Revision of the Palearctic Species of the Myrmecophilous Genus Pella(Coleoptera, Staphylinidae, Aleocharinae)
Munetoshi Maruyama
Department of Zoology, National Science Museum,
3–23–1 Hyakunin-chô, Shinjuku-ku, Tokyo, 169–0073 Japan(http://www.myrmecophile.net/)

Abstract A biosystematic study of the Palearctic species of the myrmecophilous genus Pella(Coleoptera, Staphylinidae, Aleocharinae, Lomechusini, Myrmedoniina) is presented. The adult morphology of Pella is investigated into details in comparison with several other taxa of the Lomechusini.
Pella is taxonomically revised, and is arranged as a distinct genus. Myrmelia, Pellochromonia andLepla, formerly treated as subgenera of Zyras, are synonymized with Pella. Forty-one species including 15 new species (Pella horii, P. kidaorum, P. cooterorum, P. jelineki, P. schillhammeri, P.
kinomurai, P. primorskyiana, P. hlavaci, P. iberica, P. kishimotoi, P. coreana, P. plutenkoi, P. intermedia, P. masakoae, P. zhoui) are recognised in the genus, arranged into 16 species-groups and redescribed/described with bionomical and host ant information. The following junior synonymies areproposed: Zyras (Pella) micropterus (P. jureceki), Z. (P.) almaatensis (P. bohaci), and Z. (P.) esau(P. cinctipennis). Seven species are excluded from Pella, newly assigned to the other genera/subgenera and redescribed. They are: Zyras (Zyras) coloratus, Z. (Z.) iridescens, Z. (Z.) quasar, Z. (Glossacantha) ceylonicus, Z. (G.) reelsi, Myrmoecia urartu, and Dvorakatheta wrasei (n. gen.). Life cycles and behaviour of the Japanese species of Pella are shortly reported based on the field and laboratory observations.

INTRODUCTION
The myrmecophilous beetle genus Pella Stephens, 1835, belonging to the subtribe Myrmedoniina of the tribe Lomechusini (family Staphylinidae, subfamily Aleocharinae) is represented
by about 30 named species described from the Palearctic region. They are about 3.5 to 7.0 mm inlong and associated with ants of the genera Lasius Fabricius, 1804; Formica Linnaeus, 1758;
Tapinoma Foerester, 1850; Liometopum Mayr, 1861; and Crematogaster Lund, 1831. Most ofthem are inquilines of Lasius (Dendrolasius) ants. Several species have been known as scavengers in hosts’ nests or predators on host workers (Yasumatsu, 1937; Kistner, 1972; Hölldobler
et al., 1981).
Pella beetles have been studied by several researchers from different points of view, especially because of their myrmecophily. However, our current knowledge of Pella mainly consists ofscattered descriptive works. On the other hand, myrmecophilous insects have recently been attractting attention of researchers as material for studies on host-inquiline relationships and coevolution, and works using modern methods have been published (Axen, 2000; Elmes et al.,
1999; Akino, 2002; Fraser et al., 2002; Schönrogge et al., 2002; Pierce et al., 2002, etc.). Because the Pella beetles are among the commonest myrmecophiles in their distributional areas,

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they are very useful for the study of biology of myrmecophily. A comprehensive systematic studyof the beetles, therefore, would contribute to forward the study of myrmecophily, and the present
study has been carried out in this perspective, dealing with morphology, taxonomy and bionomics of the Pella beetles.

MATERIALS AND METHODS
Material from Institutions and Private Collections
The majority of species of Pella were originally described without details useful for distinguishing them, especially among closely related species. Therefore, examination of type series isnecessary for reliable identification. Fortunately, the type specimens of most Pella species wereaccessible through loans or visits to museums or personal collections.
Most of the other specimens examined were collected by some colleagues and myself. In addition, an amount of unidentified material from areas to which I was unable to have access forthis study was supplied from some institutional or personal collections.
Depositories of the type specimens examined and the other specimens studied are as follows.
Institutions:
BMNH — The Natural History Museum, London (M. Brendell).
CNC — Canadian National Collection of Insects, Agriculture and Agri-food Canada, Ottawa, Ontario (A. Davies).
CNU — Department of Biology, Chungnam National University, Daejeon (K.-J. Ahn).
DEI— Deutsches Entomologisches Institut, Müncheberg (L. Zerche).
EEEU — Laboratory of Environmental Entomology, Ehime University, Matsuyama (N.
Ohbayashi, M. Sakai).
HUM — The Hokkaido University Museum, Sapporo (M. Ôhara).
IBSS — Institute of Biology and Soil Science, Vladivostok (V. Kuznetsov).
IRSNB — Institut Royal des Sciences Naturelles de Belgique, Bruxelles (D. Drugmand).
IZAS — Institute of Zoology, Academia Sinica, Beijing (H.-Z. Zhou).
MHNG — Muséum d’Histore Naturelle, Genève (G. Cuccodoro).
MHNL — Muséum d’Histore Naturelle, Lyon (M. Virgile).
MNHN — Muséum national d’Histore Naturelle, Paris (T. Deuve).
NHMW — Naturhistorisches Museum Wien (H. Schillhammer).
NHRS — Naturhistoriska riksmuseet, Stockholm (B. Viklund).
NMEG — Naturkundemuseum Erfurt, Erfurt (M. Hartmann).
NMP — Národní Muzeum v Praze (J. Jelínek).
MNV — Museo Civico di Storia Naturale di Verona (A. Zanetti).
NSMT — National Science Museum, Tokyo-to (S. Nomura).
SCM — Sagamihara City Museum, Sagamihara (H. Moriya).
SEHU — Systematic Entomology, Hokkaido University, Sapporo (K. Yoshizawa).
TUA— Tokyo University of Agriculture, Atsugi, Kanagawa (S. Okajima).
ZMHB — Museum für Naturkunde der Humboldt-Universität, Berlin (M. Uhlig, J. Frisch).

Palearctic Species of Pella

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Private collections:
cAss— Volker Assing (Hannover, Germany).
cDvor — Miroslav Dvořák (Praha, Czech Republic).
cHlav — Peter Hlaváč (Košce, Slovakia).
cMar — Munetoshi Maruyama (Tokyo-to, Japan).
cRoug — Guilliame de Rougemont (Londinières, France).
cSch— Michael Schülke (Berlin, Germany).
Material whose depository is not indicated belongs to my collection (cMar), SEHU, HUM
and NSMT, or non-type material of common species that are represented by more than 50 specimens in the present study. Paratypes of some new species to be described in this paper and identified specimens of common Japanese species will be distributed to the collections above.
Specimen Data
Specimen data is in many cases not followed the original spellings written on label but corrected for more precise and detailed data. But in the case of type material, and in the case of old
specimens, e.g., which was collected before the beginning of 1900’s and was historically important, the original spellings on the label were transcribed.
All the types of Pella described by Gravenhorst (1802, 1806) and Märkel (1842, 1845) deposited in ZMHB are labelled not any detailed data but a catalogue number. However, more detailed data are separately described in “Catalog. General. Musei Zoologici Berolinen” housed in
the museum, and based on which actual data are transcribed in square brackets in Type material.
All the type specimens studied in the present paper are labelled, “HOLOTYPE [or, SYNTYPE, LECTOTYPE, PARALECTOTYPE, PARATYPE] Myrmedonia...[name of species, author and year in the original description] det. Maruyama, 200.. [year of examination]”.
Collecting
I investigated more than 120 nests of Lasius ants in Japan (Hokkaidô, Honshû, Shikoku),
Korea, the Russian Far East and Slovakia, and collected about 7,000 specimens of Pella.
Pella beetles are usually abundant in the hosts’ nests, above all in the nests of the main hostants Lasius (Dendrolasius) spp. They also walk around the nests and columns (see, Chapter 3).
Therefore, the most effective collecting method is to sift dead leaves around the nest and columns
using a sieve and a white tray. Beetles dropped on the white tray should immediately be aspirated. Another method, which was sometimes adopted, is pitfall traps that were set around the nestby the use of plastic cups. Beetles collected were transferred to a tube with a piece of cotton
paper and a small quantity of ethyl acetate and were left in the tube for half a day. The killed insects were mounted each on a piece of paper card. Some beetles were kept alive for rearing.
Dissecting and Drawing
Dried or ethanol-preserved specimens were used for morphological observations. Dissection
was made for observation of mouthparts and genital parts (abdominal terminalia), and dried
specimens were softened before dissection by warming (60°C, 30–60 min.) each in a small ceramic bowl (2.5 cm in diameter) with a small amount of water.
Then the specimens were dissected in water under a high magnification stereomicroscope

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M. Maruyama(Olympus SZ-40, 6.7–80). Eighth to 10th abdominal segments were removed and dissected formost specimens.
Mouthparts of some specimens were dissected: a sharp needle was thrust into the gular plate
posterior to the mentum to destroy the muscular attachment of the mouthparts and to remove thementum, labium, labrum, and maxilla from the head.
These body parts were soaked in warm 5–8% solution of potassium chloride (60°C,
20–60 min), cleaned in 30% ethanol (5 min) and dehydrated in 99% ethanol (5 min). When necessary, mouthparts were stained in a warm mixture of acetic fuchsine (5%) and lactic acid (60°C,
60–120 min) before cleaning. The dehydrated material was mounted in Euparal (ChromaGesellschaft) on a slide glass card, which was pinned under the insect from which the parts were
removed (Maruyama, 2004).
Spermatheca was drawn in water before mounting in Euparal because it was usuallytransshaped by the osmotic pressure of the medium.
A transmission microscope (Olympus BH2, 20–600, with a drawing device, or ocularmesh) was used for observations of microstructures and line drawings.
Drawing of forebody and antenna of each species was made under the stereomicroscope
above-mentioned by the use of ocular mesh set in eyepiece. For drawings of antenna, 1st and 2nd
segments were drawn in dorsal view, and 3rd to 11th segments were drawn in lateral view.
Symbiotic Hosts
Symbiotic hosts were mostly identified by myself and some by V. Assing, F. Ito and K. Kinomura. The scientific names of ants generally follow Bolton (1995), but those of Dendrolasius
ants follows Radchenko (2005) and Maruyama (2005). Lasius (Dendrolasius) fuji was formerly
recognised as the eastern Palearctic population of Las. (D.) fuliginosus, and was erroneously assigned to “Las. (D.) nipponensis” by Espadaler et al. (2001).
The species name of symbiotic hosts are abbreviated in “Type material” and “Other material”
as follows: FP, Formica pratensis Retzius, 1783; LCAF, Lasius (Cautolasius) flavus (Fabricius,1782); LCHU, Las. (Chthonolasius) umbratus (Nylander, 1846); LCH, Las. (Ch.) sp.; LDC, Las.
(Dendrolasius) capitatus (Kuznetsov-Ugamsky, 1927); LDFJ, Las. (D.) fuji Radchenko, 2005;
LDFL, Las. (D.) fuliginosus (Latreille, 1798); LDN, Las. (D.) nipponensis Forel, 1912; LDO, Las.
(D.) orientalis Karawajew, 1912; LDS, Las. (D.) spathepus Wheeler, 1910a; LL, Las. (Lasius) sp.;
LLB, Las. (Lasius) brunneus (Latreille, 1798); LLJ, Las. (Las.) japonicus Santschi, 1941; LM, Liometopum microcephalum (Panzer, 1798); TE, Tapinoma erraticum (Latreille, 1798).
Terminology and Abbreviations
Terminology of morphology used in this study follows Blackwelder (1936), Sawada (1970a,
1972, 1974), Yosii & Sawada (1976), Naomi (1987a, 1987b, 1988a, 1988b, 1988c, 1988d, 1989a,
1989b, 1989c, 1989d, 1990, 1997) and Hanley (2002). Numbers of setae, macrosetae, pores, andcampaniform sensilla are confined to one body side, except for the pseudopores of the prementum and the pores of the medial pore field of epipharynx.
All measurements in the text are given in millimetres, and the following abbreviations areused for measurements: AL, antennal length; BL, body length, approximate whole length; EL,eye length, major axis; ELL, elytral length; ELW, elytral width; FBL, fore body length, from apexof clypeus to posterior margin of elytra; HL, head length, from apex of clypeus to the posterior

Palearctic Species of Pella5

margin of head capsule, except for preocciput; HTL, length of hind tibia; HW, head width; PL,pronotal length; PW, pronotal width.
Collecting methods are abbreviated as follows: fit, flight interception trap; pt, pitfall trap; tt,
truck trap.
Rearing
Rearing was made, above all, for observing immature stages in ant-free environment. Somebeetles were reared with ant colonies (without queen) for observing their behaviour in the presence of ants. The reared beetles belong to Pella humeralis, P. funesta, P. comes, P. socia, P. kidaorum, P. japonica, P. kinomurai, P. lugens, P. masakoae, P. spreta, P. laticollis, and P. indiscreta.
Five to ten adult beetles were kept in a plastic box (L18 cmW9 cmH4.5 cm), which was
bedded with mixture of peat moss and soil for gardening and provided with some small pieces oftree bark on the soil as shelters for beetles. The peat moss was used for preventing fungus growthand keeping moisture of the soil. The box was placed in room air temperature (18–25°C). Bothadults and larvae were fed with dead worker ants of Lasius (Dendrolasius) fuji or L. (D.) spathepus, which had been preserved in a freezer.

CHAPTER 1.

MORPHOLOGYThis chapter describes the general external morphology of the adult Pella beetle in comparison with other lomechusine species.
The tribe Lomechusini is composed of two subtribes the Lomechusina and Myrmedoniina,
and more than 200 genera and subgenera from throughout the world (Maruyama, in prep.). In the
Myrmedoniina the following species have been examined: Zyras (s. lat.) spp., Pedinopleurusspp., Diplopleurus spp., Tetrabothrus spp., Orphnebius spp., Drusilla spp., Amaurodera spp. andApteranillus spp.; in the Lomechusina: Lomechusa spp., Lomechusoides spp. and Xenodusa spp.
These species are referred to only by their generic names when character states are almost same
within each of the genera. Other species lacking tribal status are all members of the Lomechusini.
In the subfamily Aleocharinae, only a few works which study detailed morphology of wholebody are known. The present study is expected to be one of the frameworks of morphologicalstudy of the Aleocharinae.
General Structure
Body shape
The body shape (Fig. 1) in Pella species is slender or somewhat robust, more or less depressed above, and medium to large sized among aleocharine taxa. The body shape is highly generalised in spite of its myrmecophily, i.e., non-limuloid, non-myrmecoid and lacking trichome orlarge secretary gland, and thus it is not very different from that of free-living aleocharines living
in soil or under bark. However, somewhat wide pronotum in Pella species is common to many
myrmecophilous aleocharines and indicative of their myrmecophily, though it is homoplastic
states sometimes observed in non-myrmecophilous aleocharines, e.g., Aleochara spp. and Oxypoda spp.

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Fig. 1.

Pella limbata (Paykull).

Habitus.

Colour
The ground colour in Pella species is usually pale brown to black, but exceptionally the
pronotum of P. ruficollis is reddish orange. In colour or colour pattern, the members of the funesta group and P. ruficollis are very similar to their host ants, Lasius (Dendrolasius) spp. and Liometopum microcephalum, respectively. Most ants have been known to secret repellent substances or to be capable of stinging against enemies, and most predacious animals avoid ants
(Blum & Herman, 1978). Thus, it has been suggested that these colour patterns are their Batesian
mimicry to the host ants, and it was currently proven by experiments using a predator of ants,
Hyla japonica Günther, 1858 (Japanese tree-frog) (Taniguchi et al., 2005).
Body hairs
The body of Pella species is covered mainly with two kinds of hairs, the seta and themacroseta, which can be clearly distinguished. The seta is white or yellow, narrower than themacroseta and usually recumbent. The macroseta is black, stout and suberect. However, in the
coreana group, the macrosetae are poorly differentiated from the setae in length and colour.

Palearctic Species of Pella

Fig. 2.

7Pella limbata (Paykull). — A, Head, dorsal view; B, head, lateral view.

Some kinds of secondary hairs are observed on the surface of hindwing, the scutoscutellar
suture (Fig. 8 B) and the apex of female 8th sternite (Fig. 11 H) in some species, and they are referred to as pubescence or sensory setae.
Head
Head capsule
The head capsule (Figs. 2A, 2B, 3A) in Pella species is almost circular in dorsal and ventral
views, wider than long, and convex dorsally around middle or posterior part. These states arecommonly observed in most lomechusines and other aleocharines. In some species, head capsuleis depressed above, especially in male.
Occipital suture. The occipital suture (Figs. 2 A, 2 B, 3 A) is well developed in Pella and8

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Fig. 3. Pella limbata (Paykull). — A, Head, ventral view; B, antenna, dorsal view; C, 1st to 3rd segments of antenna, dorsal view; D, 10th and 11th segments of antenna, dorsal view.

extends from the ventro-lateral part of the head to near the posterior articulation of the hypostoma. This character state is observed in most aleocharine tribes.
Postoccipital suture. The postoccipital suture (Figs. 2 A, 3 B) is in a generalised state in
Pella species as in most aleocharines, and “neck” is not observed. In some genera of the Myrmedoniina, e.g., Drusilla, Tetrabothrus and Orphnebius, postoccipital suture is strongly constricted
to form a “neck”.

Palearctic Species of Pella

9Eyes
The compound eye (Figs. 2 A, 2 B) in Pella species is dorsolateral, tending to be situated on
the anterior part of the head, and 0.29–0.48 times as long as the head width. Small setae appear
among the facets. These states are common to most aleocharines.
The size of the eye is variable with species and is taxonomically informative.
The eye size probably depends on diurnal, nocturnal or day-and-night activity. While thespecies of Pella having smaller eyes (0.4 times head width), e.g., the species of the humeralis
group and the funesta group, are active day and night, the species having larger eyes (0.40
times head width), e.g., the species of the lugens group and the spreta group, appeared to be nocturnal in my field observations. The large-eyed species of the Lomechusini, e.g., Zyras spp.,
Pedinopleurus spp., Tetrabothrus spp. and Drusilla spp. are often attracted to light traps, and thissuggests that they are also nocturnal.
Antennae
The antennae (Figs. 3 B–3 D) in Pella are usually shorter than the combined length of head,pronotum and elytra, except in P. comes (longer) and P. rambouseki (almost the same in length)
and more or less depressed dorsoventrally. Fourth to 10th segments (Fig. 3 B) are usually wider
than long and subquadrate in lateral view. Their surfaces are covered with setae and macrosetae,
the setae becoming sparser and smaller toward the apex (Figs. 3 C, 3 D). These states are common to most aleocharines.
Relative lengths of antennal segments, especially those of 1st to 3rd segments and the 11th
are useful to diagnose species. Usually the 1st segment is the longest but sometimes the 11th islonger than it.
Mouthparts
LabrumThe labrum (Fig. 4 A) is relatively generalised in structure in Pella species. It is submembranous around the antero-medial part (sometimes also around the lateral margins). The surface isdensely covered with pseudopores except in the lateral area and with 10–12 setae around the apical half.
Epipharynx
The epipharynx (Fig. 4 B) in Pella is generalised in structure. The number of lateral setae isnormally 5, and these are situated on the mesolateral part of the epipharynx. The medial sensory
field have 80–90 pores, and the lateral field is covered with several pores irregularly among scalelike sculptures.
Mandibles
The mandibles in Pella species (Figs. 5 A, 5 B) are simple in shape and lack mola or any pubescent patch. However, they are slightly asymmetric, and the right mandible has a small tootharound the middle of the inner margin.
MaxillaThe maxilla in Pella species (Fig. 5 C) is generalised in shape as in most aleocharines, except
for the elongated galea.

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Fig. 4.

Pella limbata (Paykull). — A, Labrum, dorsal view; B, epipharynx (labrum, adoral view).

The elongated galea is unique to the Lomechusini and has been considered to be an autapomorphy of the tribe, though the maxilla is much shortened in the Lomechusina as in other aleocharines, and this is considered to be a secondary shortening probably due to their myrmecophilous behaviour (Maruyama, in prep.).
MentumThe general shape of the mentum in Pella species (Fig. 5 D) is trapeziform, and the anterior
and posterior margins are completely truncate. The surface of the disc is densely covered with

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Fig. 5. Pella limbata (Paykull). — A, Right mandible, dorsal view; B, ditto, ventral view; C, maxilla, ventral
view; D, mentum, ventral view.

pseudopores except for the lateral area.
LabiumThe structure of labium is highly diversified in the Myrmedoniina, and this is phylogenetically most informative character. Comparative morphology among lomechusine genera is provided
in the following lines.
Prementum, antero-medial margin. The antero-medial margin is shallowly emarginate in
Pella species (Fig. 6 A). In the Myrmedoniina, the depth and shape of the emargination is various

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Fig. 6. Pella limbata (Paykull). — A, Labium, ventral view; B, mentum and labium, lateral view; C, ligula, ventral view; D, antero-lateral part of prementum, ventral view; E, hypopharynx (labium, adoral view).

Palearctic Species of Pella

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according to species.
Prementum, pseudopores. Location: the pseudopores are located around the antero-medial
area of prementum in Pella species (Fig. 6 A). Most lomechusine species are in the same state as
Pella.
Number: the number of pseudopores occurring around the antero-medial area is about 30–50
in Pella species (Fig. 6A). Most lomechusines are similar to Pella species, but the members ofMyrmoecia and some species of Orphnebius completely lack pseudopores.
Size: the pseudopores in Pella are large and only slightly smaller than real pores. The
pseudopores of the lomechusines examined are smaller than a half of real pore in diameter.
Prementum, real pores. Two real pores are located on the mesolateral area of the prementum in Pella species (Fig. 6 A). This state is common to most lomechusines and other aleocharines. In contrast, Diplopleurus sp., Tetrabothrus spp. and members of the Lomechusina
have extra real pores, 3, 4 and 4 or 5 in total, respectively.
Prementum, setal pore. One setal pore occurs in the mesolateral area of the prementum inmost aleocharines. That in Pella species (Fig. 6 A) is located slightly anteriorly, and this state iscommon to most lomechusines. The setal pore is located around the middle of the prementum inZyras (Trachydonia) sp., Zyras (Rhynchodonia) sp. and Pedinopleurus sp.
Prementum, medial seta. A pair of medial setae are present at the antero-medial apex of theprementum in all lomechusines.
Prementum, internal ridge. Length: a pair of internal ridges are located at the backside of
the prementum in the Aleocharinae. Those in Pella species (Fig. 6 D) are very small and only
recognised around the backside of each medial seta, and a similar condition is observed also in
Myrmoecia species. The length of the internal ridge is various in the Lomechusini. The ridge inmost lomechusines is about 1/2 to 5/6 as long as the prementum, and in Diplopleurus sp. very
long, extending to the postero-medial projection of the apodeme.
Shape: the internal ridge in most lomechusines is almost straight, but that in Diplopleurusspp., Tetrabothrus spp. and Zyras (Zyras) spp. is bifid anteriorly.
Apodeme. Lateral lobes (suspensoria): the general shape of the lateral lobe can be divided
into the following two basic types in the Lomechusini: 1) not completely fused with the premental disc and visible in ventral view; 2) completely fused with the premental disc and invisible inventral view. Pella (Fig. 6 A) belongs to the type 1). The type 2) is limited to all members of the
Lomechusina and part of the Myrmedoniina.
Base of prementum: the base of the prementum is roundly emarginate postero-medially andits lateral part is projected in Pella species (Fig. 6 A: arrow). The round emargination is observedin most lomechusines and some other tribes. The round margin is serrate, dentate or projected insome species of Drusilla, Zyras (Zyrastilbus) sp. and Myrmoecia spp. In contrast, the projectionof Diplopleurus spp., Tetrabothrus spp. and Zyras (Zyras) spp. bears the internal ridges. This
state is commonly observed in most aleocharines.
Palpus. General shape: the labial palpus in Pella species (Fig. 6 A) is 3-segmented, and the
2nd segment is almost as long as the 1st. The palpus in all species of the Lomechusini is 3-segmented, but usually the 2nd segment is shorter than the 1st. In the species of the Lomechusina,
the 1st segment is much longer than the combined 2nd and 3rd, while it is shorter than the latterin the other species of the Lomechusini.
Membranous zone: the second segment of the labial palpus in Pella species bears a membranous zone (Fig. 6 A) on the ventrolateral area. This state is commonly observed in the Myrmedoniina and probably unique for the subtribe.

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Setae: the seta a (Sawada, 1972) is situated near the apex of the 1st segment in Pella species.
This state is also observed in Myrmoecia spp., Orphnebius spp. and Tetrabothrus spp.
Ligula. General shape: the ligula in Pella species (Figs. 6 A, 6 C) is bilobed, and the apex ofeach lobe is rounded. It is various in general shape in the Lomechusini. The ligulae of most
species are bilobed and similar to that of Pella species, but those of Diplopleurus sp., Zyras(Zyrastilbus) adesi and some species of Drusilla are elongated and much longer than wide, andthat of Zyras (Trachydonia) sp. is mount-shaped and only apically notched. The ligula in thespecies of the Lomechusina is very short, much wider than long and more or less flattened at the
apex.

Fig. 7. Pella limbata (Paykull). — A, Prothorax, ventral view; B, meso- and metathoraces, ventral view; C, mesoand metasterna, internal view; D, anepisternum and epimeron of metathorax, internal, dorso-lateral view; E,
ditto, internal view.

Palearctic Species of Pella

Fig. 8. Pella limbata (Paykull). — A, Scutellum, dorsal view; B, metanotum and 1st abdominal tergite, dorsal
view; C, right elytron, dorsal view; D, ditto, ventral view; E, left hind wing, dorsal view; F, flabellum.

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Apical setae, presence and number: the members of Pella has no seta at the apex of ligula(Figs. 6 A, 6 C), except for the presence of 3 campaniform sensilla at the apex of each lobe. The
absence of setae is also observed in Myrmoecia spp. but the members of this genus even lacksockets. Most lomechusines, as well as most aleocharines, have 4 apical setae on each lobe.
Diplopleurus spp. and Tetrabothrus spp. have 3 setae on each lobe.
Surface: the surface of the ligula is smooth in Pella species. In contrast, that of Zyras (Trachydonia) sp. is covered with minute granular sensilla.
Hypopharynx. A row of setae appears along the inner margin of the hypopharynx in Pella
species (Fig. 6 E). This state is commonly observed in most lomechusines.
Thorax
Pronotum
In Pella species, the pronotum is subquadrate, subtrapeziform or elliptical in dorsal view, andmore or less flattened. The disc is weakly punctured or covered with microreticulations, andmoderately to densely and uniformly covered with setae. Macrosetae are present along the posterior to lateral margins, and the number is useful in diagnosing species.
The general shape is various by species and very important in recognising Pella species at thefirst glance.
The structure of the pronotum is various in the Lomechusini. In the Myrmedoniina, the general structure is the same as in Pella species but the punctulation and the density of setae are veryvarious by species. In most species of Zyras, pronotal disc is coarsely punctured and sparsely
covered with erect and stout setae. In the canaliculata group of Drusilla, the disc is granulatepunctured and densely covered with stout setae. In contrast, the structure of pronotum is highly
modified in the Lomechusina, in which the pronotum is distinctly depressed or caved mesally andvery finely granulate-punctured.
ElytraThe elytra are subparallel-sided. The inner to posterior margin is more or less margined. The
surface is similar to that of the pronotum but with setae becoming sparser apicad. Macrosetae arepresent along the shoulder to the lateral margin and their number is used for diagnosing species.
Hind wing
The hind wing is entire in Pella species. The veins are weakly sclerotized and very obscure,but generalised as in most aleocharines. In contrast, some species of Drusilla, Tetrabothrus,
Amaurodera and Zyras, and all species of Apteranillus are brachypterous and flightless.
Meso- and metathoraces
In Pella, the mesocoxal cavity is widely separated, and the metasternum is longer thanmesosternum. These states are typical of the Lomechusini and well characterise the tribe. In most
species of the Myrmedoniina, apex of the mesosternal process is rounded, but in some species ofthe Myrmedoniina and all species of the Lomechusina it is widely truncated.
Legs
The legs are generalised in Pella (Figs. 9 A–9 C) as in many other free-living species of the
Aleocharinae, and the apices of the tibiae bear a ctenidium as in the other Myrmedoniina species.

Palearctic Species of Pella

Fig. 9.

17

Pella limbata (Paykull). — A, Foreleg; B, midleg; C, hindleg. A–C, left side, ventral view.

Each femur bears a groove from near base to apex alongside the posterior margin to receive tibia
when legs are squeezed together.

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M. Maruyama

Abdomen
Abdominal segments
General shape. In Pella, the abdomen (Figs. 10 A, 10 B) is highly generalised in shape, andits outline is subparallel-sided in dorsal view. However, P. excepta and P. maura have small projections on the 3rd and 6th abdominal tergites. Many species of the Myrmedoniina bear projections on male abdominal tergites or paratergites.
Eighth tergite. The 8th tergite is narrowed posteriad, truncated or emarginate at the apex
which is crenate or dentate (Figs. 10 A, 11 A, 11 C, 11 E, 11 F). The general shape and number ofmacrosetae covering the surface are useful in diagnosing the species. Basal projection of theapodeme of the male (Fig. 11 E) is usually longer than that of the female (Fig. 11 F).
Eighth sternite. The 8th sternite is narrowed posteriad, rounded or truncated at the apex in
Pella species (Figs. 10 B, 11 B, 11 D, 11 G, 11 H). The general shape and number of macrosetae
covering the surface are useful for diagnosing the species. The 8th sternite is larger in the malethan in the female. The apical setae in the female (Fig. 11 H) are longer and thicker than in themale (Fig. 11 G), and in some species the female apical setae are spindle-shaped and flattened.
Minute pubescence appears along the apical margin (Fig. 11 H) in the females of most species.
Ninth tergite. The 9th tergite (Figs. 12 A, 16 A) covers the aedeagus or vagina from the dorsal to the ventral side in Pella species, and this state is common to most aleocharines. The laterallobes of the male 9th tergite (Fig. 12 A) is long and asymmetric in Pella species as well as inmost aleocharines.
Ninth sternite. The male 9th sternite (Figs. 12 B, 12 C) is oblong oval in Pella species, andits apodeme is submembranous. The apex of the male 9th (Fig. 12 C) sternite is truncated orrounded, bearing a pair of macrosetae and 20–30 setae. The left side (ventral view) of the 9th
sternite is connected with the 9th tergite in the male, and the aedeagus emerges out from the rightside of the 9th sternite (Fig. 12 D). The female 9th sternite (Figs. 16 B, 16 C) is fused with the 9th
tergite except around the apex (Fig. 16 C), which bears about 20 setae. These states are common
to most aleocharines. The surface is covered with many setae and some macrosetae.
Tenth tergite. The 10th tergite in Pella species (Figs. 12 A, 16 A) is almost pentagonal andtruncated or rounded at the apex. The surface is densely covered with setae, which become sparser apicad.
These states are common to most aleocharines. The setose state is various by species in the
Lomechusini.
Male genitalia
Median lobe of aedeagus. The aedeagal median lobe in Pella species (Figs. 13 A–13 E,
14 A–14 E) is mainly composed of the basal capsule and the apical lobe. The compressor plate(Fig. 13 C) is partly or completely covered by the basal capsule. The dorsal bridge (“athetine
bridge”, sensu Seevers, 1978) is present. These structures are commonly observed in the
Lomechusini. The distal crests (Figs. 13 C–13 E) are well developed and is fused with their sides
in Pella species. The general shape is one of the most important characters in diagnosing species.
The inner sac is a tubular structure and evaginate as shown in Figs. 14 D, 14 E. In my observations of some couples of Pella beetles killed during copulation, the apical part of the apicallobe is inserted to the vagina from the ventral split line of the 9th sternite, and then the inner sacof aedeagus is evaginated in the vagina. A pair of the longitudinal bands is folded internally at
the apical opening of the median lobe, and they cover the dorsal face of the inner sac in the pos-

Palearctic Species of Pella

Fig. 10. Pella limbata (Paykull). — A, Second to 8th tergites and paratergites, male, dorsal view; B, 3rd to 8thsternites, male, ventral view. A, B, abdominal segments, location of aedeagus indicated.

19

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M. Maruyama

Fig. 11. Pella limbata (Paykull). — A, Eighth tergite, male, dorsal view; B, 8th sternite, male, ventral view; C, 8th
tergite, female, dorsal view; D, 8th sternite, female, ventral view; E, antero-lateral part of 8th tergite, male,internal view; F, ditto, female; G, apical part of 8th sternite, male, dorsal view; H, ditto, female. A–H, abdominal segments.

Palearctic Species of Pella

21

Fig. 12. Pella limbata (Paykull). — A, Ninth and 10th tergites, dorsal view; B, 9th sternite, ventral view; C, ditto,
apical part; D, 7th to 10th segments in dorso-lateral view, aedeagus emerged out from between lateral sides of
9th tergite and sternite. A–D, male abdominal segments. Abbreviations: T, tergite; S, sternite.

ture of evagination. The surface of the inner sac is covered with minute pubescence and projections.
The copulatory piece (Figs. 13 F, 13 G) is very various according to species and important in
discriminating allied species. It is not exposed when the inner sac is evaginated (Figs. 14 D,
14 E), and its dorsal part is covered with pubescence or spines.
Paramere. The paramere of aedeagus in Pella species (Figs. 15 A–15 D) is generalised instructure as in most aleocharines, in which it is attached to the median lobe as shown in Figs.
15 C, 15 D and composed of the condylite, paramerite and velum. The paramerite bears an apical22

M. Maruyama

Fig. 13. Pella limbata (Paykull). — A, Median lobe of aedeagus, dorsal view; B, ditto, ventral view; C, ditto, lateral view; D, ditto, distal crest, dorsal view; E, ditto, lateral view; F, copulatory piece, lateral view; G, ditto,
dorsal view.

lobe, with 4 setae around the apex. The apical lobe in Pella species is relatively long, exceeding
the apex of the velum.
Female genitalia
Vagina. The vagina of Pella species (Fig. 16 A) is generalised in structure as in most aleocharines. The vaginal plate is weakly sclerotized, and its size is various probably in accordancewith the size of the inner sac of aedeagus. The spermathecal duct is attached at the dorsal surfaceof the end of the vagina.

Palearctic Species of Pella

23

Fig. 14. Pella limbata (Paykull). — A, Apical lobe of median lobe of aedeagus, dorsal view; B, ditto, lateral view;C, ditto, around apex, ventral view; D, evaginated inner sac of median lobe, ventral view; E, ditto, lateral view.

Spermatheca. The spermatheca in Pella species (Figs. 16 A, 16 D–16 F) is simplified andmainly composed of two parts: basal part and apical part. The basal part is attached to the spermathecal duct at its base, and sometimes its inner wall is wrinkled. The apical part is attached tothe spermathecal gland and more or less notched at its connection with the latter. The inner wallis sparsely to densely wrinkled.
The general shape of the spermatheca is various by species and very important in diagnosing
species.
Although the spermatheca in most lomechusines are generalised as in Pella, that in thespecies of Zyras (Zyras), Diplopleurus and Tetrabothrus are coiled many times.

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M. Maruyama

Fig. 15. Pella limbata (Paykull). — A, Paramere of aedeagus, lateral view; B, ditto, far side; C, position of paramere on median lobe, ventral view; D, ditto, lateral view.

Palearctic Species of Pella

Fig. 16. Pella limbata (Paykull). — A, Ninth and 10th tergites, dorsal view, and spermatheca; B, 9th tergite andsternite, ventral view; C, 9th sternite, apical part, ventral view; D–F, spermathecae. A–C, female abdominal
segments. Abbreviations: T, tergite; S, sternite.

25

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M. Maruyama

CHAPTER 2. TAXONOMY
The identities of several Pella species are ambiguous or largely confused. For example, P.
laeviceps (Eppelsheim), P. barbara (Fairmaire), P. excepta (Mulsant & Rey) and P. maura (Fauvel), all described from the western Palearctic region, have been neither recorded nor redescribedsince the original descriptions published more than a century ago. Judging from the original descriptions, the affiliation of several species described from southern Asia and Caucasia are apparently doubtful. Faunistic studies of Pella have well been done only in a few regions, e.g., Central
Europe (Ganglbauer, 1895; Reitter, 1909; Lohse, 1974, 1989, etc.) and the Russian Far East
(Dvořák, 1981). The present study also deals with material from poorly investigated areas in the
Palearctic region, namely South Europe, North Africa, Turkey, Caucasia and East Asia. I havediscovered 15 undescribed species from these areas.
In this chapter I will revise the Palearctic species of Pella taxonomically. Recently somespecies of the genus have been found in the Nearctic region, and they are separately revised by
Klimaszewski et al. (2005). Detailed descriptions of these undescribed species and redescriptionsof all the named species including above-mentioned ambiguous species are given. Because reliable identification to Pella species is important to their ecological and behavioural studies ofmyrmecophily, this study would contribute much to biological study of Pella.
Brief History of Taxonomic Study of Pella
In this paper, I synonymize Myrmelia Mulsant & Rey, 1873b; Pellochromonia Reitter, 1909;
and Lepla Tottenham, 1939, with Pella, which are formerly arranged as the subgenera of thegenus Zyras Stephens, 1835.
The genus Pella was erected by Stephens (1835) to accommodate four British species of thegenus Aleochara without designation of the type species. Later, Westwood (1838) designatedStaphylinus limbatus Paykull, 1789, as the type species of Pella, and added two species to thegenus, which had been arranged as members of Aleochara by Stephens (1835).
Erichson (1837) established the genus Myrmedonia to accommodate several Aleochara and
Bolitochara species without designation of the type species. Duponchel (1841) designatedStaphylinus canaliculatus (Drusilla canaliculata) as the type species of Myrmedonia. However,Thomson (1859) was unconscious of Duponchel’s (1841) work and designated Aleochara humeralis (P. humeralis) as the type species of Myrmedonia. After that, under the influence of Thomson (1859), many species of Pella were described under Myrmedonia in the 1860’s to the 1920’s.
Myrmedonia is actually a junior synonym of Drusilla Leach, 1819.
Mulsant and Rey (1873b) established the subgenus Myrmelia for Myrmedonia excepta Mulsant & Rey, 1861.
Reitter (1909) regarded Pella as a subgenus of the genus Myrmedonia and established a newsubgenus Pellochromonia for Myrmedonia ruficollis Grimm, 1845.
Fenyes (1920) regarded Pella as a subgenus of the genus Zyras Stephens, 1835, together with
Myrmedonia (sensu Mulsant & Rey, 1873a), Pellochromonia and Myrmelia and referred several
species to each of them, which were described merely under Myrmedonia. Bernhauer andScheerpeltz (1926) and Scheerpeltz (1934) in their catalogue almost followed Fenyes’ (1920) list.
Tottenham (1939) established Lepla for the species, that had been included in Myrmedonia(sensu Mulsant & Rey, 1873a), and designated Myrmedonia lugens Gravenhorst, 1806 as the typespecies.

Palearctic Species of Pella

27

Kistner (1972) again raised Pella to the generic rank. However, current works (Lohse, 1974,
1989; Likovský, 1993; Smetana, 2004, etc.) did not follow Kistner’s view and regarded Pella as asubgenus of Zyras. Lohse (1972, 1989) is one of the most important identification guides of the
Aleocharinae, and many researchers, not only in Central Europe but also in other countries of the
Palearctic region, follow his arrangement.
Dvořák (1981) revised the Pella species of East Asia, but he misidentified some species andoverlooked one species.
Gürlich (1981) recognised Pella as a distinct genus on the basis of the examinations of thechaetotaxy of abdominal macrosetae. However, he examined only the type species of the genus,
and the chaetotaxy of abdominal macrosetae is highly homoplastic among lomechusine genera.
Descriptions
Tribe Lomechusini Fleming, 1821
Lomechusidae Fleming, 1821: 49 [type genus: Lomechusa Gravenhorst, 1806].
Myrmedoniini Ganglbauer, 1895: 196 [type genus: Myrmedonia Erichson, 1837].—Seevers, 1978: 151.

Diagnosis. The Lomechusini has not been well defined, but it can be distinguished from theother tribes of Aleocharinae by a combination of the following character states: 1) galea elongate; 2) metasternal process longer than mesosternal process; 3) tarsal formula: 4–5–5; 4) ventral
bridge of aedeagal median lobe present; 5) compressor plate of aedeagal median lobe covered bya plate, which extends from the bottom of the basal capsule.
Subtribe Myrmedoniina Thomson, 1867
Myrmedoniides Thomson, 1867: 209 [type genus: Myrmedonia Erichson, 1837].
Myrmedoniia Sharp, 1883: 170 [type genus: Myrmedonia Erichson, 1837].
Myrmedoniates Mulsant & Rey, 1873a: 34 [type genus: Myrmedonia Erichson, 1837].
Zyrini Bradley, 1930: 83 [type genus: Zyras Stephens, 1835].
Zyrasini Jeannel & Jarrige, 1949: 304 [type genus: Zyras Stephens, 1835].
Bolitocharinae Hatch, 1957: 146 [type genus: “Bolitochara” Westwood, 1838].
Myrmedoniinea Seevers, 1978: 18 [type genus: Myrmedonia Erichson, 1837].

Diagnosis. The Myrmedoniina has not yet been well defined and is estimated as a paraphyletic group (Maruyama, in prep.). However, it is distinguishable from the other subtribe of the
Lomechusini, the Lomechusina, by the following character states: 1) galea elongate; 2) ligula evidently bifid; 3) trichome absent on 3rd to 5th abdominal segments.
Genus Pella Stephens, 1835
Pella Stephens, 1835: 434 (description, genus) [type species: Staphylinus limbatus Paykull, 1789, fixed by Westwood(1838)].—Ganglbauer, 1895: 116 (subgenus of “Myrmedonia”, list).—Reitter, 1909: 42 (subgenus of“Myrmedonia”, description). — Fenyes, 1920: 296 (subgenus of Zyras, description).—Bernhauer & Scheerpeltz,
1926: 694 (subgenus of Zyras, list). — Kistner, 1972: 150 (distinct genus, description).—Lohse, 1974: 224 (subgenus of Zyras, description). — Gürlich, 1981: 211–212 (distinct genus, chaetotaxy).
“Myrmedonia”: Thomson, 1859: 255 (description). — Mulsant & Rey, 1873a: 50 (description).—Ganglbauer, 1895: 116(description).— Reitter, 1909: 43 (key). — Fenyes, 1920: 297 (subgenus of Zyras, description).—Bernhauer &Scheerpeltz, 1926: 694 (subgenus of Zyras, list).
Myrmelia Mulsant & Rey, 1873b: 152 (subgenus of “Myrmedonia”, description) [type species: Myrmedonia excepta
Mulsant & Rey, 1861, by monotypy]. — Ganglbauer, 1895: 116 (subgenus of “Myrmedonia”, list).—Fenyes, 1920:

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M. Maruyama

299 (subgenus of Zyras, description). — Bernhauer & Scheerpeltz, 1926: 694 (subgenus of Zyras, list). N. syn.
Pellochromonia Reitter, 1909: 43 (subgenus of “Myrmedonia”, description) [type species: Myrmedonia ruficollis
Grimm,1845, by monotypy]. — Fenyes, 1920: 298 (subgenus of Zyras, description).—Bernhauer & Scheerpeltz,
1926: 694 (subgenus of Zyras, list). — Lohse, 1974: 226 (subgenus of Zyras, description). N. syn.
Lepla Tottenham, 1939: 226 (new name for “Myrmedonia” sensu Mulsant & Rey, 1874) [type species: Aleochara lugens
Gravenhorst, 1802, fixed by Tottenham, 1939]. — Gürlich, 1981: 211 (distinct genus, chaetotaxy). N. syn.

Distribution. Holoarctic region (except for tundra, desert or non-forest areas).
Diagnosis. The following character states are considered to be basal autapomorphies of thegenus Pella (Maruyama, in prep.): 1) ligula without setae but with sensilla; 2) 2nd segment oflabial palpus almost the same as the 1st in width. In addition, Pella can be distinguished from theother genera of the tribe Lomechusini by a combination of the following character states: 1) body
3.4–7.0 mm in length; 2) head almost circular in dorsal view; 3) head without “neck”; 4) headwith occipital suture; 5) eyes 0.35–0.56 times as long as head width; 6) antenna generalised, butslightly depressed; 7) pronotum and elytra densely or moderately covered with setae; 8) pronotal
and elytral surfaces smooth or with fine reticulations, without granulation or rugose punctulation;9) hind wings entire; 10) paratergite generalised, without projection; 11) 10th tergite densely covered with setae except for mesal area; 12) aedeagus simplified, pear-shaped or oval in ventral
view.
Description. Body (Fig. 1). Slender or somewhat robust, subparallel-sided. Body length:
3.4–7.0 mm; fore body: 1.6–3.9 mm. Ground colour pale brown to black, or reddish orange.
Head. Head capsule (Figs. 2 A, 2 B, 3 A) almost circular. Occipital suture (Figs. 2 A, 2 B,
3 A) present, dorsally crossing hind part of head, continuing onto ventral side, and terminating oneach side near posterior articulation of hypostoma; surface covered with setae. Eyes (Figs. 2 A,
2 B) oval in lateral view, more or less prominent, 0.35–0.56 times as long as head width; small
setae present among facets. Antennae (Figs. 3 B–3 D) generalised in shape, almost as long ashead and pronotum combined, more or less flattened dorsoventrally; setae on surface becoming
denser and smaller apicad; 4th to 10th segments slightly dilated apicad or subparallel-sided, wellmargined postero-laterally; each segment with erect black setae, those on 3rd to 10th segments
forming a row around apical rim; 11th segment oval to elongate oval.
Mouthparts. Labrum (Figs. 4 A, 4 B) much wider than long (W/L, 1.8–2.1), truncate orslightly emarginate antero-medially, submembranous around antero-medial area; surface covered
with 100–130 pseudopores except on posterior and lateral areas; apodeme roundly projected postero-medially, with lateral arm gently curved. Epipharynx (Fig. 4 B); medial sensory field with
80–90 pores and 50–80 grain-like sensory pores laterally; each mesolateral area irregularly with
10–15 pores among round projections; 5 pairs of lateral setae stout, about five times as long as
wide. Mandibles somewhat asymmetric; right mandible (Figs. 5 A, 5 B) with a small tooth (Fig.
5 A) at middle of inner margin; 2 or 3 small setae present laterally; mesal areas of dorsal and ventral surfaces covered with numerous pseudopores; prostheca obtuse at apex, its inner margindensely pubescent. Maxilla (Fig. 5 C): cardo generalised, almost elliptical in ventral view, with
40–50 pseudopores. Stipes small, triangular, with 2 setae at base, without pseudopores. Palpifer
triangular in lateral view, with a large seta, several small setae and 15–20 pseudopores near apex.
Lacinia with 50–60 pores and 5 or 6 setae medially; apical inner margin with a row of setae
forming a comb; surface of around comb densely pubescent; subgalea small, much smaller thanlast segment of maxillary palpus; galea long, about six times as long as wide, slightly curved at
middle, with some pores near apex, densely pubescent apically. Maxillary palpus without
pseudopores; 1st segment very small, with 1 seta; 2nd segment gently curved, and much dilated

Palearctic Species of Pella

29

apically; 3rd segment longest, slightly longer than 2nd, almost straight; 4th segment small, conical. Mentum (Fig. 5 D) trapeziform; posterior and anterior margins truncate; around antero-lateral corner with a long seta; surface densely covered with pseudopores, with several setae laterally.
Labium (Figs. 3 A, 6 A–6 E): prementum (Fig. 6 A) with 2 real pores and 1 setal pore mesolaterally, and 40–50 medial pseudopores, which are relatively large and dense around the medial
setae; internal ridge (Fig. 6 D) very small, its length less than 1/8 that of prementum; apodeme(Fig. 6 A) with postero-medial margin roundly emarginate, and its lateral part acutely projected(Fig. 6 A: arrow); lateral lobe of apodeme (Fig. 6 A) gently curved, pointed apically. Ligula (Figs.
6 A, 6 C) bilobed and each lobe rounded apically; apical setae absent, but 3 sockets present. Labial palpus (Figs. 6 A, 6 B) with 1st segment shortened and as long as 2nd; 2nd segment notchedfrom base to apex, forming a membranous zone. Hypopharynx (Fig. 6 E) without apical seta; medial sensory field with small ridges around apex and scale-like sculptures near base.
Thorax. Pronotum (Figs. 1, 7 A) subtrapeziform, subquadrate or elliptical, much wider thanlong (PW/PL, 1.21–1.60); disc well margined, somewhat convex but more or less flattened above;
surface moderately to densely covered with setae uniformly. Cervical sclerites (Fig. 7 A) generalised, rectangular in lateral and ventral view. Basisternum  preepisternum (Fig. 7 A) generalised, but lacking transverse suture. Scutellum (Fig. 8 A) generalised, its apex somewhat rounded. Anepisternum and epimeron (Figs. 7 D, 7 E) of metasternum completely fused; anepisternum
narrowed posteriad; epimeron subtriangular, widened posteriad. Metanotum (Fig. 8 B) weaklysclerotized; scutoscutellar suture with a pubescent patch along inner margin; transscutal suture
obscure; scutoscutella generalised, triangular; scutelopostnotal suture almost straight. Mesosternum (Fig. 7 B) with process short, slightly projected posteriad, rounded at apex. Metaepimeron(Fig. 7 B) small, semicircular. Metasternum (Fig. 7 B) large, three times longer than metasternum; process well developed, narrowed apicad, its apex somewhat rounded. Metendosternite(Fig. 7 C) generalised; basal stalk convex dorsad, angled; furcal arm gently curved ventrad. Elytra(Figs. 8 C, 8 D) without epiplural suture, more or less margined from shoulder through inner margin to posterior margin; surface almost uniformly covered with setae, which are slightly denser
apicad. Hind wing (Figs. 8 E, 8 F) with its venation highly obscure; ScR, R1, CuPcu and
A1A2 present, very weakly sclerotized; surface uniformly and moderately pubescent, withminute pubescence along posterior margin; flabellum (Fig. 8 F) with about 20 processes.
Legs. General shape as in Figs. 9 A–9 C. Coxae: fore coxa well developed, slightly shorter
than fore femur, ventrally with a suture; mid coxa generalised, with a short suture ventrally; hindcoxa subtriangular, rounded laterally. Trochanters: fore and mid trochanters small, subconical;
hind trochanter rounded at apex. Femora stout, flattened, gently curved near apex. Tibiae dilated
apicad from base to around apical 1/3; each apical margin with a ctenidium and 2 stout setae.
Tarsi generalised; empodium with a pair of setae.
Abdomen. First segment (Fig. 8 B): tergite weakly sclerotized, with anterior margin deeply
emarginate. Second segment (Fig. 10 A): tergite with posterior margin weakly emarginate, its
postero-lateral corner rounded. Third to 6th segments (Figs. 10 A, 10 B): tergites with posterior
margins almost straight; dorsolateral plates rectangular except that of 6th segment evidently narrowed posteriad; sternite with surface moderately to densely covered with setae. Seventh segment(Figs. 10 A, 10 B): tergite with a pair of gland openings at base; lateral plates fused, narrowed
posteriad, pointed at apex. Eighth abdominal segment (Figs. 10 A, 10 B, 11A–11 H): tergite withbasal suture curved laterally and continuing apicad or weakened around lateral sides; sternite
with basal suture reaching lateral sides or weakened around lateral area. Ninth abdominal segment (Figs. 12 A, 16 A): tergite with surface densely covered with setae. Tenth abdominal seg-

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M. Maruyama

ment (Figs. 12 A, 16 A) with surface densely covered with setae except in mesal area; setae becoming sparser and longer apicad.
Male characters. Head slightly depressed above; setae on dorsal surface denser and longer
than in female. Eighth tergite (Figs. 11 A, 11 E) with basal projection of apodeme (Fig. 11 E)
more or less larger than in female; posterior margin (Fig. 11 A) dentate or crenate. Eighth sternite(Figs. 11 B, 11 G) larger than in female; macrosetae more numerous than in female; apical sensory setae (Fig. 11 G) smaller than in female; apical margin lacking minute pubescence. Lateralprojections of apodeme of 9th tergite (Fig. 12 A) asymmetrical; right projection longer than theleft. Ninth sternite (Figs. 12 B, 12 C) oblong oval; apical margin more or less truncate; surface
covered with setae, with a pair of macrosetae near posterior margin; left lateral margin attaching
to tergite (Fig. 12 D). Aedeagus (Figs. 13 A–13 G, 14 A–14 E, 15 A–15 D): median lobe (Figs.
12 A–12 E, 14 A–14 E, 15 C, 15 D) with basal capsule bulbous; distal crests (Figs. 13 D, 13 E)
fused on their sides, more or less projected; copulatory piece (Figs. 13 F, 13 G) small to mediumsized, pubescent or dentate dorsally, not exposed when inner sac evaginated; paramere (Figs.
15 A–15 D) with condylite almost straight; hinge zone distinct; apical lobe of paramerite short,sometimes exceeding apex of velum; velum densely ridged.
Female characters. Head slightly depressed above, less depressed than in male; setae on
dorsal surface sparser and shorter than in male. Eighth tergite (Figs. 11 C, 11 F) with lateral projection of apodeme (Fig. 11 F) smaller than in male; posterior margin (Fig. 11 C) crenate but
weaker than in male and its crenate margin narrower than in male. Eighth sternite (Figs. 11 D,
11 H) smaller than in male; macrosetae less numerous than in male; apical sensory setae (Fig.
11 H) larger than in male, sometimes spindle-shaped and flattened; apical margin (Fig. 11 H)
sometimes margined with minute pubescence. Ninth sternite (Figs. 16 B, 16 C) with posterior
margin truncate; surface around posterior margin covered with setae. Vagina (Fig. 16 A) withvaginal plate weakly sclerotized. Spermatheca (Figs. 16 A, 16 D–16 F) large, as long as or longer
than 10th tergite, clearly divided into basal and apical parts; spermathecal duct (Fig. 16 A) short,
3–5 times as long as spermatheca; spermathecal gland (Fig. 16 A) attaching to about middle ofapical part.
Comments. Pella has been regarded as a subgenus of Zyras by authors but is recognised asa good genus in this study. Pella is a monophyletic group and far related to Zyras (Zyras) as a result of phylogenetic analysis of the Lomechusini (Maruyama, in prep.).
I herewith synonymize Myrmelia, Pellochromonia and Lepla with Pella, which are also previously regarded as subgenera of Zyras. This view is based on the adult external structure of all
the known species of these genera/subgenera, there having been found no important difference ofgeneric or subgeneric value.
As a result of type material examinations, it has been found that several species formerly regarded as members of Pella (or Lepla) cannot be affiliated to the genus. They are: Zyras (Pella?)coloratus Cameron, 1939; Z. (P.) ceylonicus Cameron, 1939; Z. (P.) urartu Iablokoff-Khnzoryan,
1962; Z. (L.) iridescens (Sawada, 1970b); Z. (P.) wrasei Dvořák, 1988; Z. (P.) quasar Dvořák,
1996; and Z. (P.) reelsi Pace, 1998a. These species are transferred to other genera (or tribe) andredescribed at the foot of this chapter.
Check List of the Genus Pella
The limbata group
Pella limbata (Paykull, 1789)

Palearctic Species of Pella
Staphylinus divisus Marsham, 1802
Aleochara laevis Gravenhorst, 1802

P. horii Maruyama, n. sp.
The similis groupP. similis (Märkel, 1845), n. comb.
The humeralis group
P. humeralis (Gravenhorst, 1802), n. comb.
P. laeviceps (Eppelsheim, 1880), n. comb.
The funesta group
P. funesta (Gravenhorst, 1806), n. comb.
Aleochara crassicornis Stephens, 1832
Myrmedonia atrata Heer, 1839

P. comes (Sharp, 1874)
P. socia (Sharp, 1874)
P. rambouseki (Bernhauer, 1929), n. comb.
P. jureceki (Dvořák, 1981), n. comb.
Zyras (Pella) micropterus Pace, 1998a, n. syn.

P. kidaorum Maruyama, n. sp.
P. cooterorum Maruyama, n. sp.
The barbara groupP. barbara (Fairmaire, 1863), n. comb.
P. leonhardi (Bernhauer, 1912), n. comb.
P. jelineki Maruyama, n. sp.
The erratica groupP. erratica (Hagens, 1863), n. comb.
Myrmedonia mustela Rottenberg, 1870
Myrmedonia Ehlersi Eppelsheim, 1884b

The schillhammeri groupP. schillhammeri Maruyama, n. sp.
The kinomurai group
P. kinomurai Maruyama, n. sp.
P. primorskyiana Maruyama, n. sp.
The hlavaci groupP. hlavaci Maruyama, n. sp.
The cognata groupP. cognata (Märkel, 1842), n. comb.
P. iberica Maruyama, n. sp.
P. japonica (Sharp, 1888)
P. kishimotoi Maruyama, n. sp.
The ruficollis group
P. ruficollis (Grimm, 1845), n. comb.
Myrmedonia fernandi Fairmaire, 1855

The coreana groupP. coreana Maruyama, n. sp.
P. plutenkoi Maruyama, n. sp.
The excepta group
P. excepta (Mulsant & Rey, 1861), n. comb.

31

32

M. Maruyama

P. maura (Fauvel, 1898), n. comb.
Myrmedonia Ragusae Ragusa, 1921

P. kuluensis (Cameron, 1939), n. comb.
P. bohaci (Dvořák, 1984), n. comb.
Zyras (Pella) almaatensis Pace, 2002, n. syn.

P. cinctipennis (Eppelsheim, 1884), n. comb.
Zyras (Pella) esau Dvořák, 1984, n. syn.

The lugens group
P. lugens (Gravenhorst, 1802), n. comb.
P. beijingorum (Pace, 1998), n. comb.
P. intermedia Maruyama, n. sp.
P. masakoae Maruyama, n. sp.
The spreta groupP. spreta (Sharp, 1888), n. comb.
P. zhoui Maruyama, n. sp.
The laticollis groupP. laticollis (Märkel, 1845), n. comb.
P. hampei (Kraatz, 1862), n. comb.
P. indiscreta (Sharp, 1888), n. comb.
Incertae sedis
P. pumila (Fiori, 1914), n. comb.

Key to the Species Groups of the Genus Pella
1. Eleventh antennal segment long, longer than 7th to 10th segments combined. .................. 2.
— Eleventh antennal segment short to medium, shorter than 7th to 10th segments combined. ...
.............................................................................................................................................. 3.
2. Body bicoloured; pronotum reddish orange; pronotal hypomeron fully visible in lateralview; posterior margin of 8th tergite not crenate nor dentate, almost straight; macrosetae on
8th abdominal segment clearly differentiated from setae. ..............................ruficollis group.
— Body unicoloured, pale brown; pronotal hypomeron partly visible in lateral view, its visible
part less than 3/5 of pronotal length; posterior margin of 8th tergite crenate; macrosetae on
8th abdominal segment clearly differentiated from setae. ...............................coreana group.
3. Pronotal hypomeron invisible, or only slightly and narrowly visible in lateral view (its visible length less than 0.4 times as long as pronotum). ........................................................... 4.
— Pronotal hypomeron clearly visible in lateral view (its visible length more than 0.5 times aslong as pronotum). .............................................................................................................. 6.
4. Body almost unicoloured. ...............................................................................laticollis group.
— Body evidently tricoloured; black in ground colour; elytra yellow but blackish brownaround postero-lateral corner, or black but with yellow; legs reddish brown. .................... 5.
5. Eyes large and prominent, its length about 0.45–0.48 times as long as head width; lateralareas of pronotum well convex above; pronotal surface roughly punctured. ......spreta group.
— Eyes medium to large in size, its length less than 0.42 times as long as head width; lateralareas of pronotum flattened; pronotal surface finely punctured. ........................lugens group.
6. Antennae and legs yellow. ......................................................................schillhammeri group.
— Antennae and legs reddish brown to black. ......................................................................... 7.

Palearctic Species of Pella

33

7. Pronotum with postero-lateral corners, evidently narrowed posteriad, more or less parallelsided posteriorly. ................................................................................................................. 8.
— Pronotum completely rounded postero-laterally to posteriorly, not forming postero-lateralcorners. ................................................................................................................................10.
8. Elytra bicoloured, with a maculation at antero-lateral margin; posterior margin of male 8thsternite truncate; copulatory piece of aedeagal median lobe widened apicad and widely
truncate at apex. ............................................................................................humeralis group.
— Elytra unicoloured; posterior margin of male 8th sternite rounded; copulatory piece of
aedeagal median lobe narrowed apicad and not truncate at apex. ...................................... 9.
9. Body medium to large: 5.0–6.6 mm. .................................................................funesta group.
— Body small: 3.3–4.1 mm. .......................................................................barbara group (part).
10. Pronotum widest around middle, more than 1.48 times as wide as long . ........hlavaci group.
Pronotum widest anteriorly, less than 1.46 times as wide as long. .....................................11.
11. Body small: 3.3–4.1 mm, almost unicoloured. ....................................................................12.
— Body medium to large: 3.7–6.2 mm, bicoloured. ................................................................13.
12. Eleventh antennal segment almost as long as 1st segment. ...................barbara group (part).
— Eleventh antennal segment evidently shorter than 1st segment. .......................erratica group.
13. Antero-lateral areas of pronotum strongly curved ventrad; hypomeron partly visible in lateral view, its visible length less than 0.6 times as long as pronotum; pronotal surface sparsely
to moderately covered with setae; distal crest of aedeagal median lobe roundly projected. ....
...........................................................................................................................excepta group.
— Antero-lateral areas of pronotum weakly curved ventrad; hypomeron fully visible in lateralview, its visible length more than 0.7 times as long as pronotum; pronotal surface densely
covered with setae; distal crest of aedeagal median lobe gently projected, or weakly projected. ........................................................................................................................................14.
14. Male 8th sternite almost as long as that of female; apical lobe of aedeagal median lobe
much narrowed in ventral view and acutely curved ventrad at base; copulatory piece of
aedeagal median lobe aciculate around apex in ventral view. ..........................limbata group.
— Male 8th sternite evidently longer than that of female; apical lobe of aedeagal median lobe
gradually narrowed apicad and slightly curved ventrad or almost straight; copulatory pieceof aedeagal median lobe rounded, truncated or pointed at apex and not aciculate in ventral
view. ....................................................................................................................................15.
15. Legs almost uniformly reddish brown; abdomen with around posterior margin of 3rd to 5th
abdominal segments reddish brown. ............................................................kinomurai group.
— Legs reddish yellow to yellowish brown, around apical 1/2 of femora darker; abdomen withmost areas of 3rd to 5th abdominal segments yellowish brown. .........................................16.
16. Eleventh segment of antenna almost as long as 1st segment; apical lobe of aedeagal median
lobe much shorter than basal capsule, gently narrowed apicad and pointed at apex in lateralview; apical part of spermatheca without outer projection. ...............................similis group.
— Eleventh segment of antenna much shorter than 1st segment; apical lobe of aedeagal median lobe almost as long as basal capsule, subparallel-sided around middle and obtuse at apex
in lateral view; apical part of spermatheca with outer projection. ...................cognata group.
Comments. In distinguishing the species of Pella examinations of aedeagus or spermatheca are often indispensable, because most other characters used in identification are rather quantitative. It is better to refer first to the figures of aedeagus or spermatheca.

34

M. Maruyama

The limbata GroupSpecies included. Pella limbata, P. horii.
Distribution. European subregion, Mediterranean subregion, Manchurian subregion.
Diagnosis. Species of the limbata group may be characterised by a combination of the following character states: 1) eye length 0.32–0.34 times as long as head width; 2) 11th antennal
segment longer than the 1st; 3) pronotum with postero-lateral corners; 4) pronotum 1.18–1.35
times as wide as long; 5) pronotum widest anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytra with yellowish brown maculations around antero-lateral and postero-lateralcorners; 8) inner and posterior margins of elytra weakly margined; 9) male 8th sternite not longer
than in female and almost the same in size; 10) posterior margin of 8th tergite crenate; 11) posterior margin of male 8th sternite rounded; 12) lateral projection of apodeme of male 8th tergite
not evidently longer than that of female; 13) macrosetae of 8th abdominal segment generalised inlength, exceeding the posterior margin of the segment; 14) postero-medial margin of female 8thsternite with minute pubescence; 15) apical lobe of aedeagal median lobe curved ventrad at basein lateral view; 16) distal crests of aedeagus well developed, projected semicircularly in lateralview; 17) copulatory piece of aedeagal median lobe aciculate around apex in dorsal view.
Comments. The limbata group can be regarded as a monophyletic group by character
states: the apical lobe of the aedeagal median lobe is curved ventrad at the base in lateral view; itis much narrowed at base in ventral view; the copulatory piece of the aedeagal median lobe isaciculate around the apex in dorsal view. These states are unique within the genus and not observed in the other species of the Lomechusini. Thus, these states are considered to be autapomorphies of the species-group. This species-group may possibly be allied to the similis groupand the cognata group in resemblance of the shape of male 8th sternite, which is relatively widely emarginate posteriorly.
Symbiotic hosts. Lasius (Lasius) spp., L. (Cautolasius) flavus, L. (Dendrolasius) fuliginosus.
Key to the Species of the limbata Group——

Eyes smaller, 0.32–0.33 times as long as head width; pronotum wider, 1.28–1.35 times aswide as long; posterior margin of male 8th tergite deeply and roundly emarginate, and its
lateral corner projected; apical lobe of aedeagal median lobe shorter than apical part. Distribution: Europe. ........................................................................................................P. limbata.
Eyes larger, 0.33–0.34 times as long as head width; pronotum narrower, 1.18–1.25 times aswide as long; posterior margin of male 8th tergite weakly emarginate, and its lateral cornerslightly rounded; apical lobe of aedeagal median lobe longer than apical part. Distribution:
Asia. .............................................................................................................................P. horii.
Pella limbata (Paykull, 1789)(Figs. 1–17)

Staphylinus limbatus Paykull, 1789: 54 (original description).—Stephens, 1832: 162 (Aleochara, description).—Erichson, 1837: 288 (Myrmedonia, description). — Mulsant & Rey, 1873a: 55 (Myrmedonia, description).—des Gozis,
1886: 12 (Pella, list). — Ganglbauer, 1895: 123 (Myrmedonia (Pella), key, description).—Fenyes, 1918: 24 (Pella,list); Reitter, 1909: 43 (Myrmedonia (Pella), key). — Fenyes, 1920: 297 (Zyras (Pella), list).—Bernhauer & Scheerpeltz, 1926: 701 (Zyras (Pella), list). — Scheerpeltz, 1934: 1659 (Zyras (Pella), list).—Kistner, 1972: 150 (Pella, description).—Lohse, 1974: 225 (Zyras (Pella), key). — Gürlich, 1981: 211 (Pella, chaetotaxy).—Likovský, 1993: 59

Palearctic Species of Pella

Fig. 17.

35

Pella limbata (Paykull). — A, Fore body, dorsal view; B, right antenna.

(Zyras (Pella), list). — Smetana, 2004: 461 (Zyras (Pella), list).
Staphylinus divisus Marsham, 1802: 510 (original description).—Stephens, 1832: 162 (Aleochara, list).—Fenyes, 1920:
297 (synonym of limbata, list). — Bernhauer & Scheerpeltz, 1926: 701 (synonym of limbata, list).—Smetana, 2004:
461 (synonym of limbata, list).
Aleochara laevis Gravenhorst, 1802: 95 (original description).—Mulsant & Rey, 1873a: 55 (synonym of limbata, list).—des Gozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 123 (synonym of limbata, list). — Fenyes, 1920: 297 (synonym of limbata, list). — Bernhauer & Scheerpeltz, 1926: 701 (synonym of limbata, list).—Smetana, 2004: 461(synonym of limbata, list).

Type material. Staphylinus limbatus. Syntypes: Sweden: 1?, 2// , (without label, pinned)(NHRS).
Staphylinus divisus. Not examined. Type locality: Great Britain.
Aleochara laevis. Syntype: Germany: 1/, “/ laevis Gr. [“Berolin” (Berlin, Germany)]”(pinned) (ZMHB).
Type locality. Sweden.
Other material. Austria: 1 ex., Wien, Breit; 2 exs., Donauauen, Breit. Czech Republic: 4exs., Ostrava, Moravia, 11 IV 1980, L. Klíma; 1 ex., Chuchle, Bohemia, 10 IV 1944, Souček; 1ex., Cějč, Moravia, 9 V 1988, Ju° ta; 1 ex., same data but, 1 VI 1988; 1 ex., Sárka, Praha, Roubal;
5 exs., Vysoká, Bohemia, 26 III 1950, A. Smetana (CNC); 1 ex., Hradao, Bohemia, 19 III 1949,
A. Smetana (CNC); 1 ex., Hlvbočepy, 23 VI 1935, Kul (CNC); 1 ex., Bohemia, Celákovice, 21
III 1945, J. Král; 3 exs., Chuchle, Bohemia, 10 IV 1944, Souček (CNC); 1 ex., Hradec, Bohemia,
4 VIII 1944, Souček (CNC); 1 ex., same data but, 8 IV 1944, Souček (CNC); 2 exs., same databut, 9 IV 1944, Souček (CNC); 2 exs., Kukleny, Bohemia, 23 IV 1947, A. Smetana (CNC); 2

36

M. Maruyamaexs., same data but, 8 V 1947, A. Smetana (CNC); 3 exs., Silesia, Radvanice, 28 V 1963, P.
Nohel (CNC). Germany: 5 exs., Schneverdingen, Soltau, Luneburger, Niedersachsen, V 2001, V.
Assing (pt); 1 ex., Von Heidelberg, W. Heinz, Wilhelmsfeld, 25 IV 1963; 1 ex., Umg. Halberstadt, 7 V 1982, H. D. Briagmann; 1 ex., Berlin–Treptow, Sdl. Daheim, Ruderalflur, BF, 16
IV–22 V 1994, ABT 3, Uhlig; 1 ex., Baden–Württemberg, Kaiserstuhl mts., NW Freiburg, VII
1981, Kobel–Lamparski (unbaited pt); 1 ex., Mesobrometum, Steinberg, S Hildesheim, D.
Niedersachsen, 1 V 1997, Bodenfalle; 2 exs., northern Thuringia, 2 V 1917, Petry (LCAF); 1 ex.,same data but, 3 V 1908 (LLB). Slovakia: 1 ex., Boï-naplav Latonic, 11 III 1995, P. Hlaváč; 1ex., Streda n. B., 8 V 1989, M. Kuboň; 4 exs., Ku° ty, 27 III 1992, M. Kuboň; 1 ex., Mitra, VI
1930, Roubal (CNC).
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Belgium,
Bosnia Herzegovina, Czech Republic, Denmark, Estonia, Finland, France, Great Britain, Germany, Greece, Hungary, Ireland, Italy, Latvia, Lithuania, Luxembourg, Netherlands, Norway,
Poland, Russia (North, Central and South European territories), Slovakia, Spain, Sweden,
Switzerland.
Diagnosis. Pella limbata is closely similar to P. horii in general appearance and colour, buteasily distinguished from the latter by the emargination on posterior margin of male 8th tergite
much deeper and the presence of a depression on male pronotal disc. These two species are locally isolated. Among the sympatric species, Pella limbata is similar to P. similis and P. cognata ingeneral appearance, but in addition to the diagnostic characters above distinguished from P. similis by the larger body, and from P. cognata by the apical lobe of aedeagal median lobe gradually
narrowed apicad.
Redescription. Body (Fig. 1) slender. Brown in ground colour; antennae, mouthparts, legs,
2nd to 4th abdominal segments and around posterior margins of 5th to 7th abdominal segments
reddish yellow; elytron with yellow maculations around antero-lateral and postero-lateral corner;
rarely elytra and abdomen uniformly brown. Head (Figs. 2 A, 2 B, 3 A, 17 A) widest just behind
eyes; surface smooth, moderately covered with setae; setae rather long, almost as long as thoseon pronotum and elytra; length of eyes 0.32–0.33 times as long as head width. Antennae (Figs.
3 C, 3 D, 17 B) shorter than head, pronotum and elytra combined; 1st segment long, almost as
long as 2nd and 3rd combined; 2nd segment about 2/3 as long as 3rd; 3rd segment about 2/3 aslong as 1st; 4th and 5th segments almost as long as wide; 6th to 10th segments wider than long;
11th segment slightly longer than 1st; approximate relative lengths of segments from basal to apical: 30 : 13 : 18 : 7 : 7 : 7 : 7 : 6 : 7.5 : 8.5 : 32. Pronotum (Fig. 17 A) 1.28–1.35 times as wide as
long, widest around anterior 1/4, narrowed posteriorly; postero-lateral corner rounded; surface
finely punctured, densely covered with setae, with 6 macrosetae along lateral margin. Scutellum
with surface rugosely punctured and moderately covered with setae. Elytra (Fig. 17 A) slightly
broadened posteriorly; surface finely punctured, densely covered with setae, with 4 or 5 small
macrosetae laterally. Legs moderate in length; hind tibia 0.96–1.01 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 9 : 9 : 8 : 14.5 in fore tarsus;
15 : 10 : 8 : 7 : 12 in mid tarsus; 14.5 : 12.5 : 10 : 8 : 15 in hind tarsus. Abdomen subparallel-sided,slightly wider than elytra, widest around 5th and 6th segments; surface moderately punctured;
3rd to 5th tergites densely covered with setae; 6th tergites densely covered with setae but basal
area moderately covered with small setae; 7th tergite sparsely covered with minute setae; 8th tergite (Figs. 11 A, 11 C) with 7 (rarely 8) macrosetae; 9th tergite with 2 or 3 macrosetae, which arepoorly differentiated from other setae; 10th tergite with posterior margin rounded, with 1
macroseta.

Palearctic Species of Pella

37

Male: Postero-medial part of pronotal disc with a circular depression; 8th tergite (Fig. 11 A)
with posterior margin roundly and deeply emarginated, its emarginate apex weakly crenate; 8thsternite (Fig. 11 B) with 15–19 macrosetae; 9th sternite with posterior margin slightly emarginated. Aedeagus (Figs. 12 A–12 E, 13 A–13 E) somewhat gourd-shaped in ventral view; apical lobe
curved ventrad at base in lateral view, much narrowed at base, broadened at middle and roundedat apex in ventral view; basal ridge weakly convex; copulatory piece of inner sac as in Figs. 13 F,
13 G. Paramere (Figs. 15 A, 15 B) with apical lobe of paramerite narrowed apically, about 1/3 aslong as condylite.
Female: Pronotum without depression; 8th tergite (Fig. 11 C) with posterior margin slightly
truncate; 8th sternite (Fig. 11 D) with 8–10 macrosetae; sensory setae of 8th sternite generalised,similar to other setae in shape. Spermatheca (Figs. 16 D–16 F) curved three times, somewhat Mshaped; basal part 1.2–1.4 times as long as apical part, acutely curved at middle, its inner wallsparsely wrinkled around apex; apical part weakly curved, its inner wall entirely and densely
wrinkled.
Measurements. BL, 4.8–6.3; FBL, 2.4–2.8; HL, 0.69–0.79; HW, 0.80–0.88; EL, 0.262–
0.293; AL, 1.73–2.11; PL, 0.84–0.96; PW, 1.01–1.25; ELL, 0.94–1.04; ELW, 1.33–1.37; HTL,
0.95–1.00.
Bionomics. Pella limbata is commonly collected from under dead leaves around the nest of
Lasius (Dendrolasius) fuliginosus, but no detailed information on ant-association of this species
has yet been reported. The life cycle of P. limbata is unknown, but the larvae were collected inJuly. This suggests that the life cycle of P. limbata is different from those of the investigated Pella
species described in Chapter 3.
Symbiotic hosts. Lasius (Lasius) brunneus, L. (Cautolasius) flavus, L. (Dendrolasius)
fuliginosus.
Pella horii Maruyama, n. sp.
(Figs. 18–20)“Myrmedonia similis”: Sharp, 1888: 291 (misidentification, record from Japan).

Type material. Holotype: Japan: ?, “Hebi-numa, Teshio-gawa, Teshio-chô, Hokkaidô, 9
VII 1992, S. Hori leg.” (pt) (HUM).
Paratypes: Japan: Hokkaidô: 17 exs., Rebun-tô, 15 VII 1991, N. Yasuda (pt); 2 exs., Sarobetsugen’ya, 9 VI 1991, N. Yasuda (pt); 2 exs., same data but, 11 VIII 1991; 14 exs., Hebi-numa,
Teshio-gawa, Teshio-chô, 9 VII 1992, S. Hori (pt); 1 ex., Onnenai, Tsurui-mura, Akan-gun, 22
VI 1993, Y. Hirama; 4 exs., same data but, 27 VI 1993; 2 exs., same data but, 2 VI 1994; 1 ex.,
Kinashibetsu-kaigan, Otobetsu-chô, 2 IX 1995, Y. Hirama; 2 exs., Iwaobetsu, Shari-chô, 3 VIII
1989, K. Haga; 4 exs., Ishikari-gawa, Kamikawa-chô, 2 VI 1994, N. Yasuda (pt); 2 exs., samedata but, 1 VIII 1994; 4 exs., same data but, 21 VII 1997; 1 ex., Karikachi-tôge, Shintoku-chô, 24
VI 1993, K. Miyashita; 1 ex., Shiomi, Mukawa-chô, 17 IX 2001, S. Hori; 1 ex., Kaitaku-kinenkan, Nopporo-shinrin-kôen, Ebetsu-shi, 17–24 V 2000, S. Hori (fit); 1 ex., Hyakumatsu-zawa,
Sapporo-shi, 1 VI 1998, K. Mizota et al. (pt); 1 ex., same data but, 7 VII 1998; 1 ex., same databut, 14 VII 1998; 1 ex., same data but, 19 VIII 1998; 8 exs., Hamataiki, Taiki-chô, 7 VII 1993, S.
Hori (pt); 2 exs., Horokayantô-numa, Taiki-chô, 7 VII 1993, S. Hori (pt); 9 exs., same data but,
29 VII 1993; 3 exs., same data but, 29 VIII 1993; 17 exs., same data but, 12 VI 1994; 1 ex.,Ôtsu-shitchi, Toyokoro-chô, 7 VII 1993, S. Hori (pt); 1 ex., Shibetsu-gawa, Shibetsu-chô, 25–28
VI 2001, H. Yoshitomi; 1 ex., Shirogane-onsen, Biei-chô, 11 VIII 1997, T. Matsumoto; 1 ex., Hi-

38

M. Maruyama

Fig. 18.

Pella horii Maruyama, n. sp. — A, Fore body, dorsal view; B, right antenna.

gashiyama, Esashi-chô, 15 V 1999, S. Hori; 6 exs., Motoyama, Esashi-chô, 15 V 1999, K. Sasaki. Honshû: 1 ex., Funakawa, Oga-shi, Akita-ken, 26–27 V 1992, S. Saito; 1 ex., Kamuro,
Kaneyama-chô, Yamagata-ken, 26 V 1998, A. Ohkawa; 1 ex., Hirasawa-rindô, Hinoemata-mura,
Fukushima-ken, 21 V 1988, K. Haga (in a fallen tree of Fagus serratus); 1 ex., Shinano-gawa,
Ushigashima, Kawaguchi-chô, Niigata-ken, 30 IV 1995, K. Haga (under accumulated leaves on
sandy riverside); 1 ex., Kazakari-tôge, Hinohara-mura, Tokyo-to, 6 VI 1982, H. Saitoh; 1/,“Miya (written underside of paper card specimen glued on)/Miyanoshita. Miyanoshita, Hokonechô, Kanagawa-ken/Japan., G. Lewis., 1910-320./Myrmedonia similis” (recorded as “Myrmedonia similis” by Sharp (1888)) (BMNH).
Type locality. Teshio-chô, Hokkaidô, Japan.
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô, Honshû).
Etymology. Dedicated to Mr. Shigehisa Hori (Historical Museum of Hokkaidô, Japan) for
his invaluable contributions to the faunistic study in Hokkaidô, and he is a collector of most partof the type series.
Diagnosis. As Sharp (1888) misidentified Pella horii as “P. similis”, these two species areclosely similar to each other, but P. horii is distinguished from P. similis by the gourd shape of
aedeagus and the shorter basal part of the spermatheca. These two species are locally isolated.
Pella horii is also similar to P. limbata in general appearance and shape of aedeagus, but easily
distinguished from the latter by the weakness of emargination at the apex of male 8th tergite,which is very deeply emarginate in P. limbata, and the absence of depression in male pronotum.
These are locally isolated. Among the sympatric species Pella horii is most similar to P. japonica

Palearctic Species of Pella

39

Fig. 19. Pella horii Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.

in general appearance but is distinguished from the latter by the smaller body and the 11th antennal segment longer than the 1st.
Description. Body slender. Brown in ground colour; antennae, mouthparts, legs, andaround posterior margins of 3rd to 7th abdominal segments reddish yellow; elytra with reddish
yellow maculation on mesal area of each elytron; sometimes abdomen uniformly brown. Head
(Fig. 18 A) widest just behind eyes; surface smooth, moderately covered with setae; setae rather
long, almost as long as those on pronotum and elytra; length of eyes 0.33–0.34 times as long ashead width. Antennae (Fig. 18 B) shorter than head, pronotum and elytra combined; 1st segment
long, almost as long as 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments wider than long; 11th segment evidently
longer than 1st; approximate relative lengths of segments from basal to apical: 25 : 11 : 12 : 6 : 6 :40

M. Maruyama

Fig. 20. Pella horii Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca. Female.

5.5 : 6 : 5.5 : 6 : 7 : 30. Pronotum (Fig. 18 A) 1.18–1.25 times as wide as long, widest around anterior 1/4, narrowed posteriorly; postero-lateral corner rounded; surface finely punctured, densely
covered with setae, with 9 or 10 macrosetae along lateral margin. Scutellum with surface reticulated and moderately covered with setae. Elytra (Fig. 18 A) slightly broadened posteriorly; surface finely punctured, densely covered with setae, with 2 or 3 small macrosetae laterally. Legs
moderate in length; hind tibia 097–0.98 times as long as elytra; approximate relative lengths oftarsal segments from basal to apical: 7 : 7 : 6.5 : 13 in fore tarsus; 9.5 : 5.5 : 5 : 5 : 7.5 in mid tarsus;
19 : 10 : 8 : 7.5 : 12.5 in hind tarsus. Abdomen subparallel-sided, slightly wider than elytra, widest
around 4th and 5th segments; surface smooth; 3rd to 6th tergites very sparsely covered withminute setae medially, with long setae along each posterior margin; 7th tergite sparsely covered
with minute setae; 8th tergite (Figs. 19 A, 20 A) with 7 macrosetae; 9th tergite with 4 macrosetae;
10th tergite with posterior margin rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 19 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 19 B) with 11–13 macrosetae; 9th sternite with posterior
margin slightly emarginate. Aedeagus (Figs. 19 C, 19 D) somewhat gourd shaped; apical lobe
curved ventrad at base in lateral view, much narrowed at base and rounded at apex in ventral
view; basal ridge absent; copulatory piece of inner sac as in Fig. 19 E. Paramere with apical lobeof paramerite slightly narrowed apically and obtuse at apex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 20 A) with posterior margin truncate, its truncate apex weakly
crenate; 8th sternite (Fig. 20 B) with 10–13 macrosetae; sensory setae of 8th sternite generalised,Palearctic Species of Pella

41

similar to other setae in shape. Spermatheca (Fig. 20 C) curved three times, somewhat M-shaped;
basal part about twice as long as apical part, acutely curved just before middle, narrowed apicad;
apical part weakly curved, its inner wall of apical part densely wrinkled from apex to around apical 1/2.
Measurements. BL, 4.3–5.4; FBL, 2.1–2.4; HL, 0.63–0.68; HW, 0.73–0.78; EL, 0.252–
0.261; AL, 1.58–1.73; PL, 0.80–0.85; PW, 1.00–1.05; ELL, 0.89–0.95; ELW, 1.15–1.29; HTL,
0.88–0.93.
Bionomics. Most specimens examined from Hokkaidô were collected by pitfall traps, but
in two occations two specimens were collected from nests of Lasius (Cautolasius) flavus. All the
localities lie in open fields or wetlands near lakes, and Lasius flavus prefers such environment.
Therefore, Lasius flavus is most probably a symbiotic host of Pella horii. Further field investigation into Pella horii will confirm host ant. No information was available for the life cycle and theimmatures of this species.
Symbiotic host. Lasius (Cautolasius) flavus.
The similis GroupSpecies included. Pella similis.
Distribution. European subregion, Mediterranean subregion.
Diagnosis. Species of the similis group may be characterised by a combination of the following character states: 1) eye length 0.34–0.35 times as long as head width; 2) 11th antennal
segment slightly shorter than the 1st; 3) pronotum with posterior margin completely rounded, notforming corners postero-laterally; 4) pronotum 1.28–1.39 times as wide as long; 5) pronotum
widest anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytron with oblique yellowish brown maculation around antero-lateral corner to postero-medial area; 8) inner and posterior margins of elytra weakly margined; 9) male 8th sternite longer than in female and almost thesame in size; 10) posterior margin of 8th tergite crenate; 11) posterior margin of male 8th sternite
rounded; 12) lateral projection of apodeme of male 8th tergite not evidently longer than that offemale; 13) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior
margin of the segment; 14) postero-medial margin of female 8th sternite without minute pubescence; 15) apical lobe of aedeagal median lobe gently curved ventrad in lateral view; 16) distal
crests of aedeagus generalised, projected, pointed at apex in lateral view; 17) copulatory piece of
aedeagal median lobe dilated apicad and tridentate at apex in dorsal view.
Comments. The similis group is a monotypical species-group. This species-group may
possibly be allied to the limbata group and the cognata group in resemblance of the shape ofmale 8th sternite, which is relatively widely emarginate posteriorly.
Symbiotic hosts. Lasius (Dendrolasius) fuliginosus, Liometopum microcephalum.
Pella similis (Märkel, 1845), n. comb.
(Figs. 21–23)
Aleochara similis Märkel, 1845: 200 (original description).—Mulsant & Rey, 1873a: 58 (Myrmedonia, description).—des Gozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 123 (Myrmedonia (Pella), key, description).—Reitter, 1909:
43 (Myrmedonia (Pella), key). — Fenyes, 1920: 297 (Zyras (Pella), list).—Bernhauer & Scheerpeltz, 1926: 705(Zyras (Pella), list). — Scheerpeltz, 1934: 1659 (Zyras (Pella), list). — Lohse, 1974: 225 (Zyras (Pella), key).—
Likovský, 1993: 59 (Zyras (Pella), list). — Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntypes: Germany: 1?, “similis Mark* Saxon. in nid. Form. fuliginos.

42

M. Maruyama

Fig. 21.

Pella similis (Märkel). — A, Fore body, dorsal view; B, right antenna.

Märkel [“Saxon. mont., Märk” ( mountainous area of Saxonia, Germany, F. Märkel leg.)]”; 1?,[ditto].
These are deposited in ZMHB.
Type locality. Saxonia, Germany.
Other material. Austria: 1?, 1/, Opcina-Basoviz, Triestiner, Karst, 21–28 IV 1921,
Moczarski (LDFL) (cMar, ex NHMW); 1 ex., Wien, Mariabuin, Skalitzky (LDFL) (NMP); 2??,Styria [Steiermark], Reitter (BMNH). Bulgria: 1?, 1/, Trevna, V–VI 1912 M. Hilf (cMar, ex
NHMW); locality not specified: 1 ?, 3// , (BMNH). Czech Republic: 1 ex., Vtrané, Bohemia, 6
IV 1916, F. Rambousek (LDFL) (NMP); 1 ex., Čejč, Moravia, 26 III 1992, Kuboň; 1 ex., Bohemia, Kunětic. Hora, 27 IV 1947, A. Smetana (CNC); Bohemia, Chlum u Třeb., VI 1916
(CNC). Germany: 1?, 1/, Mesobromentum, Steinberg, Hildesheim, Niedersachen, 2 VII 1997,
V. Assing. Italy: 2??, 1/, Angelo, Gargano, IV 1907, A. Kniz (NMP). Slovakia: Turdošovce,
1–3 V 1986, Ju° sa; 1 ex., Kováčov, 1 V 1955, A. Smetana (CNC); 2 exs., Kamen. most, 3 V 1955,
A. Smetana (CNC). Turkey: 1?, 1/, Yaylasi (alt. 1500 m), 30 km W. Baskonus, Kahramanmaras, 28 IV 2004, Brachat & Meybohm (cAss). Yugoslavia: 1?, 1/, Brus, Kopaonik, VII 1910,
F. Rambousek (LDFL) (NMP).
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Belgium,
Bosnia Herzegovina, Croatia, Czech Republic, France, Germany, Hungary, Italy, Netherlands,
Poland, Portugal, Romania, Slovakia, Russia (South European territory), Switzerland, Turkey,
Yugoslavia. New to Turkey.
Diagnosis. Pella similis is most similar to P. horii in general appearance but distinguishedfrom the latter by the oval shape of aedeagus and the longer basal part of the spermatheca. These

Palearctic Species of Pella

43

Fig. 22. Pella similis (Märkel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

two species are locally isolated. Among the sympatric species, Pella similis is similar to P. limbata and P. cognata but distinguished from P. limbata by the diagnostic characters above and thesmaller body, and from P. cognata by the above spermathecal structure, the smaller body and theshorter apical lobe of aedeagal median lobe.
Redescription. Body slender. Brown in ground colour; antennae, mouthparts, legs slightly
paler; elytron with oblique yellowish brown maculation around antero-lateral corner to posteromedial area; reddish brown around posterior margins of 3rd to 5th abdominal segments; sometimes abdomen uniformly brown. Head (Fig. 21 A) widest just behind eyes; surface smooth,densely covered with setae, which are rather long, almost as long as those on pronotum and elytra; length of eyes 0.34–0.35 times as long as head width. Antennae (Fig. 21 B) shorter than head,pronotum and elytra combined; 1st segment long, longer than 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 as long as 1st; 4th segment almost as long

44

M. Maruyama

Fig. 23. Pella similis (Märkel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, D, spermathecae. Female.

as wide; 5th to 10th segments wider than long; 11th segment slightly shorter than 1st; approximate relative lengths of segments from basal to apical: 25 : 10 : 12 : 6 : 6.5 : 6.5 : 6 : 6.5 : 7 : 8 : 23.
Pronotum (Fig. 21 A) variable in shape, about 1.28–1.39 times wider than long, widest around
anterior 1/5 to 1/4, narrowed posteriorly; posterior margin rounded; surface finely punctured,densely covered with setae, with 7–10 macrosetae along lateral margin; a macroseta at anterior
angle evidently longer than others. Scutellum with surface smooth and densely covered withsetae. Elytra (Fig. 21 A) slightly broadened posteriorly; surface finely punctured, densely covered
with setae, with 3 or 4 small macrosetae laterally. Legs moderate in length; hind tibia 0.9 times aslong as elytra; approximate relative lengths of tarsal segments from basal to apical: 5 : 4 : 4 : 7.5 infore tarsus; 11 : 7 : 7 : 6 : 10 in mid tarsus; 16 : 9 : 8.5 : 8 : 12 in hind tarsus. Abdomen subparallelsided, slightly wider than elytra, widest around 4th and 5th segments; 3rd to 6th tergites very
sparsely covered with minute setae medially, with long setae along each posterior margin; 7th tergite sparsely covered with minute setae; 8th tergite (Figs. 22 A, 23 A) with 7 (rarely 5 or 6)
macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior margin rounded, with 4macrosetae.
Male: Eighth tergite (Fig. 22 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 22 B) with 15–17 macrosetae; 9th sternite with posterior

Palearctic Species of Pella45

margin truncate. Aedeagus (Figs. 22 C, 22 D) oval in ventral view; apical lobe slightly curved
ventrad and pointed at apex in lateral view, rounded at apex in ventral view; basal ridge small androunded; copulatory piece of inner sac as in Fig. 22 E. Paramere with apical lobe of paramerite
subparallel-sided and slightly pointed at apex, somewhat shorter than condylite.
Female: Eighth tergite (Fig. 23 A) with posterior margin slightly emarginate and crenate; 8thsternite (Fig. 23 B) with 11–13 macrosetae; sensory setae of 8th sternite generalised, similar toother setae in shape. Spermatheca (Figs. 23 C, 23 D) curved 3 times; basal part slightly longer
and much narrower than apical part, acutely curved at middle, and its base sometimes touched its
apex; apical part dilated apically, acutely curved just after middle, its inner wall wrinkled from
apex to around apical 1/3.
Measurements. BL, 4.8–5.3; FBL, 2.2–2.5; HL, 0.64–0.68; HW, 0.74–0.80; EL, 0.259–
0.278; AL, 1.55–1.68; PL, 0.71–0.78; PW, 0.98–1.08; ELL, 0.86–1.00; ELW, 1.12–1.28; HTL,
0.87–0.91.
Comments. Distinct intraspecific variations in the shape of pronotum are observed in thisspecies, e.g. it is widest around anterior 4/5 in some specimens, though it is widest around anterior 3/4 in most specimens. Since shape of pronotum is one of the most important characters for
identifying Pella species, several specimens deposited in museums are misidentified with other
Pella species, e.g., P. limbata, P. cognata and P. erratica.
Bionomics. This species is associated with Lasius (Dendrolasius) fuliginosus and Liometopum microcephalum, and is collected around the nest. Rare species (Sharp, 1888; Franc, 1992).
No information was available for the life cycle and the immature stages.
Symbiotic hosts. Lasius (Dendrolasius) fuliginosus, Liometopum microcephalum.
The humeralis GroupSpecies included. Pella humeralis, P. laeviceps.
Distribution. European subregion, Mediterranean subregion.
Diagnosis. Species of the humeralis group may be characterised by a combination of thefollowing character states: 1) eye length 0.35–0.38 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with postero-lateral corners; 4) pronotum 1.28–1.41
times as wide as long; 5) pronotum widest anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytron with yellowish brown maculation around antero-lateral corner to mesal area;8) inner and posterior margins of elytra weakly margined; 9) male 8th sternite longer than in female; 10) posterior margin of 8th tergite crenate; 11) posterior margin of male 8th sternite truncate; 12) lateral projection of apodeme of male 8th tergite evidently longer than that of female;13) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior marginof the segment; 14) postero-medial margin of female 8th sternite without minute pubescence; 15)
apical lobe of aedeagal median lobe almost straight in lateral view; 16) distal crests of aedeagus
well developed, projected, angled at middle of ventral margin in lateral view; 17) copulatory
piece of aedeagal median lobe more or less truncate at apex in dorsal view.
Comments. The two species of the humeralis group are considered to be a sister group by
the close similarity in many morphological points including aedeagal structure. However, no autapomorphy could be found in this species-group. This species-group may possibly be allied to
the funesta group in resemblance of body shape and aedeagal structure.
Symbiotic hosts. Formica (Formica) rufa group, Lasius (Dendrolasius) fuliginosus, L. (L.)
spp.

46

M. Maruyama

Key to the Species of the humeralis Group——

Brown in ground colour; posterior margin of male 8th tergite truncate at apex; pronotum
wider, 1.32–1.41 times as wide as long; apical part of spermatheca curved at middle. Distribution: Europe, Balkan, Caucasia, Siberia. .........................................................P. humeralis.
Pale brown in ground colour; pronotum narrower, 1.30–1.33 times as wide as long; posterior margin of male 8th tergite rounded at apex; apical part of spermatheca almost straight.
Distribution: Caucasia. .........................................................................................P. laeviceps.
Pella humeralis (Gravenhorst, 1802), n. comb.
(Figs. 24–26)

Aleochara humeralis Gravenhorst, 1802: 90 (original description).—Stephens, 1832: 163 (Aleochara, description).—Erichson, 1837: 287 (Myrmedonia, description). — Mulsant & Rey, 1873a: 64 (Myrmedonia, description).—desGozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 122 (Myrmedonia (Pella), key, description).—Reitter, 1909: 43(Myrmedonia (Pella), key). — Fenyes, 1920: 297 (Zyras (Pella), list).—Bernhauer & Scheerpeltz, 1926: 699 (Zyras(Pella), list).— Scheerpeltz, 1934: 1655 (Zyras (Pella), list). — Lohse, 1974: 225 (Zyras (Pella), key).—Likovský,
1993: 59 (Zyras (Pella), list). — Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntypes: Germany: 1/, “5249/humeralis Gr.* [Berolin ( Berlin, Germany)]” (pinned); 1?, 3// , [ditto] (without label, pinned); 1?, “Var. Suecia Schüpp” [“Suec.
Schüpp.” ( Switzerland, Schüppel leg.)] (pinned).
These are deposited in ZMHB.
Type locality. Berlin, Germany; Switzerland.
Other material. Austria: 2 exs. (larvae), Falkert (alt. 1600 m), Gurktaler Alpen, Kärnten,
Austria, 24 VII 1991, V. Assing; 2 exs., Petzen Ct., Ttaventa, VII 1953, Mandl Jakob; 2 exs.,
Tullnerbach, Scheerpeltz; 3 exs., Hochobir (alt. 1600 m), Kärnten, 15 VI 1965, A. Smetana(CNC); 7 exs., Hochobir (alt. 1900–2000 m), Kärnten, 20 VI 1965, A. Smetana (CNC); 2 exs.,
Gbertilliach, Lesachtal, 17 VI, A. Smetana (CNC). Bulgaria: 11 exs., 1400–1900 m Rodopi mts.,
Černatica-Goljam, Persenk, 24 VI 1996, M. Kuboň; 1ex., Rila mts., Kolarov (Belmeken) mt.,
2000–2400 m, 25 VI 1996, M. Kuboň; 3 exs., 1700 m, Pirin, 11 VIII 1949, M. Kuboň; 1 ex., SWBulg. Rila, Geb, Südl. Brovec, 1800 m, 19 VI 1988, Zerche & Behne; 3 exs., Rila, Govedarci,
1200–1500 m, 2–4 VII 1987, M. Kuboň; 1 ex., Pirin, Popinol., 8–9 VI 1989, Ju° za. Czech Republic: 5 exs., Nový Jičin, Moravia, 30 IV 1988; 3 exs., Hradec, Bohemia, 3 IV 1947, A. Smetana(CNC); 5 exs., Lovoš, Bohemia, 22 V 1956, Středohořf (CNC); 1 ex., Sumava, H. Krilda KS, Bohemia, 13 IX 1960, Hu° rka (CNC); 1 ex., Antiegel (alt. 1200–1250 m), Šumeva, Bohemia, 2. VI
1960, Hu° rka (CNC); 1 ex., same data but, alt. 1180–1200 m (CNC). Finland: 2 exs., Lken,
Kemijarai, VI 1995; 1 ex., Sewenz, Davos, Pilafus, VIII 1989, Fng. Kubinast. Germany: 2 exs.,
Perleberg, Karstadt, Brandenburg, 20 VII 1998, Sprick (pt); 2 exs., Trübenbruch, Schönhauser
Damm, Sachsen-Anhalt, V 1998, Sprick (pt); 1 ex., Schönfließ b., Berlin, Mark Brandenburg, 17
IV 1987; 2 exs., Berlin, winter 1981/82, Schulze & Uhlig; 2 exs., Branden–Württemberg, Kaiserstuhl mts., NW Freiburg, IV 1981, Kobel-Lamparski (unbaited pt). Slovakia: 1 ex., Vys. Tatry
Tatr. Lomnica, 5 VII 1947, A. Smetana (CNC); 9 exs., Dolina mnichovskeho potoka, Bardejov,
10 V 2001, M. Maruyama (FFP); 3 exs., Slov. or. Prešov solné j., 31 V 1986, Bulirsch; 1 ex.,
Košice-Bankov, 10 IV 1994, P. Hlavác; 1 ex., Horvce-preosev pri, 10 III. 1995, T. Lackner(LDFL); 3 exs., Vyšlapán ze zamokřeného trávníku, Snineké rybníky, Snine, 17 VI 1991, L.
Klíma; 1 ex., same data but, 28 V 1990, L. Klíma; 1 ex., Malé Trakany, Písčito-bcoreanatý břeh
Tisy, 10 IV 1990, L. Klíma. Turkey: Mengen, Bolu, 8 V 2001, I. Smatana. Kazakhstan: 57 exs.,

Palearctic Species of Pella

Fig. 24.

47

Pella humeralis (Gravenhorst). — A, Fore body, dorsal view; B, right antenna.

Sarymsakty riv. (alt. 1800–2200 m), Sarymsakty mt. reg., southern Altai, 22–26 VI 2000; 16 exs.,
Soldatovo vil., Bukhtarma riv., southern Altai, 21 VI 1998; 41 exs., Katon-Karagai vil. (alt.
100 m), Bukhtarma riv., southern Altai, 20 VI 1999. Russia (West Siberia): 1?, Sotonikovo vil.,Ulan-ude env., Burjatia reg., West Siberia, 4 VI 1991.
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Belgium,
Bosnia Herzegovina, Bulgaria, Belarus, Croatia, Czech Republic, Denmark, Estonia, Finland,
France, Great Britain, Germany, Georgia, Hungary, Italy, Latvia, Lithuania, Luxembourg, Netherlands, Poland, Romania, Russia (North and Central European Territories), Slovakia, Sweden,
Switzerland, Turkey, Uraine. ASIA: Russia (East and West Siberia). New to Turkey.
Diagnosis. Pella humeralis is similar to P. laeviceps in general appearance but dustinguished from the latter by the darker colour, the wider pronotum and the aedeagus having narrower apical lobe and the spermatheca having curved apical part. Among the sympatric species,
this species may be easily distinguished from the other species by the larger body and the distinct
colour.
Redescription. Body slender. Brown in ground colour; antennae, mouthparts and legs reddish brown; elytron with yellowish brown maculation around antero-lateral corner to mesal area;
3rd to 5th abdominal segments reddish brown; sometimes abdomen uniformly brown. Head (Fig.
24 A) widest just behind eyes; surface smooth, densely covered with setae; setae short, stout andrecumbent, somewhat longer than those on pronotum and elytra; length of eyes 0.36–0.37 times
as long as head width. Antennae (Fig. 24 B) shorter than head, pronotum and elytra combined;
1st segment somewhat shorter than 2nd and 3rd combined; 2nd segment shorter than 3rd; 3rdsegment about 2/3 as long as 1st; 4th segment slightly longer than wide; 5th and 6th segments al-

48

M. Maruyama

Fig. 25. Pella humeralis (Gravenhorst). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

most as long as wide; 7th to 10th segments wider than long; 11th segment much shorter than 1st;
approximate relative lengths of segments from basal to apical: 32 : 16 : 20 : 11 : 10: 10 : 11 : 10 :11 : 11 : 27 Pronotum (Fig. 24 A) 1.32–1.41 times as wide as long, widest around anterior 1/5,
narrowed posteriorly; postero-lateral corner rounded; disc with circular depression postero-medially; surface finely punctured, densely covered with stout setae, with 6 or 7 macrosetae along lateral margin; a macroseta at anterior angle evidently longer than others. Scutellum with surface
smooth and densely covered with setae. Elytra (Fig. 24 A) slightly broadened posteriorly; surface
finely punctured, densely covered with long setae, with 3 or 4 small macrosetae laterally. Legs
long; hind tibia 1.16–1.20 times as long as elytra; approximate relative lengths of tarsal segmentsfrom basal to apical: 7.5 : 8 : 8 : 15 in fore tarsus; 18 : 13 : 10 : 8 : 13 in mid tarsus; 29 : 16 : 13 :Palearctic Species of Pella

49

Fig. 26. Pella humeralis (Gravenhorst). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

10 : 16 in hind tarsus. Abdomen expanded, 1.1 times as wide as elytra, widest around 4th and 5th
segments; surface smooth; 3rd to 6th tergites very sparsely covered with minute setae medially,
with long setae along each posterior margin; 7th tergite almost glabrous except lateral area covered with minute setae; 8th tergite (Figs. 25 A, 26 A) with 10 (rarely 11 or 12) macrosetae; 9th
tergite with 4 macrosetae; 10th tergite with posterior margin rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 25 A) much shorter than 8th sternite, with posterior margin truncate, its truncate apex weakly crenate; 8th sternite (Fig. 25 B) with posterior margin truncate,with 16–21 macrosetae; 9th sternite with posterior margin slightly convex medially. Aedeagus(Figs. 25 C, 25 D) amygdaloid in ventral view; apical lobe almost straight in lateral view, pointed
at apex in lateral and ventral views; basal ridge well developed and rounded; copulatory piece of
inner sac as in Fig. 25 E. Paramere with apical lobe of paramerite subparallel-sided and obtuse atapex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 26 A) with posterior margin rounded and weakly crenate; 8thsternite (Fig. 26 B) with 13–16 macrosetae; sensory setae of 8th sternite generalised, similar toother setae in shape. Spermatheca (Fig. 26 C) curved 3 times, somewhat M-shaped; basal partabout twice as long as apical part, bulbous around base, curved at right angle at middle, narrowed
apicad; apical part slightly dilated apicad, gently curved, its inner wall densely wrinkled from
apex to around apical 2/5.
Local variation. Pella humeralis is the most widespread species among the known Pella
species, and its distribution stretches from Northern Europe, through Southern Europe, Turkey
and Caucasia, to southeastern Siberia. Small local variations have been observed among specimens from these localities. The aedeagal median lobe of specimens from Northern and Central

50

M. Maruyama

Europe are characterised by the pointed apical lobe, and it is apparently longer than in the specimens from the other distributional areas. The length of apical lobe tends to become shorter toward the eastern distributional areas.
Measurements. BL, 5.8–7.3; FBL, 2.8–3.1; HL, 0.81–0.85; HW, 0.90–0.98; EL, 0.290–
0.355; AL, 2.45–2.65; PL, 0.96–1.01; PW, 1.28–1.35; ELL, 1.13–1.33; ELW, 1.50–1.75; HTL,
1.36–1.55.
Bionomics. Pella humeralis is associated with species of Formica (Formica) rufa groupand Lasius (Dendrolasius) fuliginosus and collected around their nests. This species is common,and usually dominant myrmecophile in hosts’ nests.
Symbiotic hosts. Formica spp. (the rufa group: F. rufa, F. pratensis, etc.), Lasius (Dendrolasius) fuliginosus.
Pella laeviceps (Eppelsheim, 1880), n. comb.
(Figs. 27–29)
Myrmedonia laeviceps Eppelsheim, 1880: 459 (original description).—Fenyes, 1920: 297 (Zyras (Pella), list).—Bernhauer & Scheerpeltz, 1926: 700 (Zyras (Pella), list). — Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntypes: Azerbaijan: 1/ “Kaukas Leder Alexanderhilf/.6/c. Eppelsh.
Steind. d./laeviceps, Epp. Verh. zool. bot. Ges. Wien, 1879, p. 259/TYPUS”; 1? “? (written on
paper card specimen glued on)/Kaukas, Leder, Suram/c. Eppelsh., Steind. d./laeviceps/TYPUS”;
1? “Kaukas Leder Michailow von Suram/c. Eppelsh. Steind. d./laeviceps/laeviceps Epp det.
Bernhauer”.
These are deposited in NHMW.
Type locality. Alexanderhilf and Suram, Azerbaijan.
Other material. Armenia: 1 ex., Vedi, Khosrov reserve, 30 V 1988, M. Kalashyan (cMar);
1? “Caucasus Meskisches Geb. Leder, (Reitter)” (MNHN). Azerbaijan. 1 ex., “Caucas. Occ.
Circassien Leder. Reitter c. Epplish. Steind” (cMar). Georgia: 1? “Caucasus Abastuman Leder.
(Reitter)” (NHMW); 1/, same data (MNHN); 1 ex., Picunda, 27 VI 1969, M. Dvořák (cMar);
1ex., Zohneti (Tbilisi) (alt. 800 m), 1–10 VI 1987, D. Wrase & M. Schülke (cMar). Russia(South European territory): 2??, 2// “Caucas. occ. Circassien [Cherkessk], Leder, Reitter”(NHMW, NMP); 1?, “Ca. b. Teberda VI. 912 Roubal” (NMP).
Distribution. European subregion, Mediterranean subregion. EUROPE: Armenia, Azerbaijan, Georgia, Russia (South European territory). New to Armenia. This species is probably confined to the Caucasia.
Diagnosis. Pella laeviceps is similar to P. humeralis in general appearance but distinguished from the latter by the evidently paler colour, the narrower pronotum, the aedeagus having
wider apical lobe and the spermatheca having straight apical part. These two species are locallyisolated.
Redescription. Body slender. Pale brown in ground colour; antennae, mouthparts, legs,and 3rd to 5th abdominal segments paler; elytron with yellowish brown maculation around antero-lateral corner. Head (Fig. 27 A) widest just behind eyes; surface smooth, moderately covered
with setae; setae rather long, almost as long as those on pronotum and elytra; length of eyes
0.38–0.39 times as long as head width. Antennae (Fig. 27 B) shorter than head, pronotum andelytra combined; 1st segment long, almost as long as 2nd and 3rd combined; 2nd segment shorter
than 3rd; 3rd segment about 3/5 as long as 1st; 4th segment as long as wide; 5th to 10th segments
wider than long; 11th segment slightly shorter than 1st; approximate relative lengths of segments

Palearctic Species of Pella

Fig. 27.

51

Pella laeviceps (Eppelsheim). —A, Fore body, dorsal view; B, right antenna.

from basal to apical: 30 : 13 : 18 : 9 : 9 : 9.5 : 10 : 8 : 9 : 10 : 29. Pronotum (Fig. 27 A) 1.28–1.33
times as wide as long, widest around anterior 1/5, narrowed posteriorly; postero-lateral cornerrounded; surface smooth, densely covered with setae, with 2 or 3 macrosetae around antero-lateral margin; a macroseta at anterior angle evidently longer than others. Scutellum with surface
smooth and moderately covered with setae. Elytra (Fig. 27 A) slightly broadened posteriorly; surface smooth, densely covered with setae, with 3 or 4 small macrosetae laterally. Legs somewhat
short; hind tibia 0.84–0.88 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 8 : 8 : 7.5 : 13 in fore tarsus; 16 : 11 : 9 : 8 : 11 in mid tarsus;
26 : 12 : 11 : 10 : 12 in hind tarsus. Abdomen expanded, 1.1 times as wide as elytra, widest around
4th and 5th segments; surface smooth; 3rd to 5th tergites almost glabrous except for a row oflong setae along each posterior margin; 6th and 7th tergites almost glabrous; 8th tergite (Figs.
28 A, 29 A) with 8 (rarely 9) macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior margin rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 28 A) much shorter than 8th sternite, with posterior margin rounded and weakly crenate; 8th sternite (Fig. 28 B) with posterior margin truncate, with 20–23
macrosetae; 9th sternite with posterior margin slightly convex medially. Aedeagus (Figs. 28 C,
28 D) amygdaloid in ventral view; apical lobe almost straight in lateral view, pointed at apex inlateral and ventral views; basal ridge well developed and rounded; copulatory piece of inner sac
as in Fig. 28 E. Paramere with apical lobe of paramerite subparallel-sided and obtuse at apex,
about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 29 A) with posterior margin rounded and weakly crenate; 8thsternite (Fig. 29 B) with 14–16 macrosetae; sensory setae of 8th sternite generalised, similar to52

M. Maruyama

Fig. 28. Pella laeviceps (Eppelsheim). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

other setae in shape. Spermatheca (Fig. 29 C) curved once, somewhat L-shaped; basal part
2.0–2.5 times as long as apical part, bulbous around base, curved at right angle, narrowed apicad;
apical part dilated apicad, its inner wall densely wrinkled from apex to around apical 1/3.
Measurements. BL, 5.3–6.1; FBL, 2.5–2.8; HL, 0.73–0.77; HW, 0.76–0.85; EL, 0.299–
0.331; AL, 2.08–2.25; PL, 0.79–0.90; PW, 1.06–1.18; ELL, 0.99–1.08; ELW, 1.30–1.43; HTL,
1.10–1.26.
Bionomics. No host information has been recorded. However, in Georgia, M. Dvořák collected this species from a nest of Lasius sp., which is closely similar to L. (L.) emarginatus Olivier, 1792 (Dvořák, pers. comm.).
Symbiotic host. Lasius (Lasius) cf. emarginatus.

Palearctic Species of Pella

53

Fig. 29. Pella laeviceps (Eppelsheim). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

The funesta GroupSpecies included. Pella funesta, P. comes, P. socia, P. rambouseki, P. jureceki, P. kidaorum,
P. cooterorum.
Distribution. European subregion, Mediterranean subregion, Manchurian subregion.
Diagnosis. Species of the funesta group may be characterised by a combination of the following character states: 1) eye length 0.29–0.33 times as long as head width; 2) 11th antennal
segment longer than the 1st; 3) pronotum with postero-lateral corners; 4) pronotum 1.21–1.46
times as wide as long; 5) pronotum widest anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytra almost unicoloured, blackish brown to brown; 8) inner and posterior margins
of elytra weakly margined; 9) male 8th sternite longer than in female; 10) posterior margin of 8th
tergite crenate or dentate; 11) posterior margin of male 8th sternite rounded; 12) lateral projection of apodeme of male 8th tergite evidently longer than that of female; 13) macrosetae of 8th
abdominal segment generalised in length, exceeding the posterior margin of the segment; 14)postero-medial margin of female 8th sternite with minute pubescence; 15) apical lobe of aedeagal median lobe gently curved ventrad or almost straight in lateral view; 16) distal crests of
aedeagus well developed, projected, angled or rounded around middle of ventral margin in lateralview; 17) copulatory piece of aedeagal median lobe variable in shape, narrowed apicad in dorsal
view.
Comments. No autapomorphy could be found in the funesta group, but the members arevery similar to each other in general appearance, above all in almost uniformly black and slenderbody.
Symbiotic hosts. Lasius (Dendrolasius) spp.

54

M. Maruyama

Key to the Species of the funesta Group
1. Pronotum (1.38–1.46) times as wide as long. ..................................................................... 2.
— Pronotum (1.21–1.33) times as wide as long. ..................................................................... 3.
2. Body black, shining; pronotum almost quadrate, its anterior and posterior corners well angled. Distribution: Japan. ............................................................................................P. socia.
— Body blackish brown, slightly matted; pronotum subtrapeziform, its anterior and posterior
corners more or less rounded. Distribution: China (Beijing Shi), Korea, Russia (Far East). ..
.................................................................................................................................P. jureceki.
3. Male 8th tergite much shorter than 8th sternite, its posterior margin widely rounded andslightly crenate; apical part of spermatheca as long as basal part. Distribution: Europe. ........
...................................................................................................................................P. funesta.
— Male 8th tergite almost as long as 8th sternite, its posterior margin truncate and more orless dentate; apical part of spermatheca shorter than basal part. Distribution: Asia. ......... 4.
4. Body large: 5.9–6.6 mm; antenna long, longer than head, pronotum and elytra combined;
apical lobe of aedeagal median lobe curved ventrad; sensory setae of female 8th sternite
generalised. Distribution: Japan. ...............................................................................P. comes.
— Body small to medium: 4.4–5.3 mm; antenna shorter than or as long as head, pronotum andelytra combined; apical lobe of aedeagal median lobe almost straight; sensory setae of female 8th sternite flattened. .................................................................................................. 5.
5. Antenna almost as long as head, pronotum and elytra combined; basal part of spermatheca
five times as long as apical part. Distribution: Korea, Russia (Far East). .........P. rambouseki.
— Antenna much shorter than head, pronotum and elytra combined; basal part of spermatheca
about twice as long as apical part. ....................................................................................... 6.
6. Apical lobe of aedeagal median lobe slightly sinuate; apical part of spermatheca withoutouter projection. Distribution: Japan. ...................................................................P. kidaorum.
— Apical lobe of aedeagal median lobe almost straight; apical part of spermatheca with anouter projection. Distribution: China (Beijing Shi, Yunnan Sheng). ................P. cooterorum.
Pella funesta (Gravenhorst, 1806), n. comb.
(Figs. 30–32)
Aleochara funesta Gravenhorst, 1806: 168 (original description).—Stephens, 1832: 163 (Aleochara, description).—Erichson, 1837: 287 (Myrmedonia, description). — Mulsant & Rey, 1873a: 62 (Myrmedonia, description).—desGozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 122 (Myrmedonia (Pella), key, description).—Reitter, 1909: 43(Myrmedonia (Pella), key). — Fenyes, 1920: 297 (Zyras (Pella), list).—Bernhauer & Scheerpeltz, 1926: 698 (Zyras(Pella), list).— Scheerpeltz, 1934: 1653 (Zyras (Pella), list). — Lohse, 1974: 225 (Zyras (Pella), key).—Likovský,
1993: 59 (Zyras (Pella), list). — Smetana, 2004: 461 (Zyras (Pella), list).
Aleochara crassicornis Stephens, 1832: 113 (original description).—Fenyes, 1920: 297 (synonym of funesta, list).—Bernhauer & Scheerpeltz, 1926: 701 (synonym of funesta, list).—Smetana, 2004: 461 (synonym of funesta, list).
Myrmedonia atrata Heer, 1839: 309 (original description). —Mulsant & Rey, 1873a: 63 (synonym of funesta, list).—desGozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 122 (synonym of funesta, list). — Fenyes, 1920: 297 (synonym of
funesta, list).— Bernhauer & Scheerpeltz, 1926: 701 (synonym of funesta, list).—Smetana, 2004: 461 (synonym of
funesta, list).

Type material. Aleochara funesta. Syntypes: Germany: 1/, “5253/funesta Gr. [“Berolin”(Berlin, Germany)]” (pinned); 1 ex., [ditto] (without label 2nd to 10th abdominal segments
missing, pinned); 1/, [ditto] (without label, pinned).
These are deposited in ZMHB. According to “Catalog. General. Musei Zoologici Berolinen”

Palearctic Species of Pella

Fig. 30.

55

Pella funesta (Gravenhorst). — A, Fore body, dorsal view; B, right antenna.

ten syntypes should exist in the type series, but only three specimens were found in ZMHB.
Aleochara crassicornis. Not examined. Type locality: Great Britain.
Myrmedonia atrata. Not examined. Type locality: Switzerland.
Type locality. Berlin, Germany.
Other material. Austria: 2 exs., Lainzer, Tiergarten, 30 VI 1949, O. Scheerpeltz (LDFL).
Bulgaria: 2 exs., Trevna, V–VI 1912, M. Hilf. Croatia: 2// , 1 km E Marušići, Felsmoos, Coline, 26 VI 1981, V. Puthz. Czech Republic: 1 ex., Novỳ Jićĺí, Moravia, 30 IV 1988, Kuboň; 1ex., Nymburk, Bohemia, 1958, J. Havelka (CNC); 2 exs., Hradec, Bohemia, 3 IV 1947, A.
Smetana (CNC); 1 ex., Lovoš, Bohemia, 22 V 1956, Středohořf. Germany: 1?, 1/, Kaiserstuhl
mts., Badenwürttenberg, NW Freiberg, IV 1981, Kobel-Lamparski (pt). Slovakia: 1?, Plešivec,
15 VI 1955, A. Smetana; 1/, Lešt, VII, 1955, A. Smetana; 1 ex., Horovce, 11 VI 1994, P.
Hlaváč; 1 ex., Kováčov, 20 VII 1963, A. Smetana (CNC); 1 ex., Ruská Poruba, 4 VIII 1956, A.
Smetana (CNC); 2 exs., Ruská Poruba, 12 IX 1956, A. Smetana.
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Belgium,
Bosnia Herzegovina, Bulgaria, Belarus, Croatia, Czech Republic, Denmark, Finland, France,
Great Britain, Germany, Georgia, Hungary, Italy, Latvia, Luxembourg, Netherlands, Norway,
Poland, Romania, Russia (North, Central and South European territories), Slovakia, Sweden,
Switzerland, Ukraine. NORTH AFRICA: Algeria.
Diagnosis. Pella funesta is the largest species in the funesta group, and well characterisedby the male 8th tergite much shorter than sternite. Among the sympatric species, this species may
be easily distinguished by the uniformly black ground colour.
Redescription. Body slender. Black in ground colour; antennae, mouthparts, legs slightly

56

M. Maruyama

Fig. 31. Pella funesta (Gravenhorst). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

reddish brown to brown. Head (Fig. 30 A) slightly depressed above, wider than long; surface
smooth, moderately covered with setae; setae rather long, longer than those on pronotum and elytra in length; length of eyes 0.32–0.33 times as long as head width. Antennae (Fig. 30 B) shorter
than head, pronotum and elytra combined a; 1st segment long, almost as long as 2nd and 3rd
combined; 2nd segment about 3/4 as long as 3rd; 3rd segment about 4/7 as long as 1st; 4th to 8th
segments almost as long as wide; 9th to 10th segments slightly wider than long; 11th segment
much shorter than 1st; approximate relative lengths of segments from basal to apical: 34 : 15 :20 : 10 : 10.5 : 11 : 11 : 11 : 11.5 : 12 : 29. Pronotum (Fig. 30 A) 1.29–1.31 times as wide as long,widest around anterior 1/4, slightly constricted posteriorly; postero-lateral corner rounded; surface finely punctured, densely covered with setae, with 7 or 8 macrosetae along lateral margin.
Scutellum with surface finely punctured, moderately covered with setae. Elytra (Fig. 30 A) slightly broadened posteriorly; surface finely punctured, densely covered with setae, with 4 or 5 small

Palearctic Species of Pella

57

Fig. 32. Pella funesta (Gravenhorst). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca. Female.

macrosetae laterally. Legs long; hind tibia 1.19–1.21 times as long as elytra; approximate relative
lengths of tarsal segments from basal to apical: 10.5 : 8 : 8 : 14 in fore tarsus; 18 : 11.5 : 10 : 8 :11.5 in mid tarsus; 29 : 15 : 11.5 : 10 : 15 in hind tarsus. Abdomen expanded, 1.1 times wider thanelytra, widest around 5th and 6th segments; surface smooth; 3rd to 6th tergites glabrous except
for a row of setae along each posterior margin; 7th tergite almost glabrous except lateral area
covered with setae; 8th tergite (Figs. 31 A, 32 A) with 8 (rarely 9) macrosetae; 9th tergite with 4macrosetae; 10th tergite with posterior margin truncate, with 4 macrosetae.
Male: Eighth tergite (Fig. 31 A) with posterior margin rounded; 8th sternite (Fig. 31 B) with
28–30 macrosetae; 9th sternite with posterior margin slightly convex medially. Aedeagus (Figs.
31 C, 31 D) amygdaloid in ventral view; apical lobe almost straight in lateral view, pointed atapex in lateral and ventral views; basal ridge well developed, reaching near bottom of basal capsule; copulatory piece of inner sac as in Fig. 31 E. Paramere with apical lobe of paramerite parallel-sided, rounded apically, about 1/4 as long as condylite.
Female: Eighth tergite (Fig. 32 A) with posterior margin slightly truncate; 8th sternite (Fig.
32 B) with 15–17 macrosetae; sensory setae of 8th sternite generalised, similar to other setae inshape. Spermatheca (Fig. 32 C) curved three times; basal part almost as long as apical part,
acutely curved near base, slightly dilated apicad; apical part curved at right angle at middle, its
inner wall densely wrinkled from apex to around apical 2/3.
Measurements. BL, 5.6–6.6; FBL, 2.6–2.9; HL, 0.78–0.83; HW, 0.88–0.90; EL, 0.282–
0.300; AL, 2.28–2.50; PL, 0.88–0.93; PW, 1.15–1.20; ELL, 1.08–1.10; ELW, 1.45–1.48; HTL,
1.29–1.33.
Bionomics. This species is associated with Lasius (Dendrolasius) fuliginosus and collectedaround the nest. Common species (Franc, 1992).
Symbiotic host. Lasius (Dendrolasius) fuliginosus.

58

M. Maruyama

Pella comes (Sharp, 1874)(Figs. 33–35)
Myrmedonia comes Sharp, 1874: 10 (original description). — Bernhauer & Scheerpeltz, 1926: 696 (Zyras (Pella), list).—Fenyes, 1920: 297 (Zyras (Pella), list). — Kistner, 1972: 154 (Pella, description).—Dvořák, 1981: 54 (Zyras (Pella),key).—154: Shibata, 1985: 320 (Pella, description). — Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntypes: Japan: 1?, “TYPE (red round curator label)/Japan (yellow
round paper)/Japan. G. Lewis./Sharp Coll. 1905-313./Myrmedonia comes Type D.S./SYNTYPE(blue round curator label)”; 3// (two are glued on same paper card), “Japan (yellow round
paper)/Japan. G. Lewis./Sharp Coll. 1905-313.”.
These are deposited in BMNH.
Type locality. Japan (no other information).
Other material. Japan: Honshû: 6 exs., Tamagawa, Tazawako-machi, Akita-ken, 12–13
VI 1999, M. Sano (LDFJ); 1 ex., Gosho-ko, Morioka-shi, Iwate-ken, 13–14 VI 1999, M. Sano
(LDFJ); 4 exs., Narugo-onsen, Naruko-machi, Miyagi-ken, 14–17 VI 1999, M. Sano (LDO); 10
exs., Aobayama, Sendai-shi, Miyagi-ken, 22 IX 2001, M. Maruyama (LDN); 5 exs., Yukiwaribashi, Nishigô-mura, Fukushima-ken, 29 VII 2000, T. Kobayashi (LDC); 12 exs., Yahiko-yama,
Nishikambara-gun, Niigata-ken, 4 V 1993, M. Nishikawa (LDN); 3 exs., Tamugi, Tôkamachishi, Niigata-ken, 18 IV 1998, T. Kishimoto; 1 ex., Mizuhiki, Tateiwa-mura, Minamiaizu-gun,
Fukushima-ken, 12 VI 2004, H. Kamezawa (LDFJ); 25 exs., Aoyagidaira, Kuriyama-mura,Tochigi-ken, 1 IX 1999, T. Kobayashi (LDS ); 9 exs., same data but, 29 VIII 1999, T. Kobayashi(LDS ); 17 exs., Tobiyama-jôshi, Utsunomiya-shi, Tochigi-ken, 17 VI 1998, M. Maruyama(LDFJ); 1 ex., same data but, 17–18 VI 1998; 2 exs., Mizuhono, Utsunomiya-shi, Tochigi-ken,
17–18 VI 1998, M. Maruyama (LDS ); 11 exs., Sayado, Môka-shi, Tochigi-ken, 15 VI 2000, T.
Kobayashi & H. Obata (LDFJ); 2 exs., 4 km NW Onioshidashi, Gumma-ken, 18–21 VII 1980,
A. Smetana & Z. Smetana (CNC); 26 exs., Nakano, Shôwa-machi, Kitakatsushika-gun, Saitamaken, 4 V 2003, H. Kamezawa (LDFJ); 4 exs., Kuroishi-jinja (Kumai), Hatoyama-machi, Saitama-ken, 29 IV 2000, S. Arai (LDS); 3 exs., Shiroishi-tôge, Higashichichibu-mura, Saitama-ken,
20 VI 1999, S. Arai (LDFJ); 1 ex., Akigase-kôen (alt. 50 m), Saitama-shi, Saitama-ken, 8 V
2001, H. Sugaya (LDFJ); 4 exs., Sugaya, Ranzan-machi, Saitama-ken, 13 VII 1998, K. Toyoda(LDS); 2 exs., Shôgun-sawa, Ranzan-machi, Saitama-ken, 25 IV 1999, K. Toyoda (LDS); 8 exs.,
Takao-san (alt. 450 m), Hachiôji-shi, Tokyo-to, 1 VII 1998, M. Maruyama (LDS); 23 exs., samedata but, 4 VII 1998; 9 exs., same data but, 4 VI 2001; 9 exs., same data but, 22 VII 2003; 1 ex.,Shiroyama-kôen, Inagi-shi, Tokyo-to, 21 V 2004, A. Yoshida (LDFJ); 1 ex., same data but, 12 VI
2004; 3 exs., same data but, 13 VI 2004; 4 exs., same data but, 26 VI 2004; 1?, “Tokio [Tokyoto]”, 25–27 IX 1881, G. Lewis (BMNH); 10 exs., Aoba-chô, Chiba-shi, Chiba-ken, 24 VI 1998,
M. Maruyama (LDS); 3 exs., Ikuta-ryokuchi, Kawasaki-shi, Kanagawa-ken, 13 IV 2002, K. Matsumoto (LDFJ); 2 exs., Masukata-yama, Kawasaki-shi, Kanagawa-ken, 23 VI 1995, K. Kawada(LDS ); 1?, “Bukenji [Bukenji, Yokohama-shi, Kanagawa-ken]”, G. Lewis (no other data)(BMNH); 1?, “Yokohama [Yokohama-shi, Kanagawa-ken]”, G. Lewis (no other data) (BMNH);
1 ex., Mikage, Tsukui-machi, Kanagawa-ken, 14 IV 1979, R. Kiryu (SCM); 15 exs., Jimmuji,
Zushi-shi, Kanagawa-ken, 20 VI 2003, M. Maruyama; 1 ex., Fukazawa-jûnimagari, Nakamaru,
Nagasaka-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 1 ex., Nagasakakamijô, Nagasaka-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDS);
2 exs., Shôbuzawa, Futaba-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama(LDFJ); 5 exs., Ômukawa-keikoku, Hakushû-machi, Yamanashi-ken, 11 VII 2004, S. Nomura

Palearctic Species of Pella

Fig. 33.

59

Pella comes (Sharp). — A, Fore body, dorsal view; B, right antenna.

(LDFJ); 1/, “Seba [Seba, Shiojiri-shi, Nagano-ken]”, 30 VII 1881, G. Lewis (BMNH); 2 exs.,
Misuzu, Ina-shi, Nagano-ken, 18 IX 2000, T. Kobayashi (LDFJ); 3 exs., Tenryu-chô, Ina-shi,
Nagano-ken, 22 VII 1958, Y. Hayashi (LDS); 4 exs., Ôkubo, Komoro-shi, Nagano-ken, 29–30 IV
2001, H. Kamezawa (LDFJ); 5 exs., Shaka-shindo, Haku-san, Ishikawa-ken, 30 VII–2 VIII 2002,
H. Hoshina (pt); 30 exs., same data but, 2–22 VIII 2002; 1 ex., Hinogawa-dani, Imajô-chô,Fukui-ken, 18 VI 1981, Y. Hayashi; 12 exs., Ogamigô (alt. 800 m), Shôkawa-mura, Gifu-ken, 5–6
VIII 1998, M. Maruyama (LDS ); 1 ex., Kamiichi, Yoshino-chô, Nara-ken, 27 VII 1980, M.
Yasui; 1/, “Kobe [Kôbe-shi, Hyôgo-ken]”, 28 VI 1881, G. Lewis (BMNH); 1 ex., Gôkei, Sôjashi, Okayama-ken, 16 V 1993, T. Ito; 1 ex., Urahikimi, Shimane-ken, 6 VI 1988, S. Nomura(LDS ); 1 ex., Tokusa, Atô-chô, Yamaguchi-ken, 24 IV 1988, S. Nomura. Shikoku: 5 exs., Fujiojinja, Takamatsu-shi, Kagawa-ken, 31 V 2001, F. Itô, M. Maruyama & Y. Kamite (LDS); 1 ex.,same data but, 1 VI 2001 (LDFJ); 1 ex., Usa-jinja, Nagao-chô, Kagawa-ken, 31 V 2001, F. Itô,
M. Maruyama & Y. Kamite (LDS ); 1 ex., Daisen-zan, Kotonami-chô, Kagawa-ken, 1 VI 2001, F.
Itô & M. Maruyama (LDC ); 1 ex., Koyama, Miki-chô, Kagawa-ken, 1 VI 2001, F. Itô & M.
Maruyama (LDS); 9 exs., Atago-yama, Kotohira-chô, Kagawa-ken, 1 VI 2001, F. Itô, M.
Maruyama & Y. Kamite (LDS). Kyûshû: 1?, “Japan, Gokayama, Chikuzen [Gokayama, Nakagawa-chô, Fukuoka-ken], 14. VII. 1936” (BMNH); locality not specified: 1?, Japan, G. Lewis(no other data) (BMNH); 1/, “I, Japan/15. 5.” (no other data) (BMNH); 1 ex., Tsubaki-yama,
Saga-ken, 2 X 1977, H. Ohishi.
Distribution. Manchurian subregion. ASIA: Japan (Honshû, Shikoku, Kyûshû). This
species is most probably confined to Honshû, Shikoku and Kyûshû, Japan.
Diagnosis. Pella comes is similar to P. rambouseki, P. kidaorum and P. cooterorum in gen-60

M. Maruyama

Fig. 34. Pella comes (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

eral appearance and colour but can be distinguished from the latter three by the larger body, thelonger antennae, the narrower pronotum and the oval aedeagus. These four species are locallyisolated. Among the sympatric species, Pella comes is somewhat similar in colour to P. socia but
distinguished from it by the larger body and the pronotal corners not distinctly angled.
Redescription. Body slender. Black in ground colour; antennae, mouthparts, legs reddish
brown to brown. Head (Fig. 33 A) widest just behind eyes; surface smooth, moderately covered
with setae; setae moderate in length, as long as those on pronotum and elytra; length of eyes
0.29–0.30 times as long as head width. Antennae (Fig. 33 B) almost as long as head, pronotumand elytra combined; 1st segment slightly shorter than 2nd and 3rd combined; 2nd segment about
2/3 as long as 3rd; 3rd segment about 3/5 as long as 1st; 4th to 7th segments slightly longer thanwide; 8th and 9th segments almost as long as wide; 10th segment slightly wider than long; 11th

Palearctic Species of Pella

61

Fig. 35. Pella comes (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C–F, spermathecae. Female.

segment much shorter than 1st; approximate relative lengths of segments from basal to apical:
35 : 14 : 23 : 13.5 : 14 : 14 : 13 : 13 : 13 : 12 : 22.5. Pronotum (Fig. 33 A) 1.21–1.29 times as wide as
long, widest around anterior 1/5, evidently constricted posteriorly; lateral margins subparallelsided near base; postero-lateral corner weakly angled; surface finely punctured, moderately covered with setae, with 7 or 8 macrosetae along lateral margin. Scutellum with surface finely punctured, densely covered with setae. Elytra (Fig. 33 A) subparallel-sided; surface finely punctured,densely covered with setae, with 6 or 7 small macrosetae laterally. Legs very long; hind tibia
1.35–1.48 times as long as elytra; approximate relative lengths of tarsal segments from basal toapical: 12 : 8 : 7 : 15 in fore tarsus; 19 : 10 : 9 : 7 : 12 in mid tarsus; 33 : 12 : 11 : 10 : 14 in hind tarsus. Abdomen expanded, 1.1 times wider than elytra, widest around 5th and 6th segments; surface smooth; 3rd to 6th tergites glabrous except for a row of setae along each posterior margin;62

M. Maruyama

7th tergite almost glabrous except posterior and lateral areas; 8th tergite (Figs. 34 A, 35 A) with 8(rarely 9) macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior margin truncate,with 4 macrosetae.
Male: Eighth tergite (Fig. 34 A) with posterior margin truncate, its truncate apex dentate; 8thsternite (Fig. 34 B) with 16–18 macrosetae; 9th sternite with posterior margin truncate. Aedeagus(Figs. 34 C, 34 D) oblong-oval in ventral view; apical lobe curved ventrad and pointed at apex inlateral view, subparallel-sided in basal 1/2 and angulated at apex in ventral view; basal ridge
weakly convex, invisible in lateral view; copulatory piece of inner sac as in Fig. 34 E. Paramerewith apical lobe of paramerite parallel-sided, rounded apically, about 1/4 as long as condylite.
Female: Eighth tergite (Fig. 35 A) with posterior margin slightly emarginate, weakly dentate
around emarginate apex; 8th sternite (Fig. 35 B) with 14–16 macrosetae; sensory setae of 8th
sternite generalised, similar to other setae in shape. Spermatheca (Figs. 35 C–35 F) curved two or
three times; basal part three times as long as apical part, somewhat bulbous around base, acutely
curved, sometimes its basal part touching its middle; apical part straight or gently curved, its
inner wall densely wrinkled from apex to around apical 1/2.
Measurements. BL, 5.5–6.2; FBL, 2.5–2.7; HL, 0.76–0.89; HW, 0.89–0.95; EL, 0.273–
0.281; AL, 2.57–2.80; PL, 0.92–1.03; PW, 1.20–1.25; ELL, 0.93–1.10; ELW, 1.31–1.56; HTL,
1.38–1.49.
Bionomics. This species is commonly collected from colonies of Lasius (Dendrolasius)
spp. and widely distributed from lowlands to mountainous zones in Honshû, Shikoku andKyûshû.
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) orientalis, L. (D.)
nipponensis, L. (D.) capitatus.
Pella socia (Sharp, 1874)(Figs. 36–38)
Myrmedonia socius [sic!] Sharp, 1874: 10 (original description).—Fenyes, 1920: 297 (Zyras (Pella), list).—Bernhauer& Scheerpeltz, 1926: 705 (Zyras (Pella), list). — Kistner, 1972: 154 (Pella “socius [sic!]”, comment).—Dvořák,
1981: 54 (Zyras (Pella), key). — Shibata, 1985: 320 (Pella “socius [sic!]”, description).—Smetana, 2004: 461 (Zyras(Pella), list).

Type material. Syntypes: Japan: 1?, “? (written on paper card specimen glued on)/Type(red round curator label)/Japan (yellow paper)/Japan. G. Lewis./Sharp Coll. 1905-313./Myrmedonia socius. Type D.S./SYN-TYPE (blue round curator label)”; 1/, “//Japan (yellow paper)/Japan. G. Lewis./Sharp Coll. 1905-313./SYN-TYPE (blue round curator label)”.
These are deposited in BMNH.
Type locality. Japan (no other information).
Other material. Japan: Hokkaidô: 13 exs., Himenuma, Rishiri-tô, 4 IX 1990, T. Kishimoto (pt); 3 exs., Hebi-numa, Teshio-gawa, Teshio-chô, 9 VII 1992, S. Hori (pt); 12 exs., Makoi,
Shari-chô, 27–29 V 2001, Y. Kida (LDFJ); 2 exs., Mukôengaru, Engaru-chô, 25 V 2000, Y.
Kida; 78 exs., Taihei, Maruseppu-chô, 29–31 V 2000, Y. Kida (LDFJ); 3 exs., same data but,
11–12 VIII 2000; 23 exs., same data but, 2–3 VI 2000, Y. Kida; 7 exs., same data but, 16–17 VI
2000, Y. Kida; 26 exs., Nosaki, Bihoro-chô, 23 VI 2001, Y. Yamaga (LDC); 10 exs., Makoi,
Shari-chô, 28 V 2000, Y. Kida; 1 ex., Akkeshi, Nemuro-shi, 30 VI 1996, Y. Hirata; 1 ex.,
Kaitaku-kinenkan, Nopporo-shinrin-kôen, Ebetsu-shi, 20 IV 2003, S. Hori (LDFJ); 29 exs.,Ôsawaguchi, Nopporo-shinrin-kôen, Ebetsu-shi, 31 V 1999, M. Maruyama (LDFJ); 1 ex., same

Palearctic Species of Pella

Fig. 36.

63

Pella socia (Sharp). — A, Fore body, dorsal view; B, right antenna.

data but, 9 V 2000; 2 exs., same data but, 4 V 2001; 1 ex., same data but, 16–19 VI 2001, S.
Hori; 1 ex., Shibumbetsu, Nopporo-shinrin-kôen, Ebetsu-shi, 9 V 2000, M. Maruyama (LDFJ);
14 exs., same data but, 11 V 2000 (LDFJ); 1 ex., same data but, 18 V 2000 (LDFJ); 61 exs.,
Tomambetsu, Nopporo-shinrin-kôen, Ebetsu-shi, 4 V 2000, M. Maruyama (LDS); 49 exs., samedata but, 11 V 2000 (LDS ); 5 exs., same data but, 21 V 2000 (LDFJ); 1 ex., same data but, 18 V
2000 (LDS ); 17 exs., Hitsujigaoka, Sapporo-shi, 25 VII 1998, M. Maruyama (LDFJ); 2 exs.,
same data but, 21 VI 1998 (LDS ); 4 exs., same data but, 20 V 1999; 4 exs., same data but, 18 V
2000; 1 ex., Hakken-zan, Sapporo-shi, 31 V 2001, M. Maruyama (LDFJ); 6 exs., Kannon-zawa,
Sapporo-shi, 1 VI 2001, M. Maruyama (LDFJ); 4 exs., Asari, Otaru-shi, 1 IX 1998, K. Sugisima(LDFJ); 24 exs., Yûtoku, Ôtaki-mura, 24 VIII 2001, M. Maruyama (LDFJ); 17 exs., Akagawa,
Hakodate-shi, 15 VIII 1992, M. Nishikawa (LDFJ). Honshû: 9 exs., Mizuhiki, Tateiwa-mura,
Minamiaizu-gun, Fukushima-ken, 12 VI 2004, H. Kamezawa (LDFJ); 2 exs., Yukiwari-bashi,
Nishigô-mura, Fukushima-ken, 29 VII 2000, T. Kobayashi (LDC); 1 ex., Kashi-onsen, Nishigômura, Fukushima-ken, 16 VI 1998, M. Maruyama (LDFJ); 8 exs., Tobiyama-jôshi, Utsunomiyashi, Tochigi-ken, 17 VI 1998, M. Maruyama (LDFJ); 10 exs., Shimokomoriya, Môka-shi, Tochigi-ken, 6 VII 1999 (LDFJ); 13 exs., Sayado, Môka-shi, Tochigi-ken, 15 VI 2000, T. Kobayashi &H. Obata (LDFJ); 3 exs., Kami-imai, Hosaka-chô, Nirasaki-shi, Yamanashi-ken, 15 VIII 2000, T.
Kobayashi; 1 ex., Nakano, Shôwa-machi, Kitakatsushika-gun, Saitama-ken, 14 IV 2002, H.
Kamezawa (LDFJ); 3 exs., same data but, 25 V 2003; 3 exs., Shiroishi-tôge, Higashichichibumura, Saitama-ken, 9 V 1999, K. Toyoda (LDS); 2 exs., same data but, 17 VI 1999; 7 exs., samedata but, 20 VI 1999, S. Arai; 3 exs., Minoyama, Minano-chô, 15 VII 2000, K Toyoda (LDS); 1

64

M. Maruyama

Fig. 37. Pella socia (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view;C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

ex., Tokigawa, Kamagata, Ranzan-machi, Saitama-ken, 22 IV 1999, K. Toyoda (LDFJ); 2 exs.,
Shioyama, Kamagata, Ranzan-machi, Saitama-ken, 17 VI 2000, K. Toyoda (LDFJ); 2 exs.,
Shôgun-sawa, Ranzan-chô, Saitama-ken, 25 IV 1999, K. Toyoda (LDS); 24 exs., Takao-san (alt.
450 m), Hachiôji-shi, Tokyo-to, 1 VII 1998, M. Maruyama (LDS); 13 exs., same data but, 4 VII
1998; 11 exs., same data but, 4 VI 2001; 4 exs., Shiroyama-kôen, Inagi-shi, Tokyo-to, 23 V 2004,
A. Yoshida (LDFJ); 3 exs., same data but, 13 VI 2004; 1 ex., same data but, 26 VI 2004; 1 ex.,same data but, 11 VI 2004; 5 exs., Azeta, Sakura-shi, Chiba-ken, 20 VI 1998, M. Maruyama(LDS); 2 exs., same data but, 23–24 VI 1998; 9 exs., Masukata-yama, Kawasaki-shi, Kanagawaken, 23 VI 1995, K. Kawada (LDS ); 1 ex., Mikage, Tsukui-machi, Kanagawa-ken, 14 IV 1979,
R. Kiryu (SCM); 4 exs., Fukazawa-jûnimagari, Nakamaru, Nagasaka-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 2 exs., Nagasakakamijô, Nagasaka-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDS); 1 ex., Misuzu, Ina-shi, Nagano-ken,
18 IX 2000, T. Kobayashi (LDFJ); 9 exs., Ôkubo, Komoro-shi, Nagano-ken, 29–30 IV 2001, H.
Kamezawa (LDFJ). Shikoku: 3 exs., Fujio-jinja, Takamatsu-shi, Kagawa-ken, 31 V 2001, F. Itô,

Palearctic Species of Pella

65

Fig. 38. Pella socia (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view;C, spermatheca. Female.

M. Maruyama & Y. Kamite (LDS); 2 exs., Ôtaki-san, Shionoe-chô, Kagawa-ken, 2 VI 2001, M.
Maruyama & Y. Kamite (LDS). Kyûshû: 1?, “Nagasaki [Nagasaki-shi, Nagasaki-ken]”, 13 II–21
IV 1881, G. Lewis (no other data) (BMNH); locality not specified: 1?, 2// , “Japan” G. Lewis(no other data) (BMNH).
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô, Honshû, Shikoku, Kyûshû).
This species is most probably confined to Japan.
Diagnosis. Pella socia may be easily distinguished from the other species of Pella by the
pronotum distinctly quadrate. Among the sympatric species in Hokkaidô, Pella socia is similar to
P. kidaorum in body colour and general shape of pronotum but distinguished from it by the
pronotal setation sparser, the pronotal corners well angled and the apical lobe of aedeagal median
lobe not sinuate. In the other regions, Honshû, Shikoku and Kyûshû, P. socia is similar in bodycolour to P. comes but easily distinguished from the latter by the pronotal shape and setation
above and the smaller body.
Redescription. Body slender, shining. Black in ground colour; antennae, mouthparts, legs
reddish brown to brown. Head (Fig. 36 A) widest just behind eyes; surface smooth, moderately
covered with setae; setae moderate in length, longer than those on pronotum and elytra; length ofeyes 0.30–0.31 times as long as head width. Antennae (Fig. 36 B) shorter than head, pronotumand elytra combined; 1st segment slightly shorter than 2nd and 3rd combined; 2nd segment about
2/3 as long as 3rd; 3rd segment about 3/5 as long as 1st; 4th to 7th segments slightly longer thanwide; 8th and 9th segments almost as long as wide; 10th segment slightly wider than long; 11th
segment much shorter than 1st; approximate relative lengths of segments from basal to apical:
26 : 12 : 15 : 9.5 : 9.5 : 9.5 : 9.5 : 9.5 : 10 : 11 : 20. Pronotum (Fig. 36 A) quadrate, 1.43–1.46 times
as wide as long, widest around anterior 1/4, slightly narrowed posteriorly; postero-lateral corner
angled; surface finely punctured, moderately covered with setae, with 5 or 6 macrosetae along

66

M. Maruyama

lateral margin. Scutellum with surface finely punctured, densely covered with setae. Elytra (Fig.
36 A) slightly widened posteriorly; surface finely punctured, densely covered with setae, with 3
or 4 small macrosetae laterally. Legs long; hind tibia 1.12–1.17 as long as elytra; approximate
relative lengths of tarsal segments from basal to apical: 7 : 6.5 : 6 : 13 in fore tarsus; 12 : 8.5 :8 : 7 : 11.5 in mid tarsus; 19 : 11 : 9.5 : 8 : 12 in hind tarsus. Abdomen expanded, as wide as elytra,
widest around 4th and 5th segments; surface smooth; 3rd to 6th tergites glabrous except for a rowof setae along each posterior margin; 7th tergite almost glabrous except for posterior and lateralareas; 8th tergite (Figs. 37 A, 38 A) with 8 (rarely 9) macrosetae; 9th tergite with 4 macrosetae;
10th tergite with posterior margin slightly rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 37 A) with posterior margin truncate, its truncate apex dentate; 8thsternite (Fig. 37 B) with 17–19 macrosetae; 9th sternite with posterior margin slightly convex
medially. Aedeagus (Figs. 37 C, 37 D) somewhat gourd-shaped in ventral view; apical lobe almost straight and pointed in lateral view, narrowed at base and rounded at apex in ventral view;
basal ridge small and very weakly convex, invisible in lateral view; copulatory piece of inner sac
as in Fig. 37 E. Paramere with apical lobe of paramerite parallel-sided, rounded apically, about
1/4 as long as condylite.
Female: Eighth tergite (Fig. 38 A) with posterior margin slightly emarginateand dentate
around emarginate apex; 8th sternite (Fig. 38 B) with 14–15 macrosetae; sensory setae of 8th
sternite generalised, similar to other setae in shape. Spermatheca (Fig. 38 C) curved three times;
basal part somewhat longer than apical part, acutely curved near base; apical part gently curved,
its inner wall densely wrinkled except around basal 4/5.
Measurements. BL, 4.8–5.7; FBL, 2.1–2.3; HL, 0.70–0.72; HW, 0.78–0.81; EL, 0.240–
0.249; AL, 1.73–1.98; PL, 0.79–0.83; PW, 1.12–1.21; ELL, 0.85–0.92; ELW, 1.23–1.31; HTL,
1.00–1.03.
Bionomics. Pella socia is commonly collected from colonies of Lasius (Dendrolasius) spp.
and distributed mountainous zones in Hokkaidô, Honshû, Shikoku and Kyûshû.
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) nipponensis, L. (D.)
orientalis, L. (D.) capitatus.
Pella rambouseki (Bernhauer, 1929), n. comb.
(Figs. 39–41)
Zyras rambouseki Bernhauer, 1929: 192 (original description).—Scheerpeltz, 1934: 1658 (Zyras, subgenus status not
specified, list).— Dvořák, 1981: 54 (Zyras (Pella), key).—Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntypes: Russia (Far East): 1?, “?/As. or. 919 Vladivostok Dr.
Jureček/Zyras rambouseki Brh. det. Dr. Rambousek/Dr. Rambousek comparavit. 31. V. 1929/ex
coll., Scheerpeltz/COTYPUS Zyras Rambouseki Bernhauer.” (NHMW); 1?, “Sibir. or. Ussuri
Vladivostok Dr. Jureček 1919/COTYPUS/Zyras Rambouseki Bh./rambouseki Brnh*” (ZMHB).
Type locality. Vladivostok, Russia.
Other material. Korea: 1 ex., Jeonglyeon-chi, San’nae-myeon, Cheonlabuk-do, 12 VII
1991, S. Nomura; 5 exs., Janggoksa, Chilgab-san, Cheongyang-gun, Chungnam, 23 VI 2000, M.
Maruyama (LDFJ) (cMar, CNU); 2 exs., Oso-san, Sangdam-ri, Kwangchon-up, Hongsung-gun,
Chungnam, 19–21 VI 1999, U.-S. Hwang & H.-J. Kim (pit) (cMar, CNU); 2 exs., Donghwag-sa,
Gyeryongsan, Chungnam, 12 VI 1999, U.-S. Hwang (CNU). Russia (Far East): 5 exs., 70 km E
Vladivostok, Anisimovka, Khabarovskyi Krai, 5 VI 1993, L. Zerche (LDO) (DEI); 2 exs.,“Vladivostok, Sibri. or., Ussuria, Asia or., IV. 1919, Št. Jureček” (cDvor); 1/, Prwnaja-

Palearctic Species of Pella

Fig. 39.

67

Pella rambouseki (Bernhauer). —A, Fore body, dorsal view; B, right antenna.

Rjetschkam Tales, Vladivostok, 1918–1920, H. Farieb (NHMW); 4 exs., Kaimanovka, Ussuryisk,
Primorskyi Krai, 27 V 2005, M. Maruyama (LDFJ); 2 exs., Kamenushka, Ussuryisk, Primorskyi
Krai, 22 VII 1990, A. Putz; 1 ex., Venedivnovo, Tschermye Gory, Primorskyi Krai, 1–3 VIII
1990, A. Putz; 2 exs., Bukhta Vityaz, Poluostrov Gamov, Khasanskyi, Primorskyi Krai, 30 V
2005, M. Maruyama (LDFJ); 5 exs., same data but, 31 V 2005, M. Maruyama (LDO).
Distribution. Manchurian subregion. ASIA: Korea, Russia (Far East).
Diagnosis. Pella rambouseki is similar to P. comes, P. kidaorum and P. cooterorum in general appearance and colour but easily distinguished from the latter three by the smaller body andthe basal part of spermatheca very long. These four species are locally isolated. Among the sympatric species, P. rambouseki is somewhat similar to P. jureceki in general appearance but distinguished from it by the spermathecal structure above, the body more shining, the longer antennae,
the narrower pronotum and the apical lobe of aedeagal median lobe without a pair of projectionsin the ventro-lateral areas.
Redescription. Body slender. Black in ground colour; antennae, mouthparts, legs reddish
brown to brown. Head (Fig. 39 A) widest just behind eyes; surface smooth, moderately covered
with setae; setae short, as long as those on pronotum and elytra; length of eyes 0.31–0.33 times
as long as head width. Antennae (Fig. 39 B) almost as long as head, pronotum and elytra combined; 1st segment slightly shorter than 2nd and 3rd combined; 2nd segment about 5/7 as long as
3rd; 3rd segment about 3/5 as long as 1st; 4th to 9th segments almost as long as wide; 10th segment slightly wider than long; 11th segment slightly shorter than 1st; approximate relative
lengths of segments from basal to apical: 28.5 : 12.5 : 17 : 10 : 10 : 10 : 10 : 10 : 10 : 11 : 25.5.
Pronotum (Fig. 39 A) 1.30–1.33 times as wide as long, widest around anterior 1/6, narrowed pos-68

M. Maruyama

Fig. 40. Pella rambouseki (Bernhauer). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

teriorly; postero-lateral corner rounded; surface finely punctured, densely covered with setae,
with 5 or 6 macrosetae along lateral margin. Scutellum with surface finely punctured, densely
covered with setae. Elytra (Fig. 39 A) subparallel-sided; surface finely punctured, densely covered
with setae, with 6 or 7 small macrosetae laterally. Legs very long; hind tibia 1.26–1.31 times aslong as elytra; approximate relative lengths of tarsal segments from basal to apical: 11 : 7.5 : 7 : 14
in fore tarsus; 18 : 9 : 7 : 6.5 : 11 in mid tarsus; 26 : 11 : 9 : 8 : 11 in hind tarsus. Abdomen expanded, 1.12 times wider than elytra, widest around 5th and 6th segments; surface smooth; 3rd to 6th
tergites glabrous except for a row of setae along each posterior margin; 7th tergite almost
glabrous except posterior and lateral areas; 8th tergite (Figs. 40 A, 41 A) with 8 (rarely 7 or 9)
macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior margin rounded, with 4macrosetae.
Male: Eighth tergite (Fig. 40 A) with posterior margin truncate, its truncate apex weakly den-Palearctic Species of Pella

69

Fig. 41. Pella rambouseki (Bernhauer). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

tate; 8th sternite (Fig. 40 B) with 18–20 macrosetae; 9th sternite with posterior margin slightly
convex medially. Aedeagus (Figs. 40 C, 40 D) somewhat gourd-shaped in ventral view; apicallobe slightly curved ventrad and pointed apically in lateral view, narrowed at base and somewhat
rounded at apex in ventral view; basal ridge weakly convex; copulatory piece of inner sac as inFig. 40 E. Paramere with apical lobe of paramerite parallel-sided, rounded apically, about 1/3 aslong as condylite.
Female: Eighth tergite (Fig. 41 A) with posterior margin truncate, its truncate apex crenate;
8th sternite (Fig. 41 B) with 10–12 macrosetae; sensory setae of 8th sternite spindle-shaped andflattened. Spermatheca (Fig. 41 C) anfractuous; basal part long, more than five times as long asapical part, acutely curved at middle; apical part acutely curved near apex, its inner wall densely
wrinkled from apex to around apical 2/3.
Measurements. BL, 4.6–5.1; FBL, 2.1–2.4; HL, 0.66–0.73; HW, 0.73–0.80; EL, 0.244–
0.58; AL, 1.80–2.01; PL, 0.73–0.79; PW, 0.96–1.03; ELL, 0.84–0.92; ELW, 1.05–1.13; HTL,
1.06–1.21.
Bionomics. Pella rambouseki is collected from colonies of Lasius (Dendrolasius) spp. This
species is commonly collected in lowlands to mountainous zones in Korea and the Russian Far
East, but much less frequent than a sympatric species P. jureceki.
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) nipponensis, L. (D.)
orientalis.

70

M. Maruyama

Pella jureceki (Dvořák, 1981), n. comb.
(Figs. 42–44)
Zyras (Pella) jureceki Dvořák, 1981: 55 (original description).—Smetana, 2004: 461 (Zyras (Pella), list).
Zyras (Pella) micropterus Pace, 1998a: 976 (original description).—Smetana, 2004: 461 (Zyras (Pella), list). N. syn.

Type material. Zyras (Pella) jureceki. Paratypes: Russia (Far East): 1?, “Asia or., Ussuria, Sibir. or. 4/1919 VLADIVOSTOK Dr. Št. Jureček lgt./PARATYPUS/Zyras (Pella) jureceki
Dvořák ,81” (cDvor); 1/, same but “ALLOTYPES” (cDvor); 1?, “Sibir. or. Ussuri Vladivostok
Dr. Jureček 1919/PARATYPUS/Zyras (Pella) jureceki Dvořák ,81” (NMP).
Zyras (Pella) micropterus. Holotype: China (Beijing Shi): ?, “CHINA Beijing [Xiaolongmen, Beijing Shi, China (according to the original description)], 1100–1500 m, 1. VII. 1993, G.
de Rougemont/Zyras micropterus sp. n. det. R. Pace 1995/HOLOTYPUS Zyras micropterus m.
det. R. Pace 1995” (MHNG).
Type locality. Vladivostok, Primorskyi Krai, Russia.
Other material. China: 4??, 2// , 77 exs., Xiaolongmen (alt. 1400 m), Dongling, Yanshan, Beijing, 15–16 VI 2001, P. Hlaváč & J. Cooter (LDFJ) (IZAS, cMar, cHlav). Korea: 19
exs., Janggoksa, Chilgab-san, Cheongyang-gun, Chungnam, 23 VI 2000, M. Maruyama (LDFJ)(cMar, CNU). Russia (Far East): 8 exs., 10 km N Boitsovo, 20 km N Bikin (alt. 200 m),
Khabarovskyi Krai, 27 V 1993, L. Zerche (LDN); 1 ex., 10 km SW Samarka, 70 km N
Chuguyevka (alt. 300 m), Gordeyevskaya, 29 V 1993, L. Zerche; 28 exs., Kaimanovka, Ussuryisk, Primorskyi Krai, 27 V 2005, M. Maruyama (LDFJ); 5 exs., same data but, 28 V 2005,
M. Maruyama (LDN); 3 exs., same data but, 28 V 2005, M. Maruyama (LDO); 4 exs., Ka-

Fig. 42.

Pella jureceki (Dvořák). — A, Fore body, dorsal view; B, right antenna.

Palearctic Species of Pella

71

Fig. 43. Pella jureceki (Dvořák). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

menushka, Ussuryisk, Primorskyi Krai, 28 V 2005, M. Maruyama (LDFJ); 13 exs., Bukhta
Vityaz, Poluostrov Gamov, Khasanskyi, Primorskyi Krai, 29 V 2005, M. Maruyama (LDFJ); 14
exs., same data but, 30 V 2005; 1 ex., same data but, 31 V 2005, M. Maruyama (LDO); 80 exs.,
same data but, 31 V 2005, M. Maruyama (LDFJ); 42 exs., same data but, 31 V 2005, M.
Maruyama (LDN); 1 ex., Kravtzovka, Nadezhdinskyi, Primorskyi Krai, M. Maruyama (LDFJ); 1
ex., Okeanskaya,Vladivostok, Primorskyi Krai, 1 VI 2005, M. Maruyama (LDN); 1 ex., Arsenev,
Primorskyi Krai, VII 1991, Stiba.
Distribution. Manchurian subregion. ASIA: China (Beijing Shi), Korea, Russia (Far East).
Diagnosis. Pella jureceki may be easily distinguished from the other species of the funesta
group by the somewhat paler colour and the apical lobe of aedeagal median lobe with a pair ofprojections in the ventro-lateral areas. Among the sympatric species in China and the Russian Far

72

M. Maruyama

Fig. 44. Pella jureceki (Dvořák). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, D, spermathecae. Female.

East, Pella jureceki is similar to P. rambouseki, P. cooterorum and P. hlavaci in general appearance, but is distinguished from P. rambouseki by the shorter antennae and the shorter spermatheca, from P. cooterorum by the diagnostic character states above, the less shining body surface, thewider pronotum and the outer projection of apical part of spermatheca smaller, and from P.
hlavaci by the above spermathecal structure, the presence of pronotal postero-lateral corner andthe shorter and apically dilated apical part of spermatheca. In the other regions, Korea and theRussian Far East, Pella jureceki is somewhat similar to P. rambouseki in general appearance but
distinguished from the latter by the aedeagal structure diagnosed above, the paler colour, thewider pronotum and the shorter basal part of spermatheca.
Redescription. Body slender. Blackish brown in ground colour; antennae, mouthparts, legs
and abdomen reddish brown to brown; around posterior margin of 3rd to 5th abdominal segments
pale brown. Head (Fig. 42 A) widest just behind eyes; surface finely reticulated, densely covered
with setae; setae short, as long as those on pronotum and elytra; length of eyes 0.32–0.33 times
as long as head width. Antennae (Fig. 42 B) shorter than head, pronotum and elytra combined;
1st segment slightly shorter than 2nd and 3rd combined; 2nd segment about 5/7 as long as 3rd;
3rd segment about 3/5 as long as 1st; 4th to 9th segments almost as long as wide; 10th segment
slightly wider than long; 11th segment much shorter than 1st; approximate relative lengths ofsegments from basal to apical: 29 : 11 : 19.5 : 7.5 : 7.5 : 7.5 : 7.5 : 7 : 7.5 : 8 : 18.5. Pronotum (Fig.
42 A) 1.38–1.41 times as wide as long, widest around anterior 1/4, narrowed posteriorly; posterolateral corner rounded; surface roughly reticulated among punctures, densely covered with setae,

Palearctic Species of Pella73

with 3 or 4 small macrosetae along lateral margin. Scutellum with surface finely punctured,densely covered with setae. Elytra (Fig. 42 A) subparallel-sided; surface finely punctured, reticulated among punctures, densely covered with setae, with 6 or 7 small macrosetae laterally;
macrosetae poorly differentiated from setae. Legs long; hind tibia 1.22–1.25 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 9.5 : 7 : 6 : 10.5 in foretarsus; 13.5 : 8 : 8 : 7 : 10 in mid tarsus; 22 : 10 : 9 : 7.5 : 13.5 in hind tarsus. Abdomen expanded,
1.09 times wider than elytra, widest around 4th and 5th segments; surface weakly reticulated; 3rdsparsely covered with setae; 4th to 6th tergites glabrous except for a row of setae along each posterior margin; 7th tergite sparsely covered with minute setae; 8th tergite (Figs. 43 A, 44 A) with 8(rarely 9) macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior margin slightly
emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 43 A) with posterior margin truncate, its truncate apex dentate; 8thsternite (Fig. 43 B) with 13–15 macrosetae; 9th sternite with posterior margin slightly emarginate. Aedeagus (Figs. 43 C, 43 D) amygdaloid in ventral view; apical lobe slightly curved ventrad and pointed apically in lateral view, subparallel-sided in basal 1/2 and pointed at apex inventral view, its ventrolateral sides with a pair of projections; basal ridge weakly convex; copulatory piece of inner sac as in Fig. 43 E. Paramere with apical lobe of paramerite parallel-sided,rounded apically, about 1/2 as long as condylite.
Female: Eighth tergite (Fig. 44 A) with posterior margin truncate, its truncate apex dentate;
8th sternite (Fig. 44 B) with 13–15 macrosetae; sensory setae of 8th sternite spindle-shaped andflattened. Spermatheca (Figs. 44 C, 44 D) curved twice, S-shaped; basal part about 2–2.5 times aslong as apical part, acutely curved around basal 1/3 and gently curved apicad; apical part almost
straight, its inner wall densely wrinkled from apex to around apical 1/3.
Measurements. BL, 4.5–5.6; FBL, 2.1–2.3; HL, 0.69–0.73; HW, 0.78–0.81; EL, 0.254–
0.261; AL, 1.80–1.97; PL, 0.73–0.83; PW, 1.03–1.14; ELL, 0.76–0.88; ELW, 1.11–1.25; HTL,
0.95–1.08.
Bionomics. Pella jureceki is very common in its distributional range.
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) nipponensis, L. (D.) orientalis.
Pella kidaorum Maruyama, n. sp.
(Figs. 45–47)

Type material. Holotype: Japan: ?, “Taihei, Maruseppu-chô, Hokkaidô, Japan, 29–31 V
2000, Y. Kida leg.” (LDFJ) (HUM).
Paratypes: Japan: Hokkaidô: 1 ex., Himenuma, Rishiri-tô, 9 VII 1977, S. Naomi; 63 exs.,
same data but, 4 IX 1990, T. Kishimoto (pt); 99 exs., Hebi-numa, Teshio-gawa, Teshio-chô, 9 VII
1992, S. Hori (pt); 5 exs., Esaushi-yama, Kamikawa-chô, 8 VIII 1985, N. Yasuda (pt); 1 ex.,
Mukôengaru, Engaru-chô, 25 VI 2000, Y. Kida; 4 exs., Makoi, Shari-chô, 28 VI 2000, Y. Kida
(LDFJ); 7 exs., same data but, 27–29 V 2001; 1 ex., Nosaki, Bihoro-chô, 23 VI 2001, Y. Yamaga(LDC ); 1 ex., Kashiwagaoka, Bihoro-chô, 23 VI 2001, Y. Yamaga; 3 exs., Ochiishi, Nemuro-shi,
24–27 VIII 1993, K. Miyashita; 4 exs., Kombumori, Nemuro-shi, 26 IX 2000, H. Sugaya; 8 exs.,
Tôbaru, Nemuro-shi, 28 IX 2000, H. Sugaya (LDFJ); 38 exs., Taihei, Maruseppu-chô, 29–31 V
2000, Y. Kida (LDFJ); 7 exs., same data but, 2–3 VI 2000; 8 exs., same data but, 16–17 VI 2000;
1 ex., same data but, 11–12 VIII 2000; 2 exs., same data but, 17 VIII 2000; 1 ex., same data but,
19–21 VIII 2000; 29 exs., same data but, 25 VIII 2000, M. Maruyama; 2 exs., Onnenai, Tsuruimura, 2 VI 1994, Y. Hirama; 9 exs., Midorigaoka-kôen, Kitami-shi, 17 VI 2001, M. Maruyama;

74

M. Maruyama

Fig. 45.

Pella kidaorum Maruyama, n. sp. —A, Fore body, dorsal view; B, right antenna.

4 exs., Kamiotoineppu, Otoineppu-mura, 15–23 VII 1997, S. Hori (pt); 3 exs., Kaitaku-kinenkan,
Nopporo-shinrin-kôen, Ebetsu-shi, 20 IV 2003, S. Hori (LDFJ); 2 exs., Ôsawaguchi, Nopporoshinrin-kôen, Ebetsu-shi, 31 V 1999, M. Maruyama (LDFJ); 23 exs., Shibumbetsu, Nopporoshinrin-kôen, Ebetsu-shi, 11 VI 2000, M. Maruyama (LDFJ); 1 ex., same data but, 18 VI 2000;
14 exs., Tomambetsu, Nopporo-shinrin-kôen, Ebetsu-shi, 4 VI 2000, M. Maruyama (LDS); 3
exs., same data but, 11 VI 2000; 1 ex., same data but, 23 IX 2000, H. Sugaya; 3 exs., Hitsujigaoka, Sapporo-shi, 25 VII 1998, M. Maruyama (LDS); 9 exs., same data but, 21 VI 1998 (LDO);
10 exs., Hyakumatsu-zawa, Sapporo-shi, 24 VIII 2001, M. Maruyama & H. Sugaya (LDO); 3
exs., Kannon-zawa, Sapporo-shi, 31 V 2002, M. Maruyama (LDFJ); 61 exs., same data but, 7
VIII 2002; 1 ex., Moiwa-yama, Sapporo-shi, 16 VI 1983, Y. Hirano; 1 ex., Maru-yama, Sapporoshi, 6 VI 1998, M. Maruyama (LDFJ); 1/, “Sapporo [Sapporo-shi]”, 5–16 VIII 1880, G. Lewis(“Myrmedonia, comes var., Sapporo. Japan Lewis”, written on paper card specimen glued on)(BMNH); 6 exs., Asari, Otaru-shi, 1 IX 1998, K. Sugisima (LDFJ); 17 exs., Yutoku, Ôtaki-mura,
24 VIII 2001, M. Maruyama & H. Sugaya (LDO).
Type locality. Maruseppu-chô, Hokkaidô, Japan.
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô). This species is most probably confined to Hokkaidô, Japan.
Etymology. Dedicated to Mr. Yasunari Kida (Maruseppu Town Insectarium, Hokkaidô,Japan) and his wife Mrs. Yuriko Yamaga (Bihoro Museum, Hokkaidô, Japan) for their contribution to the study on ants and myrmecophilous insects in eastern Hokkaidô.
Diagnosis. Pella kidaorum is most similar to P. cooterorum in general appearance andPalearctic Species of Pella

75

Fig. 46. Pella kidaorum Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

shape of genitalia, but is distinguished from the latter by the apical lobe of aedeagus slightly
curved and the spermatheca lacking outer projection. Among the sympatric species Pella kidaorum is most similar to P. socia in general appearance but distinguished from the latter by the
pronotal setation denser, the pronotal corners not distinctly angled and the apical lobe of aedeagal median lobe slightly sinuate.
Discription. Body slender. Black in ground colour; antennae, mouthparts and legs reddish
brown to brown; around posterior margins of 3rd to 5th abdominal segments pale brown. Head
(Fig. 45 A) widest just behind eyes; surface finely reticulated, densely covered with setae; setae
short, as long as those on pronotum and elytra; length of eyes 0.29–0.30 times as long as headwidth. Antennae (Fig. 45 B) shorter than head, pronotum and elytra combined; 1st segment shorter than 2nd and 3rd combined; 2nd segment about 5/8 as long as 3rd; 3rd segment about 2/3 aslong as 1st; 4th segment slightly longer than wide; 5th and 6th segments as long as wide, 7th to
9th segments almost as long as wide; 10th segment slightly wider than long; 11th segment much

76

M. Maruyama

Fig. 47. Pella kidaorum Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, D, spermathecae. Female.

shorter than 1st; approximate relative lengths of segments from basal to apical: 28 : 12.5 : 17 : 11 :10 : 10 : 10 : 9.5 : 9 : 10.5 : 20.5. Pronotum (Fig. 45 A) 1.25–1.30 times as wide as long, widest
around anterior 1/4, narrowed posteriorly; postero-lateral corner rounded; surface finely reticulated among punctures, densely covered with setae, with 7 or 8 small macrosetae along lateral margin. Scutellum with surface finely punctured, densely covered with setae. Elytra (Fig. 45 A) subparallel-sided; surface finely punctured, reticulated among punctures, densely covered with setae,
with 6 or 7 small macrosetae laterally. Legs very long; hind tibia 1.27–1.35 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 11. : 7.5 : 6.5 : 10 in foretarsus; 19 : 10.5 : 8 : 6 : 10 in mid tarsus; 29 : 10 : 9 : 7.5 : 11 in hind tarsus. Abdomen expanded,
1.07 times wider than elytra, widest around 4th and 5th segments; surface weakly reticulated; 3rdsparsely covered with setae; 4th to 6th tergites glabrous except for a row of setae along each posterior margin; 7th tergite glabrous except for lateral areas sparsely covered with minute setae; 8th
tergite (Figs. 46 A, 47 A) with 10 (rarely 9 or 11) macrosetae; 9th tergite with 4 macrosetae; 10th
tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 46 A) with posterior margin truncate, its truncate apex weakly crenate; 8th sternite (Fig. 46 B) with 23–25 macrosetae; 9th sternite with posterior margin roundly
emarginate. Aedeagus (Figs. 46 C, 46 D) tear shaped in ventral view; apical lobe slightly sinuate
in ventral view, pointed at apex in lateral and ventral views; basal ridge well developed, reachingPalearctic Species of Pella77

bottom of basal capsule; copulatory piece of inner sac as in Fig. 46 E. Paramere with apical lobeof paramerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 47 A) with posterior margin truncate, its truncate apex weakly
crenate; 8th sternite (Fig. 47 B) with 12–14 macrosetae; sensory setae of 8th sternite spindleshaped and flattened. Spermatheca (Figs. 47 C, 47 D) curved twice, S-shaped; basal part about
2–2.5 times as long as apical part, acutely curved around basal 1/3 and gently curved apicad; apical part almost straight, its inner wall densely wrinkled from apex to around apical 1/3.
Measurements. BL, 4.9–5.5; FBL, 2.3–2.5; HL, 0.73–0.74; HW, 0.83–0.88; EL, 0.244–
0.257; AL, 2.08–2.15; PL, 0.88–0.92; PW, 1.12–1.19; ELL, 0.85–0.98; ELW, 1.15–1.35; HTL,
1.20–1.25.
Bionomics. Pella kidaorum is commonly collected from colonies of Lasius (Dendrolasius)
spp. and widely distributed in Hokkaidô. A specimen of Pella kidaorum was collected near a nestof Formica japonica once for all (Kida, pers. comm.), but this association is probably accidental.
Symbiotic host. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) nipponensis, L. (D.)
orientalis, L. (D.) capitatus.
Pella cooterorum Maruyama, n. sp.
(Figs. 48–50)

Type material. Holotype: China (Beijing Shi): ?, “Xiaolongmen (1400 m), Dongling
Yan-shan Beijing CHINA, 15–16 VI 2001, Hlaváč & Cooter leg.” (LDFJ) (IZAS).
Paratypes: 4??, 2// , 13 exs., same data as holotype (IZAS, cMar, cHlav); China (Yunnan
Sheng): 5 exs., Diaolin, 100 km W. Kunming, 2–22 VI 1993, E. Jendek & O. Šauša (NHMW,cMar).
Type locality. Xiaolongmen, Beijing Shi, China.
Distribution. Manchurian subregion. ASIA: China (Beijing Shi, Yunnan Sheng).
Etymology. Dedicated to Mr. Jonathan Cooter (Hereford, United Kingdom), a collector of
the type series, and his family.
Diagnosis. Pella cooterorum is most similar to P. kidaorum in general appearance andshape of genitalia, but is distinguished from the latter by the apical lobe of aedeagus almost
straight and the spermatheca having outer projection. Among the sympatric species Pella
cooterorum is somewhat similar to P. jureceki in general appearance but distinguished from thelatter by the darker colour, the body surface more shining, the narrower pronotum, the apical lobeof aedeagal median lobe without a pair of projections on the ventro-lateral areas and the outerprojection of apical part of spermatheca larger.
Description. Body slender. Black in ground colour; antennae, mouthparts and legs reddish
brown to brown; around posterior margin of 3rd to 5th abdominal segments pale brown. Head
(Fig. 48 A) widest just behind eyes; surface finely reticulated, densely covered with setae; setae
short, as long as those on pronotum and elytra; length of eyes 0.30–0.32 times as long as headwidth. Antennae (Fig. 48 B) shorter than head, pronotum and elytra combined; 1st segment shorter than 2nd and 3rd combined; 2nd segment about 5/8 as long as 3rd; 3rd segment about 2/3 aslong as 1st; 4th segment slightly longer than wide; 5th and 6th segments as long as wide, 7th to
9th segments almost as long as wide; 10th segment slightly wider than long; 11th segment much
shorter than 1st; approximate relative lengths of segments from basal to apical: 30 : 12.5 : 16 : 9 :9 : 8 : 9 : 9 : 10 : 10.5 : 22. Pronotum (Fig. 48 A) 1.23–1.30 times as wide as long, widest around
anterior 1/5, narrowed posteriorly; postero-lateral corner rounded; surface finely punctured, finely

78

M. Maruyama

Fig. 48.

Pella cooterorum Maruyama, n. sp.—A, Fore body, dorsal view; B, right antenna.

reticulated among punctures, densely covered with setae, with 7 or 8 small macrosetae along lateral margin. Scutellum with surface finely punctured, densely covered with setae. Elytra (Fig.
48 A) subparallel-sided; surface finely punctured, reticulated among punctures, densely covered
with setae, with 6 or 7 small macrosetae laterally. Legs very long; hind tibia 1.35–1.36 times aslong as elytra; approximate relative lengths of tarsal segments from basal to apical: 9 : 8 : 6 : 13 infore tarsus; 18 : 9 : 8 : 7 : 12 in mid tarsus; 22.5 : 12 : 10 : 8.5 : 12.5 in hind tarsus. Abdomen expanded, 1.09 times wider than elytra, widest around 6th and 7th segments; surface weakly reticulated; 3rd sparsely covered with setae; 4th to 6th tergites glabrous except for a row of setae alongeach posterior margin; 7th tergite glabrous except for lateral areas sparsely covered with minutesetae; 8th tergite (Figs. 49 A, 50 A) with 10 (rarely 9 or 11) macrosetae; 9th tergite with 4macrosetae; 10th tergite with posterior margin slightly rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 49 A) with posterior margin truncate, its truncate apex weakly crenate; 8th sternite (Fig. 49 B) with 20–23 macrosetae; 9th sternite with posterior margin slightly
emarginate. Aedeagus (Figs. 49 C, 49 D) amygdaloid in ventral view; apical lobe almost straight,
pointed at apex in lateral and ventral views; basal ridge well developed, reaching near bottom ofbasal capsule; copulatory piece of inner sac as in Fig. 49 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 50 A) with posterior margin somewhat truncate, its truncate apex
weakly crenate; 8th sternite (Fig. 50 B) with 12–14 macrosetae; sensory setae of 8th sternite spindle-shaped and flattened. Spermatheca (Figs. 50 C–50 E) curved three times; basal part about
1.5–2.0 times as long as apical part, acutely curved around basal 1/3 and gently curved apicad;
apical part with outer projection, slightly curved, its inner wall densely wrinkled from apex toPalearctic Species of Pella

79

Fig. 49. Pella cooterorum Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal
view. Male.

around apical 1/2.
Measurements. BL, 5.0–5.2; FBL, 2.3–2.5; HL, 0.70–0.73; HW, 0.78–0.80; EL,
0.243–0.256; AL, 2.02–2.05; PL, 0.81–0.83; PW, 1.03–1.08; ELL, 0.84–0.90; ELW, 1.12–1.25;
HTL, 1.13–1.23.
Bionomics. The type series of Pella cooterorum from Beijing Shi were collected by sifting
dead leaves accumulated around a colony of Lasius (Dendrolasius) fuji nested at a hollow of oldtree. The type series were captured with the myrmecophilous staphylinid beetles Pella jureceki, P.
hlavaci, P. beijingorum, P. zhoui, Zyras (Zyras) sp., Homoeusa chinensis Pace, 1999, Thiasophilasp., Batristilbus monstrotibialis Hlaváć, Sugaya & Zhou, 2002, and Tmesiphorus sp. (Hlaváč,pers. comm.).
Symbiotic host. Lasius (Dendrolasius) fuji.

80

M. Maruyama

Fig. 50. Pella cooterorum Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C–E, spermathecae. Female.

The barbara GroupSpecies included. Pella barbara, P. leonhardi, P. jelineki.
Distribution. Mediterranean subregion.
Diagnosis. Species of the barbara group may be characterised by a combination of the following character states: 1) eye length 0.29–0.35 times as long as head width; 2) 11th antennal
segment almost as long as, or longer than 1st; 3) pronotum with postero-lateral corners; 4) pronotum 1.25–1.35 times as wide as long; 5) pronotum widest anteriorly; 6) pronotal hypomeron fully
visible in lateral view; 7) elytra almost unicoloured, blackish brown; 8) inner and posterior margins of elytra weakly margined; 9) male 8th sternite longer than in female; 10) posterior marginof 8th tergite crenate; 11) posterior margin of male 8th sternite rounded; 12) lateral projection of
apodeme of male 8th tergite not evidently longer than that of female; 13) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior margin of the segment; 14) postero-medial margin of female 8th sternite with out minute pubescence; 15) apical lobe of aedeagal median lobe gently curved ventrad in lateral view; 16) distal crests of aedeagus well developed, projected semicircularly in lateral view; 17) copulatory piece of aedeagal median lobe spindle-shaped in dorsal view.

Palearctic Species of Pella

81

Comments. The barbara group may possibly be allied to the funesta group in resemblanceof general appearance. However, the members of this species-group are also similar to Pella erratica in general appearance, small body and shape of spermatheca, and the members of the excepta group in general structure of aedeagal median lobe.
Symbiotic host. Unknown.
Key to the Species of the barbara Group
1. Third to 6th tergites densely covered with setae, their setae overlapping each other; 8th abdominal tergite with more than 10 macrosetae in female. ......................................P. barbara.
— Third to 6th tergites moderately to sparsely covered with setae, their setae not overlapping
each other; 8th abdominal tergite with less than 8 macrosetae in female. ............................2.
2. Postero-lateral corners of pronotum well angled; pronotum and elytra densely covered withsetae as in Fig. 54 A. ............................................................................................P. leonhardi.
— Postero-lateral corners of pronotum rounded; pronotum and elytra strongly densely covered
with setae as in Fig. 56 A. ........................................................................................P. jelineki.
Pella barbara (Fairmaire, 1863), n. comb.
(Figs. 51–53)
Myrmedonia barbara Fairmaire, 1863: 638 (original description).—Bernhauer & Scheerpeltz, 1926: 695 (Zyras (Pella),list).—Fenyes, 1920: 297 (Zyras (Myrmedonia s. str.), list).—Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntype: Algeria: sex?, “barbara Fairm type/R.I.Sc.N.B. 17.819 Coll. A.

Fig. 51.

Pella barbara (Fairmaire). — A, Fore body, dorsal view; B, right antenna.

82

M. Maruyama

Fig. 52. Pella barbara (Fairmaire). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

Fauvel M. barbara” (abdomen missing) (IRSNB).
Other material. Algeria: 1?, “Bône” (NHMW); 1?, same data (IRSNB); 1?, “Bône
Philipperille/Philipperille à Horn (IRSNB); 1/, “Philipperille” (IRSNB); 1/, “Oran) Tlemcan
Dr. Normand-3” (IRSNB); 1/, “Sud-algèrien Bou Saada 25/29–v–1954 G. Fagel/G. Fagel barbarus Fairm.” (IRSNB); 2??, 1/, no data (IRSNB).
Type locality. Bône, Algeria.
Distribution. Mediterranean subregion. NORTH AFRICA: Algeria.
Diagnosis. Pella barbara is similar to P. leonhardi and P. jelineki in general appearance,
but may be easily distinguished from them by the 3rd to 6th abdominal tergites densely covered
with setae. This species is somewhat similar to the species of the funesta group in general appearance and colour but distinguished from them by the smaller body and the 11th antennal segment almost as long as, or longer than the 1st. The distribution is overlaps with that of Pella er-

Palearctic Species of Pella

83

Fig. 53. Pella barbara (Fairmaire). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

ratica, but P. barbara may be easily distinguished from P. erratica by the 3rd to 6th tergite densely covered with setae and the length of the 11th antennal segment mentioned above.
Redescription. Body slender. Blackish brown in ground colour; antennae, mouthparts andlegs reddish brown. Head (Fig. 51 A) slightly depressed above, wider than long; surface smooth,moderately covered with setae; setae rather long, as long as those of elytra in length; length ofeyes 0.29–0.30 times as long as head width. Antennae (Fig. 51 B) shorter than head, pronotumand elytra combined; 1st segment long, longer than 2nd and 3rd combined; 2nd segment about
4/5 as long as 3rd; 3rd segment about 1/3 as long as 1st; 4th to 10th segments slightly wider thanlong; 11th segment almost as long as 1st; approximate relative lengths of segments from basal toapical: 24 : 8 : 6.5 : 7 : 8 : 8.5 : 8 : 8.5 : 8 : 9 : 23.5. Pronotum (Fig. 51 A) 1.30–1.35 times as wide as
long, widest around anterior 1/5, constricted posteriorly; lateral margins subparallel-sided nearbase; postero-lateral corner weakly angled; surface densely covered with setae, with 7 or 8
minute macrosetae along lateral margin. Scutellum with surface finely punctured, moderately
covered with setae. Elytra (Fig. 51 A) slightly broadened posteriorly; surface finely punctured,densely covered with setae, with 2 or 3 small macrosetae laterally. Legs rather long; hind tibia
1.09–1.10 as long as elytra; approximate relative lengths of tarsal segments from basal to apical:
6 : 5.5 : 5.5 : 10 in fore tarsus; 9.5 : 6.5 : 5 : 4.5 : 10 in mid tarsus; 19 : 9.5 : 8 : 8 : 11 in hind tarsus.
Abdomen subparallel-sided, slightly wider than elytra, widest around 5th segment; surface moderately punctured; 3rd to 6th tergites densely covered with setae; 7th tergite moderately covered
with minute setae; 8th tergite (Figs. 52 A, 53 A) with 6–11 macrosetae; 9th tergite with 6
macrosetae; 10th tergite with posterior margin rounded, with 5 macrosetae.

84

M. Maruyama

Male: Eighth tergite (Fig. 52 A) with posterior margin rounded and crenate; 8th sternite (Fig.
52 B) with 23 or 24 macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs.
52 C, 52 D) amygdaloid in ventral view; apical lobe slightly curved ventrad and pointed at apex
in lateral view, slightly narrowed at base and pointed at apex in ventral view; basal ridge well developed, reaching near bottom of basal capsule; copulatory piece of inner sac as in Fig. 52 E.
Paramere with apical lobe of paramerite parallel-sided, rounded apically, as long as condylite.
Female: Eighth tergite (Fig. 53 A) with posterior margin somewhat rounded, its rounded apex
minutely dentate; 8th sternite (Fig. 53 B) with 11 or 12 macrosetae; sensory setae of 8th sternite
generalised, similar to other setae in shape. Spermatheca (Fig. 53 C) curved three times; basal
part about 2.5 times as long as apical part, slightly bulbous around base, acutely curved aroundbasal 2/5; apical part dilated apicad, much broader than basal part, its inner wall densely wrinkledfrom apex to around apical 2/5.
Measurements. BL, 3.7–4.1; FBL, 1.8–1.9; HL, 0.54–0.59; HW, 0.62–0.68; EL, 0.183–
0.198; AL, 1.39–1.60; PL, 0.60–0.69; PW, 0.81–0.89; ELL, 0.74–0.79; ELW, 0.96–1.05; HTL,
0.81–0.86.
Bionomics and symbiotic host. No information.
Pella leonhardi (Bernhauer, 1912), n. comb.
(Figs. 54, 55)
Zyras leonhardi Bernhauer, 1912: 110 (original description).—Fenyes, 1920: 297 (Zyras (Myrmedonia s.str.), list).—Bernhauer & Scheerpeltz, 1926: 700 (Zyras (Pella), list).—Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntype: Italy (Sicilia): 1/, “Sicilia 1906 Monte Cola O. Leonhard 12. 4.

Fig. 54.

Pella lenhardi (Bernhauer). — A, Fore body, dorsal view; B, right antenna.

Palearctic Species of Pella

85

Fig. 55. Pella lenhardi (Bernhauer). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca. Female.

ges./Syntypus/Zyras Leonhardi Bernhauer/Zyras Leonhardi Bh./Coll. DEI Eberswalde/VIDIT R.
Pace 1986” (DEI).
Type locality. Monte Cola, Sicilia, Italy.
Other material. Not examined.
Distribution. Mediterranean subregion. EUROPE: Italy (Sicilia).
Diagnosis. Pella leonhardi is closely similar to P. barbara and P. jelineki in general appearance, but distinguished from P. barbara by the well angled pronotum and the sparser setae on the
3rd to 6th abdominal tergites, from P. jelineki by the well angled pronotum and the sparser setaeon the head, pronotum and elytra. This species is somewhat similar to the species of the funesta
group in general appearance and colour, but is distinguished from the them by the smaller bodyand the 11th antennal segment longer than the 1st. The distribution overlaps with that of Pella erratica, but P. leonhardi may be easily distinguished from P. erratica by the postero-lateral cornersof pronotum well angled and the antennal structure mentioned above.
Redescription. Body slender. Blackish brown in ground colour; antennae, mouthparts andlegs reddish brown. Head (Fig. 54 A) slightly depressed above, wider than long; surface smooth,moderately covered with setae; setae rather long, almost as long as those of elytra in length;
length of eyes 0.37 times as long as head width. Antennae (Fig. 54 B) shorter than head, pronotum and elytra combined; 1st segment long, almost as long as 2nd and 3rd combined; 2nd seg-

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M. Maruyama

ment about 5/6 as long as 3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments slightly
wider than long; 11th segment much longer than 1st; approximate relative lengths of segmentsfrom basal to apical: 19.5 : 9.5 : 11 : 6 : 6 : 7 : 6.5 : 6.5 : 6.5 : 7 : 23.5. Pronotum (Fig. 54 A) 1.25
times as wide as long, widest around anterior 1/4, constricted posteriorly; lateral margins subparallel-sided near base; postero-lateral corner weakly angled; surface densely covered with setae,
with 7 or 8 minute macrosetae along lateral margin. Scutellum with surface finely punctured,
moderately covered with setae. Elytra (Fig. 54A) slightly broadened posteriorly; surface finely
punctured, densely covered with setae, with 2 or 3 small macrosetae laterally. Legs moderate inlength; hind tibia 1.02 times as long as elytra; approximate relative lengths of tarsal segmentsfrom basal to apical: 5.5 : 4.5 : 5 : 8 in fore tarsus; 9.5 : 8 : 5 : 5 : 10 in mid tarsus; 5 : 8 : 6 : 7 : 10 inhind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 5th segment; surface moderately punctured; 3rd to 6th tergites moderately to sparsely covered withsetae; 7th tergite sparsely covered with minute setae; 8th tergite (Fig. 55 A) with 6 or 7 macrosetae; 9th tergite with 6 macrosetae; 10th tergite with posterior margin rounded, with 5 macrosetae.
Male unknown.
Female: Eighth tergite (Fig. 55 A) with posterior margin somewhat truncated, its truncatedapex minutely dentate; 8th sternite (Fig. 55 B) with 8 or 9 macrosetae; sensory setae of 8th sternite generalised, similar to other setae in shape. Spermatheca (Fig. 55 C) curved three times; basal
part about three times as long as apical part, slightly bulbous around base, acutely curved aroundbasal 1/3; apical part dilated apicad, much broader than basal part, its inner wall densely wrinkledfrom apex to around apical 1/2.
Measurements. BL, 3.8; FBL, 1.9; HL, 0.55; HW, 0.64; EL, 0.238; AL, 1.46; PL, 0.67;
PW, 0.84; ELL, 0.82; ELW, 1.05; HTL, 0.84.
Bionomics and symbiotic host. No information.
Pella jelineki Maruyama, n. sp.
(Figs. 56, 57)

Type material. Holotype: Tunisia: /, “Tunis El Kef Dr. Normand/Zyras leonhardi Bernh.
det. M. Dvořák ,71” (NMP).
Type locality. El Kef [Al Kåf], Tunisia.
Distribution. Mediterranean subregion. NORTH AFRICA: Tunisia.
Etymology. Dedicated to Dr. Josef Jelínek (NMP), a curator of the type depository and a
specialist of the Nitidulidae (Coleoptera), for his cooperation in the present study.
Diagnosis. Pella jelineki is closely similar to P. barbara and P. leonhardi in general appearance, but is distinguished from them by the rounded postero-lateral corner of the pronotum, andthe denser setae on the head, pronotum and elytra. This species is somewhat similar to the speciesof the funesta group in general appearance and colour but distinguished from them by the smaller
body and the 11th antennal segment longer than the 1st. The distribution is overlaps with that of
Pella erratica, but P. jelineki may be easily distinguished from P. erratica by the antennal structure above and the denser setae on the head, pronotum and elytra.
Description. Body slender. Pale brown in ground colour; antennae, mouthparts, legs reddish brown. Head (Fig. 56 A) slightly depressed above, wider than long; surface smooth, densely
covered with setae; setae rather short, slightly longer than those of elytra in length; length of eyes
0.35 times as long as head width. Antennae (Fig. 56 B) shorter than head, pronotum and elytra
combined; 1st segment long, almost as long as 2nd and 3rd combined; 2nd segment about 5/6 asPalearctic Species of Pella

Fig. 56.

87

Pella jelineki Maruyama, n. sp. —A, Fore body, dorsal view; B, right antenna.

long as 3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments slightly wider than long;
11th segment somewhat longer than 1st; approximate relative lengths of segments from basal toapical: 17 : 7:5 : 9 : 6 : 6 : 6.5 : 6.5 : 6 : 6 : 6.5 : 19. Pronotum (Fig. 56 A) 1.31 times as wide as long,widest around anterior 1/5, slightly constricted posteriorly; postero-lateral corner rounded; surface very densely covered with setae, with 7 or 8 minute macrosetae along lateral margin. Scutellum with surface finely punctured, moderately covered with setae. Elytra (Fig. 56 A) slightly
broadened posteriorly; surface finely punctured, very densely covered with setae, with 2 or 3
small macrosetae laterally. Legs moderate in length; hind tibia 0.92 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 5.5 : 4.5 : 4.5 : 8.5 in fore tarsus; 7.5 : 6 : 6 : 4.5 : 8.5 in mid tarsus; 12 : 7.5 : 8 : 6.5 : 9.5 in hind tarsus. Abdomen subparallelsided, slightly narrower than elytra, widest around 5th segment; surface moderately punctured;
3rd to 6th tergites moderately covered with setae; 7th tergite sparsely covered with minute setae;
8th tergite (Fig. 57 A) with 7 macrosetae; 9th tergite with 6 macrosetae; 10th tergite with posterior margin rounded, with 5 macrosetae.
Male unknown.
Female: Eighth tergite (Fig. 57 A) with posterior margin truncated, its truncated apex minutely dentate; 8th sternite (Fig. 57 B) with 9 macrosetae; sensory setae of 8th sternite generalised,similar to other setae in shape. Spermatheca (Fig. 57 C) curved three times; basal part about 2.5
times as long as apical part, slightly bulbous around base, acutely curved around basal 2/5; apicalpart dilated apicad, much broader than basal part, its inner wall densely wrinkled from apex toaround apical 4/7.

88

M. Maruyama

Fig. 57. Pella jelineki Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

Measurements. BL, 3.5; FBL, 1.6; HL, 0.52; HW, 0.62; EL, 0.219; AL, 1.38; PL, 0.62;
PW, 0.81; ELL, 0.8; ELW, 0.99; HTL, 0.73.
Bionomics and symbiotic host. No information.
The erratica GroupSpecies included. Pella erratica.
Distribution. European subregion, Mediterranean subregion.
Diagnosis. Species of the erratica group may be characterised by a combination of the following character states: 1) eye length 0.34–0.37 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with postero-lateral corners; 4) pronotum 1.27–1.35
times as wide as long; 5) pronotum widest anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytra unicoloured, brown; 8) inner and posterior margins of elytra weakly margined; 9) male 8th sternite not longer than in female and almost the same in size; 10) posterior margin of 8th tergite crenate; 11) posterior margin of male 8th sternite rounded; 12) lateral projectionof apodeme of male 8th tergite longer than that of female; 13) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior margins of the segment; 14) postero-medial
margin of female 8th sternite without minute pubescence; 15) apical lobe of aedeagal median
lobe gently curved ventrad in lateral view; 16) distal crests of aedeagus well developed, projected
semicircular in lateral view; 17) copulatory piece of aedeagal median lobe almost spindle-shaped
and pointed at apex in dorsal view.
Comments. A monotypical species-group. Though its allied species-group has not beenfound, Pella erratica is similar to the members of the barbara group in the general appearance

Palearctic Species of Pella

89

and the small body, but is distinguished from them by the shorter 11th antennal segment.
Symbiotic host. Tapinoma erraticum.
Pella erratica (Hagens, 1863), n. comb.
(Figs. 58–61)
Myrmedonia erratica Hagens, 1863: 118 (original description).—Reitter, 1909: 42 (Myrmedonia (Pella), key).—Bernhauer & Scheerpeltz, 1926: 697 (Zyras (Pella), list). — Scheerpeltz, 1934: 1653 (Zyras (Pella), list).—Fenyes, 1920:
297 (Zyras (Pella), list). — Lohse, 1974: 225 (Zyras (Pella), key). — 1989: 220 (Zyras (Pella), key).—Likovský,
1981: 167 (Zyras (Pella), morphology). — Likovský, 1993: 59 (Zyras (Pella), list).—Smetana, 2004: 461 (Zyras(Pella), list).
Myrmedonia mustela Rottenberg, 1870: 25 (original description).—Fenyes, 1920: 297 (synonym of erratica, list).—Bernhauer & Scheerpeltz, 1926: 697 (synonym of erratica, list).—Smetana, 2004: 461 (synonym of erratica, list).
Myrmedonia Ehlersi Eppelsheim, 1884b: 365 (original description).—Fenyes, 1920: 297 (synonym of erratica, list).—Bernhauer & Scheerpeltz, 1926: 697 (synonym of erratica, list).—Smetana, 2004: 461 (synonym of erratica, list).

Type material. Myrmedonia erratica. Not examined, probably missing (Frisch, pers.
comm.).
Myrmedonia mustela. Not examined. Type locality: Sicilia, Italy.
Myrmedonia Ehlersi. Holotype: Morocco: /, “Maroi/Ehlaersi mihi. Marocco Quedenfeldt/
l’antenne gauche a...[indecipherable] une pression accidentelle/c. Eppelsh. Teind. D./mustela
Rottbg/Ehlersi Epp. Berl. ent. Zeit. 1884 p. 365” (NHMW).
Type locality. Germany.
Other material. Algeria: 1?, “Milianah/Syntype/R.I.Sc.N.B. 17.819 Coll. A. Fauvel M.
maura” (see, “Type material” of Pella maura) (IRSNB); 1?, “Lalla Marghria algeria” (IRSNB);

Fig. 58.

Pella erratica (Hagens). — A, Fore body, dorsal view; B, right antenna.

90

M. Maruyama

Fig. 59. Pella erratica (Hagens). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

1?, “Pajares (Esp.)/Larerduse (Algeria)/Ehlersi type Marocco/mustela Rottb.”(IRSNB). Austria:
1?, Wien (NHMW); 1?, Wien, Fauvel (NHMW); 2??, Frauenstein, Mödling, 21 IV 1928, O.
Sheerpeltz (NHMW). Slovakia: 1?, Košice, 1925, V. Machulka (NMP). Greece: 1 ex., Pelion,
Tnessalien (NHMW); France: 1?, “Vauclusa VII Ch. Fagniez/Chasses en auto/Myrmedonia erratica? Hag.” (MNHN); 1/ “Collioure au vol 6 Carcassonne F” (IRSNB). Tunisia: 1?, “T Le
Kef, Normand” (NMP). Morocco: 1?, Morocco (NHMW).
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Belgium,
Bosnia Herzegovina, Czech Republic, France, Germany, Italy (Sicilia), Romania, Slovakia,
Spain. NORTH AFRICA: Algeria, Morocco, Tunisia.
Diagnosis. Pella erratica may be easily distinguished from the sympatric species by a combination of the body smaller and unicoloured, the apical lobe of aedeagal median lobe gently
curved ventrad and the host ant Tapinoma erraticum. In body size, it is similar to Pella barbara,

Palearctic Species of Pella

91

Fig. 60. Pella erratica (Hagens). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, D, spermathecae. Female.

but may be easily distinguished from it by the paler body and the 11th antennal segment shorter
than the 1st.
Redescription. Body slender. Brown in ground colour; antennae, mouthparts, legs and 3rd
to 5th abdominal segments reddish brown. Head (Fig. 58 A) widest just behind eyes; surface finely reticulated, moderately covered with setae; setae short, shorter than those on pronotum andelytra; length of eyes 0.34–0.37 times as long as head width. Antennae (Fig. 58 B) shorter thanhead, pronotum and elytra combined; 1st segment almost as long as 2nd and 3rd combined; 2ndsegment slightly shorter than 3rd; 3rd segment about 1/2 as long as 1st; 4th segment almost as
long as wide; 5th to 10th segments wider than long; 11th segment longer than 1st; approximate
relative lengths of segments from basal to apical: 16: 7.5 : 8 : 6 : 5 : 5.5 : 6 : 5.5 : 6 : 7 : 19. Pronotum (Fig. 58 A) 1.27–1.35 times as wide as long, widest around middle, narrowed posteriorly;
posterior angle rounded; surface finely punctured, finely reticulated among punctures, densely
covered with setae, with 10–12 macrosetae along lateral margin. Scutellum with surface finely
punctured, densely covered with setae. Elytra (Fig. 58 A) slightly widened apicad; surface finely
punctured, reticulated among punctures, densely covered with setae, with 6–8 small macrosetae
laterally. Legs moderate in length; hind tibia 0.91–0.99 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 5 : 5 : 5 : 14 in fore tarsus; 9 : 8 : 7 : 6 : 9 in midtarsus; 16 : 9 : 8 : 7.5 : 9 in hind tarsus. Abdomen subparallel-sided, slightly wider than elytra,

92

M. Maruyama

Fig. 61. Pella erratica (Hagens). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male
from Algeria.

widest around 4th and 5th segments; surface smooth; 3rd to 7th tergites sparsely and uniformly
covered with minute setae; 8th tergite (Figs. 59 A, 60 A, 61 A) with 7 (rarely 8) macrosetae; 9th
tergite with 4 macrosetae; 10th tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Figs. 59 A, 61 A) with posterior margin rounded or truncated, its apicalmargin crenate; 8th sternite (Figs. 59 B, 61 B) with posterior margin rounded, with 11 or 12
macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs. 59 C, 59 D, 61 C, 61 D)
somewhat pear-shaped in ventral view; apical lobe curved ventrad, obtuse at apex in lateral view,
pointed at apex in ventral view; basal ridge roundly weakly convex; copulatory piece of inner sac
as in Figs. 59 E, 61 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex,
about 1/3 as long as condylite.

Palearctic Species of Pella

93

Female: Eighth tergite (Fig. 60 A) with posterior margin truncate, its truncate apex minutelydentate; 8th sternite (Fig. 60 B) with 9 macrosetae; sensory setae of 8th sternite generalised, similar to other setae in shape. Spermatheca (Fig. 60 C) curved three times; basal part about 2.5 times
as long as apical part, slightly bulbous around base, acutely curved around basal 1/3; apical part
dilated apicad, much broader than basal part, its inner wall densely wrinkled from apex to apical
3/5.
Measurements. BL, 3.6–4.1; FBL, 1.7–2.1; HL, 0.50–0.58; HW, 0.56–0.71; EL, 0.210–0.240; AL, 1.32–1.63; PL, 0.57–0.73; PW, 0.76–0.97; ELL, 0.74–0.90; ELW, 1.10–0.90; HTL,
0.68–0.88.
Local variation. Pella erratica is widely distributed from Middle Europe, through Southern Europe to North Africa. Local variations are observed in the 8th abdominal tergite and aedeagus. In the population of North Africa, the crenate part of the 8th abdominal tergite is narrower inmale (Fig. 61 A), and the distal crest of the aedeagal median lobe is less prominent in lateral view(Fig. 61 C).
Bionomics. Pella erratica is known to be associated with Tapinoma erraticum and collected from inside host nests, but further biological or behavioural data have not been reported any
more. This species has been regarded as one of the rarest beetles in Europe (Franc, 1992;
Likovsky, 1981). No information was available for the life cycle of this species.
Symbiotic host. Tapinoma erraticum.
The schillhammeri GroupSpecies included. Pella schillhammeri.
Distribution. Manchurian subregion.
Diagnosis. Species of the schillhammeri group may be characterised by a combination of
the following character states: 1) eye length 0.39 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with postero-lateral corners; 4) pronotum 1.27 times aswide as long; 5) pronotum widest around middle; 6) pronotal hypomeron fully visible in lateralview; 7) elytra bicoloured, yellow with black maculations around scutellum and postero-lateralcorners; 8) posterior margin of 8th tergite crenate; 9) posterior margin of male 8th sternite rounded; 10) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior margins of the segment; apical lobe of aedeagal median lobe gently curved ventrad in lateral view;14) distal crests of aedeagus well developed, projected semicircularly in lateral view; 15) copulatory piece of aedeagal median lobe almost spindle-shaped and pointed at apex in dorsal view.
Comments. The schillhammeri group is a monotypical species-group. This species-groupis quite distinct from the other species-groups in general appearance, i.e., Pella schillhammeri is
not a typical species in comparison with the other Pella species in some character states, particularly in the colour and the facies. The general appearance is characterised by the narrower pronotum and the broader elytra, and is somewhat similar to those of the Athetini.
Symbiotic host. Unknown.
Pella schillhammeri Maruyama, n. sp.
(Figs. 62, 63)

Type material. Holotype: Russia (Far East): ?, “Seitengraben des Piwnaja-Rjetschka
Tales b. Wladiwostok/Myrmoecia spec./Farieb leg. 1918–1920/ex coll. Scheerpeltz” (NHMW).
Type locality. Vladivostok, Primorskyi Krai, Russia.

94

M. Maruyama

Fig. 62.

Pella schillhammeri Maruyama, n. sp.—A, Fore body, dorsal view; B, right antenna.

Distribution. Manchurian subregion. ASIA: Russia (Far East).
Etymology. Dedicated to Dr. Harald Schillhammer (NHMW), a curator of the type depository and a specialist of the Staphylininae, for his cooperation in the present study.
Diagnosis. Pella schillhammeri may be easily distinguished from the sympatric species by
a combination of the smaller body and the narrower pronotum.
Description. Body slender. Brown in ground colour; antennae, mouthparts, legs and posterior margins of 3rd to 5th abdominal segments reddish yellow. Head (Fig. 62 A) widest just behind eyes; surface finely reticulated, sparsely covered with setae; setae short, slightly shorter thanthose on pronotum and elytra; length of eyes 0.39 times as long as head width. Antennae (Fig.
62 B) shorter than head, pronotum and elytra combined; 1st segment shorter than 2nd and 3rd
combined; 2nd segment much shorter than 3rd; 3rd segment about 3/4 as long as 1st; 4th segment almost as long as wide; 5th to 10th segments longer than wide; 11th segment shorter than
1st; approximate relative lengths of segments from basal to apical: 20.5 : 10 : 15 : 7 : 7 : 7 : 8 : 7 : 7 :7 : 17. Pronotum (Fig. 62 A) 1.27 times as wide as long, widest around middle, narrowed posteriorly; posterior angle rounded; surface roughly punctured, finely reticulated among punctures,
densely covered with setae, with 8 or 9 macrosetae along lateral margin. Scutellum with surface
finely punctured, densely covered with setae. Elytra (Fig. 62 A) slightly widened apicad; surface
finely punctured, reticulated among punctures, densely covered with setae, with 5 or 6 small
macrosetae laterally. Legs moderate in length; hind tibia 1.06 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 5.5 : 7 : 9.5 in fore tarsus; 10 : 6 :6 : 5.5 : 10 in mid tarsus; 16 : 7 : 8 : 7 : 11 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 4th and 5th segments; surface smooth; 3rd to 7th tergites sparse-

Palearctic Species of Pella

95

Fig. 63. Pella schillhammeri Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal
sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece,
dorsal view. Male.

ly and uniformly covered with minute setae; 8th tergite (Fig. 63 A) with 6 macrosetae; 9th tergite
with 4 macrosetae; 10th tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 63 A) with posterior margin truncated, its apical margin crenate;
8th sternite (Fig. 63 B) with posterior margin rounded, with 10–12 macrosetae; 9th sternite withposterior margin truncate. Aedeagus (Figs. 63 C, 63 D) somewhat pear-shaped in ventral view;apical lobe curved ventrad, obtuse at apex in lateral and ventral views; basal ridge roundly andweakly convex; copulatory piece of inner sac as in Fig. 63 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female unknown.
Measurements. BL, 4.1; FBL, 1.9; HL, 0.62; HW, 1.81; EL, 0.239; AL, 1.65; PL, 0.66;
PW, 0.84; ELL, 0.78; ELW, 1.09; HTL, 0.83.
Bionomics and Symbiotic host. No information.

96

M. Maruyama

The kinomurai GroupSpecies included. Pella kinomurai, P. primorskyiana.
Distribution. Manchurian subregion.
Diagnosis. Species of the kinomurai group may be characterised by a combination of thefollowing character states: 1) eye length 0.37–0.40 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with posterior margin completely rounded, not forming corners postero-laterally; 4) pronotum 1.37–1.46 times as wide as long; 5) pronotum widest
anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytra unicoloured, brown; 8)
inner and posterior margins of elytra weakly margined; 9) male 8th sternite slightly longer than infemale; 10) posterior margin of 8th tergite crenate; 11) posterior margin of male 8th sternite
rounded; 12) lateral projection of apodeme of male 8th tergite evidently longer than that of female; 13) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior
margin of the segment; 14) postero-medial margin of female 8th sternite without minute pubescence; 15) apical lobe of aedeagal median lobe slightly curved ventrad in lateral view; 16) distal
crests of aedeagus not projected or only weakly projected in lateral view; 17) copulatory piece of
aedeagal median lobe slender and subparallel-sided around base in dorsal view.
Comments. The kinomurai group may possibly be allied to the funesta group in resemblance of general appearance, but the body is bicoloured and the pronotum is not narrowed posteriad as in the member of the funesta group.
Symbiotic hosts. Lasius (Dendrolasius) spp.
Key to the Species of the kinomurai Group——

Body larger: 5.3–6.5 mm; eyes smaller, 0.37–0.38 times as long as head width; pronotum
narrower, 1.37–1.44 times as wide as long; apical lobe of aedeagal median lobe shorter,
about 1/2 as long as basal capsule. Distribution: Russia Japan. ......................... P. kinomurai.
Body smaller: 5.1 mm; eyes larger, 0.40 times as long as head width; pronotum narrower,
1.47 times as wide as long; apical lobe of aedeagal median lobe longer, 0.8 times as long as
basal capsule. Distribution: Japan. (Far East). .............................................P. primorskyiana.
Pella kinomurai Maruyama, n. sp.
(Figs. 64–66)

Type material. Holotype: Japan: ?, “Isshiki, Shoukawa-mura [Shôkawa-mura] Gifu
Pref., JAPAN, 27 IX 2003, K. Kinomura leg.” (LDC) (NSMT).
Paratypes: Japan: Hokkaidô: 1 ex., Himenuma, Rishiri-tô, 4 IX 1990, T. Kishimoto (pt); 1
ex., Taihei, Maruseppu-chô, 16–17 VI 2000, Y. Kida (LDFJ); 1 ex., Kaitaku-kinenkan, Nopporoshinrin-kôen, Ebetsu-shi, 2–8 VIII 2000, S. Hori (fit); 1 ex., same data but, 8–16 VIII 2000; 5
exs., Shibumbetsu, Nopporo-shinrin-kôen, Ebetsu-shi, 18 V 2000, M. Maruyama (LDFJ); 1 ex.,Tomambetsu, Nopporo-shinrin-kôen, Ebetsu-shi, 4 V 2000, M. Maruyama (LDFJ); 1 ex., samedata bu, 18 V 2000 (LDS); 18 exs., Ôsawaguchi, Nopporo-shinrin-kôen, Ebetsu-shi, 4 V 2000,
M. Maruyama (LDFJ); 24 exs., Kannnon-zawa, Sapporo-shi, 31 V 2002, M. Maruyama (LDFL);
2 exs., same data but, 1 VI 2002. Honshû: 1 ex., Gozaishi-kôsen, Nirasaki-shi, Yamanashi-ken,
18 X 1989, K. Hosoda; 9 exs., Kentô-san, Takasu-mura, Gifu-ken, 27 IX 2001, K. Kinomura(LDFJ); 21 exs., Isshiki, Shôkawa-mura, Gifu-ken, 27 IX 2001, K. Kinomura (LDC); 1 ex., Bijokôgen, Asahi-mura, Gifu-ken, 14 IX 2003, K. Kinomura (LDFJ).

Palearctic Species of Pella

Fig. 64.

97

Pella kinomurai Maruyama, n. sp.—A, Fore body, dorsal view; B, right antenna.

Type locality. Gifu-ken, Honshû, Japan.
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô, Honshû).
Etymology. Dedicated to Mr. Kyoichi Kinomura (Gifu-ken, Japan), a collector of the type
series, and a specialist of faunistics and ecology of ants and myrmecophiles in Japan.
Diagnosis. Pella kinomurai is closely similar to P. primorskyiana in general appearance but
distinguished from it by the narrower pronotum and the shorter apical lobe of aedeagal medianlobe. This species is also similar to the species of the cognata group in general appearance but
may be easily distinguished from them by the elytra almost unicoloured, by the apical lobe of
aedeagal median lobe almost straight, not curved ventrad and in ventral view somewhat pointed
at apex, and by the distal crest of aedeagal median lobe not produced. Among the sympatricspecies, Pella kinomurai is most similar to P. japonica, but is distinguished from the latter by the
diagnostic characters above and by the 3rd to 5th abdominal segments not paler than the othersegments, the posterior emargination of male 8th tergite narrower and the copulatory piece subparallel-sided around base. Pella kinomurai is also similar to P. indiscreta in the body shape but
easily distinguished from the latter by the much larger body.
Description. Body slender. Brown in ground colour; antennae, mouthparts, legs, and around
posterior margins of 3rd to 5th abdominal segments reddish brown. Head (Fig. 64 A) widest justbehind eyes; surface finely reticulated, densely covered with setae; setae short, as long as thoseon pronotum and elytra; length of eyes 0.37–0.38 times as long as head width. Antennae (Fig.
64 B) shorter than head, pronotum and elytra combined; 1st segment almost as long as 2nd and3rd combined; 2nd segment about 2/3 as long as 3rd; 3rd segment about 3/5 as long as 1st; 4th to98

M. Maruyama

Fig. 65. Pella kinomurai Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

10th segments wider than long; 11th segment much shorter than 1st; approximate relative lengthsof segments from basal to apical: 25 : 11 : 15 : 8 : 8.5 : 8 : 7 : 8 : 9 : 9 : 14. Pronotum (Fig. 64 A)
1.37–1.44 times as wide as long, widest around anterior 1/3, narrowed posteriorly; posterior margin rounded; surface finely punctured, finely reticulated among punctures, densely covered withsetae, with 7 or 8 macrosetae along lateral margin. Scutellum with surface finely punctured,densely covered with setae. Elytra (Fig. 64 A) subparallel-sided; surface finely punctured, reticulated among punctures, densely covered with setae, with 6 or 7 small macrosetae laterally. Legs
moderate in length; hind tibia 1.04–1.06 times as long as elytra; approximate relative lengths oftarsal segments from basal to apical: 7 : 6 : 6 : 11.5 in fore tarsus; 14.5 : 8.5 : 8 : 6 : 10 in mid tarsus; 24 : 10 : 8.5 : 6 : 10.5 in hind tarsus. Abdomen subparallel-sided, slightly wider than elytra,
widest around 4th and 5th segments; surface weakly reticulated; 3rd sparsely covered with setae;

Palearctic Species of Pella

99

Fig. 66. Pella kinomurai Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, D, spermathecae. Female.

4th to 6th tergites almost glabrous except for a row of setae along each posterior margin; 7th tergite glabrous except for lateral areas sparsely covered with minute setae; 8th tergite (Figs. 65 A,
66 A) with 9 (rarely 10) macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior
margin rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 65 A) with posterior margin slightly emarginate; emarginate apex
weakly crenate; 8th sternite (Fig. 65 B) with 19–21 macrosetae; 9th sternite with posterior margin
truncate. Aedeagus (Figs. 65 C, 65 D) amygdaloid in ventral view; apical lobe slightly curved
ventrad, obtuse at apex in lateral and ventral views; basal ridge weakly convex, reaching near bottom of basal capsule; copulatory piece of inner sac as in Fig. 65 E. Paramere with apical lobe ofparamerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 66 A) with posterior margin slightly emarginate; emarginate apex
weakly crenate; 8th sternite (Fig. 66 B) with 15–17 macrosetae; sensory setae of 8th sternite spindle-shaped and flattened. Spermatheca (Figs. 66 C, 66 D) curved three times; basal part about
1.5–2.0 times as long as apical part, acutely curved at middle, gently curved and narrowed apicad; apical part acutely curved around apical 1/3, narrowed apicad, its inner wall densely wrinkledfrom apex to around apical 1/2.
Measurements. BL, 5.3–6.5; FBL, 2.4–2.8; HL, 0.71–0.83; HW, 0.85–0.96; EL, 0.319–
0.366; AL, 1.90–2.15; PL, 0.87–1.02; PW, 1.26–1.45; ELL, 1.03–1.18; ELW, 1.40–1.69; HTL,
1.09–1.23.

100

M. Maruyama

Bionomics. Pella kinomurai is relatively rarely and locally collected from nests of Lasius(Dendrolasius) fuji, L. (D.) spathepus and L. (D.) capitatus.
Symbiotic host. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) capitatus.
Pella primorskyiana Maruyama, n. sp.
(Figs. 67, 68)

Type material. Holotype: Russia (Far East): ?, “RUSSIA: Primorskyi Krai, Khasanskyi,
Pluostrov Gamov, Bukhta Vityaz, 31 V 2005, Maruyama M.” (LDFJ) (IBSS).
Type locality. Khasanskyi, Primorskyi Krai, Russia.
Distribution. Manchurian subregion. ASIA: Russia (Far East).
Etymology. Named after the type locality.
Diagnosis. Pella primorskyiana is closely similar to P. kinomurai in general appearance but
distinguished from it by the wider pronotum and the longer apical lobe of aedeagal median lobe.
This species is also similar to the species of the cognata group in general appearance but may beeasily distinguished from them by the diagnostic characters of P. kinomurai mentioned above.
Description. Body slender. Brown in ground colour; antennae, mouthparts, legs, and around
posterior margins of 3rd to 5th abdominal segments reddish brown. Head (Fig. 67 A) widest justbehind eyes; surface finely reticulated, densely covered with setae; setae short, as long as thoseon pronotum and elytra; length of eyes 0.40 times as long as head width. Antennae (Fig. 67 B)
shorter than head, pronotum and elytra combined; 1st segment almost as long as 2nd and 3rd
combined; 2nd segment about 2/3 as long as 3rd; 3rd segment about 3/5 as long as 1st; 4th to

Fig. 67.

Pella primorskyiana Maruyama, n. sp.—A, Fore body, dorsal view; B, right antenna.

Palearctic Species of Pella

101

Fig. 68. Pella primorskyiana Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal
sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece,
dorsal view. Male.

10th segments wider than long; 11th segment slightly shorter than 1st; approximate relative
lengths of segments from basal to apical: 28 : 11 : 16 : 8 : 8 : 7 : 7 : 7.5 : 7.5 : 8 : 25. Pronotum (Fig.
67 A) 1.46 times as wide as long, widest around anterior 1/3, narrowed posteriorly; posterior
margin rounded; surface finely punctured, finely reticulated among punctures, densely covered
with setae, with 8 macrosetae along lateral margin. Scutellum with surface finely punctured,densely covered with setae. Elytra (Fig. 67 A) subparallel-sided; surface finely punctured, reticulated among punctures, densely covered with setae, with 6 small macrosetae laterally. Legs moderate in length; hind tibia almost as long as elytra; approximate relative lengths of tarsal segmentsfrom basal to apical: 8.5 : 6 : 6 : 11 in fore tarsus; 14 : 9 : 9 : 6 : 11 in mid tarsus; 20 : 10.5 : 9.5 : 7.5 :11 in hind tarsus. Abdomen subparallel-sided, slightly wider than elytra, widest around 4th and5th segments; surface weakly reticulated; 3rd tergite sparsely covered with setae; 4th to 6th tergites almost glabrous except for a row of setae along each posterior margin; 7th tergite sparsely
covered with small setae; 8th tergite (Figs. 68 A) with 9 macrosetae; 9th tergite with 4 macrose-

102

M. Maruyamatae; 10th tergite with posterior margin rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 68 A) with posterior margin slightly emarginate; emarginate apex
weakly crenate; 8th sternite (Fig. 68 B) with 19–24 macrosetae; 9th sternite with posterior margin
truncate. Aedeagus (Figs. 68 C, 68 D) amygdaloid in ventral view; apical lobe slightly curved
ventrad, obtuse at apex in lateral and ventral views; basal ridge weakly convex, invisible in lateralview; copulatory piece of inner sac as in Fig. 68 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female unknown.
Measurements. BL, 5.1; FBL, 2.2; HL, 1.53; HW, 0.87; EL, 0.348; AL, 1.92; PL, 0.82;
PW, 1.20; ELL, 0.98; ELW, 1.36; HTL, 0.98.
Bionomics. Pella primorskyiana was collected from nests of Lasius (Dendrolasius) fuji.
Probably, a rare species such as Pella kinomurai.
Symbiotic host. Lasius (Dendrolasius) fuji.
The hlavaci GroupSpecies included. Pella hlavaci.
Distribution. Manchurian subregion.
Diagnosis. Species of the hlavaci group may be characterised by a combination of the following character states: 1) eye length 0.32–0.33 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with posterior margin completely rounded, not forming corners postero-laterally; 4) pronotum 1.48–1.52 times as wide as long; 5) pronotum widest
at middle; 6) pronotal hypomeron fully visible in lateral view; 7) elytra unicoloured, blackish
brown; 8) inner and posterior margins of elytra weakly margined; 9) posterior margin of 8th tergite crenate; 10) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior margins of the segment; 11) postero-medial margin of female 8th sternite without minutepubescence.
Comments. The hlavaci group is a monotypical species-group. This species-group may
possibly be allied to the funesta group in resemblance of general appearance including bodycolour, but the body is not narrowed posteriad as in the member of the funesta group. The shapeof pronotum and the shape of the spermatheca are closely similar to those of the spreta group.
Symbiotic host. Lasius (Dendrolasius) fuji.
Pella hlavaci Maruyama, n. sp.
(Figs. 69, 70)

Type material. Holotype: China (Beijing Shi): /, “Xiaolongmen (1400 m), Dongling,
Yan-shan, Beijing, CHINA, 15–16 VI 2001, Hlaváč & Cooter leg.” (LDFJ) (IZAS).
Paratypes: 3// , same data as holotype (IZAS, cMar, cHlav).
Type locality. Xiaolongmen, Beijing Shi, China.
Distribution. Manchurian subregion. ASIA: China (Beijing Shi).
Etymology. Dedicated to Mr. Peter Hlaváč (Košice, Slovakia), a collector of the type series,
and an eager specialist on the systematics of the Staphylinidae.
Diagnosis. Pella hlavaci is similar to P. jureceki in general appearance and colour, but iseasily distinguished from the latter by the almost elliptical pronotum, and the apical part of spermatheca apically narrowed.
Description. Body robust. Blackish brown in ground colour; antennae, mouthparts, legs,Palearctic Species of Pella

Fig. 69.

103

Pella hlavaci Maruyama, n. sp. —A, Fore body, dorsal view; B, right antenna.

and around posterior margins of 3rd to 5th abdominal segments reddish brown. Head (Fig. 69 A)
widest just behind eyes; surface finely reticulated, densely covered with setae; setae short, as longas those on pronotum and elytra; length of eyes 0.32–0.33 times as long as head width. Antennae(Fig. 69 B) shorter than head, pronotum and elytra combined; 1st segment much longer than 2nd
and 3rd combined; 2nd segment about 1/3 as long as 1st; 3rd segment about 1/2 as long as 1st;
4th to 10th segment slightly wider than long; 11th segment much shorter than 1st; approximate
relative lengths of segments from basal to apical: 30 : 9.5 : 15 : 7 : 7 : 6 : 6.5 : 7 : 7.5 : 8 : 20. Pronotum (Fig. 69 A) 1.48–1.52 times as wide as long, widest at middle; posterior margin rounded;surface finely punctured, finely reticulated among punctures, densely covered with setae, with 7
or 8 small macrosetae along lateral margin. Scutellum with surface finely punctured, densely covered with setae. Elytra (Fig. 69 A) subparallel-sided; surface finely punctured, reticulated among
punctures, densely covered with setae, with 6 or 7 small macrosetae laterally. Legs long, almost
as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 9 : 6.5 :5.5 : 11 in fore tarsus; 13.5 : 9.5 : 8 : 6.5 : 10 in mid tarsus; 14 : 9 : 8.5 : 7.5 : 11 in hind tarsus. Abdomen expanded, 1.03 times wider than elytra, widest around 6th and 7th segments; surface
smooth; 3rd to 6th tergites glabrous except for a row of setae along each posterior margin; 7th
tergite glabrous except for lateral areas sparsely covered with minute setae; 8th tergite (Fig. 70 A)
with 9 macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior margin rounded,with 4 macrosetae.
Male unknown.
Female: Eighth tergite (Fig. 70 A) with posterior margin slightly emarginate, weakly crenate
around emarginate apex; 8th sternite (Fig. 70 B) with 11 or 12 macrosetae; sensory setae of 8th

104

M. Maruyama

Fig. 70. Pella hlavaci Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

sternite slightly flattened. Spermatheca (Fig. 70 C) curved three times, M-shaped; basal partabout 1.5 times as long as apical part, gently curved just before middle and near apex, narrowed
apicad; apical part gently curved, narrowed apicad, its inner wall densely wrinkled from apex toaround apical 2/3.
Measurements. BL, 5.5–5.7; FBL, 2.4–2.5; HL, 0.76–0.78; HW, 0.84–0.85; EL, 0.269–
0.281; AL, 1.75–1.83; PL, 0.81–0.85; PW, 1.23–1.26; ELL, 0.64–0.98; ELW, 1.38–1.42; HTL,
0.94–1.01.
Bionomics. The type series of P. hlavaci were collected from a colony of Lasius (Dendrolasius) fuji together with many other staphylinids (see, Bionomics of P. cooterorum).
Symbiotic host. Lasius (Dendrolasius) fuji.
The cognata GroupSpecies included. Pella cognata, P. iberica, P. japonica, P. kishimotoi.
Distribution. European subregion, Mediterranean subregion, Manchurian subregion.
Diagnosis. Species of the cognata group may be characterised by a combination of the following character states: 1) eye length 0.34–0.37 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with posterior margin completely rounded, not forming corners postero-laterally; 4) pronotum 1.25–1.40 times as wide as long; 5) pronotum widest
anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytron with yellowish brownmaculation around antero-lateral corner to mesal area; 8) inner and posterior margins of elytra
weakly margined; 9) male 8th sternite longer than in female; 10) posterior margin of 8th tergite

Palearctic Species of Pella

105

crenate; 11) posterior margin of male 8th sternite rounded or slightly truncate; 12) lateral projection of apodeme of male 8th tergite evidently longer than that of female; 13) macrosetae of 8th
abdominal segment generalised in length, exceeding the posterior margin of the segment; 14)postero-medial margin of female 8th sternite without minute pubescence; 15) apical lobe of
aedeagal median lobe curved ventrad, thick and parallel-sided dorsoventrally in lateral view; 16)
distal crests of aedeagus well developed, projected, rounded or truncate on ventral margin in lateral view; 17) copulatory piece of aedeagal median lobe more or less pointed at apex in dorsal
view.
Comments. The cognata group can be regarded as a monophyletic group by an autapomorphy; the apical lobe of aedeagal median lobe is thick and parallel-sided dorsoventrally. This stateis unique within the genus and not observed in the other species of the Lomechusini. Thus, thisstate is considered to be autapomorphy of the species-group. This species-group may possibly beallied to the similis group in resemblance of the shape of aedeagus, which is completely oval inventral view. This species-group also similar to the limbata group in the shape of male 8th tergite, which is widely emarginate posteriorly just as in the similis group.
Symbiotic hosts. Lasius (Dendrolasius) spp.
Key to the Species of the cognata Group
1. Occiput with a pair of projections medially; apical lobe of aedeagal median lobe longer
than basal capsule. Distribution: China (Hunan Sheng). ....................................P. kishimotoi.
— Occiput without projection medially, only somewhat convex dorsally; apical lobe of aedeagal median lobe as long as basal capsule. ........................................................................... 2.
2. Eye length 0.32–0.33 times as long as head width; copulatory piece almost circular. Distribution: Japan, Korea, Russia (Far East). ..................................................................P. japonica.
— Eye length 0.34–0.37 times as long as head width; copulatory piece subparallel-sided frombase to middle. Distribution: Europe. ................................................................................. 3.
3. Abdominal macrosetae short, those on postero-medial area of 8th tergite exceed the tergal
margin about 1/2 in their lengths. Distribution: Central Europe, Balkan. ..............P. cognata.
— Abdominal macrosetae long, those on postero-medial area of 8th tergite exceed the tergal
margin about 2/3 in their lengths. Distribution: Iberian Peninsula. .........................P. iberica.
Pella cognata (Märkel, 1842), n. comb.
(Figs. 71–73)
Aleochara cognata Märkel, 1842: 142 (original description).—Mulsant & Rey, 1873a: 69 (Myrmedonia, description).—des Gozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 122 (Myrmedonia (Pella), key, description).—Reitter, 1909:
43 (Myrmedonia (Pella), key). — Fenyes, 1920: 297 (Zyras (Pella), list).—Bernhauer & Scheerpeltz, 1926: 705(Zyras (Pella), list). — Scheerpeltz, 1934: 1651 (Zyras (Pella), list). — Lohse, 1974: 225 (Zyras (Pella), key).—
Likovský, 1993: 59 (Zyras (Pella), list). — Smetana, 2004: 461 (Zyras (Pella), list).
“Aleochara funesta”: Heer, 1839: 308 (misidentification, description).—Ganglbauer, 1895: 122 (synonym of cognata,list).—Fenyes, 1920: 297 (synonym of cognata, list).

Type material. Syntypes: Germany: 1 ex. “5250/cognata Märk, Saxon. mont. Märk[“Saxon. mont. Märk.” (mountainous area of Saxonia, Germany, F. Märkel leg.)]” (head, prothorax and 3rd to 10th abdominal segments missing, pinned); 1 ex. [ditto] (3rd to 10th abdominal
segments missing, pinned).
These are deposited in ZMHB.

106

M. Maruyama

Fig. 71.

Pella cognata (Märkel). — A, Fore body, dorsal view; B, right antenna.

Type locality. Saxonia, Germany.
Other material. Austria: 2 exs., Mark, Chorin, 22 IV 1916; 4 exs., Lainzer, Tiergarten, 30
VII 1949, O. Scheerpeltz (LDFL); 2 exs., Trübenbruch, Schönhauser Damm, Sachsen-Anhalt, V
1998, Sprick (pt); 1 ex., Koschute Aue 1300m, 13 VI 1965, A. Smetana. Bulgaria: 1 ex.,
Senokos, 20km E Tolbuchin, 4 V–19 VI 1987, Penev. Czech Republic: 1 ex., Hradec, Bohemia,
30 III 1947, A. Smetana; 1?, 1/, 23 exs., same data but, 3 IV 1947, A. Smetana; 2 exs., Ríŏany,
Bohemia, 27 III 1948, P. Ríha; 2 exs., Chlum u Třeb., Bohemia, VI 1910, J. Niedl. Greece: 1 ex.,
Kastanea, Kalambaka, Pindos, 28 III 1997, A. Riedal. Italy: 1 ex., Montecchia, Verona, 22 II
1974, A. Zanetti (LDFL) (MNV). Slovakia: 7 exs., Ruská Poruba, 4 VIII 1956, A. Smetana; 5exs., same data but, 12 IX 1956; 5 exs., same data but, 17 IX 1956.
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Belgium,
Bosnia Herzegovina, Belarus, Croatia, Czech Republic, Denmark, Finland, France, Great Britain,
Germany, Georgia, Hungary, Italy, Latvia, Lithuania, Netherlands, Norway, Poland, Romania,
Russia (North European territory), Slovakia, Sweden, Switzerland, Ukraine.
Diagnosis. Pella cognata is similar to the other species of the cognata group in general appearance, but can be distinguished from them by the copulatory piece parallel-sided around base.
The species of the cognata group are locally isolated. Among the sympatric species P. cognata issimilar to P. limbata and P. similis in general appearance, but is distinguished from P. limbata by
the postero-lateral corner of male 8th tergite not projected, the apical lobe of aedeagal median
lobe gradually narrowed apicad in ventral view and the apical part of spermatheca with outerprojection, and from P. similis by the spermathecal structure above, the apical lobe of aedeagal

Palearctic Species of Pella

107

Fig. 72. Pella cognata (Märkel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

median lobe longer and the basal part of spermatheca much longer.
Redescription. Body slender. Reddish brown in ground colour; antennae, mouthparts,
around antero-lateral corner to mesal area of each elytron, legs and 3rd to 5th abdominal segments paler. Head (Fig. 71 A) widest at middle of eyes; surface finely reticulated, densely covered
with setae; setae short, as long as those on pronotum and elytra; length of eyes 0.36–0.37 times
as long as head width. Antennae (Fig. 71 B) shorter than head, pronotum and elytra combined;
1st segment evidently shorter than 2nd and 3rd combined; 2nd segment about 1/2 as long as 1st;
3rd segment about 2/3 as long as 1st; 4th and 5th segments almost as long as wide; 6th to 10th
segments wider than long; 11th segment much shorter than 1st; approximate relative lengths ofsegments from basal to apical: 30.5 : 15 : 19 : 8.5 : 8 : 8 : 8.5 : 7 : 8 : 9 : 22.5. Pronotum (Fig. 71 A)
1.35–1.40 times as wide as long, widest around anterior 1/5, narrowed posteriorly; posterior margin rounded; surface finely punctured, finely reticulated among punctures, densely covered with

108

M. Maruyama

Fig. 73. Pella cognata (Märkel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

setae, with 10–12 macrosetae along lateral margin. Scutellum with surface finely punctured,densely covered with setae. Elytra (Fig. 71 A) subparallel-sided; surface finely punctured, reticulated among punctures, densely covered with setae, with 6–8 small macrosetae laterally. Legs
moderate in length; hind tibia 1.06–1.08 times as long as elytra; approximate relative lengths oftarsal segments from basal to apical: 9.5 : 6 : 6 : 13 in fore tarsus; 12.5 : 10 : 8.5 : 7 : 12.5 in midtarsus; 20 : 11 : 9.5 : 8.5 : 14.5 in hind tarsus. Abdomen subparallel-sided, slightly narrower thanelytra, widest around 4th and 5th segments; surface weakly reticulated; 3rd to 5th tergites
glabrous except for a row of setae along each posterior margin; 6th and 7th tergites sparsely anduniformly covered with minute setae; 8th tergite (Figs. 72 A, 73 A) with 6 macrosetae; 9th tergite
with 3 setae; 10th tergite with posterior margin slightly emarginate, with 3 setae.
Male: Eighth tergite (Fig. 72 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 72 B) with posterior margin truncate, with 18–20 macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs. 72 C, 72 D) oval in ventral view;apical lobe curved ventrad, obtuse at apex in lateral view, rounded at apex in ventral view; basal
ridge roundly convex; copulatory piece of inner sac as in Fig. 72 E. Paramere with apical lobe ofparamerite dilated apicad, rounded at apex, about 1/5 as long as condylite.
Female: Eighth tergite (Fig. 73 A) with posterior margin truncate; truncate apex weakly crenate; 8th sternite (Fig. 73 B) with 12–13 macrosetae; sensory setae of 8th sternite generalised, almost the same as the other setae in shape. Spermatheca (Fig. 73 C) curved three times; basal partalmost as long as apical part, acutely curved around middle, slightly dilated apicad; apical part
acutely curved around apical 2/5, narrowed apicad, its inner wall densely wrinkled from apex toaround apical 2/3.

Palearctic Species of Pella

109

Measurements. BL, 5.2–5.8; FBL, 2.6–2.8; HL, 0.75–0.78; HW, 0.90–0.95; EL, 0.329–
0.341; AL, 2.06–2.08; PL, 0.89–0.96; PW, 1.25–1.30; ELL, 1.08–1.10; ELW, 1.43–1.52; HTL,
1.11–1.13.
Bionomics. Pella cognata is collected from nests of Lasius (Dendrolasius) fuliginosus inEurope relatively rarely and locally (France, 1992).
Symbiotic host. Lasius (Dendrolasius) fuliginosus.
Pella iberica Maruyama, n. sp.
(Figs. 74–76)

Type material. Holotype: Spain: ?, “E. Andalusien (MA), Sierra de Palmitera, SO Ronda,
900 m, 24 III 1994 Assing 13” (cAss).
Paratypes: Spain: 2// , “Espagne: Castille Puerto de Bejar 800–1400 m V–VI 1957 G.
Fagel/G. Fagel det.. cognata Märk.” (IRSNB).
Type locality. Sierra de Palmitera, Andalusia, Spain.
Distribution. Mediterranean subregion. EUROPE: Spain. Probably confined to the Iberian
Peninsula.
Etymology. Named after the Iberian Peninsula, in which lies the type locality.
Diagnosis. Pella iberica is similar to the other species of the cognata group in general appearance but can be distinguished from them by the macrosetae and setae of the body evidently
longer, particularly on the pronotum, elytra, and 8th tergite and sternite. The species of the cognata group are locally isolated. This species can be distinguished from the sympatric species by
the diagnostic character states of the cognata group above, especially the aedeagal shape.

Fig. 74.

Pella iberica Maruyama, n. sp. —A, Fore body, dorsal view; B, right antenna.

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M. Maruyama

Fig. 75. Pella iberica Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

Description. Body slender. Brown in ground colour; antennae, mouthparts, around anterolateral corner to mesal area of each elytron, legs, and around posterior margins of 3rd to 5th abdominal segments reddish brown. Head (Fig. 74 A) widest just behind eyes; surface finely reticulated, densely covered with setae; setae long, as long as those on pronotum and elytra; length ofeyes 0.34–0.35 times as long as head width. Antennae (Fig. 74 B) shorter than head, pronotumand elytra combined; 1st segment slightly shorter than 2nd and 3rd combined; 2nd segment about
2/3 as long as 3rd; 3rd segment about 3/5 as long as 1st; 4th to 6th segments almost as long as
wide; 7th to 10th segments wider than long; 11th segment as long as 1st; approximate relative
lengths of segments from basal to apical: 29.5 : 12.5 : 18 : 7 : 7 : 6 : 7.5 : 6.5 : 6.5 : 8.5 : 29.5. Pronotum (Fig. 74 A) 1.26–1.36 times as wide as long, widest around anterior 1/5, narrowed posterior-

Palearctic Species of Pella

111

Fig. 76. Pella iberica Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

ly; posterior margin rounded; surface finely punctured, finely reticulated among punctures, densely covered with long setae, with 8 or 9 macrosetae along lateral margin. Scutellum with surface
finely punctured, densely covered with setae. Elytra (Fig. 74 A) slightly widened apicad; surface
finely punctured, reticulated among punctures, densely covered with long setae, with 6–8 small
macrosetae laterally. Legs moderate in length; hind tibia 0.98–0.99 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 5.5 : 5 : 12 in fore tarsus; 10: 7 : 6 : 6 : 12.5 in mid tarsus; 15 : 9.5 : 9.5 : 7.5 : 12 in hind tarsus. Abdomen subparallel-sided,slightly narrower than elytra, widest around 4th and 5th segments; surface weakly reticulated; 3rd
to 7th tergites glabrous except for a row of setae along each posterior margin; 8th tergite (Figs.
75 A, 76 A) with 7 (rarely 8) macrosetae; 9th tergite with 3 macrosetae; 10th tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 75 A) with posterior margin slightly emarginate, its emarginate
apex crenate; 8th sternite (Fig. 75 B) with posterior margin rounded, with 18 macrosetae; 9thsternite with posterior margin slightly emarginate. Aedeagus (Figs. 75 C, 75 D) oval in ventral
view; apical lobe curved ventrad, obtuse at apex in lateral view, rounded at apex in ventral view;
basal ridge roundly convex; copulatory piece of inner sac as in Fig. 75 E. Paramere with apicallobe of paramerite dilated apicad, rounded at apex, 1/4 as long as condylite.
Female: Eighth tergite (Fig. 76 A) with posterior margin truncate; truncate apex weakly crenate; 8th sternite (Fig. 76 B) with 11–13 macrosetae; sensory setae of 8th sternite generalised, almost the same as the other setae in shape. Spermatheca (Fig. 76 C) curved three times; basal partalmost as long as apical part, acutely curved around middle, slightly dilated apicad; apical part112

M. Maruyama

acutely curved around apical 2/5, narrowed apicad, its inner wall densely wrinkled from apex toaround apical 2/3.
Measurements. BL, 5.1–5.5; FBL, 2.3–2.5; HL, 0.75–0.76; HW, 0.90–0.91; EL, 0.315–
0.319; AL, 1.95–2.02; PL, 0.93–0.94; PW, 1.17–1.28; ELL, 1.01–1.10; ELW, 1.46–1.53; HTL,
0.99–1.09.
Bionomics and symbiotic host. No information was available. Since all the other speciesof the cognata group are associated with Lasius (Dendrolasius) spp., and since L. (D.) fuliginosus is distributed around the type locality, P. iberica is probably associated with this ant
species.
Pella japonica (Sharp, 1888)(Figs. 77–79)
Myrmedonia cognata var.?: Sharp, 1874: 10 (record from Japan, comments).
Myrmedonia cognata var. japonica Sharp, 1888, 290 (original description).—Bernhauer & Scheerpeltz, 1926: 696 (variety of Zyras (Pella) cognatus, list). — Kistner, 1972: 153 (Pella, distinct species, “Pella japonicus [sic!]”, description).—Smetana, 2004: 461 (Zyras (Pella), subspecies of cognata, list).

Type material. Syntypes: Japan: 1/, “Japan. G. Lewis. 1910-320./cognatus var?? probably n. sp./japonicus Sharp/SYN-TYPE (blue round curator label)”; 1/, “Bukenji (written underside of paper card specimen glued on)/Bukenji/Japan. G. Lewis./Sharp Coll. 1905-313./Myrmedonia cognata var. japonica Japan. Lewis/SYN-, TYPE (blue round curator label)”; 1?, “Bukenji(written underside of paper card specimen glued on)/Bukenji/Japan. G. Lewis. 1910-320./SYNTYPE (blue round curator label)”; 1?, “Bukenji/Japan. G. Lewis. 1910-320./SYN-TYPE (blueround curator label)”; 3// (two are mounted on same paper card), “Bukenji (written undersideof paper card specimen glued on)/Bukenji/Japan. G. Lewis./Sharp Coll. 1905-313./SYN-TYPE(blue round curator label)”; 1/, “Bukenji 5/4/80 (written underside of paper card specimen gluedon)/Yokohama. 20. III.–14. IV. 80./Japan. G. Lewis. 1910-320./Myrmedonia, congata var.”; 1/,“Myrmedonia congata var. Sapporo. Japan. Lewis. (written on paper card specimen gluedon)/Japan. G. Lewis./Sharp Coll. 1905-313.”; 1?, “16. 6. 8 (probably, 16. 6. 1880) (written underside of paper card specimen glued on)/Kashiwagi. 15. VI–24. VI 81./Japan. G. Lewis. 1910320.”; 1?, “2/6/81 (written underside of paper card specimen glued on)/Nagasaki., 22. V–3. VI
81/Japan. G. Lewis. 1910-320.”; 1?, “14. 4. 80 (written underside of paper card specimen gluedon)/Nagasaki. 13. II.–21. IV. 81/Japan. G. Lewis. 1910-320.”.
These are deposited in BMNH.
Type locality. Sapporo (Hokkaidô), Yokohama (Honshû), or Nagasaki (Kyûshû), Japan.
Other material. Japan: Hokkaidô: 1 ex., Himenuma, Rishiri-tô, 4 IX 1990, T. Kishimoto(pt); 2 exs., Sarobetsu-gen’ya, 9 VII 1991, N. Yasuda (pt); 9 exs., Hebi-numa, Teshio-gawa,
Teshio-chô, 9 VII 1992, S. Hori (pt); 63 exs., Makoi, Shari-chô, 27–29 V 2001, Y. Kida (LDFJ);
1 ex., Nijibetsu-rindô, Yoroushi, Nakashibetsu-chô, 7 VIII 1992, K. Miyashita; 1ex., Onnenairindô, Tsurui-mura, Akan-gun, 29 VII 1994, Y. Hirama; 1 ex., Shibetsu-gawa, Shibetsu-chô,25–28 VI 2001, H. Yoshitomi; 1 ex., Taishô, Kitami-shi, 11 V 1992, T. Katô; 23 exs., Esaushisan, Kamikawa-chô, 8 VIII 1985, N. Yasuda (pt); 1 ex., Ishikari-gawa, Kamikawa-chô, 2 VI
1994, N. Yasuda (pt); 2 exs., Taihei, Maruseppu-chô, 11–12 VIII 2000, Y. Kida (LDFJ); 1 ex.,same data but, 19–21 VIII 2000; 6 exs., Makoi, Shari-chô, 28 V 2000, Y. Kida (LDFJ); 1 ex.,
Apoi-dake, Samani-chô, 13 VII 1995, N. Yasuda (pt); 1 ex., Kimontô, Taiki-chô, 7 VII 1993, S.
Hori; 3 exs., Kamiotoineppu, Otoineppu-mura, 15–23 VII 1997, S. Hori (pt); 3 exs., Kaitaku-ki-

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113

nenkan, Nopporo-shinrin-kôen, Ebetsu-shi, 20 IV 2003, S. Hori (LDFJ); 9 exs., Tomambetsu,
Nopporo-shinrin-kôen, Ebetsu-shi, 4 V 2000, M. Maruyama (LDS); 5 exs., same data but, 11 V
2000; 4 exs., same data but, 11 V 2000 (LDFJ); 8 exs., same data but, 18 V 2000 (LDS); 14 exs.,Ôsawaguchi, Nopporo-shinrin-kôen, Ebetsu-shi, 31 V 1999, M. Maruyama (LDFJ); 2 exs., samedata but, 4 V 2000; 2 exs., same data but, 9 V 2000, H. Sugaya (LDFJ); 7 exs., Shibumbetsu,
Nopporo-shinrin-kôen, Ebetsu-shi, 11 V 2000 (LDFJ); 2 exs., Hitsujigaoka, Sapporo-shi, 21 VI
1998, M. Maruyama (LDS ); 1 ex., Kannon-zawa, Sapporo-shi, 1 VI 2001, H. Sugaya (LLJ); 4
exs., same data but, 1 VI 2002, M. Maruyama (LDFJ); 3 exs., same data but, 31 V 2002; 2 exs.,
Moiwa-yama (520 m alt.), Sapporo-shi, 27 VII 1991, A. Smetana; 3 exs., Asari, Otaru-shi, 1 IX
1998, K. Sugisima (LDFJ); 4 exs., Akagawa, Hakodate-shi, 15 VIII 1992, M. Nishikawa(LDFJ). Honshû: 1 ex., Matamatsu, Yomogita-mura, Aomori-ken, 9 VI 1999, H. Uyama (pt); 1
ex., Haguro-yama, Yomogita-mura, Aomori-ken, 11 VI 1999, H. Uyama (pt); 1 ex., Gosho-ko,
Morioka-shi, Iwate-ken, 13–14 VI 1999, M. Sano (LDFJ); 1 ex., Kuzakai, Kawai-mura, Iwateken, 9 VII 1980, K. Ogata; 1 ex., Tamugi, Tokamachi-shi, Niigata-ken, 18 IV 1998, T. Kishimoto; 5 exs., Ochinomizu, Tôkamachi-shi, Niigata-ken, 18 IV 1998, T. Kishimoto (LDFJ); 1 ex.,
Sasagamine, Myôkô-san, Niigata-ken, 14–15 VI 1980, S. Naomi; 72 exs., Tobiyama-jôshi, Utsunomiya-shi, Tochigi-ken, 17 VI 1998, M. Maruyama (LDFJ); 118 exs., Shimokomoriya,
Môka-shi, Tochigi-ken, 6 VII 1999, M. Maruyama (LDFJ); 34 exs., Mizuhono, Môka-shi, Tochigi-ken, 17–18 VI 1998, M. Maruyama (LDS); 16 exs., Sayado, Môka-shi, Tochigi-ken, 15 VI
2000, T. Kobayashi & H. Obata (LDFJ); 1 ex., 4 km NW Onioshidashi, Gunma-ken, 18–21 VII
1980, A. & Z. Smetana; 3 exs., Nakano, Shôwa-machi, Kitakatsushika-gun, Saitama-ken, 17 VI
2001, H. Kamezawa (LDFJ); 4 exs., Chichibu-kôgen-bokujô, Higashichichibu-mura, Saitamaken, 3 IV 1999, K. Toyoda (LDFJ); 4 exs., Sugaya, Ranzan-machi, Saitama-ken, 13 VII 1998, K.
Toyoda (LDS); 5 exs., Shioyama, Kamagata, Ranzan-machi, Saitama-ken, 17 VI 2000, K. Toyoda(LDFJ); 1 ex., Jômine-san, Chichibu-shi, Saitama-ken, 22 IV 1999, K. Toyoda (LDS); 1 ex.,
Hashidate-gawa (alt. 500 m), Chichibu-shi, 17 VII 1999, K. Toyoda (LDS); 1 ex., Shinjuku-ku,
Tokyo-to, 14 VII 1984, S. Kubota; 2 exs., Takao-san (alt. 450 m), Hachiôji-shi, Tokyo-to, 4 VII
1998, M. Maruyama (LDS ); 39 exs., Shiroyama-kôen, Inagi-shi, Tokyo-to, 12 VI 2004, A. Yoshida (LDFJ); 1 ex., same data but, 13 VI 2004; 1 ex., same data but, 26 VI 2004; 1 ex., same databut, 27 VI 2004; 2 exs., same data but, 4 VII 2004; 15 exs., Aoba-chô, Chiba-shi, Chiba-ken, 24
VI 1998, M. Maruyama (LDS ); 63 exs., Dan’ya, Chiba-shi, Chiba-ken, 27–31 V 1997, M.
Maruyama (LDS); 11 exs., Azeta, Sakura-shi, Chiba-ken, 20 VI 1998, M. Maruyama (LDS); 4
exs., same data but, 23–24 VI 1998; 4 exs., Shigena, Tateyama-shi, Chiba-ken, 3 IV 1998, S.
Naomi (sifting leaf litter); 2 exs., Masukata-yama, Kawasaki-shi, Kanagawa-ken, 23 VI 1995, K.
Kawada (LDS ); 1 ex., Mikage, Tsukui-machi, Kanagawa-ken, 14 IV 1979, R. Kiryu (SCM); 10
exs., Jimmuji, Zushi-shi, Kanagawa-ken, 20 VI 2003, M. Maruyama (LDS); 1 ex., Hirata-rindô,Isehara-shi, Kanagawa-ken, 8 IV 2001, S. Arai (LCH); 1 ex., Shiroishi-sawa, Yamakita-chô,Kanagawa-ken, 21 V 2000, S. Nagashima; 1 ex., Oppama, Yokosuka-shi, Kanagawa-ken, 8–9 IV
2000, T. Shimada; 1 ex., Sengokubara, Hakone-machi, Kanagawa-ken, 24 IV 1997, M. Maruyama (LCAF); 3 exs., Kamiimai, Hosaka-chô, Nirasaki-shi, Yamanashi-ken, 6 VI 2004, M.
Maruyama (LDFJ); 1 ex., Ogasawara, Akeno-mura, Kitakoma-gun, Yamanashi-ken, 6 VI 2004,
M. Maruyama (LDS); 8 exs., Fukazawa-jûnimagari, Nakamaru, Nagasaka-chô, Kitakoma-gun,
Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 1 ex., Nagasakakamijô, Nagasaka-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDS); 1 exs., Hino, Nagasaka-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDC); 2 exs., Shobuzawa, Futaba-chô,Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 1 exs., Shôyakuji, Akeno-

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M. Maruyama

Fig. 77.

Pella japonica (Sharp). — A, Fore body, dorsal view; B, right antenna.

mura, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDN); 3 exs., Kamikandori,
Akeno-mura, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 24 exs., Shakashindô, Haku-san, Ishikawa-ken, 30 VII–2 VIII 2002, H. Hoshina (pt); 1 ex., Ichinose-tôge, Shiramine-mura, Ishikawa-ken, 7–22 VI 2003, K. Nakata (fit); 96 exs., same data but, 2–22 VIII
2002; 1 ex., Ôshiba-yama, Fukui-shi, Fukui-ken, 3 V 1978, H. Sasaji; 1 ex., Hakusan-superrindô, Shirakawa-mura, Gifu-ken, 5 VII 1997, Y. Takai; 26 exs., Ogamigô (alt. 800 m), Shôkawamura, Gifu-ken, 5–6 VIII 1998, M. Maruyama (LDS); 2 exs., Hirakura, Misugi-mura, Ichishigun, Mie-ken, 25 V 1964, Y. Miyake; 1 ex., Mizuho-chô, Kyôto-fu, 11 V 1978, Y. Hayashi (LL);
1 ex., Daigo-yama, Kyôto-shi, Kyôto-fu, 17 IV 1994, T. Ito; 1 ex., Minoo-shi, Ôsaka-fu, 7 IV
1958, Y. Hayashi; 1 ex., Kôya-san, Kôya-chô, Wakayama-ken, 1 V 1985, T. Ito; 2 exs., Urahikimi, Shimane-ken, 6 VI 1988, S. Nomura (LDS ); 2 exs., Daisen, Tottori-ken, 22 V 1985, S. Nomura; 1 ex., Tokusa, Atô-chô, Yamaguchi-ken, 24 IV 1988, S. Nomura. Shikoku: 1 ex., Kotsuzan, Tokushima-ken, 29 IV 1979, M. Yoshida; 2 exs., Minokoshi, Tsurugi-san, Tokushima-ken,
3 V 1969, M. Yoshida; 1 ex., Tsurugi-san, Tokushima-ken, 5 IV 1968, M. Sakai (EEEU); 1 ex.,
Sugitate, Matsuyama-shi, Ehime-ken, 27 VI 1970, M. Tomokuni (EEEU); 1 ex., Saragamine,
Ehime-ken, 4 VII 1959, M. Miyatake (EEEU). Kyûshû: 1/, “Japan, Gokayama, Chikuzen[Gokayama, Nakagawa-chô, Fukuoka-ken]”, 14 VII 1936, K. Yasumatsu (BMNH); 20 exs.,
Hikosan, Fukuoka-ken, 7 VII 1983, Y. Sawada. Korea: 47 exs., Oso-san, Sangdam-ri, Kwangchon-up, Hongsung-gun, Chungnam, 19–21 VI 1999, U.-S. Hwang & H.-J. Kim (pt) (cMar,
CNU); 1 ex., Surak, Daedunsan, Beolgok-myeon, Nansan, Chungnam, 2–10 V 2000, U.-S.
Hwang, S.-J. Park & H.-J. Kim (pt) (cMar, CNU); 1 ex., Chôndong, Sobaek-san, Tangyang-gun,
Kyôngbuk, 8–9 V 1999, U.-S. Hwang & H.-J. Kim (CNU); 4 exs., Janggoksa, Chilgab-san,Palearctic Species of Pella

115

Fig. 78. Pella japonica (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

Cheongyang-gun, Chungnam, 23 VI 2000, M. Maruyama (LDFJ) (cMar, CNU). Russia (Far
East): Kaimanovka, Ussuryisk, Primorskyi Krai, 27 V 2005, M. Maruyama (LDFJ); 2 exs.,
Bukhta Vityaz, Poluostrov Gamov, Khasanskyi, Primorskyi Krai, 30 V 2005, M. Maruyama(LDFJ); 1 ex., same data but, 31 V 2005; 21 exs., Azajsanovka, Kraskino, Primorskyi Krai,
13–16 VII 1992, Snizek; 2 exs., “Seitengraben des Prwnaja-Rjetschka Tales b. Wladiwostok/H.
Farieb leg. 1918–1920” (NHMW).
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô, Honshû, Shikoku, Kyûshû),Korea, Russia (Far East). New to Korea and Russia.
Diagnosis. Pella japonica is closely similar to the other species of the cognata group in
general appearance but can be distinguished from them by the larger body, shorter antennae, andthe shape of the pronotum. The species of the cognata group are locally isolated. Among the
sympatric species, this species can easily be distinguished from the others species by the 3rd to
5th abdominal segments evidently paler than the other segments (reddish brown).

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M. Maruyama

Fig. 79. Pella japonica (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, D, spermathecae. Female.

Redescription. Body slender. Blackish brown in ground colour; antennae, mouthparts,
around antero-lateral corner to mesal area of each elytron, legs and 3rd to 5th abdominal segments reddish brown. Head (Fig. 77 A) widest just behind eyes; surface finely reticulated, densely
covered with setae; setae short, as long as those on pronotum and elytra; length of eyes 0.32–0.33
times as long as head width. Antennae (Fig. 77 B) shorter than head, pronotum and elytra combined; 1st segment longer than 2nd and 3rd combined; 2nd segment about 2/3 as long as 3rd; 3rdsegment about 3/5 as long as 1st; 4th segment slightly longer than wide; 5th segment almost as
long as wide; 6th to 10th segments wider than long; 11th segment much shorter than 1st; approximate relative lengths of segments from basal to apical: 33.5 : 12 : 19 : 9 : 7.5 : 7 : 8 : 8 : 9 : 10 : 27.5.
Pronotum (Fig. 77 A) 1.28–1.34 times as wide as long, widest around anterior 1/3, narrowed posteriorly; posterior margin rounded; surface finely punctured, finely reticulated among punctures,
densely covered with setae, with 10–12 macrosetae along lateral margin. Scutellum with surface

Palearctic Species of Pella

117

finely punctured, densely covered with setae. Elytra (Fig. 77 A) slightly widened apicad; surface
finely punctured, reticulated among punctures, densely covered with setae, with 6–8 small
macrosetae laterally. Legs moderate in length; hind tibia 0.93–0.99 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 9 : 7 : 7.5 : 13.5 in fore tarsus;
14.5 : 10 : 10 : 8.5 : 13 in mid tarsus; 18 : 12 : 11.5 : 9.5 : 14 in hind tarsus. Abdomen subparallelsided, slightly narrower than elytra, widest around 4th and 5th segments; surface weakly reticulated; 3rd to 5th tergites glabrous except for a row of setae along each posterior margin; 6th and7th tergites sparsely and uniformly covered with minute setae; 8th tergite (Figs. 78 A, 79 A) with
7 (rarely 8) macrosetae; 9th tergite with 3 macrosetae; 10th tergite with posterior margin slightly
emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 78 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 78 B) with posterior margin truncate, with 18–20 macrosetae; 9th sternite with posterior margin slightly emarginate. Aedeagus (Figs. 78 C, 78 D) oval inventral view; apical lobe curved ventrad, obtuse at apex in lateral view, rounded at apex in ventral
view; basal ridge roundly convex; copulatory piece of inner sac as in Fig. 78 E. Paramere withapical lobe of paramerite dilated apicad, rounded at apex, 1/4 as long as condylite.
Female: Eighth tergite (Fig. 79 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 79 B) with 12–13 macrosetae; sensory setae of 8th sternite
generalised, almost same as other setae in shape. Spermatheca (Figs. 79 C, 79 D) curved threetimes; basal part almost as long as apical part, acutely curved around middle, slightly dilated
apicad; apical part acutely curved around middle, narrowed or slightly dilated apicad, its innerwall densely wrinkled from apex to around apical 2/3.
Measurements. BL, 5.5–6.2; FBL, 2.7–2.8; HL, 0.81–0.85; HW, 0.95–1.04; EL, 0.301–
0.343; AL, 1.90–2.12; PL, 0.96–1.04; PW, 1.29–1.36; ELL, 1.10–1.20; ELW, 1.46–1.56; HTL,
1.09–1.12.
Bionomics. This species is commonly collected from colonies of Lasius (Dendrolasius)
spp. and widely distributed from lowlands to mountainous zones in Hokkaidô, Honshû, Shikoku
and Kyûshû. This species is sometimes collected from colonies of the other subgenera of Lasius,
but it may be accidental.
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) nipponensis, L. (D.)
capitatus.
Pella kishimotoi Maruyama, n. sp.
(Figs. 80–82)

Type material. Holotype: China (Hunan Sheng): ?, “Dahong-cun (1030 m), Da’an
Xiang, Longshan, Hunan Sheng, China, T. Kishimoto leg.” (NSMT).
Paratypes: 27 exs., same data as holotype; 15 exs., same data as holotype but, Yûki Imura(LDS ) (NSMT, TUA, IZAS, cMar).
Type locality. Da’an Xiang, Longshan, Hunan Sheng, China.
Distribution. Manchurian subregion. ASIA: China (Hunan Sheng).
Etymology. Dedicated to Dr. Toshio Kishimoto (Japan Wildlife Research Centre, Tokyo-to,
Japan), a collector of the type series, and a specialist of the Aleocharinae.
Diagnosis. Pella kishimotoi is similar to the other species of the cognata group in generalappearance, but can be distinguished from them by the apical lobe of aedeagal median lobe narrower in ventral view and longer. The species of the cognata group are locally isolated. No sym-

118

M. Maruyama

Fig. 80.

Pella kishimotoi Maruyama, n. sp. —A, Fore body, dorsal view; B, right antenna.

patric species have been known.
Description. Body slender. Blackish brown in ground colour; antennae, mouthparts, around
antero-lateral corner of each elytron, legs and 3rd to 5th abdominal segments reddish brown;
around antero-lateral corner of each elytron with yellow maculation. Head (Fig. 80 A) widest justbehind eyes; surface finely reticulated, densely covered with setae; setae short, as long as thoseon pronotum and elytra; length of eyes 0.34–0.36 times as long as head width. Antennae (Fig.
80 B) shorter than head, pronotum and elytra combined; 1st segment longer than 2nd and 3rd
combined; 2nd segment about 2/3 as long as 3rd; 3rd segment about 3/5 as long as 1st; 4th segment slightly longer than wide; 5th segment almost as long as wide; 6th to 10th segments wider
than long; 11th segment much shorter than 1st; approximate relative lengths of segments frombasal to apical: 32 : 11 : 17 : 7 : 7 : 7 : 7.5 : 7 : 8 : 10 : 26. Pronotum (Fig. 80 A) 1.25–1.37 times aswide as long, widest around anterior 1/3, narrowed posteriorly; posterior margin rounded; surface
finely punctured, finely reticulated among punctures, densely covered with setae, with 11 or 12
macrosetae along lateral margin. Scutellum with surface finely punctured, densely covered withsetae. Elytra (Fig. 80 A) slightly widened apicad; surface finely punctured, reticulated among
punctures, densely covered with setae, with 6–8 small macrosetae laterally. Legs somewhat short;hind tibia 0.86–0.94 times as long as elytra; approximate relative lengths of tarsal segments frombasal to apical: 7 : 6 : 6 : 12 in fore tarsus; 10.5 : 9 : 7 : 6.5 : 11.5 in mid tarsus; 12 : 6 : 6 : 5 : 10 inhind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 4th and 5th
segments; surface weakly reticulated; 3rd to 5th tergites glabrous except for a row of setae alongeach posterior margin; 6th and 7th tergites sparsely and uniformly covered with minute setae; 8th
tergite (Figs. 81 A, 82 A) with 7 (rarely 8) macrosetae; 9th tergite with 3 macrosetae; 10th tergite

Palearctic Species of Pella

119

Fig. 81. Pella kishimotoi Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal
view. Male.

with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 81 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 81 B) with posterior margin rounded, with 15–17 macrosetae; 9th sternite with posterior margin slightly emarginate. Aedeagus (Figs. 81 C, 81 D) oval inventral view; apical lobe curved ventrad, obtuse at apex in lateral view, rounded at apex in ventral
view; basal ridge roundly convex; copulatory piece of inner sac as in Fig. 81 E. Paramere withapical lobe of paramerite dilated apicad, rounded at apex, 1/4 as long as condylite.

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M. Maruyama

Fig. 82. Pella kishimotoi Maruyama, n. sp. A, 8th abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, D, spermathecae. Female.

Female: Eighth tergite (Fig. 82 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 82 B) with 14 or 15 macrosetae; sensory setae of 8th sternite generalised, almost the same as the other setae in shape. Spermatheca (Figs. 82 C, 82 D)
curved three times; basal part almost as long as or slightly longer than apical part, acutely curved
around basal 1/3, slightly dilated apicad; apical part acutely or gently curved around middle, narrowed or slightly dilated apicad, its inner wall densely wrinkled from apex to around apical 4/5.
Measurements. BL, 4.8–5.9; FBL, 2.4–2.9; HL, 0.68–0.79; HW, 0.81–0.92; EL, 0.290–
0.310; AL, 1.53–1.78; PL, 0.84–0.95; PW, 1.15–1.29; ELL, 0.96–1.14; ELW, 1.32–1.49; HTL,
0.90–0.98.
Bionomics. The type locality of Pella kishimotoi is a secondary deciduous forest situated at
the back of a small village in Longshan Xiang, northwestern Hunan Sheng (alt. 1,030 m). The
specimens were collected from the trails of Lasius (Dendrolasius) spathepus with other myrmecophilous staphylinids, Drusilla yunnanensis Pace, 1993, D. watanabei Maruyama & Kishimoto,2002, D. dendrolasii, Maruyama & Kishimoto, 2002.
Symbiotic host. Lasius (Dendrolasius) spathepus.

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121

The ruficollis GroupSpecies included. Pella ruficollis.
Distribution. European subregion, Mediterranean subregion.
Diagnosis. Species of the ruficollis group may be characterised by a combination of thefollowing character states: 1) eye length 0.30–0.31 times as long as head width; 2) 11th antennal
segment longer than the 1st; 3) pronotum with posterior margin completely rounded, not forming
corners postero-laterally; 4) pronotum 1.34–1.44 times as wide as long; 5) pronotum widest anteriorly; 6) pronotal hypomeron fully visible in lateral view; 7) elytron with yellowish brown zone
along inner and posterior margins; 8) inner and posterior margins of elytra weakly margined; 9)male 8th sternite longer than in female; 10) posterior margin of 8th tergite not crenate nor dentate, almost straight; 11) posterior margin of male 8th sternite rounded; 12) lateral projection of
apodeme of male 8th tergite not evidently longer than that of female; 13) macrosetae of 8th
abdominal segment generalised in length, exceeding the posterior margin of the segment; 14)postero-medial margin of female 8th sternite without minute pubescence; 15) apical lobe of
aedeagal median lobe almost straight and curved ventrad near apex in lateral view; 16) distal
crests of aedeagus well developed, projected, pointed at apex in lateral view; 17) copulatory pieceof aedeagal median lobe subparallel-sided from base to middle and gently narrowed apicad in
dorsal view.
Comments. The ruficollis group is a monotypical species-group. This species-group may
be allied to the coreana group in resemblance of general appearance except for its colour, especially the head convex posteriorly and the long 11th antennal segment. This species-group is very
unusual in the genus for the poor sexual dimorphism of abdominal tergites.
Symbiotic host. Liometopum microcephalum.
Pella ruficollis (Grimm, 1845), n. comb.
(Figs. 83–85)
Myrmedonia ruficollis Grimm, 1845: 126 (original description).—Mulsant & Rey, 1873a: 72 (Myrmedonia, description).—des Gozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 123 (Myrmedonia (Pella), key, description).—Reitter,
1909: 43 (Myrmedonia (Pellochromonia), key). — Fenyes, 1920: 298 (Zyras (Pellochromonia), list).—Bernhauer &Scheerpeltz, 1926: 704 (Zyras (Pellochromonia), list). —Scheerpeltz, 1934: 1659 (Zyras (Pellochromonia), list).—Lohse, 1974: 226 (Zyras (Pellochromonia), key). — Likovský, 1993: 59 (Zyras (Pellochromonia), list).—Smetana,
2004: 461 (Zyras (Pellochromonia), list).
Myrmedonia fernandi Fairmaire, 1855: 309 (original description).—Mulsant & Rey, 1873a: 73 (synonym of ruficollis,list).—des Gozis, 1886: 12 (Pella, list). — Ganglbauer, 1895: 123 (synonym of ruficollis, list).—Reitter, 1909: 43(synonym of ruficollis, list). — Fenyes, 1920: 298 (synonym of ruficollis, list).—Bernhauer & Scheerpeltz, 1926:
704 (synonym of ruficollis, list). — Smetana, 2004: 461 (synonym of ruficollis, list).

Type material. Myrmedonia ruficollis. Not found.
Myrmedonia fernandi. Holotype: Italy: sex?, “Fernandi Type Naples/Salermes” (IRSNB).
Type locality. Berlin, Germany.
Other material. Croatia: Zagreb, 23 IV 1917, Hochetlinger (LM) (NMP). Italy: 2 exs.,
Son Basilio, Murgien, Paganetti; 2 exs., Grottaglie, Murgien, Paganetti (cMar, ex NHMW); 2
exs., Grottaglie, Murgien, Paganetti (cMar, ex NHMW); 4 exs., S. Basilio, Puglia Italia, 9 V
1989, F. Angelini (MNV). Italy (Sicilia): 1 ex., Sicilia, Schramm (no other data) (NMP). Slovakia: 1 ex., Jur pri Bret., 19 V 1963, M. Dvořák (LM) (cMar, ex cDov); 1 ex., Bratislava, 10 V
1986, I. Rychlík (cMar, ex cHlav).
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Bulgaria,

122

M. Maruyama

Fig. 83.

Pella ruficollis (Grimm). — A, Fore body, dorsal view; B, right antenna.

Croatia, France, Greece, Hungary, Italia, Macedonia, Romania, Slovakia, Turkey.
Diagnosis. Pella ruficollis can easily be distinguished from all the other species of Pella by
the peculiar body colour, specially the reddish orange pronotum.
Redescription. Body robust. Blackish brown in ground colour; antennae, mouthparts andlegs reddish brown; pronotum reddish orange; elytron with yellowish brown zone along inner andposterior margins; around posterior margins of 3rd to 5th abdominal segments yellowish brown.
Head (Fig. 83 A) widest at middle of eyes, convex posteriorly; surface smooth, densely covered
with setae; setae short, as long as those on pronotum and elytra; length of eyes 0.30–0.31 times
as long as head width. Antennae (Fig. 83 B) shorter than head, pronotum and elytra combined;
1st segment much longer than 2nd and 3rd combined; 2nd segment slightly longer than 3rd; 3rdsegment about 1/3 as long as 1st; 4th to 10th segments wider than long; 11th segment much
longer than 1st; approximate relative lengths of segments from basal to apical: 29 : 11 : 10 : 6.5 :6 : 6 : 7 : 7 : 8 : 8.5 : 36. Pronotum (Fig. 83 A) 1.34–1.44 times as wide as long, widest around middle, narrowed posteriorly; posterior margin rounded; surface smooth, densely covered with setae,
with 11 or 12 small macrosetae along lateral margin. Scutellum with surface smooth, densely
covered with setae. Elytra (Fig. 83 A) slightly widened apicad; surface smooth, densely covered
with setae, with 6–8 small macrosetae laterally. Legs short; hind tibia 0.77–0.79 times as long as
elytra; approximate relative lengths of tarsal segments from basal to apical: 11 : 8 : 6.5 : 12 in foretarsus; 15 : 9.5 : 7.5 : 7 : 12 in mid tarsus; 20 : 10 : 9 : 8 : 12.5 in hind tarsus. Abdomen subparallelsided, almost as wide as elytra, widest around 4th and 5th segments; surface smooth; 3rd to 6th
tergites densely covered with setae; 7th tergite sparsely and uniformly covered with minute setae;
8th tergite (Figs. 84 A, 85 A) with 9 (rarely 8) macrosetae; 9th tergite with 5 macrosetae; 10th

Palearctic Species of Pella

123

Fig. 84. Pella ruficollis (Grimm). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

tergite with posterior margin slightly truncate, with 4 macrosetae.
Male: Eighth tergite (Fig. 84 A) with posterior margin truncate; 8th sternite (Fig. 84 B) withposterior margin rounded, with 21–25 macrosetae; 9th sternite with posterior margin truncate.
Aedeagus (Figs. 84 C, 84 D) oval in ventral view; apical lobe curved ventrad near apex, pointed
at apex in lateral view, slightly pointed at apex in ventral view; basal ridge absent; copulatory
piece of inner sac as in Fig. 84 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 85 A) with posterior margin truncate; 8th sternite (Fig. 85 B)
with 17–19 macrosetae; sensory setae of 8th sternite spindle-shaped, flattened. Spermatheca (Fig.
85 C) curved three times; basal part about 10 times as long as apical part, slightly bulbous aroundbase, acutely curved around basal 1/3; apical part dilated apicad, its inner wall densely wrinkledfrom apex to around apical 2/3.
Measurements. BL, 5.2–6.0; FBL, 2.5–2.7; HL, 0.70–0.81; HW, 0.99–1.06; EL, 0.311–

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M. Maruyama

Fig. 85. Pella ruficollis (Grimm). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

0.318; AL, 1.78–1.95; PL, 0.89–1.01; PW, 1.34–1.44; ELL, 1.14–1.23; ELW, 1.58–1.81; HTL,
0.88–0.98.
Bionomics. Pella ruficollis is collected from nests of Liometopum microcephalum sometimes together with P. hampei. This species occurs very locally and rarely in Central Europe
probably due to rarity of the host ants. This species is very similar to the host ant in colorationand probably a Batesian mimic. Although the profile of defence substances of Liometopum microcephalum is unknown, this ant probably has repellent taste against predator. No information
was available for the life cycle and the immature stages of this species.
Symbiotic host. Liometopum microcephalum.
The coreana GroupSpecies included. Pella coreana, P. plutenkoi.
Distribution. Manchurian subregion.
Diagnosis. Species of the coreana group may be characterised by a combination of the following character states: 1) eye length 0.30–0.33 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with posterior margin completely rounded, not forming corners postero-laterally; 4) pronotum 1.48 times as wide as long; 5) pronotum widest aroundmiddle; 6) pronotal hypomeron partly visible in lateral view, its visible length less than 3/5 times
as long as pronotal length; 7) elytra unicoloured, chestnut brown; 8) inner and posterior margins
of elytra weakly margined; 9) posterior margin of 8th tergite crenate; 10) macrosetae of 8th abdominal segment short, which not exceed the posterior margin of the segment; 11) postero-medial margin of female 8th sternite with minute pubescence.

Palearctic Species of Pella

125

Comments. The coreana group may be allied to the ruficollis group in resemblance of general appearance except for its colour, particular the head convex posteriorly and the long 11th antennal segment. This species-group may be easily distinguished from the other species-groups by
the short macrosetae of 8th abdominal segments, which do not exceed the posterior margins of
the segments. This character state may be an autapomorphy of this species-group.
Symbiotic host. Unknown.
Key to the Species of the coreana Group——

Antennae longer than pronotal width; basal part of spermatheca longer and curved twice.
Distribution: Korea. ................................................................................................P. coreana.
Antennae shorter than pronotal width; basal part of the spermatheca shorter and curved
once. Distribution: Russia (Far East). ...................................................................P. plutenkoi.
Pella coreana Maruyama, n. sp.
(Figs. 86, 87)

Type material. Holotype: Korea: /, “Korea: Seoul, Nowon, Sanggye-dong, Mt. Suraksan,
14 IV 1988, T. S. Kim” (CNU).
Type locality. Seoul, Korea.
Distribution. Manchurian subregion. ASIA: Korea.
Etymology. Named after the type locality.
Diagnosis. Pella coreana is most similar to P. plutenkoi in general appearance and shape of

Fig. 86.

Pella coreana Maruyama, n. sp. — A, Fore body, dorsal view; B, right antenna.

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M. Maruyama

Fig. 87. Pella coreana Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

spermatheca, but is distinguished from the latter by the longer antennae, which are longer thanthe pronotal width, and by the basal part of the spermatheca longer and curved three times.
Among the sympatric species, Pella coreana may be easily distinguished by the 11th antennal
segment longer than the combined length of 7th to 10th antennal segments.
Description. Body robust. Pale brown in ground colour; head slightly darker; 3rd to 7h tergites brown except around posterior margins. Head (Fig. 86 A) widest just behind eyes, convex
posteriorly; surface smooth, densely covered with setae; setae moderate in length, as long as
those on pronotum and elytra; length of eyes 0.30 times as long as head width. Antennae (Fig.
86 B) shorter than head, pronotum and elytra combined; 1st segment much longer than 2nd and3rd combined; 2nd segment as long as 3rd; 4th to 10th segments wider than long; 11th segment
shorter than 1st; approximate relative lengths of segments from basal to apical: 32.5 : 14 : 14 :6.5 : 6 : 4 : 5 : 4.5 : 5 : 6 : 30. Pronotum (Fig. 86 A) 1.48 times as wide as long, widest around middle; posterior margin rounded; surface finely punctured, finely reticulated among punctures,
densely covered with setae, with 9 or 10 small macrosetae along lateral margin. Scutellum withsurface smooth, densely covered with setae. Elytra (Fig. 86 A) slightly widened apicad; surface
finely punctured, finely reticulated among punctures, densely covered with setae, with 4 or 5
small macrosetae laterally. Legs short; hind tibia 0.86 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 10 : 6 : 7 : 14 in fore tarsus; 10 : 9 : 8 : 8 : 11.5

Palearctic Species of Pella

127

in mid tarsus; 16 : 10 : – : – : – in hind tarsus. Abdomen subparallel-sided, almost as wide as elytra,
widest around 4th and 5th segments; surface finely reticulated; 3rd tergite glabrous except for arow of setae along posterior margin; 4th to 7th tergites sparsely and uniformly covered withminute setae; 8th tergite (Fig. 87 A) with 7 macrosetae; 9th tergite with 5 macrosetae; 10th tergite
with posterior margin slightly truncate, with 4 macrosetae; macrosetae on tergites poorly differentiated from other setae.
Male unknown.
Female: Eighth tergite (Fig. 87 A) with posterior margin truncate, its truncate apex minutelydentate; 8th sternite (Fig. 87 B) with 7 macrosetae; sensory setae of 8th sternite generalised, similar to the other setae in shape. Spermatheca (Fig. 87 C) curved three times; basal part about 4.5
times as long as apical part, slightly bulbous around base, gently curved at basal 1/4, and acutely
curved around middle; apical part dilated apicad, much broader than basal part, its inner wallcompletely and moderately wrinkled.
Measurements. BL, 6.6; FBL, 3; HL, 0.88; HW, 1.07; EL, 0.325; AL, 1.80; PL, 1.08; PW,
1.60; ELL, 1.35; ELW, 1.86; HTL, 1.15.
Bionomics and symbiotic host. No information.
Pella plutenkoi Maruyama, n. sp.
(Figs. 88, 89)

Type material. Holotype: Russia (Far East): /, “RUSSIA: Primorie/S Artyom town env.,
100–300 m Ozemyi Kluyich riv. 20. IV.–30. V. 1976 leg. A. Plutenko/Coll. M. Schulke Berlin
Ankauf A. Plutenko Oktober 2002” (cSch).
Type locality. Artyom, Primorskyi Krai, Russia.
Distribution. Manchurian subregion. ASIA: Russia (Far East).
Etymology. Dedicated to Mr. Andrey Vital’evich Plutenko (Smolensk, Russia), collector of
the holotype, and a specialist of the Carabidae.
Diagnosis. Pella plutenkoi is most similar to P. coreana in general appearance and shape of
spermatheca, but is distinguished from the latter by the shorter antennae, which are shorter thanthe pronotal width, and the basal part of the spermatheca shorter and curved once. Among the
sympatric species, Pella plutenkoi may be easily distinguished by the 11th antennal segment
longer than the combined length of 7th to 10th antennal segments.
Description. Body robust. Pale brown in ground colour; head slightly darker; mesal areasof 3rd to 6th tergites brown. Head (Fig. 88 A) widest just behind eyes, convex posteriorly; surface
smooth, densely covered with setae; setae moderate in length, as long as those on pronotum andelytra; length of eyes 0.33 times as long as head width. Antennae (Fig. 89 A) shorter than head,pronotum and elytra combined , shorter than pronotal width; 1st segment much longer than 2nd
and 3rd combined; 2nd segment as long as 3rd; 4th to 10th segments wider than long; 11th segment shorter than 1st; approximate relative lengths of segments from basal to apical: 29.5 : 10 :12 : 6 : 5 : 4 : 4.5 : 4 : 4 : 5 : 28. Pronotum (Fig. 88 A) 1.48 times as wide as long, widest aroundmiddle; posterior margin rounded; surface finely punctured, finely reticulated among punctures,
densely covered with setae, with 9 or 10 small macrosetae along lateral margin. Scutellum withsurface smooth, densely covered with setae. Elytra (Fig. 88 A) slightly widened apicad; surface
finely punctured, finely reticulated among punctures, densely covered with setae, with 4 or 5
small macrosetae laterally. Legs short; hind tibia 0.83 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 8 : 6.5 : 6 : 12.5 in fore tarsus; 11 : 9 : 7.5 : 7.5 :128

M. Maruyama

Fig. 88.

Pella plutenkoi Maruyama, n. sp. — A, Fore body, dorsal view; B, right antenna.

11 in mid tarsus; 15 : 10 : 9.5 : 9 : 11 in hind tarsus. Abdomen subparallel-sided, almost as wide as
elytra, widest around 4th and 5th segments; surface finely reticulated; 3rd tergite glabrous except
for a row of setae along posterior margin; 4th to 7th tergites sparsely and uniformly covered withminute setae; 8th tergite (Fig. 89 A) with 7 or 8 macrosetae; 9th tergite with 5 macrosetae; 10th
tergite with posterior margin slightly truncate, with 4 macrosetae; macrosetae on tergites poorly
differentiated from other setae.
Male unknown.
Female: Eighth tergite (Fig. 89 A) with posterior margin widely emarginate, its truncate apex
minutely dentate; 8th sternite (Fig. 89 B) with 7 macrosetae; sensory setae of 8th sternite generalised, similar to other setae in shape. Spermatheca (Fig. 89 C) curved two times; basal part about
three times as long as apical part, slightly bulbous around base, acutely curved around middle;
apical part dilated apicad, much broader than basal part, its inner wall completely and moderately
wrinkled.
Measurements. BL, 6.3; FBL, 2.9; HL, 0.81; HW, 1.01; EL, 0.338; AL, 1.48; PL, 1.05;
PW, 1.55; ELL, 1.28; ELW, 1.73; HTL, 1.08.
Bionomics and symbiotic host. No information.
The excepta GroupSpecies included. Pella excepta, P. maura, P. kuluensis, P. bohaci, P. cinctipennis.
Distribution. Mediterranean subregion.
Diagnosis. Species of the excepta group may be characterised by a combination of the fol-Palearctic Species of Pella

129

Fig. 89. Pella plutenkoi Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

lowing character states: 1) eye length 0.34–0.37 times as long as head width; 2) 11th antennal
segment variable in length, longer to shorter than the 1st; 3) pronotum with posterior margincompletely rounded, not forming corners postero-laterally; 4) pronotum 1.28–1.43 times as wide
as long; 5) pronotum widest anteriorly or around middle; 6) pronotal hypomeron partly visible inlateral view, its visible length less than 3/5 times as long as pronotal length; 7) unicoloured, yellowish brown to blackish brown; 8) inner and posterior margins of elytra distinctly margined; 9)male 8th sternite longer than in female; 10) posterior margin of 8th tergite crenate; 11) posterior
margin of male 8th sternite rounded; 12) lateral projection of apodeme of male 8th tergite
evidently longer than that of female; 13) macrosetae of 8th abdominal segment generalised inlength, exceeding the posterior margin of the segment; 14) postero-medial margin of female 8thsternite without minute pubescence; 15) apical lobe of aedeagal median lobe gently curved
ventrad in lateral view; 16) distal crests of aedeagus well developed, projected semicircularly inlateral view; 17) copulatory piece of aedeagal median lobe slender, subparallel-sided or spindleshaped in dorsal view.
Comments. The excepta group can be regarded as a monophyletic group by autapomorphies: the apical lobe of aedeagal median lobe more or less elongate, constricted at base andpointed at apex in lateral view, and the body surface relatively sparsely covered with setae. These130

M. Maruyama

states are unique within the genus and not observed in the other species of the Lomechusini.
Thus, these character states are considered to be autapomorphies of this species-group. No allied
species-group could be inferred. Although the large eyes and the pronotal shape are somewhat
similar to the spreta group, these states are considered to be homoplastic.
Symbiotic hosts. Crematogaster spp.
Key to the Species of the excepta Group
1. Body large: BL: 5.1–5.8 mm, FBL: 2.2–2.6 mm; pronotum with a circular depression postero-medially in male; 3rd and 4th tergites with minute projection postero-medially in male.
Distribution: South Europe, North Africa. .......................................................................... 2.
— Body medium to small: BL: 3.7–4.8 mm, FBL: 1.7–2.2; pronotum without a circular depression postero-medially in male; 3rd and 4th tergites without minute projection posteromedially in male. Distribution: Turkey, Central Asia, Pakistan. ......................................... 3.
2. Body reddish brown; posterior margin of male 8th tergite truncate and bisinuate; apical lobeof aedeagal median lobe gently curved apicad. Distribution: France, Spain. ........ P. excepta.
— Body blackish brown; posterior margin of male 8th tergite rounded and quadrisinuate; apical lobe of aedeagal median lobe much narrowed near apex and curved ventrad. Distribution: Italy (Sicilia), Morocco. ....................................................................................P. maura.
3. Eleventh antennal segment much longer than 1st. Distribution: Pakistan. ...........P. kuluensis.
— Eleventh antennal segment shorter than or as long as 1st. Distribution: Turkey, Central Asia.
............................................................................................................................................... 4
4. Body smaller: BL: 3.7–4.2 mm, FBL: 1.7–1.8 mm; pronotum wider, 1.37–1.43 times aswide as long; 8th tergite with 7 macrosetae; apical lobe of aedeagal median lobe almost as
long as basal capsule; basal part of spermatheca spherical at base. .................P. cinctipennis.
— Body larger: BL: 4.4–4.8 mm, FBL: 1.4–1.6 mm; pronotum narrower, 1.26–1.36 times aswide as long; 8th tergite with 6 macrosetae; apical lobe of aedeagal median lobe much
longer than basal capsule; basal part of spermatheca slightly dilated around base. ................
...................................................................................................................................P. bohaci.
Pella excepta (Mulsant & Rey, 1861), n. comb.
(Figs. 90–92)
Myrmedonia excepta Mulsant & Rey, 1861: 113 (original description).—Mulsant & Rey, 1873a: 86 (Myrmedonia(Myrmelia), description). — Ganglbauer, 1895: 120 (Myrmedonia (Myrmelia), key, description).—Bernhauer &Scheerpeltz, 1926: 697 (Zyras (Myrmelia), list). — Scheerpeltz, 1934: 1653 (Zyras (Myrmelia), list).—Fenyes, 1920:
299 (Zyras (Myrmelia), list). — Smetana, 2004: 460 (Zyras (Myrmelia), list).

Type material. Syntype: France: ?, “?/Coll. C. REY 1884 Muséum de Lyon” (MHNL).
Type locality. Marseille, France (after the original description).
Other material. France: 1?, “Mont. Alaric Aude.” (NMP); 1/ “Joug de l’Aigle (val)
1.146” (MNHN); 1?, “St-Michel-l’Obs. (B.-A.) C. Dufay 8. 50” (MNHN); 1/ “excepta Rey/extype” (MNHN); 1?, “France Avril/Myrmedonia excepta Rey ?” (IRSNB); 1/, “Hyèus/
Carthagène/excepta Rey ex typ. /” (IRSNB). Spain: 1?, “Hispania Umb. v. Madrid/Zyras exceptus Rey” (NHMW); 1/, N-Hang, Sierra de Alhamilla, Andalusia, 20 III 1994, V. Assing(cAss).
The female specimens labeled “ex type” deposited in IRSNB and MNHN are actually not a

Palearctic Species of Pella

Fig. 90.

131

Pella excepta (Mulsant & Rey). —A, Fore body, dorsal view; B, right antenna.

part of the type series, since the type series are represented by only male material (Mulsant &Rey, 1861). Moreover, in their redescription of this species, Mulsant and Rey (1873a) commented “/. Nous est inconnue”.
Distribution. Mediterranean subregion. EUROPE: France, Spain.
Diagnosis. Pella excepta is closely similar to P. maura in general appearance, but is distinguished from the latter by the body reddish brown and the apical lobe of aedeagal median lobe
shorter. These two species are locally isolated. No sympatric species have been recorded.
Redescription. Body slender. Reddish brown in ground colour; head, antennae, mouthparts, pronotum, elytra and legs paler. Head (Fig. 90 A) widest just behind eyes; surface smooth,moderately covered with setae; setae moderate in length, as long as those on pronotum and elytra; length of eyes 0.35–0.36 times as long as head width. Antennae (Fig. 90 B) shorter than head,pronotum and elytra combined; 1st segment slightly longer than 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments wider
than long; 11th segment shorter than 1st; approximate relative lengths of segments from basal toapical: 25.5 : 11 : 13.5 : 8 : 8.5 : 8 : 8 : 8.5 : 7.5 : 7.5 : 22.5. Pronotum (Fig. 90 A) 1.38–1.41 times aswide as long, widest around anterior 1/6, narrowed posteriorly; posterior margin rounded; surface
smooth, moderately covered with setae, with 7 or 8 macrosetae along lateral margin. Scutellum
with surface smooth, moderately covered with setae. Elytra (Fig. 90 A) slightly widened apicad;
surface smooth, moderately covered with setae, with 3 or 4 small macrosetae laterally. Legs moderate in length; hind tibia 0.90–0.99 times as long as elytra; approximate relative lengths of tarsal
segments from basal to apical: 6 : 5 : 4.5 : 12 in fore tarsus; 8 : 8 : 6 : 5 : 10.5 in mid tarsus; 13 : 8 :7.5 : 6.5 : 11.5 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest

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M. Maruyama

Fig. 91. Pella excepta (Mulsant & Rey). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

around 4th and 5th segments; 3rd and 4th tergites with minute projection postero-medially; 7th
tergite with a ridge postero-medially which is about 1/2 as long as 7th tergite; surface smooth;
3rd to 7th tergites sparsely and uniformly covered with setae; 8th tergite (Figs. 91 A, 92 A) with 6
macrosetae; 9th tergite with 3 macrosetae; 10th tergite with posterior margin slightly emarginate,with 3 macrosetae.
Male: Eighth tergite (Fig. 91 A) with posterior margin somewhat truncate, its truncate apex
slightly bisinuate and crenate; 8th sternite (Fig. 91 B) with posterior margin rounded, with 8–10
macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs. 91 C, 91 D) somewhat
pear-shaped in ventral view; apical lobe curved ventrad near apex, pointed at apex in lateral view,
obtuse at apex in ventral view; basal ridge roundly convex; copulatory piece of inner sac as inFig. 91 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, 1/4 as long as
condylite.

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133

Fig. 92. Pella excepta (Mulsant & Rey). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

Female: Eighth tergite (Fig. 92 A) with posterior margin slightly truncate, its truncate apex
crenate; 8th sternite (Fig. 92 B) with 6 macrosetae; sensory setae of 8th sternite spindle-shaped,
flattened. Spermatheca (Fig. 92 C) curved three times; basal part almost slightly longer than apical part, acutely curved around middle; apical part gently curved around basal 1/3, narrowed
apicad, its inner wall densely wrinkled from apex to around apical 3/5.
Measurements. BL, 5.1–5.2; FBL, 2.2–2.3; HL, 0.61–0.65; HW, 0.81–0.82; EL, 0.286–
0.298; AL, 1.75–1.80; PL, 0.74–0.75; PW, 1.03–1.04; ELL, 0.87–0.91; ELW, 1.19–1.22; HTL,
0.81–0.86.
Bionomics. The type series of P. excepta were collected with ants at root of a tree
(Mulsant & Rey, 1861), but the ant species was not specified.
Symbiotic host. Unknown.
Pella maura (Fauvel, 1898), n. comb.
(Figs. 93–95)
Myrmedonia maura Fauvel, 1898: 110 (original description).—Fenyes, 1920: 299 (Zyras (Myrmelia), list).—Bernhauer& Scheerpeltz, 1926: 701 (Zyras (Myrmelia), list). — Smetana, 2004: 461 (Zyras (Myrmelia), list).
Myrmedonia Ragusae Ragusa, 1921: 88 (original description, as “Myrmedonia Ragusae Bernhauer”).—Bernhauer &Scheerpeltz, 1926: 701 (synonym of maura). — Smetana, 2004: 461 (synonym of maura).

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M. Maruyama

Fig. 93.

Pella maura (Fauvel). — A, Fore body, dorsal view; B, right antenna.

Type material. Myrmedonia maura. Not examined.
Myrmedonia Ragusae. Holotype: Italy (Sicilia): ?, “Sicilia/Myrmedonia Ragusae Bh. Type”(NMP).
I examined a specimen labelled “Milianah”, the type locality of Myrmedonia maura, and it
was identified as a syntype of M. maura by Dr. D. Drugmand of IRSNB. However, the specimen
did not fit with the description of this species in some features, particularly in body length, and I
did not regard it as syntype of Myrmedonia maura. It was Pella erratica, and its data is describedin “Other material” of P. erratica.
Type locality. Milianah, Algeria.
Other material. Italy (Sicilia): 1?, “Sicilia M. Alto E. Ragusa” (MNHN); 1/, “M. Alto
Sicilia” (MNHN); 1?, “ITALIE Madonia/Zyras Ragusae/ex coll. Bettinger”. Morocco: 1/,“Midelt Hoher Atlas Marokko/Sadostl. von Azrou Hochsteppe/unfer einem Stein./leg. Dr. R.
Ebner 2. VI. 1930/maurus ?? Fauv.” (NHMW).
Distribution. Mediterranean subregion. EUROPE: Italy (Sicilia); NORTH AFRICA: Morocco.
Diagnosis. Pella maura is closely similar to P. excepta in general appearance but distinguished from the latter by the body blackish brown and the apical lobe of the aedeagal median
lobe longer. These two species are locally isolated. Pella maura can be distinguished from thesympatric species by the presence of the abdominal projections and ridge in male.
Redescription. Body slender. Blackish brown in ground colour; head, antennae, mouthparts, pronotum, elytra, legs reddish brown. Head (Fig. 93 A) widest just behind eyes; surface
smooth, moderately covered with setae; setae moderate in length, as long as those on pronotum

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135

Fig. 94. Pella maura (Fauvel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

and elytra; length of eyes 0.34–0.35 times as long as head width. Antennae (Fig. 93 B) shorter
than head, pronotum and elytra combined; 1st segment slightly longer than 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th
segments wider than long; 11th segment shorter than 1st; approximate relative lengths of segments from basal to apical: 25 : 10.5 : 7.5 : 7 : 9 : 9 : 9 : 9.5 : 8.5 : 27. Pronotum (Fig. 93 A)
1.40–1.41 times as wide as long, widest around anterior 1/5, narrowed posteriorly; posterior margin rounded; surface smooth, moderately covered with setae, with 7 or 8 macrosetae along lateral
margin. Scutellum with surface smooth, moderately covered with setae. Elytra (Fig. 93 A) slight-

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M. Maruyama

Fig. 95. Pella maura (Fauvel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

ly widened apicad; surface smooth, moderately covered with setae, with 3 or 4 small macrosetae
laterally. Legs moderate in length; hind tibia 0.95–0.98 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 5 : 5 : 4.5 : 14 in fore tarsus; 10 : 9 : 7 : 5 : 12 in
mid tarsus; 16 : 12 : 10 : 7 : 13 in hind tarsus. Abdomen subparallel-sided, slightly narrower thanelytra, widest around 4th and 5th segments; 3rd and 4th tergites with minute projection posteromedially; 7th tergite with ridge postero-medially which is about 1/2 as long as 7th tergite; surface
smooth; 3rd to 7th tergites sparsely and uniformly covered with setae; 8th tergite (Figs. 94 A,
95 A) with 6 or 7 macrosetae; 9th tergite with 3 macrosetae; 10th tergite with posterior margin
slightly emarginate, with 3 macrosetae.
Male: Eighth tergite (Fig. 94 A) with posterior margin somewhat rounded, its apex slightly
quadrisinuate and dentate; 8th sternite (Fig. 94 B) with posterior margin rounded, with 11–13
macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs. 94 C, 94 D) somewhat
pear-shaped in ventral view; apical lobe curved ventrad near apex, pointed at apex in lateral view,rounded at apex in ventral view; basal ridge roundly convex; copulatory piece of inner sac as inFig. 94 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, 1/4 as long as
condylite.
Female: Eighth tergite (Fig. 95 A) with posterior margin rounded, its rounded apex crenate;
8th sternite (Fig. 95 B) with 8–10 macrosetae; sensory setae of 8th sternite spindle-shaped, flattened. Spermatheca (Fig. 95 C) curved three times; basal part almost slightly longer than apicalPalearctic Species of Pella

137

part, somewhat bulbous around base, acutely curved around middle; apical part gently curved
around basal 1/3, narrowed apicad, its inner wall densely wrinkled from apex to around apical
3/5.
Measurements. BL, 5.2–5.8; FBL, 2.4–2.6; HL, 0.68–0.69; HW, 0.87–0.91; EL, 0.302–
0.313; AL, 1.90–1.95; PL, 0.81–0.84; PW, 1.14–1.18; ELL, 1.00–1.03; ELW, 1.33–1.40; HTL,
0.98–0.99.
Bionomics and symbiotic host. No information.
Pella kuluensis (Cameron, 1939), n. comb.
(Figs. 96, 97)
Zyras (Pella) kuluensis Cameron, 1939: 545 (original description).—Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Holotype: Pakistan: ?, “Type (red round curator label)/Naggar Kulu. Punjab. 5,000 ft. H.G.C. [H.G. Champion]/Z. kuluensis TYPE Cam (BMNH).
Type locality. Naggar, Kulu, Punjab, Pakistan.
Other material. Not examined.
Distribution. Mediterranean subregion. ASIA: Pakistan.
Diagnosis. Pella kuluensis is closely similar to P. bohaci and P. cinctipennis but may beeasily distinguished from the latter two by the 11th antennal segment much longer than the 1st.
This species is locally isolated from P. bohaci and P. cinctipennis.
Redescription. Body slender. Brown in ground colour; antennae, posterior margin of
pronotum, around posterior margins of 3rd to 5th abdominal paratergites yellowish brown. Head

Fig. 96.

Pella kuluensis (Cameron). — A, Fore body, dorsal view; B, right antenna.

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M. Maruyama

Fig. 97. Pella kuluensis (Cameron). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.

(Fig. 96 A) widest just behind eyes; surface smooth, moderately covered with setae; setae moderate in length, as long as those on pronotum and elytra; length of eyes 0.36 times as long as headwidth. Antennae (Fig. 96 B) shorter than head, pronotum and elytra combined; 1st segment aslong as 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 aslong as 1st; 4th to 10th segments wider than long; 11th segment as long as 1st; approximate relative lengths of segments from basal to apical: 21 : 9 : 10 : 5.5 : 5 : 4.5 : 5 : 5 : 5 : 5 : 24.5. Pronotum(Fig. 96 A) 1.41 times as wide as long, widest around middle; posterior margin rounded; surface
smooth, sparsely covered with setae, with 7 or 8 macrosetae along lateral margin. Scutellum withsurface smooth, moderately covered with setae. Elytra (Fig. 96 A) slightly widened apicad; surface smooth, moderately covered with setae, with 4 small macrosetae laterally. Legs moderately

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139

short; hind tibia 0.80 times as long as elytra; approximate relative lengths of tarsal segments frombasal to apical: 4.5 : 4.5 : 5 : 10.5 in fore tarsus; 6 : 6 : 6 : 5 : 10.5 in mid tarsus; 10 : 8.5 : 6.5 : 5 : 10
in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 5th and6th segments; surface smooth; 3rd to 7th tergites glabrous except around a row of setae posterior
margin; 8th tergite (Fig. 97 A) with 6 (sometimes 7) macrosetae; 9th tergite with 3 macrosetae;
10th tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 97 A) with posterior margin somewhat rounded, its apex slightly
quadrisinuate and weakly crenate; 8th sternite (Fig. 97 B) with posterior margin rounded, with
14–16 macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs. 97 C, 97 D)
somewhat pear-shaped in ventral view; apical lobe curved ventrad in lateral view, pointed at apex
in lateral and ventral views; basal ridge roundly convex; copulatory piece of inner sac as in Fig.
97 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, 1/4 as long as
condylite.
Female unknown.
Measurements. BL, 4.6; FBL, 2.2; HL, 0.64; HW, 0.79; EL, 0.287; AL, 1.38; PL, 0.73;
PW, 1.03; ELL, 0.96; ELW, 1.29; HTL, 0.78.
Bionomics and symbiotic host. No information.
Pella bohaci (Dvořák, 1984), n. comb.
(Figs. 98–100)
Zyras (Pella) bohaci Dvořák, 1984: 195 (original description).—Smetana, 2004: 461 (Zyras (Pella), list).
Zyras (Pella) almaatensis Pace, 2002: 195 (original description). N. syn.

Type material. Zyras (Pella) bohaci. Paratypes: Uzbekistan: 1?, “USSR-Uzbekistan
Šafirkan pr. Buchere Kyźy-kum des. J. Bohač legt. IV. 79/PARATYPUS/Zyras (Pella) bohaci sp.
n. det. M. Dvořák ,81”; 1/, same data but, “ALLOTYPUS”.
These are part of type series. All the type specimens are deposited in cDvor. Each specimen
was pinned with ant specimen.
Zyras (Pella) almaatensis. Holotype: Kazakhstan: /, “KASACHSTAN Alma-Ata-Gebiet
Pristan Dubins-kaja 550 m NN 43°45N 80°13E 15. —18. VI. 1993 leg. V. & A.
Lukhtanov/HOLOTYPUS Zyras almaatensis det. R. Pace 2001/Zyras almaatensis n. sp. det. R.
Pace 2001” (NMEG).
The shape of the spermatheca in the original description (Pace, 2002) appears to be considerably different from that of Pella bohaci illustrated herein. However, the spermatheca of the type
mounted on slide, based on which illustration was prepared, was twisted 180-degree with thebasal and apical parts.
Other material. Turkmenistan: 2??, Tadzen, 12–25 III 1992, Snizek (NHMW, cMar).
Type locality. Buchere, Uzbekistan.
Distribution. Mediterranean subregion. ASIA: Kazakhstan, Turkmenistan, Uzbekistan.
New to Kazakhstan.
Diagnosis. Pella bohaci is closely similar to P. kuluensis and P. cinctipennis, but may beeasily distinguished from the latter two by the dorsal surface of the body yellowish brown. These
three species are locally isolated.
Redescription. Body slender. Brown in ground colour; head, antennae, mouthparts, pronotum, elytra, legs and around posterior margins of 3rd to 5th tergites yellowish brown. Head (Fig.
98A) widest just behind eyes; surface smooth, moderately covered with setae; setae moderate in

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M. Maruyama

Fig. 98.

Pella bohaci (Dvořák). — A, Fore body, dorsal view; B, right antenna.

length, as long as those on pronotum and elytra; length of eyes 0.35–0.36 times as long as headwidth. Antennae (Fig. 98 B) shorter than head, pronotum and elytra combined; 1st segment aslong as 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 aslong as 1st; 4th to 10th segments wider than long; 11th segment as long as 1st; approximate relative lengths of segments from basal to apical: 21 : 10 : 11 : 5.5 : 6 : 5 : 6 : 5 : 5 : 6 : 21. Pronotum(Fig. 98 A) 1.37–1.43 times as wide as long, widest around anterior 1/4, narrowed posteriorly;
posterior margin rounded; surface smooth, sparsely covered with setae, with 7 or 8 macrosetae
along lateral margin. Scutellum with surface smooth, moderately covered with setae. Elytra (Fig.
98 A) slightly widened apicad; surface smooth, moderately covered with setae, with 3 or 4 small
macrosetae laterally. Legs short; hind tibia 0.83–0.85 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 7 : 6 : 6 : 14 in fore tarsus; 9 : 9 : 8 : 7 : 12 in
mid tarsus; 12 : 10 : 9 : 7 : 12.5 in hind tarsus. Abdomen subparallel-sided, slightly narrower thanelytra, widest around 5th and 6th segments; surface smooth; 3rd to 7th tergites sparsely and uniformly covered with setae; 8th tergite (Figs. 99 A, 100 A) with 6 (sometimes 7) macrosetae; 9th
tergite with 3 macrosetae; 10th tergite with posterior margin slightly emarginate, with 3 macrosetae.
Male: Eighth tergite (Fig. 99 A) with posterior margin somewhat truncate, its truncate apex
weakly crenate; 8th sternite (Fig. 99 B) with posterior margin rounded, with 14–16 macrosetae;
9th sternite with posterior margin truncate. Aedeagus (Figs. 99 C, 99 D) somewhat pear-shaped in
ventral view; apical lobe curved ventrad in lateral view, pointed at apex in lateral and ventral
views; basal ridge roundly convex; copulatory piece of inner sac as in Fig. 99 E. Paramere withapical lobe of paramerite dilated apicad, rounded at apex, 1/4 as long as condylite.

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141

Fig. 99. Pella bohaci (Dvořák). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

Female: Eighth tergite (Fig. 100 A) with posterior margin slightly truncate, its truncate apex
crenate; 8th sternite (Fig. 100 B) with 13 or 14 macrosetae; sensory setae of 8th sternite spindleshaped, flattened. Spermatheca (Fig. 100 C) curved three times; basal part almost as long as apical part, acutely curved around middle, slightly narrowed apicad; apical part gently curved
around base, narrowed apicad, its inner wall densely wrinkled from apex to around apical 3/4.
Measurements. BL, 4.4–4.8; FBL, 1.7–1.8; HL, 0.64–0.66; HW, 0.82–0.85; EL, 0.294–
0.303; AL, 1.58–1.61; PL, 0.79–0.80; PW, 1.10–1.14; ELL, 0.95–0.99; ELW, 1.31–1.38; HTL,
0.81–0.83.
Bionomics. The type series from Uzbekistan (three specimens) were collected with Crematogaster sp. (Dvořák, 1984), but nothing was reported on its association with ants was not referred to any more. The type series of an allied species, Pella esau, was also collected with antsof the same genus. This suggests closer associations with the ants rather than merely walking

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M. Maruyama

Fig. 100. Pella bohaci (Dvořák). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

around ant colony. Thus, I tentatively regard this ant species as a symbiotic host of Pella bohaci.
The symbiotic host species is different from that of Pella esau which was also collected from
Tadjikistan. A paratype colleted from Turkmenia lacks information on the collecting situation.
No information was available for the life cycle of this species.
Symbiotic host. Crematogaster (Crematogaster) sp.
Pella cinctipennis (Eppelsheim, 1884), n. comb.
(Figs. 101–103)
Myrmedonia cinctipennis Eppelsheim, 1884a: 12 (original description).—Smetana, 2004: 460 (Zyras (Myrmelia), list).
Zyras (Pella) esau Dvořák, 1984: 197 (original description). —Smetana, 2004: 461 (Zyras (Pella), list). N. syn.

Type material. Myrmedonia cinctipennis. Holotype: Azerbaijan: /, “35/Rusano/cinctipennis/c. Eppelsh. Steind. d./cinctipennis Epp. Verh. Naturf. Ver. Brünn Bd. XXII/TYPUS”(NHMW).
Zyras (Pella) esau. Paratypes: Tadjikistan: 1?, “USSR, Tadjikistan Romit (Dušanbe) 25.–9.
4. 1981. Aldo Olexa lgt./PARATYPUS/Zyras (Pella) esau sp. n. det. M. Dvořák ,81” (cDvor).
Uzbekistan: 1/, “USSR, Uzbekistan Aktaš (Taškent) 22. 4. 1980 Aldo Olexa lgt./ALLOTY-

Palearctic Species of Pella

143

Fig. 101. Pella cinctipennis (Eppelsheim). —A, Fore body, dorsal view; B, right antenna.

PUS/Zyras (Pella) esau sp. n. det. M. Dvořák, 81” (cDvor).
These are part of the type series. Most specimens of the type series including the holotype
are deposited in cDvor. The male paratype of Zyras esau was pinned with an ant specimen.
Other material. Turkey: 1/, pass above Seki (1820 m alt.), 70 km NE Fethiye, Mugla, 7
VII 2002, V. Assing (cAss).
Type locality. Rusano, Azerbaijan.
Distribution. Mediterranean subregion. EUROPE: Turkey; ASIA: Azerbaijan, Tadjikistan,
Uzbekistan. New to Turkey.
Diagnosis. Pella cinctipennis is closely similar to P. kuluensis and P. bohaci but may beeasily distinguished from the latter two by smaller body.
Redescription. Body slender. Brown in ground colour; antennae, mouthparts, pronotum,
elytra and legs reddish brown. Head (Fig. 101 A) widest around middle of eyes; surface smooth,sparsely covered with setae; setae long, slightly longer than those on pronotum and elytra; lengthof eyes 0.36–0.37 times as long as head width. Antennae (Fig. 101 B) shorter than head, pronotum and elytra combined; 1st segment slightly longer than 2nd and 3rd combined; 2nd segment
slightly shorter than 3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments wider thanlong; 11th segment shorter than 1st; approximate relative lengths of segments from basal to apical: 23 : 10 : 11 : 6 : 5 : 5 : 4.5 : 4.5 : 6 : 5.5 : 17. Pronotum (Fig. 101 A) 1.28–1.36 times as wide as
long, widest around anterior 1/6, narrowed posteriorly; posterior margin rounded; surface
smooth, sparsely covered with setae, with 3 or 4 long macrosetae along lateral margin. Scutellum
with surface smooth, moderately covered with setae. Elytra (Fig. 101 A) slightly widened apicad;
surface smooth, moderately covered with setae, with 3 or 4 small macrosetae laterally. Legs

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M. Maruyama

Fig. 102. Pella cinctipennis (Eppelsheim). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus (evaginated), lateral view; D, ditto, ventral view; E, copulatory
piece, dorsal view. Male.

short; hind tibia 0.85–0.86 times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 5 : 4.5 : 4 : 10 in fore tarsus; 6 : 6 : 6 : 5 : 10 in mid tarsus; 10 : 7.5 : 7 :6 : 11.5 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around
4th and 5th segments; surface smooth; 3rd to 7th tergites sparsely covered with setae; 8th tergite(Figs. 102 A, 103 A) with 7 (sometimes 8) macrosetae; 9th tergite with 3 macrosetae; 10th tergite
with posterior margin slightly emarginate, with 3 macrosetae.
Male: Eighth tergite (Fig. 102 A) with posterior margin somewhat truncate, its truncate apex
dentate; 8th sternite (Fig. 102 B) with posterior margin rounded, with 10–12 macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs. 102 C, 102 D) somewhat pear-shaped in
ventral view; apical lobe curved ventrad near apex, pointed at apex in lateral view, obtuse at apex
in ventral view; basal ridge weakly convex; copulatory piece of inner sac as in Fig. 102 E. Para-

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145

Fig. 103. Pella cinctipennis (Eppelsheim). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

mere with apical lobe of paramerite dilated apicad, rounded at apex, 1/4 as long as condylite.
Female: Eighth tergite (Fig. 103 A) with posterior margin somewhat truncate, its truncate
apex weakly crenate; 8th sternite (Fig. 103 B) with 10 or 11 macrosetae; sensory setae of 8th
sternite generalised, almost the same as the other setae in shape. Spermatheca (Fig. 103 C)
curved three times; basal part almost as long as apical part, bulbous at base, acutely curved
around middle, dilated apicad; apical part slightly curved around middle, slightly narrowed apicad, with outer projection, its inner wall densely wrinkled from apex to around apical 2/5.
Measurements. BL, 3.7–4.2; FBL, 1.4–1.7; HL, 0.54–0.58; HW, 0.66–0.70; EL, 0.236–
0.260; AL, 1.21–1.28; PL, 0.58–0.63; PW, 0.78–0.83; ELL, 0.74–0.81; ELW, 0.98–1.10; HTL,
0.63–0.70.
Bionomics. A paratype of Zyras esau from Tadjikistan was collected with Crematogastersp. (DVOŘÁK, 1984). Although the collector of the type series did not report the behaviour ofZyras esau with the Crematogaster ants, I tentatively regard this ant species as symbiotic host of
this species. No information was available for the life cycle of this species.
Symbiotic host. Crematogaster (Crematogaster) sp.
The lugens GroupSpecies included. Pella lugens, P. beijingorum, P. intermedia, P. masakoae.
Distribution. European subregion, Mediterranean subregion, Manchurian subregion.

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M. Maruyama

Diagnosis. Species of the lugens group may be characterised by a combination of the following character states: 1) eye length 0.36–0.42 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with posterior margin completely rounded, not forming corners postero-laterally; 4) pronotum 1.50–1.66 times as wide as long; 5) pronotum widest
around middle; 6) pronotal hypomeron invisible in lateral view; 7) elytra bicoloured, yellow withblack maculations around scutellum and postero-lateral corners; 8) inner and posterior margins
of elytra weakly margined; 9) male 8th sternite longer than in female; 10) posterior margin of 8th
tergite crenate; 11) posterior margin of male 8th sternite rounded; 12) lateral projection of
apodeme of male 8th tergite not evidently longer than that of female; 13) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior margin of the segment; 14) postero-medial margin of female 8th sternite without minute pubescence; 15) apical lobe of aedeagal
median lobe slightly curved ventrad in lateral view; 16) distal crests of aedeagus well developed,
projected, rounded on ventral margin in lateral view; 17) copulatory piece of aedeagal median
lobe rounded around base and more or less pointed at apex in dorsal view.
Comments. The lugens group is probably a monophyletic group in view of the close similarity in the general appearance, e.g., the combination of the tricoloured body and the muchtransverse pronotum, and in the aedeagal shape with short apical lobe, which is narrowed at base.
However, each of these states is homoplastic, and no autapomorphy could be found in thespecies-group. This species-group is very similar to the spreta group in colour but aedeagal
structure is quite different. No allied species-group could be inferred.
Symbiotic hosts. Lasius (Dendrolasius) spp.
Key to the Species of the lugens Group
1. Aedeagal median lobe small, shorter than head capsule. .................................................... 2.
— Aedeagal median lobe small, longer than head capsule. ..................................................... 3.
2. Eyes larger, 0.41–0.42 times as long as head width; posterior margin of male 8th tergite
truncate; apical lobe of aedeagal median lobe parallel-sided around base. Distribution: Europe, Balkan, Caucasia. .............................................................................................P. lugens.
— Eyes smaller, 0.36–0.37 times as long as head width; posterior margin of male 8th tergite
slightly emarginate; apical lobe of aedeagal median lobe gently narrowed apicad. Distribution: China (Beijing Shi), Japan, Russia (Far East) . ........................................P. beijingorum.
3. Body larger: BL: 5.2–6.3 mm (normally, more than 6.0 mm); FBL: 2.4–2.6 mm; apical lobeof aedeagal median lobe about 1/2 as long as basal capsule. Distribution: Japan, Korea. .......
..............................................................................................................................P. masakoae.
— Body smaller: BL: 5.3 mm; FBL: 2.3 mm; apical lobe of aedeagal median lobe about 3/5 aslong as basal capsule. Distribution: Russia (Far East). ......................................P. intermedia.
Pella lugens (Gravenhorst, 1802), n. comb.
(Figs. 104–106)
Aleochara lugens Gravenhorst, 1802 (original description): 95.—Stephens, 1832: 161 (Aleochara, description).—Erichson, 1837: 289 (Myrmedonia, description). — Mulsant & Rey, 1873a: 76 (Myrmedonia, description).—Ganglbauer,
1895: 123 (Myrmedonia (Pella), key, description). — Reitter, 1909: 43 (Myrmedonia (Myrmedonia), key).—Bernhauer & Scheerpeltz, 1926: 701 (Zyras (Myrmedonia), list).—Scheerpeltz, 1934: 1656 (Zyras (Myrmedonia),list).—Fenyes, 1920: 297 (Zyras (Myrmedonia), list). —Lohse, 1974: 226 (Zyras (Pella), key).—Gürlich, 1981: 211(Lepla, chaetotaxy). — Likovský, 1993: 59 (Zyras (Lepla), list).—Smetana, 2004: 460 (Zyras (Lepla), list).

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147

Fig. 104. Pella lugens (Gravenhorst). — A, Fore body, dorsal view; B, right antenna.

Type material. Lectotype: Germany: designated herewith, ?, [“Berolin” (Berlin, Germany)] (without label, pinned).
Paralectotype: 1/, [ditto] (without label, pinned).
These are deposited in ZMHB.
Comments on lectotype designation. The syntypes of Aleochara lugens contain two
species, Pella lugens and Pella laticollis (two of the four syntypes). Thus, the lectotype is herewith designated for a specimen, which is generally recognized as Pella lugens in many articles.
Other material. Armenia: 1 ex., Nerkin And, Kafan, Syunik, 24 VII 1952, Iablokoff-Khnzoryan; 1 ex., Shagali, Kirovakan, Lori, 2 VI 1949, Iablokoff-Khnzoryan. Austria: 1/, Wien,
Breit (no other data); 1?, Herrmanstadt, 7 VI 1887, Slkalitt; 1?, Paum Garbenm, 27 V (no otherdata); 1/, Neuwalolegg, 24 IV 1904, Luze (LDFL). Bosnia Herzegovina: 1/, Ivan Sedio,
Pazario, 21 III 1913, J. Fodor. Czech Republic: 2??, 2// , Nový Jičín, Moravia, 30 IV 1988,
Kuboň (cMar). Greece: 4 exs. (2 adults, 2 larvae), Fthiotis (alt. 940 m), Oros, Kallidromo,
Lamia, 2 IV 2001, V. Assing; 5 exs., Kastanea, Kalambaka, Pindos, 28 III 1997, A. Riedal. Slovakia: 1?, Teplý vrch, 23 IV 1998, T. Lackner.
Type locality. Berlin, Germany.
Distribution. European subregion, Mediterranean subregion. EUROPE: Armenia, Austria,
Belgium, Belarus, Croatia, Czech Republic, Denmark, Finland, France, Great Britain, Germany,
Georgia, Greece, Hungary, Italy, Latvia, Lithuania, Luxembourg, Macedonia, Netherlands, Norway, Poland, Romania, Russia (North and Central European Territories), Slovakia, Sweden,
Switzerland, Ukraine, Yugoslavia. ASIA: Kazakhstan, Turkmenistan, Uzbekistan. New to Armenia.

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Fig. 105. Pella lugens (Gravenhorst). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

Diagnosis. Pella lugens is closely similar to P. beijingorum, P. masakoae and P. intermedia
in general appearance but can be distinguished from P. beijingorum by the eyes 0.47–0.42 as longas the head width and the apical lobe of the aedeagal median lobe parallel-sided, from P.
masakoae and P. intermedia by the smaller body and the small aedeagus which is much shorter
than the head capsule. Pella lugens is locally isolated from the other species of the lugens group.
This species may be easily distinguished from the sympatric species by a combination of the
pronotal hypomeron invisible in lateral view and the elytra yellow in ground colour.
Redescription. Body slender. Black in ground colour; antennae, mouthparts, legs, andaround posterior margins of 3rd to 6th tergites reddish brown; elytra yellow but with black maculations around scutellum and postero-lateral corners. Head (Fig. 104 A) widest just behind eyes;surface finely reticulated, moderately covered with setae; setae moderately long, longer thanthose on pronotum and elytra; length of eyes 0.41–0.42 times as long as head width. Antennae

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149

Fig. 106. Pella lugens (Gravenhorst). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca. Female.

(Fig. 104 B) shorter than head, pronotum and elytra combined; 1st segment as long as 2nd and3rd combined; 2nd segment about 2/3 as long as 3rd; 3rd segment about 3/5 as long as 1st; 4th to10th segments wider than long; 11th segment shorter than 1st; approximate relative lengths ofsegments from basal to apical: 25 : 10 : 15 : 6.5 : 6 : 5.5 : 5.5 : 6 : 6.5 : 7.5 : 22.5. Pronotum (Fig.
104 A) elliptical, 1.50–1.66 times as wide as long, widest just after middle; surface finely punctured, finely reticulated among punctures, densely covered with setae, with 7 or 8 macrosetae
along lateral margin. Scutellum with surface smooth, moderately covered with setae. Elytra (Fig.
104 A) widened apicad; surface finely punctured, finely reticulated among punctures, densely
covered with setae, with 4 or 5 small macrosetae laterally. Legs short; hind tibia 0.82–0.89 times
as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 6 : 6 :11.5 in fore tarsus; 11 : 8 : 7 : 6 : 11 in mid tarsus; 17.5 : 10.5 : 9.5 : 7.5 : 12.5 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 4th and 5th segments; surface smooth; 3rd to 7th tergites sparsely covered with minute setae, with a low of setae alongposterior margins; 8th tergite (Figs. 105 A, 106 A) with 7 or 8 macrosetae; 9th tergite with 5
macrosetae; 10th tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 105 A) with posterior margin somewhat truncate, its truncate apex
crenate; 8th sternite (Fig. 105 B) with posterior margin rounded, with 17–21 macrosetae; 9th sternite with posterior margin rounded. Aedeagus (Figs. 105 C, 105 D) somewhat pear-shaped in
ventral view; apical lobe gently curved ventrad in lateral view, pointed at apex in lateral and ventral views; basal ridge roundly convex; copulatory piece of inner sac as in Fig. 105 E. Paramerewith apical lobe of paramerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 106 A) with posterior margin slightly emarginate, its emarginate

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apex weakly crenate; 8th sternite (Fig. 106 B) with 13 or 14 macrosetae; sensory setae of 8th
sternite generalised, almost the same as the other setae in shape. Spermatheca (Fig. 106 C)
curved three times; basal part about 2.5 times as long as apical part, acutely curved around basal
2/5; apical part gently curved around middle, narrowed apicad, its inner wall densely wrinkledfrom apex to around apical 2/3.
Measurements. BL, 4.3–5.2; FBL, 2.1–2.5; HL, 0.61–0.69; HW, 0.78–0.86; EL, 0.331–
0.358; AL, 1.23–1.75; PL, 0.53–0.79; PW, 0.85–1.30; ELL, 0.78–1.10; ELW, 103–1.56; HTL,
0.88–0.96.
Bionomics. In Europe, this species is rather commonly collected from trails Lasius (Dendrolasius) fuliginosus and rarely from around nests of L. (L.) brunneus. Life cycle of this species
is probably similar to that of Pella masakoae.
Symbiotic host. Lasius (Lasius) brunneus, L. (Dendrolasius) fuliginosus.
Pella beijingorum (Pace, 1998), n. comb.
(Figs. 107–109)
Zyras (Pella) beijingorum Pace, 1998a: 976 (original description).—Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Holotype: China (Beijing Shi): /, “CHINA Beijing [Xiaolongmen, Beijing Shi, China (accorinding to the original description)] 1100–1500 m 1. VII. 1993 G. de Rougemont/Zyras beijingensis sp. n. det. R. Pace 1995/HOLOTYPUS Zyras beijingensis m. det. R.
Pace 1995” (MHNG).
Other material. China (Beijing Shi): 1?, 2// , Xiaolongmen (1400 m), Dongling, Yanshan, Beijing, 15–16 VI 2001, P. Hlaváč & J. Cooter (LDFJ) (IZAS, cMar, cHlav). Japan:
Hokkaidô: 4 exs., Hebi-numa, Teshio-gawa, Teshio-chô, 9 VII 1992, S. Hori (pt); 1?, 1/, 18
exs., Nozaki, Bihoro-chô, Hokkaidô, 23 VI 2001, Y. Yamaga (LDC); 2 exs., Taihei, Maruseppuchô, Hokkaidô, 29–31 V 2000, Y. Kida (LDFJ); 1 ex., same data but, 16–17 VI 2000; 10 exs.,
Kamimuri, Maruseppu-chô, Hokkaidô, 6 VI 2004, M. Satô (LCHU); 1 ex., Akita, Oketo-chô, 7V 1992, T. Kato; 1 ex., Mitsumata, Kamishihoro-chô, 22–25 V 1990, K. Haga; 2 exs., same databut, 21 VII 1995, K. Miyashita; 1 ex., Takami, Shizunai-chô, 23 VIII 1993, S. Hori; 2 exs., Shiomi, Mukawa-chô, 17 IX 2001, S. Hori; 1?, 1/, Shibumbestu, Nopporo-shinrin-kôen, Ebetsu-shi,
11 V 2000, M. Maruyama (LDFJ); 2 exs., Tomambetsu, Nopporo-shinrin-kôen, Ebetsu-shi, 11 V
2000, M. Maruyama (LDS ); 4 exs., Hakken-zan, Sapporo-shi, 31 V 2002, M. Maruyama.
Honshû: 1/, Uono-gawa, Kawaguchi-machi, Niigata-ken, 30 IV 1995, K. Haga; 1 ex., Nakano,Shôwa-machi, Kitakatsushika-gun, Saitama-ken, 14 IV 2002, H. Kamezawa (LDFJ); 6 exs.,
same data but, 25 V 2003; 1 ex., Isezaki, Môka-shi, Tochigi-ken, 14 VI 2004, M. Maruyama(LDFJ). Russia (Far East): 1/, Arsenev, Primorskyi krai, 27 V–3 VII 1991 O. Šausa (NHMW);
1/, “Prwnaja-Rjetschkam Tales, Vladivostok, 1918–1920, H. Farieb” (NHMW); 1 ex.,
Kaimanovka, Ussuryisk, Primorskyi Krai, 27 V 2005, M. Maruyama (LL); 1 ex., Kamenushka,
Ussuryisk, Primorskyi Krai, 28 V 2005, M. Maruyama (LDFJ); 1 ex., Bukhta Vityaz, Poluostrov
Gamov, Khasanskyi, Primorskyi Krai, 31 V 2005, M. Maruyama (LDFJ); 4 exs., “Seitengraben
des Prwnaja-Rjetschka Tales b. Wladiwostok/H. Farieb leg. 1918–1920” (NHMW).
Distribution. Manchurian subregion. ASIA: China (Beijing Shi), Japan (Hokkaidô,
Honshû), Russia (Far East). New to Japan and Russia.
Diagnosis. Pella beijingorum is closely similar to P. lugens, P. masakoae and P. intermedia
in general appearance but can be distinguished from P. lugens by the eyes 0 36–0.37 as long as
the head width and the apical lobe of the aedeagal median lobe gently narrowed posteriad, from

Palearctic Species of Pella

151

Fig. 107. Pella beijingorum (Pace). — A, Fore body, dorsal view; B, right antenna.

P. masakoae and P. intermedia by the smaller body and the tiny aedeagus which much shorter
than the head capsule. This species is similar in colouration and general appearance to Pella spreta, but distinguished from it by the smaller eyes and the wider pronotum.
Redescription. Body slender. Black in ground colour; antennae, mouthparts, legs, andaround posterior margins of 3rd to 6th tergites reddish brown; elytra yellow but with black maculations around scutellum and postero-lateral corners. Head (Fig. 107 A) widest just behind eyes;surface finely reticulated, moderately covered with setae; setae moderate long, longer than thoseon pronotum and elytra; length of eyes 0.36–0.37 times as long as head width. Antennae (Fig.
107 B) shorter than head, pronotum and elytra combined; 1st segment as long as 2nd and 3rd
combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 as long as 1st; 4th to10th segments wider than long; 11th segment longer than 1st; approximate relative lengths ofsegments from basal to apical: 21 : 10 : 11 : 5.5 : 5 : 4.5 : 4.5 : 4.5 : 5 : 6 : 23. Pronotum (Fig. 107A)
elliptical, 1.57–1.64 times as wide as long, widest just after middle; surface finely punctured,finely reticulated among punctures, densely covered with setae, with 7 or 8 macrosetae along lateral margin. Scutellum with surface smooth, moderately covered with setae. Elytra (Fig. 107 A)
widened apicad; surface finely punctured, finely reticulated among punctures, densely covered
with setae, with 4 or 5 small macrosetae laterally. Legs short; hind tibia 0.81–0.88 times as longas elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 7 : 5.5 : 12 infore tarsus; 10 : 8 : 7.5 : 6 : 9.5 in mid tarsus; 15 : 9 : 8 : 7 : 12 in hind tarsus. Abdomen subparallelsided, slightly narrower than elytra, widest around 4th and 5th segments; surface smooth; 3rd to
7th tergites sparsely covered with minute setae, with a low of setae along posterior margins; 8th

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M. Maruyama

Fig. 108. Pella beijingorum (Pace). —A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

tergite (Figs. 108 A, 109 A) with 7 or 8 macrosetae; 9th tergite with 5 macrosetae; 10th tergite
with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 108 A) with posterior margin somewhat emarginate, its emarginate
apex crenate; 8th sternite (Fig. 108 B) with posterior margin rounded, with 21–23 macrosetae;
9th sternite with posterior margin rounded. Aedeagus (Figs. 108 C, 108 D) somewhat pearshaped in ventral view; apical lobe gently curved ventrad in lateral view, pointed at apex in lateral and ventral views; basal ridge roundly convex; copulatory piece of inner sac as in Fig. 108 E.
Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/3 as long as
condylite.
Female: Eighth tergite (Fig. 109 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 109 B) with 13 or 14 macrosetae; sensory setae of 8th
sternite generalised, almost the same as the other setae in shape. Spermatheca (Figs. 109 C–
109 F) curved three times; basal part about 2.5 times as long as apical part, acutely curved

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153

Fig. 109. Pella beijingorum (Pace). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C–F, spermathecae (F, teneral). Female.

around basal 2/5; apical part gently curved around middle, narrowed apicad, its inner wall densely wrinkled from apex to around apical 2/3, rarely with outer projection.
Measurements. BL, 4.4–5.1; FBL, 2.2–2.4; HL, 0.58–0.64; HW, 0.72–0.83; EL, 0.264–
0.314; AL, 1.31–1.61; PL, 0.68–0.75; PW, 1.11–1.23; ELL, 0.93–1.08; ELW, 1.28–1.45; HTL,
0.78–0.88.
Bionomics. In Japan, this species is relatively rare and has been collected from colonies of
Lasius (Dendrolasius) fuji, L. (D.) spathepus and L. (D.) capitatus. Some individuals were collected from a nest of Lasius (Chthonolasius) sp. in Hokkaidô together with Pella masakoae, andone individual was collected from a nest of L. (L.) cf. platythorax Seifert, 1991, in Primorskyi
Krai, on one occasion, respectively. Life cycle of this species is probably similar to that of Pella
masakoae.
Symbiotic hosts. Lasius (Lasius) sp., L. (Chthonolasius) umbratus., L. (Dendrolasius) fuji,

154

M. Maruyama

L. (D.) spathepus, L. (D.) capitatus.
Pella intermedia Maruyama, n. sp.
(Figs. 110, 111)

Type material. Holotype: RUSSIA (Far East): ?, “RUSSIA: Primorskyi Krai, Nadezhdunskyi, Kravtzovka, 1 VI 2005, Maruyama M.” (LDFJ) (IBSS).
Type locality. Nadezhdunskyi, Primorskyi Krai, Russia.
Distribution. Manchurian subregion. ASIA: Russia (Far East).
Etymology. Named for the intermediate body size between the eastern Palearctic species ofthe lugens group, Pella beijingorum and P. masakoae.
Diagnosis. Pella intermedia is closely similar to P. masakoae in general appearance andlarge aedeagus, which is much larger than head capsule, but can be distinguished from the latterby the smaller body and the longer apical lobe of aedeagal median lobe. This species is also similar to Pella lugens and P. beijingorum in general appearance but can be distinguished from thelatter by the much larger body and large aedeagus. This species can be easily distinguished fromthe sympatric species by a combination of the following character states: the body large, the elytra yellow in ground colour, and the legs reddish brown.
Description. Body slender. Dark brown in ground colour; antennae, mouthparts, legs, andaround posterior margins of 3rd to 5th tergites reddish brown; elytra with around antero-lateralcorners yellowish brown. Head (Fig. 110 A) widest around middle of eyes; surface finely reticulated, moderately covered with setae; setae moderately long, longer than those on pronotum andFig. 110. Pella intermedia Maruyama, n. sp.—A, Fore body, dorsal view; B, right antenna.

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155

Fig. 111. Pella intermedia Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal
view. Male.

elytra; length of eyes 0.37 times as long as head width. Antennae (Fig. 110 B) shorter than head,pronotum and elytra combined; 1st segment longer than 2nd and 3rd combined; 2nd segment
about 4/5 as long as 3rd; 3rd segment shorter than 1/2 of 1st; 4th to 10th segments wider thanlong; 11th almost as long as 1st; approximate relative lengths of segments from basal to apical:
24 : 9 : 11 : 6 : 5.5 : 5.5 : 6 : 6 : 6 : 7.5 : 23. Pronotum (Fig. 110 A) elliptical, 1.62 times as wide as
long, widest just after middle; surface finely punctured, finely reticulated among punctures,
densely covered with setae, with 7 or 8 macrosetae along lateral margin. Scutellum with surface
smooth, moderately covered with setae. Elytra (Fig. 110 A) widened apicad; surface finely punctured, finely reticulated among punctures, densely covered with setae, with 6 or 7 small macrose-

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M. Maruyama

tae laterally. Legs short; hind tibia 0.84 times as long as elytra; approximate relative lengths oftarsal segments from basal to apical: 5 : 5.5 : 5.5 : 11 in fore tarsus; 10 : 7 : 6 : 5.5 : 12 in mid tarsus; 17 : 10 : 8.5 : 7.5 : 12 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra,
widest around 4th and 5th segments; surface smooth; 3rd to 7th tergites almost glabrous except
for a row of setae along posterior margins; 9th tergite with 3 macrosetae; 10th tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 111 A) with posterior margin somewhat emarginate, its emarginate
apex crenate, with 8 or 9 macrosetae; 8th sternite (Fig. 111 B) with posterior margin rounded,with 23–25 macrosetae; 9th sternite with posterior margin rounded. Aedeagus (Figs. 111 C,
111 D) somewhat pear-shaped in ventral view; apical lobe almost straight in lateral view, pointed
at apex in lateral and ventral views; basal ridge roundly convex; copulatory piece of inner sac as
in Fig. 111 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/3
as long as condylite.
Female unknown.
Measurements. BL, 5.3; FBL, 2.3; HL, 0.68; HW, 0.84; EL, 0.308; AL, 1.51; PL, 0.77;
PW, 1.26; ELL, 1.01; ELW, 1.4; HTL, 0.85.
Bionomics. Only the male holotype has been known. The type locality is an old secondary
forest of Abies and Quercus trees, and the specimen was collected from the trials of Lasius (Dendrolasius) fuji together with Pella jureceki and Homoeusa spp.
Symbiotic host. Lasius (Dendrolasius) fuji. Probably most sympatric species of Dendrolasius.
Pella masakoae Maruyama, n. sp.
(Figs. 112–114)

Type material. Holotype: Japan: ?, “Takao-san, Hachiôji-shi, Tokyo, JPN, 4 VI 2001,
Maruyama-M. leg./ex colony of Lasius spathepus, det. Maruyama, 2002” (LDS) (NSMT).
Paratypes: Japan: Hokkaidô: 1 ex., Himenuma, Rishiri-tô, 4 IX 1990, T. Kishimoto (pt); 1
ex., Makoi, Shari-chô, 28 V 2000, Y. Kida (LDFJ); 1 ex., same data but, 20 VI 2000 (LCH); 1
ex., same data but, 27–29 V 2001 (LDFJ); 3 exs., same data but, 24 VI 2002; 2 exs., same databut, 27 VI 2002; 2 exs., Miwa, Koshimizu-chô, 28 VII 2001, S. Kawahara (LDC); 1 ex., Taihei,
Maruseppu-chô, 19–21 VIII 2000, Y. Kida (LDFJ); 8 exs., Kamimuri, Maruseppu-chô,
Hokkaidô, 6 VI 2004, M. Satô (LCHU); 2 exs., Kamiotoineppu, Otoineppu-mura, 15–23 VII
1997, S. Hori (pt); 1 ex., Inasato, Hobetsu-chô, 9 V 2004, S. Hori (tt); 32 exs., Tomambetsu,
Nopporo-shinrin-kôen, Ebetsu-shi, 11 V 2000, M. Maruyama (LDS); 21 exs., same data but, 18
V 2000; 1 ex., same data but, 18 V 2000 (LDFJ); 2 exs., Kaitaku-kinenkan, Nopporo-shinrinkôen, Ebetsu-shi, 1–10 V 2000, S. Hori (fit); 1 ex., same data but, 17–24 V 2000; 1 ex., samedata but, 24–31 V 2000; 2 exs., same data but, 11–19 VII 2000; 2 exs., same data but, 19–26 VII
2000; 1 ex., same data but, 26 VII–2 VIII 2000; 1 ex., same data but, 31 V 2000 (tt); 1 ex., samedata but, 20 IV 2003 (LDFJ); 5 exs., Ôsawaguchi, Nopporo-shinrin-kôen, Ebetsu-shi, 4 V 2000,
M. Maruyama (LDFJ); 1 ex., same data but, 9 V 2000 (LDFJ); 2 exs., Hitsujigaoka, Sapporoshi, 25 VII 1998, M. Maruyama (LDS); 1 ex., same data but, 21 IX 1998; 1 ex., Hakkenzan, Sapporo-shi, 12–14 VII 2001, H. Sugaya (fit); 7 exs., same data but, 31 V 2002, M. Maruyama(LDFJ); 28 exs., same data but, 1 VI 2002; 2 exs., Kannon-zawa, Sapporo-shi, I VI 2002, M.
Maruyama (LDFJ). Honshû: 14 exs., Takao-san (alt. 450 m), Hachiôji-shi, Tokyo-to, 4 VI 2001,
M. Maruyama (LDS); 1 ex., Tanzawa, Kanagawa-ken, 14 VI 1986, M. Tao; 1 ex., Gozaishi-

Palearctic Species of Pella

157

Fig. 112. Pella masakoae Maruyama, n. sp. —A, Fore body, dorsal view; B, right antenna.

kôsen, Nirasaki-shi, Yamanashi-ken, 18 V 1990, K. Hosoda; 1 ex., Kamiimai, Hosaka-chô, Nirasaki-shi, 6 VI 2004, M. Maruyama (LDFJ); 1 ex., Fukazawa-jûnimagari, Nakamaru, Nagasaka-chô, Kitakoma-gun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ). Shikoku: 1 ex.,Ôtaki-san, Shionoe-chô, Kagawa-ken, 2 VI 2001, M. Maruyama & Y. Kamite (LDS). Korea: 1?,
Chŏndong, Sobaek-san, Tangyang-gun, Kyŏngbuk Prov., 7–9 V 1999, U.-S. Hwang & H.-J. Kim(fit) (CNU).
Type locality. Takao-san, Hachiôji-shi, Tokyo-to, Japan.
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô, Honshû, Shikoku), Korea.
Etymology. Dedicated to my grandmother Masako Maruyama, who passed away in Tokyoto, when I was finishing my Ph. D. thesis.
Diagnosis. Pella masakoae is closely similar to P. intermedia in general appearance andlarge aedeagus, which is much larger than head capsule, but can be distinguished from the latterby the larger body and the shorter apical lobe of aedeagal median lobe. This species is also similar to Pella lugens and P. beijingorum in general appearance but can be distinguished from thelatter by the much larger body and huge aedeagus. This species can be easily distinguished fromthe sympatric species by a combination of the following character states: the body large, the elytra yellow in ground colour, and the legs reddish brown.
Description. Body slender. Black in ground colour; antennae, mouthparts, legs, and around
posterior margins of 3rd to 5th tergites reddish brown; elytra yellow but with black maculations
around scutellum and postero-lateral corners. Head (Fig. 112 A) widest around middle of eyes;surface finely reticulated, moderately covered with setae; setae moderately long, longer than158

M. Maruyama

Fig. 113. Pella masakoae Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal
view. Male.

those on pronotum and elytra; length of eyes 0.39–0.40 times as long as head width. Antennae(Fig. 112 B) shorter than head, pronotum and elytra combined; 1st segment longer than 2nd and3rd combined; 2nd segment about 2/3 as long as 3rd; 3rd segment longer than 1/2 of 1st; 4th to10th segments wider than long; 11th segment shorter than 1st; approximate relative lengths ofsegments from basal to apical: 31 : 11 : 17.5 : 8.5 : 7 : 6 : 6.5 : 7 : 7.5 : 9 : 27. Pronotum (Fig. 112 A)
elliptical, 1.51–1.60 times as wide as long, widest just after middle; surface finely punctured,finely reticulated among punctures, densely covered with setae, with 7 or 8 macrosetae along lateral margin. Scutellum with surface smooth, moderately covered with setae. Elytra (Fig. 112 A)
widened apicad; surface finely punctured, finely reticulated among punctures, densely covered
with setae, with 4 or 5 small macrosetae laterally. Legs short; hind tibia 0.88–0.91 times as long

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Fig. 114. Pella masakoae Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca. Female.

as elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 6 : 6 : 11.5 infore tarsus; 11 : 8 : 7 : 6 : 11.5 in mid tarsus; 20 : 12 : 10 : 8.5 : 11.5 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 4th and 5th segments; surface smooth;
3rd to 7th tergites almost glabrous except for a row of setae along posterior margins; 9th tergite
with 5 macrosetae; 10th tergite with posterior margin slightly emarginate, with 4 macrosetae.
Male: Eighth tergite (Fig. 113 A) with posterior margin somewhat emarginate, its emarginate
apex crenate, with 15 macrosetae; 8th sternite (Fig. 113 B) with posterior margin rounded, with
27–31 macrosetae; 9th sternite with posterior margin rounded. Aedeagus (Figs. 113 C, 113 D)
somewhat pear-shaped in ventral view; apical lobe slightly curved ventrad in lateral view, pointed
at apex in lateral and ventral views; basal ridge roundly convex; copulatory piece of inner sac as
in Fig. 113 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/5
as long as condylite.
Female: Eighth tergite (Fig. 114 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate, with 10 macrosetae; 8th sternite (Fig. 114 B) with 16 or 17 macrosetae; sensory setae of 8th sternite generalised, almost the same as the other setae in shape. Spermatheca(Fig. 114 C) curved twice; basal part about twice as long as apical part, roundly curved aroundmiddle; apical part narrowed apicad, its inner wall sparsely wrinkled from apex to around apical
2/5.
Measurements. BL, 5.2–6.3; FBL, 2.4–2.6; HL, 0.71–0.75; HW, 0.86–0.93; EL, 0.342–
0.367; AL, 1.70–1.89; PL, 0.81–0.93; PW, 1.28–1.47; ELL, 1.15–1.26; ELW, 1.63–1.80; HTL,
1.01–1.15.
Bionomics. This species is relatively rarely collected from nests of Lasius (Dendrolasius)
spp., but sometimes many individuals are found in one nest. Several specimens were collected

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also from nests of Lasius (Chthonolasius) umbratus in Hokkaidô together with Pellabeijingorum.
Symbiotic hosts. Lasius (Dendrolasius) fuliginosus, L. (D.) spathepus, L. (Chthonolasius)
umbratus.
The spreta GroupSpecies included. Pella spreta, P. zhoui.
Distribution. Manchurian subregion.
Diagnosis. Species of the spreta group may be characterised by a combination of the following character states: 1) eye length 0.45–0.48 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with posterior margin completely rounded, not forming corners postero-laterally; 4) pronotum 1.51–1.56 times as wide as long; 5) pronotum widest
around middle; 6) pronotal hypomeron invisible in lateral view; 7) elytra with yellow maculations
around antero-lateral and postero-lateral corners; 8) inner and posterior margins of elytra weakly
margined; 9) male 8th sternite longer than in female; 10) posterior margin of 8th tergite crenate;11) posterior margin of male 8th sternite rounded; 12) lateral projection of apodeme of male 8th
tergite evidently longer than that of female; 13) macrosetae of 8th abdominal segment generalised in length, exceeding the posterior margin of the segment; 14) postero-medial margin of female 8th sternite without minute pubescence; 15) apical lobe of aedeagal median lobe almost
straight in lateral view; 16) distal crests of aedeagus well developed, projected, rounded on ventral margin in lateral view; 17) copulatory piece of aedeagal median lobe gently narrowed apicad
and more or less rounded at apex in dorsal view.
Comments. The spreta group is probably a monophyletic group in view of the close similarity in the general appearance, e.g., a combination of the tricoloured body and the completely
elliptical pronotum, and in the aedeagal shape with straight apical lobe. However, each of thesestates is homoplastic, and no autapomorphy could be found in the species-group. This speciesgroup is very similar to the lugens group in colour but aedeagal structure is quite different. No
allied species-group could be inferred.
Symbiotic hosts. Lasius (Dendrolasius) spp.
Key to the Species of the spreta Group——

Pronotum 1.51–1.55 times as wide as long; posterior margin of male 8th tergite almost
truncate and only slightly emarginate medially, and its lateral corner not projected; apicallobe of aedeagal median lobe about 4/5 as long as basal capsule. Distribution: Japan. ..........
....................................................................................................................................P. spreta.
Pronotum 1.55–1.56 times as wide as long; posterior margin of male 8th tergite almost
widely emarginate, and its lateral corner somewhat projected; apical lobe of aedeagal median lobe about 1/2 as long as basal capsule. Distribution: China (Beijing Shi), Russia (Far
East). ...........................................................................................................................P. zhoui.
Pella spreta (Sharp, 1888), n. comb.
(Figs. 115–117)

Myrmedonia spreta Sharp, 1888: 291 (original description).—Bernhauer & Scheerpeltz, 1926: 705 (Zyras (Myrmedonia), list).—Dvořák, 1981: 54 (Zyras (Pella), key). — Smetana, 2004: 461 (Zyras (Pella), list).

Palearctic Species of Pella

161

Type material. Lectotype: Japan: designated herewith, ?, “Myrmedonia spreta Type D.S.
Sapporo. Japan. Lewis (written on paper card specimen glued on)/SYN-TYPE (blue round curator label)/Type (red round curator label)/Sapporo 5. VIII.–16. VIII. 80./Japan. G. Lewis./Sharp
Coll. 1905-313.” (BMNH).
Paralectotype: Japan: 1?, “Myrmedonia spreta D.S., Sapporo. Japan. Lewis (written on
paper card specimen glued on)/SYN-TYPE (blue round curator label)/Type (red round curator
label)/Sapporo 5. VIII.–16. VIII. 80./Japan. G. Lewis., 1910-320” (BMNH).
Type locality. Sapporo-shi, Hokkaidô, Japan.
Comments on lectotype designation. In the present study, I was unable to examine all of
the type series. According to the original description, the remaining syntypes include material
from “Hakodate” (Hakodate-shi, Hokkaidô) other than from Sapporo, though the number of
specimens was not noted. However, any specimen labelled “Hakodate” was not found in BMNH
(M. Brendell, pers. comm.) and the other museums I have contacted. On the other hand, in
Hokkaidô, Pella spreta is often collected together with P. beijingorum and P. masakoae that arevery similar to P. spreta in general appearance and usually difficult to distinguish from one another without dissection of genitalia. Sharp (1888) did not dissect the specimens. Therefore, there isa possibility that unexamined portion of the type series include specimen of Pella beijingorum or
P. masakoae. For this reason I designate herewith the lectotype of “Myrmedonia spreta” for thestability of the scientific name.
Other material. Japan: Hokkaidô: 3 exs., Makoi, Shari-chô, 27–29 V 2001, Y. Kida
(LDFJ); 2 exs., Asahi, Koshimizu-chô, 10 XII 1992, S. Kawahara; 4 exs., Nosaki, Bihoro-chô,
23 VI 2001, Y. Yamaga (LDC ); 2 exs., Taihei, Maruseppu-chô, 16–17 VI 2000, Y. Kida (LDFJ);

Fig. 115. Pella spreta (Sharp). — A, Fore body, dorsal view; B, right antenna.

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M. Maruyama

Fig. 116. Pella spreta (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

1 ex., same data but, 19–21 VIII 2000; 16 exs., Ôsawaguchi, Nopporo-shinrin-kôen, Ebetsu-shi,
4 V 2000, M. Maruyama (LDFJ); 1 ex., same data but, 16–19 VI 2001, S. Hori; 18 exs., Tomambetsu, Nopporo-shinrin-kôen, Ebetsu-shi, 11 V 2000, M. Maruyama (LDS); 12 exs., same databut, (LDFJ); 12 exs., same data but, 18 V 2000 (LDS); 20 exs., Hitsujigaoka, Sapporo-shi, 21 IX
1998, M. Maruyama (LDS ); 2 exs., same data but, 18 V 2000 (LDS); 21 exs., Hakken-zan, Sapporo-shi, 31 V 2002, M. Maruyama (LDFJ); 42 exs., same data but, 1 VI 2002; 11 exs., Chitosegawa, Rankoshi, Chitose-shi, 18 VII 2000, T. Itô & A. Ohkawa (light trap); 1 ex., Horokappugawa, Raikishi, Shakotan-chô, 7 VII 1996, M. Ôhara (LDFJ); 1 ex., Akagawa, Hakodate-shi, 15
VIII 1992, M. Nishikawa (LDFJ). Honshû: 17 exs., Mizuhiki, Tateiwa-mura, Minamiaizu-gun,
Fukushima-ken, 12 VI 2004, H. Kamezawa (LDFJ); 1 ex., Kashi-onsen Nishigô-mura, Fukushi-

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163

Fig. 117. Pella spreta (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

ma-ken, 16 VI 1998, M. Maruyama (LDFJ); 1 ex., Shimokomoriya, Môka-shi, Tochigi-ken, 6
VII 1999, M. Maruyama (LDFJ); 1 ex., Daiyûzan, Hakone-machi, Kanagawa-ken, 27 VIII 1982,
Y. Shibata (LDFJ) (SCM); 1 ex., Fukazawa-jûnimagari, Nakamaru, Nagasaka-chô, Kitakomagun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 1 ex., Misuzu, Ina-shi, Nagano-ken, 18
IX 2000, T. Kobayashi (LDFJ); 1 ex., Sasabe, Kawanishi-shi, Hyôgo-ken, 16 VIII 2001, N. Inahata.
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô, Honshû).
Diagnosis. Pella spreta is very similar to P. zhoui in general appearance, but is distinguished from the latter by the apical lobe of aedeagal median lobe as long as basal capsule andmuch longer than that of P. zhoui and by the slightly wider pronotum. These two species are locally isolated. Among the sympatric species, Pella spreta is similar in colouration and generalappearance to P. beijingorum and P. masakoae, but is distinguished from P. beijingorum by thelarger eyes and the narrower pronotum and from P. masakoae by the smaller body.
Redescription. Body slender. Black in ground colour; antennae, mouthparts, legs, andaround posterior margins of 3rd to 6th tergites reddish brown; elytra yellow but with black maculations around scutellum and postero-lateral corners. Head (Fig. 115 A) widest around middle ofeyes; surface finely reticulated, moderately covered with setae; setae moderate in length, as longas those on pronotum and elytra; length of eyes 0.46–0.48 times as long as head width. Antennae(Fig. 115 B) shorter than head, pronotum and elytra combined; 1st segment almost as long as 2nd
and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 3/5 as long as 1st;
4th to 10th segments wider than long; 11th segment shorter than 1st; approximate relative lengthsof segments from basal to apical: 25 : 11 : 14.5 : 7 : 7 : 6.5 : 6.5 : 6.5 : 6.5 : 7.5 : 23. Pronotum (Fig.
115 A) elliptical, 1.51–1.55 times as wide as long, widest around middle; posterior margin rounded; surface roughly punctured, finely reticulated between punctures, moderately covered withsetae, with 6 or 7 macrosetae along lateral margin. Scutellum with surface smooth, sparsely covered with setae. Elytra (Fig. 115 A) slightly widened apicad; surface smooth, moderately covered

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M. Maruyama

with setae, with 5 or 6 small macrosetae laterally. Legs short; hind tibia 0.85–0.90 times as longas elytra; approximate relative lengths of tarsal segments from basal to apical: 5 : 5 : 5.5 : 11.5 infore tarsus; 10.5 : 8 : 7 : 6 : 9.5 in mid tarsus; 21 : 9.5 : 8 : 7 : 11 in hind tarsus. Abdomen subparallel-sided, somewhat wider than elytra, widest around 5th and 6th segments; surface smooth; 3rd
to 7th tergites sparsely covered with minute setae and with a row of long setae along posterior
margins; 8th tergite (Figs. 116 A, 117 A) with 8 or 9 macrosetae; 9th tergite with 4 macrosetae;
10th tergite with posterior margin slightly rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 116 A) with posterior margin somewhat emarginated, its emarginate apex crenate; 8th sternite (Fig. 116 B) with posterior margin rounded, with 15–17
macrosetae; 9th sternite with posterior margin rounded. Aedeagus (Figs. 116 C, 116 D) somewhat
pear-shaped in ventral view; apical lobe almost straight in lateral view, pointed at apex in lateraland ventral views; basal ridge roundly convex extending to near bottom of basal capsule; copulatory piece of inner sac as in Fig. 116 E. Paramere with apical lobe of paramerite dilated apicad,
rounded at apex, about 1/2 as long as condylite.
Female: Eighth tergite (Fig. 117 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 117 B) with 11 or 12 macrosetae; sensory setae of 8th
sternite generalised, almost the same as the other setae in shape. Spermatheca (Fig. 117 C)
curved three times, somewhat M-shaped; basal part slightly longer than apical part, curved
around basal 1/3, slightly narrowed apicad; apical part gently curved around middle, narrowed
apicad, its inner wall densely wrinkled from apex to around apical 3/4, with outer projection.
Measurements. BL, 4.5–5.5; FBL, 2.0–2.4; HL, 0.63–0.73; HW, 0.85–0.64; EL, 0.412–
0.444; AL, 1.63–1.68; PL, 0.78–0.88; PW, 1.15–1.33; ELL, 0.99–1.18; ELW, 1.38–1.58; HTL,
0.90–1.01.
Bionomics. This species is associated with Lasius (Dendrolasius) spp. While this species iscommonly collected in Hokkaidô, normally rare in Honshû. This species is considered to be nocturnal and often collected by light trap.
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) capitatus.
Pella zhoui Maruyama, n. sp.
(Figs. 118–120)

Type material. Holotype: China (Beijing Shi): ?, “Xiaolongmen (1400 m), Dongling,
Yan-shan, Beijing, CHINA, 15–16. VI 2001, Hlaváč & Cooter leg.” (LDFJ) (IZAS).
Paratype: 1?, same data as holotype (cMar). Russia (Far East): 1 ex., Kaimanovka, Ussuryisk, Primorskyi Krai, 27 V 2005, M. Maruyama (LDFJ) (cMar); 2exs., same data but, 28 V
2005, M. Maruyama (LDN ) (cMar, cHlav); 1 ex., Bukhta Vityaz, Poluostrov Gamov, Khasanskyi,
Primorskyi Krai, 31 V 2005, M. Maruyama (LDFJ) (cMar).
Type locality. Xiaolongmen, Beijing Shi, China.
Distribution. Manchurian subregion. ASIA: China (Beijing Shi), Russia (Far East).
Etymology. Dedicated to Dr. Hong-zhang Zhou (IZAS) for his effort for permission of thecollecting trip in China conducted by P. Hlaváč and J. Cooter.
Diagnosis. Pella zhoui is similar to P. spreta in general appearance, but is distinguishedfrom the latter by the narrower pronotum, the apical lobe of aedeagal median lobe about 1/2 aslong as basal capsule and much shorter than that of P. spreta and by the slightly narrower pronotum. These two species are locally isolated. Among the sympatric species, Pella zhoui is similar
in colour and general appearance to P. beijingorum but may be easily distinguished from it by the

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165

Fig. 118. Pella zhoui Maruyama, n. sp. —A, Fore body, dorsal view; B, right antenna.

narrower pronotum.
Diagnosis. Body slender. Black in ground colour; antennae, mouthparts, legs, and around
posterior margins of 3rd to 6th tergites reddish brown; elytra yellow but with black maculations
around scutellum and postero-lateral corners. Head (Fig. 118 A) widest around middle of eyes;surface finely reticulated, moderately covered with setae; setae moderate in length, as long as
those on pronotum and elytra; length of eyes 0.45–0.46 times as long as head width. Antennae(Fig. 118 B) shorter than head, pronotum and elytra combined; 1st segment slightly longer than
2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2 as long as
1st; 4th to 10th segments wider than long; 11th segment shorter than 1st; approximate relative
lengths of segments from basal to apical: 23 : 9.5 : 12 : 6.5 : 6 : 5.5 : 5.5 : 5.5 : 6.5 : 7 : 21. Pronotum(Fig. 118 A) elliptical, 1.55–1.56 times as wide as long, widest around middle; posterior marginrounded; surface roughly punctured, finely reticulated between punctures, moderately covered
with setae, with 6 or 7 macrosetae along lateral margin. Scutellum with surface smooth, sparsely
covered with setae. Elytra (Fig. 118 A) slightly widened apicad; surface smooth, moderately covered with setae, with 5 or 6 small macrosetae laterally. Legs short; hind tibia as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 7 : 5 : 4.5 : 9.5 in fore tarsus;
11.5 : 6 : 5 : 4.5 : 8.5 in mid tarsus; 20 : 8 : 7 : 5.5 : 9.5 in hind tarsus. Abdomen subparallel-sided,
somewhat wider than elytra, widest around 5th and 6th segments; surface smooth; 3rd to 7th tergites sparsely covered with minute setae and with a row of long setae along posterior margins;
8th tergite (Figs. 119 A, 120 A) with 8 or 9 macrosetae (10 in left in holotype); 9th tergite with 4macrosetae; 10th tergite with posterior margin slightly rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 119 A) with posterior margin widely emarginated, its emarginate

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M. Maruyama

Fig. 119. Pella zhoui Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view.
Male.

apex crenate; 8th sternite (Fig. 119 B) with posterior margin rounded, with 15–17 macrosetae;
9th sternite with posterior margin rounded. Aedeagus (Figs. 119 C, 119 D) somewhat pearshaped in ventral view; apical lobe almost straight in lateral view, somewhat pointed at apex inlateral and ventral views; basal ridge roundly convex extending to near bottom of basal capsule;
copulatory piece of inner sac as in Fig. 119 E. Paramere with apical lobe of paramerite dilated
apicad, rounded at apex, about 1/2 as long as condylite.
Female: Eighth tergite (Fig. 120 A) with posterior margin slightly emarginate, its emarginate
apex weakly crenate; 8th sternite (Fig. 120 B) with 11 or 12 macrosetae; sensory setae of 8th
sternite generalised, almost the same as the other setae in shape. Spermatheca (Fig. 120 C)
curved three times, somewhat M-shaped; basal part slightly longer than apical part, curved
around basal 1/3, slightly narrowed apicad; apical part gently curved around middle, narrowed
apicad, its inner wall densely wrinkled from apex to around apical 3/4, with outer projection.

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167

Fig. 120. Pella zhoui Maruyama, n. sp. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, spermatheca. Female.

Measurements. BL, 4.8–5.3; FBL, 2.0–2.3; HL, 0.60–0.65; HW, 0.81–0.83; EL, 0.369–
0.381; AL, 1.58–1.59; PL, 0.70–0.74; PW, 1.10–1.15; ELL, 1.94–1.00; ELW, 1.31–1.36; HTL,
0.81–0.84.
Bionomics. The type series of Pella zhoui from China were collected from a colony of Lasius (Dendrolasius) fuji together with many other staphylinids (see, bionomics of P. cooterorum).
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) nipponensis.
The laticollis GroupSpecies included. Pella laticollis, P. hampei, P. indiscreta.
Distribution. European subregion, Mediterranean subregion, Manchurian subregion.
Diagnosis. Species of the laticollis group may be characterised by a combination of the following character states: 1) eye length 0.35–0.44 times as long as head width; 2) 11th antennal
segment shorter than the 1st; 3) pronotum with posterior margin completely rounded, not forming corners postero-laterally; 4) pronotum 1.42–1.57 times as wide as long; 5) pronotum widest
around middle; 6) pronotal hypomeron invisible in lateral view; 7) elytra unicoloured, reddish
brown to blackish brown; 8) inner and posterior margins of elytra weakly margined; 9) male 8thsternite longer than in female; 10) posterior margin of 8th tergite crenate; 11) posterior margin of
male 8th sternite rounded; 12) lateral projection of apodeme of male 8th tergite evidently longer
than that of female; 13) macrosetae of 8th abdominal segment generalised in length, exceeding
the posterior margins of the segment; 14) postero-medial margin of female 8th sternite without
minute pubescence; 15) apical lobe of aedeagal median lobe slightly curved in lateral view; 16)
distal crests of aedeagus developed, projected, rounded or truncate on ventral margin in lateralview; 17) copulatory piece of aedeagal median lobe gently narrowed apicad and more or lesspointed at apex in dorsal view.
Comments. The laticollis group is probably a monophyletic group in view of the close

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M. Maruyama

similarity in the general appearance, e.g., a combination of the unicoloured body and the completely elliptical pronotum. However, each of these states is homoplastic, and no autapomorphy
could be found in the species-group. This species-group is very similar to the spreta group in the
pronotal shape but aedeagal structure is quite different. No allied species-group could be found.
Symbiotic hosts. Lasius (Dendrolasius) spp., Liometopum microcephalum.
Key to the Species of the laticollis Group
1. Body smaller: BL: 3.3–4.3 mm; FBL: 1.7–2.0 mm; dorsal bridge of aedeagal median lobe
very long, much longer than apical lobe; basal part of spermatheca longer than apical part.
Distribution: Japan. ..............................................................................................P. indiscreta.
— Body larger: BL: 3.7–4.8 mm; FBL: 1.7–2.2 mm; dorsal bridge of aedeagal median lobe
short as in most Pella species; basal part of spermatheca as long as apical part. Distribution:
Europe. ................................................................................................................................ 2.
2. Reddish brown to blackish brown in ground colour; antennae slender, 5th segment almost
as long as wide; pronotum and elytra more or less matted; 8th sternite with 19–22 macrosetae. Distribution: Europe. Symbiotic host: Lasius fuliginosus. .............................P. laticollis.
— Pale brown to reddish brown in ground colour; antennae thick, clavate, 5th segment muchwider than long; pronotum and elytra evidently shining; 8th sternite with 13–19 macrosetae.
Distribution: Europe. Symbiotic host: Liometopum microcephalum. .....................P. hampei.
Pella laticollis (Märkel, 1845), n. comb.
(Figs. 121–123)
Aleochara laticollis Märkel, 1845: 203 (original description).—Mulsant & Rey, 1873a: 81 (Myrmedonia, description).—Ganglbauer, 1895: 124 (Myrmedonia (Pella), key, description).—Reitter, 1909: 43 (Myrmedonia (Myrmedonia),key).—Fenyes, 1920: 298 (Zyras (Myrmedonia), list). —Bernhauer & Scheerpeltz, 1926: 700 (Zyras (Myrmedonia),list).—Scheerpeltz, 1934: 1655 (Zyras (Myrmedonia), list). — Lohse, 1974: 226 (Zyras (Pella), key).—Likovský,
1993: 59 (Zyras (Lepla), list). — Smetana, 2004: 460 (Zyras (Lepla), list).

Type material. Syntypes: Germany: 1?, “5258/laticollis Märk Saxon. mont. Märk[“Saxon. mont., Märk.” (mountainous area of Saxonia, Germany, F. Märkel leg.)]” (glued on
triangle paper card); 3??, 2// , [ditto] (without label, glued on triangle paper card).
These are deposited in ZMHB.
Type locality. Saxonia, Germany.
Other material. Bulgaria: 1 ex., Rodopi mts, Černatica-Goljam, Persenk, 1400–1900 m,
24 VI 1996, M. Kuboň; 1 ex., Senokos 20 km W Silistra, 17 VI–11 VII 1987, Penev. Czech
Republic: 22 exs., Nový, Jičín, Moravia, 30 IV 1988, M. Kuboň; 2 exs., Soběšile, Moravia, 29.
IX 1991, P. Ph; 1 ex., Bfaclav-Lacná, Morovia, 11 IV 1981, Jonis. Germany: 3 exs., Lainzer,
Tiergarten, 30 VII 1949, O. Scheerpeltz (LDFL); 1 ex., Laxenburg, O. Scheepeltz; 1 ex.,Eberswalde-Finow, Märk Brandenburg, 16 IV 1990, Zerche; 1 ex., Baden–Württemberg, Kaiserstuhl mts., NW Freiburg, VII 1981, Kobel-Lamparski (unbaited pt); 1 ex., Gartow, Wendland,
Niedersachsen, 22 V 1988 (LDFL). Slovakia: 1 ex., Viničky-preosevpri 11 III 1995, P. Hlaváć
(LDFL).
Distribution. European subregion, Mediterranean subregion. EUROPE: Austria, Belgium,
Bosnia Herzegovina, Bulgaria, Croatia, Czech Republic, Denmark, Estonia, Finland, France,
Great Britain, Germany, Hungary, Italy, Latvia, Luxembourg, Netherlands, Norway, Poland, Ro-

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169

Fig. 121. Pella laticollis (Märkel). — A, Fore body, dorsal view; B, right antenna.

mania, Russia (North and Central European Territories), Slovakia, Sweden, Switzerland, Ukraine.
New to Bulgaria.
Diagnosis. Pella laticollis is very similar to P. hampei not only in general appearance but
also in genitalia, but can be distinguished from the latter by the slightly darker colour, the morematted body surface, the thinner antennae, and the number of macroseta of 8th sternite: 19–22.
Host ant species is different, and this species is associated with Lasius (Dendrolasius)
fuliginosus. Among the other sympatric species, Pella laticollis is somewhat similar to P. lugens
in the wide pronotum but may be easily distinguished from the latter by the smaller body and theunicoloured elytra.
Redescription. Body slender. Reddish brown to blackish brown in ground colour; antennae,
mouthparts, legs, and around posterior margins of 3rd to 6th tergites reddish paler. Head (Fig.
121 A) widest around middle of eyes; surface finely reticulated, moderately covered with setae;
setae moderate in length, longer than those on pronotum and elytra; length of eyes 0.35–0.36
times as long as head width. Antennae (Fig. 121 B) shorter than head, pronotum and elytra combined; 1st segment almost as long as 2nd and 3rd combined; 2nd segment slightly shorter than
3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments wider than long; 11th segment
shorter than 1st; approximate relative lengths of segments from basal to apical: 24 : 9.5 : 13.5 :7.5 : 6.5 : 6 : 6 : 6 : 6 : 6.5 : 21. Pronotum (Fig. 121 A) elliptical, 1.48–1.57 times as wide as long,widest around middle; posterior margin rounded; surface finely reticulated, moderately covered
with setae, with 6 or 7 macrosetae along lateral margin. Scutellum with surface smooth, moderately covered with setae. Elytra (Fig. 121 A) slightly widened apicad; surface smooth, moderately

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M. Maruyama

Fig. 122. Pella laticollis (Märkel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

covered with setae, with 5 or 6 small macrosetae laterally. Legs short; hind tibia 0.88–0.89 times
as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 5 : 5 :9.5 in fore tarsus; 7 : 6.5 : 6 : 5 : 8.5 in mid tarsus; 17 : 9 : 7.5 : 7 : 11 in hind tarsus. Abdomen subparallel-sided, somewhat wider than elytra, widest around 5th and 6th segments; surface smooth;
3rd to 7th tergites sparsely covered with minute setae and with a row of long setae along posterior margins; 8th tergite (Figs. 122 A, 123 A) with 9 (sometimes 10 or 11) macrosetae; 9th tergite
with 4 macrosetae; 10th tergite with posterior margin slightly rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 122 A) with posterior margin truncate, its apex dentate; 8th sternite(Fig. 122 B) with posterior margin rounded, with 20–22 macrosetae; 9th sternite with posterior
margin slightly emarginated. Aedeagus (Figs. 122 C, 122 D) somewhat pear-shaped in ventral
view; apical lobe slightly curved ventrad in lateral view, obtuse at apex in lateral and ventral

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171

Fig. 123. Pella laticollis (Märkel). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

views; basal ridge roundly convex, extending to near bottom of basal capsule; copulatory pieceof inner sac as in Fig. 122 E. Paramere with apical lobe of paramerite dilated apicad, rounded atapex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 123 A) with posterior margin slightly emarginatemedially,around its emarginate apex weakly dentate; 8th sternite (Fig. 123 B) with 19–21 macrosetae; sensory setae of 8th sternite generalised, almost same as other setae in shape. Spermatheca (Fig.
123 C) curved three times; basal part slightly longer than apical part, bulbous around base,
curved and narrowed around middle; apical part gently curved around middle, dilated apicad, its
inner wall sparsely wrinkled from apex to around apical 3/5, with outer projection.
Measurements. BL, 4.0–4.8; FBL, 1.7–2.2; HL, 0.60–0.66; HW, 0.69–0.78; EL, 0.253–
0.269; AL, 1.43–1.55; PL, 0.72–0.75; PW, 1.07–1.16; ELL, 0.83–0.91; ELW, 1.21–1.30; HTL,
0.73–0.81.
Bionomics. This species is associated with Lasius (Dendrolasius) fuliginosus and is common (Franc, 1992).
Symbiotic host. Lasius (Dendrolasius) fuliginosus.
Pella hampei (Kraatz, 1862), n. comb.
(Figs. 124–126)
Myrmedonia Hampei Kraatz, 1862: 267 (original description).—Ganglbauer, 1895: 124 (Myrmedonia (Pella), key, description).—Fenyes, 1920: 298 (Zyras (Myrmedonia), list).—Bernhauer & Scheerpeltz, 1926: 699 (Zyras (Myrmedonia), list).—Lohse, 1974: 226 (Zyras (Pella), key); 1989: 220 (Zyras (Pella), key).—Likovský, 1993: 59 (Zyras(Lepla), list).— Smetana, 2004: 460 (Zyras (Lepla), list).

Type material. Syntypes: Croatia: 1/, “Croat./Hampe/Coll. Kraatz/Syntypus/Hampei

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M. Maruyama

mihi Croat. Hampe/Dtsch. Entomol. Institut Berlin/Coll. DEI Eberswalde”; 2??, “Hampei Krtz
Croatia Hampe/Syntypus/D.E.I. coll. Vo Heyden/Coll. DEI Eberswalde”.
These are deposited in DEI.
Type locality. Croatia.
Other material. Bulgaria: 1/, St. Ivan, Sofia, 3 VIII 1908, F. Rambousek (NMP).
Croatia: Reitter (no other data) (cMar); 1?, “Hampei Kr., Croatia, Kraatz/c. Eppelsh. Steind. d.”(NHMW); 1/, “Hampei Croatia Schauffs/c. Eppelsh. Steind. d./Hampei Kraatz Berl. Zeit., 1862.
p. 267” (NHMW); 1?, “Hampei Krtz Croatia Miller/D.E.I. coll. Vo Heyden/Coll. DEI Eberswalde” (DEI); 1 ex., “Croatia Apfelbeck/Hampei Kr.” (NHMW); 1 ex., “Croatia/Hampe/hampei
Kr.” (NHMW). Hungary: 1 ex., “c. Eppelsh. Steind. d./Hampei Kr. Hungar. Geb. Dr. Chyici.”(NHMW); 1 ex., “c. Eppelsh. Steind. d./Hampei Hungar” (NHMW). Yugoslavia: 1?, Topčider,
Beograd, VI 1910, F. Rambousek (NMP).
Distribution. European subregion, Mediterranean subregion. EUROPE: Albania, Austria,
Bulgaria, Croatia, Czech Republic, Greece, Hungary, Slovakia, Yugoslavia. New to Bulgaria.
Diagnosis. Pella hampei is very similar to P. laticollis not only in general appearance but
also in genitalia, but can be distinguished from the latter by the slightly paler colour, the moreshining body surface, the clavate antennae, and the number of macroseta of 8th sternite 13–19.
Host ant species is differentiated, and this species is associated with Liometopum microcephalum.
Among the other sympatric species, Pella hampei is somewhat similar to P. lugens in the wide
pronotum but may be easily distinguished from the latter by the smaller body and the unicoloured
elytra. Pella lugens is associated with Lasius (Dendrolasius) fuliginosus.
Redescription. Body slender. Pale brown to reddish brown in ground colour; antennae,

Fig. 124. Pella hampei (Kraatz). — A, Fore body, dorsal view; B, right antenna.

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173

Fig. 125. Pella hampei (Kraatz). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

mouthparts, legs, and around posterior margins of 3rd to 6th tergites reddish paler. Head (Fig.
124 A) widest around middle of eyes; surface finely reticulated, moderately covered with setae;
setae moderate in length, longer than those on pronotum and elytra; length of eyes 0.43–0.44
times as long as head width. Antennae (Fig. 124 B) shorter than head, pronotum and elytra combined; 1st segment almost as long as 2nd and 3rd combined; 2nd segment slightly shorter than
3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments wider than long; 11th segment
shorter than 1st; approximate relative lengths of segments from basal to apical: 20 : 8.5 : 11.5 :5.5 : 5.5 : 5.5 : 5.5 : 5 : 5 : 6 : 19. Pronotum (Fig. 124 A) elliptical, 1.54–1.61 times as wide as long,widest around middle; posterior margin rounded; surface finely reticulated, moderately covered
with setae, with 6 or 7 macrosetae along lateral margin. Scutellum with surface smooth, moderately covered with setae. Elytra (Fig. 124 A) slightly widened apicad; surface smooth, moderately
covered with setae, with 5 or 6 small macrosetae laterally. Legs short; hind tibia 0.82–0.86 times
as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 5 : 4.5 :174

M. Maruyama

Fig. 126. Pella hampei (Kraatz). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, D, spermathecae. Female.

9.5 in fore tarsus; 10 : 6 : 5 : 5 : 9 in mid tarsus; 17 : 9 : 7 : 6.5 : 10 in hind tarsus. Abdomen subparallel-sided, somewhat wider than elytra, widest around 5th and 6th segments; surface smooth; 3rd
to 7th tergites sparsely covered with minute setae and with a row of long setae along posterior
margins; 8th tergite (Figs. 125 A, 126 A) with 8 (sometimes 7) macrosetae; 9th tergite with 4macrosetae; 10th tergite with posterior margin slightly rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 125 A) with posterior margin truncate, its apex dentate; 8th sternite(Fig. 125 B) with posterior margin rounded, with 18 or 19 macrosetae; 9th sternite with posterior
margin slightly emarginated. Aedeagus (Figs. 125 C, 125 D) somewhat pear-shaped in ventral
view; apical lobe slightly curved ventrad in lateral view, obtuse at apex in lateral and ventral
views; basal ridge roundly convex extending to near bottom of basal capsule; copulatory piece of
inner sac as in Fig. 125 E. Paramere with apical lobe of paramerite dilated apicad, rounded atapex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 126 A) with posterior margin truncate or slightly emarginate medially, its apex weakly dentate; 8th sternite (Fig. 126 B) with 13–15 macrosetae; sensory setae of
8th sternite generalised, almost the same as the other setae in shape. Spermatheca (Figs. 126 C,
126 D) curved three times; basal part slightly longer than apical part, bulbous around base,
curved and narrowed around middle; apical part gently curved around middle, dilated apicad, its
inner wall sparsely wrinkled from apex to around apical 3/5, with outer projection.
Measurements. BL, 3.7–4.0; FBL, 1.8–1.9; HL, 0.57–0.60; HW, 0.69–0.70; EL, 0.249–
0.265; AL, 1028–1.38; PL, 0.67–0.69; PW, 1.04–1.08; ELL, 0.80–0.83; ELW, 1.20–1.22; HTL,
0.66–0.71.

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175

Bionomics. This species is collected from nests of Liometopum microcephalum usually together with Pella ruficollis. This species occurs very locally and rarely in Central Europe probably because of rarity of the host ants (Franc, 1992).
Symbiotic host. Liometopum microcephalum.
Pella indiscreta (Sharp, 1888), n. comb.
(Figs. 127–129)
Myrmedonia indiscreta Sharp, 1888: 291. — Fenyes, 1920: 298 (Zyras (Myrmedonia), list).—Bernhauer & Scheerpeltz,
1926: 700 (Zyras (Myrmedonia), list). — Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntypes. Japan: 1/, “Myrmedonia indiscreta Type D.S. Japan 31. VII.
1881. Lewis./Type (red round curator label)/Shimonosuwa 31. 7. 81./Japan. G. Lewis./Sharp
Coll., 1905-313/SYN-TYPE” (BMNH); 1/, “Sap (written underside of paper card specimen
glued on)/Sapporo. 5. VIII.–16. VIII. 80./Japan. G. Lewis./Sharp Coll. 1905-313” (BMNH).
Other material. Japan: Hokkaidô: 1?, 1/, 37 exs., Nozaki, Bihoro-chô, 23 VI 2001, Y.
Yamaga (LDC ); 1?, Hitsujigaoka, Sapporo-shi, 21 IX 1998, M. Maruyama (LDS); 4 exs., Kannon-zawa, Sapporo-shi, 31 V 2002, M. Maruyama (LDFJ); 2 exs., Akagawa, Hakodate-shi, 15
VIII 1992, M. Nishikawa (LDFJ) (cMar). Honshû: 30 exs., Aobayama, Sendai-shi, Miyagi-ken,
22 IX 2001, M. Maruyama (LDN ); 2 exs., Kinshû-ko, Yuda-machi, Waga-gun, Iwate-ken, 19 V
2004, H. Kamezawa (LDFJ); 2 exs., Yukiwari-bashi, Nishigô-mura, Fukushima-ken, 29 VII
2000, T. Kobayashi (LDC); 2??, 2// , 6 exs., Aoyagidaira, Kuriyama-mura, Tochigi-ken, 1 IX
1999, T. Kobayashi (LDN ); 5 exs., Tobiyama-jôshi, Môka-shi, Tochigi-ken, 17 VI 1998, M.
Maruyama (LDFJ); 6 exs., Shimokomoriya, Utsunomiya-shi, Tochigi-ken, 6 VII 1999, M.
Maruyama (LDFJ); 1 ex., Akebono, Shibukawa-shi, Gumma-ken, 1 VII 2004, T. Watanabe(cWat); 3 exs., Nakano, Shôwa-machi, Kitakatsushika-gun, Saitama-ken, 4 V 2003, H. Kamezawa (LDFJ); 2 exs., same data but, 25 V 2003; 1 ex., Tokigawa, Kamagata, Ranzan-machi, Saitama-ken, 22 IV 1999, K. Toyoda (LDFJ); 4 exs., Akigase-kôen, Urawa-shi, Saitama-ken, 19 V
2000, H. Sugaya (LDFJ); 3 exs., Daiyûzan, Hakone-machi, Kanagawa-ken, 31 VIII 1982, Y. Shibata (LDFJ) (SCM); 1 ex., Fukazawa-jûnimagari, Nakamaru, Nagasaka-chô, Kitakoma-gun,
Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 1 ex., Kamikandori, Akeno-mura, Kitakomagun, Yamanashi-ken, 6 VI 2004, M. Maruyama (LDFJ); 1 ex., Jôyama, Matsumoto-shi, Naganoken, 12–13 VII 1998, M. Maruyama (LDN ). Shikoku: 1 ex., Ôtaki-san, Shionoe-chô, Kagawaken, 2 VI 2001, M. Maruyama & Y. Kamite (LDS).
Type locality. Shimonosuwa, Nagano-ken, or Sapporo-shi, Hokkaidô, Japan.
Distribution. Manchurian subregion. ASIA: Japan (Hokkaidô, Honshû, Shikoku).
Diagnosis. Pella indiscreta is similar to P. laticollis and P. hamepei in general appearance,
but is distinguished from the latter two by the evidently smaller body and the dorsal bridge of
aedeagal median lobe longer. Pella indiscreta and the latter two are locally isolated. Among the
sympatric species Pella indiscreta is somewhat similar to P. kinomurai in general appearance but
distinguished from the latter by the much smaller body.
Redescription. Body slender. Pale brown to reddish brown in ground colour; antennae,
mouthparts, legs, and around posterior margins of 3rd to 6th tergites reddish paler. Head (Fig.
127 A) widest around middle of eyes; surface finely reticulated, densely covered with setae; setae
moderate in length, longer than those on pronotum and elytra; length of eyes 0.39–0.42 times aslong as head width. Antennae (Fig. 127 B) shorter than head, pronotum and elytra combined; 1stsegment almost as long as 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd seg-

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M. Maruyama

Fig. 127. Pella indiscreta (Sharp). — A, Fore body, dorsal view; B, right antenna.

ment about 1/2 as long as 1st; 4th to 10th segments wider than long; 11th segment shorter than
1st; approximate relative lengths of segments from basal to apical: 21.5 : 9.5 : 13 : 6.5 : 6.5 : 5.5 :6 : 6 : 6.5 : 7 : 19.5. Pronotum (Fig. 127 A) elliptical, 1.42–1.50 times as wide as long, widest
around middle; posterior margin rounded; surface finely reticulated, densely covered with setae,
with 4 or 5 macrosetae along lateral margin. Scutellum with surface smooth, moderately covered
with setae. Elytra (Fig. 127 A) slightly widened apicad; surface smooth, densely covered withsetae, with 3 or 4 small macrosetae laterally. Legs moderate in length; hind tibia 0.92–0.97 times
as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 6 : 5 : 4.5 :9.5 in fore tarsus; 10 : 8 : 7 : 5.5 : 9 in mid tarsus; 18 : 11 : 8.5 : 5.5 : 11.5 in hind tarsus. Abdomen
subparallel-sided, somewhat wider than elytra, widest around 5th and 6th segments; surface
smooth; 3rd to 7th tergites sparsely covered with minute setae and with a row of long setae alongposterior margins; 8th tergite (Figs. 128 A, 129 A) with 8 (sometimes 7) macrosetae; 9th tergite
with 4 macrosetae; 10th tergite with posterior margin slightly rounded, with 4 macrosetae.
Male: Eighth tergite (Fig. 128 A) with posterior margin truncate, its apex dentate; 8th sternite(Fig. 128 B) with posterior margin rounded, with 14–17 macrosetae; 9th sternite with posterior
margin slightly emarginated. Aedeagus (Figs. 128 C, 128 D) elongate pear-shaped in ventral
view; apical lobe slightly curved ventrad in lateral view, obtuse at apex in lateral and ventral
views; dorsal bridge very long; basal ridge roundly convex, extending near bottom of basal capsule; copulatory piece of inner sac as in Fig. 128 E. Paramere with apical lobe of paramerite dilated apicad, rounded at apex, about 1/3 as long as condylite.
Female: Eighth tergite (Fig. 129 A) with posterior margin truncate or slightly emarginate me-Palearctic Species of Pella

177

Fig. 128. Pella indiscreta (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, copulatory piece, dorsal view. Male.

dially, its apex weakly dentate; 8th sternite (Fig. 129 B) with 11 or 12 macrosetae; sensory setaeof 8th sternite generalised, almost the same as the other setae in shape. Spermatheca (Fig. 129 C)
curved three times; basal part about twice as long as apical part, slightly bulbous around base,
curved near base, its inner wall moderately wrinkled; apical part gently curved around base, dilated apicad, its inner wall densely wrinkled from apex to around apical 3/4.
Measurements. BL, 3.3–4.3; FBL, 1.7–2.0; HL, 0.55–0.60; HW, 0.65–0.68; EL, 0.254–
0.288; AL, 1.32–1.55; PL, 0.70–0.65; PW, 0.93–1.00; ELL, 0.71–0.81; ELW, 1.02–1.13; HTL,
0.66–0.79.
Bionomics. This species is collected from nests of Lasius (Dendrolasius) spp. mainly from
July to October (see, also Life cycle). Four ant species are known as the symbiotic hosts of this178

M. Maruyama

Fig. 129. Pella indiscreta (Sharp). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral
view; C, spermatheca. Female.

species, but very rare in colonies of Lasius (Dendrolasius) fuji and L. (D.) spathepus. In contrast,
a number of specimens were collected from colonies of L. (D.) nipponensis and L. (D.) capitatus.
Thus, this species is considered to have more or less host preference to Lasius (Dendrolasius)
nipponensis and L. (D.) capitatus.
Symbiotic hosts. Lasius (Dendrolasius) fuji, L. (D.) spathepus, L. (D.) nipponensis, L. (D.)
capitatus.
Incertae Sedis
Pella pumila (Fiori, 1914), n. comb.
Zyras pumilus Fiori, 1914: 105 (original description). — Bernhauer & Scheerpeltz, 1926: 703 (list).—Smetana, 2004:
461 (Zyras (Pella), list).

Type material. Not examined.
Type locality. Bosco (Perugia), Umbria, Italy.
Comments. Unfortunately, the type material of this species was not found. After Fiori(1914), the type specimen was collected from a nest of Liometopum microcephalum in Italy together with Pella ruficollis and P. laticollis, and this species was similar to P. cognata. These suggest that this species is most probably a junior synonym of Pella similis, which is sometimes collected from a nest of Liometopum microcephalum and considerably similar to P. cognata in general appearance. Moreover, the original description fits well with that of Pella similis.

Palearctic Species of Pella

179

The Species that should be Excluded from Pella
Genus Zyras Stephens, 1835
Zyras Stephens, 1835: 430 (original naming without description) [type species: Aleochara haworthi Stephens, 1832, bymonotypy].—Fenyes, 1920: 293 (description). — Bernhauer & Scheerpeltz, 1926: 693 (list).—Scheerpeltz, 1934:
1648 (list).

Subgenus Zyras Stephens, 1835
Zyras Stephens, 1835: 430 [type species: Aleochara haworthi Stephens, 1832].—Thomson, 1860: 258 (list).—Mulsant& Rey, 1873a: 35 (list). — Reitter, 1909 (description): 42.—Bernhauer & Scheerpeltz, 1926: 694 (list).—Scheerpeltz, 1934: 1650 (list).

Comments. Although the subgenus Zyras of the genus Zyras has not been well defined, thefollowing three species are considered to belong to the subgenus because of their close relationship with the type species, Zyras haworthi, in the following character states: body shining, without microsculpture; body irregularly covered with small setae; 1st segment of labial palpus much
longer than 2nd; ligula long, with minute setae at apex; pronotum coarsely punctured; pronotum
sparsely covered with long setae.
Many species of this subgenus are known in southern Asia. However, most of them werebriefly described and have not been redescribed. Thus, precise systematic position of the species
redescribed in the present study could not be determined at present.
Zyras (Zyras) coloratus Cameron, 1939, n. assignment(Figs. 130–131)
Zyras (Pella?) coloratus Cameron, 1939: 545 (original description).—Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Syntypes: India: 1/, “Dehra Dun, Dr. M. Cameron, 8–8, 1921/Z., coloratus, Cam, TYPE/SYNTYPE (blue round curator label)” (BMNH); 1/, “Dehra Dun, Dr. M.
Cameron, 9–9, 1922/N. GEN/SYNTYPE (blue round curator label)” (BMNH).
Type locality. Dehra Dun, India.
Other material. Not examined.
Distribution. Oriental region: Indian subregion. ASIA: India.
Diagnosis. Among the examined species of the genus Zyras, Z. coloratus is considered to
be closely allied to Z. pictus (Sharp, 1874) described from Japan and some species widely distributed in Southeast Asia in general appearance and punctulation of body surface but can be distinguished from the latter species by the clavate antennae, the larger head which is only slightly
narrower than the pronotum and the macrosetae of the pronotum, elytra and abdomen much
shorter.
Redescription. Body slender, slightly matted. Reddish brown in ground colour; pronotum,
elytra, legs, and 2nd to 4th and base of 5th abdominal segments yellow. Head (Fig. 130 A) widest
at eyes; surface smooth, moderately covered with setae; setae moderate in length, as long as
those on pronotum and elytra; eyes prominent and its length 0.46–0.48 times as long as headwidth. Antennae (Fig. 130 B) clavate, shorter than head, pronotum and elytra combined; 1st segment slightly longer than 2nd and 3rd combined; 2nd segment slightly longer than 3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments wider than long; 11th segment shorter than
1st; approximate relative lengths of segments from basal to apical: 19 : 9.5 : 8.5 : 3.5 : 3 : 3.5 : 3.5 :3.5 : 4 : 4 : 15. Pronotum (Fig. 130A) subtrapeziform, 1.19–1.28 times as wide as long, widest

180

M. Maruyama

Fig. 130. Zyras (Zyras) coloratus Cameron.—A, Fore body, dorsal view; B, right antenna.

around anterior 1/5, narrowed posteriorly; posterior margin rounded; surface sparsely covered
with setae and punctures, with 7 or 8 macrosetae along lateral margin. Scutellum with surface
smooth, with 2 or 3 setae. Elytra (Fig. 130 A) gently widened apicad; surface moderately covered
with setae and punctures, with 4 or 5 small macrosetae laterally. Legs moderate in length; hindtibia 0.91–0.92 as long as elytra; approximate relative lengths of tarsal segments from basal toapical: 3 : 4 : 4 : 12 in fore tarsus; 6 : 5 : 5 : 4.5 : 10 in mid tarsus; 12 : 8 : 7 : 6 : 10 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 5th and 6th segments; surface smooth on 2nd to 7th tergites and finely punctured on 8th tergite; 3rd to 6th tergites sparsely
and uniformly covered with minute setae, except a row of long setae along posterior margin; 7th
tergite moderately covered with long setae; 8th tergite (Fig. 131 A) with 9 or 10 macrosetae; 9th
tergite with 6 or 7 macrosetae; 10th tergite with posterior margin slightly rounded, with 3
macrosetae.
Male unknown.
Female: Eighth tergite (Fig. 131 A) with posterior margin slightly rounded; 8th sternite (Fig.
131 B) slightly emarginate postero-medially, with 7 macrosetae; sensory setae of 8th sternite
spindle-shaped, flattened. Spermatheca (Figs. 131 C, 131 D) with duct coiled about 10 times
around base and coiled once near apex; basal part and apical part gently curved around middle,
dilated apicad.
Measurements. BL, 3.9–4.3; FBL, 1.6–1.8; HL, 0.50–0.53; HW, 0.61–0.63; EL, 0.152–
0.161; AL, 1.19–1.25; PL, 0.59–0.60; PW, 0.71–0.75; ELL, 0.74–0.75; ELW, 1.00–0.96; HTL,
0.67–0.69.
Bionomics. No information.

Palearctic Species of Pella

181

Fig. 131. Zyras (Zyras) coloratus Cameron. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca; D, apex of spermatheca. Female.

Zyras (Zyras) iridescens (Sawada, 1970), n. assignment
Bolitochara iridescens Sawada, 1970b: 49 (original description).—Gürlich, 1981: 211 (Lepla, chaetotaxy).—Smetana,
2004: 460 (Zyras (Lepla), list).

Comments. This species was originally described from Japan under the genus Bolitochara(Sawada, 1970b), and at that time, Bolitochara was assigned by authors to the members of Zyras.
Gürlich (1981) regarded this species as a member of the genus Lepla based on chaetotaxy of the182

M. Maruyama

abdomen in referring the original description. However, this species is typically a member ofZyras, and is closely allied to the type species of the genus Zyras: Z. haworthi.
This species will be redescribed in a revision of the Japanese species of the genus Zyras(Maruyama, in prep.).
Zyras (Zyras) quasar Dvořák, 1996, n. assignment(Figs. 132, 133)
Zyras (Pella) quasar Dvořák, 1996: 6 (original description).

Type material. Holotype: Vietnam: ?, “Vietnam 20–30. 12. 1980 Taj Nguen [TháiNguyên] Bud Lioj 65 km N Ankche A Pokarzvekij legit./HOLOTYPE/Zyras quasar sp. n., det.
M. Dvořák, 90” (cDvor).
Type locality. Thái Nguyên, Bud Lioj, Vietnam.
Other material. Not examined.
Distribution. Oriental region: Indo-Chinese subregion. ASIA: Vietnam.
Diagnosis. Zyras quasar may be distinguished from the other species of the genus by a
combination of the following character states in addition to aedeagal shape: 1) body almost uniformly reddish brown; 2) pronotal disc with surface smooth; 3) depressions of 3rd to 6th tergites
with a row of punctures.
Redescription. Body slender, shining. Reddish brown in ground colour; antennae darker;
mouthparts, around antero-lateral corners of elytra, legs and paratergites paler. Head (Fig. 132 A)
widest at eyes; surface smooth, sparsely covered with setae; setae long, as long as those on

Fig. 132. Zyras (Zyras) quasar Dvořák. —A, Fore body, dorsal view; B, right antenna.

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183

Fig. 133. Zyras (Zyras) quasar Dvořák. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view; E, ditto, dorsal view; F, apicallobe of paramere, lateral view. Male.

pronotum and elytra; eyes prominent, large, its length 0.45 times as long as head width. Antennae(Fig. 132 B) slender, shorter than head, pronotum and elytra combined; 1st segment slightly
shorter than 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rd segment about 1/2
as long as 1st; 4th to 7th segments as long as wide; 8th to 10th segments slightly wider than long;
11th segment shorter than 1st; approximate relative lengths of segments from basal to apical: 22 :11 : 12 : 8 : 7.5 : 7 : 7 : 7 : 7 : 7.5 : 21. Pronotum (Fig. 132 A) almost circular, 1.14 times as wide as
long, widest around anterior 1/5, narrowed posteriorly; posterior margin rounded; surface

184

M. Maruyamasmooth, sparsely covered with long and suberect setae, with 5 or 6 macrosetae along lateral margin. Scutellum with surface roughly punctured, about 10 setae. Elytra (Fig. 132 A) gently
widened apicad; surface moderately punctured and covered with long setae, with 4 or 5 macrosetae laterally. Legs moderate in length; hind tibia 1.10 times as long as elytra; approximate relative
lengths of tarsal segments from basal to apical: 4.5 : 6 : 5 : 13.5 in fore tarsus; 10 : 8 : 5.5 : 5 : 11.5
in mid tarsus; 20.5 : 9.5 : 8 : 7 : 14 in hind tarsus. Abdomen subparallel-sided, slightly narrower
than elytra, widest around 4th and 5th segments; surface smooth, except a row of punctures on
depression of 3rd to 6th tergites and punctures around base of 7th tergite; 8th tergite (Fig. 133 A)
with 11 macrosetae; 9th and 10th abdominal segments missing in the holotype.
Male: Eighth tergite (Fig. 133 A) with posterior margin rounded; 8th sternite (Fig. 133 B)
with posterior margin rounded, with 18–22 macrosetae. Aedeagus (Figs. 133 C–133 E) oval inventral view; apical lobe slightly curved ventrad, somewhat pointed at apex in lateral view, obtuse at apex in ventral view; basal ridge absent. Paramere with apical lobe of paramerite dilated
apicad, rounded at apex, 1/4 as long as condylite.
Female unknown.
Measurements. BL, 5.1; FBL, 2.2; HL, 0.6; HW, 0.73; EL, 0.329; AL, 1.67; PL, 0.77; PW,
0.93; ELL, 0.89; ELW, 1.16; HTL, 0.98.
Comments. Since the middle of the 1980’s, many species of Zyras have been describedfrom southern Asia by Pace (1984a, 1984b, 1985, 1986, 1990, 1992, 1993, 1998a, 1998b, 1999,
2000). Dvořák (1996) did not compare his type of Zyras quasar with any species of Zyras of later
description since he misidentified this species as a member of Pella. Pace only briefly describedmany species often on the basis of female material alone, though identification of species of thesubgenus Zyras is usually very difficult without referring punctulation of body and structure of
aedeagus. Thus, precise systematic position and validity of this species is unknown at present.
Only the holotype has been known. The sex of the holotype was indicated as female in theoriginal description (Dvořák, 1996), but in fact it was a male. The specimen is considerably damaged, and the 9th and 10th abdominal segments are missing. The aedeagus is in a bad condition,
somewhat spoiled, and the copulatory is invisible externally.
Bionomics. No information.
Subgenus Glossacantha Gemminger & Harold, 1868
Acanthoglossa Motschulsky, 1859: 88 (original description, preoccupied name) [type species: Acanthoglossa badia
Motschulsky, 1859: 89, by monotypy].
Glossacantha Gemminger & Harold, 1868: 519 (replacement name for preoccupied Acanthoglossa). — Fenyes, 1920: 276(distinct genus, description). — Bernhauer & Scheerpeltz, 1926: 694 (subgenus of Zyras, list).—Scheerpeltz, 1934:
1649 (subgenus of Zyras, list). — Cameron, 1939: 518 (description, as subgenus of Zyras).

Diagnosis. Glossacantha has not been well defined but can be distinguished from the othergenera/subgenera of the Lomechusini by a combination of the following character states: 1) maxillary palpus elongate; 2) 2nd segment of maxillary palpus depressed; 3) lateral lobe of premental
apodeme rounded at apex; 4) 1st antennal segment depressed dorsoventrally; 5) 2nd to 11th antennal segments depressed laterally; 6) 4th to 10th antennal segments evidently widened apicad,
somewhat triangular.
Comments. Glossacantha has been regarded as a subgenus of the genus Zyras. However,Zyras is considered to be a polyphyletic group (Maruyama, in prep.), and Glossacantha is far related to it. Therefore, Glossacantha could be regarded as a distinct genus. However, I was not

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185

able to examine all the species known in Glossacantha, and tentatively arranged it as a subgenus
of Zyras in the present study.
In my preliminary research on the subgenera of Zyras, most of them should be synonymized
and amalgamated with other genera. For example, no important differences of subgeneric value
have been found among the subgenera Glossacantha, Diaulaconia Bernhauer, 1928, andTaprodonia Cameron, 1939. Moreover, they are quite distantly related to Zyras (Zyras). Thus, it
is appropriate to regard Glossacantha (the oldest subgenus) as a distinct genus, and in the future
the other subgenera should be synonymized with it.
Many similar problems remain in the taxonomy of the Lomechusini. The entire tribe needs
revision on a worldwide basis.
Zyras (Glossacantha) ceylonicus Cameron, 1939, n. assignment(Figs. 134, 135)
Zyras (Pella) ceylonicus Cameron, 1939: 544 (original description).

Type material. Syntypes: Sri Lanka: 1/, “SYN-TYPE (blue round curator label)/Kandy
Ceylon/M. Cameron Bequest. B. M. 1955-147./Z. ceylonicus TYPE Cam.” (BMNH); 1/, SYNTYPE (blue round curator label)/Ceylon/Sharp Coll 1905-313./1119 : Ceylon (round label, written on two faces)/Z. ceylonicus Cam./near but not M. planaticollis Kr.”
These are deposited in BMNH.
Other material. Not examined.
Type locality. Kandy, Sri Lanka.
Distribution. Oriental region: Ceylonese subregion. ASIA: Sri Lanka.

Fig. 134. Zyras (Glossacantha) ceylonicus Cameron.—A, Fore body, dorsal view; B, right antenna.

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M. Maruyama

Fig. 135. Zyras (Glossacantha) ceylonicus Cameron. —A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca. Female.

Diagnosis. Zyras (Glossacantha) ceylonicus is closely allied to Z. (G.) beckeri Scheerpeltz,
1963, originally described from “Bengalore” [Bangladesh], but can be distinguished from the latter by the antero-lateral angle of the pronotum more prominent and the setae on the pronotumand elytra much shorter.
Redescription. Body slender. Reddish brown in ground colour; inner and posterior margins of elytron, around posterior margin of 3rd to 5th abdominal segments yellow; abdomen
darker. Head (Fig. 134 A) widest just behind eyes; surface finely reticulated, sparsely covered
with setae; setae short, slightly longer than those on pronotum and elytra; length of eyes 0.36
times as long as head width. Antennae (Fig. 134 B) shorter than head, pronotum and elytra combined; 1st segment slightly as long as 2nd and 3rd combined; 2nd segment slightly shorter than
3rd; 3rd segment about 1/2 as long as 1st; 4th to 10th segments wider than long; 11th segment
shorter than 1st; approximate relative lengths of segments from basal to apical: 23 : 10 : 13 : 8 :8.5 : 8 : 7.5 : 8 : 8.5 : 8 : 21.5. Pronotum (Fig. 134 A) semicircular, 1.20–1.25 times as wide as long,widest just behind anterior margin, narrowed posteriorly; posterior margin rounded; surface reticulated, sparsely covered with setae, much sparser than on head, without macrosetae; hypomeron
fully visible in lateral view. Scutellum with surface rugose, moderately covered with setae. Elytra(Fig. 134 A) subparallel-sided; surface reticulated, sparsely covered with setae, almost the sameas on head in density, without macrosetae. Legs moderate in length; hind tibia 0.96–1.02 times aslong as elytra; approximate relative lengths of tarsal segments from basal to apical: 9 : 8 : 8 : 15.5
in fore tarsus; 12 : 11 : 7 : 6 : 15 in mid tarsus; 20 : 11 : 9 : 8.5 : 17 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 4th and 5th segments; surface finely reticulated; 3rd to 7th tergites sparsely and uniformly covered with minute setae; 8th tergite (Fig.

Palearctic Species of Pella

187

135 A) with 10 macrosetae; setae on lateral areas of 8th tergite thick and poorly differentiatedfrom macrosetae; 9th tergite with 3 macrosetae; 10th tergite with postero-medial margin minutely projected, with 5 or 6 macrosetae.
Male unknown.
Female: Eighth tergite (Fig. 135 A) with posterior margin truncate; 8th sternite (Fig. 135 B)
with 10 or 11 macrosetae, without sensory setae. Spermatheca (Fig. 135 C) curved twice; basal
part about 2.5 times as long as apical part, acutely curved around middle, slightly narrowed apicad; flattened at apex.
Measurements. BL, 5.9–6.3; FBL, 2.5–2.6; HL, 0.77–0.78; HW, 0.95–0.96; EL, 0.346–
0.351; AL, 1.80–1.93; PL, 0.90–0.93; PW, 1.1–1.1; ELL, 0.98–1.08; ELW, 1.39–1.43; HTL,
1.00–1.04.
Bionomics. No information.
Zyras (Glossacantha) reelsi Pace, 1998, n. assignment(Figs. 136–138)
Zyras (Pella) reelsi Pace, 1998a: 973 (original description). —Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Holotype: China (Hongkong): ?, “Hong Kong Shek Kwu Chao 3. VII. 96
U.V. light G. T. Reels/Zyras reelsi sp. n. det. R. Pace 1996/HOLOTYPUS Zyras reelsi mihi det.
R. Pace 1996” (MHNG).
Paratypes: China (Hongkong): 1?, 3// , “Hong Kong Beaufort 1. V 96, G. T. Reels
leg./PARATYPI Zyras reelsi det. R. Pace 96” (cMar, ex cRoug).
Other material. Not examined.
Type locality. Shek Kwu Chao, Hongkong, China.
Distribution. Oriental region: Indo-Chinese subregion. ASIA: China (Hongkong).
Diagnosis. Zyras (Glossacantha) reelsi is most similar in general appearance to Z. (G.) reversus Cameron, 1939, but is distinguished from the latter by the projections of 3rd tergite andparatergite reaching the posterior margin of 4th tergite and the absence of ridge on postero-medial area of 7th tergite.
Redescription. Body slender. Brown in ground colour; pronotum, elytra, legs, and posterior margins of 3rd to 7th abdominal segments reddish brown. Head (Fig. 136 A) widest just behind eyes; surface finely reticulated, sparsely covered with setae; setae moderate in length, aslong as those on pronotum and elytra; eyes very large, prominent, its length 0.45–0.46 times aslong as head width. Antennae (Fig. 136 B) shorter than head, pronotum and elytra combined; 1stsegment slightly longer than 2nd and 3rd combined; 2nd segment slightly shorter than 3rd; 3rdsegment about 1/2 as long as 1st; 4th to 10th segments wider than long; 11th segment shorter
than 1st; approximate relative lengths of segments from basal to apical: 41 : 14 : 21 : 12 : 12 : 12 :11 : 11 : 11 : 11 : 25. Pronotum (Fig. 136 A) subtrapeziform, 1.15–1.21 times as wide as long,widest just behind anterior margin, narrowed posteriorly; posterior margin rounded; surface finely reticulated, moderately covered with short setae, with 8 or 9 macrosetae along lateral margin.
Scutellum with surface smooth, without setae. Elytra (Fig. 136 A) slightly widened apicad; surface finely reticulated, moderately covered with setae, with about 15 macrosetae laterally. Legs
moderate in length; hind tibia 0.98–1.01 times as long as elytra; approximate relative lengths oftarsal segments from basal to apical: 9 : 11 : 9 : 21 in fore tarsus; 15 : 12 : 10 : 9 : 18 in mid tarsus;
20 : 16 : 13 : 12 : 22 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra,
widest around 3rd segment; surface smooth, sparsely punctured and covered with setae; 9th ter-

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M. Maruyama

Fig. 136. Zyras (Glossacantha) reelsi Pace. — A, Fore body, dorsal view; B, right antenna; C, 3rd tergite andparatergite. Male.

gite with 8 macrosetae; 10th tergite with posterior margin slightly emarginate, with 10 macrosetae.
Male: Third tergite (Fig. 136 C) with a long postero-medial projection; 3rd paratergite (Fig.
136 C) acutely produced and its apex reaching posterior margin of 4th tergite; 8th tergite (Fig.
137 A) with posterior margin moderately emarginate, with a pair of granula on postero-medial
area and 10 macrosetae; 8th sternite (Fig. 137 B) with posterior margin rounded, with 15–17
macrosetae; 9th sternite with posterior margin truncate. Aedeagus (Figs. 137 C, 137 D) elongate
oval in ventral view; apical lobe almost straight, obtuse at apex in lateral and ventral views; basal
ridge absent; copulatory piece of inner sac as in Figs. 137 C, 137 D. Paramere with apical lobe ofparamerite narrowed apicad, obtuse at apex, about 1/2 as long as condylite.
Female: Third abdominal segment generalised; 8th tergite (Fig. 138 A) with posterior margin
slightly emarginate; 8th sternite (Fig. 138 B) with 15–17 macrosetae; apical margin with a row ofminute pubescence. Spermatheca (Fig. 138 C) gently curved; basal part three times as long asapical part, bulbous around base, narrowed apicad; apical part almost straight, narrowed apicad.
Measurements. BL, 5.9–6.5; FBL, 3.1–3.2; HL, 0.82–0.83; HW, 1.13–1.16; EL, 0.51–
0.53; AL, 2.24–2.35; PL, 1.07–1.10; PW, 1.27–1.30; ELL, 1.30–1.35; ELW, 1.63–1.67; HTL,
1.28–1.40.
Bionomics. The type series of Zyras (Glossacantha) reelsi were collected by UV light trap

Palearctic Species of Pella

189

Fig. 137. Zyras (Glossacantha) reelsi Pace. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, lateral view; D, ditto, ventral view. Male.

(Pace, 1998), but nothing is known about their habitat. The male specimen of the paratypes,
which collected in May, is not fully mature, while the other specimens collected on the same day
are well mature. This suggests the multivoltine life cycle of this species.
Genus Myrmoecia Mulsant & Rey, 1874
Myrmoecia Mulsant & Rey, 1874: 98 (original description) [type species: Myrmedonia tuberiventris Fairmaire, 1855,
fixed by Mulsant & Rey (1874)]. — Fenyes, 1920: 293 (subgenus of Zyras, description).—Bernhauer & Scheerpeltz,

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M. Maruyama

Fig. 138. Zyras (Glossacantha) reelsi Pace. — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, spermatheca. Female.
1926: 694 (subgenus of Zyras, list). — Scheerpeltz, 1934: 1649 (subgenus of Zyras, list). — Lohse, 1974: 226 (subgenus of Zyras, description). — Lohse, 1989: 220 (distinct genus).—Smetana, 2004: 459 (distinct genus, list).

Comments. The genus Myrmoecia has not been well defined, and is considered to beclosely allied to Pella (Maruyama, in prep.). The members of the genus are somewhat similar to
those of Pella in general appearance, but they are easily distinguished from Pella species by themonilliform antennae, the narrower pronotum, and the pointed apex of the metasternal process.
Myrmoecia urartu (Iablokoff-Khnzoryan, 1962), n. comb.
Zyras (Pella) urartu Iablokoff-Khnzoryan, 1962: 111 (original description).—Smetana, 2004: 462 (“incertae sedis”, list).

Type material. Unfortunately, the type material was not available for my examnation. The
holotype is support to have been deposited in the Institute of Zoology, Laboratory of Entomology
National Academy of Sciences of the Republic of Armenia.
Type locality. Urartu fortress, near Nor-Aresh, Yerevan, Armenia.
Other material. Not examined.
Distribution. European subregion, Mediterranean subregion. EUROPE: Armenia.
Comments. In referring to the figure of the whole body in the original description(Iablokoff-Khnzoryan, 1962), I have concluded that this species is evidently a member of Myrmoecia and is closely allied to M. rigida (Erichson, 1839).
Bionomics. The type specimen of this species was collected from a desert zone of the Arax
riverside where no vegetation was found (Iablokoff-Khnzoryan, 1962).

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191

Tribe Athetini Casey, 1910
Athetae Casey, 1910: 2 [type genus: Atheta Thomson, 1858].

Diagnosis. The Athetini has not yet been well defined, but it can be distinguished from theother tribes of the Aleocharinae by a combination of the following character states: 1) head without occipital suture; 2) base of prementum with a projection postero-medially; 3) galea andlacinia shortened; 4) apex of the 1st segment of labial palpus obliquely truncated; 5) ligula without setae; 6) tarsi segmented 4–5–5; 7) ventral bridge of aedeagal median lobe (athetine bridge)
present.
Genus Dvorakatheta Maruyama, n.
Type species. Zyras (Pella) wrasei Dvořák, 1988.
Etymology. A combination of “Dvorak”, the author of the type species, and “Atheta”, thetype genus of the tribe Athetini. Gender feminine.
Diagnosis. Dvorakatheta may be differentiated from the other genera of the Athetini by a
combination of the following character states; 1) larger body (ca. 6.0 mm); 2) head much wider
than long, as wide as pronotum; 3) labial palpus widely separated; 4) 8th abdominal segment
widened posteriad in male; 5) aedeagal median lobe bifid at apex. The bifid apex of the aedeagal
median lobe suggests close relationship to the genus Nepalota Pace, 1987, but Dvorakatheta isclearly distinguished from it by the size and general shape of the body, the wider head, and the
8th abdominal segment widened in male.
Description. Body. Slender, subparallel-sided and flattened. Body length; 5.9–6.1 mm;
2.4–2.5 mm (from apex of clypeus to posterior margin of elytra). Colour dark brown to black.
Head. Head capsule (Fig. 139 A) large, much wider than long, as wide as pronotum. Occipital suture absent; surface covered with setae. Eyes oval in lateral view, slightly prominent, its
length 0.38 times as long as head width; small setae present among facets. Antennae filiform,
longer than combined head and pronotum, not flattened dorsoventrally; setae covering surface becoming denser and smaller apicad; 4th to 10th segments evidently dilated apicad, much longer
than wide, well margined postero-laterally; each segment with erecting black setae; 11th segment
elongate oval.
Mouthparts. Labrum much wider than long (W/L, 3.07), considerably emarginate anteromedially, submembranous around antero-medial area; surface covered with about 100 pseudopores except postero-lateral area; apodeme roundly projected postero-medially, with lateral arm
acutely curved and projected laterad. Epipharynx with medial sensory field with 40–50 pores andabout 70 grain-like sensory pores laterally; each mesolateral area randomly with 10–15 poresamong round projections; 5 pairs of lateral setae stout, about 10 times as long as wide. Mandibles
asymmetric; right mandible with a large tooth at middle of inner margin; left mandible moderately emarginate at middle of inner margin; 2 or 3 small setae present laterally; mesal areas of dorsal
surface with about 10 pseudopores mesally; ventral surface scattered with about 30 pseudoporesaround base; prostheca obtuse at apex, its inner margin densely pubescent. Maxilla: Cardo generalised, almost elliptical in ventral view, with about 80 pseudopores. Stipes large, triangular, with
2 setae at base, with about 5 pseudopores. Palpifer triangular in lateral view, with a large seta,
several small setae and 15–20 pseudopores near apex. Lacinia prominent at middle of inner margin, with 10 pores and 5 or 6 setae medially; apical inner margin with a low of setae forming acomb; surface around comb densely pubescent; subgalea small, much smaller than last segment

192

M. Maruyama

of maxillary palpus; galea long, about four times as long as wide, slightly curved, with somepores near apex, densely pubescent apically. Maxillary palpus without pseudopores; 1st segment
very small, with 1 seta; 2nd segment gently curved, and much dilated apically; 3rd segment thelongest, almost as long as 2nd, almost straight; 4th segment elongate, conical. Mentum (Fig.
140 A) trapeziform; posterior and anterior margins truncate; around antero-lateral corner with 1
seta; surface sparsely covered with pseudopores except in mesal and lateral areas, with somesetae laterally. Labium (Fig. 140 B): prementum (Fig. 140 B) with 2 real pores and 1 setal poremesolaterally, about 60 medial pseudopores and 15–20 mesolateral pseudopores; internal ridge(Fig. 141 B) entire, its length about 2/3 as long as prementum; apodeme (Fig. 140 B) with postero-medial margin roundly emarginate; lateral lobe of apodeme (Fig. 140 B) gently curved,
pointed apically. Ligula (Fig. 140 B) bilobed and each lobe rounded apically; apical setae absent,
but three sockets present. Labial palpus (Fig. 140 B) widely separated; 1st segment twice as longas 2nd, obliquely truncate at apex; 2nd the shortest. Hypopharynx (Fig. 140 B) without apicalseta; medial sensory field with small ridges around apex and scale-like sculptures near base.
Thorax. Pronotum (Fig. 139 A) subtrapeziform (PW/PL, 1.42); disc well margined, somewhat convex but more or less flattened above; surface densely covered with setae uniformly; hypomera visible in lateral view. Mesosternum with process long, pointed at apex. Metasternum
with process slightly prominent, its apex completely rounded. Elytra without epiplural suture,
margined from inner margin to posterior margin; surface almost uniformly covered with setae but
slightly denser apicad. Hind wing entire.
Legs. Coxae: fore coxa well developed, slightly shorter than fore femur, with suture ventrally; mid coxa generalised, with short suture ventrally; hind coxa subtriangular, rounded laterally. Trochanters: fore and mid trochanters small, subconical; hind trochanter rounded at apex.
Femora stout, flattened, gently curved near apex. Tibiae dilated apicad from base to around apical
1/4; each apical margin with ctenidium and 2 stout setae. Tarsi generalised; empodium with pairof setae.
Abdomen. Third to 6th segments: tergite with posterior margins almost straight; dorsolateral plates rectangular; sternite with surface densely covered with setae. Seventh segment withlateral plates fused, narrowed posteriad, pointed at apex. Eighth abdominal segment (Figs. 141 A,
141 B): tergite with basal suture curved laterally and continued apicad; sternite with basal suturereaching lateral sides. Ninth abdominal segment: tergite with surface densely covered with setae.
Tenth abdominal segment with surface densely covered with setae uniformly.
Male characters. Head depressed above. Eighth tergite (Fig. 141 A) widened apicad, withlateral projection of apodeme (Fig. 141 A) long; posterior margin (Fig. 141 A) rounded. Eighthsternite (Fig. 141 B) widened apicad; Lateral projections of apodeme of 9th tergite symmetrical;
Ninth sternite elongate oval; apical margin rounded; surface covered with setae and a pair ofmacrosetae near posterior margin. Aedeagus (Figs. 100 C–100 E): median lobe (Figs. 141 C,
141 D) bifid at apex; basal capsule somewhat bulbous; distal crests (Figs. 141 C, 142 D) fusedfrom side to side, large, much projected; copulatory piece (Fig. 141 E) very large, about 1/2 aslong as median lobe; paramere (Fig. 140 C) with condylite almost straight, as long as paramerite;
hinge zone distinct; apical lobe of paramerite short, curved; velum densely ridged.
Female characters. Female not examined. Eighth tergite narrowed posteriad, its apex
roundly emarginate (Dvořák, 1988).
Comments. The genus can be placed in the tribe Athetini by the following character statesin addition to the diagnosis of the tribe: 1) mesocoxal cavity narrowly separated; 2) metasternal
process very short, much shorter than mesosternal process. These character states are not ob-

Palearctic Species of Pella

193

served in the Lomechusini, in which the type species was formerly placed.
Dvorakatheta wrasei (Dvořák, 1988), n. comb.
(Figs. 139–141)
Zyras (Pella) wrasei Dvořák, 1988: 1. — Smetana, 2004: 461 (Zyras (Pella), list).

Type material. Paratypes: Tadjikistan: 2??, “USSR, Tadjikistan Varzob/Hissar/, 19. 7.
1984, 2000 m, D. W. Wrase legit.”/PARTYPUS/Zyras wrasei sp. n. det. M. Dvořák ,85 (ZMHB,
cDvor).
Type locality. Varzob, Tadjikistan.
Other material. Not examined.
Distribution. Meditrranean subregion. ASIA: Tadjikistan.
Diagnosis. Dvorakatheta wrasei may be easily distinguished from the other species of thetribe Athetini by a combination of the larger body and the larger head, which almost as wide as
pronotum. This species is somewhat similar in body size and general appearance to the membersof the subgenus Psammostiba Yosii & Sawada, 1976, of the genus Atheta but the head of the latter is generalised and much narrower than the pronotum.
Diagnosis. Body slender. Dark brown to black in ground colour; antennae, mouthparts,
pronotum, elytra, and legs paler. Head (Fig. 139 A) widest behind eyes; postocular part swollen

Fig. 139. Dvorakatheta wrasei (Dvořák). —A, Fore body, dorsal view; B, right antenna.

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M. Maruyama

Fig. 140. Dvorakatheta wrasei (Dvořák). — A, Mentum, ventral view; B, prementum, ventral view; C, paramereof aedeagus, lateral view.

in dorsal view; surface smooth, moderately covered with setae; setae moderate in length, as longas those on pronotum and elytra; length of eyes 0.26 times as long as head width. Antennae (Fig.
139 B) shorter than head, pronotum and elytra combined; 1st segment much shorter than 2nd and3rd combined; 2nd segment shorter than 3rd; 3rd segment about 2/3 as long as 1st; 4th to 10th
segments 2.1–2.6 times as long as wide; 11th segment shorter than 1st; approximate relative
lengths of segments from basal to apical: 32 : 20 : 24 : 19 : 20 : 20 : 18 : 17 : 17 : 15 : 23. Pronotum(Fig. 139 A) subtrapeziform, 1.42 times as wide as long, widest around anterior 1/6, narrowed
posteriorly; postero-lateral corner rounded; surface finely punctured, densely covered with setae.
Scutellum with surface smooth, moderately covered with setae. Elytra (Fig. 139 A) slightly

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195

Fig. 141. Dvorakatheta wrasei (Dvořák). — A, Eighth abdominal tergite, dorsal view; B, 8th abdominal sternite,
ventral view; C, median lobe of aedeagus, inner sac slightly evaginated, lateral view; D, ditto, ventral view; E,
ditto, apical part, postero-ventral view; F, copulatory piece, dorsal view. Male.

widened apicad; surface smooth, densely covered with setae. Legs very long; hind tibia 1.44
times as long as elytra; approximate relative lengths of tarsal segments from basal to apical: 5 :5.5 : 5.5 : 21 in fore tarsus; 7.5 : 8 : 9 : 10 : 18 in mid tarsus; 20 : 18 : 11 : 11 : 21 in hind tarsus. Abdomen subparallel-sided, slightly narrower than elytra, widest around 3rd and 4th segments; surface finely punctured; 3rd to 7th tergites densely and uniformly covered with setae; 8th tergite(Fig. 139 A) with 5 macrosetae; 9th tergite with 4 macrosetae; 10th tergite with posterior marginroundly prominent, with 5 macrosetae.

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Male: Eighth tergite (Fig. 141 A) with posterior margin slightly rounded; 8th sternite (Fig.
141 B) with posterior margin rounded, with 21–24 macrosetae; 9th sternite with posterior marginrounded. Aedeagus (Figs. 141 C, 141 D) elongate pear-shaped in ventral view, bifid at apex; apical lobe curved ventrad at middle, pointed at apex in lateral view; basal ridge thick and roundly
convex; copulatory piece of inner sac as in Fig. 141 E. Paramere with apical lobe of paramerite
dilated apicad, rounded at apex, about 1/8 as long as condylite.
Female: Not examined. Eighth tergite with posterior margin roundly emarginate (Dvořák,1988).
Measurements. BL, 5.7; FBL, 2.3; HL, 0.70; HW, 1.06; EL, 0.276; AL, 3.19; PL, 0.88;
PW, 1.08; ELL, 1.15; ELW, 1.55; HTL, 1.66.
Bionomics. No information.

CHAPTER 3.

BIONOMICS

Myrmecophily is one of very remarkable and interesting phenomena in adaptation of animals
and has been reported in numerous animal taxa, especially in the class Insecta. Among myrmecophilous insects, the majority of species are members of the Aleocharinae. In addition, theirmodes of life are diversified: scavengers, symphiles, predators, etc. However, very few myrmecophilous aleocharine life cycle and behaviour have been investigated in sufficient detail. To
know the evolutionary patterns of myrmecophilous aleocharines, descriptive data of their life cycles and behaviours are very important.
Hölldobler et al. (1981) already reported those of European species of Pella in detail, but inmy observation, those of the Japanese species are somewhat different from their report in somedetails, and I found some new information on the bionomics of the Pella beetles.
In this chapter, I am going to report life cycles and behaviours of several species of Pella (P.
socia, P. kidaorum, P. kinomurai, P. japonica, P. beijingorum, P. masakoae, P. spreta and P. indiscreta) based on my observations both in the field (Sapporo-shi, Hokkaidô) and in the laboratory
cultures.
Life Cycle
Life cycles in most species
In late April to early May, a large number of Pella beetles belong to P. socia, P. kidaorum, P.
kinomurai, P. beijingorum, P. masakoae and P. spreta were found at the entrance of Dendrolasius
nest in Sapporo. On warmer days in this period, beetles exhibited high flight activity in the laboratory. In the field, beetles were observed to have been attracted to a light trap, and also to have
copulated around the entrance to the nest.
In early May, I found the first eggs under the shelters around the field nest. Observations on
immature stages were confined to laboratory cultures. Larvae hatched about two days after theoviposition. Two or three days after the hatching, the larvae moulted into the second instar, andfurther three to four days later, they pupated. Five to seven days after the pupation, young beetles
eclosed from their pupae. The beetles stayed in their cocoons for seven to ten days, then theirbodies became gradually harder. In the field, the length of each stage may be longer than in thelaboratory as far as surmised from the appearance of young beetles.
In early June the mortality of overwintered adults in the laboratory increased. In the field, noPalearctic Species of Pella

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beetle was observed around the middle of June, and in late June to early July the first young beetles appeared above ground. They were weakly stained and not much sclerotized, with the abdomen not expanded as in overwintered adults.
In the field, on cold days (5–10°C) in late September, some beetles were observed under shelters around the nest and a few beetles walking around the nest. In early October, the number ofthe beetles found around the nest declined markedly and in late October no beetle was observed
outside the nest.
In November of 1999, I excavated a nest of Lasius (Dendrolasius) spathepus in Hitsujigaoka,
Sapporo-shi. I found many beetles within the nest entrance covered by nest material and in the
garbage dump accumulated around the nest entrance. Each beetle was in an elliptical space, withlegs squeezed together in “dormance position” expressed by Hölldobler et al. (1981). Apparently
these beetles were overwintering. Such beetles were also observed in the roots of trees, wheretrail of ants was observed on warmer days.
Pella japonica and P. indiscreta apparently have different life cycles in my field observations
as follows.
Pella japonica
Adults of Pella japonica appeared in early May to early June and actively copulated. However, they did not lay eggs in this period. My observation of the ovarium in several females showedthat the eggs did not grow in this season but fully matured in late August to early September. In
spite of my repeated trials to rear Pella japonica, I did not observe oviposition and obtained nolarvae. No newly eclosed or overwintering adults were found in my field investigations, but many
larvae were found in August. All this suggests that this species lays eggs in the summer and overwinters in the stage of larva or pupa.
Pella indiscreta
Newly eclosed adults of Pella indiscreta appeared in July, and matured eggs were observed intheir ovaria in late August to September. However, I did not observe oviposition and obtained nolarvae. Probably this species lays eggs in the autumn to just before the winter and overwinters at
the stage of egg, larva or pupa. It may pupate or eclose around late June to July.
Behaviour of Adults
General behaviour
Pella beetles are usually observed walking around the hosts’ nests but not observed withinnests. They sometimes prey upon host ants (see, Predatory behaviour), but they generally act as
scavengers, feeding on dead workers or garbage discarded around the nests. Although the Dendrolasius ants essentially feed on aphid or coccid honeydew, they also eat dead insects or earthworms when these are available. Occasionally, in the field, the beetles are observed eating deadcaterpillar or earthworm together with ant workers. The Pella beetles, therefore, can also be cleptoparasitic.
In the field, two patterns of daily rhythmic activity have been recognized among Pella
species. While Pella masakoae and P. spreta are apparently nocturnal, clustering under shelters,e.g., dead leaves or pebbles, in the daytime, P. comes, P. socia, P. kidaorum, P. kinomurai and P.
japonica are active both day and night.
In May to October, the main foraging season, Dendrolasius ants travel along trunk trails to

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aphid colonies. I saw Pella beetles walking along the trails. Although beetles were usually foundwithin a range of 10 m from the nest entrance, those of Pella comes, P. socia, P. kidaorum and P.
japonica were found at the trail more than 30 m distant from the nest. Akino (2002) verified, by abioassay, that P. comes can follow the trail pheromone of “Lasius fuliginosus” (Lasius fuji).
Mating and egg laying
I observed mating behaviour in all species that I reared in the laboratory culture, and confirmed that it was almost the same as in other aleocharines, i.e., “assault style” (Peschke, 1978).
The male beetle chases the female and inserts his aedeagus from the side no sooner than catching
up with her. Copulation usually takes a few minutes, but in Pella japonica it took about 10 minutes. The male beetle of Pella japonica is sometimes dragged by the female, inserting his aedeagus.
Eggs were laid under shelters (bark of dead tree) in the laboratory culture. I did not observe
oviposition behaviour but the females probably hid under the shelters and laid egg there. Theylaid 1 egg (rarely 2 eggs) at once and 6–8 eggs in total with an interval of two or three days. I observed the eggs in the field nest three times. They were laid under dead leaf or in the garbage
dump in front of the nest.
Predatory behaviour
According to Wasmann (1886, 1920, 1925), all species he had studied, Pella humeralis, P. funesta, P. cognata, P. similis, P. lugens and P. laticollis, prey on ants, especially disabled ants. In
contrast, Kolbe (1971) failed to find a predatory behaviour in Pella humeralis and concluded that
this species primarily fed on dead ants. Yasumatsu (1937) and Kistner (1972) made similar observations as Kolbe (1971) for Pella comes and P. japonica. However, Hölldobler et al. (1981)
observed Pella funesta, P. laticollis and P. humeralis living not only as scavengers, feeding ondead or disabled ants and debris discarded by the ants, but also as very effective predators on theants. In my field and laboratory observations, Pella comes, P. kidaorum and P. japonica also acted
as predators.
I frequently witnessed Pella beetles hunting Lasius fuji and L. spathepus workers on the foraging trail in the field. I saw the beetles hunting during the daytime on one occasion, but mostfrequently in the evening or at night.
The hunting behaviour is almost the same in Pella comes, P. kidaorum and P. japonica in myobservations and also agrees with the behaviour of P. laticollis, P. funesta and P. humeralis reported by Hölldobler et al. (1981).
When the hunting was successful, the beetles usually tried to drag the prey away from thetrail of ants to under dead leaves or dead branches on ground. Hölldobler et al. (1981) reported
that when beetles were starved for several days and were kept without foods, they occasionallychased each other and sometimes fed on the disabled.
Defence and appeasement behaviour
As reported by Hölldobler et al. (1981), when encountering an ant, the beetle flexes the abdomen even more strongly. This behaviour is frequently described in many myrmecophilous
staphylinids, especially aleocharines, and is generally considered as a defence response (Wasmann, 1886, 1920; Donisthorpe, 1927; Jordan, 1913; Koblick & Kistner 1965; Kolbe, 1971;
Hölldobler et al., 1981). It has been suggested that the beetles discharge secretions from their tergal glands when flexing the abdomen (Kistner & Blum, 1971).

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Kistner and Blum (1971) mentioned that Pella japonica and possibly also P. comes produced
citronellal in their tergal glands. In contrast, Hölldobler et al. (1981) analysed contents of tergal
glands of some European Pella species and noted that they “could not find any resemblance of
the Pella tergal gland secretions to the mandibular gland secretions of Lasius fuliginosus”. They
also found that the Pella secretions contained undecane, a hydrocarbon commonly found in the
Dufour’s glands of ants of the subfamily Formicinae, and surmised that the undecane was an
alarm pheromone of Lasius fuliginosus.
Behaviour of Larvae
In the field, the larvae are usually found near the entrances or in the garbage dumps of Dendrolasius nests. Larvae feeding on dead ants were frequently observed in the laboratory, andhatched larvae immediately start to walk around to search dead ant. When the larvae found adead ant, they grasped it using their mandibles, walked backwards and attempted to carry it
under a shelter or the nook of the rearing box. The larvae crushed their head into a slip between
abdominal segments of the dead ant and sucked its body fluid. Their abdomen expanded and became blackish with the ant’s body fluid. The larvae particularly preferred dead ants that wereplaced in room temperature for one or two days after death to fresh dead bodies.
Hölldobler et al. (1981) reported that when ants encountered Pella larvae, they usually attacked them. The defensive behaviour is similar to that of adults.
The matured second instar larvae burrowed into ground to a depth of 2–3 cm or hid immediately under shelter to make their silk cocoons before pupation.
Discussion
Life cycle
The life cycles of Pella socia, P. kidaorum, P. kinomurai, P. beijingorum, P. masakoae and P.
spreta almost agreed with the cycle of the European P. funesta reported by Hölldobler et al.
(1981), and may also be similar to the cycle of some other European species, P. humeralis, P. lugens and P. laticollis, described by some authors (Wasmann, 1886, 1920, 1925; Franc, 1992) andobserved by me in the field in southern Slovakia. Pella japonica and P. indiscreta are different
from all of them. Usually, five to eight species of Pella are observed in the same nest. For example, in Central Europe, Pella humeralis, P. funesta, P. cognata, P. lugens and P. laticollis (rarely,
also P. similis) have been collected from same ant nest, in Japan (Sapporo) P. kidaorum, P. socia,
P. kinomurai, P. japonica, P. beijingorum, P. masakoae, P. spreta and P. indiscreta, and in Beijing
P. jureceki, P. cooterorum, P. hlavaci, P. beijingorum and P. zhoui. Though any competition
among the species was not suggested in my field observations, their general behaviour were similar among them. Thus, the difference in life cycle is indicative of seasonal habitat segregation
among the species, at least among P. japonica, P. indiscreta and the others.
The period of each immature stage is remarkably short, i.e., egg (2 days); 1st instar larva(2–3 days); 2nd instar larva (3–4 days); pupa (5–7 days) in the laboratory culture. Therefore, inthe minimal length, it takes Pella beetles only 12 days to become adult from egg laying. Moreover, two instars were represented in the larval stage of all the observed Pella species. As far as
known three instars are generally represented in the larval stage of aleocharine beetles, e.g., Aleochara (Aleocharini), Atheta (Athetini) (Ashe & Watrous, 1984) and Phanerota (Homalotini)(Ashe, 1981), and the immature period is 20–25 days. The abbreviation of larval instars in Pella

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may be related with the short immature period. Although the immature period of most staphylinids have been unknown, this is considered to be very short example.
Behaviour
There are some reports on categorization of myrmecophily based on the behavioural information: Wheeler (1910b) and Kistner (1979) that include five and two kinds of categories, respectively.
The Pella beetles have several behavioural aspects. Observations revealed three food habitats
of Pella beetles: 1) scavenger feeding on dead host workers or ants’ garbage; 2) predator uponliving ants; 3) cleptoparasite eating ants’ booty.
By these behavioural data, Pella can be classified into “synecthrans” (persecuted guests) ofWheeler’s (1910b) categorisation. The “synecthrans” eat some part of their hosts’ colony (thecolony member, the brood, or the refuse) and are persecuted by the colony members when theyencountered.
In contrast, it is difficult to classify Pella beetle into neither “integrated species” nor “non-integrated species” in Kistner’s (1979) categorisation. To a stretch point, Pella beetles are rather“integrated species” because they are chemically integrated to ant society, e.g., the pheromone
following behaviour and the similarity of the cuticular profiles to the host ants (Akino, 2002).
However, Pella beetles have also aspects of “obligate predators” and “defensive types” in the category “non-integrated species”.

ACKNOWLEDGMENTS
This paper is derived from a part of my Ph.D. thesis entitled “Biosystematics of the myrmecophilous genus Pella (Coleoptera, Staphylinidae, Aleocharinae)”, accepted by the Graduate
School of Agriculture, Hokkaido University, Sapporo, in March 2003. From the original paper,
the chapters “ Phylogeny” and “Larval morphology” are omitted for further reviewal for publication.
I would like to express my cordial thanks to Prof. Masaaki Suwa, Assoc. Prof. Shin-ichi Akimoto, Asist. Prof. Kazunori Yoshizawa, and Prof. Emeritus Sadao Takagi (SEHU), Prof . Yutaka
Saito (Animal Ecology, Hokkaido University), and Assoc. Prof. Masahiro Ôhara (HUM) for theircritical reading of the earlier version of the manuscript and continuous guidance, and the otherstaff and students of SEHU for their encouragement since I entered Hokkaido University. I am
also indebted to Dr. Shuhei Nomura (NSMT), who helped me in various ways during my studies
in NSMT, and by Dr. Keiichi Matsuura (NSMT), who arranged the publication of this paper.
Thanks are also due to cooperation in the material examined and informations of the following
entomologists and museum curators: Dr. Kee-Jeong Ahn (CNU), Mr. Koji Arai [Koji Toyoda],
Mrs. Shiho Arai (Ranzan-machi, Saitama), Dr. Volker Assing (Hannover, Germany), Mr. Martin
Brendell (BMNH), Mr. Jonathan Cooter (Hereford, England), Dr. Giulio Cuccodoro (MHNG),
Mr. Anthony Davies (CNC), Dr. Thierry Deuve (MNHN), Mr. Miroslav Dvořák (Praha, Czech
Republic), Dr. Johannes Frisch (ZMHB), Mr. Hirofumi Fujimoto (Kagawa-ken, Japan), Dr.
Matthias Hartmann (NMEG), Mr. Kaoru Haga (Saitama-ken, Japan), Mr. Yasuhiko Hayashi(Hyôgo-ken, Japan), Mr. Yukihiko Hirano (Kanagawa-ken, Japan), Mr. Peter Hlaváč (Košice,
Slovakia), Mr. Shigehisa Hori (Hokkaido Historical Museum, Hokkaidô, Japan), Dr. Hideto
Hoshina (Faculty of Educatin, Fukui University, Fukui-ken, Japan), Mr. Noriaki Inahata (Hyôgo,

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Japan), Dr. Fuminori Itô (Kagawa University, Kagawa-ken, Japan), Mr. Katsuhiko Itô (Hokkaidô,Japan), Mr. Tateo Itô (Kyôto-fu, Japan), Dr. Josef Jelínek (NMP), Mr. Hiromu Kamezawa(Tokyo-to, Japan), Mr. Yuuki Kamite (Ôsaka-fu, Japan), Mr. Yasunari Kida (Maruseppu Insectarium, Hokkaidô, Japan), Mr. Kyoichi Kinomura (Gifu-ken, Japan), Dr. Toshio Kishimoto (Japan
Wildlife Research Centre, Tokyo-to, Japan), Dr. Takato Kobayashi (Yamanashi Institute of Environmental Science, Yamanashi-ken, Japan), Dr. Victor Kuznetsov (IBSS), Mr. Tomas Lackner(HUM), Dr. Kouichi Matsumoto (Tokyo-to, Japan), Dr. Ottó Merkl (HNHM), Mr. Kiminori
Miyashita (Hokkaidô, Japan), Mr. Koji Mizota (Miyagi University of Education, Miyagi-ken,
Japan), Mr. Hirofumi Moriya (SCM), Mr. Yoshiyuki Nagahata (Yamagata-ken, Japan), Mr. Seidai
Nagashima (Okayama-ken, Japan), Ms. Yoshie Nagayasu (HokkaidôHistorical Museum,
Hokkaidô, Japan), Dr. Shun-Ichiro Naomi (Natural History Museum and Institute, Chiba, Chibaken, Japan), Mr. Hiromichi Obata (Kanagawa-ken, Japan), Dr. Nobuo Ohbayashi (EEEU), Ms.
Ayuko Ohkawa (University of Tokyo, Furano Experimental Forest, Hokkaidô, Japan), Mr. Andrey Plutenko (Smolensk, Russia), Mr. Guilliame de Rougemont (Londinières, France), Mr.
Masahiro Sakai (EEEU), Mr. Masakazu Sano (SEHU), Mr. Kunihiko Sasaki (Hokkaidô, Japan),Dr. Harald Schillhammer (NHMW), Michael Schülke (Berlin, Germany), Mr. Yasutoshi Shibata(Tokyo-to, Japan), Mr. Takashi Shimada (Shimane-ken, Japan), Dr. Aleš Smetana (CNC), Mr. Hiroshi Sugaya (JAICA, Sabah, Malaysia), Mr. Kazuhiro Sugisima (SEHU), Ms. Azadeh Taghavian(MNHN), Mr. Yasushi Takai (Gifu-ken, Japan), Dr. Yuri Tshistjakov (IBSS), Dr. Manfred Uhlig(ZMHB), Mr. Bert Viklund (NHRS), Mr. Takashi Watanabe (Kanagawa-ken, Japan); Mr. Marengo Virgile (MHNL), Mrs. Yuriko Yamaga (Bihoro Museum, Bihoro-chô, Hokkaidô, Japan), Mr.
Nobuki Yasuda (Sounkyo Visitor Centre, Hokkaidô, Japan), Mr. Atsuhito Yoshida (Kanagawaken, Japan), Mr. Masataka Yoshida (Tokushima-ken, Japan), Dr. Hiroyuki Yoshitomi (Hokkaidô,Japan), Dr. Adriano Zanetti (MNV), Dr. Lothar Zerche (DEI), Dr. Hong-zhang Zhou (IZAS).
Last, not but least, I thank Dr. Shun-Ichi Uéno (NSMT) for his comments on the final version of
the manuscript, Dr. Tadashi Ishikawa (TUA) and Mr. H. Kamezawa for checking the first proof.
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