(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Bulletin of the British Museum (Natural History)"

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 
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 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of 
the series on either an Annual or Per Volume basis. Prices vary according to the contents of 
the individual parts. Orders and enquiries should be sent to: 



Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW75BD, 
England. 



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 (B 2 ) on tergite 
IX short and stout, whereas males and females of all Idolothripinae have these setae as long as 
the dorsal (B^) and lateral (B 3 ) 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 
B 2 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 


ii 




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( P Sirt)(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 
discal setae; tube width 0-68 times length (Figs 62, 71, 74, 79). 

P. citricauda. Stylets wide apart; ocellar setae small; antennal III with sense cones, VI fused to 
VII -I- VIII; meso- and metanota separate; pelta as in thomasi; sternite IV with 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 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 B 3 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 B 2 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; B 2 ?; B 3 220. 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 B v ?; B 2 ?; B 3 240. 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 B v 360, B 2 420. 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; B 2 120; B 3 160. 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/7otes(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 B 2 and 5 3 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, B l 600; B 2 550; B 3 550. 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 B l 500; B 2 450. 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; B 2 
200; B 3 200. 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; B 2 210: B 3 210. 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 B v 65; B 2 
145; B 3 228. Tergite IX setae, B^ 460; B 2 488. 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; B 2 
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). 
*aM>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). 
*scAoii(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 5 t 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 B l 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 B l 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, B 2 
180, 5 3 230. Tergite IX setae, B l 180, B 2 220, B 3 180. 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 B 2 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 B l on tergite VII thorn-like on a stout tubercle, but B 1 and B 2 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, B 2 90, B 3 
60. Tergite IX setae BI 50, B 2 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 B 2 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 B 2 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 B 2 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 B 2 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 B 2 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 B 2 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 B 2 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). 



References 

Ananthakrishnan, T. N. 19560. Erythrinothrips , nov. - A new subgenus of Agnostothrips Moulton 

(Tubulifera, Thysanoptera) from India. Indian Journal of Entomology 17 (1955): 341-342. 

19566. Studies on some Indian Thysanoptera III. Zoologischer Anzeiger 157: 130-139. 

1957. Studies on Indian Thysanoptera IV. Zoologischer Anzeiger 159: 92-102. 

1958. Two new species of Tubuliferous Thysanoptera from India (Thysanoptera, Phlaeothripidae). 

Proceedings of the Entomological Society of Washington 60: 277-280. 

1959. Studies on some Indian Thysanoptera V. Zoologischer Anzeiger 162: 313-324. 

1960. Neosmerinthothrips inquilinus sp. nov., an inquiline Tubuliferan (Thysanoptera) from India. 

Bulletin of Entomology, Loyola College, Madras 1: 32-33. 
1961a. New Tubuliferan Thysanoptera from India. Journal of the Zoological Society of India 12 

(1960): 250-258. 

19616. Studies on some Indian Thysanoptera VI. Zoologischer Anzeiger 167: 259-271. 

1964a. A contribution to our knowledge of the Tubulifera (Thysanoptera) from India. Opuscula 

EntomologicaSuppl. 25: 1-120. 

19646. Thysanopterologica Indica-I. Entomologisk TidskriftSS: 99-120. 

1966. Thysanopterologica Indica - IV. Bulletin of Entomology, Loyola College, Madras 7: 1-12. 

1967. New Indian Thysanoptera. Annales de la Societe Entomologique de France 3: 227-237. 

1968a. Studies on new and little known Indian Thysanoptera. Oriental Insects 1: 113-138. 

19686. Thysanopterologica Indica- V. Oriental Insects 2: 41-58. 

1968c. Two new species of Nesothrips with further remarks on Nesothrips robustus (Thysan. 

Tubulifera}. Annales de la Societe Entomologique de France 4: 967-973. 

1969a. Mycophagous Thysanoptera -I. Indian Forester 95: 173-185. 

19696. Mycophagous Thysanoptera -II. Oriental Insects 3: 289-299. 

1969c. The genus Polyphemothrips Schmutz from India. Oriental Insects 3: 301-310. 

I969d. Indian Thysanoptera. CSIR Zoological Monograph 1, 171 pp. New Delhi. 

1970. On some species of Nesothrips Kirkaldy with notes on the influence of oedymerism and 



98 L. A. MOUND AND J. M. PALMER 

gynaecoidism in species of allied genera of Cryptothripini (Tubulifera: Thysanoptera). Journal of the 
Zoological Society of India 22: 51-62. 

- 1971. Mycophagous Thysanoptera -III. Oriental Insects 5: 189-208. 

- 1972a. New species of Stigmothrips Ananthakrishnan with keys to Indian species. Journal of the 
Zoological Society of India 23: 175-185. 

- 19726. Mycophagous Thysanoptera -IV. Oriental Insects 6: 425^37. 

- 1972c. Mycophagous Thysanoptera - V. Oriental Insects 6: 439-447. 

1973a. Studies on some Indian species of the genus Elaphrothrips Buffa (Megathripinae: Tubulifera: 

Thysanoptera). Pacific Insects 15: 271-284. 
19736. Mycophagous Tubulifera of India (Thysanoptera: Insecta). Occasional Publications of the 

Entomological Research Unit, Loyola College, Madras 2: 144 pp. , 26 pis. 

1978. Thrips galls and gall thrips. Zoological Survey of India, Technical Monograph no. 1, 69 pp., 



26 pis. 
Ananthakrishnan, T. N. & Jagadish, A. 1970. The species of Diceratothrips Bagnall and allied genera from 

India (Thysanoptera: Megathripinae: Insecta). Oriental Insects 4: 265-280. 

Andre, F. 1940. The Nearctic species of Elaphrothrips Buffa (Thysanoptera: Phlaeothripidae). Proceed- 
ings of the Entomological Society of Washington 42: 75-90. 
Bagnall, R. S. 1908a. On some new and curious Thysanoptera (Tubulifera) from Papua. Annals and 

Magazine of Natural History (8) 1: 355-363. 
19086. On some new genera and species of Thysanoptera. Transactions of the Natural History Society 

of 'Northumberland 3: 183-217. 
1909a. Synonymical notes; with description of a new genus of Thysanoptera. Annales de la Societe 

Entomologique de Belgique 52: 348-352. 
19096. Notes on Thysanoptera (Tubulifera) new to the British fauna, with description of a new 

species oiMegathrips. Entomologist's Monthly Magazine 45: 129-132. 
1909c. On some new and little-known exotic Thysanoptera. Transactions of the Natural History 

Society of Northumberland 3: 524-540. 

I909d. On two new genera of Thysanoptera from Venezuela: Anactinothrips and Actinothrips. 



Journal of the Linnaean Society (Zoology) 30: 329-335. 

1910a. A contribution towards a knowledge of the neotropical Thysanoptera. Journal of the Linnaean 
Society (Zoology) 30: 369-387. 

19106. Thysanoptera. In: Fauna Hawaiiensis 3: 669-701. London. 

1911. Descriptions of three new Scandinavian Thysanoptera (Tubulifera). Entomologist's Monthly 
Magazine 46: 60-63. 

1912. Preliminary descriptions of three new species of Thysanoptera. Annals and Magazine of 
Natural History (8) 9: 214-217. 

1913. Brief descriptions of new Thysanoptera. - 1. Annals and Magazine of Natural History (8) 12: 
290-299. 

1914a. Brief descriptions of new Thysanoptera. - II. Annals and Magazine of Natural History (8) 13: 
22-31. 

19146. Brief descriptions of new Thysanoptera. -III. Annals and Magazine of Natural History (8) 13: 
287-297. 

1914c. Brief descriptions of new Thysanoptera. - IV. Annals and Magazine of Natural History (8) 14: 
375-381. 

1915a. Brief descriptions of new Thysanoptera. - V. Annals and Magazine of Natural History (8) 15: 
315-324. 

19156. Brief descriptions of new Thysanoptera. - VI. Annals and Magazine of Natural History (8) 15: 
588-597. 

1915c. A preliminary account of the Thysanoptera of Borneo. Sarawak Museum Journal 2: 267-272. 

1916. Brief descriptions of new Thysanoptera. -VIII. Annals and Magazine of Natural History (8) 17: 
397-^12. 

1917. On a collection of Thysanoptera from St Vincent, with descriptions of four new species. Journal 
of Zoological Research 2: 21-27. 

1918. Brief descriptions of new Thysanoptera. - IX. Annals and Magazine of Natural History (9) 1: 
201-221. 

19210. On Thysanoptera from the Seychelles Islands and Rodrigues. Annals and Magazine of Natural 
History (9) 7: 257-293. 

19216. Brief descriptions of new Thysanoptera. - XI. Annals and Magazine of Natural History (9) 7: 
355-368. 



GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 99 

1921c. Brief descriptions of newThysanoptera. -XII. Annals and Magazine of Natural History (9) 8- 
393-400. 

1923. Brief descriptions of newThysanoptera. - XIII. Annals and Magazine of Natural History (9) 12: 



624-631. 

1924. Brief descriptions of newThysanoptera. -XIV. Annals and Magazine of Natural History (9) 14: 
625-640. 

1926. Brief descriptions of newThysanoptera. -XVI. Annals and Magazine of Natural History (9) 18: 
545-560. 

1927. Contributions towards a knowledge of the European Thysanoptera. - II. Annals and Magazine 
of Natural History (9) 19: 564-575. 

1928. On some Samoan and Tongan Thysanoptera, with special reference to Ficus gall-causers and 



their inquilines. Thysanoptera. In: Insects of Samoa 7: 55-76, London. 

1929. On some new and interesting Thysanoptera of economic importance. Bulletin of Entomological 
Research 20: 69-76. 

1932. Brief descriptions of new Thysanoptera. - XVII. Annals and Magazine of Natural History (10) 
10: 505-520. 

1933a. More new and little-known British thrips. Entomologist's Monthly Magazine 69: 120-123. 

19336. Contributions towards a knowledge of the European Thysanoptera. - IV. Annals and 
Magazine of Natural History (10) 11: 647-661. 

1934a. Brief descriptions of new Thysanoptera. - XVIII. Annals and Magazine of Natural History 
(10) 13: 481-498. 

19346. Contributions towards a knowledge of the Thysanopterous genus Elaphrothrips Buffa. - I. 



Annals and Magazine of Natural History (10) 13: 627-636. 
1935. Contributions towards a knowledge of the Thysanopterous genus Elaphrothrips Buffa. - II. 

Annals and Magazine of Natural History (10) 15: 130-146. 
1936. Descriptions of some new Thysanoptera from tropical Africa and Madagascar. Revue Frangaise 

d'Entomologie 3: 219-230. 
Bergroth, E. 1888. Diagnose d'une nouvelle espece de Thysanopteres. Bulletin de la Societe Entomolo- 

gique de Belgique 1888: xxx. 
Bianchi, F. A. 1944. Nesothrips Kirkaldy supersedes Oedemothrips Bagnall. Proceedings of the Hawaiian 

Entomological Society 12: 31-38. 

1945. Introduction to the Thysanoptera of New Caledonia. Proceedings of the Hawaiian Entomo- 
logical Society 12: 249-278. 
1947. Thysanoptera Hawaiiensis - I. Proceedings of the Hawaiian Entomological Society 13: 

37-42. 
1949. New thrips records and species from the Marianas. Proceedings of the Hawaiian Entomological 

Society 13: 347-350. 

1953. Thysanoptera of Samoa. Proceedings of the Hawaiian Entomological Society 15: 93-108. 



Bournier, A. 1956. Trois especes nouvelles de Phloeothripidae (Thysanopt.). Bulletin de la Societe 

Entomologique de France 61: 160-174. 
1962. Thysanopteres de France, II. Descriptions sommaire de quelques especes nouvelles du 

Languedoc Mediterraneen. Bulletin de la Societe Entomologique de France 67: 41-43. 
1967. Thysanopteres de Madagascar. Annales de la Societe Entomologique de France (N.S.) 3: 

1015-1027. 
1968. Thysanopteres de 1' Angola. IV. Publicafdes Culturais de Companhia de Diamantes de Angola 

77: 135-164. 

1970. Thysanopteres du Gabon. Biologia Gabonica 6: 151-168. 

1971. Thysanopteres d'Afrique noire. Bulletin de 1'I.F.A.N. (A) 33: 145-167. 

1974. Thysanopteres de 1' Angola: VI. Publicagoes Culturais de Companhia de Diamantes de Angola 



88: 153-165. 
Buffa, P. 1908. Esame della raccolta di Tisanotteri Italiani esistente nel Museo Civico di Storia Naturale di 

Geneva. Redia4: 382-391. 

1909. 1 Tisanotteri esotici esistenti nel Museo Civico di Storia Naturale di Geneva. Redia 5: 157-172. 

Chen, L. S. 1980. Thrips associated with Mulberry plant (Morus sp.) in Taiwan. Proceedings of the 

National Science Council, Taiwan 4: 169-182. 
Cott, H. E. 1956. Systematics of the suborder Tubulifera (Thysanoptera) in California. University of 

California Publications in Entomology 13: 1-216. 
Crawford, D. L. 1910. Thysanoptera of Mexico and the South. II. Pomona College Journal of Entomology 

2: 153-170. 



100 L. A. MOUND AND J. M. PALMER 

Crawford, J. C. 1947. The North American species of the genus Megalothrips Uzel (Thysanoptera, 
Phlaeothripidae). Proceedings of the Entomological Society of Washington 49: 197-199. 

- 1948. A new Parallothrips from Cyprus. Proceedings of the Entomological Society of Washington 50: 
213-215. 

De Santis, L. 1943. Especies nuevas y conocidas de la Republica Argentina. Revista de la Facultad de 
Agronomia de La Plata 25: 89-96. 

- 1957. Dos nuevos Tubuliferos de la Republica Argentina (Thysanoptera: Tubulifera). Revista de la 
Uruguaya de Entomologia 2: 1-4. 

1958. Las especies Argentinas del genero Leptogastrothrips (Thysanoptera: Tubulifera). Revista de la 

Facultad de Agronomia 34: 95-102. 
1960. Un nuevo Tisanoptero gigante del Brasil. Actas y trabajos del Primer Congreso Sudamericano 

de Zoologia 3: 57-60. 
1963a. Adiciones a la Fauna Argentina de Tisanopteros III. Notas Comision de Investigacion 

Cientifica, Provincia de Buenos Aires 1: 1-14. 

19636. Tisanopteros de Tierra del Fuego. Revista de la Sociedad Entomologica Argentina 24: 63-66. 



Dyadechko, N. P. 1961. A new thrips species Embothrips tubversicolor sp. nov. (Thysanoptera). [In 
Russian.] Dopovidi Akademie Nauk Ukrayin'skoiRSR. Kiev 1961 (5): 688-690. 

1962. A new thrips species (Thysanoptera) from South Maritime Provinces - Cryptothrips maritimus 

Djadetschko sp. n. [In Russian.] Zoologicheskii Zhurnal 41: 764-765. 

1964. Thrips (Thysanoptera) of the European part of the U.S.S.R. 387 pp. Kiev. [In Russian.] 



Faure, J. C. 1925. A new genus and five new species of South African Thysanoptera. South African Journal 

of Natural History 5: 143-166. 
1933. New genera and species of Thysanoptera from South Africa. Bulletin of the Brooklyn 

Entomological Society 28: 1-20, 55-75. 
1942. Records and descriptions of South African Thysanoptera - III. Journal of the Entomological 

Society of Southern Africa 5: 74-83. 

1943. Two new species of Bolothrips (Thysanoptera) from South Africa. Journal of the Entomologi- 
cal Society of Southern Africa 6: 86-89. 
1945. Two new species of Allothrips (Thysanoptera) from South Africa. Proceedings of the Royal 

Entomological Society of London (B) 14: 150^154. 
1949a. Maxillata priesneri gen. et spec. nov. , a thrips (Thysanoptera) with exceedingly long maxillae. 

Annals and Magazine of Natural History (12) 2: 851-858. 
19496. Three remarkable new genera of Phlaeothripidae (Thysanoptera) from South Africa. 

Entomology Memoirs. Department of Agriculture, Union of South Africa 2: 204-217. 
1949c. A new species oiPygothrips Hood (Pygothripidae, Thysanoptera) from Zululand. Journal of 

the Entomological Society of Southern Africa 12: 118-122. 
19540. South African Thysanoptera - 1. Journal of the Entomological Society of Southern Africa 17: 

9-26. 
19546. South African Thysanoptera - II. Journal of the Entomological Society of Southern Africa 17: 

145-166. 
1955. South African Thysanoptera - 3. Journal of the Entomological Society of Southern Africa 18: 

13-41. 
1956. South African Thysanoptera - 5. Journal of Entomological Society of Southern Africa 19: 

313-341. 
Froggatt, W. W. 1904. Studies on Australian Thysanoptera: the genus Idolothrips, Haliday. Proceedings of 

the Linnean Society of New South Wales 1904: 54-57. 
Gaud, S. M. 1961. The Thysanoptera of Puerto Rico. Technical Papers of the Agricultural Experimental 

Station, Puerto Rico 32: 1-160. 
Gauld, I. D. & Mound, L. A. 1982. Homoplasy and the delineation of holophyletic genera in some insect 

groups. Systematic Entomology 7: 73-86. 

Girault, A. A. 1926. New pests from Australia, V. Published privately. 4 pp. Brisbane. 
- 19270. Some new wild animals from Queensland. Published privately. 3 pp. Brisbane. 

19276. New Australian animals so far overlooked by outsiders. Published privately. 2 pp. Brisbane. 

1927c. Thysanoptera nova Australiensis from Queensland. Published privately. 1 pp. Brisbane. 

I921d. Thysanoptera nova Australiensis, II. Published privately. 2pp. Brisbane. 

1927e. A discourse on wild animals. Published privately. 2 pp. Brisbane. 

19280. Aprodigeous discourse on wild animals. Published privately. 3 pp. Brisbane. 

19286. Some Insecta and a new All Highness. (Notes compiled in fear and sorrow.). Published 

privately. 4 pp. Brisbane. 



GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 101 

1928c. Notice of a curious professor and of native wasps and wood lice. Published privately. 4 pp. 



Brisbane. 

1929. New pests from Australia VI. Published privately. 1 pp. Brisbane. 
1930. New pests from Australia, VII. Published privately. 3 pp. Brisbane. 



Greenwood, P. H. 1980. Towards a phyletic classification of the 'genus' Haplochromis (Pisces, Cichlidae) 
and related taxa. Part II; The species from Lakes Victoria, Nabugabo, Edward, George and Kivu. 

Bulletin of the British Museum (Natural History) (Zoology) 39: 1-101. 
Haga, K. 1975. A revision of the genus Pyrgothrips Karny with keys to the world species (Thysanoptera, 

Tubulifera). Kontyu 43: 263-280. 
Haliday, A. H. 1852. Order III. Physapoda. In Walker, F. , List of the specimens of Homopterous insects in 

the collection of the British Museum Pt IV: 1094-1118. 
Hartwig, E. K. 1948. Six new species of South African Thysanoptera, with statistical analyses of 

measurements. Journal of the Entomological Society of Southern Africa 11: 83-126. 
1952. Taxonomic studies of South African Thysanoptera, including genitalia, statistics and a revision 

of Trybom's types. Entomology Memoirs. Department of Agriculture, Union of South Africa 2: 339^99. 
Heeger, E. 1852a. Beitrage zur Naturgeschichte der Physapoden (Blassenfusse). Sitzungsberichte der 

Kaiserlichen Akademie der Wissenschaften, Wien9: 123-141. 
18526. Beitrage zur Insecten-Fauna Osterreichs. Sitzungsberichte der Kaiserlichen Akademie der 

Wissenschaften, Wien 9: 473-490. 

Hennig, W. 1966. PhylogeneticSystematics. Translated by D. D. Davis & R. Zangerl, 263 pp. Urbana. 
Hesse, A. J. 1934. Some insects associated with the plant Gnidia (Arthrosolen) laxa Gilg. -Thysanoptera. 

Annals of the South African Museum 30: 434-440. 
Hinds, W. E. 1902. Contribution to a monograph of the insects of the order Thysanoptera inhabiting North 

America. Proceedings of the United States National Museum 26: 79-242, 11 pis. 
Hood, J. D. 1908a. New genera and species of Illinois Thysanoptera. Bulletin of the Illinois State 

Laboratory of Natural History 8: 361-379. 

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. 

Canadian Entomologist 44: 137-144. 
1912c. Descriptions of new North American Thysanoptera. Proceedings of the Entomological Society 

of Washington 14: 129-160. 
1914. Studies on tubuliferous Thysanoptera. Proceedings of the Biological Society of Washington 27: 

151-172. 
1915. A remarkable new thrips from Australia. Proceedings of the Biological Society of Washington 

28:49-51. 
1916. Oedaleothrips hookeri, a new genus and species of Thysanoptera. Bulletin of the Brooklyn 

Entomological Society 11: 64-65. 

1918a. A new Kleothrips (Thysanoptera) from North Queensland. Bulletin of the Brooklyn Entomo- 
logical Society 13: 77-79. 
19186. New genera and species of Australia Thysanoptera. Memoirs of the Queensland Museum 6: 

121-150. 

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 

Biological Society of Washington 32: 75-91. 

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, 

134-138. 
1925d. Four new Phloeothripidae from the United States (Thysanoptera). Canadian Entomologist 

57: 218-222. 
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 

Society 23: 147-150. 



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: 

497-505. 
1941. A century of new American Thysanoptera. II. Revista de Entomologia, Rio de Janeiro 12: 

139-243. 
1942. A century of new American Thysanoptera. III. Revista de Entomologia, Rio de Janeiro 12: 

547-678. 

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 

67: 277-286. 
1954c. Three new Thysanoptera from Trinidad and British Guiana. Proceedings of the Royal 

Entomological Society of London (B) 23: 205-212. 

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. 

1958. A new Zeugmatothrips from Brazil. Proceedings of the Biological Society of Washington 60: 

225-228. 
Hood, J. D. & Williams, C. B. 1915. New Thysanoptera from Florida and Louisiana. Journal of the New 

York Entomological Society, 23: 121-138. 

International Commission on Zoological Nomenclature. 1961. International Code of Zoological Nomencla- 
ture. 176 pp. London. 
Ishida, 1. 1932. Fauna of the Thysanoptera in Japan. Insecta Matsumurana 7: 1-16. 



GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 103 

Jacot-Guillarmod, C. F. 1937. Ten new species of Thysanoptera and a catalogue of the known South 

African forms. Publications of the University of Pretoria. (Series II, Natural Science) 3: 1-62. 
1939a. New species of Phlaeothripidae (Thysanoptera) from South Africa. Journal of the Entomolo- 
gical Society of Southern Africa 1: 47-77. 

- 19396. Phlaeothripidae (Thysanoptera) new to South Africa, with descriptions of new genera and 
species. Journal of the Entomological Society of Southern Africa 2: 36-62. 

1940. Studies on South African Thysanoptera - 1. Journal of the Entomological Society of Southern 



Africa 3: 131-138. 

1941. Studies on South African Thysanoptera - II. Journal of the Entomological Society of Southern 
Africa 4: 80-100. 

1942. Studies on South African Thysanoptera- HI. Journal of the Entomological Society of Southern 
Africa 5: 64-74. 

1978. Catalogue of the Thysanoptera of the World. Part 5. Annals of the Cape Provincial Museums 



(Natural History) 7: 1257-1556. 
Johansen, R. M. 1974. Siete nuevos Tisanopteros de Tabasco, Veracruz, y el Pedregal de San Angel, 

Mexico, D. F. (Thysanoptera: Terebrantia: Tubulifera). Revista de la Sociedad Mexicana de Historia 

Natural 35: 249-276. 
1975. Un nuevo Zeuglothrips Hood Panameno, y descripcion del macho de Z. guerreronis Johansen y 

Garcia Aldrete, 1973 (Thysanoptera: Phlaeothripidae). Revista de la Sociedad Mexicana de Historia 

Natural 36: 187-194. 
1976. Nuevos thrips tubuliferous (Insecta: Thysanoptera), de Mexico, I. Anales del Instituto de 

Biologia, Universidad de Mexico (Zoologia) 47: 57-68. 
1977a. Nuevos thrips tubuliferous (Insecta: Thysanoptera), de Mexico. III. Anales del Instituto de 

Biologia, Universidad de Mexico (Zoologia) 48: 37-50. 
19776. Nuevos thrips tubuliferous (Insecta: Thysanoptera), de Mexico. IV. Anales del Instituto de 

Biologia, Universidad de Mexico (Zoologia) 48: 51-69. 
1978a. Dos nuevas especies de Elaphrothrips Buffa, 1909 (Thysanoptera: Phlaeothripidae), del 

sureste de Mexico. Anales del Instituto de Biologia, Universidad de Mexico (Zoologia) 49: 87-94. 
19786. Seis nuevas especies de Elaphrothrips Buffa, 1909 (Thysanoptera: Phlaeothripidae) de Brasil 

y Peru. Anales del Instituto de Biologia, Universidad de Mexico (Zoologia) 49: 95-113. 
1978c. Notas sinonimicas acerca de Tisanopteros de Mexico. I. Anales del Instituto de Biologia, 

Universidad de Mexico (Zoologia) 49: 277-280. 

1979. Nuevos thrips tubuliferous (Insecta: Thysanoptera), de Mexico. VI. Anales del Instituto de 



Biologia, Universidad de Mexico (Zoologia) 50: 179-191. 

Johansen, R. M. & Garcia Aldrete, A. N. 1973. Una nueva especie Mexicana de Zeuglothrips Hood. 
(Thysanoptera: Phlaeothripidae). Revista de la Sociedad Mexicana de Historia Natural34: 55-61. 

1976. A new Mexican Dasythrips (Thysanoptera; Phlaeothripidae). Revista de Biologia Tropi- 
cal 24: 235-241. 

Karny, H. 1911. Neue Phloeothripiden - Genera. Zoologischer Anzeiger 38: 501-504. 

1912a. Ueber einige afrikanische Thysanopteren. Entomologische Zeitschrift, Fauna Exotica 2 (5): 

19-20, 22-26. 

19126. Gallenbewohnende Thysanopteren aus Java. Marcellia 11: 115-169. 

1912c. Einige weitere Tubuliferen aus dem tropischen Afrika. Entomologische Rundschau 29: 

130-133, 138-139, 150-151. 

1913a. Uber gallenbewohnende Thysanopteren. Verhandlungen der Zoologisch-Botanischen Gesell- 



schaft in Wien 63: 5-12. 

19136. Beitrag zur Thysanopteren-Fauna von Neu-Guinea und Neu-Britannien. Archiv fur Natur- 



geschichte 79(1): 123-136. 

1913c. H. Sauter's Formosa- Ausbeute. Supplementa Entomologica 2: 127-134. 

1913d. Thysanoptera von Japan. Archiv fur Naturgeschichte 79 (2): 122-128. 

1916. Beitrage zur Kenntnis der Gallen von Java. Zweite Mitteilung liber die javanischen Thysanop- 
terocecidien und deren Bewohner [in part]. Zeitschrift fur Wissenschaftliche Insektenbiologie 12: 84-94. 

1919. Synopsis der Megathripidae (Thysanoptera). Zeitschrift fur Wissenschaftliche Insektenbiologie 
14: 105-118. 

1920a. Die exotischen Tubuliferen (Thysanoptera) des Deutschen Entomologischen Museums 
(Berlin-Dahlem). Entomologische Mitteilungen 9: 88-94, 104-111, 186-191. 

19206. Some Thysanoptera from the Philippine Islands. Philippine Journal of Science 17: 203-211. 

1920c. Nova Australska Thysanoptera, jez nashbiral Mjoberg. Casopis Ceskoslovenske Spolecnosti 
Entomologicke 17: 35^4. 



104 L. A. MOUND AND J. M. PALMER 

1921fl. Zur Systematik der Orthopteroiden Insekten, & Beitraege zur Malayischen Thysanop- 

terenfaune. Treubia 1: 163-269, 277-291. 
19216. Ergaenzung zu Priesner's "//ap/of/mps-studien" die Australischen Haplothripinen. Treubia 

2: 21-36. 
1921c. Beitraege zur Malayischen Thysanopterenfauna. Treubia 2: 37-83. 

- 1923. Beitrage zur Malayischen Thysanopterenfauna. Treubia 3: 277-380. 

- 1924. Results of Dr E. Mjoberg's Swedish Scientific Expeditions to Australia 1910-1913. 38. 
Thysanoptera. Arkivfiir Zoologi 17A (2): 1-56. 

1925a. Die an Tabak auf Java und Sumatra Angetroffenen Blasenfusser. Bulletin van het Deli 

Proefstation, Medan, Sumatra 23: 1-55. 

19256. On some tropical Thysanoptera. Bulletin of Entomological Research 16: 125-142. 

1925c. Uber Phloeothrips sanguinolentus Bergroth nebst einer Revision der Diceratothripinen- 



Genera. Notulae Entomologicae 5: 77-84. 

Kelly, R. & Mayne, R. J. B. 1934. The Australian Thrips, a monograph of the Order Thysanoptera in 
Australia. 81 pp. Sydney. 

Kirkaldy, G. W. 1907. On two Hawaiian Thysanoptera. Proceedings of the Hawaiian Entomological 
Society 1: 102-103. 

Knechtel, W. K. 1936. Zur Kenntnis der Thysanopterenfauna Rumaniens. Bulletin de la Section Scienti- 
fique de I' Academic Roumaine 17: 159-162. 

Kudo, 1. 1975. New name for the subgenus Telothrips Kudo et Ananthakrishnan, not Priesner (Thysanop- 
tera, Phlaeothripidae). Kontyu43: 421. 

Kudo, I. & Ananthakrishnan, T. N. 1974. A new subgenus and species of Meiothrips Priesner (Thysanop- 
tera, Megathripinae) from Nepal. Kontyu 42: 385-387. 

Kurosawa, M. 1932. Descriptions of three new thrips from Japan. Kontyu 5: 230-242. 

1968. Thysanoptera of Japan. [In Japanese, English summary.] Insecta Matsumurana Suppl. 4: 1-94. 

Lieberman, J. & Gemignani, E. V. 1931. Un nuevo genero y dos nuevas especies de Thysanopteros 
argentinos. Revista de la Sociedad Entomologica, Argentina 3: 211-216. 

Maltbaek, J. 1928. Tillaeg til Thysanoptera Danica, Danske Frynsevinger. Entomologiske Meddelelser 16: 
369-381. 

Maynard Smith, J. 1958. The theory of evolution. 344pp. Harmondsworth, Middlesex. 

Morgan, A. C. 1925. A new genus, a new subgenus and seven new species of Thysanoptera from Porto 
Rico. Florida Entomologist 9: 1-9. 

Morison, G. D. 1958. Thysanoptera from South- West Arabia and Ethiopia. Journal of the Linnean Society 
of London (Zoology) 43: 587-598. 

Moulton, D. 1907. A contribution to our knowledge of the Thysanoptera of California. Miscellaneous 
Papers, Technical Series of the Bureau of Entomology, United States Department of Agriculture 12: 
39-68. 

1928a. Thysanoptera of the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society 7: 

105-134. 

19286. Thysanoptera from Abyssinia. Annals and Magazine of Natural History (10) 2: 227-248. 

1928c. New Thysanoptera from Formosa. Transactions of the Natural History Society of Formosa 18: 

287-328. 

1928d. The Thysanoptera of Japan: new species, notes, and a list of all known Japanese species. 

Annotationes Zoologicae Japonenses 2: 287-337. 

1928e. New Thysanoptera from India. Indian Forest Records 13: 285-292. 

1929a. New California Thysanoptera. Pan-Pacific Entomologist 4: 125-136. 

19296. New Mexican Thysanoptera. Pan- Pacific Entomologist 6: 11-20. 

1929c. Contribution to our knowledge of American Thysanoptera. Bulletin of the Brooklyn Entomo- 
logical Society 24: 224-244. 

1930. Thysanoptera from South Africa. Annals and Magazine of Natural History (10) 5: 414-416. 

19330. The Thysanoptera of South America (IV). Revista de Entomologia, Rio de Janeiro 3: 385-419. 

19336. New Thysanoptera from India. Indian Forest Records 19: 1-6. 

1934. New Thysanoptera of the Hawaiian Islands. Proceedings of the Hawaiian Entomological 

Society 8: 499-503. 

1935. New species of thrips from South- Western Australia. Journal of the Royal Society of Western 

Australia 21: 97-100. 

1938. Thysanoptera from Minas Geraes, Brazil. Revista de Entomologia, Rio de Janeiro 9: 374-383. 

1939. Thysanoptera collected by the Mangarevan Expedition. Occasional Papers of the Bernice P. 



Bishop Museum, Hawaii 15: 141-148. 



GENERIC AND TRIBAL CLASSIFICATION OF SPORE-FEEDING THYSANOPTERA 105 

1941. Thysanoptera from Minas Gerais, Brazil. Revista de Entomologia, Rio de Janeiro 12: 314-323. 

19420. Seven new genera of Thysanoptera from Australia and New Zealand. Bulletin of the Southern 
California Academy of Sciences 41: 1-13. 

19426. Insects of Guam-I, Thysanoptera. Bulletin of the Bernice P. Bishop Museum, Hawaii 172: 
7-16. 

1944. Thysanoptera of Fiji. Occasional Papers of the Bernice P. Bishop Museum, Hawaii 17: 267-31 1. 

1947a. Thysanoptera from New Guinea, the Philippine Islands and the Malay Peninsula. Pan-Pacific 
Entomologist 23: 172-180. 

19476. New Thysanoptera from Mexico. Anales de la Escuila National de Ciencias Biologicas 4: 
419-421. 

1949. New Thysanoptera from Africa. Annals and Magazine of Natural History (12) 2: 481-499. 
1968. New Thysanoptera from Australia. Proceedings of the California Academy of Sciences (4) 36: 



93-124. 
Moulton, D. & Andre, F. 1936. Four new Thysanoptera, with a preliminary list of the species occurring in 

Iowa. Iowa State College Journal of Science 10: 223-234. 
Moulton, D. & Steinweden, J. B. 1932. New Marquesan Thysanoptera. Bulletin of the Bernice P. Bishop 

Museum, Hawaii98: 165-168. 
1933. Thysanoptera from the Society Islands. Bulletin of the Bernice P. Bishop Museum, Hawaii 

113: 29-33. 

1935. Two new species of Cryptothrips (Thysanoptera) from the Maquesas. Bulletin of the 



Bernice P. Bishop Museum, Hawaii 114: 163-165. 
Mound, L. A. 1968. A review of R. S. Bagnall's Thysanoptera Collections. Bulletin of the British Museum 

(Natural History) (Entomology) Suppl. 11: 1-181. 
1969. Revision of three Australian genera of Phlaeothripidae (Thysanoptera) with seven new species, 

and one new generic synonym. Journal of the Australian Entomological Society 8: 173-186. 
1970. Thysanoptera from the Solomon Islands. Bulletin of the British Museum (Natural History) 

(Entomology) 24: 83-126. 
1971a. Gall-forming thrips and allied species (Thysanoptera: Phlaeothripinae) from Acacia trees in 

Australia. Bulletin of the British Museum (Natural History) (Entomology) 25: 387^66. 
19716. A review of the Melanesian genus Mecynothrips (Thysanoptera). Journal of Natural History 

5: 279-283. 

1972a. Polytypic species of spore-feeding Thysanoptera in the genus Allothrips Hood (Phlaeothripi- 
dae). Journal of the Australian Entomological Society 11: 23-36. 
19726. Species complexes and the generic classification of leaf-litter thrips of the tribe Urothripini 

(Phlaeothripidae). Australian Journal of Zoology 20: 83-103. 

1974a. Spore-feeding thrips (Phlaeothripidae) from leaf litter and dead wood in Australia. Australian 



Journal of Zoology (Suppl. Series) 27: 1-106. 

19746. The Nesothrips complex of spore-feeding Thysanoptera (Phlaeothripidae: Idolothripinae). 
Bulletin of the British Museum (Natural History) (Entomology) 31: 107-188. 

1974c. Andrethrips floydi - a remarkable new Thysanopteron. Journal of Entomology (B) 43: 
109-113. 

1977. Species diversity and the systematics of some New World leaf litter Thysanoptera (Phlaeothri- 



pinae: Glyptothripini). Systematic Entomology 2: 225-244. 
Mound, L. A., Heming, B. S. & Palmer, J. M. 1980. Phylogenetic relationships between the families of 

recent Thysanoptera (Insecta). Zoological Journal of the Linnean Society 69: 111-141. 
Mound, L. A., Morison, G. D., Pitkin, B. R. & Palmer, J. M. 1976. Thysanoptera. Handbooks for the 

Identification of British Insects 1 (11): 1-79. 
Mound, L. A. & Palmer, J. M. 1981. Phylogenetic relationships between some genera of Thripidae 

(Thysanoptera). Entomologica Scan dinavica Suppl. 15: 153-170. 
Mound, L. A. & Walker, A. K. 1982. Thysanoptera: Terebrantia. In: Fauna of New Zealand, D.S.I.R., 

Wellington 1: 1-113. 
1983. Thysanoptera: Tubulifera. In: Fauna of New Zealand, D.S.I.R., Wellington, [in 

prep.] 
Newman, E. 1855. Characters of two undescribed species of Thrips, Lin. Transactions of the Entomological 

Society of London 3: 264-267. 
Okajima, S. 1975. Notes on the Thysanoptera from the Ryukyu Islands. I. Descriptions of two new species. 

Kontyu 43: 13-19. 
1976. Notes on the Thysanoptera from the Ryukyu Islands. II. On the genus Stigmothrips Ananthak- 

rishnan. Kontyu 44: 119-129. 



106 L. A. MOUND AND J. M. PALMER 

- 1979a. A revisional study of the genus Apelaunothrips (Thysanoptera: Phlaeothripidae). Systematic 
Entomology 4: 39-64. 

- 19796. Notes on the Thysanoptera from Southeast Asia VI. A new species of the genus Mecynothrips 
from Taiwan (Phlaeothripidae). Transactions of the Shikoku Entomological Society 14: 127-130. 

- 1979c. Two new species of the genus Gastrothrips Hood (Thysanoptera: Phlaeothripidae) from Japan 
and Taiwan. Kontyu47: 511-516. 

1981. A revision of the tribe Plectrothripini of fungus-feeding Thysanoptera (Phlaeothripidae: 



Phlaeothripinae). Systematic Entomology 6: 219-336. 

Palmer, J. M. & Mound, L. A. 1978. Nine genera of fungus-feeding Phlaeothripidae (Thysanoptera) from 
the Oriental Region. Bulletin of the British Museum (Natural History) (Entomology) 37: 153-215. 

Pelikan, J. 1961. Two new species of Oedaleothrips (Thysanoptera) from Asia. Casopis Ceskoslovenske 
Spolecnosti Entomologicke 58: 302-309. 

1965. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. 29. Thysanop- 
tera. Annales Historico-Naturales Musei Nationalis Hungarici 57: 229-239. 

1970. Thysanopteren aus Nepal (Ins., Thysanoptera). Khumbu Himal3: 361-369. 



Pergande, T. 1896. Description of a new species of Idolothrips. Entomological News 7: 63-64. 

Pitkin, B. R. 1978. Lectotype designations of certain species of thrips described by J. D. Hood and notes on 

his collection (Thysanoptera). Proceedings of the Entomological Society of Washington 80: 264-295. 
Pitkin, B. R. & Mound, L. A. 1973. A catalogue of West African Thysanoptera. Bulletin de 1'I.F.A.N. (A) 

35: 407-449. 

Post, R. L. 1961. Five new Oregon Thysanoptera. Pan-Pacific Entomologist 37: 137-143. 
Priesner, H. 1919. Zur Thysanopteren-Fauna der ostadriatischen Kiistenlander. Zeitschrift des Oster- 

reichischen Entomologen-Vereines 4: 104-106. 
1920. Kurze Beschreibungen neuer Thysanopteren aus Osterreich. Sitzungberichte derAkademie der 

Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Wien 129: 71-88. 
1921. Neue und wenig bekannte Thysanopteren der neotropischen Fauna aus der Sammlung des 

Berliner Zoologischen Museums. Deutsche Entomologishe Zeitschrift 1921: 187-223. 

1922. Ueber albanische Thysanopteren. Wiener Entomologische Zeitung 39: 105-107. 

1924. Bernstein-Thysanopteren. Entomologische Mitteilungen 13: 130-151. 

1925a. Zwei neue, beachtenswerte Thysanopteren-typen aus Ungarn. Zeitschrift des Oesterreichis- 

chen Entomologen-Vereines 10: 5-7. 

19256. Neue Thysanopteren. Deutsche Entomologische Zeitschrift 1925: 13-28. 

1925c. Thysanopterologica I. Zoologische Jarbiicher, Jena 50: 305-319. 

I925d. Katalog der europaischen Thysanopteren. Konowia4: 141-159. 

- 1926a. Die Jugendstadien der malayischen Thysanopteren. TreubiaS(Suppl.): 1-264. 

19266. Un genero nuevo y curioso del orden physopodos o Thysanopteros (insectos) de Mexico. 

Memorias y Revista de la Sociedad Cientifica 'Antonio Alzate' 44: 485-489. 

1927a. Die Thysanopteren Europas pp. 343-568. Wien. 

19276. Ein neuer Zeugmatothrips (Thysanoptera: Tubulifera) aus Costarica. Bollettino del Labor- 

atorio di Zoologia generate e agraria delta R. Scuola superiore di Agricoltura di Portici 20: 189-190. 

- 1927c. Neue und wenig bekannte Thysanopteren, gesammelt in Westafrika von Prof. Dr. F. Silvestri. 
Bollettino del Laboratorio di Zoologia generate e agraria dell R. Scuola superiore di Agricoltura di Portici 
21: 61-83. 

1928a. Die Thysanopteren Europas pp. 569-755. Wien. 

19286. Uber australische Thysanopteren. Sitzungberichte derAkademie der Wissenschaften. Mathe- 

matisch-Naturwissenschaftliche Klasse, Wien 137: 643-659. 

1928c. Thysanopterologica III. Zoologische Jarbiicher, Jena 56: 43-66. 

I929a. Bernstein-Thysanopteren II. Bernstein-Forschungen 1: 111-138. 

19296. Spolia Mentawiensia: Thysanoptera. Treubia 11: 187-210. 

1932. Thysanopteren aus dem Belgischen Congo. Revue de Zoologie et de Botanique Africaines 22: 

321-344. 

1933a. Neue Thysanopteren aus Mexiko, gesammelt von Prof. Dr. A. Dampf. Wiener entomolo- 



gische Zeitung 50: 49-63. 

- 19336. Indomalayische Thysanopteren IV. Konowia 12: 69-85. 

- 1933c. Neue exotische Thysanopteren. Stylops2: 145-156. 

1934. Drei neue Phloeothripiden (Thysanoptera). Stylops3: 58-63. 

- 1935a. Indomalayische Thysanopteren VI. Konowia 14: 159-174, 241-253, 323-339. 

19356. Neue exotische Thysanopteren. Stylops4: 125-131. 

- 1935c. New or little known Oriental Thysanoptera. Philippine Journal of Science 57: 351-375. 



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- 



gique d'Egypte 20: 83-104 

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 
Entomology 13: 183-200. 

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 

History Society 30: 788-792. 
1928. A contribution to our knowledge of the Thysanoptera of India. Memoirs of the Department of 

Agriculture in India (Entomological series) 10: 217-316. 

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: 

255-258. 
Reuter, O. M. 1879. Diagnoser ofver nya Thysanoptera fran Finland. Ofversigt af Finska Vetenskaps- 

Societetens forhandlingar 1878-1879: 207-223. 
1885. Thysanoptera Fennica, descripsit. I. Tubulifera. Bidrag till Kdnnedom af Finlands Natur och 

Folk 40: 1-26. 
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 

Insects 13: 391^03. 
1979. Notes on some Cryptothripine species from Polynesia, with description of a new species 

(Phlaeothripidae: Thysanoptera). Pacific Insects 20: 313-317. 
Sakimura, K. & Bianchi, F. A. 1977. A review of the Hawaiian species of Idolothripinae (Phlaeothripidae: 

Thysanoptera). Proceedings of the Hawaiian Entomological Society 22: 495-523. 
Schmutz, K. 1909. Zur Kenntnis einiger neuen Thysanopterengenera (Tubulifera). Annalen des Natur- 

historischen Museums, Wien 23: 273-281. 342-347. 
1913. Zur Kenntnis der Thysanopterenfauna von Ceylon. Sitzungsberichte der Akademie der 

Wissenschaften in Wien 122: 991-1089. 
Seshadri, A. R. & Ananthakrishnan, T. N. 1954. Some new Indian Thysanoptera I. Indian Journal of 

Entomology 16: 210-256. 

Solowinow, P. 1924. Neue Arten Blasenfiisse. Entomologische Zeitschrift 38: 25-26. 
Stannard, L. J. 1953. Illinothrips rossi, new genus and species (Thysanoptera; Phlaeothripidae). Transac- 
tions of the Illinois Academy of Science 46: 193-196. 

1954a. Tropothrips in North America (Thysanoptera; Phlaeothripidae). Proceedings of the Biologi- 
cal Society of Washington 67: 81-84. 



108 L. A. MOUND AND J. M. PALMER 

- 1954ft. Actinothrips (Hybridothrips) oneillae, new subgenus and species. Proceedings of the Entomo- 
logical Society of Washington 56: 71-74. 

- 1955. The species and subspecies of North American Allothrips (Thysanoptera: Phlaeothripidae). 
Annals of the Entomological Society of America 48: 151-157. 

- 1956. Six new species of Adelothrips from the New World with critical remarks on this genus and 
related genera (Thysanoptera: Tubulifera). Proceedings of the Biological Society of Washington 69: 
105-114. 

- 1957. The phylogeny and classification of the North American genera of the suborder Tubulifera 
(Thysanoptera). Illinois Biological Monographs 25: 1-200. 

1974. Atractothrips mockfordi, a new species from Mexico (Thysanoptera: Phlaeothripidae). Pro- 



ceedings of the Entomological Society of Washington 76: 45-48. 

1976. A synopsis of some ant-mimicking thrips, with special reference to the American fauna 



(Thysanoptera: Phlaeothripidae: Idolothripinae). Journal of the Kansas Entomological Society 49: 

492-508. 
Stys, P. 1967. Monograph of Malcinae, with reconsideration of morphology and phylogeny of related 

groups (Heteroptera, Malcidae). Acta Entomologica Musei Nationalis, Prague 37: 351-516. 
Takahashi, R. 1935. An interesting thrips from Amami-Oshima, Loochoos. Mushi 8: 61-63. 
1937. Descriptions of new Thysanoptera from Formosa, with notes on the species found on the high 

elevations of the island. Tenthredo 1: 339-350. 
Targioni-Tozzetti, A. 1881. Relazione intoro ai lavori della R. Stazione di entomologia agraria di Firenze 

per gli anni 1877-78. Art. V. Fisapodi (Thrips). Annali di Agricoltura 34: 120-134. 
Thomasson, G. L. & Post, R. L. 1966. 1. North Dakota Tubulifera (Thysanoptera). North Dakota Insects 6: 

1-56. 
Titschack, E. 1964. Zur Variation der Borstenlange von Cryptothrips nigripes O. M. Reuter 1880 

(Thysanoptera). Verhandlungun des Vereins fur Naturwissenschaftliche Heimatforschung 36: 45-51. 

- 1965. Cryptothrips nigripes O. M. Reuter, subspec. phariacus nom. nov. Verhandlungen des Vereins 
fur Naturwissenschaftliche Heimatforschung 36: 147. 

Trybom, F. 1908. Physapoda, In Sjostedt, Wissenschaftliche Ergebnisse der schwedischen Zoologischen 
Expedition nach der Kilimanjaro und Meru, 1905-1906 16: 1-23. 

- 1910. Zwei neue Physapoden aus Madagaskar. Voeltzkow's Reise in Ostafrika in den Jahren 
1903-1905, Wissenschaftliche Ergebnisse 2: 521-525. 

1912. Physapoden aus Natal und dem Zululande. Arkivfur Zoologi 7(33): 1-52. 



Uzel, H. 1895. Monographic der Ordnung Thysanoptera. 472 pp. Koniggratz. 

Vuillet, A. 1914. Description d'un Dicaiothrips nouveau de 1'Inde (Thysanopt. Idolothripidae). Bulletin de 

la Societe Entomologique de France 1914: 276-278. 
Walker, F. 1859. Characters of some apparently undescribed Ceylon insects. Annals and Magazine of 

Natural History (3) 4: 217-224. 
Watson, J. R. 1919. New Thysanoptera from Florida-IV. Florida Buggist 2: 97-102. 

- 1920. New Thysanoptera from Florida- VII. Florida Entomologist 4: 18-23, 27-30. 

- 1921. New Thysanoptera from New York. Bulletin of the Brooklyn Entomological Society 16: 78-86. 

- 1923. Synopsis and catalog of the Thysanoptera of North America. Technical Bulletin of the 
Agricultural Experimental Station, University of Florida 168: 1-98. 

- 1925. A new species of Symphyothrips (Thysanoptera) from Argentina. Florida Entomologist 9: 
29-30, 45. 

- 1931. A collection of Thysanoptera from Western Oklahoma. Publications of the University of 
Oklahoma Biological Survey 3: 339-345. 

1933. Two new species of Oedaleothrips with notes on other species. Florida Entomologist 17: 48-50; 

63-64. 

1934/35/37. Thysanoptera of the Geenton. Florida Entomologist 18: 44-46, 55-62; 20: 12-15, 17-21. 



Woo, K. S. 1974. Thysanoptera of Korea. Korean Journal of Entomology 4: 1-90. 

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- 
tera) des siidlichen Marokko. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 515: 
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 









^ 




! ' / 


* 




\y 


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 



-QVV 




O 



o 



OOf 



08 



09C 





ore I 



O 



o o o o o 
o 

o 



* 



8 8 

C C 



o -o 5 

C C N 

is S 5 



o 



* 



OOE 



083 



092 



I 



CL, 

.2 



o 

Cu 

03 
Ofi 



. 

si 






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 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of 
the series on either an Annual or Per Volume basis. Prices vary according to the contents of 
the individual parts. Orders and enquiries should be sent to: 



Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW75BD, 
England. 

. 




" 






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 ^ MM* 

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 R s 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 R s 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 R s . 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, R l and R s 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 R s 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 fl s . 

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 ' CCCc cc cccC 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 ccccccccc CCff ^ 

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 R s , 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 fl s 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 R s 
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 R s 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 R s 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 R s 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 R s 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. 

References 

Alexander, R. D. 1962. The role of behavioral study in cricket classification. Systematic Zoology 11: 53-72. 
Bennet-Clark, H. C. 1970. A new French mole cricket, differing in song and morphology from Gryllotalpa 

gryllotalpa L. (Orthoptera: Gryllotalpidae). Proceedings of the Royal Entomological Society of London 

(B) 39: 125-132. 
Broughton, W. B. 1964. Method in bio-acoustic terminology. 7nBusnel,R. G. (ed.), Acoustic behaviour of 

animals, xx+933 pp. Amsterdam, London, New York. 

1976. Proposal for a new term 'echeme' to replace 'chirp' in animal acoustics. Physiological 

Entomology 1: 103-106. 

Bruner, L. 1915. Preliminary catalogue of the Orthopteroid insects of the Philippine Islands. The 

University Studies of the University of Nebraska 15: 195-281. 
Brunn, M. von 1901. Ostafrikanisehe Orthopteren gesammelt von Herrn Dr. F. Stuhlmann 1888 und 1889. 

Mitteilungen aus dem Naturhistorischen Museum in Hamburg 18: 213-283. 
Burmeister, H. 1834-1839. Handbiich der Entomologie 2. 1050 pp. Berlin. 
Chopard, L. 1934. Catalogues raisonnes de la faune entomologique du Congo Beige. Orthopteres- 

Gryllides. Annales du Musee du Congo Beige, Zoologie (3) 4: 1-88. 
- 1939. Gryllidae. Exploration du Pare National Albert. Mission G. F. de Wine (1933-1935) 27: 5-15. 

1948. Contribution a 1'etude des Gryllides du Congo Beige. Revue de Zoologie et de Botanique 

Africaines 41: 109-121. 

1955. Orthoptera Ensifera et Tridactyloidea. South African animal life. Results of the Lund University 

expedition in 1950-1951 2: 266-301. 

1967. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descarpentries. L. 



Orthopteres Gryllodea. Bulletin de I'lnstitut Fondamental d'Afrique Noire 29(A): 758-776. 



202 B. C. TOWNSEND 

1968. Gryllides. Fam. Gryllidae: subfam. Mogoplistinae, Myrmecophilinae, Scleropterinae, Cacho- 
plistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae. Fam. Oecan- 
thidae, Gryllotalpidae. Orthopterorum Catalogus 12. pp. 215-500. s'Gravenhage. 

1969. Orthoptera, vol. 2, Grylloidea. The fauna of India and the adjacent countries, xviii+421 pp. 



Calcutta. 
Fieber, F. X. 1851. Geschichte der Gesellschaft 1848, 1849, 1850. 10. Section fur Naturwissenschaften und 

angewandte Mathematik am 23 November 1848. Abhandlungen der Koniglichen Bohmischen Gesell- 
schaft der Wissenschaften (5) 6: 15-18. 
Gerstaecker, A. 1869. Beitrag zur Insekten-fauna von Zanzibar. 11. Orthoptera et Neuroptera. Archivfur 

Naturgeschichte 35: 202-223. 
Karny, H. 1907. Die Orthopterenfauna des Kustengebietes von Osterreich-Ungarn. Berliner Entomolo- 

gische Zeitschrift 52: 17-52. 
Kirby, W. F. 1906. A synonymic catalogue of Orthoptera 2. Orthoptera Saltatoria 1. Achetidae et 

Phasgonuridae. vii+228 pp. London. 
Latreille, P. A. 1802. Histoire naturelle, generale et particuliere, des Crustacees et des Insectes 3. 386 pp, 

37 pis. Paris. 

Leach, W. 1815. Entomology. In Brewster, D. (ed.), The Edinburgh Encyclopedia 9. 766 pp. Edinburgh. 
Lopez Seoane, V. 1878. Ortopteros de la peninsula Hispano-Lusitanica. Stettiner Entomologische Zeitung 

39: 366-376. 
Matheny, E. L. 1981. Contrasting feeding habits of pest mole cricket species. Journal of Economic 

Entomology 74: 444-445. 
Mjoberg, E. 1913. Preliminary descriptions of some new, Australian Gryllids and Forficulids. Entomolo- 

gisk Tidskrift 34: 26-34. 

Oken, L. 1815. Lehrbuch der Naturgeschichte 3. Zoologie 1: xxviii+842+xviii+ivpp., 40 pis. Jena. 
Orian, A. J. E. 1957. Saltatoria, Phasmida and Dictyoptera of Mauritius. Annals and Magazine of Natural 

History (12) 10: 513-520. 
Palisot de Beauvois, A. M. F. J. 1805-1821. Insectes recueillis en Afrique eten Amerique dans les royaumes 

d'Oware et de Benin, a Saint- Domingue et dans les Etats-Unis, pendant les annees 1786-1797. 276 pp. , 38 

pis. Paris. 

Ragge, D. R. 1955. The wing-venation of the Orthoptera Saltatoria with notes on Dictyopteran wing- 
venation. vi+159pp. London. 

- 1969. A revision of the African species of Pseudorhynchus Serville (Orthoptera: Tettigoniidae). 
Bulletin of the British Museum (Natural History) (Entomology) 23: 167-190. 

Redtenbacher, J. 1900. Die Dermatopteren und Orthopteren (Ohrwurmer und Geradflugler) von Oster- 
reich-Ungarn und Deutschland. 148 pp. Wien. 

Rochebrune, A. T. de 1884. Diagnoses d'Arthropodes nouveaux propres a la Senegambia (premier 
supplement). Bulletin de laSocietephilomathique de Paris (7) 8: 28-31. 

Saussure, H. de 1870-1879. Etudes sur les insectes Orthopteres. Mission scientifique au Mexique et dans 
I' Amerique centrale. Recherches zoologiques 6 (1). 533 pp., 8 pis. Paris. 

- 1876-1877. Melanges Orthopterologiques 5. 3 Gryllides. Memoires de la Societe de Physique et 
d' Histoire Naturelle de Geneve 25 (1): i-xxii, 1-352. 

1894. Fam. Gryllidae. Biologia Centrali Americana. Insecta. Orthoptera 1: 198-284. 



Saussure, H. de & Zehntner, L. 1894. Notice morphologique sur les Gryllotalpiens. Revue suisse de 

Zoologie 2: 403-430. 
Scudder, S. H. 1869. Revision of the large, stylated, fossorial crickets. Memoirs of the Peabody Academy of 

Science 1: 1-28. 
Sismondo, E. 1979. Stridulation and tegminal resonance in the tree cricket Oecanthus nigricornis 

(Orthoptera: Gryllidae: Oecanthinae.) Journal of Comparative Physiology 129: 269-279. 
Smithie, F. B. 1975. Naturalist's Color Guide. 23 pp. New York. 
Tindale, N. B. 1928. Australasian mole-crickets of the family Gryllotalpidae (Orthoptera). Records of the 

South Australian Museum 4: 1-42. 
Vayssiere, P. & Mimeur, J. 1926. Les insectes nuisibles au cotonnier en Afrique occidentale Frangaise. 

175 pp., 20 pis. Paris. 



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, 
and an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of 
the series on either an Annual or Per Volume basis. Prices vary according to the contents of 
the individual parts. Orders and enquiries should be sent to: 



Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW75BD, 
England. 



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, 44'S, 
14115'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 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, 
and an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of 
the series on either an Annual or Per Volume basis. Prices vary according to the contents of 
the individual parts. Orders and enquiries should be sent to: 



Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW75BD, 
England. 



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 u c ( 

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- 140- 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 5458 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, 722'S, 2050'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.