Bulletin of the
British Museum (Natural History)
^LIBRARY
Entomology series Vol 46 1983
British Museum (Natural History)
London 1983
Dates of publication of the parts
No 1 28 April 1983
No 2 26 May 1983
No 3 28 July 1983
No 4 25 August 1983
ISSN 1524-6431
Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset
Contents
Entomology Volume 46
Page
No 1 The generic and tribal classification of spore- feeding Thysanoptera
(Phlaeothripidae: Idolothripinae)
L. A. Mound & J. M. Palmer 1
No 2 A revision of the Afrotropical mole-crickets (Orthoptera:
Gryllotalpidae)
B. C. Townsend 175
No 3 Key to the genera of galerucine beetles of New Guinea, with a review
of Sastra and related new taxa (Chrysomelidae)
Sharon L. Shute 205
No 4 The Afrotropical dacetine ants (Formicidae)
Barry Bolton . 267
GENERAL
2 8 APR 1983
Bulletin of the
British Museum (Natural History)
V %/ /
>. UBRARY .jV,
The generic and tribal classification of
spore-feeding Thysanoptera
(Phlaeothripidae : Idolothripinae)
L. A. Mound & J. M. Palmer
Entomology series
Vol 46 No 1 28 April 1983
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1983
The Entomology series is produced under the general editorship of the
Keeper of Entomology: Laurence A. Mound
Assistant Editor: W. Gerald Tremewan
ISSN 0524-6431 Entomology series
Vol 46 No 1 pp 1-174
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 28 April 1983
The generic and tribal classification of spore-feeding GENERA^
Thysanoptera (Phlaeothripidae: Idolothripinae)
L. A. Mound & J. M. Palmer
>£eJ5? it vAVt>^
Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7
5BD
Contents
Synopsis 1
Introduction 2
Acknowledgements and depositories 3
Problems in constructing classifications 3
Character inheritance 4
Zoogeography 5
Patterns of speciation 5
Characters studied 6
Family-group classification of Idolothripinae 8
Key to subtribes of Idolothripinae 11
Genus-group classification of Idolothripinae 12
Key to genera of Idolothripinae 14
Tribe Pygothripini 20
Genera of Pygothripina 21
Genera of Allothripina 30
Genera of Compsothripina 34
Genera of Gastrothripina 38
Genera of Diceratothripina 40
Genera of Macrothripina 50
Tribe Idolothripini 62
Genera of Elaphrothripina subtrib. n 62
Genera of Idolothripina 71
Genera of Hystricothripina 80
Taxa transferred from Idolothripinae to Phlaeothripinae 88
Tribe Apelaunothripini 89
Tribe Docessissophothripini 90
References 97
Distribution tables of Idolothripinae 109
Index 165
Synopsis
In this paper 154 genus-group names are recognised as available in the subfamily Idolothripinae, including
six new genera. However, 75 of these names are here placed in synonymy, including 36 new generic
synonyms, and one genus was unavailable for study. Keys are provided to the 78 remaining genera, based
on a study of more than 480 of the 600 species listed here in this group. In addition, a further 200
species-group names are listed in synonymy, including eight new synonyms, and 17 new species and 1
new combinations are established. These taxa are arranged into two tribes and nine sub-tribes, with eight
family-group names being placed in synonymy. Moreover, 26 generic names are transferred from the
Idolothripinae to the subfamily Phlaeothripinae and placed in two newly recognised tribes, the Ape-
launothripini with two genera and 12 species, and the Docessissophothripini with nine genera and 99
species. Decisions on the Docessissophothripini are based on the study of 44 species, and include 13 new
generic synonyms, two new specific synonyms and 69 new combinations. Various aspects of the biology ,
distribution and structure of spore-feeding thrips are discussed, where these seem relevant to problems ot
constructing a phylogenetic classification.
Bull. Br. Mus. not. Hist. (Ent.) 46(1): 1-174 Issued 28 April 1983
2 L. A. MOUND AND J. M. PALMER
Introduction
The family Phlaeothripidae, the sole family in the Thysanoptera sub-order Tubulifera, compris-
es about 2700 described species (Mound et al., 1980). Members of this family are structurally
uniform, although diverse in superficial appearance and with a wide range of biologies. Probably
about half of them feed on leaves of green plants, in tropical countries often inducing galls
(Ananthakrishnan, 1978), although in temperate regions phlaeothripids are most commonly
observed in the flowers of Compositae and Gramineae (Mound et al. , 1976) . A number of often
unrelated species are predatory on other small arthropods. However, almost half of the
phlaeothripid species are associated with fungi - under bark, on dead twigs and branches, or in
leaf litter - some feeding on spores but the majority feeding on hyphae or possibly the external
digestion products of fungal decay. This paper concerns the classification of those species which
feed on fungal spores, most of which comprise the holophyletic, worldwide, sub-family
Idolothripinae.
Existing classifications of Phlaeothripidae derive largely from two publications by Priesner
(1949: 1961) and these in turn are derived in part from earlier studies by Karny (1921a; 1925).
Unfortunately, the tradition of work throughout this period often involved acceptance of
previously published taxa without further re-examination of the specimens involved. Thus, the
key to genera of the world by Priesner (1949) does not indicate which genera he was unable to
study personally, although it is evident that parts of the key are based solely on published
descriptions. This is also true of Priesner's 1961 classification, which is reproduced almost
unmodified by Ananthakrishnan (1969d) and Jacot-Guillarmod (1978). This approach could
only produce a typological classification, that is a classfication emphasising the importance of
single characters. Moreover, characters found to be of use in classification by later workers are
not available in the descriptions of earlier taxa, with the result that spurious comparisons are
often made.
Stannard (1957) broke with this tradition by personally examining a wide range of phlaeothri-
pid taxa. His outstanding analysis of the North American genera set entirely new standards, by
demonstrating a range of previously unobserved characters and by clearly introducing the
concept of evolutionary relationships into the systematics of the family. Following this lead,
Mound (1974ft) re-examined almost all of the 100 species comprising the Nesothrips genus-
group, thus producing a revised generic classification, and Palmer & Mound (1978) redefined a
further eight genera from the Oriental region having examined the 60 species concerned. Few
other workers have ever examined more than a small percentage of described taxa, and because
descriptions often have a low information content (comprising colour and silhouette characters
mainly) the systematic confusion is considerable. The objectives of the present study were
therefore:
1, to examine the type-species of all genera of Idolothripinae (sensu lato);
2, to examine as many species as possible described in or referred to this group (together with
any relevant species described in the Phlaeothripinae), and also to examine more recently
collected unidentified material;
3, to try to recognise clusters of related species and, from these presumably holophyletic
groupings, to construct a new classification at genus level and above based on phylogenetic
rather than typological principles;
4, to communicate this revised classification in the form of an illustrated key to genera.
The first of these objectives was achieved almost completely; only one genus remained
unstudied. The second objective was also achieved in that 75 per cent of the known species were
studied; those not examined are indicated in the lists of species under each genus by an asterisk
(*). The third objective has been achieved only partially; as discussed below, the authors have
frequently sacrificed phylogenetic principles to traditional classificatory expediency at both
tribal and generic levels. Formal diagnoses are not given for most genera, although brief
comparative notes are given for each genus to supplement the key, and at least one species of
each genus is illustrated.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 3
Acknowledgements and depositories
The authors wish to express their gratitude for the kind co-operation and forbearance of many
colleagues throughout the world, without which this study could not have been completed. We
gratefully acknowledge the help of the curators at all the collections listed below, but Steve
Nakahara of Washington, Shuji Okajima of Tokyo and Annette Walker of Auckland have been
particularly generous in providing frequent advice, information and specimens. Comprehensive
taxonomic studies are dependent on co-operation between workers of differing views, and we
are pleased to acknowledge the debts we owe to so many other biologists.
AMG Albany Museum, Grahamstown, South Africa
ANIC Australian National Insect Collection, C.S.I.R.O., Canberra, Australia
BCIQ Bureau of Commodity Inspection and Quarantine, Taipei, Taiwan
BCM Prof. A. Bournier collection, Ecole Nationale Superieure Agronomique,
Montpellier, France
BMNH British Museum (Natural History), London, U.K.
BPBM Bernice P. Bishop Museum, Honolulu, Hawaii
CAS California Academy of Sciences , San Francisco , U . S . A .
DART Department of Agricultural Research, Taipei, Taiwan
DEI Deutsches Entomologisches Institut, Eberswalde, East Germany
FSAC Florida State Arthropod Collection, Dept. of Agriculture, Gainesville, U.S.A.
INHS Illinois Natural History Survey , Urbana , U.S.A.
MACN Museo Argentine de Ciencias Naturales Bernardino Rivadavia de Buenos Aires,
Argentina
MDA Museu do Dundo, Angola
MLPA Museo de la Plata, Argentina
MNHN Museum National d'Histoire Naturelle, Paris, France
MNHO Osaka Museum of Natural History, Japan
MR AC Musee Royal de 1'Afrique Centrale, Tervuren, Belgium
NCIP South African National Insect Collection, Pretoria, South Africa
NIAT National Institute of Agricultural Science, Tokyo, Japan
NMB Naturhistorisches Museum, Basel, Switzerland
NMG Naturhistoriska Museum, Goteburg, Sweden
NMV Naturhistorisches Museum, Vienna, Austria
NRS Naturhistoriska Riksmuseet, Stockholm, Sweden
NZAC New Zealand Arthropod Collection, D.S.I.R. , Entomology Division,
Auckland, New Zealand
OCT Dr Shuji Okajima Collection, Tokyo Agricultural University, Japan
QMB Queensland Museum, Brisbane, Australia
RPAESIC Rio Piedras Agricultural Experimental Station Insect Collection, Argentina
SMF Senckenberg Museum, Frankfurt, West Germany
TM Termeszettudomanyi Muzeum (Hungarian Natural History Museum) , Budapest, Hungary
TNA Prof. T. N. Ananthakrishnan, Loyola College, Madras, India
UN AM Universidad Nacional Autonoma de Mexico , Mexico , D . F. , Mexico
USNM United States National Museum of Natural History, Washington, D.C. , U.S. A.
ZMB Zoologisches Museum an der Humboldt-Universitat zu Berlin, East Germany
Problems in constructing classifications
Biologists construct classifications for two major purposes: firstly to provide an identification
and data storage and retrieval system, secondly to reflect wherever possible those evolutionary
relationships presumed to exist between different taxa. These two purposes, although apparent-
ly independent, are often closely inter-related, because a classification derived from evolutionary
relationships has the potential for yielding further biological information whereas a typological
classification based solely on superficial resemblances has no such potential. Some Thysa-
noptera specialists have claimed that it is not possible to ascertain evolutionary pathways, and
therefore a phenetic classification is the proper practical solution. However, none of them
4 L. A. MOUND AND J. M. PALMER
follows such an extreme attitude to its logical conclusion, e.g. by placing all wingless forms in one
group and all winged forms in another, and most classifications are an uneasy and unexplained
mixture of phenetics and phylogenetics. The present authors have considered many of the
problems involved in the production of phylogenetic classifications of Thysanoptera (Mound et
al., 1980; Mound & Palmer, 1981), and have attempted to point out areas where our
assumptions have least justification. We accept that a completely phylogenetic classification of
this group is not yet possible, but our objective has been to emphasise the presumed underlying
evolutionary relationships between taxa, in contrast to most of our predecessors who have
emphasised the often startling, but frequently superficial, differences which are readily
observed.
Character inheritance
A phylogenetic classification is based on the recognition of two or more taxa jointly exhibiting
one or more derived characters - apomorphies - not found in other taxa. Two such taxa sharing
an apomorphy not exhibited by related taxa (and each itself characterised by a further unique
apomorphy), may be regarded as holophyletic sister-groups. That is, they are derived from a
single common ancestor, and moreover include all extant taxa which have evolved from that
ancestor. Shared primitive characters - plesiomorphies - although indicating relationship
cannot be used to define a natural evolutionary group (Hennig, 1966).
Unfortunately, this strictly logical approach is subject to practical difficulties when applied to
some groups of organisms (Gauld & Mound, 1982). It is often possible to characterise one
particular group through the presence of one or more apomorphies, but the sister-group may
remain unclear (Greenwood, 1980). In such instances the true sister-group appears to lie within
some residual group that is itself imperfectly characterised. Moreover, the strictly logical
approach assumes that an apomorphy will find expression in all species in a holophyletic group,
although this would not be expected from current genetical theory (Maynard Smith, 1975) and is
contrary to observation in some groups of insects (Gauld & Mound, 1982).
Within any particular evolutionary lineage it is not unusual to observe a tendency for a
particular character to be developed (Stys, 1967), although not all species will exhibit the
character and its development is not a direct measure of evolutionary relationship. Such a
character, although inherited by all members of the lineage, fails to be expressed in some species
(even in some individuals of a species) probably because its ontogenetic development is
inhibited by some other aspect of the genotype. Similarly, reversal of a character state during
evolution is apparently common in Thysanoptera, e.g. antennal segment number (Mound &
Palmer, 1981). Unfortunately, many apomorphies in this group of insects involve losses (e.g. loss
of metathoracic sternopleural sutures or praepectus) or reductions (e.g. number of antennal
sense cones or sternal discal setae), and such characters are more likely to have evolved more
than once than a new complex structure. This irregular pattern of character inheritance,
involving reversal and recurrent reductions in structures, may be a reflection of the ecology of
the group as discussed below under patterns of speciation.
Further complications of character inheritance in Phlaeothripidae result from sex-linkage and
allometry. In fungus-feeding phlaeothripids, including spore-feeding idolothripines, males are
often oedymerous (large) and develop a range of structures not found in females. For example,
Mecynothrips and Elaphrothrips males have tubercles or setae on the forelegs not found in
females, Bactrothrips males usually have abdominal tubercles which do not occur in females,
and Gastrothrips, Diceratothrips and Macrothrips all include species in which males may be
oedymerous or gynaecoid (female-like). This relationship is reversed in Machatothrips, howev-
er, in which females exhibit the secondary sexual characters. Moreover, in gall-forming
phlaeothripines, males are usually small and constant in size but females are variable; large
females may look quite different from small females due to enlargement of the forelegs and
pronotum. Similar phenomena can be associated with wing reduction, the wingless morph again
usually being female.
The difficulty for classification arises particularly when related evolutionary lineages involve
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 5
emphasis on different morphs; that is one lineage may emphasise male-ness whereas another
emphasises female-ness. For example, bicornis, the most common species of Diceratothrips, is
also the largest member of the genus with the greatest sexual dimorphism. The other members of
Diceratothrips tend to have both sexes similar in structure to female bicornis, moreover, these
females are structurally similar to females of Sporothrips. The ontogenetic threshold at which
various characters associated with wing length or sexuality may be expressed will itself be liable
to variation during evolution, and could well be independent of close phylogenetic relationships.
Emphasis by taxonomists on striking differences between males of different species will tend to
increase the number of 'genera' without producing any increased understanding of the under-
lying evolutionary relationships.
Zoogeography
The observed geographic distribution of natural groups of organisms can provide additional
evidence in support of a classification. For example, in the classification adopted here the
Hystricothripina is predominantly New World and the Idolothripina predominantly Old World.
Moreover, the Macrothripina is largely Oriental and the Pygothripina predominantly Austro-
Oriental. Such distribution patterns reassure the systematist that he is probably recognising real
evolutionary groups. However, natural distribution patterns have been disrupted in many
instances by human trading. For example, Nesothrips propinquus is now found at most points
along the old shipping route between Britain and New Zealand and was probably distributed in
hay and straw; Nesothrips lativentris, which is frequent on dead palm leaves and coconuts, is now
widespread in the tropics; other species appear to have been transported across the Atlantic by
the slave trade, or across the Indian Ocean by even earlier shipping (Mound, 1974b). Moreover,
since many species are known only from single individuals or single samples it is probable that
recognition of further synonymy will increase the number of species known to have been
distributed by man (Mound & Walker, 1982).
Patterns of speciation
There is little evidence that most spore-feeding thrips exhibit any particular host specificity,
although there tends to be some correlation between stylet diameter and the size of spores found
in the gut, and a few species are found in association with particular plants, e.g. Sporothrips on
dead leaves of Palmetto palms in Florida. Not only are two or more congeneric species found
together quite frequently, but such species are sometimes widespread, e.g. Elaphrothrips
species (Palmer & Mound, 1978). The extensive distributions of such species are probably
facilitated by the availability of suitable fungal spores widespread on dead branches and in leaf
littler. Palmer & Mound (1978) interpreted the Oriental Elaphrothrips species as structurally
variable and behaviourally vagile with extensive and broadly overlapping geographical ranges.
In contrast, Dr R. Johansen is currently describing numerous species of Elaphrothrips from
Mexico, thus implying that the biology of these insects is different in the Neotropics. Spore-
feeding species of low vagility (low dispersive ability) may develop clines, such as that of
Allothrips megacephalus across North America (Mound, 19720). Unfortunately, although at
times it may be possible to relate different biologies to different patterns of speciation, most
species are based on few specimens (e.g. Bactrothrips) and so the concept of species is itself often
poorly defined.
Commenting on the large number of co-existing congeneric species of fungus-feeding thrips in
leaf litter in southern Brazil, Mound (1977) suggested a possible relationship to a seasonal excess
of available food, and subsequent reduction in competition between species. Such a reduction in
competition, and consequent reduction in selection pressures on incipient species, may be
related to the frequency of homoplasy (reversal and parallelism) in character inheritance within
these thrips (Gauld & Mound, 1982). Host specific phytophagous thrips will almost certainly be
subject to greater selection pressures in dividing up available resources, because they must
develop the necessary behaviour patterns to find and respond to their particular niches.
Unfortunately, the ecology, and hence speciation patterns, associated with this type of
6 L. A. MOUND AND J. M. PALMER
evolutionary strategy, involving competition and resource partitioning, is more fully investi-
gated than the ecological and evolutionary strategies of non-specific leaf litter insects. If it should
prove correct that the available resources in leaf litter are periodically in excess of the demands
of the total arthropod fauna, then the classical evolutionary model involving niche separation
and competition may be inappropriate for these small organisms with a short life cycle.
Characters studied
Head
Head shape is often useful in recognising relationships between taxa, e.g. relative length/
breadth, constriction behind eyes or basally, prolongation in front of eyes, elevation in
mid-line. However, shape is readily distorted by cover-glass pressure on specimens mounted
onto microscope slides, the degree of distortion being particularly remarkable when the
posterior half of the head is deep dorso-ventrally.
Maxillary stylets are broader (5-10 /xm) in Idolothripinae than in most Phlaeothripinae
(2-3 Aim), but are intermediate in width (3-6 /am) in members of the phlaeothripine tribes
Apelaunothripini and Docessissophothripini. The plesiomorphic position of the stylets in
Idolothripinae is probably deeply retracted and parallel medially (Figs 2-11), the derived
condition being wide apart and low in the head (Figs 134-144). However, polarity of this
character is confused by homoplasy, both reversal (Zeuglothrips , Fig. 342) and parallelism (cf.
Ozothrips and Nesothrips, Figs 13, 142). The stylets of Docessissophothripini are sometimes
exceptionally long and convoluted (Figs 385-390).
Maxillary guides are thickened internal structures associated with the stylets of Phlaeothri-
pinae. In Docessissophothripini they are large bowed structures (Figs 385-390), and in
Haplothrips species they form a characteristic bridge (Mound et al., 1976). However, in
Idolothripinae these structures are developed only in some Pygothripina.
Mouth cone shape is frequently stressed in older descriptions, but is of limited systematic
value. The apparent shape depends largely on whether the mouth cone is directed posteriorly
(pointed) or ventrally (rounded).
Maxillary palps are two-segmented and generally rather large in idolothripines, but in
Allothripina the terminal sensorium is often enlarged giving the appearance of a third segment
(Fig. 77).
Compound eyes, although rounded and multifaceted in most species, are sometimes reduced
to a few facets (Anaglyptothrips , Fig. 103) or prolonged ventrally (Bolothrips, Fig. 94).
Reduction in facet number is usually associated with aptery, and ventral prolongation of the eyes
seems to be correlated with the grass-living habit.
Ocelli are almost always present in macropterae, usually absent in apterae, and frequently
reduced in micropterae. Thus, although there is a positive correlation between presence of
wings and presence of ocelli, the development of these two structures is controlled independent-
ly. When the head is prolonged in front of the eyes, the fore ocellus may be unusually distant
from the hind ocelli.
Setae are developed on the head surface in a fairly restricted pattern. There are usually about
three pairs of setae associated with the ocelli, and either the post-ocellars or preocellars are
frequently enlarged. Most species have a pair of major post-ocular setae; a few have two pairs,
the second arising on the cheeks, or medially, or on the vertex. Many Idolothripinae have a
series of stout setae on the cheeks.
Antennal structure is important in deducing relationships between taxa. The plesiomorphic
number of antennal segments in Phlaeothripidae is eight (Mound et al., 1980). However,
Idolothripinae may have evolved from species with only seven segments, because the sub-
groups which are here regarded as least advanced (Pygothripina and Allothripina) tend to have
that number. If this is so, then the number of antennal segments has undergone reversal to eight
and reduction to seven (or less) several times. Similarly the plesiomorphic sense cone formula in
idolothripines is considered to be two on HI and two on IV, but in most species the number of
sense cones on IV is doubled. The condition of three sense cones on IV has probably evolved
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 7
more than once, being found in Cryptothrips (Pygothripina), Bolothrips (Compsothripina) and
Gastrothrips (Gastrothripina). The relative lengths of antennal segments have been used
frequently for defining species, but caution is required in using such ratios because segment
lengths are sometimes affected by allometric growth.
Prothorax
Pronotal shape is often affected by sex and morph correlated allometry, and is not usually
significant at genus-level (see Diceratothrips) , although some groups exhibit a tendency for the
anterior margin or median line of the pronotum to be thickened.
Epimeral sutures are usually present posterolaterally on the pronotum, but these are
incomplete or fused in some taxa.
Setae are borne in a very regular pattern on the pronotum , there being five pairs of major setae
in most species - antero-marginals (am), antero-angulars (aa), mid-laterals (ml), epimerals
(epim) and postero-angulars (pa). The two anterior pairs are often shorter than the other three;
only rarely are these setae undeveloped (Anaglyptothrips).
Praepectal plates (or praepectus) are a pair of small sclerites on the prosternum, arising in
front of the major probasisternal plates (Figs 140, 143), but they are often reduced (Fig. 372) or
absent (Fig. 371).
Pterothorax
The mesonotum in macropterae often bears a pair of major setae laterally. In apterae this
sclerite is reduced and in extreme instances fused to the metanotum. The mesothoracic spiracle
is situated on the anterior angles of the segment and is sometimes surrounded by an area of
specialised sculpture (e.g. Dinothrips). Ventrally, the mesopraesternum is usually a boat-
shaped sclerite but is often reduced and occasionally absent.
The metanotum often bears a pair of stout setae medially, but the sclerite is reduced and
transverse in apterae. The metathoracic sternopleural sutures, which curve posteriorly from the
mid-coxal cavities (Fig. 100), are regarded as a plesiomorphic character which has been lost in
the more advanced subgroups of Idolothripinae (Fig. 99). This derived condition may have
developed more than once, because the suture is variable in position, sometimes reduced in
length, or so slender as to be almost obliterated (Carientothrips) . Although generally constant
within groups, the sternopleural suture is variable within a few species (e.g. Nesothrips
propinquus).
The katepisternum and anepisternum of the metathorax are swollen in some groups (Hystri-
cothripina) and the anepisternal suture is short (Figs 353-355). However, the plesiomorphic
condition seems to be represented by a complete suture (Fig. 127), although these sclerites are
often eroded in reduced or apterous forms (Figs 21-22).
Theforewings bear a fringe of cilia although, unlike Terebrantia, these cilia do not arise from
sockets; there are no true longitudinal veins and the surface of the wing does not bear
microtrichia (Mound etal., 1980). On the distal hind margin the forewings often bear duplicated
cilia, ranging in number from one to almost one hundred. Idolothripines show less variation in
wing length than phlaeothripines; micropterae are rare and hemimacropterae unrecorded, most
species being macropterous and/or apterous.
The forelegs often bear a small or large tarsal tooth in one or both sexes. Similarly the
foretibiae, forefemora and even the forecoxae may bear one or more tubercles in different
genera. These tubercles, and particularly the swelling of the forefemora, are subject to
allometric growth patterns, and they are also usually sex-linked. Moreover, the production of
tubercles may recur in particular groups of related genera, e.g. many species of Macrothripina
have a small tubercle at the inner apex of the foretibiae in both sexes (Figs 220-221).
Abdomen
Pelta is the term applied to the first abdominal tergite. In most species this tergite is reduced to a
small median tergite, only in Allidothrips is it completely transverse (Fig. 64). In some
8 L. A. MOUND AND J. M. PALMER
Hystricothripina the setae on the pleurites of the first segment have migrated onto the pelta (Figs
375-384), and in some Pygothripina the pelta is eroded posteromedially (Figs 36-37).
Wing-retaining setae are developed sub-medially on tergites II-VII of most macropterae.
These setae are usually sigmoid in shape (Fig. 294), but are sometimes almost straight (Fig. 326)
or even flattened (Fig. 369). The plesiomorphic condition for the subfamily Idolothripinae is
here interpreted as involving only one pair of wing-retaining setae on each tergite as in
Pygothripini (Fig. 43). The condition with two pairs of such setae as in most Idolothripini is
considered apomorphic, and the three or more pairs found in Mecynothrips species (Fig. 297) is
particularly advanced. Micropterae and apterae have the wing-retaining setae reduced or
absent.
Tergite IX setae are important in the recognition of the two subfamilies of Phlaeothripidae.
The males of almost all phlaeothripine species have the submedian pair of setae (B2) on tergite
IX short and stout, whereas males and females of all Idolothripinae have these setae as long as
the dorsal (B^) and lateral (B3) pairs.
The tube is the most characteristic structure of Phlaeothripidae, abdominal segment X being
entirely tubular with the anus emerging terminally and the genital ducts between segments IX
and X. The tube is greatly swollen in several genera of Pygothripina and Diceratothripina. In
many Idolothripini, as well as in Cleistothrips (Pygothripina) and Campulothrips (Diceratothri-
pina), the tube bears prominent lateral setae. The base of the tube is emarginate ventrally in
males, but completely cylindrical in females.
Sternal glandular areas are rarely developed in Idolothripinae, the only known species being
in the Macrothripine genera Dichaetothrips, Peltariothrips and Tarassothrips. Glandular areas
are frequently developed in the Phlaeothripinae, however - on the median sternites in both
sexes of many Plectrothripini (Okajima, 1981), on the median sternites in the males of most
species of Docessissophothripini (p. 90), and on the eighth sternite in the males of many other
Phlaeothripinae.
Sternal discal setae are usually developed as a single transverse row medially on each sternite,
but they are duplicated in some species (Actinothrips) or reduced in others (Priesnerielld) .
Family-group classification of Idolothripinae
Two sub-families are recognised in the family Phlaeothripidae, the Phlaeothripinae and the
Idolothripinae. This latter group, the spore-feeding thrips, was defined in its modern sense
(under the name Megathripinae) by Stannard (1957), who characterised it by the presence of
broad maxillary stylets and the absence in males both of sternal glandular areas and of short stout
B2 setae on tergite IX. The latter characteristic is constant throughout the Idolothripinae as
defined in the present paper; however, three species of Macrothripina are now known which
appear to have sternal glandular areas in males and/or females (see Dichaetothrips, Peltario-
thrips and Tarassothrips).
Broad maxillary stylets are a functional adaptation to feeding on fungal spores, but although
the sub-family Idolothripinae appears to be a holophyletic group the characteristic of broadened
stylets has also evolved in several groups of Phlaeothripinae. Two such groups, the tribes
Apelaunothripini and Docessissophothripini, are discussed in this paper (p. 88) because they
were treated as idolothripines by Priesner (1961). In addition, species of the quite unrelated
genus Lissothrips, associated with lichens and possibly mosses, also have rather broad stylets.
Moreover, a Neotropical species-group of Liothrips is known in which the males may be
mistaken for idolothripines in that they lack sternal glandular areas and have three pairs of
elongate setae on the ninth tergite (Mound, 19746: 182).
A further negative characteristic of idolothripine species is the apparent absence of maxillary
guides, with the exception of a few species of Pygothripina, particularly from New Zealand.
Table 1 indicates the family-group names in the Idolothripinae, and it must be emphasised
that according to the Code of Zoological Nomenclature (1961) all categories of family-group
names are co-ordinate. The various names listed in this table must therefore be employed for
groups containing their appropriate nominal taxon in order of date priority. Thus Mound
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
Table 1 Family-group names in Idolothripinae
Families
IdolothripidaeBagnall, 1908
Hystricothripidae Karny, 19130
Megathripidae Karny, 19130
Pygothripidae Hood, 1915
Subfamilies
Bactrothripinae Karny, 1919
Compsothripinae Karny, 19210
Cryptothripinae Karny, 19210
Macrothripinae Karny, 19210
Diceratothripinae Karny, 1925c
Tribes
Emprosthiothripini Priesner, 1961
Pygidiothripini Priesner, 1961
Subtribes
Allothripina Priesner, 1961
Atractothripina Priesner, 1961
Gastrothripina Priesner, 1961
Zeugmatothripina Priesner, 1961
Hartwigia Stannard, 1976
COMPSOTHRIPINI
Compsothripina [3 genera]
Hartwigia [1 genus]
EMPROSTHIOTHRIPINI [1 genus]
Table 2 Current family-group classification of Idolothripinae (Jacot-Guillarmod, 1978)
IDOLOTHRIPINI
Idolothripina [32 genera]
Atractothripina [1 genus]
Hystricothripina [3 genera]
Megathripina [2 genera]
Apelaunothripina [2 genera]
CRYPTOTHRTPTNT
Cryptothripina [22 genera] PYGIDIOTHRIPINI [1 genus]
Allothripina [6 genera]
Gastrothripina [13 genera] PYGOTHRIPINI [1 genus]
Diceratothripina [18 genera]
Table 3 Revised family-group classification of Idolothripinae
PYGOTHRIPINI Hood, 1915
PYGOTHRIPINA Hood, 1915
Cryptothripinae Karny, 1921a
Emprosthiothripini Priesner, 1961
ALLOTHRIPINA Priesner, 1961
Pygidiothripini Priesner, 1961
COMPSOTHRIPINA Karny, 19210
GASTROTHRIPINA Priesner, 1961
DICERATOTHRIPINA Karny, 1925c
MACROTHRIPINA Karny, 19210
IDOLOTHRIPINI Bagnall, 1908
ELAPHROTHRIPINA subtrib. n.
Hartwigia Stannard, 1976
IDOLOTHRIPINA Bagnall, 1908
Bactrothripinae Karny, 1919
Megathripidae Karny, 19130
HYSTRICOTHRIPINA Karny, 19130
Atractothripina Priesner, 1961
Zeugmatothripina Priesner, 1961
(19740) pointed out that the earliest available name for the spore-feeding thrips is Idolothripi-
dae not Megathripidae which previously had been widely used. Table 2 summarises the
family-group classification of Idolothripinae in current use as given by Jacot-Guillarmod (1978),
and Table 3 gives the family-group classification (with synonymies) adopted in the present
paper.
Only two tribes are recognised by the present authors: Idolothripini in which species never
have metathoracic sternopleural sutures, and (with the exception of many Hystricothripina, as
well as Elaphrothrips antennalis) have two pairs of tergal wing-retaining setae; and Pygothripini
which have only one pair of wing-retaining setae on each tergite (except for two species of
Phaulothrips} and which usually have well-developed metathoracic sternopleural sutures
(except Macrothripina, some Compsothripina and a few Diceratothripina). However, these two
tribes are probably not sister-groups. Idolothripini may be holophyletic, but its only possible
sister-group on present evidence is the sub-tribe Macrothripina in the Pygothripini (Fig. 1).
Despite this, the two tribes are retained here for traditional classificatory convenience.
10
L. A. MOUND AND J. M. PALMER
18 2 3
1 — !• • 1 — II — 1
4
|—|
5
Allothriplna
J-JHjH LJ LJ
18 23
• 1 — 1
LJ
4
1 — 1
5
n
I/H\B-Q — Compsothripina
• LJ
— LJ
LJ
\2/ 2 7
• • Gastrothrirlna
ra i — i
i—i
2 7
nirprntnthrlDlnn
18 2 3
LJ
4
5
Macrothripina
am
4
5
^m
2 3
5
tiapnroinripina
nidolnthrlolnn
4
' " 1
5
6
!• . HvRtrlrnthrlnlnn
- PYGOTHRIPINI
- IDOLOTHRIPINI
1, Maxillary stylets long D ; short •
2, Antennal segment IV with 2 sense cones D ; H sense cones B ; 3 sense cones •
3, Metathoracic sternopleural sutures present D ; absent •
4, Tergltes with 1 pair D; 2 pairs • wing retaining setae
5, Tube glabrous D ; hairy •
6, Praepectal plates present D ; absent •
7, Antennal sense cones slender D ; stout •
8, Maxillary palp sensorla small D ; large •
Fig. 1 Summary diagram of classification and character-state distribution in suprageneric taxa of
Idolothripinae.
Within the Idolothripini three sub-tribes are recognised. Idolothripina and Hystricothripina,
the species of which bear pronounced setae on the margins of the tube, are regarded as
sister-groups, and these two together possibly represent the sister-group of the third sub-tribe,
Elaphrothripina. Priesner's group Atractothripina is synonymised with Hystricothripina, and
Megathripina with Idolothripina. Moreover, Apelaunothripina is recognised as a tribe in the
Phlaeothripinae (p. 89).
The second tribe, Pygothripini, corresponds largely with the group previously known as
Cryptothripini (together with the four other smaller tribes listed in Table 2), with the major
exception of a long series of genera removed from Cryptothripina to the Phlaeothripinae -
Docessissophothripini (q.v., p. 89 & Table 5).
Within the Pygothripini six sub-tribes are recognised in the present re-classification (Table 3).
The Pygothripina, which is characterised primarily by plesiomorphies, probably represents the
closest living approximation to the 'proto-idolothripine' condition: stylets long and close
together medially; antennae often 7-segmented, usually with two sense cones on segments III
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 1 1
and IV; metathoracic sternopleural and anapleural sutures present; pelta with slender, wide
lateral wings. However, within this group there are species with one or other apomorphic
characters found otherwise only in comparatively unrelated, more advanced groups, e.g.
Cleistothrips has the tube hairy as in Idolothripina and Hystricothripina; Cryptothrips has three
sense cones on antennal segment IV as in Gastrothripina and Bolothrips (Compsothripina).
Thus certain of the most advanced characteristics seem to have made their first appearance in
certain of the least advanced genera. The alternative explanation, involving placing Cleistothrips
in Idolothripini on the basis of this one character, the hairy tube, is contrary to the information
provided by other structural characters and makes no sense zoogeographically.
The sub-tribe Allothripina comprises a small group of closely related species, derived from
Pygothripina but with a remarkable terminal sensorium on the maxillary palps. Gastrothripina
includes most idolothripine species with three short, stout sense cones on the fourth antennal
segment. Bolothrips species, here placed in Compsothripina, also have three sense cones on this
segment, but in this genus the sense cones are slender. The Compsothripina, as interpreted here,
comprises a series of grass-living species in Bolothrips and related genera, together with the
ant-mimicking Compsothrips-group. As discussed below (p. 34), intermediates between these
two genera are found in the Mediterranean region. Although both the Gastrothripina and
Compsothripina are each interpreted here as holophyletic groups, their out-group relationships
are unclear.
The Diceratothripina is a large group, comprising those Pygothripini with metathoracic
sternopleural sutures but with the maxillary stylets wide apart and four (rarely two) sense cones
on the fourth antennal segment. This sub-tribe includes a large number of Austro-Oriental and
Pacific species in the Nesothrips-group, a few Neotropical species in the Diceratothrips-group,
together with an ill-defined pan-tropical genus Neosmerinthothrips.
Finally the Macrothripina comprises most of the Pygothripini species which lack metathoracic
sternopleural sutures. This group is almost entirely Oriental with a few species in Africa,
although the two species of Diplacothrips are known only from South America. This is a very
clearly defined group; its relationship to the other Pygothripini is unclear, but Macrothripina
may be the sister-group of the Idolothripini on the basis of the apomorphy- loss of sternopleural
sutures.
Key to subtribes of I dolothr i pinae
N.B. This key is intended only to summarise the relationships discussed above based on the morphological
characters indicated; because of individual variation it is not intended to be a practical key for routine
identifications.
1 Metathoracic sternopleural sutures absent (cf Fig. 99) ; tergites usually with two or more pairs of
wing-retaining setae (except Anactinothrips and Elaphrothrips antennalis) (Figs 294-299)
and/or tube bearing long lateral setae (Fig. 374) (IDOLOTHRIPINI (p. 62)) 2
- Metathoracic sternopleural sutures present or absent (Figs 97-100) ; tergites each with only one
pair of wing-retaining setae (except two species with well-developed sternopleural sutures);
tube never with long lateral setae (except two species with well-developed sternopleural
sutures) (PYGOTHRIPINI (p. 20)) 4
2 Tube without conspicuous lateral setae; metathoracic anapleural sutures complete (Figs 280-
283) ELAPHROTHRIPINA (p. 62)
- Tube with conspicuous lateral setae ; metathoracic anapleural sutures short (Figs 353-355) 3
3 Forewing duplicated cilia well developed; praepectal plates present; tergites each with two pairs
of wing- retaining setae: cf frequently with one or more pairs of lateral abdominal tubercles
(Figs324-325) IDOLOTHRIPINA (p. 71)
- Forewing usually lacking duplicated cilia; praepectal plates usually absent; tergites usually each
with one pair of wing-retaining setae; cf without lateral abdominal tubercles
HYSTRICOTHRIPINA (p. 80)
12 L. A. MOUND AND J. M. PALMER
4 Metathoracic sternopleural sutures absent; antennal segment IV with four sense cones, these
sometimes unusually long; foretibia often with a tubercle near inner apex; head sometimes
with an isolated ommatidium-like structure on each cheek (Figs 194-196)
MACROTHRIPINA (p. 50)
Metathoracic sternopleural sutures usually present, when absent antennal sense cones short or
segment IV with three sense cones 5
5 Terminal sensorium on maxillary palps stout (Fig. 77) ALLOTHRIPINA (p. 30)
Terminal sensorium on maxillary palps not unusually stout 6
6 Maxillary stylets wide apart in head but antennal segment IV with three stout sense cones (Fig.
93) GASTROTHRIPINA(p. 38)
Not this combination of characters, if three sense cones on IV then these are slender or stylets
are close together in the head 7
7 Maxillary stylets wide apart in head, V-shaped; antennal segment IV with four (rarely two)
sense cones DICERATOTHRIPINA (p. 40)
Maxillary stylets rarely more than one-third of head width apart; antennal segment IV with 2, 3
or 4 sense cones 8
8 Eyes frequently prolonged ventrally on head, if eyes not prolonged then antennal segment IV
with three slender sense cones; maxillary stylets not touching medially in head
COMPSOTHRIPINA(p. 34)
Eyes never prolonged ventrally; antennal segment IV with two (rarely three or four) sense
cones; stylets usually close together PYGOTHRIPINA (p. 21)
Genus-group classification of Idolothripinae
The present authors consider a genus, ideally, to comprise a group of species which share a
unique apomorphy and which together represent a holophyletic lineage. Within this phylogene-
tic ideal there seems to us to be no room for the subordinate category of subgenus. We recognise
the value of species-groups in defining evolutionary relationships, but do not accord these any
status in nomenclature.
Unfortunately, for reasons discussed above in the section concerning problems in construct-
ing classifications, the phylogenetic ideal is difficult to achieve. Only half of the described genera
are accepted here (Table 4), mainly because so many monobasic genera have been placed into
larger holophyletic groupings. A more strictly phylogenetic classification would probably
recognise even fewer genera, particularly in the Hystricothripina. But there is a tradition
amongst thysanopterists of considering any unusual character, or character combination, as
meriting recognition at generic level, and we are aware that some of our colleagues would prefer
to maintain this tradition.
This paper is therefore only a preliminary step toward a phylogenetic classification. We hope
that other workers will recognise the value of such an approach, and examine the confused
systematics of the rest of the Phlaeothripidae.
The key below includes all 78 idolothripine genera recognised in this revision (Table 4), and is
based on a study of all those species indicated under each generic name in the main text. Only
one generic name is excluded; the monobasic Pinaceothrips Yakhontov (1956) was not available
for study and is unrecognisable from its description. The key is by no means easy to use, because
of the diversity of species within some genera and the variation found within many species. The
present authors themselves regularly experience difficulty in placing an unknown species to
genus, and under such circumstances frequently have to reinspect the range of variation
exhibited within one or more genera.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
13
Table 4 Generic classification of Idolothripinae
(Pinaceothrips Yakhontov, 1956 is unplaced)
Tribe PYGOTHRIPINI
Subtribe PYGOTHRIPINA
CLEISTOTHRIPS Bagnall
CR YPTOTHRIPS Uzel
EMPROSTHIOTHRIPS Moulton
HEPTA THRIPS Moulton
Ascania Faure syn. n.
Capnothrips Zur Strassen syn. n.
OZOTHRIPSgen. n.
PELINOTHRIPS Mound
PHA ULOTHRIPS Hood
Kaleidothrips Kelly syn. n.
Tetraceratothrips Bagnall
Titanothrips Karny
PRIESNERIANA Ananthakrishnan
PYGOTHRIPSHood
Barythrips Hood & Williams syn. n.
Diplochelaeothrips Moulton syn. n.
Subtribe ALLOTHRIPINA
ALLIDOTHRIPSZur Strassen
ALLOPISOTHRIPSSakimura & Bianchi
ALLOTHRIPSHood
Bryothrips Priesner
FA UREOTHRIPS Priesner
PRIESNERIELLA Hood
Embothrips Dyadechko
Pamllothrips Hood syn. n.
Pygidiothrips Hood syn. n.
PSEUDOCRYPTOTHRIPS Priesner
Subtribe COMPSOTHRIPINA
ANAGLYPTOTHRIPSgen. n.
BOLOTHRIPS Priesner
Bolothrips (Botanothrips) Hood
Boloadelothrips Moulton syn. n.
COMPSOTHRIPS Reuter
Macrothrips Buffa
Leurothrips Bagnall
Leptogastrothrips Trybom
Oedaleothrips Hood
Myrmecothrlps Watson
Formicothrips Priesner
ILLINOTHRIPS Stannard
LOYOLAIA Ananthakrishnan
Subtribe GASTROTHRIPINA
GASTROTHRIPS Hood
Goetothrips Priesner
Isopterothrips Bagnall syn. n.
Paragastrothrips Zur Strassen syn. n.
Percnothrips Ananthakrishnan syn. n.
Pharetrothrips Priesner syn. n.
Probolothrips Moulton
Syncerothrips Hood syn. n.
Subtribe DICERATOTHRIPINA
ACALLUROTHRIPS Bagnall
Diopsothrips Hood syn. n.
CAMPULOTHRIPS Moulton
CARIENTOTHRIPS Moulton
DICERA TOTHRIPS Bagnall
Diceratothrips (Endacnothrips) Priesner
Eulophothrips Schmutz
Megalomerothrips Watson
ELGONIMA Zur Strassen
NEOSMERINTHOTHRIPS Schmutz
Coenurothrips Bagnall
Galactothrips Moulton
NESWIOTHRIPS Mound
NESOTHRIPS Kirkaldy
Oedemothrips Bagnall
Rhaebothrips Karny syn. n.
PHACOTHRIPS Mound
PSEUDOEURHYNCHOTHRIPS Moulton
SPOROTHRIPS Hood
Subtribe MACROTHRIPINA
AESTHESIOTHRIPS Ananthakrishnan
CELIDOTHRIPS Priesner
Ommatidothrlps Mound
DIAPHOROTHRIPS Karny
Diaphorothrips (Cnemidothrips) Priesner
DICHAETOTHRIPS Hood
DIPLACOTHRIPSHood gen. rev.
ETHIROTHRIPS Karny
Decothrips Ananthakrishnan syn. n.
Elaphridia Ananthakrishnan syn. n.
Eurynotothrips Moulton syn. n.
Paracryptothrips Moulton syn. n.
Percipiothrips Ananthakrishnan syn. n.
Scotothrips Priesner syn. n.
Uredothrips Ananthakrishnan syn. n.
HERA THRIPS Mound
ISCHYROTHRIPS Schmutz
MACHA TOTHRIPS Bagnall
Adiaphorothrips Bagnall
Cnestrothrips Priesner
MACROTHRIPS Bagnall
PELTARIOTHRIPSgen. n.
POL YTRICHOTHRIPS Priesner
TARASSOTHRIPSgen. n.
Tribe IDOLOTHRIPINI
Subtribe ELAPHROTHRIPINA subtrib. n.
ANACTINOTHRIPS Bagnall
Lophothrips Karny
Ophidothrips Schmutz
DERMOTHRIPS Bagnall
DINOTHRIPS Bagnall
Paxillothrips Ananthakrishnan
ELAPHROTHRIPS Buffa
Dicaiothrips Buffa
Elaphridothrips Priesner
14
Table4cont.
L. A. MOUND AND J. M. PALMER
Elaphrothrips (Cradothrips)
Ananthakrishnan
Elaphrothrips (Elaphoxothrips) Bagnall
Elaphrothrips (Paraclinothrips) Priesner
Klinothrips Bagnall
Palinothrips Hood syn. n.
HARTWIGIA Faure
LAMILLOTHRIPS Bagnall
Hylothrips Priesner syn. n.
MALESIATHRIPS Palmer & Mound
MECYNOTHRIPS Bagnall
Acrothrips Karny
Dracothrips Bagnall
Kleothrips Schmutz
Kleothrips (Akleothrips) Priesner
Kleothrips (Synkleothrips) Priesner
Phoxothrips Karny
OPHTHALMOTHRIPS Hood
Derothrips Jacot-Guillarmod syn. n.
Fulgorothrips Faure syn. n.
Pyrgothrips Karny syn. n.
TIAROTHRIPS Priesner
Subtribe IDOLOTHRIPINA
BACILLOTHRIPS Buffa
BACTROTHRIPS Karny
Bactrianothrips Bagnall
Bactridothrips Karny syn. n.
Caudothrips Karny syn. n.
Cervothrips Bagnall syn. n.
Eidothrips Bagnall syn. n.
Krinothrips Bagnall
CEUTHOTHRIPS Hood
CYLINDROTHRIPS Moulton
EGCHOCEPHALOTHRIPS Bagnall
IDOLOTHRIPS Haliday
Acanthinothrips Bagnall
LASIOTHRIPS Moulton
MEGALOTHRIPS Uzel
MEGA THRIPS Targioni-Tozzetti
Siphonothrips Buffa syn. n.
MEIOTHRIPS Priesner
Meiothrips (Aculeathrips) Kudo
Subtribe HYSTRICOTHRIPINA
ACTINOTHRIPS Bagnall
Dasythrips Hood syn. n.
A TRA CTOTHRIPS Hood
AZEUGMATOTHRIPSgen. n.
CYPHOTHRIPS Hood
HOLUROTHRIPS Bagnall
HYBRIDOTHRIPS Stannard
HYSTRICOTHRIPS Karny
Zeugmatothripoides Bagnall
NEATRACTOTHRIPSgen. n.
PARACTINOTHRIPSgen. n.
SA UROTHRIPS Hood
ZACTINOTHRIPS Hood
ZEUGLOTHRIPS Hood
ZEUGMATOTHRIPS Priesner
Key to genera of Idolothripinae
1 Abdominal tergite I complete, transversely rectangular, bearing spiracles laterally and 4 pairs
ofsetaesublaterally(Fig.64) ALLIDOTHRIPS(p.30)
Abdominal tergite I reduced to a median pelta which is varied in shape but rarely transverse (cf .
Fig. 63) , rarely bearing setae (Figs 375-384) ; spiracles never associated with pelta 2
2 Tube long with numerous lateral setae (Fig. 24), also metathoracic sternopleural sutures
present and anapleural sutures complete 3
- Not this combination of characters, if tube hairy then metathoracic sternopleural sutures
absent and anapleural sutures short and incomplete 4
3 Head with stylets close together medially and 2 pairs of postocular setae (Fig. 5) [New Zea-
land] CLEISTOTHRIPS(p.22)
- Head with stylets wide apart and V-shaped, dorsal surface with one pair of postocular setae and
one pair of very stout ocellar setae (Fig. 133) [Fiji] CAMPULOTHRIPS (p. 42)
4 Tube usually long with numerous long lateral setae (these setae short in Atractothrips)
(Fig. 374); anapleural sutures short and incomplete (Fig. 353); metathoracic sternopleural
sutures absent 5
- Tube usually short, apparently glabrous or with lateral setae sparse and minute; anapleural
sutures usually complete (Fig. 282) ; metathoracic sternopleural sutures present or absent .... 28
5 Praepectus absent [Neotropics; western Africa] 6
Praepectus present, at least laterally, but often weakly sclerotised [eastern Oriental Region] .... 16
6 Median metanotal setae less than 25 //,m long 7
- Median metanotal setae more than 40 ^tm long, usually very long [Central and South America] 8
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 15
7 Tube convex laterally, with numerous long (70 /urn) setae [Africa] HYSTRICOTHRIPS (p. 84)
- Tube sides parallel to weakly concave , setae weak and scarcely 20 /jun long [Florida]
ATRACTOTHRIPS(part) (p. 82)
8 Abdominal segment I with lateral setae in normal position, arising on anterolateral sclerites,
not associated with pelta 9
- Abdominal segment I with setae arising on lateral extremities of broad pelta (Figs. 376), or on
small sclerites distinct from the anterolateral sclerites (Fig. 375) 12
9 Head with all dorsal setae shorter than distance between two ocelli; compound eyes small and
angular (Fig. 346) ATRACTOTHRIPS(pari) (p. 82)
- Head with at least one pair of elongate dorsal setae, compound eyes well developed and round
(Fig. 340) 10
10 Head with 3 pairs of stout setae dorsally (Fig. 340); pronotal am setae more than 0-5 times as
long as aa setae, pa setae reduced butpm setae large [Mexico] HYBRIDOTHRIPS (p. 84)
- Head with one pair of stout setae dorsally (Fig. 345); pronotal am setae reduced, pm setae
never enlarged 11
11 Antennal segments III-IV with numerous small sense cones near apex ventrally; foretarsal
tooth present in Cf [Peru] ZACTINOTHRIPS (p. 87)
- Antennal segments III-IV without supernumerary ventral sense cones; foretarsal tooth
present or absent in Cf ACTINOTHRIPS (p. 81)
12 Head with maxillary stylets close together medially and retracted to postocular setae (Fig. 342)
[Peru] ZEUGLOTHRIPS (p. 87)
- Maxillary stylets wide apart medially, not retracted so deeply into head 13
13 Abdominal tergites each with 2 pairs of wing-retaining setae (Fig. 368); tube more than 2-5
times as long as head [Brazil] SAUROTHRIPS (p. 86)
- Tergites each with only one pair of wing-retaining setae ; tube at most 2-0 times as long as head 14
14 Micropterous, or forewing with no duplicated cilia.
Cf without a foretarsal tooth ZEUGMATOTHRIPS(p.88)
- Macropterous, forewing with at least two duplicated cilia 15
15 Head with 2 pairs of stout setae dorsally (Fig. 339); antennal segments III-IV without stout
dorsal setae [Brazil] CYPHOTHRIPS (p. 83)
- Head with 3 pairs of stout setae dorsally (Fig. 349); antennal segments III-IV with 2 stout
dorsal setae (Fig. 357) [Trinidad] AZEUGMATOTHRIPS (p. 82)
16 Ventral length of eyes at least twice dorsal length, ventral prolongation of eyes broad (Fig.
341); head prolongation longer than eyes and bearing 2 pairs of setae [Malaya, Japan,
Queensland] HOLUROTHRIPS(p.83)
- Eyes not prolonged ventrally, or ventral prolongation shorter and narrower; head frequently
little extended in front of eyes 17
17 Tergal wing-retaining setae leaf-like (Fig. 369); antennal segment III 0-9 times as long as IV
[Malaya, Philippines] PARACTINOTHRIPS(p.85)
- Tergal wing-retaining setae acute , usually sigmoid , antennal segment III longer than IV 18
18 Mesopraesternum transverse, apparently fused to lateral sclerites (Fig. 372); abdominal
segment I with lateral setae arising anterior to lateral lobes of pelta (Fig. 377); forewing
without duplicated cilia NEATRACTOTHRIPS(p.S5)
- Mesopraesternum boat-shaped, clearly defined from lateral sclerites; pelta different; fore-
wings , when present, with duplicated cilia 19
19 Epimeral sutures complete ; cf (where known) without drepanae laterally on abdomen 20
- Epimeral sutures incomplete ; cf often with lateral drepanae on abdomen
20 Pronotum 2-0 times as broad as long; lateral lobes of pelta broad (Fig. 335) [South Africa]
CYLINDROTHRIPS(p. 75)
- Pronotum more than 2-8 times as broad as long; lateral lobes of pelta slender 21
21 Head with 2 pairs of postocular setae arising in a transverse row; dorsal surface of head grossly
swollen [New Caledonia] EGCHOCEPHALOTHRIPS(p.76)
16 L. A. MOUND AND J. M. PALMER
Head with 2 pairs of postocular setae arising one behind the other (Fig. 322); dorsal surface of
head less elevated [Australia] LASIOTHRIPS (p. 77)
22 Stylets well retracted and lying close together in middle of head (Figs 315, 318); cf with
drepanae on tergite VI 23
Stylets not well retracted, or if well retracted then not close together in middle of head
(Figs 3 14, 321) ;cf with or without drepanae 24
23 Pronotum short (cf. Lasiothrips); pelta with slender lateral lobes; eyes sometimes slightly
prolonged ventrally MEGALOTHRIPS(p.77)
Pronotum longer, only 1-5 times as broad as long; pelta with broad lateral lobes set close to
centre (Fig. 330); eyes usually small, not prolonged ventrally [Europe] . . . BACILLOTHRIPS (p. 72)
24 Stylets retracted far into head (Figs 319, 321); tibiae pale or dark, not bicoloured; wings when
fully developed, pale 25
- Stylets not retracted far into head (Figs 313-314); tibiae usually bicoloured; wings when fully
developed with a darker median basal line 26
25 Tibiae completely dark ; head and pronotal setae short except ocellars and epimerals ; pelta with
slender lateral lobes; abdominal tergites with 1 pair of sigmoid wing-retaining setae; cf
without drepanae on abdomen [Florida] CEUTHOTHRIPS(p.75)
Tibiae completely pale; pelta with broad lateral lobes set close to centre; abdominal tergites
with 1 to 2 pairs of sigmoid wing- retaining setae; cf without or with drepanae only on tergite
VI [Holarctic] MEGATHRIPS(p.78)
26 Lateral lobes of pelta narrowly joined to centre, broadest distal part of lobe sometimes 1-3,
usually more than 2-0 times as long as the narrowest proximal part (Figs 332-333); metanotal
setae 0-6 to 1-6 times as long as the distance between their bases; head 1-4 to 2-2 times as long
as broad; cf drepanae, when present, not bearing a stout terminal seta [Old World Tropics,
southern Europe and California] BACTROTHRIPS(p.72)
Lateral lobes of pelta broadly joined to centre, broadest distal part of lobe 1-1 to 1-4 times as
long as the narrowest proximal part (Figs 334, 336); metanotal setae shorter or longer; cf
drepanae, when present, bearing a stout terminal seta 27
27 Metanotal seta short, 0-3 times as long as the distance between their bases, abdominal tergites
without accessory setae anterior to antecostal line; cf with posterior angles of at least
abdominal tergite VIII produced outwards bearing a spine-like seta; head 1-4 to 2-2 times as
long as broad [Australia] IDOLOTHRIPS*(p.76)
Metanotal setae usually long, twice as long as the distance between their bases; abdominal
tergites with accessory setae anterior to antecostal line; posterior angles of abdominal
tergites of cf sometimes bearing a spine but not produced outwards; head 1-7 to 3-0 times as
long as broad [Indomalaysia] MEIOTHRIPS(p.79)
28 Abdominal tergites each with 2 (or 3) pairs of wing-retaining setae; usually macropterous,
anterior pair of wing-retaining setae sometimes reduced in apterae but rarely absent (eg.
Elaphrothrips antennalis) 29
Abdominal tergites each with only one pair of wing- retaining setae ; frequently apterous 37
29 Head grossly swollen and bearing numerous small setae, constricted to basal neck (Fig. 268);
body ant-like [South Africa] HARTWIGIA (p. 68)
Head and body different 30
30 Eyes prolonged posteriorly on ventral surface of head (Figs 264-267) OPHTHALMOTHRIPS (p. 70)
Eyes scarcely longer ventrally than dorsally 31
31 Small dark brown species with complex sculpture on head (Figs 273 , 274, 276) 32
Large to very large , brown to black species , without complex sculpture on head 33
32 Antennal segment II with 1 or 2 large dorsal setae (Figs 29 1-292) [Java to Micronesia]
MALESIATHRIPS(part) (p. 69)
Antennal segment II without enlarged setae [Hawaii] DERMOTHRIPS(part) (p. 63)
33 Abdominal tergites with three or more pairs of major wing-retaining setae, anterior pair close
to antecostal ridge (Fig. 297); cf usually with one or more tubercles on inner margin of
forefemora MECYNOTHRIPS (p. 69)
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 17
- Abdominal tergites with two pairs of major wing-retaining setae (one pair only in
Elaphrothrips antennalis), although large individuals may have several pairs of sup-
plementary sigmoid setae anterolateral to the major pairs (Fig. 299) ; cf never with a tubercle
on inner margin of forefemora 34
34 Foreocellus arising just posterior to major ocellar setae 35
- Foreocellus arising anterior to major ocellar setae 36
35 Head prolonged to front of eyes, prolongation about four times as long as wide (Fig. 269) ; pelta
not divided into three separate parts TIAROTHRIPS(P-^}
- Head scarcely prolonged in front of eyes; pelta divided into three distinct sclerites (Fig. 310)
DINOTHRIPS (p. 64)
36 Pronotum of cf with anterior angles produced into flattened plates, forefemora without a
sickle-shaped seta apically LAMILLOTHRIPS(p.68)
- Pronotum of cf variable but never produced at anterior angles; cf forefemora often with a
stout sickle-shaped seta ELAPHROTHRIPS (p. 64)
37 Anapleural sutures long but incomplete ; metathoracic sternopleural sutures absent; head with
3 pairs of stout setae dorsally; metanotum and femora with long, stout setae; cf with a
foretarsal tooth, also a stridulating mechanism between posterior angle of forefemora and
ridged surf ace of forecoxae ANACTINOTHRIPS (p. 63)
- Not this combination of characters 38
38 Antennal segment IV with 3 sense cones 39
- Antennal segment IV with 2 or 4 to 5 sense cones 45
39 Maxillary stylets deeply retracted and close together medially in head (Fig. 2) [Holarctic]
CRYPTOTHRIPS(p. 22)
- Maxillary stylets at least one-third of head width apart 40
40 Metathoracic sternopleural sutures absent BOLOTHRIPS(part)(p. 35)
- Metathoracic sternopleural sutures present 41
41 Yellow species; dorsal surface of head reticulate (Fig. 103) [New Zealand]
ANAGLYPTOTHRIPS(p. 34)
- Usually brown species; head not reticulate 42
42 Pelta trilobed (Fig. 109) [India] LOYOLAIA (p. 38)
- Pelta triangular to broadly rounded 43
43 Compound eyes longer ventrally than dorsally (Fig. 94) BOLOTHRIPS(part) (p. 35)
- Compound eyes not prolonged ventrally 44
44 Eyes small, cheeks incut behind eyes; pelta broadly rounded (Fig. 104); tube with margins
straight [North America] ILLINOTHRIPS(p.3S)
Eyes larger (Figs 120, 125); pelta usually triangular (Figs. 122-124, 129); tube frequently
constricted at apex GASTROTHRIPS (p. 38)
45 Metathoracic sternopleural sutures not developed 46
- Metathoracic sternopleural sutures present but variable in length, width and position
(Figs20-22) 64
46 Antennal segment IV with 2 sense cones 47
- Antennal segment IV with 4 sense cones 48
47 Eyes prolonged ventrally on head ; dorsal surface of head weakly sculptured BOLOTHRIPS
(part) (p. 35)
- Eyes not longer ventrally than dorsally; dorsal surface of head strongly sculptured (Fig. 276)
[Hawaii] DERMOTHRIPS (part) (p. 63)
48 Small species, dorsal surface of head with complex sculpture (Figs 67, 273) 49
- Small to very large species, head different 50
49 Yellow species with head reticulate (Fig. 67); antennal segment II without large setae [South
Africa] FAUREOTHRIPS(p.32)
18 L. A. MOUND AND J. M. PALMER
Brown species, head with complex sculpture (Figs 273-274); antennal segment II with 1 or 2
large setae MALESIATHRIPS(part)(p. 69)
50 Maxillary stylets retracted to compound eyes and close together medially in head
(Figs 195-196) ; compound eyes large and round; antennal segment HI shorter than IV 52
- Maxillary stylets often wide apart and retracted only half-way into head, sometimes about
one-third of head width apart and deeply retracted; a few species with stylets closer together
have the eyes reduced and antennal segment III longer than IV 53
51 Antennal sense cones not exceptionally long (Fig. 243); pelta without any sculpture (Fig. 211)
[Borneo] POLYTRICHOTHRIPS(p.6Q)
- Sense cones on antennal segments III-IV long and slender, about half as long as each segment
(Figs 25 1-253); pelta sculptured 52
52 Head elongate with one pair of long ocellar setae (Fig. 200) [India; Malaya]
AESTHESIOTHRIPS(p. 51)
- Head not elongate, without long ocellar setae (Fig. 196) [Malaya] TARASSOTHRIPS(p. 61)
53 Pelta triangular with posterior margin concave, anterior margin of tergite II protruding into
pelta (Fig. 212); antennal sense cones long (Fig. 242) [Malaya; Philippines]
PELTARIOTHRIPS(p. 59)
- Not this combination of characters 54
54 Pelta D-shaped (Figs 225-227) 55
Pelta broader (Fig. 215) 56
55 Antennal sense cones short (Fig. 245); preocellar setae long (Fig. 197) [South America]
DIPLACOTHRIPS(p. 54)
- Antennal sense cones long (Figs 254-256); preocellar setae not elongate (Fig. 199) [Guyana;
South East Asia] DICHAETOTHRIPS(p. 52)
56 Head about twice as long as wide ; cheeks with several stout setae and an isolated ommatidium-
like structure behind eyes; maxillary stylets retracted to postocular setae, one-third of head
width apart (Fig. 201) [Malaya to Australia and Solomon Is.] CELIDOTHRIPS (p. 51)
- Head different, stylets further apart and usually lower in head , or with cheeks different 57
57 Small, usually apterous species with eyes usually prolonged ventrally; one pair of ocellar setae
long (Figs 142-144); $ without fore tarsal tooth [Hawaii, Australia, New Zealand]
NESOTHRIPS(part) (p. 47)
- Not above combination 58
58 Small, usually apterous species with eyes reduced to about 30 facets (50 in macropterae);
cheeks narrowed behind eyes without stout setae; pelta broadly oval; 9 without foretarsal
tooth [Australia, New Zealand] CARIENTOTHRIPS (part) (p. 42)
- Not this combination, usually large dark species 59
59 Pronotum with a hook ventrally at each anterior angle (Fig. 206); forecoxae of cf with large
recurved tubercle, forefemora of cf and $ with stout pale spines on inner surface [New
Guinea] MACROTHRIPS(p.59)
- Not as above 60
60 Fore tibiae with tubercle arising subapically in cf and $ (Fig. 220); foretarsal tooth well
developed [India to Pacific] DIAPHOROTHRIPS(p. 52)
- Tubercle on foretibiae, if present, arising at inner apex, not subapically; cf sometimes without
foretarsal tooth 61
61 Maxillary stylets scarcely retracted into head capsule (Fig. 188) [Australia] HERA THRIPS (p. 58)
- Maxillary stylets retracted at least halfway into head 62
62 Preocellar setae well developed (Fig. 203); fore femora of $ (sometimes cf ) with row of stout
dark tubercles (Figs. 217-219) [India to Micronesia] MACHATOTHRIPS (p. 58)
- Preocellar setae not elongate 63
63 Forefemora of $ with 7 to 10 tubercles on inner margin [India] ISCHYROTHRIPS (p. 58)
- Forefemora of $ without tubercles [Old World Tropics] ETHIROTHRIPS (p. 54)
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 19
64 Black apterous species with head strongly sculptured, broadly produced in front of compound
eyes and constricted basally into a neck (Fig. 18); head and pronotum without long setae;
mesopraesternum absent (Fig. 20) [Australia; New Zealand] EMPROSTHIOTHRIPS (p. 23)
- Not this combination of characters, if black and apterous then pronotal setae long and
mesopraesternum developed 65
65 Maxillary palps with a large terminal sensorium which looks like a third segment (Fig. 77);
small, yellow usually apterous species with 5 to 10 eye facets dorsally; antennae with 7
segments or less (8 segments in Pseudocryptothrips) 66
- Maxillary palps without a single large sensorium terminally; mostly dark species, with eyes
larger and antennae usually 8-segmented 69
66 Antennae 8-segmented, segment IV with 4 sense cones (Fig. 73) ... PSEUDOCRYPTOTHRIPS (p. 33)
- Antennae with 6 or 7 segments; IV with 2 sense cones 67
67 Antennal segment VII broadly joined or fused to VI (Figs 79-83) PRIESNERIELLA (p. 32)
- Antennal segment VII narrower at base than VI at apex 68
68 Antennal segment VII strongly constricted to basal neck (Fig. 78) ; mesonotum well developed
with 1 to 3 pairs of major setae ALLOTHRIPS(p. 31)
- Antennal segment VII scarcely narrower at base than VI at apex; mesonotum fused to
metanotum and with no setae [Hawaii] ALLOPISOTHRIPS(p.31)
69 Maxillary stylets close together medially in head (Figs 2-11) 70
- Maxillary stylets at least one-third of head width apart 76
70 Antennal segment VIII clearly distinct from VII 71
- Antennae 7-segmented, or VII-VIII broadly joined 74
71 Dorsal surface of head strongly sculptured, bearing rows of short broad setae (Fig. 17)
[Australia] PELINOTHRIPS (p. 27)
- Head without numerous short broad setae 72
72 Head elongate, dorsal surface elevated in midline, anterior margin bearing at least one pair of
stout preocellar setae (Figs 7-11) [Australia] PHAULOTHRIPS (p. 27)
- Head different 73
73 Maxillary stylets almost touching and closely parallel medially in head (Figs 3-4); antennal
segment IV with 2 sense cones (Fig. 53) [New Zealand] HEPTATHRIPS(part) (p. 23)
- Maxillary stylets close but not parallel and touching medially (Fig. 16); antennal segment IV
with 4 sense cones [India; Australia] PJUES3MERL4JVA (part) (p. 28)
74 Tube broad, basal or medial width more than 3 times apical width (Fig. 23) PYGOTHRIPS (p. 28)
- Tube more slender and parallel-sided, sometimes constricted at apex 75
75 Maxillary stylets not parallel and touching medially in head (Fig. 16) . . PRIESNERIANA (part) (p. 28)
- Maxillary stylets close together and parallel in head (Figs 3-4) [New Zealand, South Africa,
Saudi Arabia] HEPTATHRIPS(PSirt)(p. 23)
76 Apterous species; colour black, brown or yellow, usually constricted at metathorax with a pair
of chalky white markings producing an ant-like appearance; metathoracic sternopleural
sutures exceptionally long and extending to hind coxae (Fig. 100); antennal segment IV with
2 sense cones; eyes usually prolonged on ventral surface of head (Figs 84-
86) COMPSOTHRIPS(p.36)
- Wings present or absent; metathoracic sternopleural sutures shorter; antennal segment IV
usually with 4 (or 5) sense cones; eyes rarely prolonged ventrally 77
77 Head with one pair of stout ocellar setae arising within ocellar triangle and anterior to posterior
ocelli (Fig. 134) [India, Seychelles, Solomon Is.] NESIDIOTHRIPS (p. 47)
- Ocellar setae not arising in this position within ocellar triangle 78
78 Metanotum reticulate with several pairs of scattered minor setae; all major setae including
postocellars, median metanotals and those on tergite IX with abruptly expanded apices;
epimeral sutures not complete [Africa] ELGONIMA (p. 45)
- Not this combination of characters... 79
20 L. A. MOUND AND J. M. PALMER
79 Head with one pair of preocellar setae more or less conspicuous (Figs 130, 140); species often
large but pronotum usually broad and flat, scarcely thickened medially with am, aa and ml
setae much shorter than epim and pa setae; cf with forefemora enlarged, posterior angle
flattened and extending to a stridulatory file on forecoxae (Fig. 149) 80
- Preocellar setae not larger than ocellars or postocellars; pronotum usually thickened medially
in large species; pronotal anterior setae usually not much shorter than posterior pairs; male
forefemora and forecoxae not developed into a plectrum and stridulatory file 81
80 Foretibiae of male produced into long tubercle which underlies the long, slender foretarsal
tooth (Fig. 147); forefemora not bearing stout spines on inner margin; pterothoracic
anapleural ridge sinuate and ending in a small lateral tubercle; metathorac sternopleural
sutures short (Fig. 150) [south-eastern U.S.A.] SPOROTHRIPS (p. 50)
Foretibiae of cf without apical tubercle; forefemora usually swollen and inner margin bearing
several stout spines; anapleural ridge not extending to a lateral tubercle; metathoracic
sternopleural sutures long [Florida and Texas to Brazil] DICERA TOTHRIPS (p. 43)
81 Antennae 7-segmented or with segments VII- VIII broadly joined 82
- Antennae 8-segmented 83
82 Tube greatly expanded, lateral margins convex (Fig. 161) [widespread in tropics]
ACALLUROTHRIPS (p. 40)
- Tube normal and tapering with margins straight [New Zealand] OZOTHRIPS (p . 24)
83 Head with one ommatidium-like structure on each cheek, situated midway between hind
margin of eye and posterior of head (Fig. 141); tube exceptionally broad with convex
margins, maximum width 5-0 times apical width (Fig. 158); forewing without duplicated cilia
[Brazil]
PHACOTHRIPS (p. 49)
- Cheeks without ommatidium-like structure; tube not so broad; forewing usually with dupli-
cated cilia 84
84 Tube heavy with margins convex and sometimes bearing one or more lateral setal bases
(Figs 153-156) [widespread in tropics] NEOSMERINTHOTHRIPS (p. 45)
- Tube with margins straight and tapering 85
85 Foretarsus of $ with well-developed tooth; tube long and slender [Afrotropical Region]
PSEUDOEURHYNCHOTHRIPS (p. 50)
Foretarsus of $ without a tooth (very small tooth present in N. doulli; moderate tooth present
in N. leveri but this has tube short) [Pacific & Oriental Regions] 86
86 Maxillary stylets usually deeply retracted, extending to postocular setae, usually subparallel
medially and about one-third of head width apart (Figs 135-137), if V-shaped then head
more than 1-3 times as long as wide CARIENTOTHRIPS (part) (p. 42)
- Maxillary stylets wide apart and V-shaped in head, head less than 1-2 times as long as wide
(Figs 142-144) NESOTHRIPS(pari)(p.47)
Tribe PYGOTHRIPINI
As discussed above this tribe corresponds largely to the Cryptothripini of previous authors, with
some minor additions (Tables 2 & 3). The name change is due to recognition of the close
relationship between Cryptothrips and Pygothrips species, and the priority of the family-group
name Pygothripidae (Hood, 1915) over Cryptothripinae (Karny, 19210) (Table 1). The most
important character defining the group is the presence of only a single pair of wing-retaining
setae on each tergite (except for two species of Phaulothrips) . The metathoracic anapleural
sutures are always complete, but the sternopleural sutures may be well developed or absent. The
tube is variable in structure between genera, but in only two species does it bear prominent
lateral setae.
Six sub-tribes are here recognised within the Pygothripini (Table 3) and this group at present
includes 45 genera and 331 species. The sub-tribe Pygothripina includes species with the largest
number of presumably plesiomorphic characters. Sister-group relationships between the sub-
tribes are unclear, but the Macrothripina may be the sister-group of the Idolothripini, and the
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 21
Allothripina the sister-group of the Pygothripina. The Gastrothripina and Compsothripina may
be sister-groups on the basis of the frequent presence of three sense cones on antennal segment
IV; on this assumption they would presumably have been derived from Cryptothrips-\ike
ancestors (Fig. 1). The Diceratothripina differ from the Pygothripina primarily in having the
maxillary stylets farther apart and lower in the head.
Genera of Pygothripina
This group name was proposed by Hood (1915) as Pygothripidae to include one monobasic
genus from Australia characterised by its greatly swollen tube. Hood and subsequent authors
have emphasised for systematic purposes the importance of a swollen tube, but this characteris-
tic has developed in at least two distinct phyletic lines of Idolothripinae (Acallurothrips,
Diceratothrips, Neosmerinthothrips and Phacothrips in Diceratothripina; Phaulothrips and
Pygothrips in Pygothripina) as well as in the phlaeothripine tribe Docessissophothripini
(Holothrips and Symphyothrips) (p. 96).
The species of Pygothrips have many characters in common with the species of Cryptothrips,
and as they are here both placed in the same subtribe the name Pygothripidae must take priority
over Cryptothripinae (see Table 1). Most of the genera placed in this group (as Cryptothripina)
by Priesner (1961) are now referred to the Phlaeothripinae tribe Docessissophothripini (p. 90).
The Pygothripina is used here for a group of 9 genera. Most of these come from New Zealand
and Australia, although Pygothrips has several Neotropical species and Cryptothrips is Holarctic
in distribution. The members of these genera all have two undoubtedly plesiomorphic charac-
ters, the presence of metathoracic sternopleural sutures and complete anapleural sutures.
Moreover, many members of the group, alone amongst idolothripines, have well-developed
maxillary guides. Finally the pelta is broad basally in many species. These characteristics,
together with the zoogeographic distribution, suggest that Pygothripina species are the closest
living representatives to the proto-Idolothripinae.
The New Zealand idolothripine fauna might be expected to be particularly significant when
considering the origins of the Idolothripinae, because ancestral groups of Thripidae have
recently been described from that area (Mound & Palmer, 1981; Mound & Walker, 1982). In
New Zealand there are two groups of idolothripines; a few species of Diceratothripina evidently
derived from Pacific and Australian faunas, and the Heptathrips genus-group (Cleistothrips,
Heptathrips and Ozothrips). This latter genus-group is particularly interesting because, not only
do the members retain the plesiomorphic characters of other Pygothripina but the species show a
wide range of structural diversity. This suggests that the group is relatively ancient.
The diversity within the Heptathrips-group is remarkable. Most of the species have the
antennae 7-segmented (Figs 52-53), but one (?two) species of Ozothrips and two species of
Heptathrips from New Zealand (also two South African species transferred to Heptathrips from
Ascania, and one Saudi Arabian species transferred to Heptathrips from Capnothrips) have the
eighth segment more or less developed (Figs 50-51). Most of the species have only two sense
cones on segments III and IV, but two species of Ozothrips have four sense cones on IV. Again,
most of the species have long maxillary stylets deeply retracted and touching medially (Figs 2-5),
but the stylets are further apart in Ozothrips species (Figs 12-14). Finally, most of the species
lack praepectal plates although these are weakly developed in two species of Ozothrips. It thus
seems logical (although very surprising) to deduce that the 'proto-idolothripine' condition
involved 7-segmented antennae with two sense cones on III and IV, elongate stylets with stout
maxillary guides and absence of praepectal plates.
A further unusual feature of the Heptathrips group is the diversity in form of the tube. In
Cleistothrips the tube is long and hairy (Fig. 24), a condition found otherwise only in the two
most advanced idolothripine groups - Idolothripina and Hystricothripina. The tube is variable
in Heptathrips species from long to very short. One (undescribed) species of Heptathrips is
remarkable for its reticulate sculpture similar to Faureothrips , and Ozothrips janus is equally
unusual because of the ventral prolongation of the compound eyes (Fig. 12). One species of
Ozothrips, described below as eurytis, would probably be placed in the Diceratothripina near
22 L. A. MOUND AND J. M. PALMER
Neosmerinthothrips if it were not for the fact that it is known only from native forest habitats in
New Zealand and fits logically into the pattern of diversity of the Heptathrips-group.
Of the remaining six pygothripine genera Phaulothrips species have elongate stylets as in
Pygothrips, also the head bears a pair of stout ocellar setae as in some species of Pygothrips and
Cryptothrips . The Australian genus Pelinothrips probably shared a common ancestor with
Phaulothrips, and the remarkable Australian genus Emprosthiothrips is here interpreted as an
extreme form on this phyletic line. Finally Priesneriana appears to be related to Cryptothrips but
has the stylets further apart and the compound eyes reduced (Fig. 16). Cryptothrips species are
the only members of Pygothripina with three sense cones on the fourth antennal segment
(except possibly Priesneriana amneius in which this is possibly variable).
CLEISTOTHRIPS Bagnall
(Figs 5, 24, 32, 46)
Cleistothrips Bagnall, 1932: 511. Type-species: Cleistothrips idolothripoides Bagnall, by monotypy.
This genus comprises a single large species from New Zealand which bears a long, hairy tube
similar to that found in members of the distantly related group Idolothripini (Fig. 24). Despite
this tube, and the presence of two pairs of postocular setae (Fig. 5), idolothripoides is very
similar to species of Heptathrips. The median area of the pelta is smaller than in Heptathrips or
Ozothrips, and the lateral wings of the pelta more elongate (Fig. 32). Cleistothrips forms with
these two genera the Heptathrips genus-group which is the essential idolothripine element of the
New Zealand fauna. Contrary to Mound (1968), it is not related to the Docessissophothripini,
because the tergites bear a single pair of wing-retaining setae, the pelta is wide basally, the
maxillary stylets are broad, and antennal segments HI and IV each bear only two sense cones
(Fig. 46).
SPECIES INCLUDED
idolothripoides Bagnall, 1932: 512-3. Holotype 9, NEW ZEALAND (BMNH).
CR YPTOTHRIPS Uzel
(Figs 2, 34, 47)
Cryptothrips Uzel, 1895: 228-9. Type-species: Cryptothrips lata Uzel (a synonym otnigripes Reuter), by
subsequent designation, Hood, 1916: 64.
Jacot-Guillarmod (1978) lists 24 species under Cryptothrips. However, breviventris Hood
(1927fl) is a phlaeothripine and should be placed provisionally in Hoplothrips (teste Steve
Nakahara in lift.); okamatoi Karny (1913c) is a Phlaeothrips species with long postocular setae;
daedalus Karny (1912b) apparently represents a Psalidothrips species with postocular setae
arising far behind the eyes. (Type-material of the latter two species has been studied on loan
from the Humboldt University Museum, Berlin.) Moreover, the following seven species also
belong in the Phlaeothripinae but in undetermined genera: additamentus Karny, bursarius
Karny, longicaput Girault, nigronympha Girau\l,pusillus Karny, rufiprothorax Girault, schilleri
Girault.
Two species described in Cryptothrips from Australia, pygus and shavianus, are here
transferred to Pygothrips; three further Australian species, amneius, laticeps and uptoni, are
transferred to Priesneriana; constans is now regarded as a synonym oiNesothrips niger, and latus
var. fijiensis is a synonym of Ethirothrips hibisci. Judging from the original description, sauteri
may belong in Pygothrips, but the description offlavus is too inadequate for recognition. Finally,
maritimus was described as a predator of Scolytid beetle larvae, and judging from the original
illustration may well belong in Liothrips.
Cryptothrips thus appears to be Holarctic in distribution, and it may be the sister-group of the
Indo-Australian genus Priesneriana. The latter has reduced eyes which are directed forwards
(Fig. 16), and four sense cones on the fourth antennal segment (Fig. 48). In contrast Cry/?-
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 23
tothrips species have three sense cones on the fourth segment (Fig. 47). Both genera are similar
to Pygothrips in having broad maxillary stylets deeply retracted and close together in the middle
of the head (Figs 2, 6), and the metathoracic sternopleural sutures are well developed.
SPECIES INCLUDED
*angusfusUzel, 1895: 231-2. Holotype $, CZECHOSLOVAKIA: Bohemia (? lost).
carbonariusHood, 19080: 376-7. Holotype cf , U.S.A.: Illinois (USNM).
longiceps Hood, 1912c: 153-4. Holotype ?, U.S.A.: Illinois (USNM).
*/favusSolowiow, 1924: 24. Types not indicated, U.S.S.R. (? lost).
*/naritimusDjadetschko, 1962: 764-5. Holotype $, U.S.S.R.: Ukraine (not known).
n/gripes(Reuter, 1880: 11) (Phloeothrips) . Holotype ? $, FINLAND (? lost).
lata Uzel, 1895: 230-1. Syntypes cf $, CZECHOSLOVAKIA: Bohemia (? lost).
major Bagnall, 1911: 60-1. Holotype 9, NORWAY (BMNH).
latus f. breviceps Maltbaek, 1929: 372. Types not designated, DENMARK (? lost).
williamsi Bagnall, 19330: 120-1. Holotype cf, GREAT BRITAIN: England (BMNH).
nigripesphariacusTitschack, 1965: 147, replacement name for msw/araTitschack, 1964: 51. Syntypes
Cf?,YUGOSLAVIA(SMF).
recfcmgu/ar/sHood, 1908ft: 307-9. Lectotype $, U.S.A.: Illinois (USNM).
salicis (Watson, 1921: 80-1) (Trichothrips). Syntypes cf$, U.S.A.: New York (FSAC).
*sauteriKarny, 1913c: 1278. Holotype? $, TAIWAN (unknown).
*sordidatusHood, 1927ft: 199. Lectotype $, U.S.A.: California (USNM).
EMPROSTHIOTHRIPS Moulton
(Figs 18, 19,20,31,55)
Emprosthiothrips Moulton, 19420: 12. Type-species: Emprosthiothrips niger Moulton, by monotypy.
This genus comprises six dark, apterous species from Australia which are remarkable for the
shape of their antennae (Fig. 55), and for their reduced setae and fused sclerites. As a result of
these peculiarities the systematic position of the genus has been in doubt. Priesner (1961) placed
it in a monobasic tribe, but Mound (1974a) suggested a relationship to Dermothrips and
Pelinothrips 'in the Cryptothripini'. Dermothrips is here referred to the Elaphrothripina, but
Pelinothrips and Emprosthiothrips probably represent together an Australian derivative from
early Pygothripina ancestors. The stylets are further apart than in other Pygothripina species (in
brimblecombei they are very short and wide apart), but the praepectal plates and metathoracic
sternopleural sutures are well developed, antennal segment IV bears two sense cones, and
segments VII- VIII are broadly joined. The species of this genus are found at the bases of grass
tussocks; four of them have the eyes prolonged ventrally as in some species of Bolothrips and
Carientothrips in the same habitat.
SPECIES INCLUDED
bogong Mound, 1969: 185. Holotype $, AUSTRALIA (ANIC).
brimblecombei Mound, 19740: 51-2. Holotype <j>, AUSTRALIA (ANIC).
brittoni Mound, 1969: 186. Holotype cf , AUSTRALIA (ANIC).
cs/ro Mound, 1969: 185-6. Holotype $, AUSTRALIA (ANIC).
epallelus Mound, 19740: 52-3. Holotype cf , AUSTRALIA (ANIC).
niger Moulton, 19420: 12-3. Holotype $, AUSTRALIA (CAS).
HEPTA THRIPS Moulton
(Figs 3, 4, 33, 53)
Heptathrips Moulton, 19420: 3. Type-species: Heptathrips tonnoiri Moulton, by monotypy.
Ascania Faure, 19540: 17-20. Type-species: Ascania magnified Faure, by original designation. Syn. n.
Capnothrips Zur Strassen, 1979: 99. Type-species: Capnothrips ruficaudis Zur Strassen, by monotypy.
Syn. n.
Only one species was originally described in this genus, but a further four undescribed species
from New Zealand have been studied; these will be treated in detail in an account of the
24 L. A. MOUND AND J. M. PALMER
Phlaeothripidae of New Zealand currently in preparation. One species is similar to tonnoiri but
with a longer tube. The other three are apterous; one is pale and strongly reticulate, one has a
short tube and antennal segment VIII defined by a suture, the third has a stout tube and segment
VIII clearly separated from VII. As discussed above, this remarkable structural radiation
suggests the group has been long established in New Zealand. The species all have the stylets
close together in the head with well-developed maxillary guides (Figs 3, 4), a broadly based pelta
(Fig. 33) and well-developed metathoracic sternopleural sutures.
The two species from South Africa described in Ascania cannot at present be distinguished
from Heptathrips. The heavy tube with constricted apex found in magnified is longer but
otherwise similar to that of one of the undescribed New Zealand species referred to above.
These African species have the preocellar setae stouter, the metathoracic sternopleural sutures
weaker, and the tergal wing-retaining setae straighter than the New Zealand species. The single
species described in Capnothrips is very similar to africana but has the inner sense cone on
antennal segment III much shorter, scarcely one-third as long as the outer sense cone.
SPECIES INCLUDED
africana (Moulton, 1949: 491-2) (Adelothrips). Holotype $, SOUTH Africa (BMNH). Comb, n:
magnifies (Faure, 1954a: 20-3) (Ascania). Holotype $, SOUTH AFRICA (NCIP). Comb. n.
ru/icaudis(ZurStrassen, 1979: 99-101) (Capnothrips). Holotype $, SAUDI ARABIA (NMB).Comb. n.
tonnoiri Moulton, 1942a: 3-4. Holotype $, NEW ZEALAND (CAS).
OZOTHRIPS gen. n.
Type-species: Ozothrips priscussp. n.
Small to large, brown species of Pygothripini. Antennae 7-segmented or with VII- VIII closely joined; III
with two sense cones, IV with two or four sense cones. Head usually slightly longer than wide, eyes large;
maxillary stylets broad, retracted almost to postocular setae and about one-third of head width apart,
maxillary guides stout; mouth cone broadly rounded, maxillary palps stout. Pronotum transverse,
relatively longer in large C? with median thickening, epimeral sutures complete; praepectus present or
absent; probasisternum large, mesopraesternum broadly boat-shaped, or both sclerites eroded. Foretarsi
with a large tooth in cf , with or without a small tooth in $. Fore wing broad, almost parallel-sided, with
duplicated cilia. Metathoracic sternopleural sutures well developed, anapleural sutures complete. Pelta
with broad lateral wings; tergite II eroded laterally; tergites II- VII each with one pair of weakly sigmoid
wing- retaining setae; tergite IX with three pairs of slender setae in both sexes; sternites with one row of
small discal setae, reticulate sculpture more evident in cf than in $ .
The type-species of this new genus is abundant and widespread on dead leafy twigs of
Nothofagus in New Zealand. The fact that it is found only in native forest areas is good evidence
that it is itself endemic to that country. Most of the characters listed above are shared with
Heptathrips and Cleistothrips , although the widely separated stylets of Ozothrips are regarded as
an apomorphy achieved independently of the similar stylet arrangement found in many other
less closely related idolothripines. Two of the species described below have four sense cones on
antennal segment IV, they lack a foretarsal tooth in the female, and praepectal plates are
developed. These species are thus convergent in structure on members of the widespread
tropical genus Neosmerinthothrips , but with the tube slender and antennal segments VII-VIII
broadly joined or fused. In the absence of evidence to the contrary, it seems sensible to regard
them as endemic but aberrant members of the New Zealand fauna.
Key to Ozothrips species
1 Antennal segment IV with 2 sense cones (Fig. 51); praepectus absent; $ with well-developed
foretarsal tooth; macropterous priscussp. n. (p. 26)
- Antennal segment IV with 4 sense cones (Figs 50, 52); praepectus present; $ with no foretarsal
tooth ; usually apterous 2
2 Antennal segments VII-VIII forming one unit but with suture complete (Fig. 50); eyes
narrowed but not elongate ventrally (Fig. 13); tergite IX setae B\ about half as long as tube
eurytissp. n. (p. 25)
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 25
- Antennal segments VII- VIII without a suture (Fig. 52) ; eyes narrowed and prolonged ventrally
(Fig. 12); tergite IX setae longer than tube janussp. n. (p. 26)
Ozothrips eurytis sp. n.
(Figs 13, 42, 50)
Macropterous $ . Colour mainly brown, inner apex of femora yellowish, also base and external margin of
antennal III; forewings uniformly shaded; setae dark.
Head with postocular setae arising close to eyes, ocelli far apart, postocellar setae small, eyes not large
(Fig. 13). Distal antennal segments with clearly defined pedicels, VII-VIII broadly fused but with suture
usually complete, III with 2 sense cones, IV with 4 sense cones (Fig. 50). Pronotum transverse, epimeral
sutures complete; praepectal and probasisternal plates weakly sclerotised, mesopraesternum and
mesoeusternum broadly eroded medially. Mesonotal midlateral setae minute; metanotal median setae
close together, sternopleural sutures short and broad. Forewing parallel-sided, 5 to 8 duplicated cilia. Pelta
broad basally but weakly and irregularly sclerotised (Fig. 42). Tergite IX setae acute. Tube very slightly
constricted apically and in basal third. Sternites with 3 to 9 discal setae.
Measurements (holotype $ in /x,m). Body length 1850. Head, length 200; width 190; postocular setae 70;
postocellar setae 20. Pronotum, length 120; width 240; major setae - am 20, aa ?, ml 30, epim 75, pa 60.
Forewing, length 650; median width 65; sub-basal setae 25. Tergite IX setae 70, 75, 120. Tube, length 150;
terminal setae 130. Antennal segments III- VII+VIII length 65, 62, 58, 55, 55.
Micropterous $ . Body brown, head and tube darkest; leg colour similar to macropterae but distal half of
femora sometimes yellow; antennal segments I-III sometimes paler but more or less shaded particularly at
apices.
Head with ocelli reduced or absent, postocellar setae slightly longer than in macropterae. Mesonotum
with small round lobes laterally but axillary sclerites absent. Praepectus small; pterothoracic sternites
eroded; pelta variable but broad basally; tergal wing-retaining setae absent.
Measurements (paratype $ collected with holotype in /am). Body length 2200. Head length 230; width
205; postocular setae 80; postocellar setae 30. Pronotum, length 160; width 270; major setae - am 27, aa ?,
ml 45, epim 90, pa 50. Tergite IX setae 90, 85, 120. Tube length 170. Antennal segments III-VII -I- VIII
length 75, 70, 63, 57, 73.
Apterous cf . Similar to 9 but pronotum and forefemora enlarged; foretarsal tooth as long as tarsal width
in large individuals; tube slightly constricted medially; sternites IV- VI reticulate anterior to discal setae
(except gynaecoid cf). Measurements (paratype cf collected with holotype in /Am). Body length 1700.
Head, length 195; width 170; postocular setae 90; postocellar setae 35. Pronotum, length 170; width 240;
major setae - am 30, aa 55, ml 90, epim 120, pa 90. Tergite IX setae 75, 65, 80. Tube length 130. Antennal
segments III- VII+VIII 60, 60, 57, 50, 60.
SPECIMENS STUDIED
Holotype $ macroptera, New Zealand: North Island, Wattle Bay near Auckland, on dead twigs and
branches, 23. ii. 1979 (L. A. Mound 1349) (NZAC).
Paratypes (2 $ mac. , 19 $ mic. , 7 cf apt.). New Zealand. North island: 5 $ , 2 cf collected with holotype;
Auckland, Mt Albert, 1 ?, 18.xi.1978, 1 $ mac., 16.xii.1978 (A. K. Walker);Te Aroha, 1 $ mac. 6 <j>, 2 cf
on dead twigs and branches, 14. ii. 1979 (L. A. Mound 1447, 1452); Coromandel Peninsula, Coroglen
Saddle, 2 $, 1 cf on dead branch, 13.ii.1979 (L. A. Mound 1443). South Island: Nelson, 1 $ from Thrush
nest, 6.xii.l967 (B. S. Gourley); 2 ml north of Reefton, 1 cT, 6.H.1979 (L. A. Mound 1411); Glenorchy
State Forest, Dart River, 1 $, 1 cf , 21.i.l981 (Valentine & Noyes); Invercargill, 1 $ on Rhododendron,
18.iv.1977 (A. K. Walker). Chatham Islands: South East Island, 1 $, 9.xi.l970 (/. Townsend); Chatham,
Waitangi, 1 $, 10.ii.1967 (G. W. Ramsey) (NZAC & BMNH).
Material excluded from paratype series. New Zealand. South Island: Kaihoka Lake, 10 ml west of
Collingwood, 1 $ mac., 5 $ mic., 5 cf apt. ondeadfrondofRhopalostylissapidia, l.ii.!979(L. A. Mound
1392).
COMMENTS. The series of specimens collected on a dead palm frond at Kaihoka Lake is
remarkable in that the micropterae and apterae, but not the macropterae, have antennal
segments II-III almost clear yellow, and the legs more extensively pale than typical specimens of
eurytis. In addition, six specimens have been studied from three widely separated areas of the
North Island of New Zealand (Auckland, Rotorua, Levin) with the tube clear yellow medially.
These six specimens probably represent a further new species. O. eurytis is widespread in New
Zealand, but has only been taken in native forest areas. This reinforces the impression that the
26 L. A. MOUND AND J. M. PALMER
species is related to the larger and more common species, priscus, described below, and that
they, together with janus, constitute an endemic New Zealand genus. Both eurytis and janus
(together with the species with the tube yellow referred to above) have praepectal plates unlike
priscus, the type-species of this genus. One apterous female of eurytis collected with the
holotype bore about 10 specimens of a mite species belonging to the genus Adactylidium
(Pyemotidae).
Ozothrips janus sp. n.
(Figs 12, 40, 52)
Apterous $. Colour brown, foretarsi yellow, extreme apex of forefemora and median area of foretibiae
yellowish, also extreme base of antennal segment III and apex of II; tube golden yellow with dark brown
apex; major setae weakly shaded with long fine apices.
Head about as wide as long, ocelli absent, ocellar setae elongate; eyes greatly prolonged ventrally with
two rows of large ommatidia (Fig. 12); maxillary stylets apparently wide apart, V- or U-shaped (Dis-
organised in available specimens). Antennae with segments pedicillate; 2 sense cones on III, 4 on IV, VIII
completely fused to VII (Fig. 52). Foretarsus with inner margin slightly thickened. Praepectus weak;
pterothoracic sternites heavily eroded; meso- and metanota small and transverse. Pelta broad and
flattened (Fig. 40); lateral abdominal setae elongate; wing-retaining setae short and straight; tube short
and constricted apically. Sternites with few discal setae, holotype with no discal setae on II-III.
Measurements (holotype $ in /xm). Body length 1500. Head, length 150; width 170; postocular setae
120; postocellar setae 60. Pronotum, length 120; width 240; major setae - am 20, aa 55, ml 105, epim 150,
pa 120. Tergite IX setae 135, 135, 120. Tube, length 110; terminal setae 120. Antennal segments III-VII
length 40, 45, 43, 43, 55.
SPECIMENS STUDIED
Holotype $, New Zealand: North Island, Hauraki Gulf, Noises Islands, Otata I., in seed heads of
Ghania, l.xii.1979 (A. K. Walker} (NZAC).
Paratype. 1 $ collected with holotype (BMNH).
COMMENTS. The remarkable ventral prolongation of the eyes in this species suggests that its
normal habitat is at the base of grasses. Despite this character, janus is very similar to eurytis,
although more apteriform.
Ozothrips priscus sp. n.
(Figs 14, 41, 51)
Macropterous $ . Dark brown, tarsi slightly paler; pedicels of antennal segments III-V yellow to light
brown; major setae brown, terminal setae of tube darkest; forewings strongly shaded.
Head with eyes slightly smaller ventrally than dorsally, postocular setae finely acute (Fig. 14); maxillary
stylets broad (10-12 /xm). Antennae with 7 segments, VI broadly truncate at apex, pedicels of VI-VII
narrow, III-IV each with 2 long sense cones (Fig. 51). Praepectus absent, probasisternum large,
mesopraesternum broadly boat-shaped. Forefemora slender; foretarsal tooth small and curved at inner
apical margin. Mesonotal midlateral and metanotal median setae moderately developed (50 mm).
Forewing with 4 sub-basal setae. Pelta broadly rounded medially (Fig. 41). Tergite II posteroangular setae
not developed (Fig. 41).
Measurements (holotype $ with smallest paratype $ in yam). Body length 3700 (3100). Head, length 390
(345); median width 300 (280); postocular setae 120 (96). Pronotum, length 210 (160); median width 420
(370); major setae - am 40 (40), aa 43 (40), ml 75 (60). epim 120 (120), pa 75 (66). Forewing, length 1500
(1300); distal width 150 (120); sub-basal setae 22, 65, 105, 210 (?, 60, 105, 200); number of duplicated cilia
35 (28). Tergite IX setae 240, 330, 300 (210, 300, 300). Tube, length 400 (320); longest terminal setae 210
(200). Antennal segments III-VII length 130, 110, 100, 85, 105 (110, 90, 90, 75, 85).
Macropterous cf . Colour and structure very similar to $; large males with a small tubercle ventrally on
frons (approximately underlying posterior ocelli), also with pronotal midlateral setae elongate, forefemora
enlarged, foretarsal tooth broad and about as long as tarsal width, and antennal segment III relatively long.
Measurements (large and small paratype cf - LAM 1372; LAM 1407 - in /am). Body length 3400 (2800).
Head, length 390 (345); median width 260 (240); postocular setae 165 (120). Pronotum, length 315 (180);
median width 440 (345); major setae - am 40 (25), aa 75 (30), ml 170 (60), epim 120 (130), pa 110 (60).
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 27
Forewing, length 1550 (1300). Tergite IX setae 270, 320, 300 (200, 270, 255). Tube length 380 (300).
Antennal segments III- VII lengths 145, 120, 105, 85, 90 (115, 90, 90, 80, 90).
SPECIMENS STUDIED
Holotype 9 , New Zealand: South Island, Whangamoa Saddle near Nelson, on dead leaves and branches
ofNothofagus, 27. i. 1979 (L. A. Mound 1359) (NZAC).
Paratypes (93 $ 40 cf): New Zealand. South Island: 10 $, 5 cf data as for holotype, 7 $ similar data
except LAM 1357; Nelson, 17 $, 6 cf on dead twigs and leaves, 28. i. 1979 (LAM 1367); Nelson, Dun Mt
4 $ , 4 cf , 29.U979 (LAM 1372); Nelson, Cobb Reservoir, 4 $, 1 cf , 31.1.1979 (LAM 1381); Nelson, Lake
Rotoiti, 24 $, 14 cf , 4-8.ii.1979 (L. A. Mound & A. K. Walker); Marlborough Sounds, Tennyson Inlet &
Opouri Saddle, 16 $, 8cf on dead leafy branches, 28. i. 1979 (A. K. Walker 3, 6); 20 ml NE. of Reefton, 2 $
on dead Nothofagus branches, 6.ii.l979 (LAM 1716). North Island: 20 ml S. of Turangi, 7 <j>, 2 cf on dead
Nothofagus branches, 19.ii.1979 (LAM 1478) (NZAC & BMNH).
COMMENTS. Bothpriscus and eurytis are variable in body size with several characters subject to
allometric growth particularly in males. O. priscus has been collected only on dead Nothofagus
branches and twigs which presumably support a particular fungus to which the thrips is specific.
PELINOTHRIPS Mound
(Figs 17, 38, 39, 54)
Pelinothrips Mound, 1974o: 75-6. Type-species: Rhopalothrips ornatus Girault, by original designation.
Two species, both Australian, are placed in this genus. These resemble Cryptothrips species in
having long maxillary stylets deeply retracted into the head with maxillary guides weakly
indicated medially. The metathoracic sternopleural sutures are short, the tergites each bear a
single pair of wing-retaining setae, and praepectal plates are present. However, antennal
segment IV only bears two sense cones (Fig. 54), and both sexes have a foretarsal tooth. This
genus is probably related to the Australian genus Emprosthiothrips in which the species have the
maxillary stylets slightly further apart.
SPECIES INCLUDED
brochotus Mound, 1974a: 76-7. Holotype $, AUSTRALIA (ANIC).
ornat us (Girault, 1930: 1) (Rhopalothrips). Holotype $, AUSTRALIA (QMB).
PHA ULOTHRIPS Hood
(Figs 7-1 1,25-30, 43-45)
Phaulothrips Hood, 19186: 146-7. Type-species: Phaulothrips vuilleti Hood, by monotypy.
Titanothrips Karny, 1920c: 44. Type-species: Titanothrips portentosus Karny, by monotypy. [Synonymised
by Mound, 19740:78.]
Tetraceratothrips Bagnall, 1924: 628. Type-species: Tetraceratothrips agrestis Bagnall, by monotypy.
[Synonymised by Mound, 1974a: 78.]
Kaleidothrips Kelly, in Kelly & Maine, 1934: 73. Type-species: Kaleidothrips inquilinus Kelly, by
monotypy. Syn. n.
A revision of this genus by Mound (1974a) included nine species, all of which are Australian
although specimens of vuilleti have now been studied from Tanzania. In addition, the single
species in Kaleidothrips is here interpreted as an aberrant member of Phaulothrips, undescribed
species from New Guinea and Fiji are referred to by Mound (19740: 81) and Docessis-
sophothrips magnificus Bianchi from Samoa is also here placed in Phaulothrips. This latter
species has the pelta typical of the genus (Fig. 28), a pair of slender preocellar setae, and only
one pair of wing-retaining setae on each tergite. However, the pair of postocular setae found on
the cheeks of the other Phaulothrips species arise dorsally behind the dorsal postocular setae in
magnificus (Fig. 10), and the head is very strongly elevated medially as figured by Bianchi
(1953). All Phaulothrips species have long curved metathoracic sternopleural sutures which
arise close to the mesothoracic border. Antennal segments III and IV each bear two sense cones
(Fig. 45), but due to the dark colour of the head long, curved maxillary guides have been
28 L. A. MOUND AND J. M. PALMER
observed in only a few specimens. P. agrestis and uptoni are unusual in having two or more pairs
of sigmoid wing-retaining setae (Fig. 44).
SPECIES INCLUDED
agresf/s(Bagnall, 1924: 628-9) (Tetraceratothrips). Holotype cf , AUSTRALIA (BMNH).
aiJici Mound, 1974a: 82-3. Holotype $, AUSTRALIA (ANIC).
barretti Mound, 1974a: 83. Holotype cf , AUSTRALIA (ANIC).
caudafusBagnall, 1932: 510-1. Holotype $, AUSTRALIA (BMNH).
fuscus Moulton, 1935: 100. Holotype $, AUSTRALIA (CAS).
punctatus Rayment, 1948: 257-8 (Cladothrips). Syntypes cf ?, AUSTRALIA (ANIC).
inquilinus (Kelly, 1934: 73) (Kaleidothrips) . Holotype $, AUSTRALIA (ANIC). Comb. n.
longitubusGirault, 1928: 2. Holotype ?$, AUSTRALIA (QMB).
magfli/Jcus(Bianchi, 1953: 106-7) (Docessissophothrips). Holotype $, SAMOA (BPBM).Comb. n.
sibylla Mound, 1974a: 84-5. Holotype $, AUSTRALIA (ANIC).
uptoni Mound, 1974a: 85-6. Holotype $, AUSTRALIA (ANIC).
vu///etf Hood, 19186: 147-8. Holotype cf , AUSTRALIA (USNM).
portentosus Karny, 1920c: 40-4 (Titanothrips) . Holotype cf , AUSTRALIA (NRS).
PRIESNERIANA Ananthakrishnan
(Figs 16, 48)
Priesneriana Ananthakrishnan, 19566: 138. Type-species: Gnophothrips kabandha Ramakrishna, by
monotypy.
This genus was erected for a single species, from southern India, which has the general
appearance of an Ethirothrips species (particularly to those species previously placed in
Uredothrips), but which possesses well-developed metathoracic sternopleural sutures. The
maxillary stylets of kabandha are retracted to the compound eyes, but although arched towards
each other medially they are about one-fifth of the head width apart (Fig. 16). The two
Australian species here transferred to this genus have stylets rather similar to kabandha, but
amneius (from New Guinea, Australia and New Zealand) has the stylets meeting medially.
These three species resemble kabandha in having the eyes reduced and directed forwards,
although uptoni is unusual in that antennal segments VII- VIII are partially fused. Priesneriana
is closely related to Cryptothrips but has four sense cones on antennal segment IV (Fig. 48), and
the eyes are reduced in size.
SPECIES INCLUDED
amneius (Mound, 19740: 42) (Cryptothrips). Holotype $, NEW GUINEA (ANIC). Comb. n.
kabandha (Ramakrishna, 1928: 293-4) (Gnophothrips). Holotype cf, INDIA (TNA).
7af/ceps(Hood, 19186: 142-3) (Cryptothrips). Holotype $, AUSTRALIA (USNM). Comb. n.
uptoni (Mound, 1974a: 44-5) (Cryptothrips). Holotype $, NORFOLK Is. (ANIC). Comb. n.
PYGOTHRIPS Hood
(Figs 6, 21-23, 36-37, 49)
Pygothrips Hood, 1915: 49-50. Type-species: Pygothrips rugicauda Hood, by monotypy.
Barythrips Hood & Williams, 1915: 134-5. Type-species: Barythrips sculpticauda Hood & Williams, by
monotypy. Syn. n.
Diplochelaeothrips Moulton, 1944: 284. Type-species: Diplochelaeothrips mikrommatos Moulton, by
monotypy. Syn. n.
This genus was based originally on a single apterous female collected in Queensland, Australia.
Hood (1915) figured the terminal segments of the antenna and abdomen, and Mound (19740)
gave outline drawings of the head, stylets and pelta. This species has never been collected again,
although a damaged apterous female which may be conspecific has been taken recently near
Adelaide, South Australia. Subsequently, 19 species have been described in Pygothrips, but
re-examination of all but five of these has indicated that more than one phylogenetic group is
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 29
involved. Moreover, many of the species are known only from single samples or even single
individuals, and so no concept of intraspecific variation has been applied within the group.
The type-species of Pygothrips has the maxillary stylets elongate and close together in the
middle of the head (cf Fig. 6). In contrast, most species described in the genus have the stylets
low and wide apart in the head, and all such species are here referred to the genus Acalluro-
thrips. The two genera exhibit a series of characters in common in addition to the swollen tube,
but several characters are probably functionally related to the shared habit of raising the tube
over the head, thus producing an almost spherical mite-like body outline. For example, the
sternites are longer than the tergites, and the pelta, metathorax and antennae are reduced. The
functional significance of this behaviour is not obvious, but appears to be defensive.
The species here referred to Pygothrips share the following characters. Head longer than
wide, stylets long and close together medially (Fig. 6). Antennal segments VII-VIII broadly
joined; sense cones often lateral in position, 2 on III, 4 (rarely 2) on IV (Fig. 49). Pronotal
epimeral sutures complete or incomplete, praepectus present or absent; mesopraesternum
eroded; metathoracic sternopleural sutures present but often broadly eroded (Figs 21-22), also
anapleural sutures. Foretarsal tooth usually present in both sexes. Forewing with or without
duplicated cilia. Pelta eroded at posterior margin (Figs 36-37); median sternites longer than
tergites. Wing-retaining setae often weak; tube expanded to greatly expanded with convex
margins, often ridged and constricted apically (Fig. 23).
In addition to rugicauda three species have been studied (albiceps, fortis and satanas) which
were described in Pygothrips and which agree with the above definition of the genus. Moreover,
five further species which have not been studied are retained in the genus on the basis of their
original descriptions together with information kindly provided by Steve Nakahara that the
types all have stylets deeply retracted and close together medially. Two species described
recently in Cryptothrips are here transferred to Pygothrips because of the swollen tube and
presence of four sense cones on the fourth antennal segment. Finally, the type-species of both
Barythrips and Diplochelaeothrips are also referred to Pygothrips (N.B. - B. grandicauda
belongs in Neosmerinthothrips q.v.; B. mathuri Ananthakrishnan, 19610 belongs in the
phlaeothripine genus Hoplothrips, teste Prof. T. N. Ananthakrishnan in litt.}.
Barythrips sculpticauda was based on one oedymerous male which has been studied and
compared with macropterous females and four, small apterous males from Florida. These all
have the same remarkable antennal colour with segments I-II yellow and III-V yellow except
for a median dark area on the pedicel. The head and stylets are similar to rugicauda although the
pelta of the males is almost divided into two parts (Fig. 37). Moreover, the epimeral sutures are
usually just complete, the forewings bear 7 to 10 duplicated cilia, and the female lacks a
foretarsal tooth.
D. mikrommatos has the epimeral sutures complete, and the forewings bear duplicated cilia.
Moreover, this species is sexually dimorphic, the male being unlike other males in Pygothrips.
The antennal sense cones of albiceps arise laterally (as in Acallurothrips spinicauda) whereas in
fortis they arise ventrally , although the head of these two species is similar with a pair of elongate
postocellar setae (Fig. 6). The sense cones of rugicauda are short and weak, but those of
mikrommatos, satanas and sculpticauda are short and stout. Two macropterous females from
Java (in BMNH) have been studied which appear to belong in Pygothrips, but these have
well-developed foretarsal teeth, about 14 forewing duplicated cilia and the tube, although large,
is not rugose; a similar species from Obi Island lacks the foretarsal teeth. Finally, a male has
been studied from Singapore which is similar to rugicauda females, but has the tube much less
enlarged.
Pygothrips as defined here is closely related to Cryptothrips. However, these tropical species
have four, instead of three, sense cones on the fourth antennal segment, and the tube is
enlarged. The head and stylets are similar in the two genera, and both have well-developed
metathoracic sternopleural sutures.
30 L. A. MOUND AND J. M. PALMER
SPECIES INCLUDED
albicepsHood, 1938c: 401-2. Lectotype $, U.S.A.: Florida (USNM).
*callipygusHood, 1952c: 164-5. Holotype $, BRAZIL (USNM).
fortisHood, 1938c: 402. Holotype $, U.S.A.: Florida (USNM).
*longicepsHood, 1952c: 164. Holotype $, BRAZIL (USNM).
*/nag/iJcauc/aHood, 19540: 45. Holotype $, BRAZIL (USNM).
m/Jtrominatos(Moulton, 1944: 284-5) (Diplochelaeothrips). Holotype $, FIJI (BPBM). Comb. n.
"need/iaiiMHood, 1938c: 397-401. Holotype $, U.S.A.: Florida (USNM).
pygus (Mound, 1974a: 43) (Cryptothrips). Holotype $, AUSTRALIA (ANIC). Comb. n.
rug/caudaHood, 1915: 50-1. Holotype $, AUSTRALIA (USNM).
satanasDe Santis, 1957: 3-4. Holotype cf , ARGENTINA (MLPA).
sculpticauda (Hood & Williams, 1915: 135-6) (Barythrips). Holotype C?, U.S.A.: Florida (USNM).
comb. n.
sAaviani/s(Bagnall, 1918: 216-7) (Cryptothrips). Lectotype $, AUSTRALIA (BMNH). Comb. n.
*zeteJWHood, 1934: 420. Holotype cf , PANAMA (USNM).
Genera of Allothripina
This subtribe was erected by Priesner (1961) for six genera, each of which includes species with
reduced eyes, although Illinothrips is here transferred to the Compsothripina where it appears to
be closely related to Bolothrips. Sakimura & Bianchi (1977) also referred Diopsothrips to the
Allothripina, but this genus is here placed in the Diceratothripina as a synonym of Acalluro-
thrips. Moreover, both Allidothrips and Allopisothrips have subsequently been described as
allothripines.
Four genera recognised here in this subtribe share a unique apomorphy in the form of the
terminal sensorium on the maxillary palps. These genera are: Allopisothrips, Allothrips,
Priesneriella (= Pygidiothrips and Parallothrips) , and Pseudocryptothrips. All the species in
these genera have the terminal sensorium on the maxillary palps exceptionally large, looking
like an extra segment (Fig. 77), whereas in normal Idolothripinae the palps bear a terminal and
sub-terminal sensorium which do not differ greatly in size and arise almost at right angles.
Faureothrips is retained in the Allothripina although the only species does not have enlarged
maxillary palp sensoria, the metathoracic sternopleural sutures are not developed (they are
weak in Pseudocryptothrips), and the eighth antennal segment is pedicillate and distinct from
the seventh segment. Assuming that the Allothripina have developed from Pygothripina, with
which they share the presumably plesiomorphic characteristics involved in the tendency for
fusion of antennal segments VII- VIII and the close approximation of the maxillary stylets, then
Faureothrips must be interpreted as a reversion from the trend toward sclerite reduction.
Resemblance of F. reticulatus to some Bolothrips species is probably due to convergence
associated with adaptation to the leaf litter habitat. Allidothrips is also retained in this subtribe
and is discussed below.
ALLIDOTHRIPS Zur Strassen
(Fig. 64)
Allidothrips Zur Strassen, 1968: 86-7. Type-species: Allidothrips tricolor Zur Strassen, by monotypv
Mound (1972a) transferred Allothrips cinctus Faure to Allidothrips as a second species in il.e
genus. These species do not have the terminal sensorium on the maxillary palps exceptionally
large, although they share with the other Allothripina a series of characters involving reduction
of sclerites. Antennal segment III bears only one sense cone, although segments VII- VIII are
fused. The pelta is quite unique in that it appears to be a complete transverse tergite (Fig. 64).
This structure might be considered to be part of the same transformation series which includes
Priesneriella gnomus and P. seminole in which the pelta is reduced but transverse (Figs 62-63).
Alternatively, Allidothrips might be quite unrelated to this group, its larviform appearance
being interpreted as indicating development by neotony from some entirely different ancestry.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 31
SPECIES INCLUDED
ci/icfus(Faure, 1945: 150-2) (Allothrips) . Holotype $, SOUTH AFRICA (NCIP).
tricolor Zur Strassen, 1968: 87-90. Holotype $, MOROCCO (SMF).
ALLOPISOTHRIPSSakimura & Bianchi
Allopisothrips Sakimura & Bianchi, 1977: 498-9. Type-species: Allopisothrips alakaiensis Sakimura &
Bianchi, by monotypy.
This monobasic genus, based on a single male specimen, is intermediate in structure between
Allothrips and Priesneriella. The fused antennal segments VII-VIH are broadly based (unlike
Allothrips) but distinct from VI (unlike Priesneriella). The meso- and metanota are fused and
the pelta reduced, but the major setae are longer than in Priesneriella species.
SPECIES INCLUDED
alakaiensis Sakimura & Bianchi, 1977: 489. Holotype cf , KAUAI Is (BPBM).
ALLOTHRIPS Hood
(Figs 57, 58, 78)
Allothrips Hood, 1908a: 372-3. Type-species: Allothrips megacephalus Hood, by monotypy.
Bryothrips Priesner, 1925a: 6. Type-species: Bryothrips pillichelus Priesner, by monotypy. [Synonymised
by Stannard, 1957: 92.]
In this genus Mound (19720) recognised only four species, but with two of these divided into a
total of 16 subspecies. One reason for this interpretation was the recognition by Stannard (1955)
of a cline across North America. Mound (19720) described three forms from Australia as
subspecies of megacephalus, and suggested that these had been transported artificially by ships
trading across the Pacific from the American West Coast. This hypothesis requires testing by
collecting further populations of Allothrips in western America and Panama, but further
evidence for artificial transportation is provided by the record of brasilianus in large numbers on
the Hawaiian Islands (Sakimura & Bianchi, 1977). Allothrips species frequently produce quite
large populations in leaf litter but macropterae are rare. This probably leads to reduced gene
flow between natural populations resulting in increased structural diversity between popula-
tions.
SPECIES INCLUDED
fcras/J/anusHood, 1955: 101-3. Holotype $, BRAZIL (USNM).
megacephalus Hood
m. acutus Stannard, 1955: 154-5 (watsoniacutd). Holotype $, MEXICO (INHS).
m. greensladei Mound, 19720: 30. Holotype $, AUSTRALIA (ANIC).
m. megacephalus Hood, 19080: 373. Lectotype $, U.S.A.: Illinois (USNM).
m. mexicanus Stannard, 1955: 154 (watsoni mexicand). Holotype $, MEXICO (INHS).
m. prolixus Mound, 1972a: 30-1. Holotype $, AUSTRALIA (ANIC).
m. stannardi Mound, 19720: 31-2. Holotype $, AUSTRALIA (ANIC).
m. wateon/Hood, 1939ft: 600-2. Holotype $, U.S.A.: Florida (USNM).
nubillicauda Watson, 1935: 60-1. Syntypes cf $, U.S.A.: Florida, Alabama (FSAC).
piUichellus (Priesner)
p. acac/aeFaure, 1945: 152-4. Holotype $, SOUTH AFRICA (NCIP).
p. africanusFaure, 1933: 57-9. Holotype $, SOUTH AFRICA (NCIP).
p. aureus Stannard, 1955: 155. Holotype cf , U.S.A.: California (INHS).
p. Wco7or Ananthakrishnan, 19640: 83-4. Holotype $, INDIA (TNA).
p. biminianus Stannard, 1955: 155 (watsoni biminiana). Holotype $, BAHAMAS (INHS).
p. bournieri Mound, 1972a: 35-6. Holotype $, FRANCE (MNHN).
*p. indicus Ananthakrishnan, 1958: 277-8. Holotype $>, INDIA (TNA).
p. montanus Ananthakrishnan, 1968ft: 53. Holotype $, INDIA (TNA).
p. piUichellus (Priesner, 19250: 6-7) (Bryothrips). Holotype $, HUNGARY (SMF).
32 L. A. MOUND AND J. M. PALMER
FA UREOTHRIPS Priesner
(Figs 66, 67)
Faureothrips Priesner, 1949: 116-7. Type-species: Cryptothrips reticulatus Trybom, by monotypy.
This monobasic genus from southern Africa is difficult to place phylogenetically. The antennae
are similar to Pseudocryptothrips , with two sense cones on segment III and four on IV; however,
segment VIII is slender and pedicillate. Moreover, as in Pseudocryptothrips, the stylets are fairly
wide apart, there is a pair of stout interocellar setae with expanded apices, and the eyes are
somewhat reduced (Fig. 67). However, unlike the other Allothripina, Faureothrips does not
have the terminal sensorium on the maxillary palps exceptionally large, and the metathoracic
sternopleural sutures are not developed (N.B. they are only weakly developed in Pseudocryp-
tothrips). The pronotal epimeral sutures are incomplete, the praepectus is present, the pelta
broad (Fig. 66), but the metanotum bears more than 10 minor setae in addition to a pair of
widely spaced major setae.
SPECIES INCLUDED
reticulatus (Trybom, 1912: 9-13) (Cryptothrips). Syntypes d" $, SOUTH AFRICA (NMG).
PRIESNERIELLA Hood
(Figs 59-63, 68-72, 74-77, 79-83)
Priesneriella Hood, I921b: 198-9. Type-species: Priesneriella citricauda Hood, by monotypy.
Pygidiothrips Hood, 1938c: 389-90. Type-species: Pygidiothrips seminole Hood, by monotypy. Syn. n.
Parallothrips Hood, 1939&: 602. Type-species: Parallothrips thomasi Hood, by monotypy. Syn. n.
Embothrips Dyadechko, 1961: 688-9. Type-species: Embothrips tubversicolor Dyadechko, by monotypy.
[Synonymised with Parallothrips by Dyadechko, 1964: 307.]
Seven species are listed under the above four generic names, and a further new species is
described below from New Zealand. Several characters in these eight species exhibit trans-
formation series involving reduction or fusion. Traditional interpretations of this variation
would require five genera although the present authors regard these species as constituting a
single holophyletic group. That is, the group was derived once from Allothrips-\ike ancestors.
Allopisothrips appears to be the sister-group. The apomorphy on which this conclusion is based
is the broad and close union of antennal segment VI with segments VII & VIII (Figs 79-83). The
seven species examined have the following characteristics.
P. thomasi. Maxillary stylets close together; ocellar setae stout; antennal III with 2 sense cones, VI
subequal in length to VII + VIII; meso- and metanota fused; pelta with median lobe, and wide but slender
base; sternite IV with 4 discal setae; tube width 0-9 times length (Figs 60, 68, 82).
P. mavromoustakisi. As thomasi but pelta larger; tube width 0-8 times length.
P. luctator. As thomasi; tube width 0-76 times length.
P. clavicornis. Maxillary stylets wide apart; ocellar setae stout; antennal III with 0(?1) sense cones, VI
slightly larger than VII + VIII but these have a partial suture; meso- and metanota not quite fused; pelta
with median lobe and wide base; sternite IV with 2 discal setae; tube width 0-7 times length (Figs 59, 70,
81).
P. gnomus. Maxillary stylets wide apart; ocellar setae small; antennal III with 0(or 1) sense cones, VI
shorter than VII + VIII; meso- and metanota separate; pelta slender and transverse; sternite IV with 0
discal setae; tube width 0-68 times length (Figs 62, 71, 74, 79).
P. citricauda. Stylets wide apart; ocellar setae small; antennal III with 0 sense cones, VI fused to
VII -I- VIII; meso- and metanota separate; pelta as in thomasi; sternite IV with 0 discal setae; tube width 0-8
times length (Figs 61, 80).
P. seminole. Stylets wide apart; ocellar setae small; antennal III with 1 sense cone, VI fused to
VII -I- VIII; meso- and metanota separate; pelta slender and transverse; sternite IV with 0 discal setae; tube
width 1-1 times length (Figs 63, 69, 72, 75, 76, 83).
The following characters are plesiomorphic in this group, in that they also occur in Allothrips
and Pseudocryptothrips: stylets close together; ocellar setae long and stout; antennal III with 2
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 33
sense cones, terminal segments separate; meso and metanota separate; pelta large and rounded;
sternites with numerous discal setae; tube width 0-5 times length. Most of these characters (apart
from the fusion of the meso- and metanota) are in their most derived state in the smallest species.
SPECIES INCLUDED
c/fricaudaHood, 1927ft: 199. Lectotype $, U.S.A.: California (USNM).
c7avic0rn/s(Knetchtel, 1936: 159-60) (Hoplothrips). Syntypes cf , RUMANIA (unknown). Comb. n.
tuzetae Bournier, 1956: 160-3 (Parallothrips). Holotype $, FRANCE (BCM).
gnomussp. n. Holotype $, NEW ZEALAND (NZAC).
/ucfafor (zur Strassen, 1966: 3-6) (Parallothrips). Holotype $, TENERIFFE (SMF). Comb. n.
ma vromoustakisi (Crawford, 1948: 213-5) (Parallothrips). Holotype $, CYPRUS (USNM). Comb. n.
i.flaviceps Bournier, 1962: 43. Types not specified, FRANCE (BCM).
seminole(Hood, 1938c: 390-2) (Pygidiothrips). Holotype $, U.S.A.: Florida (USNM). Comb. n.
thomasi (Hood, 1939ft: 603-5) (Parallothrips). Holotype $, U.S.A.: Texas (USNM). Comb. n.
*tubversicolor (Dyadechko, 1961: 688-9) (Embothrips). Syntypes cf $, U.S.S.R.: Kiev (Acad. Sci.,
U.S. S.R.). Comb. n.
Priesneriellagnomussp. n.
(Figs 62, 71, 74, 79)
Apterous $. Colour brown, inner margin of forefemora and apex of antennal II paler; basal two-thirds of
tube yellow. Head scarcely longer than wide; eyes small with only about 5 ommatidia ventrally;
ocellar-setae small, postocular setae long and acute (Fig. 71); stylets wide apart, maxillary palps with large
terminal sensorium (Fig. 77). Antennae with 7 segments, VI separate from VII + VIII; IV with 2 sense
cones, III without sense cones or with one small one (6 /x,m) (Fig. 79). Pronotum transverse, epimeral
sutures incomplete, anterior setae reduced. Praepectus and probasisternum absent; metathoracic sterno-
pleural sutures broad. Meso- and metanota separate. Pelta very short but wide and close to anterior margin
of tergite II (Fig. 62); tergites without sculpture, major setae long and slender (Fig. 74); tube with sides
almost straight, scarcely constricted at apex; sternites II-IV without discal setae.
Measurements (holotype 9 in Atni). Body length 1500 (extended). Head, length 150; median width 150;
postocular setae 80. Pronotum, length 110; width 220; major setae -am 25, aa 20, ml 25, epim 120, pa 120.
Tergite IX median dorsal setae 150. Tube, length 110; basal width 75; terminal setae 140. Antennal
segments II-VII length 45, 36, 40, 42, 45, 50.
SPECIMEN STUDIED
Holotype $, New Zealand: South Island, 25 ml west of Christchurch, Kowai Bush, on dead branch of
Griselinia littoralis, 13.x. 1972 (V. F. Eastop) (NZAC).
COMMENTS. This new species has the distal antennal segments similar to thomasi (type-species of
Parallothrips), but the third segment similar to citricauda and seminole. The left antenna of the
holotype has a small (6 /u-m) sense cone externally, but this is not visible on the right antenna
(similar variation occurs in clavicornis from southern France). The meso- and metanota are
separate as in citricauda, and the pterothoracic endofurca is stout as in that species and
mavromoustakisi. P. gnomus is probably not native to New Zealand.
PSEUDOCRYPTOTHRIPS Priesner
(Figs 56, 65, 73)
Pseudocryptothrips Priesner, 1919: 105. Type-species: Pseudocryptothrips meridionalis Priesner, by
monotypy.
The type-species of this genus is very similar toAllothrips species in the structure and chaetotaxy
of the head, and in the form of the pelta and maxillary palp sensoria. However, antennal
segments VII- VIII are separated by a complete suture, and IV bears four sense cones (Fig. 73).
Moreover, the meso- and metanota are rather more fused than in Allothrips species, although
the meso- and metasterna are less eroded, and the mesonotum bears a short wing lobe (60 /zm)
laterally with one or two setae. Specimens of this genus have been studied from the following
countries: Mexico (3 $, 2 cf), Barbados (l£), Trinidad (2 $), Transvaal (2 cf), Kenya (1 $),
34 L. A. MOUND AND J. M. PALMER
France (1 $). However, the present authors are not convinced that this material represents
more than one species, although three species have been described in the genus. P. remotus
Bianchi (1947) from Hawaii was transferred to Apterygothrips in the Phlaeothripinae by
Sakimura & Bianchi (1977).
SPECIES INCLUDED
fuscicauda (Trybom, 1912: 13-5) (Cryptothrips). Holotype $, SOUTH AFRICA (NMG).
proximus Faure, 1933: 55-7. Holotype (JT, SOUTH AFRICA (NCIP).
*gradatus(Hood, 19256: 64) (Cryptothrips). Holotype £, TOBAGO (USNM).
merid/ona/JsPriesner, 1919: 105-^6. Syntypes $ cf , ALBANIA (SMF).
Genera of Compsothripina
This group was erected by Karny (19210) as a subfamily to include four generic names. Two of
these are now placed in Elaphrothripina as synonyms of Anactinothrips , the other two are here
treated as a single genus. The subtribe is here reinterpreted to include an ill-defined series of
Pygothripini mainly found at soil level in association with grasses and litter. The species share a
combination of the following characters: usually apterous; antennal segment VIII distinct from
VII; antennal segment IV with 3 (or 2) sense cones, III with 2 or 1 sense cones; eyes frequently
reduced laterally, but often extended ventrally; praepectus present; mesopraesternum entire;
metathoracic sternopleural sutures well developed or absent; tube short with sides straight.
Bolothrips species, together with the related monobasic genera Illinothrips, Loyolaia and
Anaglyptothrips , are here brought into the same subtribe as the ant-mimicking species of
Compsothrips (together with Leptogastrothrips and Oedaleothrips) . This decision is based on a
comparison of Bolothrips cingulatus, which has long metathoracic sternopleural sutures (Fig.
98), with species of Compsothrips from the Mediterranean region. In this region several species
are known which are intermediate in structure between Bolothrips and Compsothrips (B.
cingulatus, B. insularis, C. albosignatus, C. maroccanus and C. uzeli). These two genera
probably constitute sister-groups, of which one has radiated mainly in the Holarctic Region, and
the other mainly in the tropics where it has produced remarkable ant-mimics.
In Compsothripina the plesiomorphic condition of the antennal sensoria is regarded as: III
with 2 sense cones, IV with 3 sense cones. However, there are four species of Bolothrips with the
ventral sense cone missing on segment IV (two species Mediterranean, two species South
African) (Fig. 118), and this derived condition is maintained in Compsothrips (Figs 113, 114).
The ventral prolongation of the eyes may be plesiomorphic in this group, but is possibly
functionally correlated with the habit of living at the base of grasses. The eyes are not prolonged
ventrally in B. pratensis (Fig. 96) and /. rossi (N. America) (Fig. 104), L. indica (India) (Fig.
105), A. dugdalei (New Zealand) (Fig. 103), and only weakly so in C. albosignatus (Fig. 84), C.
maroccanus and B. insularis. As a result the subtribe is not easy to define, but on the basis of the
antennal sense cone formula it may be the sister-group of the Gastrothripina.
In contrast to Stannard (1976) Hartwigia is here transferred to the Elaphrothripina in the
Idolothripini because of the presence of two pairs of wing-retaining setae on each tergite, and
also the absence of metathoracic sternopleural sutures.
ANAGLYPTOTHRIPS gen. n.
(Figs 97, 103, 106, 115)
Type-species: Anaglyptothrips dugdalei sp. n.
Medium sized, apterous, yellowish Pygothripini with body surface, including legs and antennae, reticulate.
Antennae 8-segmented, VIII not constricted at base, 2 sense cones on III, 3 on IV (Fig. 115). Head longer
than wide, protruding in front of small rounded eyes (Fig. 103); postocular setae of 9 scarcely longer than
minor setae , but half as long as eye in cf ; maxillary stylets V-shaped and low in head ; mouth cone short and
rounded. Pronotum with no long setae, epimeral sutures complete (Fig. 103). Foretarsal tooth absent in $ ,
present in d". Mesonotum transversely rectangular; metanotum transverse with explanate lateral margins.
Praepectus small (Fig. 103); probasisternum large; mesopraesternal posterior margin short; metathoracic
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 35
sternopleural sutures long and curved (Fig. 97). Pelta broadly oval (Fig. 106); tergal discal setae numerous,
posteromarginal setae short and blunt on anterior segments but longer on posterior segments; tube
moderately long, margins straight; sternal discal setae in one transverse row.
This new genus is considered to be related to Bolothrips because of the presence of three sense
cones on antennal segment IV, and because of the long curved metathoracic sternopleural
sutures as in B. varius and B. cingulatus (Fig. 98). It is distinguished from all other Idolothripi-
nae by the sculpture, pale body colour, rounded eyes and short postocular setae. Faureothrips
reticulatus is the only species of similar appearance, but that has four sense cones on antennal
segment IV and lacks the sternopleural sutures. The new species for which this new genus is
erected has been collected only in New Zealand, but as discussed below it is probably introduced
to that country.
Anaglyptothrips dugdaleisp. n.
Apterous $. Colour brownish yellow, posterior segments darkest; eyes and antennal segments VI- VIII
dark brown, V light brown; major setae on posterior abdominal segments pale. Sculpture of head evenly
reticulate in posterior third but irregular medially (Fig. 103); compound eyes with about 10 ommatidia.
Pronotal sculpture weaker, major setae not distinguishable from minor setae (Fig. 103). Metanotum with
one pair of major setae medially and 10 pairs of minor setae. Pelta with strong sub-basal line of sculpture
(Fig. 106). Tergites strongly reticulate.
Measurements (holotype $ in /xm). Body length (extended) 2450. Head, length 300; width across cheeks
200; postocular setae 15. Pronotum, length 195; width 270; epimeral setae 10. Metanotal median setae 20.
Tergite IX setaeZ?!-/^ 60, 70, 100. Tube, length 195; maximum width 95; terminal setae 100. Antennal
segments III- VIII length 70, 60, 55, 55, 45, 30.
Apterous d". Similar to $ except postocular setae longer (45 fim), foretarsus with stout curved tooth,
metanotal median setae short (15 /mi).
SPECIMENS STUDIED
Holotype $, New Zealand: North Island, Waiwera, at base of grasses, 21.viii.1968 (L. A. Mound 802)
(NZAC).
Paratypes. New Zealand. North Island: 5 $ collected with holotype; Huia, near Auckland, 1 C? at base of
grass tussock, 24.L1979 (L. A. Mound 1353); Auckland, Lynfield, 3 9 under dead sheep in field, 6.iii.l977
(G. Kuschel) (NZAC; BMNH).
COMMENTS. The sites at which this species has been collected all had a ground cover of European
grasses. In New Zealand it is relatively unusual for native species to be found in association with
non-native habitats (Mound & Walker, 1982). Moreover, despite extensive sampling of leaf
litter in New Zealand, A. dugdalei has not been found in areas of native flora. Therefore it seems
likely that Anaglyptothrips is itself introduced to New Zealand from some other continent.
BOLOTHRIPS Priesner
(Figs 94-96, 98, 99, 101, 102, 118, 119)
Bolothrips Priesner, 1926a: 90. Type-species: Phloeothrips bicolor Heeger, by original designation.
Bolothrips (Botanothrips) Hood, 1939ft: 605-6. Type-species: Bolothrips pratensis Hood, by original
designation.
Boloadelothrips Moulton, 1949: 489. Type-species: Boloadelothrips africanus Moulton, by monotypy.
Syn. n.
A revision of this genus, including a key to 14 species, was given by Mound (1974&). Although
widespread in the Holarctic, it is also represented in Africa by several species, lllinothrips from
North America, Loyolaia from India, and Anaglyptothrips are here regarded as derivatives
from Bolothrips because of the presence of three sense cones on the fourth antennal segment
(Fig. 119).
Botanothrips was proposed for species with the eyes not prolonged ventrally (Fig. 96), but this
varies within species (e.g. varius) and does not define a real phylogenetic group. Boloadelothrips
is also placed in synonymy here, the only species having antennae similar to Bolothrips dentis
36 L. A. MOUND AND J. M. PALMER
(1 sense cone on III, 2 on IV; VII-VIII broadly joined). Both dentis and africanus have
metathoracic sternopleural sutures, but the former has long interocellar setae and the female
bears a foretarsal tooth. Most Bolothrips species have lost the metathoracic sternopleural
sutures, but these are present in insularis and the closely related varius (N.B. not icarus, Fig. 99),
and are exceptionally long in cingulatus (Fig. 98). Bolothrips is here interpreted as the
sister-group of Compsothrips which appears to replace it ecologically in much of the tropics.
SPECIES INCLUDED
africanus (Moulton, 1949: 489-92) (Boloadelothrips) . Holotype 9, SOUTH AFRICA
(BMNH). Comb. n.
bicolor (Heeger, 1852ft: 477-8) (Phlaeothrips). Syntypes ?sex, AUSTRIA (?lost).
f. brevicornis Priesner, 1928a: 687. Holotype $, HUNGARY (SMF).
andrei Watson, 1933: 49-50 (Oedaleothrips). Syntypes $ cf , U.S.A.: Iowa (FSAC).
c/ncfusFaure, 1943: 86-7. Syntypes $, SOUTH AFRICA (NCIP).
cingulatus (Karny, 1916: 92) (Cryptothrips). Syntypes? sex, AUSTRIA (SMF).
dentipes (Reuter, 1880: 12-4) (Phloeothrips) . Syntypes ?sex, FINLAND (?lost).
bagnalli Karny, 1916: 94 (Cryptothrips). Syntypes ?sex, SARDINIA (?lost).
dentis Faure, 1954ft: 155-9. Holotype cf , SOUTH AFRICA (NCIP).
*emfeofy/Faure, 1943: 87-9. Syntypes $ cf , SOUTH AFRICA (NCIP).
guV/pes(Hood, 1914: 169-70) (Cryptothrips}. Holotype $, U.S.A.: Maryland (USNM).
litoreus Hood, 1939ft: 609-12. Holotype $, U.S.A.: Texas (USNM).
icarus (Uzel, 1895: 323-3) (Cryptothrips}. Syntypes cf $, CZECHOSLOVAKIA (?lost).
var. pallipes Uzel, 1895: 233. Syntypes $cf , CZECHOSLOVAKIA (?lost).
uisu/aris (Bagnall, 1914ft: 295) (Cryptothrips). Holotype $, CANARY ISLAND (BMNH).
icarus tuberculatus Priesner, 1922: 105 (Cryptothrips}. Holotype $,
YUGOSLAVIA (SMF).
brachyurus Bagnall, 1927: 573-4 (Cryptothrips). Lectotype cf , FRANCE (BMNH).
arenarius Priesner, 1950: 36-7. Syntypes $ cf , EGYPT & SYRIA (SMF).
italicus Mound, 1974ft: 122. Holotype $, ITALY (USNM).
prafens/sHood, 1939ft: 606-9. Holotype 9, U.S.A.: Texas (USNM).
*rac/i/p/H7usCott, 1956: 181-2. Holotype $, U.S.A.: California (?lost).
scnaferi(Thomasson & Post, 1966: 31-2) (Nesothrips). Holotype $, U.S.A.: North
Dakota (INHS).
varius Hartwig, 1948: 110-2. Holotype $, SOUTH AFRICA (NCIP).
COMPSOTHRIPS Reuter
(Figs 84-86, 100, 107, 108, 111-114)
Compsothrips Reuter, 1901: 214. Type-species: Phloeothrips albosignata Reuter, by monotypy.
Macrothrips Buffa, 1908: 4. Type-species: Phloeothrips albosignatus Reuter, by monotypy.
Leurothrips Bagnall, 1908: 196. Type-species: Leurothrips albomaculata Bagnall, by original designa-
tion. [Synonymised by Priesner, 1928. J
Leptogastrothrips Trybom, 1912: 28. Type-species: Leptogastrothrips reuteri Trybom, by monotypy.
[Synonymised by Priesner, 1964.]
Oedaleothrips Hood, 1916: 64. Type-species: Oedaleothrips hookeri Hood, by original designation.
[Synonymised by Priesner, 1964.]
Myrmecothrips Watson, 1920: 20. Type-species: Myrmecothrips querci Watson, by original designation.
[Synonymised with Oedaleothrips by Watson, 1924.]
Myrmecothrips Priesner, 1926ft: 485-8. Type-species: Myrmecothrips dampfi Priesner, by original designa-
tion. [Junior homonym of Myrmecothrips Watson, 1920] [Synonymised with Oedaleothrips by Hood,
1936.]
Formicothrips Priesner, 1927: 479. [Replacement name for Myrmecothrips Priesner.] [Synonymised with
Oedaleothrips by Hood, 1936.]
Stannard (1976) recognised three genera in this group, although apparently accepting that they
constituted together a holophyletic assemblage. Compsothrips was used for one species,
albosignatus, with the head produced into a cone over the antennal bases (Fig. 84), the
mesonotum short and transversely rectangular, and the metanotum rectangular without a
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 37
median raised area. Oedaleothrips was reserved by Stannard for the North American species of
the group on the argument that in these the head is greatly swollen behind the eyes (Fig. 86) and
the metanotum produced medially into a cone or node. Leptogastrothrips was used for the rest of
the species in the group from the Old World as well as the Neotropics. This division into three
genera is not accepted here for the following reasons. One particularly small male of C.
albosignatus from Greece has been studied which lacks a head cone; this structure probably
varies in size allometrically. Moreover, although the mesonotum of most species placed in
Leptogastrothrips and Oedaleothrips is relatively long (Fig. Ill) and quite unlike the transverse
mesonotum of albosignatus (Fig. 112), this is not true of uzeli which has a typical ' Leptogas-
trothrips' head but a mesonotum and metanotum similar to albosignatus. The species marocca-
nus is also intermediate between the two groups. Similarly, although the head and metanotum of
certain North American species are remarkable in structure, the head ofyosemitae is intermedi-
ate between Leptogastrothrips and Oedalothrips, and the metanota of reuteri and hookeri (the
two type-species) are essentially similar. Hartwigia, another ant-mimic which is superfically
similar, is here transferred to the Elaphrothripina (p. 68).
Compsothrips is here interpreted as the sister-group of Bolothrips which it largely replaces in
the tropics. The genus is circumtropical, but with most species in Africa and South America, one
species-group in North America, and a few species in the North African/Mediterranean region.
The species of continental areas appear to exist as a series of intergrading populations which are
difficult to classify, as has been reported for other groups of apterous thrips living at soil level
(Mound, 1972b). The metathoracic sternopleural sutures are more strongly developed in this
genus than in any other idolothripines (Fig. 100), possibly correlating with the narrowed ant-like
body form.
SPECIES INCLUDED
*aeneus(Hood, 1937a: 280-5) (Oedaleothrips}. Holotype $, PERU (USNM). Comb. n.
albosignatus (Reuter, 1884: 290-1) (Phloeothrips) . Syntypes $ cf , ALGERIA (?lost).
dbomaculata Bagnall, 1908: 196-8 (Leurothrips) . Holotype $, no data (BMNH).
baileyi (Hood, 1941: 193-5) (Oedaleothrips). Holotype $, U.S.A.: Kansas (USNM). Comb. n.
*Wco/orPriesner, 1921: 213-5. Syntypes $ cf , PARAGUAY (SMF).
brasiIiensis(Hood, 1952c: 166-7) (Oedaleothrips). Lectotype $, BRAZIL (USNM). Comb. n.
brunneus(Hood, 1941: 187-90) (Oedaleothrips). Holotype $, U.S.A.: Florida (USNM). Comb. n.
congoensis(Hood, 1952ft: 204-9) (Oedaleothrips). Lectotype $, CONGO (USNM). Comb. n.
* damp/i (Priesner, 1926ft: 488-9) (Myrmecothrips). Holotype cf , MEXICO (?lost). Comb. n.
graminis (Hood, 1936c: 265-9) (Oedaleothrips). Holotype $, TRINIDAD (USNM). Comb. n.
*hoodi(De Santis, 1958: 98-9) (Leptogastrothrips). Holotype $, ARGENTINA (MLPA).Comb. n.
hookeri (Hood, 1916: 64-5) (Oedaleothrips). Holotype $, U.S.A.: Texas (USNM). Comb. n.
bradleyiHood, 1937ft: 111-3 (Oedaleothrips). Holotype $, U.S.A.: Florida (USNM).
campestris Hood, 1941: 190-3 (Oedaleothrips). Holotype $, U.S.A.: Florida (USNM).
jacksoni(Hood, 1925c: 137-8) (Oedaleothrips). Holotype $, U.S.A.: Colorado (USNM). Comb. n.
hubbeli Watson, 1931: 341-2. Holotype $, U.S.A.: Oklahoma (FSAC).
maroccanus Priesner, 1964: 146. Holotype $, MOROCCO (?SMF).
*one#7aeBournier, 1974: 153-6. Holotype $, ANGOLA (MDA).
*pampicolla (De Santis, 1958: 100-2). (Leptogastrothrips). Holotype $, ARGENTINA (MLPA). Comb. n.
*querci( Watson, 1920: 20-1) (Myrmecothrips). Syntypes $ cf, U.S.A.: Florida (FSAC). Comb. n.
ramamurfM/( Ananthakrishnan, 1964ft: 111-3) (Oedaleothrips). Syntypes $ cf , INDIA (TNA). Comb. n.
ramamurthii indicus Ananthakrishnan, 1973ft: 120. [Replacement name for ramamurthii
bicolor Ananthakrishnan, 1966: 11, nee ft/co/or Priesner.]
reuteri (Try bom, 1912: 29-31) (Leptogastrothrips). Holotype $, SOUTH AFRICA (NMG).
recticeps Hood, 1925o: 293-5. Holotype $, SOUTH AFRICA (USNM).
amabilis Jacot-Guillarmod, 1942: 67-71 (Oedaleothrips). Holotype Qj, SOUTH AFRICA (AMG).
aemulus Jacot-Guillarmod, 1942: 71-4 (Oedaleothrips}. Holotype $, SOUTH AFRICA (AMG).
*sinensis(Pelikan, 1961: 306-8) (Oedaleothrips}. Holotype $, CHINA (PelikanColl.).
*sumafranus Priesner, 1928c: 54-5. Holotype $, SUMATRA (?lost).
*tfnnir(Pelikan, 1961: 302-6) (Oedaleothrips}. Holotype $, U.S.S.R. (Pelikan Coll.).
*frisfis(Cott, 1956: 186-8) (Oedaleothrips}. Holotype $, U.S.A.: California (?lost).Comb. n.
uze//(Hood, 1952a: 134-40) (Oedaleothrips). Holotype $, ITALY (USNM).
38 L. A. MOUND AND J. M. PALMER
*walteri( Watson, 1933: 48-9) (Oeddeothrips). Holotype $, ARGENTINA (FSAC). Comb. n.
yosemitae(Mou\tor\, I929a: 135-6) (Formicothrips) . Holotype $, U.S.A.: California (CAS). Comb. n.
ILLINOTHRIPS Stannard
(Figs 104, 110, 116)
Illinothrips Stannard, 1954: 193-5. Type-species: Illinothrips rossi Stannard, by monotypy.
This monobasic genus was compared originally to Pseudocryptothrips and to Gastrothrips
acuticornis. However, the pelta (Fig. 110) and abdomen are typical of Bolothrips, and the
antennae essentially similar to members of that genus (3 sense cones on IV, but only 1 on III)
(Fig. 116). The head, with the eyes small and cheeks incut behind the eyes (Fig. 104), is unlike
most species of Bolothrips, although B. pratensis from North America is intermediate. Only one
female of rossi has been examined; according to Stannard the male bears a tooth-like projection
in front of the mesothoracic spiracle. The metathoracic sternopleural sutures are retained in
rossi, which implies that the species is not derived from the present North American Bolothrips
fauna.
SPECIES INCLUDED
rossi Stannard, 1954: 195-6. Holotype $, U.S.A.: Illinois (INHS).
LOYOLAIA Ananthakrishnan
(Figs 105, 109, 117)
Loyolaia Ananthakrishnan, 1964ft: 106-7. Type-species: Loyolaia indica Ananthakrishnan, by monotypy.
As indicated in the original description, this monobasic genus from India is similar in appearance
to Illinothrips, and the antennae have a similar sense cone arrangement (1 on III, 3 on IV)
(Fig. 117). The metathoracic sternopleural sutures are present, and the prothoracic epimeral
sutures complete, but unlike Illinothrips the pelta of indica is trilobed (Fig. 109). This species is
probably derived from the holarctic genus Bolothrips.
SPECIES INCLUDED
indica Ananthakrishnan, 1964ft: 107-8. Syntypes $ cT, INDIA (TNA).
Genera of Gastrothripina
This subtribe was erected by Priesner (1961) for a series of 13 generic names, most of which
through subsequent reinterpretation are now placed elsewhere. As a result only Gastrothrips,
with seven generic synonyms, remains in the subtribe, the species of which (in common with
most Bolothrips species) are unusual amongst Idolothripinae in having three sense cones on the
fourth antennal segment (Fig. 93). In Gastrothrips species, however, these sense cones are
relatively short and stout. The metathoracic sternopleural sutures are present (Fig. 128), but
unlike Bolothrips in the Compsothripina, the eyes are usually rounded and not prolonged
ventrally (Fig. 120), and the pelta, although variable, is basically triangular with lateral wings
rather than rounded (Figs 121-124). Gastrothripina is possibly the sister-group of Compso-
thripina, the two groups having adopted different habitats; the former is common on dead twigs
and branches in the tropics, the latter is found in grass tussocks and leaf litter in both tropical and
temperate regions.
GASTROTHRIPS Hood
(Figs 87-93, 120-129)
Gastrothrips Hood, 1912c: 156. Type-species: Gastrothrips ruficauda Hood, by original designation.
Goetothrips Priesner, 1925c: 316. Type-species: Goetothrips terrestris Priesner, by monotypy. [Synony-
mised by Johansen, 1978c: 277.]
Isopterothrips Bagnall, 1926: 553. Type-species: Isopterothrips tenuipennis Bagnall, by monotypy. Syn. n.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 39
Syncerothrips Hood, 1935ft: 191-2. Type-species: Syncerothrips hard Hood, by monotypy. Syn. n.
Probolothrips Moulton, 1941: 319. Type-species: Probolothrips hambletoni Moulton (now regarded as a
synonym ofabditus), by monotypy. [Synonymised by Hood, 1952: 163.]
Pharetrothrips Priesner, 19520: 195. Type-species: Agnostochthona curvidens Karny, by monotypy.
Syn. n.
Percnothrips Ananthakrishnan, 1967: 233. Type-species: Percnothrips turbinatus Ananthakrishnan, by
monotypy. Syn. n.
Paragastrothrips Zur Strassen, 1977: 59-60. Type-species: Paragastrothrips mauli Zur Strassen, by
monotypy. Syn. n.
A definition of this genus, with a key to 16 species, was given by Mound (1974ft). The
Neotropical species referred to in couplets 5 to 15 of that key, together with gurdus, harti and
terrestris, form a closely related and presumably holophyletic group. One species, terrestris, was
placed in a separate genus Goetothrips because the interantennal projection is relatively long.
However, Gastrothrips fulviceps has a similar but shorter projection. Similarly, harti was placed
separately in Syncerothrips because of the partial fusion of antennal segments VII-VIII
(cf. Fig. 93), whereas most of the South American species have the eighth segment long and
slender (Fig. 91).
A smaller group of Old World species, in which the tube is not constricted apically (Mound,
1974ft: 136 - key couplet 2 plus acutulus} (Fig. 88), may also constitute a holophyletic group,
although no generic name has ever been proposed for it. However, the six remaining species
(curvidens, fuscatus, mauli, tenuipennis, turbinatus and xosa) seem to be less closely related, but
although four generic names are available they are here placed in synonymy until such time as
the Old World fauna is more fully investigated. Pharetrothrips was erected for a species with
long preocellar setae, and a long curved apical tubercle on the foretibiae; antennal segment VIII
of this species is narrowed basally, whereas in fuscatus (which also has a foretibial tubercle)
segments VII-VIII are broadly joined. Percnothrips, like Syncerothrips, was erected for a
species with antennal segments VII-VIII almost completely fused (Fig. 93) but with a pair of
long preocellar setae (Fig. 126). Paragastrothrips includes a single species which is similar to
several Gastrothrips species but with a long head, broad pelta and short antennal segment VIII.
Finally, Isopterothrips includes a single large species with long postocellar setae, a fan of stout
spines on the forecoxae in males, and a slender eighth antennal segment. The species xosa,
known from a single female, is very similar to tenuipennis but smaller.
All of the species listed below as examined have well-developed metathoracic sternopleural
sutures (Fig. 128), three sense cones on antennal segment IV and two (or one) on segment III
(Figs 92 , 93) , and only the three species noted above have long ocellar setae . In most New World
species antennal segment VIII is long and slender, but it is broadly joined to VII in most Old
World species.
SPECIES INCLUDED
abdrtusHood, 1935ft: 177-82. Holotype $, PANAMA (USNM).
brasiliensis Moulton, 1938: 378-9 (Hoplothrips}. Holotype $, BRAZIL (CAS).
hambletoni Moulton, 1941: 320-1 (Probolothrips). Holotype $, BRAZIL (CAS).
acuficomis(Hood, 1925ft: 65) (Cryptothrips). Holotype $, WEST INDIES: St. Croix (USNM).
cybele Girault, 1927 d: 1 (Cryptothrips}. Holotype cf , AUSTRALIA: Queensland (QMB).
noumeae Bianchi, 1945: 251-4. Holotype £, NEW CALEDONIA (BPBM).
acufu7usOkajima, 1979c: 511-3. Holotype $, JAPAN (OCT).
a7fico/aHood, 1942: 570-3. Holotype $, PERU (USNM).
anolis Morgan, 1925: 7-8. Holotype $, PUERTO Rico (USNM).
proteus Hood, 1933: 417-9. Holotype $, PANAMA (USNM).
eatfipusHood, 1935ft: 182^6. Holotype $, U.S.A.: Texas (USNM).
*citriceps (Priesner, 1921: 208-9) (Cryptothrips}. Holotype $, PARAGUAY (ZMB).
corvus Priesner, 1933: 55-7. Holotype $, MEXICO (SMF).
capitalis Hood, 1935ft: 174-7. Holotype $, U.S.A.: Texas (USNM).
curvidens (Karny, 1921c: 38-41) (Agnostochthona). Holotype $, JAVA (SMF). Comb. n.
falcatus( Ananthakrishnan, 1968c: 969-71) (Nesothrips}. Syntypes $ C?, INDIA (TNA).
/Wv/caudaHood, 1937a: 277-80. Holotype $, PERU (USNM).
40 L. A. MOUND AND J. M. PALMER
fu/vicepsHood, 1937a: 274-7. Holotype $, PERU (USNM).
fumipennisHood, 1952c: 163. Holotype $, BRAZIL (USNM).
fuscafusOkajima, 1979c: 513-5. Holotype $, TAIWAN (OCT).
harti(Hood, 1935ft: 192-4) (Syncerothrips). Holotype $, U.S.A.: Texas (USNM). Comb. n.
*heterocerus (Hood, 1925ft: 66) (Barythrips). Syntypes $ d", WEST INDIES: St. Thomas (USNM).
[Jacot-Guillarmod, 1978: 1427 states 'Should be placed in Neosmerinthothrips'.]
intonsusHood, 1941: 180-3. Holotype $, PERU (USNM).
mand/ocae(Moulton, 1941: 321-2) (Dichaetothrips). Holotype $, BRAZIL (CAS).
*oeceticola De Santis, 1943: 92-6. Holotype $, ARGENTINA (MLPA).
mau/i(Zur Strassen, 1977: 60-63) (Paragastrothrips) . Holotype $, MADEIRA (SMF). Comb. n.
m0ngo7/cus(Pelikan, 1965: 231-3) (Nesothrips). Holotype $, MONGOLIA (TM).
monf/co/aHood, 1942: 573-6. Holotype $, PERU (USNM).
procerusHood, 1956: 99-100. Holotype $, BRAZIL (USNM).
prof urus (Bagnall, 1921«: 269-70) (Acallurothrips} . Holotype $, SEYCHELLES (BMNH). Comb. n.
*pueWaeJohansen, 1979: 179-80. Holotype $, MEXICO (UNAM).
ruficaudaHood, 1912c: 156-7. Syntypes $, U.S.A.: Illinois (USNM).
stygkusHood, 1935ft: 186-91. Holotype 5 , PANAMA (USNM).
su5u/afus(Hartwig, 1948: 113-5) (Bolothrips). Holotype $, SOUTH AFRICA (NCIP).
fenuipen/i/s (Bagnall, 1926: 554) (Isopterothrips) . Holotype $, GHANA (BMNH). Comb. n.
penicillatus Priesner, 1937ft: 626-9 (Dichaetothrips). Holotype $, SIERRA LEONE (BMNH).
terrestris (Priesner, 1925c: 316-7) (Goetothrips). Syntypes $ cf, MEXICO (SMF). Comb. n.
*gurdus Johansen, 1974: 266 (Nesothrips). Holotype $, MEXICO (UNAM).
texanusHood, 1912: 157-9. Holotype $, U.S.A.: Texas (USNM).
furfeinafus(Ananthakrishnan, 1967: 233-4) (Percnothrips) . Holotype $, INDIA (TNA). Comb. n.
xosa (Jacot-Guillarmod, 1939ft: 43-6) (Dichaetothrips). Holotype $, SOUTH AFRICA (AMG). Comb. n.
Genera of Diceratothripina
Karny (1925c) erected this group as a subfamily for 11 genera, including some now placed in the
Phlaeothripinae. Priesner (1961) used it as a subtribe of his Cryptothripini, but included a wide
range of genera whose main common characteristic was the large body size of most species. The
present reclassification derives largely from recognition of the phylogenetic significance of the
presence of metathoracic sternopleural sutures. The Diceratothripina is thus defined as those
Pygothripini which possess these sutures (with a few exceptions) as well as four sense cones on
antennal segment IV and widely spaced (usually V-shaped) maxillary stylets. The group is large
and diverse, and represented in all parts of the tropics and subtropics. However, the Nesothrips-
group (Campulothrips, Carientothrips, Nesidiothrips and Nesothrips + Rhaebothrips) is found
mainly in the Australian and Pacific regions. This genus-group probably shared an ancestor with
Acallurothrips and Neosmerinthothrips, but species of these two pantropical genera exhibit a
tendency for the tube to be swollen or at least to have convex margins. This is also found in
Phacothrips and the new species of Diceratothrips described below; these two genera, together
with Sporothrips, constitute the New World element in the Diceratothripina. Two genera
described from Africa, Elgonima and Pseudoeurhynchothrips, are each based on a single
damaged individual on which a full range of characters is not visible. Pseudoeurhynchothrips has
a large foretarsal tooth in the female, as in Neosmerinthothrips-group but unlike Nesothrips-
group, and the straight-sided tube of the only known specimen probably represents a reversion
from the convex form found in Neosmerinthothrips species.
ACALLUROTHRIPS Bagnall
(Figs 145, 146, 161, 168, 179, 180)
Acallurothrips Bagnall, 1921a: 269. Type-species: Acallurothrips macrurus Bagnall, by original designa-
tion.
Diopsothrips Hood, 1934: 422-3. Type-species: Diopsothripsflavus Hood, by original designation. Syn. n.
This genus was erected for two species from the Seychelles, each of which was known only from a
single damaged specimen (Mound, 1968). Moulton described a third species, latus from Fiji, but
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 41
the genus has never been redefined. However, one of Bagnall's species, proturus, is here
transferred to Gastrothrips, and macrurus and latus are recognised as being closely related to
spinicauda Priesner together with several other species originally described in Pygothrips.
Acallurothrips , which differs from Pygothrips in head shape and stylet position, may be defined
as follows.
Head broad, maxillary stylets wide apart (Figs 145, 146). Antennae with segments VII-VIII broadly joined
(rarely fused); sense cones usually arising laterally, 2 on III, 4 on IV, sometimes long and curved in large
species (Fig. 168). Epimeral sutures usually complete. Praepectus present (Fig. 145); mesopraesternum
often eroded; metathoracic sternopleural sutures present, but area posterior to mesocoxae as well as
anapleural suture and anterior border of anepisternum often eroded into chitinous islets. Foretarsal tooth
present in both sexes. Forewing without duplicated cilia (except breviceps); sub-basal seta B3 long.
Metanotum weakly reticulate, median setae long and widely spaced. Pelta broadly reticulate, frequently
eroded at posterior margin (Figs 179, 180); median sternites usually longer than tergites. Wing-retaining
setae usually weak; tube greatly expanded with margins convex, often ridged near base, apex constricted
(Fig. 161).
Fifteen species (including two from Diopsothrips and one from Lathrobiothrips} have been
studied and found to agree with the above definition, and a further species is referred to this
genus from Diopsothrips on the basis of its original description. A short series of specimens of
metulicauda, collected in Malaya and Java, suggests that the size and colour of the tube are
variable in this species at least. Moreover, antennal segment IV usually bears four sense cones,
but individuals have been studied with only two, or even with two on one antenna and four on
the other. Contrary to Mound (1968), the damaged holotype of macrurus has four sense cones
on antennal segment IV. Unfortunately, many of the described species have only been collected
once, and so little information is available on intraspecific variation in this genus.
Most Acallurothrips species have antennal segments VII-VIII broadly joined, but these
segments are fused without trace of a suture \nflavus (Fig. 168) and louisianae (also brunneus?) .
This difference is not accepted as a basis for recognising Diopsothrips as a useful genus. The
pronotal epimeral sutures are complete in flavus but incomplete in the closely similar species
louisianae. Hood compared Diopsothrips to Symphyothrips in his original description, and this
led Stannard (1957) to place the genus in synonymy with Polyphemothrips.
Most of the species listed below in Acallurothrips are small in size. However, quadraticeps is
large, with the head almost as long as wide, the stylets only one-third of the head width apart,
and the sense cones on antennal segments III-IV not lateral in origin. This species resembles
Phacothrips ocelloides in general appearance. Acallurothrips is probably most closely related to
Neosmerinthothrips (q.v.) in which the species usually have the tube margins slightly convex.
Pygothrips is here regarded as being more distantly related, despite the many similarities in tube
and abdomen, because of the long closely approximated maxillary stylets. Faure described
judithae in Lathrobiothrips because of the enlarged tube, but that genus is here placed in the
Phlaeothripinae - Docessissophothripini as a synonym of Holothrips.
SPECIES INCLUDED
amp/us (Faure, 1949c: 118-22) (Pygothrips). Holotype $, SOUTH AFRICA (NCIP). Comb. n.
badius (Faure, 1955: 35-40) (Pygothrips). Holotype $, SOUTH AFRICA (NCIP). Comb. n.
6reviceps(Hood, 1934: 419-20) (Pygothrips). Holotype $, PANAMA (USNM). Comb. n.
*&riwineiis(Hood, 1934: 424-5) (Diopsothrips). Lectotype $, PANAMA (USNM). Comb. n.
conifer (Hood, 19256: 67) (Pygothrips). Holotype $, TRINIDAD (USNM). Comb. n.
/ascio/afus(Hood, 1952c: 165-6) (Pygothrips). Holotype $, BRAZIL (USNM). Comb. n.
flavus (Hood, 1934: 423-4) (Diopsothrips). Lectotype $, PANAMA (USNM). Comb. n.
judithae (Faure, 1956: 321-30) (Lathrobiothrips). Holotype $ , SOUTH AFRICA (NCIP). Comb. n.
7afusMoulton, 1944: 289-90. Holotype $, FIJI (BPBM).
/ouisia/iae(Hood, 1936a: 98-100) (Diopsothrips). Holotype $, U.S.A.: Louisiana (USNM). Comb. n.
macrurusBagnall, 1921a: 270-1. Holotype $, SEYCHELLES (BMNH).
/naini7/Jcaiida(Hood, 1954c: 208-10) (Pygothrips). Holotype $, TRINIDAD (USNM). Comb. n.
metulicauda (Karny, 1923: 336-^0) (Pygothrips). Syntypes ?$, JAVA (SMF). Comb. n.
noguttii( Kurosawa, 1932: 234-8) (Pygothrips). Holotype <j>, JAPAN (7NIAT). Comb. n.
42 L. A. MOUND AND J. M. PALMER
quadraticeps(Hood, 1952c: 166) (Pygothrips). Holotype $, BRAZIL (USNM). Comb. n.
spinicauda (Priesner, 1939ft: 57-9) (Pygothrips). Holotype <J>, CONGO (SMF). Comb. n.
CAMPULOTHRIPS Moulton
(Figs 133, 148, 151,162)
Campulothrips Moulton, 1944: 310-1. Type-species: Campulothrips gracilis Moulton, by monotypy.
This genus has previously been considered to be related to the Idolothripini, but this is not
accepted here because of the presence of short metathoracicsternopleural sutures (Fig. 151) and
a single pair of wing-retaining setae on each tergite. These characters, together with the large
pair of setae between the posterior ocelli (Fig. 133) and the enlarged L-shaped femora of males
(Fig. 148), suggest that the genus is derived from Nesothrips, within whose zoogeographic range
(the Pacific) it occurs. However, the elongate antennae, long median setae on the metanotum,
tube with prominent lateral setae at least in the basal half (Fig. 162) and rather swollen
katepimera (albeit with complete anapleural sutures) are remarkable examples of convergent
evolution toward the typical body form of some South American Idolothripini.
SPECIES INCLUDED
gracilis Moulton, 1944: 311. Holotype $, FIJI (BPBM).
CARIENTOTHRIPS Moulton
(Figs 135-137, 164, 181-185)
Bolothrips (Carientothrips) Moulton, 1944: 306. Type-species: Bolothrips (Carientothrips) fijiensis
Moulton, by monotypy.
This group was first recognised as a full genus by Mound (191 4a) who defined it and discussed the
generic relationships together with a key to 17 species (19746) . These species are found mainly in
the Australian and Pacific regions, although denticulatus is known only from Tierra del Fuego
and the Falkland Islands. In general appearance several species resemble Bolothrips species, the
body being slender, pelta broad (Figs 181-185) and eyes prolonged ventrally (Fig. 137), but this
is probably due to convergent evolution through adaption to a similar habitat at the base of
grasses. The species found on dead branches tend to be less slender and in macropterae the pelta
has distinctive broad lateral wings (Fig. 183). The metathoracic sternopleural sutures in most
species of Carientothrips are clearly defined, but in a few species are narrow and difficult to see,
and in badius are not developed at all. The genus appears to represent an holophyletic
species-group derived from Nesothrips, or it may represent the sister-group of that genus.
Individual species of these two groups are frequently difficult to place in a genus.
SPECIES INCLUDED
acti Mound, 1974a: 25-6. Holotype $, AUSTRALIA (ANIC).
badius (Hood, 19186: 143-4) (Cryptothrips). Holotype $, AUSTRALIA (USNM).
apterus Girault, 19286: 2 (Elaphrothrips) . Holotype cf , AUSTRALIA (QMB).
biformis (Moulton, 1939: 146-7) (Bolothrips). Holotype $, TAHITI (BPBM).
capricornis (Mound, 1974o: 23-4) (Bolothrips). Holotype $, AUSTRALIA (ANIC).
casuarinae Mound, 19740: 26-9. Holotype $, AUSTRALIA (ANIC).
denticulatus (De Santis, 19636: 66) (Nesothrips). Holotype $, ARGENTINA (MLPA).
fijiensis (Moulton, 1944: 306-7) (Bolothrips). Holotype $, FIJI (BPBM).
grayi Mound, 19746: 129. Holotype $, NEW GUINEA (BMNH).
japonicus(B agnail, 19216: 355-6) (Cryptothrips). Holotype 9, JAPAN (BMNH).
loisthus Mound, 1974a: 29-30. Holotype $, AUSTRALIA (ANIC).
magnetis Mound, 1974a: 30-1. Holotype $, AUSTRALIA (ANIC).
miskoi Mound, 1974a: 31. Holotype $, AUSTRALIA (ANIC).
mjobergi (Karny , 1920c: 42) (Cryptothrips). Holotype $, AUSTRALIA (NRS).
incisus Girault, 1927c: 1 (Cryptothrips). Syntypes $, AUSTRALIA (QMB).
australicus Priesner, 19286: 649-51 (Cryptothrips). Holotype $, AUSTRALIA (SMF).
flavitibia Moulton, 1968: 117-8 (Bolothrips). Holotype $, AUSTRALIA (CAS).
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 43
pedidllus Mound, 1914a: 32-3. Holotype $, AUSTRALIA (ANIC).
pictilis Mound, 1914a: 33-4. Holotype $, AUSTRALIA (ANIC).
reed/Mound, 1974a: 34-5. Holotype $, AUSTRALIA (ANIC).
semiru/iis(Girault, 19286: 4) (Elaphrothrips) . Holotype $, AUSTRALIA (QMB).
vesper Mound, 1974a: 35-6. Holotype $, AUSTRALIA (ANIC).
DICERA TOTHRIPS Bagnall
(Figs 35, 130, 131, 149, 152, 163, 165, 171)
Diceratothrips Bagnall, 19086: 193. Type-species: Dicer atothrips bicornis Bagnall, by monotypy.
Eulophothrips Schmutz, 1909: 278. Type-species: Eulophothrips robustus Schmutz, by monotypy.
[Synonymised by Priesner, 1949: 129.]
Megalomerothrips Watson, 1919: 99. Type-species: Megalomerothrips eupatorii Watson, by monotypy.
[Synonymised by Priesner, 1949: 136-7.]
Diceratothrips (Endacnothrips) Priesner, 1933c: 147-9. Type-species: Diceratothrips (Endacnothrips)
horridus Priesner, by monotypy.
This genus has been subject to much confusion. Mound (1968), following Stannard (1957),
treated Diceratothrips as a senior synonym of Dichaetothrips and also included the Ethirothrips-
group of species from the Old World. At that time, however, Mound (and probably Stannard)
had not examined the type-species of Dichaetothrips. Subsequently, it has been realised that
Diceratothrips is a genus of Neotropical species which can be distinguished from similar-looking
Old World species by the presence of long, well-developed sternopleural sutures on the
metathorax (Fig. 152). Males, but not females, of all species of Diceratothrips examined in this
study have a stridulatory file on the external margin of the forecoxae, and the flattened edge of
the forefemora apparently functions as a plectrum (Fig. 149). The members of this genus share
the following characteristics.
Usually large, black to dark brown species; head often with anteocellar setae long or stout, and cheeks with
stout setae; stylets wide apart (Figs 130, 131). Antennae 8-segmented, III relatively long, VII- VIII distinct
but sometimes forming a single unit (Fig. 165); III with 2 sense cones, IV with 4 sense cones; IV- VI
prolonged ventrally. Pronotum broad and flat, scarcely thickened at anterior or medially; reticulate in
anterior third; am, aa and ml setae small (Fig. 131). Praepectus present, mesopraesternum with
posterolateral corners almost forming a right-angle; metathoracic sternopleural sutures well-developed
(Fig. 152). Forefemora often with stout spines on inner surface in both sexes; foretarsal tooth large or very
reduced; forewings broad, with numerous duplicated cilia. Pelta not exceptionally broad, curving away
from tergite II laterally (Figs 35, 171); tergites each with one pair of wing-retaining setae.
In contrast to the Old World Ethirothrips group of species, allometric growth patterns are found
commonly in Diceratothrips species. Therefore, since many of these species have been described
from few specimens and, moreover, have never been compared directly with their congeners, a
number of synonyms can be expected. For example, Hood (1934: 70) in describing a new species
princeps, listed several characteristics of four species, armatus, bicornis, persimilis and robustus,
none of which he personally had ever examined. Many of the details he gives are incorrect, and
the first three of these names are here treated as synonymous. Moreover, princeps is almost
certainly the same species, robustus and cornutus are possibly only variants with exceptionally
long anteocellar setae, and even inferorum may also be the same but with antennal segment III
slightly paler. Material identified as bicornis has been examined from the following countries:
Trinidad, Venezuela, Brazil, Peru and Mexico (in BMNH). Moreover, the unique holotype of
williamsi Karny from Guatemala has been studied and is here regarded as the same species as
bicornis. Contrary to the original description and illustration of williamsi, the stout ocellar setae
arise anterolateral to the ocellar triangle, not behind the posterior ocelli in this holotype.
Not only does bicornis exhibit considerable allometric variation in both sexes, it also shows
marked sexual dimorphism in the shape of the head and forefemora. Most of the other species of
Diceratothrips, in both sexes, resemble the females of bicornis rather than the males. Moreover,
several species have the anteocellar and cheek setae reduced in size, e.g. delicatus, harti,
setigenis and validipennis . Of these, only delicatus has stout setae on the inner surface of the
44 L. A. MOUND AND J. M. PALMER
forefemora, whereas the other species have the femora long and broad. The two species setigenis
and pallidior, described from the coast of the Gulf of Mexico, may represent one variable
species. The new species bennetti, described below from Trinidad, is exceptionally small and
Pygothrips-like, with the tube swollen, antennal projections reduced, forefemora short and
swollen without stout setae on the inner surface, and head short and broad with two pairs of stout
cheek setae. Similarly, nigricauda which was described in Pygothrips has the tube enlarged, the
stylets about one-third of the head width apart, the forewing with seven duplicated cilia, pelta
not eroded (Fig. 35), and the forecoxae with transverse striations.
SPECIES INCLUDED
* ana/i uacens/sJohansen, 1976: 59-61. Holotype $, MEXICO (UNAM).
bennetti sp. n. Holotype cf , TRINIDAD (BMNH).
Wcom/sBagnall, 19086: 194-5. Holotype $, BRAZIL (BMNH).
armatus Bagnall, 1910a: 385-6. Syntypes $ cf , VENEZUELA (BMNH). Syn. n.
williamsi Karny, 1920a: 92-4 (Dichaetothrips) . Holotype $, GUATEMALA (DEI). Syn. n.
persimilis Priesner, 19256: 25-6. Holotype $, SURINAM (SMF). Syn. n.
*cormifusHood, 1952c: 156-7. Holotype $, BRAZIL (USNM).
*cu6ens/sHood, 1941: 178-80. Lectotype $, CUBA (USNM).
detfcafusHood, 1941: 171-4. Holotype $, U.S.A.: Florida (USNM).
Aart/ Hood, 1912a: 12-4. Holotype cf , U.S.A.: Texas (USNM).
*eupatorii Watson, 1919: 99-100 (Megalomerothrips). Holotype $, U.S.A.: Florida (FDA).
*hercules Johansen, 19776: 59-61. Holotype cf , MEXICO (UNAM).
horridus Priesner, 1933c: 147-9. Holotype cf , MEXICO (SMF).
inferorum (Priesner, 19330: 62-3) (Adiaphorothrips}. Holotype $ (not cf ), MEXICO (SMF).
*70ngjpesHood, 1912a: 14-5. Holotype cf , U.S.A.: Texas (USNM).
nigricauda (Hood, 19256: 67-8) (Pygothrips}. Holotype $, TRINIDAD (USNM). Comb. n.
*o5scuricornisHood, 1941: 174-6. Holotype $, CUBA (USNM).
pallidior Priesner, 1933c: 151. Holotype $, MEXICO (SMF).
picticornisHood, 1914: 166-7. Holotype $, PANAMA (USNM).
*wolcotti Morgan, 1925: 8-9. Holotype $, PUERTO Rico (USNM).
princepsHood, 1934: 68-71. Holotype cf , PANAMA (USNM).
*robustus (Schmutz, 1909: 278-81) (Eulophothrips). Holotype cf , BRAZIL (? lost).
*sakimurai Johansen, 19776: 61-2. Holotype $, MEXICO (UNAM).
setigenis Hood, 1941: 176-8. Lectotype $, U.S.A.: Texas (USNM).
*timidus Johansen, 1976: 61-2. Holotype $, MEXICO (UNAM).
validipennis(Hood, 1938c: 403-6) (Gastrothrips). Holotype $, U.S.A.: Florida (USNM).
Diceratothrips bennetti sp. n.
(Figs 131, 152, 163, 165, 171)
Macropterous cf . Colour dark brown, tube black; distal half of antennal segment II, and segment III
except at apex, brownish yellow; major setae dark brown; wings shaded particularly at base and apex, but
with no longitudinal line.
Head short and broad with two pairs of cheek setae (Fig. 131); anteocellar setae short. Antennae with
sense cones on III very short (Fig. 165). Forefemora swollen on inner surface but with no stout setae,
posterior angle acute and extending to striate area on forecoxae; foretarsal tooth almost as long as tarsal
width. Forewing sub-basal seta B2 arising posterolateral to B^. Pelta broadly triangular (Fig. 171); tergites
II- VI each with one pair of wing-retaining setae; median sternites slightly longer than tergites; postero-
lateral abdominal setae long and stout; tube stoutly conical.
Measurements (holotype cf in /xm). Body length 2350. Head, length 285 (tilted); width 220; postocular
setae 105. Pronotum, length 150; width 300; major setae - am 15, aa 15, ml 15, epim 110/15, pa 25.
Forewing, length 950; width 90; sub-basal setae 40, 60, 100; number of duplicated cilia 18. Tergite IX setae
«! 220; B2 ?; B3 220. Tube, length 240; basal width 120; terminal setae 90. Antennal segments III- VIII
length 105, 93, 75, 65, 60, 30.
Macropterous 9 • Colour and structure similar to cf but larger; forefemora swollen, with posterior angle
rounded and not extending to forecoxae; anteocellar setae stout; pronotum not as broad and flat as in most
species of this genus, with a weak line of thickening anteromedially.
Measurements (paratype $ in /u,m). Body length 2750. Head, length 270; width 255; postocular setae
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 45
140. Pronotum, length 165; width 350; major setae - am 18, aa 18, ml ?15, epim 150, pa 45. Metanotal
median setae 45. Forewing, length 1100; distal width 120; sub-basal setae 40, 60, 120; number of duplicated
cilia 22. Tergite IX setae Bv ?; B2 ?; B3 240. Tube, length 255; basal width 140; terminal setae 90. Antennal
segments III-VIII length, 115, 100, 80, 70, 70, 30.
SPECIMENS STUDIED
Holotype cf , Trinidad: Curepe, on grasses, 8.xi.l970 (L. A. Mourn/ 921) (BMNH).
Paratype. 1 $ collected with holotype (BMNH).
COMMENTS. This new species was collected whilst studying at the headquarters of the Common-
wealth Institute of Biological Control at the invitation of the Director, Dr Fred Bennett. Only
two specimens were found, and these were in rough grassland. However, the structural
adaptations of bennetti are such as might be expected of a species of Dicer atothrips adapted to
grass-living rather than the typical habitat of dead branches. The sternites are relatively long, the
posterior abdominal setae stout, and the tube enlarged, all of which suggest that the species
raises the tube over the head in life, as do species of Acallurothrips and Pygothrips, as well as
Nesothrips propinquus .
ELGONIMA Zur Strassen
Elgonima Zur Strassen, 1972: 91. Type-species: Elgonima seticeps Zur Strassen, by monotypy.
This genus is based on a single, imperfect, macropterous female. Due to contraction of the
pterothorax it is impossible to determine the presence or absence of metathoracic sternopleural
sutures. However, seticeps is apparently typical of the Diceratothripina. Antennal segment VIII
is slender, but the sense cones on III-IV are unusually long for species in diceratothripine
genera. The pronotal epimeral sutures are incomplete, and all the major setae have expanded
apices, including those on tergite IX and one pair of postocellars. The relationships of Elgonima
cannot be determined with certainty at present.
SPECIES INCLUDED
seticepsZur Strassen, 1972: 91-3. Holotype $, KENYA (NRS).
NEOSMERINTHOTHRIPS Schmutz
(Figs 132, 138, 139, 153-156, 169, 172, 173)
Neosmerinthothrips Schmutz, 1913: 1051. Type-species: Neosmerinthothrips fructuum Schmutz, by
monotypy.
Coenurothrips Bagnall, 1921a: 271. Type-species: Coenurothrips brevlcollis Bagnall, by original designa-
tion. [Synonymised by Mound, 19746: 148.]
Galactothrips Moulton, 1933a: 404. Type-species: Galactothrips diversicolor Moulton, by monotypy.
[Synonymised by Mound, 19746: 148.]
This genus was redefined and discussed by Mound (19746) with a key to 18 species. These
comprise a small New World species-group, and a larger and more diverse Old World
species-group. However, the genus is distinguished with difficulty from the equally widespread
tropical genus Acallurothrips. Members of the latter genus have the tube more strongly swollen,
antennal segments VII-VIII broadly joined or fused, and the body sclerites frequently reduced.
Thus future research may indicate that neither of these genera represent holophyletic groupings.
Pseudoeurhynchothrips bidens is similar to some Neosmerinthothrips but has the tube long with
straight margins. The poorly preserved syntypes of Barythrips grandicauda have been examined
and are here interpreted as the only known micropterous species of Neosmerinthothrips.
However, they are similar to Acallurothrips species in having the setae on tergite IX scarcely half
as long as the tube, the pelta reduced and eroded medially on the posterior margin, and the
metanotum transverse with a pair of long setae. These two syntypes were probably collected in
the Oriental Region, the name 'Frauenfeld' on the slide almost certainly refers to the
well-known entomologist who worked in southern Europe. In some Neosmerinthothrips the
postocellar setae are elongate; in nigrisetis they are as long as the sides of the ocellar triangle, but
in the new species hamiltoni described below they are even longer (Fig. 132).
46 L. A. MOUND AND J. M. PALMER
SPECIES INCLUDED
a/J?nis(Bagnall, 19216: 361-2) (Coenurothrips) . Holotype $, SRI LANKA (BMNH).
annu//pes(Hood, 1950: 13-6) (Gastrothrips). Holotype <j>, BRAZIL (USNM).
milleforme De Santis, 1963a: 12-4 (Nesothrips). Holotype $, ARGENTINA (MLPA).
ftrev/co77js(Bagnall, 1921a: 271-2) (Coenurothrips}. Syntypes cf $, SEYCHELLES (BMNH).
co77ar/s(Bagnall, 1917: 26-7) (Cryptothrips). Lectotype $, ST VINCENT (BMNH).
fuscicauda Morgan, 1925: 6-7 (Gastrothrips). Holotype cf , PUERTO Rico (USNM).
marshalli Priesner, 1934: 58-60 (Bolothrips). Lectotype $, SIERRA LEONE (BMNH).
dominicanus Hood, 19356: 170-4 (Gastrothrips). Holotype cf , DOMINICAN REPUBLIC (USNM).
diversic07or(Moulton, 1933o: 404-6) (Galactothrips). Holotype $, BRAZIL (CAS).
/J/ieiJsis(Moulton, 1944: 286-7) (Gastrothrips). Holotype $, FIJI (BPBM).
/riicfiJumSchmutz, 1913: 1052-3. Lectotype cf , SRI LANKA (Ceylon) (SMF).
ceylonicus Karny, 1925c: 137-9 (Oedemothrips). Holotype $, SRI LANKA (BMNH).
grandicauda (Priesner, 19256: 21) (Barythrips). Syntypes cf $, ?ORIENTAL REGION (SMF). Comb. n.
hamiltonisp. n. Holotype $, BRAZIL (BMNH).
hilaris (Priesner, 19376: 624-6) (Bolothrips). Holotype cf , SIERRA LEONE (BMNH).
7ioodi(Faure, 1954a: 9-13) (Gastrothrips). Holotype $, SOUTH AFRICA (NCIP).
*MigiM/Miii*Ananthakrishnan, 1960: 32-3. Holotype <j>, INDIA (?TNA).
nigrisetis(Hood, 19356: 161-5) (Gastrothrips). Holotype $, PANAMA (USNM).
parv/dens(Hood, 19356: 165-8) (Gastrothrips). Holotype $, PANAMA (USNM).
pau//srarum(Hood, 1950: 25-7) (Gastrothrips). Holotype $, BRAZIL (USNM).
picticornis(Hood, 1936c: 272-5) (Gastrothrips). Holotype $, BRAZIL (USNM).
p7aumaiifl/(Hood, 1950: 20-2) (Gastrothrips). Holotype $, BRAZIL (USNM).
rabusfus(Ananthakrishnan, 1964a: 102-3) (Nesothrips). Syntypes cf $, INDIA (TNA).
vari/pes(Hood, 1950: 16-20) (Gastrothrips). Holotype $, BRAZIL (USNM).
xylebori Priesner, 1935c: 370. Lectotype <j>, JAVA (SMF).
Neosmerinthothrips hamiltoni sp. n.
(Figs 132, 154, 169, 172)
Macropterous $. Colour dark brown, head and tube black; antennal segment III yellow with apical third
light brown, IV yellowish brown in basal half, V slightly paler at base than apex, II yellow apically; major
setae dark brown; forewing clear except around sub-basal setae.
Head slightly narrowed to base, weakly sculptured laterally (Fig. 132); postocellar setae extending to
apex of antennal segment II; maxillary stylets wide apart, retracted to postocular setae; mouth cone
broadly rounded. Antennal segment III slender with two sense cones; IV with four sense cones; VIII
slightly narrowed to base (Fig. 169). Pronotum transverse, epimeral sutures complete (Fig. 132);
anteromarginal setae short; praepectus present, mesopraesternum broadly boat-shaped. Foretarsal tooth
slender, two-thirds as long as tarsal width. Mesonotal lateral setae very small. Metanotum scarcely
sculptured medially. Anapleural sutures complete, katepisternum eroded anteriorly; metathoracic sterno-
pleural sutures long. Forewing broad, sub-basal setae arising close together in a straight line. Pelta with
short broad lateral wings (Fig. 172). Tergites II-VII with one pair of wing-retaining setae, sigmoid on
III- VI; setae on IX elongate; tube with slightly sinuate, convex margins (Fig. 154). Sternites with about 12
rather weak discal setae.
Measurements (holotype $ in /mi). Body length 3400. Head, length 360; width behind eyes 275;
postocellar setae 190; postocular setae 240. Pronotum, length 150; width 360; major setae - am 25, aa 60,
ml 135/165, epim 225, pa 195. Metanotal median setae 30. Forewing, length 1300; distal width 150;
sub-basal setae 30, 150, 180; number of duplicated cilia 20. Tergite IX setae Bv 360, B2 420. Tube, length
345; basal width 120; terminal setae 270. Antennal segments III-VIII length, 135, 120, 100, 75, 50, 40.
Macropterous cf . Colour and structure similar to $ but smaller, body length 2500, head length 270;
postocellar setae 160.
SPECIMENS STUDIED
Holotype $>, Brazil: S.P., Ribeirao Preto, FFCLRP Campus, in hollow twig of Glyricidia, 7.ix.l975
(W. D. Hamilton) (BMNH).
Paratypes. 9 $, 2 cf collected with holotype (BMNH).
COMMENTS. This belongs to the South American species-group of Neosmerinthothrips which
includes nigrisetis and variipes, but is readily distinguished by its larger size with more slender
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 47
antennae and tube, and the remarkably long postocellar setae. This new species was at first
considered to represent a Dichaetothrips , but the presence of metathoracic sternopleural sutures
as well as the structure of the pelta and short antennal sense cones indicate that it belongs in
Neosmerinthothrips .
NESIDIOTHRIPS Mound
(Figs 134, 157, 174)
Nesidiothrips Mound, 19746: 156-7. Type-species: Nesothrips alias Ananthakrishnan, by original designa-
tion.
This genus was erected for two species which have most characters similar to those oi Nesothrips
species, but with a pair of stout setae within the ocellar triangle (Fig. 134) and the females with a
large foretarsal tooth.
SPECIES INCLUDED
alius( Ananthakrishnan, 1970: 52-5) (Nesothrips) . Holotype $, INDIA (TNA).
vatfdiis (Bagnall, 1921a: 272-3) (Coenurothrips) . Holotype $, SEYCHELLES (BMNH).
NESOTHRIPS Kirkaldy
(Figs 142-144, 159, 160, 167, 175-178)
Nesothrips Kirkaldy, 1907: 103. Type-species: Nesothrips oahuensis Kirkaldy, by monotypy.
Oedemothrips Bagnall, 19106: 680. Type-species: Oedemothrips laticeps Bagnall, by monotypy. [Synony-
mised by Bianchi, 1944.]
Rhaebothrlps Karny, 1913c: 128. Type-species: Rhaebothrips lativentris Karny, by monotypy. Syn. n.
Mound (19746) discussed the relationship between Nesothrips and Rhaebothrips and gave keys
to the world species of these two genera. At that time they were distinguished on the basis of the
relatively short tube in Nesothrips and the relatively long and closely approximated ocellar setae
in Rhaebothrips, but it was predicted that further studies on the Pacific fauna would erode these
small differences. The new species described below from New Zealand further reduces the
distance between the two groups and so they are here treated as one genus.
The type-species of Rhaebothrips is now found in most parts of the tropics, associated with
coconut fronds and fruits, but it is almost certainly native to the Pacific area. This species,
lativentris, is variable and has eight junior synonyms; like nigrisetis to which it is closely related,
it has a relatively slender head and pale antennal segment III. A further Pacific species, major, is
larger with dark antennae and even longer, more closely set ocellar setae. None of these species,
in common with all the species described in Nesothrips, has a foretarsal tooth in the females. In
contrast leveri from Fiji has a shorter head but long and close-set ocellar setae, and the female
bears a foretarsal tooth. Moreover, three species from New Zealand described by Mound
(19746) have the head intermediate in length with the ocellar setae relatively far apart and
arising between or behind the posterior ocelli (Fig. 144); one of these, doulli, has a foretarsal
tooth in the female. Finally the new species rangi, described below, has the head even broader
and more typically Nesothrips-\ike , but has long ocellar setae (Fig. 143). The head shape and
length of the ocellar setae already exhibit a wide range in Nesothrips species (Figs 142-144).
Almost all the species ofNesothrips have well-developed metathoracic sternopleural sutures,
although these are not present in oahuensis and melinus and are highly variable (from
well-developed to absent) in propinquus (Mound & Walker, 1983). Carientothrips (q.v.) is
closely related to Nesothrips and cannot be distinguished on any single character. Campulothrips
with one species appears to be a specialised derivative of the lativentris-major species-group.
SPECIES INCLUDED
aoristus Mound, 1974a: 68. Holotype $, AUSTRALIA (ANIC).
arfocarpi(Moulton, 19426: 14-5) (Bolothrips). Holotype $, GUAM (BPBM).
brevicollis (Bagnall, 19146: 29-30) (Oedemothrips). Holotype $, JAPAN (BMNH).
minor Bagnall, 1921o: 287-8 (Coenurothrips). Holotype £, RODRIGUES (BMNH).
48 L. A. MOUND AND J. M. PALMER
formosensis Priesner, 1935c: 368-70 (Neosmerinthothrips). Lectotype $, TAIWAN (SMF).
formosensis var. karnyi Priesner, 1935c: 369-70. Lectotype $, JAVA (SMF).
carver/Mound, 1974a: 71. Holotype $, AUSTRALIA (ANIC).
doulli (Mound, 1974ft: 171-3) (Rhaebothrips). Holotype $, NEW ZEALAND (BMNH). Comb. n.
eastopi (Mound, 1974ft: 173-4) (Rhaebothrips). Holotype $, NEW ZEALAND (BMNH). Comb. n.
fodinae Mound, 1974ft: 163^. Holotype $, FIJI (BMNH).
hemidiscus Mound, 1974a: 71-2. Holotype $, AUSTRALIA (ANIC).
/afiVenfr/s(Karny, 1913c: 129-30) (Rhaebothrips). Holotype cf , TAIWAN (? lost). Comb. n.
claripennis Hood, 1919ft: 90 (Cryptothrips). Holotype $, AUSTRALIA (USNM).
seychellensis Bagnall, 1921a: 274-6 (Cryptothrips). Lectotype cf , SEYCHELLES (BMNH).
difficilis Bagnall, 1921a: 276 (Cryptothrips). Holotype $, SEYCHELLES (BMNH).
magnus Moulton, 1928c: 299 (Cryptothrips). Holotype $, TAIWAN (CAS).
yuasaiMoulton, 1928d: 315 (Gynaikothrips). Holotype $, TAIWAN (CAS).
ipomoeae Ishida, 1932: 12-4 (Machatothrips). Holotype $, PONAPE (Hokkaido Univ.).
fuscus Moulton 1942ft: 15-6 (Rhaebothrips). Holotype $, GUAM (BPBM).
australiensis Moulton, 1968: 118-9 (Bolothrips). Holotype $, LORD HOWE Is. (CAS),
/ever/ (Mound, 1974ft: 175) (Rhaebothrips). Holotype $, FIJI (BMNH). Comb. n.
major (Bagnall, 1928: 75-6) (Rhaebothrips). Holotype cf , SAMOA (lost).
maJaccae Mound, 1974ft: 164-6. Holotype $, WEST MALAYSIA (BMNH).
melinus Mound, 1974a: 72-3. Holotype $, AUSTRALIA (ANIC).
niger (Moulton & Steinweden, 1932: 167-8) (Bolothrips). Holotype $, MARQUESAS (BPBM).
nigrisetis (Sakimura, 1972: 400-2) (Rhaebothrips). Holotype cf , FIJI (BPBM). Comb. n.
oa/mens/sKirkaldy, 1907: 103. Syntype $, OAHU (? BPBM).
laticeps Bagnall, 1910ft: 680-1 (Oedemothrips) . Syntypes cf $, OAHU (BMNH).
hawaiiensis, lapsus for oahuensis, Bianchi, 1944: 31-8.
propinquus (Bagnall, 1916: 408-9) (Oedemothrips). Holotype $, AUSTRALIA (BMNH).
dimidiatus Hood, 1918ft: 145-6 (Cryptothrips). Holotype $, AUSTRALIA (USNM).
cestosa Karny, 1920c: 41; 1921ft: 33-6 (Bagnalliella). Holotype $, AUSTRALIA (NRS).
propinquus var. breviceps Bagnall, 1924: 634-5 (Oedemothrips). Syntypes $, NEW ZEALAND
(BMNH).
propinquus f. obscuricornis Bagnall, 1924: 634. Types not designated.
oleriae Moulton, 1949: 492-4 (Neosmerinthothrips}. Holotype $, SOUTH AFRICA (CAS).
similis Hartwig, 1948: 103-8 (Bolothrips). Holotype $, SOUTH AFRICA (NCIP).
rangisp. n. Holotype cf , NEW ZEALAND (NZAC).
rhizophorae (Girault, 1927: 2) (Cryptothrips). Syntypes cf $, AUSTRALIA (QMB).
semiffavus (Moulton, 1939: 147-8) (Bolothrips). Holotype $, RAPA (BPBM).
yanchepi Mound, 19740: 75. Holotype $, AUSTRALIA (ANIC).
zondagi (Mound, 1974ft: 176-7) (Rhaebothrips). Holotype $, NEW ZEALAND (NZAC). Comb. n.
Nesothrips rangi sp. n.
(Figs 143, 167, 178)
$ macroptera. Body dark brown, head and tube darkest; legs dark, extreme apex of femora and base
of tibiae yellow, foretibiae yellowish brown, all tarsi paler; antennae variable in colour, III yellow with
apex more or less brown, IV yellow in basal half but sometimes light brown, V with pedicel yellow or
uniformly brown, I and VI-VIII dark brown, II variably yellow at apex; forewings dark at base,
weakly shaded distally; major setae dark brown.
Head almost as wide as long, weakly projecting in front, cheeks rounded, compound eyes not large
(Fig. 143); ocelli present but rather small, ocellar setae arising between posterior ocelli and about as
long as distance between their bases; postocular setae long and fine; stylets wide apart and retracted
halfway into head; mouth cone broadly rounded. Antennae 8-segmented; VIII narrowed to base;
VI- VII with constricted pedicels; III with 2 sense cones, IV with 4 sense cones (Fig. 167).
Pronotum transverse, weakly sculptured near posterior margin; all 5 pairs of major setae present
but epimerals longest; praepectal plates well developed (Fig. 143); mesopraesternum broadly
boat-shaped. Mesonotal lateral setae small. Metanotum not elongate (i.e. not fully macropteroid),
median setae slender; sternopleural sutures elongate; anapleural sutures complete. Foretarsi with no
tooth. Forewings parallel-sided with 3 sub-basal setae arising almost in a straight line. Pelta broad,
rounded medially (Fig. 178); wing-retaining setae present on tergites II- VII, sigmoid only on IV-VI;
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 49
anterior tergites with lateral setae short; tergite IX setae more than half as long as tube; tube margins
weakly convex in distal third.
Measurements (holotype $ in /AMI). Body length 2600. Head, length 270; median width 260;
postocular setae 90; ocellar setae 60. Pronotum, length 170; median width 320; major setae - am 30, aa
35, ml 40, epim 110, pa 60. Forewing, length 1000; distal width 100; sub-basal setae 30, 65, 65; number
of duplicated cilia 8(11). Tergite IX setae fl, 120; B2 120; B3 160. Tube, length 220; basal width 95.
Antennal segments III- VIII length 105, 100, 78, 70, 50, 35.
$ microptera. Colour and structure very similar to macroptera, body size a little smaller; meso- and
metanota slightly more transverse; forewing lobe length 150.
Cf microptera. Colour similar to $; usually small in size but variable due to allometry; small cf
similar to $ but with foretarsal tooth present and weak median thickening on pronotum; large cf with
head and pronotum more elongate, epimeral setae stout and posteroangular setae long and fine,
forefemora greatly swollen, tarsal tooth large.
Measurements (small and large paratype cf collected with holotype in /u,m). Body length 1800
(2100). Head, length 225 (260). Pronotum, length 160 (200); major setae - epim 75 (75), pa 52 (90).
Tube length 180 (195).
SPECIMENS STUDIED
Holotype $ macroptera, New Zealand: South Island, 5 ml south of Blenheim, Taylors Pass, at base
oiJuncus in seepage by roadside, 7.ii.l979 (L. A. Mound 1418) (NZAC).
Paratypes (8 $ mac. , 29 $ , 6 d" mic.). New Zealand. South Island: 1 <j> mac, 4 $ mic, 2 cf collected
with holotype; 1 $ mac, 7 $ mic, 1 cf , also 1 $ mac, 5 $ mic collected at two similar sites about 10
miles south of holotype locality (L. A. Mound 1422; A. K. Walker 56); Nelson, Lee Valley, 1 $
hemimac, 1 cf onJuncus, 22.iii.1977 (A. K. Walker); Nelson, Rough Is., 1 $ mac, 1 9 mic on grass,
24. i. 1976 (A. K. Walker); St Arnaud, Lake Rotoiti, 1 $ mac, 6 9 mic, 1 cf swept from swampy
grassland, 9.xii.l980 (A. K. Walker); 10 ml north of Reefton, 1 $ onJuncus, 6.ii.l979 (L. A. Mound
1409); Beaumont, 1 $ at roadside, 17.ii.1976 (G. W. Ramsay); Invercargill, 3 $ mac, 1 <j> mic on
Juncus, 15-17.iii.1977 (A. K. Walker). North Island: Whakatane, 1 $ mic, 18.x. 1978; 10 ml north of
Helensville, 1 $, 1 cf in Cotula swamp, 23. ix. 1973 (B. A. Holloway) (NZAC & BMNH).
COMMENTS. This new species bears some resemblance to rhizophorae Girault from Australia
(Mound, 19740) but has the head much less narrowed to the base, the forewings pale, and the
mid and hind femora not yellow on the distal posterior margins. It appears to be closely related
to the other New Zealand species which were originally described in Rhaebothrips, but unlike all
of them has the median antennal segments more or less yellow basally. The species is evidently
associated with damp grassland rather than native woodland in New Zealand, but unfortunately
no attempt was made to identify the Juncus species on which it was collected to determine
whether or not this was itself native. Micropterae, as opposed to apterae, are not common in
Diceratothripina; one female paratype is hemimacropterous with the wing lobe extending to
tergite II and bearing fringe cilia.
PHACOTHRIPS Mound
(Figs 141, 158, 166, 186)
Phacothrips Mound, 1974ft: 170. Type-species: Gastrothrips ocelloides Hood, by monotypy.
The single species in this genus is difficult to interpret. The body form, with its swollen tube
(Fig. 158), elongate median sternites, and no forewing duplicated cilia, is similar to Acallur-
othrips quadraticeps . However, the mesopraesternum is well developed, antennal segment VIII
slender and distinct from VII (Fig. 166) and the pelta (Fig. 186) similar to Gastrothrips species.
Moreover, the head is unusual in bearing a pair of isolated 'ommatidia' on the cheeks (Fig. 141).
Mound (19746) interpreted the genus as being derived from Neosmerinthothrips , several species
of which have very similar antennae and also a rather 'heavy' tube.
SPECIES INCLUDED
oce/7o«tes(Hood, 1950: 9-12) (Gastrothrips}. Holotype $, BRAZIL (USNM).
50 L. A. MOUND AND J. M. PALMER
PSEUDOEURHYNCHOTHRIPS Moulton
Pseudoeurhynchothrips Moulton, 1949: 482-3. Type-species: Pseudoeurhynchothrips bidens Moulton, by
monotypy.
This genus was based on a single crushed and distorted female specimen with the following
characteristics: antennae 8-segmented, 2 sense cones on III, 4 on IV; eyes slightly reduced?;
postocellar setae about as long as distance between 2 ocelli; stylets broad, wide apart in head?;
praepectus present; mesopraesternum boat-shaped; metathoracic sternopleural sutures short;
anapleural sutures complete; pronotal epimeral sutures complete; foretarsal tooth almost as
long as tarsal width ; pelta with short slender lateral wings ; tergites II-VI with one pair of sigmoid
wing-retaining setae; tube and setae on tergite IX long. This combination of characters is also
found in Dichaetothrips mameti which is therefore also transferred to this genus. Only females of
mameti are known; these have a smaller foretarsal tooth, and the postocellar setae arise in line
with the hind margins of the posterior ocelli. A male labelled as part of the mameti type-series in
the Paris Museum apparently represents Ethirothrips stenomelas. The genus Pseudoeurhyn-
chothrips cannot be distinguished satisfactorily from Nesothrips, but is probably derived from
Neosmerinthothrips through development of a straight-sided tube.
SPECIES INCLUDED
bidens Moulton, 1949: 483. Holotype $, SOUTH AFRICA (BMNH).
mameti (Priesner, 1951: 363) (Dichaetothrips). Holotype $, MAURITIUS (SMF). Comb. n.
SPOROTHRIPS Hood
(Figs 140, 147, 150, 170)
Sporothrips Hood, 1938c: 410. Type-species: Adiaphorothrips amplus Hood, by monotypy.
The single species in this genus could equally well be considered as an aberrant member of
Diceratothrips . The similarities in body form are most evident between the females of the two
genera, although males of amplus bear a similar sound-producing structure on the forecoxae to
that found in Diceratothrips males. The metathoracic sternopleural sutures are very short in
amplus (Fig. 150), but the anapleural sutures are long and curved, ending opposite a small
tubercle, particularly in males. The antennal sense cones are short as in Diceratothrips species,
but VIII is not broadly joined to VII (Fig. 170) and the anteocellar setae are stout (Fig. 140). In
addition to a long foretarsal tooth the males have a long tubercle at the apex of the foretibiae
(Fig. 147). Females simply have the inner apex of the foretarsi slightly prolonged. Specimens of
amplus have been studied from Florida, Georgia and south Carolina (in BMNH).
SPECIES INCLUDED
amplus (Hood, 1925a: 221-2) (Adiaphorothrips). Holotype $, U.S.A.: Florida (USNM).
Genera of Macrothripina
This group was erected by Karny (19210) as a subfamily to include seven genera of large thrips
most of which are retained in the group in the present revision. Priesner (1961) did not use this
group name, and he placed the nominate genus Macrothrips in his 'Elaphrothrips-group' of the
Idolothripini. Macrothripina is used here for an apparently monophyletic group of 12 genera
from the Old World tropics plus Diplacothrips from the Neotropics. All of the included species
lack metathoracic sternopleural sutures in contrast to most Pygothripini. The area of greatest
diversity of the subtribe is evidently South East Asia.
Within the Macrothripina two major genus-groups can be distinguished, the Aesthesiothrips-
and Ethirothrips-groups. Aesthesiothrips (Fig. 200), Polytrichothrips (Fig. 195) and Taras-
sothrips (Fig. 196) all have long maxillary stylets which are close together medially, and antennal
segment III shorter than IV (Figs 251-253). Moreover, Dichaetothrips (Fig. 199) and Celi-
dothrips (Fig. 201), which resemble each other, as well as Peltariothrips (Fig. 194) and
Tarassothrips (Fig. 196), in the presence of an ommatidium-like structure on each cheek, also
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 51
have long stylets. Aesthesiothrips, Peltariothrips, Tarassothrips and some Dichaetothrips species
have unusually long antennal sense-cones (Fig. 254). The pelta is unusually variable between
species in this Aesthesiothrips genus-group, being particularly aberrant in Peltariothrips (Fig.
212), although in most Macrothripina it is either Ethirothrips-\ike or Diaphorothrips-like (Figs
228-234). Despite this structure, Peltariothrips appears to be intermediate between the Aesthe-
siothrips group and the Ethirothrips group in having the stylets further apart. Both genus-groups
exhibit a tendency for a tubercle to be developed at the inner apex of the foretibia
(Figs 220-221), this characteristic being variable within some genera.
The Ethirothrips genus-group includes many more species than the Aesthesiothrips group
referred to above. Ethirothrips itself is a large, diverse genus from which Herathrips is a
monobasic derivative with short stylets (Fig. 188). Diaphorothrips is another small derivative
from this large genus in which the four species have stout ocellar setae and a sub-apical tubercle
on the foretibiae. Machatothrips species are very similar to Diaphorothrips species in the form of
the head and pelta, although the forefemora bear a series of tubercles in females usually (Figs
217-219). Ischyrothrips is used for a single species with forelegs like Machatothrips, but with
short ocellar setae and the pelta similar to Ethirothrips species rather than Diaphorothrips
species. Finally, small females of Macrothrips are essentially similar to Diaphorothrips species,
although large males of Macrothrips are subject to allometric growth and develop large tubercles
on the head and thorax (Fig. 206).
AESTHESIOTHRIPS Ananthakrishnan
(Figs 200, 216, 221, 238, 251)
Aesthesiothrips Ananthakrishnan, I96la: 253. Type-species: Aesthesiothrips jatrophae Ananthakrishnan,
by monotypy.
This monobasic genus, from India and Malaya, was redefined by Palmer & Mound (1978). It is
similar to Polytrichothrips in having very long maxillary stylets which are close together in the
middle of the head (Fig. 200), and in having antennal segment III shorter than IV (Fig. 251).
However, the antennal sense cones are unusually long, as in Tarassothrips and some
Dichaetothrips species but unlike Polytrichothrips (Fig. 243). The pelta is roughly triangular
(Fig. 216) but recessed into the anterior margin of tergite II, and the foretibiae in both sexes bear
a small apical tubercle (Fig. 221).
SPECIES INCLUDED
jatrophae Ananthakrishnan, 1961o: 253-4. Holotype <j>, INDIA (TNA).
CELIDOTHRIPS Priesner
(Figs 201, 215, 241)
Celidothrips Priesner, 1951: 361. Type-species: Docessissophothrips adiaphorus Karny, by monotypy.
Ommatidothrips Mound, 1970: 120-2. Type-species: Ommatidothrips lawrencei Mound, by monotypy.
[Synonymised by Mound, 1974: 36.]
This genus was redefined with a key to the four known species by Mound (19740). Since then
several females from Singapore and Kuala Lumpur have been examined which may represent
adiaphorus although the postocellar setae are relatively longer than the holotype and the tube
relatively shorter. A single male from New Guinea also represents this genus and has the
metanotum raised into a flange-like tubercle as in dolichos and lawrencei. Characterisation of
species within the genus is at present unsatisfactory due to patterns of allometric growth and
sexual dimorphism. The species resemble those placed in Dichaetothrips in having an isolated
ommatidium-like structure behind the eye on each cheek (Fig. 201), although the pelta is
different in structure (Fig. 215) and the antennal sense cones short (Fig. 241).
SPECIES INCLUDED
adiaphorus (Karny, 1923: 328-31) (Docessissophothrips). Holotype $, JAVA (SMF).
52 L. A. MOUND AND J. M. PALMER
came/us (Karny, 1920c: 43) (Adiaphorothrips). Lectotype cf , AUSTRALIA (NRS).
do//c/ios(Hood, 19186: 144) (Cryptothrips). Holotype cf, AUSTRALIA (USNM).
/awrence/(Mound, 1970: 122-3) (Ommatidothrips). Holotype $, GUADALCANAL (BMNH).
DIAPHOROTHRIPS Karny
(Figs 205, 207, 208, 214, 220)
Diaphorothrips Karny, 1920a: 186. Type-species: Diaphorothrips unguipes Karny, by monotypy.
Diaphorothrips (Cnemidothrips) Priesner, 1940: 403. Type-species: Diaphorothrips hamipes Karny, by
original designation.
Palmer & Mound (1978) redefined Diaphorothrips with a key to the three Oriental species, but
Sakimura (1979) has described a further species from Fiji. A foretibial tubercle is present in both
sexes arising sub-apically (Fig. 220), whereas only a few species in related genera have such a
tubercle and in these it is apical in position. The unique holotype of kraussihas not been studied,
but the other three species have a pair of pores on the metanotum, an unusual characteristic in
the Ethirothrips-group of genera, although found in E. brevis, and the pelta is triangular with the
lateral corners recurved (Fig. 214). The type-species unguipes has the anteocellar setae longer
than the postocellars although the reverse is true in other members of the genus (Fig. 205). This
is another example of the ineffectiveness of the lengths of the ocellar setae as indicators of
relationship. Variation in length of these setae is discussed under the related genera Ethirothrips
and Dichaetothrips , as well as under Diceratothrips and Neosmerinthothrips in the Diceratothri-
pina.
SPECIES INCLUDED
clavipes Priesner, 1940: 403-5. Holotype $, RIAU Is. (SMF).
hamipes Karny, 1923: 296-9. Syntype $, JAVA (SMF).
*kraussi Sakimura, 1979: 313-5. Holotype $, FIJI (BPBM).
unguipes Karny, 1920a: 186-9. Syntype <j>, SRI LANKA (SMF).
thevetii Ananthakrishnan, 1957: 101-2. Holotype <J>, INDIA (TNA).
spinosus Ananthakrishnan, 1959: 321-2. Holotype $, INDIA (TNA).
DICHAETOTHRIPS Hood
(Figs 199, 202, 225-227, 235, 236, 248, 254-256)
Dichaetothrips Hood, 1914: 164. Type-species: Dichaetothrips brevicollis Hood, by monotypy.
This genus has had a confused nomenclatural history, although the solitary female specimen
from Guyana on which it was based does not appear to have been studied by any subsequent
author. The generic name is derived from a pair of large postocellar setae, as figured in the
original description, and because of these setae brevicollis has been associated with a variety of
other species in which the ocellar setae are more or less developed.
Stannard (1957) treated Dichaetothrips as a subgenus of Diceratothrips, but species of the
latter genus can be distinguished by the presence of well-developed metathoracic sternopleural
sutures. The unique holotype of D. brevicollis lacks these sutures as do all species of Macrothri-
pina. Moreover, no other specimen which the present authors would accept as congeneric with
brevicollis has been studied from the New World, whereas at least two species from South East
Asia appear to be very closely related. Since brevicollis has not been collected again it seems
possible that the original specimen may have been artificially introduced. However, Dipla-
cothrips, with two species from South America, is closely related to Dichaetothrips.
The holotype of brevicollis has a most unusual D-shaped pelta (Fig. 226), and despite its size
and well developed wings the tergal wing-retaining setae are straight (Fig. 235). Moreover,
antennal segments III-IV are unusually slender and slightly clubbed (Fig. 254), and the sense
cones are elongate. Finally, there is an isolated, weakly developed ommatidium on the cheek
just behind each eye. The Asian specimens here related to this species share these characters but
do not have the elongate postocellar setae of brevicollis (Fig. 202). However, the length of the
ocellar setae is variable in Ethirothrips (even within species, e.g. stenomelas), Diaphorothrips
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 53
and Diceratothrips, and also varies between species in Neosmerinthothrips. The lack of elongate
ocellar setae is therefore not accepted here as grounds for excluding these Asian species from
Dichaetothrips and the genus is redefined as follows.
Large dark Pygothripini; head (Figs 199, 202) with cheeks almost parallel and straight, with a few pairs of
stout setae and an isolated ommatidium-like structure behind each compound eye; maxillary stylets
retracted almost to postocular setae, sub-parallel and about one-third of head width apart. Antennae (Fig.
256) 8-segmented, VIII slender and constricted at base; sense cones long and curved, 2 on III, 4 on IV;
segment IV usually longer than III. Pronotum short but wide, anterior margin heavily thickened, anterior
setae short; epimeral sutures complete; praepectus present, probasisternal plates large; mesopraesternum
boat-shaped. Mesonotal lateral setae and metanotal median setae small. Metathoracic sternopleural
sutures not developed. Foretarsal tooth present in both sexes. Wings, when present, broad with many
duplicated cilia, sub-basal setae B2 and 53 elongate. Pelta occupying scarcely one-third of anterior margin
of tergite II , without lateral lobes (Figs 225-227) . Tergites II- VI with one pair of straight or weakly sigmoid
wing-retaining setae (Figs 235-236); tube variable, sometimes exceptionally broad and heavy.
SPECIES INCLUDED
brevicollisHood, 1914: 164-5. Holotype $, GUYANA (USNM).
okajimaisp. n. Holotype $, SINGAPORE (OCT).
secutorsp. n. Holotype 9 , THAILAND (OCT).
Dichaetothrips okajimaisp. n.
(Figs 202, 227, 236, 256)
Macropterous $. Colour dark brown, tube darkest, tarsi paler; median area of antennal segment HI, also
pedicels of IV-V, yellowish; forewing with apical fifth pale but extensively shaded along median margins
and with one longitudinal dark line in basal two-thirds; major setae shaded (specimens all cleared).
With the characters in the generic definition. Head slightly constricted behind eyes (Fig. 202),
ommatidium-like structure reduced or absent; postocular setae much longer than pronotal setae; postocel-
lar setae small. Foretibia with a small but stout apical tubercle; foretarsal tooth long and curved. Pelta
reticulate distally, apex constricted. Sternites IV- VI (VII in large individuals) with paired transverse areas
of specialised sculpture (Fig. 236).
Measurements (holotype $ in /mi). Body length 5500. Head, length 600; maximum width 340;
postocular setae 240. Pronotum, length 250; width 540; major setae - am 40; aa 40; ml 40; epim 90; pa 130.
Forewing, length 2200; distal width 200; sub-basal setae 30, 90, 150; number of duplicated cilia 36/39.
Tergite IX setae, Bl 600; B2 550; B3 550. Tube, length 800; basal width 200; terminal setae 200. Antennal
segments I- VIII length, 120; 100; 180; 200; 200; 150; 115; 115.
Macropterous c?. Very similar to $ in colour and structure; sculptured areas on sternites IV- VII almost
continuous medially.
Measurements (paratype d" in /tm). Body length 3200. Head, length 570; postocular setae 240.
Pronotum, length 270; width 570; major setae - epim 150; pa 180. Tube length 600.
SPECIMENS STUDIED
Holotype $, Singapore: Macritchie Park, on dead branches, 22.vii.1976 (S. Okajimd) (OCT).
Paratypes. 2 <j>, 1 d" collected with holotype (OCT; 1 $ BMNH).
COMMENTS. This species differs from the other two members of the genus in the slight
constriction of the head behind the eyes, and in the presence of sternal reticulate areas.
Moreover the pronotal midlateral setae are shorter and the median antennal segments paler
than in the other species. The tube is longer than the head in the type-series, but this is probably
subject to allometric growth and may not be true of smaller specimens. Unlike brevicollis the
foretibiae bear a stout apical tubercle.
Dichaetothrips secutorsp. n.
(Figs 199, 225, 255)
Macropterous $ . Colour dark brown, extreme base of antennal segment III and all tarsi paler; tube black;
major setae weakly shaded; forewing largely pale with two longitudinal dark lines (specimens all cleared).
54 L. A. MOUND AND J. M. PALMER
Head similar to brevicollis but with postocellar setae short and stout (Fig. 199); one pair of ommatidia-
like structures weakly developed on cheek just behind compound eyes. Antennal segment III asymmetri-
cal (Fig. 255), otherwise similar to okajimai. Foretibiae with a small apical tubercle. Pelta (Fig. 225) and
anapleural suture as in brevicollis; wing- retaining setae on tergites IV-VI straight and not directed mesad.
Tube very heavy and tapering. Sternites without reticulate areas.
Measurements (holotype $ in /*m). Body length 5400. Head, length 560; width 340; postocular setae
320. Pronotum, length 230; width 550; major setae - am 15; aa 75; ml 220; epim 320; pa 130. Mesonotal
lateral setae 6. Metanotal median setae 65. Forewing, length 2300; distal width 240; sub-basal setae 60, 180,
180; number of duplicated cilia 50. Tergite IX setae Bl 500; B2 450. Tube, length 570; basal width 250,
terminal setae 200. Antennal segments III- VIII length, 175; 195; 185; 135; 95; 80.
SPECIMENS STUDIED
Holotype <J>, Thailand: North, Doi suthep, in dead leaves, 800 m, 7.viii.l976 (5. Okajimd) (OCT).
Specimens excluded from type-series. Thailand: East, Chanta Buri, 1 £ mic. in dead leaves, 30.iii.1975
(S. Yamaguchi). Laos: Vang-Viong, 1 $ mac. in dead leaves, 21.iii.1975 (S. Yamaguchi). West Malaysia:
Tanah Rata, 1 9 mic., 5. hi. 1976 (W. Suzuki). Japan: Amami-ohshima, Hatsuno, 1 $ mic. in dead leaves,
4.vii.l972 (5. Okajimd) (OCT).
COMMENTS. The four specimens listed above are excluded from the type-series for the following
reasons. The macropterous female from Laos is slightly smaller than the holotype, with only one
dark longitudinal line on the forewing, and with the ommatidium-like structure on the cheeks
more evident; this is almost certainly conspecific with the secutor holotype. The micropterous
female from Japan has larger setae (po 400 /xm) but is also probably the same species. However,
the micropterous female from Malaya has the tube rather shorter (500 nm) with the margins
very slightly convex. This specimen has longer wing remnants and larger ocelli than the
specimen from Japan. Finally the micropterous female from eastern Thailand is almost apterous
and has the tube remarkably broad (length 390 /-im; width 260) with the lateral tubercles
distinctly more emergent. If this material all represents secutor then the range of variation,
particularly of the tube, is remarkable. From brevicollis this species can be recognised by the
tubercle on the foretibiae, and from okajimai by the head shape and antennal colour as well as
the lack of sternal reticulate sculpture.
DIPLACOTHRIPS Hood gen. rev.
(Figs 197, 222, 239, 245)
Diplacothrips Hood, 1937c: 506-7. Type-species: Diplacothrips borgmeieri Hood, by monotypy.
This genus was treated by Stannard (1957) as a synonym of Dichaetothrips which was itself
placed as a subgenus of Diceratothrips . In the course of the present studies Diceratothrips has
been recognised as a genus of Neotropical species and placed in the Diceratothripina, all species
of which have well developed metathoracic sternopleural sutures. Diplacothrips is here treated
as a valid genus, very similar to Dichaetothrips but distinguished by short antennal sensoria
(Fig. 245) and long preocellar setae (Fig. 197). Both species in the genus have the tube broadly
conical (Fig. 239), both of them have a D-shaped pelta as in Dichaetothrips (Fig. 222), but
neither of them have sternal reticulate areas. The generic definition of Dichaetothrips (q.v.)
applies to Diplacothrips with the exceptions noted above. This is the only genus of Macrothripi-
na endemic to the New World.
SPECIES INCLUDED
borgmeieri Hood, 1937c: 507-9. Holotype $, PERU (USNM).
piceusHood, 1952c: 161-2. Holotype $, BRAZIL (USNM).
ETHIROTHRIPS Karny
(Figs 187, 189-193, 228-234, 244, 249, 250)
Liothrips (Ethirothrips) Karny, 1925: 133. Type-species: Liothrips thomasseti Bagnall (here regarded as a
synonym of Phlaeothrips stenomelas Walker, 1859), by subsequent designation, Priesner, 1949: 129.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 55
Scotothrips Priesner, 1939a: 75. Type-species: Adiaphorothrips elephas Karny, by original designation.
Syn. n.
Paracryptothrips Moulton, 1944: 281. Type-species: Par aery ptothrips inermis Moulton, by original de-
signation. Syn. n.
Percipiothrips Ananthakrishnan, 1964a: 72. Type-species: Mesothrips indicus Bagnall, by monotypy.
Syn. n.
Elaphridia Ananthakrishnan, 1964a: 90. Type-species: Elaphrothrips agasthya Ramakrishnan, by
monotypy. Syn. n.
Eurynotothrips Moulton, 1968: 119. Type-species: Eurynotothrips latapennis Moulton, by monotypy.
Syn. n.
Uredothrips Ananthakrishnan, 1969a: 184-5. Type-species: Uredothrips indicus Ananthakrishnan, by
monotypy. Syn. n.
Decothrips Ananthakrishnan, 1969a: 182. Type-species: Decothrips anacardii Ananthakrishnan, by
monotypy. Syn. n.
Mound (19740: 45, 92) has indicated some of the problems of relationships between the species
of Ethirothrips, although at that time they were treated under the names Dichaetothrips and
Scotothrips. Unfortunately for nomenclatural stability, Dichaetothrips is now recognised as a
small genus of highly aberrant large species with the pelta remarkably small and D-shaped
(Figs 225-227) and the wing-retaining setae reduced (Fig. 235). The next available generic name
is Ethirothrips, and the type-species of both this and Scotothrips are very similar. Mound (1974a)
chose to distinguish between these two on the basis of the length of the postocellar setae,
although it was indicated that these setae are variable in length. Re-examination of almost all the
described species in this group, together with several undescribed species, has convinced the
authors that there is at present no way in which genera, or even satisfactory species-groups, can
be defined in this complex. As a result Ethirothrips is used here for a wide range of species in
which the postocellar setae are usually short but sometimes long, the cheeks usually bear a few
setae but these may be very reduced or as numerous and strong as in Machatothrips species, the
cheeks are usually straight but vary from convex or even sinuate to being sharply constricted
basally, and the pelta usually has weak lateral lobes but is sometimes broad and entire. A
foretarsal tooth is present in all males, but is absent in females of a few species; moreover in
some species the foretibiae bear a small apical tubercle in one or both sexes. Thus, although
apparently monophyletic, Ethirothrips is a diverse complex of species comprising a major
element of the Old World Pygothripini. Generic relationships are discussed under Macro-
thripina.
Within Ethirothrips as treated here, the only species-group which is partially distinct is that
represented by the name Uredothrips. This group includes indicus, brevisetosus and tirumalaien-
sis from India, together with tibialis from Japan and Malaya. These species have the eyes
reduced in size and the maxillary stylets deeply retracted, but about one-third of the head width
apart (Fig. 189). However, these characters do not distinguish the species sharply from
vitreipennis from Africa (and India?), and vitreipennis cannot be distinguished at more than
species level from elephas. Moreover, although tibialis and an undescribed species from Japan
have a foretibial tubercle, this is not true of the three Indian species in Uredothrips, and although
the tube is short and heavy in indicus but long in brevisetosus, it is intermediate in tibialis and
tirumalaiensis. All these species have short ocellar setae.
Decothrips is available for a single species of the Ethirothrips group in which the head is deep
and narrowed basally as in brevis (Fig. 193) but in which the metanotal pores are absent, the
pelta broad and the antennae unusually slender (Fig. 244). This species seems to be at one end of
the range of variation of Ethirothrips.
Elaphridia has been used for a single species agasthya (Figs 187, 228) and cannot be
distinguished satisfactorily from other members of Ethirothrips. Contrary to Ananthakrishnan
(19730: 282) agasthya is not only unrelated to crassiceps Bagnall (now placed in Dinothrips), but
neither of them are closely related to Elaphrothrips.
Percipiothrips was erected for a single species now in the Ethirothrips-group but at that time
regarded as belonging in Phlaeothripinae. As Mound (1968: 82) has pointed out, the postocellar
56 L. A. MOUND AND J. M. PALMER
setae of the indicus syntypes range in size from 35 to 60 /urn, and this species is very similar to
stenomelas (Fig. 190), apart from the pale antennal III. Although indicus Ananthakrishnan now
stands as a secondary homonym of indicus Bagnall, a new name is not proposed here pending
further studies at species level into potential synonyms.
Scotothrips was used by Mound (19740) for a group of species mainly from Australia with
stout cheek setae (Figs 191, 192). Within this group latapennis, the type-species of Euryno-
tothrips, is particularly large, and australiensis has a particularly short, broad head (Fig. 192).
However, the majority of species in this group cannot be distinguished readily from the Oriental
members of Ethirothrips , although a foretibial tubercle is found in several Australian but in no
Oriental species. The type-species elephas can be distinguished from stenomelas mainly by the
broader, entire pelta which lacks lateral wings.
Par aery ptothrips was erected for two species, of which the type-species has a complete suture
between antennal segments VII and VIII, although these segments are broadly joined. Apart
from this character, and the yellow legs, inermis is very similar to several Australian species
previously treated in Scotothrips. The second species (ftjiensis) which was described in Paracryp-
tothrips has 7-segmented antennae with no suture between VII and VIII, but although the
female holotype lacks a foretarsal tooth a male specimen (in BMNH) apparently representing
this species has a slender tooth almost as long as the foretarsal width.
The type-species of Ethirothrips was treated as a synonym of brevicornis by Mound (19740:
46). However, the unique holotype of Phlaeothrips stenomelas Walker (Figs 190, 234, 250) from
Sri Lanka was discovered recently in the BMNH, mounted dry on a card. This specimen, in
excellent condition, is now mounted in balsam on a microscope slide and is evidently the same
species as brevicornis and thomasseti. This species is widespread in the Old World tropics and
material has now been studied from Seychelles, Rodrigues, Mauritius, Madagascar, Sri Lanka,
Singapore, Malaya, Thailand, Java, Philippines, Ryukyus, Bonins, Fiji, Samoa, Hawaii, New
Guinea, Solomon Is. and Australia. Moreover, a female specimen in the Zoologisches Museum,
Berlin, labelled 'Liothrips gigas' and 'New Britannien/Ralum/F. Dahl S' is here accepted as the
holotype of gigas Karny and this is synonymised with stenomelas.
Another widespread species of which the name is changed in this paper is claripennis, here
accepted as a junior synonym of brevis. Mound (1968) observed that the tube of the damaged
unique holotype of brevis is heavily sculptured, but this is now interpreted as being an artefact
due to excessive bleaching of this very crushed specimen. The form of the pelta and the presence
of a pair of pores on the metanotum distinguish this species (Mound, 19746: 177).
Chen (1980) also refers to a sculptured tube when describing a new species, virgulae, from
Taiwan. Examination of a paratype of this species has revealed that this effect is also due to
excessive bleaching and crushing in a water-soluble mountant. This sculpture, although an
integral part of the structure of the tube, is not normally visible except in heavily bleached
specimens. Mounting techniques of this sort are quite unsuitable for serious taxonomic work.
The original illustration by Chen of virgulae indicates clearly the damaged condition of the
paratype studied; the head is flattened and swollen medially and the pronotum distorted.
Fortunately virgulae appears to be a common species extending from Taiwan along the Ryuku
chain of islands.
Several names from India are now placed in synonymy under obscurus, and to these are here
added fungivorus from the Congo and neivei from Cuba. This species has long postocellar setae
and a small pelta with lateral wings (Fig. 233). In contrast, uredinis, which looks similar at first
sight, has a broad pelta without lateral wings, similar to that offirmus (Fig. 229) and more typical
of species previously referred to Scotothrips. Although indicus Bagnall is similar in general
appearance to obscurus, it is unusual in having long antennal sense cones as in Dichaetothrips
(Fig. 249).
SPECIES INCLUDED
acanthus (Hood, 19196: 88-90) (Cry ptothrips). Holotype $, AUSTRALIA (USNM). Comb. n.
sjostedti Karny, 1920c: 42 (Cry ptothrips). Holotype d", AUSTRALIA (NRS).
sismondini Girault, 1926: 1 (Adiaphorothrips). Holotype $, AUSTRALIA (QMB).
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 57
nox Girault, 1928«: 2 (Adiaphorothrips} . Holotype 9, AUSTRALIA (QMB).
differens Priesner, 19286: 656-7 (Adiaphorothrips) . Holotype $, AUSTRALIA (SMF).
advenfor(Bianchi, 1945: 259-60) (Dichaetothrips). Holotype 9, NEW CALEDONIA (BPBM). Comb. n.
agasf/iya(Ramakrishna, 1934: 10-12) (Elaphrothrips). Syntypes $, INDIA (TNA). Comb. n.
a/iacardii( Ananthakrishnan, 1969a: 182-4) (Decothrips). Syntypes cf 9, INDIA (TNA). Comb. n.
ausfra/iens/s (Moulton, 1968: 95-6) (Gastrothrips). Holotype $, AUSTRALIA (CAS). Comb. n.
barretti (Mound, 1974a: 94-7) (Scotothrips) . Holotype $, AUSTRALIA (ANIC). Comb. n.
beesoni (Moulton, 1928e: 5-6) (Dichaetothrips}. Holotype 9, INDIA (CAS). Comb. n.
brevis (Bagnall, 19210: 276-7) (Adiaphorothrips}. Holotype cf , SEYCHELLES (BMNH). Comb. n.
claripennis Moulton, 1934: 503 (Dichaetothrips}. Holotype $, HAWAII (CAS). Syn. n.
trinidadensis Hood, 19356: 168-70 (Gastrothrips}. Holotype $, TRINIDAD (USNM). Syn. n.
indicus Ananthakrishnan, 1968c: 967-9 (Nesothrips). Syntype cf 9, INDIA (TNA). Syn. n.
diversus Ananthakrishnan, 19726: 434-6 (Nesothrips). Holotype $, INDIA (TNA). Syn. n.
brevisetosus (Ananthakrishnan & Jagadish, 1970: 266-8) (Diceratothrips) . Holotype 9> INDIA (TNA).
Comb. n.
distasmus (Mound, 1974a: 97-8) (Scotothrips). Holotype 9, AUSTRALIA (ANIC). Comb. n.
dracon (Karny, 1920c: 43) (Adiaphorothrips} . Holotype cf , AUSTRALIA (NRS). Comb. n.
elephas (Karny, 1920c: 43) (Adiaphorothrips). Lectotype 9, AUSTRALIA (NRS). Comb. n.
fijiensis (Moulton, 1944: 282-3) (Paracryptothrips). Holotype 9, FIJI (BPBM). Comb. n.
*firmiis(Hood, 1952c: 162) (Gastrothrips}. Lectotype 9, BRAZIL (USNM). Comb. n.
g/rau/f/(Hood, 19186: 148-9) (Adiaphorothrips). Holotype cf , AUSTRALIA (USNM). Comb. n.
hibisd (Moulton & Steinweden, 1933: 32-3) (Neoheegeria). Holotype 9, SOCIETY Is. (BPBM). Comb. n.
longus Moulton, 1944: 297-300 (Neoheegeria). Holotype 9, FIJI (BPBM).
latus fijiensis Moulton, 1944: 270-1 (Cryptothrips). Holotype cf , FIJI (BPBM).
indicus ( Ananthakrishnan, 1969a: 185) (Uredothrips}. Syntype 9 Cf , INDIA (TNA). Comb. n.
indicus (B agnail, 19216: 365-6) (Mesothrips). Lectotype 9, INDIA (BMNH). Comb. n.
inermis (Moulton, 1944: 281-2) (Paracryptothrips). Holotype 9 , FIJI (BPBM). Comb. n.
j'o (Girault, 1926: 1) (Adiaphorothrips). Lectotype cf , AUSTRALIA (QMB). Comb. n.
latapennis (Moulton, 1968: 119-21) (Eurynotothrips). Holotype 9, AUSTRALIA (CAS). Comb. n.
longisetis (Ananthakrishnan & Jagadish, 1970: 268-9) (Diceratothrips). Holotype 9> INDIA (TNA).
Comb. n.
madagascariens/s (Bagnall, 1936: 220-1) (Cryptothrips). Holotype 9> MADAGASCAR (MNHN). Comb. n.
*meridionaIis(Zur Strassen, 1976: 247) (Diceratothrips). Holotype 9, ST HELENA (MRAC). Comb. n.
obscurus(Schmutz, 1913: 1074-6) (Ischyrothrips). Holotype 9, SRI LANKA (NMV). Comb. n.
fcowy/Bagnall, 1924: 639-40 (Mesothrips). Lectotype 9, INDIA (BMNH). Syn. n.
fungivorus Priesner, 19396: 52-4 (Dichaetothrips). Syntypes 9 Cf , CONGO (SMF). Syn. n.
gloveri Ramakrishna & Margabandhu, 1939: 31-2 (Neosmerinthothrips) . Lectotype 9 , INDIA (TNA).
Syn. n.
neivei Hood, 1940a: 576-9 (Dichaetothrips). Holotype 9, CUBA (USNM). Syn. n.
usitatus Ananthakrishnan & Jagadish, 1970: 273-4 (Diceratothrips), replacement name for indicus
Ananthakrishnan, 19616: 270-1 (Dichaetothrips). Holotype 9, INDIA (TNA). Syn. n.
stenomelas (Walker, 1859: 224) (Phlaeothrips) . Holotype 9, SRI LANKA (BMNH). Comb. n.
brevicornis Bagnall, 19106: 696-8 (Diceratothrips). Holotype cf , HAWAII (BMNH). Syn. n.
niger Schmutz, 1913: 1080-4 (Ischyrothrips). Syntypes 9 Cf , SRI LANKA (NMV). Syn. n.
gigas Karny, 19136: 133-4 (Liothrips). Holotype 9, NEW Britain (ZMB). Syn. n.
thomasseti Bagnall, 1921a: 288-9 (Liothrips). Lectotype 9, RODRIGUES (BMNH). Syn. n.
nigricornis Bagnall, 1921a: 278-9 (Liothrips). Lectotype 9 , SEYCHELLES (BMNH). Syn. n.
intrepidus Bagnall, 1921a: 279 (Liothrips). Holotype 9, SEYCHELLES (BMNH). Syn. n.
setidens Moulton, 1928a: 129-30 (Mesothrips}. Holotype 9, HAWAII (BPBM). Syn. n.
niger Moulton & Steinweden, 1935: 165 (Cryptothrips}. Holotype 9 , MARQUESAS Is. (BPBM). Syn. n.
madagascariensis Bagnall, 1936: 222. Holotype 9, MADAGASCAR (MNHN). Syn. n.
sybarite (Mound, 1974a: 100) (Scotothrips). Holotype 9, AUSTRALIA (ANIC). Comb. n.
tftfefe(Okajima, 1975: 16-9) (Uredothrips). Holotype 9, RYUKU Is. (OCT). Comb. n.
tirumalaiensis (Ananthakrishnan, 19696: 298-9) (Uredothrips). Holotype 9, INDIA (TNA). Comb. n.
uredinis (Ananthakrishnan & Jagadish, 1970: 269-71) (Diceratothrips). Holotype 9' INDIA (TNA).
Comb. n.
virgu/ae(Chen, 1980: 180-1) (Scotothrips). Holotype 9, TAIWAN (BCIQ). Comb. n.
vitreipennis (Priesner, 19396: 54-5) (Scotothrips). Syntypes 9, CONGO (SMF). Comb. n.
58 L. A. MOUND AND J. M. PALMER
SPECIES REMOVED TO PHLAEOTHRIPINAE
Teuchothrips burroughs! (Girault) (Mound, I974a: 45) (Dichaetothrips).
Akainothrips pallicornis (Karny) (Palmer & Mound, 1978: 186) (Adiaphorothrips) .
HERA THRIPS Mound
(Fig. 188)
Herathrips Mound, 1914a: 54. Type-species: Adiaphorothrips nativus Girault, by monotypy.
This genus was erected for a single large Australian species with a broad pronotum as in
Macrothrips, but a small head with the stylets not deeply retracted (Fig. 188).
SPECIES INCLUDED
nativus (Girault, 1928c: 2) (Adiaphorothrips). Lectotype d", AUSTRALIA (QMB).
ISCHYROTHRIPS Schmutz
(Fig. 213)
Ischyrothrips Schmutz, 1913: 1074. Type-species: Ischyrothrips crassus Schmutz, by subsequent designa-
tion, Priesner, 1949: 134.
Of the four species described by Schmutz in this genus obscurus is placed in Ethirothrips,
spinosus in Dinothrips and niger is a synonym of Ethirothrips stenomelas. The unique female
holotype of crassus has very broad forefemora bearing 7 to 10 small tubercles on the inner
margin and the foretibia is ridged on the inner surface. Although closely related to Macha-
tothrips this genus can be distinguished by the absence of a pair of long ocellar setae, and the
form of the pelta (Fig. 213, cf. Fig. 210).
SPECIES INCLUDED
crassus Schmutz, 1913: 1076-8. Holotype $, SRI LANKA (NMV).
MACHATOTHRIPS Bagnall
(Figs 203, 204, 210, 217-219, 237, 246, 247)
Machatothrips Bagnall, 19086: 189. Type-species: Machatothrips biuncinatus Bagnall, by monotypy.
Adiaphorothrips Bagnall, 1909c: 536. Type-species: Adiaphorothrips simplex Bagnall, by original designa-
tion. [Synonymised by Priesner 1939: 75.]
Cnestrothrips Priesner, 1932: 344; 1939: 75. Type-species: Cnestrothrips dammermani Priesner, by original
designation. [Synonymised by Palmer & Mound 1978.]
This genus was revised recently by Palmer & Mound (1978) with keys to 14 species. However,
species recognition in the genus is exceptionally difficult and recently collected material in the
collections of Dr Shuji Okajima indicates that it is still not possible to distinguish satisfactorily
between intraspecific and interspecific variation in Machatothrips. In most species of this genus
females rather than males bear the obvious sexually dimorphic characteristics; however, Palmer
& Mound described two species from Malaya in which males share the female sex-linked
character of a row of teeth on the fore femora. Machatothrips is closely related to Diaphor-
othrips and Macrothrips.
SPECIES INCLUDED
antennatus (Bagnall, 19156: 594) (Adiaphorothrips). Lectotype 9, BORNEO (BMNH).
dammermani Priesner, 1932: 344 (Cnestrothrips). Holotype $, RIAU Is. (SMF).
arfocarp/Moulton, 1928c: 322. Holotype $, TAIWAN (CAS).
biuncinatus Bagnall, 19086: 189. Holotype $, NEW GUINEA (BMNH).
simplex Bagnall, 1909c: 537 (Adiaphorothrips). Lectotype d", BORNEO (BMNH).
montanus Priesner, 1932: 344. Holotype $, BORNEO: Sarawak (SMF).
braueri Karny, 1912a: 23. Holotype $, CAMEROUN (ZMB).
braueri f. karnyi Priesner, 1932: 340. Holotype $, CONGO (? lost).
multidens Bagnall, 1934a: 487. Lectotype $, GHANA (BMHN).
paucidens Bagnall, 1934a: 489. Lectotype $, GHANA (BMNH).
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 59
pauddens var. bicolorisetosus Bagnall, 1934a: 489. Lectotype $, SIERRA LEONE (BMNH).
ce/05/aMoulton, 1928c: 325. Holotype $, TAIWAN (CAS).
corticosus Ananthakrishnan, 1972c: 443. Holotype $, INDIA (TNA).
decorus Palmer & Mound, 1978: 193. Holotype $, WEST MALAYSIA (BMNH).
*d/a507us(Priesner, 1928c: 56) (Adiaphorothrips). Holotype $, EAST AFRICA (lost).
haplodon Karny, 1925: 141. Holotype $, UGANDA (BMNH).
brauerivar. buffaiKarny, 1925: 142. Holotype d", UGANDA (BMNH).
simpliddens Bagnall, 1934a: 490. Holotype $, CONGO (MNHN).
heveae Karny, 1921c: 61. Holotype $, JAVA (SMF).
imlicus Ananthakrishnan & Jagadish, 1970: 279. Holotype $, INDIA (TNA).
lentus Palmer & Mound, 1978: 194-5. Holotype $ , WEST MALAYSIA (BMNH).
quadrudentatusMoulton, 1941 'a: 179. Holotype $, NEW GUINEA (CAS).
silvaticus Ananthakrishnan, 19726: 436. Holotype £, INDIA (TNA).
MACROTHRIPS Bagnall
(Figs 206, 209, 240)
Macrothrips Bagnall, 19080: 359. Type-species: Macrothrips papuensis Bagnall, by original designation.
As discussed by Palmer & Mound (1978) this genus is very close to Machatothrips . All the
specimens which have been studied are here interpreted as representing a single variable
species. The males vary in size very considerably, the largest being bigger than any other
Thysanoptera. However, the available males do not vary greatly in structure; large and smaller
individuals all have recurved tubercles on the forecoxae, the postero-median pronotal margin
prolonged backwards, the antero-lateral pronotal margins slightly emarginate, and a pair of
short stout setae on tubercles behind the eyes (Fig. 206). Females vary much less in size, but are
structurally more variable. The forecoxae have at most a short straight tubercle and the
forefemoral tubercle is absent in small individuals. In contrast to males, however, large females
have a prominent pronotal epimeral tubercle, although this is not present in small females. The
foretibial tubercles also seem remarkably variable; the 'typical' condition involves two short,
acute tubercles, one apical, the other subapical. However, the following variants have been
studied: 2 $ lacking subapical tubercles; 1 $ lacking subapical but with apical also reduced; 1 $
lacking apical tubercle but with subapical well developed.
SPECIES INCLUDED
papuensis Bagnall, 1908a: 359-60. Holotype cf , PAPUA NEW GUINEA (BMNH).
dubius Bagnall, 19080: 361. Holotype 9, PAPUA NEW GUINEA (BMNH). Syn. n.
intermedius Bagnall, 19086: 187-9. Holotype ? cf , PAPUA NEW GUINEA (? lost).
PELTARIOTHRIPSgen. n.
(Figs 194, 198, 212, 242)
Type-species: Peltariothrips insolitus sp. n.
Dark brown species of Macrothripina. Head slightly wider than long, narrowed to base; cheeks with 5-6
spine-like setae, also an ommatidium-like structure ventro-laterally in posterior third (Fig. 194). Two pairs
of postocular setae (1 short, 1 long); postocellar and anteocellar setae short; maxillary stylets retracted to
compound eyes, about one-third of head width apart. Antennae 7-segmented, suture between VII-VIII
incomplete; III shorter than IV; sense cones long and slender, two on III, four on IV (Fig. 242). Pronotum
short, epimeral sutures complete; epimeral setae long; praepectus small. Foretarsal tooth present in both
sexes. Metanotal median setae slender; metathoracic sternopleural sutures absent. Forewings with 10
duplicated cilia. Pelta triangular but with posterior margin concave, anterior margin of tergite II
protruding into pelta (Fig. 212) ; tergites II-VII with one pair of wing- retaining setae; tergite IX setae about
0-8 times as long as tube. Tube about as long as head, sides straight and tapering. Sternites with a transverse
row of discal setae; median sternites of cf with a pair of reticulate (? glandular) areas laterally.
The type-species of this new genus is unusual in the Macrothripina in having antennal segments
VH-VIIIfused. Segment III isshorterthan IV, with thesense cones long andslender (Fig. 242), but
60 L. A. MOUND AND J. M. PALMER
entire family Phlaeothripidae) (Fig. 212), although this structure is also unusual in the related
genera Dichaetothrips , Polytrichothrips and one species of Tarassothrips . Moreover, a single
specimen of a Machatothrips has been studied (in BMNH) from Singapore, which is generally
similar to M. antennatus but has the pelta intermediate in structure between Machatothrips and
Peltariothrips .
Peltariothripsinsolitussp. n.
Macropterous $. Colour dark brown, pterothorax and pelta paler, tube black with apex pale; antennal
segment I whitish yellow, II-IV brownish yellow, V-VI brown distally; femora pale in distal half,
foretibiae and all tarsi brownish yellow; major setae dark; forewings shaded, slightly paler medially but
with a longitudinal dark line.
With the structural characters indicated in generic definition; head and median area of metanotum
without sculpture; sub-basal wing setae short.
Measurements (holotype $ in /x,m). Body length 2515. Head, length 276, maximum width 310; basal
width 240; postocular seta - inner 6/20, outer 38/52. Pronotum, length 150; width 348; major setae - am
26/40, aa 22, ml 28/38, epim 174, pa 26, pm 18. Metanotal median setae 46/56. Forewing, length 1055;
median width 104; sub-basal setae 15, 40, 40; number of duplicated cilia 9/12. Tergite IX setae, B\ 210; B2
200; B3 200. Tube, length 270; basal width 124; terminal setae 120. Antennal segments III-VII length, 80;
96; 82, 65; 90.
Macropterous d". Similar to $ in colour and structure; mesothoracic spiracle slightly enlarged and
toothed in profile; sternites III-V with a pair of reticulate areas anterolaterally.
Measurements (paratype c? in /urn). Body length 2050. Head, length 234; maximum width 252; width at
base 195; postocular setae - inner 12, outer 40. Pronotum, length 132; width 290; epimeral setae 60/88.
Forewing, length 820; number of duplicated cilia 4. Tergite IX setae, fli 195; B2 210: B3 210. Tube, length
200; basal width 98. Antennal segments III-VII length, 70, 75, 85, 50, 70.
SPECIMENS STUDIED
Holotype $, Singapore: Singapore City, on dead twigs, 15. i. 1979 (L. A. Mound) (BMNH).
Paratypes. Singapore: 1 $ , same data as holotype (BMNH); Macritchie Park, 2 $ , 1 cf on dead Areca,
22.vii.l976(S. Okajima) (OCT).
Specimens excluded from type-series. Philippines: Mindanao, Agko, Mt Apo, 57 $, 3 cf on Alpinia
sheath, 2 $ on Palmae leaves, viii.1979; Luzon, Quezon National Forest Park, 9 $ on dead Palmae leaves,
vii.1979 (S. Okajima) (OCT).
COMMENTS. The sternal reticulate areas on the male of this species are similar in appearance to
those found in Dichaetothrips and Tarassothrips. However, the production of the mesothoracic
spiracle into a dentate structure is not found in any other genus of Macrothripina. Through the
courtesy of Dr Shuji Okajima, several series of specimens have been studied from the
Philippines which are very similar to the types of insolitus. The specimens from Mindanao are
much larger than the types from Singapore and by themselves would certainly be regarded as a
distinct species. However, the specimens from Luzon are intermediate in size and structure.
Most of the Mindanao individuals were collected from Alpinia (Zingiberaceae), and the Luzon
individuals from Palmae. Since the only two individuals from Palmae on Mindanao are
themselves intermediate in size between the two main series (Fig. 198), the pattern of variation
might reflect the existence of two or more host-limited or locality-limited species. The
alternative interpretation is adopted here that only one, widely distributed and variable species
is involved. One male from Alpinia on Mindanao is micropterous with the mesothoracic
spiracular processes greatly enlarged.
POL YTRICHOTHRIPS Priesner
(Figs 195, 21 1,243)
Polytrichothrips Priesner, 1939a: 77. Type-species: Polytrichothrips pilosus Priesner (here regarded as a
synonym of Docessissophothrips laticeps Bagnall), by monotypy.
The only species in this genus lacks metathoracic sterno-pleural sutures, but unlike most
Macrothripina, it has elongate maxillary stylets (Fig. 195). As in Aesthesiothrips , these stylets
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 61
are very long, retracted to the eyes, and lie close together medially in the head; however, in
contrast to that genus the antennae do not have greatly elongate sense cones (Fig. 243). The
pelta, which is almost devoid of sculpture as is the metanotum, bears a pair of pores in the
holotype of pilosus although these are not present in the holotype of laticeps (Fig. 211). The
genus is known only from these two individuals which are here regarded as representing one
species.
SPECIES INCLUDED
/af/ceps(Bagnall, 1915a: 322-3) (Docessissophothrips). Holotype $ , BORNEO: Sarawak (BMNH). Comb. n.
pilosus Priesner, 1939a: 77-8. Holotype $, BORNEO: Sarawak (SMF). Syn. n.
TARASSOTHRIPSgen. n.
(Figs 196, 223, 224, 252, 253)
Type-species: Tarassothrips akritussp. n.
Large, blackish brown species of Macrothripina. Head about 1-5 times as long as wide; cheeks with short
spine-like setae, also, in anterior third, a pair of ommatidia-like structures; post ocular setae long, ocellar
setae small; maxillary stylets deeply retracted and close together medially (Fig. 196). Antennae 8-
segmented, III shorter than IV; sense cones long and slender, two on III, four on IV (Figs 252, 253).
Pronotum short, epimeral sutures complete, posteroangular and epimeral setae long; praepectus present.
Both sexes with forefemora enlarged, inner margin often rugose; fore tibiae with inner apical tooth;
foretarsal tooth well developed. Metanotal median setae small and slender. Forewings with two longitu-
dinal dark bands and about 70 duplicated cilia. Pelta recessed into tergite II, either similar to Diaphor-
othrips and Aesthesiothrips or broadly rounded (Figs 223, 224). Tergites II- VI each with one pair of
wing-retaining setae; setae on IX about 0-8 times as long as tube. Tube about 1-2 times as long as head,
slightly constricted at apex. Sternites with transverse row of discal setae; median sternites of both sexes
(usually) with paired reticulate areas anterolaterally.
This new genus is closely related to Aesthesiothrips but has a shorter head without long ocellar
setae. It resembles Celidothrips, Peltariothrips and Dichaetothrips in having an ommatidium-
like structure on each cheek, moreover some species of the last genus have similar reticulate
areas on the sternites. Two species of Tarassothrips are known and these differ from each other
in the form of the pelta (Figs 223, 224).
Tarassothrips akritus sp. n.
(Figs 196, 224, 253)
Macropterous $. Colour dark brown, antennal segment III and apex of tube paler. With the structural
characters given in the generic definition; antennal III shorter than IV, major sense cones on IV about as
long as that segment (Fig. 253), but three minor sense cones also present on dorsal surface. Head with one
pair of anteocellar setae; post ocellar setae small (Fig. 196). Pelta triangular with narrow lateral lobes,
recessed into anterior margin of tergite II (Fig. 224).
Measurements (holotype 9 m /*m). Body length 3900. Head, length 470; width 325; post ocular setae
96/114. Pronotum, length 200; width 438; major setae, am 46/50; aa 45/50; ml 58/90; epim 162/170; pa
176/185. Metanotal median setae 64. Forewing, length 1700; median width 136; sub-basal setae Bv 65; B2
145; B3 228. Tergite IX setae, B^ 460; B2 488. Tube, length 556; basal width 142. Antennal segments
III-VIII length, 124; 200; 194; 120; 95; 80.
Macropterous cf . Colour and structure similar to $ .
Measurements (paratype cf in /urn). Body length 3350. Head length 438; width 276; postocular setae
110. Pronotum, length 180; width 380; major-setae, epim 148/168; pa 142. Tergite IX setae, fij 396; B2
420/430. Tube length 430; basal width 128. Antennal segments III-VIII length, 115; 156; 148; 100; 76; 68.
SPECIMENS STUDIED Holotype cf, Malaya: Buklanyan, on dead branches, 26.xii.1971 (Floyd Andre)
(BMNH).
Paratypes. Malaya: 1 9 , same data as holotype; Kuala Lumpur, 2 cf on pods of 'Singapore', 27.xii.1969
(R. G. & F. Andre). Singapore: Macritchie Park, on dead twigs, 1 cf , ll.viii.1980, 1 £, 19.viii.1980 (L. A.
Mound) (BMNH).
62 L. A. MOUND AND J. M. PALMER
COMMENTS. This new species is remarkable amongst Idolothripinae for the extreme length of the
antennal sense cones. In the BMNH collections there is a single female on a slide without data
which was acquired with the Andre Collection (Mound, 1974c) together with other material
from Malaysia. Because of the lack of data this specimen is not formally named here, but it
differs from akritus in its large size (body length 5800 /am), larger antennal segment III (III
266 /am: IV 286 /am), shorter sense cones (Fig. 252), absence of small dorsal sense cones on
segment IV, broadly oval pelta (Fig. 223), and reduced foretarsal tooth. This specimen also has
paired areas of specialised reticulation on the median sternites; however, the smallest female
paratype of akritus (Macritchie Park) lacks these areas.
Tribe IDOLOTHRIPINI
Priesner (1961) and Jacot-Guillarmod (1978) recognised five subtribes in this group (Table 2).
Apelaunothrips and Dexiothrips (Apelaunothripina) are here treated as Phlaeothripinae (p. 88)
and Atractothripina is synonymised with Hystricothripina (= Zeugmatothripina). The genera
listed by Priesner under the name Megathripina are here treated with Idolothrips in the
Idolothripina. However, most of the genera listed by Priesner in Idolothripina are here treated
in Hystricothripina. Moreover, the Elaphrothrips-group of Priesner is here considered as a
subtribe although Campulothrips, Macrothrips and Sporothrips are re-assigned to the Pygothri-
pini.
As a result, only three subtribes are now recognised, involving 33 genera and 277 species. The
Hystricothripina (predominantly New World) and Idolothripina (predominantly Old World)
are treated as sister-groups, on the grounds that they are the only members of the tribe with the
tube hairy. These two probably constitute the sister-group of the Elaphrothripina, although this
subtribe cannot be defined on any single characteristic. In fact, the Elaphrothripina show
similarities to the Pygothripini in the plesiomorphic form of the anapleural sutures. Moreover,
Anactinothrips has only one pair of sigmoid setae on each tergite as in all pygothripine species.
None of the species in the Idolothripini has metathoracic sternopleural sutures developed,
although these sutures are commonly found in species of Pygothripini with the exception of the
Macrothripina.
Genera of Elaprothripina subtrib. n.
This group is used here in the same sense as that of the 'Elaphrothrips Gruppe' of Priesner
(1961), although it is treated formally as a subtribe for the first time, and is restricted to 10
genera. The generic name Hartwigia q.v. was proposed for a new, monobasic subtribe by
Stannard (1976), but this name was not constructed in accordance with the recommendations of
the Code of Zoological Nomenclature and is not accepted here as being valid. The genus
Hartwigia, however, is here treated within the Elaphrothrips-group for the first time. Species of
Elaphrothripina, in contrast to those of the Idolothripina and Hystricothripina, do not have any
dominant setae laterally on the tube (i.e. tube not hairy), and the anapleural sutures are much
longer and stronger, completely separating the anepisterna from the katepisterna. The praepec-
tal plates are well developed, the maxillary stylets widely spaced, the metathoracic sternopleural
sutures not developed, and there are usually two pairs of wing-retaining setae on each tergite.
The majority of Elaphrothripina species are placed in Elaphrothrips, a widespread tropical
genus which extends into the Nearctic. The species of Anactinothrips, a Neotropical genus, are
unique in the Idolothripini in retaining the plesiomorphic character state of a single pair of
wing-retaining setae on each tergite, and this genus may represent the sister-group of the rest of
the Elaphrothripina. Mecynothrips, from the Austro-Oriental Region, has the remarkable
apomorphic character state of three pairs of wing-retaining setae on each tergite, and may
represent the sister-group of the other Old World genera of this subtribe. Finally, Dinothrips
and Tiarothrips species share the apomorphic character state of the foreocellus just posterior to
the major ocellar setae, in contrast to Elaphrothrips and its derivatives Lamillothrips and
Ophthalmothrips which have the foreocellus arising far forward on the head. Dermothrips and
Malesiathrips are also placed in this subtribe provisionally, although they do not appear to be
closely related to the other genera.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 63
ANACTINOTHRIPS Bagnall
(Figs 263, 295, 311)
Anactinothrips Bagnall, 1909d: 329. Type-species: Anactinothrips meinerti Bagnall, by monotypy.
Ophidothrips Schmutz, 1910: 273. Type-species: Ophidothrips handlirschii Schmutz, by monotypy.
Lophothrips Karny, 1911: 503. Type-species: Lophothrips antennatus Karny, by monotypy.
Both Ophidothrips and Lophothrips were placed as synonyms of Anactinothrips by Moulton
(1933: 416), and this synonymy was retained by Stannard (1957). However, Jacot-Guillarmod
(1978) listed the three genera separately. The syntype female of antennatus has been compared
with the syntype male of meinerti, and these two specimens very probably represent the same
species. In both of them, antennal segments V-VII bear a prominent apical lobe, and the rest of
the body is very similar. These two species, together with borgmeieri and silvicola, have a
relatively long tube, and there is no doubt that Lophothrips should be placed in synonymy with
Anactinothrips. (The data on the antennatus holotype are 'Paraguay, leg. Fiebrig, 27.vii.05'.) In
contrast, the tube of distinguendus , longisetis and vigilans is relatively short, as is that of
handlirschii judging from the original illustration, but there appears to be no reason to use a
separate generic name for these species.
Fifteen species are now placed in Anactinothrips but, since most of these have been collected
only once and there is no knowledge of their structural variation, some synonymy is to be
expected. These species are rather similar in general appearance to Cyphothrips in the
Hystricothripina, with two pairs of stout setae on the vertex (Fig. 263), foretarsal tooth present
in males but absent in females, metanotum with a pair of stout setae, metathorax rather bulbous
laterally, and abdominal tergites with only one pair of wing-retaining setae (Fig. 295). However,
in contrast to most Hystricothripina, the fore wings are relatively broad, the praepectal plates are
well developed, and the tube is relatively short without conspicuous hairs laterally. The
similarities to Hystricothripina are here interpreted as being due to convergence, or more
probably, parallel evolution if Anactinothrips is accepted as the sister-group of the other genera
in Elaphrothripina. The males of Anactinothrips have a sharp angle on the posterior margin of
the forefemora and a series of ridges on the forecoxae which are probably involved in sound
production.
SPECIES INCLUDED
antennatus (Karny, 1911: 503) (Lophothrips). Syntypes cf $, PARAGUAY (ZMB).
borgmeieri Hood, 1950: 34-8. Holotype $, BRAZIL (USNM).
*brachyuraHood, 1941: 227-230. Holotype $, PERU (USNM).
*cristatusHood, 19366: 146-7. Holotype ?, PERU (USNM).
distinguendus Bagnall, 1914c: 379-380. Holotype tf, GUYANA (BMNH).
*fuscus Moulton, 1933a: 418-9. Holotype cf , BRAZIL (CAS).
gibbiferZur Strassen, 1980: 48-53. Holotype $, BRAZIL (SMF).
*graphiduraHood, I938d: 245-7. Holotype $, PERU (USNM).
* handlirschii (Schmutz, 1910: 273-276) (Ophidothrips). Syntypes $ cf , ?BRAZIL (?lost).
longisetis Bagnall, 1926: 556-7. Holotype $, GUYANA (BMNH).
*marginipennisHood, 1941: 223-7. Holotype $, PERU (USNM).
meinerti Bagnall, l9Q9d: 330-2. Syntypes cf Cj, VENEZUELA (BMNH).
*nigricornis Hood, 19366: 143-6. Holotype $ , GUYANA (USNM).
silvicola Hood, 1952c: 167-8. Holotype $, BRAZIL (USNM).
vjgtfa/isHood, 1938a: 241-5. Holotype $, PERU (USNM).
DERMOTHRIPS Bagnall
(Figs 276, 281, 290, 300)
Dermothrips Bagnall, 19106: 677-8. Type-species: Dermothrips hawaiiensis Bagnall, by monotypy.
The relationships of this monobasic genus are far from clear. It was placed in the Gastrothripina
by Priesner (1961), but the metathoracic sternopleural sutures are not developed, and there are
two sense cones on antennal segments III and IV (Fig. 290). One macroptera has been studied
64 L. A. MOUND AND J. M. PALMER
and this bears two pairs of wing-retaining setae on the median abdominal tergites. On the basis
of these characters the genus is here provisionally referred to the Elaphrothripina. As in
Malesiathrips , to which it is probably related, the metathoracic anapleural sutures appear to be
long and complete (Fig. 281), but the basal antennal segments do not bear long setae. These two
genera are not closely related to Elaphrothrips but may represent relicts associated with the
evolution of the Hystricothripina.
SPECIES INCLUDED
hawai/eiisis Bagnall, 1910ft: 678-80. Lectotype $, MAUAI Is. (BMNH).
DINOTHRIPS Bagnall
(Figs 277-279, 298, 310)
Dinothrips Bagnall, 1908ft: 190. Type-species: Dinothrips sumatrensis Bagnall, by monotypy.
Paxillothrips Ananthakrishnan, 1961a: 250. Type-species: Paxillothrips longicauda Ananthakrishnan, by
monotypy. [Synonymised by Palmer & Mound, 1978: 166.]
The five species now recognized in this genus were revised recently by Palmer & Mound (1978)
with a discussion of the patterns of variation and resultant complex synonymy. In both sexes the
pelta is divided completely into three segments (Fig. 310); however the genus is usually
recognised by the presence in males of a curiously expanded mesothoracic spiracular process
(Fig. 278). Unfortunately, although typically forked in some species this process is simple in
others, and is reduced or absent in small males (Fig. 279) as well as all females. Dinothrips
species resemble short-bodied Elaphrothrips species in general appearance, but they share with
Tiarothrips the apomorphic character state of having the foreocellus situated just posterior to
the major ocellar setae (Fig. 277).
SPECIES INCLUDED
juglandisMoulton, 1933ft: 6. Holotype cf , INDIA (BMNH).
longicauda (Ananthakrishnan, 1961o: 250-3) (Paxillothrips}. Holotype $, INDIA (TNA).
monodon Karny, 1920ft: 204. Holotype cf , PHILIPPINES (SMF).
spjnosus(Schmutz, 1913: 1078) (Ischyrothrips). Holotype $, SRI LANKA (NMV).
affinis Bagnall, 1915c: 270. Lectotype cf , BORNEO (BMNH).
crassiceps Bagnall, 1921c: 399 (Dicaiothrips). Holotype <j>, BURMA (BMNH).
jacobsoni Karny, 1921: 283. Holotype cf , JAVA (SMF).
kemneri Karny, 1923: 294. Lectotype cf , JAVA (SMF).
anodon Karny, 1923: 295. Syntypes cf $, JAVA (unknown).
gardneri Moulton, 1928e: 290. Holotype cf , INDIA (CAS).
malloti Moulton, 1933ft: 6. Holotype cf , INDIA (BMNH).
celebensis Bagnall, 1934a: 485. Holotype cf , SULAWESI (MNHN).
sumatrensis Bagnall, 1908ft: 191. Lectotype cf , SUMATRA (BMNH).
furcifer Schmutz, 1913: 1026. Holotype cf , SRI LANKA (?lost).
fulmeki Priesner, 1959: 55. Holotype cf , SUMATRA (SMF).
ELAPHROTHRIPS Buffa
(Figs 270-272, 282, 284-286, 293, 299, 307-309)
Elaphrothrips Buffa, 1909: 162-3. Type-species: Idolothrips coniferarum Pergande, by subsequent de-
signation, Andre, 1940: 76.
Dicaiothrips Buffa, 1909: 169-70. Type-species: Thrips schottii Heeger, by subsequent designation,
Bagnall, 1910: 370. [Synonymised by Hood, 1927: 238-9.]
Klinothrips Bagnall, 1918: 217-8. Type-species: Klinothrips femoralis Bagnall, by monotypy. [Synony-
mised by Priesner, 1952: 845.]
Elaphrothrips (Elaphoxothrips) Bagnall, 1932: 517. Type-species: Kleothrips athletes Karny, by mono-
typy.
Elaphridothrips Priesner, 1932: 320. Type-species: Elaphridothrips andrapterus Priesner, by monotypy.
[Synonymised by Priesner, 1952: 861.]
Palinothrips Hood, 1952c: 168. Type-species: Palinothrips palustris Hood, by monotypy. Syn. n.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 65
Elaphrothrips (Paradinothrips) Priesner, 1952: 846. Type-species: Elaphrothrips (Paraclinothrips) coniger
Priesner, by monotypy.
Elaphrothrips (Cradothrips) Ananthakrishnan, 1973a: 273. Type-species: Elaphrothrips (Cradothrips)
insignis Ananthakrishnan, by monotypy.
Palmer & Mound (1978: 172) point out that Elaphridia is unrelated to Elaphrothrips, and the
genus is here treated in the Pygothripini. Elaphridothrips was erected for a single apterous
species with small eyes, reduced wing-retaining setae and a degenerate, transverse pelta.
Dicaiothrips appears to have been erected for males, in contrast to the female characteristics
stressed for Elaphrothrips. Bagnall erected Elaphoxothrips without discriminatory characters,
adding 'should a subgeneric name appear desirable, I suggest . . .'. Klinothrips and Paradi-
nothrips were each proposed for single species with greatly enlarged forefemora in males
(Fig. 285), but as the femora are subject to allometry, that is they are not enlarged in small
males, the group names are of little significance. In contrast, Cradothrips was used for an Indian
species, remarkable in this group, that has tuberculate femora in the females. Finally, Pali-
nothrips was erected for a single species, taken from grasses in Brazil, which shows remarkable
parallelism with the Old World Ophthalmothrips in that the eyes are prolonged ventrally (Fig.
270). However, this species has very stout cheek setae and is here interpreted as an aberrant
species of Elaphrothrips.
More than 150 species-group names are available in Elaphrothrips, although 37 of these are
listed below in synonymy. Much of this synonymy was established by Mound (1968) recognising
sexual dimorphism and allometric variation, and by Palmer & Mound (1978) expanding the
concept of polytypic species each with widespread distributions. Prior to these studies, many of
the described species from the Oriental Region were known from single specimens or single
collections, and further studies on new material from the Afrotropical Region will undoubtedly
establish further synonymy. Hood (1955) and Hartwig (1948) have discussed some of the
patterns of variation which occur in this genus. Antennal segment lengths, head and tube length,
size of forelegs, also the pronotum and certain tubercles, are all subject to allometry, often being
greatly enlarged in larger males. In contrast the postocular setae are sometimes smaller in larger
individuals. This type of variation occurs within populations, but in addition different popula-
tions of a species can be expected to exhibit differences including different patterns of variation.
Elaphrothrips is the largest, most widespread, and probably the most diverse of the genera in
the Idolothripinae. It is found throughout the tropics, although apparently replaced by
Mecynothrips east of Sulawesi. There appear to be no essential differences between the
species-groups found in South America, Africa and India which would enable geographically
limited subgenera to be recognised. However, the pelta of several species in South America
tends to have the anterior sculpture differentiated from the posterior sculpture (Fig. 307); this
has not been reported in African and Oriental species. Africa is evidently the area of greatest
diversity of the genus, with numerous species ranging in body form from the large elaborate
males offemoralis to small, slender, almost featureless species on grasses. These latter species
are very similar to small species of Ophthalmothrips, whereas Lamillothrips is evidently derived
from the opposite extreme of the range of variation in Elaphrothrips. There appears to be no
justification, neither practical nor theoretical, in subdividing this large genus at present.
SPECIES INCLUDED
acanfAomerusHood, 1941: 217-20. Holotype cf, PERU (USNM).
*addendus Priesner, 1928c: 59-60. Syntypes cf , PARAGUAY (?lost).
*aethiopiae Bagnall, 1936: 225-6. Holotype cf , ? ETHIOPIA (MNHN).
affinis (Bagnall, 1908ft: 213-4) (Idolothrips). Holotype $, NICARAGUA (BMNH).
ossimilis Bagnall, 1908ft: 213 (Idolothrips}. Holotype $, NICARAGUA (BMNH).
distinctus Bagnall, 1910o: 378-9 (Dicaiothrips). Holotype cf , NICARAGUA (BMNH).
*a/ricanus(Trybom, 1908: 16-7) (Idolothrips). Syntypes $, TANZANIA (unknown).
*a«M>spinosusMoulton, 1929ft, 11-2. Holotype $, MEXICO (CAS).
amazomciisJohansen, 1978ft: 95-7. Holotype $, PERU (BMNH).
*amoenus Priesner, 1935o: 174, 241-2. Holotype cf , NORTH VIETNAM (SMF).
andrapterus (Priesner, 1932: 321-2) (Elaphridothrips). Syntypes cf , ZAIRE (MRAC).
66 L. A. MOUND AND J. M. PALMER
angusfafus (Bagnall, 1910a: 380-2) (Idolothrips) . Holotype cf , VENEZUELA (BMNH).
*angusticeps (Crawford, 1910: 168-70) (Idolothrips). Syntypes 9 cf, MEXICO; BELIZE; NICARAGUA; CUBA
(?CanadaD.of Agric.).
*angustifrons(Bergroth, 1888: xxx-xxxi) (Phloeothrips). ?Holotype, BRAZIL (?lost).
antennalis Bagnall, 1921c: 398. Holotype cf , JAPAN (BMNH).
armaf us (Hood, 1908c: 285-7) (Idolothrips). Lectotype $, U.S.A.: Illinois (USNM).
athletes (Karny, 1923: 355-8) (Kleothrips) . Holotype cf , JAVA (SMF).
aztecusHood, 1941: 208-13. Holotype cf , MEXICO (USNM).
ftagna7/iaiiiis Priesner, 1952: 863. [Replacement name for clarispinis Bagnall nee Priesner, 1935.]
darispinis Bagnall, 1935: 134-5. Holotype cf , ZAMBIA (BMNH).
bakeri (Karny, 1920ft: 206-9) (Dicaiothrips). Holotype cf , PHILIPPINES (SMF).
mentaweiensis Priesner, 19296: 201-4. Holotype cf , MENTAWEI Is. (SMF).
bakeri var. depokensis Priesner, 1935a: 159. Syntypes cf $, JAVA (SMF).
imitator Priesner, 1935a: 249-50. Holotype cf , JAVA (SMF).
*bilineatus Priesner, 1933c: 152. Holotype cf, MEXICO (SMF).
blatchleyiHood, 1938c: 410-3. Holotype ?, U.S.A.: Florida (USNM).
dorgmeieriHood, 1955: 62-6. Holotype cf , BRAZIL (USNM).
*bottegii(Butta, 1909: 170) (Dicaiothrips). ?Holotype cf , ?AFRICA (?lost).
brachypes Bagnall, 1934a: 495-7. Holotype cf , EAST AFRICA (BMNH).
jeanneli Bagnall, 1935: 140-2. Holotype $, KENYA (BMNH).
brachyurus Bagnall, 1926: 555. Holotype <J>, SOUTH AFRICA (BMNH).
brasiliensis Johansen, 19786: 104-5. Holotype cf , BRAZIL (BMNH).
breviceps (Bagnall, 1921c: 399-400) (Dicaiothrips). Holotype $, KENYA (BMNH).
brevicornis (Bagnall, 1910a: 379-80) (Dicaiothrips). Holotype $, VENEZUELA (BMNH).
brunneipennis Bagnall, 1935: 130-2. Lectotype $, SAO THOME (BMNH).
capens/sFaure, 1942: 81-3. Holotype $, SOUTH AFRICA (NCIP).
"carajoiiiBournier, 1971: 149-155. Holotype cf, CENTRAL AFRICAN REPUBLIC (MNHN).
* cognatograndis Johansen, 1976: 63-5. Holotype cf , MEXICO (UNAM).
congoensis Priesner, 1932: 334-5. Syntypes cf $, ZAIRE (MRAC).
can/aims Bagnall, 1934a: 497-8. Holotype $, VENEZUELA (MNHN).
coniferarum (Pergande, 1896: 63^) (Idolothrips). Syntypes, U.S.A.: Washington, D.C. (USNM).
coniger Priesner, 19526: 849-51 (subgenus Paraclinothrips). Holotype cf , GUINEA (MNHN).
f. gynaecoides Priesner, 19526: 851-2. Holotype cf , GUINEA (MNHN).
constrictopeltat us Johansen, 19786: 99-101. Holotype cf , PERU (BMNH).
*coreanusWoo, 1974: 69-70. Holotype $, KOREA (Seoul Univ.).
costo/ima/Hood, 1955: 57-60. Holotype cf , BRAZIL (USNM).
curvipes Priesner, 19296: 206-8. Syntype cf , MENTAWEI Is. (SMF).
karnyi Priesner, 1935a: 246-7. Holotype $, SUMATRA (SMF).
secus Ananthakrishnan, 1973o: 278. Holotype $, INDIA (TNA).
damp/JHood, 19406: 500-4. Holotype $, MEXICO (USMN).
decipiens Priesner, 1932: 331-3. Holotype cf , ZAIRE (MNHN).
defecfusHood, 1941: 213-7. Holotype cf , PERU (USNM).
denticollis (Bagnall, 1909c: 527) (Dicaiothrips). Holotype $, NIAS (BMNH).
beesoni Ramakrishna, 1934: 7. Syntypes cf $, INDIA (TNA).
mucronatus Priesner, 1935a: 167-8. Holotype cf , JAVA (SMF).
sumbanus Priesner, 1935a: 169-70. Holotype cf , SUMBA (SMF).
productus Priesner, 19350: 170-4. Holotype cf , SUMBA (SMF).
f. obscuricornis Priesner, 1935a: 171. Syntypes cf $, SUMBA (SMF).
*devius Priesner, 19526: 857-8. Holotype cf , CAMEROUN (MNHN).
distans Bagnall, 1935: 132-4. Holotype $, TANZANIA (BMNH).
*drepanatus (Priesner, 1927c: 82) (Dicaiothrips). Holotype cf, GUINEA (?lost).
drepanifer (Faure, 1925: 162-6) (Dicaiothrips). Holotype cf , SOUTH AFRICA (NCIP).
edouard/Jacot-Guillarmod, 19396: 46-52. Holotype cf , SOUTH AFRICA (AMG).
Meat us (Karny, 1912c: 150-1) (Dicaiothrips). Syntypes cf , WEST AFRICA (unknown).
*fa/7ax Priesner, 19526: 853-4. Holotype cf , CAMEROUN (MNHN).
*faurei Jacot-Guillarmod, 19390: 67-70. Holotype cf , MOZAMBIQUE (AMG).
femoralis (Bagnall, 1918: 218-9) (Klinothrips). Holotype cf , GHANA (BMNH).
flavipes(Hood, 19080: 377-8) (Idolothrips). Lectotype $, U.S.A.: Illinois (USNM).
foveicollis (Bagnall, 19086: 214-5) (Idolothrips). Lectotype $, GUATEMALA (BMNH).
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 67
championi Bagnall, 19100: 375 (Dicaiothrips) . Holotype $, GUATEMALA (BMNH).
grandis Bagnall, 19100: 373-4 (Dicaiothrips). Holotype d", GUATEMALA (BMNH).
/ii/mdW Priesner, 19350: 242-3. Holotype cf , SUMATRA (SMF).
gaboniensis Bagnall, 1936: 224-5. Holotype ?, GABON (MNHN).
genaspinosusMoulton, 19286: 245-7. Holotype $, ETHIOPIA (BMNH).
*gnidiicolus (Hesse, 1934: 434-440) (Dicaiothrips). Holotype cf , SOUTH AFRICA (?lost).
*gracilis Moulton, 19330: 410-1. Holotype $, BRAZIL (CAS).
*grav/sPriesner, 19526: 852-3. Holotype $, CAMEROUN (MNHN).
greeni (Bagnall, 19146: 289) (Dicaiothrips). Holotype cf , SRI LANKA (BMNH).
bouvieri Vuillet, 1914: 276 (Dicaiothrips). Holotype cf , INDIA (BMNH).
micidus Ananthakrishnan, 19730: 275-6. Holotype $, INDIA (TNA).
*guachichiUs Johansen, 19776: 53-5. Holotype cf , MEXICO (UN AM).
*herricki Moulton, 19330: 411-3. Holotype cf , PERU (CAS).
impensusMorison, 1958: 595-7. Holotype $, ETHIOPIA (BMNH).
indagatorHood, 1936d: 436-40. Holotype cf , PERU (USNM).
insignis Ananthakrishnan, 19730: 273-5 (subgenus Cradothrips). Holotype $, INDIA (TNA).
*insperatus Johansen, 19780: 87-9. Holotype cf , MEXICO (UNAM).
*/nsu/ar/s Priesner, 1928c: 57-8. Holotype cf , JAVA (lost).
jacofesoiH Priesner, 19350: 243-6. Holotype cf , SUMATRA (SMF).
jacofgiMtfarmodi Johansen, 19786: 101-4. Holotype cf , PERU (BMNH).
laevicollis (Bagnall, 19100: 375-6) (Dicaiothrips). Syntypes cf $, VENEZUELA (?lost).
laticeps Bagnall, 1935: 142-3. Holotype $, TANZANIA (BMNH).
*7atfcoriMs Jacot-Guillarmod, 1941: 96-100. Holotype $, SOUTH AFRICA (AMG).
longiceps (Bagnall, 19086: 211-3) (Idolothrips). Holotype cf, MEXICO (BMNH).
*/ongJspiJiJs Priesner, 1932: 329-30. Holotype cf , ZAIRE (MRAC).
*mabirensis (Priesner, 1925: 308-9) (Dicaiothrips). Holotype $, KENYA (unknown).
*macateeiHood, 1955: 60-2. Holotype $, BRAZIL (USNM).
* madagascariensis Bagnall, 1935: 138-40. Holotype 9, MADAGASCAR (MNHN).
magnus Johansen, 19786: 97-9. Holotype cf , PERU (BMNH).
mahensis (Bagnall, 19210: 283-4) (Dicaiothrips). Holotype cf , SEYCHELLES (BMNH).
rex Bagnall, 19210: 281-3 (Dicaiothrips}. Holotype cf, SEYCHELLES (BMNH).
hystrix Bagnall, 19210: 284-6 (Dicaiothrips). Holotype $, SEYCHELLES (BMNH).
malayensis (Bagnall, 1909c: 525) (Dicaiothrips). Holotype cf, NIAS (BMNH).
bruneitarsis Schmutz, 1913: 1070 (Dicaiothrips). Holotype cf , SRI LANKA (NMV).
var. levis Schmutz, 1913: 1072 (Dicaiothrips). Holotype $, SRI LANKA (NMV).
coronatus Bagnall, 19346: 631. Holotype $, SRI LANKA (BMNH).
maynei Priesner, 1932: 325-7. Syntypes cf $, ZAIRE (7MRAC).
mectfusHartwig, 1948: 85-96. Holotype cf , SOUTH AFRICA (NCIP).
microacanthomerus Johansen, 19786: 105-7. Holotype $, PERU (BMNH).
*neodampfi Johansen, 19776: 55-7. Holotype $, MEXICO (UNAM).
* neoleonensis Johansen, 19776: 51-3. Holotype cf , MEXICO (UNAM).
*neolongiceps Johansen, 19780: 89-92. Holotype cf , MEXICO (UNAM).
*niger Jacot-Guillarmod, 19396: 56-60. Holotype cf , SOUTH AFRICA (AMG).
*nigricornis (Karny, 1912c: 139, 150) (Idolothrips). Syntypes $, Rio MUNI ('Spanish Guinea') (SMF).
nigripes Jacot-Guillarmod, 1937: 28-31. Holotype cf , MOZAMBIQUE (AMG).
nitidus (Bagnall, 19100: 372-3) (Dicaiothrips). Holotype cf , BRAZIL (BMNH).
note bills Ananthakrishnan, 19730: 276-8. Holotype <j>, INDIA (TNA).
*oculatoides Priesner, 1932: 333-4. Holotype cf , ZAIRE (MRAC).
oculat us Moulton, 19286: 243-5. Holotype $, ETHIOPIA (BMNH).
orangiae Jacot-Guillarmod, 1937: 31-4. Holotype $, SOUTH AFRICA (AMG).
palustris (Hood, 1952c: 168) (Palinothrips). Lectotype <j>, BRAZIL (USNM). Comb. n.
*paradampfi Johansen, 19776: 57-9. Holotype $, MEXICO (UNAM).
parallel™ Hood, 1924: 315-7. Holotype $, U.S.A.: Florida (USNM).
*parvus Priesner, 19366: 102. Holotype cf , SUDAN (SMF).
peruviens/sHood, 1936d: 443-6. Holotype cf , PERU (USNM).
powelli Jacot-Guillarmod, 1937: 25-8. Holotype cf , SOUTH AFRICA (AMG).
*pr/esneri Bagnall, 1926: 554. [Replacement name for breviceps Priesner, nee Bagnall.]
breviceps Priesner, 1921: 219 (Dicaiothrips). Syntypes cf $, PARAGUAY (ZMB).
procer (Schmutz, 1913: 1063) (Dicaiothrips). Holotype cf , SRI LANKA (NMV).
68 L. A. MOUND AND J. M. PALMER
novus Schmutz, 1913: 1066 (Dicaiothrips}. Holotype cf , SRI LANKA (NMV).
dallatorensis Schmutz, 1913: 1067 (Dicaiothrips}. Holotype cf , SRI LANKA (NMV).
proximus Bagnall, 19146: 289 (Dicaiothrips}. Holotype cf , SRI LANKA (BMNH).
achaetus Bagnall, 19346: 633. Holotype $, SRI LANKA (BMNH).
approximatus Bagnall, 19346: 635. Holotype cf , SRI LANKA (BMNH).
chandana Ramakrishna, 1934: 9. Holotype cf , INDIA (unknown).
eranthemi Seshadri & Ananthakrishnan, 1954: 224. Holotype cf , INDIA (TNA).
propinquus (Bagnall, 1910a: 377-8) (Dicaiothrips). Holotype cf , VENEZUELA (BMNH).
*prospectorHood, 1936d: 440-3. Holotype cf , PERU (USNM).
*scAo«ii(Heeger, 1852a: 139) (Thrips). Holotype cf , BRAZIL (?lost).
*sc/K>uftt/eiii Priesner, 1932: 327-9. Holotype $, ZAIRE (MRAC).
*scAu7teei Priesner, 1933c: 152. Holotype $, MEXICO (ZMB).
seasitiviis Priesner, 19296: 204-6. Holotype cf , MENTAWEI Is. (SMF).
"separates Priesner, 1928c: 58-9. Holotype $, TANZANIA (lost).
seychellensis (Bagnall, 1921a: 280) (Dicaiothrips}. Holotype $, SEYCHELLES (BMNH).
*snodgrass/Hood, 1955: 66-9. Holotype cf , BRAZIL (USNM).
spiniceps Bagnall, 1932: 514. Holotype <j>, INDIA (BMNH).
graveleyi Bagnall, 19346: 628. Holotype $, INDIA (BMNH).
clarispinis Priesner, 1935a: 247-9. Holotype cf , JAVA (SMF).
spinipr/Vus Priesner, 19526: 855-6. Holotype cf , MALAWI (BMNH).
sp/nosusMoulton, 1933a: 413-4. Holotype $, COLOMBIA (CAS).
stenocephalus (Bagnall, 19146: 288-9) (Dicaiothrips). Holotype cf , TANZANIA (BMNH).
nigrospinosus Bagnall, 1932: 515-6. Holotype cf , TANZANIA (BMNH).
atrispinus Bagnall, 1935: 135-7. Holotype cf , EAST AFRICA (BMNH).
variispinis Bagnall, 1935: 137-8. Holotype $, TANZANIA (BMNH).
*surinamensis Priesner, 1925: 306-8. Syntype $ , SURINAM (?SMF).
*tener Priesner, 1925: 305-6. Holotype cf , MEXICO (SMF).
* transvaalensis Jacot-Guillarmod, 19396: 60-2. Holotype cf , SOUTH AFRICA (AMG).
tuberculatus(Hood, 1908c: 287-9) (Idolothrips). Lectotype $, U.S.A.: Illinois (USNM).
imico/orMoulton, 1933a: 415-6. Holotype $, BRAZIL (CAS).
*iHM/brm/sBuffa, 1909: 164. ?Holotype $, GUINEA-BISSAU (?lost).
wtt/pennisHood, 19400: 579-83. Holotype cf , U.S.A.: Arizona (USNM).
*zefetisHood, I936d: 432-6. Holotype cf, PERU (USNM).
HARTWIGIA Faure
(Figs 268, 283, 294, 304)
Hartwigia Faure, 19496: 208-10. Type-species: Hartwigia tumiceps Faure, by monotypy.
Faure compared this genus to Docessissophothrips, although the only known species has broad
maxillary stylets (Fig. 268) and the antennae bear four sense cones on segment III. Priesner
(1961) placed the genus in the Compsothripini, and Stannard (1976) erected a monobasic
subtribe 'Hartwigia' . This family-group name is rejected here on the grounds that it is incorrectly
formed. Hartwigia is recognised as related to Elaphrothrips because of the two pairs of
wing-retaining setae on each tergite (Fig. 294) and the complete absence of metathoracic
sternopleural sutures. The ant-like body form is not unlike that of some Ophthalmothrips
species, and the fore femora of the males bear a group of stout setae on the external margin
basally.
SPECIES INCLUDED
tumiceps Faure, 19496: 210-2. Holotype cf , SOUTH AFRICA (NCIP).
LAMILLOTHRIPS Bagnall
(Figs 275, 312)
Lamillothrips Bagnall, 1923: 630-1. Type-species: Lamillothrips typicus Bagnall, by monotypy.
Hylothrips Priesner, 1932: 336. Type-species: Hylothrips aethiopicus Priesner, by original designation.
Syn. n.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 69
The five African species which have been placed in these two genera remain very poorly known.
There is no doubt that the species are congeneric, the male holotype of vitulus and a female
paratype of aethiopicus having been compared with the type-material of Bagnall's three species.
There is so little difference between these few available specimens that they may, in fact,
represent only a single variable and widespread species. If this should prove to be so, then
Lamillothrips itself may best be regarded as a synonym of Elaphrothrips in view of the few
differences by which it may be distinguished. Machatothrips, to which Lamillothrips has
previously been compared, belongs in the Macrothripina.
SPECIES INCLUDED
aethiopicus (Priesner, 1932: 337-9) (Hylothrips). Holotype cf , CONGO (MRAC). Comb. n.
fypicus Bagnall, 1923: 631. Lectotype cf , GHANA (BMNH).
pennicollis Bagnall, 1923: 631. Holotype cf , GHANA (BMNH).
longidens Bagnall, 1934a: 491-2 (Machatothrips). Holotype tf, SIERRA LEONE (BMNH).
vitulus (Karny , 192(k: 109-111) (Macrothrips) . Holotype cf, CAMEROUN (SMF).
MALESIATHRIPS Palmer & Mound
(Figs 273, 274, 280, 291, 292, 296, 305)
Malesiathrips Palmer & Mound, 1978: 196. Type-species: Malesiathrips malayensis Palmer & Mound, by
original designation.
This genus, with three species from the Oriental and Pacific Regions, appears to be closely
related to Dermothrips from Hawaii. They differ in that Malesiathrips species bear a large seta
on the dorsal surface of antennal segment II (Figs 291, 292) as in some species of Hystricothri-
pina. The two genera are here included in the Elaphrothripina provisionally, because of their
lack of metathoracic sternopleural sutures, and the presence of long, complete anapleural
sutures (Fig. 280). Two species of Malesiathrips have two pairs of wing-retaining setae on each
tergite (Fig. 296) although solomoni only has one pair.
SPECIES INCLUDED
guamensis Palmer & Mound, 1978: 196-8. Holotype $, GUAM (BMNH).
malayensis Palmer & Mound, 1978: 198_9. Holotype $, MALAYA (BMNH).
solomoni (Mound, 1970: 116-8) (Atractothrips). Holotype $, SOLOMON Is. (BMNH).
MECYNOTHRIPS Bagnall
(Figs 257-262, 287, 288, 297, 306)
Mecynothrips Bagnall, 1908a: 356. Type-species: Mecynothrips wallacei Bagnall, by monotypy.
Phoxothrips Karny, 1913c: 132. Type-species: Phoxothripspugilator Karny, by monotypy.
Kleothrips Schmutz, 1913: 1057. Type-species: Kleothrips gigans Schmutz, by monotypy.
Dracothrips Bagnall, 19146: 290. Type-species: Dracothrips ceylonicus Bagnall, by monotypy.
Acrothrips Karny, 1920c: 43. Type-species: Acrothrips sorex Karny, by monotypy.
Kleothrips (Synkleothrips) Priesner, 1935a: 330. Type-species: Kleothrips (Synkleothrips) innocens Pries-
ner, by monotypy.
Kleothrips (Akleothrips) Priesner, 1935a: 332. Type-species: Kleothrips (Akleothrips) karimonensis
Priesner, by original designation.
The generic synonymy listed above, also the structural variation shown by several species, has
been discussed extensively by Palmer & Mound (1978: 200). The large number of names
available has arisen because of earlier failures to recognise the variability of several species. The
forefemora of large males, for example, often bear one or more large tubercles (Figs 287, 288),
whereas the forefemora of small males of the same species are slender and lack tubercles.
Similarly variable can be the position of the foreocellus (Figs 257-259), the form of the antennal
setae, or even the number and position of the cheek setae on the head. Palmer & Mound (1978)
recognised three species-groups in the genus: the simplex-group from East Africa to the
Philippines and Japan; the wallacei-group from New Guinea, Australia and the Solomon
Islands; and the acanthus-group from Java, Sumba and Australia. Mecynothrips is evidently
70 L. A. MOUND AND J. M. PALMER
closely related to Elaphrothrips , which it replaces in the Old World Tropics east of Sulawesi.
However, the remarkable presence of three pairs of major tergal wing-retaining setae (the
anterior pair arises, close to the tergal antecostal ridge) suggests that Mecynothrips is the
phylogenetic sister-group of the other Old World Elaphrothripina (Fig 297).
SPECIES INCLUDED
acanf/iiis(Hood, 1918a: 77) (Kleothrips}. Holotype cf , AUSTRALIA (USNM).
serex Karny, 1920c: 43 (Acrothrips) . Lectotype cf , AUSTRALIA (SMF).
gargantua Girault, 1926: 1 (Acrothrips). Lectotype cf , AUSTRALIA (QMB).
giganteus Girault, 1926: 4 (Phoxothrips) . Holotype cf , AUSTRALIA (QMB).
afrafus(Hood, 1919a: 69) (Kleothrips). Holotype $, EAST AFRICA (USNM).
zuluensis Jacot-Guillarmod, 1939a: 70 (Kleothrips- Akleothrips). Holotype cf , SOUTH AFRICA (AMG).
goliath (Priesner, 1935a: 327) (Kleothrips). Holotype cf , SUMBA (SMF).
Aardyi (Priesner, 19286: 657) (Kleothrips). Holotype $, AUSTRALIA (SMF).
*Jtano/(Takahashi, 1937: 343) (Kleothrips). Syntypes $, TAIWAN (unknown).
karimonensis (Priesner, 1935a: 332) (Kleothrips- Akleothrips). Holotype cf , JAVA (SMF).
f. parvidens Priesner, 1935a: 334. Lectotype cf , JAVA (SMF).
kraussi Palmer & Mound, 1978: 205-6. Holotype cf , SOLOMON Is. (BMNH).
lacerta (Priesner, 1935a: 329) (Kleothrips). Lectotype $, SUMBA (SMF).
innocens Priesner, 19350: 331 (Kleothrips-Synkleothrips). Holotype cf , SUMBA (SMF).
priesneri Mound, 19716: 281. Holotype cf , NEW Guinea (BPBM).
minor Mound, 19716: 282. Holotype cf , NEW GUINEA (BPBM).
pugilator (Karny, 1913c: 132) (Phoxothrips). Holotype cf , TAIWAN (unknown).
takahoshii Priesner, 1935c: 372 (Elaphrothrips). Holotype <J>, JAPAN (SMF).
s/inp7ex Bagnall, 1912: 216. Holotype cf , PHILIPPINES (BMNH).
gigans Schmutz, 1913: 1058 (Kleothrips). Syntypes cf $ , SRI LANKA (NMV).
ceylonicus Bagnall, 19146: 290 (Dracothrips). Syntypes cf , SRI LANKA (lost).
agama Priesner, 1935a: 323 (Kleothrips). Holotype cf, JAVA (SMF).
snodgrass/Hood, I952d: 294. Holotype cf , SOLOMON Is. (USNM).
to/wamisOkajima, 19796: 127. Holotype cf , TAIWAN (OCT).
wallacei Bagnall, 1908a: 357. Holotype cf , NEW GUINEA (BMNH).
magnus Girault, 1929: 1. Syntypes cf , AUSTRALIA (QMB).
bagnalli Priesner, 1935a: 335. Holotype $, KEI Is. (SMF).
f. imbecilla Priesner, 1935a: 338. Syntypes cT $, KEI Is. (SMF).
OPHTHALMOTHRIPS Hood
(Figs 264-267, 289, 302, 303)
Ophthalmothrips Hood, 1919a: 67. Type-species: Ophthalmothrips pomeroyi Hood, by monotypy.
Pyrgothrips Karny, 1924: 35-6. Type-species: Pyrgothrips conocephalus Karny, by monotypy. Syn. n.
Fulgorothrips Faure, 1933: 62-3. Type-species: Fulgorothrips priesneri Faure, by monotypy. [Synony-
mised with Pyrgothrips by Mound, 1974a: 89.] Syn. n.
Derothrips Jacot-Guillarmod, 1940: 133. Type-species: Derothrips amyae Jacot-Guillarmod, by monoty-
py. Syn. n.
This genus has not been recognised since its original description, although both Mound (19740)
and Haga (1975) have given an account of several species under the name Pyrgothrips. The
recognition of species within the genus remains problematical, due to structural variation
related to sex, morph and size, and particularly due to the rarity with which species have been
collected in series. For example, several series collected recently in eastern Africa did not
include macropterae and micropterae together at any one site. Not only is the variation within
species poorly understood at present, but the range of form produces difficulties in defining the
genus itself. In the holotype of pomeroyi, as well as in two macropterae from Malawi and two
from Lourenco Marques, the foreocellus clearly overhangs the bases of the antennae (Fig. 267).
However, this is not usually true of macropterae identified as priesneri (Fig. 264). The pelta is
characteristically triangular throughout the genus except for amyae in which it is broadly
rounded as in Bolothrips species (Figs 302, 303). The species of Ophthalmothrips are all small,
with rather slender bodies and the eyes prolonged ventrally (Figs 264-7). They apparently all
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 71
live at the base of grass tussocks, and the genus appears to have been derived from small-bodied
species of Elaphrothrips which have become specialised to this habitat. One undescribed species
of Elaphrothrips, from grass tussocks in Tanzania, has been studied which has the general body
form of Ophthalmothrips species but has small rounded eyes which are not prolonged ventrally.
O. amyae is here interpreted as representing the extreme of a tendency within this group for the
body to foreshorten, the head in particular being short and broad. The species lesnei (head
production 103/50 to 60 /mi) is very probably the same as pomeroyi (head production 110/57
/mi). However, all the available names in the genus are listed separately below with no
subjective synonymy in view of the differences of interpretation between Mound (19740) and
Haga(1975).
SPECIES INCLUDED
a/njae(Jacot-Guillarmod, 1940: 135-8) (Derothrips) . Holotype <J>, SOUTH AFRICA (AMG). Comb. n.
ftrevjceps(Bagnall, 1914c: 380-1) (Phoxothrips). Holotype d" , INDIA (BMNH). Comb. n.
conocephalus (Karny , 1924: 36) (Pyrgothrips). Holotype cf , AUSTRALIA (NRS). Comb. n.
/aure/(Ananthakrishnan, 1964a: 96) (Fulgorothrips) . Holotype 9, INDIA (TNA). Comb. n.
formosanus (Karny, 1913c: 130-1) (Idolothrips). Holotype $, TAIWAN (ZMB). Comb. n.
lesnei (Priesner, 1952ft: 878) (Fulgorothrips). Holotype $, MOZAMBIQUE (MNHN). Comb. n.
7ong/ceps(Haga, 1975: 270) (Pyrgothrips). Holotype $, JAPAN (MNHO). Comb. n.
miscanthicola (Haga, 1975: 273) (Pyrgothrips). Holotype <j>, JAPAN (MNHO). Comb. n.
pomeroyiHood, 19190: 67. Holotype $, EAST AFRICA (USNM).
priesuer/(Faure, 1933: 63-5) (Fulgorothrips). Holotype C", SOUTH AFRICA (AMG). Comb. n.
TIAROTHRIPS Priesner
(Figs 269, 301)
Tiarothrips Priesner, 1935a: 251. Type-species: Kleothrips subramanii Ramakrishna, by monotypy.
The only species in this genus shares with Dinothrips (and some Mecynothrips) the position of
the foreocellus just posterior to the major ocellar setae (Fig. 269). This may suggest that
Tiarothrips and Dinothrips are sister-groups, and together constitute the sister-group of
Elaphrothrips, Lamillothrips and Ophthalmothrips. The species subramanii is remarkable for
the extreme allometry in the length of the preocular head process as well as the third antennal
segment.
SPECIES INCLUDED
subramanii (Ramakrishna, 1925: 788) (Kleothrips). Holotype d", INDIA (? TNA).
Genera of Idolothripina
As used here, this group includes 10 genera and corresponds to the Megathripina of Priesner
(1961) with the notable addition of Meiothrips and Idolothrips. The genus Meiothrips is closely
related to Bactrothrips, females being allocated between these genera only with difficulty, and
Meiothrips is also related to Idolothrips. Priesner, however, placed these genera with Acti-
nothrips and related Neotropical genera with a slender body-form which are here treated in the
Hystricothripina.
Males in the Idolothripina frequently bear tubercles (drepanae) laterally on the abdomen
(Figs 324, 325). However, it must be emphasised that, despite this tendency, species lacking
tubercles in the male are now known in both Megathrips and Bactrothrips. Similarly, the
abdominal tube of Idolothripina species usually bears a number of distinct setae laterally (tube
hairy), but these setae are sometimes decumbent or weakly developed. The metathoracic
sternopleural sutures are not developed, as in the rest of the Idolothripini, but the metathoracic
anapleural sutures (Fig. 327) tend to be shorter in the Idolothripina than in the Elaphrothripina
and Hystricothripina.
Within the Idolothripina only one genus contains more than 10 species; this is Bactrothrips
which is widespread throughout the Old World Tropics. Megathrips is a Holarctic derivative of
72 L. A. MOUND AND J. M. PALMER
Bactrothrips from which it can only be distinguished by the superficial characteristic of the rather
shorter head with maxillary stylets more deeply retracted (Fig. 321). Similarly, Ceuthothrips
appears to be a Neotropical, monobasic, derivative of Megathrips. In contrast Meiothrips and
Idolothrips are the eastern Oriental and Australian derivatives of Bactrothrips, and Cylin-
drothrips with one species from South Africa may also be related. Badllothrips and Mega-
lothrips appear quite distinct from the other genera in the Idolothripina (except possibly
Lasiothrips) in having the maxillary stylets greatly elongate and close together in the middle of
the head. These genera are normally regarded as Holarctic, but a new species of Megalothrips is
described below from Malaya. However, elongation of the maxillary stylets is undoubtedly
polyphyletic, possibly an adaptation to feeding on fungal spores in more confined situations. For
example, the genus Zeuglothrips alone amongst the Hystricothripina has elongate stylets.
BACILLOTHRIPS Buffa
(Figs 315, 330)
Badllothrips Buffa, 1908: 385-6. Type-species: Badllothrips linearis (now regarded as a synonym of
Megalothrips longiceps Reuter), by monotypy.
This genus is usually treated as monobasic, including just the Mediterranean species longiceps.
However, the present authors consider that both Megathrips nobilis and Docessissophothrips
longiceps should also be placed in Badllothrips, and as a result the latter species is here renamed
bagnalli. These three species have the head long and slender with the maxillary stylets deeply
retracted and close together medially (Fig. 315). Moreover, the lateral lobes of the pelta are
sharply cut off from the median lobe (Fig. 330), whereas these lateral lobes in Megalothrips
species are more slender (Fig. 329). The metanotum of longiceps and nobilis has reticulate
sculpture, whereas the sculpture on bagnalli is transverse. The head of bagnalli is clearly
elevated in the mid line, although the heads of all three species appear to be essentially similar.
SPECIES INCLUDED
bagnalli nom. n. for longiceps Bagnall not longiceps Reuter.
longiceps Bagnall, 1916: 407-8 (Docessissophothrips). Holotype $ , MADEIRA (BMNH).
longiceps (Reuter, 1901: 215-6) (Megalothrips). Syntypes d", CORFU (unknown).
linearis Buffa, 1908: 386-7. Syntypes $ cf , ITALY; SARDINIA (unknown).
nobilis (Bagnall, 1909ft: 130-1) (Megathrips). Lectotype cT, ENGLAND (BMNH). Comb. n.
BACTROTHRIPS Karny
(Figs 314, 316, 332, 333)
Bactrothrips Karny, 1912c: 131. Type-species: Bactrothrips longiventris Karny, by monotypy.
Eidothrips Bagnall, 1918: 219. Type-species: Eidothrips alluaudi Bagnall, by monotypy. Syn. n.
Krinothrips Bagnall, 1918: 220. Type-species: Krinothrips divergens Bagnall, by monotypy. [Synonymised
by Bagnall, 1921.]
Bactridothrips Karny, 1919: 116. Type-species: Bactridothrips idolomorphus Karny, by monotypy. Syn. n.
Caudothrips Karny, 1921a: 230. Type-species: Caudothrips buffai Karny, by monotypy. Syn. n.
Bactrianothrips Bagnall, 1936: 226-7. Type-species: Bactrianothrips alluaudi Bagnall, by monotypy.
[Synonymised by Bournier, 1968: 157.]
Cervothrips Bagnall, 1936: 229. Type-species: Cervothrips berlandi Bagnall, by monotypy. Syn. n.
This group of genera was treated as a subfamily by Karny (1919). However, all of the genera are
recognisable only from secondary sexual characters of the male abdomen; none of them can be
recognised in the female sex. The primary characteristic of Bactrothrips is the presence in the
males of a pair of long tubercles laterally on the sixth abdominal segment. From this condition
the other genera have been defined as follows.
Bactrianothrips. A pair of truncate tubercules on VI; however, in the unique male holotype it is evident
that these represent the bases of broken long tubercles.
Cervothrips. Elongate tubercles on VI forked; however Bournier (1968) demonstrated that, in small
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 73
males, one branch of the fork is scarcely developed. This suggests that the difference is of no more than
specific value.
Eidothrips. Elongate tubercles on segments V and VI; only one species is known with this characteristic
and, in view of the similarity of the females, there seems little advantage in segregating it to a separate
genus.
Bactridothrips. Elongate tubercles on VI , also small tubercles on either or both of segments VII and VIII ;
this has been treated as a sub-genus by Bournier (1968) but in view of the variation it is not a useful
segregate.
In addition to the variation indicated above, Bactrothrips pitkini sp. n. is described below from
an apterous male without any abdominal tubercles. If the traditional generic concepts were
accepted then pitkini would need to be placed in yet another new genus.
The spiracles of abdominal segment eight are elongate dorsoventrally in the males of
divergens and kenyensis (Mound, 1968), but this is not true of the other available species.
However, the validity of many species described from the African continent is in doubt, because
so many are based on unique specimens or short series. Thus little or no account has been taken
of allometric growth patterns in the abdominal tubercles of large and small males. Moreover,
several species are based solely on females which cannot satisfactorily be associated with male
specimens. Therefore, of the 39 available names in the genus, several are likely to be recognised
as synonyms as soon as effective collecting is carried out. For this reason the homonymy of
alluaudi Bagnall, 1918 with alluaudi Bagnall, 1936 is here allowed to stand until such time as the
group can be re-examined comprehensively. It may not be entirely irrelevant to point out that
the range of variation in the abdominal tubercles of male Bactrothrips species, as interpreted
here, is no greater than that recognised in the two species of the genus Idolothrips.
The European species buffai and the Californian species Hesperus are the only species placed
in this genus from outside the Old World Tropics. These two species have rather shorter heads
than the tropical species of Bactrothrips, and as a result the ocellar triangle is more nearly
equiangular. However, the difference when measured is so slight that there seems no useful
purpose in segregating the two species to separate genera. Caudothrips is therefore placed as a
synonym of Bactrothrips, One unusual specimen has been studied from Japan with the eyes
prolonged on the ventral surface of the head, a characteristic otherwise not found in Bactrothrips
(although variable within Ophthalmothrips q.v. Elaphrothripina). The generic relationships of
Bactrothrips are discussed under Meiothrips and Idolothrips.
SPECIES INCLUDED
alluaudi (Bagnall, 1918: 219-20) (Eidothrips}. Lectotype cf , KENYA (BMNH). Comb. n.
alluaudi (Bagnall, 1936: 227-8) (Bactrianothrips). Syntypes cf $, MADAGASCAR (MNHN: 2 9 BMNH).
aferr/musPriesner, 1936a: 213-4. Holotype cf , UGANDA (BMNH).
*afrispinisPriesner, 1932: 220-1. Holotype <j>, ZAIRE (MRAC).
*oancoens/sPriesner, 1952ft: 867-8. Holotype $, IVORY COAST (MNHN).
berlandi (Bagnall, 1936: 229-30) (Cervothrips). Holotype cf , CONGO (MNHN).
brev/fu6usTakahashi, 1935: 61-3. Holotype cf , RYUKYU Is. (DART).
*bucculentus Bournier, 1968: 139-42. Holotype $, ANGOLA (MDA).
buffai (Karny, 1921a: 230) (Caudothrips). ITALY (types not designated). Comb. n.
lesnei Bagnall, 1933ft: 659-61 (Megathrips) . Holotype cf, ALGERIA (MNHN).
congoens/sPriesner, 1932: 215-6. Syntypes cf <j>, ZAIRE (MRAC).
*de/amareiPriesner, 1952ft: 868-70. Holotype cf , GUINEA (MNHN).
divergens (Bagnall, 1918: 220-1) (Krinothrips) . Syntypes cf ?, GHANA (BMNH).
ritchianus Bagnall, 1932: 517-8 (Actinothrips) . Holotype $, TANZANIA (BMNH).
* furcates Priesner, 1932: 216-8. Syntypes cf $, ZAIRE (MRAC).
*grand/sPriesner, 1932: 219-20. Holotype $, ZAIRE (MRAC).
*gu/neaens/sMoulton, 19470: 177-8. Holotype cf , NEW GUINEA (CAS).
*gu/neens/sPriesner, 1952ft: 866-7. Holotype $, GUINEA (MNHN).
hesperus(Moulton, 1907: 65-6) (Megalothrips). Syntypes cf ?, U.S.A.: California (CAS). Comb. n.
honoris (Bagnall, 1921c: 395) (Megathrips}. Holotype cf , JAPAN (BMNH). Comb. n.
*nood/ Bournier, 1968: 142-6. Holotype tf , ANGOLA (MDA).
ido/omorpnus (Karny, 1919: 117-8) (Bactridothrips). Holotype cf , MALAYA (SMF).
serraticornis Bagnall, 1921c: 397 (Bactridothrips). Holotype cf , SRI LANKA (BMNH).
74 L. A. MOUND AND J. M. PALMER
*/nermis (Karny, 1912c: 138-9) (Panurothrips) . Syntypes (sex not stated), Rio MUNI ('Spanish Guinea')
(unknown).
*Jtawamurai(Ishida, 1932: 2-3) (Idolothrips). Holotype 9> JAPAN (unknown). Comb. n.
KenyensisPriesner, 1935ft: 129-30. Syntypes cf $, KENYA (BMNH).
laingiB agnail, 1926: 558 (Bactridothrips}. Holotype cf , SIERRA LEONE (BMNH).
hargreavesi Bagnall, 1926: 555-6. (Actinothrips). Holotype $ , SIERRA LEONE (BMNH).
*7amotfe/Priesner, 1952ft: 870-1. Holotype cf , GUINEA (MNHN).
*7evidensPriesner, 1932: 218-9. Holotype cf , ZAIRE (MRAC).
*/ojigjsef/sBournier, 1968: 154-7. Holotype $, ANGOLA (MDA).
*Iongiventris Karny, 1912c: 131-2. Holotype cf , Rio MUNI ('Spanish Guinea') (ZMB).
luteus Ananthakrishnan, 1973ft: 81-4. Holotype $, INDIA (TNA; 5 cf paratypes, BMNH).
*macr0pfeiyx(Trybom, 1910: 523-5) (Megalothrips). Holotype cf , MADAGASCAR (MNHN).
ma/gassusBournier, 1967: 1022-6. Holotype cf , MADAGASCAR (MNHN; $ cf paratypes BMNH).
moultoni (Bagnall, 1932: 513-4) (Bactridothrips). Holotype cf , SOUTH AFRICA (BMNH).
nafa/eiis/sMoulton, 1930: 415-6. Holotype cf , SOUTH AFRICA (BMNH).
nativus(Girault, 1928c: 2) (Idolothrips). Holotype cf, AUSTRALIA (QMB).
*nigr/pesPriesner, 1932: 212-3. Holotype cf , ZAIRE (MRAC).
*pa//idicri/sPriesner, 1952ft: 871-2. Holotype $, CAMEROUN (MNHN).
*parvidens Priesner, 1932: 213-4. Holotype cf , ZAIRE (MRAC).
pitkinisp. n. Holotype cf , TANZANIA (BMNH).
priesner/Bournier, 1967: 1018-22. Holotype cT, MADAGASCAR (MNHN).
propinquus (Bagnall, 1936: 228-9) (Bactridothrips). Syntypes cf $, CONGO (MNHN & BMNH).
quadrituberculatus (Bagnall, 1908ft: 210-1) (Idolothrips). Holotype $, JAPAN (BMNH).
*titec/iadWBournier, 1968: 135-9. Holotype cf , ANGOLA (MDA).
Itxctrothrips pitkinisp. n.
(Figs 314, 333)
Apterous cf. Colour dark brown; antennal segments not bicoloured, III-IV yellow, V-VIII slightly
darker; distal half of tibiae, basal half of hind femora and all tarsi yellow.
Head more than twice as long as wide (Fig. 314); slightly prolonged in front of eyes; dorsal setae well
developed, cheeks with a few fine setae; maxillary stylets wide apart, retracted into head about halfway to
posterior margin of eyes.
Pronotal epimeral sutures weak; anteroangular setae small. Median metanotal setae well developed.
Pelta with lateral lobes narrowly joined to median lobe (Fig. 333). Abdominal tergites II-VIII with one
pair of small wing-retaining setae (anterior pair reduced or straight); lateral abdominal tubercles not
developed; tube surface smooth but set with fine setae, tapering more strongly at apex, about 4-0 times as
long as broad; setae B\ on tergite IX 0-25-0-30 as long as tube.
Measurements (holotype cf in ^tm). Body length 4225. Head, length 520, maximum breadth 220;
interocellar setae 70/74; postocellar setae 68/72; postocular setae I 70/74; postocular setae II 133/138.
Pronotum length 233; breadth 333; major setae, am 54/62, aa 23/25, ml 72, pa 108/114, epim 100/106.
Median metanotal setae 117/131. Tergite IX setae B± 126/157. Tube, length 527; maximum breadth 134.
Antennal segments III-VIII length, 290/295; 214; 176/181; 138/142; 90; 71.
Apterous 9 . Colour and structure similar to male. Setae BI on tergite IX longer, almost 0-5 times as long
as tube. Tube longer, 5-5 times as long as broad, and 1-4 times as long as head.
Measurements (paratype $ in ^m). Body length 4540. Head, length 520; maximum breadth 230;
interocellar setae 67/68; postocellar setae 58/62; postocular setae I 74/76; postocular setae II 148/150.
Pronotum, length 224; breadth 342; major setae, am 70/72, aa 24/36, ml 92/104, pa 134, epim 126/130.
Median metanotal setae 168/172. Tergite IX setae BI 369/372. Tube, length 728; maximum breadth 134.
Antennal segments III-VIII length, 276/285; 204/209; 171; 135/138; 95/100; 71/74.
Macropterous $ . Structure similar to apterous female, colour slightly darker. Abdominal tergites II-VII
with two pairs of wing-retaining setae, anterior pair small. Wings pale, with a pale brown median line in
basal half and 3 or 4 stout sub-basal setae.
Measurements (paratype $ in ^tm). Body length 5365. Head, length 543; maximum breadth 238;
interocellar setae 68/75; postocellar setae 60/66; postocular setae I 66/75; postocular setae II 160/168.
Pronotum, length 248; breadth 370; major setae, am 105, aa 34/38, ml 110/126, pa 144/149, epim 158.
Median metanotal setae 190. Forewings, length 1728/1746; maximum breadth 194; number of duplicated
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 75
cilia 20/23. Tergite IX setae 5t 338/363. Tube, length 855; maximum breadth 143. Antennal segments
IH-VIII length, 309/314; 223/233; 185/190; 128/143; 90/95; 71/76.
SPECIMENS STUDIED
Holotype d" aptera, Tanzania: Pare Mountains, Gonja, in grass tussock at 1000 m, 16. vi. 1974 (B R
Paratypes. 1 $ aptera taken with holotype; 1 $ macroptera with similar data, 13. vi. 1974 (BMNH).
COMMENTS. This species is unique in the genus, not only in lacking lateral abdominal tubercles in
the male, but also in the production of apterae. The apterae are smaller than the macroptera but
retain well-developed ocelli. The metaepimera of the two morphs are essentially similar, but the
tergal wing-retaining setae are reduced in the apterae. The lack of abdominal tubercles in the
male, a characteristic which is usually diagnostic of this genus, finds a parallel in Megathrips
inermis Priesner q.v. The description of pitkini includes separately the lengths of setae and
antennal segments from the left and right hand sides of the body in order to emphasise the
variation, and taxonomic problems associated with reliance on such characters.
CEUTHOTHRIPS Hood
(Figs 319, 328, 338)
Ceuthothrips Hood, 1938c: 406-7. Type-species: Ceuthothrips timuqua Hood, by monotypy.
This monobasic genus is known only from the type-series of six females and two males collected
in Florida. The metathoracic sternite and epimera are typical of the Idolothripina, but, unlike
other members of this group, both sexes bear a small foretarsal tooth. The head (Fig. 319) is
similar to Megathrips, the pelta (Fig. 328) similar to Megalothrips , but the antennae have a
curiously ill-formed, almost larval appearance (Fig. 338). The anterior pair of wing-retaining
setae is not developed on the tergites, but this is probably a secondary reduction (as in
Megathrips) associated with the evident reduction in length of the wings. Ceuthothrips is here
considered to be a Neotropical derivative of the Holarctic genus Megathrips.
SPECIES INCLUDED
f/muguaHood, 1938c: 407-10. Holotype $, U.S.A.: Florida (USNM).
CYLINDROTHRIPS Moulton
(Figs 320, 335)
Cylindrothrips Moulton, 1949: 496. Type-species: Cylindrothrips niger Mouhon, by monotypy.
This unusual monobasic genus was erected for a single male specimen collected in South
Western Africa. However, the unique male holotype of Derothrips turneri Moulton from the
same locality is here regarded as a larger specimen of the same species as niger. As first revisers,
within the meaning of the International Code of Zoological Nomenclature, the present authors
have placed turneri as a synonym of niger despite its page precedence. Moreover, the genus
Derothrips is here treated as a synonym of Ophthalmothrips q.v.
The original illustrations of the heads and pronota of both niger and turneri are inaccurate,
although both specimens are severely damaged. Despite these illustrations, the ocellar and
postocular setae of the specimens are essentially similar, and the median setae on the vertex of
niger have simply been removed in mounting. The main difference between the specimens lies in
the structure of the tube. In turneri the base of the tube bears laterally and dorsally numerous
stout teeth arising from the margins of each sculptured reticle, whereas in the much smaller niger
only a very few, small teeth are developed. This difference is here interpreted as being an
expression of allometric growth.
The genus Cylindrothrips is similar to Lasiothrips, Megalothrips and Ceuthothrips in having
the pronotum transverse. However, unlike Megalothrips the pronotal midlateral setae are well
separated from the anteroangulars, and moreover, the basal sculpture of the tube resembles that
of Idolothrips dissimilis and Meiothrips nepalensis. The head (Fig. 320) and antennae of
76 L. A. MOUND AND J. M. PALMER
Cylindrothrips niger are reminiscent of Ophthalmothrips species; however, the anapleural
suture is short and the anterior border of the anepisternum almost entire as in the other members
of the Idolothripina.
SPECIES INCLUDED
niger Moulton, 1949: 496-8. Holotype cf , SOUTH WEST AFRICA (BMNH).
turneri Moulton, 1949: 494-6 (Derothrips). Holotype cT, SOUTH WEST AFRICA (BMNH). Syn. n.
EGCHOCEPHALOTHRIPSBagnatt gen. rev.
Egchocephalothrips Bagnall, 1916: 408. Type-species: Docessissophothrips monstrosus Bagnall, by
monotypy.
Although treated as a synonym of Docessissophothrips by Mound (1968), this genus is here
accepted as valid. It is based on a single damaged specimen (? 9) which lacks the abdominal tube
as well as the distal antennal segments. However, the metathoracic sternopleural sutures are not
developed, the anapleural sutures are short and incomplete, the praepectus well developed, the
maxillary stylets are deeply retracted and parallel medially in the head, the anteocellar pair of
setae are elongate, the pelta has slender lateral wings, the median metanotal setae are very
stout, the tergites have two pairs of wing-retaining setae with one or more additional setae
directed mesad, and there are two sense cones on antennal segment III and four on segment IV.
These characters are all shared with Megalothrips . The pronotum of the unique holotype is very
short (correlating with the exceptional dorsal elevation of the head) and the fact that the
pronotal sutures are complete may be due to cover-slip pressure. Both this genus and
Lasiothrips (q.v.) may eventually prove to be synonyms of Megalothrips.
SPECIES INCLUDED
monstrosus (Bagnall, 1909c: 538-9) (Docessissophothrips). Holotype ?$, NEW CALEDONIA (BMNH).
IDOLOTHRIPS Haliday
(Figs 317, 325, 334)
Idolothrips Haliday in Walker, 1852: 1096. Type-species: Idolothrips marginatus Haliday (now regarded as
a synonym of Idolothrips spectrum Haliday), by subsequent designation, Bagnall, 1908: 356.
Acanthinothrips Bagnall, 1908: 207. Type-species: Idolothrips spectrum Haliday, by monotypy.
Froggatt (1904) demonstrated that the species spectrum Haliday is not only sexually dimorphic,
but that the males exhibit a wide range of variation in the size and the number of the lateral
tubercles and their setae depending on the overall body size. This variation is not always
bilaterally symmetrical (Mound, 1968: fig. 55), and is complicated by the fact that long tubercles
bear short stout setae whereas short tubercles bear long slender setae. This variation is reflected
in the number of available names for spectrum.
The two species currently placed in Idolothrips are both known only from Australia (Mound,
1974a). However, the females of these species, also the head and thorax of the males, are very
similar to species of Meiothrips and Bactrothrips , and no reliable characters for distinguishing
between these genera have been found apart from those given in the key. The base of the tube in
male /. dissimilis is similar to that of Meiothrips nepalensis in having a paired row of recurved
tubercles dorsally and numerous small teeth laterally (Fig. 325), and the relationships of these
genera are further discussed under Meiothrips. The tube of male Cylindrothrips is also similar
but shorter.
Jacot-Guillarmod (1978) retains halidayi Newman, 1855 under Idolothrips; however, this
species would be known more conveniently as Gigantothrips halidayi (Newman) comb. n.
(Phlaeothripinae). The two species currently remaining in Idolothrips have been keyed by
Mound (19740).
SPECIES INCLUDED
d/ssfjiu/feGirault, 19270: 2. Holotype d", AUSTRALIA: Queensland (QMB).
spectrum Haliday in Walker, 1852: 1097. ? Syntypes cf , NEW HOLLAND (depository unknown).
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 77
marginata Haliday in Walker, 1852: 1097. ? Syntypes d" , NEW HOLLAND (depository unknown).
lacertina Haliday in Walker, 1852: 1097. ? Syntypes cf , NEW HOLLAND (depository unknown).
marginatusf. invalida Priesner, 1928: 654. ? Syntypes d", AUSTRALIA: Queensland (SMF).
lacertinus f. infirma Priesner, 1928: 654. ? Syntypes cf , AUSTRALIA: Queensland (SMF).
terrigena Girault, 1928c: 2 ? Syntypes cf , AUSTRALIA: Queensland (QMB).
fasciatipennis Girault, 1930: 1. ? Syntypes (? cf , sex not stated), AUSTRALIA: Queensland (QMB).
kellyanus Bagnall, 1932: 518-9. Holotype $, AUSTRALIA: South Australia (BMNH).
LASIOTHRIPS Moulton
(Fig. 322)
Lasiothrips Moulton, 1968: 121. Type-species: Lasiothrips perplexus Moulton, by monotypy.
This monobasic genus is based on a single male specimen from Australia. The maxillary stylets
of this specimen are about one-third of the head width apart medially (Fig. 322), but since they
are extended beyond the mouth cone by at least 100 //,m it is possible that when at rest they might
lie close together. Moreover, the epimeral sutures of this specimen are apparently complete but
this could be an artefact due to coverslip pressure. If this pair of characters is disregarded then
the specimen resembles a male Megalothrips which lacks abdominal tubercles. (Zoogeographi-
cal objections to this suggestion could be waived in view of the description below of Megalothrips
andrei sp. n. from Malaya.) The pronotum of perplexus is short and wide; however, the lateral
wings of the pelta are not exceptionally slender. Until further specimens are collected the
relationships of Lasiothrips cannot be assessed. It cannot be distinguished satisfactorily from
Egchocephalothrips on present evidence, although monstrosus has the head more strongly
elevated in the mid-line with the two pairs of postocular setae arising side by side.
SPECIES INCLUDED
perplexus Moulton, 1968: 122-3. Holotype cf , AUSTRALIA: Queensland (CAS).
MEGALOTHRIPS Uzel
(Figs 318, 324, 329, 337)
Megalothrips Uzel, 1894: 224-5. Type-species: Megalothrips bonannii Uzel, by subsequent designation,
Bagnall, 1909a: 350.
This genus has been used for five species from the Holarctic Region, three North American and
two European. In contrast, andrei sp. n., described below, was collected at Kuala Lumpur in
Malaya. This extension in range is remarkable, but specimens of Megalothrips species have also
been studied in the collection of Dr Shuji Okajima (Tokyo) which were collected in Japan
(Kanagwa and Ohdaru) as well as in the Iriomote and Ishigaki Islands near Taiwan. Moreover,
one specimen similar to andrei has been seen from Sumatra. Two further genera which are
known only from single specimens, Lasiothrips from Australia and Egchocephalothrips from
New Caledonia, may also prove to be synonyms of Megalothrips eventually.
Megalothrips species have the maxillary stylets deeply retracted into the head and close
together medially (Fig. 318) as in Bacillothrips, but the pelta is characteristic with slender lateral
lobes (Fig. 329).
The tube length is sexually dimorphic, being shorter in males than females. Moreover, in
andrei sp. n. , and to a lesser extent in bonannii but not in the other species, the tube of the female
is longer than the head. The two European species bonannii and delmasi, together with schuhi
from Oregon, U.S.A., form a closely related species-group in which antennal segment III is
mainly yellow, IV and V have yellow pedicels, and even VI is pale basally. In contrast,
picticornis, from the western U.S.A., has only the basal 0-75 of segment III yellow and the
pedicel of IV slightly pale, whereas spinosus, which is widespread from Virginia to Washington
State, U.S. A. , has all the antennal segments dark. The new species, andrei from Malaya, differs
from picticornis in the greater length of the tube, and in having the basal 0-8 or more of antennal
segment III yellow but the pedicel of IV brown. Moreover, all the major setae on the body are
78 L. A. MOUND AND J. M. PALMER
dark brown or black instead of light brown to colourless. Apart from these relatively superficial
differences in colour and proportions the six species of Megalothrips are very similar to each
other in structure.
SPECIES INCLUDED
andreisp. n. Holotype cf , MALAYA (BMNH).
feonaiiiiiiUzel, 1895: 227-8. Holotype cf, CZECHOSLOVAKIA (depository unknown).
de/maaBournier, 1956: 163-9. Holotype cf , FRANCE (BCM; 2 $, 2 cf paratypes BMNH).
p/cticomisHood, 1927ft: 204. Lectotype $, U.S.A.: California (USNM).
animus Moulton, 1929c: 242^. Holotype cf , U.S.A.: California (CAS).
schuhi Crawford, 1947: 197-9. Holotype $, U.S.A.: Oregon (USNM).
spinosusHood, 1908ft: 306-7. Lectotype $, U.S.A.: Pennsylvania (USNM).
fuscus Watson, 1921: 84-5. Holotype $, U.S.A.: New York (FSAC).
Megalothrips andreisp. n.
(Figs 324, 329, 337)
Macropterous cf . Colour dark brown; antennal segment III pale, yellow, slightly darker in apical fifth;
wings pale with pale brown median line in basal half. Head 2-3 times as long as broad; interocellar setae
long, stout pair of setae on cheeks immediately behind eyes; postocular setae pair II long; maxillary stylets
close together in centre of head, retracted to posterior margin of eyes. Pronotum short, about 3 times as
broad as long; epimeral sutures weakly developed. Median metanotal setae well developed, longer than
the distance between their bases. Wings with 3 sub-basal setae. Pelta with lateral lobes narrowly joined to
median lobe (Fig. 329). Abdominal tergites II to VI with 2 pairs of well-developed sigmoid wing-retaining
setae ; tergite VI with a pair of tubercles laterally ; setae B\ on abdominal tergite IX about 0-5 times length of
tube. Tube 3-6 times as long as broad, shorter than head, tapering more strongly at apex, set with stout dark
setae (Fig. 324).
Measurements (holotype cf in /urn). Body length 3744. Head, length 575; maximum breadth 248;
interocellar setae 116/120; postocellar setae 42/46; postocular setae pair I 88/92; postocular setae pair II
164/241. Pronotum, length 124; width 364; major setae, am 116/46, aa 40/38, ml 97/76, pa 161/152, epim
184/216. Median metanotal setae 142/151. Wings, length 1630; maximum width 143; number of duplicated
cilia 28/30. Tergite IX setae Bl 284/272. Tube, length 476; maximum width 119. Antennal segments
III- VIII length, 152/157; 124/128; 128/133; 102/112; 69/71; 67/66.
Macropterous $ . Colour and structure similar to cf . Tergite VII with 2 pairs wing- retaining setae. Tube
longer, about 5 times as long as broad and longer than head.
Measurements (2 $ paratypes in /u,m). Body length 4568 (5010). Head, length 632 (633); maximum
width 296 (287); interocellar setae 135/140 (148/152); postocellar setae 56/48 (55/42); postocular setae pair I
67 (64/67); postocular setae pair II 135/137 (160/156). Pronotum, length 124 (133); maximum width 448
(422); major setae, am 38/43 (42), aa 31/34 (39), ml 53/54 (35/44), pa 1 13/143 (130/144), epim 171 (180/182).
Median metanotal setae 103/128 (117/124). Wings, length 1727 (1872); maximum width 134 (172); number
of duplicated cilia 27/30 (29/31). Tergite IX setae Bl 301/312 (315/332). Tube, length 729 (758); maximum
width 147 (162). Antennal segments III-VIII length 176/181 (180/182); 143/147 (152); 147/152 (152/157);
114 (114/119); 62/64 (66); 71 (70/71).
SPECIES STUDIED
Holotype cf , Malaya: Kuala Lumpur, on dead branches, 24.xii.1969 (R. G. & Floyd Andre) (BMNH).
Paratypes. 1 9 taken with holotype; 1 $ similar data except 29.xii.1969 (BMNH).
COMMENT. The specimen from Sumatra referred to above is much larger (head length 900 /am;
tube length 1150 /am).
MEGA THRIPS Targioni-Tozzetti
(Figs 321, 326, 331)
Megathrips Targioni-Tozzetti, 1881: 124-5. Type-species: Megathrips piccioli Targioni-Tozzetti (now
regarded as a synonym of Phloeothrips lativentris Heeger), by monotypy.
Siphonothrips Buffa, 1908: 389. Type-species: Siphonothrips elegans Buffa, by monotypy. Syn. n.
This genus, which is used here for six nominal species, appears to be a Holarctic derivative of the
large genus Bactrothrips from the Old World tropics. The species in the two genera are
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 79
essentially similar, and can be distinguished only by the slightly larger head and more deeply
retracted maxillary stylets of Bactrothrips .
The type-species, lativentris, is highly successful and widespread, but probably evolved from
Bactrothrips through an earlier phase of wing-reduction. The fore wings are pale; the anterior
pair of wing-retaining setae on each tergite is reduced (Fig. 326); the pelta has the lateral lobes
relatively broad rather than slender (Fig. 331); the head is foreshortened, less than 2-5 times as
long as wide (Fig. 321). This constellation of characters suggests that lativentris has evolved from
a micropterous or apterous ancestor, but has redeveloped the fully winged condition without
redeveloping all of the characters associated with macroptery.
Five other species are placed in Megathrips, but of these both elegans and timidus are known
only from their original descriptions. Moreover, brevis is known only from a single damaged
male which has very short straight lateral tubercles on the sixth tergite and is smaller than any
known male of lativentris. M. flavipes is the only member of the genus recorded from Pinus leaf
litter. This species closely resembles the description of timidus in colour, having completely pale
femora, but differs in the tube being shorter than the head. Finally, inermis is particularly
interesting because, like Bactrothrips pitkini sp. n. described above, it lacks all trace in both
sexes of the lateral abdominal tubercles usually regarded as diagnostic of this group of genera.
The genus Siphonothrips is here placed as a synonym of Megathrips because there is nothing in
the description which can be used to distinguish between these taxa. The original figure of
elegans is apparently of a dry, carded specimen. The head appears relatively broad and the
abdomen shrunken, thus making the length of the tube and shape of the lateral tubercles as
illustrated unreliable. Unfortunately, the original descriptions and illustrations of both picciolli
and lativentris are also equivocal, and the concept of the type-species of Megathrips owes much
to convention. The oldest available unequivocal name for this concept is longispina Reuter, and
to ensure stability of the generic name it may become necessary to apply to the International
Commission on Zoological Nomenclature to have longispina declared the type-species of
Megathrips.
SPECIES INCLUDED
5rev/s (Bagnall, 19146: 291-2) (Siphonothrips). Holotype cT, YUGOSLAVIA (BMNH).
*e/egans(Buffa, 1908: 389-90) (Siphonothrips). Holotype cf, SARDINIA (depository unknown). Comb. n.
flav/pes (Reuter, 1901: 216) (Cryptothrips). Syntypes $, CRETE (depository unknown).
inermis Priesner, 1937a: 348-50. Holotype cf , SARDINIA (DEI).
lativentris (Heeger, 18526: 479) (Phloeothrips) . Syntypes cf , AUSTRIA (depository unknown).
longispina Reuter, 1879: 214-5 (Phloeothrips). Holotype cf , SWEDEN (depository unknown).
tibialis Reuter, 1879: 215-6 (Phloeothrips). Holotype $, SWEDEN (depository unknown).
piccioli Targioni-Tozzetti, 1881: 124-5. Syntypes $, ITALY (depository unknown).
niger Schmutz, 1909: 346-7 (Megalothrips). Holotype $, RUMANIA: Herkulesbad (depository un-
known).
padewiethi Karny, 1919: 114-5 (Bacillothrips) . Syntypes cf $, YUGOSLAVIA: 'Kroatischen Littorale'
(?SMF).
* timidus Cott, 1956: 177-9. Holotype $, U.S.A.: California (depository unknown).
MEIOTHRIPS Priesner
(Figs 313, 323, 327, 336)
Idolothrips (Meiothrips) Priesner, 19296: 197. Type-species: Idolothrips (Meiothrips) annulatus Priesner
(now regarded as a synonym of Acanthinothrips annulipes Bagnall), by monotypy.
Meiothrips Priesner; Bagnall, 1934: 494. [Raised to genus.]
Meiothrips (Aculeathrips) Kudo, 1975: 421. Type-species: Meiothrips (Telothrips) nepalensis Kudo &
Ananthakrishnan, by monotypy. [Replacement name for Meiothrips (Telothrips) Kudo & Ananthak-
rishnan, 1974: 385 nee Telothrips Priesner, 19290.] [Synonymised by Palmer & Mound, 1978.]
This genus, which is used for three species found between Borneo, Thailand and India, has been
redefined by Palmer & Mound (1978). Each of these species could be placed in a separate
monobasic genus , as also could the two species of Idolothrips , if the generic concepts traditional-
80 L. A. MOUND AND J. M. PALMER
ly employed in the Bactrothrips complex were accepted. However, the Meiothrips species are
intermediate between Idolothrips and Bactrothrips both structurally and zoogeographically, and
a series of monobasic genera would obscure this important relationship. Palmer & Mound
(1978: 212) refer to a small male of nepalensis from Thailand with the metanotal setae short as in
Idolothrips. These authors also point out that the ornamentation of the tube in the males of M.
nepalensis and /. dissimilis is very similar, and that the females in these genera are difficult to
separate from some females of Bactrothrips. In all three genera the anapleural sutures are short
and straight.
SPECIES INCLUDED
annulipes(B agnail, 1914c: 378-9) (Acanthinothrips) . Lectotype cf , SARAWAK (BMNH).
annulatus Priesner, 19296: 197-201 (Idolothrips subgen. Meiothrips). Syntypes cf 9 , SUMATRA (SMF;
1 Cf BMNH).
meno/i/Ananthakrishnan, 1964a: 99-101. Holotype 9> INDIA (TNA).
nepalensis Kudo & Ananthakrishnan, 1974: 385-7. Syntypes cf $, NEPAL (TNA, 2 cf BMNH).
Genera of Hystricothripina
Most of the genera placed in this subtribe in the present paper were listed by Priesner (1961) in
the Idolothripina. However, as interpreted here, the Idolothripina comprises a group of
predominantly old world genera characterised by the presence of well-developed praepectal
plates, two pairs (or more) of tergal wing-retaining setae, well-developed forewing duplicated
cilia, and a tendency for the males to bear one or more pairs of elongate tubercles or drepanae
laterally on the abdomen. In contrast, the Hystricothripina (= Zeugmatothripina) comprises a
group of predominantly new world genera characterised by the praepectal plates being absent or
very weakly developed, the tergites usually bear only one pair of wing-retaining setae, the
forewing duplicated cilia are absent or weakly developed, and there are no lateral drepanae on
the male abdomen (although the posterolateral tergal setae usually arise on tubercles in both
sexes).
Thirteen genera are recognised here in the Hystricothripina, of which eight, involving 33
species out of a total of 42, are found only in the Neotropics. These Neotropical genera appear to
fall into two major groupings: the Actinothrips-group (including Hybridothrips and Zacti-
nothrips) of 14 species, and the Zeugmatothrips-group (including Azeugmatothrips,
Cyphothrips, Saurothrips and Zeuglothrips) of 19 species. Stannard (1954: 72) included these
genera as subgenera of Actinothrips , and it may be that there are too many genera for the
number of species involved. This phenomenon is not unusual when species differ from each
other in very obvious, rather than somewhat subtle, characters. One of our colleagues has
written to say that he would classify the group mainly on the number of elongate setae on the
dorsal surface of the head. However, in this group there are three pairs of dorsal setal-bases -
postocellars, postoculars and mid dorsals. Each of these may bear long or short setae, and,
moreover, their position is also variable. In the opinion of the present authors, the size and
position of these setae, in this group, are so variable that they are probably under relatively
simple genetic control, and are poor indicators of phyletic relationships. This suggestion may
apply also to the foretarsal tooth (absent in females) and forewing duplicated cilia in this group,
as these characters are developed in a variety of different combinations in different species with
no apparent evolutionary trend.
The five genera of Hystricothripina found outside the Neotropics are more diverse. Hystri-
cothrips, with two species from western Africa, is most closely related to the Neotropical genera,
whereas Atractothrips , with two species from Florida and Mexico, is similar to the Oriental
genera in having a pair of stout pre-ocellar setae. These Oriental genera, Holurothrips , with
three species, and the two monobasic genera described below, Neatractothrips and Paracti-
nothrips, constitute a unique group in which the mesopraesternum is transverse, parallel-sided
and apparently continuous with the sclerites laterally. In contrast, the rest of the Hystricothri-
pina are remarkable in having the mesopraesternum reduced to a small median sclerite, whereas
the rest of the Phlaeothripidae have a boat-shaped mesopraesternum which is frequently
reduced to two small lateral triangles.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 81
ACTINOTHRIPS Bagnall
(Figs 345, 353)
Actinothrips Bagnall, 19Q9d: 332-333. Type-species: Actinothrips longicornis Bagnall, by monotypy.
Dasythrips Hood, 1937c: 521-522. Type-species: Dasythrips regalis Hood, by monotypy. Syn. n.
The type-species of this genus, longicornis Bagnall, was described from a single dry specimen.
Mound (1968) stated that only the tube of this specimen survived, mounted on a microscope
slide, but subsequently the rest of the specimen was discovered dry in a glass vial at the British
Museum (Natural History). This holotype, however, is a male, not a female, as can be deduced
from the original illustration by Bagnall of the short, stout setae on tergites VII and VIII.
Moreover, there is a lapsus in the original description, because it is the meso- and metasterna, not
the 'meso and metascutum' which are 'rather closely set with numerous short hairs'.
Hood was never able to study the type-species of Actinothrips, and in fact longicornis is very
similar to a major male of Dasythrips regalis in having a pair of forwardly directed tubercles on
tergite III and a pair of stout metanotal setae arising from tubercles, as well as densely hairy
thoracic sternites. Moreover, a large paratype of Actinothrips femoralis bears as many setae
ventrally as a small paratype of Dasythrips fraterculus. For these reasons Dasythrips is here
regarded as a synonym of Actinothrips . Moreover, the description of D. chiapensis from Mexico
does not distinguish this species satisfactorily from the female of A. trichaetus from Panama.
Actinothrips is mostly closely related to Zactinothrips , although both in that genus and in
Hybridothrips the eighth abdominal segment is elongate, twice as long as wide. Duplicated cilia
are present on the fore wing in species of all three genera; however, in only three species of
Actinothrips does the male bear a foretarsal tooth: femoralis, gargantua and pedalis. This
presence of a foretarsal tooth in these species is remarkable because femoralis and polychaetus
appear to be closely related in having numerous stout setae on the inner margin of the male
forefemora, although polychaetus lacks a foretarsal tooth. Moreover, females of pedalis are
similar to females of bondari.
Allometry and sexual dimorphism in Actinothrips species, together with the few specimens
available for study, make species delineations unsatisfactory at present. One species-group
includes six nominal species which are distinguished mainly on characters associated with size.
In decreasing order of body size these are: regalis (Peru), longicornis (Venezuela), fraterculus
(Peru), chiapensis (Mexico), trichaetus (Panama, Trinidad, Ecuador), bondari (Brazil). The
first five species have three major setae in an obliquely transverse row near the apex of the
forefemora in both sexes. However, in bondari there is only one large seta, on the inner apical
margin, and two smaller setae dorsally and externally. The apices of the tibiae are dark in the
larger species, paler in bondari, but yellow in pedalis. Despite the similarity between the females
of these last two species the males are readily distinguished by the bulbous femora of pedalis, an
unusual characteristic in Actinothrips . Finally, the females oimonochaetus, although individual-
ly larger than bondari females, have a single major seta apically on the forefemora, but the males
are unusual in having all three pairs of posteroangular setae on tergite VIII short and stout.
SPECIES INCLUDED
fcoiM/ar/Hood, 1928: 147-150. Holotype 9, BRAZIL (USNM).
*e/iiapens/s(Johansen & Garcia, 1976: 235-241) (Dasythrips). Holotype $, MEXICO (UNAM). Comb. n.
femoralis Hood, 1950: 5-9. Holotype $, BRAZIL (USNM; O" 9 paratypes BMNH).
fraterculus (Hood, 1941: 236-240) (Dasythrips). Holotype d", PERU (USNM; C? 9 paratypes BMNH).
Comb. n.
gargantua Santis, 1960: 57-60. Holotype cf , BRAZIL (MLPA).
longicornis Bagnall, 1909d: 333-334. Holotype cf , VENEZUELA (BMNH).
monochaetusHood, 1935c: 252-254. Holotype 9 , GUYANA (BMNH).
pedatfsHood, 1949: 76-78. Holotype 9, BRAZIL (USNM; cf 9 paratypes BMNH).
* polychaetus Hood, 1941: 221-223. Holotype C?, VENEZUELA (USNM).
regatfs(Hood, 1937c: 522-527) (Dasythrips). Holotype cf , PERU (USNM). Comb. n.
trichaetusHood, 1935c: 248-252. Lectotype 9, PANAMA (USNM; cf 9 paralectotypes BMNH).
82 L. A. MOUND AND J. M. PALMER
ATRACTOTHRIPS Hood
(Fig 346)
Atractothrips Hood, 1938a: 27-28. Type-species: Atractothrips bradleyi Hood, by monotypy.
Two Oriental species described in this genus are now placed in Neatractothrips and Malesiathrips
q.v. The two remaining species are found in Mexico, and in Florida and the extreme south-east
of Georgia, U.S.A. They have a combination of characters which are intermediate between
those of Neotropical and Oriental Hystricothripina. Antennal segments I and II bear large
dorsal setae, moreover III is shorter than IV as in Zeugmatothrips , but in contrast VIII is short
and stout instead of lanceolate. The head has stout cheek setae, and the maxillary stylets are
wide apart (Fig. 346), but the preocellar setae are enlarged as in the Oriental members of the
group. The praepectus and mesopraesternum are absent, and the mesothoracic epimeral setae
well developed, but the metanotal setae and fore wing sub-basal setae are very small in bradleyi
(although longer in mockfordi} , and the lateral setae of tergite I arise anterior to the lateral lobes
of the pelta. The abdominal tergites bear two pairs of wing-retaining setae, although the anterior
pair is straight, not sigmoidal, and the tergal posterolateral angles are produced into two pairs of
tubercles. Unlike the Neotropical species, the tube is long with the lateral setae short and sparse.
Atractothrips appears to represent the sister-group of the three Oriental genera Holurothrips,
Neatractothrips and Paractinothrips.
SPECIES INCLUDED
ferad7ej/Hood, 1938a: 28-32. Holotype cf , U.S.A.: Florida (USNM; cf $ paratypes BMNH).
"mocWord/Stannard, 1974: 45-8. Holotype $, MEXICO (INHS).
AZEUGMATOTHRIPS gen. n.
(Figs 349, 357, 374, 382)
Type-species: Azeugmatothrips rectus sp. n.
Antennal segments III-IV with 2 stout dorsal setae, I and V with one stout dorsal seta, III shorter than IV.
Head with 3 pairs of major dorsal setae (postocellars elongate); maxillary stylets wide apart. Pro-, meso-
and metanota similar to Zeugmatothrips; foretarsus of cf with a stout tooth; forewing with duplicated cilia.
Pelta bearing two pairs of setae laterally; tergites with one pair of wing-retaining setae; tube with lateral
setae long and erect.
The two species in this new genus appear to be derived from Zeugmatothrips. The antennae,
with segment III short and VIII lanceolate, also the pelta, are similar to that genus. Moreover,
the mesothoracic epimeral setae and the mesonotal lateral setae are elongate as in Saurothrips,
and the postocellar setae are elongate as in Saurothrips, Zeuglothrips and Hybridothrips .
However, this combination of characters, together with the forewing bearing duplicated cilia,
and the male with a f oretarsal tooth , is found only in the two species treated below . One of these ,
obrieni, was described in Zeuglothrips because of the similar head chaetotaxy although the
type-species of that genus has the maxillary stylets very long and close together medially
(Fig. 342). The new species, rectus, differs from obrieni in its much smaller size, in having two
large setae on antennal segment II instead of only one, and in having four stout setae on each
forefemora instead of about eight such setae.
SPECIES INCLUDED
oftr/ejiJ(Johansen, 1975: 188-92) (Zeuglothrips). Holotype cf , PANAMA (UNAM). Comb. n.
rectos sp. n. Holotype cf , TRINIDAD (BMNH).
Azeugmatothrips rectus sp. n.
Macropterous cf . Colour dark brown to black, mid- and hindtarsi also foretibiae paler; foretarsi and apices
of foretibiae yellowish brown; antennal segment IV, V and stem of VI yellow, III with club pale but stem
brown except for extreme base; dorsal setae on antennae, fore- and midfemora, also tergites VIII-IX, dark
brown; major setae on vertex, pronotum and tergites II-VI pale or colourless; setae on tergite VII, also
genal setae, shaded; forewing weakly shaded, with a median longitudinal dark line, sub-basal setae pale.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 83
Head (Fig. 349) produced in front of rounded eyes; postocellar setae elongate, mid-dorsal setae arising
between postoculars; cheeks with two pairs of stout setae; maxillary stylets wide apart; dorsal surface
weakly sculptured with anastomosing lines. Antennae 8-segmented, VIII lanceolate, II-IV with 2 stout
dorsal setae. I and V with one stout seta (Fig. 357); 2 sense cones on III, 4 on IV.
Pronotum (Fig. 349) transverse, aa and ml setae close together on a tubercle; epimeral sutures complete;
pm setae minute ; praepectus absent , mesopraesternum reduced to a small median sclerite . Forefemur with
4 stout setae; foretarsus with stout recurved tooth. Mesonotal mid-lateral and epimeral setae well
developed. Metanotum reticulate, median setae stout. Metaepimeron moderately swollen, with one large
seta. Forewing parallel-sided, curving forward distally, sub-basal setae well developed.
Pelta (Fig. 382) broad basally, bearing 2 pairs of setae; tergites II- VII with one pair of wing-retaining
setae; lateral tergal setae well developed, II with 2 pairs, III with 3 pairs. Tube densely setose, setae long
and erect (Fig. 374). Sternites with one row of discal setae, posteromarginal setae small.
Measurements (holotype cf in /am). Body length 4000 (extended). Head, total length 390; length in front
of eyes 60; width behind eyes 230; major setae - postocellars 150, postoculars 190, mid-dorsals 110, genals
45-60. Pronotum, length 200; width 360; major setae - am 120, aa 180, ml 135, epim 165, pa 120. Mesonotal
setae - midlaterals 60, epimerals 90. Metanotal median setae 130. Forewing, length 1700; distal width 90;
sub-basal setae 120, 100, 140; number of duplicated cilia 9 (12). Tergite VI posteroangular setae B\ 260, B2
180, 53 230. Tergite IX setae, Bl 180, B2 220, B3 180. Tube, length 900; terminal setae 200; longest lateral
setae 260. Antennal segments III-VIII length, 155, 180, 190, 130, 80, 110.
SPECIMENS STUDIED
Holotype O", Trinidad: Arima Valley, Simla, dead branch of Anona, 4.xi.l970 (L. A. Mound, 908).
COMMENTS. This new species is unique in having the mid-dorsal head setae arising so far forward,
in line with the postocular setae.
CYPHOTHRIPS Hood
(Figs 339, 352, 363, 384)
Cyphothrips Hood, 1952: 172. Type-species: Cyphothrips dorsalis Hood, by monotypy.
This monobasic genus appears to be a specialised derivative of Zeugmatothrips . The head,
pronotum (Fig. 339), sternites and pelta (Fig. 384) are similar to species of that genus, although
the ventro-lateral pale tubercles on the head are similar to those found in Zactinothrips q.v. The
first antennal segment bears a stout dorsal seta, but segments II-V do not have any stout setae
(Fig. 363), although this is also true of Zeugmatothrips hoodi. The main differences from
Zeugmatothrips species are that the fore wing bears 2 to 4 weak duplicated cilia, and the lateral
setae on the tube are weak and decumbent. Moreover, in the male, the foretarsus bears a tooth,
and the metanotal median setae are borne on a pair of elongate tubercles (Fig. 352).
SPECIES INCLUDED
dorsatfsHood, 1952: 172-173. Lectotype $, BRAZIL (USNM).
HOLUROTHRIPS Bagnall
(Figs 341, 354, 361, 383)
Holurothrips Bagnall, 1914c: 376. Type-species: Holurothrips ornatus Bagnall, by monotypy.
This genus, with three species between Japan, Malaya and Queensland, is placed in the
Hystricothripina because of the following characters: antennal segment VIII lanceolate,
segments I-II with stout dorsal setae (Fig. 361); head with 2 pairs of stout cheek setae; pronotal
aa and ml setae arising close together (Fig. 341); mesothoracic epimeral setae present, although
small; tergal posteroangular setae arising from small tubercles; tergite IX setae short; tube long
with many short, widely spaced setae. Holurothrips resembles the Oriental genera Neatrac-
tothrips and Paractinothrips , and differs from the Neotropical genera in having praepectal
plates, preocellar setae, a completely transverse mesopraesternum and reduced metanotal
setae. In this genus the setae of the first abdominal segment arise close to the lateral extremities
of the pelta (Fig. 383), the thoracic sternites bear numerous setae, and the abdominal sternites
have two rows of discal setae. The most remarkable feature of the genus is the ventral
84 L. A. MOUND AND J. M. PALMER
prolongation of the eyes (Fig. 341). Contrary to Mound (1974: 57) tergites IV-VI (sometimes
also III) have three, not two, pairs of wing-retaining setae in ornatus. However, the closely
related species morikawai has only two pairs of these setae.
SPECIES INCLUDED
collessi Mound, 1974: 58. Holotype $, AUSTRALIA: Queensland (ANIC).
morikawai Kurosawa, 1968: 55. Holotype $, JAPAN (MAT),
ornafus Bagnall, 1914c: 376-377. Lectotype cf , SARAWAK (BMNH).
leeuweni Priesner, 1934: 62-63. Syntypes cf $, JAVA (SMF; BMNH).
HYBRIDOTHRIPS Stannard
(Figs 340, 360)
Actinothrips (Hybridothrips) Stannard, 1954ft: 71-74. Type-species: Actinothrips (Hybridothrips) oneillae
Stannard, by monotypy.
Hybridothrips Stannard, 1957: 100-101. [Raised to genus.]
This genus appears to be derived, with Zactinothrips q.v., from Actinothrips. The heads are
similar in the latter two genera, but the head shape of Hybridothrips is distinctive amongst
Neotropical species in that the eyes are a little reduced and flattened (Fig. 340), much as in
Atractothrips . There are three pairs of major dorsal setae on the head, as in Zeuglothrips, but the
maxillary stylets are wide apart, the pelta does not bear setae laterally, and the pronotal pa setae
are very small but the pm setae enlarged. The pronotum of Zactinothrips is intermediate in that
both pa and pm setae are very small; however, the males of Hybridothrips and Zactinothrips are
similar in having abdominal segment VIII much larger than in Actinothrips. Antennal segments
III-IV bear supernumerary sense cones ventrally near the apex in Hybridothrips (Fig. 360), but
both dorsally and ventrally in Zactinothrips, at least in the males. The holotype female of
guerreronsis has been compared with the holotype male of oneillae and they are regarded as the
same species. According to Dr Roberto Johansen (in litt.) this species is widespread in the
Quercusl Pinus forests of Mexico at the transitional zone between the Neotropics and the
Nearctic.
SPECIES INCLUDED
oneillae (Stannard, 1954ft: 74) (Actinothrips subgenus Hybridothrips}. Holotype cf , MEXICO (USNM).
guerreronsis Johansen & Garcia, 1973: 55-61 (Zeuglothrips). Holotype $, MEXICO (UNAM). Syn. n.
HYSTRICOTHRIPS Karny
(Figs 344, 358, 370, 375)
Hystricothrips Karny, 1912c: 132. Type-species: Hystricothrips phasgonura Karny, by monotypy.
Zeugmatothripoides Bagnall, 1929: 71-72. Type-species: Zeugmatothripoides africanus Bagnall, by
monotypy. [Synonymised by Mound, 1968: 124-125.]
This genus, from western Africa, appears to be the sister-group of the Neotropical Hystricothri-
pina. It differs mainly in the heavily sculptured, and densely setose tube, and the lack of elongate
setae on the metanotum and mesothoracic epimera. The forewings, when present, bear up to
about 25 duplicated cilia, although these are very fine. In the male the foretarsus bears a stout
tooth, and the posterolateral angles of the tergites are drawn out into tubercles (Fig. 370).
Antennal segments III-VIII (Fig. 358) are similar to those of Zeugmatothrips species, the
mesopraesternum is similarly reduced, but the postocellar setae are well developed (Fig. 344)
and there are two pairs of wing-retaining setae in macropterae.
Contrary to Pitkin & Mound (1973) africanus can be distinguished from phasgonura as
follows.
1 Antennal segment I with inner dorsal seta half as long as external dorsal seta ; segment II with one dorsal
seta more than half as long as segment III; cf with seta BI on tergite VII stout but rounded apically,
setae BI and B2 on VIII short and thorn-like [Sierre Leone, Ivory Coast, Nigeria, Sao Thome]
africanus
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 85
- Antennal segment I with both dorsal setae extending beyond apex of II; segment II with no long dorsal
setae; cf with Bl on tergite VII thorn-like on a stout tubercle, but B1 and B2 on VIII setaceous (Zaire,
Equatorial Guinea, Angola) phasgonura
SPECIES INCLUDED
a/r/ca7iiis(Bagnall, 1929: 72-73) (Zeugmatothripoid.es). Holotype $, SIERRA LEONE (BMNH).
phasgonura Karny, 1912c: 132-133. Holotype $, EQUATORIAL GUINEA (ZMB).
hystrix Priesner, 1932: 198-201. Holotype cf , ZAIRE (MRAC).
NEATRACTOTHRIPSgen. n.
(Figs 372, 377)
Type-species: Atmctothrips macrums Okajima.
Antennae 8-segmented, I and II with dorsal setae enlarged, III longer than IV but without stout setae.
Head elongate, prolonged in front of eyes with stout preocellar setae and cheek setae; maxillary stylets
wide apart. Pronotum with epimeral sutures not quite complete; aa and ml setae close; praepectus present
but reduced to a pair of small, setal bearing sclerites laterally. Foretarsus in both sexes without a tooth but
with inner margin slightly raised; femora with several large setae. Mesothoracic epimeral setae well
developed; mesopraesternum transverse, not clearly delimited from lateral sclerites (Fig. 372). Metanotal
setae not elongate; metathoracic episterna moderately enlarged, anapleural sutures short. Forewings
broad, without duplicated cilia. Lateral setae of abdominal segment I anterior to lateral lobes of pelta
(Fig. 377); tergites III-V with 5 pairs of wing-retaining setae; tergites produced into 2 pairs of tubercles
posterolaterally; setae on tergite IX very short; tube exceptionally long, marginal setae short and sparse;
sternites with more than one row of discal setae.
This new monobasic Oriental genus is closely related to Paractinothrips, and, together with that
genus and Holurothrips , constitutes the sister-group of Atractothrips . The holotype has an
ill-defined foretarsal tooth, but this is not present in 14 $ , 4 cf collected in the Philippines with
the paratypes of Paractinothrips peratus q.v. The unusual structure of the mesopraesternum
(Fig. 372) is similar to that found in Holurothrips and Paractinothrips, but quite different from
that found in Atractothrips and the rest of the Hystricothripina. The three Oriental genera tend
to have the tube longer and more sparsely setose, the tergites with more numerous wing-
retaining setae, and the head with larger preocellar setae than their Neotropical counterparts.
The original description oimacrurus states that the praepectus is absent. However, examina-
tion of the ventral surface of the holotype through the back of the slide mount, using a x40 water
immersion lens, has revealed the presence of two seta-bearing sclerites anterolateral to the
probasisternum (Fig. 372). These sclerites, which are also present in the specimens from the
Philippines, are here interpreted as praepectal plates because they appear to be homologous
with the external parts of the praepectus of Paractinothrips peratus (Fig. 373), and because
cervical sclerites do not usually bear setae.
SPECIES INCLUDED
macrurus (Okajima, 1975: 13-16) (Atractothrips). Holotype $, RYUKU Is. (OCT). Comb. n.
PARACTINOTHRIPS gen. n.
(Figs 347, 348, 355, 359, 369, 373, 378)
Type-species: Paractinothrips peratus sp. n.
Head with preocular projection and well-developed preocellar setae; postocellar, postocular and mid-
dorsal setae elongate; cheek setae stout; maxillary stylets wide apart. Antennae 8-segmented; III shorter
than IV; VIII slender; I with an elongate dorsal seta. Pronotal aa and ml setae fairly close; epimeral sutures
not complete; praepectus present but weak. Both sexes without a foretarsal tooth; femora with several
stout setae (Fig. 348). Mesopraesternum transverse; metanotal median setae not elongate; metathoracic
episterna swollen, with a series of setae (Fig. 355); forewings slender, without duplicated cilia. Pelta broad
with setae of abdominal segment I anterior to lateral lobes; tergites III- VII with 3 or 4 pairs of fan-shaped
wing-retaining setae; posteroangular tergal setae arising from 2 pairs of tubercles; tergite IX setae short;
tube elongate with many fine, semi-erect setae; sternites with more than one row of discal setae.
86 L. A. MOUND AND J. M. PALMER
This new monobasic Oriental genus belongs in the Hystricothripina because of the form of the
antennae and head (Figs 347, 359), the proximity of the pronotal aa and ml setae, the enlarged
metathoracic anepisterna, the posterolateral tergal tubercles (Fig. 369),, and the long setose
tube. The praepectus, which is absent in the Neotropical genera, is more fully developed than in
Neatractothrips, although these two genera are similar in the structure of the mesopraesternum.
P. peratus differs from N. macrurus most obviously in the form of the antennae with a short third
segment, and in the slender wings.
Paractinothrips peratus sp. n.
Macropterous O". Colour brown with red internal pigment, tarsi paler; antennal segments III- VI and basal
half of VII yellow; also apices of all tibiae yellow; major setae pale except on head and antennal segment I;
small dorsal setae on antennal segments III-V dark brown; wings slightly shaded, with a very dark
longitudinal median line.
Head prolonged in front of eyes; dorsal surface sculptured, bearing 2 rows of stout setae and 3 pairs of
long setae (Fig. 347); maxillary stylets wide apart. Antennae 8-segmented, I with dorsal seta extending to
apex of II; sense cones long and slender, 2 on III, 4 on IV; segment III shorter than IV, dorsal setae small,
dark, but with apices expanded (Fig. 359).
Pronotum with complex sculpture; major setae, particularly epimerals, on tubercles (Fig. 347);
praepectus weakly sclerotised (Fig. 373). Forefemur with at least 6 stout setae on tubercles, 1 or 2 of which
are on inner margin (Fig. 348); foretarsus without a tooth. Mesonotum with 2 pairs of stout setae, similar in
form to mesothoracic epimeral and metanotal median setae. Forewing with cilia arising unusually distant
from wing margin.
Pelta with lateral wings, tergite I setae stout (Fig. 378); tergites with wing- retaining setae enlarged and
fan-shaped; posterolateral tergal angles with 4 setae arising close together (Fig. 369); tergite IX setae very
short, pair B\ close together dorsally; tube long, apex constricted, lateral setae numerous and emerging at
an angle of about 30°. Sternites IV-VI each with about 16 scattered discal setae, marginal setae not
elongate.
Measurements (holotype C? in /am). Body length 4800 (extended). Head, total length 420; length of
preocular process 60; maximum width behind eyes 210; dorsal setae - preocellar 30, postocellar 120,
postocular 90, mid-dorsal 65, cheek setae 30. Pronotum, length 210; median width 300; major setae - am 30,
aa 60, ml 40, epim 110, pa 60, pm 30. Mesothoracic epimeral seta 40. Forewing, length 1800; distal width
75; sub-basal setae 50, 45, 45. Metanotal median setae 40. Tergite V posteroangular setae B\ 110, B2 90, B3
60. Tergite IX setae BI 50, B2 50. Tube, length 1350; longest lateral setae 65; terminal setae 170. Antennal
segments I- VIII length, 60 (seta 80), 50, 105, 170, 170, 115, 70, 40.
Macropterous $. With no significant difference from cT.
SPECIMENS STUDIED
Holotype d" , Malaya: Kuala Lumpur, on dead palm leaves, 29.xii. 1969 (R. G. & Floyd Andre) (BMNH).
Paratypes. Malaya: 1 $ collected with holotype; 1 cf with similar data except 27.xii.1969; 1 $,
Buklanyan, on dead branches, 26.xii.1971 (Floyd Andre) (BMNH). Philippines: Luzon, Quezon National
Park, near Lucena City, 13 $, 18 cf on dead leaves of wild Palmaceae, 16.viii.1979 (5. Okajima) (BMNH
& OCT).
COMMENTS. The paratypes from the Philippines have the tibiae and antennal segment VI darker
than the Malayan specimens. Moreover, the ratio antennal segment III/IV varies from 0-66-0-89
apparently independently of both sex and body size.
SA UROTHRIPS Hood
(Figs 343, 356, 368, 381)
Saurothrips Hood, 1952c: 171. Type-species: Saurothrips assai Hood, by monotypy.
The only species in this genus appears to be a specialised, long-bodied, derivative of Zeugma-
tothrips. However, the eighth abdominal segment is not elongate and narrowed medially as in
Zactinothrips and to a lesser extent Actinothrips . S. assai has two stout setae on antennal
segment II (Fig. 356), as well as the stout setae on segments I and III-V which are found in
Zeugmatothrips. Moreover, the lateral setae on the tube are erect although rather short, and the
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 87
forewing bears no duplicated cilia. However, there is a stout foretarsal tooth in the male, and the
postocellar setae are elongate (Fig. 343) as in Zeuglothrips, Hybridothrips, Hystricothrips and
Azeugmatothrips . Unlike the other Neotropical Hystricothripina the metanotal setae and the
forewing sub-basal setae are unusually short (50 /mi), and tergites III- VI bear a second pair of
wing retaining setae near the antecostal ridge.
SPECIES INCLUDED
assafHood, 1952c: 171-172. Lectotype $, BRAZIL (USNM).
ZACTINOTHRIPS Hood
(Fig. 362)
Zactinothrips Hood, 1936d: 446-447. Type-species: Zactinothrips elegans Hood, by monotypy.
The most remarkable feature of this genus, the numerous small sense cones near the apex of
antennal segments III and IV (Fig. 362), is also found in Hybridothrips, although in the latter
genus these sense cones are only developed ventrally even in males. Moreover, in both these
genera the forewings bear duplicated cilia and the male has a foretarsal tooth. However, the
head of Zactinothrips resembles that of Actinothrips species, whereas the head of Hybridothrips
has more than one pair of stout setae on the vertex as in the Zeugmatothrips group of genera.
Despite the differences in the head chaetotaxy Zactinothrips from Peru and Hybridothrips from
Mexico are probably sister-groups, and together constitute the sister-group of Actinothrips .
The number of small additional, antennal sense cones is sex-linked in Zactinothrips, but may
also be subject to allometry and/or variation between populations. The available samples are not
sufficient to provide firm evidence. In large males these sense cones are present on segments III,
IV and V, although they are most numerous ventrally and there are fewer on V than on III.
Small females may have no additional sense cones on the dorsal surface of IV and V, and even
ventrally there are less than half as many as are found in males. The ventrolateral tubercles on
the head, referred to by Hood, are also found in Cyphothrips and may be homologous with similar
structures referred to as ommatidia by Mound (19740; 19746) in Celidothrips species and
Phacothrips ocelloides (Hood) (Pygothripini).
Two species of Zactinothrips are known, both from Peru. The most common species, elegans,
has the apical area of the third antennal segment dark brown, whereas in modes tus this area is
not darkened according to the original description.
SPECIES INCLUDED
e/egansHood, 1936d: 447-452. Holotype cf , PERU (USNM).
*modestusHood, 1941: 230-233. Holotype cf , PERU (USNM).
ZEUGLOTHRIPS Hood
(Figs 342, 376)
Zeuglothrips Hood, 1936d: 452-453. Type-species: Zeuglothrips echinus Hood, by monotypy.
The type-species of this monobasic genus is unique in the Hystricothripina in having the
maxillary stylets greatly elongate, retracted into the head as far as the eyes, and close together
medially (Fig. 342). The postocellar setae are elongate as in Hybridothrips (also Azeugma-
tothrips, Hystricothrips and Saurothrips) , but the pronotal posteroangular setae are elongate
rather than the posteromedials as in H. oneillae. Antennal segments I-V bear enlarged dorsal
setae in Z. echinus, and the pelta bears two setae laterally (Fig. 376) as in the Zeugmatothrips
group of genera rather than the Actinothrips-group. Only two other species have been described
in Zeuglothrips; guerreronsis is here transferred to Hybridothrips as a synonym of oneillae, and
obrieni is transferred to Azeugmatothrips.
SPECIES INCLUDED
ec/iinusHood, I936d: 453-457. Holotype $, PERU (USNM).
88 L. A. MOUND AND J. M. PALMER
ZEUGMATOTHRIPS Priesner
(Figs 350, 351, 364-367, 371, 379, 380)
Zeugmatothrips Priesner, 1925c: 313. Type-species: Zeugmatothrips hispidus Priesner, by monotypy.
The 15 species described in this Neotropical genus exhibit a considerable range of variation,
although most of them can be placed into one of two species-groups. The d/icms-group includes
borgmeieri, cinctus, gracilis, pallidulus andpeltatus. These five species have the mid-dorsal setae
on the head relatively close to the postoculars (Fig. 350), antennal segments IH-IV with two
stout dorsal setae (Fig. 366), and the pelta reduced with the setae of the first abdominal segment
on small sclerites laterally (cf. Fig. 380). The priesneri-group includes annulipes, badiicornis,
badiipes, femoralis, niger, mumbaca and priesneri. These seven species have the mid-dorsal
setae arising well posterior to the postoculars (Fig. 351), antennal segments III-IV with one
stout dorsal seta (Fig. 364), and the setae of the first abdominal segment borne on the lateral
lobes of the broad pelta (Fig. 379). The other three described species are intermediate between
these two groups: bispinosus, according to the description, has the antennae of cinctus-group ,
but the head of priesneri-group; hoodi has the head and pelta of priesneri-group, but antennal
segments III-IV each bear two small, pale dorsal setae (Fig. 367); hispidus has the pelta of
cinctus-group (Fig. 380); the mid-dorsal setae small but distant from the postoculars, and
antennal segments III-IV each with one long and one short, dark dorsal seta (Fig. 365).
Moreover, the authors have studied a further species from Trinidad which differs from hoodi in
being micropterous, with mid-femora pale, and tergites II and III with only one and two
posteroangular setae respectively.
The genus Zeugmatothrips can be recognised by the form of the antennae with stout dorsal
setae, short segment III and lanceolate segment VIII; also the absence of duplicated cilia on the
forewing, the absence of a foretarsal tooth in both sexes, and the stout setose tube. The variation
between species in the colour of the legs is remarkable , but as Hood (1949 : 84-5) has pointed out
these colours may be disruptive coloration associated with their habitat and sluggish habits.
SPECIES INCLUDED
aimu/ipesHood, 1941: 233-236. Holotype $, PERU (USNM; $ paratype BMNH).
* badiicornis Hood, I936d: 457-460. Holotype $, PERU (USNM).
badiipesHood, 1937a: 292-296. Lectotype $ PERU (USNM; $ paralectotype BMNH).
* bispinosus Hood, 1937c: 527-530. Holotype 9 , PERU (USNM).
borgmeieri Hood, 1949: 80-85. Holotype $, BRAZIL (USNM).
cincfusHood, 1952c: 170-171. Lectotype $, BRAZIL (USNM: $ paralectotype BMNH).
*femora7/sHood, 1952c: 169. Holotype $, BRAZIL (USNM).
graci/isHood, 1952c: 171. Syntypes cf $, BRAZIL (USNM).
hispidus Priesner, 1925c: 314-316. Holotype $, MEXICO (SMF).
hoodi Priesner, 19276: 189-192. Syntypes $, COSTA RICA (SMF).
mumbaca Hood, 1952c: 169-170. Lectotype $, BRAZIL (USNM; cf $ paralectotypes BMNH).
njgerHood, 1952c: 168-169. Lectotype $, BRAZIL (USNM, $ paralectotype BMNH).
pallidulus Hood, 1958: 225-228. Holotype $, BRAZIL (USNM: $ paratype BMNH).
*peltatusHood, 1949: 85-88. Holotype $, BRAZIL (USNM).
priesneri Hood, 1935a: 102-106. Holotype $, PANAMA (USNM; cf $ paratypes BMNH).
Taxa transferred from Idolothripinae to Phlaeothripinae
In the opinion of the present authors, several genera listed by Priesner (1961) in his subfamily
'Megathripinae' (= Idolothripinae) are not closely related phylogenetically to that group. In
particular, a series of genera placed in the Cryptothripini by Priesner are here transferred to the
Docessissophothripini in the Phlaeothripinae (Table 5). Moreover, the subtribe Apelaunothri-
pina is here recognised as a distinct tribe and transferred to the Phlaeothripinae from the
Idolothripinae. All of the species concerned have the stylets only moderately broad (i.e. about
5 /xm), except for a few very large or extremely advanced forms (e.g. Tropothrips) , and no
member of the group has been found with large fungal spores in the abdomen. Some of the
species treated here under Holothrips have been found to contain small fungal spores and even
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
89
pieces of fungal hyphae. It is here assumed that these two tribes represent independent
evolutionary lines from fungus-feeding ancestors involving partial adaptation to the spore-
feeding habit. Almost all of the species have the typical phlaeothripine characteristic of short,
stout B2 setae on tergite nine of the males. Both tribes are therefore assumed to have arisen
independently from Hoplandrothrips-\ike ancestors in the Phlaeothripini which also had
elongate maxillary stylets and stout maxillary guides.
Table 5 Genera transferred from Idolothripinae to Phlaeothripinae
Tribe APELAUNOTHRIPINI stat. n.
APELAUNOTHRIPS Karny
Baphothrips Priesner
DEXIOTHRIPS Hartwig
Tribe DOCESSISSOPHOTHRIPINI
ABIASTOTHRIPS Priesner
Cratothrips Priesner
ASEMOTHRIPSHood gen. rev.
Coenothrips Bagnall syn. n.
Eucoenothrips Bagnall syn. n.
Empresmothrips Karny syn. n.
DOCESSISSOPHOTHRIPS Bagnall
Polyphemothrips Schmutz syn. n.
HOLOTHRIPS Karny
Adelothrips Hood syn. n.
Agnostothrips Moulton syn. n.
Cordylothrips Hood syn. n.
Erythrinothrips Ananthakrishnan syn. n.
Holmiella Zur Strassen syn. n.
Ischnothrips Moulton syn. n.
Lathrobiothrips Hood syn. n.
Panceratothrips Bagnall syn. n.
Stinothrips Ananthakrishnan syn. n.
MAXILLATA Faure gen. rev.
OIDANOTHRIPS Moulton
PONGOLA Zur Strassen
SYMPHYOTHRIPSHood & Williams
Mesopotamothrips Liebermann &
Gemignani
TROPOTHRIPSHood gen. rev.
Tribe APELAUNOTHRIPINI stat. n.
The subtribe Apelaunothripina was erected by Priesner (1961: 288) for two genera in his tribe
Megathripini. These genera were placed in 'Megathripinae' because of the moderately broad
stylets. However, Priesner indicted that unlike all other members of that subfamily the males
have setae B2 on the ninth tergite short and stout. Because of these setae, and because no large
fungal spores have been found in the gut and the maxillary stylets are only slightly broader than
in typical phlaeothripines, Mound (19740) treated Apelaunothrips in the Phlaeothripinae. This
relationship was endorsed by Okajima (1979a), who indicated that the second genus, Dexio-
thrips, is closely related to Apelaunothrips despite the different arrangement of the maxillary
stylets. At present the tribe can be defined as those Phlaeothripidae which have B2 setae on
tergite nine of the males short and stout but in which the maxillary stylets are relatively broad
and the maxillary guides stout. In contrast to the Docessissophothripini, which also share these
characters, the Apelaunothripini have long slender antennae with eight segments, and no
metathoracic sternopleural sutures.
APELAUNOTHRIPS Karny
Apelaunothrips Karny, 1925c: 82. Type-species: Ophidothrips medioflavus Karny, by monotypy.
Baphothrips Priesner, 19336: 69-70. Type-species: Baphothrips tricolor Priesner, by monotypy. [Synony-
mised by Mound, I914a: 17.]
Nineteen Old World species are known in this genus, mostly from dead leaves in the eastern
Oriental Region.
SPECIES INCLUDED
armatus Okajima, 19790: 42-4. Holotype $, MALAYA (OCT).
bhowalii( Ananthakrishnan, 19720: 183) (Stigmothrips). Holotype $, INDIA (TNA).
bicolor Okajima, 19790: 44-6. Holotype $, THAILAND (OCT).
consimilis( Ananthakrishnan, 19690: 173-4) (Stigmothrips). Holotype 9, INDIA (TNA).
femora/is Okajima, 19790: 48-9. Holotype $, SINGAPORE (OCT).
gabonensis (Bournier, 1970: 159-162) (Baphothrips). Holotype $, GABON (?BCM).
indicus( Ananthakrishnan, 19680: 125-6) (Philothrips) . Holotype ?$, INDIA (TNA).
japonicus Okajima, 19790: 49-50. Holotype £, JAPAN (OCT).
90 L. A. MOUND AND J. M. PALMER
leios (Mound, 1970: 94-6) (Baphothrips). Holotype $, SOLOMON Is. (BMNH).
Jfen/Okajima, 19790: 50-2. Holotype $, TAIWAN (OCT).
7ur/dus Okajima, 1979a: 52-3. Holotype $, MALAYA (BMNH).
macu/ipeiMi/s (Okajima, 1976: 125-8) (Stigmothrips). Holotype 9, OKINAWA Is. (OCT).
ma7ajens/.s Okajima, 1979a: 54-6. Holotype $, MALAYA (OCT).
medioffavus (Karny , 1925a: 50-2) (Ophidothrips). Holotype $, JAVA (SMF).
montanus Okajima, 1979a: 57-9. Holotype $, JAPAN (OCT).
n/gripeniHsOkajima, 1979a: 59-61. Holotype $, TAIWAN (OCT).
ocu/aris Okajima, 1979o: 61-2. Holotype $>, MALAYA (OCT).
tasmani Mound, 1974a: 18-9. Holotype $, AUSTRALIA (ANIC).
frico/or(Priesner, 1933ft: 70-2) (Baphothrips). Holotype 9, JAVA (SMF).
DEXIOTHRIPS Hartwig
Dexiothrips Hartwig, 1952: 452. Type-species: Dexiothrips pensus Hartwig, by monotypy.
This genus has been discussed by Okajima (19790) who transferred into it a second species.
SPECIES INCLUDED
madrasensis(Ananthakrishnan, 1964ft: 109-10) (Malacothrips) . Syntypes $ cf , INDIA (TNA).
pensus Hartwig, 1952: 453-457. Holotype $, SOUTH AFRICA (NCIP).
Tribe DOCESSISSOPHOTHRIPINI
This tribe was erected by Karny (19210: 257; as a subfamily) for the two genera Docessis-
sophothrips and Egchocephalothrips , although the latter is here removed to the Idolothripina
(p. 76). Nine genera are here placed in the tribe (Abiastothrips, Asemothrips, Docessis-
sophothrips, Holothrips, Maxillata, Oidanothrips, Pongola, Symphyothrips, Tropothrips) ,
although a further 15 generic names are placed in synonymy. The species in these genera share
the following characters.
Antennae with segments VII- VIII more or less fused, III with three sense cones (two in Asemothrips,
Pongola, Symphyothrips, four in Oidanothrips}, IV with four sense cones (two in Symphyothrips and
sometimes Pongola). Maxillary stylets moderately broad, retracted to compound eyes, usually parallel in
middle of head but sometimes looped; maxillary guides stout. Pronotum with epimeral sutures complete;
praepectus absent; metathoracic sternopleural sutures well-developed, anapleural sutures complete
(Figs 412, 413). Wings usually present; forewings with duplicated cilia (except Asemothrips). Pelta (Figs
404-406) usually elongate triangular (relatively broad in Pongola}; tergites with two pairs of sigmoid
wing-retaining setae, although these are sometimes reduced; tube usually with straight sides but in various
species the tube is broadened, ridged or sculptured; tergite IX setae B2 of male short and stout (except
Pongola and Symphyothrips) ; median sternites of male (usually III-V) with one or a pair of transverse
areas of reticulate sculpture which is irridescent under phase contrast microscopy (Fig. 391).
The long maxillary stylets, stout maxillary guides, narrow pelta, thoracic sutures, and the short,
stout B2 setae on tergite nine of males indicate that this group is related to the Phlaeothripini.
However, members of the latter group are usually associated with Basidiomycete fungi and
apparently feed on the external digestion products of the fungal hyphae. In contrast, small pale
spores and even branched hyphae have been found in the gut contents of some Docessis-
sophothripini, and it seems possible that this group of species has specialised on some different
source of fungal food. Unlike typical Phlaeothripini the males do not have a glandular area on
sternite eight, although they usually have characteristic reticulate areas just anterior to the discal
setae on the median sternites (Fig. 391). Somewhat similar glandular areas are found in
Plectrothripini (Okajima, 1981) and the Idolothripinae genus Dichaetothrips (p. 52).
The Docessissophothripini comprises one large, complex and world- wide genus, Holothrips,
with several small or monobasic genera each of which is geographically restricted and definable
from Holothrips only on rather weak characters. This pattern of speciation is to be expected in a
recently evolved group, and reinforces the view that this tribe has evolved relatively recently
from the Phlaeothripini and is phylogenetically distinct from the Idolothripinae.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 91
ABIASTOTHRIPS Priesner
(Fig. 398)
Trichothrips (Abiastothrips) Priesner, I925d: 153. Type-species: Trichothrips schaubergeri Priesner, by
original designation.
Abiastothrips Priesner; Priesner, 1927a: 556. [Raised to genus.]
Cratothrips Priesner, 1927a: 494-5. Type-species: Cratothrips angulatus Priesner, by monotypy. [Synony-
mised by Zur Strassen, 1974: 119-20.]
The type-species of this genus has a broad inter-antennal projection, small eyes and rounded
cheeks; however, soror has a more slender head (Fig. 398) which is similar to some species of
Holothrips. These two species appear to be a Holarctic derivative from the large tropical
complex of Docessissophothripini. Bolothrips lativerticis, from Oregon and Washington State,
in north-western U.S.A., was transferred to Adelothrips by Mound (19746: 181) but is here
recognised as a synonym of schaubergeri from Europe.
SPECIES INCLUDED
* angulatus Priesner, 19270: 495-6. Holotype $, CORSICA (destroyed).
schaubergeri (Priesner, 1920: 86-7) (Trichothrips}. Holotype $, AUSTRIA (SMF).
priesneri Bagnall, 19336: 658 (Cratothrips). Holotype £, AUSTRIA (BMNH).
lativerticis Post, 1961: 141-3 (Bolothrips). Holotype $, CANADA (CAS). Syn. n.
soror Zur Strassen, 1974: 111-20. Holotype $, MADEIRA (SMF).
ASEMOTHRIPS Hood gen. rev.
(Figs 391, 411)
Asemothrips Hood, 19196: 83. Type-species: Asemothrips picturatus Hood, by monotypy.
Empresmothrips Karny, 1920c: 40. Type-species: Empresmothrips combustipes Karny, by monotypy.
Syn. n.
Coenothrips Bagnall, 1924: 629. Type-species: Coenothrips fallax Bagnall, by monotypy. Syn. n.
Eucoenothrips Bagnall, 1926: 553. [Replacement name for Coenothrips Bagnall, see Mound, 1968: 75.]
Syn. n.
Mound (1974a), in establishing the above generic synonymies, used the name Empresmothrips
in error despite the priority of Asemothrips. The genus is used for a group of five Australian
species which share a series of characters with Holothrips species: maxillary stylets and guides,
metathoracic sternopleural and anapleural sutures, males with reticulate glandular areas on
median sternites (Fig. 391) and short B2 setae on tergite nine. However, the species of
Asemothrips lack fore wing duplicated cilia, have only two sense cones on the third antennal
segment and these both arise ventrally, and the sixth antennal segment is broadly truncate
apically (Fig. 411).
SPECIES INCLUDED
combustipes (Karny, 1920c: 41) (Empresmothrips). Holotype $, AUSTRALIA (NRS). Comb. n.
fallax (Bagnall, 1924: 629-30) (Coenothrips). Holotype $, AUSTRALIA (BMNH). Comb. n.
rhopaloides Karny, 1924: 31-2 (Cryptothrips) . Holotype $, AUSTRALIA (NRS).
froudei Girault, 1927e: 1 (Cryptothrips). Holotype $, AUSTRALIA (QMB).
silvae Girault, 1927e: 1 (Cryptothrips). Holotype $, AUSTRALIA (QMB).
finlayi (Girault, 19276: 1) (Cryptothrips). Holotype $, AUSTRALIA (QMB). Comb. n.
folii (Girault, 1928c: 2) (Empresmothrips). Holotype cf , AUSTRALIA (QMB). Comb. n.
pallipes (Karny, 1925a: 22-4) (Empresmothrips). Holotype $, JAVA (SMF). Comb, n., but see Mound,
1971a: 400.
picturatus Hood, 19196: 83-4. Holotype cf , AUSTRALIA (USNM).
longfellowi Girault, 1926: 1 (Empresmothrips). Lectotype $, AUSTRALIA (QMB).
92 L. A. MOUND AND J. M. PALMER
DOCESSISSOPHOTHRIPS Bagnall
(Figs 394, 395, 405, 413)
Docessissophothrips Bagnall, 19086: 201-2. Type-species: Docessissophothrips ampliceps Bagnall, by
monotypy.
Polyphemothrips Schmutz, 1909: 276. Type-species: Polyphemothrips brasiliensis Schmutz, by monotypy.
Syn. n.
The male holotype of ampliceps, described originally when mounted dry on a card, is now
cleared and mounted in balsam on a microscope slide (Fig. 394). This specimen has been
compared with two females from Brazil (in BMNH) which are determined as brasiliensis from
the original description of that species. In all three specimens the head is strongly elevated
medially, and the maxillary stylets lie close together but have a single lateral loop in the
prothoracic region. Moreover, the ampliceps holotype as well as two specimens determined as
brasiliensis (in USNM) have the stylets crossing over each other near the posterior margin of the
head, although the cross-over itself is scarcely wider than the width of the stylets and might be an
artefact. In the unique holotype of dotatus this cross-over is, however, more pronounced; the
stylet arrangement of this specimen thus approaches that found in the species of Tropothrips q. v.
The existence of a lateral loop in the stylets of the following species has kindly been confirmed by
Steve Nakahara (U.S.D.A., Washington): corticis, cuneatus, dotatus, tenuiceps, tibialis, travas-
sosi, woytkowski and yupanqui. In tenuiceps the stylets are angulate on one side but looped on
the other. In villicornis the stylets can only be seen on one side and that is angulate. In cuneatus
the stylets are looped laterally in the holotype (also one male in BMNH) but not in the male
labelled 'allotype'. The species bursarius has the stylets without lateral loops and is here listed
under Holothrips. Moreover, the following three species described by Bagnall in Docessis-
sophothrips are here listed under the genera indicated: laticeps (Polytrichothrips); longiceps
(Bacillothrips);frontalis (Oidanothrips) . The type-species of Docessissophothrips and Polyphe-
mothrips are very similar and must be regarded as congeneric; however, in ampliceps the
mid-vertex setae on the head lie in the same transverse plane as the postocular setae whereas
they lie posterior to the postoculars in brasiliensis and the closely related species major
(Fig. 395). D. amplus is also unusual in having yellow mid- and hindcoxae. D. dotatus is unique
in this group in having four sense cones on antennal segment III as in Oidanothrips.
SPECIES INCLUDED
ampliceps Bagnall, 19086: 202-3. Holotype cf , MEXICO (BMNH).
*annuus Moulton; nomen nudum in Priesner, 1933a: 61. NORTH AMERICA.
brasiliensis (Schmutz, 1909: 276-8) (Polyphemothrips). Holotype ? cf , BRAZIL (? lost). Comb. n.
*corf/c/s(Hood, 1914: 167-9) (Polyphemothrips). Holotype $, PANAMA (USNM). Comb. n.
cuneatus (Hood, 1939a: 217-20) (Polyphemothrips). Holotype <j>, PERU (USNM). Comb. n.
* dotatus (Hood, 1955: 108-110) (Polyphemothrips). Holotype $, BRAZIL (USNM). Comb. n.
major Bagnall, 1912: 215. Holotype $, no data (BMNH).
* tenuiceps (Hood, 1937a: 285-8) (Polyphemothrips). Holotype $, PERU (USNM). Comb. n.
tibialis (Hood & Williams, 1915: 136-7) (Polyphemothrips). Holotype $, U.S.A.: Louisiana (USNM).
Comb. n.
*fravassosi(Hood, 1949: 55-9) (Polyphemothrips). Holotype $, BRAZIL (USNM). Comb. n.
* villicornis (Hood, 1949: 59-62) (Polyphemothrips). Holotype $, BRAZIL (USNM). Comb. n.
*woytkowskyi(Hood, I931a: 288-92) (Polyphemothrips). Holotype $, PERU (USNM). Comb. n.
* yupanqui (Hood, 1937c: 509-13) (Polyphemothrips). Holotype $, PERU (USNM). Comb. n.
HOLOTHRIPS Karny
(Figs 388-390, 393, 399-404, 407, 409, 412)
Holothrips Karny, 1911: 502. Type-species: Holothrips ingens Karny, by monotypy.
Lathrobiothrips Hood, 1933: 421. Type-species: Lathrobiothrips ramuli Hood, by monotypy. Syn. n.
Panceratothrips Bagnall, 1936: 219-20. Type-species: Panceratothrips typicus Bagnall, by monotypy.
Syn. n.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 93
Adelothrips Hood, 1938c: 380. Type-species: A delothrips xanthopus Hood, by original designation. Syn. n.
Cordylothrips Hood, 1937c: 517-8. Type-species: Cordylothrips peruvianus Hood, by monotypy. Syn. n.
Ischnothrips Moulton, 1944: 305. Type-species: Ischnothrips zimmermani Moulton, by monotypy. Syn. n.
Agnostothrips Moulton, 1941a: 172-3. Type-species: Agnostothrips semiflavus Moulton, by monotypy.
Syn. n.
Agnostothrips (Erythrinothrips) Ananthakrishnan, 1956: 341. Type-species: Agnostothrips (Erythri-
nothrips] indicus Ananthakrishnan, by monotypy. [Raised to genus by Ananthakrishnan, 1964: 94.]
Syn. n.
Stinothrips Ananthakrishnan, 1969: 55. Type-species: Ischnothrips typicus Ananthakrishnan, by mono-
typy. Syn. n.
Holmiella Zur Strassen, 1972: 95-8. Type-species: Holmiella nigrita Zur Strassen, by monotypy. Syn. n.
The type-species of Holothrips does not appear to have been re-examined since its original
description. Hood (1952) described three species in the genus, but he is the only other author to
have used the name. Of the three species, onlyprocerus has been examined in the present study,
and this has the head much longer than ingens (Figs 390, 403). In fact, ingens is here regarded as
the senior synonym of fenestralis, described by Hood in Adelothrips, and it is closely related to
lanei. Both of these have the major posteroangular setae on the abdominal tergites long and pale
but flattened and remarkably wide (10 /urn medially). However, this may be subject to sexual
dimorphism because a male identified as lanei (in BMNH) has these setae more slender. Both
ingens and lanei have striate sculpture on tergite II, and the head and pelta are very similar, but
lanei has yellow markings on the hindtibiae and hind margins of the anterior tergites, and ingens
has a curious pale area ventrally on the midfemora.
Most of the species described in Adelothrips are rather small, but no good characters have
been found for segregating these small species into a separate genus from Holothrips. Pancer-
atothrips was erected for a single species with rather stout antennae and with the head elongate
and projecting a little in front of the eyes (Fig. 401). Cordylothrips (Figs 388, 409) was also
erected for a single species with stout antennae, but with segment VI broadly fused to VII + VIII.
An undescribed species (in BMNH) from Peru has been studied, however, with antennae
intermediate in structure (N.B. this species and peruvianus are very similar in head shape and
body structure to Docessissophothrips yupanqui but do not have looped stylets).
The type-series of Ischnothrips (Fig. 389) has the head elongate and elevated dorsally with the
stylets closely approximating to the Docessissophothrips condition, whereas the type-species of
Agnostothrips (Fig. 400) has the head shorter as in ingens. Neither Stinothrips (based on a
micropterous holotype, Fig. 399) nor Erythrinothrips can be distinguished from Holothrips.
Similarly the unique holotype of Holmiella nigrita is here regarded as a large member of
Holothrips with the head relatively elongate, but similar in shape to buccalis and bellulus.
Finally, Lathrobiothrips was erected originally for one species with the tube exceptionally
broad. However, not only do the two subsequent members of this genus have a more elongate
tube (Fig. 393), there are other species described in Adelothrips with the tube stout but apically
constricted (macrura) or broad and heavily sculptured (adelos). None of these can be dis-
tinguished satisfactorily from Holothrips.
The interpretation of the genus Holothrips adopted here thus involves a wide range of body
size, head shape and tube shape. However, the range of form appears to be more or less
continuous with no subgroups being evident above the level of relatively small species-groups.
All the species studied have antennal segments VII+VIII more or less fused, there being four
sense cones on IV and three on III (except phaeura and aberrans with only two on III). The
maxillary guides are well developed, the stylets moderately broad, deeply retracted and parallel
in the middle of the head. The metathoracic sternopleural sutures are well developed, and the
anapleural sutures complete. The pelta is small and bell-shaped or elongate-triangular, the
tergites usually have two pairs of wing-retaining setae (although these are sometimes reduced),
and in males the median sternites have transverse bands of reticulation which probably represent
glandular areas.
Docessissophothrips appears to represent a small group of large-bodied species derived from
Holothrips, and Pongola and Symphyothrips are probably also derived from this genus.
94 L. A. MOUND AND J. M. PALMER
SPECIES INCLUDED
*aberrans(Hood, 1955: 88-90) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
acufus(Stannard, 1956: 108-9 (Adelothrips). Holotype £, U.S.A.: Illinois (INHS). Comb. n.
*adelos (Mound, 1968: 146) (Polyphemothrips). [Replacement name for caudatus Hood.] Comb. n.
caudatus Hood, 1955: 90-2 (Adelothrips). Holotype $, BRAZIL (USNM).
ambit us (Hinds, 1902: 191-2) (Trichothrips) . Holotype $, U.S.A.: Massachusetts (? USNM). Comb. n.
*amp7usHood, 1952: 160-1. Lectotype $, BRAZIL (USNM).
*aspericaudaHood, 1952: 161. Holotype $, BRAZIL (USNM).
australis (Mound, 1974a: 12-5) (Adelothrips). Holotype $, AUSTRALIA: A.C.T. (ANIC). Comb. n.
*7>e77u7us(Hood, 1955: 92-4) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
*Wco7or(Stannard, 1956: 109) (Adelothrips). Holotype $, MEXICO (INHS). Comb. n.
Wpartifus(Hood, I954b: 281-2) (Adelothrips). Holotype $, U.S.A.: Florida (USNM). Comb. n.
bratleyi( Watson, 1935: 61-2, & 1937: 12-13) (Trichothrips) . Syntypes £ cf , U.S.A.: Florida & Alabama
(FSAC). Comb. n.
flavus Moulton & Andre, 1936: 225-6 (Hoplothrips). Holotype $, U.S.A.: Iowa (CAS).
*7wcca7/s(Hood, 1955: 94-6) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
*ftursarius(Hood, 1957: 174) (Polyphemothrips). Holotype $, BRAZIL (USNM). Comb. n.
*carifefte/cus(Stannard, 1956: 109-10) (Adelothrips). Holotype $, MEXICO (INHS). Comb. n.
caudafus(Bagnall, 1915ft: 595-6) (Allothrips) . Holotype £, SARAWAK (BMNH). Comb. n.
c/fr/coriMs(Bagnall, 1913: 296) (Cryptothrips) . Holotype cf , TANZANIA (BMNH). Comb. n.
*comcura (Hood, 1942: 611-5) (Adelothrips). Holotype $, PERU (USNM). Comb. n.
*coiMiatfcornis(Hood, 1925ft: 65-6) (Cryptothrips). Holotype cf , TRINIDAD (USNM). Comb. n.
*cormifus(Hood, 1955: 96-9) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
craceiis( Ananthakrishnan, 1968ft: 55-6) (Polyphemothrips). Holotype $, INDIA (TNA). Comb. n.
*euc7iaro(Hood, 1955: 84-8) (Adelothrips). Holotype 9, BRAZIL (USNM). Comb. n.
/brmosus(Hood, 1952c: 158-9) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
*/UJJIK/IK( Ananthakrishnan, 1972ft: 429-30) (Polyphemothrips). Holotype <j>, INDIA (TNA). Comb. n.
*graminicola (Hood, 1952c: 157) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
*grand/s(Stannard, 1956: 110-1) (Adelothrips). Holotype §, MEXICO (INHS). Comb. n.
*7ia/nmoc7a?nsis(Stannard, 1956: 111-2) (Adelothrips). Holotype $, U.S.A.: Florida (INHS). Comb. n.
indicus (Ananthakrishnan, 1956a: 341-2) (Agnostothrips: Erythrinothrips). Holotype $, INDIA (TNA).
Comb. n.
associates Ananthakrishnan, 1968ft: 56-7 (Symphyothrips). Holotype 9> INDIA (TNA).
insignis(Hood, 1938ft: 162-5) (Lathrobiothrips). Holotype $, PERU (USNM). Comb. n.
ijigensKarny, 1911: 502-3. Holotype $, PARAGUAY (DEI).
fenestralis Hood, 1949: 67-70 (Adelothrips). Holotype $, BRAZIL (USNM). Syn. n.
./ujic*us(Hood, 1912ft: 139-42) (Cryptothrips). Lectotype $, U.S.A.: Michigan (USNM). Comb. n.
quercus Moulton & Andre, 1936: 225 (Hoplothrips). Holotype $, U.S.A.: Iowa (CAS).
7ane/(Hood, 1949: 63-6) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
*lucyae (Gaud, 1961: 117-8) (Polyphemothrips). Holotype cf , PUERTO Rico (RPAESIC). Comb. n.
7uteus(Faure, 1954ft: 147-52) (Polyphemothrips). Holotype $, SOUTH AFRICA (NCIP). Comb. n.
*macrura (Hood, 1941: 185-7) (Adelothrips). Holotype $, CUBA (USNM). Comb. n.
m/nor(Hood, 1937c: 513-7) (PolyphemQthrips). Holotype cf , PERU (USNM). Comb. n.
*mirandus( Ananthakrishnan, 1969c: 305) (Polyphemothrips). Holotype $, INDIA (TNA). Comb. n.
*nepa7e/is/s(Pelikan, 1970: 366-8) (Adelothrips). Holotype $, NEPAL (Innsbruck University). Comb. n.
nigrita (Zur Strassen, 1972: 96-8) (Holmiella). Holotype $, KENYA (NRS). Comb. n.
pa7marum (Hood, 1952c: 157) (Adelothrips). Holotype $, BRAZIL (USNM). Comb. n.
*per/c7es(Hood, 1938c: 383-6) (Adelothrips). Holotype $, U.S.A.: Florida (USNM). Comb. n.
peruvianus(Hood, 1937c: 518-21) (Cordylothrips). Holotype $, PERU (USNM). Comb. n.
*p7iaeura (Hood, 1941: 183-5) (Adelothrips). Holotype $, U.S.A.: Florida (USNM). Comb. n.
procerusHood, 1952: 160. Holotype $, BRAZIL (USNM).
ramu/i(Hood, 1933: 421-2) (Lathrobiothrips). Lectotype $, PANAMA (USNM). Comb. n.
*rofeusfus(Hood, 1954ft: 280-1) (Adelothrips). Holotype $, U.S.A.: Florida (USNM). Comb. n.
*ruidus( Ananthakrishnan, 1969c: 305-6) (Polyphemothrips). Holotype $, INDIA (TNA). Comb. n.
sculptilis (Hood, .1942: 609-11) (Adelothrips). Holotype $ , PERU (USNM). Comb. n.
semifiavus (Moulton, 1947o: 173) (Agnostothrips). Holotype $, NEW GUINEA (CAS). Comb. n.
$7f warrae (Priesner, 1933c: 146-7) (Symphyothrips). Syntype $, MEXICO (SMF). Comb. n.
*sp7end/dus(Johansen, 1977a: 39-40) (Adelothrips). Holotype $, MEXICO (UNAM). Comb. n.
*sporop/iagiis(Stannard, 1956: 112) (Adelothrips). Holotype $, VENEZUELA (INHS). Comb. n.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 95
speciossissimus (Karny , 1920c: 42) (Nesothrips). Holotype cf, AUSTRALIA: Queensland (NRS). Comb. n.
*sfannard/(Ananthakrishnan, 1972ft: 431-2) (Polyphemothrips). Holotype $, INDIA (TNA). Comb. n.
*su&tf//s(Ananthakrishnan, 1972ft: 430-1) (Polyphemothrips}. Holotype $, INDIA (TNA). Comb. n.
*titschaki (Priesner, 1928c: 53-4) (Docessissophothrips). Holotype 9> SOUTH AFRICA (Hamburg, des-
troyed). Comb. n.
f. debilis Priesner, 1928c: 54. Holotype $, SOUTH AFRICA (SMF).
*tumidusDe Santis, 1963a: 7-10. Holotype <5\ ARGENTINA (MLPA).
fjp/cus(Bagnall, 1936: 220) (Panceratothrips) . Lectotype $, MADAGASCAR (MNHN).
fypjcus(Ananthakrishnan, 1967: 235) (Ischnothrips). Holotype $, INDIA (TNA). Comb. n.
*umfcr/eo7a(Hood, 1952c: 159-60) (Adelothrips). Holotype £, BRAZIL (USNM). Comb. n.
woytkowski(Hood, 1942: 615-7) (Lathrobiothrips) . Holotype $, PERU (USNM). Comb. n.
*xanthopus(Hood, 1938c: 380-3) (Adelothrips). Holotype £, U.S.A.: Florida (USNM). Comb. n.
z/mmermann/(Moulton, 1944: 305-6) (Ischnothrips). Holotype C?, FIJI (BPBM). Comb. n.
MAXILLATA Faure gen. rev.
(Fig. 385)
Maxillata Faure, 1949a: 852-3. Type-species: Maxillata priesneri Faure, by monotypy.
This genus was treated in synonymy with Tropothrips and Docessissophothrips by Stannard
(1957). However, unlike the species of those two genera, priesneri does not have any part of the
maxillary stylets parallel medially in the head. The stylets cross over each other between the
compound eyes, and each then follows an independent, undulating course to the mouth
aperature (Fig. 385). This stylet arrangement could be interpreted as part of a single transforma-
tion series, that is with Maxillata regarded as a more advanced and complicated form of
Tropothrips, and the two genera placed in synonymy. However, the alternative interpretation is
adopted here, that the two genera represent independent lines of evolution from Holothrips.
Two specimens of an unidentified species from Jamaica (in BMNH) have the stylets arranged
similarly to priesneri as figured by Faure (1949: 855). These specimens cannot, by themselves, be
taken as indicating that the Maxillata stylet arrangement has evolved independently in both Old
and New Worlds, because natural distribution patterns of fungus-feeding thrips are known to
have been disrupted by human trading activity between Africa and the West Indies (Mound,
\974b: 111). In this connection a single specimen from Ghana (in BMNH) is also of interest. The
head of this specimen is like an exaggerated form of Tropothrips (Fig. 386), but although the
stylets are parallel medially they are both displaced laterally to the right-hand side in the
posterior half of the head before producing one or more convolutions. This specimen is dark
brown and much larger than Tropothrips or Maxillata species, although the body is essentially
similar in structure to large species of Holothrips. It is here regarded as yet another independent
off-shoot of Holothrips, but cannot be formally described because the specimen lacks antennae.
SPECIES INCLUDED
* priesneri Faure, 1949a: 854-8. Holotype $, SOUTH AFRICA (NCIP).
OIDANOTHRIPS Moulton
(Figs 396, 408)
Oidanothrips Moulton, 1944: 308-9. Type-species: Oidanothrips magnus Moulton, by monotypy.
This genus is used here for four large Old World species which are similar to Holothrips species
but have four sense cones on both the third and fourth antennal segments (Fig. 408). These
species probably constitute a holophyletic group, but it is likely that this has developed from
within the genus Holothrips rather than as a true sister-group. In the type-species sigmoid
wing-retaining setae are developed only on tergites II to IV, and the anterior pair is reduced on
each segment. The unique holotype offrontalis (Fig. 396) was rediscovered recently, dry in a
tube (contrary to Mound, 1968), but lacks antennae. It is referred to this genus on the basis of
fresh material from Japan and Malaya (Fig. 408). Moreover, megacephalus is probably the same
species asfrontalis judging from the description.
96 L. A. MOUND AND J. M. PALMER
SPECIES INCLUDED
*enormis(Ananthakrishnan, 1969c: 302-3) (Polyphemothrips) . Holotype 9, INDIA (TNA). Comb. n.
/hMifatfs(Bagnall, 1914o: 26-7) (Docessissophothrips). Holotype 9, JAPAN (BMNH). Comb. n.
*femoralis Ishida, 1932: 6-7 (Machatothrips). Holotype 9> JAPAN (Hokkaido Univ.); Kurosawa,
1968: 58.
mag/iusMoulton, 1944: 309-10. Holotype 9, FIJI (BPBM).
*/negacep/ia7us(Ananthakrishnan, 1969c: 303-4) (Polyphemothrips). Holotype 9> INDIA (TNA). Comb,
n.
PONGOLA Zur Strassen
Pongola Zur Strassen, 1959: 186-7. Type-species: Pongola rufianalis Zur Strassen, by monotypy.
The only species in this genus has reduced, almost moniliform antennae with two sense cones on
segment III and 4 or 3 (even 2) on segment IV. The pronotal epimeral sutures are incomplete
and each tergite bears only a single pair of wing-retaining setae. However, the metathoracic
sternopleural sutures are present, and the maxillary guides are long and stout although not
densely sclerotised. The two most unusual (? apomorphic) characters of the species are the
relatively broad pelta (in contrast to Holothrips} and the short, medially constricted tube.
However, even the condition of these two characters could be regarded as the extremes of
transformation series found within Holothrips - from which Pongola is almost certainly derived.
The male has not been examined during the present studies.
SPECIES INCLUDED
rufianalisZur Strassen, 1959: 187-97. Holotype 9, SOUTH AFRICA (NCIP).
SYMPHYOTHRIPS Hood & Williams
(Figs 392, 397, 406, 410)
Symphyothrips Hood & Williams, 1915: 131. Type-species: Symphyothrips punctatus Hood & Williams, by
monotypy.
Mesopotamothrips Liebermann & Gemignani, 1931: 212. Type-species: Mesopotamothrips concordiensis
Liebermann & Gemignani, by monotypy. [Synonymised by De Santis, 1959: 248.]
Most of the species listed in this genus have not been studied by the present authors, and Hood
(1952) indicated that possibly only caliginosus is congeneric with punctatus . A single specimen
from Panama has been examined (in BMNH) with more slender antennal segments than
punctatus and only one sense cone on segment III (Fig. 410), and this apparently represents a
third species. The genus is closely related to Holothrips but with only 2 (or 1) sense cones on
antennal segment III and 2 on IV. The suture between antennal segments VII and VIII is poorly
developed or absent, but the pelta is shorter and broader than in most Holothrips species
(Fig. 406), being very similar to many Haplothrips species. The maxillary stylets and maxillary
guides are similar to Holothrips (Fig. 397), and the metathoracic sternopleural sutures are well
developed, but the anterior pair of wing-retaining setae is often absent on each tergite, even in
macropterae. Males of punctatus have the typical iridescent reticulate areas anterior to the discal
setae on sternites IV- VI, but setae B2 on tergite nine are long and slender. The short swollen
tube of Symphyothrips (Fig. 392) species has also evolved in Holothrips, amongst the species
described under the name Lathrobiothrips. The position of Mesopotamothrips requires further
confirmation, because the illustration of the antenna given by de Santis (1959: 249) suggests that
concordiensis might be a species of Holothrips.
SPECIES INCLUDED
*aberrans Ananthakrishnan, 1971: 201-2. Holotype 9, INDIA (TNA).
*a///anensjsBianchi, 1949: 348-50. Holotype 9, GUAM (BPBM).
* caliginosus Hood, 1952c: 163-4. Lectotype 9, BRAZIL (USNM).
* concordiensis (Liebermann & Gemignani, 1931: 213-4) (Mesopotamothrips). Syntypes 9> ARGENTINA
(MACN).
*longicauda Priesner, 1924: 150. Holotype d", Baltic amber fossil (? lost).
*/ongicoriMsPriesner, 1921: 200-2. Syntypes 9 d", PARAGUAY (SMF).
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 97
*potosiensisMou\ton, 1947ft: 419-20. Holotype $, MEXICO (CAS).
punctatus Hood & Williams, 1915: 131-3. Holotype $, U.S.A.: Florida (USNM).
*reticulatus Watson, 1925: 29-30, 45. Holotype $, ARGENTINA (FSAC).
TROPOTHRIPS Hood gen. rev.
(Fig. 387)
Tropothrips Hood, 1949: 70-1. Type species: Tropothrips borgmeieri Hood, by monotypy.
This genus has been treated as a synonym of Docessissophothrips (Stannard, 1957), but although
closely related these two genera are here distinguished by the arrangement of the maxillary
stylets. In species of both genera the stylets lie close together and parallel medially for at least a
short distance, but in Docessissophothrips each stylet has only one lateral loop whereas in
Tropothrips the stylets cross over each other to form a large median loop as well as the pair of
lateral loops (Fig. 387). This is not a fundamental difference, however, because D. dotatus and
D. ampliceps each has a small posteromedian cross-over loop. The genus Maxillata (q.v.) is also
recognised here because of the even more complex arrangement of the stylets. Apart from the
maxillary stylets, the rest of the body of Tropothrips species, including the antennae, pelta and
tergites, is essentially similar to that of Holothrips species. The male holotype of dampfi has the
typical median transverse band of reticulation (? glandular area) on sternites V-VI anterior to
the discal setae.
SPECIES INCLUDED
* borgmeieri Hood, 1949: 71-6. Holotype $, BRAZIL (USNM).
dampfi (Priesner, 1933a: 59-61) (Docessissophothrips). Holotype d", MEXICO (SMF).
*nigripes Stannard, 1954a: 84. Holotype $, COSTA RICA (INHS).
*richardsi Stannard, 1954o: 82-4. Holotype $, U.S.A.: Florida (INHS).
*tuxtlae Johansen, 1977a: 37-9. Holotype £, MEXICO (UNAM).
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19086. Three new North American Phloeothripidae. Canadian Entomologist 40: 305-309.
1908c. Two new species of Idolothrips. Annals of the Entomological Society of America 1: 285-289.
1912a. Three new Phloeothripidae (Thysanoptera) from Texas and Michigan. Proceedings of the
Biological Society of Washington 25: 11-15.
19126. On North American Phloeothripidae (Thysanoptera), with descriptions of two new species.
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1912c. Descriptions of new North American Thysanoptera. Proceedings of the Entomological Society
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1914. Studies on tubuliferous Thysanoptera. Proceedings of the Biological Society of Washington 27:
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1915. A remarkable new thrips from Australia. Proceedings of the Biological Society of Washington
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1916. Oedaleothrips hookeri, a new genus and species of Thysanoptera. Bulletin of the Brooklyn
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1918a. A new Kleothrips (Thysanoptera) from North Queensland. Bulletin of the Brooklyn Entomo-
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19186. New genera and species of Australia Thysanoptera. Memoirs of the Queensland Museum 6:
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1919o. On some new Idolothripidae. Insecutor Inscitiae Menstruus 7: 66-74.
19196. Two new genera and thirteen new species of Australian Thysanoptera. Proceedings of the
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1924. New Thysanoptera from the United States. Entomological News 35: 312-317.
1925o. On some new Phloeothripidae (Thysanoptera) from the Transvaal. Psyche 31: 292-301.
19256. New neotropical Thysanoptera collected by C. B. Williams. Psyche 32: 48-69.
1925c. Six new Thysanoptera from the western United States. Entomological News 36: 101-105,
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1925d. Four new Phloeothripidae from the United States (Thysanoptera). Canadian Entomologist
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I927a. A contribution toward the knowledge of New York Thysanoptera, with descriptions of new
genera and species II. Entomologica Americana 7: 209-245.
19276. New western Thysanoptera. Proceedings of the Biological Society of Washington 40: 197-204.
1928. A new Actinothrips (Thysanoptera) from Brazil. Bulletin of the Brooklyn Entomological
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102 L. A. MOUND AND J. M. PALMER
1933. New Thysanoptera from Panama. Journal of the New York Entomological Society 41: 407-434.
1934. Some further new Thysanoptera from Panama. Proceedings of the Biological Society of
Washington 47: 57-81.
1935a. Ten new Thysanoptera from Panama. Proceedings of the Biological Society of Washington 48:
83-106.
19356. Some new or little known Thysanoptera of the family Phlaeothripidae. Revista de Entomolo-
gia, Rio de Janeiro 5: 159-199.
— 1935c. The thysanopterous genus Actinothrips. Stylops 4: 247-254.
— 1936a. Nine new Thysanoptera from the United States. Journal of the New York Entomological
Society 44: 81-100.
— 19366. Two new Anactinothrips from South America (Thysanoptera). Proceedings of the Royal
Entomological Society of London (B) 5: 143-147.
— 1936c. Studies in Neotropical Thysanoptera. I. Revista de Entomologia, Rio de Janeiro 6: 248-279.
— 1936d. Studies in Neotropical Thysanoptera. II. Revista de Entomologia, Rio de Janeiro 6: 424-460.
— 1937a. Studies in Neotropical Thysanoptera. IV. Revista de Entomologia, Rio de Janeiro 7: 255-296.
— 19376. A new ant-like thrips from Florida. Proceedings of the Biological Society of Washington 50:
111-113.
— 1937c. Studies in Neotropical Thysanoptera. V. Revista de Entomologia, Rio de Janeiro 7: 486-530.
— 1938a. A new genus and species of Phlaeothripidae (Thysanoptera) from Palmetto. Proceedings of
the Biological Society of Washington 51: 27-32.
— 19386. Studies in Neotropical Thysanoptera. VI. Revista de Entomologia, Rio de Janeiro 8: 161-187.
— 1938c. New Thysanoptera from Florida and North Carolina. Revista de Entomologia, Rio de Janeiro
8: 348^20.
— I938d. Studies in Neotropical Thysanoptera. VII. Revista de Entomologia, Rio de Janeiro 9: 218-247.
— 1939a. A new Polyphemothrips (Thysanoptera) from Peru. Revista Chilena de Historia Natural 42:
217-220.
19396. New North American Thysanoptera, principally from Texas. Revista de Entomologia, Rio de
Janeiro 10: 550-619.
1940a. A century of new American Thysanoptera. I. Revista de Entomologia, Rio de Janeiro 11:
540-583.
1940b. The cause and significance of macropterism and brachypterism in certain Thysanoptera,
with description of a new Mexican species. Anales de la Escuela Nacional de Ciencias Biologicas 1:
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1941. A century of new American Thysanoptera. II. Revista de Entomologia, Rio de Janeiro 12:
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1942. A century of new American Thysanoptera. III. Revista de Entomologia, Rio de Janeiro 12:
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1949. Brasilian Thysanoptera. I. Revista de Entomologia, Rio de Janeiro 20: 3-88.
1950. Brasilian Thysanoptera. II. Revista de Entomologia, Rio de Janeiro 21: 1-113.
1951. A new Oedaleothrips from Italy. Memorie della Societa Entomologica Italiana 30: 133-140.
1952a. A new Oedaleothrips from the Belgian Congo. Revue de Zoologie et de Botanique Africaine
45: 204-209.
19526. Brasilian Thysanoptera. III. Proceedings of 'the Biological Society of 'Washington 65: 141-174.
1952c. Mecynothrips snodgrassi, a new thrips from the Solomon Islands. Proceedings of the
Entomological Society of Washington 54: 294-301.
1954a. Brasilian Thysanoptera. IV. Proceedings of the Biological Society of Washington 67: 17-54.
19546. New Thysanoptera, principally Floridian. Proceedings of the Biological Society of Washington
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1954c. Three new Thysanoptera from Trinidad and British Guiana. Proceedings of the Royal
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1955. Brasilian Thysanoptera. VI. Revista Brasileira de Entomologia, Sao Paulo 4: 51-160.
1957. New Brazilian Thysanoptera. Proceedings of the Biological Society of Washington 70: 129-180.
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GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 103
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1939a. New species of Phlaeothripidae (Thysanoptera) from South Africa. Journal of the Entomolo-
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GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 105
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GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 107
— 1936a. Ueber einege neue und wenig bekannte Thysanopteren. Proceedings of the Royal Entomolo-
gical Society of London (B)5: 208-214.
19366. On some further new Thysanoptera from the Sudan. Bulletin de la Societe Roy ale Entomolo-
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— 1937a. Thysanopterologica V: Zwei neue Phlaeothripiden. Arbeiten iiber morphologische und
taxonomische Entomologie 4: 347-350.
— 1937ft. Two new Phlaeothripidae from Sierra Leone. Annals and Magazine of Natural History (10)
20: 624-629.
1939a. Thysanopterologica VIII. Proceedings of the Royal Entomological Society of London (B)8:
73-78.
— 19396. Thysanopteren aus dem Belgischen Congo. Revue de Zoologie et de Botanique Africaines 33:
49-66.
— 1940. Thysanopterologica IX. Philippine Journal of Science 71: 403-411.
— 1949. Genera Thysanopterorum. Keys for the identification of the genera of the Order Thysanoptera.
Bulletin de la Societe Fouad ler d' Entomologie 33: 31-157.
— 1950. Contributions towards a knowledge of the Thysanoptera of Egypt, XV. Bulletin de la Societe
Fouad ler d' Entomologie 34: 25-37.
— 1951. Thysanopterologica XI. Annals and Magazine of Natural History (12)4: 355-371.
— 1952a. On some new genera and species of Thysanoptera from the Oriental region. Indian Journal of
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— 19526. On some central African Thysanoptera. Bulletin de I'lnstitut Franfais d'Afrique Noire 14:
842-880.
— 1959. On the genus Dinothrips Bgn. (Thysanoptera). Idea 12: 52-59.
1961. Das System der Tubulifera (Thysanoptera). Anzeiger Osterreichische Akademie der Wissen-
schaften 1960: 283-296.
1964. Ordnung Thysanoptera. Bestimmungsbiicher zur Bodenfauna Europas 2, 242 pp. Berlin.
Ramakrishna Ayyar, T. V. 1925. Two new Thysanoptera from South India. Journal of the Bombay Natural
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1928. A contribution to our knowledge of the Thysanoptera of India. Memoirs of the Department of
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1934. Entomological investigations on the spike disease of Sandal (21), Thysanoptera. Indian Forest
Records 20: 1-12.
Ramakrishna Ayyar, T. V. & Margabandhu, V. 1939. Notes on new and known Indian Thysanoptera.
Records of the Indian Museum 41: 21-33.
Rayment, T. 1948. Observations on Thrips, with description of a new species. Australian Zoologist 11:
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1901. Thysanoptera tria Mediterranea. Ofversigt af Finska Vetenskaps-Societetens Forhandlingar 43:
214-215.
Sakimura, K. 1971. A review of the genus Rhaebothrips Karny (Thysanoptera: Phlaeothripidae). Pacific
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108 L. A. MOUND AND J. M. PALMER
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1933. Two new species of Oedaleothrips with notes on other species. Florida Entomologist 17: 48-50;
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1934/35/37. Thysanoptera of the Geenton. Florida Entomologist 18: 44-46, 55-62; 20: 12-15, 17-21.
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Yakhontov, V. V. 1956. New genus and species of thrips from Zailisk Ala-Tai. [In Russian.] Zoo-
logicheskii Zhurnal35: 554-555.
Zur Strassen, R. 1959. Studies in African Thysanoptera, 2. Journal of the Entomological Society of
Southern Africa 22: 174-198.
- 1966. Neue Fransenfliiglearten aus der Unterordnung Tubulifera von den Kanarischen Inseln (Ins.,
Thysanoptera). Commentationes Biologicae 29: 1-34.
- 1968. Okologische und zoogeographische Studien iiber die Fransenfliiger-Fauna (Ins., Thysanop-
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1-125.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 109
— 1972. Athiopische Thysanopteran iiberwiegend aus dem Massiv des Mount Elgon in Kenya (Insecta,
Thysanoptera). Zoologica Scripta 1: 85-105.
— 1974. Neue silvicole Fransenfliigler von den makaronesischen Inseln (Ins.: Thysanoptera). Sencken-
bergiana Biologica 55: 105-134.
- 1976. La faune terrestre de 1'Ile de Sainte-Helene. Part 3. 18. Thysanoptera. Annales Musee Royal de
I'Afrique Centrale, Sciences Zoologiques (8) 215: 236-256.
— 1977. Internationales Forschungsprojekt Makaronesischer Raum. Studie zur Friihjahrsfauna der
Fransenfliigler auf den Madeira-Inseln im Atlantik nebst Daten zur Abundanz und Faunistik (Insecta:
Thysanoptera). Boletim do Museu Municipal do FunchalSl: 5-78.
— 1979. Insects of Saudi Arabia. Thysanoptera. Fauna of Saudi Arabia 1: 90-104.
— 1980. Anactinothrips gibbifer n. sp. aus Baumkronen im Amazonas-Gebiet (Insecta: Thysanoptera:
Phlaeothripidae). Senckenbergiana Biologica 61: 47-56.
Table 6 Distribution of species in Pygothripina genera
NT NA PA AT O A P
Cleistothrips 1
Heptathrips 3 5
Ozothrips 3
Pelinothrips 2
Phaulothrips 1 10 1
Emprosthiothrips 6
Priesneriana 2 2
Cryptothrips 34 1
Pygothrips 54 — — 21
Table 7 Distribution of species in Allothripina genera
NT NA PA AT O A P
Allidothrips 1 1
Allopisothrips 1
Allothrips 352233
Faureothrips 1
Priesneriella 34 1
Pseudocryptothrips 1 11
Table 8 Distribution of species in Compsothripina genera
NT NA PA AT O A P
Anaglyptothrips 1
Bolothrips 565
Compsothrips 87433
lllinothrips 1
Loyolaia 1
Table 9 Distribution of species in Gastrothripina
NT NA PA AT O A P
Gastrothrips 18 5 1 3 6
HO L. A. MOUND AND J. M. PALMER
Table 10 Distribution of species in Diceratothripina genera
NT NA PA AT O A P
Carientothrips 1 13
Nesothrips 3 9 13
Campulothrips 1
Elgonima
Pseudoeurynchothrips 2
Neosmerinthothrips 11 46 1
Phacothrips 1
Nesidiothrips 1 1
Acallurothrips 71 52 1
Dlceratothrips 16 5
Sporothrips 1
Table 1 1 Distribution of species in Macrothripina genera
NT NA PA AT O A P
Aesthesiothrips
Celidothrips 2
Diaphorothrips 2
Dichaetothrips 1 2
Diplacothrips 2
Ethirothrips 1 4 13 10 4
Machatothrips 3 11
Pe/tan'ctf /in/?s
Po/yfr/c/zof/iri/w
Tarassothrips 1
Table 12 Distribution of species in Elaphrothripina genera
NT NA PA AT O A P
Anactinothrips 15
Elaphrothrips 46 7 49 19
Ophthalmothrips 451
Mecynothrips 1932
Lamillothrips 3
Dinothrips 5
Tiarothrips 1
Hartwigia 1
Malesiathrips 1 2
Dermothrips 1
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 111
Table 13 Distribution of species in Idolothripina genera
NT NA PA AT O A P
Idolothrips
Meiothrips 3
Lasiothrips
Egchocephalothrips 1
Megalothrips 3 1
Bacillothrips 3
Bactrothrips 1 1 32 7 1
Megathrips 5
Ceuthothrips 1
Cylindrothrips 1
Table 14 Distribution of species in Hystricothripina genera
NT NA PA AT O
Hystricothrips 2
Holurothrips 2
Paractinothrips 1
Neatractothrips 1
Actinothrips 11
Atractothrips 1 1
Azeugmatothrips 2
Cyphothrips 1
Hybridothrips 1
Saurothrips 1
Zactinothrips 2
Zeuglothrips 1
Zeugmatothrips 15
112
L. A. MOUND AND J. M. PALMER
Figs 2-6 Pygothripina. 2, Cryptothrips nigripes mac. ; 3, Heptathrips magnified 9 paratype; 4, H. tonnoiri
$ mac.; 5, Cleistothrips idolothripoides $ mac.; 6, Pygothrips fortis cf apt.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
113
8
Figs 7-1 1 Pygothripina: Phaulothrips species. 7, P. vuilleti $ ; 8, P. inquilinus $ ; 9, P. wptom $ holotype ;
10, P. magnificus cf allotype; 11, P. barretti $ holotype.
114
L. A. MOUND AND J. M. PALMER
16
' x \
/ ~\ys
\
c 1
I/ /
1
'"',' M
•
^
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*
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1
V
1
! 1
Figs 12-17 Pygothripina. 12, Ozothrips janus 9 holotype; 13, 0. eurytis $ holotype; 14, 0. priscus $
holotype; 15, Pygothrips mikrommatos $ holotype; 16, Pries neriana kabandha $; 17, Pelinothrips
brochotus $ holotype.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
115
21
22
23
Figs 18-24 Pygothripina. 18, Emprosthiothrips brimblecombei 9 holotype; 19, E. epallelus $ holotype;
20, E. niger thoracic sternites; 21-22, Pygothrips fortis eroded metathoracic sternopleural sutures in $
(21) and cf (22); 23, P. fortis cf tube; 24, Cleistothrips idolothripoides $ tube.
25
32
33
34
28
29
Figs 25-39 Pelta of Pygothripina (and Diceratothripina) species. 25, Phaulothrips sibylla $ mac. ; 26, P.
vuilleti; 27, P. barretti holotype; 28, P. magnificus (3" allotype; 29, P. anici holotype; 30, P. sibylla 9 mic. ;
31, Emprosthiothrips niger; 32, Cleistothrips idolothripoides; 33, Heptathrips tonnoiri; 34, Cryptothrips
nigripes; (35, Diceratothrips nigricauda;) 36, Phaulothrips sculpticauda 9; 37, P. sculpticauda C?
holotype; 38, Pelinothrips brochotus holotype; 39, P. ornatus.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
117
Figs 40-46 Pygothripina. 40, Ozothripsjanus, holotype pelta; 41, O. prisons holotype pelta & tergite II;
42, O. eurytis holotype; 43, Phaulothrips vuilleti tergite III; 44, P. agrestis tergite III; 45, P. agrestis
antennal segments I-IV; 46, Cleistothrips idolothripoides.
118
L. A. MOUND AND J. M. PALMER
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
119
56
Figs 56-66 Allothripina. 56, Pseudocryptothrips sp.; 57, Allothrips megacephalus stannardi; 58, A. m.
stannardi pelta; 59, Priesneriella clavicornis; 60, P. thomasi; 61, P. citricauda; 62, P. gnomus holotype
pelta & tergite II; 63, P. seminole pelta & tergite II; 64, Allidothrips tricolor tergite I (pelta); 65,
Pseudocryptothrips sp.; 66, Faureothrips reticulatus.
120
L. A. MOUND AND J. M. PALMER
Figs 67-73 Allothripina. 67, Faureothrips reticulatus; 68, Priesnerlella thomasi; 69, P. seminole head &
pronotum; 70, P. clavicornis; 71, P. gnomus holotype; 72, P. seminole tergite IX & tube; 73,
Pseudocryptothrips sp.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
121
74
77
78
Figs 74-83 Allothripina. 74, Priesneriella gnomus tergite IV; 75, P. seminole tergite IV; 76, P. seminole
tergite VII; 77, Priesneriella sp. left maxillary palp; 78, Allothrips megacephalus stannardi; 79,
Priesneriella gnomus; 80, P. citricauda; 81, P. clavicornis; 82, P. thomasi; 83, P. seminole.
122
L. A. MOUND AND J. M. PALMER
Figs 84-93 Compsothripina and Gastrothripina. 84, Compsothrips albosignatus; 85, C. reuteri; 86, C.
hookeri; 87-90, tubes of (87) Gastrothrips proturus holotype; (88) G. falcatus; (89) G. ruficauda; (90) G.
intonsus; 91, G. ruficauda antennal segments VII-VIII; 92, G. ruficauda antennal segment III; 93, G.
turbinatus.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
123
94
95
T
i!
1 L
n
S M
// v! /i
/
I .'
I/ '
/ -
I '
J (
T
\ ll
U:
;
100
Figs 94-102 Compsothripina. 94, Bolothrips bicolor; 95, B. italicus; 96, B. pratensis; 97-100, metaster-
num of (97) Anaglyptothrips dugdalei; (98) Bolothrips cingulatus; (99) B. icarus; (100) Compsothrips
reuteri; 101, Bolothrips bicolor pelta; 102, B. italicus pelta.
124
L. A. MOUND AND J. M. PALMER
-vrvV ^ '?*-Jt
' , ' -f
~f X »- x^ /-> ^~ >
""T
104
105
107
109
108
110
Figs 103-110 Compsothripina. 103. Anaglyptothrips dugdalei $ ; 104, Illinothrips rossi $ ; 105, Loyolaia
indica; 106-110, peltaof (W6) Anaglyptothrips dugdalei; (107) Compsothrips reuteri; (108) C. albosigna-
tus; (109) Loyolaia indica; (110) Illinothrips rossi.
111
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
119
125
Figs 111-119 Compsothripina. Ill, Compsothrips reuteri metanotum; 112, C. albosignatus metanotum;
113, C. albosignatus; 114, C. reuteri; 115, Anaglyptothrips dugdalei; 116, lllinothrips rossi; 117, Loyolaia
indica; 118, Bolothrips bicolor; 119, B. dngulatus III-IV.
126
L. A. MOUND AND J. M. PALMER
120
Figs 120-129 Gastrothripina. 120, Gastrothrips anolis cf ; 121-124, pelta of (121) G. anolis- (122) G.
ruficauda; (123) G. mandiocae; (124) G. intonsa; 125, G. ruficauda; 126, G. turbinatus; 127, G. mauli
anapleural suture; 128, G. acuticornis metasternum; 129, G. fulviceps pelta.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
127
130
132
Figs 130-133 Diceratothripina. 130, Diceratothrips bicornis cf; 131, D. bennetti $ paratype; 132,
Neosmerinthothrips hamiltoni $ holotype; 133, Campulothrips gracilis .
128
L. A. MOUND AND J. M. PALMER
134
135
136
138
137
139
Figs 134-139 Diceratothripina. 134, Nesidiothrips alius; 135, Carientothrips magnetis holotype; 136, C.
loisthus holotype; 137, C. pedicillus holotype; 138, Neosmerinthothrips fructuum; 139, A/, hilaris.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
140
129
143
Figs 140-144 Diceratothripina. 140, Sporothrips amplus cf ; 141, Phacothrips ocelloides; 142, Nesothrips
malaccae; 143, N. rangi 9 holotype; 144, N. eastopi $ holotype.
130
L. A. MOUND AND J. M. PALMER
145
146
152
147
Figs 145-152 Diceratothripina. 145, Acallurothrips spinicauda; 146, A. flavus $ paratype; 147, Spor-
othrips amplus cf foretarsus; 148, Campulothrips gracilis d" foreleg; 149, Diceratothrips bicornis cf
forecoxal stridulatory ridges; 150-152, metasternum of (150) Sporothrips amplus; (151) Campulothrips
gracilis; (152) Diceratothrips bennetti.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
131
155
159
160
Figs 153-160 Diceratothripina tubes. 153, Neosmerinthothrips hoodi;154, N. hamiltoni; 155, N. variipes;
156, N. xylebori; 157, Nesidiothrips alius; 158, Phacothrips ocelloides; 159, Nesothrips brevicollis; 160,
A/, eastopi.
132
L. A. MOUND AND J. M. PALMER
161
164
Figs 161-170 Diceratothripina. 161, Acallurothrips spinicauda tube; 162, Campulothrips gracilis tube;
163, Diceratothrips bennetti $ tube; 164, Carientothrips grayi; 165, Diceratothrips bennetti $; 166,
Phacothrips ocelloides; 167, Nesothrips rangi $ holotype; 168, Acallurothrips flavus; 169, Neosmerin-
thothrips hamiltoni 9 holotype; 170, Sporothrips amplus.
171
Figs 171-186 Pelta of Diceratothripina species. 171, Dicer atothrips bennetti C? holotype; 172, Neosmer-
inthothrips hamiltoni 9 holotype; 173, N. fructuum; 174, Nesidiothrips alius; 175, Nesothrips brevicollis;
176, N. fodinae; 111, N. leveri; 178, N. rangi $ holotype; 179, Acallurothrips badius; 180, A. flavus; 181 ,
Carientothrips acti; 182, C. mjobergi $ apt.; 183, C. mjobergi $ mic.; 184, C. magnetis; 185, C. mjobergi
$ mac.; 186, Phacothrips ocelloides.
134
187
L. A. MOUND AND J. M. PALMER
188
>-•*;: : :-::«r-
Figs 187-192 Macrothripina. 187, Ethirothrips agasthya; 188, Herathrips nativus; 189, Ethirothrips
tibialis; 190, £. stenomelas 9 holotype; 191, £. sybarita; 192, E. australiensis .
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
135
193
Figs 193-197 Macrothripina. 193, Ethirothrips anacardii; 194, Peltariothrips insolitus V holotype; 195,
Polytrichothrips laticeps $ holotype; 196, Tarassothrips akritus $ holotype; 197, Diplacothrips piceus .
136
L. A. MOUND AND J. M. PALMER
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GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
137
Figs 199-202 Macrothripina. 199, Dichaetothrips secular $ holotype; 200, Aesthesiothrips jatrophae;
201, Celidothrips lawrencei; 202, Dichaetothrips okajimai 9 holotype.
138
L. A. MOUND AND J. M. PALMER
203
Figs 203-206 Macrothripina. 203, Machatothrips biuncinatus; 204, M. antennatus; 205, Diaphorothrips
clavipes', 206, Macrothrips papuensis 9-
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
139
207
208
212
213
214
209
215
210
216
211
Figs 207-216 Macrothripina. 207, Dlaphorothrips hamipes; 208, D. clavipes III-IV; 209-216, pelta of
(209) Macrothrips papuensis; (210) Machatothrips biuncinatus; (211) Polytrichothrips laticeps; (212)
Pdtariothrips insolitus; (213) Ischyrothrips crassus $ holotype; (214) Diaphorothrips clavipes; (215)
Celidothrips ? adiaphorus; (216) Aesthesiothrips jatrophae .
140
L. A. MOUND AND J. M. PALMER
217
223
219
220
222
221
224
\
226
225
227
Figs 217-227 Macrothripina. 217, Machatothrips haplodon foreleg; 218, M. antennatus forefemur; 219,
M. heveae forefemur; 220, Diaphorothrips davipes foretarsus; 221 , Aesthesiothrips jatrophae foretarsus;
222-227, Pelta of (222) Diplacothrips borgmeieri; (223) Tarassothrips sp. indet.; (224) T. akritus; (225)
Dichaetothrips secutor; (226) D. brevicollis holotype; (227) D. okajimai.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
141
228
235
Figs 228-238 Macrothripiaa. 228-234, pelta of (228) Ethirothrips agasthya; (229) £. /zwws; (230) E.
australiensis; (231) £. distasmus; (232) E. sybarita; (233) £. obscurus (neivai holotype); (234) £.
stenomelas; 235, Dichaetothrips brevicollis tergite V; 236, D. okajimai sternite V; 237, Machatothrips
lentus tergite III; 238, Aesthesiothrips jatrophae tergite III.
142
L. A. MOUND AND J. M. PALMER
248
240
244
Figs 239-248 Macrothripina. 239, Diplacothrips borgmeieri tube; 240, Macrothrips papuensis; 241,
Celidothrips lawrencei; 242, Peltariothrips insolitus holotype; 243, Polytrichothrips laticeps (left); 244,
Ethirothrips anacardii; 245, Diplacothrips borgmeieri III-IV; 246, Machatothrips biuncinatus III-IV;
247, M. antennatus III-IV; 248, Dichaetothrips brevicollis anapleural suture.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
249 I / * 252
I / II / / 1 zo<f tr\ ^x
256
143
Figs 249-256 Macrothripina. 249, Ethirothrips indicus (Bagnall) III-IV; 250, E. stenomelas (left); 251,
Aesthesiothrips jatrophae; 252, Tarassothrips sp. indet. III-IV; 253, T. akritus; 254, Dichaetothrips
brevicollis holotype III-V; 255, D. secular III-IV; 256, D. okajimai.
144
L. A. MOUND AND J. M. PALMER
257
259
258
261
260
262
Figs 257-262 Elaphrothripina. 257, Mecynothrips acanthus; 258, M. pugilator; 259, M. kraussi; 260, M.
atratus (zuluensis holotype) pronotum; 261, M. hardy i; 262, M. kraussi.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
263
264
145
265
266
267
Figs 263-267 Elaphrothripina. 263, Anactinothrips vigilans; 264, Ophthalmothrips ? priesneri mac. from
Kenya; 265, O. ? priesneri mic. from Tanzania; 266, O. amyae mac.; 267, O. pomeroyi mac. from
Malawi.
146
L. A. MOUND AND J. M. PALMER
270
Figs 268-272 Elaphrothripina. 268, Hartwigia tumiceps; 269, Tiarothrips subramanii; 270, Elaphrothrips
palustris; 271, E. greeni (bouvieri holotype); 272, E. spiniceps.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
147
273
Figs 273-276 Elaphrothripina. 273, Malesiathrips guamensis; 274, M. malayensis; 275, Lamillothrips
typicus; 276, Dermothrips hawaiiensis .
148
L. A. MOUND AND J. M. PALMER
277
283
282
Figs 277-284 Elaphrothripina. 277, Dinothrips spinosus $ ; 278, D. monodon d" mesothoracic spiracular
process; 279, D. spinosus small cf mesothoracic spiracle; 280-283, anapleural sutures of (280) Male-
siathrips malayensis; (281) Dermothrips hawaiiensis; (282) Elaphrothrips laevicollis; (283) Hartwigia
tumiceps; 284, Elaphrothrips jacobsoni cf pronotum.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
149
285
Figs 285-292 Elaphrothripina. 285, Elaphrothrips femoralis large cf forefemur; 286, E. palustris C?
forefemur; 287, Mecynothrips priesneri cf foreleg; 288, M. simplex cf foreleg; 289, Ophthalmothrips ?
priesneri; 290, Dermothrips hawaiiensis; 291, Malesiathrips guamensis; 292, M. malayensis.
150
L. A. MOUND AND J. M. PALMER
296
293
Figs 293-299 Elaphrothripina. 293, Elaphrothrips denticollis (productus holotype) cf foreleg; 294,
Hartwigia tumiceps mac. tergite III; 295, Anactinothrips vigilans tergite III; 296, Malesiathrips malayen-
sis mac. tergite IV; 297, Mecynothrips simplex tergite III; 298, Dlnothrips sumatrensis tergite III; 299,
Elaphrothrips bakeri O" tergite III.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
151
300
Figs 300-312 Pelta of Elaphrothripina species. 300, Dermothrips hawaiiensis; 301, Tiarothrips subrama-
nii; 302, Ophthalmothrips amyae; 303, O. longiceps; 304, Hartwigia tumiceps; 305, Malesiathrips
malayensis; 306, Mecynothrips atratus; 307, Elaphrothrips laevicollis; 308, E. denticollis; 309, E.
spiniceps; 310, Dinothrips sumatrensis; 311, Anactinothrips vigilans; 312, Lamillothrips typicus.
152
L. A. MOUND AND J. M. PALMER
313
314
317
315
316
Figs 313-317 Idolothripina. 313, Meiothrips annulipes; 314, Bactrothrips pitklni $ paratype; 315,
Bacillothrips nobilis; 316, Bactrothrips idolomorphus cf from Java; 317, Idolothrips spectrum $.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
153
318
320
Figs 318-322 Idolothripina. 318, Megalothrips andrei cf holotype; 319, Ceuthothrips timuqua; 320,
Cylindrothrips niger holotype; 321, Megathrips lativentris; 322, Lasiothrips perplexus holotype.
154
L. A. MOUND AND J. M. PALMER
323
324
325
326
327
Figs 323-327 Idolothripina. 323, Meiothrips annulipes <3" tergite V; 324, Megalothrips andrei cf holotype;
325, Idolothrips dissimilis cT; 326, Megathrips lativentris $ mac. tergite IV; 327, Meiothrips annulipes
anapleural suture.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
155
328
330
329
331
337
338
332
333
334
335
336
Figs 328-338 Idolothripina. 328-336, pelta of (328) Ceuthothrips timuqua; (329) Megalothrips andrei;
(330) Bacillothrips longiceps; (331) Megathrips lativentris; (332) Bactrothrips idolomorphus; (333) E.
pitkini 9 mac. ; (334) Idolothrips nativus; (335) Cylindrothrips niger (turneri holotype); (336) Meiothrips
annulipes; 337, Megalothrips andrei; 338, Ceuthothrips timuqua.
L. A. MOUND AND J. M. PALMER
Figs 339-346 Hystricothripina. 339, Cyphothrips dorsalis cf ; 340, Hybridothrips oneillae holotype; 341,
Holurothrips ornatus; 342, Zeuglothrips echinus holotype; 343, Saurothrips assai; 344, Hystricothrips
phasgonura cf; 345, Actinothrips fraterculus 9; 346, Atractothrips bradleyi.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
157
347
Figs 347-355 Hystricothripina. 347, Paractinothrips peratus; 348, P. peratus cf foreleg; 349, Azeugma-
tothrips rectus; 350, Zeugmatothrips cinctus; 351, Z. priesneri; 352, Cyphothrips dorsalis C? metanotum;
353, Actinothrips femoralis anapleural suture; 354, Holurothrips ornatus anapleural suture; 355,
Paractinothrips peratus anapleural suture.
158
L. A. MOUND AND J. M. PALMER
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
159
368
369
370
374
371
372
373
Figs 368-374 Hystricothripina. 368, Saurothrips assai tergite III; 369, Paractinothrips peratus tergite III;
370, Hystricothrips phasgonura cf VII- VIII; 371-373, mouth cone and prosternal plates of (371)
Zeugmatothrips priesneri; (372) Neatractothrips macrurus; (373) Paractinothrips peratus; 374, Azeugma-
tothrips rectus tube.
160
L. A. MOUND AND J. M. PALMER
375
Figs 375-384 Pelta of Hystricothripina species. 375, Hystricothrips phasgonura; 376, Zeuglothrips
echinus; 377, Neatractothrips macrurus; 378, Paractinothrips peratus; 379, Zeugmatothrips priesneri;
380, Z. hispidus; 381, Saurothrips assai; 382, Azeugmatothrips rectus; 383, Holurothrips ornatus; 384,
Cyphothrips dorsalis.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
**C2?
161
387
390
393
Figs 385-393 Docessissophothripini. 385, Maxillata sp. indet from Jamaica; 386, ? Maxillata from Ghana;
387, Tropothrips dampfi holotype (setae omitted); 388, Holothrips peruvianus $ paratype; 389, H.
zimmermanni holotype; 390, H. procerus $ paratype; 391, Asemothrips picturatus cf sternite IV; 392,
Symphyothrips nr. punctatus tube; 393, Holothrips (? Lathrobiothrips} sp. indet. tube.
162
L. A. MOUND AND J. M. PALMER
394
395
Figs 394-398 Docessissophothripini. 394, Docessissophothrips ampliceps cf holotype; 395, D. major $
holotype; 396, Oidanothrips frontalis 9 holotype; 397, Symphyothrips near punctatus from Panama;
398, Abiastothrips soror.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
399 n 400
163
402
401
403
Figs 39SM03 Docessissophothripini. 399, 400, outline sketches of (399) Holothrips typicus (Anantha-
krishnan) holotype; (400) H. semiflavus holotype; 401, H. typicus (Bagnall) holotype; 402, H. ambitus;
403, H. ingens $ holotype.
164
L. A. MOUND AND J. M. PALMER
404
\
405
407
406
408
409
,-"",A
410
411
Figs 404-413 Docessissophothripini. 404, Holothrips ingens pelta; 405, Docesslssophothrips ampliceps
pelta; 406, Symphyothrips near punctatus pelta; 407, Holothrips ingens; 408, Oidanothripsfrontalis; 409,
Holothrips peruvianus; 410, Symphyothrips near punctatus; 411, Asemothrips finlayi; 412, Holothrips
ingens anapleural suture; 413, Docessissophothrips ampliceps anapleural suture.
GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA
Index
165
Synonyms are in italics; original generic combinations are indicated for identical specific epithets.
abditus 39
aberrans Adelothrips 94
aberrans Symphyothrips 96
Abiastothrips 91
acaciae 31
Acallurothrips 40
Acanthinothrips 76
acanthomerus 65
acanthus Cryptothrips 56
acanthus Kleothrips 70
achaetus 68
Acrothrips 69
acti 42
Actinothrips 81
Aculeathrips 79
acuticornis 39
acutulus 39
acutus Adelothrips 94
acutus Allothrips 31
addendus 65
adelos 94
Adelothrips 93
Adiaphorothrips 58
adiaphorus 51
adventor 57
aemulus 37
aeneus 37
Aesthesiothrips 51
aethiopiae 65
aethiopicus 69
affinis Coenurothrips 45
affinis Dinothrips 64
affinis Idolothrips 65
africana Adelothrips 24
africanus Allothrips 31
africanus Boloadelothrips 36
africanus Idolothrips 65
africanus Zeugmatothripoides 85
agama 70
agasthya 57
Agnostothrips 93
agrestis 28
Akleothrips 69
akritus 61
alakaiensis 31
albiceps 30
albomaculata 37
albosignatus 37
albospinosus 65
alifanensis 96
alius 47
alluaudi Bactrianothrips 73
alluaudi Eidothrips 73
Allidothrips 30
Allopisothrips 31
Allothrips 31
alticola 39
amabilis 37
amazonicus 65
ambitus 94
amneius 28
amoenus 65
ampliceps 92
amplus Adiaphorothrips 50
amplus Holothrips 94
amplus Pygothrips 41
amyae 71
anacardii 57
Anactinothrips 63
Anaglyptothrips 34
anahuacensis 44
andrapterus 65
andrei Megalothrips 78
andrei Oedaleothrips 36
angulatus 91
angustatus 66
angusticeps 66
angustifrons 66
angustus 23
anici 28
animus 78
annulatus 80
annulipes Acanthinothrips 80
annulipes Gastrothrips 45
annulipes Zeugmatothrips 88
annuus 92
anodon 64
anolis 39
antennalis 66
antennatus Adiaphorothrips 58
antennatus Lophothrips 63
aoristus 47
Apelaunothrips 89
approximatus 68
apterus 42
arenarius 36
armatus Apelaunothrips 89
armatus Diceratothrips 44
armatus Idolothrips 66
artocarpi Bolothrips 47
artocarpi Machatothrips 58
Ascania 23
Asemothrips91
aspericauda 94
assai 87
assimilis 65
associatus 94
aterrimus 73
athletes 66
Atractothrips 82
166
atratus 70
atrispinis Bactrothrips 73
atrispinis Elaphrothrips 68
aureus 31
australicus 42
australiensis Bolothrips 48
australiensis Gastrothrips 57
australis 94
Azeugmatothrips 82
aztecus 66
Bacillothrips 72
Bactrianothrips 72
Bactridothrips 72
Bactrothrips 72
badiicornis 88
badiipes 88
badius Cryptothrips 42
badius Pygothrips 41
bagnalli Bacillothrips 72
bagnalli Cryptothrips 36
bagnalli Mecynothrips 70
bagnallianus 66
baileyi 37
bakeri 66
bancoensis 73
Baphothrips 89
barretti Phaulothrips 28
barretti Scotothrips 57
Barythrips 28
beesoni Dichaetothrips 57
beesoni Elaphrothrips 66
bellulus 94
bennetti 44
berlandi 73
bhowalii 89
bicolor Adelothrips 94
bicolor Allothrips 31
bicolor Apelaunothrips 89
bicolor Compsothrips 37
bicolor Oedaleothrips 37
bicolor Phloeothrips 36
bicolorisetosus 59
bicornis 44
bidens 50
biformis 42
bilineatus 66
biminianus 31
bipartitus 94
bispinosus 88
biuncinatus 58
blatchleyi 66
bogong 23
Boloadelothrips 35
Bolothrips 35
bonannii 78
bondari 81
borgmeieri Anactinothrips 63
borgmeieri Diplacothrips 54
INDEX
borgmeieri Elaphrothrips 66
borgmeieri Tropothrips 97
borgmeieri Zeugmatothrips 88
Botanothrips 35
bottegii 66
bournieri 31
bouvieri 67
brachypes 66
brachyura Anactinothrips 63
brachyurus Cryptothrips 36
brachyurus Elaphrothrips 66
bradleyi Atractothrips 82
bradleyi Oedaleothrips 37
brasilianus 31
brasiliensis Elaphrothrips 66
brasiliensis Hoplothrips 39
brasiliensis Oedaleothrips 37
brasiliensis Polyphemothrips 92
bratleyi 94
braueri 58
breviceps Cryptothrips 23
breviceps Bagnall Dicaiothrips 66
breviceps Priesner Dicaiothrips 67
breviceps Oedemothrips 48
breviceps Phoxothrips 71
breviceps Pygothrips 41
brevicollis Coenurothrips 45
brevicollis Dichaetothrips 53
brevicollis Oedemothrips 47
brevicornis Bolothrips 36
brevicornis Dicaiothrips 66
brevicornis Diceratothrips 57
brevis Adiaphorothrips 57
brevis Siphonothrips 79
brevisetosus 57
brevitubus 73
brimblecombei 23
brittoni 23
brochotus 27
brunneipennis 66
brunneitarsis 67
brunneus Diopsothrips 41
brunneus Oedaleothrips 37
Bryothrips 31
buccalis 94
bucculentus 73
buffai Caudothrips 73
buffai Machatothrips 59
burroughsi 58
bursarius 94
caliginosus 96
callipygus 30
callipus 39
camelus 52
campestris 37
Campulothrips 42
capensis 66
capitalis 39
INDEX
167
Capnothrips 23
capricornis 42
carayoni 66
carbonarius 23
caribbeicus 94
Carientothrips 42
carve ri 48
casuarinae 42
caudatus Adelothrips 94
caudatus Allothrips 94
caudatus Phaulothrips 28
Caudothrips 72
celebensis 64
Celidothrips 51
celosia 59
Cervothrips 72
cestosa 48
Ceuthothrips 75
ceylonicus Dracothrips 70
ceylonicus Oedemothrips 46
championi 67
chandana 68
chiapensis 81
cinctus Allothrips 31
cinctus Bolothrips 36
cinctus Zeugmatothrips 88
cingulatus 36
citricauda 33
citriceps 39
citricornis 94
claripennis Cryptothrips 48
daripennis Dichaetothrips 57
darispinis Bagnall Elaphrothrips 66
darispinis Priesner Elaphrothrips 68
clavicornis 33
clavipes 52
Cleistothrips 22
Cnemidothrips 52
Cnestrothrips 58
Coenothrips 91
Coenurothrips 45
cognatigrandis 66
collaris 46
collessi 84
combustipes 91
Compsothrips 36
concordiensis 96
congoensis Bactrothrips 73
congoensis Elaphrothrips 66
congoensis Oedaleothrips 37
conicura Adelothrips 94
conicurus Elaphrothrips 66
conifer 41
coniferarum 66
coniger 66
connaticornis 94
conocephalus71
consimilis 89
constrictopeltatus 66
Cordylothrips 93
coreanus 66
cornutus Adelothrips 94
cornutus Diceratothrips 44
coronatus 67
corticis 92
corticosus 59
corvus 39
costalimai 66
cracens 94
Cradothrips 65
crassiceps 64
crassus 58
Cratothrips 91
cristatus 63
Cryptothrips 22
csiro 23
cubensis 44
cuneatus 92
curvidens 39
curvipes 66
cybele 39
Cylindrothrips 75
Cyphothrips 83
dallatorensis 68
dammermani 58
dampfi Docessissophothrips 97
dampfi Elaphrothrips 66
dampfi Myrmecothrips 37
Dasythrips 81
debilis 95
decipiens 66
decorus 59
Decothrips 55
defectus 66
delamarei 73
delicatus 44
delmasi 78
denticollis 66
denticulatus 42
dentipes 36
dentis 36
depokensis 66
Dermothrips 63
Derothrips 70
devius 66
Dexiothrips 90
diabolus 59
Diaphorothrips 52
Dicaiothrlps 64
Diceratothrips 43
Dichaetothrips 52
differ ens 57
diffidlis 48
dimidiatus 48
Dinothrips 64
Diopsothrips 40
Diplacothrips 54
168
Diplochelaeothrips 28
dissimilis 76
distans 66
distasmus 57
distinctus 65
distinguendus 63
divergens 73
diversicolor 46
diversus 57
Docessissophothrips 92
dolichos 52
dominicanus 46
dorsalis 83
dotatus 92
doulli 48
dracon 57
Dracothrips 69
drepanatus 66
drepanifer 66
dubius 59
dugdalei 35
eastopi 48
echinus 87
edouardi 66
Egchocephalothrips 76
Eidothrips 72
Elaphoxothrips 64
Elaphridia 55
Elaphridothrips 64
Elaphrothrips 64
elegans Siphonothrips 79
elegans Zactinothrips 87
elephas 57
Elgonima 45
Embothrips 32
embotyi 36
Empresmothrips 91
Emprosthiothrips 23
Endacnothrips 43
enormis 96
epallelus 23
eranthemi 68
Erythrinothrips 93
Ethirothrips 54
eucharis 94
Eucoenothrips 91
Eulophothrips 43
eupatorii 44
Eurynotothrips 55
eurytis 25
falcatus Dicaiothrips 66
falcatus Nesothrips 39
fallax Coenothrips 91
fallax Elaphrothrips 66
fasciatipennis 77
fasciolatus 41
faurei Elaphrothrips 66
INDEX
faurei Fulgorothrips 71
Faureothrips 32
femoralis Actinothrips 81
femoralis Apelaunothrips 89
femoralis Klinothrips 66
femoralis Machatothrips 96
femoralis Zeugmatothrips 88
fenestralis 94
fijiensis Bolothrips 42
fijiensis Cryptothrips 57
fijiensis Gastrothrips 46
fijiensis Paracryptothrips 57
finlayi 91
firmus 57
flaviceps 33
flavipes Cryptothrips 79
flavipes Idolothrips 66
flavitibia 42
flavus Cryptothrips 23
flavus Diopsothrips 41
flavus Hoplothrips 94
fodinae 48
folii 91
Formicothrips 36
formosanus 71
formosensis 48
formosus 94
fortis 30
foveicollis 66
fraterculus 81
frontalis 96
froudei 91
fructuum 46
Fulgorothrips 70
fulmeki Dinothrips 64
fulmeki Elaphrothrips 67
fulvicauda 39
fulviceps 40
fumidus 94
fumipennis 40
fungivorus 57
furcatus 73
furcifer 64
fuscatus 40
fuscicauda Cryptothrips 34
fuscicauda Gastrothrips 46
fuscus Anactinothrips 63
fuscus Megalothrips 78
fuscus Phaulothrips 28
fuscus Rhaebothrips 48
gabonensis 89
gaboniensis 67
Galactothrips 45
gardneri 64
gargantua Acrothrips 70
gargantua Actinothrips 81
Gastrothrips 38
genaspinosus 67
INDEX
169
gibbifer 63
gigans 70
giganteus 70
gigas 57
gilvipes 36
giraulti 57
gloveri 57
gnidiicolus 67
gnomus 33
Goetothrips 38
goliath 70
gracilis Campulothrips 42
gracilis Elaphrothrips 67
gracilis Zeugmatothrips 88
gradatus 34
graminicola 94
graminis 37
grandicauda 46
grandis Bactrothrips 73
grandis Dicaiothrips 67
grandis Adelothrips 94
graphidura 63
graveleyi 68
gravis 67
grayi 42
greeni 67
greensladei 31
guachichilis 67
guamensis 69
guerreronsis 84
guineaensis 73
guineensis 73
gurdus 40
gynaecoides 66
halidayi 76
hambletoni 39
hamiltoni 46
hamipes 52
hammockensis 94
handlirschii 63
haplodon 59
hardyi 70
hargreavesi 74
harti Diceratothrips 44
harti Syncerothrips 40
Hartwigia 68
hawaiiensis Dermothrips 64
hawaiiensis Nesothrips 48
hemidiscus 48
Heptathrips 23
Herathrips 58
hercules 44
herricki 67
hesperus 73
heterocerus 40
heveae 59
hibisci 57
hilaris 46
hispidus 88
Holmiella 93
Holothrips 92
Holurothrips 83
honoris 73
hoodi Bactrothrips 73
hoodi Gastrothrips 46
hoodi Leptogastrothrips 37
hoodi Zeugmatothrips 88
hookeri 37
horridus 44
hubbeli 37
Hybridothrips 84
Hylothrips 68
Hystricothrips 84
hystrix Dicaiothrips 67
hystrix Hystricothrips 85
icarus 36
idolomorphus 73
idolothripoides 22
Idolothrips 76
Illinothrips 38
imbecilla 70
imitator 66
impensus 67
incisus 42
indagator 67
indica Loyolaia 38
indicus Agnostothrips 94
indicus Allothrips 31
indicus Dichaetothrips 57
indicus Machatothrips 59
indicus Mesothrips 57
indicus Nesothrips 57
indicus Oedaleothrips 37
indicus Philothrips 89
indicus Uredothrips 57
inermis Megathrips 79
inermis Panurothrips 74
inermis Paracryptothrips 57
inferorum 44
infirma 77
ingens 94
innocens 70
inquilinus Kaleidothrips 28
inquilinus Neosmerinthothrips 46
insignis Elaphrothrips 67
insignis Lathrobiothrips 94
insolitus 60
insperatus 67
insularis Cryptothrips 36
insularis Elaphrothrips 67
intermedius 59
intonsus 40
intrepidus 57
invalida 77
io57
ipomoeae 48
170
INDEX
Ischnothrips 93
Ischyrothrips 58
Isopterothrips 38
italicus 36
jacksoni 37
jacobsoni Dinothrips 64
jacobsoni Elaphrothrips 67
jacotguillarmodi 67
janus 26
japonicus Apelaunothrips 89
japonicus Cryptothrips 42
jatrophae 51
jeanneli 66
judithae 41
juglandis 64
junctus 94
kabandha 28
Kaleidothrips 27
kanoi 70
karimonensis 70
karnyi Elaphrothrips 66
karnyi Machatothrips 58
karnyi Mesothrips 57
karnyi Neosmerinthothrips 48
kawamurai 74
kellyanus 77
kemneri 64
kenyensis 74
Kleothrips 69
Klinothrips 64
Krlnothrips 72
kraussi Diaphorothrips 52
kraussi Mecynothrips 70
lacerta 70
lacertina 77
laevicollis 67
laingi 74
Lamillothrips 68
lamottei 74
lanei 94
Lasiothrips 77
lata 23
latapennis 57
Lathrobiothrips 92
laticeps Cryptothrips 28
laticeps Docessissophothrips 61
laticeps Elaphrothrips 67
laticeps Oedemothrips 48
laticornis 67
lativentris Phloeothrips 79
lativentris Rhaebothrips 48
lativerticis 91
latus 41
lawrencei 52
leeuweni 84
leios 90
lentus 59
Leptogastrothrips 36
lesnei Fulgorothrips 71
lesnei Megathrips 73
Leurothrips 36
leveri 48
levidens 74
levis 67
lieni 90
linearis 72
litoreus 36
loisthus 42
longfellowi 91
longicauda Paxillothrips 64
longicauda Symphyothrips 96
longiceps Cryptothrips 23
longiceps Docessissophothrips 72
longiceps Megalothrips 72
longiceps Idolothrips 67
longiceps Pygothrips 30
longiceps Pyrgothrips 71
longicornis Actinothrips 81
longicornis Symphyothrips 96
longidens 69
longipes Diceratothrips 44
longisetis Anactinothrips 63
longisetis Bactrothrips 74
longisetis Diceratothrips 57
longispina Phloeothrips 79
longispinis Elaphrothrips 67
longitubus 28
longiventris 74
longus 57
Lophothrips 63
louisianae 41
Loyolaia 38
luctator 33
lucyae 94
luridus 90
luteus Bactrothrips 74
luteus Polyphemothrips 94
mabirensis 67
macateei 67
Machatothrips 58
macropteryx 74
Macrothrips Bagnall 59
Macrothrips Buffa 36
macrura Adelothrips 94
macrurus Acallurothrips 41
macrurus Atractothrips 85
maculipennis 90
madagascariensis Cryptothrips 57
madagascariensis Elaphrothrips
67
madagascariensis Ethirothrips 57
madrasensis 90
magnetis 42
INDEX
171
magnicauda 30
magnifica Ascania 24
magnificus Docessissophothrips
28
magnus Cryptothrips 48
magnus Elaphrothrips 67
magnus Mecynothrips 70
magnus Oidanothrips 96
mahensis 67
major Cryptothrips 23
major Docessissophothrips 92
major Rhaebothrips 48
malaccae 48
malayensis Apelaunothrips 90
malayensis Dicaiothrips 67
malayensis Malesiathrips 69
Malesiathrips 69
malgassus 74
malloti 64
mameti 50
mamillicauda 41
mandiocae 40
marginata 77
marginipennis 63
maritimus 23
maroccanus 37
marshalli 46
mauli 40
mavromoustakisi 33
Maxillata 95
maynei 67
Mecynothrips 69
medioflavus 90
medius 67
megacephalus Allothrips 31
megacephalus Polyphemothrips
96
Megalomerothrips 43
Megalothrips 77
Megathrips 78
meinerti 63
Meiothrips 79
melinus 48
menoni 80
mentawaiensis 66
meridionalis Diceratothrips 57
meridionalis Pseudocryptothrips
34
Mesopotamothrips 96
metulicauda 41
mexicanus 31
micidus 67
microacanthomerus 67
mikrommatos 30
milleforme 46
minor Coenurothrips 47
minor Mecynothrips 70
minor Polyphemothrips 94
mirandus 94
miscanthicola 71
miskoi 42
mjobergi 42
mockfordi 82
modestus 87
mongolicus 40
monochaetus 81
monodon 64
monstrosus 76
montanus Allothrips 31
montanus Apelaunothrips 90
montanus Machatothrips 58
monticola 40
morikawai 84
moultoni 74
mucronatus 66
multidens 58
mumbaca 88
Myrmecothrips 36
natalensis 74
nativus Adiaphorothrips 58
nativus Idolothrips 74
Neatractothrips 85
needhami 30
neivei 57
neodampfi 67
neoleonensis 67
neolongiceps 67
Neosmerinthothrips 45
nepalensis Adelothrips 94
nepalensis Meiothrips 80
Nesidiothrips 47
Nesothrips 47
niger Bolothrips 48
niger Cryptothrips 57
niger Cylindrothrips 76
niger Elaphrothrips 67
niger Emprosthiothrips 23
niger Ischyrothrips 57
niger Megalothrips 79
niger Zeugmatothrips 88
nigricauda 44
nigricornis Anactinothrips 63
nigricornis Idolothrips 67
nigricornis Liothrips 57
nigripennis 90
nigripes Bactrothrips 74
nigripes Elaphrothrips 67
nigripes Tropothrips 97
nigrisetis Gastrothrips 46
nigrisetis Rhaebothrips 48
nigrita 94
nigrospinosus 68
nitidus 67
nobilis 72
nogutti 41
notabilis 67
noumeae 39
172
INDEX
novus 68
nox57
nubillicauda 31
oahuensis 48
obrieni 82
obscuricornis Diceratothrips 44
obscuricornis Elaphrothrips 66
obscuricornis Oedemothrips 48
obscurus 57
ocelloides 49
ocularis 90
oculatoides 67
oculatus 67
oeceticola 40
Oedaleothrips 36
Oedemothrips 47
Oidanothrips 95
okajimai 53
oleriae 48
Ommatidothrips 51
oneillae Actinothrips 84
oneillae Compsothrips 37
Ophidothrips 63
Ophthalmothrips 70
orangiae 67
ornatus Holurothrips 84
ornatus Rhopalothrips 27
Ozothrips 24
padewiethi 79
Palinothrips 64
pallicornis 58
pallidicrus 74
pallidior 44
pallidulus 88
pallipes Cryptothrips 36
pallipes Empresmothrips 91
palmarum 94
palustris 67
pampicola 37
Panceratothrips 92
papuensis 59
Paraclinothrips 65
Paracryptothrips 55
Paractinothrips 85
paradampfi 67
Paragastrothrips 39
Parallothrips 32
parallelus 67
parvidens Bactrothrips 74
parvidens Gastrothrips 46
parvidens Kleothrips 70
parvus 67
paucidens 58
paulistarum 46
Paxillothrips 64
pedalisSl
pedicillus 43
Pelinothrips 27
Peltariothrips 59
peltatus 88
penicillatus 40
penicollis 69
pensus 90
peratus 86
Percipiothrips 55
Percnothrips 39
pericles 94
perplexus 77
persimilis 44
peruvianus 94
peruviensis 67
Phacothrips 49
phaeura 94
Pharetrothrips 39
phariacus 23
phasgonura 85
Phaulothrips 27
Phoxothrips 69
piccioli 79
piceus 54
picticornis Diceratothrips 44
picticornis Gastrothrips 46
picticornis Megalothrips 78
pictilis 43
picturatus 91
pillichellus 31
pilosus 61
pitkini 74
plaumanni 46
polychaetus 81
Polyphemothrips 92
Polytrichothrips 60
pomeroyi 71
Pongola 96
portentosus 28
potosiensis 97
powelli 67
pratensis 36
priesneri Bactrothrips 74
priesneri Cratothrips 91
priesneri Elaphrothrips 67
priesneri Fulgorothrips 71
priesneri Maxillata 95
priesneri Mecynothrips 70
priesneri Zeugmatothrips 88
Priesneriana 28
Priesneriella 32
princeps 44
priscus 26
Probolothrips 39
procer 67
procerus Gastrothrips 40
procerus Holothrips 94
productus 66
prolixus 31
propinquus Bactridothrips 74
INDEX
173
propinquus Dicaiothrips 68
propinquus Oedemothrips 48
prospector 68
proteus 39
proturus 40
proximus Dicaiothrips 68
proximus Pseudocryptothrips 34
Pseudocryptothrips 33
Pseudoeurhynchothrips 50
pueblae 40
pugilator 70
punctatus Cladothrips 28
punctatus Symphyothrips 97
Pygidiothrips 32
Pygothrips 28
pygus 30
Pyrgothrips 70
quadraticeps 42
quadrituberculatus 74
quadrudentatus 59
querci 37
quercus 94
rachiphilus 36
ramamurthii 37
ramuli 94
rangi 48
rectangularis 23
recticeps 37
rectus 82
reedi 43
regalis 81
reticulatus Cryptothrips 32
reticulatus Symphyothrips 97
reuteri 37
rex 61
Rhaebothrips 47
rhizophorae 48
rhopaloides 91
richardsi 97
ritchianus 73
robustus Adelothrips 94
robustus Eulophothrips 44
robustus Nesothrips 46
rossi 38
rufianalis 96
ruficauda 40
ruficaudis 24
rugicauda 30
ruidis 94
sakimurai 44
salicis 23
satanas 30
Saurothrips 86
sauteri 23
schaferi 36
schaubergeri 91
schottii 68
schoutedeni 68
schuhi 78
schultzei 68
Scotothrips 55
sculpticauda 30
sculptilis 94
secus 66
secutor 53
semiflavus Agnostothrips 94
semiflavus Bolothrips 48
seminole 33
semirufus 43
sensitivus 68
separatus 68
serex 70
serraticornls 73
seticeps 45
setldens 57
setigenis 44
seychellensis Cryptothrips 48
seychellensis Dicaiothrips 68
shavianus 30
Sibylla 28
silvae 91
silvaticus 59
silvicola 63
similis 48
simplex Adiaphorothrips 58
simplex Mecynothrips 70
simplicidens 59
sinensis 37
Siphonothrips 78
sismondini 56
sjostedti 56
skwarrae 94
snodgrassi Elaphrothrips 68
snodgrassi Mecynothrips 70
solomoni 69
sordidatus 23
soror 91
speciosissimus 95
spectrum 76
spinicauda 42
spiniceps 68
spiniprivus 68
spinosus Diaphorothrips 52
spinosus Elaphrothrips 68
spinosus Ischyrothrips 64
spinosus Megalothrips 78
splendidus 94
sporophagus 94
Sporothrips 50
stannardi Allothrips 31
stannardi Polyphemothrips 95
stenocephalus 68
stenomelas 57
Stinothrips 93
stygicus 40
174
subtilis 95
subulatus 40
subrammanii 71
sumbanus 66
sumatranus 37
sumatrensis 64
surinamensis 68
sybarita 57
Symphyothrips 96
Syncerothrips 39
Synkleothrips 69
taiwanus 70
takahashii 70
Tarassothrips 61
tasmani 90
Telothrips 79
tener 68
tenuipennis 40
tenuiceps 92
terrestris 40
terrigena 11
Tetraceratothrips 27
texanus 40
thevetil 52
thomasi 33
thomassetti 57
Tiarothrips 71
tibialis Phloeothrips 79
tibialis Polyphemothrips 92
tibialis Uredothrips 57
timidus Diceratothrips 44
timidus Megathrips 79
timuqua 75
timur 37
tirumalaiensis 57
Titanothrips 21
titschacki Bactrothrips 74
titschacki Docessissophothrips 95
tonnoiri 24
transvaalensis 68
travassosi 92
trichaetus 81
tricolor Allidothrips 31
tricolor Baphothrips 90
trinidadensis 57
tristis 37
Tropothrips 97
tuberculatus Cryptothrips 36
tuberculatus Idolothrips 68
tubversicolor 33
tumiceps 68
tumidus 95
turbinatus 40
turneri 76
tuxtlae 97
tuzetae 33
INDEX
typicus Ischnothrips 95
typicus Panceratothrips 95
typicus Lamillothrips 69
umbricola 95
unguipes 52
unicolor 68
uniformis 68
uptoni Cryptothrips 28
uptoni Phaulothrips 28
uredinis 57
Uredothrips 55
usitatus 57
uzeli 37
validipennis 44
validus 47
variipes 46
variispinis 68
varius 36
vesper 43
vigilans 63
villicornis 92
virgulae 57
vittipennis 68
vitreipennis 57
vitulus 69
vuilleti 28
wallacei 70
walteri 37
watsoni 31
williamsi Cryptothrips 23
williamsi Dichaetothrips 44
wolcotti 44
woytkowskyi Lathrobiothrips 95
woytkowskyi Polyphemothrips 92
xanthopus 95
xosa 40
xylebori 46
yanchepi 48
yosemitae 37
yuasai 48
yupanqui 92
Zactinothrips 87
zeteki 30
zetetis 68
Zeuglothrips 87
Zeugmatothripoides 84
Zeugmatothrips 88
zimmermanni 95
zondagi 48
zuluensis 70
British Museum (Natural History)
Chance, change & challenge
Two multi-author volumes from one of the foremost scientific institutions in the world.
General Editor: P. H. Greenwood
The Evolving Earth
Editor: L. R. M. Cocks
The Evolving Biosphere
Editor: P. L. Forey
In the first volume, The Evolving Earth, twenty scientists have been asked to review the present
state of knowledge in their particular field, ranging from the origin of the Earth, through ocean
sediments and soils to continental drift and palaeogeography.
In the companion volume, The Evolving Biosphere, museum scientists have chosen an
evolutionary concept — speciation, coevolution, biogeography etc. and related this to the group
of animals or plants in which they are specialising. Thus beetles and birds exemplify sympatric
and allopatric speciation, butterflies mimicry and certain fishes explosive evolution.
In both volumes the text is supplemented by over one hundred specially commissioned pieces of
two-colour artwork.
These two books will be invaluable to all sixth-form and undergraduate biology and geology
students.
The Evolving Earth: 276 x 219 mm, 280pp, 138 line illustrations, 42 halftones
The Evolving Biosphere: 276 x 219 mm, approx. 320pp, 133 line illustrations
Published: May 1981
Co-published by the British Museum (Natural History), London and Cambridge University
Press, Cambridge.
Titles to be published in Volume 46
The generic and tribal classification of spore-feeding Thysanoptera (Phlaeothripidae: Idolo-
thripinae).
By L. A. Mound & J. M. Palmer.
A revision of the Afrotropical mole-crickets (Orthoptera: Gryllotalpidae).
ByB. C. Townsend.
Key to the genera of galerucine beetles of New Guinea, with a review of Sastra and related new
taxa (Chrysomelidae).
By Sharon L. Shute.
The Afrotropical dacetine ants (Formicidae).
By Barry Bolton.
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd. , Dorchester
Bulletin of the
British Museum (Natural History)
A revision of the Afrotropical mole-crickets
(Orthoptera: Gryllotalpidae)
B.C. Townsend
Entomology series
Vol 46 No 2 26 May 1983
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,
and an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
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specialists from elsewhere who make use of the Museum's resources. Many of the papers are
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Parts are published at irregular intervals as they become ready, each is complete in itself,
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••.
"
World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
©Trustees of the British Museum (Natural History), 1983
2 6 MAY 1983
] J
The Entomology series is produced under the general editorship of the
Keeper of Entomology: Laurence A. Mound
Assistant Editor: W. Gerald Tremewan
ISSN 0524-6431
British Museum (Natural History)
Cromwell Road
Entomology series
Vol46No2ppl75-203
xby' *«^
r^ «M«M*
2 6 MAY 1985
A revision of the Afrotropical mole-crickets
(Orthoptera: Gryllotalpidae)
B. C. Townsend ^
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW75BD
Contents
Synopsis 175
Introduction 175
Material 176
Taxonomic characters 176
Methods 177
Gryllotalpidae 178
Key to the subfamilies and genera 179
Gryllotalpa Latreille 179
Synonymic list of the Afrotropical species 180
Non- Afrotropical species of Gryllotalpa also covered 180
Key to the Afrotropical species 180
Descriptions of the Afrotropical species 182
Acknowledgements 201
References 201
Index 203
Synopsis
The 12 Afrotropical species of Gryllotalpidae, all members of Gryllotalpa, are revised, with six new
species, one new synonymy, and one species revalidated. The songs of five of these species, and that of an
Oriental species, are described for the first time. Keys are provided to the two subfamilies and five genera
of Gryllotalpidae. Gryllotalpa minuta Burmeister, previously thought to occur in Africa, is shown to be
absent there.
Introduction
The Gryllotalpidae, or mole-crickets, occur throughout the tropical and warmer temperate
regions of the world. They are closely related to the Gryllidae, the true crickets, from which they
differ mainly in being highly specialised for a subterranean existence. The fore legs are modified
for digging, and bear two to four strongly sclerotised dactyls, and the body is covered in a dense
mat of hair. Mole-crickets dig a complex of burrows within which they live, feed, sing, mate and
breed, and which includes a nest chamber and a special singing burrow. They fly only rarely,
usually to search for a mate. Specimens are most commonly taken at night during such flights,
and often a high proportion of those so captured are females. The diet of mole-crickets varies
according to the species (Matheny, 1981); they may be mainly carnivorous, mainly vegetarian or
truly omnivorous. The life cycles of all the African species are totally unknown.
Several species of Gryllotalpa, in common with those of other gryllotalpid genera, become
serious crop pests when occurring in large numbers (see, for example, Vayssiere & Mimeur,
1925). Even species which are principally carnivorous can cause extensive mechanical damage to
crops by their burrowing activities (Matheny, 1981). Mole-crickets have been reported as
damaging tobacco, rice, sugar cane, potatoes and other crops, as well as lawns, seed beds and
ornamental plants. Since most of the common African, Asian and Australian species have
Bull. Br. Mus. not. Hist. (Ent.) 46 (2): 175-203 Issued 26 May 1983
176 B. C. TOWNSEND
previously been lumped together under the name 'Gryllotalpa africancC , it is usually this species
which is blamed for the damage. However, this study has shown that true africana does not occur
outside Africa, and even in Africa it is likely that other species also cause damage. Since
different species seem to require different soil conditions, particularly with respect to moisture
content (Bennet-Clark, 1970), it is likely that crops requiring different soil conditions will be
affected by different species. For example, a species occurring in very wet conditions might be
found damaging rice crops, but is unlikely to affect potatoes. It is hoped that the present study
will facilitate investigation of the relative economic importance of the various species.
Material
In addition to material in the British Museum (Natural History), I have examined specimens
from a number of other depositories, through the kindness of the specialists mentioned. The
most important and numerous were from the Musee Royal de 1'Afrique Centrale, Tervuren.
The depositories from which I have seen material are listed below, together with the abbrevia-
tions I have used for them.
ANS Academy of Natural Sciences of Philadelphia, U.S.A.
BMNH British Museum (Natural History), London, England
IAR Institute of Agricultural Research, Samaru, Nigeria
IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium
MNH Museum d'Histoire Naturelle, Geneva, Switzerland
MLU Martin-Luther-Universitat, Halle, East Germany
MNHN Museum National d'Histoire Naturelle, Paris, France
MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, East Germany
MRAC Musee Royal de 1'Afrique Centrale, Tervuren, Belgium
MZSUS Museo Zoologico della Specola, Universita degli Studi, Florence, Italy
NM Naturhistorisches Museum, Vienna, Austria
NMK National Museum of Kenya, Nairobi, Kenya
SAM South African Museum, Cape Town, South Africa
TM Transvaal Museum, Pretoria, South Africa
UZM Zoologisk Museum, Copenhagen, Denmark
ZL Zoologisk Laboratorium, Aarhus University, Aarhus, Denmark
I have examined the types of all the described species except those of minor and africana,
which must be considered lost. The type-series of minor is in neither the MNHU, nor the ZM,
between which the collection containing it was divided; and that of africana is not in the MNHN
where Palisot de Beauvois' collection is deposited.
I have also examined seven specimens labelled as syntypes of G. orientalis Burmeister: two
males and two females from the MLU, and one male and two females from the MNHU. Of
these , only the three from the MNHU have data agreeing with the original description . I am here
designating as LECTOTYPE theinale, which was originally from The Tranquebar Museum and
was collected in Manila, Philippines.
Taxonomic characters
The main characters used here for distinguishing between species are the male stridulatory file
and the venation of the male fore wing. The male genitalia, which are usually a valuable
character in the Gryllidae, are less useful in the African Gryllotalpa, except for the characteristic
genitalia of africana. The females are largely indeterminable, and no key to them is given,
although those of some species can be recognised with practice.
The shape and length of the dactyls of the fore tibiae, often used in the past, do not provide
reliable taxonomic characters. Although they show some variation between species, the dactyls
evidently wear down considerably with use.
AFROTROPICAL MOLE-CRICKETS 177
The stridulatory file of the female shows great intra-specific variation, and does not appear to
be useful in distinguishing between species. Although female mole-crickets are known to
stridulate, the sounds they produce are not pure frequencies, and are probably not used for mate
recognition.
Most African species of Gryllotalpa apparently occur only in the macropterous form, in which
the hind wings extend well beyond the tip of the abdomen in dried specimens. However, all
specimens of microptera and some of debilis are micropterous, their hind wings being shorter
than the abdomen, and often shorter than the fore wings.
Methods
The stridulatory file was examined directly using a binocular microscope. It was exposed by
raising the uppermost fore wing, usually the right one, after relaxing it with a few drops of 10%
ammonia solution to which a little detergent had been added. Drawings of the file were
prepared, using a microprojector, from replicas made using the method described by Ragge
(1969: 172) for Tettigoniidae. The terminology used for the wing venation is that of Ragge
(1955).
All drawings other than those of the stridulatory files were made using a Wild M5 microscope
and camera lucida. Brief diagnoses of previously described species are given, and all new species
are described in full.
Genitalia preparations were made in the following way. The tip of the abdomen was relaxed
using a drop or two of distilled water, together with steam from a water bath. A longitudinal
mid-ventral incision was made along the last three or four abdominal sternites. The viscera were
removed and cleared in cold 20% KOH, and rinsed several times in distilled water. The genitalia
were separated out, and eventually preserved in a tube of glycerine pinned underneath the
specimen.
For identification purposes, the characteristic long ventral processes of africana (Figs 3, 4)
may be exposed in situ, after the tip of the abdomen has been relaxed, by pulling back the
subgenital plate and the covering membrane. The term 'ventral process' is used in the absence of
established terminology for gryllotalpid genitalia or any clear homology with parts of gryllid
genitalia.
The term 'stridulatory area' is used for the pair of large cells of the male fore wing, the anterior
of which is the harp.
The dimensions of the stridulatory area and stridulatory file were measured using a Vickers
Steros II microscope with eyepiece graticule. The length of the stridulatory area was taken to be
equal to the length of the harp, and the width of the area was measured at its maximum.
All other measurements were made using vernier callipers. The body length was measured
from the front of the head to the tip of the abdomen. This measurement is influenced both by the
attitude of the head and by the degree of shrinkage of the abdomen in drying, and is therefore
less reliable than the other measurements given. In most cases, 50 males and 50 females of each
species were measured, where these were available. All measurements are given in millimetres.
The oscillograms shown in Figs 52-63 were made using a Mingograf 34T. The following
acoustic terms are used in song descriptions. A syllable is the sound produced by a single wing
stroke, and an echeme is a discrete group of syllables. The syllable repetition rate is the number
of syllables per unit time, and in complex songs it is measured within a single echeme. The
echeme repetition rate is the number of echemes per unit time. These definitions are those of
Broughton (1964; 1976), and are illustrated in Fig. 52. The carrier frequency is the frequency
within each syllable, and is probably equal to the tooth impact rate (Sismondo, 1979). This is the
frequency of the musical note heard.
Information on distributions is based entirely on specimens studied. Previously published
records are considered unreliable. The term 'Afrotropical Region' is used here but excludes the
Malagasy Region, and many of the offshore islands are not specifically treated due to lack of
material.
178 B. C. TOWNSEND
My approach is entirely phenetic, with no attempt to trace any possible phylogenetic
relationships.
GRYLLOTALPIDAE Leach
Gryllotalpida Leach, 1815: 119. Type-genus: Gryllotalpa Latre'Me.
Scariphasteae Fieber, 1851: 17. [Not based on the name of a contained genus and therefore unavailable
under Article ll(e) of the International Code of Zoological Nomenclature.]
Gryllotalpina; Fieber, 1852: 6.
Gryllotalpiens; Saussure, 1874: 333.
Gryllotalpites; Saussure, 1874: 334.
Gryllotalpidae; Lopez-Seoane, 1878: 375.
Gryllotalpinae; Saussure, 1894: 199.
Gryllotalpini; Redtenbacher, 1900: 140.
Curtillinae 'A'; Kirby, 1906: 1. Type-genus: Curtilla Oken.
Curtillidae; Bruner, 1915: 259.
Gryllotalpoidea; Karny, 1907: 32.
Cf $. Head with two ocelli and two compound eyes. Fore legs highly modified for digging, tibiae bearing
two to four dactyls. Male fore wings lacking mirror. Ovipositor absent.
DISCUSSION. The Gryllotalpidae consist of five easily recognisable genera. Two of these,
Neocurtilla and Gryllotalpella, are restricted to the New World. One mainly New World genus,
Scapteriscus , has two representatives in the Oriental region. The remarkable genus Triamescap-
tor contains a single, wholly apterous species found only in New Zealand. All the remaining
species belong to the largest, entirely Old World genus Gryllotalpa.
Although the Gryllotalpidae have not usually been subdivided, some authors (Zeuner, 1939;
Ragge, 1955; Vickery, 1977) have recognised two subfamilies, placing Scapteriscus in its own
subfamily, Scapteriscinae, and leaving the remaining four genera in the Gryllotalpinae. This
division is based on a difference in the origin of the basal spur of the fore leg, which arises from
the trochanter in Scapteriscus, and from the femur in the other genera; I consider this division to
be justified, and in the key to genera the two subfamilies are separated accordingly.
No major revisionary work on the African species has previously been undertaken. Scudder
(1869) attempted a world revision of the group, but was apparently in possession of only three
African specimens. Chopard (1968) recognised seven species from Africa. In the present
revision, the number of known species is increased to twelve, of which six are new, and one new
specific synonym is established. All species are placed in Gryllotalpa.
Kirby (1906) and Chopard (1955; 1968) placed the Old World Gryllotalpa devia Saussure and
Curtilla madecassa Chopard in the otherwise New World genus Neocurtilla, because of the lack
of spines on their hind tibiae . However, the armature of the hind tibiae is highly variable , and is
generally an extremely unreliable character at specific, let alone generic, level. The main
difference between Gryllotalpa and Neocurtilla is the orientation of the veins of the lateral field
of the fore wing (Figs 1,2), and both species have the Gryllotalpa condition. G. madecassa
comb. n. is endemic to Madagascar, and as such is not included in this study. In size and wing
venation it is more like the European G. gryllotalpa (L.) than any of the African species. G.
devia is dealt with fully in the text.
Figs 1, 2 Lateral field of right male fore wing of (1) Gryllotalpa africana, (2) Neocurtilla hexadactyla.
AFROTROPICAL MOLE-CRICKETS
179
Key to the subfamilies and genera
1 Basal spur of fore leg arising from femur; fore tibia with 3 or 4 dactyls (Gryllotalpinae) 2
Basal spur of fore leg arising from trochanter; fore tibia with 2 dactyls. New World & India
(Scapteriscinae) SCAPTERISCUS Scudder
2 Fore tibia with 4 dactyls; fore and hind wings present in both sexes 3
Fore tibia with 3 dactyls; both sexes apterous. New Zealand TRIAMESCAPTOR Tindale
3 Fore tibia with covered tympanum, opening in the form of a slit; stout-bodied insects 4
Fore tibia with exposed tympanum; slender, delicate insects. South America
GRYLLOTALPELLA Rehn
4 Veins of lateral field of fore wing as in Fig. 1 , all pointing towards wing-tip. Old World
GRYLLOTALPA Latreille
Veins of lateral field of fore wing as in Fig. 2, the more distal ones pointing towards the
wing-base. New World NEOCURTILLA Kirby
Figs 3-6 Male genitalia of (3) Gryllotalpa africana, ventral view, (4) G. africana, lateral view, (5) G.
robusta, ventral view and (6) G. robusta, lateral view. v.p. = ventral process.
GRYLLOTALPA Latreille
Gryllotalpa Latreille, 1802: 275. Type-species: Gryllus Acheta gryllotalpa Linnaeus, by monotypy.
Curtilla Oken, 1815: 445. Type-species: Gryllus Acheta gryllotalpa Linnaeus, by monotypy.
Austrotalpa Mjoberg, 1913: 30. Type-species: Austrotalpa pluvialis Mjoberg [= Gryllotalpa nitidula
Serville], by monotypy. [Synonymised by Tindale, 1928: 4.]
O" $. Fore tibiae with four dactyls. Tympana covered, opening in the form of a slit. Basal spur of fore leg
arising from femur. Fore and hind wings present. Veins of lateral field of fore wings all pointing towards
wing-tips.
DISCUSSION. The African species of Gryllotalpa fall into two quite distinct groups. In male fore
wings of the africana-group , comprising africana, bulla, debilis, devia, robusta and rufescens, the
stridulatory teeth are much more widely spaced at the centre of the file than at its extremities,
and the radius is divided distally into two branches. In male fore wings of the parva-group,
comprising brevity ra, elegans, microptera, parva, pluridens andspissidens, the stridulatory teeth
are more or less evenly spaced, and the radius is undivided. These characters are constant in all
180 B. C. TOWNSEND
species except rufescens, and possibly devia, in which the form of the radius is somewhat
variable ; these two species are placed in the first group on the basis of their stridulatory files. The
form of the radius of the females is similar to that of the males, but is rather inconsistent, and is
not a reliable character for identification purposes.
The species of africana-group are separated by a variety of male characters, as indicated in the
key; those of parva-group chiefly by the shape of the stridulatory area. A large proportion of the
females of rufescens, spissidens and elegans may be identified with practice using characters
mentioned under those species, but females of the other species cannot be reliably identified.
The females of devia and bulla are unknown.
DISTRIBUTION. Throughout the tropical and warmer temperate regions of the Old World.
Synonymic list of the Afrotropical species
a/r/cana-group
africana Palisot de Beauvois
colini Rochebrune
confusa Chopard syn. n.
fossor Scudder
bulla sp. n.
dchilis Gerstaecker sp. rev.
minor Brunn
devia Saussure comb. rev.
robust u sp. n.
rufescens Chopard
parva-group
brevilyra sp. n.
elegans Chopard
microptera Chopard
parva sp. n.
p/ii ride/is sp. n.
spissidens sp. n.
Non- Afrotropical species of Gryllotalpa also covered
madecassa Chopard comb. n. (p. 178)
min tun Burmeister (p. 185)
or/enfa7/s Burmeister (pp. 176, 183)
Key to the Afrotropical species
Males
1 Stridulatory teeth much more widely spaced at centre of file than at ends (Figs 9-15). Radius of
fore wing of macropterous specimens usually divided distally into RI and R$ (Figs 22-27)
(africana-group) 2
Stridulatory teeth fairly evenly spaced (Figs 16-21). Radius of fore wing of macropterous
specimens never divided (Figs 29-33) (parva-group) 7
2 Stridulatory area very oblong (Figs 22, 23, 25-28). Mesonotum covered by pronotum and base
of fore wings, scutum never enlarged (Fig. 7) 3
Stridulatory area almost square (Fig. 24). Mesonotum usually wholly or partly exposed,
scutum usually enlarged (Fig. 8) • G. bulla (p. 183)
3 Pronotum and legs plain brown , from light sandy-coloured to almost black , never rufous 4
Pronotum and legs very conspicuously rufous brown 6
4 Genitalia about 3-0 mm long, with long ventral processes (Figs .3, 4) G. africana (p. 182)
Genitalia about 1-5 mm long, with short ventral processes (Figs 5,6) 5
5 Larger, stouter species, body length 21 -3-43 -6 mm. Length of fore wings 10-0-13-9 mm; width
of stridulatory area 2-1-3-2 mm G. robusta (p. 186)
Smaller, less stout species, body length 15-9-25-2 mm. Length of fore wings 4-4-11-0 mm;
width of stridulatory area 1-0-2-2 mm G. debilis (p. 184)
AFROTROPICAL MOLE-CRICKETS
181
6 Hind tibiae with 3-5 dorsal spines. Equatorial Africa G. rufescens (p. 189)
Hind tibiae without dorsal spines. Southern Africa G. devia (p. 185)
7 Fore wings not reduced, more than 7 mm long, venation as in Figs 29-33. Hind wings long,
extending well beyond tip of abdomen 8
Fore wings much reduced, less than 7 mm long, venation as in Fig. 34. Hind wings vestigial,
shorter than or a little longer than fore wings G. microptera (p. 193)
8 Stridulatory area not reduced, shaped as in Figs 29-31 or Fig. 33, 3-2-4-4 mm long; or if shorter,
then density of teeth less than 35 per mm 9
Stridulatory area reduced, shaped as in Fig. 32, 2-3-3-5 mm long; density of Stridulatory teeth
more than 35 per mm G. brevilyra (p. 189)
9 Main veins and cross veins of apical field of fore wing more or less equally prominent.
Stridulatory area as in Figs 29, 30 or 33 10
Main veins of apical field of fore wing unusually prominent, cross-veins indistinct. Stridulatory
area as in Fig. 30 or Fig. 31 12
10 Stridulatory file with less than 71 teeth, density of teeth 26-4-33-3 per mm. Stridulatory area as
in Fig. 33 G. parva (p. 195)
Stridulatory file with more than 71 teeth, density of teeth 30-4-^5-2 per mm. Stridulatory area
as in Fig. 29 or Fig. 30 11
11 Stridulatory area narrower, 1-8-2-2 mm wide, shaped as in Fig. 30. West Africa & Cameroon
G. spissidens (part) (p. 200)
Stridulatory area broader, 2-1-3-0 mm wide, shaped as in Fig. 29 G. pluridens (p. 197)
12 Density of Stridulatory teeth less than 32 per mm. Stridulatory area as in Fig. 31 . Zaire Basin
G.elegans(p. 191)
Density of Stridulatory teeth more than 32 per mm. Stridulatory area as in Fig. 30. West Africa
& Cameroon G. spissidens (part) (p. 200)
Females
Because the most useful characters in African Gryllotalpa are male sexual characters, identifica-
tion of isolated females is invariably difficult and usually impossible. Even the form of the radius
of the fore wing, although conforming to the same general pattern as that of the male, is not
consistent enough in the female to provide a reliable character. Females of some species,
however, may be recognised by a variety of non-sexual characters.
The female of rufescens can be recognised by its very conspicuous rufous-brown coloration
(see couplet 3 of key to males) . In addition, the fore wings are unusually long, often reaching the
Figs 7, 8 Meso- and metanotum of (7) Gryllotalpa africana, (8) G. bulla.
182 B. C. TOWNSEND
tip of the abdomen, with very straight and parallel veins (Fig. 37). The unknown female oidevia
is probably similar, but lacking dorsal spines on the hind tibiae.
Females of the allopatric (Fig. 50) elegans and spissidens have characteristically prominent
fore wing veins on the dorsal field, with cross-veins indistinct or absent, although this character is
often not well marked in spissidens.
Females of microptera and micropterous females of debilis have characteristically short fore
and hind wings. Macropterous females of debilis are similar to parva and brevilyra. africana,
robusta andpluridens are generally larger than these three, but are indistinguishable from each
other, pluridens has a rather more restricted distribution than africana and robusta.
The female of bulla is so far unknown. If it has the same remarkable form of the metascutum
as the male (Fig. 8), it should not be hard to recognise.
Descriptions of the Afrotropical species
The africa/ia-group
Gryllotalpa africana Palisot de Beauvois
(Figs 1, 3, 4, 7, 15, 25, 38, 48, 55, 61)
Gryllotalpa africana Palisot de Beauvois, 1805: 229. Syntypes, NAMIBIA (lost) (see p. 176). NEOTYPE cf ,
SOUTH AFRICA (ANS), here designated [examined].
Gryllotalpa fossor Scudder, 1869: 21. LECTOTYPE cf, SOUTH AFRICA (ANS), here designated [ex-
amined]. [Synonymised by Chopard, 1968: 450.]
Gryllotalpa colini Rochebrune, 1884: 30. LECTOTYPE $, SENEGAL (MNHN), here designated
[ examined]. [Synonymised by Chopard, 1968: 450.]
[Gryllotalpa formosana Shiraki; Chopard, 1934: 14. Misidentification.]
Gryllotalpa confusa Chopard, 1967: 776. LECTOTYPE cf , ZAIRE (IRSNB), here designated [ examined].
Syn. n.
DIAGNOSIS, cf . Venation of right fore-wing as in Fig. 25, RI and Rs separated distally. Stridulatory file of
right fore wing as in Fig. 15 with 25-52 teeth (mean of 50 examined: 34-6) more widely spaced in centre of
file than at ends, 11-4-20-8 per mm (mean of 50 examined: 15.6). Genitalia very large, with long ventral
processes (Figs 3, 4). Song as in Figs 55, 61, a continuous thrill, mean syllable repetition rate 49-1-57-8/s,
mean carrier frequency 2-1-2-4 kHz (based on 4 recordings).
9 • Right fore wing as in Fig. 38, RI and Rs separated distally.
MEASUREMENTS
Males Females
Body length (50): 22-0-35-0, mean 28-4 (50): 21-9-33-0, mean 28-3
Median length of pronotum (50): 7-5- 9-9, mean 8-6 (50): 7-6- 9-7, mean 8-6
Length of hind femur (50): 7-2-10-4, mean 8-6 (50): 7-0- 9-9, mean 8-5
Length of fore wing (50): 9-6-13-9, mean 12-0 (50): 9-2-14-8, mean 11-9
Length of Stridulatory area (50): 3-9- 5-8, mean 4-9
Width of Stridulatory area (50): 2-1- 2-7, mean 2-4
Length of Stridulatory file (50): 1-6- 2-5, mean 2-2
DISCUSSION The identity of africana cannot be established from Beauvois' original description,
which applies equally well to any African species. In the past, its identity has been far from
settled, although two distinct genitalic forms have long been recognised among African
specimens similar in external morphology; one is small and has short ventral processes typical of
the genus (Figs 5,6), the other is much larger and has long ventral processes (Figs 3,4). Chopard
(1939) referred to the first, typical form as fossor Scudder, and the second, atypical one as
africana. Later (1967), following Saussure & Zehntner (1894), he called the typical form
africana, and gave the atypical form a new name, confusa. In 1968 he synonymised/ossor with
africana.
However, two important facts have emerged in the course of this study. Firstly, the single
species having the atypical genitalic form is by far the commonest species occurring in Africa,
comprising about one-third of all the specimens examined. Secondly, the type-locality for
AFROTROPICAL MOLE-CRICKETS 183
africana is 'Royaume Oware', apparently referring to the region of the Oware River, a seasonal
river running into the Etosha Pan of northern Namibia; the only specimens I have seen from this
area, from the collection of the ANS, are from the Etosha Pan itself, and have the atypical
genitalicform.
For these reasons I am considering the species having the atypical form of genitalia (Figs 3, 4)
to be africana. This preserves the traditional position of africana as Africa's commonest
Gryllotalpa, ensures the greatest possible stability in the nomenclature, and is most likely to be
true to Palisot de Beauvois' syntypes.
Although the specimens from the Etosha Pan are nearest to the type-locality, they are
unusually small for the species (body length 22-0-27-7 mm), perhaps as a result of the hostile
environment. In addition, recent research has shown that the song is often the most important
single character in the Grylloidea, so that where possible type-specimens should be selected
from populations from which song recordings have been made. For these reasons, I have
selected as neotype a male from such a population at Howick, South Africa, in preference to the
specimens from Etosha Pan.
Contrary to the belief of most previous authors, the single male syntype of fossor, here
designated lectotype, possesses the atypical genitalic form, fossor is therefore a synonym of
africana, as is confusa.
The difference in genitalic structure between G. africana and the other species may be related
to a difference in copulatory behaviour (Alexander, 1962). A series of specimens in the BMNH
from Nurtiti, Sudan is labelled as 'damaging potatoes'.
G. colini is included as a synonym of africana; the female lectotype cannot be definitely
identified, but is most likely to belong to this species. The female paralectotype of colini, and
that of fossor, are indeterminable.
MATERIAL EXAMINED
Gryllotalpa africana Beauvois, neotype cf , South Africa: Natal, Mkuze Game Reserve, Nsumu Pan,
19.xi.1980 (One} (ANS). Gryllotalpa fossor Scudder, lectotype cf, South Africa: Cape of Good Hope
(ANS,). Gryllotalpa colini Rochebrune, lectotype $, Senegal: Kita, 1904 (Mabille) (MNHN). Gryllotalpa
confusa Chopard, lectotype cf , Zaire: Rutshuru, 16-30.X.1934 (de Wine) (IRSNB).
Zaire: 1 $, Rutshuru, 17-24.vi.1934 (de Witte); 1 cf, Bitashimwa, Sesero, 17.viii.1934 (de Witte); 1 $
Kahojo, 16.ii.1934 (de Witte); 1 9, S. Bishoke, 2400 m, 8-19. ii. 1935 (de Witte); 3 cf , Lac Mugunga, Nzulu,
1500 m, 25. i. 1934 (de Witte) 1 $, Kibati, 1700 m, 17.1.1934 (de Witte); In Camp Ruindi, 1000 m,
20-28.xi.1934 (de Witte); 3 cf, 3 $, Lac Mugunga, Nzulu, 1500 m, 25.1.1934 (de Witte) (MRAC). (All
paralectotypes of Gryllotalpa confusa Chopard.) In IRSNB unless otherwise stated.
In addition, about 600 adults from localities too numerous to list, from the following countries:
South Africa, Namibia, Zimbabwe, Mozambique, Zambia, Angola, Tanzania, Kenya, Uganda, Rwanda,
Zaire, Somalia, Ethiopia, Sudan, Cameroon, Nigeria, Benin, Ghana, Liberia, Senegal, Socotra.
DISTRIBUTION (Fig. 48). Throughout the African continent, north to Egypt, Libya and Morocco,
and also in the Canary Islands. Although previously thought to occur throughout the Old World
tropics and sub-tropics, africana does not occur in Australia (Dr D. Otte, pers. comm.), and in
Asia and Indonesia it is apparently replaced by G. orientalis, previously thought to be a synonym
of africana. Its presence in southern Spain, Saudi Arabia and the Malagasy Region is likely, but
unconfirmed.
Gryllotalpa bulla sp. n.
(Figs 8, 11,24,49,53,59)
Cf. Fairly uniform brown in colour, veins of fore wings darker, slightly rufous. Pronotum not unusually
large compared with head. Mesoscutum more or less exposed between pronotum and base of fore wings,
often greatly enlarged (Fig. 8). Fore wings broad; venation of right fore wing as in Fig. 24; stridulatory area
very broad, almost square; radius divided distally into R{ and Rs. Stridulatory file of right fore wing as in
Fig. 11, with 35-49 teeth (mean of 9 examined: 41-3) more widely spaced in centre of file than at ends,
12-9-15-2 per mm (mean of 9 examined: 14-1). Hind wings long, extending well beyond tip of abdomen.
Hind tibiae armed above with 1-4 spines on internal margin, or unarmed. Genitalia similar to robusta (Figs
184 B. C. TOWNSEND
5, 6). Song as in Figs 53, 59, a continuous trill; mean syllable repetition rate 128-5/s, mean carrier frequency
4-8 kHz (based on 1 recording).
$ unknown.
MEASUREMENTS
Males
Body length (8): 27-7-32-8, mean 27-3
Median length of pronotum (9): 7-4- 8-8, mean 8-3
Length of hind femur (9): 6-6- 7-8, mean 7-3
Length of fore wing (9): 11-5-13-3, mean 12-3
Length of stridulatory area (9): 3-8- 4-5, mean 4-2
Width of stridulatory area (9): 2-9- 3-3, mean 3-1
Length of stridulatory file (9): 2-4- 3-5, mean 2-9
DISCUSSION. This species is remarkable in the form of the mesonotum (Fig. 8); the scutum is
sometimes greatly enlarged, and this is the only species in which it is exposed. In all other species
the pronotum and base of the fore wings cover the scutum.
The single male from Kenya excluded from the type-series differs in having a longer
stridulatory area (6-3 mm) and fewer (23), less densely packed stridulatory teeth (6-6 per mm).
This specimen may represent an extreme variant, or a separate species. A recording of the song
of this form might confirm its status.
MATERIAL EXAMINED
Holotype cf , Tanzania: Serengeti N.P., Seronera, 14.x. 1980 (One] (ANS).
Paratypes. Zaire: 3 cf , Katanga, Lubumbashi ('Elisabethville'), 1911, xi. 1911, 1930 (Buttgenbach, Miss.
Agric., Lamoral) (MRAC; BMNH); 1 cf, Kapiri, ix.1912 (Miss. Agric.) (MRAC); 5 cf, Katanga,
Kasenia, 15.ix.-15.x.l930 (de Witte) (MRAC; BMNH).
Material excluded from the type-series. Kenya: 1 cf , Masai Reserve, 7.H.1935 (Benson) (UZM).
DISTRIBUTION (Fig. 49). Central and East Africa; holotype found in very wet soil.
Gryllotalpa debilis Gerstaecker sp. rev.
(Figs 13, 14,27,28,40,41,48)
Gryllotalpa debilis Gerstaecker, 1869: 211. Holotype cf , TANZANIA (MNHU) [examined]. [Synonymised
with G. minuta by Chopard, 1968: 451.]
Gryllotalpa minor Brunn, 1901: 276. Syntypes, TANZANIA (lost) (see p. 176). NEOTYPE cf , TANZANIA
(MNHU), here designated [examined; same specimen as holotype of G. debilis Gerstaecker, 1869: 211].
[Synonymised with G. minuta by Chopard, 1968: 451.]
DIAGNOSIS, cf . Venation of right fore wing as in Figs 27, 28, RI and /?s separated distally. Stridulatory file of
right fore wing as in Figs 13, 14, with 21^8 teeth (mean of 50 examined: 31-5) more widely spaced in centre
of file than at ends, 11-6-30-0 per mm (mean of 50 examined: 19-6). Hind wings variable, sometimes
extending well beyond tip of abdomen, sometimes slightly shorter than fore wings, more often intermedi-
ate. Hind tibiae armed above with 2-4 spines on internal margin. Genitalia similar to robusta (Figs 5,6).
9- Right fore wing as in Figs 40, 41, Rl and Rs separated distally.
MEASUREMENTS
Males Females
Body length (50): 15-9-25-2, mean 20-8 (50): 19-2-26-5, mean 22-9
Median length of pronotum (50): 5-2- 8-2, mean 6-7 (50): 6-1- 7-9, mean 7-0
Length of hind femur (50): 5-1-7-6, mean 6-4 (50): 5-9- 7-9, mean 6-6
Length of fore wing (50): 4-4-11-0, mean 7-9 (50): 3-9-10-2, mean 7-9
Length of stridulatory area (50): 2-3- 4-6, mean 3-6
Width of stridulatory area (50): 1-0- 2-2, mean 1-7
Length of stridulatory file (50): 1-2- 2-5, mean 1-6
DISCUSSION. G. debilis is very similar to robusta (see p. 186), but is smaller and less stout. Both
species show unusually wide variations in minor characters such as colour, colour pattern, size
AFROTROPICAL MOLE-CRICKETS 185
and shape, and in the form of the stridulatory area and stridulatory file, suggesting that they may
in fact be complexes of several very similar species. However, I have not been able to subdivide
them satisfactorily on the basis of morphological characters.
This species has been confused with G. minuta, from which it differs chiefly in the smaller
number of stridulatory teeth (number on holotype of minuta: 63). minuta is common in the
Oriental region, but does not occur in Africa.
The type-series of minor was from Zanzibar, the type-locality of debilis, and since these two
names have been associated for several years, I have decided to treat them as synonyms. In order
to establish their synonymy firmly, I have designated the holotype of debilis as neotype of minor,
the name debilis taking priority.
G. debilis is also morphologically indistinguishable from G. orientalis. The two may be
synonymous, and if so this would apparently be the only species of Gryllotalpa common to the
Afrotropical and Oriental regions. The two taxa are treated as specifically distinct until a
recording of the song can be compared with that of orientalis (Figs 57, 63).
G. debilis is the only Afrotropical species of which both macropterous and micropterous
forms are known. The micropterous form is superficially similar to microptera, but differs in the
form of the stridulatory file (Figs 14, 19) , and in having the radius divided distally into R^ and /?s.
MATERIAL EXAMINED
Holotype cf , Tanzania: Zanzibar (MNHU).
Zambia: 3 cf, 5 $, In, Lake Bangweulu, Mbawala Is., x.-xi.!946 (Steele) (BMNH). Namibia: 1 cf,
Naukluft, 1300-1500 m, 7-10. xii. 1933 (Jordan] (BMNH). Uganda: 1 cf, Mwiri, Turtle Pool, 20.xi.1954
(Corbet} (BMNH). Zaire: 1 cf, Bas-Kasai, ix.1920 (Vanderijst); 1 cf, Equateur, Boende, 13. iv. 1926
(Hulstaert); 2 cf, Sankuru, Komi, iv.-v.1930 (Ghesquiere); 1 cf, Katanga, Katompe, 1-15. vi. 1930
(Gerard); 1 cf, Eala, 22.x. 1931 (Bredo}\ 1 $, Ruwenzori, Mutwanga, ii.-iii.1937 (Hackars); 3 cf, 1 ?,
Mutsora, 1939 (Hackars} (IRSNB); 1 cf , 3 $, Kasika, R. Ngombe, 8-10.vi.1949 (Laurent}; 64 cf , 111 $,
Upemba NP, Ganza, 8.vi.l949, 5.vii.l949 (de Witte} (15 cf, 23 $ in BMNH); 9 cf, 1 $, Garamba NP,
xi.1949, 2.1.1950, 6.J.1950, 18.viii.1950, 6.x. 1951, 28.xi.1951, 21.viii.1952 (Demoulin, De Saeger} (1 cf in
BMNH); 1 cf , Kwango, Popakabaka, i. l952(Pierquin);9 cf 12 $ , 8 n, Albert NP, Ruwenzori Massif, near
Kalonge, Kisesa, 23.V.1953 (Vanschuytbroeck & Kekenbosch} (3 cf , 4 $, 2 nn in BMNH); 12 cf , 4 ?, 8 n,
Albert NP, various localities, 2.iii.-23.vii.l957 (Vanschuytbroeck) (3 cf in BMNH); 3 Cf, 11 $, Stanley
Pool, 3-10.X.1957, 7.X.1957 (Bouillon) (1 cf, 2 $ in BMNH); 1 cf, Mayumbe, Singa to Mbomba, T.
Kipanzu, v.-vi.!958 (Laurent); 1 cf, 3 $, Mayumbe, Vemba to Minionzi, T. Tshela, vi.-vii.1958
(Laurent}. Ethiopia: 1 cf , Adda shore of L. Hora Harsadi, 3. xii. 1936 (Omer-Cooper) (BMNH). Nigeria: 1
Cf , Sokoto, 1921 (Moiser} (BMNH); 1 cf , 1 $, Ibadan, i.-vi.!954 (Clausen) (UZM); 1 cf , 8 $, Western
Province, 3-5 miles N. of Oyo, near Idode, 16. xii. 1960 (Jago} (BMNH); 1 cf, Western Pronce, Ibadan,
University College, 17. xii. 1960 (Jago) (BMNH); 1 cf , Zaria, Samaru, 1979 (Deeming) (IAR). Ghana: 1
Cf, Accra, Legon, 9.iii.l969 (Richards) (BMNH). Chad: 1 cf, Bebedjia, xi.1965 (Schmitz). In MRAC
unless otherwise stated.
DISTRIBUTION (Fig. 48). Tropical Africa; also known from Mauritius, Rodriguez, the Seychelles
and Saudi Arabia. The record from Namibia, which is based on a single male in the BMNH, is in
need of confirmation.
Gryllotalpa devia Saussure comb. rev.
(Figs 10, 22, 48)
Gryllotalpa devia Saussure, 1877: 25. Holotype cf , SOUTH AFRICA (MHN) [examined].
Neocurtilla devia (Saussure) Kirby, 1906: 2.
DIAGNOSIS, cf. Pronotum and legs rufous-brown, colour 38 (Tawny) or 40 (Cinnamon-Rufous) in
Naturalist's Colour Guide (Smithie, 1975). Pronotum very large compared with head. Venation of right
fore wing as in Fig. 22, radius divided distally into R{ and Rs in holotype, possibly sometimes undivided as in
rufescens. Stridulatory file of right fore wing as in Fig. 10, with 40 teeth, more widely spaced in centre of file
than at ends, overall density 12 per mm. Hind tibiae without dorsal spines,
unknown.
186 B. C. TOWNSEND
MEASUREMENTS
Male holotype
Body length 35-0
Median length of pronotum 12-0
Length of hind femur 10-0
Length of fore wing 15-0
Length of stridulatory area 5 • 8
Width of stridulatory area 3 • 7
Length of stridulatory file 3 • 3
DISCUSSION. Kirby (1906: 2) placed this species in his New World genus Neocurtilla, presumably
because of the absence of dorsal spines on its hind tibiae. However, this is not a reliable generic
character, and the venation of the lateral field of the fore wing clearly places devia in Gryllotalpa
(see p. 178). The fragile condition of the holotype prevents examination of the genitalia, but
those of the only other recorded specimen have been figured by Chopard (1955: fig. 16), and do
not appear to have any unusual characteristics. The orientation of some of the stridulatory
teeth appears to be reversed in the holotype (Fig. 10), but this may be abnormal.
MATERIAL EXAMINED
Holotype cf , South Africa: Cape of Good Hope (MHN).
DISTRIBUTION (Fig. 48). Southern Africa, known only from the Cape of Good Hope and
Lesotho. Apparently associated with drier regions than is usual for mole-crickets.
Gryllotalpa robusta sp. n.
(Figs 5, 6, 12,26,39,49,54,60)
[Gryllotalpa africana Palisot de Beauvois; Saussure & Zehntner, 1894: 406; Chopard, 1967: 775.
Misidentifications.]
[Gryllotalpa fossorScudder; Chopard, 1939: 6. Misidentification.]
Cf . Fairly uniform in colour, light brown to black, veins of fore wings darker. Pronotum not unusually large
compared with head. Mesoscutum not exposed, concealed by pronotum and base of fore wings, never
enlarged. Venation of right fore wing as in Fig. 26; stridulatory area more or less rectangular; radius
divided distally into R^ and /?s. Stridulatory file of right fore wing as in Fig. 12, with 30-42 teeth (mean of 7
examined: 35-7) more widely spaced in centre of file than at ends, 11-0-16-0 per mm (mean of 7 examined:
12-9). Hind wings long, extending well beyond tip of abdomen. Hind tibiae armed above with 2-4 spines on
internal margin. Genitalia small, with short ventral processes (Figs 5, 6). Song as in Figs 54, 60, a
continuous trill, mean syllable repetition rate 98-5/s, mean carrier frequency 1-6 kHz (based on 1
recording).
9 . Right fore wing as in Fig. 39, radius usually divided distally into RI and fls.
MEASUREMENTS OF HOLOTYPE
Body length
Median length of pronotum
Length of hind femur
Length of fore wing
Length of stridulatory area
Width of stridulatory area
Length of stridulatory file
35-2
13-7
6-0
3-0
2-9
OVERALL MEASUREMENTS
Body length (50): 21-3-35-2, mean 26-3 (11): 23-4-35-1, mean 29-5
Median length of pronotum (50): 7-0-10-0, mean 8-0 (11): 6-6- 8-9, mean 8-0
Length of hind femur (50): 6-7-10-0, mean 7-9 (11): 7-0-10-0, mean 8-6
Length of fore wing (50): 10-0-13-9, mean 11-7 (11): 10-9-13-5, mean 11-7
Length of stridulatory area (50): 4-0- 6-1, mean 5-1
Width of stridulatory area (50): 2-1-3-2, mean 2-6
Length of stridulatory file (50): 1-6- 3-0, mean 2-3
Number of stridulatory teeth (50): 17-0-42-0, mean 29-4
AFROTROPICAL MOLE-CRICKETS 187
Overall density of
stridulatory teeth (50): 9-0-18-0, mean 13-0
DISCUSSION. This is the most nondescript of all the African mole-crickets. It was previously
confused with africana, and also misidentified asfossor, which is now a synonym of africana.
Externally, it is indistinguishable from africana but lacks the characteristic male genitalia of that
species (Figs 3-6) and differs radically from it in song (Figs 54, 55, 60, 61). It is also very similar to
debilis, differing chiefly in its larger size and more robust shape.
Both robusta and debilis show a much greater range of variation in minor characters, and in
the form of the stridulatory area and stridulatory file, than that normally found within a single
species. These characters, such as colour, colour pattern, size and shape, appear to be quite
consistent within single populations, and strongly suggest that both robusta and debilis are in fact
complexes of several very similar species. However, I have been unable to subdivide them
satisfactorily on the basis of the morphology.
Because of the similarities between robusta and debilis in major characters, particularly the
fore wing venation and the gross form of the stridulatory file (Figs 12, 13, 14, 26, 27, 28), and
because of the great variation within both taxa, a number of specimens cannot be definitely
identified as one or the other. The changes in nomenclature adopted in this paper, which leave
the present species without a name, offer an opportunity simply to treat the two taxa as a single
species, under the name debilis. However, I have decided against this for the following reasons.
1. Such a step would involve combining two taxa previously recognised as separate. This
would be misleading, in view of the evidence suggesting that there are more than two species,
not less.
2. debilis is more similar to orientalis than to robusta, but orientalis and robusta have different
songs (Figs 54, 57, 60, 63). This provides further circumstantial evidence for a specific difference
between debilis and robusta, though the song of debilis itself is unknown.
3. The holotype of robusta is clearly different from that of debilis, and its song is known.
Despite the variation within each taxon, and the areas of overlap between them, the majority of
specimens can be assigned to one or the other on the basis of the characters described.
Because of the great variation involved, I have not designated paratypes of robusta. I have
given separate series of measurements for the holotype, and for all the specimens grouped under
robusta.
MATERIAL EXAMINED
Holotype cf , Tanzania: Serengeti N.P., Musabi Plains, c. 30 miles NW. of Serona, 20.X.1980 (Otte)
(ANS).
Material excluded from type-series. South Africa: 1 cf , KrugerN.P.,c. 70 miles N. of Skukuza, Olifant's
Camp, 7.vii.l974 (Pitkin) (BMNH); Tanzania: 1 cf, Ukerewe Is. (Conrad) (NMK); 1 cf, Kabolo,
5.vii.l947 (Poll)- 3 cf , Sumbawanga, xii.1980 (Moyer) (BMNH). Zambia: 1 cf, Lochimvar, 6-26.V.1964
(Van Noteri). Kenya: 10 cf , 1 n, Kinangop, i.1930, xi.1930 (Turner) (7 cf , 1 n in NMK; 3 cf in BMNH);
1 Cf , Kaimosi, iii-iv.1932 (NMK); 2 cf , 3 $ , Lake Baringo, Molo R. mouth, 17.vi.1934 (Rehn) (2 cf , 2 $ in
ANS, 1 £ in BMNH). Uganda: 1 cf , Kagora Plains, vi.1911 (Marshall) (BMNH); 1 cf, Kalinzu Forest,
x.1948 (Jackson) (NMK); 2 cf , Mpanga Forest, Fort Portal, ii.1957 (Carcasson) (NMK). Rwanda: 1 cf,
Rubengeri, 1911 (Lestrade); 1 cf, Kisenyi, i.1954 (Bertrand). Zaire: 1 cf, Katanga; 1 cf, Haut Congo
(IRSNB); 5 cf , Kambove, ix.1906-iii.1907 (Neave) (1 in BMNH); 16 cf, Bunkeya, x.1907 (Neave) (3 in
BMNH); 2 cf , Kambove to Chitura, xi.1907 (Neave); 1 cf, xii.1907 (Neave); 1 cf , Kasenyj, 19.vii.1911
(Stoppers); 1 cf, Kapiri, ix.1912 (Miss. Agric.); 3 Cf, 1 9, Katanga, Mwema, vii.1927 (Bayet) (1 cf in
BMNH); 1 cf , Ituri, Butembo, xii.1928 (Van Kiel); 1 cf , Semliki Plain, 900-1100 m, iv.-x.1937 (Hackars)
(IRSNB); 1 cf, Kunungu, 1941 (N'Kele); 1 cf, Bas Congo, Lemfu, x.-xii.!944 (Beir); 1 cf, Kivu,
Kitwabalazi, 1946 (Herrinck); 1 cf, Kindia, 2.V.1948 (Olsen) (ZL); 5 cf, Katanga, Kundelungu, Affl.
Lualaba II, L. Moero Basin, 1680 m, 17-19.X.1951 (Leleup) (2 in BMNH); 9 cf , Garamba N.P. , 30.xi.1951
(De Saeger) (3 in BMNH); 1 cf , L. Albert, Mahagi Port, 16.ii.1954 (Verbeke) (IRSNB). Somalia: 1 cf ,
Iscia Baidoa, 12-28. vi. 1978 (MZSUS). Malawi: 1 cf, 2 $, Namalindi, 12-14.xii.1969 (BMNH).
Cameroon: 2 cf , 6 ?, M'Bakaou, Hi. 1967 (Chemin) (1 cf , 2 9 in BMNH); 2 cf , 1 $, Koum, 20-22.U976.
(Puylaert). Ethiopia: 6 cf, Zegi Tsana, v.-vi.!902 (Degen) (BMNH). Nigeria: 1 cf, 2 $, Zaria, Samaru,
1979 (Deeming) (1 cf, 1 $ in IAR; 1 $ in BMNH). Togo: 1 cf, 3 $, Piya, 18-22.V.1963 (Schach) (1 9 in
188 B. C. TOWNSEND
BMNH). Ghana: 1 cf, Gold Coast (Woodward) (BMNH). Sierra Leone: 2 cf, 2 $, Rokupr, 1977
(BMNH). In MRAC unless otherwise stated.
DISTRIBUTION (Fig. 49). Africa south of the Sahara, and the Canary Islands. Holotype found in
very wet soil.
9 '°CCCccccccC C C C C C t C C C C C C C
10 C C C C r
c c c c c c c
„ .ce.ccceccccccc ccc c««cce,ec
12 c c c c c c c c c c c cccccccccCCff ^
13 ' C <c c = c c c c c c c eccct**" c Cc',% U "
t|
15 c c c c c r
C C C c C
16
17
18
19 "—««.«„«.„, WW,«H-.— . , 20
"X
21 c'^,.
Figs 9-21 Right male stridulatory file of (9) Gryllotalpa rufescens, (10) G. devia, (11) G. bulla, (12) G.
robusta, (13) G. debilis (macropterous), (14) G. debilis (micropterous), (15) G. africana, (16) G.
pluridens, (17) G. spissidens, (18) G. elegans, (19) G. microptera, (20) G. brevilyra, (21) G. parva.
AFROTROPICAL MOLE-CRICKETS 189
Gryllotalpa rufescens Chopard
(Figs 9, 23, 37, 49, 52, 58)
Gryllotalpa rufescens Chopard, 1948: 110. LECTOTYPE cf, ZAIRE (MRAC), here designated [ex-
amined].
DIAGNOSIS, cf. Pronotum and legs rufous brown, colour 38 (Tawny) or 40 (Cinnamon-Rufous) in
Naturalist's Colour Guide (Smithie, 1975). Pronotum very large compared with head. Venation of right
male fore wing as in Fig. 23, radius sometimes divided distally into RI and Rs, sometimes undivided.
Stridulatory teeth of right male fore wing as in Fig. 9, with 38-68 teeth (mean of 12 examined: 51-1) more
widely spaced in centre of file than at ends, overall density 9-2-18-9 per mm (mean of 12 examined: 13-1).
Hind tibiae armed above with 4-5 spines on internal margin. Song as in Figs 52, 58, a continuous trill
consisting of repeated echemes of three syllables; mean syllable repetition rate 80-0-100-0/s, mean echeme
repetition rate 15-9-17-6/s, mean carrier frequency 2-7-2-8 kHz (based on two recordings). Genitalia
similar to robusta (Figs 5,6).
9- As male except for song and genitalia, and fore wings as in Fig. 37, long, often reaching tip of
abdomen, venation variable, radius sometimes divided distally into R{ and /?s, sometimes undivided,
sometimes joined with Sc.
MEASUREMENTS
Males Females
Body length (10): 25-3-32-9, mean 29-4 (4): 26-1-31-2, mean 29-0
Median length of pronotum (12): 9-8-12-0, mean 10-7 (4): 9-6-11-2, mean 10-2
Length of hind femur (10): 8-4-10-6, mean 9-6 (4): 9-0-10-2, mean 9-6
Length of fore wing (12): 12-7-15-2, mean 14-0 (4): 14-5-16-6, mean 15-6
Length of Stridulatory area (12): 5-0- 6-8, mean 5-6
Width of Stridulatory area (12): 2-9- 4-0, mean 3-2
Length of Stridulatory file (12): 3-2- 5-1, mean 3-9
DISCUSSION. This species is similar to devia, from which it may be distinguished by the presence
of spines on the dorsal surface of the hind tibiae. The song is unusual in having the syllables
grouped in threes (Fig. 52).
MATERIAL EXAMINED
Lectotype cf , Zaire: Kunungu, 1937 (NkeleforSchouteden) (MRAC).
Zaire: 1 cf, Mongbwalu, Kilo, 1930 (Milliau); 1 cf , Rutshuru, xi.1937 (Ghesquiere); 1 9 (paralecto-
type), Mongbwalu, Kilo, vii.1938 (Scheitz); I n, Bambesa, 10.ii.1939 (Vrydagh); 1 cf, Kivu, Matale,
U.\.l949(Marlier);l $, Bunyakiri 1800m, 5-7.vi. 1949 (Laurent) (all in MRAC). Cameroon: 1 cf, Metet,
ii.l922(L/ppe/-0;2cf,Lolodorf,iv.l925,18.xii.l926(Goorf)(allinANS);lcf,D'JaPosten, l-30.vii.1936
(Merfield). Uganda: 1 cf, near Kisoro, Busanza, 13.xii.1970 (Bailey}; 3 cf, 3 $, Kigezi, Kinanira,
22.xi.1973 (Ngirumwe) (all in BMNH).
DISTRIBUTION (Fig. 49). Strictly equatorial, in moist woodland and rainforest of central Africa.
The parva-group
Gryllotalpa brevilyra sp. n.
(Figs 20, 32, 45, 51)
Cf . Fairly uniform in colour, light to dark brown, veins of fore wings a little darker. Pronotum not unusually
large compared with head. Lobes of mesonotum not exposed, concealed by pronotum and base of fore
wings, never enlarged. Right fore wing as in Fig. 32; Stridulatory area small, particularly posterior cell; R\
and fls fused; Stridulatory file of right fore wing as in Fig. 20, with 51-94 teeth (mean of 50 examined: 66-6),
fairly evenly spaced, 36-1-54-2 per mm (mean of 50 examined: 41-4). Hind wings long, extending well
beyond tip of abdomen. Hind tibiae armed above with 3-4 spines on internal margin. Genitalia small, with
short ventral processes.
$ . Right fore wing as in Fig. 45, /?, and /?s fused.
190
B. C. TOWNSEND
22
23
radius
26
Figs 22-36 Right male fore wing of (22) Gryllotalpa devia, (23) G. rufescens, (24) G. bulla, (25) G.
africana, showing position of radius, R\ and /?s, and boundary of stridulatory area, (26) G. robusta, (27)
G. debilis (macropterous), (28) G. debilis (micropterous), (29) G. pluridens, (30) G. spissidens, (31) G.
elegans, (32) G. brevilyra, (33) G. parva, (34) G. microptera, (35) G. orientalis, (36) G. minuta.
AFROTROPICAL MOLE-CRICKETS 191
MEASUREMENTS
Males Females
Body length (50): 17-4-27-2, mean 22-7 (36): 17-6-30-1, mean 23-7
Median length of pronotum (50): 5-7- 8-0, mean 7-1 (37): 5-4- 8-6, mean 6-8
Length of hind femur (50): 5-8- 8-2, mean 7-0 (36): 5-7- 8-2, mean 6-9
Length of fore wing (50): 7-5-11-0, mean 9-2 (37): 8-0-11-7, mean 9-6
Length of stridulatory area (50): 2-3- 3-5, mean 2-9
Width of stridulatory area (50): 1-3- 1-9, mean 1-5
Length of stridulatory file (50): 1-2- 2-5, mean 1-6
DISCUSSION. G. brevity ra may be identified by the characteristic shape of the male stridulatory
area (Fig. 32). The male stridulatory file is somewhat intermediate between those oi elegans and
parva.
MATERIAL EXAMINED
Holotype cf . Nigeria: Jos, 1968 (Bot-Gwong) (MRAC).
Paratypes. Nigeria: 2 cf, 6 $, same data as holotype (1 cf, 1 $ in BMNH); 2 cf , Zaria, Samaru, 1979
(Deeming) (1 in IAR; 1 in BMNH). Zaire: 1 cf, 1 9, Wenga Ifomi (Quineaux) (1 $ in IRSNB, 1 cf in
BMNH); 1 cf , Haut Congo (IRSNB); 1 cf , Kwango (IRSNB); 1 cf , Kabinda (Muller) (IRSNB); 1 cf ,
Kikwit (de Caters) (IRSNB); 1 cf, Camp Lukula, 1911 (Daniel); 1 cf, Mobwasa, ix.1911 (Giorgi); 1 $,
Eala, Hi. 1917 (Mayne); 2 cf, Kikwit, 1920 (Vanderijst) (1 in BMNH); 1 cf, Haut-Uele, Moto, 1920
(Burgeon); 1 cf , 1 $, Kisantu, 1927 (Vanderijst) (1 9 in BMNH); 2 cf , Kisangani ('Stanleyville'), xi.1929,
1949 (Collart, Miller}; 6 cf , Kasai, Tshikapa, 1930 (Fourche) (3 in BMNH); 1 cf , Sankuru, Komi, v.1930
(Ghesquiere); 1 cf , Kinshasa ('Leopoldville'), 31.viii.1930 (de Witte); 1 cf , Katanga, Kakyelo, 1-9. xi. 1930
(de Witte); 1 cf , Kunungu, 1932 (Nkele for Schouteden); 1 cf , Lomami, Kaniama, iii.-iv.1932 (Massart); 2
Cf, 7 9, Eala, v.1921, 4.xi.l930, iv.1932, 17. iv. 1932, 30.viii.1933, i.1935, ix.1935, xi.1936 (Ghesquiere,
Bredo, Corbisier) (2 9 in BMNH); 1 $, Lualua, Kapanga, 1934 (Overlaet); 1 cf , Katanga, Tshipama, 1936
(Drion); 1 cf , Mpese, 21-26.ix.1936 (Cooreman) (IRSNB); 1 cf , Kunungu, 1938 (Nkele for Schouteden); 1
9, Lubunday, Albertville, 25.vii.1938 (Pojer) (IRSNB); 1 cf , N. Rosso Norma, Lake Tumba, 31.vii.1938
(Loreux) (IRSNB); 1 cf, Katanga, Mukabe to Kasavi, 1939 (De Donckere); 1 cf, Lokandu, iii.1939
(Maree); 1 cf , Mongbwalu, v.1939 (Lepersonne); 1 cf , Lubunday, Albertville, 2.vii.l939 (Pojer) (IRSNB,
Brussels); 1 cf, Lisala, ix.-x.1939 (Leontovitch); 3 cf, Mayidi, 1942, 1945 (Van Eyen); 5 cf, Lemfu,
x.-xii.!945, xii.1945 (De Beir); 1 cf , Tshuapua, Flandria, 1946 (Hulstaert); 13 cf, 18 9, 1 n, UpembaN.P.,
various localities, 4-24.xi.1947, 6.ix-16.x.l948, 10.vi.-7.vii.1949 (de Witte) (3 cf , 6 $ in BMNH); 1 cf,
Titule, ix.-x.1949 (Verbeke) (IRSNB); 1 cf, 1 9, Gandajika, 27.xi.1950 (de Francquen)! cf, 1 9,
Maniema, Mobanga, 1952 (Henrard); 4 cf, 12 9, Bokuma, i.-ii.!952, ii.1952, iii.1952, iv.1952, vii.1952,
1953, 1954, 1955 (Lootens) (1 cf in BMNH) 1 cf, 2 $, Kalina, Kinshasa ('Leopoldville'), 1952
(Theunissen); 2 cf, Lake Tanganyika, Albertville, 14.viii.1953 (Verbeke) (IRSNB; BMNH); 1 cf,
Bokalakala, Bolobo, 1954 (Eloy); 1 cf, Albert N.P., Ruwenzori Mts., Kombo, 1550 m, 19.vii.1954
(Vanschuytbroeck & Synave); 1 cf , Albertville, xi.1954 (Bomans); 1 9, Bokuma, 1955 (Lootens); 1 cf,
Tshuapua, Ikela, 1955 (Lootens); 1 cf , Mt. Hoyo, nearKivu, iv.-v.1955 (Hostie) (IRSNB); 1 cf , Sankuru,
Djeka, 1955-1956 (Roiseaux); 1 cf , Katanga, Busumba, viii.-ix.1957 (de Caters); 1 cf, Kasongo, ix.1959
(Benoit). Tanzania: 1 cf , L. Malawi, Mbamba Bay, 12-16.iv.1936 (Zerny) (NM). Zimbabwe: 1 cf , Balla
Balla, 30.xi.1913 (Jones) (BMNH). In MRAC unless otherwise stated.
DISTRIBUTION (Fig. 51). This species occurs mainly in the central African rainforest, spreading
west as far as Nigeria, east to Lake Victoria, and south into Zambia. The record for Zimbabwe is
based on a single male in the BMNH, and is in need of confirmation.
Gryllotalpa elegans Chopard
(Figs 18, 31, 44, 50)
Gryllotalpa elegans Chopard, 1934: 14. LECTOTYPE cf , ZAIRE (MRAC), here designated [examined].
DIAGNOSIS, cf. Venation of right fore wing as in Fig. 31, main veins unusually prominent, cross-veins
indistinct or absent; R\ and /?s fused; stridulatory area somewhat triangular, narrowing posteriorly.
Stridulatory file of right fore wing as in Fig. 18, with 44-74 teeth (mean of 22 examined: 59-2) fairly evenly
spaced, 23-5-31-0 per mm (mean of 22 examined: 27-2).
9- Venation of right fore wing as in Fig. 44, main veins unusually prominent, cross-veins indistinct or
absent; /?, and /?s fused.
192
B. C. TOWNSEND
Figs 37-47 Right female fore wing of (37) Gryllotalpa rufescens, (38) G. africana, (39) G. robusta, (40) G.
debilis (macropterous), (41) G. debilis (micropterous), (42) G. pluridens, (43) G. spissidens, (44) G.
elegans, (45) G. brevilyra, (46) G. parva, (47) G. microptera.
MEASUREMENTS
Body length
Median length of pronotum
Length of hind femur
Length of fore wing
Length of stridulatory area
Width of stridulatory area
Length of stridulatory file
Males
(17): 19-3-28-5, mean 23-1 (11)
(17): 6-5- 8-5, mean 7-3 (11)
(17): 6-1- 7-7, mean 6-9 (11)
(17): 8-0-10-8, mean 9-6 (11)
(17): 2-8- 3-5, mean 3-2
(17): 1-4- 1-9, mean 1-7
(22): 1-6- 2-7, mean 2-2
Females
21-7-27-8, mean 23-9
6-9- 8-3, mean 7-5
6-7- 7-9, mean 7-2
8-0-11-1, mean 9-7
DISCUSSION. This distinctive species is recognisable by its prominent fore wing veins. The shape
of the stridulatory area is rather similar to that of spissidens, but its fewer, less densely arranged
AFROTROPICAL MOLE-CRICKETS 193
stridulatory teeth distinguish it from that species. The distribution otspissidens is quite different
(see below).
MATERIAL EXAMINED
Lectotype cf , Zaire: Bas-Congo, Yumbi, 1.x. 1929 (Bredo)
Zaire: 1 9 (paralectotype) same data as lectotype; 1 $, L. Leopold II (Hollebeke); 2 cf, Kinshasa,
13.x. 1896 (Waelbroeck) (IRSNB); 1 9, Congo R., Mongala, 22.xii.1919 (Tinant); 1 9, Luebo to
Luluabourg, 1921 (Ghesquiere); 2 cf , Bas-Congo, Yumbi, 1.x. 1929 (Bredo, Fini) (1 in BMNH); 1 cf , Eala,
xi,1934 (Ghesquiere); 1 9, Kunungu, 1941 (N'Kele); 1 cf , Kalina, 12. xi. 1942 (Fiasse); 3 cf , 3 $, Kinshasa
('Leopoldville'), 1942, i.1947, ii.1947, 20-30. ix. 1950, 15.xii.1950 (Fiasse, Dartevelle, Jobels) (1 cf in
BMNH); 2 cf , Boma, 19. xi. 1952 (Basilewsky) (1 in BMNH); 1 $ , Bokuma, i.-ii.!952 (Lootens) (BMNH);
1 9, Kasai, Djeka, 1954 (Roiseaux); 1 cf, Ishango, vii.1954 (Semliki) (IRSNB); 30 cf, 26 $, Tshuapua,
Bamanya, x. 1951, 1952, x. 1952, i. 1955, 1960, x. 1961, xii. 1961, 1968 (Hulstaert,Sibbens-Pollet); (4 cf, 5 $,
in BMNH); 1 cf , Tshuapua, Boende, 1960 (Sibbens-Pollet). In MRAC unless otherwise stated.
DISTRIBUTION (Fig. 50). Restricted to the basin of the Congo River and its tributaries. In the
coastal rainforest of West Africa and Cameroon it is replaced by spissidens.
Gryllotalpa microptera Chopard
(Figs 19, 34, 47, 51)
Gryllotalpa microptera Chopard, 1939: 6. LECTOTYPE cf , ZAIRE (MRAC), here designated [examined].
DIAGNOSIS, cf. Fore wings much shorter than abdomen, sometimes not extending beyond second
abdominal segment in larger specimens, occasionally reaching fifth or sixth segment in smaller ones;
venation of right fore wing as in Fig. 34, R\ and /?s fused; stridulatory area somewhat triangular.
Stridulatory file of right fore wing as in Fig. 19, with 40-60 teeth (mean of 20 examined: 52-5), fairly evenly
spaced, 26.7-47.5 per mm (mean of 20 examined: 34-0). Hind wings vestigial, usually shorter than fore
wings, sometimes a little longer.
9 . Fore wings sometimes a little longer than those of males; venation of right fore wing as in Fig. 47, R\
and RS fused.
MEASUREMENTS
Males Females
Body length (27): 15-6-29-0, mean 20-6 (24): 15-7-29-6, mean 21-1
Median length of pronotum (27): 5-2- 8-6, mean 7-0 (24): 5-0- 9-2, mean 7-2
Length of hind femur (27): 5-0- 7-7, mean 6-2 (24): 5-0- 8-1, mean 6-4
Length of fore wing (27): 3-3- 6-8, mean 4-8 (24): 3-6- 6-8, mean 5-3
Length of stridulatory area (24): 1-8- 3-0, mean 2-4
Width of stridulatory area (24): 1-0- 1-7, mean 1-4
Length of stridulatory file (20): 1-0- 2-1, mean 1-6
DISCUSSION. G. microptera is the only known Afrotropical Gryllotalpa which is invariably
micropterous. Superficially it resembles the micropterous form of debilis, but differs from it in
having fewer, more evenly spaced stridulatory teeth (Figs 14, 19), and in the radius being
undivided.
MATERIAL EXAMINED
Lectotype cf , Zaire: Rutshuru, 13. vi. 1934 (de Witte) (MRAC).
Zaire: 2 9, Rutshuru, 26.xi.-16.xii. 1933, 17-25.xii.1933 (de Witte) (IRSNB); 1 9, Lac Mugunga,
25.i.-3.ii.l933 (de Witte) (IRSNB); 3 cf, 2 9, Lac Mugunga, 25. i. 1934 (de Witte) (MRAC); 1 9, Bulengo,
29.1.1934 (de Witte) (MRAC) (all paralectotypes); 1 9, Katanga, Nyonga, v.1925 (de Witte); 1 cf,
Katanga, L. Kabamba, v.1927 (Bayet); 1 cf, Katanga, Mwema, vii.1927 (Bayet); 1 n, L. Mugunga,
25.i.-3.ii.l933 (de Witte); 2 n, Rutshuru, 26.xi.-16.xii. 1933, 26.xii.1933 (de Witte) (IRSNB); 2 n, L.
Mugunga, 25. i. 1934 (de Witte) (IRSNB); 1 cf, 1 9, Niangara-Dungu, Ekibondo, R. Uele, 28.ix.1934
(Rehn) (ANS); 1 cf, Lisala, 6.x. 1937 (Leontovitch); 2 cf, 1 9, Rutshuru, x.1937 (Ghesquiere) (1 cf in
BMNH); 2 cf , Lokandu, I. Biawa, vii.1939 (Vissers) (1 in BMNH); 1 9, Yangambi, 1940 (I.N.E.A.C.); 1
Cf , Bas-Congo, Mayidi, 1942 (Van Eyen); 2 cf , 1 9, Upemba, N.P., Gorges de la Pelenge, 6-10.vi.1947
(de Witte) (1 cf in BMNH); 1 cf , Upemba N.P.-, Kaswabilenga, 16.x. 1947 (de Witte); 1 9, Upemba N.P.,
Kilwezi, R. Lufira, 9-14.viii.1948 (de Witte); 1 cf, 2 $, Kivu, Matale, 8-14.V.1949 (Laurent) (1 9 in
BMNH); 1 cf , Kavuma-Costermansville, 16.vi.1949 (Laurent); 1 cf , Costermansville-Nyagezi, 20.vi.1949
194
B. C. TOWNSEND
G. africana
G. debilis
G. devia
Fig. 48 Map of known distribution of Gryllotalpa africana, G. debilis and G. devia.
(Laurent); 1 9, Urundi, Kigwena, 780 m, 9.xii.l949 (Laurent); 1 cf, Urundi, Kibaro, 1250 m, 15-
19.xii.1949 (Laurent) (BMNH); 1 cf, 1 $, Kivu, Kalehe Makwe, ii.1950 (Bomans); 9 cf, 6 $, 1 n,
Garamba N.P., 18.iv.1950, 30.viii.1950, 12.x. 1950 (De Saeger, Demoulin) (2 cf, 2 9 in BMNH); 1 9,
Albert N.P., plaine Baulendu, Semliki, 21. ii. 1951 (de Wilde); 1 9> Equateur, Bokuma, i.-ii.!952
(Lootens); 1 cf , Albert N.P., Kibanda, Lume, 7.xii.l956 (Vanschuytbroeck); 1 cf , Kivu, Uvira, Luvunyi,
5.xii.l961 (Kiss). Kenya: 1 9, Kakamega, xi.1976 (Clifton) (NMK). Tanzania: 1 cf (Lemaire). Rwanda: 3
9, Nyangwe, viii.-ix.1946 (Scholl). In MRAC unless otherwise stated.
DISTRIBUTION (Fig. 51). Equatorial Africa.
AFROTROPICAL MOLE-CRICKETS
195
G. robust a
G. buffo
G. rufescens
Fig. 49 Map of known distribution of Gryllotalpa robusta, G. bulla and G. rufescens.
Gryllotalpa parva sp. n.
(Figs 21, 33, 46, 50, 56, 62)
C? . Fairly uniform in colour, usually fairly light brown, veins of fore wings a little darker. Pronotum not
unusually large compared with head. Lobes of metanotum not exposed, concealed by pronotum and base
of fore wings, never enlarged. Right fore wing as in Fig. 33; stridulatory area fairly rectangular; /?i and Rs
fused. Stridulatory file of right fore wing as in Fig. 21, with 40-70 teeth (mean of 41 examined: 55-5), fairly
evenly spaced, 26-4-33-3 per mm (mean of 41 examined: 30-0). Hind wings long, extending well beyond tip
of abdomen. Hind tibiae armed above with 2-4 spines on internal margin. Genitalia small, with short
ventral processes. Song as in Figs 56, 62, a continuous trill; mean syllable repetition rate 76-4-81 -4/s, mean
carrier frequency 2-9-3-3 kHz (based on 2 recordings).
$ . Right fore wing as in Fig. 46, RI and Rs fused.
MEASUREMENTS
Body length
Median length of pronotum
Length of hind femur
Length of fore wing
Length of stridulatory area
Width of stridulatory area
Length of stridulatory file
Males
(41):
19
•3-26
•0,
mean 22
•1
(41):
5
•8-
7
•7,
mean
6
•7
(41):
5
•8-
7
•7,
mean
6
•8
(41):
8-2-11
•o,
mean
9
•3
(40):
3'
•0-
3
•7,
mean
3
•4
(40):
1
•5-
2
•1,
mean
1
•9
(41):
1
•3-
2
•2,
mean
1
•9
Females
(50): 17-8-28-0, mean 22-5
(50): 5-7- 8-3, mean 6-8
(50): 5-8- 8-5, mean 6-9
(50): 7-6-12-0, mean 9-6
196
B. C. TOWNSEND
A G. e/egans
• G. sptssidens
• G. parva
Fig. 50 Map of known distribution of Gryllotalpa elegans, G. spissidens and G. parva.
DISCUSSION. G. parva is most similar to brevity ra, from which it may be distinguished by the
shape of the stridulatory area and the density of the stridulatory teeth of the males, although
females are inseparable. G. parva has a characteristically rectangular stridulatory area, and
lacks the prominent fore wing veins of elegans.
MATERIAL EXAMINED
Holotype cf , South Africa: Natal, Eshowe, 30.X.1980 (Otte) (ANS).
Paratypes. South Africa: 1 cf , 1 n, same data as holotype (ANS). Tanzania: 1 cf , Lake Tanganyika
(Cunnington) (BMNH); 1 cf , Bukoba, xii.1921 (Miller) (BMNH); 1 cf , Mahagi Peninsula, Kasoge, 2550
feet, viii.-ix.1959 (Oxford University Expedition) (BMNH). Zaire: 1 cf, Kasai, Lukenge (Fontainas)
(MRAC); 1 cf , Ed. Luja, Kondue (BMNH); 1 cf , Sankuru, Beni Bendi (Cloetens); 1 cf , 1 $, Bohor; 1 cf ,
Kinshasa ('Leopoldville') (Wilverth); 1 cf , Mayumbe, 3.xi.l912 (Verschueren) (MRAC); 1 cT, 2 $ , Boma,
28.iii.1913, 5.vii.l920 (Styczynski, Schouteden) (MRAC); 2 cf, Kisangani ('Stanleyville'), xii.1929 (Mul-
ler); 1 cf, Kisangani ('Stanleyville'), 6.vii.l932 (Vrydagh) (MRAC); 1 cf, 2 $, 1 n, Lulua, Kapanga,
iii.1933 (Overlaet) (1 cf, 1 $, 1 n in MRAC; 1 $ in BMNH); 1 cf, Haute Tshuapua, Skela, 1936
(Buckinckx) (MRAC); 1 cf , Buta, 1939 (Vanbreuscghem) (MRAC); 1 cf , Sankuru, Bakwanga, 5.vi.l939
(Bequaert) (MRAC); 1 cf, Yangambi to Yakusu, 12.iii.1940 (MRAC); 1 cf, Kisangani ('Stanleyville'),
5. v. 1948 (ZL); 1 cf, Tshuapua, Imbonga, xii.1952 (Lootens) (MRAC); 2 cf, 1 $, Ubangi, Bumba,
ii.-xii.1952 (Basilewsky) (1 cf , 1 $ in MRAC; 1 cf in BMNH); 1 cf , 3 $, Lake Tanganyika, Albertville,
14.viii.1953 (Verbeke) (1 $ in BMNH); 6 cf , 20 $ , Vitshumbi, i.1953, 31.iii.1953, x.1953 (Verbeke) (1 cf , 4
9 in BMNH); 10 cf , 31 £, Kasenyi, vi.1953, ll.xii.1953, 13.xii.1953, 15.xii.1953 (Verbeke) (3 cf , 7 $ in
AFROTROPICAL MOLE-CRICKETS
197
A. G. brevilyra
* G. m/cropfera
• G. pluridens
Fig. 51 Map of known distribution of Gryllotalpa brevilyra, G. microptera and G. pluridens.
BMNH); 1 Cf , Lake Albert, Ituri Sabe, 16.xii.1953 (Verbeke); 1 cf , 5 $ $>, Bezaka, 19.xii.1953 (Verbeke)
(1 $ in BMNH); 2 cf, Tshuapua Bamanya, vi.1954, iv.1961 (Hulstaert} (MRAC); 5 C?, 12 $, Ishango,
viii.1954 (Semliki) (1 cf , 3 $ in BMNH); 4 cf , 1 ?, Sankuru, Djeka, 1955-1956 (Roiseaux) (3 cf , 1 9 in
MRAC; 1 cf in BMNH). Somalia: 1 cf, Afmadu, 20-24.viii.1970 (MZSUS). Ethiopia: 1 cf , Lake Hora
Harsadi, Addas, c. 7000 ft, 4.xii.l926 (Scott) (BMNH). Benin: 1 cT,l $, Cotonow (MHN). In IRSNB
unless otherwise stated.
DISTRIBUTION (Fig. 50). Equatorial and southern Africa, also Madagascar. G. parva is the only
member of the parva-group which occurs south of the Southern Tropic.
Gryllotalpa pluridens sp. n.
(Figs 16, 29, 42, 51)
Cf . Usually fairly uniform in colour, sandy yellow to dark brown, veins of fore wings a little darker, head
dark even in lighter specimens. Pronotum not unusually large compared with head. Lobes of mesonotum
not exposed, concealed by pronotum and base of fore wings, never enlarged. Right fore wing as in Fig. 29;
stridulatory area rather square, sometimes narrowing posteriorly; R{ and Rs fused. Stridulatory file of right
fore wing as in Fig. 16, with 76-107 teeth (mean of 23 examined: 98-0) fairly evenly spaced, 30-4-43-3 per
mm (mean of 23 examined: 35-8). Hind wings long, extending well beyond tip of abdomen. Hind tibiae
armed above with 3-4 spines on internal margin. Genitalia small, with short ventral processes.
$. Right fore wing as in Fig. 42, R{ and /?s normally fused; if separated, then R{ generally rather faint.
198
B. C. TOWNSEND
N
X
o
o
O
o
'£
>^
fi
8.S
•S §
V5 fl-
C * \ i '
II
51
a
AFROTROPICAL MOLE-CRICKETS
199
N
I
o
o
3
-O
c
cr
o -c
00 00
•2 .S
1 o
-
CN
CO
200 B. C. TOWNSEND
MEASUREMENTS
Males Females
Body length (24): 24-0-34-6, mean 28-4 (13): 21-5-32-9, mean 28-9
Median length of pronotum (24): 7-8-10-2, mean 9-2 (13): 8-0- 9-3, mean 8-8
Length of hind femur (23): 7-6-10-1, mean 8-8 (13): 7-9-10-3, mean 8-9
Length of fore wing (24): 10-5-14-2, mean 12-0 (13): 10-7-13-3, mean 12-0
Length of stridulatory area (21): 3-2- 4-4, mean 3-9
Width of stridulatory area (21): 2-1-3-0, mean 2-4
Length of stridulatory file (23): 2-4- 3-2, mean 2-7
DISCUSSION. The venation of the fore wing, arrangement of stridulatory teeth, and width of the
stridulatory area distinguish males of pluridens from all other species except spissidens. They
differ from this species principally in the overall shape of the stridulatory area, which is much
squarer in pluridens, and in never having unusually prominent veins. Females of pluridens and
spissidens are inseparable from each other, and cannot be distinguished reliably from those of
the africana-group.
MATERIAL EXAMINED
Holotype cf , Chad: Bebedjia, xi.1965 (Schmitz) (MRAC).
Paratypes. Chad: 4 cf, 6 $, same data as holotype (2 cfcf, 2 9$ in BMNH); Ghana: 1 cf, 3 $,
Trans- Volta-Togoland, Kpandu, 23.xii.1959, 28.xii.1959, 3. i. 1960 (Jago) (BMNH). Cameroon: 3 <7,
M'Bakaou, iii.1971 (Chemin) (1 in BMNH). Zaire: 1 cf, Mayumbe, Luki (Engleberi) (IRSNB); 1 cf,
Katomoja (Lemaire); 1 cf , Eala, 2.H.1923 (Oye) (BMNH); 1 cf , Kibali-Ituri, Mahagi, 1934 (Scops); 1 $,
Eala, xi.1934 (Ghesquiere); 1 cf , Lulua, Luashi, 1936 (Faiyne); 1 cf , 1 9, Upemba N.P., Masombwe R.,
Kanakakazi, 1120 m, 4-16.X.1948 (de Witte) ($ in BMNH); 1 cf, Garamba N.P., Akam, 13.xii.1949
(Demoulin); 3 cf , 2 $, Garamba N.P., 2.L1950, 23. i. 1950, 3.V.1950, 27.ix.1950 (DeSaeger, Demoulin) (2
Cf in BMNH); 1 cf , 1 $, near Bolobo, Dwa, 1950, xii.1951 (N'Gwe); 1 cf , L. Leopold II, Bokoro, 1952
(Jans); 1 cf , 1 $, Mayumbe, T. Kipanzu, Singa to Mbomba, v.-vi.!958 (Laurent); 1 cf , Mayumbe, Kitadi,
viii.1958 (Laurent); 1 cf, Buie, 27.iii.-5.iv.1975 (s.b.z.) (BMNH) Bioko (Fernando Poo): 1 cf, 2 $
(Nicholls) (BMNH). In MRAC unless otherwise stated.
DISTRIBUTION (Fig. 51). Central Africa, its range overlapping that of spissidens from Ghana to
Cameroon.
Gryllotalpa spissidens sp. n.
(Figs 17, 30, 43, 50)
Cf . Fairly uniform in colour, light to dark brown, veins of fore wings a little darker. Pronotum not unusually
large compared with head. Lobes of mesonotum not exposed, concealed by pronotum and base of fore
wings, never enlarged. Right fore wing as in Fig. 30; stridulatory area rectangular; RI and Rs fused.
Stridulatory file of right fore wing as in Fig. 17, with 72-96 teeth (mean of 19 examined: 85-9), fairly evenly
spaced, 33-5^5-2 per mm (mean of 19 examined: 39-3). Hind wings long, extending well beyond tip of
abdomen. Hind tibiae armed above with 3-4 spines on internal margin. Genitalia small, with short ventral
processes.
9- Right fore wing as in Fig. 43, RI and Rs normally fused; if separated, then RI generally rather faint.
MEASUREMENTS
Males Females
Body length (19): 23-0-31-0, mean 27-8 (23): 23-6-31-8, mean 28-4
Median length of pronotum (19): 7-6- 9-5, mean 8-6 (23): 7-0- 9-2, mean 8-3
Length of hind femur (18): 7-2- 8-7, mean 8-2 (21): 7-5- 9-0, mean 8-2
Length of fore wing (19): 9-2-11-8, mean 10-6 (23): 9-8-12-8, mean 11-0
Length of stridulatory area (19): 3-4- 4-4, mean 3-9
Width of stridulatory area (19): 1-8- 2-2, mean 2-0
Length of stridulatory file (19): 1-9- 2-8, mean 2-2
DISCUSSION. This species is very similar to elegans, from which it may be distinguished by its
many, densely packed stridulatory teeth. Some specimens have rather prominent fore wing
veins, with poorly marked cross-veins, as in elegans, but this feature is less pronounced in
AFROTROPICAL MOLE-CRICKETS 201
spissidens. The distribution oielegans is quite different (see below). G. spissidens is also close to
pluridens, differing in the shape of the stridulatory area (Figs 29, 30).
MATERIAL EXAMINED
Holotype cf , Cameroon: Efulun, 10. ii. 1923 (Weber) (ANS).
Paratypes. Cameroon: 1 cf, 1 $, Bitye, Ja River, 3 cf, 7 $, 2 n, Efulun, 25. ix. 1920, 20.xi.1920,
22.xi.1920, 24.X.1922, 10.ii.1923, 10.iii.1923, 13.iii.1923, 18.iii.1923, 21.xi.1924 (Weber) (1 cf, 1 $ in
BMNH); 5 cf, 7 $, Lolodorf, 2.iv.l920, 10.xii.1920, 16.ii.1921, ii.1921, 27.ix.1922, 2.ii.l923, x.1923,
2.H. 1924, 4.vii. 1924, Hi. 1925 (Good) (2 cf , 2 $ in BMNH) ; 1 cf (Conrad);2 $ , Harbel, 26.xii. 1944, 1945-7
(Fox, Beany); 1 cf, 1 $, Victoria, 15. xi. 1949 (Oldroyd) (BMNH). Nigeria: 1 cf, Oban District (Talbot)
(BMNH). Liberia: 1 cf, Monrovia, 23.xi.1947 (Olsen) (ZL); 2 cf, 5 $, Marshall Territory, 10. i. 1955,
21.iv.1955, 12.V.1955, 27. v. 1955, 2.xi.l955, 3.ii.l956, 7.L1957 (Fox) (2 $ in BMNH). Sierra Leone: 1 cf
(Morgan) (BMNH). Guinea: 5 $, near Serokoro ('Zerekore'), 16.iii.-18.iv.1950 (Olsen) (4 in UZM; 1 in
BMNH); 2 Cf , 8 $, Kindia, 1964-5 (Dedycker) (1 cf , 7 $ in MRAC; 1 cf , 1 $ in BMNH). In ANS unless
otherwise stated.
DISTRIBUTION (Fig. 50). Restricted to the coastal rainforest of West Africa and the Cameroon,
where it replaces elegans.
Acknowledgements
I thank the following who kindly lent type-material or other specimens from their respective
institutions:
Dr P. Basilewsky, Mr M. P. Clifton, Mr J. Deeming, DrsM. Donskoff, M. Dorn, H. Enghoff,
K. K. Gunther, B. Hauser, A. J. Hesse, P. Johnsen, D. Otte, MrM. J. Scoble, DrsG. Schmitz,
P. Vanschuytbroeck, M. Vannini and S. Mascherini.
Mr D. Moyer, Sumbawanga Mission, Tanzania, sent me live specimens ofrobusta. Dr D. Otte
generously made available to me tape recordings and specimens from his recent visit to South
Africa. Dr W. J. Bailey kindly allowed me to use his tape recordings of rufescens, and presented
them to the British Museum (Natural History). Prof. H. Striimpel provided information on the
syntypes of minor.
I am also grateful to Mr W. J. Reynolds for providing song analyses from the tape recordings
used, and to my wife for typing the manuscript.
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AFROTROPICAL MOLE-CRICKETS
Index
Principal references are in bold; synonyms are in italics.
203
africana 176, 177, 179, 182, 183, 187
africana-group 179, 180, 182, 200
Austrotalpa 179
brevilyra 179, 182, 189, 191, 196
bulla 179, 180, 182, 183, 184
colini 182, 183
confusa 182, 183
Curtilla 178, 179
debilis 177, 179, 182, 184, 185, 187, 193
devia 178, 179, 180, 182, 185, 186, 189
elegans 179, 180, 182, 191, 192, 193, 196, 200, 201
fossor 182, 183, 186, 187
Gryllidae 175
Gryllotalpa 175, 176, 177, 178, 179, 180, 181, 183.
185, 186, 193
gryllotalpa 178, 179
Gryllotalpella 178
Gryllotalpidae 175, 178
Gryllotalpinae 178
madecassa 178
microptera 177, 179, 182, 185, 193, 194
minor 176, 184, 185
minuta 175, 184, 185
Neocurtilla 178, 186
orientalis 176, 183, 185, 187
parva 179, 182, 191, 195, 196, 197
parva-group 179, 180, 189, 197
pluridens 179, 182, 197, 200, 201
robusta 179, 182, 184, 186, 187, 188
rufescens 179, 180, 181, 189
Scapteriscinae 178
Scapteriscus 178
spissidens 179, 180, 182, 192, 193, 200, 201
Triamescaptor 178
British Museum (Natural History)
Chance, change & challenge
Two multi-author volumes from one of the foremost scientific institutions in the world.
General Editor: P. H. Greenwood
The Evolving Earth
Editor: L. R. M. Cocks
The Evolving Biosphere
Editor: P. L. Forey
In the first volume, The Evolving Earth, twenty scientists have been asked to review the present
state of knowledge in their particular field, ranging from the origin of the Earth, through ocean
sediments and soils to continental drift and palaeogeography.
In the companion volume, The Evolving Biosphere, museum scientists have chosen an
evolutionary concept — speciation, coevolution, biogeography etc. and related this to the group
of animals or plants in which they are specialising. Thus beetles and birds exemplify sympatric
and allopatric speciation, butterflies mimicry and certain fishes explosive evolution.
In both volumes the text is supplemented by over one hundred specially commissioned pieces of
two-colour artwork.
These two books will be invaluable to all sixth-form and undergraduate biology and geology
students.
The Evolving Earth: 276 x 219 mm, 280pp, 138 line illustrations, 42 halftones
The Evolving Biosphere: 276 x 219 mm, approx. 320pp, 133 line illustrations
Published: May 1981
Co-published by the British Museum (Natural History), London and Cambridge University
Press, Cambridge.
Titles to be published in Volume 46
The generic and tribal classification of spore-feeding Thysanoptera (Phlaeothripidae: Idolo-
thripinae).
By L. A. Mound & J. M. Palmer.
A revision of the Afrotropical mole-crickets (Orthoptera: Gryllotalpidae).
By B. C. Townsend.
Key to the genera of galerucine beetles of New Guinea, with a review of Sastra and related new
taxa (Chrysomelidae).
By Sharon L. Shute.
The Afrotropical dacetine ants (Formicidae).
By Barry Bolton.
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd., Dorchester
29 JUL
Bulletin of the ^"
British Museum (Natural History)
Key to the genera of galerucine beetles
of New Guinea, with a review of
Sastra and related new taxa
(Chrysomelidae)
Sharon L.Shute
Entomology series
Vol 46 No 3 28 July 1983
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,
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Papers in the Bulletin are primarily the results of research carried out on the unique and
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Wo rid List abbreviation: Bull. Br. Mas. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1983
28 JUL5733
The Entomology series is produced under the general editorship of the
Keeper of Entomology: Laurence A. Mound
Assistant Editor: W. Gerald Tremewan
ISSN 0524-6431 Entomology series
Vol46No3pp205-266
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 28 July 1983
1
Key to the genera of galerucine beetles of New
Guinea, with a review of Sastra and related new
taxa (Chrysomelidae)
Sharon L. Shuttle
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis 205
Introduction 205
Depositories 206
Acknowledgements 207
Key to genera of Galerucinae from New Guinea 207
Dreeusgen. n 215
Marmina gen. n 217
Key to species of Marmina 219
Polysastra gen. n 220
Key to species groups based on external characters 222
Key to described species and subspecies of Polysastra 229
SastraBaly 245
Species remaining in Sastra 247
Species removed from Sastra 256
Check list of genera, species and subspecies treated here 257
References 257
Plant index 264
Index 265
Synopsis
A key is given to the 47 genera of the chrysomelid subfamily Galerucinae represented in New Guinea. The
current status of the 34 species previously listed in the genus Sastra Baly is reviewed; 22 new combinations
are proposed and four combinations are reinstated. Three genera, 14 species and two subspecies are
described as new. One specific synonym is newly established, and one species is recalled from synonymy.
Two of the genera have been erected primarily for the inclusion of the New Guinean species with toothed
pronota which were previously assigned to the genus Sastra. Keys, figures of the diagnostic features and
distribution maps are provided.
Introduction
The Chrysomelidae is one of the largest coleopterous families with upwards of 50,000 described
species distributed throughout the world. The adults and larvae are entirely phytophagous,
therefore the family is one of the most economically important amongst the Coleoptera. The
majority of species tend to be host-specific or are restricted to one particular plant family
although, compared with the number of species in the family, relatively few of the host plants
have been recognised.
The genera dealt with in this paper belong to the Galerucinae, which is the largest of the 17
subfamilies recognised in the Chrysomelidae (for key, see Gressitt & Kimoto 1963) and has over
5,000 described species distributed throughout the world; these are most abundant in tropical
regions. Adult Galerucinae feed on the parenchyma of the leaves and many species are known to
visit flowers (Knuth, 1908; Maulik, MS.). The larvae may be either leaf- or root-feeders. In the
majority of species pupation takes place in the soil but in others the pupae are attached to leaves.
As a result of making routine identifications of Galerucinae from New Guinea it became
Bull. Br. Mus. not. Hist. (Ent.) 46 (3): 205-266 Issued 28 July 1983
206 SHARON L. SHUTE
apparent that many of the genera occurring in this region are inaccurately and poorly defined in
the literature. This has resulted in new species being assigned to the wrong genera, and genera
being placed in the wrong tribe. Diagnosis of genera which are widespread in Asia and occur in
New Guinea are provided by Gressitt & Kimoto (1963), while Gressitt & Hornabrook (1977)
give a key to the principal beetle families represented in New Guinea. The latter work includes a
section on the Galerucinae, with a list of the principal genera found in New Guinea. However,
many problems remain in identifying Galerucinae from this region. The present work aims to
rectify the most serious omission by providing a comprehensive key to the New Guinean genera
of the subfamily. Wherever possible this key is based on type-species and takes into account
undescribed species and genera as well as those already described. Several genera not previously
recorded from this region are included.
In the present study special attention has been paid to Sastra which was found to constitute a
particularly heterogeneous assemblage of species; of the 35 species listed by Wilcox (1971) only
nine are now retained in this genus. The type-material of these nine species is listed and a
redefinition of Sastra is provided. All nine species are from New Guinea, and a study of further
undescribed species indicates that the geographical range of this genus is confined to this region
(Map 1). Eleven species from New Guinea, previously assigned to Sastra, were found to
constitute two new genera which are described here; these are principally characterised by
having a tooth-like process on the lateral margin of the pronotum. The largest of the two genera,
Polysastra, has been erected primarily for the inclusion of nine of the above mentioned species,
but I have examined a further 43 undescribed species. A full revision of these is not included in
the present paper but 16 are described below as representatives of the 'species groups' (see
discussion under Polysastra) into which the genus has been divided. It is hoped that this
preliminary study will facilitate further research by future workers.
A third new genus from New Guinea has also been described here as the species it contains are
frequently incorrectly assigned to Sastra. This genus is also interesting as it exhibits characters
uncommon in the Galerucinae. All genera, groups and species have been dealt with in
alphabetical order. Genitalia and other diagnostic features are illustrated for as many species as
possible.
Depositories
AMNH American Museum of Natural History, New York, U.S.A.
ANIC Australian National Insect Collection, Division of Entomology, CSIRO, Canberra, Austra-
lia.
BMNH British Museum (Natural History), London, U.K.
BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A.
DPI Department of Primary Industry, Konedobu, Papua New Guinea.
IP Institut fur Pflanzenschutzforschung, Eberswalde , D .D . R.
ITZ Institut voorTaxonomischeZo61ogie,Zoologisch Museum, Amsterdam, Netherlands.
MCSN Museo Civico di Storia Naturale, Genoa, Italy.
MCZ Museum of Comparative Zoology , Harvard University , Cambridge , Massachusetts , U . S . A .
MIZSU Museo ed Istituto di Zoologia Sistematica dell' Universita, Turin, Italy.
MNHN Museum National d'Histoire Naturelle, Paris, France.
MNHU Museum fur Naturkunde der Humbolt-Universitat, Berlin, D.D.R.
NMV National Museum of Victoria, Melbourne, Victoria, Australia.
NR Naturhistoriska Riksmuseet, Stockholm, Sweden.
OIP Ohlmus collection, Department of Primary Industry, Konedobu, Papua New Guinea.
RHN Rijksmuseum van Natuurlijke Historic, Leiden, Netherlands.
RWH R. W. Hornabrook collection, Wellington, New Zealand.
SAM South Australian Museum, Adelaide, Australia.
SMT Staatliches Museum fur Tierkunde, Dresden, D.D.R.
USNM National Museum of Natural History, Washington, D.C. , U.S.A.
UQ University of Queensland, St Lucia, Queensland, Australia.
GALERUCINE BEETLES OF NEW GUINEA 207
Acknowledgements
I extend my grateful thanks to the following specialists who have been kind enough to lend me
type-specimens or other material from their institutions listed above. Mile N. Berti, the late Mrs
D. H. Blake, Dr J. P. Duffels, Dr F. Hieke, Dr R. W. Hornabrook, Dr J. Krikken, Dr E. G.
Matthews, Dr G. B. Monteith, Dr A. Neboiss, Dr T. Nyholm, Mr H. Ohlmus, Prof. U. Parenti,
Dr J. N. L. Stibick (formerly DPI), Dr E. Tortonese, Dr J. White (formerly MCZ), and Dr R. E.
White. I also thank all my colleagues who have offered help and advice. My very special thanks
are due to Dr G. A. Samuelson of the Bishop Museum without whose help and patience this
paper would not have been possible. I am also indebted to him for the loan of the principal part
of the material on which the work is based. Finally I thank Mr T. Nagatani (BPBM) for the
figures of the whole specimens, Mrs B. Wavell-Craven for typing the species descriptions and
the BMNH photography unit for the photographs.
Key to genera of Galerucinae from New Guinea
1 At least three-quarters of length of antennal socket behind mid-line of eye. Length of
frons from anterior margins of antennal sockets to base of labrum not less than that of
socket.
Tarsal claws appendiculate . Frons never forming a transverse ridge 2
At least three-quarters of length of antennal socket in front of mid-line of eye . Length of
frons from anterior margins of antennal sockets to base of labrum not more than that
of a socket 14
2 (1) Pronotum with distinct dorsal depressions. Apex of tibiae with or without a ventral spur 3
Pronotum without dorsal depressions. Apex of tibiae with a ventral spur 6
3 (2) Length of gena at narrowest point less than half that of eye . Apex of tibiae with a ventral
spur. Basal segment of antenna distinctly longer than third segment. Elytron becom-
ing broader in apical half. Basal segment of hind tarsus equal to or longer than rest of
segments combined 4
Length of gena at narrowest point clearly exceeding half that of eye. Apex of tibiae
without a ventral spur. Basal segment of antenna only slightly longer than third
segment. Elytra parallel-sided. Basal segment of hind tarsus not longer than segments
2and3 combined COLEOCRANIA Jacoby
4 (3) Distance between antennal sockets at least half width of socket. Basal segment of
antenna equal to or shorter than segments 2 and 3 combined. Length of frons from
anterior margins of antennal sockets to base of labrum not greater than 2-5 x that of a
socket 5
Distance between antennal sockets less than half width of socket. Basal segment of
antenna at least l-5x as long as segments 2 and 3 combined. Length of frons from
anterior margins of antennal sockets to base of labrum at least 3-0 x that of a socket.
LUPERUSMiMer
5 (4) Body length not exceeding 10 mm. Basal segment of hind tarsus l-5x as long as rest of
segments combined. Eye at least 4-Ox the length of an antennal socket. Antennal
segments slender and individually straight. Pronotum not more than l-4x broader
than long, posterior half with shallow sublateral depressions which are sometimes
confluent, forming a single transverse depression. Elytron not more than 3-5 x length
ofpronotum NEOLEPTA Jacoby
Body length at least 15 mm. Basal segment of hind tarsus shorter than rest of segments
combined. Eye not more than 3-Ox length of an antennal socket. Antennal segments
slender, segments 4-11 individually curved. Pronotum not less than l-9x broader
than long. Elytron at least 5-Ox length ofpronotum (unevenly convex).
PALAEOSASTRA Jacoby
6 (2) Basal segment of antenna distinctly shorter than segments 2 and 3 combined, third
segment 4-5 x as long as second, segments 3-8 individually curved. Pronotum not
more than 1 -7 x broader than long. Elytra subparallel-sided. Body length not exceed-
ing7-0mm ALOPENA Baly
Basal segment of antenna as long as or longer than segments 2 and 3 combined. Other
characters not so combined . . 7
208 SHARON L. SHUTE
7 (6) Lateral margin of pronotum with a small rounded tooth at about middle. Length of frons
from anterior margins of antennal sockets to base of labrum at least 2-5 x length of
socket. Basal segment of antenna at least twice as long as frons. Elytron c. 4-Ox length
of pronotum. Antenna and legs slender. Basal segment of hind tarsus slightly longer
than segments 2 and 3 combined. Dorsum appearing glabrous. . . ASTRIDELLA Laboissiere
Lateral margins of pronotum without a median tooth . Other characters not so combined 8
8 (7) Length from hind margin of antennal sockets to hind margin of eyes not more than 0-25 x
length of socket. Length of frons from anterior margins of antennal sockets to base of
labrum 2- 5-3 -Ox that of socket 11
Length from hind margin of antennal sockets to hind margin of eyes at least 0-75 x that of
socket. Length of frons from anterior margins of antennal sockets to base of labrum
not more than 2-25 x that of socket 9
9 (8) Frons with a longitudinal median carina. Length of gena at narrowest point more than
half that of antennal socket. Apex of tibiae without a ventral spur. Basal segment of
hind tarsus shorter than segments 2 and 3 combined YULENIA Jacoby
Frons without a longitudinal median carina. Length of genat at narrowest point distinctly
less than half that of antennal socket. Apex of tibiae with a ventral spur. Basal segment
of hind tarsus longer than segments 2 and 3 combined 10
10 (9) Outer margin of antennal socket contiguous with inner margin of eye, eye large, width
greater than that of area between eyes. Distance between antennal sockets not more
than half width of socket. Length of third antennal segment equal to or shorter than
second. Pronotum widest at middle, sides evenly rounded. Elytra parallel-sided and
evenly convex. Basal segment of hind tarsus as long as rest of segments combined
NEODRANA Jacoby
Outer margin of antennal socket separated from inner margin of eye by at least half a
sockets width. Eye moderately large, width less than that of area between eyes.
Distance between antennal sockets more or less equal to width of socket. Length of
third antennal segment 1-5-2-Ox that of second. Pronotum widest in anterior half,
width decreasing gradually towards base. Elytra subparallel-sided, greatest convexity
of elytron about middle . Basal segment of hind tarsus 1 -25 x length of rest of segments
combined LOMIRANA Laboissiere
11 (8) Length of gena at narrowest point equal to or exceeding half that of antennal socket 12
Length of gena at narrowest point distinctly less than half that of antennal socket,
emargination of gena usually contiguous with anterior margin of eye.
Body length not exceeding 7 mm 13
12 (11) Body length exceeding 7 mm. Pronotum at least 2-2 x broader than long. Width between
antennal sockets at least that of socket. Antennal segments 4-11 individually curved
PRASYPTERA Baly
Body length not exceeding 7 mm. Pronotum not more than l-4x broader than long.
Distance between antennal sockets not more than half width of socket. Antennal
segments not curved LUPERUS Miiller
13 (11) Apical segment of maxillary palp stout, shorter than preceding segment. Frons with a
small median longitudinal carina. Third antennal segment equal in length to second,
fourth segment at least 4-5 x length of third. Pronotum not less than l-4x broader
than long. Basal segment of hind tarsus longer than rest of segments combined
NEODRANA Jacoby
Apical segment of maxillary palp slender, longer than preceding segment. Frons without
median carina. Third antennal segment at least 3x as long as second, fourth segment
l-25x length of third. Pronotum not more than l-2x broader than long. Basal
segment of hind tarsus shorter than rest of segments combined MICROLEPTA Jacoby
14 (1) Lateral margins of pronotum with a distinct, acute or rounded tooth-like projection at
about middle.
Third antennal segment distinctly longer than fourth. Fore coxal cavities open
behind . Tarsal claws bifid . Primary setal pores usually tuberculate 53
Lateral margins of pronotum never with a distinct median projection 15
15 (14) Tarsal claws bifid 16
Tarsal claws appendiculate 39
16 (15) Width of elytral epipleuron at level of mid coxa 2-0-3-Ox greater than at level of hind
coxa, epipleuron becoming obsolete or distinctly narrower around level of hind coxa.
GALERUCINE BEETLES OF NEW GUINEA 209
Elytron sometimes expanded laterally beyond epipleura. Pronotum with or without
dorsal depressions 17
Width of elytral epipleuron at level of mid coxa never more than l-5x greater than at
level of hind coxa, width of epipleuron either remaining more or less the same for at
least three-quarters of its length or becoming gradually narrower from around hind
coxa towards apices. Elytron not expanded laterally beyond epipleura. Pronotum
with one or more dorsal depressions 28
17 (16) Elytron distinctly costate. Pronotum transverse, greatest width equal to that of elytra at
humeral angles. Fore coxal cavities closed behind. Legs stout. Basal segment of hind
tarsus shorter than segments 2 and 3 combined. Dorsum setose PLERONEXIS Weise
Elytron not costate . Other characters not so combined 18
18 (17) Pronotum without doral depressions. Dorsum appearing glabrous 19
Pronotum with one or more dorsal depressions . Dorsum glabrous or setose 22
19 (18) Elytra broadly ovate. Elytron often expanded laterally beyond epipleura. Pronotum at
least 2-Ox broader than long 20
Elytra subparallel-sided. Elytron not expanded laterally beyond epipleura. Pronotum
never more than 1 -7 x broader than long NOTONICEA Hincks cf
20 (19) Apical segment of maxillary palp globose, almost entirely contained within preceding
segment, apex concave. Metasternum with a large anterior peg-like process projecting
forward between mid coxae. Expanded area of elytron not more than l-2x wider than
epipleuron. Legs and antennal segments stout. Elytra often banded or bicoloured
ANOIDES Weise
Apical segment of maxillary palp conical or subglobose, not more than half length of
segment contained within preceding segment. Other above characters not so com-
bined 21
21 (20) Metasternum with a large anterior peg-like process projecting forward between mid-
coxae. Elytron strongly convex, expanded area lateral to epipleura at least 1-5-2-Ox
wider than epipleura PARANOIDES Vachon
Metasternum without anterior peg-like process.. Elytron moderately convex, expanded
area lateral to epipleura not more than l-2x wider than epipleura MOROKASIAJacoby
22 (18) Elytra distinctly setose. Apex of tibiae without a ventral spur. Dorsum usually unicolor-
ous, rarely metallic 23
Elytra glabrous. Apex of tibiae with or without a ventral spur. Dorsum usually with
transverse bands or patches, sometimes partially or entirely metallic 24
23 (22) Head with vertex strongly convex, appearing glabrous and impunctuate. Pronotum at
least 2-5 x broader than long, maximum width less than that of elytra at humeri,
surface with deep transverse sublateral depressions and shallow median depression,
sparsely setose. Body length not usually exceeding 7 mm. Legs and antennal segments
slender BUPHONIDABaly
Head without vertex convex, densely setose and punctured throughout. Pronotum not
more than l-9x broader than long, maximum width equal to that of elytra at humeri,
surface densely setose with two more or less circular sublateral depressions in anterior
half, median area convex. Body length usually exceeding 10 mm. Legs and antennal
segments stout MENIPPUS Clark
24 (22) Elytra broadly ovate , elytron expanded laterally beyond epipleuron . Pronotum with two
small more or less circular sublateral depressions or a single, narrow transverse
depression in anterior half OWES Weber
Elytra not broadly ovate or elytron expanded laterally. Pronotum with sublateral
depressions which are either confluent or occupy almost two-thirds of the surface area 25
25 (24) Frons triangular, anterior third either depressed or declivous. Inter-antennal area with a
longitudinal median carina. Pronotum narrowing at level of dorsal depressions,
depressions narrow, often confluent, anterior third strongly convex. Elytra often
pyriform 27
Frons forming a more or less evenly convex ridge. Inter-antennal area without a median
carina. Pronotum not becoming narrower at level of dorsal depressions, depressions
broad, occupying almost two thirds of the surface area, anterior half convex at sides.
Elytra never pyriform 26
26 (25) Apical segment of maxillary palp subglobose. Male with a deep triangular emargination
at apex of abdomen; female with a small curved or triangular notch (may be obscured
210 SHARON L. SHUTE
by setae). Pronotal depressions present in both sexes GALERUMAEA Hincks
Apical segment of maxillary palp conical. Male with a shallow curved emargination at
apex of abdomen; female with apex evenly rounded. Male with pronotal depressions
absent or very weak NOTONICEA Hincks
27 (25) Apex of tibiae with a ventral spur (may be small or obscured by setae). Posterior
two-thirds of elytron distinctly convex, humeral area delimited by a distinct transverse
depression PAPUANIA Jacoby
Apex of tibiae without a ventral spur. Posterior two-thirds of elytron not distinctly
convex, humeral area never delimited by a transverse depression
AULACOPHORA Chevrolat
28 (16) Length of gena at narrowest point at least 0-67x that of antennal socket 29
Length of gena at narrowest point less than 0-5 x that of antennal socket (emargination
of gena usually contiguous with anterior margins of eye (Fig. 15) 33
29 (28) Elytra strongly rugose throughout, rugosities often forming long irregular carinae.
Males of some species with a large tooth on hind femur.
Eyes prominent. Frons triangular with a median longitudinal carina. Pronotum with
distinct lateral margins, dorsal depressions narrow. Legs and antennal segments
slender. (Figs 114, 115) genus A
Elytra not rugose or carinate . Hind femur never toothed in male 30
30 (29) Apex of tibiae with a ventral spur. Elytra pyriform, posterior two-thirds of elytron
distinctly convex, humeral area often delimited posteriorly by a narrow transverse
depression. Lateral margins of pronotum distinct. Third antennal segment shorter
than fourth. Dorsum appearing glabrous PAPUANIA Jacoby
Apex of tibiae without a ventral spur. Elytra parallel-sided to subovate, posterior
two-thirds of elytron not distinctly convex, humeral area never delimited by a
transverse depression. Lateral margins of pronotum absent or barely discernible.
Third antennal segment longer than fourth. Dorsum setose or glabrous 31
31 (30) Dorsum appearing glabrous. Puncturation indistinct on head and pronotum. Elytra
distinctly and evenly punctured, often metallic, sometimes bicoloured or banded
MOMAEA Baly
Dorsum distinctly setose. Puncturation strong and dense on head and pronotum. Elytra
strongly and densely punctured 32
32 (31) Frons medially convex with a strong anterior declivity. Eye at least three times as long as
the narrowest part of gena. Elytra parallel-sided, usually bi-coloured. Basal segment
of hind tarsus equal in length to that of segments 2 and 3 combined A TYSA Baly
Frons with sides convex, no anterior declivity. Eye not more than one and a half times as
long as the narrowest part of the gena. Elytron becoming gradually wider from base of
humeri onwards. Basal segment of hind tarsus shorter than segments 2 and 3
combined GALERUCELLA Crotch
33 (28) Frons with a median triangular depression, sides distinctly convex. Apical segment of
maxillary palp conical, longer than preceding segment. Eye large and prominent, at
least 3 x length of an antennal segment. Antennal segments slender, third segment at
least one and a half times longer than fourth segment. Pronotum transverse, sublateral
depressions distinct but often ill-defined. Elytra setose, densely and more or less
uniformly punctured throughout, surface evenly convex. Legs slender. Basal segment
of hind tarsus 0-5 x longer than preceding segment. (Figs 1 , 102-1 12) SASTRA Baly (p. 247)
Frons without median depression, being either medially convex or forming a more or
less evenly convex ridge. Other characters not so combined 34
34 (33) Elytra densely setose , puncturation distinct and dense 35
Elytra glabrous or only very sparsely setose , puncturation fine 37
35 (34) Anterior third of pronotum strongly convex across entire width, median region of
convex area narrower than rest, convex area distinctly elevated above posterior
two-thirds and less coarsely punctured. Eyes large and protuberant. Frons more or
less triangular, often declivous anteriorly. Elytra 3-4x longer than broad, parallel-
sided or becoming broader just before apices, some species with narrow longitudinal
ribs DKS/ODESWeise
Not as above 36
36 (35) Eye not more than twice the length of an antennal socket. Pronotum without lateral
margins, sublateral depression broad but not confluent. Inner ungue of tarsal claws
GALERUCINE BEETLES OF NEW GUINEA 211
equal in length to or only slightly shorter than outer ungue. Elytra parallel-sided, some
species with narrow longitudinal ribs. Antennal segments enlarged and flattened in
some species ATYSABa\y
Eye at least three times as long as antennal socket. Pronotum with narrow but distinct
lateral margins, sublateral depressions confluent, occupying at least two-thirds of
surface area. Inner ungue of tarsal claw not more than half length of outer segment.
Elytra becoming gradually broader from humeral area onwards. Antennal segments
filiform PONERIDEAWeise
37 (34) Frons triangular, sometimes medially convex and declivous, length from anterior
margins of antennal sockets to base of labrum usually exceeding that of a socket.
Antennal segments 3 and 4 subequal 38
Frons forming a more or less evenly convex ridge, length from anterior margins of
antennal sockets to base of labrum not exceeding that of a socket. Antennal segment
three at least 1 -4 x length of fourth MOMAEABaly
38 (37) Pronotum not more than l-4x broader than long, sides subparallel, sublateral depress-
ion indistinct, often confluent. Elytra parallel- or subparallel-sided. Antennal seg-
ments elongate and slender. Male with basal segment of all tarsi greatly enlarged.
Abdomen with two long conical appendages at middle of posterior margin of second
segment HOPLASOMA Jacoby
Pronotum at least 2-Ox broader than long, sides rounded, sublateral depressions
distinct, rarely confluent. Elytron becoming gradually broader from base of humeral
area onwards, apical quarter often broadly rounded. Antennal segments 6-11 short
and stout. Male with only basal segment of fore tarsus enlarged. Abdominal appen-
dages absent PAUMOMUA Jacoby
39 (15) Elytron with at least six narrow longitudinal ribs, punctures situated between ribs
(maybe weakly indicated) 40
Elytron without any indication of ribs, puncturation confused 42
40 (39) Apex of tibiae without a ventral spur. Pronotum at least 2-4x broader than long.
Distance between antennal sockets greater than width of a socket. Underside densely
setose. Legs moderately stout. Basal segment of hind tarsus shorter than segments 2
and 3 combined PLESISTIA Maulik
Apex of tibiae with a ventral spur. Pronotum not more than l-5x broader than long.
Distance between antennal sockets not more than width of a socket. Underside
sparsely setose . Legs slender. Basal segment of hind tarsus longer than segments 2 and
3 combined 41
41 (40) Elytral epipleuron becoming distinctly narrower above level of hind coxa, width then
decreasing gradually until epipleuron becomes obsolete prior to apex. Frons never
modified in male LOMIRANA Laboissiere
Elytral epipleuron not distinctly narrowed above hind coxa, width decreasing gradually
below level of hind coxa, epipleuron continuing to apex. Frons often excavated or
modified in male DIAPHAENIDEA Laboissiere
42 (39) Apical segment of maxillary palp distinct from preceding segment, being either subglo-
bose or conical . Third segment subcylindrical . Frons never excavated in male 43
Apical segment of maxillary palp situated in the apex of the inner angle of the preceding
segment, third segment greatly enlarged and triangular. Frons excavated in male.
Fore coxal cavities open behind. Distance between antennal sockets greater than
width of a socket. Basal segment of antenna longer than segments 2 and 3 combined.
Basal segment of hind tarsus longer than rest of segments combined
PALPAENIDEA Laboissiere
43 (42) Maximum width of elytral epipleuron at level of metepisternum equal to or exceeding
that of metepisternum , epipleuron becoming distinctly narrower at about level of hind
coxa . Dorsum appearing glabrous 44
Maximum width of elytral epipleuron at level of metepisternum not more than half that
of metepisternum, epipleuron either abbreviated at about level of mid metasternum
or fairly constant in width for entire length .Dorsum with or without setae 49
44 (43) Length of gena at narrowest point distinctly less than half that of antennal socket.
Pronotum without dorsal depressions 45
Length of gena at narrowest point not less than half that of antennal socket. Pronotum
with or without dorsal depressions 47
212 SHARON L. SHUTE
45 (44) Apex of tibiae with a ventral spur. Elytron becoming gradually broader from humeral
angles onwards, apical half convex, epipleuron becoming obsolete before apex. Legs
slender, hind tibia evenly curved. Basal segment of hind tarsus longer than rest of
segments combined 46
Apex of tibiae without a ventral spur. Elytron ovate, weakly convex, epipleuron distinct
to apex. Legs moderately slender, hind tibia straight. Basal segment of hind tarsus
shorter than segments 2 and 3 combined CASSENA Weise
46 (45) Pronotum not more than l-4x broader than long, sublateral depressions weak, some-
times confluent, forming a narrow transverse depression, maximum convexity of
pronotum in apical half, posterior margin evenly curved NEOLEPTA Jacoby
Pronotum at least l-6x broader than long, dorsal depressions absent, surface evenly
convex, posterior margin sinuate at sides. Vertex may be excavated in cf
MONOLEPTA Chevrolat
47 (44) Apex of tibiae without a ventral spur. Pronotum without dorsal depressions. Third
segment of maxillary palp enlarged, length at least 2-5 x that of apical segment.
Antennal segments 9-1 1 distinctly enlarged in cf CNEORANE Baly
Apex of tibiae with a ventral spur (may be small). Pronotum with or without dorsal
depressions. Third segment of maxillary palp not obviously enlarged, length not
more than l-5x length of apical segment. Apical segments of antenna not enlarged
incf 48
48 (47) Antennal segments stout, third segment 2-0-2-5 x length of fourth segment. Pronotum
at least l-7x broader than long, surface with two small more or less circular sublateral
depressions. Legs stout. Basal segment of hind tarsus slightly longer than segments 2
and3 combined SYNODITA Chapuis
Antennal segments slender, segments 3 and 4 subequal. Pronotum not more than l-3x
broader than long, dorsal depressions absent. Legs slender. Basal segment of hind
tarsus longer than rest of segments combined LOMIRANA Laboissiere
49 (43) Pronotum evenly convex, dorsal depressions absent. Apical segment of maxillary palp
subglobose, apex truncate. Antennal segments short and stout. Dorsum appearing
glabrous.
Length of gena at narrowest point not less than half that of antennal socket.
Maximum width of elytron in apical third. Puncturation fine. Basal segment of hind
tarsus shorter than segments 2 and 3 combined ASTENA Baly
Pronotum unevenly convex, dorsal depressions distinct. Apical segment of maxillary
palp conical. Antennal segments elongate and slender. Dorsum glabrous or setose 50
50 (49) Frons triangular, maximum convexity at middle, longitudinal median carina present.
Dorsum appearing glabrous or sparsely setose. Elytron usually pyriform, numeral
area delimited by a narrow transverse depression PARIDEA Baly
Frons forming a transverse ridge, maximum convexity at sides, longitudinal median
carina absent. Dorsum densely setose (if glabrous then length not exceeding 6 mm).
Elytron never pyriform , humeral area not delimited by a transverse depression 51
51 (50) Head and pronotum appearing impunctate. Elytron finely punctured throughout.
Dorsum either glabrous throughout or elytra sparsely setose. Length of eye not more
than twice that of an antennal socket. Body length not usually exceeding 5 mm. Hind
tibia evenly curved PHYLLOCLEPTIS Weise
Dorsum distinctly punctured and setose throughout. Length of eye at least 3 -Ox that of
antennal socket. Body length exceeding 6 mm. Hind tibia straight 52
52 (51) Length of gena at narrowest point equal to that of three eye facets combined. Maximum
convexity of frons at middle. Distance between antennal sockets not less than width of
a socket. Interantennal area with a small pit-like depression. Lateral margins of
pronotum barely discernible, sublateral depressions small and shallow, rarely con-
fluent. Elytra usually weakly metallic, punctures contiguous APOPHYLIA Thomson
Length of gena at narrowest point less than that of two eye facets combined. Maximum
convexity of frons at sides. Distance between antennal sockets less than width of a
socket. Interantennal area without depression. Lateral margins of pronotum distinct,
at least two-thirds of surface area depressed. Elytra non-metallic, punctures separated
by at least their own width PONERIDEAWeise
53 (14) Length of eye never more than 2-25 x that of an antennal socket or twice the width.
Length of gena at narrowest point at least half that of an antennal socket 54
GALERUCINE BEETLES OF NEW GUINEA
213
Length of eye at least 3-Ox that of an antennal socket and 2-5-3-Ox the width. Length of
gena at narrowest point less than half that of an antennal socket.
Male with a deep triangular emargination at apex of abdomen 56
54 (53) Antennae less than half the length of body, segments short and stout. Body shape
suggestive of Galeruca sp. Pronotal primary setal pores not tuberculate. Length of
metasternum not more than twice that of intercoxal process of mesosternum. (Fig. 40)
genus B
Antennae exceeding half body length, segments elongate and slender. Body elongate,
elytra parallel- or subparallel-sided. Pronotal primary setal pores tuberculate. Length
of metasternum at least 3-5 x that of intercoxal process of mesosternum 55
55 (54) Eye at least three times longer than shortest point of gena. Pronotum indistinctly
punctured, submedian lateral tooth small and rounded, anterior third and area
between outer margin of sublateral depression and base of tooth equally convex, disc
sometimes with a narrow ovate depression which may become confluent with sublater-
al depressions. Elytra more or less evenly convex, never rugose or carinate, rarely
metallic, usually bicoloured or with colour pattern. Basal segment of hind tarsus
evenly convex dorsally. (Figsll6-119) MARMINA gen. n. (p. 217)
Eye not more than twice as long as the shortest point of the gena. Pronotum strongly and
distinctly punctured, lateral margin with tooth well developed, surface unevenly
convex, sublateral depressions ill-defined, disc strongly convex with a small shallow
anterior and posterior depression. Elytra robust, surface rugose and unevenly convex.
Basal segment of hind tarsus tectiform GRONOVIUS Jacoby
56 (53) Male. Apex of abdomen emarginate, basal segment of fore or mid tarsi enlarged.
Antenna may have enlarged or modified segments 57
Female. Apex of abdomen evenly rounded or with small triangular emargination. Basal
segment of tarsi not enlarged . Antennal segments filiform , none enlarged or modified 58
57 (56) Antennal segments 4-11 short and stout, segments 9 and 11 distinctly enlarged and
ventrally flatterned (Figs 5, 6), these segments glabrous beneath. Mid tibia irregularly
curved (Fig. 4). Basal segment of mid tarsus with greater enlargement than fore tarsus
(Fig. 12). Elytra more or less evenly convex. (Fig. 3) DREEUSgen. n. (p. 215)
Antennal segments 4-11 elongate and slender, no modifications present (Figs 7, 8).
Basal segment of fore tarsus with greatest enlargement. Elytra with variable facies,
sometimes rugose, carinate or evenly convex. (Figs 12Q-150)POLYSASTRA gen. n. (p. 220)
58 (56) Antennal segments 4-11 stout, segment 11 twice as long as segment 10. Apex of
abdomen with a triangular emargination (Fig. 20). Elytron more or less evenly
convex. (Fig. 3) DREEUSgen. n. (p. 215)
Antennal segments 4-11 elongate and slender, segment 11 not more than l-25x longer
than segment 10. Apex of abdomen evenly rounded. Elytra with variable facies,
sometimes rugose, carinate or evenly convex. (Figs 120-150) POL YSASTRA gen. n. (p. 220)
Figs 1-3 General habitus of Galerucini genera treated here. 1, Sastra sp. cf. 2, Polysastra sp.
3, Dreeussp. cf.
214
SHARON L. SHUTE
14
Figs 4-15 Distinguishing generic characters. 4, Dreeus distinctus, mid tibia of male showing sinuation.
5-10, comparative proportions of antennal segments of (5) Dreeus sp. cf; (6) Dreeus sp. $; (7)
Polysastra sp. C? ; (8) Polysastra sp. $; (9) Marmina sp. cf; (10) Marmina sp. 9- H-13, Dreeus sp.,
comparative proportions of basal segment of fore, mid and hind tarsi. 14, 15, diagram of head showing
comparative emargination of genae of (14) Polysastra sp.; (15) Sastra sp.
GALERUCINE BEETLES OF NEW GUINEA 215
DREEUSgen. n.
Type-species: Dreeus distinctus sp. n.
GENERAL FORM. Body elongate, elytra subparallel-sided, appearing glabrous. Combined length of antennal
segments slightly less than half that of body, third segment at least 1 -5 x length of third segment, segments 9
and 10 enlarged and glabrous beneath in male (Figs 5-6). Eyes large and prominent. Pronotum transverse,
lateral margins produced into a tooth in anterior half, width at anterior margin less than that of head plus
eyes, maximum width more or less equal to that of head plus eyes and slightly less than that of elytra at
humeri, disc convex medially, with a small shallow depression anteriorly and posteriorly, and a sublateral
depression on either side. Primary setal pores weakly tuberculate. Elytra more or less uniformly convex,
usually unicolorous, epipleura becoming obsolete at about apical quarter. Underside setose, fore coxal
cavities open behind. Apex of abdomen in male with a triangular emargination which almost reaches hind
margin of segment (Fig. 20); female with a small triangular emargination at apex (Fig. 21). Legs slender,
mid tibia in male distinctly and irregularly sinuate in basal half (Fig. 4), hind tibia weakly curved, apices in
both sexes without ventral spurs or lateral spines. Basal segment of hind tarsus slightly longer than that of
segments 2 and 3 combined, basal segment of fore and mid tarsi slightly shorter, basal segment of mid
tarsus in male distinctly enlarged, length l-3x greater than that of basal segment of fore tarsus (Fig. 12).
Claws bifid.
DIAGNOSIS. Gena deeply emarginate medially, emargination reaching anterior margin of eye, maximum
width of gena less than half that of antennal socket. Apical segment of maxillary palp conical, length more
or less equal to that of preceding segment. Labrum small , not obscuring sides of mandibles. Pseudoclypeus
distinct, length almost equal to that of labrum. Frons with sides convex, length less than that of antennal
socket, sockets separated by less than the width of a socket, anterior margins level with those of eyes.
Postantennal swellings convex, distinctly elevated above level of vertex, contiguous with frons between
antennal sockets and separated medially by a narrow longitudinal groove which terminates just above
median depression of frons. Vertex with irregular puncturation, coronal 'suture' present. Eyes large and
prominent, length at least 2-4x that of antennal socket. Proportional lengths of antennal segments similar
in both sexes (Figs 5, 6), segments 9 and 11 enlarged in male. Pronotum irregularly punctured, width at
lateral tooth on average l-2x greater than minimum posterior width, tooth usually with apex acute,
depressions shallow, median depressions often joined by a narrow longitudinal groove. Scutellum
triangular, maximum width more or less equal to length, apex rounded. Elytra almost parallel-sided,
average length 3-6x greater than width, maximum width not more than l-2x greater than width at level of
apex of scutellum, posthumeral sublateral depression absent, surface more or less evenly convex,
appearing glabrous but minute setae confined within margins of punctures, puncturation irregular.
Underside minutely punctured throughout, metasternum convex, 2-8x longer than mesosternum, fore
coxal cavities open behind. Legs slender, hind leg 0-7x length of elytron and l-4x length of fore and mid
legs. Length of fore tibia slightly less than that of mid and hind tibiae, male with sinuate mid tibia (Fig. 4)
and weakly curved hind tibia. All femora more or less equal in width. Tarsi as in Figs 11-13. Aedeagus
forming a chitinous tube with a large complex apical process (Figs 22, 23). Basic structure of female
genitalia as in Polysastra; the principal differences are a distinct constriction between the vagina and bursa
(Fig. 34), distinctly longer styli (Fig. 37), and the proportions of the hemitergites (Figs 34, 37) and ligular
largely exposed (Fig. 34).
DISTRIBUTION. Irian Jaya, Papua New Guinea.
DISCUSSION. Compared with other genera found in this region, this genus is unusual in having
both the antennal segments and the mid tibiae modified in the male. It is also unusual amongst
the Galerucinae in having the maximum enlargement of the basal tarsal segment in the male on
the mid leg, not on the fore leg.
Several New Guinean species referable to this genus have been examined but only one is
described here as the others are not represented by sufficient material to make accurate
diagnoses. This genus does not appear to be closely related to the other genera from this region,
but belongs to the same group of primarily Australasian Galerucini, including Sastra and the
new genera described here, which are characterised by their elongate form and antennal
structure (Figs 1,2).
216 SHARON L. SHUTE
Dreeus distinctus sp. n.
(Figs 3, 4-6, 11-13, 20-24, Map 10)
GENERAL FORM. Length 6-9-11-3 mm. Elytra parallel to subparallel-sided, evenly convex, posthumeral,
sublateral, longitudinal carinae absent; elytral setae minute, not extending beyond margins of punctures,
sometimes a few sparse erect setae on intervals, puncturation confused. Pronotum transverse with a
distinct tooth on the lateral margins. Male with antennal segments 9 and 11 enlarged (Fig. 5). Head and
pronotum concolorous, dark reddish to pitchy brown. Elytron reddish brown, pitchy brown or dark green
with a weak aeneous lustre. Underside and legs in reddish species concolorous, dark testaceous to
orange-brown; pitchy brown and green specimens with abdomen pale testaceous and rest of underside
dark reddish to pitchy brown. Male with mid femur irregularly curved (Fig. 4), basal segment of fore and
mid tarsi enlarged, maximum enlargement in mid tarsi (Fig. 12).
DIAGNOSIS. Head with frons medially depressed, sides convex, elevation more or less equal to that of
postantennal swellings; swellings well defined and elevated above level of vertex, derm smooth and
shinning with weak microsculpture. Vertex coarsely and irregularly punctured, punctures large, shallow
and often confluent, derm distinctly microsculptured, setae long, fine and appressed. Pronotum trans-
verse, male on average l-6x broader than long, female l-75x broader than long, development of lateral
tooth variable, more acute in small specimens, one side sometimes more developed than the other in male,
lateral depressions distinct but shallow, median anterior depression shallow, delimited posteriorly by two
small admedian convexities, posterior median depression often ill-defined; puncturation dense and
irregular, size and density of punctures variable, size not normally greater than twice that of punctures on
humeral area of elytra, width apart not normally exceeding that of a puncture, derm distinctly microsculp-
tured, particularly in median depressions, setae minute, not extending beyond margins of punctures.
Scutellum triangular with broadly rounded apex, median area minutely punctured with short, appressed,
grey setae, derm microsculptured. Elytron 3-6x longer than broad. Elytra in male more or less
parallel-sided, sometimes slightly broader in apical quarter, female subparallel-sided, becoming gradually
broader from around middle to a maximum width just anterior to apical quarter. Maximum width c. l-3x
greater than the width at the level of the apex of the scutellum, apices evenly rounded in both sexes, derm
densely and irregularly punctured throughout giving a slightly rugose appearance, punctures not more than
their own width apart , setae minute , not extending beyond the margins of the punctures , epipleura narrow ,
extending to apex. Underside densely and minutely punctured, metepisternum weakly granulate, punc-
tures not separated by more than their own width, setae short and appressed, sides of metasternum with
puncturation and microsculpture as metepisternum, setae slightly longer and finer, median area of
metasternum, sparsely punctured, microsculpture composed of irregular, converging, horizontal lines,
setae longer and more erect than at sides. Femora with large shallow punctures 1-0-1-Sx their own width
apart on basal three-quarters, becoming denser at apices, setae short, fine and appressed. Tibiae granulate,
setae increasing in density towards apices, short and stout, basal segment of mid tarsus in male 1 -5 x as long
as basal segment of fore tasus. Apex of abdomen incised in both sexes (Figs 20-21). Wing fully developed.
Genitalia (Figs 22-24).
Holotype cf, Papua New Guinea: NE, Laiagam, 2180 m, 18-19. vi. 1963, m.v. light (/. Sedlacek)
(BPBM).
Paratypes. Papua New Guinea: 25 cf , 19 $ , same data as holotype (10 d", 6 $ BMNH); 7 cf , 15 $ , Lake
Siruki, 2550 m, 14-17.vi.1965 (3 cf , 3 $ BMNH); 10 cf, 10 ?, Kepilam, 2420-2490 m, 20-23. vi. 1963,
light-trap (all/. Sedlacek) (5 cf , 5 ? BMNH); 10 $ , Kotuni, south slope Mt Otto, 2200 m, 10-15.viii.1959
(5 $ BMNH); 1 cf , Purosa, camp Okapa, 1950 m, 27.ix.1959; 1 $, Kimi, Creek Camp, NE. slopes Mt
Michael, 1980 m, 29.viii.1959 (all L. T. Brass), (AMNH); 2 cf , Moke, i.x.1957 (/. Smart); !$,!?, Waisa
nr Okapa, c. 5000 ft, 15.ii.1965; 1 $, Okapa, c. 5000 ft, 4-13.ii.1965 (all M. E. Bacchus) (BMNH); 1 cf ,
1 $, Okapa, Purosa, 1700-2000 m, 18.i.l966; 4 cf, 3 $, 6-4 km W. Wabag, 2020 m, 13.vi.1963 (all /.
Sedlacek) (1 cf , 2 $ BMNH); 3 $ , Malgi, Mt Giluwe, 2400 m, 25-30.V.1961, light-trap, heavy rain (/. L. &
M. Gressitt); 3 cf , 6 $, Dimifa, SE. Mt Giluwe, 2200 m, 9-12.X.1958, light-trap (1 cf , 2 $ BMNH); 3 cf ,
10 9, ridge W. of Dimifa, S. Mt Giluwe, 2350m, ll.x.1958; 1 cf , SE. slope Mt Giluwe, 2450m, 12.x. 1958;
4 Cf, 4 9, Mt Otto, 2200 m, 24.vi.1955 (all J. L. Gressitt) (1 cf, 1 $ BMNH); 1 cf, Yaibos 1650 m,
10-ll.vi.1963 (/. Sedlacek); 1 $, Ahl Valley, Nondugle, 1750 m, 8.vii.l955 (J. L. Gressitt); 1 $, Korgua,
1450 m, 30.V.1963 (H.C.); 1 cf , Daulo Pass, 2400 m (Asaro Chimbu Div.) 14.vi.1955, light-trap (/. L.
Gressitt); II $, Tambul, 2200 m, 26.v.-7.vi.l963, light-trap (J. Sedlacek) (5 $ BMNH); 1 cf, 25,600,
Moke, 6400 ft, m.v. light, 17.iv.1962; 1 cf , 1 $, 19,025, 19,026, Aiyura, 1959 (J. H. Barrett) (DPI); 1 $,
Wau Creek, Wau, 1200-1500 m, 16-18.ix.1964 (M. Sedlacek); 1 cf, Mt Michael, Lufa, 10.iii.1974 (R.
GALERUCINE BEETLES OF NEW GUINEA 217
Hornabrook); 1 $ Samedata, 17.iii.1974, at light; 1 $, W. Highlands, Kandep, 25.iii.1971 (R. Hornab-
rook) (all specimens BPBM unless otherwise stated).
COMMENTS. This species appears to be fairly common throughout the central region of NE.
Papua New Guinea. Apart from colour variation there appears to be a certain amount of
inter-population variation. It is possible that more than one species may have been included
here, although no specific differences could be found in the genitalia of these forms.
MARMINA gen. n.
Type-species: Sastra quadripustulata Jacoby.
GENERAL FORM. Body elongate, elytra subparallel-sided to subovate. Dorsum appearing glabrous. Length
of antenna just exceeding 0-5 x that of body, third segment at least l-3x length of fourth segment and l-8x
length of second. Eyes not protuberant. Pronotum transverse, lateral margins with a small rounded tooth
just above midline, width at anterior margin equal to or just exceeding that of head plus eyes, maximum
width not greater than that of elytra at humeri. Primary setal pores distinctly tuberculate; pronotal
depressions occupying at least two-thirds of surface area, sublateral depression transverse, often confluent
with ovate depression on disc. Elytra unicolorous, bicoloured or patterned. Epipleura present to apical
quarter, broad in basal half then narrowing towards apices from around level of hind coxae onwards.
Elytron sometimes explanate laterally beyond epipleuron. Underside setose. Fore coxal cavities open
behind. Apex of abdomen in cf with a shallow even emargination (Fig. 18), 9 with a short longitudinal
bifurcation (Fig. 19). Legs moderately robust, similar in both sexes, apex of tibiae without ventral spur,
lateral apices with a comb-like row of short spines. Length of basal segment of fore and mid tarsi more or
less equal to combined length of the two preceding segments; basal segment of hind tarsus l-2x as long as
preceding segments combined. Claws distinctly bifid. Basal segment of fore tarsus in cf only slightly more
enlarged than basal segment of mid tarsus.
DIAGNOSIS. Head with gena weakly emarginate medially, minimum length of gena not less than 0-5 x
length of antennal socket. Apical segment of maxillary palp subconical, length slightly greater than
preceding segment. Labrum small, not obscuring sides of mandibles. Pseudoclypeus distinct, length
slightly greater than half that of labrum. Frons medially convex and somewhat declivous anteriorly, length
not greater than that of an antennal socket. Antennal sockets separated by not more than half width of a
socket, anterior margins level with those of eyes but not contiguous. Postantennal swellings transverse,
delimited behind by a narrow horizontal groove and separated medially by a longitudinal extention of the
coronal 'suture', anterior margins of swellings confluent with frons between antennal sockets and of similar
elevation. Vertex with puncturation sparse or almost absent. Antennae similar in both sexes (Figs 9, 10),
segments filiform, basal segment l-25x length of second, segment 3 at least 2-Ox length of second and l-3x
fourth, fifth segment slightly shorter than fourth, segments 6, 7 and 8 subequal and slightly shorter than
fifth, ninth and tenth segments equal, slightly shorter than eighth, segment 11 l-2x length of tenth, all
segments setose, density of setae increasing from segment 4 onwards. Scutellum on average l-2x longer
than broad, apex subtruncate, often medially convex. Elytron on average 2-5-3-Ox longer than broad,
average maximum width l-2x width at apex of scutellum. Post humeral sublateral longitudinal depression
almost absent or weakly indicated by raised outer margin, surface without carinae or rugosities. Underside
minutely punctured throughout. Metasternum transverse, length at least 2x that of mesosternum,
medially convex. Basic structure of $ genitalia, bursa, position and structure of styli etc. as in (Fig. 38).
Median lobe of aedeagus in c? forming an elongate chitinous tube (Figs 25-31). General structure as in
Polysastra (Fig. 32).
DISTRIBUTION. Papua New Guinea, Irian Jaya, Trobriand Islands, D'Entrecasteaux Islands
(Map 1).
DISCUSSION. This genus is superficially similar to Paumomua; it differs principally in having a
tooth on the lateral margin of the pronotum, antennal segments of different proportions, e.g.
third segment longer than fourth, and a longer apical segment of the maxillary palp. Species with
subparallel-sided elytra may also appear similar to certain Momaea species, but again can be
distinguished from these by the structure of the pronotum and the distance between the anterior
margin of the frons and the eyes. Nothing is known about the biology of this genus.
218
SHARON L. SHUTE
Figs 16-31 Polysastra, Dreeus and Marmina, taxonomic characters. 16-21, apical segment of abdomen
showing emargination of (16) Polysastra sp. cf; (17) Polysastra sp. $; (18) Marmina sp. cf; (19)
Marmina sp. $; (20) Dreeus sp. cf ; (21) Dreeus sp. $. 22, 23, Dreeus distincta cf , aedeagus, dorsal and
lateral view. 24, D. distincta 9 , spermatheca. 25, 26, Marmina quadripustulata cf , aedeagus, dorsal and
lateral view. 27, M. quadripustulata $, spermatheca. 28, 29, Marmina sp. B cf, aedeagus, dorsal and
lateral view. 30, 31, Marmina sp. A cf, aedeagus, dorsal and lateral view. (Figs 25-31 drawn to same
scale.)
GALERUCINE BEETLES OF NEW GUINEA
219
Map 1 Range of genera treated here.
Key to species of Marmina
1 Elytron with colour pattern, puncturation distinct but not strong, punctures on humeral area
1-2 x their own width apart 2
- Elytron unicolorous or with humerus similar in colour to pronotum, puncturation fine, punc-
tures minute , 3-4 x their own width apart 3
2 Elytra subparallel-sided , deep submetallic green with irregular transverse median and subapical
yellow patches, these not contiguous with lateral margin. Head, pronotum and scutellum dull
testaceous to golden yellow. Vertex posterior to post antennal swellings usually dark reddish
brown. Tibiae and tarsi deep reddish brown to pitchy black. Length 8-5-11-5 mm (Fig. 116).
Genitalia as in Figs 25-27. (Trobriand Islands, D'Entrecasteaux Islands) quadripustulata (p. 220)
- Elytra subovate, lateral margin explanate beyond epipleuron, colour testaceous with irregular
dark brown markings (Fig. 118). Pronotum testaceous with ill-defined reddish brown patches
on disc, lateral tooth and sublateral posterior convex areas. Head with frons, post antennal
swellings and vertex dark reddish brown, intermediate areas testaceous. Underside tes-
taceous. Basal three-quarters of femora and basal quarter of tibiae testaceous, rest of legs
dark reddish to pitchy brown. Antennal segments with base of each segment testaceous.
Length d" 9-0 mm. (Fig. 1 18.) Aedeagus as in Figs 30, 31. (NE. Papua New Guinea) species A (p. 220)
3 Elytron deep bluish to pitchy brown, reddish along margins, humerus light orange-brown.
Head, pronotum and scutellum dull testaceous. Metasternum and abdomen pitchy brown to
black, rest of underside testaceous. Tibiae usually darker in colour than femora. Antennal
segments 1-5 dull orange-brown, rest of segments becoming darker towards apices. Length
9- 1-9-3 mm. (Fig. 1 17. )(SE. Papua New Guinea) basalis(p. 220)
220 SHARON L. SHUTE
Elytron deep purplish black with a weak brassy lustre on humeral area, lateral margins and
epipleuron testaceous. Head, pronotum and scutellum dull testaceous. Coloration of under-
side as in basalis. Legs testaceous. Antennal segments 1-4 testaceous, rest dark reddish
brown. Length c? 7-5 mm. (Fig. 1 19.) Aedeagus as in Figs 28, 29. (SE. Irian Jaya) . . . species B (p. 220)
Marmina quadripustulata (Jacoby) comb. n.
(Figs 25-27, Map 10)
Sastra quadripustulata Jacoby, 1904: 402. LECTOTYPE $, TROBRIAND ISLANDS (BMNH), here desig-
nated [examined].
ADDITIONAL MATERIAL EXAMINED
Trobriand Islands: 1 cf (paralectotype) (MCZ). D'Entrecasteaux Islands: 1 cf , 1 $ (paralectotypes),
Fergusson Island (BMNH); 2 $, Goodenenough Island, xii. [18]96 (A. S. Meek); 3 $, Fergusson Island,
ix-xii. [18]95 (A. S. Meek) (MCZ).
Marmina basalis (Jacoby) comb. n.
(Fig. 117, Map 10)
Sastra basalis Jacoby, 1886: 72. LECTOTYPE cf , NEW GUINEA: Fly River (L. N. D'Albertis) 1876-77
(MCZ), here designated [examined].
ADDITIONAL MATERIAL EXAMINED
New Guinea: 1 $ (Wallace) (BMNH).
Two further species belonging to this genus are included in the key but are not described as they
are represented by only single specimens. Collecting data of these species are as follows.
Marmina sp. A
(Figs 30, 31, 118, Map 10)
Papua New Guinea: 1 C", Morobe District, Mt Kaindi nr Wau, montane forest, c. 8000 ft, 16.iv.1965
(J. J. H. Szent-Ivany) no. 19031 (DPI).
Marmina sp. B
(Figs 28, 29, 119, Map 10)
Irian Jaya: 1 cf , Sabang, 12.vii.1907 (Lorentz) (ITZ).
POLYSASTRAgen.n.
Type-species: Sastra costatipennis Jacoby.
GENERAL FORM. Body elongate, elytra subparallel-sided to pyriform. Dorsum either distinctly setose or
with setae confined within margins of punctures. Combined length of antennal segments equal to at least
three-quarters of body length, third segment at least l-4x length of fourth segment and l-8x second,
segments filiform in both sexes (Figs 7, 8). Eyes large and protuberant. Pronotum transverse, lateral
margins produced into a distinct tooth in anterior half, apex acute or broadly rounded, anterior width not
exceeding that of head plus eyes, maximum width at least l-3x greater than that of head plus eyes but not
exceeding that of elytra at humeri; sublateral depressions present either side of disc, disc with small
anterior and posterior depressions which may be confluent in some species. Primary setal pores tubercu-
late. Elytra variable, evenly convex, costate or rugose, with uniform or irregular puncturation, derm
translucent to metallic. Epipleuron present to apical quarter, a supra-costal flange present in some species.
Underside setose. Fore coxal cavities open behind. Apex of abdomen in male with a deep triangular
emargination, evenly rounded in female (Figs 16, 17). Legs slender, similar in both sexes, apices of tibiae
without ventral spurs, lateral edges with a comb-like row of short spines. Basal segment of fore and mid
tarsi slightly shorter than segments two and three combined, basal segment of hind tarsus slightly longer
than segments two and three combined. Claws distinctly bifid. Basal segment of fore tarsus enlarged in
male.
GALERUCINE BEETLES OF NEW GUINEA 221
DIAGNOSIS. Head with gena emarginate medially, emargination almost reaching anterior margin of eye
(Fig. 14) . Apical segment of maxillary palp conical , length more or less equal to that of preceding segment.
Labrum small, not obscuring sides of mandibles. Pseudoclypeus distinct, length not more than half that of
labrum . Frons either medially convex or entirely depressed, length not greater than half that of an antennal
socket, sockets separated by less than width of socket, anterior margins level with those of eyes.
Post-antennal swellings elevated above level of vertex, contiguous with frons between sockets and
separated medially by a narrow longitudinal groove which terminates at frons. Vertex with or without
irregular puncturation, coronal 'suture' usually distinct. Eyes large and prominent, at least 3-Ox longer
than antennal socket. Antenna with basal segment becoming gradually broader towards apex, length at
least twice that of second segment, third segment 2-5 x length of second and 1 -5-2-0 x that of fourth;
fourth, fifth, sixth and seventh segments subequal, eighth segment slightly shorter than seventh, segments
eight, nine and ten subequal, segment eleven l-5x length of tenth segment, all segments setose. Pronotum
transverse, degree of convexity and development of depressions variable, lateral tooth well developed with
apex acute or weakly developed and broadly rounded, lateral margins always produced, never evenly
rounded, type and density of puncturation variable. Scutellum triangular, length either equal to or slightly
greater than width, apex rounded or subtruncate. Average length of elytron 3-3 x longer than broad, facies
variable (see notes on species groups). Underside finely punctured throughout, metasternum convex, at
least 3-Ox as long as mesosternum. Legs of similar form in both sexes, slender, length of hind leg equal to or
slightly longer than that of elytron, length of fore and mid legs slightly less than that of hind leg, all femora
more or less equal in width, tibiae more or less straight.
Aedeagus a narrow, dorsally curved, chitinous tube, the apical section dorsally flattened. Viewed
dorsally (Fig. 32) it occupies the length of the abdomen and lies laterally, slightly to the right of the gut. The
median lobe and the basal struts are fused, forming a single unit. The median orifice is large and opens on
the dorsal side of the apical section. The apex usually has some form of projection, ranging from a blunt
knob to a large tooth. The aedeagus lies within a membranous bag, the genital atrium, to which the arms of
the tegman (Sharp & Muir, 1912) or spiculum (Varmer, 1969) are attached at the apical quarter (Fig. 32),
the remaining strut of the spiculum remaining free of the atrium wall until it enters the basal orifice where it
curves upwards between the curved basal struts. The non-eversible portion of the endophallus, which is
continuous with the median ejaculatory duct at the base of the aedeagus, appears as a dark tube within the
median section of the median lobe; it becomes confluent with the internal sac in the apical quarter which
appears as a dark amorphous shape and can be seen protruding at the base of the median orifice together
with the projecting apex (which varies in shape) of the supporting sclerite , which lies in an inverted S-shape
inside the sac at rest. On evagination the sclerite runs through the centre of the sac and through the
phallotreme where it fuses with the outer wall of the sac, acting as a supporting strut (Fig. 33). The
evaginated sac is small and simple in shape but the walls are fairly robust, being covered with setae and
spicules (Fig. 33). The sac has proved very difficult to evaginate in this genus, principally due to the narrow
structure of the aedeagus and the rigidity of the supporting sclerite which lies in an inverted S-shape at
rest and has to be 'popped' straight. Therefore the sac has not been used here as a specific diagnostic
character.
The female genitalia (of which the spermatheca is used here as a specific diagnostic character) are
enclosed in a membranous tube, the apices of which are the chitinous plates of the eighth tergite and
sternite (hemitergites); long stout setae are present along the anterior margin of both plates; with the
ventrites removed the gut can be seen lying in front of the genitalia (Fig. 35). The anterior margin of the
entrance to the gut forms a shallow membranous fold which is attached to the posterior region of the eighth
tergite, the hind region of the gut entrance forming a deep blind fold, the hind margin of which is attached
to the anterior margins of the base of the styli (Fig. 36). The styli comprise two elongate hollow tubes
which, at the base, open into the region between the fold formed by the hind margin of the gut and the
anterior wall of the vagina (Fig. 36). There are no distinct coxities or supporting valvifers. Each stylus has a
tuft of long, curved, sensory setae at the apices. The hind margins of the base of the styli are attached to the
anterior margin of the wall of the vagina; the entrance to the vagina (vulva) lies between the styli and the
ligular (Fig. 36). The vagina and the bursa copulatrix (Fig. 35) form a convoluted membranous sack; there
is no distinct elongated constriction between the vagina and the bursa as in Dreeus (Fig. 34). The
spermatheca (Fig. 35), which is a strongly chitinous curved structure, is attached to the dorsal side of the
anterior end of the bursa by a short membranous duct (Fig. 35). The spermathecal gland may be seen
attached to the posterior region of the spermatheca above the attachment of the spermathecal duct
(Fig. 35). The oviduct arises from the median region of the ventral wall of the bursa. The principal
interspecific diagnostic characters are the shape of the spermatheca and ligular, length of styli and the
density and position of the setae on the hemitergites. The general structure, i.e. , proportions of the vagina,
bursa and plates etc. are generic characters (Figs 34-38).
222 SHARON L. SHUTE
DISTRIBUTION. Papua New Guinea, Irian Jaya, New Britain, Trobriand Islands, D'Entrecas-
teaux Islands, N. Australia (Map 1).
DISCUSSION. Polysastra is closely allied to Sastra. Both genera belong to a group of primarily
Australasian Galerucini which is characterised by an elongate form (Figs 1-3), emarginate
genae (Fig. 14) and the third antennomere being at least l-3x longer than the fourth. Polysastra
differs from Sastra (Fig. 1) in having a distinct tooth on the lateral margin of the pronotum
(Fig. 2) less emarginate genae, and variable elytral facies ranging from evenly convex to rugose
or carinate; the dorsal puncturation and setal length are also very variable. The two genera have
similar geographic ranges (Map 1).
Very little is known about the biology of Polysastra. Species occur in a wide range of habitats
throughout New Guinea, from high moss forest to lowland marshy areas. Various species have
been found on cultivated plants in gardens and plantations but there is no precise record of
feeding. No species have as yet been found to be of any significant economic importance. The
majority have been collected at various forms of light, which suggests that they may be most
active at night.
In comparison with most other genera of Galerucinae the species of Polysastra exhibit an
unusually wide range of external variation. A study of those treated here and c. 40 undescribed
species showed that certain groupings, based on external characters, were evident within the
genus. Eleven such species groups have been defined here, to facilitate the preliminary
identification of members of this genus. Further groupings based on additional characters may
be apparent within these species groups and additional notes have been provided on each group
to supplement the information derived from the primary group key. At present it is not feasible
to describe all the new species examined, but at least one new species from each 'group' is
described here as a representative example.
At first sight the external characters exhibited by these species groups may suggest that they
represent monophyletic groups. However, lack of geographic correlation between the species of
a group, the occurrence of intermediate species, and the pattern of variation and overlap in
genitalic characters lends little to support this supposition. The occurrence of similar aedeagal
forms in externally dissimilar groups which show relative constancy of spermathecal structure
suggests that genitalic characters may prove more useful in elucidating phyletic relationships of
Polysastra species than similarities in external form. It is hoped that this preliminary study will
provide a helpful basis for future work on this genus.
Groups and species are dealt with in alphabetical order. Type-data etc. of those species
transferred from Sastra are included with the descriptions of the new taxa. The general form and
genitalia of all the species dealt with in this genus are illustrated.
Key to species groups based on external characters
Note. The term pubescent is used here in a specific context to describe a fine, dense covering of setae.
1 Elytron appearing distinctly setose or pubescent, setae extending well beyond margins of
punctures 2
Elytron appearing glabrous, majority of setae minute and confined within the margins of
the punctures.
Sometimes with sparse, erect setae at intervals 4
2 (1) Pronotum very densely punctured throughout, punctures fine to granulate; setae dense or
forming pubescence. Elytron non-metallic, setae dense or forming pubescence.
C. 8-0-11-Omm 3
Pronotum with irregular puncturation, punctures large and mostly shallow, setae sparse
but distinct. Elytron metallic or with a metallic lustre, one or both margins of the
post-humeral sublateral longitudinal depression forming cariniform ridges; setae dis-
tinct but not forming pubescence. C. 8-0-11-Omm bicostata- group (p. 226)
3 (2) Elytron with outer margin of post-humeral sublateral longitudinal depression forming a
distinct ridge, lateral margin somewhat explanate beyond epipleuron in median third;
elytral puncturation fine to granular, punctures usually contiguous, derm usually
32
Figs 32-37 Polysastra and Dreeus, genitalia. 32, Polysastra sp. , diagram of abdomen showing position of
aedeagus at rest. 33, Polysastra sp., apex of aedeagus showing internal sac evaginated. 34, Dreeus sp.,
general structure of female genitalia. 35, Polysastra sp., general structure of female genitalia. 36,
Polysastra sp. , position of styli and ligular with eight tergite removed. 37, Dreeus sp. , position of styli and
ligular with eighth tergite removed.
aw - anterior wall of vagina; be - bursa copulatrix; bo - basal orifice; bs - basal struts; e - endophallus; et
- eighth tergite; ev - entrance to vagina; g - gut; ga - genital atrium wall; ht - hemitergites; is - internal
sac; iss - internal supporting sclerite; lig - ligular (eighth sternite); med - median ejaculatory duct; ml -
median lobe (aedeagus); mo - median orifice; o - oviduct; p - phallotreme; s - spermatheca; sd -
spermathecal duct ; sg - spermathecal gland ; sp - spiculum (tegmen) ; st - styli ; v - ventrites ; vag - vagina .
224
SHARON L. SHUTE
39
41
pas
Figs 38, 39 Dreeus sp. , female genitalia. 38, general structure. 39, position of styli and ligular with eighth
tergite removed.
be - bursa copulatrix ; et - eighth tergite ; g - gut ; ligular (eighth sternite) ; o - oviduct ; s - spermatheca ; sd
- spermathecal duct; sg - spermathecal gland; st - styli; vag - vagina.
Fig. 40 General form of genus B.
Fig. 41 Polysastra sp. (costatipennis-group) illustrating general taxonomic characters.
ac - ad-lateral carina ; me - carinae formed by margins of depression ; pas - post-antennal swellings ; pd -
pronotal depressions; sd - post-humeral sublateral depression.
GALERUCINE BEETLES OF NEW GUINEA 225
microsculptured. Pronotum moderately convex, depressions shallow but distinct.
C. 8-0-1 1-0 mm exp/anafa-group (p. 226)
Elytron with post-humeral sublateral longitudinal depression weak to absent, margins
never forming ridges, lateral margin not explanate; elytron finely punctured, punctures
either contiguous or not more than l-5x their own width apart, majority of species with
distinct micropuncturation on intervals.
Underside may have a supra-costal flange running parallel to epipleuron, or a
preapical pit or depression which appears as a convexity on the dorsal surface.
C. 7-5-14-0 mm micropunctata-group (p. 228)
4 ( 1 ) Elytron either irregularly convex with irregular puncturation or with the punctures situated
around irregular non-punctate areas, which usually form distinct convexities or rugosi-
ties.
Outer margins of post-humeral sublateral longitudinal depression may or may not
form carinae /rregu7ar/s-group (part) (p. 227)
Elytron more or less uniformly punctured throughout, puncturation fine to coarse, no
rugosities or irregularities present 5
5 (4) Elytron with one or both margins of post-humeral sublateral longitudinal depression
forming cariniform ridges, admedian carinae sometimes present 6
Elytron with post-humeral sublateral longitudinal depression weak to absent, margins
never forming distinct ridges or carinae 9
6 (5) Frons entirely depressed to concave.
Elytra often with a metallic lustre. C. 8-0-13-0 mm var/a-group (p. 229)
Frons triangular, medially convex or weakly to moderately convex, sometimes with a weak
median depression, sides always elevated to some degree 7
7 (6) Elytron with both margins of post-humeral sublateral longitudinal depression forming
cariniform ridges which are present from base of humeri to apical quarter, one or more
entire or partial admedian carinae sometimes present 8
Elytron with outer margin of post-humeral sublateral longitudinal depression forming a
ridge, inner margin sometimes partially developed.
Elytron broadly rounded to pyriform; metallic or with a metallic or weakly metallic
lustre; some species may have a slight crumpled appearance but puncturation more or
less evenly distributed throughout. C. 10-5-14-0 mm mefa7//ca-group (p. 227)
8 (7) Frons weakly elevated or with a shallow median depression. Elytron somewhat unevenly
convex, sometimes weakly rugose, puncturation dense throughout, punctures may form
horizontal or oblique rows of c. 5-7 punctures, derm usually microsculptured, humeral
area often with one or more partial carinae, derm non-metallic. Vertex and pronotum
densely and coarsely punctured throughout. C. 9-0-15-0 mm irregularis-group (part) (p. 227)
Frons triangular, medially convex. Elytron more or less evenly convex, puncturation more
or less uniformly distributed throughout, admedian carina sometimes present, running
parallel to sublateral carinae, derm often with a distinct metallic or submetallic lustre.
Vertex and pronotum weakly and indistinctly to moderately punctured, punctures
usually large and shallow. C. 8-0-11-0 mm costatipennis-group (p. 226)
9 (5) Elytron broadly rounded, sublateral post-humeral longitudinal depression almost absent;
very strongly microsculptured, giving a dull appearance, punctures fine, tending to form
irregular groups. C. 8-0-10-0 mm irregularis-group (part) (p. 227)
Elytron not as above 10
10 (9) Pronotum broad and weakly convex, puncturation coarse and dense throughout, lateral
tooth weakly developed and rounded, median depression joined by a shallow longitu-
dinal narrow groove. Elytron strongly and densely punctured throughout, derm often
with an aeneous lustre, post-humeral sublateral depression shallow. C. 10-0-13-0 mm
abdominalis-group (p. 225)
Pronotum not as above, tending to be elongated, lateral tooth rounded to subacute,
punctures irregular, size fairly large and shallow, sometimes confluent in places. Elytra
irregularly punctured, derm smooth and shining. C. 9-0-12-0 mm inhabilis-group (p. 226)
The abdominalis-group
Characterised principally by a broad, very weakly convex, strongly and densely punctured pronotum,
which has a broad, rounded, weakly developed lateral tooth. Vertex coarsely and densely punctured
226 SHARON L. SHUTE
throughout, size of punctures as on pronotum. Frons forming a more or less evenly convex ridge. Elytron
with a shallow post-humeral sublateral longitudinal depression. Puncturation strong and dense through-
out, giving a slightly rugose appearance, punctures distinctly smaller than those on pronotum, not more
than l-3x their own width apart, derm smooth and shining, often with a weak submetallic lustre; setae
minute, confined within margins of punctures, sometimes a few short, erect setae on intervals. Sperma-
thecal form distinctive, differing from that in other groups by being narrow and distinctly elongated (Fig.
83). None of the aedeagal forms found in other groups has been found in this group. General colour dark
green to brown, often with lighter coloured lateral margins to the elytron.
COMMENTS. The species representing this group and dealt with here is abdominalis (Figs 144,
145). A further six undescribed species that fall into this group have been examined.
The fr/cosfafa-group
Resembling the obscuricornis-group but distinguished by the distinctly setose elytra, finer puncturation
and weakly developed margins of the post-humeral, sublateral longitudinal depression. Admedian carinae
sometimes present.
COMMENTS. The species representing this group and dealt with here is bicostata (Fig. 149). A
further two undescribed species have been examined.
The costatipennis-group
Characterised principally by having both margins of the post-humeral sublateral depression forming
distinct cariniform ridges from the base of the humeri to the apical quarter, one or more additional
admedian carinae sometimes present. Derm metallic or with a submetallic lustre. Puncturation fairly
uniform throughout, fine to moderate, punctures not more than their own width apart, setae minute and
confined within the margins of the punctures. Vertex and pronotum with fairly large, shallow, irregular
punctures which may be confluent in places, derm usually smooth and shining. Pronotum weakly to
moderately convex, depressions shallow, lateral tooth broad and weakly developed. Frons distinctly
convex medially.
COMMENTS. Representative species dealt with here are costatipennis, fuscitarsis, helleri, kam-
peni, laetabilis, obscuricornis, purpurasco (Figs 120-126). A further four undescribed species
belonging to this group have been examined.
The exp/aiiafa-group
Distinctly setose dorsally, with all setae extending beyond margins of punctures. Vertex and pronotum
with fine to granulate, dense puncturation throughout. Pronotum with broad, weakly developed lateral
tooth. Elytron pubescent and very finely punctured throughout, punctures not more than half their own
width apart, derm usually distinctly microsculptured. Outer margin of post-humeral sublateral longitu-
dinal depression forming a distinct ridge, lateral margins somewhat explanate beyond the epipleura (more
pronounced in female). Frons convex, elevation more or less equal to that of post-antennal swellings.
Elytral coloration dull testaceous to dark reddish brown. Species with these external characteristics exhibit
a similar type of aedeagus which predominates in the costatipennis-group.
COMMENTS. The species representing this group and dealt with here is explanata (Fig. 128). A
further six undescribed species that fall into this group have been examined.
The inhabilis-group
Slender in form with distinct, irregular dorsal puncturation. Punctures on head and pronotum slightly
larger than those on elytra, usually not more than twice their own width apart. Pronotum elongate, lateral
tooth broad and weakly developed, derm smooth and shining, setae minute. Elytron with post-humeral
sublateral longitudinal depression almost absent, outer margin sometimes forming a weakly developed
ridge, particularly over humeri; punctures irregularly distributed, l-0-l-5x their own width apart, derm
smooth and shining, setae confined within margin of punctures. Testaceous to light reddish brown.
COMMENTS. The species dealt with here and representing this group is inhabilis (Fig. 143). A
further five undescribed species have been examined.
GALERUCINE BEETLES OF NEW GUINEA 227
The irregularis-group
Elytron strongly microsculptured, or uneven or rugose. Punctures in majority of species situated between
irregular non-punctate areas which take the form of irregular, sinuate, elongate, confluent rugosities which
may be smooth and shining or distinctly microsculptured. Setae confined within margins of punctures but
some species, i.e. confusa, have long suberect setae on the intervals. Vertex and pronotum distinctly
punctured, pronotal setae usually confined within margins of punctures.
COMMENTS. The majority of species that fall into this group may be further divided into
subgroups by the additional characters detailed below. Examples of the irregularis-group dealt
with here are confusa (Fig. 138), duplicator (Fig. 135), irregularis (Fig. 136), sedlaceki
(Fig. 137), rugulosa (Fig. 140). A further 35 undescribed species of this group have been
examined.
Subgroup 1
Elytron appearing distinctly rugose, non-punctate areas smooth and shining, derm usually with a distinct
submetallic lustre, weakly developed partial carinae may be present on humeral area. Punctures situated in
small groups of 2-5 between elevated areas which tend to be sinuate and confluent in places. Vertex and
pronotum moderately punctured, pronotum convex with a well-developed lateral tooth which tends to be
rounded, area around posterior primary setal pore often produced into a blunt 'tooth'. Frons tends to be
very weakly convex or with a median depression and sinuate anterior margin. The species tend to range
from reddish to bluish purple or bluish green or a combination of both, derm usually distinctly sub-metallic
(Fig. 140).
Subgroup 2
Elytron with dense, moderate to coarse puncturation throughout, interspersed with mostly isolated, small,
irregular, distinctly convex areas which tend to be slightly darker in colour than surrounding area. Irregular
rows of long erect setae usually present, rest of setae minute. Vertex and pronotum coarsely and densely
punctured throughout. Pronotum convex with a well-developed lateral tooth, usually with a subacute
apex. Frons tending to form a more or less evenly elevated convex ridge. The species tend to be testaceous
to reddish brown with an aeneous lustre (Figs 138, 139).
Subgroup 3
Elytron with weak to moderately developed, elongate, sinuate, confluent non-punctate areas which are
moderately to strongly microsculptured, majority of punctures situated in groups between non-punctate
areas, rest irregularly distributed. Vertex and pronotum with moderately dense, large, shallow well-
defined punctures, derm usually microsculptured, pronotum usually weakly convex, lateral tooth well
developed, often with an acute apex. Frons moderately convex to somewhat flattened dorsally. The species
tend to be dull and darkly coloured, non-metallic (Figs 135, 136, 137).
Subgroup 4
-
Elytron unevenly convex, often with rugosities, puncturation irregular, derm often distinctly microsculp-
tured. Margins of post-humeral, sublateral longitudinal depression forming cariniform ridges, one or more
partial carinae may also be present on humeral area. Vertex and pronotum distinctly punctured, punctures
sometimes coarse and confluent in some species (Figs 141, 150).
Subgroup 5
Elytron more or less evenly convex, derm very strongly microsculptured, puncturation fine, usually with a
weak indication of grouping. Vertex and pronotum with distinct, large, shallow, usually non-confluent
punctures. Usually small, black to dark green species with broadly rounded elytra (Fig. 142).
The metallica-group
Elytron distinctly metallic or with a metallic lustre; broadly rounded to pyriform, outer margin of
post-humeral sublateral longitudinal depression forming a distinct ridge, inner margin sometimes partially
developed. Dorsal setae minute, confined within margins of punctures, giving a glabrous appearance.
228 SHARON L. SHUTE
Pronotum with lateral tooth broad, weakly developed. Frons medially convex. The group can be further
divided into three subgroups based on the additional characters detailed below.
COMMENTS. The species representing this group and dealt with here are metallica, suavis. A
further seven undescribed species have been examined.
Subgroup 1
Elytron distinctly metallic with fine to moderate puncturation throughout, surface sometimes slightly
uneven, giving a weakly crumpled appearance, outer margin of post-humeral sublateral longitudinal
depression forming an irregular, weakly sinuate ridge, inner margin sometimes partially developed.
Vertex and pronotum smooth and shining with fairly large, shallow, sparse to moderately dense
puncturation. Elytron deep metallic green/blue or dark purplish blue (Fig. 132).
Subgroup 2
Elytron submetallic or with a strong metallic lustre, puncturation moderate to coarse throughout, giving a
slight rugose appearance in some species, ridge formed by outer margin of sublateral depression sinuate.
Vertex and pronotum with large coarse punctures which may be confluent in places, pronotum narrow,
depressions distinct, lateral tooth sometimes slightly more acute than in group 1. Elytron dark brownish
green to purplish brown with a dark green or purplish lustre (Fig. 133).
Subgroup 3
Elytron with a metallic lustre, puncturation strong and dense throughout, surface more or less even
throughout. Pronotum broad, weakly convex; outer margin of post-humeral sublateral depression forming
a well-developed, more or less straight cariniform ridge. Elytron dark brownish green with an aeneous
lustre or bright green/blue (Fig. 134).
The micropunctota-group
Characterised by the distinct, fine, dense elytral setae that extend beyond margins of punctures and the
presence of micropuncturation on the intervals between the regular elytral puncturation (subgroup 3).
Post-humeral sublateral depressions absent, elytron evenly convex, puncturation fine and more or less
uniform throughout . Vertex and pronotum densely punctured throughout , sometimes granulate , setae fine
and distinct. Pronotum with a well-developed lateral tooth, usually with acute apices. Some species may
have a distinct supracostal flange running parallel to the epipleura on the underside of the elytron, e.g.
micropunctatus (Fig. 130), or a preapical pit-like depression; both may appear as a ridge or convexity on
the upper surface of the apical quarter. This group divides into three principal subgroups based on the
additional characters detailed below.
COMMENTS. The species representing this group and dealt with here are montana (Fig. 129) and
micropunctata. A further nine undescribed species have been examined.
Subgroup 1
Vertex and pronotum with fine to granulate puncturation, setae long, fine and appressed. Pronotum often
with two small admedian and adlateral convexities, lateral tooth well developed, apices subacute to acute.
Elytron with very fine regular puncturation, punctures 0-5-1 -5 x their own width apart, intervals with
distinct micropunctures from which the majority of the setae arise, setae short, fine and appressed,
extending well beyond margins of punctures. Sometimes a supracostal flange or preapical depression on
underside of elytron. Dull, black to reddish brown species.
Subgroup 2
Vertex and pronotum irregularly and densely punctured throughout, puncturation confluent, never fine
and granulate as above. Setae long and appressed. Pronotum somewhat flattened and elongated, lateral
tooth large and acute, wholly in anterior half, lateral margin between base of tooth and posterior primary
setal pore more or less straight, derm with a slight vitreous lustre, setae long and appressed. Elytron with
slightly larger and less dense puncturation than in subgroup 1, micropuncturation slightly less distinct due
to surface reflection, setae very fine and appressed. The species tend to be elongate and slender with almost
parallel-sided elytra, mostly pale testaceous to light reddish brown.
GALERUCINE BEETLES OF NEW GUINEA 229
Subgroup 3
Vertex and pronotum as in subgroup 1, but elytron with fine contiguous puncturation throughout,
micropuncturation almost absent due to density of regular punctures.
The var/a-group
Very similar in appearance to the obscuricornis-group, but separated by the following characters. Frons
entirely depressed, weakly concave in some species. Carinae formed by margins of post-humeral,
sublateral longitudinal depression never joined at base of humeri, sometimes linked by an irregular,
oblique convexity at around level of hind coxae, this convexity may also link ad-median carina to adjacent
carina. Colour range variable as in the costatipennis-group.
COMMENTS. The species representing this group and dealt with here are varia (Figs 146-148) and
venusta (Fig. 127). A further three undescribed species have been examined.
Key to described species and subspecies of Polysastra
[Further undescribed species are referred to in the text.]
1 Elytron distinctly pubescent throughout, all setae extending well beyond diameter of
punctures . Puncturation more or less uniform throughout , derm never rugose 2
Elytron not pubescent, majority of setae minute, confined within margins of punctures,
sparse erect setae sometimes present on intervals, puncturation irregular or uniform,
derm rugose or costate 5
2 (1) Pronotal puncturation fine to granulate throughout, setae distinct. Elytron evenly
convex , post-humeral sublateral longitudinal depression indistinct or absent 4
Pronotal puncturation not as above. Elytron with a distinct post-humeral sublateral
longitudinal depression , the outer margin of which forms a distinct ridge 3
3 (2) Elytron pale reddish brown with an overall light metallic green lustre. Head, pronotum
and scutellum testaceous to light orange-yellow. Underside and legs testaceous.
Vertex and pronotum irregularly punctured with large, shallow, mostly non-confluent
punctures. Pronotal depressions well defined. Lateral margin of elytron not explanate
beyond epipleura, setae not dense. Length 9-0-11-0 mm. (Fig. 149.) Genitalia as in
Fig. 62. (Map 2.) bicostata(p. 232)
Elytron dull orange to deep reddish brown, pronotum normally slightly lighter in colour.
Head either concolorous with pronotum or dark as in elytron. Underside and legs
testaceous to light orange-brown. Vertex, pronotum and scutellum with dense con-
fluent punctures throughout. Lateral margin of elytron somewhat explanate beyond
epipleura, setae dense. Length 9-5-12-0 mm. (Fig. 128.) Genitalia as in Figs 51, 84.
(Map 3.)
explanata (p. 233)
4 (2) Underside of elytron with a distinct supracostal flange running parallel to epipleuron
from base to apical quarter where it joins sutural margin above apex. Dorsum dark
reddish to pitchy brown, lateral margin of elytron reddish orange, distinctly lighter
than rest of elytron. Length 10-5-11-5 mm. (Fig. 130.) Genitalia as in Figs 52, 85. (Map
3.)
micropunctata (p. 238)
Underside of elytron without a supracostal flange but with a small, distinct ovate pit just
prior to apex. Dorsum dull yellow to light reddish brown. Head and pronotum slightly
darker in colour than elytron. Metasternum often distinctly darker in colour than rest
of underside. Length 10-0-13-0 mm. (Fig. 129.) Genitalia as in Figs 53, 86. (Map 3.)
montana (p. 239)
5 (1) Surface of elytron uneven with irregular smooth usually non-punctate raised areas or
rugosities which are often distinctly microsculptured. Head and pronotum with large
irregular punctures 6
Surface of elytron without such raised areas, puncturation more or less uniform
throughout 10
6 (5) Length 6-9-9-0 mm. Dorsum dark yellowish to orange-brown. Elytron usually slightly
lighter in colour than pronotum, with a golden to pinkish lustre. Pronotum c. l-6x
broader than long. Vertex, including post-antennal swellings, and pronotum with
230 SHARON L. SHUTE
large coarse confluent puncturation. Elytron with irregular longitudinal rows of long
sparse setae. (Fig. 138.)GenitaliaasinFigs68,96. (Map 3.) confusa(p. 232)
Length 10-0-15-0 mm. Coloration not as above, pronotal punctures not confluent
throughout. Other characters not so combined 7
7 (6) Length 8 • 5 mm . Ely tron pale reddish pink with a slight purplish lustre . Metasternum and
tibiae dark pitchy brown. Rest of body pale testaceous. (Fig. 140.) rugulosa(p. 241)
Length 10-0-15-0 mm. Coloration not as above 8
8 (7) Length 11-0-15-0 mm. Pronotum at least 2-Ox broader than long. Dorsum dark
orange-brown to pitchy black. Elytron may have a faint purplish lustre and pale lateral
margins. (Fig. 137.)GenitaliaasinFigs64,95. (Map7.) sedlaceki(p. 242)
Length 10-0-11-5 mm. Pronotum not more than l-8x broader than long. Elytron dark
purplish to reddish brown . Head and pronotum pitchy brown to black 9
9 (8) Femora except for apices distinctly lighter in colour than rest of legs. Derm between
humeri and sutural margin of elytron somewhat rugose and uneven. (Fig. 136.)
GenitaliaasinFigs66-67,93. (Map 5.) irregularis (p . 236)
Femora concolorous with tibiae, deep reddish to pitchy brown. Derm between humeri
and sutural margin of elytron not rugose. (Fig. 135.) Genitalia as in Figs 65, 94.
(Map3.) duplicator (p. 233)
10 (5) Frons convex, general elevation equal to or exceeding that of post-antennal swellings.
Elytron with or without carinae 12
Frons entirely depressed, general elevation less than that of post-antennal swellings.
Elytron carinate 11
11 (10) Elytron light orange to deep reddish brown, often with a strong, dark green lustre on
humeral area and over longitudinal sublateral depression. Head and pronotum
testaceous to light brownish red. Margins of post-humeral sublateral longitudinal
depression forming carinae which are joined at base of humeri, a third weakly
developed carina also present (more distinct in 9) and running more or less parallel to
inner carina, beginning at base of humeral area and becoming obsolete in apical
quarter at about same level as termination of inner carina. Length 8-0-10-0 mm.
(Fig. 127.) Genitalia as in Fig. 6. (Map 5.) venusta (p. 245)
Coloration variable, elytron light reddish brown to deep purplish blue or varying shades
of dark green. Pronotum testaceous to reddish orange or black. Margins of post-
humeral sublateral longitudinal depression forming carinae, not joined at base of
humeri, a third indistinct irregular carina also present and linked to inner carina of
depression by an oblique irregular convexity just below humeral area. All longitudinal
carina becoming obsolete in apical quarter. Length 7-0-12-0 mm. (Figs 146, 147, 148.)
Genitalia as in Figs 58-60, 100, 101. (Map 6.) varia(p. 243)
12 (10) Elytron not carinate, puncturation irregular, derm weakly rugose, dark testaceous or
green to pitchy brown 13
Elytron carinate , with fine regular puncturation , strong metallic blue/green 14
13 (12) Elytron dull greenish to pitchy brown, lateral margins normally lighter, derm weakly
rugose, puncturation more or less evenly distributed throughout. Head and pronotum
testaceous to light yellowish brown. Pronotum with dense coarse puncturation, derm
microsculptured. Scutellum triangular, apex acute. Length 7-5-12-0 mm. (Figs 144,
145.) Genitalia as in Figs 50, 83. (Map 4.) abdominalis(p.232)
Body except for head opaque testaceous to light orange-brown, elytron often with
irregular pigmentation. Head normally darker in colour than pronotum. Pronotum
with large non-confluent punctures, derm smooth and shining. Scutellum distinctly
longer than broad, apex truncate. Length 7-5-11-0 mm. (Fig. 143.) Genitalia as in
Figs 63, 97. (Map 3.) inhabilis(p. 235)
14 (12) Elytron with margins of post-humeral sublateral longitudinal depression forming cari-
nae which are joined at base of humeri, a weakly developed admedian carina may also
be present, running parallel to inner carina, derm often with a submetallic lustre 16
Elytron with only the outer margin of the post-humeral sublateral longitudinal depress-
ion forming a carina or ridge, inner margin sometimes partially developed but never
joined to outer, derm deep metallic blue or green.
Head, pronotum, scutellum and metasternites either dark orange to reddish
brown or black 15
15 (14) Elytron almost pyriform, bright metallic blue or green, outer margin of sublateral
GALERUCINE BEETLES OF NEW GUINEA 231
longitudinal depression forming an irregularly developed ridge, inner margin only
partially developed, sinuate. Pronotumc. l-8x broader than long with small indistinct
punctures 1-5-2-Ox their own width apart. Metepisternum microsculptured, punc-
tures minute, 2-5-3-Ox their own width apart. Median area of vertex raised and
impunctate. Length 9-5-13-0 mm. (Fig. 131.) Genitalia as in Figs 54, 98. (Map 7.)
suavis(p. 243)
Elytron subparallel-sided, dark metallic green, lateral margin often with bluish or
purplish lustre, outer margin of post-humeral sublateral depression forming a distinct,
well-developed, more or less straight ridge, inner margin indicated by a short
indistinct sinuate ridge. Pronotum at least 1-95X broader than long, punctures mostly
situated along anterior margin and in lateral area of depressions. Metapisternum
strongly microsculptured, punctures minute, indistinct due to microsculpture. Vertex
irregularly punctured. Length 9-5-13-0 mm. (Fig. 132.) Genitalia as in Figs 55-57, 99.
(Map?.) metallica(p. 238)
16 (14) Elytron dark submetallic green or brownish green, sometimes with distinctly lighter
lateral margins, but never lighter sutural margin. Pronotum testaceous to light
orange-brown. Head with vertex reddish to pitchy brown or concolorous with
pronotum 17
Coloration not as above 19
17 (16) At least three-quarters of lateral area of elytron, from outer margin of post-humeral
sublateral longitudinal depression to lateral margin, light reddish brown. Tibiae and
apices of femora pitchy brown . N. Australia, D'Entrecasteaux Islands 18
Elytron entirely dark metallic green. Legs unicolorous.
Head and pronotum testaceous to light reddish brown. Length 8-5-10-5 mm. (Fig.
123.) Genitalia as in Fig. 70. (Map 2.) laetabilis(p.231)
18 (17) Anterior area of head including post-antennal swellings testaceous, usually concolorous
with pronotum, vertex deep reddish brown. Lateral area of elytron entirely red to
reddish brown. Third admedian carina beginning at level equal to that of apex of
scutellum, present to apical quarter. Length 8-5-11-0 mm. (Fig. 121.) Genitalia as in
Figs 71, 87. (Map8.) obscuricornis(p. 239)
Head entirely dark reddish brown. Lateral margin of elytron reddish brown from level
equal to that of apex of metepisternum to apices. Third ad-median carina present from
level equal to that of base of humeri where lateral carinae join, to apical quarter.
Length 7-0-10-5 mm. (Fig. 125.) Genitalia as in Figs 74, 91 . (Map 2.)
purpurasco viridis(p. 241)
19 (16) Elytron light orange to reddish brown with a distinct, submetallic dark green longitudi-
nal band extending from humeral area to apical quarter, basal quarter sometimes
entirely dark green. Head anterior to post-antennal swellings, pronotum, scutellum,
underside and femora opaque, testaceous; rest of head, antennae, tibiae and tarsi
deep reddish to pitchy brown. Frons broadly triangular, not strongly convex. Elytral
puncturation never coarse or confluent. Length 6-9-10-0 mm. (Fig. 120.) Genitalia as
in Figs 72, 88. (Map 2.) costatipennis(p. 233)
Coloration not as above . Frons convex, usually with a horizontal median keel 20
20 (19) Elytron light orange to dark reddish brown with a distinct, pale submetallic pinkish
lustre which sometimes has a greenish reflection, lateral margins often lighter in
colour than rest of elytron. Pronotum, scutellum and underside testaceous to pale
orange-yellow, rest of body dark reddish brown, underside of femora often lighter in
colour than rest of legs. Length 8-9-12-5 mm. (Fig. 126.) Genitalia as in Figs 69, 95.
(Map2.) ftisdterafe(p.234)
Elytron reddish purple to deep bluish purple, sometimes with a bright bluish green lustre
on humeral area (in deep purple specimens this tends to extend over the whole
elytron). Head and pronotum concolorous, testaceous to purplish red. Tibiae norm-
ally darker in colour than femora. Teneral specimens orange to dull reddish brown
with a slight green or purplish lustre
21 (20) Elytron strongly or coarsely punctured, punctures sometimes confluent in places,
particularly on humeral area, admedian carina well developed in basal half. Head,
pronotum, scutellum, underside and legs either concolorous testaceous, or head,
pronotum and scutellum red with antenna and tibiae dark reddish to pitchy brown,
232 SHARON L. SHUTE
underside brownish orange. Elytron orange-brown to deep purplish red with a green
submetallic lustre over humeral area and often extending over sublateral depressions
(deep purple specimens have the dark green lustre extending over whole elytron).
Teneral specimens orange to dull reddish brown with a slight green or purple lustre.
Length 7-5-10-5 mm. (Fig. 122.)GenitaliaasinFigs75-76,89. (Map4.) Ae//eri(p. 235)
Elytron never coarsely punctured, punctures never confluent, on average 1 -0-1 -5 x their
own width apart, admedian carina not always well developed. Head, pronotum and
scutellum testaceous to light orange-yellow, underside testaceous to light orange-
brown. Antenna and tibiae dark reddish brown or testaceous to light reddish brown.
Elytron light purplish pink to deep reddish purple with a purplish blue or greenish blue
lustre. Teneral specimens with elytron light orange-brown with a light green or
purplish lustre. Length 8-0-11-0 mm. (Fig. 124.) Genitalia as in Figs 73, 90. (Map 2).
purpurascopurpurasco(p. 240)
Polysastra abdominalis (Jacoby) comb. n.
(Figs 50, 83, 144, 145, Map 4)
Sostra abdominalis Jacoby, 1904: 503. Syntype d", PAPUA NEW GUINEA: Moroka, 1300 m, vii-xi. [18]93
(Loria) (BMNH) [examined].
ADDITIONAL MATERIAL EXAMINED
Papua New Guinea: 112 ex., various localities (see Map 4) (AMNH, BMNH, IP, OIP, RWH, SAM).
COMMENTS. This species is representative of the abdominalis-group but it is not closely related to
any of the taxa described here. All the undescribed species of the abdominalis-group examined
are very similar in appearance, so care should be taken to check the genitalia of any specimen
that is assigned to this species.
Polysastra bicostata (Jacoby) comb. n.
(Figs 62, 149, Map 2)
Sostra bicostata Jacoby, 1894: 305. LECTOTYPE cf , IWAN JAVA: Andai, 1892 (W. Doherty) (MCZ), here
designated [examined].
ADDITIONAL MATERIAL EXAMINED
Irian Jaya: 1 $, Dor[ey] (Wallace) (BMNH).
COMMENTS. This species is representative of the bicostata-group. It is similar in appearance to
species of the costatipennis-group but may be distinguished by the distinct elytral setae.
Polysastra confusa sp. n.
(Figs 68, 96, 138, Map 3)
GENERAL FORM. Length 6-9-9-0 mm. Elytra subparallel-sided; post-humeral sublateral depression absent,
outer margin present as an irregular broken ridge, well developed on humeri in $ , surface of elytron with
irregular smooth raised areas throughout, areas between these rugosities strongly punctured, majority of
setae on elytron confined within margins of punctures, rest very long, fine and erect, forming irregular
longitudinal rows on interspaces. Head dark orange-brown, pronotum and scutellum either concolorous
with head or slightly lighter in colour; elytron either concolorous with pronotum or deeper, more reddish
brown, raised areas usually slightly darker in colour than rest of derm; underside unicolorous testaceous or
metasternum deep orange to reddish brown, basal three-quarters of femora testaceous, apical area dark
orange to reddish brown, tibiae and tarsi either concolorous with apices of femora or slightly lighter in
colour, setae translucent or pale golden.
DIAGNOSIS. Head with frons more or less evenly convex, elevation less than that of post-antennal swellings;
post-antennal swellings distinctly elevated above vertex, hind margins ill-defined due to coarse punctura-
tion; vertex with large, irregular, coarse, confluent punctures, derm strongly microsculptured, setae very
fine. Antenna with long fine suberect setae. Pronotum transverse, c. l-6x as broad as long, lateral tooth
acute , wholly in anterior half and apices inclining towards anterior margin , width across tooth 1 • 1 x greater
GALERUCINE BEETLES OF NEW GUINEA 233
than at its base, dorsal depressions ill-defined due to coarse puncturation. Puncturation coarse and
confluent throughout, punctures similar in size to those on vertex, setae very fine and suberect. Scutellum
triangular, apex subtruncate, length more or less equal to maximum width, punctures irregular, smaller
and shallower than those on pronotum but distinctly coarser than those on adjacent area of elytron; setae
fine and adpressed. Elytron c. 3-4x as broad as long, width increasing gradually from humeral angles to a
maximum at around apical quarter, maximum width l-3x greater than that at apex of scutellum, apices
evenly rounded, area between irregular raised areas strongly punctured, punctures on average not more
than their own width apart, those on humeral area slightly larger and denser than rest, but smaller and less
coarse than those on pronotum, derm with a vitreous lustre. Underside finely and minutely punctured
throughout. Metepisternum somewhat granulate, derm strongly microsculptured; punctures on metaster-
num 2-3 x their own width apart, becoming slightly denser and irregular along lateral margins, derm
smooth and shining, weakly microreticulate along lateral margins, setae on disc long, fine and suberect,
becoming slightly shorter towards sides; abdomen indistinctly punctured, punctures 3-4 x their own width
apart, becoming slightly irregular and denser at sides, derm smooth, weakly microsculptured at sides, setae
as on metasternum; femora finely punctured, punctures slightly larger and shallower than those on
metasternum, 3-4x their own width apart, becoming denser at apices, microsculpture becoming stronger
towards apices; tibiae finely granulate, setae shorter and more erect than on femora. Genitalia
(Figs 68, 96).
Holotype cf (dissected), Papua New Guinea: Bulldog Road, c. 14 km S. Edie Creek, 1405 m,
4-10.vii.1966, light-trap (G. A. Samuelson) (BPBM).
Paratypes. Papua New Guinea: 1 9 , same data as holotype; 1 9 , Wau, 2400 m, 9-12. i. 1962 (/. Sedlacek)
(BMNH, BPBM).
COMMENTS. This species belongs to the irregularis -group. Care must be taken not to confuse it
with an undescribed species from the same area that has similar facies, and a strongly
microsculptured pronotum with weaker non-confluent puncturation than in confusa; the 9 is
without the ridge on the elytral humerus (Fig. 139).
Polysastra costatipennis (Jacoby) comb. n.
(Figs 72, 88, 120, Map 2)
Sostra costatipennis Jacoby, 1886: 73. LECTOTYPE cf , PAPUA NEW GUINEA: Fly River, 1876-77 (L. M.
D'Albertis) (MCZ), here designated [examined].
ADDITIONAL MATERIAL EXAMINED
Papua New Guinea: 1 cf , 5 $ (paralectotypes), same data as lectotype (BMNH, MCZ); 1 $, between
Port Moresby and Brown River, 30 m, 29.x-l-xi.1965 (/. Sedlacek) (BPBM).
COMMENTS. This species is representative of the costatipennis-group and is most similar in
appearance to obscuricornis .
Polysastra duplicator sp. n.
(Figs 65, 94, 135, Map 3)
Length 9-5-12-2 mm. Externally very similar to irregularis. The only distinct difference is in the coloration
of the legs which, in duplicator, are unicolorous dark reddish. In comparison with irregularis, duplicator
has a less coarsely punctured pronotum and the elytron is not so rugose. The two species can be easily
separated by the genitalia characters (Figs 65, 94).
Holotype cf , Irian Jaya: Star Range, 2360 m, bivak 40, 21.vii.1959 (Neth. New Guinea Exp.) (RNH).
Paratypes. Irian Jaya: 3 cf , 7 $, same data as holotype (2 cf , 1 $ BMNH); 4 cf, 3 9 same data as
holotype except 22.vii.1939 (1 cf , 1 $ BMNH); 2 cf, same data as holotype except IS.vii., 20.vii.; 3 $,
same data as holotype except bivak 39a, 20.vi., 12.vii. and 23.vii.; 5 cf , 9 $, Paniai, 22.viii.-17.ix. 1939
(2 cf , 2 9 BMNH); 1 cf , Dejeresa, 31.ix.1939. (All RNH unless otherwise stated.)
Polysastra explanata sp. n.
(Figs 51, 84, 128, Map 3)
GENERAL FORM. Length 9-5-12-0 mm. Elytra subovate, setose, setae extending beyond margins of
punctures, lateral margins explanate, puncturation more or less uniform throughout, post-humeral
234 SHARON L. SHUTE
longitudinal sublateral depression almost absent but outer margin forming a well-developed ridge which
begins at base of elytron, extends over humerus and continues more or less straight to apical quarter, apex
evenly rounded; vertex, pronotum and scutellum coarsely and confluently punctured throughout. Head,
pronotum and scutellum concolorous testaceous to dark orange-brown; elytron light orange-brown to dark
reddish brown with a very weak, deep purplish lustre; underside testaceous to light orange-brown, femora
concolorous with underside or slightly darker, apices often dark pitchy brown, tibiae, tarsi and antennal
segments dark reddish brown; setae pale golden.
DIAGNOSIS. Head with frons convex, maximum elevation more or less equal to that of post-antennal
swellings, post-antennal swellings microsculptured, vertex punctured throughout, punctures confluent,
more or less equal in size to largest on pronotum, setae long and fine, adpressed. Pronotum transverse, on
average l-8x as broad as long, lateral margins with a broad, weakly developed tooth, width across tooth
c. 1-1 x greater than minimum posterior width, dorsal depressions distinct, puncturation coarse and
confluent throughout, derm strongly microsculptured, setae long, fine and suberect. Scutellum finely
punctured and strongly microsculptured, setae as on pronotum. Elytron becoming broader and lateral
margin becoming explanate from numeral angle to apical quarter, maximum width c. l-3x (cf) and l-5x
(?) greater than width at apex of scutellum, median area of basal quarter moderately convex with a small,
round shallow depression at base of scutellum; puncturation fine and uniform throughout, punctures not
more than their own width apart, size more or less equal to smallest on pronotum, derm finely
microsculptured; setae shorter and somewhat stouter than those on pronotum. Underside finely and
minutely punctured throughout. Punctures on disc of metasternum 3-4 x their own width apart, derm
weakly microsculptured, setae on disc shorter and finer than those at margins, puncturation of abdomen
similar to that of metasternum, setae becoming longer towards middle and sides of segments. Legs with
basal three-quarters of femora finely punctured, punctures 1-0-1-Sx their own width apart, apical quarter
granulate, setae long, fine and adpressed; tibiae granulate, setae short and stout, becoming denser towards
apices. Wing fully developed. Genitalia as in Figs 51, 84.
Holotype cf, Papua New Guinea: Kokoda, viii.1933, lower rain forest, 1300 ft (L. E. Cheesman)
(BMNH).
Paratypes. Papua New Guinea: 2 cf , same data as holotype; 3 cf, 3 $, Kokoda, 1200 ft, vi-viii.1933
(L. E. Cheesman); 1 cf, Maprik, 24.X.1957 (J. Smart) (BMNH); 1 cf, Wau, Morobe District, 1200 m,
25.xii.1961 (G. Monteith & J. Sedlacek); 1 $, Bulolo Vatus, 700-800 m, l-7.vi.1969 (/. Sedlacek); 2 £,
Kokoda-Pitoki, 450 m, 25.iii.1956 (J. L. Gressitt); 5 $, Tsenga 1200 m, Upper Jimmi V, 15.vii., 13.viii.,
14.viii.1955 (/. L. Gressitt) (1 Cf , 2 $ BMNH; 2 cf , 3 $ BMNH).
COMMENTS. This species is representative of the explanata-group. It does not bear any close
affinity to the other Polysastra species described here. Three undescribed species which closely
resemble this species and belong to this group have been examined, so care should be taken to
check the genitalia of specimens when assigning them to this species.
Polysastra fuscitarsis sp. n.
(Figs 69, 95, 126, Map 2)
GENERAL FORM. Length 8-1-12.1 mm. Elytra subparallel-sided, outer margins of post-humeral, sublateral
longitudinal, depressions forming distinct cariniform ridges which are joined at the base of the humeral
angle; a longitudinal admedian carina is also weakly indicated running parallel to inner ridge. Puncturation
of elytron uniform and dense, setae confined within margins of punctures. Head and pronotum distinctly
punctured. Head either entirely dark reddish to yellowish brown, or with area anterior to post-antennal
swellings distinctly lighter in colour than vertex. Pronotum and scutellum testaceous to orange-brown.
Elytron light orange to reddish brown with a submetallic, pale purplish pink lustre which is often dark
green over humeri in mature specimens. Underside testaceous to light orange-brown. Legs either entirely
dark orange to reddish brown, or femora testaceous with dark apices, tarsi normally darker brown than
tibiae. Setae pale golden. Fully winged.
DIAGNOSIS. Head with frons triangular and medially convex, weakly declivous anteriorly, maximum
elevation slightly greater than that of post-antennal swellings; post-antennal swellings well defined,
elevated above level of vertex, hind margins oblique, lateral margins distinctly separated from inner
margins of eyes by a shallow groove, derm smooth and shining, sparsely punctured. Vertex irregularly
punctured with groups of ill-defined, often confluent, shallow punctures, derm around punctures irregular-
ly depressed and weakly microreticulate, rest of derm smooth and shining, setae very fine, small and
adpressed. Pronotum transverse, on average 2-lx broader than long, lateral tooth well developed, apex
GALERUCINE BEETLES OF NEW GUINEA 235
subtruncate, width at tooth on average l-2x greater than minimum posterior width; lateral depressions
shallow and ill-defined due to lateral declivity of pronotum, median depressions shallow but distinct,
puncturation irregular, greatest density at sides where they may be contiguous or confluent, size variable,
largest more or less equal to largest on vertex, derm smooth and shining, weakly microsculptured in
strongly punctured areas, setae minute, fine and adpressed, those at sides confined within margins of
punctures. Scutellum slightly longer than broad, apex subtruncate, weakly convex medially, sparsely and
finely punctured, derm weakly microreticulate, setae short, fine and adpressed. Elytron c. 3-Ox longer
than broad, width increasing gradually from base of humeri to a maximum around middle which is 1-2 x
greater than width at apex of scutellum; outer margin of post-humeral sublateral longitudinal depression
forming a distinct ridge, extending from base of elytron over humerus to apical quarter where it becomes
obsolete, inner margin also forming a ridge which is joined to the outer at the base of the humeral angle and
continues to apical quarter where it becomes obsolete just above termination of outer margin, an admedian
longitudinal ridge also present, running parallel to inner ridge of depression, which is often indistinct or
obsolete for parts of its length. Puncturation more or less uniform throughout, punctures 2-0-2-5 x their
own width apart, their size less than half that of lateral pronotal punctures, derm smooth and shining,
weakly microreticulate in basal half, epipleura becoming obsolete about middle of apical quarter, anterior
half with a shallow median groove. Underside finely and densely punctured throughout, punctures on disc
of metasternum 3^tx their own width apart, becoming denser at sides, derm weakly microreticulate at
sides; metepisternum densely microsculptured, punctures 1-0-1-5X their own width apart, setae long and
fine on disc, becoming shorter and adpressed at sides. Abdomen finely punctured throughout, punctures
5-6 x their own width apart , setae long and suberect , stouter than those on metasternum , derm smooth and
shining, femora finely punctured, punctures on basal three-quarters 3-0-3-5 x their own width apart,
increasing in density to 0-5-1 -Ox width apart at apices, derm weakly microsculptured at apices, setae long,
fine and adpressed, tibiae finely and densely punctured, becoming granulate towards apices, setae
becoming shorter, stouter and more erect towards apices. Genitalia as in Figs 69, 95.
Holotype cf , Irian Jaya: Bernhard camp, 50 m, vii-xi.1939 (/. Olthof) (Neth. Ind.-Amer. New Guinea
Exped.) (BPBM).
Paratypes. Irian Jaya: 15 cf , 44 $, same data as holotype (7 O", 20 9 BMNH); 3 cf , 4 $, lebele camp,
2250 m, xi.1938 (L. J. Toxopeus); I $, Idenburgh River, 400 m, 15.vii.-15.xi.1938 (/. Olthof); 1 $,
letterbox camp, 3600 m, 27. ix. 1938 (L. J. Toxopeus) (BPBM unless otherwise stated).
Non-paratypic material. 12 specimens, same data as holotype (BPBM).
COMMENTS. P. fuscitarsis belongs to the costatipennis-group. The species described here to which
it bears the closest affinity is purpurasco .
Polysastra helleri (Weise) comb. n.
(Figs 75, 76, 89, 122, Map 4)
Sastm helleri Weise, 1917: 207. LECTOTYPE $, PAPUA NEW GUINEA: Toricelli Mts, i.1910 (Schlagin-
haufen) (SMT), here designated [examined].
ADDITIONAL MATERIAL EXAMINED
Papua New Guinea: 1 cf , 5 $ (paralectotypes), same data as lectotype (1 cf , 3 $ SMT; 2 $ NR). Irian
Jaya, Papua New Guinea: 6 cT, 11 $ , various localities (Map 4) (BMNH, BPBM, MCZ, NR, RHW, SMT).
COMMENTS. This species belongs to the costatipennis-group. Of the species dealt with here, it is
closest to laetabilis.
Polysastra inhabilis sp. n.
(Figs 63, 97, 143, Map 3)
GENERAL FORM. Length 7-5-11-0 mm. Elytra subparallel-sided, post-humeral sublateral depressions
almost absent, outer margins forming weak ridges which are distinct over humeri in $, surface of elytron
irregularly but uniformly punctured, very weakly rugose, setae minute, hardly extending beyond margins
of punctures. Head and pronotum distinctly punctured. Head and pronotum either concolorous testaceous
to dull reddish brown or with head distinctly darker in colour than pronotum. Elytron opaque testaceous to
dull brownish yellow, basal sutural angle may be dark reddish brown; derm with a weak vitreous lustre in
basal half. Underside testaceous to dull brownish yellow. Legs pale testaceous, apex of femora often dark
reddish brown, setae pale golden. Fully winged.
236 SHARON L. SHUTE
DIAGNOSIS. Head with broad triangular frons, slightly flattened dorsally, elevation less than that of
post-antennal swellings, interantennal groove deep; post-antennal swellings strongly convex and elevated
above level, of vertex, hind margins rounded, sometimes ill-defined due to strong puncturation of vertex,
lateral margins of swellings delimited by a narrow groove, surface irregularly punctured; vertex with
strong, coarse, irregular puncturation which is particularly dense and often confluent behind post antennal
swellings, punctures on centre of vertex often elongated, size variable, largest punctures more or less equal
to largest on pronotum; derm shining, weakly microsculptured in areas of strong puncturation; median
'suture' almost absent; setae short and fine, adpressed or suberect. Pronotum transverse, c. l-9x broader
than long, lateral tooth broad, moderately well defined but often unevenly developed, width at tooth l-2x
greater than minimum posterior width; sublateral depression weak and ill-defined, median depressions
distinct, anterior depression de-limited behind by two small adjacent convexities; surface distinctly
punctured throughout, puncturation irregular, punctures c. 0-5-1-Ox their own width apart, contiguous in
places, in size more or less equal to those on vertex; derm smooth and shining; setae minute and adpressed.
Scutellum l-4x longer than broad, derm with small shallow punctures, setae short and adpressed. Elytron
c. 4-5 x longer than broad, width increasing gradually from below humeral angles to a maximum just below
mid line, maximum width l-2x greater than that at apex of scutellum; puncturation irregular, size more or
less uniform throughout, punctures small and deep, less than half size of those on pronotum, tending to
form small, horizontal oblique groups of c. five punctures separated from one another by not more than
their own width; interspaces very weakly convex, giving a slight rugose appearance; setae minute, only just
extending beyond margin of punctures; those in vestigial striae indistinct, sparse, short and decumbent.
Epipleuron narrow, not becoming completely reflexed until apical quarter where it becomes obsolete.
Underside finely and densely punctured throughout, disc of metasternum almost impunctate, width
between punctures decreasing from 4-5 x width of a puncture to less than 2x at sides of metasternum;
derm becoming microsculptured towards sides; setae long and erect on disc, short and decumbent at sides,
metepisternum very finely and densely punctured, derm strongly microsculptured, setae adpressed,
punctures on abdominal segments fine and shallow, 1-5-2-Ox their own width apart, sometimes ill-defined
due to strong microsculpture on derm; setae long, fine and subadpressed, length decreasing slightly
towards sides. Femora finely and sparsely punctured, punctures 3-6 x their own width apart; derm
microsculptured; setae long fine and adpressed, becoming shorter at apices. Tibiae finely and densely
punctured, punctures 0-5-1 -Ox their own width apart, setae short, dense and stout. Genitalia as in Figs 63,
97.
Holotype cf , Irian Jaya: Lake Habbema, 3250-3300 m, vii-viii.1938 (L. J. Toxopeus} (Neth. Ind.-
Amer. New Guinea Exp. 1938) (BPBM).
Paratypes. Irian Jaya: 3 $ , same data as holotype; 3 cf, 4 $ , same data except 4.ix. (2 cf , 2 $ BMNH);
4 Cf , 2 $, Moss Forest Camp, 2800 m, 9.x.-5.xi.l938 (L. J. Toxopeus) (1 Cf , 1 $ BMNH).
COMMENTS. P. inhabilis is representative of the inhabilis-group. Several undescribed and similar
species have been examined, therefore care should be taken to check the genitalia when
assigning specimens to this species.
Polysastra irregularis sp. n.
(Figs 66-67, 93, 136, Map 5)
GENERAL FORM. Length 10-0-11-5 mm. Elytra subparallel-sided, post-humeral sublateral longitudinal
depressions shallow and ill-defined, outer margins forming ridges, surface of elytron rugose, punctures
grouped between irregular non-punctate raised areas, majority of setae confined within margins of
punctures, rest long, forming sparse, erect longitudinal rows. Head, pronotum and scutellum deep reddish
to pitchy brown, elytron either concolorous with pronotum or lighter purplish red. Femora, except for
apices, and abdomen testaceous. Antennae, apices of femora, tibiae and tarsi reddish to pitchy brown.
Metasternum pitchy brown to black. Setae grey or pale golden. Fully winged.
DIAGNOSIS. Head with frons somewhat flattened medially, weakly declivous anteriorly, sides more or less
equal in elevation to post-antennal swellings which are distinctly elevated above level of vertex, hind
margins of swellings ill-defined due to coarse, irregular, mostly confluent puncturation of vertex, derm
microsculptured, setae very fine. Pronotum transverse, c. l-8x broader than long, lateral tooth well
developed, width across tooth l-2x minimum posterior width, lateral depressions distinct, median
depressions fairly well defined, puncturation irregular, punctures becoming denser at sides, majority of
GALERUCINE BEETLES OF NEW GUINEA 237
punctures not more than half their own width apart, sparser in and around median depressions, derm
microsculptured, setae fine and adpressed, majority not extending much beyond margins of punctures.
Scutellum triangular, l-2x longer than broad, punctures slightly smaller and shallower than those on
pronotum, mostly confined to median area, setae fine and adpressed, extending beyond margins of
punctures. Elytron c. 3-8x as long as broad, width increasing gradually from humeral angle to a maximum
just anterior to apical quarter, maximum width c. l-3x greater than width at apex of scutellum, apices
evenly rounded, surface irregularly rugose, raised areas between punctures distinctly microsculptured,
punctures on average not more than their own width apart, size fairly uniform throughout, those on
humeral area slightly larger than rest. Underside finely and densely punctured throughout, metepisternum
strongly microsculptured. Punctures on anterior region of metasternum 1 -5-2-0 x their own width apart,
becoming sparse and almost absent on median area of posterior half, microsculpture present towards sides
of metasternum, setae long and fine, becoming shorter and more adpressed at sides. Punctures on
abdominal segments l-O-l-Sx their own width apart. Femora finely and irregularly punctured, punctures
4-0-5-Ox their own width apart on basal three-quarters and 1-5-2-Ox their own width apart on apical
quarter, setae long, fine and adpressed. Tibiae finely granulate, setae shorter and stouter than on femora,
density increasing towards apices. Genitalia as in Figs 66-67, 93.
Holotype cf , Papua New Guinea: NE. Morobe, Mt Kaindi, 2350 m, 18.vi.1973 (/. L. Gressitt) (BPBM).
Paratypes. Papua New Guinea: 2 cf , 9 9 , same data as holotype (1 cf , 4 9 BMNH); 5 9 , Morobe District
(E), Mt Kaindi nr Wau, 2350 m, 3.ix.l973 (J. L. Gressitt); 2 cf, 7 9, Mt Kaindi, 2400 m, 27.U963,
28.L1963 (/. Sedlacek) (1 cf, 3 9 BMNH); 5 9, Mt Kaindi, 2300 m, 2.iii.l966, 6.iv.l966, 4.iv.l966,
4.V.1967, 15.ix.1973, 2.U974 (/. L. Gressitt); (2 9 BMNH); 1 9, Mt Kaindi, 2350 m, 7.iv.l966, Malaise
trap (J. L. & M. Gressitt); 2 9, Mt Kaindi, 2350 m, 22.vii.1974, 9-1.X.1974 (A. D. Hart); 3 9, Mt Kaindi,
2300m, lO.i. 1962, 8-9.vi. 1962, 1-5.1962(7. Sedlacek)(l 9 BMNH) ; 2 9, Mt Kaindi, 2250m, 10.v.l968(/.
Sedlacek); 1 9 , Mt Kaindi, 2300 m, 22.iii.1964 (Josef Ku); 1 cf , 3 9 , Mt Kaindi, on Meari Creek 9-5 mi[les]
from Wau, 2050 m, 12.iii.1959 (L. T. Brass) (2 9 BMNH); 2 9, Wau, Edie Creek, 189 m, 8.viii.l963
(P.S.E.L.); 1 9, Wau, Edie Creek, 2000 m, 4-10.X.1961, m.v. light-trap (J. &J. H. Sedlacek); 1 9, Edie
Creek, 14 km SW. Wau, 2000m, 27. v. 1962 (J. L. Sedlacek), 1 9, Edie Creek, 7000ft, stn no. 6, 17.ix.1964
(M. E. Bacchus) (BMNH); 1 9, Wau, 1000-1250 m, 3.iii.l964 (/. Sedlacek); 1 9, Wau, 2400 m,
9-12.U962 (/. & J. H. Sedlacek, G. Monteith & native collector); 1 9, Wau, 1200 m, 26-27. ix. 1964, m.v.
light-trap (J. Sedlacek); 2 cf , 1 9, Owen Stanley Range, Goilala, Borne, 1950 m, 24.ii-15.iii. 1958 (W. W.
Brandt); 1 cf, same data except Loloipa, 21-31. xii. 1957; 1 9, Central District, Mt Goliath, 5000-7000 ft
(S. Meek) (MCZ); 1 9 , Juliana bivak, 1800 m, l.ix.1959 (RHN). (All BPBM unless otherwise stated.)
COMMENTS. This species is representative of the irregularis-group. The species described here to
which it bears the closest affinity is duplicator.
Polysastra kampeni (Weise) comb. n.
Sastra kampeni Weise, 1917: 207. Type, PAPUA NEW GUINEA: Hoofdbivak [on Sepik River, 4°4'S,
141°15'E], xi.1910 (depository unknown).
COMMENTS. This species belongs to the costatipennis-group. It has not been possible to locate the
type or any specimens determined as such, and the transfer is based on characters given in the
original description.
Polysastra laetabilis (Weise) comb. n.
(Figs 70, 123, Map 2)
Sastra laetabilis Weise, 1917: 206. LECTOTYPE cf , PAPUA NEW GUINEA: Hoofdbivak (Sepik River),
12.x. 1910 (Kampen) (NR). (NR), here designated [examined].
ADDITIONAL MATERIAL EXAMINED
Papua New Guinea: 1 9, same data as lectotype except 12-16.xi.1910 (NR); 1 cf , May River, 6.vi.l963
(R. Stratman) (BMNH); 1 cf , Mindimbe, Sepik River, 25.iv.1963 (R. Stratman) (BPBM).
COMMENTS. This species belongs to the costatipennis-group and has the same type of aedeagus as
purpurasco.
238 SHARON L. SHUTE
Polysastra metallica (Jacoby) comb. n.
(Figs 55-57, 99, 132, Map 7)
Sastra metallica Jacoby, 1886: 72. Holotype $, NEW GUINEA: 'Ramoi, ging [18]72' (L. M. D'Albertis)
(MCSN) [examined].
Gronovius andaiensis Jacoby, 1905: 500. Holotype $, IRIAN JAVA: Andai (BMNH) [examined]. Syn. n.
ADDITIONAL MATERIAL EXAMINED
1 $, No. 88, 4.viii.l903 (no further data) (ITZ). Papua New Guinea, Irian Jaya: 127 examples, various
localities (Map 7) (BMNH, BPBM, MCZ, OIP, SMT).
COMMENTS. This species belongs to the metallica-group. Of the species described here it is closest
to suavis; however, there are also a number of undescribed species which are similar in
appearance to metallica, therefore the genitalia should be examined when determining speci-
mens.
Polysastra micropunctata sp. n.
(Figs 52, 85, 130, Map 3)
GENERAL FORM. Length 10-5-11-5 mm. Elytra subpyriform, setae confined within margins of punctures,
puncturation fine and dense throughout, post-humeral sublateral depressions absent, apices rounded,
underside of elytron with a well developed supracostal flanage which runs parallel to epipleuron, from base
of elytron to apical quarter where it curves inwards to meet sutural margin (Fig. 130), appearing as a
narrow groove on dorsal surface. Vertex, pronotum and scutellum densely and confusely punctured
throughout. Head, antennae, pronotum, scutellum, legs and underside except for abdomen, deep purplish
brown to black, abdomen testaceous; elytron deep purplish brown to pitchy brown with a very weak, deep
purplish green lustre, lateral margins bright orange-red from below humeral quarter (at a level equal to
that of apex of metepisternum) to apex. Setae grey.
DIAGNOSIS. Head with frons convex, weakly declivous anteriorly, elevation more or less equal to that of
post-antennal swellings, post-antennal swellings well defined, distinctly elevated above vertex, vertex with
an area of irregular, large, shallow confluent punctures directly behind swellings, rest of vertex finely and
densely punctured throughout, punctures similar in size to smallest punctures on pronotum, derm strongly
microsculptured, median area of vertex posterior to eyes with a shallow, oblique, ovate depression either
side of 'suture'; setae very fine. Pronotum transverse, c. l-8x as broad as long, lateral tooth well
developed, width at tooth on average l-2x greater than minimum posterior width, sublateral depressions
well defined, median depressions shallow and ill-defined, punctures on disc shallow, distinctly larger than
rest, remainder of pronotum finely and densely punctured throughout, often ill-defined due to strong
microsculpturing of derm, setae long and fine, slightly shorter and stouter in median depressions.
Scutellum, finely and densely punctured and strongly microsculptured, setae as on pronotum. Elytron
3-4x longer than broad, becoming gradually broader from just below humerus to a maximum width at
around middle, after which it gradually decreases towards apex, maximum width in 9 c. l-3x greater than
width at apex of scutellum, slightly less in cf ; dorsum more or less evenly convex, slight depression just
prior to humeri below apex of scutellum. Puncturation irregular and dense throughout, punctures 1-2 x
their own width apart, more or less uniform in size, derm strongly microsculptured, setae shorter and
denser than on pronotum, majority pale grey interspersed with slightly stouter white setae, ventral surface
of elytron with a well developed supracostal flange which runs parallel to epipleuron from base to apical
quarter where it curves inward to meet sutural margin. Underside finely and densely punctured through-
out, punctures not more than l-5x their own width apart, derm microsculptured, setae on abdomen
slightly longer than those on rest of segments. Legs with femora minutely and densely punctured,
punctures not more than their own width apart, setae long, fine and adpressed, tibiae granulate, setae short
and stout, increasing in density towards apices. Wing fully developed. Genitalia as in Figs 52, 85.
Holotype cf , Papua New Guinea: Morobe District, Arabuka, 1500-2000 m, 7.J.1968 (/. & M. Sedlacek)
(BPBM).
Paratypes. 2 $ , same data as holotype (BPBM, BMNH).
COMMENTS. This species represents the micropunctata-group and is one of those defined in
subgroup 1. The species described here to which it bears the closest affinity is montana from
GALERUCINE BEETLES OF NEW GUINEA 239
which it can be readily distinguished by the presence of a supracostal flange on the underside of
the elytron.
Polysastra montana sp. n.
(Figs 53, 86, 129, Map 3)
GENERAL FORM. Length 10-0-13-1 mm. Elytra subparallel-sided, post-humeral sublateral depressions
absent, setae extending well beyond margins of punctures, puncturation fine, dense and more or less
uniform throughout; head, pronotum and scutellum densely and confusely punctured throughout. Head,
antenna, pronotum and legs dark orange-brown to reddish brown, scutellum usually darker brown than
pronotum, elytron either concolorous with pronotum or lighter yellowish to orange-brown, derm with a
slight aeneous lustre; abdomen testaceous, rest of underside dark reddish to pitchy brown; setae pale
golden.
DIAGNOSIS. Head with frons flattened dorsally, post-antennal swellings with hind margins ill-defined,
surface coarsely punctured with large shallow confluent punctures; surface of vertex irregular, area directly
behind swellings coarsely and irregularly punctured, rest of area finely and densely punctured throughout,
punctures not more than their own width apart, median area of vertex posterior to eyes with a shallow
oblique, ovate admedian depression, derm strongly microsculptured throughout, setae long, fine and
adpressed. Pronotum transverse, 1-7-1-Sx as broad as long, sometimes slightly less in cf , lateral tooth
acute, width at tooth l-2x greater than minimum posterior width, sublateral depressions present but
shallow and ill-defined, puncturation fine and dense throughout, punctures not more than half their own
width apart, sometimes indistinct due to strong microsculpture on derm, setae similar to those on vertex.
Scutellum c. l-4x longer than broad, weakly convex medially, finely and densely punctured throughout,
derm strongly microsculptured, setae as on elytron. Elytron 4x as long as broad, width increasing very
gradually from below humeral quarter to a maximum at about apical quarter, maximum width 1-2-3-Ox
greater than that at apex of scutellum, apex evenly rounded, underside of elytron with a small, shallow
ovate pit just prior to apicosutural angle. Puncturation minute and more or less uniform throughout,
punctures not more than their own width apart, derm microsculptured, setae adpressed, slightly shorter
than those on pronotum. Underside finely and minutely punctured throughout, punctures not more than
l-5x their own width apart, those on metepisternum and adjacent area of metasternum becoming
granulate, derm more strongly microsculptured in these areas, setae on disc of metasternum and centre of
abdominal segments longer than those at sides. Legs with femora minutely punctured throughout,
punctures 2-3 x their own width apart, derm distinctly microsculptured, setae on lateral surfaces distinctly
longer and finer than rest; tibiae granulate, setae short and stout. Genitalia as in Figs 53, 85.
Holotype cf (dissected), Papua New Guinea: Mt Dayman, Maneau Range, 2230 m, N. slope no. 4,
19.v.-19.vi.l953 (G. M. Tote) (AMNH).
Paratypes . Papua New Guinea : 2 cf , 7 $ , same data as holotype ( 1 cf , 1 $ BMNH , rest AMNH) ; 1 $ , Mt
Suckling, Exp. Camp Mau 2, 1700 m, 2.vii.l972, black light (T. L. Fenner} (BMNH).
COMMENTS. This species belongs to the micropunctata-group and represents those species of
subgroup 1 that lack the supracostal flange on the underside of the elytron. The species
described here to which it bears the closest affinity is micropunctata from which it can be readily
distinguished by the absence of a supracostal flange on the underside of the elytra.
Polysastra obscuricornis (Blackburn) comb. n.
(Figs 71, 87, 121, Map 8)
Sastra obscuricornis Blackburn, 1896: 84. Holotype $, AUSTRALIA: N. Queensland], no. 6040 (BMNH)
[examined].
ADDITIONAL MATERIAL EXAMINED
Australia: 65 examples, N. Queensland, various localities (Map 8) (ANIC, BMNH, MCZ, NMV, NR,
SAM,UQ).
COMMENTS. This species belongs to the costatipennis-group and is most closely related to
costatipennis .
PLANT ASSOCIATE. Laportea sp.
240 SHARON L. SHUTE
Polysastra purpurasco sp. n.
(Figs 73, 74, 90, 91, 124, 125, Map 2)
GENERAL FORM. Length 8-0-11-3 mm. Elytra subparallel-sided, post-humeral sublateral longitudinal
depressions present, outer margins forming cariniform ridges which are joined at base of humeri; an
irregularly developed ad-median carina present, running more or less parallel to inner margin of
depression; median area of basal quarter convex; puncturation more or less uniform throughout, vestigial
strial setae short and erect, rest of setae minute and confined within margins of punctures. Head and
pronotum distinctly punctured. Head testaceous to pitchy black, pronotum testaceous to dull reddish
brown, underside and femora testaceous to orange-brown, tibiae distinctly darker in colour than femora;
elytron light reddish brown to pinkish purple or deep submetallic green, setae pale golden. Fully winged.
DIAGNOSIS. Head with frons triangular, strongly convex medially, maximum elevation slightly greater than
that of post-antennal swellings, post-antennal swellings not strongly convex, hind margins often ill-defined
and confluent with vertex. Vertex irregularly punctured with large, shallow punctures which are mostly
concentrated behind post-antennal swellings, confluent in places; derm weakly microsculptured with a
vitreous lustre, setae short and fine, mostly adpressed. Pronotum transverse, on average 2-lx broader
than long, lateral tooth not well developed, apex rounded, width at tooth l-2x greater than minimum
posterior width. Dorsal depressions distinct, puncturation dense and irregular, average puncture size more
or less equal to largest on vertex, often indistinct due to vitreous lustre of derm; setae minute, decumbent.
Scutellum with maximum width more or less equal to length, apex rounded, derm microsculptured with
small shallow irregular punctures, setae minute, decumbent. Elytron on average 3-7x longer than broad,
width increasing gradually from humerus to a maximum around middle, maximum width l-2x greater than
width at apex of scutellum; cariniform margins of post-humeral sublateral depression present from base of
humeral angle, where they are joined, to apical quarter where they become obsolete, carinae more or less
parallel and separated by the width occupied by c. 6-7 punctures, an irregularly developed ad-median
carina present, running more or less parallel to inner margin of depression and becoming obsolete in apical
quarter, width between ad-median carinae and inner marginal ridge slightly less than c. width of 5
punctures, area on basal quarter between ad-median carina and sutural margin convex, puncturation more
or less uniform throughout, slightly coarser on basal quarter, punctures slightly smaller than those on
pronotum, 0-5-0- 15 x their own width apart, derm smooth and shining. Underside finely and densely
punctured throughout, punctures on metasternum 3-4 x their own width apart, becoming slightly denser
along lateral margins; derm smooth and shining, setae short, fine, decumbent, becoming adpressed along
margins; metepisternum strongly microsculptured, punctures dense, 1-0-1-5X their own width apart, setae
short and adpressed. Abdomen minutely punctured, punctures 3-4x their own width apart, derm weakly
microsculptured, setae very fine, longer than on metasternum, particularly at sides, femora finely
punctured with small shallow punctures 3-4 x their own width apart on basal three-quarters, dense at
apices, setae long fine and adpressed; tibiae finely granulate, setae becoming shorter and more erect
towards apices.
COMMENTS. This species belongs to the costatipennis-group and is mostly closely related to
obscuricornis, having the same type of facies and aedeagus; apart from genitalia and coloration
it differs principally by having the elytral margins less explanate and the carinae better
developed, and the elytra are also slightly less densely punctured. For genitalic differences see
Figs 71, 73. P. purpurasco also bears close affinity to fuscitarsis .
Polysastra purpurasco purpurasco subsp. n.
(Figs 73, 90, 124, Map 2)
Length 8-0-11-0 mm. Head, basal segments of antenna and scutellum pale testaceous to dull yellowish
brown; elytra light reddish brown to deep pinkish purple, mature specimens with a deep bluish purple
lustre over humeral area and sublateral depression. Underside testaceous, abdomen often dull orange-
brown, femora testaceous to brownish yellow, apices of femora and rest of legs dark reddish brown. Head
and pronotum moderately to densely punctured. General form slightly less robust than island subspecies.
Genitalia as in Figs 73, 90.
Holotype d", Papua New Guinea: Wau, Morobe District, Hospital Creek, 1200 m, i.1965 (J. Sedlacek)
(BMNH).
Paratypes. Papua New Guinea: 4 cf , 8 $, same data as holotype (2 $, 3 $ BMNH); 2 $, Wau, 1050 m,
GALERUCINE BEETLES OF NEW GUINEA 241
4.xi.l961; 1 O", Wau, 1150 m, 12.ix.1961 (all J. Sedlacek); 1 £, Wau, 1200-1300 m, 23.xii.1961 (G.
Monteith); 3 cf, 2 $, Wau, 1200 m, 13.viii.-14.x.l961 (/. Sedlacek); 4 cf, 4 $>, Wau, 1200 m, light-trap,
14.vi.-22.x.l961; 2 cf , 7 $, Wau, 1200 m, m.v. light-trap, Il.x.-25.xi.l961 (all/. J. & M. Sedlacek) (2 cf ,
3 9 BMNH); 1 $, Wau, 1700-1800 m, 17.xi.1961; 1 cf,5 $, Wau, 1200m, 1.V.-10.X.1962; 1 cf,4 $, Wau,
1200m, light-trap, 16.ii.-15.vii.1962; 1 cf,2 $, Wau, 1200m, m.v. light-trap, 2.x., 28.xii. 1962; 7 $, Wau,
1250 m, 5. i. 1963, m.v. light-trap (2 $ BMNH); 1 $, Wau, 1200 m, 5.x. 1962, Malaise trap; 4 cf, 2 $>,
1250 m, 14. i., 21. i. 1963 (1 CM $ BMNH); 2 cf , 9 9, Wau, 1200 m, m.v. light-trap, 4.i.-23.x.l963 (1 cf , 2
$ BMNH); 6 $, Wau, 1200-1250 m, 4.ii.-19.viii.l964 (all /. Sedlacek); 3 cf, 3 9, 1200 m, 26.iii.-
30.vii.1964 (J. & J. Sedlacek); 3 cf , 12 $, same data except m.v. light-trap (2 $ BMNH); 1 cf , Wau,
1200-1300 m, 22.x. 1965 1 $, Wau, 1700-1800 m, 27.ix.1965; 1 cf, Wau, 1100-1300 m, l.ii.1966 (all/.
Sedlacek); 1 $, Wau, 1200 m, 25.viii.1966 (G. A. Samuelson); 1 $, Wau, 1200 m, iv.1966, light-trap (/. L.
Gressitt); 2 cf , Wau, 1200m, 14.iii.1966 (/. L. Gressitt & Wilkes); 1 $, Wau, 1200m, 26.iii.1966, light-trap
(/. L. Gressitt) (BMNH); 2 cf, Wau, 1200 m, 3.H.1966 (/. Sedlacek); 1 $, Wau, 1200 m, 24.iii.1968,
Acalypha sp. (J. Sedlacek); 1 cf , Wau, 1200 m, i.1968 1 ?, Wau, 8.ix.l968 (all 7. & M. Sedlacek); 1 $,
Wau, 1200 m, 28. i. 1974, K 606, Pipturus; 1 cf , 1200 m, 27.V.1974, at light (A. D. Hart) (WEI); 1 cf , Wau,
1150 m, 6.HU974, beaten from Coffea arabica (J. J. H. Szent-Ivany); 1 cf , Wau, 1200 m, 24.vii.1974,
Myrtaceae, Eugeni stipularis (G. Otaweto) (WEI); 1 cf , Wau, Wau Creek, 1200-1500 m, 28.iii.1963; 1 cf ,
6 km W. of Wau, Nami Creek, 1700 m, 12.vi.1962 (all /. Sedlacek); 1 cf , Wau, Coviak Ridge, 763 m,
7.xii.l963 (H.C.); 1 9, 32 km SW. of Lae, 100 m, 23.iii.1963 (/. Sedlacek); 2 cf, Garaina, 550-750 m,
16.1.1968; 1 Cf, Garaina-Saureli, 900-1400 m, 5. i. 1968 (all/. & M. Sedlacek); 1 $, Karimui, 4.vi.l961,
light-trap (/. L. Gressitt); 1 $, Kassam, 1350 m, 48 km E. of Kainantu, 7.xi.l959 (T. C. Mad); 1 cf , Owen
Stanley Range, Goilala-Loloipa, 16-30.xi.1958 (W. W. Brandt); 1 $, Tsenga, 1200 m, Upper Jimmi V,
15.vii.1955, light-trap (all/. L. Gressitt); 2 $, Finisterre Range, Saidor, Matoko village, 6-24. ix. 1958 (W.
W. Brandt); 1 $, Wau, 14.ii.1970 (OIP); 2 $, Gadsup, ix.1973 (all H. Ohlmus); 1 $, Popondetta area,
8-10.iv.1966 (R. Rodzyork), 25521; 1 cf , Melambi River, Lae, Mirilunga village, 4500 m, 29.xii.1956 (/.
H. Ardley), 19073; Kaparvia-Sangi, or Kododa, 600 m, 26.111. 1956 Curcuma; Icf 1 $, Wapenamanda
School, 5700 ft, West Highlands, 21.iii.1960 (/. H. Barrett), 25596; 1 $ , Luth, Mission Garden, Wau, 3500
ft, 2.vi.l957 (/. /. H. Szent-Ivany), 19050; 1 $, Bisianumu-Sogeri, Agri. Exp. Sta., 20.iii.1955 (/. /. H.
Szent-Ivany & A. Himson), 19051 (allPNGDA); 1 cf, 1 $,MadangDist.,FinisterreMts,Budemu,c. 4000
ft, 15-24.X.1964 (BMNH); 4 $, Madang Dist., Finisterre Mts, Damanti, 3550 ft, 2-11. x. 1964 (all M. E.
Bacchus), 1 cf, Kokoda, 1200ft, ix. 1933; 1 $, Mondo, 5000ft, iii. 1934 (all L. E. Cheesman) (all BMNH); 1
Cf, New Britain, Gazelle Pen., Gaulim, 130 m, 28.xi.1962 (/. Sedlacek). (All BPBM unless otherwise
stated.)
PLANT ASSOCIATES. Coffea arabica, Curcuma sp. , Pipturus sp. , Eugeni stipularis, Acalypha sp.
Polysastra purpurasco v/ridis subsp. n.
(Figs 74, 91, 125, Map 2)
Length 8-0-11-3 mm. Differs from the nominate mainland form as follows. Head deep reddish brown to
pitchy black. Pronotum and scutellum opaque, testaceous, vitreous lustre lacking. Elytra dark submetallic
green with a slight aeneous lustre, longitudinal area between lateral margin and carina formed by outer
margin of sublateral depression bright reddish to orange-brown. Underside testaceous to brownish orange.
Femora except for apices testaceous, apices and rest of legs pitchy brown. Head and pronotum tend to be
more strongly and densely punctured than in nominate mainland form. No other obvious structural
differences have been observed. Genitalia as in Figs 74, 91.
Holotype cf (dissected), D'Entrecasteaux Islands: Normanby I., Wakaiuna, Sewa Bay, 1-18. i. 1957
(W. W. Brandt) (BPBM).
Paratypes. D'Entrecasteaux Islands: 2 $, same data as holotype (BPBM, BMNH); 3 $, Fergusson I.,
ix-xii.1894 (A. S. Meek) (MCZ); 1 $, Fergusson L, mountains between Agamoia and Ailuluai, 900 m,
no. 4, 5-7.vi.1957 (L. T. Brass) (AMNH).
Polysastra rugulosa (Weise) comb. n.
(Fig. 140)
Sastra rugulosa Weise, 1912: 438. Holotype $, NEW GUINEA: Erima (MNHU) [examined].
COMMENTS. This species belongs to the irregular is -group and is representative of subgroup 4.
242 SHARON L. SHUTE
Polysastra sedlaceki sp. n.
(Figs 64, 95, 137, Map 7)
GENERAL FORM. Length 11 -0-15-0 mm. Elytra subparallel-sided, post-humeral sublateral depression
present from base of humeri to just above apical quarter, outer margin of depressions forming a weak
irregular ridge , surface of elytra weakly rugose with irregular smooth non-punctate raised areas , punctura-
tion dense and irregular. Vertex and pronotum with large, shallow irregular punctures. Setae on elytron
minute, majority confined within margins of punctures, few sparse long setae on intervals. Dorsum dull
pinkish or reddish brown to deep purple-brown, outer margins of elytron often paler in colour, humeral
area often with a slight greenish or pinkish lustre. Antenna and legs deep reddish to pitchy brown.
Abdomen testaceous , rest of underside pitchy brown to black ; basal three-quarters of femora often lighter
in colour than rest of legs in male. Apices of elytron emarginate just before apico-sutural margin, more
pronounced in female. Setae grey or pale golden. Fully winged.
DIAGNOSIS. Head with frons evenly convex, sometimes somewhat flattened anteriorly, elevation less than
that of post-antennal swellings. Post-antennal swellings distinctly elevated above level of vertex, margins
well defined, lateral margins not contiguous with inner margins of eyes, surface microsculptured and
sparsely punctured. Vertex irregularly punctured throughout with large, shallow, often confluent punc-
tures, derm weakly microsculptured, median 'suture' distinct, setae long and fine. Pronotum transverse, on
average 2-Ox (9), l-8x (cf) broader than long, lateral tooth well developed and broadly rounded, width at
tooth on average l-2x greater than minimum posterior width, depressions shallow and poorly defined,
puncturation irregular, size of punctures similar to those on vertex, punctures on average not more than
half their own width apart, those at sides often confluent and usually larger than those on disc, derm
microsculptured with a weak aeneous lustre, setae fine, mostly confined within margins of punctures.
Scutellum 1-lx longer than broad, apex subtruncate, central area densely punctured with small shallow
punctures, setae extending beyond margins of punctures. Elytron on average 3-8x longer than broad,
width increasing gradually from base of humeral angles to a maximum at around apical quarter, maximum
width l-3x greater than that at apex of scutellum; elytral puncturation irregular due to rugosities,
punctures, in groups, not more than their own width apart; size of elytral punctures less than that of the
smallest punctures on pronotum, punctures on area around scutellum somewhat larger and coarser than
rest, derm shining, raised areas weakly microsculptured. Underside finely and densely punctured
throughout, metepisternum finely granulate, derm microsculptured, setae dense, punctures on posterior
half of metasternum around median groove, 2-0-2-Sx their own width apart, setae almost absent, derm
smooth and shining, rest of surface densely punctured, those at sides less than their own width apart, derm
distinctly microsculptured, setae dense, short and fine; punctures on abdominal segments not more than
their own width apart, derm microsculptured, setae becoming slightly shorter and denser at sides, setae
slightly stouter than those on metasternum; basal three-quarters of femora irregularly punctured with
small shallow punctures up to 3 x their own width apart, apical quarter densely punctured, punctures not
more than their own width apart, microsculpture becoming stronger towards apices, lateral setae long, fine
and adpressed, dorsal setae slightly shorter and stouter; tibiae finely granulate, setae becoming shorter and
stouter towards apices. Genitalia as in Figs 64, 95.
Holotype cf (dissected), Papua New Guinea: Morobe Dist., Wau, 1200 m, 20-26.V.1962 (/. Sedlacek)
(BPBM).
Paratypes. Papua New Guinea: 1 cf, Wau, 1100-1300 m, 2.U966 (/. Sedlacek); 1 cf, Wau, 1200 m,
l.ix.1961, light-trap (/. Sedlacek) (BMNH); 1 cf , no. 10, Purosa Camp, Okapa area, 1950 m, ix-27-1959
(AMNH); 1 $, Owen Stanley Range, Goilala, Tapini, 975 m, 16-25.xi.1957 (W. W. Brandt); 1 $, same
data except Tororo, 1560 m, 15-20.xi.1958; 1 $, Wau, 1050 m, 4.xi.l961 (J. H. Sedlacek) (BMNH); 1 $,
Wau, 1200 m, 15. ix. 1961, light-trap (J. & M. Sedlacek); 1 9, Wau, 1200 m, 4-7. i. 1963, m.v. light-trap (/.
Sedlacek) (BMNH); 1 $, same data except 29.viii.1963; 1 $, Wau, 1200 m, 16-17.viii.1964 (J. Sedlacek);
1 9, Wau, 1200-1300 m,4.ix. 1965 (/. Sedlacek); 1 $, same data except 15. viii. 1965; 1 $, same data except
15.viii.1965; 1 $, same data except l-9.ix.65 (BMNH); 1 $, Wau, 1200 m, 9.iii.l965 (/. Sedlacek); 1 J,
Wau, 1200 m, 24.vii.1965 (/. & M. Sedlacek). (All BPBM unless otherwise stated.)
COMMENTS. P. sedlaceki is representative of the irregularis-group and is related to irregularis and
duplicator.
GALERUCINE BEETLES OF NEW GUINEA 243
Polysastra suavis sp. n.
(Figs 54, 98, 131, Map 7)
GENERAL FORM. Length 9-8-12-0 mm. Elytra pyriform, post-humeral sublateral depressions weak, outer
margin of depression forming an irregular ridge, inner margin indicated by a short sinuate ridge just below
basal quarter, puncturation dense and irregular, setae confined within margins of punctures, 'striaF setae
on interspaces short and erect. Head and pronotum indistinctly punctured. Head deep reddish brown to
black, pronotum and scutellum either concolorous with head or light reddish brown. Elytron dark metallic
green with a weak aeneous lustre, or deep metallic blue with a purplish lustre. Underside and femora either
brownish orange or with metasternites and legs pitchy brown to black. Abdomen testaceous; setae pale
golden. Fully winged.
DIAGNOSIS. Head with frons convex, maximum elevation equal to that of post-antennal swellings,
post-antennal swellings convex, margins distinct, hind and lateral margins oblique, anterior outer angle
contiguous with inner margin of eye, derm smooth and shining, vertex with large, shallow irregular
punctures surrounding a central, slightly raised impuctate area, derm microsculptured, setae long and fine,
confined to punctate areas. Pronotum on average l-8x broader than long, lateral tooth broad and weakly
developed, width at tooth 1-Ox greater than minimum posterior width, lateral depressions shallow but
distinct, median depressions well defined, linked by a narrow longitudinal groove, posterior depression
almost circular, puncturation shallow and indistinct due to vitreous lustre, punctures 1-0-1-5X their own
width apart, derm weakly microsculptured, setae minute and decumbent. Scutellum more or less as broad
as long, apex rounded, minutely punctured, setae adpressed, extending beyond margins of punctures.
Elytron slightly explanate at side, on average 3-lx longer than broad, maximum width l-5x greater than
width at apex of scutellum, apices evenly rounded; basal quarter between humeri and sutural margin,
delimited laterally and posteriorly by shallow depressions, median area of elytron weakly convex;
post-humeral sublateral longitudinal depression shallow, outer margins forming an irregularly developed
ridge from humeri to apical quarter, inner margin only partially indicated by a very short sinuate ridge just
below basal convexity, lateral margins weakly explanate from below humeral angles to just above apical
quarter, punctures 1-2 x their own width apart, shallow and indistinct in places, derm weakly and
irregularly punctured. Underside finely punctured throughout, metepisternum minutely punctured,
punctures 2-5-3-Ox their own width apart, derm microsculptured, setae long and adpressed, adjacent area
of metasternum weakly alutaceus with minute punctures 2-3 x their own width apart, setae long and
decumbent, slightly finer than those on metepisternum; abdomen minutely punctured, punctures 5-6 x
their own width apart, derm weakly microreticulate, setae very fine, length of those in centre of segments
more or less equal to longest on metasternum, those at sides shorter, decumbent or suberect, femora
except for apices minutely punctured, punctures 5-6 x their own width apart, dense at apex, derm strongly
microsculptured at apex, setae long, fine and adpressed, short at apices. Tibiae granulate, setae shorter,
stouter and suberect. Genitalia as in Figs 54, 98.
Holotype cf (dissected), Papua New Guinea: Morobe district, Wau, 1300 m, 22.xii.1961 (/. & J. H.
Sedlacek) (BPBM).
Paratypes. Papua New Guinea: 1 cT, Wau, 1450m, 6.ii.l963 (J. Sedlacek) (BMNH); 1 cf , 1 $, Finisterre
Range, Saidor, Matoko Village, 6-24.ix.1958 (W. W. Brandt) (1 $ BMNH); 2 $ Finisterre Range, Saidor,
Kambavi Village, 1-28. viii. 1958 (W. W. Brandt) (1 $ BMNH); 1 $, Wau, 1450 m, 20.xii.1961 (J. & M.
Sedlacek); I $,MtKaindi, 2350m, 23. iii. 1966, light-trap (/. L. Gressitt);! $, Wau, Kunai Creek, 1270m,
22. viii. 1963 (/. Sedlacek) (BPBM); 1 $ , Finisterre Mts, Budemu, c. 4000ft, 15-24.X.1964 (M. E. Bacchus)
(BMNH).
COMMENTS. P. suavis is representative of the metallica-group. The species included here to
which it bears the closest affinity is metallica (see specific key for distinguishing characters,
p. 225).
Polysastra varia sp. n.
(Figs 58-60, 100, 101, 146-148, Map 6)
GENERAL FORM. Length 8-5-12-0 mm. Elytra subparallel-sided, margins of post-humeral sublateral
longitudinal depressions forming cariniform ridges, puncturation confused, dense throughout, setae
minute, confined within margins of punctures. Head and pronotum with irregular, shallow, distinct
puncturation. Coloration variable, specimens from the Huon Peninsula and the south-east have the elytron
varying from light pinkish brown to deep purple with a deep blue or green submetallic lustre on humeral
244 SHARON L. SHUTE
area, which may extend over whole elytra in darker specimens. Teneral specimens light orange-brown with
a slight green lustre on humeral area; head, pronotum and scutellum testaceous to orange-red, underside
usually pale testaceous; antenna, tibiae and tarsi either concolorous with pronotum or dark reddish brown.
Specimens from the NE. highland areas (Aiyura) have the elytron dark metallic green with dark brown
lateral margins, the abdomen testaceous and the rest of the body black. All specimens seen from north-east
of the Huon region , including Japen I . , have the elytron bright metallic green with a brassy lustre , the head ,
pronotum, scutellum and femora deep orange to reddish orange; underside testaceous to light yellowish
orange, antennal segments and tibiae usually darker in colour than femora. No distinct morphological
differences were noted between the colour forms. Size tends to be very variable within populations.
DIAGNOSIS. Head with frons entirely depressed to weakly concave. Vertex irregularly punctured, punc-
tures large and shallow, confluent in places, average size slightly larger than largest pronotal punctures,
derm surrounding punctures weakly microsculptured, rest smooth and shining, majority of setae confined
within margins of punctures; post-antennal swellings elevated above level of vertex, hind margins
ill-defined, derm almost impunctate. Pronotum transverse, c. 1-7-1-Sx (cf) and c. 1-9-2-Ox ($) broader
than long. Lateral tooth weakly developed, width at tooth c. 0-75 x greater than minimum posterior width;
punctures shallow, concentrated in and around sublateral depressions where they may become confluent,
size variable, largest more or less equal to those on adjacent area of elytron, punctures 1-0-1-2X their own
width apart, derm smooth and shining, weakly microsculptured around punctured areas. Scutellum weakly
convex, punctures minute, 3-4x their own width apart, setae distinct, extending beyond margins of
punctures, derm microreticulate. Elytron on average 3-3 x longer than broad, maximum width at level
more or less e.qual to that of second abdominal segment, width at this point c. 1 -3 x greater than that at level
of apex of scutellum. Outer cariniform margin of post-humeral sublateral longitudinal depression distinct
to apical quarter, inner cariniform margin indistinct until it converges with outer margin at a level more or
less equal to that of apex of metepisternum, continuing till apical quarter where it becomes obsolete, width
between carinae decreasing slightly towards apical quarter, a third, weakly developed admedian carina
sometimes present, particularly in female, running more or less parallel to inner adlateral carina and
becoming obsolete in apical quarter, linked to inner adlateral carina by a short, oblique, irregular
convexity just below convergence of inner and outer carinae. Puncturation of elytron confused, density
more or less uniform throughout, punctures 1-1-1-2X their own width apart, basal third sometimes slightly
more coarsely punctured than rest of derm, particularly around scutellum and on humerus, derm weakly
microsculptured, sometimes few sparse erect setae on intervals. Ventral surfaces minutely punctured
throughout, metasternum convex, punctures 2-0-2-5 x their own width apart, setae long, pale golden,
derm smooth and shining; metepisternum distinctly microsculptured, punctures shallow, distinctly larger
than those on metasternum, c. 0-5-1-Ox their own width apart. Setae similar to those on sides of
metasternum. Punctures on abdomen 2-3 x their own width apart, derm weakly microsculptured, setae
slightly longer than those on disc of metasternum. Legs with basal three-quarters of femora minutely
punctured, punctures 2-0-2-5 x their own width apart, apical quarter slightly more densely and coarsely
punctured, derm microsculptured, setae long and appressed. Tibiae granulate, setae becoming shorter and
denser towards apices. Fully winged. Genitalia as in Figs 58-60, 100, 101.
Holotype cf, Papua New Guinea: Morobe District, Wau, 1200 m, 17.viii.1961, light-trap (/. & M.
Sedlacek) (BPBM).
Paratypes. Papua New Guinea: 1 $ , same data as holotype (BPBM); 12 cf , 9 $ , Morobe District, Wau,
1200-1300 m, 9.vii.-20.xi.l961, light- or Malaise trap (6 cf, 3 $ BMNH); 1 cf, 8 $, Wau, 1200 m,
Il.iii.-10.x.l962, light- or Malaise trap (5 $ BMNH); 6 $, Wau, 1200 m, 21.i.-29.viii.l963, light- or
Malaise trap (all/. & M. Sedlacek); 1 cf, Wau, 1200m, 23.x. 1963, m.v. light-trap (/. L. Gressitt);2tf,l $,
Wau, 1200 m, 3.iv.-16.ix.l964, light-trap (J. & M. Sedlacek) (1 cf , 2 $ BMNH); 1 $, Wau, no. 1187,
4000 ft, 21 .x. 1965 (A. H. Kistner); I $ , Wau, Big Wau Creek, 1200 m, xii. 1965, Malaise trap (J. Sedlacek);
1 Cf , Wau, 1200 m, 22.ii.1966 (/. & M. Sedlacek); 1 £, Wau, 7.iii.l970 (H. Olmus); 1 $, Lae, 17.vii.1970
(H. Olmus) (OIP); 1 $, Karimui, 3.vi.l961, light-trap (/. L. & M. Gressiti); 1 cf, 1 ?, Owen Stanley
Range, Goilala, Loloipa, 11. -20. xii. 1957; 1 <£, Owen Stanley Range, Goilala, Tapini, 975 m, 16-
25. xi. 1957 (all W. W. Brandt); 2 cf, Huon Peninsula, Pindu, 20.iv.1963 (/. Sedlacek); 1 $, Finisterre
Range, Saidor, Sibong Village, 6-16. vi. 1958 (W. W. Brandt); 3 cf, Finisterre Mts, Madant District,
Damanti, 3550 ft, 2-1 1.x. 1964 (M. E. Bacchus) (BMNH); 6 $, Torricelli Mts, Mokai Village, 750 m,
8-15.xii.1958 (2 BMNH); 1 $, Torricelli Mts, Mobitei, 750m, 5-15. iii. 1959 (all W. W. Brandt); 1 cf , 1 $,
Hollandia, 300-600 m, i. 1937 (W. Stuber) (BMNH); 1 $ , Guega, W. of Swart Valley, 1200 m, 15. xi. 1958; 1
Cf, Upper Jimmi Valley, 1000m, 13.vii.1955; 1 $, Upper Jimmi Valley, 1300m, Korop, 12. vii. 1955 (all/.
L. Gressitt); 1 $, 19002, Menyama, 4.U960 (/. H. Ardley); 1 $, 19059 N. Papua, Sangara Estate,
6.viii.l958, at rest on cacoa tree (J. J. H. Szent-Ivany) (DPI); 1 $>, 2219, Afore, Boikik Plantation,
GALERUCINE BEETLES OF NEW GUINEA 245
18.xii.1974 (E. S. C. Smith) ex cardomom (BMHN); 1 cf , Aiyura, 25524, 5400 ft, 21. ii. 1959, regrowth
area; 2 cf, 19043, 19008, Aiyura, 5400 ft, at light (no. 19008 BMNH); 1 cf, 1 $, Aiyura, 6000 ft,
22.vii.1960, u.v. light; 1 $, 19054, Aiyura, i.ix.1960 (all/. H. Barrett) (DPI); 1 cf , Garaina; 1 $, Goroka,
ii.1975, at light (all R. Hornabrook) (RHW); 1 cf , 2 $, Kainantu, 1650 m, 20-26.X.1959, m.v. light (T.
Mad) (1 $ BMNH). (All specimens BPBM unless otherwise stated.)
COMMENTS. P. varia is representative of the vana-group. The species described here to which it
bears the closest affinity is venusta; superficially it is very similar topurpurasco. It appears to be
one of the most variable species examined and exhibits geographical colour variation.
PLANT ASSOCIATES. Theobroma cacao, Elettaria cardamomum, Coffeaarabica.
Polysastra venusta sp. n.
(Figs 61, 127, Map 5)
GENERAL FORM. Length 8-0-10-0 mm. Elytra subparallel-sided, post-humeral sublateral longitudinal
depression present, outer margins forming distinct ridges which are joined below humeri at a point level
with middle of metasternum, outer ridge distinct from base of elytron to apical quarter, inner ridge
becoming obsolete just above termination of outer ridge in apical quarter, a third longitudinal ridge is
weakly indicated parallel to inner ridge, lateral separation equal to space occupied by five punctures,
surface densely and regularly punctured throughout, pronotum almost impunctate, punctures shallow and
ill-defined due to vitreous surface. Head testaceous to bright purplish red, pronotum either concolorous
with head or slightly darker; elytron deep yellowish orange to purplish red, humeral area with a strong,
dull, dark green submetallic lustre which extends posteriorly over sublateral depression in purplish
specimens. Underside testaceous to orange, legs either entirely testaceous to yellowish brown or with
tibiae and tarsi pitchy brown. Majority of elytral setae confined within margins of punctures, interspersed
with a few long, erect setae on interspaces. Fully winged.
DIAGNOSIS. Head with frons completely depressed anteriorly. Post-antennal swellings distinct from vertex,
posterior margins rounded, anterior outer angle contiguous with inner margin of eye; vertex irregularly
punctured with moderately large shallow punctures, majority situated around middle of vertex, derm
weakly microsculptured, setae very fine and adpressed, median 'suture' distinct. Pronotum transverse, on
average l-9x broader than long, lateral tooth not well developed, apex subacute, width at tooth 1-lx
greater than minimum posterior width, depressions shallow and ill-defined, puncturation sparse, indistinct
due to vitreous surface of derm, punctures shallow, similar in size to those on vertex, majority situated
along anterior margin, setae very small and fine. Scutellum triangular, length more or less equal to
maximum width, derm weakly microsculptured, punctures minute, setae short and fine. Elytron on
average 3-3x longer than broad, width increasing gradually from base of humeral angles, reaching a
maximum in apical quarter, maximum width l-3x greater than that at apex of scutellum, puncturation
more or less uniform throughout, punctures not more than their own width apart, slightly coarser in basal
quarter, derm microsculptured. Underside finely and densely punctured throughout, punctures on
metasternum 1-0-1-5X their own width apart, derm smooth and shining, becoming slightly microsculp-
tured at sides, metepisternum distinctly microsculptured, puncturation tending to become granulate, setae
fine and decumbent, punctures on abdominal segments minute, 2-0-2-5X their own width apart, derm
shining, very finely microsculptured, setae very fine, slightly longer than those on metasternum, particular-
ly towards middle of segments. Femora densely and finely punctured throughout, punctures not more than
their own width apart, setae long, fine and adpressed, tibiae finely granulate, setae dense, stouter and more
erect than on femora, density increasing towards apices. Genitalia as in Fig. 61.
Holotype cf (dissected), Papua New Guinea: Torricelli Mts, Wantipi, vill[age], 30.xi-8.xii.1958 (W. W.
Brandt) (BPBM).
Paratypes. Papua New Guinea: 2 cf , same data as holotype; 1 cf , Kumur, Upper Jimmi Vfalley], 1000 m,
13.vii.1955 (J. L. Gressitt); 1 cf , 1 $, Humboldt Bay, Bewani Mts, 400 m, vii.1937 (W. Stiiber) (BMNH).
COMMENTS. P. venusta is representative of the varia-group. The species described here to which
it bears the closest affinity is varia.
SASTRA Baly
(Figs 1, 15, 42-49, 77-82, Maps 1,9)
Sastra Baly, 1865: 253; Chapuis, 1875: 198, 206; Maulik, 1936: 254 (description based on species from Asia,
246 SHARON L. SHUTE
transferred here to other genera); Gressitt & Kimoto, 1963: 404 (description based on single species
incorrectly listed as Chinese due to data error). Type-species: Sastra placida Baly, by original designa-
tion.
Eriosardella Chujo, 1935: 219. Type-species: Eriosardella costata Chujo, by original designation. [Synony-
mized by Gressitt & Kimoto, 1963: 404.]
GENERAL FORM. Body elongate, elytra subparallel-sided (Fig. 1). Dorsum setose, elytron with sparse or
dense, distinct setae throughout. Elytron more or less evenly convex, post-humeral sublateral longitudinal
depression shallow to absent, derm more or less uniformly and densely punctured throughout. Length of
antenna equal to at least three-quarters of body length, third segment at least l-4x length of fourth
segment and 2-Ox that of second, all segments filiform, similar in both sexes. Eyes large and protuberant.
Frons with sides distinctly convex, median area with a shallow depression or groove. Gena emarginate to
eye. Pronotum either almost as broad as long or distinctly transverse, lateral margins evenly rounded in
anterior half, maximum width not exceeding that of head plus eyes, distinctly less than that of elytra at
humeri, primary setal pores distinctly tuberculate, dorsal depressions present, sublateral depressions
shallow to well defined, each depression occupying at least two-thirds of area between disc and lateral
margin, disc with a small anterior and posterior depression, anterior depression largest and usually better
defined. Derm either with a vitreous reflection or dull without lustre, elytron very rarely metallic,
sometimes a very weak submetallic lustre on humeral area in some species. Elytral epipleura present to at
least apical quarter. Underside setose and finely punctured throughout. Fore coxal cavities open behind.
Apex of abdomen in male with a deep triangular incision, evenly rounded in female. Legs slender, similar
in both sexes, apices of tibiae with a comb-like row of short spines on the lateral edges, ventral spurs absent.
Basal segment of fore tarsi in male slightly more enlarged than mid and hind, basal segment of fore and mid
tarsi slightly shorter than length of segments 2 and 3 combined, basal segment of hind tarsus slightly longer
than segments 2 and 3 combined. Claws distinctly bifid. Fully winged.
DIAGNOSIS. Head with gena emarginate medially, emargination reaching anterior margin of eye (Fig. 15).
Apical segment of maxillary palp conical , length slightly greater than that of preceding segment. Mandibles
large and distinctly toothed. Labrum small and rounded anteriorly, not obscuring mandibles from above,
width less than that of frons. Pseudoclypeus distinct, length not more than half that of labrum. Frons with
sides distinctly convex, median area with a triangular depression or shallow groove extending from
interantennal area, length not more than l-5x that of antennal socket, anterior margin level with that of
eyes. Eyes large and protuberant, length at least 3-5 x that of antennal socket. Antennal sockets separated
by less than width of a socket. Post-antennal swellings distinctly convex and elevated above level of vertex,
confluent with frons between antennal sockets and delimited behind by a narrow horizontal groove. Vertex
with a distinct coronal 'suture', derm either sparsely and indistinctly punctured or with an area of dense to
confluent puncturation behind post-antennal swellings. Pronotum varying from almost as broad as long to
distinctly transverse, maximum width and convexity in anterior half, lateral margins of anterior two-thirds
evenly rounded, becoming straighter as pronotum narrows in posterior third, posterior margin sinuate,
degree and density of puncturation variable, setae either sparse and indistinct or long and moderately
dense, size and development of dorsal depressions variable. Elytron on average 3-4x longer than broad,
surface more or less evenly convex (some species may have a very shallow transverse depression just
behind humeral area and just above apical quarter) , apices of elytra evenly rounded or with an apicosutural
projection, maximum width in apical quarter. Underside minutely and densely punctured throughout,
setae long and distinct, extending well beyond margins of punctures, metasternum convex with a narrow,
shallow longitudinal groove, length at least 2-5x greater than that of mesosternum. Legs slender, length of
hind leg plus tarsus slightly greater than that of elytron, fore and mid legs c. 1-5 mm shorter than hind leg,
femora more or less of equal width, tibiae more or less straight, no sexual dimorphism, puncturation
tending to become granulate towards apices, setae short and stout. Position in abdomen, and structure of
male and female genitalia similar to that of Polysastra. Male aedeagus taking the form of a narrow tube,
either with a spatulate apical region with a broad median orifice, or of fairly uniform width with a small
median orifice and acute apex. Female genitalia very similar in structure to those of Polysastra, styli of
similar, short rounded type.
DISCUSSION. The present study shows that only nine of the 35 species included in Sastra by
Wilcox (1971) were correctly assigned: these are listed below with their type-data. The
remaining species that have not been assigned to new genera in this paper are listed (p. 256) with
their new generic combinations.
Very little is known about the biology of Sastra. Specimens have been collected at various
forms of light, including black light. An undescribed north Australian species was beaten from
GALERUCINE BEETLES OF NEW GUINEA 247
dead branches by Dr G. B. Monteith during the day, and another undescribed species from
Afore, Papua New Guinea was collected on Cardamom. Apart from these records no other
bionomic data are known for this genus.
All the species remaining in Sastra are from the New Guinea region. In addition to the nine
listed below I have examined more than 34 undescribed species, all from the same area (Map 1),
indicating that the genus is restricted to the region. 5. costata Chujo, described from a single
specimen which bears the locality 'China', appears to have been incorrectly labelled as all other
specimens examined of this species are from Irian Jaya. Enquiries and examination of Chinese
Galerucinae material has not revealed further alleged specimens of this species from China.
Species remaining in Sastra
Sastra beccarii Jacoby
(Figs 49, 82, 110, Map 9)
Sastra beccarii Jacoby, 1886: 76. Holotype 9, IRIAN JAVA: Hatam, vi.1875 (Beccari) (MIZSU) [examined].
ADDITIONAL MATERIAL EXAMINED
Irian Jaya: 1 $,Etnabaai, 1904; 1 $, Heuvel Bivak, 750 m,xi.[19]09(Lor£>rtfz) (ITZ); 1 9, Central Mts,
Archbold Lake, 760 m, 26.xi-3.xii. 1961 (BPBM).
Japen Island: 1 cf , R. Manai-Undei, 500 ft, x.1938 (L. E. Cheesman); 2 9, Japen Camp 2, Mt Eiori,
2000ft, ix.1938 (L. E. Cheesman) (BMNH).
Sastra costata (Chujo)
(Figs 43, 78, 105, 106, Map 9)
Eriosardella costata Chujo, 1935: 219. Holotype $, 'China': (Kraatz) (IP) [examined].
Sastra costata (Chujo) Gressitt & Kimoto, 1963: 404 [genera synonymized].
ADDITIONAL MATERIAL EXAMINED
New Guinea: 1 cf, N[ew Guinea] (Wallace); 1 9, New Guinea; 1 $, 1 9 (Wallace); 1 9 (no data)
(BMNH); 1 Cf , 1 9, Ramoi, 'ging 12' (L. M. D'Albertis); 1 $, same data plus iv.[18]73; 1 $ , New Guin[ea]
(MCZ).
Sastra depressa Weise
(Figs 46, 79, 107, Map 9)
Sastra depressa Weise, 1917: 208. LECTOTYPE $, IRIAN JAYA: Bivak Elland, i.[19]10 (Lorentz) (ITZ),
here designated [examined].
ADDITIONAL MATERIAL EXAMINED
Irian Jaya: 4 9 (paralectotypes), same data as lectotype (ITZ, MNHU) (1 9 without date); 1 cf , 3 $
(paralectotypes), Noord River, ix.[19]09 (Lorentz) (ITZ, MNHU); 1 9, Noord Riv[er], 8. v. 1907
(Lorentz) (ITZ). Papua New Guinea: 1 9, Fly River, x.ii.[18]95 (L. M. D'Albertis); 1 $, Paumomu
Riv[er], ix-xii.[18]92 (Loria) (MCZ); 7 9, Karimui, 1080 m, 10-13. vii. 1963 (/. Sedlacek); 1 9, same data
except l.v.1969; 1 9, Fly River, Oslobip, 100-600 m, viii.1969 (/. Sedlacek) (BPBM).
Sastra elegans Weise
(Figs 45, 104, Map 9)
Sastra elegans Weise, 1912: 437. Holotype cf , IRIAN JAYA: Bivak Elland, x.[19]09 (Lorentz) (ITZ).
Sastra limbata Baly
(Figs 44, 111, 112, Map 9)
Sastra limbata Baly, 1865: 254. LECTOTYPE cf , NEW GUINEA (Wallace) (BMNH), here designated
[examined].
ADDITIONAL MATERIAL EXAMINED
New Guinea: 2 cf (paralectotypes), same data as lectotype (BMNH).
248
SHARON L. SHUTE
ft
43
44
u
Figs 42-53 Sastra and Polysastra, aedeagi, dorsal and lateral view. 42, Sastra olivacea. 43, 5. costata. 44,
5. limbata. 45, S. elegans. 46, 5. depressa. 47, 5. viridipennis. 48, 5. rugicollis. 49, 5. beccarii. 50,
Polysastra abdominalis. 51, P. explanata. 52, P. micropunctata. 53, P. montana.
GALERUCINE BEETLES OF NEW GUINEA
249
VJ
62
Figs 54-65 Polysastra, aedeagi, dorsal and lateral view. 54, P. sauvis. 55, P. metallica (SE. form). 56,
metallica (NW. form). 57, P. metallica (NE. form) (Map 7). 58, P. wzr/a (P.N.G. form). 59, P. wm'a
(NW.-NE. form). 60, P. var/a (green highland form). 61, P. venusta. 62, P. bicostata. 63, P. inhabilis.
64, P. sedlaceki. 65, P. duplicator.
250
SHARON L. SHUTE
66
\J
74
\J
Figs 66-76 Polysastra, aedeagi, dorsal and lateral view. 66, P. irregularis (western form) (Map 5). 67, P.
irregularis (eastern form). 68, P. confusa. 69, P. fuscitarsis. 70, P. laetabilis. 71, P. obscuricornis. 72, P.
costatipennis. 73, P. purpurasco purpurasco. 74, P. purpurasco viridis. 75, P. helleri (NW. form). 76, P.
helleri (NE. form) (Map 4).
GALERUCINE BEETLES OF NEW GUINEA
Figs 77-101 Sastra and Polysastra, female spermathecae. 77, Sastra placida. 78, S. costata. 79, 5.
depressa. 80, 5. olivacea. 81, S. rugicollis. 82, 5. beccarii. 83, Polysastra abdominalis. 84, P. explanata.
85, P. micropunctata. 86, P. montana. 87, P. obscuricornis. 88, P. costatipennis. 89, P. helleri. 90, P.
purpurasco purpurasco. 91, P. purpurasco viridis. 92, P. fuscitarsis. 93, P. irregularis. 94, P. duplicator.
95, P. sedlaceki. 96, P. confusa. 97, P. inhabilis. 98, P. swavw. 99, P. metallica. 100, P. var/a (typical
form). 101, P. von'fl (green highland form).
252 SHARON L. SHUTE
Sastra olivacea Jacoby
(Figs 42, 80, 109, Map 9)
Sastra olivacea Jacoby, 1904: 503. LECTOTYPE cf, D'ENTRECASTEAUX ISLANDS: Fergusson Island
(MCZ), here designated [examined].
ADDITIONAL MATERIAL EXAMINED
D'Entrecasteaux Islands: 1 d", 2 9 (paralectotypes), same data as lectotype (BMNH, MCZ). Trobriand
Islands: 1 cf,l $, Kriwini Island, iii.v.[18]95 (A. S. Meek) (MCZ).
Sastra placida Baly sp. rev.
(Figs 77, 102, Map 9)
/ Sastra placida Baly, Ijf65: 254. Holotype $, MYSOL: (Wallace) (BMNH) [examined; abdomen missing].
[Synonymised with Sastra viridipennis Boisduval by Weise, 1917: 208.]
ADDITIONAL MATERIAL EXAMINED
Mysol: 1 9 (Wallace) (BMNH); 1 $, 'Malaya pen.' (MCZ).
Sastra rugicollis Jacoby
(Figs 48, 81, 108, Map 9)
Sastra rugicollis Jacoby, 1904: 502. LECTOTYPE $, TROBRIAND ISLANDS: (Loria) (BMNH), here
designated [examined].
ADDITIONAL MATERIAL EXAMINED
Trobriand Island: 1 $ (paralectotype), same data as lectotype (BMNH). D'Entrecasteaux Islands: 2 cf ,
2 9, Ferguson Island, ix, x, xi, xii.[18]94 (A. S. Meek) (MCZ).
Sastra viridipennis (Boisduval)
(Figs 47, 103, Map 9)
Galleruca viridipennis Boisduval, 1835: 559. Holotype $ , NEW GUINEA (D. Lesson) (MIZSU) [examined].
ADDITIONAL MATERIAL EXAMINED
Irian Jaya: 1 cf , Dorey (Wallace); 4 9 , Dorey (BMNH, 1 MCZ) 2 cf , Andai, 1892 (W. Doherty) (MCZ).
Nomen dubium
Sastra suturalis Jacoby
(Fig. 113)
Sastra suturalis Jacoby, 1886: 75. ?Type 9, AUSTRALIA (MCZ) [examined].
ADDITIONAL MATERIAL EXAMINED
Australia: 2 9, Cooktown (BMNH).
COMMENTS. 5. suturalis was described from Somerset, N. Queensland (L. M. D'Albertis).
However the MCZ type-specimen labelled as being from the Jacoby collection bears no locality
label or labels consistent with a D'Albertis specimen, and some uncertainty remains concerning
its status.
This species is provisionally retained in Sastra as it appears to belong to an undescribed genus
near Gallerucella. No additional species congeneric with suturalis are known so it is not possible
to establish the range of this genus.
GALERUCINE BEETLES OF NEW GUINEA
253
KEY SYMBOLS
bicostata
fuse itarsis
A laetabilis
*purpurasco
purpurasco
**costatipennis
'purpurasco viridis
• inhabilis
explanata
• duplicator
Tconfusa
cropunctata
helleri
• abdominalis
Maps 2-4 Distribution of Polysastra species.
254
SHARON L. SHUTE
KEY SYMBOLS
T venusta
' • irregularis
i metallica
"frsedlaceki
Maps 5-7 Distribution of Polysastra species.
GALERUCINE BEETLES OF NEW GUINEA
255
KEY SYMBOLS
* Polysastra
obscuricornis
Marmina sp. B
mina basalis
Marmina sp.A
Marmina
quadripustulata
9 Dreeus distinctus
Maps 8-10 Distribution of Polysastra, Sastra, Marmina and Dreeus species.
256 SHARON L. SHUTE
Species removed from Sastra
Coelocrania rubya (Maulik) comb. n.
Sastra rubya Maulik, 1936: 256. Holotype $, BURMA: Ruby mines (Doherty) (BMNH) [examined].
Galerucella ceylonensis Jacoby comb. rev.
Galerucella ceylonensis Jacoby, 1887: 105: 105. Syntypes, SRI LANKA: 1 cf, Dikoya, 3800-4200 ft,
6.xii.[18]81-16.i.[18]82 (G. Lewis); 1 $, (G. Lewis); (BMNH) [examined].
Sastra ceylonensis (Jacoby) Maulik, 1936: 261.
Galerucella lateralis iacoby comb. rev.
Galerucella lateralis Jacoby, 1887: 106. Holotype $, SRI LANKA: (G. Lewis) (BMNH) [examined].
Sastra lateralis (Jacoby) Maulik, 1936: 258.
Galerucella marginata Jacoby comb. rev.
Galerucella marginata Jacoby, 1887: 107. Holotype $, SRI LANKA: Bogawantalawa, 4900-5200 ft,
21.iii.-4.iv.[18]82 (G. Lewis) (BMNH) [examined].
Sastra marginata (Jacoby) Maulik, 1936: 260.
Galerucella rugosa (Jacoby) comb. n.
Sastra rugosa Jacoby, 1886: 71. Type cf , SUMATRA: Singkara, x.1878 (Beccari) (MCZ) [examined].
Galerumaea ffavomarginata (Jacoby) comb. n.
Sastra flavomarginata Jacoby, 1886: 74. Holotype <j>, NEW GUINEA: Fly River (L. M. D'Albertis) (MCSN)
[examined].
Motnaea fasciata (Jacoby) comb. n.
Sastra fosciata Jacoby, 1886: 77. Syntype $, NEW GUINEA: Ramoi, ii.1875 (Beccari) (MCZ) [examined].
Momaea meijerei (Weise) comb. n.
Sastra meijerei Weise, 1908: 320. Syntype $ , NEW GUINEA: Jamur (MNHU) [examined].
Sastracella fulvicornis (Jacoby) comb. n.
Sastra fulvicornis Jacoby, 1892: 958. Holotype $, BURMA: Karen Mts (Fea) (MCSN) [examined].
Sastracella harmandi (Laboissiere) comb. n.
Sastra harmandi Laboissiere, 1932: 961. Holotype cf , INDIA: Sikkim (Harmand) (MNHN) [examined].
Sastroides acutipennis (Laboissiere) comb. n.
Sastra acutipennis Laboissiere, 1932: 960. Syntype cf , INDIA: Sikkim (Harmand) (MNHN) [examined].
Sastroides hirtipennis (Jacoby) comb. n.
Sastra hirtipennis Jacoby, 1891: 33. Holotype $, INDIA: Assam (BMNH) [examined].
Sastroides metallescens (Jacoby) comb. n.
Sastra metallescens Jacoby, 1894: 304. Holotype $, BURMA: Martapura, 1891 (Doherty) (MCZ) [ex-
amined].
Sastroides purpurascens (Hope) comb. rev.
Galleruca purpurascens Hope, 1831: 29. Holotype cf , INDIA (BMNH) [examined].
GALERUCINE BEETLES OF NEW GUINEA
257
Gastroides [sic] purpurascens (Hope) Bryant, 1923: 146.
Momaea purpurascens (Hope) Weise, 1924: 69.
Sastra purpurascens (Hope) Maulik, 1936: 267.
Sastroides purpurascens (Hope) Bryant, 1937: 101.
Sastra purpurascens (Hope) Wilcox, 1971: 52.
Yulenia discoidalis (Baly) comb. n.
Sastra discoidalis Baly, 1886: 35. Syntypes 1 cf, 1 $, BORNEO: Sar[awak] (Wallace); 1 <J>, MALAYA:
Sing[apore] (Wallace) (BMNH) [examined].
Check list of genera, species and subspecies treated here
DREEUSgen. n.
distinctussp. n.
MARMINA gen. n.
basalis (Jacoby) comb. n.
quadripustulata (Jacoby) comb. n.
POLYSASTRA gen. n.
abdominalis (Jacoby) comb. n.
bicostata (Jacoby) comb. n.
confusa sp. n.
costatipennis (Jacoby) comb. n.
duplicator sp. n.
explanata sp. n.
fuscitarsissp. n.
helleri (Weise) comb. n.
in ha bills sp. n.
irregularissp. n.
kampeni (Weise) comb. n.
laetabilis (Weise) comb. n.
metallica (Jacoby) comb. n.
andaiensis (Jacoby) syn. n.
micropunctata sp. n.
montana sp. n.
obscuricornis (Blackburn) comb. n.
purpurasco sp. n.
purpurascopurpurascosubsp. n.
purpurasco viridis subsp. n.
rugulosa (Weise) comb. n.
sedlacekisp. n.
suavis sp. n.
varia sp. n.
venusta sp. n.
SASTRA Baly
beccarii Jacoby
costata Chujo
depressa Weise
elegans Weise
limbata Baly
olivacea Jacoby
placida Baly sp. rev.
rugicollis Jacoby
viridipennis (Boisduval)
Nomen dubium
suturalis Jacoby
References
Allard, E. 1888. Synopsis des Galerucines a corselet sillonne transversalement. Annales de la Societe
Entomologique de France (6) 8: 305-332.
Baly, J. S. 1865. Descriptions of new genera and species of Galerucidae. Annals and Magazine of Natural
History (3) 16: 247-255.
1886. Descriptions of new genera and species of Galerucidae. Transactions of the Entomological
Society of London 1886: 27-39.
Blackburn, T. 1896. Further notes on the Australian Coleoptera with descriptions of new genera and
species. Transactions and Proceedings and Report of the Royal Society of South Australia 20: 35-109.
Bryant, G. E. 1923. Notes on synonymy in the Phytophaga (Coleoptera). Annals and Magazine of Natural
History (9) 12: 130-147.
1937. Notes on Synonymy in the Phytophaga (Coleoptera). Annals and Magazine of Natural History
(10)20:97-101.
Boisduval, J. B. A. de 1835. Voyage de decouvertes de L' Astrolabe faune Entomologique du L'Ocean
Pacifique, Part II, Coleopteres. 608 pp. Paris.
Chujo, M. 1935. Descriptions of a new genus and two new species of Chrysomelidae (Col.). Arbeiten iiber
morphologische und taxonomische Entomologie aus Berlin-Dahlem 2: 219-220.
Gressitt, J. L. & Kimoto, S. 1961. The Chrysomelidae of China and Korea. Pacific Insects Monograph
Parts 1,2, 1A, IB.
Gressitt, J. L. & Hornabrook, R. W. 1977. Handbook of the common New Guinea Beetles. Wau Ecology
Institute, Handbook no. 2. 87 pp.
258 SHARON L. SHUTE
Hope, F. W. 1831. Synopsis of the new species of Nepaul insects in the collection of Major General
Hardwicke, pp. 21-32. In Gray, J. E., The Zoological Miscellany . London.
Jacoby, M. 1886. Description of new genera and species of phytophagous Coleoptera from the Indo-
Malayan and Austro-Malayan sub-regions, contained in the Genoa Civic Museum. Annali del Museo
Civico di Storia Naturale. 4: 41-128.
- 1887. Descriptions of the phytophagous Coleoptera of Ceylon obtained by Mr. George Lewis during
the years 1881-1882. Proceedings of the Zoological Society of London 1887: 65-118.
- 1889. List of the phytophagous Coleoptera obtained by Signori Fea at Burmah and Tenasserim, with
descriptions of the new species. Annali del Museo Civico di Storia Naturale 27: 147-237.
- 1892. Descriptions of the new genera and species of the phytophagous Coleoptera obtained by Sign.
L. Fea in Burma. Annali del Museo Civico di Storia Naturale 32: 869-999.
- 1894. Descriptions of new genera and species of phytophagous Coleoptera obtained by W. Doherty
in the Malayan archipelago. Novitates Zoologicae 1: 267-330.
1899. Descriptions of the new species of phytophagous Coleoptera obtained by Dr. Dohrn in
Sumatra. Entomologische Zeitung, Stettin 60: 259-364.
1904. Descriptions of new genera and species of phytophagous Coleoptera obtained by Dr. Loria in
New Guinea. Annali del Museo di Storia Naturale 41: 498-504.
Knuth, P. 1909. Handbook of flower pollination 3: 545-546. Oxford.
Laboissiere, M. V. 1932. Galerucini de la collection du Museum National D'Histoire Naturelle, recueillis
dans L'Himalaya par le Dr. J. Harmand. Bulletin du Museum National d'Histoire Naturelle 4: 960-970.
Maulik, S. 1936. The Fauna of British India including Ceylon and Burma. Coleoptera, Chrysomelidae,
Galerucinae, pp. 254-268. London.
— Manuscript (on cards). Alphabetical list of plants attacked by Chrysomelid beetles. BMNH, London.
Sharp, D. & Muir, F. 1912. The comparative anatomy of the male genitalia tube in Coleoptera.
Transactions of the Entomological Society of London 1912: 477-642.
Varmer, K. K. 1969. Functional and developmental organs in the male of Galerucella birmanica Jacoby
(Coleoptera: Chrysomelidae). Annales des Sciences Naturelles (Zoologie) 11: 139-234.
Weise, J. 1908. Resultats de 1'expedition scientifique Nederlandaise a la Nouvelle-Guinee en 1903.
Chrysomelidae. Nova Guinea 5: 311-350.
1912. Chrysomeliden und Coccinelliden. Nova Guinea 9: 437-438.
1917. Chrysomeliden und Coccinelliden aus Nord Neu-Guinea. Tijdschrift voor Entomologie 60:
192-224.
1922. Chrysomeliden und Coccinelliden aus Queensland. Arkivfur Zoologi 15 (12): 1-150.
Wilcox, J. A. 1971. Coleopterorum Catalogus, supplementa Pars 78, Chrysomelidae: Galerucinae.
Gravenhage.
GALERUCINE BEETLES OF NEW GUINEA
259
Figs 102-110 Sastra. 102, S. placida $, 6-5 mm. 103, 5. viridipennis O", 7-5 mm. 104, 5. elegans cf,
9-1 mm. 105, S. costata $,8-1 mm. 106, 5. costata, showing emargination of epipleura. 107, 5. depressa
$, 10-1 mm. 108, 5. rugicollis $,9-1 mm. 109, 5. olivacea $,8-9 mm. 110, 5. beccarii $, 8-5 mm.
260
SHARON L. SHUTE
111
114
Figs 111-119 Sastra, genus A and Marmina. Ill, 112, Sastra limbata $?, 8-0 mm, showing two colour
forms. 113, ?S. suturalis, 6-5 mm. 114, male of genus A. 115, female of genus A. 116, Marmina
quadripustulata 9 , 10-1 mm. 117, M. basalis $ , 9-1 mm. 118, Marmina sp. A cf , 9-0 mm. 119, Marmina
sp. B C?, 7-5 mm.
GALERUCINE BEETLES OF NEW GUINEA
261
122
123
Figs 120-128 Polysastra. 120, P. costatipennis d", 7-1 mm. 121, P. obscuricornis cf, 8-5 mm. 122, P.
helleri $, 9-5 mm. 123, P. laetabilis $, 10-5 mm. 124, P. purpurasco purpurasco $, 11-0 mm. 125, P.
purpurasco viridis $, 10-1 mm. 126, P. fuscitarsis £, 12-0 mm. 127, P. venusta & , 8-5 mm. 128, P.
explanata 9, 10-5 mm.
262
SHARON L. SHUTE
Figs 129-137 Polysastra. 129, P. montana $ , 12-5 mm. 130, P. micropunctata $ , 11-2 mm. 131, P. sauvis
$,11-5 mm. 132, P. metallica $>, 12-5 mm. 133, P. sp. $, metallica subgroup 2, 11 -2 mm. 134. P. sp. $,
metallica subgroup 3, 10-2 mm. 135, P. duplicator $, 10-5 mm. 136, P. irregularis £, 10-2 mm. 137, P.
sedlaceki $, 13-2 mm.
GALERUCINE BEETLES OF NEW GUINEA
263
138
139
140
141
142
145
146
Figs 138-146 Polysastra. 138, f. confusa cf , 7-8 mm. 139, P. sp. cf near confusa, 8-0 mm. 140, P.
rugulosa $ , 8-5 mm. 141 , P. sp. $ , Irregularls subgroup 4, 8-1 mm. 142, P. sp. $ , irregularis subgroup 5,
10-3mm. 143, P. inhabilisQ, ll-2mm. 144, P. abdominalis $, 10-5 mm (pale form). 145, P. abdominalis
$, 10-3 mm (dark form). 146, P. varia $,11-3 mm (typical form).
264
SHARON L. SHUTE
150
Figs 147-150 Polysastra. 147, P. varia $ , 12-0 mm (NW.-NE. metallic form). 148, P. varia $ , 12-2 mm
(NE. highland form). 149, P. bicostata $,11-0 mm. 150, P. sp. $, irregularis subgroup 4, 13-5 mm.
Acalypha sp. 241
Coffeasp. 241,245
Curcuma sp. 241
Elettariasp. 245,247
Eugeni sp. 241
Plant index
Laportea sp. 239
Pipturus sp. 241
Theobroma sp. 245
abdominalis 230, 232, 257
abdominalis-group 225-226
acutipennis 256
Alopena 207
andaiensis 238, 257
Anoides 209
Apophylia 212
Astena 212
Astridella 208
Atysa210,211
Aulacophora 210
basalis 220, 257
beccarii 247, 257
bicostata 229, 232, 257
bicostata-group 222, 226
Buphonida 209
Cassena 212
ceylonensis 256
Cneorane 212
Coelocrania 207, 256
confusa 230, 232-233, 257
costata 247, 257
costatipennis 231, 233, 239, 257
costatipennis-group 225, 226, 232
depressa 247, 257
Diaphaenidea211
discoidalis 257
distinctus 216-217, 257
Dreeus 213, 215-217, 257
duplicator 230, 233, 237, 242, 257
Dysiodes 210
elegans 247, 257
explanata 229, 233-234, 257
explanata-group 225, 226
fasciata 256
flavomarginata 256
fulvicornis 256
fuscitarsis 231, 234-235, 240, 257
Galerucella 210, 256
Galerumaea 210, 256
Gronovius213,238
harmandi 256
helleri 232, 235, 257
hirtipennis 256
Hoplasoma211
inhabilis 230, 235-236, 257
inhabilis-group 225, 226
irregularis 230, 233, 236-237, 242, 257
irregularis-group 225, 227
GALERUCINE BEETLES OF NEW GUINEA
Index
Iaetabilis231,235,237,257
lateralis 256
limbata 247, 257
Lomirana208,211,212
Luperus 207, 208
marginata 256
Marmina 213, 217-220, 257
Marmina species A 219, 220
Marmina species B 220
meijerei 256
Menippus 209
metallescens 256
metallica 231, 238, 243, 257
metallica-group 225, 227-228
Microlepta 208
micropunctata 229, 238-239, 257
micropunctata-group 225, 228-229
Momaea 210, 21 1,256
Monolepta 212
montana 229, 238, 239, 257
Morokasia 209
265
Neodrana 208
Neolepta 207, 212
Notonicea209,210
obscuricornis 231, 233, 239, 240, 257
Oides 209
olivacea 252, 257
Palaeosastra 207
Palpaenidea211
Papuania 210
Paranoides 209
Paridea 212
Paumomua211
placida 252, 257
Phyllocleptis 212
Pleronexis 209
Plesistia211
Polysastra 213, 220-247, 257
Poneridea 21 1,212
Prasyptera 208
purpurascens 256
purpurasco 232, 235, 237, 240-241, 245, 257
quadripustulata 217, 219, 220, 257
rubya 256
rugicollis 252, 257
rugosa 256
rugulosa230,241,257
Sastra 210, 245-247, 252, 257
Sastracella 256
Sastroides 256
266 SHARON L. SHUTE
sedlaceki 230, 242, 257 varia-group 225, 229
suavis 231 , 243, 257 venusta 230, 245, 257
suturalis 252, 257 viridipennis 252, 257
Synodita212 viridis231, 241, 257
varia 230, 243-245, 257 Yulenia 208, 257
British Museum (Natural History)
Blue butterflies of the Lycaenopsis-group
J. N. Eliot and A. Kawazoe
A definitive taxonomic study and revision of the Lycaenopsis-group of the Polyommatini,
butterflies that occur throughout North America, Britain and northern Europe, Asia and the
Orient to Japan, New Guinea and the Philippines.
This book will be a 'must' for lepidopterists interested in the Lycaenidae of the Holarctic and
Oriental regions. The 560 figures include six pages of colour illustrating unique specimens and
rarer species while the drawings of genitalia are of exceptional quality. There are keys and
descriptions for identification to subspecific level, chapters on variation and a checklist.
296pp, 6pp colour illustrations, 560 figs
ISBN 0 565 00860 9
Hardback £28.00
Published by the British Museum (Natural History),
Cromwell Road, London SW7 5BD. May 1983.
Titles to be published in Volume 46
The generic and tribal classification of spore-feeding Thysanoptera (Phlaeothripidae: Idolo-
thripinae).
By L. A. Mound & J. M. Palmer.
A revision of the Afrotropical mole-crickets (Orthoptera: Gryllotalpidae).
By B.C. Townsend.
Key to the genera of galerucine beetles of New Guinea, with a review of Sastra and related new
taxa (Chrysomelidae).
By Sharon L. Shute.
The Afrotropical dacetine ants (Formicidae).
By Barry Bolton.
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd., Dorchester
GENERAL
Bulletin of the * =
British Museum (Natural History)
LIBRARY
The Afrotrooical dacetine ants
Barry Bolton
Entomology series
Vol 46 No 4 25 August 1983
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,
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2 5 AUG1983
T 1
World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1983
The Entomology series is produced under the general editorship of the
Keeper of Entomology: Laurence A. Mound
Assistant Editor: W. Gerald Tremewan
ISSN 0524-6431 Entomology series
Vol 46 No 4 pp 267-416
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 August 1983
/ y\>w uc(
The Afrotropical dacetine ants (Formicidae)
Barry Bolton i
LIBRARY -V
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis 267
Introduction 267
Measurements and indices 271
Abbreviations of depositories 272
Diagnosis of Afrotropical dacetine ants 272
Key to Afrotropical dacetine genera (workers) 273
Smithistruma Brown 274
Key to species (workers) 276
Key to species-groups (workers) 280
Trichoscapa Emery 319
Glamyromyrmex Wheeler 320
Key to species (workers) ? 321
Serrastruma Brown 335
Key to species (workers) 337
Cladarogenys Brown 353
Epitritus Emery 354
Key to species (workers) 354
Strumigenys Smith 358
Key to species (workers) 360
Quadristruma Brown 400
Microdaceton Santschi 401
Key to species (workers) 402
Acknowledgements 403
References 403
Index 416
Synopsis
The nine genera (107 species) of Afrotropical dacetine ants are revised; keys to the genera and to the
species of each genus are presented. The genus-level name Miccostruma Brown is newly synonymized with
Smithistruma Brown, of which 35 species are recognized and 27 are described as new. Two species formerly
placed in Codiomyrmex Wheeler are transferred to Glamyromyrmex Wheeler, of which a total of 11
Afrotropical species are described. Eleven species of Serrastruma are recognised of which five are new (one
is an inquiline form). Six new synonyms are proposed in this genus and one previously synonymized name
is returned to specific status. Four species of Epitritus Emery and two of Microdaceton Santschi are known,
and one species each of Quadristruma Brown, Cladarogenys Brown and Trichoscapa Emery, the last
recorded for the first time from sub-Saharan Africa. Of the 41 recognized Afrotropical Strumigenys Smith
23 are described as new in this paper and two previously synonymized names are returned to specific status.
Introduction
Modern taxonomic work on the dacetine ants dates back only to Brown (1948) who published a
revisionary survey of the tribe as it was then understood. This was followed by a series of papers
refining the ideas of the original study by defining some of the genera more accurately and
delimiting other new genera (Brown, 1949a; 19496; 1950a). During the course of these studies it
was recognized that a number of genera originally placed with the dacetines did in fact
constitute a separate but convergently similar tribe, the Basicerotini, which was established by
Bull. Br. Mus. not. Hist. (Ent.) 46 (4): 267-416 Issued 25 August 1983
268 BARRY BOLTON
Brown (1949c). The genera of this tribe, whose members resemble the higher dacetines in many
features, were later fully revised by Brown & Kempf (1960), with keyed additions to the Old
World fauna by Taylor (19680).
These pioneering studies of Brown were followed by a series of revisionary papers aimed at
single dacetine genera or at the fauna of a particular region, along with many papers describing
new species from all over the world. These last are too numerous to list here but the main generic
and faunistic studies are as follows.
Acanthognathus , revised by Brown & Kempf (1969); Epitritus, keyed by Bolton (1972);
Glamyromyrmex and Gymnomyrmex, keyed by Kempf (I960); Kyidris, discussed by Wilson &
Brown (1956); Mesostruma, first revised by Brown (19526) with later additions by Taylor (1973);
Neostmma, revised by Brown (19596); Orectognathus , revised by Brown (19536) and later also
by Taylor (1980, and included references); Pentastruma, discussed by Brown & Boisvert (1978);
Serrastruma, revised by Brown (19520); Smithistruma, revised by Brown (19530; 1964);
Strumigenys of the Afrotropical region, revised by Brown (1954), of the Neotropical region,
revised by Brown (19626, and included references), and a continuing series of papers revising
the Indo- Australian fauna, the latest being Brown (1973c, and included references). A paper
discussing the evolution of the dacetines, which also includes an ecological synopsis of the
genera, has been produced by Brown & Wilson (1959). The Polynesian dacetine fauna is keyed
by Wilson & Taylor (1967) and the fauna of Japan, China and Taiwan is treated by Brown
(19490). The entire Neotropical fauna has been catalogued by Kempf (1972) and the Nearctic
fauna by Krombein et al. (1979). Dacetine ant larvae have been investigated by Wheeler &
Wheeler (1954).
Apart from short notes included in some of the above references papers dealing with the
biology of dacetine ants, either entirely or in part, include those of Brown (1950c), Carlin
(1981), Creighton (1937), Dejean (19800; 19806), Holldobler (1981), Kennedy & Schramm
(1933), Weber (19526), Wesson (1936), Wesson & Wesson (1939) and Wilson (1950, 1954,
1962).
On a world-wide basis 27 dacetine genera (384 species) are presently recognized, split into
four subtribes which were first proposed and defined by Brown (19526; 19530), and later
summarized by Brown & Wilson (1959). They are as follows.
Subtribe Dacetiti. Genera in which the eyes are dorsal or lateral and which lack antennal
scrobes. The antennae have 11 segments and the palp formula is 5,3. This subtribe includes only
the two small Neotropical genera Daceton and Acanthognathus.
Subtribe Orectognathiti. Contains only the Australian/New Guinean genus Orectognathus
which has the eyes lateral and lacks antennal scrobes. The antennae have 5 segments and of the
four funicular segments the second is the longest. The palp formula is 5,3.
Subtribe Epopostrumiti. The eyes are dorsolateral, placed above the scrobes when such are
present. The antennae have 4 or 6 segments and of the funicular segments the second is not the
longest. The palp formula is 5,3 or 3,2. Included here are the Afrotropical Microdaceton
(PF 3,2), and the Australasian Epopostruma, Mesostruma and Colobostruma (all with PF 5,3).
Subtribe Strumigeniti. The eyes are ventrolateral, placed within or beneath the scrobes which
are universally present though shallow and reduced in some. The antennae are 4 or 6 segmented
and the apical segment is much the longest of the funiculars. The palp formula is 1,1.
This subtribe holds 19 of the 27 genera and is split into two groups based on the presence or
absence of a spiniform apical fork on the mandibles. Those genera with the fork are termed
strumigeniform (Strumigenys, Neostruma, Quadristruma] , those without it smithistrumiform
(Asketogenys, Chelystruma, Cladarogenys, Codiomyrmex, Codioxenus, Dorisidris, Dysedrog-
nathus, Epitritus, Glamyromyrmex, Gymnomyrmex, Kyidris, Pentastruma, Serrastruma,
Smithistruma, Tingimyrmex, Trichoscapa) . The core-genera of this smithistrumiform group can
be regarded as Smithistruma, Trichoscapa and Pentastruma together with their close relatives
Gymnomyrmex, Kyidris and Tingimyrmex. In these genera the mandibles tend to be relatively
short, basically triangular, dorsoventrally flattened structures which may be quite delicate, are
not strongly downcurved, and in which the teeth are relatively small. The maximum number of
teeth is usually 12 but rarely may be as high as 19, following a strongly differentiated basal
THE AFROTROPICAL DACETINE ANTS 269
lamella. Mandibular variation in this complex includes reduction in number of teeth, variation in
size and arrangement of teeth, modifications in the development of the basal lamella and the
development in some of a diastema, of very variable extent, between the teeth and the basal
lamella (Brown, 1948; 19490; 19530; 1964; Brown & Boisvert, 1978; Wilson & Brown, 1956-
Kempf, 1960).
Modified away from these core-genera are several lines. In one of these the mandibles become
more massively constructed and strongly downcurved, and usually accompanying this is a
reduction in the number of teeth coupled with an increase in size in the teeth that remain
(Brown, 19500; 19530). Basal to this complex seem to be Chelystruma and Codiomyrmex in
which the mandibles are enlarged but a more or less full set of teeth is retained. Other genera
included here are Codioxenus, Glamyromyrmex and Dorisidris, the last showing a secondary
elongation of the mandibles, the second with marked variation in the number of teeth present.
A second line, represented by Serrastruma and Cladarogenys, shows an elongation of the
mandibles but with retention of their basically triangular shape. In these the teeth are initially
reduced and incorporated in a long series of denticles. The long basal lamella also becomes
denticulate and is pressed into service as part of the masticatory margin (Serrastruma; Brown,
19520). Further elongation of the blades coupled with a secondary reduction in dentition gives
the condition seen in Cladarogenys (Brown, 1976).
A third line shows elongation of the mandibles with eventual loss of the triangular shape and
their development into long narrow blades. This line, including Dysedrognathus and Epitritus,
was postulated by Taylor (19686). It involves an initial increase in mandible length coupled with
an increase in the number of teeth. With continuing increase in length the teeth on the main part
of the blade become spaced out or lost and only those crowded near the apex remain.
The last smithistrumiform genus to be considered here, Asketogenys, appears to be an
independent relatively long-mandibulate derivative of Smithistruma in which the teeth at about
the midlength of the masticatory margin have been enlarged (Brown, 1972).
Finally there is the anomalous Neotropical genus Phalacromyrmex and an apparently related
undescribed genus from the Indo- Australian region which do not fit any of the above subtribes.
At first glance they appear to fall into the smithistrumiform group but they have 9-11 antennal
segments, a palp formula of 3,2 (Phalacromyrmex), and lack other characters which may be
considered as typically smithistrumiform such as spongiform appendages on the pedicel
segments, a transverse lamellar or spongiform strip across the base of the first gastral tergite and
a basal lamella on their massively constructed bear trap-like mandibles. Indeed, the massive
mandibles seen in these genera are reminiscent of some Glamyromyrmex species but also have
some resemblance to the Malagasy genus Pilotrochus Brown, so there is a very strong possibility
that Phalacromyrmex and its undescribed relative may be convergent on the smithistrumiform
dacetines from some other part of the Myrmicinae.
The distribution of the 27 genera includes all the zoogeographical regions, but dacetines are
absent from the northern parts of the Palaearctic and Nearctic. Three of the genera are very
widespread (Smithistruma, Strumigenys, Epitritus) and two small genera include efficient
tramp-species (Quadristruma, Trichoscapa) whose members have been introduced by human
commerce over much of the tropical and subtropical zones. Of the remaining 22 genera nine are
restricted to the Neotropical region, two to the Afrotropical, one to the Oriental, two to the
Indo- Australian and two to the Australasian region. The remaining six genera are shared by
two, usually adjacent, zoogeographical regions.
The table below summarizes the number of described dacetine species of the world and
indicates their distribution. For the purposes of this study the Afrotropical and Malagasy are
regarded as separate regions and in the table the Indo Australian region is taken to include New
Guinea and the Pacific island systems. Tramp-species or species shared by two regions are
entered in the table only in their presumed region of origin. Thus Serrastruma ludovici (Forel),
5. simoni (Emery), Strumigenys scotti Forel, St. rogeri Emery, Quadristruma emmae (Emery),
and Trichoscapa membranifera (Emery) are all recorded in the Afrotropical column alone,
although all have been found in other parts of the world. Smithistruma dubia Brown is recorded
as Indo-Australian although also present in Australia.
270 BARRY BOLTON
Genus-level names which are now regarded as synonyms are excluded from the table. Apart
from those listed in this paper under the appropriate generic headings, the following are
recognized synonyms.
Alistruma Brown is a synonym of Colobostmma. [Synonymy by Brown, 1959c.]
Arnoldidris Brown is a synonym of Orectognathus. [Synonymy by Brown, 1973ft. ]
Clarkistmma Brown is a synonym of Colobostruma. [Synonymy by Brown, 1959c.]
Hexadaceton Brown is a synonym of Epopostruma. [Synonymy by Brown, 1973ft.]
Polyhomoa Azuma is a synonym of Kyidris. [Synonymy by Brown & Yasumatsu, 1951.]
The fossil genus Hypopomyrmex Emery, formerly considered a dacetine, has been re-
examined by Brown & Carpenter (1978) and excluded from the tribe.
In the table the zoogeographical regions are abbreviated as follows. Af. Afrotropical, Au.
Australasian, In. Indo- Australian, Ma. Malagasy, Ne. Nearctic, No. Neotropical, Or. Oriental,
Pa. Palaearctic.
Genus Ne. No. Pa. Af. Ma. Or. In. Au. Total
Acanthognathus Mayr
Asketogenys Brown
Chelystruma Brown
Cladarogenys Brown
Codiomyrmex Wheeler
Codioxenus Santschi
Colobostruma Wheeler
Daceton Perty
Dorisidris Brown
6
1
2
1
1
1
1
1
1
2
8
6
1
1
1
4
1
9
1
1
Dysedrognathus Taylor
-
_
-
1
-
1
Epitrltus Emery
-
1 4
1
1
-
7
Epopostruma Forel
-
_
-
-
4
4
Glamyromyrmex Wheeler
7
11
-
-
-
18
Gymnomyrmex Borgmeier
6
- - -
-
-
-
6
Kyidris Brown
-
- - -
2
2
-
4
Mesostruma Brown
- -
_ _ _
-
-
6
6
Microdaceton Santschi
- -
2
-
—
-
2
Neostruma Brown
6
_ _ _
-
-
-
6
Orectognathus Smith
-
_ _ _
-
10
19
29
Pentastruma Forel
- -
_ _
2
-
-
2
Phalacromyrmex Kempf
1
_ _
-
-
-
1
Quadristruma Brown
-
1
-
1
-
2
Serrastruma Brown
- -
11
-
-
-
11
Smithistruma Brown
24 19
3 35
5
11
—
97
Strumigenys Smith
1 54
41 1
8
48
8
161
Tingimyrmex Mann
1
_
-
-
-
1
Trichoscapa Emery
-
1
-
-
-
1
Total
25 106
4 107 1
18
76
47
384
Discounting papers whose sole purpose was the mass description of new forms the history of
Afrotropical dacetine studies prior to Brown's (1948) publication consisted only of the mono-
graphic study of South African ants by Arnold (1917), the catalogue of Wheeler (1922) and the
key presented by Santschi (19130). Arnold and Wheeler both recognized three genera in the
Afrotropical region, Microdaceton, Strumigenys and Epitritus, of which only the first has
remained unchanged to the present day.
The genus Strumigenys, as recognized by Wheeler (1922), contained not only the long-
mandibulate forms with a spiniform apical fork which constitute the genus as it is presently
understood, but also a number of short-mandibulate species which lacked an apical fork and
which were grouped under a subgenus Cephaloxys. Brown (1948) recognized that these
short-mandibulate forms were fundamentally different from the foregoing group and also noted
THE AFROTROPICAL DACETINE ANTS 271
that Cephaloxys, beside being a preoccupied name, itself contained two disparate groups of
species which differed consistently in the structure of their mandibles. Erecting Smithistruma to
replace the name Cephaloxys, Brown (1948) proceeded to remove those African species which
had multi-denticulate mandibles to a separate subgenus of Smithistruma, Serrastmma, which he
later elevated to generic status (Brown, 19490) and then revised both genera (Brown, 19520,
19530).
The two African species placed in Epitritus by their original authors and retained there by
Wheeler (1922) were recognized by Brown (1948) as falling outside the limits of that genus. He
transferred them to a separate genus, Miccostruma, which is now regarded as a synonym of
Smithistruma (see discussion of that genus). True Epitritus was later discovered in Africa
(Brown, 19620) and four species are now known from that continent. Brown (19530) described a
species of Codiomyrmex from Africa, and Taylor (1965) another. These two, plus nine newly
discovered species, are best referred to Glamyromyrmex as noted under the discussion of that
genus. Finally, species of Quadristruma and Trichoscapa have been recorded from Africa, the
former by Bolton (1973), the latter newly reported here; and the monotypic genus Cladarogenys
has recently been described by Brown (1976).
Thus the Afrotropical region currently has nine dacetine genera containing a total of 107
species. The vast majority of these are found in the leaf litter and topsoil layers where they
constitute an important fraction of the fauna. Nests are made either in compressed leaf litter, in
the soil, or in pieces of wood or stumps embedded in the litter and topsoil layers. A couple of
Strumigenys species are known which nest and forage arboreally and some Serrastruma may
ascend tree trunks to a considerable distance above the ground.
This study of the dacetine ants of sub-Saharan Africa is the latest part in a series of papers
aimed towards a revision of the entire myrmicine ant fauna of the Afrotropical region.
Previously published parts include Bolton (1974; 1976; 1980; 19810; 19816; 1982).
Measurements and indices
Total Length (TL). The total outstretched length of the ant from the mandibular apex to the
gastral apex.
Head Length (HL). The length of the head proper, excluding the mandibles, measured in a
straight line from the mid-point of the anterior clypeal margin to the mid-point of the
occipital margin, in full-face view. In species where the clypeal margin or the occipital
margin (or both) is concave the measurement is taken from the mid-point of a transverse
line spanning the anteriormost or posteriormost projecting points respectively
Head Width (HW). The maximum width of the head in full-face view, measured behind the eyes.
(In Microdaceton ignoring the projecting tubercles.)
Cephalic Index (CI). HWxlOO
HL
Mandible Length (ML). The straight-line length of the mandible, measured in the same plane for
which the HL measurement is taken, from the mandibular apex to the transverse through
the anteriormost point or points of the clypeal margin.
Mandibular Index (MI). ML x 100
HL
Scape Length (SL). The maximum straight-line length of the antennal scape excluding the basal
constriction or neck close to the condylar bulb. (In Epitritus measured from the tip of the
subbasal lobe to the scape apex.)
Scape Index (SI). SLxlOO
HW
Pronotal Width (PW). The maximum width of the pronotum in dorsal view.
Alitrunk Length (AL). The diagonal length of the alitrunk in profile from the point at which the
pronotum meets the cervical shield to the posterior base of the metapleuron.
272 BARRY BOLTON
Abbreviations of depositories
AMNH American Museum of Natural History, New York, U.S.A.
BMNH British Museum (Natural History), London, U.K.
CAS California Academy of Sciences, San Francisco, California, U.S.A.
ENS A Ecole Nationale Superieure Agronomique, Toulouse, France.
IE Istituto di Entomologia del'Universita, Bologna, Italy.
MCSN Museo Civico di Storia Naturale 'Giacomo Doria', Genoa, Italy.
MCZ Museum of Comparative Zoology , Cambridge , Massachusetts , U . S . A .
MHN Museum d'Histoire Naturelle, Geneva, Switzerland.
MNHN Museum National d'Histoire Naturelle, Paris, France.
MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, Germany (D.D.R.).
MR AC Musee Royal de 1'Afrique Centrale, Tervuren, Belgium.
NMB Naturhistorisches Museum, Basle, Switzerland.
NMV Naturhistorisches Museum, Vienna, Austria.
SAM South African Museum, Cape Town, South Africa.
TM Termeszettudomanyi Muzeum, Budapest, Hungary.
USNM United States National Museum , Washington ,D.C.,U.S.A.
Diagnosis of Afrotropical dacetine ants
WORKER. Myrmicine ants in which the antennae have only 4 or 6 segments, the funiculus ending in a
2-segmented club. Pedicel segments with spongiform or lamelliform appendages; sometimes the appen-
dages small but always present (Figs 13, 16, 22, 30, 38-44, 68-70, 77, 78, 80, 81). Mandibles of two basic
forms , either produced into a pair of long narrow linear blades (Figs 45-67 , 7 1-79) with or without an apical
fork of spiniform teeth, or the mandibles shorter, usually subtriangular, always lacking an apical fork and
armed with 8->30 teeth or denticles (Figs 1-12, 14, 15, 17-21, 23-37). Clypeus broad and shield-like,
broadly inserted between the widely separated frontal lobes; the latter sometimes projecting beyond the
lateral margins of the head. Palp formula usually 1,1 but higher in Microdaceton (PF 3,2). Antennal
scrobes usually present (not in Microdaceton, Figs 78, 79), situated above the eye, the latter generally small
to moderate in size and commonly on the ventrolateral margin of the head. Propodeum usually with a pair
of teeth or spines, rarely otherwise. Bizarre pilosity frequently developed.
Among the Afrotropical myrmicine ants the dacetines are easily identified by their low
antennomere count of 4 or 6 and their possession of spongiform or lamellate appendages on the
pedicel segments. Only one other genus in the region has an antennomere count as low as 6,
Melissotarsus Emery, but this differs from the dacetine genera as follows.
Dacetine genera
Spongiform or lamellate appendages
present on pedicel segments.
Frontal lobes widely separated,
situated laterally on anterior
half of head.
Clypeus projecting back between
frontal lobes.
Mandibles blade-like or subtriangular;
if the latter then always with more
than 4 teeth.
Antennal scrobes present except in
Microdaceton but here the
mandibles are linear.
Propodeum usually armed with a pair
of spines or teeth, often with a
strong infradental lamella.
Anterior coxae as large as or
larger than the middle and hind
coxae.
Melissotarsus
Spongiform or lamellate appendages absent
from pedicel segments.
Frontal lobes confluent, situated
centrally and high on dorsum of head.
Clypeus not projecting back between
frontal lobes.
Mandibles short, at most with 4 teeth,
the apical long and finger-like
when unworn.
Antennal scrobes absent.
Propodeum evenly rounded and unarmed.
Anterior coxae much smaller than the
massively developed middle and hind
coxae (Bolton, 1982: 334, fig. 23).
THE AFROTROPICAL DACETINE ANTS 273
Dacetine genera - cont. Melissotarsus - cont.
Basitarsal leg segments not swollen, Basitarsal leg segments greatly swollen,
without an apical circlet of teeth with an apical circlet of teeth on
on the anterior edge of the middle the anterior edge of the middle
and hind basitarsi. and hind basitarsi.
Key to Afrotropical dacetine ant genera (workers)
1 Mandibles elongate and linear, produced into narrow projecting blades (Figs 45-67, 71-79):
never triangular/subtriangular, never serially multidentate or denticulate 2
- Mandibles triangular or subtriangular, not produced into narrow projecting blades; apical
(masticatory) margin serially multidentate or denticulate but teeth sometimes reduced
(Figs 1-15, 17-37) 6
2 Apex of each mandibular blade armed with a fork of 2 or 3 spiniform teeth set in a more or less
vertical series, with or without intercalary denticles between the spiniform fork teeth
(Figs 49-67, 71-79) 3
- Apex of each mandibular blade either with a single long tooth at the dorsal apex subtended by a
series of minute denticles , or with a series of minute denticles only (Figs 45-48) 5
3 Apical fork of mandibles with 3 spiniform teeth; blades of mandibles without preapical teeth.
Maxillary palp 3-segmented. Antennal scrobes absent, the eyes dorsolateral. Petiole node
with a pair of teeth or short spines, postpetiole with lamellate appendages (Figs 78-81)
MICRODACETON (p. 401)
Apical fork of mandibles with 2 spiniform teeth; blades of mandibles usually with preapical
teeth. Maxillary palp 1-segmented. Antennal scrobes present, the eyes ventrolateral. Petiole
node unarmed , postpetiole with spongiform appendages (Figs 49-77) 4
4 Antennae with 4 segments (Fig. 67) QUADRISTRUMA (p. 400)
Antennae with 6 segments STRUMIGENYS (p. 358)
5 Antennal scapes with a broad anteriorly projecting subbasal lobe . Clypeal margin with spatulate
or strap-like projecting hairs. Head with large orbicular hairs present; the head broad,
CI>100(Figs46-48) EPITRITUS (p. 354)
- Antennal scapes linear, without a projecting lobe. Clypeal margin without spatulate or
strap-like projecting hairs. Head only with simple hairs present; the head narrower, CI<80
CLADAROGENYS (p. 353)
6 Differentiated prominent basal lamella of mandible absent. Apical (masticatory) margin of
mandible with >20 denticles, the basal 4-8 of which may be enlarged. Mandibles relatively
long, MI>25 (Figs 34-37) SERRASTRUMA (p. 335)
- Differentiated prominent basal lamella of mandible present. Apical (masticatory) margin of
mandible with 17 or fewer teeth or denticles of varying size. Mandibles relatively short,
MK25 (Figs 1-15, 17-33) 7
7 Fully closed mandibles with a strongly defined transverse basal border which is separated
from the anterior clypeal margin by a conspicuous impression or gap (Fig. 21). Basal
lamella of mandible situated ventral to the basalmost tooth, in a plane almost at right-
angles to the anterior portion of the mandible, not visible in full-face view with the mandibles
open TRICHOSCAPA (p. 319)
- Fully closed mandibles without a strongly defined basal border, the basal region of the mandible
contiguous with or overlapped by the anterior clypeal margin, the two not separated by an
impression or gap (Figs 1-12, 14, 15, 17-20, 23-29). Basal lamella of mandible following
basalmost tooth in the same plane , visible in full-face view with the mandibles open 8
8 With the head in profile the mandibles increasing in width from base to apex and the distal
portion of the blades passing into a strong downcurved arc so that part or most of the apical
margin is at right-angles to the long axis of the head (Figs 30-33). Masticatory margin of
mandible armed with a basal lamella plus 8-11 teeth , the basal 5-8 of which may be very strong
(Figs23-29) GLAMYROMYRMEX (p. 320)
- With the head in profile the mandibles with their upper and lower margins approximately
parallel for most of their length or evenly tapering anteriorly. At most the extreme tip
of the mandible downcurved, without a major part of the apical margin at right-angles to
the long axis of the head (Fig. 13). Masticatory margin of mandible armed with a basal
lamella plus 12-17 teeth or denticles, the apicalmost group of which are minute (Figs 1-12, 14,
15, 17-20) SMITHISTRUMA (p. 274)
274 BARRY BOLTON
SMITHISTRUMA Brown
(Figs 1-20)
Cephaloxys F. Smith, 1865: 76. Type-species: Cephaloxys capitata F. Smith, 1865: 77, by monotypy.
[Junior homonym of Cephaloxys Signoret, 1847: 294 (Hemiptera).]
Smithistruma Brown, 1948: 104. Type-species: Strumigenys pulchella Emery, 1895b: 327, pi. 8, fig. 19, by
original designation.
Wessonistruma Brown, 1948: 106 [as subgenus of Smithistruma]. Type-species: Strumigenys pergandei
Emery, 18956: 326, pi. 8, figs 17, 18, by original designation. [Synonymy by Brown, 1973a: 35.]
Weberistruma Brown, 1948: 106 [as subgenus of Smithistruma]. Type-species: Strumigenys (Cephaloxys)
leptothrix Wheeler, 1929: 55, fig. 7, by original designation. [Synonymy by Brown, 19730: 35.]
Miccostruma Brown, 1948: 123. Type-species: Epitritus mandibularis Szabo, 1909: 1, fig. 2, by original
designation. Syn. n.
Platystruma Brown, 1953a: 112 [as subgenus of Smithistruma]. Type-species: Strumigenys (Cephaloxys)
depressiceps Weber, 1934: 47, fig. 6, by original designation. [Synonymy by Brown, 19730: 35.]
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles triangular to narrowly subtriangular and
short (MI 7-20), serially dentate or denticulate and lacking an apical fork of spiniform teeth. When fully
closed at least the base of the mandible, but sometimes much of its length, concealed by the clypeus;
without a sharply defined transverse basal margin which is separated from the anterior clypeal margin by a
conspicuous impression or gap. In profile the mandibles with their upper and lower margins approximately
parallel for most of their length or evenly tapering anteriorly (Fig. 13), at most with the tip of the mandible
downcurved, never with a major part of the apical (masticatory) margin strongly arched-downcurved or at
right-angles to the long axis of the head. Apical (masticatory) margin of the mandible with 12-17 teeth
following a conspicuously differentiated prominent basal lamella, the lamella concealed by the clypeus
when the mandibles are closed. Arrangement of teeth either with 5 larger members, distal to the basal
lamella, forming a principal tooth row in which the teeth may be about the same or of different sizes; these
followed by two somewhat smaller teeth and a series of 4 minute denticles before the apical tooth or
denticle; or with a principal row of 7 teeth followed by 4 minute denticles and an apical tooth or denticle; or
with a principal row of 7-8 teeth separated from the basal lamella by a long diastema, the distal member of
this series by far the largest tooth on the mandible and followed sequentially by 3 small teeth, a slightly
larger tooth, 4 denticles and an apical tooth.
Species of Smithistruma have been described from all the zoogeographical regions except the
Australian and Malagasy. Undescribed species from the latter region are represented in the
BMNH and MCZ, Cambridge collections, but the genus is represented in the Australian region
only by the introduced 5. dubia Brown (R. W. Taylor, pers. comm.). On a worldwide basis 97
Smithistruma species have been described, of which 35 are Afrotropical. As indicated in the
table of dacetine species presented in the introduction to this paper (p. 270), all other
zoogeographical regions now fall behind the Afrotropical in terms of number of Smithistruma
species, but this picture is somewhat distorted as many new species from other regions await
description in the museums of the world.
The modern taxonomic study of Smithistruma dates back only to 1953 when Brown (19530)
published a world revision of the genus as it was then understood, having previously defined the
genus and a number of subgenera in an earlier introductory paper (Brown, 1948). Later Brown
(1964) produced a supplement to the world revision and subsequently indicated (Brown, 1973a)
that the subgenera should be regarded as synonyms of Smithistruma, except for Serrastruma
which he had previously raised to generic status (Brown, 1949«). The collapse of the subgenera
was due solely to the continuing discovery of species linking groups which originally seemed
quite distinct, and this process is still in operation as species reducing or bridging the gaps
between many of the genera of short-mandibulate dacetines continue to be found. Indeed, the
position of Smithistruma itself is not assured. It stands central to, and is the largest single
member of, a group of closely related mostly small genera of dacetines with short mandibles
which also includes Pentastruma, Trichoscapa, Tingimyrmex, Kyidris, Chelystruma, Codiomyr-
mex, Codioxenus, Dysedrognathus, Glamyromyrmex and Gymnomyrmex. Closely linked to
these are a number of forms with more specialized and usually longer mandibles which appear to
be derived from various members of the Smithistruma-group, namely Asketogenys, Serrastru-
ma, Cladarogenys, Dorisidris and Epitritus. In recent years Brown (1973a) and Brown &
THE AFROTROPICAL DACETINE ANTS 275
Boisvert (1978) have discussed a number of these names and generally concluded that
Serrastruma, Tingimyrmex, Epitritus and Kyidris are valid genera, but that the remainder are
dubious and in a state of flux as modern collecting techniques continue to reveal previously
unknown species which are gradually filling the gaps originally invoked to separate the genera.
In the present paper a single generic name, Miccostmma, is newly synonymized with
Smithistruma. Miccostruma was originally erected by Brown (1948) to include two Afrotropical
species, mandibularis and marginata, which had both been regarded previously as members of
Epitritus; later Brown (19730) added a third species, tigrilla. The characters which Brown used
to separate Miccostruma from Smithistruma were the possession of 4-segmented antennae and
relatively very short mandibles by the former, as opposed to 6-segmented antennae and longer
mandibles in the latter. With the description of S. cavinasis by Brown (19506) it became
apparent that species with 6-segmented antennae could also have very short mandibles, as
Brown (19530) mentioned in his world revision. This discovery seriously eroded the strength of
the character and the present survey has indicated that it has no value at genus level as short
mandibles (MI 10 or less) are by no means confined to species with 4-segmented antennae but
occur in a wide range of forms from several species-groups.
Concerning the reduced antennal segmentation Brown (19730) has already pointed out that it
is a weak character as the fusion of segments which takes place to reduce the antennomere count
is not always complete. In tigrilla, with correct lighting, the limits of the former segments 3-5,
which fuse to form segment 3 in tigrilla, can be seen. At present seven Afrotropical species with
4-merous antennae are known. An analysis of their characters indicates that the reduction in
antennal segmentation from 6 to 4 has occurred in three separate lines derived from different
groups within Smithistruma. Of these species with 4-merous antennae fulda, mandibularis,
ninda and tigrilla form a single group which is very closely related to, and most probably directly
descended from, the emarginata-group. In these four species the clypeus is broad and prominent
anteriorly and laterally, is fringed by a continuous row of large specialized hairs and has the
anterior margin concave; the mandibles have a high truncated-triangular basal lamella and a
principal tooth row of 5. Body pilosity is extremely sparse or absent and flagellate hairs are
lacking, but the leading edges of the scapes have projecting strong specialized hairs. The
pronotum lacks both lateral margination and a median dorsal longitudinal carina.
Compared to these fundamental shared characters of mandibularis and its allies the other
three species with 4-merous antennae are very different. In marginata the clypeus lacks hairs of
any description, has the anterior margin broadly convex and the sides parallel. The mandibles
have the basal lamella shaped as a long low lobe and have a principal tooth row of 7. Body
pilosity is present and long flagellate hairs occur on the head and alitrunk, but the leading edges
of the scapes lack projecting hairs. The pronotum is sharply marginate laterally and has a strong
median carina. 5. tacta and vodensa share most of the characters of marginata but have the
clypeus differently constructed. In tacta-group the clypeus is narrow, has convergent sides with a
produced and narrowly rounded anterior margin, and is densely clothed with fine hairs. 5.
marginata and tacta share more characters between them than either one does with mandibularis
and its allies, but the fundamental difference in clypeal form indicates that they have arisen from
separate origins within Smithistruma.
The disparity of these species-group level characters, between mandibularis and its allies on
the one hand and marginata on the other, shows that Miccostruma contained, from its inception,
elements from fundamentally different origins within Smithistruma. The discovery of tacta and
vodensa, from yet another group, makes it clear that reduction in antennomere count has little
or no value at genus level amongst the short-mandibulate dacetines. The removal of marginata
from Miccostruma does leave a uniform group of species centring on mandibularis, but the
overwhelming similarity of these species to the members of the emarginata-group , and the
collapse of the original separating characters of Miccostruma, confirms that the species can no
longer be regarded as constituting a separate genus.
All known Afrotropical species of Smithistruma inhabit the leaf litter and topsoil layers,
usually nesting directly into the ground or in rotten wood. No species is remarkably common and
collections of many species are only of a few individual workers. In recent years increased
276 BARRY BOLTON
collecting by funnelling techniques has shown that Smithistruma is by no means as poorly
represented in Africa as was thought only a few years ago, and many more species probably
await discovery; but as Brown (1952a, 1953a) has pointed out, Smithistruma in Africa is more or
less eclipsed by the much more common, widely distributed and versatile species of Serrastmma
which, though having fewer species, greatly outnumber Smithistruma in terms of numbers of
individuals.
List of Afrotropical Smithistruma
mandibularis-group weberi-group
fulda sp. n. arahana sp. n.
mandibularis (Szabo) comb. n. enkara sp. n.
nincln sp. n. / 'enkara sp. n.
tigrilla (Brown) comb. n. kerasma sp. n.
emarginata-group malaplaxsp.n.
behasyla sp. n. mekaha sp. n.
ca vinasis B ro wn minkara sp . n .
chyatha sp. n. nykara sp. n.
datissa sp. n. placora sp. n.
dendexa sp. n. synkara sp. n.
emarginata (Mayr) tolomyla sp. n.
gatudasp. n. weberi Brown
hensekta sp. n. marginata-group
impidora sp. n. marginata (Santschi) comb. n.
sharra sp. n. rusta sp. n.
truncatidens Brown oxysma-group
transversa-group anarta sp. n.
transversa (Santschi) oxysma sp. n.
ferrouAgroup tacta-group
terronisp. n. tacta sp. n.
vodensa sp. n.
Key to species (workers)
1 Antennae with 4 segments 2
- Antennae with 6 segments 8
2 Anterior clypeal margin convex in full-face view (Fig. 20). Pronotum with a median longitudin-
al carina and pronotal humeri each with a single flagellate hair 3
Anterior clypeal margin concave in full-face view (Figs 1,2). Pronotum without a median
longitudinal carina and pronotal humeri without flagellate hairs 5
3 Clypeus broad, in full-face view expanded laterally far beyond the line of the outer margins of
the closed mandibles (shaped as in Fig. 17). Dorsum of clypeus without hairs. (Ivory Coast,
Kenya, Zimbabwe) marginata (p. 312)
Clypeus narrow, in full-face view more or less continuing the line of the outer margins of the
closed mandibles (Fig. 20) . Dorsum of clypeus with abundant short curved hairs 4
4 Pronotum sharply marginate laterally, the dorsum unsculptured. Head relatively broad and
scapes short, CI>60,SI<75. (Ivory Coast, Ghana, Cameroun, Zaire) tacta (p. 317)
- Pronotum not marginate laterally, the dorsum weakly sculptured. Head relatively narrow and
scapes long, CI<60,SI>100. (Cameroun) vodensa (p. 317)
5 Entire body coloured with broad alternating yellow and black transverse bands. Basigastral
costulae arising in a continuous row across the tergite, without a central gap. (Ivory Coast,
Cameroun) tigrilla (p. 284)
Body uniformly coloured, without alternating yellow and black broad transverse bands.
Basigastral costulae radiating from each side of a central gap 6
6 With the head in profile the dorsum with a pair of short erect hairs close to the occipital margin.
(Kenya, Tanzania, Angola) mandibularis (p. 283)
With the head in profile the dorsum without erect hairs 7
7 Anterior half of clypeus with a broad longitudinal impression mid-dorsally which is filled with
short scale-like hairs, these hairs directed towards the midline. Subbasal elbows of scapes
THE AFROTROPICAL DACETINE ANTS 277
extensively developed and strongly angular (Fig. 1). Pronotal dorsum with dense fine
longitudinal rugulae or costulae. (Ivory Coast) fulda (p. 282)
- Anterior half of clypeus without an impression mid-dorsally, this area without short scale-like
hairs directed towards the midline. Subbasal elbows of scapes weakly developed, evenly
rounded and not angled (Fig. 2). Pronotal dorsum smooth or at most with faint superficial
shagreening on the glossy surface. (Ivory Coast, Ghana, Nigeria, Chad, Cameroun)
ninda (p. 284)
8 Dorsum of pronotum extremely coarsely sculptured with strong rugae or sulci which are close
packed and give a very coarse overall appearance; without broad shining areas between the
rugae or sulci 9
- Dorsum of pronotum varying from smooth to densely reticulate-punctate. Feeble striate
sculpture or extremely fine superficial rugulae may occur but coarse rugae or sulci are always
absent 20
9 Disc of postpetiole densely and strongly longitudinally costulate 10
- Disc of postpetiole smooth and unsculptured or at most uneven and feebly punctate, never
longitudinally costulate 12
10 Head exceptionally long and narrow, CI 54-58 (Fig. 14); antennal scapes relatively long,
SI 73-78. (Ivory Coast) minkara (p. 306)
- Head shorter and broader, CI >60; antennal scapes relatively short , SI 68-73 11
11 Disc of postpetiole in dorsal view surrounded on all sides by dense spongiform material.
Transverse spongiform strips behind petiole and postpetiole broad and complete. Pronotal
dorsum coarsely longitudinally rugose. (Ivory Coast, Ghana, Cameroun) enkara (p. 301)
- Disc of postpetiole in dorsal view with spongiform material only developed posteriorly and
posterolaterally. Transverse spongiform strips behind petiole and postpetiole interrupted
medially. Pronotal dorsum irregularly reticulate-rugose. (Zimbabwe) nykara (p. 307)
12 With the alitrunk in profile the metanotal groove distinctly impressed. (Fig. 16) 13
- With the alitrunk in profile the metanotal groove not impressed , the dorsal outline continuous 15
13 Posterior spongiform margin of postpetiole very deeply indented medially, the indentation
reaching to the margin of the disc. (Zaire) weberi (p. 311)
Posterior spongiform margin of postpetiole very shallowly indented medially, the indentation
not approaching the margin of the disc, always a thick band of spongiform material
remaining between the margin of the disc and the innermost point of the indentation in the
spongiform tissue 14
14 Hairs on dorsum of head strongly arched forward so that their apices are in contact or almost in
contact with the surface, without hairs which are erect and sharply angled at about their
midlengths. (Cameroun) mekaha (p. 305)
- Hairs on dorsum of head, especially on posterior half, with the basal half to two-thirds erect,
the apical portion of each hair sharply angled forward. (Cameroun) kerasma (p. 303)
15 Dorsum of head behind clypeus only with fine soft flexuous looped or arched simple hairs,
without specialized strong hairs which are similar to those on the clypeal dorsum. (Nigeria,
Zaire , Angola) malaplax (p. 304)
- Dorsum of head behind clypeus usually with some fine hairs but also with long stout very
conspicuous hairs which are erect and curved anteriorly and which are similar to those on the
clypeal dorsum 16
16 With the head in profile the longest hairs arising from the clypeal dorsum (the posteriormost
row) at most only half the length of the longest hairs on the cephalic dorsum, which arise just
behind the level of the eyes. (Cameroun) placora (p. 308)
- With the head in profile the longest hairs arising from the clypeal dorsum about the same length
as those situated on the cephalic dorsum just behind the level of the eyes, or only fractionally
different 17
17 Posterior spongiform strip of petiole enormously developed , in dorsal view the thickness of the
strip from front to back greater than the exposed length of the petiole node; in appearance
the strip obviously densely spongiform rather than lamellar. (Cameroun) arahana (p. 300)
- Posterior spongiform strip of petiole narrow and lamellar, in dorsal view the thickness of the
strip from front to back conspicuously less than the exposed length of the petiole node 18
18 With the head in posterior view the long hairs arising from the dorsum and sides distinctly
swollen apically, increasing markedly in width from base to apex. (Angola) fenkara (p. 302)
- With the head in posterior view the long hairs arising from the dorsum and sides of uniform
width throughout their length, not increasing in width from base to apex 19
278 BARRY BOLTON
19 Median indentation in posterior margin of the spongiform strip bordering the postpetiole
posteriorly not reaching the sclerotized portion of the disc. Larger species, HW 0-50.
(Gabon) synkara (p. 309)
Median indentation in posterior margin of the spongiform strip bordering the postpetiole
posteriorly reaching to the sclerotized portion of the disc. Smaller species, HW <0-40.
(Cameroun) tolomyla (p. 310)
20 Pronotal humeri each with a single long flagellate hair, the pronotal dorsum with a median
longitudinal ridge or carina, at least anteriorly. Leading edges of scapes lacking anteriorly
projecting strong hairs, those present being short and appressed. Anterior margin of clypeus
convex in full-face view and the lateral clypeal margins lacking a continuous fringe of
anteriorly curved spatulate or spoon-shaped hairs (Figs 17-19) 21
Pronotal humeri without flagellate hairs, the pronotal dorsum without a median longitudinal
ridge or carina. Leading edges of scapes with a row of anteriorly projecting strong hairs.
Anterior margin of clypeus transverse to concave in full-face view and the lateral clypeal
margins with a continuous fringe of anteriorly curved spatulate or spoon-shaped hairs
(Figs 3-12) 23
21 Clypeal dorsum in profile without hairs. In full-face view the anterior clypeal margin broadly
and evenly convex; sides of clypeus approximately parallel, not forming a more or less
continuous line with the outer margins of the closed mandibles (Fig. 17). (Zimbabwe)
rusta (p. 313)
Clypeal dorsum in profile with posteriorly or posteromedially curved hairs which are weakly
clavate apically. In full-face view the anterior clypeal margin narrowly convex; sides of
clypeus converging anteriorly and forming a more or less continuous line with the outer
margins of the closed mandibles (Figs 18,19) 22
22 Elongate hairs on first gastral tergite restricted to a transverse row of four close to the base.
Flagellate hairs absent from upper scrobe margins (Fig. 18). (South Africa) anarta (p. 314)
Elongate hairs on first gastral tergite numbering 12 or more, not restricted to area close to the
base. Two or three flagellate hairs present on eachfupper scrobe margin (Fig. 19). (South
Africa, Lesotho) oxysma (p. 315)
23 Mandibles with 12 teeth of which one of the basal row of five is the largest . From the base to the
apex the mandible with five relatively large teeth followed by two slightly smaller teeth, four
minute denticles and a small apical tooth. Diastema between basal lamella and basalmost
tooth minute or absent, always smaller than the height of the basalmost tooth (Figs 3-6,
8-12) 24
Mandibles with 16-17 teeth of which the seventh or eighth from the base is by far the largest.
From the base to the apex the mandible with six or seven small teeth followed by a relatively
very large tooth, three small teeth, a single slightly larger tooth, four minute denticles and an
apical tooth. Diastema between basal lamella and basalmost tooth long, distinctly much
longer than the height of the basalmost tooth (Fig. 7). (Cameroun) terroni (p. 299)
24 With the head in full-face view the entire dorsum with large flattened very broadly scale-like to
suborbicular hairs (Fig. 3) 25
With the head in full-face view the dorsum without large flattened broadly scale-like to
suborbicular hairs or at most with such hairs occurring in one or two sharply defined
transverse bands 26
25 Postpetiole and first gastral tergite with erect hairs present. CI 63-67. Scale-like hairs
inconspicuous or absent on disc of postpetiole. (Cameroun, Zaire, Angola) .... cavinasis (p. 287)
Postpetiole and first gastral tergite without erect hairs. CI 56-63. Scale-like hairs sparse but
obvious on disc of postpetiole. (Ivory Coast, Ghana, Cameroun, Angola) sharra (p. 295)
26 First gastral tergite without standing hairs 27
First gastral tergite with standing hairs which are usually numerous but which may be restricted
to a single basal pair and a single apical pair 28
27 Dorsum of head with a transverse band of broadly scale-like to suborbicular hairs just in front
of the occipital margin (Fig. 5). (Ghana) chyatha (p. 288)
Dorsum of head lacking hairs of any description except for the fringe around the clypeal
margins (Fig. 6). (Ivory Coast) impidora (p. 294)
28 Base of first gastral tergite sharply impressed medially, the sclerite with a dented appearance.
Scapes relatively long, SI 72-80 (Fig. 8). (Ivory Coast, Ghana, Togo, Burundi, Zimbabwe,
Angola, South Africa) emarginata (p. 291)
Base of first gastral tergite not impressed medially. Scapes shorter, SI 58-67 (Figs 4, 9-12) 29
THE AFROTROPICAL DACETINE ANTS 279
29 Pronotal disc glassy smooth between very widely scattered small punctures. (Rwanda)
gatuda (p. 292)
- Pronotal disc uniformly closely sculptured, subopaque to opaque 30
30 Dorsum of head in full-face view with a transverse band of broadly scale-like hairs in front of
the occipital margin and another just behind the level of the frontal lobes (Fig. 4).
(Cameroun) behasyla (p. 286)
- Dorsum of head in full-face view without two bands of broadly scale-like hairs as described
above 31
31 With the head in full-face view the lateral margins behind the level of the eyes with conspicuous
freely projecting hairs (Figs 10,12) 32
- With the head in full-face view the lateral margins behind the level of the eyes without freely
projecting hairs or at most with a single short hair at the scrobal apex; any other hairs present
are strongly curved anteriorly and closely applied to the surface, not freely projecting (Fig. 9) 34
32 Anterior clypeal margin transverse (Fig. 12). Dorsal alitrunk with six or seven pairs of erect
hairs. Base of first gastral tergite with a broad transverse spongiform strip, the basigastral
costulae commencing behind the strip and not impinging upon it. (Ivory Coast, Ghana,
Cameroun, Gabon, Angola) hensekta (p. 293)
- Anterior clypeal margin concave (Fig. 10). Dorsal alitrunk with one or two pairs of erect hairs.
Base of first gastral tergite with a transverse lamellate strip, the basigastral costulae running
across the strip to the basal margin 33
33 Pronotal dorsum punctate. Slightly larger species, HW 0-46-0-54. (Rwanda, Burundi, Kenya,
Tanzania) truncatidens (p. 296)
- Pronotal dorsum finely longitudinally rugulose. Slightly smaller species, HW 0-40. (Came-
roun) dendexa (p. 290)
34 Dorsum of postpetiole finely longitudinally costulate. Infradental lamella of propodeum
reduced to a mere carina on each side. Larger and with broader head, HW 0-54, CI 77
(Fig. 9). (Rwanda) datissa (p. 289)
- Dorsum of postpetiole unsculptured and smooth. Infradental lamella of propodeum
broad and conspicuous. Smaller and with narrower head, HW 0-42-0-46, CI 68-71. (South
Africa) transversa (p. 297)
The species-groups
The 35 known Afrotropical species of Smithistruma are divided into eight species-groups; with
four species in the mandibularis-group , 11 in the emarginata-group, 12 in the weberi-group, two
each in the marginata-, oxysma-, and tacta-groups, and one each in the transversa- and
terrom'-groups.
The mandibularis-group (Figs 1, 2) and emarginata-group (Figs 3-6, 8-13) are very closely
related, the former apparently being directly derived from the latter by reduction of the antennal
segmentation from 6 to 4 and by shortening of the antennal scapes. Most of the characters of
these two groups (see diagnoses) are also shared by the transversa-group but in this last-named
group the basal lamella of the mandible is different in shape and more extensive than in either of
the foregoing groups. In transversa the basal lamella of the mandible is a broadly rounded lobe
which is visible even when the mandibles are completely closed, whereas in both the mandibu-
laris- and emarginata-group^ the lamella is a truncated high triangle or high rectangle which is
concealed by the clypeus when the mandibles are fully closed. Brown (1953«: 125) included
transversa in the emarginata-group but I consider that the difference in structure of the basal
lamella of the mandible is sufficient to exclude it. Reinforcing this decision is the presence of a
broad infradental lamella on the propodeum in transversa, a character not developed in the
emarginata-group (but present in the mandibularis-group) .
On a broader basis the emarginata-group may be cognate with the Holarctic rostrata-group of
Brown (1953a: 81), or with part of it. Until a fuller investigation of the rostrata-group species
from the U.S.A., China and Japan can be made it seems most advisable to keep the groups
separate, especially as rostrata (Emery) itself has a long stout simple hair at each of the pronotal
humeri, a character not encountered in any species of the emarginata-group as defined in this
paper.
The terroni-group , with its single species (Fig. 7), has been derived directly from the
280 BARRY BOLTON
emarginata-group by modification of the mandibular structure. In terroni the mandibular blades
have narrowed and lengthened, opening a long diastema between the basal lamella and the
basalmost tooth, and the number of teeth present has been increased from 12 to 16-17. Apart
from these developments the remaining diagnostic characters conform with those of the
emarginata-group.
The weberi-group constitutes a peculiarly Afrotropical assemblage of striking species which
are immediately recognized by their very coarse heavy sculpture and fine dense simple pilosity
(Figs 14-16). In mandibular structure they resemble the members of the emarginata-group,
having a high truncated basal lamella followed by a row of five principal teeth, two slightly
smaller teeth, four minute denticles and a small apical tooth. However, here the similarity ends
and the members of the weberi-group are not obviously closely related to any other group, either
in Africa or elsewhere.
The oxysma-group (Figs 18, 19), containing two species, has a characteristic clypeal form and
pilosity. The sides of the clypeus are convergent anteriorly and the anterior margin is prominent
and narrowly rounded so that the outer margins of the mandibles and the clypeus form a more or
less continuous line in full-face view. The clypeal dorsum is equipped with feebly clavate hairs
which characteristically curve posteriorly or posteromedially. This form of clypeus approxi-
mates closely to the Nearctic/Neotropical ornata-group (Brown, 1953a: 64), but in the three
known species of this group (ornata (Mayr), dietrichi (M. R. Smith), hyphata Brown) the
mandibles have a long diastema between the basal lamella and the first tooth of the principal
row. In the Afrotropical species oxysma and anarta no such diastema is developed.
The marginata-group contains only the two species marginata and rusta (Fig. 17). The first of
these was included as a member of the now synonymized genus Miccostruma because of its
4-merous antennae and relatively short mandibles. It is now apparent that the the reduction of
antennal segmentation from 6 to 4 has occurred independently three times among Afrotropical
Smithistruma (in the tacta-group, the mandibularis-group, and in marginata), in species that are
otherwise broadly dissimilar, and as a result its value as a genus-level character has disappeared
(see the discussion of the genus, above). The shape of the clypeus in marginata is characteristic
and is not matched by members of the mandibularis- or the tacta-group. Only one other species,
rusta, has a clypeus shaped like that of marginata and so I have grouped them together here. In
both species the lateral margins of the clypeus are more or less straight and parallel and the
anterior margin is broadly and shallowly convex; the clypeus is devoid of hairs both dorsally and
on its margins. For further characters in which the two species coincide, and those in which they
differ, see the diagnosis of the marginata-group.
Finally the tacta-group (Fig. 20), another group having only 4 antennal segments, must be
considered. In clypeal form, structure of the mandibular teeth, presence of long flagellate hairs
on the pronotal humeri, lack of projecting hairs on the leading edges of the scapes, presence of a
median pronotal carina and presence of propodeal infradental lamellae, tacta-group members
resemble the oxysma-group. However, the reduced antennal segmentation and presence of
dense simple clypeal pilosity without specialized hairs argue against its inclusion with oxysma
and for the present it is left on its own.
Key to species-groups (workers)
1 Leading edges of antennal scapes without a series of freely anteriorly projecting strong erect to
suberect hairs (Figs 17-20). Pronotal humeri with a long flagellate hair on each side. Pronotal
dorsum with a median longitudinal ridge or carina at least anteriorly 2
Leading edges of antennal scapes with a series of freely anteriorly projecting strong erect to
suberect hairs which may be simple or bizarre (Figs 1-12, 14, 15). Pronotal humeri without
flagellate hairs. Pronotal dorsum without a median longitudinal ridge or carina 4
2 Clypeus without hairs; in full-face view the clypeal margins lacking fringing pilosity, in profile
the clypeal dorsum without hairs. Anterior clypeal margin broadly and shallowly convex in
full-face view, the sides more or less parallel and not converging anteriorly (Fig. 17)
marginata-group (p. 311)
- Clypeus with hairs; in full-face view the clypeal margins with fringing pilosity or at least with a
THE AFROTROPICAL DACETINE ANTS 281
few hairs projecting; in profile the clypeal dorsum with hairs present. Anterior clypeal margin
narrowly rounded in full-face view, the sides more or less evenly convergent anteriorly
(Figsl8-20) 3
3 Antennae with 6 segments. Clypeus with specialized long recurved hairs present (Figs 18, 19)
oxjsma-group (p. 314)
Antennae with 4 segments. Clypeus without specialized long recurved hairs (Fig. 20)
facfa-group (p. 316)
4 Pronotal dorsum extremely coarsely sculptured with rugae or sulci. With the clypeus in full-face
view the fringing pilosity not consisting of a regular row of curved broad spatulate to
spoon-shaped hairs but rather of irregular long cylindrical simple hairs which may or may not
be curved (Figs 14, 15) wefceri-group (p. 300)
- Pronotal dorsum finely sculptured to smooth, never with coarse rugae or sulci. With the clypeus
in full-face view the fringing pilosity consisting of a regular row of curved broad spatulate to
spoon-shaped hairs (Figs 1-12) 5
5 Mandibles with a long diastema between the basal lamella and the basalmost tooth , the diastema
much longer than the height of the basalmost tooth. 16-17 teeth present, the seventh or eighth
tooth from the base by far the largest (Fig. 7) terroni-group (p. 298)
- Mandibles without a diastema or at most with a minute diastema between the basal lamella and
the basalmost tooth; when present the length of the diastema distinctly much shorter than the
height of the basalmost tooth. 12 teeth present, one of the basal series of 5 the largest
(Figs 1-6, 8-12) 6
6 Basal lamella of mandible an evenly rounded broad lobe which is visible even when the
mandibles are fully closed. Anterior clypeal margin transverse transversa-group (p. 297)
- Basal lamella of mandible either a high triangle which may be truncated apically, or a high
rectangle which may have concave sides; the lamella concealed by the clypeus when the
mandibles are closed. Anterior clypeal margin usually concave, only rarely transverse 7
7 Antennae with 4 segments. Scapes relatively short, SI 50-57. Propodeum with a conspicuous
infradental lamella mandibularis-group(p. 281)
- Antennae with 6 segments. Scapes longer, SI 58-80. Propodeum without or with only a slender
infradental lamella emarginata-group(p. 285)
The mandibularis-group
(Figs 1,2)
Antennae with 4 segments. Basal lamella of mandible a high narrow triangle, usually truncated apically;
tooth row of mandible without or with a minute diastema, the principal tooth row of 5. Sculpture of head
and body fine, without coarse rugae or sulci on the pronotum. Anterior clypeal margin always concave in
full-face view. Lateral and anterior margins of clypeus fringed by a continuous row of projecting flattened
or spoon-shaped large hairs which are smaller on the anterior than on the lateral margins. These hairs are
curved anteriorly on the sides of the clypeus, medially on the anterolateral angles, and are directed
anteriorly or are curved slightly towards the midline on the anterior margin. Body hairs very sparse to
absent. Flagellate hairs absent. Leading edges of scapes with strong anteriorly projecting hairs. Dorsal
(outer) surfaces of middle and hind tibiae without projecting hairs. Pronotum not marginate laterally and
without a median longitudinal ridge or carina on the dorsum. Propodeal spines or teeth subtended by a
broad infradental lamella. Postpetiole in dorsal view with spongiform appendages restricted to a posterior
transverse strip which is broadest at the posterolateral angles; the disc not completely surrounded by
spongiform tissue.
The four closely related species presently recognized in this group appear to be descended
directly from the emarginata-group and share most characters with that group. They differ
primarily by their reduced antennal segmentation, short antennal scapes, development of a
broad infradental lamella on the propodeum and by their drastic reduction of body pilosity,
although this last character is paralleled by the chyatha-complex of the emarginata-group. For
the separation of the mandibularis-group from the remaining species-groups of Africa see the
key to groups above and the diagnoses of the individual groups.
As discussed under the diagnosis of the genus the two previously described members of this
group (mandibularis and tigrilla) constituted two-thirds of the now abandoned genus Miccostru-
ma. For the third species formerly placed in Miccostruma see under the marginata-group , below.
282 BARRY BOLTON
Three of the four species in the group occur in West and Central Africa where they constitute a
part of the leaf litter and topsoil fauna. S. tigrilla, with its distinctive black and yellow transverse
bands and continuous row of basigastral costulae, is known from Ivory Coast and Cameroun,
and most probably also occurs in the territories between these countries. The other two species
occurring in the same area as tigrilla, ninda and fulda are uniformly coloured and have the
basigastral costulae originating on each side of a central gap or clear area. S. fulda has only been
recorded from Ivory Coast, but ninda is much more widely distributed, material having been
seen from Ivory Coast, Ghana, Nigeria, Chad and Cameroun. The only representative of the
mandibularis-group known from East and southern Africa is mandibularis itself, recorded to the
present from Kenya, Tanzania and Angola. Like fulda and ninda it has basigastral costulae
which radiate from each side of a central gap, but unlike them it possesses a pair of standing hairs
on the cephalic dorsum close to the occipital margin. A fifth, as yet undescribed, species
belonging to this group occurs in the Malagasy region.
Smithistrumu fulda sp. n.
(Fig. 1)
HOLOTYPE WORKER. TL 2-0, HL 0-53, HW 0-42, CI 79, ML 0-04, MI 8, SL 0-24, SI 57, PW 0-26, AL 0-54.
Basal lamella of mandible concealed by clypeus, dentition as described for mandibularis. Anterior
clypeal margin very deeply concave medially, the inner margin of the concavity with 5 pairs of scale-like
(inner 3 pairs) to spoon-shaped (outer 2 pairs) hairs which project inward over the mandibles. Anterolater-
al clypeal angles broadly convex on each side of the median impression, equipped with a series of medially
curved spatulate to spoon-shaped hairs which continue along the lateral clypeal margins to about the
midlength of the sides of the clypeus. Dorsum of clypeus with the area behind the marginal concavity
transversely quite deeply depressed, this concave area occupying about the central third of the dorsum and
just over half its total length, from the anterior margin to about the level of the frontal lobes. Areas of the
clypeus on each side of this central concave area are convex, as is the posterior portion of the clypeus
between the frontal lobes. Dorsum of clypeus densely clothed with small flattened to scale-like hairs which
are closely applied to the surface and are directed towards the clypeal midline except on that portion of the
clypeus between the frontal lobes where they are directed anteriorly. Cephalic dorsum with numerous
short spatulate to scale-like hairs which are larger than those on the clypeus and are all closely applied and
directed forward. Dorsum of head without projecting hairs of any description. Antennae with 4 segments.
The scape narrow at the extreme base but then suddenly and very powerfully expanded, flattened and
extremely broad, the leading edge passing through a strongly anteriorly projecting right-angle and
equipped with a series of strong projecting hairs, the longest of which is situated at the apex of the
projection. Eyes small, maximum diameter about 0-lOxHW, markedly smaller than the maximum
diameter of the scape. Dorsum of head reticulate-punctate, the clypeal dorsum finely granular but the
sculpture partially concealed by the pilosity. Pronotum not marginate laterally, the dorsum without a
median longitudinal ridge or carina but the anterior pronotal border weakly marginate. In profile the
dorsal alitrunk consisting of three separate very shallow convexities, the mesonotal slightly higher than the
pronotal or propodeal, but the propodeal anteriorly the most strongly convex. Metanotal groove absent
but the dorsum with a very slight indentation where the mesonotal convexity meets the propodeal.
Propodeal spines not elevated but upcurved along their length, the basal halves of their ventral margins
confluent with the broad infradental lamellae. Alitrunk in dorsal view with the metanotal groove
represented by a transverse line and change in sculpture. Dorsal surfaces of alitrunk and petiole without
standing hairs of any description but the postpetiole with a single pair of simple hairs which are directed
posteriorly. First gastral tergite near base with a pair of very stout appressed hairs which are weakly clavate
apically. It is possible that these hairs should be erect but have been flattened down as an accident of
preservation. Gastral segments behind the first with sparse hairs. Dorsal alitrunk with scattered minute
appressed spatulate hairs. Dorsal surfaces of petiole, postpetiole and first gastral tergite with similar or
even smaller appressed pubescence. Pronotal dorsum finely longitudinally costulate to rugulose. Mesono-
tum finely rugulose anteriorly and punctulate posteriorly, the two forms of sculpture blending together
centrally. Propodeal dorsum unsculptured except for some fine punctures laterally; declivity unsculptured.
Dorsum of petiole node with the faintest vestiges of punctulate sculpture, the postpetiole unsculptured.
First gastral tergite unsculptured except for the basal costulae which radiate from the anterolateral margin
on each side of a median area which is clear. Spongiform appendages of pedicel segments well developed in
profile. In dorsal view the petiole node bordered posteriorly by a broad transverse spongiform strip, the
concave anterior face of the postpetiolar disc bordered by a vestigial lamina. Ventrolateral spongiform
THE AFROTROPICAL DACETINE ANTS 283
tissue of postpetiole does not project beyond the lateral margins of the disc in dorsal view. Convex
posterior margin of postpetiole with an appendage which is broad and spongiform posterolaterally but
narrowed and laminar medially where the posterior margin of the disc itself is flattened. Base of first gastral
tergite with a lamellar transverse margin which is smooth medially but traversed by the strong basigastral
costulae on each side. Colour dark brown to blackish brown, the clypeus and appendages lighter.
Holotype worker, Ivory Coast: Issoneu, 12.x. 1980 (V. Mahnert & J.-L. Ferret) (MHN).
In the mandibularis-group fulda is immediately recognized by the massive angular extension of
the antennal scape, the form and pilosity of the clypeus and the costulate-rugulose pronotal
sculpture. Beside this fulda lacks the conspicuous yellow and black bands oitigrilla and does not
have the pair of erect hairs on the vertex characteristic of mandibularis .
Smithistruma mandibularis (Szabo) comb. n.
Epitritus mandibularis Szabo, 1909: 1, fig. 2. Syntype workers, TANZANIA: Mto-ya-kifaru (K. Katona)
(TM) [examined].
Miccostruma mandibularis (Szabo) Brown, 1948: 123.
WORKER. TL 1-7-1-8, HL 0-46-0-52, HW 0-35-0-41, CI 75-80, ML 0-04-0-06, MI 8-11, SL 0-18-0-21, SI
50-57, PW 0-20-0-23, AL 0-44-0-48 (10 measured).
Mandibles armed with a high narrow-based triangular basal lamella which is truncated apically and
concealed by the clypeus when the mandibles are closed. Basal lamella without or with only a minute
diastema between itself and the principal row of five relatively large teeth, the lamella slightly longer than
the largest of these teeth. Distally the principal tooth row followed by two slightly smaller teeth, a series of
four minute denticles and a small apical tooth. Anterior clypeal margin concave, the concavity here
shallower than in other members of the group, the margin equipped with 3-4 pairs of scale-like hairs which
project over the mandibles. Lateral margins of clypeus convergent anteriorly and equipped with a freely
projecting fringe of large anteriorly curved spatulate to spoon-shaped hairs. Dorsum of clypeus and
cephalic dorsum in full-face view with scattered minute appressed flattened hairs which are directed
anteriorly. In profile the dorsal surface with a single pair of erect feebly clavate hairs which are weakly
curved forward and are situated just behind the highest point of the vertex. Antennae with 4 segments.
Scape narrow basally, bent at about the basal quarter and the anterior border expanded at about this level.
Leading edges of scapes flattened and rounded, expanded but not projecting as a strong lobate or angular
prominence, equipped with a projecting row of spatulate to spoon-shaped strong hairs. Maximum
diameter of eye 0-llxHW. Dorsum of head finely and densely reticulate-punctate to punctate-granular,
the sculpture weaker or effaced on the clypeus. Anterior border of pronotum weakly marginate. Sides of
pronotum not marginate, the dorsum without a median longitudinal ridge or carina. With the alitrunk in
profile the metanotal groove faintly marked, the propodeal teeth with a broad and very conspicuous
infradental lamella. All dorsal surfaces of body with scattered minute appressed pubescence. Alitrunk
without standing hairs but a pair present on the posterior margin of the petiole node, a second pair on the
posterior margin of the postpetiole, a third pair on the base of the first gastral tergite and a fourth pair at the
apex of that sclerite. Remaining gastral tergites with sparse erect hairs. Dorsum of pronotum and
mesonotum with very faint superficial granular or punctulate sculpture, the propodeal dorsum smooth or
with vestigial punctures. Petiole node dorsally with vestigial punctures but disc of postpetiole smooth. First
gastral tergite unsculptured except for the basigastral costulae which arise on each side of a median clear
area. Spongiform appendages of pedicel segments well developed in profile. In dorsal view the petiole
node with a posterior transverse spongiform strip. Anterior margin of postpetiole with a vestigial strip
which is less than half the width of that on the petiole. Posterior margin of postpetiole with a transverse
strip which is broadest laterally, narrowed medially. Base of first gastral tergite with a lamellar strip which
is narrowest medially where its anterior margin is concave, and broadest laterally where its anterior free
margin is convex and traversed by the basigastral costulae. Colour yellow to light brownish yellow.
To the present this is the only species of the group to be found in eastern and southern Africa.
The other three species are more or less restricted to the rain forest zones of West and Central
Africa.
MATERIAL EXAMINED
Kenya: Tana R., Wema (V. Mahnert & J.-L. Ferret); Kilife dist., Jilore (V. Mahnert & J.-L. Ferret);
Lamu, nr Witu (V. Mahnert & J.-L. Ferret); Kisumu, Chemelil (V. Mahnert}. Angola: Salazar (P.
Hammond). Tanzania: Mto-ya-kifaru (K. Katona}.
284 BARRY BOLTON
Smithistruma ninda sp. n.
(Fig. 2)
HOLOTYPE WORKER. TL 1 -8, HL 0-52, HW 0-38, CI 73, ML 0-04, MI 8, SL 0-20, SI 53, PW 0-24, AL 0-48.
Anterior clypeal margin deeply concave medially, the concavity fringed with 4 pairs of scale-like hairs
which project over the mandibles. Sides of clypeus distinctly convergent anteriorly, fringed by a continuous
row of spatulate to spoon-shaped hairs which are curved anteriorly. Mandibles closed in holotype but from
a paratype the dentition consisting of a high narrowly triangular basal lamella which is blunted apically and
distinctly longer than the largest tooth. A minute diastema separates the basal lamella from the principal
row of 5 relatively stout teeth, and these are followed distally by two slightly smaller teeth, 4 minute
denticles and a small apical tooth. Dorsum of clypeus and cephalic dorsum without standing hairs of any
description, with widely scattered and somewhat flattened minute appressed hairs which are directed
anteriorly. Antennae with 4 segments. Scape narrow at base, bent and suddenly broadened in its basal
quarter; the leading edge broadly convex at the bend and equipped with a row of freely projecting spatulate
to spoon-shaped hairs, but the margin not projecting forward into a broad free lobe or strong angle.
Maximum diameter of eye 0-13xHW. Dorsum of head finely and densely reticulate-punctate, the
sculpture becoming finer and more granular anteriorly. Clypeal dorsum granular to merely shagreened.
Pronotum without a median longitudinal ridge or carina dorsally, not marginate laterally, the dorsum
shallowly transversely convex and rounding broadly and evenly into the sides. With the alitrunk in profile
the mesonotum forming a shallow convexity separate from that of the pronotum and propodeum, the
metanotal groove extremely faintly indicated on the dorsum, not impressed in profile. Propodeal teeth
short and triangular, the anterior half or slightly more of the ventral margin confluent with the broad
infradental lamellae. Dorsal surfaces of alitrunk, petiole and first gastral tergite only with very sparse
minute appressed pubescence, without standing hairs of any description. Posterior border of postpetiole
with a single pair of feebly clavate standing hairs and similar hairs are present on the gastral tergites behind
the first. Sides of alitrunk smooth, with a few feeble marginal punctures. Dorsal alitrunk smooth except for
a narrow band of punctures just behind the anterior pronotal margin. The alitrunk frequently with a dull
slightly rough appearance due to a superficial waxy deposit which when removed leaves the surface smooth
and highly polished. Petiole and postpetiole unsculptured dorsally, the first gastral tergite unsculptured
except for the basigastral costulae which arise on each side of a central clear area. Spongiform appendages
of pedicel segments moderately developed in profile. In dorsal view the petiole node with a distinct
spongiform strip posteriorly. Anterior margin of postpetiole with a strip which is about half the width of
that on the petiole. Posterior margin of postpetiole with a spongiform strip which is broadest laterally and
narrowed centrally. Base of first gastral tergite with a laminar strip which is broadest laterally where it is
traversed by the basigastral costulae. Colour dark brown.
PARATYPE WORKERS. TL 1-7-1-8, HL 0-48-0-52, HW 0-37-0-38, CI 73-79, ML 0-03-0-05, MI 7-10, SL
0-18-0-20, SI 50-54, PW 0-22-0-24, AL 0-47-0-50 (9 measured).
As holotype, the maximum diameter of the eye 0- 12-0- 14xHW.
Holotype worker, Cameroun: Nkoemvon, 28.ix.1980, no. N 33 (D. Jackson) (BMNH).
Paratypes. 6 workers with same data as holotype; 3 workers with same data but 6.x. 1980, no. N 34; 1
worker with same data but l.viii.1980, no. N 18 (BMNH; MCZ; MHN; ENSA).
Non-paratypic material examined. Ivory Coast: Gregbeu (V. Mahnert & J.-L. Ferret); Monogaga (V.
Mahnert & J.-L. Ferret). Ghana: Tafo (D. Leston); Mampong (P. M. Room). Nigeria: Gambari (B.
Taylor); Ibadan (B. R. Critchley); Ibadan (A. Russell-Smith). Chad: Umg. Maundou (H. Franz).
The non-paratypic material from Ivory Coast has the alitrunk light brown and the gaster much
darker brown , and has a single pair of stout erect hairs close to the base of the first gastral tergite .
Such hairs are absent in the type-series but are frequently seen in Ghanaian and Nigerian
specimens. The colour of the Ghana material is intermediate between that of the holotype and
the ligher Ivory Coast samples.
Smithistruma tigrilla (Brown) comb. n.
Miccostruma tigrilla Brown, 1973o: 32, figs 1, 2. Holotype worker, IVORY COAST: nr Divo, 18.iii.1963,
berlesate from rain forest leaf litter (L. Brader); and paratype worker, Banco Forest Res., nr Abidjan,
circuit 1, i.1963, berlesate from rain forest leaf litter (W. L. Brown) (MCZ; BMNH) [examined].
WORKER. TL 2-0-2-2, HL 0-56-0-62, HW 0-48-0-51, CI 82-88, ML 0-05-0-06, MI 9-11, SL 0-23-0-26, SI
48-52, PW 0-30-0-32, AL 0-55-0-62 (4 measured).
THE AFROTROPICAL DACETINE ANTS 285
Dentition of mandibles apparently as described under mandibularis . Anterior clypeal margin very
broadly and deeply arched-concave, the excavation semicircular in full-face view and the concave margin
with 5 pairs of projecting scale-like to spatulate small hairs which are curved medially. Sides of clypeus
weakly convergent anteriorly, equipped with a fringe of freely projecting large spatulate to spoon-shaped
hairs which are curved anteriorly. Because of the width and depth of the clypeal concavity the anterolateral
angles seem narrow and strongly prominent anteriorly. Dorsum of clypeus and cephalic dorsum with
widely scattered decumbent to appressed anteriorly directed minute flattened hairs which are very
inconspicuous; without standing hairs of any description. Antennae with 4 segments. Scape narrow basally
but then the leading edge suddenly broadened into a large anteriorly prominent rounded lobe. Leading
edges of scape with a row of large freely projecting spatulate to spoon-shaped hairs. Maximum diameter of
eye O14-16xHW. Cephalic dorsum densely and quite sharply reticulate-punctate, the posterior clypeus
similarly but more finely sculptured, the sculpture tending to fade out towards the anterior clypeal margin.
Pronotal dorsum more or less flat transversely, the dorsum meeting the sides in a bluntly rounded angle.
Pronotum without a median longitudinal ridge or carina. With alitrunk in profile the mesonotum forming a
shallow convexity which is separate from the pronotum and propodeum. Metanotal groove extremely
feebly marked by an impression, its location more obviously indicated by a change of colour. Propodeal
teeth without any portion which is free of the infradental lamella. Dorsal surfaces of alitrunk, petiole,
postpetiole and first gastral tergite without standing hairs of any description, only with minute appressed
slightly flattened pubescence which is very sparse. Gastral tergites behind the first with weakly clavate
hairs. Sides of alitrunk mostly smooth, with marginal punctation. Dorsal surfaces of alitrunk, petiole and
postpetiole finely and densely reticulate-punctate to granular, the postpetiole also with fine longitudinal
costulae or rugulae at least on the anterior half of the disc. Spongiform appendages of pedicel segments
only moderately developed in profile, the ventral appendage of the petiole represented only by a small
posteroventral lobe below the node. In dorsal view the petiole node with a narrow posterior strip and the
postpetiole with an anterior strip of about the same width or even narrower. Posterior margin of
postpetiole with a spongiform strip which is broadest laterally and very narrow or even interrupted
medially. Posteromedian area of postpetiole disc impressed. Base of first gastral tergite with a narrow
spongiform strip which is concave anteromedially. Basigastral costulae arising right across the base of the
first tergite, without a broad central gap. Mandibles, clypeus and antennae yellow; remainder of head
black. Pronotum, mesonotum and forelegs yellow; propodeum, pleurae, middle and hind coxae black.
Middle and hind femora dusky at least basally, remainder of legs yellow. Petiole and postpetiole black.
Basal third of first gastral tergite yellow or yellowish white, the rest of the gaster black.
Rendered very distinctive by its conspicuous black and yellow colour pattern, tigrilla is also
characterized by its lack of dorsal pilosity, rugulose-costulate postpetiolar dorsum, basigastral
costulae which arise in a continuous row without a central clear area, and evenly sculptured
dorsal alitrunk.
MATERIAL EXAMINED
Ivory Coast: nr Divo (L. Brader). Cameroun: Korup Res. (D. Jackson).
The emarginate-group
(Figs 3-6, 8-13)
Antennae with 6 segments. Basal lamella of mandible a high triangle which is narrowly truncated apically,
or a concave-sided high rectangle; never a low rounded lobe; never with a marked diastema. Principal
dental row of mandible with 5 teeth. Sculpture of head and body fine, without coarse rugae or sulci on the
pronotum. Anterior clypeal margin usually concave in full-face view, rarely transverse and never convex.
Lateral and anterior margins of the clypeus fringed by a continuous row of large projecting specialized hairs
which are usually flattened, spatulate or spoon-shaped and which are very conspicuous, curving anteriorly
on the sides, medially on the anterolateral angles and towards the midline on the anterior clypeal margin.
Pilosity of head behind clypeus very variable but never consisting solely of fine simple hairs. Frequently
very few hairs are present and sometimes none. Flagellate hairs absent from head and alitrunk. Leading
edges of scapes with anteriorly projecting stout or bizarre specialized hairs. Dorsal (outer) margins of
middle and hind tibiae lacking projecting hairs, any hairs which do occur here are decumbent to appressed.
Pronotum not sharply marginate laterally and without a median longitudinal ridge or carina. Propodeum
usually without an infradental lamella but sometimes a very narrow to vestigial lamella present.
286 BARRY BOLTON
Eleven species are currently recognized in this group, falling into three complexes of closely
related forms.
The chyatha-complex (Figs 3-6), containing behasyla, cavinasis, chyatha, impidora and
sharra, is characterized by the presence of exceptionally fine and dense pronotal sculpture which
usually appears as minute close-packed longitudinal striolae or rugulae upon a granular surface.
Coupled with this the head is usually granular, erect hairs are absent from the dorsal alitrunk,
and in most suborbicular or very broadly scale-like hairs are developed on the head. These hairs
are best developed in cavinasis and sharra, in which the whole head is covered with them; they
are present in transverse bands in chyatha and behasyla. In impidora such hairs are absent but it
is not known whether this condition is basal for the chyatha-complex as a whole or represents a
stage where these specialized hairs have been secondarily lost. However, in chyatha and
impidora the head is strongly dorsoventrally flattened, which is certainly a secondary adapta-
tion, and the sequence sharra-behasyla-chyatha-impidora, showing increased flattening of the
head and gradual disappearance of the specialized hairs, implies that the second alternative is
most probably correct.
This complex, all the members of which are restricted to West and Central Africa, appears to
be the stock from which the terroni-group is descended. Apart from the modified mandibles of
terroni its overall resemblance to the members of the chyatha-complex is striking.
The truncatidens-complex (Figs 9-13), containing datissa, dendexa, gatuda, hensekta and
truncatidens , is defined by the predominance of punctate sculpture on the head and alitrunk,
presence of erect hairs on the dorsal alitrunk and lack of broadly scale-like or suborbicular hairs.
The head in profile is depressed anteriorly but strongly thickened at the vertex. Members of this
complex approach the rostrata-group, as noted in the discussion of the species-groups, and also
seem to represent the stock from which both the mandibularis-group and the transversa-group
developed. This complex tends to be more widely distributed than the above, with a preponder-
ance of species in East Africa.
The emarginata-complex (Fig. 8), containing only emarginata itself, is defined by its abundant
spoon-shaped cephalic pilosity and basigastral costulae that radiate from each side of a median
broad indentation on the first tergite. The species is perhaps the most successful member of the
genus in Africa, being found to the present in Ivory Coast, Ghana, Togo, Burundi, Zimbabwe,
Angola and South Africa.
Smithistruma behasyla sp. n.
(Fig. 4) '
HOLOTYPE WORKER. TL 2-5, HL 0-67, HW 0-42, CI 63, ML 0-11, MI 16, SL 0-28, SI 67, PW 0-27, AL 0-68.
Mandibular dentition as described for cavinasis. Anterior clypeal margin very shallowly evenly concave,
equipped with a row of broad short flattened hairs which project forward over the mandibles. Anterolater-
al clypeal angles rounded, the sides feebly divergent posteriorly and with a continuous row of anteriorly
curved large spatulate to spoon-shaped hairs. In full-face view the preocular laminae feebly convergent
posteriorly. Upper scrobe margins divergent posteriorly and with a row of anteriorly directed scale-like to
broadly spoon-shaped hairs which are strongly curved. Occipital margin deeply evenly concave. Clypeal
dorsum in full-face view with minute appressed stubble-like ground-pilosity, the individual hairs widely
scattered. Cephalic dorsum just behind the level of the frontal lobes with a transverse band of broadly
scale-like to suborbicular hairs. Behind this band the head only with stubble-like ground-pilosity like that
on the clypeus but the zone between the highest point of the vertex and the occipital margin with a second
transverse band of broadly scale-like hairs. Dorsum of head without simple fine hairs, without flagellate
hairs. Scape bent in the basal third, somewhat flattened and broadest just distal to the bend, the leading
edge equipped with a row of freely projecting spatulate to narrowly spoon-shaped hairs, the longest of
which occurs at the bend of the scape. Dorsum of head finely and densely punctate, with a granular
appearance; clypeal dorsum similarly but less strongly sculptured. Anterior border of pronotum margin-
ate, the sides not marginate and without a median longitudinal ridge or carina dorsally. Metanotal groove
represented by a faint transverse line on the dorsum, not impressed. Outline of dorsal alitrunk in profile
with the pronotum and anterior part of the mesonotum sloping upwards to the highest point, which is
shallowly convex; the posterior portion of the mesonotum and the propodeum forming a single extremely
shallowly concave surface which is weakly sloped posteriorly. Propodeal teeth triangular and acute, the
THE AFROTROPICAL DACETINE ANTS 287
infradental lamella represented only by a narrow concave crest down each side below the teeth. Sides of
alitrunk weakly superficially punctate, densest on the mesopleuron, the metapleuron almost smooth.
Pronotal dorsum with extremely fine superficial but quite dense scratch-like longitudinal striation and with
a few scattered punctures. Mesonotum anteriorly sculptured as pronotum but posteriorly only weakly
punctate. Propodeal dorsum almost smooth, with only the faintest vestiges of sculpture. Dorsal alitrunk
without standing hairs, flagellate hairs or any form of specialized pilosity, only with sparse appressed
minute ground-pilosity. Spongiform appendages of pedicel segments strongly developed in profile, the
subpetiolar process curtain-like and with a deep indentation at about its midlength. Postpetiolar ventral
appendage large and lobate. Dorsum of petiole node superficially very shallowly punctate, the posterior
spongiform strip lamellate. Postpetiole dorsum smooth, its posterior spongiform strip broadly and
shallowly indented medially. Basal spongiform strip of first gastral tergite narrow but dense, not traversed
by the basigastral costulae, the latter, however, are sharply defined on the tergite behind the spongiform
tissue. Petiole and postpetiole dorsally with appressed short very narrowly spatulate hairs, the posterior
margins of each segment with one or two pairs of much larger spatulate hairs which project backwards over
the spongiform material. First gastral tergite with fine appressed very sparse ground-pilosity, and with two
pairs of longer stout hairs. The first, basally situated pair are erect or nearly so, the second pair, situated
close to the apical margin of the tergite, are subdecumbent. Colour light brown.
PARATYPE WORKER. TL 2-5, HL 0-67, HW 042, CI 63, ML 0-11, MI 16, SL 0-28, SI 67, PW 0-27, AL 0-68.
As holotype.
Holotype worker, Cameroun: nr Yaounde, sample 1768 (G. Tenon) (ENSA).
Paratype. 1 worker with same data as holotype (BMNH).
In the emarginata-group four out of the five members of the chyatha-complex have broadly
scale-like to suborbicular hairs on the head. In cavinasis and sharra such hairs are evenly
distributed over the surface (Fig. 3). In chyatha the hairs are restricted to a single transverse
band just in front of the occipital margin (Fig. 5), but in behasyla they are arranged in two
transverse bands, one close to the occiput as in chyatha and another situated just behind the level
of the frontal lobes (Fig. 4).
Smithistruma cavinasis Brown
Smithistrwna (Smithistruma) cavinasis Brown, 19506: 42. Holotype worker, ZAIRE: Ituri Forest between
Beni and Irumu, ii.1948, no. 2129 (N. A. Weber) (AMNH) [examined]. [See also Brown, 1953a: 129.]
WORKER. TL 1-9-2-1, HO 0-52-0-56, HW 0-34-0-37, CI 63-67, ML 0-04-0-06, MI 7-10, SL 0-22-0-26, SI
63-70, PW 0-23-0-25, AL 0-50-0-56 (10 measured).
Mandibular dentition of 5 large teeth following the basal lamella without a diastema, then two slightly
smaller teeth and a series of 4 small denticles before the apical tooth. Anterior clypeal margin broadly
deeply and evenly concave, the concavity including the whole of the anterior margin except for the
anterolateral corners. Lateral margins of clypeus convergent anteriorly and fringed with a continuous row
of large flattened spatulate to roughly spoon-shaped projecting hairs which are curved anteriorly. Anterior
clypeal margin with a row of 6 broadly scale-like to suborbicular hairs which project out over the
mandibles. Dorsum of clypeus and of head with numerous broadly scale-like to suborbicular hairs, densest
on the clypeus; such hairs also fringing the lateral borders of the head in full-face view. Flagellate hairs or
other pilosity absent. Preocular laminae broad in full-face view and somewhat divergent anteriorly.
Antennal scapes narrow basally, bent at about the basal quarter and suddenly broadened, broadest at
about this level and the leading edge bluntly subangulate. Dorsal surface of scape with scale-like hairs but
leading edge with a series of freely projecting longer narrower hairs, the longest of which occurs at the
angle. Eyes of moderate size, 0-ll-0-14xHW, the diameter less than the maximum width of the scape.
Dorsum of head finely and densely reticulate-punctate, with a granular appearance. Dorsal alitrunk with
scattered but conspicuous scale-like to suborbicular hairs, smaller versions of which also occur on the
petiole dorsum but which are sparse or absent from the postpetiolar disc and absent from the gaster.
Elongate simple hairs absent from alitrunk but present on the petiole (1 pair), postpetiole (2-3 pairs) and
base of the first gastral tergite (usually 1-2 pairs but sometimes 3 pairs). Flagellate hairs absent. Alitrunk
not marginate laterally, the pronotum without a median longitudinal ridge or carina dorsally. With the
alitrunk in profile the mesonotum very slightly raised above the level of the pronotum and propodeum.
Metanotal groove not impressed but its site marked by a small step-down from the mesonotal to the
propodeal dorsum. Propodeal teeth strong and broad basally, the infradental lamellae very narrow and
with a distinctly concave outline. Mesopleuron reticulate-punctate, the remainder of the sides of the
288
BARRY BOLTON
alitrunk unsculptured or only with faint superficial sculpture. Pronotal dorsum finely superficially
longitudinally striolate to feebly rugulose , the remainder of the dorsum and the petiole dorsum punctulate .
Disc of postpetiole smooth and first gastral tergite unsculptured except for the basigastral costulae.
Spongiform appendages massively developed in profile. In dorsal view the sides and posterior margin of
the petiole node surrounded by continuous thick spongiform material. The postpetiole with an anterior
spongiform transverse strip and with the lateral spongiform material projecting beyond the sides of the disc
in dorsal view. Posterior margin of postpetiole disc with a continuous broad spongiform strip which is
slightly narrower centrally than at the sides. Base of first gastral tergite with a transverse spongiform band
as wide as that on the posterior margin of the postpetiole , and like the postpetiolar strip this is also broadest
at the sides and narrow centrally. Colour medium to light brown.
One of only two species in the emarginata-group to show broadly scale-like to suborbicular hairs
all over the head, cavinasis shares this character with sharra. Workers of the two species are
separated as follows.
cavinasis
Head absolutely and relatively
shorter, HL 0-52-0-56, CI 63-67.
Antennal scapes absolutely and
relatively shorter, SL 0-22-0-26,
SI 63-70.
Posterior margin of postpetiole disc
without a row of spatulate to
squamate hairs on each side of
the midline.
Simple elongate hairs present on the
postpetiolar disc.
Simple elongate hairs present on the
basal portion of the first gastral
tergite.
sharra
Head absolutely and relatively longer,
HL 0-58-0-64, CI 56-63.
Antennal scapes absolutely and
relatively longer, SL 0-26-0-28.
SI 73-78.
Posterior margin of postpetiolar disc
with a row of 5-6 spatulate to
squamate hairs on each side of the
midline which project backward over
the spongiform strip.
Simple elongate hairs absent from the
postpetiolar disc.
Simple elongate hairs absent from the
basal portion of the first gastral
tergite.
MATERIAL EXAMINED
Cameroun: Nkoemvon (D. Jackson); nr. Yaounde (G. Tenon). Zaire: Ituri Forest (N. A. Weber).
Angola: Dundo (L. de Carvalho).
Smithistruma chyatha sp. n.
(Fig. 5)
HOLOTYPE WORKER. TL 2-1, HL 0-62, HW 0-39, CI 63, ML 0-08, MI 13, SL 0-25, SI 64, PW 0-24, AL 0-59.
Mandibles with 5 relatively large teeth followed by two slightly smaller teeth, 4 denticles and an apical
small tooth. Basal lamella concealed by clypeus. Anterior clypeal margin broadly and evenly concave,
equipped with a row of 7 scale-like hairs which project forward over the mandibles. Of the seven the three
central hairs are the smallest, the next one on each side is slightly larger and the outermost on each side
(closest to the anterolateral angle) is much larger, transitional in size and shape to the continuous fringe of
spatulate anteriorly curved long hairs which project from the lateral clypeal margins. Upper scrobe
margins with a single row of adherent suborbicular hairs. Dorsum of head between highest point of vertex
and occipital margin with suborbicular hairs present in a transverse band. Remainder of cephalic dorsum
and clypeal dorsum without hairs, equipped only with minute pubescence which is somewhat flattened and
is only visible under very high magnification. Flagellate hairs absent. Lateral margins of clypeus shallowly
convex and convergent anteriorly, the preocular laminae slightly divergent anteriorly in full-face view.
Antennal scapes bent very close to the base, broadest at about the point of maximum curvature, with the
leading edge broadly rounded and equipped with a series of projecting flattened hairs, the longest of which
is about at the point of maximum scape width. Dorsum of scape just behind the leading edge with a sparse
row of suborbicular hairs. Maximum diameter of eye about 0- 13 x HW, less than the maximum width of the
scape. Head very conspicuously dorsoventrally flattened, in profile the ventral surface not strongly convex
posteriorly and the dorsum only shallowly convex at the vertex. Sides of alitrunk not marginate, the
pronotum without a median longitudinal ridge or carina dorsally. Metanotal groove absent. Propodeal
teeth strong, broad basally and upcurved at the tips. Dorsal alitrunk without flagellate or simple hairs of
any description, only with very widely scattered extremely small inconspicuous short flattened hairs,
THE AFROTROPICAL DACETINE ANTS 289
appearing hairless under low magnification. Petiole node with a few small flattened hairs on the dorsum but
fringed posteriorly and down the sides, in dorsal view, by a row of larger spatulate to squamate hairs.
Lateral and posterior margins of postpetiole fringed with a similar row of spatulate to squamate hairs
which project over the spongiform tissue. Dorsum of postpetiole and first gastral tergite only with minute
pubescence, without hairs of any description. Spongiform appendages of pedicel segments massively
developed in profile. In dorsal view the petiole node bounded posteriorly by a transverse spongiform strip
which continues down the sides of the node posterolaterally, the lateral margins of the node in front of this
with a few decumbent spatulate hairs but without spongiform tissue. Disc of postpetiole in dorsal view
surrounded by spongiform tissue, with a transverse strip on the shallowly concave anterior margin and a
broader spongiform strip on the convex posterior margin which is narrowed centrally. The spongiform
material visible at the sides of the postpetiole disc is narrower and lower than the anterior and posterior
strips but can be seen projecting beyond the lateral margins throughout their length. Base of first gastral
tergite with a transverse spongiform strip which is overlapped by that on the posterior margin of the
postpetiole and which is not narrowed centrally. Pronotal dorsum with dense but very fine low superficial
longitudinal rugulae. Mesonotum superficially punctulate. Propodeal dorsum mostly smooth, with
scattered small punctulae. Mesopleuron finely punctulat, metapleuron smooth but the sides of the
propodeum with scattered quite large punctures. Petiole node faintly punctulate dorsally, the postpetiole
smooth and shining. First gastral tergite unsculptured except for the short widely spaced basigastral
costulae.
Holotype worker, Ghana: Tafo, 15.x. 1970, cocoa leaf litter (B. Boltori) (BMNH).
Along with impidora in the emarginata-group chyatha shares the characters of strongly reduced
pilosity and markedly dorsoventrally flattened head, where the maximum depth of the head is
0-60xHW and the posteroventral convexity of the surface is vestigial. The retention by chyatha
of some suborbicular hairs on the cephalic dorsum links this species with behasyla, sharra and
cavinasis, where these specialized hairs are much better developed. In the worker chyatha and
impidora are easily separated as the former has a row of adherent suborbicular hairs lining the
upper scrobe margins and has a transverse band of suborbicular hairs on the cephalic dorsum
between the vertex and the occipital margin. Such suborbicular hairs are absent in impidora.
Smithistruma datissa sp. n.
(Fig. 9)
HOLOTYPE WORKER. TL 2-7, HL 0-70, HW 0-54, CI 77, ML 0-12, MI 17, SL 0-34, SI 63, PW 0-34, AL 0-74.
Mandibles armed with 5 relatively large teeth following the basal lamella (which is concealed by the
clypeus in the holotype). Distal to the 5 principal teeth are 2 slightly smaller teeth, a row of four denticles
and a small apical tooth. Anterior clypeal margin broadly and evenly concave between the anterolateral
corners, the margin equipped with 10 scale-like hairs which project forward over the mandibles. These
hairs become gradually larger away from the midline but the outermost, at the anterolateral corner, is
much the largest and forms an intermediate between the shorter hairs on the anterior margin and the large
spatulate to spoon-shaped anteriorly curved hairs which form a fringe on the lateral clypeal margins. In
full-face view the preocular laminae slightly divergent anteriorly. Clypeal dorsum more or less smooth
centrally but feebly sculptured laterally and anteriorly. Cephalic dorsum densely shallowly reticulate-
punctate everywhere. Dorsum of head with numerous small, widely spaced flattened hairs which are
subdecumbent to decumbent and are mostly directed anteriorly. In full-face view the sides of the head with
a few such hairs projecting, curved anteriorly, most conspicuous on the sides of the occipital lobes.
Flagellate hairs absent. Antennal scapes narrow basally, shallowly bent at about the basal third and
broadest at this point . The leading edges of the scapes evenly curved at the bend and equipped with a row of
projecting spatulate to broadly clavate hairs and an interspersed row of shorter much finer simple hairs.
Maximum diameter of eye about 0-13xHW, approximately equal to the maximum width of the scape.
Head in profile roughly wedge-shaped, the vertex forming a high narrowly rounded convexity, the ventral
surface evenly shallowly convex. Pronotal dorsum without a median longitudinal ridge or carina, not
sharply marginate laterally. Alitrunk without flagellate hairs. With the alitrunk in profile the posterior part
of the pronotum and anteriormost section of the mesonotum raised into a broad shallowly convex tumulus,
the remainder of the mesonotum and the propodeum, which form a single surface without trace of a
metanotal groove, markedly depressed below the level of this tumulus. Propodeal teeth short and broadly
triangular, the infradental lamellae narrow but clearly visible. Dorsal alitrunk with scattered short
flattened hairs which are decumbent to appressed, without standing pilosity of any description. Petiole and
290 BARRY BOLTON
postpetiole dorsally with similar but extremely sparse hairs, the latter also with 4 clavate suberect hairs
projecting from the posterior margin. In the holotype a single clavate hair is also present on the left side of
the postpetiolar disc, appressed to the surface; this is not matched in the paratypes. First gastral tergite with
numerous suberect to erect stout hairs which are simple to weakly clavate apically. Pronotal dorsum
predominantly broadly reticulate-punctate but anteriorly and laterally the margins of the punctures
tending to run together and form very fine rugulae. Mesonotum and propodeal dorsum more sharply
punctate, the sculpture on the latter running between the propodeal teeth and ending about halfway down
the declivity. Petiole dorsum minutely rugulose, the disc of the postpeiiole finely longitudinally costulate.
Basigastral costulae sharply developed and conspicuous, the tergite otherwise unsculptured. Alitrunk
pleurae mostly smooth centrally but punctulate marginally, a line of punctures separating meso- and
metapleuron and a relatively densely punctured patch on the mesopleuron behind the upper half of the
front coxa. Spongiform appendages of pedicel segments moderately large in profile. In dorsal view the
petiole node with a posterior spongiform strip which is continued down the sides. Postpetiole with a narrow
spongiform strip anteriorly and a broader strip bordering the posterior margin which is slightly narrowed
medially. Colour dark brown.
PARATYPE WORKERS. TL 2-6-2-7, HL 0-70, HW 0-54, CI 77, ML 0-12, MI 17, SL 0-34, SI 63, PW 0-32-0-34,
AL 0-73-0-74 (2 measured).
As holotype but maximum diameter of eye 0-13-0- 15 xHW.
Holotype worker, Rwanda: Rangiro, ix.1976 (P. Werner) (MHN).
Paratypes. 2 workers with same data as holotype (BMNH; MCZ).
The truncatidens-complex of this group contains five species. Four of these, gatuda, hensekta,
dendexa and truncatidens , possess conspicuously projecting hairs on the sides of the occipital
lobes. In datissa, however, such projecting hairs are absent, any hairs which occur on the sides of
the occipital lobes being small and curved, usually closely adherent to the head and not freely
projecting. 5. datissa is also separated from gatuda, dendexa and hensekta by having the
postpetiolar disc longitudinally costulate; it is glassy smooth in the last three named. A few
specimens of truncatidens do show costulae either on part or all of the postpetiole but here the
mesonotum has a pair of long erect hairs which are not seen in datissa.
Smithistruma dendexa sp. n.
HOLOTYPE WORKER. TL 2- 1 , HL 0-58, HW 0-41 , CI 71 , ML 0-08. MI 14, SL 0-26, SI 63, PW 0-26, AL 0-58.
Mandibles closed but dentition apparently as described for truncatidens, certainly with 5 large teeth
basally, the fourth of which, counting from the base, is the smallest, about the same size as teeth six and
seven. Anterior clypeal margin broadly shallowly concave, with a row of small scale-like hairs which
project over the mandibles. Lateral margins of clypeus slightly convergent anteriorly, the preocular
laminae slightly convergent posteriorly in full-face view. Lateral margins of clypeus equipped with a row of
freely projecting anteriorly curved large spatulate to spoon-shaped hairs. Upper scrobe margins divergent
posteriorly, with projecting anteriorly curved spatulate to narrowly spoon-shaped hairs which are not as
dense nor as broad as those on the lateral margins of the clypeus, and which mostly also curve upwards from
their points of origin. Clypeal dorsum and cephalic dorsum with numerous small flattened hairs which are
curved anteriorly and appear scale-like in full-face view. Antennal scape bent at the basal third, broadest at
or just distal to the bend. Leading edge of scape with a row of freely projecting long stout hairs. Head
without flagellate hairs. Dorsum of head densely punctulate and matt, the clypeal dorsum less strongly
sculptured. Pronotum not marginate laterally, without a median longitudinal ridge or carina. Metanotal
groove absent. With the alitrunk in profile the anterior half of the mesonotum forming the highest point of
the outline as a low tumulus. Posterior half of mesonotum and propodeum forming a single feebly sinuate
surface. Propodeal teeth triangular and acute, weakly elevated and with a narrow infradental lamella.
Dorsal alitrunk with a number of appressed short fine hairs. Large erect hairs on alitrunk restricted to a
single long stout pair which are weakly clavate apically and situated on the raised anterior portion of the
mesonotum. Without other standing pilosity on the alitrunk and without flagellate hairs. Dorsal surfaces of
petiole, postpetiole and first gastral tergite with numerous suberect to erect stout hairs which are feebly
clavate apically. Sides of alitrunk mostly smooth but mesopleuron punctate anteriorly and sides of
propodeum punctate. Pronotal dorsum finely longitudinally rugulose, without punctate sculpture; meso-
notum and propodeal dorsum densely reticulate-punctate. Dorsum of petiole node punctate, the post-
petiolar disc unsculptured. First gastral tergite unsculptured except for the sharply defined basal costulae.
Spongiform appendages of pedicel segments strongly developed in profile. In dorsal view the petiole node
THE AFROTROPICAL DACETINE ANTS 291
with a narrow lamellar spongiform strip posteriorly and a similar but narrower strip on the anterior margin
of the postpetiole. Sides of postpetiole without projecting spongiform material in dorsal view but the
posterior margin with a transverse strip which is broad posterolaterally but considerably narrowed
medially. Base of first gastral tergite with a transverse lamellar strip which is traversed by the basigastral
costulae. Colour dull yellow to light yellowish brown.
PARATYPE WORKER. TL 2- 1 , HL 0-60, HW 0-42, CI 70, ML 0-08, MI 13, SL 0-27, SI 64, PW 0-28, AL 0-58.
Asholotype.
Holotype worker, Cameroun: nr Yaounde, sample 1779 (G. Terron) (ENSA).
Paratype. 1 worker with same data as holotype (BMNH).
5. dendexa is very closely related to truncatidens , an East African species known from Rwanda,
Burundi, Kenya and Tanzania. The two species agree in most diagnostic characters but dendexa
is smaller (compare measurements) and has fine longitudinal rugulae on the pronotal surface,
without punctures. In truncatidens the pronotum is usually punctate but at most only a few feeble
scattered rugulae occur, due to alignment of the walls of adjacent punctures.
Smithistruma emarginata (Mayr)
(Fig. 8)
Strumigenys emarginata Mayr, 1901: 26. Syntype workers, SOUTH AFRICA: Port Elizabeth (H. Brauns)
(NMV) [examined].
Smithistruma (Smithistruma) emarginata (Mayr) Brown, 1948: 105; 1953a: 126.
WORKER. TL 2-4-2-8, HL 0-64-0-70, HW 0-39-0-42, CI 58-64, ML 0-11-0-12, MI 16-19, SL 0-29-0-32, SI
72-80, PW 0-24-0-28, AL 0-62-0-72 (18 measured).
Mandibles with a high truncated triangular basal lamella (concealed by clypeus when mandibles are
closed), followed without a diastema by a row of 5 relatively large teeth, 2 slightly smaller teeth and 4 small
denticles before the apical tooth. Anterior clypeal margin in full-face view varying from almost transverse
to evenly shallowly concave. Lateral margins of clypeus slightly convergent anteriorly, the anterolateral
clypeal angles bluntly rounded. Anterior clypeal margin fringed by a series (usually of 6-8) broad scale-like
hairs, the lateral margins and corners with an unbroken sequence of long fringing hairs which are flattened
to spoon-shaped and which are curved anteriorly on the sides and medially on the anterolateral corners.
Dorsum of clypeus and of head behind clypeus with numerous spoon-shaped curved hairs which appear
scale-like in full-face view. Sometimes the occipital region with a few simple curved hairs present but these
variable in number and degree of development; flagellate hairs never developed. With the head in full-face
view the upper scrobe margins and occipital lobes laterally fringed with anteriorly curved spoon-shaped
hairs, the head long and narrow (CI <65) and with the eyes plainly visible, projecting beyond the level of
the upper scrobe margins. Eyes larger than in any other other known Afrotropical species, their maximum
diameter 0-21-0-25xHW, greater than the maximum width of the scape. Scapes long (SI >70), narrow
basally, shallowly curved at about the basal third and broadest just distal to this where the leading edge is
bluntly subangulate. Leading edges of scapes with projecting flattened to spoon-shaped strong hairs. With
the head in profile the dorsum very shallowly impressed between clypeus and vertex, highest at the vertex
and sloping down posteriorly to the occipital margin. Dorsum of head finely and densely reticulate-
punctulate to granular everywhere. With the alitrunk in profile the central portion of the mesonotum
extremely feebly impressed. The metanotal groove not impressed but sometimes represented as a line.
Propodeal teeth long and narrow, often slightly upcurved and sometimes weakly sinuate along their length.
Infradental lamellae narrow and inconspicuous down the propodeal declivity. Sides of alitrunk not
marginate, the pronotal dorsum without a median longitudinal ridge or carina, the pronotal humeri evenly
rounded. Pilosity of dorsal alitrunk variable, usually with curved spoon-shaped hairs on pronotum and
anterior mesonotum but behind this the hairs longer and finer, subspatulate to cylindrical and simple, and
often with one or two pairs suberect to erect. Variation from this more or less median position is shown on
the one hand in samples where all the hairs are spoon-shaped and merely vary in size (becoming larger
posteriorly), there being no subspatulate or simple hairs developed; and on the other hand by the
suppression of the spoon-shaped hairs and their replacement everywhere by simple suberect to erect
pilosity. Flagellate hairs never present. Pronotal dorsum very finely and faintly striate, this sculpture
sometimes virtually effaced. Mesonotum and usually also propodeal dorsum finely punctulate; sides of
alitrunk punctulate. Spongiform appendages of petiole and postpetiole massively developed in profile. In
dorsal view the petiole with a spongiform strip on its posterior margin which is strongest posterolaterally.
292 BARRY BOLTON
Anterior margin of postpetiole in dorsal view with a spongiform strip but the sides without. The broadly
convex posterior margin of the postpetiole with spongiform tissue very broadly developed at the sides but
strongly indented or even interrupted medially, usually the posterior margin of the spongiform material
touching the margin of the postpetiolar disc centrally. Petiole dorsum very faintly punctulate to smooth,
the disc of the postpetiole always unsculptured and smooth. Dorsal surfaces of petiole, postpetiole and first
gastral tergite with elongate simple curved hairs present. First gastral tergite impressed mediobasally,
usually sharply so, the impressed area usually including both the central portion of the basal lamellar band
of the tergite and the tergal area immediately behind it. Basigastral costulae absent from the impressed
area, radiating from each side of it; gaster otherwise unsculptured. Colour yellow to medium brown,
sometimes the gaster distinctly darker than the head and alitrunk.
Within its species-group emarginata stands very much alone, lacking the pilosity and other
characters of the various species-complexes discussed under the species-group diagnosis, but
possessing an elongate narrow head, long scapes and a basally indented first gastral tergite
coupled with the largest eyes known for a member of this genus in the Afrotropical region.
Despite the wide range of the species it shows relatively little variation, the only notable
changes occurring in the form of the alitrunk pilosity as discussed above. Brown (1953a: 126)
first drew attention to this but noted that, even though his material was sparse, intergrades
between the forms were apparently present. This study has confirmed that alitrunk pilosity is by
no means stable in emarginata and, like Brown, I am of the opinion that the differences observed
only represent variation between populations and are not significant at species-level.
MATERIAL EXAMINED
Ivory Coast: Lamto, Toumodi (J. Levieux). Ghana: Mampong (P. Room). Togo: Palime, Kpime Forest
(Vif). Burundi: Imbo Plain (A. Dejeari). Zimbabwe: Sawmills (G. Arnold), Gwebi (K. J. Wilson);
Chishawasha (A. Watsham). Angola: Dundo, Luachino (Machado). South Africa: Cape Prov., Port
Elizabeth (H. Brauns); Algoa Bay (H. Brauns); Grahamstown (L. Weatherill); Natal, Zululand, Richard's
Bay (/. C. Faure); St Lucia Lake, Bird Island (/. C. Faure).
Smithistruma gatuda sp. n.
(Fig. 11)
HOLOTYPE WORKER. TL 2-2, HL 0-59, HW 0-42, CI 71 , ML 0-06, MI 10, SL 0-28, SI 67, PW 0-27, AL 0-60.
Mandibles with 5 relatively large teeth followed distally by two slightly smaller teeth and a series of 4
denticles before the apical tooth. Basal lamella of mandible concealed by clypeus and not visible. Anterior
clypeal margin broadly and shallowly concave between the broadly rounded anterolateral angles, the
margin equipped with 6 spatulate to spoon-shaped broad hairs which are strongly curved towards the
midline and arranged in three pairs; the innermost pair so strongly curved together that the apices are
almost touching, the outermost pair intermediate in size to the very large spoon-shaped hair at the
anterolateral clypeal corner. Lateral clypeal margins shallowly convex and very feebly convergent
anteriorly, equipped with a fringe of anteriorly curved large spatulate to spoon-shaped hairs. Clypeal and
cephalic dorsa with a ground-pilosity of small spoon-shaped curved hairs. Sides of occipital lobes in
full-face view with freely projecting long feebly clavate hairs. With head in profile the dorsum from the
highest point of the vertex back to the occipital margin with a number of long erect to suberect anteriorly
curved hairs which are feebly clavate to weakly remiform. Preocular laminae in full-face view weakly
divergent anteriorly but reaching a maximum width before meeting the clypeus, and from there to the
clypeus slightly convergent. Antennal scapes narrow basally, bent at about the basal third and broadest at
about this point, the leading edge subangulate at the point of maximum width and with a series of freely
projecting spatulate hairs. Flagellate hairs absent. Clypeus, area between frontal lobes and a short median
strip behind that smooth and unsculptured. Remainder of head densely punctate to reticulate-punctate,
the punctures and spaces between them glossy. Head in profile with vertex moderately high and narrowly
rounded, the ventral surface behind the level of the eye broadly and evenly convex. Maximum diameter of
eye distinctly less than maximum width of scape. Pronotum with lateral margins bluntly narrowly rounded
but not marginate, the dorsum without a median longitudinal ridge or carina and the humeri lacking
flagellate hairs. Anterior portion of mesonotum raised up to level of pronotal dorsum , the remainder of the
mesonotum and the propodeal dorsum markedly depressed. Highest point of raised anterior portion of
mesonotum with a pair of long erect stout hairs which constitute the only standing pilosity on the dorsal
alitrunk. Metanotal groove absent, the mesonotal and propodeal dorsa forming a single surface. Propodeal
THE AFROTROPICAL DACETINE ANTS 293
teeth long and narrow, slightly upcurved, the infradental lamellae narrow. Alitrunk dorsum with scattered
minute appressed hairs and the long mesonotal pair mentioned above . Petiole with one pair of long straight
simple hairs, postpetiole with 3-4 pairs. Gastral tergites with numerous straight hairs which are simple to
feebly clavate, erect to suberect. Dorsal alitrunk glassy smooth, highly polished with widely separated
small punctures. Petiole and the voluminous postpetiole glassy smooth dorsally, the first gastral tergite
unsculptured except for the basigastral costulae. Pleurae of alitrunk smooth except for a patch of evenly
spaced punctures occupying the central third or slightly more of the mesopleuron. Extreme base of first
gastral sternite with sparse but sharply incised punctures. Spongiform appendages of pedicel segments
strongly developed in profile. In dorsal view the posterior margin of the petiole node with a transverse strip
which is broad and spongiform postrolaterally but which is very narrow and lamellar centrally. The concave
anterior margin of the postpetiole with a narrow translucent lamellar strip, the posterior margin with a
transverse strip which is broad laterally but contracted down to a narrow isthmus medially. Transverse
basal strip of first gastral tergite lamellar rather than spongiform and traversed by the raised basigastral
costulae. Colour light glossy brown.
PARATYPE WORKERS. TL 2-2-2-3, HL 0-58-0-60, HW 0-41-0-42, CI 70-71, ML 0-06, MI 10, SL 0-26-0-28, SI
63-67, PW 0-26-0-28, AL 0-60-0-62 (5 measured). As holotype.
Holotype worker, Rwanda: Rangiro, 10.vii.1973, 1800m (P. Werner) (MHN).
Paratypes. 4 workers with same data as holotype; 1 worker with same data but 6.viii.l973 (MHN;
BMNH; MCZ).
A very distinctive species of the emarginata-gioup , gatuda is characterized by its distribution of
simple pilosity, reduced alitrunk sculpture and glassy smooth body.
Smithistruma hensekta sp. n.
(Fig. 12)
HOLOTYPE WORKER. TL 2-1 , HL 0-60, HW 0-41 , CI 68, ML 0-10, MI 17, SL 0-24, SI 59, PW 0-25, AL 0-58.
Mandibles with 5 relatively large teeth following the basal lamella (concealed by the clypeus), distal to
which are 2 slightly smaller teeth followed by 4 minute denticles and a small apical tooth. Anterior clypeal
margin transverse, equipped with 3 pairs of medially curved flattened hairs of which the outermost pair is
the largest, forming a transition to the long anteriorly curved spatulate hairs which form a continuous fringe
along the lateral clypeal margins. Sides of clypeus feebly convergent anteriorly. With the head in full-face
view the sides with numerous straight to slightly curved freely projecting stout hairs which are feebly
clavate. Dorsum of clypeus and cephalic dorsum from posterior clypeal margin to highest point of vertex
with numerous short scale-like hairs which are curved anteriorly. Scale-like hairs absent from highest point
of vertex back to the occipital margin, replaced by numerous distinctly longer erect to suberect hairs which
are simple to feebly clavate and mostly slightly curved anteriorly. This broad band of simple hairs occupies
the dorsum from the vertex to the occiput and the surfaces of the occipital lobes. Flagellate hairs absent.
Antennal scapes curved and broadened at about the basal third, the leading edge of the scape with a freely
projecting row of long stout hairs. Maximum diameter of eye 0-15xHW. Cephalic dorsum reticulate-
punctate everywhere, with a rough granular appearance. Clypeal dorsum less strongly sculptured than
remainder of head. Pronotum not sharply marginate laterally, lacking a median longitudinal ridge or carina
dorsally and without flagellate hairs at the humeri. Metanotal groove absent but posterior half of
mesonotum very shallowly concave. Propodeal teeth long and narrow, slightly upcurved along their
length. Infradental lamellae vestigial. Flagellate hairs absent but pronotal humeri each with a laterally
projecting stout hair which is feebly clavate. Dorsal alitrunk with 6 pairs of elongate stout erect hairs which
are simple to feebly clavate, and similar hairs are numerous on the petiole, postpetiole and first gastral
tergite but tending to be curved posteriorly on the pedicel segments; without other pilosity. Sides of
alitrunk densely punctate. Dorsal alitrunk densely punctate, the punctures slightly smaller and more
widely spaced on the pronotum than on the mesonotum or propodeum. Dorsum of petiole finely punctate;
disc of postpetiole unsculptured and glassy smooth. First gastral tergite unsculptured except for the sharply
defined row of basigastral costulae. Spongiform appendages of pedicel segments strongly developed in
profile. In dorsal view the petiole node with a spongiform strip along the posterior margin, the spongiform
material also extending down the posterolateral surfaces of the node. Anterior postpetiolar margin
transverse to exceptionally feebly concave, with a narrow spongiform strip; the posterior margin with a
much broader spongiform band which is broadest posterolaterally and narrowed medially. In dorsal view
the lateral spongiform tissue of the postpetiole can be seen projecting beyond the outline of the disc. Base
294
BARRY BOLTON
of first gastral tergite with a transverse spongiform band which is overlapped by that on the posterior
margin of the postpetiole. Colour uniform dull yellow.
PARATYPE WORKERS. TL 2-0-2-1, HL 0-58-0-60, HW 0-39-0-41, CI 67-68, ML 0-10-0-11, MI 17-18, SL
0-24-0-25, SI 59-63, PW 0-24-0-26, AL 0-58-0-60 (4 measured).
As holotype but in two workers the mandibles are open and the basal lamella is visible as a broad-based
high triangle which tapers strongly to a narrow truncated apex; there is no diastema between the basal
lamella and the basalmost tooth of the principal row.
Holotype worker, Ghana: Mampong, 9.ii.l970 (P. Room) (BMNH).
Paratypes. 4 workers with same data as holotype (BMNH; MHN; MCZ).
Non-paratypic material examined. Ivory Coast: Divo (L. Brader). Ghana: Tafo (D. Leston); Mampong
(P. Room). Cameroon : nr Yaounde (G. Tenon). Gabon: Plateau d'Ipassa (/. A. Barrd). Angola: Salazar
(P. Hammond).
The dimensions of the seven specimens constituting the non-paratypic material are HL
0-56-0-60, HW 0-37-0-40, CI 66-69. MI 17-18, SL 0-24-0-25, SI 60-65. Apart from slight
variation in intensity of punctate sculpture on the alitrunk the main variation is only that 6-7
pairs of hairs may be present on the dorsal alitrunk and that the outermost pair of hairs on the
anterior clypeal margin (at the corners) may be relatively small, so that the anterior margin may
have 3-4 medially curved pairs of hairs.
Within the emarginata-group hensekta is quickly diagnosed by its transverse anterior clypeal
margin and characteristic pilosity as described above. It apppears closest related to truncatidens
but the two are separated as follows in the worker.
hensekta
HW 0-37-0-41, CI 66-69.
Anterior clypeal margin transverse.
Dorsum of head from highest point of
vertex to occiput with erect to
suberect simple to clavate hairs.
Pronotal humeri each with a laterally
projecting stout feebly clavate hair.
Dorsal alitrunk with 6-7 pairs of
stout erect hairs.
Base of first gastral tergite with a
broad spongiform strip, the basigastral
costulae not traversing it, not running
up to the basal margin.
Posterior spongiform appendage of
postpetiole not interrupted medially.
truncatidens
HW 0-46-0-54, CI 72-76.
Anterior clypeal margin concave.
Dorsum of head from highest point of
vertex to occiput with anteriorly
curved flattened hairs which are
suberect only close to occipital
margin.
Pronotal humeri without laterally
projecting hairs.
Dorsal alitrunk with 1-2 pairs of
stout erect hairs.
Base of first gastral tergite without
a broad spongiform strip, the
basigastral costulae running up to
the basal margin.
Posterior spongiform appendage of
postpetiole interrupted medially.
Smithistruma impidora sp. n.
(Fig. 6)
HOLOTYPE WORKER. TL 2- 1, HL 0-64, HW 0-40, CI 63, ML 0-06. MI 9, SL 0-25, SI 63, PW 0-25, AL 0-58.
Mandibles equipped with 5 relatively large teeth following the basal lamella (which is concealed by the
clypeus). Distal to these are two slightly smaller teeth followed by 4 minute denticles and a small apical
tooth. Anterior clypeal margin broadly and evenly concave, equipped with a series of 7 scale-like hairs
which project forward over the mandibles. Of these the three central hairs are the smallest and the
outermost pair, almost at the anterolateral angles, are by far the largest and form a transition to the fringe
of large spatulate to spoon-shaped projecting hairs which line the lateral clypeal margins. Dorsum of
clypeus and cephalic dorsum devoid of hairs of any description; upper scrobe margins and sides of head
posteriorly devoid of hairs of any description. The cephalic dorsum and clypeal dorsum with scattered
minute pubescence which is only visible at high magnification. Sides of clypeus shallowly convex and
convergent anteriorly in full-face view. Preocular laminae slightly divergent anteriorly in full-face view.
Antennal scapes broad and flattened, bent very close to the base and broadest at the level of the bend or
just distal to it. Leading edges of scapes evenly rounded at the bend and with a series of projecting large
THE AFROTROPICAL DACETINE ANTS 295
spatulate hairs. Cephalic dorsum finely punctulate-granular everywhere, the clypeus shagreened. Head in
profile very obviously dorsoventrally flattened, the ventrolateral margin of the head almost flat and the
mid-posteroventral convexity of the head vestigial. Eyes of moderate size, the maximum diameter about
0-13xHW, less than the maximum width of the scape. Pronotum not marginate laterally, without a median
longitudinal ridge or carina dorsally. With alitrunk in profile the mesonotum and propodeum forming a
single almost flat surface, without trace of metanotal groove. Propodeal teeth only slightly elevated from
the line of the mesonotal-propodeal dorsa, somewhat upcurved along their length. Infradental lamellae
vestigial, merely a minute crest between the propodeal teeth and the metapleural lobes. Dorsal alitrunk
and dorsal surfaces of petiole , postpetiole and first gastral tergite lacking standing hairs of any description .
The alitrunk and first gastral tergite only with minute appressed pubescence but the petiole and postpetiole
bordered posteriorly by a sparse row of indistinct appressed hairs which project backward over the
posterior spongiform appendages of the segments. Pronotal dorsum exceedingly feebly sculptured with
vestiges of low minute longitudinal rugulae which are almost completely effaced. Mesonotum and
propodeal dorsum smooth. Dorsum of petiole node shagreened, postpetiolar disc smooth. First gastral
tergite unsculptured except for the feeble and widely spaced basigastral costulae. Sides of alitrunk mostly
smooth but with punctures on the mesopleuron and sides of the propodeum. Spongiform appendages of
pedicel segments massively developed in profile. In dorsal view the posterior border of the petiole node
with a transverse spongiform strip which continues down the sides. Anterior margin of postpetiole
shallowly concave and with a transverse spongiform strip, the posterior margin convex at the sides but
flattened or slightly indented medially and with a transverse spongiform strip which is narrowed centrally.
Sides of postpetiole disc with the more ventrally situated spongiform material projecting beyond the
outline and visible in dorsal view; the spongiform tissue outline diverging from front to back. Base of first
gastral tergite with a transverse spongiform band which is overlapped by that on the postpetiole. Colour
uniform light brown.
PARATYPE WORKERS. TL 2- 1-2-2, HL 0-62-0-64, HW 0-39-0-40, CI 63, ML 0-06, MI 9-10, SL 0-25-0-26, SI
64-65, PW 0-24-0-25, AL 0-56-0-60 (2 measured).
As holotype but maximum diameter of eye 0- 1 1-0-13 x HW.
Holotype worker, Ivory Coast: Abidjan, Banco Nat. Park, primary forest, dead trunk, 3.iii.l977 (7.
Lobl) (MNH).
Paratypes. 2 workers with same data as holotype (BMNH; MCZ).
The flattened head and lack of specialized pilosity isolate impidora from the other members of
the emarginata-group. The closest related species is chyatha, but here the dorsum of the head
retains a transverse band of suborbicular hairs between the vertex and the occipital margin. In
behasyla, another close species, two such bands of hairs are present, one occipitally and one just
behind the frontal lobes, and the head is by no means as strongly dorsoventrally flattened.
Smithistruma sharrs sp. n.
(Fig. 3)
HOLOTYPE WORKER. TL 2-1, HL 0-62, HW 0-38, CI 61, ML 0-06, MI 10, SL 0-28, SI 74, PW 0-24, AL 0-60.
Mandibles with 5 relatively strong teeth following the basal lamella without a diastema. Distal to these
main teeth are two slightly smaller teeth, followed by 4 minute denticles and an apical small tooth. Anterior
clypeal margin strongly and evenly concave, the concavity involving the entire border between the
anterolateral angles. Lateral margins of clypeus feebly convex and convergent anteriorly, fringed with a
continuous series of large flattened spatulate to spoon-shaped hairs which project freely and are curved
anteriorly. Anterior clypeal margin with a row of 6 broadly scale-like to suborbicular hairs which project
out over the mandibles. Dorsum of clypeus and remainder of head densely covered with broadly scale-like
to suborbicular hairs which are densest on the clypeus; such hairs also fringe the lateral borders of the head
in full-face view. Flagellate hairs or other pilosity absent. Preocular laminae broad in full-face view and
somewhat divergent anteriorly. Antennal scapes narrow basally, bent at about the basal quarter and
suddenly broadened, broadest at about this level and the leading edge evenly rounded with a narrow
prominent lamina. Dorsal surface of scape with scale-like hairs but leading edge with a series of freely
projecting longer narrower hairs, the longest of which occurs at about the broadest part of the scape. Eyes
of moderate size, about 0-11 xHW, smaller than the maximum width of the scape. Dorsum of clypeus and
area immediately posterior to it very finely reticulate-punctate, with a granular appearance. Remainder of
dorsum similarly sculptured but also with scattered very short low rugulae. Dorsal alitrunk with scattered
but conspicuous scale-like hairs, which are also conspicuous on the petiole and postpetiole, though
296 BARRY BOLTON
averaging smaller in size. In dorsal view the posterior margin of the postpetiolar disc with a row of 5-6
scale-like to spatulate hairs on each side of the midline which project out over the spongiform tissue; the
posterior petiolar margin with a similar row of 4 scale-like hairs (2 on each side of the midline). Standing
simple hairs absent from alitrunk, petiole, postpetiole and first gastral tergite; flagellate hairs never
developed. Alitrunk not marginate laterally, the pronotum flattened and without a median longitudinal
ridge or carina dorsally. With the alitrunk in profile the mesonotum very slightly raised above the level of
the pronotum and propodeum. Metanotal groove not impressed but its site marked by the small step-down
from the mesonotal to the propodeal dorsum. Propodeal teeth strong, broad basally and slightly upcurved
along their length, the infradental lamella very narrow and its outline distinctly concave. Pleurae and sides
of propodeum densely punctulate. Pronotal dorsum exceedingly feebly rugulose, the rugulae tending to be
irregular but with an overall longitudinal trend. Spaces between the minute rugulae shagreened and dull.
Remainder of dorsal alitrunk and petiole dorsum finely punctulate. Disc of postpetiole smooth and shining
and first gastral tergite unsculptured except for the basigastral costulae which form an uninterrupted band
across the base of the sclerite. Spongiform appendages massively developed in profile. In dorsal view the
sides and posterior margin of the petiole node surrounded by continuous thick spongiform material. The
postpetiole with an anterior spongiform transverse strip and with the lateral spongiform material
projecting beyond the lateral outlines of the disc in dorsal view. Posterior margin of postpetiole disc with a
continuous broad spongiform strip which is narrower centrally than at the sides but not broken . Base of first
gastral tergite with a transverse spongiform band which is as broad as, and is overlapped by, that on the
posterior postpetiolar margin. Colour medium to dark brown.
PARATYPE WORKERS. TL 2-0-2-2, HL 0-58-0-64, HW 0-35-0-38, CI 56-63, ML 0-06-0-08, MI 9-12, SL
0-26-0-28, SI 73-78, PW 0-22-0-26, AL 0-54-0-62 (20 measured).
As holotype but with maximum diameter of eye 0-ll-0-15xHW. In some the longitudinal nature of the
minute pronotal rugulae is better shown than in others, which are less regular and like the holotype. The
maximum number of scale-like hairs fringing the posterior dorsal margin of the petiole node appears to be 6
(3 on each side of the midline) though the outermost on each side may actually arise on the lateral margin of
the node and project backward. The basal lamella of the mandible consists of a high rectangle with concave
sides, visible in one of the paratypes.
Holotype worker, Ivory Coast: Issoneu, 12.x. 1980 (V. Mahnert & J.-L. Ferret) (MHN).
Paratypes. Ivory Coast: 14 workers and 1 female with same data as holotype; 9 workers, Man, 7.x. 1980
(V. Mahnert & J.-L. Ferret); 1 workers and 1 female, Dropleu, 10.x. 1980 (V. Mahnert & J.-L. Ferret); 4
workers, Tai Forest, 17.x. 1980 (V. Mahnert & J.-L. Ferret) (MHN; BMNH; MCZ).
Non-paratypic material examined. Ghana: Tafo (D. Leston); Mampong (P. Room). Cameroun:
Nkoemvon (D. Jackson); nr Yaounde (G. Terron). Angola: Salazar (P. Hammond); nr Gubela (P.
Hammond).
Dimensions of the non-paratypic samples fall within the range given for the paratypes.
The closest relative of sham is cavinasis which shares the remarkable dense suborbicular
pilosity. Details for the separation of the two are given under the latter name.
Smithistrumu truncatidens Brown
(Figs 10, 13)
Smithistruma (Smithistruma) truncatidens Brown, 19506: 43, pi. 3, fig. 1. Holotype and paratype workers,
TANZANIA: Lupembe (K. Bock) (paratype in MCZ) [examined]. [See also Brown, 1953a: 127.]
WORKER. TL2-4-3-0, HLO-62-0-72, HWO-46-0-54, CI 72-76, MLO- 10-0- 12, MI 14-18, SLO-28-0-34, SI
58-65, PW 0-30-0-35, AL 0-64-0-78 (12 measured).
Basal lamella of mandible a high rectangle, truncated apically and with shallowly concave sides. The
lamella is followed without a diastema by 5 relatively large teeth, two slightly smaller teeth, a row of 4
minutes denticles and a small apical tooth. Anterior margin of clypeus concave and equipped with a series
of 8 scale-like hairs, arranged in 4 pairs, which are curved towards the midline and project out over the
mandibles. Lateral margins of clypeus with a continuous row of anteriorly curved spatulate to spoon-
shaped large hairs which form a fringe around the clypeus. Clypeal dorsum and cephalic dorsum with
numerous small flattened hairs which are curved anteriorly and appear scale-like in full-face view. The
posteriormost one or two rows of hairs, close to the occipital margin, are longer, narrower and more erect.
Sides of head with freely projecting elongate feebly clavate hairs which curve upwards and often weakly
forwards from their points of origin. Antennal scapes bent at the basal third, broadest at or just distal to the
THE AFROTROPICAL DACETINE ANTS 297
bend. Leading edge of scape quite evenly rounded at the bend and with a series of freely projecting long
stout hairs. Dorsum of head densely punctate and matt, usually with a coarsely granular appearance but
sometimes with the punctures more widely spaced. Clypeal dorsum much less strongly sculptured,
frequently shining. Maximum diameter of eye 0- 14-0-16xHW. Pronotum not sharply marginate laterally,
the dorsum lacking a median longitudinal ridge or carina. With the alitrunk in profile the anterior half of
the mesonotum elevated and on the same level as the pronotal dorsum. Posterior half of mesonotum and
the propodeum markedly depressed below this level. Metanotal groove absent, the posterior half of the
mesonotum and the propodeal dorsum forming a single uninterrupted surface. Propodeal teeth triangular
and acute, weakly elevated and with a narrow infradental lamella. Dorsal alitrunk with a number of
appressed short hairs which are most easily seen on the pronotum. Erect hairs on alitrunk restricted to a
single long stout pair which are weakly clavate apically, situated on the raised anterior portion of the
mesonotum. In some samples a shorter second pair of erect hairs is present further back on the
mesonotum. Flagellate hairs absent. Dorsal surfaces of petiole, postpetiole and first gastral tergite with
numerous suberect to erect stout hairs which are simple or feebly clavate. Sides of alitrunk punctate but the
upper and posterior portions of the mesopleuron, the metapleuron and the anteriormost part of the
propodeal side forming an extensive smooth area on which punctures are extremely sparse or absent.
Dorsum of pronotum punctate, usually densely so but may be more widely scattered in some. Frequently
the walls of the individual punctures align to form feeble rugulae and the surface appears granular.
Remainder of dorsal alitrunk more sharply reticulate-punctate, the punctures extending onto the pro-
podeal declivity between the teeth. Petiole node punctate dorsally. Disc of postpetiole commonly
unsculptured but in many showing faint longitudinal costulae towards the sides. In some, often larger
individuals, the costulae are more extensive and occur over most or all of the postpetiolar dorsum. First
gastral tergite unsculptured except for the conspicuous basal costulae. Spongiform appendages of pedicel
segments strongly developed in profile. In dorsal view the petiole node with a transverse spongiform strip
on the posterior margin and the postpetiole with a narrow strip which is broadest posterolaterally but which
becomes narrower medially and is vestigial or interrupted at the centre of the margin. At this point the
posterior face of the postpetiole disc is itself indented . Base of first gastral tergite with a transverse lamellar
strip which is not spongiform and which is traversed by the basigastral costulae, the latter arising at the
anterior margin of the sclerite. Colour dull yellow to light yellowish brown.
S. truncatidens is closely related to datissa, dendexa, gatuda and hensekta. Characters separating
tmncatidens from the last of these are tabulared under hensekta. S. datissa is a more darkly
coloured species which lacks the freely projecting hairs on the sides of the head which are seen in
truncatidens. It also lacks standing hairs on the dorsal alitrunk and has the postpetiolar disc
strongly costulate. In gatuda the alitrunk and petiole dorsum are glassy smooth with widely
scattered punctures and the sides of the alitrunk are unsculptured except for the central
mesopleuron. The eyes are distinctly smaller, measuring only 0-09 x HW in gatuda as opposed to
0- 14—0- 16x HW in truncatidens. Finally dendexa, a smaller species, is separated by its possession
of fine longitudinal rugulose sculpture on the pronotum, which is not present in truncatidens.
MATERIAL EXAMINED
Rwanda: Kayove (P. Werner). Burundi: Bujumbura (A. Dejean); Bugarama (A. Dejean); Imbo Plain
(A. Dejean). Kenya: Nairobi (V. Mahnert); nr Narok (V. Mahnert &J.-L. Ferret); Embu, Irangi Forest
Sta. (V. Mahnert & J. -L. Ferret). Tanzania: Lupembe (K. Bock).
The fransversa-group
With the characters of the emarginata-group but the basal lamella of the mandible is an evenly rounded
broad lobe , visible in full-face view even when the mandibles are closed. Infradental lamella on propodeum
broad and conspicuous.
Recorded only from South Africa, transversa is certainly a direct derivative of the emarginata-
group which has modified the shape of the mandibular basal lamella from the high triangular or
rectangular structure seen in that group to the low broad lobe which it possesses, without
altering the basic dental pattern of the parent stock.
Smithistruma transversa (Santschi)
Strumigenys transversa Santschi, 1913a: 258 (diagnosis in key). Holotype worker, SOUTH AFRICA: Natal
(not in NMB; presumed lost).
298 BARRY BOLTON
Smithistruma (Smithistruma) transversa (Santschi) Brown, 1948: 105; 1953a: 127.
WORKER. TL 2-2-2-4, HL 0-62-0-66, HW 0-42-0-46, CI 68-71, ML 0-12-0- 13, MI 19-20, SL 0-27-0-28, SI
60-64, PW 0-26-0-28, AL 0-58-0-62 (5 measured).
Basal lamella of mandible an elongate high broadly rounded lobe whose length along the base is
approximately the same as the length of the masticatory margin occupied by the principal row of 5 teeth,
and which is clearly visible even when the mandibles are closed. Height of the basal lamella equal to that of
the longest of the teeth. Principal row of 5 teeth followed distally by 2 slightly smaller teeth and 4 minute
denticles before the small apical tooth. Anterior clypeal margin transverse to very shallowly evenly convex,
equipped with 8 scale-like hairs which project forwards over the mandibles and which are usually slightly
curved medially. Lateral margins of clypeus very slightly convergent anteriorly and with an unbroken
series of long broad spatulate to spoon-shaped hairs which project freely and are curved anteriorly.
Dorsum of clypeus and cephalic dorsum with numerous scale-like to spoon-shaped anteriorly curved short
hairs; those on the clypeus smaller than those on the cephalic dorsum. Upper scrobe margins and sides of
the occipital lobes with an unbroken sequence of sharply anteriorly curved spoon-shaped hairs which are
closely applied to the surface. Flagellate hairs absent. Antennal scape narrow basally, bent at about the
basal third and broadest just beyond the bend. Leading edge of scape with a projecting row of strong
spatulate to spoon-shaped hairs, the dorsum of the scape also with spatulate hairs present. Eyes relatively
large, maximum diameter 0-18xHW, greater than the maximum width of the scape. Entire dorsum of head
finely and densely reticulate-punctate. Pronotum not marginate laterally and without a median longitudin-
al ridge or carina dorsally. Flagellate hairs absent. Metanotal groove not impressed and propodeal teeth
subtended by a broad and conspicuous infradental lamella. Dorsal alitrunk with narrowly spatulate short
hairs, most of which are reclinate but a few of which may be subdecumbent. Petiole and postpetiole with
similar pilosity and also with elongate quite stout simple hairs which are directed posteriorly. First gastral
tergite with a transverse row of 4 erect hairs basally, grouped in pairs on each side of a central broad gap.
Remainder of first tergite hairless except for a widely separated pair close to the apical margin. Pleurae of
alitrunk mostly smooth, usually with some punctures basally and on upper anterior portion of the
mesopleuron. Pronotal and mesonotal dorsa finely and densely reticulate-punctate but dorsum of
propodeum mostly or entirely smooth; usually with some laterally situated punctures and with punctures
on the declivity between the propodeal teeth. Dorsum of petiole node distinctly broader than long,
sometimes indented medially; unsculptured or at most with superficial vestiges of punctate sculpture.
Postpetiole smooth and shining. First gastral tergite unsculptured except for the basigastral costulae, which
radiate from each side of a smooth median area. Spongiform appendages of pedicel segments strongly
developed in profile. In dorsal view the petiole node with a broad posterior strip; the anterior margin of the
postpetiole with a narrow strip, the posterior margin bordered by a spongiform strip which is broad
posterolaterally but rapidly narrowing medially, very narrow or interrupted centrally where the post-
petiole itself is indented. Base of first gastral tergite with a transverse strip which has its anterior free
margin concave medially and convex at the sides where the costulae arise. Colour dark brown to blackish
brown.
As the holotype of this species has not been found it is necessary to rely on the inadequate
descriptions presented by Santschi (19130; 1914c). Accordingly I attach the name transversa to
four short series from Natal and Cape Province in South Africa which match the available
descriptions tolerably well.
This species is peripheral to the emarginata-group but is isolated by its uniquely shaped
mandibular basal lamella and broad infradental lamellae on the propodeum.
MATERIAL EXAMINED
South Africa: Natal, Zululand, Eshowe (R. E. Turner); Dukuduku Forest Res. (W. L. & D. E. Brown);
Cape Prov., Pondoland, Port St Johns (R. E. Turner); Alexandria Forest Res. (L. Weatherill & W. L.
Brown).
The terroni-group
(Fig. 7)
Sharing the characters noted in the emarginata-group diagnosis (and particularly resembling the chyatha-
complex) but with a very distinctly modified mandible form. Blade of mandible narrow and somewhat
elongated (MI, however, within range of emarginata-group members), with a long diastema between the
basal lamella and the basalmost tooth, the diastema much longer than the height of the basalmost tooth.
Basal dental series usually of 6 but rarely of 7 small teeth (the number may vary on opposite mandibles in
THE AFROTROPICAL DACETINE ANTS 299
the same specimen), the next tooth relatively large, by far the largest tooth on the masticatory margin.
Distal to this large tooth are 3 small teeth, a single slightly larger tooth, 4 minute denticles and an apical
tooth, making a total of 16-17 teeth in all on the margin.
This group, represented by the single species terroni from Cameroun, appears to be derived
directly from the chyatha-complex of the emarginata-group by the modification of the mandible
outlined above. In the emarginata-group the basal lamella is a high truncated triangle or a
concave-sided high rectangle which is followed immediately by a row of 5 relatively large teeth,
without a diastema between the basal lamella and the basalmost tooth. The 5 teeth constituting
the principal row vary in size both between species and sometimes within species, but always one
of these five is the largest tooth on the margin. Distal to the row of 5 are 2 smaller teeth
(sometimes only fractionally smaller) which are themselves followed by 4 minute denticles and
an apical tooth, making a total of 12 teeth in all on the margin. Comparing the mandibles of the
two groups it appears that the basal row in terroni, which terminates in the relatively very large
tooth 7 (or rarely 8), is homologous to the basal row of 5 enlarged teeth in the emarginata-group
which have been spread out because of the elongation of the blade, and small secondary teeth
have developed in terroni to fill the gaps so formed. Teeth 6 and 7 of the emarginata-group are
represented in terroni by four teeth, the apicalmost of which is the same as tooth 7 in the
emarginata-group. Apically both have 4 denticles and a small apical tooth.
Smithistruma terroni sp. n.
(Fig- 7)
HOLOTYPE WORKER. TL 2-6, HL 0-74, HW 0-51, CI 69, ML 0-14, MI 19, SL 0-32, SI 63, PW 0-32, AL 0-76.
Mandibles narrowly triangular, with a distinct diastema separating the basal lamella (concealed by the
clypeus but visible in anterior view) from the basalmost tooth, the length of the diastema conspicuously
much greater than the height of the basalmost tooth. Counting from the base the mandibles with 6
relatively small teeth followed by a much larger seventh tooth, this tooth by far the largest on the
masticatory margin and more than twice larger than those proximal to it. Of the row of six teeth preceding
the enlarged seventh the first, third and fifth are larger than the second and fourth, the sixth is slightly
smaller than the fifth but slightly larger than the fourth. The large seventh tooth is followed distally by 3
small teeth, a larger tooth, 4 minute denticles and a small apical tooth, making a total of 16 teeth altogether.
Both mandibles similarly armed in the holotype but in one of the paratypes the left mandible has an
additional minute denticle between the third and fourth tooth from the base, and this mandible thus has a
total of 17 teeth. Anterior clypeal margin shallowly concave, with a series of 8 short scale-like hairs which
are truncated apically; the outermost pair the largest. Anterolateral angles of clypeus rounded, with
medially curved spoon-shaped hairs, the sides of the clypeus divergent posteriorly and with larger
anteriorly curved spoon-shaped hairs. In full-face view the preocular laminae divergent anteriorly, the
upper scrobe margins divergent posteriorly, the lateral margins of the occipital lobes evenly convex and the
occipital margin deeply but evenly concave. Upper scrobe margins just behind the frontal lobes shallowly
depressed on each side of a central higher area. Antennal scapes bent near base, flattened and broadest just
distal to the bend, the leading edges with a row of freely projecting long spatulate to spoon-shaped hairs.
Entire dorsum of head densely punctate. Ground-pilosity everywhere of minute decumbent to appressed
stubble-like hairs, without standing pilosity of any description on the cephalic dorsum and without
flagellate hairs. Eye moderate in size, with more than 15 ommatidia. Pronotum marginate anteriorly, the
alitrunk without lateral margination but the propodeal dorsum separated from the sides by bluntly rounded
angles. In dorsal view the pronotum without a median longitudinal ridge or carina and the metanotal
groove absent. In profile the mesonotum shallowly convex, confluent with the shallowly sloping surface of
the pronotum anteriorly but sloping more steeply posteriorly. The posterior half of the mesonotum
forming a single surface with the propodeal dorsum. Propodeal teeth strong and stout, the infradental
lamella vestigial and represented only by a narrow rim down the concavity of the declivity below the
propodeal teeth. Sides of alitrunk mostly with scattered quite sharply defined relatively large punctures,
the spaces between which are smooth, but the anterior portion of the mesopleuron finely reticulate-
punctate, the punctures much smaller than elsewhere. Dorsal alitrunk with scattered punctures, the
pronotum also with vestiges of exceptionally fine rugulae. Dorsal alitrunk only with scattered minute
appressed hairs, without standing pilosity of any description and lacking flagellate hairs. Pedicel segments
in profile with spongiform appendages massively developed. Dorsum of petiole node sculptured with
strong scattered punctures and bordered posteriorly by a continuous transverse spongiform strip which is
300 BARRY BOLTON
densest posterolaterally. Postpetiole in dorsal view with the disc smooth and unsculptured, completely
surrounded by dense spongiform material. Posterior transverse spongiform strip of postpetiole feebly
sinuate medially but not distinctly indented. Base of first gastral tergite with a narrow but dense transverse
spongiform strip. Basigastral costulae narrow and sharply defined, not traversing the basal spongiform
tissue. Pedicel segments and gaster without standing pilosity, with minute appressed ground-pilosity and
the posterior margins of the petiole and postpetiole with 1-2 pairs of larger appressed spatulate hairs which
project backwards over the spongiform strips. Colour black.
PARATYPE WORKERS. TL 2-5-2-6, HL 0-72-0-74, HW 0-49-0-51, CI 68-69, ML 0-13-0-14, MI 18-19, SL
0-31-0-32, SI 62-63, PW 0-32-0-33, AL 0-74-0-78 (3 measured).
As holotype but with 7-8 short hairs bordering the concave anterior clypeal margin and one paratype
with an extra mandibular tooth as discussed above.
Holotype worker, Cameroun: nr Yaounde, sample 1911 (G. Tenon) (ENS A).
Paratypes. 4 workers with same data as holotype (ENS A; BMNH).
The unique construction of the mandible immediately separates terroni from all its Afrotropical
congeners.
The wefteri-group
(Figs 14-16)
Antennae with six segments. Basal lamella of mandible a high triangle which is truncated apically or a high
rectangle with concave sides, never a low rounded lobe. Often a small diastema present between the basal
lamella and the basalmost tooth. Principal dental row of mandible with 5 teeth but the next two teeth
distally may sometimes also be enlarged. Sculpture very coarse on body and usually also coarse on head,
very characteristic, the pronotum with strong rugae or sulci which are usually longitudinal. Anterior
clypeal margin in full-face view most often approximately transverse but sometimes extremely shallowly
concave or convex. Lateral and anterior margins of clypeus with a series of irregular projecting simple hairs
which may be acute, truncated or even feebly clavate apically, but which are never spatulate nor
spoon-shaped and which do not form an orderly fringing row such as is characteristic of the emarginata- and
related groups. Body pilosity consisting of an array of fine simple hairs which are usually dense and are
generally wavy, twisted, bent or otherwise deformed, but without bizarre hairs. Flagellate hairs absent
from pronotal humeri or at least indistinguishable from the other pilosity. Leading edges of antennal scapes
with freely projecting simple hairs. Pronotum not marginate laterally and lacking a median dorsal ridge or
carina. Propodeum without or at most with a vestigial infradental lamella.
The obvious outstanding character of this peculiarly African group of species is the heavy coarse
sculpture. No other species or species-group known in the world approaches the members of the
weberi-group in this aspect and this character alone will serve to separate the twelve members of
the group from their Afrotropical congeners.
The group falls into two informal complexes depending on whether the postpetiolar disc is
sculptured or smooth. In the minkara-complex (enkara, mirtkara, nykard) the disc of the
postpetiole is strongly and clearly longitudinally costulate. Of the three species in this complex
two are West African, with niinkara known only from Ivory Coast and enkara from Ivory Coast
and Ghana; the third species, nykara, has only been found in Zimbabwe to the present. In the
second, weberi-complex (arahana, f enkara, kerasma, malaplax, mekaha, placora, synkara,
tolomyla, weberi), the postpetiolar disc is smooth and shining. All the species of this complex are
from West or central Africa and the species range from Nigeria to Angola.
Smithistruma arahana sp. n.
HOLOTYPE WORKER. TL 2-5, HL 0-65, HW 0-40, CI 61, ML 0-05, MI 8, SL 0-28, SI 70, PW 0-30, AL 0-64.
Anterior clypeal margin transverse, the anterolateral angles rounded and the lateral margins feebly
divergent posteriorly. Outline of preocular laminae shallowly convex in full-face view, broadest at about
their midlength and slightly convergent both anteriorly and posteriorly. Lateral margins of clypeus with
simple projecting hairs, the shorter of which are curved anteriorly and the longer of which curve upwards or
forwards and upwards. Clypeal and cephalic dorsa equipped with simple fine ground-pilosity, the hairs of
the ground-pilosity short and arched forward so that their apices are in contact or nearly in contact with the
surface. Projecting above the ground-pilosity are longer stouter simple hairs which are erect or nearly so.
THE AFROTROPICAL DACETINE ANTS 301
On the clypeal dorsum most of these hairs curve forward then upward but the posteriormost clypeal row
are shallowly sinuate and are also the longest. Behind the clypeus similar erect curved to sinuate hairs are
present, none of which are longer than the posterior clypeal row. Close to the occipital margin are a few
hairs which are angled and have their apical portions narrowly flagellate. In full-face view the sides of the
head with numerous projecting simple hairs. Antennal scapes slightly bent in the basal third, broadest at
about the midlength , the leading edge with anteriorly projecting simple hairs most of which are upcurved in
their distal halves. Dorsum of head coarsely and very densely reticulate-rugulose everywhere, the clypeus
less strongly sculptured. Promesonotum not marginate, the pronotal dorsum without a median longitudin-
al carina. Metanotal groove not impressed. Propodeal teeth broad and triangular, short, the infradental
lamellae vestigial and represented only by a narrow rim; outline of the propodeal declivity in profile
distinctly concave. Sides of pronotum and propodeum coarsely rugose, the pleurae smooth. Mesopleuron
with a dense vertical band of fine punctulae close to its junction with the metapleuron but otherwise the
pleurae only with very widely scattered fine punctulae. Promesonotal dorsum coarsely and densely rugose,
the propodeal dorsum smooth except for a few feeble anteriorly situated punctures. Dorsal alitrunk with
numerous fine simple hairs. Spongiform appendages of pedicel segments massively developed in profile.
Outline of petiolar ventral process concave at about its midlength, the ventral postpetiolar lobe very large
indeed. In dorsal view the petiole node rugose and distinctly broader than long, with a very thick posterior
ruff of dense spongiform material, the thickness of which is greater than the length of the exposed dorsum
of the node. Laterally the spongiform tissue laps around the sides of the node almost to the anterolateral
angles. Disc of postpetiole in dorsal view completely surrounded by dense thick spongiform material, the
disc uneven and with scattered punctures, not glassy smooth as is usual in the weberi-complex but lacking
the strong costulae characteristic of the minkara-comp\ex. Anterior margin of postpetiolar disc bordered
by a dense spongiform strip, the sides with dense spongiform tissue which is narrowest anteriorly and
extremely broad posterolaterally. Posterior spongiform strip broad and with a narrow median cleft. Base
of first gastral tergite with a broad very finely and densely spongiform transverse strip, the tergite posterior
to this with short basigastral costulae. Dorsal surfaces of petiole, postpetiole and gaster with numerous fine
simple hairs. Colour dull brownish yellow to light brown.
PARATYPE WORKER. TL 2-6, HL 0-70, HW 0-43, CI 61 , ML 0-06, MI 9, SL 0-30, SI 70, PW 0-32, AL 0-70.
As holotype but disc of postpetiole less noticeably punctate.
Holotype worker, Cameroun: nr Yaounde, sample D2 (G. Tenon) (ENSA).
Paratype. 1 worker, Cameroun: nr Yaounde, sample 2419 (G. Terrori) (BMNH).
Among the members of the weberi-group six species combine the characters of having the
postpetiolar disc without dense costulate sculpture and having the metanotal groove unim-
pressed. Of these only arahana has the spongiform tissue behind the petiole node very densely
and massively developed. In dorsal view the spongiform material is thicker than the length of the
node in arahana, whereas in the five other species (fenkara, malaplax, placora, synkara,
tolomyla) it is decidedly narrower than the length of the node, in some being merely a lamella.
Smithistrumu enkara sp. n.
(Fig. 15)
HOLOTYPE WORKER. TL 2-4, HL 0-63, HW 0-41, CI 65, ML 0-08, MI 13, SL 0-28, SI 68, PW 0-28, AL 0-62.
Dentition as described for minkara but basal lamella of mandible (from non-paratypic material) a high
truncated rectangle with concave sides; a small diastema present between the basal lamella and the
basalmost tooth. Anterior clypeal margin transverse, the anterolateral angles of the clypeus rounded and
the sides slightly convergent anteriorly. Lateral margins of clypeus with numerous projecting curved to
flagellate simple fine hairs which are also present bordering the sides of the head. Dorsal surface of clypeus
and dorsal surface of head with abundant fine simple hairs which are irregular to flagellate and mostly
arched over so that the apices of most of them are directed back down towards the surface or are roughly
parallel with the surface. All the cephalic hairs are fine and simple, without erect to suberect longer stouter
straight hairs pointing up from the dorsum. Entire dorsum of head strongly reticulate-rugulose. Antennal
scapes only very feebly bent at about the basal third, broadest at about the midlength, the leading edges
arched convex and equipped with projecting simple hairs similar to those on the cephalic dorsum.
Maximum diameter of eye 0-12xHW. With the alitrunk in profile the mesonotum slightly elevated, the
metanotal groove not impressed. Sides of pronotum not sharply marginate and the dorsum lacking a
median longitudinal ridge or carina. Propodeal teeth narrow and acute, subtended by a vestigial
infradental lamella. Pronotum, mesonotum, petiole, postpetiole and gastral tergites with numerous fine
302 BARRY BOLTON
simple flagellate hairs, many of which are arched over towards the surface, as on the head. Sides of
pronotum rugose, pleurae and sides of propodeum punctate. Pronotal dorsum densely coarsely longitudi-
nally rugose, with a few cross-meshes; spaces between the rugae mostly narrow and smooth. Mesonotum
with irregular strong rugae the spaces between which are punctate. Propodeal dorsum punctate, the
declivity smooth. Petiole node irregularly but strongly rugose dorsally, the postpetiolar disc strongly
longitudinally costulate-rugose. Basigastral costulae dense and strongly developed, extending almost the
length of the sclerite centrally, less extensive on the sides. Spongiform appendages of pedicel segments
massively developed in profile. In dorsal view the posterior margin of the petiole node with a broad
spongiform strip whose posterior border is concave medially and which is broadest posterolaterally where
it forms a rounded lobe. Postpetiole in dorsal view with the disc completely surrounded by thick
spongiform material which is broadest posterolaterally and has the posterior strip indented medially. First
gastral tergite with a broad spongiform band basally which is overlapped by the posterior spongiform strip
of the postpetiole. Colour dark brown, the gaster blackish brown.
PARATYPE WORKER. TL 2-4, HL 0-63, HW 040, CI 63, ML 0-07, MI 11, SL 0-28, SI 70, PW 0-28, AL 0-62.
As holotype but maximum diameter of eye 0-13xHW.
Holotype worker, Ivory Coast: Abidjan, Banco Nat. Pk., primary forest, 3. Hi. 1977, in dead trunk (/.
Lobl) (MHN).
Paratype. 1 worker with same data as holotype (BMNH).
Non-paratypic material examined. Ghana: Tafo (D. Leston); Tafo (C. A. Collingwood). Ivory Coast:
Lamto (W. H. Gotwald). Cameroon: nr Yaounde (G. Tenon).
The five specimens constituting the non-paratypic material are very close to the holotype but
have some minor differences. Principal among these is a rugulose propodeal dorsum, not seen in
the type-series. With so little material available I cannot assess the significance of this and I am
not prepared to split them further at present.
Of the three species in this group which possess a sculptured postpetiolar disc, minkara is
easily differentiated by its very long narrow head, CI 54—58 as opposed to CI 63-68 in enkara and
nykara. These last two species are differentiated by the characters given in the key plus the fact
that nykara has long stout evenly curved clypeal hairs as well as the finer pilosity, such long hairs
being absent in enkara. With the pedicel segments in profile the lateral spongiform appendage of
the postpetiole touches or is confluent with the transverse strip bordering the anterior postpetio-
lar margin in enkara; in nykara there is a distinct gap between them.
Smithistruma f enkara sp. n.
HOLOTYPE WORKER. TL 2-4, HL 0-67, HW 0-43, CI 64, ML 0-07, MI 10, SL 0-31 , SI 72, PW 0-30, AL 0-66.
Dentition not clearly visible as mandibles closed but apparently like that described for malaplax.
Anterior clypeal margin transverse, the sides irregular, shallowly convex and weakly convergent anterior-
ly. With the head in full-face view the lateral clypeal margins with a few simple short anteriorly curved hairs
on the posterior half, but the pilosity dominated by the numerous stout hairs which project anterolaterally
are clavate apically and upcurved in their distal half to third. Sides of head with numerous similar
projecting clavate hairs which are curved forwards or upwards, the posterior curve of the occipital lobes
with weakly flagellate hairs replacing the clavate pilosity. In profile the clypeal dorsum with clavate hairs
anteriorly which curve upwards. The surface of the clypeus behind these hairs is shallowly concave and
hairless. Posteriorly the clypeal dorsum with a transverse row of sinuate clavate erect hairs which are
slightly longer than those situated anteriorly. Dorsum of head from posterior margin of clypeus to vertex
with simple short ground-pilosity which is curved anteriorly and closely applied to the surface, and with
longer stout clavate hairs which are erect to suberect, feebly inclined or curved anteriorly, all of about the
same length and stature and about equal in length to the posterior clavate clypeal row. Sloping portion of
head behind the vertex and in front of the occipital margin with weakly flagellate hairs replacing the clavate
pilosity. Antennal scapes feebly bent at about the basal third , the leading edge with a projecting row of long
curved hairs which are weakly clavate apically. Maximum diameter of eye 0-16xHW. Entire dorsum of
head densely reticulate-rugulose. Pronotum not marginate laterally, without a median longitudinal ridge
or carina dorsally. With the alitrunk in profile the metanotal groove absent, the propodeal teeth narrowly
triangular and subtended by a narrow infradental lamella whose free margin is evenly concave. Sides of
pronotum and propodeum irregularly rugulose, the pleurae punctate but the metapleuron mostly smooth
centrally. Pronotal dorsum strongly longitudinally rugose, with a few cross-meshes and with the inter-
spaces weakly punctate to granular. Mesonotum more strongly reticulate-rugose than pronotum, especial-
THE AFROTROPICAL DACETINE ANTS 303
ly posteriorly. Propodeal dorsum punctate, with rugulae at the sides and one or two weak transverse
rugulae close to the declivity, the latter smooth. Petiole node irregularly rugose dorsally, the postpetiole
smooth and shining. First gastral tergite unsculptured except for the strong basigastral costulae. Dorsal
surfaces of pronotum, 'mesonotum, petiole, postpetiole and gaster with numerous fine weakly flagellate
hairs. Spongiform appendages of pedicel segments massively developed in profile. In dorsal view the
petiole node with a broad posterior strip which is narrowed posteromedially. Postpetiole completely
surrounded by thick spongiform material in dorsal view, the posterior strip deeply indented medially. First
gastral tergite with a thick basal spongiform ruff. Colour medium brown, the gaster blackish brown.
PARATYPE WORKER. TL 2-3, HL 0-68, HW 0-43, CI 63, ML 0-07, MI 10, SL 0-30, SI 70, PW 0-30, AL 0-63.
As holotype.
Holotype worker, Angola: Dundo, Carisso Park, gallery forest, R. Luachimo, 7°22'S, 20°50'E,
26. iv. 1963, 'berlesate by native collector' (MCZ).
Paratype. 1 worker with same data as holotype (BMNH).
5". fenkara is closest related to placora, tolomyla and synkara. The characters linking them and
those which separate them are noted under synkara.
S. fenkara is separated from arahana by its massive development of the posterior petiolar
spongiform appendage, as discussed under the latter name; fenkara is differentiated from
malaplax by the lack of specialized hairs on the head behind the clypeus in the latter.
Smithistruma kerasma sp. n.
(Fig. 16)
HOLOTYPE WORKER. TL 2-5, HL 0-68, HW 0-44, CI 65, ML 0-06, MI 9, SL 0-30, SI 68, PW 0-32, AL 0-69.
Mandibular dentition (from a paratype) consisting of a high truncated rectangular basal lamella with
concave sides, followed by a small diastema and a principal row of 5 relatively large teeth. Distal to this with
2 slightly smaller teeth, 4 minute denticles and a small apical tooth. Anterior clypeal margin broadly
shallowly convex, sides of the clypeus irregular and only very weakly convergent anteriorly to the rounded
anterolateral angles. Preocular laminae weakly convex in full-face view, the lateral margins of the head
rugular and uneven. Lateral margins of clypeus in full-face view with a few simple anteriorly curved short
hairs and with longer stouter simple hairs which project anterolaterally from the margin and are curved
upwards. Sides of head with abundant fine simple projecting hairs which are curved anteriorly in their
apical halves. Hairs on clypeal dorsum fine, more or less vertical and curved towards the midline. Dorsum
of head with abundant fine simple hairs which are erect or suberect basally but which are angled anteriorly
in their apical halves, those situated more posteriorly on the dorsum being in general more strongly bent
forward than those situated more anteriorly. The most strongly bent hairs are inverted L-shaped. All hairs
on dorsal head approximately the same size and stature, without hairs which are obviously longer and
stouter than others. Dorsum of head coarsely irregularly reticulate-rugose, the clypeus similarly but less
intensely sculptured. Antennal scapes scarcely bent basally, broadest at about the midlength and the
leading edge with projecting curved simple hairs which also occur on the dorsum of the scape. Maximum
diameter of eye 0-16xHW. Pronotum not marginate laterally, without a median longitudinal ridge or
carina dorsally. In profile the alitrunk with the mesonotum strongly convex, sloping down posteriorly to a
broad, shallow but distinctly impressed metanotal groove. Dorsal outline of propodeum raised behind the
metanotal groove, then sloping downwards to the triangular propodeal teeth. Infradental lamellae of
propodeum vestigial, their free margins strongly concave. Pronotal and mesonotal dorsa with numerous
erect to suberect long fine simple hairs which are bent in their apical halves and often directed anteriorly.
Dorsal surfaces of petiole, postpetiole and gaster with elongate simple hairs which are subflagellate to
flagellate or sometimes arched over. Dorsal (outer) surfaces of middle and hind tibiae with projecting
simple subflagellate hairs. Sides of pronotum and propodeum coarsely rugose, the pleurae punctate; the
punctures of the mesopleuron smaller denser and more sharply defined than those on the metapleuron.
Pronotal dorsum coarsely longitudinally rugose, the rugae broad and high and the spaces between them
smooth. Mesonotum, metanotal groove and base of propodeal dorsum strongly rugose but the rugae less
massive and less regular than on the pronotum. Central area of propodeal dorsum with irregular punctures,
declivity smooth. Petiole dorsum coarsely rugose, postpetiole dorsum smooth and shiny. First gastral
tergite unsculptured except for the regular strong short basal costulae. With pedicel segments in profile the
spongiform appendages massively developed. In dorsal view the posterior margin of the petiole with a very
broad spongiform strip which has its free posterior margin shallowly concave medially and which is
broadest posterolaterally where its length is equal to that of the free side of the node in front of it. Disc of
304
BARRY BOLTON
postpetiole thickly surrounded by spongiform material on all sides in dorsal view. The broadly and
shallowly concave anterior margin of the postpetiole is equipped with a thick ruff-like transverse
spongiform band which is contiguous with the lateral spongiform material on each side. Convex posterior
margin of postpetiolar disc indented medially and bearing an extremely broad spongiform band whose
posterior margin is also indented medially. The spongiform material on each side of the median
indentation is as broad as the disc is long. Base of first gastral tergite with a thick spongiform ruff from
which the basigastral costulae emerge. Colour dark brown.
PARATYPE WORKERS. TL 2-5-2-6, HL 0-68-0-70, HW 0-44-0-45, CI 64-66, ML 0-06-0-07, MI 9-10, SL
0-30-0-31, SI 68-70, PW 0-32-0-33, AL 0-68-0-72 (9 measured). As holotype.
Holotype worker, Cameroun: Nkoemvon, 16.iii.1980 (D. Jackson) (BMNH).
Paratypes. 9 workers with same data as holotype (BMNH; MHN; MCZ; ENSA).
Among the nine species of this group which have the postpetiole unsculptured only three
(kerasma, mekaha, weberi) have the metanotal groove impressed. S. kerasma and mekaha differ
together from weberi as follows.
kerasma and mekaha
Median indentation of posterior
spongiform appendage of postpetiole
shallow, not approaching the margin
of the disc.
Spongiform material bordering margin
of postpetiole posteriorly as wide
from front to back as the disc of
the postpetiole is long.
Propodeal teeth long, the infradental
lamella vestigial and its free margin
evenly concave.
All hairs on dorsum of head of same
construction and approximate size,
not divided into appressed small
ground-pilosity and much larger
erect subclavate hairs.
Larger species, HL 0-68-0-70
HW 0-44-0-47.
Second tooth of principal mandibular
row the longest, the first
(basalmost) and third about equal
in length.
weberi
Median indentation of posterior
spongiform appendage of postpetiole
reaching the margin of the disc.
^pongiform material bordering margin
of postpetiole posteriorly distinctly
narrower from front to back than the
disc of the postpetiole is long.
Propodeal teeth short and broad, the
infradental lamella conspicuous and
its free margin straight or feebly
sinuate, not evenly concave.
Hairs on dorsum of head of two forms,
divided into small appressed
ground-pilosity and much larger
erect subclavate hairs.
Smaller species, HL 0-61, HW 0-39.
Second tooth of principal mandibular
row the longest but the first
(basalmost) very much smaller than
the third.
S. kerasma and mekaha are a very closely related pair but are quickly separated by the form of
the cephalic pilosity . In kerasma the principal cephalic hairs are erect or suberect basally but pass
through an obtuse angle so that their apical halves are directed anteriorly. In mekaha the
cephalic hairs lack this structure, instead being evenly arched forward from base to apex, their
apices generally in contact with the surface of the head some distance in front of their point of
origin.
Smithistruma malaplaxsp. n.
HOLOTYPE WORKER. TL2-1, HLO-64, HWO-40, CI63, ML 0-07, MI 11, SLO-28, SI 70, PWO-28, ALO-62.
Basal lamella of mandible a high truncated rectangle with concave sides, separated from the principal
tooth row by a small diastema. Of the 5 teeth following the diastema the first is the shortest and the second
is the longest. The principal row of 5 teeth is followed by two slightly smaller teeth , 4 minute denticles and a
small apical tooth. Anterior clypeal margin transverse, the sides feebly convergent. In full-face view the
lateral clypeal margins with a more ventrally situated series of projecting fine simple hairs which are curved
anteriorly, often sharply so. Situated above this row on the sides of the clypeus are numerous longer stouter
weakly clavate hairs which project laterally or anterolaterally and are upcurved or backcurved in the distal
third to half of their length. Clypeal dorsum with very sparse short anteriorly curved simple ground-pilosity
THE AFROTROPICAL DACETINE ANTS 305
and with numerous erect to suberect long stout weakly clavate hairs. Anteriorly on the clypeus the stout
hairs curve forward from their bases then upwards and usually slightly backwards. Posteriorly on the
clypeus is a single transverse row of stout clavate hairs which are much longer than those situated anteriorly
and which are vertical, weakly sinuate throughout their length and weakly directed anteriorly at their
apices (from the non-paratypic material as the posterior row of clavate hairs is crushed down in the
holotype). Dorsum of head behind clypeus only with simple fine pilosity, without the long weakly clavate
hairs which are so obvious on the clypeus; the fine hairs simply anteriorly curved and closely applied to the
surface in the area behind the clypeus but posterior to that, approaching the vertex and beyond, the hairs
are arched, looped or weakly flagellate. With the head in full-face view the sides with projecting simple
hairs similar to those on the dorsum, weakly flagellate, arched or looped. Scape feebly bent at its basal
third, broadest at about the midlength and the leading edge with a row of projecting simple curved hairs.
Maximum diameter of eye 0-16xHW. Pronotum not marginate laterally, without a median dorsal ridge or
carina. With alitrunk in profile the metanotal groove absent, not impressed. Propodeal teeth triangular and
acute, subtended by a narrow evenly concave infradental lamella. Dorsal surfaces of pronotum, mesono-
tum, petiole, postpetiole and gaster with numerous fine simple hairs which are arched, looped or weakly
flagellate. Sides of pronotum and propodeum reticulate-rugulose, the pleurae punctate. Promesonotal
dorsum densely and strongly rugulose. Propodeal dorsum rugulose and with vestigial punctures. Dorsum
of petiole node rugulose, the postpetiole smooth and shining. First gastral tergite unsculptured except for
the strong basigastral costulae. Spongiform appendages of pedicel segments strongly developed in profile.
In dorsal view the broad posterior strip of the petiole node concave medially. Postpetiole surrounded by
spongiform material in dorsal view, the broad posterior strip indented medially. Spongiform material at
base of first gastral tergite forming a narrow band which is mostly overlapped by the posterior postpetiolar
spongiform tissue, the area of the first tergite immediately behind the spongiform material lamellar and
traversed by the basigastral costulae. Colour orange-brown, the gaster blackish brown.
Holotype worker, Angola: nr Gubela, 17.iii.1972, forest litter (P. Hammond) (BMNH).
Non-paratypic material examined. Nigeria: Ibadan (B. R. Critchley). Zaire: Yangambi (M. Maldague).
Angola: R. Kahingo (Mwaoka).
The non-paratypic material consists of three specimens, one from each locality, which resemble
the holotype in all main characters but which show some sculptural variation. With so few
specimens available I cannot guess at the significance, or lack of significance, of this variation
and so leave all as a single species for the time being. The species is characterised and separated
from other members of the group by having a smooth postpetiolar disc, no metanotal groove , and
by having specialized long stout clypeal hairs which are absent from the dorsum of the head
behind the clypeus where only fine simple hairs are present. Other members of the group having
a smooth postpetiole and lacking a metanotal groove (arahana, fenkara, placora, synkara,
tolomyld) all have very obvious specialized hairs on the cephalic dorsum which are similar to or
even longer than those on the clypeus.
Smithistruma mekaha sp. n.
HOLOTYPE WORKER. TL 2-6, HL 0-70, HW 0-46, CI 66, ML 0-06, MI 9, SL 0-31, SI 65, PW 0-32, AL 0-71.
Principal dental row of 5 teeth, dentition as described for kerasma. Anterior clypeal margin extremely
shallowly convex, the anterolateral angles rounded. Lateral margins of clypeus very feebly divergent
posteriorly, the preocular lamellae continuing the lines of the clypeal margins in full-face view but slightly
convergent posteriorly. Lateral and anterior margins of clypeus with fine simple hairs which are directed
forward or forward and upward, the clypeal dorsum with some erect curved fine hairs. Behind the level of
the clypeus all hairs on the cephalic dorsum are fine, simple and strongly arched forward so that their apices
are in contact with the surface some distance in front of their bases. Lateral margins of head with some
freely projecting fine hairs and with curved hairs like those on the dorsum. Upper scrobe margins divergent
behind the frontal lobes, the sides of the head behind the level of the scrobes irregularly convex. Occipital
margin concave and with a narrow bordering rim or flange. Clypeus irregularly rugose, the sculpture much
weaker than on the cephalic dorsum. Dorsum of head coarsely irregularly rugose to coarsely punctate-
rugose, the rugae in places surrounding small foveolate punctures from which the hairs arise. Scapes
narrow at base, broadening to a maximum at about the midlength then narrowing again to the apex.
Leading edges of scapes with fine projecting simple hairs. Pronotum not marginate laterally, without a
median longitudinal ridge or carina. Metanotal groove shallowly but conspicuously impressed. In profile
the propodeal teeth short and stout, the infradental lamellae very narrow and with concave free margins.
306 BARRY BOLTON
Sides of pronotum, metapleuron and propodeum coarsely irregularly rugose, contrasting strongly with the
mesopleuron which is sculptured with fine sharply incised small separate punctures on a smooth surface.
Pronotal dorsum very coarsely irregularly longitudinally rugose, the rugae and the small spaces between
them smooth. Mesonotum similarly sculptured, propodeum rugose towards the sides but the centre of the
dorsum with a few deformed punctures. Dorsal alitrunk with numerous fine simple hairs. Spongiform
appendages of pedicel segments massively developed. In profile the ventral spongiform appendage of the
petiole forming a lobe anteriorly which is suddenly narrowed at about the level of the ascending face of the
node and then broadened again behind, as if a broadly triangular notch had been cut in the ventral margin
of the spongiform tissue. Ventral spongiform lobe of postpetiole very large. Petiole node in dorsal view
coarsely sculptured, with a thick posterior ruff of spongiform material which is almost as thick at its
midlength (its narrowest point) as the dorsum of the node is long, the spongiform material becoming even
thicker laterally. Postpetiolar disc unsculptured, smooth and shining, surrounded on all sides by dense
spongiform tissue. Anterior margin of postpetiole bounded by a transverse spongiform strip, the sides
bounded by projecting spongiform tissue which is narrowest anteriorly. Posterior spongiform strip of
postpetiole with its posterior margin very weakly indented medially, the indentation very shallow and not
approaching the margin of the disc; with a thick band of spongiform material separating the posteriormost
point of the disc from the base of the impression. Base of first gastral tergite with a broad dense spongiform
strip which is not traversed by the basigastral costulae; the latter short but strongly defined on the base of
the tergite proper. Pilosity of petiole, postpetiole and gaster entirely of fine simple hairs. Colour brown.
PARATYPE WORKER. TL 2-6. HL 0-70, HW 0-47, CI 67, ML 0-07, MI 10, SL 0-31, SI 66, PW 0-33, AL 0-73.
Asholotype.
Holotype worker, Cameroun: nr Yaounde, sample ABH (G. Terrori) (ENSA).
Paratype. 1 worker with same data as holotype (BMNH).
Among the known species of the weberi-complex only 3, kerasma, mekaha and weberi, have the
metanotal groove impressed. Of these weberi is recognised by the very strong impression in the
posterior margin of the spongiform strip bordering the postpetiole posteriorly. This impression
is so deep in weberi that it reaches to the margin of the postpetiolar disc, whereas in kerasma and
mekaha the impression is shallow and there is always a wide expanse of spongiform material
between the posterior margin of the postpetiolar disc and the deepest point of the impression.
Other differences from weberi are tabulated under kerasma. S. kerasma and mekaha are
separated by the form of the cephalic pilosity , which in the former consists of numerous standing
hairs which are erect basally but pass through an obtuse angle near their midlengths so that their
upper portions are directed forwards. In mekaha, on the other hand, all the cephalic hairs are
strongly arched forwards from base to apex so that their apices are in contact with the surface
some distance in front of their bases.
Smithistruma minkara sp. n.
(Fig. 14)
HOLOTYPE WORKER. TL 2-5, HL 0-73, HW 0-40, CI 55, ML 0-06, MI 8, SL 0-31, SI 78, PW 0-28, AL 0-68.
Head very long and narrow, CI range of entire type-series 54-58; CI range for all other known species of
the weberi-group is 61-68. Mandibles (from a paratype) armed with a high truncated basal lobe which is
slightly longer than any of the teeth in the principal row. Distal to the basal lamella is the principal row of 5
relatively large teeth, separated from the lamella by a small diastema. Following these are two slightly
smaller teeth, 4 minute denticles and a small apical tooth. Anterior clypeal margin transverse to
exceedingly shallowly convex, the lateral clypeal margins very slightly converent anteriorly and with
broadly rounded anterolateral angles. Sides and dorsum of clypeus with short curved ground-pilosity and
also with numerous much longer stouter curved simple hairs. The long stout simple hairs arising from the
lateral clypeal margins are directed outwards from the margin but then curve upwards or forward and
upwards. On the dorsum of the clypeus the hairs are shorter centrally than at the sides, directed vertically
or slightly curved. In profile the dorsum of the head behind the clypeus with short fine anteriorly curved
ground-pilosity which is decumbent, and with stouter longer straighter hairs which are vertical or nearly so,
these hairs shorter anteriorly than posteriorly on the head. In full-face view the sides of the head with
abundant long simple projecting hairs, most of which are curved or sinuate. Median portion of clypeus
from anterior tumulus to frontal lobes smooth or nearly so, the rest of the clypeus irregularly punctate.
Dorsum of head coarsely reticulate-punctate, with well developed rugulae between the punctures on the
THE AFROTROPICAL DACETINE ANTS 307
vertex. Occipital concavity bounded on each side by a small flange or tooth in full-face view. Antennal
scapes relatively long, narrowest at base but gradually increasing in width through the basal third, then
slightly bent and broadened, the evenly curved leading edge with a series of freely projecting curved long
simple hairs. Eyes of moderate size, maximum diameter 0-15xHW. Head flattened in profile, the dorsum
depressed and shallowly concave between clypeus and vertex, the eye bulging slightly beyond the ventral
margin of the scrobe. Dorsal surfaces of alitrunk (except propodeum), petiole, postpetiole and first gastral
tergite with numerous erect irregular to flagellate fine simple hairs, shorter more reclinate forms of which
also project from the dorsal (outer) surfaces of the middle and hind tibiae. Dorsum of promesonotum and
sides of pronotum strongly longitudinally rugose, the rugae smooth and rounded dorsally but the spaces
between them punctate to shagreened and dull. Propodeal dorsum unsculptured except for a few small
punctures, the declivity smooth. Pleurae mostly smooth, with a sparse median punctulate patch; the sides
of the propodeum irregularly strongly rugose. With the alitrunk in profile the metanotal groove very feebly
indicated, the propodeal teeth strong and broadly triangular, without infradental lamellae. Pronotum not
sharply marginate laterally and lacking a median dorsal longitudinal carina. Dorsum of petiole node
strongly irregularly rugose. Dorsum of postpetiole everywhere very strongly longitudinally costate to
rugose, the sculpture very regular and almost sulcate. Basigastral costulae fine and very numerous,
extending back almost to the apex of the segment in the centre of the sclerite, less extensive at the sides.
Spongiform appendages of pedicel segments massively developed in profile. In dorsal view the petiole
node surrounded posterolaterally and posteriorly by a thick spongiform strip. Disc of postpetiole in dorsal
view completely surrounded by thick spongiform tissue which is broadest posterolaterally. Base of first
gastral tergite with a thick spongiform transverse band which is overlapped by that on the posterior margin
of the postpetiole. Colour medium brown.
PARATYPE WORKERS. TL 2-4-2-6, HL 0-70-0-76, HW 0-40-0-44, CI 54-58, ML 0-06-0-08, MI 8-11, SL
0-30-0-33, SI 73-78, PW 0-28-0-29, AL 0-67-0-74 (14 measured).
As holotype but maximum diameter of eye 0-15-0- 18 xHW. In some the pleural punctate area is
somewhat more extensive than in others and frequently the mesonotum is rather more swollen in profile
than is the case in the holotype. One or two vestigial rugulae may be present on the propodeal dorsum,
especially towards the sides. The basigastral costulae may cover only about half of the first gastral tergite on
the centre of the sclerite.
Holotype worker, Ivory Coast: Monogaga, 24.x. 1980 (V. Mahnert & J.-L. Ferret) (MHN).
Paratypes. Ivory Coast: 11 workers with same data as holotype; 21 workers and 3 females, Tai Forest,
17.x. 1980 (V. Mahnert & J.-L. Ferret); 1 worker, Sassandra, 10 km from Monogaga, 16. iii. 1977(7. Lobl); 1
worker, Abidjan, Banco Forest, ii.1963 (W. L. Brown) (MHN; BMNH; MCZ; ENSA).
Of the three known species of this group which have the postpetiolar disc sculptured, minkara is
immediately identifiable by its very long narrow head and relatively long scapes.
Smithistruma nykara sp. n.
HOLOTYPE WORKER. TL 2-4, HL 0-66, HW 0-43, CI 65, ML 0-07, MI 12, SL 0-31, SI 72, PW 0-29, AL 0-63.
Basal lamella of mandible not visible, what can be seen of dentition as described for enkara. Anterior
clypeal margin transverse, the lateral margins very shallowly convex and feebly convergent anteriorly.
Clypeus laterally and dorsally with fine short simple ground-pilosity which is mostly anteriorly curved and
quite closely applied to the surface, and also with conspicuous much longer simple stouter hairs which are
blunt apically. In profile these long hairs arise almost vertically from the clypeal dorsum, are shorter
anteriorly and longest posteriorly where they form a transverse row of 4. In full-face view the long hairs
project laterally or anterolaterally from the margins and are upcurved in the apical half to one-third of their
length. Sides of head with numerous projecting fine simple hairs which are feebly flagellate, arched or
looped. Dorsum of head behind clypeus with short anteriorly curved ground-pilosity such as is seen on the
clypeus but towards the vertex and from the vertex to the occipital margin with fine simple hairs which are
short flagellate, arched or looped. Long stout hairs such as those described on the clypeus are absent from
the cephalic dorsum proper. Dorsum of head reticulate-rugulose, the clypeus less regularly rugulose.
Antennal scapes feebly bent at about the basal third, broadest just distal to this. Leading edge of scape with
a series of simple long projecting curved hairs. Maximum diameter of eye 0-14xHW. Pronotum not
marginate laterally, without a median longitudinal ridge or carina dorsally. In profile the metanotal groove
not impressed, the propodeal teeth broad basally but narrowly triangular at apex, and with a narrow but
distinct infradental lamella. Dorsal surfaces of pronotum, mesonotum, petiole, postpetiole and gaster with
numerous fine simple hairs which are mostly short flagellate but some of which are curved or looped
308 BARRY BOLTON
apically. Dorsal (outer) surfaces of middle and hind tibiae with numerous simple projecting hairs, many of
which are curved or subflagellate. Sides of pronotum reticulate-rugose, pleurae densely punctate. Dorsal
alitrunk everywhere finely but strongly reticulate-rugose, the spaces between the rugae not punctate
except posteriorly on the propodeum where they form the main sculpture between the bases of the teeth.
Petiole dorsum reticulate-rugose and the anterior face with a narrow transverse crest; the disc of the
postpetiole strongly longitudinally rugose. Basigastral costulae strongly developed, covering the basal
third or slightly more of the tergite. With pedicel segments in profile the spongiform appendages strongly
developed. In dorsal view the petiole node with a narrow posterior strip which is broadest posterolaterally
and interrupted medially. Sides of postpetiole disc not bounded by spongiform tissue in dorsal view.
Posterior margin of postpetiole with a spongiform strip which is broad posterolaterally but concave and
much narrowed medially, and interrupted centrally. Base of first gastral tergite with a transverse strip
which is mostly laminar and is traversed by the basal costulae. Colour medium brown.
PARATYPE WORKERS. TL 2-4-2-6, HL 0-65-0-70, HW 0-44-0-47, CI 66-68, ML 0-07, MI 10-12, SL
0-31-0-34, SI 71-73, PW 0-28-0-32, AL 0-62-0-70 (4 measured).
As holotype, the maximum diameter of the eye 0-14-0- 16 xHW.
Holotype worker, Zimbabwe: Umtali, Melsetter, 1700 m, ii.1969 (R. Mussard) (MHN).
Paratypes. 4 workers with same data as holotype (MHN; BMNH; MCZ).
Related to enkara and minkara by its possession of a sculptured postpetiolar disc, nykara is
separated from the latter by its shorter broader head, punctate pleurae and different cephalic
pilosity. From the former nykara is differentiated by the characters given in the key and noted
under enkara.
Smithistruma placora sp. n.
HOLOTYPE WORKER. TL 2- 1, HL 0-58, HW 0-39, CI 67, ML 0-04, MI 7, SL 0-28, SI 72, PW 0-27, AL 0-58.
Dentition of mandible (from a paratype) as described for malaplax. With the head in full-face view the
anterior clypeal margin very shallowly concave. Sides of clypeus irregular and feebly convex, somewhat
convergent anteriorly and with rounded anterolateral angles. Lateral margins of clypeus in full-face view
with numerous projecting hairs; a lower series of more slender hairs present which are curved anteriorly
and are densest on the posterior halves of the margins, the more anteriorly placed members of this series of
slender hairs may be upcurved apically. Above these finer hairs is a series of much longer stouter cylindrical
hairs which project anterolaterally, are curved upwards or upwards and backward in the apical half to
one-third of their length, and which are feebly clavate apically. The anterior clypeal margin with a few pairs
of weakly clavate very short hairs which are directed towards the midline. Sides of head in full-face view
irregular, with projecting long hairs which are stoutest and most rigid anteriorly on the upper scrobe
margins but which become finer and more flexuous posteriorly on the sides and are weakly flagellate on the
posterior curves of the occipital lobes. With the head in profile the clypeal dorsum with a shallow median
concavity which lacks hairs. In front of this the anterior clypeal convexity is equipped with numerous short
stout weakly clavate hairs which are directed anterodorsally from their bases but which are then curved so
that their apieces point vertically or even posteriorly; the more anteriorly situated members of this group of
hairs are shorter than those nearest the median concavity. Behind the median clypeal concavity is a single
transverse row of longer sinuate weakly clavate hairs whose apices tend to point weakly forwards. Behind
these , at the level of the frontal lobes are similarly constructed but shorter hairs, about half the length of the
posterior clypeal row or slightly more. Dorsum of head behind clypeus with fine simple short ground-
pilosity which is closely applied to the surface and strongly curved anteriorly, and with numerous very long
specialized hairs which are arranged roughly in arched-transverse rows. The anteriormost specialized row
contains the stoutest most rigid hairs, which are slightly curved anteriorly and at least twice longer than the
longest hairs on the clypeal dorsum. The more posterior rows are no shorter but become progressively finer
and more flexuous; those behind the vertex are feebly flagellate. Scape weakly bent at its basal third and
broadest just distal to this, the leading edge and dorsal surface with curved simple projecting hairs.
Maximum diameter of eye 0- 15 x HW. Dorsum of head densely reticulate-rugose. Pronotum not marginate
laterally, without a median longitudinal ridge or carina. Alitrunk in profile lacking a metanotal groove or
impression. Propodeal teeth narrowly triangular and subtended by a narrow evenly concave infradental
lamella. Sides of pronotum and propodeum irregularly rugulose, the pleurae punctate. Promesonotal
dorsum longitudinally rugose with weakly punctulate interspaces. Propodeal dorsum densely punctate
with only vestiges of fine rugulae, the declivity smooth. Dorsum of petiole node irregularly rugose, the
postpetiolar disc smooth and shining. First gastral tergite with dense conspicuous basal costulae. Dorsal
THE AFROTROPICAL DACETINE ANTS 309
surfaces of pronotum , mesonotum , petiole , postpetiole and gaster with numerous long fine flagellate hairs .
Spongiform appendages of pedicel segments massively developed in profile. In dorsal view the petiole
node with a broad posterior spongiform strip which is concave posteromedially. Postpetiole disc complete-
ly surrounded by spongiform material, the margin of the posterior spongiform strip sharply indented
medially. Spongiform band traversing base of first gastral tergite thick and ruff-like. Colour medium
brown, the gaster blackish brown.
PARATYPE WORKERS. TL 2-0-2-1, HL 0-56-0-60, HW 0-36-0-38, CI 63-66, ML 0-04-0-05, MI 7-9, SL
0-24-0-28, SI 68-74, PW 0-24-0-27, AL 0-52-0-58 (3 measured).
As holotype but maximum diameter of eye 0-14-0- 16 xHW.
Holotype worker, Cameroun: Nkoemvon, 2.xi.l980, N49 (D. Jackson) (BMNH).
Paratypes. Cameroun: 2 workers with same data as holotype but 2.iii.l980; 1 worker with same data but
12.x. 1980, N45 (BMNH; MCZ; MHN).
Among the species of the weberi-complex of this group, as characterized by their unsculptured
postpetiolar discs, placora is isolated by its remarkable cephalic pilosity and lack of an impressed
metanotal groove. The closest relatives of placora, fenkara, tolomyla and synkara, are discussed
under the last name.
Smithistruma synkara sp. n.
HOLOTYPE WORKER. TL 2-7, HL 0-76, HW 0-50, CI 66, ML 0-07, MI 9, SL 0-34, SI 68, PW 0-34, AL 0-74.
Dentition of mandible not clearly visible but apparently as described for mdaplax. Anterior clypeal
margin transverse to feebly sinuate. Sides of clypeus irregular, slightly convergent anteriorly and with
rounded anterolateral angles. In full-face view the posterior halves of the sides of the clypeus with a few
simple projecting anteriorly curved fine hairs which are acute apically. Above and forward of these fine
hairs are a number of much longer stouter cylindrical hairs which are blunt apically and which project
anterolaterally, being sharply upcurved in the apical half to one-third of their length. Anterior clypeal
margin with a few much shorter straight hairs which project forward over the mandibles. Sides of head
behind clypeus irregular and with numerous projecting fine hairs, the posteriormost of which are weakly
flagellate. In profile the clypeal and cephalic dorsa with short fine ground-pilosity which is curved anteriorly
and closely applied to the surface, and with long specialized hairs which are stout and simple and pointed to
blunt apically, but not clavate. On the anterior portion of the clypeus the specialized hairs are relatively
short and curve forwards and upwards. Behind them is a shallowly concave area of the clypeus which lacks
hairs and behind this is a transverse row of long erect feebly sinuate hairs which are two or more times
longer than those on the anterior part of the clypeal dorsum. From this level to the vertex all the specialized
long hairs are stout and simple, acute apically and slightly curved forward, all about the same length,
roughly equal to the longest hairs on the clypeal dorsum except for those which are adjacent to the frontal
lobes, which are slightly shorter. Behind the vertex the hairs shorter and more strongly curved, those
closest to the occipital margin finer and weakly flagellate. Entire dorsum of head strongly reticulate-
rugulose. Scapes weakly bent at about the basal third, broadest distal to this and the leading edge and
dorsal surface with long projecting cylindrical curved hairs. Maximum diameter of eye 0-16xHW.
Pronotum not marginate laterally, lacking a median longitudinal ridge or carina dorsally. In profile the
alitrunk lacking a metanotal groove or impression, with narrow sharply triangular propodeal teeth
subtended by a slender infradental lamella whose free posterior margin is concave. Sides of pronotum and
propodeum irregularly rugose, the pleurae punctate. Pronotal dorsum longitudinally rugose with a few
cross-meshes; mesonotal dorsum strongly reticulate-rugose. Propodeal dorsum predominantly punctate,
with faint rugular vestiges. Dorsum of petiole node strongly rugose, the postpetiolar disc smooth and
shining. First gastral tergite smooth and shining except for the dense sharply defined basal costulae. Dorsal
surfaces of pronotum, mesonotum, petiole, postpetiole and gaster with fine dense hairs which are arched,
looped or flagellate. Spongiform appendages of pedicel segments strongly developed in profile. In dorsal
view the petiole node with a thick posterior spongiform strip which is narrowest medially. Disc of
postpetiole completely surrounded by thick spongiform material, the posterior band deeply indented
medially. Base of first gastral tergite with a thick ruff-like transverse spongiform band. Colour dark brown,
the gaster blackish brown.
PARATYPE WORKER. TL 2-8, HL 0-76, HW 0-50, CI 66, ML 0-07, MI 9, SL 0-34, SI 68, PW 0-34, AL 0-72
As holotype.
Holotype worker, Gabon: Makokou, x.1972, rain forest (/. Lieberburg) (MCZ).
Paratype. 1 worker with same data as holotype (BMNH).
310 BARRY BOLTON
Within the weberi-complex four species, fenkara, placora, tolomyla and synkara form a close
association by their mutual lack of postpetiolar sculpture, lack of a metanotal impression and
possession of long specialized hairs on the cephalic dorsum which are similar to those on the
clypeal dorsum. In fenkara, tolomyla and synkara these specialized cephalic hairs tend to be
about equal in size and shape and equal to the longest hairs on the clypeal dorsum, whereas in
placora the size and shape of the specialized hairs are very variable, and those on the cephalic
dorsum tend to be very much longer than any found on the clypeus. 5. fenkara is a smaller more
lightly coloured species than synkara and has the long cephalic hairs conspicuously clavate
(simple in synkara). Finally tolomyla, a smaller species, has a deep median indentation in the
spongiform strip bordering the posterior margin of the postpetiole and has the anterior clypeal
margin shallowly but evenly concave.
Smithistruma tolomyla sp. n.
HOLOTYPE WORKER. TL 2- 1 , HL 0-58, HW 0-39, CI 67, ML 0-06, MI 10, SL 0-27, SI 69, PW 0-27, AL 0-60.
Dentition (from paratype) of a high basal lamella followed by a small diastema, 5 relatively large teeth
forming the principal row, two slightly smaller teeth, 4 minute denticles and a small apical tooth. Anterior
clypeal margin evenly shallowly concave, the anterolateral angles rounded and the sides very feebly
divergent posteriorly. Lateral margins of clypeus with projecting simple hairs, the shorter hairs curved
anteriorly, the longer hairs projecting outwards and upcurved in their apical halves. Sides of head with
numerous long fine projecting hairs. In profile the clypeal dorsum with a few upcurved hairs anteriorly and
a transverse row of much longer erect sinuate hairs across the posterior clypeal margin. Ground-pilosity of
cephalic dorsum behind clypeus of short fine anteriorly arched hairs whose apices are in contact or nearly in
contact with the surface. Specialized pilosity of erect curved to sinuate hairs similar to those on the
posterior clypeus are present on the cephalic dorsum, the longest of them no longer than those on the
posterior clypeus or only very slightly longer. Scape slightly bent in basal third, the leading edge with a
series of freely projecting simple hairs which are upcurved apically. Cephalic dorsum densely and coarsely
reticulate-rugose, the clypeus more finely sculptured. Pronotum not marginate laterally, without a median
longitudinal carina dorsally. Metanotal groove absent. Propodeal teeth fine and narrow, the infradental
lamella reduced to a mere carina which follows the concavity of the declivity. Sides of pronotum and
propodeum reticulate-rugose, the mesopleuron with scattered small sharply incised punctures on a smooth
surface, the metapleuron mostly smooth. Promesonotal dorsum coarsely rugose, the propodeal dorsum
densely punctate. Spongiform appendages of pedicel segments well developed in profile. Ventral appen-
dage of petiole with a broad indentation in its ventral margin at about the midlength. Ventral lobe of
postpetiole massive. Petiole node rugulose in dorsal view, the posterior spongiform strip narrow medially,
its thickness distinctly much less than the dorsal length of the node. Postpetiole in dorsal view smooth and
shining, surrounded on all sides by spongiform material. Anteriorly the postpetiole with a relatively
narrow transverse spongiform strip, laterally the spongiform material increasing thickness posteriorly, the
tissue thickest at the posterolateral angles. Margin of posterior spongiform appendage of postpetiole
indented medially, the indentation reaching the posterior margin of the disc. Base of first gastral tergite
with a dense spongiform strip, the tergite behind this level with short basigastral costulae present. Dorsal
surfaces of alitrunk, petiole, postpetiole and gaster with numerous simple fine hairs. Colour brown.
PARATYPE WORKER. TL 2- 1 , HL 0-58, HW 0-38, CI 66, ML 0-05, MI 9, SL 0-26, SI 68, PW 0-26, AL 0-58. As
holotype.
Holotype worker, Cameroun: nr Yaounde, sample K2 (G. Terrori) (ENSA).
Paratype. 1 worker, Cameroun: nr Yaounde, sample FF (G. Terron) (BMNH).
In the weberi-group six species are known in which the postpetiolar disc is without costulate
sculpture and the metanotal groove is not impressed. These two characters are combined in
arahana, fenkara, malaplax, placora, synkara and tolomyla. The first named is easily dis-
tinguished from the rest as it has the spongiform trip which borders the petiole node posteriorly
very thick indeed, thicker than the dorsal length of the node. In the remainder this strip is quite
narrow, not even approaching the length of the node. Two other species which are quickly
differentiated from tolomyla; malaplax, which lacks specialized long hairs on the cephalic
dorsum similar to those on the clypeal dorsum, and placora, in which such specialized hairs are
present but very much longer on the cephalic dorsum than on the clypeus. The remaining
species, fenkara, synkara and tolomyla, form a close triad. S. fenkara is characterized by the
THE AFROTROPICAL DACETINE ANTS 311
conspicuously swollen nature of the specialized cephalic hairs and synkara is differentiated from
tolomyla by the characters mentioned in the key and the shape of the anterior clypeal margin,
which is concave in the latter species.
Smithistruma weberi Brown
Smithistruma weberi Brown, 1959c: 7, fig. 4. Holotype worker, ZAIRE: Ango, ii.-iii.1948, no 2170 (N. A.
Weber} (MCZ) [examined].
WORKER. TL 2-3, HL 0-61, HW 0-39, CI 64, ML 0-07, MI 11, SL 0-28, SI 72, PW 0-28, AL 0-58.
Basal lamella of mandible a high truncated rectangle with concave sides. Basalmost tooth on mandible
small, separated from the basal lamella by a small diastema. Second tooth from base the longest, the third
about twice longer than the basalmost. The three teeth of the principal row following the second (longest)
tooth are about the same size and are followed distally by 2 smaller teeth, 4 minute denticles and a small
apical tooth. Anterior clypeal margin more or less transverse, only very feebly sinuate. Lateral clypeal
margins irregular, feebly convergent anteriorly and with rounded anterolateral corners. With the head in
full-face view the lateral clypeal margins with a few anteriorly curved simple short hairs and with a number
of anterolaterally or laterally projecting stout long hairs which are upcurved in their apical halves and
feebly clavate apically. Such hairs also present on clypeal dorsum where they curve posteromedially, and
on the sides of the head where they curve upwards and forwards. Dorsum of head behind clypeus with small
simple anteriorly curved hairs which are closely applied to the surface and with longer stout hairs similar in
shape and size to those on the clypeus, the longer subclavate hairs feebly curved anteriorly or anterome-
dially. Cephalic dorsum strongly densely reticulate-rugulose. Antennal scape weakly bent in its basal third,
broadest at about the midlength and having the leading edge equipped with freely projecting curved hairs
which also occur on its dorsal surface. Maximum diameter of eye 0-18xHW. Pronotum not marginate
laterally and without a median longitudinal ridge or carina dorsally. With the alitrunk in profile the
metanotal groove distinctly impressed. Propodeal teeth very small and triangular, subtended by a
conspicuous infradental lamella whose free posterior margin is almost straight, not evenly concave as is
usual in this group. Dorsal surfaces of pronotum, mesonotum, petiole, postpetiole and gaster with
numerous simple fine hairs which may be subflagellate, looped or arched, without large subclavate hairs
similar to those on the head. Sides of pronotum and propodeum rugulose, the mesopleuron finely punctate
and the metapleuron almost smooth. Dorsum of pronotum and mesonotum densely reticulate-rugulose,
the propodeal dorsum densely punctate and the declivity smooth. Petiole dorsum irregularly rugulose;
postpetiolar disc smooth. First gastral tergite with sharply defined but short basal costulae. Spongiform
appendages of pedicel segments strongly developed in profile. In dorsal view the petiole node with a broad
posterior spongiform strip whose free margin is shallowly concave medially. Disc of postpetiole sur-
rounded by spongiform material, the strip bordering the posterior margin broadest posterolaterally,
narrowing medially and sharply indented at the midpoint, the indentation reaching the margin of the disc
itself. Base of first gastral tergite with a broad band of spongiform material from which the basigastral
costulae emerge. Colour medium brown.
Known only from the holotype weberi is one of three species in the group which combine an
unsculptured postpetiolar disc and an impressed metanotal groove. The other species showing
these two characters together are kerasma and mekaha\ details for separating these two from
weberi are tabulated under kerasma.
MATERIAL EXAMINED
Zaire: Ango (N. A. Weber).
The marginata-group
(Fig. 17)
Antennae with 4 or 6 segments. Basal lamella of mandible a long low lobe followed by a principal dental
row of 7 teeth, without a diastema. Anterior clypeal margin broadly and shallowly convex in full-face view
and the sides of the clypeus roughly parallel, not convergent anteriorly throughout their length. Lateral
and anterior margins of clypeus, and clypeal dorsum, lacking hairs of any description. Body hairs sparse,
fine and simple. Long flagellate hairs present on dorsal margins of the antennal scrobes and on the pronotal
humeri. Leading edges of scapes without projecting hairs, any hairs which do occur here are minute and
decumbent to appressed. Pronotum not marginate laterally in rusta but sharply marginate in marginata,
both with a median longitudinal carina on the pronotal dorsum. Infradental lamellae on propodeum broad.
312 BARRY BOLTON
Of the two species recognized in this small group rusta, known only from Zimbabwe, has 6
antennal segments and lacks lateral pronotal margination, whilst the more widely distributed
marginata, from Ivory Coast, Kenya and Zimbabwe, has only 4 antennal segments and possesses
strong lateral pronotal margination. Despite these marked differences I regard both species as
belonging in the same group as they have the same very characteristic clypeal structure, head
shape, body pilosity and distribution of flagellate hairs.
5. marginata was previously included in the now disbanded genus Miccostruma, as discussed
in the introduction to the genus.
Smithistruma marginata (Santschi) comb. n.
Epitritusmarginatus Santschi, 1914o: 114, fig. 21. Syntype workers, KENYA: Shimoni, st. no. 9,xi.l911 (Ch.
Alluaud & R. Jeannel) (NMB) [examined].
Miccostruma marginata (Santschi) Brown, 1948: 123.
WORKER. TL 1-2-1-3, HL 0-40-0-43, HW 0-26-0-28, CI 64-68, ML 0-04, MI 8-10, SL 0-17-0-18, SI 64-67,
PW 0-16-0-18, AL 0-37-0-39 (10 measured).
Mandible with a low basal lamella, not a high triangle or high rectangle with concave sides, the lamella
not or just visible when the mandibles are closed. No diastema between basal lamella of mandible and
basalmost tooth. Principal dental row consisting of 7 teeth, followed by 4 minute denticles and a small
apical tooth. In full-face view the outer margins of the fully closed mandibles diverging posteriorly but
intersecting the shallowly convex anterior margin of the clypeus well in from the rounded anterolateral
angles; the outer margins of the mandibles and the lateral clypeal margins not forming a more or less
continuous line. Clypeus absolutely devoid of hairs, without fringing pilosity and lacking dorsal pilosity.
Lateral margins of clypeus straight and parallel, rounding anteriorly into the shallowly convex anterior
margin, continuous posteriorly with the parallel preocular laminae. Disc of clypeus without a tumulus, with
scattered minute pubescence visible under high magnification and with its posteriormost portion slightly
raised into a low blunt prominence between the frontal lobes. Upper scrobe margins in full-face view
evenly curved-divergent behind the frontal lobes, with a maximum of three laterally projecting flagellate
hairs on each side, though these seem to be lost easily by abrasion. Dorsum of head with very fine sparse
simple curved short ground-pilosity and with two pairs of long curved to flagellate hairs. Dorsum of clypeus
finely shagreened or granular, dorsum of head finely reticulate-punctate. Antennae with 4 segments, the
scapes curved in the basal third, not dorsoventrally flattened beyond the curve; their leading edges with fine
apically curved simple hairs which are decumbent to appressed. Eyes small, their maximum diameter only
about 0-07XHW and distinctly less than the maximum width of the scape. Pronotum sharply marginate
anteriorly and laterally and with a strong mid-dorsal longitudinal ridge or carina which may be doubled for
part or most of its length. Mesonotum laterally less strongly marginate than pronotum but propodeum
sharply marginate to the base of the teeth. Mesonotal dorsum usually with a continuation of the pronotal
median carina but this may be poorly developed or faint in some individuals. The dorsal alitrunk with a
transverse crest or slightly raised step between the mesonotum and propodeum. Dorsal alitrunk with
scattered sparse ground-pilosity which is short fine and decumbent, and with three pairs of flagellate hairs
distributed as follows. First pair on pronotal humeri, directed dorsolaterally; second pair at approximate
midlength of lateral pronotal margination, directed dorsally; third pair on mesonotal margin posteriorly,
close to the transverse crest, directed dorsally. Propodeal teeth laminar and continuous with the broad
infradental lamellae which run the length of the declivity on each side. Sides of alitrunk unsculptured.
Pronotal dorsum mostly smooth but in some with the faintest vestiges of patchy superficial sculpture.
Mesonotum posteriorly with faint vestiges of reticular sculpture; propodeum smooth. Spongiform
appendages of petiole and postpetiole strongly developed and very voluminous in profile, but in dorsal
view only the posterior margin of each segment bounded by spongiform tissue and on the postpetiole the
transverse spongiform material is interrupted posteromedially. Disc of postpetiole unsculptured, its
posterior margin slightly indented medially. Both pedicel segments with fine curved hairs, some of which
are long and subflagellate. First gastral tergite with 5 or 6 basigastral costulae on each side of the midline,
otherwise the gaster unsculptured. Gastral pilosity simple and sparse, consisting of scattered fine short
hairs which are decumbent to appressed and even sparser suberect to erect fine hairs which are longer. Legs
with appressed pubescence only, without standing hairs. Colour uniform dull yellow to yellowish brown.
One of the few Smithistruma species to have 4-segmented antennae, marginata is separated from
all others with this antennomere count by the shape of its clypeus and lack of clypeal pilosity, by
THE AFROTROPICAL DACETINE ANTS 313
its strongly marginate pronotum and possession of a median longitudinal ridge or carina on the
pronotal dorsum.
MATERIAL EXAMINED
Ivory Coast: Man (V. Mahnert & J.-L. Ferret); Adiopodoume (V. Mahnert & J.-L. Ferret). Kenya:
Shimoni (Ch. Alluaud & R. Jeannel); Lamu, nr Witu (V. Mahnert & J.-L. Ferret). Zimbabwe: Umtali,
Melsetter (R. Mussard).
Smithtstruma rusta sp. n.
(Fig. 17)
HOLOTYPE WORKER. TL 2-0, HL 0-55, HW 0-37, CI 67, ML 0-07, MI 13, SL 0-27, SI 73, PW 0-24, AL 0-52.
Mandible with a principal dental row of 7 teeth of approximately the same size, followed distally by 4
minute denticles and a small apical tooth. Basal lamella of mandible concealed by clypeus. Anterior clypeal
margin evenly broadly shallowly convex, the lateral margins more or less straight and parallel, not evenly
convergent anteriorly throughout their length. Outer margins of closed mandibles in full-face view
intersecting the anterior clypeal margin some distance in from the anterolateral angles, the outer
mandibular margins and lateral clypeal margins not forming a more or less continuous line. Anterior and
lateral clypeal margins without projecting hairs of any description, dorsum of clypeus without hairs.
Dorsum of head behind clypeus with scattered simple fine hairs which are arched and decumbent, or
appressed. Sides of head with long flagellate hairs present. Each member of the type-series has lost
some flagellate hairs, which seem easily displaced by abrasion; the maximum number of flagellate cephalic
hairs appears to be as follows. One pair posterolaterally on the occipital lobes which may be directed
upwards or outwards; one pair directed laterally from the posteriormost point of the upper scrobe margins;
one pair arising from the side of the head just above the last-mentioned pair and tending to be directed
upwards rather than outwards; one pair more anteriorly situated on the upper scrobe margin, about on a
level with the anterior margin of the eye. Preocular laminae in full-face view more or less parallel.
Antennae with 6 segments, the scape narrow and not strongly flattened, bent approximately at its basal
third. Leading edge of scape lacking a series of anteriorly projecting hairs, only with short decumbent to
appressed fine pubescence. Maximum diameter of eye 0-llxHW. Dorsum of clypeus closely punctulate,
cephalic dorsum strongly reticulate-punctate everywhere except for a narrowly triangular smooth area
running back from the posterior clypeal margin between the frontal lobes. Anterior border of pronotum
sharply transversely marginate, the sides of the pronotum not marginate. Pronotal and mesonotal dorsa
with a median longitudinal ridge or carina. Posterior half of mesonotum and all of propodeum narrowly
marginate laterally. With alitrunk in profile the lateral mesonotal-propodeal margination continuous,
without trace of a metanotal groove; however, mid-dorsally the median mesonotal ridge or carina ends at a
distinct step-down at its junction with the propodeum. Propodeal teeth broadly triangular and with a
conspicuous infradental lamella. Pronotal humeri each with a long fine flagellate hair. Dorsal alitrunk with
2-3 pairs of long fine curved hairs which are simple and erect, and with several pairs of decumbent to
appressed fine simple short hairs. Dorsal surfaces of petiole and postpetiole with sparse but conspicuous
erect to suberect fine hairs. Sides of alitrunk smooth and shining, with marginal feeble sculpture dorsal to
and posterior to the extensive smooth area. Pronotal dorsum with 2-3 feeble longitudinal costulae on each
side of the median ridge or carina, the spaces between the costulae filled with broad shallow superficial
punctures. Mesonotum, propodeal dorsum, propodeal declivity between the teeth and petiole node
densely punctate. Disc of postpetiole glassy smooth. First gastral tergite unsculptured except for the sparse
widely spaced basigastral costulae. Spongiform appendages of pedicel segments strongly developed in
profile. Petiole node in dorsal view with a narrow posterior lamina. Postpetiole in dorsal view with a narrow
lamina on the anterior margin and with the lateral spongiform material visible projecting beyond the lateral
margins of the disc. Posterior margin of postpetiole with a laminar rather than spongiform transverse strip;
broadest laterally and narrowing medially where the posterior margin of the postpetiolar disc itself is
indented. Base of first gastral tergite with a narrow laminar strip which is traversed by the sparse basigastral
costulae. Colour glossy light brown.
PARATYPE WORKERS. TL 2-0-2-1, HL 0-54-0-58, HW 0-36-0-38, CI 63-69, ML 0-07-0-08. MI 12-14, SL
0-26-0-28, SI 70-78, PW 0-24-0-26, AL 0-52-0-56 (10 measured). As holotype.
Holotype worker, Zimbabwe: Umtali, Melsetter, 1700 m, ii.1969 (R. Mussard) (MHN).
Paratypes. 10 workers with same data as holotype (MHN; BMNH; MCZ; ENSA).
Related to marginata by the characters discussed in the species-group diagnosis, rusta is quickly
314 BARRY BOLTON
separated from all other Afrotropical Smithistruma presently known by its combination of these
characters with 6-segmented antennae.
The oxysma-group
(Figs 18, 19)
Antennae with 6 segments. Basal lamella of mandible a low lobe, principle dental row of 7 teeth; no
diastema between the basal lamella and the basalmost tooth. Sculpture of head and body fine. Anterior
clypeal margin prominent and narrowly rounded in full-face view, the sides more or less evenly convergent
anteriorly and forming an approximately continuous line with the outer margins of the closed mandibles.
Sides of clypeus without a fringing row of spatulate or spoon-shaped anteriorly curved hairs. Dorsum of
clypeus with weakly clavate hairs which are curved posteriorly or posteromedially, the anteriormost one or
two pairs of these being visible in full-face view as they project beyond the clypeal margin, close to the point
where the clypeus overlaps the mandibles, and curve outwards and backwards. Body pilosity sparse, fine
and simple. Long flagellate hairs present on pronotal humeri, present or absent on upper scrobe margins.
Leading edges of antennal scapes without projecting hairs, any hairs which do occur here are short and
decumbent to appressed. Pronotum not marginate laterally but with a median dorsal longitudinal carina
present. Propodeal infradental lamellae broad and well developed.
The two species of this small group, anarta and oxysma, are presently known only from South
Africa. They are characterized primarily by the form and pilosity of the clypeus, characters not
shared with any other Afrotropical species, although the shape is duplicated in the tacta-group.
In this last-named group, however, the clypeal pilosity is radically different and the antennae
have only 4 segments.
The closest relatives of the oxysma-group appear to belong to the New World ornate-group,
which contains three species showing the clypeal shape and pilosity noted above (Brown, 1953a:
64), but in ornata and its relatives the mandibles have a long diastema between the basal lamella
and the basalmost tooth, a character not observed in the oxys ma-group. At present I am
uncertain how important this character is, so I feel it is best to keep the New World and
Afrotropical species in separate groups until it can be investigated in more detail.
Smithistruma anarta sp. n.
(Fig. 18)
HOLOTYPE WORKER. TL 1 -9, HL 0-54, HW 0-34, CI 63, ML 0-06, MI 11 , SL 0-27, SI 79, PW 0-21, AL 0-48.
Principal dental row of mandible with 7 teeth, followed by 4 minute denticles and a small apical tooth.
Basal lamella of mandible (concealed by clypeus in holotype) a long low rounded lobe which is only as high
as the basalmost tooth; no diastema between basal lamella and basalmost tooth. In full-face view the
clypeus with shallowly convex sides which are evenly convergent anteriorly and with a strongly convex
anterior margin which is narrowly rounded medially; the anterior margin on each side of the midpoint
forms a single evenly convex line which is continuous with the lateral margins, without trace of an
anterolateral angle. Outer margins of the fully closed mandibles forming a more or less continuous line
with the outer margins of the clypeus in full-face view. Clypeal margins without a fringe of anteriorly or
medially curved large spatulate hairs but one or two simple short hairs may occur posterolaterally. The
dorsum of the clypeus along the anterior margin with 3 pairs of short recurved spatulate hairs which curve
upwards and backwards from the clypeal edge. Behind this anterior row the clypeal dorsum with 12 similar
curved hairs. These 12 make up four pairs which are situated on each side of the midline and which curve
backwards and towards the midline, the posteriormost pair being at the posteromedian clypeal apex; a pair
on the posterior clypeal margin immediately in front of the anteriormost part of the frontal lobes, curved in
the direction of the clypeal margin; and a pair situated posterolaterally on the clypeus, curved towards the
midline and slightly backwards. Cephalic dorsum behind clypeus with subdecumbent to decumbent short
narrowly spatulate hairs which are curved towards the highest point of the vertex. Flagellate hairs absent
from dorsum of head and from upper scrobe margins. Antennal scapes slender, not flattened, narrowed
basally and bent at about the basal quarter. Leading edges of scapes without a freely projecting row of long
hairs, only with short fine pubescence which is decumbent to appressed. Maximum diameter of eye
0-12xHW. Clypeus finely punctulate, cephalic dorsum reticulate-punctate. Anterior pronotal border
sharply transversely marginate. Sides of pronotum not marginate but sides of mesonotum and propodeum
angulate. Pronotum with a median longitudinal ridge or carina dorsally. Metanotal groove visible on the
THE AFROTROPICAL DACETINE ANTS 315
dorsal alitrunk but not impressed in profile. Propodeal teeth short and confluent with the broad infradental
lamellae. Pronotal humeri each with a long flagellate hair. Posterodorsally on the mesonotum is a pair of
somewhat flattened hairs which are markedly curved towards the midline and are notched apically. Dorsal
ground-pilosity of alitrunk consists of a very few decumbent to appressed scattered short hairs, most easily
visible on the anterior half of the pronotum. Dorsal surfaces of petiole and postpetiole with numerous
back-curved hairs. First gastral tergite with 4 standing hairs only, which are blunted or notched apically and
arranged in a transverse row close to the base of the sclerite. Behind this are sparse flattened short
appressed hairs on the remainder of the tergite which are directed towards the midline. Sides of alitrunk
unsculptured. Dorsal alitrunk unsculptured apart from the median carina and some extremely faint, almost
effaced, sculptural vestiges on the promesonotum. Dorsum of petiole node densely punctate, postpetiolar
disc glassy smooth. First gastral tergite unsculptured except for the sparse basal costulae which are
arranged on each side of a central clear area. Spongifirm appendages of pedicel segments strongly
developed in profile. In dorsal view the petiole node bounded posteriorly by a narrow lamellar strip.
Shallowly concave anterior margin of postpetiole with a narrow lamellar strip. Ventrolateral spongiform
appendages of postpetiole not visible in dorsal view. Posterior margin of postpetiole with a lamellar strip
and the margin of the disc indented medially. Basal border of first gastral tergite with a sinuate lamella
whose free margin is concave medially and convex towards the sides. Colour dull glossy yellow.
PARATYPE WORKER. TL 1-9, HL 0-55, HW 0-36, CI 65, ML 0-06, MI 11, SL 0-28, SI 78, PW 0-22, AL 0-50.
Asholotype.
Holotype worker, South Africa: Natal, Dukuduku Forest Res., 12-15 km E. of Mtubatuba, coast vine
forest on sand, 26.1.1977 (W. L. & D. W. Brown) (MCZ).
Paratype. 1 worker with same data as holotype (BMNH).
To the present only two species of this group are known. They are separated easily as in anarta
the clypeal dorsum has 18 recurved hairs, the upper scrobe margins lack flagellate hairs, the
cephalic dorsum lacks flagellate hairs, and the first gastral tergite has only 4 standing hairs
arranged in a transverse row close to the base of the sclerite. In contrast oxysma has 12 recurved
hairs on the clypeal dorsum, has 3 pairs of flagellate hairs on the upper scrobe margins, has
flagellate hairs present on the cephalic dorsum, and has 12 or more standing hairs on the first
gastral tergite which are distributed all over the sclerite.
Smithistruma oxysma sp. n.
(Fig. 19)
HOLOTYPE WORKER. TL 2- 1 , HL 0-58, HW 0-36, CI 62, ML 0-07, MI 12, SL 0-27, SI 75, PW 0-22, AL 0-54.
Principal dental row of mandible with 7 teeth followed by 4 minute denticles and a small apical tooth.
Basal lamella of mandible (concealed by clypeus in holotype) a long low rounded lobe which is only as high
as the basalmost tooth; without a diastema between the basal lamella and the basalmost tooth. In full-face
view the clypeus with shallowly convex sides which are evenly convergent anteriorly and with a strongly
convex anterior margin which is narrowly rounded medially; the anterior margin on each side of the
midpoint forms a single evenly convex line which is continuous with the lateral margins, without trace of
anterolateral angles. Outer margins of the fully closed mandibles forming a more or less continuous line
with the outer margins of the clypeus in full-face view. Clypeal margins without a fringe of curved spatulate
to spoon-shaped hairs but one or two simple short hairs may occur laterally. The dorsum of the clypeus
along the anterior margin with three pairs of flattened, apically gradually clavate, recurved hairs. The
innermost pair arises very close to the clypeal apex and curves up and back without breaking the clypeal
outline. The two outer pairs curve outwards as well as upwards and back and project beyond the clypeal
outline in full-face view. Dorsum of clypeus behind the anterior margin also with 6 hairs which are similar in
shape to those just described, arranged in a transverse band of 4 behind the midlength which curve
backwards and towards the midline, and a pair close to the posterior clypeal margin which are strongly
arched towards the midline and only feebly curved backwards. Ground-pilosity of cephalic dorsum of
numerous simple to very narrowly spatulate fine hairs which are subdecumbent to decumbent and
generally curved towards the highest point of the vertex. Upper scrobe margins with three pairs of
flagellate hairs; the posterior occipital lobes with a pair of flagellate hairs; 4 other pairs of flagellate hairs
present on the dorsum of the head, making a total of 16 cephalic flagellate hairs. Antennal scapes slender
and more or less cylindrical, slightly narrowed basally and very weakly bent at the basal quarter. Leading
edges of scapes without strong freely projecting hairs, only with decumbent fine pubescence. Maximum
316 BARRY BOLTON
diameter of eye 0-llxHW. Clypeal dorsum very finely punctate-granular, the cephalic dorsum strongly
reticulate-punctate. Anterior border of pronotum transversely marginate; sides of pronotum not margin-
ate but sides of mesonotum and propodeum angular. A mid-dorsal longitudinal ridge or carina present on
pronotum and mesonotum, absent from propodeal dorsum. With the alitrunk in profile the metanotal
groove absent and the propodeal teeth confluent with the broad and conspicuous infradental lamellae.
Pronotal humeri with a pair of long flagellate hairs which are directed predominantly laterally. Pronotal
dorsum behind the humeri with a pair of shorter flagellate hairs and with 2-3 pairs of anteriorly situated fine
decumbent shorter hairs. Mesonotal dorsum with two pairs of fine hairs. Fine simple hairs numerous on
dorsal surfaces of petiole and postpetiole, and such hairs widely distributed on the first gastral tergite where
12 or more are present. Sides of alitrunk unsculptured except for a few feeble longitudinal striae close to the
anterior pronotal margination. Pronotum and mesonotum dorsally with scattered weak longitudinal
rugulae or costulae most of which are short, the spaces between them mostly smooth on the pronotum but
on the mesonotum with vestiges of punctate sculpture also visible. Propodeal dorsum smooth, with two
weak rugulae running from the apex of the median mesonotal ridge across the dorsum to the bases of the
propodeal teeth. Dorsum of petiole node densely punctate; disc of postpetiole smooth and shining. First
gastral tergite unsculptured except for the basigastral costulae which arise in two patches, on each side of a
median clear area. Spongiform appendages of pedicel segments strongly developed in profile. In dorsal
view the petiole node bounded posteriorly by a narrow lamellate strip. Postpetiole in dorsal view with the
anterior margin sharply concave medially and with a short narrow transverse lamella bordering the
concave section. The ventrolaterally situated spongiform appendages of the postpetiole are visible in
dorsal view, projecting anterolaterally and laterally beyond the outline of the disc. Posterior margin of
postpetiole bordered by a lamellate strip which is very broad posterolaterally. First gastral tergite with a
sinuous basal lamellar strip which has its anterior free border convex at the sides and concave medially, and
which is traversed by the basigastral costulae. Colour glossy medium brown.
PARATYPE WORKERS. TL 1-9-2-1, HL 0-50-0-58, HW 0-32-0-38, CI 62-67, ML 0-06-0-08, MI 11-14, SL
0-25-0-30, SI 72-80, PW 0-21-0-26, AL 0-50-0-56 (15 measured).
Maximum diameter of eye 0-11-0-14XHW. Mostly as holotype but propodeal dorsum may be unsculp-
tured and the mesonotum may lack any trace of punctate sculpture. The long recurved hairs on the clypeus
appear to be easily lost by abrasion, especially those of the anterior row, and the long flagellate hairs of the
head may be flattened to the surface by accident of preservation.
Holotype worker, South Africa: Natal, 75 km WSW. Estcourt, Cathedral Peaks Forest Sta.,
7-31.xii.1979, Her. 8, 17.xii.1979, podocarp forest rotted stump of Cussonia spicata (S. &J. Peck) (MCZ).
Paratypes. South Africa: 8 workers and 1 female with same data as holotype; 5 workers with same data
but Ber. 19, 24.xii.1979, podocarp forest rotted wood, moss, fleshy and woody fungi, 1500 m; 2 workers
with same data but Ber. 18, 24.xii.1979, podocarp forest rotted fruit bait 1500 m (MCZ; BMNH; MHN).
Non-paratypic material examined. South Africa: Natal, Pietermaritzburg (W. L. & D. E. Brown).
Lesotho: Mamathes (C. Jacot-Guillarmod).
The only closely related species is anarta; details of their separation are given under that name.
The tecte-group
(Fig. 20)
Antennae with 4 segments, the second funicular long and obviously a fusion segment. Basal lamella of
mandible a low lobe, the principal dental row of 7 teeth, without a diastema between the basal lamella and
the basalmost tooth. Sculpture of head and body fine. Anterior clypeal margin prominent and narrowly
rounded in full-face view, the sides more or less evenly convergent anteriorly and forming an approximate-
ly continuous line with the outer margins of the closed mandibles. Margins and dorsum of clypeus with
dense fine simple short hairs, without specialized or bizarre pilosity. Body pilosity fine and simple. Long
flagellate hairs present on pronotal humeri and upper scrobe margins. Leading edges of scapes without
projecting stout hairs. Pronotum dorsally with a median longitudinal carina. Propodeal infradental
lamellae present.
The two species in this group, tacta and vodensa from West and central Africa, may be derived
from the oxysma-group. The clypeal structure is strikingly similar in the two groups and most
other characters diagnostic at species-group level are in accord. The main differences between
the groups lie in the reduced antennomere count in tacta and vodensa, and their lack of
specialized strong hairs on the clypeus, which are so obvious in the oxysma-group.
THE AFROTROPICAL DACETINE ANTS 317
Smithistruma tacta sp. n.
HOLOTYPE WORKER. TL 1-7, HL 046, HW 0-31, CI 67, ML 0-07, MI 15, SL 0-20, SI 65, PW 0-20, AL 0-46.
Principal dental row of mandible with 7 teeth followed by 4 minute denticles and a small apical tooth.
Basal lamella of mandible a long low rounded lobe which is no higher than the basalmost tooth; without a
diastema between basal lamella and the basalmost tooth. In full-face view the clypeus with shallowly
convex sides which are evenly convergent anteriorly and with a strongly convex anterior margin which is
narrowly rounded medially. The anterior margin on each side of the midpoint forms a single evenly convex
line which is continuous with the lateral margins, without trace of an anterolateral angle. Outer margins of
the fully closed mandibles forming a more or less continuous line with the outer margins of the clypeus in
full-face view. Dorsum of clypeus densely clothed with short spatulate hairs which are curved, decumbent
and directed anteriorly. Lateral and anterior clypeal margins similarly densely clothed. Dorsum of head
with decumbent curved fine hairs, those in front of the vertex directed towards the midline, the remainder
directed towards the highest point of the vertex. Upper scrobe margins with a number of fine curved hairs
similar to those on the dorsum of the head, and also with three pairs of long laterally projecting flagellate
hairs; the cephalic dorsum near the occipital margin with another pair of flagellate hairs which are directed
vertically. Antennae with 4 segments; the scape slender and only very weakly curved basally, not flattened.
Leading edge of scape without a freely projecting row of strong hairs, only with fine curved pubescence
which is subdecumbent to decumbent. Eyes small, the maximum diameter 0-06xHW. Cephalic dorsum
reticulate-punctate everywhere, clypeal dorsum more finely punctate but the sculpture partially concealed
by the dense pilosity. Pronotum strongly and sharply marginate anteriorly and laterally, the pronotal
dorsum with a strong median longitudinal ridge or carina which does not extend onto the mesonotum.
Sides of mesonotum angular, of propodeum sharply marginate. Alitrunk in dorsal view with a transverse
straight line between the mesonotum and propodeum. Pronotal humeri each with a long laterally directed
flagellate hair. Vertically directed flagellate hairs present in pairs on dorsum at midlength of pronotal
lateral margination and posterolaterally on mesonotum. Pilosity of dorsal alitrunk otherwise of fine simple
strongly curved hairs on the pronotum and mesonotum. Fine simple hairs, some of which may be looped
apically, present on dorsal surfaces of petiole, postpetiole and first gastral tergite. Sides of alitrunk
unsculptured, the propodeal teeth lamelliform and continuous with the infradental lamellae. Pronotal
dorsum unsculptured apart from the strong median carina. Mesonotum reticulate-punctate. Propodeal
dorsum and declivity glassy smooth. Dorsum of petiole node finely punctate, postpetiole glassy smooth.
First gastral tergite unsculptured except for the basigastral costulae. Spongiform appendages of pedicel
segments strongly developed in profile. Petiole node in dorsal view with a broad strip of spongiform
material posteriorly. Anterior margin of postpetiole with a narrow spongiform strip, the posterior margin
with a much broader band of spongiform tissue which is indented medially. Base of first gastral tergite with
a spongiform to lamellar strip which is concave medially. Colour yellow to yellowish brown.
PARATYPE WORKERS. TL 1-7-1-8, HL 0-42-0-46, HW 0-29-0-31, CI 65-70, ML 0-06-0-08, MI 14-17, SL
0-18-0-20, SI 61-67, PW 0-19-0-22, AL 0-44-0-48 (10 measured). As holotype.
Holotype worker, Ivory Coast: Droplieu, lO.x.1980 (V. Mahnert &J.-L. Ferret) (MNH).
Paratypes. Ivory Coast: 2 workers with same data as holotype; 7 workers, Monogaga, 24.x. 1980 (V.
Mahnert & J.-L. Ferret); 1 worker, Man, Mt Tonkoui, 900 m, 13.x. 1980 (V. Mahnert & J.-L. Ferret)
(MHN; BMNH; MCZ).
Non-paratypic material examined. Ghana: Tafo (B. Bolton). Cameroun: Nkoemvon (D. Jackson); nr
Yaounde (G. Terron). Zaire: Yangambi (M. Maldague).
Of the seven known Afrotropical species with 4-segmented antennae only two, tacta and
marginata, have the pronotum sharply marginate laterally and equipped medially with a
longitudinal carina. These two are separated by the shape and pilosity of the clypeus as indicated
in the key, and by the characters noted in the species-group diagnoses. Characters separating
tacta and vodensa are tabulated under the latter.
Smithistruma vodensa sp. n.
(Fig. 20)
HOLOTYPE WORKER. TL 3-0, HL 0-74, HW 0-38, CI 51 , ML 0-09, MI 12, SL 0-44, SI 1 16, PW 0-30, AL 0-80.
Principal dental row of mandible with 7 teeth, followed by 4 minute denticles and a small apical tooth.
Basal lamella of mandible a low rounded lobe which is not as high as the basalmost tooth. No diastema
between the basal lamella and the basalmost tooth. In full-face view the clypeus with shallowly convex sides
318 BARRY BOLTON
which are evenly convergent anteriorly and with a strongly convex anterior margin which is narrowly
rounded medially. The anterior clypeal margin on each side of the midpoint forms a single evenly convex
line which is continuous with the lateral margins, without trace of anterolateral angles. Outer margins of
the fully closed mandibles forming a more or less continuous line with the outer margins of the clypeus in
full-face view. Dorsum and margins of clypeus densely clothed with elevated fine simple hairs which are
directed anteriorly. Dorsum of head with decumbent curved fine hairs, those in front of the vertex directed
towards the midline, the remainder directed towards the highest point of the vertex. Upper scrobe margins
with projecting long flagellate hairs. (One pair is visible in the holotype, at the level of the scrobal apex;
more may be present in undamaged specimens, but the holotype is somewhat abraded.) Head long and
narrow, the CI of 51 is the lowest yet recorded in any Afrotropical Smithistruma. Between the posterior
clypeal margin and the frontal lobes, and running back between the lobes, the surface depressed into a
shallow inverted V-shaped trench. Upper scrobe margins pinched in behind the frontal lobes, evenly
convex behind this and confluent with the convex sides of the head. Occipital margin concave and bordered
by a raised lamelliform ridge or flange which is traversed by a number of ribs. In profile the antennal
scrobes reduced and shallow, the preocular laminae low and inconspicuous. Clypeus very densely
punctate-granular, the inverted V-shaped impression smooth. Cephalic dorsum to level of eyes finely
reticulate-punctate, behind this the surface more grossly reticulate-punctate, the punctures becoming
larger both posteriorly and away from the midline, and blanketing the entire surface. Antennae with 4
segments, the scape relatively very long (SI 116, the longest yet recorded among Afrotropical Smithistru-
ma). The second funicular (third antennal) is an extremely long fusion segment constituting funicular
segments 2-4 of the normal 5-merous funiculus ; this second funicular segment almost two times longer than
the first. On the right funiculus vague vestigial constrictions can be seen marking the original segmental
limits, but on the right even these traces are absent. Scapes with simple fine hairs present, without bizarre
pilosity; shaft of scape not bent nor flattened but increasing in thickness from base to apex. Pronotum
marginate anteriorly and with a median longitudinal carina. Sides of pronotum and mesonotum not
marginate but propodeal dorsum meeting the sides in an angle. Metanotal groove broad, deeply
impressed, the impressed area blocked off at each side by a short longitudinal lamina running from the
mesonotum to the propodeum. Propodeal dorsum with a sharp transverse rim bordering the metanotal
impression posteriorly. In profile the propodeal teeth very thin apically, subtended by narrow translucent
infradental lamellae. Sides of alitrunk unsculptured except for some weak peripheral punctation. Pronotal
dorsum smooth near the median carina but laterally with some low disorganized sculpture. Mesonotum
densely punctate. Metanotal groove, propodeal dorsum and declivity smooth except for a few vestigial
punctures which are scarcely visible, situated on the propodeum just behind the transverse rim which
borders the metanotal groove. Pronotal humeri with long flagellate hairs. Dorsal alitrunk with numerous
simple hairs, without bizarre pilosity. Petiole missing from holotype. Postpetiole in profile broadly convex,
the spongiform appendages poorly developed. Ventral appendage of postpetiole represented by a lobate
translucent thin lamina which contains a few stiffening veins but is not spongiform. First gastral sternite
with a basal felt-like fibrous pad which runs across the width of the sclerite, the fibres constituting the felt
running longitudinally. In dorsal view the postpetiole unsculptured, bordered anteriorly by a narrow
ribbed lamina, without lateral appendages. Posteriorly the postpetiole margin bordered by a ribbed lobate
lamina on each side of a broad median gap. Basal lamella of first gastral tergite longitudinally concave and
traversed by costulae which do not impinge upon the sclerite proper. This last only with very faint
scratch-like markings near the base. Postpetiole with a few fine simple hairs and with fine appressed
ground-pilosity. First gastral tergite only with fine appressed ground-pilosity. Colour medium brown.
Holotype worker, Cameroun: Nr Yaounde, sample 3123 (G. Tenon) (ENS A).
The only known species closely related to vodensa is tacta; the two are separated as follows.
tacta vodensa
Head relatively broad, CI 65-70. Head relatively narrow, CI 51 .
Scapes relatively short, SI 61-67. Scapes relatively long, SI 116.
Smaller species, HL 0-42-046. Larger species, HL 0-74.
Hairs on clypeal dorsum spatulate. Hairs on clypeal dorsum simple.
Pronotum sharply marginate laterally. Pronotum not marginate laterally.
Metanotal groove shallow to absent. Metanotal groove broadly, deeply impressed.
Ventral appendage of postpetiole Ventral appendage of postpetiole
spongiform. laminar.
THE AFROTROPICAL DACETINE ANTS 319
TRICHOSCAPA Emery
(Figs 21, 22)
Trichoscapa Emery, 18696: 24 [as subgenus of Strumigenys] . Type-species: Strumigenys (Trichoscapa)
membranifera Emery, 18696: 24, fig. 11, by monotypy.
Trichoscapa Emery; Brown, 1948: 112. [ Raised to genus.]
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles short triangular (MI 16-20), serially dentate
and lacking an apical fork of spiniform teeth. When fully closed the dorsal surface of the mandible with a
sharp conspicuous transverse basal margin which is separated from the anterior clypeal margin by a distinct
impression or gap. Apical (masticatory) margin of mandible with 12 teeth following a basal lamella, the
lamella inflected below the basalmost tooth, not visible when the mandibles are closed. Antennae with 6
segments.
This monotypic genus is very close to Smithistruma, being separated only by the differently
constructed mandibles. The apparent strong transverse basal margin seen in Trichoscapa
(Fig. 21) but not in Smithistruma (Figs 1-12, 14, 15, 17-20) is a secondary development caused
by the dorsal surface of the mandible passing through a sudden downward near right-angled
bend at the level of the basalmost tooth, this sharp downward angle running across the entire
width of the mandible. The basal lamella, which follows the basalmost tooth in approximately
the same plane in Smithistruma, is in Trichoscapa on the inner margin of the descending surface
below the basalmost tooth, and so is usually invisible in full-face view even when the mandibles
are open.
The single species included in Trichoscapa, membranifera, is an accomplished tramp species
in the tropics and the warm temperate zones. Brown & Wilson (1959) suggested an African
origin for the species but at that time no specimens of membranifera had been reported from the
Afrotropical region. A single series from Sierra Leone (in BMNH) shows that the species does
indeed occur in sub-Saharan Africa but whether this continent represents its place of origin
remains to be seen, for this series may also represent an introduction.
The tramping ability of this small ant has ensured that it has accrued more than its fair share of
synonyms. These are dealt with by Brown (1948), and I have no changes nor additions to make
to the list.
Trichoscapa membranifera (Emery)
(Figs 21, 22)
Strumigenys (Trichoscapa) membranifera Emery, 18696: 24, fig. 11. Holotype worker, ITALY: Napoli,
Portici (MCSN) [examined].
Strumigenys membranifera race simillima Emery, 1890: 69, pi. 8, fig. 5. Holotype worker, ST THOMAS I.
(West Indies) (MCSN). [Synonymy by Brown, 1948: 114.]
Strumigenys membranifera var. santschii Forel, 1904o: 6. Syntype workers, TUNISIA: Kairouan (F.
Santschi) (MHN). [Synonymy by Brown, 1948: 114.]
Strumigenys (Cephaloxys) vitiensis Mann, 1921: 461, fig. 22c. Syntype workers, FIJI Is: Vanua Levu,
Lasema (W. M. Mann} (MCZ) [Synonymy by Brown, 1948: 114.]
Strumigenys (Cephaloxys) silvestriana Wheeler, 1928: 27. Syntype workers, MACAO: (F. Silvestri) (MCZ)
[Synonymy by Brown, 1948: 114.]
Strumigenys (Cephaloxys) foochowensis Wheeler, 1928: 28. Holotype female, CHINA: Foochow (F.
Silvestri) (location of type unknown). [Synonymy by Brown, 1948: 114.]
Strumigenys (Cephaloxys) membranifera var. marioni Wheeler, 1933: 276. Syntype workers, U.S.A.:
Mississippi (M. R. Smith) (MCZ). [Synonymy by Brown, 1948: 114.]
Strumigenys (Cephaloxys) membranifera var. \villiamsi Wheeler, 1933: 276. Syntype workers, HAWAII: S.
of Olaa, off the road to Puna, iv. 1932, under moss etc. (F, X. Williams] (MCZ). [Synonymy by Brown,
1948: 114.]
Trichoscapa membranifera (Emery) Brown, 1948: 113.
WORKER. TL 1 -9-2- 1 , HL 0-46-0-50, HW 0-40-0-44, CI 84-90, ML 0-08-0- 10, MI 16-20, SL 0-22-0-24, SI
51-57, PW 0-23-0-26, AL 0-50-0-53 (10 measured).
Mandibles with 12 teeth, arranged in a series of 7 larger teeth basally followed by a series of 4 denticles
and a small apical tooth; the 7 basal teeth not all the same size. Dorsal surface of mandible sharply angled
320 BARRY BOLTON
downwards immediately behind the basalmost tooth, this angle running across the width of the mandible
and forming a sharp transverse basal margin which is separated from the anterior clypeal margin by a gap or
impression. Basal lamella of mandible situated on the descending inner margin below the basalmost tooth
and not visible in full-face view. Anterior clypeal margin transverse to broadly shallowly convex, the
clypeal margins both anteriorly and laterally lacking projecting hairs of any description. Dorsum of clypeus
shining, sometimes with faint sculptural vestiges; dorsum of head behind clypeus reticulate-punctate and
dull. Pilosity of head restricted to a single pair of standing hairs at the highest point of the vertex, otherwise
the dorsum only having minute appressed pubescence which is very sparse and difficult to see. Clypeus and
lateral margins of head hairless, without flagellate or other specialized hairs. Antennal scapes short , bent in
the basal third and broadest at the bend, the leading edge with a row of spatulate to spoon-shaped freely
projecting hairs. Eyes small, of only a few ommatidia, situated at the ventral scrobe margin. Pronotum
strongly marginate anteriorly and laterally, mesonotum and propodeum not marginate. Dorsal outline of
alitrunk in profile very shallowly concave between mesonotum and propodeum but the metanotal groove
absent. Propodeum descending posteriorly to the broad strongly spongiform infradental lamellae.
Separated propodeal teeth absent, either indistinguishable from the large infradental lamellae or at most
forming a minute point close to the dorsum of the lamella. Sides of alitrunk smooth. Pronotal dorsum
smooth or at most with only the very faintest vestiges of sculpture, which may include an extremely faint
median longitudinal ridge. Mesonotum with some fine superficial punctures but these may be very feeble
and difficult to see. Propodeal dorsum and declivity smooth. Standing hairs absent from dorsal alitrunk;
scattered sparse minute appressed pubescence present. Humeral angles of pronotum without flagellate or
other hairs. Spongiform appendages of the pedicel segments massively developed in profile. Petiole
ventrally with a deep curtain-like appendage, the dorsum of the peduncle with a narrow spongiform strip
which runs up almost to the highest point of the node. Lateral spongiform appendages of the petiole node
large and strongly prominent. Lateral and ventral spongiform lobes of postpetiole very large, much larger
than the exposed area of the disc. In dorsal view both petiole and postpetiole smooth, bounded laterally by
dense spongiform tissue. Petiole node also with a posterior spongiform strip linking the large lateral
appendages. Postpetiole also with transverse lamellate spongiform tissue bounding the anterior and
posterior margins. Base of first gastral tergite with a transverse strip which is spongiform laterally but
lamellate medially where it is overlapped by the convex posterior strip of the postpetiole. Basigastral
costulae present, grouped on each side of a median clear area; the gaster otherwise unsculptured. Dorsal
surfaces of petiole, postpetiole and gaster without hairs of any description but with minute appressed very
sparse pubescence. Colour dull yellow to yellowish brown.
Superficially similar to some Smithistruma species, T. membranifera is easily distinguished from
members of that genus by the characteristic form of the mandibles and strongly marginate
pronotum, and by the near absence of standing hairs. Feeding behaviour of membranifera was
investigated by Wilson (1954) who found that it would eat a wide range of small soft bodied
arthropods.
MATERIAL EXAMINED
Sierra Leone: Njala(£. Hargreaves). Egypt: noloc. (Min. of Agriculture coll.); Banage (Alfieri). U.S.A.:
Miss., West Point (E. E. Byrd). Italy: Napoli, Portici.
GLAMYROMYRMEX Wheeler
(Figs 23-33)
Glamyromyrmex Wheeler, 1915: 487. Type-species: Glamyromyrmex beebi Wheeler, 1915: 488, fig. 2, by
monotypy.
Borgmeierita Brown, 1953a: 23. Type-species: Codiomyrmex excisus Weber, 1934: 51, fig. 9, by original
designation. [Synonymy by Brown, 1973a: 35.]
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles relatively short (MI 8-24), subtriangular in
full-face view and powerfully developed, serially dentate and lacking an apical fork of spiniform teeth.
When fully closed the mandibles are overlapped basally by the clypeus. In profile the mandibles with their
upper and lower margins diverging from base to apex and the distal portion passing into a strong
downcurved arc so that part or most of the apical margin is at right-angles to the long axis of the head.
Apical (masticatory) margin of mandible with 8-11 teeth following a conspicuously differentiated
prominent basal lamella; the basal series of teeth following the lamella large and strong, the lamella itself
partially or wholly concealed by the clypeus when the mandibles are closed. Antennae with 6 segments.
THE AFROTROPICAL DACETINE ANTS 321
Two species from the Afrotropical region which were formerly included in Codiomyrmex are
here transferred to Glamyromyrmex. Brown (1973£) has indicated that the former name may be
a junior synonym of the latter. Whether this proves to be correct or not, the previously described
Afrotropical species are certainly closer to the type-species of Glamyromyrmex than they are to
the type-species of Codiomyrmex (C. thaxteri Wheeler). At present there are 11 Afrotropical
and 7 Neotropical species of Glamyromyrmex known, though it is most likely that some or all of
the species currently placed in Codiomyrmex and Chelystruma, from Australia and the
Neotropical region, may be referred to Glamyromyrmex in the future. Glamyromyrmex and its
relatives are closely related to Smithistruma, differing in the much more powerful construction
of the mandibles in the former. The species currently placed in Glamyromyrmex are a fairly
diverse assemblage and may even represent several separate lines of descent from Smithistruma-
like ancestral forms. As mentioned under Smithistruma the stability of the short-mandibulate
dacetine genera is in question and a world revision of them would most probably show some
marked changes in generic limits.
Previous work on Glamyromyrmex has mostly been limited to the description of new species
but the genus has been defined by Brown (19500) on the basis of the sparse material then
available, and the Neotropical species have been keyed by Kempf (1960).
List of Afrotropical Glamyromyrmex
tetragnathus-group loveridgei (Brown) comb. n.
africanus sp. n. sistrurus sp. n.
tetragnathus (Taylor) comb. n. ravidurus-group
dagon-group ravidurus sp. n.
dagon sp. n. thuvidus-group
sahurussp.n. thuvidussp.n.
loveridgei-group trymalussp.n.
crypturussp.n. tukultussp. n.
Key to species (workers)
1 Clypeal dorsum without appressed small hairs of any description , all hairs present on the clypeal
dorsum conspicuously elevated. (Cameroun) trymalus (p. 333)
- Clypeal dorsum with appressed small hairs which may be simple , spatulate or scale-like 2
2 Appressed hairs on clypeus simple , fine and minute 3
- Appressed hairs on clypeus flattened, spatulate or scale-like 6
3 Posteromedian area of cephalic dorsum raised into a broad tumulus which is bounded laterally
by an approximately flat area on each side (Fig. 27). Pronotum laterally sharply marginate
throughout, the margins overhanging the sides (Fig. 30). Postpetiolar disc finely longitudinal-
ly costulate 4
- Posteromedian area of cephalic dorsum not raised into a broad tumulus, instead the dorsum is
more or less evenly convex from side to side. Pronotum laterally not sharply marginate
throughout. Postpetiolar disc smooth or at most with lateral shagreening 5
4 Lateral margins of head in full-face view with projecting flagellate hairs. Postpetiole and first
gastral tergite with numerous long flagellate hairs. (Gabon) africanus (p. 322)
- Lateral margins of head in full-face view without flagellate hairs. Postpetiole and first gastral
tergite with sparse short straight hairs. (Cameroun, Angola) tetragnathus (p. 323)
5 With the head in profile the edges of the clypeal lobes enormously thickened , much thicker than
the maximum width of the scape (Fig. 32) . Anterior clypeal margin with a semicircular median
impression flanked by a lobe on each side (Fig. 25). CI 70-71, MI 8-9. (Rwanda) dagon (p. 325)
- With the head in profile the edges of the clypeal lobes narrow, narrower than the maximum
width of the scape (Fig. 33). Anterior clypeal margin broadly evenly shallowly concave from
corner to corner (Fig. 26). CI 75-78, MI 16-17. (Rwanda) sahurus (p. 326)
6 Entire cephalic dorsum densely clothed with appressed scale-like hairs, without other pilosity of
any description and the sides of the head without projecting flagellate hairs
Cephalic dorsum not clothed with appressed scale-like hairs, with other pilosity present, and the
sides of the head with projecting flagellate hairs 8
7 First gastral tergite with erect fine simple hairs present. (Ivory Coast) tukultus (p. 334)
- First gastral tergite with appressed spatulate to scale-like hairs only. (Cameroun) . . ravidurus (p. 331)
322 BARRY BOLTON
8 Cephalic dorsum behind clypeus with widely spaced broad foveolate punctures, with a cratered
appearance, the surface smooth between the punctures. Minute yellow species, HL 0-43-
0-44, HW 0-29. (Kenya) thuvidus (p. 332)
Cephalic dorsum behind clypeus reticulate-punctate, without widely spaced broad foveolate
punctures, without a cratered appearance. Larger black or blackish brown species, HL 0-49 or
more, HW 0-34 or more 9
9 With postpetiole in dorsal view the sides of the disc completely enclosed by dense spongiform
tissue. Dorsum of petiole node sculptured, at least in part. Basal lamellate band of first gastral
tergite broad and continuous , indented anteromedially but not interrupted 10
- With postpetiole in dorsal view the spongiform tissue restricted to the posterolateral angles of
the disc and fading out anteriorly. Dorsum of petiole node unsculptured and smooth. Basal
lamellate band of first gastral tergite narrow at sides and interrupted medially. (Malawi)
loveridgei (p. 328)
10 Mandible with 6 enlarged teeth, without medially projecting broad spatulate hairs between the
basalmost tooth and the basal lamella (Fig. 24). (Cameroun) sistrurus (p. 329)
- Mandible with 5 enlarged teeth, with medially projecting broad spatulate hairs between the
basalmost tooth and the basal lamella (Fig. 23). (Ghana) crypturus (p. 327)
The tetragnathus-group
(Figs 27, 30)
Outline shape of head as Fig. 27. Mandibles with 8 teeth consisting of 5 large slightly recurved spiniform
teeth following the basal lamella, and an apical series of 3 denticles which share a common base. Anterior
clypeal margin indented medially. Lateral clypeal margin not expanded into a broad lobe on either side,
the outer margins of the mandibles at full closure intersecting the anterior clypeal margin at or very close to
the anterolateral corners. Head dorsoventrally flattened, the dorsum posteromedially raised into a broad
tumulus. Broad rounded occipital lobes present which are strongly prominent posteriorly. Pronotum
sharply marginate laterally, the marginations overhanging the sides and the dorsum between the
marginations shallowly transversely concave. Sculpture of cephalic dorsum behind clypeus weak and
superficial, without well-developed rugulose or punctate sculpture. Clypeus with minute appressed
pubescence.
The two species placed in this group, africanus and tetragnathus , are closely related forms, very
conspicuous in appearance and easily distinguished from all the other Afrotropical members of
Glamyromyrmex. They are the members of this genus which least resemble Smithistruma and
the construction of the head renders them immediately recognizable.
Both species are of central African origin, with africanus known from Gabon and tetragnathus
from Cameroun and Angola.
Glamyromyrmex a fricanussp. n.
(Fig. 27)
HOLOTYPE WORKER. TL 3-0, HL 0-82, HW 0-70, CI 85, ML 0-16, MI 20, SL 0-30, SI 43, PW 0-36, AL 0-68.
Mandibles with a broad basal lamella (partially visible as mandibles slightly opened) followed by a
principal dental row of 5 large triangular teeth which are slightly recurved and evenly spaced on the
strongly downcurving arch of the mandibular masticatory margin. Basalmost tooth following the lamella
without a diastema, slightly smaller than the second tooth. Third tooth from the basal lamella the largest,
fourth and fifth tooth slightly smaller. Distal to the principal dental row is a series of three denticles which
share a common base. In profile the mandibles rapidly increasing in width from base to apex, the upper and
lower borders strongly divergent, the former arching up above the level of the anterior clypeal margin, the
latter shallowly concave and downcurved; apical margin as seen in profile strongly arched-convex. With the
head in full-face view the anterior clypeal margin with a concave median indentation, the clypeus broad but
the lateral free margins not extended into lobes and noticeably narrower than the sides of the head behind
the clypeus. Sides of head evenly shallowly convex, broader behind than in front. Occipital margin
extended backwards as a rounded lobe on each side of a central broad impression, the median portion of
which is transverse . The margin of the occipital impression bounded on the dorsum by a continuous low rim
or crest. Dorsum of head behind clypeus raised medially into a broad tumulus which is convex in both
directions but does not reach the lateral margins; rather the tumulus is surrounded on all sides by more or
less flat areas of cuticle. Frontal lobes and frontal carinae fused to form a continuous upper scrobe margin
THE AFROTROPICAL DACETINE ANTS 323
which is strongly prominent laterally, conceals the scrobes from dorsal view, and is continuous with the
flattened posterolateral portions of the head. Antennal scrobes deep and extensive, anteriorly divided into
upper and lower compartments by the weak preocular laminae and with the small eye situated on the
ventral scrobe margin. In profile the dorsal cephalic tumulus is balanced by an even more strongly
prominent mid-ventral tumulus whose maximum convexity occurs at about the level of the eye and behind
which the ventral surface is markedly concave. Antennal scapes short (SI 43) and feebly clavate, not bent
near the base, the leading edge evenly shallowly convex and lacking projecting hairs. Dorsum of head with
dense abundant decumbent to appressed fine simple hairs, shortest near the clypeus (which has only
minute fine pubescence) and longest occipitally, directed anteriorly or anteromedially and densest on the
lateral margins; the latter also with 3 pairs of long projecting flagellate hairs. Dorsum of head with minute
shallow pits from which the hairs arise, otherwise unsculptured except for a feeble superficial shagreening
in places; clypeus smooth. Sides of alitrunk sharply laterally marginate throughout, most strongly so on the
pronotum where the marginations are prominent and overhang the sides. Pronotum also marginate
anteriorly and with a median longitudinal ridge or carina dorsally. Dorsum of mesonotum separated from
the short propodeal dorsum by a low transverse crest; metanotal groove absent. Propodeal teeth very
broad basally, rapidly tapering apically and with the extreme apices upcurved. Infradental lamellae much
narrower than the propodeal teeth , the latter with more than half their length standing free of the lamellae .
Sides of alitrunk unsculptured except for the punctate mesopleuron. Dorsal alitrunk with a few superficial
rugular vestiges on the promesonotum but only the median carina conspicuous. Lateral margination of the
alitrunk with 2-3 long flagellate hairs on each side, otherwise the dorsum and margins only with fine
scattered simple pilosity. Peduncle of petiole long, the node bluntly triangular in profile. Spongiform
ventral process of petiole peduncle massively developed and curtain-like, about as deep as the node is high.
Other spongiform material on petiole reduced to a pair of short aliform prominences situated lateroven-
trally when the node is viewed from above and from which a narrow crest arises which follows the posterior
margin of the node. Petiole node sparsely rugulose, disc of postpetiole sparsely irregularly longitudinally
costulate. Ventral spongiform appendages of postpetiole moderate, the lateral appendages narrow in
dorsal view, broadest at the posterolateral angles. Anterior face of postpetiolar disc with a narrow
bordering lamella, the posterior margin without spongiform material, bordered instead only by a sharp
narrow and shallowly convex rim which abuts a similar but concave rim bordering the base of the first
gastral tergite. First gastral tergite without basal spongiform material but with a lamellate area laterobasal-
ly, immediately behind the lateral appendages of the postpetiole; this lamellate area thrown into strong
ridges which form the origins of the lateral basigastral costulae. First gastral sternite without a basal
spongiform pad. Primary basigastral costulae, arising at the base of the tergite, few in number and mostly
lateral in origin; more posteriorly numerous finer costulae arise which form a dense band over about
one-third of the length of the tergite. Petiole, postpetiole and first gastral tergite with numerous long fine
flagellate hairs. Black, the spongiform appendages pale.
Holotype worker (gold-palladium coated), Gabon: Makokou, berlese no. 17, x-xii. 1972, rain forest (/.
Lieberburg) (MCZ).
The only known close relative of africanus is tetragnathus , from Cameroun and Angola.
Differences to separate these two species are tabulated under tetragnathus.
Glamyromyrmex tetragnathus (Taylor) comb. n.
(Fig. 30)
Codiomyrmex tetragnathus Taylor, 1965: 225, figs 1, 2. Holotype worker, ANGOLA: Dundo, Route
Turismo, approx 7°02'S, 20°51'E, gallery forest, 28.iii.1962, no. 16888, R. Luachimo, 'berlesate by
native collector' (MCZ) [examined].
WORKER. TL 2-4-2-8, HL 0-60-0-70, HW 0-49-0-58, CI 79-83, ML 0-14-0-16, MI 22-24, SL 0-25-0-28, SI
47-51, PW 0-28-0-34, AL 0-58-0-62 (4 measured).
Mandibles with a broad basal lamella followed by a row of 5 large triangular teeth which are slightly
recurved and evenly spaced on the strongly downcurving arch of the apical margin. Distal to this tooth row
is a series of 3 denticles which share a common base on a low process. In profile the mandibles rapidly
increasing in width from base to apex, the upper and lower borders strongly divergent, the former arching
up above the level of the anterior clypeal margin, the latter shallowly concave and downcurved; apical
margin as seen in profile strongly arched-convex. With the head in full-face view the anterior clypeal
margin with a median indentation, the clypeus broad but the lateral free margins not extended into lobes
and noticeably narrower than the sides of the head behind the clypeus. General shape of head in profile and
324
BARRY BOLTON
full-face views as described for africanus. Antennal scapes short (SI 47-51) and feebly clavate, not bent
near the base, with the leading edge more or less evenly convex and lacking projecting hairs. Clypeal
dorsum with minute appressed pubescence which is directed anteriorly. Dorsum of head with anteriorly or
anteromedially directed minute appressed hairs which are approximately the same length everywhere on
the dorsum and no longer than the clypeal pubescence. Lateral margins of head without flagellate hairs.
Clypeus smooth. Dorsum of head with minute scattered pits from which the hairs arise, otherwise
unsculptured except for a feeble superficial shagreening in places. Sides of ali trunk sharply laterally
marginate throughout their length, most strongly so on the pronotum where the marginations are
prominent and overhang the sides. Pronotum also marginate anteriorly and with a median longitudinal
ridge or carina dorsally. Dorsum of mesonotum separated from the short propodeal dorsum by a low
transverse crest; metanotal groove absent. Propodeal teeth very broad basally, rapidly tapering apically
and with the extreme apices suddenly upcurved. Infradental lamellae much narrower than the propodeal
teeth, the latter with more than half their length standing free of the lamellae. Sides of alitrunk
unsculptured except for the punctate mesopleuron and some weak peripheral sculpture round the margins
of the segments. Dorsal alitrunk with some superficial rugular vestiges on the pronotum and mesonotum
beside the median carina. Lateral margins of alitrunk with two pairs of long simple hairs, the first pair at the
pronotal humeri, the second pair just in front of the transverse crest that separates mesonotum from
propodeum. Dorsal alitrunk otherwise with only sparse appressed simple hairs which are very short.
Spongiform ventral appendage of petiole peduncle massively developed and curtain-like, about as deep as
the node is high. Other spongiform material on petiole reduced to short aliform prominences situated
lateroventrally when the node is viewed from above. Petiole node sparsely rugulose, disc of postpetiole
finely densely and quite regularly longitudinally costulate. Lateral spongiform appendages of petiole
narrow in dorsal view, broadest posterolaterally. Anterior face of the postpetiolar disc with a narrow
bordering lamella, the posterior margin bordered by a convex rim which abuts a similar but concave rim
bordering the base of the first gastral tergite. First gastral tergite lamellate basally, not spongiform, with
numerous strong basigastral costulae and with many secondary costulae arising between them on the body
of the tergite. First gastral sternite without a basal spongiform pad. Petiole, postpetiole and first gastral
tergite with scattered short straight simple hairs. Colour blackish brown to black.
The only known relative of tetragnathus is africanus; the two are separated as follows in the
worker.
africanus
Larger species with shorter antennal
scapes, HW 0-70, SI 43.
Lateral margins of head in full-face
view with 3 pairs of projecting
flagellate hairs.
Appressed cephalic pilosity longer
posteriorly than anteriorly; much
longer than the clypeal pubescence.
Postpetiole and first gastral tergite
with numerous long fine flagellate
hairs.
tetragnathus
Smaller species with longer antennal
scapes, HW 0-49-0-58, SI 47-51.
Lateral margins of head in full-face
view without flagellate hairs.
Appressed cephalic pilosity very short,
of approximately equal length
everywhere; no longer than the
clypeal pubescence.
Postpetiole and first gastral tergite
with simple sparse short straight
hairs.
MATERIAL EXAMINED
Cameroun: Nkoemvon (D. Jackson); nr Yaounde (G. Terrori). Angola: Dundo.
The dagwi-group
(Figs 25, 26, 32, 33)
Outline shape of head as Figs 25, 26. Mandibles with 8 teeth consisting of a small denticle close to the basal
lamella followed by a series of 5 large teeth and an apical pair of denticles which arise from a common base.
Anterior clypeal margin broadly and evenly concave or sharply indented medially. Lateral clypeal margins
expanded into a lobe on each side, the outer margins of the mandibles at full closure intersecting the
anterior clypeal margin some distance medially of the anterolateral corners. Head not dorsoventrally
flattened, without a broad convex tumulus posteromedially on the dorsum. Posteriorly projecting rounded
occipital lobes absent. Pronotum not sharply marginate laterally, the dorsum transversely flat to shallowly
THE AFROTROPICAL DACETINE ANTS 325
convex. Cephalic sculpture behind clypeus finely and very densely reticulate-punctate, with a granular
appearance. Clypeus with minute appressed pubescence.
The two species in this group, dagon and sahurus, are both known only from Rwanda. They are
characterized chiefly by their dentition and their possession of a broad clypeus whose free lateral
margins are convex and expanded into a lobe on each side. The character is much more strongly
expressed in dagon than in sahurus and is accompanied in the former by a massive thickening of
the lateral free margins of the clypeus.
Glamyromyrmex dagon sp. n.
(Figs 25, 32)
HOLOTYPE WORKER. TL 2-0, HL 0-57, HW 0-40, CI 70, ML 0-05, MI 9, SL 0-24, SI 60, PW 0-26, AL 0-52.
Mandibles appearing very short in full-face view (MI 8-9) as the apical (masticatory) margin is at a
right-angle to the long axis of the head from the second tooth to the apex. Basal lamella of mandible, which
is concealed by the clypeus at full closure, followed by a short diastema and a denticle. Distal to the denticle
is a row of 5 large teeth which are slightly recurved, and apically two denticles which share a common base
are present. In profile the upper mandibular margin curves upwards above the level of the anterior clypeal
margin and the apical (masticatory) margin forms a near-vertical shallowly convex arch. Anterior clypeal
margin in full-face view convex on each side of a deep median concavity, the lateral free margins of the
clypeus expanded into a smoothly rounded prominent broad lobe on each side so that the outer margins of
the closed mandibles intersect the anterior clypeal margin some distance medially of the anterolateral
corners. Upper scrobe margins shallowly concave and feebly divergent posteriorly, the lateral margins of
the occipital lobes behind this are shallowly convex and round behind into the smoothly concave occipital
margin. In profile the edges of the laterally expanded clypeal lobes are greatly thickened in front of the
level of the antennal insertions, the maximum thickness distinctly greater than the maximum width of the
scape. Eyes present, small, situated on the ventral margin of the deep scrobe. Scapes of moderate length
(SI 60), narrowly clavate and lacking projecting hairs on the leading edges. Dorsum of head with a single
pair of erect fine hairs situated on each side of the midline close to the occipital margin. Otherwise the head
only with very short fine simple hairs everywhere which are appressed and directed anteriorly; flagellate
long hairs absent. Clypeus with scattered minute appressed pubescence only. Dorsum of clypeus, and of
head in a band immediately behind the clypeus, glassy smooth. Remainder of cephalic dorsum finely and
densely reticulate-punctulate. Anterior border of pronotum narrowly marginate, the sides of the pro-
notum immarginate anteriorly but separated from the dorsum by a blunt angle posteriorly. In profile the
promesonotum dorsally forming a single convex outline which is separated from the propodeal dorsum by a
small step, which appears as a transverse rim in dorsal view, the propodeum being on a slightly lower level
than the mesonotum and marginate laterally. Infradental lamellae of propodeum broad, the propodeal
teeth represented only by a short narrow denticle standing free of the lamella. Sides of alitrunk glassy
smooth except for peripheral punctate sculpture. Pronotal dorsum unsculptured, mesonotum densely
punctate, propodeum smooth anteriorly but with some punctures between the bases of the teeth.
Pronotum and mesonotum each with a single pair of long erect simple hairs, the dorsum otherwise only
having scattered short appressed hairs which are directed roughly towards the midline. Spongiform
appendages of pedicel segments massively developed in profile. Petiole node in dorsal view unsculptured,
broader than long, flanked on each side by a prominent spongiform process, the two linked across the
posterior margin of the node by a narrow lamella. Disc of postpetiole glassy smooth, very broad and
surrounded by spongiform or lamellate tissue on all sides. Spongiform tissue broadest posterolaterally,
narrowest medially where it is contracted down to a very narrow rim along the posterior margin of the disc.
Base of first gastral tergite with a spongiform strip which is thickest laterally. Basigastral costulae sparse in
centre of tergite, denser laterally. Dorsal surfaces of petiole, postpetiole and gaster with sparse erect
simple pilosity and also with much shorter appressed widely scattered simple hairs. Colour yellowish
brown.
PARATYPE WORKER. TL2-1, HLO-59, HWO-42, CI 71, ML 0-05, MIS, SLO-25, SI 60, PWO-28, ALO-56. As
holotype.
Holotype worker, Rwanda: Rangiro, i.1976, forest humus, 1800 m (P. Werner) (MHN).
Paratype. 1 worker with same data as holotype (BMNH).
G. dagon is immediately recognizable as no other African species has such short mandibles or
such a bizarre clypeal structure. Its only close relative is sahurus and characters separating the
two are tabulated under the latter name.
326 BARRY BOLTON
Glamyromyrmex sahurus sp. n.
(Figs 26, 33)
HOLOTYPE WORKER. TL 2-3, HL 0-56, HW 0-42, CI 75, ML 0-09, MI 16, SL 0-25, SI 57, PW 0-28, AL 0-58.
Mandibles in full-face view with the basal lamella mostly concealed by the clypeus at full closure but its
margin continued as an oblique edge leading to the first tooth, which is thus some distance from the clypeal
margin. Basalmost tooth small, reduced to a denticle. Distal to this is a row of 5 large teeth which are
slightly recurved, and apically two denticles which share a common base are present. In profile the upper
and lower mandibular margins are strongly divergent from base to apex, the dorsal margin curving upwards
until it is above the level of the anterior clypeal margin and the apical (masticatory) margin forming a
shallowly convex arch which is approximately at a right-angle to the long axis of the head. Anterior clypeal
margin shallowly concave in full-face view, the lateral free clypeal margins expanded into a broad but
shallow lobe on each side. Upper scrobe margins divergent posteriorly, straight to very shallowly concave.
Posteriorly the upper scrobe margins merge with the weakly convex lateral occipital lobes which round
posteriorly into the broadly shallowly concave occipital border. In profile the lateral free margins of the
clypeal lobes not grossly thickened, narrower than the maximum width of the scape. Eyes present, small,
situated on the ventral margin of the deep scrobe. Scapes of moderate length (SI 57-60), narrowly clavate
and lacking projecting hairs on the leading edges. Dorsum of head with a pair of erect fine hairs situated
close to the midline and close to the occipital margin. Head otherwise devoid of elongate hairs; lacking
flagellate hairs but fairly densely clothed with curved to hooked anteriorly directed short simple hairs.
Clypeus with minute appressed pubescence only. Clypeus shiny with a superficial punctulate patterning;
dorsum of head behind clypeus finely but very densely and conspicuously reticulate-punctate and dull.
Anterior pronotal border weakly marginate, the sides not marginate and rounded anteriorly but separated
from the dorsum by a weak blunt angle posteriorly. Mesonotum not, and propodeum only weakly bluntly
marginate laterally. Mesonotum separated from propodeum on dorsum by an extremely feeble ridge or
crest. In profile the mesonotum slightly raised but not separated from the propodeum by a groove.
Propodeal teeth vestigial to absent, not or only weakly differentiated from the infradental lamellae as
minute points. Sides of alitrunk glassy smooth, without sculpture except on the extreme periphery.
Pronotal dorsum unsculptured and shining; mesonotum with a smooth median longitudinal strip but
punctate on each side of it; propodeum smooth anteriorly but punctate at the top of the declivity.
Pronotum and mesonotum each with a single pair of long fine subflagellate hairs, otherwise only short
simple appressed hairs which are directed towards the midline are present. Spongiform appendages of
pedicel segments massively developed in profile. Petiole node in dorsal view unsculptured, broader than
long, equipped with a strongly prominent spongiform process on each side and the two linked by a narrow
lamella which runs across the posterodorsal margin of the node. Postpetiolar disc with a smooth broad
median longitudinal strip but with faint shagreening on each side, the postpetiole surrounded by
spongiform or lamellate material. The shallowly convex anterior margin of the postpetiole is bordered by a
narrow lamella which is confluent with the lateral spongiform tissue on each side. The latter is thickest
posterolaterally but narrows down to a vestigial strip posteromedially where the posterior postpetiolar
margin is most strongly convex. Base of first gastral tergite with a spongiform transverse trip which narrows
medially behind the posteriormost point of the postpetiole. First gastral tergite with conspicuous dense
basigastral costulae grouped on each side of a median smooth area. Dorsal surfaces of petiole, postpetiole
and first gastral tergite with scattered erect fine hairs and with scattered short simple appressed hairs.
Colour brown.
PARATYPE WORKERS. TL 2-1-2-3, HL 0-52-0-56, HW 0-40-0-42, CI 75-78, ML 0-09, MI 16-17, SL
0-24-0-25, SI 57-60, PW 0-26-0-28, AL 0-54-0-58 (3 measured).
As holotype but the mesonotal sculpture may consist of punctures everywhere, obliterating the median
clear area seen in the holotype.
Holotype worker, Rwanda: Rangiro, ix.1976 (P. Werner) (MHN).
Paratypes. 3 workers with same data as holotype (MHN; BMNH; MCZ).
Along with dagon, sahurus forms a close species-pair known only from Rwanda and characte-
rized by the form of the mandibles and clypeus, though the modification of the latter is much
more extreme in dagon than in sahurus (Figs 25, 26). The two species are separated as follows in
the worker.
dagon sahurus
CI 70-71, MI 8-9 CI 75-78, MI 16-17.
THE AFROTROPICAL DACETINE ANTS 327
dagon - cont. sahurus - cont.
Anterior clypeal margin with a deep Anterior clypeal margin shallowly
median impression flanked on each concave.
side by a convex lobe.
Lateral free margins of clypeal lobes Lateral free margins of clypeal lobes
greatly thickened, thicker in not thickened, in profile much
profile than the maximum width of narrower than the maximum width of
the scape. the scape.
Lateral portions of postpetiolar Lateral portions of postpetiolar disc
disc smooth. finely sculptured.
The /overidgei-group
(Figs 23, 24)
Outline shape of head as Figs 23, 24. Mandibles with 7-8 teeth. Either with 6 teeth plus a pair of apical
denticles which share a common base (sistrurus), or with 5 teeth plus an apical series of 3 denticles which
share a common base (loveridgei), or with 5 teeth plus a minute apical pair of denticles (crypturus).
Anterior clypeal margin shallowly convex to extremely feebly evenly concave, not indented medially nor
deeply concave. Lateral clypeal margins not expanded into lobes on each side, the outer margins of the
mandibles at full closure intersecting the anterior clypeal margin at the anterolateral corners. Head not
dorsoventrally flattened and without a posteromedian broad tumulus dorsally. Posteriorly projecting
rounded occipital lobes absent. Pronotum not sharply marginate laterally, the dorsum transversely convex
to approximately flat. Sculpture of head behind clypeus strongly reticulate-punctate. Clypeus with
scale-like to spatulate appressed hairs. Flagellate hairs present on lateral margins of head.
G. loveridgei from Malawi, and the westerly distributed species sistrurus from Cameroun and
crypturus from Ghana, have the head more strongly sculptured than in other Afrotropical
representatives of the genus. Basically this sculpture is a strong dense reticulate-punctation but a
tendency to rugulation is present due to the alignment of adjacent punctures, whose walls form
rugule-like ridges.
Glamyromyrmexcrypturussp. n.
(Fig. 23)
HOLOTYPE WORKER. TL 2-0, HL 0-53, HW 0-37, CI 70, ML 0-10, MI 19, SL 0-21, SI 57, PW 0-24, AL 0-54.
Mandibles with a principal row of 5 enlarged teeth. Basally the mandible with a broad and extensive
lamella which has in the small diastema between itself and the basalmost tooth a medially directed long
broadly spatulate hair. An even longer but not so broadly spatulate hair projects medially from a point
closer to the clypeus but more remote from the masticatory margin than the broadly spatulate hair. If the
mandibles are fully closed (ajar in the holotype) these hairs may be difficult to see. Basalmost tooth smaller
than the more strongly recurved second tooth, the second smaller than the third and fourth, the fifth
slightly smaller than the fourth. Distal to the fifth tooth detail is difficult to see but there appears to be a pair
of vestigial denticles. In profile the upper and lower mandibular margins are strongly divergent from base
to apex and the upper margin curves up above the level of the anterior clypeal margin. The apical margin,
from the second tooth to the apex, is almost at right-angles to the long axis of the head. Anterior clypeal
margin transverse, the lateral free margins diverging anteriorly from the frontal lobes and with a few
projecting spatulate to spoon-shaped hairs. Maximum width of clypeus greater than the width across the
frontal lobes. Upper scrobe margins diverging posteriorly, the sides of the head convex and the occipital
margin concave. Eyes small, situated on the ventral scrobe margin. Antennal scapes weakly clavate, the
leading edges shallowly convex and equipped with apically curved spatulate hairs. Clypeus densely clothed
with appressed small scale-like hairs which are distinctly snorter than the basalmost tooth. Hairs on dorsum
of head immediately behind the clypeus short spatulate and appressed, but moving towards the occiput the
hairs become more elevated and more narrowly spatulate, anteriorly arched or curved. At the highest
point of the vertex the hairs are narrowly spatulate but further back they become simple, though still
curved anteriorly. Clypeus smooth and shining, remainder of cephalic dorsum broadly and strongly
reticulate-punctate. Three pairs of long flagellate hairs present; one pair on the dorsum close to the
occipital margin, one pair at the occipital corners and one pair at the apices of the scrobes. Anterior border
of pronotum marginate , the sides not marginate . Sides of mesonotum and propodeum marginate , the latter
more strongly so than the former, the two not separated by a ridge or crest across the dorsum.
328 BARRY BOLTON
Promesonotal dorsum in profile forming a more or less even shallow convexity which is on a slightly higher
level than the propodeum. Metanotal groove absent. Propodeal teeth incorporated in the infradental
lamellae. Sides of alitrunk smooth except for a few longitudinal rugulae anteriorly on the pronotum.
Dorsal alitrunk smooth except for some weak sculptural vestiges on the mesonotum which, however, is
mostly smooth. Pronotum and mesonotum each with a single pair of long flagellate hairs, otherwise the
dorsum only with scattered fine simple hairs which are arched towards the midline. Spongiform appen-
dages massively developed in profile. Petiole node in dorsal view roughly transversely rectangular, slightly
broader than long and fractionally broader behind than in front, the anterolateral angles approximately
right-angular and the surface with scattered weak punctures. Spongiform material strongly prominent
posterolaterally, linked across the posterior margin of the node by a broad translucent lamella. Postpetio-
lar disc smooth and shining, surrounded on all sides by dense Spongiform tissue. Posteromedially the
spongiform tissue narrowing down to a slender lamella. Base of first gastral tergite with a broad transverse
ridged lamellate strip which is shallowly concave anteromedially but not interrupted. Basigastral costulae
short, arranged in two groups, on each side of a clear median strip; the costulae shortest near this clear area
and longer laterally. Petiole dorsum with simple long fine hairs which are curved posteriorly. Postpetiole
with simple fine hairs which are mostly erect or suberect and hooked or curved apically. Gaster with
scattered simple erect hairs. Colour blackish brown.
Holotype worker, Ghana: Tafo, 29.xi.1969, ant ecology sample AES 433 (D. Lestori) (BMNH).
Closest related to sistrurus, characters separating sistrurus and crypturus are tabulated under
the former name.
Glamyromyrmex loveridgei (Brown) comb. n.
Codiomyrmex loveridgei Brown, 1953a: 21. Holotype worker, MALAWI: N. Prov., Nyika Plateau, above
Nchenachena, 5000 ft (1524 m), 1948 (A. Loveridge) (MCZ) [examined].
WORKER. TL 2-2, HL 0-54, HW 0-39, CI 72, ML 0-11, MI 20, SL 0-22, SI 56, PW 0-27, AL 0-54.
Mandibles with a broad basal lamella which is followed without a diastema by 5 enlarged teeth and an
apical series of 3 denticles which share a common base. Upper and lower mandibular margins in profile
strongly divergent from base to apex, the apical (masticatory) margin a strongly downcurved arch but the
dorsal mandibular border not upcurved beyond the level of the anterior clypeal margin. In full-face view
the anterior clypeal margin shallowly convex; posteriorly the clypeus narrowing to the frontal lobes and the
preocular laminae visible. Upper scrobe margins divergent posteriorly from the small frontal lobes, not
strongly expanded laterally. Behind the level of the scrobes the sides shallowly convex and rounding into
the more or less transverse occipital border. Eyes small, situated on the ventral scrobe margin. Antennal
scapes slightly curved and feebly clavate, the leading edge lacking long projecting hairs but with numerous
fine apically directed simple decumbent hairs. Clypeal dorsum densely clothed with elongate flattened
scale-like hairs which are directed forwards and are closely appressed to the surface. Remainder of cephalic
dorsum with decumbent fine dense anteriorly arched simple hairs. Close behind the clypeal posterior
margin the hairs are intermediate in shape between the scale-like clypeal pilosity and the simple cephalic
hairs, being narrowly spatulate or feebly clavate apically. A laterally projecting long flagellate hair present
at the apex of the antennal scrobe on each side and another at each occipital corner. Clypeus shiny and very
smooth, the remainder of the head densely reticulate-punctate. Anterior border of pronotum marginate,
the pronotal sides bluntly marginate posteriorly but the sides broadly rounding into the dorsum anteriorly.
Mesonotum not and propodeum only weakly laterally marginate. On the dorsum the mesonotum and
propodeum separated only by a change of sculpture, without a transverse ridge or crest. In profile the
metanotal groove absent, the propodeal teeth triangular and confluent with the conspicuous infradental
lamellae. Sides of alitrunk unsculptured except for a few strong punctures anteriorly on the pronotum and
feeble peripheral sculpture dorsally and posteriorly. Dorsal surfaces of pronotum and propodeum smooth
and shiny, the mesonotum densely punctate. A pair of long fine subflagellate hairs present at the pronotal
humeri, and another pair on the mesonotum; otherwise the dorsal alitrunk only with scattered fine hairs
which are decumbent to appressed and directed approximately towards the midline. Spongiform appen-
dages of pedicel segments conspicuously developed in profile. In dorsal view the petiole node much
broader than long, with a pair of small lateral spongiform lobes connected across the posterior margin of
the node by a vestigial lamellar strip. Postpetiole in dorsal view with a narrow lamella bordering the
anterior margin, the lateral spongiform tissue broad behind but fading out anteriorly; the posterior margin
only with a narrow transverse lamellar strip joining the two posterolateral spongiform masses. First gastral
tergite basally with a narrow transverse strip of spongy lamellar tissue which is concave and interrupted
medially. Dorsal surfaces of petiole and postpetiole smooth; first gastral tergite with basal costulae dense
THE AFROTROPICAL DACETINE ANTS
329
on each side of a median strip where they are short and sparse. Simple fine standing hairs present on dorsal
surfaces of petiole, postpetiole and first gastral tergite together with shorter sparse decumbent to appressed
simple hairs. Colour brown.
Known only from the holotype worker, the Malawian loveridgei is closest related to the West
African sistmrus and crypturus. In the worker they are quickly separated by the following
characters.
loveridgei
Mandibles with 3 apical denticles
sharing a common base.
Mesonotum densely punctate.
Petiole node dorsally much broader
than long, the surface smooth.
Disc of postpetiole in dorsal view
with thick spongiform material
posterolaterally, fading out to
nothing anteriorly.
Basal transverse lamellate strip of
first gastral tergite narrow and
interrupted medially.
sistrurus and crypturus
Mandibles with 2 apical denticles
sharing a common base.
Mesonotum mostly or wholly smooth.
Petiole node dorsally only marginally
broader than long, the surface
sculptured.
Disc of postpetiole in dorsal view with
thick spongiform material visible all
along the sides.
Basal transverse lamellate strip of
first gastral tergite broad, shallowly
concave medially but not interrupted.
MATERIAL EXAMINED
Malawi: Nyika Plateau, above Nchenachena (A Loveridge).
Glamyromyrmex sistrurus sp. n.
(Fig. 24)
HOLOTYPE WORKER. TL 2-0, HL 0-50, HW 0-35, CI 70, ML 0-10, MI 20, SL 0-20, SI 57, PW 0-23, AL 0-52.
Mandible with a principal dental row of 6 enlarged teeth. Basal tooth of mandible slightly smaller than
the second, following the basal lamella without a diastema. (The basal lamella partially visible as
mandibles are not fully closed.) Second tooth distinctly smaller than the third and the third noticeably
smaller than the fourth and fifth teeth which are the largest. Sixth tooth about the same size as the third,
followed apically by a pair of denticles which share a common base. In profile the upper and lower
mandibular margins divergent from base to apex, the apical (masticatory) margin a strongly downcurved
arch but the mandibular dorsal border not upcurved so that it overlaps the level of the anterior clypeal
margin. Anterior clypeal margin extremely shallowly concave, almost transverse in full-face view. Upper
scrobe margins diverging evenly behind the relatively narrow frontal lobes, not strongly expanded, the
preocular laminae just visible in full-face view. Sides of occipital lobes evenly shallowly convex behind the
level of the scrobes, rounding to the occipital margin which is shallowly concave and lacks strongly
prominent posteriorly projecting lobes. Eyes small, of about 10 ommatidia, situated on the ventral scrobe
margin. Antennal scapes weakly clavate, broadest at about the midlength, with the leading edge shallowly
convex and equipped with a number of very narrowly spatulate hairs which are subdecumbent to
decumbent and directed towards the apex of the scape. Clypeal dorsum densely clothed with short
spatulate appressed hairs. Behind the clypeus the hairs more narrowly spatulate and posteriorly becoming
gradually even narrower so that by the level of the eye the hairs are simple. All cephalic hairs behind the
clypeus are strongly arched forwards and subdecumbent to decumbent. Three pairs of long flagellate hairs
present; one pair dorsally close to the occipital margin, one pair at the occipital corners and one pair at the
apices of the scrobes. Clypeus smooth and highly polished, remainder of cephalic dorsum strongly
reticulate-punctate. Pronotum marginate anteriorly but not laterally. Mesonotum and propodeum margin-
ate laterally, the latter more sharply so than the former, the two areas not separated by a ridge or crest
across the dorsum. In profile the dorsal alitrunk convex in outline, highest at the mesonotum and without
trace of a metanotal groove, the mesonotum and propodeum forming an even shallow convexity.
Propodeum with the teeth incorporated in the infradental lamella. Sides of alitrunk unsculptured except
for anterior part of pronotum and some very weak peripheral vestiges. Pronotum and propodeum smooth
dorsally, the mesonotum mostly smooth but with faint scattered punctures. Pronotum and mesonotum
each with a pair of flagellate hairs. Otherwise dorsal alitrunk only with sparse simple hairs of varying length
which arch across the dorsum or curve towards the midline and are subdecumbent to decumbent.
Spongiform appendages of pedicel segments massively developed in profile. Petiole node rugulose on the
330 BARRY BOLTON
sides and dorsum, only very slightly broader than long in dorsal view and equipped with strongly prominent
lateral spongiform appendages which are linked posteriorly by a broad lamina running across the posterior
face of the node. Postpetiole smooth and shining, completely surrounded by spongiform tissue which is
laminar anteriorly and posteriorly, thickest posterolaterally and narrowest medially. Base of first gastral
tergite with a broad strongly ridged transverse band of lamellar spongiform material which has its anterior
margin shallowly concave behind the posteromedian margin of the postpetiole. First gastral tergite with
short basal costulae. Dorsal surfaces of petiole, postpetiole and first gastral tergite with simple fine hairs,
some of those on the petiole subflagellate and others strongly back-curved. Colour black to blackish
brown.
PARATYPE WORKERS. TL 1-9-2-0, HL 0-49-0-53, HW 0-34-0-36, CI 67-70, ML 0-10-0-11, MI 19-21, SL
0-18-0-21, SI 53-58, PW 0-22-0-25, AL 0-52-0-55 (11 measured).
As the holotype but the basal and second teeth on the mandible may be about the same size.
Holotype worker, Cameroun: Nkoemvon, 2.xi.l980, N49 (D. Jackson) (BMNH).
Paratypes. 11 workers with same data as holotype and 1 worker with same data but 6.x. 1980, N34
(partially dissected to show mandibles) (BMNH; MCZ; MHN; ENSA).
Non-paratypic material examined. Cameroun: nr Yaounde (G. Terron).
The separation of sistrums and its closest relative cryptums from loveridgei is tabulated under
the last name. G. sistrurus differs from crypturus as follows.
sistrurus
Mandible with 6 enlarged teeth, the
basalmost following the lamella
without a diastema.
Basal tooth of mandible not followed
by a long broadly spatulate medially
projecting hair before the basal
lamella.
Lateral margins of clypeus more or less
parallel anterior to the frontal
lobes; clypeus narrower (Fig. 24).
Appressed spatulate hairs on clypeus
relatively large, as long as or
longer than the basalmost tooth
of the mandible.
Arched cephalic hairs at highest
point of vertex simple.
crypturus
Mandible with 5 enlarged teeth, with
a short diastema between the
basalmost tooth and the lamella.
Basal tooth of mandible followed by a
long broadly spatulate medially
projecting hair before the basal
lamella.
Lateral margins of clypeus divergent
anterior to the frontal lobes;
clypeus broader (Fig. 23).
Appressed scale-like hairs on clypeus
minute, conspicuously shorter than
the basalmost tooth of the mandible.
Arched cephalic hairs at highest point
of vertex narrowly spatulate.
The rav/duras-group
(Fig. 28)
Outline shape of head as Fig. 28. Mandibles with 11 teeth, the principal dental row of 6 teeth (counting
from the basalmost) is followed by 2 slightly smaller teeth and an apical series of 3 denticles which share a
common base. Anterior clypeal margin weakly impressed medially. Lateral clypeal margins not expanded
into a convex lobe on each side, the outer margins of the mandibles at full closure intersecting the anterior
clypeal margin at the anterolateral corners. Head not dorsoventrally flattened and without a posterome-
dian broad tumulus. Posteriorly projecting rounded occipital lobes present. Pronotum not marginate
laterally, the dorsum shallowly convex. Cephalic sculpture behind the clypeus reticulate-punctate to
granular, partially concealed by the pilosity. Clypeus with scale-like appressed hairs. Flagellate hairs
absent.
The single species included in this group, ravidurus from Cameroun, combines the general
appearance of the members of the loveridgei-group with a more generalized mandible (11 teeth
as opposed to a maximum of 8 in loveridgei and allies), a more specialized pilosity involving the
loss of flagellate hairs and development of uniform appressed scale-like hairs all over the head,
and the presence of broad rounded posteriorly projecting occipital lobes. The high dental count
is the same as that noted in the thuvidus-group but in the latter the mandibles are by no means as
strikingly arched-downcurved as in ravidurus and the head lacks rounded occipital lobes.
THE AFROTROPICAL DACETINE ANTS 331
Glamyromyrmexravidurussp. n.
(Fig. 28)
HOLOTYPE WORKER. TL 2- 1 , HL 0-54, HW 0-41 , CI 76, ML 0- 1 1 , MI 20, SL 0-22, SI 54, PW 0-24, AL 0-58.
Mandible with 1 1 teeth following the basal lamella without a diastema, the dentition consisting of a basal
series of 6 relatively large teeth followed distally by 2 slightly smaller teeth and an apical series of three
denticles which share a common base (from a paratype with open mandibles). Masticatory margin
obscured in dorsal view by the numerous scale-like hairs arising from the blade which project medially over
the teeth. In profile the mandibles strongly arched-downcurved, the upper margin strongly curved and its
highest point above the level of the anterior clypeal margin. Median portion of clypeal dorsum depressed
and more or less flat, the lateral margins weakly elevated. Anterior clypeal margin very feebly indented
medially, almost transverse, the lateral portions of the anterior margin slightly elevated. In full-face view
the sides of the head behind the level of the antennal insertions evenly divergent to the apices of the scrobes
then passing through an obtuse angle and gradually converging occipitally . Rounded posteriorly projecting
occipital lobes present. Mandibles, clypeus, entire cephalic dorsum and antennal segments all with
numerous conspicuous appressed scale-like hairs, without other pilosity of any description. Lateral
margins of head without flagellate hairs. With head in profile the eyes very small, with only 4-5 ommatidia.
Scale-like hairs present on the sides of the head behind the deep scrobes and on the preocular laminae and
the lateral portion of the clypeus in front of the antennal insertions. Clypeus superficially and very faintly
reticulate, the cephalic dorsum weakly reticulate-granular between the scale-like hairs. Antennal scapes
weakly clavate in full-face view, broadest close to the midlength and their leading edges equipped with a
series of appressed scale-like hairs which lie almost nose-to-tail, the apex of one hair nearly touching the
base of the next. Pronotum marginate anteriorly but not marginate laterally. Mesonotum not marginate
but the short propodeum laterally angulate between sides and dorsum; the dorsum broader than long.
Metanotal groove not impressed but present across the dorsal alitrunk as a narrow transverse line. In
profile the dorsal alitrunk outline more or less evenly convex, highest at about the mesonotal midlength.
Propodeal teeth short, upcurved at the extreme apex, confluent ventrally with the broad infradental
lamellae, the latter with their free margins more or less vertical, not evenly concave. Sides of alitrunk
unsculptured, with scattered scale-like hairs on sides of pronotum. Dorsum of promesonotum with
superficial faint reticular patterning and scattered appressed scale-like hairs which are similar to those on
the head but narrower and in general more widely spaced. Propodeal dorsum smooth and without
scale-like hairs. Flagellate hairs absent from alitrunk, without pilosity of any description other than the
scale-like hairs. Pedicel segments in profile with spongiform appendages enormously developed. Petiole
with ventral appendage forming a deep narrow lobe anteriorly but in its posterior half the spongiform tissue
is much expanded laterally so that it is clearly visible in dorsal view. Ventral spongiform appendage of
postpetiole massive and base of first gastral sternite with a well-developed spongiform pad. Petiole node in
dorsal view approximately as broad as long, with a broad lamellate posterior strip. Disc of postpetiole
surrounded on all sides by lamellar or spongiform material, the disc broadest in front of the midlength, the
sides convergent posteriorly and the posterior margin not indented medially. Base of first gastral tergite
with a lamellate spongiform strip which is traversed by the basigastral costulae, the latter very short and
scarcely extending onto the tergite proper. Petiole, postpetiole and gaster unsculptured, the only pilosity
present on all these surfaces being appressed scale-like hairs. Posterior margins of petiole and postpetiole
with a series of large scale-like hairs which overlap the lamellar strips, these hairs much larger than those on
the dorsal surfaces of the petiole and postpetiole. Colour dark brown.
PARATYPE WORKER. TL 2-1, HL 0-56, HW 0-43, CI 77, ML 0-12, MI 21, SL 0-22, SI 51, PW 0-25, AL 0-60.
As holotype.
Holotype worker, Cameroun: nr Yaounde, sample SQ (G. Terron) (ENSA).
Paratypes. 1 worker, nr Yaounde, sample SV (G. Terron); 1 worker, nr Yaounde, sample YM (G.
Terron) (BMNH; ENSA).
The combination of characters noted in the species-group diagnosis quickly separates mvidurus
from all its Afrotropical congeners. The scale-like hairs on the head and body are very
conspicuous and at first sight are the most obvious feature of this species . Only one other African
species in the genus is similarly covered with scale-like hairs, tukultus, although several have
appressed scale-like hairs on the clypeal dorsum alone.
332 BARRY BOLTON
The f/mvidus-group
(Figs 29, 31)
Outline shape of head as Fig. 29. Mandibles with 11 teeth, the principal dental row containing 8 teeth of
about the same size, not strikingly enlarged but larger than the three apical denticles. Mandibles not as
strongly arched-downcurved as in preceding groups. Anterior clypeal margin transverse. Lateral clypeal
margins not expanded into a convex lobe on each side, the outer margins of the mandibles at full closure
intersecting the anterior clypeal margin at the anterolateral corners. Head not dorsoventrally flattened and
without a dorsal posteromedian broad tumulus. Posteriorly projecting rounded occipital lobes absent.
Pronotum not sharply marginate laterally, the dorsum transversely flat to shallowly convex. Cephalic
sculpture behind the clypeus either finely granular or with coarse foveolate punctures. Clypeus usually with
scale-like to spatulate hairs which are appressed, but with elevated pilosity in trymalus.
The three species included in this group, thuvidus from Kenya, tukultus from Ivory Coast and
trymalus from Cameroun, are those members of Glamyromyrmex in Africa which seem closest
to Smithistruma. At the start of this survey I was undecided about the correct generic assignment
for these three species but, considering the appearance of the mandibles in profile (Fig. 31), I
decided that Glamyromyrmex was the best generic fit that could be achieved at present. With the
revision of the short mandibulate dacetines of the world at generic level the position of these
species may change.
Although sharing the characters listed above two of the three species included here have
radically different sculpture and pilosity from the third, which may not be closely related. In
tukultus the mandibles and the entire cephalic dorsum are blanketed by appressed scale-like
hairs, which partially conceal the underlying fine dense reticulate-punctate sculpture of the
head. In contrast, thuvidus and trymalus lack this coat of scale-like hairs and have the cephalic
dorsum coarsely foveolate, the foveolae being so strongly developed that the head has a cratered
appearance.
Glamyromyrmex thuvidus sp. n.
(Fig. 31)
HOLOTYPE WORKER. TL 1-6, HL 0-43, HW 0-29, CI 67, ML 0-08, MI 19, SL 0-19, SI 66, PW 0-21, AL 041.
On the mandible the first tooth following the basal lamella small, about half the size of the second tooth.
Remaining teeth of the principal dental row showing some slight variation in size but none radically
reduced. In profile the upper and lower mandibular margins diverging from base to apex, the upper margin
weakly elevated anteriorly above the level of the anterior clypeal margin; the apical (masticatory) margin
arched-downcurved. Anterior clypeal margin transverse, the lateral margins not expanded, converging
behind to the convex frontal lobes, the width across which, from side to side, is only slightly less than the
maximum width of the clypeus. Upper scrobe margins distinctly indented behind the frontal lobes, convex
and divergent posteriorly. Occipital margin shallowly concave . Eyes vestigial, consisting of only one or two
ommatidia, situated on the ventral scrobe margins and the diameter of the eye less than that of the
foveolate punctures which occur on the side of the head behind the scrobe. Antennal scapes very weakly
clavate, the leading edges gently convex and with apically directed fine curved short hairs. Clypeus with
closely appressed short spatulate to scale-like hairs. Behind the clypeus the cephalic dorsum with fine
standing hairs which are arched or curved, mostly directed towards the midline; posteriorly some of the
arched hairs are directed forwards. Behind the clypeus the curved hairs are narrowly spatulate, more
posteriorly they are simple but some are minutely bifurcated at the apex. Sides of head with 4 pairs of
laterally projecting long flagellate hairs, the occipital margin with another pair towards the outer edges but
these tend to be directed upwards. Clypeus smooth and shining. Central strip of cephalic dorsum from
between the frontal lobes approximately to the highest point of the vertex mostly smooth, with a few
extremely minute fine rugular vestiges which are very indistinct. Dorsum behind and on each side of this
area with broad coarse foveolate punctures which give the surface a cratered appearance. The head shiny
and smooth between the punctures. Anterior border of pronotum marginate, the sides rounded and not
marginate. Sides of mesonotum and propodeum weakly marginate, the two confluent dorsally, not
separated by a transverse ridge or crest and the metanotal groove absent. Propodeal teeth mostly
incorporated in the infradental lamellae, with just a minute point projecting. Sides of alitrunk smooth
except for a few foveolate punctures anteriorly on the pronotum. Dorsal alitrunk smooth and shining
everywhere, devoid of sculpture. Pronotum at the humeri with a pair of laterally directed long flagellate
THE AFROTROPICAL DACETINE ANTS 333
hairs, the dorsum with two vertically directed pairs, one situated anteriorly and the other posteriorly.
Mesonotum with a single flagellate pair directed vertically. Otherwise the dorsal alitrunk only with
scattered fine hairs which arch towards the midline, some of these hairs minutely bifurcate apically. With
the pedicel segments in profile the ventral petiolar appendage reduced to an anteriorly situated broad lobe
beneath the peduncle , the lobe petering out about on a level with the highest point of the node . Lateral and
ventral appendages of postpetiole large but delicate and blister-like, translucent and with minute weak
veins present. Petiole in dorsal view with the node transverse, much broader than long, smooth and
shining. Lateral spongiform appendages vestigial, reduced to a minute and scarcely visible strip on each
side, without a transverse lamella connecting them across the posterior face of the node. Postpetiole in
dorsal view smooth and shining, the blister-like appendages prominent on each side and the anteriormost
parts of the ventral appendage visible, projecting in front of the anterior margin of the lateral appendage on
each side. Anterior and posterior margins of postpetiolar disc bordered by a narrow lamella, the disc itself
roughly trapezoidal in shape, narrowing posteriorly and with broadly rounded anterolateral corners, the
anterior margin deeply concave. Base of first gastral tergite lamellar, the lamella deeply concave and
almost interrupted medially. Basigastral costulae arise on each side of this concavity. Dorsal surfaces of
petiole, postpetiole and first gastral tergite each with scattered fine simple hairs which are mostly erect to
suberect. Colour glossy dull yellow.
PARATYPE WORKER. TL 1-6, HL 0-44, HW 0-29, CI 66, ML 0-09, MI 20, SL 0-20, SI 69, PW 0-22, AL 0-43.
As holotype.
Holotype worker, Kenya: Embu, Kirimiri Forest, W. of Runyenje, 1550 m, 3.x. 1977 (V. Mahnert &
J.-L. Ferret) (MRN).
Paratype. 1 worker with same data as holotype (BMNH).
G. thuvidus and trymalus are closely related and together are separated from tukultus as
tabulated below. Differences between thuvidus and trymalus are given under the latter.
thuvidus and trymalus
Mandibles not clothed in scale-like
hairs.
Cephalic dorsum behind clypeus with
arched simple hairs, and with
flagellate hairs present (5 pairs).
Sides of pronotum with a few anteriorly
situated foveolate punctures.
Ventral appendage of petiole lobiform,
not running the length of the
segment.
Petiole node much broader than long
in dorsal view, the posterior margin
without a transverse bordering
lamella.
Anterior margin of postpetiolar disc
concave in dorsal view.
tukultus
Mandible clothed in scale-like hairs.
Cephalic dorsum behind clypeus with
appressed scale-like hairs;
flagellate hairs absent.
Sides of pronotum unsculptured.
Ventral appendage of petiole massively
spongiform, running the length of
segment.
Petiole node about as broad as long
in dorsal view, the posterior
margin with a broad transverse
bordering lamella.
Anterior margin of postpetiolar
disc transverse in dorsal view.
Glamyromyrmex trymalus sp. n.
HOLOTYPE WORKER. TL 1-7, HL 0-43, HW 0-30, CI 70, ML 0-10, MI 23, (antennae lost), PW 0-22, AL 0-44.
Extremely closely related to thuvidus and answering to the description given for that species. In
particular, trymalus shows the same highly characteristic cephalic sculpture as thuvidus, having numerous
broad foveolate punctures with smooth spaces between them so that the head appears cratered. This very
distinctive sculpture coupled with minute size, yellow colour, and the characters noted in the species-group
diagnosis serve to isolate thuvidus and trymalus from all other Afrotropical Glamyromyrmex. The two are
separated as follows.
thuvidus
Clypeal dorsum with appressed short
scale-like to spatulate hairs.
trymalus
Clypeal dorsum without appressed hairs
of any description.
334
thuvidus - cont.
Elevated relatively long hairs absent
from clypeal dorsum.
Lateral and ventral appendages of
postpetiole delicate and blister-like,
translucent and with minute veins
present.
With postpetiole in dorsal view the
anterior margin of the disc
broadly and deeply concave.
Basalmost tooth of mandible only half
the size of the second tooth from
the base.
BARRY BOLTON
trymalus - cont.
Elevated relatively long hairs present
on clypeal dorsum.
Lateral and ventral appendages of
postpetiole conspicuously spongiform.
With postpetiole in dorsal view the
anterior margin of the disc evenly
shallowly concave.
Basalmost tooth of mandible only very
slightly smaller than the second
tooth from the base.
Holotype worker, Cameroun: nr Yaounde, sample AAU (G. Terrori) (ENSA).
Glamyromyrmex tukultus sp. n.
(Fig. 29)
HOLOTYPE WORKER. TL 1-6, HLO-41, HWO-29, CI 71, ML 0-10, MI 24, SLO-16, SI 55, PWO-20, ALO-44.
Mandibles covered in small scale-like appressed hairs. Basalmost tooth following the basal lamella
without a diastema, slightly smaller than the second tooth; all teeth after the second approximately the
same size except teeth nine and ten which are much reduced. There is a tendency for alternating slightly
smaller and slightly larger teeth in the principal dental row. In profile the mandibles broadening from base
to apex, the upper margin not conspicuously raised above the level of the anterior clypeal margin and with a
plateau-like slightly convex outline before arching steeply downwards. Anterior clypeal margin transverse.
Frontal lobes not distinctly convex, their margins more or less parallel, the width across the frontal lobes
from edge to edge conspicuously less than the maximum width of the clypeus; the preocular laminae plainly
visible in full-face view. Sides of head behind the frontal lobes shallowly convex and divergent. Occipital
margin evenly concave. Eyes small, situated on the ventral margin of the short but deep antennal scrobe.
Antennal scapes in full-face view with the leading edges moderately convex but distinctly indented near the
base, without projecting hairs but equipped with appressed small scale-like hairs which are also present on
the funicular segments. Clypeus and entire dorsum of head densely clothed with appressed scale-like hairs
which are directed anteriorly, the hairs situated more posteriorly on the head narrower and more spatulate
than those sited further forwards. Head without other pilosity of any description. Clypeus smooth, the
cephalic dorsum behind the clypeus finely reticulate-punctate but the sculpture largely concealed by the
pilosity. Anterior pronotal border marginate, the sides not marginate. Sides of mesonotum extremely
feebly marginate and sides of propodeum slightly more strongly so, the two segments confluent on the
dorsum, not separated by a transverse ridge or crest; the metanotal groove absent. Propodeal teeth entirely
incorporated in the infradental lamellae. Sides of alitrunk unsculptured and shining. Dorsal alitrunk
smooth and shining everywhere. Pronotum with three pairs of short erect simple hairs, one pair at the
humeri, one pair anterodorsally and another posterodorsally; mesonotum with a single pair of short erect
hairs. Apart from these the dorsal alitrunk with decumbent to appressed small hairs which are directed
towards the midline; those on the pronotum narrowly spatulate. In profile the spongiform appendages of
the pedicel segments massively developed, the petiolar ventral appendage running the length of the
segment, finely and densely spongiform and almost as deep as the maximum height of the node. Lateral
and ventral postpetiolar appendages thick and the basisternal pad on the gaster conspicuous. Petiole node
in dorsal view smooth, about as long as broad and with sharply defined anterior and posterior borders.
Lamellate appendage of node continuous across the posterior margin. Disc of postpetiole smooth and
shining, much broader than long and thickly surrounded at the sides by dense spongiform tissue, a lobe of
which projects strongly forwards from below the disc on each side of the petiole-postpetiole junction.
Anterior border of postpetiolar disc with a narrow lamellate margin. Base of first gastral tergite with a thick
costulate-spongiform transverse strip, the anterior margin of which is very feebly concave medially.
Basigastral costulae mostly confined to this strip, only feebly encroaching onto the tergite proper at the
sides. Petiole, postpetiole and gaster with fine simple hairs, the first also with a few narrowly spatulate
reclinate hairs. Colour glossy dull yellow.
V
PARATYPE WORKERS. TL 1-6, HL 0-42, HW 0-29-0-30, CI 69-71, ML 0-10, MI 24, SL 0-16-0-17, SI 55-59,
PW 0-20, AL 0-44 (3 measured).
THE AFROTROPICAL DACETINE ANTS
335
As holotype but infradental lamellae may have a point developed apically.
Holotype worker, Ivory Coast: Bingerville, 29.X.1980 (V. Mahnert &J.-L. Ferret) (MHN).
Paratypes. 3 workers with same data as holotype (MHN; BMNH; MCZ).
Closest related to thuvidus, differences separating the species are tabulated under that name.
The most obvious character separating tukultus from all other members of the genus except
ravidurus is the dense covering of scale-like hairs everywhere on the head and its appendages. G.
ravidurus is a much larger darker species than tukultus and the characters given in their
respective species-group diagnoses will differentiate the two.
SERRASTRUMA Brown
(Figs 34-44)
Serrastruma Brown, 1948: 107 [as subgenus of Smithistruma]. Type-species: Strumigenys simoni Emery,
1895a: 42, pi. 2, fig. 21, by original designation.
Serrastruma Brown; Brown, 19490: 6. [Raised to genus.]
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles elongate-triangular (MI 26-50), serially
finely and densely denticulate along the entire masticatory margin, with >20 denticles; without an apical
fork of spiniform teeth. Mandibular denticles either uniformly small or with the basal 4-8 enlarged,
commonly the basalmost denticles enlarged in either case. Mandibles lacking a differentiated prominent
basal lamella and shallowly curved in profile. Antennae with 6 segments.
All 11 presently recognized species of Serrastruma are found in sub-Saharan Africa. Most are
restricted to that region but a couple of species (ludovici and simoni) have successfully invaded
the Malagasy region. Three species (lujae, serrula and may net) have been found on the islands in
the Gulf of Guinea. All nest in rotten wood, in the leaf litter or topsoil layer, or in the stumps of
trees. Foraging is mainly conducted in the leaf litter or topsoil layers but individuals may ascend
bushes and trees to a height of a metre or more, and maynei may form nests in trunks some
distance above the ground. The predatory behaviour of S. serrula has been investigated in some
detail by Dejean (19800; 19806).
The genus Serrastruma has its origins in Smithistruma, being derived and differentiated from
that genus by the specialized form of the mandibles in Serrastruma. Brown (1952a: 71) has
postulated an origin for Serrastruma in the Smithistruma alberti-group or capitata-group and
maintains that the elongate basal lamella of the mandible seen in these groups has in Serrastruma
become denticulate all along its inner free margin and has been pressed into service as part of the
masticatory surface, the original smithistrumiform dentition being represented only by the
apical group of denticles in Serrastruma and most of the length of the mandible being
represented by this secondary development. This explanation effectively accounts for the
mandibular form and dentition seen in Serrastruma and also accounts for the lack of a
differentiated basal lamella in the genus, the presence of which is characteristic of Smithistruma
and all its close relatives. The differences in mandibular form between Serrastruma and
Smithistruma in the Afrotropical fauna may be tabulated as follows.
Serrastruma
Mandibles elongate triangular,
relatively long, MI 26-50.
Mandibles without a differentiated
projecting basal lamella.
Masticatory margin of mandible with
>20 (usually at least 30) fine
small denticles, sometimes with the
basal 4-8 enlarged, often with the
basalmost enlarged.
Smithistruma
Mandibles triangular to subtriangular,
relatively short, MI 7-20.
Mandibles with a differentiated
projecting basal lamella (usually
concealed by clypeus when closed).
Masticatory margin of mandibles usually
with 12 teeth or denticles, in one
species with 16-17, the basal 5-7
of which form the principal dental
row.
336
Serrastruma - cont.
Base of mandibles not extensively
overlapped by the clypeus in
full-face view, the latter with
a false anterior margin above the
true margin.
BARRY BOLTON
Smithistruma - cont.
Base of mandibles extensively
overlapped by the clypeus in
full-face view, the latter lacking
a false anterior margin above the
true margin.
Historically Serrastruma dates back only to 1948 when Brown first defined the group as a
subgenus of Smithistruma. Prior to that date the members of Serrastruma had been treated
either as belonging to Strumigenys (Arnold, 1917) or grouped together with Smithistruma in a
subgenus of Strumigenys called Cephaloxys (Wheeler, 1922). Brown (1948) showed that several
disparate groups were included in Strumigenys. He erected the genus Smithistruma to hold most
of the short-mandibulate forms previously included in Strumigenys (the name Cephaloxys being
preoccupied) and treated Serrastruma as a subgenus of Smithistruma. The next year Brown
(1949a) recognized that Serrastruma and Smithistruma had differently constructed mandibles
and raised the former name to full generic status.
Subsequently Brown (19520) issued an extensive revision of Serrastruma which cut down the
27 named forms then in the genus to seven recognized species plus two species inquirendae.
Since the publication of this revision much type-material unobtainable by Brown has become
available for study, along with a great many more samples of Serrastruma, all of which has
greatly facilitated the present investigation. Fundamentally Brown's treatment of the genus
remains unchanged except for the addition of five new species and a reinterpretation of a few
names, most of these belonging to forms where the type-series was not available to Brown while
conducting his survey. The changes from his revision are summarized as follows. Brown's
(1952a: 85) species inquirendae included the names ludovici and calypso. The types of these
were not available for his study but he speculated that ludovici was the senior synonym of
alluaudi. Brown has informed me that he confirmed this synonymy some time ago and my
examination of the types agrees with his conclusion. As for calypso, Brown supposed it might be
synonymous with lotti. However, an examination of the holotype, which was not available for
study at the time of Brown's revision, shows it to be a straight synonym of lujae.
S. lotti and bequaerti, treated by Brown (1952a: 76, 80) as valid species, are now included as
synonyms of ludovici and lujae respectively. Some time ago Brown informed me that he had
recognized both these synonyms, and I have merely confirmed them here.
S. alluaudi st. nigeriensis was given by Brown (1952a: 83) as a synonym of simoni but a
re-examination of the mandibles of the syntype-series has revealed the characteristic dentition
of ludovici, and nigeriensis has now been referred to the synonymy of ludovici.
Brown (1952a: 75) recognized that the type-series of raymondi was a mixture of alluaudi (now
ludovici) and simoni, and synonymized raymondi under the former name. However, the
specimen labelled as holotype by Donisthorpe, the author of raymondi, is plainly a specimen of
simoni, and raymondi is therefore transferred to the synonymy of that species.
Finally concolor is removed from the synonymy ofserrula and reinstated as a valid species, for
reasons given in the discussion of concolor, below.
In terms of number of species Serrastruma falls well behind Smithistruma and Strumigenys,
but in terms of numbers of individuals and the range of distribution of the species Serrastruma far
outstrips the rest. Its species are easily the most successful dacetine ants in sub-Saharan Africa
and in terms of number of samples encountered in collections Serrastruma material amounts to
more than that of all the other genera combined. It is this commonness, coupled with an innate
tendency of the most widely distributed species to show variation in sculpture and size, which has
been mostly responsible for the extensive synonymy.
List of Afrotropical Serrastruma
concolor (Santschi) sp. rev. ludovici (Forel)
dotaja sp. n. alluaudi Santschi syn. n.
geoterra sp. n. alluaudi st. nigeriensis Santschi syn. n.
inquilina sp. n. rothkirchi Wasmann
escherichi subsp. lotti Weber syn. n.
THE AFROTROPICAL DACETINE ANTS
337
lujae (Forel)
reticulata Stitz
glanduscula Santschi
bequaerti Santschi syn. n.
gerardi Santschi
calypso Santschi syn. n.
aequalis Menozzi
maynei (Forel)
maynei var. latiuscula Forel
miccata sp. n.
serrula (Santschi)
uelensis Santschi
simoni (Emery)
escherichi Forel
cognata Santschi
biconvexa Santschi
cognata st. boerorum Santschi
escherichi r. limbata Forel
escherichi var. cliens Forel
escherichi st. cognata var.
obscuriventris Santschi (unavailable)
escherichi st. cognata var.
fusciventris Santschi (unavailable)
raymondi Donisthorpe syn. n.
sulumana sp. n.
Key to species (workers)
Note. The parasitic 5. inquilina, known only from a series of females found in a nest of lujae, is omitted
from the key.
1 With the head in full-face view the upper scrobe margin just behind the level of the eyes without
a projecting long hair on each side 2
- With the head in full-face view the upper scrobe margin just behind the level of the eyes with a
long flagellate or simple fine hair projecting on each side 5
2 Dorsal surfaces of head and alitrunk with very conspicuous dense thickly spatulate to scale-like
coarse ground-pilosity (Figs 36, 37) . Antennal scapes relatively short , SI 73-83 3
- Dorsal surfaces of head and alitrunk with inconspicuous slender very narrowly spatulate
ground-pilosity. Antennal scapes relatively long, SI 1 16-127 4
3 Dorsal alitrunk sculptured. Sides of head not broadly convex posteriorly (Fig. 37). (Ivory
Coast, Ghana, Nigeria, Cameroun, Annobon I. , Zaire, Angola, Uganda, Tanzania)
maynei (p. 347)
Dorsal alitrunk glassy smooth. Sides of head broadly convex posteriorly (Fig. 36). (Cameroun)
geoterra (p. 341)
4 Pronotal humeri each with a short stout clavate hair. Leading edges of scapes with projecting
short clavate hairs. Mandibles shorter, MI 26, the basal 4-5 denticles suddenly and
conspicuously enlarged. Metanotal groove not impressed. (Ghana) miccata (p. 348)
- Pronotal humeri each with a long fine flagellate hair. Leading edges of scapes with fine simple
outcurved short hairs. Mandibles longer, MI 44-45, the basal 4-5 denticles not enlarged.
Metanotal groove impressed. (Cameroun) sulumana (p. 352)
5 Principal sculpture of pronotal dorsum dense reticulate-punctation, with or without overlying
costulate or striate sculpture. When such overlying sculpture is present it is distinctly
secondary to the dense punctate component 6
- Principal sculpture of pronotal dorsum consisting of longitudinal to oblique striae or costulae,
or the surface mostly to entirely unsculptured and smooth; punctate sculpture usually absent
but when present is very feeble and distinctly secondary to the striate or costulate component 9
6 With the head in full-face view the basal series of denticles on the mandible suddenly enlarged ,
broader longer and coarser than those preceding; usually this enlarged series very distinct
(Fig. 34). (Extremely widespread in Afrotropical region, Madagascar, Mauritius)
ludovici (part, p. 343)
- With the head in full-face view the basal series of denticles on the mandible not enlarged, the
denticles approximately the same size throughout the length of the margin or very evenly and
gradually increasing in size basally 7
7 Mesonotum with 3-4 pairs of short erect relatively slender hairs, the posteriormost pair
situated at or close to the metanotal groove. Ground-pilosity of promesonotum strong, not
reclinate, conspicuous in profile (Fig. 42). Punctate sculpture of head and alitrunk strong and
sharply defined . (West and central Africa, Sao Tome I. ) serrula (p. 349)
- Mesonotum usually with a single pair of erect apically thickened to strongly clavate hairs which
are situated anteriorly on the segment. Very rarely a second pair present but if so these are
close to the first pair and remote from the metanotal groove. Ground-pilosity of promesono-
tum feeble, mostly or entirely reclinate, inconspicuous in profile (Figs 43, 44). Punctate
sculpture on head and alitrunk usually fine 8
338 BARRY BOLTON
8 Dorsum of head behind highest point of vertex with 4 relatively slender simple standing hairs
and with a further pair situated in front of the highest point of the vertex. With the head in
full-face view the upper scrobe margins not indented at the site of the projecting hair.
Postpetiole with more than one pair of standing hairs. (Extremely widespread in Afro-
tropical region) lujae (p. 345)
Dorsum of head behind highest point of vertex with 2-4 thickened apically clavate standing
hairs; without standing pilosity in front of the highest point of the vertex. With the head in
full-face view the upper scrobe margins usually conspicuously indented at the site of the
projecting hair. Postpetiole with only a single pair of standing hairs. (West and Central
Africa, Uganda) concolor (p. 338)
9 Pronotum (at the humeri) and mesonotum each with a pair of exceptionally long fine flagellate
hairs (Fig. 38). First gastral tergite with only a single pair of long simple hairs, which are not
thickened or clavate apically and are situated close to the base of the sclerite. (Cameroun,
Gabon) dotaja (p. 339)
- Pronotum (at the humeri) with flagellate hairs but mesonotum never with such hairs. Usually
the mesonotum with a single pair (very rarely 2 pairs) of stout hairs which are thickened or
clavate apically (Figs 40, 41). First gastral tergite with hairs present which are thickened or
clavate apically 10
10 Basal series of mandibular denticles suddenly and conspicuously enlarged (Fig. 34). With
postpetiole in profile the area of the ventral spongiform lobe less than that of the exposed
portion of the postpetiolar disc (Fig. 41). (Extremely widespread in Afrotropical region,
Madagascar, Mauritius) ludovici (part, p. 343)
- Mandibular denticles small fine and regular to base. With postpetiole in profile the area of the
ventral spongiform lobe distinctly greater than that of the exposed portion of the postpetiolar
disc (Fig. 40). (Eastern and southern Africa, Nigeria, Mauritius) simoni (p. 350)
Serrastruma concolor (Santschi) sp. rev.
(Fig. 43)
Strumigenys (Trichoscapa) concolor Santschi, 1914ft: 375. Syntype workers, female, GHANA: Aburi
(NMB) [examined]. [Previously synonymized with serrula by Brown, 1952a: 81.]
WORKER. TL 2-0-2-2, HL 0-48-0-52. HW 0-40-0-46, CI 80-90, ML 0-16-0-18, MI 32-36, SL 0-30-0-33, SI
67-80, PW 0-28-0-32, AL 0-56-0-62 (12 measured).
Mandibular denticles evenly sized to the base, without a basal series of 4-8 denticles which are enlarged
though the basalmost may be larger than those preceding. Upper scrobe margins well developed, with a
flagellate hair at each side just behind the level of the eye and with the margin indented or impressed at the
site of the hair. Width and depth of this indentation variable, usually conspicuous in full-face view but
sometimes only shallow. Ground-pilosity of head short and narrowly spatulate, closely applied to the
surface and indistinct. Dorsum of head behind highest point of vertex with 2 or 4 elongate standing hairs
which are anteriorly curved, stout and narrowly clavate apically. Samples with only one pair of these hairs
generally have the upper scrobe margins more strongly impressed at the site of the flagellate hair than do
those samples with two pairs of hairs. Entire dorsum of head densely finely reticulate-punctate. Alitrunk in
profile with promesonotum convex, sloping down posteriorly to the impressed metanotal groove.
Propodeal dorsum convex, sloping behind to a pair of teeth which are variable in size. Infradental lamellae
present on propodeal declivity. Sides of pronotum superficially punctulate to smooth, the pleurae usually
smooth but in some their upper halves finely punctulate. Dorsal alitrunk finely reticulate-punctate
everywhere but the intensity of sculpture on the pronotum variable and sometimes the punctures filled by a
waxy superficial layer. Occasionally weak superficial rugulae, generally formed by alignment of puncture
margins, may occur, but these are always distinctly secondary to the punctate sculpture. Pronotum with
flagellate hairs present at the humeri. Mesonotum with a single anteriorly situated pair of standing hairs
which are stout long conspicuous and clavate apically. Dorsal alitrunk otherwise only with sparse
ground-pilosity which is short, inconspicuous and generally closely applied to the surface. Spongiform
appendages of pedicel segments moderately developed in profile. Petiole in dorsal view usually punctate
but the sculpture very faint in some specimens. Postpetiole usually superficially punctulate but almost
smooth in some, with a posterior spongiform strip which abuts a similar narrow strip on the base of the first
gastral tergite. Basigastral costulae present, the tergite otherwise unsculptured. Petiole and postpetiole
dorsally each with a single pair of elongate stout hairs which are clavate apically, the first gastral tergite with
numerous similar hairs. Colour yellow to yellowish brown.
THE AFROTROPICAL DACETINE ANTS
339
This small species was earlier treated as a synonym of serrula by Brown (1952a: 81), but on
examining the numerous series assigned to serrula it became apparent that two distinct species
were in fact present. One of these matched the holotype of serrula (= uelensis) and the other the
type-series of concolor, which is therefore returned to species rank. The two are separated by
the following characters in the worker.
concolor (Fig. 43)
Mesonotum with a single anteriorly
situated pair of elongate stout
apically clavate standing hairs.
Petiole and postpetiole each with only
a single pair of standing hairs.
Ground-pilosity of head and alitrunk
short sparse and inconspicuous,
closely applied to the surface; the
standing hairs on the cephalic dorsum
very obviously longer and stouter
than the ground-pilosity.
Alitrunk pleurae unsculptured at
least on lower halves.
Punctate sculpture of pronotal
dorsum fine and not sharply defined.
serrula (Fig. 42)
Mesonotum with 3 (rarely 4) pairs of
more slender standing hairs which
are weakly or scarcely clavate, the
posteriormost pair the shortest and
situated at the metanotal groove.
Petiole usually and postpetiole always with
more than one pair of standing hairs.
Ground-pilosity of head and alitrunk
moderately long and dense, very
conspicuous and elevated; the
standing hairs on the cephalic dorsum
only slightly longer and stouter
than the ground-pilosity.
Alitrunk pleurae punctate but sometimes
only superficially so.
Punctate sculpture of pronotal dorsum
strong and sharply defined.
One other species is close to concolor, the common lujae, a species that shows a remarkable
size range but whose diagnostic characters remain quite stable. S. concolor is persistently small,
overlapping only the lower end of the range of lujae, and it differs by having 2-4 standing hairs
posteriorly on the cephalic dorsum which are distinctly clavate apically, as opposed to the 6 fine
hairs on the cephalic dorsum in lujae which are scarcely or not at all swollen apically. In lujae the
6 hairs are arranged in a posterior row of 4 and an anterior pair, situated just in front of the
highest point of the vertex. The postpetiole in concolor always bears a single pair of stout hairs;
in lujae two or more pairs occur.
I consider it possible that concolor, as presently constituted, may consist of two sibling
species. It is noticeable that the samples with a single pair of standing cephalic hairs have the
upper scrobe margins more strongly impressed at the site of the projecting hair than do those
with two pairs of standing hairs. Whatever significance this may have cannot be investigated at
the present time as not enough material of the two forms is available for comparison.
MATERIAL EXAMINED
Ivory Coast: Man (V. Mahnert & J.-L. Ferret); Sassandra (7. Lobl). Ghana: Mampong (P. Room);
Aburi. Cameroun: Mt Cameroun, Buea slope (B. Malkin); nr Yaounde (G. Terron). Gabon: Plateau
d'Ipassa (J. A. Band). Zaire: Ituri Forest, Beni Irumu (N. A. Weber); Ituri Forest, Epulu (T. Gregg);
Niangara (N. A. Weber); Ruwenzori, Mwenda (J. C. Bradley). Angola: R. Camudembele (L. de
Carvalho); R. Mussungue (L. de Carvalho); Dundo; Gubela (P. Hammond). Chad: Haut Mbomu (N. A.
Weber). Uganda: Ft Portal (N. A. Weber).
Serrastruma dotaja sp. n.
(Figs 35, 38)
HOLOTYPE WORKER. TL 2-3, HL 0-53, HW 0-42, CI 79, ML 0-22, MI 42, SL 0-36, SI 86, PW 0-32, AL 0-62.
Mandibular denticles small even and regular, without a basal series of 4-8 enlarged denticles; only the
basalmost denticle enlarged. Upper scrobe margins narrow and petering out posteriorly, not composed of
a broad continuous lamellate granular flange; equipped just behind the level of the eye with an extremely
long flagellate hair which basally projects laterally from the margin. Clypeus smooth on the disc, with
appressed small spatulate hairs, the anteriormost row of spatulate hairs situated on the false margin of the
clypeus and freely projecting forwards over the mandibular bases. Ground-pilosity of head to highest point
of vertex consisting only of sparse narrow inconspicuous hairs which are decumbent to appressed and
340
BARRY BOLTON
directed anteriorly. Dorsum of head behind highest point of vertex with similar but slightly longer
ground-pilosity and also with two pairs of anteriorly curved long simple hairs, one pair situated just behind
the highest point and the other latero-occipitally. Dorsum and sides of head densely and strongly
reticulate-punctate everywhere. Pronotum marginate anteriorly but laterally the sides of the alitrunk
separated from the dorsum only by smoothly rounded blunt angles. In profile the pronotum and anterior
part of the mesonotum forming an even shallow convexity, the posterior part of the mesonotum sloping
down to the shallowly impressed metanotal groove. Propodeal dorsum curving down posteriorly to the
bases of the strong propodeal teeth which are elevated and slightly upcurved. Infradental lamellae present
down the depth of the propodeal declivity, its width equal to or slightly greater than the diameter of the
propodeal spiracle. Sides of alitrunk glassy smooth, devoid of sculpture. Dorsal surfaces of pronotum,
anterior mesonotum and propodeum glassy smooth. Posterior (sloping) portion of mesonotum smooth
centrally but with some weak lateral punctulae and posteriorly with some irregular sculpture just in front of
the metanotal groove. Propodeal declivity reticulate-punctate between the bases of the teeth. Pronotum
and mesonotum each equipped with a pair of extremely long fine flagellate hairs, each hair arising from a
small papilla. Dorsum of promesonotum otherwise only with very sparse short ground-pilosity which is
subdecumbent. With the pedicel segments in profile the spongiform appendages well developed. Ventral
petiolar process spongiform posteriorly but more solid and opaque anteriorly. Ventral and lateral
spongiform lobes of postpetiole about equal in size, lateral appendage of petiole node smaller. In dorsal
view both petiole and postpetiole smooth and very shiny, the former with a distinct spongiform strip
running across the posterior face. Disc of postpetiole transversely roughly oval in dorsal view, the anterior
face with a narrow transverse lamellate strip, the sides bordered with spongiform material which becomes
broader posteriorly, and the posterior margin bordered with a broad spongiform strip the posterior margin
of which is shallowly concave medially. Extreme base of first gastral tergite lamellate spongiform, with a
continuous band of short basigastral costulae which do not run further back than the pair of long gastral
hairs; remainder of tergite glassy smooth. Petiole and postpetiole each with a single pair of stout simple
posteriorly curved long hairs. First gastral tergite with a single pair of simple hairs which are situated close
to the base of the sclerite. Colour jet black, glossy behind the head, the appendages brown.
PARATYPE WORKERS. TL 2-3-2-5, HL 0-50-0-54, HW 0-41-0-43, CI 78-82, ML 0-20-0-23, MI 38-43, SL
0-34-0-37, SI 81-88, PW 0-31-0-34, AL 0-60-0-66 (20 measured).
As holotype but specimens from Gabon are slightly lighter in colour, blackish brown with somewhat
lighter pedicel segments.
Holotype worker, Cameroun: Nkoemvon, 16.iii.1980 (D. Jackson) (BMNH).
Paratypes. Cameroun: 12 workers with same data as holotype; 3 workers with same data but 25. xi. 1980,
N52. Gabon: 11 workers, Plateau d'Ipassa, 6, IPA CI9 (/. A. Band); 1 worker with same data but IVI5,
IPA 8 (BMNH; MCZ; MHN).
Non-paratypic material examined. Cameroun: nr Yaounde (G. Terron).
The presence of very long fine flagellate hairs on the mesonotum as well as on the pronotum
immediately distinguishes dotaja from all other species as in those the mesonotum has one or
more pairs of stout hairs which are thickened or clavate apically. The species lujae, serrula,
concolor and maynei are also distinguished from dotaja by their possession of conspicuous
pronotal sculpture, and maynei, sulumana and geoterra lack cephalic flagellate hairs. From
ludovid samples in which the enlarged basal series of mandibular denticles is not very strongly
developed dotaja differs in having the pleurae and propodeal dorsum smooth, as well as by
having simple cephalic, petiolar, postpetiolar and gastral pilosity, all of which are clavate in
ludovid. S. simoni, the closest relative of dotaja, is separated as follows.
dotaja (Fig. 38)
Standing hairs behind highest point
of cephalic vertex simple.
Mesonotum with a pair of very long
flagellate hairs similar to those
on the pronotum.
Pilosity of petiole, postpetiole and
gaster each of a single pair of
simple curved hairs.
Pronotal dorsum glassy smooth.
simoni (Fig. 40)
Standing hairs behind highest point
of cephalic vertex clavate.
Mesonotum with a pair of stout clavate
hairs contrasting with the flagellate
hairs on the pronotum.
Pilosity of petiole, postpetiole and
gaster each of one or more pairs of
clavate hairs.
Pronotal dorsum at least with scattered
vestiges of sculpture.
THE AFROTROPICAL DACETINE ANTS 341
dotaja (Fig. 38) - cont. simoni (Fig. 40) - cont.
Dorsum of propodeum and of petiole Dorsum of propodeum punctate, very
node glassy smooth. very rarely with a smooth median area;
petiole node punctate dorsally even if
only feebly so.
Disc of clypeus smooth (when clean) . Disc of clypeus sculptured (when clean) .
MI 38-43; ML is 0-48-0-53 xHW. MI 33-39; ML is 0-42-0-48 xHW.
A single worker of simoni from Burundi (in MCZ) with very reduced sculpture is responsible
for the qualifying 'at least' and 'very rarely' in the sculpture characters listed above. In this
specimen the sculpture is more reduced than is usual in simoni and it is more like dotaja in this
respect than any other sample examined. However, even in this individual the pilosity characters
are absolutely those of simoni.
Serrastruma geoterra sp. IK
(Fig. 36)
HOLOTYPE WORKER. TL 2-6, HL 0-58, HW 0-52, CI 90, ML 0-23, MI 40, SL 0-38, SI 73, PW 0-34, AL 0-74.
Mandibles with denticles very slightly enlarging basally, the increase in size minute and gradual, without
a suddenly much enlarged basal series. False anterior margin of clypeus with a row of very conspicuous
broad flattened hairs which project straight forwards over the real clypeal margin and the bases of the
mandibles. Upper scrobe margins expanded into a very broad conspicuous lamella which runs the length of
the scrobe, is slightly elevated and is of approximately equal width throughout its length. This upper
scrobal lamella is so broad posteriorly that the eyes are concealed and not visible in full-face view; the
margin without a pair of projecting long hairs just behind the level of the eyes. Antennal scapes feebly
curved, broadened after the basal third and narrowing again apically, their leading edges with a projecting
row of short broad flattened hairs which are truncated apically. Ground-pilosity of head consisting
everywhere of very thick short blunt off-white to yellowish hairs which in profile can be seen to be
dorsoventrally flattened and strongly curved anteriorly. Close to the occipital margin is a single pair of
similarly constructed but longer, slightly more erect hairs. Eyes small, of 5-6 ommatidia. Clypeus with very
feeble superficial reticular sculpture, a narrow strip behind the clypeus and following the shape of the
posterior clypeal margin depressed and smooth. Behind this the head weakly reticulate-punctate, this
sculpture fading posteriorly and the area between the highest point of the vertex and the occipital margin
smooth except for the pits from which the hairs arise. Broad flange of upper scrobe margin densely
reticulate-granular. Pronotum not marginate laterally, the humeri without flagellate hairs. Pronotum in
dorsal view almost twice wider than the mesonotum, the two separated by a shallow arched-transverse
impression. Metanotal groove represented by a transverse line across the dorsum but not impressed. With
the alitrunk in profile the pronotal outline separated by a shallow impression from the mesonotum, the
anterior half of the latter raised and on the same level as the pronotum. Behind this the mesonotum
descending almost vertically and its posterior half flat or shallowly concave to the level of the metanotal
groove, behind which the propodeal dorsum is shallowly convex and sloping posteriorly. Propodeal teeth
triangular, with a narrow but conspicuous infradental lamella. Dorsum and sides of alitrunk glassy smooth
everywhere, the propodeal declivity with reticular vestiges on its upper half. Pilosity of dorsal alitrunk as
on head, the hairs curved anteriorly or medially on the pronotum and mesonotum, posteriorly on the
propodeum. Mesonotum with a pair of similarly constructed but larger hairs at the point where the surface
begins its sudden descent. Spongiform appendages of pedicel segments small, the subpetiolar process
reduced to a thin strip and the subpostpetiolar lobe much smaller than the exposed area of the postpetiolar
disc in profile. Dorsum of petiole node smooth but the peduncle and sides reticulate; posterior spongiform
strip of node lamellate and narrow. Postpetiole smooth in dorsal view, posteriorly with a thin spongiform
strip which abuts a similar thin strip on the base of the first gastral tergite. Basigastral costulae very short,
no longer on the tergite than the width of basal spongiform strip; gaster otherwise smooth. Dorsal surfaces
of petiole, postpetiole and gaster with ground-pilosity similar to that described for the head, and with
paired longer more erect hairs which are thick, sturdy and broadly clavate. Colour glossy brown, the legs
with dense scale-like decumbent to appressed pilosity.
PARATYPE WORKER. TL 2-7, HL 0-60, HW 0-54, CI 90, ML 0-23, MI 38, SL 0-40, SI 74, PW 0-37, AL 0-75.
As holotype.
Holotype worker, Cameroun: nr Yaounde, sample 2513 (G. Terrori) (ENSA).
Paratype. 1 worker with same data but sample TS (BMNH).
342
BARRY BOLTON
This very distinctive species appears closest related to maynei, the two having broad upper
scrobe margins, conspicuous cephalic ground-pilosity, lacking flagellate hairs on the head and
pronotum, lacking dense punctate pronotal sculpture, and having antennal scapes of moderate
length. They are easily separated asgeoterra has the alitrunk absolutely smooth whilst in maynei
sculpture is present. Apart from this the alitrunk is characteristically shaped in geoterra, the
cephalic pilosity is even broader and coarser than in maynei, and the sides of the head are
conspicuously more convex because of the wide upper scrobe margin in geoterra, compare
Figs 36, 37.
Serrastruma inquilina sp. n.
HOLOTYPE FEMALE. TL 3-0, HL 0-65, HW 048, CI 74, ML 0-22, MI 34, SL 0-52, SI 108, PW 0-48, AL 0-80.
Mandibular denticles very gradually increasing in size towards base. Antennal scrobes vestigial, the
upper scrobe margins not differentiated behind the level of the anterior margin of the eye ; behind this point
the dorsum of the head rounding into the sides. Flagellate hairs absent from the head. Clypeus glassy
smooth, dorsum of head finely and densely punctate, the punctures superficial and the surface shining.
Pilosity of head consisting entirely of fine simple curved hairs which are directed anteriorly except in the
vicinity of the ocelli where they are directed approximately towards the mid-dorsal point. Dorsum of head
without elongate standing specialized hairs. With the alitrunk in profile the propodeum unarmed, without
trace of teeth. Sides of pronotum densely reticulate-punctate, the lateral portions of the mesoscutum,
above the pronotum, more finely punctulate. Mesopleuron smooth except for the strip immediately below
the wing insertion which is punctate. Metapleuron punctate in the upper half, smooth below. Sides of
propodeum densely reticulate-punctate. Mesoscutum with a broad central smooth area but the periphery
of the sclerite tending to be punctulate. Scutellum weakly punctulate, the propodeum densely reticulate-
punctate. With the pedicel segments in profile spongiform appendages are absent from the petiole and very
reduced on the postpetiole where they are represented only by a small lateral and ventral lobe. In dorsal
view the petiole node smooth, as long as broad and lacking a transverse lamellar or spongiform strip
posteriorly. Postpetiole slightly broader than long, smooth, and having a narrow lamellate strip traversing
the posterior margin. First gastral tergite smooth, without trace of basal costulae. Pilosity everywhere on
dorsal surfaces of alitrunk, petiole, postpetiole and gaster consisting of quite dense fine simple soft curved
hairs which are subdecumbent to decumbent, pointed apically and directed approximately towards the
midline on the alitrunk and posteriorly on the pedicel segments and the gaster; without long flagellate or
any other specialized hairs. Colour yellow.
PARATYPE FEMALES. TL 2-9-3-1, HL 0-64-0-68, HW 0-46-0-48, CI 71-74, ML 0-20-0-22, MI 30-34, SL
0-50-0-52, SI 108-113, PW 0-46-0-48, AL 0-78-0-85 (7 measured).
As holotype, the petiole node in dorsal view may be fractionally longer than broad.
Holotype female (alate), Zaire ('B. Congo' on data label): S. slope of Mt Kahuzi, 1900 m, 5.ix.l957, in
nest of Serrastruma lujae (Forel) (E. S. Ross & R. E. Leech) (CAS).
Paratypes. 7 females with same data as holotype (CAS; MCZ; BMNH).
This series of females, found in a nest of the common S. lujae, constitutes the first known socially
parasitic dacetine in the Afrotropical region. It is easily distinguished from the female of the
host-species, as follows.
inquilina
Scrobes vestigial, upper scrobe
margins not sharply defined behind
level of anterior margin of eye.
Head without laterally projecting long
fine hairs on upper scrobe margins;
without specialized standing hairs
on cephalic dorsum.
Alitrunk and first gastral tergite
with fine soft curved hairs only,
without standing hairs.
Pronotal humeri without flagellate hairs.
Propodeum unarmed.
Basigastral costulae absent.
lujae
Scrobes present, upper scrobe margin
sharply defined to beyond level of
posterior margin of eye.
Head with laterally projecting long
fine hairs on upper scrobe margins;
with specialized standing hairs
on cephalic dorsum.
Alitrunk and first gastral tergite
without fine soft curved hairs,
with standing hairs.
Pronotal humeri with flagellate hairs.
Propodeum bidentate.
Basigastral costulae present.
THE AFROTROPICAL DACETINE ANTS 343
inquilina - cont. lujae - com.
Petiole node in dorsal view as long Petiole node in dorsal view broader than
as or slightly longer than broad, long, punctate, with a transverse
smooth, lacking a transverse spongiform strip posteriorly,
spongiform strip posteriorly.
Scapes long, SI >100 Scapes short, SI <100.
Serrastruma ludovici (Forel)
(Figs 34, 41)
Strumigenys ludovici Forel, 19046: 369. Syntype workers, MADAGASCAR: 'Madagascar meridional', 1899
(M. Sikoro) (MHN) [examined].
Strumigenys alluaudi Santschi, 19106: 360. Syntype workers, TANZANIA: Tanga Cave, Kulumuzi, iv.1909
(Ch. Alluaud) (NMB) [examined]. Syn. n.
Strumigenys (Trichoscapa) alluaudi st. nigeriensis Santschi, 19146: 376. Syntype workers, NIGERIA:
Olokemeji (F. Silvestri) (NMB) [examined]. [Wrongly synonymized with simoni by Brown, 1952a: 83.]
Syn. n.
Strumigenys rothkirchiWasmann, 1918: 142, pi. 2, figs 9, 10. Syntype workers, CAMEROUN: noloc., 1912 (v.
Rothkirch) (MNHU). [Synonymized with alluaudi by Brown, 1952a: 75.]
Strumigenys (Cephaloxys) escherichisubsp. lotti Weber, 1943: 378, pi. 15, fig. 13. Syntype workers, SUDAN:
Equatoria, Lotti Forest, 5.viii.l939, no. 1451 (N. A. Weber) (BMNH; MCZ; MRAC) [examined].
Syn. n.
Serrastruma alluaudi (Santschi) Brown, 1952a: 75.
Serrastruma lotti (Weber) Brown, 1952o: 76. [Raised to species.]
Serrastruma ludovici (Forel) Brown, 1952o: 85. [Species inquirenda.]
WORKER. TL 1-9-3-0, HL 0-46-0-66, HW 0-36-0-49, CI 71-80, ML 0-17-0-30, MI 35-50, SL 0-32-0-50, SI
86-115, PW 0-26-0-38, Al 0-55-0-80 (85 measured).
Basal 4-8 denticles on mandibular masticatory margin enlarged, usually very conspicuously coarser
broader and longer than the preceding denticle row. In a few samples the enlarged denticles not so obvious
and sometimes the enlarged series may be better developed on one mandible than on the other. Upper
scrobe margins defined by a narrow flange or rim which is broadest just behind the frontal lobes and peters
out posteriorly, frequently the rim not even extending to the apex of the scrobe. Upper scrobe margins
each with a fine flagellate hair just behind the level of the eye. Clypeus usually finely reticulate to punctate,
only rarely the sculpture reduced. Dorsum of head behind clypeus finely and densely reticulate-punctate.
Ground-pilosity of cephalic dorsum of short narrow to moderately broad spatulate hairs which are
decumbent and directed anteriorly. A row of longer spatulate hairs present on the false margin of the
clypeus which project forwards over the bases of the mandibles. Dorsum of head with a single pair of
standing stout hairs which are narrowly clavate apically, situated behind the highest point of the vertex.
Pronotum narrowly marginate anteriorly, not marginate laterally. Mesonotum in profile sloping posterior-
ly to the impressed metanotal groove, the propodeal dorsum shallowly convex and sloping posteriorly to
the propodeal teeth; these latter variable in size, usually triangular but sometimes reduced to small
rounded lobes. Infradental lamella usually narrow and inconspicuous, only rarely moderately broad but
sometimes vestigial. Sides of pronotum usually smooth but sometimes with sculpture, especially on the
upper portions. Mesopleuron, metapleuron or both punctate, sides of propodeum punctate. Frequently
the central area of the mesopleuron with sculpture reduced or absent, more rarely with the central area of
the metapleuron smooth. Sculpture of pronotal dorsum very variable, ranging from almost smooth to
moderately strongly sculptured. At one end of the range the pronotum is smooth except for a median
longitudinal carina, at the other a number of oblique to longitudinal fine striae or costulae are present and
the spaces between them may be finely punctulate. All grades between these extremes have been noted,
including a few sample where the pronotum is predominantly punctate. Frequently the striate component
is reduced leaving weak punctures as the predominant component, but in this case they are by no means as
strongly developed nor as conspicuous as the punctures on the mesonotum. Dorsal surface of mesonotum
and propodeum reticulate-punctate. Pronotum with humeral flagellate hairs. Mesonotum with a pair of
stout curved standing hairs which are clavate apically. Ground-pilosity of dorsal alitrunk of short curved
hairs which are sparse and subdecumbent to decumbent. With the pedicel segments in profile the
spongiform appendages reduced. Petiole with a narrow ventral strip and small lateral lobe. Postpetiole
with a moderately developed ventral lobe which is larger than the lateral spongiform appendage but
smaller, and usually obviously smaller, than the exposed area of the postpetiolar disc. In dorsal view the
petiole node reticulate to reticulate-punctate, with a narrow posterior spongiform strip. Postpetiole usually
344 BARRY BOLTON
reticulate to reticulate-punctate, rarely the sculpture reduced and superficial, bordered posteriorly by a
lamellate spongiform strip which abuts a similar narrow strip on the anterior margin of the first gastral
tergite. Basigastral costulae present, usually short. Petiole, postpetiole and gaster dorsally with stout
standing hairs which are clavate apically. Colour yellow to mid-brown.
One of the most successful members of the genus and of the Afrotropical dacetines as a whole,
ludovici ranges very widely over the whole continent and is also established in Madagascar and
Mauritius. It is closely related to simoni but most ludovici samples are instantly distinguishable
by the enlarged basal series of mandibular denticles in the latter. In those few samples where the
enlargement of the denticles is not marked the differentiating characters tabulated under simoni
will separate the two.
Apart from the diagnostic enlarged denticle series (Fig. 34) ludovici is separated from dotaja
by the presence in the latter of flagellate mesonotal hairs and simple gastral hairs; homsulumana
and maynei by those species' lack of cephalic flagellate hairs and strongly developed upper
scrobe margins. Usually ludovici is distinguished from lujae, serrula and concolor by their
blanketing reticulate-punctate pronotal sculpture, which generally is not seen in ludovici, but in
the few populations of the latter with a predominantly punctate pronotum the enlarged basal
mandibular denticles of ludovici will separate them. S. miccata shares the character of enlarged
basal denticles with ludovici but this minute species is very easily separated by the characters
noted in the discussion of the former.
In his revision of Serrastruma Brown (1952a) treated ludovici under the names alluaudi and
lotti, leaving ludovici as a species inquirenda as he had not been able to examine the type-series.
However, he speculated that ludovici might be the senior synonym of alluaudi and that alluaudi
and lotti may grade into one another. A few years ago he informed me that, having examined the
types of ludovici, he was convinced that the synonymy ludovici = alluaudi was in order, and
further that the characters which he had invoked to separate lotti from alluaudi (Brown, 1952a:
86) were indeed gradient, so that that species should also fall into the synonymy of ludovici. The
present investigation, utilizing much more material than was available to Brown, has served to
confirm all these findings.
Two other changes to the synonymies listed by Brown should be mentioned here. Firstly, the
name alluaudi st. nigeriensis was included by Brown in the synonymy of simoni, but a
re-examination of the type-series of nigeriensis shows them to have the characteristic dentition
of ludovici and the name is herewith transferred to the synonymy of this species. Secondly,
raymondi was described from a mixed series of ludovici and simoni originating in Mauritius,
where both are well established. As Brown (19520: 75) pointed out the 'type' (= holotype)
designated by Donisthorpe was referable to simoni. This is the only specimen in the entire series
bearing a type-label and there seems no reason to doubt that this was the specimen chosen as
holotype by Donisthorpe; thus raymondi is correctly referred to the synonymy of simoni, and
not to the synonymy of alluaudi (= ludovici) where it was left by Brown.
MATERIAL EXAMINED
Guinea-Bissau: Rio Cassine (L. Fed). Ivory Coast: Sassandra (/. Lobl); Man (/. Lobl); Man (V. Mahnert
& J.-L. Ferret); Abidjan, Banco Nat. Pk. (/. Lobl); Banco Forest (W. L. Brown); Issoneu (V. Mahnert &
J.-L. Ferret); Agboville, Yapo Forest (/. Lobl); Anguededou Forest Res. (W. L. Brown); Lamto (W. H.
Gotwald & R. Schaefer); Sangrobo (W. L. & D. E. Brown); Divo (L. Brader). Ghana: Mampong (D.
Leston);Tafo (D. Leston); Tafo (B. Bolton); Mt Atewa (C. A. Collingwood); Kade (W. H. Gotwald & R.
Schaefer). Togo: Palime, Klouto Forest (Vit). Nigeria: Olokemeji (F. Silvestri); Gambari (B. Bolton).
Cameroun: Korup (D. Jackson); Nkoemvon (D. Jackson); nr Yaounde (G. Terron). Gabon: Makokou
(W. H. Gotwald); Plateau d'Ipassa (/. A. Barra). Congo: Nkogo (L. Fed). Zaire: Irangi (E. S. Ross & R. E.
Leech); Ruwenzori, Mwenda (/. C. Bradley); Ituri Forest, Beni Irumu (N. A. Weber); Epulu (T. Gregg).
Sudan: Equatoria, Kagelu (N. A. Weber); Lotti Forest (N. A. Weber); Imatong Mts (N. A. Weber).
Uganda: Kampala (N. A. Weber). Tanzania: Amani (E. S. Ross & R. E. Leech); Tanga Cave Kulumuzi
(Ch. Alluaud). Angola: Duque de Braganca Falls (P. Hammond); Dundo, Carrisso Pk. (L. de Carvalho);
Saurimo (L. de Carvalho). Mozambique: Amatongas Forest (G. Arnold). Zimbabwe: Melsetter (R.
Mussard); Chirinda Forest (E. S. Ross & R. E. Leech). South Africa: Natal, Zululand, Eshowe (R. E.
Turner); Eshowe (G. Arnold); Gillitts (W. L. & D. W. Brown). Madagascar: Perinet (E. S. Ross); no loc.
(Sikora). Mauritius: Le Pouce Mt (R. Mamet); Le Pouce Mt (W. L. Brown); W. of Petria (W. L. Brown).
THE AFROTROPICAL DACETINE ANTS 345
Serrastruma lujae (Forel)
(Fig. 44)
Strumigenys lujae Forel, 1902: 294 (footnote), pi. 1, fig. 1. Syntype workers, MOZAMBIQUE: Zambesi,
Morrumballe (E. Lujd) (MHN) [examined].
Strumigenys reticulata Stitz, 1910: 141. Syntype workers, female, CAMEROUN: Bibundi (Tessmann)
(MNHU). [Synonymy by Brown, 19520: 78.]
Strumigenys (Cephaloxys) glanduscula Santschi, 1919; 88. Syntype workers, ZAIRE: Yambuya, 25. ii.,
no. 79 (Bequaert) (MRAC; MCZ) [examined]. [Synonymy by Brown, 19520: 78.]
Strumigenys (Xephaloxys) [sic] bequaerti Santschi, 1923: 286. Syntype workers, ZAIRE: Ruwenzori,
10.vii.1914 (Bequaert) (BMNH; MRAC; MCZ) [examined]. Syn. n.
Strumigenys (Cephaloxys) gerardi Santschi, 1923: 287. Syntype workers, ZAIRE: Katanga, R. Kasa,
Manyema, 1918 (Gerard) (MRAC) [examined]. [Synonymy by Brown, 19520: 78.]
Strumigenys (Cephaloxys) calypso Santschi, 1923: 288, fig. 4a. Holotype worker, TANZANIA: Ouha (Meyer)
(NMB) [examined]. Syn. n.
Strumigenys (Cephaloxys) aequalis Menozzi, 1942: 177. Syntype workers, EQUATORIAL GUINEA: Fernando
Po I., Concepcion (H. Eidmann) (NMB; MCZ) [examined]. [Synonymy by Brown, 19520: 78.]
Serrastruma lujae (Forel) Brown, 19520: 78.
Serrastruma bequaerti (Santschi) Brown, 19520: 80.
Serrastruma calypso (Santschi) Brown, 19520: 85. [Species inquirenda.]
WORKER. TL 2-2-3-3, HL 0-50-0-80, HW 0-42-0-62, CI 75-86, ML 0-18-0-32, MI 33-42, SL 0-32-0-58, SI
75-100, PW 0-30-0-66, AL 0-58-0-98 (55 measured).
Mandibular denticles regular, without a suddenly enlarged basal series but sometimes the denticular row
very evenly and gradually becoming slightly larger towards the base; the basalmost denticle frequently
larger than those preceding. Upper scrobe margins with a long simple or weakly flagellate hair projecting
laterally just behind the level of the eyes. Dorsum of head reticulate-punctate everywhere, the sculpture
fine and even. Ground-pilosity of dorsal head consisting of narrowly spatulate anteriorly curved short hairs
which are decumbent or closely applied to the surface. Standing pilosity of head consisting of a posterior
transverse row of 4 hairs situated close to the occipital margin and a more anteriorly situated pair just in
front of the highest point of the vertex. The standing hairs are usually simple and more or less cylindrical,
often pointed apically but sometimes very weakly swollen at their apices. Alitrunk in profile with the
promesonotum convex and sloping posteriorly to the impressed metanotal groove. Behind the metanotal
groove the propodeum is shallowly convex and sloping to the teeth posteriorly. Propodeal teeth very
variable in shape and size, varying from obtuse angles to strong triangular teeth. Infradental lamellae
present on the propodeal declivity but often narrow. Sides of alitrunk often reticulate-punctate everywhere
but with some variation in intensity. The sides of the pronotum may be only faintly sculptured and, in some
cases, the mesopleuron may be partially or entirely smooth. A central smooth patch may also occur on the
metapleuron. Dorsal alitrunk reticulate-punctate everywhere, the punctation on the pronotum usually fine
and very dense. Sometimes the intensity of the sculpture reduced so that the punctures are less well defined
than usual. Pronotum with humeral flagellate hairs present. Mesonotum with a single pair (or exceptionally
with 2 pairs) of stout standing hairs situated anteriorly on the sclerite. Ground-pilosity of dorsal alitrunk of
scattered finely spatulate hairs which are decumbent to appressed and inconspicuous. With the pedicel
segments in profile the spongiform appendages moderately developed. In dorsal view the petiole and
postpetiole both bounded posteriorly by a narrow lamellate spongiform strip, both usually punctate or
finely reticulate dorsally though on the postpetiole the sculpture may be almost effaced. Base of first gastral
tergite with a narrow lamellar strip and with basigastral costulae present. Petiole with one or more pairs of
standing stout hairs, the postpetiole with 2 or more pairs; first gastral tergite with numerous similar hairs.
Colour yellow to medium brown.
This very successful, common, widely distributed species shows a size range which is notably
greater than in any other Serrastruma. Brown (1952«: 79) has discussed the size variation and
concluded that only a single species is represented, and the present survey finds no argument
with that conclusion.
The closest relatives of lujae are serrula and concolor, the three together sharing the
characters of dense reticulate-punctate sculpture on the pronotal dorsum and lack of an enlarged
series of basal denticles on the mandible. Beside these features other easily observed characters
useful in separating lujae and' its immediate allies from the other species of the genus are as
follows. In dotaja the mesonotum is equipped with a single pair of extremely long flagellate hairs
346 BARRY BOLTON
and the gaster has only a single pair of simple hairs. In maynei, sulumana and geoterra the upper
scrobe margins lack projecting hairs. In simoni the spongiform appendages of the pedicel
segments are more massively developed (Figs 40, 44). In ludovici, beside the enlarged
mandibular denticles, the head has only a single pair of standing hairs, situated posteriorly. The
minute miccata has short mandibles, lacks projecting hairs on the upper scrobe margins and has
straight clavate hairs at the pronotal humeri.
S. serrula and concolor are both persistently small species, only overlapping the lower end of
the size range given for lujae. S. serrula is separated from lujae by having 3 or 4 pairs of
mesonotal standing hairs and elevated ground-pilosity on the head and alitrunk. S. concolor has
2 or 4 clavate standing hairs on the head (as opposed to 6 relatively fine hairs in lujae), and has
only a single pair of hairs on the postpetiole. Beside this most concolor specimens show a marked
indentation or impression of the upper scrobe margin at the site of origin of the projecting
flagellate hair.
The synonymy of lujae given by Brown (I952a: 78) is extended here to include the names
calypso and bequaerti. S. calypso was included by Brown as a species inquirenda as the holotype
of this form was not available for study at the time of his revision . An examination of the calypso
holotype places it firmly in the synonymy of lujae. S. bequaerti was treated as a rather doubtful
separate species by Brown (I952a: 80) who said This form may eventually prove to be a
montane subspecies or even a synonym of lujae.' He has since informed me that he is now
convinced that bequaerti and lujae grade together without possibility of any meaningful division
being made, and I fully concur with this opinion.
S. lujae nests in rotten wood and forages there and in the leaf litter layer. In a letter William L.
Brown has informed me that the prey are collembolans. Speaking of his observations in Banco
Forest, Ivory Coast, made in 1963, he says The 5. lujae nest was in the split end of a log and
consisted of many, perhaps a thousand or more, workers and copious brood. I found the nest by
following a single-file trail of workers along the top of the log, spaced out at intervals of 4—10 cm,
almost every one of which carried in its jaws a dead (or at least motionless) entomobryid
collembolan with furcula extended. The Collembola were about the same size as the ants
carrying them, or slightly larger. In the space of about 10 minutes I counted 40 springtails being
carried along the log on what appears to have been a trunk foraging trail. No other kinds of
insect prey were seen being carried or lying within the dissected nest.'
MATERIAL EXAMINED
Ivory Coast: Banco Forest (/. Lobl); Banco Forest (W. L. Brown); Man (/. Lobl); Man (V. Mahnert &
J.-L. Ferret); Yapo Forest, Agboville (/. Lobl); Anguededou Forest (W. L. Brown); Sangrobo (W. L. &
D. E. Brown). Ghana: Tafo (C. A. Collingwood); Tafo (B. Bolton); Aburi (D. Leston); Mamfe Scarp (D.
Leston); Bolgatanga (E. S. Ross & K. Lorenzen). Nigeria: Gambari (B. Bolton). Cameroun: Nkoemvon
(D. Jackson); nr Yaounde (G. Terron); Mt Cameroun (L. Fed); Mt Cameroun (B. Malkiri); Batanga (G.
Schwab). Gabon: Makokou (W. H. Gotwald); Plateau d'Ipassa (J. A. Barm). Zaire: Yambuya (J.
Bequaert); Ruwenzori (J. Bequaert); Ruwenzori (N. A. Weber); Burunga (J. Bequaert); Katanga,
Manyema (Gerard); Ituri Forest, Irumu (N. A. Weber); Ituri, Mt Hoyo (E. S. Ross & R. E. Leech);
Albertville (E. S. Ross & R. E. Leech); Thysville (E. A. Ross & R. E. Leech); Mt Kahuzi (E. S. Ross & R.
E. Leech); Lwiro R., Bukavu (E. S. Ross & R. E. Leech). Equatorial Guinea: Fernando Po I., Moka (L.
Fea); Fernando Po I. , Concepcion (H. Eidmann). Sao Tome L: Rib. , Palma (L. Fea); no loc. (Bl Malkiri).
Angola: Gubela (P. Hammond); Salazar (P. Hammond); Duque de Braganza Falls (P. Hammond);
Dundo (A. Machado); Dundo (L. de Carvalho); Tchimana R. (A. Machado). Sudan: Equatoria, Khor
Aba (N. A. Weber). Uganda: Ruwenzori, Mubuku (G. O. Evans); Ft Portal (N. A. Weber). Rwanda:
Rangiro (P. Werner). Burundi: Bujumbura (A. Dejean). Kenya: Embu, Irangi Forest Sta. (V. Mahnert &
J.-L. Perret); Mau Forest (F. Meneghetti); Kaimosi Mission (E. S. Ross & R. E. Leech); Tanzania: Ouha
(Meyer); Mt Meru (E. S. Ross & R. E. Leech); Amani (E. S. Ross & R. E. Leech); Uluguru Mts. Bunduki
(E. S. Ross & R. E. Leech). Mozambique: Morrumballe (E. Luja). Zimbabwe: Umtali, Melsetter (R.
Mussard); Vumba Mts (W. L. Brown). South Africa: Cape Prov., Port St John (R. E. Turner).
THE AFROTROPICAL DACETINE ANTS 347
Serrastruma maynei (Forel)
(Fig. 37)
Strumigenys (Trichoscapa) maynei Forel, 1916: 427. Syntype workers, female, male, ZAIRE: Stanleyville
(= Kisangani) (Kohl) (MHN; MRAC; MCZ) [examined].
Strumigenys (Trichoscapa) maynei var. latiuscula Forel, 1916: 428. Syntype workers, ZAIRE: Eala,
20.viii.1912 (R. Mayne) (MHN; MRAC; MCZ) [examined]. [Synonymy by Brown, 1952a: 77.]
Serrastruma maynei (Forel) Brown, 19520: 77.
WORKER. TL 2-3-3-0, HL 0-52-0-66, HW 0-44-0-54, CI 78-86, ML 0-18-0-24, MI 32-39, SL 0-34-0-42 SI
73-83, PW 0-31-0-41, AL 0-62-0-82 (45 measured).
Mandibular denticles regular, without a suddenly enlarged series of 4-8 denticles basally but often with
the basalmost denticle larger than those preceding. Upper scrobe margins strongly developed into broad
projecting flanges which run the length of the scrobe and do not peter out posteriorly, the margins without
flagellate hairs present. Clypeus sculptured, finely punctulate to granular, with short but broad spatulate
ground-pilosity and a row of longer spatulate hairs projecting forwards over the bases of the mandibles
from the false anterior clypeal margin. Dorsum of head strongly reticulate-punctate, with conspicuous
short but broad spatulate ground-pilosity. In profile the cephalic dorsum behind the highest point of the
vertex with 4-6 stout standing hairs which are narrowly to moderately clavate apically, the variation in
number occurring in single nest-series. Usually 4 such hairs are present, situated in a row in front of the
occipital margin; rarely a pair is present anterior to this row. Alitrunk in profile with promesonotum
shallowly convex and sloping more steeply posteriorly to the impressed metanotal groove. Propodeum
convex dorsally , on a lower level than the" promesonotum and terminating posteriorly in a pair of triangular
teeth which are subtended by conspicuous spongiform infradental lamellae. Sides of pronotum longitudi-
nally costulate, usually distinctly so but sometimes the costulae effaced in places or becoming weaker lower
down the sides; the spaces between the costulae smooth. Pleurae and sides of propodeum densely
reticulate-punctate. Pronotal dorsum longitudinally to obliquely strongly costulate, the spaces between the
costulae smooth; this sculpture usually weaker in small workers than in large. Remainder of dorsal alitrunk
strongly reticulate-punctate. Flagellate hairs usually absent from pronotal humeri (present in only a single
specimen of those examined). Mesonotum with a single pair of curved standing clavate hairs. Ground-
pilosity of dorsal alitrunk dense spatulate and very conspicuous. Spongiform appendages of pedicel
segments moderately developed in profile. The petiole with a ventral lamella and small lateral processes,
the postpetiole with a well developed ventral lobe and smaller lateral lobe. Petiole in dorsal view
reticulate-punctate, sometimes only superficially so, with a narrow transverse spongiform strip posteriorly.
Postpetiole dorsally smooth to superficially punctulate , sometimes with a suggestion of minute longitudinal
striae; with a narrow posterior spongiform strip which abuts a similar strip on the base of the first gastral
tergite. Basigastral costulae short. Dorsal surfaces of petiole, postpetiole and gaster with stout apically
clavate hairs. Colour dull brownish yellow to mid-brown.
S. maynei is characterized by its absence of flagellate hairs, generally strong costulate pronotal
sculpture, broad upper scrobe margins and conspicuous ground-pilosity; it is unlikely to be
confused with any other species. In distribution it appears to be restricted to the forest zones of
West and central Africa, and Uganda. Georges Terron's collection has yielded a couple of small
workers from Cameroun in which the pronotal sculpture is very reduced indeed. These may
represent a separate sibling species but I suspect that they are more likely to be members of the
first brood of a new colony.
The closest relative of maynei is geoterra , a species which shares the characters of broad upper
scrobe margins, lack of flagellate hairs and very broad conspicuous ground-pilosity. However, in
geoterra the entire alitrunk is glassy smooth and without trace of sculpture, the upper scrobe
margins are very broad and strikingly convex (compare Figs 36, 37), the cephalic ground-pilosity
is even coarser than in maynei, there is an obvious impression separating the pronotum and
mesonotum, the pronotal dorsal pilosity is strong and elevated, and the petiole node is smooth
dorsally.
Although often found nesting in rotten wood in the leaf litter layer maynei may also nest in rot
holes in trees some distance above the ground. In 1970 at Tafo in Ghana I observed a nest in a
cocoa branch about 1-7 m above the ground. One of the workers returning to this nest was
carrying a small nematoceran fly.
348 BARRY BOLTON
MATERIAL EXAMINED
Guinea-Bissau: Bolama (L. Fed). Ivory Coast: Lamto (W. L. & D. E. Brown). Ghana: Aburi (D.
Leston); Wiawso (D. Leston); Tafo (C. A. Collingwood);Tafo (B. Boltori). Nigeria: Gambari (B. Bolton).
Cameroun: nr Yaounde (G. Terron). Equatorial Guinea: Annobon I. (Cambridge Univ. Expd.). Zaire:
Kisangani (Kohl); Eala (R. Mayne); Ruwenzori (/. C. Bradley). Angola: Dundo (A. Machado). Uganda:
Maragambo Forest (E. S. Ross & R. E. Leech). Tanzania: Lk. Manyara (E. S. Ross & R. E. Leech).
Serrastruma miccata sp. n.
(Fig. 39)
HOLOTYPE WORKER. TL 1-7, HL 0-46, HW 0-32, CI 70, ML 0- 12, MI 26, SL 0-37, SI 116, PW 0-24, AL 0-46.
Basal 4-5 denticles on mandibular masticatory margin suddenly and conspicuously enlarged, distinctly
much coarser and broader than those preceding. Head narrow, antennal scapes long (CI and SI, above).
Upper scrobe margins very feebly developed, merely an edge without a projecting lamina; flagellate hairs
absent from upper scrobe margins. Clypeus finely punctulate, with curved narrowly spatulate small hairs
and with an anteriorly projecting row of such hairs on the false anterior clypeal margin. Ground-pilosity of
cephalic dorsum of minute anteriorly curved inconspicuous hairs which are narrowly spatulate and
subdecumbent, those situated posteriorly tending to be splayed or forked at the apex. Dorsum of head
without specialized standing longer hairs which are differentiated from the ground-pilosity. Dorsum of
head densely reticulate-punctate everywhere. Pronotum narrowly marginate anteriorly, not marginate
laterally. In dorsal view the pronotum and mesonotum separated by a shallow faint impression, the
mesonotum and propodeum separated by a narrow fine transverse line. In profile the mesonotum weakly
raised above the level of the pronotum and sloping shallowly downwards posteriorly. Metanotal groove a
narrowly incised line, not impressed. Propodeum continuing the slope of the mesonotum posteriorly and
ending in a minute triangular tooth (left tooth broken). Infradental lamellae absent. Pleurae mostly
smooth, with some peripheral punctulae. Sides of pronotum with very feeble vestiges of sculpture.
Pronotal dorsum with vestigial superficial reticulation, the mesonotum and propodeum finely punctulate.
Flagellate hairs absent from alitrunk, the pronotal humeri with a pair of short straight hairs which are quite
stout and clavate apically. Dorsal surfaces of pronotum and mesonotum with dense short ground-pilosity
consisting of anteriorly or medially curved elevated hairs, those on the mesonotum appearing clavate in
profile. Spongiform appendages of pedicel segments in profile very reduced, the ventral petiolar appen-
dage represented only by a minute crest. Ventral lobe of postpetiole very small and lateral process reduced
to a thin strip. Petiole node in dorsal view slightly longer than broad, superficially punctulate and with a
minute transverse crest on the posterior border which represents the last vestige of the spongiform strip.
Postpetiole in dorsal view marginally longer than broad and with its posterior margin sharply indented
medially. Spongiform material absent laterally but posteriorly with a narrow lamelliform strip which abuts
a similar narrow strip on the base of the first gastral tergite. Postpetiole punctulate-granular and the first
gastral tergite with extremely fine short basal costulae. Dorsal surfaces of petiole, postpetiole and gaster
with numerous short apically clavate hairs. Colour yellow.
Holotype worker, Ghana: Mampong, 26.U970 (P. M. Room) (BMNH).
Although sharing with ludovici the character of an enlarged basal series of mandibular denticles,
I suspect that miccata has acquired it independently as otherwise the two share very few
diagnostic characters. In fact miccata is remote from all the known species of Serrastruma on a
number of counts. Most obvious of these is the differently shaped alitrunk (compare Figs 39-44).
Whereas in all other species the posterior portion of the mesonotum slopes to an impressed
metanotal groove and the promesonotum forms a surface on a higher level than the propodeum,
in miccata the mesonotum and propodeum form a more or less uniform slope, the metanotal
groove is not impressed and the promesonotum is not at a higher level than the propodeum. In
miccata the petiole and postpetiole in dorsal view are fractionally longer than broad, whereas in
all other species they are broader, in the case of the postpetiole much broader, than long. The
mandibles of miccata are short but the scapes are long, a combination not found elsewhere in
Serrastruma, and the lack of elongate specialized hairs on the head and dorsal alitrunk in miccata
is not repeated elsewhere in the genus, where at least a single cephalic and a single mesonotal
pair occur. These characters, along with the unique development of straight clavate hairs at the
pronotal humeri in miccata in place of the more usual flagellate hairs, and the small size of the
species, render it immediately recognizable.
THE AFROTROPICAL DACETINE ANTS 349
Serrastruma serrula (Santschi)
(Fig. 42)
Strumigenys lujae var. serrula Santschi, 191(k: 390. Holotype worker, CONGO: Brazzaville (A Weiss)
(NMB) [examined].
Strumigenys serrula Santschi; Santschi, 19106: 361. [Raised to species.]
Strumigenys (Cephaloxys) uelensis Santschi, 1923: 289, fig. 4b. Syntype workers, ZAIRE: Haul Uele,
Watsa, xi.1919 (L. Burgeon) (MRAC) [examined]. [Synonymy by Brown, 1952a: 81.]
Serrastruma serrula (Santschi) Brown, I952a: 81.
WORKER. TL 1 -9-2-3, HL 0-44-0-54, HW 0-34-0-44, CI 75-88, ML 0- 15-0-18, MI 32-37, SL 0-26-0-33 SI
65-82, PW 0-24-0-30, AL 0-48-0-56 (50 measured).
Mandibular denticles evenly sized to the base or minutely and very gradually increasing in size basally,
the basalmost denticle usually enlarged but never with a series of 4-8 obviously enlarged basal denticles.
Upper scrobe margins regular or with a shallow impression at the site of the flagellate hair. Ground-pilosity
of head narrowly spatulate, quite dense and very conspicuous, curved anteriorly and elevated, not closely
applied to the surface. Dorsum of head with an occipital transverse row of 4 standing longer hairs which are
usually cylindrical and tapered apically, only very rarely with their apices slightly swollen. Commonly a
more anteriorly situated pair of similar hairs is present , j ust in front of the highest point of the vertex. All of
these standing hairs are only slightly longer and stouter than the curved hairs of the ground-pilosity. Entire
dorsum of head sharply and strongly reticulate-punctate. Dorsal alitrunk with the convex promesonotum
sloping posteriorly to the impressed metanotal groove. Propodeal dorsum convex and sloping down to the
teeth, the latter usually acutely triangular but variable in size; infradental lamellae present down the sides
of the propodeal declivity. Sides of pronotum and the pleurae punctate, the punctures on the mesopleuron
often more superficial and more widely spaced than elsewhere; infrequently the punctures superficial
everywhere on the sides. Dorsal alitrunk strongly reticulate-punctate everywhere, the punctures sharply
defined and the pronotum often with feeble longitudinal rugulae or striae which when present are very
obviously secondary to the punctate component. Pronotum with elongate flagellate hairs at the humeri.
Mesonotum with 3 (rarely 4) pairs of standing hairs; these are relatively slender, at most only feebly
expanded apically. The posteriormost of these hairs is situated at or very close to the metanotal groove and
is very variable in size. In some samples it is almost as long as the preceding mesonotal hairs but frequently
is only as long as the ground-pilosity; whatever its length it is always directed posteriorly. Ground-pilosity
on promesonotum quite long and conspicuous, dense and elevated, not closely applied to the surface. In
profile the spongiform appendages of the pedicel segments moderately developed, the ventral appendage
of the petiole may be reduced to a narrow ridge but is usually spongiform. Lateral and ventral spongiform
lobes of postpetiole present. In dorsal view the surfaces of both the petiole and postpetiole reticulate to
reticulate-punctate, sometimes the sculpture superficial and faint. Posterior spongiform strips of both
segments narrow as is the basal strip on the first gastral tergite. Basigastral costulae present but frequently
short and widely spaced, the gaster otherwise unsculptured. Petiole and postpetiole always with more than
one pair of standing hairs, the gaster with numerous hairs. These vary from almost cylindrical to very
weakly expanded apically, not strongly clavate. Colour dull yellow to yellowish brown.
This widespread persistently small species is related to concolor and lujae. Together the three
are characterized by their lack of an enlarged basal series of mandibular denticles, presence of
cephalic flagellate hairs and dense reticulate-punctate pronotal sculpture. The differences
separating serrula and concolor are tabulated under the latter name.
5. serrula is separated from lujae by its size and pilosity. The largest specimens of serrula only
overlap the very smallest individuals of lujae (serrula HW 0-34-0-44, SL 0-26-0-33; lujae HW
0-42-0-62, SL 0-32-0-58). Ground-pilosity everywhere on the head and promesonotum is
relatively short, inconspicuous and closely applied to the surface in lujae. This gives the ant a
rather smooth appearance and emphasises the long specialized hairs which stand out very
conspicuously. In serrula the ground-pilosity is quite long and very distinctive, being markedly
elevated from the surface so that the long specialized hairs which project from the ground-
pilosity are by no means as distinctive in appearance. On the mesonotum long hairs are
restricted to a single anteriorly placed pair in lujae (or very exceptionally a second pair may be
present, sited very close to the first) whereas in serrula three pairs are generally present
distributed along the length of the mesonotum and with the posteriormost pair at or very close to
the metanotal groove. Finally the reticulate-punctate pronotal sculpture is usually more strongly
350 BARRY BOLTON
developed and more sharply defined in serrula than in lujae and serrula frequently has
superimposed fine longitudinal striae or rugulae on the punctate surface.
5. serrula nests in pieces of rotting wood embedded in the leaf litter and topsoil layers, and
preys on the isotomid collembolan Folsomia Candida Willem. Its predatory behaviour has been
investigated in some detail by Dejean (19800; 1980ft).
MATERIAL EXAMINED
Ivory Coast: Tai Forest (V. Mahnert&J. -L. Ferret); Bingerville (V. Mahnert&J.-L. Ferret); Lamto (J.
Levieux); Anguededou (W. L. Brown); Banco Forest (W. L. Brown); Divo (L. Brader). Ghana:
Mampong (P. Room); Tafo (B. Bolton). Cameroun: Nkoemvon (D. Jackson); nr Yaounde (G. Tenon).
Gabon: Makokou (/. Lieberburg); Makokou (W. H, Gotwald). Chad: Haut Mbomu (N. A. Weber). Sao
Tome I.: Rib. Palma (L. Fed); Vista Algere (L. Fed); no loc. (/. Derron). Congo: Brazzaville (A. Weiss).
Zaire: Ituri Forest, Irumi (N. A. Weber); Niangara (N. A. Weber); Hout Uele, Watsa (L. Burgeon);
Tchikapa (A. Machado). Angola: Salazar (P. Hammond); Duque de Braganca Falls (P. Hammond);
Gubela (P. Hammond); Dundo (L. de Carvalho); Camissombo (A. Machado). Sudan: Equatoria, Kagelu
(N. A. Weber). Burundi: Bujumbura (A. Dejean); Imbo Plain (A. Dejean); Bugarama (A. dejean).
Serrastruma simoni (Emery)
(Fig. 40)
Strumigenys simoni Emery, 18950: 42, pi. 2, fig. 21. Holotype worker, SOUTH AFRICA: Transvaal,
Makapan, 1893 (£. Simon) (MCSN) [examined].
Strumigenys escherichi Forel, 1910: 261. Syntype workers, ETHIOPIA: Eritrea, Ghinda (K. Escherich)
(MHN) [examined]. [Synonymy by Brown, 19520: 82.]
Strumigenys cognata Santschi, 1910ft: 362. Syntype workers, female, ANGOLA: Benguela, Cucala, 1910 (J.
Cruchet) (NMB; MR AC) [examined]. [Synonymy by Brown, 19520: 82.]
Strumigenys biconvexa Santschi, 19130: 258. Syntype workers, KENYA: Cheteni, xi.1911, st. 4 (Alluaud &
Jeannel) (NMB) [examined]. [Synonymy by Brown, 19520: 83.]
Strumigenys cognata st. boerorum Santschi, 19130: 259. Syntype workers, SOUTH AFRICA: Zululand,
Dukuduku (/. Trdgdrdh) (NMB) [examined]. [Synonymy by Brown, 19520: 83.]
Strumigenys escherichi race limbata Forel, 1913c: 222. Syntype workers, ZIMBABWE: Bulawayo (G.
Arnold) (MHN) [examined]. [Synonymy by Brown, 19520: 83.]
Strumigenys escherichi var. cliens Forel, 1913d: 317. Syntype workers, ZAIRE: Katanga, Elizabethville,
1912 (Bequaert) (MHN; MRAC; MCZ) [examined]. [Synonymy by Brown, 19520: 83.]
Strumigenys (Trichoscapa) escherichi st. cognata var. obscuriventris Santschi, 1914ft: 376. Syntype
workers, NIGERIA: Olokemeji (F. Silvestri) (NMB) [examined]. [Unavailable name.]
Strumigenys (Trichoscapa) escherichi st. cognata var. fusciventris Santschi, 1915: 261. [Unnecessary
replacement name for obscuriventris Santschi.] [Unavailable name.]
Strumigenys (Cephaloxys) raymondi Donisthorpe, 1945: 779. Holotype worker, MAURITIUS: Corps de
Garde Mt, 17. i. 1944, no. 20 (R. Mamet) (BMNH) [examined]. [Wrongly synonymized with alluaudi by
Brown, 1952a: 75. S. raymondi series is a mixture of ludovici (= alluaudi) and simoni, but holotype of
raymondi belongs to the latter species.] Syn. n.
Serrastruma simoni (Emery) Brown, 19520: 82.
WORKER. TL 2-4-3-0, HL 0-54-0-64, HW 0-44-0-52, CI 74-85, ML 0-20-0-22. MI 33-39, SL 0-34-0-41, SI
76-86, PW 0-28-0-36, AL 0-64-0-75 (60 measured).
Mandibular denticles small even and very regular, without an enlarged basal series but often with the
basalmost denticle somewhat enlarged. Upper scrobe margins conspicuous, forming a punctulate or
granular flange which extends back to the apex of the scrobe before petering out, the margins each with a
single long flagellate hair arising just behind the level of the eye and projecting laterally. Clypeus
sculptured on the disc, finely punctulate or granular, or sometimes feebly striate, with sparse spatulate
appressed hairs. False anterior clypeal margin with a row of spatulate hairs which project forwards over the
mandibular bases. Ground-pilosity of head of scattered but distinctive anteriorly directed spatulate hairs
which are decumbent to appressed. Dorsum of head behind highest point of vertex with two pairs of stout
standing hairs which are anteriorly curved and apically clavate. Dorsum of head densely reticulate-
punctate. Pronotum not marginate laterally, the pronotum and anterior portion of mesonotum convex and
on a higher level than the propodeum, the posterior portion of the mesonotum sloping steeply down to the
metanotal groove which is broadly and shallowly impressed. Propodeal dorsum shallowly convex, the teeth
usually broad and strongly developed, with a conspicuous infradental lamella. Alitrunk pleurae smooth,
THE AFROTROPICAL DACETINE ANTS
351
devoid of sculpture or at most with punctulate vestiges peripherally. Sides of pronotum usually smooth but
sometimes with one or two striae which extend onto the sides from the dorsum. Sculpture of pronotal
dorsum variable, ranging from almost smooth with only vestiges of sculpture to quite strongly sculptured.
Commonly the sculpture consists of few to several longitudinal to oblique fine costulae or striae on an
almost or quite smooth surface. Variation away from this occurs either by reduction in number and
intensity of the costulae or striae until the dorsum is almost unsculptured, or by intensification of the
costulae or striae, or by the appearance of fine punctulation between them. However, in samples where
punctulate sculpture occurs it is always very obviously secondary to the costulate or striate component, and
in samples where the pronotum is smooth a median fine longitudinal carinula usually remains. Dorsal
surfaces of mesonotum and propodeum finely and densely reticulate-punctate, as is the upper portion of
the propodeal declivity between the teeth. Pronotum equipped at the humeri with a pair of long fine
flagellate hairs, the mesonotum with a pair of stout curved hairs which which are clavate apically.
Ground-pilosity of promesonotum consisting of a few scattered narrowly spatulate hairs which are
appressed or nearly so. Spongiform appendages of pedicel segments well developed in profile, the
subpetiolar appendage running the length of the segment; the ventral and lateral spongiform lobes of the
postpetiole large, the former usually somewhat larger than the latter. Petiole node in dorsal view finely
reticulate-punctate, the sculpture sometimes faint, broader than long and with a conspicuous posterior
collar of spongiform material. Postpetiole smooth dorsally, with spongiform material posterolaterally and
with a broad posterior strip. Base of first gastral tergite bordered with a lamellate strip upon which the
basigastral costulae arise and run posteriorly for a short distance on the tergite proper. Petiole and
postpetiole each with a single pair of stout hairs which are clavate apically, the first gastral tergite with 1-8
pairs of such hairs. Colour ranging from yellowish brown to blackish brown.
A very successful and widely distributed species, simoni ranges throughout eastern and southern
Africa. It has also been found in Nigeria but otherwise there are no records of its presence in
West or central Africa, so this sample may represent an introduction or a mislabelled series. It is
also known from Mauritius where it was most probably introduced by man.
S. simoni is separated from dotaja by the characters tabulated under the latter name. It is
quickly distinguished from lujae and the related concolor and serrula by the presence in these
three of dense blanketing reticulate-punctate sculpture on the promesonotum, and it is
separated from maynei, geoterra and sulumana by those species' lack of cephalic flagellate hairs,
which are very conspicuous in simoni. The minute miccata is readily separated, not only by its
enlarged basal series of mandibular denticles and small size, but also by its possession of short
clavate hairs at the pronotal humeri and lack of a metanotal impression. In simoni the long
humeral flagellate hairs are obvious and the metanotal impression broad. Most samples of
ludovici are easily distinguished from simoni by the presence in the former of a series of 4-8
enlarged denticles basally on the mandibular masticatory margin. However, in some ludovici
series the denticles are not nearly so sharply defined as is usual, and these may be more difficult
to tell apart. The following contrasting characters serve to separate the two, the first character
noted is usually sufficient alone.
simoni
Mandibular denticles small fine and
regular to base, the basalmost
may be enlarged.
Dorsum of head behind highest point
of vertex with two pairs of standing
curved clavate hairs.
Upper scrobe margins in full-face view
forming a conspicuous flange which
runs to the apex of the scrobe.
Mesopleuron and metapleuron smooth
except for peripheral weak sculpture.
With postpetiole in profile the area
of the ventral spongiform lobe
exceeding that of the exposed area
of the postpetiolar disc (Fig. 40).
ludovici
Mandibular denticles with basal
4-8 enlarged.
Dorsum of head behind highest point
of vertex with a single pair of
standing curved clavate hairs.
Upper scrobe margins in full-face view
forming a narrow weak rim which
rapidly peters out posteriorly.
Metapleuron and usually also mesopleuron
with reticular or punctate sculpture.
With postpetiole in profile the area
of the ventral spongiform lobe usually
much less than that of the exposed
area of the postpetiolar disc; only
rarely the two subequal (Fig. 41).
352
simoni - cont.
Infradental lamellae on propodeum
broad and conspicuous.
Ranges of indices. SI 76-86, CI 74-85,
MI 33-39 (ML = 0-42-0-48 xHW).
BARRY BOLTON
ludovici - cont.
Infradental lamellae on propodeum
usually narrow or vestigial.
Range of indices, SI 86-115, CI 71-80,
MI 35-50 (ML = 0-45-0-68 xHW).
The extensive synonymy of simoni is basically as Brown (19520: 82-83) left it except for a
couple of minor modifications. The name alluaudi st. nigeriensis, formerly included in the
synonymy of simoni, has been transferred to ludovici as its type-series shows the characteristic
dentition of that species. 5. raymondi has been brought into the synonymy of simoni from that of
ludovici. The original series of raymondi consisted of a mixture of both species but the holotype
is a very oridinary specimen of simoni.
MATERIAL EXAMINED
Nigeria: Olokemeji (F. Silvestri). Ethiopia: Eritrea, Ghinda (K. Escherich). Kenya: Cheteni (Alluaud &
Jeannel). Burundi: Imbo Plain (A. Dejean). Uganda: Entebbe (G. Arnold). Zaire: Shaba, Lubumbashi
(Bequaert). Malawi: Lk. Nyasa, Urundi (E. S. Ross & R. E. Leech); Mzimba (E. S. Ross & R. E. Leech).
Zambia: Kasama (E. S. Ross & R. E. Leech). Mozambique: Amatongas Forest (G. Arnold). Zimbabwe:
Bulawayo (G. Arnold); Sawmills (G. Arnold); Victoria Falls (G. Arnold); Victoria Falls (W. L. Brown).
Matopos (G. Arnold); Harare, Chishawasha (A. Watsham). Angola: Dundo (A. Machado); Benguela,
Cucala (Cruchet). Botswana: Okavango, Maxwee (A. Russell-Smith). South Africa: Transvaal, Makapan
(E. Simon); Acornhoek (Tucker); Natal, Dukuduku (/. Tragdrdh); Dukuduku (W. L. & D. E. Brown);
Durban (H. B. Marley); Hlabisa (J. C. Faure); Richards Bay (/. C. Faure); Sordwana (/. C. Faure).
Mauritius: Corps de Garde Mt (R. Mamet).
Serrastruma sulumana sp. n.
HOLOTYPE WORKER. TL2-0, HLO-50, HWO-33, CI66, ML 0-22, MI 44, SLO-40, SI 121, PW 0-23, ALO-58.
Small narrow-headed species with relatively long mandibles and very long scapes. Mandibles slender,
serially finely denticulate, the apical tooth subspiniform and the denticles becoming gradually slightly
larger towards the base. Rounded angle separating masticatory and jasal margins of mandible surmounted
by a thin translucent crest which follows the curve and represents the remains of the basal lamella. Anterior
clypeal margin translucent and convex, overhung by the false clypeal margin which is equipped with
anteriorly projecting hairs. Antennal scrobes vestigial, the dorsum rounding into the sides except in front
of the level of the eye where a feebly angular upper scrobe margin remains. The eyes freely visible in
full-face view on the ventrolateral margins of the head because of the disappearance of the upper scrobe
margins which partially or wholly obscure them in most other members of the genus. Occipital corners
evenly rounded, the occipital margin shallowly concave. Antennal scapes very long and slender, subcylin-
drical, slightly increasing in thickness from base to apex, the leading edges only with fine curved hairs,
without bizarre projecting pilosity. Ground-pilosity of head of inconspicuous curved narrowly spatulate
hairs. A transverse row of 4 stouter more erect hairs present paralleling the occipital margin but the head
without flagellate hairs. Dorsum of head finely and densely reticulate-punctate. Pronotum not marginate
laterally, the humeri broadly rounded and each with a long fine flagellate hair. In profile the posterior half
of the mesonotum descending steeply to the broadly impressed metanotal groove. Propodeal dorsum
elevated behind the level of the metanotal groove then sloping shallowly backwards. Propodeal teeth
elevated, long and narrowly triangular, without an infradental lamella. Sides and dorsum of alitrunk evenly
and densely reticulate-punctulate everywhere. Dorsal alitrunk with inconspicuous curved narrowly
spatulate pilosity, the mesonotum without standing specialized hairs such as are usually seen in Serrastru-
ma species. Petiole and postpetiole reticulate-punctate everywhere, the gaster smooth and with vestigial
basal costulae. Spongiform appendages of pedicel segments very reduced. In profile the subpetiolar
appendage reduced to a narrow translucent strip and the subpostpetiolar lobe represented only by a thin
laminar fringe around the curve of the sternite. In dorsal view the petiole and postpetiole each with minute
vestiges of their respective transverse posterior spongiform strips, the best developed section being at the
posterolateral angles of the postpetiole. Petiole, postpetiole and gaster dorsally with a number of erect to
suberect quite stout hairs which are thickened apically. Colour pale yellow.
PARATYPE WORKER. TL not measurable as pedicel segments and gaster missing, HL 0-56, HW 0-36, CI 64,
ML 0-25, MI 45, SL 0-46, SI 127, PW 0-24, AL 0-62. As holotype.
Holotype worker, Cameroon : nr Yaounde, sample SK (G. Terron) (ENSA).
Paratype. 1 worker with same data as holotype (BMNH).
THE AFROTROPICAL DACETINE ANTS 353
A very distinctive small species of Serrastruma immediately separated from all its congeners by
its combination of narrow head with relatively long mandibles and extremely long scapes,
reduced upper scrobe margins which lack flagellate hairs, inconspicuous cephalic ground-
pilosity, presence of humeral flagellate hairs but lack of specialized large hairs on the mesono-
tum, and vestigial spongiform tissue on the pedicel segments.
CLADAROGENYS Brown
(Fig. 45)
Cladarogenys Brown, 1976: 33. Type-species: Cladarogenys lasia Brown, 1976: 34, figs. 1-5, A-D, by
original designation.
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles elongate and narrow (MI 46), produced into
long narrow projecting blades which taper apically, lacking an apical fork of spiniform teeth. Instead the
mandible is equipped apically with a crowded series of small denticles and the distal two-thirds of the blade
has irregularly spaced minute denticles. Proximal one-third of mandible edentate and lacking a differenti-
ated basal lamella. Antennae with 6 segments, the scapes cylindrical. Orbicular hairs absent.
The single species included in this genus, C. lasia, is certainly a direct derivative of Serrastruma,
separated only by its more specialized mandibles which, whilst appearing longer in Cladar-
ogenys, are in fact within the known range of mandible relative length of Serrastruma (MI
26-50), and only seem longer because they are narrower. Basically the mandibles are the same
as in Serrastruma, with a rounded angle separating the basal and masticatory margins and
lacking a differentiated basal lamella, but, whereas the continuous rows of denticles on the
masticatory margins are opposable in Serrastruma throughout the length of the blade, in
Cladarogenys the narrowing of the blades has meant that the mandibles are only properly
opposable at the extreme apex and for most of their length are separated by a gap. This has lead
to, or has been accompanied by, a reduction in the denticle row down the length of the blades
which probably indicates that these areas are no longer used in prey seizure, this function having
devolved upon the apical clump of denticles alone.
The remainder of the head and the form of the body in general agree so closely with
Serrastruma that I doubt whether a separate genus is really necessary for C. lasia, and I strongly
suspect that a revision of the short-mandibulate dacetine genera on a world wide basis will see
the synonymy of Cladarogenys under Serrastruma. Known only from the holotype collected in
Gabon, C. lasia has been well described and profusely illustrated by Brown (1976). The
following notes (and Fig. 45) are abstracted from that original description.
Cladarogenys lasia Brown
(Fig. 45)
Cladarogenys lasia Brown, 1976: 34, figs 1-5, A-D. Holotype worker, GABON: nr Makokou, Laboratoire
de Primatologie et d'Ecologie Equatoriale, ix.-xii.1972, berlesate of rain forest litter and humus (/.
Lieberburg) (MCZ).
WORKER. TL 3-2, HL 0-70, HW 0-52, CI 74, ML 0-32, MI 46, SL 0-55, SI 106, AL 0-84.
Mandibles as noted under the generic diagnosis and Fig. 45. Anterior clypeal margin prominent medially
but not overlapping the bases of the mandibles. Posterodorsal margins of head forming a raised rim which
extends to the posterolateral margin, expanded into a small flat truncated tubercle on each side and with a
similar but smaller tubercle in front of this, just dorsal to the eye on each side. Dorsum of head densely
clothed with simple fine hairs and with a long flagellate hair arising from the tubercle above the eye on each
side. Dorsum of head irregularly rugulose, the spaces between the rugulae reticulate-punctate; clypeal
dorsum reticulate-punctate. Posterodorsal corners of vertex with a sulcus which parallels the rim and is
composed of 3-4 partially confluent foveae with concave shining bottoms. Pronotal humeri bluntly
tuberculate and equipped with a pair of long flagellate hairs. Metanotal groove impressed. Pronotal
dorsum shining, with quite dense shallow foveolae, the remainder of the dorsal alitrunk reticulate-punctate
with superimposed rugulae, some of them enclosing irregular pits. Sides of pronotum finely punctate,
mesopleuron finely punctulate and remainder of pleuron smooth; sides of propodeum reticulate-punctate.
Petiole and postpetiole punctulate-rugulose. Gaster smooth with conspicuous basal costulae. Entire
354 BARRY BOLTON
dorsum of body densely clothed with fine simple pilosity, and with paired long flagellate hairs on
mesonotum and petiole. Postpetiole with several and gaster with more than 30 flagellate hairs.
EPITRITUS Emery
(Figs 46-48)
Epitritus Emery, 1869a: 136. Type-species: Epitritus argiolus Emery, 1869a: 136, fig. 1, by monotypy.
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles elongate and linear (MI 31-48), produced
into long narrow projecting blades but lacking an apical fork of spiniform teeth. Instead the mandibular
apex is equipped with either a more or less vertical array of denticles or with a single apicodorsal spiniform
tooth subtended by a row of denticles. Preapical teeth or denticles present or absent. Antennae 4 or 6
segmented. In all Afrotropical species the scape is strongly bent backwards and has a large anteriorly
projecting subbasal lobe at the bend. Orbicular hairs present on the head. Labral lobes elongate-conical
and visible in full-face view in the space enclosed by the mandibular blades.
In its modern restricted sense Epitritus, as redefined by Brown (19496), contains only seven
species restricted to the Old World. Bolton (1972) presented a key dealing with all the species
known to that date. Prior to Brown's redefinition a number of unrelated species were associated
under Epitritus but these were dispersed to various other genera by Brown (1948; 19496), who
also showed (Brown, 1958) that the antennae in this genus may have 4 or 6 segments (until then
the known species had 4-merous antennae). Two Afrotropical species originally described in
Epitritus, marginatus and mandibularis , which were retained there by Arnold (1917) and
Wheeler (1922), were placed in a separate genus, Miccostruma, by Brown (1948), a generic
name now synonymized under Smithistruma.
Apart from the four West African species dealt with here the remaining three show a wide
distribution, with argiolus Emery from a number of Mediterranean lands, hexamerus Brown
from Japan, and murphy i Taylor from West Malaysia and Sarawak. An undescribed species
from Nepal is also known.
The origins of Epitritus appear to lie among the short-mandibulate dacetines related to
Smithistruma rather than with the long-mandibulate allies of Strumigenys. Brown (1958) first
postulated that Epitritus was a long-mandibulate genus independently derived from an ancestral
short-mandibulate Smithistruma-like stock, and the later description of Dysedrognathus by
Taylor (19686) produced a plausible intermediate stage which strengthened the hypothesis.
List of Afrotropical Epitritus
laticeps Brown room! Bolton
minimus Bolton tiglath sp . n .
Key to species (workers)
1 Antennae with 4 segments. Postpetiole in profile with a large spongiform ventral lobe which is as
deep as the height of the postpetiolar node. Minute species, HW 0-29-0-31 (Fig. 46).
(Ghana, Cameroun) minimus (p. 355)
- Antennae with 6 segments. Postpetiole in profile without or with a vestigial ventral lobe. Larger
species, HW>0-35 2
2 Tooth at dorsal apex of mandible long and spiniform, strongly crossing over the tooth from the
opposite mandible when the blades are closed; MI 35-38, HW >0-50. Area of head
immediately behind the posterior clypeal margin without orbicular hairs (Fig. 47). (Ghana,
Ivory Coast, Cameroun) room! (p. 356)
- Tooth at dorsal apex of mandible short, not spiniform, not crossing over the tooth from the
opposite mandible when the blades are closed; MI 35^48 but if MI <40 then HW only about
0-39. Orbicular hairs on head present immediately behind the posterior clypeal margin 3
3 Dorsal surface of each mandibular blade with two large flattened hairs arising on the distal half.
Posterior half of mesonotum not humped in profile. Minute species with shorter mandibles,
HW 0-39, MI 35. (Ivory Coast) tiglath (p. 357)
- Dorsal surface of each mandibular blade without flattened hairs arising from them. Posterior
half of mesonotum distinctly humped in profile. Larger species with longer mandibles, HW
>0-45, MI 42-48. (Ivory Coast, Nigeria, Cameroun) laticeps (p. 355)
THE AFROTROPICAL DACETINE ANTS
Epitritus laticeps Brown
(Fig. 48)
355
Epitritus laticeps Brown, 1962«: 77, figs 1-4. Holotype and paratype workers, NIGERIA: nr Zungeru on the
Kaduna road, 19.xii.1956, base of dead tree, S780 (IV. A. Sands) (BMNH; MCZ; USNM) [examined].
WORKER. TL 1-7-2-2, HL 0-40-0-48, HW 0-46-0-59, CI 115-125, ML 0-18-0-22, MI 42-48, SL 0-22-0-29,
SI 47-50, PW 0-26-0-32, AL 0-41-0-50 (12 measured).
Mandibular blades with a single preapical denticle, situated close to the apex. Apical armament of
mandible consisting of a series of 7-8 denticles, without a conspicuously elongate spiniform tooth at the
dorsal end of the series. Dorsal surfaces of mandibular blades naked, without large flattened hairs arising
from them. Anterior clypeal margin with 4 long strap-like spatulate hairs which project forwards, the outer
pair longer than the inner; and also with a short spatulate hair close to the inner base of each mandibular
blade. Clypeus shining, with numerous very small appressed spatulate hairs. Remainder of head finely
densely reticulate-punctate and equipped with orbicular hairs which occur from the posterior clypeal
margin to the occiput; without a large space behind the clypeus which is free of orbicular hairs. Antennae
6-segmented, the scapes strongly back-curved and with a large anteriorly directed subbasal lobe at the
bend, the leading edge with a row of projecting large flattened to spoon-shaped hairs. Alitrunk in profile
with the mesonotum strongly humped posteriorly, the highest point of the outline being at or just behind
the mesonotal midlength. Anteriorly the mesonotum slopes down to its junction with the pronotum and
posteriorly it joins the still more steeply sloped propodeal dorsum. Propodeal declivity bordered by a
broad lamella, usually without teeth but rarely an angular tooth is developed, projecting posteriorly from
the dorsal end of the lamella. Sides of alitrunk densely reticulate-punctate everywhere. In dorsal view the
alitrunk with a very shallow and feeble impression between pronotum and mesonotum and with a distinct
transverse line between mesonotum and propodeum, the entire surface densely reticulate-punctate.
Alitrunk lacking bizarre pilosity but dorsally with scattered minute simple hairs. Petiole and postpetiole
reticulate-punctate. In dorsal view both pedicel segments conspicuously broader than long, the postpetiole
with a concave anterior face and a shallow median longitudinal impression. Spongiform appendages
vestigial, reduced to a narrow posterior transverse strip on the petiole node, a similar but somewhat
broader strip on the postpetiole and with a short strip on the anterior postpetiolar border which traverses
the most concave part of the margin. In profile both segments lacking ventral spongiform appendages, the
lateral appendage of the postpetiole very reduced and present only at the posterior angle. Dorsal surfaces
of pedicel segments with minute hairs as on pronotum, the first gastral tergite with larger long strong hairs
which are clavate apically; the surface of the tergite weakly reticulate-shagreened and with short feeble
basal costulae. Colour medium brown.
The closest relative of laticeps is tiglath, which shows most of the characters of laticeps including
the 6-segmented antennae, lack of a spiniform tooth dorsally in the apical armament of the
mandible, orbicular hairs which occur immediately behind the clypeus and vestigial spongiform
appendages on the petiole and postpetiole. Features separating the two species are as follows.
laticeps
Larger species with longer mandibles,
HW 0-46-0-59, MI 42-18.
Dorsal surfaces of mandibular blades
lacking large flattened hairs.
Subbasal lobe of scape broad and
bluntly rounded.
Mesonotum strongly swollen and
humped posteriorly.
tiglath
Smaller species with shorter mandibles,
HW 0-39, MI 35.
Dorsal surfaces of each mandibular blade
with two large flattened hairs.
Subbasal lobe of scape narrow and
narrowly rounded.
Mesonotum not swollen, not humped
posteriorly.
MATERIAL EXAMINED
Ivory Coast: Abidjan, Banco Forest (/. Lobl); Adiopodoume (V. Mahnert & J.-L. Ferret); Bingerville
(V. Mahnert & J.-L. Ferret). Gregbeu (V. Mahnert & J.-L. Ferret). Nigeria: Ibadan (B. R. Critchley); nr
Zungeru (W. A. Sands). Cameroun: nr Yaounde (G. Tenon).
Epitritus minimus Bolton
(Fig. 46)
Epitritus minimus Bolton, 1972: 205, figs 1, 2. Holotype and paratype workers, GHANA: Eastern Region,
356 BARRY BOLTON
Akwapim Dist., Mampong, litter sample in cocoa farm, 27.vii.1970 (P. M. Room) (BMNH; MCZ)
[examined].
WORKER. TL 1 -2, HL 0-29, HW 0-29-0-31 , CI 100-107, ML 0-09, MI 31 , SL 0- 17, SI 55-59, PW 0-20-0-21 ,
AL 0-32 (2 measured).
Mandibles without preapical teeth and without an elongate spiniform tooth at the dorsal apex. Apical
armament of mandible a more or less vertical series of 6 small denticles of which the basalmost is the
largest , the prebasal approximately half this length and the upper group of 4 only about a quarter the length
of the basal. Dorsal surfaces of each mandibular blade with two large flattened hairs, the distal hair slightly
smaller than the proximal. Anterior clypeal margin with 4 large flattened hairs which project anteriorly,
and with a pair of smaller hairs. Clypeus with minute scale-like hairs only. Dorsum of head with numerous
large orbicular hairs which occur from the posterior clypeal margin to the occiput, the dorsum with a
narrow median longitudinal strip which is free from such hairs. Head without any other form of pilosity.
Antennae with 4 segments, the scape with a very strongly prominent subbasal lobe and fringed with large
flattened to spoon-shaped projecting hairs. Eyes minute, of a single ommatidium. Anteromedian portion
of clypeus shining, the remainder of the head finely and very densely punctulate-granular and dull.
Promesonotum fused in profile, the point of junction marked by a slight impression. Mesonotum behind
the impression shallowly convex and weakly inflated, ending posteriorly on a slightly higher level than the
propodeum. Propodeal dorsum convex and sloping downwards posteriorly, without teeth or spines but
the declivity margined by a conspicuous lamella on each side. Dorsum of pronotum, and to a lesser ex-
tent the mesonotum, with scattered minute stubble-like erect hairs which are shorter than the diameter of
the propodeal spiracular orifice. In dorsal view the alitrunk densely and finely punctulate-granular
everywhere, the shallow impression separating pronotum and mesonotum feebly visible medially, but the
mesonotum and propodeum separated by a distinct transverse line. Petiole and postpetiole both broader
than long in dorsal view, the latter much broader than the former; the petiole without spongiform
appendages and such appendages restricted on the postpetiole to a posterior transverse strip which is
broadly interrupted medially. In profile the petiole peduncle ventrally with a small anteriorly situated
lamella, without spongiform material. Postpetiole in profile with moderately developed spongiform
appendages posterolaterally and with a conspicuous ventral spongiform lobe. Petiole and postpetiole
punctulate-granular, with sparse minute hairs such as are present on the pronotum. Gaster with short weak
basal costulae, with short erect weakly clavate straight hairs. Colour dull yellow to yellowish brown.
Known only from two series, minimus is easily separated from its Afrotropical congeners by its
4-merous antennae, small size and strongly developed spongiform ventral lobes on the post-
petiole.
MATERIAL EXAMINED
Ghana: Mampong (P. M. Room). Cameroun: nr Yaounde (G. Terron).
Epitritus roomi Bolton
(Fig. 47)
Epitritus roomi Bolton, 1972: 206, figs 3, 4. Holotype worker and paratype female, GHANA: Eastern
Region, Akwapim Dist., Mampong, litter sample in cocoa farm, 10. iv. 1970 (P. M. Room) (BMNH)
[examined].
WORKER. TL 1 -8-2- 1 , HL 0-42-0-47, HW 0-54-0-58, CI 120-129, ML 0-15-0-17, MI 35-38, SL 0-25-0-28,
SI 45-48, PW 0-32-0-35, AL 0-46-0-52 (7 measured).
Mandibles with a single short recurved preapical tooth which is situated very close to the apex. Apical
armament of mandibular blades consisting of a dorsally situated elongate spiniform tooth, which crosses
the opposite mandible at full closure, subtended by an edentate or microscopically serrate lamina and
ending ventrally in a denticle. Dorsal surface of each mandibular blade with two large flattened hairs
arising on the distal half, the apicalmost hair the largest, the second narrower and tending to be directed
towards the midline between the mandibles. Anterior clypeal margin with 4 anteriorly projecting long
spatulate or strap-like hairs, the outer pair the longest. A much smaller pair of spatulate hairs also present,
projecting forwards from the clypeus between the larger hairs on each side. Clypeus dully shining,
equipped with minute appressed spatulate hairs. Remainder of head densely reticulate-punctate and with
conspicuous large orbicular hairs. Space on head behind the clypeus free of orbicular hairs, such hairs not
commencing immediately behind the posterior clypeal margin. Antennae with 6 segments, the scape with a
prominent anteriorly projecting subbasal lobe and fringed with large flattened to spoon-shaped hairs on the
leading edges. Alitrunk in dorsal view with a broad but shallow arched impression between pronotum and
THE AFROTROPICAL DACETINE ANTS 357
mesonotum. Propodeum separated from mesonotum by a feebly impressed line. In profile the dorsal
outline of the alitrunk is impressed at the pro-mesonotal junction and the mesonotum is convex posterior to
this. Propodeum shallowly convex and sloping downwards posteriorly to the broad laminae which border
the declivity. Alitrunk everywhere finely and densely reticulate-punctate and dull. Specialized hairs absent
from alitrunk but the dorsum, especially on the pronotum, with a scattered stubble of minute simple hairs.
Petiole and postpetiole in dorsal view both distinctly broader than long, the latter with a conspicuous
median impression. Spongiform appendages restricted to a narrow lamellar strip behind the petiole node,
another behind the postpetiole and one across the anterior margin of the postpetiole which is short and
restricted to the median concave portion of the margin. In profile the petiole with a narrow ventral carina,
the postpetiole with a vestigial ventral lobe and a larger posterolateral lobe. Pedicel segments and first
gastral tergite reticulate-punctate to granular, the latter with short feeble basal costulae but equipped with
a number of erect clavate hairs. Colour medium brown.
A very conspicuous species, easily separated from both its Afrotropical congeners which have
6-merous antennae by the form of the mandibles and distribution of orbicular hairs on the head.
In roomi the mandibular apex terminates dorsally in an elongate spiniform tooth which strongly
crosses over its counterpart on the opposite mandible at full closure, and the orbicular hairs do
not commence immediately behind the posterior clypeal margin. In both laticeps and tiglath the
mandibles do not have an enlarged spiniform tooth at the dorsal mandibular apex and the
orbicular hairs commence immediately behind the posterior clypeal margin.
MATERIAL EXAMINED
Ghana: Mampong (P. M. Room). Ivory Coast: Issoneu (V. Mahnert & J.-L. Ferret). Cameroon: nr
Yaounde (G. Terrori).
Epitritus tiglath sp. n.
HOLOTYPE WORKER. TL 1-4, HLO-34, HWO-39, CI 115, MLO-12, MI35, SLO-20, SI51, PWO-25, ALO-38.
Mandibles each with a single minute preapical denticle which is situated very close to the apex and may
be obscured by the flattened hairs. Apical mandibular armament consisting of a series of denticles arranged
in a more or less vertical row, the basalmost of which appears to be the largest. Without an elongate
spiniform tooth at the dorsal end of the series. Two large flattened hairs arise from the dorsal margin of
each mandibular blade on their distal halves; the hair closest to the apex is slightly smaller than the one
sited nearer the mandibular midlength. Anterior clypeal margin with 4 large spatulate to strap-like hairs
which project anteriorly, the outer pair, at the anterolateral angles, is the longest. A pair of much smaller
spatulate hairs also projects from the anterior clypeal margin above the mandibular bases and between the
larger hairs. Clypeus dully shining and with minute appressed spatulate hairs. Remainder of head
reticulate-punctate to granular and densely clothed with large orbicular hairs which occur from immediate-
ly behind the posterior clypeal margin to the highest point of the vertex. Antennae with 6 segments, the
scapes with a large anteriorly projecting subbasal lobe and fringed around the leading edges with large
flattened to spoon-shaped hairs. Outline shape of head capsule as shown for laticeps (Fig. 48). Alitrunk in
profile with the anterior portion of the mesonotum extremely shallowly concave , the posterior portion very
weakly convex just in front of the narrowly incised metanotal groove, the mesonotum not strongly swollen
or humped posteriorly. Propodeal dorsum sloping downwards posteriorly, without teeth but the declivity
margined on each side by a broad finely spongiform lamella. In dorsal view the pronotum and mesonotum
separated by a shallow impression, the mesonotum and propodeum separated by the conspicuous finely
incised line of the metanotal groove. Entire alitrunk finely and densely reticulate-punctate, without
specialized hairs but the dorsum with stubble-like microscopic erect simple hairs. Petiole node in dorsal
view broader than long, bordered posteriorly by a narrow lamellate strip. Postpetiole broader than long,
the anterior margin concave, the posterior margin convex and with a distinct median longitudinal
impression. The posterior margin of the postpetiole with a narrow lamellate strip, the anterior margin with
a short lamellate strip traversing the most concave portion of the border. Ventral surface of petiole with a
narrow longitudinal ridge, the postpetiole ventrally without lamellate or spongiform lobes but postero-
laterally with a small spongiform appendage. Both petiole and postpetiole reticulate-punctate to granular
and equipped dorsally with minute stubble-like hairs such as are seen on the alitrunk. First gastral tergite
with straight hairs which are clavate apically, the basigastral costulae short and weak; sculpture of fine
reticulation or shagreening. Colour light yellowish brown.
Holotype worker, Ivory Coast: Tai Forest, no. 45, 12.viii.1975, in sample of Oligomyrmex sp. (T.
Diomande) (BMNH).
358 BARRY BOLTON
This small species, the fourth Epitritus to be found in sub-Saharan Africa, is closest related to
laticeps. Characters separating the two are tabulated under laticeps.
STRUMIGENYS F. Smith
(Figs 49-66, 68-77)
Strumigenys F. Smith, 1860: 72. Type-species: Strumigenys mandibularis F. Smith, 1860: 72, by monotypy.
Labidogenys Roger, 1862: 249. Type-species: Labidogenys lyroessa Roger, 1862: 251, pi. 1, fig. 17, by
monotypy. [Synonymy by Brown, 1959a: 38.]
Pyramica Roger, 1862: 251. Type-species: Pyramica gundlachi Roger, 1862: 253, pi. 1, fig. 18, by
monotypy. [Synonymy by Brown, 1959a: 37.]
Proscopomyrmex Patrizi, 1946: 294. Type-species: Proscopomyrmex londianensis Patrizi, 1946: 295, figs 1,
2, by monotypy. [Synonymy by Brown, I949a: 15.]
Eneria Donisthorpe, 1948: 598. Type-species: Eneria excisa Donisthorpe, 1948: 598, fig. 1 (= Strumigenys
loriae Emery), by original designation. [Synonymy by Brown, 1949o: 15.]
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles extended into elongate narrow linear blades
(MI 26-65) which terminate in an apical fork of two spiniform teeth arranged in a more or less vertical
series, the dorsal fork tooth the longest. Intercalary teeth between the fork teeth sometimes present. Each
mandibular blade with one to two preapical teeth on the inner margin. Palp formula 1,1. Eyes ventrolater-
al, below the antennal scrobes. Antennae with 6 segments, sometimes funicular segments 2-3 very
reduced. Petiole node not bidentate dorsally. Postpetiole with spongiform appendages present. Special-
ized body pilosity frequently present.
By far the largest dacetine genus, Strumigenys has endemic species in all the zoogeographical
regions except the Palaearctic, and in all regions except the Nearctic it has a greater number of
species than any other dacetine genus (see table, p. 270).
Modern taxonomic understanding of Strumigenys depends almost entirely upon the works of
Brown who, beginning in 1948, has sorted the great diversity of forms previously included in the
genus and has completed a large number of descriptive, faunistic and revisionary works on
Strumigenys and its allies, on a world wide basis. Key works in this series include Brown (1948;
19490; 1949ft; 1953a; 1954; 1959«; 1959ft; 1962c; 1973c; and their included references), as well as
those other papers discussed in the introductory section of the present paper.
The first revisionary treatment of Afrotropical Strumigenys was that of Brown (1954), who
recognised 14 valid species. Prior to this date the only synthesizing studies of the genus in Africa
were those of Arnold (1917) on the South African fauna, and the regional catalogue of Wheeler
(1922). Each of these author's concept of Strumigenys included members of three genera by
modern reckoning, Strumigenys, Smithistruma and Serrastruma; the last two of these being
grouped together by Wheeler in a subgenus Cephaloxys. Brown (1948) realised that Strumigenys
as thus constituted contained several discrete evolutionary lines and proceeded to split the genus
into the groups which we recognize today. Collections made since Brown's (1954) revision have
greatly increased the number of African species, which now stands at 41.
As Brown (1954) indicated, it seems that the entire Afrotropical fauna of the genus belongs in
a single species-group, the rogeri-group, which has undergone an extensive adaptive radiation in
sub-Saharan Africa. The core-species of the group are represented by the /aura-complex, and
what I assume to be the most generalized character states within the complex (and thus within
the group as a whole) are shown byfaurei, petiolata and rufobrunea, as follows.
Mandibles with two preapical teeth on each blade, the proximal of which is the largest; inner margins of
blades without tumuli, lamellae or other excrescences. Apical fork of mandible without intercalary teeth,
the upper spiniform tooth of the fork longer than the lower, the lower spiniform fork tooth with an
extremely minute denticle or prominent angle ventrobasally which may only be visible when the
mandibular apices are viewed from behind.
Preocular notch present, the eye detached anteriorly from the side of the head. A ventral preocular
impression present behind and separate from the postbuccal impression, the ventral preocular impression
running from the preocular notch towards the ventral midline but not reaching the midline.
Eyes relatively large, generally at least equal to the maximum width of the scape. Antennal scapes linear
or slightly expanded at about the middle third, not strongly dorsoventrally flattened nor with the leading
THE AFROTROPICAL DACETINE ANTS 359
edges strongly convex. Mesonotum depressed posteriorly, with a single pair of standing hairs; alitrunk
without dense erect standing pilosity.
Upper scrobe margins bordered by a narrow lamina, sinuate or curved in full-face view.
Ground-pilosity small to minute on cephalic dorsum, smaller than the hairs lining the upper scrobe
margins.
Flagellate hairs absent from head.
Dorsum of head with 6 standing hairs, arranged in a row of 4 transversely close to the occipital margin,
and a more anteriorly situated pair at or close to the highest point of the vertex.
Resembling these very closely are the three species pretoriae, shaula and dromoshaula, which
conform to the above characterization but have the upper scrobe margins diverging
posteriorly almost in a straight line. In pretoriae the eyes are very large and the pronotal humeri
lack the flagellate hairs shown in the other two. Completing the faurei-complex is a cluster of
seven species which shows a gradual reduction and loss of the preocular notch and ventral
preocular impression whilst retaining the other characters listed above. Of these seven relahyla,
dy shaula, xenohyla and totyla show a small preocular notch; adrasora and rukha have the notch
vestigial to absent, variably developed even in a single series; absent in ettillax. Also in these
seven a tendency to broaden and flatten the antennal scapes is shown, perhaps best developed in
xenohyla.
Closely related to the 13 species of the /aura-complex is a species-pair consisting of bernardi
and vazerka which, whilst retaining most of the listed characters, have narrowed the head and
lengthened the mandibles and scapes beyond the range shown by the/awre/-complex, thus.
CI MI SI
faurei-complex 70-83 40-54 59-77
bernardi + vazerka 65-72 50-65 79-92
In the rogeri-complex (cacaoensis, londianensis, sarissa, rogeri) the upper scrobe margins lack a
bordering lamina, are strongly pinched in behind the frontal lobes and are concave to deeply
indented above the eyes. The scapes are long and slender. S. rogeri forms an intermediate stage
between the generalized condition shown in the/flwm-complex and the more strongly modified
remaining members of the rogeri-complex both in terms of the condition of the upper scrobe
margin and in the dentition. In rogeri the mandibles are armed as defined above, but in
cacaoensis, sarissa and londianensis intercalary teeth are present at least in the left apical fork,
and the preapical teeth are modified. In cacaoensis the proximal preapical teeth are reduced, at
most as large as the distals and usually smaller. In sarissa and londianensis the left mandible has
lost its distal preapical tooth. The cephalic ground-pilosity in rogeri is as infaurei and its allies,
whereas in the remaining species of the rogeri-complex the hairs lining the upper scrobe margins
are no broader than the ground-pilosity of the dorsum.
Perhaps derived from the rogeri-complex the five species of the ^corn-complex (scotti, hasty la,
zandala, murshila, helytruga) lack a lamina on the upper scrobe margin, have elongate
cylindrical scapes (SI 75-95), and have the hairs lining the upper scrobe margins slender, not or
only fractionally larger than the cephalic ground-pilosity. The mandibles are as defined for the
faurei-complex but the preocular notch is vestigial or absent and there is no ventral preocular
impression in the head. The eyes are relatively large in the first three species listed but are much
reduced in murshila and helytruga.
The 15 species of the arnoldi-complex themselves show a wide range of adaptations but
appear to be derived as a unit from ancestral forms related to relahyla and ettillax in the
faurei-complex. In the arnoldi-complex all species lack a preoccular notch and ventral preocular
impression, have the eyes Very small or vestigial (much smaller than the maximum width of the
scape), and show the development of conspicuous scale-like cephalic ground-pilosity. Within
the complex havilandi and korahyla have retained the relatively long mandibles of the ancestral
/flwre/-complex and have increased the scape length, so that their respective indices are MI
45-50, SI 80-90, as opposed to MI 26-45, SI 52-75 in the remainder of the arnoldi-complex. The
central species of the complex, represented by arnoldi, bitheria, omalyx, traegaordhi, mesahyla,
stygia and nimbrata, retain the mandibular dentition described toifaurei but in dextra, paranax
360 BARRY BOLTON
and katapelta the distal preapical tooth of the left mandible has been lost, and in irrorata the
distal preapical tooth of the right mandible is also missing. In all these species the antennal
scapes tend to be dorsoventrally flattened and have the leading edge arched convex. This
character is taken to extremes in tetraphanes where the leading edge of the scape is massively
expanded into a forward pointing lobe. In the arnoldi-comp\e\ in general there is an irregular
trend towards broadening the head and shortening the mandibles as one moves away from the
species closest to the/flMra'-complex, as follows.
CI MI
havilandi + korahyla 67-74 45-50
dextra + paranax 70-77 37-41
omalyx + stygia 74-84 35-41
spathoda + tetraphanes 77-97 26-37
It is certain that Quadristruma emmae represents a continuation of this trend (CI 80-85, MI
26-32) , further specialized by the development of cephalic orbicular hairs and the loss of the two
smallest funicular segments (numbers 2 and 3) so that the antennae have only 4 segments in all
(Fig. 67). This last specialization is not shown in the species listed above, but in nimbrata
funicular segments 2 and 3 are vestigial and often difficult to discern.
Finally the marleyi-complex (marleyi and pallestes), although related to the above and most
probably developed from them, has a striking overall convergence to the members of the
Oriental and Indo- Australian lyroessa-group (see Brown, 1948). As in the arnoldi-complex the
preocular notch and ventral preocular impression are absent in the marley /-complex, and the
scale-like pilosity is also present on the head. The eyes, however, are relatively larger in marleyi
and pallestes and the mandibular basal areas are much broadened and have an accentuated
basal-external angle. Additional teeth have developed on the mandibular apical fork, growing
from the ventral base of the lower spiniform fork tooth. The preapical mandibular armament is
as in the/flwm-complex. Both members of this complex are arboreal, a feature shared only with
cacaoensis in the Afrotropical fauna.
List of Afrotropical Strumigenys
rogeri-group omalyx sp. n.
adrasora sp. n. pallestes Bolton
arnoldi Forel paranax sp. n.
bernardi Brown petiolata Bernard sp. rev.
bitheria sp. n. pretoriae Arnold
cacaoensis Bolton relahyla sp. n.
dextra Brown rogeri Emery
dromoshaula sp. n. incisa Godfrey
dyshaulasp. n. sulfurea Santschi
ettillax sp. n. rufobrunea Santschi
faurei Arnold sp. rev. rukha sp. n.
hastyla sp. n. sarissa sp. n.
havilandi Forel scotti Forel
helytruga sp. n. shaula sp. n.
irrorata Santschi spathoda sp. n.
katapelta sp. n. stygia Santschi
korahyla sp. n. tetraphanes Brown
londianensis (Patrizi) totyla sp. n.
marleyi Arnold traegaordhi Santschi
mesahyla sp. n. vazerka sp. n.
murshila sp. n. xenohyla sp. n.
nimbrata sp. n. zandala sp. n.
Key to species (workers)
1 Preocular notch present; the ventrolateral margin of the head impressed, notched or indented
immediately in front of the eye, even if only feebly so (Figs 49-58, 71-74) 2
Preocular notch absent; the ventrolateral margin of the head continuous to the anterior margin
THE AFROTROPICAL DACETINE ANTS 361
of the eye, without trace of an impression, notch or indentation immediately in front of the
eye (Figs 59-66, 75-77) 21
2 Blade of left mandible with 1 preapical tooth 3
Blade of left mandible with 2 preapical teeth 5
3 Apical fork of left mandible without an intercalary small tooth between the spiniform fork
teeth. Small species with relatively slightly longer mandibles, HW 0-36-0-41, MI 55-65.
(Cameroun , Gabon , Zaire , Angola) bernardi (part , p. 366)
Apical fork of left mandible with an intercalary small tooth between the spiniform fork teeth
(Fig. 50). Larger species with relatively slightly shorter mandibles, HW 0-50-0-70, MI 51-55 4
4 Pronotal humeri with straight feebly clavate stout hairs, the anterior pronotal margin without a
second pair of hairs between the humeral pair. Propodeum without sharp teeth. Leading
edge of scape with hairs on the proximal half directed basally (except for the 2 basalmost),
hairs on the distal half directed apically . (Kenya) londianensis (p. 377)
Pronotal humeri with fine flagellate hairs and the anterior pronotal margin with a second pair of
stouter hairs between the humeral pair. Propodeum with sharp teeth. All hairs on leading
edge of scape directed apically (Fig. 50). (Zaire, Rwanda, Burundi) sarissa (p. 390)
5 Apical fork of left mandible with a strong intercalary tooth and a denticle between the
spiniform fork teeth (Fig. 49). With the head in full-face view the upper scrobe margin
strongly notched above the eye. Large species, HW >0-60, with the preapical teeth about
equal in size or the distal larger than the proximal. (Ghana, Nigeria) cacaoensis (p. 367)
Apical fork of left mandible without intercalary tooth or denticle between the spiniform fork
teeth. With the head in full-face view the upper scrobe margin continuous above the eye.
Smaller species, HW <0-60, with the proximal preapical tooth larger than the distal 6
6 Preocular notch on each side continued onto the ventral surface of the head as a transverse
impression or groove of varying length, which runs towards but does not reach the midline;
this impression situated behind the post-buccal groove or impression and independent of it
(Figs 71, 72) 7
Preocular notch on each side not continued onto the ventral surface of the head, the ventral
surface convex and without impressions behind the post-buccal groove or impression
(Figs73,74) 15
7 Antennal scapes relatively long, SI 79-92 8
Antennal scapes relatively short, SI 64-74 10
8 Mandibles more or less straight in full-face view, not conspicuously bowed outwards (Fig. 51).
Larger species, HW 0-42-0-52. (Cosmopolitan tramp species, very widespread in Afrotro-
pical region) rogeri (p. 387)
Mandibles conspicuously bowed outwards in full-face view (Fig. 52). Smaller species,
HWO-36-0-41 9
9 Dorsum of propodeum densely reticulate-punctate and dull. (Cameroun, Gabon, Zaire,
Angola) bernardi (part, p. 366)
Dorsum of propodeum smooth and shining. (Ivory Coast, Ghana, Nigeria) vazerka (p. 397)
10 Eyes very large, their maximum diameter 0-23-0-24 xHW or more (Fig. 53). (Botswana, South
Africa) pretoriae (p. 385)
Eyes smaller, the maximum diameter less than 0-20 x HW 11
11 Pronotal humeri without flagellate hairs. (Guinea, Ivory Coast, Ghana, Nigeria, Cameroun,
Gabon, Angola, Sudan, Central African Republic) petiolata (p. 384)
- Pronotal humeri with flagellate hairs 12
12 Disc of postpetiole very finely and densely longitudinally superficially sculptured. (South
Africa) faurei (p. 371)
Disc of postpetiole smooth and shining (when clean, frequently the surface with a waxy bloom
present) 13
13 Distal preapical tooth of left mandible short and separated from the proximal by a distance
which at least is equal to the length of the distal tooth but which is usually much more.
(Guinea, Ivory Coast, Togo, Nigeria) rufobrunea (p. 389)
Distal preapical tooth of left mandible long and separated from the proximal by a distance
which is distinctly much less than the length of the distal tooth 14
14 Extension of preocular notch on ventral surface of head forming a parallel-sided groove which
is narrower than the maxirnum diameter of the eye ; the edges of the groove sharply defined.
(Zimbabwe) shaula (p. 392)
- Extension of preocular notch on ventral surface of head forming a broad dish-like impression
362 BARRY BOLTON
which is at least as broad as the maximum diameter of the eye and usually broader; the edges
of this impression are rounded and not sharply defined. (Burundi) dromoshaula (p. 369)
15 Eyes small, with only 8 ommatidia. Mandibles in full-face view with outer margins of blades
straight. Antennal scapes with SI 79. (Angola) helytruga (p. 374)
Eyes larger, with more than 8 ommatidia. Mandibles in full-face view with outer margins of
blades convex, the mandibles usually bowed outwards. Antennal scapes with SI 65-77 16
16 Pronotal humeri without flagellate hairs. (Cameroun) totyla (p. 395)
Pronotal humeri with flagellate hairs 17
17 In full-face view the projecting hairs on the leading edges of the scapes large and broadly
spoon-shaped, about equal in size to the large broadly spoon-shaped hairs fringing the upper
scrobe margins. Upper scrobe margins with a broad lamellate rim or flange (Fig. 54).
(Cameroun, Zaire) xenohyla (p. 398)
In full-face view the projecting hairs on the leading edges of the scapes slender, either distinctly
narrower than those fringing the upper scrobe margins or with the hairs in both places
conspicuously slender, spatulate to narrowly elongate spoon-shaped. Upper scrobe margins
with a narrow rim or flange 18
18 In profile the area of the ventral postpetiolar spongiform lobe conspicuously much less than the
visible area of the postpetiolar disc (Fig. 68) . Infradental lamellae of propodeum very narrow
or vestigial, the teeth free or nearly free of the lamellae. (Rwanda, Burundi) adrasora (part, p. 364)
- In profile the area of the ventral postpetiolar spongiform lobe equalling or exceeding the visible
area of the postpetiolar disc. Infradental lamellae of propodeum broad, engaging half or
more of the length of the teeth 19
19 With the postpetiole in dorsal view spongiform material is visible projecting laterally at and in
front of the midlength of the disc. (Uganda, Kenya) rukha (part, p. 389)
With the postpetiole in dorsal view projecting spongiform material is restricted to the apices of
the posterior transverse strip and does not occur in front of the midlength of the disc 20
20 Basigastral costulae sparse and forming a continuous row across the tergite, without a broad
clear central area and usually without secondary fine costulae arising some distance behind
the basal strip of the tergite. Scapes shorter, SI 65-69. Occipital margin broadly and
shallowly impressed (Fig. 57). (Cameroun, Zaire, Angola) relahyla (p. 386)
Basigastral costulae dense and very obviously radiating from each side of a broad clear central
area, with secondary fine costulae present which arise some distance behind the basal strip of
the tergite. Scapes longer, SI 70-74. Occipital margin narrowly and deeply impressed
(Fig. 55). (Zimbabwe) : dyshaula (p. 370)
21 Blade of left mandible with 1 preapical tooth 22
Blade of left mandible with 2 preapical teeth 25
22 Blade of right mandible with 1 preapical tooth. (Zimbabwe, South Africa) irrorata (p. 375)
Blade of right mandible with 2 preapical teeth 23
23 Apical fork of left mandible with an intercalary denticle between the spiniform teeth. Dorsum
of head without a transverse row of 4 standing hairs close to the occipital margin. Head
broad, CI 85-90 (Fig. 61). (Burundi, Kenya) katapelta (p. 375)
Apical fork of left mandible without an intercalary denticle between the spiniform teeth.
Dorsum of head with a transverse row of 4 standing hairs close to the occipital margin and
with a pair situated anterior to this row. Head narrower, CI <80 24
24 Pronotal humeri each with a straight stout thickly clavate projecting hair. (Cameroun, Gabon)
paranax (p. 383)
Pronotal humeri each with a long fine flagellate projecting hair. (Cameroun, Gabon, Angola,
Central African Republic, Uganda) dextra (p. 368)
25 Ventral tooth of left mandibular apical fork with an adventitious tooth and an intermediate
denticle arising from its ventrobasal surface (Fig. 59). Sides of alitrunk densely reticulate-
punctate everywhere. Arboreal species 26
Ventral tooth of left mandibular apical fork without additional teeth arising from the
ventrobasal surface or at most with an extremely minute denticle-like point at the extreme
base. Sides of alitrunk usually with at least the pleurae smooth and shining, not densely
reticulate-punctate everywhere 27
26 Pronotal humeri each with a stout straight laterally directed hair which is clavate apically.
Petiole node weakly transversely striate dorsally. (Ghana, Nigeria) pallestes (p. 383)
Pronotal humeri without projecting hairs of any description. Petiole node punctate dorsally.
(South Africa) marleyi (p. 378)
THE AFROTROPICAL DACETINE ANTS 363
27 Pronotal humeri without flagellate hairs 28
- Pronotal humeri with flagellate hairs 31
28 Leading edge of antennal scape grossly expanded into an enormous anteriorly projecting lobe
(Fig. 60). (Uganda, Cameroun, Gabon) tetraphanes (p. 395)
- Leading edge of antennal scape not expanded into a gross projecting lobe 29
29 Antennal scape broadest close to the base, thereafter evenly tapering to the apex (Fig. 64).
Mandibles and scapes longer, head narrower, MI 48-50, SI 82-85, CI 67-71 . (Cameroun)
korahyla (p. 376)
- Antennal scape broadest at or just beyond the midlength, narrowing both proximally and
distally (Fig. 63). Mandibles and scapes shorter, the head broader, MI 35-41, SI 65-75, CI
74-84 30
30 Postpetiolar disc smooth and shining. (Zimbabwe, Kenya) arnoldi (p. 365)
- Postpetiolar disc sculptured. (Kenya) omalyx (p. 382)
31 Funicular segments 2 and 3 vestigial and difficult to see so that the funiculus appears to have
only 3 segments altogether. Combined length of funicular segments 2 and 3 less than half the
length of segment 4 (the penultimate segment) 32
- Funicular segments 2 and 3 reduced but easily visible, the funiculus distinctly with 5 segments.
Combined length of funicular segment 2 and 3 more than half the length of segment 4 33
32 Width of lamellate flange bordering upper scrobe margins distinctly greater than the maximum
diameter of the eye, approaching the maximum width of the scape. Petiole node in dorsal
view as broad as long. (Cameroun) hithcria (p. 367)
- Width of lamellate flange bordering upper scrobe margins distinctly less than the maximum
diameter of the eye , only a fraction of the maximum width of the scape . Petiole node in dorsal
view much broader than long. (Ivory Coast, Ghana) nimbrata (p. 381)
33 Disc of postpetiole sculptured 34
- Disc of postpetiole smooth and shining (when clean , frequently with a waxy bloom present) 35
34 Ground-pilosity of head, upper scrobe margins and leading edges of scapes of narrow spatulate
hairs. Scapes slender and cylindrical, the leading edges not expanded and convex. MI 48, SI
77, CI 76. (Rwanda) murshila (p. 380)
- Ground-pilosity of head, upper scrobe margins and leading edges of scapes of broadly
spoon-shaped or scale-like to suborbicular hairs. Scapes flattened, the leading edges
expanded and convex. MI 36-40, SI 63-68, CI 80-84. (Kenya, Zimbabwe, Cameroun,
Angola) stygia (p. 394)
35 Maximum diameter of eye distinctly much less than maximum width of scape 36
- Maximum diameter of eye at least equal to and usually distinctly greater than maximum width
of scape 39
36 Mandibles and scapes very short, MI 26-30, SI 55-61 (Fig. 62). (Ivory Coast, Togo, Cameroun)
spathoda (p. 393)
Mandibles and scapes longer, MI 42-50, SI 70-90 37
37 Scapes long, SI 80-90. Proximal preapical tooth of left mandible situated close to midlength of
the blade so that the distance from the basal midpoint of the tooth to the clypeal margin is less
than twice the distance from the basal midpoint of the tooth to the distal base of the dorsal
fork tooth (Fig. 65). (South Africa) havilandi (p. 373)
- Scapes shorter, SI 70-75 . Proximal preapical tooth of left mandible situated far along the blade
so that the distance from the basal midpoint of the tooth to the clypeal margin is more than
twice the distance from the basal midpoint of the tooth to the distal base of the dorsal fork
tooth 38
38 Dorsum of head with 6 standing hairs. Cephalic ground-pilosity much narrower on the
posterior half of the head than on the anterior half, the former narrowly spatulate, the latter
spoon-shaped to scale-like. Ventral spongiform lobe of postpetiole at most equal to the
exposed area of the postpetiolar disc in profile. (South Africa) traegaordhi (p. 396)
- Dorsum of head with 4 standing hairs. Cephalic ground-pilosity spoon-shaped to scale-like
everywhere, not much narrower on the posterior half of the head. Ventral spongiform lobe
of postpetiole much larger than the exposed area of the postpetiolar disc in profile (Fig. 69) .
(Zimbabwe) mesahyla (p. 379)
39 Ventral spongiform lobe of postpetiole in profile distinctly smaller than the exposed area of the
postpetiolar disc (Fig. 68). Ventral appendage of petiole not spongiform. (Rwanda, Burun-
di) adrasora (part, p. 364)
- Ventral spongiform lobe of postpetiole in profile as large as or larger than the exposed area of
364 BARRY BOLTON
the postpetiolar disc. Ventral appendage of petiole spongiform 40
40 Upper scrobe margins fringed by many inconspicuous fine spatulate to narrowly spoon-shaped
hairs which are the same size as the hairs of the cephalic ground-pilosity (Fig. 66). Head
narrower and scapes longer, CI 64-75 , SI 74-95 41
Upper scrobe margins fringed by few conspicuous broadly spatulate to spoon-shaped large
hairs which are distinctly much larger than the minute hairs of the cephalic ground-pilosity.
Head broader and scapes shorter, CI 75-81, SI 59-73 43
41 Mandibles and scapes long, MI 46-50, SI 88-95. Petiole node in dorsal view at least as long as
broad, often longer than broad. (Sao Tome I., Seychelles) scotti (p. 391)
Mandibles and scapes shorter, MI 42-46, SI 75-82. Petiole node in dorsal view broader than
long 42
42 In dorsal view some or all of the hairs on the petiole, postpetiole and base of the first gastral
tergite thickened or clavate. Smaller species, HW 0-34-0-39. (Ivory Coast, Nigeria, Came-
roun, Gabon, Angola, Burundi) hastyla (p. 372)
- In dorsal view all hairs on the petiole, postpetiole and base of the first gastral tergite simple.
Larger species, HW 0-42-0-44. (Annobon I.) zandala (p. 399)
43 Yellow species with longer mandibles and scapes , MI 47-49 , SI 67-73 . (Uganda , Kenya)
rukha (part, p. 389)
- Blackish brown species with shorter mandibles and scapes, MI 40-43, SI 59-60. (Cameroun)
ettillax (p. 371)
Strumigenys adrasora sp. n.
(Fig. 68)
HOLOTYPE WORKER. TL 2-4, HL 0-60, HW 0-45, CI 75, ML 0-32, MI 53, SL 0-33, SI 73, PW 0-29, AL 0-58.
Mandibles in full-face view evenly shallowly bowed outwards. Apical fork of each mandible consisting of
two spiniform teeth, the upper largest, without intercalary teeth or denticles. Each mandibular blade with
2 preapical teeth, the proximal the largest. The distal tooth approximately half the length of the proximal
and slightly shorter to about equal in length to the distance separating their bases. Upper scrobe margins
not bordered by a projecting flange, the eyes visible in full-face view. Maximum diameter of eye slightly
greater than maximum width of scape. Preocular notch present but small and shallow, not continued onto
the ventral surface of the head as a transverse impression or groove. Antennal scapes slender, very
shallowly curved basally, their leading edges equipped with a row of elongate narrowly spatulate hairs
which are curved apically. Upper scrobe margins with an anteriorly curved row of elongate spatulate hairs
which are narrow, only fractionally broader than those on the scapes. Dorsum of head with 6 standing hairs
arranged in a transverse row of 4 close to the occipital margin and a more anteriorly situated pair. Cephalic
ground-pilosity of inconspicuous simple short hairs anteriorly but with the hairs tending to become
narrowly spatulate on the occipital lobes. Dorsum of head reticulate-punctate. Pronotal humeri each with a
single long fine flagellate hair. Mesonotum with a single pair of standing hairs. Dorsal alitrunk otherwise
only with sparse minute ground-pilosity which is appressed. Metanotal groove obsolete, scarcely discern-
ible on the dorsal alitrunk and not impressed in profile. Posterior portion of mesonotum only shallowly
depressed behind the level of the standing hairs. Propodeal teeth lamellate and narrowly triangular, the
infradental lamella narrow, engaging one-quarter or less of the length of the tooth. Sides of pronotum with
faint sculpture anteriorly and posteriorly but smooth medially. Pleurae mostly glassy smooth but with
peripheral weak punctulation. Sides of propodeum below the level of the spiracle punctate. Pronotum
dorsally very finely longitudinally striolate-costulate, the remainder of the dorsal alitrunk punctate. Petiole
node punctate dorsally, the postpetiole mostly smooth but with faint vestiges of shagreening towards the
sides. Spongiform appendages of pedicel segments very reduced, the subpetiolar strip vestigial, commenc-
ing only at the midlength of the peduncle. Ventral lobe of postpetiole small, in profile distinctly smaller
than the exposed portion of the postpetiole and about the same size as the lateral spongiform lobe. In
dorsal view the petiole node with a narrow posterior collar, the postpetiole with a narrow posterior strip;
the sides in front of the midlength not showing projecting spongiform tissue. Base of first gastral tergite
with a narrow lamellate strip from which the basigastral costulae arise. Petiole, postpetiole and first gastral
tergite with stout standing hairs, most of which are thickened apically. Colour blackish brown to black.
PARATYPE WORKERS. TL 2-3-2-4, HL 0-59-0-62, HW 0-44-0-47, CI 73-78, ML 0-30-0-33, MI 50-54, SL
0-33-0-36, SI 72-77, PW 0-29-0-31, AL 0-58-0-62 (10 measured).
As holotype but in some the preocular notch is vestigial and very difficult to see, to all intents and
purposes absent. In most the distal preapical tooth is as described above but in a few is distinctly shorter
than the distance separating the bases of the two preapical teeth. The subpetiolar spongiform appendage
THE AFROTROPICAL DACETINE ANTS 365
may be reduced to a small lobe under the node or reduced to a narrow short carina. In a few individuals the
propodeal teeth are almost free of the infradental lamellae.
Holotype worker, Rwanda: Rangiro, ix.1976 (P. Werner) (MHN).
Paratypes. 26 workers with same data as holotype (MHN; MCZ; BMNH; ENSA).
Non-paratypic material examined. Burundi: Bujumbura (A. Dejean); Bugarama (A. Dejeari).
The non-paratypic material is lighter than the type-series, being yellowish brown with a dark
brown gaster, but otherwise matches the above description. 5. adrasora belongs to the
faurei-complex where it is diagnosed by its poorly defined to vestigial preocular impression,
reduced spongiform appendages on the pedicel segments and narrow hairs on the upper scrobe
margins. The variably developed preocular notch has led me to key this species in two places,
firstly amongst the faurei-comp\ex members where it really belongs, but secondly amongst the
members of the scom'-complex towards the end of the key, where its close relatives rukha and
ettillax also occur. 5. adrasora is separated from all of these forms by its reduced spongiform
appendages on the pedicel segments.
Strumigenys arnoldi Forel
Strumigenys arnoldi Forel, 19136: 114. Holotype worker, ZIMBABWE: Bulawayo, under stone in nest of
Bothroponera krugeri (Forel) (G. Arnold) (MHN) [examined].
Strumigenys arnoldi Forel; Brown, 1954: 26.
WORKER. TL 2-0-2-1 , HL 0-54-0-60, HW 0-39-0-46, CI 74-78, ML 0-22-0-23, MI 38-41 , SL 0-30-0-32, SI
68-75, PW 0-24-0-26, AL 0-54-0-58 (3 measured).
Mandibles in full-face view broadest near the base and gradually tapering towards the apex. Apical fork
of each mandible with 2 spiniform teeth , without intercalary teeth or denticles . Preapical armament of each
mandibular blade of 2 teeth, the proximal preapical much longer than the distal and the distance separating
their bases less than the height of the distal preapical tooth. Both preapical teeth situated in the apical third
of the length of the blade. Upper scrobe margins forming a feeble rim or flange, the eyes not visible in
full-face view. Eyes small, the maximum diameter distinctly less than the maximum width of the scape.
Preocular notch absent, the ventral surface of the head without a preocular transverse impression or
groove on each side. Antennal scapes shallowly curved in the basal third, the leading edges weakly convex
and equipped with a row of apically curved spoon-shaped hairs which are about the same size as those
fringing the upper scrobe margins. Dorsum of head with dense anteriorly curved hairs which are scale-like
to stud-like in full-face view, the upper scrobe margins fringed with similar hairs. Dorsum of head
everywhere finely and densely reticulate-punctate. Pronotal humeri without flagellate hairs. Mesonotum
with a single pair of stout standing hairs. Ground-pilosity of dorsal alitrunk like that of cephalic dorsum but
the hairs tending to be smaller and sparser. Posterior portion of mesonotum depressed, the metanotal
groove absent. Propodeal teeth broadly triangular and conspicuous, subtended by broad sinuate in-
fradental lamellae. Sides of alitrunk smooth except for some punctures on the upper portion of the
mesopleuron. Entire dorsal alitrunk finely reticulate-punctate, on the pronotum this sculpture overlaid by
some fine longitudinal rugulation. Dorsum of petiole node finely punctate, the postpetiolar disc smooth
and shining. Spongiform appendages of pedicel segments well developed, the petiole with a broad ventral
strip which has its ventral free margin indented before the midlength. Ventral spongiform lobe of
postpetiole larger than the exposed area of the postpetiolar disc in profile and distinctly larger than the
lateral spongiform lobe. Basigastral costulae short and sparse, widely spaced on each side of a broad
central clear area. Petiole, postpetiole and first gastral tergite with stout standing hairs which are swollen to
clavate apically. Colour dull yellow.
Within the arnoldi-complex the species tetraphanes, korahyla, arnoldi and omalyx are charac-
terized by lacking pronotal flagellate hairs whilst retaining the usual mandibular dentition of 2
preapical teeth on each blade. Of the four tetraphanes is instantly recognized by its short broad
head and enormous plate-like lobate extension of the antennal scapes. S. korahyla has long
narrow mandibles and scapes (MI 48-50, SI 82-85), and has the scapes evenly tapering from
base to apex. S. arnoldi is separated from omalyx by details of sculpture as in the latter the sides
of the pronotum and the postpetiolar disc are strongly sculptured, and the reticulate-punctate
sculpture of the pronotal dorsum is not overlaid by longitudinal rugulae. In arnoldi, on the other
hand, the pronotal sides and postpetiolar disc are smooth, and the pronotal dorsum has
longitudinal rugulae overlying the reticulate-punctate sculpture.
366 BARRY BOLTON
MATERIAL EXAMINED
Zimbabwe: Bulawayo (G. Arnold). Kenya: Eldoret (Patrizi).
Strumigenys bernardi Brown
Strumigenys new species, Bernard MS; Brown, 1954: 16 (described but not named).
Strumigenys bernardi Brown, 1960: 206. Holotype worker, ZAIRE: 10 miles (16 km) E. of Stanleyville
(= Kisangani), iii.1948, no. 2225 (N. A. Weber) (MCZ) [examined]
WORKER. TL 1-7-2-1, HL 0-52-0-58, HW 0-36-0-41, CI 67-72, ML 0-30-0-36, MI 55-65, SL 0-30-0-36, SI
82-92, PW 0-22-0-26, AL 0-44-0-54 (25 measured).
Mandibles long and slender, conspicuously bowed outwards in full-face view. Apical fork of both
mandibles without intercalary denticles. Left mandible usually with only a single spiniform preapical tooth
(the proximal) but extremely rarely a minute almost invisible distal preapical denticle can be seen very
close to the dorsal tooth of the apical fork . Right mandible with 2 preapical teeth , the proximal much longer
than the distal and the latter frequently concealed by the opposing upper fork tooth when the mandibles are
closed. Upper scrobe margins close together immediately behind the frontal lobes, usually feebly sinuate
close to the frontal lobes and then evenly divergent behind, not concave or impressed above the eyes and
without a conspicuous bordering lamella or flange. Preocular notch strongly developed and deep, the
anterior portion of the eye detached from the side of the head. Preocular notch continuing onto ventral
surface of head as a transverse impression. Maximum diameter of eye equal to or greater than maximum
width of scape. Antennal scapes straight and slender, the leading edge with a row of apically curved
narrowly spatulate hairs. Cephalic ground-pilosity of inconspicuous narrowly spatulate hairs which are
curved anteriorly. Laterally projecting curved spatulate to spoon-shaped hairs which border the upper
scrobe margins distinctly larger than the ground-pilosity. Vertex of head with 6 standing hairs arranged in a
row of 4 close to the occipital margin and a more anteriorly situated pair. Dorsum of head finely and
shallowly reticulate-punctate everywhere. Pronotal humeri each with a long fine flagellate hair and the
mesonotum with a single pair of stout standing hairs. Ground-pilosity on alitrunk minute and appressed.
Metanotal groove represented by a transverse line on the dorsum, not or only vestigially impressed in
profile. Posterior portion of mesonotum depressed and on same level as propodeum, the latter armed with
a pair of triangular teeth subtended by narrow infradental lamellae. Pleurae of alitrunk smooth and shining
or at most with faint peripheral punctulae. Sides of propodeum usually punctate, less commonly virtually
unsculptured. Pronotal dorsum usually with fine longitudinal rugulae which diverge posteriorly and are
superimposed on a punctate ground-sculpture, but one or the other component may be emphasised so that
at one extreme the pronotum is punctate dorsally and at the other almost entirely rugulose. Mesonotum
and propodeum reticulate-punctate dorsally. Petiole node punctate dorsally, the postpetiole smooth or
weakly sculptured, often with feeble longitudinal costulae and sometimes with vestigial punctures. Petiole
with a vestigial ventral appendage which at most is represented by a narrow carina, the node posteriorly
with a slender transverse collar. Ventral spongiform lobe of postpetiole moderately developed, larger than
the lateral lobe in profile. In dorsal view the postpetiole with a very narrow anterior and broader posterior
spongiform strip. Base of first gastral tergite with a narrow lamellar strip from which the basigastral
costulae arise. Petiole, postpetiole and gaster with strong hairs which are weakly clavate apically. Colour
dull yellow.
Among the Afrotropical Strumigenys in which the preocular notch is developed bernardi and
vazerka form a close species-pair characterized by their long bowed mandibles, long scapes, lack
of intercalary teeth in the mandibular apical forks, and by having the preocular notch continued
as an impression on the ventral surface of the head. Most samples of bernardi are instantly
distinguishable from vazerka as the former has only a single preapical tooth on the left
mandibular blade whilst the latter has two. However, now and again a specimen of bernardi with
a minute vestige of the left distal preapical tooth is found, but here bernardi is recognized by its
reticulate-punctate propodeal dorsum, which in vazerka is smooth.
MATERIAL EXAMINED
Cameroun: Nkoemvon (D. Jackson); nr Yaounde (G. Terron). Gabon: Plateau d'Ipassa (J. A. Barra);
Makokou (W. H. Gotwald); Makokou (/. Lieberburg). Zaire: Ituri Forest (N. A. Weber); Stanleyville
(N. A. Weber). Angola: R. Kahingo; R. Chicapa, Saurimo (L. de Carvalho); Dundo (L. de Carvalho);
Camudembele (L. de Carvalho); R. Mussungue (L. de Carvalho); R. Chinana (A. Machado); Salazar
(P. Hammond).
THE AFROTROPICAL DACETINE ANTS 367
Strumigenys bitheria sp. n.
HOLOTYPE WORKER. TL 1 -9, HL 047, HW 0-37, CI 79, ML 0-20, MI 43, SL 0-28, SI 76, PW 0-22, AL 0-47.
Outer margins of mandibles very feebly convex in full-face view, broadest at about the level of the
proximal preapical tooth and weakly tapering towards the base. Apical fork of 2 spiniform teeth, without
intercalary teeth or denticles. Each mandible with 2 preapical teeth, the proximal of which is the largest and
is situated just distal of the midlength of the blade. The distance separating the bases of the preapical teeth
is less than the length of the distal preapical tooth. Anterior two-thirds of upper scrobe margins with a very
conspicuous broad bordering translucent lamella or flange which is distinctly broader than the maximum
diameter of the eye and approaches the maximum width of the scape. Eyes very small, with only 3-4
ommatidia, the maximum diameter of the eye distinctly less than the maximum width of the scape.
Preocular notch absent, the ventral surface of the head without a transverse preocular groove or
impression on each side. Antennal scapes narrow, slightly curved at the base and broadest at about the
midlength, not distinctly flattened nor with the leading edges convex. Hairs fringing the leading edges of
the scapes very slender, much smaller than those fringing the upper scrobe margins. Funicular segments 2
and 3 very reduced , vestigial , their combined length less than half that of the fourth (penultimate) funicular
segment. Dorsum of head from posterior margin of clypeus to about the midlength densely clothed with
broad conspicuous scale-like hairs. Behind this level mediodorsally are only much smaller sparse hairs but
towards the sides of the occipital lobes are hairs similar in construction but smaller than those on the
anterior half. Upper scrobe margins fringed with large anteriorly curved spoon-shaped hairs. Dorsum of
head with 6 standing hairs arranged in a transverse row of 4 close to the occipital margin and a more
anteriorly situated pair. Cephalic dorsum reticulate-punctate. Pronotal humeri each with a single fine
flagellate hair. Mesonotum with a single pair of stout standing hairs. Ground-pilosity of dorsal alitrunk of
small curved hairs. Posterior portion of mesonotum depressed behind the level of the standing hairs.
Propodeal teeth triangular, elongate and narrow, longer than their basal width and subtended by narrow
infradental lamellae. Sides of pronotum punctate, remainder of sides of alitrunk mostly smooth, with some
punctate patches above and below a large smooth central area. Dorsal alitrunk reticulate-punctate, the
pronotum also with some fine longitudinal rugulae. Petiole node as long as broad in dorsal view, the surface
punctate. Postpetiole smooth. Spongiform appendages of pedicel segments poorly developed, the petiole
with a narrow ventral strip and the node with a lateral lobe which is scarcely broader than the posterior
collar. Ventral spongiform lobe of postpetiole about equal in size to the exposed area of the propodeal disc
in profile. Basigastral costulae very short but stout, arising across the width of the tergite rather than on
each side of a central clear area. Petiole, postpetiole and gaster with stout standing hairs which are swollen
or thickened apically, colour light brown.
Holotype worker, Cameroon: Nkoemvon, 2.iii.l980 (D. Jackson) (BMNH).
Closely related to nimbrata and sharing the striking reduction of the second and third funicular
segments seen in that species, bitheria is distinguished by its broad lamellate upper scrobe
margins, longer distal preapical teeth, stronger pronotal sculpture, longer narrower propodeal
teeth, and by possessing a petiole node which is as long as broad in dorsal view.
Strumigenys cacaoensis Bolton
(Fig. 49)
Strumigenys cacaoensis Bolton, 1971: 59, fig. 1. Holotype worker, paratype workers and female, NIGERIA:
GambariExp. Sta. (Cocoa Res. Inst. of Nigeria), 10. vii. 1969, rot hole in trunk of cocoa tree (B. Bolton)
(BMNH; MCZ) [examined].
WORKER. TL 2-7-3-3, HL 0-83-0-92, HW 0-72-0-80, CI 86-91, ML 0-38-0-45, MI 44-50, SL 0-48-0-50, SI
62-67, PW 0-36-0-40, AL 0-76-0-88 (11 measured).
Apical fork of left mandible with a dorsally situated small tooth and a ventrally situated denticle between
the upper and lower spiniform fork teeth. Apical fork of right mandible with an intercalary denticle only.
Preapical armament of both mandibular blades consisting of a pair of teeth or denticles. In general the
proximal preapical tooth is very small, reduced to a denticle, and the distal is distinctly larger, but in a few
specimens the two are of approximately equal size on each blade. The usual configuration seen in the
rogm-group, with the distal preapical tooth distinctly smaller than the proximal, is not found in this
species. Upper scrobe margins not bordered by a lamella or flange , narrowly concave and with a pinched-in
appearance immediately behind the convex frontal lobes. Posterior to this the upper scrobe margins are
evenly divergent then suddenly and deeply excavated above the eye, the site of this excavation directly
368 BARRY BOLTON
above the strongly developed preocular notch so that the two together form a broad deep groove running
down the side of the head in front of the eye. Anterior portion of the eye detached from the head and the
preocular notch continued onto the ventral surface of the head as a broad transverse impression. Antennal
scapes narrow in the basal fifth than broadened, the short curved stout hairs on the leading edge directed
apically and broadly spatulate to spoon-shaped. Head reticulate-punctate everywhere, the ground-pilosity
of dense short broadly spatulate to scale-like hairs which are curved anteriorly and quite closely applied to
the surface, the hairs bordering the upper scrobe margins no longer than those sited elsewhere on the
dorsum. Vertex of head without standing hairs of any description. Pronotal humeri without projecting
hairs of any description, the mesonotum bearing a single pair of stout hairs which represent the only
standing pilosity on the dorsal alitrunk. Ground-pilosity of alitrunk of short sparse spatulate hairs which
are closely applied to the surface. Propodeum armed with a pair of acute spines, the infradental lamellae
very narrow or vestigial. Sides and dorsum of alitrunk and of pedicel segments reticulate-punctate
everywhere, often with a granular appearance on the latter. Metanotal groove shallowly impressed.
Spongiform appendages absent from petiole, present on postpetiole as a small ventral lobe and a narrow
posterior collar. Transverse basal strip on first gastral tergite reduced to a narrow rim or carina from which
the fine basigastral costulae arise. Petiole, postpetiole and gaster with stout hairs which increase in
thickness from base to apex. Colour dull yellow to light yellowish brown.
This relatively large species is arboreal, nesting in rot holes in the trunks and branches of trees. It
is immediately separated from all its Afrotropical congeners by its unique dentition and strongly
excavated upper scrobe margins.
MATERIAL EXAMINED
Ghana: Tafo (D. Lesion); Tafo (C. A. Collingwood); Tafo (B. Bolton); Tafo (A. B. S. King). Nigeria:
Gambari (B. Bolton).
Strumigenys dextra Brown
Strumigenys dextra Brown, 1954: 27. Holotype and paratype workers, UGANDA: 5 miles (8 km) N.
Kamapala, Kawanda Exp. Sta., 15. ii. 1949, no. SS 30, soil sample under elephant grass (G. Salt) (MCZ)
[examined].
WORKER. TL 1-6-1-8, HL 0-41-0-47, HW 0-31-0-34, CI 71-77, ML 0-16-0-19, MI 38-42, SL 0-22-0-25, SI
68-75, PW 0-21-0-24, AL 0-40-0-48 (10 measured).
Mandibles relatively slender, shallowly and evenly curved along the outer margins. Apical fork of each
mandible of two teeth, without intercalary teeth or denticles. Preapical armament of left mandible of a
single tooth, the right mandible with 2 preapical teeth. Upper scrobe margins concealing the eyes in
full-face view. Eyes small, with only 3-5 ommatidia, the maximum diameter of the eye distinctly less than
the maximum width of the scape. Preocular notch absent, the ventral surface of the head without a
transverse preocular groove or impression on each side. Antennal scape shallowly curved basally,
somewhat expanded in the median third, the leading edges with a row of apically curved narrow
spoon-shaped hairs. Ground-pilosity of cephalic dorsum reduced, consisting of a few inconspicuous small
spatulate hairs. Upper scrobe margins with a triple row of larger narrowly spoon-shaped hairs. Dorsum of
head with 6 standing hairs arranged in a transverse row of 4 close to the occipital margin and a more
anteriorly situated pair. Pronotal humeri each with a single fine flagellate hair. Mesonotum with a single
pair of standing stout hairs which are clavate apically. Ground-pilosity of dorsal alitrunk of inconspicuous
narrowly spatulate hairs like those on dorsum of head. In profile the mesonotum feebly or not depressed
behind the level of the standing hairs. Propodeal teeth short and subtended by moderately developed
infradental lamellae. Sides of pronotum almost smooth to weakly longitudinally rugulose. Pleurae and
sides of propodeum smooth except for weak peripheral punctures. Dorsal alitrunk with sparse widely
separated longitudinal rugulae on pronotum, the spaces between the rugulae smooth or with vestiges of
superficial sculpture. Remainder of dorsal alitrunk reticulate-punctate. Dorsum of petiole node punctate,
the postpetiole smooth. Spongiform appendages of pedicel segments only moderately developed. In
profile the petiole with a narrow ventral strip and small lateral appendage on the node. Ventral spongiform
lobe of postpetiole equal to or slightly smaller than the exposed area of the postpetiolar disc in profile.
Basigastral costulae short but sharply defined, arising across the width of the first tergite basally, not
radiating on each side of a broad central clear area. Dorsal surfaces of petiole, postpetiole and gaster with
stout standing hairs which are narrowly clavate apically. Colour dull yellow.
In the arnoldi-complex three other species beside dextra have lost the distal preapical tooth of
the left mandible, irrorata, katapelta andparanax. The first of these is distinguished from dextra
THE AFROTROPICAL DACETINE ANTS 369
and the rest by also lacking the distal preapical tooth on the right mandible so that both blades
have only a single preapical tooth. S. katapelta is the only species of the four which possesses
intercalary teeth between the spiniform teeth of the apical forks, andparanax is easily separated
from dextra by its possession of a straight stout projecting hair at the humeri where dextra has a
long fine flagellate hair present.
Elsewhere in the genus bernardi, sarissa and londianensis also have only a single preapical
tooth on the left mandible, but in all of these there is a large and very distinct preocular notch
present.
MATERIAL EXAMINED
Cameroun: nr Yaounde (G. Terron). Gabon: He aux Singes (J. A. Band). Angola: Dundo. Uganda:
Kampala, Kawanda Exp. Sta. (G. Salt); Busnia (A/. A. Weber). Central African Republic: Haut Mbomu
(A/. A. Weber).
Strumigenys dromoshaula sp. n.
HOLOTYPE WORKER. TL 2- 1 , HL 0-55, HW 0-41 , CI 75 , ML 0-27, MI 49, SL 0-29, SI 70, PW 0-27, AL 0-54.
Mandibles in full-face view stout and shallowly bowed outwards. Apical fork of each mandible of
2 spiniform teeth, without intercalary teeth or denticles. Blade of each mandible with 2 preapical teeth, the
proximal much the largest in each case and the space separating the preapical teeth shorter than the length
of the distal tooth; both preapical teeth situated in the apical third of the length of the blade. Upper scrobe
margins bordered by a narrow rim or flange throughout their length, evenly divergent posteriorly and more
or less straight rather than sinuate. Eyes of moderate size, the maximum diameter about 0-18xHW. In
full-face view the eyes plainly visible and their maximum length distinctly greater than the maximum width
of the scape. Preocular notch conspicuous and strongly developed, the anterior portion of the eye detached
from the side of the head. Preocular notch extended onto ventral surface of head as a broad impression
whose width is about equal to or slightly larger than the maximum diameter of the eye and whose margins
are rounded and not sharply defined. Antennal scapes very weakly bent in the basal third and somewhat
thickened in the median third, the leading edges with an apically curved row of narrow spoon-shaped hairs.
Ground-pilosity of cephalic dorsum consisting of inconspicuous spatulate to spoon-shaped minute hairs,
the upper scrobe margins fringed with a row of much larger very distinctive spoon-shaped hairs which are
curved anteriorly. Dorsum of head posteriorly with 6 standing hairs arranged in a transverse row of 4 close
to the occipital margin and a more anteriorly situated pair. In the holotype this anteriorly situated pair,
close to the highest point of the vertex, is flattened to the surface, but erect in the paratypes. Dorsum of
head finely and densely reticulate-punctate. Pronotal humeri each with a single fine flagellate hair, the
mesonotum with a single pair of stout standing hairs. Ground-pilosity of dorsal alitrunk of minute flattened
hairs which are closely applied to the surface. Posterior portion of mesonotum slightly depressed behind
the level of the pairs of hairs; metanotal groove forming a line across the surface but not impressed.
Propodeal teeth broadly triangular and lamellate, sharply elevated and subtended by narrow infradental
lamellae. Sides of pronotum showing vestigial striolate sculpture, the pleurae smooth except for some
peripheral punctation, which is best developed on the uppermost parts. Sides of propodeum finely
punctate. Pronotal dorsum finely longitudinally rugulose with a few punctures visible laterally but the
remainder of the dorsal alitrunk reticulate-punctate. Petiole node faintly punctulate dorsally, the sculpture
almost effaced; postpetiole smooth. Spongiform appendages of pedicel segments moderately developed.
Subpetiolar process consisting of a thickened cuticular longitudinal ridge which is shallowly concave
ventrally and from which a narrowly lunate spongiform strip is dependent. Ventral spongiform lobe of
postpetiole larger than lateral lobe, about as large as the exposed area of the postpetiolar disc in profile. In
dorsal view the postpetiole node with a broad lamellate strip posteriorly which abuts a similar but narrower
strip across the base of the first gastral tergite. Petiole node distinctly broader than long in dorsal view.
Basigastral costulae arising from the lamellate strip on each side of a central clear area. Petiole, postpetiole
and gaster with stout standing hairs which are somewhat thickened apically. Colour brownish yellow, the
gaster dark brown.
PARATYPE WORKERS. TL 2-1, HL 0-54-0-55, HW 0-40-0-41, CI 74-75, ML 0-26-0-27, MI 48-49, SL
0-28-0-29, SI 68-73, PW 0-27, AL 0-52 (2 measured).
As holotype but sides of pronotum may be more obviously striolate.
Holotype worker, Burundi: Bugarama, 2200 m, 1976-7, under Eucalyptus (A. Dejean) (MCZ).
Paratypes. 2 workers with same data as holotype (MCZ; BMNH).
Within the 13 species of the faurei-complex six have the preocular notch extended onto the
370 BARRY BOLTON
ventral surface of the head as a transverse groove or impression. Of the six pretoriae and
petiolata lack flagellate hairs at the pronotal humeri. 5. faurei has the postpetiole sculptured. 5.
shaula is separated from dromoshaula by the key character concerning the shape of the ventral
extension of the preocular notch, and rufobrunea is distinguished by its dental characteristics as
the distal preapical tooth is short and widely separated from the proximal.
Strumigenys dyshaula sp. n.
(Fig. 55)
HOLOTYPE WORKER. TL 2-0, HL 0-53, HW 0-40, CI 75, ML 0-25, MI 47, SL 0-28, SI 70, PW 0-26, AL 0-50.
Mandibles in full-face view with the outer margins shallowly convex. Apex of each mandible with a fork
of two spiniform teeth, without intercalary teeth or denticles. Preapical armament of 2 teeth on each
mandible, situated in the apical third of the length of the blade, the proximal preapical tooth distinctly
larger than the distal. Upper scrobe margins evenly divergent posteriorly, shallowly convex and bordered
by a thin projecting rim or flange. Eyes visible in full-face view, not wholly hidden by the upper scrobe
margins. Preocular notch present but only weakly developed, merely an impression in the side of the head
immediately in front of the eyes; the anterior portion of the eye not detached from the side of the head.
Preocular notch ending below at the ventrolateral cephalic margin, not extending onto the ventral surface
of the head as a transverse groove or impression. Maximum diameter of eye greater than the maximum
width of the scape. Antennal scapes very shallowly curved in the basal third, broadest just distal of the
midlength, the leading edges with a row of narrow spoon-shaped hairs which are curved apically and are
distinctly smaller than those on the upper scrobe margins. Occipital margin of head narrowly and deeply
impressed in full-face view. Ground-pilosity of cephalic dorsum consisting of inconspicuous minute
spatulate hairs which are closely applied to the surface. Upper scrobe margins with a row of large
spoon-shaped hairs which are curved anteriorly and very distinct. Dorsum of head with 6 standing hairs
arranged in a transverse row of 4 close to the occipital margin, and a more anteriorly situated pair. Head
reticulate-punctate. Pronotal humeri each with a fine flagellate hair. Mesonotum with a single pair of
standing hairs. Ground-pilosity of dorsal alitrunk of sparse closely applied hairs similar to those on the
head. Posterior portion of mesonotum depressed behind the level of the standing hairs. Metanotal groove
forming a line across the dorsum but not impressed in profile. Propodeal teeth triangular and subtended by
infradental lamellae. Pleurae mostly smooth, with weak peripheral punctulation which is best developed
laterodorsally. Sides of propodeum with a fine punctulate strip above and below the spiracle, otherwise
smooth. Sides of pronotum with vestigial striolate and punctulate sculpture. Pronotal dorsum with feeble
longitudinal costulae or striae, the remainder of the dorsal alitrunk finely reticulate-punctate. Dorsum of
petiole node broader than long and superficially punctate, the postpetiole smooth. Spongiform appen-
dages of pedicel segments moderately developed. In profile the petiole with a shallow ventral process
which is about half as deep as the depth of the peduncle at its midlength. Lateral lobe of petiole node
bluntly triangular. Ventral and lateral spongiform lobes of postpetiole about equal in size, the former
about equal to the exposed portion of the postpetiolar disc in profile. In dorsal view the postpetiole with a
lamellar posterior strip which abuts a similar strip traversing the base of the first gastral tergite. Basigastral
costulae dense and sharply defined, radiating from each side of a central clear area which is free of costulae.
Secondary costulae present which arise between the main costulae, these latter originating on the basal
strip and the secondaries arising some distance behind it. Petiole , postpetiole and gaster with stout standing
hairs which are broadened to narrowly clavate apically. Colour dull yellow.
PARATYPE WORKERS. TL 2-0-2-1, HL 0-54-0-56, HW 0-40-0-43, CI 74-78, ML 0-25-0-27, MI 46-48, SL
0-28-0-31, SI 70-74, PW 0-26-0-28, AL 0-50-0-54 (7 measured). As holotype.
Holotype worker, Zimbabwe: Victoria Falls, spray forest, rotten wood M440, 7.iii.l969 (W. L. Brown)
(MCZ).
Paratypes. 8 workers and 1 female with some data as holotype (MCZ; BMNH; MHN).
Of the species possessing a preocular notch and relatively large eyes a number of forms do not
have the preocular notch extended onto the ventral surface of the head as a transverse
impression. Among the six species in this category dyshaula is distinguished by having a
postpetiole whose sides are not completely lapped around by spongiform tissue, having the
ventral lobe of the postpetiole well developed, having six standing hairs on the cephalic dorsum
and having basigastral costulae which do not arise as a regular row across the width of the tergite
but rather radiate out from each side of a central clear area.
THE AFROTROPICAL DACETINE ANTS 371
Strumigenys ettillaxsp. n.
HOLOTYPE WORKER. TL 2-0, HL 0-57, HW 0-46, CI 81 , ML 0-23, MI 40, SL 0-27, SI 59, PW 0-30, AL 0-52.
Mandibles stout, the outer margins shallowly convex in full-face view and sharply incurved basally where
the blades suddenly narrow to their insertions. Apical fork of each mandible with 2 spiniform teeth,
without intercalary teeth or denticles. Each mandibular blade with 2 stout preapical teeth, the proximal
about one-third longer than the distal and their bases separated by a distance which is slightly less than the
length of the distal preapical tooth. Upper scrobe margins very feebly sinuate in full-face view, bordered by
a narrow rim or flange and meeting the sides of the occipital lobes (at the scrobal apices) in an obtuse but
distinct angle, the upper scrobe margins not merging smoothly into the sides of the occipital lobes. Eyes of
moderate size, with 5 ommatidia across the greatest diameter and with 15 or more ommatidia in all.
Maximum diameter of eye greater than the maximum width of the scape. Preocular notch absent, ventral
surface of head without a transverse preocular groove or impression. Antennal scapes narrow and
shallowly curved basally, expanded in the median third and with the leading edges equipped with a row of
apically curved spoon-shaped hairs which are distinctly smaller than those fringing the upper scrobe
margins. Ground-pilosity of cephalic dorsum of inconspicuous small flattened hairs, the upper scrobe
margins with a row of large spoon-shaped hairs. Dorsum of head with 6 standing hairs, arranged in a
transverse row of 4 close to the occipital margin and a more anteriorly situated pair. Dorsum of head
densely reticulate-punctate. Pronotal humeri each with a fine flagellate hair. Mesonotum with a single pair
of stout standing hairs which are thickened apically. Ground-pilosity of dorsal alitrunk of inconspicuous
small hairs like those on the head. Posterior portion of mesonotum very slightly depressed behind the level
of the pair of hairs, the metanotal groove forming a weak line across the dorsum. Propodeal teeth short and
triangular, subtended by relatively broad infradental lamellae which at their broadest extend posteriorly
almost as far as the apices of the teeth . Sides of pronotum feebly punctate-striolate , the pleurae and sides of
propodeum mostly smooth but with some peripheral punctate sculpture. Pronotum finely and quite
densely longitudinally rugulose, the remainder of the dorsal alitrunk reticulate-punctate. Dorsum of
petiole node finely punctate, the postpetiole smooth. Ventral spongiform strip of petiolar peduncle
narrow, broadening beneath the node. Lateral spongiform lobe of postpetiole equal to or slightly
exceeding the exposed area of the disc in profile. In dorsal view the lateral spongiform material is visible,
projecting beyond the outline of the sides of the postpetiole. Basigastral costulae sparse, 5-6 arising on
each side of a narrow clear central area. Petiole, postpetiole and gaster dorsally with stout standing hairs
which are thickened apically. Colour blackish brown.
PARATYPE WORKERS. TL 2-0, HL 0-53-0-54, HW 0-43-0-44, CI 81, ML 0-23, MI 42^3, SL 0-26, SI 59-60,
PW 0-27, AL 0-50-0-52 (2 measured).
As holotype but one lighter in colour than the holotype and probably teneral.
Holotype worker, Cameroun: nr Yaounde, series IY (G. Terron) (ENSA).
Paratypes. 2 workers with same data as holotype but one series AM, the other series TR (ENSA;
BMNH).
Strumigenys faurei Arnold sp. rev.
Strumigenys faurei Arnold, 1948: 226, figs 12, 12a. Syntype workers and females, SOUTH AFRICA: Natal,
Zululand, Sordwana, 21. v. 1946 (J. C. Faure) (SAM) [examined]. [Previously synonymized with
rufobrunea by Brown, 1954: 17.]
WORKER. TL 2-2-2-3, HL 0-55-0-56, HW 0-43-0-44, CI 77-79, ML 0-25-0-26, MI 46-47, SL 0-30-0-32, SI
70-72, PW 0-25-0-27, AL 0-55-0-57 (6 measured).
Outer margins of mandibles shallowly convex, the blades weakly bowed outwards in full-face view.
Apical fork of each mandible with 2 spiniform teeth, without intercalary teeth or denticles. Preapical
armament of 2 teeth on each blade, the proximal spiniform and much larger than the distal. Length of left
distal preapical tooth about equal to the distance separating the bases of the preapical teeth. Upper scrobe
margins bordered by a narrow rim or flange, not concealing the eyes in full-face view. Maximum diameter
of eye slightly greater than the maximum width of the scape. Preocular notch strongly developed, the
anterior portion of the eye detached from the side of the head. Preocular notch continued onto ventral
surface of head as a broad shallow impression whose margins are quite sharply defined close to the eye but
are more rounded and indistinct elsewhere; the maximum width of the impression the same as or slightly
more than the maximum diameter of the eye. Antennal scapes very feebly bent in the basal third, broadest
at about the midlength and the leading edges equipped with a row of apically curved spatulate to narrowly
spoon-shaped hairs which are distinctly smaller and finer than those on the upper scrobe margins.
Ground-pilosity of head of minute spatulate hairs which are curved, inconspicuous and closely applied to
372 BARRY BOLTON
the surface. Upper scrobe margins with a row of projecting large anteriorly curved narrowly spoon-shaped
hairs. Dorsum of head with 6 standing hairs arranged in a transverse row of 4 close to the occipital margin
and a more anteriorly situated pair. Dorsum of head reticulate-punctate. Pronotal humeri each with a
single fine flagellate hair. Mesonotum with a pair of stout standing hairs. Ground-pilosity of alitrunk similar
to that on the head but the hairs sparse. Posterior half of mesonotum depressed behind the level of the pair
of hairs and with a shallow transverse impression at the base of the slope, behind which the remainder of
the mesonotum rises slightly to the level of the propodeum. Metanotal groove forming a transverse line on
the dorsum, extremely weakly impressed. Propodeal teeth triangular and subtended by an infradental
lamella. Sides of pronotum feebly punctate. Pleurae smooth except for sparse peripheral punctures and a
patch on the upper half of the mesopleuron which is punctate. Pronotal dorsum finely longitudinally
rugulose with weak punctures between the rugulae posteriorly. Remainder of dorsal alitrunk reticulate-
punctate. Petiole node in dorsal view finely punctate, twice broader than long. Postpetiole with the disc
finely and densely longitudinally costulate except for the posteromedian area which is smooth. Subpetiolar
process reduced to a very thin laminar strip. Ventral spongiform lobe of postpetiole slightly larger than the
lateral lobe but smaller than the exposed area of the postpetiolar disc in profile. In dorsal view the
postpetiole with a thin transverse laminar strip posteriorly which abuts a similar but even narrower strip
across the base of the first gastral tergite from which the fine but sharply defined basigastral costulae arise.
Petiole, postpetiole and gaster with standing hairs which are broadened apically. Colour dull yellow.
Close to the West African rufobrunea but slightly larger and evenly yellow in colour, faurei also
differs by having the postpetiole densely costulate, the subpetiolar process very reduced, the
petiole node twice broader than long dorsally, and the ventral preocular impression broader and
less sharply defined.
Included by Brown (1954) as a synonym of rufobrunea, I consider faurei to be sufficiently
distinct to be regarded as a separate species on the strength of the characters noted above.
MATERIAL EXAMINED
South Africa: Natal, Sordwana (/. C. Faure); St Lucia Lake (J. C. Faure); Richards Bay (J. C. Faure);
Dukuduku Forest Res. (W. L. & D. E. Brown).
Strumigenys hastyla sp. n.
HOLOTYPE WORKER. TL 2-0, HL 0-52, HW 0-38, CI 73, ML 0-23, MI 44, SL 0-29, SI 76, PW 0-24, AL 0-52.
Mandibles slender in full-face view, the outer margins evenly and very shallowly curved, the blades only
slightly tapering from their broadest point near the base to the apex. Apical fork of each mandible with 2
spiniform teeth, without intercalary teeth or denticles. Each mandible with 2 preapical teeth which are
stout and situated within the apical quarter of the length of the blade , the proximal tooth slightly longer and
thicker than the distal. The preapical teeth on each blade close together so that the length of the distal tooth
is more than twice the distance which separates their bases. Upper scrobe margins shallowly divergent and
evenly curved, confluent with the sides of the occipital lobes through an even smooth curve in full-face
view. Eyes quite large, with about 15 ommatidia, the maximum diameter of the eye distinctly greater than
the maximum width of the scape. Preocular notch absent, the ventral surface of the head without a
preocular transverse groove or impression on each side. Antennal scapes slender and almost cylindrical,
very shallowly curved in the basal third, their leading edges with a row of apically curved slender hairs
which are somewhat flattened to very feebly and narrowly spoon-shaped , smaller than the hairs fringing the
upper scrobe margins. Cephalic ground-pilosity of numerous narrowly spatulate to slender spoon-shaped
hairs, the upper scrobe margins fringed with a row of similar or very slightly larger hairs. Dorsum of head
with 6 standing hairs, arranged in a transverse row of 4 close to the occipital margin and a more anteriorly
situated pair. Dorsum of head reticulate-punctate. Pronotal humeri each with a single fine flagellate hair.
Mesonotum with a single pair of standing hairs. Ground-pilosity of dorsal alitrunk like that of head but the
hairs smaller and more widely scattered. Mesonotum shallowly depressed behind the level of the pair of
hairs. Propodeal teeth triangular and subtended by conspicuous infradental lamellae. Sides of pronotum
weakly rugulose. Pleurae and sides of propodeum mostly smooth but with punctate areas around the
periphery. Pronotum dorsally finely longitudinally rugulose, the spaces between the rugulae superficially
punctate. Remainder of dorsal alitrunk reticulate-punctate. Dorsum of petiole node reticulate-punctate,
the propodeum smooth. Spongiform appendages of pedicel segments well developed, the petiole with a
broad ventral strip and a distinct lateral lobe on the node, the postpetiole with large ventral and lateral
spongiform lobes. In dorsal view the disc of the postpetiole appears surrounded by spongiform tissue on all
sides as the lateral and ventral lobes project beyond the outline of the disc. Basigastral costulae arise on the
first tergite on each side of a central clear area. Dorsal surfaces of petiole, postpetiole and gaster with
THE AFROTROPICAL DACETINE ANTS 373
standing hairs, some or all of which are thickened or flattened apically in dorsal view. Colour brownish
yellow.
PARATYPE WORKERS. TL 1-8-2-0, HL 0-49-0-52, HW 0-36-0-38, CI 73-75, ML 0-22-0-23, MI 43-45, SL
0-28-0-29, SI 76-78, PW 0-22-0-24, AL 0-48-0-52 (3 measured). As holotype.
Holotype worker, Burundi: Bujumbura, 1977, no. 17 (A. Dejean) (BMNH).
Paratypes. 3 workers with same data as holotype (BMNH; MCZ).
Non-paratypic material examined. Ivory Coast: Mongaga (V. Mahnert & J.-L. Ferret); Dropleu
(V. Mahnert & J.-L. Ferret); Man (V. Mahnert & J.-L. Ferret); Banco Forest (/. Lobl); Anguededou (W.
L. Brown); Nzi Noua (W. L. & D. E. Brown). Nigeria: Gambari (B. Bolton). Cameroun: Victoria (B.
Malkiri); nr Yaounde (G. Tenon). Gabon: Plateau d'Ipassa (J. A. Barra). Angola: Gubela (P. Hammond);
Dundo (L. de Carvalho). Burundi: Imbo Plain (A. Dejean).
Variation in this non-paratypic material is as follows. TL 2-0-2-1, HL 0-48-0-54, HW 0-34-0-39,
CI 70-74, ML 0-21-0-24, MI 42-46, SL 0-27-0-31, SI 76-82, PW 0-22-0-25, AL 0-48-0-56 (18
measured). The pronotum in some is wholly reticulate-punctate dorsally, the fine longitudinal
rugulae seen in the type-series being suppressed. Much of the West African material is darker in
colour, medium brown, so that the cephalic ground-pilosity is more conspicuous.
5. hasty la is one of three very closely related species in the scorn-complex. Along with scotti
and zandala, hasty la is characterized by the form of the mandibles, the elongate almost
cylindrical scapes, the relatively slender fringing hairs on the upper scrobe margins (which are
only slightly or not at all larger than the cephalic ground-pilosity), the large eyes, lack of a
preocular notch and ventral preocular groove or impression, and evenly rounded sides of the
head which round into the upper scrobe margins without trace of an angle. These three species
are very similar and difficult to separate . 5. scotti is the largest of the three and has relatively long
mandibles and scapes; it also tends to have the petiole node longer than broad in dorsal view
whereas in the other two the node is very obviously broader than long. The measurements
compare as follows.
HL HW MI SI
scotti 0-62-0-70 0-42-0-46 46-50 88-95
zandala 0-59-0-61 0-42-0-44 43-45 75-81
hastyla 0-48-0-54 0-34-0-39 42-46 76-82
As can be seen, hastyla is the smallest species of the three in absolute terms. It also differs from
zandala as the gastral hairs are simple in the latter but mostly flattened or thickened apically in
hastyla.
Strumigenys havilandi Forel
(Figs 65, 70)
Strumigenys havilandi Forel, 1905: 13. Syntype workers, SOUTH AFRICA: Natal, 5300 ft (1615 m) (Haviland)
(MHN; BMNH) [examined].
Strumigenys havilandi Forel; Brown, 1954: 25.
WORKER. TL 2-5-2-6, HL 0-62-0-68, HW 0-44-0-50, CI 69-74, ML 0-28-0-33, MI 45-50, SL 0-36-0-40, SI
80-90, PW 0-28-0-32, AL 0-62-0-68 (13 measured).
Mandibles in full-face view slender and almost straight, broadest basally and evenly tapering towards the
apex. Apical fork of each mandible with 2 spiniform teeth, without intercalary teeth or denticles. Each
mandibular blade with 2 spiniform preapical teeth, the distal only slightly shorter than the proximal and the
distance separating their bases distinctly less than the length of the distal preapical tooth. Eyes small, with
only 4-5 ommatidia, their maximum diameter much less than the maximum width of the scape. Preocular
notch absent, the ventral surface of the head without a transverse preocular groove or impression.
Antennal scapes elongate and relatively narrow, SI 80-90, matched only by korahyla in the arnoldi-
complex. The scapes only shallowly and very gently curved, slightly expanded beyond the curve and with
their leading edges having an apically directed row of narrow spoon-shaped hairs which are about the same
size as those fringing the upper scrobe margins. Ground-pilosity of cephalic dorsum of numerous but
inconspicuous narrowly spoon-shaped hairs, the upper scrobe margins fringed by a row of hairs which are
the same as those on the dorsum of the head. Occipital margin dorsally with a transverse row of 4 short
374 BARRY BOLTON
curved standing hairs and a pair of similar but even shorter hairs situated anterior to this row. Dorsum of
head reticulate-punctate. Pronotal humeri each with a single fine flagellate hair. In this species the hair
appears to be very delicate and easily lost by abrasion. Mesonotum with a single pair of standing hairs.
Ground-pilosity of dorsal alitrunk of numerous small curved spatulate to spoon-shaped hairs similar to
those on the head but slightly smaller. Metanotal groove a transverse line across the dorsum, weakly
impressed in profile. Propodeal teeth slender and triangular, subtended by a broad infradental lamella on
each side. Sides of alitrunk smooth, with only peripheral punctation present. Pronotal dorsum finely
superficially longitudinally rugulose, the rugulae low and inconspicuous, frequently with faint punctures
between them. Remainder of dorsal alitrunk finely reticulate-punctate. Petiole node reticulate-punctate
dorsally, the postpetiole smooth. Postpetiole distinctly swollen and inflated, subglobular. Spongiform
appendages of petiole consisting of a fairly broad ventral strip and a narrow posterior collar. Postpetiole in
profile with the lateral lobe much reduced, obviously smaller than the ventral spongiform lobe, and the
latter itself relatively small, smaller than the exposed area of the postpetiolar disc in profile. Basigastral
costulae short but distinct. Dorsal surfaces of petiole, postpetiole and gaster with standing short hairs
which are expanded apically. Colour yellow.
In the arnoldi-complex havilandi is characterized by its long scapes, the structure of its
mandibles and the form of its inflated postpetiole and spongiform appendages. It resembles the
two closely related species traegaordhi and mesahyla but, apart from the mandibular character
quoted in the key, may be separated from both by its long scapes and form of the postpetiole and
its appendages; compare Figs 69, 70.
MATERIAL EXAMINED
South Africa: Natal (Haviland); Natal, Gillitts (W. L. & D. E. Brown).
Strumigenys helytruga sp. n.
HOLOTYPE WORKER. TL 2-2, HL 0-59, HW 043, CI 73, ML 0-29, MI 49, SL 0-34, SI 79, PW 0-26, AL 0-55.
In full-face view the mandibular blades approximately straight. Apical fork of each mandible with 2
spiniform teeth, without intercalary teeth or denticles. Preapical armament of 2 teeth on each blade, the
proximal only fractionally longer than the distal and with both teeth situated in the apical third of the length
of the blade. Upper scrobe margins evenly divergent posteriorly and without a bordering rim or flange,
posteriorly grading evenly into the sides of the head. Preocular notch vestigially present, represented only
by a feeble indentation of the ventrolateral margin immediately in front of the eye, the notch not extended
onto the ventral surface of the head as a transverse groove or impression. Eye relatively small, about
0-12xHW and with 8 ommatidia, the maximum diameter of the eye about equal to the maximum width of
the scape or fractionally larger. Antennal scapes slender and roughly cylindrical, shallowly curved basally
and with their leading edges equipped with a row of apically curved narrow spatulate hairs. Ground-
pilosity of head consisting of quite dense conspicuous spatulate hairs which are broader than those on the
leading edges of the scapes. Upper scrobe margins without larger hairs but fringed with spatulate hairs the
same size and shape as those on the dorsum. Elongate standing hairs on cephalic dorsum restricted to a
single pair situated close to the midline near the occipital margin. Dorsum of head reticulate-punctate.
Pronotal humeri each with a single fine flagellate hair. Mesonotum with a single pair of standing stout hairs.
Ground-pilosity of dorsal alitrunk of sparse narrowly spatulate hairs. Posterior portion of mesonotum
shallowly depressed behind the level of the hairs. Metanotal groove represented by a feebly impressed line.
Propodeal teeth triangular and subtended by conspicuous infradental lamellae. Sides of alitrunk mostly
smooth but with some punctate sculpture dorsally and posteriorly. Pronotal dorsum longitudinally sparsely
rugulose, with feeble superficial punctures between the rugulae. Mesonotum and propodeum densely
punctate. Dorsum of petiole node punctate, the postpetiole smooth. In profile the petiole with a
well-developed spongiform ventral strip and a narrow lateral lobe. Postpetiole with large ventral and
lateral lobes. In dorsal view the petiole node with a broad posterior spongiform strip, the postpetiole with
spongiform material projecting beyond the sides and posteriorly with a laminar transverse strip connecting
the spongiform lateral lobes. Transverse basal strip of first gastral tergite laminar, the basigastral costulae
arising from it sharply defined but sparse and short, with only 4 or 5 on each side of a central clear area.
Petiole, postpetiole and gaster dorsally with stout standing hairs which are thickened apically. Colour dull
brownish yellow.
Holotype worker, Angola: Bruco, 26.ii.-2.iii.1972, forest litter (P. Hammond) (BMNH).
A member of the scorn-complex, helytruga is separated from its four close relatives (scotti,
hastyla, murshila and zandala) by its retention of a vestigial preocular notch and relatively very
THE AFROTROPICAL DACETINE ANTS 375
small eyes. Of the five species only murshila has eyes which approach the small size seen in
helytruga, but here the preocular notch is absent, the mandibles are conspicuously bowed
outwards, the cephalic dorsum has six standing hairs, the postpetiole is sculptured and the distal
preapical tooth of the left mandible is less than half the length of the proximal.
Strumigenys irrorata Santschi
Strumigenys irrorata Santschi, 1913a: 257 (diagnosis in key). Holotype worker, SOUTH AFRICA: Natal,
Zululand, Lake Sibayi (/. Tragardh) (NMB) [examined].
Strumigenys irrorata Santschi; Santschi, 1914c: 29, fig. 5 (description).
Strumigenys irrorata Santschi; Brown, 1954: 33.
WORKER. TL 1-9-2-1, HL 0-47-0-54, HW 0-38-0-43, CI 76-83, ML 0-19-0-23, MI 40-43, SL 0-24-0-28, SI
62-70, PW 0-24-0-27, AL 0-50-0-56 (10 measured).
Apical fork of each mandibular blade with 2 spiniform teeth, without intercalary teeth or denticles.
Preapical armament of a single spiniform tooth on each blade which corresponds to the proximal preapical
tooth in related species; the distal preapical teeth lost. Upper scrobe margins strongly divergent behind the
frontal lobes, the eyes not visible in full-face view. Eyes very small, conspicuously much smaller than the
maximum width of the scape. Preocular notch absent, the ventral surface of the head without a preocular
transverse groove or impression. Scapes short, weakly curved, their anterior margins shallowly convex and
equipped with a row of short spoon-shaped to scale-like hairs. Ground-pilosity of cephalic dorsum of dense
short spoon-shaped hairs which are broad and appear scale-like in full-face view. Hairs fringing the upper
scrobe margins the same as those on the dorsum, and about equal in size to the projecting hairs on the
leading edges of the scapes . Dorsum of head with 6 standing hairs , arranged in a transverse row of 4 close to
the occipital margin and a more anteriorly situated pair. Dorsum of head densely reticulate-punctate.
Pronotal humeri each with a single long fine flagellate hair. Mesonotum with a single pair of stout standing
hairs which are weakly clavate apically. Ground-pilosity of dorsal alitrunk of sparse small hairs which are
spatulate to narrowly spoon-shaped. Propodeal teeth subtended by broad infradental lamellae. Metanotal
groove not impressed. Sides of alitrunk mostly smooth, sometimes with vague traces of sculpture on the
pronotal sides and usually with weak punctulae on the pleurae and propodeum. Pronotal dorsum
longitudinally rugulose, the rest of the dorsal alitrunk punctate. Dorsum of petiole node punctate, the
postpetiole smooth and shining. Spongiform appendages of pedicel segments well developed, the petiole
with a broad ventral spongiform strip and the postpetiole with large ventral and lateral lobes. Basigastral
costulae short but sharply defined. Dorsal surfaces of petiole, postpetiole and gaster with stout standing
hairs which are weakly clavate apically. Colour medium to dark brown.
Immediately recognized by its unique (in Africa) preapical dentition, irrorata is the only
member of the rogeri-group having a single preapical tooth on each mandibular blade. The usual
count in the group is 2 preapical teeth on each blade but a few species have one left and two right
preapical teeth.
MATERIAL EXAMINED
Zimbabwe: Umtali, Melsetter (R. Mussard). South Africa: Natal, Lake Sibayi (/. Tragardh); Gillitts, nr
Durban (W. L. &. D. E. Brown); Town Bush, nr Pietermaritzburg (W. L. & D. E. Brown).
Strumigenys katapelta sp. n.
(Figs 61, 76)
HOLOTYPE WORKER. TL 2- 1 , HL 0-51 , HW 0-46, CI 90, ML 0-20, MI 39, SL 0-27, SI 59, PW 0-27, AL 0-56.
Mandibles short and conspicuously bowed outwards in full-face view. Apical fork of each mandible
consisting of a long spiniform tooth dorsally, a smaller spiniform tooth ventrally and an intercalary denticle
between the two longer teeth. Blade of left mandible with a single long spiniform preapical tooth, the
proximal; right mandible with a -similar proximal preapical tooth and with a very small distal preapical
tooth also present. Head broad, the upper scrobe margins strongly divergent and the eyes not visible in
full-face view. Eyes small, with only 3-4 ommatidia, their maximum diameter distinctly much less than the
maximum width of the scape. Preocular notch absent, the ventral surface of the head without a preocular
transverse groove or impression. Antennal scapes short and curved, their anterior margins flattened and
expanded in the median third, broadest at the midlength; their convex leading edges with a row of
shallowly spoon-shaped to flattened large broad hairs. Cephalic ground-pilosity of inconspicuous short
stubby flattened hairs which are closely applied to the surface; without the scale-like or broadly
376 BARRY BOLTON
spoon-shaped hairs usually seen in the arnoldi-complex. Upper scrobe margins fringed by similar small
hairs, these hairs distinctly very much smaller than those on the leading edges of the scapes. Dorsum of
head with a row of 4 standing hairs close to the occipital margin, without a further pair of hairs situated
close to the highest point of the vertex. Dorsum of head reticulate-punctate. Pronotal humeri without
flagellate hairs, instead each with a projecting straight stout hair which is thickened and flattened apically,
directed laterally and slightly elevated. Mesonotum with a single pair of stout standing hairs. Ground-
pilosity of dorsal alitrunk of small hairs similar to those on the head. Metanotal groove represented by a
line across the dorsum. Propodeal teeth short and triangular, subtended by infradental lamellae whose free
margins are concave. Sides of alitrunk smooth except for weak punctures around the periphery. In dorsal
view the pronotum sharply marginate anteriorly, weakly longitudinally rugulose and with feeble punctures
between the rugulae. Mesonotum reticulate-punctate. Propodeal dorsum smooth anteriorly but punctate
posteriorly and between the teeth. Dorsum of petiole node almost smooth, with only the faintest vestiges of
reticular patterning; postpetiole smooth. Spongiform appendages of pedicel segments well developed. The
petiole in profile with a ventral strip and triangular lateral lobe. Postpetiole with large ventral and lateral
spongiform lobes, the former larger than the exposed area of the postpetiolar disc in profile. In dorsal view
the disc of the postpetiole surrounded on all sides by projecting spongiform tissue. Basigastral costulae
short but sharply defined. Petiole, postpetiole and gaster dorsally with stout standing hairs. Colour
brownish yellow.
PARATYPE WORKERS. TL 2-0-2-2, HL 0-50-0-53, HW 0-44-0-46, CI 88-90, ML 0-20-0-21, MI 38-40, SL
0-26-0-27, SI 57-60, PW 0-25-0-28, AL 0-54-0-56 (4 measured). As holotype.
Holotype worker, Burundi: Bujumbura, 1977, no. 42 (A. Dejean) (BMNH).
Paratypes. 4 workers with same data as holotype (BMNH; MCZ).
Non-paratypic material examined. Kenya: Embu, Kirimiri Forest W. of Runyenje (V. Mahnert & J.-L.
Ferret); Mau Forest (V. Mahnert & J.-L. Ferret).
The non-paratypic material from Kenya answers to the description of the holotype but shows the
following size range. HL 0-52-0-55, HW 0-43-0-48, CI 85-87, ML 0-20-0-23, MI 38-41, SL
0-26-0-30, SI 58-63 (5 measured).
The arnoldi-complex has only four species in which the distal preapical tooth of the left
mandible has been lost, katapelta, irrorata, dextra andparanax. S. katapelta is easily separated
from the other three as it is the only species to have an intercalary denticle between the apical
fork teeth. Apart from this it separates from irrorata as that species has also lost the distal
preapical tooth of the right mandible; from dextra as that species has flagellate hairs at the
pronotal humeri; and from paranax as that species is much smaller, with a narrower head and
longer antennal scapes.
Strumigenys korahyla sp. n.
(Fig. 64)
HOLOTYPE WORKER. TL 2-3, HL 0-62, HW 0-44, CI 71 , ML 0-30, MI 48, SL 0-36, SI 82, PW 0-29, AL 0-60.
Mandibles slender and moderately long, the blades broadening slightly from base to apex in full-face
view, not bowed outwards. Apical fork of each mandible of 2 spiniform teeth, without intercalary teeth or
denticles. Preapical armament of 2 teeth on each mandible, the proximal tooth about one-third longer than
the distal and the distance separating their bases equal to or slightly greater than the length of the distal
preapical tooth. Upper scrobe margins bordered by a broad translucent rim or flange, the eyes not visible in
full-face view. Eyes very small, with only 4-5 ommatidia, their maximum diameter distinctly less than the
maximum width of the scape. Preocular notch absent, the ventral surface of the head without a preocular
groove or impression on each side. Antennal scapes long and slender, approximately straight, very narrow
in the basal eighth then with the anterior margin suddenly broadened. Beyond this the scape evenly
tapering towards the apex. Leading edges of scapes with a series of small apically curved spoon-shaped
hairs which are very obviously much smaller than those on the upper scrobe margins. Dorsum of head from
the posterior clypeal margin to the midlength with large anteriorly curved shallowly spoon-shaped hairs
which appear scale-like in full-face view. Similar but even larger hairs are present fringing the upper scrobe
margins. Behind the midlength of the head the hairs of the cephalic ground-pilosity are much smaller and
contrast with the larger anterior hairs. Dorsum of head with a transverse row of 4 clavate standing hairs
close to the occipital margin, and with a pair of similar but smaller hairs close to the highest point of the
vertex (this abraded in the holotype but present in the paratypes). Dorsum of head densely punctate, the
THE AFROTROPICAL DACETINE ANTS 377
walls of the punctures aligned in places on the occipital lobes and showing as fine rugulae. Pronotal humeri
hairless. Mesonotum with a single pair of clavate hairs. Metanotal groove represented by a faint line across
the dorsum, extremely weakly impressed in profile. Mesonotum slightly depressed behind the level of the
hairs, the base of the declivous portion forming a shallow transverse impression. Propodeal teeth
triangular and acute, subtended by narrow infradental lamellae. Sides of pronotum sculptured, remainder
of sides of alitrunk smooth except for weak peripheral punctures . Pronotal dorsum longitudinally rugulose ,
the remainder of the dorsal alitrunk reticulate-punctate. Dorsum of petiole node reticulate-punctate, the
postpetiole smooth. Spongiform appendages of pedicel segments moderately developed. In profile the
petiole with a straight ventral strip and small lateral appendages. Postpetiole with a small lateral
spongiform lobe and a slightly larger ventral lobe, the latter, however, slightly smaller than the exposed
area of the postpetiolar disc in profile. In dorsal view the postpetiolar disc about l-5x broader than the
petiole node. Basigastral costulae short and widely spaced, but sharply defined. Dorsal surfaces of petiole,
postpetiole and gaster with stout clavate hairs. Colour brownish yellow.
PARATYPE WORKERS. TL 2-2-2-3, HL 0-61-0-64, HW 0-41-0-43, CI 67, ML 0-30-0-32, MI 49-50, SL
0-35-0-36, SI 84-85, PW 0-27-0-28, AL 0-58-0-60 (2 measured). As holotype.
Holotype worker, Cameroun: nr Yaounde, series MT (G. Tenon) (ENSA).
Paratypes. 2 workers with same data as holotype but one series OV, the other series EP (ENSA;
BMNH).
Among the species of the arnoldi-complex, characterized by their small to minute eyes, lack of a
preocular notch and ventral preocular impression, and scale-like cephalic ground-pilosity, only
two species have elongate mandibles and scapes. These two, korahyla and havilandi, compare
with the remaining 11 species of the complex as follows.
MI SI
havilandi 45-50 80-90
korahyla 48-50 82-85
remainder of arnoldi-complex 26-45 52-75
Apart from this korahyla is characterized within the arnoldi-compiex by its complete preapical
dentition of 2 teeth on each mandible, lack of intercalary or adventitious teeth in the mandibular
apical fork, lack of pronotal flagellate hairs at the humeri, slender antennal scapes and hairs on
the leading edges of the scapes which are much smaller than those on the upper scrobe margins.
Strumigenys londianensis (Patrizi)
Proscopomyrmex londianensis Patrizi, 1946: 295, figs 1, 2. Syntype workers, KENYA: Londiani, q. 2260
m.s.m., 4.ix.l943 (5. Patrizi); and Mau Forest, 16.U946 (Meneghetti) (BMNH; MCZ) [examined].
Strumigenys (Proscopomyrmex) londianensis (Patrizi) Arnold, 1948: 227.
Strumigenys londianensis (Patrizi); Brown, 1954: 14.
WORKER. TL 3-5-4-2, HL 0-84-0-92, HW 0-62-0-70, CI 74-77, ML 0-44-0-47, MI 51-52, SL 0-52-0-58, SI
82-87, PW 0-38-0-44, AL 0-82-0-94 (8 measured).
Apical fork of left mandible with a small intercalary tooth between the upper and lower fork teeth; right
apical fork without an intercalary tooth. Blade of left mandible with a single spiniform preapical tooth
present (the proximal) ; blade of right mandible with 2 preapical teeth , a larger proximal and a smaller distal
tooth which is situated close to the apical fork and may be hidden by the opposing left apical fork when the
mandibles are fully closed. Upper scrobe margins not bordered by a continuous projecting lamina, close
together on anterior third of head, the eyes clearly visible in full-face view. Upper scrobe margins concave
immediately behind the convex frontal lobes, with a pinched-in appearance. Behind this the scrobe
margins shallowly concave above the eyes and then diverging posteriorly. Preocular notch deep and
strongly developed, the anterior portion of the eyes detached from the side of the head. Preocular notch
continued onto the ventral surface of the head as a conspicuous broad transverse impression. Antennal
scapes roughly cylindrical, very slightly broadened in the median third and with a characteristic arrange-
ment of strong hairs projecting from the leading edge. The basalmost 1-3 (usually 2) projecting hairs are
curved apically, the next 3-4 are curved basally and the distalmost 3-4 are curved apically. Ground-pilosity
of head short, broadly spatulate to scale-like everywhere and curved anteriorly. In profile the vertex
usually with a single pair of stout standing hairs which are weakly clavate, but these are easily lost by
abrasion. Dorsum of head reticulate-punctate. Pronotal humeri each with a long stout straight hair which is
378
BARRY BOLTON
remiform to weakly clavate apically. Mesonotum with a single pair of shorter stout straight hairs which are
somewhat more strongly clavate apically; the dorsal alitrunk otherwise without standing pilosity but with
sparse narrowly spatulate appressed ground-pilosity. In profile the posterior portion of the mesonotum
sharply depressed below the level of the anterior portion and pronotum, forming a single surface with the
propodeum. Metanotal groove absent. Propodeum without differentiated angular teeth, instead the
infradental lamellae merely bulge slightly and form blunt angles dorsally. Sides of alitrunk mostly punctate
but with some smooth shining areas on the pleurae. Dorsal alitrunk predominantly punctate but the
pronotum generally with a few posteriorly divergent rugulae superimposed on the punctures. Petiole node
weakly punctulate dorsally, the postpetiole generally smooth but sometimes with the weakest vestiges of
punctulate sculpture visible Spongiform appendages of petiole represented by a thin ventral strip and a
narrow posterior collar on the node. Postpetiole in profile with moderately well-developed ventral and
lateral spongiform lobes and in dorsal view with a very narrow anterior and posterior spongiform strip.
Basigastral costulae short and sparse, radiating from the narrow basal spongiform strip of the first tergite.
Petiole, postpetiole and gaster dorsally with stout strong hairs which are clavate apically. Colour light
brown, gaster darker.
A relatively large and easily recognized species, londianensis is known only from Kenya.
Together with its close relative sarissa, londianensis is characterized by its distinctive mandibular
dentition, deep preocular notch and detached dye. The only species coming close to londianensis
and sarissa is bernardi, but this is a smaller species with relatively long mandibles which lacks
intercalary teeth in the apical fork. The other two are separated as follows.
londianensis
HW 0-62-0-70, HL 0-84-0-92.
Some hairs on leading edge of antennal
scape curved basally.
Vertex of head in profile with a
single pair of stout clavate standing
hairs, the cephalic ground-pilosity
short and broadly spatulate.
Pronotal humeri with stout straight
hairs which are clavate apically.
Anterior pronotal margin between the
humeral hairs without other standing
hairs.
Propodeum without triangular teeth.
sarissa
HW 0-50-0-60, HL 0-72-0-82.
All hairs on leading edge of antennal
scape curved apically.
Vertex of head in profile without standing
hairs, the cephalic ground-pilosity
elongate, dense and narrowly spatulate.
Pronotal humeri with elongate fine
flagellate hairs.
Anterior pronotal margin between the
humeral hairs with a pair of stouter
standing hairs; rarely with two pairs.
Propodeum with triangular teeth.
MATERIAL EXAMINED
Kenya: Londiani (5. Patrizi); Nyandura, Njabini (V. Mahnert & J.-L. Ferret); Mt Elgon Nat. Pk.,
Koitoboa Peak (V. Mahnert).
Strumigenys marleyi Arnold
Strumigenys havilandi race marleyi Arnold, 1914: 31, fig. 10. Syntype workers, SOUTH AFRICA: Natal,
Durban, ii.1914, 'in nest of Pheidole punctulata" (F. B. Marley) (SAM) [examined].
Strumigenys marleyi Arnold; Arnold, 1926: 286. [Raised to species.]
Strumigenys marleyi Arnold; Brown, 1954: 24.
WORKER. TL 2-4-2-5. HL 0-62-0-66, HW 0-48-0-50, CI 75-77, ML 0-22-0-24, MI 35-37, SL 0-30-0-32, SI
60-63, PW 0-28-0-31, AL 0-62-0-66 (2 measured).
Mandibles in full-face view broad basally and narrowing to the apex, with an exaggerated basal external
angle and with a basal internal rounded lamina, both of which serve to increase the basal width of the
blades. Apical fork of left mandible without intercalary teeth but the lower spiniform fork tooth with an
adventitious tooth arising from its ventral basal surface which is about half the length of the lower fork
tooth, and with a minute denticle between this adventitious tooth and the lower fork tooth. Apical
armament of right mandible as left but the minute denticle may be absent. Both mandibular blades with 2
preapical teeth set close to the apex, the proximal of these larger than the distal. Upper scrobe margins
shallowly convex and divergent from just behind the frontal lobes. Preocular notch absent. Eyes large,
their maximum diameter distinctly greater than the maximum width of the scape. Ventral surface of head
without a preocular transverse groove or impression. Antennal scapes weakly curved in the basal third,
broadest at about the midlength, the leading edge shallowly convex and with a series of apically curved
THE AFROTROPICAL DACETINE ANTS
379
spoon-shaped hairs. Dorsal surfaces of scapes with numerous short spatulate to narrowly spoon-shaped
hairs present. In full-face view the cephalic dorsum densely clothed with conspicuous short scale-like to
spoon-shaped hairs which are curved anteriorly; those hairs bordering the upper scrobe margins no larger
than those on the dorsum. Cephalic dorsum without simple standing hairs of any description. Head finely
but sharply punctate everywhere. Pronotal humeri without projecting hairs of any description, the
mesonotum without standing hairs. Dorsal alitrunk only with short hairs similar to but sparser than those
on the head. In profile the pronotum and mesonotum forming a single even convexity, the posterior
portion of the mesonotum not suddenly depressed. Metanotal groove present across the dorsum as a very
feebly impressed line, the impression visible in profile but extremely shallow and narrow. Propodeal teeth
broad, laminar and confluent through most of their length with the broad sinuate infradental lamellae, both
the teeth and the laminae appearing reticulate or even spongiform . Entirety of sides and dorsum of alitrunk
densely punctate to reticulate-punctate. Spongiform appendages of pedicel segments strongly developed.
Ventral appendage of petiole in profile deeper than the depth of the peduncle at its midlength and abruptly
truncated posteriorly, the end of the spongiform appendage occurring directly below the highest point of
the node. Lateral spongiform lobe of petiole large. Ventral spongiform lobe of postpetiole in profile large,
its area distinctly greater than the exposed area of the postpetiolar disc. Lateral postpetiolar lobe almost as
large as the ventral. Petiole node punctate dorsally, the postpetiole smooth. Posterior face of petiole node
bordered by a translucent lamella. Postpetiolar disc in dorsal view with projecting spongiform tissue
present all down the sides, posteriorly with a narrow translucent laminar strip. Base of first gastral tergite
with a narrow laminar transverse strip, the basigastral costulae radiating from the lateral portions of this
strip , on each side of a central clear area . Petiole , postpetiole and gaster with numerous stout standing hairs
which are thickened apically. Colour yellowish brown.
Together with pallestes, marleyi forms a close species-pair characterized by their broad-based
mandibles, distinctive apical mandibular armament, complete set of preapical teeth, relatively
large eyes, absence of a preocular notch and completely reticulate-punctate sides to the alitrunk.
The following characters separate the two species.
pallestes
HW 0-38-0-44, HL 0-52-0-58.
Pronotal humeri with a single straight
hair which is clavate apically and
is directed laterally.
Promesonotum at each side bordered
by a longitudinal row of 4-5 short
clavate standing hairs.
Metanotal groove not impressed.
Dorsum of petiole node weakly
transversely striate.
Pronotal dorsum with longitudinal
rugular sculpture.
marleyi
HW 0-48-0-50, HL 0-62-0-66.
Pronotal humeri without projecting hairs
of any description.
Promesonotum not bordered by a row
of standing hairs.
Metanotal groove feebly impressed.
Dorsum of petiole node punctate.
Pronotal dorsum reticulate-punctate,
without rugular sculpture.
Strumigenys mesahyla sp. n.
(Fig. 69)
HOLOTYPE WORKER. TL 2-0, HL 0-57, HW 0-41, CI 75, ML 0-25, MI 43, SL 0-32, SI 75, PW 0-26, AL 0-58.
Apical fork of each mandible with 2 teeth, without intercalary teeth or denticles. Preapical armament of
2 teeth on each blade, the proximal longer than the distal and both teeth situated in the apical quarter to
third of the length of the blade . Upper scrobe margins bordered by a narrow rim or flange which is broadest
behind the frontal lobes and slowly peters out posteriorly; the eyes not visible in full-face view. Eyes very
small, with only 4 ommatidia, the maximum diameter of the eye less than the maximum width of the scape.
Preocular notch absent, the ventral surface of the head lacking a preocular transverse groove or
impression. Scapes relatively slender, only moderately broadened in the median third and evenly shallowly
curved in the basal third. Leading edges of scapes with an apically curved row of shallowly spoon-shaped
hairs. Ground-pilosity of cephalic dorsum everywhere of broad flattened to spoon-shaped hairs which are
curved anteriorly and which appear scale-like in full-face view. These hairs approximately the same size
everywhere on the dorsum, not becoming much smaller on the posterior half. Hairs fringing the upper
scrobe margins the same shape and size as those on the dorsum. Four stout standing hairs which are
thickened apically are present in a transverse row close to the occipital margin; there is no pair of standing
380 BARRY BOLTON
hairs situated anterior to this row. Dorsum of head reticulate-punctate. Pronotal humeri each with a single
long flagellate hair, the mesonotum with a single pair of stout standing hairs. Ground-pilosity of alitrunk of
sparse scale-like hairs which are similar to those on the head but smaller. Metanotal groove represented by
a faint line across the dorsum, the mesonotum sharply depressed behind the level of the pair of hairs.
Propodeal teeth narrowly triangular and acute apically, confluent in their basal halves with the shallowly
convex broad infradental lamellae. Sides of alitrunk mostly smooth, with vestigial traces of rugular
sculpture anteriorly on the pronotum and with scattered peripheral patches of punctures on the pleurae
and propodeum. Pronotal dorsum longitudinally rugulose, the remainder of the dorsal alitrunk punctate.
Petiole node punctate dorsally, the postpetiole smooth and shining. Spongiform appendages of pedicel
segments well developed. In profile the petiole with a broad ventral strip which projects into a lobe below
the spiracle, and with a lateral lobe on the node. Postpetiole with large lateral and ventral spongiform
lobes, the latter much larger than the exposed area of the postpetiolar disc in profile. Basigastral costulae
sparse but quite sharply defined, arising on each side of a broad central clear area. Petiole, postpetiole and
gaster with stout standing hairs which are thickened apically. Colour yellow.
PARATYPE WORKER. TL 2- 1 , HL 0-52, HW 041 , CI 79, ML 0-22, MI 42, SL 0-30, SI 73, PW 0-24, AL 0-54.
As holotype but the infradental lamella of the propodeum not as evenly convex as indicated in Fig. 69
and the ventral spongiform lobe of the petiole more broadly triangular and only narrowly spongiform in
front of the lobe.
Holotype worker, Zimbabwe: Bulawayo, Hillside, 8.ii.l914, in nest of Solenopsis sp. (G. Arnold)
(BMNH).
Paratype. 1 worker, Zimbabwe: Victoria Falls, spray forest, iii.1969 (W. L. Brown) (MCZ).
The closest relative oimesahyla is traegaordhi, known only from South Africa. Details separating
the two are tabulated under the latter name.
Strumigenys murshila sp. n.
(Fig. 77)
HOLOTYPE WORKER. TL 2-3, HL 0-58, HW 0-44, CI 76, ML 0-28, MI 48, SL 0-34, SI 77, PW 0-27, AL 0-58.
Mandibular blades slender and shallowly but distinctly bowed outwards in full-face view. Apical fork of
each mandible with a pair of spiniform teeth, without intercalary teeth or denticles. Two preapical teeth
present on each blade, the proximal longest and the distal tooth on the left mandible slightly smaller than
that on the right. Length of left distal preapical tooth about equal to the distance separating its base from
that of the proximal preapical tooth. Upper scrobe margins narrow anteriorly, the preocular laminae
prominent, strongly divergent and evenly convex posteriorly but without a projecting bordering rim or
flange. Eyes only partially visible in full-face view. Eyes very small, with only 5-6 ommatidia, their
maximum diameter less than the maximum width of the scape. Preocular notch and ventral preocular
transverse impression absent. Antennal scapes slender and more or less cylindrical, very weakly bent near
the base where they are slightly narrowed. Leading edges of scapes equipped with a row of apically curved
narrow spatulate hairs. Ground-pilosity of cephalic dorsum and hairs bordering the upper scrobe margins
the same; hairs approximately the same length and thickness on all parts of the head, curved and narrowly
spatulate, conspicuous. Upper scrobe margins without a row of much broader larger hairs which contrast
with the ground-pilosity. Dorsum of head with 6 standing hairs, arranged in a transverse row of 4 close to
the occipital margin, and a more anteriorly situated pair close to the highest point of the vertex. Dorsum of
head sharply reticulate-punctate. Pronotal humeri each with a long fine flagellate hair. Mesonotum with a
single pair of stout standing hairs. Sparse ground-pilosity of dorsal alitrunk of slender spatulate hairs which
are shorter and narrower than those on the head. Metanotal groove a narrow transverse impression across
the dorsum. In profile the anterior mesonotum slightly raised above the level of the posterior pronotum.
Posterior portion of mesonotum depressed behind the level of the standing hairs, the metanotal groove
impressed and the propodeum raised and convex behind the groove. Propodeal teeth narrowly triangular,
the infradental lamellae narrow, confluent with only the basal third or so of the tooth. Sides of pronotum,
extreme upper portions of the pleurae and propodeum punctate, the sides of the alitrunk otherwise
smooth. Entire dorsal alitrunk reticulate-punctate, the pronotum also with a few weak overlying rugulae
which are irregularly longitudinal. Petiole node punctate dorsally, the postpetiole longitudinally costulate-
rugulose. Spongiform appendages of pedicel segments strongly developed. In profile the petiole with a
curtain-like ventral process, the postpetiole with large lateral and ventral lobes of which the latter is larger
than the exposed area of the postpetiolar disc in profile. Disc of postpetiole surrounded on all sides by
spongiform material in dorsal view, the lateral spongiform lobes strongly prominent at the sides and the
THE AFROTROPICAL DACETINE ANTS 381
posterior transverse strip broad. Basal strip of first gastral tergite lamellate spongiform, the basigastral
costulae arising from it almost parallel and only weakly directed towards the midline, not conspicuously
radiating from the lateral portions of the strip. Petiole, postpetiole and first gastral tergite with standing
hairs which are slightly thickened apically. Colour brownish yellow, the gaster somewhat broader than the
head and alitrunk.
Holotype worker, Rwanda: Rangiro, 10.vii.1973, 1800 m (P. Werner) (MHN).
Known only from the holotype murshila is nonetheless a very distinctive species of the
scotti-complex characterized by its cephalic pilosity, small eyes and lack of a preocular notch,
sculptured postpetiole and slender antennal scapes.
Strumigenysnimbratasp. n.
HOLOTYPE WORKER. TL 1-5, HLO-43,HWO-31, CI72,MLO-18, MI42, SLO-22,SI71,PWO-20, ALO-37.
Outer margins of mandibles shallowly convex in full-face view, the blades narrowing basally and
broadest at about the midlength. Apical fork of each mandible with 2 spiniform teeth, without intercalary
teeth or denticles. Each mandible with 2 preapical teeth, a larger proximal tooth which is situated very
close to the midlength of the blade, and a smaller distal preapical tooth which is close to the apical fork. The
distance separating the bases of these two teeth is distinctly greater than the length of the distal preapical
tooth. Upper scrobe margins with a narrow inconspicuous bordering rim or flange which is distinctly
narrower than the maximum diameter of the eye. Eyes small, with only 4 ommatidia, the maximum
diameter equal to or slightly less than the maximum width of the scape. Preocular notch absent, the ventral
surface of the head without a preocular transverse groove or impression. Antennal scapes slender at the
base and very weakly curved, the medial third slightly expanded and the leading edges with a row of
apically curved narrow spoon-shaped hairs which are smaller than those fringing the upper scrobe margins.
Funicular segments 2 and 3 vestigial and difficult to see, the separation of the two segments almost invisible
and the length of segments 2 and 3 together less than half the length of segment 4 (the penultimate
segment); under low magnification or in poor light the funiculus appears to consist of only 3 segments
rather than the usual 5. Dorsum of head from posterior clypeal margin to about the midlength with
conspicuous narrowly spoon-shaped pilosity which is curved anteriorly, and a double to triple row of these
hairs border the upper scrobe margins. Behind the midlength the hairs are much smaller and sparser,
narrow and inconspicuous; the pilosity of the two areas contrasting strongly. Dorsum of head with 6
standing hairs arranged in a transverse row of 4 close to the occipital margin and a more anteriorly situated
pair. Dorsum of head reticulate-punctate. Pronotal humeri each with a single long fine flagellate hair.
Mesonotum with a single pair of stout standing hairs. Ground-pilosity of dorsal alitrunk of sparse small
hairs which are closely applied to the surface. Metanotal groove a feeble transverse line on the dorsum
which is minutely impressed. Dorsum of mesonotum very shallowly concave in profile behind the level of
the standing hairs, not sharply depressed. Propodeal teeth small and narrowly triangular, the infradental
lamellae very narrow and petering out ventrally, broadest where they join the teeth. Sides of alitrunk
unsculptured. Pronotal dorsum very weakly and irregularly longitudinally rugulose, the remainder of the
dorsal alitrunk and the petiole node reticulate-punctate. Postpetiole smooth in centre of disc but elsewhere
with faint superficial reticulation. Spongiform appendages of pedicel segments much reduced, the
peduncle of the petiole with a narrow ventral strip and the lateral lobe of the node minute. Petiole node
broader than long in dorsal view. Ventral spongiform lobe of postpetiole smaller than the exposed area of
the postpetiolar disc in profile. Basigastral costulae widely spaced and short, but sharply defined. Petiole,
postpetiole and gaster with stout standing hairs which are thickened apically. Colour dull yellow to
brownish yellow.
PARATYPE WORKERS. TL 1-5-1-8, HL 0-42-0-46, HW 0-31-0-35, CI 72-77, ML 0-17-0-20, MI 40-45, SL
0-22-0-24, SI 68-73, PW 0-19-0-23, AL 0-35-0-43 (12 measured).
As holotype, the eyes with 4-6 ommatidia and the sculpture showing some variation in intensity. The
postpetiole may be as described above, or wholly smooth, or even have a few faint longitudinal rugulae
towards the outer edges of the disc.
Holotype worker, Ivory Coast: Tai Forest, 17.x. 1980 (V. Mahnert & J.-L. Ferret) (MHN).
Paratypes. 31 workers and 1 female with same data as holotype (MHN; BMNH; MCZ; ENSA).
Non-paratypic material examined. Ivory Coast: Banco Forest (/. Lobl); Banco Forest (W. L. Brown);
Divo (L. Brader); Monogaga (V. Mahnert & J.-L. Ferret); Tai Forest (V. Mahnert & J.-L. Ferret);
Languededou (V. Mahnert & J.-L. Ferret); Adiopodoume (V. Mahnert & J.-L. Ferret). Ghana: Tafo
(B. Bolton).
382 BARRY BOLTON
The size range of the non-paratypic material is HL 042-048, HW 0-31-0-37, CI 73-77, ML
0-17-0-21, MI 40-44, SL 0-22-0-26, SI 69-71. All this material matches the holotype. 5.
nimbrata is easily diagnosed by its very reduced funicular segments 2 and 3. Other characters
aiding its recognition within the arnoldi-complex include the cephalic pilosity, position of the
proximal preapical teeth and size of the distals, development of the infradental lamellae and
spongiform appendages, and minute size. The only other species sharing the character of very
reduced funicular segments is bitheria, but in this species the flange bordering the upper scrobe
margins is very broad, the distal preapical tooth of the mandible is longer, the pronotal dorsum
has distinct punctate sculpture between the rugulae, the petiole node is as broad as long in dorsal
view and the propodeal teeth are much longer than in nimbrata.
Strumigenysomalyxsp. n.
(Fig. 63)
HOLOTYPE WORKER. TL 2-3, HL 0-57, HW 045, CI 79, ML 0-23, MI 40, SL 0-31, SI 69, PW 0-26, AL 0-58.
Mandibles in full-face view of approximately the same width to the proximal preapical tooth, not evenly
tapering from base to apex. Apical fork of 2 spiniform teeth on each mandible, without intercalary teeth or
denticles. Each mandibular blade with 2 preapical teeth, the proximal much longer than the distal in each
case. Upper scrobe margins with a narrow bordering rim or flange, the eyes not visible in full-face view.
Eyes small, with only 5-6 ommatidia, the maximum diameter of the eye conspicuously much less than the
maximum width of the scape. Preocular notch absent, the ventral surface of the head without a transverse
preocular groove or impression. Antennal scapes shallowly bent at about the basal third, somewhat
dorsoventrally flattened and broadest at about the midlength, their leading edges distinctly convex and
equipped with a row of apically curved large spatulate to spoon-shaped hairs which are as large as or slightly
larger than those fringing the upper scrobe margins. Dorsum of head in full-face view clothed with broad
scale-like to stud-like hairs which do not decrease in size posteriorly on the dorsum. Cephalic dorsum with a
transverse row of 4 stout standing hairs close to the occipital margin, without a more anteriorly situated pair
close to the highest point of the vertex. Head densely and strongly reticulate-punctate everywhere.
Pronotal humeri without flagellate hairs, lacking projecting hairs of any description. Mesonotum with a
single pair of stout standing hairs which are broadly clavate apically. Ground-pilosity of dorsal alitrunk of
sparse scattered scale-like hairs. Metanotal groove feebly marked across the dorsum but not impressed.
Propodeal teeth triangular, broad in profile and confluent for approximately their basal halves with the
broad sinuate infradental lamellae. Sides of pronotum densely reticulate-punctate, the pleurae and sides of
the propodeum smooth except for peripheral puncturation. Dorsal alitrunk densely reticulate-punctate
everywhere, the pronotum not overlaid by longitudinal rugulae. Dorsum of petiole node reticulate-
punctate, the postpetiolar disc longitudinally striolate to punctate-striolate, the sculpture denser towards
the sides of the disc than at the centre. Spongiform appendages of pedicel segments well developed, the
ventral spongiform lobe of the postpetiole larger than the lateral lobe and equal to or slightly larger than
the exposed area of the postpetiolar disc in profile. Basigastral costulae arising on each side of a central
clear area. Petiole, postpetiole and first gastral tergite with standing stout hairs which are clavate apically.
Colour light brown.
PARATYPE WORKERS. TL 2-0-2-4, HL 0-49-0-60, HW 0-41-0-46, CI 75-80, ML 0-19-0-23, MI 35-40, SL
0-26-0-32, SI 65-71, PW 0-24-0-28, AL 0-49-0-60 (15 measured).
As holotype but in some the postpetiolar disc is more strongly sculptured, the central portion
reticulate-punctate and the lateral portions striolate.
Holotype worker, Kenya: TanaR., Sankuri, 160m, 18.X.1977 (V. Mahnert &J.-L. Ferret) (MHN).
Paratypes. 42 workers and 4 females with same data as holotype (MHN; BMNH; MCZ; ENS A).
Non-paratypic material examined. Kenya: Lamu, nr Witu (V. Mahnert & J.-L. Ferret).
Closest related to arnoldi and sharing that species' lack of pronotal flagellate hairs whilst
retaining a complete mandibular dentition of 2 preapical teeth on each blade. 5. omalyx is
separated from arnoldi by the presence in the former of sculptured pronotal sides and
postpetiolar disc, and a lack of longitudinal rugulae on the pronotal dorsum. In arnoldi the
pronotum is smooth laterally, the disc of the postpetiole is smooth and longitudinal rugulae are
present on the pronotal dorsum.
THE AFROTROPICAL DACETINE ANTS 383
Strumigenys pallestes Bolton
(Fig. 59)
Strumigenys pallestes Bolton, 1971: 62, figs 2, 3. Holotype worker, paratype workers and female, GHANA:
Eastern Region, New Tafo, Cocoa Res. Inst. Ghana, mossy rot hole in trunk of cocoa tree, 22.vii.1970
(B. Bolton) (BMNH; MCZ) [examined].
WORKER. TL 2-0-2-2, HL 0-52-0-58, HW 0-38-0-44, CI 70-77, ML 0-18-0-21, MI 32-36, SL 0-24-0-26, SI
57-62, PW 0-24-0-30, AL 0-52-0-60 (20 measured).
Mandibles in full-face view broad basally and tapering towards the apices, the inner margin with a large
basal lamellate lobe whose apex is directed posteriorly and is concealed by the clypeus when the mandibles
are closed. External margins of mandibles with an accentuated basal angle, the blades enclosing a central
vacuity at full closure, the vacuity broadest distally and tapering towards the base. Apical fork of each
mandible consisting of a pair of spiniform teeth, lacking intercalary teeth or denticles. Ventral margin of
lower fork tooth with a smaller adventitious tooth arising near its base, and with a minute denticle present
basally between this adventitious tooth and the lower fork tooth. Each mandibular blade with 2 preapical
teeth, the proximal the longest. Eyes not visible in full-face view, concealed by the projecting upper scrobe
margins. Preocular notch absent, the ventral surface of the head without a preocular groove or impression.
Eyes moderate in size, with 5-6 ommatidia in the greatest diameter, their maximum diameter equal to or
only fractionally less than the maximum width of the scape. Antennal scapes weakly curved basally, slightly
expanded and broadest at about the midlength, their leading edges with a row of curved spoon-shaped
small hairs. Dorsum of head densely clothed with short broad flattened hairs which appear scale-like to
short spatulate in full-face view, the upper scrobe margins densely fringed by hairs similar in shape and size
to those on the leading edges of the scapes. Dorsum of head with a transverse row of 4 short standing hairs
close to the occipital margin. Head reticulate-punctate everywhere. Pronotal humeri each with a laterally
projecting straight clavate hair. Lateral margins of promesonotal dorsum with a row of 4-5 clavate hairs on
each side, the first 1-2 of these curve towards the midline, the remaining 3 are more or less straight.
Ground-pilosity of dorsal alitrunk like that on head but the hairs sparser, frequently somewhat smaller and
slightly more elevated. Metanotal groove absent. Mesonotum not depressed posteriorly, instead the
promesonotum forming a single more or less evenly curved surface in profile. Propodeal teeth subtended
by broad convex infradental lamellae. Sides of alitrunk uniformly reticulate-punctate everywhere.
Pronotal dorsum longitudinally rugulose, usually with punctures between the rugulae. Remainder of
dorsal alitrunk densely reticulate-punctate. Dorsum of petiole node weakly transversely striate, the
postpetiole smooth. Spongiform appendages of pedicel segments large, the petiole ventrally with a
spongiform strip which is as deep as the peduncle. Ventral spongiform lobe of postpetiole distinctly larger
than the exposed area of the postpetiolar disc in profile, and larger than the lateral lobe. In dorsal view the
postpetiole narrow, only slightly broader than the petiole. Basigastral costulae dense, radiating on each
side of a central clear area. Dorsal surfaces of petiole, postpetiole and gaster with numerous short standing
hairs which are clavate apically. Colour dull yellow to brownish yellow.
In the Afrotropical region only pallestes and marleyi share the strange mandibular shape and odd
dentition described above. The characters separating these two species are listed under marleyi.
S. pallestes is one of the very few arboreal species of Strumigenys known from Africa. All the
series listed below were collected from rot holes in tree trunks or branches, or from isolated
workers wandering on the bark of a tree.
MATERIAL EXAMINED
Ghana: Tafo (B. Bolton); Baudua (D. Leston). Nigeria: Gambari (B. Bolton); Gambari (B. Taylor}.
Strumigenys paranax sp . n .
HOLOTYPE WORKER. TL 1-8, HL 0-47, HW 0-34, CI 72, ML 0-19, MI 40, SL 0-25, SI 74, PW 0-23, AL 0-46.
Mandibles slender and shallowly curved along their external borders, tapering gradually from base to
apex. Apical fork of each mandible of 2 teeth, without intercalary teeth or denticles. Left mandibular blade
with only 1 preapical tooth (the proximal), right mandible with 2 preapical teeth present. Upper scrobe
margins gradually divergent, the eyes partially visible in full-face view. Eyes small, with only 4 ommatidia,
their maximum diameter less than the maximum width of the scape. Preocular notch absent, the ventral
surface of the head without a transverse preocular groove or impression . Antennal scapes shallowly curved
in the basal third, only slightly broadened medially; the leading edges equipped with a row of apically
curved spoon-shaped hairs which are slightly smaller than those fringing the upper scrobe margins.
384 BARRY BOLTON
Ground-pilosity of head reduced and sparse, consisting of inconspicuous small flattened hairs. Upper
scrobe margins with a double or triple row of large spoon-shaped hairs which are curved anteriorly and are
much more conspicuous than the ground-pilosity. Dorsum of head with 6 standing hairs arranged in a
transverse row of 4 close to the occipital margin and a more anteriorly situated pair. Cephalic dorsum
reticulate-punctate. Pronotal humeri each with a single straight stout hair which is clavate apically , without
the elongate fine flagellate hair usually seen in this position. Mesonotum with a single pair of short broadly
clavate standing hairs. Ground-pilosity of dorsal alitrunk consisting of small flattened hairs which are
closely applied to the surface, similar to those found on the head. Metanotal groove a transverse line on the
dorsum, weakly impressed in profile. Mesonotum in profile not sharply depressed behind the level of the
pair of hairs, instead the dorsum of the mesonotum forming a more or less even slope to the metanotal
groove. Propodeal teeth small and triangular, subtended by infradental lamellae. Sides of alitrunk smooth
except for peripheral punctures round the pleurae and propodeum. Dorsum of pronotum with widely spaced
longitudinal rugulae, the remainder of the dorsal alitrunk reticulate-punctate. Petiole node punctate,
the postpetiole smooth. Spongiform appendages of pedicel segments small, the petiole with only a very
narrow ventral strip and the ventral spongiform lobe of the postpetiole smaller than the exposed area of the
postpetiolar disc in profile. Basigastral costulae short but sharply defined, arising across the width of the
tergite rather than radiating from each side of a broad central clear area. Petiole, postpetiole and gaster
dorsally with stout clavate standing hairs. Colour brownish yellow.
PARATYPE WORKERS. TL 1-7-1-8, HL 0-45-0-46, HW 0-32-0-34, CI 70-74, ML 0-17-0-19, MI 37-41, SL
0-24-0-25, SI 73-75, PW 0-22-0-24, AL 0-42-0-46 (4 measured).
As holotype but some have the mesonotal dorsum shallowly concave in profile.
Holotype worker, Cameroun: Nkoemvon, 1979 (D. Jackson) (BMNH).
Paratypes. Gabon: 1 worker, He aux Singes, IS 1-4 (/. A. Band) (MCZ). Cameroun: 1 worker, Fo Tabe,
19.U937 (no collector's name) (BMNH).
Non-paratypic material examined. Cameroun: nr Yaounde (G. Terron).
Of the four arnoldi-complex species in which only a single preapical tooth is present on the left
mandible, irrorata is identified by its possession of only a single preapical tooth on the right
mandible also, and katapelta by its possession of intercalary small teeth between the teeth of the
apical mandibular fork. Of the two species remaining, which have 2 preapical teeth on the right
mandible and lack intercalary teeth, dextra is recognized by having a long fine flagellate hair at
each of the pronotal humeri, whilst paranax has a stout straight strongly clavate hair in this
position.
Strumigenys petiolata Bernard sp. rev.
(Figs 58, 71)
Strumigenys petiolata Bernard, 1952: 254, figs 14 H-J. Syntype workers, GUINEA: Mt Nimba, 700 m, in
termitary in forest (Villiers) (not found in MNHN; presumed lost). [Previously synonymized with
rufobrunea by Brown, 1954: 17.]
WORKER. TL 2-0-2-5, HL 0-54-0-68, HW 0-40-0-53, CI 75-83, ML 0-24-0-32, MI 44-50, SL 0-28-0-36, SI
64-72, PW 0-25-0-32, AL 0-50-0-64 (38 measured).
Mandibles in full-face view with the outer margins shallowly convex, the blades feebly bowed outwards.
Apical fork of each mandible consisting of a pair of spiniform teeth, without intercalary teeth or denticles.
Preapical armament of each mandibular blade of 2 teeth, the proximal spiniform and the longest in each
case. Right distal preapical tooth usually larger than the left. Upper scrobe margins shallowly sinuate in
full-face view and bordered by a narrow rim or flange throughout their length. Eyes visible in full-face view,
their maximum diameter equal to or greater than the maximum width of the scape . Preocular notch present
and strongly developed, the anterior portion of the eye detached from the side of the head. Preocular notch
continued onto ventral surface of head as a transverse groove which is narrower than the maximum
diameter of the eye and which usually has quite sharply defined margins. Antennal scapes not or only
extremely shallowly bent basally, broadest at about the midlength and their leading edges equipped with a
row of apically curved narrowly spoon-shaped hairs which are slightly smaller than those on the upper
scrobe margins. Cephalic ground-pilosity of minute inconspicuous spatulate to spoon-shaped hairs which
are closely applied to the surface. Upper scrobe margins with a row of anteriorly curved large spoon-
shaped hairs. Dorsum of head with 6 standing hairs which are arranged in a transverse posterior row of 4
close to the occipital margin and a more anteriorly situated pair. Head finely and usually very sharply
THE AFROTROPICAL DACETINE ANTS 385
reticulate-punctate but in some samples the sculpture is less intensely developed, the punctures not so
sharply incised. Pronotal humeri lacking flagellate or any other kind of outstanding hair. Mesonotum with
a single pair of stout standing hairs. Ground-pilosity of alitrunk of minute hairs similar to those on the
cephalic dorsum. Posterior half of mesonotum sharply depressed behind the level of the standing hairs.
Metanotal groove represented by a line across the dorsum but not or only very feebly impressed. Propodeal
teeth triangular and subtended by an infradental lamella on each side. Sides of alitrunk with the pleurae
smooth except for peripheral punctulae which are best developed dorsolaterally. Sides of propodeum
above and behind the spiracle punctulate. Sides of pronotum varying from smooth to very weakly striolate,
sometimes also with vestigial punctures. Pronotal dorsum usually finely longitudinally striolate or
costulate, often with fine superficial punctures between the longitudinal sculpture. Frequently the costulae
or striae are poorly defined and the punctures more conspicuous, and in some samples dense punctures
constitute the principal component. Dorsal alitrunk behind pronotum densely reticulate-punctate. Petiole
node punctate dorsally, the postpetiole often with some fine longitudinal striolae but these are very
variable in development and frequently are absent. Petiole with a narrow ventral spongiform strip whose
depth is less than half the depth of the peduncle at its midlength. Sides of petiole node with a small
triangular appendage. Ventral spongiform lobe of postpetiole usually marginally larger than the lateral
lobe in profile. Petiole, postpetiole and gaster with standing hairs. Colour often uniform, varying in shade
from yellow to dark brown or even blackish brown; sometimes with the gaster considerably darker in
colour than the head and alitrunk.
One of the most successful and widely distributed Strumigenys of the Afrotropical region,
petiolata nests in rotten wood, under the bark of more recently fallen timber, in log mould, or
sometimes directly into the soil. The workers forage singly in the topsoil, leaf litter or in rot
tunnels in wood.
As I have not been able to find the types of petiolata my interpretation of this name and its
application to this common species must remain somewhat shadowed with doubt. The inter-
pretation is based on Bernard's insufficient original description and figure and supplemented by
the notes in Brown's (1954) revision, in which he treated petiolata as a synonym of rufobrunea.
Since that time considerably more material has been amassed and it was noted that Brown's
rufobrunea consisted of more than one species. In particular a number of West African samples
with pronotal flagellate hairs, narrower heads and overall smaller size, were found to match the
rufobrunea types perfectly, and the South African faurei type (also included as a synonym of
rufobrunea by Brown) has also proved to be a separate species. This left the common species
which lacked pronotal flagellate hairs, and which formed the bulk of Brown's concept of
rufobrunea, with the possible available name of petiolata, now applied here. Admittedly
Bernard's description could apply to any of the names mentioned above but his figure does not
show pronotal flagellate hairs and neither are such hairs mentioned in the description. Because
of this, and because the species is so common in West Africa, I have decided that the name
petiolata is most probably applied to the following material, with the diagnostic characters
described above.
MATERIAL EXAMINED
Ivory Coast: Man (V. Mahnert & J.-L. Ferret); Tai Forest (V. Mahnert & J.-L. Ferret); Issoneu (V.
Mahnert & J.-L. Ferret); Sassandra (/. Lobl); Banco Forest (/. Lobl); Divo (L. Brader); Gagnoa (L.
Brader). Ghana: Enchi (D. Leston); Legon (D. Leston); Tafo (D. Lesion); Tafo (B. Bolton); Tafo (C. A.
Collingwood); Mampong (P. Room); Mt Atewa (B. Bolton). Nigeria: Ibadan (A. Russell-Smith); Gambari
(B. Bolton); Apoje (B. Taylor). Cameroun: Nkoemvon (D. Jackson); nr Yaounde (G. Tenon); Batanga
(G. Schwab). Gabon: Makokou (/. Lieberburg); Makokou (W. H. Gotwald); Plateau d'Ipassa (/. A.
Barra); He aux Singes (/. A. Barra). Central African Republic: Haut Mbomu (N. A. Weber). Angola:
Dundo (L. de Carvalho}; R. Chicapa (L. de Carvalho). Sudan: Khor Aba (N. A. Weber).
Strumigenys pretoriae Arnold
(Fig. 53)
Strumigenys pretoriae Arnold, 1949: 267, fig. 8. Syntype workers, SOUTH AFRICA: Transvaal, Pretoria,
22. i. 1946 (E. K. Hartwig) (SAM) [examined].
Strumigenys pretoriae Arnold; Brown. 1954: 15.
386 BARRY BOLTON
WORKER. TL 2-3-3-0, HL 0-59-0-70, HW 0-43-0-54, CI 71-77, ML 0-26-€-31 , MI 40-44, SL 0-33-0-40, SI
69-77, PW 0-27-0-33, AL 0-58-0-70 (5 measured).
Mandibular blades broad and powerful, the outer margins shallowly convex. Apical fork of each
mandible of 2 teeth, without intercalary teeth or denticles. Each mandibular blade with 2 preapical teeth,
crowded close to the mandibular apex, the proximal teeth larger than the distal. Upper scrobe margins in
full-face view constricted immediately behind the frontal lobes; behind the constriction diverging pos-
teriorly in an almost straight line on each side which passes directly above the inner margin of the eye on
each side so that the latter is clearly visible in full-face view. Eyes very large, larger than in any other
Afrotropical Strumigenys, their maximum diameter 0-23-0-24xHW; in full-face view the maximum eye
diameter more than twice the width of the scape at its broadest. Preocular notch present, the anteriormost
portion of the eye detached from the side of the head. Preocular notch continued onto ventral surface of
head as an extensive impressed area. Antennal scapes quite slender, very feebly bent in the basal third, the
leading edges with a row of apically curved spoon-shaped hairs. Dorsum of head clothed with short broad
spoon-shaped hairs which are curved anteriorly and appear scale-like in full-face view. Hairs bordering the
upper scrobe margins the same as those on the cephalic dorsum but slightly larger. Vertex of head with 4
simple standing hairs arranged in a transverse row close to the occipital margin, without a pair situated
anterior to this row. Dorsum of head densely reticulate-punctate. Pronotal humeri without flagellate or any
other kind of projecting hairs. Mesonotum with a single pair of standing hairs. Ground-pilosity of dorsal
alitrunk of small broadly spoon-shaped to scale-like hairs, like those on the head but not so dense. In profile
the posterior portion of the mesonotum slightly depressed, the metanotal groove minutely impressed.
Propodeal teeth lamellate, subtended by broad infradental lamellae which are confluent with the basal
margins of the teeth for about half of their length. Central areas of pleurae smooth but peripherally with
punctate sculpture. Sides of pronotum with faint striolate markings. Pronotal dorsum finely longitudinally
rugulose, the remainder of the dorsal alitrunk finely reticulate-punctate. Petiole node superficially
reticulate-punctate, the postpetiole smooth or with traces of faint longitudinal costulae or striolae.
Spongiform appendages of pedicel segments strongly developed. In profile the petiole with a large ventral
strip which is almost as deep as the peduncle at its midlength; the lateral lobe extensive. Ventral and lateral
lobes of postpetiole large and spongiform, the former larger than the exposed area of the postpetiolar disc
in profile. Sides of postpetiole surrounded by projecting spongiform material in dorsal view. Base of first
gastral tergite with a lamellate transverse strip from the more lateral portions of which the dense and
sharply defined basigastral costulae arise. Petiole, postpetiole and gaster with standing hairs which are
more or less simple or slightly expanded apically. Colour dull yellow to light yellowish brown.
5. pretoriae is immediately separated from its Afrotropical congeners by its very large eyes; no
other species even approaches the ocular development seen here. Its closest relatives are shaula,
dromoshaula and dyshaula but in all of these the pronotal humeri are equipped with flagellate
hairs and the cephalic dorsum lacks the dense scale-like ground-pilosity characteristic of
pretoriae.
MATERIAL EXAMINED
Botswana: Maxwee (A. Russell-Smith). South Africa: Transvaal, Nelspruit (M. Samways); Pretoria
(£. K. Hartwig).
Strumigenys relahyla sp. n
(Figs 57, 74)
HOLOTYPE WORKER. TL 2-0, HL 0-56, HW 0-41, CI 73, ML 0-24, MI 43, SL 0-28, SI 68, PW 0-26, AL 0-52.
Apical fork of mandibles with 2 spiniform teeth, without intercalary teeth or denticles. Preapical
armament of 2 teeth on each blade, the proximal longest, the distal about as long as, or slightly longer than,
the distance separating the bases of the preapical teeth. Outer margins of mandibles shallowly convex in
full-face view and the occipital margin broadly concave. Upper scrobe margins with a very narrow
bordering rim or flange, the eyes mostly visible in full-face view. Maximum diameter of eye about
0-15xHW, slightly greater than the maximum width of the scape. Preocular notch present, narrow but
distinct, the anterior portion of the eye not detached from the side of the head and the preocular notch not
extended onto the ventral surface of the head as a transverse groove or impression. Antennal scapes feebly
bent basally, broadened in the middle third, the leading edge equipped with a row of narrow spoon-shaped
hairs which are smaller than those fringing the upper scrobe margin. Cephalic ground-pilosity of
inconspicuous small spatulate hairs, the upper scrobe margins with a projecting row of large anteriorly
curved spoon-shaped hairs. Dorsum of head with 6 standing hairs arranged in a transverse row of 4 close to
the occipital margin and a more anteriorly situated pair. Dorsum of head finely reticulate-punctate.
THE AFROTROPICAL DACETINE ANTS 387
Pronotal humeri each with a single long fine flagellate hair. Mesonotum with a single pair of stout standing
hairs. Ground-pilosity of dorsal alitrunk of fine spatulate hairs similar to those on the head. Posterior
portion of mesonotum depressed behind the level of the pair of hairs, with a shallow transverse impression
immediately behind the descending slope. Remainder of mesonotum and dorsum of propodeum convex.
Metanotal groove forming a transverse line on the dorsum. Propodeal teeth lamellate, confluent for
slightly more than half their length with the conspicuous infradental lamellae. Sides of alitrunk unsculp-
tured except for some faint scratch-like costulae on the pronotum and some feeble peripheral punctulation
on the pleurae and propodeum. Pronotal dorsum longitudinally finely costulate-rugulose, the remainder of
the alitrunk punctate. Dorsum of petiole node shallowly punctate, the postpetiole smooth. Spongiform
appendages of pedicel segments moderately developed, the petiole with a thin ventral strip and small
lateral lobe. Ventral spongiform lobe of postpetiole larger than the lateral lobe and larger than the exposed
area of the postpetiolar disc in profile. Basigastral costulae relatively sparse, without secondary costulae
arising between those which have their origins on the basal gastral strip. Petiole, postpetiole and gaster
dorsally with standing hairs. Colour yellowish brown, the gaster darker.
PARATYPE WORKERS. TL 1-9-2-0, HL 0-54-0-56, HW 0-40-0-41, CI 73-76, ML 0-23-0-24, MI 43-44, SL
0-26-0-28, SI 65-68, PW 0-25-0-27, AL 0-51-0-54 (4 measured). As holotype.
Holotype worker, Angola: Duque de Braganca Falls, 12.iii.1972, riverbank (P. Hammond) (BMNH).
Paratypes. 4 workers with same data as holotype (BMNH; MCZ).
Non-paratypic material examined. Cameroun: nr Yaounde (G. Terron). Zaire: Ituri Forest, Beni-Irumu
(A/. A. Weber). Angola: Dundo (L. de Carvalho); R. Camudembele (L. de Carvalho); R. Mussungue (L.
de Carvalho); Dundo (A. Machado).
Size range of non-paratypic material is HL 0-50-0-53, HW 0-36-0-41, CI 72-77, ML 0-22-0-24,
MI 44-46, SL 0-25-0-27, SI 66-69 (10 measured). Resembling the holotype but with variable
colour ranging from uniform pale brown, through medium brown with the gaster darker, to
uniform dark brown. The size of the ventral spongiform lobe of the postpetiole shows some
variation but is always at least as large as the exposed area of the disc in profile. The distal
preapical teeth of the mandibular blades are usually as described above but in a few samples they
are shorter than the distance separating the bases of the two preapical teeth. 5. relahyla belongs
to a small aggregation of species in which the preocular notch is present but small, and is not
extended onto the ventral surface of the head as a groove or impression. Of the species thus
defined relahyla is distinguished by a lack of specialized characters when compared to the others.
In totyla the pronotal humeri lack flagellate hairs; mxenohyla the scape hairs are very large and
spoon-shaped, like those on the upper scrobe margins; in adrasora the spongiform appendages
of the petiole and postpetiole are much reduced; in rukha the spongiform appendages are
strongly developed; and in dyshaula the head is more narrowly and deeply impressed at the
occipital margin.
Strumigenys rogeri Emery
(Figs 51, 72)
Strumigenys rogeri Emery, 1890: 68, pi. 7, fig. 6. Holotype worker, ST THOMAS I. (West Indies) (MCSN)
[examined].
Strumigenys incisa Godfrey, 1907: 102 [attributed to Forel]. Syntype workers, GREAT BRITAIN: Scotland,
Edinburgh, hothouse in Royal Botanic Garden, 10. vi. 1904 (R. Godfrey) (BMNH) [examined].
[Synonymy by Donisthorpe, 1915: 341.]
Strumigenys sulfurea Santschi, 1915: 261. Syntype workers, GABON: Samkita (F. Faure) (NMB) [ex-
amined]. [Synonymy by Brown, 1954: 20.]
Strumigenys rogeri Emery; Brown, 1954: 20.
WORKER. TL 2-3-2-8, HL 0-58-0-74, HW 0-42-0-52, CI 69-75, ML 0-31-0-40, MI 51-58, SL 0-36-0-46, SI
82-89, PW 0-27-0-32, AL 0-58-0-68 (40 measured).
Mandibular blades almost straight and at full closure nearly parallel, not obviously bowed outwards.
Apical fork of each blade with 2 spiniform teeth, without intercalary teeth or denticles. Preapical
armament of 2 teeth on each blade , set in the distal third of the blade's length ; the proximal preapical teeth
larger than the distals. Upper scrobe margins narrowly concave immediately behind the frontal lobes, with
a pinched-in appearance in full-face view. Behind this the upper scrobe margins feebly divergent to the level
of the eye and relatively close together, sometimes even shallowly concave directly above the eye, then
388 BARRY BOLTON
diverging strongly to the scrobal apices. Eyes plainly visible in full-face view, the preocular notch strongly
developed and the anterior portion of the eye detached from the side of the head. Preocular notch
continued onto the ventral surface of the head as a broad impression which runs transversely immediately
in front of the level of the eye, but not reaching the ventral midline. Antennal scapes long and slender,
approximately straight , the leading edges equipped with a row of narrowly spatulate hairs which are angled
towards the apex. Dorsum of head with short narrowly spatulate ground-pilosity which is directed
anteriorly, the upper scrobe margins with a row of larger anteriorly curved spoon-shaped hairs. With the
head in profile the dorsum with 6 standing hairs which are arranged as a row of 4 transversely close to the
occipital margin and a more anteriorly situated pair. Dorsum of head reticulate-punctate. Pronotal humeri
each with a long fine flagellate hair and the mesonotum with a single pair of stout standing hairs. Otherwise
the dorsal alitrunk without standing hairs, the ground-pilosity of sparse narrow hairs which are closely
applied to the surface. With the alitrunk in profile the posterior portion of the mesonotum sharply
depressed, the metanotal groove represented by a transverse line across the dorsum but not or only
minutely impressed. Propodeal teeth triangular and subtended by narrow infradental lamellae. Sides of
alitrunk sometimes completely smooth but usually the propodeum punctulate and the pronotum with faint
traces of striolate or costulate sculpture anteriorly. Pronotal dorsum longitudinally striolate or costulate on
a finely punctate surface, but in some the costulae may be very feeble and indistinct; the median costula is
usually stronger and more sharply defined than any other and in many samples forms a weak median
longitudinal carina at least on the anterior half of the pronotum. Remainder of dorsal alitrunk reticulate-
punctate. Dorsum of petiole node weakly reticulate-punctate, the postpetiole generally smooth but
sometimes with vague sculptural vestiges. Petiole in profile with a spongiform ventral strip and the node
with a transverse collar posteriorly. In profile the postpetiole with large ventral and lateral spongiform
lobes. In dorsal view the postpetiole with a posterior spongiform strip which abuts a similar but narrower
strip on the base of the first tergite. Basigastral costulae sparse but sharply defined. Dorsal surfaces of
petiole, postpetiole and gaster with stout standing hairs which are weakly swollen apically. Colour dull
yellow to light medium brown.
Among the members of the rogeri-complex in which the preocular notch is strongly developed
and extends onto the ventral surface of the head as a transverse groove or impression, rogeri is
characterized by its simple dentition (without intercalary teeth in the apical fork and with a full
complement of preapical teeth), relatively long straight mandibles, long antennal scapes,
presence of pronotal flagellate hairs, and presence of characteristically shaped upper scrobe
margins which lack a projecting laminar rim or flange.
5. rogeri is a well known and very efficient tramp species, probably of West African origin but
very widely distributed in the tropics by human commerce. It has also been recorded from
hothouses and other constantly heated buildings in the temperate zone. Brown (1954) gives
observations on the biology of rogeri which were made by Wilson in Cuba. In West Africa the
species usually nests in rotten wood on the ground or under the bark of larger fallen trunks or
branches, but on occasion it will nest directly in the soil or in wood which has crumbled almost to
powder. The Neotropical distribution of rogeri is summarized by Brown (19626) and Kempf
(1972), and the Pacific distribution by Wilson & Taylor (1967).
MATERIAL EXAMINED
Afrotropical material. Ivory Coast: Tai Forest (V. Mahnert & J.-L. Ferret}; Bingerville (V. Mahnert &
J.-L. Ferret); Languededou (V. Mahnert & J.-L. Ferret); Issoneu (V. Mahnert & J.-L. Ferret); Sassandra
(/. Lobl); Man (/. Lobl); Sangouine (/. Lobl); Divo (L. Brader); Banco Forest (W. L. Brown); Nzi Noua
(W. L. & D. E. Brown). Ghana: Tafo (B. Bolton); Tafo (D. Leston). Nigeria: Gambari (B. Bolton).
Cameroun: Nkoemvon (D. Jackson); nr Yaounde (G. Tenon). Gabon: Samkita (¥. Faure); Plateau
d'Ipassa (J. A. Barrd); Makokou (/. Lieberburg). Angola: Gubela (P. Hammond); R. Chicapa (L. de
Carvalho); Cossa (L. de Carvalho). Burundi: Bujumbura (A Dejean).
Other regions. Solomon Is: Guadalcanal (E. S. Brown). Hawaii (F. X. Williams). New Hebrides: Santo
(L. Weatherill); Port Vila (L. Weatherill). Wallis Is: Uvea (G. Hunt). Fiji Is: Viti Levu (W. L. & D. E.
Brown). Malaysia: Sarawak, Gunong Mulu Nat. Pk. (B. Bolton). Great Britain: Scotland, Edinburgh (R.
Godfrey). West Indies: St Thomas I. ; Montserrat (N. A. Weber); Dominica (N. A. Weber). Cuba: Trinidad
Mts (W. M. Mann); Soledad (W. M. Mann). Puerto Rico: Mayaquez (M. R. Smith); Como Springs (W. M.
Wheeler); El Yunque (E. O. Wilson). Jamaica: St Ann Parish (M. Fiske); James Hill. Trinidad: Tumpuna
Res. (J. Noyes); Pitch Lake (N. A. Weber). Haiti: Moea (Russo); Mts N. of Jacmel (W. M. Mann).
Panama: Barro Colorado I. (A. Newton). Guiana: R. Mazaruni (N. A. Weber). Equador: Pichincha, St
Domingo (5. & J. Peck). Seychelles: Cousin I. (G. M. Bathe).
THE AFROTROPICAL DACETINE ANTS 389
Strumigenys rufobrunea Santschi
Strumigenys rufobrunea Santschi, 19146: 373. Lectotype female (designated by Brown, 1954: 17), and
paralectotype worker, GUINEA: Conakry (F. Silvestri) (NMB) [examined].
Strumigenys rufobrunea Santschi; Brown, 1954: 17.
WORKER. TL 1-8-2-0, HL 0-48-0-53, HW 0-36-0-40, CI 73-76, ML 0-22-0-25, MI 45-48, SL 0-25-0-29, SI
69-74, PW 0-23-0-25, AL 0-42-0-49 (12 measured).
Mandibles in full-face view with the outer margins shallowly convex, the blades slightly bowed outwards.
Apex of each mandible with 2 spiniform fork teeth, without intercalary teeth or denticles. Preapical
armament of each mandible of 2 teeth, the proximal spiniform and longer than the distal; in general the
distal preapical tooth of the left mandible slightly shorter than that of the right. Distance separating the
bases of the preapical teeth on the left mandible at least as great as the length of the distal preapical tooth
and usually greater. Upper scrobe margins shallowly sinuate rather than straight, bordered by a narrow rim
or flange throughout their length. Eyes of moderate size, not concealed by the upper scrobe margins and
visible in full-face view, their maximum diameter greater than the maximum width of the scape. Preocular
notch present and strongly developed, the anterior portion of the eye detached from the side of the head.
Preocular notch extended onto the ventral surface of the head as a transverse impression in front of the eye.
Antennal scapes very shallowly bent in the basal third, slightly expanded in the median third and broadest
at about the midlength. Leading edges of scapes with a row of apically curved hairs which are spatulate to
narrowly spoon-shaped and conspicuously smaller than the hairs bordering the upper scrobe margins.
Ground-pilosity of cephalic dorsum of inconspicuous small spatulate hairs which are closely applied to the
surface, the upper scrobe margins with a very obvious row of larger broadly spoon-shaped hairs which are
curved anteriorly. Dorsum of head with 6 standing hairs arranged in a transverse row of 4 close to the
occipital margin and a more anteriorly stituated pair. Dorsum of head reticulate-punctate. Pronotal
humeri each with a single fine flagellate hair. Mesonotum with a single pair of standing hairs, the dorsal
alitrunk otherwise with only sparse appressed minute pilosity similar to that which forms the cephalic
ground-pilosity. Posterior portion of mesonotum depressed behind the level of the standing hairs.
Metanotal groove present as a line across the dorsum which is not or only very feebly impressed. Propodeal
teeth triangular and subtended by a moderately developed infradental lamella which is confluent with the
tooth for about half its length . Sides of pronotum showing vestigial costulate or striolate sculpture , or traces
of punctures which are almost effaced. Pleurae smooth except for some peripheral punctulation. Pronotal
dorsum finely longitudinally costulate or rugulose, sometimes with feeble punctures between the costulae.
Remainder of dorsal alitrunk densely reticulate-punctate. Dorsum of petiole node densely and quite
strongly reticulate-punctate, the node itself slightly broader than long but not a narrow transverse
rectangle. Postpetiole smooth or rarely with vestiges of longitudinal striolate sculpture laterally. Spongi-
form appendages of pedicel segments moderately developed, the ventral petiolar strip spongiform but
usually confined to the posterior two-thirds of the length. Ventral spongiform lobe of postpetiole
fractionally larger than the lateral lobe. Basigastral costulae sharply defined. Petiole, postpetiole and
gaster dorsally with stout standing hairs which are swollen to feebly clavate apically. Colour usually with
head and alitrunk medium brown, the gaster darker brown, but uniformly dark individuals also occur.
In Brown's (1954) study of the African Strumigenys he synonymized two names, petiolata and
faurei, under rufobrunea. Since then a considerable amount of material has been amassed and it
now appears that each of these names represents a separate species, not for the reasons put
forward by their original authors but based upon characters which have only become apparent as
the number of samples available for study has increased. The bulk of the material referred by
Brown to rufobrunea belongs in fact to petiolata, quickly separable as it lacks flagellate hairs at
the pronotal humeri. The remainder is split between the genuine West African rufobrunea and
the South African faurei, known at present only from Natal, both of which possess humeral
flagellate hairs. Characters separating these two are given under faurei.
MATERIAL EXAMINED
Guinea: Conakry (F. Silvestri). Ivory Coast: Lamto (W. H. Gotwald); Man (V. Mahnert &J.-L. Ferret);
Adiopodoume (V. Mahnert & J.-L. Ferret). Ghana: Legon (D. Lesion); Mampong (P. Room). Togo:
Palime, Kpime Forest (Vit). Nigeria: Gambari (B. Taylor).
Strumigenys rukha sp. n.
HOLOTYPE WORKER. TL 2-4, HL 0-60, HW 0-45, CI 75, ML 0-28, MI 47, SL 0-33, SI 73, PW 0-29, AL 0-58.
390
BARRY BOLTON
Mandibular blades with their outer margins shallowly convex, feebly bowed outwards in full-face view.
Mandibular apices each with a fork of 2 teeth, without intercalary teeth or denticles. Each blade with
preapical armament of 2 teeth, the proximal the longest. Distal preapical tooth of left mandible longer than
the distance separating its base from that of the proximal preapical tooth. Upper scrobe margins evenly
divergent posteriorly, the eyes visible in full-face view. Maximum diameter of eye about equal to or very
slightly greater than the maximum width of the scape, the eye with 14-15 ommatidia. Preocular notch
present but vestigial, represented only by an inconspicuous shallow indentation of the ventrolateral margin
immediately in front of the eye ; the preocular notch not continued onto the ventral surface of the head as a
transverse groove or impression. Antennal scapes very shallowly curved in the basal third, the median third
expanded to about twice the basal width. Leading edges of scapes equipped with an apically curved row of
slender spatulate hairs which are smaller than the projecting hairs fringing the upper scrobe margins.
Ground-pilosity of cephalic dorsum of inconspicuous short flattened to spatulate curved hairs, the upper
scrobe margins fringed by an anteriorly curved row of much larger hairs which are spatulate to narrowly
spoon-shaped. Dorsum of head with 6 standing hairs arranged in a transverse row of 4 close to the occipital
margin and a more anteriorly situated pair. Cephalic dorsum reticulate-punctate. Pronotal humeri each
with a fine flagellate hair. Mesonotum with a single pair of stout standing hairs. Ground-pilosity of dorsal
alitrunk of sparse inconspicuous hairs similar to those on the head. Posterior portion of mesonotum
depressed behind the level of the pair of hairs. Metanotal groove represented by a feebly marked line
across the dorsum, not impressed. Propodeal teeth short and broadly triangular, subtended by a broad
infradental lamella on each side which is confluent with the tooth for half or more of its length. Sides of
alitrunk mostly smooth, the pronotal sides with vestigial superficial reticulate markings and the pleurae
with faint peripheral punctures. Pronotal dorsum longitudinally finely costulate-rugulose, the remainder of
the dorsal alitrunk punctate. Petiole node punctate dorsally, the postpetiole smooth and shining.
Spongiform appendages of pedicel segments well developed, the petiole ventrally with a broad spongiform
strip which follows a basal low broad translucent triangular lobe. Ventral spongiform lobe of postpetiole
larger than the lateral lobe and larger than the exposed area of the postpetiolar disc in profile. Petiole node
in dorsal view transversely rectangular, with a posterior transverse lamella. Postpetiole surrounded on all
sides with spongiform material, the lateral and ventral lobes projecting beyond the outline of the disc and
visible in dorsal view. First gastral tergite with a narrow basal strip from which the sharply defined
basigastral costulae arise. Petiole, postpetiole and gaster dorsally with standing stout hairs. Colour yellow.
PARATYPE WORKERS. TL 2-3-2-4, HL 0-57-0-60, HW 0-44-0-46, CI 75-79, ML 0-27-0-28, MI 47-49, SL
0-30-0-33, SI 67-73, PW 0-28-0-30, AL 0-56-0-60 (10 measured).
As holotype but in some the mesonotum with a shallow transverse impression at the base of the
descending portion of the sclerite. The translucent lobe at the base of the petiole ventrally is very variable
in shape and size. In most workers it is a triangular to rounded lobe but in some is much reduced and rarely
it may be absent. The preocular notch, weakly developed at best, may be undetectable.
Holotype worker, Kenya: Embu, Kirimiri Forest, W. of Runyenje, 1550 m, 3.x. 1977 (V. Mahnert &
J.-L. Perret) (MHN).
Paratypes. 42 workers and 2 females with same data as holotype (MHN; BMNH; MCZ; ENSA).
Non-paratypic material examined. Kenya: Embu, Kirimiri Forest (V. Mahnert & J. -L. Perret). Uganda:
Ft. Portal (N. A. Weber).
S. rukha is most closely related to adrasora, relahyla, and dyshaula. It is easily separated from
the first of these as in adrasora the spongiform appendages of the pedicel segments are small, the
ventral postpetiolar lobe being smaller than the exposed portion of the postpetiolar disc in
profile. In rukha the spongiform appendages are better developed than in dyshaula and relahyla,
but in dyshaula the distal preapical tooth of the left mandible is much more slender than the
proximal and only about half of its length, whereas in rukha the distal preapical tooth of the left
mandible is only marginally narrower than the proximal and is three-quarters or more of its
length. In relahyla the mandibles are slightly shorter (MI 43-46) and stouter than in rukha (MI
47-49) and the preocular notch is much more strongly impressed.
Strumigenys sarissa sp. n.
(Fig. 50)
HOLOTYPE WORKER. TL 2-9, HL 0-72, HW 0-52, CI 72, ML 0-38, MI 53, SL 0-46, SI 88, PW 0-33, AL 0-74.
Apical fork of each mandible of 2 spiniform teeth, the fork of the left mandible with an intercalary small
THE AFROTROPICAL DACETINE ANTS 391
tooth between the upper and lower spiniform teeth; right apical fork without an intercalary tooth. Blade of
left mandible with a single preapical tooth, the proximal; right mandibular blade with 2 preapical teeth, a
spiniform proximal (which is equal in size to that on the left blade) and a much smaller distal preapical tooth
which is situated very close to the apical fork and is hidden from view by the right dorsal fork tooth when the
mandibles are closed. Upper scrobe margins irregular in full-face view, not fringed by a lamina throughout
their length. Behind the convex frontal lobes the upper scrobe margins are sharply concave and have a
pinched-in appearance. Posterior to this the upper scrobe margins expand and diverge, are shallowly
concave above the eyes so that the latter are clearly visible, and diverge more strongly behind this.
Preocular notch deep and strongly developed, the anterior portion of the eye detached from the side of the
head. Preocular notch continued onto ventral surface of head as a shallow but broad impression. Antennal
scapes elongate and narrow, subcylindrical and with all the hairs on the leading edges directed apically.
Ground-pilosity of head everywhere of fairly dense narrowly spatulate hairs which are curved anteriorly,
the hairs fringing the upper scrobe margins not noticeably larger than those elsewhere on the vertex. In
profile the vertex lacking larger prominent hairs which project above the ground-pilosity. Entire cephalic
dorsum finely punctate. Pronotal humeri each with a long fine flagellate hair and the curved anterior
margin of the pronotum between the flagellate hairs with a pair of shorter but stouter erect simple curved
hairs. Mesonotum with a pair of strong standing hairs, the dorsal alitrunk otherwise without standing
pilosity except that in a few paratypes a second pair of simple erect hairs may occur on the pronotum.
Ground-pilosity of dorsal alitrunk of sparse curved narrow hairs which are closely applied to the surface. In
profile the pronotum and anterior mesonotum high and convex, the posterior mesonotum and propodeum
depressed. Metanotal groove not impressed. Propodeum with a pair of triangular teeth which are
subtended by narrow infradental lamellae. Sides of alitrunk feebly punctate peripherally, the main area of
the pleurae smooth. Dorsal alitrunk punctate everywhere. Dorsum of petiole punctate, the postpetiole
showing vestigial punctate sculpture. Petiole in profile without a ventral spongiform appendage, with a
narrow posterior collar on the node. Postpetiole with a moderate ventral spongiform lobe and a smaller
lateral lobe. In dorsal view the postpetiole with a narrow posterior spongiform strip. Base of first gastral
tergite with a narrow transverse strip, with numerous fine basal costulae. Petiole, postpetiole and gaster
dorsally with strong standing pilosity which is clavate apically. Colour light brown.
PARATYPE WORKERS. TL 2-9-3-2, HL 0-72-0-82, HW 0-50-0-60, CI 68-75, ML 0-38-0-44, MI 53-55, SL
0-46-0-52, SI 83-92, PW 0-30-0-38, AL 0-74-0-80 (15 measured).
As holotype but some with a second pair of simple hairs on the pronotum which are sited beside the
flagellate hairs. Postpetiolar sculpture may be intense so that the disc is as strongly punctate as the petiole,
and the basigastral costulae may be more strongly defined. The petiole ventrally usually lacks a spongiform
appendage but in some a very narrow strip may be present.
Holotype worker, Burundi: Bujumbura, no. 82, 1977 (A. Dejean) (BMNH).
Paratypes. Burundi: 1 worker with same data as holotype; 2 workers with same data but no. 86. Rwanda:
17 workers and 1 female, Kayove, 2100 m, 15. v. 1973 (P. Werner);^ workers with same data but 25. v. 1973;
2 workers with same data but 23. iv. 1973; 2 workers, Kamiranzovu, 1900 m, i.1976 (P. Werner) (BMNH;
MHN;MCZ;ENSA).
Non-paratypic material examined. Zaire: Lwiro (P. J. Curtis).
In the Afrotropical region the characteristic apical and preapical dentition where an intercalary
tooth is present in the left apical fork but not in the right, and the left blade has one preapical
tooth but the right blade has two, is restricted to the two species sarissa and londianesis . Details
for separating them are tabulated under the latter name.
Strumigenys scotti Forel
(Fig. 75).
Strumigenys scotti Forel, 1912: 159. Syntype workers, SEYCHELLE Is: Silhouette, Mare auxCochons, 1000ft
(305 m), 1905 (H. Scott) (BMNH; MHN) [examined].
Strumigenys scotti Forel; Brown, 1954: 23.
WORKER. TL 2-4-2-6, HL 0-62-0-70, HW 0-42-0-46, CI 64-70, ML 0-30-0-33, MI 46-50, SL 0-39-0-42, SI
88-95, PW 0.26-0-29, AL 0-60-0-66 (8 measured).
Mandibles in full-face view with the outer margins shallowly and evenly convex, the width of the blade
approximately constant from the level of the proximal preapical tooth to near the base where the mandibles
are somewhat narrowed. Apical fork of each mandible of 2 stout teeth, without intercalary teeth or
392
BARRY BOLTON
denticles. Each mandible with 2 stout preapical teeth which are situated in the apical third of the length of
the blade. The proximal preapical teeth slightly longer than the distal, the distals longer than the distance
separating the bases of the preapical teeth. Upper scrobe margins evenly and shallowly convex, rounding
cleanly into the sides of the occipital lobes without trace of an angle, the two together forming a single
evenly curved surface in full-face view. Upper scrobe margins not bounded by a rim or flange, the eyes
clearly visible in full-face view. Eyes large, with about 20 ommatidia, the maximum diameter of the eye
distinctly greater than the maximum width of the scape. Preocular notch absent, the ventral surface of the
head without a transverse preocular groove or impression. Antennal scapes long slender and subcylindri-
cal, only very feebly curved near the base and with their leading edges equipped with a row of slender small
hairs which curve towards the apex and which are slightly flattened or spoon-shaped apically. Cephalic
dorsum densely clothed with curved narrow spatulate to spoon-shaped ground-pilosity, the upper scrobe
margins fringed with a dense row of hairs which are the same shape and size as those on the dorsum.
Cephalic dorsum with 6 simple standing hairs arranged in a transverse row of 4 close to the occipital margin
and a more anteriorly situated pair. Pronotal humeri each with a single long fine flagellate hair. Mesonotum
with a single pair of standing hairs. Ground-pilosity on alitrunk as on head but the hairs smaller and
sparser. Posterior portion of mesonotum shallowly depressed behind the level of the standing hairs.
Propodeal teeth triangular and subtended by narrow infradental lamellae. Sides of alitrunk superficially
punctulate peripherally, the pleurae mostly smooth. Pronotal dorsum longitudinally feebly rugulose and
punctate. Remainder of dorsal alitrunk reticulate-punctate. Petiole node in dorsal view reticulate-punctate
and at least as long as broad, often longer than broad. Postpetiole smooth and shining. Spongiform
appendages of pedicel segments well developed, the petiole with a ventral strip and conspicuous lateral
lobe on the node. Postpetiole with large lateral and ventral spongiform lobes of which the ventral is the
larger, about as large as the exposed area of the postpetiolar disc in profile. In dorsal view the spongiform
material not or only very slightly projecting beyond the lateral outline of the disc. Basigastral costulae
arising on each side of a central clear area. Dorsal surfaces of petiole, postpetiole and gaster with standing
hairs which are simple or very slightly thickened apically. Colour yellowish brown to medium brown.
The affinities and differentiation of scotti are discussed under hasty la, a closely related species.
S. scotti is still only known from a couple of collections, one made in the Seychelles and the
other on Sao Tome island. This implies that scotti is most probably an Afrotropical species of
limited tramping ability, but to date no samples have been found on the continental mainland.
MATERIAL EXAMINED
Seychelles: Silhouette I. (H. Scott). Sao Tome & Principe: Sao Tome I., Mkambrera (B. Malkiri).
Strumigenysshaulasp. n.
HOLOTYPE WORKER. TL 2-2, HL 0-57, HW 0-44, CI 77, ML 0-25, MI 44, SL 0-29, SI 66, PW 0-28, AL 0-56.
Mandibles in full-face view weakly bowed outwards. Apical fork of each mandible with 2 teeth, without
intercalary teeth or denticles. Preapical armament of 2 teeth on each mandibular blade, both situated close
to the apex, the proximal longer than the distal in each case. Space separating the proximal and distal
preapical teeth distinctly shorter than the length of the distal tooth. Upper scrobe margins bordered by a
narrow rim or flange throughout their length, evenly divergent posteriorly and approximately straight
rather than sinuate. Eyes relatively large, plainly visible in full-face view, the maximum diameter of the eye
0-18xHW, and in full-face view the length of the eye distinctly much greater than the maximum width of
the scape. Preocular notch present and conspicuous, the anteriormost portion of the eye detached from the
side of the head. Preocular notch continued onto ventral surface of head as a deep transverse groove which
is narrower than the maximum diameter of the eye and has approximately parallel quite sharply defined
margins. Antennal scapes feebly bent in the basal third and slightly thickened medially, the leading edges
with a row of apically curved spoon-shaped hairs. Ground-pilosity of cephalic dorsum of inconspicuous
small curved hairs, the upper scrobe margins fringed by a row of much larger spoon-shaped hairs which are
curved anteriorly. Dorsum of head with 6 standing hairs arranged in a transverse row of 4 close to the
occipital margin and a more anteriorly situated pair. Cephalic dorsum shallowly reticulate-punctate.
Pronotal humeri each with a single fine flagellate hair, the mesonotum with a single pair of standing hairs.
Ground-pilosity of dorsal alitrunk of minute hairs which are closely applied to the surface. With the
alitrunk in profile the posterior portion of the mesonotum only very feebly depressed behind the level of
the hairs, the metanotal groove weakly impressed. Propodeal teeth subtended by infradental lamellae
which are about half as wide as the length of the tooth. Sides of propodeum superficially punctate, the
pleurae mostly smooth except for some peripheral fine punctures which are best developed laterodorsally.
Sides of pronotum with traces of punctate sculpture anteriorly and dorsally. Pronotal dorsum finely
THE AFROTROPICAL DACETINE ANTS 393
longitudinally rugulose, the spaces between the rugulae inconspicously punctulate. Remainder of dorsal
alitrunk reticulate-punctate. Dorsum of petiole node narrow from front to back and very broad, finely
punctate; the postpetiole smooth. Spongiform appendages of pedicel segments moderately developed, the
subpetiolar process narrower than the depth of the peduncle at its midlength. Ventral and lateral
spongiform lobes of postpetiole well developed, the former only marginally larger than the latter and about
equal in size to the exposed area of the postpetiolar disc in profile. In dorsal view the postpetiole with a
narrow laminar posterior transverse strip; on the sides projecting spongiform material restricted to the
posterior halves. Base of first gastral tergite with a laminar transverse strip from which the sharply defined
basigastral costulae arise. Petiole, postpetiole and gaster dorsally with stout standing hairs which are
thickened to feebly clavate apically. Colour yellowish brown to light brown.
PARATYPE WORKERS. TL 2-1-2-3, HL 0-54-0-58, HW 0-42-0-47, CI 75-81, ML 0-25-0-27 MI 44-47 SL
0-27-0-31, SI 64-69, PW 0-27-0-31, AL 0-52-0-58 (3 measured). As holotype.
Holotype worker, Zimbabwe: Gwebi, 1971, ace. 14746, pitfall trap (K. J. Wilson) (BMNH).
Paratypes. 3 workers with same data as holotype (BMNH; MCZ).
The closest relative of shaula appears to bepretoriae, but in that species the eyes are very large,
the head has uniform scale-like pilosity and the pronotal humeri lack flagellate hairs. In the
closely related dromoshaula from Burundi the extension of the preocular notch onto the ventral
surface of the head forms a broad shallow dish-like impression with feebly denned rounded
margins, rather than the narrow groove with sharp edges seen in shaula. In dyshaula the
preocular notch is reduced, small and shallow in full-face view and not extended onto the ventral
surface of the head. 5. shaula also shows some relationship with the West African rufobrunea
but the latter species is smaller, has longer scapes, smaller eyes, more sinuate upper scrobe
margins and a petiole node which in dorsal view is only marginally broader than long, as well as
the dental character given in the key
Strumigenys spathoda sp. n.
(Fig. 62)
HOLOTYPE WORKER. TL 2- 1 , HL 0-55, HW 0-44, CI 80, ML 0- 16, MI 29, SL 0-27, SI 61 , PW 0-27, AL 0-56.
Mandibles very short, stout and powerfully constructed, their outer margins convex. Apical fork of each
mandible with 2 spiniform teeth, the upper of which is very long, its length distinctly greater than 0-5 x ML;
apical forks without intercalary teeth or denticles. Each mandible with 2 preapical teeth, the proximal by
far the longest (just less than 0-5 x ML) and situated at or just distal of the midlength of the blade. Distal
preapical tooth less than half the length of the proximal . Upper scrobe margins bordered by a narrow rim or
flange whose free margins are irregular, the eyes not visible in full-face view. Eyes very small, their
maximum diameter conspicuously very much less than the maximum width of the scape. Preocular notch
absent, the ventral surface of the head without a preocular transverse groove or impression on each side.
Antennal scapes flattened and expanded, the leading edge broadly convex and prominent, equipped with a
row of large spoon-shaped hairs which are about equal in size to those bordering the upper scrobe margins.
Dorsum of head from the posterior clypeal margin to about the midlength densely clothed with broad
anteriorly curved spoon-shaped hairs which appear scale-like in full-face view. Hairs of the same shape and
size fringe the upper scrobe margins. Behind the midlength of the head the hairs are distinctly much smaller
and narrow, and contrast strongly with those on the anterior half. Dorsum of head with a transverse row of
4 short stout standing hairs close to the occipital margin, without a more anteriorly situated pair. Cephalic
dorsum reticulate-punctate to granular. Pronotal humeri each with a fine flagellate hair (apparently easily
lost by abrasion in this species). Mesonotum with a single pair of standing hairs. Ground-pilosity of dorsal
alitrunk consisting of sparse small hairs similar to those on the posterior half of the head. Metanotal groove
represented by a short line on the dorsum, very feebly impressed in profile. Mesonotum not sharply
depressed behind the level of the pair of hairs, instead its surface forming a fairly even slope. Propodeal
teeth triangular and subtended by broad infradental lamellae. Sides of pronotum superficially sculptured,
the pleurae and sides of propodeum mostly smooth, with some faint peripheral punctation. Pronotal
dorsum sparsely longitudinally rugulose , the remainder of the dorsal alitrunk reticulate-punctate . Dorsum
of petiole node punctate, the postpetiole smooth (when clean, in the holotype the surface is obscured by a
thin layer of wax or dirt). Spongiform appendages of pedicel segments well developed. Petiole with a broad
ventral strip which at its broadest is equal to the depth of the peduncle. Ventral and lateral spongiform
lobes of postpetiole subequal, the former marginally larger and about the same size as the exposed area of
394 BARRY BOLTON
the postpetiolar disc in profile. Basigastral costulae short but quite sharply defined. Dorsal surfaces of
petiole, postpetiole and gaster with stout standing hairs which are thickened to clavate apically. Colour
medium brown.
PARATYPE WORKERS. TL 2-0-2-1, HL 0-53-0-55, HW 0-41-0-44, CI 77-81, ML 0-14-0-16, MI 26-30, SL
0-24-0-26, SI 55-61, PW 0-25-0-28, AL 0-48-0-56 (5 measured).
As holotype. All members of the type-series are covered to some extent by a thin layer of dirt or a waxy
deposit which obscures some features. In particular the sculpture of the dorsal body is difficult to discern
and the layer tends to obscure the pilosity.
Holotype worker, Togo: Palime, Klouto Forest, 20-25. iv. 1974 (Vit) (MHN).
Paratypes. 5 workers with same data as holotype (MHN; BMNH; MCZ).
Non-paratypic material examined. Ivory Coast: Man (V. Mahnert & J.-L. Ferret). Cameroun: nr
Yaounde (G. Tenon).
This distinctive species has the shortest mandibles yet recorded for a member of Strumigenys in
the Afrotropical region. It is related to tetraphanes but does not possess the massively lobate
expansions of the anterior scape margins seen in that species and has pronotal flagellate hairs
present.
Strumigenys stygia Santschi
Strumigenys stygia Santschi, 1913a: 257 (diagnosis in key). Syntype workers, KENYA: Cave A at Shimoni,
st. no. 9, xi.1911 (Alluaud & Jeannel) (NMB) [examined].
Strumigenys stygia Santschi; Santschi, 1914a: 113, fig. 20 (description).
Strumigenys stygia Santschi; Brown, 1954: 29.
WORKER. TL 1-9-2-1, HL 0-50-0-53, HW 0-40-0-43, CI 80-84, ML 0-18-0-21, MI 36-40, SL 0-26-0-28, SI
63-68, PW 0-24-0-25, AL 0-48-0-52 (7 measured).
Apical fork of each mandible with 2 teeth, without intercalary teeth or denticles. Each mandibular blade
with 2 preapical teeth, the proximal longer than the distal in each case. Upper scrobe margins convex, the
eyes not visible in full-face view. Eyes small, the maximum diameter distinctly less than the maximum
width of the scape. Preocular notch absent, ventral surface of head without a transverse preocular groove
or impression. Antennal scapes curved in the basal third, the median third expanded and somewhat
flattened, the convex leading edges of the scapes with a row of apically curved short broadly spoon-shaped
hairs. Ground-pilosity of head relatively broad spoon-shaped hairs which are curved anteriorly and appear
stud-like or scale-like in full-face view, the hairs on the dorsum anterior to the highest point of the vertex
somewhat larger and more conspicuous than those posterior to this point. Hairs fringing the upper scrobe
margins the same shape as those on the dorsum. Cephalic dorsum reticulate-punctate. Pronotal humeri
each with a single fine flagellate hair. Mesonotum with a single pair of clavate standing hairs. Ground-
pilosity of dorsal alitrunk of sparse spoon-shaped to scale-like small hairs. Mesonotum depressed behind
the level of the pair of hairs. Metanotal groove represented on the dorsum by a faint line. Sides of
pronotum superficially reticulate or granular, pleurae and propodeum laterally smooth except for some
weak patches of punctate sculpture peripherally. Pronotal dorsum densely punctate, usually with overlying
weak longitudinal rugulae. Remainder of dorsal alitrunk reticulate-punctate. Dorsum of petiole node
reticulate-punctate, postpetiole superficially granular to reticulate-punctate. Ventral spongiform strip of
petiole narrow and inconspicuous. Ventral spongiform lobe of postpetiole equal to or slightly less than the
exposed area of the postpetiole. Basigastral costulae conspicuous. Dorsal surfaces of petiole, postpetiole
and gaster with stout standing hairs which are thickened or clavate apically. Colour light brown to medium
brown.
S. stygia belongs to the core-species of the arnoldi-complex, which also includes arnoldi,
traegaordhi, mesahyla and nimbrata. These four are separated from stygia as arnoldi lacks
pronotal flagellate hairs, nimbrata has funicular segments 2 and 3 vestigial, and both mesahyla
and traegaordhi do not have the postpetiole sculptured.
MATERIAL EXAMINED
Kenya: Shimoni (Alluaud & Jeannel). Zimbabwe: Umtali, Melsetter (R. Mussard). Cameroon: nr
Yaounde (G. Tenon). Angola: Bruco (P. Hammond).
THE AFROTROPICAL DACETINE ANTS 395
Strumigenys tetraphanes Brown
(Fig. 60)
Strumigenys tetraphanes Brown, 1954: 30. Holotype worker, UGANDA: 5 miles (8 km) N. of Kampala,
KawandaExp. St., 15. ii. 1949, soil sample under elephant grass (G. Salt) (MCZ) [examined].
WORKER. TL 2-0-2-2, HL 0-54-0-60, HW 0-51-0-57, CI 91-97, ML 0-19-0-22, MI 34-37, SL 0-28-0-30, SI
52-55, PW 0-30-0-32, AL 0-50-0-58 (4 measured).
Mandibular apices each with a fork of 2 spiniform teeth, without intercalary teeth or denticles, the upper
tooth of the apical fork very long, more than 0-5 x ML. Preapical armament of each blade of 2 teeth, the
proximal long and strongly spiniform, 2-3 times longer than the small distal preapical tooth. Upper scrobe
margins sharply divergent behind, the head broad behind the midlength and almost as broad as long. Eyes
not visible in full-face view, small, conspicuously much smaller than the maximum width of the scape.
Preocular notch absent, the ventral surface of the head without a transverse preocular groove or
impression. Antennal scapes flattened and enormously expanded anteriorly into a large lobe which about
equals the clypeus in area. Leading edges of scapes with a row of broad shallowly spoon-shaped hairs.
Dorsum of head from posterior clypeal margin to about the midlength densely clothed with very broad
shallowly spoon-shaped hairs which appear scale-like to suborbicular in full-face view; such hairs also
fringe the upper scrobe margins. Behind the midlength the cephalic ground-pilosity is much smaller, about
the same as on the clypeus; the difference in size between these hairs and the broad scale-like hairs is
striking. Dorsum of head with a transverse row of 4 longer narrowly clavate hairs close to the occipital
margin, without a pair situated close to the highest point of the vertex. Head densely reticulate-punctate.
Pronotal humeri lacking flagellate or any other kind of projecting hair. Mesonotum with a single pair of
strongly clavate hairs. Ground-pilosity of dorsal alitrunk of small flattened hairs which are almost
appressed. With the alitrunk in profile the sides of the pronotum thickly and bluntly marginate. Anterior
portion of mesonotum shallowly convex, the posterior portion depressed behind the level of the clavate
hairs and shallowly transversely impressed. Propodeal teeth subtended by broad infradental lamellae.
Sides of pronotum reticulate-punctate, the pleurae and sides of the propodeum mostly smooth, with
punctures peripherally. Dorsal alitrunk, petiole and postpetiole reticulate-punctate. Spongiform appen-
dages of pedicel segments well developed. In profile the petiole with a straight narrow ventral strip; ventral
spongiform lobe of postpetiole equal to or slightly smaller than the exposed area of the postpetiolar disc in
profile, equalling or slightly larger than the lateral lobe. Basigastral costulae sharply developed but short.
Petiole, postpetiole and gaster dorsally with stout hairs which are swollen or clavate apically. Colour
brown.
The enormously expanded antennal scapes make tetraphanes one of the most easily recognized
Afrotropical Strumigenys and this character, coupled with the form of the mandibles, pilosity,
head width and sculpture, should make confusion of tetraphanes with any other species
impossible.
MATERIAL EXAMINED
Uganda: Kampala, Kawanda Exp. Sta. (G. Salt). Cameroun: Nkoemvon (D. Jackson); nr Yaounde (G.
Terron). Gabon: Plateau d'Ipassa (J. A. Band).
Strumigenys totyla sp. n.
(Fig. 56)
HOLOTYPE WORKER. TL 2-3, HL 0-64, HW 0-45, CI 70, ML 0-28, MI 44, SL 0-34, SI 76, PW 0-29, AL 0-58.
Apical fork of each mandible with 2 spiniform teeth, without intercalary teeth or denticles. Preapical
armament of each mandible of 2 stout teeth, the distal tooth about 0-75 x the length of the proximal. Outer
margins of mandibles shallowly convex in full-face view. Upper scrobe margins bordered by a conspicuous
laminar rim or flange throughout their length, the eyes visible in full-face view. Maximum diameter of eye
about 0-17xHW, the maximum diameter of the eye distinctly greater than the maximum width of the
scape. Preocular notch present and distinct on the ventrolateral cephalic margin but the anterior portion of
the eye not detached from the side of the head. Preocular notch ending at the ventrolateral margin, not
extending across the ventral surface as a groove or impression. Antennal scapes slightly bent in the basal
third, broadest at about the midlength and the leading edges equipped with a row of apically curved
narrowly spoon-shaped hairs which are only slightly smaller than those fringing the upper scrobe margins.
With the head in full-face view the sides behind the apices of the scrobe margins approximately straight and
convergent posteriorly. Ground-pilosity of head consisting of inconspicuous small spatulate to narrowly
396 BARRY BOLTON
spoon-shaped hairs. Upper scrobe margins with a row of anteriorly curved large spoon-shaped hairs.
Dorsum of head with an occipital transverse row of 4 stout standing hairs, without a pair situated anterior to
this row (it is possible that an anterior pair should be present but has been lost by abrasion in the holotype).
Head finely reticulate-punctate everywhere. Pronotal humeri without flagellate hairs. Mesonotum with a
single pair of stout standing hairs. Metanotal groove represented by a transverse line on the dorsum. In
profile the posterior portion of the mesonotum depressed behind the level of the standing hairs. Propodeal
teeth small, almost completely merged with the infradental lamellae and only with their extreme apices
projecting. Sides of pronotum finely superficially punctate, the pleurae mostly smooth but with peripheral
punctures; sides of propodeum finely punctate. Pronotal dorsum punctate and with irregular rugulae
formed by alignment of the punctures, postero-central portion with some longitudinal costulae. Re-
mainder of dorsal alitrunk and dorsum of petiole node reticulate-punctate, the postpetiole with some
scratch-like faint striae towards the sides of the disc but smooth medially. Spongiform appendages of
pedicel segments moderately developed, the petiole with a narrow ventral strip. Ventral spongiform lobe
of postpetiole slightly larger than the exposed area of the disc in profile. In dorsal view the postpetiole is
bounded by narrow spongiform strips both in front and behind, but spongiform material does not freely
project beyond the outline of the sides except posterolaterally. First gastral tergite with a lamellate basal
strip, the basigastral costulae short and radiating from the basal strip on each side of a clear central area.
Dorsal surfaces of petiole, postpetiole and gaster with standing stout hairs. Colour yellow.
Holotype worker, Cameroun: nr Yaounde, sample no. 1784 (G. Tenon) (ENSA).
Known only from the holotype, totyla belongs to that section of the/awra'-complex in which the
preocular notch is present but does not extend ventrally as an impression across the ventral
surface of the head. Among the six species falling into this category (relahyla, dyshaula,
xenohyla, totyla, adrasora, rukhd) the totyla holotype is easily recognized by its lack of pronotal
flagellate hairs and presence of only 4 standing hairs on the cephalic dorsum. In all the others
flagellate hairs are present on the pronotum and the cephalic dorsum has 6 standing hairs.
Unfortunately the universality of these characters among the closest relatives of totyla makes me
suspect that perhaps the hairs have been abraded away.
Strumigenys traegaordhi Santschi
Strumigenys traegaordhi Santschi, 19130: 257 (diagnosis in key). Syntype workers, SOUTH AFRICA: Natal,
Piertermaritzburg (/. Trdgardh) (NMB) [ examined].
Strumigenys traegaordhi Santschi; Santschi, 1914c: 28, fig. 4 (description).
Strumigenys traegaordhi Santschi; Brown, 1954: 26.
WORKER. TL 2-0-2-1, HL 0-54-0-55, HW 0-43, CI 78-80, ML 0-24, MI 44, SL 0-30, SI 70, PW 0-28, AL
0-59-0-60 (2 measured).
Apical fork of each mandible with 2 teeth, without intercalary teeth or denticles. Preapical armament of
each blade of 2 teeth, the proximal distinctly longer than the distal and both teeth situated in the apical third
of the length of the blade. Upper scrobe margins bordered by a narrow rim, the eyes not visible in full-face
view. Eyes small, their maximum diameter distinctly less than the maximum width of the scape. Preocular
notch absent, the ventral surface of the head without a preocular transverse groove or impression on each
side. Antennal scapes shallowly curved in the basal third and slightly expanded in the median third,
broadest at about the midlength. Leading edges of scapes shallowly convex and with a row of apically
curved narrowly spoon-shaped hairs. Dorsum of head with ground-pilosity which becomes narrower and
finer from front to back. Anteriorly, from the level of the posterior clypeal margin to about the level of the
ends of the preocular laminae the hairs are spoon-shaped and quite broad, appearing almost scale-like in
full-face view. Behind this level, and posteriorly over the highest point of the vertex to the occipital margin,
the hairs are much narrower and less conspicuous, narrowly spatulate in shape. Hairs fringing the upper
scrobe margins spoon-shaped and as large as those on the anterior portion of the cephalic dorsum. In
profile the dorsum of the head with 6 standing hairs arranged in a row of 4 close to the occipital margin and a
more anteriorly situated pair which are about at the highest point of the vertex. Dorsum of head
reticulate-punctate. Pronotal humeri each with an elongate fine flagellate hair, the mesonotum with two
pairs of standing hairs, the posterior pair of which is only half as long as the anterior pair. Ground-pilosity
of dorsal alitrunk of sparse narrowly spatulate hairs similar to those on the head behind the highest point of
the vertex. Metanotal groove present as a feeble line across the dorsum. Apical portions of propodeal
teeth, which are free from the narrow infradental lamellae, very narrowly triangular and almost spiniform.
Sides of alitrunk mostly smooth, the pleurae and sides of the propodeum with some peripheral punctures.
THE AFROTROPICAL DACETINE ANTS
397
Dorsal alitrunk reticulate-punctate, the pronotum also with some weak longitudinal rugulae. Dorsum of
petiole node reticulate-punctate, the postpetiole smooth or at most with a few feeble punctures posteriorly.
Spongiform appendages of pedicel segments moderately developed. In profile the petiole with a narrow
ventral strip and a small lateral lobe, the postpetiole with the ventral lobe slightly smaller than the exposed
area of the postpetiolar disc in profile. Basigastral costulae short but quite strongly defined. Petiole,
postpetiole and gaster with standing stout hairs which are swollen or feebly clavate apically . Colour brown.
The closest relative of traegaordhi is mesahyla, known from Zimbabwe. The two are superfi-
cially very similar but differ as follows.
traegaordhi
Dorsum of head with 6 standing hairs,
the hairs slender.
Hairs of cephalic ground-pilosity much
narrower posteriorly than anteriorly.
Mesonotum with 2 pairs of stout standing
hairs.
Reticulate-punctate sculpture predominant
on pronotum.
Ventral spongiform lobe slightly smaller
than exposed area of disc in profile.
Basigastral costulae arising across
entire width of first tergite.
MATERIAL EXAMINED
South Africa: Pietermaritzburg (/. Tragardh).
mesahyla
Dorsum of head with 4 standing hairs,
the hairs thick.
Hairs of cephalic ground-pilosity the
same everywhere on the head.
Mesonotum with a single pair of stout
standing hairs.
Longitudinal rugular sculpture predominant
on pronotum.
Ventral spongiform lobe much larger than
exposed area of disc in profile.
Basigastral costulae arising on each side
of a clear central area on first tergite.
Strumigenys vazerka sp. n.
(Fig. 52)
HOLOTYPE WORKER. TL 2-2, HL 0-58, HW 0-38, CI 66, ML 0-34, MI 59, SL 0-32, SI 84, PW 0-26, AL 0-54.
Mandibles in full-face view long, noticeably divergent from base to apex, and with the outer margins of the
blades convex. Apical fork of each mandible with 2 spiniform teeth, without intercalary teeth or denticles.
Preapical armament of each mandible of 2 teeth , a long spiniform proximal and a short distal . Upper scrobe
margins not bounded by a projecting rim or flange, close together behind the frontal lobes and evenly
divergent posteriorly; not concave or impressed above the eyes but still quite close together so that the eyes
are clearly visible in full-face view. Preocular notch present, deep and conspicuous, the anterior portion of
each eye detached from the side of the head. Preocular notch continued onto ventral surface of head as a
broad groove or impression. Maximum diameter of eye greater than the maximum width of the scape.
Antennal scapes relatively long, straight and slender, their leading edges with a row of narrowly spatulate
hairs which are directed apically. Ground-pilosity of cephalic dorsum inconspicuous, of short narrowly
spatulate hairs which are curved anteriorly. Upper scrobe margins bordered by a row of anteriorly curved
spoon-shaped hairs which are distinctly much larger than the cephalic ground-pilosity. In profile the
dorsum of the head with 6 approximately erect simple hairs which are arranged in a transverse row of 4
close to the occipital margin and a more anteriorly situated pair. Dorsum of head finely and shallowly
reticulate-punctate. Pronotal humeri each with a fine flagellate hair. Mesonotum with a single pair of stout
erect hairs which are broadened apically; dorsal alitrunk otherwise without standing pilosity; the
ground-pilosity of minute sparse hairs which are closely applied to the surface. Posterior portion of
mesonotum depressed and on the same level as the propodeum, the metanotal groove represented by a
transverse line on the dorsum but not impressed. Propodeum armed with a pair of short triangular teeth
which are subtended by a narrow infradental lamella on each side. Sides of alitrunk smooth, unsculptured
except for some vestigial punctulation on the posterior propodeum. Pronotal dorsum feebly longitudinally
costulate or striate, without punctate sculpture. Anterior portion of mesonotum with vestigial punctures,
the depressed posterior portion more strongly punctate. Propodeal dorsum mostly smooth, with a very few
vestigial punctures laterally. Petiole node punctate-granular dorsally, the postpetiole with some sparse
vestigial longitudinal costulae, most conspicuous towards the sides of the disc. In profile the petiole with a
narrow ventral spongiform strip. Lateral and ventral spongiform lobes of postpetiole moderately de-
veloped. In dorsal view the postpetiole with a transverse narrow spongiform strip posteriorly and the first
gastral tergite with a similar but even narrower strip anteriorly. Basigastral costulae sharply defined.
398 BARRY BOLTON
Petiole, postpetiole and gaster with stout standing hairs which are weakly clavate apically. Colour dull
yellowish brown.
PARATYPE WORKERS. TL 1-9-2-2, HL 0-52-0-60, HW 0-36-0-40, CI 65-70, ML 0-28-0-34, MI 50-59, SL
0-29-0-34, SI 79-86, PW 0-22-0-28, AL 0-46-0-54 (15 measured).
As holotype but sculpture of alitrunk showing some variation. Sides usually smooth but in some
peripheral faint punctulae are present. Dorsum of pronotum usually as holotype but in some the costulae
are more pronounced and quite strong, and in others there is a faint punctulate component visible between
the costulae. Anterior portion of mesonotum sometimes as distinctly punctate as the depressed posterior
portion. Propodeal dorsum usually smooth but often with fine faint lateral or peripheral punctulae, but
never punctulate all over the surface.
Holotype worker, Ivory Coast: Man, Mt Tonkoui, 900 m, 13.x. 1980 (V. Mahnert & J.-L. Ferret)
(MHN).
Paratypes. Ivory Coast: 11 workers with same data as holotype; 22 workers and 2 females, Tai Forest,
17.x. 1980 (V. Mahnert & J.-L. Ferret) (MHN; BMNH; MCZ; ENSA).
Non-paratypic material examined. Ivory Coast: Divo (L. Brader); Abidjan, Banco Forest (W. L.
Brown); Banco Nat. Pk. (V. Mahnert & J.-L. Ferret); Sassandra (V. Mahnert & J.-L. Ferret). Ghana: Mt
Atewa (D. Leston). Nigeria: Gambari (B. Bolton).
The closest relative of vazerka is the Central African bernardi, but in the latter the left
mandibular blade has lost the distal preapical tooth and the propodeal dorsum is reticulate-
punctate.
Strumigenys xenohyla sp. n.
(Figs 54, 73)
HOLOTYPE WORKER. TL 2-3, HL 0-60, HW 0-47, CI 78, ML 0-29, MI 48, SL 0-33, SI 70, PW 0-27, AL 0-58.
Apical fork of each mandible with a pair of spiniform teeth, without intercalary teeth or denticles.
Preapical armament on each mandibular blade of 2 teeth, both spiniform but the proximal much the
longest . Length of the distal preapical tooth more than twice that of the distance separating the bases of the
2 preapical teeth. Upper scrobe margins bordered by a relatively broad conspicuous projecting lamina
which has an irregular free margin and which partially conceals the eyes in full-face view. Eyes of moderate
size, about 0-15xHW but only fractionally larger than the maximum width of the scape because of the
broadening of the latter. Preocular notch present but shallow, the anterior portion of the eye not detached
from the side of the head and the notch not extending onto the ventral surface of the head as a transverse
groove or impression. Antennal scapes shallowly curved basally, broadened in the middle and slightly
dorsoventrally flattened. Leading edges of scapes convex and weakly undulate, the undulate rim forming a
narrow flange from which the large spoon-shaped hairs arise; these hairs are about equal in size to those on
the upper scrobe margins. Ground-pilosity of cephalic dorsum inconspicuous, of narrow spoon-shaped
hairs. Upper scrobe margins with an anteriorly curved row of large spoon-shaped hairs. Cephalic dorsum
with 6 standing hairs arranged in a transverse row of 4 close to the occipital margin and an anterior pair
close to the highest point of the vertex. Dorsum of head reticulate-punctate. Pronotal humeri each with a
long fine flagellate hair. Mesonotum with a single pair of stout standing hairs which are broadly clavate
apically. Ground-pilosity of dorsal alitrunk of sparse spatulate to narrowly spoon-shaped hairs which are
closely applied to the surface. Mesonotum suddenly and steeply depressed behind the level of the hairs.
Metanotal groove represented by a line across the dorsum. Propodeal teeth lamellate and confluent with
the broad infradental lamellae for more than half their length. Sides of pronotum with a few faint
scratch-like marks. Pleurae and sides of propodeum smooth. Pronotal dorsum finely longitudinally
costulate, without punctures. Remainder of dorsal alitrunk almost smooth, with only the vaguest traces of
punctulate sculpture present. Dorsum of petiole node granular, the postpetiole smooth. Spongiform
appendages of pedicel segments well developed, the petiole with a ventral spongiform strip which is more
than half the depth of the peduncle, and with a broadly triangular lateral lobe. Ventral spongiform
appendage of postpetiole large, larger than the lateral lobe and distinctly much larger than the exposed
area of the postpetiolar disc in profile. In dorsal view the petiole node with a lamellate collar posteriorly.
Sides of postpetiole with projecting spongiform tissue visible. Base of first gastral tergite with a lamellar
transverse strip from which the sparse basigastral costulae radiate on each side of a central smooth area.
Petiole, postpetiole and gaster dorsally with stout standing hairs which are thickened apically. Colour dull
yellow.
THE AFROTROPICAL DACETINE ANTS 399
PARATYPE WORKER. TL 2-3, HL 0-61, HW 0-48, CI 79, ML 0-30, MI 49, SL 0-34, SI 71, PW 0-27, AL 0-58.
As holotype.
Holotype worker, Cameroun: Nkoemvon, N22, 7.ix.l980 (D. Jackson) (BMNH).
Paratype. 1 worker with same data as holotype (BMNH).
Non-paratypic material examined. Cameroun: nr Yaounde (G. Terrori). Zaire: Ituri Forest, vie. Epulu
(T. Gregg}.
Measurements of the two non-paratypic specimens show HL 0-64-0-67, HW 0-53, CI 79-83, ML
0-32-0-33, MI 48-51, SL 0-36, SI 68.
Among the species in which the preocular notch is present but not extended as a groove or
impression across the ventral surface of the head, xenohyla is recognized by its broad flange or
rim bordering the upper scrobe margins, flattened scapes, large spoon-shaped hairs on the
scapes and upper scrobe margins which are about equal in size, reduced sculpture on the dorsal
alitrunk and well-developed spongiform appendages.
Strumigenys zandala sp. n.
(Fig. 66)
HOLOTYPE WORKER. TL 2-3, HL 0-60, HW 0-42, CI 70, ML 0-27, MI 45, SL 0-34, SI 81, PW 0-27, AL 0-60.
Mandibles slender in full-face view, the external margins very shallowly evenly convex and the blades
about the same width from the proximal preapical tooth to the base where they are somewhat narrowed
and inflected. Apical fork of each blade with 2 teeth, without intercalary teeth or denticles. Each
mandibular blade with 2 preapical teeth, the proximal longer and slightly stouter than the distal, both teeth
distinctly within the apical third of the length of the blade. Distance separating the bases of the preapical
teeth less than the length of the distal tooth . Upper scrobe margins evenly and shallowly convex in full-face
view, the eyes visible, the apices of the upper scrobe margins confluent with the sides of the occipital lobes
through an even curve, without an angle separating the two. Eyes moderate, with 19-20 ommatidia, the
maximum diameter greater than the maximum width of the scape. Preocular notch absent, the ventral
surface of the head without a transverse preocular groove or impression. Antenna! scapes slender and
subcylindrical, curved weakly near the base and with their leading edges equipped with a row of slender
flattened hairs which are narrowly spatulate to spoon-shaped and curved towards the apex. Dorsum of
head densely clothed with curved narrowly spatulate to slender spoon-shaped ground-pilosity, the upper
scrobe margins fringed by a dense row of similar hairs, these hairs slightly larger than those on the scapes.
Dorsum of head with 6 standing hairs arranged in a row of 4 close to the occipital margin and a more
anteriorly situated pair. Dorsum of head densely reticulate-punctate. Pronotal humeri each with a single
fine flagellate hair. Mesonotum with a single pair of simple standing hairs. Ground-pilosity of dorsal
alitrunk of narrow curved flattened hairs. Posterior portion of mesonotum shallowly depressed behind the
level of the hairs. Metanotal groove feebly impressed. Propodeal teeth triangular and subtended by narrow
infradental lamellae. Sides of pronotum superficially punctulate and with some feeble rugulae anteriorly.
Upper third to half of mesopleuron, upper third of metapleuron and portion of propodeum above and
behind the spiracle densely punctate; lower portions of these segments smooth. Pronotal dorsum very
feebly longitudinally rugulose and with punctate sculpture, remainder of dorsal alitrunk and petiole node
reticulate-punctate. Postpetiole smooth. Spongiform appendages of pedicel segments well developed, the
petiole with a broad ventral strip. Postpetiole with large lateral and ventral lobes, the latter larger than the
former and larger than the exposed area of the disc in profile. In dorsal view the petiole node broader than
long and the postpetiole surrounded by spongiform tissue, the lateral lobes projecting sideways beyond the
outline of the disc. Basigastral costulae fine, arising on each side of a central clear area. Dorsal surfaces of
petiole, postpetiole and gaster with simple standing hairs. Colour light brownish yellow.
PARATYPE WORKERS. TL 2-2-2-3, HL 0-59-0-61, HW 0-42-0-44, CI 70-73, ML 0-26-0-27, MI 43-45, SL
0-33-0-35, SI 75-81, PW 0-24-0-30, AL 0-57-0-63 (10 measured). As holotype.
Holotype worker, Equatorial Guinea: Annobon I., 400-500 m, v.1902 (L. Feo) (MCSN).
Paratypes. 27 workers with same data as holotype (MCSN; BMNH; MCZ).
Non-paratypic material examined. Cameroun: nr Yaounde (G. Terron).
S. zandala is closely related to scotti and hastyla. It is separated from the first of these by its
shorter mandibles and scapes and by the shape of the petiole node in dorsal view which is
broader than long in zandala and at least as long as broad (sometimes longer than broad) in
400 BARRY BOLTON
scotti. S. hastyla is a smaller species than zandala and has the standing hairs on the gaster
distinctly swollen or flattened apically in dorsal view. There is a possibility that hastyla and
zandala may represent extremes of a single species but for the present I am treating them as
separate species.
QUADRISTRUMA Brown
(Fig. 67)
Quadristruma Brown, 1949ft: 47. Type-species: Epitritus emmae Emery, 1890: 70, pi. 8, fig. 6, by original
designation.
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles linear and curved, relatively short (MI
26-32) , produced into narrow blades and equipped apically with a strong fork of two long spiniform teeth in
a vertical series, the dorsal tooth the longest. Inner margin of each mandibular blade with a long spiniform
proximal preapical tooth and a small distal preapical denticle. Antennae with 4 segments. Scape with the
leading edge angled but without a subbasal lobe. Head with scale-like to orbicular hairs present. Labral
lobes very short and inconspicuous.
This small genus was erected to hold the species emmae and eurycera (Emery), both of which
were originally described in the genus Epitritus because of their 4-merous antennae. Brown
(19496) showed that these two species, although seemingly similar to Epitritus, were in reality
convergently so and had been derived from Strumigenys rather than from a Smithistruma-\ike
ancestor as is the case with Epitritus. With the description of so many new Strumigenys and
Epitritus since 1949 this discovery has been amply confirmed and in fact the differences between
Strumigenys and Quadristruma have been narrowed down to the antennomere count, with 6
segments in the former and 4 in the latter genus. As seen in other dacetine genera (Epitritus,
Smithistruma) this difference is insignificant at genus-level and it is likely that Quadristruma will
eventually fall into the synonymy of Strumigenys.
Of the two species in Quadristruma one, eurycera, is known only from New Guinea, but the
second species, emmae, is very widespread in the tropics by dint of being a successful
tramp-species. Initially Brown (19496) thought that the original range of emmae lay in the
Indomalayan-Papuan area but later he revised this opinion (Brown, 1954) in favour of the
Afrotropical region as the place of origin, as a derivation from the Strumigenys rogeri-group (in
the sense of the 1954 paper, the arnoldi-comp\ex of the present study) seemed certain. At that
time emmae had not been found in sub-Saharan Africa but it was known from Hawaii, Guam,
the U.S.A. (Florida), Puerto Rico, the West Indies, Cuba, Surinam, Sumatra, Singapore, and
New Guinea (listed by Brown, 19496). Wilson & Taylor (1967) added the Philippines, New
Hebrides and Australia (Queensland) to the list, and Kempf (1972) the Bahamas. Soon after this
the species was detected for the first time in West Africa, being recorded from Ghana (Bolton,
1973). The present paper adds localities in India, Malaysia, Sulawesi and Equatorial Guinea to
the list, showing that emmae is indeed a very accomplished tramp-species and likely to be found
in any tropical area of the world.
Quadristruma emmae (Emery)
(Fig. 67)
Epitritus emmae Emery, 1890: 70, pi. 8, fig. 6. Holotype worker, ST THOMAS I. (West Indies) (MCSN)
[examined].
Epitritus dypeatus Szabo, 1909: 1, figs la, c. Syntype workers, NEW GUINEA: Berlinhafen (L. Biro); and
SINGAPORE (L. Biro) (TM). [Synonymy by Brown, 1949ft: 48.]
Epitritus dypeatus var. malesiana Forel, 1913a: 83. Syntype workers and female, INDONESIA: Sumatra
(Buttel-Reepen) (MHN). [Synonymy by Brown, 1949ft: 48.]
Epitritus wheeleri Donisthorpe, 1916: 121. Holotype worker, HAWAII: Oahu, Honolulu (R. C. L. Perkins)
(not in BMNH; presumed lost). [Synonymy by Brown, 1949ft: 48.]
Quadristruma emmae (Emery) Brown, 1949ft: 48.
THE AFROTROPICAL DACETINE ANTS 401
WORKER. TL 1-7-1-9, HL 0-43-0-46, HW 0-35-0-39, CI 80-85, ML 0-12-0- 14, MI 26-32, SL 0-18-0-22 SI
52-58, PW 0-21-0-25, AL 0-42-0-48 (15 measured).
Mandibles a pair of narrow linear outcurved blades, armed apically with a fork of 2 spiniform teeth of
which the upper is the longer. Between the fork teeth the left mandible has 2, and the right mandible 1 or 2,
minute intercalary denticles which cannot be seen when the mandibles are closed. Preapical armament
consisting on both blades of a single long spiniform tooth at about the apical third of the length and usually
also a minute denticle on the margin between the spiniform preapical tooth and the upper tooth of the
apical fork, though in some samples this denticle is extremely small and inconspicuous. Anterior clypeal
margin broad, projecting well beyond the mandibular bases on each side, with a feeble median impression
and with numerous small spatulate to spoon-shaped hairs which are curved towards the midline. Lateral
margins of clypeus short and with 2-3 anteriorly curved small spoon-shaped hairs. Preocular laminae
broad, running back from the clypeus and anteriorly forming a strong floor below the antennal insertions.
Median portion of clypeus broad, finely punctulate and with scale-like to suborbicular hairs present.
Dorsum of head behind clypeus reticulate-punctate and with numerous broadly scale-like to orbicular
hairs. Outer margins of frontal lobes and divergent upper scrobe margins behind them with a continuous
row of scale-like to orbicular hairs, the row terminating at the posterior end of the scrobe in a more or less
straight clavate hair on each side. Eyes very small, situated just above the ventral scrobe margin. Antennae
with 4 segments, the scapes narrow basally but broadening to the midlength then narrowing again to the
apex, the leading edge angular and prominent at about the midlength , with a row of projecting scale-like to
spoon-shaped hairs. Pronotum more or less flat dorsally, anteriorly rounding into the sides, posteriorly
meeting the sides in a broad blunt angle. In profile the mesonotal dorsum very shallowly convex anteriorly,
very shallowly concave posteriorly before meeting the propodeum. Metanotal groove absent. Propodeal
dorsum shallowly convex anteriorly, sloping posteriorly to the declivity. Propodeal teeth mostly incor-
porated in the infradental lamellae, with only a small point projecting. Sides of alitrunk smooth to
superficially reticulate. Dorsal alitrunk and at least the upper half of the propodeal declivity reticulate-
punctate, the punctures more strongly defined and denser on the pronotum than on the propodeum, where
they may be superficial. Pronotal humeri each with a straight clavate hair and mesonotum with a similar but
shorter pair of hairs. Ground-pilosity of dorsal alitrunk consisting of numerous small scale-like to broadly
spoon-shaped hairs, most of which are closely applied to the surface. In profile the pedicel segments with
moderately developed spongiform appendages. The ventral petiolar strip broad and distinct but the lateral
lobe of the petiole node small. Lateral and ventral lobes of the postpetiole moderate and a spongiform pad
present at the base of the first gastral sternite. Dorsal surface of petiole node finely punctate to reticulate,
the postpetiole superficially reticulate to smooth. Posterior spongiform strip of petiole node very narrow,
narrower than the strip bordering the anterior margin of the postpetiole. Sides of postpetiole in dorsal view
surrounded by spongiform tissue and posteriorly with a narrow bordering strip. Base of first gastral tergite
lamellar centrally, spongiform towards the sides, with a continuous row of basal costulae. Petiole,
postpetiole and gaster with short straight narrowly clavate hairs. Colour dull yellow to pale brown.
MATERIAL EXAMINED
Ghana: Bunso (P. Room). Equatorial Guinea: Annobon I., Dint, del Pueblo (L. Fed). Seychelles:
Aldabra I., Grande Terre (V. Spaul); Pt Hodoul (V. Spaul); Picard (V. Spaul); Big Sister I. (U. Mutter}.
India: no loc. (P. C. W. Westall). West Malaysia: Sg Patani (G. H. Lowe). Indonesia: Sulawesi Tengah,
Morowali (M. /. D. Brendall).
MICRODACETON Santschi
(Figs 78-81)
Microdaceton Santschi, 19136: 478. Type-species: Microdaceton exornatum Santschi, 19136: 478, by
monotypy.
DIAGNOSIS OF WORKER. Afrotropical dacetine ants. Mandibles extended into elongate linear blades (MI
55-69) which terminate in an apical fork of 3 spiniform teeth arranged in a vertical series. Mandibular
blades without preapical teeth or denticles. Palp formula 3,2 (as opposed to 1 ,1 in other African dacetines).
Antennal scrobes absent; antennae with 6 segments. Petiole node armed with a pair of teeth or short spines
dorsally. Postpetiole lacking spongiform appendages but with lateral alar extensions. Specialized body
pilosity absent. Eyes dorsolateral.
Microdaceton, the only Afrotropical member of its genus-group, is closely related to the
primarily Australian genera Colobostruma, Mesostruma and Epopostruma (Brown, 19526;
1953a; Brown & Wilson, 1959; Taylor, 1973). Within the Dacetini this group of genera, the
402 BARRY BOLTON
subtribe Epopostrumiti, is defined by having the eyes placed dorsolaterally, above the scrobes
when such are present; the antennae 4- or 6-segmented (the second funicular segment not longer
than the rest); the palp formula 3,2 or 5,3; the postpetiole usually with lateral alar extensions.
Both known species of Microdaceton are restricted to the Afrotropical region, tibialis being
found in West and central Africa, and exornatum being widely distributed in East and South
Africa.
List of Afrotropical Microdaceton
exornatum Santschi
leakeyi Patrizi syn. n.
exornatum var. laevior Arnold syn. n.
tibialis Weber
Key to species (workers)
1 Postpetiole in dorsal view very broad, spanning almost the entire basal width of the first gastral
tergite (Fig. 80). The width of the postpetiole 0-60-0-65 times the maximum width of the first
gastral tergite. Basigastral costulae usually distinct, rarely faint. Body colour yellow. (Kenya,
Zambia, Zimbabwe , South Africa) exornatum (p. 402)
- Postpetiole in dorsal view narrow, spanning 0-70 or less of the basal width of the first gastral
tergite (Fig. 81). The width of the postpetiole 0-46-0-56 times the maximum width of the first
gastral tergite. Basigastral costulae absent. Body colour blackish brown to black. (Ivory
Coast, Ghana, Zaire) tibialis (p. 403)
Microdaceton exornatum Santschi
(Figs 78, 80)
Microdaceton exornatum Santschi, 19136: 478. Holotype worker, SOUTH AFRICA: Natal Zululand (/.
Tragardh) (not in NMB; presumed lost).
Microdaceton leakeyi Patrizi, 1947: 219, figs 1, 2. Holotype female, KENYA: Masai Reserve, Olorgasalic,
iv.1945 (S. Patrizi) (IE) [not examined]. Syn.n.
Microdaceton exornatum var. laevior Arnold, 1948: 225. Syntype workers and female, SOUTH AFRICA:
Natal, Zululand, Dukuduku; and Natal, Richard's Bay, February (/. C. Faure) (SAM) [examined].
Syn. n.
WORKER. TL 3-0-4-0, HL 0-79-1 -00, HW 0-76-0-94, CI 92-96, ML 0-48-0-58, MI 55-61 , SL 0-50-0-64, SI
66-70, PW 0-40-0-50, AL 0-70-0-90 (10 measured).
Mandibles elongate and linear, without preapical armament but armed apically with a fork of 3 long
spiniform teeth set in a more or less vertical series, the apical fork teeth without intercalary denticles.
Anterior clypeal margin with a small median notch or indentation. Eyes large and conspicuous, clearly
visible in full-face view. Antennal scrobes absent, frontal carinae absent, the antennal fossa ventrally on
each side with a small laterally projecting tubercle in front of the eye. Outline shape of head as in Fig. 79.
Occipital lobes with 2 pairs of tubercles which are variable in size, the first pair laterodorsal, the second pair
at the posteriormost point of the lobes. Clypeus coarsely punctate to narrowly foveolate, with appressed to
slightly elevated fine simple ground-pilosity, without standing or specialized hairs of any description.
Dorsum of head foveolate, with a fine short simple hair arising from the centre of each foveola, the hairs
appressed or nearly so, the head without specialized or standing pilosity . In profile the dorsal surface of the
head rising and shallowly convex from the posterior clypeal margin to about the midlength, then suddenly
depressed. Sides of head foveolate as dorsum. Dorsal outline of alitrunk dominated by the strong
subconical mesonotal teeth or tubercles and the long propodeal spines (Fig. 78), the latter without or only
with a vestige of an infradental lamella. Metapleural lobes long and broad, slightly upcurved. Sides of
alitrunk foveolate but on the mesopleuron the sculpture may be partially or almost wholly effaced. Dorsal
surfaces of pronotum and mesonotum strongly foveolate, the metanotal groove with short longitudinal
cross-ribs. Propodeal dorsum reticulate-punctate, sometimes with one or two laterally situated partial
foveolae. Alitrunk without specialized or bizarre pilosity, only with fine short simple hairs arising from the
foveolar centres. Usually these hairs are very short inconspicuous and appressed, but in some they may be
longer and slightly elevated. Petiole in profile without spongiform or alar appendages, armed dorsally with
a pair of spines and posterodorsally with a sharp triangular elevation. Postpetiole without spongiform
tissue but with strong lateral alar prominences which appear in profile as thick longitudinal crests. In dorsal
THE AFROTROPICAL DACETINE ANTS 403
view the postpetiole very broad, spanning almost all of the basal width of the first gastral tergite.
Basigastral costulae present, usually fine dense and distinctive, only rarely reduced in intensity. Petiole and
postpetiole densely reticulate-punctate to granular. Petiole, postpetiole and first gastral tergite without
standing hairs of any description, only with minute appressed sparse pubescence. Colour yellow, the
appendages paler than the body.
MATERIAL EXAMINED
Zambia: Kipushi (//. 5. Evans). Zimbabwe: Gwebi (K. J. Wilson), Chishawasha (A Watsham). South
Africa: Natal, St Lucia Lake (/. C. Faure).
Microdaceton tibialis Weber
(Figs 79, 81)
Microdaceton tibialis Weber, I952a: 30, fig. 25. Holotype worker, ZAIRE: 37 km N. of Stanleyville (=
Kisangani), lat. 0°45'N, long. 25°15'E, 15.iii.1948, rain forest, no. 2218 (N. A. Weber) (AMNH)
[examined].
WORKER. TL 3-2-3-8, HL 0-88-0-98, HW 0-80-0-90, CI 88-92, ML 0-56-0-66, MI 63-69, SL 0-62-0-72, SI
75-81, PW 0-38-0-47, AL 0-78-0-90 (10 measured).
Answering to the description ofexornatum in all major features, tibialis is distinguished as follows.
exornatum
Mandibles relatively shorter, MI 55-61.
Scapes relatively shorter, SI 66-70.
Postpetiole in dorsal view spanning
almost the entire basal width of the
first gastral tergite (Fig. 80).
Width of postpetiole in dorsal view
0-60-0-65 x maximum width of first
gastral tergite.
Basigastral costulae usually dense and
distinct, rarely reduced in intensity.
Body colour yellow.
Laterodorsal cephalic tubercles large
and conspicuous.
Petiolar armament spiniform.
Propodeal dorsum with reticulate-
punctate sculpture.
tibialis
Mandibles relatively longer, MI 63-69.
Scapes relatively longer, SI 75-81.
Postpetiole in dorsal view spanning 0-70
or less of the basal width of the first
gastral tergite (Fig. 81).
Width of postpetiole in dorsal view
0-46-0-56x maximum width of first
gastral tergite.
Basigastral costulae absent, at most with
faint shagreening near gastral base.
Body colour black to blackish brown.
Laterodorsal cephalic tubercles vestigial
to absent.
Petiolar armament dentiform.
Propodeal dorsum without reticulate-
punctate sculpture.
MATERIAL EXAMINED
Ivory Coast: Man, Mt Tonkoui (V. Mahnert & J.-L. Ferret). Ghana: Mampong (P. Room); Bunso (D.
Lesion). Zaire: Kisangani (N. A. Weber).
Acknowledgements
I would like to express my thanks and gratitude to the following people, who have greatly
facilitated this study by lending me types and other material and who have given me free access
to their collections and other data: Dr Cesare Baroni Urbani (NMB); Dr Claude Besuchet
(MHN); Professor William L. Brown Jr (Cornell University, Ithaca); Dr Jean Decelle
(MRAC); Mrs Marjorie Favreau (AMNH); Dr Max Fischer (NMV); Dr Dorothy A. Jackson
(Oxford University); Dr Frank Koch (MNHU); Dr Egidio Mellini (IE); Mr Alfred F. Newton
(MCZ); Dr Jeno Papp (TM); Dr Roberto Poggi (MCSN); Dr Wojciech Pulawski (CAS); Dr
David R. Smith (USNM); Dr George Terron (ENSA); Dr Janine C. Weulersse (MNHN);
Dr V. B. Whitehead (SAM).
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404 BARRY BOLTON
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406 BARRY BOLTON
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THE AFROTROPICAL DACETINE ANTS 407
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1916. Ants collected in Trinidad by Professor Roland Thaxter, Mr F. W. Urich, and others. Bulletin
of the Museum of Comparative Zoology at Harvard College 60: 323-330, 1 fig.
1922. Ants of the Belgian Congo, parts 1-8. Bulletin of the American Museum of Natural History 45:
1-1139, 23 pis, 76 figs.
1928. Ants collected by Professor F. Silvestri in China. Bollettino del Laboratorio di Zoologia
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1933. An ant new to the fauna of the Hawaiian Islands. Proceedings of the Hawaiian Entomological
Society 8: 275-278, 1 fig.
408 BARRY BOLTON
Wheeler, G. C. & Wheeler, J. 1954. The ant larvae of the myrmicine tribes Basicerotini and Dacetini.
Psyche, a Journal of Entomology 61: 111-145.
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America 46 (1953): 479-495.
1962. Behaviour of Daceton armigerum (Latreille), with a classification of self-grooming movements
in ants. Bulletin of the Museum of Comparative Zoology at Harvard College 127: 401-422, 2 figs.
Wilson, E. O. & Taylor, R. W. 1967. The ants of Polynesia. Pacific Insects Monograph 14: 1-109, 84 figs.
Wilson, E. O. & Brown, W. L., Jr 1956. New parasitic ants of the genus Kyidris, with notes on ecology and
behaviour. Insectes Sociaux 3: 439-454, 5 figs.
THE AFROTROPICAL DACETINE ANTS
409
Figs 1-13 Smithistruma workers. 1-12, heads of(\)fulda, (2) ninda, (3) sharra, (4) behasyla, (5) chyatha,
(6) impidora, (7) terroni, (8) emarginata, (9) datissa, (10) truncatidens , (11) gatuda, (12) hensekta. 13,
profile of truncatidens . Fringing pilosity only indicated except in 3-6.
410
BARRY BOLTON
Figs 14-26 14-20, Smithistruma workers. 14, 15, heads of (14) minkara, (15) enkara. 16, alitrunk and
pedicel of kerasma. 1 7-20, heads of (17) rusta, \8anarta,(19)oxysma,(2Q)vodensa.21,22, Trichoscapa
workers. 21, head of membranifera, 22, profile of membranifera. 23-26, Glamyromyrmex workers,
heads of (23) crypturus, (24) sistrurus, (25) dagon, (26) sahurus. Fringing pilosity only indicated.
THE AFROTROPICAL DACETINE ANTS
411
Figs 27-39 27-33. Glamyromyrmex workers. 27-29, heads of (27), africanus, (28) ravidurus, (29)
tukultus. 30, profile of tetragnathus . 31-33, profile of head of (31) thuvidus, (32) dagon, (33) sahurus.
34-39. Serrastruma workers. 34-37, heads of (34) ludovici, (35) dotaja, (36) geoterra, (37) maynei.
38-39, alitrunk and pedicel of (38) dotaja, (39) miccata. Fringing pilosity only indicated except in 36, 37.
412
BARRY BOLTON
o
Figs 40-48 40-44. Serrastruma workers, alitrunk and pedicel of (40) simoni, (41) ludovici, (42) serrula,
(43) concolor, (44) lujae. 45, mandibles of Cladarogenys lasia worker (from Brown, 1976). 46-48.
Epitritus workers, heads of (46) minimus, (47) room/, (48) laticeps.
THE AFROTROPICAL DACETINE ANTS
413
56
Figs 49-59 Strumigenys workers. Heads of (49) cacaoensis, (50) sarissa, (51) rogeri, (52) vazerka, (53)
pretoriae, (54) xenohyla, (55) dyshaula, (56) tofy/a, (57) relahyla, (58) petiolata, (59) pallestes . Fringing
pilosity only indicated.
414
BARRY BOLTON
Figs 60-73 60-66 Strumigenys workers. Heads of (60) tetraphanes, (61) katapelta, (62) spathoda, (63)
omalyx, (64) korahyla, (65) havilandi, (66) zandala. 67, head of Quadristruma emmae worker. 68-73
Strumigenys workers. 68-70, alitrunk and pedicel of (68) adrasora, (69) mesahyla, (70) havilandi. 71-73,
profile of head of (ll)petiolata, (72) rogeri, (73) xenohyla. Fringing pilosity^only indicated except in 67.
THE AFROTROPICAL DACETINE ANTS
415
78
81
Figs 74-81 74—77 Strumigenys workers. 74-76, profile of head of (74) relahyla, (75) scotti, (76) katapelta.
77, profile oimurshila. 78-81 Microdaceton workers. 78, profile ofexornatum. 79, head oltibialis. 80-81,
petiole, postpetiole and base of first gastral tergite in dorsal view of (80) exornatum, (81) tibialis.
416
BARRY BOLTON
Index
Synonyms are in italics.
adrasora 364
fenkara 302
membranifera 319
sharra 295
aequalis 345
foochowensis 319
mesahyla 379
shaula 392
africanus 322
fulda 282
miccata 348
silvestriana 319
alluaudi 343
fusciventris 350
Miccostruma 274
similllma 319
anarta 314
Microdaceton 401
simoni 350
arahana 300
gatuda 292
minimus 355
sistrurus 329
arnoldi 365
geoterra 341
minkara 306
Smithistruma 274
gerardi 345
murshila 380
spathoda 393
behasyla 286
Glamyromyrmex 320
Strumigenys 358
bequaerti 345
glanduscula 345
nigeriensis 343
stygia 394
bernardi 366
nimbrata 381
sulfurea 387
biconvexa 350
hastyla 372
ninda 284
sulumana 352
bitheria 367
havilandi 373
nykara 307
synkara 309
boerorum 350
helytruga 374
Borgmeierita 320
hensekta 293
obscuriventris 350
tacta 317
omalyx 382
terroni 299
cacaoensis 367
impidora 294
oxysma 315
tetragnathus 323
calypso 345
incisa 387
tetraphanes 395
cavinasis 287
inquilina 342
pallestes 383
thuvidus 332
Cephaloxys 274
irrorata 375
paranax 383
tibialis 403
chyatha 288
petiolata 384
tiglath 357
Cladarogenys 353
katepelta 375
placora 308
tigriUa 284
cliens 350
kerasma 303
Platystruma 274
tolomyla 310
clypeatus 400
korahyla 376
pretoriae 385
totyla 395
cognata 350
Proscopomyrmex 358
traegaordhi 396
concolor 338
Labidogenys 358
Pyramica 358
transversa 297
crypturus 327
laevior 402
Trichoscapa 319
lasia 353
Quadristruma 400
truncatidens 296
dagon 325
laticeps 355
trymalus 333
datissa 289
latiuscula 347
ravidurus331
tukultus 334
dendexa 290
leakeyi 402
raymondi 350
dextra 368
limbata 350
relahyla 386
uelensis 349
dotaja 339
londianensis 377
reticulata 345
dromoshaula 369
to/ft' 343
rogeri 387
vazerka 397
dyshaula 370
loveridgei 328
roomi 356
vitiensis 319
ludovici 343
rothkirchi 343
vodensa 317
emarginata 291
lujae 345
rufobrunea 389
emmae 400
rukha 389
weberi311
Eneria 358
malaplax 304
rusta313
Weberistruma 274
enkara 301
malesiana 400
Wessonistruma 274
Epitritus 354
mandibularis 283
sahurus 326
wheeled 400
escherichi 350
marginata 312
santschii 319
williamsi 319
ettillax 371
marioni3l9
sarissa 390
exornatum 402
marleyi 378
scotti 391
xenohyla 398
maynei 347
Serrastruma 335
faurei 371
mekaha 305
serrula 349
zandala 399
British Museum (Natural History)
Chance, change & challenge
Two multi-author volumes from one of the foremost scientific institutions in the world.
General Editor: P. H. Greenwood
The Evolving Earth
Editor: L. R. M. Cocks
The Evolving Biosphere
Editor: P. L. Forey
In the first volume, The Evolving Earth, twenty scientists have been asked to review the present
state of knowledge in their particular field, ranging from the origin of the Earth, through ocean
sediments and soils to continental drift and palaeogeography.
In the companion volume, The Evolving Biosphere, museum scientists have chosen an
evolutionary concept — speciation, coe volution, biogeography etc. and related this to the group
of animals or plants in which they are specialising. Thus beetles and birds exemplify sympatric
and allopatric speciation, butterflies mimicry and certain fishes explosive evolution.
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These two books will be invaluable to all sixth-form and undergraduate biology and geology
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The Evolving Earth: 276 x 219 mm, 280pp, 138 line illustrations, 42 halftones
The Evolving Biosphere: 276 x 219 mm, approx. 320pp, 133 line illustrations
Published: May 1981
Co-published by the British Museum (Natural History), London and Cambridge University
Press, Cambridge.
Titles to be published in Volume 46
The generic and tribal classification of spore-feeding Thysanoptera (Phlaeothripidae: Idolo-
thripinae).
By L. A. Mound & J. M. Palmer.
A revision of the Afrotropical mole-crickets (Orthoptera: Gryllotalpidae).
By B. C. Townsend.
Key to the genera of galerucine beetles of New Guinea, with a review of Sastra and related new
taxa (Chrysomelidae).
By Sharon L. Shute.
The Afrotropical dacetine ants (Formicidae).
By Barry Bolton.
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd., Dorchester
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