Full text of "Revue suisse de zoologie"

See other formats

aig Ts
TU TH

CAT
oi abe oh op ot Em wire
RAP Baars

oh ant À M athe Baer
Er Eng Paz
A pier das
19;
PACE
AT Rene
od Aa IA
DE Li se A AA.
A
POCA

PAT

EHE,
CECA PE
CLONI

OT
ali

re
parta ran
nee
PURE TE ET tm te Ae
de an abot Pen À ne
RE EHEN

Den
Era

“ne E,

ce
ee
ps

BEE
CAE ate FE
pertimztizai Der = Se EEE Fr
Po 2 7 PP ee
PANNE ge de a ae 4
TRE
RTE >
Gah CE Frater ZI
ee Te ee

vette
MENT

argine dina
Sie

Ye à
LR NE È
x ditta an,
Sas
INECTERO pren SE

ees

>
suas

eds ae eres
ROSES

in.

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 LÔBL — 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

MUSEUM 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 étre 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

SOCIETE SUISSE DE ZOOLOGIE

et du

MUSEUM D'HISTOIRE NATURELLE
de la Ville de Geneve

tome 105
fascicule 3
1998

E :
EI GENEVE SEPTEMBRE 1998 ISSN 0035 - 418 X

SMIT HSonza

REVUE SUISSE DE ZOOLOGIE

TOME 105 — FASCICULE 3

Publication subventionnee par l'Académie suisse des Sciences naturelles
et la Societe suisse de Zoologie

VOLKER MAHNERT
Directeur du Museum d'histoire naturelle de Geneve

FRANCOIS BAUD
Conservateur au Museum d'histoire naturelle de Geneve

CHARLES LIENHARD
Chargé de recherche au Muséum d’histoire naturelle de Geneve

Comite de lecture

President: Ivan LOBL — Museum de Geneve

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du
Muséum de Geneve et de représentants des Instituts de zoologie des universites
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 étre adressées
à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse

REVUE SUISSE DE ZOOLOGIE 105 (3): 465-485; septembre 1998

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

Stefano POZZI", Yves GONSETH? & Ambros HANGGI

! Muséum d'histoire naturelle, Case postale 6434, CH-1211 Genève 6.
2 CSCF, Terreaux 14, CH-2000 Neuchâtel.

3 Naturhistorisches Museum, Augustinergasse 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 seches constituent les milieux herbacés les plus riches en especes
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 these du premier auteur a l’Université de Genève sur
|’ étude des araignées des prairies sèches en tant que bioindicateur de la qualité du milieu, projet
753-V A-1116/00 de I’ 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 Ecosystemes sont semi-naturels, ils
ont été créés et maintenus par les activités agricoles traditionnelles non mécanisées,
essentiellement le paturage et la fauche.

En Suisse, la disparition progressive des prairies sèches a encouragé 1’ 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; Pozzı 1996).
Present dans tous les biotopes terrestres, ce groupe d’arthropodes très abondants
comprend beaucoup d’espèces dont les peuplements sont révélateurs de conditions
écologiques précises (HANGGI er 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), LÖRTSCHER er al. (1994), ANTOGNOLI et al. (1995), BAUR
et al. (1996) ont utilisé les araignées pour évaluer la qualité de prairies seches 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
seches par le biais des araignées, basée sur les travaux de HANGGI (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’intérét 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; HANGGI 1993a;

EVALUATION 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) paturage bovin (6 stations), b) paturage 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 ef al. 1995).

RÉCOLTES DES ARAIGNÉES

La récolte des données arachnologiques a été effectuée avec des pièges au sol
(Barber): gobelets blancs en polypropylène de 7 cm de haut et 7 cm de diamètre,
remplis au tiers d’un liquide conservateur (formaldéhyde 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 HANGGI 1989). Le matériel sera déposé
au Muséum d'histoire naturelle de Genève et au Naturhistorisches Museum de Bâle.

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
CLVMI et CLVM2 classes des indices VMI et VM2

CEV ST: classes de valeur stationnelle globale (VST)
CLVSTH classes de valeur stationnelle théorique (VSTH)
EI stations entretenues irrégulièrement

F indice de fidélité des espèces

BE stations fauchées précocement

ET 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’especes

PB stations paturées 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 VMI et VM2

SVP milieu sans valeur particuliere

THVL milieu de tres haute valeur

UGBN charge en bétail, nombre UGB pour 100 jours de pature
V valorisation: SVP, IVL, VAL, HVL et THVL

VAL milieu de valeur

VM indices VMI et VM2

VMI 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 dies VSP d’une station

EVALUATION DES STATIONS

Plusieurs méthodes ont été proposées pour |’ évaluation d’un milieu du point de
vue de la protection de la nature (littérature compilée dans MARTI & STUTZ 1993). Elles
présentent différents objectifs: Evaluation qualitative d’un site, Evaluation des effets des
mesures de gestion, évaluation a différents niveaux de perception de l’espace: paysage,
milieu complexe ou station. En Allemagne, plusieurs auteurs ont proposé des méthodes
pour des évaluations plutöt globales en considérant différents facteurs abiotiques et
divers groupes biologiques (PLACHTER 1992, 1994; HOLSTEIN 1995; BEINLICH ef al.
1995):

La methode utilisée ici se base sur les travaux de HANGGI (1987) et sur leurs
développements ultérieurs (HANGGI 1990). Elle n’est pas basée sur l’ensemble de la
faune mais est au contraire focalisée sur les araignées. Cette approche est fondee 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 especes «piégées» dans une station. Nous pensons en effet, que la valeur d’une
station ne peut étre définie seulement a partir de quelques especes 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 méme si toutes les especes 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

EVALUATION DE L’ENTRETIEN DES PRAIRIES SECHES 469

pas sous-estimer la qualité des biotopes tres homogenes renfermant peu d’especes mais
dont les liens avec les conditions du milieu sont très étroits (espèces sténoèces) et
parallélement de ne pas surestimer les stations riches en especes ubiquistes (euryeces).
Par exemple, une roseliére renferme peu d’especes mais de valeur spécifique élevée
tandis que certains milieux artificiels bien structurés ont un grand nombre d’especes
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 especes capturées. Ces indices sont les suivants:

RARETE (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 & HANGGI 1990; HANGGI 1993b; BAUR ef al.
1996; Pozzı 1996). Certaines modifications ont donc été apportées aux indices retenus
par HANGGI (1990). Ces modifications ont en outre entraîné une révision des limites des
classes préalablement établies.

classes rareté NOS % N % NSP

1 espece banale NOS > 30 PET 9.6

2 espèce très commune 30 > NOS > 20 20.0 1226

3 espece commune 20 > NOS > 12 15.8 18.4

4 espèce peu commune 12 > NOS > 6 1742 21.8

5) espece rare 6 =NOS > 2 16.3 25.4

6 espèce très rare 22 NOS 13.0 al
NOS Nombre d’observations en Suisse (versant nord des Alpes uniquement)
TN 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? + x avec x = 1, 2, 3,4 et 5

FIDELITE (F)

La fidélité d’une espèce a 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 & HANGGI 1990 et HANGGI er 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 atrribué a une
espèce donnée ne peut être reprise sans autre pour l’Evaluation de stations appartenant a
une autre region. De manière générale, un indice de faible valeur est attribué a une
espèce peu exigeante fréquentant différents milieux de structure peu précise; un indice
de valeur élevée est attribué a une espèce liée a un milieu de structure précise. La

signification de l’indice attribué a chaque espèce est la suivante:

470 S. POZZI, Y. GONSETH & A. HANGGI

CRITERES TRIVIALITE RARETE STATUT

pourcentage des
espèces euryéces

Nb de stations connues
au nord des alpes

Expression des

Spee ER Fidélité au biotope
differents criteres

pEU

Valeur spécifique
VSP=R*F

Standardisation des
critéres

Valeur stationnelle globale
VST => VSP

Valeur stationnelle
théorique
VSTH = VST/ NSP

See onecoencoenoh=

Classes de % des espèces! Classes de valeur
euryéces stationnelle globale
CLpEU CLVST

Classes de valeur
stationnelle théorique
CLVSTH

VM2
CLpEU * CLVSTH

Classification
CLVM2

Estimation de la valeur
de chaque station

Classification
CLVMI

SVP: SUM =< 8
IVL: SUM =< 18
VAL: SUM =< 32
HVL: SUM =< 50

THVL: SUM =< 72

Fic. 1. Méthode d’évaluation des stations. SVP= sans valeur particuliere; IVL= interessant, de
valeur limitée; VAL= de valeur; HVL= de haute valeur; THVL= de très haute valeur.

EVALUATION DE L’ENTRETIEN DES PRAIRIES SECHES 471

JE espèce très peu exigeante et/ou tolérant les milieux artificiels - euryece (sans
exigence), ubiquiste - ex: forêts et milieux ouverts, secs et humides, zones très
artificielles

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

SI 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

DE espèce exigeante, liée à un type de milieu bien précis - sténoèce modérée - ex:
prairies seches

6: espèce très exigeante, liée à des conditions bien particulières d’un type de milieu
bien précis - sténoèce stricte - ex: prairies seches 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 espece
(VSP = valeur spécifique). La valeur stationnelle globale (VST) est calculée en effec-
tuant la somme des valeurs des espéces 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 VMI et VM2 (Fig. 1).

COMPARAISON DES INDICES VM1 ET VM2

Les indices VMI et VM2 expriment des aspects différents de la qualité d’une
station donnée. VMI favorise les stations ayant une haute diversité faunique (milieux
mosaiques, écotones); VM2 favorise plutöt les milieux abritant une faune particuliere
(milieux extrêmes: homogènes, très stables). Ces deux indices sont donc complé-
mentaires (et non contradictoires). De maniére 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 particulierement bien la présence de
milieux extrémes dans une étude donnée (par exemple: une roseliere aura VM2 tres
haut et VMI bas; tandis qu’un milieu mosaïque aura VM2 bas et VMI 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 = VMI+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 particuliere 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 trés haute valeur SUM < 72

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

Le recours aux variables suivantes permet d’obtenir les indices de valeur stationnelle
VMI et VM2:

— 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)

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

HANGGI (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 ESPECES EURYECES (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 (HANGGI 1987). Sous le terme d’euryèces sont comprises les
espèces appartenant a 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), a 4 (F=R=2 VSP=4 VST=180), a 6.25 (F=R=2.5 VSP=6.25 VST=
291.25), 29 (F—R=3 VSP=9 VSil—405) et a 12°25 (F=R=3:5 VSP-P25NSIT 5525)

1S ASSESS ODES) 2: 101.25 < VST < 180 3: 180 < VST < 281.25
4: 281.25 < VST < 405 5: 405 < VST < 551.25 6: VS7>53125

EVALUATION DE L’ENTRETIEN DES PRAIRIES SECHES 473

CLASSES DE VALEUR STATIONNELLE THEORIQUE (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
E=R—3 12 25 pour —R=3.5)

IE VS THs 2.25 2: 2.25 <NSTH<4 3: 4 < VSTH < 6.25
4: 6.25 < VSTH $9 5s VW SE S25 6: VSTH >12:25

CLASSES DES INDICES VM1 ET VM2 (CLVMI ET CLVM2)

1: VM=1 2:1<VMS4 3:4<VM<9
4:9<VM< 16 5: 16<VM<25 6:25 < VM < 36

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

Le choix des facteurs d’estimation des milieux et les notes attribuées aux diffé-
rentes espéces 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’especes
recensées par station. De plus, les araignées des milieux ouverts sont assez bien
connues (THALER 1985; GONSETH 1985; DELARZE 1987; BAUCHHENSS 1990; MAURER &
Hänccı 1990; HANGGI 1992; LÖRTSCHER et al. 1994; HANGGI et al. 1995, 1996; BAUR
et al. 1996).

RESULTATS

Au total, 22057 individus adultes appartenant a 215 especes ont été récoltés avec
les pieges Barber. Les résultats bruts (nombre d’individus de chaque espéce par station)
sont publiés ailleurs (Pozzi 1997). Pozzi & HANGGI (1998) présentent les principaux
résultats taxonomiques et faunistiques. Les analyses présentées dans le Tab. 1 ont été
faites a partir du statut de rareté et de fidélité de chaque espece (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 HANGGI (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 Evaluation des sites et
de leur entretien.

Premiérement, 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 séches c’est-a-dire des stations qui a priori
étaient de valeur.

474 S. POZZI, Y. GONSETH & A. HÄNGGI

TAB. 1. Tableau des indices des différentes stations regroupées par type d’entretien.

st15 | st27 | st33 | st34 | st36 | st45 | st49

6.4 8.8

23.7 | 15.9 | 28.1 | 31.8
een

CHISRENSISEESENEN (m ||:

VAL | VAL | IVL

NIND= nombre d’individus; NSP= nombre d’especes; 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; VM1/2= indice VM1/2; CLVM1/2= classe
de VM1/2; SUM= VM1+VM2; V= valorisation; stl= station no.1; 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 paturées par ovins; XPB= moyennes des stations paturées par bovins; SVP= sans
valeur particulière; IVL= intéressant, de valeur limitée; VAL= de valeur; HVL= de haute valeur;
THVL= de très haute valeur.

475

EVALUATION DE L’ENTRETIEN DES PRAIRIES SECHES

‘SUQUUQ =f {UO[QUEUD =] ‘pue]o) =H :oddıy e] =9 10 4 ‘uopuoypy = ‘seg[mano =q ‘»uuog
-9P-914 =O {THOA-9P-UINON =q !DIPA-PP-UInNoW =V ‘96 19 SH US sddIpMa SUONE]S = „ :uoneJs e] op OLQWINU = [] !(suoneJs Sap %) UONESLIOJEA
=A TNA+IINA =WNS ‘neea amey son op =TAHL ‘INOJPA oiney op =TAH :In9feA op =TVA :Pallun] Ina op ‘JUESSOIQUI =TA] ‘o1orpnonsed
INg[va sues =qAS ‘saguuopuege SUOHEIS =GVY ‘JUQUIDAIPIE] sagyone} SUONEIS = A JUIWI90991d sagyone} SUONEIS =44 ‘USWIdIIT[NSIL1 sonuaja.nus
suonels =]q ‘sutAo sed saainjed suonrys =Od ‘sulaog Jed saainjed suogels =qq ‘USNANUS | 2P UONIUOJ Ud sUONEIS Sap sIMo[ea Sap neajgeL ‘7 “Ol

(% St) TAH (% 0€) TVA (% OD) TAI

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 & HANGGI (1990); F= indice de fidélité; R= indice de rareté.

espèce

Walckenaeria dysderoides

Walckenaeria furcillata

Dysdera crocata

Walckenaeria monoceros

Bathyphantes gracilis

Harpactea hombergi

Centromerita bicolor

Harpactocratus drassoides

Centromerita concinna

Zodarion italicum

Centromerus dilutus

Pachygnatha degeeri

Centromerus incilium

Aculepeira ceropegia

Centromerus serratus

Araneus diadematus

Centromerus sylvaticus

Hypsosinga albovittata

Diplostyla concolor

Hypsosinga sanguinea

Lepthyphantes arenicola

Neoscona adianta

Lepthyphantes keyserlingi

Lepthyphantes leprosus

Lepthyphantes mengei

Acartauchenius scurrilis

Lepthyphantes pallidus

Araeoncus humilis

Ceratinella brevis

Lepthyphantes tenuis

Meioneta mollis

Ceratinella scabrosa

Meioneta rurestris

Cnephalocotes obscurus

Meioneta saxatilis

Dicymbium brevisetosum

Meioneta simplicitarsis

Diplocephalus latifrons

Microlinyphia pusilla

Eperigone trilobata

Microneta viaria

Erigone atra

Neriene furtiva

Erigone dentipalpis

Porrhomma microphthalmum

Erigonella hiemalis

Sintula cornigera

Erigonoplus globipes

Stemonyphantes lineatus

Gonatium rubens

Theonina cornix

Gongylidiellum latebricola

Dipoena coracina

Jacksonella falconeri

Enoplognatha thoracica

Metopobactrus prominulus

Episinus truncatus

OIBWINIWINIEINININ WIE LM le Dire Row ne le | wow ID Do Sn ln | ww ID |"

Micrargus herbigradus

Euryopis flavomaculata

Micrargus subaequalis

Euryopis laeta

WM} | OV]

Minicia marginella

Euryopis quinqueguttata

Mioxena blanda

Neottiura bimaculata

Monocephalus fuscipes

Neottiura suaveolens

Oedothorax apicatus

Robertus lividus

Ostearius melanopygius

Robertus neglectus

Panamomops sulcifrons

Steatoda albomaculata

Pelecopsis parallela

Steatoda phalerata

Pocadicnemis juncea

Silometopus bonessi

WIR IRIE IN HR 0 | NONA en Do nm ln US D ID IR BR |m|S nm SSID ID Ww] Bi) MN) ty

Theridion impressum

Theridion nigrovariegatum

Tapinocyboides pygmaeus

jAlopecosa accentuata

Tiso vagans

Trichoncus saxicola

Alopecosa cuneata

Alopecosa fabrilis

Trichopterna cito

Alopecosa pulverulenta

Typhochrestus digitatus

Alopecosa striatipes

Typhochrestus simoni

Alopecosa trabalis

Walckenaeria acuminata

Arctosa figurata

Walckenaeria antica

Aulonia albimana

Walckenaeria corniculans

db RRRRÀ

Dolvialainıalivirialw)rlwiunl-lnviulu ini Raw unies ln ai PIRI IS ein uw IR a IS UD ID Ie lR IS 0 DIU So IUT

Pardosa agrestis

mIDIUIU|IBR|Im IRIS IR HR WIN IR ID | IND En

Diminwluniminin|h|uin | lalwleiuininulalw|wlniuiulw || Rlalsnbiulpislw ls ls now iulals|s|wlR | Rubin lan) |

EVALUATION DE L’ENTRETIEN DES PRAIRIES SECHES

espece

Pardosa bifasciata

Pardosa hortensis

Micaria guttulata

Pardosa saltans

Micaria pulicaria

Pardosa monticola

Pardosa nigriceps

palustris

Poecilochroa variana

Zelotes apricorum

Zelotes atrocaeruleus

Zelotes civicus

Zelotes erebeus

Pardosa vittata

Zelotes exiguus

Pirata hygrophilus

Zelotes latreillei

Pirata latitans

Tricca lutetiana

Zelotes lutetianus
Zelotes pedestris

Trochosa robusta

DIWIM DIS Hw IR RK |HIH ID IR IQ TT

Trochosa ruricola

Zelotes petrensis
Zelotes praeficus

Trochosa terricola

D |

Xerolycosa miniata

Zelotes pumilus

Zelotes pusillus

Pisaura mirabilis

Zelotes villicus

Oxyopes lineatus

Cicurina cicur

Coelotes inermis
Coelotes terrestris

Zora nemoralis

Zora parallela

Zora silvestris

DUR IAIN) RIB) CS WW la 8 | CR Olan

Zora spinimana

Histopona torpida

Philodromus rufus

Mastigusa arietina

Thanatus formicinus

Tegeneria silvestris

Thanatus atratus

Antistea elegans

Oxyptila atomaria

Hahnia nava

Hahnia pusilla

Argenna subnigra
Dictyna arundinacea

Oxyptila blackwalli
Oxyptila claveata

Oxyptila pullata

Oxyptila scabricula

Titanoeca quadriguttata
Agroeca brunnea

Oxyptila simplex

Oxyptila trux

\Agroeca cuprea

Xysticus acerbus

Agroeca proxima

Xysticus audax

Apostenus fuscus

Xysticus bifasciatus

Phrurolithus festivus

Xysticus cambridgei

Phrurolithus minimus

Xysticus cristatus

Phrurolithus nigrinus

{Xysticus erraticus

Scotina palliardi

UlnAlwlelwlalalmol/aAlalElrolwlml—lAlelelmlelalalalelelalmlelelalalelalelalwlel|elalz

Xysticus kempeleni

Cheiracanthium virescens

Xysticus kochi

Clubiona brevipes

Xysticus lineatus

Clubiona coerulescens

Xysticus robustus

Clubiona diversa

Ballus chalybeius

Clubiona frutetorum

Bianor aurocinctus

Clubiona neglecta

Euophrys aequipes

Clubiona pseudoneglecta

Euophrys aperta

Clubiona terrestris

Callilepsis schuszteri

Euophrys frontalis

Evarcha arcuata

ov/iwvlivieiVvieirlulivdeelwiwvn!leiulwiun win) elwiun/eA/wIn

Drassodes cupreus

Evarcha laetabunda

Drassodes lapidosus

Heliophanus cupreus

Drassodes pubescens

Heliophanus flavipes

Gnaphosa lucifuga

Myrmarachne formicaria

Haplodrassus dalmatensis

Pellenes tripunctatus

WINN D Ian a/| D IS IR |IRID IRD | [MO ID IRIDIDIRIQIRIRID AIS IDR Ua ID IRD ID ID IRD auf]

Haplodrassus kulczynskii

Phlegra fasciata

Haplodrassus signifer
Micaria albimana

Phlegra insignata

Synageles hilarulus

ra von Io uw)

Www

Micaria formicaria

EEREERR ERRE RES

JARS RRB RAA

477

478 S. POZZI, Y. GONSETH & A. HÄNGGI

COMPARAISON DES STATIONS PAR TYPE D’ENTRETIEN

PATURAGE BOVIN

Les stations paturées par les bovins se sont révélées les moins riches en especes,
et celles dont les valeurs stationnelles théorique et globale étaient les plus basses
(Tab. 1). Soulignons qu’en plaine la pature 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 ef 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)! 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.

PATURAGE 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

! La charge en bétail est exprimée en UGBN, nombre UGB pour 100 jours de päture; 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.

EVALUATION 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 a 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 er 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 ef 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 m&me prairie,
la premiere est fauchée tardivement toutes les années alors que la deuxieme est fauchée
tous les deux ans. Malgré leur proximite, 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 a 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 méme 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-
site 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
recelent 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 481

Dans cette étude, les stations abandonnées sont de deux types: des prairies très
xeriques a Evolution lente et des prairies en voie d’embroussaillement prononcé. Le
premier cas est typique des zones xériques qui devraient étre entretenues le moins
possible. Les stations du plateau du Moulin-de-Vert (st. 1 et 45) et de 1’ Allondon (st. 11
et 49) illustrent bien cette situation. Pour ces milieux de grande taille, l’absence
d’entretien leur convient bien parce que les conditions extrémes (climat, sol) empéchent
un changement microclimatique trop rapide (végétation herbacée dense, buissons). La
tendance au feutrage, due au manque de fauche, n’entraine pas la disparition des
especes les plus xérophiles, puisqu’il existe encore assez de zones ouvertes (terre,
cailloux) dans la station. Deux autres stations xériques (st. 24 et 14) entretenues
regulierement pourraient étre améliorées en intervenant plus extensivement.

En ce qui concerne les zones moins seches 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’apres nos
résultats, le fauchage tardif pourrait étre 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 VMI et VM2. L’extréme 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 VMI 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 a celle de VMI. 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 VMI 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

systeme prairie seche et les especes qui le caractérisent. L’abandon d’une station
ne devrait intervenir que dans les zones a 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 étre préjudiciable pour la faune a 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 tres 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 étre 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-

EVALUATION 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 (VMI 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 a 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 seches. Par leur position élevée dans la chaine 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 a la suite d’un entretien plus extensif. Ces résultats sont
importants mais devront étre confirmés par des études a plus long terme. A l’avenir, il
semble donc important d’utiliser les araignées pour caractériser un milieu et son évo-
lution (dynamique) en parallele de la phytosociologie. Pour cela, il est nécessaire de
poursuivre les études autécologiques fines sur la faune arachnologique, en particulier
sur les espèces liées a 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 a vérifier le bien-fondé des buts de protection et 4 améliorer les
mesures de gestion. Le succès de ces différents pas dépend de l’obtention de connais-
sances scientifiques sur les prairies seches qui doivent englober des aspects biotiques et
abiotiques ainsi que leurs interactions.

Par ce travail, nous espérons avoir contribué a 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
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 foréts, de la
faune et de la protection de la nature (Canton de Geneve), du Naturhistorisches
Museum Basel et du Muséum d’histoire naturelle de Genève.

484 S. POZZI, Y. GONSETH & A. HANGGI

BIBLIOGRAPHIE

ANTOGNOLI, C., LORTSCHER, 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.

BAUCHHENSS, E. 1990. Mitteleuropäische Xerotherm-Standorte und ihre epigäische Spinnenfauna
— eine autökologische Betrachtung. Abhandlungen des Naturwissenschaftlichen Vereins
in Hamburg NF 31/32: 153-162.

BAUR, B., JosHI, J., SCHMID, B., HANGGI, A., BORCARD, D., STARY, J., PEDROLI-CHRISTEN, A.,
THOMMEN, G. H., LUKA, H., RUSTERHOLZ, H.-P., OGGIER, P., LEDERGERBER, S. &
ERHARDT, A. 1996. Variation in species richness of plants and diverse groups of
invertebrates in three calcareous grasslands of the Swiss Jura mountains. Revue suisse de
Zoologie 103(4): 801-833.

BEDFORD, S. E. & USHER, M. B. 1994. Distribution of arthropod species across the margins of
farm woodlands. Agriculture, Ecosystems and Environment 48: 295-305.

BEINLICH, B. HERING, D. & PLACHTER, H. 1995. Ein standardisiertes Bewertungsverfahen für die
Kalkmagerrasen der Schwäbischen Alb. Beihefte. Veröffentlichungen für Naturschutz und
für Landschaftspflege Baden-Würtenberg 83: 425-439.

CLAUSEN, I. H. S. 1986. The use of spiders (Araneae) as ecological indicators. Bulletin British
Arachnological Society 7: 83-86.

DELARZE, R. 1987. La faune des pelouses steppiques valaisannes et ses relations avec le tapis
végétal. II: Les araignées (Araneida) et les mille-pattes (Myriapoda Diplopoda). Bulletin
romand d’entomologie 5: 1-14.

GALLAND, P. & GONSETH, Y. 1990. Typologie des milieux de Suisse. Ligue suisse pour la pro-
tection de la nature, Bale, Centre suisse de cartographie de la faune, Neuchatel, 48 pp.

GIBSON, C. W. D., HAMBLER, C. & BROWN, V. K. 1992. Changes in spider (Araneae) assemblages
in relation to succession and grazing management. Journal of Applied Ecology 29:
132-142.

GONSETH, Y. 1985. Influence de l’entretien de trois pelouses sèches du Jura neuchätelois sur leurs
peuplements arachnologiques. Mitteilungen der Schweizerischen Entomologischen
Gesellschaft 58: 77-86.

GONSETH, Y. 1994. La faune des Lépidoptères diurnes (Rhopalocera) des pâturages, des pelouses
seches et des prairies de fauche du Jura neuchätelois. Mitteilungen der Schweizerischen
Entomologischen Gesellschaft 67: 17-36.

GONSETH, Y. & MULHAUSER, G. 1995. Bioindication et surfaces de compensation écologique.
Cahier de l'Environnement No 261, Nature et paysage, BUWAL, Bern, 135 pp.

HANGGI, A. 1987. Die Spinnenfauna der Feuchtgebiete des Grossen Mooses, Kt. Bern - II. Beur-
teilung des Naturschutzwertes naturnaher Standorte anhand der Spinnenfauna. Mit-
teilungen der Naturforschenden Gesellschaft in Bern 44: 157-185.

HANGGI, A. 1989. Erfolgskontrollen in Naturschutzgebieten. Natur und Landschaft 64: 143-146.

HANGGI, A. 1990. Les Araignées, pp. 70-109. In: GONSETH, Y. (éd.). Etude d’impact sur les
milieux naturels a l’aide de la faune invertébrée I. Ouvrage RNI6 Transjurane,
Laboratoire d'écologie animale et d’entomologie, Université de Neuchâtel, 156 pp.

HANGGI, A. 1992. Spinnenfange in Magerwiesen und Brachen aus dem Tessin - Unkommentierte
Artenlisten. Arachnologische Mitteilungen 4: 59-78.

HANGGI, A. 1993a. Minimale Flächengrösse zur Erhaltung standorttypisher Spinnengemein-
schaften - Ergebnisse eines Vorversuches. Bulletin de la Société Neuchäteloise des
Sciences naturelles 116(1): 105-112. C.R. XIII® Colloque européen d’Arachnologie,
Neuchätel 2-6 sept. 1991.

HANGGI, A. 1993b. Nachträge zum «Katalog der schweizerischen Spinnen» - 1. Neunachweise
von 1990 bis 1993. Arachnologische Mitteilungen 6: 2-11.

EVALUATION DE L’ENTRETIEN DES PRAIRIES SECHES 485

HANGGI, A., STÖCKLI, E. & NENTWIG, W. 1995. Habitats of central european spiders -
Characterisation of the habitats of the most abundant spider species of Central Europe
and associated species. Miscellanea Faunistica Helvetiae 4, CSCF, Neuchätel, 459 pp.

HANGGI, A., DELARZE, R. & BLICK, T. 1996. Beitrag zur Kenntnis der Spinnenfauna des Kantons
Wallis. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 69: 189-194.

HEUBLEIN, P. 1983. Räumliche Verteilung, Biotoppräferenzen und kleinräumige Wanderungen
der epigäischen Spinnenfauna eines Wald-Wiesen Okotons; ein Betrag zum Thema
“Randeffekt”. Zoologische Jahrbücher für Systematik 110: 473-519.

HOLSTEIN, J. 1995. Die Spinnen- und Käferzönosen zweier Streuobstwiesen in Oberschwaben.
Dissertation Abteilung Okologie und Morphologie der Tiere (Biologie III), Universität
Ulm, 144 pp.

KIRBY, P. 1992. Habitat management for invertebrates: a pratical handbook. Royal society for the
protection of birds, Berdforshire, 150 pp.

LORTSCHER, M., HANGGI, A. & ANTOGNOLI, C. 1994. Zoological Arguments for managing the
abandoned grasslands on Monte San Giorgio - based on data of three invertebrate groups
(Lepidoptera, Araneae, Saltatoria). Mitteilungen der Schweizerischen Entomologischen
Gesellschaft 67: 421-435.

MAELFAIT, J.-P., DESENDER, K. & BAERT, L. 1989. Some examples of the practical use of spiders
and carabid beetles as ecological indicators. Comptes rendus du Symposium «Invertébrés
de Belgique»: 437-442.

MAERTENS, T., WAHLER, M. & Lutz, J. 1990. Landschaftspflege auf gefährdeten Gründland-
standorten. Schriftenreihe Angewandter Naturschutz der Naturlandstiftung Hessen 9:
1-167.

MARTI, F. & Stutz, JH.-P. 1993. Zur Erfolgskontrolle im Naturschutz. Literaturgrundlagen und
Vorschläge für ein Rahmenkonzept. Berichte der Eidgenüssischen Forschungsanstalt für
Wald, Schnee und Landschaft 336, 171 pp.

MAURER, R. & HANGGI, A. 1990. Katalog der schweizerischen Spinnen. Documenta Faunistica
Helvetiae 12, CSCF, Neuchätel, 412 pp.

PLACHTER, H. 1992. Grundzüge der naturschutzfachlichen Bewertung. Veröffentlichungen für
Naturschutz und für Landschaftspflege Baden-Würtenberg 67: 9-48.

PLACHTER, H. 1994. Methodische Rahmenbedingungen für synoptische Bewertungsverfahren im
Naturschutz. Zeitschrift für Okologie und Naturschutz 3: 87-106.

PLATEN, R. 1993. A method to develop an ‘indicator value’ system for spiders using canonical
correspondence analysis (CCA). Memoirs of the Queensland Museum 33(2): 621-627.

Pozzi, S. 1996. Les invertébrés de lisieres naturelles et dégradées du Canton de Genève. Bulletin
romand d’entomologie 14: 1-38.

Pozzi, S. 1997. Spinnenfänge aus Magerwiesen der Kantone Genf und Waadt (Schweiz) -
Unkommentierte Artenlisten. Arachnologische Mitteilungen 14: 55-80.

Pozzi, S. & HANGGI, A. 1998. Araignées nouvelles ou peu connues de la Suisse (Arachnida:
Araneae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 71: 33-47.

Pozzi, S. & BORCARD, D. (soumis). Effets de l’entretien des prairies sèches du plateau occidental
suisse sur les peuplements d’araignées. Ecologie.

SCHULTZ, W. & FINCH, O. D. 1996. Biotoptypenbezogene Verteilung der Spinnenfauna der
nordwestdeutschen Küstenregion — Charakterarten, typische Arten und Gefährdung.
Cuvillier Verlag, Göttingen, 141 pp.

THALER, K. 1985. Über die epigäische Spinnenfauna von Xerothermstandorten des Tiroler Inn-

tales (Österreich) (Arachnida: Aranei). Veröffentlichungen des Museum Ferdinandeum in
Innsbruck 65: 81-103.

veti Re
an! Mr n

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 A. 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. Löbl)
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: 14, 42, 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: 14,39, North Carolina, Watauga Co., Boone, 12.IIL./ 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: 34, 19, Bjelasnica planina (MHNG).

Bulgaria: 26, 29, Pirin, leg. Weirather (MHNG); 32, Ali Botusch, ‘N-Seite’ near
Goleschowo, 1015m, 41°42’13N, 23°35’21E, Fagus wood, 15.VI.1997, leg. Behne (DED; 14,
Sredna Gora, S Koprivschtiza, 1110m, 42°35’12N 24°22°19E, Fagus wood, 29.V1.1997, leg.
Zerche & Behne (cAss).

Greece: 15,3%, Crete, Zakros, ‘Tal der Toten’, 28.11.1973, leg. Fülscher & Meybohm
(MHNG); 54,42, Rédhos, M. Kariona, 400m, 11.1V.1977, leg. Besuchet (MHNG, cAss); 29,
Rodhos, Petaloudes, 8.1V.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: 24, 19, E. Asturias, Cibes, Sierra de la Serrantina, 800m, 16.V.1997, leg.
Starke (coll. Feldmann, cAss); 19, Albacete, leg. Comellini (MHNG); 14, Prov. Albacete,
Villaverde, 15.1V.1959, leg. Besuchet (MHNG); 16, Prov. Tarragona, Sierra de Moutsant (?),
23.11.1959, leg. Besuchet (MHNG); 16, 19, Prov. Lèrida, 2.VI.1965, leg. Comellini
(MHNG); 26, 19, Prov. Teruel, Rubielos de Mora, 19.IX.1971, leg. Comellini (MHNG); 19,
Prov. Murcia, Sierra de Espuña, 26.111.1959, leg. Besuchet (MHNG); 36, 19, Prov. Jaen,
Sierra de Cazorla, 12.1V.1959, leg. Besuchet (MHNG).

Previously, only the type specimens of A. 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 489

Habrocerus neglectus sp. n. Figs 1-4

Habrocerus rougemonti Pace: ASSING & WUNDERLE 1995 (partim, Figs 9a-g, 11g)

Holotype d : THAILAND. NE Bangkok, Khao Yai Nat. Park, Khao Khieo, 1150m, leg.
Burckhardt & Löbl, 28.X1.85 (MHNG).

Paratypes: 116, 192, same data as holotype (MHNG, cAss, cWun); 3d, Chiang Mai,
Mae Nang Kaeo, 900m, 54 km NE Chiang Mai, leg. Burckhardt & Löbl, 3.XI.1985 (MHNG,
cAss).
DiAGNosis: Closely related to and externally indistinguishable from H. rougemonti
Pace, its sister species.
d : tergum and sternum VII slightly broader than in A. 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).
2: 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: 54, 19, mountains near Umphang, 1250 m, 10.11.1993, leg. Schwendinger (MHNG,
CASS).

DIAGNOSIS: d : tergum and sternum VII slightly more oblong than in A. 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).

2: 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: 96, 49, Alberta, George Lake, 53°57’N, 114°06’W, ex mushrooms, 11./
22.V111.1977, leg. Ashe (UKNHM, cAss); 16, 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

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

Nomimocerus longispinosus Assing & Wunderle

Chile: 24, Chiloé, Isla Chiloé, San Juan de Chadmo, 20.11.1997, leg. Cekalovic
(UKNHM, cAss); 29, 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: 46, 42, Osorno, 14 km E Termas de Puyehue, 40°40’S, 71°14’ W, 350 m, ex
fungus:on live vine, 30.XT.1994, leg. Leschen & Carlton (UKNHM, cAss); 12, same data, but
450 m, ex sifting leaf litter (UKNHM); 14, 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); 14, Cautin, 8 km S Pucon, 1075 m, 39°21’S, 71°58’ W, 23.X1.1994, leg. Leschen
& Carlton (UKNHM); 14, Cautin, 26.7 km E Pucon, 625 m, 39°39’S, 71°47’W, 24.X1.1994,
leg. Leschen & Carlton (UKNHM); 16, Cautin, Termas de Palguin, Salto Puma, 725 m,
39°22’S, 71°50’ W, 25.X1.1994, leg. Leschen & Carlton (cAss).

N. 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 ¢: 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: 54, 39, same data as holotype (UKNHM, cAss); 68, 42, 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); 26, 92 [22 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).

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

49]

A REVISION OF THE HABROCERINAE OF THE WORLD

Will CA A
COPINE

492 VOLKER ASSING

DrAGNosis: Total length: 3.8 - 4.8 mm; pronotal length: 0.68 - 0.76 mm; pronotal
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.

d: 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.

ACKKNOWLEDGEMENTS

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 Löbl 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 SAKAI 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. Löbl danke ich sehr, daß er mir die Bearbeitung dieser Tiere
ermöglichte sowie für die Überlassung einiger der Käfer für meine Sammlung. Herrn
Dr Volker Mahnert, 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 05.02.1998

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. Löbl & 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-Malaysıa (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, confl. Sun Oyan and Mujong riv., E Kapit, 50 m, 18.5.1994,
# 5 a, 1 Ex., leg Löbl & 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. Fliigeldecken 3,0 lang, 2,1 gemeinsam breit.

Färbung: Kopf und Halsschild rotbraun. Fliigeldecken 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. Löbl & 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 Löbl & Burckhardt,
Paratypus. E-Malaysia, Sarawak, confl. Sun Oyan and Mujong riv., E Kapit, 50 m, 18.5.1994,
#5 a, 1 Ex., leg. Löbl & Burckhardt, Paratypus.

496 GERHARD UHMANN

Î \ 10 fl
À an SU») N
f N Ÿ
J N
fl 8 \
ABB. 6-11

Derarimus compacticornis sp. n. (6-7): 6. Habitus; 7. Halsschildprofil. — D. sabahensis sp. n.
(8-11): 8. Habitus; 9. Halsschildprofil; 10. Aedeagusspitze, dorsal; 11. 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 Malaysıa (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, 11 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-Malaysıa (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.

i, Raa

REVUE SUISSE DE ZOOLOGIE 105 (3): 499-555; septembre 1998

A taxonomic revision of the family Oncopodidae 1.
New genera and new species of Gnomulus Thorell
(Opiliones, Laniatores)

Jochen MARTENS! & Peter SCHWENDINGER?

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

? 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, 1. 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. fenuis 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

thoush they 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 revisional 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 arbitrary manner. Each seemingly allopatric population with

NEW GENERA AND SPECIES OF ONCOPODIDAE 501

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 9
and absent in dd. 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.
Sf) is of the same type. A detailed account of this shall be given in our next paper.

In contemporary taxonomy of arthropods d 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 SG,
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 d d, 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 ef al. 1981). This trait holds true not only for Oncopus
(MARTENS ef 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 er 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 Gnomulus/Oncopus 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

Fic. 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.); 1) 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.); 0) 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.j; 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

Fic. 4. Scanning electron micrographs showing external characters of Oncopodidae species. - a)
Palaeoncopus gunung sp. n., left 2 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 507

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

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

Types: SUMATRA, Northern Sumatra Province, Langkat, Bukit Lawang Nature
Reserve, 180 m, 4 holotype (MHNG Sum-85/49). - Deli Serdang, north of Brastagi, 1400 m,
d paratype (MHNG Sum-85/47); both leg. B. Hauser, 8.-20.X1.1985.

New material: SUMATRA, Aceh Province, Mt. Leuser National Park, Ketambe
Research Station, 300-500 m, 23.-30.X1.1989, 4 3, 13 2; 800 m, 28.X1.1989, 1 9; all leg. D.
Agosti, I. Löbl & D. Burckhardt (MCSNG, MHNG). - North Sumatra Province, Langkat, Bukit
Lawang Nature Reserve, 11.-12.X.1990, 1 d, 1 2; leg. A. Riedel (MAR). - Bohorok,
7.VII.1982, 1 9,1 2, 13.X1.1983, 1 2, 30.x11.1993, 2 2; all leg. C. Deeleman-Reinhold & P.
Deeleman (CCD). - Langkat, Bukit Lawang Nature Reserve, Bohorok river, 5.VII.1984, 1 3, 2
2; leg. J. Robert (MHNG). - Forest 7 km north of Brastagi, 1500 m, 2.X11.1989, 1 4,6 9: all
leg. D. Agosti, I. Lob] & 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 2 © (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 4)
available (SCHWENDINGER 1992: 197).

Variation: Measurements range: d: 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; 2: body length 3.59-4.39 (x =
LOS SD 0238) width 2.072.853 (x = 2.40, SD = 0.18 )En = 27. Body size
gradually decreases in between the three populations.

Population: mean d body length (SD) mean 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, d holotype (MHNG), 9 d, 4 © paratypes (MAR, MCSNG, MHNG), 20.-21.X1.1989. -
Palopo Nature Reserve, north of Bukittinggi, 900 m, 1 d paratype (MHNG), 18.-20.X1.1989.
All specimens leg. D. Agosti, I. Löbl & 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); 4 holotype (8), © (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 dd; 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: 3 (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

= =

9 À

Fics 9-16

Caenoncopus tenuis sp. n.; 6 holotype (9-12), 8 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.

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

510 J. MARTENS & P. SCHWENDINGER

Biel

Caenoncopus tenuis sp. n., 3 holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale
line | mm.

NEW GENERA AND SPECIES OF ONCOPODIDAE 511

Measurements (8, in brackets 2): 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:

‘itr 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)
LegI 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: 8 2.26-2.52/1.38-1.58
(n= 12); 2 2.46-2.55/1.50-1.57 (n = 4). Some d d (including the holotype) have a
smaller ventro-basal process on the palpal femur than shown in Fig. 15. In several d d
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, 4 holotype (MHNG), 2
3,2 2 paratypes (MAR, MHNG), 19.X1.1989; all leg. D. Agosti, I. Löbl & D. Burckhardt. -
Road from Lubuksikaping to Panti, ca. 700 m, 26.X. 1991, 1-2 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: & (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 I 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.

SI? 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.

2. As the 3; no external sexual dimorphism discernible.

Measurements (d, in brackets 9 ): 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:

‘Tt Fe Pa Ti Mt Ta Total
Palp 0.37 (0.38) 0.53 (0.52) 0.44 (0.43) 0.35 (0.34) - - 0.55 (0.56) 2.24 (2.23)
LegI 0.29 (0.31) 0.87 (0.81) 0.60 (0.58) 0.47 (0.46) 0.92 (0.90) 0.05 (0.06) 3.20 (3.12)
Leg II 0.34 (0.38) 1.16 (1.08) 0.75 (0.72) 0.78 (0.75) 1.40 (1.31) 0.08 (0.11) 4.51 (4.35)
Leg III 0.31 (0.29) 0.69 (0.66) 0.59 (0.58) 0.47 (0.47) 0.99 (0.95) 0.06 (0.08) 3.11 (3.03)
Leg IV 0.34 (0.39) 1.07 (1.01) 0.78 (0.76) 0.85 (0.82) 1.34 (1.28) 0.06 (0.08) 4.44 (4.34)

Variation: Measurements of body length/width range: d 3.04-3.25/1.90-2.00
(n= 3); 2 3.20-3.24/1.96-2.12 (n = 3). One d paratype with interocular tubercle
lower than in Fig. 26c.

Fics 18-25

Caenoncopus affinis sp. n.; d holotype (18-21), & paratype (22-23), 2 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

Fic. 26

Caenoncopus affinis sp. n., d 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 "0" 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, 1.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, d holotype (MHNG), | 9,4 © 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: 3 (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

Fics 27-38

Palaeoncopus gunung sp. n.; & holotype (27-30, 34), & paratype (31-32, 35), 2 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

Fic. 39

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

NEW GENERA AND SPECIES OF ONCOPODIDAE 517

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 d; no external sexual dimorphism discernible.

Measurements (6, in brackets 2): 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)
Leg I 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)
Leg II 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 IIT 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: d 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.X1.1989, 3 holotype and ® paratype; leg. D. Agosti, I. Löbl &
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

40, 41

Fics 40-45

Palaeoncopus kerdil sp. n.; & holotype (40-43), © 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: 3 (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

Fic. 46

Palaeoncopus kerdil sp. n., d 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.

2. As the d; no external sexual dimorphism evident.

Measurements (4, in brackets 9): 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:

Ahr Fe Pa Tı 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)
Leg I 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)
34)
43)

SSIS

Leg HI 0.17 (0.18) 0.39 (0.40) 0.32 ( i 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. 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.V11.1984, 5 holotype and d 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: è (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.

2. Unknown.

NEW GENERA AND SPECIES OF ONCOPODIDAE 52]

Fics 47-57

Palaeoncopus katik sp. n.; 3 holotype (47-50, 55-56), 8 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 (3): body 1.85 long, 1.24 wide; carapace 0.50 long, 0.72 wide.

- Palp and legs:

Tr Fe Pa Ti Mt Ta Total
Palp 0.20 0.26 0.25 0.15 = 0.36 1.22
Leg I 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 5 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.

322 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, ¢ holotype (MHNG), 2 8,4 9
paratypes (MAR, MHNG), 19.11.-9.1II.1991, leg. W. Schawaller & J. Martens. - Visca, N of
Baybay, 200-500 m, 1 4 paratype (MAR), 2.1IL.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

uo
ive)

Fics 58-69

Biantoncopus fuscus n. sp.; 6 holotype (58-61, 68-69), 4 paratypes (62-65) [Lake Danao (62.
64-65), N of Baybay (63)], ? 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

Fic. 70

Biantoncopus fuscus sp. n., 3 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: è (paratype). Coloration: Body amber, dark reticulations on:
carapace, margin and elevations of scuta, legs (except light orange tarsı) 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 IH 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).

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

Measurements (à, 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: 4 2.25-2.48/1.60-1.69 (n = 4), 2 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 d (from the type locality) has a thin cylindrical stylus and a narrowly
rounded median plate with a smooth margin (Fig. 62). In all other 5 d 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; 4, 2 2 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 d d from Sumatra in CCD, SMF, ZMA,
ZMT examined.

G. aborensis (Roewer, 1913), transferred from Pelitnus by ROEWER (1923: 61-62)
[NE-India] - 2 4 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 |Malaysıa, Penang
Island] - 3 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-1.

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.

crucifer sp. n. - Philippines, Luzon.

maculatus sp. n. - Philippines, Luzon and Mindoro (?).

coniceps sp. n. - Philippines, Luzon.

. leyteensis sp. n. - Philippines, Leyte.

. laruticus sp. n. - Malaysia, Perak.

. asli sp. n. - Malaysia, Perak.

. hirsutus sp. n. - Malaysia, Selangor and Pahang.

O0 000.0

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.

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

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

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

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

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

G. thorelli (Sgrensen, 1932) comb. n., transferred from Pelitnus [Java].

G. palawanensis (Suzuki, 1982) comb. n., transferred from Pelitnus [Philippines,
Palawan)].

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

Q Q Q Q

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,
d 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

Fics 71-80

Gnomulus crucifer sp. n., 4 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: 3 (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.

529

NEW GENERA AND SPECIES OF ONCOPODIDAE

Fic. 81
Gnomulus crucifer sp. n., 4 holotype, body, dorsal (a), ventral (b) and lateral view (c). - Scale

line 1 mm.

530 J. 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.

9. Unknown.

Measurements (6 ): body 3.12 long, 1.78 wide; carapace 0.88 long, 1.02 wide.
- Palp and legs:

Tr Fe Pa Ti Mt Ta Total
Palp 0.31 0.43 0.35 0.26 2 0.49 1.84
Leg I 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 II 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 (d 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, d holo-
type (MHNG), 21.XII.1979; 1 6, 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 © paratype (MAR), 2.-4.1.1979 (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]

Fics 82-88

Gnomulus maculatus sp. n.; 4 holotype (82-85), 4 paratype (86), © 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 II. 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: è (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.

J. MARTENS & P. SCHWENDINGER

532

Fic. 89
Gnomulus maculatus sp. n., 3 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.

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

Measurements (3, 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:

Dr Fe Pa li 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: 4 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 2 examined, in one of
them missing on the left side. The median plate of the glans penis is narrower in the
d 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. goodnight.

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. i Figs 90-96

Material: PHILIPPINES, Luzon, Baguio, near Cristal Caves, 1500 m, & 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

2 \lwy

Fics 90-95

Gnomulus coniceps sp. n., 3 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: 3 (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 II: palpal coxae with large ventral pro-
cesses; genital operculum longer than wide (Fig. 96b).

595

NEW GENERA AND SPECIES OF ONCOPODIDAE

Fıc. 96
‘eps Sp. n., d holotype, body, dorsal (a), ventral (b) and lateral v

(c). - Scale

1eW

S CONIC

Gnomulu

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.

2. Unknown.

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

fit Fe Pa Ti Mt Ta Total
Palp 0.42 0.61 0.51 0.33 - 0.65 DS?)
Leg I 0.33 1.05 0.56 0.64 1.03 0.59 4.20
Leg II 0.35 1235 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, d holotype
(MHNG), 10.11.1991, 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: 3 (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

zZ
=>

Fics 97-103

Gnomulus leyteensis sp. n., 3 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-VI;
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

Fic. 104

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

NEW GENERA AND SPECIES OF ONCOPODIDAE 539

2. Unknown.
Measurements (3 ): body 3.27 long, 2.39 wide; carapace region 0.76 long, 1.24
wide. - Palp and legs:

Tr Fe Pa Ti Mt Ta Total
Palp 0.32 0.50 0.43 0.31 - 0.64 2.20
Leg I 0.27 0.85 0.53 0.56 0.87 0.55 3.63
Leg II 0.35 1.14 0.69 0.88 1633 0.59 4.98
Leg HI 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 0.50 997

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. n. Figs 105-113

Material: MALAYSIA, Perak, Taiping, Maxwell Hill (= Bukit Larut), 1320 m,
d holotype (MHNG), 26.1.1995, leg. P. Schwendinger.

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

Fics 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, 109-112), 0.1 mm (107-108).

540 J. MARTENS & P. SCHWENDINGER

leven IIs)

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

NEW GENERA AND SPECIES OF ONCOPODIDAE 54]

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: & (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 IT 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. 113c). 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. 113b).

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 111-112), 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 (6 ): 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
Leg I 0923 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 3252
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 3:59

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.

Fics 114-121

Gnomulus asli sp. n.; 4 holotype (114-117), d paratypes (118-119), 9 paratype (120-121). -
Penis, dorsal (114) and lateral view (115); apex of penis, dorsal (116, 118-119) 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,
3 holotype (MHNG), 2 d, 1 £ paratypes (MAR, MHNG), 22.-31.1.1994; 330-400 m, 6 9
paratypes (MAR, MHNG), 15.-22.1.1995; 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

PR RSS

Fic. 122

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

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: ¢ (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 H with distinct anterio- and posterio-
proximal processes, coxa II 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.

2. As the d, except for an indistinct hair cover on ventral scutum.

Measurements (3, in brackets 9 ): 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:

br Fe Pa TE 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: & 2.29-2.34/1.56-1.59 (n = 3), 2 2.28-
SAISIE CE);

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

= = AR

R
N

132

Fics 123-132

Gnomulus hirsutus sp. n.; 3 holotype (123-126), & paratype (127-128, 131-132), ? 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, d holotype and | ® paratype
(MHNG), 1-3.XII.1990, leg. C. Deeleman-Reinhold. - Ulu Gombak, University of Malaya
Field Centre, 200 m, 1 5, 1 @ paratypes (MAR, MHNG), 26.IX.1991, leg. D. Agosti. -
Pahang, on the road to Genting Highlands, 500-800 m, 1 © 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

Fie. 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 sclerites further apart
from each other, median plate without marginal teeth (Figs 123-128).

Description: 3 (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.

2. As the d, except for less elevated, more strongly pigmented ventral scutal
areas.

Measurements (Ö, in brackets 9): 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)
Leg I 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: & 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 & 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 © from Ulu Gombak shows a faint pattern in
the central region of its dorsal scutum. The d 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 © 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 2 2. We are reluctant to describe
new species from 9 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 4 specimens.

1. 1 ©, Sumatra, 7 km north of Brastagi, 1500 m, 2.XII.1989, leg. I. Löbl 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 ©, 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 2, Sumatra, Mt. Kerinci, 1750-1900 m, 11.-13.X1.1989, leg. I. Löbl er al.
(MHNG) --- close to C. affinis sp. n. but different in colour pattern; eye tubercle more
rounded; genital operculum much larger; ventral process on palpal trochanter smaller.

4.2 2, Sumatra, Panti, 250 m, 19.X1.1989, leg. I. Löbl er 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. affinis sp. n.

5. 1 ©, Sumatra, Mt. Kerinci, 1750-1850 m, 11.-12.X1.1989, leg. I. Löbl er 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 ©, 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-like; 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 2, Philippines, Leyte, Visca north of Baybay, 200-500 m, 23.11.1991, leg.
J. Martens & W. Schawaller (MAR) --- similar to B. fuscus sp. n. (tarsı 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 2, 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 2, Malaysia, Perak, Taiping, Maxwell Hill, ca. 1200 m, 10.1V.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

sE In
À x en
(7 t
SS, da GE
N ee a

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]

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 fuscus 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 ¢ to ejaculate deeper into the 2 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 £ © 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, ıs 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 d 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 | (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 d d 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 Löbl (MHNG), Dr Hirotsugu Ono (National Science Museum,
Tokyo), Mr Alexander Riedel (Zoologische Staatssammlung, München), Dr
Wolfgang Schawaller (Staatliches Museum fiir 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 Kathe 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. /n: 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
fiir Systematik 16: 163-216.

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: 117-120.

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). /n: 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 fiir 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+1116 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.

SORENSEN, W. 1932. Descriptiones laniatorum (Arachnidorum Opilionum Subordinis) fecit
William Sgrensen opus posthumum recognovit 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 1889 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 VIII + IX.

Zink, R.M. 1997. Species concepts. Bulletin of the British Ornithologists' Club 117: 97-109.

REVUE SUISSE DE ZOOLOGIE 105 (3): 557-568; septembre 1998

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

TAN Heok Hui! & Maurice KOTTELAT ! 2

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

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

Redescription of Betta picta (Teleostei: Osphronemidae) and descrip-
tion of B. falx sp. n. from central Sumatra. - Betta picta 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. picta 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 trifasciata 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. picta. The type material of B. picta is lost and the
type(s) of B. trifasciata is(are) apparently mixed up with other Berta 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. picta, from Jambi, central
Sumatra, is here described as a new species. A redescription of B. picta is also
presented.

The B. picta group was first established by WITTE & SCHMIDT (1992) and later
emended by TAN & NG (in press). All members of the B. picta 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. picta 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. picta, 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 & Woop
(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 Berta,
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 preorbital 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, Geneve
(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 PICTA AND B. FALX 559

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

560 TAN HEOK HUI & MAURICE KOTTELAT

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

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

BETTA PICTA AND B. FALX 56]

DESCRIPTIONS

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

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

Betta trifasciata Bleeker, 1850: 12; GUNTHER 1861: 388; REGAN 1910: 781.

Betta picta: 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:
SS, 100, SI

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, maie; 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 picta 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. picta are listed under Affinities.

DESCRIPTION. General appearance is shown in Fig. 1; meristics and morphometrics of
the B. picta 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 preorbital 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 picta 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 picta
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 picta 1s 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
(51/2, 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 picta 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
(9-91/5, mode 9, vs. 91/2-11!/2, mode 11!/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 picta 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 preorbital black stripe (vs. thick); more modal dorsal rays (9,
vs. 8); fewer modal subdorsal scales (51/3, 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).

sample size

MERISTICS
[range (mode)]

vertebrae

anal-fin rays
dorsal-fin rays
caudal-fin rays

pelvic-fin rays
pectoral-fin rays
subdorsal scales
transverse scales
lateral scales
lateral scale below
dorsal-fin origin
lateral scale above
anal-fin origin
predorsal scales
postdorsal scales

MORPHOMETRICS

percentage of SL [range]

total length

predorsal length
postdorsal length
caudal peduncle depth
preanal length

head length

body depth

pelvic-fin length
anal-fin base length
dorsal-fin base length

percentage of HL [range]

orbit diameter
postorbital length
interorbital width

B. picta B. falx B. taeniata
12 12 16
2+8+ 17-18 2+8+ 17-18 2+8+ 17-19
total: 27-28 (27) total: 27-28 total: 27-29 (28)
II,19-21 II, 19-21 I-II, 22-24
total: 21-23 (22) total: 21-23 (22) total: 23-26 (26)
0-I, 7-9 I-II, 7-8 0-II, 8-9
total: 8-9 (9) total: 8-10 (8) total: 8-10 ( 9)

1,5+6,i ii, 5 + 6, i-li 11,5 + 6-7, 1-11
(ii, 5 + 6, li) (ii, 5 + 7, 1)
I, 1,4 I, 1,4 iets a
12 12 11-12 (12)
51/5-61/3 (51/5) St/5-61/5 (51/3 or 6) 5-6 (6)
9-91/; (9) 9-91/5 (91/5) 9-91/5 (91/5)
27-30 (28) DI 271/5-30 (29)
12-14 (13) 11-12 (12) 12-15 (14)
6-8 (7) 6-8 (6) Serien)
19-21 (20) 19-20 (19) 19-21 (19)
9-11 (10) 9-11 (10) 9-11 (10)
135.4-143.9 135.8-142.2 132.9-141.6
63.0-67.6 60.0-65.7 62.4-67.7
19.5-25.5 DYDD 19.5-24.8
14.4-17.3 15.8-18.3 14.8-19.7
48.7-53.6 48.7-52.3 47.2-54.1
31.1-36.0 33.1-37.9 30.3-34.5
29-255 22.6-28.0 25.3-30.1
23.8-33.0 24.2-34.3 23.5-37.3
42.6-48.4 46.5-50.3 47.9-52.7
10.2-13.0 10.5-13.3 10.8-15.1
23.4-27.1 24.2-28.] 23.8-30.7
47.3-53.4 46.1-53.4 45.2-49.2
26.0-31.1 24.7-30.5 29.2-36.2

B. simplex

2+8+18
total: 28
II, 21-22
total: 23-24 (24)
0-I, 8-9
total: 9
1,6+7,1

I, 1,4
11-12 (12)
5)

91/5-1 11/5 (111%)
27-28 (28)

20 (20)
9-10 (10)

1321-1372
66.7-69.8
2322250
17.8-19.3
SES
34.6-37.2
29.3-32.2
28.1-28.7
48.3-50.0
11.2-11.8

TABLE |. Meristic and morphometric data of the Betta picta species group

564 TAN HEOK HUI & MAURICE KOTTELAT

REMARKS. Betta picta 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. picta 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 picta: 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. | 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 11034, 5 ex.; Jambi: Danau Pinang, lake
connected to Sungai Pijoan, | boat hour upstream of Pıjoan (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, 11 ex., 15.8-
26.5 mm SL, CMK 11136, 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 11071, I 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°35.07S
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. picta 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. picta) and distinct caudal transverse bars in female (vs. absent in B. taeniata and
B. simplex, faint or absent in B. picta). Other characters distinguishing B. falx are
listed under Affinities.

BETTA PICTA AND B. FALX 565

DESCRIPTION. General appearance as shown in Figs. 2-3; meristics and morphometrics
of the B. picta 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 preorbital 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 preorbital 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 falx 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. picta 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 vittata. 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 11-12th 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 91/3, vs. 91/3-11!/, mode 111/:); 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. picta 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. Sa and Sb); thick preorbital
black stripe (vs. narrow); fewer modal dorsal-fin rays (8 vs. 9); fewer modal lateral
scales (27 vs. 28); dorsal fin origin above 11-12th 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. picta 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. picta group is presently known from Lampung, the
southernmost province of Sumatra. Betta falx might occur there, but B. picta 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. picta and B. falx are apparently
closely related, their niche preference, however, is quite marked. Betta picta 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. picta 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 Isbrücker (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 THH’s 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.

EMBRADURE EITED

BLEEKER, P. 1850. Bijdrage tot de kennis der visschen met doolhofvormige kieuwen van den
Soenda-Molukschen Archipel. Verhandelingen van het Bataviaasch Genootschap van
Kunsten en Wetenschappen 23: 1-15.

BLEEKER, P. 1879. Mémoire sur les poissons à pharyngiens labyrinthiformes de l'Inde
archipélagique. Natuurkundige Verhandelingen van de Koninklijke Nederlandsche
Akademie van Wetenschappen 9: 1-56.

CRACRAFT, J. 1989. Speciation and its ontology: the empirical consequences of alternative
species concepts for understanding patterns and processes of differentiation, pp. 28-59.
In: OTTE, D. & ENDLER, J. A. (eds). Speciation and its consequences. Sinauer Asso-
ciates, Sunderland, MA.

CUVIER, G. & VALENCIENNES, A. 1846. Histoire naturelle des poissons, tome dix-huitieme.
Levrault, Paris, xix + 505 pp., pls. 520-553.

GUNTHER, A. 1861. Catalogue of the Acanthopterygian fishes in the collection of the British
Museum, vol. 3. British Museum, London, xxv + 586 pp.

KOTTELAT, M. 1984. Revision of the Indonesian and Malaysian loaches of the subfamily
Noemacheilinae. Japanese Journal of Ichthyology 31: 225-260.

568 TAN HEOK HUI & MAURICE KOTTELAT

KoTTELAT, M. 1994. Diagnoses of the two new species of fighting fishes from Thailand and
Cambodia (Teleostei: Belontiidae). /chthyological Exploration of Freshwaters 5:
297-304.

KOTTELAT, M. 1995a. The fishes of the Mahakam River, East Borneo: an example of the
limitations of biogeographic analyses and the need for extensive fish surveys in
Indonesia. Tropical Biodiversity 2 (3 [1994 (1995)]): 401-426.

KOTTELAT, M. 1995b. Systematic studies and biodiversity: the need for a pragmatic approach.
Journal of Natural History 29: 565-569.

KOTTELAT, M. 1997. European freshwater fishes. An heuristic checklist of the freshwater fishes
of Europe (exclusive of former USSR), with an introduction for non-systematists and
comments on nomenclature and conservation. Biologia, Bratislava, Section Zoology 52
(Suppl. 5): 1-271.

KOTTELAT, M. & NG, P. K. L. 1994. Diagnoses of five new species of fighting fishes from
Banka and Borneo (Teleostei: Belontiidae). /chthyological Exploration of Freshwaters
5: 65-78.

KOTTELAT, M., WHITTEN, A. J., KARTIKASARI, S. N. & WIRJOATMODIO, S. 1993. Freshwater
fishes of Western Indonesia and Sulawesi. Periplus, Hong Kong, 259 pp., 84 pls.

LINKE, H. 1990. Labyrinthfische - Farbe im Aquarium. Ein Handbuch fiir Bestimmung, Pflege
und Zucht. Tetra, Melle, Germany, 174 pp.

MAYDEN, R. L. & Woop, R. M. 1995. Systematics, species concepts, and the evolutionarily
significant unit in biodiversity and conservation biology. American Fisheries Society
Symposium 17: 58-113.

REGAN, C. T. 1910. The Asiatic fishes of the family Anabantidae. Proceedings of the
Zoological Society, London, 1909 [1910]: 767-787, pls. 77-79.

ROBERTS, T. R. 1993. The freshwater fishes of Java, as observed by Kuhl & van Hasselt in
1820-23. Zoologische Verhandelingen 285: 1-94.

TAN, H. H. & KOTTELAT, M. 1998. Two new species of Betta (Teleostei: Osphronemidae) from
the Kapuas basin, Kalimantan Barat, Borneo. Raffles Bulletin of Zoology 46: 41-51.

TAN, H. H. & NG, P. K. L. (in press). Revision of the fighting fish of the genus Betta (Teleostei:
Osphronemidae) from Malaysia, Singapore and Brunei. Raffles Bulletin of Zoology,
Supplement.

WARREN, M. L. 1992. Variation of the spotted sunfish, Lepomis punctatus complex (Centrar-
chidae): meristics, morphometrics, pigmentation and species limits. Bulletin of the
Alabama Museum of Natural History 12: 1-47.

WEBER, M. & DE BEAUFORT, L. F. 1912. Die Fische, pp. 522-541, pls. 11-12. In: Maass, A.
(ed.). Durch Zentral-Sumatra, Vol. 2. Behr, Berlin & Leipzig.

WEBER, M. & DE BEAUFORT, L. F. 1922. The fishes of the Indo-Australian archipelago, vol. 4.
Brill, Leiden, xiii + 410 pp.

Witte, K.-E. & SCHMIDT, J. 1992. Betta brownorum, a new species of anabantoids (Teleostei:

Belontiidae) from northwestern Borneo, with a key to the genus. /chthyological
Exploration of Freshwaters 2: 305-330.

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 (Neuchätel, Switzerland)!

Sylvie BARBALAT
Institut de Zoologie, Emile Argand 11, CH-2007 Neuchätel, Switzerland.

Importance of forest structures on four beetle families (Col.: Bupre-
stidae, Cerambycidae, Lucanidae and phytophagous Scarabaeidae) in
the Areuse Gorges (Neuchätel, 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).

| 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 Neuchatel (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 m2, 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

Les Grattes
880 m @7

Les Tablettes !
288m ©?

SEHR, retereules ®8
È 830m &
e°

57 m Lake Neuchatel

7 Champ-du-Moulin 7

1
x Creux du Van _ -
i a

watershed
railway
road

river
locality
trapping site

Fic. 1. Study area. After BARBALAT (1997)

_ station locality coordonnates altitude expo _ forest type cover habitat type edge type
“i Gab 556125204250 695m S oak 80% edge natural
Colombier, Chanet 555600 202950 555m beech 40% artificial clearing clean
Chambrelien, Plan du Bois 553250 202700 670m SE mixed 60% artificial clearing clean
Boudry, Chanet 553000 201150 550m oak 30% artificial clearing clean
Chambrelien, Chassagne 552000 201900 725m SE mixed 40% artificial clearing clean
Chambrelien, Chassagne 551900 202250 770m SE mixed 60% artificial clearing clean

Rochefort, Les Grattes 552470 204070 880m SE mixed 40% edge natural
Fretereules 549050 201600 860m SE beech 90% edge natural
Brot-Dessous 547600 200950 890 m S beech 90% edge clean
Brot-Dessous 547850 200900 790 m S beech 50% natural clearing natural
Chambrelien, Bois-Devant 553500 202450 620 m mixed 70% artificial clearing clean
Rochefort, Chaumes 551375 202625 840 m 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).

SYLVIE BARBALAT

Author Distribution Level Habitat Main host plants Site

Agrilus angustulus
Agrilus biguttatus
Agrilus cyanescens
Agrilus laticornis
Agrilus olivicolor
Agrilus sulcicollis
Agrilus viridis

Anthaxia helvetica
Anthaxia nitidula
Anthaxia quadripunctata
Anthaxia salicis
Anthaxia similis
Chrysobothris affinis
SCARABAEIDAE
Amphimallon atrum
Cetonia aurata

Hoplia argentea
Omaloplia ruricola
Phyllopertha horticola
Rhizotrogus aestivus
Serica brunnea

Trichius fasciatus
LUCANIDAE
Platycerus caprea
Platycerus caraboides
Sinodendron cylindricum
CERAMBYCIDAE
Agapanthia villosoviridescens
Agapanthia violacea
Alosterna tabacicolor
Anaglyptus mysticus
Anastrangalia dubia
Anastrangalia sanguinolenta
Anoplodera sexguttata
Arhopalus rusticus
Cerambyx scopolii
Clytus arietis

Clytus lama

Cortodera femorata
Corymbia rubra
Dinoptera collaris
Gaurotes virginea
Grammoptera abdominalis
Grammoptera ruficornis
Leiopus nebulosus
Leptura maculata
Molorchus minor
Obrium brunneum
Oxymirus cursor
Pachytodes cerambyciformis
Parmena balteus
Phymatodes testaceus
Phytoecia cylindrica
Pidonia lurida
Plagionotus arcuatus
Pogonocherus fasciculatus
Pogonocherus hispidulus
Prionus coriarius
Pseudovadonia livida
Pyrrhidium sanguineum
Rhagium bifasciatum
Rhagium inquisitor
Rhagium mordax
Stenocorus meridianus
Stenostola dubia
Stenurella bifasciata
Stenurella melanura
Tetropium castaneum

(111.,1803) eurosiberian, NA col
(F.,1777) holomediterranean col
Ratzb.,1837 european col-mon
(111.,1803) european, ME col
Kiesw.,1857 eurosiberian col
Lacord.,1835 eurosiberian col
(L.,1758) eurosiberian col-mon
Stierl.,1868 mountain col-sub
(L.,1758) holomediterranean col-mon
(L.,1758) mountain mon-sub
(F.,1777) holomediterranean col
Saund.,1871 mountain mon-sub
(F.,1794) eurosiberian, NA col-mon
(Hbst.,1790) W. european col-mon
(L.,1761) palearctic col-sub
(Poda,1761) C. and S. european col-sub
(F.,1775) C. european col-mon
(L.,1758) eurosiberian col-sub
(01.,1789) pontomediterranean col-mon
(L.,1758) eurosiberian col-mon
(L.,1758) eurosiberian col-mon
(Deg.,1774) C. european mon-sub
(L.,1758) C. european col-mon
(L.,1758) eurosiberian col-mon
(Deg.,1775) eurosiberian col-mon
(F.,1775) pontomediterranean col-mon
(Geer,1775) palearctic col-mon
(L.,1758) european col-mon
(Scop.,1763) mountain mon-sub
(L.,1761) C. and E. european mon-sub
(F.,1775) european col
(L.,1758) holarctic col-mon
Füssl.,1775 european, NA col-mon
(L.,1758) european, ME col-mon
Muls.,1847 mountain mon-sub
(F.,1787) C. and E. european col-sub
(L.,1758) palearctic col-sub
(L.,1758) palearctic col-mon
(L.,1758) boreo-alpine mon-sub
(Steph.,1831) C. and S. european, ME co
(F.,1781) european, ME co
(L.,1758) european col-mon
(Poda,1761) european, ME col-mon
(L.,1758) palearctic mon-sub
(F.,1792) european, ME mon-sub
(L.,1758) E. european, ME mon-sub
(Schrk.,1781) C. european, ME col-mon
(L.,1767) W. mediterranean co
(L.,1758) holarctic col-mon
(L:;1758) palearctic col-mon
(F.,1792) mountain mon-sub
(L.,1758) european, ME, NA co
(Deg.,1775) palearctic col-sub
(Pill.Mitt.,1783) european col-mon
(L.,1758) palearctic co
(F.,1776) palearctic col
(L.,1758) european, NA col
F.,1775 european col-mon
(L.,1758) holarctic col-mon
(Deg., 1775) eurosiberian col-mon
(L.,1758) eurosiberian co)
(Laich.,1784) C. and N. european col-mon
(Müll.,1776) palearctic col
(L.,1758) palearctic col-mon
(L.,1758) palearctic mon-sub

broadl/tran
broadl/tran
broadl/tran
broadl/tran
broadl/tran
broadl/tran
broadl/tran
conif/tran
tran/orch
conif/tran
broadl/tran
conif/tran
broadl/tran

open/tran
broadl/tran
open/tran
open
open/tran
open/tran
broadl/tran
broadl/tran

broadl
broadl
broadl

open/tran
open
broadl
broadl/tran
conif/tran
conif/tran
broadl/tran
conif
broadl/orch
broadl/tran
conif/tran
conif
conif/tran
broadl/tran
conif/tran
broadl/tran
broadl/tran
broadl/mix
broadl/tran
conif/mix
conif/mix
conif/mix
broadl/tran
broadl/tran
broadl
open
conif/mix
broadl/mix
conif/tran
broadl/mix
broadl/mix
open/tran
broadl
conif/mix
conif/mix
broadl/mix
broadl
broadl/tran
mix/tran
broadl/mix
conif

Quercus
Quercus
Lonicera
Quercus
Corylus, Carpinus
Quercus
broadl
conif
Frunus
conif
Quercus
conif
broadl

broadl

Poaceae, div
div

div

broadl

broadl
broadl
broadl

Asteraceae, Apiaceae
Dipsacaceae
broadl
broadl

conif

conif
Quercus
conif

broadl
broadl

conif

Picea
conif/broadl
broadl

conif
Quercus, Castanea
broadl
broadl
broadl

conif

conif
conif/broadl
broadl/conif
broadl
broadl
Apiaceae
Picea, Fagus
Quercus
conif

broadl
broadl/conif
soil
Quercus
conif/broadl
conif/broadl
broadl/conif
broadl
broadl
broadl/conif
broadl/conif
conif

Distribution: ME = Middle East, NA = North Africa, C. = central, E. = east, N. = north, S. = south, W. = west;

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

3,4,7

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

3,6,11
1,2,3,4,8,9,11

7,8,9,10

3,4,7,8,11

4,5,8,9,10

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

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

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

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

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

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

3,5,6,12

1,2,12

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

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

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

3,4,7,9

7,10

1,4,11

5,6,12

3,4

3,5

1,3,7,12

5,6,7,12

2,3,6,11

4,5,8,10,12

5,6,8,9

8,10,12

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

Habitat: broadl = 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, broadl = 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

St.1 Su4 Still St.3 St.2 St.5 St.6 St.12 St.7 St.8 St.10 St.9| total |

A. villosoviridescens
A. rusticus
A. olivicolor

SoSe

PIN D WwWAIAN WwW

A. sanguinolenta
C. rubra

P. cerambyciformis

>
I © NY O0 D = mm WwW

(A
je}
(>)

wore
Re nm mr LU 00 SW

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 analysıs 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 (< Im, 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 | 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

A. salicis à

Oak stand ®

G. rufi o P. arcuatus
o S. bifasciata o G. ruficornis

e Natural edge

o A. nitidula

A. sexguttata o

St. 4

Sus a A. atrum
3 M. minor A. angustulus
SE u ° P.cylindrica c a
St9 = St.10 = P. horticola o EURE ©
S. brunnea o © H. argentea C. femorata
S. dubia , © P. caprea Parestaceus BA. laticornis
D. collaris © P.sanguineum , © o C. affinis

Axis I A. tabacicolor ° oT. fasciatus

Altitude | maculata R. mordax

S. meridianus A. helvetica o | o D Chemin

*Beech stand PI o P. coriarius
A. sanguinolenta © P. caraboides
o o C. rubra

o À. quadripuncata

A. viridis Artificial clearing
C. lama o © ® À. dubia “Stell

R. bifasciatum ° : i
Mixed forest St. 3

A. olivicolor

oA. bigutattus
9 A. sulcicollis

O. brunneum o
L. nebulosus o
P. balteus 0

G. Yirginea _ A. similis a St.2 |
St. 12 = A mysticus È R, inquisitor
P. hispidulus
Li U
St. 6 St

Fic. 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 | 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, 11 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
cylindrica, 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 cylindrica 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 cylindrica. 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.

REFERENCES

ALLENSPACH, V. 1970. Coleoptera: Scarabaeidae, Lucanidae. Insecta helvetica, Catalogus 2,
186 pp.

BARBALAT, S. 1995. Efficacité comparée de quelques méthodes de piégeage sur certains Colé-
opteres saprophages ou xylophages et influence de l’anthophilie sur le résultat des
captures. Bulletin de la Société neuchäteloise des Sciences Naturelles 118: 39-52.

IMPORTANCE OF FOREST STRUCTURES ON BEETLES 579

BARBALAT, S. 1996: Influence de l’exploitation forestiere sur trois familles de Coléopteres lies au
bois dans les Gorges de l’Areuse (Canton de Neuchatel, Suisse). Revue suisse de
Zoologie 103 (2): 1-12.

BARBALAT, S. 1997. Faunistique de 47 Cérambycides (Col. Cerambycidae) capturés dans les
Gorges de l'Areuse (Neuchatel, Suisse). Bulletin de la Société neuchäteloise des Sciences
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 (Neuchatel, 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.

DaJ0z, R. 1980. Ecologie des insectes forestiers. Bordas, Paris, 489 pp.

GEISER, R. 1984. Rote Liste der Käfer. In: BLAB, J., NOWAK, E., TRAUTMANN, W. & SUKOPP, H.
(eds). Rote Liste der gefährdeten Tiere und Pflanzen in der Bundesrepublik Deutschland.
Naturschutz Aktuell 1. Kilda. Greven, 270 pp.

GUTOWSKI, J. 1985. Distribution of cerambycid beetles (Coleoptera: Cerambycidae) in various
forest site types in Bialowieza Primeval Forest. Parki Narodowe i Rezerwaty Przyrody
6 (1): 77-94. (in Polish)

GUTOWSKI, J. 1986. Species composition and structure of the communities of longhorn beetles
(Col., Cerambycidae) in virgin and managed stands of Tilio-Carpinetum stachysetosum
association in the Bialowieza Forest (NE Poland). Journal of Applied Entomology 102:
380-391.

GUTOWSKI, J. 1995. Changes in communities of longhorn and buprestid beetles (Coleoptera:
Cerambycidae, Buprestidae) accompanying the secondary succesion of the pine forests of
Puszcza Bialowieska. Fragmenta Faunistica 38 (20): 389-409.

HARTMANN, K. & SPRECHER, E. 1990. Ein Beitrag zur Insektenfauna des Arlesheimer Waldes
unter Berücksichtigung der Holzbewohnenden Käfer. Tätigkeitsberichte der Naturfor-
schenden Gesellschaft Baselland 36: 75-124.

Horion, A. 1958. Faunistik der mitteleuropäischen Käfer. Band 6: Lamellicornia. Selbstverlag,
Uberlingen, 343 pp.

Horton, A. 1974. Faunistik der mitteleuropäischen Käfer. Band 12: Cerambycidae - Bockkäfer.
Selbstverlag, Uberlingen, 228 pp.

KocH, K. 1992. Die Käfer Mitteleuropas, Ökologie. Band 3, Goecke & Evers, Krefeld, 389 pp.

Louse, G. A. & LUCHT, W. 1992. Die Käfer Mitteleuropas. Band 13, Goecke & Evers, Krefeld,
375 pp.

SPEIGHT, M. 1989. Les invertébrés saproxyliques et leur protection. Collection sauvegarde de la
nature 42, Strasbourg, 77 pp.

STARZYK, J. 1976. Grouping of cerambycids (Coleoptera, Cerambycidae) on the background of
types of environnement of the Niepolomice Forest near Krakow. Acta Agraria et
Silvestria, Series Silvestris 16: 131-152. (in Polish)

STARZYK, J. 1977. Influence of the stand age upon the quality of the composition and the
numerousness of the timber beetles (Col., Cerambycidae) in the Niepolomice Forest. Acta
Agraria et Silvestria, Series Silvestris 17: 117-135. (in Polish)

STARZYK, J. 1979. Cerambycidae communities (Col. Cerambycidae) occurring in various phyto-
sociological forest types of Niepolomice Forest near Krakow. Journal of Applied Ento-
mology 88: 44-55.

STARZYK, J. & Witkowski, Z. 1981. Changes of the parameters describing the cambio- and
xylophagous insect communities during the secondary succession of the oak-hornbeam

association in the Niepolomice Forest near Krakow. Journal of Applied Entomology 91:
525-533.

580 SYLVIE BARBALAT

TER BRAAK, C. J. F. 1986. Canonical correspondence analysis. A new eigenvector technique for
multivariate direct gradient analysis. Ecology 67: 1167-1179.

TER BRAAK, C. J. F. 1988a. Partial canonical correspondence analysis (pp. 551-558). In: Block,
H. H. (ed.). Classification and related methods of data analysis. North Holland Press,
Amsterdam.

TER BRAAK, C. J. F. 1988b. CANOCO - an extension of DECORANA to analyze species-envi-
ronment relationships. Vegetatio 75: 159-160.

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-4051 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. favosus. 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-4130-1 (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 Investigaciön de Recursos Biolögicos Alexander von Humboldt, Villa de
Leiva, Santafé de Bogota, Colombia. Courtesy Fernando Fernandez-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 Löbl.

MIZA Museo del Instituto de Zoologia 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 Sao Paulo, Brazil. Courtesy of Prof. Carlos
Roberto Ferreira Brandao.

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

Fıc. |
Dominican amber. Specimen Do-4130. Profile (top); dorsal view (bottom).

The measurements and Indices I used are those defined by BROWN (1975); these

and other measurements defined here are:

HW
BE
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 pronotal 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

PeW
HFeL

HFeW
HTiL

HTiW
HBaL

HBaW

CI
SI
HFel
HTil
HBal

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

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

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

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

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

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

(Cephalic Index): HW/HL x 100

(Scape Index): SW/SL x 100

(Hind Femur Index): HFeW/HFeL x 100

(Hind Tibia Index): HTiW/HTiL x 100

(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 (MADDISON & MADDISON
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 IT 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 RI. Mf2 and r-m medially interrupted. Mf4 and CuAl 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

IRA

WIN
Ne

DD
Nn pw

IS)
CA

ne)
=

MARIA L. DE ANDRADE

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

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 "?".

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).

Worker and gyne. Base of the mandibles not angulate laterally (0), or angulate laterally
(1).

Worker and gyne. Occiput high (Fig. 17) (0), or low (Figs. 8, 31) (1).

Hypostomal bridge narrow (Fig. 9) (0), or broad (Fig. 32) (1).

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).

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 "?".

Worker and gyne. Ventral process of the petiole different shape but never triangular (0),
or broad, triangular (1).

Worker and gyne. Dorsum of the petiole with more than 7 long pointed hairs (0), or with
at most 3 long, pointed hairs (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).

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).
Worker and gyne. Mid tibiae with no or with one pectinate spur (0), or with two pectinate
spurs (1).

Worker and gyne. Fore basitarsi longer than mid basitarsi (0), or fore basitarsi at most as
long as the mid basitarsi (1).

Worker and gyne. Fore basitarsi shorter than hind basitarsi (0), or fore basitarsi as long as
or longer than hind basitarsi (1).

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).

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).

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).

Gyne. Compound eyes largely behind the mid line of the head (0), or on the mid line of
the head (1).

Gyne. Scutellum smooth, foveolate, or with very thin striae (0), or scutellum with very
thick striae (1).

Gyne. Hind femora Index < 48 (0), or > 50 (1).

Male. Frontal carinae subparallel (0), broad anteriorly and narrower posteriorly (1), or
strongly broad anteriorly and touching each other posteriorly (2).

Male. Antero-median border of the clypeus convex (0), or straight (1).

Male. Anterior face of femora densely covered by hairs (0), or with only few hairs (1).
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).
Male. Hypopygium not strongly constricted distally (Figs. 23, 27, 34) (0), or strongly
constricted distally (Figs. 7, 11, 13) (1).

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

72T371#15[6 1718 [8 [10] 14] 12] 13] 14] 15] 16] 17] 18] 19] 20] 24] 22] 23] 24] 25] 26}

C. boliviae TER OT DIR 0202070 0 0 00 0071 0 2 0
C. brasiliensis il Wee Oa D Oe. 10010 10 1 0 16200 (0) Oi 4
C. brevitarsus 11000 1 2 0&1 0 1&2 1 00 Du ORMONI 0822250
C. darlingtoni 1.3°0°0 © J 2.9270 1% 0 0 2B} OO? 2 4
C. escobari 17 OM O m I 2 ay oO 0 4 0 0 2 1 vd D PNR?
C. godmani 110101040021 0 0 01 002 Oele
C. longiceps 1.0 OT CONO -2 4 CA 01 Qo oe PR
C. meinerti 1 @-@ Y U CIO O- 2.4 0 1 0 1 Oele (0): 4
C. striatus OY Ut Oo ONTO a oO 1 0 10 i =) 1 -@ 4
C. whymperi OT hae OF yoo OW Oo 1 0 1 0 150250 1
1 C. antillanus 120) O1 Oui 20.0.2 4 0 1 0 1 00? Ct 2

11 OO Ta @ a4 0 0 2 0 OO? Le 0 00

200202070077. OF 10) 0! 0 0 0 0 0 0 0 0 0 0 0

2 020 OO O42 0.0.0.0 0 0 0 0 00 ? D. Pot

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 | 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 (/ongiceps, 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), nec gyne
from Lima = whymperi (nec MAYR 1870). Misidentification.

590 MARIA L. DE ANDRADE

Cylindromyrmex brasiliensis Emery, 1901: 53. Worker and male (Santa Catarina). Original
description. Type locality: Brazil. Type material: 3 syntype workers labelled: "S.
Catharina, Schmalz, typus", in MCSN: 1 syntype worker labelled: "Bresil, Mayr, typus,
Cylindromyrmex brasiliensis Em (striatus Mayr 1887)", in MCSN, examined.

Cylindromyrmex brasiliensis Emery, BORGMEIER 1937: 218. Gyne.

Cylindromyrmex brasiliensis Emery, JAFFE 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 | — du en

Fic. 3

C. brasiliensis Emery. Worker from Capào Bonito, S40 Paulo, Brazil. Apical funicular joint with
spine-like setae.

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 59]

Fic. 4

Fic. 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.

MARIA L. DE ANDRADE

592

LO)
N
©)
N
O
(©)
Oo
oO

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 593

0.5mm

Fic. 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.

Fic. 6. C. brasiliensis Emery. Worker from Capäo Bonito, Sao 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. Pronotal 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 pronotal 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
pronotal 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; HBal 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]. SAo PAULO: Piracicaba, Escola Superior de Agricultura
"Luiz de Queiröz", 25.X.1974, 10 workers, 1 gyne, E. Berti Filho [MZSP, MPEG]; Agudos,
VIII.1958, 5 workers, R. Mueller [MZSP]; Barueri, 03.VI.1967, 88 workers, K. Lenko [MZSP];
Botucatu, 13.X.1987, 1 worker, L. C. Forti & I. M. P. Rinaldi [MZSP]; Sao Paulo, Butantan 23-
31.VII.1969, 13.VIII.1969, 31.VII.1973, 5 workers, L. Travassos Filho [MZSP]; Ilha de Sao
Sebastiäo, VII.1987, 500 m, 13 workers, C. R. F. Brandäo [MZSP]; Salesöpolis, Estaçäo
Biolögica de Boracéia, 3-5.V.1996, 1 worker, C. R. F. Brandäo et al. [MZSP]; Capäo 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 [NHMWI]; 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, | 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), nec worker and male (=brasi-
liensis). Nec 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. Nec 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 MARIA L. DE ANDRADE

| 00007981 …

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 599

i... L.A | °—[REM-Labor
e 100 BO. Uni Basel

Fic. 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.

Fic. 8. C. whymperi (Cameron). Worker from Ecuador. Head in full dorsal view (top), body in
full dorsal view (middle), body in profile (bottom).

MARIA L. DE ANDRADE

600

oO)
ice)
oO)
N
O
[æ)
oO
[@)

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 601

0.5mm

Fic. 11

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.

Fic. 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 0.09-0.11; 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 CYLINDROMYRMEX 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, 24.11.1963, 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. Quirös [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, | 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, 25-27.11.1970, 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,
111.1938, 1 gyne, H. Hanson & W. H. W. Komp [MCZC]. Guayas: Guayaquil, X.1922, 1 worker,
F. X. Williams [MCZC]; same locality, | 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. Pätapo, 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 (EGG). 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
NHMwW 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. Nec 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).
Nec 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). Nec 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.

606 MARIA L. DE ANDRADE

Uni Basel

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 607

Fic. 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.

Fic. 12. C. striatus Mayr. Worker from Parad, Rio Curuä-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 Curuä-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

0000799

MARIA L. DE ANDRADE

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 611

| 00007960 | | ‘Unt Basel

612 MARIA L. DE ANDRADE

0.5mm

Fic. 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.

Fic. 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 613

Cylindromyrmex boliviae Wheeler Figs 4, 14-16

Cylindromyrmex boliviae Wheeler, 1924: 104, fig. 20. Gyne. Original description. Type locality:
Mapiri, Bolivia. Type material: I 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). Nec 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 tarsı 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-
0.11; 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 10.8-11.3.

Material examined. COLOMBIA: CUNDINAMARCA: Medina, Quebrada Ardita, 1475 m,
28.11.1997, 1 gyne, F. Escobar [IAVH]. VENEZUELA: ARAGUA: Rancho Grande, 17.VII.1972,
1 gyne, N. A. Weber [MCZC]; same locality, 1100 m, IV.1987, 1 gyne, C. Bordon [MIZA];
same locality, 1200 m, 15.V & 22.V1.1987, 3 males, C. Bordon [MIZA]. PERU: Lima:
Calanga, 1 male, Staudinger [MCSN]. BOLIVIA: Songo, 1 male [MCSN]. La Paz: Mapiri, |
gyne (holotype), Staudinger [MCZC].

Discussion. Boliviae 1s 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
11 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.

616 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. Nec EMERY

1901. Misidentification.

Holotype: Worker labelled: "Colombia, Nariño, Ricaurte La Planada, 1°17' N 78°15' W,
1800 m, interior bosque, bmh-PM, Col: F. Escobar", in IAVH.

Derivatio 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.

Fic. 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

618 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 619

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.

MARIA L. DE ANDRADE

620

‘(WONOA) Mara [esIop [ny ur peau ‘(do7) apyoid ur Apog ‘Joquie uesıumwog WO} Suo, "SpeIpuy ap sn14199]2 I

81 SIN

ULI |

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 621

Legs. Femora slightly inflated. Mid basitarsi with parallel sides. Hind basitarsı
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: | gyne (holotype) without reference number
[MCZC].

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

Cylindromyrmex 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.V1.1936, 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-

Fic. 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).

623

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX

REM-Labor
Uni Basel

00008089

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: "Bresil, 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. Pronotal 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

mn
3
A
zZ
<
in
A
SI
S
Z
<
=

627

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX

REM-Labor
Uni Basel

1 mm

00007986

628 MARIA L. DE ANDRADE

00007958

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 629

ÉIG 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.

Fic. 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).

Fic. 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).

Fic. 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

0.5mm

Fic. 24

C. brevitarsus Santschi. Male from Butantan, Sao 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 631

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 basitarsı.

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: ARAGUA: 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-4.1II.1994, 1 gyne, 1 male, P.
Hibbs [LACM]. PERU: APURIMAC: no further locality, 14.VIII.1962, 1 worker, M. Dourojeanni
[USNM]. BRAZIL: PARA: Serra Norte, Estacäo. Manganés, 5-9.IX.1983, 1 gyne, F. F. Ramos
[MPEG]. Goras: Jataî, XII.1972, 1 gyne, F. M. Oliveira [MZSP]. BAHIA: Ilhéus, Reserva
Botanica, CEPEC, 23-24.1V.1987, 1 male [CPCC]. Mato Grosso: Sinop, 55°37' W, 12°31' S,
X.1974, 1 male, M. Alvarenga [MZSP]. Rio DE JANEIRO: Ilha 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, VIII.1974, 1 gyne, F. M. Oliveira [MZSP]. SAo PAULO: Sao 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 Jardım (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, pl. 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

MARIA L. DE ANDRADE

634

‘QUSLI) MOIA fesıop [ny ur peasy ‘(1J9) apyoud ur Apog eweurg “ınbruryy ULI]OA WO] SUÂO '[210g IUDWPOB ‘I
SC PIT

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 635

iii

| 00007963 || — 1mm Arabo:

| Uni Basel

Fic. 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 CYLINDROMYRMEX 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 /ongiceps 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-4130-1 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.

MARIA L. DE ANDRADE

640

‘(WONOI) MATA [ESIOp [[n} ur peau ‘(do7) apyoud ur Apog ‘Joquie UPOIUIOG Wo JUAN) 'apeapuy ap SAUPJJHUD ‘I
80 PIT

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;
SW 0.15; WL1.68; PeL 0.524 PeW 0.56; HFeL 0.53; HFeW 0.24; HTiL 0.44; HTiW 0.16;
HBaL 0.25: HBaW 0.07; CI 77.8; SI 46.9; HFel 45.3; HTil 36.4; HBal 28.0.

Material examined. Dominican amber: | gyne (reference number Do-4130) [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 Andre, 1892: 47. Worker. Original description. Type locality: Brazil.

Type material: 1 worker labelled: "Bresil, Type, Museum Paris, Collection Ernest Andre,

1914, longiceps Andre", 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

Fic. 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

00008084

~ REM-Labor
1 mm Uni Basel

644 MARIA L. DE ANDRADE

REM-Labor
30 um Uni Basel

00008083
Fic. 30
C. longiceps Andre. 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 CYLINDROMYRMEX 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-
0.11; 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 0.11; CI 64.1; SI 42.8; HFel 52.0; HTil 36.0; HBal 25.0.

Material examined. BRAZIL: no further locality, 1worker (holotype), MNHN. SAo
PAULO: Sao Paulo, 5.1.1974, 1 gyne, R. L. Araujo [MZSP]. Rio DE JANEIRO: Rio de Janeiro,
25.VIIL.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. Nec 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. 117 & 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

| 00007966 a

Fic. 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 647

00009891 | — 100 um .

ni Basel

Fic. 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 scapes1/2 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

A =

REM-Labor :
Uni Basel |

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 649

0.5mm

r—21z_x1ÀjHzkzmmux=

Fic. 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 VII.

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.

Fic. 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 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.
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; HTil 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 651

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 Mayrıan and
parapsidal furrows superficially marked. Impar Mayrıan 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, | gyne, D. H. Janzen & I.
Gauld [BMNH]; Hambrug Farm, Santa Clara Province, 23.1V.1926, 1 worker, 2 gynes, 2 males,
F. Nevermann [USNM]; same locality, Reventazön 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.V1.1935, 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 [[AVH,
MZSP]; same locality, 1 worker, in termite nest, [WEMC]; same locality, III. 1V.1949, Zetek, 1
male [USNM]. VENEZUELA: ZuLta: 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, 23.111.1938, 1 male, G. Vivas-Berthier
[WEMC]. BoLIvar: Las Trincheras, in altem Baume, 4 workers (syntypes of meinerti), Meinert
[MHNG, MCZC, MCSN]. BRAZIL: Amazonas: Ilha de Curari, varzea, 2.1X.1976, 1 gyne, J.
Adis [LACM].

a a
8 5 2 8 § 0000 ae
a 2 S =
2 È S è S ® N = a D 3 S 8 5
= 2 2 2 E S 8 È = S E = > =
® S G as > = OS G © E 8 (SÌ < ©
S Se Ss = < Si D = I g E S E
& 2 5 ü ES a © & Ss D © G S
ao < ©) ©) ©) © ©) + ©) ©) È) + ©) ©)
116: 1->0
|
F =
2: 120 18150 601
[54 0-1 DI_0=1]
1270-51)
B: 1->0
1: 6->1] GT: O>II (15: 0->1]
19: 0->1]
4: 0->1 =
57-159 ET 0-54
19: 0->1] 8: 0->1] 22: 1->2
25: 2->1]
14: 0->1
16: 0->1]
17: 0-51) 15051
(20; 0->1]
12: 0->1
(26: 0->1] (p>: 0->1]
25: 0->2
Fic. 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

U) PSG EE 2)
(Cb) > a SSR SE
= ® ccs È =
SPILLESESSLE LI
De a eS wo ge a See ce
> Sl eS oo Ba 2
@ 2S) SS) Bug Sten eyes SS
PMO ROLO OOO OMO,
99 30 88
53 92
74 88
100
68
77
99
Fic. 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

% brasiliensis
à M striatus
A whymperi

Fig. 37
Distribution of the species of the striatus clade.

656 MARIA L. DE ANDRADE

% boliviae

Fic. 38
Distribution of the species of the boliviae clade.

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 657

A brevitarsus
Ÿ &* darlingtoni

SX electrinus

X escobari

Fic. 39
Distribution of the species of the brevitarsus clade.

658 MARIA L. DE ANDRADE

> antillanus
A godmani
X longiceps
EB meinerti

Fic. 40
Distribution of the species of the longiceps clade.

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 659

IDENTIFICATION KEYS

WORKERS

The workers of antillanus, boliviae, electrinus and godmani are not included in

this key because they are not yet known.

ile

Eyes large and convex (2 400 ommatidia). Ocelli present and well
detinedH(lüigsz ON AID) Vent OTE COCR END ach 2
Eyes small or of medium size (> 16 and < 200 ommatidia), flat or
slightly convex. Ocelli absent or represented by superficial pits (Figs 17,

ORDNER TEE EN MARNE BIS TONER ED ae JO FORSE 3
Legs dark orange to light brown. Dorsum of the petiole with thin,
imecularstniae (False 0) BrazilYParaguayız Dramen ee 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

Posterior third of the head dorsum at most with 25 striae. Postpetiole
with 19-25 longitudinal striae. Guatemala, Costa Rica, Galapagos Is.,

Ecuador Penn Bolivia, ChilesBrazil Re mr See Be whymperi
Posterior third of the head dorsum with more than 34 striae. Postpetiole
with about 29-30 striae. Surinam, French Guyana, Brazil............ striatus

Head length (mandibles excluded) ca. 1/3 longer than broad; frons,
slightly more than 1/3 of the head width. Mandibles dorsally flat
(OS DONS Drei ae Sara Sa ee MET IR À 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

dorsalliyiconvers (es AIT M0F20) TERE 0 SACE eer. RE EN IE 6
Frontal carinae not reaching the anterior clypeal border. Pygidium with a
semieireleoMfeethroß similar size. Brazil. pes DENE EEE longiceps

Frontal carinae reaching the anterior clypeal border. Pygidium with a
semicircle of teeth with two larger ones over the sting. Costa Rica,
Panama VenezuelasBrazile san cease tanks er a meinerti
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 Elandmemoralndext (diel) =14 ST FI EINE EIN ee 7
Frontal carinae surpassing the anterior clypeal border. Mandibles with 9-
JOdEniclesaScapelndeA(SD = S7NEUDA RR ON e darlingtoni

Frontal carinae reaching the anterior clypeal border. Mandibles with 6-7
denticles. Scape Index (SI) > 42. Venezuela, Ecuador, Peru and Brazil
PRE SAO! DLL ESC LUN RAR Dili AQU LIS) DITE DE brevitarsus

660

GYNES

MARIA L. DE ANDRADE

The gyne of escobari 1s not included in this key because it is not yet known.

Finst gastric tergite:simoothi a. salat see 0 RRSSAA 2
Kirstgastricitergite sculptured "1" used doa Son a Gee eee I
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

dorsal awithmaximumwidenticles "stat Sain. 2: ee 3)
Legs dark yellowish-orange to light brown. Body striation more irre-
eulargBrazilsbBarasuayarı sad SRE Paced ESS brasiliensis
Legs dark ferrugineous to black with large part of the tibiae yellowish.
Bodysstriationme cular ru. na al A) te beh re 4

Cephalic Index (CI) = 80. Posterior third of the head dorsum at most with
25 striae. Guatemala, Costa Rica, Galapagos Island, Ecuador, Peru,

B'oliviaX@hilesBrazil Kar na Ale sa ante ee whymperi
Cephalic Index (Cl) < 77. Posterior third of the head dorsum with more
than 34 striae. Surinam, French Guyana, Brazil.................... striatus
Frons at most slightly more than 1/3 the head width. Eyes on the middle
ofsthesheadisides..... 1.2.42... ek oa ses a ee oe Le eee 6
Frons at least slightly less than 1/2 or more than 1/2 of the head. Eyes
behindithe middle ofthe headisides tere ERRATE 8

Frontal carinae not reaching the anterior clypeal border. Pygidium api-
cally without a distinct pair of large teeth. Distal border of hind basitarsi
witht spine-like.setae. Brazil Ween: sete Se eee 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. +... ESE ee 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} Li a th sehe CR TT SRE meinerti
Outer apical edge of mid basitarsi with 5 spine-like setae on the outer
face. Cephalic Index (CI) > 77. HFel < 46. Dominican amber...... antillanus

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 2
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(Cl) >83. are Cet athe) Sere eee 9
Coxae and femora black. Mid basitarsi with parallel sides and more than
half longer than the hind basitarsi. Dominican amber.............. electrinus

10.

MALES

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 661

Coxae, femora dark brown. Mid basitarsi broad apically and half as long
asıtheshindibasitarsin 9. PI. E BT UN RS ENT ART EN PARENT LSPA ed o 10
Frontal carinae not surpassing the anterior clypeal border. Mandibles
with 6-8 small denticles. Femora yellowish to light brown. Venezuela,

Beuadorsbensand- Brazil Bat Peu PAR I 9 brevitarsus
Frontal carinae surpassing the anterior clypeal border. Mandibles with 9-10
smallrdentielesy Eemorardarksbrown Cuba Fe ee darlingtoni

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.

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
ASA) ee U O NEE 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

(LUIS J Dal ide Re 3
Total length (TL) > 9.5 cm. Mesosoma massive. Petiole with traces of

striae only anteriorly. Postpetiole smooth. Costa Rica, Panama, Ecuador
e ATE ORE AS ote Te Er Rem DATS godmani
Total length (TL) < 8.5 cm. Mesosoma elongate. Petiole entirely striate.
Postpetiole superficially striate. Costa Rica, Panama, Venezuela, Brazil .
RA SCARS AC NARRA, STARB ee SE DPR hse RS PE AGNO meinerti
Head and basal face of the propodeum with thick striae (Fig. 10),
sometimes with thick foveae between the striae. Hypopygium smooth or

ine lyadenticulate beiweenstheidistalapodemes,.. um. nun ea 4
Head and basal face of the propodeum with thinner striae (Figs 15, 22).
Hypopygium with a simple, umpair, median projection between the apo-

GETS SALE PAL Le SPER TER IONI GE ORTEN REN Be A LA ND 5
Anterior clypeal border slightly convex. Scape Index (SI) < 53.1. Hypo-
pygium finely denticulate between the distal apodemes (Fig. 7b). Brazil

An CRB ATASW As 8 E SMe ae Teme EL TINE A, I Me brasiliensis
Anterior clypeal border straight (Fig. 10). Scape Index (SI) > 57.1.
Hypopygium smooth between the distal apodemes. (Figs 11b, 13b).
Guatemala, Costa Rica, Galapagos Island, Ecuador, Peru, Bolivia, Chile,

| SRV lo ANTE te ape SOT eater Per whymperi
SurtamAErenchkGuyanayBrazilieerer ee OE striatus

662 MARIA L. DE ANDRADE

Si Coxae black with the remaining parts of the legs yellow to light brown.
Ventral border of the aedeagus with at least 42 denticles. Venezuela,
Ecuadomeerurandbraziless SERIO SEIN NA 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#RerulandiBolivia Wi, 2 ner. ya atten eee 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.

REFERENCES

ANDRADE, M. L. DE. First description of fossil Acanthostichus from Dominican amber (Hyme-
noptera: Formicidae). Mitteilungen des Schweizerischen entomologischen Gesellschaft
(in press).

ANDRE, E. 1892. Matériaux myrmécologiques. Revue d' Entomologie 11: 45-56.

BARONI URBANI, C. & SAUNDERS, J. B. 1982. The fauna of the Dominican Republic amber: the
present status of knowledge. Transaction of the Ninth Caribbean Geological Conference,
Santo Domingo 1: 213-223, 1 pl.

BARONI URBANI, C. 1995. Invasion and Extinction in the West Indian ant fauna revised: the
example of Pheidole (Amber Collection Stuttgart: Hymenoptera, Formicidae. VIII:
Myrmicinae, partim). Stuttgarter Beiträge zur Naturkunde (B) 222: 1-29.

BOLTON, B. 1994. Identification guide to the ant genera of the world. Cambridge, Massachusetts,
222
222 pp.

BORGMEIER, T. 1937. Formigas novas ou pouco conhecidas da America do Sul e Central,
principalmente do Brasil (Hym. Formicidae). Archivos do Instituto de Biologia Vegetal 3
(2): 217-255.

Brown, W. L., JR. 1973. A comparison of the Hylean and Congo-West African rain forest ant
fuanas. Pp. 161-185. In: MEGGERS, B. J., AYENSU, E. S. & DUCKWORTH, W. D. (eds).
Tropical forest ecosystems in Africa and South America, a review. Washington D. C.
(Smithsonian Inst. Press).

Brown, W. L. JR. 1975. Contributions toward a Reclassification of the Formicidae. V. Pone-
rinae, Tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini, and
Aenictogitini. Search, Agriculture, Entomology (Ithaca) 5 (15): 1-116.

CAMERON, P. 1891. Appendix. Hymenoptera, Formicidae. Pp. 89-95. In: WHYMPER, E. (ed.).
Travels amongst the Great Andes of the Equator. London, 147 pp.

CLARK, D. B., GUAYASAMIN, C., PAZMINO, O., Donoso, C. & PAEZ DE VILLACIS, Y. 1982. The
tramp ant Wasmannia auropunctata: autoecology and effects on ant diversity and distri-
bution on Santa Cruz Island, Galapagos. Biotropica 14: 196-207.

FOSSIL AND EXTANT SPECIES OF CYLINDROMYRMEX 663

Emery, C. 1901. Notes sur les sous-familles des dorylines et ponérines (famille des formicides).
Annales de la Societe Entolologique de Belgique 45: 32-54.

FELSENSTEIN, J. 1985. Confidence limits on phylogenies: an approach using bootstrap. Evolution
39: 379-404.

FERNANDEZ-C. F. & ESCOBAR-S., F. 1997. Primer registro de Cylindromyrmex Mayr (Hymen-
optera: Formicidae) para Colombia. Caldasia 19 (1-2): 347.

FOREL, A. 1892. Critique de: Peter Cameron. Hymenoptera, Formicidae. Extracted from supple-
mentary appendix to travels amongst the Great Andes of the Equator by Edw. Whymper.
London, 1891. Annales de la Societe Entomologique de Belgique 36: 255-256.

FOREL, A. 1899. Biologia Centrali-Americana. Insecta. Hymenoptera. Vol. III. (Formicidae).
London (Taylor and Francis), 169 pp., 4 pl.

FOWLER, H. G. & DELABIE, J. H. C. 1995. A new record of Cylindromyrmex striatus and range
extension of C. brasiliensis in Brazil (Hymenoptera: Formicidae). Revista de Biologia
Tropical 43: 327-328.

HÖLLDOBLER, B. & WILSON, E. O. 1990:. The ants. Cambridge/Mass. (The Belknap Press of
Harvard University Press), 732 pp.

JAFFE, K. C. 1993. El mundo de las hormigas. Baruta, Venezuela: Equinoccio (Ediciones de la
Universidad Simon Bolivar), 185 pp., 52 pl.

KEMPF, W. W. 1968. Miscellaneous studies on Neotropical ants. IV. (Hymenoptera, Formicidae).
Studia Entomologica 11: 369-415.

KEMPF, W. W. 1972. Catalogo abreviado das formigas da regiäo neotropical. Studia Entomolo-
gica 15: 3-334.

LUBIN, Y. D. 1984. Changes in the native fauna of the Galapagos Islands following invasion by
the little red fire ant, Wasmannia auropunctata. Biological Journal of the Linnean Society
21: 229-242.

Mackay, W. P. 1996. A revision of the ant genus Acanthostichus (Hymenoptera: Formicidae).
Sociobiology 27: 129-179.

Mappison, W. P. & MAppison, D. R. 1992. MacClade: analysis of phylogeny and character

evolution. Version 3.1.1, xi + 398 pp. + floppy disk.

Mayr, G. L. 1870. Neue Formiciden. Verhandlungen der k. k. Zoologisch-Botanischen Gessell-

schaft in Wien 20: 939 - 996.

Mayr, G. L. 1887. Siidamerikanische Formiciden. Verhandlungen der k. k. Zoologish-Bota-

nischen Gesellschaft in Wien, 37: 511-632.

MENOZZI, . 1931. Qualque nuova Formica di Costa Rica. (Hym.). Stettiner Entomologische
Zeitung 92: 188-202.

OVERAL, W. L. & BANDEIRA, A. G. 1985. Nota sobre habitos de Cylindromyrmex striatus Mayr,
1870, na Amazonia (Formicidae, Ponerinae). Revista brasileira de Entomologia 29: 521-
522.

SANTSCHI, F. 1924. Nouvelles Fourmis brésiliennes. Annales de la Societé Entomologique de
Belgique 64: 5-20.

SANTSCHI, F. 1932. Quelques fourmis inédites de l'Amérique centrale et Cuba. Revista de
Entomologia 2: 410-414.

SNELLING, R. R. & Hunt, J. H. 1975 The ants of Chile (Hymenoptera: Formicidae). Revista
chilena de Entomologia 9: 63-129.

STITZ, H. 1932. The Norwegian Zoological Expedition to the Galapagos Islands 1925, conducted
by Alf Wollebek. 5. Formicidae. Meddelelser fra det Zoologiske Museum Oslo 31: 367-
572;

SWOFFORD, D. L. 1993. PAUP, Phylogenetic Analysis Using Parsimony. Version 3.1.1. A
computer program distributed by the Illinois Natural History Survey, Champagne/Illinois.

664 MARIA L. DE ANDRADE

TOMOTAKE, M. E. M. & CAETANO, F. H. 1997. Morphology of the digestive tract of Acanthosthi-
chus serratulus and Cylindromyrmex brasiliensis (Hymenoptera: Formicidae). Naturalia
2299213:

WHEELER, W. M. 1910. Ants, their structure, development and behavior, xxv+663 pp. New York
(Columbia Univ. Press).

WHEELER, W. M. 1914. The ants of the Baltic amber. Schriften der Physikalisch-ökonomischen
Gesellschaft zu Königsberg 55: 1-142.

WHEELER, W. M. 1919. Expedition of the California Academy of Sciences to the Galäpagos
Islands, 1905-1906. Part XIV. The ants of the Galapagos Islands. Proceedings of the
California Academy of Sciences 14: 259-297.

WHEELER, W. M. 1924. The Formicidae of the Harrison Willians Galapagos Expedition.
Zoologica 5: 101-122.

WHEELER, W. M. 1934. Neotropical ants collected by Dr. Elisabeth Skwarra and others. Bulletin
ofthe Museum of Comparative Zoology 77: 157-240.

WHEELER, W. M. 1936. Ecological relations of ponerine and other ants to termites. Proceedings
of the American Academy of Arts and Sciences 71: 159-243.

WHEELER, W. M. 1937. Ants mostly from the mountains of Cuba. Bulletin of the Museum of
Comparative Zoology 81: 441-465.

WHEELER, W.M. & MANN, W. M. 1914. The ants of Haiti. Bulletin of the American Museum of
Natural History 3: 1-61.

WILSON, E. O. 1985. Ants of the Dominican amber (Hymenoptera: Formicidae). 1. Two new
myrmicine genera and an aberrant Pheidole. Psyche 92: 1-9.

REVUE SUISSE DE ZOOLOGIE 105 (3): 665-732; septembre 1998

Aleocharinae della Cina: Parte III
(Coleoptera, Staphylinidae)

Roberto PACE
Via Vittorio Veneto 13, I-37032 Monteforte d’Alpone (Verona), Italia.

Aleocharinae from China: Part III (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 d, 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’estremita delle antenne dell’holo-
typus d 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 d, China, Yunnan, Dali, 9.11.1993, de Rougemont leg. (MHNG).

Paratypi: 2 & e 1 2, 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-

PY&A.

Atheta (Philhygra) dalijiensis sp. n. Figg. 10-12

Holotypus d, 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. 11-12.

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 6, China, Gansu, M. ts 25 Km E Xiahe, 3000 m, 5.VIII.1994, A. Smetana
leg. (MHNG).
Paratypi: 2 d, stessa provenienza.

667

ALEOCHARINAE DELLA CINA

01 mm

1mm

Fico. 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) conferta sp. n.

668 ROBERTO PACE

Fico. 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 d, China, Zhejiang Tianmushan, 29.1V.1993, de Rougemont leg. (MHNG).

Paratypus: 1 ®, 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 nell’edeago della nuova specie.

Atheta (Philhygra) ideogrammifera sp. n. Figg. 20-22

Holotypus d, 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 nell’edeago
di pseudoelongatula. Inoltre l’edeago della nuova specie, in visione ventrale, è
strettissimo, mentre è largo in pseudoelongatula.

670 ROBERTO PACE

Fıcc. 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

F1GG. 20-25

Habitus ed edeago in visione laterale e ventrale. 20-22: Atheta (Philhygra) ideogrammifera
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 6, 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 basalı 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 d, China, Shanxi, Wutaishan, 4-5.V1.1993, de Rougemont leg. (MHNG).

Paratypus: 1 ©, 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 d, China, Yunnan, Kunming, I-II.1993, de Rougemont leg. (MHNG).
Paratypi: 2 ©, 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

01 mm

FIGG. 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 ®, 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 1
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 d, China, Yunnan, Ruili, ca. 700 m, 3.11.1993, de Rougemont leg. (MHNG).

Paratypus: 1 d, stessa provenienza.

DESCRIZIONE. Lunghezza 1,8 mm. Corpo lucido e bruno con pronoto bruno-
rossiccio e con 1 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

01 mm

FicG. 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 d, China, Beijing, B.N.U., flight interception trap, 10.VI-10.VII.1993, de
Rougemont leg. (MHNG).

Paratypi: 1 d e 1 ©, stessa provenienza; 1 d, China, Shanxi, Wutaishan, 4-5.VI.1993,
de Rougemont leg.

DESCRIZIONE. Lunghezza 2,4 mm. Corpo lucido e bruno con elitre ed estremita
addominale bruno-rossicci; antenne nere con antennomero basale bruno; zampe
giallo-brune con tarsi gialli. La reticolazione e distinta sul capo e svanita sul resto del
corpo. I tubercoletti dell’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 piu 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 4, 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-Amgog, 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

Feel
dr
EE Cy
SGRIUTVAN
ESATA

0,Imm

Imm

FIGG. 39-46

Habitus, edeago in visione laterale e ventrale e spermateca. 39-42: Atheta (Acrotona) 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 € 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 d, China, Zhejiang, Tianmushan, 29.1V.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 meta 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 d, China, Zhejiang, Tianmushan, 29.1V.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 d, China, Shaanxi, Nanwutai, 17.1X.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

01 mm
Imm

Ù E
A à
I, La

i
2a!

Imm

F1GG. 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 d, 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 estremita 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 d, 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 di fletcheri.

Atheta (Acrotona) singularis sp. n. Figg. 73-75

Holotypus à, 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

Fico. 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) filia sp. n.

682 ROBERTO PACE

01mm

FıGc. 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 é fitta, simile a quella di molte specie di
Myllaena. Edeago figg. 74-75.

COMPARAZIONI. La nuova specie € 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
nell’edeago di fletcheri.

Atheta (Acrotona) lingicola sp. n. Figg. 76-77

Holotypus 9, 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 d, China, Beijing, Xishan, [X.1992, de Rougemont leg. (MHNG).

Paratypi: 14 es., stessa provenienza; 1 ®, 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

Fico. 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) lingicola 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 d, Hong Kong, N.T., Kadoorie Farm, VI.1992, Malaise trap, G. Ades leg.
(MHNG).

Paratypi: 1 d e 1 2, 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, 1 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
nell’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 di 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

ROBERTO PACE

686

01 mm

r
r
Ih
di UNI uns
SAD wy Pg
lls
RS Coa
= ee "er E
= Eire sE
Ah Ur UA
tr 1 QUES
ni AR
i Suir tr UN ie Hi À
- d hay
ER

Imm

0mm

Fico. 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

in fungi).

Atheta (Acrotona) parasuspiciosa sp. n. Figg. 88-89

Holotypus 2, China, Gansu, M. ts 25 Km E Xiahe, 2805-2925 m, 3.VII.1994, A.
Smetana leg. (MHNG).

Paratypus: 1 ©, 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 2, China, Gansu, Yonghai, ca. 20 Km SW Yuzhong, 2700-2800 m,
9.VIH.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 d, China, Gansu, M. ts 25 Km E Xiahe, 2805-2925 m, 3.VIII.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

FIGG. 86-91

Habitus e spermateca. 86-87: Atheta (Acrotona) pseudofungi sp. n.; 88-89: Atheta (Acrotona)
parasuspiciosa sp. n.; 90-91: Atheta (Acrotona) yuzhongensis sp. n.

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 1 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 d, China, Beijing, Xishan, IX.1992, de Rougemont leg. (MHNG).

Paratypi: 1 d e 1 ?, stessa provenienza; 1 ®, Beijing, Songshan, 15.V.1993, de
Rougemont leg.; 23 es., Beijing, B.N.U., at light, V-VI.1993, de Rougemont leg.; 1 d e 1 9,
Beijing, B.N.U. campus, from birds nesting box, 7.VII.1993, de Rougemont leg.; 1 4, Beijing,
B.N.U., flight interception trap, 10.VI-10.VII.1993, de Rougemont leg.; 2 2, China, Henan,
Luoyang, 10.[X.1994, de Rougemont leg.; 1 4, China, Jiangsu, Suzhoo, 3.IX.1994, de Rouge-
mont leg.; 13 es., China, Hebei Prov., Yongnian, 6.X.1995, Shugiang 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

ROBERTO PACE

690

005mm

Fico. 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 d, China, Jiangsu, Nanjing, 17.VIII.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.
Edeago figg. 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 ci6 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

01 mm

100 101 102

Figc. 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. 108-110

Holotypus d, China, Beijing, Xishan, IX.1992, de Rougemont leg. (MHNG).

Paratypus: 1 d, 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. 109-110.

COMPARAZIONI. La forma dell’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, nè robusta armatura genitale
interna, come in vicaria, nè l'apice dell’edeago stesso, in visione ventrale, ad ogiva
come in vicaria.

Atheta (Acrotona) mimannuliventris sp. n. Figg. 111-114

Holotypus 3, Hong Kong, Tai Po, III.1996, de Rougemont leg. (MHNG).

Paratypus: 1 ©, 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

T
AE
NAME
NIRO

Figc. 106-114

Habitus, spermateca ed edeago in visione laterale e ventrale. 106-107: Atheta (Acrotona)
guandongensis sp. n.; 108-110: Arheta (Acrotona) fraudolenta sp. n.; 111-114: 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. 119-120

Holotypus ?, 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 1 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 d, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 26.VII.1994,
A. Smetana leg. (MHNG).

Paratypi: 1 2, 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

01 mm

F1GG. 115-121

Habitus e spermateca. 115-116: Atheta (Acrotona) inornata (Kraatz); 117-118: Arheta
(Dimetrota) insinuata sp. n.; 119-120: Atheta (Dimetrota) sericoides sp. n.; 121: Atheta
(Dimetrota) apicalis sp. n.

ALEOCHARINAE DELLA CINA 697

Atheta (Dimetrota) paritetica sp. n. Figg. 125-127

Holotypus 6, China, Yunnan, Kunming, 1.11.1993, 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 d, 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 1 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 dé, China, Shaanxi, Nanwutai, 17.1X.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

FIGG. 122-128

Edeago in visione laterale e ventrale, spermateca e habitus. 122-124: Arheta (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 d, China Shaanxi, Nanwutai, 17.1X.1995, de Rougemont leg. (MHNG).

Paratypi: 3 2, 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 1 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 d, 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

131

01 mm

133 134

135

Here

01 mm

FicG. 129-136

Edeago in visione laterale e ventrale, spermateca e habitus. 129-131: 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 4, 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 d, China, Yunnan, Ruili, ca. 700 m, 3.11.1993, de Rougemont leg. (MHNG).

Paratypi: 2 d, 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

01 mm

Fico. 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

FIGG. 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 d, China, Beijing, Xiaolongmen, 1100-1500 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 ©, China, Zhejiang, Tianmushan, 29.1V.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 d, China, Yunnan, Dali, 9.1.1993, 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

\
Vue

Ficc. 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 d, China, Shanxi, Wutaishan, 4-5.VI.1993, de Rougemont leg. (MHNG).

Paratypi: 2 d e 1 ©, 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 ®, 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

161

164

01 mm

F1GG. 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 d, 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 1
due antennomeri basali rossicci; zampe rossicce. La reticolazione del capo é 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.
Edeago figg. 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 dell’edeago 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 2, 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’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 d, 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 é netta, quella del pronoto e dell’addome é distinta (a
maglie lievemente trasverse sull’addome) e quella delle elitre & assente. I tubercoletti
della superficie del capo e delle elitre sono distinti, quelli del pronoto sono svaniti e

ALEOCHARINAE DELLA CINA 709

172

Ficc. 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 profondita ventrale, per 1 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 9, Hong Kong, Kadoorie Farm, V.1996, de Rougemont leg. (MHNG).

Paratypus: 1 ©, Hong Kong, Tai Po, VII.1996, de Rougemont leg.

DESCRIZIONE. Lunghezza 2,7 mm. Corpo lucido e nero con pronoto, metà
basale delle elitre, 1 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 d, China, Gansu, pass btw Herzuo-Amgog, 3300 m, 12.VII.1994.
A. Smetana leg. (MHNG).

Paratypi: 9 ®, stessa provenienza; 11 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

Ficc. 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.

712 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 d, 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.,
11.V.1996, J. Cooter leg. (MHNG).

Paratypi: 1 d 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

Fico. 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 1 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 © , 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 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 d, China, Beijing, Xishan, IX.1992, de Rougemont leg. (MHNG).

Paratypus: 1 d, 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 WANS)

FicG. 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.

TG ROBERTO PACE

Atheta (Datomicra) viduoides sp. n. Figg. 199-202

Holotypus d, China, Yunnan, Xishuangbanna, Jinghong, II.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 d, China, Zhejiang, Tianmushan, 29.IX.1993, de Rougemont leg. (MHNG).

Paratypi: 1 d e 1 9, 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 d, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 26.VII.1994,
A. Smetana leg. (MHNG).

Paratypi: 23 es., stessa provenienza; 1 d, Gongga Shan, above camp 3, 3050 m,
22.VII.1994, 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

Ficc. 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 di 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 d, 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 2, Hong Kong, Kadoorie Agricultural Research Centre, Malaise trap, 19-
31.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. I 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 di formosa. Inoltre l’introflessione apicale del bulbo distale della
spermateca della nuova specie è emisferica, mentre è conica in formosa.

ALEOCHARINAE DELLA CINA 719

1mm

205 206

211

FIGG. 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

FIGG. 213-219

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio.
213-217: Atheta (Datomicra) bimacula sp. n.; 218-219: Atheta (Datomicra) mimoformosa sp. n.

ALEOCHARINAE DELLA CINA 72]

01 mm

Ph Gg Mp hh pt
el!

224

Olmm

228

FIGG. 220-229

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio
220-224: Atheta (Datomicra) canescens (Sharp); 225-229: Arheta (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 d, China, Zhejiang, Tianmushan, 29.1V.1993, de Rougemont leg. (MHNG).

Paratypi: 3 d 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 2, 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

“RATORI
Hines AA

0. mm

235

Olmm

Fic. 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

FIGG. 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, nè 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 d, China, Sichuan, Gongga Shan, above capo 3, 3050 m, 22.VII.1994,
A. Smetana leg. (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 é, China, Gansu, Xinlong Shan, ca. 70 Km S Lanzhou, 2225-2380 m,
7.VII.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

Fico. 241-246

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio.
241-245: Atheta (Datomicra) parainopinata sp. n.; 246: Atheta (Datomicra) xinlongensis sp. n.

ALEOCHARINAE DELLA CINA TOT

Figc. 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) regressa 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) regressa sp. n. Figg. 251-255

Holotypus d, China, Gansu, Xinlon Shan, ca. 70 Km S Lanzhou, 2225-2380 m, 7.VII.
1994, A. Smetana leg. (MHNG).

Paratypus: 1 2, 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 d, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 25.VII.1994,
A. Smetana leg. (MHNG).

Paratypi: | 9, stessa provenienza; 1 & e 1 2, China, Gansu, Xinlong Shan, ca. 70 Km
S Lanzhou, 2225-2380 m, 7.VIII1994, 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

FIGG. 254-260

Edeago in visione laterale e ventrale, habitus, spermateca e sesto urotergo libero del maschio.
254-255: Atheta (Datomicra) regressa 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 3, China, Zhejiang, Tianmushan, 29.1V.1993, de Rougemont leg.; 18 es. China, Zhejiang,
Hangzhou, 27.1V.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, 1100-1500 m, 1.VII.1993, de Rougemont leg.; 1 d e 1 9,
China, Ganzu, Dalijia Shan, 46 Km W Linkia, 2980 m, 10.VII.1994, A. Smetana leg.; 1 d e
1 2, 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.[X.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 (Datostiba) poroshirica Sawada, 1978: 243, syn. n.
Atheta (Datomicra) shuteae Pace, 1987: 420, syn. n.
1 de2 9, China, Gansu, Pass btw Hezuo-Amgog, 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 dell’“Institut Royal des Sciences Naturelles” di Bruxelles e il Dr L.
Zerche del D.E.I. di Eberswalde.

ALEOCHARINAE DELLA CINA 73]

BIBLIOGRAFIA

BERNHAUER, M. 1907. Zur Staphylinidenfauna von Japan. Verhandlungen der Zoologisch-
Botanischen Gesellschaft in Wien 57: 371-414.

BRUNDIN, L. 1943. Zur Kenntnis einiger in die Atheta-Untergattung Metaxya MR. gestellten
Arten (Col. Staphylinidae). Lunds Universitets Arsskrift N.F. Ad. 2, 39: 3-37.

CAMERON, M. 1939. The Fauna of British India, including Ceylon and Burma. Coleoptera,
Staphylinidae 4: 410 pp., London.

CAMERON, M. 1943. New species of Staphylinidae (Col.) from Borneo. The Entomologist's
monthly Magazine 79: 39-42.

CAMERON, M. 1944a. Descriptions of new Staphylinidae (Coleoptera). The Entomologist's
monthly Magazine 76: 103-106.

CAMERON, M. 1944b. New species of Staphylinidae (Col.) from China and Japan. The
Entomologist’s monthly Magazine 80: 158-159.

EPPELSHEIM, E. 1893. Beitrag zur Staphylinidenfauna des südwestlichen Baikalgebiete.
Deutsche Entomologische Zeitschrift 37: 17-67.

ERICHSON, G. 1839. Genera et Species Staphylinorum Insectorum Coleopterorum Familiae 1:
400 pp., Berlin.

GRAVENHORST, J. J. C. 1806. Monographia Coleopterorum Micropterorum. 236 pp., Gottingae.

KIESENWETTER, E. A. H. 1844. Die Staphylinidenfauna von Leipzig’s Umgegend. Stettiner Ent.
Zeitung 5: 307-320.

KRAATZ, G. 1858. Naturgeschichte der Insecten Deutschlands. Coleoptera 2: 1080 pp., Berlin.

KRAATZ, G. 1859. Die Staphyliniden-Fauna von Ostindien, insbesondere der Insel Ceylan.
Archiv fiir Naturgeschichte Berlin 25: 1-196.

Louse, G. A. 1974. Staphylinidae II (Hypocyphtinae und Aleocharinae). In: Die Käfer
Mitteleuropas. Band 5: 304 pp., Krefeld.

MANNERHEIM, C. G. 1830. Précis d’un nouvel arrangement de la Famille des Brachélytres de
l’Ordre des Insectes Coléoptères. Mémoires de l’Académie Impériale des Sciences de
St. Pétersbourg 1: 415-511.

MOTSCHULSKY, U. 1859. Insectes des Indes orientales et de contrées analogues. Etudes Entomo-
logiques 8: 25-118.

PACE, R. 1984. Aleocharinae della Thailandia e della Birmania riportate da G. de Rougemont.
Bollettino del Museo civico di Storia naturale di Verona 11: 427-468.

PACE, R. 1987a. Aleocharinae riportate dall’ Himalaya dal Prof. Franz. Parte III. Nouvelle Revue
d’Entomologie 4: 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. Jn: 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.

SHARP, D. 1869. A Revision of the British species of Homalota. Transactions of the
Entomological Society of London 1869: 91-272.

THOMSON, C. G. 1867. Skandinaviens Coleoptera 9: 407 pp., Lund.

REVUE SUISSE DE ZOOLOGIE

Tome 105 — Fascicule 3

Pozzi, Stefano, Yves GONSETH & Ambros HANGGI. Evaluation de l’entre-
tien des prairies seches du plateau occidental suisse par le biais de
leurs peuplements arachnologiques (Arachnida: Araneae).........

ASSING, Volker. A revision of the Habrocerinae of the world. Supplement
Ne(ColeopteratStaphylinidae)e EEE EPP ioe bee

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

MARTENS, Jochen & Peter SCHWENDINGER. A taxonomic revision of the
family Oncopodidae I. New genera and new species of Gnomulus
ihorelli(OpilionesyEaniatores)arr.2..... ee SORA

Tan, Heok Hui & Maurice KOTTELAT. Redescription of Betta picta
(Teleostei: Osphronemidae) and description of B. falx sp. n. from cen-
UR SUTENE, LR ME ne

BARBALAT, Sylvie. Importance of forest structures on four beetle families
(Col.: Buprestidae, Cerambycidae, Lucanidae and phytophagous Sca-
rabaeidae) in the Areuse Gorges (Neuchâtel, Switzerland). .........

DE ANDRADE, Maria L. Fossil and extant species of Cylindromyrmex
(iiymenopteras Formicidae)... ETRO IDR

PACE, Roberto. Aleocharinae della Cina: Parte III (Coleoptera, Staphy-
HG A a e SNO Ce A Re ae ee

465-485

487-492

493-497

499-555

557-568

569-580

581-664

665-732

REVUE SUISSE DE ZOOLOGIE

Volume 105 — Number 3

Pozzi, Stefano, Yves GONSETH & Ambros HANGGI. Evaluation of dry
grassland management on the Swiss occidental plateau using spider
Communities (ArachnidaAraneae). O OO

ASSING, Volker. A revision of the Habrocerinae of the world. Supplement
IMl@oleopterazStaphylinidae). "CEE PRES

UHMANN, Gerhard. Description of four new species of the genus Dera-
rimus (Coleoptera, Anthicidae) from Malaysia ..................

MARTENS, Jochen & Peter SCHWENDINGER. A taxonomic revision of the
family Oncopodidae I. New genera and new species of Gnomulus
ihorell (Opiliones,Laniatores)......... 2.0.2.2. RR ee

TAN, Heok Hui & Maurice KOTTELAT. Redescription of Betta picta
(Teleostei: Osphronemidae) and description of B. falx sp. n. from cen-
CARS UT AT A ee era nen ker N

DE ANDRADE, Maria L. Fossil and extant species of Cylindromyrmex
(Hymenoptera: Formicidae) i RO eee

PACE, Roberto. Aleocharinae from China: Part III (Coleoptera, Staphy-
MES i Rn AI en: AREE

Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX

Pages

465
487

493

499

569
581

665

PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE
CATALOGUE DES INVERTEBRES DE LA SUISSE

Base SARCODINÉS parlE SPENARDI LISI TE a, 12—
Pee EEE eORODESIparalihy SMINGELING Hey een 12
SARARAATEGNEESIpaBRSDELCESSERIEREIE SNO 42.—
ASS OPODES| pared GAR I ee SIIT 8.—
Se SEUDOSCORRIONS pare ReDETEESSERD jae oeioe rs oie I 5.50
CEMINEUSOIRESIPAREFANDRE RM II II 18.—
URS OPISOEHENESIParEBIEUENIEUKTBRETSCHERE I. er =
Se COLERODES pan MANTIEIERAUDE O RR O NI 18.—
IEORIEIONSIPALREIDENBESSERTE De ee 11.—

lO; SGORAONIS ae IRS DS ILIISIINN Gs opces OO I O 3550)
ME ROVAREURS pate SE "WEBER CG MONTE ER CE SIE 38.—
VED ECAPODES pated MEARS sy tuoi ee ee 11.—
Be ACAN HOGER HAICES spar eANDRE rescission eee tte nate eee eee 11.—
A GASTRÉROMRICHES PAR GA MONTE O eileen ec 18.—
ISSEAMBENIBODESIPARTHEARTN A ee 12.—
16. HIRUDINEES, BRANCHIOBDELLES

EWBOTSYEHEINESE Pan EFANDRE FE TER TO TE 17.50
IFRCESTODESIDARONEUERMANNE ARR EE iim eric ote rane CC. 30.—

REVUE E PALE ORIOLOGIE: nenn. Echange ou par fascicule Fr. 35.—

LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris)....... par fascicule Fr. 35.—

CATALOGUE ILLUSTRE DE LA COLLECTION LAMARCK Ù

APPARTENANT AU MUSEUM D'HISTOIRE NATURELLE DE GENEVE

lÉpartie Fossiles il) vols4> avec lii7-planches; ON nn Fr. 300.—
COLLEMBOLENFAUNA EUROPAS

von H. GISIN, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck 1984) .............. Fr. 30.—
THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND

DISTRIBUTION WITH KEYS FOR DETERMINATION

byARINOSEKS S46 pages, Lill tieuresimtext 1973 ee Fr. 30.—
CLASSIFICATION OF THE DIPLOPODA

DAMRICRATAMENEOEEMAN, 235i pages MOTO ee eels ees Fr. 30.—
LES OISEAUX NICHEURS DU CANTON DE GENEVE

par P. GEROUDET, C. GUEX et M. MAIRE

Slepacessnombreuses cartesset-tieures 1983, ra ee Fr. 45.—
CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS

CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES,

SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ

A LA CONNAISSANCE DES ECHINODERMES ACTUELS

PawMichelWANGOUX, 67 pages, lil planches 1985) PAM NE NO n Fr. 15.—
RADULAS DE GASTEROPODES LITTORAUX DE LA MANCHE

(COTENTIN-BAIE DE SEINE, FRANCE)

Pane BINED WUESTCEKe MAREDA 62 pages 1 99 Er ee ee Fr. 10.—

GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN;

SHELLS AND RADULAS

byAYeIRINED se WiWESian dike sNIARED ACIS SD I Sener Fr. 30.—
O. SCHMIDT SPONGE CATALOGUE

par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992.......... Fr. 40.—
ATLAS DE REPARTITION DES AMPHIBIENS

ET REPTILES DU CANTON DE GENEVE

PARA INELCER, VE AELLENCLNV IMAENERT, 48 pages, 199302 nee aime) Se
THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS:

A DOCUMENTED FAUNAL LIST

DAME SEINE ISO A LSM SER OE E ne Fr. 30.—
NOTICE SUR LES COLLECTIONS MALACOLOGIQUES

DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE

pagane Claude CATRRIEZAA 9 pa Pe SAI IS EE Fr. 22—
PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS

OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr. 160.—

CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE)

(Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997 ................... Fr. 50.—

A inniz DIT
Fr RITA SI

un ona fed Ÿ

PAL ES,

tote, vota ont
= i x ai a

Re ne vurlige de Za

Volume 105 - Number 3 - 1998
Revue suisse de Zoologie: Instructions to Authors

The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific
results based on the collections of the Muséum d’histoire naturelle, Geneva. Submission of a manuscript implies
that it has been approved by all named authors, that it reports their unpublished work and that it is not being
considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of
colour plates and large manuscripts. All papers are refereed by expert(s).

In order to facilitate publication and avoid delays authors should follow the Instructions to Authors and refer to
a current number of R.S.Z. for acceptable style and format. Manuscripts not conforming with these directives are
liable to be returned to the authors. Papers may be written in French, German, Italian and English. Authors should
aim to communicate ideas and information clearly and concisely in language suitable for the moderate specialist.
Authors not writing in their native language should pay particular attention to the linguistical quality of the text.

Manuscripts must be typed, on one side only and double-spaced, on A4 (210 x 297 mm) or equivalent paper
and all pages should be numbered. All margins must be at least 25 mm wide. Authors must submit one original
and two copies, including tables and figures, in final fully corrected form, and are expected to retain another copy.

Papers should conform to the following general layout:

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces,
name(s) in full and surname(s) of author(s), and full address(es).

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should
summarise the contents and conclusions of the paper. The abstract is followed by less than 10 key-words,
separated by hyphens, which are suitable for indexing.

Introduction. A short introduction to the background and the reasons for the work.

Materials and methods. Sufficient experimental details must be given to enable other workers to repeat the
work. The full binominal name should be given for all organisms. The International Code of Zoological Nomen-
clature must be strictly followed. Cite the authors of species on their first mention. Use SI units and the appropriate
symbols.

Results. These should be concise and should not include methods or discussion. Text, tables and figures should
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp.
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types
should be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed
by synonyms, material examined and distribution, description and comments. All material examined should be listed
in similar, compact and easily intelligible format; the information should be in the same language as the text.

Sex symbols should be used rather than “male” and “female”.

Discussion. This should not be excessive and should not repeat results nor contain new information, but
should emphasize the significance and relevance of the results reported.

References. The Harvard System must be used for the citation of references in the text, e.g. White & Green
(1995) or (White & Green 1995). For references with three and more authors the form Brown et al. should be
used. Authors’ names should not be written in capitals. The list of references must include all publications cited in
the text but only these. References must be listed in alphabetical order of authors, and both the title and name of
the journal must be given in full in the following style:

Penard, E. 1888. Recherches sur le Ceratium macroceros. These, Geneve, 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.

References should not be interspaced and, in the case of several papers by the same author, the name has to be
repeated for each reference.

The text should be in roman (standard) type face throughout, including headings, with genus and species
names underlined with pencil; bold, small capitals, large capitals, italics and other type faces should not be used.
Footnotes and cross-references by page should be avoided.

Also, we encourage authors to submit the text on floppy disk (3,5’’ or 5 1/4’’, Macintosh or IBM compatible,
ath “Microsoft Word” or similar programmes). The authors should refrain from using special codes for text
ormating.

Tables. These should be self-explanatory, with the title at the top organised to fit 122 x 180 mm. Each table
should be typed, double spaced, on a separate page and numbered consecutively and its position indicated in the text.

Figures. These may be line drawings or half tones and all should be numbered consecutively, and their position
indicated in the text. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and
lettering should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors
should refrain from mixing the drawings and half tones. Original drawings will not be returned automatically. The
Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents.

Legends to figures. These should be typed in numerical order on a separate sheet.

Proofs. Page proofs only are supplied, and authors may be charged for alterations (other than printer’s errors) if
they are numerous.

Offprints. The authors receive 25 offprints free of charge; more copies may be ordered at current prices when
proofs are returned.

Correspondence. All correspondence should be addressed to

Revue suisse de Zoologie
Muséum d'histoire naturelle
CP 6434
CH-1211 Genève 6
Switzerland.
Phone: +41 22 418 63 33 - Fax +41 22 418 63 01
e-mail: volker.mahnert @ mhn.ville-ge.ch
Home page RSZ: http://www-geneva-city.ch:80/musinfo/mhng/publications/revues.htm

CAT

DECEMBRE 1998 ISSN 0035 - 418 X

REVUE SUISSE DE ZOOLOGIE

TOME 105 — FASCICULE 4

Publication subventionnée par l'Académie suisse des Sciences naturelles
et la Societe suisse de Zoologie

VOLKER MAHNERT
Directeur du Muséum d'histoire naturelle de Genève

FRANCOIS BAUD
Conservateur au Muséum d'histoire naturelle de Genève

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

Comité de lecture

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

Président: Ivan LOBL — Muséum de Genève

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

SUISSE Fr. 225.—

PRIX DE L'ABONNEMENT:

(en francs suisses)

Les demandes d'abonnement doivent étre adressées
à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse

UNION POSTALE Fr. 230.—

ANNALES

de la

SOCIETE SUISSE DE ZOOLOGIE

et du

MUSEUM D'HISTOIRE NATURELLE
de la Ville de Genéve

tome 105
fascicule 4
1998

REVUE SUISSE DE ZOOLOGIE

E À Z
kl GENEVE DECEMBRE 1998 ISSN 0035 - 418 X

REVUE SUISSE DE ZOOLOGIE

TOME 105 — FASCICULE 4

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

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

Président: Ivan LOBL — Museum de Genève

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

SUISSE Fr. 225.—

PRIX DE L'ABONNEMENT:

(en francs suisses)

Les demandes d'abonnement doivent étre adressées
à la rédaction de la Revue suisse de Zoologie,
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse

UNION POSTALE Fr. 230.—

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
19-20 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 ef 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 al.
BALLOUX Francois, 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 individual 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 MUHLHAUSER Claudia et al.
Bray Rodney A., see HoBERG Eric P. er al.
BRUNNER 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 Jeröme & 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-Francois, LOIZEAU Pierre-André, ANDREWS S. &
SPICHIGER Rodolphe
Molecular Phylogeny and Biogeography of the Genus /lex L. (Aquifoliaceae)
Curcic Bozidar P. M. & MAKAROv Slobodan
[Endemic differentiation of millipedes (Diplopoda, Myriapoda) in Serbia,
Montenegro and Macedonia: taxonomic implications
2On geographic distribution and historical development of some cave-dwelling
diplopods (Myriapoda) in Serbia, Montenegro and Macedonia (FYROM)
DAJOZ Isabelle, see NADOT Sophie er al.
DE BRUIJN Anette, see SAVOLAINEN Vincent et al.
DE VARGAS Colomban!, ZANINETTI Louisette & PAWLOWSKI Jan
Cryptic diversity in the open Ocean
Isee 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.! & 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 annuus

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 Francois, 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 PAABO Svante et al.

GRIFFIN Andrea
Updating the path integrator through a visual fix

GROSSENBACHER Kurt, see LUSCHER Beatrice et al.

GUGGENHEIM Richard, see KAUPP Andreas et al.

GUNTERT Marcel, see LUSCHER Beatrice et al.

HAUSSER Jacques, see LUGON-MOULIN Nicolas and WUST SAUCY Anne-Gabrielle ef 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)

786 ZOOLOGIA ET BOTANICA ‘98

KELLER Laurent, see BERNASCONI Giorgina et al.

KRINGS Matthias, see PAABO Svante et al.

LOCHER Rolf & BAUR Bruno
Is the sperm pool in the hermaphroditic land snail Arianta arbustorum limited?

LOIZEAU Pierre-André, see CUENOUD Philippe et al.

LOSADA Freddy, see Vizoso Dita et al.

LUGON-MOULIN Nicolas, GOUDET Jeröme, BRUNNER Harald & WYTTENBACH

Andréas
Microsatellites reveal the fine-scale structure of a hybrid zone in Sorex
araneus (Insectivora, Soricidae)

LUSCHER Beatrice, GROSSENBACHER Kurt, GUNTERT Marcel & SCHOLL Adolf
Genetic differentiation of the common toad (Bufo bufo) in the Alps

MAKAROV Slobodan, see CURCIC Bozidar P. M. et al. |?

MANEN Jean-Francois, 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 predation 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?

MULLER 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.

PAABO 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 ‘98 787

PAWLOWSKI Jan!, DE VARGAS Colomban, FAHRNI José & ZANINETTI Louisette
Calibrating ribosomal clocks in foraminifera
Isee also DE VARGAS Colomban er 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 PAABO 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 ef 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 Jeröme et al., PERRET
Mathieu et al. and SPICHIGER Rodolphe et al.

SAVOLAINEN Vincent, CHASE Mark W., MORTON Cynthia M., Hoor 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! & EXCOFFIER Laurent
Estimation of past demographic parameters using the mismatch distribution
!see also DUPANLOUP Isabelle er 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 ef al. and LUSCHER 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! & SAVOLAINEN Vincent
Teaching botany in a molecular world
Isee 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 er 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.
WUEST 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!, RINGsBY Thor-Harald? & SAETHER Bernt-Erik?

! 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 Neuchatel; Rue Emile-Argand 11; CH-2007 Neuchatel;
Switzerland

740 ZOOLOGIA ET BOTANICA ‘98

Oviposition preference and larval performance in a leaf beetle
No abstract available.

BALLOUX Francois, GOUDET 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! & KELLER Laurent?

! Behavioral Ecology; University of Berne; Hinterkappelen
2 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 invicta) 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 (1)
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!?, VALSANGIACOMO Claudio?, PIFFARETTI Jean-Claude? &

WARD Paul!

! Zoologisches Museum der Universität; Abt. Ökologie; Winterthurerstrasse 190; CH-8057
Zürich; Switzerland

? Istituto Cantonale Batteriosierologico; Via Ospedale 6; CH-6904 Lugano; Switzerland

ZOOLOGIA ET BOTANICA ‘98 74]

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 anımal 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: (1) 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-Felix!, Dixon Paul? & SUGIHARA George?

! Institut de Zoologie; Rue Emile Argand, 11; CH-2007 Neuchatel; 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? 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 Ausgleichsflachen» 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
Ausgleichsflachen» and bigger extensively cultivated ares in combination with erecting more
perches.

CASTELLA Vincent, GOUDET Jeröme & PERRIN Nicolas

Institut de Zoologie et d’Ecologie Animale; Bätiment 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 (L&man and
Neuchätel) and six sites being of pond habitats. All individuals were genotyped using six
polymorphic allozyme locı.

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 selfing 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!, MANEN Jean-Francois!, LOIZEAU Pierre-André!, ANDREWS S.?

& SPICHIGER Rodolphe!

| 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 /lex L. (Aquifoliaceae)

The genus /lex 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 //ex 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 (Stylo-
brachydesmus) ljubetensis Attems, Xestoiulus (Oroiulus) macedonicus (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 Colomban!, ZANINETTI Louisette! & PAWLOWSKI Jan! 2

| Department de Zoologie et Biologie Animale; Université de Genève; CH-1224 Chéne-
Bougeries; Switzerland
2 Muséum d’histoire naturelle; CP 6434; CH-1211 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-1211 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 Ziirich; Winterthurerstrasse 190; CH-8057 Ziirich, 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 (Col->Coll, NINFA->CB3FC) to amplify the mt-DNA sequences. To cut them we
used three restriction enzymes Dral, SspI and Taq].

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 NINFA->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 Neuchatel; 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
Geneve 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 Genéve; 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; Universitat 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; Universitat 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

(1) comparing sequence divergence within acridoid lineages;

(11) comparing sequence divergence between other orthopteran groups;

(111) 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 Ziirich; Switzerland

RAPDs reveal extensive genetic variability in the apomictic Erigeron annuus

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; Batiment 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 (Brullé, 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 £. 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 Jeröme & SAVOLAINEN Vincent

Institut de Zoologie et d’Ecologie Animale & Institut de Botanique Systematique et Géobo-
tanique; Bätiment 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 ‘98

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.

Hoserc Eric P.!, MARIAUX Jean”, JONES Arlene? & BRAY Rodney A?

| Biosystematics and National Parasite Collection Unit; USDA; Agricultural Research Service:
Beltsville, MD 20705; USA

2 Muséum d’histoire naturelle; CP 6434; CH-1211 Genéve 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 12 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!, PILLER Werner2, ZANINETTI Louisette? & PAWLOWSKI Jan*

F Institut für Paläontologie der Universität Wien; Austria

5 2 Institut fiir Geologie und Palaeontologie der Universitàt Graz; Austria
„Departement de Geologie: Université de Genéve; Switzerland
4 Departement 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. | 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 ‘98 751

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 Chéne-
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!, GUGGENHEIM Richard, WirTz Sabine? & NAGEL Peter!

! Universität; Institut für Ntur-, Landschafts- und Umweltschutz / Biogeographie; St. Johanns-
Vorstadt 10; CH-4056 Basel; Switzerland
2 Universitit 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

152 ZOOLOGIA ET BOTANICA ‘98

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, BRUNNER 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 bybrid 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
R-statistics and one migrant per generation using F-statistics. Hierarchical F- and R-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,, 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!--, GROSSENBACHER Kurt?, GÜNTERT Marcel? & SCHOLL
Adolf!

! Institute of Zoology; University of Berne; Switzerland
2 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 Arles (Provence, France, B. b.
spinosus), respectively.

A 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, FLOOK Paul K. & ROWELL C. H. F.
Zoologisches Institut; Universitat 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 DI-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 Jérôme

Institut de Zoologie et d’Ecologie animale; Université de Lausanne; Batiment de Biologie; CH-
1015 Lausanne; Switzerland

Selection and life-history responses to size-dependent predation in Physa acuta
(Gastropoda)
In order to investigate the evolutionary response of life-histories to a size-dependent predation,

we performed two long-term (20 months) laboratory experiments involving the freshwater snail
Physa acuta.

754 ZOOLOGIA ET BOTANICA ‘98

1. A selection experiment on juvenile length - simulating predation - 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.

. The second experiment involved controlled predation by two flatworm species on artificial
populations, and also showed a significant increase in juvenile size in the populations under
predation.

3. These two results show that important life-history parameters of the species under study

may evolve quite rapidly (12 generations) under size-dependent predation.

MONTOYA-BURGOS Juan-Ignacio & PAWLOWSKI Jan
Museum d’histoire naturelle; 1, route de Malagnou; CP 6434; CH-1211 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 ITSI - 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 (1.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-1211 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 11 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.

Napot Sophie!, CREACH Jean-Baptiste!, BALLARD Harvey? & DAJOZ Isabelle?

| Laboratoire Evolution et Systématique; Université Paris 11; France
9 . . . .

“ 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.

PAABO 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 Zürich-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 tRNAH!S, tRNAS®, and
tRNAFEU 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 rosalia,
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!:, DE VARGAS Colomban!, FAHRNI José! & ZANINETTI Louisette!

l Département de Zoologie et Biologie animale; Université de Genève; CH-1224 Chéne-
Bougeries; Switzerland
2 Muséum d’histoire naturelle; CP 6434; CH-1211 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 |’ 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-1211 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 showed 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 Chéne-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!, SMITH Margaret F.? & PATTON James L.?

! Institut de Zoologie et d’Ecologie Animale; UNIL; CH-1015 Lausanne; Switzerland
? 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 7. bottae from New Mexico, we show
that apparent similarity is due to an introgression of 7. b. ruidosae mtDNA into 7. 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!-2, CHASE Mark W.!, Morton Cynthia M.’, Hoot Sara B.*,
Souris Douglas E.’, BAYER Clemens®, Fay Michael F.!, DE BRUIJN Anette!,
SULLIVAN Stuart! & Qru Yin-Long’

! 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; Universitat 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 Geneve; CH-1292 Chambesy; 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 Biometrie; 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; Universitat 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.

WACKERS 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.

WUEST Jean
Museum d’histoire naturelle; CP 6434; CH-1211 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 lineola, 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. lineola. 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. lineola, 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 A. 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!, HAUSSER Jacques!, TABERLET Pierre? & SAUCY
Francis’

! Institute of Zoology and Animal Ecology; University of Lausanne; CH-1015 Lausanne;
Switzerland

? Laboratoire des Populations d’ Altitude; CNRS EP55; University Joseph Fourier; F-38041
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 Neuchatel; Rue Emile Argand, 11; CH-2007 Neuchatel;
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 DNA 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

e the two traditional families: Monticelliidae and Proteocephalidae are not recognised

e 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

e 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.

e nn FTSE ©

\sorgealarÄgne

i) ve 2 | Ne à È
- pr » Lia À rw 1 if dvi jiu

nite MEN Fer x : ( (UNS amule sie
Fri rg k

ed ne À | mie rate as veel) Ai ft Det DA

REVUE SUISSE DE ZOOLOGIE 105 (4): 767-770; decembre 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 Geneve.

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 és sciences en 1976 (zoologie, botanique, physiologie, biochimie) 11
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 these de doctorat
sous la direction du Prof. W. Huber de l’université de Berne. En 1983 il défend sa
these sur le sujet «Biologie et gestion d’une population de chamois du Jura vaudois»
et obtient le grade de docteur ès 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 (1948-1998)

Zoologue de terrain, ıl a parcouru le Jura durant des semaines en toutes saisons
a la recherche des chamois, sujets de sa these. Dormant souvent a 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 micromammiferes 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 maitrisait 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° 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, ıl n’eut pas le temps de réaliser tous ses buts. Dans son bureau, plus d’une
douzaine de travaux resteront inacheves...

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...

BIBLIOGRAPHIE

BECK, C., FOREST, V., OLIVE, C. & DE ROGUIN, L. 1993. Animaux, techniques et paysages
d'élevage dans les pays bourguignons et rhodano-alpins (1° au XVII 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
Monneaux. Bulletin de la Société vaudoise des sciences naturelles 73 (351): 247-262.

BURNAND, J.-D., CHERIX, D., MORET, J.-L. & DE ROGUIN, L. 1977. La faune du marais des
Monneaux. I. Batraciens, Oiseaux et Mammifères. Bulletin de la Société vaudoise des
sciences naturelles 73 (352): 351-368.

BURNAND, J.-D., CHERIX, D., MORET, J.-L. & DE ROGUIN, L. 1978. La faune du marais des
Monneaux. III. Les Lépidoptères. Bulletin de la Société vaudoise des sciences natu-
relles 74 (354): 125-138.

CHAIX, L., OLIVE, C., DE ROGUIN, L., SIDI MAAMAR, H. & STUDER, J. (éds) 1995. L’anımal dans
l’espace humain, l'homme dans l’espace animal. Actes du 5° colloque international
HASRI, Muséum de Genève, 23-25.11.1994. Anthropozoologica 21, 287 pp.

LOUIS DE ROGUIN (1948-1998) 769

DE ROGUIN, L. 1978. «In parvo multa» ou le voyage du naturaliste Henri de Saussure aux
Antilles et au Mexique de 1854 à 1856. Musées de Genève 188: 18-22.

DE ROGUIN, L. 1980. La chasse au chamois dans le Jura vaudois. Diana 9: 363.

DE ROGUIN, L. 1982. Les chamois du Jura vaudois. SOS Nature 10: 14-15.

DE ROGUIN, L. 1983. Biologie et gestion d’une population de chamois (Rupicapra rupicapra L.)
du Jura vaudois. Thèse de doctorat ès sciences, Université de Lausanne (dir.: Prof. W.
Huber, Uni. Berne). Conservation de la faune et Section protection de la nature et des
sites du canton de Vaud, éds, Lausanne, 63 pp.

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.

Revue suisse de Zoologie 93 (4): 1009-1022.

DE ROGUIN, L. 1988. Expéditions Thai-Maros 85 et Thaï 87: Mammifères. Exp. A.P.S. Asie sud-

est, travaux scientifiques l: 47-52.

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, Besancon, 84 pp.

DE RoGuIN, L. 1990. Les micromammiferes de Portout, pp. 54-56. In: PERNON, J. & C. (eds).
«Les potiers de Portout, production, activités et cadre de vie d’un atelier au V° s. ap.
J.-C. en Savoie». Revue d’Archéologie Narbonnaise, suppl. 20, Editions CNRS, Paris,
220 pp.

DE ROGUIN, L. 1990. Mammifères et Oiseaux. /n: «Sentier naturaliste, Vallon de la Roulavaz
(Dardagny)», pp. 48-52. Série documentaire no. 27, Conservatoire et Jardin bota-
niques, Genève, 89 pp.

DE ROGUIN, L. 1991. Données historiques nouvelles sur la présence du rat noir Rattus rattus
(L.) en Europe occidentale. Jn: LE BERRE & LE GUELTE (Eds). «Le Rongeur et
Espace». Actes Colloque International, Lyon 1989, pp. 323-325. Editions R.
Chabaud, Paris, 362 pp.

DE ROGUIN, L. 1992. Les micromammiferes, p. 135. In: RAYNAUD, F. (éd.). Documents d’Ar-
chéologie en Rhône-Alpes No 6: «Le Chateau 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. Jn: «Formes et fonctions de l’habitat castral 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“ siècle.» Ed. Maison des Sciences de
l'Homme, Paris, 416 pp.

770 LOUIS DE ROGUIN (1948-1998)

DE ROGUIN, L. 1995. Le Rat noir Rattus rattus L., pp. 288-292. In: HAUSSER, J. (éd.).
Mammiferes de la Suisse. Birkhäuser, Basel.

DE RoGuIN, L. 1995. Le Rat surmulot Rattus norvegicus (Berk), pp. 283-287. In: HAUSSER, J.
(éd.). Mammiferes de la Suisse. Birkhäuser, Basel.

DE ROGUIN, L. & STUDER, J. 1991. Le rat noir a l’äge du Bronze final. Revue de Paléobiologie,
Geneve 10 (1): 79-83.

DE ROGUIN, L. & WEBER, C. 1985. Stenoderma tolteca Saussure, 1860 (Mammalia, Chiroptera):
proposed suppression of the neotype and validation of the rediscovered holotype. Z. N.
(S) 2466. Bulletin of zoological nomenclature 42 (3): 302-303.

DE ROGUIN, L. & WEBER, C. (eds) 1993. Henri de Saussure: Voyage aux Antilles et au
Mexique, 1854-1856. Editions Olizane, Genéve, 513 pp.

JUSTINE, J.-L. & DE ROGUIN, L. 1990. Capillaria murissylvatici (Nematoda, Capillariinae),
parasite d’un rongeur du Baluchistan iranien. Bulletin du Museum national d’Histoire
naturelle, Paris, 4° ser., sect. A. 12 (1): 19-33.

MOESCHLER, P., STUDER, J., BENIER, C., BURCKHARDT, D., LÖBL, I., DE ROGUIN, L. & WEBER, C.
1991. Nature enjeux. Musees de Geneve 314: 13-17.

WEBER, C. & DE ROGUIN, L. 1983. Notes inédites de Henri de Saussure sur les types de deux
rongeurs du Mexique: Microtus m. mexicanus (Saussure) et Reithrodontomys s. sumich-
rasti (Saussure). Revue suisse de Zoologie 90 (3): 747-750.

Francois 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! & Renata DE ANDRADE?

! Muséum d'histoire naturelle, case postale 6434, CH-1211 Genève 6, Switzerland

? Departamento de Zoologia, Instituto de Biociencias da USP, c.p. 11461,
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
strinatii, 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: 19 (holotype) 34 19 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) (23 1% 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 clavate, 4+1 nearly smooth preocular ones on anterior,
7-10 on posterior margin; tergites (XI excepted) divided, scaly sculpture, setae clavate,
longer on last tergites, 5-6 setae on posterior margin, IH-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, IT 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 clavate 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, serrula 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 (34/39 ): 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 773

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.

ist cib
oest ZISD cesh
oeb

Fics 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 clavate 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 I
subapical branch, serrula exterior 10 blades, flagellum 4 setae. Palps: femur 2,2 times
as long as broad (0.21mm/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 ¢ 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)

I 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
RAULONCAVESEE PNR DS A ER LORS ee 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
CATATINANE tato Lao PRE PAS IR MON ER RARES plaumanni Beier

- Length of palpal femur 0.40-0.42mm; anterior transverse furrow in
basal third of carapace; serrula exterior with 15 blades. Argentina,

Re RE NN ia AGE EN birabeni Feio

BIOLOGY

Maxchernes iporangae n. sp. is very abundant on frugivorous bat guano piles
in Alambarı 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
of Arachnology, 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.

PINTO-DA-ROCHA, R. 1995. Sinopse da fauna cavernicola do Brasil (1907-1994). Papeis
Avulsos de Zoologia, Sao Paulo, 39 (6): 61-173.

TRAJANO, E. & GNASPINI-NETTO, P. 1991. Composigao da fauna cavernicola brasileira, com
uma analise preliminar da distribucao dos taxons. Revista brasileira de zoologia 7(3):
383-407.

PMR EL ENTER AT AD OT WAN

aA in Ihm?
nf hi to quit! L'ala che 11 LA,
pray hd | Wu siti ie [TOI Ps AT

REVUE SUISSE DE ZOOLOGIE 105 (4): 777-787; décembre 1998

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 sıx 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 | 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 2 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 d 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,
14.1V.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,
14.1V.1985, leg. Smetana (MHNG, cAss); 7 ex., Nepal, Annapurna mountains, Lamjung Himal,
below Taunja Danda, 2350m, 6.V.1996, leg. O. Jager (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 d 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

Fics 1 a - d: Masuria plumbea Cameron: labium (a); maxilla (b); mandibles (c); labrum (d).
Scales: 0.2 mm.

780 VOLKER ASSING

I, | I
St LI

Fics 2 a - g: Masuria loebli Pace: aedeagus in lateral and in ventral view (a); flagellum of internal
sac (b); spermatheca (c); posterior part of & tergum (d) and sternum VIII (e); posterior part of £
tergum (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.

3: 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).

2: hind margin of tergum VIII weakly pointed, that of sternum VIII more
strongly convex than in d (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 d: NEPAL, Khandbari Distr., Induwa Khola Valley, 2050m, 16.1V.1984, Smetana &
Löbl (MHNG).

Paratypes: 1d, 29 2, 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

Fics 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
d tergum (d) and sternum VIII (e); posterior part of 2 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).

3: 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).

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 d : NEPAL, Khandbari Distr., Induwa Khola Valley, 2050m, 16.1V.1984, Smetana &
Löbl (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

Fics 4 a - h: Masuria 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
d tergum (d) and sternum VIII (e); posterior part of 2 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.
d : 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.
2: 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. Figs5a-g
Holotype d: NEPAL, Rasuwa Dis., Langtang Kh. Vall., 2.5km E Syabru, 1730m, 14.1V.1985,
A. Smetana (MHNG).

Paratypes: 1d, 12, same data as holotype (d 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.

d: 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).

2: 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. Sc.

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

Fics 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 d tergum (d) and sternum VIII (e); posterior
part of 2 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.

ACKKNOWLEDGEMENTS

I am indebted to Dr. Ivan Löbl (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.

rn Lie ws ao 5
rd WU too | DA
box cs

fi | = titi) er
ta Apia ALES ; "né È AR
Ab send en 0 TR Ber Lu

lat

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. LOURENCO* & 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-1211 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.
SıssoM (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
(Birula), 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 11.05.1998

790 W. R. LOURENCO & 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), I/1962 (Anderson leg.): 6 males, 4 females. Kavahari, XII/1958
(Anderson leg.): 1 male-juvenile, 1 female. Mirpur-Sakro, south of Karachi, II/1962 (Anderson
leg.): 2 males. Near Karachi, 11/1962 (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: pectinal 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

Fic. 1

Compsobuthus rugosulus, female from Hyderabad in dorsal view.

792 W. R. LOURENCO & L. MONOD

un
SUES”

REDESCRIPTION OF COMPSOBUTHUS RUGOSULUS 793

Soo 09°.
250%
So

000
00__00000
is)

Le)

oo 0

der

000 00PQ,

doo,

(E)

CIS
° o DISS

Fics 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

external aspects.

Fics 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. LOURENCO & 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, orthobothriotaxy (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 3,0 4,3
- anterior width IS) 2,8
- posterior width 3,4 4,8
Metasomal segment I:
- length 169 2,4
- width 2,1 2,6
Metasomal segment V:
- length 32 4,0
- width 1,5 DID,
- depth IES ZA
Vesicle:
- width 1,2 1,8
- depth 151 165
Pedipalp:
- Femur length 25 SÒ]
- Femur width 0,8 12
- Tibia length 352 4,1
- Tibia width 13 19,
- Chelae length DD. 12
- Chelae width 1,4 1,8
- Chelae depth 165 197

Movable finger:
- length 3,6 SI

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
15-15 I 3
16-16 Il
16-17 |
17-16 |
17-17 2
17-18 I
18-17 I

18-18
19-19

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 “Departement des affaires culturelles de la Ville de
Geneve” for financial support.

REFERENCES

BIRULA, A. 1905. Beiträge zur Kenntniss der Skorpionenfauna Persiens. Bulletin de l’Académie
Imperiale des Sciences de St. Pétersbourg, 5° serie 23 (1-2): 119-148.

BIRULA, A. 1910. Ueber Scorpio maurus Linné und seine Unterarten. Horae Societatis Entomo-
logicae Rossicae 35: 115-192.

BIRULA, A. 1917. Arachnoidea Arthrogastra Caucasica. Pars I. Scorpiones. Zapiski Kavkazs-
kogo Muzeya (Mémoires du Musée du Caucase), Tiflis Imprimerie de la Chancellerie
du Comite pour la Transcaucasie, ser. 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.

HaAB1B1, 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): 113-140.

796 W. R. LOURENCO & L. MONOD

PEREZ, S. M. 1974. Un inventario preliminar de los escorpiones de la region paleärtica y claves
para la identification de los géneros de la region paleärtica occidental. Universidad
complutense de Madrid, Faculdad de ciencias, Departamento de zoologia, Catédra de
artröpodos, 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: PoLis,
G. A. (ed.). The Biology of Scorpions. Stanford University Press.

SıssoM, 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’Algerie 27 (1):
66-100.

VACHON, M. 1952. Etudes sur les scorpions. Publications de l’Institut Pasteur d’Algerie, 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° ser. 35
(2): 161-166.

VACHON, M. 1966. Liste des scorpions connus en Egypte, Arabie, Israll, Liban, Syrie, Jordanie,
Turquie, Irak, Iran. Toxicon 4: 209-218.

VACHON, M. 1973. Etude des caracteres utilisés pour classer les familles et les genres de Scor-
pions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et
types de trichobothriotaxie chez les Scorpions. Bulletin du Museum National d'Histoire
Naturelle, Paris, 3° ser. 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, ser. 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)

Najla ZEIDAN-GEZE! & Daniel BURCKHARDT?
| Faculté des Sciences (2), Université Libanaise, BP 90656, Jdeidth El Maten, Lebanon.
2 Naturhistorisches Museum, Augustinergasse 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 olivina”, 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 & ONUCAR 1993; GÜCLÜ & BURCKHARDT 1996); Jordan 33
(AL-KHAWALDEH et al. 1997); Syria 6 (TALHOUK 1969; MUSTAFA 1991); Israel 66
(BODENHEIMER 1937; HALPERIN ef al. 1982; HALPERIN 1986; HALPERIN er 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 er 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-GEZE & 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.

N
MICRO
DE 9

È
= \
Tripoli & = 5
ad {
S \
N \
Batroun x

9 Bcharré — >
4
Jbail n BEKAA ,

Jounie \ Baalbek ©

Beyrouth

or MATEN >

Saida D

Fic. 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, Geneve (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

I Antennal flagellar segments flattened bearing long black setae. Male
proctiger distinctly 2-segmented. On Ficus carica.
SA RR RARE Homotomidae: Homotoma ficus (L.)
- Antennal flagellar segments more or less cylindrical. Male proctiger
jEseemented=sometimessindistinctly subdivided e n 2
2 Forewings with vein R+M+Cu, bifurcating into Rand M+Cu;; if tri-
furcating then anal break close to apex of vein Cu,,, and metabasitarsus
with 1 or 2 black spurs. Costal break and/or pterostigma often deve-
lame dR swiidaes gcse crete ea 2 N a Ta 3,
- Forewings with vein R+M+Cu, trifurcating into R, M and Cu, or
bifurcating into R+M and Cu;; anal break distant from apex of vein
Cu,,; costal break and pterostigma always absent. Metabasitarsus

WIN OVADA CKESPUTSSEANTTOZICACI ae E Re CO 24
3 Metacoxae without meracanthus; trochanteral cavity with weakly scle-

rowsedtubercles | RhinOcOlinde oars cus ee eI ee ee eee oi n.
- Metacoxae with horn-shaped meracanthus; trochanteral cavity without

ADERIRE oi E i RI NE SM SER 5
- 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.

SERRE TESA: A 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

RISIOCIGSPpr en: Agonoscena pistaciae Burckhardt & Lauterer
5 MertexsslonserithanlarseIiyınaezlwian eine RO RI 6
= WICTLO XN ALO CTC An Once PP ence PE II NI 7
6 Antennal segment 2 long pear-shaped. Forewings oval. On Juncus spp.

i RIE EEE RIT EINEN AR ane Ser er Livia juncorum (Latreille)

800

N. ZEIDAN-GEZE & D. BURCKHARDT

Antennal segment 2 short, cylindrical. Forewings oblong with sub-
parallel fore and hind margins. Host plant unknown.

RA ee kien. teks co tars one Have LR AT Livia mediterranea Loginova
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
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

(NIG ADAG) NUT Ae ee Asse 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.

TE N PEN A Euphyllura phillyreae Foerster
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... nasser u nein gi lite las NE 10
Basal metatibial spine often developed. Apical metatibial spurs always
srouped. Vertex trapezoidal: head with genal cones. Renn 12
Proepimeron much larger than proepisternum; propleural suture dia-

sonale Onwiamanix sppsse see Ar. Colposcenia kiritshenkoi Loginova
Proepimeron and proepisternum subequal; propleural suture vertical.
Craspedoleptar. «sar Mar ana es ti ae Sue nue ee TE 11

Antennae 10-segmented. On Leontodon autumnale.

SERGE re Re RAE Ape EN a de my Craspedolepta sonchi (Foerster)
Antennae 9-segmented. On Achillea, Anthemis, Pyrethrum spp.

SEES ASE SRE AE © a Craspedolepta pontica Dobreanu & Manolache

Metabasitarsus without or with a single black spur. .................... 13
Metabasitarsus with two, black Spurs. SNA IS

Male proctiger with posterior wing-like lobes. Forewings with long cell

mj, and high cell cu,,, vein Cu; longer than Cu,. On Acacia, Albizia

Sol SO e ee Acizziinae: Acizzia uncatoides (Ferris & Klyver)
Male proctiger without posterior wing-like lobes. Forewings with shor-

ter and lower M,,, and Cu,, cells, vein Cu,,, shorter than Cu,.
Arytaininae.Pp.:P. u... nen wenn ne rayon ie ie CU TOC ET 14
Genal cones about half vertex length. On Calicotome spp.

MA ni IRE Arytainilla cytisi (Puton)
Genal cones longer than vertex length. On Spartium junceum.

VR RI e o AE NEVA PER Lala ae Livilla spectabilis (Flor)

PSYLLOIDEA FROM LEBANON 801]

Forewings with long cell m,,, and high cell cu,,. Male parameres
truncate apically. On Astragalus spp.

PICEA RAE Re Arytaininae: Cyamophila stoklosai Klimaszewski & Lodos
Forewings with shorter and lower m,,, and cu,, cells. Male parameres

diferentes yilinae ser Ste shi. aree. ie were at el pale 34. 16
Forewing membrane yellow, bearing irregularly, densely spaced sur-
HAC ERS PILI C Seeley as ae Ae Casa OR RR meee o> 17

Forewing membrane colorless or with brown pattern; surface spinules
sparse forming regular rhombi or squares, or reduced to narrow stripes

mmathesmiddleofuthe:cellssGacopsyllapape ne an tate ten ee 18
Genal processes more than half vertex length. Antennae less than 2.0
times head width. On Ostrya carpinifolia.............. Psylla colorata Low
Genal processes less than half vertex length. Antennae more than 2.0
timesiheadmwidths@n Alnus SpD AE rss NE Psylla foersteri Flor

Pattern of forewings consisting of well-defined dark patches at the

apices of pterostigma and veins Rs, M,,5, M3,4, Cu, and Cuj,, as well

as a more or less interrupted band from apex of vein Rs to the middle of
vemuCurr On Cercissiliquastrum. Der Teti Cacopsylla pulchella (Löw)
Battembolstore wines differentave aber Han oe Am Il Pe. 19
Forewings with brown band along outer margin. On Crataegus spp.

FORTAN UOTE NT. DIGO SE RER TE SER Cacopsylla mariannae (Baeva)
Forewings without brown band along outer margin. ................... 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
Combinanononceharaciers absente gate. eee 22
Hindmargin of parameres, in lateral view, convex. Dorsal margin of
female proctiger weakly convex. On Salix spp.

ob) og RANDE Ee REE RRR har ce Cacopsylla brunneipennis (Edwards)
Hindmargin of parameres, in lateral view, concave. Dorsal margin of
female proctiger weakly concave. On Salix spp.

Dur li] DRS APTE ae EROI al ce ae PRL à Cacopsylla subpropinqua (Loginova)

Forewings with brown clavus. On Pyrus spp....... Cacopsylla bidens (Sulc)
Forewings with clavus of the same colour as surrounding membrane. ... . . 23
Surface spinules forming broad fields in apical half of forewings. On

IBN GUSAST Docc ee hee ae RGR ees RI EE Cacopsylla pyrisuga (Foerster)
Surface spinules forming narrow stripes in apical half of forewings. On

Khamnus.spD... aan. Ba I or. ee. ve Cacopsylla myrthi (Puton)
Forewings with vein R+M+Cu, bifurcating in veins R+M and Cu..

kos@plantunknownk eee RESA. Eutrioza opima Loginova
Forewings with vein R+M+Cu, strictly trifurcating into R, M and Cu,..... 25
Metdtibiaesvuithalkraiblacktapiealispurs2Triozanpapr rue. 26

Meratibiae with 2 iblacksapicalispurses 2 nn Da

N. ZEIDAN-GEZE & D. BURCKHARDT

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.

BEE N RNA WERE EEE FIRE Trioza urticae (L.)
Bifurcation of vein M of forewings distinctly distal to line from apices
ofevemsRsito Cu) En TS 28
Bifurcation of vein M of forewings on, or proximal to line from apices

OF veins RSito CU. 2. NEUTRE OI ARE EEE TE
Vein Rs of forewings sinuous to almost straight; vein M,,, more than
twice the length of vein M;,4; fore margin of forewings strongly
arched, hind margin only weakly curved. Male proctiger with large
posterior lobes. Female proctiger long. On Rubus spp.

RUE DEE ARE SAA. RATTEN Phylloplecta trisignata (Löw)
Vein Rs of forewings concave; vein M,,, less than twice as long as

M,,,4; fore margin of forewings only slightly stronger curved than hind
margin. Hind margin of male proctiger weakly produced. Female geni-

Sil

tal Short Triozaip! pl. TIER NOUS RE ee 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
Surface spinules present in all cells of forewings, forming large fields.

ONdeGiduous iO er CUSISpp ES EEE Trioza remota Foerster
Surface spinules largely reduced in apical half of forewings. On ever-
CECCHI OUT CUSEILCK IN Pane ey OCRA Trioza ilicina (de Stefani)

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

AN GUSTHONGs 05 S050 ARI RAR 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.................... 52
Forewings bearing surface spinules. On Salix spp.

SEIN Yen RSR atea ee petite MER Bactericera albiventris (Foerster)
Forewings without surface spinules in apical half...................... 38
Genal cones longer than half vertex length. On Salix spp.

RE RI MR TA ek ER A Bactericera curvatinervis (Foerster)
Genal cones shorter than half vertex length. Polyphagous.

RL NE RUSSE. EU VARS e at Bactericera nigricornis (Foerster)

PSYLLOIDEA FROM LEBANON 803

BISMORSPECIES

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, 24.1X.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.V1.1981 and 2.V.1983; Dahr El Souan: Maten,
1.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 & ONUCAR 1993).

Host plants. Astragalus spp. (Fabaceae).

804 N. ZEIDAN-GEZE & D. BURCKHARDT

Livilla 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.V11.1995 and 2.VIII.1996; Kartaba: Kesrouan, 22.1V.1985; Meyrouba: Kesrouan,
24.1X.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.V1.1981 (MHNL).

Distribution. Armenia, Azerbaijan, former Czechoslovakia, Georgia, Greece,
Israel, Kazakhstan, Kyrgyzstan, Romania, Russia (European part), Tajikistan, Turkey,
Turkmenistan (GEGECHKORI & LOGINOVA 1990).

Host plants. Achillea, Anthemis, Pyrethrum spp. (Asteraceae).

Craspedolepta sonchi Foerster

Material examined. Bhamdoun: Aley, 21.1X.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, 9.11.1981 (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 21.1X.1979; Kfar-
Katra: Chouf, 8.11.1981; Bhannés: Maten, 4.1V.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.V1.1986 and 22.VI.1986; Yah-
chouch: Kesrouan, 16.VI.1981; Zaarour: Maten, 1.VI.1997; Naas: Maten, 19.V1.1997;
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-GEZE & 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.[X.1994; Baskinta: Kesrouan, 22.1X.1994
(MHNL, NHMB, MHNG).

Distribution. Tadzhikistan, Turkey (GEGECHKORI & LOGINOVA 1990;
BURCKHARDT & ONUCAR 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,
17.1X.1994; Baskinta: Kesrouan, 19. IX.1994; Hajar El Afrit: Kesrouan, 17.1X.1994,
24.1X.1994 and 16.1X.1994; Hrajel: Kesrouan, 27.VI.1994; Jabal Moussa: Kesrouan,
15.V1.1997 and 21.VI.1994; Kartaba: Kesrouan, 22.X1.1985; Meyan: Kesrouan,
23.1V.1983; Meyrouba: Kesrouan, 5.V.1985 and 24.[X.1994; Yahchouch: Kesrouan,
16.V1.1981; 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 (Löw)

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 siliquastrum 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, 24.1X.1994; Achkout: Kesrouan,

17.1X.1994; Jabal Moussa: Kesrouan, 15.VI.1997 and 21.VI.1994; Jaj: Batroun, IV-
V1.1984 (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.V1.1997 (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,

16.1X.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.V111.1985 (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.V1.1986 and 15.VI.1997; Naas: Maten, 14.VI.1997; Broummana: Maten,
14.V1.1997 (MHNL, NHMB, MHNG).

Distribution. Israel, Jordan (AL-KHAWALDEH 1997).

Host plant. Pistacia vera L. (Anacardiaceae).

808 N. ZEIDAN-GEZE & D. BURCKHARDT

TRIOZIDAE

Bactericera albiventris (Foerster)

Material examined. Zaarour: Maten, 28.VI.1997; Kfar Debiane: Kesrouan,
31.V11.1985 (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.VII.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.X1.1983; 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 809

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.X1.1985
(MHNL).

Distribution. Croatia, France, Israel, Italy, Spain (CONCI er al. 1996).

Host plant. Rhamnus alaternus L. (Rhamnaceae).

Trioza neglecta Loginova

Material examined. Broumana: Maten, 12.V1.1997 (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, 31 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-GEZE & 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); SY = Syria (based on records by TALHOUK 1969; MUSTAFA
1991).

Lebanese species PA ME EM IN IS SM

Acizzia uncatoides (Ferris & Klyver) + +
Agonoscena pistaciae Burckhardt & Lauterer +
Arytainilla cytisi (Puton) + +
Bactericera albiventris (Foerster) +
Bactericera curvatinervis (Foerster)
Bactericera ni gricornis (Foerster)
Cacopsylla bidens (Sulc)

Cacopsylla brunneipennis (Edwards)
Cacopsylla mariannae (Baeva)
Cacopsylla myrthi (Puton) + +
Cacopsylla pulchella (Löw) + +
Cacopsylla pyrisuga (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 +
Eutrioza opima Loginova +
Homotoma ficus (L.) + + +
Livia juncorum (Latreille) +

Livia mediterranea Loginova +

Livilla spectabilis (Flor) +
Megagonoscena gallicola Burckhardt & Lauterer +
Phylloplecta trisignata (Löw) +

Psylla (Baeopelma) colorata Löw +

Psylla (Baeopelma) foersteri Flor +

Trioza alacris Flor + +
Trioza ilicina (de Stefani)
Trioza marginepunctata Flor st +
Trioza neglecta Loginova ali

Trioza remota Foerster +

Trioza urticae (L.) + +

ail

++++++

+ +++ ++
+
+
w

+ + + +

x Recorded as Agonoscena targionii (Lichtenstein) by TALHOUK (1969).

“ Recorded as Psylla pyricola 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 Fast. A similar disjunct known distribution has Cacopsylla
saligna (Loginova) in Spain and Kazakhstan (LAUTERER & BURCKHARDT 1997).

PSYLLOIDEA FROM LEBANON 81]

Two species, viz. Acizzia uncatoides and Trioza neglecta, have been intro-
duced from Australia and East Asia respecitively. Homotoma ficus 1s 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 (Löw) (HODKINSON & HOLLIS 1987). The species was originally
recorded from “Syria: Kaifa (Reitter)” (Löw 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.

REFERENCES

AL-KHAWALDEH, M., KATBEH-BADER, A. & BURCKHARDT, D. 1997. Psylloidea (Hemiptera) of
Jordan. Zoology in the Middle East 15: 71-82.

Anonymous 1997. Carte du Liban. Echelle 1 : 200'000. 3" edition, GEOprojects, Reading and
Beyrouth.

BODENHEIMER, F. S. 1937. Prodromus Fauna Palestinae. Mémoires de l'Institut d'Egypte Cairo
33: 286 pp.

BURCKHARDT, D. 1987. Jumping plant lice (Homoptera: Psylloidea) of the temperate neo-
tropical region. Part 1: Psyllidae (subfamilies Aphalarinae, Rhinocolinae and Aphala-
roidinae). Zoological Journal of the Linnean Society 89: 299-392.

BURCKHARDT, D. 1989. Les Psylles (Insecta, Homoptera, Psylloidea) de I’ Algérie. Archives des
Sciences, Genéve 42: 367-424.

BURCKHARDT, D. 1994. Psylloid pests of temperate 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.

812 N. ZEIDAN-GEZE & D. BURCKHARDT

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. & Mirsup, D. 1998. Psylloidea (Insecta: Hemiptera) of the Arabia Peninsula.
Fauna of Arabia 17: in press.

BuRCKHARDT, D. & Onucar, A. 1993. A review of Turkish jumping plant lice (Homoptera,
Psylloidea). Revue suisse de Zoologie 100: 547-574.

Concı, 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.

Concl, 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.

GÜÇLÜ, S. & BURCKHARDT, D. 1996. New records of jumping plant-lice from Turkey
(Hemiptera, Psylloidea). Entomofauna, Zeitschrift fiir Entomologie 17: 381-384.
HALPERIN, J. 1986. An Introduced Psyllid Injurious to Acacia Trees. Phytoparasitica 14: 234-

235)

HALPERIN, J., 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). /srael 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. Cambrideo
University Press, reprinted with corrections, 707 pp.

MUSTAFA, T. M. 1991. The Psyllids (Homoptera: Psylloidea) of Jordan. /ragi Journal of
Science 31: 139-145.

OSSIANNILSSON, F. 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. Fauna
Entomologica Scandinavica 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 countries.
Monographien zur angewandten Entomologie. Beihefte zur Zeitschrift für angewandte
Entomologie 21: 99- 102.

WHITE, I. M. & Hopkinson, I. D. 1985. Nymphal taxonomy and systematics of the Psylloidea
(Homoptera). Bulletin of the British Museum (Natural History), Entomology 50: 153-
SOI

REVUE SUISSE DE ZOOLOGIE 105 (3): 813-822; decembre 1998

Les organes producteurs de phéromones de quelques Hespérides
(Lepidoptera, Hesperiidae, Hesperiinae)!

Jean WUEST
Muséum d’histoire naturelle, Case postale 6434, CH-1211 Geneve 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 7h. 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
phéromone des males de quelques papillons de jour du genre Argynnis sensu lato.
Nous avons poursuivi notre étude au moyen du microscope électronique à balayage
sur quelques especes de la famille des Hespérides, sous-famille des Hesperiinae, qui
présentent un appareil tout a 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 a l’Assemblée annuelle de la Société entomologique suisse
(Geneve, 13-14 mars 1998).

Manuscrit accepté le 29.06.1998

814 JEAN WÜEST

le röle de support “macroscopique” de l’odeur. Ces éléments se comportent comme
une poudre et peuvent étre dispersés par le male 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 plutöt 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
lineola 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; GRASSÉ 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 1c1. Notre étude porte
sur trois espèces qui illustrent bien la complexification de l’appareil et des écailles.

MATÉRIEL ET MÉTHODES

Les trois espèces étudiées [Thymelicus lineola (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 a
lor, dans un microscope électronique à balayage Zeiss 940A.

L'APPAREIL A PHEROMONE

Chez les males des Hespérides de couleur générale orange (représentant la
sous-famille des Hesperiinae, genres Hesperia et Thymelicus présentés ici), on peut
remarquer a 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’écailles 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 a la face inférieure de l’aile, les massifs d’androconies s’y
retrouvent chez Th. lineola. Les tissus de l’aile sont fortement épaissis a 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 lineola, 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 A PHEROMONES DES HESPERIDES 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 a la zone des osmophores (Figs 5 & 6), qui deviennent plus larges et
s’orientent perpendiculairement a la nervure médiane. Mais la zone des écailles a
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’arriere (Figs 11 & 12). Les
massifs d’osmophores, qui ont pourtant la méme 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 trés 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 a 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 À PHEROMONE

Chez Thymelicus lineola, les androconies ne sont pas modifiées en écailles
porteuses d’osmophores. Les éléments diffuseurs de pheromone 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 déhiscence 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’écaille a tendance à se boursoufler, préfigurant une zone de
déhiscence (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’écailles très allongées, filiformes, présentant sur leur longueur un
certain nombre de restrictions ou zones de déhiscence. 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 (GRASSÉ 1975).

816 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 troncons 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 des la base
la structure des segments osmophores (Figs 7 & 14).

La forme des osmophores est assez constante d’une espece a l’autre. Par
contre, leur longueur est extrémement variable d’un osmophore a 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
déhiscence. |

D'une manière générale, les osmophores présentent une partie renflée terminée
à chaque extrémité par les zones de déhiscence. 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 déhiscence, 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 emmmélées, sans qu’un réseau d’osmophores collés se
mette en place comme chez H. comma. La zone d’écailles à osmophores consiste
presque exclusivement en ce type d’écailles, 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 A PHEROMONES DES HESPERIDES 817

présentent un pas plus ou moins serré et peuvent faire 1,5 a 3,5 fois le tour d’un
segment de 20 um de long. Les extrémités du segment sont fusiformes et |’ ouverture
est laterale (Fig. 15). Par contre, les cötes présentent une ornementation transversale
particulière sous la forme de petits bourrelets (Fig. 16). La base des écailles a
osmophores est beaucoup plus courte que les osmophores, mais présente la même
ornementation superficielle. Les osmophores de cette espece ont la faculté de se
recoller 4 un autre segment et de former un réseau tres 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 a se localiser
en surface de la zone a androconies, peut-étre poussé vers le haut par les écailles
banales (Fig. 13).

DISCUSSION

Parmi les trois especes étudiées, on peut distinguer une gradation nette dans la
complexité de l’appareil a phéromone. Chez Thymelicus lineola, la zone a andro-
conies se présente simplement comme une zone a 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 espece 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 a osmophores sont plus larges et s’orientent
perpendiculairement a la zone a osmophores, préfigurant l’apparition d’écailles de
recouvrement. Mais les osmophores restent bien visibles de l’extérieur. C’est chez
Hesperia comma enfin, que |’ 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’écailles 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’écailles 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

Fics 1-4. Thymelicus lineola. - 1. Zone a phéromone de l’aile antérieure (face supérieure). On
distingue directement les massifs d’androconies. G = 40x. - 2. Detail de la surface des massifs
d’androconies. Il n’y a pas d’osmophores. G = 600x. - 3. Zone a androconies en coupe,
montrant quelques rares restrictions sur la longueur des écailles filiformes. G = 430x. -
4. Androconie présentant une restriction et amorgant le processus de formation des osmo-
phores. G = 6300x.

Fics 5-10. Thymelicus acteon. - 5. Zone a phéromone. G = 12x. - 6. Détail de la zone a
pheromone. 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 Ä PHEROMONES DES HESPERIDES 819

JEAN WÜEST

820

EI,

4% À

GIG,
7; 4
208

ORGANES A PHEROMONES 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’eviter un gaspillage des principes actifs, ou d’assurer leur diffusion seulement dans
certaines phases du comportement de cour. Mais il est bien sür illusoire de penser
vérifier cette hypothèse en visualisant la position des écailles de recouvrement
pendant les différentes étapes de l’accouplement, méme si BARTH (1944) a décrit les
déplacements des écailles de l’appareil des Argynnes pendant le vol, permettant la
libération des phéromones. Il serait interessant de vérifier si on retrouve une telle
architecture des appareils 4 phéromone (massifs d’androconies recouverts par des
écailles de couverture) chez d’autres groupes de rhopaloceres: chez les papillons de
jour, les phéromones jouent un röle 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 déhiscence des autres
espèces, et PIVNICK ef al. (1992) ont montré que ces zones de déhiscence semblent se
multiplier au cours de la vie imaginale, sans en expliquer le mécanisme. Thymelicus
acteon, lui, présente deux types d’écailles 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 Th. acteon, avec des
extremites fusiformes. Les écailles normales parsemées dans la zone 4 osmophores
sont filiformes ou légèrement aplaties, mais ne présentent pas d’intermédiaires avec
les androconies à osmophores.

Un autre critere concerne la fragilité des androconies a osmophores. Chez
Th. lineola, les limites entre segments de type osmophore n’étant que partiellement
indiquées, ces Ecailles ne se rompent pratiquement pas et restent entieres dans la zone
a androconies, quoique PIVNICK et al. (1992) démontrent que les cassures, comme les
zones de déhiscence, se multiplient avec l’äge de l’imago. Chez Th. acteon, les

Fics 11-16. Hesperia comma. - 11. Zone a 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. Detail d’une coupe de chambre a androconies montrant les écailles filiformes
non modifiées et les écailles 4 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. À 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. /n: GRASSE P. P. 1975. Traité de Zoologie, tome
VII, fasc. III, 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 Societe lepidoptero-
logique de Geneve 3: 122-128.
SELLIER, R. 1971. Etude morphologique en microscopie électronique à balayage de quelques

types d’androconies alaires chez les Lépidoptéres 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 a 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.

WEST, 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 espéces asiatiques du genre Bryaxis et quelques données
sur des especes connues (Coleoptera: Staphylinidae: Pselaphinae)

Sergei A. KURBATOV! & Ivan LOBL?
! Severodvinskaya 9-84, Moscou 129224, Russie.
2 Muséum d’histoire naturelle, Case postale 6434, CH-1211 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 & Löbl
and B. panda Kurbatov & Löbl.

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. koltzei (Reitter), est largement
répandue (COULON & Li 1995; KURBATOV & LOBL 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 (LOBL 1977; NOMURA & LEE 1993), 19 a Taiwan (LOBL &
KURBATOV 1996) et 34 au Japon (LOBL et al. 1998).

Basée sur des récoltes récentes, cette étude comporte la description d’une
espèce nouvelle de Bryaxis de |’ Altai russe et de six espèces nouvelles de Chine.

Manuscrit accepté le 11.09.1998

824 S. A. KURBATOV & I. LÖBL

D’autre part, une espece vraisemblablement nouvelle pour la science, mais non-
décrite en l’absence des caracteres essentiels du mäle, est signal&e de Birmanie, et de
nouvelles données sur la distribution de deux especes de Bryaxis sont fournies.

Abreviations utilisees:

CSKM Collection privee de S. A. Kurbatov, Moscou

MHNG Museum d’histoire naturelle, Geneve

ZMAN Muséum Zoologique de la Division Sibérienne de I’ Académie Russe de Sciences,
Novosibirsk

ZMUM Museum Zoologique, Moscou

DESCRIPTIONS

Bryaxis kimi sp. n. Figs 1, 12

Holotype d: 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
l’holotype, 2 4,10 2 (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. Téte densément et presque entierement ponctuée, les diamétres des points plus
grands que les intervalles entre eux. Moitie basale de la dépression frontale et
tubercules antennaires lisses. Lobe frontal large de 0,19-0,20 mm, très légèrement
rétréci en arriere. 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 d nettement plus longs que les tempes en
vue latérale, comportant 20-25 ommatidies; yeux © 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 a peine transverses, 9 aussi long et plus large que 8, 10 plus long et
plus large que 9, 11 plus long que 8 a 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, diamétres des points le plus souvent plus grands que
les intervalles entre eux.

d. 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.

2. 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 & Löbl, 1995. Elle en offre
l’ensemble des caractères, notamment la forme de l’édéage, très similaire à ceux de
B. emeicus dont elle differe nettement par le nombre et la forme des pieces sclérifiées
du sac interne de l’édéage, en particulier par les branches du sclérite apical plus
étroites et moins asymetriques.

Etymologie. L’espece 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 BRYAXIS 825

Fics 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 d: 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 d, 7 2; mêmes données mais 18.VII.1996, 4 8, 5 2; mêmes
données mais 2500 m, 23.VII.1996, 2 8,2 2; mêmes données mais 2500 M, 24.VII.1996, 1 3
(ZMUM, MHNG, CSKM).

Longueur: 1,45-1,55 mm; largeur maximale: 0,65-0,69 mm. Corps d’un brun
rougeätre 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 d plus longs que les tempes en vue latérale,
comportant 20 ommatidies environ; yeux 2 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 a peine transverses,
8 de la méme longueur, mais a peine plus large, 9 plus large et a peine plus long que
8, 10 plus long et plus large que 9, 11 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 diametres des points généra-
lement plus grands que les intervalles entre eux.

d. 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.

2. Scape cylindrique, environ 2 fois plus long que large, pédicelle ovale, plus
long que large, plus long que l’article 3.

Espece ressemblant a B. kimi par les caracteres externes, mais en différant par
la proportion des articles antennaires 6-11 et par la forme du scape chez le à.
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 pieces
sclérifiées symétriques du sac interne de |’ édéage.

Etymologie. L’espece 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 d: 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 © (ZMUM, MHNG).

Longueur: 1,5 mm: largeur maximale: 0,64-0,70 mm. Corps brun. Pubescence
comme chez les deux especes précédentes. Ponctuation sur la tete 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 d 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 827

tidies. Autres caracteres de la téte 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.

Sd. Articles 1 à 3 de l’antenne comme Fig. 14; scape long de 0,165 mm,
pédicelle long de 0,06 mm. Fémurs renfles; 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.

2. Scape et pédicelle comme chez les espèces précédentes.

Cette espèce ressemble a B. kimi par la forme du scape et du pédicelle 3. Elle
en diffère par la ponctuation de l’avant-corps plus dense et par l’édéage beaucoup
plus étroit, a sac interne symétrique, orné de nombreux stylets et d’une plaque apicale.

Etymologie. L’espece est dédiée a Vladimir A. Pletnev, entomologiste de
Moscou et ami du premier auteur.

Bryaxis ruidus sp. n. Figs 4, 15

Holotype d: Chine, province de Zhejiang, Tienmushan, 2.1X.1994, leg. G. de Rouge-
mont (MHNG). Paratypes: mêmes données que l’holotype, 1 4,2 2 (MHNG).

Longueur: 1,5-1,55 mm; largeur maximale: 0,63-0,70 mm. Corps brun, élytres
parfois plus clairs, brun rougeätres. Pubescence comme chez les espèces précédentes.
Téte comme chez B. pletnevi, mais dépourvue d’une carène occipitale. Yeux compor-
tant 22-25 ommatidies chez le ¢ 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 a peine plus long
que 9, 11 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.

d. 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
metatibias plus courte. Edéage (Fig. 4) long de 0,37 mm.

2. Scape cylindrique, à peu près 1,5 fois plus long que large, pédicelle ovale,
d’un tiers plus court que le scape.

L’espece ressemble a B. holciki Lobl, 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 tres fins et de pieces sclérifiées pus larges, progressivement
rétrécies vers I’ apex.

828 S. A. KURBATOV & I. LÖBL

Bryaxis mendax sp. n. Figs 5, 16

Holotype 9: Chine, province de Zhejiang, Tienmushan, 2.IX.1994, leg. G. de Rouge-
mont (MHNG). Paratypes: mémes données que l’holotype, 2 2 (MHNG).

Longueur: 1,45-1,55 mm; largeur maximale: 0,64-0,67 mm. Uniformément
brun, pubescence comme chez B. kimi. Téte comme chez B. ruidus. Yeux deux fois
plus longs que les tempes en vue latérale et comportant 25 ommatidies environ chez
le d; yeux £ 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 a peine
plus larges que longs, égaux, 9 plus large et plus long que 8, légérement transverse, 10
aussi long et plus large que 9, 11 plus long que 8-10 réunis. Pronotum a peine plus
large que long, sa ponctuation nettement moins dense que celle du vertex. Elytres a
ponctuation aussi dense que celle du pronotum, mais formée de points plus grands et
plus superficiels; les diametres des points en général un peu plus grands que les
intervalles entre eux.

d. Articles 1 à 3 de l’antenne comme Fig. 16; scape long de 0,10 mm, pédi-
celle long de 0,09 mm. Protibias tres legerement échancrés sur leur quart distal,
dépourvus de dent a 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 a l’apex.

2. Scape cylindrique, à peine plus long que large, pédicelle allongé, un peu
plus court que le scape.

Cette espece ressemble a B. ruidus, notamment par la conformation de la t£te.
Elle en differe par le pronotum a ponctuation nettement moins dense et, surtout, par
l’édéage. Les paramères sont échancrés subapicalement chez B. mendax, semblables a
ceux de B. smetanai Löbl; les pièces sclérifiées du sac interne, constituées d’une paire
de grandes dents latérales et d’une plaque centro-apicale, sont tres caractéristiques
chez ces especes.

Bryaxis fictor sp. n. Figs 6, 17

Holotype d: Chine, province de Shaanxi, Nonwutai, à peu pres 25 km sud de Xian,
400-800 m, 17.1X.1995, debris 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 subparalleles.
Bord antérieur du front incliné en avant. Centre des fossettes du vertex situé en arriere
du niveau du bord antérieur des yeux. Caréne occipitale presque absente, a 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, ä peine transverses, 9 plus long et a peine plus large que 8,
legerement 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

Fics 5a 11

Bryaxis, édéages, vue dorsale, et métatibias d, 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 (Aubé), édéage (8) et métatibia (9). Echelles = 0.1 mm (Figs 5 a 8); = 0.2 mm
(Figs 9 à 11).

830 S. A. KURBATOV & I. LÖBL

plus grands et plus superficiels que sur le pronotum; les diametres des points plus
grands que les intervalles entre eux.

d. 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 particulierement dense. Profémurs légerement mais nettement
déprimés a la base du bord inférieur, sur une surface a peu pres 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 espece présente des caractéres uniques sur le métasternum, les pro-
femurs, les antennes et l’édéage. La ponctuation particulière du métasternum et les
dépressions profémorales sont considérées comme étant liées au sexe d. Bryaxis
fictor semble étre isolé au sein du genre. Les lobes dorsaux sur les parameres
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 sclérite circulaire basal joint au sclérite
apical bifide.

Bryaxis valentulus sp. n. Figs 70s ENS

Holotype d: Russie, Altaï occidental, chaîne Ivanovsky, à 10 km au sud de Lenino-
gorsk, forêt clairsemée d’Abies, 1400 m, 30.V.1996, leg. R. Dudko (ZMAN). Paratypes: mêmes
données que l’holotype, 3 4, 1 2 (ZMAN, MHNG, CSKM).

Longueur: 2,05-2,2 mm; largeur maximale: 0,77-0,85 mm. Corps brun rou-
geätre ou uniformément brun. Pubescence double, à soies mi-dressées assez denses et
a soies plus longues, plus dressées et beaucoup moins nombreuses. Téte 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. Caréne occipitale atteignant
le bord postérieur de la dépression frontale. Yeux aussi longs que les tempes en vue
latérale, comportant 12-16 ommatidies chez d, un peu plus petits chez 9. Articles 2
et 3 de palpes maxillaires dépourvus de tubercules. Articles antennaires: 3-8 de la
méme largeur, 3 plus long que large, 4-5 égaux, plus longs que larges, plus courts que
3, 6-8 aussi longs que larges ou a peine plus longs que larges, plus courts que les
précédents, 9 aussi long que large, plus grand que 8 (8 et 9 a peine transverse chez
2), 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.
Elytres à ponctuation formée de grands points; les diamètres des points aussi grands
que les intervalles entre eux.

d. 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 83]

18 19

Fics 12 à 19

Bryaxis, articles 1 à 3 de l’antenne d: 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 (Aubé). 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.

2. 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 espece semble proche de B. femoratus (Aubé) par ses caracteres 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 parameres 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 légerement transverses, 9-10 nettement transverses), la forme
des métatibias chez le d (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 LOCALITES

Bryaxis sacrificus Kurbatov & Löbl, 1995

Matériel: 4 ¢, 2 2, 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’espece n’était connue que de la province chinoise de Shaanxi.

Bryaxis panda Kurbatov & Löbl, 1995

Matériel: 1 d, Chine, ouest de la province de Hubei, Shennongjia Nat. Res.,
2000-2200 m, débris végétaux, 7.VI.1995, leg. S. A. Kurbatov (CSKM).

Notes. Espece décrite et connue seulement de Sichuan.

Le d 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 2, Birmanie, Shan province, Namhsan, 1600 m, litière, 19.02.
1996, leg. S. A. Kurbatov (CSKM).

L’absence de d ne permet pas de donner une description basée sur des carac-
teres sürs. Toutefois, la découverte d’une espece de Bryaxis en Birmanie est impor-
tante. Le genre est inconnu de la region himalayenne, du Tibet, de Yunnan et de
Thailande.

LITTERATURE

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. & LOBL, 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

LößL, I. 1964. Neue ostasiatische Arten der Gattung Bryaxis Kugelann (Col., Pselaphidae).
Acta Societatis entomologicae cechosloveniae 61: 43-46.

LÔBL, I. 1977. Weitere Pselaphiden von der Koreanischen Volksdemokratischen Republik.
Bulletin de l’Académie Polonaise des Sciences. Serie des sciences biologiques Cl. V,
25: 235-241.

Losi, I. & KURBATOV, S. A. 1996. The Bryaxis of Taiwan (Coleoptera: Staphylinidae:’
Pselaphinae). Bulletion of the National Museum of Natural Science (Taichung) 8: 1-22.

Los, I., KURBATOV, S. A. & Nomura, S. 1998. On the Japanese species of Bryaxis (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.

i
sr FEMON

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, I-72021 Francavilla Fontana, (Brindisi), Italy.
** 19 Mount Crescent, Hereford, HRI INQ, 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 d). 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 $th 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. Sutural 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+Sth 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 = 1:74). 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 1:92 mm and height 1:10 mm.

Fics 1-5

Stetholiodes magnifica n. sp. 6: body outline 1, dorsal and 2, lateral, scale line = 1-0 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 2:15 mm, width 2:10 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: 6, Taiwan, Taichung Hsien, Hseuhshan, above Shan-Lin-Gien Hut,
3360 m, 10.v.1991, leg A. Smetana, in Geneva Natural History Museum collection.

Discussion: Stetholiodes magnifica sp. n. 1s 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 Löbl, 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)!

Sändor 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: Paracarinogalumna 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 110 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, forét),
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’Esperance River, riviere presque a sec, dans forét de Lauriers et Manguiers; (st.2)
50 m; 15.V1.1979; 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, pres de Thomaso, E de Morne Panache, forét claire, prélevement 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 Derniére Riviére,
récolte de feuilles dans forét dense, tres en pente, avec falaises et abris sous roche;
(st.4), 300 m; 17.V1.1979; 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, pres 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, forét 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, pres de Thomaso, E de Morne Panache, forét claire, prelevement sur souche
d’arbres pourris, tamisage, Berlese a Genève: (st.3.), 50 m; 16.VI.1979; leg. T. Jaccoud
et L. de Roguin (WB)*.

'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.V1.1979; leg. T. Jaccoud et L. de Roguin.

STL-79/11: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt
de pluie typique, très bien conservée (protégée), très humide pe ansa de sol; (st.6.),
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B)3.

STL-79/13: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forét
de pluie typique, très bien conservée (protégée), très umide 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, a 500 m de Piton St. Esprit, forét
de pluie typique, très bien conservée (protégée), très humide prelevement de sol; (st.6),
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B)?.

STL-79/17: 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.V1.1979; leg. T. Jaccoud et L. de Roguin.

STL-79/18: Saint Lucia (Micoud/Soufriére): Quiless Reserve, à 500 m de Piton St. Esprit, forêt
de pluie typique, très bien conservée (protégée), très humide RTE de sol; (st.6.),
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B)°.

STL-79/19: Saint Lucia (Micoud/Soufrière): Quiless Reserve, à 500 m de Piton St. Esprit, forêt
de pluie typique, très bien conservée (protégée), très ne prélèvement de sol; (st.6),
200-350 m; 19.VI.1979; leg. T. Jaccoud et L. de Roguin (B)°.

3 (B): extraction par appareils Berlese à Genève.
+ (WB): extraction par appareils Winkler-Moczarski sur place et par appareils Berlese à a
Genève.

ORIBATIDS FROM ST. LUCIA 84]

LIST OF IDENTIFIED SPECIES

Mesoplophoridae Ewing, 1917

Mesoplophora hauseri Mahunka, 1982
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) subtilis Niedbata, 1981
Localities: STL-79/12: 5 specimens; STL-79-18: 3 specimens.
Distribution: Well distributed in the Neotropical and Oriental Region
(see Niedbata 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.

842 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: I 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
Prorat ye Sle 79/93 specimens:
Distribution: Cuba (known from the type localities only); first record
for St. Lucia.
Microtegeus hauseri sp. n.
roicalliters -2SDE 79 STE-79/M:
Microtegeus lucianus sp. n.

roreanlaieers: STE-79/27S1E279/77SDE79/172S:11-791192

Eremaeozetidae Piffl, 1962

Eremaeozetes lineatus Mahunka, 1985
Poicalnye SE 79/821Especimen:
Distribution: St. Lucia (known from the type locality only); second
record.

ORIBATIDS FROM ST. LUCIA 843

Eremaeozetes roguini sp. n.
Boat EST 79/7

Microzetidae Grandjean, 1936

Berlesezetes auxiliaris Grandjean, 1936
Localities: STL-79/2: | 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.
rorcranlatays: STE-79/19;

Eremulidae Grandjean, 1965

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.
Poches SIE79/2;STE-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-79/13: 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.
Kogalıt y: STE79/19;
Gittella insularis sp. n.
Locality: STL-79/17.
Machuella ventrisetosa Hammer, 1961
Locality; STL-79/19: 2 specimens.
Distribution: Neotropics and Oriental Region; first record for
St. Lucia.
Moritzoppia subfallax sp. n.
OI ANNE RS ITIE 791107:
Multioppia insularis Mahunka, 1985
Locality: STL-79/19: 2 specimens.
Distribution: St. Lucia (known from the type locality only); second
record.
Oppiella nova (Oudemans, 1902)
[Evorcea litt es: SIE 79/522 specimens: SIE 79/10: Ssspecimenss) Siile=
79/13: 1 specimen.
Distribution: Cosmopolitan; second record for St. Lucia.
Oxyoppia antillensis sp. n.
ioc alte s = SIE 79/62 S1E-7913:
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, 1985
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; STE-79N22SIE79472

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.

Kocalities: STL-79/1; STE-79/19:

Oripodidae Jacot, 1925

Benoibates minimus Mahunka, 1985
Localities: STL-79/12: I 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
PO CAEN Sie 9/222 specimens:
Distribution: South- and Middle-America; first record for St. Lucia.
Peloribates antillensis (Mahunka, 1985)
Boealities.: STL-79/4: 1 specimen; STL-79/7: 1 specimen: SIE 7911:
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.
MocaliityeasTE-79/"

Galumnidae Jacot, 1925

Galumna flabellifera Hammer, 1958
Locality: STL-79/1: 15 specimens.
Distribution: Circumtropical; second record for St. Lucia.
Galumna hamifer Mahunka, 1985
Localities: STL-79/5: 3 specimens; STL-79/8: 9 specimens.
Distribution: Guadeloupe, Brazil; first record for St. Lucia.
Paracarinogalumna genavensium gen. n., sp. n.
rOrceay bit ers, SL Ol SMIE=7. 9/575 BE.7978:2S 7797125 eas Oil:
STE-79/17.
Pergalumna cucheae sp. n.
Po ait se SUR
Pilogalumna antillensis sp. n.
ocaliity. STE 70/8

848 SANDOR MAHUNKA

Fics 1-3

Mesoplophora (Parplophora) subtilis Niedbata, 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$, 1 paratype (1556-PO-96): HNHM’.

Measurements.-Length of body: 198-214 um, width of body: 126-134
um.

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 um, width of body: 105-139
um.

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

Fics 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

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 8-11

Material examined: Holotype: St. Lucia: STL-79/7; 1 paratype: from the same sample.
Holotype: MHNG, | paratype (1557-PO-96): HNHM.

Measurements. - Length of body: 346-367 um, width of body: 205-226
um.

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).

SANDOR MAHUNKA

852

Fics 8-11

Eremaeozetes roguini sp. n. — 8: body in dorsal aspect, 9: body in ventral
10: posteromedian part of the coxisternal region, 11: tarsus and tibia of leg. I.

aspect,

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.

Gnathoso ma: 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 - | - 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
(ad,, ad,) 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 @, and 9,
of leg I arising on a large process (Fig. 11), @, very small.

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 um, width of body: 166-186
um.

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

Fics 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 of podosoma: 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 um, width of body: 155-180
um.

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 of podosoma: 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

Fics 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: | - | - 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 ne
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 um, width of body: 155-171
um.

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, short,
setae h, much longer than setae p, and p3.

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 /c 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 um, width of body: 255-297
um.
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

Fics 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

ine

Fics 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 of podosoma: Pedotecta I narrow, setae /c 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, ap. 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 - 442 - 2042 - 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: Itis 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 um, width of body: 145-171
um.
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 à
pair of humeral apophyses conspicuous. Setae c, not shorter than the other nine pairs
of notogastral setae (Fig. 27). .
Lateral part of podosoma: Well granulated, with some stronger
laths (Fig. 29). Exobothridial setae hardly shorter than the interlamellar ones.

ORIBATIDS FROM ST. LUCIA

Fics 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.

861

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 /c 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,, 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: Itis 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: 193-211 um, width of body: 91-102
um.

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,.

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

Wear

ER >
AR N
55 \
N \
\

DA EOS
oa SUE

sauer

Fics 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 11 paratypes: MHNG, 6 paratypes: (1566-PO-96): HNHM.

Measurements. - Length of body: 291-376 um (female), 331-352 um
(male), width of body: 160-211 um (female), 175-181 um (male).

864 SANDOR MAHUNKA

Fics 34-37

Parakalumma piton sp. n. — 34: body in dorsal aspect (2), 35: body in dorsal aspect (d),
36: lamella and sensillus (©).
Parakalumma foveolata Balogh & Mahunka, 1969. — 37: lamella and sensillus (9).

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, Sa 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 - | - 2 - 3. Lyri-
fissures ad in adanal position. All setae in the ventral regions short and simple.

Legs: All legs monodactylous, with large claws. Seta ff” and € inserted
behind solenidia w, and w,. @, and @, arising on a projection. A well developed
blade-like formation on femora III and IV ventrally.

Remarks: The new species stands very near Parakalumma foveolata
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; | paratype: STL-79/1. Holotype and 3 paratypes: MHNG, 2 paratypes: (1564-PO-96):
HNHM.

Measurements. - Length of body: 560-691 um, width of body: 477-544
um.

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

Fics 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 of podosoma (Fig. 40): Tutorium well developed, with
a short cusp, rostral seta inserted close to it. Exobothridial setae conspicuously long
and thin. Pedotecta II-II 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 Unguizetes 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 um, width of body: 372 um.

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 /p 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

868 SANDOR MAHUNKA

Fics 43-44
Eupelops spongiosus sp. n. — 43: body in dorsal aspect, 44: body in ventral aspect.

adjacent setae /p and h,, 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 nominis: 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, | paratype: (1565-PO-96): HNHM.

ORIBATIDS FROM ST. LUCIA 869

Fics 45-46

Truncozetes rugosus sp. n. — 45: body in dorsal aspect, 46: body in ventral aspect.

Measurements. - Length of body: 233-244 um, width of body: 171-176
um.

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,
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,
Truncozetes 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 7. 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 - | - 2 - 3. Lyriffissures iad located far laterally from the
anal aperture. Porose area postanalis absent. All legs tridactylous, heterodactyly weak.
Solenidium w, and e stand behind w..

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 (Carinogalumna 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
Carinogalumna 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 Carinogalumna.

Derivatio nominis: The new genus is very near to Carinogalumna
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-79/11; 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 um, width of body: 589-642
um.

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, narrow, long.
Median pori absent, lyrifissures im located near to porose area A).

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 - 0-2-2.
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 - 341 - 442 - 2042 - 3
IV: 1-2-2-341-12-3

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 um, width of body: 460-552
um.

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

Fics 47-48

Paracarinogalumna genavensium gen. n., sp. n. — 47: body in dorsal aspect, 48: body in ventral
aspect.

ORIBATIDS FROM ST. LUCIA 873

Fics 49-51

Paracarinogalumna genavensium gen. n., sp. n. — 49: podosoma in lateral aspect, 50: lamellar
line, 51: 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: | - 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

Fics 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 clericata (Berlese, 1914), which was
designated by PEREZ-INIGO & BAGGIO (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 ın 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: 683-781 um, width of body: 526-622
um.

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, characteristically elon-
gated, A, irregular. Ten pairs of minute notogastral setae or their alveoli present, at
least setae c,, da and p, 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 - 0 - 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

Fics 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.

GRANDIJEAN, 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: 93-117.

PEREZ-INIGO, C. & Baggio, D. 1994. Oribatides édaphiques du Brésil (VII). Oribates de l’état
de Sao Paulo (Cinquième partie). Acarologia 35: 181-198.

SUBIAS, 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, Curacao and Aruba im
Jahre 1930. No. 10. Trimalaconothrus pilipes, eine neue Oribatide aus Westindien.
Zoologische Jahrbücher. Abteilung für Systematik 64: 447-452.

; AI BIER
es si wh mus uN Wien bi IA 015 HE ta) xa:
We atl, ti A if. 7 à I, | y | Bei hi 7 7

= sain nt ys

1°

Oy yew ca LEPRI CAIO
‘“eforo Nr T

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!
51, rue du Faubourg Saint-Martin, F-21121 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 d and 2 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 Departement 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 étre cités d’Asie et des îles du
Pacifique. L’espece 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 émérite de l’Université de Bourgogne, Equipe d’Ecologie et Dynamique
des Populations, F-21100 Dijon.

Manuscrit accepté le 01.07.1998

880 JEAN PAGES

L’énigmatique P. (G.) grassianus var. indica Silv. du Sikkim (SILVESTRI 1913)
est unique Parajapygidé signalé de la région indienne. D’ Australie, WOMERSLEY
(1934, 1945) a fait connaitre P. (P.) swani Wom. d Australie occidentale et P. (G.)
queenslandica Wom. du Queensland; enfin, des iles du Pacifique, on peut citer, outre
l’ubiquiste Parajapyx (P.) isabellae (Grassi) des Îles Hawaï, une © 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
figures justifient leur attrıbution 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 recemment 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) (PAGÉS 1954) dont la dj droite n’est plus
qu'un minuscule denticule, le reste de la marge étant normal?. 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 |’ Alabama ne montrerait plus que la d,, le reste
des marges étant totalement lisse (EWING 1941). P. paucidentis pourrait étre considéré
comme le stade ultime de la régression, les marges ne présentant plus qu’un minus-
cule denticule pres de l’apex des cerques ne correspondant a 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 Hal. d’Italie dont la marge interne du
cerque droit est complètement dépourvue d’armature. Il me semble inutile de nommer

2 La fig. 6 représentant cette anomalie a été attribuée par erreur à un variant IV d’In
Salah.

JAPYGOIDEA DU SUD-EST ASIATIQUE 881

de telles anomalies 4 moins que la méme 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 étre établie avec certitude actuellement.

Dans cette note, je répartis les 14 spécimens de Parajapygidés récoltés en 1987
a Java et Bali (Indonésie) et en 1988 a 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 caracteres ambigus et
trop peu nets qui ne me semblent pas permettre pour l’instant une determination
précise.

La découverte chez Parajapyx (P.) hauseri n. sp. d’un nouveau stade du déve-
loppement postembryonnaire des 6, 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 ESPECES

On trouvera la description des chétotaxies typiques, la definition des rapports
et des abréviations utilisés dans PAGES 1952a, 1952b et 1954. La nouvelle numé-
rotation des stades du développement postembryonnaire des d et des 9 est exposée
ci-apres.

Parajapyx (Parajapyx) hauseri n. sp. Figs 1-18

Matériel étudié: 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 Belait), forêt “Kerangas” (= “Tropical heath forest”), prélèvemenet de sol au pied de
Casuarina nobilis Whitmore (Casuarinaceae), 30 m; 20.X1.1988; leg. B. Hauser (B,); holo-
type: 5 4, de 2,14 mm; paratypes: 1 5 3, de 1,94 mm, 1 ¢ „, de 1,66 mm, 1 9 st. Î de 1,55
mm, 1 © de 2,02 mm, 1 © st.4 de 2,22 mm, 1 sexe? (urites 8 à 10 et les cerques absents).

TETE

Vertex: chétotaxie typique.

Pli oral: sans la soie 4.

Antennes: de 18 articles assez pileux, pas d’aires pileuses différenciées:
sensilles recourbées présentes a partir du Se article; 4 sensilles placoides 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) (B,), à Bandar
Seri Begawan (Brunei) (B,), Hong Kong (B;) ou à Genève (B,).

882 JEAN PAGES

Fics 1-6

Parajapyx (Parajapyx) hauseri n. sp., d holotype. - 1. Vertex, e = 105 um. - 2. Pronotum, e =
105 um. - 3. Mésonotum, e = 105 um. - 4. Métanotum, e = 105 um. - 5. Tergite 1, e = 105 um.
- 6. Tergites 7 à 10 et les cerques, e = 105 um.

JAPYGOIDEA DU SUD-EST ASIATIQUE 883

THORAX

Face tergale

Pronotum: les 5+5M typiques, M; 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, les plus longs, les M, les plus courts, les 5+5 s typiques assez
courtes, 1+1 soies supplémentaires entre sj et s, et 1+1 autres entre sy et ss.

Métanotum: 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
aigués, dépassant de peu l’apex de la griffe postérieure; celle-ci est a peine plus
longue que l’antérieure; unguiculus minuscule, aigu.

ABDOMEN

Tergite 1: Préscutum: 2+2 soies assez longues et environ 5+5 microsoies.
Scutum: les 5+5 M typiques, les M, les plus longs, 2 fois aussi développés que les M,
qui sont les plus courts de tous; les 6+6 s typiques assez courtes, sauf les s; 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, et/ou la présence d’une soie supplémentaire
entre M, et sg.

Tergites 2 à 7: Prescutum: 1+1+1 M typiques; le 3 holotype présente
2+2 M sur le préscutum du tergite 6. Scutum: les 8+8 M typiques longs, les M, et M;
les plus développés; les 7+7 s peuvent étre 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 9 st.1 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,, 55-57, de la s, gauche
et d’une soie supplémentaire près de la s, gauche; cette soie supplémentaire peut être
présente chez les individus plus âgés, ainsi que deux autres entre M, et sj ou M, et sg.

Tergite 8 : un peu moins de 1,25 fois aussi large que long (1/L = 1,25 chez
le 4 holotype, 1,22 chez la 9 st.1, 1,16 chez la 9 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 (sj nulles) chez les d et la 2 st.4, seules s3 et s; sont présentes chez
la ast

Tergite 9: 1/L= 2,24 en moyenne (2,25 chez l’holotype, 2,07 chez la 2
st.1, 2,48 chez la © st.4); les 3+3 M typiques longs; aucune s chez le & holotype;
chez les autres spécimens, seules les sj sont présentes si l’on admet que cette paire de
phaneres est venue se placer entre M, et M,.

884 JEAN PAGES

Fics 7-14. Parajapyx (Parajapyx) hauseri n. sp., 4 holotype. - 7. Urosternite 1, e = 105 um. -
8. Urosternite 3, e = 105 um. - 9. Hypopyge, e = 84 um. - 10. 9 st.1, tergites 7 à 10 et les
cerques, e = 105 um. - 11. @ st. 3, tergites 7 à 10 et les cerques, e = 105 um. - 12. idem,
acropyge, e = 51 um. - 13. idem, angle externe de l’organe subcoxal gauche et style, e = 42 um.
- 14. idem, hypopyge, e = 51 um.

JAPYGOIDEA DU SUD-EST ASIATIQUE 885

Fics 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. d holotype (= d 4,), € = 60 um. - 16. d 3.
e = 60 um. - 17. d ,,,e = 51 um. - 18. © st. 2, e = 46 um.

Tergite 10: L/1 = 1,61 en moyenne (v. ex. = 1,54-1,74), 1,56 chez le 4
holotype; 6+1+6 M (M, nuls) longs, le M, droit manque chez la 9 st.4; 444 s (s4
nulles) courtes, la sj droite est nulle chez la 9 st.1; le phanère situé entre M, et s,
peut étre interprété comme un M, qui est typiquement inséré au-dessus de la ligne
joignant M; et s,, ce qui le place entre M, et M-, ou bien comme une s3 typiquement
située presqu’exactement entre M; et s,; sur certains individus, se phanère peut être
considéré comme un M, a droite et comme une s3 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 8à 10: 63-33-100 en
moyenne; d holotype: 61-33-100; 3 ,,: 61-29-100; 2 st.1: 64-35-100; ® st.4: 64- 29-
100.

Acropy ge: triangulaire, simple, aussi long que large à la base; chez le d
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
a rapprocher de ce que SILVESTRI (1929) représente chez son P. (P.) dorianus ou moi-
méme chez P. (P.) genavensium (PAGES 1978).

Sternite 1: Préscutum: chétotaxie typique, la soie médiane peut étre
absente. Scutum: les 10+10 M typiques assez longs, My, M, et M, 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 a gauche de la ligne médiane: 1 entre M, et M,, | entre M, et b, 1 ou 2 entre M, et b.

886 JEAN PAGES

Organes subcoxaux latéraux: ils occupent chacun environ le
1/3 de la largeur interstylaire et sont peu saillants; de 14 a 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 holotype, 1,61 en moyenne chez les autres exemplaires.

La partie médiane postérieure du sternite porte les 1+1 minuscules soies
typiques.

Sternites 2 à 7 : 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 cöne secondaire aigu, Egal aux 2/5 de la
longueur du cône principal; aux st,_, la soie recourbée (s’) est subégale à la soie droite
(s); S,/st, =s’,/st, = 0,47; sj/st7 = 0,38; s7/st7 = 0,34; st, /st7 = 0,83.

Vésicules exsertiles: typiques du genre.

Hypopyge: Chezle d ,,, 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 Lg : j'ai défini dans ma note de 1975 sur Parajapyx
(P.) botosaneanui, 4 stades du développement postembryonnaire des d caractérisés
par le nombre de soies présentes sur la papille génitale: d | = 6 soies, d , = 12 soies,
33 = 14 soles, Sy = 16 soies. Or, parmi les 3 d reconnus ici, un a 12 soies, un autre
14 et le troisième en présente 10, ce qui le rend intermédiaire entre mes d , et d5. Je
crois qu’il faut donc maintenant definir 5 stades dans le développement des d et
modifier par conséquent la nomenclature précédente. Si l’on part toujours d’un 4, a6
soies (1+1 latérales, 1+1 submédianes antérieures et 1+1 submédianes postérieures),
le nouveau d , 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 d 3 c’est le dédoublement des 1+1
soies latérales qui donne bien 12 soies; quant aux 1+1 soies supplémentaires du d 4,
elles proviennent du dédoublement des 1+1 soies sublatérales antérieures, ce qui
corrobore ce que j’Ecrivais pour botosaneanui: “les nouvelles soies apparaissent dans
les angles latéraux de l’orifice génital, médialement par rapport aux soies latérales”;
enfin le d 5 voit cette fois le dédoublement des 1+1 soies sublatérales postérieures. En
resume, la nouvelle succession des stades postembryonnaires des d de Parajapyx
s’etablit ainsi:

din = 6 soies; d ;, = 10 soies; 43, = 12 soies; d 4, = 14 soies; ds, = 16 soies.
On a donc les équivalences suivantes entre la nouvelle nomenclature d,, et l’an-
cienne d ;:

Sin = 15 San nouveau, 53, = 59, Sun= 93: Ê5n = da

JAPYGOIDEA DU SUD-EST ASIATIQUE 887

Papille génitale 2 : typique de la famille. En 1952a, grace à
l’abondant matériel en ma possession, mais ne comprenant que des individus pourvus
de cerques indurés, j'ai défini | stade asexué st.I et 4 stades du développement post-
embryonnaire des @, st.Il à st.V caractérisés par le plus ou moins grand déve-
loppement du canal antérieur de la spermatheque. 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 troisieme possede des cerques indurés; il s’ensuit que logi-
quement, mon st.I doit étre denomme st.III par analogie avec celui des Japygides;
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 2 de 1,55 mm ne posséde qu’une seule et unique soie sur chacune des
valves supérieures, aucun autre phanere sur la papille et la spermatheque 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 2; je le nomme st.1. La deuxième raison est |’ extraordinaire développement de la
spermatheque des 2 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 |’ an-
cienne notation): st.1 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, /Ling = 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 d3 égale
en moyenne le 1/3 de leur longueur, le 1d,/1cq = 0,58 chez le d holotype alors qu'il
est en moyenne égal a 0,73 (v. ex. = 0,70-0,76) chez les paratypes. Un sinus peu
profond entre d, et d3 chez les paratypes, plus net chez l’holotype.

Marges internes: celles du d,, et des ? sont semblables; toutes les
dents sont aiguës, à sommet dirigé vers la base du cerque; d3 la plus forte, les autres
plus petites subégales; d, avec un très faible épaulement postérieur, obsolète chez le
Sn, seule la d3 montre un épaulement postérieur net; en faisant l’intervalle d,-d, égal
a 100 ceux entre les dents et entre d,-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 méme individu, mais les écarts sont minimes; les plus grands se rencontrent chez
le 3 ,,: 100-105-63-63-168 au cerque droit, 100-90-60-60-173 au gauche. Les marges
internes des 2 autres d sont bien différentes par la forme de la d, d’une part, et
d’autre part, par le déplacement des d; a d, les unes vers les autres et vers l’apex du
cerque: chez le d 3, lad, devient prépondérante, son sommet est saillant et arrondi, la
d, s’aplatit et ne forme plus qu’une légère élevure mousse, peut saillante sur le sinus,
les d, à d, 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 dj, il y a une déformation remarquable des deux dj 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, sont réduites 4 de minuscules tuber-
cules arrondis, les d, a d, sont assez semblables a celles des 2, 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 à ds.

Plaques d‘évaporation: les 9 et le d,, ne montrent qu’une seule
plaque d’évaporation assez grande située sur une ligne joignant M, à c et près de cette
dernière; le d 3, en possède 1 à gauche et 2, plus ou moins jointives, à droite, quant au
34, il en présente 3 à droite et 4 à gauche assez petites; chez ces 2 d les plaques
tendent à se déplacer d’une part vers la minuscule soie insérée au niveau de d, 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 (PAGÉS 1952a).

Chétotaxie: les 10 M typiques présents, M, 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 d 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 d, à ds. Le nombre d’articles antennaires, l’absence de M ou de s sur certains
tergites, en particulier le dixiéme et, sur les cerques, la présence d’une seule plaque
d’évaporation chez les © et les & 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-
seum d’histoire naturelle de Genève. Grace à ses nombreuses missions vouées avant
tout a la récolte de la faune du sol de régions peu prospectées ou écologiquement sen-
sibles, il a réuni, parmi d’autres, une trés 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”), prelevement de sol dans les angles formés par les contreforts de grands
arbres morts, 80 m; 24.X1.1988; leg. B. Hauser (B3); holotype: © st. 4 de 2,35 mm.

TETE
Vertex: les 11+11 soies typiques assez courtes, aucune soies supplé-
mentaires.

JAPYGOIDEA DU SUD-EST ASIATIQUE 889

Fics 19-26

Parajapyx (Grassjapyx) temburong n. sp., 2 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

Pronotum: 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, et M, assez courts, les autres longs; 5+5 s typiques assez
courtes; 2+2 soies supplémentaires dont 1+1 entre sj et s, et 1+1 entre sj et ss.

Métanotum: Préscutum: 2+2 M. Scutum: en tout point identique a celui
du mésonotum.

Face sternale

Chétotaxie: 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 aigués dépassant la griffe postérieure; celle-ci 1 fois 1/3 aussi longue
que l’antérieure; unguiculus normal.

ABDOMEN

Tergite 1: Préscutum: 2+2 M assez longs. Scutum: les 5+5 M typiques,
M, 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, M,, M; et M, longs, les autres assez longs; sj et s5_7 en paires
typiques, s, et s, présentes d’un seul côté de la ligne médiane, en général l’une d’elles
à droite et l’autre a gauche; 1+1 soies supplémentaires entre sg et 57.

Tergite 8: un peu moins de | fois 1/5 aussi large que long (1/L = 1,15);
6+6 M longs (M, nuls); 4+4 s typiques (s, nulles), s; 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 10: I fois 3/4 aussi long que large, à côtés parallèles; 6+6 M (M,
nuls), M, courts, les autres longs ou assez longs; 3+3 s (s3 et s, nulles), les remarques
faites chez l’espèce précédente a propos du choix à faire entre M, et s3 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.

Sternite 1: Préscutum: 4+1+4 soies assez longues. Scutum: les 10+10 M
typiques. M, et M,, plus longs que les autres qui sont assez longs; 1+2 soies typiques
assez courtes (c nulles); I+1 soies supplémentaires entre M4 et M,; 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, = 0,77; 5 soies sensorielles régulièrement
espacées, SS/st] = 0,46; SG/SS = 1,65.

JAPYGOIDEA DU SUD-EST ASIATIQUE 89]

La partie médiane postérieure du sternite porte les 1+1 minuscules soies
typiques.

Sternites 2 a 7: Préscutum: 5+1+5 soies typiques longues, les 1+1 les
plus externes les plus longues. Scutum: les 12+12 M typiques, M, et M, longs, les
autres assez longs ou assez courts; 4+3+4 s typiques assez courtes ou courtes; une
soie supplémentaire peut s’ observer entre Ms et My.

Styles: typiques du genre, le cöne 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,_3 est a
peine plus courte que la soie droite (s): s,/st,; = 0,38; s’/st, = 0,35; sJ/s7 = 1,3; stj/st7
= 0,90; s,/stz = 0,34.

Vésicules exsertiles: typiques du genre.

Hypopy ge: iloccupe les 3/4 de la largeur intercondylaire, saillant, a sinus
large et peu profond, orné de nombreuses dents aigués.

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 a la base; leur largeur au
niveau de la d, égale le 1/3 de la longueur d’un cerque et les 3/4 de sa largeur a la
base.

Marges internes: dents aigués a sommet dirigé vers la base du cerque;
d, et d; les plus fortes, les 3 autres subégales, petites; épaulements nuls; les intervalles
entre les d et entre la d, et l’apex du cerque sont assez différentes d’un cerque a
l’autre, a droite ils sont comme 100-120-78-53-240 et a gauche comme 100-83-67-61-
211; cela peut indiquer que les d s’engrenent bien les unes dans les autres quand les
cerques se referment; les fortes valeurs des intervalles d,-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, à M, et M, ac.

Chétotaxie: les 10 M typiques, M, 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 Pagés, 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: INDONESIE: 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: © st.6 de 2,22 mm. Sar-87/8: INDONESIE: Java: Bogor, Jardin Botanique,
prelevement de sol dans les angles formes par les contreforts de grands arbres pres du “Guest
House”, env. 250 m; 24.X1.1987; leg. B. Hauser (B,); paratype: © st.6. de 2, 20 mm.

TETE

Vertex et pli oral: chétotaxie typique.

Antennes: typiques de 18 articles peu pileux, sans aires pileuses
différenciées; 4 sensilles placoides en position typique sur l’article apical.

Pièces bucales: typiques du genre.

THORAX

Face tergale

Pronotum: 5+5 M typiques, M, et M, assez courts, les autres longs; 3+3 s
typiques courtes, sj à mi-distance de M, et M,, s, et s3 rapprochées l’une de l’autre,
seules les s3 sont à leur place normale.

Mésonotum: Préscutum: 1+1 M assez courts et 3+3 microsoies. Scutum:
les 5+5 M typiques, M, les plus longs de tous, M, les plus courts, les autres subégaux,
longs; les 5+5 s typiques assez courtes; 1+1 soies supplémentaires entre les s| et 55,
celle de droite nulle chez la £ de Java qui présente une soie supplémentaire à droite
entre M, et s5..

Métanotum: Préscutum: 2+2 M assez longs et 6+6 microsoies. Scutum:
les 5+5 M typiques, M, longs, M4 courts, les autres subégaux et assez longs; la 9 de
Bali possède les 5+5 s typiques et 3+3 soies supplémentaires entre les s, et s,, M, et
Sy, Sq et sg; chez la © 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. et la seconde paire de soies supplémentaires de la © de Bali et 1+1 entre ©
les s, 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
| supplémentaire entre les 2 postérieures; en outre, on observe une petie soie
supplémentaire dans la rangée antérieure de la plage médiane des mémes sternites.

Pattes: peu pileuses; 2+2 soies sternales aux tarses, les subapicales sont
aigués, atteignant au plus le milieu des griffes qui sont typiques; unguiculus petit,
aigu.

ABDOMEN

Tergite 1: Prescutum: 2+2 M assez longs. Scutum: les 5+5 M typiques

assez longs, les M, un peu plus développés que les autres, M, les plus courts; les 6+6

s typiques assez courtes, la s3 droite manque chez la 2 de Java; 1+1 soies supple-
mentaires entre M, et M, et une autre paire entre 54 et sg.

JAPYGOIDEA DU SUD-EST ASIATIQUE 893

j e
by > \ 2 gt =

Fics 27-37. Parajapyx (Grassjapyx) reniformis n. sp., 2 st. 6, holotype. - 27. Vertex, e = 126
um. - 28. Mésonotum, e = 126 um. - 29. Métanotum, e = 126 um. - 30. Tergite 1, e = 126 um. -
31. Tergites 7 à 10 et les cerques, e = 126 um. - 32. Marges internes des cerques, e = 63 um. -
33. Plaque d’évaporation du cerque gauche, e = 32 um. - 34. Urosternite 1, e = 105 um. -
35. Urosternite 3, e = 105 um. - 36. Canal de la spermathèque caracteristique du st. 6, e =
238 um. - 37. Hypopyge, e = 51 um.

894 JEAN PAGES

Tergites 2&7: Préscutum: les 1+1+1 M typiques. Scutum: les 8+8 M
typiques, M, assez courts, les autres longs ou assez longs; les 7+7 s typiques présentes,
assez courtes, sj et s, les plus longues, une des s, manque le plus souvent chez la 2 de
Java; 1+1 soies supplémentaires entre M, et Mg et une autre paire entre sg et 57.

Tergite 8: à peine plus large que long chez la $ de Java, chez celle de
Bali 1/L = 1,20; 6+6 M longs, My; les plus longs, M nuls; 3+3 s (s, et ss nulles), 53
longues, sj et sy courtes, sj nulles chez la ? de Java; 1+1 soies supplémentaires
courtes entre M, et M et 1+1 entre M, et M.

Tergite 9: entre 1,70 et 2 fois aussi large que long; les 3+3 M typiques
longs; seules les sj sont présentes et longues.

Tergite 10: en moyenne | fois 3/4 aussi long que large; 6+1+6 M (M,
nuls) longs; on constate la même ambiguïté entre M, et s, que chez P. (G.) hauseri
n. sp.; il y a au plus 4+4 s (sy nulles) ou 3+3 si s3 est nulle elle aussi.

Longueurs relatives des segments 8 à 10: 65-35-100 en
moyenne.

Acropyge: saillant, triangulaire, un peu plus long que large à la base.

Sternite 1: Préscutum: 5+1+5 phanères longs à assez courts. Scutum:
8+8 M, Mg et M;q nuls, M, assez courts, les autres longs, les Mg les plus développés;
1+2+1 s courtes; 1+1 soies supplémentaires entre M, et Mg et 1+1 autres entre Ms et
b; 6-7 + 6-7 soies en avant des organes subcoxaux latéraux.

Organes subcoxau x : 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/st, =
0,86, SS/st, = 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 © 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 s3.

Styles: typiques du genre; la soie recourbée des sf, ; est égale à la soie
droite. sj/st] = 0,40; s’,/st] = 0,40; s/st7 = 0,32; s7/st = 0,40; st,/stz = 0,80.

Vésicules exsertiles: typiques du genre.

Hypopyge: Iloccupe 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 ® : Typique du genre; chez la 2 de Bali, le canal de
la spermathèque est extraordinairement long formant un peloton läche 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 2 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 (2 de Bali) à 2 fois 1/4 (9 de

JAPYGOIDEA DU SUD-EST ASIATIQUE 895

Java) aussi longs que larges a 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 a la base.

Marges internes: les dents sont aigués a sommet dirigé vers la base:
d, et d; 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,-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
réniforme lisse; au milieu de son bord concave s’ouvre un pore glandulaire qui est le
débouché d’un canal a 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) a la plaque d’évaporation du cerque droit.

Chétotaxie: les 10+10 M typiques, M,, M, et M, 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;, M; et h comme un M, uniquement parce qu'il est
inséré en dessous de la ligne joignant M, et i et sur celle joignant f et i, la soie e se
plaçant typiquement au dessus de la ligne M,-i et plus près de M; que de Mg.

AFFINITES

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 Parajapygides.

DERIVATIO NOMINIS

Allusion à la forme très particulière de l’unique plaque d’évaporation des
cerques.

Parajapyx (Grassjapyx) sabahnus Pagés Figs 38-44

Matériel étudié: Bru-88/70: SINGAPOUR: Sentosa Island (partie orientale), forét
pres de la “Earth Satellite Station”, prélevement de sol dans les angles formés par les
contreforts d’un grand arbre, 70 m; 6.XII.1988; leg. B. Hauser (B,); 3 ? st.3 de 2,17 mm,
2,34 mm et 2,54 mm.

Ces exemplaires correspondent parfaitement a la description de cette espéce du
Sabah (PAGEs 1987). La chétotaxie tergale, les rapports 1/L et L/1 des segments
abdominaux et des cerques entrent dans les limites de variations de l’espece; 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 d, à ds 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

Fics 38-44. Parajapyx (Grassjapyx) sabahnus Pages, 2 st. 3 de 2,54 mm. - 38. Tergite 1, e =
112 um. - 39. Tergites 7 à 10 et les cerques, e = 112 um. - 40. Marges internes des cerques, e =
49 um. - 41. Urosternite 1, e = 112 um. - 42. Angle lateral 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
probleme reste entier.

Plaques d’évaporation: tres petites, mais plus nettes que chez les
exemplaires du Sabah; au nombre de 4-5 par cerque, elles sont inscrites dans un
trapeze dont les sommets sont les M, M;, Ms et la soie c comme au Sabah.

Chétotaxie: caractérisée par l’absence des soies d et e et la brièveté des
Mg.

JAPYGOIDEA DU SUD-EST ASIATIQUE 897

Parajapyx (subgen. ?) sp.? Figs 45-47

Matériel étudié: Bru-88/32: BRUNEI (Belait District): “Labi Hills Forest
Reserve”, “Teraja”, a 42 km au sud de Sungai Liang (= 12 km au Sud de Labi), environs de
“Rumah Panjang” (= Longhouse du Kampong Teraja), forét primaire (“Mixed dipterocarp
forest”), prelevement de sol dans les angles formés par les contreforts d’un tres grand arbre,
40 m; 22.X1.1988; leg. B. Hauser (BJ); 1 2 st.3 de 1,80 mm.

Fics 45-47

Parajapyx (subgen. ?) sp.?, 9 st. 3. - 45. Tergites 7 a 10 et les cerques, e = 79 um. - 46. Marges
internes des cerques, e = 45 um. - 47. Hypopyge, e = 105 um.

898 JEAN PAGES

Cet exemplaire possede des cerques dont les caractéristiques ambigués me font
hésiter a le classer spécifiquement.

Les cerques sont nettement plus trapus que chez les espéces précédentes et se
rapprocheraient de ceux de P. (G.) sabahnus, mais la forme de la d, et la présence
d’une seule plaque d’évaporation située sur une ligne joignant M4 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 d;, même aussi peu
marqué que chez la © 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’évaporation situées elles aussi, il est
vrai, sur une ligne joignant M, et c, enfin les acropyges sont identiques, présentant
une dent sur le côté gauche. Je ne crois ni utile, nı souhaitable de créer un taxon
nouveau en l’absence d’autres exemplaires, en particulier de 4 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

CHou, I. 1966. Studies on Japygidae IV (Insecta: Diplura). Acta Zootaxonomica Sinica 3:
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.

NAKAMURA, O. 1994. Diplura (Insecta) from the Northern Mariana Islands, Micronesia. Natural
History Research, Special Issue 1: 219.

PAGÉS, J. 1952a. Parajapyginae (Insecta, Entotrophi, Japygidae) de l’Angola. Publicagöes
Culturais da Companhia de Diamantes de Angola 13: 53-96.

PAGES, J. 1952b. Contribution à l’étude des Japygidae (Insecta, Entotrophi) d’Algerie et de
Tunisie. Bulletin de la Société Zoologique de France 77: 125-148.

Paces, J. 1954. Parajapyginés (Diploures Japygidés) du Maroc et du Sahara. Bulletin de la
Société des Sciences Naturelles du Maroc 33: 129-144.

PAGES, J. 1975. Un Parajapygide inédit des plages de la cöte caraibe de Cuba récolté par Mr L.
Botosaneanu au cours de la seconde expédition biospéléologique cubano-roumaine a
Cuba en 1973. International Journal of Speleology 6: 339-352.

PAGES, J. 1978. Dicellurata Genavensia IV. Especes inédites de Japygoidea du Guatemala.
Revue suisse de Zoologie 84: 807-814.

PAGES, J. 1987. Dicellurata Genavensia XIV. Japygoidea du Sud-Est asiatique No 5. Revue
suisse de Zoologie 94: 41-47.

SILVESTRI, F. 1913. On some Thysanura in the Indian Museum. Record of the Indian Museum
9: 51-62.

JAPYGOIDEA DU SUD-EST ASIATIQUE 899

SILVESTRI, F. 1928. Japygidae (Thysanura) dell’Estremo Oriente. Bolletino del Laboratorio di
Zoologia Generale e Agraria della R. Scuola Superiore d’Agricoltura, Portici 22:
49-80. 3

SILVESTRI, F. 1929. Nuevos Parajapiginos de Africa (Thys. Japygidae). Memorias de la Real
Sociedad Espanola de Historia Natural 15: 221-235.

SILVESTRI, F. 1930. Contribuzione alla conoscenza degli Japygidae (Thysanura) della regione
Australiana. Bolletino del Laboratorio di Zoologia Generale e Agraria della R. Scuola
Superiore d’Agricoltura, Portici 23: 210-226.

SILVESTRI, F. 1948a. Intorno ad alcune anomalie di Japygidae (Insecta, Diplura). Bollettino del
Laboratorio di Entomologia Agraria di Portici 8: 209-213.

SILVESTRI, F. 1948b. Specie di Japygidae (Insecta Diplura) finora raccolti nel Messico. Bol-
lettino del Laboratorio di Entomologia Agraria di Portici 8: 297-320.

SMITH, L. M. 1961. Japygidae of North America, 8. Postembryonic development of Paraja-
pyginae and Evalljapyginae (Insecta, Diplura). Annals of the Entomological Society of
America 54: 437-441.

WOMERSLEY, H. 1934. On the Australian species of Japygidae. Transactions of the Royal
Society of South Australia 58: 37-47.

WOMERSLEY, H. 1945. New species of Diplura (Insecta, Apterygota) from Australia and New
Guinea. Transactions of the Royal Society of South Australia 69: 223-228.

XIE, R. & YANG, Y. 1990. A new species of Parajapyx from Xishuangbanna, China (Diplura:
Japygidae). Contributions from Shanghai Institute of Entomology 9 (1989-1990):
189-192.

XIE, R., YANG, Y. & YIN, W. 1990. A new species of Parajapyx from the Tianmu Mountain,
China (Diplura: Japygidae). Contributions from Shanghai Institute of Entomology 8
(1988): 229-233.

ZIMMERMAN, E. C. 1948. Order Diplura Borner, 1904. Insects of Hawaii 2: 38-42.

ZWALUWENBURG, R. H. VAN, 1934. Parajapyx isabellae, Proceedings of the Hawaiian Ento-
mological Society 8: 382.

fr Zi
u ni str sah
slot. 7,0% j ar: Muah ae x
1 nye. 4 u sli gue f , | r a

why untl

KA
w

Le a
ER

= an | Re ey ve if
ae de;

REVUE SUISSE DE ZOOLOGIE 105 (4): 901-910; decembre 1998

Redescription of Diplomystes mesembrinus
(Siluriformes, Diplomystidae)

Maria de las Mercedes AZPELICUETA! & Atila Esteban GOSZTONYI?

l Depto. Cientifico Zoologfa de Vertebrados, Museo de La Plata, 1900 La Plata,
Argentina

2 Centro Nacional Patagénico, 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 1s
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 (GOSZTONY! 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 Patagönico (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., | 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. Taylor. MLP uncat., 20 ex.,
63.0-135.3 mm (2 ex. C&S), rio Chubut en canadön 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
Cöndor (43° 19° 1°°-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 l+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 Sth parapophysis. A well-developed rib joined to Sth. 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
(Eis. 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

Fic. |

Diplomystes mesembrinus, rio Chubut near cerro Condor, 150.5 mm SL.

Fic. 2

Young specimen of Diplomystes mesembrinus, rio Chubut near cerro Céndor, 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).

LS ore -

——

SE
/

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

Fics 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: | 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 DIPLOMYSTES MESEMBRINUS 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 radials in the pectoral fin. The number of pectoral
distal radials 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 |

Measurements of 35 specimens of Diplomystes mesembrinus in percentage of indicated lengths.
SL = 33.8-215.0 mm.

x min max

Percentage of standard length

Predorsal-fin length 37.8 3501 39m
Preadipose fin-length 64.2 57.8 68.2
Preventral-fin length 48.6 44.5 51.0
Preanal-fin length 66.7 599 69.0
Dorsal-fin base IB 123 16.4
Adipose-fin base 25.9 21.4 31,9
Anal-fin base 14.0 11.0 175
Pelvic-fin length 16.6 13.6 20.0
Pectoral-spine length 18.4 18.2 215
Dorsal-spine length 18.0 Mes 2167
Greatest body depth 20.0 14.4 2575
Greatest head depth SZ 13.0 20.0
Head length 26.6 24.2 29.0
Head width 18.4 16.2 222
Mouth width TESTI 6.0 Well
Interorbital length 8.1 6.8 8.1
Preorbital length eZ 8.3 13.1
Percentage of head length

Head width 69.1 61.6 86.7
Mouth width 20.0 22.6 SO)
Greatest head depth 592 50.0 74.5
Interorbital length 30.5 DS 35.8
Preorbital length 42.1 338 49.2
Orbital length 16.4 ILS 29.8
Maxillary length 2359 17/87, 28.4
Premaxillary length 22) 15.6 23

Percentage of mouth width
Premaxillary tooth plate 82.7 65.8 999

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. Neotröpica 39:
101-102.

AZPELICUETA, M. DE LAS M. 1994a. Los diplomistidos 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). /chthyological Exploration of Freshwaters 5: 223-240.

CLARAZ, J. 1988. Diario de viaje de exploraciön 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), Seccion B
46: 41-50.

RINGUELET, R. A. 1982. Una nueva subespecie de bagre patagönico Diplomystes viedmensis Mac
Donagh, 1931 en el rio 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: 107-119.

REVUE SUISSE DE ZOOLOGIE, 105 (4): 911-982; décembre 1998

Aleocharinae della Cina: Parte IV
(Coleoptera, Staphylinidae)

Roberto PACE
Via Vittorio Veneto 13; I-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 Mimoxypoda). 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, c), 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 d, 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 estremita 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 é 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 1 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, 29.1V.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 d, China, Zhejiang, Hangzhou, 27.1V.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

Fico. 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 € 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 dé, 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 d, 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

0! mm

DISS \
DEN

Ol mm

Fico. 7-14

Edeago in visione laterale e ventrale, sesto urotergo libero del maschio, spermateca e habitus.
7-10: Atheta (Microdota) tricoloroides sp. n.; 11-14: 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 copulatori 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 d, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 25.VII.1994,
A. Smetana leg. (MHNG).

Paratypi: 2 d, 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 d, China, Gansu, Xinlong Shan, ca. 70 Km S Lanzhou, 2225-2380 m,
7.VII.1994, A. Smetana leg. (MHNG).

Paratypus; 1 d, 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 d, China, Gansu, 120 Km S Lanzhou, Guanghe Xian Mai Jia, 2300 m,
8.VII.1994, A. Smetana leg. (MHNG).

ALEOCHARINAE DELLA CINA

01 mm

oimm

FIGG. 15-21

Habitus, edeago in visione laterale e ventrale e spermateca. 15-18: Arheta (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
l’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

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 trasverse, 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

oimm

N ey,
1”

Fıcc. 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 4, 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, 9.11.1993, de Rougemont leg. (MHNG).

Paratypi: 1 & (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. placita 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 placita). 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
placita), priva di introflessione apicale del bulbo distale (presente invece nel bulbo
distale della spermateca di placita) e per la parte prossimale della stessa spermateca,
breve e non lungamente protratta come in placita.

ALEOCHARINAE DELLA CINA 92]

05 m

Ficc. 32-40

Habitus, edeago in visione laterale e ventrale e spermateca. 32-33: Arheta (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 d, China, Zhejiang, Hangzhou, 27.1V.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
figg. 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 d, China, Zhejiang, Trianmushan, 29.1V.1993, de Rougemont leg. (MHNG).

Paratypi: 6 d e 2 2, 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 ¢, 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

FIGG. 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 d, China, Yunnan, Xishuangbanna, Chayanhe F.P., 24.1.1993, de
Rougemont leg. (MHNG).

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 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 sclerificata 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 4, 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

imm

Fic. 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 € 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 d, China, Yunnan, Ruili, ca. 700 m, 3.11.1993, de Rougemont leg. (MHNG).

Paratypus: 1 ® 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 2, China, Yunnan, Kunming, 1.11.1993, de Rougemont leg. (MHNG).
Paratypi: 1 d, stessa provenienza; 1 d, China, Yunnan, Dali, 9.11.1993, de Rougemont
leg.

ALEOCHARINAE DELLA CINA 927

FIGG. 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.

ROBERTO PACE

928

01mm

E
E
Ss

FIGG. 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 d, 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 4, China, Sichuan, Gongga Shan, above camp 2, 2800 m, 26.VII.1994,
A. Smetana leg. (MHNG).

Paratypi: 10 d, 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

Fico. 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 & larghissimo.

Atheta (Bessobia) pergranulosa sp. n. Figg. 83-85

Holotypus dé, 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
dell’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 d, China, Gansu, Yonghai ca. 20 Km SW Yuzhong, 2700-2800 m,
9.VIIL.1994, A. Smetana leg. (MHNG).

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

RES

Be.

TIS

FiGG. 79-86

Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del maschio. 79-82: Atheta
(Bessobia) peranomala sp. n.; 83-85: Atheta (Bessobia) pergranulosa 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 ¢, China, Gansu, 120 Km S Lanzhou, Guanghe Xian Ma Jia, 2300 m,
8.VII.1994, A. Smetana leg. (MHNG).

Paratypus: | d, 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 2, 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 d, Gansu, Dalijia Shan, 60 Km W Linxia, 3475 m, 11.VII.1994, A. Smetana
leg. (MHNG).

ROBERTO PACE

934

Olmm

0) mm

FIGG. 87-92
Edeago in visione laterale e ventrale, spermateca e habitus. 87-89: Arheta (Oreostiba) yonghai-

cola sp. n.; 90-92: Atheta (Oreostiba) semitibialis sp. n.

ALEOCHARINAE DELLA CINA 935

Olmm

Fico. 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 d, China, Gansu, pass btw Hezuo-Amgog, 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 ifurupensis), 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. I] 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 4, 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

Fısc. 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 € 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 2, China, Sichuan, Gongga Shan, above camp 3, 3050 m, 22.VII.1994,
A. Smetana leg. (MHNG).

Paratypi: 1 9, stessa provenienza; 1 92, 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. angusticollis 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 d, Russia, Sakhalin, Aniva distr. 5 Km W Petropaulovskiy, tributary of
Lyutoga river, 20-21.VII.1993, Putz & Wrase leg. (CASS).

Paratypus: 1 ®, 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. 115, edeago figg. 116-117.

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

PTT; È
in Prref
2) r
DL ER il
FE 4 ]
44

108

Olmm

Ficc. 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

Fısc. 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. 118-121

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, 19-31.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.:
1del 9, 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 deil’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. 119-120, 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, e a P. 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 d, Hong Kong, XII.1995-1.1996, de Rougemont leg. (MHNG).

Paratypi: 18 es., stessa provenienza; 2 d, Hong Kong, Kadoorie Agricultural Research
Centre, 19-31.V.1996, de Rougemont leg.; 1 d, stessa provenienza, ma flight interception trap,
VI.1996, de Rougemont; 1 ®, stessa provenienza, ma VIII.1996, de Rougemont leg.; 1 à,
Hong Kong, Kadoorie Farm, V.1996, de Rougemont leg.; 6 es., Hong Kong, Tai Po, II.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) lii sp. n. Figg. 126-127
Holotypus ©, China, Hebei Prov., Yongniang, 6.X.1995, Shugiang Li leg. (MHNG).
Paratypi: 2 9, 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

01mm

Fico. 118-125

Habitus, edeago in visione laterale e ventrale e spermateca. 118-121: 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. Shugiang Li
di Stuttgard (Germania).

Pelioptera (Geostibida) kowloonensis sp. n. Figg. 128-132

Holotypus ©, Hong Kong, XII.1995-I.1996, de Rougemont leg. (MHNG).
Paratypi: 1 ©, 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 2, Hong Kong, Kadoorie Agricultural Research Centre, flight interception
trap, 19-31.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 d, 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

een

Fico. 126-134

Habitus, spermateca, edeago in visione laterale e ventrale e sesto urotergo libero del maschio.
126-127: Pelioptera (s. str.) lit 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 1 due antennomeri basali
giallo-rossicci; zampe gialle. La reticolazione del capo e del pronoto é 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, 3.11.1993, 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, nè 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 d, 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.VIIL.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 di fessa. 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

Ol mm

140

FIGG. 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 2, 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 d, China, Zhejiang, Tianmushan, 23.1V.1993, de Rougemont leg. (MHNG).

Paratypi: 49 es., stessa provenienza; 1 d e 2 2, 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 ¢, China, Zhejiang Prov., Lin’an County, 1000 m, W Tianmu
Shan N.R., 18.V.1996, J. Cooter leg.; 1 2, China, Yunnan, Ruili, ca. 700 m, 3.11.1993, de
Rougemont leg.; 1 d e 1 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 di fessa 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 dé, China, Beijing, Panshan, 8.V.1993, 8.V.1993, de Rougemont leg.
(MHNG).
Paratypus: 1 9, stessa provenienza.

ROBERTO PACE

948

144

Ol mm

FIGG. 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

Ficc. 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 d, 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 4, Hong Kong, Tai Po, V.1996, flight interception trap, de Rougemont leg.
(MHNG).
Paratypus: 1 d, stessa provenienza.

ALEOCHARINAE DELLA CINA 95]

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. . .
FERRO ee eo dae Pia Mimoxypoda s. str.
(Typus subgeneris: Mimoxypoda (s. str.) rufa 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 2, 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
superficie 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

THESEN
URTO
a ı f
SES VU UL.

160

156

00
4 Imm

166

FIGG. 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 ¢, China, Yunnan, Dali, 9.11.1993, 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 & 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 & 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

Olmm

FIVE
tin

OE
fener Ase î BS

ESME SÄCER
A TARA: 9

ES 41 1 1 ite

Fısc. 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

178

10727,

Ficc. 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 d, 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, 15.1X.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 d, Hong Kong, Kadoorie Farm, flight interception trap, 15.1X.1996, de
Rougemont leg. (MHNG).

Paratypus: 1 ®, 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

<a
NN
ANNAN

Olmm

imm

185

Figc. 183-188

Labio con palpo labiale, maxilla con palpo mascellare, spermateca, edeago in visione laterale e
kongensis sp. n.

ventrale e habitus. 183-184: Mesomegaskela adesi gen. n., sp. n.; 185-188: Doryloxenus hong-

958 ROBERTO PACE

Fico. 189-194

Habitus, spermateca, edeago in visione laterale e ventrale e nono segmento addominale. 189-
190: Doryloxenus rougemonti sp. n.; 191-194: Odontoxenus rougemonti sp. n.

ALEOCHARINAE DELLA CINA 959

COMPARAZIONI. La nuova specie é 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 € dedicata al suo raccoglitore studioso di Staphy-
linidae Guillaume de Rougemont di Londra.

Odontoxenus hongkongensis sp. n. Figg. 195-196

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 O. rougemonti sp. n. sopra
descritta.

Cephaplakoxena gen. n. Figg. 197-208

DIAGNOSsI. 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 1 posteriori di 5); piastra apicale dei parameri
minuscola (fig. 202).

960 ROBERTO PACE

0,05 mm

Fısc. 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. I] genere grammaticale € femminile.

Cephaplakoxena rougemonti sp. n. Figg. 197-208

Holotypus ¢, 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 estremita
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 é 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 striatura 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. 211-214

Holotypus d, 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

Fıcc. 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.

963

ALEOCHARINAE DELLA CINA

Olmm

ESA
Ta
PT

FIGG. 209-213

Habitus, spermateca ed edeago in visione laterale e ventrale. 209-210: Chaetosogonocephus
chinensis sp. n.; 211-213: Tetrabothrus rougemonti sp. n.

964 ROBERTO PACE

COMPARAZIONI. La nuova specie € simile a T. indicus Cameron, 1939, dell’ India,
ma ha antenne chiaramente fusiformi (non fusiformi in indicus), elitre unicolori
(rossicce con meta 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 7. 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 ¢, China, Yunnan, Ruili, ca. 700 m, 3.11.1993, 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 d, China, Zhejiang, Tianmushan, 29.1V.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 4 ; Hong Kong, N.T., Kadoorie Agricultural Research Centre, Malaise trap,
VIIL.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 1 tre antennomeri basali giallo-rossicci;
zampe giallo-rossicce. La superficie del corpo non é 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 é fine. Gli
uroterghi liberi 2° e 3° hanno un’ampia bozza basale mediana, il 4° e il 5° hanno una
concavita 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. assamensis (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 & 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 é stata raccolta la nuova specie.

Drusilla gibberella sp. n. Figg. 225-228

Holotypus 9, Hong Kong, Kadoorie Agricultural Research Centre, flight interception
trap, 19-31.V.1996, de Rougemont leg. (MHNG).

Paratypus: 1 d, 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

ta
>
=,

Imm
POSS

224

F1GG. 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 2, 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), nè parte prossimale della spermateca così ampiamente avvolta a matassa.

Zyras (s. str.) fratrumkadooriorum sp. n. Figg. 231-234

Holotypus d, Hong Kong, Kadoorie Farm, Malaise trap, VI.1991, G. Ades leg.
(MHNG).

Paratypi: 1 d e 1 2, 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

Imm

228

0) mm

Fısc. 225-229

Habitus, edeago in visione laterale e ventrale e spermateca. 225-228: Drusilla gibberella sp. n.;
229: Diplopleurus cooteri sp. n.

ROBERTO PACE

970

Fıcc. 230-234
Spermateca, habitus ed edeago in visione laterale e ventrale. 230: Diplopleurus cooteri sp. n.;

231-234: Zyras (s. str.) fratrumkadooriorum sp. n.

ALEOCHARINAE DELLA CINA 971

Zyras (s. str.) notaticornis sp. n. Figg. 235-238

Holotypus ¢, Hong Kong, Kadoorie Agricultural Research Centre, flight interception
trap, 19-31.V.1996, de Rougemont leg. (MHNG).

Paratypi: 4 es., stessa provenienza, ma III.1996, VI.1996, de Rougemont leg.: 1 d,
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 1 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 d, China, Shaanxi, Nanwutai, 17.1X.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 1 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 d, 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

ROBERTO PACE

972

238

237

241

FIGG. 235-

Habitus, spermateca ed edeago in visione laterale e ventrale. 235

cornis Sp. n.; 239-241: Zyras (s. str.) shaanxiensis sp. n.

(s. str.) notati-

-238: Zyras

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 d, Hong Kong, N.T., Kadoorie Agricultural Research Centre, Malaise trap,
VIII.1991, G. Ades leg. (MHNG).

Paratypi: 1 d e 1 ©, stessa provenienza, ma VI.1991, G. Ades leg.; 1 ©, Hong Kong,
Shing Mun, 8.VII.1996, G. Reels leg.; 1 2, 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 d, 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, 1.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

FIGG. 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

F1GG. 249-256

Habitus, edeago in visione laterale e ventrale e spermateca. 249-252: Zyras (Diaulaconia)
orientalis Bernhauer; 253-256: Zyras (Pella) 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 4, China, Beijing, Xiaolongmen, 1100-1500 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 ®, 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 1 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

O,lmm

3
TR RES AL

0
NI
OOD
Yon
NRA

Figc. 257-263

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del maschio.
257-261: Zyras (Pella) micropterus sp. n.; 262-263: Zyras (Pella) beijingorum sp. n.

978 ROBERTO PACE

COMPARAZIONI. La nuova specie é 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 d, 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 reticolazione sulle elitre e sugli
uroterghi. I tubercoletti della superficie del capo sono indistinti, quelli delle elitre sono
fini. La metà posteriore del quinto urotergo libero è coperta da robusta punteggiatura e
il sesto urotergo presenta una punteggiatura ancor più robusta. Edeago figg. 265-266,
spermateca fig. 267.

COMPARAZIONI. La nuova specie è distinta da Z. almorensis Cameron, 1939,
dell India, per gli occhi più lunghi delle tempie (occhi più corti delle tempie in almo-
rensis), per gli antennomeri 4° a 10° non trasversi (trasversi in almorensis) e per
l’ultimo antennomero lungo quanto i tre antennomeri precedenti compresi insieme
(ultimo antennomero lungo quanto i due antennomeri precedenti insieme in almo-
rensis).

ETIMOLOGIA. La nuova specie è dedicata a Garry Ades, zoologo che ha raccolto
Aleocharinae in Cina.

Pedinopleurus hongkongicola sp. n. Figg. 268-271

Holotypus 6, Hong Kong, XII.1995-I.1996, de Rougemont leg. (MHNG).

DESCRIZIONE. Lunghezza 3,8 mm. Corpo lucido e rossiccio con pronoto giallo-
rossiccio ed elitre giallo-brune con margine suturale largamente oscurato di bruniccio;
antenne rossicce con i due antennomeri basali e la base del terzo giallo-rossicci; zampe
rossicce. La reticolazione della superficie del corpo è distinta, quella del pronoto e
dell’ addome è a maglie trasverse. L’avancorpo è coperto di tubercoletti salienti, essi sul
pronoto sono radi in avanti e lateralmente e discretamente fitti dal disco al margine
posteriore. Il capo presenta il disco concavo e due pori discali salienti. La stria suturale
delle elitre è poco distinta. Edeago figg. 269-270, spermateca fig. 271.

ORIBATES D’UNE TOURBIERE JURASSIENNE 979

Fico. 264-271

Habitus, edeago in visione laterale e ventrale e spermateca. 264-267: Zyras (Zyrastilbus) adesi
sp. n.; 268-271: Pedinopleurus hongkongensis sp. n.

980 ROBERTO PACE

COMPARAZIONI. La nuova specie è distinta da P. indicus Cameron, 1939,
dell’ India, per la taglia corporea minore, per gli occhi ridotti e per la presenza di un
tubercolo mediano lungo quanto largo al margine posteriore del quinto urotergo libero
del maschio (tubercolo trasverso in indicus).

ADDENDA

All’elenco delle specie note e nuove per la Cina dato nella parte I della
presenta serie di lavori sulle Aleocharinae della Cina vanno aggiunte le seguenti
specie:

Medeterusa minima Pace Figg. 170-175

Medeterusa minima Pace, 1987c: 427
13 es., Hong Kong, Tai Po, IV.V.1996, de Rougemont leg.; 1 2, Hong Kong, N.T., sifted litter,
IX.1996, de Rougemont leg.

Specie finora nota solo del Nepal.

Zyras (Diaulaconia) orientalis Bernhauer Figg. 249-252

Zyras (Diaulaconia) orientalis Bernhauer, 1929: 3
1 d, Hong Kong, N.T. Shek Kong, 19.XI.1990, G. Ades leg.; 2 d e 1 2, Hong Kong, N.T.
Kap Lung, 26.IX.1996, G.T. Reels leg., at light.

Hong Kong é la localita tipica di questa specie che é finora nota solo di questa
localita.

RINGRAZIAMENTI

Rivolgo i miei piu sentiti ringraziamenti a quanti mi hanno affidato in studio le
Aleocharinae della Cina oggetto del presente lavoro e frutto di recenti raccolte: il
Dr. Ales Smetana di Ottawa, ı colleghi Guillaume de Rougemont di Londra, Jonathan
Cooter di Hereford (Gran Bretagna), Garry Ades, Graham Reels (Hong Kong), il
Dr. Jeng-Tze Yang della “National Chung Hsing university” di Taiwan, Volker Assing
di Hannover e il Dr. Shugiang Li di Stuttgard. Per il prestito di tipi e di materiale di
confronto, ringrazio molto il Dr. A.F. Newton del “Field Museum of Natural History”
di Chicago, il Dr. P.M. Hammond del “Natural History Museum” di Londra e il
Dr. L. Zerche del D.E.I. di Eberswalde.

BIBLIOGRAFIA

BERNHAUER, M. 1907. Zur Staphylinidenfauna von Japan. Verhandlungen der Zoologisch-
Botanischen Gesellschaft in Wien 57: 371-414.

BERNHAUER, M. 1915a. Beitrag zur Staphylinidenfauna von Neuguinea. Deutsche Entomo-
logische Zeitschrift 1915: 179-202.

BERNHAUER, M. 1915b. Zur Staphyliniden-Fauna des tropischen Afrika (7. Beitrag). Annales
Historico-Naturales Musei Nationalis Hungarici 13: 95-189.

BERNHAUER, M. 1929. Neue Kurzfliigler aus China. Entomologisches Nachrichtenblatt,
Troppau 3: 2-4.

ALEOCHARINAE DELLA CINA 981

BERNHAUER, M. 1939a. Zur Staphyliniden-Fauna von China und Japan (10. Beitrag).
Entomologisches Nachrichtenblatt, Troppau 12: 97-109.

BERNHAUER, M. 1939b. Neuheiten der Chinesischen Staphylinidenfauna (12. Beitrag). Mit-
teilungen der Münchner Entomologischen Gesellschaft 29: 585-602.

CAMERON, M. 1925. New Staphylinidae from Dutch East Indies. Treubia 6: 174-198.

CAMERON, M. 1933. Staphylinide (Col.) from Mount Kinabalu. Journal of Federal Malay State
Museum 17: 338-360.

CAMERON, M. 1938. New myrmecophilous Staphylinidae (Col.) from East Africa. The
Entomologist’s Monthly Magazine 74: 270-271.

CAMERON, M. 1939. The fauna of British India, including Ceylon and Burma. Coleoptera
Staphylinidae. Vol. 4. London, 410 pp.

CAMERON, M. 1950. New species of Staphylinidae (Col.) from the Malay Peninsula. Annual
Magazin of Natural History 12: 89-131.

ERICHSON, W. F. 1839. Die Käfer Mark Brandeburg. Vol. 1. Berlin, F. H. Morin, 740 pp.

GRAVENHORST, J. L. C. 1802. Coleoptera Microptera Brunsvicensia. Brunsvigae, 206 pp.

HEER, O. 1842. Fauna Coleopterorum Helvetica. Turici, 652 pp.

KISTNER, D. H. & JACOBSON, R. 1975. The Natural History of the Myrmecophilous Tribe
Pygostenini (Coleoptera: Staphylinidae). Sociobiology 1: 155-384.

KRAATZ, G. 1857. Beitrag zur Kenntniss der Termitophilen. Linnaea Entomologica 11: 44-56.

KRAATZ, G. 1858a. Naturgeschichte der Insecten Deutschlands. Coleoptera. Berlin, 1080 pp.

KRAATZ, G. 1858b. Einige neue und ausgezeichnete Staphylinen-Gattungen. Berliner Entomo-
logische Zeitschrift 2: 361-368.

KRAATZ, G. 1859. Die Staphyliniden-Fauna von Ostindien, insbesondere der Insel Ceylon.
Archiv fiir Naturgeschichte 25: 1-196.

MAEKLIN, F. G. 1880. Ytterligare Diagnoser Oefver Nagra Nya Coleopter-Arten. Öfversigtv af
Finska Vettenskaps Akadademie Förhandlingen 22: 79-86.

MOTSCHULSKY, V. 1858. Enumération des nouvelles especes de Coléoptéres rapportées de ses
voyages. Bulletin de la Société impériale des Naturalistes de Moscou 3: 204-264.

PACE, R. 1984. Aleocharinae della Thailandia e della Birmania riportate da G. de Rougemont.
Bollettino del Museo civico di Storia naturale de Verona 2: 427-468.

PACE, R. 1986. Aleocharinae dell’ Asia Sudorientale reccolte da G. de Rougemont. Bollettino
del Museo civico di Storia naturale di Verona 13: 138-237.

PACE, R. 1987a. Aleocharinae riportate dall’Himalaya raccolte da Marc Tronquet e Georges
Ledoux. Bollettino del Museo civico di Storia naturale di Verona 14: 403-419.

PACE, R. 1987b. Aleocharinae riportate dall’ Himalaya dal Prof. Franz. Parte III. Nouvelle
Revue d’Entomologie 4: 117-131.

PACE, R. 1987c. Aleocharinae riportate dall’Himalaya Nepalese. Aleocharinae raccolte dal
Prof. Dr. J. Martens. Courrier Forschungs-Institut Senckenberg 93: 384-441.

PACE, R. 1990a. Aleocharinae delle Filippine. 82° Contributo alla conoscenza delle Aleo-
charinae. In: BERTI, N. (ed.) Miscellanées sur les Staphylins. Mémoires du Museum
National d’Histoire Naturelle (A) 147: 57-113.

PACE, R. 1990b. Aleocharinae del Nepal. 101° Contributo alla conoscenza delle Aleocharinae.
In: BERTI, N. (ed.), Miscellanées sur les Staphylins. Mémoires du Muséum National
d’Histoire Naturelle (A) 147: 155-169.

PACE, R. 1991. Aleocharinae Nepalesi del Museo di Ginevra. Parte III. Autaliini ad Athetini
(Coleoptera, Staphylinidae). Revue suisse de Zoologie 98: 107-158.

PACE, R. 1992. Aleocharinae Nepalesi del Museo di Ginevra. Parte IV. Myrmedoniini
(Coleoptera, Staphylinidae). Revue suisse de Zoologie 99: 125-145.

PACE, R. 1993. Aleocharinae della Cina. Bollettino del Museo civico di Storia naturale di
Verona 17: 69-126.

982 ROBERTO PACE

Pace, R. 1998a. Aleocharinae della Cina: Parte I. Revue suisse de Zoologie 105: 139-220.

PACE, R. 1998b. Aleocharinae della Cina: Parte II. Revue suisse de Zoologie 105: 395-463.

PACE, R. 1998c. Aleocharinae della Cina: Parte III. Revue suisse de Zoologie 105: 665-732.

SEEVERS, C. H. 1953. Two new genera of Myrmecophilous Staphylinidae form the Philippines.
Philippine Journal of Science 81: 125-131.

STEPHENS, J. F. 1832. Illustration of British Entomology. Mandibulata. Vol. 5. London, V.
Baldwin & Cradock, 448 pp., 4 pls.

THOMSON, C. G. 1852. Oefversiat af de i Sverige Funna Arter af Slaegtet Homalota (Manner-
heim). Ofversigt af Kungliga Svenska Vetenskaps. Akademiens Foerhandlingar 9:
131-146.

THOMSON, C. G. 1858. Skandinaviens Coleoptera, synoptiskt bearbeitate. Tom III. Lund,
Berlinska Boktryckeriet, 278 pp.

WASMANN, E. 1896. Neue Termitophilen und Termiten aus Indien. Annali del Museo civico di
Storia naturale di Genova 36: 612-632.

WASMANN, E. 1898. Eine neue dorylophile Tachyporinen-Gattung aus Siidafrika. Wiener Ento-
mologische Zeitung 17: 101-103.

REVUE SUISSE DE ZOOLOGIE

Tome 105 — Fascicule 4

ZOOLOGIA ET BOTANICA ‘98, Geneva, 18-20 February 1998 (Joint meeting
of the Swiss Society of Zoology and the Swiss Society of Botany). . .

BAup, François J. Nécrologie Louis de Roguin (1948 - 1998). .........

MAHNERT, Volker & Renata DE ANDRADE. Description of a new troglo-
philous species of the genus Maxchernes Feio, 1960 (Pseudoscor-
piones, Chernetidae) from Brazil (Sao Paulo State)...............

AssınG, Volker. New species and records of Masuria Cameron from Nepal
(Coleoptera, Staphylinidae, Aleocharinae). .....................

LOURENÇO, Wilson R. & Lionel Mono». Redescription of Compsobuthus
rugosulus (Pocock, 1900) (Scorpiones, Buthidae) based on specimens
RO AKT SEAMEN Re TOI RE

ZEIDAN-GEZE, Najla & Daniel BURCKHARDT. The jumping plant-lice of
ebanon (Hemipteras Esyilloidea) 6 tain: ee

WUEST, Jean. Les organes producteurs de phéromones de quelques Hespé-
ndesi(bepidoptera, Hespeniidae, Hesperinae)tr en ee

KURBATOV, Sergei A. & Ivan LOBL. Nouvelles espèces asiatiques du genre
Bryaxis et quelques données sur des espèces connues (Coleoptera:
Staphylinidaes Pselaphinae) es ark ee ee ee eee ere eee ©

ANGELINI, Fernando & Jonathan Cooter. A new species of Stetholiodes
Fall, 1910 (Coleoptera, Leiodidae, Agathidiini) from Taiwan.......

MAHUNKA, Sandor. New data on Oribatids (Acari: Oribatida) from St.
Lucia (Antilles). (Acarologica Genavensia LXXXIX).............

PAGES, Jean. Japygoidea (Diplura) du Sud-Est asiatique n° 8: Indonésie
(Java, Bali), Singapour et Brunei - Dicellurata Genavensia XXIII -. .

AZPELICUETA, Maria de las Mercedes & Atila Esteban GoszTONYI.
Redescription of Diplomystes mesembrinus (Siluriformes, Diplo-
IMNYSUCIO RR AIR à hs ERI MEL DIOS, pi. Flex

PACE, Roberto. Aleocharinae della Cina: Parte IV (Coleoptera, Staphy-
Ina): eRe RS RE ER e a ARR NER ;

Pages

733-765

767-770

771-775

777-787

789-796

797-812

813-822

823-833

835-837

839-877

879-899

901-910

911-982

REVUE SUISSE DE ZOOLOGIE
Volume 105 — Number 4

ZOOLOGIA ET BOTANICA ‘98, Geneva, 18-20 February 1998 (Joint meeting
of the Swiss Society of Zoology and the Swiss Society of Botany)...

BAUD, François J. Nécrologie Louis de Roguin (1948 - 1998)..........

MAHNERT, Volker & Renata DE ANDRADE. Description of a new troglo-
philous species of the genus Maxchernes Feio, 1960 (Pseudoscor-
piones, Chernetidae) from Brazil (Sao Paulo State)...............

AssınG, Volker. New species and records of Masuria Cameron from Nepal
(Coleoptera, Staphylinidae, Aleocharinae)......................

LOURENÇO, Wilson R. & Lionel Monop. Redescription of Compsobuthus
rugosulus (Pocock, 1900) (Scorpiones, Buthidae) based on specimens
dromePakastans 2 eo bos EPS LE ae IA RE

ZEIDAN-GEZE, Najla & Daniel BURCKHARDT. The jumping plant-lice of
Lebanon (Hemiptera: /Psylloidea). . RR EEE

WUEST, Jean. The pheromone dispersing apparatus in some Hesperiinae
(Lepidoptera, Hesperiidae, Hesperiinae)...................

KURBATOV, Sergei A. & Ivan LOBL. New Asian species of the genus
Bryaxis, and data on previously known species (Coleoptera: Staphy-
Imidae#Bselaphinae). .. art) ie Ae

ANGELINI, Fernando & Jonathan CooTER. A new species of Stetholiodes
Fall, 1910 (Coleoptera, Leiodidae, Agathidiini) from Taiwan.......

MAHUNKA, Sandor. New data on Oribatids (Acari: Oribatida) from St.
Lucia (Antilles). (Acarologica Genavensia LXXXIX).............
PAGES, Jean. Japygoidea (Diplura) from South-East Asia n° 8: Indonesia
(Java, Bali), Singapore and Brunei - Dicellurata Genavensia XXIII -.
AZPELICUETA, Maria de las Mercedes & Atila Esteban GOSZTONYI.
Redescription of Diplomystes mesembrinus (Siluriformes, Diplo-
DUTY /S C1 LAC) LITI EINES LISTE
PACE, Roberto. Aleocharinae from China: Parte IV (Coleoptera, Staphy-
Landa) iran eee

Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX

Pages

153)

767

771

(0/7)

789

TST

813

823

835

839

879

901

911

PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE
CATALOGUE DES INVERTEBRES DE LA SUISSE

ÉaS CES ARCODINÉ SIPARE NPENARDE 3 LITTA OTT Fr. 12.—
2MPEANMELORODES Path STINGELINERE Ce ee Le Cr ce CEE Cr 12.—
SRARAIGNEESIPAER“ DEIEESSERTEG Eee ee 42.—
BUS ORODES paral CAR Uy ER RO ose de cite clei tae Le Cite 8.—
S'MPSEUDOSCORPIONSPA TRADE KESSERTEE N M ee oe ae ee ete deren 5.50
CMINEUSOIRESIPARNE WANDRE sere eae Ne 18.—
7, OULGOECHETES parE. PIGUET et Ko BRETSCHER. LT n 18.—
SEX CORERODES panNISnnIEBAUD IR ee nad... he Moet ee ee 18.—
SEO LIEIONSIPareREDEIESSERIEIERE IS IO I 11.—

lDMSCORPIONSIPATRIDENLESSERTE Re Eee CU 3.50
11. ROTATEURS par E.-F. WEBER et G. MONTET .............................. 38.—
12. DÉCAPODES DA Ve CARTE EE ee ee et EN RT: 11.—
13. ACANTHOCEPHALES PAGE ANDRE au TRI seele IE 11.—
14, GAS TERORRICHES panGIMONTET ee oe O 18.—
1S Se ANMIPHIPODES par CARL Gb eee ee te ee a enon aoe CIE 12.—
16. HIRUDINEES, BRANCHIOBDELLES

et POLYCHETES Pan wANDRE ee A see a IR 17.50
ACESRODESIPARONFUARMANNERE RITI OCHO BUS 30.—

REVUEIDEIRALEOBIOLOGIE Ns... nn Echange ou par fascicule Fr. 35.—

LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris)....... par fascicule Fr. 35.—

CATALOGUE ILLUSTRE DE LA COLLECTION LAMARCK h

APPARTENANT AU MUSEUM D'HISTOIRE NATURELLE DE GENEVE

IT partie — Fossiles — 1 vol. 4° avec 117 planches, 1918........................ Fr. 300.—
COLLEMBOLENFAUNA EUROPAS

von H. GISIN, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck 1984).............. Fr. 30.—
THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND

DISTRIBUTION WITH KEYS FOR DETERMINATION

by2ISNOSER, 346 pages, II figures intext; 1973. eo geese cc Fr. 30.—
CLASSIFICATION OF THE DIPLOPODA

PANRIChATANESEOEEMAN 237 pases UOT IE cre cic ee Fr. 30.—
LES OISEAUX NICHEURS DU CANTON DE GENEVE

par P. GEROUDET, C. GUEX et M. MAIRE

351 pages, nombreuses cartes et figures, 1983 ................................. Fr. 45.—
CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS

CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES,

SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ

A LA CONNAISSANCE DES ECHINODERMES ACTUELS

par Michel JANGOUX, 67 pages, 11 planches, 1985 ............................. Fr. 15.—
RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE

(COTENTIN-BAIE DE SEINE, FRANCE)

par Y. FINET, J. WÜEST et K. MAREDA, 62 pages, 1991 .......................... Fr. 10.—
GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN;

SHELLS AND RADULAS

bYAYAEINER Is WiUESTpand KEMAREDAMIOO2 RER I, PIER O Fr. 30.—
O. SCHMIDT SPONGE CATALOGUE

par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992 .......... Fr. 40.—
ATLAS DE RÉPARTITION DES AMPHIBIENS

ET REPTILES DU CANTON DE GENEVE

par A. KELLER, V. AELLEN et V. MAHNERT, 48 pages, 1993 ...................... Fr. 15.—
THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS:

A DOCUMENTED FAUNAL LIST

PAAVESIEINE RMS ODA Le SUORE oboe IE Fr. 30.—
NOTICE SUR LES COLLECTIONS MALACOLOGIQUES

DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE

pamlean Claude CAILLIEZ 49]pares 1 II ST ON Fr. 22.—
PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS

OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr. 160.—

CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE)

(Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997 ................... Fr. 50.—

| ALES

Ber

È Rida

eon EN Kar AI
Wa SVC I: |

Volume 105 - Number 4 - 1998
Revue suisse de Zoologie: Instructions to Authors

The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific
results based on the collections of the Museum d’histoire naturelle, Geneva. Submission of a manuscript implies
that it has been approved by all named authors, that it reports their unpublished work and that it is not being
considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of
colour plates and large manuscripts. All papers are refereed by expert(s).

Papers should conform to the following general layout:

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces,
name(s) in full and surname(s) of author(s), and full address(es).

Introduction. A short introduction to the background and the reasons for the work.

Materials and methods. Sufficient experimental details must be given to enable other workers to repeat the
work. The full binominal name should be given for all organisms. The Zoological Code must be strictly followed.
Cite the authors of species on their first mention. Use SI units and the appropriate symbols.

Sex symbols should be used rather than “male” and “female”.

Discussion. This should not be excessive and should not repeat results nor contain new information, but
should emphasize the significance and relevance of the results reported.

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.

References should not be interspaced and, in the case of several papers by the same author, the name has to be
repeated for each reference.

Also, we encourage authors to submit the text on floppy disk (3,5°° or 5 1/4”’, Macintosh or IBM compatible,
with “Microsoft Word” or similar programmes). The authors should refrain from using special codes for text
formating.

Figures. These may be line drawings or half tones and all should be numbered consecutively, and their position
indicated in the text. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and
lettering should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors
should refrain from mixing the drawings and half tones. Original drawings will not be returned automatically. The
Revue suisse de Zoologie declines responsability for lost or damaged slides or other documents.

Legends to figures. These should be typed in numerical order on a separate sheet.

Proofs. Page proofs only are supplied, and authors may be charged for alterations (other than printer’s errors) if
they are numerous.

Offprints. The authors receive 25 offprints free of charge; more copies may be ordered at current prices when
proofs are returned.

Correspondence. All correspondence should be addressed to

Revue suisse de Zoologie
Muséum d histoire naturelle
CP 6434
CH-1211 Genève 6
Switzerland.
Phone: (022) 418 63 33 - Fax (022) 418 63 01
e-mail: volker.mahnert @ mhn.ville-ge.ch
Home page RSZ: http://www-geneva-city.ch:80/musinfo/mhng/publications/revues.htm

HECKMAN
BINDERY INC.

APR 99

Round To: N. MANCHESTER,
To Plast INDIANA 46962

PRET PENTIER
PAL

LL Hn
Funken

ATOME

DES

th
raditeie”

FLIEHEN

dei i adria tt Bat ATE
PESTO

AP
bye a bady
Al si

TION LIBRARIE

1993

Internet Archive Audio

Featured

Top

Images

Featured

Top

Software

Featured

Top

Books

Featured

Top

Video

Featured

Top

Mobile Apps

Browser Extensions

Archive-It Subscription

Save Page Now

Full text of "Revue suisse de zoologie"

See other formats