il margine posteriore mediano del 
sesto urotergo libero del maschio della nuova specie è arcuato, mentre quello di 
smetanaorum presenta due prominenze arcuate mediane. 

Atheta (Bessobia) peranomala sp. n. Figg. 79-82 

Holotypus S, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 26. VII. 1994, 
A. Smetana leg. (MHNG). 

Paratypi: 10 â, stessa provenienza. 

Descrizione. Lunghezza 2,6 mm. Corpo lucido e bruno con capo nero-bruno e 
con addome giallo-rossiccio con uriti liberi 3° e 4° neri; antenne brune con i tre 
antennomeri basali giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo è 
netta, quella del pronoto è nettissima, quella delle elitre è distinta, quella dei tre 
uroterghi basali è svanita e composta di maglie lievemente trasverse e quella degli 
uroterghi liberi 4° e 5° è distinta. Il quinto urotergo libero del maschio presenta dei 
robusti tubercoli salienti. La punteggiatura del capo è svanita. I tubercoletti che coprono 
la superficie del pronoto e delle elitre sono distinti. Edeago figg. 80-81, sesto urotergo 
libero del maschio fig. 82. 



930 



ROBERTO PACE 




Figo. 74-78 

Sesto urotergo libero del maschio, habitus ed edeago in visione laterale e ventrale. 74: Atheta 
(Bessobia) gentilior sp. n.; 75-78: Atheta (Bessobia) wutaishanensis sp. n. 



ALEOCHARINAE DELLA CINA 93 \ 

Comparazioni. Il margine posteriore del sesto urotergo libero del maschio della 
nuova specie è molto simile a quello di A. smetanaorum Pace, 1991, ma l'edeago delle 
due specie presenta numerosi caratteri differenziali, tra cui l'apice strettissimo, in 
visione ventrale, della nuova specie, mentre in smetanaorum è larghissimo. 

Atheta (Bessobia) pergranulosa sp. n. Figg. 83-85 

Holotypus S, China, Sichuan, Gongga Shan, above camp 3, 3050 m, 22. VII. 1994, 
A. Smetana leg. (MHNG). 

Descrizione. Lunghezza 2,6 mm. Avancorpo debolmente lucido, addome 
lucido. Corpo bruno con capo bruno scuro e addome giallo-rossiccio con gli uriti liberi 
3° e 4° neri; antenne brune con i quattro antennomeri basali giallo-rossicci; zampe 
giallo-rossicce. La reticolazione del capo e del pronoto è netta, quella delle elitre e 
deir addome è distinta. La punteggiatura del capo è indistinta. I tubercoletti che stanno 
sulla superficie del pronoto e delle elitre sono poco salienti. Il quinto urotergo libero del 
maschio presenta robusti tubercoli salienti. Edeago figg. 84-85. 

Comparazioni. La nuova specie per la forma del margine posteriore del sesto 
urotergo libero del maschio sembra affine ad A. smetanaorum Pace, 1991, del Nepal, 
ma l'edeago ha minore sviluppo rispetto all'edeago di smetanaorum, ha lati, in visione 
ventrale, rettilinei e non profondamente sinuati come in smetanaorum ad apice brusca- 
mente ristretto a triangolo (in visione ventrale) e non ristretto gradualmente come in 
smetanaorum. 

Atheta (Oreostiba) yonghaicola sp. n. Figg 86-89 

Holotypus 6, China, Gansu, Yonghai ca. 20 Km SW Yuzhong, 2700-2800 m, 
9. Vili. 1 994, A. Smetana leg. (MHNG). 

Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 2,2 mm. Corpo lucido e nero-bruno con capo e addome 
neri; antenne brune con i due antennomeri basali bruno-rossicci; zampe bruno-rossicce. 
La reticolazione del capo e degli uroterghi liberi 5° e 6° è netta, quella del pronoto è 
distinta e quella delle elitre e dei quattro uroterghi basali è svanita. La punteggiatura del 
capo è svanita, quella del pronoto è fine e distinta. Tubercoletti superficiali coprono la 
superficie delle elitre. Edeago figg. 87-88, spermateca fig. 89. 

Comparazioni. La nuova specie è affine ad A. tibialis (Heer, 1842) e specie 
affini diffuse dalla Scozia alla regione alpina, Pirenei e Caucaso. Infatti alcuni caratteri 
dell'edeago e della spermateca della nuova specie come il colore nero o nero bruno del 
corpo, si rinscontrano anche in tibialis. La nuova specie è distinta da essa, oltre che per 
la differente armatura genitale interna dell'edeago, per avere la parte apicale ventrale 
sporgente del bulbo basale, molto più sviluppata in modo tale che la lunghezza della 
"crista apicalis" è uguale in lunghezza alla lunghezza del segmento che prende origine 
dalla stessa "crista apicalis" e termina all'angolo ventrale, mentre in tibialis e specie 
affini, tale segmento è nettamente molto più breve della lunghezza della corrispondente 
"crista apicalis". Inoltre l'apice dell'edeago della nuova specie, in visione laterale, non 
presenta alcun dentino smussato come si osserva in tibialis. La spermateca della nuova 



932 



ROBERTO PACE 




FlGG. 79-86 

Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del maschio. 79-82: Atheta 
(Bessobia) peranomala sp. n.; S3-S5: Atheta (Bessobia) pergramilosa sp. n.; 86: Atheta (Oreo- 
stiba) yonghaicola sp. n. 



ALEOCHARINAE DELLA CINA 933 

specie non possiede una netta introflessione apicale del bulbo distale, ben presente e 
profonda in tibialis, e ha la parte prossimale con evidenza molto più sviluppata in 
lunghezza. 

Atheta (Oreostiba) semitibialis sp. n. Figg. 90-93 

Holotypus 6, China, Gansu, 120 Km S Lanzhou, Guanghe Xian Ma Jia, 2300 m, 
8.VII.1994, A. Smetana leg. (MHNG). 

Paratypus: 1 6 , stessa provenienza. 

Descrizione. Lunghezza 2,6 mm. Corpo lucido e nero-bruno; antenne nere; 
zampe giallo-brune. La reticolazione è svanita sul capo, sulle elitre e sui quattro 
uroterghi basali, è distinta sul pronoto ed è netta sul quinto urotergo libero. La 
punteggiatura del capo è ombelicata e svanita. I tubercoletti della superficie del pronoto 
sono distinti, quelli delle elitre sono superficiali. Edeago figg. 91-92, sesto urotergo 
libero del maschio fig. 93. 

Comparazioni. La nuova specie è probabilmente affine ad A. tibialis (Heer, 
1842) per i caratteri dell'edeago, per il colore del corpo e per il profilo del margine 
posteriore del sesto urotergo libero del maschio. E' tuttavia da essa ben differenziata 
per avere la parte apicale dell'edeago nettamente più stretta, in visione ventrale, per 
avere la parte mediana del margine posteriore del sesto urotergo libero del maschio 
nettamente protratta all' indietro (appena protratta in tibialis) e per l'armatura genitale 
interna dell'edeago, composta di un pezzo copulatore ricurvo e stretto, assente 
all'interno dell'edeago di tibialis. 

Atheta (Oreostiba) shanicola sp. n. Figg- 94-95 

Holotypus 9, China, Sichuan, Lang Musi, 3500-3600 m, 13.VII.1994, A. Smetana leg. 
(MHNG). 

Descrizione. Lunghezza 3,2 mm. Corpo lucido e nero-bruno con addome nero; 
antenne brune con i tre antennomeri basali bruno-rossicci; zampe giallo-rossicce. La 
reticolazione del capo e del pronoto è netta, quella delle elitre e dell'addome è svanita, 
sull'addome è a maglie molto trasverse. I tubercoletti della superficie del capo e del 
pronoto sono distinti, quelli delle elitre sono superficiali. Spermateca fig. 94-95. 

Comparazioni. La forma della spermateca della nuova specie è simile a quella 
di A. sibirica Maeklin, 1880, diffusa in Siberia, ma il bulbo distale della spermateca 
della nuova specie è nettamente più robusto, con forte introflessione apicale, mentre 
in sibirica non vi è distinta introflessione apicale. Inoltre la lunghezza della sutura 
delle elitre è maggiore della lunghezza del pronoto (indice 1,16), mentre in sibirica la 
sutura delle elitre è un po' più corta della lunghezza del pronoto (indice 0,90). 

Etimologia. Il nome della nuova specie significa "colei che abita i monti", dal 
sostantivo cinese "shan" che significa monti. 

Atheta (s. str.) linxiensis sp. n. Figg. 96-99 

Holotypus o\ Gansu, Dalijia Shan, 60 Km W Linxia, 3475 m, 1 1. VII. 1994, A. Smetana 
leg. (MHNG). 



934 



ROBERTO PACE 




Figo. 87-92 

Edeago in visione laterale e ventrale, spermateca e habitus. 87-89: Atheta (Oreostiba) yonghai- 
cola sp. n.; 90-92: Atheta (Oreostiba) semitibialis sp. n. 



ALEOCHARINAE DELLA CINA 



935 






Figo. 93-96 

Sesto urotergo libero del maschio, habitus e spermateca. 93: Atheta (Oreostiba) semitibialis 
sp. n.; 94-95: Atheta (Oreostiba) shanicola sp. n.; 96: Atheta (s. str.) linxiensis sp. n. 



936 ROBERTO PACE . 

Descrizione. Lunghezza 3,4 mm. Corpo lucidissimo e nero pece con addome 
nero; antenne nere; zampe bruno-rossicce con femori bruni. La reticolazione del capo e 
del pronoto è netta, quella delle elitre è superficiale e quella dell'addome è molto 
svanita. Il corpo è coperto di tubercoletti distinti. Edeago figg. 97-98, sesto urotergo 
libero del maschio fig. 99. 

Comparazioni. La nuova specie si pone in posizione tassonomica intermedia tra 
A. triangulum (Kraatz, 1858a) e A. aquatica (Thomson, 1852), entambe della regione 
paleartica occidentale. Della prima la nuova specie ha il profilo ventrale dell' edeago, 
della seconda una prominenza ventrale presso la "crista apicalis". In visione ventrale 
l'apice dell 'edeago è più largo di quello di entrambe le specie note. Inoltre il colore 
nero del corpo è ben differente da quello di triangulum che mostra elitre chiare con 
fascia obliqua oscura a partire dagli omeri e diretta verso la sutura. 

Atheta (s. str.) serraculter sp. n. Figg- 100-104 

Holotypus S, China, Gansu, pass btw Hezuo-Amqog, 3300 m, 12. VII. 1994, 
A. Smetana leg. (MHNG). 

Paratypi: 19 es., stessa provenienza. 

Descrizione. Lunghezza 3,9 mm. Corpo lucido e bruno con elitre giallo-brune e 
con addome nero; antenne brune con i tre antennomeri basali bruno-rossicci; zampe 
rossicce. La reticolazione del capo è netta, quella del pronoto e delle elitre è distinta e 
quella dell'addome è molto svanita, composta di maglie molto trasverse e ondulate. La 
punteggiatura del capo è molto svanita. I tubercoletti della superficie del pronoto e delle 
elitre sono distinti. Edeago figg. 101-102, spermateca fig. 103, sesto urotergo libero del 
maschio fig. 104. 

Comparazioni. In base alla forma dell'edeago e particolarmente per la presenza 
del robusto dente ventrale dell'edeago stesso, la nuova specie appare affine ad 
A. iturupensis Bernhauer, 1907, del Giappone. Se ne distingue per il profilo ventrale 
apicale dell'edeago, sinuato (arcuato in iturupensis), per il lato distale del dente ventrale 
seghettato (non seghettato in iturupensis), per l'apice dell'edeago, in visione ventrale, 
stretto e lungo (corto e largo in iturupensis) e per il margine posteriore del sesto 
urotergo libero del maschio polilobato (non lobato, ad andamento quasi rettilineo in 
iturupensis). 

Etimologia. Il nome della nuova specie significa "coltello a sega" a motivo del 
margine seghettato del dente ventrale dell'edeago. 

Atheta (s. str.) kazakhstanensis sp. n. Figg- 105-108 

Holotypus ó\ Kazakhstan, Alma Ata. 1000 m. 18.IX.1994, de Rougemont leg. (MHNG). 

Paratypi: 5 9, stessa provenienza. 

Descrizione. Lunghezza 3,8 mm. Corpo lucido e nero con elitre brune con zona 
periscutellare nera; antenne brune con antennomero basale bruno-rossiccio; zampe 
giallo-rossicce con femori posteriori bruni e i medi con lato esterno bruno. La retico- 
lazione del capo e del pronoto è distinta, quella delle elitre è netta, quella degli uro- 
terghi anteriori è molto svanita e quella degli uroterghi posteriori è superficiale. I 



ALEOCHARINAE DELLA CINA 



937 




Figo. 97-103 

Edeago in visione laterale e ventrale, sesto urotergo libero del maschio, habitus e spermateca. 
97-99: Atheta (s. str.) linxiensis sp. n.; 100-103: Atheta (s. str.) serraculter sp. n. 



938 ROBERTO PACE 

tubercoletti della superficie del capo sono poco salienti e assenti sulla fascia mediana, 
quelli del pronoto sono distinti e quelli delle elitre sono salienti. Edeago figg. 106-107, 
spermateca fig. 108. 

Comparazioni. La presenza del dente ventrale dell' edeago permette di porre la 
nuova specie in posizione tassonomica vicina ad A. castanoptera (Kraatz, 1858a), 
diffusa nella regione paleartica occidentale. Ma il lungo dente ventrale dell'edeago 
della nuova specie è robusto e più corto (lungo e sottile in castanoptera) e la struttura 
della spermateca della nuova specie ha parte prossimale a matassa meno aggrovigliata. 

Pelioptera (Tropimenelytron) viatica sp. n. Figg- 109-113 

Holotypus 9, China. Sichuan, Gongga Shan, above camp 3, 3050 m, 22.VII.1994, 
A. Smetana leg. (MHNG). 

Paratypi: 1 2, stessa provenienza; 1 2, stessa provenienza, ma 3300-3350 m, 23. VII. 
1994, A. Smetana leg.; 8 es., stessa provenienza, ma camp 2, 2800 m, 28. VII. 1994, A. Smetana 
leg. 

Descrizione. Lunghezza 3,6 mm. Corpo lucido con addome lucidissimo. 
Corpo nero pece con addome nero; antenne nero pece con i tre antennomeri basali 
bruni; zampe rossicce. La reticolazione dell'avancorpo è netta, quella dell'addome è 
svanita, composta di maglie poligonali irregolari. I tubercoletti della superficie del 
capo sono svaniti, quelli dell'addome sono netti. La punteggiatura del pronoto è netta, 
quella delle elitre è indistinta. Edeago figg. 110-111, spermateca fig. 112, sesto 
urotergo libero del maschio fig. 113. 

Comparazioni. La nuova specie appare simile a P. angustie ollis Cameron, 
1939, dell'India, se si osservano edeago e spermateca. Ma le elitre e il quinto urotergo 
libero del maschio non presentano carene come in angusticollis. Inoltre il bulbo 
distale della spermateca è molto più trasverso in angusticollis che nella nuova specie e 
la parte prossimale è molto protratta in angusticollis e breve nella nuova specie. 

Pelioptera (Tropimenelytron) sakhalinensis sp. n. Figg. 114-117 

Holotypus 3, Russia, Sakhalin, Aniva distr. 5 Km W Petropaulovskiy, tributary of 
Lyutoga river, 20-21. VII. 1993, Putz & Wrase leg. (CASS). 
Paratypus: 1 2 , stessa provenienza. 

Descrizione. Lunghezza 2,8 mm. Corpo lucido e bruno con pronoto, elitre, 
margine posteriore dei tre uroterghi basali, metà posteriore del quarto urite libero e 
l'intero urite quinto giallo-bruni; antenne brune con i tre antennomeri basali bruno- 
rossicci; zampe gialle. La reticolazione del capo è più netta in avanti che all' indietro, 
quella del pronoto e dell'addome è pure netta, quella delle elitre è svanita. I tubercoletti 
della superficie del capo sono svaniti, quelli del pronoto e delle elitre sono salienti. 
Spermateca fig. 1 15, edeago figg. 116-1 17. 

Comparazioni. La nuova specie è simile a P. angusticolis Cameron, 1939, 
dell'India, ma ha occhi ed elitre molto ridotti (occhi lunghi quanto le tempie ed elitre 
molto più lunghe del pronoto in angusticollis). Il bulbo distale della spermateca è 
voluminoso e piriforme nella nuova specie, schiacciato e reniforme è quello di 
angusticollis. 



ALEOCHARINAE DELLA CINA 



939 




Figo. 104-108 

Sesto urotergo libero del maschio, habitus, edeago in visione laterale e ventrale e spermateca. 
104: Atheta (s. str.) serraculter sp. n.; 105-108: Atheta (s. str.) kazakhstanensis sp. n. 



940 



ROBERTO PACE 




FlGG. 109-117 

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 
109-113: Pelioptera (Tropimenelytron) viatica sp. n.; 114-117: Pelioptera (Tropimenelytron) 
sakhalinensis sp. n. 



ALEOCHARINAE DELLA CINA 94] 



Pelioptera (s. str.) samchunensis sp. n. Figg. 1 18-121 

Holotypus 9, Hong Kong, XII. 1995-1. 1996, de Rougemont leg. (MHNG). 

Paratypi: 4 es., stessa provenienza; 3 2 , Hong Kong, Kadoorie Agricultural Research 
Centre, flight interception trap, 19-3 I.V. 1996, de Rougemont leg.; 1 2, stessa provenienza, ma 
VI. 1996, de Rougemont leg.; 69 es., Hong Kong, Kadoorie Farm, V.1996, de Rougemont leg.; 
1 e? e 1 2, China, Zhejiang Prov., Lin'an County, 1000 m, W Tianmu Shan N.R., 18.V.1996, 
J. Cooter leg. 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e nero con elitre giallo-brune 
sul disco; antenne nere; zampe giallo-rossicce. L'avancorpo è coperto di reticolazione 
molto svanita, la superficie dell'addome è priva di reticolazione. La punteggiatura del 
capo è superficiale. I tubercoletti della superficie del pronoto sono fini e poco distinti, 
quelli delle elitre sono salienti. Edeago figg. 1 19-120, spermateca fig. 121. 

Comparazioni. La nuova specie, in base alla forma dell'edeago e della sper- 
mateca, è affine a P. opaca Kraatz, 1857, dello Sri Lanka, eaP. exasperata (Kraatz, 
1859), dell'India, dello Sri Lanka e del Nepal. Essa è distinta da entrambe per avere 
gli occhi più lunghi delle tempie (occhi nettamente più corti delle tempie in opaca ed 
exasperata) e per l'introflessione apicale del bulbo distale della spermateca volu- 
minosa e non conica come in opaca o strettissima e profonda come in exasperata. 

Etimologia. La nuova specie prende nome da Samchun, il fiume di Hong 
Kong. 

Pelioptera (s. str.) kwantungensis sp. n. Figg 122-125 

Holotypus ó\ Hong Kong, XII. 1995-1. 1996, de Rougemont leg. (MHNG). 

Paratypi: 18 es., stessa provenienza; 2 6 , Hong Kong, Kadoorie Agricultural Research 
Centre, 19-3 I.V. 1996, de Rougemont leg.; 1 â , stessa provenienza, ma flight interception trap, 
VI. 1996, de Rougemont; 1 2, stessa provenienza, ma Vili. 1996, de Rougemont leg.; 1 6, 
Hong Kong, Kadoorie Farm, V.1996, de Rougemont leg.; 6 es., Hong Kong, Tai Po, III. 1996 e 
V.1996, de Rougemont leg. 

Descrizione. Lunghezza 2,2 mm. Avancorpo debolmente opaco, addome 
lucido. Corpo bruno con apice addominale bruno-rossiccio; antenne brune; zampe 
gialle. La reticolazione del capo è netta, quella del pronoto e delle elitre è svanita. La 
punteggiatura del capo e del pronoto è quasi indistinta. Tubercoletti svaniti coprono la 
superficie delle elitre. I caratteri sessuali secondari dell'addome del maschio possono 
essere assenti. Edeago figg. 123-124, spermateca fig. 125. 

Comparazioni. Per la forma dell'edeago e della spermateca, la nuova specie è 
affine sia a P. unituberculata (Bernhauer, 1915a), della Nuova Britannia, che da 
P. sagadensis Pace, 1990a, delle Filippine e del Vietnam. Da entrambe è distinta per 
avere la spermateca più sviluppata, con parte prossimale più protratta. 

Etimologia. La nuova specie prende nome dalla provincia cinese di Kwang- 
tung confinante con Hong Kong, sua località tipica. 

Pelioptera (Geostibida) Hi sp. n. Figg. 126-127 

Holotypus 9, China, Hebei Prov., Yongniang, 6.X.1995, Shuqiang Li leg. (MHNG). 
Paratypi: 2 2 , stessa provenienza. 

Descrizione. Lunghezza 2,0 mm. Corpo lucidissimo e giallo-rossiccio con capo 

e uriti liberi 3°, 4° e 5° (tranne il margine posteriore rossiccio) bruni, elitre giallo-brune; 



942 



ROBERTO PACE 




FlGG. 118-125 

Habitus, edeago in visione laterale e ventrale e spermateca. 1 18-121: Pelioptera (s. str.) 
samchunensis sp. n.; 122-125: Pelioptera (s. str.) kwantungensis sp. n. 



ALEOCHARINAE DELLA CINA 943 

antenne giallo-rossicce con undicesimo antennomero rossiccio; zampe gialle. La 
reticolazione della superficie del corpo è svanita. L'avancorpo è coperto di tubercoletti 
estremamente svaniti o indistinti, l'addome ha tubercoletti distinti. Spermateca fig. 127. 

Comparazioni. La nuova specie è distinta da P. indica Cameron, 1939, 
dell'India, per avere le elitre giallo-rossicce e non brune come in indica, per il quarto 
antennomero molto trasverso (lungo quanto largo in indica) e per la presenza di 
introflessione apicale del bulbo distale della spermateca (introflessione assente nel 
bulbo distale della spermateca di indica). 

Etimologia. La nuova specie è dedicata al suo raccoglitore, il Dr. Shuqiang Li 
di Stuttgard (Germania). 

Pelioptera (Geostibida) kowloonensis sp. n. Figg. 128-132 

Holotypus 9 , Hong Kong, XII. 1995-1. 1996, de Rougemont leg. (MHNG). 

Paratypi: 1 9, stessa provenienza; 5 es., Hong Kong, Tai Po, III. 1996, de Rougemont leg. 

Descrizione. Lunghezza 1,9 mm. Corpo lucidissimo e nero-bruno, antenne 
comprese; zampe gialle con femori giallo-bruni. La reticolazione del capo è molto 
svanita, quella del pronoto è superficiale e quella delle elitre e dell'addome è distinta, 
sull'addome a maglie trasverse. La punteggiatura del capo e del pronoto è indistinta, 
quella delle elitre è rada e fine. I tubercoletti della superficie dell'addome sono distinti. 
Edeago figg. 129-130, sesto urotergo libero del maschio fig. 131, spermateca fig. 132. 

Comparazioni. La nuova specie è distinta da P. indica Cameron, 1939, 
dell'India, per avere l'edeago molto meno sviluppato e la spermateca con bulbo distale 
ellittico e con parte prossimale a spirale e non con bulbo distale sferico e con parte 
prossimale rettilinea come in indica. 

Etimologia. La nuova specie prende nome da Kowloon, la penisola di Hong 
Kong. 

Pelioptera (Geostibida) eremita sp. n. Figg. 133-134 

Holotypus 9 , Hong Kong, Kadoorie Agricultural Research Centre, flight interception 
trap, 19-3 I.V. 1996, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,1 mm. Corpo lucido e giallo-bruno (immaturo); 
antenne brune; zampe gialle. La reticolazione delle superficie del capo e dell'addome è 
superficiale, quella del pronoto e delle elitre è molto svanita. I tubercoletti che coprono 
la superficie dell'avancorpo sono molto superficiali. Spermateca fig. 133. 

Comparazioni. Per la forma della spermateca e per gli occhi molto sviluppati, la 
nuova specie è forma affine a P. championi Cameron, 1939, dell'India. Se ne distingue 
per la minore dimensione della spermateca che ha bulbo distale poco dilatato e parte 
prossimale breve (bulbo distale assa dilatato e parte prossimale lunga della sperma- 
teca di championi). 

Nepalota gansuensis sp. n. Figg. 135-138 

Holotypus 6 , China, Gansu, Xinlong Shan, ca. 70 Km S Lanzhou, 2225-2380 m, 
7.VIII.1994, A. Smetana leg. (MHNG). 

Paratypi: 17 es., stessa provenienza. 



944 



ROBERTO PACE 




Figo. 126-134 

Habitus, spermateca, edeago in visione laterale e ventrale e sesto urotergo libero del maschio. 
126-127: Pelioptera (s. str.) IH sp. n.; 128-132: Pelioptera (Geostibida) kowloonensis sp. n.; 
133-134: Pelioptera (Geostibida) eremita sp. n. 



ALEOCHARINAE DELLA CINA 945 

Descrizione. Lunghezza 4,1 mm. Corpo lucido e bruno con addome bruno- 
rossiccio avente gli uriti liberi 4° e 5° bruni; antenne brune con i due antennomeri basali 
giallo-rossicci; zampe gialle. La reticolazione del capo e del pronoto è netta, quella 
delle elitre è distinta. L'addome è quasi privo di reticolazione: su alcune aree ristrette è 
a maglie molto trasverse e molto svanite. I tubercoletti della superficie del capo e del 
pronoto sono molto svaniti, quelli delle elitre sono indistinti. Edeago figg. 136-137, 
spermateca fig. 138. 

Comparazioni. La nuova specie è affine a N. franzi Pace, 1987b, del Nepal. Ne 
è distinta per l' edeago più ampiamente arcuato al lato ventrale e avente armatura 
genitale interna composta da due lamine ricurve di cui una corta e altri pezzi copulatori, 
mentre in franzi è presente un'unica lamina ricurva e di dimensioni intermedie. 

Nepalota globifera sp. n. Figg. 139-140 

Holotypus 9, China, Yunnan, Ruili, ca. 700 m, 3.II.1993, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 3,4 mm. Corpo lucidissimo e nero-bruno con uriti 
liberi 4° e 5° neri; antenne nere con i due antennomeri basali nero-bruni; zampe bruno- 
rossicce. Una reticolazione svanita è presente solo sulla superficie delle elitre, sul resto 
del corpo non vi è traccia di reticolazione. La punteggiatura del capo e del pronoto è 
fine, quella delle elitre poco saliente. Spermateca fig. 140. 

Comparazioni. L'addome della nuova specie non è con evidenza ristretto 
all'indietro, né il pronoto è più fortemente ristretto in avanti che all'indietro, pertanto la 
nuova specie sembra affine a N. pernitida (Pace, 1984) della Brimania. La nuova specie 
se ne differenzia essenzialmente per la forma asimmetrica del bulbo distale della 
spermateca, con introflessione apicale brevissima e non come in pernitida con bulbo 
distale della spermateca simmetrico, con profonda e larga introflessione apicale. 

Nepalota smetanai sp. n. Figg. 141-144 

Holotypus 6, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 25. VII. 1994, 
A. Smetana leg. (MHNG). 

Paratypi: 15 es., China, Gansu, Xinlong Shan, ca. 70 Km S Lanzhou, 2225-2380 m, 
7.VIII.1994, A. Smetana leg. (MHNG). 

Descrizione. Lunghezza 3,8 mm. Corpo lucido e bruno; antenne brune con i due 
antennomeri basali giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo è 
distinta, quella del pronoto è netta, quella delle elitre è superficiale e quella 
dell'addome è assai svanita e a maglie molto trasverse. La punteggiatura del capo è 
svanita. I tubercoletti che coprono il pronoto e le elitre sono molto superficiali. Edeago 
figg. 142-143, spermateca fig. 144. 

Comparazioni. Per la forma dell' edeago e della spermateca, la nuova specie 
sembra affine a N. fessa Pace, 1987c, del Nepal. Tuttavia l'addome non è evidente- 
mente molto ristretto all'indietro e gli occhi sono più sviluppati di quelli difessa. Le 
differenze più vistose sono nell' edeago che ha uno sviluppo minore e il profilo 
ventrale, in visione laterale, largamente arcuato, ma in modo poco profondo e non 
arcuato strettamente e profondamente come in fessa. 



946 



ROBERTO PACE 




Figo. 135-140 

Habitus, edeago in visione laterale e ventrale e spermateca. 135-138: Nepalota gansuensis sp. n. 
139-140: Nepalota globifera sp. n. 



ALEOCHARINAE DELLA CINA 947 

Etimologia. La nuova specie è dedicata al suo raccoglitore, il noto studioso di 
Staphylinidae Dr. Ales Smetana di Ottawa. 

Nepalota granulosella sp. n. Figg. 145-146 

Holotypus 9, China, Beijing, Xialongmen, 1100-1500 m, 1. VII. 1993, de Rougemont 
leg. (MHNG). 

Descrizione. Lunghezza 3,2 mm. Corpo lucido e bruno con elitre e margine 
posteriore dei tre uriti basali bruno-rossicci e con uriti liberi 4° e 5° nero-bruni; antenne 
brune con i quattro antennomeri basali giallo-rossicci; zampe rossicce. La reticolazione 
del capo è molto svanita, quella del pronoto è poco superficiale, quella delle elitre è 
distinta e quella dell'addome è assente. La punteggiatura del capo è svanita. Il pronoto 
presenta due punti isolati molto superficiali e superficie coperta di distinti tubercoletti 
che sulle elitre sono svaniti. Spermateca fig. 146. 

Comparazioni. L'habitus della nuova specie è molto simile a quello di N. fessa 
Pace, 1987c, del Nepal, ma gli occhi sono lunghi quanto le tempie (e non molto più 
corti come in fessa) e la spermateca è priva di introflessione apicale del bulbo distale e 
di parte prossimale molto lunga, con bulbo prossimale molto sviluppato come in fessa. 

Nepalota chinensis sp. n. Figg. 147-150 

Holotypus â , China, Zhejiang, Tianmushan, 23.IV. 1993, de Rougemont leg. (MHNG). 

Paratypi: 49 es., stessa provenienza; 1 $ e 2 9, stessa provenienza, ma 2. IX. 1994, de 
Rougemont leg.; 18 es., China, Zhejiang Prov. Anji County, ca. 500 m, Long Wan Shan N.R., 
12. V. 1996, J. Cooter leg.; 1 S , China, Zhejiang Prov., Lin'an County, 1000 m, W Tianmu 
Shan N.R., 18.V.1996, J. Cooter leg.; 1 9, China, Yunnan, Ruili, ca. 700 m, 3.II.1993. de 
Rougemont leg.; 1 del 9, China, Shaanxi, Nanwutai, 17.IX.1995, de Rougemont leg. 

Descrizione. Lunghezza 4,6 mm. Corpo lucido e nero-bruno con elitre brune; 
antenne nero-brune con antennomero basale rossiccio; zampe rossicce. La reticolazione 
del capo è netta, quella del pronoto è nettissima, quella delle elitre è distinta e quella 
dell'addome è assente. I tubercoletti del disco del capo sono svaniti, quelli del resto 
della superficie epicraniale sono distinti, quelli del pronoto sono distinti, quelli delle 
elitre sono svaniti e quelli dell'addome sono assenti. Il primo urotergo libero del 
maschio mostra una carena mediana affilata. Edeago figg. 148-149, spermateca 
fig. 150. 

Comparazioni. La nuova specie è chiaramente distinta da N. fessa Pace, 1987c, 
del Nepal, se si osservano l'edeago e la spermateca. La parte apicale dell'edeago della 
nuova specie è a profilo ventrale pressoché rettilineo, mentre in fessa è profondamente 
arcuato. L'armatura genitale interna dell'edeago difessa è più sviluppata e più robusta 
di quella della nuova specie. La spermateca della nuova specie è molto meno svi- 
luppata, con introflessione apicale del bulbo distale profonda (appena sporgente in 
fessa). 

Alevonota sericata sp. n. Figg- 151-154 

Holotypus ó\ China, Beijing, Panshan, 8.V.1993, 8.V.1993, de Rougemont leg. 
(MHNG). 

Paratypus: 1 9 , stessa provenienza. 



948 



ROBERTO PACE 




Figo. 141-146 

Habitus, edeago in visione laterale e ventrale e spermateca. 141-144: Nepalota smetanai sp. n. 
145-146: Nepalota granulosella sp. n. 



ALEOCHARINAE DELLA CINA 



949 




Figo. 147-154 

Habitus, edeago in visione laterale e ventrale e spermateca. 147-150: Nepalota chinensis sp. n. 
151-154: Alevonota sericata sp. n. 



950 ROBERTO PACE 

Descrizione. Lunghezza 1,9 mm. Corpo debolmente lucido e nero bruno; 
antenne brune; zampe giallo-brune. L'intera superficie corporea è coperta di tubercoletti 
fittissimi e distinti, posti su un fondo non reticolato. Edeago figg. 151-152, spermateca 
fig. 154. 

Comparazioni. La nuova specie è distinta da A. chinensis Pace, 1993, della 
Cina, per la taglia corporea nettamente minore (1,9 mm invece di 2,8 mm), per gli occhi 
lunghi quanto le tempie (molto più corti delle tempie in chinensis) e per la parte 
prossimale della spermateca descrivente un semicerchio e non una spira completa come 
in chinensis. 

Gastropaga (Rougemontia) rougemonti sp. n. Figg. 155-156 

Holotypus 9 , Hong Kong, N.T., IX. 1996, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 1,7 mm. Avancorpo lucido, addome lucidissimo. 
Corpo giallo-rossiccio comprese le antenne e le zampe. La reticolazione del capo e 
delle elitre è svanita, quella del pronoto è molto svanita e quella dell'addome è assente. 
La punteggiatura del capo è distinta e composta di punti grandi, quella del pronoto è 
svanita. I tubercoletti della superficie delle elitre sono svaniti, quelli dell'addome sono 
fini e distinti. Spermateca fig. 156. 

Comparazioni. La nuova specie si distingue da G. siamensis Pace, 1984, della 
Thailandia e della Cina, per gli occhi più sviluppati, per le elitre larghe quanto il 
pronoto (elitre più larghe del pronoto in siamensis) e per la parte prossimale della 
spermateca più prolungata. 

Etimologia. La nuova specie è dedicata al suo raccoglitore, il noto studioso di 
Staphylinidae Guillaume de Rougemont di Londra. 

THAMIARAEINI 

Mimacrotona (s. str.) taiwanensis sp. n. Figg. 157-160 

Holotypus 6 . Taiwan, 95° 323, epiphytes, Ieng-Tze Yang leg. (MHNG). 
Paratypi: 60 es., stessa provenienza. 

Descrizione. Lunghezza 1,35 mm. Corpo molto convesso, lucido e giallo- 
rossiccio con capo, elitre e parte posteriore del quarto urotergo libero bruni; antenne e 
zampe gialle. La superficie corporea è coperta da tubercoletti fitti e assai superficiali, 
posti su un fondo non reticolato. Edeago figg. 158-159, spermateca fig. 160. 

Comparazioni. La nuova specie è distinta da M. orousseti Pace, 1990b, del 
Nepal, per la taglia corporea minore (1,70 in orousseti), per l'edeago poco arcuato al 
lato centrale e per il bulbo distale della spermateca poco sviluppato (molto sviluppato in 
orousseti). 

Mimacrotona (s. str.) rougemonti sp. n. Figg. 161-163 

Holotypus S , Hong Kong, Tai Po, V.1996, flight interception trap, de Rougemont leg. 
(MHNG). 

Paratypus: 1 â , stessa provenienza. 



ALEOCHARINAE DELLA CINA 95 1 

Descrizione. Lunghezza 1,7 mm. Corpo debolmente lucido e rossiccio con 
elitre bruno-rossicce; antenne e zampe rossicce. La reticolazione del capo è estrema- 
mente superficiale, quella del pronoto e delle elitre è svanita e quella dell'addome è 
assente. I tubercoletti della superficie del capo e del pronoto sono fini e distinti, quelli 
delle elitre sono svaniti. Solo il terzo urotergo libero è coperto di scultura a distinte 
squame, gli altri l'hanno confusa o ne sono privi. Edeago figg. 162-163. 

Comparazioni. La nuova specie è affine a M. orousseti Pace, 1990b, del Nepal, 
a motivo della forma dell'edeago. Ma la nuova specie ha l'undicesimo antennomero del 
maschio lungo quanto i tre precedenti antennomeri considerati insieme, mentre in 
orousseti è più corto dei tre precedenti presi insieme. L' edeago della nuova specie ha 
minore sviluppo, è più ampiamente arcuato al lato ventrale e, in visione ventrale, ha 
l'apice più largo. 

Etimologia. La nuova specie è dedicata al suo raccoglitore il collega Guillaume 
de Rougemont di Londra. 

Aidemonusa subgen. n. di Mimoxypoda Cameron, 1925 

Il nuovo sottogenere Aidemonusa del genere Mimoxypoda Cameron, 1921 si 
distingue come segue; 
1 Capo nascosto sotto il pronoto (in fig. 165 il capo è stato forzatamente 

tratto fuori da sotto il pronoto); ligula intera, ma larghissima e corta; 

spermateca piegata come la lettera C Aidemonusa subgen. n. 

(Typus subgeneris: Mimoxypoda (Aidemonusa) fasciatipennis sp. n.) 

Capo normalmente sporgente da sotto il pronoto; ligula intera, ma 

stretta e di media lunghezza; spermateca piegata secondo la lettera S. . . 

Mimoxypoda s. str. 

(Typus subgeneris: Mimoxypoda (s. str.) ruf a Cameron 1925) 

Etimologia. Il nome del nuovo sottogenere significa "Essenza vereconda", 
dato che il capo è nascosto sotto il pronoto, tale atteggiamento suggerisce timore e 
verecondia. 

Mimoxypoda (Aidemonusa) fasciatipennis sp. n. Figg- 164-166 

Holotypus 9 , Hong Kong, Tai Po, V.1996, de Rougemont leg. (MHNG). 
Paratypi: 2 9 , stessa provenienza. 

Descrizione. Lunghezza 1,6 mm. Corpo debolmente lucido e giallo-rossiccio 
con occhi bruni, con elitre brune aventi base, sutura e margine posteriore giallo-rossicci 
e con addome rossiccio avente estremità giallo-rossiccia; antenne con i tre antennomeri 
basali gialli, gli antennomeri 4° a 9° giallo-bruni e il decimo e l'undicesimo giallo- 
rossicci; zampe giallo-rossicce. La reticolazione del pronoto è molto svanita, quella sul 
resto del corpo è assente. La punteggiatura del capo è rada e distinta. I tubercoletti della 
superfìcie del pronoto sono assai poco distinti, quelli delle elitre sono fini e salienti. 
L'addome è coperto di fitta pubescenza sericea. Spermateca fig. 166. 



952 



ROBERTO PACE 



158 159 




Figo. 155-166 

Habitus, spermateca, edeago in visione laterale e ventrale, spermateca e labio con palpo labiale. 
155-156: Gastropaga (Rougemontia) rougemonti sp. n.; 157-160: Mimacrotona (s. str.) taiwa- 
nensis sp. n.; 161-163: Mimacrotona (s. str.) rougemonti sp. n.; 164-166: Mimoxypoda (Aide- 
monusa subg. n.) fasciatipennis sp. n. 



ALEOCHARINAE DELLA CINA 953 

Mimoxypoda (s. str.) chinensis sp. n. Figg. 167-169 

Holotypus S, China, Yunnan, Dali, 9.II.1993, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 1,9 mm. Corpo lucido e nero-bruno con addome nero; 
antenne brune con antennomero basale bruno rossiccio; zampe brune. La reticolazione 
delle elitre è molto svanita (le elitre forse non sono pertinenti dato che staccate dal 
corpo sono state recuperate nel liquido di conservazione in provetta), quella del capo e 
del pronoto è assente. I tubercoletti della superficie dell'avancorpo sono superficiali, 
quelli dell'addome sono salienti. Edeago figg. 168-169. 

Comparazioni. La nuova specie è simile a M. indica Cameron, 1939, dell'India, 
ma la taglia corporea è maggiore (1,6 mm in indica). L'edeago della nuova specie è 
meno ampiamente arcuato al lato ventrale, ha "crista apicalis" più lunga e ha apice, in 
visione ventrale, a punta e non ampiamente semicircolare come in indica. 



PYGOSTENINI 

Mesomegaskela gen. n. Figg. 176-184 

Diagnosi. Il nuovo genere è affine al genere Pygostenus Kraatz, 1858b, unica- 
mente diffuso nella regione etiopica. Con questo genere, il nuovo condivide la forma 
fusiforme del corpo, ma le antenne non sono corte e fusiformi. 

Descrizione. Occhi sviluppati; le procoxe e le mesocoxe sono molto larghe; 
femori molto dilatati, particolarmente i pro-mesofemori; tibie cortissime; formula 
tarsale 4-5-5 (figg. 179-189); palpi labiali di tre articoli (fig. 183); ligula larghissima, 
divisa in due lobi; paraglosse molto prominenti in avanti; palpi mascellari di quattro 
articoli (fig. 184); meso-metasterno come da fig. 178; nono segmento addominale fig. 
182; parte prossimale della spermateca avvolta in quattro spire (al massimo con una 
stretta spira in Pygostenus). 

Typus generis: Mesomegaskela adesi sp. n. 

Etimologia. Il nome del nuovo genere significa "Coxe intermedie grandi". 



Mesomegaskela adesi sp. n. Figg. 176-184 

Holotypus 9, Hong Kong, Kadoorie Agricultural Research Centre, N.T., IX. 1990, 
G. Ades leg. (MHNG). 

Descrizione. Lunghezza 5 mm. Corpo lucido e bruno-rossiccio con metà 
posteriore delle elitre bruna e con addome gradualmente giallo rossiccio verso l'apice; 
antenne e zampe bruno-rossicce. L'avancorpo è privo di reticolazione ed è coperto da 
pubescenza cortissima e fittissima. Le elitre presentano una fila di lunghe setole al 
margine posteriore. La pubescenza dell'addome è coperto da una pubescenza aderente, 
posta su un fondo non reticolato. Spermateca fig. 172. 

Etimologia. La nuova specie è dedicata al suo raccoglitore Garry Ades noto 
zoologo inglese. 



954 



ROBERTO PACE 




Figo. 167-175 

Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del maschio, labio con palpo 
labiale e spermateca. 167-169: Mimoxypoda (s. str.) chinensis sp. n.; 170-175: Medeterusa 
minima Pace. 



ALEOCHARINAE DELLA CINA 



955 





Figo. 176-182 

Habitus, spermateca, pro-mesometasterno, zampa anteriore (179), media (180) e posteriore 
(181) e nono segmento addominele. 176-182: Mesomegaskela adesi gen. n., sp. n. 



956 ROBERTO PACE 

Doryloxenus hongkongensis sp. n. Figg. 185-188 

Holotypus 6, Hong Kong, Kadoorie Farm, flight interception trap, 15.IX.1996, de 
Rougemont leg. (MHNG). 

Paratypus: 1 9, stessa provenienza, ma 26.V.1996, de Rougemont leg. 

Descrizione. Lunghezza 1,7 mm. Corpo lucidissimo e bruno-rossiccio con elitre 
brune; antenne brune; zampe non visibili da sopra. Sul corpo non vi è traccia di 
reticolazione. La punteggiatura del capo è assai rada e debole, quella del pronoto e delle 
elitre è fine e distinta. Una corta pubescenza è presente solo sulla metà posteriore di 
ciascun urotergo. Edeago figg. 186-187, spermateca fig. 185. 

Comparazioni. La nuova specie è la prima specie asiatica del genere 
Doryloxenus Wasmann, 1898, finora noto solo della regione etiopica. Una specie che 
ha la parte prossimale della spermateca avvolta in numerose spire come quella della 
nuova specie, non è nota. Solo D. castaneus Cameron, 1938, diffuso in Tanzania, 
Uganda, Rodesia, Angola e Ghana, ha parte prossimale avvolta in spire simili a quelle 
della nuova specie, ma esse sono solo due. 

Doryloxenus rougemonti sp. n. Figg. 189-190 

Holotypus 9 , Hong Kong, Kadoorie Farm, 15. IX. 1996, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,1 mm. Corpo lucidissimo, non reticolato e rossiccio 
con estremità addominale giallo-rossiccia; antenne bruno-rossicce; zampe non visibili 
dall'alto. La punteggiatura del capo e del pronoto è molto rada ed estremamente super- 
ficiale, quella delle elitre è meno superficiale. Le setole posteriori marginali di ciascun 
urotergo sono erette. I tubercoli del margine posteriore dei quattro uroterghi basali sono 
ben salienti. Spermateca fig. 190. 

Comparazioni. La nuova specie è la seconda specie asiatica del genere 
Doryloxenus Wasmann, 1898, dopo quella descritta sopra. Essa si distingue da tutte le 
specie africane per la parte prossimale della spermateca avvolta a fitta matassa e dalla 
specie sopra descritta, oltre che per la spermateca di taglia maggiore con bulbo distale 
sferico, per la taglia corporea maggiore. 

Etimologia. La nuova specie è dedicata al suo raccoglitore, il noto studioso di 
Staphylinidae Guillaume de Rougemont. 

Odontoxenus rougemonti sp. n. Figg. 191-194 

Holotypus 6, Hong Kong, Kadoorie Farm, flight interception trap, 15.IX.1996, de 
Rougemont leg. (MHNG). 

Paratypus: 1 9 , stessa provenienza. 

Descrizione. Lunghezza 1,9 mm. Corpo lucido e giallo-bruno con elitre brune e 
addome giallo rossiccio; antenne bruno-rossicce; zampe rossicce con tibie bruno- 
rossicce. La punteggiatura del capo è doppia costituita da radi punti superficiali più 
grandi e fitti puntini fini, quella del pronoto è rada e netta come quella delle elitre. Gli 
uroterghi liberi 3° a 5° mostrano delle setoline fitte allineate trasversalmente. Sul corpo 
non vi è traccia di microscultura reticolare. Edeago figg. 192-193, nono segmento 
addominale fig. 194. 



ALEOCHARINAE DELLA CINA 



957 




Figo. 183-188 

Labio con palpo labiale, maxilla con palpo mascellare, spermateca, edeago in visione laterale e 
ventrale e habitus. 183-184: Mesomegaskela adesi gen. n., sp. n.; 185-188: Doryloxenus hong- 
kongensis sp. n. 



958 



ROBERTO PACE 




Figo. 189-194 

Habitus, spermateca, edeago in visione laterale e ventrale e nono segmento addominale, li 
190: Doryloxenus rougemonti sp. n.; 191-194: Odontoxenus rougemonti sp. n. 



ALEOCHARINAE DELLA CINA 959 

Comparazioni. La nuova specie è distinta da O. krishnai Kistner & Jacobson, 
1975, della Birmania, la specie geograficamente più vicina alla nuova, per avere la 
punteggiatura del pronoto e delle elitre netta. Non è noto il maschio di krishnai. 

Etimologia. La nuova specie è dedicata al suo raccoglitore studioso di Staphy- 
linidae Guillaume de Rougemont di Londra. 

Odontoxenus hongkongensis sp. n. Figg. 195-196 

Holotypus ?, Hong Kong, Kadoorie Farm, flight interception trap, 15. IX. 1996, de 
Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,1 mm. Corpo lucido, senza traccia di reticolazione, 
bruno-rossiccio con pronoto e addome giallo-rossicci; antenne rossicce con undicesimo 
antennomero giallo-rossiccio; zampe rossicce con tarsi gialli. La punteggiatura del capo 
è distinta e fitta, quella del pronoto è estremamente svanita, quella delle elitre è doppia 
composta da punti grandi assai radi e svaniti e puntini fitti e fini, quella dell'addome è 
fine e fitta. Spermateca fig. 196. 

Comparazioni. Non esistono specie note con spermateca avvolta in più spire, 
come la spermateca della nuova specie. La doppia punteggiatura delle elitre, oltre che al 
pronoto poco trasverso, distinguono la nuova specie da 0. rougemonti sp. n. sopra 
descritta. 

Cephaplakoxena gen. n. Figg- 197-208 

Diagnosi. Il nuovo genere è vicino al genere Aenictoxenus Seevers, 1953, delle 
Filippine, per la forma delle antenne. Se ne distingue per avere le antenne composte di 
9 antennomeri (invece di 8 come in Aenictoxenus), per il capo compresso e per il 
pronoto molto poco trasverso e quasi emisferico (pronoto fortemente trasverso in 
Aenictoxenus). 

Descrizione. Pronoto convesso; addome appiattito; capo nascosto sotto il 
pronoto, fortemente compresso e concavo (figg. 200-201) al fine di rinserrare il robusto 
primo antennomero basale; antenne di 9 antennomeri, in stato di riposo sporgenti dal 
contorno del pronoto mediante il solo antennomero terminale che è lunghissimo; palpi 
labiali di tre articoli (fig. 199); ligula larga e divisa; paraglosse appena sporgenti; palpi 
mascellari di quattro articoli (fig. 205); lobo esterno delle maxille molto più lungo 
dell'interno; pronoto molto sviluppato, convesso e senza angoli posteriori distinti; elitre 
ed ali non presenti, forse perdute in fase di raccolta: l'esemplare essendo stato raccolto 
al volo deve essere provvisto di ali ed elitre; prosterno non carenato sulla linea 
mediana; procoxe poco convesse (fig. 206); mesocoxe medie grandi e ovali, contigue 
fra loro dato che il processo mesosternale è appena accennato da una sinuosità mediana 
volta all' indietro (fig. 208); metasterno non visibile perché coperto dalle metacoxe che 
sono molto trasverse; formula tarsale 4-5-(5) (la parentesi indica che le zampe 
posteriori sono andate perdute forse nella fase di raccolta dell'esemplare; è indicato 5 
anche per i tarsi posteriori perché nelle Aleocharinae tutte le specie che presentano tarsi 
medi di 5 articoli, sempre presentano i posteriori di 5); piastra apicale dei parameri 
minuscola (fig. 202). 



960 



ROBERTO PACE 




0,05 mm 



Figo. 195-202 

Habitus, spermateca, antenna, labio con palpo labiale, capo in visione frontale everticale e 
piastra apicale di paramero. 195-196: Odontoxenus hongkongensis sp. n.; 197-202: Cepha- 
plakoxena rougemonti gen. n., sp. n. 



ALEOCHARINAE DELLA CINA 961 

Typus generis: Cephaplakoxena rougemonti sp. n. 

Etimologia. Il nome del nuovo genere significa "Ospite con capo schiacciato". 
Ospite di formicai. Il genere grammaticale è femminile. 

Cephaplakoxena rougemonti sp. n. Figg. 197-208 

Holotypus S, Hong Kong, Kadoorie Farm, flight interception trap, 12.X.1996, de 
Rougemont leg. (MHNG). 

Descrizione. Lunghezza 2,8 mm. Corpo lucidissimo e giallo-bruno con capo 
bruno, con pronoto traslucido avente lati di un giallo-rossiccio sporco e con estremità 
addominale giallo-rossiccia; antenne brune con i due antennomeri basali giallo-rossicci 
ed apice del nono antennomero (l'ultimo) giallo-bruno; zampe giallo-rossicce, non 
visibili dall'alto, tanto sono corte. La reticolazione del capo è svanita, quella del pro- 
noto è assente, le elitre sono andate perdute, quella dell'addome è distinta e composta di 
maglie molto trasverse, sul quinto libero la reticolazione è meno trasversa. La pun- 
teggiatura del pronoto è sparsa, fine e superficiale. Il quinto urotergo libero ha un 
profondo solco a ciascun lato. Edeago figg. 203-204. 

MYRMEDONIINI 

Chaetosogonocephus chinensis sp. n. Figg. 209-210 

Holotypus 9 , China, Zhejiang, Tianmushan, 2.IX.1994, de Rougemont leg. (MHNG). 

Descrizione. Lunghezza 3,5 mm. Corpo lucidissimo e rossiccio con elitre 
soffuse di bruno; antenne rossicce con i tre antennomeri basali giallo-rossicci; zampe 
rossicce. Sul corpo non vi è traccia di reticolazione. La punteggiatura del capo è dis- 
tinta, irregolarmente distribuita e composta di punti grandi. Il pronoto e le elitre sono 
coperti di tubercoletti salienti, essi sono assenti sulla fascia mediana del pronoto. La 
metà posteriore del quinto urotergo libero della femmina e il sesto sono coperti di 
profonde strie longitudinali. Spermateca fig. 210. 

Comparazioni. La nuova specie è simile a C. rougemonti Pace, 1986 della 
Malaysia. Ne è distinta per la taglia corporea maggiore, le tempie nettamente divergenti 
all'indietro (appena divergenti in rougemonti), per le elitre meno larghe, per l'addome 
più appuntito, per la stilatura longitudinale della metà posteriore del quinto urotergo 
libero della femmina (assente nella femmina di rougemonti) e per la notevole lunghezza 
della parte prossimale della spermateca, descrivente una spira (parte prossimale della 
spermateca di rougemonti cortissima). 

Tetrabothrus rougemonti sp. n. Figg. 21 1-214 

Holotypus S , Hong Kong, Tai Po, flight interception trap, V.1996, de Rougemont leg. 
(MHNG). 

Paratypus: 1 9 , Hong Kong, Ng Fai Tin, at light, 22.VIII.1996, G.T. Reels leg. 

Descrizione. Lunghezza 5,0 mm. Corpo lucido, senza reticolazione e rossiccio, 
antenne comprese; zampe rossicce con femori gialli aventi l'estremità distale bruna. La 
punteggiatura dell'avancorpo è quasi indistinta. Gli uroterghi sono privi di punteggia- 
tura e di setole, tranne alcune isolate. Edeago figg. 211-212, spermateca fig. 214. 



962 



ROBERTO PACE 




203 



204 



206 




Figo. 203-208 

Edeago in visione laterale e ventrale, maxilla con palpo mascellare, zampa anteriore (206) e 
media (207) e meso-metasterno. 203-208: Cephaplakoxena rougemonti gen. n., sp. n. 



ALEOCHARINAE DELLA CINA 



963 




FiGG. 209-213 

Habitus, spermateca ed edeago in visione laterale e ventrale. 209-210: Chaetosogonocephus 
chinensis sp. n.; 21 1-213: Tetrabothrus rougemonti sp. n. 



964 ROBERTO PACE 

Comparazioni. La nuova specie è simile a T. indicus Cameron, 1939, dell'India, 
ma ha antenne chiaramente fusiformi (non fusiformi in indicus), elitre unicolori 
(rossicce con metà posteriore brune in indicus), edeago nettamente meno sviluppato, 
con apice, in visione ventrale, a margini preapicali nettamente sinuati (quasi rettilinei in 
indicus). La nuova specie è pure distinta da T. laticornis (Wasmann, 1896), per avere la 
spermateca più semplice, con parte prossimale avvolta in tre o quattro spire e non 
avvolta in numerosissime spire (circa 15) come in laticornis. 

Amaurodera yunnanensis sp. n. Figg. 215-217 

Holotypus ó\ China, Yunnan, Ruili, ca. 700 m, 3.II.1993, de Rougemont leg. (MHNG). 
Paratypi: 3 9, stessa provenienza. 

Descrizione. Lunghezza 3,7 mm. Corpo lucido, tranne il pronoto molto opaco 
d'aspetto vellutato, tranne la parte anteriore che è lucida. Corpo rossiccio con elitre 
bruno-rossicce e con addome giallo avente la metà posteriore del terzo urotergo libero, 
il quarto e macchie ai lati dei due uriti basali bruni, quinto urite libero bruno-rossiccio; 
antenne rossicce con antennomero basale giallo; zampe anteriori bruno-rossicce con 
tarsi gialli, medie e posteriori brune con base dei femori e i tarsi gialli ed estremità 
distale delle tibie rossiccia. Il capo presenta punteggiatura fine e molto svanita e un 
debole e largo solco mediano. Il pronoto ha un solco mediano profondo, nel fondo di 
una larga depressione. I tubercoletti della superficie delle elitre sono distinti. Edeago 
figg. 216-217. 

Comparazioni. In base alla forma dell'edeago, la nuova specie è affine ad 
A. veluticollis (Motschulsky, 1858), dell'India, ma è attera, perciò ha elitre più corte e 
più strette. Il profilo ventrale dell'edeago della nuova specie è bisinuato in modo molto 
più accentuato rispetto il corrispondente profilo ventrale dell'edeago di veluticollis e le 
espansioni preapicali laterali dello stesso edeago, sono più ampie nella nuova specie 
che in veluticollis. 

Drusilla zhejiangensis sp. n. Figg. 218-220 

Holotypus ó\ China, Zhejiang, Tianmushan, 29.IV. 1993, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 5,0 mm. Avancorpo debolmente lucido, addome 
lucido. Avancorpo nero pece, addome bruno con uriti liberi 4° e 5° neri; antenne 
rossicce con i cinque antennomeri basali bruno-rossicci; zampe rossicce con metà 
distale dei femori medi e posteriori nero-brune e la metà basale gialla. La reticolazione 
del capo è distinta, quella del pronoto e delle elitre è svanita. La punteggiatura del capo 
è distinta e fitta. I tubercoletti della superficie del pronoto e delle elitre sono fitti e 
superficiali. Il pronoto presenta un fine solco mediano profondo. Edeago figg. 218-220. 
Comparazioni. La nuova specie differisce da D. obliqua (Bernhauer), della 
Birmania e dell'India, per il capo nettamente più stretto del pronoto (appena più stretto 
del pronoto in obliqua), per la minore taglia dell'edeago che presenta una notevole 
gibbosità preapicale ventrale (poco marcata in obliqua), per l'assenza di "crista api- 
calis" (presente in obliqua) e per lo scarso sviluppo dell'armatura genitale interna 
(sviluppo molto marcato in obliqua). 



ALEOCHARINAE DELLA CINA 



965 




FiGG. 214-218 

Spermateca, habitus ed edeago in visione laterale e ventrale. 214: Tetrabothrus rougemonti sp. 
n.; 215-217: Amaurodera yunnanensis sp. n.; 218: Drusilla zhejiangensis sp. n. 



966 ROBERTO PACE 

Drusilla kadooriorum sp. n. Figg. 221-224 

Holotypus â; Hong Kong, N.T., Kadoorie Agricultural Research Centre, Malaise trap, 
Vili. 1991, G. Ades leg. (MHNG). 

Paratypi: 2 es., stessa provenienza. 

Descrizione. Lunghezza 4,1 mm. Corpo lucidissimo e giallo-rossiccio con capo 
e uriti liberi 3°, 4° e base del 5° bruni, tranne il margine posteriore degli uroterghi liberi 
3° e 4° che è rossiccio; antenne brune con i tre antennomeri basali giallo-rossicci; 
zampe giallo-rossicce. La superficie del corpo non è reticolata. La punteggiatura del 
capo è distinta e diradata sul disco, quella delle elitre è ben conformata. Tubercoletti 
distinti sono addensati in avanti e all' indietro del pronoto che ha un netto solco mediano 
e un'ampia depressione laterale a ciascun lato. La punteggiatura dell'addome è fine. Gli 
uroterghi liberi 2° e 3° hanno un'ampia bozza basale mediana, il 4° e il 5° hanno una 
concavità mediana. Una depressione laterale obliqua sta sul quinto urotergo libero. 
Edeago figg. 222-223, spermateca fig. 224. 

Comparazioni. In base alla forma della spermateca, la nuova specie sembra 
tassonomicamente vicina a D. assamensìs (Cameron, 1939), della Birmania e dell' 
India, ma l'introflessione apicale del bulbo distale della spermateca della nuova specie 
è profonda e robusta ed esile in assamensis. Ma è la parte apicale stretta e lunga 
dell' edeago che distingue nettamente la nuova specie da assamensis che ha edeago 
larghissimo in visione laterale e l'apice a forma di larga ogiva, in visione ventrale. 

Etimologia. La nuova specie è dedicata ai fratelli Kadoorie, noti filantropi di 
Hong Kong, nel cui centro agricolo di ricerca è stata raccolta la nuova specie. 

Drusilla gibberella sp. n. Figg. 225-228 

Holotypus 9 , Hong Kong, Kadoorie Agricultural Research Centre, flight interception 
trap, 19-3 I.V. 1996, de Rougemont leg. (MHNG). 

Paratypus: 1 S, Hong Kong, Tai Po, VII. 1996, de Rougemont. 

Descrizione. Lunghezza 4,2 mm. Corpo lucido e giallo-bruno con capo nero e 
uriti liberi 3°, 4° e 5° bruni; antenne brune con il secondo antennomero e la base del 
terzo basali giallo-rossicci; zampe giallo-brune. La reticolazione del capo e del pronoto 
è distinta, quella delle elitre e del pronoto è svanita. La punteggiatura del capo è fine, 
quella del pronoto è distinta. Tubercoletti salienti coprono la superficie delle elitre. Il 
pronoto presenta una larga depressione mediana posteriore e a ciascun lato di essa 
un'altra larga depressione della superficie. Edeago figg. 226-227, spermateca fig. 228. 

Comparazioni. In base alla forma dell' edeago e della spermateca, come dei 
caratteri dell'esoscheletro, la nuova specie è tassonomicamente vicina a D. aerea 
(Cameron, 1933), del Borneo. I penultimi antennomeri della nuova specie sono molto 
trasversi, mentre quelli di aerea sono appena trasversi. Il quinto urotergo libero del 
maschio della nuova specie non ha un rilevante tubercolo, presente in due file trasverse 
in aerea. L' edeago della nuova specie ha dimensioni minori, ha l'appendice preapicale 
ventrale poco saliente (ben distinta in aerea) e situata più vicina alla "crista apicalis" 
che all'apice dell' edeago stesso (situata più vicina all'apice che alla "crista apicalis" in 
aerea). 



ALEOCHARINAE DELLA CINA 



967 




Figo. 219-224 

Habitus, edeago in visione laterale e ventrale e spermateca. 219-220: Drusilla zhejiangensis 
sp. n.; 221-224: Drusilla kadooriana sp. n. 



968 ROBERTO PACE 

Diplopleurus cooteri sp. n. Figg. 229-230 

Holotypus 9, China, Zhejiang Prov., Lin'an County, W Tianmu Shan N.R., 16-22. V. 
1996, J. Cooter leg. (MHNG). 

Descrizione. Lunghezza 4,2 mm. Corpo lucido e nero con margine posteriore 
degli uroterghi liberi 2° a 5° bruno-rossiccio come l'apice addominale; antenne nero- 
brune con i tre antennomeri basali rossicci e l'undicesimo bruno-rossiccio; zampe 
giallo-rossicce. La superficie del corpo non è reticolata. La punteggiatura del capo è 
profonda e assente sulla fascia mediana, quella del pronoto è pure profonda, ma a punti 
contigui, quella delle elitre è come quella del pronoto e con punti fusi tra loro ai lati 
esterni. La punteggiatura degli uroterghi è netta e assente in alcuni tratti della super- 
ficie. Il disco del capo è lievemente concavo. Il pronoto presenta una concavità mediana 
posteriore, un'impressione a ciascun lato e uno spigolo basale senza punteggiatura. Il 
margine posteriore del sesto urosterno della femmina è incavato a metà. Spermateca 
fig. 4,2 mm. 

Comparazioni. Per la presenza della plica basale del pronoto che è fittamente e 
profondamente punteggiato, la nuova specie è attribuibile al genere Diplopleurus 
Bernhauer, 1915b, finora noto solo della regione etiopica. Non è nota ancora una specie 
che presenti occhi minori delle tempie (occhi molto più lunghi delle tempie nelle specie 
africane), né parte prossimale della spermateca così ampiamente avvolta a matassa. 

Zyras (s. str.) fratrumkadooriorum sp. n. Figg- 231-234 

Holotypus 6, Hong Kong, Kadoorie Farm, Malaise trap, VI. 1991, G. Ades leg. 
(MHNG). 

Paratypi: 1 6 e 1 9, stessa provenienza, ma VII. 1992, G. Ades leg.; 8 es., stessa 
provenienza, ma V.1996, VIII. 1996, IX. 1996, de Rougemont leg. 

Descrizione. Lunghezza 4,0 mm. Corpo lucidissimo e non reticolato. Capo e 
pronoto neri, elitre giallo-rossicce con angolo posteriore esterno largamente bruno, 
addome giallo-rossiccio con una macchia bruna a metà degli uroterghi liberi 2°, 3° e 4°, 
con gli uroterghi liberi 5° e 6° bruni e con una macchia pure bruna ai lati degli uroterghi 
liberi 2°, 3° e 4°; antenne brune con i tre antennomeri basali rossicci; zampe gialle. La 
punteggiatura del capo è netta e assai rada, quella del pronoto è distinta e irregolar- 
mente distribuita e quella delle elitre è profonda e assente lungo il margine posteriore. Il 
pronoto ha una profonda fossetta mediana posteriore. Spermateca fig. 232, edeago 
figg. 233-234. 

Comparazioni. In base alla forma dell' edeago e per alcuni caratteri dell'eso- 
scheletro, la nuova specie sembra simile a Z. chinkiangensis Bernhauer, 1939a, pure 
della Cina. Ma i tre antennomeri apicali non hanno il colore giallo paglierino come 
quelli di chinkiangensis e l'antennomero terminale del maschio della nuova specie è 
lunghissimo, mentre è corto in chinkiangensis. Inoltre l' edeago della nuova specie ha 
dimensione di una metà inferiore e la spina basale dell'armatura genitale interna 
dell' edeago è lunga nella nuova specie e breve in chinkiangensis. 

Etimologia. La nuova specie è dedicata ai fratelli Kadoorie, insigni benefattori 
di Hong Kong, nella cui azienda agricola è stata raccolta la nuova specie. 



ALEOCHARINAE DELLA CINA 



969 




225 





Figo. 225-229 

Habitus, edeago in visione laterale e ventrale e spermateca. 225-228: Drusilla gibberella sp. n. 
229: Diplopleurus cooteri sp. n. 



970 



ROBERTO PACE 




Figo. 230-234 

Spermateca, habitus ed edeago in visione laterale e ventrale. 230: Diplopleurus cooteri sp. n. 
231-234: Zyras (s. str '.) fratrumkadoorionan sp. n. 



ALEOCHARINAE DELLA CINA 97 j 

Zyras (s. str.) notaticornis sp. n. Figg. 235-238 

Holotypus S , Hong Kong, Kadoorie Agricultural Research Centre, flight interception 
trap, 19-3 I.V. 1996, de Rougemont leg. (MHNG). 

Paratypi: 4 es., stessa provenienza, ma III. 1996, VI. 1996, de Rougemont leg.; 1 3, 
China, Zhejiang Prov., Anji County, ca. 480 m, Long Wan Shan N.R., 13.V.1996, J. Cooter leg. 

Descrizione. Lunghezza 5,4 mm. Corpo lucido e non reticolato. Capo e pronoto 
nero-bruni, elitre brune con omeri e sutura rossicci, addome bruno con margine pos- 
teriore e base degli uroterghi giallo-rossicci; antenne nere con il secondo antennomero 
basale e l'undicesimo giallo-rossicci; zampe gialle. La punteggiatura dell'avancorpo è 
profonda, assente su una fascia mediana del capo e del pronoto, irregolarmente distri- 
buita su quest'ultimo e radi sulla metà posteriore delle elitre. Spermateca fig. 236, 
edeago figg. 237-238. 

Comparazioni. La nuova specie è simile a Z. chinkiangensis Bernhauer, 1939a, 
pure della Cina, ma l' antennomero apicale è giallo rossiccio e lungo nella nuova specie 
e non giallo-paglierino come i due precedenti e corto come in chinkiangensis. L' edeago 
della nuova specie è minore e più ampiamente arcuato al lato ventrale, con armatura 
genitale interna robusta (esile in chinkiangensis). 

Zyras (s. str.) shaanxiensis sp. n. Figg- 239-241 

Holotypus 3, China, Shaanxi, Nanwutai, 17.IX.1995, de Rougemont leg. (MHNG). 
Descrizione. Lunghezza 6,4 mm. Corpo lucido e non reticolato. Capo e pronoto 
neri, elitre giallo-rossicce con angolo posteriore esterno largamente bruno, addome 
bruno avente i lati e il margine posteriore degli uroterghi giallo-rossicci; antenne brune 
con i tre antennomeri basali giallo-rossicci e l'undicesimo rossiccio; zampe gialle. La 
punteggiatura dell'avancorpo è netta e profonda, assente sulla fascia mediana del capo e 
del pronoto e lungo il margine posteriore di quest'ultimo che presenta una profonda 
fossetta mediana posteriore e due punti discali robusti. Edeago figg. 240-241. 

Comparazioni. La nuova specie è ben distinta da Z. chinkiangensis Bernhauer. 
1939a, pure della Cina, per avere l'undicesimo antennomero rossiccio (e non i tre 
antennomeri apicali di colore giallo paglierino come in chinkiangensis), per l'edeago 
meno sviluppato e per la sua robusta armatura genitale interna (esile armatura in 
chinkiangensis). 

Zyras (Rynchodonia) longwangmontis sp. n. Figg. 242-244 

Holotypus 3 , China, Zhejiang Prov., Anji County, ca. 400 m, 13.V.1996, J. Cooter leg. 
(MHNG). 

Descrizione. Lunghezza 9,0 mm. Capo opaco, resto del corpo lucido. Capo ed 
elitre bruni, pronoto rossiccio, addome bruno-rossiccio con base rossiccia; antenne 
rossicce con margine distale di ciascun antennomero bruno; zampe rossicce. La 
reticolazione del capo è vigorosa e d'aspetto di velluto, ai suoi lati e all' indietro è 
svanita, quella del pronoto è estremamente svanita, quella delle elitre è superficiale e 
quella dell'addome è distinta. La punteggiatura del capo è distinta, quelle del pronoto 
e delle elitre è netta, fitta e profonda e quella dell'addome è doppia: a punti robusti e a 



972 



ROBERTO PACE 




Figo. 235-241 

Habitus, spermateca ed edeago in visione laterale e ventrale. 235-238: Zyras (s. str.) notati- 
cornis sp. n.; 239-241: Zyras (s. str.) shaanxiensis sp. n. 



ALEOCHARINAE DELLA CINA 973 

punti fini. Il rilievo mediano posteriore del terzo urotergo libero del maschio è privo 
di punteggiatura. Edeago figg. 243-244. 

Comparazioni. L' edeago della nuova specie è simile a quello di Z. nepalensis 
Pace, 1992, del Nepal, ma l'estremità distale dell'armatura genitale sporgente 
dell 'edeago ha forma di due lame ricurve, di cui una più stretta, mentre in nepalensis 
l'estremità distale di detta armatura genitale ha forma di due larghi lobi. Inoltre il 
secondo urotergo libero del maschio presenta due spine larghe e corte in nepalensis e 
lunghe nella nuova specie. 

Zyras (Diaulaconia) kadoorianus sp. n. Figg. 245-248 

Holotypus 3 , Hong Kong, N.T., Kadoorie Agricultural Research Centre, Malaise trap, 
Vili. 1991, G. Ades leg. (MHNG). 

Paratypi: 1 del 9, stessa provenienza, ma VI. 1991, G. Ades leg.; 1 9, Hong Kong, 
Shing Mun, 8.VII.1996, G. Reels leg.; 1 9, Hong Kong, N.T., Shek Kong, 21. VI. 1990, 
G. Ades leg. 

Descrizione. Lunghezza 10,0 mm. Corpo lucidissimo e giallo rossiccio, 
antenne comprese; zampe gialle. La reticolazione del capo è netta, quella del pronoto 
e dell'addome è assente, quella delle elitre è obliqua, molto trasversa e distinta. La 
punteggiatura del capo è distinta, ombelicata e assente sulla fascia mediana, quella del 
pronoto è netta e assente sulla linea mediana e a ciascun lato di essa, quella delle elitre 
è netta e profonda e quella dell'addome è fine. Il quarto urotergo libero del maschio 
presenta una concavità mediana, il quinto ha un tubercolo mediano spianato su un 
appiattimento triangolare della superficie che è distintamente reticolata. Edeago 
figg. 245-246, spermateca fig. 247. 

Comparazioni. La nuova specie è simile a Z. orientalis Bernhauer, 1929, pure 
di Hong Kong. Ne è distinta perché non ha una fila di punti profondi a ciascun lato 
della linea mediana del pronoto, come in orientalis e perché il margine posteriore del 
terzo urotergo libero del maschio è sinuoso e non profondamente incavato a metà 
come in orientalis (fig. 249). 

Etimologia. La nuova specie è dedicata ai fratelli Kadoorie noti filantropi di 
Hong Kong, nel cui centro agricolo è stata raccolta la nuova specie. 

Zyras (Pella) reelsi sp. n. Figg- 253-256 

Holotypus 6, Hong Kong, Shek Kwu Chao, 3.VII.1996. M.V. light, G.T. Reels leg. 
(MHNG). 

Paratypi: 12 es., stessa provenienza; 34 es., Beaufort Island, I.V. 1996, M.V. light, G.T. 
Reels leg. 

Descrizione. Lunghezza 7,0 mm. Avancorpo lucido, addome lucidissimo. 
Capo e addome nero-bruni, pronoto, elitre e lati dell'addome rossicci; antenne brune 
con i tre antennomeri basali rossicci; zampe rossicce con femori giallo-rossicci. La 
reticolazione è vigorosa sul disco del capo e svanita sul resto della sua superficie, 
quella del pronoto è netta, quella delle elitre è svanita e quella dell'addome è assente. 
Netta è la punteggiatura sull'intera superficie del corpo. Il pronoto presenta una 
depressione a ciascun lato. Il quinto urotergo libero del maschio ha una saliente 



974 



ROBERTO PACE 




Figo. 242-248 

Habitus, edeago in visione laterale e ventrale e spermateca. 242-244: Zyras (Rynchodonia) 
longwangmontis sp. n.; 245-248: Zyras (Diaulaconia) kadoorianus sp. n. 



ALEOCHARINAE DELLA CINA 



975 




Figo. 249-256 

Habitus, edeago in visione laterale e ventrale e spermateca. 249-252: Zyras (Diaulaconia) 
orientalis Bernhauer; 253-256: Zyras (Pelici) reelsi sp. n. 



976 ROBERTO PACE 

carena mediana, il sesto presenta due tubercoli mediani. Edeago figg. 253-254, sper- 
mateca fig. 255. 

Comparazioni. La nuova specie è distinta da Z coloratus Cameron, 1939, 
dell'India, di cui condivide parte del colore del corpo, per la maggiore taglia corporea 
(4,0 mm in coloratus), per gli occhi più lunghi delle tempie (occhi lunghi quanto le 
tempie in coloratus), per la presenza di netta reticolazione del pronoto (reticolazione 
assente sul pronoto di coloratus) e per la presenza di una spina mediana al margine 
posteriore del primo urotergo libero del maschio e di un'altra a ciascun lato posta sul 
margine laterale del primo urotergo libero (spine assenti in coloratus). 

Etimologia. La nuova specie è dedicata al suo raccoglitore, lo zoologo inglese 
Graham Reels. 



Zyras (Pella) micropterus sp. n. Figg- 257-261 

Holotypus 6 , China, Beijing, Xiaolongmen, 1 100-1500 m. 1. VII. 1993, de Rougemont 
leg. (MHNG). 

Paratypi: 15 es., stessa provenienza. 

Descrizione. Lunghezza 4,2 mm. Corpo lucido e nero-bruno con margine 
posteriore dei due uroterghi basali rossiccio; antenne rossicce con antennomero basale 
bruno; zampe bruno-rossicce con femori bruni, tarsi anteriori giallo-rossicci. La 
reticolazione del capo e del pronoto è netta, quella delle elitre è distinta e quella 
dell'addome è trasversa. La punteggiatura del capo è fitta e distinta. I tubercoletti della 
superficie del pronoto e delle elitre sono fitti e distinti, quelli dell'addome sono salienti 
e radi. Edeago figg. 258-259, spermateca fig. 260, sesto urotergo libero del maschio 
fig. 261. 

Comparazioni. La nuova specie è distinta da Z. ceylonicus Cameron, 1939, 
dello Sri Lanka, per gli occhi più corti delle tempie (occhi lunghi quanto le tempie in 
ceylonicus), per l'assenza di una fossetta mediana basale del pronoto (presente sul 
pronoto di ceylonicus) e per le elitre più corte del pronoto (elitre lunghe quanto il 
pronoto in ceylonicus). 

Zyras (Pella) beijingorum sp. n. Figg. 262-263 

Holotypus 9. China. Beijing. Xiaolongmen, 1100-1500 m, 1. VII. 1993, de Rougemont 
leg. (MHNG). 

Descrizione. Lunghezza 4,8 mm. Corpo lucido. Capo e pronoto bruni, elitre 
giallo-brune con macchia laterale bruna, addome bruno con margine posteriore dei due 
uroterghi basali largamente giallo-bruni, degli uroterghi terzo e quarto strettamente 
rossiccio, apice addominale bruno-rossiccio; antenne bruno-rossicce con i tre antenno- 
meri basali rossicci; zampe rossicce. La reticolazione del capo e dell'addome è distinta, 
quella addominale è a maglie molto trasverse, quella del pronoto è superficiale e quella 
delle elitre è netta. Il capo presenta una fossetta trasversa tra le antenne e i tubercoletti 
che coprono la superficie salienti. Il pronoto presenta una superficie coperta di 
tubercoletti distinti, le elitre li ha quasi indistinti. Spermateca fig. 263. 



ALEOCHARINAE DELLA CINA 



977 




Figo. 257-263 

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio. 
257-261: Zyras (Pella) microptems sp. n.; 262-263: Zyras {Fella) beijingorum sp. n. 



978 ROBERTO PACE 

Comparazioni. La nuova specie è distinta da Z. coloratus Cameron, 1939, 
dell'India, per le elitre giallo-brune con macchia laterale bruna (elitre bruno-rossicce 
in coloratus), per il capo nettamente più stretto del pronoto (capo poco più stretto del 
pronoto in coloratus) e per gli uroterghi basali privi di punteggiatura al di fuori dei 
tubercoletti marginali. 

Zyras (Zyrastilbus) adesi sp. n. Figg. 264-267 

Holotypus S, Hong Kong, Kadoorie Agricultural Research Centre, flight interception 
trap, 19-31.V.1996, de Rougemont leg. (MHNG). 

Paratypi:415 es., stessa provenienza, ma anche IX-X.1991, G. Ades leg., VI. 1992, 
G. Ades leg., V-VIII.1996, de Rougemont leg., IX. 1996, de Rougemont; 6 es., Hong Kong, Tai 
Po, VIII.1992, J. Fellow leg. e VI.1996, de Rougemont leg.; 12 es., Hong Kong, Ng Kai Tin, 
Malaise trap, 23.VIII.1996, G.T. Reels leg. 

Descrizione. Lunghezza 3,9 mm. Capo ed elitre debolmente lucidi, pronoto 
opaco, addome lucidissimo. Capo ed elitre bruni, pronoto bruno-rossiccio, addome 
giallo-rossiccio; antenne bruno-rossicce con gli antennomeri terminali da 8° a 11° 
giallo-rossicci; zampe giallo-rossicce. La reticolazione del capo è vigorosa. Il pronoto è 
coperto da tubercoletti contigui a diametro maggiore sulla linea mediana: essi danno un 
aspetto vellutato o pruinoso alla superficie. Non vi è retic