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ISSN 0968-0454

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18 JUN 1998

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The Bulletin of The Natural History Museum (formerly: Bulletin of the British
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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Ent.)

Entomology Series
ISSN 0968-0454 Vol. 67, No. 1, pp. 1-152

The Natural History Museum
Cromwell Road
London SW7 5BD Issued 25 June 1998

Typeset by Ann Buchan (Typesetters), Middlesex
Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset

Bull. nat. Hist. Mus. Lond. (Ent.) 67(1): 1-64 Issued 25 June 1998

Mealybugs of the genera
Eumyrmococcus Silvestri and
Xenococcus Silvestri associated
with the ant genus Acropyga Roger
and a review of the subfamily
Rhizoecinae (Hemiptera,
Coccoidea, Pseudococcidae)

THE NATURAL
HISTORY MUSEUM

18 JUN 1998

D.J. WILLIAMS__ GENERAL LIBRARY

Department of Entomology, The Natural History Museum, Cromwell Road, London

SW7 SBD.
CONTENTS
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SyNopsIS. The hypogeic mealybug genera Eumyrmococcus Silvestri and Xenococcus Silvestri,
their distribution throughout southern Asia and Australasia, and their association with the ant
genus Acropyga Roger are discussed. Eumyrmococcus is also discussed from Europe and South
Africa. Descriptions or redescriptions of 17 species of Eumyrmococcus are presented and
illustrated, including E. corinthiacus Williams, E. falciculosus sp. n., E. kolombangarae sp. n., E.
kruiensis sp. n., E. kusiacus sp. n., E. lamondicus sp. n., E. lanuginosus sp. n., E. maninjauensis
sp. n., E. neoguineensis sp. n., E. nipponensis Terayama, E. queenslandicus sp. n., E. recalvus sp.
n., E. sarawakensis sp. n., E. scorpioides (De Lotto), E. smithii Silvestri, E. sulawesicus sp. n. and
E. taylori sp. n.The genus Xenococcus includes two species, X. acropygae sp. n. andX. annandalei
Silvestri. It is now known that X. annandalei is a local species from a small area in India. The
female pupal instar, already known in Xenococcus, 1s also discussed inEumyrmococcus. Full page
illustrations for each species and keys to species of Eumyrmococcus and Xenococcus are provided.

D.J. WILLIAMS

The position of Eumyrmococcus and Xenococcus within the subfamily Rhizoecinae is discussed
and a review of all the genera included in the subfamily is provided. A new genus and species,
Leptorhizoecus deharvengi, is described from Sumatra and the generic name Radicoccus

Hambleton is synonymised with Rhizoecus.

INTRODUCTION

In a list of family-group names of the Coccoidea,
Williams (1969a) erected the tribe Rhizoecini, within
the mealybug family Pseudococcidae, for Rhizoecus
Kiinckel d’Herculais and a few related genera, all
inhabiting soil, leaf litter, rotting logs or feeding on
roots. Some of these genera are now known to be
identical with others and some other genera have been
added since. After a comprehensive study of the la-
bium of the Coccoidea, Koteja (1974a, 1974b) accepted
the group as the subfamily Rhizoecinae, mainly on the
basis that the labium is very narrow. This group also
lacks cerarii and the antennae are usually strongly
geniculate, with never more than 6 antennal segments,
the terminal segment always tapering and pointed. The
subfamily status has been accepted by most workers.
The other subfamilies Trabutininae, Pseudococcinae
and Sphaerococcinae, discussed by Koteja, have also
been accepted by many students of the Coccoidea
although the status of each subfamily is still ill-de-
fined.

The purpose of this work is to revise the genera
Eumyrmococcus and Xenococcus. A few species of
Eumyrmococcus had been studied already for the au-
thor’s current work on the mealybugs of southern Asia.
Some interesting collections, however, sent by Dr
R.W. Taylor, CSIRO, Canberra, from Australasia, as-
sociated with the ant genus Acropyga, have revealed
some remarkable species. Any attempt to publish on
southern Asian species without a knowledge of all the
others would be undesirable. It is for this reason that a
study is presented here of all the known species of
Eumyrmococcus, now totalling 17.

It is clear that most of the records of the related
genus Xenococcus have been based on a misconcep-
tion. The type species X. annandalei Silvestri, appears
to be a local species, at present living only on Barkuda
I., Orissa Province, India. Another species, previously
identified as X. annandalei from India, most of south-
ern Asia and Australasia, is different, and is here
described as new.

Eumyrmococcus and Xenococcus were also included
in the subfamily Rhizoecinae by Tang (1992) who
erected the tribe Xenococcini for them. According to
Tang, this tribe differs from the Rhizoecini, the only
other tribe, in lacking ostioles. The distinction may
hold for Chinese and neighbouring species but not in
the subfamily as a whole as known worldwide. It is
difficult to separate Eumyrmococcus and Xenococcus,
for instance, without reference to Neochavesia

Williams & Granara de Willink, a South American
genus. This study of Eumyrmococcus and Xenococcus
is followed, therefore, by a discussion of all genera of
the subfamily Rhizoecinae. Included genera accepted
here are Rhizoecus, Capitisetella Hambleton,
Pseudorhizoecus Green, Geococcus Green,
Leptorhizoecus gen. nov., Pygmaeococcus McKenzie,
Neochavesia, Eumyrmococcus and Xenococcus. A key
to these genera is also included.

Few specimens of Eumyrmococcus have been avail-
able for this study. Collecting these mealybugs is not
easy and acknowledgement is always due to the col-
lectors who have searched for them in ants’ nests or
have taken the trouble to extract them from soil sam-
ples. Species records represent isolated collections
over a wide area. All the species are minute and their
preparation on to microscope slides is often tedious.
Distinguishing adults from immatures preserved in
vials is almost impossible and it is often disappointing
to find only one or two adult females in any sample.
Even more disappointing is the complete absence of
adults.

The present study of Eumyrmococcus and
Xenococcus must be recognized as a preliminary iden-
tification guide and not a phylogenetic study. Some
possible species-groups can be distinguished and these
are discussed in the introduction to the section on
“Species and Genera’ (p. 7). Itis hoped that the present
study of these mealybugs and their ant relationships
will form the basis for a future cladistic study. Such a
study could also be extended to the other rhizoecine
genera discussed here, at least to some of the monotypic
genera which need further investigation.

MATERIAL AND METHODS

The slide-mounting techniques and the methods of
illustration described by Williams & Watson (1988)
and Williams & Granara de Willink (1992) are fol-
lowed. The terminology in these works is adopted here
except for the terms bitubular and tritubular pores. In
the present work they are referred to as bitubular
cerores and tritubular cerores, terms that were widely
used by Hambleton (1976) and adopted from
MacGillivray (1921).

Measurements of the body length and width are in
millimetres but all other measurements, even if larger
than the body length, are given in microns.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 3

ABBREVIATIONS OF INSTITUTIONS

The following acronyms are used throughout for the
depositories of specimens.

ANIC Australian National Insect Collection, Canberra,
Australia.

BMNH _ The Natural History Museum, London, UK.

IEAUN Istituto di Entomologia Agraria dell’ Universita di
Napoli, Portici, Italy.

MNHN- Muséum National d’ Histoire Naturelle, Paris,
France.

NIAES _ National Institute of Agro-Environmental

Science, Tsukuba-shi, Japan.
QM Queensland Museum, South Brisbane, Queens-
land, Australia.

SANC National Collection of Insects, Pretoria, South
Africa.

USNM _ National Museum of Natural History [Scale
insect Collection], Beltsville, Maryland, USA.

ACKNOWLEDGEMENTS

Much of the material discussed in this work was sent by Dr
R.W. Taylor, CSIRO, Canberra, Australia, as part of his ant
studies of southern Asia and Australasia. | am much indebted
to Dr Taylor for the opportunity of examining this material.
Daniele Matile-Ferrero, Muséum National d’Histoire
Naturelle, Paris, France, kindly sent all the mealybug material
extracted from soil samples collected by Drs Louis Deharveng
and Anne Bedos, Université Paul Sabatier, Toulouse. Dr D.R.
Miller, Systematic Entomology Laboratory, USDA, Beltsville,
Maryland, USA, arranged the loan of material from the US
National Museum of Natural History housed at Beltsville,
and has kindly spent time examining specimens and giving
useful information. Dr Mamoru Terayama, Department of
Biology, College of Arts and Sciences, University of Tokyo,
Japan, has generously sent material of Eumyrmococcus
nipponensis Terayama and E. smithii Silvestri used in the
descriptions for this work. lan M. Millar, Plant Protection
Research Institute, Pretoria, South Africa, has kindly sent for
study the holotype and paratypes of E. scorpioides (De Lotto).
Dr Penny J. Gullan, Australian National University, Can-
berra, Australia, read the entire draft manuscript, suggested
improvements and offered perceptive criticism, which I have
heeded throughout. I am much indebted to Dr Gullan for
spending her time and showing keen interest in this work.

HABIT AND ECONOMIC
IMPORTANCE

It has been been postulated by Koteja (1984, 1985) and
well summarised by Gullan & Kosztarab (1997) that
the ancestors of present-day Coccoidea lived on the
leaf-litter layer, feeding on dead and decaying matter,

or even on associated fungi and bacteria. Some soil-
inhabiting coccoids are, apparently, secondarily
inhabitants but Koteja suggests that Margarodes
Guilding (Margarodidae) and Rhizoecus
(Pseudococcidae) might have spent all of their
phylogeny in soil or litter. The underground habit of
Rhizoecus could not have been acquired secondarily
from an aerial habit on stems, twigs and leaves (Koteja,
1985).

Endosymbionts of scale insects are diverse and may
have been acquired via the alimentary canal during the
original leaf-litter habit. Mealybugs may have even
changed their feeding behaviour by living a symbiotic
relationship with ants (Tremblay, 1989) but the
endosymbionts of most mealybugs, although differ-
ent, are nevertheless related (Buchner, 1969). The
symbionts of Puto Signoret and Macrocerococcus
Leonardi are different from others in the
Pseudococcidae and those of Rastrococcus Ferris are
so distinct that Rastrococcus should be separated from
the Pseudococcidae (Buchner, 1957, 1969). Tremblay
(1977) recognizes two main symbiotic adaptations
among the pseudococcids, the Pseudococcus
Westwood type and the Puto type. No symbionts of
Rhizoecus and its close relatives have been studied but
Silvestri (1924) showed that Xenococcus has a
pseudococcid type of endosymbiosis. Buchner (1957,
1969) apparently studied symbionts of Eumyrmococcus
(without mentioning the species) and commented that
these are a pseudococcid type, near those of
Phenacoccus Cockerell.

Rhizoecinae live underground, mostly feeding on
rootlets. Many species have only been collected from
Berlese funnel apparatus but many are known from
leaf litter and rotting logs. The only aerial habit re-
ported is that of Rhizoecus mexicanus (Hambleton) on
leaves of Zygocactus truncatus; otherwise this
mealybug normally feeds on cactus roots (Hambleton,
1979):

Species of many genera of Rhizoecinae are associ-
ated with ants of the genus Acropyga Roger which
attend the mealybugs and feed on their excreted hon-
eydew. The association may be an old one because
some of these ant species are known to be obligate
coccidophiles and many of the mealybug species are
probably obligate myrmecophiles, although some
mealybug species reported with ants are capable of
living without them. Rhizoecus coffeae Laing is a pest
of coffee in Brazil where it lives on the roots in
association with Acropyga (Rhizomyrma)
paramibensis Borgmeier (Biinzli, 1935). The
mealybug is well protected from excessive change in
dryness and wetness caused by the weather because
the nests are well drained and the mealybugs are
capable of moving to suitable feeding sites. Accord-
ing to Biinzli, the situation is different from that of
the ant-free mealybugs. Flanders (1957) reported that

the myrmecophilous mealybug Eumyrmococcus (=
Neochavesia) sp. only feeds on roots exposed by the
ant A. (Rhizomyrma) fuhrmanni Forel and that the
number of mealybugs feeding at any one time is
regulated by the ant.

Ants associated with Eumyrmococcus and
Xenococcus are also species of Acropyga and these
mealybugs have been collected within the ants’ nests,
in ant tunnels, or from the mandibles of flying queens.
A few Eumyrmococcus species described here as new,
have been collected without ant data or simply labelled
‘with ants’ but it is reasonable to suggest that all these
mealybugs may be associated with Acropyga (see list
of Acropyga-Rhizoecinae associations p. 30).

When carrying mealybugs in the mandibles, ants
grip species of Rhizoecus and Geococcus at a point
near the metathorax (Biinzli, 1935). Ants apparently
grasp individuals of Eumyrmococcus at the dilated
cephalothorax (Roepke 1930; Prins 1982; Buschinger
et al., 1987).

Ants of the genus Acropyga are discussed here in the
four subgenera listed by Bolton (1995a). Some of the
unidentified species of Australasian Acropyga may be
undescribed if not discussed by Taylor and Brown
(1985) orTaylor (1992). Of the 56 species ofAcropyga,
about half occur in the Neotropical Region and the
remainder are distributed mainly in the Indo-Australa-
sian Region, with a few described from the Palaearctic,
Afrotropical and Oriental Regions (Bolton, 1995b).
This ant distribution appears to be correlated with the
number of mealybug species found in these regions. A
few non-rhizoecine mealybugs may be associated with
Acropyga in South America (Biinzli, 1935), and it
seems unlikely that species of Acropyga in other re-
gions will not exploit mealybug trophobionts in other
subfamilies.

Biinzli (1935) has listed R. coffeae, R. caladii
Green, R. moruliferus Green (= R. falcifer Kiinckel
d’Herculais), Geococcus coffeae Green, Pseudo-
rhizoecus proximus Green and Capitisetella migrans
(Green) as attacking coffee roots in Brazil. All the
indigenous host plants of these mealybugs
are grasses, and Biinzli has shown that the mealy-
bugs have migrated to the new host plant Coffea
liberica.

Other rhizoecine mealybugs associated with
Acropyga in SouthAmerica, attacking economic plants,
are Neochavesia caldasiae (Balachowsky) on roots of
coffee in Colombia, NV. eversi (Beardsley) on roots of
banana in Panama, and N. trinidadensis (Beardsley)
on roots of cacao in Trinidad.

In India, Dr C.A. Viraktamath, University of Agri-
cultural Sciences, Bangalore, has sent specimens of X.
acropygae sp. n., that were feeding on the rootlets of
grape, causing a severe reduction in yield. The
mealybug also attacked many weed species in the
vicinity of the vines.

D.J. WILLIAMS

LIFE CYCLES

In the family Pseudococcidae, there are usually four
instars in the female and five in the male. Normally the
female is regarded as neotenic and all instars feed. In
the male, feeding is restricted to the first two instars
followed by the non-feeding prepupa, pupa and adult.
Koteja (1985) suggests that loss of wings in the female
occurred only once and was an adaptation to the
original leaf-litter or hypogeic habitat. Normally adult
males are much smaller than the adult female. Koteja
(1985) hypothesizes that in the original leaf litter,
small males were able to crawl among soil particles to
reach the females. This small size could be realized
only by a cessation of feeding, i.e. at the end of the
second instar. In order for the males and females to
emerge simultaneously, the last male instars became
resting and non-feeding instars known as the prepupa
and pupa.

The life cycles of Rhizoecus and near relatives are
virtually unknown. Biinzli (1935) discusses four fe-
male instars in R. coffeae and three male instars but he
may have overlooked the pupal instars. Schmutterer
(1952) discusses four female instars in R. albidus
Goux in Germany and mentions nymphs and adults in
the male without further details.

The adult males in some of the Rhizoecus group are
wingless and morphologically degenerate but in other
species they are winged (Schmutterer, 1952; Beardsley,
1962). Surprisingly, winged males in slide collections
of Rhizoecus are not uncommon (D.R. Miller, per-
sonal communication).

The first record of a female pupal instar was de-
scribed by Williams (1988) for the genus Xenococcus.
This instar replaces the normal third or feeding instar
and probably acts as a resting stage for the female to
develop in the presence of attending ants. A similar
type of pupa has now been found and is discussed here
in at least five species of Eumyrmococcus.

Unfortunately, no first-instar nymphs of Eumyrmo-
coccus are known from any of the material at hand
except for a single specimen still within the egg mem-
brane. This first-instar nymph is densely covered in
short setae and shows unusual developmemt of many
characters as in the first instar of Xenococcus (Fig. 24).
Second-instar nymphs are available in 11 species of
Eumyrmococcus but these remain unsexed. Normally
second-instar mealybugs show strong dimorphism,
with the second-instar male usually possessing more
tubular ducts and pores to form a waxy covering for the
prepupa and pupa. These characters are usually absent
or fewer in the second-instar female. In
Eumyrmococcus, the female pupal instar and the male
prepupal and pupal instars are without any coverings
and any sexual differences in the second instar are not
apparent.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 5

Female pupae show a well developed labium and
this is evidenced by at least four species of female
pupae with the developing adult female still inside
(Fig. 1B). Itis thus easy to separate female pupae from
male prepupae and pupae which lack the well devel-
oped labium (Figs 21, 25). Male prepupae are available
with the developing pupae still inside and pupae are at
hand with the developing adult male inside just before
the final moult (Fig. 1A). All female pupae, and male
prepupae and pupae, lack dermal structures such as
pores and setae. Unlike most male pupae in the
Pseudococcidae, which usually possess longer limbs
and antennae than those of the prepupae, these charac-
ters in the pupa of Xenococcus are shorter, presumably
in preparation for the adult male which possesses
minute tubercle-like antennae and short legs.

Studies of the adult male of Rhizoecus are remark-
ably few. Beardsley (1962) described the adult male of
R. falcifer in detail and showed that the head bears
only two pairs of eyes and lacks the normal lateral
ocelli. Furthermore, the head is not separated from the
thorax by a constricted neck. The penial sheath, ac-
cording to Beardsley, is related to that of Phenacoccus
Cockerell. Similar features are present in the male of
R. albidus as shown by Schmutterer (1952).

The adult male of Neochavesia eversi (Beardsley) is
morphologically degenerate, without wings, eyes or
thoracic sclerotization, and the penial sheath and
aedeagus resemble those of Puto (Beardsley, 1970).
Adult males of three species of Eumyrmococcus are
available, one of which is not described here because
no adult females were collected. The males are dispa-
rate. All are without eyes, wings and thoracic
sclerotization and possibly resemble males of
Neochavesia. The genital capsule in the Australian
species studied, possesses a long dorsal style and a
long pointed aedeagus. It is not clear why there should
be such development when the tip of the female abdo-
men is normally curled dorsally, presenting easy access.
Adult males of E. sarawakensis sp. n. are elongate,
curled ventrally, with most of the genital capsule
internal. Both the male and female in any one species
of Eumyrmococcus are about the same size.

Peculiar development in the adult male of
Xenococcus is described on p. 24. The legs are placed
at the anterior end of the body and the claws have the
strangest development known in any scale insect, with
apparent digitules larger and stouter than the actual
claw.

PARASITISM

Of about 160 species described in the subfamily
Rhizoecinae, only a single record of a parasitoid has
been published. Schmutterer (1952) reported a very

effective chalcid parasitising R. albidus Goux in Ger-
many. The parasitoid was recorded later as
Anomalicornis tenuicornia Mercet (Hymenoptera:
Encyrtidae) by Ferriere (1956), a fairly common Old
World species.

The Encyrtidae have successively parasitised the
other three mealybug subfamilies Trabutininae,
Pseudococcinae and Sphaerococcinae recognized by
Koteja (1974a, 1974b) as evidenced by the host
parasitoid index of encyrtid parasitoids of mealybugs
in Noyes and Hayat (1994). The views of Koteja
(1984, 1985) that Rhizoecus, which we may substitute
as the Rhizoecinae, could have spent the whole of their
phylogeny underground, may be a reason for the lack
of parasitism. Many of the mealybugs in the other
three subfamilies live underground or are concealed
and are known to be parasitised. The hypogeic habit is,
therefore, not a barrier. Many Rhizoecinae have a
symbiotic relationship with ants which could success-
fully exclude parasitoids but conversely many
Rhizoecinae are not associated with ants. Parasitoids
may not recognize rhizoecine mealybugs as suitable
hosts. J.W. Beardsley (personal communication) has
commented that the Rhizoecinae may have branched
off from the main mealybug lineage before the evolu-
tion of the Encyrtidae and may not be recognized as
normal encyrtid hosts. It is possible, however, that
parasitoids have simply not been collected from the
Rhizoecinae.

MORPHOLOGY OF
EUMYRMOCOCCUS AND
XENOCOCCUS (ADULT FEMALES)

Bopy. All descriptions of the adult female in life
indicate that the cephalothorax is strongly dilated, or
shows some sign of dilation, with a narrow tapering
abdomen which curls dorsally. When prepared on
microscope slides, the anal lobes are poorly developed
and are recognizable by infolding on both the dorsum
and venter with the apex of each lobe bearing three
long setae, collectively referred to here as anal lobe
setae. In E. neoguineensis sp. n., each lobe has 8 anal
lobe setae and in E. /anuginosus sp. n., the anal lobe
setae are not differentiated from other setae on the anal
lobes. Usually there are 3 anal lobe setae that are long
and stout at the bases with wide setal collars. They are
similar to those in many other genera of the
Rhizoecinae. In Eumyrmococcus and Xenococcus,
there are usually one ventral and two dorsal setae on
each anal lobe but occasionally two are placed ven-
trally and one dorsally.

ANAL RING. The anal opening projects slightly be-
tween the anal lobes and the anal ring is normally

simple with a crescentic sclerotized band without
cells. Three pairs of anal ring setae appear to be usual
but occasionally there are 4~7 pairs. The anal ring
setae are normally detached from the anal ring, lying
just anterior to it except in E. corinthiacus Williams,
which possesses 4 pairs, apparently attached to the
outer edge of the ring. In all species the anterior pair of
setae are short and slender, the second pair either the
same length as the anterior pair or longer, and the
posterior pair often long and stout, sometimes as long
as the anal lobe setae.

ANTENNAE. The antennae are 2—4-segmented and
differ in shape and position. In E. lanuginosus, they
are 2-segmented, short and tubercle-like. In most spe-
cies of Eumyrmococcus the antennae are 2-segmented,
with a short basal segment and a long tapering second
segment. They are placed fairly wide apart on either
the ventral or dorsal margin of the head. In E.
sarawakensis sp. n. and E. sulawesicus sp. n., the
antennae are situated well on the dorsum of the swol-
len cephalothorax. The 4-segmented antennae of E.
neoguineensis are unusual in possessing spine-like
setae on the leading edge. Xenococcus has 4-seg-
mented antennae that are long, sometimes as long as
the body, with strong articulation between the first and
second segments. The second segment is provided
with small points at the proximal end which fit into
grooves at the distal end of the first segment. This
articulation allows the antennae to fold against the
dorsum of the body. A pair of peg-like setae are present
at the tip of each fourth segment.

LeGs. The legs are always well developed with few
setae. The setae are stout and flagellate but sometimes
a few on the outer edge of the femur in Eumyrmococcus
are blunt and sensory. In Xenococcus, the leg setae
may be unusually long on the distal end of the femur,
almost as long as the femur. In most species the tibia is
swollen distally before narrowing towards the tarsus.
The tarsus is usually swollen near the base then tapers
gradually, sometimes for nearly half its length. In one
species of Xenococcus, the tibia and proximal end of
the tarsus are straight. The claw in E. scorpioides (De
Lotto) is broad but in all other species it is long,
pointed and slender with a pair of short setose digitules.
These digitules are difficult to discern in some species
except for the bases. A campaniform sensillum is
present at the base of all tarsi.

Eyes. These are completely absent in all species of
Eumyrmococcus and Xenococcus.

LABIUM. Inmost species the labium is narrow with 3
segments, often longer than the clypeolabral shield.
The setae on the anterior surface of the apical segment
are often widely spaced and the subapical setae are
sometimes reduced to 3 pairs. A few species possess 2
pairs of setae on the clypeolabral shield.

D.J. WILLIAMS

CEPHALIC PLATE. This sclerotized area on the frons
in many species of Rhizoecus is not apparent in any
species of Eumyrmococcus or Xenococcus.

CIRCULI. The type species of Eumyrmococcus, E.
smithii Silvestri, is the only species studied without a
circulus. In other species the circuli number 1-3,
situated within the borders of the segments. When
only a single circulus is present it is situated either on
abdominal segment II or III. When 2 are present they
lie on abdominal segments II and III, and a third in
Eumyrmococcus is sometimes present on the
metathorax. In Xenococcus, when a third is present, it
lies on abdominal segment IV. Each circulus is round
and may be small (the smallest studied only 8.75 um in
diameter) or unusually large (up to 85.0 um in diam-
eter). The outer edge of each circulus may be flat or
raised slightly from the surrounding derm. In one
species the circulus is almost conical but the centre in
all circuli is cupped, sometimes deeply in
Eumyrmococcus, or shallowly in Xenococcus, so that
the circulus resembles a crater. A similar type of
circulus is present in Neochavesia, and Beardsley
(1970) has cautiously termed these ‘circulus-like pro-
jections’. The form of the circulus in all these genera is
different from that of other genera in the Rhizoecinae
which possess circuli that are usually truncate-conical
with the centre part flat and often reticulate or faveo-
late.

A function of the circulus as an adhesive organ has
been demonstrated for Pseudococcus adonidum (L.)
(=P. longispinus (Targioni Tozzetti)) and Planococcus
citri (Risso) by Pesson (1939) and in other species by
Lloyd and Martini (1957). Ferris & Murdock (1936),
from histological sections, discussed the circulus in
Pseudococcus maritimus (Ehrhorn) as a secretory or-
gan. Pesson (1939), in a detailed study, found that the
circulus of Pseudococcus 1s an exsertile organ, lacking
secretory pores between the organ and the cuticle and
that the epithelium is only a part of the hypoderm. The
epithelium is responsible for the formation of the
peripheral chitin, which protects it, and of a mucin
which hardens after secretion. In the circulus, the
substance secreted appears to be a chitin which re-
mains soft longer with the gluey properties of a mucin
and hence gives the circulus adhesive properties. It is
easy to see the exsertile action of the circulus, accord-
ing to Pesson, by placing the mealybug on a microscope
slide so that it has difficulty in taking hold with its
claws. The mealybug can then hold on to the surface
by the circulus.

It is not clear 1f the circulus of Eumyrmococcus and
Xenococcus has the same function as those studied
above. Silvestri (1924) studied histological sections of
the circulus of X. annandalei and concluded that there
are numerous elongate-pyriform cells directly under
the shallow central cup. Silvestri thought that the

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI il

circulus secretes some kind of liquid which may be
attractive to the ants attending the mealybugs. Any
exsertile properties of the circulus in Eumyrmococcus
and Xenococcus, or even in any Rhizoecinae, remain
unproved. The term circulus is retained for the species
under discussion.

OSTIOLES. These structures are always absent in
Eumyrmococcus, Xenococcus and Neochavesia but
are usually well defined in some other genera of the
Rhizoecinae.

SETAE. One of the striking characters of Eumyr-
mococcus and Xenococcus is the abundance of short
setae on the dorsum of the abdomen. They extend to
the venter of the abdomen in Eumyrmococcus and may
be present on the head and thorax of both surfaces. In
Xenococcus, they extend to the head and thorax of the
dorsum and to the ventral lateral margins of the head
and thorax. They lie in well defined bands across the
segments leaving clear areas on the intersegmental
lines. Normally these setae are flagellate, often slightly
curved, but in E. falciculosus sp. n., they are mostly
sickle-shaped, and in Xenococcus, there are patches of
sickle-shaped setae among the flagellate setae on the
dorsum of the thorax.

Stout and longer flagellate setae are usually present
on the mid-venter of the thorax. In Xenococcus, these
extend to the entire venter of the abdomen also. Sen-
sory setae are sometimes present in Eumyrmococcus.
These are slightly swollen at the apices and may be the
only setae present on the head and thorax in some
species. In other species, they are either long or short
and sometimes mingle with the short, flagellate setae.
The sensory setae may also be lanceolate or extremely
minute and can be detected with certainty only by oil-
immersion studies. Some short setae in E.
neoguineensis are set in dermal pockets, forming clus-
ters on the venter of the thorax.

MICROTRICHIA. Elongate microtrichia, resembling
setae, are so abundant in some species that they almost
completely cover the insect so that the setae are ob-
scured. The setae in these species are only recognizable
by the presence of setal collars. At present, species
with dense microtrichia are only known from the more
equatorial areas, Sumatra, Sarawak and Sulawesi.

PORES AND DUCTS. Some genera of Rhizoecinae
contain species possessing trilocular pores, multilocu-
lar disc pores and tubular ducts. Normally, trilocular
pores secrete fine wax to protect the surface from
contamination by the insect’s own honeydew. Ex-
creted honeydew balls are often coated with this wax.
Multilocular disc pores secrete wax to help in the
production of ovisacs or to protect eggs. They are
sometimes present in second-instar males to secrete
wax in the formation of the cocoon (Cox and Pearce,
1983).

Tubular ducts in the Rhizoecinae are usually small,
often narrower than the trilocular pores. Normally,
tubular ducts in the adult female secrete wax to form
the ovisac, or in the second-instar male, to help forma
cocoon.

Pores and ducts are completely absent in
Eumyrmococcus and Xenococcus. Honeydew excreted
is probably taken entirely by the ants without need for
the mealybugs to secrete wax as an anti-contaminent.
Apparently the species are ovoviviparous and first-
instar nymphs would be protected and transported by
ants along with any other instars. Second instars lack
pores and ducts and succeeding male prepupae and
pupae are not enclosed in cocoons. The dense covering
of short setae on the abdomen of all species and the
complete covering of microtrichia in some species
may help to protect the mealybugs from any possible
contamination of honeydew and from excessive hu-
midity in the nests and tunnels formed by the ants.

In Neochavesia, trilocular pores are found in the
type species only but these pores are unlike any other
trilocular pores in possessing a minute filament at the
centre. Normal trilocular pores and tubular ducts are
present in Leptorhizoecus gen. nov. (Fig. 28).

GENERA AND SPECIES

Eumyrmococcus and Xenococcus are closely related
and it could be argued that Xenococcus is a subordi-
nate clade withinEumyrmococcus. Both genera occupy
similar geographical areas in southern Asia and Aus-
tralasia (Figs 29, 30) and the species in both genera are
mostly associated with specific ants of the same genus
(p. 30). Xenococcus possesses unusual 4-segmented
antennae, almost as long as the body, with remarkable
articulationn between the first and second segments.
In Eumyrmococcus, the antennae are much shorter and
lack this special articulation. Furthermore, all species
of Eumyrmococcus have a dense covering of short
setae on both the dorsum and venter of the abdomen.
In Xenococcus, short setae are restricted to the dorsum
of the abdomen only and any ventral abdominal setae
on the abdomen are long and stout. Although both
genera have many shared characters, the genera are
regarded here as distinct, in the interests of nomen-
clatural stability, pending future cladistic studies. A
key to separate both genera is included in the general
key on p. 30.

Some species of Eumyrmococcus share a large type
of circulus, others a small type, and two species lack
any type. Although it may be possible to recognise
species-groups based on the type of circulus, the circu-
lus is not a stable character in some other genera of
Rhizoecinae when any single species may possess 0—
2 (Williams, 1996).

Species-groups could be recognised on other char-
acters. The E. smithii-group contains species with
abundant short flagellate setae covering the dorsum
and venter and lacking any sensory setae. In addition
to the type species, E. smithii, this group also contains
E. kolombangarae and E. kusiacus. Another group,
found near the equator, comprises E. kruiensis, E.
lanuginosus, E. maninjauensis, E. sarawakensis and
E. sulawesicus. All are densely covered in elongate
microtrichia. In possessing similar 4-segmented an-
tennae, E. corinthiacus in Europe, is probably related
to E. scorpioides from South Africa even though the
claws in E. scorpioides are stout and different from the
long slender claws in any other species. The Australian
species, E. lamondicus, E. queenslandicus, E. recalvus
and E. taylori appear to be related to the Japanese
species, E. nipponensis. All possess similar 2-seg-
mented antennae, with the second segment elongate
and tapering, and with elongate posterior setae on the
anal ring.

Most of the abundant abdominal setae in E.
falciculosus are sickle-shaped, unlike the setae in any
other species, although similar setae are present in
small numbers in both species of Xenococcus. A final
species, E. neoguineensis, has an unusual combination
of characters. The 4-segmented antennae are the long-
est of any species and, although the articulation between
the first and second segments is normal, these anten-
nae possibly come nearest in structure to those of
Xenococcus. Furthermore, the anal lobes in E.
neoguineensis contain numerous setae and the body
shape is unusual, without such a marked dilated
cephalothorax. This species may form a distinct group.

The two adult males described here in
Eumyrmococcus are so disparate that if discovered
without accompanying adult females, it would be
difficult to place them in the same genus. The differ-
ence between these adult males is as great as the
difference between either and the adult male of
Xenococcus acropygae. Similar differences in male
morphology are expected among the different species-
groups of Ewmyrmococcus discussed here.

Eumyrmococcus Silvestri
Eumyrmococcus Silvestri, 1926: 271.

Type species Eumyrmococcus smithii Silvestri, by
original designation.

DESCRIPTION

Adult female. Body pyriform, broadly pyriform, or
elongate-pyriform, with cephalothorax usually strongly
dilated, often constricted between second and third
abdominal segments, then abdomen tapering to nar-
row abdominal segment VIII. In life, tip of abdomen
curled dorsally. Posterior end rounded, anal lobes not
developed but recognizable by inner edges slightly

D.J. WILLIAMS

grooved, each lobe usually terminating in long, stout
flagellate setae forming a group of 3, sometimes nearly
as long as body, sometimes stout and short or in groups
of numerous slender flagellate setae. Antennae each
with 2-4 segments, placed fairly wide apart on venter
of head margin or displaced to dorsum of head when
prepared on microscope slides; terminal segment usu-
ally tapering, sometimes tubercle-like or long and
slender. Legs well developed, tibia and tarsus subequal
in length, tarsus swollen from joint with tibia then
tapering to narrow distal end, all segments with fairly
stout flagellate setae, set well apart; occasionally with
sensory setae on outer edges of femur and tibia. Claw
normally long and slender, pointed, except in one
species with claw relatively stout at base; each with a
pair of short, slender setose digitules but these difficult
to recognise in some species. Labium longer than
wide, often with 3 pairs of subapical setae, widely
spaced. Clypeolabral shield with 1 or 2 pairs of setae.
Anal ring dorsal, sclerotized, usually crescentic except
in one species when circular; simple, without cells;
with normally 3 pairs of setae; first 2 pairs usually
detached, lying just anterior to ring, second pair some-
times longer than anterior pair, posterior pair sometimes
stout and as long as anal lobe setae; sometimes ring
with 4~7 pairs of setae, variously placed. Circuli present
or absent; when present numbering 1-3, all round,
small to unusually large and conspicuous, always
deeply cupped.

Body setae short and abundant, at least on abdomen,
sometimes present on entire body, usually flagellate
but sometimes most abdominal setae replaced by sickle-
shaped setae.

Setae on head and thorax often longer and stouter.
Sensory setae, either lanceolate or with slightly swol-
len blunt tip, sometimes distributed over entire body
with the flagellate setae, or situated at anterior end of
body only: occasionally minute bulbous setae present.
Microtrichia sometimes present, covering entire body,
these often nearly as long as setae giving the insect a
woolly appearance and often masking the short body
setae. Ostioles absent. Tubular ducts, multilocular disc
pores and trilocular pores absent.

First instar. Well clothed in short setae as evi-
denced by single specimen still within egg membrane.

Second instar. Similar to adult female but with
fewer setae and usually shorter antennae and legs.

Female pupa. Recognizable by elongate labium,
antennae showing segmentation, legs well developed.

Male prepupa and pupa. Similar to female pupa
but with shorter limbs and antennae. Labium not de-
veloped.

Adult male. Wingless and morphologically degen-
erate, of diverse appearance. Genital capsule well
developed, either exposed or mostly enclosed within
abdomen; when exposed, with conspicuous anal open-
ing and dorsal style, penial sheath rounded apically,

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 9

ventral slit rounded near base then widening apically,
basal ridge of penial sheath well defined ventrally,
aedeagus long and pointed. Posterior edge of abdomi-
nal segment VIII forming possible anal ring as in
female, with 3 pairs of setae. Legs well developed,
slender. Body setae numerous. Antennae 2-segmented.
Eyes absent.When genital capsule mostly enclosed
within abdomen then ventral slit almost square, anal
opening oval at posterior end of abdomen at base of
sclerotized extension, possibly representing a short
style. Legs squat. Antennae short, 2-segmented. Body
almost naked except for a few minute setae.

COMMENTS.

The description of the adult male is

based on two species and another still within the pupal
instar. The wide disparity in shape and form may
indicate a much wider range of variation.

Key to Species of Eumyrmococcus (Adult

Females)
PAIL AG SEPINIENLCH 5---.--.-..-.-ccorecceecceanssrargrececcerersers 2
SOILED UAG 2 -SEPIMENLEa..:<s<aceis.-a-bervsenequerexe<nskspuwonentebss 4
2 Anal lobe setae in groups of 3.Anal ring with 6-8 pairs of

QELS, = cotdcaned ree ROLE EPEC EEE ALE EERE EERE CELE Pen ret ae ar ee 3

Anal lobe setae in groups of 8 or 9. Anal ring with 10
pairs of setae (Fig. 10) ................ neoguineensis (p. 14)

All setae on head, thorax and abdomen, minute and
flagellate. Claw elongate, slender (Fig. 2) .............::000+
stontan <ec SR RAR ORR Eee corinthiacus (p. 10)

Most setae on head and thorax sensory, blunt, expanded
apically, only abdominal setae flagellate. Claw stout and
ISHN (EM SO) ca an aaruasioxsaisse Pap ox saicvans al scorpioides (p. 19)

Surface of body densely covered in minute hair-like
microtrichia in addition to abundant slender setae ...... 5

Surface of body not densely covered in minute hair-like
microtrichia. Abundant setae present only, although there
may be a few stout microtrichia present in some areas

Anal lobe setae not differentiated from other setae on
anal lobes. Antennae each with segment 2 short, scarcely
longer than segment | (Fig. 8)........ lanuginosus (p. 13)

Anal lobe setae in groups of 3 on each anal lobe. Anten-
nae each with second segment noticeably longer than
ISTE: CSL Ta Re ee enon Oe Re ae 6

Anal lobe setae and posterior anal ring setae short, about
70-75 \m long. All setae flagellate, blunt sensory setae
SO tat (EU GD) (2 Fa teaches Sarsteeceeechcn se cess kruiensis (p. 11)

Anal lobe setae and posterior anal ring setae much longer,
each 220-525 um long. Blunt sensory setae present, at
least on head, thorax and outer edges of legs .............. y

Antennae distinctly placed dorsally on microscope prepara-
tions. Blunt sensory setae present on dorsum of head and
thorax in addition to short minute flagellate setae ...... .. 8

Antennae distinctly placed ventrally on microscope prepa-

rations. Blunt sensory setae absent from dorsum of head
and thorax, only short minute flagellate setae present
(Bigs 9) seca Reece maninjauensis (p. 14)

Long, blunt sensory setae, at least 17.5—22.5 um long,
present on dorsum anterior to antennae only, near to head
margin. Shorter blunt setae present on dorsum posterior
to antennae on thorax and abdominal segment I (Fig. 14)
Seda ge CEPA VFO CER UGS EC AEE EES sarawakensis (p. 18)

Long blunt sensory setae, at least 45—50 um long, more
numerous on dorsum, present anterior and posterior to
antennae. Shorter blunt setae absent from dorsum on
thorax and abdominal segment I (Fig. 18).............008
Bs seed ac secrets nan diG ee eee ean Sines Si sulawesicus (p. 20)

Most setae on abdomen short, stout and sickle-shaped
(Rie) z eevee AR: ee. dates falciculosus (p. 10)

Most setae on abdomen short and flagellate, sickle-
SHAPE! SETAC: ADSCM LE) wcxccenccurescnddvcesscescsccxsconesicaseteaeores 10

Blunt sensory setae present, in addition to dense, short
Mapellatesetaey 5 sete sees ens, eles) es 13

Blunt sensory setae absent, dense, short flagellate setae
PRCSEDM OMY race ccare-tocde user ceatesanscerstnssehieveverscesesesteraccs I]

Circuli absent. Anal lobe setae and posterior anal ring
setae long, at least S00 um long (Fig. 17) «0.0.0... eee
BE Pe Ree eect eee eee ee smithii (p. 19)

Circuli present, small, on abdominal segments II and III.
Anal lobe setae and posterior anal ring setae much shorter,
AUIMOST GSMO O MMO Sieve. -t sceccecenenerreceieverecssnanaesees 12

Long, stout flagellate setae, at least 80-90 Lm long,
present in rows at posterior edges of dorsal and ventral
abdominal segments, these in addition to abundant minute
flapellate Setae (E10. ©) ccn.-cscccaneeede caren kusiacus (p. 12)

Long, stout flagellate setae shorter, at most 35 um long,
present medially on venter of abdomen only, in addition
to abundant minute flagellate setae (Fig. 4) .......0....0.
es SOE RS See? Se kolombangarae (p. 11)

Most sensory setae lanceolateniic.0:icreesvesesstecsess-e: 14
All sensory setae blunt and expanded apically .......... 15

A single circulus present only, on abdominal segment III,
wide and conspicuous, at least 52-60 [tm wide. Slender,
blunt sensory setae present on dorsum and venter of
El oral ovaunt atalllis(ern ant syan iil Ul (PSS 102) Weeperceerereee earn cere cecocceecee:
co eaoemntenpeerets penile et aster he queenslandicus (p. 16)

Two circuli present, these on abdominal segments II and
III, small, at most 11.0-12.5 um wide. Slender, blunt
sensory setae absent from abdominal segment II (Fig. 7)
Be yest UU Ek Re ta te ch areca lamondicus (p. 12)

Short slender setae abundant on head and thorax. Obanal
setae stout and long, about as long as anal lobe setae. A
single circulus present, this on abdominal segment III
(iiss Ul) at Bee fet or tees nipponensis (p. 15)

Short slender setae sparse on head and thorax. Obanal
setae short and slender. Two circuli present, these on
abdominal segments II and III ..........eeeeeeeeeeeeeeeeeees 16

16 Dorsal setae on head and thorax all sensory, of various
Sizes) (ig sil) pee. Seen ees recalvus (p. 17)

— Dorsal setae on head and thorax mostly long and flagel-
late, sensory setae few and minute (Fig. 19) .................
Stith 2 ee ee ee taylori (p. 20)

SPECIES DESCRIPTIONS

Eumyrmococcus corinthiacus Williams

(Fig. 2)

Eumyrmococcus corinthiacus Williams, 1993: 218.
Holotype adult, Greece, Corinth (BMNH) [exam-
ined].

DISTRIBUTION

Greece: Corinth, Perachora, near Lautraki, carried by
swarming ant Acropyga sp., 7.x.1985 (A. Buschinger);
without locality, carried by queens of Acropyga sp.
during mating flight, x. 1992 (W.H.O. Dorow).

COMMENTS. This species was described recently by
Williams (1993). Important characters are the 4-seg-
mented antennae and thick setae in groups of 3 on the
anal lobes, 2 in each group distinctly longer and
stouter than the other. The dorsal and ventral setae are
abundant, all flagellate, becoming less numerous on
the thorax and head. As in other species, the circulus is
concave and cup-shaped when viewed laterally.

The original material, the first known from Europe,
was sent for identification by ProfessorA. Buschinger,
Institut fiir Zoologie Technische Hochschule,
Darmstadt, Germany. When discussing the habit,
Buschingeretal., (1987) recorded the ant as Plagiolepis
sp. and this name was used by Williams (1993). Pro-
fessor Buschinger (personal communication) has since
indicated that the ant is actually a species of Acropyga.
The most likely species is A.(Rhizomyrma) paleartica
Mennozzi, the only species known in Greece.

The accompanying figure first appeared in the Ento-
mologist’s Gazette and is reproduced here from the
original illustration with slight modification.

Eumyrmococcus falciculosus sp. n.
(Fig. 3)

DESCRIPTION

Adult female on microscope slide membranous, elon-
gate, largest specimen 1.15 mm long, 0.50 mm wide,
widest at about mesothorax; head and thorax rounded,
constricted slightly between abdominal segments II and
III, abdomen tapering and narrowing between abdomi-
nal segments VII and VIII, posterior end of body
rounded, abdominal segment VIII 200 im wide at base.

D.J. WILLIAMS

Positions of each anal lobe with 2 stout dorsal setae, each
about 188m long, and | ventral setaabout 125umlong,
forming a group of 3. Antennae situated on ventral head
margin, each 65-77 um long, with 2 segments; basal
segment wide, second segment tapering. Legs well
developed, slender; hind trochanter + femur 115—
118 um long, hind tibia+ tarsus about 100um long, claw
slender, about 20 um long. Ratio of lengths of hind tibia
+ tarsus to hind trochanter + femur 0.84—0.86. Ratio of
lengths of hind tibia to tarsus 0.73—0.81.Tibiae swollen
then tapering, tarsi widening near base then narrowly
tapering. Leg setae pointed. Labium about 117—125 um
long, as long as clypeolabral shield, 77-80 um wide;
ratio of length to width 1.51—1.56. Circulus present,
distorted in available specimens but at least 50m wide,
situated near middle of abdominal segment III.Anal ring
about 75 um wide, with 6 slender setae, the anterior pair
each about 32.5 um long, the second and posterior pairs
longer but not complete in available material.

Dorsal surface with unusual sickle-shaped setae,
fairly crowded, present as far forward as abdominal
segment III, those posteriorly on abdominal segment
VIII each about 20 Um long, others on abdomen
mostly 12.5 tm long. A few flagellate setae each
12.5 um long, present among the sickle-shaped setae.
Setae on head, thorax and abdominal segments I and
II, curved but fairly stiff and pointed, mostly about
15 um long, moderately numerous. Thicker sensory
setae, each bluntly tipped and with large setal collars,
few, distributed mainly around margins.

Ventral surface with a similar distribution of sickle-
shaped setae, pointed setae and thick sensory setae, to
those on dorsum. A few long, stout sensory setae
present in medial area of thorax.

MATERIAL

HOLOTYPE. Adult 9, Indonesia: Sumatra, Krui,
Kubu Prau, among primary forest litter, 800 m,
29.v.1991 (L. Deharveng & A. Bedos) (MNHN).

PARATYPE. Indonesia: Sumatra, same data as
holotype. 1 adult 2 (BMNH).

Non-type material, Indonesia: Sumatra, Jambi Proy-
ince, Rantau Pandan, in soil in Hevea plantation,
9.vi.1991 (L. Deharveng & A. Bedos).

COMMENTS. This is a curious species with peculiar
sickle-shaped setae on the abdomen and short 2-seg-
mented antennae. It has no close relatives thus far
discussed here. The antennae are similar to those of E.
lanuginosus sp. n., also described from Sumatra, but
E. lanuginosus is extensively clothed in long
microtrichia whereas inE. falciculosus any microtrichia
are short and barely perceptible.

The specimen recorded from Rantau Pandan is
extremely poor but is recognisable as this species by
the sickle-shaped setae.

The epithet ‘falciculosus’ is based on the Latin word

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 11

‘falcicula’ meaning small sickle, and the Latin suffix
‘-osus’ meaning ‘abundance of’, referring to the num-
erous setae of this shape.

Eumyrmococcus kolombangarae sp. n.
(Fig. 4)

DESCRIPTION

Adult female on microscope slide broadly oval, about
0.90 mm long, 0.60 mm wide, widest at mesothorax,
tapering to a constriction between abdominal seg-
ments VII and VIII, base of abdominal segment VIII
180 tm wide. Posterior end of body rounded; posi-
tions of each anal lobe with 2 ventral setae and | dorsal
seta forming a group of 3, each seta about 80 Um long.
Antennae lying on ventral head margin, each about
110 um long, with two segments, the second tapering.
Legs well developed; hind trochanter + femur about
168 um long, hind tibia + tarsus 142 ttm long, claw
slender, unusually long, about 45 um long. Ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 0.84. Ratio of lengths of hind tibia to tarsus
1.84. Tarsi each swollen near base then tapering to
narrow distal end. All leg setae stout and flagellate.
Labium distorted in available specimen but at least
100 ttm long and fairly wide, about as long as
clypeolabral shield. Circuli numbering 2, each about
15 um wide, situated near posterior edges of abdomi-
nal segments II and III but within borders of segments.
Anal ring about 75 lm wide, with 6 setae, each about
65 um long.

Dorsal surface with crowded flagellate setae occu-
pying most of surface except for well marked
intersegmental areas; setae on abdominal segment
VIII mostly 15 {1m long, anteriorly about 12.5 um
long; a few setae on any segment with larger setal
collars.

Ventral surface with crowded setae as on dorsum
except in medial area of thorax where they are longer
and not numerous. Long stout setae, each about 35 um
long, present in pairs near mid-line of abdominal
segments II-VI.

MATERIAL

HoLoryPe. Adult ?, Solomon Islands: New Geor-
gia Group, Kolombangara, E. Kusi, 600 m, associated
with Acropyga lauta in log, 30.vili.1965 (Isiah)
(ANIC).

PARATYPES. Solomon Islands: same data as
holotype, 1 2nd instar (ANIC), 1 2 pupa (ANIC), |
pupa (BMNH).

COMMENTS

This species is related to the type species E. smithii in
the arrangement of the densely crowded flagellate
setae, at least on the dorsum, but differs in possessing

two small circuli. E. smithii lacks circuli completely.
Furthermore, in E. kolombangarae, the anal lobe setae
are at most only 80 um long and the posterior anal ring
setae are about 65 {1m long, whereas in E. smithii,
these setae are noticeably much longer, at least SOO um
long.

E. kolombangarae is very close to E. kusiacus,
herein described, differing mainly in lacking long
stout setae in rows at the posterior edges of the dorsal
and ventral abdominal segments. InE. kolombangarae,
long stout setae are present in pairs only near the
midline of the venter of abdomen.

The description of this species has also been based
on an almost perfect adult female still within the pupal
instar. A single second instar is also available. This is
similar to the adult female, differing mainly in having
shorter limbs, a single circulus present on abdominal
segment III, and in the body setae which are not so
dense.

The epithet is based on the Latin genitive of the
place name meaning ‘of’ or ‘from’.

Eumyrmococcus kruiensis sp. n.
(Fig. 6)

DESCRIPTION

Adult female on microscope slide, elongate, membra-
nous 0.8l1mm long, 0.38 mm wide, widest at
mesothorax, abdomen gradually tapering, slightly con-
stricted between abdominal segments VII and VIII,
base of abdominal segment VIII 120 um wide, poste-
rior end of body fairly straight. Positions of each anal
lobe with 2 stout dorsal setae and | ventral seta, each
about 70 um long, forming a group of three. Antennae
placed on ventral head margin, each 82.5 um long,
with 2 segments, the second segment 60 Lm long,
tapering. Legs well developed; hind trochanter + fe-
mur about 117.5 um long, hind tibia + tarsus 95 Um
long, claw slender, about 27.5 um long, 5 um wide at
base. Ratio of lengths of hind tibia + tarsus to hind
trochanter + femur ().80. Ratio of lengths of hind tibia
to tarsus 1.11. Tibiae swollen then tapering to narrow
distal ends. All legs with stout flagellate setae. Labium
about 92.5um long, same length as clypeolabral shield,
65 um wide; ratio of length to width 1.42. Circulus
large, about 45 ttm wide, situated near posterior end of
abdominal segment III but within borders of segment.
Anal ring about 70 um wide, with 6 setae; anterior and
middle pairs slender, each 25—37 {1m long, posterior
pair stout, each about 75 um long, similar to anal lobe
setae.

Dorsal surface densely covered in hair-like
microtrichia and slender flagellate setae except for
intersegmental areas. Most setae on abdominal seg-
ment VIII about 15 tm long, those on anterior
abdominal segments 10.0—12.5 um long, most on head
and thorax 5.0-—12.5 um long.

Ventral surface with similar microtrichia and setae
to those on dorsum; on abdominal segment VIII, some
setae as long as 40 um, most other setae on abdomen
about 25 um long, marginal areas of head and thorax
with short slender setae each about 12.5 tm long.
Long, stout flagellate setae, each 25—30 Um long, with
large setal collars, present on head and submarginal
and medial areas of thorax.

MATERIAL

HOLOTYPE. Adult 9, Indonesia, Sumatra, Krui,
Pahmungan, in soil of dammar plantation, 23.v.1991
(L. Deharveng & A. Bedos) (MNHN).

PARATYPES. Indonesia, Sumatra, same data as
holotype, 3 2nd instars (MNHN), 2 2nd instars
(BMNH).

COMMENTS. This is a distinctive species within the
group extensively clothed in hair-like microtrichia.
The stout setae in groups of three on the anal lobes and
the posterior pair on the anal ring are much shorter
than those in E. maninjauensis, E. sarawakensis and
E. sulawesicus. E. kruiensis also differs from these
three species in lacking any sensory setae with swollen
tips.

There are also available five second instars. These
are similar to the adult female in the dense covering of
microtrichia but the limbs are shorter and the body
setae are fewer.

The epithet is based on the place name ‘Krui’ and
the Latin suffix ‘-ensis’ denoting origin.

Eumyrmococcus kusiacus sp. n.
(Fig. 6)

DESCRIPTION

Adult female on microscope slide broadly oval,
1.12 mm long, 0.65 mm wide, widest at mesothorax,
abdomen tapering, abdominal segment VIII 220 Um
wide at base. Positions of each anal lobe with 2 stout
dorsal setae and | ventral seta forming a group of 3,
each about 100 um long. Antennae placed near ventral
head margin, each 112—120 um long, with 2 segments,
the second tapering, both segments with long stout
setae. Legs well developed, slender; hind trochanter +
femur about 200 um long, hind tibia + tarsus 158-162
uum long, claw slender, about 58 um long. Ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 0.79-0.81. Ratio of lengths of hind tibia to
tarsus 1.02—1.16. Tarsi each swollen near base then
tapering to narrow distal end. All legs with stout
flagellate setae. Labium 150—162tm long, about same
length as clypeolabral shield, 75—80 [1m wide; ratio of
length to width 1.50-1.62. Circuli numbering 2 situ-
ated near posterior edges of abdominal segments II
and III but within borders of segments, diameter on
abdominal segment II about 10 Lm, on abdominal

D.J. WILLIAMS

segment III about 17.5 um, each deeply cupped. Anal
ring about 100 um wide with 6 setae; anterior and
second pairs each 75 Um long, posterior pair each
about 87 Lum long, the anterior pair with setal collars
touching.

Dorsal surface with crowded flagellate setae; on
abdominal segment VIII mostly about 40 um long,
anteriorly on abdomen and remainder of body mostly
about 15 um long except towards posterior edges of
abdominal segments, mostly about 25 um long. Setae
on abdomen in fairly narrow bands leaving extensive
intersegmental bare areas. Setae on head and thorax
much more crowded than on abdomen. Long stout
setae, each 80—90 um long, present in rows at posterior
edges of abdominal segments II—-VII.

Ventral surface with similar setae to those on dor-
sum, sparse on venter of thorax. Long stout setae as on
dorsum, present at posterior edges of abdominal seg-
ments I—VII.

MATERIAL

HoLotyPe. Adult ?, Solomon Islands: New Geor-
gia Group, Kolombangara, Kusi, 1220 m, associated
with Acropyga lauta, in log, 29.vii.1965 (Isiah)
(ANIC).

PARATYPE. Solomon Islands: same data as holotype,
1 adult 2? (BMNH).

COMMENTS. E.kusiacus is related to E. smithii but
differs in possessing two small circuli, and short stout
setae on the anal lobes. These setae are scarcely longer
than the width of the anal ring. In E. smithii, the circuli
are absent and the setae on the anal lobes are conspicu-
ously longer than the width of the anal ring. E. kusiacus
is very close to E. kolombangarae, differing mainly in
possessing long stout setae on the dorsum and venter at
the posterior edges of the abdominal segments. In E.
kolombangarae, there are long stout setae in the me-
dial areas of the ventral abdominal segments only.

The name is based on ‘Kusi’, the place of origin, and
the Latin suffix “-acus’ meaning “belonging to’.

Eurmyrmococcus lamondicus sp. n.
(Fig. 7)

DESCRIPTION

Adult female on microscope slide elongate, slender,
largest specimen 1.28 mm long, 0.53 mm wide
(holotype 1.40 mm long, 0.40 mm wide), widest at
about mesothorax, head and thorax rounded, narrow-
ing at abdominal segment III then narrowly tapering to
small constriction between abdominal segments VII
and VIII, base of abdominal segment VIII about 100 um
wide. Positions of each anal lobe with 2 stout dorsal
setae and | ventral seta, each about 325 um long,
forming a group of 3. Antennae placed on dorsal head
margin, each 330-340 um long, with 2 segments, the

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 13

second 260-280 um long, tapering. Legs well devel-
oped, slender; hind trochanter + femur 193-218 um
long, hind tibia + tarsus 112-122 um long, claw
slender, about 27.50 um long, 6.25 um wide at base.
Ratio of lengths of hind tibia + tarsus to hind tro-
chanter + femur 0.91—1.00. Ratio of lengths of hind
tibia to tarsus 1.38—1.43. Tibiae swollen then tapering
abruptly to narrow distal ends. All legs with stout
pointed setae. Labium 127.5—137.5 tm long, slightly
longer than clypeolabral shield, 80-105 um wide;
ratio of length to width |.21—1.66. Circuli numbering
2, situated near middle of abdominal segments I and
II, cup-shaped, about as deep as wide; on abdominal
segment II about 12.5 tum wide, on abdominal seg-
ment III about 11.0 um wide. Anal ring about 45 um
wide, with 6 setae; anterior pair slender, each about
72 um long, second and posterior pairs stouter, each
about 187 tm long, resembling anal lobe setae.

Dorsal surface with short crowded setae in bands
across segments as far forward as prothorax; setae at
posterior end of abdominal segment VIII each about
30 um long, most on abdominal segment VII about
12.5 um long, anteriorly mostly 7.5—10.0 um long; a
few on most segments with larger setal collars; on
prothorax and mesothorax setae occupying medial
areas except for marginal band, absent from head and
in noticeable intersegmental areas. Stout flagellate
setae, 32.5-37.5 tum long, present in medial area of
head, interspersed with a few smaller setae. Short
lanceolate sensory setae, each about 10 Um long, not
numerous, mingled with the slender setae posteriorly
to abdominal segment VI. Slender sensory setae, each
about 10 um long, and bluntly tipped, represented by
a few on abdominal segment II only, these in addition
to lanceolate setae.

Ventral surface with similar slender setae to those
on dorsum, those on venter of thorax in deep marginal
bands. Lanceolate sensory setae as on dorsum, few,
present on most segments. Slender sensory setae oc-
curring on abdominal segment II only. Stout flagellate
setae present on head and medial area of thorax.

MATERIAL

HoLotyPe. Adult 2, Australia: Queensland, Mt
Lamond, 12°44’S 143°18’E, associated with Acropyga
sp., in rotting wood, 9—15.vi.1971 (R.W. Taylor & G.J.
Feehan) (ANIC).

PARATYPES. Australia: Queensland, same data as
holotype, 3 adult9? (ANIC), 2 adult 9? (BMNH), 4 2nd
instars (ANIC), 3 2nd instars (BMNH).

COMMENTS. The distribution of the dorsal and ven-
tral slender setae and the shape of the antennae, place
this species near E. nipponensis. E. lamondicus, how-
ever, comes closest to E. queenslandicus sp. n. in
possessing minute lanceolate sensory setae, whereas
those of E. nipponensis are blunt and swollen apically.

E. lamondicus differs from E. queenslandicus in pos-
sessing also some slender blunt sensory setae on
abdominal segment II and having two small circuli. In
E. queenslandicus the blunt sensory setae are absent
and there is only a single but large circulus. The
slender setae on the venter of the thorax in E.
lamondicus are also in much deeper marginal bands
than in E. queenslandicus.

The epithet ‘/amondicus’ is based on the place name
Mt Lamond and the Latin suffix *-icus’ meaning ‘be-
longing to’.

Eumyrmococcus lanuginosus sp. n.
(Fig. 8)

DESCRIPTION

Adult female on microscope slide elongate-oval, mem-
branous, largest specimen 1.00 mm long, 0.50 mm
wide, widest at mesothorax then gently tapering to
abdominal segment VII, with a constriction between
abdominal segments VII and VIII, posterior end of
body rounded. Positions of each anal lobe with numer-
ous long stout setae, mostly about 75 [Um long.
Antennae placed on ventral margin of head, short,
each 35,0-37.5 tum long, with 2 segments, second
segment tapering, almost conical. Legs well devel-
oped, robust, hind trochanter + femur 145-150 um
long, hind tibia + tarsus 97.5-110.0 um long, claw
slender, 10.0-12.5 tum long, 7.5 tum wide at base.
Ratio of lengths of hind tibia + tarsus to hind tro-
chanter + femur 0.67—0.73. Ratio of lengths of hind
tibia to tarsus 1.00-1.09. Tibiae each swollen near
base then tapering to narrow distal end. All legs with
stout flagellate setae. Labium 95—100 um long, 50.0—
62.5 um wide, ratio of length to width 1.6—1.9. Circuli
numbering 2 or 3, when 3 then situated near middle of
mesothorax and abdominal segments II and II]. When
only 2 then absent from mesothorax; each circulus
about 12.5 um in diameter except on abdominal seg-
ment III when about 10 Um in diameter,
truncate-conical but deeply cupped from apex. Anal
ring 70—74 um wide, with 6 setae; anterior pair slen-
der, each about 30 um long, second pair each about
50 um long, posterior pair stouter, each about 80 um
long; anal ring setae difficult to discern because of
surrounding setae.

Dorsal surface extensively covered in fine hair-like
microtrichia and short fine setae except for bare in-
tersegmental areas. Short setae on abdominal segment
VII mostly about 15 um long, those anteriorly mostly
about 7.5 um long except for a few about 12.5 um long
with slightly larger setal collars. Long stout setae
present towards posterior edges of abdominal seg-
ments IV—VUI, those on abdominal segment VIII
numerous, 55—75 1m long; anteriorly mostly 55 um
long, distributed across the segments except on ab-
dominal segment IV where they are sparse.

Ventral surface with similar microtrichia and setae to
those on dorsum, covering most of surface; the short
setae fine in medial area of thorax. Long flagellate
setae, much more numerous than on dorsum, present
towards posterior edges of abdominal segments and in
medial areas of head and thorax. A few stout sensory
setae, each with swollen tip, present laterally on thorax.

MATERIAL

HoLotyPe. Adult ?, Indonesia: Sumatra, Si Antar
[2°40’N 98°43’E], with ants (Smithsonian Expedi-
tion), 1937(W.M. Mann) (USNM).The holotype is the
middle specimen of five, all mounted in a row on the
same slide and is clearly marked.

PARATYPES. Indonesia: Sumatra, same data as
holotype, 3 adult 9, clearly marked in row with
holotype, 1 2nd instar to right of other specimens
(USNM).

COMMENTS. E. lanuginosus belongs to the group of
species extensively covered in minute hair-like
microtrichia. It differs from all the other species in
possessing only short, almost conical antennae, and
numerous stout setae on the positions of each anal
lobe, not differentiated into groups of three.

The specimens were first studied by Harold Morrison
who noted on the envelope that the species had been
discussed by Roepke (1930). Although Roepke’s illus-
tration of the mealybug is rather simple, it does show
the small tubercle-like antennae. The ant species was
provisionally identified as Cladomyrma sp. but the
equally simple illustration of the ant with the mealybug
held in the mandibles, may be a species of Acropyga.

The epithet ‘lanuginosus’ is a Latin adjective mean-
ing downy or woolly, referring to the dense covering of
microtrichia.

Eumyrmococcus maninjauensis sp. n.
(Fig. 9)

DESCRIPTION

Adult female on microscope slide, membranous, pyri-
form, about 0.83 mm long, 0.48 mm wide, widest at
mesothorax, narrowing abruptly at about abdominal
segment II, then gently tapering to rounded posterior
end; abdominal segment VIII 150 tm wide at base.
Positions of each anal lobe with 2 stout dorsal setae
and | ventral seta, each about 275 um long, forming a
group of 3. Antennae situated on ventral head margin,
each 95 im long, with 2 segments, the second segment
about 62.5 um long, tapering, | antenna partly divided
on | side only. Legs well developed, slender, hind
trochanter + femur about 107.5 tum long, hind tibia +
tarsus about 105 tm long, claw, slender, about 25 um
long, 5.0 um wide at base. Ratio of lengths of hind
tibia + tarsus to hind trochanter + femur 0.97. Ratio of
lengths of hind tibia to tarsus 1.0. Tibiae each narrow

D.J. WILLIAMS

at base, widening, then tapering abruptly to narrow
distal end. All legs with outer setae sensory, each with
slightly swollen tip, inner setae flagellate. Labium
fairly wide, difficult to measure in available specimen,
about same length as clypeolabral shield. Circulus
large, about 32.5 lum wide, placed near middle of
abdominal segment III. Anal ring about 75 um wide,
with 6 setae; anterior pair slender, each 67.5 tum long,
second pair similar, each about 70 um long, posterior
pair stout and long, about 220 um long, resembling
anal lobe setae.

Dorsal surface densely covered in hair-like
microtrichia and short, fine setae, except for interseg-
mental areas. Setae on abdominal segment VIII
flagellate, mostly about 20 tm long; similar setae 10—
20 um long, present towards posterior edges of
abdominal segments [V—VII; anteriorly, setae much
more slender, 6.25—12.00 long, sometimes difficult to
discern amongst the dense microtrichia.

Ventral surface with similar microtrichia and setae
to those on dorsum. Obanal and cisanal setae present,
each about 50 um long; stout flagellate setae present
on abdominal segment VIII, as long as 67 um but most
at posterior edge of abdominal segments about 50 um
long, mixed with others about 20 um long. Minute
slender setae distributed around margins of thorax.
Medial area of thorax with stout flagellate setae; me-
dial area of head and lateral area of thorax with a few
stout sensory setae, each 15—20 um long, with slightly
expanded tips and with wide thick setal collars.

MATERIAL

HOLoTyPe. Adult 9, Indonesia: Sumatra, Barat
Province, Kotomalintang, nr Lake Maninjau, in soil,
extracted by Berlese funnel apparatus, 19.xii.1994 (L.
Deharveng & A. Bedos) (MNHN).

COMMENTS. Among the group of species with a
dense covering of hair-like microtrichia, E.
maninjauensis seems to be related to E. sarawakensis
and E. sulawesicus in possessing long stout setae in
groups of three on the positions of each anal lobe, and
a pair of similar posterior setae on the anal ring. The
antennae of E. sarawakensis and E. sulawesicus, how-
ever, are placed well on the dorsum of the
cephalothorax, whereas in E. maninjauensis they are
placed on the ventral head margin. Furthermore, any
stout sensory setae with swollen tips in E.
maninjauensis are distributed on the venter only,
whereas in the other two species they occur on the
dorsum also.

The epithet is based on the place name ‘Lake
Maninjaw’ and the Latin suffix ‘-ensis’ denoting local-

ity.
Eumyrmococcus neoguineensis sp. n.
(Fig. 10)

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 15

DESCRIPTION

Adult female on microscope slide membranous except
for abdominal segment VIII lightly sclerotized; elon-
gate-oval, largest specimen 1.15 mm long, 0.57 mm
wide, widest at about abdominal segments II and III;
head and thorax rounded, anterior margin of head
sometimes straight, constricted slightly at mesothorax,
then widening to anterior abdominal segments, re-
mainder of abdomen tapering, base of abdominal
segment VIII about 230 ttm wide, apparent anal lobes
set well apart, apex of abdomen protruding. Position
of each anal lobe with 3 or 4 dorsal setae and usually 5
ventral setae forming a group of 8 or 9; inner ventral
setae slender, each about 150 Um long, others increas-
ing in length distally, outer ventral and dorsal setae
stout, 550-600 tum long. Antennae situated on dorsal
head margin, each 420-440 um long, with 4 segments,
second segment longest with a row of short conical
setae on anterior edge, increasing in length distally;
setae on third and fourth segments long, longest on
third segment, about 325 lum long. Legs well devel-
oped; hind trochanter + femur 188—232 um long, hind
tibia + tarsus 180-220 um long, claw slender, about
34 um long, 7.5m wide. Ratio of lengths of hind tibia
+ tarsus to hind trochanter + femur 0.93—0.95. Ratio of
lengths of hind tibia to tarsus 1.20—1.29. First legs with
noticeably larger coxae than in other legs and longer
trochanter + femur, 232—245 um long, and longer tibia
+ tarsus, 260-272 um long. Tibiae of all legs swollen
near bases and narrowly tapering distally. All legs with
long slender setae, except for some stout setae on
posterior surface of femur of each first leg. Labium
162-175 tm long, 87-100 tum wide, longer than
clypeolabral shield; ratio of length to width 1.62—1.86.
Circulus present near anterior end of abdominal seg-
ment II but placed well within borders of segment,
20-25 um in diameter, deeply cupped, about as deep
as wide or slightly deeper. Anal ring about 90 um
wide, with 14 slender setae, each about 110 Lm long.

Dorsal surface densely covered in slender flagellate
setae except around antennal bases and intersegmental
areas; some setae at posterior end of abdominal seg-
ment VIII each about 87 um long, those at anterior end
of segment about 25 um long. Anteriorly on abdomen,
as far forward as abdominal segment V, mostly 17—
25 um long; in medial area of abdominal segment VI,
medial and anterior areas of abdominal segment V, and
all anterior segments of abdomen, thorax and head,
extensively covered in minute setae 10.0-12.5 um
long. Posterior edges of abdominal segments each
with a row of stout setae, each about 100 [1m long, a
few at lateral edges of abdominal segments each about
125 um long.

Ventral surface with a similar distribution of setae to
that on dorsum but mainly bare on medial areas of
head and thorax except for a few short setae. A bunch
of invaginated setae present lateral to each first and

second coxa, these lightly sclerotized when prepared
on slides.

MATERIAL

HoLotyPe. Adult?, Papua New Guinea, East Sepik
Province, Yawasora [ Yauwasoru], near Wewak, asso-
ciated with Acropyga (Atopodon) ambigua Emery, in
rotting log, 4—6.vu1.1972 (R.W. Taylor) (ANIC).

PARATYPES. Papua New Guinea, same data as
holotype, 3 adult 9? (ANIC), 2 adult 9? (BMNH); same
data but in mountain log, 1 adult ? (BMNH); Morobe
Province, 16 km NW Lae, “Timber Track’, in rotting
wood, with A. (Atopodon) ambigua, 12.vi.1976 (R. W.
Taylor), \ adult 9 (ANIC).

COMMENTS. This species shares withE. corinthiacus
and E. scorpioides the character of 4-segmented an-
tennae but the antennae of E. neoguineensis are much
longer, with long setae on the third and fourth seg-
ments, and conical setae on the anterior edge of the
second segment. Furthermore, the long stout setae on
each anal lobe number eight or nine, whereas those in
the other two species are in groups of three. The
affinities of E. neoguineensis are obscure but
Xenococcus annandalei also possesses large groups of
long stout setae on the anal lobes and the antennae are
also 4-segmented with long setae on the third and
fourth segments. In Xenococcus, however, the anten-
nae are much longer, about as long as the body, and
there is well developed articulation between the first
and second segments. The bunches of invaginated
setae in E. neoguineensis have not been seen in any
other species. Despite the unusual characters, the true
affinities of this spectacular species appear to be with
the genus Eumyrmococcus to which it is here assigned.

The epithet is a Latinized word for New Guinea
with the Latin suffix ‘-ensis’ meaning ‘belonging or
pertaining to’.

Eumyrmococcus nipponensis Terayama

(Fig. 11)

Eumyrmococcus nipponensis Terayama, 1986: 509;
1988: 645; Ben-Dov, 1994: 152. HOLOTYPE 9°,
Japan, Noboritachi, Mikura-jima Is, Tokyo, on
rootlets of plants in nests of Acropyga [Atopodon]
nipponensis Terayama, 30.vi.1980 (K. Masuko)
(NIAES).

DESCRIPTION

Adult female on microscope slide pyriform, largest
specimen 1.40 mm long, 0.60 mm wide, widest at
mesothorax, abdomen tapering, constricted slightly
between abdominal segments VII and VIII, base of
abdominal segment VIII about 180 um wide. Position
of each anal lobe with 2 long, stout ventral setae and |
dorsal seta, forming a group of 3, each seta about

450 um long. Antennae situated on ventral head mar-
gin, each 2-segmented, 260-270 \1m long, second
segment tapering, 210-215 um long. Legs well devel-
oped, slender; hind trochanter + femur 180-200 um
long, hind tibia + tarsus 182-195 tm long, claw
slender, about 30 um long. Ratio of lengths of hind
tibia + tarsus to hind trochanter + femur 0.95-1.01.
Ratio of lengths of hind tibia to tarsus 1.05—1.21.
Tibiae swollen near middle then narrowly tapering. All
legs with outer setae stout, sensory, swollen apically;
inner setae flagellate. Labium 138—150um long, 75.0-
77.5um wide, slightly longer than clypeolabral shield;
ratio of length to width 1.84—1.93. Circulus present,
round to slightly elliptical, 20-30 um wide, deeply
cupped, about as deep as wide, situated near middle of
abdominal segment II. Anal ring 68-75 tm wide,
with 6 setae; anterior pair slender, each about 27 um
long, second pair stouter, each about 62 [im long,
posterior pair stout and long, each about 450 um long,
resembling anal lobe setae.

Dorsal surface extensively covered with short slen-
der flagellate setae, sparse near head margin and absent
in wide intersegmental areas; at posterior end of body
each seta about 15 um long, anteriorly on abdominal
segment VIII mostly 12.5 um long, elsewhere mostly
about 7.5 tum long. Short stout sensory setae, each
with noticeable swollen tip, fairly numerous across
segments and mixed with the short flagellate setae; at
posterior end of body, each about 11 um long but most
anteriorly each about 7.5 um long. Head with longer
stout sensory setae 12.5—25.0 um long. Most setae
with setal collars probably slightly raised from sur-
rounding derm so that setae appear to be surrounded
by halos.

Ventral surface with similar short setae as on dor-
sum but not so dense; absent from medial area of
thorax. Long sensory setae, each about 35 [1m long,
present at posterior edge of abdominal segment VII;
similar setae, each about 25 tim long, present on head.
Other sensory setae shorter, those around margins,
each 7.5m long as on dorsum, others 12—28 um long,
distributed across segments. Long, stout flagellate
setae present in medial area of thorax. Three very
slender sensory setae sometimes present on each side
of circulus. Obanal setae stout and long, about as long
as anal lobe setae. Many flagellate setae at posterior
end of body with thick setal collars.

MATERIAL

Japan: Yakushima Is, Amboh, on rootlets of plants in
nests of Acropyga (Atopodon) nipponensis Terayama,
17.x.1984(M. Terayama); Tokunoshima Is, Tampatsu-
yama, same data but 8. viii. 1984 (all labelled paratypes).

COMMENTS. E. nipponensis differs from E. smithii,
the only other species known from Japan, in possess-
ing a circulus and sensory setae with expanded tips. In
E. smithii, the circulus is absent and all setae are

D.J. WILLIAMS

flagellate. E. nipponensis seems to have affinities with
two Australian species, E. lamondicus and E. recalvus,
in possessing similar 2-segmented antennae, each with
the second segment long and tapering; also with minute
flagellate setae on the dorsum distributed at least as far
forward as the prothorax. Most of the sensory setae in
the two Australian species, however, are lanceolate,
whereas those in E. nipponensis are blunt and widely
expanded distally.

The ant attending this mealybug was described
originally by Terayama (1985), from material col-
lected at the same time.

The accompanying illustration has been prepared
from paratypes kindly made available by Dr M.
Terayama.

Eumyrmococcus queenslandicus sp. n.
(Fig. 12)

DESCRIPTION

Adult female on microscope slide elongate, largest
specimen 1.30 long, 0.65 mm wide, widest at
mesothorax, head and thorax rounded, abdomen taper-
ing to rounded posterior end, constricted between
abdominal segments VII and VIII, segment VIII about
150 um wide at base; positions of each anal lobe with
2 dorsal setae and | ventral seta, each about 300 um
long, forming a group of 3. Antennae situated on
dorsal head margin, each 270-300 um long, with 2
segments, the second tapering, 220-240 tm long.
Legs well developed, slender; hind trochanter + femur
190-192 tm long, hind tibia + tarsus 110-117 um
long, claw slender, about 27.5 um long, 5.0m wide at
base. Ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 0.97—1.02. Ratio of lengths of hind
tibia to tarsus 1.37—1.41. Tibiae swollen then tapering
towards narrow distal ends. All legs with stout pointed
setae. Labium 130-138 um long, slightly longer than
clypeolabral shield, 75—100 um wide; ratio of length
to width 1.38-1.70. Circulus present near middle of
abdominal segment II, large and conspicuous, 52—
60 tm in diameter, deeply cupped with almost parallel
sides but wider than deep. Anal ring 52.5-60.0 Um
wide, with 6 setae; anterior pair slender, each about
100 um long, second pair thicker, each about 220 um
long, posterior pair each about 250 Um long, almost as
thick and as long as anal lobe setae.

Dorsal surface with crowded slender flagellate se-
tae as far forward as thorax, those on prothorax
represented by a medial band, absent laterally on
mesothorax leaving bare areas on head and laterally on
prothorax, mesothorax and intersegmentally. Setae
towards posterior end of abdominal segment VIII each
about 37.5 um long, anteriorly on next few segments
mostly 10-20 tm long, on thorax and anterior ab-
dominal segments mostly 10 Um long; some setae on
any segment with larger setal collars than others. A

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 17

few sensory lanceolate setae, each about 10 [1m long,
present on thorax and abdominal segments I—V. Setae
on head sparse, mostly stout and flagellate, 35-42 um
long.

Ventral surface with similar setae to those on dor-
sum. Short slender setae present around margins only
of thorax and first abdominal segment, few. Long stout
setae, 35—42 um long, not numerous, present on head
and medial area of thorax. Sensory lanceolate setae,
same as on dorsum, few, distributed on abdominal
segments II—V.

MATERIAL

HoLoryePe. Adult 9, Australia: Queensland, Kir-
rama Range, NE slope of Mt Pershouse, 700 m,
associated with Acropyga sp., 5.v.1969 (E. Kennedy &
R.W. Taylor) (ANIC).

PARATYPES. Australia: Queensland, same data as
holotype, 7 adult 92 (ANIC), 5 adult 9? (BMNH), 1
adult? (USNM), | 2nd instar (ANIC), | 2nd instar
(BMNH); 4.5 km W of Cape Tribulation, 760 m, taken
in the mandibles of Acropyga sp. in flight, 20—
23.iv.1983 G.B. Monteith & D.K. Yeates), | adult 2
(QM).

COMMENTS. Among the species already described,
E. queenslandicus comes nearest to E. nipponensis in
possessing similar antennae, each with a long second
segment, and with the thorax and abdomen densely
covered in slender setae. In E. nipponensis, the sen-
sory setae are blunt and expanded apically, whereas in
E. queenslandicus they are lanceolate. E.
queenslandicus comes closest to E. lamondicus sp. n.
Both species have short lanceolate sensory setae but in
E. queenslandicus there is a large single circulus only,
present on abdominal segment III. In E£. lamondicus
there are two small circuli present on each of abdomi-
nal segments II and III. Furthermore, E. lamondicus
possesses some short, blunt sensory setae on abdomi-
nal segment II in addition to lanceolate sensory setae.
These blunt setae are absent from E. queenslandicus.

Two second instars are also available. These have a
similar appearance to the adult females but the short
flagellate setae are present only as far forward as
abdominal segment IV.

The epithet is based on the locality ‘Queensland’
and the Latin suffix *-icuws’ meaning ‘belonging to’.

Eumyrmococcus recalvus sp. n.
(Fig. 13)

DESCRIPTION

Adult female on microscope slide, narrow, elongate,
1.40 mm long, 0.65 mm wide, widest at about
metathorax; head and thorax rounded, constricted
slightly between abdominal segments III and IV, then
tapering to posterior end of body, sharply narrowing

between abdominal segments VII and VIII; abdominal
segment VIII 162 tum wide at base before tapering
abruptly, projecting beyond positions of anal lobes.
Each anal lobe with a group of 3 long stout setae, 2 on
dorsum and | on venter, each seta at least 240 um long
but difficult to measure in available specimen. Anten-
nae each about 270 um long, with 2 segments, the
second segment curved, 205 tm long. Legs well de-
veloped, slender; hind trochanter + femur about 170 um
long, hind tibia + tarsus about 140 tm long, claw
slender, about 28 um long. Ratio of lengths of hind
tibia + tarsus to hind trochanter + femur 0.82. Ratio of
lengths of hind tibia to tarsus 1.8. Inner leg setae
flagellate, outer setae sensory, with slightly swollen
tips. Tibiae swollen towards distal ends then narrow-
ing, tarsi expanding then tapering to narrow distal
ends, no thicker than bases of claws. Labium about
112 um long, 82.5 im wide, about same length as
clypeolabral shield; ratio of length to width 1.35; all
labial setae flagellate. Circuli numbering 2, each about
10 um wide, situated near middle of each of abdomi-
nal segments II and III, deeply cupped. Anal ring about
48 um wide with 6 setae; 2 anterior pairs slender,
curled in available specimen but about twice length of
diameter of anal ring, posterior pair thicker, almost as
long as apical setae.

Dorsal surface with crowded slender setae as far
forward as abdominal segment II; those on abdominal
segment VIII each about 25 tm long, on abdominal
segments V—-VII mostly 17.5 um long, anteriorly about
12.5 um long; some on any segment with larger setal
collars. Setae on head, thorax and abdominal segment
I, all sensory with only minutely swollen tips, not
numerous; many on head and medially on thorax each
up to 37.5 um long: others on thorax and abdominal
segment I 12.5 um long, mingled with minute and
slightly swollen setae each about 3.7 Lim long.

Ventral surface with similar setae to those on dor-
sum as far forward as abdominal segment II; anteriorly
not numerous; metathorax and abdominal segment I
with short sensory setae as on dorsum, on margins of
head and thorax mostly long and sensory, up to 37.5 um
long; others on medial areas of thorax, long and flag-
ellate.

MATERIAL

HoLoryePe. Adult 2, Australia, Queensland, Byfield,
22°51’S, 150°39’E, associated with Acropyga sp.,
26.x.1976 (R.W. Taylor, T-A. Weir) (ANIC).

PARATYPES. Australia, same data as holotype, 4
2nd instars (ANIC), 2 2nd instars (BMNH).

COMMENTS. This species is very close to E. taylori
sp. n., differing mainly in the shape of the setae
anterior to abdominal segment II. In E. reclavus, these
setae on the dorsum and around the ventral margins are
all sensory and slightly expanded apically, the only

flagellate setae are those situated medially on the
venter of the thorax. In E. taylori, all setae in these
positions are flagellate.

Material at hand also contains six second instars.
These resemble the adult female but the minute flagel-
late setae are not so crowded and the appendages are
slightly smaller.

The epithet ‘recalvus’ is the Latin adjective mean-
ing “bald in front’, referring to the paucity of setae at
the anterior end of the body.

Eumyrmococcus sarawakensis sp. n.
(Figs. 14, 15)

DESCRIPTION

Adult female on microscope slide extremely slender,
largest specimen 1.35 mm long, 0.52 mm wide, widest
at about prothorax, then gently tapering, abdominal
segment VII about 115 um wide at base. Position of
each anal lobe with | stout dorsal seta and 2 ventral
setae forming a group of 3, each 430-488 um long.
Antennae placed well on dorsum of cephalothorax,
each 260-267 tm long, with 2 segments, second
tapering, 210-215 tm long. Legs well developed,
slender; hind trochanter + femur 162.5—182.0um long,
hind tibia + tarsus 160-185 um long, claw slender,
about 22.5 um long, 6.25 lum wide at base. Ratio of
lengths of hind tibia + tarsus to hind trochanter + femur
0.97-1.01. Ratio of lengths of hind tibia to tarsus 1.00—
1.05. Outer setae on legs sensory, with swollen tips,
inner setae flagellate. Labium about 137 tm long,
longer than clypeolabral shield, 70.0-82.5 um wide,
ratio of length to width 1.66—1.96. Circulus large,
round to slightly elliptical, 75—85 um wide, situated in
middle of abdominal segment III, cupped or bowl-
shaped, not so deep as wide. Anal ring 65-70 Um wide,
with 6 setae; anterior pair each about 32.5 um long,
second pair each about 62.5 tm long, posterior pair
about 390 um long, similar to anal lobe setae.

Dorsal surface with minute setae and hair-like
microtrichia covering entire surface except for in-
tersegmental areas. Setae on abdominal segment VIII
each about 10.0—12.5m long, others anteriorly mostly
about 5 wm long, difficult to discern among
microtrichia. Stout sensory setae with swollen tips,
present on head, each 17.5— 22.5 um long, not numer-
ous. A few sensory setae, each about 12.5 um long,
present on thorax and abdominal segment I.

Ventral surface with similar setae and microtrichia
to those on dorsum. Stout sensory setae present on
head and thorax, not numerous; short sensory setae,
10-15 um long, few, distributed across abdominal
segments II-VI.

Second instar
Similar in shape to adult female. Legs and antennae
shorter. Hair-like microtrichia present on abdomen

D.J. WILLIAMS

only. Short sensory setae extending to abdominal seg-
ment VII. Circulus large as in adult female.

Adult Male (Fig. 15).

Body of adult male degenerate, elongate, slender,
largest specimen 1.35 mm long, 0.50 mm wide, widest
at mesothorax, head and thorax rounded, then gently
tapering to rounded posterior end, base of abdominal
segment VIII about 150 mm wide. Area between an-
tennae on venter tessellated. Antennae short, almost
conical, 2-segmented, 37.5-45.0 mm long, situated
near ventral head margin. Legs squat, coxae wide;
trochanter + femur fused, unsegmented, stout, about
67.5 mm long; tibia + tarsus slender, 50-57 mm long,
sometimes with indistinct segmentation; claw dis-
torted, stout and almost triangular, about 12.5 mm
long. Genital capsule retracted inside abdomen, ven-
tral slit almost square. Aedeagus about 100 mm long,
stout, almost trumpet-shaped at apex. Anus oval, situ-
ated at posterior end of abdomen at base of sclerotized
extension to abdominal segment VIII, this extension
about 1.5 mm wide and long, hook-shaped in profile,
equivalent to long style in male of E. taylori.

Dorsal and ventral surfaces practically naked except
for some minute setae, each with setal length consider-
ably smaller than width of collar, except on dorsum of
thorax where setae slightly longer.

COMMENTS. ‘There is available also a single male
pupa with an adult male inside. Also available are
some female pupae, one of which shows the develop-
ing adult female inside.

MATERIAL

HoLoryPe. Adult 9, Malaysia, Sarawak, 1°38’N
113°35’E, associated with Acropyga sp., 25.11.1963
(A. Emerson) (ANIC).

PARATYPES. 9, Malaysia, Sarawak, same data as
holotype, 1 adult ?, 7 adult dic’ (ANIC), 2 adult 92, 7
adulto'o' (BMNH), 2 2nd instars (ANIC), 2 2nd instars
(BMNH), 5 pupae 9? (ANIC), 3 pupae 99 (BMNH), 3
pupae dc’ (ANIC), 2 pupae d'o' (BMNH).

COMMENTS. This singular species has an unusual
shape, evenly tapering posteriorly from the prothorax.
It belongs to a group collected within the tropics and is
extensively clothed in hair-like microtrichia and minute
setae. E. sarawakensis comes closest to E. sulawesicus
from which it differs mainly in possessing shorter and
fewer stout sensory setae on the cephalothorax, mostly
17.5—22.5 um long. In E. sulawesicus, these setae are
noticeably longer, 45-50 um long and more numerous.

Most of the differences are small but both species
are known from widely isolated localities and at present
this separation is justified.

The epithet is based on the name of the locality and
the Latin suffix ‘-ensis’ indicating origin.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 19

Eumyrmococcus scorpioides (De Lotto)
(Fig. 16)

Xenococcus scorpioides De Lotto, 1977: 33.
HOLOTYPE, SouthAfrica, Cape Province, Jacobs
Bay (SANC) [examined].

Eumyrmococcus scorpioides (De Lotto), Williams,
1993: 217; Ben-Dov, 1994: 152.

DESCRIPTION

Appearance in life not recorded. Adult female on
microscope slide elongate, body membranous except
for moderately sclerotized abdominal segment VIII,
largest specimen 1.55 mm long, 0.77 mm wide, widest
at mesothorax, head and thorax rounded, with a con-
striction between abdominal segments III and IV,
narrowing abruptly, remainder of abdomen tapering
with a further constriction between abdominal seg-
ments VII andVIII; segment VIII about 170m wide at
base, posterior end rounded. Positions of each anal lobe
with 2 stout ventral setae and | dorsal seta, each about
800 um long, forming a group of 3.Antennae slender, 4-
segmented, each 175-190 um long. Legs well
developed; hind trochanter + femur 170—180 um long,
hind tibia + tarsus 150-162 um long, claw about
27.5 um long, stout, 12.5 um wide at base. Ratio of
lengths of hind tibia + tarsus to hind trochanter + femur
0.87—0.92. Ratio of lengths of hind tibia to tarsus 1.41—
1.61. Outer setae on legs sensory, slightly swollen at
tips; inner setae flagellate. Labium 147-150 um long,
about as long as clypeolabral shield, 50-65 um wide;
ratio of length to width 2.26—3.00. Circuli numbering 2,
situated anteriorly on abdominal segments II and II but
within borders of segments; circulus on abdominal
segment II about 25 1m in diameter, on segment III
about 15 {1m in diameter, both cup-shaped. Anal ring
57-60 tum wide, with 6 slender setae; posterior pair
each about 87 [im long, anterior 2 pairs shorter but
curled in available specimens.

Dorsal surface with crowded flagellate setae on
abdomen, those at posterior end of abdominal segment
Vill each about 75um long, on abdominal segment VII
mostly about 37 Um long, anteriorly about 25 um long.
Head and thoracic setae mostly sensory, with slightly
swollen tips, 12.5—25.0 um long but some flagellate
setae present mingled with the sensory setae.

Ventral surface with similar setae to those on dor-
sum, crowded and flagellate as far forward as
abdominal segment III but a few sensory setae also
present on abdominal segment III. Anteriorly, setae
mostly sensory with swollen tips except in medial area
of thorax where they are mostly flagellate but a few
sensory setae also present.

MATERIAL

South Africa: Cape Province, Jacobs Bay, in nests of
Acropyga (Malacomyrma) arnoldi Sanschi, 7.viii. 1969
(A.J. Prins).

COMMENTS. In possessing 4-segmented antennae and
in the general distribution of body setae, this species is
related to E. corinthiacus. Most dorsal and ventral
setae on the head and thorax of E. scorpioides, how-
ever, are sensory with slightly swollen tips, whereas in
E. corinthiacus they are all flagellate. Moreover, the
long setae on the anal lobes of E. scorpioides are each
over 800 ttm long and those of E. corinthiacus are
thicker but at most only about 200-230 um long. E.
scorpioides differs from all other known species of
Eumyrmococcus in possessing stout claws, those in
other species being much more slender.

The accompanying illustration is based on the
holotype and two paratypes kindly made available for
this work by Ian Millar, Plant Protection Research
Institute, Pretoria, South Africa.

Eumyrmococcus smithii Silvestri

(Fig. 17)

Eumyrmococcus smithii Silvestri, 1926: 273; Williams,
1970: 138; 1978: 63; Ben-Dov. 1994: 152.
LECTOTYPE, China: Macao, [on roots of plants,
attended by the antAcropyga (Rhizomyrma) sautori
Forel], designated by Williams, 1978: 63 (IEAUN)
[examined].

COMMENTS. ‘This species was described and illus-
trated in detail by Williams (1970) from material
collected in various localities in China and Japan. Dr
M. Terayama has kindly made available some fresh
material collected in Okinawa and a new illustration is
presented here based on this material.

The most important characters are the 2-segmented
antennae, the absence of a circulus, long stout setae in
groups of three on each apparent anal lobe, each seta
about 500 tm long in the species illustrated but up to
700 Lim long in other specimens. The anal ring pos-
sesses 6 setae, the 2 anterior pairs short and slender
and the posterior pair stout, about as long as the anal
setae. All the body setae are short, slender and flagel-
late, occupying all the dorsal surface except for
intersegmental areas. A striking character, not seen in
any other species, is a long wide extension on each
hind coxal process.

In lacking any sensory setae with blunt or swollen
tips, E. smithii is related to E. kolombangarae and E.
Kusiacus, described here from the Solomon Islands. It
differs from both of these species in lacking a circulus.

MATERIAL
China: Macao, in nest of Acropyga (Rhizomyrma)
sauteri. Taiwan: Taichu, on sugarcane, 24.11.1933 (M.
Yanagihara). Japan: Ryukyu Is, Okinawa Is, in nest of
A. (Rhizomyrma) sauteri, 15.vii.1984(M. Terayama);
without locality, ex coll. R. Takahashi.

The species was also recorded from Shanghai by

Silvestri (1926, 1927) and from Taiwan by Takahashi
(1934).

Eumyrmococcus sulawesicus sp. 0.
(Fig. 18)

DESCRIPTION

Adult female on microscope slide narrowly elongate,
membranous, largest specimen 1.50 mm long, 0.60
wide, widest at mesothorax; head and thorax rounded
then constricting slightly between abdominal segments
VII and VIII, abdominal segment VIII 162 um wide at
base. Position of each anal lobe with 2 stout dorsal
setae and | ventral seta, forming a group of 3, each
510-525 um long. Antennae placed well on to dorsum,
each 275-295 wm long, with 2 segments, second
tapering, 215-232 um long. Legs well developed,
slender; hind trochanter + femur 200-205 tm long,
hind tibia + tarsus 182.5—205.0 um long, claw slender,
about 27.5 um long, 7.5 tum wide at base. Ratio of
lengths of hind tibia + tarsus to hind trochanter +
femur 0.90-1.02. Ratio of lengths of hind tibia to
tarsus 0.87—0.95. Leg setae short and sensory on outer
edges, each with swollen tip, flagellate on inner edges.
Labium about 142.5 um long, longer than clypeolabral
shield, 95 Um wide, ratio of length to width 1.5.
Circulus large, round to slightly elliptical. Anal ring
72-78 wm wide with 6 setae; anterior pair slender,
each about 45 um long, second pair each about 75 um
long, posterior pair stout, each about 475 um long,
resembling anal lobe setae with which they are some-
times intertwined.

Dorsal surface entirely covered with minute slender
setae and fine microtrichia. Setae on abdominal seg-
ment VIII mostly about 15 um long, anteriorly about
6-10 um long, many of longer setae with larger setal
collars, density less towards head and thorax. Long
sensory setae, each with swollen tip, 45-50 um long,
present on head and thorax. Microtrichia dense, hair-
like, short and stouter on abdominal segment VIII,
very slender anteriorly. Setae and microtrichia absent
from intersegmental areas.

Ventral surface with similar setae and microtrichia
to those on dorsum, covering entire surface except for
intersegmental areas. Long, stout sensory setae, as on
dorsum, present on head and thorax, fairly numerous,
becoming shorter, 17.5-25.0 um long, towards ante-
rior abdominal segments except for one or two, each
about 32 um long near circulus. Sensory setae towards
abdominal segment V mostly about 15m long, some-
times blunt, without swollen tips. Setae stout and
flagellate on medial area of thorax.

MATERIAL
HOLOTYPE. Adult 2, Indonesia, Sulawesi, Sulawesi
Utara, Toraut forest, Dumoga-Bone National Park,

with Acropyga (Atopodon) sp., 15.11.1985 (R.HLL.
Disney) (BMNH).

D.J. WILLIAMS

PARATYPES. Indonesia, Sulawesi, same data as
holotype, 3 adult 9? (BMNH), 6 2nd instars (BMNH).

COMMENTS. This species, covered in hair-like
microtrichia, is noticeably elongate with the antennae
placed well on the dorsum of the cephalothorax. It is
closely related to E. sarawakensis in most characters
but the stout sensory setae on the head and thorax are
longer, up to 50 um long, and more numerous. In E.
sulawesicus these setae are at most about 22.5um long.

A few second instars are also available. They differ
from the adult female in having much fewer minute
setae and microtrichia, but the stout sensory setae are
more numerous, extending posteriorly as far as ab-
dominal segment VII.

The epithet is based on the country of origin,
Sulawesi, and the Latin suffix ‘-icus’ meaning ‘per-
taining to’.

Eumyrmococcus taylori sp. n.
(Figs 19-22)

DESCRIPTION

Appearance in life not recorded. Adult female (Fig. 19)
on microscope slide elongate, narrow, membranous
except for posteriorend of body moderately sclerotized;
longest specimen 1.30 mm long, 0.50 mm wide, widest
at mesothorax; constricting between abdominal seg-
ments IV and V, widening at abdominal segment VI,
then narrowly tapering between abdominal segments
VIL andVIIL, base of abdominal segment VII 117-125
um wide; posterior end of body rounded, projecting
beyond actual anal lobes, these each with 2 ventral setae
and | long stout seta forming a group of 3, each seta
about 315 um long. Antennae each situated on dorsal
margin when flattened on slide, 310-390 um long, with
2 segments, second segment long, about 290-330 um
long, curved and tapering. Legs well developed, slen-
der, hind trochanter + femur 170-210 Lm long, hind
tibia + tarsus 150—190 Um long, claw slender, about 27
uum long. Ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 0.88—0.90. Ratio of lengths of hind
tibia to tarsus 1.14—1.25. Tibia swollen towards distal
end then tapering. Tarsus swollen then tapering to
narrow distal end, about as wide as base of claw. Leg
setae all flagellate. Labium aboutas long as clypeolabral
shield, 117-120 um long, 77.5—82.5 tum wide; ratio of
length to width 1.41—1.51. Circuli numbering | or 2,
situated within borders of abdominal segments II and
III; if only 1 circulus then present on abdominal seg-
ment III; circulus on abdominal segment II about 8.75
tum wide, on abdominal segment III about 11.25 um
wide, each circulus deeply cup-shaped. Anal ring 53—
57 tum wide, with 6 setae; anterior pair each about 67
uum long, second pair each about 1 12 um long, posterior
pair thicker, each about 275 um long, resembling the
apical setae.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI PAN |

Dorsal surface with crowded flagellate setae as far
forward as abdominal segment II, mostly about 17.5—
25.0 um long on abdominal segment VIII, those on
anterior segments each about 15 tm long, very slen-
der. Minute swollen setae present, each about 3.75 um
long, sparse, mingled with the flagellate setae on
abdominal segments I-IV. Setae anterior to abdomi-
nal segment II mostly thick and flagellate, 25-50 um
long, not numerous.A few minute swollen setae present
on head, each about 3.5 um long, slightly narrower
than those on abdomen.

Ventral surface with similar setae to those on dor-
sum, crowded as far forward as abdominal segment II,
with a few minute swollen setae on abdominal seg-
ments II-VI. Anteriorly a few thicker flagellate setae
present as on dorsum, and a few minute swollen setae
also present on head and around anterior spiracles.

Second instar (Fig. 20)

Body pyriform, 1.05—1.10 mm long, 0.45—0.58 mm
wide, widest at about mesothorax, tapering abruptly to
abdominal segment IV then gradually to posterior end
of body; base of abdominal segment VIII 75—92 Um
wide; posterior end of body projecting beyond actual
anal lobes, each lobe with | dorsal and 2 stout ventral
setae each 375-470 ttm long. Antennae each 290—
350 um long, with 2 segments; first 40-50 Um long,
second curved, with setae of various lengths, longest at
distal end about 100 Lm long. Legs well developed;
hind trochanter + femur 155—195 um long, hind tibia +
tarsus 157.5—185.0 um long, claw slender, about 22.5
uum long. Ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 0.94—1.04. Ratio of lengths of hind
tibia to tarsus 1.03—1.25. First pair of legs longer,
trochanter + femur 170-200 tm long, tibia + tarsus
185—207 um long. Labium broad, 105—115 tm long,
longer than clypeolabral shield. Circulus present within
borders of abdominal segment III, 7.50-11.25 Um in
diameter, deeply cupped. Anal ring 35-40 um wide,
with 6 setae; 2 anterior pairs slender, each about 75 um
long, posterior pair stout but not so stout as anal lobe
setae, each about 350 um long.

Dorsal surface with long stout setae 95-120 um
long at posterior edges of abdominal segments I[I—
VII: other abdominal setae, each about 25 Lm long,
present on abdominal segment VIII, and others, each
about 20 tum long, at anterior edges of abdominal
segments IV—VII; slenderer setae on these segments
mostly about 15 um long. Other setae on head, thorax
and abdominal segments I-III, each 20-55 tum long,
not numerous. Minute clavate setae present on ab-
dominal segments II-IV, and others, about same size
but with smaller collars, present on head and lateral
area of metathorax.

Ventral surface with similar setae to those on dor-
sum. Abdominal segments V and VI each with a few
elongate sensory setae; abdominal segments III and IV

each with a few shorter sensory setae. Minute clavate
setae present anterior to each spiracle and posterior to
each second spiracle.

COMMENTS. Material from Baroalba Spring pos-
sesses legs and antennae slightly longer than that from
Sawcut Gorge but the proportions of the segments are
about the same. At present, specimens from both areas
are treated here as conspecific.

Female pupa (Figs 1B, 21A)

Body membranous, elongate-pyriform, 0.90—0.99 mm
long, 0.38-0.43 mm wide, widest at mesothorax, ab-
domen gently tapering, posterior end projecting only
slightly, segmentation distinct on abdomen. Antennae
situated on dorsal margin, 2-segmented, curved, 262—
270 um long, apex rounded. Legs tapering to pointed
developing claws, 325-390 um long, with faint seg-
mentation. Labium distinct, 100—120 um long, longer
than clypeolabral shield. Anal ring dorsal, at apex of
abdomen. Spiracles present.

COMMENTS. Specimens are available showing the
adult female inside, almost ready to emerge (Fig. 1B).

Adult Male (Fig. 22)

Body of adult male elongate-pyriform, 104 Um long,
40 um wide, widest at mesothorax, head and thorax
rounded, tapering abruptly to anterior end of abdomi-
nal segment IV, remainder of abdomen narrow, tapering
gradually, base of abdominal segment VII 90 um wide.
Apparent anal lobes each with | stout dorsal seta and
2 ventral setae, each 200-250 ttm long, forming a
group of 3. Antennae 2-segmented, 310—400 tm long;
second segment curved, 270-350 1m long, longest
setae about 90 um long. Legs well developed, slender;
hind trochanter + femur 155—175 um long; hind tibia
+ tarsus 147.5— 175.0 um long; claw slender, 25 um
long, 5 um wide at base. Ratio of lengths of hind tibia
+ femur to trochanter + femur 0.94-1.01. Ratio of
lengths of hind tibia to tarsus 1.03—1.33. First legs
longer; trochanter + femur 180-210 Lim long, tibia +
tarsus 180-200 um long. Mouthparts represented by
tentorium, remains of clypeolabral shield with a few
setae, and a small vestigial labium. Genital capsule
almost triangular, 65 um wide, length about the same,
venter of penial sheath rounded apically, ventral slit
rounded near base then widening slightly apically;
basal ridge of penial sheath well defined ventrally;
aedeagus long and pointed, 250—262 1m long. Dorsal
aspect with an apparent anal ring and anal ring setae at
posterior edge of abdominal segment VIII; 2 anterior
pairs of setae each about 45 um long, posterior pair
thicker, each about 58 Lim long, situated near curved
ends of anal ring and lateral to large, almost triangular
anus, about 30 um wide. Dorsal part of capsule ex-
tending from anus as a slender elongate style, about
125 um long, shorter than aedeagus, slightly expanded

apically then pointed, tip bearing a few spicule-like
extensions; base of style with 2 pairs of short setae.

Dorsal surface of body with short, flagellate setae,
each about 12.5—15.0 um long, on abdominal segment
VIII. Anteriorly on all segments including head, with
transverse rows of long, slender flagellate setae 62—
70 um long, these sometimes absent; posterior
abdominal segments also with a few short flagellate
setae. Minute sensory setae, each with slightly swollen
tip and scarcely longer than a setal collar, present in
moderate numbers across segments anterior to ab-
dominal segment VIII.

Ventral surface with similar distribution of setae to
those on dorsum, the long flagellate setae sometimes
absent. Minute sensory setae present as on dorsum but
fewer on thorax.

Male prepupa (Fig. 21B)

Body elongate-pyriform, about 0.90 mm long, 0.40 mm
wide at mesothorax; abdomen with distinct segmenta-
tion, tapering to rounded apex. Antennae situated on
dorsal margin, 2-segmented, tapering, apex rounded,
220-240 um long. Legs 305—330 um long, segmented,
developing claws pointed. Anal ring present at apex of
abdomen. Mouthparts represented by remains of
clypeolabral shield and small lobes of labium. Spiracles
distinct.

COMMENTS. Specimens are available showing the
developing pupa inside.

Male pupa (Fig. 21C)

Body narrowly pyriform, 1.00—1.27 mm long, 0.40-
0.43 mm wide, widest at mesothorax, narrowing to
abdominal segment V then tapering to developing
genital capsule, posterior end elongate, pointed; seg-
ments distinct on abdomen. Antennae tapering to
pointed distal end, 285-300 tm long, situated on
dorsal head margin. Legs tapering to pointed claws,
320-360 um long. Mouthparts represented by indis-
tinct clypeolabral shield. Spiracles present. Anal ring
situated dorsally towards anterior end of abdominal
segment VIII.

COMMENTS. Some specimens available show the
adult male complete, almost ready to emerge (Fig. 1A).

MATERIAL

HOLOTYPE. Adult 9, Australia: Northern Territory,
Sawcut Gorge, 12°55’S, 132°56'E, associated with
Acropyga sp., 19.xi.1972 (R.W. Taylor) (ANIC).

PARATYPES. Australia: same data as holotype, 1
adult 2, 2 adult bo' (ANIC), 1 2nd instar, 2 éo' pupae,
1 2 pupa (ANIC), | adult 9, 3 adult do’, 1 3 pupa
(BMNH): same data but 13.vi.1973, 1 adult &, 4 2nd
instars, 2 prepupae do’, | pupa o’, 2 pupae 9? (ANIC):
2 2nd instars, 1 prepupac' (BMNH). Baroalba Spring,

D.J. WILLIAMS

12°47’S, 132°51’E, with Acropyga sp., 17.xi.1972
(R.W. Taylor), 1 adult o', 7 2nd instars, 7 pupae 9
(ANIC), | 2nd instar, 3 pupae 99 (BMNH); same data
but 17.x1.1971, 1 adult @ (ANIC).

COMMENTS. In possessing a general distribution of
short crowded setae in the adult female, only as far
forward as abdominal segment II, this species closely
resembles E. recalvus sp. n. In E. taylori, however, all
the dorsal setae on the head, thorax and first abdominal
segment are flagellate except for a few minute swollen
sensory setae. In E. recalvus, all the dorsal setae at the
anterior end of the body are sensory and bluntly tipped
or slightly swollen. Furthermore, the outer setae on the
legs of E. taylori are flagellate, whereas those of E.
recalvus are sensory.

There are available some second instars from all
localities. These differ from the adult female in lack-
ing the wide bands of dense setae across the segments
and instead there are long slender setae, similar to
those of the adult male.

Adult males appear to be of two forms, with or
without long flagellate body setae. Most of the adult
males from Sawcut Gorge possess the long setae ex-
cept in one specimen. One of the adult males from
Baroalba Spring also lacks these long setae as does the
specimen still enclosed within the pupal instar. All
specimens are here regarded as representing the same
species. Such wide differences occur also in adult
females of Molluscococcus fibrillae Hall, redescribed
by Miller and Williams (1995). This species normally
possesses extremely long setae, 812-928 um long
over most of the dorsum, but there are also specimens
with short setae only, 44-61 um long.

The only adult females available for study from
Baroalba Spring are still within the pupal instar and
they appear to be identical with adult females from
Sawcut Gorge. If there is evidence in future that the
two forms of adult male discussed here, represent two
distinct species, then the matter can easily be cor-
rected.

The species is named after the collector, R.W. Taylor,
CSIRO, Canberra, Australia, who sent many of the
mealybugs discussed in this work.

Xenococcus Silvestri

Xenococcus Silvestri, 1924: 312; Williams, 1978: 63.

TYPE SPECIES. Xenococcus annandalei Silvestri, by
original designation and monotypy.

DESCRIPTION

Body of adult female broadly oval, membranous, ab-
domen tapering abruptly to narrow sclerotized apical
segment. Anal lobes not developed, recognizable by
inner ventral grooves, position of each lobe with a
group of 3 long, stout setae, 2 ventral and | dorsal.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 23

Anal ring protruding between anal lobes, represented
by crescentic dorsal band without cells, with 8 anal
ring setae, the 2 anterior pairs of setae slender, the third
pair thicker and longer, detached from ring, the fourth
pair ventral in position, about as long as anal lobe
setae. Antennae 4-segmented, placed on dorsal mar-
gin, tapering, about as long as body with strong
articulation between first and second segments. Legs
well developed, long and slender; claw elongate, slen-
der. Labium elongate, longer than wide, setae on upper
surface well spaced. Circuli present, round and
shallowly cupped. Dorsal body setae minute and abun-
dant, extending to lateral ventral margins on thorax.
Ventral setae mostly long and stouter. Sickle-shaped
setae usually present on thorax. Eyes absent. Ostioles
absent. Pores and ducts absent.

COMMENTS. In life, the abdomen of Xenococcus
curls slightly to the dorsum as in Eumyrmococcus. The
genus differs from Eumyrmococcus in possessing very
long antennae with well developed articulation be-
tween the first and second segments. Although the
abdomen tapers, it narrows abruptly to a narrow ab-
dominal segment VIII and the body is widest at about
the metathorax. In Eumyrmococcus, the whole
cephalothorax is dilated and the abdomen tapers gradu-
ally. Furthermore, only the dorsal abdominal setae in
Xenococcus are short and crowded, those on the venter
of the abdomen, although numerous, are long and
stouter, similar to the medial ventral setae on the head
and thorax. The ventral abdominal setae in
Eumyrmococcus are always short and crowded, simi-
lar to the dorsal abdominal setae.

At present, two species are recognized, always as-
sociated with the ant genus Acropyga. The female
possesses a pupal instar and the adult male of one of
the species is described on p. 24.

Key to Species of Xenococcus (Adult
Females)

1 Legs with tibia shorter than tarsus. Antennae each about
860-1090 Lm long ou... ee eeeeeeeeeeeee acropygae (p. 23)

— Legs with tibia about twice as long as tarsus. Antennae
about 1480-1550 um long............... annandalei (p. 25)

Xenococcus acropygae sp. n.

(Figs 23-26)

Xenococcus annandalei Silvestri, Williams, 1985: 390;
Williams and Watson, 1988: 221 (mis-identifica-
tions).

DESCRIPTION

Adult female (Fig. 23) on microscope slide pyriform,
narrowing at abdominal segment VII; 1.27—1.65 mm
long, 0.74—1.08 mm wide, widest at about metathorax;

body membranous except for sclerotized abdominal
segment VIII, posterior end of body projecting beyond
anal lobes; base of abdominal segment VIII 180-
220 um wide; anal lobes each with | dorsal and 2
ventral setae, all stout, forming a group of 3, each 875—
920 um long. Antennae conspicuous, each
860-1090 um long, with 4 segments; segment | 170-
230 um long (type series 90-120), segment 2
280-300 tum long (type series 330-360), segment 3
90-130 um long (type series 90-120), segment 4 200-
300 um long (type series 240-300); segment | 170-230
lim wide, segments progressively narrowerer to seg-
ment 4 60-100 tum wide, segment 4 tapering. All
antennal segments with long stout setae, segment 2
with about 30 setae on ventral surface; longest setae on
segment 4 400 Um long, setal bases 3—5 um wide with
correspondingly wide setal collars. Segments | and 2
strongly articulated with grooves at distal end of seg-
ment | and minute projections at proximal end of
segment 2. Tip of segment 4 with a pair of peg-like
setae each 10.0—12.5 um long. Legs well developed;
hind trochanter + femur 270-380 um long (type series
290-340), hind tibia + tarsus 280-415 um long (type
series 310-360), claw slender about 35—45 um long.
Ratio of lengths of hind tibia + tarsus to hind tro-
chanter + femur 1.01—1.13. Ratio of lengths of hind
tibia to tarsus 0.66—0.95; tarsus always longer than
tibia. Hind tarsus swollen then narrowing distally. All
legs with long stout setae, the longest at distal end of
femur, 250-300 um long. Labium 165-220 um long,
longer than clypeolabral shield, setae well separated.
Circuli normally numbering 2 within borders of ab-
dominal segments If and II, each 27.5—47.0 ttm in
diameter, with rim projecting from surrounding derm,
inner part cup-shaped, shallow. Occasionally a small
third circulus present on abdominal segment IV. Anal
ring 77.5—85.0 um wide with 8 setae; 2 anterior pairs
slender, each about 130 um long, 2 posterior pairs
stout, each about as long as anal lobe setae, | pair on
dorsum and the posteriormost pair usually placed on
venter.

Dorsal surface densely covered with slender setae
except in intersegmental areas and on head, mostly
curved, each about 15 tum long towards anterior edges
of segments, and about 20 um long towards posterior
edges of segments; a few longer setae, each about
25 um long with larger collars, present across all
segments; abdominal segment VIII with fewer but
longer setae. Sickle-shaped setae usually present on
thorax and anterior abdominal segments but some-
times absent entirely, each with slender tip. Setae near
each antennal base slender, stouter on head margin.

Ventral surface with stout setae on abdomen and
medial areas of head and thorax, 40-165 «1m long
except for some on margins of abdominal segment
VIII, each about 300 um long. Short slender setae as
on dorsum, present around margins of anterior ab-

dominal segments and thorax. A few sickle-shaped
setae usually present near margin of thorax.

COMMENTS. This species differs from X, annan-
dalei mainly in the lengths and ratios of the tibiae and
tarsi. In X. acropygae, the tarsus is longer than the
tibia but in X. annandalei, the tibia is about twice as
long as the tarsus. Furthermore, the setal bases on the
antennae of A. acropygae are mostly narrower than
those of X. annandalei. Moreover, the antennae and
legs of A. acropygae are shorter than those of A.
annandalei.

First instar (sex not determined) (Fig. 24)

COMMENTS. ‘This instar was described by Williams
(1978) from specimens collected in India, Mysore, on
coconut roots under the name X. annandalei. Speci-
mens from southern Asia and Australasia agree with
these first instars but show wider variation. The body
varies from 0.70—1.05 mm long and 0.27-—0.58 mm
wide. Antennae 4-segmented, 740-880 um long, with
long stout setae, the longest on segment 4 about
250 um long. Hind trochanter + femur 175—250 um
long, hind tibia + tarsus 215-270 um long. Ratio of
lengths of hind tibia + tarsus to trochanter + femur
1.08-1.27. Ratio of lengths of hind tibia to tarsus
0.58—0.74, tarsus always noticeably longer than tibia.
All segments with long stout setae, longest at distal
end of femur about 160 um long. Anal ring with 4
pairs of setae; 2 anterior pairs slender, each about 60—
110 um long, third pair stouter, 295-360 Um long,
fourth pair stoutest, 535-750 um long, resembling
the anal lobe setae. Anal lobe setae, stout, usually
curled when prepared on slides, 675—700 Lm long,
forming a group of 3 on each anal lobe, | on venter
and 2 on dorsum. Body setae short and curved on
head and thorax, each 12—15 tum long, accompanied
by a few sickle-shaped setae on dorsum. Other body
setae 40-88 um long except on margins where they
are 100-120 um long.

A new illustration has been prepared for this work
based on specimens collected at the type locality,
Sulawesi.

Third-instar female (pupa) (Fig. 25A)

Body broadly oval, 0.84—0.90 mm long, 0.55—0.67 mm
wide, membranous, with only faint signs of segmenta-
tion at posterior end of abdomen. Antennae dorsal,
660-760 tm long, tapering. Legs tapering, 425—
530 um long, segmentation faint. Labium distinct,
135-145 um long, longer than clypeolabral shield.
Spiracles distinct.

COMMENTS. The female pupal instar is always rec-
ognizable by the well developed labium and the long
dorsal antennae. Some specimens available have the
developing adult female still within the pupal instar.

D.J. WILLIAMS

Adult male (Fig. 26)

Body normally curved ventrally and distorted when
prepared on microscope slides. When flattened,
broadly oval, about 1.0 mm long, 0.70 mm wide,
sides sub-parallel, widest at about abdominal seg-
ments II and V. Head and abdominal segment I
sclerotized dorsally except at anterior end surround-
ing antennae; ventrally, sclerotized laterally on thorax.
Antennae situated on dorsal membranous area, each
1-segmented, tubercle-like, with 6 stout setae, each
30-35 um long. Legs placed well anterior on body, of
an unusual shape, squat and robust. Coxae large. Tro-
chanter + femur stout, about 70 Lim long, fused except
for faint line apparent in some specimens, trochanter
with the usual 2 pairs of sensory pores. Tibia + tarsus
much narrower, tapering, about 75 um long. Claw
about 20 Lim long, unusual, with well developed wide
base, actual claw pointed and slender with apparently
a pair of stout blunt digitules. Genital capsule about
290 um long, 110 um wide, mostly internal, the
external venter of penial sheath about 170 um long;
ventral slit indistinct, at distal end of sheath; tip of
sheath with a few minute setae. Aedeagus strongly
sclerotized, elongate. Base of genital capsule dorsally
with a pair of lobes, each with a group of 3 stout
setae, 30-37 ttm long, and usually 4 short setae, the
lobes situated on each side of a minute projecting
plate; anus, when viewed laterally, opening under the
plate; entire area normally lying between the genital
capsule and dorsal surface of abdomen when flat-
tened on slide.

Body setae of 2 types.A band of setae present across
anterior edges of venter of abdominal segments II and
Ill, each seta minute and shorter than diameter of
heavily sclerotized collar. A minute type of seta with
membranous collar, present across dorsal and ventral
segments; setae in a row near ventral edge of abdomi-
nal segment IV either with membranous or sclerotized
collars.

Third-instar male (prepupa) (Fig. 25B)

Body almost rotund, 0.78—0.86 mm long, 0.66—
0.70 mm wide. Segmentation discernible on dorsum
and venter of abdomen. Antennae dorsal, tapering to
blunt apex, 185-260 um long. Legs 180-240 um long,
tapering, with faint segmentation. Remains of
clypeolabral shield and labium present. Spiracles well
developed.

COMMENTS. The prepupal instar is recognizable by
the dorsal developing antennae, only about one-third
the length of those of the female pupa. There are faint
signs of the labium and clypeolabral shield.

Fourth-instar male (pupa) (Fig. 25C)
Body broadly oval, 0.86—0.98 mm long, 0.65—0.75 mm
wide, with signs of segmentation on abdomen. Anten-

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI Pa)

nae reduced to small triangular flaps on venter of head,
25-30 uum long, 50-65 tum wide. Legs 155-200 um
long, stout and tapering to sclerotized points or devel-
oping claws, segmentation not discernible. Small area
of clypeolabral shield present only. Spiracles distinct.

COMMENTS. In the male pupa the legs are usually
shorter than those of the prepupa but the lengths
sometimes overlap. The most distinctive pupal charac-
ter is the shape of the antennae which are short,
triangular and present on the anterior edge of the head
to contain the short developing antennae which in the
adult male lie dorsally.

MATERIAL

HOLOTYPE. Adult 2, Indonesia, Sulawesi, Sulawesi
Utara, Dumoga-Bone National Park, Toraut forest,
with Acropyga (Acropyga) acutiventris Roger, 19.111.
1985 (R.H. Disney) (BMNH).

PARATYPES. Indonesia, Sulawesi, same data as
holotype but (R.H. Disney or R.H. Disney & J.H.
Martin), 3 adult 99 (BMNH), 10 adult éo&' (BMNH), |
adults’ (ANIC), | adulto' (MNHN), | adulto’(USNM),
18 Ist instars, 24 pupae 92, 4 prepupae do’, 3 pupae do
(BMNH).

Non-type material (all 99). Indonesia, Krakatau,
Anak Krakatau, Rakata (in mandibles of A.
acutiventris). India, Mysore, on roots of Cocos
nucifera, 21.vi.1937 (T.V. Subramanian) (first instar);
Karnataka, Bangalore, on roots of Vitis vinifera,
18.1.1996 (A. Virakramath & B.K. Rajagopal). Singa-
pore; Botanical Gardens, in soil under wood, with A.
acutiventris, 21.v.1968. Philippine Islands, Palawan
(spirit material). Malaysia, Sabah, Tawau, Quoin
Hill, with A. acutiventris, 10.vi.1968. Papua New
Guinea, Morobe Province, 16 km N.W. Lae, in soil
under wood, with A. acutiventris, 12.vi.1972 (R.W.
Taylor); Wau, McAdam Park, in rotting log, with A.
acutiventris, 14.vi.1972 (R.W. Taylor); Bulolo (spirit
material, with A. acutiventris and A. (Atopodon)
ambigua): East Sepik Province, Angoram, from rot-
ting log, 2.vi.1972 (R.W. Taylor); Yawasora, nr
Wewak, from rotting log, 4—6.vii.1972 (R.W. Taylor);
Hayfield, nr Maprik, in soil under wood (all with A.
acutiventris): West Sepik Province, Pes, nr Aitape, in
soil under wood with A. acutiventris, 8—9.vi1.1972:
Northern Province, Kokoda, associated with dying
Theobroma cacao, ix.1990 (G. Lockwood); nr
Kokoda, in soil under wood, with A. acutiventris,
l.vi.1972 (R.W. Taylor). Solomon Islands, San
Cristobal, Guadalcanal, Nggela (spirit material), with
A. acutiventris and A. (Rhizomyrma) lauta Mann.
Australia, Queensland, Iron Range, 12°42’S
143°18’E, 9-15.vi.1971 (R.W. Taylor & J. Feehan);
Waugh, from rotten log, 11.vi.1962 (R.W. Taylor);
Cape York Peninsular, Bemaga, from mandibles,
xii.1983 (J. Sedlack); Finch Hatton Gorge, 21°05’S

148°38’E, 11.xi.1976 (R.W. Taylor & TA. Weir);
Josephine Falls (spirit material): Northern Territory,
Baroalba Spring, 12°47’S 132°51’E, in soil under
wood, 16, 17, 20.x1.1972, 13.vi.1973 (all with A.
acutiventris).

COMMENTS. Specimens recorded under the name X.
annandalei from Hong Kong, Vietnam and Penang by
Williams (1978) are probably this species and await
verification.

Xenococcus annandalei Silvestri

(Fig. 27)

Xenococcus annandalei Silvestri, 1924: 312; 1926:
275; 1927: 253. LECTOTYPE 2, INDIA: Barkuda
I. [Chilka (Chilika) Lake, Madras District (now
Orissa Province), on roots of Ficus obtusa, with
antsAcropyga acutiventris Roger] (IEAUN, Portic1)
designated by Williams 1978: 66 [examined].

COMMENTS. Williams (1978) described the adult
female in detail and the illustration is reproduced here
with slight modification. Some further notes are now
added to distinguish the species from X. acropygae.

The antennae are 1480-1550 [tm long, longer than
in any specimen of X. acropygae so far studied. In X.
acropygae, the range is 860-1090 um long. Further-
more, although the distribution of the antennal setae
are about the same in the two species, there are many
more shorter and more slender setae in X. annandalei.
The bases of the setae in the antennae of X. annandalei
vary in width from 2.5 um to 7.5 um with a corre-
sponding difference in size of the setal collars, whereas
in X. acropygae there is less variation, with the width
of the setal bases 3—5 tm. The legs of X. annandalei
differ from those of X. acropygae. In X. annandalei,
they are longer, with the hind trochanter + femur
440-480 um long and the hind tibia + tarsus 530-560
uum long. The ratio of lengths of hind tibia + tarsus to
those of the hind trochanter + femur is 1.16—1.18 and
the most striking difference is the ratio of the hind
tibia to tarsus, with the tibia always about twice as
long as the tarsus. Besides, the tibiae and tarsi are
fairly uniform in width for most of their lengths,
although the tarsi narrow and taper distally. In X.
acropygae, the tarsus is always longer than the tibia
and the tibia is swollen before narrowing abruptly to
a long slender distal end.

The short flat setae on the dorsum discussed by
Williams (1978) are, in fact, sickle-shaped in profile,
similar to those of X. acropygae.

The third instar discussed by Williams (1978) is
now known to be the second instar (Williams, 1988). It
is also evident that the first instar discussed by Williams
(1978) from Mysore, on coconut roots, is the first
instar of X. acropygae.

MATERIAL

At present the species is only known from India,
Orissa Province, Barkuda I., on roots of Ficus obtusa
and F: religiosa.

REVIEW OF GENERA IN THE
RHIZOECINAE

In addition to Eumyrmococcus and Xenococcus, al-
ready discussed, the following genera have been
included in the subfamily at one time or another and
their current status is discussed.

Tang (1992) erected the subtribes Rhizoecina,
Prorhizoecina and Pseudorhizoecina for some of the
genera, but this action seems unnecessary. These
names, nevertheless, remain available.

The opportunity is taken to describe a new genus
from southern Asia because it resembles
Eumyrmococcus superficially but is clearly more re-
lated to Rhizoecus.

Brevicoccus Hambleton

Brevicoccus Hambleton, 1946a: 10; Williams &
Granara de Willink, 1992: 68. Type species:
Brevicoccus clavisetosus Hambleton, by original
designation and monotypy.

DIAGNOsIS. Body broadly oval, anal lobes poorly
developed, each without any development of long
setae. Antennae placed close together, strongly genicu-
late, 4-segmented, last segment triangular. Legs well
developed; claws long and narrow, Anal ring fairly
simple, with a few cells and 8—16 short knobbed setae.
Ostioles present. Eyes absent. Cephalic plate present.
Body setae short, clavate. Trilocular pores present.
Multilocular disc pores present on venter. Minute oval
disc pores present.

COMMENTS. The characters of the anal ring and
antennae are distinct and the genus is recognized by
most workers as belonging to the Rhizoecinae. This is
a monotypic genus, known only from Brazil, and the
single species is reported to live on roots of Gramineae
and Cyperaceae in ant tunnels.

Capitisetella Hambleton

Capitisetella Hambleton, 1977: 40; Williams &
Granara de Willink, 1992: 71. Type species:
Pseudorhizoecus migrans Green, by original des-
ignation and monotypy.

DIAGNOSIS. Body almost pyriform, abdomen con-

D.J. WILLIAMS

stricted slightly near posterior end. Anal lobes not
developed, without differentiated anal lobe setae. An-
tennae placed fairly wide apart, 3-segmented, the third
segment tapering, antennal setae clavate. Legs stout,
with capitate setae, claws elongate. Anal ring heavily
sclerotized, with a few elongate cells and 6 setae.
Ostioles absent. Cephalic plate absent. Eyes absent.
Body setae all capitate. Trilocular pores present. Mul-
tilocular pores absent. Large granular discoidal pores
present at posterior end of body.

COMMENTS. The combination of a constriction at
the posterior end of the body, reduction of antennal
segments to three, the capitate body setae and the
absence of ostioles, link this South American genus
with Eumyrmococcus. In Capitisetella, there are a few
elongate cells on the anal ring and these are absent in
Eumyrmococcus. Furthermore, Capitisetella lacks the
abundant setae on the abdomen, present in
Eumyrmococcus or even in Neochavesia, also possi-
bly related to Capitisetella. The single species lives in
association with Acropyga (Rhizomyrma) paramari-
bensis (Biinzli, 1935). At present, the genus is accepted
in the Rhizoecinae.

Geococcus Green

Geococcus Green, 1902: 262; Williams, 1969b: 508.
Type species: Geococcus radicum Green, by origi-
nal designation.

DIAGNOSIS. Body elongate to oval, anal lobes well
developed, sclerotized, each terminating in a stout,
spine-like seta. Antennae geniculate, 6-segmented.
Legs well developed, inner edges of tibia and tarsus
with stout setae, claws long and slender. Anal ring
with large cells and 6 setae. Ostioles present. Circuli
present, raised. Cephalic plate present. Multilocular
disc pores present. Trilocular pores present. Large
trilocular pores present, usually much larger than the
normal trilocular pores. Body setae fairly abundant,
short and flagellate.

COMMENTS. The genus is widely accepted in the
Rhizoecinae. At present six species are assigned to the
genus from South-East Asia, southern Asia and the
Indo-Australian Region but there are many species
awaiting to be described. The genus is well known for
the spread of G. coffeae throughout much of the trop-
ics.Although this species was described from Surinam,
where it is associated with Acropyga (Rhizomyrma)
paramaribensis on coffee roots and other plants, it is
undoubtedly native to southern Asia.

The peculiar large trilocular pores, always present,
may be modified tritubular cerores. It is often difficult
to study the anal ring because it is concealed between
the sclerotized anal lobes.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 27

Leptorhizoecus gen. nov.

TYPE SPECIES. Leptorhizoecus deharvengi sp. n.

DESCRIPTION

Body membranous, elongate, with dilated
cephalothorax, abdomen narrow, subparallel, narrow-
ing abruptly to abdominal segment VIII; posterior end
of body sclerotized, rounded. Anal lobes not devel-
oped; setae on positions of anal lobes not differentiated
from other setae on abdominal segment VIII. Anal ring
ventral, crescentic, with a single row of minute elon-
gate cells and 6 spine-like setae. Antennae placed
close together on venter of head margin, 6-segmented.
Legs well developed; with thick pointed setae on inner
edge of tibia and tarsus. Claw elongate and slender.
Labium narrow, longer than wide. Ostioles and circu-
lus present. Body setae flagellate, mostly short and
fairly numerous. Trilocular pores present on dorsum
and venter. Quadrilocular pores present next to circu-
lus. Tubular ducts absent.

COMMENTS. This genus is typically rhizoecine but
the body shape resembles species of Eumyrmococcus
in exhibiting a dilated cephalothorax. It differs from
Eumyrmococcus in possessing 6-segmented antennae,
placed close together, and in having ostioles and trilocu-
lar pores. The ventral anal ring is unlike any so far
described in the Rhizoecinae.

Leptorhizoecus is probably intermediate between
the Rhizoecus and Eumyrmococcus groups and the
shape suggests it is possibly attended by ants.

The name Leptorhizoecus is based on the Greek
word ‘Leptos’, meaning fine and delicate, combined
with the extant genus name Rhizoecus.

Leptorhizoecus deharvengi sp. n.
Fig. 28.

DESCRIPTION

Appearance in life not recorded. Adult female on
microscope slide elongate, cephalothorax dilated, ab-
domen subparallel, widening slightly to abdominal
segment VII, narrowing abruptly to rounded posterior
end, body membranous except for rounded posterior
half of apical segment of abdomen; 0.80-0.92 mm
long, 0.27—0.36 mm wide; anal lobes not developed,
anal lobe setae not differentiated from others on apical
segment. Antennae placed close together on ventral
head margin, each 115—135 um long, with 6 segments;
falcate setae well developed on segments 5 and 6. Legs
well developed; hind trochanter + femur about 92.5
tum long, hind tibia + tarsus 97.5—100.0 um long, claw
slender, elongate and slightly curved, about 27.5 um
long. Ratio of lengths of hind tibia + tarsus to hind
trochanter + femur 1.05—1.08. Ratio of lengths of hind
tibia to tarsus 0.97—1.00. Leg setae flagellate, placed
well apart, inner edges of tibia and tarsus with thicker

setae. Labium 77.5-82.0 um long, about 37.5 um
wide, ratio of length to width 2.06—2.18; longer than
clypeolabral shield. Clypeolabral shield with 2 pairs
of setae. Circulus slightly oval, about 17.5 um wide,
situated within borders of abdominal segment IJ; struc-
ture difficult to determine in available material. Ostioles
present, posterior pair fairly well developed but with-
out associated setae; anterior pair represented by
indistinct slits. Anal ring ventral in position, oval to
triangular, 25—30 Lim wide, with a single row of minute
elongate cells in posterior arc, accompanied by 6 stout,
elongate-conical setae, each 17.5 um long (cells and
setae actually anterior if ring transferred to dorsum).
Eyes present.

Dorsal surface with long setae, each about 60 um
long, on sclerotized apical segment of abdomen; shorter
setae, each about 25 um long, situated at anterior end
of apical segment. Anteriorly on abdomen, setae all
about 25 um long, in moderate numbers in well de-
fined bands across middle of segments leaving bare
intersegmental areas. Shorter setae, each 12.5—18.0 um
long, present on head and thorax. All setae flagellate.
Trilocular pores present, not numerous, evenly distrib-
uted among the setae.

Ventral surface with similar distribution of setae to
those on dorsum. Multilocular disc pores represented
by a few quadrilocular pores, near circulus, each about
5 um in diameter. Trilocular pores scattered, not so
numerous as on dorsum.

MATERIAL

HOLOTYPE. Adult 2, Indonesia, Sumatra, Rantau
Pandan, in soil of Hevea sp., 9.vi.1991 (L. Deharveng
& A. Bedos) (MNHN).

PARATYPE. Indonesia, Sumatra, same data as
holotype. | adult ? (BMNH).

COMMENTS. The species is named after Louis
Deharveng, Université Paul Sabatier, Toulouse, one of
the collectors. Dr Deharveng has sent interesting sam-
ples from southern Asia collected from Berlese funnel
apparatus.

Neochavesia Williams & Granara de Willink

Chavesia Balachowsky, 1957: 158; Beardsley, 1970: .
[Preoccupied by Chavesia Dollfus, 1889, Isopoda].

Neochavesia Williams & Granara de Willink, 1992:
232 [replacement name]. Type species Chavesia
caldasiae Balachowsky, by original designation.

DIAGNOSIS. Body in life with tip of abdomen curled
to dorsum, scorpion-like. Slide-mounted specimens
with cephalothorax dilated, abdomen narrowing to a
pair of protuberant, rounded anal lobes bearing many
long flagellate setae. Anal ring situated at base of anal
lobes, simple, with 6 or a few setae around anterior

sclerotized half, without cells. Antennae each with 4 or
5 segments. Legs well developed, tarsi tapering, claws
long and slender. Ostioles absent. Circuli conical, each
with centre cupped. Labium with 3 pairs of subapical
setae. Eyes absent. Body setae short and abundant on
cephalothorax, longer and in bands across abdominal
segments. Trilocular pores present in type species
only, each with a minute internal filament arising from
centre of pore. Multilocular disc pores and tubular
ducts absent.

COMMENTS. In the strong development of the
cephalothorax, the simple anal ring without cells, the
absence of ostioles, and the presence of cupped circuli,
this South American genus comes closest to
Eumyrmococcus and Xenococcus. Many species origi-
nally and provisionally identified as Eumyrmococcus
in South America, associated with ants of the genus
Acropyga, refer to Neochavesia. The genus is here
accepted as belonging to the Rhizoecinae.

Prorhizoecus Miller & McKenzie

Prorhizoecus Miller & McKenzie, 1971: 583. Type
species Prorhizoecus atopoporus Miller &
McKenzie, by original designation and monotypy.

DIAGNOSIS. Body oval, anal lobes not developed;
positions of each anal lobe with a single normal apical
seta. Antennae each with 4 or 5 segments. Legs well
developed, slender, hind coxae with translucent pores.
Anal ring heavily sclerotized with thick rim, cells, and
6 setae. Cerarii present on anal lobes, each cerarius
with 2 conical setae and a group of trilocular pores.
Ostioles present. Eyes present. Body setae short and
abundant. Trilocular pores present. Multilocular disc
pores present, each with 12 loculi. Tubular ducts
present, each with narrow oral rim and heavily
sclerotized bulbous duct.

COMMENTS. Although the antennae resemble those
of Rhizoecus, the other characters, cerarii, translucent
pores on the hind coxae, an anal ring with a broad rim,
and peculier oral rim ducts, exclude this genus from
the Rhizoecinae. The single species is known only
from Mexico and feeds on grass roots. The genus may
be related to Cryptoripersia Cockerell or Syrmococcus
Ferris.

Pseudorhizoecus Green

Pseudorhizoecus Green, 1933: 55; Hambleton, 1977:
38; Williams & Granara de Willink, 1992: 463.
Type species Pseudorhizoecus proximus Green, by
original designation.

DIAGNOSIS. Body rotund; anal lobes not developed,
without differentiated anal lobe setae. Antennae placed
close together, each with 5 segments, tapering. Legs
well developed, tarsi tapering, claws long and slender.

D.J. WILLIAMS

Anal ring terminal, wider than long, irregularly out-
lined, with 2—5 short setae, a few small cells, and
numerous protuberances. Ostioles, eyes and cephalic
plate absent. Body setae abundant, short and flagel-
late. Trilocular pores present.

COMMENTS. This unusual genus is probably related
to Capitisetella but the anal ring possesses numerous
protuberances and all the body setae are flagellate. The
only included species has a fairly wide distribution in
Central and South America and is associated with
Acropyga (Rhizomyrma) rutgersi and A. paramarib-
ensis.

The adult male is morphologically degenerate, with-
out wings, resembling the adult female in body shape
and possessing similar 5-segmented antennae.
Beardsley (1970) remarked that the genitalia appeared
to be similar to the genitalia of Capitistella and
Neochavesia.

Pygmaeococcus McKenzie

Pygmaeococcus McKenzie, 1960: 741. Type species
Pygmaeococcus morrisoni McKenzie, by original
designation and monotypy.

DIAGNOSIS. Body of adult female minute, elongate,
anal lobes not developed, position of each lobe with |
long ventral and 2 long dorsal setae, forming a group
of 3. Antennae each with 5 segments. Legs well devel-
oped, with stout, spinose setae present on inner edges
of tibiae and tarsi; claw long and slender, digitules
clavate, equalling or surpassing claw in length. Ce-
phalic plate present. Anal ring terminal, with
elongate-oval cells and 6 long setae. Ostioles present.
Circulus present. Eyes present. Body setae short and
sparse. Trilocular pores present. Tubular ducts present,
each heavily sclerotized and slightly dome-shaped at
inner end with, apparently, a septum. Bitubular and
tritubular cerores absent.

COMMENTS. The only distinguishing characters sepa-
rating this monotypic genus from Rhizoecus is the
unusual type of duct. The genus, known from Califor-
nia, is accepted by most workers on scale insects but
further related species may help to clarify its position.

Rhizoecus Kiinckel d’ Herculais

Rhizoecus Kinckel d’Herculais. 1878: 163. Type spe-
cies Rhizoecus falcifer Kiinckel d’Herculais, by
monotypy.

Ripersiella Tinsley, in Cockerell 1899:278. Type spe-
cies Ripersia rumicis Maskell, by subsequent
designation of Cockerell, 1901: 165. Synonymised
by Hambleton, 1974: 147.

Pararhizoecus Goux, 1941: 197, as a subgenus of
Rhizoecus. Type species Rhizoecus (Pararhizoecus)
petiti Goux, by original designation and monotypy.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 29

Synonymised with Ripersiella by Morrison &
Morrison, 1966: 147, and with Rhizoecus by Ben-
Dov, 1994: 286.

Morrisonella Hambleton, 1946a: 16. Type species
Morrisonella poensis Hambleton, by original des-
ignation. Homonym of Morrisonella Bartsch, 1920.
Synonymised by Ferris, 1953: 426.

Radicoccus Hambleton, 1946a: 47. Williams &
Granara de Willink, 1992: 492. Type species
Rhizoecus globosus James, by original designa-
tion. Syn. nov.

Coccidella Hambleton, 1946b: 177. Replacement name
for Morrisonella Hambleton. Synonymised by
Ferris, 1953: 426.

Neorhizoecus Hambleton, 1946a: 40. Type species
Rhizoecus coffeae Laing, by original designation.
Synonymised by Lindinger, 1957: 550.

DIAGNOSIS. Body normally small, elongate to ro-
tund, anal lobes usually not developed, their positions
each usually with 3 long anal lobe setae, or more
rarely, with numerous setae. Antennae usually placed
close together, short, strongly geniculate, each with 5
or 6 segments, these often wider than long, terminal
segment tapering, almost triangular; with sensory
falcate setae well developed on terminal and penulti-
mate segments, these in addition to normal flagellate
setae. Labium elongate, longer than wide. Anal ring
with 6 setae, these usually flagellate or occasionally
clavate; with fairly large elongate to triangular cells
that can easily be counted to distinguish between
species. Legs normally well developed; tarsus taper-
ing to a long slender claw with short setose or dilated
digitules, sometimes about as long as claw; without
translucent pores but occasionally with large vacu-
olate pores; setae on inner edges of tibia and tarsus
often elongate spine-like. Eyes present or absent. Cir-
culus present or absent, when present usually truncate
conical, distal surface often reticulate or faveolate,
usually situated on abdominal segment II within bor-
ders of segment; sometimes as many as 6 circuli
present distributed singly among other abdominal seg-
ments and occasionally present on metathorax. Ostioles
present, placed well on dorsum, sometimes reduced to
posterior pair only and occasionally barely percepti-
ble. Frons often with sclerotized cephalic plate.

Body setae usually short and flagellate, often abun-
dant, rarely very sparse. Bilocular or trilocular cerores
present, never both types present together, rarely ab-
sent altogether. Trilocular pores present, often
abundant, sometimes few. Tubular ducts present or
absent, when present usually minute with parallel or
subparallel sides. Multilocular disc pores present or
absent. Medioventral pore groups sometimes present
on abdomen. Mushroom bodies occasionally present
on dorsum and venter.

COMMENTS. The above description should be ad-

equate to separate the genus from all others discussed
here. The tritubular and bitubular cerores are unique to
the genus and some attempts have been made to distin-
guish between Rhizoecus with tritubular cerores and
Ripersiella with bitubular cerores but there are many
variations of the cerores. Sometimes the tubes of the
bitubular cerores are so appressed as to appear
unitubular and often their true structure can only be
determined by the use of oil immersion. Besides, R.
andensis (Hambleton) andR. colombiensis Hambleton,
both South American species, and R. cobelopus
Williams from Australia, are without either type but
appear to be congeneric in other characters.

Emphasis on the presence or absence of eyes, the
length and type of the claw digitules, and the shape of
the bitubular and tritubular cerores to distinguish among
genera has been challenged by Takagi & Kawai (1971)
who commented on the variability of these characters.

The position of Radicoccus needs special mention.
Hambleton (1946a) erected this genus for five species,
each with a fairly stout body, the legs and antennae,
although well developed, are minute for the size of
body, and body setae are sparse. The species show
widely diverse characters but these fall within the
present concepts of Rhizoecus. These five species and
another transferred since, are discussed as follows:

Rhizoecus globosus James. This African species,
was illustrated by De Lotto (1957) and, considering
the poor material on which the illustration was based,
the illustration is fairly comprehensive. Some further
notes are now added although a study of fresh material
is needed. The species possesses 3 circuli and large
tritubular cerores of two distinct sizes, each with short
stout ducts giving the whole ceroris a flat appearance.
Anterior and posterior ostioles are present although
they are small and almost imperceptible.

Rhizoecus incrassatus James. This African species
is similar to R. globosus, differing in possessing only 2
circuli and tritubular cerores of one size. Anterior and
posterior ostioles are present. The species was illus-
trated by De Lotto (1957).

Rhizoecus kelloggi Ehrhorn & Cockerell.This North
American species has sparse body setae, lacks bitubular
or tritubular cerores but possesses distinctive tubular
ducts, each with the orifice elliptical; nevertheless, the
sides of each duct are parallel. Ostioles are present
although they are represented by mere slits (D.R.
Miller, personal communication). There are no good
reasons to exclude this species from Rhizoecus.

Radicoccus hawaiiensis Hambleton. It was shown
by Beardsley (1966) that this species possesses all the
characters of Rhizoecus to which it was transferred.

Rhizoecus poltavae Laing. For some reason this
species, described from Ukraine, has always been
regarded as distinct in lacking bitubular or tritubular
cerores but bitubular cerores are plainly discernible in
the original material, a character noted by Matile-

Ferrero (1976) who included the species in Ripersiella.
The species is here retained in Rhizoecus.

Rhizoecus cocois Williams. Although this species,

described from India, was transferred to Radicoccus
by Tang (1992) because of the globular body and
paucity of body setae, the action was unnecessary. The
species 1s related toR. globosus and R. incrassatus and
should be retained in Rhizoecus.

All the species, therefore, that have been assigned to

Radicoccus can be comfortably included in Rhizoecus
and the name Radicoccus, as listed by Ben-Dov (1994),
is here synonymised with Rhizoecus.

Key to Genera of the Subfamily
Rhizoecinae (Adult Females)

Anal ring with protuberances; anal ring setae short, much
shorter than diameter of anal ring, numbering 2—5, un-
evenly distributed .................. Pseudorhizoecus (p. 28)

Anal ring without protuberances, anal ring setae con-
spicuous, usually at least as long as diameter of anal ring
or almost as long, numbering 6-16, distributed evenly on
ring or detached from ring and lying in an arc anterior to
ring (sometimes anal ring masked by heavily sclerotized
anal Lobes) bere vescas Sins sonasneanna cath oun tedeepetoeeeeaeveraneedis 2

Anal ring distinctly ventral in position, with a single row
of elongate cells and thick spine-like anal ring setae
around posterior edge (actually anterior edge if trans-
HETHRSLS | (HO) GORA WHOM) oo eecocosecnceere Leptorhizoecus (p. 27)

Anal ring always dorsal in position, anal ring setae not
spine-like, at least the 2 anterior pairs slender. Cells
PLESCME OL ASEM ti A reeenese ene ceee cee acento eee 3)

Anal ring with large elongate to triangular cells, anal ring
setae arising from surface of anal ring. Body rotund,
elongate-oval or weakly pyriform, never with
cephalothorax strongly dilated. Circulus, if present, never
cupped in centre. Ostioles present or absent. Tubular
GuctsipresentiomabSscm tases -eeee ee eee eee ere eee 4

Anal ring simple, without elongate cells, anal ring setae
detached from ring or at edge of ring, usually lying in an
are anterior to ring, sometimes difficult to distinguish

D.J. WILLIAMS
sosneasptez row. Poh Bo, ee, Capitisetella (p. 26)

Anal lobes well developed, protruding well beyond pos-
terior end of body, each lobe terminating in a stout
SPINE -Wike\SCtal a wesmnee eee Geococcus (p. 26)

Anal lobes barely perceptible or not developed. Any
setae on anal lobes or on normal positions of anal lobes,
flagellate sro tas 1m SKC se cece reese ee ners ae tece eee 7

Body with conspicuous dome-shaped tubular ducts, each
heavily sclerotized. Bitubular and tritubular cerores al-
Way StaDSEM ities --s eee oe ene Pygmaeococcus (p. 28)

Body without conspicuous dome-shaped tubular ducts; if
tubular ducts are present they are minute with parallel or
subparallel sides. Bitubular or tritubular cerores often
present. Body setae usually abundant, rarely sparse .....
$9. Se CE Lee. | ee i eee ee ae Rhizoecus (p. 28)

Anal lobes protruding, rounded, each with numerous
flagellate setae. Antennae each with 4 or 5 segments.
Anal opening situated at base of anal lobes. Trilocular
pores present or absent. Circuli present .................2:000
AS A ee RA ee Re Neochavesia (p. 27)

Anal lobes not protruding, posterior end of body rounded,
each lobe often with long setae, sometimes almost as
long as body. Circuli present or absent. Antennae each
with 1-4 segments. Trilocular pores always absent.....9

Antennae short, each with 24 segments; any articulation
between first and second segments poorly developed.
Claws normally long and slender except in one species
with stout claws. Minute setae abundant, always present
on dorsum and venter of abdomen, most on abdomen
sometimes replaced with short sickle-shaped setae. Cla-
vate sensory setae present or absent on body
Eumyrmococcus (p. 8)

Antennae conspicuously long, always with 4 segments,
almost as long as body, the first and second segments
with well developed articulation, the proximal end of
second segment with small points which fit into grooves
at distal end of first segment. Claws long and slender.
Minute setae abundant on dorsum of abdomen only,
absent from venter of abdomen; ventral setae on abdo-
men mostly long and stout. If sickle-shaped setae are
present they form small groups on thorax only. Clavate

from dorsal setae of abdominal segment VIII. sensory setae always absent........... Xenococcus (p. 22)
Cephalothorax dilated, abdomen narrow, tapering (in E.
neoguineensis, the cephalothorax is dilated but the abdo-
men widens before tapering). Circulus, if present, cupped
in centre, even if only shallowly. Ostioles always absent.
Tubularductsyalway S| abSentt cue .c--eesse.eeecveeseatense eee 8 RAT CTR SETS Tes
4 All body setae knobbed. Antennae each with 3 or 4 WITH DISCUSSION OF
SESMENtS LN ae ee ee ee: 5 SPECIFICITY OF THE MEALYBUG-
— All body setae flagellate. Antennae each with 5 or 6 ANT ASSOCIATIONS

SEG ITEM EG Soc. spac nec tat base pce vse teres aoe ee ote eRe 6

5 Antennae each with 4 segments. Multilocular disc pores
present. Anal ring with 8—16 short capitate setae ..........
an Sas a tn eine ein Brevicoccus (p. 26)

Eumyrmococcus corinthiacus, Greece, with Acropyga
(Rhizomyrma) paleartica, with queen in flight.
Eumyrmococcus falciculosus, Sumatra with ? ant, pri-

mary forest litter, Hevea soil.
Eumyrmococcus kolombangarae, Solomon Islands,

— Antennae each with 3 segments. Multilocular disc pores
absent. Anal ring with 6 pointed setae ..............0000000

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 31

withAcropyga (Rhizomyrma) lauta in log, queen in
flight.

Eumyrmococcus kruiensis, Sumatra, with ? ant, in
soil.

Eumyrmococcus kusiacus, Solomon Islands, with
Acropyga (Rhizomyrma) lauta in log, queen in
flight.

Eumyrmococcus lamondicus, Australia, withAcropyga
sp. in rotting wood.

Eumyrmococcus lanuginosus, Sumatra, with ‘ants’.

Eumyrmococcus maninjauensis, Sumatra, with ? ants,
in soil.

Eumyrmococcus neoguineensis, Papua New Guinea,
with Acropyga (Atopodon) ambigua, in rotting log.

Eumyrmococcus nipponensis, Japan, with Acropyga
(Atopodon) nipponensis, on rootlets of plants or in
rotten wood.

Eumyrmococcus recalvus, Australia, with Acropyga
sp.

Eumyrmococcus sarawakensis, Sarawak, with
Acropyga sp.

Eumyrmococcus scorpioides, South Africa, with
Acropyga (Malacomyrma) arnoldi, in nests and
mandibles.

Eumyrmococcus smithii, SE. Asia, with Acropyga
(Rhizomyrma) sauteri, in nests.

Eumyrmococcus sulawesicus, Sulawesi, withAcropyga
sp., rain forest.

Eumyrmococcus taylori, Australia, Acropyga sp., in
nests.

Eumyrmococcus sp., Papua New Guinea, with
Acropyga (Rhizomyrma) oceanica, in log.

Eumyrmococcus sp., Papua New Guinea, with
Acropyga sp., in rainforest.

Xenococcus acropygae, Southern Asia, Australasia,
with Acropyga (Acropyga) acutiventris, A.
(Atopodon) ambigua, A. (Rhizomyrma) lauta, in
nests and mandibles of flying queens.

Xenococcus annandalei, India, with Acropyga
(Acropyga) acutiventris, in nests and with flying
queens.

Neochavesia caldasiae, Colombia, Trinidad, with
Acropyga (Rhizomyrma) robae.

Neochavesia eversi, Colombia, Panama, withAcropyga
(Rhizomyrma) kathryna.

Neochavesia trinidadensis, Colombia, Trinidad, with
Acropyga sp.

Neochavesia weberi, Guyana, with Acropyga
(Rhizomyrma) paludis.

Capitisetella migrans, Colombia, Guyana, Surinam,
Trinidad, with Acropyga (Rhizomyrma) rutgersi.

Pseudorhizoecus proximus, Colombia, Costa Rica,
Ecuador, Guatemala, Surinam, with Acropyga
(Rhizomyrma) rutgersi, A. (Rhizomyrma) para-
mibensis.

Rhizoecus coffeae, Brazil, Costa Rica, Surinam, with
Acropyga (Rhizomyrma) paramibensis.

Rhizoecus caladii, Brazil, Colombia, Guyana, Surinam,
with Acropyga (Rhizomyrma) paramibensis.

Rhizoecus moruliferus (= R. flalcifer), Surinam, Trini-
dad, with Acropyga (Rhizomyrma) paramibensis.

Geococcus coffeae, South America, with Acropyga
(Rhizomyrma) paramibensis.

Where possible the ant species are listed in the four
subgenera of Acropyga presented by Bolton (1995a).
Other Rhizoecinae associated with Acropyga species
in South America have been recorded mentioning
mealybug genera only. It is not certain if voucher
specimens were kept and at present the species cannot
be identified.

There is no doubt that Xenococcus acropygae and
X. annandalei are always attented by Acropyga
(Acropyga) acutiventris and the mealybug may be
found eventually wherever the ant is located. The
symbiotic association suggests that neither ant nor
mealybug can live without the other. Taylor (1992)
discusses the distribution of the ant in more detail with
special reference toAustralia. Records of A. (Atopodon)
ambigua in Papua New Guinea and of A. (Rhizomyrma)
lauta in the Solomon Islands, associated with X.
acropygae, cannot be taken too seriously because they
are listed from spirit material only. The type locality of
A. acutiventris is Sri Lanka but so far no species of
Xenococcus have been reported from there despite
extensive collecting of mealybugs by E.E. Green at the
beginning of this century.

A much better picture of ant-mealybug specificity
would have emerged if some of the ants had been
identified and recorded. Present records indicate that
each species of Eumyrmococcus may be attended by a
different ant species. The type species of Eumyrmo-
coccus, E. smithii, has an apparent wider distribution
than any other in the genus and it is always attended by
A. (Rhizomyrma) sauteri. E. kolombangarae and E.
kusiacus, two closely-related species, are attended by
A. (Rhizomyrma) lauta in the Solomon Islands but the
mealybugs are known from only a few specimens. E.
neoguineensis is attended by A. (Atopodon) ambigua
in Papua New Guinea and an interesting species of
Eumyrmococcus, known from immatures only, is as-
sociated with A. (Rhizomyrma) oceanica. these
mealybugs in the Solomon Islands and Papua New
Guinea live close to colonies of A. acutiventris attend-
ing X. acropygae.

In Europe, E. corinthiacus is apparently associated
withA. (Rhizomyrma) paleartica and E. scorpioides in
SouthAfrica is attended by A. (Malacomyrma) arnoldi.

Central and South American mealybug species in
other genera are always associated with Rhizomyrma,
the only subgenus in the area, and normally each
mealybug species appears to be specific to a particular
ant species.

The wide gap in records of Eumyrmococcus be-

tween Europe and South Africa in the west and south-
ern Asia and Australasia in the east may be due to lack
of collecting. Another possibility could be that spe-
cies of Eumyrmococcus have been collected there but,
at present, are lying in ant vials waiting to be sorted or
are present in soil sample material extracted by stu-
dents of Acari or Collembola.

There must be instances when ant species overlap
and it seems difficult to believe that any species of
Acropyga would reject a strange species of
Eumyrmococcus if it were offered. Geococcus coffeae
is readily attended by A. (Rhizomyrma)
paramaribensis in Brazil although the mealybug origi-
nated in southern Asia.

These listed associations probably represent only a
fraction of similar Acropyga-mealybug associations
throughout southern Asia and Australasia. The asso-
ciations may be widespread and just await further
study.

The mealybug—ant system offers itself as an excel-
lent future candidate for detailed coevolutionary
studies based on independent cladistic analyses of the
partner groups.

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34 D.J. WILLIAMS

Fig. 1 Eumyrmococcus taylori sp. n. A. Male pupa enclosing adult male. B. Female pupa enclosing adult female.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 35

\ \
WAAL Wary.
SEQ NAN Ue

ii i AG MK fan HiAAT Ta j -
tii in
vif titli uy i th i a

pany

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iit TAA)

Fig. 2} Ewmyrmococcus corinthiacus Williams. Adult female.

36 D.J. WILLIAMS

f /t FN Tay
tre 2 OUR
WNP UPA I IAS

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es ’ tr Dy PEN TASST NAAN =
Be LOAN OSC ee a olia nat OATS Se
e FONG MUNA ALA VINE. Tay OTS YN TT IGY ~
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Fig. 3 Eumyrmococcus falciculosus sp. n. Adult female.

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LLL

UG
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Fig. 4 Eumyrmococcus kolombangarae sp. n. Adult female.

Fig. 5 Eumyrmococcus kruiensis sp. n. Adult female.

D.J. WILLIAMS

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

bis) WIZZ

i iy

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Fig. 6 Eumyrmococcus kusiacus sp. n. Adult female.

ye cietc bts tif,
GEG I LENG ohn
Leathe Wh ih j
CAAA 7
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\

Fig. 7 Eumyrmococcus lamondicus sp. n. Adult female.

S
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D.J. WILLIAMS

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

Pp i Pac AS} k len Sr
i f Goce x7 ile oF Ad My re a
t Peph ep ft Li Rat a) .
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ra aviay negeart U Dl Aare) AW aire as eee
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thitlyrs\e ant <i)
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: NI

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r ade at tite int iat; i) ip GAT ie a) i; a ih hebjeleli “IR
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ZZ

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Fig. 8 Eumyrmococcus lanuginosus sp. n. Adult female.

D.J. WILLIAMS

Fig.9 Eumyrmococcus maninjauensis sp. n. Adult female.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

4 fam
, 1

Pe UA) 1
ee ANN
ater PLT (hays

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Fig. 10 =Eumyrmococcus neoguineensis sp. n. Adult female.

44 D.J. WILLIAMS

Zi: i
heey Pt

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a
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D.J.W.

Fig. 11 9 Eumyrmococcus nipponensis Terayama. Adult female.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

(fitter y
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F |
a zg = ee eS TAN es
itt

Fig. 12} Eumyrmococcus queenslandicus sp. n. Adult female.

46 D.J. WILLIAMS

C
aS
S
SS
.
X
a= — —— ~
= y
inl mY ~
=a gL! Ui ~
i NTA +
ih ~
ee
iF /. ne ey x
SS SSS
4
= ~
to
WANG ; H
= Vid )\ \ mgt th FY j u uy
= Mf 1 Vi ‘
we ATI NN NH RA \ hat Front h AT Ml ee Mr tt
Loe vir! wit! Tite Viti p gly, Nye iy
ee USS =
— rili'
~
<
Ne
i
Nea ia
EPA VEG a
Mal hs S
\ ai N
Si atl tatiana Titi ~
Nau ~
S
X
= S
~
~
TAWA
f ANH RN
Hifi WVtPSR 7
{
i] Vr a _ Ss
C - if ~
A Till ht Ss Sr
Lily (yin ~ ~
Hy =
PAI a &
x
Gre
~
a
; 3

Fig. 13. Eumyrmococcus recalvus sp. n. Adult female.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

Fig. 14 Eumyrmococcus sarawakensis sp. n. Adult female.

D.J. WILLIAMS

Fig. 15 Eumyrmococcus sarawakensis sp. n. Adult male.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

CE, a

(
‘
\
- i ( a\j
i

‘ al
= i —o— ——_ ¢
iS : Jc L eo | =e |
Sage ao ee '. NS
ee Sse \ mee eee
me < ae |
OE SSSe=
AS \ ey ‘

Fig. 16 Eumyrmococcus scorpioides (De Lotto). Adult female.

D.J. WILLIAMS

7:
AH
AM Mil

mit
at

eS ks
. AY
NS

NN
&

\\ me
IAS AA \\

Was
a

a A ‘\

S
INT NAS
\ AK hex

\ i\

enn i i

NY AW

sua NNER TE lh ed
i uh Aan ttt) i inna
al Aiea gar
A Ty
A SN etintatiny i

/ S
CA Us

Fig. 17 Eumyrmococcus smithii Silvestri. Adult female.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

Ft)
y

bgt

ian]
if

f j yy eee aa a aa } HEN
‘ li aes
wy Pea

Fig. 18 Eumyrmococcus sulawesicus sp. n. Adult female.

all

52 D.J. WILLIAMS

it Le
Zz Vikt
E WV =
- 2S = 3 S—
wag a ee
i [Sees
2
s
y j
& LD Vy
5, ri i‘
ae yh ut
- | ’
2
zZ
Ze
S
S
=
x
<
<
~
~
“N
S
<
<
<
<
x
~
S
mS
=

Fig. 19 Eumyrmococcus taylori sp. n. Adult female.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

D.J.W.

Fig. 20 Eumyrmococcus taylori sp. n. Second instar.

1
7
4
¢

pe
a/
W
a
Va

Fig.

Eumyrmococcus taylori sp. n. A.

Female pupa.

B. Male prepupa. C. Male pupa.

D.J. WILLIAMS

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

Fig. 22 Eumyrmococcus taylori sp. n. Adult male.

Wt
carat

Leg apy

eens
A iph
fit

PAP TT

Fig. 23 Xenococcus acropygae sp. n. Adult female.

D.J. WILLIAMS

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

D.J.W.

de sp. n. First instar.

Fig. 25 Xenococcus acropygae sp. n. A. Female pupa. B. Male prepupa. C. Male pupa.

D.J. WILLIAMS

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

Fig. 26 Xenococcus acropygae sp. n. Adult male.

60 D.J. WILLIAMS

Ut
PRiy
ul anit

allt
TU

me 7
Tyg Cl
mnt

TULA
Ve A

Fits
alr

TMT
ia Hiysthy
1 aut
i K
TANS
it

a Aa i
a ARAN) GIN ea aa TTL THY
AA ALTA eH TTA
NY,

Fig. 27 Xenococcus annandalei Silvestri. Adult female.

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

Fig. 28 Leptorhizoecus deharvengi sp. n. Adult female.

D.J. WILLIAMS

‘snod0I0ULAKUN JO satdads jo dew uonNgINsIq 67 “SIA

Mojany “LI
SNIISAMDINS “7 O]
HYNUs “FT CI
saploidioos “7 p|
SISUBYDMDADS “FT ¢]
snajp2a4 “J TZ]
snoipupjsuaanb “FZ ||
sisuauoddiu “7 (|
SIsuaaUInsoau “FT 6
sisuanviuiupu “J g
snsourgnup] “J 1,
snoipuoun] ‘yg
SnovISny “y ¢
sIsuaIndy “7 p
apaDsunquojoy “FT ¢
snsojnoi20f “q 7
SNIDIYIUIAOD “7 |

NS ON

MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI

‘sno2020uaX JO saidads Jo deul uonngmnsIq = O¢ “SIA

lajppupuun “¥ 6]
av3sddo.ov “¥ 3]

D.J. WILLIAMS

INDEX

Synonyms and misidentifications are in italics; main citations in bold

acropygae 4, 23, 31
albidus 4, 5
andensis 29
annandalei 23, 25, 31
annandalei 23
atopoporus 28

Brevicoccus 26, 30

caladii 4, 31

caldasiae 4, 27, 31
Capitisetella 26, 30
Chavesia 27
clavisetosus 26
cobelopus 29
Coccidella 29

cocois 30

coffeae Geococcus 4, 31
coffeae Rhizoecus 4, 29, 31
colombiensis 29
corinthiacus 9, 10, 30

deharvengi 27

Eumyrmococcus 8, 30, 31
eversi 4, 5, 31

falciculosus 9, 10, 30
falcifer 4,5, 28, 31

Geococcus 26, 30

globosus 29
hawaiiensis 29
incrassatus 29

kelloggi 29
kolombangarae 9, 11, 30
kruiensis 9, 11, 31
kusiacus 9, 12, 31

lamondicus 9, 12, 31
lanuginosus 9, 13, 31
Leptorhizoecus 27, 30

maninjauensis 9, 14, 31
migrans 4, 26, 31
Morrisonella 29
morrisoni 28
moruliferus 4, 31

Neochavesia 27, 30
neoguineensis 9, 14, 31
Neorhizoecus 29
nipponensis 9, 15, 31

Pararhizoecus 28
petiti 28

poensis 29
poltavae 29
Prorhizoecina 26

Prorhizoecus 28
proximus 4, 28, 31
Pseudorhizoecina 26
Pseudorhizoecus 28, 30
Pygmaeococcus 28, 30

queenslandicus 9, 16

Radicoccus 29
radicum 26
recalvus 10, 17, 31
Rhizoecina 26
Rhizoecinae 2, 26
Rhizoecini 2
Rhizoecus 28, 30
Ripersiella 28
rumicis 28

sarawakensis 9, 18, 31
scorpioides 9, 19, 31
smithii 9, 19, 31
sulawesicus 9, 20, 31

taylori 10, 20, 31
trinidadensis 4, 31

weber 31

Xenococcini 2
Xenococcus 22, 30

Bull. nat. Hist. Mus. Lond. (Ent.) 67(1): 65-78 Issued 25 June 1998

Monophyly of the dacetonine tribe-
group and its component tribes
(Hymenoptera: Formicidae)

BARRY BOLTON

Department of Entomology, The Natural History Museum, Cromwell Road, London
SW7 SBD, UK (e-mail: bb@nhm.ac.uk)

CONTENTS
MRO CLIN C HOM copes sso en snaps os pacndids<csieceen da sesteeecienaepohu2 cn ects Geegee tsa ae vnc tupeeehe: San comms Rate Uns oS 65
BRAUN COIN ONAL G US SCONY secs arses snsne ao dci a axcionsanvnanend sanadesusscdanavdasatus leas tadeevts dite sateite Sevas dup adnan stacyiasvaniaens 66
Monophvilyion thle acetone (lb 6 OUOMD cece ececxs ce ¥aeenagenscucheessievey n=aetlsue neunsasues, cbekvs esaes-.2a¥ 67
Syuapomorpoies Of the GacetomiMe Ee PTOUP! cores cxceecee- -tedewneetrencanvavaceeaech ten antace- esters <oree es 68
Some characters considered but not used at tribe-group rank ...........cccccceeceseeeeeseeseeseeseeeeeees 70
IVEOTLO PLLVLys Ot OEE ASICEN CULT eee te ee enter cscs cxcnes cvtuna hex tatar Farosteb a ueeaaeleenaseess ceanupsceteasare coders 70
FSDOMONPMIGS OL ULE EAS ICCLOLIU ee steeesaes.: sac: efutacectosntsnacenss sacrsayedasaaescts saeanosvaseacechteganseaness 70
Mono PIL y Ly OLDE DD ACSLOMIMI ere: eatetetet ctee acs n<o shades teeaunn snaxtrsstensenserceesta Tons wtncpscsecennteneeesenace 71
APOMONPRIES OM LDS) ACELOMINM cert ye.stetcsss.oncntvenosnctunavectcrondeebesstav ober steasttseioesecticasestenteters? 71
Monophyly ont belPhalaGromiynnxeGini tires. ....... seve edeevie-eunasachadvied.csddcenaatde<stmteatOeses taeversse 72

Apomorphies of tribe Phalacromyrmecini
Potential synapomorphies between pairs of component tribes .............

Potential synapomorphies of Dacetonini + Phalacromyrmecini ...... er Ae
Potential synapomorphies of Basicerotini + Phalacromyrmecinl ........2:cc0cccseeeeeesereeseeseerees 73
Potential synapomorphies of Dacetonini + Basicerotint .............:cceccesscssceeecssscsecsneeseeseeseneens 73
PRNGIXHSDECIESICISSE CTEM cap x rscucveer ceva, et owsdcvescsecstee ree otece ee wetanU NT Seas easee toe Saas 73
PNORMOWIEG SEMEN Grae chievncbecs ced ek Mk dex Lewap vas denscaxncsccazudssaueediens tian uea ta payed atctruces sasaverasisescaderiua tte 73
FRE LELEMICE SV Me uh hue att Rene ho PME 2G nnnesabhas Styne mnaeen CA ena dd Made utente ana eces ties easeuatnes 74

SYNOPSIS. Within the large subfamily Myrmicinae three independent tribes of ants, Basicerotini,
Dacetonini and Phalacromyrmecini, are each recognised as monophyletic and are shown together
to constitute a monophyletic tribe-group, here termed the dacetonine-group. Synapomorphies
uniting the tribe-group and autapomorphies isolating each tribe are identified and discussed;
comments on potential synapomorphies among the three tribes are given.

done little to clarify the overall position, although the
latter does indicate a number of potentially interesting
groupings. An investigation of myrmicine compara-
tive morphology which is currently taking place is

INTRODUCTION

For many years it has been obvious that the classifica-

tion of the ant subfamily Myrmicinae at tribe rank is
inadequate, with genera or groups of genera being
associated on flimsy evidence or merely on superficial
similarity. The most recent synoptic classifications
(Holldobler & Wilson, 1990; Bolton, 1994) only serve
to stress the rather decrepit nature of the structure.
Detailed analysis of the subfamily to produce an accu-
rate higher classification at this rank is in its infancy.
Past work on myrmicine larvae (G. C. Wheeler & J.
Wheeler, 1976) and sting structure (Kugler, 1979) has

unearthing many characters of value in establishing a
natural tribe-rank classification. The aim of this paper
is to establish, through morphological characters, the
monophyly of three tribes within the Myrmicinae,
namely Basicerotini, Dacetonini and Phalacromy-
rmecini, and to show that the three together form a
monophyletic group.

Most members of all three tribes nest and forage in
leaf litter, topsoil or rotten wood where they form
small, usually monogynous, colonies. Only few for-

age openly on the surface of the ground and few are
arboreal or occur very deep in the soil. All known
species are predatory, mainly catching
entomobryomorph Collembola but with numerous spe-
cies also preying on a wide range of other small
arthropods such as sminthurid Collembola, Diplura,
Symphyla, Chilopoda, Pseudoscorpiones, Acarina,
Araneae, Isopoda, Amphipoda, and many orders of
small Insecta and their larvae (Wilson, 1953, 1956;
Carlin, 1982; Masuko, 1985; Wilson & Brown, 1985;
Dejean, 1987a, 1987b and included references). Spe-
cies and individuals are common in Berlese or Winkler
bag samples of leaf litter or rotten wood throughout the
tropics. They may be numerous or locally abundant
(e.g. Belshaw & Bolton, 1994; Fisher, in press).

Dacetonini is a large tribe with a world wide distri-
bution. At the last count it included 395 described
species (Bolton, 1995a) but many more species have
been diagnosed since and their taxonomy is currently
being studied by the author. The vast majority of
dacetonine species are tropical or subtropical but the
tribe is also well represented in zones with a
mediterranean climate. Some species occur
endemically as far north as Switzerland and Georgia in
the West Palaearctic (Kutter, 1977; Arakelian &
Dlussky, 1991), and Japan in the East Palaearctic
(Morisita, Kubota, et al., 1992). In North America one
species ranges as far north as the USA — Canada border
(Brown, 1953a). In the southern hemisphere dacetonine
species occur as far south as New Zealand (Brown,
1953b), the southernmost parts of South Africa (Bol-
ton, 1983), and central Argentina (Brown, 1962).

Basicerotini mostly shows a Gondwanic distribu-
tion, with many species occurring in the Neotropical
and Australasian regions (Brown & Kempf, 1960). In
the New World basicerotines have been recorded as far
north as Florida in the USA (Deyrup, Johnson, et ai.,
1989) and as far south as Argentina (Kempf, 1972).
The tribe is also well represented in Indonesia, Malay-
sia, the Philippines and the Pacific islands (Taylor,
1990) but is absent from the Holarctic, Afrotropical,
Malagasy and Oriental regions. Currently there are 64
described species (Bolton, 1995a), but more await
description in various collections.

Phalacromyrmecini is a widely distributed but small
and apparently relict tribe that contains only 3 de-
scribed species (Bolton, 1984), each comprising a
monotypic genus. Two of these have only been col-
lected once. They have been found in Brazil,
Madagascar and Malaysia.

The three tribes together thus have 462 described
species, representing over 10% of the subfamily
Myrmicinae, itself easily the largest subfamily in the
family Formicidae. The Myrmicinae, which contains
4377 species, represents over half the total described for
the entire family. The monophyly of Myrmicinae is
probably certain and is documented elsewhere (Baroni

B. BOLTON

Urbani, Bolton & Ward, 1992). Bolton (1994) gives a
modern definition of the subfamily and keys to its genera,
and all its species are catalogued in Bolton (1995b).

TAXONOMIC HISTORY

Dacetonini is the oldest suprageneric name in the
group, initiated by Forel (1892) to include the genera
(in the order that he gave them) Strumigenys,
Orectognathus, Epitritus, Hypopomyrmex [a fossil
taxon], Daceton, Acanthognathus, Rhopalothrix,
Ceratobasis [now a synonym of Basiceros] and
Cataulacus. By modern reckoning this is a fairly
disparate group and Forel gave no definitive diagnosis.
The next year Forel (1893) listed these same genera, as
Myrmicinae tribe Dacetonii, in a synoptic classifica-
tion. Emery (1895) also produced a synopsis in which
he rendered the tribe name as Dacetii. He did not
mention the fossil Hypopomyrmex and rightly ex-
cluded Cataulacus from the tribe. The first formal
diagnosis of Dacetonini was produced in key form by
Emery (1896) with the above inclusions and exclu-
sions. The tribe remained stable with these seven
genera for a number of years (W. M. Wheeler, 1910).

In later synopses and classifications other genera,
described in the intervening years and referred to
Dacetonini, were added, for instance Stegomyrmex,
Microdaceton, Pentastruma by the time of Emery
(1914), who spelled the tribe name as Dacetini;
Glamyromyrmex and Epopostruma by the time of
Forel (1917); Blepharidatta by the time of Emery
(1922); Codiomyrmex by the time of W. M. Wheeler
(1922), who also correctly excluded Stegomyrmex and
Blepharidatta.

Over the twenty years that followed this a number of
authors added further genera to the tribe, until by 1945
the number of genera stood at about 21. Shortly after-
wards Brown (1948) began a series of revisionary
studies on the generic composition of the tribe, adding
a number of new genera. Subsequent papers (Brown,
1949a, 1949b, 1949c, 1950a, 1950b, 1952, 1953a,
1954) refined his concept of the tribe and its compo-
nent genera. Several more new genera were described,
some were synonymised, a number of incorrectly
placed taxa were excluded, and the ranks of several
genus-group taxa were adjusted. The tribe was subdi-
vided into five subtribes (Brown, 1952) and a proposed
evolutionary sequence within the tribe was produced
(Brown & Wilson, 1959). A very old name, the fossil
Hypopomyrmex that had been one of the original
members of the tribe (and sole member of subtribe
Hypopomyrmiciti), was finally reassesed (Brown &
Carpenter, 1979) and decisively excluded.

In one paper of his revisionary series Brown (1949c)
recognised that a number of closely related genera

MONOPHYLY OF THE DACETONINE TRIBE-GROUP

formed a uniform group that could be excluded from
Dacetonini. These genera (Acanthidris, Basiceros,
Creightonidris, Heptastruma, Octostruma,
Rhopalothrix and Talaridris in the 1949 paper) were
grouped together as the tribe Basicerotini. The species
of all these were later fully revised on a world basis by
Brown & Kempf (1960).

At about the same time Kempf (1960) described an
oddly dacetonine-like genus (Phalacromyrmex) which
together with two other small genera was diagnosed as
a discrete genus group by Bolton (1984). A tribe rank
name, Phalacromyrmecini, was applied to this group
by Dlussky & Fedoseeva (1988), citing the Bolton
(1984) definition. [Earlier uses of the name
Phalacromyrmecini by G. C. Wheeler & J. Wheeler
(1976, 1985) are unavailable as they do not satisfy
Article 13 (a) of the International Code of Zoological
Nomenclature, third edition, 1985.]

Thus by the early 1990s the classification and ge-
neric content of the three tribes had reached the state
outlined below, full references for which can be
obtained from Bolton (1995b). The synopsis given
here indicates all names in the genus group that are
currently included in the three tribes. The names listed
in parentheses are current junior synonyms. Figures in
square brackets after the tribe names indicate the
number of species examined for characters visible
without dissection; a list of species dissected is ap-
pended below.

Tribe Dacetonini [569]

Subtribe Dacetoniti [5]
Genera: Acanthognathus, Daceton (=Dacetum).

Subtribe Epopostrumiti [15]
Genera: Colobostruma (=Alistruma, =Clarkistr-
uma), Epopostruma (=Hexadaceton), Mesostruma,
Microdaceton.

Subtribe Orectognathiti [14]
Genus: Orectognathus (=Arnoldidris).

Subtribe Strumigenyiti [535]
Genera: Asketogenys, Chelystruma, Cladarogenys,
Codiomyrmex, Codioxenus, Dorisidris,
Dysedrognathus, Epitritus, Glamyromyrmex
(=Borgmeierita), Gymnomyrmex, Kyidris
(=Polyhomoa), Neostruma, Pentastruma,
Quadristruma, Serrastruma, Smithistruma
(=Cephaloxys, =Miccostruma, =Platystruma,
=Weberistruma, =Wessonistruma), Strumigenys
(=Eneria, =Labidogenys, =Proscopomyrmex,
=Pyramica), Tingimyrmex, Trichoscapa.

Tribe Basicerotini [49]

Genera: Basiceros (=Ceratobasis, =Aspididris),
Creightonidris, Eurhopalothrix, Octostruma,
Protalaridris, Rhopalothrix (=Acanthidris,
=Heptastruma), Talaridris.

67
Tribe Phalacromyrmecini [3]

Genera: Ishakidris, Phalacromyrmex, Pilotrochus.

Until 1994 characters defining the tribes tended to
be of an inclusive nature. They were organised in such
a way that all taxa showing a particular combination of
characters were included in the tribe, those not show-
ing such a combination were excluded. Usually none
of these characters could be obviously pointed out as
apomorphic. Along these lines the most recent inclu-
sive definitions of the tribes were Brown (1953a) for
Dacetonini, Brown & Kempf (1960) for Basicerotini
and Bolton (1984) for Phalacromyrmecini.

Baroni Urbani & de Andrade (1994) identified the
first synapomorphy exhibited by, and thus unifying, all
three tribes: opposing, as distinct from overlapping,
mandibles. They also demonstrated that some charac-
ters regarded in the past as potential synapomorphies
(head shape, specialised hairs) had no real value.
Failing to find apomorphies for the individual tribes
within the bounds of their synapomorphy, they termi-
nated the independent existence of Basicerotini and
Phalacromyrmecini, treating them as junior synonyms
of Dacetonini. Although the Baroni Urbani & de
Andrade (1994) position was summarily reversed in
Bolton’s (1995b) catalogue, their approach provided a
useful starting point for a detailed investigation. Based
on the result of their work, the hypothesis postulated
was that only a single real tribe is represented. Charac-
ters were then sought and analysed to validate or
disprove the results of that assumption. The results
obtained indicate that the hypothesis is incorrect and
that three tribes should be recognised. The diagnosis
and discussion of the relevant characters follows this
section.

The present investigation is based upon the worker
caste, although most if not all of the characters men-
tioned are also applicable to known queens. Males are
excluded from the survey for the simple reason that
they remain utterly unknown in phalacromyrmecines
and in several genera of the other tribes. Even in the
larger genera males are only very scantily represented.
This means that selection of universal characters can-
not even be implied, let alone be guaranteed. For the
sake of this paper each currently recognised genus in
the classification outlined above is assumed to be a
valid taxon at genus rank, although work in progress
indicates that many should not have such status.

MONOPHYLY OF THE DACETONINE
TRIBE-GROUP

The tribes Basicerotini plus Dacetonini plus
Phalacromyrmecini share five worker/queen synapo-

morphies and therefore together form a monophyletic
group within the subfamily Myrmicinae. The synapo-
morphies are listed and discussed below, each under
its own heading. Character polarity here and in subse-
quent sections is determined by comparison with those
groups currently recognised as the most morphologi-
cally generalised Myrmicinae (Myrmicini, Tetramori-
ini, Formicoxenini). Characters and states across the
entire subfamily have been taken into consideration to
account for cases of convergence.

Synapomorphies of the dacetonine tribe-
group

Masticatory margins of mandibles oppose but do not
overlap at full closure
Throughout the three tribes the masticatory (inner)
margins of the mandibles fail to overlap or cross over
when the mandibles are fully closed. In taxa with
serially dentate mandibles the main dental rows inter-
lock tightly at full closure (Figs. 5, 10, 15, 23) rather
than the dental row on one mandible passing over the
other (Figs. 1, 2, 4), although a few teeth at the extreme
apices may cross over (Figs. 11, 22). In taxa with
elongate or linear mandibles the inner margins oppose
each other, or touch for part of their length at full
closure, but do not overlap (Figs. 13, 14, 24). In such
forms where a long preapical spiniform tooth occurs,
or where an apical fork of spiniform teeth is devel-
oped, the teeth themselves may interlock or cross, but
the margins from which they arise do not do so (Figs.
13, 14, 33).

This is the character first recorded by Baroni Urbani
& de Andrade (1994) and is apomorphic through the
groups under consideration. With one exception other
Myrmicinae, including all the tribes regarded as mor-
phologically most generalised on other grounds
(Myrmicini, Tetramoriini, Pheidolini, Formicoxenini),
have the mandibles plesiomorphically overlapping at
full closure (Figs. 1, 2, 4). The exception is the peculiar
monotypic genus Tatuidris (Fig. 6), the sole extant
member of tribe Agroecomyrmecini, which also has
opposing mandibles. That this is a non-homologous
parallelism with the dacetonine-group is easily dem-
onstrated.

Tatuidris has the following apomorphies; the corre-
sponding plesiomorphies exhibited throughout the
dacetonine-group are given in square brackets.

1. Tergite and sternite of first gastral segment (ab-
dominal segment 4) are fused. [First gastral tergite
and sternite unfused. ]

2. First gastral sternite reduced, very much shorter
than tergite, so that first gastral segment is directed
ventrally with respect to the postpetiole. [First
gastral sternite full-sized; first gastral segment not
directed ventrally. ]

B. BOLTON

3. Mandible with a thick brush of stout setae on inner
surface. [Mandible without setal brush. ]

4. Eyes at extreme posterior apex of scrobe. [Eyes not
at posterior apex of scrobe.]

5. Antennal sockets extremely widely separated.
[Antennal sockets relatively close together. ]

6. Alitrunk very short and compact, in profile higher
than long. [Alitrunk elongate; in profile usually
longer than high. ]

7. Apicotibial brush of thick dense setae present on
foretibia opposite strigil. [Apicotibial brush ab-
sent. ]

Conversely Tatuidris, beside lacking other
dacetonine-group apomorphies, has the following char-
acters exhibiting plesiomorphic states; their
corresponding apomorphic expressions in the
dacetonine-group (and mostly elsewhere in the
Myrmicinae) are in square brackets.

1. Postpetiole very large, very broadly articulated to
gaster. [Postpetiole small and narrowly articulated
to gaster.]

2. Sternite of postpetiole large, in profile almost as
extensive as tergite. [Sternite of postpetiole re-
duced, much smaller than tergite. ]

3. Petiole sessile. [Petiole pedunculate. ]

4. Tibial spurs pectinate, strongly developed on mid-
dle and hind legs. [Tibial spurs not pectinate,
vestigial to absent on middle and hind legs. ]

Anterior head capsule is narrowed from side to side
In the three dacetonine-group tribes the head in full-
face view, anterior to the antennal sockets, is relatively
narrow. A line drawn parallel to the long axis of the
head, touching the outermost point of the torulus, will
pass outside or very close to the point where the outer
margin of the fully closed mandible intersects the
anterior clypeal margin (Figs. 5, 7, 10, 13, 14, 15, 22,
23, 24, 33). In all other Myrmicinae (except for the few
with radically migrated toruli discussed below) a line
so constructed will pass considerably mesad of the
point where the outer margin of the fully closed man-
dible intersects the anterior clypeal margin (Fig. 1).
The anterior narrowing of the head can be expressed
as aratio, the Mandibular-Torular Index (MTI), which
with the head in full-face view may be defined as:
distance between points where outer margins of fully
closed mandibles intersect anterior clypeal margin
divided by distance between outermost points of lower
margins of toruli. Thus when MTI =1.0 the two meas-
urements are equal. MTI range of 0.5—1.3 is shown by
the dacetonine-group of tribes, and also by Tatuidris
(Fig. 6), Cataulacus (Fig. 4) and some Cephalotini.
Other tribes of Myrmicinae together have MTI range
1.5—>3.0. The reason why these three non-dacetonine
taxa have a similar MTI range to the dacetonine-group

MONOPHYLY OF THE DACETONINE TRIBE-GROUP

is that whilst their heads have remained
plesiomorphically broad anteriorly, their toruli have
secondarily migrated outwards on the cephalic dor-
sum, bringing them into line with the clypeal
intersection points of the mandibles. Thus two com-
pletely different evolutionary routes, static toruli
combined with narrowed anterior head (the dacetonine-
group), versus static broad anterior head combined
with outwardly migrated toruli (Tatuidris, Cataulacus,
some Cephalotini), will give the same overall MTI
result. For an illustration of this character spanning the
entire subfamily compare the full-face view photo-
graphs of the various myrmicine genera in Bolton
(1994).

As a direct result of the narrowing of the anterior
head the dacetonine-group shows a number of other
derived characters that are unavoidable corollaries of
this modification.

1. Width of labrum at base, and consequently also of
clypeo-labral hinge, is reduced (see below).

2. Width of buccal cavity, and consequently also of
labio-maxillary complex, is reduced (compare Fig.
3 with Figs. 12, 26, 28).

3. Anterolateral surface of head tends to be vertical,
or nearly so, and is very close to or more or less
directly below the antennal socket (e.g. Figs. 11,
182 19532):

Preocular carina is present
With the head in profile or dorsolateral view there is a
ridge, carina or lamella that originates at the posterola-
teral termination of the clypeus, extends posteriorly
below the antennal socket and usually continues for
some distance towards the level of the eye (Figs. 11,
18, 19, 30, 32). In many taxa this preocular carina is
also visible in full-face view, at least anteriorly (Figs.
13, 23, 25, 31), but usually cannot be seen in this view
in those species or genera where the frontal lobes and
frontal carinae are strongly expanded laterally (Fig. 7).
Morphologically the carina represents the exagger-
ated outer margin of the antennal fossa and is another
corollary of the narrowing of the head (particularly of
number 3, immediately above). At its weakest the
carina 1s a ridge that emphasises the dorsalmost line of
the more or less vertical side of the head capsule. At its
strongest it is a broad, laterally projecting lamella. The
structure is perhaps better termed the subtorular carina
or subantennal carina as this area is always where it is
strongest developed, but earlier literature has used
preocular carina so much that the name will probably
be retained.

Specialised range-finder/trigger hairs are developed
on the mouthparts

In the dacetonine-group hunting always seems to in-
volve an open-mandible approach to the prey, followed

by a rapid strike with the mandibles (see references
above, particularly Masuko, 1985). The strike may be
followed by static pressure of the mandibles, merely to
retain a grip on the prey until the sting can be brought
into use (‘strike-hold-sting’ technique), or dissipation
of the kinetic energy of the strike itself may be suffi-
cient to shock the prey into immobility, even if only
temporary, until the sting can be brought into play,
particularly if the prey is lifted clear of the ground
immediately after the strike (‘strike-lift-sting’ tech-
nique). Whatever the technique, the explosive closing
of the mandibles is initiated by activation of a trigger
which consists of one or more specialised hairs on the
mouthparts. Contact with prey by these hairs com-
mences a sequence of events that causes the mandibles
to snap shut (Masuko, 1985; Gronenberg, 1996). In the
vast majority of dacetonine-group taxa these hairs
arise from the anterior margin of the labrum or the
apices of the labral lobes (Figs. 16, 25-31 (broken off
short in 26), 27, 33), more rarely from the mandible
itself; such trigger hairs are not found elsewhere in the
Myrmicinae.

Specialisation of the labrum

Except for the dacetonine-group of tribes the structure
of the labrum is very uniform and generalised in the
Myrmicinae (Gotwald, 1969 and present investiga-
tion), and this same structure is prevelent throughout
the Formicidae. Outside the dacetonine-group the la-
brum is a simple sclerite (Fig. 3) that is broader than
long and broadly hinged to the clypeus. Its posterior
margin, which is attached to the clypeus, is more or
less straight. Its anterior (free) margin is indented or
cleft medially so that in general the labrum appears
broadly B-shaped, or D-shaped with a median inden-
tation in the anterior free margin. The clypeo-labral
hinge is mobile so that at rest the labrum folds back
and down with respect to the clypeus. In this position
it tightly overlaps and protects the delicate apical
portions of the labio-maxillary complex. In the
Myrmicinae this is the plesiomorphic shape and posi-
tion. It is encountered in all the morphologically more
generalised tribes (e.g. Myrmicini, Tetramoriini,
Pheidolini, Pheidologetonini, Formicoxenini,
Solenopsidini), as well as in those that are rather more
specialised morphologically (e.g. Attini, Cataulacini,
Cephalotini, Crematogastrini, Stenammini,
Agroecomyrmecini).

Tribes in the dacetonine-group show a number of
modifications away from the generalised structure
illustrated in Fig. 3, but all are based on labral narrow-
ing and elongation, and loss of the basic broadly B- or
D-shaped outline.

In taxa with mandibles that use static pressure,
regardless of whether the mandibles are long or short,
the labrum becomes longer and narrower, sometimes
longer than broad. The sclerite usually terminates in

one or two exaggerated anterior lobes or linguiform
prominences that bear the trigger hairs (Figs. 12, 16,
25, 28, 29, 31). In these forms the labrum can usually
still flex down very slightly, but it is not capable of
tight closure against the labio-maxillary complex (Figs.
12, 28). Conversely, in some taxa the labrum is hyper-
trophied and elongate-linguiform; instead of concealing
only the apex of the labio-maxillary complex it covers
the entire buccal cavity (Fig. 32). Taxa with long
kinetic mandibles tend to have the labrum very narrow
basally, and more distally to develop a pair of laterally
projecting processes or arms; the sclerite therefore
tends to be roughly T-shaped and its apex functions to
prop open the mandibles prior to striking (Figs. 26, 27,
30). One long-mandibulate genus (Acanthognathus)
has eliminated the labrum from this function and here
it is represented only by a very slender, inverted Y-
shaped sclerite.

Some characters considered but not used
at tribe-group rank

Presence of 2-segmented antennal club

An antennal club of two segments is universal in the
tribes Basicerotini, Dacetonini and Phala-
cromyrmecini, though only very poorly expressed in
Daceton. This may be the result of a single evolution-
ary event in the dacetonine-group, but a two-segmented
club is also developed in a wide range of non-
dacetonine myrmicine taxa. For example, a strongly
defined two-segmented club is universal in
Melissotarsini (Melissotarsus, Rhopalomastix), present
in all the core genera of Pheidologetonini (Afroxyidris,
Carebara, Oligomyrmex, Paedalgus, Pheidologeton),
present in some but by no means all Stenammini
(Adelomyrmex, Baracidris, Lachnomyrmex, Mayriella,
Tetheamyrma), occurs in a few Solenopsidini
(Carebarella, Solenopsis) and is present in the single
extant genus of Agroecomyrmecini (Zatuidris). It is
weakly developed in some Blepharidattini
(Blepharidatta, Wasmannia), and in some but not all
species of Cardiocondyla (Formicoxenini) and
Crematogaster (Crematogastrin1).

Antennomere count

Although very useful as acharacterin keys, antennomere
count was ignored in this survey because in some
myrmicines there is variation within genera (e.g. 10, 11
or 12 in Tetramorium (Tetramoriini); 4, 5 or 6 in
Smithistruma (Dacetonini)). Also, within the tribes of
the dacetonine-group there is a wide range of counts
that makes analysis very difficult. For example in
Dacetonini counts of 4, 5, 6 and 11 have been recorded;
in Basicerotini 7,8,9 and 12, andin Phalacromyrmecini
8, 9 and 11. Iam convinced that these counts indicate
independent morphoclinal reductions within each tribe,
with no significance at tribe rank.

B. BOLTON

Absence of tibial spurs

Throughout the tribes of the dacetonine-group spurs
are absent from the middle and hind tibiae. This condi-
tion is apomorphic among the Myrmicinae but
unfortunately is so extremely widespread through the
subfamily that it has very little analytical value in the
current survey.

MONOPHYLY OF TRIBE
BASICEROTINI

Basicerotini Brown

Basicerotini Brown, 1949c: 86. Type-genus: Basiceros
Schulz, 1906: 156. [Basicerotini relegated as junior
synonym of Dacetonini by Baroni Urbani & de
Andrade, 1994: 10; revived from synonymy by
Bolton, 1995b: 9.]

Apomorphies of tribe Basicerotini

Each worker apomorphy is briefly described, in some
cases with added comments where unrelated taxa
show similar but non-homologous modifications. The
plesiomorphic state of each character is given in square
brackets; in some cases alternative apomorphic condi-
tions are also noted for comparative purposes.

LABRUM

Distal of the hinge with the clypeus the labrum has a
deeply incised transverse groove or trench across its
entire dorsal width. The distal margin of this groove is
defined by a sharp edge, ridge or crest (Fig. 16; also
visible between the closed mandibles in Figs. 13, 14).
{Labrum without sharply defined transverse groove
distal of hinge. ]

In the basicerotines this groove, or at least its sharp
distal margin, is usually located far enough anteriorly
on the labrum as to be visible in ordinarily mounted
specimens which have the mandibles ajar and the
labrum slightly depressed.

TORULUS

The dorsal lobe of the torulus is hypertrophied and
strongly curved downwards (Figs. 18, 19); its outer
surface is nearly vertical so that it conceals part to most
of the condylar bulb of the scape and the antennal
socket itself. [Torulus a simple annulus or with a small
lobe present dorsally that is horizontal and does not
conceal the condylar bulb of the scape and the antennal
socket. |

SCAPE NECK ARTICULATION

Because of the size and shape of the torulus the space
in which the basal neck of the scape (the short narrow
section between condylar bulb and scape shaft proper)

MONOPHYLY OF THE DACETONINE TRIBE-GROUP

can move is narrow and directed fore and aft (Figs. 18,
19). [Space in which basal neck of scape can move is
not restricted to a narrow fore and aft motion. ]

BASE OF SCAPE

Scape shaft near base is bent downwards through a
right-angle or near right-angle; the articulatory condyle
at the extreme base projects forward from this through
another right-angle that is not in the same plane as the
first bend but rather is rotated through about 90 degrees
(Figs. 8, 9). [Scape shaft in a straight line with basal
condyle; or if scape angled downward near base then
angle of basal condyle remains in the same plane as the
shaft and is not rotated through 90 degrees. ]

ANTENNAL FOSSA

Antennal fossa separated from scrobe by at least a
cuticular rim or crest; the depressed fossal area sur-
rounded on all sides by raised or prominent cuticle
(Figs. 18, 19). [Antennal fossa and scrobe confluent
(when the latter is developed). ]

SCROBE

Scrobe always present and located below the eye
(Figs. 18, 19), the latter usually situated on the extreme
dorsolateral rim of the scrobe, more rarely towards
underside of upper scrobe rim (very rarely eye absent).
[Scrobe absent, or present but extending above the eye
(eye sometimes absent). ]

Some other myrmicine taxa have a scrobe that
extends below the eye, namely Cataulacus, some
Cephalotini, and the Dacetonini related toEpopostruma
(Epopostrumiti in the synoptic classification above).
Cataulacus and the few cephalotines with this condi-
tion can be dismissed as obvious convergence. Not
only is the basic structure of the scrobe dissimilar but
also the detailed structure of the head is very different.
They lack, of course, the apomorphies of the
dacetonine-group of tribes and exhibit their own series
of apomorphies. The Epopostrumiti, on the other hand,
belong in the dacetonine-group. Apart from possess-
ing the apomorphies of Dacetonini and lacking those
of Basicerotini, the scrobes in Epopostrumiti lack
sharply defined posterior margins, usually lack vent-
rolateral margins and, except in a very few species, fail
to extend forward to the mandibular articulation (Fig.
32); all of these are developed in Basicerotini. Finally,
a morphocline of species in the genera Colobostruma
— Mesostruma — Epopostruma is present that exhibits
a gradual increase in development and definition of the
scrobe; no basicerotine could be inserted into the
series, nor added to either end.

OCCIPITAL FORAMEN

The occipital foramen is set in a deep depression on the
occipital surface of the head; cuticular margination is
continuous around the depressed area (Fig. 17). [Oc-
cipital foramen not set in a continuously marginate
deep depression. ]

A very few Dacetonini have a transverse ventral rim
of cuticle below the occipital foramen. This structure
is very different from the basicerotine organisation.

HELCIUM

Helcium arises from the base of a broad, deeply con-
cave depression or excavation in the anterior surface of
the postpetiole (Fig. 20). [Helcium not set in a concave
depression. |

GASTER

First gastral tergite and sternite each distinctly trans-
versely marginate basally, immediately behind the
postpetiole (Fig. 21). [First gastral tergite and sternite
not marginate basally. ]

Members of the strumigenyite group of Dacetonini
genera have a specialised transverse crest on the first
gastral tergite, the limbus, which is an apomorphy of
that group. The limbus is located prebasally and is
inclined towards the base proper, where it overhangs
the presclerites of the segment. Itis nota homologue of
the basal margination developed in Basicerotini. No
Dacetoninihavea basally marginate first gastral sternite.

SCULPTURE
First gastral tergite and sternite with characteristic
sculpture of dense, sharply incised, separated punc-
tures (e.g. Fig. 21). [First gastral tergite and sternite
unsculptured, or with different sculpture. |

This character may seem rather imprecise, but the
form of sculpture is striking and not obviously re-
peated elsewhere. A very few individual basicerotine
species have secondarily reduced or effaced the sculp-
ture. Elsewhere in the dacetonine-group, and in the
Myrmicinae as a whole, are many species with sculp-
tured gasters. Even when the sculpture in these is of a
basically punctate form it tends to be reticulate-punc-
tate or sparse, or on one sclerite but not the other. It is
not the dense deeply-incised punctation exhibited by
the Basicerotini on both sclerites.

MONOPHYLY OF TRIBE
DACETONINI

Dacetonini Forel

Dacetonini Forel, 1892: 344. Type-genus: Daceton
Perty, 1833: 136.

Apomorphies of tribe Dacetonini

The two worker apomorphies are briefly described and
comments added. The plesiomorphic state of each
character is given in square brackets.

MANDIBLE
Mandible with a medially projecting cuticular process
present on the inner margin close to the base

(basimandibular process); the process not merely a
modified tooth (Figs. 25, 26, 27, 31, 33). [Mandible
without a basimandibular process. ]

In some dacetonine taxa the basimandibular process
is visible in ventral view in ordinarily mounted speci-
mens, but in most the mandibles need to be opened
quite widely or the labrum depressed. The process is
variably shaped in different dacetonine taxa, taking
the form of a lobe, a short or long spur, or a lamella
(basal lamella). In all dacetonines except
Acanthognathus the basimandibular process inserts
between the clypeus (above) and the labrum (below)
when the mandibles are closed and is part of the jaw
locking mechanism. In Acanthognathus the process is
hypertrophied but passes ventral to the labrum as the
latter is vestial in this genus and no longer serves in the
jaw locking mechanism.

A few basicerotine species in the genera
Eurhopalothrix and Octostruma have the basal tooth
of the mandible flattened or lengthened. This is not
homologous with the dacetonine basimandibular proc-
ess, which is derived from the mandible itself and not
from a tooth. Basicerotine species with a modified
basal tooth are exceptional and certainly best regarded
as independent acquisitions. The usual condition is to
have the basal tooth quite normal; presence/absence of
a flattened basal tooth cannot be used diagnostically
either at genus or species-group rank.

LABRUM
Dorsal surface of labrum with an impression or pair of
impressions located medially on the labral shield,
distal of the basal hinge but proximal of the labral
lobes (Figs. 27, 30). [Labrum without mid-dorsal im-
pression. |

The labral impression receives the basimandibular
processes of the mandibles when they are fully closed.
The character is not repeated anywhere else in the
Myrmicinae. It is secondarily lost in Acanthognathus
because, as pointed out above, the labrum is secondar-
ily extremely reduced in this genus.

MONOPHYLY OF TRIBE
PHALACROMYRMECINI

Phalacromyrmecini Dlussky & Fedoseeva

Phalacromyrmecini Dlussky & Fedoseeva, 1988: 80
[based on diagnosis in Bolton, 1984: 381]. Type-
genus: Phalacromyrmex Kempf, 1960: 89.
[Phalacromyrmecini relegated as junior synonym
of Dacetonini by Baroni Urbani & de Andrade,
1994: 10; revived from synonymy by Bolton, 1995b:
9.)

B. BOLTON

Apomorphies of tribe
Phalacromyrmecini

Each worker apomorphy is briefly described and com-
ments added. The plesiomorphic state of each character
is given in square brackets.

MANDIBLE

Dentition of alternating large and small teeth from
base to apex; largest tooth usually the basal. [Dentition
not of alternating large and small teeth; largest tooth
usually the apical. ]

A few isolated species or small species-groups in
the dacetonine genus Glamyromyrmex have the basal
tooth the largest on the masticatory margin, but these
lack alternating dentition.

MESOPLEURON

Katepisternum with an impression or groove extend-
ing obliquely downward from posterior margin of
mesopleural hair-filled gland towards metapleuron;
impressed area usually bounded by ridges or carinae.
[Katepisternal oblique groove absent. ]

SCAPE
Scape slender basally, clavate apically; entire scape
roughly Indian-club shaped. [Scape subcylindrical.]

POTENTIAL SYNAPOMORPHIES
BETWEEN PAIRS OF COMPONENT
TRIBES

The object of this paper has been to establish the
monophyly of the tribes and the tribe-group, rather
than to produce a formal phylogeny. However, anumber
synapomorphies potentially linking pairs of tribes
within the group have been noticed and these are
mentioned below. The plesiomorphic state expressed
by the isolated tribe in each set is given in square
brackets.

Potential synapomorphies of Dacetonini
+ Phalacromyrmecini

POSTPETIOLE-GASTER ARTICULATION

Diameter of presclerites of abdominal segment 4 (=first
gastral segment) small and constricted so that the
postpetiole-gaster articulation is relatively narrow.
[Basicerotini: diameter of these presclerites broad so
that the postpetiole-gaster articulation is relatively
wide. ]

PRESCLERITE OF FOURTH ABDOMINAL SEGMENT
Pretergite of abdominal segment 4 (=first gastral seg-
ment) subtended by a short narrow neck-like
constriction. [Basicerotini: pretergite subsessile to ses-
sile.]

MONOPHYLY OF THE DACETONINE TRIBE-GROUP

BASIMANDIBULAR GLAND
Basimandibular gland present. [Basicerotini:
basimandibular gland absent. ]

This gland is variously developed in many genera of
these two tribes (/shakidris, Microdaceton,
Strumigenys, Glamyromyrmex); it is universally ab-
sent from basicerotines. Its absence from some
dacetonines (Orectognathus, Acanthognathus) may be
secondary.

MESOPLEURAL GLAND
Anterolateral angle of mesopleuron bears a hair-filled
glandular structure set in an emargination of the rim of
the sclerite. [Basicerotini: mesopleural gland absent. ]
This supposed gland varies from absent to mas-
sively hypertrophied in Dacetonini and
Phalacromyrmecini. It is universally absent in
Basicerotini.

Potential synapomorphies of
Basicerotini + Phalacromyrmecini

BASE OF MANDIBLE

Base of mandible with a long stiff ventrally directed
seta on ventral margin. [Dacetonini: such a seta ab-
sent. ]

This specialised seta is present in all Basicerotini. In
Phalacromyrmecini it is obvious in /shakidris. Not
recorded in the single specimen of Phalacromyrmex
currently available, but this is in poor condition and
badly mounted.

PROPODEAL SPIRACLE
Propodeal spiracle low on side of sclerite, abutting the
margin of the small metapleural gland bulla.
{Dacetonini: propodeal spiracle high on side, widely
separated from metapleural gland. |

In the dacetonine genus Acanthognathus the spiracle
abuts the metapleural gland bulla but here the spiracle
is high on the side and the bulla is secondarily ex-
tended upwards.

Potential synapomorphies of Dacetonini
+ Basicerotini

None detected.

APPENDIX: Species dissected

The parts of all dissected specimens, whether partially
or completely disarticulated, have been remounted on
green-flagged card points and are deposited in The
Natural History Museum, London. [List does not in-
clude the many taxa mounted with mandibles open
and mouthparts displayed, but not otherwise dissected;
these have been mounted on blue-flagged card points
in the Natural History Museum, London, collection. ]

Basicerotini

Basiceros. discigera, manni, militaris, singularis.

Eurhopalothrix: australis, biroi, bolaui, dubia,
floridana, gravis, heliscata, insidiatrix, jennya,
omnivaga, procera, punctata, speciosa, spectabilis,
szentivanyl.

Octostruma: balzani, betschi, iheringi, inca, rugifera,
stenognatha.

Protalaridris: armata.

Rhopalothrix: ciliata, isthmica, plus | unidentified
species.

Dacetonini

Acanthognathus: brevicornis, ocellatus, rudis.

Codiomyrmex: thaxteri.

Colobostruma: alinodis, cerornata, plus | unidenti-
fied species.

Daceton: armigerum.

Epitritus: argiolus, hexamerus, laticeps, roomi.

Epopostruma: frosti.

Glamyromyrmex: beebei, excisa, flagellatus,
semicomptus, sistrurus, tukultus, plus 2 unidenti-
fied species.

Gymnomyrmex: villiersi.

Kyidris: mutica, plus | unidentified species.

Mesostruma: browni, turneri.

Microdaceton: exornatum, tibialis, plus | unidentified
species.

Neostruma: brevicornis, crassicornis, myllorhapha,
zeteki.

Orectognathus: antennatus, clarki, mjobergi,
szentivanyi, versicolor.

Pentastruma: sauteri.

Quadristruma: emmae.

Serrastruma: lujae, ludovici, serrula.

Smithistruma: alberti, angulata, dohertyi,
fridericimuelleri, microthrix, ornata, transversa,
truncatidens, plus 6 unidentified species.

Strumigenys: biolleyi, chyzeri, denticulata, doriae,
elongata, gundlachi, godmani, koningsbergeri,
louisianae, micretes, nidifex, pallestes, perplexa,
precava, prospiciens, rogeri, rukha, saliens,
signeae, subedentata, tigris, trieces, trudifera, plus
12 unidentified species.

Trichoscapa: membranifera.

Phalacromyrmecini

Phalacromyrmex: fugax.
Ishakidris: ascitaspis.

ACKNOWLEDGEMENT

My sincere thanks to Suzanne Lewis (Department of
Entomology, The Natural History Museum) for taking
the SEM photographs used to illustrate this paper.

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Brown, W. L. Jr. 1954. A preliminary report on dacetine ant studies
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Brown, W. L. Jr. 1962. The Neotropical species of the ant genus
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238-267.

Brown, W. L. Jr. & Carpenter, F. M. 1979. A restudy of two ants
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Carlin, N. F. 1982. Polymorphism and division of labor in the dacetine
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Dejean, A. 1987a. Behavioral plasticity of hunting workers of
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Dejean, A. 1987b. Etude du comportement de prédation dans le genre
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91-103.

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Dlussky, G. M. & Fedoseeva, E. B. 1988. Proiskhozhdenie i rannie
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Kugler, C. 1979. Evolution of the sting apparatus in the myrmicine
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Masuko, K. 1985. Studies on the predatory biology of Oriental
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Morisita, M., Kubota, M., Onoyama, K., Ogata, K., Terayama,
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H.T. 1992. A guide for the identification of Japanese ants. 3.
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224. Monachii.

Taylor, R. W. 1990. New Asian ants of the tribe Basicerotini, with an
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Memoirs of the Entomological Society of Washington 7: 1-108.
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Formicidae. Transactions of the American Entomological Society

111; 255-264.

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663 pp. New York, Columbia University Press.

Wheeler, W. M. 1922. The ants of the Belgian Congo. Bulletin of the
American Museum of Natural History 45: 1-1139.

Wilson, E. O. 1953. The ecology of some North American dacetine

MONOPHYLY OF THE DACETONINE TRIBE-GROUP 75

ants. Annals of the Entomological Society of America 46: 479-495. Wilson, E. O. & Brown, W. L. Jr. 1985. Behavior of the cryptobiotic
Wilson, E. O. 1956. Feeding behavior in the ant Rhopalothrix biroi predaceous ant Eurhopalothrix heliscata, n. sp. Insectes Sociaux
Szabo. Psyche 63: 21-23. 31: 408-428.

76 B. BOLTON

Figs. 1-11. Worker ants: 1-3, Head of Myrmica rubra; 1, tilted slightly back from full-face view; 2, oblique close-up of
fully closed mandibles; 3, ventral view to show buccal cavity; 4, head of Cataulacus lujae, tilted back from full-face view;
5-7, heads in full-face view of 5, Pilotrochus besmerus; 6, Tatuidris tatusia; 7, Ishakidris ascitaspis; 8-9, scape of
Basiceros singularis, condyle uppermost; 8, ventral view; 9, dorsal view; 10-11, head of Eurhopalothrix platisquama; 10,
full-face view; 11, oblique frontal view.

MONOPHYLY OF THE DACETONINE TRIBE-GROUP 77

Figs. 12-22. Worker ants: 12, ventral head to show buccal cavity of Eurhopalothrix platisquama; 13-15, heads in full-face
view of 13, Rhopalothrix ciliata, 14, Protalaridris armata; 15, Basiceros discigera; 16, oblique frontal view of head of
Basiceros singularis, mouthparts extended and right mandible removed; 17, occipital foramen of Eurhopalothrix heliscata;

18-19, lateral view of head with antenna removed, of 18, Basiceros singularis; 19, Eurhopalothrix platisquama; 20,

frontal view of helcium of Eurhopalothrix procera; 21, base of gaster in oblique ventral view of Basiceros singularis; 22,

| head in full-face view of Dysedrognathus sp. n.

B. BOLTON

Figs. 23-33. Worker ants. 23-25, head in full-face view of 23, Smithistruma sp. n.; 24, Daceton armigerum; 25,
Smithistruma reliquia, mandibles open; 26-28, ventral view of head to show buccal cavity of 26, Microdaceton sp. n.,
mandibles open; 27, Daceton armigerum, right mandible and labio-maxillary complex removed; 28, Smithistruma
truncatidens;, 29-30, oblique frontal view of head to show labrum, mandibles removed, of 29, Smithistruma truncatidens;
30, Strumigenys sp. n.; 31, head in full-face view of Smithistruma kerasma, mandibles open; 32, head in ventrolateral view
of Colobostruma sp.; 33, mandibles in oblique frontal view of Microdaceton tibialis.

Bull. nat. Hist. Mus. Lond. (Ent.) 67(1):79-152 Issued 25 June 1998

An annotated checklist of bumble
bees with an analysis of patterns of
description (Hymenoptera: Apidae,
Bombini)

PAUL H. WILLIAMS

Department of Entomology, The Natural History Museum, Cromwell Road, South
Kensington, London SW7 5BD

CONTENTS

| Hii OTG IC COLAC 01 SA SPR ee ORE nc RRRPEY cece ce ert pe SScR PPA Rae ORME 08 dc. ORE Sm ok Ore onan eee 80
PAS EALISE SEO IES IECLES socass vaaeteraccee dr caamevusa scaversussaeroncetaxcnes cade caps sauseizeccsee ured Tatars meen gone otsers = ane 80
FUISLOUY ON CIS COVEUY OL SPECIOS fences onseteteseascsncoseesasgatusch cust daceacoasy shes eatovedadec¥oates diospsoeesucece teas 80
ANSTO yal Chin UNIT GALCL OM OL TALES) « cecps deen ce <oeesk fab nes ch nectar asae cuca org <tsaccrab sad esexansuecheainssdcnancpesséee 81
Summary of historical and regional trends in describing bumble bees ...... 86
WEVElonmmemt Ola eVvISed CHECKING Opietececcrecct acecassevaraceseesasvedscceatitaactaroarccckacesseutnestacancecnencee 87
PMCMSIICHMLE CE CHIC IBS) nee tencerscerserrtcsestreteseraressteeesca ta esersgsefuxiurcrasteveseiasvadisesczsssesaosiaueeseaseescanes 88

SEONOMONY Meeteee tener cere ctu c cee cc M Scere ts settee acci ce sancancednevaessesved eaddzncs sassppocuyesneesagiai«spcd/seapenwascen 88
Phylogeny, supra-specific taxa and ordering Of SPeCieS ...........:eescceseesseeseeeeeeeeeeeeeneeeeeeeeeneees 89
Te MaNiGNALSOMMUMINALE SPECLOS ter Meech ics ccasconcectcascsescaccusscacussssussveccoassaseestvacensnetsatanesasnons 90
Sub-specific taxa

INomenGlatune?.2. ic .ccteit.cccees.s
ATSIC) cred py kiteelll COMVEMIMOMS 4. meseee sa teeters csessxencsvevacespescaaecteatssdcscsesdeseccecddscesussvenrsdheustscceneers
SEI SG HIT OV, 7900) PBB Ree eee eR oe se a ee A ee Rh nent et anh 97
MPU GATOS GLE ZN x oe, ete eae ean. on ccuvsschteederce acecceeracedees hl usctee MMe ctebenssecstlaedeaascs 97

IC YiS (ar oH iro) ol sTa FO) SA Reece a5 nt RR er AR ie ee Eo Fr ee ee 97

IBIS (ROIBSTIG CLES en eee eee, Soe Pe tearebeteeibe St... i LORE R aren Lt eden ee EI age oe EE 99

GRCKENGCES crrescess ve santh restr sd vansh cuban Mee ee Tyas oon saa acsht estate dade edassdeausseeshsasdddubendeealdsnseaeeteeeanea 140

SYNOPSIS. Bumble bees are among the minority of groups of organisms for which there is some
evidence that most species have already been described. Nonetheless, a synoptic revision of the
group has been delayed, in part by the difficulties imposed by an unusually high ratio of names to
species (averaging more than | 1). To explore some of the factors contributing to this phenomenon,
historical and geographical trends in the naming of bumble bees are summarised. This shows that
most taxa were named by European authors, beginning with the most widespread European
species, moving later to not only the more narrowly distributed species and to species from other
parts of the world, but also to taxa at progressively lower nomenclatural ranks, particularly within
the more widespread European species. Nearly half of all of these names have been published
since the last world-wide checklist in 1922. In attempting to bring this up to date, the present
checklist adopts broad interpretations of species and recognises a total of 239 recent species
(including the social parasites but excluding fossil taxa), with 24 new synonyms and 29 provi-
sional synonyms. The list also includes notes on alternative interpretations of taxonomic status and
on nomenclatural problems, drawing attention to those cases where further research is most
urgently needed. In particular, suggestions are presented for an application to the International
Commision on Zoological Nomenclature to use its Plenary Power in order to conserve current
usage of the commonly used names atratus, balteatus, distinguendus, flavifrons, humilis,
hyperboreus, mesomelas, mixtus, norvegicus, polaris, pyrenaeus, soroeensis and variabilis.

INTRODUCTION

Bumble bees have long been popular with collectors.
Just as with butterflies, part of the attraction may be
explained by their bright colours, large body size,
activity during daylight hours, and abundance in the
north-temperate regions where most collectors have
lived. As a result, large samples of bumble bee speci-
mens have now been assembled, even from remote
parts of the world.

A problem for biologists trying to identify bumble
bee species, all the more apparent because of the large
amount of material available, is that while bumble
bees can be described as morphologically relatively
‘monotonous’ (Michener, 1990), they are often ex-
traordinarily variable in the colour patterns of their
pubescence. The situation is made worse by a strong
tendency for species to converge locally on different
colour patterns (Plowright & Owen, 1980).

Faced with this variation, generations of taxono-
mists since the starting point of Linnaean nomenclature
in 1758 have described differing individuals under a
plethora of more than 2800 formal names (Williams,
unpublished catalogue, including names for species,
subspecies and synonyms, as well as infrasubspecific
names, misspellings and other unavailable names).
Most of these names are for taxa below the rank of
species, and just 239 taxa are interpreted here as
separate species. Arguably, the nomenclatural burden
of more than 11 names per species (median 5, maxi-
mum 186) has slowed progress towards a complete
revision of the group. Hence there is a need for an
overview which, although bound to require revision,
will provide an improved framework for more detailed
regional studies. It is also important to understand any
regional or taxon-directed bias in patterns of taxo-
nomic description when seeking to interpret patterns
in diversity, ecology and biogeography. The present
checklist begins to address these needs.

Past lists of species

There have been few attempts to present complete
revisions, catalogues or checklists of all bumble bee
species from which to see summaries of past views.
Latreille (1809) included 13 species in his genus
Bombus. Most of the early lists included just those
species seen by their authors, usually from particular
collections, and often from just one region. For exam-
ple, Smith (1854) catalogued 87 bumble bee species
(79 Bombus + 8 Apathus [= Psithyrus]) in the collec-
tion of the British Museum. The only truly synoptic
catalogue of bumble bees was published by Dalla
Torre (1896), with 255 (non-fossil) species (228
Bombus + 27 Psithyrus). It included many varietal
names, synonyms and early references. The reason

P.H. WILLIAMS

why Dalla Torre’s species count exceeds the total now
recognised as described before 1899 (159 species, Fig.
1) is of course that many of his species are now treated
as synonyms or subspecies. Later, Skorikov (1922a)
listed 237 species (plus 70 ‘Bombi incertae sedis’ ), but
with few synonyms and without including Psithyrus.
Nonetheless, Skorikov’s list did arrange most of the
known species within his genera and subgenera, which
form the basis of the current subgeneric system
(Richards, 1968).

Taken together, the few past lists of bumble bees
show that the number of taxa accepted as species at a
particular date grew rapidly during the nineteenth
century, but has since remained relatively stable, with
a slight decline to the conservative estimate of 239
species in the present list. Undoubtedly part of the
explanation for this decline lies in the relatively con-
servative species concept accepted at present (see
below). This reflects a gradual shift in emphasis among
criteria for recognising species from the use of colour
characters to the use of morphological characters,
particularly to using characters of the male genitalia
(see the introductory comments by Radoszkowski,
1884). A similar pattern of growth and decline has
been found for past numbers of milkweed butterfly
species (Ackery & Vane-Wright, 1984). However, there
might now be another period of rapid growth if mo-
lecular characters and phylogenetic species concepts
(discussed below) were to be applied (cf. discussion of
the number of bird species by Martin, 1996; Patterson,
1996; Zink, 1996, 1997; Snow, 1997).

History of discovery of species

The dates of first formal description for the currently
recognised bumble bee species show that the highest
rates of species discovery were in the latter half of the
nineteenth century through to the First World War
(Fig. 1, median date 1877). These species are recog-
nised retrospectively from the present list, rather than
as the numbers accepted within each time period. The
larger dips in overall rate of discovery may be associ-
ated with factors such as war and its aftermath (e.g.
Napoleonic and Second World Wars), presumably
through constraints on resources and on freedom of
travel.

Some authors described many more bumble bee
species than others: 45% of presently accepted species
were described by just 10% of the authors who de-
scribed these species (Smith 32 species, Skorikov 19,
Cresson 17, Morawitz 15, Radoszkowski 13, and Friese
12). Similarly, Gaston, Scoble & Crook (1995) found a
skewed pattern of activity among authors describing
geometrid moths. But of the six authors who described
the most bumble bee species, only Ezra Cresson (Snr)
actually worked in the New World, whereas the other
five were based in Europe (including European Russia).

CHECKLIST OF BUMBLE BEES
250

200

150

Species

100

1750 1770 1790 1810 1830 1850

1870" 1890" 1810) ~ 1930" 1950" 1970 = 1990

Date of description

Fig. 1 Rate (lower grey) and cumulative number (upper white) of first formal descriptions of presently recognised bumble
bee species (dates from the oldest available names in the sense of ICZN, 1985).

Rates of discovery of bumble bee species vary to
some extent among biogeographic regions (Fig. 2).
Again, this was also found for geometrid moths by
Gaston, Scoble & Crook (1995). For bumble bees, the
recent proportional discovery rates have been lowest
in the New World and highest in the Oriental Region.
The Neotropical Region appears to have a small known
bumble bee fauna for its large area. There is no obvious
evidence that descriptive effort has been lacking, al-
though detailed revisionary work is needed and species
with small range sizes may remain to be discovered. In
contrast, the Oriental Region’s high recent propor-
tional rate of species discovery, despite its smaller area
than the other regions, is possibly explained in part
because it has been studied intensively for a shorter
period.

The world-wide rate of discovery of genuinely un-
known bumble bee species appears now to be slowing
down (Fig. 1). Undiscovered species are very likely to
remain, although there is no evidence that large num-
bers of species are awaiting description in collections
(although some known subspecific taxa might yet be
recognised as species if changes were to occur in
species concepts or in the availability of character
evidence, see Martin, 1996; Patterson, 1996; Zink,
1996). The sigmoidal pattern of species discovery in
Fig. 1 isalsoshownby afew other relatively well known
groups such as birds, although for most large groups
(including Hymenoptera as a whole) the rates of de-
scription continue to be high or are even increasing
(Hammond in Groombridge, 1992; Tennessen, 1997).

History of publication of names

The present checklist is intended only to address the
question of taxa at the rank of species (see below). For
this purpose it is not necessary to consider concepts of
taxa at the rank of subspecies and below and subspecific
names may be treated in analyses as further synonyms
of species (Gaston & Mound, 1993). This is not to say
that subspecific taxa ought not to be recognised if they
were considered useful in the context of other studies.
In addition, some authors have applied classical names
to taxa at even lower nomenclatural ranks, for example
in referring to ‘varieties’ or ‘forms’ within subspecies.
These are now interpreted as infrasubspecific names
and are ‘unavailable’ for use in the sense of the /nter-
national Code of Zoological Nomenclature (ICZN,
1985). They have had to be included in a manuscript
catalogue (unpublished) in order to avoid confusion by
explicitly resolving questions of nomenclatural status
and availability. Infrasubspecific names are included
in this analysis as a category separate from specific or
subspecific names because of their particular signifi-
cance for understanding historical patterns in the
description of diversity at the lowest nomenclatural
ranks.

Bumble bees have the highest known levels of
synonymy (83%, or 92% if infrasubspecific names
were to be included) in comparison with the range of
insect taxa reviewed by Gaston & Mound (1993).
Their results showed synonymy levels ranging from
7% for Siphonaptera to 80% for Papilionidae and

Palaearctic Region
120

0)
1750) 1790! 18380) 1870) 19110) 1950) 1'9S0

Oriental Region
60
40

0
1750) 177/90) 1880) 1870) 1910) 19505) 1390

P.H. WILLIAMS
Nearctic Region

48
32

(0)
1750) 11790) «1830 1870" 1910) SaiS50)) S80

Neotropical Region

0
1750; > 1790), 1830) | 1870) 190 ISSO RT SS0

Fig. 2 Cumulative number of first formal descriptions of presently recognised bumble bee species with centres of area of
occupancy (so species lists do not overlap) in each of the four principal biogeographic regions occupied by bumble bees
(dates from the oldest available names in the sense of ICZN, 1985; regions defined in Williams, 1996b: fig. 1; Oriental
includes northern and southern Oriental Regions; Nearctic includes northern, central and southern Nearctic Regions;
Neotropical includes northern, central and southern Neotropical Regions; the Arctic Region is excluded; species that are
exclusively peri-Tibetan Oriental but which nevertheless have range centres in Palaearctic central Tibet by simple range

averaging are included as Oriental species).

Pieridae combined. It must be born in mind that the
insect taxa that they surveyed are all more speciose
than the bumble bees by a factor of at least four, and
extreme values for larger groups are less likely. None-
theless, Gaston & Mound (1993) also noted that the
two families of most brightly coloured butterflies have
the highest levels of synonymy and that these families
have many more subspecific names than the smaller
and duller-coloured hesperiid butterflies. R. I. Vane-
Wright (pers. com.) suggests that synonymy rates may
be particularly high among the large, colourful butter-
flies of the Danainae and Parnassius.

In contrast to the discovery of currently recognised
species, the greatest activity in publishing names for
all supposed bumble bee taxa at the rank of species and
below was concentrated slightly later than for pres-
ently recognised species, in the first half of the twentieth
century (Fig. 3, median date 1922). This difference
may be explained in part by the logical inevitability
that synonyms and names for taxa below the rank of
species can only be published subsequently to valid

species names (i.e. the oldest available names, exclud-
ing junior homonyms, in the sense of ICZN, 1985). If
these names were in effect to represent the redescription
of known species at random, then the earlier described
species might be expected to have accumulated more
names. Studies of other taxa have also shown that both
the date of first description and the number of syno-
nyms per species may be affected by variation in the
size of a species’ geographic range (as well as by other
factors such as body size). Large range size is likely to
affect the date of first description because it contrib-
utes to a greater ‘apparency’ of the species to collectors
(Gaston, Blackburn & Loder, 1995), particularly as
broad correlations between range size and abundance
suggest that widespread species also tend to have
higher local densities (Brown, 1984; Gaston, 1994; for
bumble bees, see Hanski, 1982; Williams, 1988).Apart
from enhancing the chances of random redescription,
large range size is also likely to affect the number of
synonyms because there is a greater likelihood that
specimens collected in one area will be regarded as

CHECKLIST OF BUMBLE BEES
3000

2500

2000

Names
3
+

1000 +

1750

1770 1790 1810 1830 1850

1870 1890 1910 1930 1950 1970 1990

Date of description

Fig. 3 Rate (lower grey) and cumulative number (upper white) of all descriptions with classical names for bumble bee
species, subspecies and infrasubspecies since the starting point of zoological nomenclature in 1758 (from a manuscript

catalogue, unpublished).

distinct from specimens collected from another distant
area, because character variation is apt on average to
be greater (Gaston, Blackburn & Loder, 1995).

For the bumble bee catalogue data, the number of
synonyms (including subspecies, but excluding
infrasubspecies) is correlated with both the date of first
description and the range size of a species independ-
ently of one another, although slightly more of the
variation is explained by variation in range size (par-
tial r, Table 1). Many of the species with large range
sizes, early dates of first formal description and many
synonyms are found in western Europe (i.e. triangles
at the left and upper part of Fig. 4). Most of these
species occur in either the lowland areas of Europe
where early naturalists were most active, such as
Britain, or else are nearly circumpolar in their distribu-
tion.

Curiously, all of the infrasubspecific names (34% of
all names as interpreted at present) belong to the
bumble bee species of the Old World (Fig. 5). Species
of the Old World also have more synonyms and sub-
species per species than do the species of the New
World (numbers of names log-transformed and ex-
cluding 6 Holarctic species, 1,,,.= 3.81 with separate
variance estimates, p< 0.001).
| One possible explanation for the greater numbers
of names per species for bumble bees of the Old
World is that they might have broader distributions

than the species of the New World (see above). This
could arise because the Old World has a slightly
larger total area of suitable habitat (bumble bees oc-
cupy 131 of the 611,000 km? grid cells in the Old
World and 117 in the New), which is apparently
subdivided into fewer well differentiated
biogeographic assemblages of bumble bee species
(e.g. Williams, 19965: fig. 1). However, this explana-
tion is not strongly supported by the bumble bee
data, which show the difference in range sizes
between the two hemispheres to be not significant,
(range sizes log-transformed and excluding 6
Holarctic species, t,,,= —1.24 with separate variance
estimates, p= 0.22). Consequently, while an effect of
differences in habitat area will deserve future consid-
eration, other effects are likely to be more important.

A second possibility is that whereas bumble bee
taxa of uncertain rank may have tended to be re-
garded more often as subspecies in the Old World, in
the New World they may have tended to be regarded
as species (see the discussion below of criteria to
recognise species). While this factor could have con-
tributed to the observed patterns, it is unlikely to
explain why (at a lower rank) so many infrasub-
specific names were described exclusively for taxa
from the Old World.

A third possibility is that the diversity of languages
used for taxonomic publications in the Old World may

Table 1

P.H. WILLIAMS
Results of multiple regression of numbers of synonyms/subspecific names (infrasubspecific names are excluded;

from a manuscript catalogue, unpublished) on date of first formal description for presently accepted species and range size

(number of occupied 611,000 km? grid cells world-wide). Partial r values indicate the correlations with the synonyms
variable after adjusting for the other predictor variable in each case.

log, ,(synonyms+1) = 6.316(+0.969) — 0.003(+0.0005).date + 0.401(+0.057).log, (range)
multiple = 0.72

Fg leo 16 p< 0.0001
partial r Ure Dp
date —0.390 -6.51 < 0.0001
log,,(range) 0.418 7.08

< 0.0001

pascuorum
a

lucorum s.]

humilis

guahuoukS

soroeensjs

terrestris

Fig. 4 Scatterplot of 239 presently accepted bumble bee species by range size (number of occupied 611,000 km? grid cells
world-wide), date of first formal description and numbers of synonyms/subspecific names (infrasubspecific names are
excluded; from a manuscript catalogue, unpublished). The British fauna is distinguished as filled triangles, the nearly

circumpolar fauna (B. hyperboreus, B. balteatus, B. polaris and B. lapponicus) as squares, and some British and
widespread European species are labelled individually.

CHECKLIST OF BUMBLE BEES

1200 |

1000 |

800 }

600

Numbers

400

200

Old World

MMM Species
Synonyms
[= Infrasubspecies

New World

Fig.5 Number of presently recognised bumble bee species, synonyms/subspecific names and infrasubspecific names for
the Old World and the New World (from a manuscript catalogue, unpublished).

have impeded communication and lead to more fre-
quent re-description of taxa than in the New World,
where English was much more dominant (C. O’ Toole,
pers. com.). Again, while this factor is likely to have
contributed to the observed patterns of synonyms, it
does not explain why (at a lower rank) so many
infrasubspecific names were described exclusively for
taxa from the Old World.

Another possible interpretation, which might ex-
plain more of the differences in description dates
between Figs. 1-3 as well as the differences in the
distribution of bumble bee subspecies, synonyms and
infrasubspecific names between hemispheres (Fig. 5),
is that during the twentieth century, effort for describ-
ing the variety of these insects may have become, in
effect, re-directed towards finer distinctions and lower
nomenclatural ranks within known species. This is
perhaps likely as undescribed species became inevita-
bly more difficult to find close to home for the most
active taxonomists, who were based in Europe. Three
lines of evidence are consistent with this explanation.
First, slightly more of the variation in richness of
infrasubspecific names among species is accounted
for by variation in the date of first description of the
species (partial r, Table 2), rather than by variation in
total range size. This is in contrast to the pattern for
synonyms alone (cf. Table 1), although species that are
sufficiently widespread in lowland Europe to include
Britain within their distributions still tend to have high
numbers of both synonyms and infrasubspecific names
(Fig. 6, e.g. B. pascuorum, B. lucorum). A second
intriguing observation is that compared to the number
of authors who have published presently accepted
species names, only one third the number of authors
(20) have published infrasubspecific names, even
though there are nearly four times as many
infrasubspecific names. Indeed, just three of these
authors (Bruno Pittioni, Edgar Kriiger and Alexander

Skorikov) are responsible for 70% of the infra-
subspecific names (all of the species with many
infrasubspecific names had been described before these
three authors became active in publishing infra-
subspecific names between 1910 and 1960, see Fig. 7).
Many similar examples are known from work on
butterflies (R. I. Vane-Wright, pers. com.), with au-
thors choosing a particular favoured species and
describing large numbers of infrasubspecific names
(e.g. Bright & Leeds, 1938). The third point is that the
three most prolific authors all worked in Europe, and
there is a correlation among all 239 bumble bee spe-
cies between the number of infrasubpecific names and
the breadth of the species’ distributions just within
Europe (measured as the number of occupied 611,000
km? grid cells between Britain and the Urals, but
excluding Atlantic islands, North Africa, Turkey and
the Caucasus; Spearman r= 0.67,1,,,= 13.99,p< 0.001).
Thus, a high proportion of the many infrasubspecific
names were published by very few European authors,
for previously described species that are also particu-
larly widespread in Europe.

High numbers of synonyms and infrasubspecific
names for B. terrestris and B. lucorum (subgenus
Bombus) and for B. humilis and B. pascuorum
(subgenus Thoracobombus) in Fig. 6 raise the possi-
bility that large numbers of names are associated with
particular groups of species, perhaps with particular
subgenera. Number of names per species is plotted
against range size per species for subgenera in Fig. 8.
These properties are correlated (log-transformed data,
correlation r= 0.58, F, ,.= 18.16, p< 0.001), but it is the
subgenera with high scores that are more informative.
The subgenus Kallobombus includes many names, but
only a single, very variable species B. soroeensis,
which is broadly distributed in Europe (see below and
Reinig, 1939: fig. 10). The subgenera Alpinobombus
and Laesobombus also have broadly distributed

P.H. WILLIAMS

Table 2 Results of multiple regression of numbers of infrasubspecific names (from a manuscript catalogue, unpublished)
on date of first formal description for presently accepted species and range size (number of occupied 611,000 km? grid
cells world-wide). Species without infrasubspecific names were excluded from the analysis. Partial r values indicate the
correlations with the infrasubspecific names variable after adjusting for the other predictor variable in each case.

log Ginfrasubspecifics+1) = 14.169(+3.638) — 0.007(+0.002).date + 0.742(+0.212).log, (range)

multiple r= 0.66

partial r
date —0.364
log, (range) 0.340

eee 36:57 p< 0.0001

to4 P

-3.79 < 0.001
3.50 < 0.001

20 Ne —

140

120

100
7)
®
5
By 80
L
Oo
®
o
Te) 60
>|
@
= rupestris
= ruderarius
40 s

pomorum

lapponicus

pratorum

0) 10 20 30

lucorum s.|
a

terrestris
A

soroeensis

a humilis
A

pascuorum
a

40 50 60 70 80

Synonyms

Fig.6 Scatterplot of 239 presently accepted bumble bee species by numbers of infrasubspecific names and numbers of
synonyms/subspecific names (from a manuscript catalogue, unpublished). The British fauna is distinguished as filled
triangles, the nearly circumpolar fauna (B. hyperboreus, B. balteatus, B. polaris and B. lapponicus) as squares, and some

widespread European species are labelled individually.

species but relatively few names, perhaps because they
are absent or not abundant in those parts of Europe
where the authors publishing most bumble bee names
have worked, despite several of the species being very
variable in colour pattern (e.g. B. balteatus). In con-
trast, the high ratio of names per species for the
subgenus Bombus shows the keen interest by some
European authors such as Kriiger (1951, 1954, 1956,
1958) in describing the finer points of variation, not so

much within the North American species, but particu-
larly within the widespread European species, B.
terrestris and B. lucorum.

Summary of historical and regional
trends in describing bumble bees

Based on the evidence of asymptotic tendencies in
species-discovery curves, a higher proportion of all

CHECKLIST OF BUMBLE BEES
140

120

100
n
oO
§
S 80
fc)
i
(Ss)
oO
a
a 60
=I
Y .
© soroeensis
= rupesins
= a humilis
Ta ruderanus fa
40 a

pascuorum

pomorum
lapponicus ©
a

consobrinus

a
Se as

1750 1775 1800 1825

1875

keriensis

asiaticus O

hypocrita

1900 1925

1950

1975

2000

Date of first description of species

Fig. 7 Scatterplot of 239 presently accepted bumble bee species by numbers of infrasubspecific names (from a manuscript
catalogue, unpublished) and date of first formal description of species. The British fauna is distinguished as filled triangles,
the nearly cireumpolar fauna (B. hyperboreus, B. balteatus, B. polaris and B. lapponicus) as squares, and some species

with more infrasubspecific names are labelled individually.

species appear to be known for bumble bees than for
many other groups of organisms. Most of these bum-
ble bee species have been described by authors working
in Europe (including European Russia). The species
with the largest geographic range sizes, and particu-
larly the European species with the largest ranges,
have tended to be described first. The same species
have also attracted the highest numbers of synonyms
and subspecific names. As a group, bumble bees have
an unusually high ratio of synonyms and subspecific
names per species, which is otherwise known for some
of the groups of larger and more colourful butterflies.

A few European authors were disproportionately
prolific between 1910 and 1960 in describing finer
variation at infrasubspecific rank, which now accounts
for one third of all bumble bee names. Again, this more
detailed effort has been largely concentrated on the
earlier-described species that are more widespread
within Europe (in contrast, New World bumble bees
have been ignored at this level), presumably because
large samples were more readily accessible to the most
active authors. Determining whether this re-direction
of activity towards lower nomenclatural ranks was a
| logical progression in the recognition of useful taxa, a
| fashion in taxonomic concepts, or in some cases merely

a less disruptive channelling of the enthusiasm of

some authors to publish more names (the ‘mihi itch’),
is beyond the scope of this preliminary review.

All of these patterns in the descriptions of bumble
bees must, as yet, be interpreted with caution. Much
work still remains to be done on the rates of descrip-
tion of taxa at different nomenclatural ranks (species,
subspecies, infrasubspecies), on rates of recognition
of synonymy and of changes in rank, and particularly
on how this activity is partitioned among different time
periods, different geographic regions, different taxo-
nomic subgroups and different authors.

Fundamental to almost all analyses are taxonomic
revisions and checklists of bumble bee species. A
revised checklist is now overdue, because nearly half
(49%) of all names for bumble bees have been pub-
lished since the last synoptic checklist (Skorikov,
1922a).

Development of a revised checklist

To begin to bring a checklist up to date, a draft was
made in 1980 and first circulated for comment in 1985
(Williams, 1985a). This project was developed during
a more detailed study of the west Himalayan fauna
(Williams, 1991) and as part of continuing work on the
large fauna of China in collaboration with Wang S.-f.

Kallobombus

Median names per species

Confusibombus
fo)

Thoracobombus

Bombus s.str
a

0 10 20 30

P.H. WILLIAMS

Laesobombus
O°

Alpinobombus
°

40 50 60 70

Median range size per species

Fig. 8 Scatterplot of 38 bumble bee subgenera by median numbers of all names per species (including synonyms,
subspecific and infrasubspecific names; from a manuscript catalogue, unpublished) and median range size per species
(number of occupied 611,000 km? grid cells world-wide). The subgenera represented in the British fauna are distinguished
as filled triangles and some subgenera are labelled individually.

and Yao J. (unpub.). Some of the broader revisions that
have had the greatest influence on this include works
by Vogt (1909, 1911), Franklin (1913), Stephen (1957),
Milliron (19706, 1971, 1973a, b), Loken (1973, 1984),
Pekkarinen (1979), Reinig (1981), Wang (1982, 1987,
1988), Rasmont (1983, 1988), Thorp ef al. (1983),
Labougle (1990), and especially the publications by
Skorikov (1910-1938) and Tkalcti (1959-1989). In-
evitably, the present checklist cannot be expected to
solve all biological and nomenclatural problems, but it
is hoped that by identifying some of the major prob-
lems it will stimulate further research.

Acknowledgements

My grateful thanks to all who have contributed to the
discussion of this and previous lists, including Donald
Baker, Andreas Bertsch, Barry Bolton, Sydney
Cameron, Gabriela Chavarria, Sally Corbet, Liz Day,
Mick Day, Anne Divers, George Else, Kevin Gaston,
Chris Humphries, Ian Kitching, Astrid Loken, Rod
Macfarlane, Jim Mallet, Russell Miller, Chris O’ Toole,
Antti Pekkarinen, Chris Plowright, Oliver Prys-Jones,
Robert Prys-Jones, Pierre Rasmont, Malcolm Scoble,
Chris Starr, Bill Stephen, Robin Thorp, Borek Tkalcii,

Dick Vane-Wright, Doug Yanega, Yao Jian and Wang
Shu-fang, although they do not necessarily share the
opinions expressed here. I particularly appreciate the
help of Philip Tubbs, Executive Secretary to the ICZN,
for advice on the application of the current Code
(ICZN, 1985) to nomenclatural problems. My thanks
to Julie Harvey and Lorna Mitchell of the Entomology
Library (Dept of Library and Information Services,
NHM) for all their help. I would also like to thank
Wang Shu-fang, Chen Wei and Yao Jian for their
generous hospitality during my visit to China, as well
as the Dept of Botany (NHM) for funding the visit.

TAXONOMY

Bumble bees are a monophyletic group (Williams,
1985b, 1995), constituting the tribe Bombini. They
may be distinguished from other bees (family Apidae)
by the following diagnosis (from Williams, 1991,
which includes descriptions of the characters and dis-
cussion of homologies):

Bombini have the labrum at least twice as broad as long. The

CHECKLIST OF BUMBLE BEES

labrum lacks a longitudinal median ridge, although for the
females it has a strong transverse basal depression. The
clypeus has a transverse subapical depression and the apico-
lateral corners are curved back towards the occiput. A malar
area (= malar space) separates the compound eye from the
base of the mandible, often by a distance greater than the
breadth of the mandible at its base. The hind wings lack a
jugal lobe (= anal lobe). The volsella (= lacinia) of the male
genitalia is greatly enlarged and is produced apically beyond
the gonostylus (= squama).

Bumble bees are large (body length 7-27 mm) robust
insects. Their bodies have a dense covering of variously
coloured long plumose hairs, although these are few or absent
on some parts of the ventral surface of the gaster, on parts of
the propodeum, on parts of of the anterior face of gastral
tergum I, and on parts of the head. The sclerites are usually
black, or lighter brown on the distal parts of the limbs, but are
never marked with bright yellow, red or metallic (= interfer-
ence) colours. The wings may be transparent (= hyaline) to
strongly darkened (= infuscated), but rarely show strongly
metallic reflections.

Female bumble bees have 12 antennal ‘segments’ (= scape,
pedicel and 10 flagellomeres) and six visible gastral terga and
sterna (abbreviated to TI-VI, SI-VI). Males have 13 antennal
‘segments’ (= scape, pedicel and 11 flagellomeres) and seven
visible gastral terga and sterna (abbreviated to TI-VII, SI-
VID).

Where possible, a divisive, ‘top-down’ approach to
the description of bumble bee diversity has been fol-
lowed, in the sense of concentrating initially on
higher-rank relationships and then distinguishing pro-
gressively the species groups, species and then variation
within species (as opposed to beginning with de-
scribed infraspecific taxa and searching ‘upwards’ for
close relatives). At the rank of species, this accepts
those putative species or species complexes that are
supported by consistent evidence for separate status,
and which can be reliably identified throughout their
range for the purpose of mapping distributions. This
kind of broad over-view at least has the potential to
apply consistent criteria across all taxa, even though it
is appreciated that not all taxa at the rank of species are
necessarily of the same kind (Ackery & Vane-Wright,
1984; de Queiroz & Donoghue, 1988). Specialists will
need to modify this list as further information becomes
available for particular species groups.

Phylogeny, supra-specific taxa and
ordering of species

From available cladistic evidence (Williams, 1991,
1995), use of Psithyrus as a genus for the social
parasites separate from the remainder of the social
bumble bees in Bombus can no longer be justified, so
a single genus Bombus is used for all of the species of
bumble bees (see the comments under the subgenus
Psithyrus).

A system of subgenera has become widely used by

specialists who wish to label assemblages of the more
closely similar species. This system is summarised
with subgeneric diagnoses and keys by Richards
(1968). For a review of supraspecific classifications of
bumble bees, see Ito (1985).

The subgeneric system would be more useful if the
names were applied only to strictly monophyletic
groups. Unfortunately, Richards’s (1968) concepts of
the bumble bee subgenera do not always agree well
with recent estimates of phylogeny, because some of
these assemblages now appear to be paraphyletic (e.g.
Mendacibombus) or even polyphyletic (e.g.
Sibiricobombus in the sense of Richards, 1968, in-
cludes Obertobombus, whereas he placed B.
(Sibiricobombus) flaviventris in Subterraneobombus)
(Williams, 1991).

Furthermore, the system of subgenera would prob-
ably be more useful if it were simplified (e.g. Menke &
Carpenter, 1984; and reply by Williams, 1985c). For
example, in the New World, both the monophyletic

fraternus-group of subgenera and the subgenus

Fervidobombus are endemic, and these are the only
two groups represented south of the Panama isthmus.
But whereas Fervidobombus has been treated nearly
consistently as a single, relatively large subgenus (20
species in this list), the fraternus-group (18 species in
this list) has regularly been split into as many as nine
subgenera.

However, no attempt is made in this checklist to
revise radically the subgeneric system, because stabil-
ity will only be served when a revision can be supported
by a comprehensive cladistic analysis. This should
include not only a broad sample of species, but also a
broad range of morphological and molecular charac-
ters. Minor modifications from the subgeneric system
described by Richards (1968) are detailed in the list
after the subgeneric names.

Full synonymy of supraspecific names is included
in this checklist, along with details of type species,
because these have been revised since Richards (1968).
The given generic combination for subgeneric names
is Shown. Where a genus-group name was published at
the rank of genus and subsequently treated at subgeneric
rank, the first such action is listed separately. The two-
letter abbreviations for subgeneric names are based on
those used by Ito (1985).

Species are listed in an order (Table 3) that repre-
sents their phylogenetic relationships (after the
sequencing convention of Nelson, 1972) as these are
currently understood from cladistic studies of the adult
morphology of both sexes (Williams, 1995, and many
references therein). Within subgenera, this informa-
tion is still of a very preliminary nature (e.g. Williams,
1991). Many other estimates of relationship exist and
would result in different sequences of species names.
An alphabetic index is provided as an aid to finding
names in this list.

Table 3 List of names for subgenera of the genus
Bombus, with numbers of species recognised in this
checklist. The subgeneric classification is based on
Richards (1968), modified to accommodate recent
publications (see text; no attempt is made to revise the
subgeneric system, because stability will only be served
when a revision can be supported by a comprehensive
cladistic analysis). Subgenera are listed in an order that
represents their phylogenetic relationships (after the
sequencing convention of Nelson, 1972) as these are
currently understood from cladistic studies of the adult
morphology of both sexes (Williams, 1995).

Subgenus Number of species

1 Mendacibombus 12

2, Bombias 2

3 Confusibombus 1

4 Mucidobombus 1

5 Eversmannibombus 1

6 Psithyrus 29

7 Laesobombus 1

8 Orientalibombus 3

9 Exilobombus 1
10 Thoracobombus 19
11 Tricornibombus 3
12 Fervidobombus 20
13 Senexibombus 4
14 Diversobombus 4
15 Megabombus 14
16 Rhodobombus 3
17 Kallobombus 1
18 Alpinobombus 3)
19 Subterraneobombus 9
20 Alpigenobombus 6
2] Pyrobombus 43
22 Festivobombus 1
23 Rufipedibombus 2
24 Pressibombus 1
5 Bombus s.str. 10
26 Cullumanobombus 4
27 Obertobombus 2,
28 Melanobombus 14
29 Sibiricobombus 5
30 Fraternobombus 1
31 Crotchtibombus 1
32 Robustobombus 3)
33 Separatobombus 2
34 Funebribombus 2}
35 Brachycephalibombus 2
36 Rubicundobombus 1
37 Coccineobombus 2
38 Dasybombus 2

Criteria to discriminate species

It is not possible or appropriate to discuss species
concepts in detail in this paper (though the selected
references provide some introduction; for recent re-
views, see Claridge et al., 1997; Mallet, 1997).
However, in order to interpret the checklist, where
possible it would be useful to make the species-dis-
criminating criteria explicit. It is equally important to
convey the present belief that there is no simple solu-

P.H. WILLIAMS

tion to the problem, and that no single known approach
can resolve all of the cases in a uniform and entirely
satisfactory manner.

Species concepts (ideas or general notions of the
class of objects) and species diagnoses (operational
determinations of individual objects) are contentious
and probably unresolvable issues. Therefore there is
arguably no single ‘true’ list of species, only more or
less valid interpretations from different viewpoints.

Unresolveable conflicts may arise from opposing
views of the nature of species. Species have been
regarded either as typological classes, with member-
ship to be defined by some shared essence (reviewed
by Templeton, 1981), or as individuals, to be discoy-
ered (Ghiselin, 1975). There are also conflicting
opinions concerning criteria (characteristics or stand-
ards by which an object may be judged) for recognising
species, based in part on differing emphasis on pattern
or process (de Queiroz & Donoghue, 1988).

Species may be considered not to differ from taxa at
other ranks (e.g. genera, subspecies) in any qualitative
way. There may be quantitative differences in the
numbers of character differences that distinguish them
in comparison with taxa of lower rank. For example,
according to Mallet (1995:294), Darwin (1859) held
this view. The problem with quantitative criteria
(whether applied to genetic or phenotypic characters)
is there is no reason to believe that any choice of
threshold in the degree of difference used to recognise
taxa at the rank of species is anything other than
essentially arbitrary and thus idiosyncratic to particu-
lar authors.

In another view, species may be considered to differ
qualitatively from taxa at other ranks. It is widely
accepted, though often implicitly, that taxa at the rank
of species should be recognised so as to mark the
boundary between, on the one hand, reticulate rela-
tions (for sexually reproducing organisms), and on the
other, more consistently divergent genealogical rela-
tions. One problem is that this distinction may require
predictions as to whether or not currently distinct
groups of individuals are likely to show reticulate
relationships again in the future.

Interbreeding and the associated genetic recombi-
nation is an important part of Dobzhansky’s (1937)
‘modern synthesis’ of Mendelian genetics with Dar-
win’s natural selection theory for evolution.
Emphasising interbreeding as a criterion for recognis-
ing species characterised what Mayr (1940, 1963)
called the ‘biological’ species concept. These ideas
have been modified in the recognition concept of
species (Paterson, 1985). One problem with inter-
breeding or mate recognition as criteria for recognising
species is that direct and reliable evidence is rarely
available and the results of tests under artificial condi-
tions cannot necessarily be generalised (Splitter, 1982).
Another is that the capacity for interbreeding is an

——eE

CHECKLIST OF BUMBLE BEES

ancestral condition (i.e. not an homology) and so
cannot provide support for recognising taxa in the
phylogenetic sense (Rosen, 1979).

In practice, all that is usually available to discrimi-
nate species as ‘different’ is evidence from character
differences and their patterns of concordance among
individuals. The phylogenetic species concept
(Cracraft, 1989) is popular because it also embodies
the notion that species mark the boundary between
different patterns of relationship among individuals
and yet it does not rely on inference of interbreeding.
The problem is that discovery of phylogenetic species
as minimum cladistically-diagnosable (discrete) groups
of individuals requires that these groups uniquely
share homologies (synapomorphies), which may not
always be the case (Ackery & Vane-Wright, 1984;
Frost & Kluge, 1994).

Mallet (1995) has argued for minimising the number
of assumptions built into species concepts. He sug-
gests that two nominal taxa should be considered
conspecific until it can be demonstrated that data for
multiple characters distinguish consistent subgroups
of individuals with few or no intermediates (the char-
acter-cluster concept of species). Although he was
arguing against the use of the widely-held biological
species concept, he recognised that his prescription
differs little from recent common practice. The prob-
lem with the cluster concept is how to decide on a
threshold for permissable numbers of intermediate
individuals between taxa for them still to be consid-
ered separate species.

Ultimately, species may be seen as useful conven-
tions to aid in the communication of information
gathered about the individuals that are their parts. It
may be argued that the most important initial goal is to
describe the nature of the variation in each particular
case and to avoid presenting only theory-laden (and
constrained) interpretations. In this way, basic infor-
mation on variation will remain available for
re-interpretation as theory changes.

For the sake of illustration, four principal classes of
problems in geographical variation may be distin-
guished within the spectrum of kinds of relationships,
with the following examples:

Broad co-occurrence of differing individuals

Skorikov (1931) and Reinig (1935) recognised that
throughout much of the range of B. keriensis, both
yellow-banded and cream- or white-banded individu-
als with indistinguishable morphology co-occur (Fig.
9). From available evidence, it is possible that B.
niveatus / vorticosus may show a similar pattern of

_ yellow/white variation, as may B. impetuosus/potanini,

although with differing degrees of geographical varia-
tion in colour-form frequency (see the comments on
these species). Consequently, taxa in these pairs are

also treated as conspecific for the present (it is possible
that in some cases such colour differences may be
controlled by alleles at a single locus, see Owen &
Plowright, 1980, on B. melanopygus; and Williams,
1991, on B. asiaticus; or by small numbers of loci, see
Plowright & Owen, 1980, on B. rufocinctus). In con-
trast, although the yellow-banded B. shaposhnikovi
and the white-banded B. handlirschianus also show a
broadly-overlapping pattern of distribution, the one
white-banded male that I have seen is distinct from the
yellow-banded males in the morphology of its genita-
lia (Williams, 1991).

Broad clinal variation

Many species show broad trends in variation across
continents, most obviously in colour pattern (e.g. B.
cingulatus, Fig. 10; and the trifasciatus-group, Fig.
13, which may be combined with locally convergent
colour variation, e.g. within the haemorrhoidalis-
group, breviceps-group and rotundiceps-group, see
Sakagami & Yoshikawa, 1961; Tkalcii, 19685, 1989).
In North America, several pairs of nominal taxa were
described originally from individual type-specimens
with differing colour patterns from eastern and west-
ern regions respectively (e.g. B. auricomus /nevadensis,
B. fervidus / californicus, B. pensylvanicus / sonorus,
B. terricola / occidentalis). These taxon pairs have
long caused difficulties, for example with Franklin
(1913:239) commenting on a list including these taxa
and others that are now considered conspecific that “it
must be entirely a matter of personal opinion whether
they should be given full species rank or be considered
as only subspecies’ (although, intriguingly, B.
auricomus /nevadensis were not included in Franklin’s
list). In at least some of these cases, many individuals
with what appears to be a continuum of intermediate
colour patterns are now known from broad intervening
areas, so that threshold criteria for distinguishing these
taxa appear to be essentially arbitrary (e.g. making
decisions based on whether a particular tergum has the
pubescence entirely yellow, rather than having a few
black hairs present). In consequence, taxa in these
taxon pairs are treated here as conspecific and maps
are compiled for the more clearly recognisable, more
inclusive taxa (but see the comments on B. auricomus
/ nevadensis).

Narrow hybrid zones

In some cases, otherwise discrete colour forms with
closely similar morphology meet in narrow zones (of
the order of a few km in breadth), where there may be
evidence of intermediate or genetically recombinant
individuals. In Europe this is best known for B.
ruderatus / argillaceus (Fig. 11; Scholl, Obrecht &
Zimmermann, 1992), and inAsia it has been suggested

oD P.H. WILLIAMS

| § ‘

Fig.9 Approximate distribution range (area within the dotted line) and principal colour variation for B. keriensis from Reinig
(1939: fig. 23). Many more records are available now, but the pattern remains similar, with broad overlap of yellow- and
white-banded individuals in Mongolia, Tien Shan, Pamir and western Himalaya. Yellow and cream pubescence is shown on
the bees by crosses; red pubescence by vertical hatching.

Fig. 10 Distribution records (spots), approximate range (area within the line) and principal colour variation for B.
cingulatus in the northern Palaearctic Region from Reinig (1939: fig. 7). The lightest individuals occur in the east
(Kamchatka) and the darkest individuals (with the black thoracic band) occur in the west, with intermediate individuals in
intervening areas.

CHECKLIST OF BUMBLE BEES 93

ombus
: /uderans
suderatis abr OTGUMACEUS SCOP.

Fig. 11 Distribution records (spots), approximate range (area within the cross-hatching, left, and line, right) and principal
colour variation between queens of B. ruderatus and B. argillaceus in Europe from Reinig (1939: fig. 7). These taxa were
regarded as subspecies by Reinig, but have recently been treated as separate species. Although there is evidence of a hybrid
zone between some areas of parapatry, the hybrid individuals are very rare (Scholl, Obrecht & Zimmermann, 1992).
Yellow pubescence is shown on the bees by crosses.

Fig. 12 Distribution records (spots) and principal colour variation for B. asiaticus in Kashmir from Williams (1991: map
48). There is evidence of a hybrid zone between some areas of parapatry, such as some high passes along the divide of the
Great Himalaya Range, where there are abundant hybrid individuals. The spot symbols show the locally most abundant
colour pattern. Yellow pubescence is shown on the bees by fine stippling; red pubescence by vertical hatching.

P.H. WILLIAMS

Fig. 13 Distribution records and principal colour variation within the trifasciatus-group in Asia (updated from Williams,
1991: fig. 11). The individuals may all be considered parts of a single species, B. trifasciatus, depending on which
species-defining criterion is accepted. The dashed line shows the 1000 m contour above sea level and the solid line
shows the 4000 m contour. Yellow pubescence is shown on the bees by fine stippling, orange pubescence by coarse

stippling, red pubescence by vertical hatching.

CHECKLIST OF BUMBLE BEES

«
On | in
On

for B. asiaticus / longiceps (Fig. 12; Williams,
1991). Other possible examples include B.
lapponicus / monticola in Europe and B. pyrosoma /
friseanus / miniatus in China. In the case of B.
asiaticus / longiceps, | have treated them as
conspecific, because intermediate individuals greatly
outnumber ‘typical’ individuals at some localities.
For the other cases, I have followed earlier treat-
ments of these taxa as separate species, because
intermediate individuals are rare or not well known
(although this may be a consequence of poor sam-
pling in some inaccessible areas).

Disjunct peripheral populations

Some peripheral populations on offshore islands or
habitat islands (e.g. mountains) show some diver-
gence in colour pattern with little morphological
divergence. European examples include B. terrestris
/ canariensis and B. hortorum / reinigiellus. Asian
examples include B. schrencki / honshuensis, B.
trifasciatus / maxwelli (Fig. 13, Peninsular Malay-
sia), B. trifasciatus / wilemani (Fig. 13, Taiwan), B.
breviceps / angustus, B. parthenius / sonani, B.
flavescens / rufoflavus and B. flavescens /
baguionensis. For the application of the biological
species concept, in these cases there is no ‘natural’
meeting of individuals between the taxon pairs and
so no admissible evidence on interbreeding (Splitter,
1982). For the application of Mallet’s (1995) cluster
concept, quantitative analysis of patterns of variation
is urgently needed. Where this information is absent,
I agree with his prescription of treating taxa in these
taxon pairs as provisionally conspecific. Bombus
honshuensis and B. schrencki are mapped separately
here because, from published accounts and a small
sample of material examined, their colour differ-
ences appear to coincide with stronger and more
consistent morphological distinctions.

It is hoped that further information may help to
clarify these cases. In the interests of pluralism, I aim
to report not only a preferred interpretation in the
comments on each species, but also at least the more
widely-held alternative interpretations.

Sub-specific taxa

For this checklist the interest is primarily in problems
of recognition and nomenclature for taxa at the rank of
species. Subspecific names refer to parts of species,
and so for present purposes these can be treated as
synonyms of specific names (e.g. Schwarz ef al.,
1996). This is not to say that subspecific taxa should
not be recognised if they are considered useful, and of
course other biologists may add subspecies to this list
(cf. Rasmont et al., 1995).

P.H. WILLIAMS

NOMENCLATURE

Nomenclature should be seen as the servant of biol-
ogy: its purpose is to provide labels that enable
biologists to communicate information about organ-
isms with minimal confusion concerning the organisms
to which they refer. Accounts of the history of nomen-
clature for many groups of organisms (e.g. on British
bumble bees: Alford, 1975; Prys-Jones & Corbet,
1987:82) show that this is not a trivial matter and that
rules are necessary.

Treatment of names follows the /nternational Code
of Zoological Nomenclature (International Commis-
sion on Zoological Nomenclature [{CZN], 1985). The
Principle of Priority is generally adhered to, although
regard is given to the stated purpose of priority (ICZN,
1985: Article 23b): namely that it should be used to
promote stability and is not intended to be used to
upset a long-accepted name in its accustomed mean-
ing (Article 79c) through the introduction of an unused
name that is its senior synonym. Similar action is also
suggested where cases of homonymy affect current
usage, although this action cannot be taken when it is
felt desirable to maintain availability of a senior homo-
nym. My suggestions for applications to ICZN for
conservation of names in current use are indicated by

stars (@).

Typographical conventions

Bombus b-us valid name in the species group,

c-us available name in the species group,
including synonyms of a valid spe-
cies name,

2d-us available name in the species group,

a provisional synonym of a valid

species name,

unavailable name, informally asso-

ciated with a valid species name,

jf-us examined type material for species-group name
f-us examined (in whole or in part),

[e-us]

@ comments on status of species,
O comments on application of names,
Ok) suggestion for application to ICZN.

?Bombus g-us_ valid name in the species group, fora
taxon that is recognised provision-
ally as a separate species from B.

b-us.

A question mark (?) before a valid name shows that,
while it refers to a taxon that is considered likely to be
a separate species, it may be conspecific with the
preceding taxon in the list (1.e. while Bombus g-us may
be conspecific with Bombus b-us, Bombus d-us is
much more likely to be conspecific with Bombus b-
US).

CHECKLIST OF BUMBLE BEES

Names in the more detailed references are followed
by names of authors, date of first publication (within
the meaning of ICZN, 1985), and page reference.
Wherever possible, the true first date of publication is
given in preference to any purported date of publica-
tion when these differ. If a name were published
originally in a different generic combination, then the
original genus is shown in brackets. If the name had
been published originally with a different termination,
or with capital initial letters, diacritic marks etc., then
the original form is shown without the mandatory
changes (with the exception that small capital letters
are reduced to lower case).

Selection of synonyms

This checklist is based on a much longer catalogue of
over 2800 names. As a checklist, it is not required to
include the full list of synonyms, so synonyms are
selected for this list primarily where they help to
clarify the identity and scope of the species (including
the subspecies included by some authors), particularly
with reference to those names in most common use in
the literature of the last 25 years. Misidentifications
are not included with the lists of synonyms and are
discussed only when necessary to clarify the applica-
tion of problematic names.

Applications to ICZN

Flexibility in interpretation of the status of taxa is
possible where the evidence to distinguish among
interpretations is absent, inconclusive, or may permit
different interpretations under different species con-
cepts. Otherwise flexibility in the application of names
depends on whether systematists are eager to apply to
the International Commission on Zoological Nomen-
clature to use its Plenary Power in order to conserve a
preferred usage of names (e.g. Laken et al., 1994;
ICZN, 1996).

I propose that this could be achieved in a single
application to include all names for which action is
currently known to be required (atratus, balteatus,
distinguendus, flavifrons, humilis, hyperboreus,
mesomelas, mixtus, norvegicus, polaris, pyrenaeus,
soroeensis and variabilis). Comments on this proposal
would be welcomed.

DISTRIBUTION MAPS

This checklist was compiled in conjunction with dis-
tribution data in support of biogeographic studies.

Maps of world-wide distribution at a coarse grain size
were designed for use in comparisons of regional
bumble bee faunas (e.g. Williams, 1989, 1991, 1993,
1995, 1996a, b; Williams & Seddon, 1993; Williams
& Humphries, 1996).

Aside from any difficulties in identifying species or
localities, comparisons among faunas are complicated
by two principal factors: first, by differences in sam-
pling effort (as illustrated by ‘species-accumulation
curves’, e.g. Colwell & Coddington, 1994); and sec-
ond, by differences in the extent of sampling areas
(‘species-area effects’, e.g. Connor & McCoy, 1979).
Fortunately for the first problem, the attractiveness of
bumble bees to collectors has ensured that they have
been relatively intensively sampled, so that most fau-
nas are relatively well known. But in order to reduce
this problem further, rather than extrapolate local rich-
ness and lose information on individual species, the
expected distributions of some species are interpo-
lated on the basis of knowledge of their habitat
associations (see the legend to Fig. 14). To reduce the
second problem of species-area effects, equal-area
grid cells were established using a cylindrical, equal-
area projection of the world, marked at intervals of 10°
of longitude and calculated intervals of latitude (Fig.
14). However, this does not ensure equal land areas
among grid cells, or equal areas of habitat suitable for
bumble bees.

Because the intention is to study biogeographic
patterns, maps are required to show all historical
records, including data from areas where species may
now be extinct. On the other hand, data exclude fossil
taxa (reviewed by Zeuner & Manning, 1976) and
documented introductions (e.g. Oliff, 1895; Frison,
1925b; Gurr, 1957; Prys-Jones et al., 1981; Arretz &
Macfarlane, 1982; Cardale, 1993).

The maps for every species are not included with
this checklist because many data are still being col-
lected, although for each subgenus a preliminary map
of species richness is included as a general guide (or
for monotypic subgenera, a map of records for the
single species is included). The numerical values for
the grey-scale classes differ between maps and are
not shown. This is because I have adopted an alterna-
tive approach of using equal frequency classes, which
have the advantage that each grey-scale class remains
consistent in its relative richness among all maps (e.g.
dark grey always shows the richest one fifth of occu-
pied cells excluding the maximum etc.). The sources
of the distribution data have not been included be-
cause this will be included in a later atlas.

98 P.H. WILLIAMS

Key to map symbols:

Maps for single species

specimens examined,

precise literature records (e.g. ‘Dungeness TRO1, UK’),

vague locality data (e.g. “Florida’),

interpolations of expected distribution (following common practice for range-filling maps; the
rules adopted here are to fill cells between occupied cells when filled cells are known to have had
a high proportion of suitable habitat within recorded history; these records amount to < 10% of all
gridcell records at this scale, Williams, 1993).

OOS®e

Maps for multiple species

maximum species counts are shown in black, otherwise counts are divided into five grey-scale
classes of approximately equal size by numbers of grid cells.

Fig. 14 Map of the world (excluding Antarctica) using a cylindrical equal-area projection that is orthomorphic (minimum
shape distortion) at 46° North and South (where bumble bee records are particularly plentiful). Intervals of 10° longitude
(top of map) are used to calculate intervals of latitude (right of map) that provide equal-area grid cells of c. 611,000 km?.
The portion of the grid shown covers the known, native distribution of bumble bees. Map symbols are shown above for (a)
plotting individual species, for which different spots distinguish different data categories (Map 3); or (b) for plotting
coincidence maps for multiple species, using a grey scale for variation in species richness (Map 1).

CHECKLIST OF BUMBLE BEES

LIST OF SPECIES

(plot of total species richness with grey scale, for
explanation see Fig. 14)

Genus BOMBUS Latreille in the broad sense

[Bremus [Jurine], 1801:164, type-species Apis terrestris
Linnaeus (= Bombus terrestris (Linnaeus)) by subse-
quent designation of Morice & Durrant, 1915:429,
suppressed by ICZN, 1939]

Bombus Latreille, 1802a:437, type-species Apis terrestris
Linnaeus (cited asApis terrestris F.) (=Bombus terrestris
(Linnaeus)) by monotypy

Bombus Latreille, 1802b:385, type-species Apis terrestris
Linnaeus (=Bombus terrestris (Linnaeus)) by monotypy,
redescribed

[Bremus Panzer, 1805:pl. 19-21, type-speciesApis agrorum
Fabricius (= Bombus pascuorum (Scopoli)) by subse-
quent designation of Sandhouse, 1943:532, suppressed
by ICZN, 1954]

[Bombellus WE, 1931:248, incorrect subsequent spelling]

Subgenus MENDACIBOMBUS Skorikov
Mendacibombus Skorikov, 1914a:125, type-species
Bombus mendax Gerstaecker by subsequent designa-
tion of Sandhouse, 1943:572
Bombus (Mendacibombus) Kriiger, 1917:62

COMMENT. The species of Mendacibombus appear
to be paraphyletic with respect to the rest of the

oY)

bumble bees and in consequence are not a ‘natural’
group (Williams, 1991, 1995).

Bombus (Md.) avinoviellus (Skorikov)
avinoviellus (Skorikov, 1914a:126 [Mendacibombus]) ex-
amined
callophenax Cockerell, 1917:122, examined

Bombus (Md.) mendax Gerstaecker
mendax Gerstaecker, 1869:323, examined
latofasciatus Vogt, 1909:50, not of Vogt, 1909:42 (= B.
lucorum (Linnaeus))
pyrenes (Tkalcit, 1975:173 [Mendacibombus]) replace-
ment name for latofasciatus Vogt, 1909:50

Bombus (Md.) makarjini Skorikov
makarjini Skorikoy, 1910a:329, examined

Bombus (Md.) superbus (Tkalcit)

superbus (Tkalctt, 1968a:22 |Mendacibombus]) examined

Bombus (Md.) himalayanus (Skorikov)

?varius (Skorikoy, 1914a:125 [Mendacibombus]) exam-
ined, not of Lepeletier, 1832:381 (= B. campestris
(Panzer))

himalayanus (Skorikov, 1914a:127 [Mendacibombus})
examined

Bombus (Md.) marussinus Skorikov
marussinus Skorikov, 1910a:330, examined
afghanus Reinig, 1940:230, examined

Bombus (Md.) turkestanicus Skorikov
turkestanicus Skorikoy, 1910a:329, examined

Bombus (Md.) defector Skorikov
defector Skorikov, 1910a:330
?altaicus Skorikov, 1910a:329, not of Eversmann,
1846:436 (= B. melanurus Lepeletier)
?margreiteri Vogt, in Skorikov, 1910a:330, examined

@ TAXONOMIC sTATUS. Skorikov’s (1910a) de-
scriptions of varieties of B. mendax are all of females.
Many of these nominal taxa have subsequently been
treated as separate species (e.g. Skorikov, 1931;
Rasmont, 1988).

However, I have examined type material or other
material identified by Skorikov for all of these taxa
and find some of them to be morphologically closely
similar. The females of defector, altaicus and
margreiteri differ from one another principally in

100

colour, and the only males I have seen associated with
them (collections in London, Petersburg, Beijing) have
very similar genitalia (which are distinct from B.
mendax).

Until more evidence to the contrary is available
from critical studies of patterns of variation, I shall
continue to treat B. defector, B. altaicus and B.
margreiteri as parts of a single variable species, B.
defector (Williams, 1985a, 1991).

O NOMENCLATURE. Williams (1991) regarded B.
defector, B. altaicus and B. margreiteri as likely to be
conspecific and following the Principle of First Re-
viser (ICZN, 1985: Article 24) chose B. defector as the
name for the species.

Bombus (Md.) handlirschianus Vogt
Handlirschianus Vogt, 1909:49

Bombus (Md.) shaposhnikovi Skorikov
shaposhnikovi Skorikoy, 1910a:329

Bombus (Md.) waltoni Cockerell
chinensis Skorikov, 1910a:330, examined, not of Morawitz,
1890:352 (= B. chinensis (Morawitz))
waltoni Cockerell, 1910b:239, examined

Bombus (Md.) convexus Wang
lugubris Morawitz, 1880:339, examined, not of
Kriechbaumer, 1870:159 (= B. maxillosus Klug)
convexus Wang, 1979:190, examined

Subgenus BOMBIAS Robertson

Bombias Robertson, 1903:176, type-species Bombias
auricomus Robertson (?= Bombus nevadensis Cresson)
by original designation

Bombus (Bombias) Franklin, 1913:138

Nevadensibombus Skorikoy, 1922a:149, type-species
Bombus nevadensis Cresson by subsequent designation
of Frison, 1927:64

Bremus (Boopobombus) Frison, 1927:59 (proposed as a

P.H. WILLIAMS

section name but stated by Frison to include those forms
considered by Franklin, 1913, to belong to the subgenus
Bombias Robertson), type-species Bombias auricomus
Robertson (= Bombus auricomus (Robertson)) by sub-
sequent designation of Williams, 1995:339.

Bombus (Bi.) nevadensis Cresson
nevadensis Cresson, 1874:102

COMMENT. A single queen of B. nevadensis has
been reported from Hidalgo, Mexico, by Milliron
(1971) and Hurd (1979), although the species is not
listed for Mexico by Labougle (1990).

?Bombus (Bi.) auricomus (Robertson)
auricomus (Robertson, 1903:176 [Bombias])

@ TAXONOMIC STATUS. B. nevadensis and B.
auricomus have been regarded both as conspecific
(e.g. LaBerge & Webb, 1962; Milliron, 1971; Thorp er
al., 1983; Laverty & Harder, 1988) and as separate
species (e.g. Franklin, 1913; Rasmont, 1988; Scholl,
Thorp, Owen & Obrecht, 1992; Poole, 1996).

B. nevadensis from western North America was not
mentioned in the original description of B. auricomus
(lectotype worker from Illinois by designation of
Milliron, 1971:78), although the latter was described
using characters of morphology and of colour pattern.
The two taxa have generally been distinguished on the
basis of the extent of the black pubescence on the
dorsum of the female thorax and laterally on the male
gastral terga (e.g. Franklin, 1913).

The only study to investigate variation in characters
used to distinguish the two taxa at a fine spatial scale in
their area of overlap was by LaBerge & Webb (1962).
They reported (p. 26) that “Throughout the broad
middle half of Nebraska nevadensis seems to be rather
rare and most specimens, although referable to sub-
species auricomus show some indication of
intergrading with the typical subspecies [nevadensis]
in the west. . .. Many specimens from Nebraska in the
range of the typical subspecies [nevadensis| show
some tendency toward the darker coloration of subspe-
cies auricomus. They concluded that these variable
bees are all parts of the same species.

Recently, Scholl et al. (1992) distinguished two
groups of individuals on the basis of differing mobility
morphs of five enzymes. The individuals in one en-
zyme group were all extensively dark-banded, and
Scholl et al. associated these with the name B.
auricomus. However, individuals in the other enzyme
group, which Scholl et al. associated with the name B.
nevadensis, apparently included not only the contrast-
ing, extensively pale individuals (B. nevadensis), but
also a few of the extensively dark-banded individuals
(B. auricomus) similar to those in the first group (8/49

CHECKLIST OF BUMBLE BEES

individuals had gastral tergum I almost completely
black; 3/49 individuals had the scutellum predomi-
nantly black). Thus the enzyme evidence does identify
two groups of individuals, but (1) these do not appear
to correspond precisely to the two traditional colour
groups; (2) some of the key areas likely to support
intermediate or recombinant individuals still need to
be sampled for enzyme variation (e.g. in the Dakotas,
L. Day in litt.); and (3) inheritance of enzyme and
colour states needs to be better understood, including
the unusual enzyme morphs of the heterozygous bees
(detected in 20/141 queens). They concluded that
these bees represent two species.

A. Scholl (in litt.) reports a further intriguing
morphometric study. A random subsample of 20 queens
from the enzyme study was scored for 15 characters
and analysed by linear discriminant analysis. This
method seeks a combination of characters that best
discriminates any two a priori sets, in this case using
three measurements of parts of the radial cell, eye and
antenna. However, although this approach may be
useful for discriminating previously recognised taxa,
it does not provide evidence that they are necessarily
separate species (it could also be used to discriminate
morphological subsets within a single, variable popu-
lation, e.g. among breeds of domestic dogs).

From an examination of 41 females, so far I have
found only one subtle morphological character to
distinguish eastern, banded bees (B. auricomus), on
the one hand, from western unbanded (B. nevadensis)
and banded (e.g. Vancouver Island) bees, on the other.
This concerns the anterior part of a band of large
punctures along the inner eye margin, dorsally oppo-
site the ocelli, just before these punctures meet a more
anterior, very dense patch of small punctures. The
western bees have areas between the large punctures
conspicuously shining, with few fine punctures and
lacking microsculpture. In contrast, the eastern bees
have these areas appearing rather dull, often with more
of the fine punctures, and more particularly with a very
fine, wrinkled or reticulate microsculpture. A similar
difference may be present in the males, posterio-
laterally to the ocelli, though the sample sizes available
to me are too small for much confidence.

I regard the conflicting evidence available at present
as not entirely conclusive as to whether these bees are
parts of the same population or two separate species.
As far as is known, both the variations of the colour
pattern and of the enzyme mobilities are inherited and
genetically determined, but details of patterns of in-
heritance and of the spatial aspects of any association
between these characters are unknown. In view of the
multiple enzymes differences found and of the appar-
ent association between the enzyme groups and the
morphological character states, I shall follow the treat-
ment of these taxa as two separate species until more
evidence is available.

(plot of records for a single species, for explanation and key
see Fig. 14)

Subgenus CONFUSIBOMBUS Ball

Bombus (Confusibombus) Ball, 1914:78, type-species
Bombus confusus Schenck by monotypy

Bombus (Sulcobombus) Kriiger, 1917:65, type-species
Bombus confusus Schenck by subsequent designation
of Sandhouse, 1943:602

Confusobombus Skorikoy, 1922a:156, type-species
Bombus confusus Schenck by subsequent designation
of Richards, 1968:214

Bombus (Cf.) confusus Schenck
confusus Schenck, 1859:135
paradoxus Dalla Torre, 1882:18
festivus Hoffer, 1882:80, not of Smith, 1861:152 (= B.
festivus Smith)

@ TAXONOMIC STATUS. B. confusus and B.
paradoxus differ in the colour pattern of the pubes-
cence (e.g. Reinig, 1939: fig. 19). Rasmont (1988)
reports that in north western Europe, the yellow-banded
and white-tailed B. paradoxus occurs only as rare
individuals within the population of predominantly
unbanded and red-tailed B. confusus. In contrast, all of
the individuals that I have seen from the disjunct
population in Central Asia have the yellow-banded
and white-tailed B. paradoxus colour pattern.

Subgenus MUCIDOBOMBUS Kriiger
Mucidobombus Kriiger, 1920:350, type-species Bombus
mucidus Gerstaecker by monotypy
Bombus (Mucidobombus) Pittioni, 1937:97

102

Bombus (Mc.) mucidus Gerstaecker
mucidus Gerstaecker, 1869:324
atratus Friese, 1911:572, examined

Subgenus EVERSMANNIBOMBUS Skorikov
Agribombus (Eversmannibombus) Skorikoy, 1938a:145,
type-species Mucidobombus eversmanniellus (=

Bombus persicus Radoszkowski) by monotypy

Bombus (Eversmannibombus) Richards, 1968:214

Bombus (Ey.) persicus Radoszkowski

calidus Eversmann, 1852:133, examined, not of Erichson
in Middendorff, 1851:65 (= B. hypnorum (Linnaeus))

persicus Radoszkowski, 1881:v, examined

Persicus Radoszkowski, 1883:214, redescribed

eversmanni Friese, 1911:572, not of Skorikov, 1910¢:581
( B. modestus Eversmann), replacement name for
calidus Eversmann, 1852:133

eversmanniellus (Skorikoy, 1922a:149 [Mucidobombus])
replacement name for eversmanni Friese, 1911:572

Subgenus PSITHYRUS Lepeletier
Psithyrus Lepeletier, 1832:373, type-species Apis rupestris
Fabricius (= Bombus rupestris (Fabricius)) by subse-
quent designation of Sandhouse, 1943:572
Apathus Newman, 1835:404, replacement name for
Psithyrus Lepeletier, incorrectly stated to be a junior
homonym of Psithyros Hiibner, [1819]:132 (=
Macroglossum Scopoli, 1777:414)

P.H. WILLIAMS

?Psithyrus (Laboriopsithyrus) Frison, 1927:69, type-spe-
cies Bombus laboriosus Fabricius (= Emphoropsis
laboriosus (Fabricius) in the sense of Frison (=Bombus
citrinus (Smith), a misidentification, see Milliron,
1960:99, requiring designation by ICZN) by original
fixation &

Psithyrus (Ashtonipsithyrus) Frison, 1927:69, type-spe-
cies Apathus ashtoni Cresson (= Bombus ashtoni
(Cresson)) by original designation

Psithyrus (Fernaldaepsithyrus ) Frison, 1927:70, type-spe-
cies Psithyrus fernaldae Franklin (= Bombus fernaldae
(Franklin)) by original designation

Psithyrus (Eopsithyrus) Popov, 1931:134, type-species
Apathus tibetanus Morawitz (= Bombus tibetanus
(Morawitz)) by original designation

Psithyrus (Metapsithyrus) Popov, 1931:135, type-species
Apis campestris Panzer (= Bombus campestris (Pan-
zer)) by original designation

Psithyrus (Allopsithyrus) Popov, 1931:136, type-species
Apis barbutella Kirby (= Bombus barbutellus (Kirby))
by original designation

Psithyrus (Ceratopsithyrus) Pittioni, 1949:270, type-spe-
cies Psithyrus klapperichi Pittioni (= Bombus cornutus
(Frison)) by original designation

Psithyrus (Citrinopsithyrus ) Thorp inThorp et al., 1983:50,
type-speciesApathus citrinus Smith (=Bombus citrinus
(Smith)) by original designation

Bombus (Psithyrus) Williams, 1991:44

[Psithyrus (Fernaldepsithyrus) Amiet, 1996:86, incorrect
subsequent spelling]

@ TAXONOMIC sTATUS. It has long been consid-
ered useful to regard Psithyrus as a separate genus in
recognition of the distinctive behaviour of the species,
as social parasites in colonies of the remaining
Bombini, and in recognition of their distinctive mor-
phology. However, most recent studies have shown (if
phenograms are interpreted along with cladograms as
phylogenetic estimates) that, although Psithyrus is
itself very likely to be monophyletic, the remaining
bumble bees are not (Plowright & Stephen, 1973;
Obrecht & Scholl, 1981; Ito, 1985: Williams, 1985),
1991, 1995; Pamilo et al., 1987).

I have previously attempted to retain the use of the
names Psithyrus and Bombus for monophyletic genera
by recognising a third genus, Mendacibombus
(Williams, 1985b). However, further study of all of the
species of Mendacibombus (Williams, 1991, 1995)
showed that it is likely to be paraphyletic with respect
to all other bumble bees, with the consequence that as
many as another nine genera (mostly for single spe-
cies) might be required to maintain monophyly
alongside a genus Psithyrus. In the face of this evi-
dence, a pragmatic solution was recommended,
recognising a single genus Bombus for all bumble
bees, to include Psithyrus as a subgenus. This is a
return to an emphasis of the more widely shared
characters and the more distant affinities for the ge-
neric concept, encouraged by the opinion of Michener
(1990) that bumble bees are ‘morphologically mo-

CHECKLIST OF BUMBLE BEES

notonous’ in comparison with variation among species
within closely related groups such as Euglossini (or-
chid bees) and Meliponini (stingless bees). One
advantage of a single genus for all bumble bees is that
it recognises a group for which evidence of monophyly
is particularly strong, so that nomenclature is most
likely to remain stable in the future. Use of a single
genus Bombus for all bumble bees (Williams, 1991)
has now been accepted by most recent authors (e.g.
Rasmont & Adamski, 1995; Rasmont ef al., 1995;
Schwarz et al., 1996).

The subgenera within the former genus Psithyrus
have often been considered less distinct from one
another than have the other subgenera of Bombus
(Pittioni, 1939a; Ito, 1985; Williams, 19855; Michener,
1990) and therefore may be treated as synonyms of
Psithyrus (Milliron, 1961; Williams, 1991, 1995). In
an alternative treatment, Rasmont et al. (1995) include
the former subgenera of the former genus Psithyrus as
separate subgenera within the genus Bombus.

O NOMENCLATURE. The names of six species of
the subgenus Psithyrus from Kashmir were explicitly
stated to be new combinations with the genus Bombus
by Williams (1991). Rasmont ef al. (1995) have since
listed the other European species in this combination.
No formal statements of new combination are made
here for the remaining species of the subgenus
Psithyrus because a principle of implied combinations
(Poole, 1996) is followed after the change in status of
Psithyrus from genus to a subgenus of Bombus.

APPLICATION TO ICZN. Because the type spe-
cies of Laboriopsithyrus was misidentified (discussed
by Milliron, 1960:99), ICZN is required to designate
as type species whichever species will best serve
nomenclatural stability (ICZN, 1985: Art. 70b). It is
suggested that, in the interests of stability (ICZN,
1985: Article 23b), an application be made to ICZN to
use its Plenary Power to select the species actually
involved (Bombus laboriosus in the sense of Frison, =
Bombus citrinus (Smith)), which was wrongly named
in the type fixation (ICZN, 1985: Art. 70b(i)).

COMMENT. The highest richness of species of the
subgenus Psithyrus occurs in the Old World (there
are no species known from south of Panama), al-
though the earliest-diverging species appear to be
North American (unpublished). This is the opposite
pattern to that shown by species of the largest
subgenus, Pyrobombus (see the comments on the
subgenus Pyrobombus).

All species of the subgenus Psithyrus are believed
to be obligate social parasites in colonies of other
Bombus species (reviewed by Alford, 1975; Fisher,
1987). There is variation in the degree of host
specificity. See also the comments on B. inexspectatus
and B. hyperboreus.

103

Bombus (Ps.) insularis (Smith)

interruptus Greene, 1858:11, not of Lepeletier, 1832:381
(= B. rupestris (Fabricius))

insularis (Smith, 1861:155 [Apathus]) examined

consultus (Franklin, 1913:459 [Psithyrus])

? bicolor (Franklin, 1913:460 [Psithyrus]) not of H6ppner,
1897:33 (=B. soroeensis (Fabricius)) (provisional syno-
nym)

crawfordi (Franklin, 1913:464 [Psithyrus])

@ TAXONOMIC stTaTUS. According to D. Yanega
(in litt.), who has examined the type material, B.
bicolor Franklin is conspecific with B. interruptus.

Bombus (Ps.) citrinus (Smith)
citrinus (Smith, 1854:385 [Apathus]) examined
contiguus (Cresson, 1863:112 [Apathus])

Bombus (Ps.) variabilis (Cresson) ®

intrudens (Smith, 1861:154 [Apathus]) examined

variabilis (Cresson, 1872:284 [Apathus]) new synonym

?guatemalensis (Cockerell, 1912:21 [Psithyrus}) (provi-
sional synonym)

?sololensis (Franklin, 1915:173 [Psithyrus]) (provisional
synonym)

?mysticus (Frison, 1925a:138 [Psithyrus]) (provisional
synonym)

@ TAXONOMIC STATUS. Specimens in the NHM
collection from Mexico and Guatemala labelled
‘intrudens’ and ‘sololensis’ appear to me to be closely
similar to B. variabilis. Frison (1925a) believed that B.
sololensis is acolour form of B. guatemalensis. Never-
theless, he proceeded to distinguish B. mysticus as a
separate species on the basis of colour pattern alone. I
am unaware of any reason (other than minor differ-
ences in colour pattern) why B. variabilis, B. intrudens,
B. sololensis, or B. guatemalensis and B. mysticus
(judging from the published descriptions at least),
should not be considered conspecific.

O NOMENCLATURE. A female in the NHM collec-
tion has three labels “Apathus / intrudens / Smith.’,
*58.135 MEX. / (Oajaca.)’, “Holo- / type’ and I am
unaware of any problems with this designation. If this is
correct and the type is conspecific with B. variabilis,
then B. intrudens is the oldest available name for this
species. D. Yanega (in /itt.) agrees with this interpreta-
tion.

€ APPLICATION TOICZN. Although B. intrudens is
the oldest available name for the present interpretation
of this species, the name B. variabilis has been in
common use for the species since 1947 (e.g. Stevens,
1948; Chandler, 1950; LaBerge & Webb, 1962;
Mitchell, 1962; Medler & Carney, 1963; Hobbs, 1966;
Plowright & Stephen, 1973; Hurd, 1979; Husband et
al., 1980; Michener, 1990; Poole, 1996). I know of no
publications using the name B. intrudens since 1947. It

104

is suggested that, in the interests of stability (ICZN,
1985: Article 23b), an application be made to ICZN to
use its Plenary Power to suppress the unused senior
synonym (ICZN, 1985: Article 79) (see the comments
on B. muscorum). However, the consequence of this
action would be that B. intrudens would no longer be
available for a species or for a subspecies of B. variabilis
(Cresson).

Bombus (Ps.) suckleyi Greene
Suckleyi Greene, 1860:169

Bombus (Ps.) vestalis (Geoffroy)
veftalis (Geoffroy in Fourcroy, 1785[see Hagen
1862:246]:450 [Apis])

Bombus (Ps.) perezi (Schulthess-Rechberg)
perezi (Schulthess-Rechberg, 1886:275 [Psithyrus])

Bombus (Ps.) ashtoni (Cresson)
Ashtoni (Cresson, 1864:42 [Apathus])

Bombus (Ps.) bohemicus Seidl

nemorum (Fabricius, 1775:380 [Apis]) examined, not of
Scopoli, 1763:307 (= B. subterraneus (Linnaeus)), not
of Fabricius, 1775:382 (= B. distinguendus Morawitz)

bohemicus Seidl, 1837:73

?chinganicus (Reinig, 1936:8 [Psithyrus]) (provisional
synonym)

hedini (Bischoff, 1936:26 [Psithyrus]) not of Bischoff,
1936:15 (= B. hedini Bischoff)

@ TAXONOMIC sTATUS. Iam unaware of any rea-
son (other than the small body size of the holotype
female and three paratype females of B. chinganicus)
why B. bohemicus and B. chinganicus should not be
considered conspecific. Consistent with this, body
sizes do appear to vary considerably within British
species of the subgenus Psithyrus, including B.
bohemicus.

Bombus (Ps.) coreanus (Yasumatsu)
coreanus (Yasumatsu, 1934:399 [Psithyrus])

Bombus (Ps.) barbutellus (Kirby)
Barbutella (Kirby, 1802:343 [Apis]) examined
?richardsi (Popov, 1931:150,190 [Psithyrus]) not of Frison,
1930:6 (= B. rufipes Lepeletier)
2icenti (Maa, 1948:34 [Psithyrus]) examined

O NOMENCLATURE. Loken (1984) interpreted B.
saltuum (Panzer, 1801) as conspecific with B.
barbutellus. Consequently, B. saltuum would appear
to be the oldest available name for this species. How-
ever, Lgken made no further comment on this and used
the name Psithyrus barbutellus (= B. barbutellus),
possibly because she remained unsure of the identity
of B. saltuum. In contrast, Warncke (1986) interpreted

P.H. WILLIAMS

B. saltuum as conspecific with B. subterraneus. See
the comments on B. subterraneus.

?Bombus (Ps.) maxillosus Klug
maxillosus Klug in Germar, 1817:269
lugubris (Kriechbaumer, 1870:159 [Psithyrus])
unicolor (Kriechbaumer, 1870:159 [Psithyrus])
mixta (Kriechbaumer, 1870:160 [Psithyrus])
Susterai (May, 1944:267 [Psithyrus]) not infrasubspecific
after Tkalci, 1977:224

@ TAXONOMIC STATUS. As Rasmont (1988) notes,
B. maxillosus 1s closely similar to B. barbutellus in
morphology and habitat, so that specimens cannot
always be distinguished reliably. Consequently these
nominal taxa might be considered conspecific. More
evidence is awaited.

Bombus (Ps.) cornutus (Frison)
cornutus (Frison, 1933:338 [Psithyrus])
pyramideus (Maa, 1948:19 [Psithyrus]) examined
acutisquameus (Maa, 1948:21 [Psithyrus]) examined
Klapperichi (Pittioni, 1949:273 [Psithyrus]) examined, not
of Pittioni, 1949:266 (= B. picipes Richards)
?canus (Tkalcit, 1989:42 [Psithyrus])

Bombus (Ps.) expolitus Tkalcit
expolitus (Tkalct, 1989:44 [Psithyrus]) examined

Bombus (Ps.) turneri (Richards)
turneri (Richards, 1929a:141 [Psithyrus]) examined
?monozonus (Friese, 1931:304 [Psithyrus]) not of Friese,
1909:674 (= B. lucorum (Linnaeus))
?decoomani (Maa, 1948:26 [Psithyrus]) examined
?martensi (Tkalct, 1974b:314 [Psithyrus]) (provisional
synonym)

@ TAXONOMIC sTATUS. Several of these nominal
taxa have been treated as separate species. However,
aside from differences in colour pattern, they are
closely similar in morphology. Until more evidence to
the contrary is available from critical studies of pat-
terns of variation, I shall treat them as parts of a single
variable species.

Bombus (Ps.) tibetanus (Morawitz)
tibetanus (Morawitz, 1886:202 [Apathus])
?latefasciatus (Friese, 1931:304 [Psithyrus])

Bombus (Ps.) chinensis (Morawitz)
chinensis (Morawitz, 1890[April 30]:352 [Apathus])
morawitzi (Friese, 1905:516 [Psithyrus]) not of
Radoszkowski, 1876:101 (= B. morawitzi Rado-
szkowski)
hénei (Bischoff, 1936:26 [Psithyrus]) not of Bischoff,
1936:10 (= B. friseanus Skorikoy)

CHECKLIST OF BUMBLE BEES

Bombus (Ps.) novus (Frison)
novus (Frison, 1933:340 [Psithyrus])
?nepalensis (Tkalcti, 19746:318 [Psithyrus]) examined

Bombus (Ps.) branickii (Radoszkowski)
Branickii (Radoszkowski, 1893:241 [Psithyrus]) exam-
ined
chloronotus (Morawitz, 1894:6 [Apathus])
elisabethae (Reinig, 1940:231 [Psithyrus]) examined
[branichi (Kim & Ito, 1987:32 [Psithyrus]) incorrect sub-
sequent spelling]

Bombus (Ps.) rupestris (Fabricius)
rupeftris (Fabricius, 1793:320 [Apis])
Pyrencus (Lepeletier, 1832:375 [Psithyrus])
Interruptus (Lepeletier, 1832:381 [Psithyrus])
armeniacus (Reinig, 1970:77 [Psithyrus]) not of
Radoszkowski, 1877b:202 (= B. armeniacus
Radoszkowski)

Bombus (Ps.) ferganicus (Radoszkowski)
ferganicus (Radoszkowski, 1893:241 [Psithyrus]) exam-
ined
ochraceus (Morawitz, 1894:5 [Apathus])
indicus (Richards, 1929a:139) examined

Bombus (Ps.) morawitzianus (Popov)
morawitzianus (Popov, 1931:148,183 [Psithyrus]) exam-
ined
redikorzevi (Popov, 1931:160,181 [Psithyrus])

O NOMENCLATURE. Griitte (1937) regarded B.
morawitzianus and B. redikorzevi as conspecific and,
following the Principle of First Reviser (ICZN, 1985:
Article 24), chose B. morawitzianus as the name for
the species.

Bombus (Ps.) campestris (Panzer)

campestris (Panzer, 1801(74):11 [Apis])

Varius (Lepeletier, 1832:381 [Psithyrus])

flavus (Pérez, 1884:265 [Psithyrus])

flavo-thoracicus (Hoffer, 1889:49 [Psithyrus])

?Susterai (Tkalcti, 1959:251 [Psithyrus]) examined, not of
May, 1944:267 (= B. maxillosus Klug) (provisional
synonym)

?susteraianus (Tkalcu, 1977:224 [Psithyrus]) replacement
name for susterai Tkalcii, 1959:251 (provisional syno-
nym)

@ TAXONOMIC stTaTUS. Iam unaware of any rea-
son (other than minor differences) why B. campestris
and B. susteraianus should not be considered
conspecific.

105

Bombus (Ps.) bellardii (Gribodo)
Bellardti (Gribodo, 1892:108 [Psithyrus]) examined
pieli (Maa, 1948:29 [Psithyrus]) examined, new synonym
tajushanensis (Pittioni, 1949:277 [Psithyrus]) examined,
not of Pittioni, 1949:244 (= B. kulingensis Cockerell),
new synonym

@ TAXONOMIC sTATUS. B. bellardii, B. pieli and
B. tajushanensis are closely similar in morphology
and I am unaware of any reason why these nominal
taxa should not be considered conspecific.

O NOMENCLATURE. For this species, the oldest
available name is B. bellardii, which becomes the
valid name. The only subsequent publications using
the name B. pieli of which I am aware are by Maa
(1948), Sakagami (1972), Tkalciit (1987) and Williams
(1991), so this change of valid name is not a serious
disruption of common usage.

Bombus (Ps.) norvegicus (Sparre-Schneider)®
norvegicus (Sparre-Schneider, 1918:40 [Psithyrus]) not of
Friese, 1911:571 (= B. monticola Smith)
transhaicalicus (Popov, 1927:269 [Psithyrus])

O NOMENCLATURE. With Psithyrus regarded as
being a subgenus of the genus Bombus (Williams,
1991, 1995), P. norvegicus Sparre-Schneider (1918)
becomes a junior secondary homonym in Bombus of
B. lapponicus var. norvegicus Friese (1911) (deemed
subspecific, see ICZN, 1985: Article 45g(ii)), and
therefore the name P. norvegicus Sparre-Schneider is
invalid (ICZN, 1985: Article 57c). For this species, the
oldest available name of which I am aware is P.
norvegicus var. transbaicalicus Popov, 1927 (deemed
to be subspecific, see ICZN, 1985: Article 45g(ii)), so
B. transbaicalicus would become the valid name.

© APPLICATION TO ICZN. Although B. trans-
baicalicus is the oldest available name for this
species, the name B. norvegicus has been in com-
mon use for the species since 1947 (e.g. Faester &
Hammer, 1970; Delmas, 1976; Ito & Tadauchi, 1981;
Pekkarinen ef al., 1981; Reinig, 1981; Loken &
Framstad, 1983; Rasmont, 1983; Loken, 1984; Ito,
1985; Pekkarinen & Teras, 1993; Rasmont ef al.,
1995). It is suggested that, in the interests of stabil-
ity, an application be made to ICZN to use its Plenary
Power to suppress the senior homonym (ICZN, 1985:
Article 79) (see the comments on B. muscorum).
However, the consequence of this action would be
that norvegicus Friese would no longer be available
for a subspecies of B. monticola.

Bombus (Ps.) fernaldae (Franklin)
fernalde (Franklin, 1911:164 [Psithyrus]) examined

106

Bombus (Ps.) flavidus Eversmann
flavidus Eversmann, 1852:131
lissonurus (Thomson, 1872:49 [Apathus])

@ TAXONOMIC STATUS. Rasmont (1988) reports
that the Pyrenean population of B. flavidus is
morphometrically distinct from the disjunct Scan-
dinavian population (comparable distinctions are not
known within its close relatives B. norvegicus and B.
sylvestris, which share these areas of distribtuion).
Nevertheless he continues to treat them as conspecific
and I shall follow this, at least until further evidence in
support of two separate species is available.

Bombus (Ps.) skorikovi (Popov)
skorikovi (Popov, 1927:267 [Psithyrus]) examined
?gansuensis (Popov, 1931:202 [Psithyrus])
?kuant (Tkalcu, 1961b:362 [Psithyrus])

Bombus (Ps.) quadricolor (Lepeletier)
Quadricolor (Lepeletier, 1832:376 [Psithyrus])
globosus (Eversmann, 1852:126 [Psithyrus])
meridionalis (Richards, 1928b:351 [Psithyrus]) not of
Dalla Torre, 1879:13 (= B. hortorum (Linnaeus))

Bombus (Ps.) sylvestris (Lepeletier)

Sylvestris (Lepeletier, 1832:377 [Psithyrus])

Brasiliensis (Smith, 1854:385 [Apathus]) examined, not of
Lepeletier, 1836:470 (= B. brasiliensis Lepeletier)

citrinus (Schmiedeknecht, 1883[see Baker,
1996c:297]:23[407] [Psithyrus]) not of Smith, 1854:385
(= B. citrinus (Smith))

[silvestris (Dalla Torre, 1896:571 [Psithyrus]) incorrect
subsequent spelling]

e00e000ee0e er
Pats eecsece: *
ee se 00% oe one f

Subgenus LAESOBOMBUS Kriiger
Bombus (Laesobombus) Kriiger, 1920:350, type-species
Bombus laesus Morawitz by monotypy
Agrobombus (Laesobombus) Skorikov, 1922b:20, type-
species Bombus laesus Morawitz by monotypy
Agribombus (Laesibombus) Skorikoy, 1938a:145, unjusti-
fied emendation

P.H. WILLIAMS

Bombus (Ls.) laesus Morawitz
laesus Morawitz in Fedtschenko, 1875:3
Mocsaryi Kriechbaumer, 1877:253
?maculidorsis (Skorikoy, 1922b:23 [Agrobombus]) not
infrasubspecific after Panfilov, 1956:1328
?tianschanicus Panfilov, 1956:1327 (provisional synonym)
ferrugifer Reinig, 1971:158

@ TAXONOMIC STATUS. Panfilov (1956) regarded
B. laesus, B. mocsaryi, B. maculidorsis and B. tian-
schanicus as separate species, differing particularly in:
(1) the colour of the pubescence on the thoracic dor-
sum; (2) the number of large punctures on the clypeus;
(3) the strength of the median keel on gastral sternum
VI; and (4) the length of the hair of the dorsum.
However, from the material I have examined (collec-
tions in London, Beijing), these character states do not
appear to be either discreet or strongly associated.
Until more evidence to the contrary is available from
critical studies of patterns of variation, I shall treat
them as parts of a single variable species.

Subgenus ORIENTALIBOMBUS Richards
Bombus (Orientalibombus) Richards, 1929c:378, type-
species Bombus orientalis Smith (= Bombus
haemorrhoidalis Smith) by original designation
Bombus (Orientalobombus) Kruseman, 1952:102, unjus-
tified emendation

Bombus (Or.) funerarius Smith
funerarius Smith, 1852b:47, examined
priscus (Frison, 1935:349 [Bremus])
birmanus (Tkalct, 1989:47 [Orientalibombus]) examined

Bombus (Or.) braccatus Friese
braccatus Friese, 1905:512, examined
metcalfi (Frison, 1935:357 [Bremus]) examined

Bombus (Or.) haemorrhoidalis Smith
heemorrhoidalis Smith, 1852a:43
orientalis Smith, 1854:402, examined
assamensis Bingham, 1897:550, examined

CHECKLIST OF BUMBLE BEES

montivolans Richards, 1929c:382, examined
semialbopleuralis (Tkalcit, 1974b:322 [Orientalibombus])
cinnameus (Tkalcii, 1989:47 [Orientalibombus]) examined

@ TAXONOMIC sTATUS. Several of these nominal
taxa have been treated as separate species, most recently
in the case of B. montivolans [Burma to southern China]
(e.g. Tkalcii, 1968b, 1989). However, aside from differ-
ences in colour pattern, they are all closely similar in
morphology with a range of variation (Williams, 1991).
Until more evidence to the contrary is available from
critical studies of patterns of variation, I shall treat them
as parts of a single variable species.

Subgenus EXILOBOMBUS Skorikov
Mucidobombus (Exilobombus) Skorikov, 1922a:150, type-
species Mucidobombus exil Skorikoy (cited as exiln.) (=
Bombus exil (Skorikoy)) by monotypy
Megabombus (Exilnobombus) Milliron, 1973a:81, unjus-
tified emendation

Bombus (Ex.) exil (Skorikov)

exiln. nov. (Skorikovy, 1922a:150 |[Mucidobombus}) {not a
replacement name]

[exul (Skorikov, 1931:216 [Mucidobombus}) incorrect sub-
sequent spelling]

exil (Milliron, 1961:56 [Megabombus]) justified emenda-
tion

[exilis Richards, 1968:254, incorrect subsequent spelling]

exul (Tkalcti, 1974a:42 [Megabombus]) unjustified emen-
dation

Subgenus THORACOBOMBUS Dalla Torre
Bombus (Thoracobombus) Dalla Torre, 1880:40, type-

107

species Apis sylvarum Linnaeus (= Bombus sylvarum
(Linnaeus)) by subsequent designation of Sandhouse,
1943:604

Bombus (Chromobombus) Dalla Torre, 1880:40, type-spe-
cies Apis muscorum Linnaeus (= Bombus muscorum
(Linnaeus)) by subsequent designation of Sandhouse,
1943:538

Bombus (Agrobombus) Vogt, 1911:52, type-species Apis
agrorum Fabricius (=Bombus pascuorum (Scopoli)) by
subsequent designation of Sandhouse, 1943:523

[Agrabombus Skorikov, 1914a:119, incorrect subsequent
spelling]

Bombus (Ruderariobombus) Kriiger, 1920:350, type-spe-
cies Apis ruderaria Miiller (= Bombus ruderarius
(Miiller)) by subsequent designation of Yarrow, 197 1:27

Agrobombus (Adventoribombus) Skorikoy, 1922a:150,
type-speciesA grabombus adventor Skorikov (=Bombus
filchnerae Vogt) by subsequent designation of
Sandhouse, 1943:522, new synonym

[Agrobombus (Adventoriobombus) Skorikoy, 1931:218,
incorrect subsequent spelling]

Agribombus Skorikovy, 1938a:145, unjustified emendation

[Bombus (Thoraocbombus) Esmaili & Rastegar, 1974:52,
incorrect subsequent spelling]

[Bombus (Thoracibombus) Schwarz et al., 1996:197, in-
correct subsequent spelling]

@ TAXONOMIC STATUS. Richards (1968) treated
Thoracobombus and Adventoribombus as separate
subgenera, although he questioned whether they should
be kept separate. I have followed Tkalcti (1974a) in
treating B. adventor (=B. filchnerae) as part of a single
subgenus Thoracobombus.

Bombus (Th.) filchnerae Vogt
Filchnerae Vogt, 1908:100, examined
adventor (Skorikov, 1914a:119 [Agrabombus})
lii Tkalcit, 1961b:355

Bombus (Th.) muscorum (Linnaeus)

Mujcorum (Linnaeus, 1758:579 [Apis]) examined

pallidus Evans, 1901:47, not of Cresson, 1863:92 (= B.
pensylvanicus (DeGeer))

[fulvofasciatus Friese, 1905:520, infrasubspecific]

laevis Vogt, 1909:63

?nigripes Pérez, 1909:158, not of Haliday in Curtis ef al.,
1837:321 (= B. dahlbomii Guérin-Méneville)

?pereziellus (Skorikoy, 1922a:150 [Agrobombus]) replace-
ment name for nigripes Pérez, 1909:158

?bannitus (Skorikov in Popov, 1930:98 [Agrobombus])

?liepetterseni Loken, 1973:152

celticus Yarrow, 1978:15, replacement name for pallidus
Evans, 1901:47

@ TAXONOMIC STATUS. B. bannitus (= B. smith-
ianus of authors, a misidentification (=B. pascuorum))
has been regarded as a separate species by some
authors (e.g. Richards, 1935; Tkalct, 1987; Rasmont
& Adamski, 1995) on the basis of its semi-melanic
colour pattern and more coarsely sculptured surface of

108

gastral terga [V-V. However, Loken (1973: fig. 81)
found no difference between these taxa in a
morphometric study (other authors reporting no clear
morphological differences include Richards, 1935;
Alford, 1975; Pekkarinen, 1979; Rasmont, 1982;
Baker, 1996a). Furthermore, I have collected many
specimens with a range of intermediate colour patterns
on the Isle of Skye in western Scotland. Until more
evidence to the contrary is available from critical
studies of patterns of variation, I shall treat them as
parts of a single variable species.

B. pereziellus has also been regarded as a separate
species by Rasmont & Adamski (1995), because of its
dark colour pattern (even darker than B. bannitus, B.
pereziellus has the thoracic dorsum black rather than
red-brown, and has more black hairs on gastral tergum
II, whereas these black hairs tend to be more frequent
on tergum I for B. bannitus) and because it is endemic
to the island of Corsica. Morphologically it was con-
sidered by Rasmont (1982) to show no perceptible
differences from B. muscorum or B. bannitus. Further-
more, a male with a colour pattern apparently
intermediate between B. muscorum and B. pereziellus
is mentioned by Delmas (1976:271). Depending on
the species concept embraced, some differences might
be expected for a peripheral population such as this
even if it were conspecific and I shall treat them as
parts of a single variable species. Further evidence is
awaited.

O NOMENCLATURE. Richards (1935, 1968),
Yarrow (1968) and Léken (1973) recognised that none
of the admissable syntypes in the Linnean collection
agreed with the traditional interpretation of B.
muscorum, which is very rare in the parts of Sweden
where Linnaeus collected (Richards, 1935; Loken,
1973; Day, 1979), but took no action. When Day
(1979) came to fix the application of the name, he had
no reason to believe that Linnaeus had not described
his A. muscorum from the syntype specimen that was
subsequently described as lectotype (= B. humilis
Illiger).

To reaffirm the traditional usage of B. muscorum, a
case was made to ICZN by Loken et al. (1994). This
sought an Opinion from ICZN (ICZN, 1996) that set
aside by use of its Plenary Power (ICZN, 1985: Arti-
cles 78b, 79) the lectotype designation forA. muscorum
by Day from application of the Code (ICZN, 1985)
and then designated a neotype (ICZN, 1996: 64) to
conserve the traditional usage of the name for even the
narrowest concept of the taxon (ICZN, 1985: Article
75).

Bombus (Th.) anachoreta (Skorikov)
anachoreta (Skorikov, 1914a:121 [Agrobombus])

P.H. WILLIAMS

Bombus (Th.) opulentus Smith
opulentus Smith, 1861:153, examined

Bombus (Th.) zonatus Smith
zonatus Smith, 1854:389

Bombus (Th.) humilis MligerS
fulvefcens (Schrank, 1802:367 [Apis])
humilis Wliger, 1806:171
?tristis Seidl, 1837:69
?variabilis Schmiedeknecht, 1878:424, not of Cresson,
1872:284 (= B. variabilis (Cresson))
?subbaicalensis Vogt, 1911:42,54

OQ NOMENCLATURE. When Day (1979) came to
fix the application of A. muscorum Linnaeus (see the
comments on B. muscorum), he had no reason to
believe that Linnaeus had not described this taxon
from the syntype specimen that was subsequently
described as lectotype (= B. humilis Mlliger). This
action brought B. humilis Mliger into subjective junior
synonymy with B. muscorum (Linnaeus).

To reaffirm the traditional usage of B. muscorum
and B. humilis, a case was made to ICZN by Loken et
al. (1994). This sought an Opinion from ICZN (ICZN,
1996) that set aside by use of its Plenary Power (ICZN,
1985: Articles 78b, 79) the lectotype designation forA.
muscorum by Day from application of the Code (ICZN,
1985) and then designated a neotype (ICZN, 1996: 64)
to conserve the traditional usage of B. muscorum and
B. humilis ((CZN, 1985: Article 75).

However, Warncke (1986) recognised B. fulvescens
(Schrank) as questionably conspecific with B. humilis.
I have seen no type specimens, but the description is
consistent with this interpretation. B. fulvescens is
therefore likely to be the oldest available name for this
species.

€ APPLICATION TO ICZN. Although B. fulvescens
may be the oldest available name for the present
interpretation of this species, the name B. humilis has
been in common use for the species since 1947 (e.g.
case and references in Lgken et al., 1994). In contrast,
I know of no publications using the name B. fulvescens
(Schrank) since 1947. Warncke (1986:98) followed
the listing of this name with ‘Art. 23b’, which is a
reference to purpose of the Principle of Priority (ICZN,
1985). I agree that, in the interests of stability, an
application be made to ICZN to use its Plenary Power
to suppress the unused senior synonym (ICZN, 1985:
Article 79) (see the comments on B. muscorum).

Bombus (Th.) deuteronymus Schulz
senilis Smith, 1879:131, examined, not of Fabricius,
1775:382 (= B. pascuorum (Scopoli))
deuteronymus Schulz, 1906:267, replacement name for

CHECKLIST OF BUMBLE BEES

senilis Smith, 1879:131
velox (Skorikoy, 1914a:120 [Agrobombus}])
[superequester (Skorikov, 1914c:405 [Agrobombus])
infrasubspecific ]
superequester (Skorikov, 1925:116 [Agrobombus})
bureschi Pittioni, 1939b:1, examined

Bombus (Th.) schrencki Morawitz
Schrencki Morawitz, 1881:123
Schrencki Morawitz, 1881:250, redescribed
konakovi Panfiloy, 1956:1330

?Bombus (Th.) honshuensis (Tkalcii)
honshuensis (Tkalcti, 1968a:47 [Megabombus])

@ TAXONOMIC stTATUS. B. honshuensis and B.
schrencki have allopatric distributions in northern Ja-
pan (Sakagami & Ishikawa, 1969; Ito & Munakata,
1979: fig. 6; Ito, 1993), with B. honshuensis being
possibly a disjunct peripheral population of B.
schrencki. The two taxa are closely similar, and yet
despite some variation in morphology, apparently con-
sistent differences have been described (Tkalcii, 1968a;
Sakagami & Ishikawa, 1972). Nonetheless, some dif-
ferences might be expected even if they were
conspecific, depending on the species concept ac-
cepted (see the comments on B. ruderatus), so further
evidence is awaited.

Bombus (Th.) impetuosus Smith
impetuosus Smith, 1871:249, examined
Potanini Morawitz, 1890:350, new synonym
yuennanensis Bischoff, 1936:14, examined
combai Tkalcti, 1961b:357, new synonym

@ TAXONOMIC STATUS. The white-banded B.
potanini is morphologically closely similar to the
yellow-banded B. impetuosus. Some individuals from
Sichuan are intermediate in colour pattern in that they
have the pale bands of the thorax and gastral tergum I
white, and the pale band of tergum II yellow. There is
considerable variation in the male gonostylus, but this
variation appears to overlap between the the colour
forms and I shall treat them as parts of a single variable
species. S.-f. Wang and J. Yao (in litt.) also believe that
the two taxa may be conspecific. Further evidence is
awaited.

Bombus (Th.) remotus (Tkalcii)
remotus (Tkalcti, 1968a:45 [Megabombus]) examined

Bombus (Th.) pseudobaicalensis Vogt
Pseudobaicalensis Vogt, 1911:43,53
gilvus (Skorikov, 1925:117 [Agrobombus})

109
Bombus (Th.) hedini Bischoff

unicolor Friese, 1905:514, examined, not of Kriechbaumer,
1870:159 (= B. maxillosus Klug)
hedini Bischoff, 1936:15

Bombus (Th.) ruderarius (Miiller)
ruderaria (Miller, 1776:165 [Apis])
Derhamella (Kirby, 1802:363 [Apis]) examined
montanus Lepeletier, 1836:463
simulatilis Radoszkowski, 1888:317, examined

Bombus (Th.) inexspectatus Tkalcit
lutescens Kriiger, 1939:105, not of Pérez, 1890:154 (= B.
flavidus Eversmann)
inexspectatus (Tkalct, 1963:187 [Agrobombus})
[inexpectatus (Reinig, 1981:161 [Megabombus}]) incorrect
subsequent spelling]

COMMENT. On the grounds of its peculiar morphol-
ogy, B. inexspectatus has been suggested to be an
obligate social parasite in colonies of other Bombus
species, with B. ruderarius being the most likely host
(Yarrow, 1970). As yet, there is no direct evidence for
this behaviour (Rasmont, 1988). See the comments on
the subgenus Psithyrus and on B. hyperboreus.

Bombus (Th.) sylvarum (Linnaeus)
/ylvarum (Linnaeus, 1761:425 [Apis]) examined
Daghestanicus Radoszkowski, 1877a:vii
Dagestanicus Radoszkowski, 1877b:211, redescribed

Bombus (Th.) veteranus (Fabricius)
veterana (Fabricius, 1793:324 [Apis])
arenicola Thomson, 1872:31

Bombus (Th.) mlokosievitzii Radoszkowski

Mlokosievitzii Radoszkowski, 1877a:viii

Mlokassewiczi Radoszkowski, 1877b:212, redescribed

pérezi Vogt, 1911:55, not of Schulthess-Rechberg,
1886:275 (= B. perezi (Schulthess-Rechberg))

vogtiellus (Tkalcu, 1977:224 [Megabombus]) replacement
name for perezi Vogt, 1911:55

[mlokossowiczi (Reinig, 1981:161 [Megabombus]) incor-
rect subsequent spelling]

O NOMENCLATURE. There are particularly many
incorrect subsequent spellings of B. mlokosievitzii.

Bombus (Th.) pascuorum (Scopoli)
Pafcuorum (Scopoli, 1763:306 [Apis])
Jenilis (Fabricius, 1775:382 [Apis])
agrorum (Fabricius, 1787:301 [Apis]) not of Schrank,
1781:397 (= B. mesomelas Gerstaecker)
thoracicus Spinola, 1806:30

110

arcticus Dahlbom, 1832:50, not of Quenzel in Acerbi,
1802:253 (= B. hyperboreus Schonherr)

cognatus Stephens, 1846:17, examined

smithianus White, 1851:158

@ TAXONOMIC STATUS. Warncke (1986) listed B.
cognatus as a synonym of B. muscorum, possibly
following Stephens (1846), who wrote of B. cognatus:
‘Closely allied to Bo. Muscorum, of which the exam-
ples I possess may be immature specimens’. Pagliano
(1995) listed B. cognatus as a species separate from
both B. muscorum and B. pascuorum, but without any
explanation.

Saunders (1896:366—367) wrote ‘I have re-exam-
ined the type of cognatus, Steph., . . . F. Smith placed
it in the British Museum collection. ... Saunders
considered this specimen to be conspecific with B.
agrorum (Fabricius), continuing: ‘It is certainly not
the species known on the Continent as cognatus’.

A female in the NHM collection bears the following
labels: (1) a red-edged printed ‘Type’; (2) ‘cognatus.’
in handwriting identical to that of F. Smith; (3) ‘=
agrorum / I.H.H.Y.’ in handwriting identical to that of
I. Yarrow; (4) ‘B.M. Type / HYM. / 17B.1163’. I have
examined this specimen and am unaware of any reason
why it should not be considered the type of B. cogna-
tus and (ignoring minor differences in colour pattern)
conspecific with B. pascuorum.

OQ NOMENCLATURE. L@ken (1973) listed B. cogna-
tus Stephens, 1846, as anomen nudum, citing Sherborn
(1925). However, the reference by Sherborn is to an
earlier paper by Stephens (1829), so this does not affect
the use of the name B. cognatus Stephens, 1846.

Subgenus TRICORNIBOMBUS Skorikov
Agrobombus (Tricornibombus ) Skorikoy, 1922a:151, type-
species Bombus tricornis Radoszkowski by monotypy
Bombus (Tricornibombus) Tkalcti, 1960:70

Bombus (Tr.) tricornis Radoszkowski
tricornis Radoszkowski, 1888:319, examined

Bombus (Tr.) atripes Smith
atripes Smith, 1852a:44, examined

P.H. WILLIAMS

Bombus (Tr.) imitator Pittioni
imitator Pittioni, 1949:251, examined
flavescens Pittioni, 1949:254, not of Smith, 1852a:45 (=B.
flavescens Smith)

Subgenus FERVIDOBOMBUS Skorikov

Fervidobombus Skorikov, 1922a:153, type-species Apis
fervida Fabricius (= Bombus fervidus (Fabricius)) by
subsequent designation of Frison, 1927:69

Bombus (Fervidobombus) Franklin, 1954:47

Bombus (Digressobombus ) Laverty etal., 1984:1051, type-
species Megabombus digressus Milliron (= Bombus
digressus (Milliron)) by original designation

@ TAXONOMIC STATUS. The subgenus Digresso-
bombus was described subsequent to Richards (1968).
I have treated Digressobombus as part of a single
subgenus Fervidobombus (Williams, 1995), as has
Labougle (1990). While this study found no evidence
for monophyly of the combined group, I believe that
this is more likely (unpublished data) than monophyly
of the subgenus Fervidobombus excluding Digresso-
bombus.

COMMENT. This is the only early-diverging and large
subgenus of bumble bees to occur in the New World
other than the subgenus Psithyrus. Although it makes
up only a small part of the fauna of America north of
Mexico, it makes up most of the low- to mid-altitude
bumblebee fauna of Central and SouthAmerica. It also
includes the only species of bumble bees genuinely
occurring in tropical lowland wet forest (e.g. Moure &
Sakagami, 1962; Milliron, 1973a; Cameron &
Whitfield, 1996). The species with more temperate
distributions appear to occupy similar habitats and
show similar flower-depth preferences to species of
subgenera such as Thoracobombus and Megabombus
in the Old World.

Bombus (Fy.) fervidus (Fabricius)
feruida (Fabricius, 1798:274 [Apis])
?Californicus Smith, 1854:400, examined
Dumoucheli Radoszkowski, 1884:78
sonome Howard, 1902:pl. II

@ TAXONOMIC STATUS. B. fervidus and B.

CHECKLIST OF BUMBLE BEES

californicus have been regarded both as conspecific
(e.g. Milliron, 1973a; Labougle, 1990) and as separate
species (e.g. Franklin, 1913; Stephen, 1957; Thorp et
al., 1983: Poole, 1996). Both Franklin (1913:239) and
Stephen (1957) also considered the possibility that
they are conspecific as quite reasonable.

Many specimens from the north west of North
America show a reduction in the extent of the yellow
bands on the scutellum and gastral terga I-III and
appear to be intermediate or recombinant individuals.
Indeed, Stephen’s (1957:32) figure 2 shows several
patterns that could represent a continuum in variation
between the two forms. Thorp ef al. (1983) found no
intermediate females in California, although some of
the males of B. californicus were said to approach the
pattern of B. fervidus.

In view of the existence of apparent intermediates
between these nominal taxa in at least part of their
range, they are treated here as likely to be conspecific.
More evidence is awaited.

OQ NOMENCLATURE. Apis feruida is the original
spelling in Fabricius (1798). The orthography of this
publication employs ‘u’ in place of “v’ widely, a com-
mon practice of the period. This convention has since
changed and subsequent authors have consistently
used ‘vy’ for B. fervidus.

In fact, whatever the interpretation of the Code,
pragmatically it matters little which spelling offervidus
is used unless either of the spellings were to be pub-
lished as the name of another taxon in Bombus. See the
comments on the spelling of B. pensylvanicus.

Bombus (Fy.) pensylvanicus (DeGeer)
penfylvanica (DeGeer, 1773:575 [Apis])
americanorum Fabricius, 1804:346
?sonorus Say, 1837:413
pallidus Cresson, 1863:92
Pensylvanicus Cresson, 1863:94
flavodorsalis Franklin, 1913:409
pennsylvanicus Hurd, 1979:2204, unjustified emendation

@ TAXONOMIC sTATUS. B. pensylvanicus and B.
sonorus have been regarded both as conspecific (e.g.
Milliron, 1973a; Labougle ef al., 1985; Labougle,
1990; Poole, 1996) and as separate species (e.g.
Franklin, 1913 [but see p. 239]; Stephen, 1957; Thorp
et al., 1983; S. Cameron in litt.).

From the few males from the United States (not
Mexico) that I have examined in detail, there appear to
be subtle differences in the male genitalia (e.g. in the
shape of the penis valve head). However, Labougle
(1990) reports that the two ‘forms are geographically
intermixed in México, and chromatically intermediate
specimens occur, mainly in northeastern México and
southwestern Texas’. He went on to say that ‘In fact, it
is sometimes difficult to place a Mexican specimen in
either subspecies because there are specimens with the

111

coloration of the scutellum and the punctation of the
clypeus intermediate between the two taxa. Average
differences of certain proportions are found . . . but do
not differentiate all specimens’. G. Chavarria (pers.
com.) also believes that intermediate specimens occur
in Mexico and that they are conspecific. Taking an
extreme viewpoint, it is even possible to see the ‘typi-
cal’ B. sonorus colour pattern as intermediate between
B. pensylvanicus (in the strict sense) and the extreme
pale form that has the thoracic dorsum and gastral
tergum I entirely yellow (flavodorsalis, see Thorp et
al., 1983: fig. 137b).

In view of the existence of apparent intermediates
between these nominal taxa in at least part of their
range, they are treated here as likely to be conspecific.
More evidence is awaited.

O NOMENCLATURE. Apis penfylvanica is the origi-
nal spelling in DeGeer (1773). The orthography of this
publication employs ‘Pin place of ‘s’ widely, a com-
mon practice of the period. This convention has since
changed and subsequent authors (e.g. Cresson, 1863)
have consistently used ‘s’ for B. pensylvanicus.

Technically, according to the Code (ICZN, 1985:
Article 32b), pensylvanicus with just two ‘n’s is the
correct original spelling, to be preserved unaltered
unless it is demonstrably incorrect under Article 32c.
Article 32c(ii) states that clear evidence of an inadvert-
ent error is only admissable if it lies within the original
publication, without recourse to any external source of
information (DeGeer, 1773, spelled Penjylvanie and
penfylvanica consistently in this way). Any intentional
change to that spelling in a subsequent publication is
an unjustified emendation under Article 33b(iii).

In fact, whatever the interpretation of the Code,
pragmatically it matters litthe which spelling of
pensylvanicus is used unless either of the spellings
were to be published as the name for another taxon in
Bombus. No doubt many will prefer to use B.
pennsylvanicus, although the name does appear as B.
pensylvanicus in the recent checklist by Poole (1996)
(and by analogy, the similar spelling of Vespula
pensylvanica (Saussure) has been accepted, e.g. by
Akre et al., 1980; Edwards, 1980).

COMMENT. This species was deliberately introduced
into the Philippines, but is not known to have persisted
(Frison, 1925b).

Bombus (Fy.) excellens Smith
excellens Smith, 1879:133, examined

Bombus (Fy.) dahlbomii Guérin-Méneville
Dahlbomii Guérin-Méneville, [1835, see Cowan,
NO TE2 epi
nigripes Haliday in Curtis et al., 1836:321

112

O NOMENCLATURE. Cowan (1971), considering
Guérin-Méneville’s insect volume, states that ‘it is
quite certain that valid publication [of the Insectes
text] under the International Code of Nomenclature
did not take place until August or September 1844.’
However, he lists plate 75, on which B. dahlbomii
appears as figure 3 together with a legend containing
the name, as having been published in livraison 39 in
June 1835. This meets the criteria for valid publication
(ICZN, 1985: Article 8). Therefore B. dahlbomii ts the
oldest available name for this species.

Bombus (Fyv.) morio (Swederus)

morio (Swederus, 1787:283 [Apis]) examined

velutinus Mliger, 1806:175

violaceus Lepeletier, 1836:473

carbonarius Handlirsch, 1888:241, not of Menge, 1856:27
[fossil]

Kohli Cockerell, 1906:75, replacement name for
carbonarius Handlirsch, 1888:241

Bombus (Fy.) diligens Smith
diligens Smith, 1861:154, examined
dolichocephalus Handlirsch, 1888:244

Bombus (Fy.) opifex Smith
opifex Smith, 1879:133, examined

Bombus (Fy.) rubriventris Lepeletier
rubriventris Lepeletier, 1836:472, examined

@ TAXONOMIC sTATUS. B. rubriventris is known
from a single female specimen from ‘St. Domingue’
(2= Sao Domingos, Goids) (Milliron, 1973a). This
specimen has dark brown wings and the pubescence is
black, except that most of the hairs of the thorax are
grey-tipped, and the hairs of gastral terga II-IV are
bright ‘coppery’ red.

This colour pattern resembles the Andean B.
excellens, although the pubescence of B. rubriventris
is much shorter and more even; the oculo-malar area is
nearly square rather than nearly twice as long as the
basal breadth of mandible; and tergum VI is raised
subapically. Franklin (1913) had not seen B.
rubriventris but suggested that it was probably a ‘freak
specimen’ of B. carolinus (a misidentification, = B.
excellens). Milliron (1973a) had examined B.
rubriventris and considered the morphological char-
acters to be very much like those of B. bellicosus.
However, B. rubriventris can be distinguished by the
much finer punctures in the centre of the clypeus and
by an absence of a median ridge on tergum VI. I
consider B. rubriventris to be more similar in these
characters to B. opifex, although it can be distin-

P.H. WILLIAMS

guished from that species by a pair of characteristi-
cally slightly recessed bands of fine punctures
extending anteriorly from the ocello-ocular areas and
by a shallow median groove in the subapically raised
area of tergum VI.

The colour pattern is very distinctive among non-
Andean bumble bees in South America and does not
appear to be the result of abnormal colour develop-
ment. The specimen has had the gaster glued back into
place, although the characters of both the head and
gaster appear to be distinctive, so there is no reason to
believe that the specimen is a composite and not
genuine.

COMMENT. Milliron (1973a) researched the history
of this specimen and believed that it may have been
collected as early as 1800. He concluded that it was
probably a highland species from southeastern Brazil
and that it may now be extinct. If so, and accepting that
it is very difficult to establish the absence of a species,
this would be one of the few known cases of complete
extinction of an insect species.

Bombus (Fyv.) bellicosus Smith
thoracicus Sichel, 1862:121, not of Spinola, 1806:30 (=B.
pascuorum (Scopoli))
bellicosus Smith, 1879:131, examined
Emiliae Dalla Torre, 1890:139, replacement name for
thoracicus Sichel, 1862:121

Bombus (Fy.) pullatus Franklin
pullatus Franklin, 1913:122

Bombus (Fy.) transversalis (Olivier)
tran/verfalis (Olivier, 1789:65 [Apis])
Cajennenjis (Fabricius, 1798:273 [Apis])
incarum Franklin, 1913:131

Bombus (Fy.) atratus Franklin®
azurea (Christ, 1791:129 [Apis])
?atratus Franklin, 1913:118, not of Friese, 1911:572 (=B.
mucidus Gerstaecker) (provisional synonym)
?niger Franklin, 1913:120, examined (provisional syno-
nym)
?nigriventris Friese, 1913:87 (provisional synonym)

@ TAXONOMIC stTaTUS. Atleast four species of the
subgenus Fervidobombus from Central and South
America have many individuals for which the pubes-
cence is almost entirely black. The genitalia of the
males are quite distinctive, but association of the
conspecific females with these males has caused prob-
lems.

In the original description of B. niger, Franklin
stated that ‘atratus is possibly the male of niger (p.

CHECKLIST OF BUMBLE BEES

121), whereas in the original description of B. atratus
he stated both that ‘Niger may represent the females of
this species’ (p. 118) and that “This may be the true
male of kohli’ (p. 119). B. niger was described from a
syntype series of four queens and four workers, of
which one queen in the Smithsonian collection carries,
amongst others, a red label ‘LECTOTYPE / Bombus /
niger Franklin/H.E.Milliron ‘59’ and a label “Boquete
/ Chiriqui’. This Central American locality was men-
tioned by Franklin, but is outside the known distribution
of the species (Milliron, 1973a) to which the specimen
belongs. In my opinion, this lectotype of B. niger is not
conspecific with B. pullatus (contrary to the sugges-
tion by Labougle, 1990, see also Milliron, 1962) but
rather is conspecific with B. atratus Franklin.

Another possibility is that this variable species is the
Apis azurea of Christ (1791). I know of no type
specimens and the type locality was said to be inAfrica
(Ist in Afrika am Vorgebiirg der guten Hofnung zu
Haus’). The description and figure of the colour pat-
tern do not agree with any African bees that Ihave been
able to trace, but do resemble closely the yellow-
banded individuals of the SouthAmerican B. niger, the
Central American B. medius Cresson, and the South
American B. transversalis (Olivier) (although for the
last named species the yellow bands on the thorax are
usually broader). Among the specimens to hand, the
wings do appear slightly more “Schwarzblaue’ for B.
niger, as described for A. azurea, although these
grounds seem slim justification from which to estab-
lish the application of a name.

O NOMENCLATURE. B. azureus is possibly the old-
est available name for this species.

Milliron (1962), without mention of the name B.
azureus, first regarded B. atratus and B. niger as
conspecific and, following the Principle of First Re-
viser (ICZN, 1985: Article 24), chose B. atratus as the
valid name for the species.

Unfortunately, B. atratus Franklin, 1913, is a junior
primary homonym of B. mucidus var. atratus Friese,
1911 (deemed to be subspecific, see ICZN, 1985:
Article 45g(ii)), therefore the name B. atratus Franklin
is invalid (ICZN, 1985: Article 57b).

© APPLICATION TOICZN. The name B. azureus has
not been used since the original publication. The name
B. atratus has been used for this species since 1947
(e.g. Moure & Sakagami, 1962; Sakagami & Zucchi,
1965; Sakagami ef al., 1967; Milliron, 1971, 1973a;
Sakagami, 1976; Ito, 1985; Labougle, 1990; Varela,
1992; Silveira & Cure, 1993). It is suggested that, in
the interests of stability (ICZN, 1985: Article 23b), an
application be made to ICZN to use its Plenary Power
to suppress both the unused senior synonym (ICZN,
1985: Article 79) and the senior homonym. This would
achieve both an unambiguous, valid name for this
species (see the comments on B. muscorum) and also

113

help to protect the validity of the names B. medius and
B. transversalis from future change. However, the
consequence of this action would be that atratus Friese
would no longer be available for a subspecies of B.
mucidus.

Bombus (Fv.) digressus (Milliron)
digressus (Milliron, 1962:730 [Megabombus]) examined

Bombus (Fy.) brasiliensis Lepeletier
brasiliensis Lepeletier, 1836:470, examined

Bombus (Fy.) steindachneri Handlirsch
Steindachneri Handlirsch, 1888:239

@ TAXONOMIC sTATUS. B. medius andB. steindac-
hneri have been regarded both as separate species
(Milliron, 1973a; Labougle, 1990) and as conspecific
(G. Chavarria, pers. com.).

Labougle (1990) reports that ‘Although the chro-
matic differences between B. medius and B.
steindachneri are conspicuous, the male genitalia are
extremely similar’. Labougle listed four character dif-
ferences from the male genitalia and I can confirm two
of these: (1) that the head of the penis valve of B.
steindachneri has fewer fine teeth or serrations; and
(2) that the interior process of the volsella (misinter-
preted as the preapical tooth of the “gonostylus’; for
discussion of homologies see Williams, 1991) of B.
steindacheri is narrower. However, I have examined
only a few males and these characters might be ex-
pected to vary among other individuals. Labougle
(1990) continued: ‘The lack of chromatic and morpho-
logical intermediates supports the idea of two different
species’.

Until more evidence to the contrary is available
from critical studies of patterns of variation, I shall
treat them as two separate species.

Bombus (Fy.) medius Cresson
medius Cresson, 1863:97

Bombus (Fy.) weisi Friese
laboriosus Smith, 1861:153, examined, not of Fabricius,
1804:352 (= Emphoropsis laboriosus (Fabricius))
weisi Friese, 1903:253, examined
nigrodorsalis Franklin, 1907:90

O NOMENCLATURE. The lectotype female of B.
weisi by designation of Milliron (1960:98) was recog-
nised as conspecific with B. nigrodorsalis by Labougle
(1990) (Ihave examined the lectotype of B. weisi at the
MNHU, Berlin, and agree with Labougle). He then

114

used B. weisi (the oldest available name) as the valid
name for this species. However, a case could be made
in favour of the use of either name.

For Labougle’s (1990) use of this previously unused
senior synonym to be considered by ICZN as a prima
facie case of upsetting the use of a long-accepted name
in its accustomed meaning (ICZN, 1985: Article 23b),
the name B. weisi should not have been used in this
sense in the preceding fifty years; and at least five
authors should have used the junior name, B.
nigrodorsalis, in at least ten publications during the
same period (ICZN, 1985: Article 79c). As far as 1am
aware, no other admissable publications have used B.
weisi (Williams, 1995, disclaimed any nomenclatural
action in a list of names for material examined), al-
though publications using the junior name B.
nigrodorsalis Franklin for this species since 1947 are
more common, including Milliron (1961, 1962, 1971,
1973a), Laverty et al. (1984), Labougle et al. (1985),
Williams (19855) andAsperen de Boer (1992b). Other
such references may exist, therefore this may be seen
as a borderline case, requiring an application to be
made to ICZN to use its Plenary Power if suppression
of the unused senior synonym, B. weisi, is required
(see the comments on B. muscorum).

On the other hand, a change of valid name from B.
nigrodorsalis to B. weisi does not appear to be a
serious disruption of common usage, so there is no
obvious need for action to retain B. nigrodorsalis and
I have continued to use B. weisi.

Bombus (Fy.) trinominatus Dalla Torre
modestus Smith, 1861:153, examined, not of Eversmann,
1852:134 (= B. modestus Eversmann)
trinominatus DallaTorre, 1890:139, replacement name for
modestus Smith, 1861:153
xelajuensis Asperen de Boer, 1992b:162, examined (pro-
visional synonym)

@ TAXONOMIC STATUS. The description of B.
xelajuensis shows that this nominal taxon, known
from a single location, diverges only slightly in col-
our pattern and morphology from the otherwise
restricted and uncommon mountain species B.
trinominatus. Therefore it seems most likely to be
conspecific with B. trinominatus, with a slightly dif-
ferent colour pattern. However, the information
available at present is not conclusive, and it remains
possible that it represents a separate species, and
further evidence is awaited.

Bombus (Fy.) mexicanus Cresson
mexicanus Cresson, 1878:187

P.H. WILLIAMS

Bombus (Fyv.) brevivillus Franklin
brevivillus Franklin, 1913:119
?abditus (Tkalcit, 1966:271 [Megabombus}))

@ TAXONOMIC STATUS. The single known female
of B. abditus was described as originating from “Rep.
de Guinée Beyla’ (equatorial Africa). However, it is
indistinguishable from B. brevivillus according to
Sakagami (1976:427) and probably represents an in-
troduced or mislabelled individual (Michener, 1979).

Subgenus SENEXIBOMBUS Frison
Bremus (Senexibombus) Frison, 1930:3, type-species
Bombus senex Vollenhoven by original designation
[Bombus (Senecibombus) Kruseman, 1952:101 incorrect
subsequent spelling]
Bombus (Senexibombus) Richards, 1968:217

Bombus (Sx.) kulingensis Cockerell
kulingensis Cockerell, 1917:266
tajushanensis Pittion1, 1949:244

Bombus (Sx.) bicoloratus Smith
bicoloratus Smith, 1879:132, examined

Bombus (Sx.) senex Vollenhoven
Senex Vollenhoven, 1873:229

Bombus (Sx.) irisanensis Cockerell
irisanensis Cockerell, 1910a:416, examined

CHECKLIST OF BUMBLE BEES

Subgenus DIVERSOBOMBUS Skorikov
Bombus (Diversobombus ) Skorikov, 1914c:406, type-spe-
cies Bombus diversus Smith by subsequent designation
of Sandhouse, 1943:546
Diversibombus Skorikov, 1938b:2, unjustified emenda-
tion

Bombus (Dy.) trifasciatus Smith
trifasciatus Smith, 1852a:43, examined
montivagus Smith, 1878:168, examined
montivagus Smith, 1879:131, redescribed
?wilemani Cockerell, 1911:100, examined
albopleuralis Friese, 1916:108, examined
?maxwelli Pendlebury, 1923:67, examined
mimeticus Richards, 1931b:529, examined
malaisei (Skorikov, 1938b:2 [Diversibombus]) not of
Bischoff, 1930:4 (= B. sporadicus Nylander)
atropygus (Tkalcti, 1989:58 |Megabombus]}) examined

@ TAXONOMIC sTaTUS. Several of these nominal
taxa have been treated as separate species, for example
as B. albopleuralis (= B. mimeticus) {Himalaya}, B.
montivagus {northern Burma to southern China], B.
maxwelli [Peninsular Malaysia] and B. wilemani [Tai-
wan] (Tkalcii, 1968b, 1989). However, aside from
differences in colour pattern (Fig. 13), they are closely
similar in morphology and show a range of variation
(Williams, 1991). Until more evidence to the contrary
is available from critical studies of patterns of varia-
tion, I shall continue to treat them as parts of a single
variable species.

Bombus (Dyv.) longipes Friese
longipes Friese, 1905:511
hummeli Bischoff, 1936:18, examined

Bombus (Dy.) diversus Smith
diversus Smith, 1869:207, examined
tersatus Smith, 1869:207, examined

OQ NOMENCLATURE. Tkalcti (1965) first explicitly
regarded B. diversus and B. tersatus as conspecific
and, following the Principle of First Reviser (ICZN,
1985: Article 24), chose B. diversus as the valid name
for the species.

115

Bombus (Dv.) ussurensis Radoszkowski
Ussurensis Radoszkowski, 1877b:196
{ussuriensis Morawitz, 1881:254, incorrect subsequent
spelling]

Subgenus MEGABOMBUS Dalla Torre
Bombus (Megabombus) Dalla Torre, 1880:40, type-spe-
cies Bombus ligusticus Spinola (= Bombus argillaceus
(Scopoli)) by monotypy
Bombus (Megalobombus) Schulz, 1906:267, unjustified
emendation

Bombus (Hortobombus) Vogt, 1911:56, type-species Apis
hortorum Linnaeus (= Bombus hortorum (Linnaeus))
by subsequent designation of Sandhouse, 1943:559

Bombus (Odontobombus) Kriiger, 1917:61,65 (proposed
as a section name but stated by Milliron, 1961:53, to be
equivalent to his concept of the subgenus Megabombus
Dalla Torre), type-species Apis argillacea Scopoli (=
Bombus argillaceus (Scopoli)) by subsequent designa-
tion of Williams, 1995:339

[Nortobombus Skorikoy, 1922b:map 3, incorrect subse-
quent spelling]

Hortibombus Skorikov, 1938a:146, unjustified emenda-
tion

Bombus (Mg.) supremus Morawitz
supremus Morawitz, 1886:196
linguarius Morawitz, 1890:351

Bombus (Mg.) gerstaeckeri Morawitz
Gerstdckeri Morawitz, 1881:242
Gerstaeckeri Hoffer, 1883:55, mandatory correction
(ICZN, 1985: Article 32d)

Bombus (Mg.) consobrinus Dahlbom
consobrinus Dahlbom, 1832:49

Bombus (Mg.) tichenkoi (Skorikoy)
[tichenkoi (Skorikoy, 1922a:156 [Hortobombus]) published
without description]
tichenkoi (Skorikov, 1925:115 [Hortobombus])

116

?yezoensis Matsumura, 1932:pl. 1
?przewalskiellus (Skorikov, 1933a:59 [Hortobombus])
kurilensis Sakagami, 1954:92

@ TAXONOMIC STATUS. B. tichenkoi and B.
yezoensis have apparently been regarded both as
conspecific and as separate species.

Sakagami (1954) described kurilensis as a (semi-
melanic) subspecies of what he called ‘B. tersatus’ (a
misidentification, = B. diversus). Following Tkalcit
(1962), Sakagami subsequently identified his ‘B.
tersatus’ as B. yezoensis (Ito & Sakagami, 1980). In
this later paper he went on to recognise kurilensis as
conspecific with B. tichenkoi, but then, despite having
regarded kurilensis as a subspecies of what he now
believed to be B. yezoensis in the earlier paper, listed
B. tichenkoi as a species separate from B. yezoensis.
The apparent contradiction was not explained, al-
though morphological comparisons between these taxa,
B. argillaceus and B. sushkini were tabulated (which
show primarily that B. argillaceus 1s very different).
He even noted the allopatric distributions of B. tichenkoi
and B. yezoensis between the northern and southern
Kurile Islands and the ‘resemblance of tichenkoi and
dark individuals of yezoensis, especially in workers.’

Until more evidence to the contrary is available
from critical studies of patterns of variation, I shall
treat B. tichenkoi and B. yezoensis as parts of a single
variable species.

Bombus (Mg.) sushkini (Skorikov)
[saltuarius (Skorikov, 1922a:156 [Hortobombus]) pub-
lished without description]
sushkini (Skorikoy, 1931:235 [Hortobombus]) examined
saltuarius (Skorikov, 1931:235 |Hortobombus])

@ TAXONOMIC STATUS. B. sushkini and B.
saltuarius have been regarded both as conspecific
(Bischoff, 1936) and as separate species (Skorikov,
1931; Tkalct, 1974a). I have as yet seen no evidence
that more than one species is involved. More evidence
is awaited.

O NOMENCLATURE. Skorikov (1931) provided the
first valid publication of the names B. sushkini and B.
saltuarius as two separate species. Subsequently,
Bischoff (1936) regarded the two as conspecific and,
following the Principle of First Reviser (ICZN, 1985:
Article 24), chose B. sushkini as the valid name for the
species.

Bombus (Mg.) portchinsky Radoszkowski
Portchinskij Radoszkowski, 1883:208
Portchinsky Radoszkowski, 1883:208[210], 209[211],
210[212]

O NOMENCLATURE. Radoszkowski (1883) pub-
lished two different spellings of B. portchinsky,

P.H. WILLIAMS

repeating this second form several times (page num-
bers 207 and 208 are repeated twice for different
pages, so the two spellings do not occur on the same
page). Precedence of the correct original spelling
should be determined using the Principle of the First
Reviser (ICZN, 1985: Article 24), but to date I have not
found an author who has correctly cited both names
and then chosen one in precedence to the other (there
are many incorrect subsequent spellings, e.g. Dalla
Torre, 1896). Since the second form of the name has
been used more recently (e.g. Baker, 1996b), I suggest
that it should be given precedence. This form is a
simple noun in apposition and so retains the same
ending whatever the gender of the generic name with
which it is combined (ICZN, 1985: Article 31b(11)).

Bombus (Mg.) hortorum (Linnaeus)

hortorum (Linnaeus, 1761:424 [Apis]) examined

meridionalis Dalla Torre, 1879:13

hispanicus Pittioni, 1939c:244, not of Friese, 1911:571 (=
B. monticola Smith)

asturiensis (Tkalct, 1975:181 [Megabombus]) replace-
ment name for hispanicus Pittioni, 1939c:244

?reinigiellus (Rasmont, 1983:43 [Megabombus])

@ TAXONOMIC STATUS. The SpanishB. asturiensis
has been considered a separate species from B.
hortorum by Rasmont (1983, 1988), although they
have been treated as conspecific by Pittioni (1939c),
Tkalcti (1975), Ornosa (1986a, 1986b, 1991), Castro
(1988, 1993) and, more recently, by Rasmont ef al.
(1995). The two taxa are closely similar.

The Spanish B. reinigiellus has also been consid-
ered both as conspecific with B. hortorum (Castro,
1987) and as a separate species (e.g. Rasmont, 1983;
Castro, 1988; Ornosa, 1991).The two taxa are allopatric
(Rasmont, 1983), with B. reinigiellus being narrowly
restricted to the Sierra Nevada of Spain, possibly as a
disjunct peripheral population. B. reinigiellus is closely
similar to B. hortorum, although subtle differences in
characters of colour and morphology have been de-
scribed (e.g. Rasmont, 1983; Castro, 1988; Ornosa,
1991). From the material I have examined, the mor-
phological differences appear to be analogous to the
variation between mainland and island populations of
B. terrestris (see the comments on B. terrestris).

Depending upon the species concept embraced,
such subtle differences as those between B. reinigiellus
and B. hortorum might be expected even within a
single population and [| shall treat all three taxa as
conspecific for the present. More evidence is awaited.

COMMENT. B. hortorum has been introduced into
New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr,
1995) (see the comments on B. ruderatus, B.
subterraneus and B. terrestris). It occurs in Iceland,
where it has also probably been introduced (Prys-
Jones et al., 1981) (see the comments on B. Jucorum).

CHECKLIST OF BUMBLE BEES

Bombus (Mg.) argillaceus (Scopoli)
Argillacea (Scopoli, 1763:305 [Apis])
ligusticus Spinola, 1806:29

?Bombus (Mg.) ruderatus (Fabricius)
ruderata (Fabricius, 1775:380 [Apis]) examined
Perniger (Harris, 1776:131 [Apis])
villarricaensis Asperen de Boer, 1992a:133

@ TAXONOMICSTATUS. B. argillaceus and B. rude-
ratus are similar in most characters and differ
principally in the colour patterns of the queens (e.g.
Reinig, 1939; Fig. 11). Scholl, Obrecht & Zimmer-
mann (1992) found that hybrid queens between B.
argillaceus and B. ruderatus do occur in parts of
southeastern France, but are very rare. Whether or not
the taxa on either side of this hybrid zone are consid-
ered to be separate species therefore depends on which
species concept is preferred. Because Scholl, Obrecht
& Zimmermann (1992) estimated that only slight gene
flow is occurring, I shall continue to treat them as
separate species.

COMMENT. This species has been introduced into
New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr,
1995) (see the comments on B. hortorum, B.
subterraneus and B. terrestris) and Chile (Arretz &
Macfarlane, 1982; Asperen de Boer, 1993b). B.
ruderatus also occurs on the Azores (which have never
had a continental connection), where it may be pre-
sumed to be an introduction (Yarrow, 1967).

Bombus (Mg.) czerskii Skorikov
czerskii Skorikov, 1910b:413, examined

Bombus (Mg.) koreanus (Skorikov)
koreanus (Skorikoy, 1933a:59 [Hortobombus])
pekingensis Bischoff, 1936:21, examined
?notocastaneus Tkalcit, 1961a:52 (provisional synonym)

@ TAXONOMIC STATUS. B. notocastaneus was de-
scribed from a single male from Hubei. From the
description, it appears most likely to be conspecific
with B. koreanus.

Bombus (Mg.) melanopoda Cockerell

melanopoda Cockerell, 1910a:416, examined

@ TAXONOMIC STATUS. B. melanopoda is known
from a single female specimen (labelled ‘Sumatra’,
reverse *92.182.’) in the NHM collection in London.

According to the accessions catalogue, the number
on the label of this specimen refers to 8 Hymenoptera
presented in November 1892 by H. O. Forbes. The
area of origin is given as Borneo, although this has

117

been crossed out and Sumatra added. Forbes’ (1885)
account of his travels of 1878-1883 in Indonesia
shows that he did not visit Borneo, although he did
visit the mountains of southern Sumatra. There is no
direct account of the collection of this specimen,
although he recorded bees from at least three possible
localities at higher altitudes: first, in late 1880 he
climbed Gunung Tenggamus, where he recorded (p.
159) ‘a few bees’ at 7200 ft (2160 m); second, in 1881
he visited Gunung Dempa, where he recorded (p. 208)
‘a fine grey-haired humble-bee (Bombus senex)’ (iden-
tification by Forbes) between 7000-7700 ft
(2100-2310 m); and later in the same year, near the
summit of Gunung Kaba (1983 m), he recorded again
(p. 228) ‘A large humble-bee (Bombus senex)’.

B. melanopoda appears to be a morphologically
distinct species. It can be distingished from the only
other long-tongued bumblebee species known from
Sumatra, B. senex, by the longer oculo-malar area of
B. melanopoda, which is more than 1.5 times longer
than the basal breadth of the mandible. Otherwise,
the most closely related long-tongued bumble bee in
any neighbouring area is B. trifasciatus from the
Cameron Highlands of Peninsular Malaysia. How-
ever, like most queens of the subgenus Megabombus,
the holotype of B. melanopoda is easily distin-
guished by its narrow longitudinal median groove
subapically on gastral tergum VI (for B. trifasciatus
this area is uniformly convex). The colour pattern of
the B. melanopoda female is predominantly black,
but the hairs of terga [V—V are very slightly paler, so
there is some similarity to the darkest queens of B.
koreanus, which have this pubescence brownish
cream. However, unlike the few queens of B.
koreanus available to me, the type of B. melanopoda
has the unpunctured areas around the ocelli extend-
ing to less than half the ocello-ocular distance; the
dorsal furrow of the gena (between the vertex and
the post-ocular area) is strongly marked anteriorly;
and the dorsal face of the labral tubercles (the face
adjacent to the clypeus) is more sharply separated
from the anterior ventral face and more densely
marked by moderate-sized punctures.

Since no further individuals have been found, the
possibility that the holotype of B. melanopoda is a
mislabelled melanic specimen of another species of
the subgenus Megabombus ought to be explored, per-
haps initially through a morphometric analysis.

COMMENT. Like most other Sumatran bumble bees
(with the notable exception of the extensively greyish-
white queens of B. senex, see Sianturi et al., 1995), the
holotype of B. melanopoda is almost entirely black. It
is likely that females of this species would be particu-
larly easily mistaken for black individuals of B. senex
(although males of B. melanopoda might be paler, as
for B. koreanus).

118

Bombus (Mg.) securus (Frison)
securus (Frison, 1935:346 [Bremus]) examined
yuennanicus Bischoff, 1936:23, examined

Bombus (Mg.) religiosus (Frison)
religiosus (Frison, 1935:344 [Bremus]) examined

Subgenus RHODOBOMBUS Dalla Torre

Bombus (Rhodobombus) Dalla Torre, 1880:40, type-spe-
cies Bremus pomorum Panzer (= Bombus pomorum
(Panzer)) by subsequent designation of Sandhouse,
1943:596

Bombus (Pomobombus) Kriiger, 1917:65, type-species
Bremus pomorum Panzer (= Bombus pomorum (Pan-
zer)) by subsequent designation of Sandhouse, 1943:589

Pomibombus Skorikov, 1938a:145, unjustified emenda-
tion

Bombus (Rh.) armeniacus Radoszkowski
armeniacus Radoszkowski, 1877b:202

Bombus (Rh.) mesomelas Gerstaecker®
Agrorum (Schrank, 1781:397 [Apis])
arvenjfis (Gmelin in Linnaeus, 1790:2786 [Apis]) unjusti-
fied replacement name for agrorum Schrank, 1781:397
mesomelas Gerstaecker, 1869:321

O NOMENCLATURE. Warncke (1986) listed B.
agrorum (Schrank) as questionably conspecific with
B. distinguendus, but without any explanation. Al-
though I know of no extant type specimens, Schrank’s
(1781) description of his B. agrorum of ‘Habitat rurv
from Austria appears to me to be almost certainly of
the same species as B. mesomelas, because the head is
described as black and the pale hairs of the thorax and
of gastral tergum I are described as grey, with the
remainder of the gaster rusty or tawny-yellow (the
head and the pale pubescence of the thorax and gaster
are more uniformly dull yellowish fa B.
distinguendus). See the comments onB. distinguendus.

& APPLICATION TO ICZN. Although B. agrorum is
the oldest available name for the present interpretation
of this species, the name B. mesomelas has been in

P.H. WILLIAMS

common use for the species since 1947 (e.g. Tkalcii,
1969, 1975; Delmas, 1976; Reinig, 1974, 1981; Ozbek,
1983; Rasmont, 1983; Ornosa, 1986a, b; Rasmont et
al., 1987, 1995). I know of no publications using the
name B. agrorum (Schrank) since 1947, although the
name B. agrorum (Fabricius) was in widespread use
for another species (= B. pascuorum) until Richards
(1968). It is suggested that, in the interests of stability
(ICZN, 1985: Article 23b), an application be made to
ICZN to use its Plenary Power to suppress the unused
senior synonym, B. agrorum, and its unjustified re-
placement name, B. arvensis (ICZN, 1985: Article 79)
(see the comments on B. muscorum).

Bombus (Rh.) pomorum (Panzer)
pomorum (Panzer, 1805(86):18 [Bremus])
Lefebvrei Lepeletier, 1836:461

Subgenus KALLOBOMBUS Dalla Torre

Bombus (Kallobombus) Dalla Torre, 1880:40, type-spe-
cies Apis soroeensis Fabricius (= Bombus soroeensis
(Fabricius)) by subsequent designation of Sandhouse,
1943:561

Bombus (Callobombus) Dalla Torre, 1896:503, unjustified
emendation

Bombus (Soroeensibombus) Vogt, 1911:63, type-species
Apis soroeensis Fabricius (= Bombus soroeensis
(Fabricius)) by monotypy

[Bombus (Soroensibombus) Ball, 1914:78, incorrect sub-
sequent spelling]

[Sorocoénsibombus Skorikoy, 1922a: map 15, incorrect
subsequent spelling]

Bombus (KI.) soroeensis (Fabricius) @

Cardui (Miller, 1776:165 [Apis])

Joroeenfis (Fabricius, [1777, see Baker, 1996a:9]:246
[Apis})

Proteus Gerstaecker, 1869:325

perplexus Radoszkowski, 1884:82, not of Cresson, 1863:91
(= B. perplexus Cresson)

Radoszkowskyi Dalla Torre, 1890:139, replacement name
for perplexus Radoszkowski, 1884:82

miniatocaudatus Vogt, 1909:56

CHECKLIST OF BUMBLE BEES

that the name B. cardui has narrow priority over B.
soroeensis by publication date.

© APPLICATION TOICZN. AlthoughB. cardui is the
oldest available name for the present interpretation of
this species, the name B. soroeensis has been in com-
mon use for the species since 1947 (e.g. Tkalct, 1969,
1975; Lgken, 1973; Alford, 1975; Delmas, 1976;
Pekkarinen, 1979; Reinig, 1981; Ozbek, 1983;
Rasmont, 1983; Ito, 1985; Ornosa, 1986a; Rasmont et
al., 1995). The only publications using the name B.
cardui since 1947 are those of Baker (1996a, b). Using
this name contrary to the purpose of priority is not
accepted as usage in the sense of the Code (ICZN,
1985: Article 23b), and so cannot justify the continued
use of the name B. cardui in place of B. soroeensis. It
is suggested that, in the interests of stability, an appli-
cation be made to ICZN to use its Plenary Power to
suppress the unused senior synonym (ICZN, 1985:
Article 79) (see the comments on B. muscorum).

Subgenus ALPINOBOMBUS Skorikoy
Alpinobombus Skorikov, 1914a:123, type-species Apis
alpina Linnaeus (= Bombus alpinus (Linnaeus)) by
subsequent designation of Frison, 1927:66
Bombus (Alpinobombus) Kriiger, 1917:62
Alpinibombus Skorikoy, 1937:53, unjustified emendation

COMMENT. Species of the subgenus Alpinobombus
make up the most northerly distributed of all bee
faunas (e.g. K. W. Richards, 1973). Indeed, three of the
five species have a nearly circumpolar distribution, as
a major component of an Arctic bumble bee fauna
(Williams, 1996b). This relative homogeneity of the
Arctic fauna among northern continents resembles the
pattern in the Arctic flora, which shows little regional
differentiation in comparison with more southern flo-
ras (Hooker, 1861; Walker, 1995).

Bombus (Al.) hyperboreus Schonherr
Arctica (Quenzel in Acerbi, 1802:253 [Apis])
hyperboreus Schénherr, 1809:57, unjustified replacement
name for arcticus Quenzel, 1802:253
clydensis Yarrow, 1955:151, examined

119

@ TAXONOMIC STATUS. The identity of B. arcticus
(Quenzel) has been uncertain. Warncke (1986) listed
B. arcticus (Quenzel) as conspecific withB. lapponicus
without any explanation. Presumably this was because
B. lapponicus is extensively pale on the dorsum, al-
though the pale pubescence is differentiated into yellow
and red areas and much of it is much paler than
Quenzel’s description. | agree with Lgken (1973) that,
from the original description and the illustration (no
type specimen is known to exist), B. arcticus (Quenzel)
is most likely to be conspecific with B. hyperboreus,
which has the pale pubescence uniformly brownish
yellow.

OQ NOMENCLATURE. The name B. arcticus has
rarely been used for this species in preference to B.
hyperboreus, and perhaps only as a misidentification
of B. arcticus Kirby (see e.g. Franklin, 1913; Richards,
193 1a). Loken (1973) considered B. arcticus (Quenzel)
to be a nomen oblitum, so she continued to use the
name B. hyperboreus. However, nomina oblita are not
supported for a publication of this date by the present
Code (ICZN, 1985: Article 79c(ii1)), although it does
allow that B. arcticus (Quenzel) could be suppressed
by use of the Plenary Power. See the comments on B.
polaris Curtis.

€ APPLICATION TO ICZN. Although B. arcticus is
the oldest available name for the present interpretation
of this species, the name B. hyperboreus has been in
common use for the species since 1947 (e.g. Loken,
1973; Milliron, 1973a; K.W. Richards, 1973; Svensson
& Lundberg, 1977; Hurd, 1979; Pekkarinen, 1979;
Pekkarinen ef al., 1981; Reinig, 1981; Rasmont, 1983;
Pekkarinen & Teras, 1993). It is suggested that, in the
interests of stability (ICZN, 1985: Article 23b), an
application be made to ICZN to use its Plenary Power
to suppress the unused senior synonym (ICZN, 1985:
Article 79), in order to confirm usage of B. hyperboreus
as the valid name (see the comments on B. muscorum).

COMMENT. B. hyperboreus has been suggested to be
a social parasite in colonies of B. polaris, at least
facultatively in some parts of its range (Milliron &
Oliver, 1966; Loken, 1973: K. W. Richards, 1973). See
the comments on Psithyrus and B. inexspectatus.

Bombus (Al.) balteatus Dahlbomes
balteatus Dahlbom, 1832:36
nivalis Dahlbom, 1832:40
tricolor Dahlbom, 1832:41
?Kirbiellus Curtis in Ross, 1835:1xii
kirbyellus Dalla Torre, 1896:527, unjustified emendation
tristis Sparre-Schneider in Friese, 1902:495, not of Seidl,
1837:69 (= B. humilis Iliger)

@ TAXONOMIC status. B. balteatus and B. kirbi-
ellus have been considered conspecific by most authors
(e.g. Thomson, 1872; Richards, 1931a; Skorikov, 1937;

120

Pittioni, 1942; Loken, 1973; Hurd, 1979; Thorp et al.,
1983), although Milliron (1973a) considered them to
be separate species that co-occur in some areas, par-
ticularly in Alaska.

Milliron (1973a) described several characters by
which to discriminate B. balteatus and B. kirbiellus,
placing particular emphasis on the shape of male
gastral sternum VIII and the female malar area.

From the small samples I have examined, I have
been unable to find convincing evidence of discrete
differences in these characters. Until more evidence to
the contrary is available from critical studies of pat-
terns of variation, I shall treat them as parts of a single
variable species.

O NOMENCLATURE. Richards (193 1a) believed B.
balteatus, B. nivalis and B. tricolor to be conspecific
and selected the name B. balteatus to have precedence
because it was published on an earlier page (page
priority is not a mandatory part of the Code, only a
recommendation, see ICZN, 1985: Recommendation
24A). However, Thomson (1872:35) had already cho-
sen the name B. nivalis in precedence to B. balteatus
and, following the Principle of the First Reviser (ICZN,
1985: Article 24), Thomson’s action should now stand.
Consequently, the valid name for this species is B.
nivalis, although the Code (ICZN, 1985) allows that this
name could be suppressed by use of the Plenary Power.

€ APPLICATIONTOICZN. AlthoughB. nivalis is the
valid name for this species, the name B. balteatus has
been in common use for the species since 1947 (e.g.
Loken, 1973; Milliron, 1973a; Plowright & Stephen,
1973; Hurd, 1979; Pekkarinen, 1979; Reinig, 1981;
Rasmont, 1983; Thorp er al., 1983; Laverty & Harder,
1988; Pekkarinen & Teras, 1993). Itis suggested that, in
the interests of stability, an application be made to ICZN
to use its Plenary Power to suppress the unused name
(ICZN, 1985: Article 79) (see the comments on B.
muscorum). However, the consequence of this action
would be that nivalis Dahlbom would no longer be
available for a subspecies of B. balteatus.

Bombus (Al.) neoboreus Sladen
strenuus Cresson, 1863:102, not of Harris, 1776:131 (=B.
lapidarius (Linnaeus))
neoboreus Sladen, 1919:28

O NOMENCLATURE. B. strenuus Cresson (1863) is
a junior secondary homonym in Bombus of Apis
strenuus Harris (1776), and therefore the name B.
strenuus Cresson is invalid (ICZN, 1985: Article 57c).
For this species, the oldest available name is B.
neoboreus, which becomes the valid name. The only
publications using the name B. strenuus Cresson since
1947 of which I am aware are by Hurd (1979), Milliron
(1973a) and Poole (1996), so this change of valid
name is not a serious disruption of common usage.

P.H. WILLIAMS

Bombus (AL) polaris Curtis
Arcticus Kirby in Parry, 1824:ccexvi, examined, not of
Quenzel in Acerbi, 1802:253 (= B. hyperboreus
Sch6nherr)
Polaris Curtis in Ross, 1835:1xiii, examined
diabolicus Friese, 1911:571
alpiniformis Richards, 1931a:13

© NOMENCLATURE. Loken (1973) used the name
B. arcticus Kirby for this species because she consid-
ered B. arcticus (Quenzel) to be a nomen oblitum.
However, this is not supported by the present Code for
a publication of this date (ICZN, 1985: Article 79c(iii)),
although it does allow B. arcticus (Quenzel) to be
suppressed by use of the Plenary Power. See the
comments on B. hyperboreus.

APPLICATION TO ICZN. It is suggested above
that, in the interests of stability ICZN, 1985: Article
23b), an application be made to ICZN to use its
Plenary Power to suppress B. arcticus (Quenzel), the
unused senior synonym (ICZN, 1985: Article 79) of B.
hyperboreus. This would free B. arcticus Kirby from
junior primary homonymy with B. arcticus (Quenzel)
(ICZN, 1985: Article 57b), so that it would become the
valid name for this species (see the comments on B.
hyperboreus). However, although the name B. arcticus
Kirby has been in use for this species (e.g. Loken,
1973; Sakagami, 1976; Svensson & Lundberg, 1977;
Reinig, 1981), the more frequently used name has
been B. polaris (e.g. Milliron & Oliver, 1966; Milliron,
1973a: K.W. Richards, 1973; Hurd, 1979; Pekkarinen,
1979; Pekkarinen ef al., 1981; Rasmont, 1983;
Pekkarinen & Teris, 1993). In the interests of stability,
the application to ICZN might be extended to suppress
B. arcticus Kirby, in order to conserve the current
usage of B. polaris as the valid name.

Bombus (AI.) alpinus (Linnaeus)
alpina (Linnaeus, 1758:579 [Apis]) examined

Subgenus SUBTERRANEOBOMBUS Vogt
Bombus (Subterraneobombus) Vogt, 1911:62, type-spe-
ciesApis subterranea Linnaeus (=Bombus subterraneus
(Linnaeus)) by subsequent designation of Frison,
1927:68

CHECKLIST OF BUMBLE BEES

Subterraneibombus Skorikov, 1938a:145, unjustified
emendation

Bombus (St.) melanurus Lepeletier
melanurus Lepeletier, 1836:469, examined
?difficillimus Skorikov, 1912:609, examined
subdistinctus Richards, 1928a:333, examined

@ TAXONOMIC sTaTUS. Several of these nominal
taxa have been treated as separate species and at least
B. difficillimus may indeed prove to be a separate
species. However, aside from differences in colour
pattern, they are closely similar in morphology with a
range of variation (Williams, 1991). Until more evi-
dence to the contrary is available from critical studies
of patterns of variation, I shall treat them as parts of a
single variable species.

Bombus (St.) fragrans (Pallas)
fragrans (Pallas, 1771:474 [Apis])
mongol Skorikoy, 1912:607, examined
?charharensis Yasumatsu, 1940:94 (provisional synonym)

@ TAXONOMIC STATUS. This taxon is interpreted
here in the broadest sense, to include a complex of
poorly-known taxa (Williams, 1991). More evidence
is awaited.

Bombus (St.) amurensis Radoszkowski
Amurensis Radoszkowski, 1862:590, examined

@ TAXONOMIC sTaTUS. I have seen no males of
this species and its precise relationships remain un-
clear.

Bombus (St.) fedtschenkoi Morawitz
Fedtschenkoi Morawitz in Fedtschenko, 1875:5

Bombus (St.) personatus Smith
personatus Smith, 1879:132, examined
Roborowskyi Morawitz, 1886:197, examined

Bombus (St.) subterraneus (Linnaeus)
fubterranea (Linnaeus, 1758:579 [Apis]) examined
Nemorum (Scopoli, 1763:307 [Apis])

? faltuum (Panzer, 1801(75):21 [Apis])

O NOMENCLATURE. Loken (1984) interpreted B.
saltuum as being conspecific with B. barbutellus (see
the comments on B. barbutellus). Warncke (1986)
interpreted B. saltuum as having been described from
a male (presumably because the antennae were de-
scribed as rather long) conspecific with B.
subterraneus. No type specimen is known. The de-

121

scription of the anterior part of the gaster of B. saltuum
as ashen and the middle part as nearly bald is perhaps
slightly closer to B. subterraneus, because although
both species may have gastral tergum I with pale hair
and terga I-III sparsely haired, this hair is much shorter
on B. subterraneus and the posterior fringing hairs of
the terga often appear greyish-white.

If B. saltuum were not accepted as most likely to be
conspecific with B. subterraneus, then further action
would be required. If both interpretations were con-
sidered to remain supportable, then it might be
considered appropriate (ICZN, 1985: Article 75b) to
designate a specimen of B. subterraneus as neotype
of Apis saltuum in order to conserve the current usage
of B. barbutellus. Alternatively, if B. saltuum were
considered more likely to be conspecific with B.
barbutellus, then B. saltuum would become the oldest
available name for that species, even though the name
has not been used in the last 50 years. In the interests
of stability (ICZN, 1985: Article 23b), an application
could then be made to ICZN to use its Plenary Power
to suppress the unused senior synonym (ICZN, 1985:
Article 79).

COMMENT. This species has been introduced into
New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr,
1995).

Bombus (St.) distinguendus Morawitz
nemorum (Fabricius, 1775:382 [Apis]) not of Scopoli,
1763:307 (= B. subterraneus (Linnaeus)), not of
Fabricius, 1775:380 (?= B. bohemicus Seidl)
elegans Seidl, 1837:67
distinguendus Morawitz, 1869:32

O NOMENCLATURE. The nameB. elegans has been
applied to several taxa by different authors. Tkalcit
(1969:901-903) reasoned that Seidl had originally
described B. elegans from an individual of the species
that has more recently been known by the name B.
distinguendus, although Seidl’s original type is lost.
According toTkalcti, a specimen of B. mesomelas may
then have been substituted as the type, but now this
cannot be found either. Any remaining confusion could
be resolved by the designation of an appropriate
neotype. See the comments on B. mesomelas.

€ APPLICATIONTOICZN. AlthoughB. elegans may
be the oldest available name for the present interpreta-
tion of this species, the name B. distinguendus has
been in common use for the species since 1947 (e.g.
Tkalcti, 1969, 1974a; Loken, 1973; Alford, 1975;
Delmas, 1976; Sakagami, 1976; Pekkarinen, 1979;
Reinig, 1981; Pekkarinen er al/., 1981; Rasmont, 1983;
Pekkarinen & Teréas, 1993; Rasmont ef al., 1995). I
know of no publications using the name B. elegans for
this taxon (only for B. mesomelas Gerstaecker as a
misidentification) since 1947. It is suggested that, in

122

the interests of stability (ICZN, 1985: Article 23b),
and to prevent confusion with B. mesomelas, an appli-
cation be made to ICZN to use its Plenary Power to
suppress the unused senior synonym (ICZN, 1985:
Article 79) (see the comments on B. muscorum).

Bombus (St.) appositus Cresson
appositus Cresson, 1878:183

Bombus (St.) borealis Kirby
borealis Kirby, 1837:272

Subgenus ALPIGENOBOMBUS Skorikov

Alpigenobombus Skorikov, 1914a:128, type-species
Alpigenobombus pulcherrimus Skorikov (= Bombus
kashmirensis Friese) by subsequent designation of
Williams, 1991:65

Bombus (Mastrucatobombus) Kriger, 1917:66, type-spe-
cies Bombus mastrucatus Gerstaecker (= Bombus
wurflenii Radoszkowski) by monotypy

Bombus (Alpigenobombus) Frison, 1927:64

[Nobilibombus Skorikov, 1933a:62, published without fixa-
tion of type-species]

[Bombus (Nobilibombus) Bischoff, 1936:12, type-species
Nobilibombus morawitziides Skorikov (= Bombus
nobilis Friese) by monotypy, published as a junior
synonym]

Alpigenibombus Skorikov, 1938b:1, unjustified emenda-
tion

[Pyrobombus (Nobilibombus) Milliron, 1961:54, type-spe-
cies Bombus nobilis Friese (cited as Bombus nobilis
Skorikov) by original designation, published as a junior
synonym]

Bombus (Nobilibombus) Richards, 1968:222, type-species
Bombus nobilis Friese by original designation (see
Williams, 1991)

[Alpegenobombus Wang, 1979:188, incorrect subsequent
spelling]

@ TAXONOMIC sTATUS. Richards (1968) treated
Alpigenobombus and Nobilibombus as separate
subgenera. Following Bischoff (1936), I have treated
them as a single subgenus Alpigenobombus (Williams,
1991), for which the evidence for monophyly is strong
(Williams, 1995).

P.H. WILLIAMS

Bombus (Ag.) kashmirensis Friese
kashmirensis Friese, 1909[September, Tkalcii, 1974b]:673
examined
stramineus Friese, 1909[September, Tkalct, 1974b]:673
tetrachromus Cockerell, 1909[November, Tkalci,
1974b]:397, examined
pulcherrimus (Skorikov, 1914a:128 [Alpigenobombus])

@ TAXONOMIC sTaTUS. Several of these nominal
taxa have been treated as separate species. However,
aside from differences in colour pattern, they are
closely similar in morphology with a range of varia-
tion (Williams, 1991). Until more evidence to the
contrary is available from critical studies of patterns of
variation, I shall treat them as parts of a single variable
species.

O NOMENCLATURE. Tkalcti (1974b) first regarded
B. kashmirensis and B. stramineus as conspecific and,
following the Principle of First Reviser (ICZN, 1985:
Article 24), chose B. kashmirensis as the name for the
species.

Bombus (Ag.) wurflenii Radoszkowski
Wurflenii Radoszkowski, 1859:482, examined
[Wurfleini Radoszkowski, 1877b:191, incorrect subsequent
spelling]
mastrucatus Gerstaecker, 1869:326, examined
alpigenus Morawitz, 1874:132

Bombus (Ag.) nobilis Friese
?validus Friese, 1905:510, examined (provisional syno-
nym)
nobilis Friese, 1905:513
?sikkimi Friese, 1918:82, examined (provisional syno-

nym)

[morawitziides Skorikov, 1922a:159, published without
description]

[moravitziides Skorikov, 1931:203, published without de-
scription]

?morawitziides (Skorikov, 1933a:62 [Nobilibombus]) ex-
amined (provisional synonym)

Dxizangensis Wang, 1979:188, examined (provisional syno-
nym)

chayaensis Wang, 1979:189, examined, new synonym

@ TAXONOMIC STATUS. Several of these nominal
taxa have been treated as separate species.

Friese (1905) described B. validus initially (p. 510)
as having a quadrate malar area and untoothed mandi-
bles, but went on (p. 517) to place it within the
mastrucatus (= B. wurflenit) group, which he charac-
terised as having a short malar area and toothed
mandibles. Tkalcii (1987) designated as lectotype of
B. validus a female with a quadrate malar area and
multi-toothed mandibles. He also synonymised B.
morawitziides with B. validus.

The type specimens of B. nobilis have also been in
some doubt (Richards, 1968). In the same publication

CHECKLIST OF BUMBLE BEES

as the description of B. validus, Friese (1905) de-
scribed the female of B. nobilis as having a‘quadratisch’
malar area and 4—5 teeth on the mandible (even though
he placed it [p. 519] ina group withB. lapidarius).The
original description lists several females (particularly
from Sichuan), but the only putative type female that I
have been able to examine (although it carries no
Friese ‘type’ label) is in the Berlin museum collection
and is a specimen of B. friseanus labelled ‘Kashgar’
(this locality is outside the known distribution range of
either B. nobilis or B. friseanus). The specimen does
not match the original description of the mandibles of
B. nobilis and so cannot be considered a valid syntype.
Nonetheless, the identity of B. nobilis is clear from the
original description, so the designation of a neotype is
not justified (ICZN, 1985: Article 75b).

B. chayaensis appears to me to be very closely
similar to the yellow banded B. nobilis (in the strict
sense) and I am unaware of any reason to treat them as
separate species.

B. nobilis is interpreted here in the broadest sense,
to include a complex of morphologically closely simi-
lar taxa (Williams, 1991). At least some of the taxa
included may prove to be separate species from B.
nobilis. The most obvious variation is in the colour of
the pale thoracic bands, which may be yellow (B.
nobilis), yellow-white (B. sikkimi), gey-white (B.
morawitziides), or almost completely replaced by black
(B. validus). However, aside from these differences in
colour pattern, they are similar in morphology with a
range of variation. Until more evidence to the contrary
is available from critical studies of patterns of varia-
tion, I shall treat them as parts of a single variable
species.

O NOMENCLATURE. Following the Principle of
First Reviser (ICZN, 1985: Article 24), and as the first
author to regard these taxa as conspecific, I select the
name B. nobilis as the valid name in preference to B.
validus from the available names for this species from
Friese (1905).

Bombus (Ag.) genalis Friese

genalis Friese, 1918:84, examined

@ TAXONOMIC sTaTUS. Ihave seen no males of this
species and its precise relationships remain unclear.

Bombus (Ag.) grahami (Frison)
grahami (Frison, 1933:334 [Bremus])

Bombus (Ag.) breviceps Smith
nasutus Smith, 1852a:44, examined
breviceps Smith, 1852a:44, examined
dentatus Handlirsch, 1888:227
simulus Gribodo, 1892:114, examined

123

orichalceus Friese, 1916:107

rufocognitus Cockerell, 1922:4, examined

pretiosus Bischoff, 1936:11, examined, not of Friese,
1911:571 (= B. polaris Curtis)

2angustus Chiu, 1948:59 (provisional synonym)

bischoffiellus (Tkalct, 1977:224 [Alpigenobombus]) re-
placement name for pretiosus Bischoff, 1936:11

@ TAXONOMIC sTATUS. Several of these nominal
taxa have been treated as separate species. At least B.
dentatus [Himalaya] and B. angustus [Taiwan] may
prove to be separate species (e.g. Tkalct, 1968), 1989).
However, aside from differences in colour pattern,
they are similar in morphology with a range of varia-
tion (Williams, 1991). Until more evidence to the
contrary is available from critical studies of patterns of
variation, I shall treat them as parts of a single variable
species.

O NOMENCLATURE. Tkalcii (1968d) first regarded
B. nasutus and B. breviceps as likely to be conspecific
and, following the Principle of First Reviser (ICZN,
1985: Article 24), chose B. breviceps as the name for
the species.

Subgenus PYROBOMBUS Dalla Torre

Bombus (Pyrobombus) Dalla Torre, 1880:40, type-species
Apis hypnorum Linnaeus (= Bombus hypnorum
(Linnaeus)) by monotypy

Bombus (Poecilobombus) Dalla Torre, 1882:23, type-spe-
cies Bombus sitkensis Nylander by subsequent
designation of Sandhouse, 1943:589

[Bombus (Pyrrhobombus) Dalla Torre, 1882:28, incorrect
subsequent spelling]

Bombus (Pyrrhobombus) Dalla Torre, 1896:503, unjusti-
fied emendation

Bombus (Pratobombus) Vogt, 1911:49, type-species Apis
pratorum Linnaeus (= Bombus pratorum (Linnaeus))
by subsequent designation of Frison, 1927:67

[Bombus (Pratibombus) Ball, 1914:78, incorrect subse-
quent spelling]

Bombus (Anodontobombus ) Kriiger, 1917:61,65 (proposed
as a section name but stated by Milliron, 1961:53, to be
synonymous with his concept of the subgenus
Pyrobombus Dalla Torre), type-species Apis hypnorum
Linnaeus (= Bombus hypnorum (Linnaeus)) by subse-
quent designation of Williams, 1991:69

124

Bombus (Uncobombus) Vogt in Kriiger, 1917:65 (pro-
posed as a group name but stated by Milliron, 1961:53,
to correspond to his concept of Pyrobombus Dalla
Torre), type-speciesApis hypnorum Linnaeus (=Bombus
hypnorum (Linnaeus)) by subsequent designation of
Williams, 1991:69

Bombus (Lapponicobombus) Quilis-Pérez, 1927:19, type-
speciesApis lapponica Fabricius (=Bombus lapponicus
(Fabricius)) by subsequent designation of Milliron,
1961:58

[Bombus (Hypnorubombus) Quilis-Pérez, 1927:19, incor-
rect original spelling]

[Bombus (Laponicobombus) Quilis-Pérez, 1927:63, incor-
rect original spelling]

Bombus (Hypnorobombus) Quilis-Pérez, 1927:97, type-
species Apis hypnorum Linnaeus (= Bombus hypnorum
(Linnaeus)) by monotypy

Pratibombus Skorikoy, 1938b:1, unjustified emendation

COMMENT. This is the largest subgenus of Bombus.
The highest richness of Pyrobombus species occurs in
the New World (there are no species known from south
of Panama), although the earliest-diverging species
within the subgenus appear to be in the Old World
(Williams, 1991). This is the opposite pattern to that
shown by the next-largest subgenus (of social para-
sites), Psithyrus (see the comments on the subgenus
Psithyrus).

Bombus (Pr.) abnormis (Tkalci)
abnormis (Tkalcii, 1968a:33 [Pyrobombus]) examined

Bombus (Pr.) hypnorum (Linnaeus)

Hypnorum (Linnaeus, 1758:579 [Apis]) examined

leucopygus Wlliger, 1806:172

calidus Erichson in Middendorff, 1851:65

fletcheri Richards, 1934:90, examined

insularis Sakagami & Ishikawa, 1969:180, not of Smith,
1861:155 (= B. insularis (Smith))

Koropokkrus Sakagami & Ishikawa, 1972:610, replace-
ment name for insularis Sakagami & Ishikawa, 1969:180

@ TAXONOMIC sTaTuS. B. hypnorum is a broadly
distributed species with a fairly easily recognised
brown-black-white colour pattern (e.g. Reinig, 1939;
Williams, 1991). Recently, Starr (1992) has described
what appears to be a divergent, brownish-black or-
ange-tailed colour form from a disjunct peripheral
population on the island of Taiwan.

Bombus (Pr.) perplexus Cresson
perplexus Cresson, 1863:91

Bombus (Pr.) haematurus Kriechbaumer
haematurus Kriechbaumer, 1870:157

Bombus (Pr.) subtypicus (Skorikov)
leucopygus Morawitz in Fedtschenko, 1875:3, not of Illiger,

P.H. WILLIAMS

1806:172 (= B. hypnorum (Linnaeus))

[leucopygos (Skorikov, 1914b:294 [Pratobombus]) incor-
rect subsequent spelling]

subtypicus (Skorikoy, 1914b:294 [Pratobombus]) exam-
ined

leucurus Bischoff & Hedicke, 1931:391, replacement name
for leucopygus Morawitz in Fedtschenko, 1875:3

kohistanensis (Tkalcti, 1989:49 [Pyrobombus]) examined

Bombus (Pr.) mirus (Tkalcii)
mirus (Tkalcu, 1968a:37 [Pyrobombus]) examined
?tibetanus Friese, 1913:86, examined, not of Morawitz,
1886:202 (= B. tibetanus (Morawitz))

Bombus (Pr.) lemniscatus Skorikov
lemniscatus Skorikov, 1912:607, examined
flavopilosus Friese, 1918:84, examined
peralpinus Richards, 1930:646, examined

Bombus (Pr.) lepidus Skorikoy
lepidus Skorikoy, 1912:606, examined
genitalis Friese, 1913:85, examined
tetrachromus Friese, 1918:85, examined, not of Cockerell,
1909:397 (= B. kashmirensis Friese)
?yuennanicola Bischoff, 1936:7, examined

Bombus (Pr.) infirmus (Tkalcit)
leucurus Bischoff, 1936:8, examined, not of Bischoff &
Hedicke, 1931:391 (= B. subtypicus (Skorikov))
infirmus (Tkalct, 1968a:24 [Pyrobombus]) replacement
name for /eucurus Bischoff, 1936:8

Bombus (Pr.) parthenius Richards
parthenius Richards, 1934[14April, Williams & Cameron,
1993]:89, examined
?sonani (Frison, 1934[30 April, Williams & Cameron,
1993]:175 [Bremus]) examined
?infrequens (Tkalctt, 1989:56 [Pyrobombus]) examined
(provisional synonym)

@ TAXONOMIC STATUS. Several of these nominal
taxa have been treated as separate species. At least B.
sonani [Taiwan] and B. infrequens [northern Burma to
southern China] may prove to be separate species.
However, aside from differences in colour pattern,
they are closely similar in morphology (Williams,
1991). Until more evidence to the contrary is available
from critical studies of patterns of variation, I shall
treat them as parts of a single variable species.

Bombus (Pr.) luteipes Richards
luteipes Richards, 1934:89, examined
?avanus (Skorikoy, 1938b:2 [Pratibombus]) (provisional
synonym)
signifer (Tkalctt, 1989:52 [Pyrobombus]), examined

@ TAXONOMIC STATUS. The identity of B. avanus

CHECKLIST OF BUMBLE BEES

is in doubt because the type cannot be found. The
description of the colour pattern resembles B.
parthenius and B. luteipes, and the description of the
longer than usual squama (= gonostylus) and the half-
crooked apex of the sagitta (= penis valve) appear to be
closely similar to B. luteipes, so these taxa are very
likely to be conspecific. More evidence is awaited.

Bombus (Pr.) flavescens Smith
flavescens Smith, 1852a:45, examined
mearnsi Ashmead, 1905:959
baguionensis Cockerell, 1920:631, new synonym
tahanensis Pendlebury, 1923:65, examined
?rufoflavus Pendlebury, 1923:66, examined (provisional
synonym)

@ TAXONOMIC sTATUS. Several of these nominal
taxa have been treated as separate species. B. rufoflavus
[Peninsular Malaysia] and B. baguionensis [Philip-
pines] are particularly distinct in colour pattern. They
may prove to be separate species, but from the material
available from a few sites, they appear to me to be
closely similarin morphology to B. flavescens (Williams,
1991). Until more evidence to the contrary is available
from critical studies of patterns of variation, I shall treat
them as parts of a single variable species.

Bombus (Pr.) rotundiceps Friese
rotundiceps Friese, 1916:108, examined
montivolanoides Sakagami & Yoshikawa, 1961:431
Shillongensis (Tkalct, 1974b:334 [Pyrobombus]) exam-
ined

Bombus (Pr.) beaticola (Tkalcii)
beaticola (Tkalcti, 1968a:28 [Pyrobombus]) examined

Bombus (Pr.) picipes Richards
flavus Friese, 1905:517, examined, not of Pérez, 1884:265
(= B. campestris (Panzer))
picipes Richards, 1934:90, examined
klapperichi Pittioni, 1949:266, examined
?nikiforuki Tkalct, 1961b:354 (provisional synonym)

@ TAXONOMIC STATUS. B. nikiforuki was described
from a single worker from Qinghai. From the descrip-
tion, it appears to be closely similar to B. picipes and is
likely to be conspecific.

O NOMENCLATURE. With Psithyrus regarded as
being a subgenus of the genus Bombus (Williams,
1991, 1995), B. pratorum subsp. flavus Friese (1905)
becomes a junior secondary homonym in Bombus of
Psithyrus campestris var. flavus Pérez (1884) (deemed
to be subspecific, see ICZN, 1985: Article 45g(ii)),
and therefore the name B. flavus Friese is invalid
(ICZN, 1985: Article 57c). For this species, the oldest
available name of which I am aware is B. parthenius

125

var. picipes Richards, 1934 (deemed to be subspecific,
see ICZN, 1985: Article 45g(11)), which becomes the
valid name, B. picipes. The only publications using the
name B. flavus Friese since 1947 of which I am aware
are by Sakagami (1972), Ito (1993) and Yao & Luo
(1997), so this change of valid name is not a serious
disruption of common usage.

Bombus (Pr.) ardens Smith
ardens Smith, 1879:133, examined
andreae Friese, 1910:405, examined

Bombus (Pr.) modestus Eversmann
modestus Eversmann, 1852:134
Baikalensis Radoszkowski, 1877b:203
nymphae Skorikoy, 1910b:409
eversmanni Skorikoy, 1910c:581, not infrasubspecific af-
ter Skorikov, 1922a:149

Bombus (Pr.) cingulatus Wahlberg
cingulatus Wahlberg, 1854:208

COMMENT. The distribution of B. cingulatus accord-
ing to Reinig (1939) is shown in Fig. 10.

?Bombus (Pr.) oceanicus Friese
oceanicus Friese, 1909:675, examined
oceanicus Friese & Wagner, 1910:52, redescribed

@ TAXONOMICSTATUS. B. oceanicus is known only
from the Kurile Islands. A particularly close relation-
ship with the otherwise broadly distributed B.
cingulatus (absent from the Kuriles, but present in
Kamchatka, Reinig, 1939; Ito & Sakagami, 1980; Fig.
10) has been suggested by Ito & Sakagami (1980) and
it is possible that they are conspecific. More evidence
is awaited.

Bombus (Pr.) brodmannicus Vogt
Brodmannicus Vogt, 1909:49, examined

Bombus (Pr.) pratorum (Linnaeus)
pratorum (Linnaeus, 1761:424 [Apis]) examined

COMMENT. This species was deliberately introduced
into Sydney, Australia, although it is not known to
have persisted (Oliff, 1895). Until the twentieth cen-
tury, B. pratorum was not known from Ireland, where
it is now well established (see references in Alford,
1975, 1980) (see comments on B. monticola).

Bombus (Pr.) jonellus (Kirby)
Jonella (Kirby, 1802:338 [Apis]) examined
alboanalis Franklin, 1913:385

126

@ TAXONOMIC status. B. alboanalis has been
regarded both as a separate species (Franklin, 1913;
Frison, 1927) and as conspecific with either B. frigidus
(Burks, 1951; Hurd, 1979; Poole, 1996) or B. jonellus
(Williams, 1991 [as B. jonellus from western Canada];
Scholl et al., 1995).

Recently, Scholl et al. (1995) concluded from stud-
ies of enzyme mobility morphs that whereas B.
alboanalis and B. frigidus have separate gene pools, in
contrast, B. alboanalis and B. jonellus show a low
level of genetic differentiation. They also noted the
lack of colour gradation between sympatric B.
alboanalis and B. frigidus.

From the limited amount of material I have exam-
ined, I believe that B. alboanalis and B. jonellus are
morphologically closely similar. Until more evidence
to the contrary is available from critical studies of
patterns of variation, I shall treat them as parts of a
single variable species.

Bombus (Pr.) pyrenaeus Pérez
pyrenaeus Pérez, [1880, see Baker, 1996d:300]:127, not of
Lepeletier, 1832:375 (= B. rupestris (Fabricius))
tenuifasciatus Vogt, 1909:49
[pyreneus Pagliano, 1995:23, incorrect subsequent spell-
ing]

O NOMENCLATURE. With Psithyrus regarded as
being a subgenus of the genus Bombus (Williams,
1991, 1995), B. pyrenaeus Pérez (1880) becomes a
junior secondary homonym in Bombus of Psithyrus
pyrenaeus Lepeletier (1832), and therefore the name
B. pyrenaeus Pérez is invalid (ICZN, 1985: Article
57c). The next available name, tenuifasciatus, was
used by Vogt (1909) for individuals with particular
colour patterns from both B. pyrenaeus Pérez and B.
sichelii. The choice of which of these two homonyms
should have precedence depends on the Principle of
the First Reviser (ICZN, 1985: Article 24). As far as I
have been able to discover, Tkalciti (1973:266) is the
first author to have recognised this problem. He recog-
nised precedence for B. pyrenaeus ssp. tenuifasciatus
Vogt. Consequently, the oldest available name for this
species, and therefore the valid name, is B.
tenuifasciatus.

& APPLICATION TO ICZN. Although B.
tenuifasciatus is the oldest available name for this
species, the name B. pyrenaeus has been in common
use for the species since 1947 (e.g. Krusemen, 1958;
Tkalct, 1969, 1973, 1975; Reinig, 1972, 1981; Delmas,
1976; Rasmont, 1983; Ornosa, 1986; Williams, 1991;
Rasmont ef al., 1995). It is suggested that, in the
interests of stability, an application be made to ICZN
to use its Plenary Power to suppress the senior homo-
nym (ICZN, 1985: Article 79) (see the comments on B.
muscorum). However, the consequence of this action

P.H. WILLIAMS

would be that pyrenaeus (Lepeletier) would no longer
be available for a subspecies of B. rupestris.

Bombus (Pr.) biroi Vogt
biroi Vogt, 1911:51, examined
nursei Friese, 1918:84, examined
Yagnatus Skorikov, 1933b:248, examined, not of Skorikov,
1912:97 (= B. monticola Smith)
?kotzschi Reinig, 1940:227, examined

@ TAXONOMIC sTATUS. Several of these nominal
taxa have been treated as separate species. However,
aside from differences in colour pattern, they are
closely similar in morphology with a range of varia-
tion (Williams, 1991). Until more evidence to the
contrary is available from critical studies of patterns of
variation, I shall treat them as parts of a single variable
species.

Bombus (Pr.) frigidus Smith
frigidus Smith, 1854:399, examined

Bombus (Pr.) sandersoni Franklin
sandersoni Franklin, 1913:353

Bombus (Pr.) flavifrons Cresson
pleuralis Nylander, 1848:231, examined
flavifrons Cresson, 1863:105, new synonym

@ TAXONOMIC STATUS. B. pleuralis and B.
flavifrons were regarded as separate species by Franklin
(1913), and Poole (1996) also lists them as separate
species, without explanation. In my opinion, the
lectotype of B. pleuralis designated by Milliron
(1960:95) is an individual of the dark form of B.
flavifrons (see descriptions of variation by e.g. Stephen,
1957; Thorp et al., 1983). See also the comments on B.
mixtus.

O NOMENCLATURE. B. pleuralisis the oldest avail-
able name for this species.

€ APPLICATION TOICZN. Although B. pleuralis is
the oldest available name for the present interpretation
of this species, the name B. flavifrons has been in
common use for the species since 1947 (e.g. Stephen,
1957; Thorp, 1969, 1970; Plowright & Stephen, 1973;
Macior, 1975; Sakagami, 1976; Hurd, 1979; Plowright
& Owen, 1980; Thorp et al., 1983; Laverty & Harder,
1988). I know of no publications using the name B.
pleuralis since 1947, apart from the list by Poole
(1996). It is suggested that, in the interests of stability
(ICZN, 1985: Article 23b), an application be made to
ICZN to use its Plenary Power to suppress the unused
senior synonym (ICZN, 1985: Article 79) (see the

CHECKLIST OF BUMBLE BEES

comments on B. muscorum). However, the conse-
quence of this action would be that pleuralis would no
longer be available for a species or for a subspecies of
B. flavifrons.

Bombus (Pr.) centralis Cresson
centralis Cresson, 1864:41

Bombus (Pr.) vandykei (Frison)
vandykei (Frison, 1927:375 [Bremus])
cascadensis (Milliron, 1970a:382 [Pyrobombus])

Bombus (Pr.) caliginosus (Frison)
caliginosus (Frison, 1927:376 [Bremus])

Bombus (Pr.) vagans Smith
vagans Smith, 1854:399, examined

Bombus (Pr.) mixtus Cresson®
Praticola Kirby, 1837:274
mixtus Cresson, 1878:186, not of Kriechbaumer, 1870:160
(= B. maxillosus Klug), new synonym

@ TAXONOMICSTATUS. The identity of B. praticola
has remained uncertain (e.g. Cresson, 1863; Franklin,
1913). Recently, Poole (1996) has listed B. praticola,
B. mixtus and B. flavifrons as separate species without
explanation.

Although I know of no type material, Kirby pro-
vided a description of B. praticola from northern
Canada (latitude 65° North) with a colour pattern
(including anterior half of abdomen yellow, posterior
ferruginous) that for individuals from this area is most
likely to be conspecific either with B. mixtus (some
individuals have few black hairs on gastral terga II-
III), or with B. flavifrons (which has terga V—VI black,
although this is not always apparent from the dorsal
view). In his original description of B. flavifrons,
Cresson (1863) conceded that this might be the same
species as Kirby’s B. praticola, and he went on to write
(p. 106) that he had not yet identified B. praticola.
Franklin (1913:371) wrote that he had ‘been unable to
decide whether the original description of B. praticolus
[sic] referred to this species [B. flavifrons] or to the
colour variant of pleuralis [intermediate colour pat-
terns between B. flavifrons and B. pleuralis].’ Milliron
(1971:42) subsequently listed Pyrobombus praticola
flavifrons (Cr.) as a member of his ‘Praticola Group’.

However, here I follow R. Miller (in Jitt.), who
believes that the original material was more likely to
have been of the species that has come to be known as
B. mixtus. See the comments on B. flavifrons.

O NOMENCLATURE. B. praticola is probably the
oldest available name for this species. Any remaining

127

confusion could be resolved by the designation of an
appropriate neotype (e.g. see the comments on B.
subterraneus).

© APPLICATION TOICZN. Although B. praticola is
probably the oldest available name for this species, the
name B. mixtus has been in common use for the
species since 1947 (e.g. Stephen, 1957; Thorp, 1970;
Plowright & Stephen, 1973; K. W. Richards, 1973;
Macior, 1975; Sakagami, 1976; Hurd, 1979; Plowright
& Owen, 1980; Thorp et al., 1983; Laverty & Harder,
1988; Macfarlane et al., 1994). It is suggested that, in
the interests of stability, an application be made to
ICZN to use its Plenary Power to suppress the senior
synonym and homonym (ICZN, 1985: Article 79) (see
the comments on B. muscorum). However, the conse-
quence of this action would be that mixtus (Kriech-
baumer) would no longer be available for a subspecies
of B. maxillosus.

Bombus (Pr.) sitkensis Nylander
Sitkensis Nylander, 1848:235

Bombus (Pr.) melanopygus Nylander
melanopyge Nylander, 1848:236
Edwardsii Cresson, 1878:184
melampygus Handlirsch, 1888:231, unjustified emenda-
tion
[melanopygus Viereck, 1904:99, incorrect subsequent spell-
ing]
melanopygus Franklin, 1913:334, justified emendation
@ TAXONOMIC STATUS. B. melanopygus and B.
edwardsii were shown by Owen & Plowright (1980) to
differ principally by a single allele controlling the
colour of the pubescence on gastral terga II-III. There
can be little doubt that they are conspecific.

Bombus (Pr.) lapponicus (Fabricius)
lapponica (Fabricius, 1793:318 [Apis])
?sylvicola Kirby, 1837:272
zhaosu Wang, 1985:162, examined, new synonym

@ TAXONOMIC sTATUS. B. sylvicola is morpho-
logically closely similar to B. lapponicus, and it has
been suggested repeatedly that they may be conspecific
(e.g. Sladen, 1919; Skorikoy, 1922a, 1937; Pittioni,
1942, 1943; Thorp, 1962; Thorp et al., 1983).

B. zhaosu was described from material from
Xinjiang, China, and is closely similar toB. lapponicus.

These three nominal taxa have been treated as sepa-
rate species. However, aside from differences in colour
pattern, they are closely similar in morphology. Until
more evidence to the contrary is available from critical
studies of patterns of variation, I shall treat them as
parts of a single variable species.

128

?Bombus (Pr.) monticola Smith

montanus Smith, 1844:549, not of Lepeletier, 1836:463 (=
B. ruderarius (Miiller))

monticola Smith, 1849:lx, replacement name formontanus
Smith, 1844:549

lugubris Sparre-Schneider, 1909:155, not of Kriechbaumer,
1870:159 (= B. maxillosus (Klug))

scandinavicus Friese, 1912:684, replacement name for
lugubris Sparre-Schneider, 1909:255

@ TAXONOMIC STATUS. B. scandinavicus (= B.
monticola) and B. lapponicus are names that were
applied initially to two colour forms in Scandinavia.

Lgken (1973) reported that these two taxa overlap
narrowly in distribution and intergrade. However, they
have been found to differ consistently (for samples
analysed) in the composition of cephalic secretions
(Bergstr6m & Svensson, 1973; Svensson & Bergstré6m,
1977). Svensson (1973, 1979) also described subtle
differences in morphological characters, although other
morphological studies by Lgken (1973) and Pekkarinen
(1979) found no distinct differences. Pekkarinen (1982,
in litt.) now believes that they are separate species.

It remains possible that there is a hybrid zone where
the colour forms intergrade, with some gene flow. In this
case, depending on the species concept embraced, these
taxa might be considered conspecific (see the com-
ments on B. ruderatus). Until further evidence is avail-
able, I shall continue to treat them as separate species.

COMMENT. Until the twentieth century B. monticola
was not known from Ireland, where it is now estab-
lished (see references in Alford, 1975, 1980) (see
comments on B. pratorum).

Bombus (Pr.) bimaculatus Cresson
bimaculatus Cresson, 1863:92

Bombus (Pr.) impatiens Cresson
impatiens Cresson, 1863:90

Bombus (Pr.) vosnesenskii Radoszkowski
Vosnesenskii Radoszkowski, 1862:589

Bombus (Pr.) bifarius Cresson
bifarius Cresson, 1878:185
andamanus Gribodo, 1882:268, examined
fernaldi Franklin, 1911:157, not a replacement name

@ TAXONOMIC sTaTUS. B. andamanus was de-
scribed as originating from ‘Andaman’ (= Andaman
Islands, Indian Ocean), but appears to be a mislabelled
queen of B. bifarius from western North America
(Tkalcti, 1966). I have examined this specimen and
agree with this identification (i.e. contrary to Richards,
1929, itis nota species of the subgenus Bombus s. str.).

P.H. WILLIAMS

Bombus (Pr.) huntii Greene
Huntii Greene, 1860:172

Bombus (Pr.) ternarius Say
ternarius Say, 1837:414
ornatus Smith, 1854:398, examined

Bombus (Pr.) ephippiatus Say
ephippiatus Say, 1837:414
formosus Smith, 1854:403, examined
lateralis Smith, 1879:134, examined
?wilmatte Cockerell, 1912:21, examined
?alboniger Franklin, 1915:409, examined
folsomi (Frison, 1923:322 [Bremus]) examined

@ TAXONOMIC STATUS. B. wilmattae, B.
alboniger and B. ephippiatus have been regarded
both as conspecific and as separate species. Recently,
B. wilmattae and B. ephippiatus were regarded as
separate species by Labougle ef al. (1985) and
Labougle (1990), who described diagnostic charac-
ters of colour pattern and morphology. However, D.
Yanega (in litt.) and G. Chavarria (pers. com.) believe
that all of these nominal taxa are part of the wide-
spread and variable B. ephippiatus. Until more
evidence to the contrary is available from critical
studies of patterns of variation, I shall treat them as
parts of a single variable species.

B. folsomi was described as originating from “Kina
Bala / N. Borneo’ (= Gunung Kinabalu, Sabah), but
appears to be a mislabelled queen of B. ephippiatus,
probably from Costa Rica or Panama (Starr, 1989). I
have examined this specimen and agree with this
identification.

Subgenus FESTIVOBOMBUS Tkalcii
[Atrocinctob.{ombus | Skorikov, 1933b:244, published with-
out description]
Pyrobombus (Festivobombus) Tkalcu, 1972:26, type-spe-
cies Bombus festivus Smith by original designation
Bombus (Festivobombus) Williams, 1985b:240

@ TAXONOMIC STATUS. Richards (1968) treated
B. atrocinctus (= B. festivus) as a species of the

CHECKLIST OF BUMBLE BEES

subgenus Pyrobombus, even though this required nu-
merous exceptions in the diagnosis. I have followed
Tkalcii(1972, 1974) in treating Festivobombus and
Pyrobombus as separate subgenera (Williams, 1991),
because together they do not form a monophyletic
group (Williams, 1995).

Bombus (Fs.) festivus Smith
festivus Smith, 1861:152, examined
atrocinctus Smith in Horne, 1870:193, examined
terminalis Smith in Horne, 1870:193, examined

Subgenus RUFIPEDIBOMBUS Skorikov
Rufipedibombus Skorikov, 1922a:156, type-species
Bombus rufipes Lepeletier by monotypy
Bombus (Rufipedibombus) Richards, 1930:638
Bombus (Rufipedobombus) Kruseman, 1952:102, unjusti-
fied emendation

Bombus (Rf) rufipes Lepeletier
rufipes Lepeletier, 1836:473
richardsi (Frison, 1930:6 |Bremus])

Bombus (Rf.) eximius Smith
eximius Smith, 1852b:47, examined
latissimus Friese, 1910:405

Subgenus PRESSIBOMBUS Frison
Bremus (Pressibombus) Frison, 1935:342, type-species

129

Bremus pressus Frison (= Bombus pressus (Frison)) by
original designation

Bombus (Pressobombus) Kruseman, 1952:102, unjusti-
fied emendation

Bombus (Pressibombus) Richards, 1968:217

Bombus (Pe.) pressus (Frison)
pressus (Frison, 1935:342 [Bremus])

Subgenus BOMBUS in the strict sense

Bombus (Leucobombus) Dalla Torre, 1880:40, type-spe-
cies Apis terrestris Linnaeus (= Bombus terrestris
(Linnaeus)) by subsequent designation of Sandhouse,
1943:564

Bombus (Terrestribombus) Vogt, 1911:55, type-species
Apis terrestris Linnaeus (= Bombus terrestris
(Linnaeus)) by subsequent designation of Frison,
1927:67

Bombus (Bo.) sporadicus Nylander
sporadicus Nylander, 1848:233
malaisei Bischoff, 1930:4

Bombus (Bo.) tunicatus Smith
tunicatus Smith, 1852a:43, examined
vallestris Smith, 1878:8
gilgitensis Cockerell, 1905:223, examined

Bombus (Bo.) franklini (Frison)
franklini (Frison, 1921:147 [Bremus])

@ TAXONOMIC STATUS. This species has been
treated as conspecific with B. occidentalis (= B.
terricola) by Milliron (1971), but has since been shown
to be very distinct in morphology by Plowright &
Stephen (1980) and Williams (1991), and in enzyme
mobilities by Scholl, Thorp & Obrecht (1992).

COMMENT. 8B. franklini has one of the narrowest
distributions of any bumble bee species world-wide.
All recent specimens have been collected within a 60
mile (38 km) radius of Grants Pass, Oregon (Thorp,
1970; Thorp et al., 1983).

130

Bombus (Bo.) affinis Cresson
affinis Cresson, 1863:103

Bombus (Bo.) ignitus Smith
ignitus Smith, 1869:207, examined
terminalis Smith, 1873:206, examined, not of Smith in
Horne, 1870:193 (= B. festivus Smith)
japonicus Dalla Torre, 1890:139, replacement name for
terminalis Smith, 1873:206

Bombus (Bo.) terrestris (Linnaeus)
terreftris (Linnaeus, 1758:578 [Apis])
Audax (Harris, 1776:130 [Apis]) not of Harris, 1776:137
(= Anthophora sp.)
canariensis Pérez, 1895:191
maderensis Erlandsson, 1979:191

@ TAXONOMICSTATUS. B. terrestris, B. maderensis
and B. canariensis have been regarded both as
conspecific and as separate species.

Erlandsson (1979) argued that the dark individuals
from the Canary Islands, previously placed within B.
terrestris by for example Kriiger (1954, 1956), are a
separate species, B. canariensis. Erlandsson also ar-
gued that individuals from the island of Madeira,
previously placed within B. terrestris by Bischoff
(1937), are a separate species, B. maderensis. In both
cases the morphological characters used to support
these distinctions are not strongly divergent from the
broad variation within B. terrestris in the broad sense.
Rasmont (1984) regarded these three taxa as separate
species, but Pekkarinen & Kaarnama (1994) treated
them as conspecific.

Recent work by Estoup et al. (1996) has found that
although European mainland populations do not vary
significantly among themselves in mitochondrial
genes, all island populations studied (from six Medi-
terranean islands in addition to B. canariensis) show
significant differences from the mainland populations.

Consequently, viewing these three nominal taxa as
separate species may be one interpretation, but this
appears to depend on adopting a species concept that
admits little colour, morphological or genetic varia-
tion within a species and regards current geographical
isolation as highly indicative. I prefer to regard these
taxa as conspecific until further evidence is available.

O NOMENCLATURE. Day (1979) described how
none of the admissable syntypes of A. terrestris
Linnaeus is in agreement with the current usage of the
name.

To reaffirm the traditional usage of this particularly
widely used name, a case was made to ICZN by Loken
et al. (1994). This sought an Opinion from ICZN
(ICZN, 1996) that set aside, by use of its Plenary
Power (ICZN, 1985: Articles 78b, 79), the lectotype
designation for A. terrestris by Day from application

P.H. WILLIAMS

of the Code (ICZN, 1985), and then designated a
neotype (ICZN, 1996: 64) to conserve the traditional
usage of the name for even the narrowest concept of
the taxon (ICZN, 1985: Article 75).

COMMENT. This species has been introduced into
New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr,
1995) (see the comments onB. hortorum, B. ruderatus,
and B. subterraneus), Tasmania (Cardale, 1993), and
Japan (I. Washitani, in /itt.). It was also apparently
introduced into mainland Australia (New South Wales)
without persisting (W. Froggatt in Franklin, 1913).

Bombus (Bo.) hypocrita Pérez
hypocrita Pérez, 1905:30

Bombus (Bo.) patagiatus Nylander
patagiatus Nylander, 1848:234
vasilievi Skorikov, 1913:172

Bombus (Bo.) lucorum (Linnaeus)

lucorum (Linnaeus, 1761:425 [Apis]), examined

2cryptarum (Fabricius, 1775:379 [Apis])

?modestus Cresson, 1863:99, not of Eversmann, 1852:134
(= B. modestus Eversmann)

?moderatus Cresson, 1863:109, replacement name for
modestus Cresson, 1863:99

monozonus Friese, 1909:674

?magnus Vogt, 1911:56

2jacobsoni Skorikov, 1912:610, examined

?burjaeticus Kriiger, 1954:277

?florilegus Panfilov, 1956:1334

?reinigi Tkalct, 1974b:322, examined

@ TAXONOMIC sTaTUS. These bees have received
particularly close attention by authors describing the
minutiae of colour variation, using at least 186 classi-
cal names (see the introduction). At least some of these
nominal taxa have been regarded as separate species
by some authors (e.g. Rasmont, 1983, 1984, 1988;
Scholl & Obrecht, 1983; Scholl et al., 1990; Scholl,
Thorp & Obrecht, 1992; Rasmont et al., 1995; Amiet,
1996; Ozbek, 1997; Pamilo et al., 1997). In contrast,
B. cryptarum and the North American B. moderatus
have recently been treated as conspecific with B.
lucorum by Poole (1996).

There are conflicting patterns of variation among
some characters of these taxa, which are not fully
understood (Pekkarinen, 1979; Pamilo ef al., 1984;
Amiet, 1996; Pamilo et al., 1997). As far as I can tell
from the evidence available at present, separation of
the taxa for mapping is still not reliable, at least in
some areas of their distribution, and particularly in
Central and Eastern Asia, where there appears to be a
broad range of variation with some intergradation of
character combinations (Williams, 1991). Therefore,

CHECKLIST OF BUMBLE BEES

because complete mapping of separate taxa is not yet
possible for me, B. /ucorum is interpreted here in the
broadest sense, to include a complex of similar taxa.
However, these taxa require more critical work to
clarify population patterns of variation and inherit-
ance, even in relatively well known areas such as
Britain.

O NOMENCLATURE. When Day (1979) came to
fix the application of A. terrestris Linnaeus (see the
comments on B. terrestris), he had no reason to believe
that Linnaeus had not described this taxon from the
syntype specimen that was subsequently described as
the lectotype (=A. cryptarum Fabricius, see Rasmont,
1988:52, ?= B. lucorum (Linnaeus)). This action
brought B. Jucorum (Linnaeus) into subjective junior
synonymy with B. terrestris (Linnaeus).

To reaffirm the traditional usage of B. terrestris and
B. lucorum, a case was made to ICZN by Loken et al.
(1994). This sought an Opinion from ICZN (ICZN,
1996) that set aside, by use of its Plenary Power
(ICZN, 1985: Articles 78b, 79), the lectotype designa-
tion for A. terrestris by Day from application of the
Code (ICZN, 1985), and then designated a neotype
(ICZN, 1996: 64) to conserve the traditional usage of
B. terrestris and B. lucorum (ICZN, 1985: Article 75).

COMMENT. This species occurs in Iceland, where it
has probably been introduced (Prys-Jones et al., 1981)
(see the comments on B. hortorum).

Bombus (Bo.) terricola Kirby
Terricola Kirby, 1837:273
occidentalis Greene, 1858:12

@ TAXONOMIC sTaTUS. B. terricola and B.
occidentalis have been regarded both as conspecific
(e.g. Milliron, 1971; Poole, 1996) and as separate
species (e.g. Franklin, 1913 [but see p. 239]; Stephen,
1957; Thorp et al., 1983; Scholl et al., 1990).

Many specimens from the north west of North
America show a reduction in the extent of the yellow
bands on gastral terga I and II, with an expansion of
the pale pubescence on tergum IV, and so appear to be
intermediate or recombinant individuals. Indeed,
Stephen’s (1957:74) figure 4 shows several patterns
that could represent a continuum of variation between
the two forms. Furthermore, Thorp ef al. (1983: fig.
140a) illustrate individuals of ‘B. occidentalis’ from
California that look very similar to eastern B. terricola.
In view of this, Stephen’s conclusion that there is no
intergradation may result from adopting colour crite-
ria (identifying B. terricola in the strict sense either by
completely black pubescence of female terga V—VI [p.
15] and male tergum IV [p. 19], or by completely
yellow pubescence of tergum II [pp. 19, 71], two
character states that do not always occur together, even

131

in the east) that could be considered as essentially
arbitrary points on a continuum (see the comments on
B. fervidus).

In view of the existence of apparent intermediates
between these nominal taxa in at least part of their
range, they are treated here as likely to be conspecific.
Until more evidence to the contrary is available from
critical studies of patterns of variation, I shall treat
them as parts of a single variable species.

Subgenus CULLUMANOBOMBUS Vogt

Bombus (Cullumanobombus) Vogt, 1911:57, type-species
Apis cullumana Kirby (= Bombus cullumanus (Kirby))
by subsequent designation of Frison, 1927:66

Bremus (Rufocinctobombus) Frison, 1927:78, type-spe-
cies Bombus rufocinctus Cresson by monotypy

Cullumanibombus Skorikov, 1938a:145, unjustified emen-
dation

Bombus (Cu.) rufocinctus Cresson
rufo-cinctus Cresson, 1863:106

Bombus (Cu.) cullumanus (Kirby)
Cullumana (Kirby, 1802:359 [Apis]) examined
serrisquama Morawitz, 1888:224
Silantjewi Morawitz, 1892:132
apollineus Skorikovy, 19106:412

@ TAXONOMIC sTATUS. Several of these nominal
taxa have been treated as separate species. However,
aside from differences in colour pattern, they are
closely similar in morphology (Panfilov, 1951).
Rasmont (1988) has drawn attention to the co-occur-
rence of the white-banded B. apollineus with the
yellow-banded B. serrisquama in northern Iran, ap-
parently without intermediate individuals. But by
analogy, it is possible that this colour difference
could be the effect of a single allele for pigment (cf.
B. melanopygus, see also the comments on B.
keriensis). Until more evidence to the contrary is
available from critical studies of patterns of varia-
tion, I shall treat them as parts of a single variable
species.

Bombus (Cu.) unicus Morawitz
unicus Morawitz, 1883:235
controversus Skarikov, 1910b: 411

@ TAXONOMIC STATUS. B. unicus is similar to B.
cullumanus and could possibly be conspecific. How-
ever, the male genitalia appear to be more distinct
(Panfilov, 1951) from those of the other taxa tradition-
ally considered subspecies of B. cullumanus.

Bombus (Cu.) semenoviellus Skorikov
semenoviellus Skorikoy, 1910b:410

Subgenus OBERTOBOMBUS Reinig
Bombus (Obertobombus) Reinig, 1930:107, type-species
Bombus oberti Morawitz by monotypy
[Obertibombus Skorikoy, 1931:239, incorrect subsequent
spelling]
Bombus (Obertibombus) Reinig, 1934:167, unjustified
emendation

@ TAXONOMIC STATUS. Richards (1968) treated
Obertobombus as a synonym of the subgenus
Sibiricobombus. I have recognised Obertobombus and
Sibiricobombus as separate subgenera, because to-
gether they do not form a monophyletic group
(Williams, 1995).

Bombus (Ob.) morawitzi Radoszkowski
Morawitzi Radoszkowski, 1876:101, examined
hydrophthalmus Morawitz, 1883:240, examined

Bombus (Ob.) oberti Morawitz
Oberti Morawitz, 1883:238, examined
Semenovi Morawitz, 1886:198, examined
xionglaris Wang, 1982:432, examined, new synonym
duanjiaoris Wang, 1982:444, examined
zhadaensis Wang, 1982:444, examined, new synonym

@ TAXONOMIC STATUS. B. xionglaris and B. zha-
daensis are closely similar to B. oberti in morphology
and in colour pattern. These bees occur at high alti-
tudes and are not common in collections (Williams,
1991). However, I know of no reason why these

P.H. WILLIAMS

nominal taxa should not be considered conspecific.

Subgenus MELANOBOMBUS Dalla Torre

Bombus (Melanobombus) Dalla Torre, 1880:40, type-spe-
cies Apis lapidaria Linnaeus (= Bombus lapidarius
(Linnaeus)) by subsequent designation of Sandhouse,
1943:569

Bombus (Lapidariobombus) Vogt, 1911:58, type-species
Apis lapidaria Linnaeus (= Bombus lapidarius
(Linnaeus)) by subsequent designation of Sandhouse,
1943:562

Kozlovibombus Skorikov, 1922a:152, type-species Bombus
kozlovi Skorikov, 1910b (=Bombus keriensis Morawitz)
in the sense of Skorikov, 1922a (based on males =
Bombus pyrosoma Morawitz, a misidentification, see
Reinig, 1934:169, requiring designation by ICZN), by
subsequent fixation of Sandhouse, 1943:561

Bombus (Kozlowibombus) Bischoff, 1936:10, unjustified
emendation

Lapidariibombus Skorikov, 1938a:145, unjustified emen-
dation

?Bombus (Tanguticobombus) Pittioni, 1939d:201, type-
species Bombus tanguticus Morawitz by original
designation (provisional synonym)

[Bombus (Lapedariobombus )Esmaili & Rastegar, 1974:52,
incorrect subsequent spelling]

Bombus (MI.) tanguticus Morawitz
tanguticus Morawitz, 1886:200

@ TAXONOMIC STATUS. Queens of B. tanguticus
are morphologicaly very distinctive (discussed in
Williams, 1991), so much so that Pittioni (1939d)
considered the species warranted a subgenus of its
own. The male remains apparently unknown (the spe-
cies occurs at high altitudes in Tibet [= Xizang] and is
very rare in collections), so that its precise relation-
ships are difficult to resolve at present and a separate
subgenus seems premature.

Bombus (M1.) simillimus Smith
simillimis Smith, 1852b:48, examined
[similis Smith, 1854:403, incorrect subsequent spelling]
[simillimus Dalla Torre, 1896:548, incorrect subsequent
spelling]
grossiventris Friese, 1931:303, examined

CHECKLIST OF BUMBLE BEES

oculatus (Frison, 1933:335 [Bremus]) examined
tonsus (Skorikov, 1933b:248 [Sibiricobombus]) examined
simillimus Williams, 1991:99, justified emendation

Bombus (ML.) richardsiellus (Tkalcii)
richardsiellus (Tkalci, 1968a:42 [Pyrobombus]) exam-
ined

Bombus (ML.) pyrosoma Morawitz
pyrosoma Morawitz, 1890:349, examined
pyrrhosoma Dalla Torre, 1896:544, unjustified emenda-
tion
wutaishanensis (Tkalcit, 1968a:39 [Pyrobombus]) exam-
ined

@ TAXONOMICSTATUS. B. pyrosomahas been con-
sidered conspecific with B. friseanus (Bischoff, 1936)
and has been considered conspecific with B.
formosellus, B. friseanus and B. flavothoracicus (= B.
miniatus) (Williams, 1991). From a preliminary analy-
sis of colour variation, S.-f. Wang and J. Yao report (in
litt.) that these taxa appear to remain discrete and are
likely to be separate species. More evidence is awaited.

?Bombus (MI.) formosellus (Frison)

formosellus (Frison, 1934:163 [Bremus]) examined

@ TAXONOMIC status. B. formosellus has been
considered conspecific with B. pyrosoma, B. friseanus
and B. flavothoracicus (= B. miniatus) (Williams,
1991), as a disjunct peripheral population on Taiwan.
From a preliminary analysis of colour variation, S.-f.
Wang and J. Yao report (in /itt.) that these taxa appear
to remain discrete and are likely to be separate species.
More evidence is awaited.

?Bombus (ML.) friseanus Skorikov
friseanus Skorikov, 1933a:62, examined
hénei Bischoff, 1936:10, examined

@ TAXONOMIC STATUS. _ B. friseanus has been con-
sidered conspecific with B. pyrosoma (Bischoff, 1936;
Tkalcti, 1961b; Sakagami, 1972) and has been consid-
ered conspecific with B. pyrosoma, B. formosellus and
B. flavothoracicus (= B. miniatus) (Williams, 1991).
From a preliminary analysis of colour variation, S.-f.
Wang and J. Yao report (in /itt.) that these taxa appear
to remain discrete and are likely to be separate species.
More evidence is awaited.

?Bombus (ML.) miniatus Bingham
flavothoracicus Bingham, 1897:552, examined, not of
Hoffer, 1889:49 (= B. campestris (Panzer))
miniatus Bingham, 1897:553, examined
eurythorax Wang, 1982:435, examined, new synonym

133
stenothorax Wang, 1982:439, examined, new synonym

@ TAXONOMIC STATUS. B. miniatus has been con-
sidered conspecific with B. pyrosoma, B. formosellus
and B. friseanus (Williams, 1991).

Evidence of intermediates between B. miniatus and
B. friseanus is not strong, but not least because so little
material is available from where these taxa occur in
close proximity in the eastern Himalaya The few
workers and males from this area that I have seen are
difficult to assign to either taxon with any confidence,
although the queens are closer to the colour pattern of
B. miniatus (Williams, 1991). More evidence is
awaited.

B. eurythorax and B. stenothorax are closely similar
in morphology and colour pattern to B. miniatus. I
know of no reason why these nominal taxa should not
be considered conspecific.

O NOMENCLATURE. With Psithyrus regarded as
being a subgenus of the genus Bombus, B.

flavothoracicus Bingham (1897) becomes a junior

secondary homonym in Bombus of Psithyrus camp-
estris var. flavothoracicus Hoffer (1889) (deemed to
be subspecific, see ICZN, 1985: Article 45g(ii)), and
therefore the name B. flavothoracicus Bingham is
invalid (ICZN, 1985: Article 57c).

For this species, the oldest available name is B.
miniatus, which becomes the valid name. The only
subsequent publications of which I am aware that use
the name B. flavothoracicus for this taxon as a species
are by Tkalcti (1974b), Wang (1982) and Macior (1990),
so this change of valid name is not a serious disruption
of common usage.

Bombus (M1.) rufofasciatus Smith

rufo-fasciatus Smith, 1852b:48, examined

Prshewalskyi Morawitz, 1880:342

rufocinctus Morawitz, 1880:343, examined, not of Cresson,
1863:106 (= B. rufocinctus Cresson)

chinensis Dalla Torre, 1890{June 25]:139, replacement
name for rufocinctus Morawitz, 1880:343; not of
Morawitz, 1890[April 30]:352 (= B. chinensis
(Morawitz))

waterstoni Richards, 1934:88, examined

Bombus (MI.) ladakhensis Richards
ladakhensis Richards, 1928a:336, examined, not
infrasubspecific after Tkalcti, 1974b:335
phariensis Richards, 1930:642, examined, not
infrasubspecific after Tkalcti, 1974b:336
variopictus Skorikov, 1933b:248, examined
reticulatus Bischoff, 1936:7, examined

Bombus (MI.) semenovianus (Skorikov)
semenovianus (Skorikov, 1914a:127 [Lapidariobombus])
examined

134

Bombus (MIL.) incertus Morawitz
incertus Morawitz, 1881:229

Bombus (ML.) lapidarius (Linnaeus)
Lapidaria (Linnaeus, 1758:579 [Apis]) examined
Strenuus (Harris, 1776:131 [Apis])
eriophorus Klug, 1807:265, examined
caucasicus Radoszkowski, 1859:482, examined

Bombus (ML.) keriensis Morawitz

keriensis Morawitz, 1886:199, examined

separandus Vogt, 1909:61, examined

kohli Vogt, 1909:61, examined, not of Cockerell, 1906:75
(= B. morio (Swederus))

kozlovi Skorikoy, 1910b:413, replacement name for kohli
Vogt, 1909:61

tenellus Friese, 1913:86

[alagesianus (Skorikov, 1922a:152 [Lapidariobombus])
published without description]

alagesianus Reinig, 1930:89

richardsi Reinig, 1935:341, not of Frison, 1930:6 (= B.
rufipes Lepeletier)

tibetensis Wang, 1982:439, replacement name forrichardsi
Reinig, 1935:341

trilineatus Wang, 1982:441, examined, new synonym

@ TAXONOMIC STATUS. Several of these nominal
taxa have been treated as separate species (e.g.
Skorikov, 1931), although B. keriensis has also long
been considered a broadly-distributed and variable
species, including both yellow-banded and white-
banded individuals throughout much of its range
(Reinig, 1935, 1939; Williams, 1991; Fig. 9).

B. trilineatus is morphologically closely similar to
B. keriensis. | know of no reason why these nominal
taxa should not be considered conspecific.

Bombus (ML1.) sichelii Radoszkowski

Sichelii Radoszkowski, 1859:481, examined

[Sicheli Radoszkowski, 1877b:213, incorrect subsequent
spelling]

tenuifasciatus Vogt, 1909:49, not of Vogt, 1909:49 (= B.
pyrenaeus Pérez) after Tkalcii, 1973:266

chinganicus Reinig, 1936:6, not of Reinig, 1936:8 (?= B.
bohemicus Seidl)

erzurumensis (Ozbek, 1990:209 [Pyrobombus]) examined,
new synonym

@ TAXONOMIC sTaTUS. Until recently, the white-
banded form of B. sichelii has been known from west
of the Caspian Sea only from the Caucasus (Reinig,
1935). Now that B. erzurumensis (morphologically
closely similar to B. sicheli-*

i and with white bands) has been described from
Turkey, it could be interpreted as another white-banded,
western colour form. By analogy (cf. comments on B.
melanopygus), the difference in colour could be the

P.H. WILLIAMS

effect of a single allele for pigment.

Until more evidence to the contrary is available
from critical studies of patterns of variation, I shall
treat them as parts of a single variable species.

Subgenus SIBIRICOBOMBUS Vogt

Bombus (Sibiricobombus) Vogt, 1911:60, type-speciesApis
sibirica Fabricius (= Bombus sibiricus (Fabricius)) by
subsequent designation of Sandhouse, 1943:599

Sibiricibombus Skorikov, 1938a:145, unjustified emenda-
tion

[Bombus (Sibericobombus) Kruseman, 1952:101, incor-
rect subsequent spelling]

Bombus (Sb.) sibiricus (Fabricius)
fibirica (Fabricius, 1781:478 [Apis]) examined
flaviventris Friese, 1905:514, examined, new synonym
ochrobasis Richards, 1930:655, examined, new synonym

@ TAXONOMIC sTaTuUS. B. sibiricus and B. flavi-
ventris have been regarded as separate species. Females
of B. flaviventris are morphologically closely similar
to those of B. sibiricus, but differ in having the orange
pubescence dorsally between the wing bases and on
gastral terga IV—VI replaced with black. S.-f. Wang
and J. Yao have kindly shown me the male of B.
flaviventris, which 1s closely similar in its genitalia to
B. sibiricus.

B. ochrobasis appears to differ from B. flaviventris
only in the lighter hue of the yellow pubescence of B.
ochrobasis.

At present I know of no good biological reason why
these three nominal taxa should not be regarded as
conspecific. More evidence is awaited.

COMMENT. B. flaviventris has long been placed in
the subgenus Subterraneobombus (e.g. Skorikov,
1922a; Richards, 1930, 1968), although the characters
of the females (Williams, 1991) and the males (Wang
& Yao, unpublished) agree with the species of the
subgenus Sibiricobombus.

Bombus (Sb.) obtusus Richards
obtusus Richards, 1951:196, examined

CHECKLIST OF BUMBLE BEES

Bombus (Sb.) asiaticus Morawitz
asiatica Morawitz in Fedtschenko, 1875:4, examined
longiceps Smith, 1878:8
Regeli Morawitz, 1880:337, examined
regelii Dalla Torre, 1896:544, unjustified emendation
[miniatocaudatus Vogt, 1909:50, infrasubspecific]
miniatocaudatus Vogt, 1911:61, examined, not of Vogt,

1909:56 (= B. soroeensis (Fabricius))

heicens Wang, 1982:430, examined, new synonym
huangcens Wang, 1982:430, examined, new synonym
flavicollis Wang, 1985:163, examined, new synonym
baichengensis Wang, 1985:164, examined, new synonym

@ TAXONOMIC STATUS. Several of these nominal
taxa have been treated as separate species.

B. heicens, B. huangcens, B. flavicollis and B.
baichengensis are morphologically closely similar to
B. .asiaticus and differ onlyin details of the colour
pattern. In the case of the yellow unbanded colour
form and the grey banded colour form in Kashmir
(Fig. 12), there is evidence of interbreeding, with
many recombinant individuals in some localities
(Williams, 1991).

Aside from differences in colour pattern, these taxa
are similar in morphology with a range of variation
(Williams, 1991). Until more evidence to the contrary
is available from critical studies of patterns of varia-
tion, I shall treat them as parts of a single variable
species.

Bombus (Sb.) niveatus Kriechbaumer
niveatus Kriechbaumer, 1870:158
?vorticosus Gerstaecker, 1872:290, examined (provisional
synonym)

@ TAXONOMIC STATUS. B. niveatus and B. vorti-
cosus have been regarded both as conspecific
(Schmiedeknecht, 1883; Handlirsch, 1888; DallaTorre,
1896; Schulz, 1906) and, more recently, as separate
species (e.g. Skorikov, 1922a; Pittioni, 1938; Tkalcii,
1969; Reinig, 1981; Rasmont, 1983).

As far as I am aware, the white-banded B. niveatus
occurs only within the broader distributional bounds
of the yellow-banded B. vorticosus (within its “extent
of occurrence’ in the sense of Gaston, 1994). Although
they differ in the colour of the pale pubescence (Pittioni,
1939a), they are closely similar in morphology
(Williams, 1991; Baker, 19965). Pittioni (1938) and
Baker (1996b) report that they occur at different alti-
tudes, without intermediate colour forms. However,
the significance of this is unclear, because Baker
(1996b) notes that the white-banded B. niveatus co-
occurs with other bumble bees (B apollineus (= B.
cullumanus), B. simulatilis (= B. ruderarius)) that also
show strong convergences in these areas towards the
white-banded colour pattern, while elsewhere they are
more broadly distributed in yellow-banded colour
forms. By analogy with other species (cf. comments

135

on B. melanopygus, B. keriensis), the difference in
colour could be the effect of a single pair of alleles for
pigment. It is suspicious that both colour forms show
identical variation in the extent of pale fringes to the
pubescence on the posterior of tergum ILI.

Until more evidence for differences between these
nominal taxa other than colour is available from criti-
cal studies of patterns of variation, I shall treat them as
parts of a single variable species.

Bombus (Sb.) sulfureus Friese
sulfureus Friese, 1905:521, examined

Subgenus FRATERNOBOMBUS Skorikov
Alpigenobombus (Fraternobombus) Skorikov, 1922a:156,
type-species Apathus fraternus Smith (= Bombus
fraternus (Smith)) by subsequent designation of Frison,
1927:63
Bombus (Fraternobombus) Franklin, 1954:44

Bombus (Fr.) fraternus (Smith)
fraternus (Smith, 1854:385 [Apathus]) examined

Subgenus CROTCHIIBOMBUS Franklin
Bombus (Crotchiibombus) Franklin, 1954:51, type-spe-
cies Bombus crotchii Cresson by original designation

136

Bombus (Cr.) crotchiti Cresson
Crotchii Cresson, 1878:184

Subgenus ROBUSTOBOMBUS Skorikov

Volucellobombus Skorikov, 1922a:149, type-species
Bombus volucelloides Gribodo (?= B. melaleucus
Handlirsch) by monotypy

Alpigenobombus (Robustobombus) Skorikov, 1922a:157,
type-species Bombus robustus Smith by subsequent
designation of Sandhouse, 1943:597

Bombus (Robustobombus) Richards, 1968:217

COMMENT. Variation within and among the species
of this subgenus is particularly poorly understood and
a critical review is urgently needed.

Bombus (Rb.) melaleucus Handlirsch
melaleucus Handlirsch, 1888:228, examined
?volucelloides Gribodo, 1892:119 (provisional synonym)
?vogti Friese, 1903:254 (provisional synonym)
?nigrothoracicus Friese, 1904:188, examined (provisional
synonym)
melanoleucus Schulz, 1906:267, unjustified emendation

@ TAXONOMIC STATUS. Several of these nominal
taxa have been treated as separate species.

B. volucelloides is closely similar to B. melaleucus,
but has been considered to be a separate species (e.g.
Milliron, 19735). B. vogti is also closely similar to B.
volucelloides, and these two taxa have been consid-
ered both as conspecific (e.g. Franklin, 1913; Labougle,
1990) and as separate species (e.g. Milliron, 1973).
G. Chavarria (pers. com.) also believes that B.
melaleucus, B. volucelloides and B. vogti are all
conspecific.

In addition, it seems to me that B. nigrothoracicus is
more likely to be conspecific with B. vogti than with B.
ecuadorius (see the comments on B. ecuadorius).

Thus B. melaleucus is interpreted here in a very
broad sense, to include much variation that is not yet
well understood. Until more evidence to the contrary
is available from critical studies of patterns of varia-
tion, I shall treat them as parts of a single variable
species.

P.H. WILLIAMS

O NOMENCLATURE. For this species, the oldest
available name of which I am aware is B. melaleucus,
which becomes the valid name. The name B.
volucelloides has been in most common use, although
for just part of this species. However, it seems prema-
ture to conserve B. volucelloides by suppressing B.
melaleucus until the taxa are better understood, be-
cause the name B. melaleucus might yet be required
for a separate species or subspecies.

Bombus (Rb.) ecuadorius Meunier
Ecuadorius Meunier, 1890:66
?butteli Friese, 1903:254, examined (provisional syno-
nym)

@ TAXONOMIC STATUS. B. butteli is closely simi-
lar to B. ecuadorius. They have been considered to
be separate species (e.g. Franklin, 1913; Milliron,
1973b), although Franklin conceded that B. butteli
(which has grey hairs intermixed on the thoracic
dorsum) might be ‘only a variety or subspecies’ of
B. ecuadorius (which has the thoracic dorsum en-
tirely black).

B. ecuadorius females are very rare in collections.
For example, Milliron (19736) had seen only five
putative specimens (as opposed to 42 specimens of
B. butteli). Of these five specimens, four were fe-
males, and just one was a male, which is the same
specimen as the holotype of B. nigrothoracicus (see
the comments on B. melaleucus). This male is la-
belled ‘Bolivia / ?Peru’, whereas the rest of
Milliron’s B. ecuadorius are from Ecuador, with the
exception of one queen from ‘Peru’ (it carries no
further locality data). This putative male of B.
ecuadorius differs from the females in having yellow
hairs intermixed on the front and rear of the thorax.
This was not mentioned in the original description of
this male (under the name B. nigrothoracicus) by
Friese (1904), which Franklin (1913) used subse-
quently as the sole basis for associating the male
with B. ecuadorius.

Currently I favour another possible interpretation.
This views the male holotype of B. nigrothoracicus
instead as a semi-melanic male of B. melaleucus
(the males of B. volucelloides [= B. melaleucus] that
I have seen have the thoracic dorsum extensively
yellow). This might explain the difference in colour
pattern and distribution of this male from other B.
ecuadorius. However, a consequence of this inter-
pretation would be that the only remaining known
difference between B. ecuadorius and B. butteli
would be in colour pattern, because the main mor-
phological justification for regarding them as
separate species (the broader apical process of the
gonostylus of the putative male B. ecuadorius, now
B. melaleucus in the broad sense) would have been
removed. Further evidence is awaited.

CHECKLIST OF BUMBLE BEES

Bombus (Rb.) robustus Smith
robustus Smith, 1854:400, examined

? Bombus (Rb.) hortulanus Friese

hortulanus Friese, 1904:188, examined

[{hortulans Frison, 1925a:155, incorrect subsequent spell-

ing]

@ TAXONOMIC STATUS. B. robustus and B. hortu-
lanus have been considered both as conspecific (e.g.
Franklin, 1913; Frison, 1925a; G. Chavarria, pers.
com.) and as separate species (e.g. Milliron, 1973;
Asperen de Boer, pers.com.).

B. robustus and B. hortulanus are morphologically
similar. Among the specimens I have seen, individu-
als that have the sides of gastral terga I-II yellow (B.
robustus) also have pubescence extending to the mid-
dle or almost to the middle of tergum I, and the males
have the space between the inner basal process of the
gonostylus and the inner apical process narrower
than the apical process. Conversely, individuals with
the sides of terga I-II black (B. hortulanus) have at
least the medial third of tergum I hairless, and the
space between the inner processes of the male gono-
stylus is wider than the breadth of the apical process.

Until more evidence to the contrary is available
from critical studies of patterns of variation, I shall
treat them as separate species.

Bombus (Rb.) tucumanus Vachal
tucumanus Vachal, 1904:10

Subgenus SEPARATOBOMBUS Frison
Bremus (Separatobombus) Frison, 1927:64, type-species
Bombus separatus Cresson (= Bombus griseocollis
(DeGeer)) by original designation
Bombus (Separatobombus) Franklin, 1954:44

Bombus (Sp.) morrisoni Cresson
Morrisoni Cresson, 1878:183

Bombus (Sp.) griseocollis (DeGeer)
grifeo-collis (DeGeer, 1773:576 [Apis])
separatus Cresson, 1863:165

Subgenus FUNEBRIBOMBUS Skorikov
Alpigenobombus (Funebribombus) Skorikoy, 1922a:157,
type-species Bombus funebris Smith by monotypy
Bombus (Funebribombus) Richards, 1968:214

Bombus (Fn.) funebris Smith
funebris Smith, 1854:400, examined

Bombus (Fn.) rohweri (Frison)
rohweri (Frison, 1925a:144 [Bremus])

@ TAXONOMIC sTATUS. B. funebris and B. rohweri
have been regarded both as conspecific (Milliron,
1962) and as separate species (Frison, 1925a; Asperen
de Boer, 1993a; G. Chavarria, pers. com.). They have
been distinguished with reference to subtle morpho-
logical characters as well as to the consistently and
strongly differing colour patterns. Both Asperen de
Boer (1993a) and G. Chavarria (pers. com.) found that
they co-occur at some localities without intermediate
colour patterns. Further evidence is awaited.

Subgenus BRACHYCEPHALIBOMBUS Williams
Bombus (Brachycephalibombus) Williams, 1985b:247,
type-species Bombus brachycephalus Handlirsch by
original designation

138

@ TAXONOMICSTATUS. B. brachycephalus was not
explicitly placed in any subgenus by Richards (1968).
I described a separate subgenus Brachycephalibombus
for B. brachycephalus and B. haueri (Williams, 19855),
in order to maintain monophyletic groups (Williams,
1995).

Bombus (Br.) brachycephalus Handlirsch
brachycephalus Handlirsch, 1888:244
neotropicus (Frison, 1928:151 [Bremus])
krusemani Asperen de Boer, 1990:1, examined, new syno-
nym

@ TAXONOMIC STATUS. The description of B.
krusemani shows that this nominal taxon, known from
a single location, diverges slightly in colour pattern
from the otherwise widespread, common and variable
Central American species, B. brachycephalus. The
information available at present for B. krusemani is
consistent with the known range of variation within B.
brachycephalus (e.g. Labougle, 1990). Until more
evidence to the contrary is available from critical
studies of patterns of variation, I shall treat them as
parts of a single variable species.

Bombus (Br.) haueri Handlirsch
Haueri Handlirsch, 1888:234

COMMENT. Franklin (1913) and Labougle (1990)
believed that this species is closely related to B. crotchii
(although Labougle had not examined any males).
Surprisingly, Milliron (1973b) placed B. haueri in his
‘Dentatus-group’, without explanation (B. dentatus is
a junior synonym of the Indo-Chinese B. breviceps of
the subgenus Alpigenobombus). Possibly Milliron, at
least, may have been influenced by Skorikoy (1922a),
who placed B. haueri in the subgenus Alpigenobombus
(as Alpigenobombus (Alpigenobombus) haueri, which
he also listed next toAg. (Ag.) crotchii). However, both
sexes of species of the subgenus Alpigenobombus, as it
has been accepted recently (Richards, 1968; Williams,
1991), are easily distinguished from any New World
bumble bees because they have more teeth on the
mandibles.

I have examined the morphology of both sexes and,
on the basis of cladistic analysis, have grouped B.
haueri withB. brachycephalus (Williams, 1985b, 1995)
and with B. rubicundus (Williams, 1995). Further
evidence is awaited.

P.H. WILLIAMS

Subgenus RUBICUNDOBOMBUS Skorikov
Fervidobombus (Rubicundobombus ) Skorikoy, 1922a:154,
type-species Bombus rubicundus Smith by subsequent
designation of Sandhouse, 1943:597
Bombus (Rubicundobombus) Richards, 1968:217

Bombus (Rc.) rubicundus Smith
[Napensis Spinola in Osculati, 1850:201, published with-
out description]
rubicundus Smith, 1854:400, examined

Subgenus COCCINEOBOMBUS Skorikov
Alpigenobombus (Coccineobombus )Skorikovy, 1922a:157,
type-species Bombus coccineus Friese by subsequent
designation of Sandhouse, 1943:539
Bombus (Coccineobombus) Richards, 1968:214

Bombus (Cc.) coccineus Friese
coccineus Friese, 1903:254, examined

Bombus (Cc.) baeri Vachal
Baeri Vachal, 1904:10

CHECKLIST OF BUMBLE BEES

Subgenus DASYBOMBUS Labougle & Ayala
Bombus (Dasybombus) Labougle & Ayala, 1985:49, type-
species Bombus macgregori Labougle & Ayala by
original designation

@ TAxonomic status. B. handlirschi was not ex-
plicitly placed in any subgenus by Richards (1968),
and B. macgregori had yet to be described. I have
grouped B. handlirschi with B. macgregori in the
subgenus Dasybombus (Williams, 1995).

139

Bombus (Ds.) macgregori Labougle & Ayala
macgregori Labougle & Ayala, 1985:50, examined
menchuae Asperen de Boer, 1995:47, examined, new syno-

nym

@ TAXONOMIC STATUS. B. menchuae was de-
scribed from a single location and, on the basis of the
worker and male I have examined, appears to diverge
from B. macgregori only in colour pattern. Until more
evidence to the contrary is available from critical
studies of patterns of variation, I shall treat them as
parts of a single variable species.

Bombus (Ds.) handlirschi Friese
handlirschi Friese, 1903:255, examined

COMMENT. Franklin (1913) believed that this spe-
cies is closely related to B. rubicundus. Milliron
(1973b) knew ‘of no closely related species in the
Western Hemisphere’. I have examined the morphol-
ogy of both sexes and, on the basis of cladistic analysis,
have grouped B. handlirschi with B. macgregori as
sister species (Williams, 1995).

140

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CHECKLIST OF BUMBLE BEES

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150

The index includes references to names of bumble bees in the list, but not to those in the introduction or in the comments on each

INDEX

P. H. WILLIAMS

species. Valid names are shown in bold. Names in the genus group are shown in capitals.

abditus 114

abnormis 124
acutisquameus 104
adventor 107
ADVENTORIBOMBUS 107
affinis 130

afghanus 99

agnatus 126
AGROBOMBUS 107
agrorum 109, 118
alagesianus 134
alboanalis 125
alboniger 128
albopleuralis 115
ALLOPSITHYRUS 102
ALPIGENOBOMBUS 122
alpigenus 122
alpiniformis 120
ALPINOBOMBUS 119
alpinus 120

altaicus 99
americanorum 111
amurensis 121
anachoreta 108
andamanus 128
andreae 125

angustus 123
ANODONTOBOMBUS 123
APATHUS 102
apollineus 131
appositus 122

arcticus 110, 119, 120
ardens 125

arenicola 109
argillaceus 117
armeniacus 105, 118
arvensis 118

ashtoni 104
ASHTONIPSITHYRUS 102
asiaticus 135
assamensis 106
asturiensis 116

atratus 102, 112
atripes 110
ATROCINCTOBOMBUS 128
atrocinctus 129
atropygus 115

audax 130

auricomus 100
avanus 124
avinoviellus 99
azureus 112

baeri 138
baguionensis 125
baichengensis 135
baikalensis 125
balteatus 119
bannitus 107
barbutellus 104
beaticola 125
bellardii \05
bellicosus 112
bicolor 103
bicoloratus 114
bifarius 128
bimaculatus 128

birmanus 106

biroi 126
bischoffiellus 123
bohemicus 104
BOMBELLUS 99
BOMBIAS 100
BOMBUS 99, 129
BOOPOBOMBUS 100
borealis 122
braccatus 106

BRACHYCEPHALIBOMBUS 137

brachycephalus 138
branickii 105
brasiliensis 106, 113
BREMUS 99
breviceps 123
brevivillus 114
brodmannicus 125
bureschi 109
burjaeticus 130
butteli 136

cajennensis 112

calidus 102, 124
californicus 110
caliginosus 127
callophenax 99
campestris 105
canariensis 130

canus 104

carbonarius 112

cardui 118

cascadensis 127
caucasicus 134

celticus 107

centralis 127
CERATOPSITHYRUS 102
charharensis 121
chayaensis 122
chinensis 100, 104, 133
chinganicus 104, 134
chloronotus 105
CHROMOBOMBUS 107
cingulatus 125
cinnameus 107
CITRINOPSITHYRUS 102
citrinus 103, 106
clydensis 119
COCCINEOBOMBUS 138
coccineus 138

cognatus 110

combai 109
CONFUSIBOMBUS 101
confusus 101
consobrinus 115
consultus 103

contiguus 103
controversus 132
convexus 100

coreanus 104

cornutus 104

crawfordi 103

crotchii 136
CROTCHIIBOMBUS 135
cryptarum 130

CULLUMANOBOMBUS 131

cullumanus 131

ezerskii 117

daghestanicus 109
dahlbomii 111
DASYBOMBUS 139
decoomani 104

defector 99

dentatus 123
derhamellus 109
deuteronymus 108
diabolicus 120
difficillimus 121
DIGRESSOBOMBUS 110
digressus 113

diligens 112
distinguendus 121
DIVERSOBOMBUS 115
diversus 115
dolichocephalus 112
duanjiaoris 132
dumoucheli 110

ecuadorius 136
edwardsii 127
elegans 121
elisabethae 105
emiliae 112
EOPSITHYRUS 102
ephippiatus 128
eriophorus 134
erzurumensis 134
eurythorax 133
eversmanni 102, 125

EVERSMANNIBOMBUS 102

eversmanniellus 102
excellens 111

exil 107

exilis 107
EXILOBOMBUS 107
eximius 129
expolitus 104

exul 107

fedtschenkoi 121
ferganicus 105
fernaldae 105

FERNALDAEPSITHYRUS 102

fernaldi 128

ferrugifer 106
FERVIDOBOMBUS 110
fervidus 110
FESTIVOBOMBUS 128
festivus 101, 129
filchnerae 107
flavescens 110, 125
flavicollis 135

flavidus 106

flavifrons 126
flaviventris 134
flavodorsalis 111
flavopilosus 124
flavothoracicus 105, 133
flavus 105, 125
fletcheri 124

florilegus 130

folsomi 128
formosellus 133

CHECKLIST OF BUMBLE BEES

formosus 128

fragrans 121

franklini 129
FRATERNOBOMBUS 135
fraternus 135

frigidus \26

friseanus 133
fulvescens 108
fulvofasciatus 107
FUNEBRIBOMBUS 137
funebris 137
funerarius \06

gansuensis 106
genalis |23
genitalis 124
gerstaeckeri 115
gilgitensis 129
gilvus 109
globosus 106
grahami 123
griseocollis 137
grossiventris 132
guatemalensis 103

haematurus 124
haemorrhoidalis 106
handlirschi 139
handlirschianus 100
haueri 138

hedini 104, 109
heicens 135
himalayanus 99
hispanicus 116
hoenei 104, 133
honshuensis 109
HORTOBOMBUS 115
hortorum 1|16
hortulanus 137
huangcens 135
humilis 108
hummeli 115

huntii 128
hydrophthalmus 132
hyperboreus 119
HYPNOROBOMBUS 124
hypnorum 124
hypocrita \30

ignitus 130
imitator 110
impatiens 128
impetuosus 109
incertus 134
indicus 105
inexspectatus 109
infirmus 124
infrequens 124
insularis 103, 124
interruptus 103, 105
intrudens 103
irisanensis 114

Jacobsoni 130
japonicus 130
Jonellus 125

KALLOBOMBUS 118
kashmirensis \22
keriensis 134
kirbiellus 119
kirbyellus 119
klapperichi 104, 125
kohistanensis 124

kohli 112, 134
konakovi 109
Koreanus 117
koropokkrus 124
kotzschi 126
kozlovi 134
KOZLOVIBOMBUS 132
krusemani 138
kuani 106
Kulingensis 114
kurilensis 116

LABORIOPSITHYRUS 102
laboriosus 113
ladakhensis \33
LAESOBOMBUS 106
laesus 106

laevis 107
LAPIDARIOBOMBUS 132
lapidarius 134
LAPPONICOBOMBUS 124
lapponicus |27
latefasciatus 104
lateralis 128
latissimus 129
latofasciatus 99
lefebvrei 118
lemniscatus \24
lepidus 124
LEUCOBOMBUS 129
leucopygus 124
leucurus 124

licenti 104
liepetterseni 107
ligusticus 117

lit 107

linguarius 115
lissonurus 106
longiceps 135
longipes 115
lucorum 130
lugubris 100, 104, 128
luteipes |24
lutescens 109

macgregori \39
maculidorsis 106
maderensis 130
magnus 130

makarjini 99

malaisei 115, 129
margreiteri 99

martensi: 104
marussinus 99
MASTRUCATOBOMBUS 122
mastrucatus 122
maxillosus 104
maxwelli 115

mearnsi 125

medius 113
MEGABOMBUS 115
melaleucus \36
MELANOBOMBUS 132
melanopoda \\7
melanopygus \27
melanurus 121
menchuae 139
MENDACIBOMBUS 99
mendax 99
meridionalis 106, 116
mesomelas 118
METAPSITHYRUS 102
metcalfi 106
mexicanus 114

mimeticus 115
miniatocaudatus 118, 135
miniatus 133

mirus 124

mixtus 104, 127
mlokosievitzii 109
mocsaryi 106
moderatus 130
modestus 114, 125, 130
mongol 121
monozonus 104, 130
montanus 109, 128
monticola 128
montivagus 115
montivolanoides 125
montivolans 107
morawitzi 104, 132
morawitzianus 105
morawitziides 122
morio \12

morrisoni 137
MUCIDOBOMBUS 101
mucidus \02
muscorum \07
mysticus 103

napensis 138
nasutus 123
nemorum 104, 121
neoboreus 120
neotropicus 138
nepalensis 105
NEVADENSIBOMBUS 100
nevadensis 100
niger 112

nigripes 107, 111
nigriventris 112
nigrodorsalis 113
nigrothoracicus 136
nikiforuki 125
nivalis 119
niveatus 135
NOBILIBOMBUS 122
nobilis \22
norvegicus 105
notocastaneus 117
novus \05

nursei 126
nymphae 125

oberti \32
OBERTOBOMBUS 32
obtusus \34
occidentalis 131
oceanicus |25
ochraceus 105
ochrobasis 134
oculatus 133
ODONTOBOMBUS 115
opifex 112

opulentus 108
orichalceus 123
ORIENTALIBOMBUS 106
orientalis 106

ornatus 128

pallidus 107, 111
paradoxus 101
parthenius \24
pascuorum 109
patagiatus 130
pekingensis 117
pensylvanicus \11

151

152

peralpinus 124
perezi 104, 109
pereziellus 107
perniger 117
perplexus 118, 124
persicus \02
personatus 121
phariensis 133
picipes 125

pieli 105

pleuralis 126
POECILOBOMBUS 123
polaris 120
POMOBOMBUS 118
pomorum 118
portchinsky 116
potanini 109
praticola 127
pratorum \25
PRESSIBOMBUS 129
pressus 129
pretiosus 123
priscus 106

proteus 118
prshewalskyi 133
przewalskiellus 116
pseudobaicalensis 109
PSITHYRUS 102
pulcherrimus 122
pullatus 112
pyramideus 104
pyrenaeus 105, 126
pyrenes 99
PYROBOMBUS 123
pyrosoma 133

quadricolor 106

radoszkowskyi 118
redikorzevi 105

regeli 135

reinigi 130

reinigiellus 116

religiosus 118

remotus 109

reticulatus 133
RHODOBOMBUS 118
richardsi 104, 129, 134
richardsiellus 133
roborowskyi 121
ROBUSTOBOMBUS 136
robustus 137

rohweri 137

rotundiceps 125
RUBICUNDOBOMBUS 138
rubicundus 138
rubriventris 112
RUDERARIOBOMBUS 107
ruderarius 109

ruderatus 117
RUFIPEDIBOMBUS 129
rufipes 129
RUFOCINCTOBOMBUS 131
rufocinctus 131, 133
rufocognitus 123
rufofasciatus 133
rufoflavus 125

rupestris 105

saltuarius 116
saltuum 121
sandersoni 126
scandinavicus 128
schrencki 109
securus 118
semenovi 132
semenovianus 1|33
semenoviellus |32
semialbopleuralis 107
senex 114
SENEXIBOMBUS 114
senilis 108
separandus 134
SEPARATOBOMBUS 137
separatus 137
serrisquama 131
shaposhnikovi 100
shillongensis 125
SIBIRICOBOMBUS 134
sibiricus 134
sichelii 134

signifer 124

sikkimi 122
silantjewi 131
simillimus 132
simulatilis 109
simulus 123
sitkensis 127
skorikovi 106
smithianus 110
sololensis 103
sonani 124

sonomae 110
sonorus 111
SOROEENSIBOMBUS 118
soroeensis 118
sporadicus \29
steindachneri 1\13
stenothorax 133
stramineus 122
strenuus 120, 134
subbaicalensis 108
subdistinctus 121
SUBTERRANEOBOMBUS
subterraneus 121
subtypicus 124
suckleyi 104
SULCOBOMBUS 101
sulfureus 135
superbus 99
superequester 109
supremus 115
sushkini 116
susterai 104, 105
susteraianus 105
sylvarum 109
sylvestris 106
sylvicola 127

tahanensis 125
tajushanensis 105, 114
TANGUTICOBOMBUS 132
tanguticus 132

tenellus 134

tenuifasciatus 126, 134
terminalis 129, 130
ternarius 128
TERRESTRIBOMBUS 129

P. H. WILLIAMS

terrestris 130
terricola 131

tersatus 115
tetrachromus 122, 124
thoracicus 109, 112
THORACOBOMBUS 107
tianschanicus 106
tibetanus 104, 124
tibetensis 134
tichenkoi 115

tonsus 133
transbaicalicus 105
transversalis \12
tricolor 119
TRICORNIBOMBUS 110
tricornis 110
trifasciatus 115
trilineatus 134
trinominatus 114
tristis 108, 119
tucumanus 137
tunicatus 129
turkestanicus 99
turneri 104

UNCOBOMBUS 124
unicolor 104, 109
unicus 132
ussurensis 115

vagans 127
validus 122
vallestris 129
vandykei 127
variabilis 103, 108
variopictus 133
varius 99, 105
vasilievi 130
velox 109
velutinus 112
vestalis 104
veteranus 109
villarricaensis 117
violaceus 112
vogti 136
vogtiellus 109
VOLUCELLOBOMBUS 136
volucelloides 136
vorticosus 135
vosnesenskit 128

waltoni 100
waterstoni 133
weisit 113
wilemani 115
wilmattae 128
wurflenii 122
wutaishanensis 133

xelajuensis 114
xionglaris 132
xizangensis 122

yezoensis 116

yuennanensis 109
yuennanicola 124
yuennanicus 118

zhadaensis 132
zhaosu 127
zonatus 108

Bulletin of The Natural History Museum
Entomology Series

Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside front
cover). Where the complete backlist is not shown, this may also be obtained from the same address.

Volume 58

No. | The mealybug genus Planococcus (Homoptera: Pseudococcidae). J.M. Cox. 1989. Pp. 1-78, 40 figs.

No. 2 The Simuliidae (Diptera) of the Santiago onchocerciasis focus of Ecuador. A.J. Shelley, M. Arzube & C.A.
Couch. 1989. Pp. 79-130, 153 figs (including 2 plates in colour).

Volume 59

No. 1 The songs of the western European bush-crickets of the genus Platycleis in relation to their taxonomy
(Orthoptera: Tettigoniidae). D.R. Ragge. 1990. Pp. 1-35.
A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). C.M.F. von Hayek. 1990.
Pp. 37-115.

No. 2 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). S.J. Brooks & P.C.
Barnard. 1990. Pp. 117-286.

Volume 60
No. | The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). P.H. Williams. 1991.
Pp. 1-204.

No. 2 Sattleria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae). L.M. Pitkin & K.
Sattler. 1991. Pp. 205-241.
A review of wing reduction in Lepidoptera. K. Sattler. 1991. Pp. 243-288.

Volume 61
No. | Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76.
No. 2 Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Scaffer and Perissopteryx Warren

(Lepidoptera: Geometridae). M. Kruger & M.J. Scoble. 1992. Pp. 77-148.

Volume 62

No. | Caloptilia \eaf-miner moths (Gracillariidae) of South-East Asia. Decheng Yuan and Gaden S. Robinson.
1993. Pp. 1-37.

No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular
reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Linda M. Pitkin. 1993.
Pp. 39-159.

Volume 63

No. | A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of the

immature stages of economically important insect species (mainly Hemiptera and Lepidoptera). D.W.
Huang and J.S. Noyes. 1994. Pp. 1-135.

No. 2 A taxonomic review of the common green lacewing genus Chrysoperla (Neuroptera: Chrysopidae). S.J.
Brooks. 1994. Pp. 137-210.

Volume 64

No. 1 Revision of the neotropical genus Oospila Warren (Lepidoptera: Geometridae) M.A. Cook and M.J.
Scoble. 1995. Pp. 1-115.

No. 2 Encyrtidae of Costa Rica (Hymenhoptera: Chalcidoidea): the genus Aenasius Walker, parasitoids of
mealybugs (Homoptera: Pseudococcidae). J.S. Noyes and H. Ren. 1995. Pp. 117-164.

Volume 65

No. | A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae).
Y. Zhang and M.D. Webb. 1996. Pp 1-103.

No. 2 Encyrtidae (Hymenoptera: Chalcidoidea) of Costa Rica: the genera and species associated with jumping

plant-lice (Homoptera: Psylloidea). J.S. Noyes and P. Hanson. 1996. Pp. 105-164.

Volume 66

No. | A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae).
Y. Zhang and M.D. Webb. 1997. Pp 1-121.

No. 2 Microtermes in East Africa (Isoptera: Termitidae: Macrotermitinae) S. Bacchus. 1997. Pp. 123-171.

Mealybugs of the genera Eumyrmococcus Silvestri and Xenococcus —
Silvestri associated with the ant genus Acropyga Roger and a review

_ of the subfamily Rhizoecinae (Hemiptera, Coccoidea, Pseudococeidas
D.J. Williams
Monophyly of the dacetonine tribe-group and its component tribes
(Hymenoptera: Formicidae)
Barry Bolton
An annotated checklist of bumble bees with an analysis of patterns se
description (Hymenoptera: Apidae, Bombini)

Paul H. Williams

ENTOMOLOGY SERIES

Vol. 67, No. 1, June 1998

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