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DEEL 117 MUS. COMP. ZOOL 1974 i LIBRARY
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HARVARD UNIVERSITY
TIJDSCHRIFT VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING
| Tijdschrift voor Entomologie, deel 117, 1974 |
Afleveringen 1-2 verschenen 4.VI.1974 Afleveringen 3-4 verschenen 23.VIII.1974 Afleveringen 5-6 verschenen 15.X.1974 Aflevering 7 verscheen, 31.X11.1974
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INHOUD VAN DEEL 117
CHVALA, M. J. Doskoür, J. H. Mook & V. Pokorny. — The genus Lipara Meigen (Diptera, Bu systematics, morphology, behaviour, and CODE ss nt. . AAN A IO en A 1
Doskotit, J., zie CHVALA.
Hure Ris LAMBERS, D. — On American Aphids, with descriptions of a new genus and some new species (Homoptera, Aphididae). . . . . . . . . . 103
Jong, R. DE. — Systematics and evolution of the Palaearctic Spzalia species (Lepi- hot, HER Ne EE 0 0. 225
LiEFTINCK, M. A. — Review of Central and East Asiatic Habropoda F. Smith with Habrophorula, a new genus from China (Hymenoptera, Anthophoridae) 157
LitH, J. P. VAN. — Revision of the Psenini of the Ethiopian Region, including Malagasy (Hymenoptera, Sphecidae, Pemphredoninae) . . . . . . . . 39
Mook, J. H., zie Chvála.
MUNROE, E. — Pyralidae of the Third Archbold ee IE Ma Odon- Bimaenkenidoptera) 2 e... li : PA 27
Pokorny, V., zie Chvala. RossEM, G. VAN. — The Gravenhorst, Schigdte and Foerster types belonging to the genus Megastylus Schigdte, 1838, with keys to the Be a
Ichneumonidae, Microleptinae) . . . . . . : LT
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TIJDSCHRIFT Sr VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING
INHOUD
x M. CHVALA, J. Doskoär, J. H. Mook & V Pokorny. — The genus Lipara Meigen a | (Diptera, Chloropidae), systematics, ae behaviour, and ecology, pei 1-25, EE 1—22, Pl. 1—7.
| Tijdschrift voor Entomologie, deel 117, afl. 1
. Gepubliceerd 4-VI-1974
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THE GENUS LIPARA MEIGEN (DIPTERA, CHLOROPIDAE), SYSTEMATICS, MORPHOLOGY, BEHAVIOUR, AND ECOLOGY
by
M. CHVÁLA *), J. DOSKOCIL'), J. H. MOOK?) & V. POKORNY ©)
ABSTRACT
Four species of the genus Lipara, viz.: L. lucens, similis, rufitarsis and pullitarsis, occur in Europe, where they cause galls in Common Reed (Phragmites communis). The specific differences of eggs, larval, and adult stages are described, and a general account is given of the life cycle, gall formation, precopulatory and ovipositional behaviour, habitat relations, and influence of parasites and predators with emphasis on the known differences between the species.
CONTENTS
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1) Department of Systematic Zoology, Faculty of Natural Sciences, Charles University, Viniëná 7, 128 44 Praha 2, Czechoslovakia.
2) Institute of Ecological Research, Kemperbergerweg 11, Arnhem, the Netherlands.
3) Institute of Scientific and Technical Information, Czechoslovak Akademy of Agriculture, Slezská 7, 120 56 Praha 2, Czechoslovakia.
2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
1. INTRODUCTION
The galls on Common Reed (Phragmites communis Trin.), caused by flies of the genus Lipara, have attracted the attention of many generations of entomologists in Europe. During more than one hundred years, after several entomologists in Vienna had collected reed galls and studied their inhabitants (Schiner 1854, Heeger 1856, Loew 1858, Giraud 1863), three species were recognized: L.lucens, rufitarsis, and similis.
In 1957 Ruppolt mentioned the occurrence of two types of larvae of L. rufitarsis, one in typical galls and the other in galls indentical to those of L. /acens. He supposed that these might be two different species.
From 1967 to 1970, Pokorny took up the problem again in Czechoslavakia, and found that the two types of L. rzfitarsis make distinctive galls. The differences he observed also in the life cycle and the behaviour of the larvae were such that he expected the insects to belong to two different species. He submitted the adults, bred from the galls, to Doskocil and Chvála who indeed recognized them as two different species, one being L. rufitarsis and the other, a previously undescribed species which they named L. pullitarsis (Doskotil and Chväla, 1971).
In recent years Mook studied the ecology of Lipara lucens in the Netherlands (Mook, 1961, 1967) and also collected data on the ecology of L. rufitarsis and pullitarsis (L. similis does not occur in the Netherlands). In co-operation with Miss C. G. Bruggemann he studied the behaviour of the adult of L. /zcens (Mook and Bruggemann, 1968, 1971) and also collected data on the behaviour of the other species. These data corroborate the view that four different species are involved.
It seemed worth while to collect both the published and the unpublished data in order to give a survey of the similarities and differences in the biology of the four species. The results are presented here. The authors have each treated those aspects of the biology with which they are most familiar, and they remain responsible for the relevant section: Chväla and Doskoëil for section 2, Pokorny, for section 3 and Mook, for sections 4 and 5. It should be kept in mind that the different geographical localities where the authors collected their data may have influenced their findings.
2. SYSTEMATICS AND MORPHOLOGY a. Adults Genus Lipara Meigen, 1830
Lipara Meigen, 1830: 1. Type species, Lipara lucens Meigen, 1830 (monobasic).
Homalura Wiedemann, 1830: 573 (nec Meigen, 1826), junior homonym. Type species, ee grisea Wiedemann, 1830 (monobasic).
? Gymnopoda Macquart, 1835: 503, subjective synonym. Type species, Gymnopoda tomentosa Macquart, 1835 (monobasic).
The genus Lipara Meigen belongs to the family Chloropidae, subfamily Oscinellinae, and comprises medium-sized to large blackish species, 3.3—7.3 mm, with dense pale pubescence on mesonotum. Arista thin, not pubescent, antennae broadly separated at base by a wide pollinose facial keel, very prominent in lateral aspect. Eyes rather small and finely pubescent, face very deep, almost as long as one half of the eye-height; vibrissal angles absent.
M. CHVALA, J. DoskoCr, J. H. Mook & V. Pokorny: The genus Lipara 3
The genus nearest related to Lipara is Calamoncosis Enderlein wich comprises dis- tinctly smaller species, 2-3 mm, with equally oval head in profile, with facial “keel” only indistinctly developed. In Lipara the facial keel is prominent and very broad over the whole length, as far as the mouth cavity, or, if it is narrowed in the middle between the antennae, again widening out below; viewed from the side, the keel is strongly prominent above the antennae, as is the lower part of the frons, so that the head in profile ist strongly prominent anteriorly.
The larvae of all Lipara species cause more or less distinct galls on stems of the Common Reed (Phragmites communis Trin.).
The genus has been recently revised by Doskoëil and Chvála (1971) and as a result four Palaearctic species (viz., lucens, similis, rufitarsis, and pullitarsis) were introduced as essentially European species. Two of them, /zcens and similis, have been found separately as immigrants in North America.
The four Palaearctic Lipara species may be keyed as follows:
1 Thoracic pubescence long, arranged in ee stripes, A a ridged
appearance. Large or smaller species . . RE 2
— Thoracic pubescence shorter, uniformly spread not angel i in Stripes Smaller Speciesy I . . to sac 3
2 Thoracic eee: bes Helen uo species 5. Je 3 mm, gel keel very progam Pa. .\. ee ee ens
— Thoracic pubescence ele Smaller Ser 3. 34. 6 mm, ia keel narrower A similis
3 Facial keel a a Ato i Sii ol as banal as one-half of antennal segment 3. Male genitalia with fused cerci. Length 3.3—5.0 mm pullitarsis
— Facial keel narrowed in the middle, widening above and below, less broad than one-third of antennal segment 3. Male genitalia with paired cerci. Length 3.5— ENE ee NN af ee 4 AN raftarsis
Lipara lucens Meigen, 1830
Lipara lucens Meigen, 1830, Syst. Beschr. 6: 1. Homalura grisea Wiedemann, 1830, Aussereurop. zweifl. Ins., 2: 573. ? Gymnopoda tomentosa Macquart, 1835, Hist. nat. Ins. Dipt., 2: 502.
Diagnosis. A conspicuously large and robust species, 5.3—7.3 mm, the largest species of the genus and family in the Palaearctic region. Thorax with long, brassy-yellow pubescence on mesonotum which is arranged in distinct longitudinal stripes, producing a ridged appearance. Antennae yellowish on basal segments, segment 3 mostly blackish. Facial keel conspicuously broad and with almost parallel sides. Legs extensively darkened. Male genitalia (Fig. 2) large, with small but rather widely separated, apically pointed cerci.
Distribution. A widely distributed and everywhere common species in Europe, ranging in the North from England through the Netherlands, Denmark, and southern Sweden, eastward as far as the northern regions of the European part of the USSR, a very common species in central and southern Europe, immigrant ín North America (Con- necticut).
4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
Lipara similis Schiner, 1854
Lipara similis Schiner, 1854, Verh. zool.-bot. Ges. Wien, 4: 172.
Diagnosis. Closely related to /zcens, having the same type of long pubescence on thoracic disc which is arranged in distinct longitudinal stripes, producing a ridged appearance. In contrast to /zcens, the thoracic pubescence is distincly whitish to silvery- grey; similis is a much smaller species, 3.3—4.6 mm. Facial keel rather broad, slightly widened above and below. Antennae and palpi yellowish. Legs mostly yellow on tibiae and tarsi, but in some specimens tarsi extensively darkened. Male gentitalia (Fig. 4) small, cerci not very wide apart, small, and apically distinctly rounded.
Distribution. The species is rather common and widely distributed in Central Europe (Poland, Germany, Czechoslovakia, Austria, and Hungary), and has also been recorded from England, France, and the western regions of the European part of the USSR.
Figs. 1—8. Head in profile and male genitalia (caudal view) of Lipara: 1, 2, Iucens Meig.; 3, 4, similis Schin.; 5, 6, L. pullitarsis Dosk. et Chv.; 7, 8, L. rufitarsis Loew. Scale = 0.3 mm
M. CHVALA, J. DoskoCr, J. H. Mook & V. POKORNY : The genus Lipara 5
According to Sabrosky (1958), the species was intercepted in the USA (Hoboken) in Phragmites packing from a ship from the Netherlands. It was also mentioned for the Netherlands by Docters van Leeuwen (1957), but his description and illustration of the gall do not fit. The species has not been reared from Dutch galls (de Meijere, 1949). Dr. Th. van Leeuwen, Dr. W. J. Kabos, Dr. G. Kruseman (Amsterdam) and Dr. S. J. van Ooststroom (Leiden), who were so kind as to inform us of their experiences, also did not find the species in the Netherlands.
Lipara pullitarsis Doskoëil et Chvála, 1971
Lipara pullitarsis Doskotil et Chvála, 1971, Acta ent. bohemoslov., 68: 102.
Diagnosis. A smaller species, 3.3—5.0 mm, with mesonotum densely covered with mostly brassy-yellow short pubescence, not arranged in longitudinal ridges. Facial keel very broad, the narrowest part in the middle about as broad as half the width of antennal segment 3. Antennae mostly black, only basal segments sometimes very dark brown; palpi black. Legs mostly black, extreme tips of femora and extreme bases of tibiae slightly brownish; fore tarsi black, posterior four tarsi yellowish-brown on basal segments, apical segments extensively darkened. Male genitalia (Fig. 6) rather large, with broad parameres and entirely fused long cerci. The Central European and southern Scandinavian pop- ulations have extremely darkened antennae and legs, but we have seen paler specimens from the Netherlands that superficially resemble r#fitarsis, differing from the latter by a much broader facial keel and quite distinct male genitalia.
Distribution. This species has long been confused with rwfitarsis, and it is highly probable that many records on the distribution of r#fitarsis refer, in fact, to pullitarsis. The species was described very recently from Czechoslovakia but we have also seen documentary material from the Netherlands, Denmark, and Sweden.
Lipara rufitarsis Loew, 1858
Lipara rufitarsis Loew, 1858, Wien. Ent. Monatschr., 2: 57.
Diagnosis. A smaller species, 3.5-—5.3 mm, resembling pullitarsis in having the same type of pubescence on thoracic disc; the hairs are short and not arranged in longitudinal ridges but, in contrast to pullitarsis, the pubescence is distinctly silvery-grey and the hairs on the hind part of mesonotum and on the scutellum are distinctly longer. Facial keel is much narrower than in pullitarsis, widening above and below, the narrowest part in the middle being hardly as broad as one-third the width of antennal segment 3. Antennae reddish-brown except for outer side of segment 3 and arista which are extensively darkened or almost black; palpi black. Femora and tibiae predominantly black, the apical tips of femora, both rather broader tips of tibiae and whole tarsi yellowish- to reddish-brown. Wings with cubital and discal veins almost straight, not curved as in pullitarsis. Male genitalia (Fig. 8) rather large, with narrower parameres, long cerci apically distinctly separated.
Distribution. L. rufitarsis is recorded in the literature as a common species throughout Europe, but at least some of these records must concern pullitarsis. We have seen the documentary material from the Netherlands, Czechoslovakia, Austria, and Italy, and
6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
according to Dr. Hackman (letter of 24.8.1971), the material from southwestern Finland also belongs to rfitarsis. On the other hand, having revised in Copenhagen and Lund all of the accessible vufitarsis material from Denmark and southern Sweden, one of the authors (Chvála) comes to the conclusion that all of it belongs to pullitarsis.
b. Immature stages
There are few older data on immature stages of Lipara; a summary of all the records up to 1950 is given by Hennig (1952). He found only four records of larva and puparium of /zcens in the literature, but the record of larva by Vimmer (1925), must, in fact, refer to rufitarsis judging from the Czech description and figures. Ruppolt (1957) described eggs and larval stages of /cens. The most complete treatment of the morphology of immature stages of lwcens is by Waitzbauer (1969). No other records are available.
Most of the eggs, larvae, and puparia used for the study discussed in this section, belong to material collected and bred by J. H. Mook and V. Pokorny.
Eggs
The eggs of the four Palaearctic Lipara species are very distinctive. In general, they may easily be separated into two different groups, viz., (1) the /ucens type and (2) the rufitarsis type. The former type (/ucens and similis) is characteristically rather broader, more oval, and light yellow in colour. The differences between the eggs of lucens (Plate 2b) and similis (Plate 2d) are very slight; the egg of similis is almost as long and as wide as that of /cens, but the latter has more distinct sculpture, con- sisting of rather deep longitudinal furrows (Plate 2a). The measurements of the eggs of these two species are as follows: /ucens: (N = 40) length 1.37—1.69 mm (M = 1.57 mm), width 0.25—0.36 mm (M = 0.32 mm); similis: (N = 20) length 1.35— 1.46 mm (M = 1.42 mm), width 0.23—0.32 mm (M = 0.27 mm). The ratio of length to width is about 5:1 in both /zcens and similis.
The second type of egg (rufitarsis and pullitarsis) is characteristically long and narrow, and distinctly blackish in colour. The egg of pullitarsis (Plate 3b) is distinctly narrower, slightly shorter, rather circular in cross-section, and the longitudinal furrows (Plate 3a) are quite distinct. The eggs of rufitarsis (Plate 3d) is less slender but slightly longer, somewhat triangular in cross-section (rounded above, flattened below), and the long- itudinal furrows are less numerous and barely visible; the whole egg is therefore rather smooth. The measurements of these two eggs are: pullitarsis: (N = 30) length 1.13— 1.26 mm (M = 1.21 mm), width 0.07—0.11 mm (M = 0.09 mm); rwfitarsis: (N = 40) length 1.39—1.53 mm (M = 1.50 mm), width 0.15—0.20 mm (M = 0.18 mm). The ratio of length to width in rwfitarsis is 8 : 1, in pullitarsis 13 : 1.
Larvae Larvae are elongate, light yellowish, with one or both tips more or less sclerotized; the length of mature larvae varies from 5.5 to 12 mm; all measurements were taken from larvae collected in the winter or early spring. The whole body is rather smooth, integument exclusively with regularly spaced flower-like not sclerotized papillae and sometimes with striae, especially on anterior segments. The sclerotization of anterior and posterior segments, including the number of buds on anterior spiracle, seem to be
M. CHVALA, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 74
the best specific characters. Posterior spiracle consists of three simple radiating slits, interstigmatal processes are branched (? absent in rzfitarsis).
The mature larva of /ucens (Plate 6a) is conspicuously large and stout, 8—12 mm in length and 2—3 mm in width. Only anterior segments dorsally sclerotized (Fig. 9), more heavily on segment 1 as far as the anterior spiracle, a slight sclerotization con- tinues over anterior spiracle on dorsum of segment 2 and anterior half of segment 3. Segment 1 sometimes also slightly sclerotized ventrally. Posterior margin of anterior three segments on dorsum with small sclerotized papillae in a row; on venter also present but hardly visible (not sclerotized). Anterior spiracle (Fig. 13) with 11 to 12 (exceptionally up to 14) buds, their number not constant on each side; the usual com- binations are 11—11, 11—12, or 12—12, in rare cases, 12—13 or 12—14.
Figs. 9—12. Four anterior segments of mature larva of Lipara (lateral view). 9, L. lucens Meig.; 10, L. similis Schin.; 11, L. pullitarsis Dosk. et Chv., 12, L. rufitarsis Loew. Scale = 1 mm
8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
The mature larva of similis (Plate 6b) generally is smaller and especially more slender, 5.5—10 mm in length and 1—2 mm in width; it is the only species of the genus with distinct sclerotization of both tips. Segment 1 (Fig. 10) heavily sclerotized dorsally, except for a narrow anterior strip, segments 2 and 3 heavily sclerotized dorsally, almost blackish, as is segment 1. Segment 4 dorsally with a slight sclerotization, venter of four anterior segments pale, membraneous. Segments 5 to 10 membraneous, segment 11 slightly sclerotized dorsally and ventrally, almost membraneous laterally. Segment 12 heavily sclerotized throughout, having the same blackish colour as anterior three segments dorsally. Posterior margins of segments 2, 3, and 10 dorsally with a row of slightly sclerotized papillae, these membraneous only on venter. Anterior spiracle (Fig. 14) with 9 tot 11 buds with usual combinations 9—9, 9—11, 10—11, 11—11, or 9—10.
The mature larva of pullitarsis (Plate 6c) is 6.5—9 mm long and 1.5—2 mm wide, with sclerotization on segment 1 similar to that in rwfitarsis. Dorsum of segment 1 (Fig. 11) more or less heavily sclerotized, lateral area almost membraneous, and venter evenly and slightly sclerotized, without any distinct pattern. Slight sclerotization also visible on the anterior ridge. No rows of membraneous papillae have been found on posterior margins, but distinct papillae are visible on anterior ridge and among the striation on segment 1 ventrally; this segment distinctly striated both ventrally and dorsally, segment 2 on anterior half dorsally and ventrally, segment 3 on anterior half ventrally only. Otherwise the integument is smooth, except for regularly spaced, small, flower-like papillae. Anterior spiracle (Fig. 15) with 4 to 5 buds, the number on each side often differing.
The mature larva of rufttarsis (Plate 6d) has in general much in common with pullitarsis; it is rather smaller, 6—8.5 mm long and 1.5—2.5 mm wide, more or less sclerotized only on segment 1. The dorsal sclerotization (Fig. 12) is not very distinct (light brown in colour), almost interrupted laterally near anterior spiracle and again more distinct on a triangular patch ventrally. Papillae on posterior margins always mem- braneous and therefore inconspicuous. Whole of segment 1 distinctly striated except for the anterior ridge, which bears very small, not sclerotized papillae. Segment 2 with fine striae on anterior half ventrally, otherwise segments smooth. Anterior spiracle (Fig. 16) with 6 to 9 buds in combinations 7—8, 8—9, 7—7, 6—7, or 6—6.
The mature Lipara larvae may be keyed as follows: 1 Larva heavily sclerotized (blackish-brown in colour) on both tips, anteriorly on 4, posteriorly on 2 segments. Anterior ons with 9 to 11 buds. Larva about 8 mm
[On ce A See rito — Larva less AT oi oh in lo), ci antes at most on the three anterior segments. Posterior segments pale, membraneous. . . 2
2 Three anterior segments slightly sclerotized on dorsum. All thoracic segments with a row of heavily sclerotized papillae on posterior margin, segment 1 without a ridge on anterior margin. Anterior spiracle with 11 to 12 (or rarely up to 14) buds. Larva about 10 mm long and very stout. . . . PM acer’
-— Only segment 1 slightly sclerotized. Sclerotized papillae on ost margins of thoracic segments absent, segment 1 with a distinct ridge on anterior margin.
Fig. 13—16. Anterior spiracle of mature larva of Lipara. 13, L. lucens Meig.; 14, L. similis Schin; 15, L. pullitarsis Dosk. et Chv.; 16; L. rufitarsis Loew. Scale = 0.1 mm
M. CHVALA, J. Dosko£ır.. I. H. Mook & V. POKORNY : The genus Lipara
10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
Anterior spiracle with at most 9 buds. Larvae generally smaller, about 8 mm long, and ‘slender 1, EREN EE PSE ER ARR ER TAO 3 Anterior spiracle with 6 to 9 buds. Segment 1 usually slightly sclerotized, also ventrally, distinctly striated on the whole surface, except the smooth anterior ridge; segment 2 with slight striae on anterior half ventrally; segment 3 smooth, without Straen etn PAN AS A EREN EEE I e E PN OE — Anterior spiracle with 4 to 5 buds. Segment 1 usually membraneous beneath, at most quite indistinctly sclerotized, but with distinct striae throughout, also on anterior ridge; segment 2 with striae on anterior half both ventrally and dorsally; segment 3 with striae on anterior half ventrally. . . . . . . . pallitarsis
Puparia
The puparia are brown to reddish-brown, with the same specific characters as in mature larvae, i.e. of the same size, different colour of tips, and the same number of buds on the anterior spiracle. L. /zcens has a large and stout puparium, 8 to 12 mm long, the anterior two segments distinctly darkened dorsally, anterior spiracle with 11 to 14 buds; puparium of similis slightly smaller on average, distinctly blackish over three anterior and two posterior segments, anterior spiracle with 9 to 11 buds; rufitarsis and pullitarsis have smaller puparia, about 6 to 9 mm long, slightly darkened on segment 1 only, anterior spiracle with 6 to 9 buds in r#fifarsis and 4 to 5 buds in pullitarsis.
3. LIFE CYCLE AND INFLUENCE ON THE REED STEMS a. Life cycle and gall formation
By their action in the larval stage, all four Lipara species induce the formation of a terminal gall on the infested shoot, thus altering the length of the stem, the number and dimensions of internodes and the formation of a panicle. Each gall is ultimately inhabited by a single larva, even when shoots are infected by more than one egg.
The formation of the gall is principally the same in all four species, and is closely related to the behaviour of the larvae. The egg-laying behaviour of the fly is described in section 4. Several days after the eggs are laid the larvae hatch. In L. Iwcens the larvae hatch after about 9 days. Hatching takes place virtually only during the night. This is probably important for survival, because in order to escape drying out the larvae require a very high humidity (Ruppolt, 1957, Mook, 1967). The young larva crawls upward on the surface of the shoot untill it reaches the top part, and then crawls under the edge of a leaf sheath into the roll of young leaves. There it feeds on the young leaf parenchyma and bores downward toward the growing point. Just above this point it continues to feed on the newly-formed leaves. Here the larva molts twice. The growing point is affected by the feeding of the larva and probably also by its metabolic products. Internodes formed subsequently remain shorter than the internodes of a healthy stem.
The shortening of the internodes leads to the formation of the terminal gall in the stem. The gall itself is formed (except in the case of L.lwcens) by the leaves growing from the agglomerated nodes. The leaf-sheaths do not have sufficient room to grow parallel with the stem axis and are pushed off at an angle to it; in this way the typical cigar- or spike-like inflation develops.
Up to this point the life cycle is basically the same for all the four species. The
M. CHVALA, J. DoskoCit, J. H. Mook & V. Pokorny : The genus Lipara 11
differences become apparent at a later stage, and therefore the rest of the life cycle will be described for each species separately.
During July or August the larva of L. /ucens (Plate 6a) bites through the growing point and lives in the shortened internodes where it feeds on the soft parenchyma filling these internodes. In due time it gnaws a chamber passing into the 6th to 8th internode from above. At the end of the summer and in the beginning of the autumn the larva turns 180° in the chamber, so that its head is pointing upward. This turn is very important (and takes place in the other species, too), because it enables the adult to leave the gall. After turning, the larva shifts its position 2 to 3 cm higher, living without nutrition approximately 1 to 2 cm under the remnants of the growing point; at that time it is 8—12 mm long, thick, and ivory white. In this position it also hibernates and pupates in the spring. In Czechoslovakia and in the Netherlands, depending on the climatic conditions, the adults emerge as the second Lipara species at the end of May or beginning of June (Fig. 17).
I Q D Q
FEBR. MARCH APRIL MAY JUNE JAN. FEBR. ERBUÜCENS ee L. RUFITARSIS L.SIMILIS L. PULLITARSIS
Fig. 17. Life cycle of Lipara species in Czechoslovakia
The growing point of the reed is also perforated by the larva of L. rufitarsis (Plate 6d), but in this species the chamber reaches only to the 3rd or 4th (shortened) inter- node from the top. The walls of the shortened internodes are not thickened and their width is the same as that of the adjacent not shortened ones. Once arrived below the growing point the larva feeds until some time around August, after which it turns and rests until spring, when it pupates. At that time it is 6—8.5 cm long, slightly yellowish, and more flattened than the larvae of the other species. The adult emerges as the third Lipara species at the beginning of June.
12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
These two species are characterized by a common behaviour pattern viz., biting through the growing point and life in the chamber in the shortened internodes. The other two species leave the growing point intact, living throughout their whole lifespan between the leaves above this.
In the beginning, the larva of L. pullitarsis (Plate 6c) feeds on young rolled leaves in which it “chops” its chamber. Later on, when the panicle is beginning to be formed, it feeds on this rudiment often completely destroying the panicle. Sometimes, probably when infestation occurs later, a panicle large enough to survive is formed in the gall, only to have the larva gnaw through it, too, to maintain its chamber. Such panicles never develop fully. Before turning, the larva usually reaches the growing point, but after having turned, it shifts itself 3 to 5 cm above the growing point and hibernates there. At that time it is 6.5—9 mm long and white. In the spring, as soon as the days become warmer, the larva pupates as the first of the Lipara species and the adults emerge also as the first, in the second half of May.
Similarly, the larva of L. similis (Plate 6b) lives in the tube of leaf sheaths. In the beginning it feeds on the rolled leaves, which later becomes completely severed from the growing point, so that they can be pulled out by the tips. The space between the leaves is relatively narrow and the adult larva fills it completely. As soon as the panicle begins to develop the larva starts to feed on the rudiments, thus destroying the panicle com- pletely. Even in the narrow cleft in the tube the larva turns and shifts itself upward. During the winter and early spring, the two sheaths become hard, thus protecting the hibernating larva, which at that time lies 5 to 7 cm above the remnants of the growing point. In this period it reaches a length of 5.5 to 10 mm, and is slim, milky white, and glossy, with both ends black. In the same position, the larva pupates in spring as the last of the Lipara species. The adults emerge, also as the last, in the middle of June.
b. Description of the galls
The number of shortened internodes is more or less characteristic for the species and thus also for the shape of the gall (Fig. 18 and Plate 4). The most massive and prominent galls result from the influence of the larva of L. lucens (Plate 5a), and this corresponds with the greatest number of shortened internodes, usually 10—13. The shortened internodes are 2 to 3 times wider than the adjacent normal ones, and their walls are thickened and strongly lignified (Plate 4a). All of the 6 to 8 uppermost shortened internodes are brownish and covered with a continuous hairy layer. The agglomerated nodes are also provided with a wreath of hairs, and the leaves growing from them have short broad sheaths and weaker short blades. The highest leaves are relatively broad and short, again covered with a continuous hairy layer; these lena do not differentiate into sheath and blade.
Due to the activity of the larvae of L. rzfitarsis or L. pullitarsis, only 5 to 6 internodia are manifestly shortened (Plate 4c). The galls (Plate 5c, d) in these two species are usually similar, and can only be conclusively differentiated by a longitudinal section through the gall. The galls are medium-sized to large, the pullitarsis galls being usually larger, since pullitarsis occurs on wider shoots. This is why its galls are sometimes larger than those of /zcens. But if the galls of lwcens and pullitarsis appear simultaneously on equally wide shoots, those of lwcens are always larger (Fig. 20).
The stem infested by L. similis has the smallest number of shortened internodes, i.e.
PLAAT 1
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
P
*(AUIOYOG “A ‘30Yg) MaOT szsargufma
"I ‘p
“AUD 39 ‘YSOQ Szsartmnd “JT ‘D SUIS szprwis "TÈ
rie,
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e)
The genus Lipara
2
KORNY
M. CHVALA, J. Doskocit, J. H. Mook & V. Po
PLAAT 2
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
+
IJ
X 160 (Phot.
lis Egg: X 70, detail:
mt
5
a UGC DSS @ chel,
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b
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a
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Plate 2. Lipara eggs with deta
J. Chalupsky)
M. CHVALA, J. Doskocit, J. H. Mook & V. POKORN
‘a
The genus Lipar
x
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 3
Plate 3. Lipara eggs with detail. a ‚b, L. pullitarsis; c, d, L. rufitarsis. Egg: X 70, detail X 160 (Phot. J. Chalupskÿ)
M. CuvAra, J. Doskoém, J. H. Mook & V. POKORNY : The genus Lipara
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 4
Plate 4. The highest internodes of reed stems with a gall of Lipara. a, L. lucens; b, L. similis; c, L. pullitarsis (Phot. V. Pokorny)
M. CHVALA, J. DoskoCit, J. H. Mook & V. POKORNY : The genus Lipara
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 5 er
Plate 5. Galls typical of the individual species of Lipara. a, L. lucens; b, L. similis; c, L. pullı- tarsis; d, L. rufitarsis. The size relations are maintened (Phot. V. Pokorny)
M. CHVALA, J. Doskocit, J. H. Mook & V. Pokorny: The genus Lipara
PLAAT 6
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
DI
Yor”
4
d, L. rufitarsis (Phot. V.
L. pullitarsis;
)
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b, L. similis;
,
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Plate 6.
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The genus Lipara
Yi
POKORN
KOCIL,
ALA, J. DOS
ALA
. CHV
M
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 PLAAT 7
Plate 7. Signals of Lipara species registered with an UV recorder. a, signal of male L. /zcens with part of answering signal of female; b-e signals of males of different species: b, L. /ucens; c, L. similis; d, L. pullitarsis; e, L. rufitarsis
M. CHVALA, J. DoskoCit, J. H. Mook & V. POKORNY : The genus Lipara
M. CHVALA, J. Doskocit, J. H. Mook & V. Pokorny: The genus Lipara 13
AVERAGE LENGTH OF INTERNODES (CM)
4 2 3 4 5,6 TMD meten On 4e WAZ. INNETNETRENNONDNESS
Fig. 18. Length of the highest internodes (counted from above) of reed stems attacked by larvae of Lipara species. (average of 30 stems per species)
3 to 4. This is why the gall is the smallest and least noticeable (Plate 5b). Due to this small number of shortened internodes and also to the fact that the shortened internodes are narrower than the adjacent normal ones but rather telescopic (Plate 4b), the sheaths have enough room and embrace the stem closely. The gall itself is formed only by the leaf sheaths growing from the two highest visible nodes which form an approx- imately 9 cm long tube concealing the remnants of the rolled leaves, the larva and the remainders of the panicle and the growing point. This is why the gall is not inflated and resembles the top of a sterile shoot, from which it differs only by the fact that the sheaths of the two uppermost leaves covering the gall are almost equally long. By summarizing these descriptions we arrive to the following key to the galls: 1 There is a terminal inflation on the stem: a cigar- or spike-shaped gall. . . 2 — There is no terminal inflation of the stem, it looks like the top of sterile shoot. In the tube formed by the leaf sheaths growing from the two highest visible nodes there is a slim, milky white, glossy larva with both ends black (or brown pupa with both ends black) . . . MORT 0 2 Ina longitudinal section through the Di a Abe uo doni the young rolled
14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
leaves can be seen. In the chamber there is a big white larva (or brown pupa). The chamber never passes SE the “ua B to the shortened internodes .
pullitarsis — The lano n tai ie rest ci the growing point in order to reach the shortened internodes. Its inner walls blak. . . . 3
3 Walls of the shortened internodes thickened, their tn two or Bin dim that of adjacent not shortened ones. The chamber passing through 6 to 9 shortened internodes, containing a big, thick, ivory white larva (or a brown pupa). The shortened internodes brown . . . EN lacens
— Walls of shortened internodes neither Lidi nor bon their width equal to that of the adjacent not shortened ones. The chamber passing through 2 tot 4 shortened internodes only and containing a big, yellowish, and somewhat dorsoven- trally flattened larva (or brown pipa), A 4
4. BEHAVIOUR OF ADULTS
The discussion of behaviour and ecology will be somewhat one-sided, because Lipara lucens is by far the best known species of the four and the outline of its biology will therefore be the largest. The fragmentary information concerning the biology of the other species will be compared with that of /ucens.
a. Precopulatory behaviour
The adults of Lipara lucens force themselves upward between the leaf sheaths and emerge from tip of the gall during the early morning hours. In this process an important role is played by the ptilinum, a large sac of the head, which can be expanded, so pushing aside any obstacle in front of the head. It is also useful for bursting the puparium, and in other species of higher flies, for forcing a way up through the soil (Oldroyd, 1964). In the hardening process after emergence, the ptilinum is retracted inside the head.
The flies emerge over a period of two or three weeks in the end of May or the beginning of June. Our observations do not indicate that the female flies emerge a few days before the males, as is stated by Waitzbauer (1969).
The female flies are probably very sedentary before copulation, but detailed ob- servations on the behaviour in the field are not yet available. In laboratory experiments, where the flies are observed in small cages provided with some old reed stems, the females remain in the same place for a long time unless the temperature becomes high (above about 25° C), when they become restless. The males move about much more, and it is often obvious that they are in search of a female. They fly from reed stem to reed stem, on each stem giving a signal by vibrating. A virgin female fly sitting on the same substrate as the male perceives this signal as a vibration of the substrate and im- mediately answer by vibrating in its turn (Plate 7a). This in turn induces in the male increased searching on the reed stem. It starts to walk over the stem, stops after a few seconds to give a signal again, and when this is answered, begins to walk again. In this way the male finally finds the female, and copulation often takes place without obvious further display.
When a male gives its signal on a stem and does not receive an answer, it soon leaves the stem and flies to another. Therefore the function of this behaviour seems obvious. This procedure requires much less time than the male would need for searching the whole reed stem.
M. CHVALA, J. Doskocit, J. H. Mook & V. Pokorny: The genus Lipara 15
The signals of both the male and the female fly cannot normally be heard by the human ear, but the male’s signal is perceived when seated on a reverberating substrate. Both signals can be made audible by placing a crystal gramophone pick-up element in contact with the substrate, for instance a reed stem, and by connecting this element to a tape recorder. The tape recordings can then be converted to visual signals with the help of an oscilloscope or an UV recorder (Plate 7).
This description is based primarily on observations of L.lwcens (Mook and Brugge- mann, 1968, 1971), but behaviour of other species is known to be similar. Important differences between species were found in the signals of the males, characteristic examples of which are given in Plate 7b-d. The main pitch of these sounds is about the same (a few hundred Hz.), but there are wide differences in the rhythm making it easy to recognize the different species by their signals. There are no great differences between the signals of the females. They all have a pattern similar to the one shown in Plate 7a; only the duration varies, that of L. pullitarsis being characteristically short (one or two seconds) while that of the other species is usually longer (up to over 10 seconds). No differences were found between the signals of flies collected in Czechoslovakia and the Netherlands.
Because such marked differences were found between the signals of the males, an experiment was set up in order to investigate whether these differences play a role in reproductive isolation. In this experiment, sounds of three species of Lipara and two species of Haplegis (small Chloropid flies, the larvae of which occur as inquilines in Lipara galls) were played back to females of the three Lipara species. The female flies rested on a strip of paper in contact with a small loudspeaker (output 0,5 watt). The tape gave each of the signals ten times in a random sequence, with a 30 second interval between signals. The tape was played back twice to each of five females of the three different species, so that each female was subjected to each signal twenty times, and their response was observed with a binocular microscope. The number of responses (Table I) was high only for the own species, which makes it probable that the signals do indeed play a role in reproductive isolation.
Table I. Reactions of 5 females of 3 Lipara species to 20 male signals of 5 different Chloropidae
3 signal Reactions of 9
lucens pullitarsis rufitarsis L. lucens 18 16 18 19 18 — — — — — — — — — — L. pullitarsis u Ze a ea EEE CL CERN L. rufitarsis Zeelse mer. — — 19, 19720: 17 17 Haplegis spec. A SS SD ol, ai er Haplegis spec. B - Jos oo ol oo do —
b. Egg-laying behaviour
The egg-laying behaviour of Lipara lucens has been investigated extensively (Mook, 1967). The species is autogenic: it does not need food to lay eggs. At emergence the ovaries contain almost mature eggs, and egg-laying can start two or three days later. The number of eggs in the ovaries of L. lucens lies between 64 and 96, with a mean of 84 (data from 40 Dutch specimens). However, egg production is not limited to this
16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
number. In laboratory experiments the flies were often observed laying eggs in two or three periods of one or two days, each period separated by two or three days in which no eggs were laid. The number of eggs laid during the first of these periods cor- responded to the number of mature eggs in the ovaries at emergence. In the later periods a smaller number of eggs was laid, so that the total production of eggs often attained twice the number of eggs originally present in the ovaries. Under favourable circumstances (the flies were provided daily with fresh drinking water and pieces of fresh reed shoot of the preferred diameter; light and a temperature of 20° C were maintened for 7 hours each day) the number of eggs laid was between O and 199, with a mean of 133.6 (median 147).
Comparable data for the other species are lacking. At emergence they too have almost mature eggs in the ovaries. For L. pullitarsis the number of eggs in the ovaries lies between 98 and 157 in 20 Dutch specimens and for L. rufitarsis between 54 to 83 in 17 Dutch specimens. No data are available for L. similis.
Lipara lucens lays its eggs during the warmest hours of the day. The female flies to a reed shoot and walks on it, first upward. After reaching the top of the shoot it often turns and walks down again. During this walking the ovipositor may be slightly extended, but does not touch the shoot. After a while the female begins to walk on a more restricted part of the shoot, a few centimeters’ distance. By this time the ovipositor is fully extended and its tip now and then touches the surface of the shoot. The fly then pauses for about 60 seconds with its head pointing upward, after which the egg emerges from the tip of the ovipositor and is pressed against the surface of the shoot. After the egg has been laid the female walks upwards and usually flies away to another shoot.
The eggs are usually laid on the stem, sometimes but not always on or under the hairy ligules, but in an outdoor cage an appreciable proportion of the eggs was found on the reed blades. The flies show a marked preference for ovipositing on shoots of a certain diameter; nonpreferred shoots are generally deserted during the walking stage of this ovipositional behaviour. Because this preference influences the habitat relationships, it will be treated in the next section.
5. ECOLOGY a. Habitat relations
In the literature it is often stated that the galls of Lipara lucens are most numerous in reed stands located in dry places (Wagner, 1907; Docters van Leeuwen, 1957; Ruppolt, 1957), but this does not necessarily imply that the water level itself influences the species. Together with the water level, characteristics of the vegetation vary consider- ably. The number of shoots per square metre, for instance, tends to be higher in drier places (but is lower again where the soil becomes very dry). The dimensions of the reed shoots are also influenced by the water level, the shoots being shorter and thinner in dry than in wet places.
A partial regression analysis of samples of three different vegetations showed that the variation in gall density is not primarily due to variations in water level or shoot density, but to variations in the dimensions of the shoots. In this analysis and in the subsequent experimental work the basal diameter of the shoots was chosen as the most characteristic shoot dimension, because this is the most constant measure throughout the year and is not influenced by gall formation. The basal diameter does not alter during
M. CHVALA, J. DoskoCit, J. H. Mook & V. PoKoRNY: The genus Lipara 17
the growing season and when the reed dries in the autumn it diminishes only by about 10 per cent.
Experimental work in the field and in the laboratory showed that the diameter of the reed shoots plays an important role in the biology of Lipara lucens, in some cases directly affecting the behaviour of the fly or its parasites, in other cases correlated only with the real operating factor. The correlation of life functions of Lipara lucens with the diameter of the reed shoots has been treated elswhere (Mook, 1967); only a short survey will be given here.
The female flies lay their eggs preferentially on shoots with a width of 4 and 5 mm at the base. In Fig. 19a oviposition is expressed as the mean number of eggs per shoot, thus eliminating the influence of the number of shoots available in each class. Waitzbauer (1969) found a distribution of eggs in the field with an optimum on shoots with a diameter of 4—4.5 mm, but he did not take into account the distribution of shoot
0.6 04
0.2 d.
eggs per shoot
0.0 ovıposition preference
0.6
04
b.
survival of larvae up 0.0 to gall formation
0.2
0.8 0.6
0.4 C.
fraction of surviving larvae
0.2 survival of larvae 0.0 in the gall 2 3 4 5 6 4 8 mm
basal shoot diameter
Fig. 19. Lipara lucens Meig. Correlation of oviposition and survival of larvae with diameter of reed shoots
18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
diameter in the vegetation. He states that Mook (1967) found a preference for shoots with a diameter of 2.5—2.9 mm, but this is derived from a laboratory experiment in which the diameter of the shoot top was used. This value corresponds to a basal diameter of about 4 mm.
Once a shoot has received an egg, the probability that a gall will be formed is also correlated with shoot width (Fig. 19b). It is in the shoots with a diameter of about 2 and 3 mm that the young larvae have the greatest chance to induce a gall and thus the greatest chance to survive. The differences in survival of larvae in the gall (fig. 19c) are due to a heavier parasitism by Stenomalina and more predation by birds on the thinner shoots.
Multiplication of the estimates of oviposition preference and survival up to gall
=} ©
diameter of the gall (mm)
6.0
40
20 30 40 50 diameter of the shoot base (mm)
M. CHvÂLA, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 19
formation in one diameter class gives the probability that a shoot in that class will develop a gall. This appears to be highest on shoots with a diameter of 3 and 4 mm, and this estimate agrees with the fractions of galled shoots in each class found in the field. In the field a low number of galls is also found on shoots with a diameter of 7 or 8 mm and sometimes even more, so that the chance of gall formation cannot be zero as in the experiment of Fig. 19b. The experiments were of course, all carried out with a limited number of shoots, so that chance variation can be important.
This analytical approach explains why the highest density of galls can be found in
E 100
diameter of the gall (m
6.0
4.0
20 30 40 50 diameter of the shoot base (mm)
Fig. 20—21. Correlation between basal diameter of reed shoot and diameter of gall occurring on
that shoot, for different types of galls: lignified galls containing larvae of L. /ucens (LJ) and
L. rufitarsis (WA), and soft walled galls with larvae of L. rufitarsis (©) and L. pullitarsis (9).
Galls collected in Voorsterbos (Fig. 20: parcel T 35, Fig. 21 parcel S 18), N.O. Polder, The Netherlands 11-1-1963
20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974
places where a large proportion of the reed shoots have a width of about 3 or 4 mm.
It may be concluded that most galls of Lipara lucens are found in reed with sub- optimal growth, but not in reed with the worst growth. Although the water level has no direct influence, it can be said that reed with suboptimal growth is found mainly in dry places. On the basis of experience with a wide variety of reed vegetations in the Netherlands it can be said that there is no predilection for reed growing in extensive vegetations, as compared to reed in smaller patches, for instance in small bogs, along ditches, etc.
Lipara rufitarsis and especially L. pullitarsis are not as abundant in the Netherlands as L. lucens. Only superficial observations on their habitat are available, but comparison with L. lacens is possible. In reed vegetations where two or three species are present the galls of L. rufitarsis and L. lucens occur on shoots with about the same diameter, but th shoots with galls of L. pullitarsis on the average have a larger diameter. This can be seen, for instance, in Fig. 20, but has also been found in many other vegetations.
Galls of L. rufitarsis are sometimes present in low numbers in closed reed vegetations on dry soil, together with L. lwcens, but occur much more abundantly in smaller reed vegetations in dry places at the edge of woods or along roadsides or other somewhat shaded places. In such places they are often more numerous than L. lucens.
Galls of L. pullitarsis were found in small reed stands with a low shoot density (often located between trees or bushes) in wetter, marshier places than those where L. rufitarsis has its optimum. In the Netherlands no galls of pullitarsis were found in large closed reed vegetations. This shows clearly that shoot dimension is not the only important factor determining the habitat of Lipara species.
In places where L. rufitarsis and L.lucens occur together, the larvae of the former species are sometimes found in galls that are indistinguishable from those of the latter species. This peculiar phenomenon has already been reported by Ruppolt (1957), but he did not describe L. rufitarsis in its typical galls. The thin galls with larvae he describes under the name of L. rufitarsis, undoubtedly refer to L. pullitarsis.
The two types of galls in which rwfitarsis is found are quite different. That no inter- mediate forms occur is evident, e.g. in Figs. 20 and 21, showing two distinct groups of dots. The upper group corresponds with the wide galls with thickened internodes and lignified walls, the lower group, with narrow galls of which the internodes are not thickened. For both types of gall there is a strong positive correlation between shoot and gall width illustrating that the diameter and other dimensions of a gall are significant only in relation to shoot diameter.
Analysis of the morphology of the larvae and adults and also of the acoustical signals of the male flies, proved that the same species (rufitarsis) occurs in both types of gall. A few galls of the /ucens type have also been found in which only a pullitarsis larva was present (above the growing point).
It is difficult to believe that one species can induce different galls on the same food plant. We know that when a shoot is infected by more than one egg, never more than one larva will survive. When these eggs are of different species, it cannot be said beforehand which species will survive. In the case of multiple infection by /zcens and by one of the smaller species, if the /ucens larva dies the result will be a typical /ucens gall with a surviving rufitarsis or pullitarsis larva. This only holds, however, if the /ucens larva does not die before the gall reaches maturity, since otherwise we would find a gall of intermediate size. We know that gall growth stops when a /zcens larva dies before
M. CHVALA, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 21
gall maturation, because we find rudiments of galls above which the shoot has resumed normal growth. This does not imply, however, that /zcens will always succumb to the smaller species. Without extensive breeding experiments, it will be impossible to judge the outcome of interspecific competition.
b. Parasites
Hymenopterous parasites belonging to four families attack the different species of Lipara. We have not tried to give a comprehensive list of these parasites. Only the common species with which we are acquainted will be treated here, because the records of occasional parasites are difficult to check. It is important to rear from material found in opened galls, to be sure that the parasites originate from Lipara larvae and not from the numerous species of inquilines that are also present in the galls.
The occurence of parasites differs widely between the types of galls. Table II shows the numbers of hosts and parasites found in galls collected in a number of localities in the central and eastern part of the Netherlands. The /zcens galls included here originate exclusively from places where this species occurred together with one or both of the other fly species. More figures on parasitism of L./ucens have been given elswhere (Mook, 1967).
The findings of Table II cannot be generalized for other regions. The relative im- portance of the parasites in different regions is indicated below under the different species.
Table II. Distribution of host and parasite larvae over 3 types of galls collected in various places in the Netherlands
Type of gall
Species of larvae lucens rufitarsis pullitarsis Lipara lucens 191 — — Lipara rufitarsis 31 240 — Lipara pullitarsis 9 — 119 Polemon liparae 83 me ae Stenomalina liparae 58 8 13 Scambus phragmitidis 1 1113 4 Total 373 361 136
Ichneumonidae: From Table II it is apparent that Scambus phragmitidis Perk. is primarily a parasite of L. rufitarsis. Waitzbauer (1969) reared one specimen from /ucens in Austria (Neusiedler See). In the Netherlands we have found a few in galls of /ucens, but always in places where rufitarsis also occurred in /ucens galls. This parasite was erroneously named Pimpla arundinator F. var. similis Bridgm. in Mook (1967). The new identification was made by Mr. K. W. R. Zwart, Wageningen, who compared specimens with paratypes from the British Museum (Natural History), London.
The distribution is difficult to assess. Pokorny did not find Scambus in Czechoslovakia. Other species of the same genus have been mentioned in the literature: Giraud (1863) reared Scambus (= Pimpla) arundinator F. as a parasite of L. rufitarsis or L. pullitarsis in Austria, while Wagner (1907) in Germany (Hamburg) and Blair (1944b) in England mention Scambus (= Pimpla) detritus Holmgr. from galls of L. /ucens.
22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 Cnemopogon apicalis 5 demen 10. Scambus phragmitidis | À 0 ERK
is Lipara pullitarsis
ol
Polemon liparae
number of specimens Ui oO
(©)
= Lipara lucens x PA
Lipara rufitarsis
Stenomalina liparae
5 chr i li
24 26 28 30 2 4 6 8 10 12 14 16 18 20 22 24 26 28 APRIL ~ MAY
Fig. 22. Emergence in the laboratory of adult flies, predators and parasites from galls collected in N.O. Polder and near Dalfsen, The Netherlands, 20-IV-1962
M. CHVALA, J. DoskoCr, J. H. Mook & V. Pokorny: The genus Lipara 23
Of the life history of Scambus phragmitidis nothing more is known than that it emerges early in the season (see Fig. 22), long before L. rufitarsis. The oviposition behaviour has not been studied.
Braconidae: Two species of Polemon have been described: P. liparae Gir. and P. melas Gir. In the Netherlands P. liparae is primarily a parasite of L. lucens. In Austria (Waitz- bauer, 1969) and Czechoslovakia (Pokorny) it is by far the most important parasite of L. lucens, while in Hungary it was the only parasite found in this host (Erdös, 1961). Giraud (1863) mentions this species as the only parasite of Lipara similis.
Polemon melas is the most important parasite of L.rufitarsis in Czechoslovakia (Pokorny). Giraud (1863) found it in Austria. It was reared occasionally in the Nether- lands (Reynvaan and Docters van Leeuwen, 1906).
Polemon liparae emerges from the tip of the gall just as its host and at the same time as L. lucens. It oviposits in the eggs of /ucens (Mook, 1961). Larvae of lwcens parasitized by Polemon form a puparium in September instead of in the spring, as they normally do. A puparium containing a parasite can be distinguished from the normal by being reddish-brown and slightly more slender. Varley and Butler (1933, see also Askew, 1971) have suggested that the premature pupation might be caused by wounding by the ovipositing parasite. This is improbable because of the long time lying between infection and pupation.
Pteromalidae. Stenomalina (= Pteromalus) liparae Gir. was found in the Netherlands in galls of lucens, rufitarsis and pullitarsis (see Table II), but most commonly in /zcens. In England (Blair, 1932) and western Germany (Hamburg, Wagner, 1907) it is the most common parasite of /wcens. In Czechoslovakia it is not common (Pokorny); it was not found in eastern Austria (Neusiedler See) by Waitzbauer (1969) and in Hungary by Erdös (1961). Giraud (1863) has described it from material collected near Vienna and indicates that it was rather common there.
The adults of Stenomalina liparae leave the gall by biting a small round hole in the leaf sheaths just above the gall chamber. They begin to emerge about 8—10 days later than L. lucens. They parasitize the young larvae living above the growing point, by boring through the leaf sheaths. In view of the circumstance that the first larvae of /zcens arrive above the growing point in the period that Stexomalina starts to emerge and that this species does not need much time between emergence and oviposition, it seems that its life cycle is primarily synchronized to that of L. /ucens.
Eulophidae. Tetrastichus legionarius Gir. is mentioned as a parasite of L. /ucens in Czechoslovakia (Pokorny), Austria (Giraud, 1863), and the DDR (Greifswald, Ruppolt, 1957), but seems nowhere to be common. Erdös (1961) mentions the species from Hungary as a parasite of Haplegis flavitarsis Mg., a small chloropid fly that is an in- quiline in Lipara galls. In the Netherlands 7. legionarius is rare in lucens. We have found only recently two parasitized puparia, after having opened more than 10.000 galls from various places, although mainly from the eastern part of the country. Mr. M. J. Gijswijt ('s-Graveland), who kindly identified our specimen, informed us that the species has also been reared from /wcens galls from three localities in the central and southern parts of the Netherlands. It is a gregarious parasite: our puparia contained 25 and 28 larvae respectively.
24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 1, 1974 c. Predators
Birds attack the galls of L.r#fitarsis in particular and also the smaller galls of L. lucens occurring on narrow shoots and, less often, the galls of L. pullitarsis. The intensity of this predation differs widely between winters. Blue Tits (Parus caeruleus L.) are known to predate in winter on all kinds of insects in reed stems (Tischler, 1943). Actual observations of predating tits are scarce, but they are at least known to be able to open the galls (Mook, 1967). Kramer (1917) observed a Great Spotted Woodpecker (Dryobatus major L.), opening a gall of L. lucens.
Puparia of Cnemopogon apicalis Wiedemann (Diptera, Scatophagidae) are found in the winter in small numbers between the top blades of all three types of galls. The host larvae are always absent, presumably having been consumed by the Cremopogon larvae (Wagner, 1907; Theowald, 1961).
6. AKNOWLEDGEMENTS
We are indebted to Miss C. G. Bruggemann (Arnhem) for doing much of the work on the acoustical behaviour, to Dr. H. PeSkova (Praha) for her generous help in the investigation of the immature stages, and to Mrs. J. A. van Noort and Mr. D. Landsman (Arnhem) for assistance in collecting data for sections 4 and 5. Thanks are also due to Dr. J. Chalupsky (Praha) for making the photographs of Plates 2 and 3, to Prof. Dr. L. M. Schoonhoven (Wageningen) for making the UV recordings of Plate 7 and to Mr. W.J. N. M. Verholt for drawing Fig. 19—22.
7. REFERENCES
Askew, R. R., 1971, Parasitic insects. Heinemann, London, 316 pp.
Balachowsky, A. et L. Mesnil, 1935, Les insectes nuisibles aux Plantes cultivées. Vol. 1, Paris, 1137 pp.
Becker, T., 1910, Chloropidae. Eine monographische Studie I. Arch. Zool. 10: 33—174.
Blair, K. G., 1932, Some notes on the galls of Lipara lucens. Entomologist’s mon. Mag. London.
68: 10—13.
, 1944a, Further notes on the galls of Lipara lucens Mg. Entomologist’s mon. Mag. 80: 6—7.
, 1944b, Some galls of Lipara lucens Mg. Entomologist's mon. Mag. 80: 189—190.
Collin, J. E., 1946, The British Genera and Species of Oscinellinae (Diptera, Chloropidae). Trans. R. Ent. Soc. Lond., 97: 117—148.
Docters van Leeuwen, W. M., 1957, Gallenboek, 2 nd. ed. Zutphen: Thieme. 332 pp.
Doskoäil, J. et Chvála, M., 1971, A revision of Lipara Meigen (Diptera, Chloropidae), including the description of a new species from Europe. Acta ent. bohemoslov., 68: 100—107.
Duda, O., 1933, Chloropidae, in: E. Lindner Ed., Die Fliegen der palaearktischen Region. Stuttgart. 248 pp.
Enderlein, G., 1913, Analytische Tabelle der Gattungen der Chloropiden-Subfamilie Oscinellinae.
Zool. Anz. 38: 10—13.
, 1961, Beobachtungen über die Insektencönose des Schilfes. 8. Wanderversammlung
Deutscher Entomologen, 171—177.
Giraud, J., 1863, Mémoire sur les insectes qui vivent sur le Roseau Commun, Phragmites communis Trin. et plus spécialement sur ceux de l'ordre des hyménoptéres. Verh. zool. bot. Ges. Wien 13: 1251—1288.
Heeger, E., 1856, Neue Metamorphosen einiger Dipteren, Sber. K. Akad. Wiss. Wien. 20: 335— 345.
Hennig, W., 1952, Die Larvenformen der Dipteren. 3 Teil, Berlin: Akad.-Verlag, 628 pp.
Erdos, J.
M. CiivALa, J. Doskocit, J. H. Mook & V. POKORNY : The genus Lipara 25
Kramer, H., 1917, Die Musciden der Oberlausitz. Abh. Naturforsch. Ges. Görlitz. 28: 334— 338. Loew H., 1858, Zwanzig neue Diptern. Wien. Ent. Monatschr., 2: 57—62. De Meijere, J. C. H., 1949, Achtste supplement op de nieuwe naamlijst van Nederlandse diptera van 1898. Tijdschr. Ent. 92: 1—14. Mook, J. H., 1961, Observations on the oviposition behaviour of Polemon liparae Gir. (Hym., Braconidae). Archs. néerl. Zool. 14: 423—430. — , 1967, Habitat selection by Lipara lucens Mg. (Diptera, Chloropidae) and its survival value. Archs. néerl. Zool. 17: 469—549.
, and C. G. Bruggemann, 1968, Acoustical communication by Lipara lucens (Diptera, Chlo- ropidae). Entomologia exp. appl. 11: 397—402.
, and C. G. Bruggemann, 1971, Acoustical behaviour of Chloropid flies living on reed (Phragmites communis Trin.). Proceedings XIII int. Congress of Entomology, I; 418—419.
Oldroyd, H., 1964, The natural history of flies. Weidenfeld and Nicholson, London.
Pokorny, V., 1970a, Bionomie druhu Lipara lucens Meig. a L. similis Schin. na räkosu. Ziva, Praha, 3: 101-103.
—., 1970b, The influencing of reed (Phragmites communis Trin.) by gall forming Diptera of the genus Lipara Meig. PT - PP Report No. 1 (1964—1969) Productivity of terrestrial ecosystems, Praha, 135—136.
——, 1971, Flies of the genus Lipara Meigen on Common Reed, Hidrobiologia, Bucarest, 12:
287—292. , 1972, Bionomie druhu Lipara rufitarsis Loew. a L. pullitarsis Dosk. et Chvála na rákosu. Ziva, Praha (in press).
Reynvaan, J. and W. M. Docters van Leeuwen, 1906, Die Entwicklung der Galle von Lipara lucens. Recl. Trav. Bot. néerl. 12: 235--261.
Ruppolt, W., 1956, Uber die cecidogene Diptere Lipara lucens Meigen (Chloropidae). Naturwissen- schaften, 43: 260.
—., 1957, Zur Biologie der Cecidogenen Diptere Lipara lucens (Chloropidae), Wiss. Z. Ernst Moritz Arndt-Univ. Greifswald, Mat.-Naturwiss. Reihe, 6: 280—292.
Sabrosky, C. W., 1941, An Annotated List of Genotypes of the Chloropidae of the World. Ann.
ent. Soc. Am., 34: 735—765. , 1958, A Phragmites Gall-maker new to North America (Diptera, Chloropidae). Proc. ent. Soc. Wash., 60: 231.
Schiner, J. R., 1854, Dipterologische Fragmente IV. Verh. zool.-bot. Ges. Wien 4: 169—176.
Theowald, Br., 1961, Diptera uit de sigaargal van het riet. Ent. Ber., Amst. 21: 108—109.
Tischler, F., 1943, Schilfrohr als Nahrungsquelle für insekten-fressende Vögel im Winter. Vogelzug. 14: 69—71.
Varley, G. C. and C. G. Butler, 1933, The acceleration of development in insects by parasitism. Parasitology, 25: 263—268.
Vimmer, A., 1925, Larvy a kukly dvoukfidlého hmyzu stfedoevropsk&ho. Praha: Ceská graf. Unie, 348 pp.
Wagner, W., 1907, Uber die Galle der Lipara lucens Meig. Verh. Ver. naturw. Unterh. Hamb. 13: 120—135.
Waitzbauer, W., 1969 Lebensweise und Produktionsbiologie der Schilfgallenfliege Lipara lucens Mg. (Diptera, Chloropidae). Sber. dst. Akad. Wiss. Mathem.-Naturw. Kl., Abt. I, 178: 175—242.
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PYRALIDAE OF THE THIRD ARCHBOLD EXPEDITION. 1. SUBFAMILY ODONTIINAE (LEPIDOPTERA)
BY EUGENE MUNROE Ottawa, Canada*)
ABSTRACT
Nine species in six genera are recorded from the material of the Third Archbold Expedition to Western New Guinea and are figured. The following taxa are described as new: Trigo- noorda gen. nov., type-species T. triangularis spec. nov. also includiing T. iebelealis spec. nov; Pseudonoorda gen. nov., type-species P. minor spec. nov., also including P. brunneiflava spec. nov. and the previously described P. nigropunctalis (Hampson) comb. nov., P. distigmalis (Hampson) comb. nov. and P. metalloma (Lower) comb. nov., ex Noorda, and P. noordimimalis (Hampson) comb. nov., P. brunneifusalis (Hampson) comb. nov, P. photina (Tams) comb. nov. and P. lampra (Tams) comb. nov., ex Clupeosoma; Hyalinarcha gen. nov., type-species H. hyalina (Hampson) comb. nov., ex Boeotarcha. Thesaurica notodontalis (Hampson) comb. nov. and T. accensalis (Swinhoe) comb. nov. are transferred from Sameodes and Noorda, respectively.
INTRODUCTION
The rich material of Pyralidae collected by the Netherlands Indian — American Exped- ition to the interior of Dutch New Guinea (now West Irian), better known as the Third Archbold Expedition, has been lent to me for study for a number of years, thanks to the generosity and sorely tried patience of the Rijksmuseum van Natuurlijke Historie, Leiden. During this time the whole of this material has been prepared and sorted to species and the problems of identity and relationship have for the most part been solved. I now propose to give lists of the material, subfamily by subfamily, with remarks on relation- ships and with descriptions of new taxa where required. A general faunistic analysis will be deferred until after the taxonomic treatment is complete, but the following state- ments will serve for introduction.
The collection was made mostly in relatively undisturbed habitats at a variety of elevations from near sea level to 3800 m. The localities are described in detail by Brass (1941), Archbold, Rand & Brass (1942) and Toxopeus (1940). The most intensive collecting was done at middle and higher altitudes, from which the species represen- tation is good. Many species were collected at lower levels also, but they probably represent a considerably smaller fraction of the fauna there. The fauna shows pronounced altitudinal zonation. In a general way this corresponds well to what is known from eastern New Guinea. The same groups: Scopariinae, certain Nymphulinae, etc., are well developed at high altitudes, with such groups as Epipaschiinae and certain Phycitinae and Pyraustinae becoming important in the 1500—2500 m zone. At both levels some species are known from eastern New Guinea, whereas others are known only from the
*) Biosystematics Research Institute, Canada Department of Agriculture, Central Experimental Farm, Ottawa, Ontario KIA OC6, Canada. 27
28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974
present collections. At the lower levels, especially Araucaria Camp at 800 m, at which a large collection was made, the fauna is generally similar to that in other parts of New Guinea.
The high-altitude material is the most extensive so far collected in New Guinea, but the whole collection is of very great interest. In the main it reflects the hard and careful work of the late L. J. Toxopeus, whose untimely death in 1951 interrupted a brilliant entomological career.
This first paper deals with the subfamily Odontiinae. Although the subfamily has a number of genera and species in New Guinea, the representation in the present collection is poor. I can only speculate as to the reasons, but there are probably relatively few species at high altitudes, while at moderate and low altitudes they may have been eclipsed by the many more conspicuous and exciting species of other groups.
SUBFAMILY ODONTIINAE
Clupeosoma Snellen Clupeosoma Snellen, 1880: 203.
Type-species C. pellucidalis [recte, pellucidale} Snellen, 1880, Celebes, by monotypy.
This genus was used by Hampson, 1897: 216, for a rather mixed assemblage of odontiine and pyraustine species, which he placed in the Hydrocampinae (= Nymphu- linae) because of the stalking of R, with Rz 4 4. In my “Synopsis of the North American Odontiinae” (Munroe, 1961: 5) I referred Clupeosoma to the Odontiinae without dis- cussion. As will be seen from the figure, the genitalia are typically odontiine, with bilobed setose uncus, with gnathos arms fused basally with tegumen, with strong coremata associated with the juxta, and with rounded, radially fluted valves.
I restrict the genus to a series of species closely related to C. pellucidale, including C. astrigale Hampson, emend., from Queensland; C. subrufale (Rothschild), emend., comb. nov., from Ceram, described in Noorda; C. atristriatum Hampson, emend., from Rookfe] Island; C. rufistriatum Hampson, emend., from the Oetakwa River, Snow Mountains, New Guinea; C. glaucinale Hampson, emend., from Sumbawa; and C. la- niferale Hampson, emend., from St. Aignan, as well as a number of undescribed species. Snellen’s characterization of the genus is good, and will serve to distinguish it from the discordant elements placed in it by Hampson. The maculation is characteristic and is shared only by the closely related genus Neogenesis Hampson, discussed below.
Clupeosoma rufistriatum Hampson Figs. 1, 10 Clupeosoma rufistriatum {sic} Hampson, 1917: 277. The single specimen of the genus in the present collection appears to belong to
Hampson’s species, described from the Oetakwa River at 3.000 ft [ca. 900 m]. 1g, Araucaria Camp, 800 m, 24 March 1939, L. J. Toxopeus.
Neogenesis Hampson
Neogenesis Hampson, 1907: 19. Type-species N. flaviplagialis Hampson, from New Guinea, by monotypy. As Hampson noted, this genus has clearly arisen from Clupeosoma by exaggeration
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 29
of the secondary sexual characters and by stalking and fusion of veins in both sexes. The single known species is widely distributed in New Guinea.
Neogenesis flaviplagialis Hampson Figs. 2, 11, 19
Neogenesis flaviplagialis Hampson, 1907: 19.
The species should not be mistaken for anything else. It was described from material from Milne Bay (type locality) and Moroka, British New Guinea [now Papua}. Meek collected it in the Snow Mountains and George Holland and I found it at several localities in eastern New Guinea. I can see no significant geographical variation.
1g, Hollandia, 1938, Brinkmann; 14, Araucaria Camp, 800 m, 22 March 1939, L. J. Toxopeus; 1g°, Mist Camp, 1800 m, 15 Jan. 1939, L. J. Toxopeus.
Trigonoorda gen. nov.
Type-species Trigonoorda triangularis spec. nov.
External characters. Frons slightly prominent, either rounded or flat, smoothly scaled. Vertex short, with rough, erect scaling. Labial palpus porrect, exceeding frons by about dength of head, scaling flattened, blade-like, third segment largely hidden in scaling of second. Maxillary palpus prominent, with an expanded distal tuft of scales. Proboscis well developed, scaled at base. Eye large. Ocellus present but small. Antenna filiform, in male with feathery ciliations, several times as long as diameter of shaft; in female with short even ciliations; in both sexes smoothly scaled dorsally. Body short but not especially robust. Praecinctorium prominent, with transversely rounded distal tuft. Legs slender, without unusual specializations.
Forewing broadly triangular; costa and termen somewhat convex; apex acute; posterior margin sub-basally convex and with a weak scale tuft. Cell rather narrow, a, little over half of length of wing. R, from cell basad of anterior angle. R, stalked with R3 + 4, the latter long-stalked. R, from anterior angle of cell, weakly curved and approximated to R3 + 4 at base. Discocellular straight, somewhat oblique. M, from somewhat behind anterior angle of cell, diverging from R;. M, and M, from posterior angle of cell, their origins close but slightly separated, their basal parts not or scarcely curved and approx- mated. Cu, from well basad of posterior angle. Cu, from cell at two-thirds to three-fifths from base. Anal loop large, very weak distally.
Hind wing of moderate width, termen rounded. Sc and R, anastomosed. R, and M, stalked for some distance. Cell about half length of wing. Discocellular concave, posterior limb strongly oblique distad. M, and M, from posterior angle of cell, their basal parts usually weakly curved and approximated. Cu, from cell at two-thirds to three-fifths. Cubital and anal area with slender, weakly spatulate, hairlike scales.
Male genitalia. Uncus rather narrowly subtriangular, tip rounded, sides rolled down and weakly setose. Gnathos Y-shaped. Transtilla narrow, arched. Juxta large and pyriform. Vinculum ventrally rounded and expanded, bearing moderately developed submedial and lateral paired coremata. Valve of moderate width, ovoidal, with a sinuate ridge well below costa and with inflated sacculus; distal and costal parts radially fluted; a variably developed group of short spines on ventrodistal curvature at end of sacculus. Penis slender and curved, with fairly long basal caecum; armature of vesica weak or absent.
30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974
Female genitalia. Ovipositor with high narrow lobes bearing relatively few, variably developed setae. Apophyses moderate to long. Eighth tergite rectangular or with anterior margin medially produced, with scattered setae. Ductus bursae membranous, evenly narrow, of moderate length, with a sclerotized collar at junction with bursa. Bursa round or pyriform, membranous, with a small signum near junction of ductus bursae.
Remarks. This genus would key to Zebronia Hübner in Hampson’s Nymphulinae, but it has only certain superficial characters in common with that pyraustine genus. It is related in a general way to Clupeosoma and Autocharis. It differs from the former in the shorter palpi, broadly triangular wings and in characters of the wing venation, maculation and genitalia. From the latter it differs, inter alia, in the stalking of Ro with Rg + 4.
In addition to the two species described here there are several smaller undescribed species from New Guinea and the Sunda Archipelago, which will be dealt with else- where.
Trigonoorda triangularis spec. nov. Figs. 3, 12, 20
External characters. Frons somewhat rounded, dark reddish brown. Vertex contrastingly light buff. Maxillary palpus reddish brown. Basal scaling of proboscis buff. Eye and ocellus brownish fuscous. Antenna buff; dorsal scaling with fuscous segmental bands. Thorax above anteriorly reddish brown, posteriorly light buff. Abdomen above buff. Body beneath light buff. Legs whitish buff, with femora reddish brown and outer surfaces of front tibiae and tarsi broadly banded with reddish brown.
Forewing above somewhat thinly scaled, light, weakly pinkish buff with fine fulvous- buff dusting. Costa and termen narrowly dark reddish brown. Antemedial line narrow weak, almost straight; oblique distad from costa at one-third from base to posterior margin at two-fifths. Discocellular bar fine, fuscous, connected anteriorly with costal line. Postmedial line narrow, weak, slightly dentate, tending to be interrupted at veins, dark reddish brown or brownish fuscous; oblique distad from costa at four-fifths from base as far as middle of cell Mg, there angled and curved increasingly basad to just in front of Cuz, there obtusely angled and erect to posterior margin at three-fourths from base. Fringe light buff basally, white distally.
Hind wing above translucent pale yellowish buff. Fringe whitish buff.
Forewing beneath light pinkish buff, with silky lustre. Costal line as above. Terminal line more diffuse. Discocellular bar strong. Transverse lines visible only by transparency. Fringe as above.
Hind wing beneath as above.
Expanse 21 to 24 mm.
Male genitalia. Uncus less than three times as long as wide. Gnathos with median element tapering to a point distally. Dorsal part of juxta narrow. Patch of spines at ventrodistal curvature of valve strong.
Female genitalia. Ovipositor lobe high and narrow, with long setae. Apophyses short. Eighth tergite rectangular, with few fine setae, most numerous posterolaterally. Ostium membranous. Ductus bursae short but slender and of even width. Bursa round, signum near junction of ductus bursae.
Life history. Unknown.
Types. Holotype, g', allotype, 9, and 4 paratypes, Moss Forest Camp, 2600—2800
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 31
m, 11—24 Oct. 1939, L. J. Toxopeus. Holotype, allotype, and two paratypes in Leiden Museum; one male and one female paratype in Canadian National Collection, Type No. 12.861.
Remarks. This species closely resembles T. iebelealis but is slightly smaller, with fore- wings sharper and less warmly coloured and with finer, sharper markings; the hindwings are paler and less strongly yellowish in the present species. The genitalia differ in the two species in a number of characters as noted in the descriptions.
Trigonoorda iebelealis spec. nov. Figs. 4, 13, 21
External characters. Frons flat and oblique, dark brown medially, lighter towards sides but with a dark border. Vertex reddish brown anteriorly, yellowish buff posteriorly. A whitish-buff line between eye and antenna. Labial palpus with mixed fuscous and fer- rugineous scales, its base beneath narrowly but contrastingly white. Maxillary palpus fuscous; base of proboscis buff. Antenna light buff with segmental fuscous bands dorsally. Thorax above reddish brown anteriorly, shading to warm buff posteriorly. Abdomen above light greyish buff. Body beneath and legs whitish buff; outer surface of front legs broadly banded with purplish fuscous.
Forewing broadly triangular; costa and termen somewhat convex; apex subrectangular; termen slightly oblique, tornus rounded. Ground colour above warm yellowish buff, finely dusted with ferrugineous. Costa ferrugineous. Antemedial line weak, narrow, somewhat sinuated, ferrugineous; oblique distad from costa at one-third from base to posterior margin at three-fifths. A weak fuscous discocellular bar.
Postmedial line slightly oblique distad from costa at five-sixths to vein Mg, there weakly retracted to an obtuse angulation on Cu, and erect to termen near tornus. Termen with a narrow, somewhat diffuse, ferrugineous band. Fringe yellowish at base, whitish distally.
Hind wing above translucent yellowish buff, a little darker terminally. Faint traces of a reddish-brown postmedial line in cubito-anal area. Fringe as on forewing.
Wings beneath much as above. Forewing a little darker, with transverse lines lost and other dark markings reduced. Hind wing as above.
Expanse 23 to 25 mm.
Male genitalia. Uncus more than three times as long as wide. Gnathos with median element finger-like and not tapering. Dorsal part of juxta wider than in T. triangularis. Patch of spines at ventrodistal curvature of valve very weak.
Female genitalia. Ovipositor lobe high and narrow, but somewhat reduced and with few setae. Apophyses long and strong. Eighth tergite elongate and heavily sclerotized, produced forward medially and at anterolateral angles, its surface with scattered, fairly strong setae. Ductus bursae wider than in T. triangularis, pyriform and with signum far from junction of ductus bursae.
Life history. Unknown.
Types. Holotype, &, allotype, 9, and 1 9 paratype, TehelE Camp, 2250 m, 7 and 26 Nov. 1938, L. J. Toxopeus. 1 @ paratype, Sigi Camp, 1500 m, 23 Feb. 1939, L. J. Toxopeus. Holotype, allotype and one paratype in Leiden Museum; one paratype in Canadian National Collection, Type No. 12.862.
Remarks. This species is very similar in appearance to T. triangularis, but is a little
32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974
larger; the forewing is more orange and the hind wing is yellower; the transverse lines of the forewing are a little more diffuse; the antemedial line is more oblique and less regular. There are obvious differences in the male genitalia, especially the much weaker development in the present species of the spinose patch on the ventrodistal curvature of the valve.
Pseudonoorda gen. nov.
Type-species: Pseudonoorda minor spec. nov.
External characters. Frons flat and oblique, smoothly scaled. Vertex with rather smooth vestiture of raised and often broad scales, arising laterally and curving anteromesad. Labial palpus long and porrect, with flattened, blade-like, triangular scaling; third seg- ment wholly or partly hidden in scaling of second. Maxillary palpus prominent, with triangular distal scale-tuft. Proboscis well developed, scaled at base. Eye large. Ocellus present. Antenna filiform, ventrally short-pilose in both sexes, dorsally scaled. Body robust, abdomen slightly exceeding anal angle of hind wing. Praecinctorium with transverse, rounded, distal fan of scales. Legs slender, outer tibial spurs somewhat shorter than inner.
Forewing triangular, costa and termen weakly convex, posterior margin almost straight. Cell about half as long as wing, of moderate width. Discocellular erect anteriorly, curved increasingly distad, posterior angle of cell moderately acute. R, from near end of cell, closely apposed to Ro 4. Ry stalked for a considerable distance with R; + 4. Rz and R, stalked to near apex. R; from anterior angle of cell, its basal part curved and approx- imated to Rs 4 4. M; from anterior angle of cell, but straight and diverging from base of Rs. Mo, Mg, and Cu, from posterior angle of cell, their basal parts curved and approximated. Cu, from cell somewhat basad of end. Ist A absent. 2nd A strong. Anal loop large and complete.
Hind wing with Sc and R, anastomosed for some distance. Sc and M, separate or short-stalked. Cell less than half length of wing. Discocellular weak, its anterior part concave, its posterior limb strongly oblique. Mo, M, and Cu, from posterior angle of cell; M, and M, with basal parts curved and strongly approximated; basal part of Cu, weakly curved and slightly approximated to that of Mg. Cu, from cell at about two-thirds from base. Three anals present.
Male genitalia. Uncus subtriangular, bilobed, not or weakly setose. Gnathos with lateral arms arising from tegumen, median element spike-like, with weak terminal denticles. Transtilla angled dorsad medially. Juxta large and weakly sclerotized, flanked by coremata bearing strong, specialized scales. Vinculum ventrally broadly rounded, medially expanded. Valve oval, with complex basal sclerotization and strong subcostal ridge; distal part fluted and sometimes with somewhat specialized terminal setae.
Female genitalia. Ovipositor with elongate sparsely setose lobes. Posterior apophysis with cross-bar set at a shallow angle to shaft but not expanded; shaft long and slender, but strong. Eighth segment long, slender, cylindrical; tergite more or less rectangular, its anterior margin slightly excised and strengthened. Anterior apophysis as long as posterior, with triangular sub-basal flange. Ostium unarmed. Ductus bursae slender, membranous, with a sclerotized collar near bursa. Bursa globular or oval, membranous, with unevenly distributed spinules.
Life history. Unknown, but the ovipositor appears adapted to inserting eggs into plant tissues or crevices. The larva of the closely related Decelia terrosalis Snellen bores in mango seeds.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 1
Plate 1. Fig. 1—9. New Guinea Odontiinae. 1, Clupeosoma rufistriatum Hampson, &; 2, Neogenesis flaviplagialis Hampson, 4; 3, Trigonoorda
triangularis spec. nov, holotype, & ; 4, Trigonoorda iebelealis spec. nov, paratype, à ; 5, Pseudonoorda minor spec. nov., holotype, & ; 6, Pseudonoor-
da nigropunctalis (Hampson), 6; 7, Pseudonoorda brunneiflava spec. nov, holotype, &; 8, Hyalinarcha hyalina (Hampson), &; 9, Thesaurica noto- dontalis (Hampson), 4
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 ge" —
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 2
Plate 2.
Fig. 10—12. New Guinea Odontiinae, & genitalia. 10, Clupeosoma rufistriatum Hampson; 11, Neogenesis flaviplagialis Hampson; 12, Trigonoorda triangularis spec. nov.
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 3
Plate 3. Fig. 13—15. New Guinea Odontiinae, 4 genitalia 13, Trigonoorda iebelealis spec, nov.; 14.
Pseudonoorda minor spec. nov.; 15, Pseudonoorda nigropunctalis (Hampson)
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 PLAAT 4
Plate 4.
Fig. 16—18. New Guinea Odontiinae, 4 genitalia. 16, Pseudonoorda brunneiflava spec. nov.; 17, Hyalinarcha hyalina (Hampson); 18, Thesaurica notodontalis (Hampson)
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1
PLAAT 5
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974 Si sid AS bit
mr
N Vi - I È fe, \ jé;
| po | Fo $ \ sa i $ À ee È
A gii x + \ : "x
LÉ : / È
N \
Plate 5.
Fig. 19-22. New Guinea Odontiinae, 9 genitalia. 19, Neogenesis flaviplagialis Hampson; 20, Trigonoorda triangularis spec. nov.; 21, Trigonoorda iebelealis spec. nov.; 22, Pseudonoorda minor spec. nov.
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1
dann ns on
LE
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 33
Remarks. This genus contains a number of species which collectively range from Samoa through tropical Asia to Africa. They have been put in C/upeosoma in the Nymphulinae when authors noticed the stalking of R, with R34 4 or in Noorda when they did not. The most closely related named genera are: Decelia Snellen, unplaced by Hampson, though he put one of the nominal species in Noorda; Autocharis Swinhoe (see Amsel, 1965, 1970), whose species were also included by Hampson in Noorda; and Noordodes Hampson, placed by him in Endotrichinae. Decelza differs in having the third segment of the labial palpus clearly marked off from the second, not hidden in its scales, in having the tip of the forewing curved downward, in having the abdomen produced some distance beyond the anal angle of the hind wing, in having specialized scale-tufts on the front and middle legs of the male, in having the ovipositor lobes and eighth tergite of the females less elongate and in having the bursa copulatrix coiled. Autocharis has vein Ry of the forewing free, not stalked with R; + 4; the cell of the forewing is long and the discocellular is only weakly curved; in the hind wing veins M, and M, are stalked.
In addition to the type-species at least the following described species belong to this genus: Noorda distigmalis Hampson, from Africa, Clupeosoma noordimimalis Hampson, from Africa, Noorda nigropunctalis Hampson, from Perak; Clupeosoma brunneifusalis Hampson, from New Guinea, Clupeosoma photina Tams and C. lampra Tams, from Samoa; and Noorda metalloma Lower, from Australia and New Guinea, if the last species is correctly identified at the British Museum. All these become new combinations in Pseudonoorda.
Pseudonoorda minor spec. nov. Bigs!) > las 22
External characters. Frons fuscous, with a sharply defined, clay-coloured, postero- median triangle. Vertex ferrugineous. Labial palpus fuscous; base beneath contrastingly light grey. Eye and ocellus fuscous. Antenna dull buff. Thorax above ferrugineous anteriorly, shading to clay-colour posteriorly. Abdomen above brownish clay-colour, with posterior margins of segments light grey. Body beneath and legs grey, coxae, femora and front tibia infuscated.
Forewing above translucent greyish fuscous to postmedial line, brown-tinged beyond. Costa narrowly dark fuscous. Weak traces of an outwardly oblique, diffuse, dark-fuscous antemedial line. A weak fuscous streak on discocellular, strongest at anterior and posterior angles of cell. Postmedial line fine, fuscous, somewhat irregular, denticulate basad on veins, the denticulations somewhat accentuated. A diffuse, inconspicuous, dark terminal line, with faint dark rays extending a short distance basad on and between veins. Fringe fuscous, with a yellowish-buff line at extreme base, followed by a darker line.
Hind wing with a wedge from M to anal fold coloured like forewing; costal and anal areas pale translucent yellowish grey. No antemedial or discal markings. Postmedial line obsolescent anteriorly, generally straight but slightly denticulate from M, to Ist A, there nearly touching termen and ending abruptly. Fringe as on forewing.
Wings beneath as on upperside but shinier, less strongly marked and with terminal area hardly differentiated in colour.
Expanse 18 mm.
Male genitalia. Uncus about 21/ times as long as wide. Gnathos with median element rod-like, somewhat decurved and spinose at tip. Marginal scales of corema flanking juxta rather slender, in strong bunches. Valve irregular in shape, both dorsal and ventral
34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974
margins somewhat incised before tip; subcostal ridge sinuous, rather weak; fluting strong. Penis rather thick, with an imbricated group of small spine-like cornuti.
Female genitalia. As described for the genus.
Life history. Unknown.
Types. Holotype, g', and 1 & paratype, Mist Camp, 1600 m, 18 Jan. 1939, L. J. Toxopeus. Allotype, 9, same locality and collector, 11 Jan. 1939. 1 g' paratype, Bernhard Camp B, 100 m, 10 April 1939, L. J. Toxopeus. Holotype, allotype and one paratype in Rijksmuseum van Natuurlijke Historie, Leiden; one paratype in Canadian National Collection, Type No. 12.863.
Remarks. This species closely resembles P. brunneifusalis (Hampson), from the Oetakwa River, Snow Mountains, but is appreciably smaller (18 as against 22 mm) and much paler than the holotype of N. brunneifusalis. I think it better to classify the present series as a distinct species.
Pseudonoorda nigropunctalis (Hampson) comb. nov. Figs. 6, 15
Noorda nigropunctalis Hampson, 1899: 222.
The present collection contains two specimens and the Canadian National Collection | has one from Aiyura, Eastern Highlands, New Guinea, ca. 1850 m, which agree well with Hampson's types in spite of the different provenance of the latter.
1 2 (without abdomen), Araucaria Camp, 800 m, 21 March 1939, L. J. Toxopeus; 1 g, Iebele Camp, 2250 m, 28 Nov. 1938, L. J. Toxopeus.
Pseudonoorda brunneiflava spec. nov. Eps. 16
External characters. Frons yellow to yellowish buff, with black anterolateral wedges. Vertex yellow or yellowish buff. Labial palpus black, white at base beneath. Maxillary palpus black. Basal scaling of proboscis white. Eye and ocellus fuscous. Antenna buff, dorsally with dark-brown segmental stripes. A fuscous stripe from eye to wing-base. Thorax above yellow.
Abdomen above light orange-buff, with posterior margins of segments yellow. Body beneath and legs whitish buff; front and middle tibiae with weak fuscous bands.
Forewing above yellow. Costa to cell contrastingly fuscous with blue metallic lustre. Black antemedial dot, orbicular dash and comma-shaped discocellular bar, all adjoining costal stripe. Postmedial line faint, fuscous, denticulate inward with black accentuation on veins, oblique distad from costal stripe to cell R;, thence regularly convex to termen before tornus. Whole area beyond postmedial line evenly copper brown with purple lustre, except slightly paler just beyond postmedial line anteriorly. A row of fuscous terminal dots between veins. Fringe shining fuscous with a slightly darker line just basad of middle.
Hind wing above coloured like forewing, but with costal area whitish and its maculation obsolescent. No markings basad of postmedial line. The latter almost straight from M, to Ist A, there ending, but succeeded by a weak dark terminal smudge in anal area. Fringe as on forewing, but becoming paler toward anal area.
Wings beneath as above but somewhat suffused with brownish and with dark areas considerably less contrasting.
Expanse 15 to 17 mm.
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 35
Male genitalia. Uncus about twice as long as wide. Gnathos with median element tapering somewhat irregularly from base to narrowly rounded tip. Scale tufts of corema flanking juxta narrow. Valve broadly and rather evenly rounded; subcostal ridge straight; fluting weak. Penis straight and weakly sclerotized.
Female genitalia. Unknown.
Life history. Unknown.
Types. Holotype, &,and 1 g° paratype, Araucaria Camp, 800 m, 17 and 19 March 1939, L. J. Toxopeus. Holotype in Rijksmuseum van Natuurlijke Historie, Leiden; paratype in Canadian National Collection, type no. 12.864.
Remarks. As will be seen from the figures, the genitalia of this species differ con- siderably from those of P. #igropunctalis, especially in the shape of the valve. A species identified as P. metalloma (Lower) in the British Museum (Natural History) and represented in the Canadian National Collection from Padwe, Sepik District, New Guinea, is somewhat similar, but has the terminal area purplish fuscous and the post- medial line of the forewing evenly curved. Another, apparently undescribed, species from Papua and New Guinea has the postmedial line differently shaped but otherwise is very close to P. brunneiflava.
Hyalinarcha gen. nov.
Type-species Boeotarcha hyalina Hampson.
External characters. Frons flat and oblique, smoothly scaled, somewhat produced, but not conical. Vertex short, with rather small erect scales. Behind occiput a collar of large, wide, smooth scales, probably arising from neck. Labial palpus weakly decumbent, with broad, compressed, distally pointed scaling, exceeding frons by somewhat more than length of cranium, the segments with their scaling not visibly divided. Maxillary palpus prominent, with triangularly dilated distal tuft of scales. Proboscis moderately well developed, scaled at base. Eye large. Ocellus well developed, separated by a considerable distance from eye. Chaetosema marked by a tuft of slender scales. Antenna of male filiform, dorsally scaled, ventrally fasciculate. Body rather short and thick. Legs mod- erately slender, outer tibial spurs of male about half length of inner. Praecinctorium with simple, transverse, rounded distal scale tuft.
Male genitalia. Uncus flattened, pyriform, the sides downcurved and more heavily setose than the median part. Gnathos with wide lateral arms and heavy, somewhat trough- shaped median piece, the latter shorter than uncus and with rows of spines on dorsal surface distally. Juxta large, pyriform, dorsally rounded. Vinculum shallow, ventrally broadly flattened, bearing a moderately developed scale-like structure on each side, flanked mesally and ectally by narrower striated sclerites. Valve of moderate size, weakly fluted; a very strong subcostal sclerotized bar, ending in a prominent, free, spine-like process extending beyond body of valve; dorsad of this a weak zone of setae on distal part of costa; sacculus broadly sclerotized, bounded dorsally by a weak ridge. Penis thick, curved, with spirally pleated vesica and with a group of very heavily sclerotized spine-like cornuti, large, few and in a single row distally, becoming small, numerous and multiserial basally.
Female genitalia. Unknown.
Life history. Unknown.
Remarks. Although at present I propose this genus for the type-species only, there are additional closely related species, which may have to be placed here when the group
36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 2, 1974
has been studied more comprehensively. The type-species is recorded only from West Irian, but its relatives range through the Indo-Papuan region.
Hyalinarcha hyalina (Hampson) comb. nov. Figs. 8, 17
Boeotarcha hyalina Hampson, 1913: 527.
This species is represented in the present collection by a single male, which exactly matches the type male from the Mimika River in the British Museum (Natural History). Though Hampson may have meant to designate a holotype, his citation is ambiguous, reading “2 g' type”. I therefore designate as lectotype the specimen in the British Museum (Natural History) with the label “Boeotarcha hyalina type g'. Hmpsn”.
1 6, Hollandia, Brinkmann.
Thesaurica Turner
Thesaurica Turner, 1915: 49.
Type-species Sameodes argentifera Hampson, 1913, from Queensland, by monotypy and original designation.
This genus was erected for a single Australian species, but Hampson correctly asso- ciated with this in the British Museum collection the additional species Sameodes noto- dontalis Hampson, from Sandakan, Borneo, and Noorda accensalis Swinhoe, from Thai- land. The generic transfers appear not to have been published, and these species ac- cordingly become Thesaurica notodontalis (Hampson) comb. nov, and Thesaurica accensalis (Swinhoe) comb. nov. One of these is represented in the present collection.
Thesaurica notodontalis (Hampson) Figs. 9, 18
Sameodes notodontalis Hampson, 1899: 175.
Epipagis notodontalis: Klima, 1939: 316.
The exact identity of this species is uncertain, as topotypical material is not available at the time of writing and there are at least three superficially very similar species in New Guinea. The species represented in the present collection is at least closely similar to T. notodontalis. There are three specimens in the collection.
2 G', Bernhard Camp, 100 m, 10 and 12 April 1939, L. J. Toxopeus; 1 g Araucaria Camp, 800 m, 28 March 1939, L. J. Toxopeus.
ACKNOWLEDGEMENTS
My thanks are due to Dr. A. Diakonoff and to the authorities of the Rijksmuseum van Natuurlijke Historie, Leiden, for their patience in allowing me to retain this material for study for a very long period of time. It would have been impossible to work out the taxonomy without the continued support of the British Museum (Natural History) and especially the active help of Messrs. P. E. S. Whalley and Michael Shaffer. The photo- graphs were made by Messrs. Orin Hanright and Tom Stovell of the Bio-Graphic Unit, Research Branch, Canada Department of Agriculture, with the help of Mr. D. H. Kritsch. Mr. Kritsch also made preparations of genitalia and rendered other technical assistance.
E. MUNROE : Pyralidae of the 3rd Archbold Expedition. 1 37
LITERATURE
AMSEL, H. G., 1965. — Pyraliden-Studien 2. Die Noorda blitealis Wlk.-Gruppe (Lep.: Pyralidae).
— Stuttgarter Beitr. zur Naturk. 144: 1—7. , 1970. — Afghanische Pyraliden (Lepidoptera: Pyralidae). — Beitr. natürk. Forsch.
Südwestdeutschl. 29: 25—66.
ARCHBOLD, R., A. L. RAND, & L. J. BRASS, 1942. — Results of the Archbold Expeditions. No. 41, Summary of the 1938—39 New Guinea Expedition. — Bull. Amer. Mus. nat. Hist. 79: 197—288.
Brass, L. J., 1941. — The 1938—1939 expedition to the Snow Mountains, Netherlands New Guinea. — J. Arnold Arboretum 22: 271—342.
HAMPSON, Sir G. F., 1897.— On the classification of two subfamilies of moths of the family Pyralidae, the Hydrocampinae and Scoparianae. — Trans. ent. Soc. London 1897: 127—240.
—, 1899. — A revision of the moths of the subfamiliy Pyraustinae and family Pyralidae. Part II. — Proc. zool. Soc. London 1899: 172—291.
, 1907. — Descriptions of new Pyralidae of the subfamilies Hydrocampinae and Scoparianae [concluded}. — Ann. Mag. nat. Hist. (7) 19: 1—24. , 1913. — Descriptions of new species of Pyralidae of the subfamily Pyraustinae [con- tinued}. — Ann. Mag. nat. Hist. (8) 11: 322—342. —————,, 1917. — Descriptions of new Pyralidae of the subfamilies Hydrocampinae, Scoparia-
nae, &c. [concluded}. — Ann. Mag. nat. Hist. (8) 20: 265—282.
Kuma, A., 1939. — Pyralididae: Subfam. Pyraustinae II. — Lepidopterorum Catalogus. Pars 94.
MUNROE, E., 1961. — Synopsis of the North American Odontiinae, with descriptions of new genera and species (Lepidoptera: Pyralidae). — Can. Ent. Suppl. 24: 1—93.
SNELLEN, P. C. T., 1880. — Nieuwe Pyraliden op het eiland Celebes gevonden door Mr. M. C. Piepers. — Tijdschr. v. Ent. 23: 198—250.
Toxopeus, L. J. 1940. — Nederlandsch-Indisch-Amerikaansche expeditie naar Nederlandsch Nieuw Guinea (3e Archbold-Expeditie naar Nieuw Guinea 1938—1939): lijst van verzamel- stations. — Treubia 17: 271—279.
TURNER, A. J., 1915. — Studies in Australian Lepidoptera. — Proc. roy. Soc. Queensland 27: 11-57.
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| J. P. VAN LrrH. — Revision of the Psenini of the Ethiopian Region, including Malagasy a (Hymenoptera, Sphecidae, Pemphredoninae), p. 39—101, fig. 1—78.
| | Tijdschrift voor Entomologie, deel 117, afl. 3 Gepubliceerd 23-VIII-1974
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REVISION OF THE PSENINI OF THE ETHIOPIAN REGION, INCLUDING MALAGASY (HYMENOPTERA, SPHECIDAE, PEMPHREDONINAE) by J. P. VAN LITH
Rotterdam
ABSTRACT
The following new forms are described and illustrated: Psen (Psen) leclercgi (9); Psen (Mimesa) empeyi (4); Psenulus alveolatus (4), P. aurifasciatus (Q), P. bidentatus pallidus (9 and 8), P. freetownensis (2 and 3), P. jacoti (9 and &), P. leoninus (2), P. oweni (9 and 4), P. rugifrons (2), P. sapobaensis (2), P. uelleburgi (2). A supplementary description of earlier named species is given and males of some of these species are described for the first time. The subgeneric position of the species of Psen and the relationships of the species of Psenulus are discussed. Provisional keys and a list of Ethiopian species are presented.
A few years ago Dr. H. Townes, Ann Arbor, kindly sent me about sixty African Psenini, mainly from Sierra Leone, for identification. Usually material of this group is not found in great numbers. This relatively large collection therefore stimulated the undertaking of a new revision of the Ethiopian Psenini.
In 1961 Prof. J. Leclercq published his catalogue of the species of the Ethiopian region including Malagasy. In that paper of fundamental importance, he recorded four species of the subgenus Psen and twenty species of Psenulus, of which ten were newly de- scribed. A description of Psenulus garambae (Leclercq, 1961b) followed soon after- wards.
The study of nearly all types or paratypes of known species, along with some small fresh collections, enabled me to give supplementary descriptions, where necessary. Furthermore, eleven new species and one subspecies have been added and the systematic status of a few forms, regarded as subspecies by Leclercq, has been reconsidered. At present 41 species and subspecies of Psenini are known from the Ethiopian region.
In all about 450 specimens have now been recorded, the majority of which I had at my disposal. The material not seen by me includes a few large series of the commoner species, like a series of over sixty Psenulus capensis from Garamba. It is evident that the above total is far from being representative for this large area. Nevertheless we have obtained a somewhat better idea of its Psenine fauna. Some species have dispersed over large areas, consisting of savanna as well as forest regions. Psenulus capensis Brauns, for instance, is found from the extreme South of Africa to as far north as Sierra Leone and Ethiopia, with very little variation in structure or colour. Of Psenulus bidentatus (Came- ron) at least three subspecies can be distinguished. The nominate form occurs mainly in the savanna region, pallidus subsp. nov. in the forest region and the subspecies rabrocau- datus Turner in South Africa.
The subgenus Mimesa was already known from North Africa; the first representative of this subgenus of Psen south of the Sahara has now been found in Transvaal. The
39
40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
subgeneric status of a few species, thus far thought to belong to Psen s.str., is discussed at the end of the present paper, where also an attempt is made to recognize some higher groups of Psenulus.
The few data we have concerning the bionomics of the Ethiopian Psenini, their prey (cf. Psen silvaticus Arnold, Psen madecassus Arnold, Psenulus stevensoni Arnold and Psenulus capensis Brauns) or their nesting (cf. Psenulus patei Arnold and Psenulus capensis) agree with our present knowledge of the bionomics of their Palaearctic or Indo- Australian relatives.
For generously sending me their material I am much indebted to the authorities and staff of the institutions as well as to the private entomologists mentioned below, preceded by abbreviations used in the following text:
AMG — Albany Museum, Grahamstown, Cape Province, South Africa; Dr. F. W. Gess
AMNH — The American Museum of Natural History, New York, U.S.A.; Dr. J. G. Rozen and Mrs. M. Favreau
BM — British Museum (Natural History), London, United Kingdom; Dr. I. H. H. Yarrow and Mr. C. R. Vardy
CNC — Canadian National Collection of Insects, Ottawa, Canada; Dr. C. M. Yoshi- moto
EC — Collection H. N. Empey, Highlands North, Transvaal, South Africa
FAG — Faculté des Sciences Agronomiques de l'Etat, Gembloux, Belgium; Prof. J. Leclercq
HT — Collection H. Townes, Ann Arbor, U.S.A.
IRSNB — Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium; Dr. P. Dessart
MRAC — Musée Royal de l’Afrique Centrale, Tervuren, Belgium; Dr. J. Decelle
MS — Naturhistoriska Riksmuseet, Stockholm, Sweden; Dr. P. I. Persson
NMB — Naturhistorisches Museum, Basle, Switzerland; Dr. C. Baroni Urbani
OSUM — Oregon State University, Entomological Museum, Corvallis, U.S.A.; Dr.
G. R. Ferguson PMFV — Collection P. M. F. Verhoeff, Utrecht, The Netherlands
RM — National Museum of Rhodesia, Bulawayo, Rhodesia; Mr. F. C. de Moor
SAM — South African Museum, Cape Town, South Africa; Dr. A. J. Hesse
TM — The Transvaal Museum, Pretoria, Transvaal, South Africa; Dr. J. A. van Reenen
USNM — National Museum of Natural History, Smithsonian Institution, Washington D.C., U.S.A; Dr. K. V. Krombein, Dr. P. D. Hurd and Dr. A. S. Menke
ZMB — Zoologisches Museum der Humboldt-Universität, Berlin, D.D.R.; Dr. E. Königsmann
Special thanks are due to Prof. Jean Leclercq, Gembloux, and to Mr. F. C. de Moor, Bulawayo for their assistance. Mr. de Moor kindly also sent me some interesting un- published notes by Arnold. I am also much obliged to Dr. J. G. Betrem, Deventer, Drs. J. P. Duffels of the Institute of Taxonomic Zoology, Amsterdam, Mr. H. N. Empey, Transvaal, Dr. Per Inge Persson of the Natural History Museum, Stockholm, and to the South African Embassy, The Hague, for courteous aid in securing bibliographic, topo- graphic or other valuable information.
J. P. vAN LITH: Revision of Ethiopian Psenint 41
KEY TO THE SPECIES OF Psen S.L.
(the females of P. empeyi, P. patellatus, and the males of P. lecleregi and P. silvaticus
N
Ww
=
are still unknown)
Belles lode Ug PaO TEP Mn 2 MAES iur: TONE POE Pee) Mesopleura Reledingi pe il area at striato- nance) San densely, coarsely punctate, interstices narrower than punctures. Petiole dorsally smooth . 3 Mesopleura much finer and more sparsely punctate. Scutum coarsely punctate . 4
Propodeum finely reticulato-carinate, enclosed area ill-defined, more coarsely reticu- lato-carinate. Frons medially rugoso-punctate. Thorax black, pronotal tubercles and parts of pronotum red. Length 12 mm. Male unknown. Rhodesia :
Psen (Psen) silvaticus not peten sa ek carinate, ai area well-defined. Frons densely coarsely punctate, partly striato-punctate. Thorax largely black, pronotum, pronotal tubercles, anterior corners and narrow lateral margin of scutum, and upper two- thirds of anterior en of hora reddish. Length 11.5 mm. Male unknown. Malagasy . . . mu... Psen (Psen) leclercgi sp. nov. Frons coarsely striato- tate meg more distinctly and more densely punctate, interstices 3 or 4 times diameter of puncture. Petiole dorsally smooth. Pygidial area finely coriaceous with a few irregular punctures along sides. Thorax and gaster completely black, legs dark. Length 14 mm. Zaire . 5
si Psen (Psen) ao asta Eh ali stel ate deb finely and very sparsely, indistinctly, punctate. Scutum coarsely punctate, interstices usually at most as large as punctures. Petiole with fine latero-dorsal groove, in which a row of punctures, each with erect long hair. Pygidial area shining, with a few punctures along sides. Thorax including mesopleura largely dark reddish, gaster including petiole reddish-brown, tibiae reddish. Length 10— 11 mm. Malagasy . . Psen (subgenus?) madecassus Arnold Apical margins of third and fourth gastral sternites with fasciculate hairs. Scutum striato-punctate. Mesopleura distinctly punctate. Black, fore and mid tibiae brownish. Length 12—13 mm. Zaire . . . . + + . Psen (Psen) congolus Leclercq Apical margins of third and fone sternites without fasciculate hairs. Thorax largely dark reddish, or gaster partly reddish, or tarsi whitish. . . . . . . 6 Frontal carina distinct. Mesopleura including hypo-epimeral area finely, very sparsely, punctate. Petiole with fine latero-dorsal groove, in which row of punctures, each with erect long hair. Thorax including mesopleura largely dark reddish, tibiae reddish, i including petiole reddish-brown. Length 8.5—9.5 mm. Malagasy . 3 Psen (subgenus?) madecassus Arnold Frontal carina ln Hate densely punctate. Thorax black . . . . 7 Hypo-epimeral area dull, coarsely coriaceous. Mesopleura densely punctate, with narrow interstices. Scutum shining, densely punctate, interstices mostly larger than punctures. Antennae normal, gradually broadening towards apex. Legs normal. Ventral plate of petiole, first gastral tergite and base of second tergite reddish. Length 6.5 mm. Female unknown. South Africa . Psen (Mimesa) empeyi sp. nov.
— Hypo-epimeral area shining, with distinct, separate, punctures. Mesopleura densely
—
N
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
finely punctate, narrow interstices. Scutum shining, densely somewhat irregularly punctate, interstices sometimes larger than punctures. Apical half of flagellum broadened, segments much concave below. Fore femora, fore tibiae and especially fore basitarsi broadened. Gaster black, all basitarsi whitish. Length 7 mm. Female unknown. South Africa . . . . . . . Psen (subgenus?) patellatus Arnold
KEY TO THE FEMALES OF Psenulus (the females of P. alveolatus, P. avernus and P. trevirus are still unknown)
Tempora with coarse sculpture. Interantennal carina much broadened. Transverse carina below antennae. Prescutal sutures long, on apical half at least slightly indi- cated. Propodeal enclosure with wide central area. Petiole usually short, cylindrical, base flattened. No distinct pygidial area. Apical margins of sternites 4—5 with dense fringe of long pale hairs . . . . . . (Group of Psenulus turneri) 2 Tempora with much finer sculpture, in some species finely striate. Sternites 4—5 on apical margin with short pubescence, or with fringes of medium length (see
leoninus; veticulosas and sapobaensas). culi Pils © NE, ST, ADITO Second submarginal cell of fore wings petiolate. . . . u Second submarginal cell of fore wings not petiolate, if alan see aso turnert or paulisae . . 5 5
Apical part of fourth belgie Blas: ee zes NA (ad also So ui tergite. Fore legs reddish, mid legs more or less brown, hind legs black. Antennae dorsally reddish-brown. Frons obliquely striato-punctate, vertex punctate with faint striae. Scutum rather coarsely punctate with traces of transverse striae. Propodeum behind enclosure irregularly striate. Mesopleura strongly punctate with tendency to
striation. South Africa. . . . . twrneri Arnold Tergites 1—6 or 2—6 reddish. an Blackich don all At least base of hind tibiae reddish. Upper part of propodeum obliquely striate . . . . . . . 4
Frons below ocelli distinctly obliquely striato-punctate. Vertex shining, finely punctate. Clypeus distinctly bidentate. Mesopleura and scutum distinctly densely punctate with tendency to transverse striation. Fore femora yellowish-red, sometimes with brown streak behind; mid femora brown behind; hind femora black, hind tibiae dorsally brown with pale reddish base. Zaire, Sierra Leone . pazlisae Leclercq Frons finely densely punctate, slightly striate near median carina. Vertex shining, very finely punctate. Clypeal margin slightly emarginate, teeth less distinct. Scutum and mesopleura much finer and more sparsely punctate. Fore and mid legs and also hind tibiae yellowish-red; hind femora partly brown. Male unknown. Zaire, Sierra L'CONcH ARA «ll. | perusasiLecleteq Second racial cl i o wines Ario or Ron so (see also twrneri and paulisae); if slightly open, at least mid basitarsi pale yellowish. Upper part of propodeum striate. Frons more or less obliquely striato-punctate, or distinctly punctate. Tergites 1—2 or 1—3 black. . . . . step En Second submarginal cell distinctly open dorsally (see also: Par me saltitans). Frons striato-punctate or finely punctate. Gaster completely black or tergites 1—6 red, or upper part of propodeum coarsely reticulate. . . . aak 8 Apical margin of second tergite and tergites 3—6 completely ea Legs cu greater part of all femora dark brown, also back of mid tibiae and hind tibiae; mid
10.
bts
J. P. vAN LITH: Revision of Ethiopian Psenini 43
basitarsi yellowish-brown; hind tibiae with yellowish basal ring. Antennae slightly darkened dorsally. Mandibles yellowish-red. Vertex shining. Scutum shining, finely punctate, anteriorly somewhat striato-punctate. Mesopleura distinctly punctate, more sparsely than in paulisae, interstices smaller than punctures. Male unknown. Zaire
garambae Leclercq Tele Dig radish den most allo ek oF Ti ito) rar) bte Add ER
Second submarginal cell usually triangular or nearly so. Scutum shining, finely punctate, laterally with tendency to transverse striation. Tergites 4—6 reddish. Fore legs yellowish-red, underside of femora and trochanters brown; mid femora dark brown, underside of mid tibiae pale reddish-brown, basal fifth yellowish; hind femora and tibiae brownish-black, basal fourth of hind tibiae straw-yellow; all basitarsi pale yellowish, apex of hind basitarsi more or less darkened. Antennae dorsally slightly brownish. Male unknown. Rhodesia. . . . . . pate? Arnold Second submarginal cell not completely closed. Scutum shining, slightly stronger, but still finely punctate, with very little tendency to transverse striation. Apical margin of third tergite and whole tergites 4—6 reddish. Fore femora almost completely brown, fore tibiae yellowish-red, fore tarsi yellowish; mid femora and tibiae brown, mid basitarsus pale yellowish; hind legs brown, tibiae without distinct basal ring. Antennae brownish dorsally. Male unknown. Rhodesia . . . . saltitans Arnold
Gaster black, last tergite brownish. Legs black, fore tibiae, apex of fore femora and back of mid tibiae reddish-brown. Frons on either side of median carina convex, shining, very finely and sparsely punctate. Scutum shining, finely sparsely punctate. Prescutal sutures indistinct on posterior three-quarters part of scutum, indicated only by a line of punctures. Mesopleura distinctly punctate, interstices mostly larger than punctures. Upper part of back of propodeum finely obliquely striate. Larger species, length 8 mm. Petiole about one and one-quarter times as long as first tergite, in dorsal aspect. Male unknown. Sierra Leone . . . . . . aurifasciatus sp. nov. At least tergites 5—6 reddish . . . . a Bte Tergites 1—6 reddish. All legs including pira tok i, ini Antennae dark brown above; scape, last segment completely and rest of segments below yellowish-brown. Frons, scutum and mesopleura finely punctate. Upper part of propodeum smooth and shining. Petiole shorter than first tergite, in dorsal aspect. Sierra Leone, Equatorial Guinea . . . . freetownensis sp. nov. Only tergites 4—6 or 5—6 completely il ie lest mid and hind femora brown, fore tibiae and foreside of mid tibiae reddish, hind tibiae brown with yellowish-grey base. Antennae dorsally blackish-brown, scape and antennae below brownish. Frons above antennal sockets obliquely striato-rugose. Scutum and mesopleura striato- punctate. Upper part of propodeum coarsely reticulate, laterally somewhat obliquely
striate. Petiole about as long as first tergite. Zaire, Sierra Leone . . ealae Leclercq First gastral tergite not distinctly nodose; or thorax with Ka 5 marking and gaster reddish or partly reddish-brown . . Le Ze
First gastral tergite nodose. Frontal carina narrow bebe antennae. No sharp transverse carina below antennae, at most a low ridge. Anterior margin of clypeus blunt. Prescutal sutures short. Thorax and Di black. (Geo of Psenulus latian- mula) (ats: 4 30 Raised part of nanna carina more or re Binden sometimes not excavate.
44
12)
13
14.
19%
16.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
Prescutal sutures long (except in reticulosus), sometimes partly indistinct. Some species with reddish petiole and gaster . . Ana TZ Raised part of interantennal carina narrow Ge slightly ied in dilectus) between antennae. Prescutal sutures short. Never gaster including petiole reddish
Gaster and petiole reddish. Fore and mid legs whitish or yellowish. Distinct trans- verse carina below antennae . . . Larga DE) At least petiole black, if RES andy fie cate gite Ch Bess, legs not whitish or yellowish . . . . “nes o)
e part of carina between antennae eh aan ratio Sort length about
: 1.5. Prescutal sutures long but not sharply defined. Scutum shining, distinct, Ro punctation, sometimes with tendency to rugosity. Upper part of back of propodeum smooth. Petiole about one and one-third times as long as first tergite, in dorsal aspect. Pygidial area distinct, narrow. Pronotal dorsum, pronotal tubercles, hind margin of scutellum and metanotum ivory-white. Fore and mid legs including coxae ivory-white; hind = yellowish-red, hind trochanters more or less yellowish-
white. Malagasy . . . + . fulgidus Arnold Carina between antennae een iii sometimes Maltees: sharp. Petiole over one and a half times as long as first tergite, in dorsal aspect. . . . . . . 14
Prescutal sutures distinct only on anterior third of scutum. Scutum with distinct separate punctures. Propodeum behind enclosure smooth, back reticulate, above with oblique striae. Pygidial area narrow, with parallel unsharp lateral carinae. Pronotum yellow dorsally, in var. alboscutellatus Arnold also scutellum and metanotum yellow. Fore and mid legs including trochanters yellowish, hind legs reddish. Antennae reddish, base dorsally slightly darkened. Malagasy. . . . . reticulosus Arnold Prescutal sutures distinct on about two-fifths part of scutum, continued by a row of punctures. Scutum with variable, distinct separate punctures. Behind propodeal enclosure narrow smooth margin, back-side obliquely striate. No pygidial area. Mid tibiae on outer apex with longitudinal groove. Pronotum medially and metanotum somewhat brownish (discoloured?). Fore and mid legs including trochanters and mid coxae straw-yellow, hind legs including coxae reddish. Antennae dorsally dark brown, last segment above and flagellum below reddish-brown. Male unknown. INSERT IA NE STE sapobaensis sp. nov. Apex of iu Sini Sigi ans and Tone a longitudinal carina. Frontal carina between antennae only slightly broadened. Frons finely punctate. Distinct transverse carina below antennae. Prescutal sutures long. Back of propodeum smooth. Petiole about one and a half times as long as first tergite, in dorsal aspect. Petiole except ventral plate black, gaster reddish. Pronotal tubercles and tegulae yellowish- red. Fore and mid legs except back of femora and base of hind tibiae yellowish, hind femora and apex of hind tibiae brown. Zaire . . . . . benoiti Leclercq Pygidium without single longitudinal carina. . . NE EURO No transverse carina below antennae. Broadened part Fd raced tarten carina indistinctly closed below. Protruding median part of clypeal margin slightly emargi- nate, almost blunt. Frons and vertex densely finely punctate. Scutum densely finely punctate, slightly transversely rugose, prescutal sutures with short transverse striae and deep alveoles. Petiole at least one and a half times as long as first gastral tergite, in dorsal aspect. Pygidial area elongate-triangular. Gaster black. All tibiae and tarsi
197%
18.
19.
20.
21e
20%
J. P. van LITH: Revision of Ethiopian Psenini 45
whitish; fore and mid tibiae below and apical half of tibiae, except a narrow line
above, brownish. Male unknown. Rhodesia. . . . ale. bob Arnold Distinct transverse carina below antennae, sometimes fine and only visible in ventral aspect. At least tergites 5—6 reddish or hind tibiae dark. . . . 17
Petiole little longer than first tergite. Apical margins of fourth aad) fifth one with fringe of yellowish-golden hairs, at least as long as width of base of hind basitarsi, but shorter than in group of turneri. Frontal carina ending normally as single carina in transverse carina below antennae, transverse carina almost rectangular in frontal aspect. Scutum partly transversely striato-punctate and also with some very large punctures. Prescutal sutures ill-defined. Apical half of pygidial area narrow with parallel carinae. Apical margin of first tergite and tergites 2—6 completely reddish. Legs reddish, back of mid femora and hind femora except underside dark brown. Fore and mid basitarsi GET Length about 6 mm. Male unknown. Sierra
Benet OU. Sule hee) eon nis sp. NOV: Petiole much lonen Apical margins 1. fourth i fifth sternites without fringes of long hairs. . . ls
Raised part of Bone) carina ) meh Broadenedt betweet aften proadened part usually not closed below but ending with two parallel carinae in transverse carina below antennae. Upper part of propodeum finely obliquely striate. Pygidial area distinct. Fore and mid legs not whitish, at most mid basitarsi . . 5 dl) Carina between antennae much less broadened, or broadened part drat and inter- antennal carina ending as a single carina in transverse carina. Lateral carinae of pygidial area not well-defined, if more distinct, fore and mid legs extensively
whitish. Petiole about twice as long as first tergite, in dorsal aspect . . . . 23 Gaster black, at most last tergite brownish, or petiole completely and first tergite parilygredcish-browntt Mt... 2,2,.2,2 2 ae es dien 20 Gaster partly reddish . . . . #21
Frons very finely obliquely striate, vertex ih call sculptures Scum del super- ficially transversely striate and with some larger punctures. Petiole more than twice as long as first tergite, in dorsal aspect. Antennae reddish, apical half darkened. Fore and mid legs reddish-brown; at least mid basitarsi yellowish-white; hind legs entirely brown. Pronotal tubercles blackish; black. Length 7 mm. Rhodesia .
3 stevensoni Arnold Ea ona finely ee, ek mat meer Scutum shining, anterior two-thirds sparsely punctate. Petiole less than twice as long as first tergite. Antennae reddish-brown, segments 7—11 more or less darkened dorsally. Fore and mid legs including mid basitarsi and base of hind tibiae reddish-brown; fore and mid femora partly darkened, hind femora black, hind tibiae dark brown. Pronotal tubercles reddish-brown. Gaster black, petiole at least ventrally and laterally partly reddish- brown, also first tergite more or less brown, apex of last segment yellowish-red. Length 7 mm. South Africa . . . . . jfacoti sp. nov. Apical margin of fourth tergite and Late 56 zoeen reddish. Mid basitarsi brown. Scutum dull, finely punctate. Petiole over twice as long as first tergite, in dorsal view. Pygidial area broader, lateral carinae slightly converging towards apex. Length 8 mm. Male unknown. Rhodesia . . . . . . . . luciuosus Arnold Gastral tergites 1—6 reddish. Pygidial area narrow . . bide 22 Frons very finely punctate, vertex almost impunctate. Petnle over twice as long as
46
208
24.
25%
26.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
first tergite, in dorsal aspect. Antennal segment 1—12 reddish below. Greater part of mandibles reddish-brown. Legs brown, fore tibiae and tarsi reddish, mid basitarsi testaceous. Length 7.75 mm. Male unknown. Zaire. . . . . lusingae Leclercq Frons finely densely striato-punctate, vertex indistinctly striate. Petiole slightly over one and three-quarters times as long as first tergite, in dorsal aspect. Antennae black, only small central part of segments 5—12 below reddish-brown. Mandibles darker. Legs black, last tarsal segments, underside of fore tibiae and base of hind tibiae
brownish. Length 7 mm. Male unknown. Ethiopia. . . . . rugifrons sp. nov. Tergites 1—6 or 2—6 reddish . . . Mrt 24 At least tergites 1—3 black or very ane prow mare rad end aaf edi brown". at MERG
Pronotal ande at most hy reddish or Rare beo ae rarely yellowish; if so cf. bidentatus pallidus). Fore and mid legs reddish, underside femora brownish, base of fore tibiae often somewhat yellowish, rarely a narrow yellowish streak on foreside, fore and mid basitarsi yellowish or yellowish-red; hind legs reddish on inner side, dark brown on outer side, base of hind tibiae usually dark. Mandibles reddish. Antennae reddish, apical half dorsally brown. First gastral tergite mostly dark brown (reddish in female from Angola). Punctation of scutum superficial, interstices larger than punctures, slight tendency to transverse striation. Propodeum with narrow smooth margin behind enclosed area, back dorsally obliquely striate, below irregularly carinate. Pygidial area dull, lateral carinae not sharp, parallel. Length 8.5—9.5 mm. Tanzania, Kenya, Rhodesia, Angola, Zaire . ike . bidentatus bidentatus (Cn At ies: posero: half of TA tubereles Aaa foreside of fore tibiae distinctly yellowish. Tergites 1—6 or apex of first tergite and tergites 2—6 completely reddish. Length 7.5—8 mm . . . LIFE RISORSE 25 Only pronotal tubercles, foreside of fare “ao Ba DE aid apie and fore and mid basitarsi yellowish or whitish. Base of hind tibiae paler than rest of hind tibiae. Punctation of scutum coarser than in bidentatus bidentatus, more striato-punctate with interstices larger than punctures and more strongly transversely striate. Sculpture on back of propodeum reaching enclosed area, except laterally. Pygidial area as in nominate subspecies. Length 7.5 mm. Zaire, Rhodesia, Malawi, Equatorial Guinea bidentatus pallidus subsp. nov. PORC partly, tonen bereide and metanotum yellow. Fore and mid legs yellowish-white, femora with light brown streak behind; greater part of hind legs brown. Scutum finely punctate, slight tendency to transverse striation. Propodeum behind enclosed area smooth, back with oblique striae above and irregular carination below. Pygidial area dull, more distinctly defined, almost rectangular and with two rows of fine punctures. ae 8 mm. Male unknown. Nigeria, Sierra Leone . nigeriae Leclercq At east cite 56 as Sui as apical half or four rei reddish. Thorax black, pronotal tubercles partly dark brown. Fore legs except trochanters reddish-brown, underside of fore femora darker; mid legs reddish-brown, underside of femora and tibiae dark brown; hind legs dark brown or black. Mandibles reddish. Scutum with large, somewhat superficial punctures and distinct transverse rugae especially on outer side of prescutal sutures. Narrow smooth margin behind enclosed area of pro- podeum, back with fine oblique striae, medially sometimes reaching enclosed area.
27:
28:
29.
30.
J. P. van LITH: Revision of Ethiopian Psenini 47
Pygidial area as in bidentatus bidentatus. Length 8.75 mm. Male unknown. South ARIANE „IR. . . + . bidentatus rubrocaudatus Turner Tergites 4—6 geek be Thorax black including pronotal tubercles. Legs some- what darker reddish-brown, except fore basitarsi. Mandibles reddish. Scutum finer punctate, very little tendency to transverse striation. Narrow smooth margin behind enclosed area, back side dorsally with distinct fine oblique striae. Length 9.5 mm. Male’unknown. Ethiopia . . „Ti et. . . . bidentaius subsp. nov.?
No well-defined pygidial area. Scutum transversely rugoso-punctate. Petiole less than twice as long as first tergite. This tergite of usual shape. Upper part of pro- podeum laterally with oblique striae, on either side of median sulcus a smooth area. Back of propodeum with coarse reticulate carination. Distinct transverse carina below antennae. Median part of pronotum, and whole metanotum, yellowish. Petiole black, tergites 1—6 reddish. Fore and mid legs yellow, fore and mid femora brown below, hind femora completely brown. Antennae dorsally dark brown. Length 7.5
mm. Male unknown. Zaire, Uganda. . . . een wghesgutenenlteclereg Distinct narrow pygidial area. Scutum oi finely He First gastral tergite Sliphitly}modoseltt. tu. |. 128
Hind legs and tergites 1—6 een Pronotum JU oi ess small marks on scutum, scutellum, metanotum, large part of back of propodeum and a mark on anterior plate of mesepisternum pale yellow, also fore and mid legs. Petiole black with yellowish base. Transverse carina below antennae lacking or indistinct. Back of propodeum almost sega smooth. Length 8—9 mm. Sierra Leone
3 oweni sp. nov. Hind LA ac aie dea ria or ati Sa coe dark brown and only Apicdemarcinsidancreddishers wah. banusnanl dert de store wer stun mul 29 Pronotum, pronotal tubercles, four marks on scutum, scutellum, metanotum, back of propodeum and parts of sides of thorax yellow. Base of petiole yellow, remainder dark brown, gaster brownish. Fore legs mostly yellow; foreside mid legs yellowish, back-side brownish-red; hind legs brown but back of femora partly yellow. Fine transverse carina below antennae. Upper part of propodeum smooth, lower two- thirds of back-side with fine reticulate carination. Petiole about twice as long as first
tergite. Length about 10.5 mm. Male unknown. Equatorial Guinea . 2 uelleburgi si nov.
Pronotum dorsally, pronotal tubercles, scutellum and metanotum yellowish-red. Petiole reddish-brown or darker brown, base of tergites dark brown, apical margins dark reddish. Fore and mid legs including trochanters reddish, mid femora brownish behind, hind legs brown, base of tibiae reddish. Transverse carina below antennae fine or indistinct, vertical carinae downwards from outer side of antennal sockets distinct. Propodeum behind enclosed area smooth, back-side finely punctate, apex finely reticulate. Petiole slightly longer than first tergite. Length 8 mm. Male unknowg.Malapasy; Eee. honest werd deg ver wll ‘nd air dlectzs Saussure
Lateral corners of pronotum obtuse. No sharp transverse carina below antennae, at most an indistinct ridge, but below outer side of each antennal socket a short vertical carina, in dorsal aspect protruding like a tooth. Punctation of scutum variable, sometimes very coarse. Upper part of back of propodeum usually distinctly obliquely striate. Pygidial area indistinct. Foreside of fore and mid tibiae yellow, also
48
Dlt
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
fore and mid basitarsi and base of hind tibiae. Length about 7 mm. South Africa, Mozambique, Malawi, Zambia, Tanzania, Uganda, Zaire, Sierra Leone. The sub- species /aevior Arnold, Ethiopia, female unknown, is distinguished by the brown femora and weaker sculpture of scutum and propodeum . capensis capensis Brauns Lateral corners of pronotum sharp. No distinct transverse or vertical carinae below antennae, sometimes a small low shining tubercle below antennae. Punctation of scutum variable but usually finer than in capensis. Propodeum behind enclosed area usually smooth. At least foreside of fore and mid tibiae more or less yellowish. hanger: waal fee de elec We athe Side" ee Se chi a RE Fore tarsi, a part of mid basitarsi, and base of hind tibiae yellowish-white. Puncta- tion of scutum variable, usually fine and sparse. Pygidium with two indistinct, close, parallel carinae at apex. Length 8—9 mm. Tanzania, Rhodesia, Uganda, Zaire, Nigeria, Sierra Leone. . |. . +. . datiannulatus latiannulatus (Cameron) Mid basitarsi and base of kia tibiae black or dark brown. Punctation of scutum stronger. Distinct narrow pygidial area, with very close, parallel carinae. Length 7.75 mm. Zaire, Rwanda, Cameroon. . . . . . latiannulatus basilewskyi Leclercq
KEY TO THE MALES OF Psenulus
(the males of P. aurifasciatus, P. dilectus, P. erusus, P. garambae, P. ghesquierei, P. kobli, P. leoninus, P. luctuosus, P. lusingae, P. nigeriae, P. patei, P. rugifrons, P. saltitans,
P. sapobaensis, and P. velleburgt are still unknown)
Tempora coarsely, mostly longitudinally, striato-reticulate. Raised part of carina between antennae more or less broadened. Antennae with tyloidea. Anterior margin of pronotum alveolate. Prescutal sutures long, almost reaching hind margin of scutum. At least thorax and petiole black . . . (Group of Psenulus turneri) 2 Tempora much less coarsely sculptured, sometimes finely striate (see benoiti) . 6
Upper part of back of propodeum smooth. Frons shining, finely punctate. Antennal segments long, segments 3 (or 4)—8 with distinct tyloidea. Scutum shining, finely punctate. Second submarginal cell of fore wings open dorsally. Tergites 1—6 reddish, only last sternites slightly brownish, all legs reddish. Sierra Leone, Equa-
torial Guinea . . . . freetownensis sp. nov. Upper part of back a de sail si At least first tergite com- pletely.blackind. hab sum: pane ahi bone NES
Second submarginal cell not petiolate or triangular, dorsally open. Frons obliquely striate. Antennal segments 8—12 at least one and a half times as long as broad at apex, third segment with very small tyloidea, segments 4—10 with distinct long tyloidea, on third and 11th segment at most a small point. Scutum strongly trans- versely striate. Petiole about one and a half times as long as first tergite, in dorsal aspect. Tergites 1—3 black or dark brown. Fore and mid tibiae and tarsi and fore- side of fore and mid femora reddish; back of fore and mid femora, as well as hind legs completely, brown except pale yellowish base of hind tibiae. Zaire, Sierra Leone
: ealae Leclercq Soi Sebra cell idee te ae la in turner? or paulisae). Base of hind:tibiaeidarkn: Dei yao WIR en SD ai Arr
CI
J. P. van LITH: Revision of Ethiopian Psenint 49
Petiole nearly one and one-third times as long as first tergite, in dorsal aspect. Frons obliquely striate. Antennae moniliform, segments 8—12 little longer than broad, segments 3—10 with distinct tyloidea, small on tenth segment. Scutum distinctly punctate, slight tendency to transverse striation. Prescutal sutures with large alveoles. First tergite, basal two-thirds of second tergite and sides of tergites 3—5 partly black. Fore and mid legs including trochanters reddish, also hind tibiae and tarsi, hind femora brown. Zaire, Sierra Leone m based on male from Sierra
BEONE) = . i 02) 2) paulisaeleelereq Petiole usually diaz lugo ou one and a ale x or twice as long as first tergite, in dorsal aspect, or scutum more coarsely sculptured. . . . . . . 5
Petiole about one and a half times as long as first tergite, in dorsal aspect. At least tergites 1—3 fully black, fourth tergite more or less darkened. Fore legs including trochanters reddish-brown, femora brown below, mid legs reddish-brown, darkened below, trochanters brown; hind legs for greater part dark brown. Antennal segments 3—8 about one and a half times as long as wide at apex, segments 4—8 with distinct small tyloidea, 9—10 with indistinct tyloidea. Frons obliquely striate, vertex more superficially striato-punctate. Scutum more coarsely punctate than in pazlisae, with some transverse rugae, especially across the prescutal sutures, and large alveoles. South Africa . . . woes urnen Arnold Petiole nearly twice as iene as ae ent Set O and two-thirds of second tergite black, greater part of sternites 2—4 blackish-brown. Fore legs including trochanters reddish-brown, mid legs slightly darker brown, hind legs dark brown. Antennal segments 3—8 about one and a half times as long as wide at apex, segment 3 with small tyloidea, segments 4—8 with longer, distinct, tyloidea. Frons and interocellar area coarsely, frons obliquely, striato-punctate, vertex between ocelli and oculi more finely striato-punctate. Anterior half of pronotal dorsum with stronger longitudinal short cärinae, alveolate. Scutum distinctly punctate, tendency to transverse striation, rugae across prescutal sutures very coarse, alveoles larger than in turneri. Female unknown. Rhodesia . . . Uni... alveolatas'sp. nov. First gastral tergite not distinctly nodose, or arie 1—6 or 2—6 reddish . . 7 First gastral tergite distinctly nodose. Prescutal sutures short. Raised part of frontal carina between antennae sharp. Gaster black, fore and mid tibiae at least partly yellowish or yellowish-white . . . . . (Group of Psenulus latiannulatus) 16 Petiole and gaster reddish. Interantennal carina not distinctly broadened between antennae, as it is in female. Upper part of back of propodeum smooth behind enclosed area. Petiole about one and a half times as long as first tergite. . . 8 Petiole black, at most base yellowish, or sides dark reddish-brown. . . . . 9 Scutum coarsely transversely striato-punctate. Mesopleura coarsely punctate, hypo- epimeral area more finely. No tyloidea on antennal segments. Scape of antennae yellow, rest of antennae yellowish-brown, brownish above. Pronotum, pronotal tuber- cles, tegulae, hind margin of scutellum and metanotum yellow. Fore and mid legs including trochanters yellow; hind legs i coxae a ner Length 6 mm. Malagasy . . . 5 . . + fulgidus Arnold Scutum shining, Ca Mare A variable size, interstices mostly larger than punctures. Mesopleura finely punctate. Antennal segments 3 (or 4)—13 with distinct oblique tyloidea. Scape of antennae yellow, rest of antennae reddish. Two marks on pronotum, and pronotal tubercles, yellow or yellowish-white. In var.
50
10.
12%
13:
14.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
alboscutellatus Arnold also scutellum with yellowish-white mark. Fore and mid legs including trochanters whitish, femora brownish below and last tarsal segments reddish; hind legs ee: coxae reddish. Length 8.5 mm. Malagasy .
reticulosus Arnold mora: ata aast ple at most Sa ped nad sides of petiole reddish- brown. Interantennal carina broadened. Median part of propodeal enclosure rather broad. Upper part of back of propodeum finely Eh striate. Petiole long . 10 Gaster more or less reddish . . . CANARIA Petiole at least twice as long as dt i Hat in dl ar Frons with superficial oblique striae. Antennal segments 10—12 about as long as broad; seg- ments 3—13 with tyloidea. Scutum transversely striato-punctate, prescutal sutures almost reaching hind margin. Pronotal tubercles dark brown. Legs and veins of fore wings reddish-brown. Petiole completely black. Gaster with somewhat appressed silvery pubescence. Length 6 mm. Rhodesia. . . . + . stevensoni Arnold Petiole little more than one and a half times as one: as first tergite. Frons shining, finely punctate. Antennal segments 10—12 about one and a half times as long as broad; segments 4—11 with tyloidea, indistinct tyloidea on segment 12. Scutum shining, scarcely finely punctate, prescutal sutures posteriorly slightly indicated. Pronotal tubercles reddish-brown, sides of petiole somewhat reddish-brown. Legs black, fore tibiae and tarsi and underside of mid and hind tibiae yellowish-brown. Veins of wings black. No appressed silvery Pena on gaster. Length nearly 7 mmi:South;Africa vem è . . facoti sp. nov. Scutum distinctly or ei transversely es, Une Propodeum black. Petiole at least twice as long as first tergite. Antennae very strongly moniliform,
with tyloidea. Interantennal carina slightly or not broadened. . . . . . . 12 Scutum smooth, finely punctate, or coarsely punctate and back of propodeum with yellow marking. Antennae not strongly moniliform . . . Licata «13
Pronotal tubercles dark brown. Fore and mid legs reddish- Bon basitarsi paler, femora below dark brown, hind legs dark brown. Mandibles reddish. Base of petiole dark brown or black. Gastral tergites 2—6 reddish. Pronotum and metanotum black. Scutum with coarse punctures and light transverse rugae. Narrow smooth margin behind propodeal enclosure, sometimes lacking. Length 8 mm. Tanzania, Rhodesia, Kenya,rAngola, Zaire In MIE . . +. . bidentatus bidentatus (Cameron) Pronotal tubercles yellowish or lo red. Fore and mid tibiae yellow, basitarsi whitish. Mandibles yellowish. Base of petiole somewhat yellowish-red (not in female). Gastral tergites 1—6 or 2—6 reddish. Pronotum partly and metanotum more or less yellow, rarely black. Scutum with distinct punctures and feeble trans- verse rugae. Broader smooth margin behind enclosed area of propodeum. Length 7 mm. Zaire, Equatorial Guinea, Rhodesia and Malawi . bidentatus pallidus subsp. nov. Thorax black. Prescutal sutures long. Frontal carina more or less broadened between antennae. Scutum finely punctate . . . ses “age A Thorax and propodeum with yellow maine. Pu sutures na one-third or one-half of length of scutum. Raised part of frontal carina narrow between antennae. First gastral tergite slightly nodose. No tyloidea . . . . . . . 15 Upper part of propodeum smooth, back-side coarsely reticulate. Petiole about one and one-third times as long as first gastral tergite, in dorsal aspect. Antennal segments slightly rounded below, no distinct tyloidea. Pronotal tubercles yellowish-brown.
15:
16.
17.
J. P. van LITH: Revision of Ethiopian Psenini 51
Tergites 1—6 reddish. Fore and mid legs reddish, hind legs brown. Zaire .
ma benoiti Leclercq Upper Bar of ren ie indistinctly Blans striate, back with coarse reticulate carination. Petiole about two and one-third times as long as first tergite. Antennal segments more cylindrical, less rounded below, segments 3—12 with very fine narrow shining tyloidea, indistinct on twelfth segment. Pronotal tubercles dark brown. First gastral tergite brownish, 2—6 reddish. All femora blackish-brown, fore and mid tibiae brown. Female unknown. Zaire . . . . . . avernus Leclercq Anterior lateral corners of pronotum obtuse. Scutum coarsely punctate, scutellum and metanotum almost impunctate. Upper part of propodeum behind enclosed area smooth, rest coarsely reticulate. Sides of propodeum with narrow smooth anterior margin. Mesopleura distinctly punctate, interstices about three or four times as wide as punctures. Petiole about one and one-third times as long as first tergite, in dorsal aspect. Pronotum, pronotal tubercles, two square marks on scutellum, metanotum, two marks on back of propodeum and small spot on anterior plate of mesepisternum, brownish-yellow. Base of petiole yellowish-white, tergites 2—6 reddish, base of second tergite black. Fore and mid legs including trochanters brownish-yellow, back of femora brown; hind trochanters and femora brown, median part of hind tibiae brownish, rest of hind legs reddish. Female unknown. Zaire . . trevirus Leclercq Anterior lateral corners of pronotum sharp. Scutum shining, very finely punctate. Upper part of propodeum smooth, back coarsely reticulate, sides of propodeum with broad smooth anterior margin. Mesopleura finely punctate, interstices larger. Petiole about one and a half times as long as first gastral tergite. Pronotum, pronotal tuber- cles, two or four small longitudinal marks on scutum, scutellum, metanotum, four marks, connected below, on back of propodeum and a small spot on anterior plate of mesepisternum, whitish-yellow. Base of petiole yellowish-red, tergites 1—6 reddish, first tergite more or less darkened. Fore and mid legs including trochanters pale yellow, femora brown below; hind legs including trochanters dark brown, underside femora and basal half of tibiae more reddish. Sierra Leone . . . oweni sp. nov. Anterior lateral corners of pronotum with obtuse angle. Antennal segments 4—12 behind, on line between dark brown upper side and yellowish-brown underside, with low but distinct tyloidea. Scutum with coarse variable punctation, apically finer but more densely punctate. Upper part of propodeum behind enclosed area completely rugoso-striate. Black; fore and mid tibiae, except brown mark below, and tarsi yellow, base of hind tibiae whitish. Length about 7 mm. South Africa, Mozambique, Malawi, Zambia, Tanzania, Uganda, Zaire, Sierra Leone, Ethiopia (subspecies laevior Arnold, distinguished by brown femora and weaker sculpture of scutum and Propodeum m. SUE : : . . « capensis capensis Brauns Anterior lateral corners be Bro more of Te sharp, never obtuse. Antennae without distinct tyloidea. Upper part of propodeum behind enclosed area usually at least partly smooth, or base of hind tibiae dark. Length 7.5—8 mm . . . 17 Mid basitarsi and base of hind tibiae whitish, apart from other yellowish-white parts of fore and mid legs. Antennae dorsally black, reddish-brown below. Punctation of scutum distinct, interstices large, punctures usually small, rarely larger. Upper part of propodeum behind enclosed area near median sulcus usually smooth. Anterior lateral corners of pronotum more or less sharp. Length 8 mm. Tanzania, Rhodesia, Uganda, Zaire, Nigeria, Sierra Leone . . Jatiannulatus latiannulatus (Cameron)
52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
— Mid basitarsi and base of hind tibiae dark brown. Antennae dorsally blackish-brown, below on basal half reddish-brown, on apical half brown. Punctation of scutum coarser. Upper part of propodeum behind enclosed area obliquely striate or rugose. Anterior lateral corners of pronotum sharp. Length 7.5 mm. Zaire, Rwanda, Came- toon | te 2 et enten ao Ane nr benleusey reel
Psen (Psen) silvaticus Arnold
Arnold, 1924: 38—39, 9 (Psen silvaticus; Rhodesia). Leclercq, 1961a: 15, Q (Psen (Psen) silvaticus; Rhodesia).
Material studied: Rhodesia: 1 9 (holotype), “Cloudlands, 6000 ft, Vumba Mts, 6—17 April, 1923”, 1 ©, Vumba Mts, Umtali dist, 7 March, 1930, coll. R. H. R. Stevenson, 1 ©, Vumba Mts, March, 1930, coll. R. H. R. Stevenson, 2 9, Vumba, March, 1930, coll. R. H. R. Stevenson, 1 9, Vumba Mts, 1 March, 1938, coll. G. Arnold (RM); 1 ©, Vumba, March, 1930, coll. R. H. R. Stevenson (SAM).
Arnold (1924) recorded the holotype as follows: “Mt. Chinzewa, 6200 ft, Vumba Range, near Umtali, S.R. One female taken flying round a species of wild raspberry in the hollow stems of which it probably nests. Type in R.M. coll.’ Mr. F. C. de Moor kindly checked his topocadasteral map of the Vumba Mountains area because of the discrepancy between the locality label under the type and Arnold’s record. In his letter, 10 May, 1973, he writes: “Cloudlands is a Private Game reserve, the highest point on this farm is an unnamed mountain with a beacon “Castle Beacon 6245 feet”. I can find no reference to Mt. Chinzewa however”.
Mr. de Moor also reported to me that in March, 1930 in total ten specimens were collected in the Vumba Mts., with the following note written by Arnold: “Nesting in cracks in old rotten tree stump (prone). All ten specimens taken from same spot in dull weather.”
Under one of these specimens the prey has been pinned, an adult Homopteron be- longing to the Cercopidae, about 7 mm long. Cercopidae have been recorded earlier as the prey of Palaearctic and Indo-Australian species of Psen.
Supplementary description. — Frontal carina distinct on lower half of frons, on upper half almost absorbed by the coarse close punctation. Arnold’s figure of the head wrongly gives the impression that there is a transverse carina below the antennae. In reality two very fine carinae connect the interantennal tooth with the inner side of the antennal sclerites (Fig. 1). Propodeal enclosure ill-defined, with irregular reticulate carination, back very finely reticulato-carinate. Hypo-epimeral area striato-punctate, like adjacent part of mesopleura. Outer epicnemial carinae below hardly receding backwards. No acetabular carina. Mesosternum dull, finely densely superficially punctate, fine median longitudinal carina. Mesopleura on lower part shining, with large punctures and tendency to striation. Upper longitudinal half of hind femora smooth, separated from normally pubescent lower part by a narrow longitudinal area of dense fine punctation and fine short hairs, somewhat broadened near apex, not distinctly depressed there. Petiole cylindrical with long lateral hairs, directed obliquely downwards, sometimes worn off.
This species much resembles P. leclercgi sp. nov. from Malagasy.
J. P. vAN LITH: Revision of Ethiopian Psenini 53
Psen (Psen) congolus Leclercq
Leclercq, 1961a: 15—16, 2 and  (Zaire).
I have examined one of the paratypes (a male) from Lubumbashi (Elisabethville), Dec., 1933, coll. M. Bequaert (FAG). The structure of the epicnemial area and of the mesosternum agrees with that of the Indo-Australian representatives of the subgenus Psen. The acetabular carina is short. Hind margins of third and fourth sternites with tufts of long brownish hairs.
Psen (Psen) leclercqi spec. nov.
Female. — Head black; mandibles reddish-brown except dark tips, labrum reddish- brown, palpi yellowish-brown, antennae black but underside of scape, of second, third and 12th segment reddish-brown. Thorax black, with following parts reddish-brown: pronotum dorsally and upper part of foreside, pronotal tubercles, anterior corners of scutum and a narrow margin along tegulae, upper two-thirds of anterior plate of mesepisternum and tegulae. Legs black with following parts reddish-brown: fore tibiae and tarsi, underside of mid and hind tibiae, and tarsi. Calcaria yellowish-brown. Petiole reddish, gaster black but greater part of first and second gastral tergites, apex of last segment and greater part of sternites reddish-brown. Veins of wings black but basal part and stigma reddish-brown.
Clypeus feebly convex, like supra-clypeal area densely but superficially punctate, protruding median part slightly emarginate (Fig. 2, 3). Frontal carina reaching groove around median ocellus but almost absorbed by sculpture of frons. Carina ending below antennae in low triangular tooth, connected with inner side of antennal sclerites by fine carinae. Frons flat, up to ocelli densely striato-punctate, interstices shining, very narrow margin along oculi finer and sparsely punctate. Punctation between ocelli fine, interstices about as wide as punctures. Vertex behind ocelli with close and large punctures, laterally coarser and somewhat striato-punctate, with only a few smooth places about as large as punctures. Postocellar region not distinctly raised. Temples finely but distinctly punctate, interstices about as wide as punctures or slightly more. Occipital carina ending in hypostomal carina, both narrow and high, but lower than in P. madecassus Arnold. Scape of antennae long, at least three times as long as wide, third segment nearly four times, fourth segment more than twice, segments 5—7 about one and a half times, eighth segment about one and a third times as long as wide at apex, segments 9—11 slightly longer than wide, 12th segment about twice as long as broad at base.
Pronotal corners obtuse. Scutum densely and coarsely rugoso-punctate (Fig. 4), inter- stices shining, medially not more than a narrow line. Prescutal sutures indicated by a row of much finer punctures. Scutellum somewhat striato-punctate, interstices larger than punctures, metanotum with irregular longitudinal carinae. Propodeal enclosure depressed, lateral parts with oblique longitudinal carinae, central part with irregular carination; back of propodeum with coarse irregular reticulate carination reaching enclosed area. Sides of propodeum with oblique carinae and fine punctures between. Mesopleura coarsely striato-punctate, striae posteriorly more striking, lower part finer sculptured, hypo-epimeral area coarsely densely punctate. Epicnemial areas almost smooth, inter- epicnemial area finely punctate, mesosternum depressed behind short acetabular carina, antero-laterally obliquely striato-punctate, medially with strong transverse carinae. Meta-
54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
pleura shining, with some low longitudinal rugae on posterior half. Anterior plate of mesepisternum with irregular long, partly vertical, striae; anterior oblique suture foveolate, widened upper part with transverse striae, continuing on hypo-epimeral area. Epicnemial carina below normally curved backwards. Metasternum with deep triangular emargination.
Femora heavy, hind tibiae dorsally with row of short thick thorns and thin long white spines. Base of second submarginal cell of fore wings about one and a half times as long as upper side, receiving first recurrent vein at about one-third from inner side; second recurrent vein ending in this cell very near outer side.
Petiole as long as first two tergites and half third tergite, dorsally completely smooth, impunctate, apically somewhat flattened, sides longitudinally slightly depressed, only lower keel distinct, ventrally an indistinct blunt median longitudinal keel. Tergites basally very finely punctate, on third tergite deeper than on second tergite; interstices about as wide as punctures, on apical two-thirds interstices a few times as wide as punctures. Margin of fourth tergite narrowly, of fifth broadly impunctate. Pygidial area elongate-triangular, defined by distinct carina, apex rounded, surface shining but extreme apex dull, laterally a few small punctures, each bearing a short stiff hair (Fig. 5). Sternites sparsely punctate, somewhat more densely at base, especially antero-laterally on second sternite.
Clypeus with appressed golden pubescence and long erect hairs. Supra-clypeal area and lower part of frons with less appressed golden pubescence; upper part of head with longer, brownish pubescence. Temples with erect greyish pubescence, pronotum with appressed greyish-white pubescence, thorax with long and dense pubescence, brownish on scutum, whitish on rest of thorax, denser on mesosternum. Legs with greyish hairs, base and apex of hind tibiae on inner side and underside of hind basitarsus with very dense and short, yellowish-golden pubescence. Upper half of back of hind femora smooth and shining, separated from normally pubescent lower half by a dorsally sharply defined narrow line of fine dense punctation and short pubescence. Petiole below lower lateral longitudinal carina with row of fine punctures, each bearing a long obliquely downwards directed hair and also with a row of dense short hairs, densest apically. Tergites sparsely reddish-brown pubescent, dense on lateral parts of hind margin. Apex of fifth tergite before smooth hind margin with a few long stiff hairs, also sixth segment with long stiff hairs. Base of second sternite with dense pale pubescence, on disk and on hind margins of following sternites a few long stiff hairs.
Length about 11.5 mm.
Male unknown.
Malagasy: 1 ® (holotype), “Madagascar, Rogez, II.44” (FAG).
This is the second species of Psen known from Malagasy. It is easily distinguished from P. madecassus Arnold, which is smaller, more reddish, and much more finely, very sparsely, punctate, especially on front, vertex, scutum and sides of thorax.
P. leclercqi is very similar to P. silvaticus Arnold from Rhodesia but it differs not only in the reddish anterior corners of the scutum, but especially in the sculpture of the propodeum. In P. si/vaticus the enclosed area is hardly distinguishable, with irregular reticulate carination; back-side of propodeum very finely reticulato-carinate. In P. leclercgi the propodeal enclosure is well-defined, lateral parts of enclosure shining, with oblique distinct carinae, median part with coarse irregular sculpture, back of propodeum coarsely irregularly reticulato-carinate.
J. P. van LITH: Revision of Ethiopian Psenini 55
3,
Fig. 1. Psen (Psen) silvaticus Arnold, @. Face. Fig. 2—5. Psen (Psen) leclercqi sp. nov., 2, holo- type. 2, face; 3, clypeal margin; 4, punctation median part of scutum; 5, pygidial area
56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
Psen (subgenus?) madecassus Arnold
Arnold, 1945: 152—153, $ and & (Psen madecassus; Malagasy: Bekily, Ranomafana, Ivondro). Leclercq, 1961a: 15, $ and 4 (Psen (Psen) madecassus; Malagasy: Bekily, Rogez). Leclercq, 1961c: 117, 2 (Malagasy: Mandraka).
Material studied: Malagasy: 1 ©, paratype and 1 G', paratype, both labelled “in cop. B.”, Bekily, Febr., 1937, coll. A. Seyrig, 2 &, Bekily, Jan., 1937, coll. A. Seyrig, 1 9, Perinet, “Forêt côte est”, Febr., 1939, coll. A. Seyrig (RM); 1 9, Bekily, Jan., 1942 (BM); 1 &, Bekily, Jan., 1937, coll. A. Seyrig (author's collection); 1 g', Rogez, June, 1944, coll. A. Seyrig (MRAC); 1 9, Madagascar Est, Mandraka (Tamatave), 4 April, 1958, coll. F. Keiser (NMB).
The excellent description by Arnold may be supplemented with the following char- acters. — Frontal carina fine, reaching anterior ocellus, slightly widening at lower end and ending in a low small tubercle. Lateral parts of frons with large low tubercle. In both sexes occipital carina narrow and high, especially lower part, ending normally in hypostomal carina. Apical margin of clypeus thick, slightly emarginate.
Scutum coarsely punctate, interstices as large as, sometimes larger than punctures. Mesopleura not closely and finely punctate, as stated by Arnold, but finely and sparsely, including hypo-epimeral area, interstices many times larger than punctures. Mesosternum depressed and smooth behind interepicnemial area, with high median longitudinal carina, anteriorly defined by a distinct short acetabular carina. Metasternum deeply emarginate. Petiole dorsally smooth, latero-dorsally with shallow narrow groove, sides of petiole with distinct groove, also ventro-laterally a distinct longitudinal groove; in all these grooves a row of fine punctures, each with a long erect fine hair. Hind coxae with sharp inner longi- tudinal carinae. Upper third of hind femora smooth, medially a broad, densely finely punctate and pubescent, longitudinal line, lower part of femora sparsely longer pubescent. Pygidial area of female (Fig. 13) elongate-triangular, apex emarginate, lateral carinae high. Antennae of male without distinct tyloidea. No fasciculate hairs on apical margins of third and fourth gastral sternites, but margins of second and following sternites with fine pale, obliquely inwards directed, long erect hairs. Genitalia of male: Fig. 6, 7, 8, 9. Seventh sternite: Fig. 10. Eighth sternite: Fig. 11, 12.
P. madecassus does not belong to the subgenus Pser because of the somewhat different shape of the sternum, the fine, punctate, latero-dorsal sutures of the petiole and the absence of fasciculate hairs on the apical margins of the third and fourth gastral sternites of the male. Probably it should best be placed into the subgenus Punctipsen, of which representatives are known from the Palaearctic region, Ryukyu Islands, Formosa and S India. The characters they have in common are the punctation of the petiole, the almost vertical foreside of the first tergite, the continued longitudinal rugae on the back of the propodeum, behind the enclosed area, the thick apical margin of the clypeus, the absence of fasciculate hairs on the sternites of the male and to some extent also the shape of the male genitalia.
Fig. 6—12. Psen madecassus Arnold, &. 6, outer valve of genitalia, dorsal aspect; 7, the same,
lateral aspect; 8, the same, inner aspect, with volsella; 9, right penis valve, outer side; 10, 7th
sternite, ventral aspect; 11—12, 8th sternite, lateral and ventral aspect. Fig. 13. Psen madecassus
Arnold, 9, pygidial area. Fig. 14—18. Psen patellatus Arnold, 6, paratype. 14, face; 15, 6th, 7th and 8th sternites, ventral aspect; 16—18, genitalia, ventral, lateral and posterior aspect
J. P. van LITH: Revision of Ethiopian Psenini
58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
P. madecassus differs from the earlier described species belonging to Punctipsen in the reduced punctation of the dorsal side of the petiole and in having no tooth on mandibles or mid basitarsi of the males, but I think that these characters do not have any sub- generic value.
The hind coxae have a sharp inner carina as in the subgenus Pseneo, but there is no oval patch on the posterior apex of the hind femora, the clypeus is bidentate and the petiole has latero-dorsal grooves with fine punctures and long erect lateral and ventral hairs.
Arnold (1945) records one specimen with prey, a small adult Cercopid.
Psen (subgenus?) patellatus Arnold
Arnold, 1924: 39—40, & (Psen patellatus; SW Africa: Onoolonga and Mafa). Leclercq, 1961a: 15 (Psen (Psen) patellatus)).
Material studied: South-West Africa: 1 & (holotype), Mafa, Febr., 1923, 1 {, Onoolonga, Febr., 1923, det. Arnold (SAM); 1 g', Onoolonga, Febr., 1923, with green paratype label (RM).
Dr. Hesse of the South African Museum informed me that there are two more males in his collection, respectively one from Mafa and one from Onoolonga. Arnold (1924) recorded in total three males from Onoolonga and Mafa, he did not mention which was the holotype.
Little can be added to the detailed original description. — Face broad (Fig. 14), hypostomal carina ending normally in occipital carina. Mandibles unidentate at apex. Sternum very densely punctate, no acetabular carina, lateral epicnemial carinae not curved backwards as in subgenus Psen. Petiole short, not only dorsally, but also laterally with irregular longitudinal carinae; similar carinae on ventral side which are converging back- wards; ventral plate with fine longitudinal rugae. Fourth and fifth gastral sternites with- out fasciculate hairs. Sternites 6—8: Fig. 15, ventral aspect. Apical spine short. Genitalia short, base and apex dark brown, median part and penis valves testaceous (Fig. 16, 17 and 18).
Arnold already doubted whether this species belongs to the subgenus Psen. The structure of the epicnemial areas and of the petiole, the depressed posterior side of the propodeum, the absence of fasciculate hairs on the apical margins of the fourth and fifth sternites, and the short genitalia make it very unlikely that P. patellatus should be placed in the subgenus Psen. As it does not fit in any of the other subgenera, perhaps a new subgenus has to be erected to receive this species.
Psen (Mimesa) empeyi spec. nov.
Male. — Head and thorax black; anterior margin of lower part of pronotum somewhat thicker and yellowish-white; apices of mandibles reddish, palpi very dark brown, antennal segments 5—12 below, last two segments completely, orange-reddish, segments 5—11 brownish dorsally. Tarsi brownish, tibial spurs of hind legs whitish. Veins of wings brown, base of veins and base of stigmata of fore wings paler brown. Petiole black, ventral plate red, also first tergite and base of second tergite, following tergites black.
No distinct frontal carina, small low tubercle between antennae. Frons and vertex densely punctate, interstices on vertex larger, at least as wide as diameter of punctures,
J. P. vAN LITH: Revision of Ethiopian Psenini 59
shining. Temples rather densely punctate, surface with fine striae. Underside of head shin- ing, occipital carina distinctly ending in hypostomal carina. Face below antennae, including clypeus, dull, finely very densely punctate, protruding median part of clypeus with small triangular emargination and slight depression (Fig. 19). Mandibles narrow. Third antennal segment nearly three times, fourth segment about twice, fifth segment about one and a half times, segments 6—8 about one and a quarter times as long as broad at apex, segments 9—12 about as long as broad at apex, last segment about one and three-quarter times as long as wide at base. Interocellar area almost impunctate, distance between posterior ocelli longer than distance between ocelli and oculi.
Sides of pronotum converging roundly backwards (Fig. 20). Scutum shining, densely punctate, interstices mostly larger than diameter of punctures, prescutal sutures indistinct, scutellum punctate as scutum, posteriorly somewhat longitudinally striate. Metanotum dull, with coarse and very close reticulate carination. Propodeum also dull, with coarser reticulate carination, coarsest on lower half. Propodeal enclosure ill-defined, propodeal longitudinal groove deep, coarsely reticulato-carinate. Sides of propodeum dull, finely closely reticulato-carinate, more striate on anterior half. Upper part of metapleura dull, lower part almost shining. Mesopleura, anterior plate of mesepisternum, greater part of epicnemial areas and mesosternum densely punctate, with narrow shining interstices; hypo-epimeral area dull, finely closely reticulato-carinate or coarsely coriaceous, epimeral suture ill-defined. First recurrent vein of fore wings interstitial, second recurrent vein ending near end of second submarginal cell. Legs slender. Petiole about one and a half times as long as first tergite, thin, cylindrical, slightly widened at base and apex, with small pit dorsally at apex. Gaster slender, finely alutaceous, last tergite dorsally flattened, broad-triangular with rounded apex (Fig. 21, 22), densely punctate with short appressed hairs.
Sixth sternite dentate apically (Fig. 23). Seventh sternite whitish transparent, tip dark brown with triangular emargination (Fig. 24). Eighth sternite whitish transparent, apical margin dark brown, spine short and dark brown (Fig. 25, 26). Genitalia dark brown: Fie227,28.
Pubescence silvery-whitish, dense and appressed below antennae.
Length about 6.5 mm.
Female unknown.
South Africa: 3 & (holotype and paratypes), Ellisras, N. Transvaal, 19 June, 1962, 1 & (paratype), 22 June, 1963, coll. H. N. Empey (holotype deposited by Mr. Empey in the Transvaal Museum, paratypes EC, PMFV and in the author's collection).
P. empeyi is the first known Ethiopian species of the subgenus Mimesa, a subgenus well represented in the arctic and Mediterranean regions. It is distinguished by the remarkable female-like pygidial area which resembles that of the Nearctic Psen (Mimesa) pygidialis Malloch (1933).
Psenulus turneri Arnold
Arnold, 1927: 121, 2 (South Africa: Cape Province). Leclercq, 1961a: 26.
Supplementary description of female. — Outer half of mandibles reddish. Fourth tergite varying from completely reddish to basally or almost completely black. All of the eleven females from Grahamstown and Kenton-on-Sea, mentioned below, have darker
60
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
J. P. VAN LITH: Revision of Ethiopian Psenint 61
legs than the holotype, the hind legs are completely black. The holotype has a completely black fourth tergite and all legs are reddish.
Frontal carina raised and much broadened between antennae, connected with high transverse carina below antennae by two closely parallel or confluent carinae. Prescutal sutures long, on anterior three quarters of scutum with short transverse carinae (alveoles). Petiole cylindrical, little longer than first tergite in dorsal aspect. No distinct pygidial area. One of the six females from Howison’s Poort, Grahamstown, has a triangular, instead of petiolate, second submarginal cell, in both wings.
Apical margin of fourth and fifth gastral sternites with dense fringe of long whitish hairs.
First description of male. — Resembling female. Mandibles almost completely reddish. Basal half of fourth tergite black in all three specimens. Hind legs partly brownish. Antennal segments 3—13 brown above; scape, second segment and underside of flagel- lum reddish.
Antennae moniliform, segments 3—8 about one and a half times, segments 9—12 about one and a third times as long as broad at apex, last segment about one and three quarters times as long as broad at base, segments 4—8 with small narrow tyloidea, segments 9—10 with indistinct tyloidea. Petiole of second submarginal cell shorter than in female. Back of propodeum coarsely reticulate, more or less striate behind enclosed area. Petiole slightly shorter than in female, below with distinct lateral keel. No conspic- uous fringes on apical margins of fourth and fifth gastral sternites.
Apex of seventh sternite with triangular emargination (Fig. 31) and fringe of long fine hairs. Eighth sternite: Fig. 29, 30. Genital apparatus (Fig. 32, 33) short, broad, pale yellowish-brown, stipes broad and rounded at apex.
The holotype is a female from South Africa, Cape Province, Mossel Bay, 18—30 Nov., 1921, coll. R. E. Turner (BM).
New records from South Africa, Cape Province: 6 9, Howison’s Poort, Grahamstown, 19—30 Nov. and 14—20 Dec., 1971, 2 9, Belmont Valley, Grahamstown, 28—31 Oct. and 5—9 Nov., 1971, all collected by F. W. Gess, 3 9, Kenton-on-Sea, 1—9 Nov., 1971, 26-31 Nov. 1971 and 1—14 March, 1972, coll. R. A. Jubb; 1 g', Howison’s Poort, Grahamstown, 24—26 Nov., 1971, coll. F. W. Gess, 3 d, Kenton-on-Sea, 23— 29 Oct., 1970, 14—21 Oct., 1971, 21—26 Oct., 1971, coll. R. A. Jubb, all collected in Malaise traps (AMG).
Leclercq (1961a) rightly assumed that P. twrneri and P. paultsae Leclercq are very closely related, but they are still distinctly different.
Psenulus paulisae Leclercq Leclercq, 1961a: 23—26, 2 and & (Zaire).
New records from Sierra Leone: 1 9, Njala, Dec., 1934, coll. E. Hargreaves, pres. by Com. Inst. Ent. B.M. 1973—1 (BM); Freetown, 4 ©, March, May, July, 1967, 3 3, Febr., April, 1968, coll. D. F. Owen (HT).
Fig. 19—28. Psen (Mimesa) empeyi sp. nov., 6, 19, holotype, 20—28, paratype. 19, face; 20,
pronotum; 21—22, 7th segment, dorsal and latero-dorsal aspect; 23, apex 6th sternite, dorsal aspect;
24, 7th sternite, latero-ventral aspect; 25—26, 8th sternite, ventral and lateral aspect; 27, genitalia,
dorsal aspect; 28, apex penis valve, lateral aspect. Fig. 29-33. Psenulus turneri Arnold, &. 29—30,
Sth sternite, with part of musculature, ventral and lateral aspect; 31, 7th sternite, lateral aspect; 32—33, genitalia, right stipes bent, dorsal and lateral aspect
62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
1 have compared these specimens with a paratype (female) from the type-locality Paulis (FAG). In the females the transverse striae on the sides of the scutum are less distinct, but in one of the males they are distinct. Leclercq has given a very good and detailed description but I would like to mention the following additional characteristics.
Female. — Anterior oblique suture of mesopleura foveolate, widened upper part with transverse carinae. Stiff hairs of fringes on apical margins of fourth and fifth sternites about as long as fourth segment of posterior tarsus.
Supplementary description of male based on the males from Sierra Leone. — Antennal segments moniliform. Scape, second segment and underside of following segments yellowish-red, dorsal side of flagellum dark brown, third, 12th and 13th segment some- what paler. Third antennal segment about one and a half times as long as broad at apex, length of segments then gradually decreasing, segments 11—12 about as long as broad at apex, last segment about one and a half times as long as broad at base. Segments 3—9 with distinct, narrow and oblique, pale tyloidea, on tenth segment much smaller. Raised part of carina between antennae somewhat narrower than in female.
There is some variation in the colour of the gaster. One of the females from Sierra Leone, collected March, 1967, has also the first tergite completely reddish, and the apex of the ventral plate of the petiole. In one of the males from Sierra Leone not only the first tergite and at least the base of the second tergite are deep black, but the following tergites also are somewhat darkened on the sides. Apical margins of sternites 2—6 brownish. Also the sculpture of scutum and propodeum may vary somewhat.
Psenulus erusus Leclercq
Leclercq, 1961a: 26—27, Q (Zaire).
New records: Sierra Leone: Freetown, 1 ©, July, 1967, 2 9, Jan., 1968, coll. D. F. Owen (HT).
These specimens are identical with the holotype from Zaire (Congo), Eala, Nov., 1936 (MRAC), but they are slightly smaller and the upper part of the back of the propodeum is less densely striate.
Length about 6.5 mm.
There are fringes of long hairs on the apical margins of the fourth and fifth sternites like in the other species belonging to the group of P. turneri.
Psenulus garambae Leclercq
Leclercq, 1961b: 57—58, 9 (Zaire).
In the holotype the dilated raised part of the interantennal carina is wide, ratio width- length being about 1 : 2.5, carina ending below antennae in a distinct transverse carina. Tempora coarsely striato-reticulate. Petiole about one and one-third times as long as first gastral tergite.
Apical margins of fourth and fifth sternites with fringe of long, yellowish golden hairs.
Length about 6 mm.
Leclercq recorded this specimen from the National Park of Garamba, in wooded savanna surrounding the camp of Bagbele, 20 March, 1950, coll. H. de Saeger (MRAC). There are no further records. He has already recognized the close relationships of P. garambae with P. saltitans Arnold, as well as with P. turneri, P. paulisae, and P. erusus.
J. P. vAN LITH: Revision of Ethiopian Psenint 63
Psenulus patei Arnold
Arnold, 1940: 132—133, £ (Rhodesia: Bulawayo and Khami). Leclercq, 1961a: 29, 9 (Rhodesia: Matopos).
Arnold based his description of this species on 6 females from Bulawayo and Khami. The holotype, which Mr. F. C. de Moor of the National Museum at Bulawayo kindly sent me on loan, is labelled: “Khami, S. Rhodesia, 30.10.1938, Nat. Museum S. Rhodesia” and also bears a red type-label of G. Arnold.
The remaining five females recorded by Arnold are undoubtedly included in the following material from the type localities: 2 9, Bulawayo “Kh” (= Khami?), 16 Sept., 1928, 1 9, Bulawayo, 25 Dec., 1929, 1 9, Khami Dam, 14 Sept., 1938, 2 9, Khami, 1 Nov., 1938, all collected by R. H. R. Stevenson (RM).
I could also study the female recorded by Leclercq (1961a) from Rhodesia: Matopos, 20 Nov., 1938, leg. et det. G. Arnold (BM).
New records from Rhodesia: 1 9, Murampara, 9 Sept., 1927, 1 9, Umguza Valley, Bulawayo dist., 1 Febr., 1930, both coll. R. H. R. Stevenson (RM).
Supplementary description, based on holotype. — Mandibles dark reddish except blackish tips. Labrum dark reddish. Palpi pale brown. Fore legs yellowish-red, trochanters completely and base of femora below brown, basitarsi yellowish-white. Mid trochanters and femora brown, apex of mid femora yellowish-red below, base of mid tibiae yellowish, underside pale reddish-brown. Hind trochanters, femora and tibiae brownish-black, basal fourth of tibiae straw-yellow, base of hind basitarsi pale yellowish. Pronotal tubercles dark brown, tegulae yellowish-transparent. Veins of wings dark brown.
Frons below antennae with tendency to transverse striation. Vertex shining, very finely punctate. Labrum not distinctly tridentate. Antennae short, third segment about one and a half times as long as broad at apex, segments 4—6 little longer than broad, segments 7—11 shorter than broad at apex, last segment about one and a half times as long as broad at base.
Prescutal sutures almost reaching hind margin of scutum, indicated only by a row of irregular short transverse grooves. No pygidial area. Apical margins of fourth and fifth sternites with fringes of long yellowish-golden hairs, sixth sternite densely yellowish- golden pubescent. Epicnemial areas below with patch of dense short whitish pubescence.
The female from Umguza Valley has darker mandibles, reddish fore tarsi, light brown mid tibiae, straw-yellowish mid basitarsi, rest of mid tarsi pale reddish, brownish-black hind legs and yellowish-brown base of hind tibiae.
P. patei is recognized by the triangular (rarely slightly open) second submarginal cell of the fore wings, striate upper part of propodeum and reddish tergites 4—6. It is closely related to P. saltitans Arnold, which also belongs to the group of P. turner? Arnold but P. saltitans has no distinct pale base of hind tibiae.
According to Arnold (1940) this species was found nesting in a decayed tree trunk.
Psenulus saltitans Arnold Arnold, 1958: 135, 9 (Rhodesia). Leclercq, 1961a: 29. Material studied: 1 Q, “Cashel, S. Rhodesia”, 30 Dec., 1947, with green paratype- label (RM). Supplementary description. — Middle part of mandibles dark reddish. Fore trochanters
64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
and at least outer side of fore femora brown, not reddish-brown. Mid basitarsi straw- yellow. Veins of wings brown.
Frons shining, oblique rugae superficial. Antennae short, third segment about one and one-quarter times as long as broad at apex, segments 4--6 little longer than broad, segments 7—11 wider than long, last segment about one and three-quarter times as long as broad at base. Interantennal carina raised and broadened between antennae, lateral edges of widened part converging below and ending as two very close parallel carinae in the transverse carina below antennae. Transverse carina more angular than figured by Arnold (1958: 134). Lower half of clypeus shining, impunctate.
Prescutal sutures as in P. patei Arnold. Anterior oblique suture foveolate, widened upper part with transverse carinae reaching upper anterior corner of episternal area. No pygidial area. Petiole about one and a half times as long as first tergite in dorsal aspect, cylindrical, base flattened dorsally.
Petiole dorsally with lateral row of very short hairs, ventro-laterally a few long out- standing hairs. Apical margins of fourth and fifth sternites with fringe of long yellowish- golden hairs, sixth sternite yellowish-golden pubescent.
P. saltitans belongs to the group of P. turneri Arnold, but differs from latter species in the non-petiolate but triangular second submarginal cell of the fore wings. It differs from P. patei in the dark bases of mid and hind tibiae.
According to Arnold this species was caught “jumping about on the leaves of a shrub, probably in search of aphids”.
Psenulus ealae Leclercq
Leclercq, 1961a: 27—29, 9 (Zaire).
New records: Sierra Leone: Freetown, 1 9, May, 1967, 1 g', April, 1968, coll. D. F. Owen (HT).
A series of seven females is known from Eala; through the kind intermediary of Prof. J. Leclercq I could study three paratypes (FAG).
Supplementary description of female based on these paratypes. — All legs brown, apex of fore femora, fore tibiae, and underside of mid tibiae more yellowish-brown, base of hind tibiae yellowish-grey. Veins of fore wings dark brown.
Transverse carina below antennae not long, reversed-V-shaped with an angle of about 100 degrees. Foreside of pronotum with transverse striae. Anterior oblique suture broad, foveolate, widened upper part with transverse carinae. Base of second sternite with semi- circular depression. No pygidial area. Apical margins of fourth and fifth sternites with fringe of long yellowish-golden hairs.
Length about 6.5 mm.
First description of male (Sierra Leone). — Resembling female. Base of fourth tergite also brown. Raised and broadened part of interantennal carina narrower, ratio width- length about 1:4. Antennal segments convex below, third segment about one and three-quarter times, segments 4—12 about one and a half times as long as broad at apex, last segment nearly two and a half times as long as broad at base. Segments 4—10 behind with distinct narrow oblique tyloidea, on segments 5—6 about half as long as segment, all reddish like underside of flagellum, on segments 3 and 11 at most a small point.
Second submarginal cell of fore wings sessile, but its upper width shorter than in
J. P. van LITH: Revision of Ethiopian Psenini 65
Fig. 34—36. Psenulus aurifasciatus sp. nov. 2, holotype. 34, face; 35, frontal (interantennal)
carina, dorsal aspect; 36, propodeal enclosure. Fig. 37—41. Psenulus freetownensis sp. nov, 9,
holotype. 37, face; 38, frontal carina, dorsal aspect (39, the same, paratype); 40, 2nd and 3rd sub- marginal cell of left fore wing; 41, propodeal enclosure
female. Petiole about one and a half times as long as first tergite. This is much longer than in the female, in which the petiole is about equal in length. However, this may be a sexual difference.
The female from Sierra Leone is somewhat smaller than the paratypes from Eala, the scutum is slightly less strongly sculptured and the fore femora are more reddish.
P. ealae is closely related to P. turner? Arnold, but is distinguished easily by the open second submarginal cell.
Psenulus aurifasciatus spec. nov.
Female. — Head and thorax black; mandibles partly brownish, palpi yellowish-brown. Underside of antennae reddish-brown. Apex of fore femora, fore tibiae completely and tarsi reddish; back-side of mid tibiae and tarsi reddish, rest of mid legs and hind legs completely black or very dark brown. Gaster black, apical margins of tergites 2—5 brownish transparent laterally; last tergite, greater part of fifth and whole of sixth sternite brown. Veins of wings dark brown, tegulae reddish-brown.
66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
Frontal carina (Fig. 35) raised and broadened between antennae, ratio width-length about 1 : 2.5, lower narrow part of carina about as long as broadened part and ending below in a transverse carina, its length about half the distance between the eyes, obtusely angular in the middle (Fig. 34). Basal half of clypeus dull, apical half shining, median part much protruding with low triangular emargination, forming two triangular teeth. Mandibles normal. Occipital carina ending in hypostomal carina. Frons near oculi with large convex, shining and very finely punctate areas, below anterior ocellus on both sides of carina very densely finely punctate. Vertex shining, very finely punctate, punctures between ocelli coarser. Tempora with coarse, striato-reticulate longitudinal carination, especially along the eyes. Antennae short, third segment about one and a half times, segments 4—5 about one and one-third times as long as broad at apex, segments 6—10 almost square, 11th segment about one and one-quarter times as long as broad at apex, last segment about twice as long as broad at base.
Pronotal corners rounded, foreside above with fine longitudinal striation. Scutum shining, finely punctate, prescutal sutures linear on anterior fourth, on posterior part until near hind margin of scutum indicated by a line of punctures larger than those on rest of scutum. Scutellum and metanotum very finely punctate, interspaces a few times as large as punctures. Lateral parts of propodeal enclosure medially separated from metanotum by two triangular, dull areas; median part broad, almost heart-shaped and connected with anterior margin by a small separate area (Fig. 36). Back of propodeum shining, dorsally with superficial oblique striae, followed by a more coarsely and some- what irregularly striate area, lower third with fine irregular reticulation and a few stronger carinae. Median longitudinal groove broad. Posterior half of sides of propodeum reticulate, separated from back-side by a broad, dorsolateral, irregularly foveolate groove; anterior part of sides dull. Metapleura dull over greater part. Mesopleura and mesoster- num shining, distinctly punctate, interstices mostly larger than punctures, punctures very fine on hypo-epimeral area, a foveolate groove along metapleura. Anterior oblique suture broad and foveolate, widened upper part with transverse carinae. Legs normal, stoutly built, a row of four small teeth on apical two-fifths of mid tibiae, hind tibiae with two very small thorns on apical half. Second submarginal cell of fore wings sessile, upper side about one-fifth of length of lower side, first recurrent vein ending in second sub- marginal cell. Petiole about one and one-quarter times as long as first tergite, cylindrical, base above distinctly, and sides only slightly, flattened, apex above with a small pit. Gaster very finely punctate. No pygidial area. Second sternite at base with a rounded- triangular depression.
Face with appressed silvery pubescence, sparse on anterior third; also with long whitish hairs. Head, thorax and legs whitish pubescent, more yellowish-grey and longer on vertex, dense and somewhat yellowish-grey on lower part of epicnemial areas and on mesoster- num. Gaster with yellowish-brown pubescence. Pygidium with many long stiff hairs, apical margins of fourth and fifth gastral sternites with fringe of long, yellowish-golden hairs.
Length 8 mm.
Male unknown.
Sierra Leone: 1 9 (holotype), Freetown, April, 1967, coll. D. F. Owen (HT).
This species belongs to the group of P. turneri and is easily distinguished by its dark gaster.
J. P. vAN LITH: Revision of Ethiopian Psenini 67
Psenulus freetownensis spec. nov.
Female. — Head and thorax black; mandibles reddish, palpi testaceous. Scape of antennae and underside of flagellum yellowish-brown, flagellum above dark-brown. Pronotal tubercles and tegulae yellowish-brown. Veins of wings dark-brown. Legs in- cluding trochanters reddish, fore and mid legs slightly paler than hind legs. Petiole, except ventral plate, black, following segments of gaster reddish.
rontal carina (Fig. 38, 39) raised and widened between antennae, length of widened part about 3.5 times width, carina ending below antennae in transverse carina, which has the shape of a reversed V with an angle of about 120 degrees (Fig. 37). Clypeus very finely and densely punctate, lower margin less densely punctate and more shining; protruding median part with two distinct small teeth, distance between apices of teeth about one sixth of total distance there between eyes. Frons and vertex shining, distinctly punctate. Tempora with coarse reticulato-striate sculpture behind eyes. Occipital carina ending normally in hypostomal carina. Mandibles normal. Third antennal segment at least one and a half times, segments 4—5 about one and a third times as long as broad at apex, segments 6—7 slightly longer than broad, segments 8—11 about as long as broad at apex, last segment about one and three quarters times as long as broad at base.
Lateral corners of pronotum obtuse. Scutum densely finely punctate, prescutal sutures not sharp, almost reaching hind margin, parapsidal sutures distinct. Scutellum and meta- notum very finely and densely punctate. Propodeal enclosure triangular, central part broad, separated from metanotum by a small square area (Fig. 41), lateral areas with at least three oblique carinae. Median longitudinal groove broad, oval, with narrow deeper groove. Upper half of back of propodeum smooth and shining, lower half with fine reticulate carination. Posterior half of sides of propodeum with reticulate carination, anterior half opaque, propodeum postero-laterally with a broad foveolate groove. Meta- pleura opaque, somewhat coriaceous, mesopleura and mesosternum shining, very finely punctate. Anterior oblique suture broad, foveolate, broadened upper part with transverse carinae. Upper side of second submarginal cell about one third length of lower side of cell, first recurrent vein ending just in second submarginal cell, second recurrent vein ending well in third submarginal cell (Fig. 40). Legs normal, mid tibiae at apex behind with 3—5 short reddish thorns, on back of hind tibiae in the middle two or three short reddish thorns.
Petiole about two-thirds of length of first tergite (in dorsal view), almost cylindrical, base flattened and with lateral carina, apex with small pit. Gaster very finely punctate. No pygidial area. Second sternite at base with semi-circular depression.
Face with silvery appressed pubescence and also with long erect silvery hairs. Pubes- cence of head and thorax whitish, epicnemial areas below with round patch of dense appressed pubescence. Petiole with long lateral hairs and dorsally with two backwardly convergent rows of very short hairs. Pubescence of legs and gaster yellowish-golden, apical margins of fourth and fifth sternites with fringe of long, yellowish-golden hairs, sixth sternite with short dense yellowish-golden pubescence.
Length about 6.5—7 mm.
Male. — As female, but underside of antennae paler, transverse carina below antennae longer and less angular in frontal aspect, segments 3—12 about one and a half times as long as broad at apex, last segment about two and a half times as long as broad at base, underside of segments convex, segments 4—8 on back with indistinct, oblique, reddish
68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
tyloidea. Circular patches of appressed pubescence at lower side of epicnemial areas less conspicuous and sternites 4—5 without fringes of long hairs. Upper, horizontal part of propodeum smooth and shining, back and sides of propodeum more coarsely sculptured than in female. Petiole on apical half below with distinct ridge, sides of petiole slightly depressed.
Length about 5.5 mm.
Sierra Leone: Freetown, 1 9 (holotype), Aug., 1967, 1 g' (allotype), July, 1967, 1 & (paratype), Jan., 1968, all collected by D. F. Owen (HT, paratype in author's collection).
Equatorial Guinea (Spanish Guinea): 1 9 (paratype), ‘“Uelleburg”, June—Aug., 1908, coll. Tessmann (ZMB).
In the allotype the second submarginal cell of the left wing is almost triangular. On the contrary this cell is more open above in the male paratype than in the holotype.
The petiole of the female from Equatorial Guinea is brownish, its apex reddish dorsally. The interantennal carina is slightly broader and the upper side of the second submarginal cell is about one fourth of the length of the lower side.
P. freetownensis is in both sexes easily recognized by the completely red gaster and legs, and the broad smooth upper part of the propodeum behind the enclosed area. It evidently belongs to the group of P. turneri, of which some species normally have triangu- lar or even petiolate second submarginal cells.
Psenulus alveolatus spec. nov.
Male. — Head and thorax black; antennae brown, underside and scape completely reddish-brown. Palpi pale brown. Tegulae dark reddish-brown. Labium and apical half of mandibles reddish-brown. Fore legs including trochanters reddish-brown, mid and hind legs brown. Tibial spurs of mid and hind legs yellowish-white. Petiole including ventral plate deep black, also first tergite and basal two-thirds of second tergite, rest of dorsal side of gaster reddish; sternites 2—4 very dark brown, nearly black, following sternites reddish; apical spine dark reddish. Veins of wings dark brown.
Frontal carina raised and much broadened between antennae, ending below antennae in high transverse carina. Clypeal margin medially with two low triangular teeth. Frons with coarse oblique rugae, interocellar area with transverse rugae, vertex more shining, striato-punctate, striae weak. Tempora coarsely striato-reticulate with a few strong longi- tudinal carinae. Occipital carina ending in hypostomal carina, high, alveolate along fore- side. Antennal segments 3—8 about one and a half times, segments 9—12, about one and a quarter times width at apex, last segment one and three quarters times width at base; third segment with small tyloidea, segments 4—8 with longer and distinct, narrow tyloidea.
Anterior margin of pronotum alveolate; scutum shining, with irregular rather dense punctation, and centrally with weak transverse rugae; prescutal sutures with deep and large alveoles and strong transverse rugae, reaching posterior margin which shows many short longitudinal rugae; parapsidal sutures distinct; lateral margins of scutum alveolate. Scutellum shining, sparsely finely punctate. Metanotum somewhat more densely punctate. Enclosed area of propodeum shining, with rather short oblique carina on lateral parts, wide central area; back coarsely reticulate, median longitudinal carina deep and wide but irregular. Sides of propodeum less coarsely reticulate. Metapleura smooth. Mesopleura
J. P. vAN LITH: Revision of Ethiopian Psenini 69
and mesosternum shining, densely coarsely punctate, partly somewhat rugose. Anterior plate of mesepisternum shining, less densely punctate, not rugose. Anterior oblique suture broad, foveolate, widened upper part with transverse carinae.
Legs slender. Second submarginal cell distinctly petiolate, first recurrent vein ending in second submarginal cell, second recurrent vein ending well in third submarginal cell. Petiole long, nearly twice length first tergite; sides slightly depressed, with distinct lower keel, dorsally convex, but base and apex somewhat flattened, narrow triangular pit at apex.
Face with appressed silvery pubescence and erect longer hairs, rest of head, and thorax, with whitish hairs, gaster yellowish-grey pubescent, apical margins of sternites with thin fringe of short, backwards directed stiff hairs, disk of sixth sternite with dense, back- wards directed, golden pubescence.
Length about 7 mm.
Female unknown.
Rhodesia: 1 & (holotype), Bulawayo, 25 Dec., 1929, coll. R. H. R. Stevenson (RM).
P. alveolatus is certainly closely related to P. twrneri Arnold, differing however by having a longer petiole, a more coarsely sculptured scutum, and by the fact that more gastral segments are reddish.
A male from Angola (A30), 7 miles W. of Gabela, 16—18 March, 1972, Southern African Exp. B.M. 1972—1 (BM) is very similar and certainly belongs to the group of P. turneri. It has also a long petiole and tergites 2—6 are completely reddish, but it differs from alveolatus in being somewhat smaller, with the scutum less coarsely sculp- tured and the second submarginal cell being open dorsally. As the antennae are missing I prefer to wait for additional material before describing this form as a new species.
Psenulus fulgidus Arnold
Arnold, 1945: 156—157, ® and 4 (Malagasy: Bekily, Behara and Ranomafana). Leclercq, 1961a: 35, 9 (Malagasy: Fort-Dauphin).
Material studied: 2 9, Bekily, Dec., 1932, Febr., 1940, 1 &, Bekily, May, 1934, (FAG); 1 9, Bekily, March, 1937, coll. A. Seyrig, det. Arnold (?) (author’s collection).
The material from the collections of the Faculty of Agricultural Sciences at Gembloux was kindly sent to me by Prof. J. Leclercq; it forms part of the material collected by Seyrig and was mounted after Arnold’s study (1945) and identified by Leclercq (1960).
P. fulgidus and P. reticulosus Arnold are much alike as regards their colour patterns, but they differ in a few important characteristics. P. fulgidus is smaller (female 6.3—8.5 mm, male 6—6.5 mm, cf. Arnold, 1945, who based his description on 16 females and 9 males). In the female of P. fulgidus the broadened raised part of the interantennal carina is lozenge-shaped, petiole about one and one-third times as long as first tergite, apex of petiole about twice as wide as base, pygidial area with distinct lateral carinae, parallel on apical half, gradually diverging towards base. I have seen one male, recorded above, from Bekily (det. J. Leclercq). This differs from the male of P. reticulosus, apart from being smaller, in having no tyloidea. Raised part of interantennal carina hardly broadened. Arnold described the punctation of the scutum as being denser than in the female, inter- spaces between larger punctures about twice as wide as punctures themselves and some specimens with a trace of transverse rugosity. He did not mention the interantennal carina.
70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
The above male has a strongly transversely rugose scutum, apical margin of scutellum yellow, remaining part brown and yellowish-brown. Petiole about one and a half times as long as first tergite.
Psenulus reticulosus Arnold
Arnold, 1945: 154—156, 9 and & (Psenulus reticulosus and Psenulus reticulosus var. albo- scutellatus; Malagasy: Rogez, Perinet, Ivondro, Ranomafana).
Leclercq, 1961a: 35, 2 (Malagasy: Mandraka, Rogez, Analandraraka).
Leclercq, 1961c: 117, 9 (Malagasy: Ranomafana, Perinet, Joffre-ville).
Material studied: Rogez, 1 9, Jan., 1931, 1 2,1932, 1 S, April, 1931 (FAG); 1 9, Rogez, “forêt côte est”, Febr., 1934, coll. A. Seyrig (author's collection); Rogez, 1 9, Dec., 1931, 1 9, 1932, 1 g', May, 1932 (FAG). The last mentioned three specimens belong to the var. alboscutellatus Arnold.
Petiole over one and a half times as long as first tergite, in dorsal aspect. Prescutal sutures distinct only on basal third of scutum. Raised part of interantennal carina slightly broadened in female, hardly or not broadened in male.
Pygidial area of female narrow with indistinct, parallel, carinae.
Underside of fore and mid femora pale brown in male, in female fore and mid legs completely pale yellow. Antennal segments 3 (or 4)—13 with distinct tyloidea, which are reddish like flagellum.
Psenulus sapobaensis spec. nov.
Female. — Head and thorax black; mandibles yellowish with brown tips, labrum reddish-brown, palpi testaceous. Scape of antennae yellowish, following segments dark brown above, last segment reddish-brown above, underside of antennae yellowish-red. Pronotal dorsum medially, and metanotum, blackish-brown. Posterior part of pronotal tubercles yellowish, tegulae yellowish-red. Fore and mid legs including trochanters and mid coxae pale yellowish-red or straw-yellow; hind legs including coxae darker yellowish- red. Veins of wings dark brown. Gaster including petiole yellowish-red.
Frontal carina distinct, raised part between antennae very slightly broadened (Fig. 43), ending below antennae in a distinct transverse carina. Supraclypeal area and basal half of clypeus superficially densely punctate, apical half of clypeal disk not very shining but without distinct sculpture or punctation. Protruding median part of clypeal margin distinctly bidentate, distance between apices of teeth about one-ninth of total distance there between the eyes (Fig. 42). Frons below anterior ocellus with distinct separate punctures, near oculi almost smooth. Vertex shining, sparsely and minutely punctate. Tempora with fine longitudinal striae. Occipital carina ending normally in hypostomal carina. Mandibles normal. Third antennal segment about two and one-third times, fourth segment about twice, fifth segment about one and three-quarter times, segments 6—8 about one and a half times, segments 9—11 about one and one-third times as long as broad at apex, last segment at least twice as long as broad at base (all in frontal view).
Pronotal corners sharp. Scutum fairly strongly punctate, finer on anterior part between prescutal sutures. Prescutal sutures distinct on anterior two-fifths of scutum, continued until near apical margin by a row of relatively large punctures. Scutellum and metanotum scarcely finely punctate. Propodeal enclosure triangular, well-defined, median part rather
J. P. VAN LITH: Revision of Ethiopian Psenini 71
47 46
Fig. 42—44. Psenulus sapobaensis sp. nov., 2, holotype. 42, face; 43, frontal carina, dorsal aspect; 44, propodeal enclosure. Fig. 45—47. Psenulus leoninus sp. nov., 9, holotype. 45, face; 46, frontal carina, dorsal aspect; 47, pygidial area
wide with median longitudinal carina (Fig. 44), lateral parts with few oblique carinae. Upper part of propodeum with narrow smooth and shining area, back with fine oblique striae, transverse on lower part. Dorso-lateral edge of propodeum more coarsely reticulate, posterior part of sides finely reticulate, anterior part smooth. Metapleura shining, some- what striate below, mesosternum and mesopleura smooth, with scattered punctures. Anterior oblique suture foveolate, widened upper part smooth. Legs stout, fore and mid femora and tibiae rather heavily built, apex of mid tibiae on outer side with narrow longitudinal depression, margined anteriorly by a reddish oblique low carina and behind by four short reddish spines. Hind tibiae on outer side with three (left tibia) or two (right tibia) longer spines. Second submarginal cell above about half as wide as lower side. First recurrent vein ending just in second submarginal cell, second recurrent vein ending well in third submarginal cell, about one-fifth of base of third submarginal cell. Petiole long, slightly more than one and a half times length of first tergite, cylindrical, apex dorsally with small triangular pit. Gaster minutely punctate, no well-defined pygidial area, just a narrow smooth and shining area bordered by a few long hairs. Base of second sternite somewhat depressed.
Pubescence of face and tempora silvery, appressed, leaving structure of clypeus well visible, face also with a number of long erect hairs. Vertex with long greyish pubescence.
72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 117, AFL. 3, 1974
Pubescence of thorax whitish, long on propodeum, dense on mesosternum. Epicnemial areas below with round patch of dense appressed silvery pubescence. Legs and gaster with yellowish-grey pubescence, denser and rather long on apical margins of fourth and fifth sternites but not fringe-like, sixth sternite densely yellowish-golden pubescent. Petiole with a few long, obliquely directed downwards, hairs, dorsum bare.
Length about 9 mm.
Male unknown.
Nigeria: 1 ® (holotype), Sapoba, 16 Aug., 1962, coll. D. C. Eidt, in Malaise trap (CNC).
This species is probably closely related to P. reticulosus.
Psenulus leoninus spec. nov.
Female. — Head and thorax black; mid part of mandibles brownish, palpi testaceous, antennae dark-brown, scape and underside of following segments reddish-brown. Pronotal tubercles slightly brownish, tegulae reddish. Fore and mid legs reddish, including tro- chanters, mid femora brown below, fore and mid basitarsi yellowish; hind trochanters brown, hind femora brown, apical two-thirds reddish below, rest of hind legs also reddish. Veins of wings dark-brown. Petiole including ventral plate black; gaster reddish, first tergite with large black mark almost reaching hind margin.
Frontal carina raised and widened between antennae, about two and a half times as long as broad (Fig. 46), ending below antennae in a short transverse carina, in the shape of a reversed V with obtuse angle. Face below transverse carina dull, finely aciculate; clypeal margin ending in two distinct small teeth, distance between these teeth about one eighth of total distance there between eyes (Fig. 45). Mandibles normal. Occipital carina ending in hypostomal carina. Frons densely punctate, vertex between eyes with more widespread punctures. Tempora aciculate, partly finely striate. Antennae short, third segment about one and a half times, segments 4—6 about one time, segments 7—9 about two-third times, segments 10—11 about half as long as broad at apex; last segment about one and a third times as long as broad at base.
Pronotal lateral corners with sharp anterior angle. Scutum finely densely punctate, relatively few very large punctures and on sides of anterior half transversely striate. Prescutal sutures almost reaching hind margin of scutum, rather ill-defined. Scutellum and metanotum with sparse fine punctures. Propodeal enclosure narrow, lateral parts with a few oblique carinae. Median longitudinal groove wide, with transverse carinae. Upper part of back of propodeum immediately behind enclosed area finely obliquely striate, lower half and posterior two-thirds of sides of propodeum with irregular reticulate carina- tion; anterior part of sides dull. Metapleura shining. Mesopleura and mesosternum with few fine punctures. Anterior oblique suture coarsely foveolate, widened upper part with a few transverse carinae. Legs normal, flattened apical outer side of mid tibiae with short row of three small reddish teeth. First recurrent vein of fore wings interstitial, upper side of second submarginal cell about half length of lower