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TIJDSCHRIFT 
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UITGEGEVEN DOOR 


DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING 


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DEEL 114 AFLEVERING 8 1971 


TIJDSCHRIFT 
VOOR ENTOMOLOGIE 


UITGEGEVEN DOOR 


DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING 


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| <hrift voor Entomologie, deel 114, 1971 | 


Aflevering 1 verscheen 15 juni 1971 
Afleveringen 2—4 verschenen 27 september „ 
5—8 5 29 december ,, 


LE 


INHOUD VAN DEEL 114 


HOLLANDER, J. DEN. — Life histories of species in the Pardosa pullata group, 
a study of ten populations in the Netherlands (Araneae, Lycosidae) . 


KRIKKEN, J. — New species of the Papuan genus Tafaza Valck Lucassen (Coleo- 
ptera, Cetoniidae) 


LEMAIRE, CLAUDE. — Descriptions d’Attacidae (Saturniidae) nouveaux d’Ameri- 
que Centrale et du Sud (Lepidoptera) . 


LIEFTINCK, M. A. — Studies in Oriental Corduliidae (Odonata) I. 


LIEFTINCK, M. A. — A catalogue of the type-specimens of Odonata preserved 
in the Netherlands, with a supplementary list of the Odonata types described 
by Dutch scientists deposited in foreign institutional collections . 


LukoscHus, F. S. & F. M. DRIESSEN. — Amorphacarus parvisetosus spec. nov. 
(Myobiidae, Trombidiformes) from Neomys fodiens Pennant (Soricidae) . 


LukoscHus, F. S., A. W. A. M. DE Cock & A. Fain. — Life cycle of Meleso- 
dectes auricularis Fain & Lukoschus (Glycyphagidae, Sarcoptiformes) . 


LUKOSCHUS, F. S., A. W. A. M. DE Cock & F. M. DRIESSEN. — Four new 
species of the genus Psorergates Tyrell, from European hosts (Acarina, 
Psorergatidae) 

ROSSEM, G. VAN. — The genus Buathra Cameron in Europe (Hymenoptera, Ich- 
neumonidae) 

ROSSEM, G. VAN. — A new species of Cryptus from the Canary Islands (Hy- 


menoptera, Cryptinae) 


ROSSEM, G. VAN. — Additional notes on the genus Trychosis Foerster in Europe 
(Hymenoptera, Cryptinae) . 


THEOWALD, B. — Die Tipuliden der Benelux-Länder (Diptera, Tipulidae) . 


Register 


255 


163 


173 


185 


201 


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DEEL 114 AFLEVERING 1 MUS. COMP. 24931. 
LIBRARY 


JUL 30 1971 


HARVARD 
UNIVERSITY 


TIJDSCHRIFT 
VOOR ENTOMOLOGIE 


UITGEGEVEN DOOR 


DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING 


INHOUD 
M. A. LIEFTINCK. Studies in Oriental Corduliidae (Odonata) I, p. 1—63, Fig. 1—50. 


Tijdschrift voor Entomologie, deel 114, afl. 1 Gepubliceerd 15-VI-1971 


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nn eS eut té 


STUDIES IN ORIENTAL CORDULIIDAE (ODONATA) I 
by 


M. A. LIEFTINCK 
Rhenen, Netherlands 


ABSTRACT 


An attempt is made to prove the incongruity of attributing family rank to the Macromia alliance 
and the classification of Corduliinae as a subfamily of Libellulidae. Instead of this, arguments 
are put forward in favour of restoring and maintaining, the group as a subfamily of Cor- 
duliidae. Idionychidae and Synthemistidae are regarded as groups so closely affiliated with 
Macromiinae that their family status is called in question. Contrasting characters are tabulated 
of Idiophya, Idionyx and Macromidia. The male of Idionyx philippa Ris and both sexes of 
I. murcia spec. nov. (Sumbawa) are described and a key is constructed for the S. E. Asiatic species, 
followed by a description and figures of the larva of I. montana Karsch. Some taxa in Macromidia 
are re-characterized and M. asahinai spec. nov. (Palawan) is added to the list. In Macromia several 
species are discussed and the list of Malaysian taxa is made up to date; new species are M. dione 
(Sumatra), and in the Papuan group M. lachesis (Bismarcks) and M. astarte (S. E. New Guinea). 
Lastly, a review is given of Synthemis in New Caledonia, with definitions of S. campioni spec. nov., 
the females of S. fenella Campion and S. montaguei Campion hitherto unknown, and a key to all 
insular species. Descriptions and illustrations of two New Caledonian Synthemis larvae lead to 
comments on the acquisition of adaptive structural features during larval development. 


Contents 
I. Remarks on the subfamily Macromiinae . . . 1 
II. Notes on the Idionyx-Macromidia alliance, with an account en ie Iba Br 
Idionyallagen 2. SP RE da cc 2 
III. Some Malaysian species of Mann Ramıpur ERBE N co WED ys se 22 
IWeeebhe Papuasian group of Macromia Rambur. „os on .. +. . 30 
V. Remarks on the subfamily Synthemistinae . . 45 
VI. Notes on Synthemis Selys from New Caledonia, ns an account RICE eee 
larvae. NES BR SP © Sieh ad Alten 0 CAS 
VII. Material and ruben Oee Een IAN O1 
VIT Selen Tal EEEN Te gn de, Meden GD 


I. REMARKS ON THE SUBFAMILY MACROMIINAE 


In a historical survey of the “Macromia Group” of genera, a new family name, 
Macromiidae, was proposed by Mrs. L. K. Gloyd (1959). The diagnosis was based 
primarily on three genera, Macromia, Didymops and Epophthalmia, but the author 
added that the venational characteristics, at least, seemed to apply to Macromidia also. 
It was admitted that the features enumerated were not fully checked for species in all these 
genera, while the author also felt that her diagnosis and the composition of Macromiidae 
might require revision with further study. It is not the purpose of the present paper to 


1 


2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


attempt any definitions of suprageneric taxa within the Corduliidae; but I merely wish 
to point out that the removal of the Macromia group from the Corduliidae and its 
elevation to family rank is based on the forsaken conception that “Corduliinae and 
Libellulinae are so closely related that they belong in one family, the Libellulidae”, to 
quote Mrs. Gloyd’s own words. I am unable to share this view, adhering instead to the 
well-considered arguments put forward by St. Quentin (1939) 1), in whose publication 
the reader will find substantial evidence in support of the view that Corduliidae and 
Libellulidae are different families, each with well-founded and unmistakable characters. 
In fact only few of the basic characters separating the extremely diversified Corduliidae 
from the Libellulidae were mentioned by Gloyd; she apparently failed to observe that 
her diagnosis, apart from the Macromia group, applies equally well to many (other) 
corduliid genera. Under these circumstances it is only natural that the author’s family 
diagnosis includes but few characters serving to distinguish the Macromia group from 
its nearest relatives, i.e. the remainder and major part of the corduliids. Genera like 
Macromidia and Synthemis are mentioned only in passing, and the reader is left in doubt 
as to which of the rest of the corduliid genera the author prefers to leave in her Libellu- 
lidae (“including the Corduliinae, or Corduliidae of some authors’) and which are 
considered “Macromiidae”. In any case it would seem to me to be irrelevant to construct 
a family diagnosis upon characters incompatible with the morphology of the remotely 
allied Libellulidae, an independent family which, in its restricted sense as accepted by 
most present-day authors, omits the Macromiinae and all the rest of the Corduliidae. As 
to the morphology and anatomy of the corduliid larvae, I believe to have shown earlier 
(Lieftinck, 1950, 1952) that the remarkable diversity of structure, notably in tropical 
species of Macromia, is unmistakably correlated with a particular mode of life, many 
instances being known of larvae having modified their entire organization to adapt them- 
selves to some peculiar environment. In the case of Macromia this is especially striking 
as whole sets of morphological adaptations go hand in hand with varying environmental 
conditions. Thus larval features like the development of posterolateral tubercles at the 
head, shape of labial ligula, gizzard armature, as well as palpal and mandibular dentition, 
are to a large extent so variable within the limits of each genus, that they can at best be 
used as group characters, certainly not as a means of subfamily division or even generic 
separation. 


II. NOTES ON THE Idionyx - Macromidia ALLIANCE, WITH AN ACCOUNT OF THE LARVA 
OF Idionyx HAGEN 


Idiophya Fraser, 1934 


Idiophya was proposed by Fraser (1934: 553—554, fig. 1, wings © ) for a single 
enigmatic corduliid, first described as Phyllomacromia nilgiriensis Fraser (1918: 383— 
384), and known only from seven females, all collected at the Burliyar river in the 
Nilgiri Hills (South India). The author subsequently (1957) expressed doubt as to the 
validity of a monobasic genus comprising a species of which the male had never been 
discovered, the more so because Idiophya was stated to differ from Idionyx only by 


1) With many other judicious publications of fundamental importance simply ignored in the revised 
edition of F. C. Fraser’s “Reclassification of the Order Odonata” (1957). 


M. A. LIEFTINCK: Studies in Corduliidae I 3 


having the costal side of the discoidal triangle of the fore wing fractured, “although this 
feature is not always present” (Fraser, loc. cit, 1957: 111). Indeed, the excellent wing 
photograph of I. nilgiriensis in Fraser's 1934 publication shows a venation so similar to 
that of Idionyx that Idiophya could easily be regarded as a slightly aberrant member, 
were it not that a second species had been recorded from Luzon I. (Philippine Is.). This 
is I. salva Needham & Gyger (1937: 57—58, pl. 1 fig. 16, Q genit. & pl. 3 fig. 58, 
Q wings), once again described after a single female and for that reason placed doubt- 
fully in Idiophya. Through the kindness of Dr. Pechuman I have been able to examine 
this immature specimen. (CUI). Owing to the fact that it was preserved in poor alcohol, 
the body has fallen to pieces, lost all colours and become practically unrecognizable. 
There are a pair of wings on a slide, evidently the same as those reproduced on pl. 3 
fig. 58 in Needham’s publication. 

Needham (loc. cit.: 58) refers to two characters that would serve to distinguish his 
Idiophya salva from Idionyx philippa. In the former the single row of cells beyond the 
4-sided triangle of the fore wing continues beyond the level of the nodus, whereas in 
I. philippa beyond a 3-sided triangle the cell-row is doubled well before the level of the 
nodus. This is true only for the right fore wing of philippa (the one figured by Ris), 
the double row of cells on the left wing beginning only a single cell-breadth before that 
level. The second character refers to the course of the veins Cu, and Cz, on the fore 
wing, which in I. salva diverge to the wing margin while in I. philippa they converge. 
In the left pair of wings of the type of I. philippa (not photographed by Ris), these 
two veins run parallel to each other up to a point two cells before reaching the anal 
margin, at which point Czy gives off two weak branches, one running parallel to Cv, 
the other curving away from it and meeting the anal margin at a right angle. This is 
obviously also a variable character, for not one of the other examples of philippa is 
exactly alike in this respect. 

Yet it is impossible to associate Idionyx philippa Ris with this dragonfly. The venation 
is very similar, but the wings of I. salva appear more broadened midway their length 
and more pointed than in J. philippa, the type of which possesses a normal fore wing 
triangle. The yellow spots at the sides of the thorax seem to be arranged differently in 
the two species, philippa having no “long triangle pointing downward covering a good 
part of the metepimeron”’; the femora of the latter are black, not pale “thrice faintly 
ringed with brown”, as described for I. salva. It would appear, therefore, that salva and 
philippa are specifically distinct, although the remaining venational differences between 
the two, enumerated by Needham at the end of his description, are nonapparent (see 
under I. philippa). 

It would be interesting to know more of the morphology of the occipital region and 
genital organs of the females of Idiophya since little information has been given about 
these structures in the existing descriptions. As to I. nilgiriensis, Fraser only says that 
the vulvar scale is “small, triangular, not visible in profile”, while the sketch of the 
subgenital plate of I. salva, accompanying Needham’s account, suggests Idionyx. 

Considering the above, it remains impossible to establish the generic status of the two 
species described in Idiophya. All the same, for each of them we may expect a male 
resembling Idzonyx more closely than any other genus, because the venation in nearly all 
respects corresponds with this, not with Macromidia. 


4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Idionyx Hagen, 1867 


This genus was first placed in a subfamily of its own, viz. Idionychinae, by Tillyard 
& Fraser (1940), only the aberrant Idiophya Fraser (see above) also forming part of it. 
The characters employed to distinguish Idionychinae from Macromiinae (= Epophthal- 
miinae Fraser et auct.) ate such that they can be applied only to a limited number of 
representatives of either group, and then only when the extremes of both are taken into 
account. The differences between some of the larger species of Idionyx, i.e. those which 
are at home in continental south-east Asia, and the smaller-sized and slenderly built 
members of Macromia inhabiting the Oriental tropics, are certainly of no greater 
importance than those separating the latter from the sturdily built members of Epophthal- 
mia, which no one would exclude from the Macromia assemblage. These characters in the 
writer's opinion are of generic rather than subfamilial or even tribal value. I do not 
hesitate, therefore, to discard Idionychinae and follow the example set by de Selys Long- 
champs (1878), who included Idionyx in the Macromia group of genera; if de Selys had 
also known Macromidia, he would certainly have added this genus too. In point of fact 
Macromidia, as we will see, neatly bridges the gap between Macromia and Idionyx, so that 
nothing can be gained by placing the last-mentioned genera in two separate subfamilies. 

The larva of Idionyx, described below, shows many characters found also in other 
macromiines and bears a close resemblance to that of some sand-dwelling species of 
Macromia (Fig. 5). 

In the Malaysian Subregion and ‘Wallacea’ the genus is represented only by a limited 
number of species, grouped around I. yolanda Selys, 1871, the type-species, first des- 
cribed after a female from Singapore. As has been pointed out by Lieftinck (1939), 
I. dohrni Krueger, 1899, from Sumatra, and its “subspecies” I. dohrni borneensis Laid- 
law, 1913, from Borneo, are both synonyms of I. yolanda Selys. A second mem- 
ber of the group is I. montana Karsch, 1891, originally described from Java but sub- 
sequently discovered also in Sumatra and the Mentawei Is. A third species, I. philippa 
Ris, 1912, was described from the Philippines; this was known only from a single female 
but a characterization of both sexes will now be found in the next pages. A fourth 
regional species is I. orchestra Lieftinck, 1953b, which was reported from Sumba I. (Lesser 
Sunda Is.). Lastly, a species new to science, I. murcia spec. nov., from the island of 
Sumbawa, can be added to the list. Leaving aside the nondescript Idiophya salva, dis- 
cussed before, this brings the number of southeastern Idionyx up to five. 1) 

The distribution of these species, so far as at present known, is as follows: 

I. yolanda Selys (= dohrni Krueger) — Malay Peninsula (terr. typ.) ; Sumatra; Bil- 

liton; Borneo; Basilan I. (Philippine Is.) ; Hongkong. 

I. montana Karsch — Java (terr. typ.); Sumatra; Mentawei Is.; Malay Peninsula (?). 

I. orchestra Lieftinck — Sumba I. (Lesser Sunda Is.). 

I. murcia spec. nov. — Sumbawa I. (Lesser Sunda Is.). 

I. philippa Ris — Mindoro, Leyte and Mindanao Is. (Philippine Is.). 

These species have the following characters in common: 

Head large, considerably broader than thorax; eyes globular, broadly contiguous, 
median eye-line longer than occipital triangle. This triangular area raised perpendicularly 


1) I agree with Fraser (1936a) that I. laidlawi Fraser, known only from the solitary female collected 
in Pahang (Malay Peninsula), belongs to a different species group, viz. that of I. optata Selys. 


M. A. LIEFTINCK: Studies in Corduliidae I 5 


above level of eyes, finely pointed anterad with sharply acute side margins; smooth 
dorsal surface clothed with long erect hairs and vertical surface also with fringe of very 
long hairs behind rounded posterior margin. Vertex low and broad, evenly convex and 
of simple structure in both sexes. Labrum and anteclypeus at least partly chrome. Syn- 
thorax with or without incomplete yellow mesepisternal (juxtahumeral) spot, with 
yellow lateral spots or stripe at level of metaspiracle, and with variable spots or complete 
stripe on posterior portion of metepimeron. Legs at least with hind tibia partly yellow; 
all tibiae with membranous distal keel on flexor surface, the one at mid tibia very short; 
tarsal claws acute, bifid, the inferior tooth stronger and usually slightly longer than the 
apex (Fig. 12). Es 

Wing neuration open. Nodus placed far distad, space between nodus and pterostigma 
on fore wing only half as long or less than that between nodus and wing base. An- 
tenodal coplex between C and R + M complete: no distinct primary (thickened) 
antenodals. Fork of M,-„—M3 strongly asymmetrical in fore and hind wing. Only the 
first, or first two, postnodal nervures of second series missing in all wings. Fore wing 
triangle placed far beyond arculus, distinctly smaller than that of hind wing, equilateral 
but costal side very rarely slightly fractured (5-sided) in one of the wings; hind wing 
triangle also distal to arculus, but distance separating it from arculus invariably shorter 
than its proximal (shortest) side. Subtriangle (1) in fore wing nearly always irregular, 
its proximal side (distal Cux) very oblique, forming a 4- or 5-sided cell with proximal 
side of triangle, hence only 1 transverse Cxx in fore wing (very rarely an additional 
cross-vein present in one of the wings); 2 transverse Cux in hind wing. Discoidal field 
of fore wing subparallel-sided, with single row of cells as far as level of nodus, or even 
further, thereafter frequently slightly expanded. Rsp/ present, its course mostly 
slightly fractured. Anal loop of hind wing elongate, fully twice longer than broad 
without toe-like prolongation, midrib slightly zigzagged, apex obtuse, not extending 
beyond level at apex of triangle. Proximal side of anal triangle of male hind wing 
strongly sinuous; cross-vein in triangle placed well beyond midway its length; anal angle 
rounded. Pterostigma small, less than 2 mm long but usually overlying two cross-veins. 
Membranula large, in male extending as far as apex of triangle or a little beyond. 

Abdomen slender, slightly spindle-shaped; colour black with metallic gloss on proximal 
tergites. Yellow markings reduced to lateral and lateroventral spots and streaks on tergites 
1—3 and 7—9 only. Male superior anal appendages long and slender, longer than segm. 
9 + 10; appendix inferior subequal to or slightly shorter than superior pair. Branches 
of posterior genital hamulus subequally long, outer branch thick and strong, the inner 
branch a slender sickle-shaped hook. Female with apex of 8th sternite decidedly prom- 
inent in lateral view, longitudinally carinate, not projecting beyond apex of 8th tergite; 
9th sternite long and likewise carinate; styli tubercular; supra-anal plate declivous and 
tapered; anal appendages shorter than 10th segment, conical. 


It must be stressed that the above diagnosis applies only to the Malaysian species- 
group, the great length of the male appendages, for instance, being no peculiarity of 
all: on the contrary, several continental Asiatic species exhibit appendages distinctly 
approximating to the type found in Macromia. See also Laidlaw (1912) in his discussion 
of Metaphya Laidlaw. 

Apart from the fact that Idionyx and Macromidia have several characters in common, 
it should not be forgotten that interspecific variation is considerable in both, a circum- 


6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


stance impeding their distinction from Macromia, which itself is far from homogeneous 
in character. To demonstrate the existing difficulty to construct an all-embracing diagnosis 
for each of these three genera, it is only necessary to consult Fraser's “Revision of genus 
Idionyx” (1934). In that paper not less than six definite groups are recognised, and 
from a consultation of the group characters employed it is obvious that corresponding 
characters can be used to split up the large genus Macromia. This indicates, in my 
opinion, that the two genera are genuinely related. Another peculiarity pointing in the 
same direction is found in the structure of the tarsal claws, which in all regional Idionyx 
are shaped similarly to those in certain groups of Macromia (see under that genus). 

The five species presently included can be distinguished from each other by means of 
the following 


Key to the males of Idionyx 


1. Front of synthorax (mesepisterna) on either side with sharply defined, oval yellow 
juxtahumeral spot equal in size to and confluent with adjacent mesinfraepisternal 
yellow spot. Postclypeus bright yellow in middle. Sup. anal apps. in dorsal view at 
first slightly outbent, then gradually incurved, each bearing a minute subapical 
internal tooth followed by a shallow emargination, the apex somewhat swollen and 
downcurved, with small external fringe of longish, erect, golden brown hairs. Inf. 
app. with lateral prominence situated at about 24 length from base, its terminal 
portion broadened towards apex, which bears a deep V-shaped emargination, the 
branches being upturned and pointed (Fig. 4). Dorsum of 10th abdominal segment 
with distinct, though low, median carina on each side of a depression, but lacking 
a triangular boss. Main body of posterior genital hamule swollen, its outer border 
in side view distinctly convex proximal to the hook-like inner branch (Fig. 3). 
Apicoventral border of abdominal tergite 7 slightly protuberant, end portions of 
7 and 8 as well as sternite 8 sparsely clothed with soft erect yellowish pubescence. 
A complete, broad, almost parallel-sided yellow stripe at thoracic sides crossing the 
spiracle, continuous ventrally over metinfraepisternum; a similar, though slightly 
broader stripe, widest at middle, upon posterior portion of metepimeron. Larger 
species: abd. + app. 31.7 mm, hind wing 30.0 mm. Hab.: Sumbawa . . murcia 


—. Front of synthorax either entirely metallic green or lower lateral area non-metallic 
and filled out diffusely with reddish-brown. Postclypeus wholly brown or black . 2 

2. Inferior appendix very slender, simply tapered or with vestige only of a marginal 
tubercle placed slightly beyond halfway its length; apex gently upcurved, tip flattened 
dorsally, terminating abruptly in a short bluntly triangular point. Sup. anal apps. 
shaped much as in I. murcia but subapical internal prominence barely indicated and 
followed by a shallower emargination, the knob-like apex of each bearing a con- 
spicuous external tuft of long golden brown hairs. Dorsum of 10th abdominal 
tergite raised, the middorsal carina replaced by a low triangular boss. Main body of 
posterior genital hamule in side view with straight outer border, but with tubercular 
swelling at base of hook-like inner branch. Apicoventral border of 7th abdominal 
tergite swollen and markedly projecting ventrad, clothed with conspicuous tuft of 
long erect golden hairs. Yellow stripes at thoracic sides complete, though narrower 
than in I. murcia, the spiracular stripe slightly more undulated. Larger species: abd. 


M. A. LIEFTINCK: Studies in Corduliidae I 7 


+ app. 31.0—33.0 mm, hind wing 29.0—31.7 mm. Hab.: West Malaysia 
A montana 
5 Inferior arend ende, ies sio roi ad Beus a distinct marginal 
prominence or conspicuous tooth placed on either side beyond halfway its length. 
Erect pubescence at ventral borders of 7th abdominal tergite and apices of sup. anal 
apps. more or less ee but hairs shorter and less closely set than in I. mon- 
VED on 3 u CEE ES O) 
. Sup. anal ms Aden at hass, but at a Sei SI ti, then 
gradually incurved, each bearing a tiny subapical internal prominence followed by 
a shallow emargination, the apex slightly swollen, curved inward and downward, 
its extremity subtruncated or bluntly rounded. Inf. app. with distinct tooth-like 
lateral projection a little beyond halfway its length, the terminal portion slightly 
expanded before the upturned apex, which is bifid, the points shorter and more 
closely approximated than in I. murcia and separated by a crescentic emargination. 
Genitalia much as described for I. murcia. Middorsal crest of 10th abdominal seg- 
ment as described for I. murcia. Abdomen shorter: abd. + app. 29.0—30.5, hind 
wing 29.0—31.5 mm. Hab.: Sumba . . . . . . . orchestra 
. Sup. anal apps. in dorsal view widest at base, hen outer hot: weakly angulated at 
about 1/ the length from base, then straight and parallel-sided and finale abruptly 
and obliquely bent inward with knob-like external protuberance at the angulation, 
tips compressed and bluntly pointed. Apex of inf. app. gradually narrowed, hol- 
lowed out above and with pointed tip; lateral projections strong, subtriangular, 
placed well beyond midway length of appendix . . . . i aly hd TA 
. Sides of synthorax with two complete broad yellow stripes the Suini stripe 
somewhat narrowed above but reaching upper margin of meso-metapleurae, the met- 
epimeral stripe often somewhat undulated or irregular but never interrupted. Main 
body of posterior genital hamule greatly swollen, outer border in side view strongly 
convex proximal to the hook-like inner branch. Middorsal crest of 10th abdominal 
segment obtuse, forming a low, bluntly triangular boss. Size smaller: abd. + app. 
28.0—29.0 mm, hind wing 27.5—28.0 mm. ee Basilan (P.I.) and Hong- 
Kong Zu . . . yolanda 
. Yellow spiracular ces at Monet sites narrower, er at end below the spiracle 
but frequently constricted or interrupted at about halfway its length, the dorsal 
portion not quite reaching upper margin of meso-metapleurae; yellow stripe on 
posterior portion of metepimeron also obliterated, consisting of a large oval anterior 
(lower) and a smaller subtriangular posterior (upper) spot. Sup. anal apps. almost 
straight in side view, apex shorter, less abruptly inbent, external subbasal angulation 
barely indicated and subterminal protuberance likewise less pronounced than in 
I. yolanda (Fig. 2). Main body of genital hamule less bulging, outer border in side 
view but slightly convex proximal to the hook-like inner branch (Fig. 1). Middorsal 
crest of 10th abdominal segment more strongly raised, forming an acuminate ridge 
which slopes down steeply posteriorly. Size larger: abd. + app. 31.0—32.0 mm, 
hind wing 30.0—30.3 mm. Hab.: ? Luzon; Mindoro; Leyte; Mindanao (P.l.) 
D lc a MERE RE A SR philippa 


8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Idionyx philippa Ris 
Fig. 1—2 


1912. Ris, Suppl. Entom. 1: 80 (® key), 81—82, fig. 16 (2 right wings, ventral side), 9 
Naujan, Mindoro I., P.I. — ? 1937. Needham & Gyger, Philipp. J. Sci. 63: 57 (key Idiophya and 
Idionyx), 58—59, 2 Luzon, Los Baños. 


Material. — Philippine Is: 1 ® (adult), Naujau (recte Naujan), Mindoro, 
Phil. Rolle 1910, phot. (in Ris’s writing), with red label typus, holotype I. phtlippa Ris 
(SMF). 1 g 1 @ (adult), Mindanao I., Misamis Or., Bal-ason, 2.1V.1960 (g') and 
Mt. Emgagatao, 13.IV.1961 (2), H. Torrevillas; 1 ® (adult), Mindanao I., Cotabato, 
Parang, 23.III.1953, Henry Townes; 1 4 1 9 (adult), Leyte I., So. Leyte, Anahawan, 
Mainit Spring, 31.V.1970, Cristobal Plateros. 


Previously known only from the type, I. philippa has remained a somewhat puzzling 
insect ever since it was described. Dr. H. Schröder has been kind enough to send me on 
loan Ris’s female for comparison with recently acquired specimens of Philippine Idionyx, 
and this enabled me to decide upon its proper status. There are some discrepancies in 
the original description which need clarification. Thus the occipital triangle of the type 
is said to be “flach” and to have no “frontalwärts verlängerte Leiste der I. claudia 9”, 
a rather misleading statement since the whole structure, though indeed smaller and less 
protruding than in I. claudia, is raised well above level of compound eyes, the slightly 
convex upper surface being marked off from the steeply sloping sides by an acute carina, 
i.e. a shape quite similar to that seen in the remaining species dealt with in this paper. 
The tibiae, stated by Ris to be “trüb rötlichbraun” are, indeed, much obscured, yet the 
hinder pair are distinctly yellowish externally, as they are in all other regional species. 
Ris’s description of the coloured spots at the thoracic sides is incomplete: the yellow 
metaspiracular stripe is broadest at the spiracle and, though tapering to a point and 
leaving off a short way above it, continues upward as a lanceolate streak placed in line 
upon the first suture; the much larger mark on the metepimeron is oval, not “fast 
kreisrund’’, thus conforming to the shape it has in the other individuals. 

The specimens from Leyte and Mindanao listed above are undoubtedly conspecific 
with I. philippa. However, they differ among themselves, showing not only considerable 
variation in the extent of yellow thoracic markings but also in their dimensions and wing 
venation, no two specimens of either sex being exactly alike. As pointed out before, under 
Idiophya salva, even the left and right pair of wings in a single individual (e.g. the type 
of I. philippa) may show differences in the venation. This may go hand in hand with 
dense venation, the wings of the type being somewhat more closely reticulated than in 
the other females, the anal loop containing no less than 4 + 6 cells (see below). 

The following descriptions and figures are based on the pair from Misamis Or. 
(Mindanao), which are rather larger than the remaining specimens. 


Male. — Labium with the median lobe and basal one-third of the lateral lobes out- 
wardly chrome, for the rest dark chestnut, except a narrow line bordering the free margin 
of the lateral lobes, which is black; marginal fringe of golden yellow bristles. Base of 
mandibles black, distal portion dark brown. Labrum deep black with a large bright 
ochreous bilobate marking, broadly connected with the base and occupying most of the 


M. A. LIEFTINCK: Studies in Corduliidae I 9 


surface. Anteclypeus blackish with a subtriangular pale yellow dot placed in the middle. 
Postclypeus deep black, the lateral lobes with slight metallic green lustre basally. Frons 
anteriorly black with brilliant emerald green reflex on either side of the smooth deep 
black sulcus, the metallic colour changing to deep purplish-blue above; surface irregularly 
wrinkled by large superficial punctures. Vertex low, surface convex, barely impressed 
medially and more finely punctate. Occipital triangle somewhat raised and acutely ridged 
above level of compound eyes, surface smooth, black; rear of the head glossy black. 

Prothorax brownish-black, anterior border and a spot low down upon the sides, yellow. 
Synthorax metallic green marked with ochreous, as follows. Posterior half of mesin- 
fraepisternum; an adjoining indistinctly brownish-yellow whiff ventrally on the 
mesepisterna; an almost complete, slightly irregular, though barely constricted, stripe at 
the first lateral suture crossing the spiracle, widest at and below the latter and continued 
downward upon whole posterior half of metinfraepisternite; a large oval twin-spot placed 
in the long axis of the body and occupying the latero-ventral surface of the metepimeron. 
Ventral surface purplish black, with a yellow L-shaped stripe at the median and lateral 
dividing lines of the basal parts of the poststernum; yellow are also the metapostepimera 
and apical portion of the poststernum. Ante-alar triangles dull black. 

Coxae and trochanters of fore and middle legs yellow, but outer surface of mid 
trochanter brown; those of the hind legs brownish black, the coxa with small yellow 
postero-basal spot. All femora brownish black, the tibiae yellowish with definite obscura- 
tion towards base and apex; keels light yellow, extending along more than distal one- 
third on fore tibia (36 : 100), about one-tenth on mid tibia and along more than seven- 
tenths (76 : 100) on hinder pair. 


Fig. 1-—2. Idionyx philippa Ris, & from Mindanao. 1, genitalia; 2, anal appendages, dorsal, and left 
side view 


10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Wings unspotted, membrane slightly tinged with greyish-yellow, especially towards 
the tips. Antenodals 12-13 on fore wings, 8 on hind wings; postnodals 6 and 9, res- 
pectively. Pterostigma unbraced, short, covering about 11/ cells, deep black. Arc at Axg, 
sectors originating low down at a point scarcely removed from Cu + A, fused together 
almost as far as first cross-vein beyond Arc. Only 1 cross-vein in ht and 2 Cux on all 
wings (inclusive of the 4- or 5-sided “internal triangles”) ; discoidal field of fore wing 
parallel-sided, with a single row of 13—14 cells followed by 3—4 double cells well 
beyond level of nodus, and 4—5 marginal cells; the same of hind wing with single row 
of 6 cells as far as level of nodus, then expanded, with 7 marginal cells; 2—3 Bxs on 
fore as well as hind wings. Anal loop of hind wing elongate, made up of 7 cells. Anal 
triangle two-celled, the angle after a distinct concavity rounded; only one basal cell 
between loop and anal triangle and two rows between triangle and anal margin of hind 
wing. Membranula about equal in width to the triangle midway its length, extending 
all along the strongly undulated marginal vein; colour grey, growing paler (almost 
white) towards base. 

Abdomen slender, spindle-shaped, basal segments but slightly expanded, from base 
to apex of 7 widened, thereafter almost parallel-sided as far as end of 9, then again a 
little narrowed. Colour deep black, tergites 2—3 smooth and shiny, progressively less 
so posteriorly; 7—10 almost lustreless. Segm. 1 with tiny yellow lateral spot, 2 with 
complete broad yellow band bordering ventral margin but excluding the auricles; the 
colour also occupies the basal portion of the lobus posterior; a similar, though much 
narrower, stripe along basal half of lower margin of tergite 3, the distal half of 7 
ventrally, and whole lengths of 8 and 9, likewise on their ventral surfaces (widest at end 
of 8). No dorsal yellow markings save for tiny transverse yellowish streaks on the 
intersegmental rings of segments 3—5. Genital organs mainly black, plate-shaped 
anterior lamina with crescent-shaped apical emargination; anterior hamulus with distinct 
conical apex; posterior hamulus with its outer branch yellow on each side of the black 
rim (Fig. 1). Suberect marginal pubescence fringing ventral surfaces of tergites 7 and 8 
and sternite of 8 only little more conspicuous or longer than elsewhere, golden yellow, 
all hairs being directed obliquely mesiad and caudad. Segment 10 strongly raised and 
forming an acute middorsal crest, whose basal portion is convexly rounded in lateral 
view. 

Anal appendages black, shaped as in Fig. 2; bristly pubescence neither very dense 
nor tufty, yellowish brown. 

Female. — Resembling the male in most respects, even in the shape of the vertex and 
occipital structure. Differs slightly in that the bipartite yellow marking upon the posterior 
portion of the metepimeron is divided into two spots, one more or less oval, the hinder- 
most spot rather more triangular in outline. Femora somewhat lighter brown, the outer 
faces of the first two pairs of tibiae also brown. Wings a little broader than in male. 
Membrane hyaline, except the bases as far out as Ax and Arc, diffusely and not deeply 
saffronated. Neuration very similar to male. Antenodals 13 on fore wings, 8—9 on hind 
wings; postnodals 6—7 and 7—8, respectively. Arc slightly distal to Ax, on all wings; 


122 12 DE sb : ‘ è ; è ; 
ht Et Cux 50 Bxs 1 = Discoidal field of fore and hind wings as in male, with single 


row of 12—14 cells to beyond nodus in fore wing and 6—7 as far as level of nodus in 
hind wing. Anal loop consisting of 8 or 9 cells; three cell-rows between loop and anal 


M. A. LIEFTINCK: Studies in Corduliidae I 11 


margin of hind wing. Membranula a little longer than first marginal cell, coloured as in 
male. 

Abdomen of the usual cylindrical shape, nature and colour of tergal integument as 
described for the male. Yellow markings reduced to a point at the sides of 1, a stripe 
along tergal margins of 2 and 3 (widest basally on 2) and a fine yellow line bordering 
the lower margin of tergites 4—8, the 8th in addition having a small postmedian spot 
near ventral margin. Valvula vulvae not projecting beyond apical border of 8th tergite, 
apparently shaped similarly to that described and figured for I. yolanda Selys by Lieftinck 
(1939, Fig. 2), although the terminal segments are rather compressed, preventing appro- 
priate comparisons. 

Measurements: & abdomen + app. 32.0 mm, hind wing 30.3 mm, pt. fore wing 
1.2 mm; ® 32.0, 34.0, 1.8 mm, respectively. 

Male (Leyte). — Differs from the former only in being a little smaller and in that 
the yellow metaspiracular stripe is so much constricted as to become almost interrupted 


(VIZIO 12 eZ wy be 
Wol NOCH eer discoidal 
8. 9. 8.10 1.1 22 


at its middle. Wings entirely hyaline. Nodal index 


field of fore and hind wings as in the Mindanao example; Bxs — anal loop with 
4 + 5 and 3 + 5 cells. Genitalia and anal appendages as in the previous male. 
Measurements: abd. + app. 31.0 mm, hind wing 30.0 mm, pt. fore wing 1.5 mm. 
Female (Leyte). — Generally similar to the male. First pleural thoracic stripe 
completely divided into two portions, as described for the type, but separate spots are a 
little larger. Wing membrane coloured similarly to the type, the basal spots equally 
2.1390: pease Cun: discoidal 


extensive but not so deeply stained. Nodal index ="; à 
LOR SMS: Mell 2.2 


i DER. 
field of fore wing with single row of cells up to level of nodus; Bxs LE anal loop with 


3 + 5 cells in both hind wings. 

Measurements: abdomen 30.0 mm, hind wing 32.2 mm. 

Female (Cotabato, Mindanao). — A small-sized specimen with rather broad and 
uninterrupted first lateral thoracic stripe but with the metepimeral spots similar to those 
of the others. Wings deeply saffronated, basal spots as in the type. Nodal index 
641241256 1.1 

emt 
8. 8. 8. 8 1.1 
4 +5,4 +95. 

Measurements: abdomen 30.5 mm, hind wing 30.0. 

The three females recorded from the island of Luzon by Needham & Gyger (loc. cit.), 
are referred here with some misgivings because the synthorax is described to bear “a row 
of four yellow spots low on the sides, the first above the middle coxa, the second on the 
spiracle, the other two on the metepimeron”. This would mean that the metaspiracular 
stripe is devoid of its upward prolongation, which in all other specimens examined by 
me is present, either in the form of a more or less isolated streak or fused together with 
the spot at the spiracle. 


DD dE : È 2 DD, 
Cux 53: discoidal field as in the Leyte specimen; Bxs mi anal loop 


12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Idionyx murcia spec. nov. 
Fig. 34 


Material. — Lesser Sundals.:1 & 2 9 (1 9 immature), Sumbawa I., Central 
Sumbawa, Semongkat-atas, 21.1V.1961 (¢), Semongkat, 300 m, 27.1V.1961 (2), and 
Route Batu dulang, 600 m, 10.IV. 1961 (2 juv.), all P. Jauffret & R. Pujol, ex coll. 
A. Heymer. The G' is the holotype. 


The characterization of this new species is the result of a direct comparison with 
I. philippa Ris, described in the previous pages. 

Male. — Lateral lobes of labium with angulated yellow extero-basal spot, the 
remaining parts of labium brownish-black. Labrum entirely chrome, narrowly bordered 
anteriorly with black. Anteclypeus likewise chrome, only the somewhat impressed side- 
angles obscured. Postclypeus dark metallic green with conspicuous chrome spot in the 
middle, the latter shaped like a broad triangle whose apex is transversely cut off. Frons 
emerald green changing to blue-green above; anterior surface less protuberant, more 
closely and deeply corrugated on either side of the black sulcus and also less shiny, than 
in I. philippa, but occipital triangle and vertex much as in that species. Rear of the head 
glossy black. 

Wings unspotted, membrane slightly tinged with brownish-yellow. Neuration much as 
described for I. philippa and I. orchestra Lieft. (from Sumba I.), except for the fol- 
lowing slight differences. Antenodals 12—13 on fore wings, 8 on hind wings; post- 
nodals 6—7 and 9, respectively. Pterostigma covering little more than the underlying cell, 
dark brown. One cross-vein in ht and 2 Cux on all wings (inclusive of the 4- or 5-sided 
12), discoidal field of fore wing with a single row of 11—12 cells to a little beyond level 
of nodus, followed by 3—4 double (or triple) cells and 5 marginal cells; the same of 
hind wing with single row of 6 cells to level of nodus and with 6 marginal cells; 1 Bxs 
on fore, 2 on hind wings. Shape of anal triangle, angulation and membranula exactly as 
in I. philippa. 

Anterior lobe of prothorax clear yellow, pleurae largely yellow, posterior lobe brown. 
Synthorax brilliant metallic green, extensively marked with bright ochreous, as follows. 
A sharply defined, elongate-oval juxtahumeral band that occupies also the posterior two- 
thirds or more of the mesinfraepisternum, extending two-fifths up the dorsum; a 
similarly coloured, broad and almost parallel-sided stripe (ca. 0.7 mm broad) crossing 
the spiracle and extending from just below the dorsal crest right down onto the posterior 
part of the hind coxa, on which it is much narrower; a still broader, somewhat curved, 
metepimeral band covering only little less than its posterior half. Ventrally these metep- 
imeral bands are confluent across metapostepimera and apical portions of the post- 
sternum, forming together a U-shaped marking which itself encloses a metallic blue-black 
spot of the same form, the central area of the ventral surface remaining largely yellow. 
Ante-alar triangles black. 

Legs coloured as described for I. philippa; tibial keels yellow, those on fore tibia 
occupying their apical two-fifths (40: 100), on mid tibia about one-eleventh and on 
hinder pair about seven-tenths (71: 100). 

Abdomen with the terminal segments less expanded than in I. orchestra and philippa 
but for the rest very similar in shape and colouring, lacking metallic reflections. Segm. 1 


M. A. LIEFTINCK: Studies in Corduliidae I 13 


Fig. 3—4. Idionyx murcia spec. nov., & holotype from Sumbawa. 3, genitalia; 4, anal 
appendages, dorsal and left side view 


with large cuneiform yellow lateral spot; 2 with complete very broad band running along 
ventral margin not covering the auricles but occupying also the lobus posterior, which is 
obscured only at its apex; a similar though narrower stripe, tapered posteriorly along 
basal one-third of lower margin of tergite 3; broader stripes along whole lengths of 
tergites 7, 8 and 9 on their ventral surfaces. No dorsal pale markings. Genital organs 
shaped and coloured much as in the allied species but anterior hamulus triangular; lobus 
posterior rounded, with dense fringe of long radiating golden yellow bristles (Fig. 3). 
Marginal pubescence on ventral surface of tergites 7 and 8 and of sternite 8 erect, rather 
long but not very dense, all hairs brown and directed ventrad. Segment 10 impressed on 
either side of a small and rather low (though acute) middorsal crest, which is barely 
visible in lateral view. . 

Anal appendages black, shaped as in Fig. 4; pubescence short and inconspicuous, 
except a subapical fringe of much longer light brown hairs at the outer faces of the 
superior pair. 

Female. — Differs from the male only in having all yellow markings on thorax and 
abdominal tergites a little more expanded; also in that only the hind tibiae (save the 
bases and apices) are bright yellow. Wings broader, bases bright orange-yellow as far 
distally as Ax2, the arculus and base of discoidal triangles in cv; membrane otherwise 
either entirely hyaline (immature 9 ) or smoky yellowish-brown (adult 9 ). Antenodals 
12—14 on fore wings, 8—9 on hind wings; postnodals 6—7 and 8, respectively. Arc 

2.1 LL Zal 


Weal 11 M 6 
at or a little proximal to Ax; ht —- and —; Cux —; Bxs and —~. Discoidal field 
eee aak ALE MITE 1 DD 2.2 2.3 


14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


as in male, with single rows of 12—13 cells in fore wings and 5—6 in hind wings. 
Anal loop made up of 8—9 cells; three cell rows between loop and anal margin of hind 
wing. Pterostigma black. Membranula a little longer than first marginal cell, coloured as 
in male. 

Abdomen cylindrical; yellow lateral spots and bands of 1—3 and 7—8 (absent on 9) 
more conspicuous, especially the one on 2, which is widest at extreme base. Valvula 
vulvae indistinguishable from that of I. yolanda Selys (see description of I. philippa Ris). 

Measurements: g abdomen + app. 31.7 mm, hind wing 30.0 mm, pt. fore wing 1.0 
mm; ® (adult) 30.5, 33.0, 1.8 mm, respectively; @ (juv.) —, 31.5, 1.7 mm, res- 
pectively. 


The larva of Idionyx does not seem to have been described with absolute certainty. 
Fraser (1936) only says that it is “distinctly Libelluline in character”. In their work on 
the Odonata of the Philippines, Needham & Gyger (1939: 59—60, pl. 8 figs. 9395) 
published an account of a larva from Luzon which they ascribed to “Idzonyx spec.?” 
Although it was stated that the venation in the wings was well preserved and generic 
determination positive, there remains doubt as to the correct identification of this 
specimen. This is evident not only from the size, Needham’s example measuring 19—20 
mm, i.e. much too large for regional Idionyx, but also from the description and figures, 
which suggest some other genus. This Philippine larva is peculiar in that the crenulated 
labial palpus bears only “six coarse teeth on the terminal border (five only being shown in 
the figure!), the three large middle teeth wider than high, obliquely rounded, each armed 
with about a dozen spinules in double array longer and shorter’. Lateral abdominal 
spines are present on 8 and 9, and there are no dorsal hooks, features applying to 
Idionyx. Other details in the description do not fit any other regional corduliid genus, 
although the facies is reminiscent of Hemicordulia and Procordulta. However, the larvae 
of Procordulia sambawana Foerster and artemis Lieftinck possess seven rounded teeth at 
the distal border of the palpus (inclusive of the divided outermost tooth) and dorsal 
abdominal hooks are present on segm. 4—8 in sambawana but absent in artemis (Lief- 
tinck, 1933: figs. 9—10). The larva of Hemicordulia mumfordi Needham also has 
seven palpal teeth while middorsal hooks on the abdominal segments are wanting (Need- 
ham, 1932, fig. 34). Hemicordulia australiae (Ramb.) is similar but possesses middorsal 
hooks on 3—9 or 4—9 (Watson, 1962: 14). It is evident, therefore, that it remains 
impossible to express any definite opinion on the identity of Needham’s larva from 
Luzon. For a description of Macromidia donaldi Fraser (1936), see under that genus. 

The following authenticated material of Idionyx larvae has been studied by the present 
author. 

Idionyx yolanda Selys. — Malay Peninsula: 1 & (with exuviae), Central Perak, 10 mi. 
N. of Ipoh, Sungai Chepor, 6.III.1963, M. A. Lieftinck; 1 9 (with exuviae), Selangor, 
6 mi. Bukit Tiga-Subang Rd., Klang distr., Sungai Pelumut, 5.VI.1963, emerged Kuala 
Lumpur, 15.VIIL.1963, J. I. Furtado. 

Idionyx spec. indet. — Malay Peninsula: 3 ex. ult, 1 ex. penult, Perak, Sungai Yum, 
15.11.1933, M. W. F. Tweedie. A number of other unidentified specimens, collected by 
J. I. Furtado and the author during their explorations of streams in various provinces of 
the Malay Peninsula (1963—64), are not now available for comparison. 

Idionyx montana Karsch. — Central W. Sumatra: 1 g' 1 © ult (reared from larvae), 
Benkulen, S.W. slope of Mt. Dempo, 300 m, received 24.X.1941, from W. C. Verboom; 


M. A. LIEFTINCK: Studies in Corduliidae I 15 


Fig. 5—7. Idionyx montana Karsch. 5, ultimate instar larva from Mt. Dempo (Sumatra), live speci- 
men; 6, inner view of exuvial labium; 7, left palpus, more enlarged 


emerged in Buitenzorg laboratory, 15.X11.1941 (@), and 23.XII.1941 (g'). West Java: 
1 G ult, Djasinga, Tjibarangbang (stream), ca. 150 m, sifted from algal growth on 
bottom of tiny rivulet, 31.VIIL.1937, M. A. Lieftinck; died in captivity at Buitenzorg 
(Bogor), 10.X.1937. 

The above larvae resemble each other so much in outward appearance and structure, 
that I have abstained from making a careful study of all. The illustration of the whole 
larva (Fig. 5) was made from one of the Sumatran specimens shortly after its penulti- 
mate ecdysis. 


16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 
Description of full grown larva of I. montana Karsch from Sumatra (Fig. 5—7) 


Body compact, sparsely pubescent dorsally. Head large, directed obliquely upwards 
during life so as to expose part of the labium (Fig. 5); eyes knob-like and distinctly 
protuberant when looked at from above with mask concealed from view. Upper surface 
of frontal plate concave, its anterior border evenly convex. Postocular lobes evenly 
rounded, surface set with short, thick spicules, which are crowded together posterolater- 
ally. Labium of large size, postmentum (hinge) reaching back to about halfway between 
mid and hind coxae. Median lobe (ligula) rather short, forming a low triangle, with a 
row of 11—13 short spiniform setae on each side and a small bunch of these at the apex. 
Premental setae 5 + 5, placed in a curve. Labial palpus with 6—7 deep triangular 
crenations and 7—8 rounded dentations, the outermost of the latter divided and all 
carrying about 8 spiniform marginal setae at the apex; palpal setae long, 5 on either side, 
outer margin of palpus with minute spiniform setae; movable hook rather short and 
only slightly curved. Antenna short, seven-segmented. Propleural lobes of thorax almost 
as broad as the transverse diameter of head, slanting forward, borders fringed with very 
long hair. Wings widely divaricate, hind wing rudiments reaching as far as halfway 
length of segm. 6. Legs long and slender, sparsely clothed with long soft hair; claws 
simple. Abdomen oval, rather depressed, greatest width 5.0 mm at end of 6th segment; 
all segments fringed laterally with long hair. Dorsal hooks absent. Lateral spines acute, 
present only at segm. 8 and 9 and of small size. Segment 10 very short and annular. Anal 
appendages short, epiproct and paraprocts straight, triangular, acuminate, the latter only 
a trifle longer than epiproct; cercoids shorter than either, incurved with convex outer 
border, tips acute. 

Ground colour of body pale yellowish green, the coxae almost white; pattern distinct, 
varicoloured grey-brown to dark brown, the abdominal mottling lighter than colour of 
wings; two distinct broad dark bands on all femora, as shown in Fig. 5. 

Measurements. Total length 13.7 mm; width of head behind eyes 4.5 mm; hind femur 
4.5 mm; hind tibia 4.7 mm; width of abdomen at end of 6th segment 5.0 mm. 


Macromidia Martin, 1907 


Syn.: Indomacromia Fraser, type-species I. donaldi Fraser, 1924. 


In current classifications, Macromidia figures as a member of the Macromiinae (= 
Epophthalmiinae of Fraser, 1957). More recently still, this was done, with some mis- 
givings, also by Gloyd (1959), who gave Macromiinae family rank. Idionyx, on the other 
hand, which Needham (1937) still left in the Corduliinae, was removed therefrom by 
Tillyard & Fraser (1940), who erected a new subfamily, Idionychinae, to hold it. Their 
arguments were the long fusion of the arculus sectors, the shape of the anal loop, the 
distal position of the nodus on the fore wing, the incompletely recessed discoidal triangle 
on the hind wing; and, coupled with these venational characteristics, the large head, 
weak sclerotization of the body, and the highly organized and intricate anal appendages 
of the male (often correlated with a peculiar form of the vertex in the female). 

Taken on the whole, the venation of Macromidia resembles that of Macromia more 
closely than Idionyx. Yet Macromidia differs strikingly from the majority of Macromia 


M. A. LIEFTINCK: Studies in Corduliidae I 17 


in its more rounded wings and the less markedly sinuous primary veins, two characters 
which it shares with Idionyx. Almost all venational characteristics that served Fraser to 
define his Idionychinae are equally applicable to Macromidia. Concerning the morpholo- 
gy of other body parts, with their manifold appendages, it is impossible to find a set 
of clear-cut characters that would guarantee every species to be put into its proper genus. 

In view of the fact that Macromidia shares so many characters with both Idionyx and 
Macromia — not to mention the problematical Idzophya! — there can be no doubt about 
their affinity, Macromidia taking rather an intermediate position between Idionychinae 
and Macromiinae but lying closest to Idionyx. This will inevitably lead to a revaluation 
of subfamily characters. Seeing how many corduliid genera are still insufficiently 
characterized, it seems best therefore to attempt first a summary of the principal generic 
characters serving to distinguish between Idionyx and Macromidia and to leave in abey- 


ance Fraser's subfamily divisions proposed in the “Reclassification” (1957). 
These characters can be recapitulated in tabular form, as follows: 


Idionyx 


Fore wing triangle equilateral, smaller 
than that of hind wing; fore wing # 
irregularly trapezium-shaped, 4- to 5- 
sided and placed in the long axis, i.e., 
Cu + A almost in line with anal side of 
ti. No distinct intercalated supplementary 
sectors in discoidal field of fore wing, 
which is parallel-sided for a long dist- 
ance. Anal loop of hind wing elongate 
(sausage-shaped), more than two times 
longer than broad, traversed lengthwise 
by a bisector. Proximal (inner) side of 
anal triangle of male hind wing strongly 
sinuous. Pterostigma small, less than 2 
mm long. Nodus placed far distad: post- 
nodal space (distance between nodus and 
pterostigma) at most half as long as 
antenodal. Only 1 (fore wing) or 2 
(hind wing) proximal postnodal cross- 
veins discontinuous into adjacent space 
between R and M,. 

Apical keel on mid tibia present 
though vestigial. Tarsal claws with robust 
inferior tooth longer and stronger than 
the apical tooth (Fig. 12). 


Outer branch of male posterior genital 
hamule thick and equal in length to 
sickle-shaped inner branch. 


Macromidia 


Fore and hind wing triangles of equal 
size, the distal side on fore wing longest; 
fore wing ? 4- or even 5-sided, but 
placed more transversely, ie, Cw + A 
reaching anal angle of triangle after 
being strongly deflected around inner 
angles of #. Discoidal field of fore wing 
more evenly widened distad, with distinct 
and rather long intercalated supplemen- 
taries (including Msp/). Anal loop of 
hind wing short and more rounded, less 
than two times as long as broad and 
lacking a bisector. Proximal (inner) side 
of anal triangle of male hind wing not 
markedly sinuous. Pterostigma larger, 
over 2 mm long. Nodus usually placed 
more proximad, antenodal space only 
twice as long as postnodal, or less. 
Usually 3 or 4 proximal postnodal cross- 
veins not continued into adjacent space 
between R and M, in all wings. 

Keel on mid tibia extending along its 
distal half. Tarsal claws with inferior 
tooth distinctly shorter and less acute than 
apical tooth, both about equally strong 
(Fig. 13). 

Outer branch of male posterior genital 
hamule longer, thinner and incurved, 
projecting much beyond hook-shaped in- 
ner branch. 


18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Male anal apps. long and slender, Male anal apps. of normal length, 
longer than segm. 9 + 10. shorter than segm. 9 + 10. 

Apical portion of female 8th sternite Valvula vulvae of female well marked 
abbreviated but prominent in side view off, extending caudad almost or fully 
and scoop-shaped, not at all emarginate half the length of 9th segment, deeply 
and lacking leaf-like lobes. emarginate forming two pointed lobes. 


From this table it will be clear that there is no evidence whatsoever that would justify 
a classification of Idionyx and Macromidia in separate subfamilies, as has been done in 
the past. An interesting feature which the two genera share alike with Macromia is the 
characteristic shape of the occipital triangle, with its acute-angulate lateral borders, a 
character perhaps found also in other allied genera of doubtful position. For instance, in 
the present context it should be emphasized that this part of the occiput is shaped 
similarly in most, if not all, species of Synthemis, and I have observed the same structural 
feature in the Australian Micromidia Fraser. 


The genus now includes the following species: 


M. rapida Martin, 1907, type-species. — g ® Tonkin. 

M. fulva Laidlaw, 1915. — g' N. Borneo; 9 Sarawak (Laidlaw, 1920). 
M. g. genialis Laidlaw, 1923. — g' Malay Peninsula; © unknown. 

M. g. erratica Lieftinck, 1948, stat. nov. — g' Q S. Sumatra; g' S. Java. 
M. donaldi (Fraser, 1924). — G © Peninsular W. India. 

M. shanensis Fraser, 1927. — G 9 Upper Burma. 

M. atrovirens Lieftinck, 1935. — © S.W. Sumatra; 4 unknown. 

M. samal Needham & Gyger, 1937. — ® Mindanao (P.I.); $ unknown. 
M. asahinai spec. nov. — ¢ Palawan (P.I.). 


Brief diagnoses of six of these have been given by Lieftinck (1935: 195—196). The 
species are more diversified in their venation and markings than Idzonyx. Two of them, 
M. rapida and fulva have more densely veined wings than the rest, the discoidal field in 
both fore and hind wings commencing with two rows of cells, whereas in all others only 
a single row extends outwards from a point varying between 3—4 cells proximal to Nod 
to about as far as that level on the fore wings, and to 3—5 cells beyond # in the hinder 
pair. M. samal (3 unknown) is intermediate in this respect inasmuch as the field in the 
fore wing expands to two cell-rows already midway between ¢ and Nod, there being only 
two undivided cells distal to ¢ in the hind wing. M. rapida, donald: and fulva also differ 
from the other species in that the body is metallic black with well defined middorsal 
yellow spots on segm. 3—7 (rapida and donaldi), or marked with light brown rings 
(fulva). M. samal seems to come nearest shanensis from Burma; the female genital 
organs of the latter have recently been figured by Asahina (1970). M. atrovirens is a 
large and very dark-coloured species with no near allies. The rediscovery of the much 
smaller M. genialis in the island of Penang, off the westcoast of Malaya, leads me to 
point out some features separating it from M. erratica, which I now consider to be only 
a subspecies of the former. Lastly, one new species from Palawan can be added to the 
list. Asahina (1968) supposed this to be the undescribed male of Idionyx philippa Ris, 
but it is a true Macromidia, which I have renamed M. asahinai spec. nov. 


M. A. LIEFTINCK: Studies in Corduliidae I 19 


The larva of Macromidia donaldi (Fraser) was briefly described and figured by 
Fraser (1936b: 208, fig. 48A, A’, whole larva and labial mask). The following charac- 
ters are taken from the description. 

Eyes rather prominent (more strongly so than in Idionyx, according to Fraser’s sketch) ; 
head across the eyes only little less broad than the thorax. Midlobe of labium produced 
slightly; 13 mental setae in a curved row on each side; palpi bordered with 7 crenulate 
spined teeth, the last one duplicated, and with 8 marginal lateral setae; movable hook 
strong. Abdomen broadly fusiform, rather strongly carinated middorsally but lacking 
dorsal spines; a robust apical spine on each side of segments 8 and 9. Legs of moderate 
length, almost naked, and with two broad dark brown annules on all femora. Total 
length 15 mm, head 6 mm wide, abdomen 9 mm long and 5 mm in width. 


Macromidia genialis Laidlaw 
Fig. 13 


1923. Laidlaw, J. Mal. Br., Roy. Asiatic Soc. 1: 231—232, pl. 5 (4 wings, genit. & apps.), 
6 Malaya. i 


Material — Malay Peninsula: ¢ (holotype), Pahang, Gunong Tahan, 
10.X11.1921, M. genialis & Type, det. F. F. Laidlaw (BM); & (adult), Penang Island 
[off Prov. Wellesley}, W. Malaya, stream in Catchment area near Batu Feringgi, 
1.111.1963, M. A. Lieftinck. 

The wing venation of the type is well shown in Laidlaw’s photograph of the right 
pair. To his description may be added that the labium is bright orange-yellow. Wings 
unspotted; discoidal field in fore wing with a single row of eight-nine cells to about two 
cells proximal to level of nodus, in hind wing with four and five single cells following 
the triangle. Cross-veins in cu = inclusive of the 4-sided “internal triangle’; ht = 
The rather brief description can be further emended by a number of characters taken 
from the second specimen, which is a little larger than the type. 

Male (Penang). — Labium orange-yellow; mandible-bases blackish, surface shiny; 
labrum black, sides faintly metallic blue with a pair of slightly impressed areas on either 
side of the middle; anteclypeus yellowish brown, postclypeus metallic blue, frontoclypeal 
groove black. Frons and vertex brilliant metallic blue-green, rather densely punctate; 
frons short, hardly projecting beyond the vertex when viewed laterally, its surface 
flattened anteriorly, pubescence black; vertex broader than long, surface evenly and 
strongly convex and clothed densely with long erect dark hair. Occipital triangle raised 
above level of compound eyes, shorter than median eye-line with sharply acute sides, its 
shape exactly as described for the regional species of Idionyx. Prothorax entirely chrome. 
Synthorax brilliant metallic green; dorsum with a bright yellow spot between the 
axillaries of the fore wings and the metallic ante-alar triangles. Outer half of each 
mesepisternum with well-defined, broadly oval, brown spot continued ventrad on to the 
mesinfraepisternum and extending dorsad for about half the length of the dorsal surface. 
Metinfraepisternum yellow, its anteroventral two-fifths metallic green; metepisternum 
with slightly oblique, yellow, lanceolate stripe, pointed at either end, situated well behind 
the spiracle, incomplete above and extending about as far upward as 1 mm before the 


20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


base of fore wing. Metepimeron ventrolaterally with short upward extension from the 
yellow colour underneath and with the metapostepimeron also yellow. Ventral surface 
mainly yellow but anterior portions of metepimera (except medially) metallic green and 
posterior belt of metasternum with transverse streak of metallic purple. 

Coxae of fore legs yellow, the remainder grey-brown; legs brownish black or black; 
tibial keels yellow, those on fore tibia occupying about the apical two-fifths (40: 100), 
on mid tibia slightly less than one-half (44: 100) and on hind tibia about the apical 
two-thirds (70: 100). 

Gol NMG, 3 


10 10 10 10 ° Arc slightly distal to Ax, on all wings; Cux 


Wings clear; nodal index 


3 ht = Bxs = anal loop of hind wing with 8.7 cells. A single row of 6—7 cells 
in discoidal field of fore wing up to level of fork M, ;—M,, only 3 or 4 continuous 
cross-veins in that of hinder pair. Proximal side of anal triangle only slightly sinuous; 
anal angle bluntly angulated, rounded.1) Pterostigma brownish-black surmounting 21% 
or even 3 cells. Membranula extending as far as the cross-vein in anal triangle, light grey 
at base, growing darker apically. 

Abdomen deep black, dorsum and part of the sides of segm. 2 metallic green. Yellow 
are: the auricles; a continuous stripe and fine lines along lateroventral margins of tergites 
2—3 (including whole lobus posterior) and 4—6, respectively; a fine middorsal line on 
2 and basal one-third of 3; a narrow but sharply defined longitudinal lance-shaped 
median spot on posterior three-fourths of segm. 6; and complete stripes bordering lateral 
margins of tergites 7 and 8, ventrally. Innermost portions of the ventral surfaces of these 
latter tergites clothed rather densely with short dark hairs, which are directed obliquely 
mesiad and caudad. 

Genitalia and anal appendages shaped much as described and figured for M. erratica 
Lieft. (loc. cit.), the outer branches of the hamuli bright yellow. 

Measurements: abd. + app. 33.0 mm, hind wing 29.0 mm, pt. fore wing 2.7 mm. 

The specimen from Penang was taken by chance in dense jungle, flying rapidly low 
over a trickle in a leafy tunnel formed by overhanging shrubs. 


Macromidia genialis erratica Lieftinck, 1948, stat. nov. 


1948. Lieftinck, Treubia 19: 274—278 (as species), figs. 18—19 (@@ genit. structures), 4 
S. Java. 1954. Lieftinck, Treubia 22 Suppl.: 121. 


This dragonfly resembles M. genialis Laidlaw so much in the majority of its characters 
that I am of the opinion that erratica can hardly be maintained as a full species. It is, 
therefore, classified more appropriately as a subspecies of genialis. The Javan example 
is a freshly emerged male, but all errafzca from Sumatra differ from the two Malayan 
individuals in having the lower half of the thoracic dorsum entirely metallic, lacking the 
sharply delimited oval brown patch of genialis. They are also distinguished from 
genialis by the design of the abdomen: the dorsal carina of abdominal segments 3—7, as 
well as the tergal margins of 3—8 ventrally, are lined finely with citron-yellow, whereas 


1) The strong angulation shown in the photograph of the type is exaggerated and caused by the 
wing-base having been damaged. 


M. A. LIEFTINCK: Studies in Corduliidae I 21 


in genialis these lines are unapparent on the dorsum of 4—5 but broadened to form a 
definite lanceolate spot on the 6th. Lastly, the superior anal appendages of erratica are 
slightly but distinctly more expanded after their middle than they are in typical genialis. 

I took both sexes of M. g. erratica over a small stream in a jungly habitat resembling 
greatly the one which yielded the example of M. g. genialis in the island of Penang. The 
Sumatran insects flew in company with males of Chlorogomphus magnificus Selys, which 
I first mistook for some of the larger Macromia. 


PE 


Macromidia asahinai spec. nov. 


1968. Asahina, Jap. Jour. Zool. 15: 357—358, pl. 2 fig. 11 & figs. 5—10 (& wings, head & 
thorax, and abd. structures), 3 Palawan I. (sub Idionyx philippa Ris). Holotype in B. P. Bishop 
Museum, Honolulu. 


This little species has been so well described and figured by Asahina that it will suf- 
fice to summarize its principal characters, which are taken from the description. The 
insect was misidentified by its author, who thought it to be the underscribed male of 
the puzzling I. philippa Ris, a species now definitely placed in Idionyx. 

Male. — Anteclypeus yellow, labrum and postclypeus blackish-brown. Synthorax 
with four yellow spots low down on the sides: one mesinfraepisternal posterior spot; a 
second, metinfraepisternal, coalescent with a smaller, triangular metepisternal spot term- 
inating just above (and including) the spiracle; a third elliptical stripe traversing 
obliquely the middle of the metepimeron; and a much smaller, triangular mark in the 
posterior corner of the metepimeron and metapostepimeron. Tibial armature not des- 
cribed. 

Wing neuration typical for the genus. Nodal index (partly of right pair only) 
n de t zu, Cux Zn Br WDiseoidal field commencing with 8 single 
270910 25 Le ES 
cells as far as about 3 cells proximal to nodus on right fore wing, and 3 undivided cells 
beyond ¢ on right hind wing. Anal loop comprising 7—8 cells. Pterostigma relatively 
long, as for genus. 

Abdomen brown, broadened at segments 7—8, with narrow middorsal line on 2—4, 
a lateroventral marginal stripe running from the auricles to as far as tergite 9 (very 
narrow on 4—9), and a pair of conspicuous, closely approximated, circular spots of 
yellow just before the middle on the dorsum of 7. Genital organs and anal appendages 
shaped much as in the other members of the genus. Superior pair sub-parallel, somewhat 
sinuous in dorsal view with acute incurved tips, which are a little upturned in lateral 
view; no subapical inferior ventral projection. 

Measurements: abdomen + app. 29 mm, hind wing 27 mm. 


This is the smallest known species of Macromidia, which I have much pleasure in 
naming after my colleague, Dr. S. Asahina, who first described it. 


22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


III. SOME MALAYSIAN SPECIES OF Macromia RAMBUR 


Macromia gerstaeckeri Krueger 


1954. Lieftinck, Treubia 22 Suppl.: 118 (full references). 


Since last discussing this species and its immature stages (Lieftinck, 1950), more 
material of both adults and larvae have come under my notice. In 1963, during our joint 
investigation of the Malayan Odonate fauna, Dr. J. I. Furtado and I found gerstaeckeri 
to be widely-spread at forest streams in the Malay Peninsula; but owing to their rapid 
flight and remarkable inconspicuousness, only few imagines (mostly ovipositing fema- 
les) were obtained, the remainder having been reared from larvae. 

Additional material. — Malay Peninsula: 1 9 (adult), Selangor, Ulu Langat 
area, Dusun Tua, 2—500 m, 9.IV.1963, M. A. Lieftinck; 2 9 (id.), Selangor, Sungai 
Gombak, 1114 mi. Kuala Lumpur-Bentong Rd., 22.V and 27.X1.1963, J. I. Furtado; 
1 & (juv., with exuviae), Selangor, same locality, 16.111.1963, J. I. Furtado & M. A. 
Lieftinck, emerged at Kuala Lumpur, 30.111.1963; 1 ® (adult), Perak, 8 mi. E. of 
Sungai Siput, kp. Talong, Sungai Korbu, 9.1V.1964, J. I. Furtado; 1 & (juv., with 
exuviae), Pahang, 2914 mi. Kuala Lumpur-Bentong Rd., Sungai Tangli, 13.V.1965, 
emerged at Kuala Lumpur, 5.VI.1963, J. I. Furtado. Borneo: 1 © (adult), North 
Borneo, Keningan, 1000—1500 m, 24.XII.1962, Y. Hirashima (BISH). 

A detailed characterization is to be found in the writer’s 1950 publication. Were it 
not for the distinctive form of its posterior hamule, the male of gerstaeckerz would be 
easily mistaken for M. callisto or erato, which are otherwise closely similar and may 
occur together in one locality. This proved to be so with callisto and gerstaeckeri in 
Malaya, where we captured both indiscriminately. 

Female (Malaya). — Labium dark chestnut to brownish-black, except the median lobe 
which carries a pair of transverse citron-yellow spots at base; a circular spot of the same 
colour at the mandible-bases. Anteclypeus dark brown; postclypeus either entirely yellow 
or rather intermingled with brown on the anterior part, the yellow being restricted to an 
ill-defined undulated basal stripe, the side portions for the greater part also remaining 
yellow. Thorax beneath brownish-black, slightly metallic, lateroventral ridge of met- 
epimeron and all sutures sharply outlined in yellow. 
6.15.15.6 SD 
9.10.10.8° 9.10.11.9° 
6.15.16.6 N) PL , 43 4.4 Il > GE fd 5.6 LO 2 nb 
DMO MIS) Yo Dells} DDE Dig 2D 222 44 4.4 4.4 4.5 
loop 9.10, 6.7, 9.10 and 8.8, all except the second including 1 (rarely 2) central cells. 
Costa black with fine posterior yellow line from base about half-way up to nodus. 
Pterostigma black. Membranula ashy grey, almost black. 

Abdomen sparsely marked with yellow: 2 with transverse lanceolate streak, widest at 
middle and tapering on either end, upon middorsum and a large, oblique, slightly 
irregular and constricted patch at the sides not attaining the jugal suture; 3 and 4 each 
with a pair of very small, oval middorsal spots placed immediately in front of the jugal 
suture, those on 4 vestigial; sides of 3 moreover with longitudinal stripe along ventral 
margin but lacking a transverse streak at its base; 7 with large prejugal dorsolateral mark, 
widest at base, rounded off laterally and slightly indented by black from behind at the 


Neuration open. Nodal indices and other details of 4 adulti: 


M. A. LIEFTINCK: Studies in Corduliidae I 23 


crest, occupying from one-fifth to one-fourth of the segment’s length; in one specimen 
this basal marking is much smaller and finely interrupted in the median line. Terminal 
segments unmarked. Valvula vulvae undeveloped: apical border in ventral view some- 
what swollen, very slightly undulated, the receding part of the sternite sloping down 
abruptly (dorsad) and forming a pair of parallel swollen ridges fitting closely against 
the body wall. Anal appendages awl-shaped with finely acuminate tips. 

Female (discoloured). — Similar to Malayan individuals but median lobe of labium 
and basal edges of palpi entirely ochreous. Yellow areas of postclypeus darker and 
invaded by brown on either side of middle. Nodal index Sl. ht: 2: > Cux - ie ‚anal 

IHS 22: 44 
loop with 9—10 cells inclusive of a central cell. Genitalia and appendages as described. 

Measurements: ® abdomen 38.3—40.0 mm, hind wing 34.7— 36.0 mm, pt. fore wing 
2.0 mm (Malaya); 40.0, 36.0, 2.0 mm, respectively (Borneo). 

The females from Malaya differ somewhat from the allotype described from Java and 
also from the one taken at Jor Camp in Perak, figured in my previous paper (1950, fig. 
7). In them the sides of the 3rd abdominal segment are unmarked at base, while minute 
paired dorsal spots are present on 3—4. I am unable to Dj whether or not these dif- 
ferences are due to individual variation. 


Macromia callisto Laidlaw 
Big. 10 18 1A 


1954. Lieftinck, Treubia 22 Suppl.: 116 (full references). 


Material. — Malay Peninsula: 1 g, 1 ® (ovipositing, both adult), Selangor, 
1114 mi. Kuala Lumpur-Bentong Rd., Sungai Gombak, 12.VI.1963, J. I. Furtado; 1 9 
(adult), Central Perak, 10 mi. N. of Ipoh, Sungai Chepor, 6.111.1963, M. A. Lieftinck. 

These specimens are undoubtedly conspecific with the immature type described by 
Laidlaw. Similar to M. erato but smaller, the male with slenderer and more spindle- 
shaped abdomen. Immediately distinguished by the darker clypeus and other details of 
coloration but resembling erato in the shape of the male genital organs and anal append- 
ages. 

Male (adult, Malaya). — Labium and mandibles as described for gerstaeckeri and 
erato. Labrum black, its extreme base on each side of the impressions chestnut. Antecly- 
peus dark brown; postclypeus likewise brown but diffuse dark yellow areas are clearly 
discernible in the middle, in the depressions and at base of the lateral divisions. Frons 
black with slight metallic gloss, shape and surface sculpture of the tubercles as in erato. 

Ground colour of thorax deep black, dorsally with steely blue, laterally with purplish 
and blue reflections; all yellow markings a little narrower and more restricted than in 
erato. Legs as in that species, length of hind femur 8.0 mm; keel on fore tibia a little 
shorter, 37.5 : 100 (erato: 44 : 100). 

Wings clear, nervures black, only the costa finely yellow posteriorly for about half the 

5.14.14.6 


distance from base to nodus on fore wing. Neuration open. Nodal index — OM ht 


24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


3.3. 
DD 
running directly from M, to Cz, in hind wing. Anal loop made up of 6 or 7 cells. 
Whole anal area of hind wing, including the triangle and rounded angle, shaped 
similarly to erato. Pterostigma black. Membranula dark grey, slightly lighter basally, 
extending to a little beyond the transverse cross-vein in anal triangle, as in erato. 

Abdomen with the basal and terminal inflated segments much as in erato, but inter- 
mediate segments distinctly narrower; from the base as far as the apex of 7 markedly 
expanded in lateral dimension, the next segments diminishing in breadth from the base 
of 8 as far as the end of 9, the 10th being narrower than 9 (erato: end segments 
expanded gradually and widest across segm. 10). Middorsal carina of 10th segment 
acute, extending caudad as far as about half-way length of segment but becoming in- 
distinct posteriorly. Markings of proximal segments much as in gerstaeckeri, hence much 
more reduced than in erato: transverse middorsal yellow mark of 2 isolated, constricted 
in the median line, lateral patch including the auricles narrowly detached from it; 3 with 
very narrow latero-ventral line and lacking a transverse lateral stripe at base; middorsal 
antejugal paired spots small and present only on 3, the succeeding segments unmarked; 
basal spot of 7 small, cordiform, not quite touching base or jugal suture, bluntly pointed 
caudad at the median crest; 8 with vermiform latero-ventral spot extending from the 
base obliquely upward and backward, terminating about half-way the segment’s length. 
Remainder of abdomen, inclusive of the sternites, black. 

Genitalia and lobus posterior practically identical in shape to those of erato (Fig. 10). 
Anal appendages, superior pair a little shorter and broader than in erato but otherwise 
very similar, the subterminal tooth perhaps a trifle less strong than in that species. 

Male (ad., Borneo; in fragments). — Agrees with the above description in almost 
every respect. Differs in that the labium is of a lighter brown, and in that the incomplete 
antehumeral thoracic stripe extends only half-way up the dorsum, tapering to a point. 
SEAT A 
9 LOOS NE 


Cux En Two cell-rows in discoidal field of fore wing; only a single cross-vein 


Wing membrane strongly tinged brownish-yellow. Nodal index Cux 


5.53 
44° 

Anal appendages and genital organs not differing from those of the Malayan example 
(Kiez). 

Female (Malaya). — Labium brown, the median lobe with a pair of suboval lemon- 
yellow spots coalescent at extreme base. Mandibles brown, main body with yellow external 
dot. Labrum black, with crescentic brown median spot at its base. Anteclypeus brownish- 
black, postclypeus dirty ochreous, save for a cloudy brown spot placed on each side of the 
impressed areas. Frons, vertex and rear of the head deep black, not metallic. Yellow 
markings of thorax and abdomen as in the male; the orange prejugal patch at the base of 
the 7th abdominal segment is broader than long, placed transversely, somewhat narrowed 
posteriorly and rounded off at the sides in one example, occupying slightly less than one- 
third of the segment’s length; latero-ventral border of same tergite finely lined with 
yellow and traces of yellow also basally at the lower margin on the 8th. 

Wing membrane in one old specimen strongly dark flavescent, in the second example 
hyaline Nodal indices nd ee py ae Gr 

8.10.11.8 6. 9. 9.8 DDA 227 4.4 
8.8 or 9.9 cells, without or including 1 or 2 central cells. 


6 cells in the anal loop, no central cell. 


DIO : 
Aa Anal loop with 


M. A. LIEFTINCK: Studies in Corduliidae I 25 


Valvula vulvae at 8th sternite scarcely developed, not reaching apex of the corres- 
ponding tergite; the lower margin (i.e. the upper, when looked at from below) bears a 
distinct V-shaped median incision, instead of being straight or slightly undulated, the 
rest of the thickened sternite being directed upwards to fit against the body wall as a 
pair of rounded parallel-ridges, similar in form to that in the allied species. 


Fig. 8—9. Macromia dione spec. nov. 6 holotype from N. E. Sumatra. 8, genitalia; 9, anal append- 
ages, right lateral and dorsal view. — Fig. 10—11. Macromia callisto Laidlaw, & genitalia; 10, 
from Malaya; 11, anal appendages of & from Borneo 


26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Measurements: & abd. + app. 41.0 mm, hind wing 33.8 mm, pt. fore wing 2.0 mm 
(Malaya), 41.5, 32.5, 2.0 mm, respectively (Borneo); ® 37.0—38.8, 34.0—36.0, 2.0 
mm, respectively. 

Except for the dissimilar shape of the vulvar lamina, it was found impossible to 
separate the female of callisto from that of gerstaeckeri. The individual differences in 
the venation of each have been given merely to demonstrate the existing variation and 
are evidently of no importance specifically. 

Structurally, M. callisto is most nearly related to erato. The female of the latter has the 
postclypeus entirely bright yellow, the paired dorsal spots upon the segments 3 and 4 
more enlarged, with an additional pair on 5, and differs further from callisto in that the 
ventro-apical margin of the valvula vulvae is not excised. 


Macromia dione spec. nov. 
Fig. 8—9 


Material. — N.E. Sumatra: 1 g' (adult), labelled 6594/Deli, Sumatra, L. Martin 
G. (print), Macromia ? westwood: Selys (unknown writing) (MNB). 


Of the M. septima Martin group. 

Male (holotype). — Labium dark chestnut, the basal portion of the broad midlobe 
pale yellow forming a complete transverse band which in the middle is narrowed and 
invaded by brown from the distal border. Mandibles glossy brown with large external 
yellow patch, centred with dark brown, placed upon its swollen basal portion. Labrum 
and clypeus dark brown, the anteclypeus with a tiny light brown median spot at extreme 
base; sides of postclypeus lighter brown, the median area with a pair of vestigial 
brownish-yellow spots filling out the impressions on either side of the middle. Frons and 
vertex black with low metallic blue lustre, pyramidal processes of frons not flattened or 
framed anteriorly, surface superficially and rugosely striato-punctate, sides smooth, almost 
impunctate; vertex in frontal view with raised tubercles more widely distant than in 
M. septima and even more so than in erato (fig. 1 and 5 in Lieftinck, 1950). Raised 
occipital triangle and rear of the head glossy black. 

Synthorax metallic green. Incomplete yellow antehumeral bands, ante-alar triangles and 
lateral thoracic bands as described for M. erato, the dorsal bands extending about three- 
fifths up the mesepisterna with slightly incurved and rounded apices; metaspiracular 
band broad, widest at the spiracle (1.2 mm) and almost twice as broad as this; posterior 
metepimeral band perfectly straight, parallel-sided, and only little narrower than the 
latter; ventral surface of thorax as described for M. erato. 

Legs long and slender, black; all coxae bright yellow posteriorly; hind femur 7.3 mm 
long. Fore legs for the greater part missing; tibial keel absent on mid tibia but occupying 
full length of hind tibia, though not extending quite to its base. Tarsal claw with inner 
ramus longer and more robust than apex. 

Wings unspotted, membrane faintly suffused with greyish-yellow all over, especially 
toward the tips. Costa with fine interior yellow line extending outwards from base 
GIES) oe 44 


006 on 


almost as fat as the nodus. Neuration open, nodal index 


M. A. LIEFTINCK: Studies in Corduliidae I Di, 


Two or only a single cross-vein running directly from M, to Cz, in hind wing. Only 
one large basal cell between anal triangle and anal loop of hind wing, the latter with 
only 6 cells, without central cell. Two cell-rows between loop and posterior border of 
hind wing. Distal side of anal triangle almost straight from base to tornal angle, which 
is rounded; margin of hind wing between membranula and anal angle concave. Ptero- 
stigma black. Membranula dark grey with distinct, almost pure white spot at extreme 
base. 

Abdomen slender, slightly spindle-shaped, rather longer relatively than in the allied 
species; basal segments, as well as 7 from base to apex, broadened in both dimensions, 
then parallel-sided as far as the end of 8 and finally again slightly narrowed from base to 
apex of 9. Dull black, sides of 1—3 slightly shiny and with faint dark bronze-green lustre 
in certain lights. Yellow markings greatly reduced, much as in M. gerstaeckeri and the 
Bornean M. corycia, as follows. Anterior half of segm. 2 with large oblique lateral patch, 
broadly attached to base of segment and including the auricles; this mark well distant 
from a pair of closely approximated, transverse oval spots placed dorsally in front of the 
jugal suture; there is, besides, a tiny spot situated just dorsal to the lobus posterior, which 
itself is black. Segm. 3 with vestige of a yellow lateral streak at the basal margin, a 
yellow line terminating at the jugal suture bordering the ventral margin of the tergite, 
and a pair of barely visible middorsal specks, one on either side of the median carina, 
placed also in front of the transverse suture; 4—6 unmarked; 7 with transverse orange 
marking, narrowly detached from base, longest at the median carina and with its pointed 
apex directed caudad; it extends back for about one-fourth the segment’s length, Le. 
as far as the jugal suture; laterally, the same spot descends to about one-half the height 
of the tergite, at which level it is well rounded off. Segm. 8 with conspicuous orange 
latero-ventral mark attaining exactly half the segment’s length from base, the apex of the 
spot subtruncated and about three times broader than its width at base; 9—10 black. 
Segm. 10 strongly longitudinally carinate, this dorsal crest in side view gradually more 
strongly raised towards the base so as to form a rather blunt, though distinctly elevated 
ridge (Fig. 11). 

Genitalia shaped much as in M. callisto and erato, especially the pear-shaped lobus 
posterior of 2nd segment; posterior hamuli more strongly convex in profile view, with 
the narrow and very slender distal portion a little shorter, and each of them at the same 
time more definitely outcurved, than in these species (Fig. 8). 

Anal appendages black, shorter than in the allied species, apices of superior pair only 
slightly and gradually incurved in dorsal view, the distal portion, beyond the strong 
transversely placed tooth, longer than in any of the other Malaysian species of the same 
group; bristles strong, of great length and rather numerous, especially at the blunt tips. 
Inferior appendage subequal in length to the superior pair, apex carrying two minute 
dorsal denticles (Fig. 9). 

Measurements: abdomen + app. 39.0 mm, hind wing 32.0 mm, pt. fore wing 2.0 
mm. 

Female unknown. 


A very distinct species that runs out into couplet 12 of my key (1950: 681), to near 
M. callisto and erato, with which it shares the slender form of the posterior genital 
hamulus and the pear-shaped lobus posterior. It differs from both in the more proximal 
position of the external tooth at the superior appendage, the presence of an elevated 


28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


middorsal crest on the 10th abdominal segment, as well as in details of coloration. The 
open neuration and the relatively great length of the abdomen are additional specific 
characters. 


Macromia arachnomima Lieftinck 


1953. Lieftinck, Treubia 22: 395—406, figs. 4—7, & sttuct., whole larva and larval struct. (4 
S. Borneo); 1954. Lieftinck, Treubia 22 Suppl.: 116; 1955. Lieftinck, Zool. Meded. 33: 263—266, 
fig. 14, ult larva (2 S. Borneo); 1969. Lieftinck, De Lev. Natuur 72: 103—109, figs. 2—3, whole 
larva & struct., ecol. notes (Borneo). — 1969. Furtado, Verh. Intern. Verein. Limnol. 17: 881. 


Additional material. — Malay Peninsula: 1 @ (ad.), Central Perak, Plus 
river area, Sungai Chior, low country, 7.111.1963, M. A. Lieftinck. — Borneo: 1 9 
(juv.), W. Borneo, Singkawang, stream at Montrado Rd., 24.111.1932, L. Coomans de 
Ruiter; 2 Q (ad.), Central E. Borneo, Samarinda distr., 250 m inland, 125 m, Tabang, 
Sungai Bengen (trib. of Belajan river), 4.IX and 15.X.1956, A. M. R. Wegner. 


The Malay Peninsula is a new record for this extraordinary species. The types, male 
and female, were described after specimens reared from larvae collected in southern 
Borneo (loc. cit., 1953, 1955). A second immature female from western Borneo, col- 
lected as early as 1932, had to be left unnamed by the absence of a male; this can now 
be recorded also. The discovery of the perfect dragonfly in the Malay Peninsula 
definitely confirms previous determinations of a number of larvae extracted from sub- 
merged rootlets and collected by Messrs. D. S. Johnson and J. I. Furtado (1969: 881) 
from various lowland streams (not specified here) in Selangor and Johore. 

The following notes are based on the first mature examples of either sex. 

Male (ad., Malaya). — Conforms to the type, except for the following colour differ- 
ences and other details. Mouth-parts brown, a cloudy yellowish patch at the mandible- 
bases. Clypeus uniform dark brown. Wings hyaline; rusty blackish-brown spots at 
extreme base vestigial in c-sc and cz on fore wings, slightly larger (though smaller than 
DAS Sel 255 6.6 
="; cross-veins in ht — ; CUX_ 
Cb 9 Dus) 1.1 DID 
cells in anal loop 6.6 without central cell. Anal triangle wider at base than in most other 
species; membranula almost pure white. Pterostigma black. 

Abdomen of normal length, segments parallel-sided from base of 4 almost as far as 
end of 6, thereafter gradually broadened and widest at apex of 7, then again a little 
narrowed to the end of 8; 9—10 parallel. Segment 9 of normal shape, its dorsal crest 
acute. Abdominal markings bright yellow on a deep black ground; no metallic reflections. 
Undulated middorsal band across segm. 2 subinterrupted in the median line, otherwise 
all abdominal markings as described for the type, except that the prejugal annular spot 
on segm. 7 is rather obscured basally, and there is no yellow colouring on 9—10 ven- 
trally. 

Female. — All specimens differ conspicuously from the other sex in that the dark 
spots at the bases of all wings are much more extensive: deep rusty brown in the 
immature female and extending to beyond Ax, in fore wing and as far as Axo 3 in the 
hinder pair, all surrounded by a golden yellow areola covering the entire anal area; in 
the adult females the dark spots occupy a little less of the basal cell surfaces and lack 


in the type) on the hinder pair. Nodal index 


M. A. LIEFTINCK: Studies in Corduliidae I 29 


the golden areolae. The rest of the wings in the mature examples is hyaline, except in 
one female, which has the apices cloudy yellow. Venation denser than in the males but 
indices variable; there are from 12 to 15 cells in the anal loop of the hind wing, which 
invariably includes 1—2 central cells. Head in front uniform dark brown including the 
mandibles; upper portion of frons anteriorly and above, as well as the vertex, metallic 
blue-black. Yellow juxtahumeral thoracic bands obsolete. Abdominal markings greatly 
enlarged on segm. 2, though varying in extent, occupying almost the entire anterior 
(prejugal) part of the dorsum and sides in the juvenile specimen, indented by black 
posteriorly or obliterated middorsally, in the adulti. Basal spot of segm. 7 larger than in 
male, covering basal two-fifth of segment, otherwise as in the opposite sex. 

Measurements: g (Malaya) abd. + app. 43 mm, hind wing 37.7 mm, pt. fore wing 
2.1 mm; © 45.2, 43.5, 3.0 mm (W. Borneo) and 42.5—43.5, 40.0—40.5, 2.2 mm, 
respectively (E. Borneo). 

Possibly widely distributed in the lowlands of Malaysia but so far known only from 
Borneo and the Malay Peninsula. 


GEOGRAPHICAL DISTRIBUTION OF Macromia IN MALAYSIA 


It was foreseen at the end of an earlier summary (loc. cit., 1950: 676—677) that 
explorations in Malaysia during the next 20 years would result in the discovery of some 
more species and a better knowledge of the distribution of those already described. The 
following table gives an idea of the present state of our knowledge. Of the 15 species 
listed there are still eight which are known only from one of the six geographical 
“provinces” of Malaysia here included (all, except one, insular). Endemic species are 
marked with an asterisk * 


Species Malay Sumatra Java Borneo Bangka Billiton Further 
Peninsula range 

arachnomima + + 
callisto * + 
cincta + + + Palawan 
corycia * SE 
cydippe a 4 | 
dione * 
erato * + 
euterpe * + 
gerstaeckeri + + + + ?Philippines 
jucunda * + 
mnemosyne * 
moorei fumata a. + + Celebes 
polyhymnia* + 
Septima | Tonkin 
westwoodi + + + + 
Total 7 7 8 8 2 2 


30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 
IV. THE PAPUASIAN GROUP OF Macromia RAMBUR 


Since the last revision of the Papuasian representatives of this genus (Lieftinck, 1952) 
several fresh captures of these impressive insects from all over the Papuan Region have 
come into my hands. Besides a fair number of additional specimens pertaining to earlier 
described forms, these collections yielded three (and possibly more) species new to 
science. One of these was described only few years afterwards (Lieftinck, 1955), but the 
rest still awaited closer examination. It is the purpose of this chapter to characterize the 
novelties, to give further information, where necessary, on related species, and to supply 
new locality records for the species already known. 

All regional species are very much alike and obviously closely interrelated; I have, 
therefore, merely accumulated comparative notes and, in one instance, also a key, to 
facilitate their recognition, a more elaborate descriptive table for eight of the then known 
taxa being found in my 1952 paper (loc. cit.: 438—444). The present survey deals with 
eight species known to inhabit those islands and land masses of the Indo-Australian 
archipelago which are situated east of the Philippines and Celebes. 


DAR 


13 


= ae 


Fig. 12—16. Metatarsal claws of &. 12, Idionyx montana Karsch, from 

Java; 13, Macromidia genialis erratica Lieft., from Java; 14, Macromia 

callisto Laidlaw, from Malaya; 15, Macromia celaeno Lieft., from New 
Guinea; 16, Synthemis miranda Selys, from New Caledonia 


The Australo-Papuan members of Macromia form a distinct section of the genus. All 
agree in being of large size and by having most of the body brilliant metallic green 
(except the more terminal segments of abdomen), yellow markings being much reduced: 
a single orangish patch at the base of the 7th tergite being the only conspicuously 
coloured area ornamenting the abdomen dorsally. All are characterized by the exceed- 
ingly small pterostigmata and the remarkably diminutive size of the discoidal triangle of 
the hinder wing. Special attention must be paid to the extraordinary form of the tarsal 
claws in this and probably several other sections of Macromia 1). The claws are deeply 


1) Up to now only few other species were examined upon this character, i.e. all members of the 
Malaysian section of M. septima Martin, which appear to be similar in this respect; the Palaearc- 
tic M. splendens Pictet, however, possesses normally shaped claws. 


M. A. LIEFTINCK: Studies in Corduliidae I 31 


cleft; but whereas in most other cordultids the inner ramus (or inferior tooth) is usually 
shorter and slenderer than the outer (main) branch, in them the inner branch follows 
the strong main curve, being at the same time the most robust and longest of the two, 
whilst the outer is finer, branching off slightly asymmetrically (Fig. 15). Distinctive 
characters common to all males are the slender and very narrow genital hamule and the 
sharply pronounced acute anal angle of the hind wing. Like the majority of Corduliidae 
useful criteria for the recognition of species are most evident in the males. In this sex 
the shape and armature of the 10th abdominal segment, the accessory genital organs and 
anal appendages, are quite distinctive. 

The distribution centre is New Guinea, only three species occurring outside the 
continent including its satellite islands, viz. one on Halmahera and Batjan in the northern 
Moluccas, a second in New Britain and New Ireland (Bismarck Archipelago), while a 
third, the rather enigmatic M. viridescens Till, from the Cape York peninsula in 
northern Australia, is known only from a single female. 

The regional section can be split up conveniently into three groups based on colour 
characters shared by both sexes; these do not, however, appear to reflect relationships. 


Group I of M. terpsichore Foerster 


Postclypeus with conspicuous transverse yellow band, or at least spotted with yellow 
on either side of the middle. Dorsum of thorax (mesepisterna) with a pair of clearly 
defined yellow antehumeral bands extending at least half-way up the dorsum. 

The species are: 

M. amymone Lieft., 1952 (W. and S.W. New Guinea) 

M. celaeno Lieft., 1955 (E. New Guinea; d’Entrecasteaux and Louisiade Arch.) 

M. eurynome Lieft., 1942 (S. New Guinea) 

M. hermione Lieft., 1952 (Misool I.) 

M. lachesis spec. nov. (Bismarck Arch.) 

M. sophrosyne Lieft., 1952 (Waigeu I.) 

M. terpsichore Foerster, 1900 (N. and N.E. New Guinea) 

M. viridescens Tillyard, 1911 (Cape York, N. Australia). 


Group II of M. melpomene Ris 


Postclypeus reddish- to dark brown or almost black, agreeing in colour with that of 
the rest of the head anteriorly. Dorsum of thorax metallic green lacking well defined 
antehumeral bands, but lower part of mesepisterna frequently reddish brown, dorsal- 
wards acquiring a metallic green lustre. 

The species are: 

M. astarte spec. nov. (S.E. New Guinea) 

M. euphrosyne Lieft., 1952 (Waigeu I.) 

M. melpomene Ris, 1913 (New Guinea, universal ?) 

M. spec. indet. (d’Entrecasteaux Is.). 


32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Group III of M. chalciope Lieftinck 


Postclypeus as in Group II. Dorsum of thorax with a pair of short, lanceolate, orange 
or yellow antehumeral bands tapering dorsally to a blunt point. 

There is only one species: 

M. chalciope Lieft., 1952 (N. Moluccas), with one subspecies (?) off N.W. New 

Guinea (Schouten Is.). 


It is of interest to note that the structural characters separating the members of each 
of the two main groups are allotted analogously, so that if these group characters were 
lacking, specific recognition would become much more difficult. 


Macromia amymone Lieftinck 


1952. Lieftinck, Treubia 21: 444 (& key), 446—449, 465—467, figs. 2, 6, 19—21 (& genit. & 
apps.; head & labium of larva), and 448—449 (remarks on supposed synonymy), 3 8 W. New 
Guinea (Vogelkop, Sorong). — 1915. Campion, Trans. zool. Soc. Lond. 20: 488, ¢ S. New Guinea, 
Utakwa River (M. terpsichore Foerst.). 


Further material. — S. New Guinea: 2 d 1 @ (adult), Utakwa River, Feb. 
1913, A. F. R. Wollaston Exped., over drawer-label M. terpsichore Foerst., probably 
named by H. Campion (BM). 3 & (adult), S. New Guinea, Fly River, Kiunga, 6, 14 
and 21.X.1957, W. W. Brandt (1 & BISH); 1 © (adult), Papua, Western Div., same 
area, Fly River, 5 mi. below Palmer Junction, 20.V.1936, coll.? previously named M. 
terpsichore Foerst., by the present writer. 


When describing this species in 1952, I believed Campion’s examples of “M. terpsi- 
chore Foerst.” to be wrongly associated with that species and to belong to M. amymone 
instead. Since re-examining them in the British Museum collection this supposition 
proved to be correct, as the males do agree with amymone by possessing the peculiar tuft 
of long hairs at the apex of the superior appendages and tallying with the description 
also in other characters. Now the discovery of this species at Kiunga (low country, 6° 07’ 
S lat, 141° 17’ E long.), extends its range even further eastwards so that the species is 
at present known to occur from Sorong at the western extremity of the continent 
probably all along the southcoast as far as the Fly River area. 

Male (Kiunga). — Conforming with the description and figures. Postclypeus with 
broad, uninterrupted orange band. The very broad and almost parallel-sided yellow 
stripes crossing the spiracle at the thoracic sides extend upward almost to the dorsal crest 
of the metepisternum. The orange band at the base of the 7th abdominal segment 
encircles the whole tergite and is intersected laterally only by a blackish line at the jugal 
suture, thus occupying only little less than one-half the segment’s length and on the 
ventral surface reaching even further caudad. Other specific features worth attention are 
the unusually prolonged and very acute anal angles of the hinder wings and the spindle- 
shaped abdomen, the breadth at the junction of the 7th and 8th segments measuring 
about 4 mm. 


M. A. LIEFTINCK: Studies in Corduliidae I 33 


Female (Palmer Junction). — Except for its much smaller size (the specimen from 
Sorong measuring 48.7, 49.5, 1.8—1.9 mm, respectively), the two females agree in all 
essential characters; the present specimen is a very mature one, the wing membrane being 
strongly coloured, brownish yellow all over, with the bases more deeply and diffusely 
stained with orangish. 

Measurements: & abd. + app. 49.2—49.5 mm, hind wing 47.0—47.2 mm, pt. 1.2 
mm; 9 abdomen 45.0 mm, hind wing 43.8 mm, pt. fore wing 1.2 mm. 


> 


Macromia eurynome Lieftinck 


1942. Lieftinck, Treubia 18: 563-564 (nom. nov. pro M. terpsichore Ris nec Foerster; descr. notes 
& supposed synonymy), & Kloof-bivak, Lorentz River; 1952, Lieftinck, Treubia 21: 443 (key &), 
445, Ris's 2 allotype, Lorentz River. — 1913. Ris, Nova Guinea 9 Zool. 3: 494—495, fig. 13 (2 
wings), ® Bivak-Insel, sub M. terpsichore Foerst.; 1915. Ris, Nova Guinea 13 Zool. 2: 84—85, figs. 
2—3 (genit. & apps.), ¢ Lorentz River, sub M. terpsichore Foerst. 


Material. — S. New Guinea: 4 (lectotype by present designation), Lorentz 
River, Kloof-bivak, 17.XI.1912, G. M. Versteeg, Lorentz Exp., M. terpsichore Foerst., 
det. F. Ris, ex SMF IV.1970 (MA). 

This is the male discussed by me in 1942, at that time erroneously called ‘“allotype” 
and stated to be lodged in the Amsterdam Museum. In fact, this example was shown to 
me in 1929 by the late Dr. Ris at his home in Rheinau, from where it was finally 
returned to the Amsterdam Museum via the Natur-Museum Senckenberg at Frankfurt 
a/M. 
ee species is immediately distinguished from other Papuan Macromia by the 
characteristic shape of the genital organs. Another peculiarity not mentioned before is 
the presence of a small tubercular swelling at the lower border of the second abdominal 
segment, which is beset with radiating stiff light brown bristles. An indication of this 
tiny tubercle is perceivable also in some individuals of M. melpomene and M. celaeno 
(Fig. 19 and 22). 

The lectotype and the female first described by Ris (1913) are the only specimens of 
M. eurynome so far known. 


Macromia chalciope Lieftinck subspec. ? 
Bis 18 


1952. Lieftinck, Treubia 21: 438—439 (key &), 456—460, fig. 5, 10 (& genit. & apps.), & ? 
Halmahera I. (N. Moluccas). 


Further material. — N.W. New Guinea: 1 & (slightly immature), 3 9 
(adult), Schouten Is. Biak I., Base, 29.XII.1954 (9), and Sorido, 28.X11.1954 
(2 2), all L. B. Holthuis. 

Very similar in most respects to our series from Halmahera (and an additional g' 
from Batjan I.), but a direct comparison with typical M. chalciope reveals the following 
slight differences. 


34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Male. — Lateral yellow stripe crossing the spiracle at the sides of thorax a little 
narrower and more distinctly constricted at some distance above the spiracle, the latter 
being, however, completely surrounded by yellow. Orange basal spot on dorsum of 7th 
abdominal segment distinctly larger and, though still detached from basal margin of 
segment and also more or less diamond-shaped, prolonged triangularly about half-way 
down the tergite, the spot being widest at the transverse suture; posteriorly it projects a 
little beyond the suture so as to form a tiny, arrow-shaped off-shoot, the middorsal length 
of the spot taking approximately 3/8 of the whole length: hence distinctly larger than in 
any of the 19 Moluccan examples compared. The “penis” is exerted but unfortunately 


Fig. 17—18. Macromia chalciope Lieft. subspec., 6 from Biak I. 17, genitalia; 18, anal appendages, 
dorsal and right lateral view 


damaged at the apex of the terminal segment; otherwise the accessory genitalia are 
shaped similarly to those of typical chalciope, even the lobus posterior having the same 
blunt form (Fig. 17). Anal appendages also resembling greatly those of the Moluccan 
insect, yet the superior pair are more slender, especially the distal portion when viewed 
obliquely from within, each furnished with a minute exterior tooth at about half-way 
their length (lacking in typical chalciope!); also the tips are slightly more upturned in 
the Biak specimen and do not project beyond the apex of the appendix inferior, which 
in Moluccan chalciope is somewhat longer and appears to be less strongly upcurved (Fig. 
18). The slender middorsal spine on the 10th segment is equally well developed and 
situated at the same place of the tergite. 

Female. — All three individuals are nearly identical morphologically with the Hal- 
mahera examples, the first three abdominal segments showing the same brilliant metallic 
lustre. Diamond-shaped orange spot on dorsum of 7 about twice as large as in typical 


M. A. LIEFTINCK: Studies in Corduliidae I 35 


chalciope, attached to the basal margin and not constricted directly beyond the suture (as 
in the male) but projecting posteriorly well beyond it and extending caudad to nearly 
half the segment’s entire length. Valvula vulvae shaped as described earlier, the almost 
circular lobes being equally characteristic. 

Measurements: 4 abdomen + app. 48.4 mm, hind wing 44.0 mm; © abdomen 
46.2—48.3 mm, hind wing 47.5—49.0 mm; pterostigma as in typical chalciope. 


In spite of the above differences, the present individuals are determined, with some 
reserve, as M. chalciope. More material, especially of the male, will be necessary to 
establish its proper identity, — most likely as a subspecies of the better known Moluccan 
dragonfly. This seems at present the best surmise in view of the widely different habitats 
and discontinuous distribution of the insects. 


Macromia celaeno Lieftinck 


1955. Lieftinck, Zool. Meded. Leiden, 33: 274—277, figs. 27—28 (4 genit. & apps.), &2 NE 
New Guinea. — 1900. Foerster, Termész. Füzetek 23: 86—88 (pars, excl. fig.), ¢ Kaiser Wilhelm- 
Land, Bongu (M. terpsichore n.sp., partim). — 1942, Lieftinck, Treubia 18: 563 (note on Foerster’s 
males of M. terpsichore Foerst.). 


Further material. — N.E. New Guinea: 4, Astrolabe Bay, labelled “Im Urwald 
bei Bongu bei Constantinhafen, D. Neuguinea 10—12/1898”, and “Macromia Terpsi- 
chore Foerster Type” (UMMZ); 1 g, Madang distr., Saidor subdistr., Finisterre Range, 
Kiambavi, 5000 ft., 22.VII.1958, W. W. Brandt. 2 Z., Papua, 75 km W. of Pt. Moresby, 
Doa Estate, 2.IX and 14.X.1962, R. Straatman (1 & BISH); 1 9, Papua, 60 mi. W. 
of Pt. Moresby, 17.IX.1961, R. Straatman; 2 G', Papua, Sogeri, N. of Pt. Moresby, 500 
m, 27—28.X.1969, R. Straatman; 1 2 (juv.), Papua, Kokoda, 25—29.VII.1964, R. 
Zweifel & K. Sluder, 7th Archbold Exped. (AMNH). 1 g', d’Entrecasteaux Is., Fer- 
gusson I., Agamoia, 200 m, 21.VI.1956, L. J. Brass, 5th Archbold Exped. 2 &, 
Louisiade Archip., Sudest I., Mt. Riu, west slopes, Camp 10, 250—350 m, 26 & 
29.VIII.1956, L. J. Brass, 5th Archbold Exped. 

The 10 males presently available for comparison are not quite alike. They were col- 
lected in a variety of habitats all over the eastern part of the mainland and in more or 
less isolated island groups far beyond. The structural differences observed are slight and 
possibly due to clinal variation; yet it would be unwise to ignore them. 

The original description of M. terpsichore Foerster was based on two males originating 
from the same locality. Since it became evident that these specimens were not conspecific 
(see Lieftinck, 1942: 562—563), I decided in 1957 to write to Dr. E. Kormondy, who 
at that time studied the Foerster collection in the Michigan Museum, Ann Arbor. He 
kindly complied with my request by sending me one of Foerster’s males for comparison 
with allied species, and this example proved, indeed, to be quite distinct from M. terps7- 
chore auct. We then agreed that a selection of a lectotype of M. terpsichore should be 
made. This was done by Kormondy, who wrote (August 21, 1957): “The lectotype, here 
designated, lacks the spine [on the 10th segment of abdomen} but has the protuberances 
on the 10th segment, as described by Foerster; the genitalia and anal appendages of the 


36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


lectotype agree with those figured by Lieftinck in 1942. The paratype [syntype} here 
designated, bears the spine, as described by Foerster; it also differs from the lectotype 
by bearing on each superior anal appendage a pronounced laterally directed angle at 
one-half the distance to the tip”. Meantime, a new species, M. celaeno Lieft., had been 
described from the N.E. Division of Papua (Kwagira River). As this turned out to be 
conspecific with Foerster’s terpsichore, from the eastern part of the Astrolabe Bay, no 
new name is required for the latter. It corresponds closely with the holotype of M. 
celaeno, this being only slightly larger. The latter also has the yellow thoracic markings 
and the spot on the 7th abdominal segment a little more reduced; otherwise the two are 
very similar. Unfortunately, the type of M. celaeno, and Foerster’s Bongu male as well, 
are the sole individuals known from their respective localities. They differ from all 
others in the relatively great length of the inferior appendage, which in the males from 
the Madang district (N.E. New Guinea) and those from the islands of Fergusson and 
Sudest, are only little longer than the superior pair. The type also differs from the rest 
by having a more parallel-sided and broadly rounded lobus posterior (genital lobe), 


Fig. 19—22. Genitalia of & Macromia. 19, M. celaeno Lieft., from Pt. Moresby; 20, M. lachesis 
spec. nov., from New Britain; 21, M. astarte spec. nov., from Kokoda trail; 22, M. melpomene Ris, 
from Upper Jimmi 


M. A. LIEFTINCK: Studies in Corduliidae I 37 


which in all others appears more drawn out and tapering towards a narrower apex 
(Fig. 19). 

Our males from the environs of Pt. Moresby (Doa Estate and Sogeri) are smaller in 
size and decidedly more slenderly built than the remaining specimens. They were first 
thought to represent a distinct species on account of the short inferior appendage, and 
based also on the fact that the external tooth at the superior pair is situated just a 
little more proximad than in most others, including the type. However, since we know 
little still about the amount of individual variation existing with regard to the above 
characters, I hesitate to lay too much stress on them as specific criteria. To take an other 
example: the presence or absence of a bristled tubercle at the base of the ventral border 
of the 2nd abdominal tergite, figured for one of these southern males (Fig. 19), does 
not seem to be a reliable character either: it is present in all of them, but barely indicated 
in the type and some other males. 

Summarizing the above, it seems best for the present to consider M. celaeno a polytypic 
species that can, perhaps, be split up later into a number of geographical races. 

Female. — A full description of the topotypical specimen is found in the original 
description. The female from the Pt. Moresby area, though being a good deal smaller, 
agrees with it in all essential characters and so does the Kokoda specimen, which is 
rather immature; as to its dimensions the latter nearly corresponds with the allotype. 
Wings hyaline; bases diffusely yellow about as far out as Ax; tips of fore wings deep 
golden yellow from a little beyond nodus as far as the apex. Anal loop of hind wing 
enclosing 14—16 (Pt. Moresby) or 18—21 cells (Kokoda). 

In the fully adult Pt. Moresby specimen the leaf-like blades of the genital valve are a 
little broader than in the allotype and extend back about three-fourths the length of the 
9th tergite, measured along the middorsal crest; they are directed obliquely ventrad and 
caudad, each blade being hollowed out with upcurved tip. Anal appendages hairy, 
distinctly outcurved; tips slender, acuminate. 


Measurements (in mm) 


abd. + app. hind wing pterostigma 
d' Kwagira River (holotype) 49.8 44.4 1.4—1.5 
g Madang 48.4 44.0 1.3—1.4 
& Bongu (syntype terpsichore) 47.8 43.0 15 
d' Fergusson I. 50.0 45.0 1.9 
d Sudest I. 48.0 45.2 1.3 
d' Sudest I. 48.0 46.0 1.4 
d' Papua (Sogeri) 45.8 42.9 1.2 
d' Papua (Sogeri) 45.5 42.7 1.3—1.4 
d' Papua (Pt. Moresby) 45.8 41.8 1.2 
d' Papua (Pt. Moresby) 44.3 41.3 12 
9 Kwagira River (allotype) 51.0 49.3 1025) 
Q Papua (Kokoda, juv.) 50.0 50.0 1.0—1.2 


9 Papua (Pt. Moresby) 45.5 46.0 152 


38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Macromia lachesis spec. nov. 
Fig. 20, 25—26 


1942. Lieftinck, Treubia 18: 562—563, pars: 9 Herbertshöh, New Britain, sub M. terpsichore 
Foerst.; 1949. Lieftinck, Treubia 20: 361, no descr., sub M. terpsichore Foerst. 


Material. — Bismarck Archipelago: 7 & (one imperfect) 1 9, New 
Britain, Gaulim, 130—160 m, 20—28.X.1962 (2 & 1 9) and Illugi, 8—15.XIL.1962 
(5 &), all J. Sedlacek. Holotype g', New Britain, Illugi, 8.X11.1962, J. Sedlacek; para- 
types d and ® (BISH), from the above localities. 2 g', New Britain, Yalom, 1000 m, 
15.V.1962, Noona Dan Exped. 1961—62 (MC); 1 &, New Britain, 15 km SE of 
Kokopo, 10.VII.1962, Noona Dan Exped. 1961—62. 1 9, New Ireland, Danu, Kalili 
Bay, 29.1V.1962, Noona Dan Exped. 1961—62 (MC). 

Very similar to M. celaeno Lieft. and obviously nearly related with it. 

Male (ad., holotype). — Head a little smaller than in celaeno, width across the eyes 
8.9 mm; shape, texture and colour of face, frons and vertex not differing from that 
species. Labium brown, generally darker than in celaeno. Labrum and mandibles deep 
black, surface shiny. Anteclypeus dark brown with an indistinct greyish-yellow spot upon 
middle. Postclypeus blackish-brown marked with a pair of large, bright chrome lateral 
spots, each of which occupies the whole of the side portions save the anterior border 
narrowly, the latter remaining black; these transverse spots are more sharply defined than 
in celaeno, each tapering to a point inward, the distance separating them subequal to, or 
a little less than, their own breadth. Frons and vertex brilliant metallic green. Occipital 
triangle black, raised slightly above level of compound eyes, the lateral margins acute, as 
in the allied species; rear of the head glossy black. 

Thorax brilliant metallic green; yellow markings as in ce/aeno, median crest, ante-alar 
triangles (except black anterior rims) as well as a transverse lanceolate spot on each side 
in front of the latter, chrome; metaspiracular stripe moderately broad, a little shorter than 
in celaeno, leaving off fully 1 mm before reaching dorsal margin of metepisternum; 
latero-ventral border of metepimeron more narrowly yellowish but ventral surfaces 
coloured as in that species. 

Legs a little shorter than in celaeno, coloured as described for that species; hind femur 
(incl. troch.) 10.0 mm long, not reaching beyond apex of lobus posterior of genitalia. 
Keel on flexor side of fore tibia extending less than half (41%) the total length, i.e. 
a little shorter than in celaeno, armature otherwise similar, the inner ramus of the tarsal 
claws equally robust and longer than the apex itself. 

Shape and neuration of wings, including the anal area and excavated angle, hardly 
differing from the allied species (see description of M. celaeno), but as compared with 
the latter the venation is somewhat denser and the membranula is of a darker grey. 

CRTC AR 4. 4 5.6. 
Nodal index ; cross-veins in ht — M ; anal loop with 11 and 9 cells, 
TOA al 44 
with or without central cell. Pterostigma brownish-black, almost black. 

Abdomen relatively shorter than in celaeno, with the intermediate segments less 
constricted and the terminal ones more gradually expanded: in celaeno the greatest 
breadth of the abdomen is attained at the apex of the 9th segment, whereas in Jachesis 
it is widest at the end of the 7th, the 8th being almost parallel-sided while the apical 


M. A. LIEFTINCK: Studies in Corduliidae I 39 


Fig. 23—24. Macromia celaeno Lieft., from Pt. Moresby, & anal appendages. 23, left side; 24, 
dorsal view. — Fig. 25—26. Macromia lachesis spec. nov., from New Britain, d anal appendages. 
25, left side; 26, dorsal view 


segments again diminish gradually in width toward the end. Moreover, the segments 8, 
9 and 10 are markedly less highly crested when viewed from the side, 8 and 9 being less 
protuberant posteriorly. Colour and markings not different, except that the yellow spots 


40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


on 2—3 and those on the ventral surface of 8 are even more restricted; 9 and 10 entirely 
black. Orange spot a base of 7th tergite occupying roughly the upper two-thirds in lateral 
view, its lower border convex; posteriorly, the spot is narrowed, projecting apicad beyond 
the jugal suture as a low triangle, the coloured area extending back for about two-fifth 
of the segment’s length. Strong, short, reddish brown bristles are present and directed 
ventrad along basal one-fourth of lower margin of 2nd tergite, but there is no small 
marginal tubercle, as is seen in the majority of celaeno. Segm. 10 dull black, not metallic, 
lacking the shine of ce/aeno; middorsal carina running from base to apex, strong, acute 
and abruptly produced beyond half-way its length into a laterally compressed triangular 
boss ending in a short spine, the whole prominency distinctly smaller than in celaeno 
(Fig. 25). Genitalia as figured for one of the paratypes (Fig. 20); posterior lobe 
markedly shorter and more bluntly rounded than in any ce/aeno, even including the type, 
shorter than its width at base. Hamulus similar in form to that of the type of celaeno 
though straighter than in most other individuals referred to that species, one of which 
is here figured (Fig. 19). Anal appendages relatively small, superior pair straighter and 
less outcurved apically than in celaeno and with indication only of a minute external 
tooth a little before half-way its length; internal fringe of subapical hairs of normal 
length and density. Appendix inferior somewhat longer than the superiors, less strongly 
upcurved than in celaeno (Fig. 25—26). 

Female (New Britain). — Similar to male except for the following differences. 
Anteclypeus wholly black; postclypeal spots more closely approximated mesiad, reddish- 
brown area separating inward extensions about 1.5 mm broad. Yellow lateral thoracic 
stripe broader, almost parallel-sided, barely attenuated above the spiracle. Wings clear 
hyaline, bases of all strongly saffronated as far as midway between Arc and #7 in all 
wings but not entering anal loop in hind wing, this colour disappearing gradually out- 

9.18.18.10 


wards. Neuration closer than in male, nodal index"; cross-veins in hf —: 
IS Dd 


Cux 2, anal loop comprising 18 and 18 cells. 


Valvula vulvae shaped exactly as described for celaeno, though the blades appear to be 
a trifle broader with bluntly pointed apices. Anal appendages very small, acuminate. 

Measurements: & (holotype) abd. + app. 47.7 mm, hind wing 43.2 mm, pt. fore 
wing 1.0 mm; & (paratypes) 44.0—50.8, 41.0—47.8, 1.1—1.3 mm, respectively; 9 
(New Britain) abd. 44.3, hind wing 46.8, pt. fore wing 1.4 mm; ® (New Ireland) 
45.5, 47.5, 1.8 mm, respectively. The dimensions of the ® from Herbertshöh cannot 
now be given. 


A fairly homogeneous series showing little variation except as regards the dimensions. 
In the female from New Ireland the dorsal spot at the base of the 7th abdominal segment 
is oval, widest at its middle and placed in the long axis of the body; it occupies only the 
upper one-fourth of the tergite in lateral aspect. Unlike M. celaeno, this sex shows no 
sign of a postnodal golden yellow area on the fore wings. M. lachests, along with 
celaeno, runs out in my key (1952: 441) to near sophrosyne from Waigeu Island. The 
males of these three species can be held apart most satisfactorily by a comparison of their 
anal appendages and accessory genital organs, illustrations of the parts being now 
available for all. Here again, the differences are slight and should be evaluated in con- 
junction with other characters, such as body form and markings. 


M. A. LIEFTINCK: Studies in Corduliidae I 41 
Macromia viridescens Tillyard 


1911. Tillyard, Proc. Linn. Soc. N.S. Wales 36: 380—381, pl. 10 fig. 11 (pterostigma) ® Cape 
York; 1913. Tillyard, ibid. 37 (1912): 584, sub M. terpsichore Foerst. — 1942. Lieftinck, Treubia 
18: 563—564, note = ? M. eurynome Lieft.; 1952. Lieftinck, ibid. 21: 444—445, holotype 
redescribed. — 1960. Fraser, Handb. Drag. Austr.: 49, sub M. terpsichore Foerst. — 1968. Kimmins, 
Bull. Brit. Mus. nat. Hist. (Ent.) 22: 301, note on holotype (& err. pro 2). 


Apparently known only from the type. 

Although it would appear from Fraser's kev (Joc. cit. 1960) that he had seen more 
Australian specimens of this dragonfly to justify his opinion that viridescens is only a 
synonym of terpsichore, there is no proof that Fraser had actually seen the insect. Most 
likely M. viridescens is specifically distinct from the known Papuan members of the 
genus. 


Macromia melpomene Ris 
Fig. 22, 29—30 


1913. Ris, Nova Guinea 9 Zool. 3: 496—499, figs. 14—17 (4 genit. & apps, ¢ 2 wings), 
Lorentz River territ., S. New Guinea; 1915, Ris, ibid. 13 Zool. 2: 85 (no descr.), Beaufort River, 
same area. — 1929, Lieftinck, Tijdschr. v. Ent. 72: 63 (key), 68—69; 1942, Lieftinck, Treubia 18: 
564-565 (addit. descr., variation & habits), 4 $ N. & Central W. New Guinea; 1952, Lieftinck, 
ibid. 21: 441 (descr. key), 461 (key) & 464—465, figs. 14—15, 17 (descr. & larval struct.). 


Additional and re-examined material. — New Guinea: & (holotype), Nova 
Guinea Exp. Lorentz, Bivak No. 3 [= Heuvelbivak, between Beaufort and Lorentz 
Rivers in Went Mts.}, 700—750 m, 12.XI (rect. X) 1909, M. melpomene n. sp. Ris 
Type &, det. F. Ris (MA); © (allotype) with same labels, 13. XI (rect. X) 1909, det. 
F. Ris (MA); 1 g 1 ©, same region, Beaufort River, 12.XII and 8.X11.1912, Lorentz 
Exp., M. melpomene Ris, det. F. Ris, ex SMF IV.1970. Series g' 9, N. New Guinea, 
Cycloop Mts. and localities in Humboldt Bay area, sea level to 1100 m, various dates, 
19311938, W. Stüber & L. J. Toxopeus; 1 g', Central W. New Guinea, Idenburg 
River valley, Rattan Camp, 1150 m, 10.11.1939, and 8 3, Araucaria Camp & River, 
800 m, III—IV.1939, L. J. Toxopeus (ML & AMNH). N.E. New Guinea: 2 dg’, 
Central region, Upper Jimmi Valley, Tsenga, 1200 m, 14.VII.1955, J. L. Gressitt (BISH 
& ML); 1 9, Western Highlands, Adalbert Mts., W. of Madang, Wanuma, 750—1050 
m, 25—29.X.1958, J. L. Gressitt (BISH).. 
| 

As I have pointed out earlier (1942: 565), this species varies somewhat in respect of 
dimensions and in the development of a minute extero-lateral denticle at the male 
superior appendage, which latter occasionally may be obsolete. Variation also exists in 
the extent of the basal mark upon the 7th abdominal segment. This occupies almost the 
basal half of the dorsum in both male and female from the typical locality but is much 
smaller in northern individuals and in those from the Upper Jimmi valley in the east, 
where the spot barely projects beyond the jugal suture. These eastern specimens differ 
from the others also in that the lateral thoracic stripe is much narrower, being hardly 
0.5 mm broad at its upper portion. In them the posterior hamule is not so strongly 


42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


curved as in the type, and inone the dorsal process of the 10th segment is less com- 
pressed, approaching the form it has in M. astarte spec. nov.; this is, however, a smaller 
and more obscurely coloured insect. 

I must confess not to feel sure about the correctness of the identification of the insects 
from north-central and eastern localities in New Guinea, reported above. Unless M. mel- 
pomene, like some others, varies individually all over the continent and even within a 
restricted area, all of them may prove ultimately to represent geographical subspecies, 
each with a limited range, or even belong to distinct species intimately allied with it. 

The measurements of the examples from N.E. New Guinea are: & abd. + app. 
49.0— 50.0 mm, hind wing 47.0—47.5 mm, pt. fore wing 1.1—1.2 mm; 9 48.5, 
49.0 and 1.3 mm, respectively. The genital organs and anal appendages of a Tsenga 
male are here figured (Fig. 22, 29—30). 


Macromia spec. indet. 


Material. — D’EntrecasteauxIs.: 2 © (ad.), Goodenough I., 900 m Camp, 
East slopes, 25 & 26.X.1953, K. M. Wynn & L. J. Brass. 1 @ (ad.), Normanby I., Mt. 
Pabinama, 820 m, 10.V.1956, L. J. Brass. 

This is another near ally of M. melpomene, collected by members of the 4th and 5th 
Archbold Expeditions in the islands east of the Papuan mainland and received by 
courtesy of the American Museum of Natural History, New York. The specimen from 
Normanby is an aged individual having the entire wing membrane coloured a smoky 
brown. The two females taken on Goodenough have hyaline wings with a cloudy yellow 
spot occupying most of the postnodal portion of the fore wings. They are darker than 
the allotype female of M. melpomene and even superior in size to the latter, differing 
also by having an almost black pterostigma. 

By the absence of a male I prefer to leave these examples without a name. Their 
proportions are: abd. 49.0—50.0, hw. 49.0—51.0, pt. fore wing 1.2 mm (Goodenough), 
50.0, 49.0, 1.2 mm, respectively (Normanby). 


Macromia astarte spec. nov. 
ge, Di, 2728 


Material. — SE. New Guinea: 4 8 (adult), Papua, Kokoda trail, 700 m, 
14.X.1969 (1 8) and 4—7.X1.1969 (3 G'), R. Straatman; 1 9 (semi adult), Papua, 
Sogeri, N. of Pt. Moresby, 500 m, 3.XI.1969, R. Straatman. Holotype g', Kokoda trail, 
5.X1.1969, R. Straatman. Paratypes: 3 & from Kokoda trail (BISH and ML); Q from 
Sogeri. 

As compared with the two males (type and paratype) of M. melpomene that are 
known from the hill-country of southwest New Guinea, this new species is readily 
distinguished by its smaller size, slenderer forms, much narrower metepisternal stripe, 
and reduced orange marking at the base of the 7th abdominal tergite. The structure of 
the facial parts, leg armature, wing venation and body colour, are all so much alike in 
the two species that there is no need for descriptions. The principal features by which 
the two can be held apart are summarised below and are best appreciated by a comparison 
of the existing illustrations, including those given by Ris (1913). 

M. astarte. — Smaller, & abdomen + app. not exceeding 46 mm, hind wing 4344 


M. A. LIEFTINCK: Studies in Corduliidae I 43 


2 
WG 


Mf m 


Fig. 27—28. Macromia astarte spec. nov., from Kokoda trail, & anal appendages. 27, left side; 28, 
dorsal view. — Fig. 29—30. Macromia melpomene Ris, from Upper Jimmi, & anal appendages. 
29, left side; 30, dorsal view 


44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


mm. Metepisternal yellow stripe crossing metaspiracle linear, 0.2—0.4 mm, widest at 
and not broader than the greatest diameter of the spiracle, tapering on either end and 
separated from upper margin by a space twice broader than its own width. Posterior 
hamuli slightly twisted lengthwise, converging from base to apex, tips more nearly 
parallel, not outcurved (Fig. 21). Basal half of 10th segment deeply impressed on either 
side of median carina, which turns up abruptly to form a slender spine which is com- 
pressed only at its base. Orange basodorsal mark of segm. 7 small, irregular in shape, 
whole spot occupying about two-seventh of the segment’s length. Sup. anal apps. 
perfectly straight in side view, external tooth vestigial or absent, extremities a little 
upcurved and only slightly outbent, internal bristly hairs not much longer than breadth 
of apex (Fig. 27—28). Pterostigma generally brownish-black or black; membranula dark 
grey. 

M. melpomene. — Larger, 4 abdomen + app. 49—52,5, hind wing 44—49 mm. 
Metepisternal yellow stripe broader and more nearly parallel-sided, 0.4—0.8 mm, broader 
than greatest diameter of metaspiracle, separated from upper margin by a space subequal 
to its own width. Posterior hamuli still more slender, not twisted, usually converging from 
base as far as about three-fourths their length, then frequently a little diverging with 
slightly outcurved tips (Fig. 22). Basal half of 10th segment as before but median 
carina more pinched and gradually raised to form a compressed triangular boss of 
variable shape and height. Orange basodorsal mark of segm. 7 very variable but usually 
larger and more regular, whole spot in the type almost reaching half the segment’s length 
middorsally. Sup. anal apps. very similar to astarte but tips a little more upcurved and 
distinctly more outbent at the extremities, internal bristly hairs more conspicuous and 
longer than breadth of apex (Fig. 29—30). Pterostigma generally light to dark reddish- 
brown; membranula lighter, greyish white. 

VISI 8 7.163166 8.16.16. 8 


The nodal indices are: = (_ holotype), ; and 
10.10. 9.11 8.11.10.10 Wily Ge Oki 
VD 7 6.17.16. 8 
10. 9.10.10 (9 Paratypes); Go C°) 


The only, slightly immature, female has a narrow lateral metaspiracular stripe that 
widens a little at level of spiracle but otherwise is scarcely broader than the greatest 
diameter of the latter. Wings hyaline with small, diffuse, golden yellow spots barely 
reaching Ax, at base, and with a conspicuous apical patch of the same colour filling out 
the apex of the fore wings as far inward as Px,. The anal loop of the hind wings 
comprises 16 and 15 cells, each including 3 central cells. Yellow lateral spot at jugal 
suture of 2nd abdominal tergite shaped like an inverted comma; orange dorsal mark of 
the 7th broadly attached to base of segment, then widening gradually as far as the jugal 
suture and beyond the latter tapering abruptly to a pointed triangle. The rounded leaf- 
shaped blades of the valvula vulvae are very broad, attaining a little over one-third the 
length of 9th sternite, and are separated by a deep V-shaped cleft almost reaching the 
base. 

Measurements: & (holotype) abd. + app. 45.0 mm, hind wing 43.0 mm, pt. fore 
wing 1.0 mm; & (paratypes) 43.2—46.0, 43.8—44.0, 1.2 mm, respectively; Q abd. 
47.0, hind wing 47.0, pt. fw. 1.2 mm. 

As explained before, populations of M. melpomene from geographically remote 
habitations show slight differences in one way or other; but considering all characters of 
both, there can be no doubt that astarte should be regarded as a distinct species. 


M. A. LIEFTINCK: Studies in Corduliidae I 45 
V. REMARKS ON THE SUBFAMILY SYNTHEMISTINAE 1) 


Following Tillyard’s classification in the “Biology of Dragonflies’ (1917), the much 
discussed and very diversified Synthemis group is here segregated from the Corduliidae 
(or Corduliinae of Tillyard) and placed next to the Macromiinae (or Macromiini of 
Tillyard) in the subfamily Synthemistinae (or Synthemini of Tillyard). In an attempt to 
unravel the phylogeny of the Corduliidae, Tillyard (1910) had already placed the 
Synthemis group at the base of the macromian assemblage, with which there exist 
undeniable affinity. The above units are thus merely given subfamily instead of tribal 
rank. As regards venation, its members present a mixture of very archaic and secondarily 
developed characters. For example, the alternate strength of veins in the antenodal 
complex, incomplete basal antenodal cross-nerve and constant presence of cross-veins in 
the median space of the wings are without counterpart among associated genera. Several 
authors have emphasized the remarkable similarity between the larvae of Synthemis 
(sens. lat.) and Chlorogomphinae among the cordulegasterids, and the great phylo- 
genetic importance of this. 

As with Macromia, considerable differences exist between tropical species of Syn- 
themis and those living in the subtropical or temperate climatic zones. This clearly finds 
expression in corresponding differences in colour pattern, mode of flight, habitat selec- 
tion and general behaviour. The larvae are as variable morphologically as are the mature 
dragonflies but relationships among the various specific groups are equally difficult to 
interpret. 

Publications of special interest dealing in particular with Synthemis and allied genera 
(or subgenera), subsequent to Tillyard’s monograph (1910), are those given by Fraser 
(1960), Lieftinck (1935b, 1953a) and Watson (1962, 1967). The last-mentioned 
papers also contain valuable information on the immature stages of the species occurring 
in South-west Australia, including good pictures of whole larvae and details of their 
structure. 


VI. NOTES ON Synthemis SELYS FROM NEW CALEDONIA, WITH AN ACCOUNT OF 
THEIR LARVAE 


The last general information on the Odonata of New Caledonia is contained in a 
synopsis published by Campion (1921), who on the same occasion revised the other 
Corduliidae then known to occur in this Oceanic island. Campion was the first to 
describe the male of Synthemis miranda Selys and re-characterized both its sexes. In the 
same paper he supplied descriptions of three additional species, all of them new to 
science. These descriptions are accompanied by good figures and beautiful wing-photo- 
graphs of all New Caledonian taxa, although the females of two of the novelties had to 
remain unknown. 


1) It has been pointed out by Cowley (1942: 63) that the names of all suprageneric taxa based on 
generic names ending in -themis should be emended. The present writer has accepted Cowley’s 
suggestion to adopt themist- as being the Homeric form which occurs in names like Themistocles, 
instead of regarding the stem to which the group-ending is added as them-, the former selection 
being the more correct one. 


46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


In the following pages some more data can be supplied concerning the imagines of 
New Caledonian Syrfhemis, with descriptions of the females of two species already 
dealt with by Campion and both sexes of one previously unknown. I have taken the 
present opportunity also to give an account of the highly peculiar larvae of two dif- 
ferent species, with drawings of their most remarkable morphological parts. Campion 
(loc. cit.) had already given a brief diagnosis of an unidentified Synthemis larva, the 
first of its kind found in New Caledonia. This larva was like that of the Australian 


Fig. 31—32. Proventriculi of ultimate instar larvae of New Caledonian species. 31, Synthemis fenella 
Campion (supposition); 32, S. miranda Selys (supposition), showing dentition of dorsal and ventral 
plates from within 


M. A. LIEFTINCK: Studies in Corduliidae I 47 


Eusynthemis guttata in having the median lobe of the labium produced anteriorly and a 
conspicuous semicircular plate projecting from the frons. In other ways, however, it 
failed to agree with that of any other known species in the subfamily, especially on 
account of its smooth body and scantiness of setae at the labial palpus. These features 
are manifest also in the two kinds presently discussed, but all three are easily distin- 
guished from one another and obviously belong to three distinct species. At the same 
time they can be distinguished from most, if not all, other congeners from outside New 
Caledonia. 

The proventriculus (gizzard) of the two available species was cut open longitudinally 
and spread out, showing from within the set of four teeth-bearing folds arranged in 
bilateral symmetry. The armature of the plates is much alike in the two species described 
(Fig. 31—32) and needs no further explanation, except that their shape conforms in 
almost every detail to that shown by a host of other Corduliidae and Libellulidae, 
inclusive of Macromia and Idionyx among members of the first family. 


Synthemis miranda Selys 
F10.916032 


1871. Selys, Bull. Acad. Belg. (2) 31: 557—558 (121—122 sep.), 2 Nouvelle Calédonie. — 
1921, Campion, Ann. Mag. Nat. Hist. (9) 8: 34, 47—55, figs. 6—8 (4 and ® app. & ovip.), 
pl. 8 fig. 12 (& wings.). 


Besides a number of specimens which had been examined at an earlier date and 
returned to the Bishop Museum (Honolulu), the following are still before me. 

Material. — New Caledonia: 2 g, Bourai-Houailou Rd., III.1959, N. & L. H. 
Krauss; 1 ©, La Foa, 17.111.1961, J. Sedlacek; 1 ©, La Crouen, 16.III.1961, J. Sed- 
lacek; 1 9, Forêt de Thi, 550 m, 6.111.1960, J. L. Gressitt; 1 &, Mt. Mou, 11.11.1962, 
N. & L. H. Krauss; 1 g', Col de Pirogue, 13.11.1962, N. & L. H. Krauss; 1 5, Nakety, 
400 m, 23.XII.1963, R. Straatman; 1 g', Col de Rousettes, 550 m, 6.11.1963, J. L. 
Gressitt; 1 9, Mt. des Sources, 550 m, 30.XII.1963, R. Straatman; 1 9, Canala, 
23.XII.1963, R. Straatman; 2 9, Pouébo, 12.1.1964, R. Straatman. (All in BISH & 
ML). 

Both sexes of this extremely handsome dragonfly have been discussed at some length 
by Campion, who gave good figures of the terminalia of either sex as well as a wing- 
photograph of the male. The same author commented upon the variation in extent and 
depth of the coloured areas on the wings of the female. I have examined good series of 
this species from various localities in New Caledonia and it seems that S. miranda is the 
least rare of its large-sized congeners inhabiting the island. From the present series it 
would appear that there are two main colour forms of the female, which differ markedly 
from one another as far as the wing colour is concerned, evidently quite independent 
from the age of the individual, intermediates occurring at the same time, though ap- 
parently more rarely. 

Following Campion’s notes, some additional characteristics can now be given, which 
are based on five males and seven females. 

Male. — Agreeing with Campion’s decription of the allotype but for the following 
details. Anteclypeus dirty yellow, postclypeus in the middle likewise, but lateral divisions 


48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


wholly cream-coloured. Frons metallic blue-black with a pair of small creamy oval spots, 
one on either side towards the eye-margin, and often also a very small, slightly impressed 
yellow spot on each of the frontal tubercles anteriorly. Creamy yellow bands at the 
thoracic sides complete, the one at the first suture leaving off shortly before reaching the 
dorsal crest, the metepimeral stripe slightly curved and complete. Legs black, the coxae 
and trochanters somewhat more brownish; fore tibia with yellow-green interior stripe, 
incomplete basally. Tibial keels whitish, extending along about distal one-half on fore 
and mid tibiae, along distal 7/9 on hind tibia. Pterostigma deep black. 

Abdominal segment 1 usually with two small yellow dots low down at the sides, 2 
with a broad subinterrupted yellow stripe bordering lower margin of tergite, and 5 
occasionally with additional tiny yellow spot on either side at extreme base. Genital 
organs black; hamulus strongly protuberant, thick and subcylindrical for its basal three- 
fourth, then rather abruptly narrowed, directed caudad and tapered, with slender out- 
curved tip. 

Female. — All specimens are fully coloured. Three of these differ from the type in 
respect that the deep golden area in the wings in no case extends beyond the level of 
the arculus, though in all of them brown rays are present also in sc, fading away at 
about the same level. In three other females the wing membrane is as conspicuously 
coloured as in the holotype: in them not only the bases of fore and hind wings are 
deeply stained with golden yellow as far as the arculus and rich brown in sc to beyond 
the nodus, but there are also broad oblique brown bands which extend as far out as 
about midway between base and nodus. One specimen (from Pouébo) in this respect is 
exactly intermediate between the extremes just mentioned. 

The shape of the ovipositor valves corresponds with the description given by 
Campion, inasmuch as the paired outer and inner processes, although fused together for 
their whole length on each side of a narrow V-shaped emargination, are still recog- 
nizable by a longitudinal furrow separating them. 

Measurements. — 4 abdomen + app. 48.0—51.0 mm, hind wing 39.0— 42.0 mm, 
pt. fore wing 2.5—2.8 mm; 9 abdomen 50.0—53.0 + 1.5—1.7 mm, hind wing 
46.0—47.5 mm, pt. fore wing 3.0 mm. 


Synthemis campioni spec. nov. 
Fig. 33—35 


Material. — New Caledonia: & (adult, holotype), N.E.-side, Pouébo, 200 m, 
24.1.1964, R. Straatman; & (subadult), Mt. Mou, 11.11.1962, N. & L. H. Krauss 
(BISH); 9, La Crouen, 16.III.1961, J. Sedlacek; 2 (adult), N.E.-side, Pouébo, ca. 
250 m, near mountain stream, 23.1.1964, R. Straatman (1 ex. BISH). 


Very near S. montaguei Campion and of the same slender build. 

Male (holotype). — Labrum ochraceous-orange, broadly bordered with black; low 
median longitudinal ridge also blackish anteriorly. Clypeus olivaceous-brown, the post- 
clypeus with oval pale greenish-yellow spot on either side. Frons anteriorly and round- 
about the prominent tubercles blackish brown, marked each side with a squarish pale 
greenish-yellow spot towards margin of compound eye; anterior surface of tubercles 
red-brown. Vertex brownish-black. (In S. montaguei the light areas of labrum, post- 


M. A. LIEFTINCK: Studies in Corduliidae I 49 


clypeus as well as those upon summit and sides of frons are definitely larger, more 
sharply delimited and bright greenish-yellow in colour; the roundish spots on top of 
frontal tubercles are well marked off posteriorly from a thick bluish-black band at the 
base of frons). 

Pro- and synthorax metallic brown, less obscured than in S. montaguei and with 
metallic green reflections less evident on mesepisternum and posterior to first lateral 
suture. No yellow markings at spiracle and first lateral suture (S. montaguei: an undul- 
ated yellowish stripe dividing mesepimerum and metepisternum, separated from a 
yellow spot placed a little more ventrad at the spiracle). 

Legs lighter brown than in S. montaguei, femora less obscured apically. Tibial keels 
as in that species, extending along distal one-half on fore and middle pair and a little 
more than distal three-fourths on hind tibia. 

Wings hyaline with rusty brown spots, ill-defined distally, in c-sc as far as the basal 
postcostal nervure, and traces of similarly coloured spots in m and cz. Shape of wings 
very similar to S. montaguez; neuration dense. Fore wings with 22 Ax of first series and 
13—15 Px, hind wings with 15—16 Ax and 17 Px. Triangle with 2 cross-veins in fore 
wing, only 1 (or # 3-celled!) in hind wing; # 3-celled in fore wing; ht with 3 cross- 
veins in fore wings, 2 in hinder pair; m with 5 in fore wings, 4 in hind wings; cw with 
8—9 in fore wings, 7—8 in hinder pair. Bridge cross-veins 7—8 in fore as well as in 
hind wings. Discoidal field of fore wings commencing with 4 cells, then expanded and 
followed by double cells up to a point one cell distal to the separation of M, 9; 15—18 
marginal cells. Primary (distal) anal loop consisting of 6—7 cells without central cell, 
the secondary (proximal) enclosure with 4—5 cells. Pterostigma a little longer than in 
S. montaguei, without brace vein; colour dark brown. Membranula white (smoky grey 
to black in adult S. montague7), extending as far as the cross-vein in anal triangle. 

Shape and markings of abdomen much as in S. montaguei but all light spots a little 
smaller, less sharply defined and of a darker yellow tint. Bases of segm. 2—5 unmarked, 
lacking the paired transverse yellow streaks, interrupted in the median line, of S. mon- 
taguei; paired middorsal subcircular spot at the jugal sutures of 2—8 present and similar 
to that species, except that they are more roundish anteriorly, projecting markedly beyond 
the suture; middorsal spots on 8 subtriangular and a little smaller. Segm. 9—10 blackish 
brown. Genitalia (Fig. 33) prominent; basal two-thirds of hamulus straighter and more 
inflated at extreme base than in S. montaguei, undulated in profile view, the apical 
portion weakly S-shaped with curled, outbent tips; shiny dark brown, the slender distal 
part reddish brown. Posterior division of vesicle in caudal view circular and hollowed 
out deeply so as to form a tiny yellowish brown crater-like pit (similar to that of 
S. montaguei). Anal appendages very long, superior pair distinctly longer than segm. 
9 + 10 (ratio ca. 36 : 26) and also longer than in S. montaguei (ratio ca. 32 : 27); in 
dorsal view the superiors are shaped similarly in the two species but in profile those of 
S. campioni ate straighter, neither upcurved nor flattened, and depressed in distal half, 
but rather sinuous and swollen towards apex; colour uniform brown (not yellow in 
distal two-thirds!) with slight darkening at extreme base and a blackish obscuration all 
along inner border from a point slightly beyond half-way their length to apex; bristles 
black. Appendix inferior less upcurved than in S. montaguei; colour brown (Fig. 34). 

Female (adult). — Very similar to the male but differing from it in the following 
respects. Pale areas of labrum ochraceous-orange, the free margin less broadly black; 
clypeus and base of frons brown, the lateral pale spots dirty ochreous; swollen tubercles 


50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


Fa ) 

: S N 

NOONE 
NX \ > 
VOR DIE 


Fig. 33—34. Synthemis campioni spec. nov, & from New Caledonia. 33, genitalia, right side; 34, 
anal appendages, right side and dorsal view 


of frons orangish, turning darker with distinct metallic blue reflections posteriorly and 
in the sulcus. Vertex metallic blue-black. Occipital triangle brown, its side margins raised 
above level of compound eyes but dorsal surface and posterior division convex. 

Pro- and synthorax as well as the legs as in male, except that there are no tibial keels. 
Wing membrane hyaline. In one specimen the extreme bases of both fore and hind 
wings are strongly spotted with dark rusty brown in c-sc, m and cz, these spots being 
ill-defined outwards and in sc extending from base as far as the first complete Ax, in the 
other spaces only about half-way into the basal cells. In the second female the basal 
spots are greatly enlarged, in c-sc reaching as far as Ax7 in fore wing and Axg in hind 
wing, in m as far as Arc and in cz to about C4x3_4; there is, in addition, a cloudy post- 
nodal spot of brown extending from the nodus outwards for about 5—6 cells (fore wing) 
and 2 cells (hind wing) between C and M,. Venation denser than in male (e.g. 23 Ax 
in fore and 15 Ax in hinder wings, 14—15 Px in fore and 15—17 Px in hinder pair). 
All ¢ at least with 1 cross-vein and all # of fore wing 3-celled. Anal loop consisting of 
three enclosures, the two proximal ones containing 4—7 cells, the distal (primary) loop 
made up of 11—13 cells with 2—3 central cells. Pterostigma light brown between black 
nervures. Membranula pure white, extending to the end of the two basal cells. 

Abdomen (compressed) long, slender and cylindrical, the basal segments only slightly, 
the terminal ones not at all inflated. Colour uniform brown, only the dorsal parts of the 
intersegmental rings deep black. Pale spots on 2—7 yellowish, quite distinct, arranged 


M. A. LIEFTINCK: Studies in Corduliidae I 51 


as in the male but considerably reduced in size and in the form of tiny transverse streaks 
placed at the jugal sutures on either side of the dorsal crest; 8—10 dark reddish-brown, 
mixed in places with cloudy brown areas. 

Genital valve short, directed almost straight backward; much swollen basally and at 
some distance from base deeply divided to form a pair of slightly less inflated triangular 
processes; median processes of ovipositor arising from the 9th sternite short, black, 
finger-like and subparallel, directed obliquely ventrocaudad, concealed in ventral view 
by the genital valve. Anal appendages long, lanceolate, depressed, unicoloured light 
brown with black pubescence; tuberculum supra-anale subtriangular, not strongly pro- 
tuberant in dorsal view, directed ventrad between the appendages almost at a right angle 
A 55) 

Measurements: & (holotype) abdomen + app. 50.5 mm, hind wing 40.5 mm, pt. 
fore wing 3.2 mm; ¢ paratype 50.3, 41.3, 3.5 mm, respectively; 9 abdomen 44.8— 
47.5 + 4.0 mm, hind wing 40.5—44.0 mm, pt. fore wing 3.5—3.7 mm. 


Synthemis montaguei Campion 
Fig. 36 


1921. Campion, Ann. Mag. Nat. Hist. (9) 8: 55—57, textfig. 9 (& app.), pl. 8 fig. 13 (& 
wings), 6 Mt. Mou, New Caledonia. 


Material. — New Caledonia: 2 & (1 immature) 2 9, N.E.-side, 10 km S. of 
Pouébo, 400 m, 24.1.1964 (&), 15 km S. of Pouébo, 480 m, 22.1.1964 (3), Pouébo, 
200—250 m, 19 & 21.1.1964 (2 9 ), all R. Straatman; g* (adult), Col d’Anuen, 550 
m, 3.111.1960, J. L. Gressitt; 3 (adult), St. Louis Valley, 24.III.1945, H. E. Milliron. 
(All in BISH and ML). 


This is undoubtedly the same insect as the one characterized by Campion, the male only 
being known so far. All of our males are fully coloured but apparently slightly less 
mature than the holotype, with which Mr. Kimmins has been kind enough to compare a 
well preserved specimen from Pouébo. He tells me in a letter that owing, perhaps, to 
inferior optical apparatus, Campion erred a little in his description. The sides of the 
thorax of the type in a good light show obscure pale spots and stripes in the same places 
as in fresh examples, while the markings on the frons, though more obscure, are similar. 
In the type the pale area on the superior appendage is dull yellowish and does not extend 
so far basally; the pterostigma also is blacker, as is the dark ground of the abdomen. 
These are, however, evidently characters varying with the age of the individual, as is 
evident from the present specimens. The genital hamule is of the same shape in all 
individuals, not at all undulated as in S. campioni. 

Female (hitherto undescribed). — Similar to male and differing but slightly, as 
follows. 

Labrum still more broadly bordered with black, the ochreous spots subinterrupted by 
brown in the median line. Frons yellow only on summit of tubercles, which are otherwise 
reddish-brown anteriorly and dark metallic greenish-black posteriorly. Entire thorax dark 
brown with violet and bronze reflections; pale yellow stripe at first lateral suture in one 
specimen as in male, in the second broader and of even width but similarly leaving off 


52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


N 
OA 
fie 


Fig. 35—38. Ventral view of ® terminal abdominal segments of New Caledonian species. 35, Syz- 
themis campioni spec. nov. 36, S. montaguei Campion; 37—38, S. fenella Campion 


ventrad almost 1 mm above the spiracle, which also carries a tiny yellow spot. Membrane 
of wings in adult female subhyaline, distal portion gradually acquiring a pale yellowish- 
brown tint from nodus as far as the apex. Anal area of hind wing posterior to triangle 
with three loop enclosures, the two secondary (basal) ones made up of 4—5 cells, the 
largest (distal) enclosure consisting of 7—9 cells with or without central cell. Pterostig- 
ma brown. Membranula dark grey-brown. 

Transverse yellow lines at extreme bases of abdominal segments 2—5 or 2—6 distinct 
though linear; central spots also narrower and placed more transversely than in male. 
Sexual organs as in Fig. 36; branches of genital valve similar to those of campioni but 
not outcurved. Anal appendages a little longer than segm. 9—10, depressed, lanceolate, 
slightly broader and more swollen than in campioni; colour light yellow, with the bases 
for about 0.5 mm dark brown and the extremities also finely obscured; pubescence black. 

Measurements: 4 abdomen + app. 51.0—53.0 mm, hind wing 42.0—42.6 mm, pt. 
fore wing 3.0 mm; 9 abdomen 45.0—45.5 + 3.8 mm, hind wing 44.0—44.5 mm, pt. 
fore wing 3.0—3.2 mm. 


M. A. LIEFTINCK: Studies in Corduliidae 1 53 


Synthemis fenella Campion 
Fig. 31, 37—38, 44—50 


1921. Campion, Ann. Mag. Nat. Hist. (9) 8: 61—62, textfig. 11 (& app.), pl. 9 fig. 16 (& 
wings), & Mt. Mou, New Caledonia. 


Material. — New Caledonia: 2 & (adult, 1 imperfect) 2 9 (adult), Forêt de 
Thi, 100—300 m, 9.III.1961 (g) and 28—29.III.1961, J. Sedlacek; 2 g 1 2, N.E- 
side, Pouébo, 500 m, 12.1.1964 (g'), 10 and 15 km S. of Pouébo, 400—480 m, 22— 
24.1.1964 (JS 9), R. Straatman (1G 1 © BISH). 


Much smaller than the other four New Caledonian species. Only a single male has been 
described. Our specimens correspond with the existing description so far as it goes; the 
following emendations can now be supplied. 

Coxae of legs blackish anteriorly, yellowish posteriorly; for the rest the legs are black 
except that the inner faces of the fore femora are light green as far as about 0.7 mm 
before the apex. Tibial keels also yellow, extending along distal one-half on fore tibia, 
almost three-fifths on mid tibia, and from near base as far as the apex on hind tibia. 

Wings either entirely colourless or slightly tinged yellowish all over, the proximal area 
to the arculus in one male more deeply so, especially at the bases. Neuration open. All 
triangles and # free (left hind wing 7 with transverse cross-vein in one male); ht with 
a single cross-vein. Ax coincident in c-sc but basal subcostal cross-vein present on all 
wings. Cross-veins in #7, cw and bridge space slightly variable though not appreciably 
differing in number from those of the type as shown in Campion’s photograph. 

Abdomen deep black marked with bright yellow as described for the type. Genital 
organs not very prominent, vesicle of penis shaped as in the other species; hamulus 
shiny black, thick at base, evenly downcurved, and carrying a triangular inferior off-shoot 
at the basal portion, which is very similar to that seen in the allied species. Anal 
appendages deep black, shaped as described and figured by Campion, except that in one 
specimen the appendix superior after the subapical constriction is a little shorter, more 
swollen and distinctly rounded apically, the internal prominence being best visible in 
lateral view. 

Female (adult; hitherto undescribed). — Resembles the male in almost every respect. 
Clear yellow reniform patches on either side on top of frons a little larger and more 
approximated. Yellow band crossing the spiracle at the thoracic sides broader, widest 
dorsally but similarly curving back along upper margin of metepisternum, only little 
narrower than the blue-black area separating it from the metepimeral band. Wing mem- 
brane more strongly flavescent, especially at the bases, which are deeply stained with 
golden yellow about as far as level of triangles. Neuration as described for the male, 
save that in one specimen the internal triangle in one fore wing possesses a single cross- 
vein. Basal portion of hind wing broader and more densely veined: primary (distal) loop 
of large size, comprising 11—12 cells with 1—2 central cells; the much smaller basal 
loop 6-celled. 

Abdomen subcylindrical, basal and terminal segments 8—9 a little inflated. Deep 
black, yellow markings sharply outlined, slightly larger than in male, the transverse basal 
streaks broader and all paired central spots subcircular, surpassing the jugal sutures 
anteriorly. Genital organs black, shaped as in Fig. 37—38. Anal appendages only 1.3 


54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


mm long, subequal in length to segm. 9, straight and cylindrical, gradually tapered, tips 
acuminate. 

Measurements: § abdomen + app. 28.5—29.0 mm, hind wing 25.5—26.5 mm, pt. 
fore wing 1.2—1.3 mm; 9 abdomen (incl. app.) 29.0—31.2, hind wing 28.0—30.0 
mm, pt. fore wing 1.3—1.6 mm. 


Despite the fact that no fresh examples of S. flexicauda have come to our knowledge, 
the five New Caledonian species at present known can be easily held apart by the 
following 


KEY TO THE NEW CALEDONIAN SPECIES OF Synthemis 


1. Large insects, & and 9 abdomen (incl. apps.) measuring 42—55 mm, hind wing 
37—47.5 mm. . . 2 
— Smaller species, 4 and 9 abdomen Ga. ane) 28. 531 mm, a “fine 25. SE 
30 mm. A pair of clear yellow mesepisternal (antehumeral) stripes. present on 
thoracic dorsum. Frons anteriorly with pair of large, curved chrome yellow patches. 
Both sexes with anal appendages black, superior pair of $ not exceeding 2 mm in 
length. Triangles and internal triangles normally without cross-veins and hyper- 
triangle once traversed. Pterostigma small, black, about 1.5 mm long. Terminal seg- 
ments and ovipositor valve of ® as in Fig. 37—38. . . . . . . . fenella 

2. No yellow mesepisternal (antehumeral) stripes present on dorsum of thorax. Wings 
more densely reticulated: triangles and internal triangles traversed by cross-veins, 

at least so in fore wings, and hypertriangles with more than one cross-vein in fore 
wing. Pterostigma much longer and not quite black. Anal appendages in both sexes 
brown, or partly yellow, the superior pair in g' about 3 mm long or more. . 3 

3. Male abdomen with large, erect, pointed black spine on dorsum of segm. 10. Sup. 
anal apps. 4 mm long; in dorsal view broad, almost straight, with acute internal 
black spine at about mid-length, followed first by an emargination, then by a 
dilatation; colour black as far as the emargination, pale yellow beyond. Genital 
hamule in profile view evenly downcurved, with strongly convex outer margin. 
Body dark, almost black, including the legs. Yellow lateral thoracic band present at 
level of spiracle as well as paired basal yellow spots on abdominal tergites 3—4 or 
3—5. Wings at least with the extreme bases spotted with golden yellow, in 9 more 
extensively so and often with brown rays also extending as far as the arculus. Genital 
valve of © short, but outer and inner ovipositor valves, though fused together, still 
discernible by a tie furrow. Female re short, upcurved, mainly 


yellow nr: . . . miranda 
— No robust black sie on dorsum of 3 ab donminal cem 10. Combiacd characters 
not as above . . 4 


4. Male sup. anal apps. M 9 nd at Tease A yellow or POSER Seem 
3—4 of abdomen with pair of yellow spots forming more or less part of a ring at 
extreme base. Sides of thorax at least with traces of a yellow stripe de spot at 
level of first suture and the spiracle. . . 5 

— Male sup. anal apps. and 9 appendages mesta doni Sori as in iis. 33— 
34 and 35. Segm. 2—4 or 2—5 lacking pale-coloured spots at extreme base. Sides 


M. A. LIEFTINCK: Studies in Corduliidae I 55 


of thorax without any trace of yellow colour at first suture and spiracle. Male genital 
hamule Se sinuous Ee, (Fig. 33). Genital valve of © as in fig. 35 

: campioni 

SÌ Mal sua sal os on a mm lone Sel in enc And ehaped similarly to 
those of S. campioni in dorsal view, though slightly dilated internally before the 
middle (not at middle), then emarginate, and dilated just before the rather obtuse 
apex. Appendix inferior about 24 as long as the superior pair. Thoracic sides with 
narrow yellowish line at the first suture eee from a pale spot at the spiracle. 
Genital valve of 9 as in Fig. 36 . . . . . montaguei 
— Male sup. anal apps. over 5 mm long, in Kotal view ent ei in basal half, 
then more sharply convergent, and ending by the tips becoming dilated, parallel, 
and almost in contact with one another; in lateral view depressed and dilated 
ventrally in the middle. Appendix inferior about half as long as superior pair, 
almost straight, abruptly reduced in breadth at some distance from apex, which is 
pointed. Thoracic sides with two broad uninterrupted creamy stripes or bands, one 

at the first suture enclosing spiracle, and a second crossing the metepimeron. Shape 
PROB senital_ valve not known pt. en an une ven, flexicauda 


Discussion and illustrations of the immature stages of some New Caledonian 
Synthemis 


Synthemis miranda Selys (supposition) 
Fig. 32, 39—43 


Material. — ¢ larva (ult), New Caledonia: Bondé region, coastal area, middle 
course of Diahot river, near village Ouénia (chapelle de Saint-Joseph), 70 m, 
16.IX.1965, F. Starmühlner et al, coll. no. FNK 105/1. The specimen is unique and in 
good condition. 

Principal features. 

Measurements: total length 22.5 mm; greatest width of head across eyes 5.7 mm; 
posterior femur 5.2 mm; length of abdomen 13.7 mm, its breadth at apex of segm. 4 
6.5 mm, highest point 4.0 mm. 

Facies of S. macrostigma Selys (Watson, 1962, pl. 9 fig. 49). Body moderately hairy. 

Labium with postmentum reaching backwards to about midway between meso- and 
metacoxae. Prementum broad, gradually widened anterad, deeply concave dorsally; 
median lobe low, not prominent but projecting markedly ventrad, its thickened margin 
minutely and bluntly serrulate, lacking setae; ligula subtriangular, abruptly projecting 
anterad; premental setae clearly differentiated into major and minor sets, ie. 4 + 8— 
7 + 4, the former rather short, the latter minute. Labial palpus strongly, deeply cupped, 
its borders strengthened and transversely wrinkled, outer border with row of rounded 
tubercles bearing longish fine setae; apical border with 4 large crenations, the last (outer- 
most) divided, distal margin of each microscopically serrulate, lacking setae; palpal setae 
4-4, lacking basal spiniform setulae; movable hook short and strong. Frontal plate 
between antennae porrect, transverse, only about one-fourth as broad as greatest width 
of head across eyes and twice broader than long, its borders rounded; surface flat, wholly 
and densely covered with minute swollen papillae, the marginal ones longest and inter- 


56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


mixed with few long setae. Antenna setiferous, 6-segmented, the second segment longest 
(Fig. 43). Head widest across anterior part of eyes, the latter moderately prominent, 
sides behind the eyes only slightly converging, postorbital lobes broadly rounded, covered 
rather densely with microscopical setiferous warts. 

Prothorax transverse, upper pronotal lobes well marked off from the lower parts, their 
surface flat with evenly rounded, slightly raised, almost acute, side margins, which are 
beset with numerous minute setiferous warts. Wing sheaths strongly divergent, reaching 
caudad as far as end of segment 5. Legs relatively short, unmodified and hairy. 

Abdomen elongate-oval, moderately flattened, pubescence especially long and dense at 
side margins, still longer and tufty at posterolateral angles of segments. Middorsal hooks 
conspicuous and in the form of raised bluntly rounded tubercles, those on 2—3 minute 
and knob-like, the next ones progressively higher and longest on 6 and 7, the terminal 
ones directed slightly caudad. Lateral spines wanting, replaced by dense tufts of longish 
hair. Anal pyramid pointed; processes sharply acute, subequal in length, but cerci shortest 
and epiproct a little shorter than the slender paraprocts. 

Integument brown, darkest on head and pronotum; lighter brown are the coxae, 


y) 


40 


Fig. 39—43. Larval structures (ult instar) of Synthemis miranda Selys (supposition) from New 
Caledonia; 39, external view of labium; 40, partial internal view of prementum, showing setae; 41, 
ligula of same, more enlarged; 42, internal view of left labial palpus; 43, antenna 


M. A. LIEFTINCK: Studies in Corduliidae I 57 


trochanters, tibiae towards the end, tarsi, and basal segments of abdomen. Thoracic 
pleurae and terminal segments of abdomen mottled with blackish-brown forming a 
distinct pattern on the abdomen, the dark areas centred with, or surrounded by, light 
brown of the ground colour. 


The present larva exhibits some unusual features by which it differs markedly from 
other species including the two other New Caledonian forms. It is placed in this species 
by supposition only, based on the evidence of the fully developed venation, the body size 
being also in agreement with this assumption. On that basis it might be, however, one 
of the other large-sized species quite as well. It differs from Campion’s larva in many 
respects and is unique in that the penultimate abdominal segments are furnished with 
conspicuous, almost finger-shaped “nodding’’ tubercles middorsally. Campion’s example 
agrees with Australian species like S. eustalacta or leachi by having the two basal joints 
of the 7-segmented antennae short and rather rounded, those of the supposed miranda 
have 6 segments, only the basal one being short and rounded (Fig. 43). Also, Campion’s 
Synthemis is less hairy, has distinct postero-lateral spines at segments 2—9 and no dorsal 
abdominal hooks; lastly, it is said to have more palpal teeth and a greater number of 
submental setae. 1 


Synthemis fenella Campion (supposition) 
Fig. 31, 4449 


Material. — Many larvae, all stages, New Caledonia: 4 larvae (ult) and one 
specimen (probably 4th instar) more closely investigated, dissected and measured. 
Localities: W.-coast, Poya-Nekliai region, right tributary of river Nekliai, 2 km above 
Catholic Mission, 36 m, 10.VIII.1965, coll. no. FNK 63 (1 very young larva); S.W.- 
coast, right tributary of river Dumbéa, 117 m, 15.VII.1965, coll. no. FNK 9 (1 @); 
S.-part, Mont des Sources, near Ouénarou, tributary of river Bleue, primeval forest, 250 
m, 17.VII.1965, coll. no. FNK 15 (1 ex.); same area, streamlet at tributary of river 
Blanche, near forest bridge, 160 m, 21.VII.1965, coll. no. FNK 26 (1 ex.); E-coast, 
Hienghène area, 8 km N., 3 km S. of Ouaiéme Bay, waterfall stream, ca. 130 m, 
7.1X.1965, coll. no. FNK 97 (1 9); all F. Starmühlner et al. 


Principal features of mature larva. 

Resembles the preceding species but can be easily distinguished by much smaller size, 
almost hairless and lighter coloured body, 5-segmented antennae, absence of dorsal 
abdominal protuberances and details of labial structure. 

Measurements: total length 15.0—16.0 mm; greatest width of head across eyes 3-8— 
4.0 mm; posterior femur 3.4—3.5 mm; length of abdomen 9.4—10.0 mm, its breadth 
at apex of segm. 4 4.2—4.6 mm, highest point 4.5 mm. 

Postmentum of labium reaching backwards to the base of metacoxae. Prementum 
distinctly narrower, still more deeply cupped with abruptly downbent midlobe, and more 
heavily sclerotized, than in the former species; asetose margins similar but apical border 
on either side at base of palpus. prolonged to form a broadly tongue-shaped lobe whose 
free margin is entire; premental setae reduced to a pair of inconspicuous short setae 
(occasionally only one!) on either side, the placement of these indicated in Fig. 45 by 
two dots. Labial palpus strongly hollowed out, solid and hard, subtriangular in outline, 


58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


outer border with row of rounded tubercles bearing short spinulose setae; apical border 
entire with strengthened margin and lacking setulae; palpal setae relatively short, 2—2 
only; movable hook long and markedly curved. Frontal plate between antennae projecting 
anterad but distinctly sloping down, longer than in the previous species and about one- 
third (34: 100) as wide as the greatest breadth of head across eyes; surface somewhat 
hollowed out with upturned rounded margins, densely covered with minute swollen 
papillae but lacking setae. Antenna short, almost bare, only 5-segmented, the second 
segment longest (Fig. 49). Prothorax (Fig. 44) shaped much as described for the 
previous species, anterolateral angles of pronotum almost pointed. Integument of all 
thoracic segments and legs covered with minute warts. Wing-sheaths strongly divergent, 
reaching caudad as far as a little beyond apical margin of segment 4 (fore wing) or 
midway between 4 and 5 (hind wing). Legs relatively short, unmodified, lacking long 
setae or hairs but all tibiae with outer row of minute spinulose setae. 

Abdomen elongate-oval, narrower and also higher than in the former species, practi- 
cally hairless. Middorsal hooks wanting, replaced by very low and faintly indicated apical 
tuberculae. Lateral spines likewise absent, though indicated by a vestigial tooth-like 
projection at posterior angle of segment 9, which is devoid of hairs. Anal pyramid 
pointed; processes sharply acute, cerci only half as long as epiproct, which is broadly 
triangular in outline and only little shorter than the paraprocts. 

Integument light brown, darkest on the wing-buds and femora, the latter being 
decidedly lighter apically; the pale yellowish-brown tibiae ringed with dark brown at 
their bases. Abdominal segments 4—9 with a row of rather closely approximated dark 
brown dots placed lengthwise on either side of the dorsal line slightly in advance of the 
middle of segments. 


This is by far the most aberrant Synthemis larva so far known. By its almost naked 
body and elongate, rather flattened forms, it bears a ‘prima facie’ resemblance to certain 
gomphid larvae. Its most outstanding characters are best understood from the outline 
drawings (Fig. 44—49). Head, thoracic segments and divergent wing buds are almost 
exactly similar in shape to the other two regional species. It also resembles the latter in 
the shape of the frontal plate and advanced median lobe of the labium. The strongly 
spoon-shaped labium with its extraordinary hook-like palpus lacking crenations at its 
distal border is, however, surprisingly different from anything so far observed in 
Synthemis. Another peculiarity is the reduced 5-segmented antenna with its single short 
basal joint, present in all individuals examined. Both specimens whose fully developed 
wing venation could be studied in detail, proved to be females. On comparing the neural 
characters with those of the adult dragonfly, I found full agreement in all essential 
respects. There are 12 or 13 cells in the anal loop of the hind wings and the nodal 
indices are those typical of S. fenella. The only discrepancies are (1) all triangles with 
1 cross-vein; (2) discoidal field of both fore and hind wings commencing with 2 cell- 
rows; (3) 2 rows of cells also at the beginning of the field Cz,-Cu, of hind wing. These 
irregularities in the venation may be due either to a certain amount of variation existing 
in the female wings — so much in evidence throughout the Synthemis group — or else, 
may point to some undescribed species closely allied to fenella, the last possibility being 
unlikely considering the great number of larvae collected. This is out of proportion to 
the few adult dragonflies captured and may be due to the diminutive size and dark 
colours of the latter, causing it to be an insect much less conspicuous than its larger 
congeners in the island. 


M. A. LIEFTINCK: Studies in Corduliidae I 59 


Fig. 44—49. Larval structures (ult instar) of Synthemis fenella Campion (supposition) from New 

Caledonia. 44, dorsal view of head and prothorax; 45, external view of labium; 46, left side view 

of same; 47, ligula of submentum, more enlarged; 48, internal view of left labial palpus; 49, 
antenna 


60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 
The early instar larva of S. fenella (Fig. 50) 


The smallest and youngest individual present in Dr. Starmühlner’s collection is 
presumably a 4th instar larva, measuring only 3.1 mm total length. The body is pale 
brownish-yellow, the thorax and legs bearing few fine longish hairs. The frontal plate is 
well developed, flat, but shaped similarly to that of the mature larva. The antennae are 
5-segmented. Fore and mid tarsi consist of a single segment, the hinder pair are 2-seg- 
mented, the basal one being the shortest. The labium is of large size, already markedly 
hollowed out and sclerotized, reaching back to the end of the mesocoxae, shaped and 
armed as in Fig. 50. Wing-buds are still undeveloped and there are no traces of dorsal 
abdominal hooks or lateral spines. 

Judging by the form of the palpus and the complete lack of dorsal abdominal 


Fig. 50. Synthemis fenella Campion (supposition) from New Caledonia. 
Antenna and internal view of labium of 4th instar larva 


prominences, this specimen is almost certainly conspecific, yet differs less from the full- 
grown individuals than might have been expected with such a young creature. As shown 
in the figures, the ligula of the submental midlobe is very similar while crenations at the 
distal border of the labial palpus, though feebly indicated, are non-apparent. One may 
well ask if well-developed teeth exist at all in the earliest instar larva? On the other 
hand, the two premental and two palpal setae are relatively longer than in the mature 
larva, and the antenna — though already 5-segmented — hase two instead of only one 
small basal segments. 

In Oriental larvae of Macromia we have met with a variety of characters developed 
independently in species living under different circumstances, habitat and behaviour 
varying correspondingly from species to species. In our Syzthemis larvae the specific 


M. A. LIEFTINCK: Studies in Corduliidae I 61 


differences are even more striking and less easy to explain. Here we meet with the 
simultaneous development of deeply fissured and straight-bordered labial palpi coupled 
with well developed or non-existent dorsal abdominal hooks and variously segmented 
antennae. It would appear that the firm toothless border of the labial palpus of S. fenella 
with its claw-like movable hook has arisen as a secondary modification of the indented 
form retained by its congeners and generally regarded as a more primitive type. Hence 
we obviously observe combinations of archaic and recently acquired characters resulting 
in reductions as well as specializations allotted differently in species which are never- 
theless closely interrelated. Exaggerated modifications like those of the S. fenella larva 
are presumably purely adaptive, in any case have no phylogenetic counterpart in the adult 
evolution. 

It is my hope before long to investigate more fully the whole collection of Synthemis 
larvae made in 1965 by members of the Oesterreichische Neukaledonien Expedition, the 
same source as the one from which the present notes were derived. 


VII. MATERIAL AND ACKNOWLEDGEMENTS 


Most of the Corduliidae reported upon in the present paper have been obtained in 
recent years by various expeditions, field parties and private collectors throughout the 
Indo-Australian Archipelago. Besides the invaluable help and hospitality I have received 
in 1963 from my friends J. I. Furtado and H. T. Pagden, during our joint field trips in 
Malaya and Penang, many other colleagues have generously supplied me with material 
from different sources and/or given access to specimens of Odonata in various museum 
collections. In the accumulation of material I have enjoyed the kind co-operation of the 
following persons, to whom it is a pleasant duty to acknowledge my gratitude for the 
assistance they have rendered. It goes to J. L. Gressitt and Miss S. Nakata, of the Bishop 
Museum, Honolulu (BISH), for the loan and gift of many specimens particularly from 
the Papuan Region; to K. K. Günther of the Museum f. Naturkunde, Humboldt Uni- 
versitat, Berlin (MNB), and D. E. Kimmins of the British Museum (Nat. Hist.), 
London (BM); to H. Schréder, of the Natur-Museum Senckenberg, Frankfurt a.M. 
(SMF), for the loan of the type of Idionyx philippa; to L. L. Pechuman, of the Dept. of 
Entomology, Cornell University, Ithaca (CUI), for the loan of several of the late J. G. 
Needham types of Odonata from the Philippines; to L. Coomans de Ruiter (Hilversum), 
M. W. F. Tweedie (Rye, Sussex), R. Straatman (Pt. Moresby, Papua), and C. Plateros, 
of the University of San Carlos, Cebu City, P.I., for valuable material of dragonflies and 
their larvae from all over the Archipelago ever since 1933; to J. G. Rozen, of the 
American Museum of Natural History, New York (AMNH), for the privilege of 
studying all Odonata brought home by members of the successive Archbold New Guinea 
expeditions; to A. Heymer, of the Museum National d'Histoire Naturelle, Brunoy, for 
the partial gift of Odonata collected recently in the Lesser Sunda Islands; to Borge 
Petersen, of the Universitetets Zoologiske Museum, Kopenhagen (MC), for specimens 
obtained during 1962 by the Danish Noona Dan expedition in the Bismarcks; and lastly, 
to D. St. Quentin and F. Starmühlner (Vienna), for arranging the loan of an important 
collection of larvae made in 1965 by the Oesterreichische Neukaledonien Expedition. 

Unless indicated otherwise, i.e. by means of the above symbols denoting institutions 


62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 1, 1971 


and museums abroad, the specimens dealt with in this paper are lodged in the Rijks- 
museum van Natuurlijke Historie, Leiden (ML). 

The writer gratefully acknowledges his thanks to the Uyttenboogaart-Eliasen Stichting 
and the Netherlands Organization for Pure Scientific Research (Z.W.O.), for supplying 
a travel grant in connection with entomological investigations carried out by him during 
1963 in the Malay Peninsula. 


VIII. SELECTED REFERENCES 


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1—2. 
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Ceriagrion species. — Jap. Jour. Zool. 16: 99—126, 68 figs., 1 pl. 
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Mag. Nat. Hist. (9) 8: 33—67, pls. 8—9, figs. 12—17. 
Cowley, J., 1942. Descriptions of some genera of fossil Odonata. — Proc. R. ent. Soc. Lond. (B) 
11: 63—78. 
Foerster, F., 1900. Odonaten aus Neu-Guinea. — Termész. Fiizetek 23: 81—108, figs. 
Fraser, F. C., 1918. Two new Indian dragonflies. — J. Bombay Nat. Hist. Soc. 25: 383—385. 
1924. A. survey of the Odonate (Dragonfly) fauna of Western India, etc. — Rec. Ind. 
Mus. 26: 423522, 8 figs., pls. 25—27. 
——, 1934. Indian dragonflies 42. Revision of genus Idionyx. — J. Bombay Nat. Hist. Soc. 37: 
553—572, 3 figs., 2 pls. 
——, 1936a. New Oriental dragonflies. — J. Bombay Nat. Hist. Soc. 38: 700—701, 1 phot. 
———, 1936b. Odonata III iz The Fauna of British India &c. London, 461 pp., 125 figs., 2 pls., 
map. 
——., 1957. A reclassification of the Order Odonata. — Roy. Zool. Soc. N.S. Wales, Sydney. 133 
pp., 62 figs. 
—, 1960. A handbook of the dragonflies of Australasia. With keys for the identification of all 
species. — Roy. Zool. Soc. N.S. Wales, Sydney. 67 pp., 27 pls. 
Furtado, Jose I., 1969. Ecology of Malaysian odonates: Biotope and association of species. — Verh. 
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Laidlaw, F. F., 1912. On a new genus and species of Odonata from Sarawak. — Sar. Mus. Journ. 2: 
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399— 435, 14 figs., 1 pl. 
——., 1935a. New and little known Odonata of the Oriental and Australian regions. — Treubia 
15: 175—207, 15 figs. 
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Nova Guinea 17, Zool.: 203—300, 50 figs. 
——., 1939. Critical notes on the Malaysian species of Idionyx Hagen. — Treubia 17: 199—204, 
2 figs. 
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Treubia 18: 441—608, 2 figs., 19 pls. 
——., 1948. Descriptions and records of South-east Asiatic Odonata. — Treubia 19: 221—278, 
19 figs. 


2 


M. A. LIEFTINCK: Studies in Corduliidae I 63 


Lieftinck, M. A., 1950. Further studies on Southeast Asiatic species of Macromia Rambur, with notes 
on their ecology, habits and life history and with descriptions of larvae and two new 
species. — Treubia 20: 657—716, 61 figs. 

—, 1952. On the Papuasian representatives of the genus Macromia Rambur, with descriptions 
of five new species and some larval forms. — Treubia 21: 437—468, 22 figs. 

— , 1953a. Revision of the Australasian species of Synthemis Selys. With descriptions of four 
new species and a key to their identification. (O.S.R. publ. 38). — Idea (Bogor) 9: 70— 
88, 14 figs. 

———, 1953b. The Odonata of the island Sumba, with a survey of the dragonfly fauna of the 
Lesser Sunda Islands. — Verh. Naturf. Ges. Basel 64: 118—228, map & 72 figs. 

——., 1954. Handlist of Malaysian Odonata; etc. — Treubia 22 Suppl. xiii + 202 pp., map. 

, 1955. Further inquiries into the Old World species of Macromia Rambur. — Zool. Meded. 

Leiden 33: 251—277, 28 figs. 

Martin, R., 1907. Cordulines #2 Cat. Coll. Selys Longchamps 17, 89 pp., 3 col. pls. 

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Hemicordulia assimilis Hagen. — Bull. B. P. Bishop Mus. 98: 111—114, figs. 33—34. 

Needham, J. G. & M. K. Gyger, 1937. The Odonata of the Philippines. — Philipp. J. Sci. 63: 21— 
101, 2 figs., 10 pls. 

Ris, F., 1912. Neue Libellen von Formosa, Südchina, Tonkin und den Philippinen. — Suppl. Ent. 1: 

4484, 19 figs., pls. 3—5. 

, 1913. Die Odonata von Dr. H. A. Lorentz’ Expedition nach Siidwest-Neu-Guinea 1909 und 

einige Odonata von Waigéu. — Nova Guinea 9 Zool. 3: 471-512, 28 figs. 

Selys Longchamps, E. de, 1878. Secondes additions au Synopsis des Cordulines. — Bull. Acad. Belg. 
(2) 45: 183—222. 

St. Quentin, D., 1939. Die systematische Stellung der Unterfamilie der Corduliinae Selys. — Proc. 
VII. Intern. Kongr. Entom. Berlin, I. Sekt. Syst. u. Tiergeogr.: 345—360, 25 figs. 

Starmühlner, F., 1968. Etudes hydrobiologiques en Nouvelle-Caledonie (Mission 1965 du Premier 
Institut de Zoologie de l'Université de Vienne). I. Généralités et description des Stations. 
— Cah. O.R.S.T.O.M., ser. Hydrobiol. 2: 3—27, 18 figs. 

Tillyard, R. J., 1910. Monograph of the genus Synthemis. — Proc. Linn. Soc. N.S. Wales 35: 312— 

377, fig. 1 & pls. 49. 
, 1911. Further notes on some rare Australian Corduliinae, with descriptions of new species. 
— Proc. Linn. Soc. N.S. Wales 36: 366—387, pl. 10. 

Tillyard, R. J. & F. C. Fraser, 1938—1940. A reclassification of the order Odonata. Based on some 
new interpretations of the venation of the dragonfly wing. — Austr. Zool. 9: 125—169, 
27 figs. (1938), 195—221, 11 figs. (1939), 359—396, 15 figs. (1940). 

Watson, J. A. L., 1962. The dragonflies (Odonata) of South-Western Australia, etc. — Handbook 

7, W. Austr. Natur. Club, Perth, 72 pp., 1 col. pl., 21 pls., 3 tab. 
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Synthemidae. — The West Austr. Naturalist 10: 86—91, 9 figs. 


No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.Fl. 45.—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae)). 350 pp. en 98 pl. D.Fl. 70.—. 


No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov 
and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae, Cantantopinae), 77 pp., 1 map, 6 pl. 
D.FI. 25.—. 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. 
DFI. 75.—. 


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No. 1. A. J. Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) 
(The Hydrachnellae of the Netherlands), 199 pp., 333 figs, D.Fi. 30.—. 


En 


A CATALOGUE OF THE TYPE-SPECIMENS OF ODONATA 
PRESERVED IN THE NETHERLANDS 


With a supplementary list of the Odonata types described by Dutch 
scientists deposited in foreign institutional collections 


by 


M. A. LIEFTINCK!) 


Rhenen 
CONTENTS 
I. Introduction . . 65 
II. Annotated list of ee nen in the Anne voor (TORCIA Zeslesıe Co 
Zoölogisch Museum), Amsterdam . . . . TEEN ET HOT 


III. Annotated list of type-specimens in the Rome van Nan ile Tita, Leiden . 70 
IV. Annotated list of type-specimens in foreign institutional collections described by Dutch 


ANDER vo brive At A KEN A A Lal RE COMM AE eed ere he, CALLEN D 
V. References . . be lose Ver me TU EE AND MEN AAE a pet 132 
VI. Addenda to Pa Iv RC el TT em Vener e ES Ree oe ace He 10917, 


I. INTRODUCTION 


Following the example set by D. E. Kimmins, who before his retirement published 
four lists of the type-specimens of Odonata lodged in the collections of the British 
Museum (Natural History)2), the purpose of the present paper is to offer somewhat 
similar lists of Odonata types to be found in other museum collections. The publication 
of type catalogues is advocated by the International Commission on Zoological Nomen- 
clature (Recomm. 72 D4), and consultation of such lists has proved to be both con- 
venient and interesting. 

The present catalogue consists of three parts: in the first two parts are enumerated 
the types of all species-group taxa of Odonata deposited in institutional collections in 
the Netherlands, i.e. the Instituut voor taxonomische Zoölogie (formerly Zoölo- 
gisch Museum), afdeling Entomologie, Zeeburgerdijk 21, Amsterdam (ZMA) 
and the Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden (ML). 
Similarly, the third part comprises a list of primary types of the dragonflies 
presently found in museum collections outside the Netherlands, notably only those 
which have originally been described by Dutch authors. As to this supplementary 
catalogue, it may be considered pretentious that such a selection should have been made. 
After due consideration, however, the preparation of it seemed justified and rewarding 
for the following reasons. Firstly, it was the only way to list all nominal taxa so far 
proposed by the few specialists of Dutch nationality, and by accumulating these the 


1) The author's address: Nieuwe Veenendaalseweg 224, Rhenen (U.), Netherlands. 
?) Bull. Brit. Mus. (Nat. Hist.) Ent. 18 (1966), 22 (1968), 23 (1969) and 24 (1970). 


65 


66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


writer could arrive at some sort of individual completeness not only with respect to his 
own work. In the second place, publication of this list may prove advantageous to future 
students inasmuch as paratypes, syntypes or duplicates of the new taxa listed are, in the 
great majority of cases, also available for study in the collection of the Rijksmuseum 
van Natuurlijke Historie at Leiden. I must confess that the main reason for having 
prepared these catalogues is because many authors, including myself, did not always 
mention or indicate a type-specimen in their earlier works. In point of fact they 
frequently preferred to define a new taxon from the total, or average, of all characters 
observed — provided that good series of specimens were available — rather than 
describe it from a single individual. 

In all categories the names of taxa in the species-group are listed alphabetically. Only 
the names of Holotypes and Lectotypes are printed in clarendon (bold) type, all other 
names in /falics. Perhaps more than in any other insect order, odonatologists are con- 
cerned about the sex of their specimens. In the majority of species, male and female 
are very different morphologically, and it is only too well known that the identification 
of females often meets with considerable difficulty. In view of this, the catalogue 
includes not only the taxa represented by holotypes and lectotypes of either sex but also 
a reference for the first described example(s) of the opposite sex. However, despite 
common usage among entomologists of the term Allotype, to denote the other sex of 
the described taxon, this term has been avoided as depending too much on the individual 
point of view; besides, this subjective qualification is not entitled to have the status of a 
primary type and zpso facto has no validity in nomenclature. The term in question is, 
therefore, replaced by “First described female” (or male), except when the sexes were 
actually found pairing (“in cop.”). Consequently, all described allotypes occurring in 
the literature, whether designated by the author of the nominal species-group, or selected 
subsequently by others, are entered in the lists by the afore-mentioned indication, 
followed by all necessary data concerning its whereabouts. When only the so-called allo- 
type of some earlier described taxon is represented in the collection, then the writer 
has contented himself with mentioning the taxon’s name, its first describer, original 
generic combination and year of publication; full information then follows on the 
opposite sex, as for a primary type. 

In the lists of primary types deposited in the Amsterdam and Leiden museums, the 
first described examples of the opposite sex which are found in foreign institutional 
collections, are incorporated simultaneously and provided with the appropriate depository 
symbols listed on page 120. The names of these “allotypic’’ individuals are not repeated 
in the third list just mentioned. 

Each entry begins with the specific or subspecific name, the author of the genus in 
which it was described, and the reference by year, page and illustrations, if any. The 
status of the type is followed by the data on the labels and those found in the original 
description. In several instances the latter will be found to differ somewhat in arrange- 
ment from the original. For example, the spelling (or paraphrasing) of locality names 
are occasionally slightly altered or amplified in order to facilitate the location of places 
in modern as well as old atlases. On the other hand no attempt has been made to keep 
pace with the exertions of political advisers to change well-established geographical 
names of whole continents or parts of islands, such as Borneo, Celebes and New Guinea, 
into disputable terms like Kalimantan, Sulawesi and Irian, respectively. The changes 
involved are slight and always remain recognizable by comparing the dates of capture 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 67 


with those given in the original or marked on the labels. In some exceptional cases it 
was considered advisable to transcribe the original labels parenthetically in full. The 
label data are followed by the family name, comments concerning the present status 
of the taxon, its generic assignment where differing from the original combination, 
and additional information if considered helpful. 

Needless to say that, wherever in original descriptions unique specimens of either sex 
bear no indication as to their status of type, they are considered Holotypes and referred 
to as such in the corresponding reference. Similarly, in all cases where holotypes have 
been selected and indicated as such in the original or subsequent publications by the 
author of the taxon, they are also referred to in the lists as Holotypes. In those cases 
where in the original description, or in any other subsequent article published by the 
same author, a specimen was marked as “Type” (e.g., “Type in coll. Ris”), this is 
considered equivalent to a designation of Lectotype. No difference has been made in 
this catalogue between lectotypes already designated as such by the author (either 
originally or on a later occasion) and those presently selected by the writer. The new 
lectotype designations perhaps outnumber those recorded previously, because the concept 
of lectotype was not known until comparatively recently. 

Besides types representing taxonomically available species and actually present in the 
collections as of 1971, the valid names of taxa whose types were stated to be found in 
museums but now are lost or destroyed, are included also, but enclosed in square brackets. 

Types of secondary status, such as paratypes (and equivalents) or plesiotypes, etc., 
have not been included in the catalogue’). 

To enliven the dry subject, portraits are added of 23 Zygoptera and 18 Anisoptera 
whose types are found in the collections mentioned in this catalogue. These pictures are 
arranged in Plates 1—7, and for briefness’ sake provided only with the species-group 
name printed beside each picture. Full information on these specimens is to be found 
in the original publications cited. These references are given at the end of the paper 
for all primary types. 

This catalogue was completed July 31, 1971. 


II. ANNOTATED LIST OF TYPE-SPECIMENS IN THE 
INSTITUUT VOOR TAXONOMISCHE ZOÖLOGIE 
(FORMERLY ZOÖLOGISCH MUSEUM), AMSTERDAM 


albardae Selys (Macrogomphus), 1878a: 416—417. Lectotype 4 and first described 
9. Bangka I., labelled by H. Albarda “Macrogomphus Albardae Selys. 4. Banka” 
( 3 ), and [E. Sumatra} “Macrogomphus Albardae Selys. @. Sumatra (Lahat)” (2). 
— Gomphidae. 
Now M. parallelogramma albardae Selys. 
albula Ris (Teinobasis), 1915b: 83, fig. 1. Holotype 4. S.W. New Guinea, Lorentz 
River, 22.1X.1912, G. M. Versteeg, H. A. Lorentz Exped. — Coenagrionidae. 


1) In his “Was versteht man unter einem Typus”, Heinicke (1959, Mitt. Insektenkde 3: 136— 
139) has rather confused the significance of the type concept by the re-introduction of the term 
“Typoid” and its derivatives. This irrelevant quibble has found its admirers among cataloguers on the 
European continent and led to ludicrous terms like “Metatypoide”, ‘“Paratypoide” and other absur- 
dities, which are, of course, not to be taken seriously. 


68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


buruensis Lieftinck (Nannophlebia), 1926: 284—286, fig. 7—8. Lectotype & (very 
immature) and first described 9. Buru I. (S. Moluccas), Fakal, 30.VIII.1921, L. J. 
Toxopeus, labelled by M. A. L. androtype and gynetype, respectively. — Libellulidae. 
Plate 6. 

A second pair, also from Buru, were collected at Mnges’wain, 875 m, 12-14.IV. 
1921, by the same collector, the adult in better condition having been described 
and figured by Lieftinck (1930a: 308—309, fig. 2) (ML). 

ciliata Ris (Nesobasis), 1913: 485—487, fig. 6. Holotype 4 (semiad.). S.W. New 
Guinea, Lorentz River, Bivak Eiland, no date [IX.1909}, H. A. Lorentz Exped. — 
Coenagrionidae. 

First described 9. S. New Guinea, Digul River, Mappi Post, IV.1938, J. M. van 
Ravenswaay Claasen (ML). 

Type-species of Plagulibasis Lieftinck, 1949a: 176—178. 

convergens Lieftinck (Argiolestes), 1949a: 47—48, fig. 13, 24. Lectotype &. N.W. 
New Guinea, Berau Peninsula (Vogelkop), Manokwari, 9.V.1903, Nieuw Guinea 
Exped.; with additional labels: Wahnesia sp. (unknown hand) and Argiolestes 
(Wahnesia) obscura Selys, det. Dr. F. Ris. — Megapodagrionidae. 

dives Ris (subsp. of Tetrathemis cladophila Tillyard), 1913: 505—506. Lectotype &' 
and first described 9 . S.W. New Guinea, Lorentz River, Bivak Eiland, IX.1909, H. A. 
Lorentz Exped. — Libellulidae. 

eurynome Lieftinck (Macromia), 1942: 563—564 (nom. nov.). Lectotype g'. S.W. 
New Guinea, Lorentz River territ., Kloofbivak, ca. 700 m, 17.X1.1912, G. M. Ver- 
steeg, H. A. Lorentz Exped. — Corduliidae. 

New name proposed for misidentified examples of either sex described and figured 
by Ris (1915b: 84—85, fig. 2—3) as M. terpsichore Foerster. The lectotype g' here 
selected was wrongly stated to be the allotype (Lieftinck, 1942: 564), the topotypical 
® having been treated as such more appropriately on a later occasion (Lieftinck, 
1952a: 445). A description and wing photograph of the latter (from Bivak Eiland, 
IX.1909, Lorentz Exped.) were published also by Ris (1913: 494—495, fig. 13, sub 
M. terpsichore Foerster). 

| forcipata Morton (Ischnura), 1907 (&). First described 2 (juv., heterochrom.). 
N. India, W. Himalaya, Sutlej River valley, Sholtu, 2300 m, 4 miles from Kilba, 
1.VII.1926, W. G. N. van der Sleen; destroyed. See Lieftinck, 1927: 91—92. — 
Coenagrionidae. | 

fumosus Hagen (Anax), 1867: 43—44. Lectotype g'. Ternate I. (N. Moluccas), with 
two labels in H. A. Hagen’s writing: Ternate/fumosus Hagen. — Aeshnidae. 

Selected by Lieftinck, 1942: 585, 594— 595, fig. 2 and pl. 40 fig. 147. Specimens 
of this and many other species of Odonata were collected in the island by H. A. 
Bernstein, either in Dec. 1860, April-June 1861, or October 1861 (see Bernstein, 
1865, Ned. Tijdschr. Dierk. 2: 329 q.v.). 

First described 9. Buru I. (S. Moluccas), various locs., V—VI.1921, L. J. Toxopeus 
(ML). See Lieftinck, 1926 : 277—279. 

Now A. fumosus fumosus Hagen. 

hirundo Ris (Lyriothemis), 1913: 509—510. Holotype @. S.W. New Guinea, Lorentz 
River, Bivak Eiland, IX.1909, H. A. Lorentz Exped., with identification label: L. 
hirundo n. sp., det. Dr. F. Ris. — Libellulidae. 

First described 9. N. New Guinea, Humboldt Bay area, Hollandia, I—II.1931, 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 69 


W. Stüber (ML), see Lieftinck, 1933d: 59—62, fig. 34. 

hypsophila Lieftinck (Huonia), 1963c: 775—777, fig. 33—37. Holotype g' and 
first described 9. S.W. New Guinea, Lorentz River basin, Bivak no. 3, 700—750 m, 
16.X1.1909 (g') and 8.XI.1909 (2), H. A. Lorentz Exped. — Libellulidae. 

Described and figured also by Ris (1912b: 743—744, fig. 422), sub H. epinephela 
Foerster. 

lorentzi Ris (Oreagrion), 1913: 480—482, fig. 4a. Holotype ® (in fragments). 
Central W. New Guinea, Snow Mountain (Nassau) Range, Wichmann Mts., 3000 m, 
XI.1909, H. A. Lorentz Exped. Redescribed and figured by Lieftinck, 1949a: 205— 
206, fig. 244, 247. — Coenagrionidae. 

Type-species of Oreagrion Ris (op. cit.). 

macrostylis Ris (Argiolestes), 1913: 475. Holotype g' (immature). S.W. New Guinea, 
Lorentz River, Bivak Island, 14—28.IX.1909, H. A. Lorentz Exped. — Megapoda- 
grionidae. 

melpomene Ris (Macromia), 1913: 496—497, fig. 14—17. Holotype 8 and first 
described 9. S.W. New Guinea, Lorentz River territ., Bivak no. 3 (= Heuvelbivak), 
Went Mts., between Northwest (Beaufort) River and Lorentz River, 700—750 m, 
12.X.1909 (g') and 13.X.1909 (9), H. A. Lorentz Exped. — Corduliidae. 

nigrifrons Ris (Caconeura), 1913: 479. Holotype 8 (terminal abd.-segments missing). 
S.W. New Guinea, Lorentz River, Bivak Eiland, 1.1910, H. A. Lorentz Exped., with 
F. Ris’s identification and type label: Caconeura nigrifrons n. sp. Type. — Protoneuri- 
dae. 

Currently known as Notoneura nigrifrons (Ris). 

nitescens Lieftinck (Teinobasis), 1935c: 253, 262—263, fig. 27. Holotype 8 and 
first described 9. S.W. New Guinea, Lorentz River, Bivak Eiland, 1.1910 (g') and 
Lorentz River, IX.1909 (9), H. A. Lorentz Exped. — Coenagrionidae. 

Nom. nov. pro T. metallica metallica Ris (1913: 487, fig. 8), nec T. metallica 

Foerster. 

paula Ris (Diplacina), 1915b: 114. Lectotype g' (abd.-segments 5—10 missing). 
S.W. New Guinea, Lorentz River, Bivak Eiland, 1909, H. A. Lorentz Exped., Dipla- 
cina fulgens g', det. F. Ris (see Ris, 1913: 508—509, ¢ sub D. fulgens Ris). See 
Lieftinck, 1933d: 44, fig. 26. — Libellulidae. 

First described complete 4 and 9. S. New Guinea, Digul River basin, Mappi 
Post, near Ederat, 11.1939, J. M. van Ravenswaay Claasen (JS) and S.W. New 
Guinea, Sungei Arja (E. tributary of Umar River) W. of Uta, 26.VI.1941, E. 
Lundqvist (® ) (ML). See Lieftinck, 1953d: 159—161, fig. 1. 

Now D. paula paula Ris. 

petaurina Lieftinck (Microtrigonia), 1949a: 8. Holotype 9. S.W. New Guinea, 
Lorentz River territ., Bivak no. 3, 700 m, 6.XI.1909, H. A. Lorentz Exped. — 
Libellulidae. 

Nom. nov. pro M. marsupialis Ris, 1913: 506, and 1919: 1049—1050, fig. 609, 

nec M. marsupialis Foerster, 1903. 

pseudexul Ris (Caconeura), 1913: 478—479, fig. 3. Lectotype 4 (immature) and 
first described 9. S.W. New Guinea, Lorentz River, Bivak Eiland (no date, g') 
and Rivier Eiland (= Bivak Eiland) (no date, © ), H. A. Lorentz Exped. Identified 
on envelopes by F. Ris with: Caconeura spec. n. e, pseudexul, Type 4 and 9. — 
Protoneuridae. 


70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Currently placed as Notoneura pseudexul (Ris). 

reductum Ris (Papuagrion), 1913: 483, 484—485. Holotype 4 (immature), S.W. 
New Guinea, Lorentz River, Bivak Eiland, no date, H. A. Lorentz Exped. Iden- 
tified on envelope by F. Ris with: Stenobasis? spec. C ¢ juv., Papuagrion (n.g.) 
reductum nov. spec. Type, einziges Expl. — Coenagrionidae. 

signiferum Lieftinck (Orthetrum), 1926: 288—290, fig. 12—14. Lectotype 4 and 
first described 9. Buru I. (S. Moluccas), Nal’Besi, 11.V.1921 (4) and 5.VII.1921 
(2), L. J. Toxopeus. — Libellulidae. 

A second pair, also from Buru, were taken by the same collector at Wai Eno, be- 
ginning of 1V.1921, g with pinned slide preparation of penile organ by K. F. 
Buchholz (ML). 

silvarum Ris (Hemicordulia), 1913: 500, 504—505, fig. 24—26. Lectotype d' and 
first described 9. S.W. New Guinea, Lorentz River territ., Bivak no. 3, 700—750 
m, 8.XI.1909 (g') and 12.XI.1909 (2), H. A. Lorentz Exped. — Corduliidae. 
tillyardi Ris (Metaphya) 1913: 497—499, fig. 18—19. Holotype 9. S.W. New 
Guinea, Lorentz River, Bivak Eiland, IX.1909, H. A. Lorentz Exped. — Cor- 
duliidae. è 
First described g', see Anacordulia stueberi Lieft. (ML). 


III. ANNOTATED LIST OF TYPE-SPECIMENS IN THE 
RIJKSMUSEUM VAN NATUURLIJKE HISTORIE, LEIDEN 


abbreviata Lieftinck (Prodasineura), 1951a: 76—80, 83, fig. 1, 2, 5, 6, 10—13. 
Holotype g' and first described 9. S.E. Borneo, Kandangan distr., Ampah, 0—20 
m, IV—V.1948, Liem Swie Liong. — Protoneuridae. 

aciculare Lieftinck (Aciagrion), 1929b: 117—121, fig. 10—11. Holotype g' and 
first described 9. W. Java, Batavia, XI.1907 (g') and X.1907 (9), Edw. Jacob- 
son. — Coenagrionidae. 

acuta Lieftinck (Plattycantha), 1937a: 70—72, fig. 38, 41, 43—45. Lectotype G' 
and first described 9. N. New Guinea, Humboldt Bay area, southern slopes of 
Mt. Cycloop, ca. 1000 m, 7.X.1932 (g') and IV.1935 (9), W. Stüber. — Aesh- 
nidae. Plate 5. 

acuticauda Lieftinck (Ischnura), 1959b: 222—225, fig. 5—6. Holotype g' and first 
described Q. N.E. New Guinea, Upper Chimbu Valley, Mt. Wilhelm, above 
Keglsugl, 2700 m, 4.VIII.1955, J. L. Gressitt. — Coenagrionidae. Plate 2. 

adami Fraser (Libellago), 1939 (3). First described 9. Ceylon, Central Prov., Hara- 
gama, 500 m, Mahaweliganga River, 18.IX.1938 (in cop.), M. A. Lieftinck. See 
Lieftinck, 1940a: 84—88, sub L. miae Lieft. — Chlorocyphidae. 

aderces Lieftinck (Agriocnemis), 1932b: 597, 598—601, fig. 67. Lectotype g'. N. 
New Guinea, Humboldt Bay area, Hollandia, 27.VIII—4.1X.1930, W. Stüber. 
First described 9, same loc., I—II.1931, W. Stüber. — Coenagrionidae. 

adonira Lieftinck (Nannophlebia), 1938: 118—119, fig. 48—49. Holotype g and 
first described 9. N. New Guinea, upper course of Tami River, Bewani Hills, 
300—500 m, Arso, 11.1V.1937, W. Stüber. — Libellulidae. 

adusta Lieftinck (subsp. of Rhinocypha tincta Ramb.), 1949b: 332—334, fig. 1. 
Holotype g'. Solomon Is., Shortland Is., Hisiai River, 22.X.1936, R. A. Lever. — 
Chlorocyphidae. 


EO true 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 71 


aemulus Lieftinck (Onychogomphus), 1937b: 115—119, fig. 31—32. Holotype &. 
S. Sumatra, Lampong distr., Terbanggi-hilir near Menggala, 18.VIII.1936, M. 
Bartels Jr. — Gomphidae. 

aerostiba Lieftinck (Nannophlebia), 1955d: 305—309, 312, fig. 1, 8—9. Holotype 
d and first described 9. Ambon I. (S. Moluccas), Soja di Atas, 400 m, 11.X.1949, 
M. A. Lieftinck. — Libellulidae. 

agalma Lieftinck (Nannophlebia), 1963c: 755—757, fig. 5—7. Holotype & and first 
described 9. Papua, Central Province, 75 km W. of Pt. Moresby, Doa Estate, 
20.X.1962 (7) and 24.IX.1961 ( 9 }, R. Straatman. — Libellulidae. 

aglaia Lieftinck (Nannophlebia), 1948a: 239—243, fig. 5, 8. Holotype g' and first 
described 9. S. Celebes, Bantimurung near Maros (Makassar distr.), 22.V.1941 
(8) and 8.VI.1941 (2), L. Coomans de Ruiter. — Libellulidae. 

aglaia Lieftinck (Selysioneura), 1953b: 646—647, 657, fig. 6, 10. Holotype &. 
Morotai I. (off Halmahera I.), SW-part, 10 m, V.1949, A. J. Kostermans. — 
Isostictidae. Plate 1. 

[albardae Selys (Sieboldius), 1886: CLXXXI—CLXXXII. Holotype &. “Pékin”; 
lost. — Gomphidae. } 

The type was neither recovered in de Selys’ collection (IRSN) nor in the Leiden 

Museum. 

albinensis Belle (Aphylla), 1970: 47—50, fig. 77—80, pl. 8a-b, 9b. Holotype 4 and 
first described 9. Surinam, Albina, 6.1V.1964 (&) and 7.IV.1964 (9), J. Belle. 
— Gomphidae. 

alcedo Lieftinck (Palaiargia), 1949a: 115, 119—123, fig. 153—155. Lectotype gd 
and first described 9. Central N. New Guinea, Sahuweri River basin, Araucaria 
River, 700 m, 22.III.1939 (g'), and above Bernhard Camp, 750 m, 27.111.1939 
(2), L. J. Toxopeus (3rd Archbold Exped.) — Coenagrionidae. 

alecto Lieftinck (Synthemis), 1953a: 72, 74, 84—88, fig. 12—14. Holotype 4 and 
first described 9. Halmahera I. (N. Moluccas), Mt. Sembilan, 600 m, 8.X.1951 
(3) and Mt. Siu, 600—700 m, 28.IX.1951 (9), Sundanese collectors. — Cor- 
duliidae. 

alexia Lieftinck (Nannophlebia), 1933d: 14, 15, 23—24, fig. 14—15. Lectotype & 
and first described 9. N. New Guinea, Humboldt Bay area, Hollandia, 26.VIII. 
1932, W. Stiiber. — Libellulidae. Plate 6. 

alfurus Lieftinck (Argiolestes), 1956b: 66, 72—75, fig. 1—2. Holotype g and 
first described 9. Batjan I. (N. Moluccas), Salawaku River, 50—150 m, 12— 
13.V1.1953, A. M. R. Wegner et al. — Megapodagrionidae. 

alfurus Lieftinck (Lestes, sg. Indolestes), 1960c: 143—147, pl. 2 fig. 2, fig. 11, 
18—20. Holotype & and first described 9. Halmahera I. (N. Moluccas), Mt. 
Sembilan, 600 m, 30.IX.1951, A. M. R. Wegner. — Lestidae. 

alia Calvert (Aphylla), 1948 (3). First described 9. Venezuela, Guanoco, Sucre, 25. 
VIII.1951, J. Racénis. See Belle, 1970: 60—61, fig. 93. — Gomphidae. 

aliena Selys (Urothemis), 1878b: 294—305 (holotype 9, New Guinea). First described 
d'. N. New Guinea, Humboldt Bay area, Hollandia, 0—400 m, XI—XII.1930, W. 
Stiiber (see Lieftinck, 1942: 541—542, pl. 38 fig. 126). — Libellulidae. 

Currently placed as U. signata aliena Selys. 

alternans Lieftinck (Teinobasis), 1935c: 251, 260—262, fig. 26. Lectotype g' and 

first described 9. N. New Guinea, Humboldt Bay area, Tami River plain, Nafri 


72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Road, 14.11.1933, W. Stüber. — Coenagrionidae. 

amabilis Lieftinck (Amphicnemis), 1940b: 363, 366, 368—370, fig. 15, pl. 15 fig. 2. 
Holotype g and first described 9. E. Borneo, N. Kutai, Sangkulirang distr., 
Kariorang, IV.1937, M. E. Walsh. — Coenagrionidae. 

amabilis Lieftinck (Vestalis), 1965b: 337, 349, fig. 4, 5, 9. Holotype g and first 
described 9. N.W. Borneo, Sabah-Sarawak boundary, Brunei Bay area, Dent 
Province, Mt. Marapok, coll. G., 4 labelled “Vestalis amoena g'”’ by R. Martin. 
— Calopterygidae. 

amanda Lieftinck (subsp. of Rhinocypha tincta Ramb.), 1938: 68—69, fig. 1, 3 D-G, 
4 L-M, 5E. Holotype ¢ and first described 9. N. New Guinea, Humboldt Bay 
area, Hollandia, V. 1930, W. Stiiber. — Chlorocyphidae. 

amaryllis Lieftinck (Nannophlebia), 1955d: 316—318, fig. 13—14. Holotype &. 
N. New Guinea, Humboldt Bay area, Bewani River territ., ca. 500 m, Fumb River, 
26.1V.1937, W. Stüber. — Libellulidae. 

amaryllis Lieftinck (Vestalis), 1965b: 338, 345—348, fig. 2, 4, 6, 9a. Holotype dg 
and first described 9. N.W. Borneo, W. Sarawak, slope of Mt. Santubong near 
Kuching, 30.IX.1950, M. A. Lieftinck. — Calopterygidae. 

amethystina Lieftinck (Vestalis), 1965b: 338, 343—345, fig. 3, 6, 9. Holotype gd 
and first described @. S. Sumatra, Lampong distr., S-slope of Mt. Tanggamus, 
Giesting, 400 m, 24.XI1.1934, M. A. Lieftinck. — Calopterygidae. 

amnicola Lieftinck (Vestalis), 1965b: 338, 349—351, fig. 4, 7, 9. Holotype &. N. 
Borneo, Sabah, Mt. Kina Balu, coll. Staudinger, acq. 1903, labelled Vestalis amoe- 
na Selys, by R. Martin. — Calopterygidae. 

amnosia Lieftinck (Nannophlebia), 1955d: 313—315, fig. 10—12. Holotype g and 
first described 9. W. New Guinea, western extremity, Sorong, near Remu, ca. 8 
mi. inland, 24—31.X.1948, M. A. Lieftinck. — Libellulidae. 

amoena Lieftinck (subsp. of Diplacina phoebe Ris), 1953d: 158, 180, fig. 9. Holo- 
type d' and first described ©, Halmahera I. (N. Moluccas), Tuguaer-Tasoa, 
100—150 m, 23.IX.1951, Sundanese collectors. — Libellulidae. 

amphicyllis Lieftinck (Nannophlebia), 1933d: 15, 25—27, fig. 16—17. Lectotype 
d and allotype ® (in cop.). N. New Guinea, Humboldt Bay area, Cycloop Mts., 
southern slopes, 1000 m, 9.IX.1932, W. Stüber. — Libellulidae. 
Central N. New Guinea, Idenburg River basin, above Bernhard Camp, 700—750 
Holotype &. Central N. New Guinea, Sahuweri River basin, Rattan Camp, 1200 
m, 1.111.1939, L. J. Toxopeus (3rd Archbold Exped.). — Megapodagrionidae. 

amphidactylis Lieftinck (Arrhenocnemis), 1949a: 92—93, fig. 99, 111—112, 128. 
Holotype g'. Central N. New Guinea, Sahuweri River basin, Rattan Camp, 1200 
m, 1.111.1939, L. J. Toxopeus (3rd Archbold Exped.). — Megapodagrionidae. 

amphistylus Lieftinck (Argiolestes), 1949a: 41—42, fig. 22, 34, 35. Holotype d'. 
Central N. New Guinea, Idenburg River basin, above Bernhard Camp, 700—750 
m, 29.III.1939, L. J. Toxopeus (3rd Archbold Exped.). — Megapodagrionidae. 

ampycteria Lieftinck (Nannophlebia), 1933d: 15, 22—23, fig. 13. Lectotype 9. 
N. New Guinea, Humboldt Bay area, Hollandia, 17.1V.1931, W. Stüber. First 
described g'. N. New Guinea, Upper course of Korimé River, Nonno (Japu) 
Hills, ca. 15 km S. of Bougainville Range, 400 m, 7—21.III.1936, W. Stüber (see 
Lieftinck, 1942: 455—456, pl. 25 fig. 29—31). — Libellulidae. 

amymone Lieftinck (Macromia), 1952a: 444, 446—449, 461, 465—467, fig. 2, 6, 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 73 


19—21. Holotype 4 and first described 9. W. New Guinea, western extremity, 
Remu, S. E. of Sorong, ca. 50 m, 28.VIII.1948 (3) and 26.X.1948 (9), M. A. 
Lieftinck. — Corduliidae. 

anacharis Lieftinck (Nannophlebia), 1955d: 309—311, 313, fig. 5—7. Holotype 
g and first described 9. Halmahera I. (N. Moluccas), Tolewang, 50 m, 13.X. 
1951 (8) and Mt. Sembilan, 600 m, 8.X.1951 (9), A. M. R. Wegner. — Libel- 
lulidae. 

anacolosa Lieftinck (Vestalis), 1965b: 336, 353—354, fig. 2, 4, 7, 9. Holotype gd. 
N. Borneo, Sabah, E.-slope of Mt. Kinabalu, 10 mi. N. of Ranau, 1570 ft., Paring, 
9.X.1958, T. C. Maa. — Calopterygidae. 

anatya Lieftinck (Nannophlebia), 1933d: 14, 18—19, fig. 9—10. Holotype g'. N. 
New Guinea, Humboldt Bay area, Hollandia, plain country, IV—VI.1931, “Augen 
grin” (W.S.), W. Stüber. — Libellulidae. 

andromeda Selys (Cyclophylla), 1869b (2). First described GJ. Surinam, Sipaliwini 
River, 4.11.1961, D. C. Geijskes. See Belle, 1970: 145—150, fig. 237247, 256 
(8% and larva), pl. 19b, 20a, 21c (sub Negomphoides). — Gomphidae. 

annae Lieftinck (Amphicnemis), 1940b: 365, 367, 370—371, pl. 15 fig. 4, pl. 16 
fig. 4. Holotype 8 and first described 9. W. Borneo, Singkawang, forest marsh 
near Tjapkala, 16.1.1934, L. Coomans de Ruiter. — Coenagrionidae. 

annulata Fraser (Protosticta), 1926: 492. Holotype & (juv., terminal abd.-segments 
lost). N. Celebes, Manado, labelled “Mohari” [= collectors name!}, with F. C. 
Fraser's identification, marked “Type”. — Platystictidae. 

Syn. of Protosticta simplicinervis Selys, 1885. — Syn. nov. 

annulipes Lieftinck (Argiolestes), 1956b: 72, 102—105, fig. 39. Holotype & and 
first described 9 (both immature). Goodenough I. (d'Entrecasteaux Archip.), 900 m 
Camp, 27.X.1953, K. M. Wynn & L. J. Brass (4th Archbold Exped.). — Mega- 
podagrionidae. 

anthaxia Lieftinck (subsp. of Diplacina phoebe Ris), 1933d: 43, 49—52, fig. 29. 
Lectotype ¢ and first described 9. N. New Guinea, Humboldt Bay area, 120 km 
E. of Hollandia, Kressi, 300 m, 1.1932, W. Stüber. — Libellulidae. 

Now N. anthaxia Lieft. 

antiacantha Lieftinck (Nannophlebia), 1963c: 752—755, fig. 1—4. Holotype & and 
first described 9. N.E. New Guinea, Eastern Highlands, Kratke Mts., Lae- 
Goroka Rd., Kassam, 1370 m, 26—29.IX.1959, L. J. Brass (6th Archbold Exped.). 
— Libellulidae. 

antigone Lieftinck (Diplacina), 1933d: 42, 45—47, fig. 27. Lectotype ¢ and 
first described 9. N. New Guinea, Humboldt Bay area, Kressi (up to 120 km 
S.W. of Hollandia), ca. 300 m, 1.1932, W. Stüber. — Libellulidae. 

appendiculatum Lieftinck (Mortonagrion), 1937b: 102—104, fig. 24. Lectotype gd 
and first described ©. Billiton I., Tandjong Pandan, 6—17.1.1937, F. J. Kuiper. 
— Coenagrionidae. 

approximatus Belle (Progomphus), 1966a: 25—28, fig. 46—50. Holotype g' and 
first described 9. Surinam, Mooi Wanna (Weyneweg), 3.1.1964 (&) and 4.1.1964 
(2), J. Belle. — Gomphidae. 

aquila Lieftinck (Tramea), 1942: 526, 530, pl. 33 fig. 95, pl. 34 fig. 101. Holotype 
d' and first described 9. N. New Guinea, southern Bewani Hills, upper course 


74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


of Tami River (5 days’ march S. of Hollandia), Ampas distr., Pauwasi River, 
200—300 m, 7.VI.1939, W. Stüber. — Libellulidae. 
Now placed as Trapezostigma aquila (Lieft.), Hagen’s earlier proposed generic 
name having priority. 
arachne Ris (Planiplax), 1912b (3). First described 9. Surinam, Coropina Creek at 
Republiek, 15.11.1959 (g £), J. Belle. See Geijskes, 1964: 43—45, fig. 21. — 
Libellulidae. 
arachnomima Lieftinck (Macromia), 1953f: 395—406, fig. 4—7. Holotype dg 
(reared from larva: with exuviae). S. Borneo, Sampit distr., ca. 50 m, Pemantan, 
150 km inland, 26.VII.1953, forest brook, M. A. Lieftinck (larva ult); transformed 
at Bogor (Java), 2—3.IX.1953. — First described 9 (reared from larva; with 
exuviae). Same data with holotype; transformed at Bogor (Java), 4.XI.1953. See 
Lieftinck, 1955c: 263—266, fig. 14. — Corduliidae. 
arboricola Lieftinck (Selysioneura), 1959a: 286—288, fig. 4, 9, 17—18. Holotype 
3. Fergusson I. (d'Entrecasteaux Archip.), mountain camp between Agamoia and 
Ailuluai, southcoast, 900 m, 14.VI.1956, L. J. Brass (Sth Archbold Exped.). — 
Isostictidae. Plate 1. 
arborophila Lieftinck (Huonia), 1942: 487—488. Lectotype ¢ and first described 9. 
N. New Guinea, Humboldt Bay area, 120 km E. of Hollandia, Kressi, 300 m, I. 
1932, W. Stüber. — Libellulidae. Plate 6. 
See also Lieftinck, 1935c: 277—279 (partim), fig. 36C, 38—39 (sub H. thalasso- 
phila Foerster). 
Now classified as H. arborophila arborophila Lieft. (1963c: 777—778). 
arcuata Lieftinck (Drepanosticta), 1934d: 469—470, 471, pl. 10 fig. 1, 2, 4. Lecto- 
type d' and first described 9. S.W. Sumatra, Lampong distr., Waitebu near 
Talangpadang, 500 m, 24.VI.1934 ( 4 ) and same area, Wailaläan, 250 m, 22.VI. 
1934 (9), L. J. Toxopeus. — Platystictidae. 
arcuata Lieftinck (Coeliccia), 1940b: 353—355, fig. 7—8. Holotype g and first 
described 9. E. Borneo, N. Kutai, Sangkulirang distr., Batu Besi, V—VI.1934, 
M. E. Walsh. — Platycnemididae. 
arethusa Lieftinck (Nannophlebia), 1948a: 237—239, 243, fig. 5, 7. Holotype & 
and first described 9. S. Moluccas, Ambon I., 20.XI.1921, L. J. Toxopeus (g') and 
Ambon, Sungei Waitumu, 23.1V.1941, Sundanese collector Manis ( 9). — Libel- 
lulidae. Plate 6. 
The holotype was first described and figured by the same author in 1926, sub 
N. lorquini lorquini Selys (see Lieftinck, 1926: 284, fig. 5—6). 
argiocnemis Lieftinck (Teinobasis), 1949a: 165—167, fig. 210—211. Holotype & 
and first described 9. N. Central New Guinea, Idenburg River, Bernhard Camp 
B, 100 m, 11.IV.1939 (g') and Bernhard Camp, 50 m, 7.11.1939 (9), L. J. 
Toxopeus (3rd Archbold Exped.). — Coenagrionidae. 
ariel Lieftinck (Ischnura), 1949a: 223—225, fig. 276, 279—280, 285—286. Holo- 
type g and first described 9. West Central New Guinea, Wissel Lakes group, 
Lake Paniai, 1740 m, 21.VIII.1939, H. Boschma (Le Roux Exped.). — Coena- 
grionidae. Plate 2. 
armeniacum Lieftinck (Oreagrion), 1949a: 211—216, fig. 248, 254—255, 258—259, 
267, 271—272. Holotype 8 and first described 9. Central W. New Guinea, 
Snow Mountain (Nassau) Range, Lake Habbema, 3250 m, 1—4.IX.1938, L. J. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata TO 


Toxopeus (3rd Archbold Exped.). — Coenagrionidae. Plate 3. 

armeniacus Lieftinck (Argiolestes), 1956b: 70, 90—93, fig. 25—27. Holotype g' 
and first described 9. Goodenough I. (d’Entrecasteaux Archip.), 900 m Camp, 
east slopes, 26—27.X.1953, K. M. Wynn & L. J. Brass (4th Archbold Exped.). 
— Megapodagrionidae. 

arses Lieftinck (Palaiargia), 1957: 54—58, fig. 14. Holotype g' and first described 
9. W. New Guinea, Berau Peninsula (Vogelkop), Beraur River area, Klamono 
oilfields (50 km from westcoast inland), 50 m, 22—24.VIII.1948, M. A. Lieftinck. 
— Coenagrionidae. 

arsinoe Lieftinck (Gynacantha), 1948c: 422—424, 426—427, fig. 3. Holotype d. 
Talaud Is. (off N.E. Celebes), Salebabu I., “Salibaboe, coll. Staudinger, acq. 1903”, 
and “Gynacantha basiguttata”, in R. Martin's handwriting. — First described 9: 
with label “Gynacantha n. sp.! Salibaboe” (unknown hand) (NRS), see Lieftinck, 
1948c: 422—427. — Aeshnidae. 

artemis Lieftinck (Procordulia), 1930b: 159—162, 164, fig. 23—25. Lectotype & 
and first described ® (ovipositing). W. Java, Priangan, old crater-lake at Kawah 
Kamodjan near Garut, 1650 m, 19.1V.1930, M. A. Lieftinck. — Corduliidae. 

arthuri Lieftinck (Burmagomphus), 1953e: 251—252, fig. 7. Holotype 9. SE. 
Borneo, Kutai, Balikpapan, Sungai Mentawir, sea-level, 6.X.1950, A. M. R. 
Wegner. — Gomphidae. 

In Lieftinck, 1954: 88, for q read 9. 

arthuri Lieftinck (Gynacantha), 1953c: 177—179, fig. 46. Holotype 4 and allotype 
® (in cop.), W. Sumba I. (Lesser Sunda Is.), Waimangura, Matakori, 436 m, 
21.VIII.1949, A. M. R. Wegner (Sumba Exped.). — Aeshnidae. 

aruana Lieftinck (Huonia), 1942: 491, 492, pl. 26 fig. 38—39. Holotype @. Aru 
Is., Dobo, V.1932, native coll., M. E. Walsh ded. — Libellulidae. 

assidua Lieftinck [subsp. of Cratilla lineata (Brauer)}, 1953c: 202—206, fig. 71. 
Holotype g and first described 9. Central Java, Banjumas, Mt. Slamat, Batur- 
raden, 800 m, 25.1V.1929, F. C. Drescher. — Libellulidae. 

assimilis Selys (Hemicordulia?), 1871: 251—252 (17—18 sep.). Holotype ® (im- 
mature). N. Celebes: Rosenberg Celebes (print), Limbotto (written), 6 Hag. 65 
(written, yellow label), over drawer-label Hemicordulia assimilis Hagen. — 
Corduliidae. 

Three males bear identical locality labels, one adult $ carrying a label Hemicor- 
dulia assimilis Hagen, in de Selys’ handwriting. 

Correctly placed in Hemicordulia. 

astarte Lieftinck (Macromia), 1971b: 42—44, fig. 21, 27-28. Holotype ¢ and first 
described @. Papua, Kokoda Trail, 700 m, 5.X1.1969 (') and Papua, Sogeri, N. 
of Pt. Moresby, 500 m, 3.XI.1969 (Q semiad.), both R. Straatman. — Corduliidae. 

asthenes Lieftinck (Phaenandrogomphus), 1964b: 6—14, fig. 1—13. Holotype d'. 
Malay Peninsula, Selangor, Templer Park, 12—13 mi. from Kuala Lumpur, road- 
side stream, 26.III.1963, M. A. Lieftinck. — Gomphidae. 

First described Q (op. cit.). Malay Peninsula, “Wasserscheide zwischen Perak u. 
Pahang, Inner Malakka, Camp Jor, Albert Grubauer 1901”, ex coll. F. Förster 
(UMMZ). 

Type-species of Phaenandrogomphus Lieftinck (op. cit.). 

astridae Lieftinck (Procordulia), 1935a: 189—190. Lectotype ® (semiadult) and 


76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


first described g (immature, in fragments). N. New Guinea, Humboldt Bay area, 
Cycloop Mts., 1000 m, 19.V.1932 (2) and 141V.1935 (8), W. Stüber. — 
Corduliidae. 

Although this insect, upon the discovery of a topotypical g', was recognized as a 
species distinct from P. sylvia Lieft., I have selected the 9 as lectotype (mentioned 
as “type” in my later account, 1942: 559), the & being in poor condition. The 
terminalia of fully adult topotypical g and 9 were figured in the paper last 
mentioned (1942, pl. 37 fig. 120—122). 

atlanticus Belle (Negomphoides), 1970: 133—136, fig. 218—222, pl. 20b. Holotype 
d'. Surinam, Coppename river, Raleigh Falls, forest 1 km from river, 21.VIII. 
1957, D. C. Geijskes. — Gomphidae. 

atomarius Lieftinck (Podolestes), 1950c: 40—44, fig. 1—2. Holotype & and first 
described Q. S.E. Borneo, Kandangan distr., Ampah, 0—20 m, IV—V.1948, 
Liem Swie Liong. — Megapodagrionidae. 

atrovirens Lieftinck (Macromidia), 1935a: 194—196. Holotype 9. S. Sumatra, Ben- 
kulen distr., Bukit Itam, 6—700 m, 11—15.VI.1935, M. E. Walsh. — Corduliidae. 

attala Lieftinck (Drepanosticta), 1934d: 472—474, fig. 2—3. Holotype g and first 
described 9. W. Borneo, Singkawang-Bengkajang Rd., forest brook near Serukan, 
hill country, 16.1V.1934 (g') and 28.1.1934 (9), L. Coomans de Ruiter. — 
Platystictidae. 

audax Lieftinck (subsp. of Gomphidia abbotti Williamson), 1948a: 263—264, 266, 
fig. 13—15. Lectotype 4. S. Sumatra, Lampong distr., Terbanggi-hilir near Meng- 
gala, 19.VIII.1936, M. Bartels Jr. — Gomphidae. 

aulicus Lieftinck (Argiolestes), 1949a: 43—44, fig. 23. Holotype 9. Central W. 
New Guinea, Bernhard Camp B, 150 m, “waterfall”, 13.1V.1939, J. Olthof (3rd 
Archbold Exped.). — Megapodagrionidae. 

aurantiaca Lieftinck (Notoneura), 1938: 94—95, fig. 29— 30. Lectotype ¢ and first 
described 9. W. New Guinea, Onin Peninsula (western extremity), Fak Fak, 
28—29.XII.1912, H. Elgner. Both misidentified with Caconeura plagiata: Ris, 
1915b: 90—91, fig. 10; nec Alloneura plagiata Selys, 1886 (3 Misool I.). — Pro- 
toneuridae. 

aurea Lieftinck (Teinobasis), 1932b: 585—587, fig. 63. Lectotype d' and first 
described 9. N. New Guinea, Humboldt Bay area, Pim River, near Hollandia, 
200 m, 30.V.1931, W. Stüber. — Coenagrionidae. 

aurea Lieftinck (Pachycypha), 1950a: 631—638, fig. 1—3. Holotype g and first 
described @. S.E. Borneo, Kandangan distr., 7 km N.E. of Ampah, near Rana- 
mun, 19.V.1948 (g') and 22.V.1948 (2), Liem Swie Liong. — Chlorocyphidae. 

Type-species of Pachycypha Lieftinck (op. cit.). 

aureofrons Lieftinck (Cyanocnemis), 1949a: 107—112, fig. 104, 114, 117—119, 
125—127. Holotype g and first described 9. Central W. New Guinea, Sahuweri 
River valley, Araucaria Camp, 800 m, 17.111.1939, L. J. Toxopeus (3rd Archbold 
Exped.). — Platycnemididae. 

Type-species of Cyanocnemis Lieftinck (op. cit.). 

auriculatum Lieftinck (Papuagrion), 1937a: 27, 39—41, fig. 27—28. Holotype ¢ 
and first described 9. N. New Guinea, Humboldt Bay area, southern slope of 
Mt. Cycloop, 1000 m, 2.IX.1933 (4) and 20.IX.1934 (9), W. Stüber. — Coen- 
agrionidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata zu 


auripennis Geijskes (Aeschnosoma), 1970: 27—28, fig. 26—27, pl. 2a. Holotype 9. 
Surinam, upper Coropina creek near Zanderij, 6.11.1946, D. C. Geijskes. — Cor- 
duliidae. 

austeni Lieftinck (Drepanosticta), 1940a: 89—91, fig. 2. Lectotype @ and first 
described g'. Waigeu I., off West New Guinea, labelled “Waigeoe, Bernstein”). 
— Platystictidae. 

australis Guérin-Méneville (Agron), 1832: 196 (holotype 9 Offak, Waigeu I.). First 
described g'. Waigeu I., off West New Guinea, labelled ‘‘Waigeoe, Bernstein”’1). 
One of two males redescribed, figured and selected as allotype by Lieftinck, 1938: 
70—73, fig. 6, 7a, 8. — Megapodagrionidae. 

Type-species of Argiolestes Selys, 1862. 

australis Hagen (Epophthalmia), 1867: 61—62. Holotype &. “Celebes [N. Celebes, 
Minahasa}, Limbotto, Rosenberg” (print) and “7 Hag. 65” (written, orange 
label). Designated by Lieftinck, 1931: 64. — Corduliidae. 

austrosundanus Lieftinck (subsp. of Burmagomphus williamsoni Foerster), 1964b: 
25—26. Holotype g and first described 9. E. Sumba I. (Lesser Sunda Is.), 
Laluku, 4.VII.1949 (G') and Wai Lekabe, 28.VI.1949 (2), A. M. R. Wegner et 
al. (Sumba Exped.). — Gomphidae. 

axiagasta Lieftinck (Nannophlebia), 1933d: 14, 15, 16—17, fig. 7—8. Lectotype { 
and first described 9. N. New Guinea, Humboldt Bay area, Hollandia, III.1931 
(8) and IV—VI.1931 (2), W. Stüber. — Libellulidae. 


balteatum Lieftinck (Orthetrum), 1933d: 63—66, fig. 35. Lectotype g' and first 
described 9. N. New Guinea, Humboldt Bay area, Hollandia, plain country, 
III.1931, W. Stüber. — Libellulidae. 

banteng Lieftinck (Onychogomphus), 1929b: 133—136, fig. 25—27. Holotype d'. 
W. Java, Djampangs, Pandan Arum [Estate on slope of Mt. Salak], 1000 m, 
VI.1916, native coll., ex coll. W. Roepke. — Gomphidae. 

barbatula Lieftinck (Drepanosticta), 1940b: 351—353, fig. 6. Holotype &. E. Bor- 
neo, N. Kutai, Sangkulirang distr., Batu Besi, VI.1937, M. E. Walsh. — Platy- 
stictidae. 

bartelsi Lieftinck (Drepanosticta), 1937b: 70—72, fig. 8. Holotype g' and first 
described 9. S.W. Java, Tjidamar near Tjitoé, ca. 40 m, 13.IX.1935, M. Bartels 
Jr. — Platystictidae. 

bartelsi Lieftinck (Heliaeschna), 1940b: 386—390. Holotype ¢ and first described 
2. W. Borneo, Singkawang distr., near Bakuan, 9—10.1V.1934, L. Coomans de 
Ruiter. — Aeshnidae. 

basilanensis Laidlaw (subsp. of Devadatta podolestoides Laidlaw), 1934: 102—103. 
Lectotype & and first described 9. Philippine Is., Basilan I., Maloong, XII.1932, 
K. Kuwasima. — Amphipterygidae. 

basitincta Lieftinck [subsp. of Amphiaeschna ampla (Rambur) }, 1940b: 385—386. 
Holotype ¢ and first described 9. S.W. Sumatra, Benkulen distr., Tandjong Sakti 
near Pagaralam, 600 m, 27.V.1935 (g') and Muara Tenam, 250 m, 16—23.VI. 
1935 (9), M. E. Walsh. — Aeshnidae. 

batjanum Asahina (Ceriagrion), 1967: 267—269, 321, fig. 32—37. Holotype gd 


1) H. A. Bernstein collected in the island of Waigeu from about mid March till the 13th May, 1863. 


78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


and first described 9. Batjan I. (N. Moluccas), Wajaua, Birabiraketjil, 19.VI. 
1953, A. M. R. Wegner et al. — Coenagrionidae. 

beatricis Lieftinck [subsp. of Rhyothemis phyllis (Sulzer) |, 1942: 506, 509—510, pl. 
28 fig. 50—53, pl. 29 fig. 54-56. Holotype g and first described 9. N. New 
Guinea, Humboldt Bay area, Hollandia, 27.VIII—4.1X.1930, W. Stüber. — Libel- 
lulidae. 

beatrix Lieftinck (Notoneura), 1949a: 63—64, fig. 67—70, 85—86. Holotype ¢ 
and first described 9. Central N. New Guinea, Idenburg River valley, Bernhard 
Camp, 50 m, 31.VII.1938 (G') and 26.IX.1938 (£), J. Olthof (3rd Archbold 
Exped.). — Protoneuridae. 

bellicosa Lieftinck [subsp. of Epophthalmia vittigera (Ramb.)}, 1948e: 19—21. 
Holotype g'. Burma, Thaton distr., Mokpalin, 16.V.1923, J. Elton Bott, ex coll. 
T. B. Fletcher. — First described 9. S. Burma, Tenasserim, Mergui, 1922, J. Elton 
Bott, ex coll. T. B. Fletcher. — Corduliidae. 

billitonis Lieftinck (Amphicnemis), 1940b: 365, 367, 374, pl. 15 fig. 4, pl. 16 fig. 1. 
Holotype g and first described @. Billiton I. (west), Tjerutjuk, 24.X11.1935, 
F. J. Kuiper. — Coenagrionidae. 

biroi Foerster (Tetrathemis), 1900 (g', not 9). First described Q (in cop.). N. New 
Guinea, Humboldt Bay area, Cycloop Mts., 400 m, I—II.1931, W. Stüber. See Lief- 
tinck, 1933d: 19—21, fig. 11. — Libellulidae. 

Now Nannophlebia biroi (Foerster). 

borneensis Lieftinck (Heliogomphus), 1964a: 90—93, fig. 3—5. Holotype g. E. 
Borneo, Kutai, Nunukan, 1.1953, forest stream, L. Hentig. — First described 9. 
E. Borneo, Kutai, Samarinda distr., ca. 100 km upstream Tabang River, kali Ben- 
gen, ca. 100 m, 17.IX.1956, A. M. R. Wegner. — Gomphidae. 

brachycnemis Needham (Progomphus), 1944 (2). First described g'. Surinam, Upper 
Para River, 25.111.1962, J. Belle. See Belle, 1966a: 2—7 (dg 9 and larva), fig. 
1—12, pl. 1. — Gomphidae. 

brasiliensis Belle (Aphylla), 1970: 51—52, fig. 81—82. Holotype ¢ and allotype 9 
(in cop.). Brazil, Mato Grosso, Barre do Tapirapé, forest edge, 11.1.1963, B. 
Malkin. — Gomphidae. 

buettikoferi Ris (Eleuthemis), 1910: 382—384, fig. 231—232. Lectotype G'. Liberia, 
labelled “Bavia v/d Prul [?} Liberia feb. 1880”, under drawer-label büttikoferi 
Ris. — Libellulidae. 

Type-species of Eleuthemis Ris (op. cit). 

buwaldai Lieftinck (Podolestes), 1940b: 347— 348, fig. 4b. Holotype &. E. Sumatra, 
Riouw distr, Rengat, Pangkalan kasai, 2.1V.1939, P. Buwalda. — Megapod- 
agrionidae. 

buwaldai Lieftinck (Teinobasis), 1949a: 171—172, fig. 214— 216. Lectotype g' and 
first described 9. Aru Is, Wokam I., Dosi Wamalau, 18.V.1938, P. Buwalda. — 
Coenagrionidae. 


calamineum Lieftinck (Ceriagrion), 1951c: 185—186, 189—193, fig. 2. Holotype 
d and first described 9. W. Java, Bogor, Botanic Garden, 250 m, 23.VIII.1930, 
M. A. Lieftinck. — Coenagrionidae. 
See Asahina (1967: 272—274, figs.). 
callisphaena Lieftinck (Notoneura), 1937a: 16—18, fig. 10—11. Lectotype g' and 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 79 


first described 9. N. New Guinea, Humboldt Bay area, Nonno Hills, 400 m, 
15—20.X1.1935, W. Stiiber. — Protoneuridae. 

callisto Laidlaw (Macromia), 1922 (3). First described 9. Malay Peninsula, Selangor, 
1114 mi. Kuala Lumpur-Bentong Rd., Sungai Gombak, 12.VI.1963, J. I. Furtado. 
See Lieftinck (1971b: 23—26). — Corduliidae. 

calosomum Lieftinck (Pseudagrion), 1936a: 124—126, fig. 8, 9, 11. Lectotype d. 
Sumba I. (Lesser Sunda Is.), E. Sumba, Kananggar, 700 m, V.1925, K. W. Dammer- 
man. — First described 9. E. Sumba, Baing, 100—200 m, 21.VI.—3.VII.1949 
(Sumba Exped. 1949, NMB). 

calverti Geijskes (Misagria), 1951: 71—74, 76, fig. A—F. Holotype g' and first 
described 9. Surinam, Brownsberg, 14.1X.1938 (g') and 19.IX.1938 (2), D. C. 
Geijskes. — Libellulidae. 

campioni Lieftinck (Synthemis), 1971b: 48—51, fig. 33—35. Holotype ¢ and first 
described 9. New Caledonia, N.E.-side, Pouébo, 200 m, 24.1.1964 (g') and same 
loc., 250 m, near mountain stream, 23.1.1964 ( © ), both R. Straatman. — Corduliidae. 

capillaris Lieftinck (Tanymecosticta), 1959a: 294—295, fig. 11, 29—32. Holotype 3. 
Tanimbar Is. (Timorlaut), Jamdena I., IV.1938, P. Buwalda. — Isostictidae. 

capreola Lieftinck (Selysioneura), 1932b: 542, 545—547, fig. 40—41. Lectotype ¢ 
and first described 9. N. New Guinea, Humboldt Bay area, Hollandia, I—11.1931, 
W. Stüber. — Isostictidae. 

carnifex Lieftinck (Palaiargia), 1932b: 559, 563—565, fig. 53. Lectotype g and 
first described 9. N. New Guinea, Humboldt Bay area, Hollandia, 27.VIII—4.IX. 
1930. W. Stüber. — Coenagrionidae. 

casuarina Lieftinck (Podopteryx), 1949a: 39— 41, fig. 21, 33. Holotype 4. Central 
W. New Guinea, Idenburg River basin, above Bernhard Camp, ca. 400 m, “in heavy 
jungle far away from water”, 11.11.1939, L. J. Toxopeus (3rd Archbold Exped.). — 
Megapodagrionidae. 

celaeno Lieftinck (Macromia), 1955c: 274—277, fig. 27-28. Holotype ¢ and first 
described 9. N.E. New Guinea, Papua, Kwagira River, Peria Creek, 50 m, 3 and 
2.1X.1953, respectively, G. M. Tate (4th Archbold Exped.). — Corduliidae. 

celebense Lieftinck (subsp. of Anax fumosus Hagen), 1942: 581, 586, 595—597, pl. 
40 fig. 144, 149, pl. 41 fig. 152. Holotype ¢ and first described @. Central Celebes, 
Palu distr., S. Kulawi, Kalamanta, 1000 m, 10.X.1940 (G') and Kulawi, 550 m, 
15.1.1941 (9), P. M. Felix. — Aeshnidae. 

celebense Lieftinck (Pseudagrion), 1937b: 90—92, fig. 18c, 19. Holotype g' and 
first described 9. N.W. Celebes, Palu, 1.1937, R. Awibowo (g') and Centr. E. 
Celebes, Kali Tominanga, 22.VIII.1932, Prof. Dr. R. Woltereck (9). — Coen- 
agrionidae. 

cerinomelas Lieftinck (Ceriagrion), 1927: 88—91, fig. 4. Holotype g' and allotype 
® (in cop.). N. India, W. Himalaya, Sutlej valley, Sholtu, 2300 m, 4 miles from 
Kilba, 1.VII.1926, W. G. N. van der Sleen. — Coenagrionidae. 

Currently placed as a subspecies of C. fallax Ris (see Asahina, 1967: 284—286, 

321, fig. 100—106). 

cervula Lieftinck (Synthemis), 1938: 119—122, fig. 50. Holotype g' and first 
described 9. N. New Guinea, Bewani Hills, upper course of Tami River, 300—500 
m, Josko, 22—30.X1.1936. W. Stüber. — Corduliidae. 

ceyx Lieftinck (Palaiargia), 1949a: 115, 123—125, fig. 135, 156—157. Holotype g' 


80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


and first described 9. Central N. New Guinea, Idenburg River valley, above Bern- 
hard Camp, 700 m, 27—29.111.1939, L. J. Toxopeus (3rd Archbold Exped.). — 
Coenagrionidae. 
Now P. ceyx ceyx Lieft. (1949a). 
chalciope Lieftinck (Macromia), 1952a: 439, 456—460, fig. 5, 10. Lectotype g' and 
first described 9. Halmahera I. (N. Moluccas), Mt. Sembilan, 600 m, 27.IX.-6.X. 
1951, Sundanese collectors. — Corduliidae. 
chalcochiton Ris [ subsp. of Agrionoptera insignis (Rambur)], 1915a: 15—16. Lecto- 
type d and first described Q (androchrom.). Simalur I. (off N.W. Sumatra), Pulu 
Babi, IV.1913, E. Jacobson, with Ris’s identification labels. — Libellulidae. 
chalcosoma Lieftinck (Nannophyopsis), 1935a: 183—188, fig. 4—6. Holotype ¢ 
and first described ©. Billiton I. (east), Gantung, 17.IX.1935 (g') and 13.X.1935 
(2), F. J. Kuiper. — Libellulidae. Plate 7. 
Type-species of Nannophyopsis Lieftinck (op. cit.). 
chalybeostoma Lieftinck (Notoneura), 1932b: 521, 525, 528—529, fig. 20—22. 
Lectotype 4 and allotype ® (in cop.). N. New Guinea, Humboldt Bay area, Hol- 
landia, I—II.1931, W. Stüber. — Protoneuridae. 
charmosyna Lieftinck (Palaiargia), 1932b: 559, 560—563, fig. 52. Lectotype g' and 
first described 9. N. New Guinea, Humboldt Bay area, Hollandia, 400 m, IX.1930, 
W. Stiiber. — Coenagrionidae. 
Now P. charmosyna charmosyna Lieft. (1949a). 
chloropleura Lieftinck (Idiocnemis), 1932b: 502—503, fig. 6. Lectotype g' and first 
described 9. N. New Guinea, Humboldt Bay area, Hollandia, IV—VI.1931, W. 
Stüber. Selected by Lieftinck, 1958: 267. — Platycnemididae. 
civicum Lieftinck (Pseudagrion), 1932b: 568, 570—572, fig. 55. Lectotype g' and 
allotype 9 (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, 27.VIII— 
4.1X.1930, W. Stüber. — Coenagrionidae. 
claaseni Lieftinck (Drepanosticta), 1938: 80—81, 87, fig. 17, 19a. Holotype g'. N.W. 
New Guinea, Berau Peninsula (Vogelkop), Jenau near Aja Maru, 350 m, XII.1937, 
J. M. van Ravenswaay Claasen. — Platystictidae. 
clavata Lieftinck (Drepanosticta), 1932b: 515—517, fig. 18. Lectotype g' and first 
described 9. N. New Guinea, Humboldt Bay area, Hollandia, plain forest, 400 m, 
IX.1930, W. Stüber. — Platystictidae. 
clymene Lieftinck (Diplacina), 1963c: 767—768, fig. 20— 23. Holotype g'. Fergusson 
I. (d'Entrecasteaux Archip.), Agamoia, 200 m, 20.VI.1956, L. J. Brass (Sth Arch- 
bold Exped.). — Libellulidae. 
coarctatum Lieftinck (Pseudagrion), 1932b: 567, 578—579, fig. 57, 60. Holotype 
d'. N. New Guinea, Humboldt Bay area, Hollandia, IX—XII.1930, W. Stüber. 
First recorded 9. Same area, Lake Sentani, 2—17.X11.1932 (in cop.), same collector. 
— Coenagrionidae. 
coccinea Lieftinck (Onychothemis), 1953e: 264—266. Holotype 8 and first described 
Q.S.E. Borneo, Kutai, Balikpapan, Sungai Mentawir, sea-level, 23.X.1950, A. M. R. 
Wegner. — Libellulidae. 
conchinus Williamson (Cyanogomphus), 1916 (8). — First described 9. Surinam, 
Upper Para River, 1.1V.1962, J. Belle. See Belle, 1966b: 52—56 (¢ 9), fig. 83—90, 
pl. 8 (sub Ebegomphus). — Gomphidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 81 


Later, Belle (1970) placed Ebegomphus Needham in the synonymy of Cyanogom- 
phus Selys. 
conjectus Belle (Progomphus), 1966a: 17—20, fig. 32, 44, pl. 4. Holotype ©. Suri- 
nam, Coppename River, 10.X.1901, H. A. Boon. — Gomphidae. 
connectens Lieftinck (Argiolestes), 1956b: 69, 79—81, fig. 8—9, 12—13. Lectotype 
d' and first described 9. W. New Guinea, western extremity, Sorong, ca. 100 m, 
24.X.1948 (¢) and 30.X.1948 (9), M. A. Lieftinck. — Megapodagrionidae. 
coomansi Laidlaw (Leptogomphus), 1936: 267—269, fig. 1. Holotype 8 and first 
described 9. W. Borneo, Mampawah near Singkawang, forest between Pakmiong- 
theo and Pandjaoa, 3.1V.1932 (g') and Sungei Bagak (G. Raja-complex), 10.VI. 
1933 (2), L. Coomans de Ruiter. — Gomphidae. 
coomansi Lieftinck (Pseudagrion), 1937b: 86—90, fig. 17—18. Holotype g and 
first described 9. W. Borneo, Singkawang, Montrado, 22.X.1931 (g') and Paten- 
gahan Road, 18.11.1932 (2), L. Coomans de Ruiter. — Coenagrionidae. 
coomansi Lieftinck (Coeliccia), 1940b: 355—356, fig. 9—10. Holotype g and first 
described 9. W. Borneo, Singkawang, Mt. Poteng, 400 m, 31.1.1932, L. Coomans de 
Ruiter. — Platycnemididae. 
coomansi Lieftinck (Elattoneura), 1937b: 79—80, fig. 13. Lectotype g and first 
described 9. W. Borneo, Singkawang area, Mt. Ambawang, 13.11.1931 (4), and 
Singkawang, forest marsh near Bakuan (Selakau River), 20.VII.1932 (9), L. Coo- 
mans de Ruiter. — Protoneuridae. 
coomansi Lieftinck (Podolestes), 1940b: 348—350, fig. 4c, 5. Holotype g and first 
described ©. S.E. Sumatra, Palembang, 25.1X.1939, L. Coomans de Ruiter. — Mega- 
podagrionidae. 
cornelia Selys (Calopteryx), 1853 : 15. Holotype 4 (adult). With printed label ’’v. 
Siebold, Japan”, under old drawer-label “Cornelia”. First described Q (ad. legs 
missing), labelled “Burger, Japan” (white disk, unknown handwriting), and a second 
® (ad., intact), with rectangular label “Japonia Agrion?” (as before). — Calo- 
pterygidae. 
The 9 was first recorded in the Monographie (1854 : 48), as follows: “9. 
Je crois me rappeler que la femelle que j'ai vu à Leyde, porte un faux ptérostigma 
blanc”, an observation evidently made after studying the above specimens. The first 
full description is found in Selys (1869a: 648). 
Type-species of Anaciagrion Kennedy, 1920. 
cornelia Lieftinck (Selysioneura), 1953b: 647—651, 656, 658—668, fig. 2, 7, 11, 13, 
15—26. Holotype g' and first described 9. W. New Guinea, western extremity, 
Sorong, 50 m, 30.X.1948, M. A. Lieftinck. — Isostictidae. Plate 1. 
cornelii Lieftinck [subsp. of Rhinocypha fenestrata (Burm.) }, 1947: 218—220, pl. 2 
fig. 1. Holotype g'. S.W. Bali I. (Lesser Sunda Is.), Pulukan forest-reserve, ca. 200 
m, 3.1V.1936, C. G. G. J. van Steenis. — First described ©. Central S. Bali I., Mt. 
Batukau, ca. 700 m, Wonggaja-gede, 13.11.1940, Walter Spies. — Chlorocyphidae. 
corniculatum Lieftinck (subsp. of Papuagrion pesechem Lieft.), 1949a: 152—153, 
fig. 194, 200. Lectotype Z and first described 9. Central N. New Guinea, Sahuweri 
River valley, Rattan Camp, 1150 m, 12.11.1939, L. J. Toxopeus (3rd Archbold 
Exped.). — Coenagrionidae. 
cornuta Foerster (Karschia), 1900 ( 9 ). First described &. N. New Guinea, Humboldt 
Bay area, S. Cycloop Mts., 700—1000 m, IX.1932, 1933 & 1934; II, IV & IX.1935, 


82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


W. Stüber. See Lieftinck, 1937a: 66—70, fig. 44. — Aeshnidae. 
Type-species of Karschia Foerster, 1900 (nom. praeocc.). 
Now Plattycantha cornuta Foerster, 1908, isogenotypic. 
coropinae Geijskes (Micrathyria), 1963: 78—81, fig. 17. Holotype g and first 
described 9. Surinam, Coropina Creek, near Vier Kinderen, 26.1.1947, D. C. Geijs- 
kes. — Libellulidae. 
corruptum Lieftinck (Papuagrion), 1938: 105—107, fig. 40. Lectotype g' and first 
described 9. N. New Guinea, Bewani Hills, upper course of Tami River, 5 days’ 
journey S.E. of Hollandia, Sawia, 200 m, 10.X.1936, W. Stüber. — Coenagrionidae. 
corvina Lieftinck (Caconeura), 1930b: 138—141, 151, fig. 1—4. Holotype g and 
allotype Q (in cop.). Java mer., res. Banjumas, Djeruklegi, 7.1.1929, F. C. Drescher. 
— Protoneuridae. 
Syn. of Prodasineura autumnalis (Fraser), see Lieftinck (1934e: 390). 
crenitis Lieftinck (Drepanosticta), 1933a: 288—292, 296, fig. 3. Holotype g' and 
first described 9. W. Borneo, Singkawang, G. Poteng, 325 m, forest brook, 28.II. 
1932, L. Coomans de Ruiter. — Platystictidae. 
cristatus Needham (Gomphoides), 1944 (3). First described © (in cop.). Surinam, 
Para River, 4.1.1959, J. Belle. See Belle, 1970: 126—130, fig. 202—210 (SQ and 
larva), pl. 17b, 18a, 21a (sub Negomphoides). — Gomphidae. 
crucigera Lieftinck (Archibasis), 1949a: 185, 192—193, fig. 176, 177, 226, 227, 239. 
Holotype g and first described 9. N. New Guinea, Humboldt Bay area, Tami River 
valley, 25—31.1.1933, W. Stüber. — Coenagrionidae. 
cruentata Lieftinck (Notoneura), 1932b: 522, 540—541, 601, fig. 38. Holotype ¢. 
N. New Guinea, Humboldt Bay area, near Lake Sentani, 9.III.1931, W. Stüber. 
First described 9, topotypical, XI.1931 & III.1932, same collector. — Protoneuridae. 
Now N. rosea cruentata Lieft. 
cyanura Lieftinck (Notoneura), 1932b: 521, 529-530, 601—602, fig. 23. Holotype 
d'. N. New Guinea, Humboldt Bay area, Hollandia, I—11.1931, W. Stüber. — First 
described 9. N. New Guinea, Humboldt Bay area, 400 km S. of Hollandia, 300 m, 
25.111.1932, W. Stüber. See also Lieftinck, 1937a: 11—12, fig. 5. — Protoneuridae. 
cyclopica Ris (Lanthanusa), 1912b: 746—747, fig. 424. Holotype 9. N. New 
Guinea, Humboldt Bay area, “N. Guinea Exp., Cycloop Gebergte, 14 April 1903” 
(two written labels), Mus. Leiden (print), Orthetrum? (unknown hand), “nov. gen. 
nov. spec. @”, det. Dr. F. Ris. — Libellulidae. 
Type-species of Lanthanusa Ris (op. cit.). 
cyclopica Lieftinck (subsp. of Palaiargia charmosyna Lieft.), 1949a: 118. Holotype gd 
and first described 9. N. New Guinea, Humboldt Bay area, Cycloop Mts., southern 
slopes, 900 m, 26.VI.1938, J. Olthof (3rd Archbold Exped.). — Coenagrionidae. 
cyclopica Lieftinck (Hemicordulia), 1942: 546—548, pl. 35 fig. 108—109. Holo- 
type g'. N. New Guinea, Humboldt Bay area, Cycloop Mts., 1000 m, 8.VIII.1935, 
W. Stiiber. — Corduliidae. 
cydippe Laidlaw (Macromia), 1922 (3). First described 9. W. Java, Djasinga, Tji- 
barangbang River, 150 m, 1.XI1.1935, M. A. Lieftinck. See Lieftinck, 1950b: 680, 
704—705. — Corduliidae. 
cynthiae Lieftinck [ subsp. of Agrionoptera insignis (Ramb.)], 1942: 472— 473, pl. 24 
fig. 19. Holotype g and first described 9. Tanimbar Is. (Timorlaut), P. Jamdena, 
IV.1938, P. Buwalda. — Libellulidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 83 


cyrene Lieftinck (Diplacina), 1953d: 155, 164—169, fig. 3—4. Lectotype &. W. 
New Guinea, Berau Peninsula (Vogelkop), Beraur distr., Klamono Oilfields, 40 km 
inland, ca. 50 m, 19.VIII.1948, M. A. Lieftinck. — First described 9. Misool I. 
(off W. New Guinea), Fakal, ca. 50 m, 23.1X.1948, M. A. Lieftinck. — Libel- 
lulidae. 


dactylogastra Lieftinck (Celebophlebia), 1936b: 401—403, fig. 1—2. Holotype à. 
Central S.W. Celebes, northeastern slopes of Quarles Mts., Todjambu, 1000 m, 
17.VIL.1936, L. J. Toxopeus. — Libellulidae. 

Type-species of Celebophlebia Lieftinck (op. cit.). 

dactylostyla Lieftinck (Amphicnemis), 1953f: 386—388, fig. 2. Holotype & and first 
described 9. S. Borneo, Sampit distr., 50— 100 m, Pemantan, 150 km inland, 26.VII. 
1953, forest marshes, M. A. Lieftinck. — Coenagrionidae. 

dagnyae Lieftinck (Idiocnemis), 1958: 258—260, 279, fig. 22—26. Holotype & and 
first described 9. W. New Guinea, western extremity, Malano, 15 km inland of 
Sorong, 27.VIII.1948 (4), and Remu near Sorong, 28.VIII.1948 (9), M. A. 
Lieftinck. — Platycnemididae. 

daphne Lieftinck (Huonia), 1953d: 186, 198—200, fig. 10B, 11, 12, 15, 17. Holo- 
type & and first described 9. Halmahera I. (N. Moluccas), Mumar River, 200—300 
m, 25.1X.1951 (4), and Mt. Sembilan, 600 m, 8.X.1951 ( © ), Sundanese collectors. 
— Libellulidae. 

debeauforti Lieftinck (Teinobasis), 1938: 107—109, 113, fig. 41—43. Holotype & 
and first described 9. N.-coast of New Guinea, estuary of Sermowai River, Moaif, 
26.VI—4.VII.1903, L. F. de Beaufort & H. A. Lorentz. Identified by H. W. van 


der Weele with “Telebasis spec. no. 1”. — Coenagrionidae. 
debeauxi Lieftinck (Teinobasis), 1938: 113—114, fig. 45—46. Holotype g'. Papua, 
Fiume Purari [Purari River}, 4.1.1894, Lamberto Loria. — Coenagrionidae. 


Erroneously recorded in the original description as from “Astrolabe Range”. 
declaratum Lieftinck [subsp. of Pseudagrion pilidorsum (Brauer) }, 1936a: 130, 132— 
134, fig. 13a-b. Lectotype g' and first described ©. W. Flores I. (Lesser Sunda Is.), 
Labuan Badjo, 100 m, XI.1937, J. K. de Jong. — Coenagrionidae. 
deflexum Lieftinck [subsp. of Pseudagrion pilidorsum (Brauer) }, 1936a: 130, 134, fig. 
13c. Lectotype &. S. Timor I. (Lesser Sunda Is.), Amarasi, XII.1931—1.1932, Ed. 
Handschin. — Coenagrionidae. 
degeneratum Lieftinck (Papuagrion), 1937a: 34—35. Holotype 9. N. New Guinea, 
Humboldt Bay area, Korimé River plain, Ajiep, I. 1931, W. Stiiber. — First described 
d. Same area, southern Bewani Hills, 300—500 m, Arso, 2.111.1937, W. Stüber 
(see Lieftinck, 1949a: 162, fig. 207-208). — Coenagrionidae. 
The ® was first described sub P. magnanimum (Selys), but later recognized as a 
distinct species (see Lieftinck, 1935c: 245, fig. 19, 247). 
delicatula Lieftinck (Caconeura), 1930b: 141—143, 151, fig. 5—8. Holotype & and 
first described 9. Java mer., res. Banjumas, Djeruklegi, 11.VIII and 20.X.1929, 
respectively, F. C. Drescher. — Protoneuridae. 
Now Prodasineura delicatula (Lieft.). 
demerarae Selys (Cyanogomphus), 1894 (& incomplete). First described 9 in good 
condition. Surinam, Zanderij, Troelinde Creek, 1.VIII.1959, J. Belle. See Belle, 1966b: 
46—52 (3 2), fig. 72—82, pl. 7 (sub Ebegomphus). — Gomphidae. 


84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Later, Belle (1970) placed Ebegomphus Needham in the synonymy of Cyanogom- 
phus Selys, and E. strumens Needham, 1944, as a synonym of C. demerarae Selys. 
dentata Geijskes (Coryphaeschna), 1943: 64—71, fig. 1 & 2 (larva). Holotype gd. 

Surinam, Palumeu River near Indian village Julu, 3.1.1941, L. Schmidt. — Aeshnidae. 
Currently placed in Triacanthagyna Selys. 
diadesma Lieftinck (Notoneura), 1936a: 120—121, fig. 5, 7. Holotype & (teneral). 
E. Sumba I. (Lesser Sunda Is.), Kananggar, 700 m, V.1925, K. W. Dammerman. — 
First described 9. E. Sumba, Mau Marru, 500 m, 18—23.VII.1949 (Sumba Exped. 
1949, NMB), see Lieftinck, 1953c: 152—154, fig. 7—9. Adults of both sexes from 
same locality also in ML. — Protoneuridae. 
dictatrix Lieftinck (Oreaeschna), 1937a: 77—81, fig. 46—47. Holotype & and first 
described 9. N. New Guinea, Humboldt Bay area, southern slopes of Mt. Cycloop, 
3700 ft., 2.1X.1933 (4 ) and id., 1000 m, 2.IX.1935 (9), W. Stüber. — Aeshni- 
dae. Plate 5. 
Type-species of Oreaeschna Lieftinck (op. cit.). 
digitiferum Lieftinck (Papuagrion), 1949a: 155—156, fig. 196—198, 202. Holotype 
o and first described 9. Central N. New Guinea, Sahuweri River basin, Sigi Camp, 
1500 m, 20.11.1939 (g') and Mist Camp, 1800 m, 9.1.1939 (2), L. J. Toxopeus 
(3rd Archbold Exped.). — Coenagrionidae. 
dimidiata Selys (Dysphaea), 1853 (&). First described 9 (semiad.), Sumatra, Central 
Sumatra, labelled “Silago tot 12/7 77”, “Sumatra Exp.”, and “Dysphaea dimidiata 
Sel. 9”, in H. Albarda’s writing. See Albarda, 1881: 6, pl. 2 fig. 2. — Euphaeidae. 
Type-species of Dysphaea Selys (op. cit.). 
diminuta Lieftinck (subsp. of Huonia arborophila Lieft.), 1963c: 778, fig. 38. 
Holotype g. Misima I. (St. Aignan, Louisiade Archip.), north slopes, Sisa, 350 m, 
19.VII.1956, L. J. Brass (Sth Archbold Exped.). — Libellulidae. 
dioxippe Lieftinck (Diplacina), 1963c: 764—765, fig. 16—19. Lectotype g and first 
described ©. Central W. New Guinea, Central Range, Sterren Mts, Ok Temna, 
1500 m, 19.V.1959 (Neth. New Guinea Exped.). — Libellulidae. 
dissoluta Lieftinck (subsp. of Agrionoptera longitudinalis Selys), 1963c: 771—772. 
Holotype 8 and first described 9. New Ireland (Bismarck Archip.), Kandan, 
25.X11.1959, W. W. Brandt. — Libellulidae. 
divaricatus Lieftinck (Burmagomphus), 1964b: 16, 26—29, fig. 29— 35. Holotype 
d and first described 9. Malay Peninsula, Kelantan, Ulu Kelantan, Sungai Nengiri, 
Fort Brooke, 700 m, 31.VII.1963, J. I. Furtado (g') and Selangor, 1114 mi. 
Kuala Lumpur-Bentong Rd., Sungai Gombak, 16.11.1963, J. I. Furtado & M. A. 
Lieftinck (9, reared from larva, emerged at Kuala Lumpur). — Gomphidae. 
dohrni Krueger (Idionyx), 1899: 326—330. Lectotype g (head gone) and first 
described 9, [N.E.} Sumatra {Deli} Soekaranda Dohrn (print), Idionyx dohrni 
Krüger, L. Krüger, d und 9, determ. 1927 (written), ex Mus Stettin. — Corduliidae. 
Synonymous with I. yolanda Selys. 
dominula Lieftinck (Teinobasis), 1937a: 46—47, fig. 32. Holotype 8 and first 
described 9. N. New Guinea, Upper course of Korimé River, Tarafia, ca. 600 m, 
10.V.1935, W. Stüber. — Coenagrionidae. 
dorcadion Lieftinck (Drepanosticta), 1949a: 59—61, fig. 32, 45—49, 52. Holotype 
d and first described 9. Central W. New Guinea, Sahuweri River valley, Rattan 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 85 


Camp, 1150 m, 12.11.1939, L. J. Toxopeus (3rd Archbold Exped.). — Platystictidae. 

dorsocyana Lieftinck (Libellago), 1937b: 56—57, fig. 1. Holotype g'. S. Borneo, 
Sampit area, Kota Waringin River, Riamtiwata, XII.1935, J. J. Menden. — Chloro- 
cyphidae. 

dorsosanguinea Lieftinck (Rhinocypha), 1961a: 124—125. Holotype &. S. Philippine 
Is., Basilan I., Maloong, 25.IX.1935, K. Kuwasima. — Chlorocyphidae. 

dravida Lieftinck (Gynacantha), 1960a: 252—253. Holotype ¢ and first described 9. 
S.W. Peninsular India, Coorg, Hallery near Mercara, 4000 ft., 20.V.1922 (4), 
and Coorg, Fraserpet, 2000 ft, 25.IX.1923, “brook emptying in Lotus-tank” (9), 
F. C. Fraser, both misidentified by F. C. Fraser as G. hyalina Selys. — Aeshnidae. 

drescheri Lieftinck (Heliogomphus), 1929b: 121—123, fig. 12—13. Holotype dg 
and first described 9. Central Java, Banjumas, G. Slamet, Baturraden, 2500 ft., 27 
and 28.VII.1928, respectively, F. C. Drescher. — Gomphidae. Plate 4. 

drusilla Lieftinck (Drepanosticta), 1934d: 474—476, fig. 4—5. Holotype & and 
first described 9. W. Borneo, Singkawang-Bengkajang Rd., forest brook near Seru- 
kan, hill-country, 30.VIII.1932 (g') and same district, Sungei Bagak (Mt. Raja 
complex), hill country, 7.IX.1932 (9), L. Coomans de Ruiter. — Platystictidae. 

drymobia Lieftinck (Selysioneura), 1959a: 284—286, fig. 3, 8, 15—16. Holotype &. 
Misima I. (St. Aignan I., Louisiade Archip.), Sisa, north slopes, 350 m, 22.VII.1956, 
L. J. Brass (Sth Archbold Exped.). — Isostictidae. Plate 1. 

dupophila Lieftinck (Drepanosticta), 1933a: 286—288, 296, fig. 2. Holotype {. 
W. Borneo, Singkawang, swampy forest near Pandjoa, 3.1V.1932, L. Coomans de 
Ruiter. — Platystictidae. 


eclecta Lieftinck (Palaiargia), 1949a: 114, 127—131, fig. 149—152. Holotype gd 
and first described 9. West Central New Guinea, Wissel Lakes group, Arabu 
Bivouac, 10 mi. N.E. of Lake Paniai, ca. 1800 m, 15.X.1939 (g') and 23.X.1939 
(2), H. Boschma (Le Roux Exped.). — Coenagrionidae. 

efasciata Lieftinck (subsp. of Notoneura erythrura Lieft.), 1949a: 73. Holotype à. 
Central N. New Guinea, Idenburg River valley, Bernhard Camp, 50 m, 23.VIII.1938, 
J. Olthof (3rd Archbold Exped.). — Protoneuridae. 

egregia Lieftinck (Notoneura), 1937a: 24, fig. 17. Holotype 4 and first described 9. 
Boeroe (Buru I.), S. Moluccas, Bah’lalé, 600 m, 10.V.1921 (4, abdomen missing) 
and 22.1.1922 (9), L. J. Toxopeus. — Protoneuridae. 

Nom. nov. pro Caconeura eburnea: Ris, 1929: 144, fig. 3—4, g' Q Buru I, nec 

C. eburnea Foerster, § Kei Is., a congeneric species. 

ekari Lieftinck (Papuagrion), 1949a: 148—150, fig. 182—185, 191. Holotype & 
and first described 9. West Central New Guinea, Wissel Lakes group, Arabu Bi- 
vouac, 10 mi. N.E. of Lake Paniai, ca. 1800 m, 25.X.1939 (G'}) and 15.X.1939 (9), 
H. Boschma (Le Roux Exped.). — Coenagrionidae. 

elegans Selys (Aeschnosoma), 1871 (9). First described g'. Surinam, Nassau Mts., 
bush creek 2 km from Marowijne River, 17.11.1949, D. C. Geijskes. See Geijskes, 
1970: 19—22 (8 2), fig. 7, 11, 1416, 20—23, pl. 1b. — Corduliidae. 

elegans Belle (Aphylla), 1970: 58—60, fig. 90—92. Holotype g'. Venezuela, Apure, 
2.11.1937. — Gomphidae. 

elegans Lieftinck (Leptogomphus), 1948a: 254—258, fig. 12 & pl. 8. Holotype 8 
and first described 9. E. China, Prov. Fu Kien, Kuatun, 2300 m, 7.VII. 1938 (4) 
and 1.VII.1938 (9), J. Klapperich. — Gomphidae. 


86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


ellioti Lieftinck (Lokia), 1969c: 15—16. Holotype 3. Zambia, Upper Zambezi Region, 
North Mwinilunga, Ikelenge distr., Zambezi Rapids, 21.1.1965, E. C. G. Pinhey. — 
Libellulidae. 

emarginata Lieftinck (Teinobasis), 1949b: 344—345, fig. 6. Holotype 9. Solomon 
Is., Shortland I., Hisiai River, 22.X.1936, R. A. Lever. — Coenagrionidae. 

emphyla Lieftinck (Notoneura), 1936a: 118—120, fig. 5—6. Holotype ¢ (teneral). 
Centr. Flores I. (Lesser Sunda Is.), Badjawa, 1200 m, XII.1931, Ed. Handschin. 
First described © (and adult 9). Flores I., Soa, 22—25.VII.1950, G. Kramer 
(Lieftinck, 1953c: 151—152). — Protoneuridae. 

enganoense Lieftinck [subsp. of Pseudagrion pilidorsum (Brauer) }, 1948b: 290—291, 
fig. 6. Lectotype 8 and first described 9. Engano I. (off S.W. Sumatra), Buahbuah, 
30.V.1936 (4 ) and Kiojoh, 13.VI.1936 (@), J. K. de Jong et al. — Coen- 
agrionidae. 

ensifera Lieftinck (Argiocnemis), 1932b: 588, 589—591, fig. 64. Lectotype @ and 
first described 9. N. New Guinea, Humboldt Bay area, Hollandia, 27.VIII—4.IX. 
1930, W. Stüber. — Coenagrionidae. 

eos Lieftinck (Palaiargia), 1938: 95—96, fig. 31—32. Holotype g'. N.W. New 
Guinea, Berau Peninsula (Vogelkop), Vengatap near Aja Maru, 500 m, XII.1937, 
J. M. van Ravenswaay Claasen. — Coenagrionidae. 

ephippiata Lieftinck (Drepanosticta), 1937b: 72—74, fig. 9. Lectotype g'. N. Ce- 
lebes, Tondano near Manado, IV.1935, C. van Braekel, acq. 11.VI.1935. — Platy- 
stictidae. 

erato Lieftinck (Macromia), 1950b: 673—676, 689, 693—699, fig. 4, 5, 14, 15, 23, 
31, 38, 45—47. Holotype g'. W. Java, Bogor (Buitenzorg, olim), 250 m, Botanic 
Garden, 17.11.1945, native collector. First described 9. W. Java, Djasinga, Tjibarang- 
bang River, 150 m, 4.XI1.1938, M. A. Lieftinck. — Corduliidae. 

ericae Belle (Ischnogomphus), 1966b: 39—44, fig. 61—67 & 68—71 (larva, sup- 
position), pl. VIa-b. Holotype 4 and first described 9. Surinam, Upper Para River, 
8.IX.1962 (3) and 10.VIII.1960 (@), J. Belle. — Gomphidae. 

ericetorum Lieftinck (Hemicordulia), 1942: 556—558, pl. 36 fig. 112—113, pl. 37 
fig. 118—119. Holotype @ and first described 9. Central N. New Guinea, Baliem 
River valley, nr. Baliem Camp, 1650 m, 14.X11.1938, L. J. Toxopeus (3rd Archbold 
Exped.). — Corduliidae. 

erigone Lieftinck (Diplacina), 1953d: 157, 177—179, fig. 3—4 Holotype g' and 
first described ©. Misool I. (off W. New Guinea), Fakal, ca. 150 m, 4.X.1948, M. 
A. Lieftinck. — Libellulidae. 

erminea Lieftinck (Amphicnemis), 1953e: 247—251, fig. 6c-e. Holotype & and first 
described ©. S.E. Borneo, Kandangan distr., Ampah, 0—20 m, IV—V. 1948, Liem 
Swie Liong. — Coenagrionidae. 

errans Lieftinck (subsp. of Zygonyx iris Selys), 1953e: 266—268. Holotype g' and 
first described 9. W. Borneo, Singkawang, Mt. Poteng, 400 m, X.1932, L. Coomans 
de Ruiter. — Libellulidae. 

erratica Lieftinck (Macromidia), 1948: 274—278, fig. 18—19. Holotype g' and 
first described ©. S. Sumatra, Lampong distr., Giesting near Talangpadang, Wai 
Tebu, ca. 500 m, 19—31.III.1940, M. A. Lieftinck. — Corduliidae. 

Now classified as a subspecies of M. genitalis Laidlaw (see Lieftinck, 1971b: 

202221) 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 87 


erythromma Lieftinck (Elattoneura), 1953f: 383—385, fig. 1a-e. Holotype & and 
first described 9. S. Borneo, Sampit distr., 0—50 m, near Sampit, about 50 km 
inland, 21.VII.1953, M. A. Lieftinck, with collector's note: “Very dull, except bril- 
liant cherry-red eyes”. — Protoneuridae. 
erythrostigma Lieftinck (Paramecocnemis), 1932b: 504—508, fig. 9—13. Lectotype 
d and allotype @ (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, ca. 
300 m, IV—VI.1931, W. Stiiber. — Platycnemididae. 
Type-species of Paramecocnemis Lieft. (op. cit.). 
erythrura Lieftinck (Notoneura), 1932b: 522, 525, 530—532, fig. 2426. Lectotype 
d' and allotype ® (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, I— 
11.1931, W. Stüber. — Protoneuridae. 
Now N. erythrura erythrura Lieft. 
esuriens Lieftinck (Argiolestes), 1956b: 71, 93—94, fig. 28—29. Holotype &. 
Papua, Maneau Range, N. slopes of Mt. Dayman, 1550 m, 6.VII.1953, G. M. Tate 
(4th Archbold Exped.). — Megapodagrionidae. 
eucera Lieftinck (Drepanosticta), 1949a: 56—57, fig. 30, 41—44, 50. Holotype & 
and first described 9. Central W. New Guinea, Idenburg River basin, above Bern- 
hard Camp, 700 m, 27—29.111.1939, L. J. Toxopeus (3rd Archbold Exped.). — 
Platystictidae. 
euglena Lieftinck (Teinobasis), 1934a: 8—11, fig. 3. Holotype ¢ and first described 
Q. Mid Java, South Banjumas, Djeruklegi, 7.11.1931, F. C. Drescher. — Coena- 
grionidae. 
excelsa Lieftinck [subsp. of Neurothemis intermedia (Ramb.)}, 1934c: 269—270. 
Holotype &. East [rect. Central N.] Java, Rembang, Kedangan, 40 m, 2.X11.1937, 
F. Th. Verbeek. — First described 9. Java, Kangean I. (Java Sea) and Sumba; not 
specified (see Lieftinck, 1953c: 213—214). — Libellulidae. 
exoleta Lieftinck (Drepanosticta), 1932b: 515, 517—519, fig. 19. Lectotype g' and 
first described 9. N. New Guinea, Humboldt Bay area, Hollandia, IV—VI.1931, 
W. Stüber. — Platystictidae. 
exul Selys (Alloneura), 1878 (&). First described 9. Halmahera I. (N. Moluccas), 
labelled “Bernstein, Noord Halmahera” (print), and old drawer-label “exul Hagen” 
in H. A. Hagen’s writing. Collected in the years 1861 or 1862. See Lieftinck, 1938: 
89— 90. — Protoneuridae. 
Currently placed as Notoneura exul (Selys). 


falcatum Lieftinck (Mortonagrion), 1934a: 12—15, fig. 5. Lectotype g' and first 
described ©. Karimon Djawa Is. (Java Sea), P. Karimondjawa, mangrove-swamps 
near debouchment of small brook Todjero, 28.X1.1930, M. A. Lieftinck. — Coen- 
agrionidae. 

farinicolle Lieftinck (Pseudagrion), 1932b: 568, 572—573, fig. 56. Lectotype 4 and 
allotype @ (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, XI—XN. 
1930, W. Stüber. — Coenagrionidae. 

fasciculare Lieftinck (Aciagrion), 1934a: 15—17, fig. 6—7. Holotype g'. W. Java, 
South Preanger, Djampangs, ca. 800 m, near Sukanegara, 24.X11.1931, M. A. Lief- 
tinck. — Coenagrionidae. 

fenella Campion (Synthemis), 1921 (¢). First described 9. New Caledonia, Forêt de 


88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Thi, 100—300 m, 28—29.III.1961, J. Sedlacek. See Lieftinck (1971b: 53—54, fig. 
37—38). — Corduliidae. 
ferentina Lieftinck (Huonia), 1953d: 189, 205—207, fig. 17—18. Holotype d. 
Halmahera I. (N. Moluccas), Mt. Sembilan, 600 m, 14.X.1951, Sundanese collector. 
| — Libellulidae. 
feronia Lieftinck (Protosticta), 1933a: 281—285, fig. 1. Holotype g and allotype 9 
(in cop.). W. Borneo, Singkawang, G. Poteng, 325 m, forest brook, 1.X1.1932, L. 
Coomans de Ruiter. — Platystictidae. 
feronia Lieftinck (Synthemis), 1938: 122—124, fig. 51. Holotype g' and first 
described 9. N. New Guinea, Bewani Hills, upper course of Tami River, 300— 500 
m, Josko, 22—30.X1.1936, W. Stüber. — Corduliidae. 
feuerborni Schmidt (Aciagrion), 1934: 344—346, fig. 4346. Lectotype g' and first 
described 9. N.E. Central Sumatra, Lake Toba area, Huta Gindjang, 3.1V.1929 
(Deutsch. limnol. Sunda-Exped. 1928—29). Selected by E. Schmidt and presented 
to author in 1935. — Coenagrionidae. 
filicornis Lieftinck (Torrenticnemis), 1949a: 104—107, fig. 101, 113, 129—130, 
137—138. Holotype ¢ and allotype 2 (in cop.). Central W. New Guinea, Sahuweri 
River basin, Sigi Camp, 1500 m, 15.11.1939, L. J. Toxopeus (3rd Archbold Exped.). 
— Platycnemididae. 
Type-species of Torrenticnemis Lieft. (op. cit.). 
filostyla Martin (Heliaeschna), 1907b: 221—222. Holotype & (slightly immature, 
head missing). Rosenberg, Celebes (printed), and “Heliaeschna filostyla Martin 
9 [Type au Musée de Leyden”, in R. Martin's handwriting. First described © (im- 
mature). Same locality label with additional Panybie (written), and Heliaeschna 
filostyla Q Type, in R. Martin's writing. — Aeshnidae. 
(See also Martin, 1909b: 165—167, fig. 167—168, incl. H. gladiostyla Martin, 3 
Celebes. — Syn. nov.). 
fissicollis Lieftinck (Stenosticta), 1932b: 551—557, fig. 48—51. Lectotype g' and 
allotype ® (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, Pim River, 
ca. 200 m, III.1931, W. Stüber. — Isostictidae. 
Type-species of Stenosticta Lieftinck, nom. praeocc. (op. cit.). 
Now Tanymecosticta fissicollis (Lieft.). 
fissidens Lieftinck (Idiocnemis), 1958: 260—261, 281, fig. 1—5. Holotype & 
and first described 9. Waigeu I., off N.W. New Guinea, labelled “Waigeoe [H. 
A.} Bernstein”. — Platycnemididae. 
flammula Lieftinck (Prodasineura), 1948a: 227—231, fig. 1, 3. Holotype d. 
E. Borneo, Kutai, Batau Besi, near Sangkulirang, VI.1937, M. E. Walsh. — Pro- 
toneuridae. 
flammula Lieftinck (subsp. of Palaiargia ceyx Lieft.), 1949a: 115, 125—126, fig. 
136. Holotype 8 and first described 9. Central N. New Guinea, Sahuweri River 
valley, Sigi Camp, 1500 m, 17.11.1939 (g) and 20.11.1939 (@), L. J. Toxopeus 
(3rd Archbold Exped.). — Coenagrionidae. 
flavipedum Lieftinck (Papuagrion), 1949a: 156—157, fig. 186—187. Lectotype d'. 
W. New Guinea, Onin Peninsula, Fak Fak, sea-level, VII.1939, R. G. Wind. — 
Coenagrionidae. 
floresianus Lieftinck (Lestes, sg. Indolestes), 1960c: 141—143, pl. 2 fig. 1, fig. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 89 


16, 17. Holotype g'. Flores I. (Lesser Sunda Is.), Rana Mese, W. Flores, 1200 m, 
5.1V.1958, A. M. R. Wegner. — Lestidae. 

fluviatilis Lieftinck (Chalybeothemis), 1933b: 132—137, pl. 13 fig. 1, pl. 14 fig. 
2, pl. 15 fig. 3. Lectotype g' and first described 9. W. Borneo, Pontianak, Peniti 
River, 24.11.1931, L. Coomans de Ruiter. — Libellulidae. 

Type-species of Chalybeothemts Lieftinck (op. cit). 

foliacea Lieftinck (Oligoaeschna), 1968a: 149, 153, 169—171, fig. 4, 6, 7. Holo- 
type of and first described 9. W. Borneo, Singkawang area, Mt. Poteng, 400 m, 
15.1V.1934 (G') and Patengahan-Bakuan road, 20.VII.1931 (2), L. Coomans de 
Ruiter. — Aeshnidae. 

foliosa Navas (Platycnemis), 1932: 9—10, fig. 21a-c. Holotype 4. E. China, Shang- 
hai: Zô-sè (China) 13.V.30, and Platycnemis foliosa 4 Nav., P. Navás S. J. det., 
both labels in L. Navas’ writing; with pink label Typus (id.). Ex coll. L. Navas, 
pres. 1934. — Platycnemididae. 

fonticola Lieftinck (Notoneura), 1932b: 524, 528, 533535, fig. 29—31. Lectotype & 
and allotype @ (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, III.1931, 
W. Stüber. — Protoneuridae. 

fontinalis Lieftinck (Argiolestes), 1956b: 69, 84—85, fig. 11, 16; 109—110, fig. 
49— 52. Holotype 4 and first described 9. W. New Guinea, Berau Peninsula 
(Vogelkop), Beraur River area, Klamono oilfields (50 km from westcoast inland), 
50 m, 20—24.VIII.1948, M. A. Lieftinck. — Megapodagrionidae. 

forficulatum Lieftinck (Mortonagrion), 1953f: 392—395, fig. 3, 3a. Holotype & 
and first described 9. S. Borneo, Sampit distr., 50—100 m, Pemantan, 150 km 
inland, 27.VII.1953, M. A. Lieftinck. — Coenagrionidae. 

fraterculum Lieftinck (Papuagrion), 1937a: 26, 41—43, fig. 29—30. Lectotype & 
and allotype Q (in cop.). N. New Guinea, southern slope of Mt. Cycloop, 1000 m, 
18—23.IX.1935, W. Stüber. — Coenagrionidae. 

fraterna Albarda (Trithemis), 1881: 4. Holotype g'. Central Sumatra, Sungei Abu, 
labelled “vfan} Soengfei} aboe nfaar] Moeafra} Lab[oe} 3—10.10.1877”, Su- 
matra Exp., “Trithemis fraterna Albarda @”, in H. Albarda’s writing, coll. Albarda 
acq. 1892 (print). With Ris’s identification label: Trithemis aurora Bm., det. Dr. F. 
Ris. — Libellulidae. 

Currently placed as a synonym of T. aurora (Burm.), see Ris (1912b: 778), who 
considers it an individual variation. It is, however, possibly a distinct subspecies, or 
even a good species. 

Preoccupied by T. fraterna (Hagen, 1873) Kirby, 1890, a synonym of Erythrodi- 
plax c. connata (Burm.); and, if Albarda’s insect proves to be distinct taxonomically, 
it would require a new name. 

frubstorferi Krueger (Alloneura), 1898 (3). First described 9. Central Java, Ban- 
jumas, Mt. Slamet, Baturraden, 760 m, 16.1V.1928, F. C. Drescher. See Lieftinck, 
1930b: 147—151, fig. 13—16. — Protoneuridae. 

Currently known as Notoneura insignis (Selys). 

fulgens Ris (Diplacina), 1898: 323—324 (incomplete 4 and 9 New Britain). First 
described complete &. New Britain (Bismarck Arch.), Vanubakan, near Kerawa, 
11—20.X1.1959, J. L. Gressitt. See Lieftinck, 1963c: 768—769, fig. 24—27. — 
Libellulidae. 
Holotype & (incomplete), New Britain (MNB). 


90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


fulgens Lieftinck (Teinobasis), 1949a: 172—174, fig. 220—221. Holotype &. S.W. 
New Guinea, Sungei Arja near Umar, W. of Uta (southcoast), 26.VI.1941, E. 
Lundqvist (Negumy Exped.). — Coenagrionidae. 

fulvia Fraser (Gynacantha), 1926: 477478. Holotype 9. S. New Guinea, Frederik 
Hendrik Eiland [Marind Anim, south of Digul River debouchment], III.1910, coll. 
unknown. — Aeshnidae. 

Syn. of Gynacantha kirbyi Krueger. — Syn. nov. 

funereum Lieftinck (Thaumatagrion), 1932b: 508—514, fig. 14—17. Lectotype ¢ 
and first described 9. N. New Guinea, Humboldt Bay area, Pim River, 200 m, III. 
1931, W. Stüber. — Platycnemididae. 

Type-species of Thaumatagrion Lieftinck (op. cit.). 

furcifer Lieftinck (Podolestes), 1950c: 40, 44—47, fig. 3—4. Holotype g'. S. Borneo, 

Sampit, 0—50 m, 11.1950, W. Buyn. — Megapodagrionidae. 


gazella Lieftinck (Drepanosticta), 1929b: 110—112, 115, fig. 1—3. Holotype 4 and 
first described 9. Central Java, Banjumas, Mt. Slamet, Baturraden, 2500 ft., 13.VI. 
1928 (&) and 9.VI.1928 (9 }, F. C. Drescher. — Platystictidae. 
geijskesi Needham (Progomphus), 1944 (9). First described G'. Surinam, Suriname 
River, Aroesobanja Falls, 27.V.1959, J. Belle. See Belle, 1966a: 7—11, fig. 13—20, 
pl. 2. — Gomphidae. 
geijskesi Belle (Rhodopygia), 1964b: 48—54, fig. 22—25. Holotype 8 and first 
described ©. Surinam, Republiek, 26.XII.1961 (4) and Zanderij (Bos Bivak), 
1.XII.1962 (9), J. Belle. — Libellulidae. 
geminata Fraser [subsp. of Caconeura salomonis (Selys) }, 1926: 492— 493. Lectotype 
d and first described 9. Kei Is, Groot Kei, Gg. Daab, no. 149, 1922 (8) and 
Elat, no. 156 (@), H. C. Siebers. — Protoneuridae. 
Syn. of Notoneura eburnea (Foerster), see Lieftinck, 1937a: 22—23, fig. 15—16 
(4 $ types of Caconeura salomonis geminata Fraser). 
Currently placed in Notoneura Tillyard. 
geometricus Selys (Onychogomphus), 1854: 31. Holotype 9. Java, labelled “Java 
Kluhl} & v. Hfasselt}” (white disk), with identification label of M.A.L. and 
“holotype” (see also Lieftinck, 1929b: 131—133). — Gomphidae. 
First described g from Java, by de Selys & Hagen (1858: 280—282, 20—22 sep., 
pl. 1 fig. 1) (IRSN). 
glochidion Lieftinck (Bironides), 1963c: 759—761, fig. 8—11. Holotype g and 
first described @. Papua, Central Province, 75 km W. of Pt. Moresby, Doa Estate, 
13.X.1962 (g') and 19.IX.1962 (© }, R. Straatman. — Libellulidae. 
gloriosa Lieftinck (Archboldargia), 1949a: 145—146, fig. 144—144a, 147, 162. 
Holotype & (immature). Central N. New Guinea, Sahuweri River valley, Top 
Camp, 2100 m, 29.1.1939, L. J. Toxopeus. — Coenagrionidae. 
gomphoides Lieftinck (Microtrigonia), 1933d: 36—39, fig. 23—25. Lectotype & 
and first described 9. N. New Guinea, Humboldt Bay area, foot of Cycloop Mts., 
400 m, IV—VI.1931, W. Stiiber. — Libellulidae. Plate 6. 
goniocercus Lieftinck (Lestes, sg. Indolestes), 1960c: 154—155, 159, fig. 24. Holo- 
type d. W. New Guinea, western extremity, Malano, 15 km S.E. of Sorong, 2.IX. 
1948, M. A. Lieftinck. — Lestidae. 
gracilenta Lieftinck (Synthemis), 1935c: 285, 290—293, fig. 45. Lectotype & and 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 91 


first described 9. N. New Guinea, Humboldt Bay area, Cycloop Mts., 1000 m, 
13.1X.1932 (d') and 9.IX.1932, W. Stüber. — Corduliidae. 
gracilis Albarda #7 Selys (Libellula), 1887: 15—16. Lectotype & and first described 
Q. Iran, both labelled “N. Perzië, Sibakuh” and on the reverse side, “Libella gra- 
cilis Albarda”, in H. Albarda’s writing, Coll. Albarda acq. 1892 (print). Both with 
Ris’s identification label Orthetrum ransonnetii Br., det. Dr. F. Ris. — Libellulidae. 
Syn. of Orthetrum ransonneti (Brauer). 
gracillima Fraser (Teinobasis), 1926: 494, fig. 7. Holotype 8. A papered specimen, 
marked “T(9)” on the envelope and described as from “Java” (err. pro Celebes!) 
— Coenagrionidae. 
Erroneously considered synonymous with T. superba (Selys) by Lieftinck, 1935c: 
254. Bona species. 
guyanensis Belle (Progomphus), 1966a: 20—24, fig. 33—43, 45, pl. 3. Holotype gf 
and first described 9. Surinam, Upper Coropina River (Dauwdropkamp), 26.XII. 
1959 (8), and Upper Para River, 30.1V.1963 (£), J. Belle. — Gomphidae. 
haematosoma Lieftinck (Prodasineura), 1937b: 84—86, fig. 16. Holotype & and 
first described 9. W. Borneo, Singkawang, forest brook near Bengkajang, 15.VI (g') 
and 13.X.1933 (9), L. Coomans de Ruiter. — Protoneuridae. 
halcyon Lieftinck (Palaiargia), 1938: 97—99, fig. 33. Lectotype g' and first described 
9. N. New Guinea, southern Bewani Hills, upper course of Tami River, 5 days’ 
journey S.E. of Hollandia, 250 m, Fumb (Keerom?) River, 25—27.1V.1937, W. 
Stüber. — Coenagrionidae. 
hamifera Lieftinck (Macromia), 1955c: 253—256, fig. 1—4. Holotype 4. E. China, 
Prov. Fu Kien, Kuatun, Chung-an Hsien, 2300 m, 22—25.VIII.1943, T. C. Maa. — 
Corduliidae. 
harrissoni Lieftinck (Podolestes), 1953e: 233— 236, fig. 1—2. Holotype & and first 
described 9. N.W. Borneo, Sarawak, Kuching, Matang Road, 3rd mile, 22.1X.1950, 
M. A. Lieftinck. — Megapodagrionidae. 
helvola Lieftinck (Teinobasis), 1930b: 155—157, fig. 21. Holotype g' and allotype 
® (in cop.), S. Celebes, Maros, VI.1929, G. Overdijkink. — Coenagrionidae. 
hermione Lieftinck (Macromia), 1952a: 442, 454—456, 461, 467—468, fig. 4, 9, 18, 
22. Lectotype g' and first described 9. Misool I., off W. New Guinea, surroundings 
of Fakal, ca. 50 m, 18.X.1948 (7) and 23.IX.1948 (9), M. A. Lieftinck. — 
Corduliidae. 
heterostylus Lieftinck (Platylestes), 1932c: 248—251, fig. 1—2. Holotype &. 
S.W. Java, res. Banjumas, Djeruklegi, 28.1.1931, “Head and eyes dark brown. Thorax 
grass-green with black spots. Abdomen grass-green with black rings at articulations. 
Appendages white” (living colours, F. C. Drescher leg.). — Lestidae. 
First described 9 from Malaya and Sunda Islands (Lieftinck, 1960c: 135—136, 
fig.). 
Now Lestes (Platylestes) heterostylus Lieft. 
hilbrandi Lieftinck (Hemicordulia), 1942: 548—549, pl. 35 fig. 110—111. Holo- 
type d'. West Central New Guinea, Wissel Lakes group, Lake Paniai, 1742 m, 
4.1X.1939, H. Boschma (Le Roux Exped.). — Corduliidae. 
hippolyte Lieftinck (Diplacina), 1933d: 44, 57—58, fig. 31, 33. Lectotype & and 
first described 9. N. New Guinea, Humboldt Bay area, Cycloop Mts., 1000 m, 
13.IX.1932, W. Stüber. — Libellulidae. 


92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


hoogerwerfi Lieftinck (Ceriagrion), 1940b: 360—361, fig. 14. Holotype &. N. 
Sumatra, Atjeh, Meluwak, Laut Tiga Sagi (mountain lake), ca. 1500 m, 24.IV.1937, 
A. Hoogerwerf. — First described 9. S.W. Sumatra, Benkulen, Mt. Dempo, Tebat 
Kerindjing-ketjil, 1100 m, X.1941, W. C. Verboom. (See Asahina, 1967: 271—272, 
322, fig. 46—52). 

Now classified as a subspecies of C. bellona Laidlaw. 

humeralis Lieftinck (subsp. of Teinobasis serena Lieft.), 1949a: 168—169, fig. 172, 
217. Lectotype 4 and first described 9 (dark colour-phase). N. New Guinea, Hum- 
boldt Bay area ‚upper reaches of Bewani River system, Keerom River distr., Tompar, 
25.VI.1939, W. Stüber. — Coenagrionidae. 

humphriesi Lieftinck (Lestes), 1952b: 126—128, fig. 1—2. Holotype g and first 
described @. S. Australia, Murray River near Renmark, 13.IX.1949, L. Humphries. 
— Lestidae. 

Synonymous with Lestes (Austrolestes) aridus Tillyard, see Fraser, 1960: 24. 

huonensis Lieftinck (Idiocnemis), 1958: 267—270, 284, fig. 27—31. Holotype & 
and first described 9. N.E. New Guinea, Huon Gulf, labelled “Sattelberg (Gega- 
galu), Sommer 1900, Carl Wahnes” (F. Förster's writing), identified by F. Förster 
with “I. inornata Selys Subrasse huonensis Foerster, Type” [nom. nud.}, nos. 2253 and 
2254, respectively, ex UMMZ. — Platycnemididae. 

hurley Tillyard (Rhyothemis), 1926 (&). First described 9. N. New Guinea, Hum- 
boldt Bay area, Hollandia and env., 1931—1937, various locs., no dates selected, 
W. Stüber. See Lieftinck, 1942: 502—504, pl. 30 fig. 62—65. — Libellulidae. 

The above females included the ‘“allotype”, which should be, however, assigned to 
a different (undescribed) subspecies; only the males reported from S. New Guinea 
(Digul River basin) are nominotypical hurley. 

hyalinus Selys (Chlorogomphus), 1869b: 202 (39 sep.). Nom. nov. pro C. magnificus 
Selys g'. Holotype g'. Muller Java (print). — Cordulegasteridae. 

The specimen is in a very dilapidated state, lacking its head, most of the thoracic 
segments and terminalia of abdomen. It was photographed in that condition by the 
late R. van Eecke at Leiden, who sent a copy to F. C. Fraser for publication of a wing- 
photograph reproduced in his revision (1929: 144, fig. 25A). 

Conspecific with C. magnificus Selys, of which it is the g'. 

hylophila Lieftinck (Huonia), 1942: 489, 492, pl. 26 fig. 36—37. Holotype à. 
Central N. New Guinea, Sahuweri River basin, Araucaria Camp, 800 m, 28.111.1939, 
L. J. Toxopeus. — Libellulidae. 

First described ® (incomplete). W. New Guinea, Vogelkop, Taminabuan (S.W.- 
coast), 16.III.1949, Sten Bergman. 


ianthinipennis Lieftinck (Neurobasis), 1949a: 17—18, 22—23, fig. 6. Holotype & 
and first described 9. N. New Guinea, Humboldt Bay area, southern slopes of 
Cycloop Mts., 900 m, 26.VI.1938, J. Olthof. — Calopterygidae. 

inaequale Lieftinck (Ceriagrion), 1932b: 592—594, fig. 65. Lectotype g and allo- 
type £ (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, 27.VII4.IX. 
1930, W. Stüber. — Coenagrionidae. 

See Asahina (1967: 289—291, fig. 116—122). 

incisura Lieftinck (Archibasis), 1949a: 185, 186, 188, fig. 229, 233. Holotype 4 

and first described 9. W. Borneo, Singkawang, forest-marsh near Bakuan, 19.VII. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 93 


1932 (g') and Sungei Bagak, 29.X11.1932 (2), L. Coomans de Ruiter. — Coena- 
grionidae. 

indicus Lieftinck (Anax), 1942: 589—591. Lectotype g'. Peninsular India, Coorg, 
Hallery near Mercara, 15.V.1923, F. C. Fraser leg. & don., misidentified as A. gut- 
tatus Burm. — Aeshnidae. 

First recognized as a distinct species upon a illustrated by Laidlaw (1921: 
83—84, fig. 2, g abdomen, sub A. guttatus Burm., Series A), from Barkuda I, 
1479/H2 (N.E. Peninsular India, Orissa, Chilka Lake, N. Annandale leg.), ex 
Indian Mus., and probably no more in existence. See also the correspondence between 
the late J. Cowley and the author, 77 Lieftinck, 1955e: 84—86. 

infans Lieftinck (subsp. of Nannophlebia imitans Ris), 1963c: 757—758. Holotype 
3. New Ireland (Bismarck Archip.), Kandan, 22.XII.1959, W. W. Brandt. — Li- 
bellulidae. 

infracavum Schmidt (Pseudagrion), 1934: 349—350, fig. 49 (g' E. Java). Lecto- 
type G'. E. Java, Ranu (Lake) Lamongan near Klakah, S.S. Hotel, X—X1.1928 
(Deutsch. limnol. Sunda-Expedition 1928—29). Selected by E. Schmidt and in 1935 
presented to the author. — Coenagrionidae. 

Syn. of P. nigrofasciatum Lieftinck, 1934 (see Lieftinck, 1936a: 124, footnote). 

inornata Selys (Idiocnemis), 1878b (g'). First described 9 (in cop.). W. New Guinea, 
Vogelkop (western extremity), env. of Sorong, ca. 50 m, forest stream, 24.X.1948, 
M. A. Lieftinck. See Lieftinck, 1958: 264—266, fig. 10. — Platycnemididae. 

inscriptus Hagen im Selys (Onychogomphus?), 1878a: 422—423. Holotype 9. 
“Java Kfuhl} & v. Hfasselt}” (white disk), over drawer-label “Inscriptus Hagen/ 
Java”. See Lieftinck, 1929b: 130—131, fig. 22. — Gomphidae. Plate 4. 

Currently known as Burmagomphus inscriptus (Hag. & de Selys) (= B. jacobsoni 

Ris, 1912, & holotype in SMF, g syntype in ML). 

insulare Lieftinck (Papuagrion), 1949a: 160, fig. 204—205. Holotype @. Schouten 
Is. (off N.W. New Guinea), Biak I., Bosnik, 15.V1.1938, L. J. Toxopeus. — Coen- 
agrionidae. 

interposita Lieftinck (Phasmosticta), 1951b: 17—24, fig. 8—10. Holotype g' (im- 
mature; wings and penile organ detached and papered separately). N. Australia, N. 
Queensland, Redlynch, 5.X1.1938, R. G. Wind. — Isostictidae. 

Type-species of Phasmosticta Lieftinck (op. cit.). 

A recent comparison (by J. A. L. Watson and the writer) of this individual with 
authentic specimens of Oristicta filicicola Tillyard, 1913 (terr. typ. Cooktown, N.Q.), 
a monotypic and inadequately described and figured zygopteron, reveals that these 
insects are probably conspecific, in which case P. interposita falls as a synonym of 
Tillyard’s species (J. A. L. Watson, unpublished). 

interrogata Selys (Agrionoptera), 1878b (9). First described g'. Biak I. (Schouten Is., 
off N.W. New Guinea), Bosnik, 16.V1.1939, L. J. Toxopeus. See Lieftinck, 1942: 
463, pl. 24 fig. 15. — Libellulidae. 

Now Nesoxenia mysis interrogata (Selys). 

inversa Lieftinck (Drepanosticta), 1949a: 55—56, fig. 29, 58—59. Holotype g'. W. 
New Guinea, Berau peninsula (Vogelkop), Manokwari, 15.VI.1938, L. J. Toxopeus. 
— Platystictidae. 

io Fraser (Rhinocypha), 1926: 485—486. Holotype g. “Sumatra” [err. pro Java!}, 
without further particulars. — Chlorocyphidae. 


94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Syn. of R. heterostigma Ramb., see Lieftinck, 1954: 11, footnote. 

irene Lieftinck (Notoneura), 1949a: 64—67, fig. 62—66, 87—88. Holotype & and 
first described 9. Central N. New Guinea, Sahuweri River valley, Araucaria Camp, 
700—800 m, 17 and 31.111.1939, respectively, L. J. Toxopeus & J. Olthof (3rd 
Archbold Exped.). — Protoneuridae. 

irene Lieftinck (subsp. of Rhyothemis princeps Kirby), 1942: 515—516, pl. 31 fig. 
70—73, pl. 33 fig. 88—89. Lectotype g' and first described 9. N. New Guinea, 
Humboldt Bay area, Hollandia, VII.1930, W. Stüber. — Libellulidae. 

irina Lieftinck (Macromia), 1950b: 714—716. Holotype 3. S. Celebes, foot of Mt. 
Lompobatang, Borong Rapoa, 800 m, VIII.1949, A. Diakonoff. — Corduliidae. 

irregularis Brauer (Tetrathemis), 1868 ( 9). First described G'. Basilan I. (S. Philippine 
Is.), Maloong, 15.X1.1932, K. Kuwasima. See Lieftinck, 1942: 448—450, pl. 
23 fig. 5 and 9. — Libellulidae. 

Now T. i. irregularis Brauer. 

ismene Lieftinck (Diplacina), 1933d: 44, 55—57, fig. 31—32. Lectotype g' and first 
described @. N. New Guinea, Humboldt Bay area, Hollandia, IV—VI.1931, W. 
Stüber. — Libellulidae. 

isoetes Lieftinck (Ischnura), 1949a: 225—229, fig. 277— 278, 281—284. Lectotype 
d and first described 9. Central W. New Guinea, Snow Mountain (Nassau) Range, 
Lake Habbema, 3250 m, 13.VIII. 1938, L. J. Toxopeus (3rd Archbold Exped.). — 
Coenagrionidae. 

[japponicus Selys (Sieboldius), 1854: 83. Types 4 and 9, given as from “Japon. 
(Musée de Leyde)”; & destroyed. — Gomphidae. 

This species is a resident of Malaya, Sumatra and Borneo. The type 4 was er- 
roneously recorded from Japan, the description being based on a specimen labelled 
“Muller Borneo”. This was examined by me in 1929, but has since been destroyed. 
An old @ (still extant) and labelled “Burger Japan” is possibly the specimen first 
described by de Selys as japponicus and may be conspecific with S. albardae Selys 
(1886), described from “Pékin”. 

In the Brussels Museum (IRSN) are three males of later date, labelled “Lansbg. [ = 
van Lansberge} Borneo” and “Bukau, N. Borneo”. 

javana Lieftinck [subsp. of Copera vittata (Selys)}, 1940c: 292, 298—303, fig. 1°, 
2—4, pl. 10 fig. 23—25, pl. 11 fig. 3, pl. 12 fig. 3—4, pl. 13 fig. 4, pl. 14 fig. 4. 
Holotype g and first described 9. S.W. Java, southcoast, Udjung Genteng, 26— 
29.111.1937, M. A. Lieftinck. — Platycnemididae. 

javica Fraser (Gynacantha), 1926: 479— 480. Holotype 9. W. Java, Buitenzorg 
{Bogor}, 22.111.1920, coll. unknown, with Fraser’s identification and “Type” written 
on envelope. — Aeshnidae. 

Syn. of Gynacantha basiguttata Selys, see Lieftinck, 1930b: 165. 

jejuna Lieftinck (Tanymecosticta), 1959a: 295— 299, fig. 34—37. Holotype 4 and 
first described 9. W. New Guinea, western extremity, Sorong, ca. 40 m, 29.VIII. 
1948, forest brook on hill slope, M. A. Lieftinck. — Isostictidae. 

jubilaris Lieftinck (Acrogomphus), 1964a: 97—102, fig. 13—16. Holotype d'. 
N.W. Borneo, Sarawak, Tabang, 7.IX.1958, M.B. 30 L, ex BISH. First described ©. 
E. Borneo, Kutai, Samarinda distr., ca. 100 km upstream, Tabang River, kali Bengen, 
ca. 100 m, 1.XI.1956, A. M. R. Wegner. — Gomphidae. 

jucunda Lieftinck (Macromia), 1955c: 270—274, fig. 25—26. Holotype g'. W. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 95 


Java, Bogor (Buitenzorg), 250 m, Botanic Garden, 2.11.1954, M. A. Lieftinck. — 
Corduliidae. 

jucundus Lieftinck (Rhipidolestes), 1948e: 9—12, fig. 6. Holotype g and first 
described 9. E. China, prov. Fu Kien, Kuatun, 2300 m, 2.VI.1938 (g') and 
7.V1.1938 (@), J. Klapperich. — Megapodagrionidae. 

juliana Lieftinck [subsp. of Rhyothemis regia (Brauer) }, 1942: 517, pl. 31 fig. 74—77, 
pl. 33 fig. 86—87. Holotype g'. N. New Guinea, Humboldt Bay area, near Lake 
Sentani, 300 m, 9.1.1933, “waterlily pond”, W. Stüber. — Libellulidae. 


karnyi Fraser (Argiolestes), 1926: 486—487. Lectotype g'. Java [err. pro Celebes], 
without further indication, with F. C. Fraser’s identification and TYPE indication. — 
Megapodagrionidae. 

Described from “three males and one female, collected by Dr. H. Karny”, an 
erroneous statement, as Karny did not collect in Celebes, the only specimen recovered 
in the Bogor museum being a g' undoubtedly originating from northern Celebes and 
caught by some Indonesian collector. 

The specimen is conspecific with Celebargiolestes cincta (Selys). — Syn. nov. 

karnyi Fraser (Procordulia), 1926: 472—473 (composite description!). Lectotype ¢ 
(head and prothorax detached, tips of superior apps. broken off). E. Java, Tengger 
Mts., 1200 m, 8.XII.1920, Hans Docters van Leeuwen, labelled ‘‘Procordulia karnyi 
d Type” in F. C. Fraser’s writing. — Corduliidae. 

In the Buitenzorg Museum were 1 G', returned by Fraser and 1 9, both from the 
Tengger Mts., the 9 dated 14.1.1921, same collector as lectotype, and labelled 
“Allotype”. A second g', from S. Sumatra, Lampong distr., Wai Lima, XI—XII. 
1921, H. H. Karny and H. C. Siebers (recorded in the original description) and 
labelled by Fraser “Procordulia javica sp. n. g' Type”, is a misidentified example of 
Zygonyx ida Selys. Fraset’s description of the § terminalia of P. karnyz applies to 
the organs of Z. ida, with which it was evidently confounded. See also Lieftinck 
(1930b: 162—164), who considered it conspecific with P. sambawana Foerster; it is, 
however, probably subspecifically distinctive. 

karnyi Laidlaw [subsp. of Caconeura verticalis (Selys)}, 1926: 230—231. Lecto- 
type &. Mentawei Is. (off W. Sumatra): Siberut I, X.1924, H. H. Kfarny}. — 
Protoneuridae. 

Now Prodasineura verticalis (Selys), see Lieftinck, 1954: 43—44. 

kimminsi Lieftinck (Lieftinckia), 1963a: 537—541, pl. 26 fig. 3, fig. 17—18. Holo- 
type g'. Solomon Is., S. Bougainville I., central part, Crown Prince Range, W. of 
Kieta, Kokorei (= Kokure), 900 m, 12.VI.1956, E. J. Ford Jr. First described 9 
(incomplete), same loc., 9.VI.1956, J. L. Gressitt. — Platycnemididae. 

kimminsi Lieftinck (Neurobasis), 1955a: 163—166, pl. 3 fig. 5. Holotype & and 
first described 9. Papua, Kwagira River, Peria Creek, 50 m, 14.VIII—6.1X.1953, 
G. M. Tate (4th Archbold Exped.). — Calopterygidae. 

krugeri Laidlaw (Drepanosticta), 1926: 228—229, fig. 2a-c. Lectotype & (headless, 
otherwise in perfect condition). Mentawei Is. (off W. Sumatra): Sipora I., 21.X. 
1924, H. H. Kfarny}, no. 95. — Platystictidae. 

One intact & from N. Pagai I. (Mentawei Is.), now also in the Leiden museum, 
according to Laidlaw (op. cit.), is a syntype (see Lieftinck, 1934d: 468— 469, pl. 10 
fig. 1—4). 


96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


kuiperi Lieftinck (Amphicnemis), 1937b: 99—101, fig. 23. Holotype 8 and first 
described ©. Billiton I. (west), Tjerutjuk, VIII.1935, F. J. Kuiper. — Coen- 
agrionidae. 


lachesis Lieftinck (Macromia), 1971b: 38—40, fig. 20, 25—26. Holotype g and 
first described 9. New Britain (Bismarck Archip.), Illugi, 8.X11.1962, J. Sedlacek 
(&, ML), and Gaulim, 160 m, 20—28.X.1962, J. Sedlacek (9, BISH). — 
Corduliidae. 

[lacteola Selys (Platycnemis?), 1863: 167— 168. Holotype g', destroyed. “Le Japon. 
Musée de Leyde, où j'ai pris le signalement trés-incomplet qu’ont vient de lire”. — 
Platycnemididae. } 

laglaizei Selys (Telebasis), 1878b (3). First described 9. N. New Guinea, Humboldt 
Bay area, Hollandia and env., I—II.1931, XII.1932, II, IV and X.1933, W. Stüber. 
See Lieftinck, 1935c: 259— 260. — Coenagrionidae. 

Now Teinobasis laglaizei (Selys). 

lairdi Lieftinck (Lieftinckia), 1963a: 534—537, 542, pl. 26 fig. 2, fig. 13—16. 
Holotype g (immature, in alcohol). Solomon Is., Guadalcanal I., Suta, 680 m, 
on ridge above Sutakiki River, 27.VI.1956, J. L. Gressitt. — Platycnemididae. 

First described Q (immature). Solomon Is., Guadalcanal I., Tapenanje, 10—23. 
XII.1953, J. D. Bradley, BM. 1954—222, Lieftinckia? sp., det. D. E. Kimmins 
(BM). 

lamberti Lieftinck (Lanthanusa), 1942: 497—499, pl. 27 fig. 42—43, 45. Holotype 
d'. Central N. New Guinea, Snow Mountain (Nassau) Range, moss-forest N.E. of 
Lake Habbema, 2800 m, 25.X.1938, L. J. Toxopeus (3rd Archbold Exped.). — 
Libellulidae. 

laminatum Lieftinck (Papuagrion), 1937a: 28, 37—39, fig. 25—26. Lectotype G' 
and first described 9. N. New Guinea, Humboldt Bay area, upper course of Korimé 
River, Mameda-Warim, ca. 250 m, about 15 km W. of Lake Sentani, 24.1X.1934, W. 
Stüber. — Coenagrionidae. 

lamprostomus Lieftinck (Argiolestes), 1949a: 49—51, fig. 26, 36. Holotype g and 
first described 9. Central N. New Guinea, Idenburg River basin, Bernhard Camp 
B, 100 m, 9.1V.1939 (g') and 6.1V.1939 (© ), L. J. Toxopeus et al. (3rd Archbold 
Exped.). — Megapodagrionidae. 

lara Krueger (Euphaea), 1898 ( & ). First described 9. E. Sumba (Lesser Sunda Is.), 
Kananggar, 700 m, V.1925, K. W. Dammerman. See Lieftinck, 1936a: 106—167, 
fig. 1. — Euphaeidae. 

Now E. lara lara Krueger. 

latericium Lieftinck (Ceriagrion), 1951c: 185—186, 193—196, fig. 3. Holotype & 
and first described 9. S.E. Sumatra, Palembang, Bukit Besar, 25.IX.1937, “dark 
rubber plantation” (g) and Palembang, 3.IX.1937, “own garden” (9), L. Coo- 
mans de Ruiter. — Coenagrionidae. 

Now C. latericium latericium Lieft. (see Asahina, 1967: 291—293, fig. 123—135). 

leonardi Lieftinck (Idiocnemis), 1958: 275—276, 282, fig. 52—54. Holotypeg. 

Tagula I. (Sudest I., Louisiade Archip.), N.-coast, Joe Landing, 0—100 m, 18— 
21.VIII.1956, L. J. Brass (Sth Archbold Exped.). — Platycnemididae. 

leopoldi Fraser (Procordulia), 1932 (& juv.). First described 2 and adult &. N. New 
Guinea, Humboldt Bay area, Cycloop Mts., 1000 m, IX—X.1932, W. Stüber. See 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 97 


Lieftinck, 1935c: 295—298, fig. 47— 49. — Corduliidae. 
leptalea Lieftinck (Tanymecosticta), 1959a: 299—300, fig. 5, 13, 38—41. Holotype 
dg. Woodlark I. (Murua I.), Kulumadau, 0—100 m, 2.X1.1956, L. J. Brass (Sth 
Archbold Exped.). — Isostictidae. Plate 1. 
lepyricollis Lieftinck (Drepanosticta), 1949a: 57—59, fig. 31, 53—57. Holotype dg. 
Central N. New Guinea, Idenburg River valley, above Bernhard Camp, 750 m, 
25.11.1939, L. J. Toxopeus (3rd Archbold Exped.). — Platystictidae. 
lethe Lieftinck (subsp. of Diplacina paula Ris), 1953d: 155, 161—164, fig. 1—2. 
Lectotype g'. Misool I. (off New Guinea), Solal, source of Gu River, ca. 50 m, 
15.X.1948, M. A. Lieftinck. — Libellulidae. 
liberata Lieftinck (Rhinocypha), 1949a: 27—29, fig. 9. Holotype g'. Solomon Is, 
Ugi I., off the northcoast of San Cristoval, 6.V.1934, no. 2513, R. A. Lever. — 
Chlorocyphidae. 
lieftincki Asahina (Ceriagrion), 1967: 297—298, 325, fig. 144149. Holotype g'. 
Philippine Is., Basilan I., Maloong, 29.IX.1932, K. Kuwasima. — Coenagrionidae. 
lieftincki Laidlaw (Coeliccia), 1932: 32—33, pl. 1 fig. 8, 14; pl. 2 fig. 13—14, pl. 
3 fig. 2. Holotype & and allotype 9 (in cop.). S. Java, prov. Banjumas, S.-coast, 
Nusa Kambangan, 18.VI.1927, F. C. Drescher. — Platycnemididae. 
See also Kimmins, 1970: 180. 
lieftincki Belle (Negomphoides), 1970: 154—158, fig. 258—264. Holotype g'. Perú, 
environs of Tingo Maria on Guayaga river, 700 m, 1962, L. Gómez Alonso. First 
described 9. Perú, dept. Huanuco, Huallaga river area, 6—700 m, Rio Rondos, 
X.1962—XI.1963, L. Gómez Alonso. — Gomphidae. 
lieftincki Watson (subsp. of Trapezostigma liberata Lieft.), 1967: 396, 400, fig. 19, 
43. Holotype g and first described 9. Central W. New Guinea, Wissel Lakes 
(Paniai), Lake Tigi, Waghete, 1740 m, 13.1.1955 (G', no. 840) and 12.1.1955 (9, 
no. 841), L. D. Brongersma et al. — Libellulidae. _ 
liesthes Lieftinck (Bironides), 1937a: 51—53, fig. 36. Holotype ¢ and first describ- 
ed 9. N. New Guinea, ca. 15 km south of Bougainville Hills, Nonno (Japu) 
hills, ca. 400 m, 12—21.11.1936, W. Stüber. — Libellulidae. 
lilacina Foerster (Trithemis), 1899a (3). First described 9. E. Sumba (Lesser Sunda 
Is.), Kananggar, 700 m, V.1925, K. W. Dammerman. See Lieftinck, 1936a: 144—145. 
— Libellulidae. 
linsleyi Lieftinck (Lestes, sg. Indolestes), 1960c: 148—151, 160, pl. 5 fig. 9—10; 
fig. 15, 26—27. Holotype G. Papua, Central Province, Brown Range, E. of Pt. 
Moresby, 8.V1.1955, J. L. Gressitt. — First described 9. S. New Guinea, Digul River 
territ., Obaa distr., Mappi Post, IV.1938, J. M. van Ravenswaay Claasen. — Lestidae. 
lombockensis Mac Lachlan (var. of Euphaea lara Krueger), 1898 (&). First described 
9. W. Flores (Lesser Sunda Is.), Wai Radjang, 140 m, 16—18.X1.1929, J. K. de 
de Jong. See Lieftinck, 1936a: 108—110, fig. 2 (name miss-spelt lombokensis). — 
Euphaeidae. 
Now E. lara lombockensis MacLachlan. 
longipennis Selys 77 Selys & Hagen (Calopteryx), 1854: 50. Lectotype 4 (immature). 
With printed label “v. Siebold, Japan”, under old drawer-label “Atrata de Selys” 
(unknown handwriting). — Calopterygidae. 
Described in some detail, yet recognized as juvenile individuals of C. atrata Selys, 
1853, originally described from China. Of these Japanese specimens the author says: 


98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


“Croyant ces exemplaires [deux & du Musée de Leyde] distincts, je les avais d’abord 
nommés C. longipennis (De Selys)”. 

Synonymous with, or a subspecies of, C. atrata Selys. 

longispina Lieftinck (Elattoneura), 1937b: 76—79, fig. 10—12. Holotype g and 
first described 9. W. Borneo, Singkawang, forest marsh near Bakuan, 17.11.1932, 
L. Coomans de Ruiter. — Protoneuridae. 

lorentzi Lieftinck (Notoneura), 1938: 91—92, fig. 24—26. Holotype J (ex alco- 
hol). Central N.W. New Guinea, Boven [Upper] Jamur, between Geelvink Bay and 
Etna Bay, 4.VIII.1903, L. F. de Beaufort & H. A. Lorentz. — First described 9. 
Berau Peninsula (Vogelkop, eastern interior), Manikiong territory, 14— 28.11.1903, 
same collectors. — Protoneuridae. 

lorquini Selys (Telebasis), 1877 (2). First described &. E. Celebes, Malili, with label: 
K 68b, VIIL.1932, R. Woltereck. See Lieftinck, 1935c: 256—257. — Coenagrionidae. 

Now Teinobasis lorquini (Selys). 

louisiadensis Lieftinck (Idiocnemis), 1958: 274—275, 283, fig. 43—51, 55. Holo- 
type d' and first described 9. Misima I. (St. Aignan, Louisiade Archip.), Sisa, north 
slopes, 350 m, 18.VII.1956, L. J. Brass (Sth Archbold Exped.). — Platycnemididae. 

luciae Lieftinck (Teinobasis), 1937a: 44—46, fig. 31. Holotype 4 and first described 
9. N. New Guinea, ca. 15 km south of Bougainville Hills, Nonno (Japu) hills, ca. 
400 m, 12.11.1936, W. Stüber. — Coenagrionidae. 

luctuosa Lieftinck (Neurothemis), 1942: 480—482. Holotype g. S. New Guinea, 
Digul River territ., Mappi Post, low country, IV.1938, J. M. van Ravenswaay 
Claasen. — Libellulidae. 

lugens Albarda 77 Selys (Vestalis), 1879: 359—360 (15—16 sep.). Lectotype g and 
first described 9. Central Sumatra, with written labels in Snelleman’s hand: “Su- 
matra Exped./Moeara laboe tot 28/11.1877” (¢) and “v. Soengei aboe naar Moeara 
laboe, 3—10/10.1877” (@), with Vestalis lugens Albarda & and 9, in Albarda’s 
writing. Ex coll. H. Albarda. — Calopterygidae. 

The application of priority of publication would demand that the specific name 
should be attributed to de Selys Longchamps, not to Albarda, whose beautifully 
illustrated account was considerably delayed in publication and did not appear in 
print until the beginning of April 1881. Identical cases are Libellago snellemanni 
(Alb.) and L. sumatrana (Alb.). However, the authorship of these three taxa was 
explicitly stated to be Albarda’s by de Selys himself (1879: 350—351, 361—362, 
397—399; and again in 1889: 445, 447, 478—479). Since de Selys’ first descriptions 
(1879) were extracted from Albarda’s manuscript, there can be no doubt that the 
names are correctly accredited to Albarda and that lectotype selections should be 
made from the material in his collection. 

lundquisti Lieftinck (Lestes sg. Indolestes), 1949a: 31—33, fig. 10—12, 97. Holo- 
type &. S.W. New Guinea, Sungei Aindua, W. of Uta (southcoast), 50—70 km 
inland, 4—9.VII.1941, E. Lundqvist et al. (Negumy Exped.). First described 9. 
S. New Guinea, Merauke, 12.111.1939, R. G. Wind (see Lieftinck, 1960c: 147—148, 
fig. 14 & 25, pl. 4 fig. 7—8, g' 2). — Lestidae. 

luteipes Lieftinck (Argiolestes), 1956b: 71, 94—97, fig. 30—32. Holotype &. Papua, 
Cape Vogel Peninsula, Menapi, up to 150 m, 9.IV.1953, G. M. Tate (4th Archbold 
Exped.). — Megapodagrionidae. 

luxatus Lieftinck (Lestes), 1932b: 492, 495—497, fig. 2. Lectotype & and first des- 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 99 


cribed 9. N. New Guinea, Humboldt Bay area, Hollandia, XI—XII.1930, W. Stü- 
ber. — Lestidae. 

Now Lestes (Indolestes) luxatus Lieft. 

lygisticercus Lieftinck (Lestes), 1932b: 492, 497, fig. 3. Holotype g'. N. New Guinea, 

Humboldt Bay area, Hollandia, 1111931, W. Stüber. First recorded 9. N. New 
Guinea, Humboldt Bay area, Njau Sanké, 15 km S. of Bougainville Mts., 15—25.XI. 
1935, W. Stüber (Lieftinck, 1949a: 30). — Lestidae. 

Now Lestes (Indolestes) lygisticercus Lieft. 


maclachlani Foerster (Anax), 1898 ( 4). First described 9. N. New Guinea, Humboldt 
Bay area, Hollandia, V.1930, W. Stiiber. See Lieftinck, 1942: 587, 605, pl. 39 fig. 
140. — Aeshnidae. 

madelenae Laidlaw (Amphicnemis), 1913 (8). First described 9. W. Borneo, Sing- 
kawang, forest-marsh near Bakuan, 7.XII.1931, L. Coomans de Ruiter. See Lieftinck, 
1940b: 376—377. — Coenagrionidae. 

magica Lieftinck (Hemicordulia), 1937b: 107—109, fig. 26. Holotype g'. Bali I, 
East Bali, Mt. Abang, 1900 m, 7.1V.1936, on mountain ridge, C. G. G. J. van 
Steenis. — Corduliidae. i 

Synonymous with H. australiae (Ramb.), see Lieftinck, 1953c: 183—184. 

magnificus Selys (Chlorogomphus), 1854: 99 (80 sep.), J Q “Sumatra”. Lectotype 
9. Muller Sumatra (print). First described g (incomplete). Muller Java (print); 
this example is the holotype & of C. hyalinus Selys, a new name first proposed for 
the § in the Secondes Additions (1869b: 202), not in the Monographie (1858), as 
stated by de Selys. — Cordulegasteridae. Plate 5 (g'). 

A syntypic ®, also from “Sumatra”, bears J. C. H. de Meijere’s label “Waar- 
schijnlijk type” (MA). This is in a much better condition than the lectotype, which 
now lacks its head, legs and abdomen. 

After publication of his “Fissilabioidea” (1929), Fraser informed me in a letter 
that he wrongly gave the ¢ as the type of C. magnificus and that the 9 from Su- 
matra should instead be designated lectotype, an opinion with which I concur. 

Type-species of Chlorogomphus Selys. 

Cf. also Chlorogomphus hyalinus Selys, in the present paper. 

malacodora Lieftinck (Lochmaeocnemis), 1949a: 100—104, fig. 103, 116, 120—124, 
131—133. Holotype g and first described 9. Central W. New Guinea, Idenburg 
River valley, above Bernhard Camp, 700 m, 11.11.1939 (g') and 25.111.1939 (2), 
L. J. Toxopeus (3rd Archbold Exped.). — .Platycnemididae. 

Type-species of Lochmaeocnemis Lieftinck (op. cit.). 

malkini Belle (Phyllocycla), 1970: 75—77, fig. 112—115. Holotype g and first 
described ©. Brazil, Maranhäo, Aldeia Yavaruhu (Aracu), 50 km E. of Canindé, 
11-25.11.1966, B. Malkin & J. C. Pinheiro. — Gomphidae. 

malleifera Lieftinck (Macromia), 1955c: 256—258, fig. 5—7. Holotype g'. E. China, 
prov. Fu Kien, Kuatun, 2300 m, 6.VII.1938, J. Klapperich. — Corduliidae. 

mariae Lieftinck (Amphicnemis), 1940b: 366, 374—375, pl. 15 fig. 1. Holotype d'. 
E. Borneo, N. Kutai, Sangkulirang distr., Pelawan Besar, V.1937, M. E. Walsh. — 
Coenagrionidae. 

mariae Lieftinck (Leptogomphus), 1948a: 249—251, fig. 10 and pl. 8. Holotype 


100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


9. E. Borneo, N. Kutai, Sangkulirang distr., Dagaunan, VI.1937, M. E. Walsh. — 
Gomphidae. 
mariae Lieftinck (Rhinocypha), 1930b: 136—138. Lectotype g'. S. Sumatra, Benkulen, 
Lake (Danu) Ranau distr., E. of Surabaja, 600 m, 28.X.1929, M. J. van Steenis- 
Kruseman & C. G. G. J. van Steenis. First described 9. S. Sumatra, near locality 
of lectotype, at Sukanegri, 31.X11.1937, L. Coomans de Ruiter. See Lieftinck, 1948a: 
270. — Chlorocyphidae. 
maroccanus Lieftinck (subsp. of Gomphus simillimus Selys), 1966c: 34—38, fig. 6. 
Holotype &, Morocco, Moyen-Atlas, El Hajeb, between Meknès and Azrou, ca. 
1050 m, 25.V.1966, M. A. Lieftinck. — Gomphidae. 
First described 9. Morocco, Moyen Atlas, Ifrane, 1600 m, 15-16.VII.1935, A. 
Ball (IRSN) (op. cit.). 
{materna Selys (Agriocnemis), 1877: 151—152. Holotype g and first described 9. 
“Sumatra. Un couple au musée de Leyde”; destroyed. — Coenagrionidae. | 
Currently considered synonymous with A. femina (Brauer), but probably subspeci- 
fically distinct. 
[melaenops Selys (Gomphus), 1854: 48. Syntype &. “Japon. (Musée de Leyde et 
Collect. Selys)”; destroyed. — Gomphidae. } 
A lectotype g' should be selected from de Selys’ collection (IRSN). 
melanostoma Lieftinck (Anaciaeschna), 1949b: 354— 356, fig. 10. Holotype & (im- 
mature). Solomon Is., Guadalcanal I., Aola, VIII.1938, unknown collector (ex NMB). 
— Aeshnidae. 
melanoxantha Lieftinck(Notoneura), 1949a: 68—70, fig. 80—81, 89— 90. Holotype 
g and allotype Q (in cop.). Central N. New Guinea, Sahuweri River valley, 
Araucaria Camp, 800 m, 21.111.1939, L. J. Toxopeus & J. Olthof (3rd Archbold 
Exped.). — Protoneuridae. 
merope Lieftinck (Diplacina), 1963c: 761—763, fig. 12—15. Holotype g'. W. New 
Guinea, western extremity, Sorong, 2.III.1959, G. F. Mees. — Libellulidae. 
[metallica Fraser (Parathemis), 1926: 470—471, fig. 1. Holotype &. “Java” (err. 
pro Celebes); lost. Described from a single erroneously labelled 4 ex Buitenzorg 
Museum (Mus. Zool. Bog.). — Libellulidae. 
Type-species of Parathemis Fraser (op. cit.) }. 
Syn. of Celebothemis delecollei Ris (see Lieftinck, 1936c: 403). 
miae Lieftinck (Libellago), 1940a: 84— 88, fig. 1. Lectotype g and allotype @ (in 
cop.). Ceylon, Central Prov., Haragama, 500 m, Mahaweliganga River, 18.IX.1938, 
M. A. Lieftinck. — Chlorocyphidae. 
Syn. of Libellago adami Fraser, 1939: 23—24, fig. la-c, & only, from the same 
locality. 
micans Lieftinck (Teinobasis), 1949a: 174—175, fig. 219. Holotype g' and first 
described 9. W. New Guinea, Berau Peninsula (Vogelkop), Sungei Tisa (south- 
coast), 9.V.1941, E. Lundqvist (Negumij Exped.). — Coenagrionidae. 
micans Lieftinck (Diplacina), 1953d: 175—177, fig. 8—9. Holotype 4 (abd.-segm. 
6—10 and apps. missing). Waigeu I. (off N. W. New Guinea), with printed label 
“Waigeu Bernstein”, and “Diplacina smaragdina Selys”, in de Selys’ handwriting. — 
Libellulidae. 
micropsitta Lieftinck (Palaiargia), 1957: 60—64, fig. 16 & pl. 4 fig. 2. Holotype 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 101 


d and first described ©. Misool I. (off W. New Guinea), env. of Fakal, 50—100 
m, 2.X.1948, M. A. Lieftinck. — Coenagrionidae. 
migratum Lieftinck [subsp. of Orthetrum pruinosum (Burm.)], 1951b: 35—37, fig. 
14. Holotype g', N. Australia, Cape York Peninsula, Coen, 10.X1.1947, H. L. 
Pottinger. — Libellulidae. 
miniata Fraser (Macromia), 1924: 450—451, pl. 25 fig. 7. Lectotype 3. S. Peninsular 
India, Coorg, Somwarpet, 1.VII.1923, F. C. Fraser. — Corduliidae. 
For particulars, see Kimmins, 1966: 204. 
miniata Lieftinck (subsp. of Palaiargia charmosyna Lieft.), 1949a: 119, fig. 158. 
Holotype & and first described 9. Central N. New Guinea, Sahuweri River basin, 
Araucaria Camp, 800 m, 30.111.1939, L. J. Toxopeus (3rd Archbold Exped.). — 
Coenagrionidae. 
minutus Belle (Cyanogomphus), 1970: 23—27, fig. 25, 42—47, pl. 3b, 4a-b. Holo- 
type d and first described 9. Surinam, Mooi Wanna, Weijne road near Albina, 
3.V1.1965, J. Belle. — Gomphidae. 
mirifica Lieftinck (Archboldargia), 1949a: 142—145, fig. 139—143, 148, 161, 171. 
Holotype 4 and first described @. West Central New Guinea, Wissel Lakes group, 
Enarotali, E.-shore of Lake Paniai, 1740 m, 27.IX.1939 ({) and Arabu Bivouac, 
10 mi. N.E. of Lake Paniai, ca. 1800 m, 22.X.1939 (2), H. Boschma (Le Roux 
Exped). — Coenagrionidae. Plate 2. 
Type-species of Archboldargia Lieftinck (op. cit.). 
misoolensis Lieftinck (subsp. of Neurobasis australis Selys), 1955a: 161—163, pl. 3 fig. 
4. Holotype g' and first-described 9. Misool I. (off W. New Guinea), Wartama 
River near Fakal, ca. 40 m, 2.X.1948, M. A. Lieftinck. — Calopterygidae. 
mnemosyne Lieftinck (Macromia), 1935a: 190—191, fig. 7. Lectotype g'. W. Bor- 
neo, Singkawang, swampy forest near Bakuan, low country, “at dusk”, 22.1.1934, 
L. Coomans de Ruiter. — Corduliidae. 
modesta Belle (Phyllocycla), 1970: 70—74, fig. 106—111, pl. 10b, 11a, 12b. Holotype 
d'. Surinam, Surinam river (Afobakka), 28.X11.1963, J. Belle. First described 9, 
Surinam, Saramacca, right Toekoemoetoe (Tafelberg creek), 6.X.1944, L. Schmidt. 
— Gomphidae. 
moerens Lieftinck (Huonia), 1963c: 772—774, fig. 30— 32. Holotype g'. New Britain 
(Bismarck Archip.), N.-slope of Nakanai Mts, 150 m, 22.VII.1956, E. J. Ford 
Jr. — Libellulidae. 
moluccana Lieftinck (Anaciaeschna), 1930a: 316—318, fig. 4. Holotype 3. Buru I. 
(S. Moluccas), Wai Eno, beginning of IV.1921, L. J. Toxopeus. — Aeshnidae. 
moluccana Lieftinck (Drepanosticta), 1938: 82—83. Lectotype g' and first described 
2. Buru I. (S. Moluccas), Nal’besi, 26.1V.1921 (4 ) and Buru I., Station 2, 1.1922 
(2), L. J. Toxopeus. — Platystictidae. 
Nom. nov. pro Drepanosticta auriculata: Ris (1929: 141—142, g' 9 Buru I.), nec 
Platysticta auriculata Selys, 3 N.W. New Guinea, a congeneric species. 
moluccana Lieftinck [subsp. of Nesoxenia mysis (Selys) }, 1942: 461, 464, (not pl. 24, 
fig. 13—14!). Lectotype 4 and first described 9. Noord Halmaheira [H.A] 
Bernstein [N. Moluccas, Halmahera I.]; © with additional pin-label “Agr. mysis 
Selys, ann. Dresden”, in de Selys’ writing. — Libellulidae. 
This is a composite taxon, examples from the southern Moluccas and Kei Islands 


102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


(figured in the original description) belonging to a different and still undescribed 
subspecies of N. myszs Selys. 
monoceros Lieftinck (Drepanosticta), 1965a: 185—186, fig. 12—15. Holotype ¢ 
and first described 9. E. Borneo, Kutai, Bengen River, Tabang, 125 m, 28.IX.1956 
(3) and 6.X.1956 (2), A.M.R. Wegner. — Platystictidae. 
montaguei Campion (Synthemis) 1921 (&). First described 9. New Caledonia, 
N.E.-side, Pouébo, 200—250 m, 19-21.1.1964. R. Straatman. See Lieftinck (1971b: 
51—52, fig. 36). — Corduliidae. 
monticola Lieftinck (subsp. of Tramea eurybia Selys), 1942: 537—538, pl. 38 fig. 124. 
Holotype & and first described 9. Central W. New Guinea, Baliem River valley, 
Baliem Camp, 1600 m, 16.X11.1938 (4) and 30.X1.1938 (9), L. J. Toxopeus 
(3rd Archbold Exped.). — Libellulidae. 
Now Trapezostigma eurybia monticola (Lieft.). 
montivagans Lieftinck (Anaciaeschna), 1932c: 251—253, fig. 3. Holotype g and 
first described 9. W. Java, G. Gedeh-complex, Puntjak pass, Telaga Saat, 1450 m, 
111.1930 (6, bred from larva) and 15.X11.1929 (© ad.), M.A. Lieftinck. — 
Aeshnidae. 
mungo Needham (Cacus), 1940 (&). First described 9. Surinam, upper course of 
Coropina Creek, near Zanderij I (Airport), 20.1.1957, D. C. Geijskes. See Geijskes, 
1964: 37—40, fig. 15—16. — Gomphidae. 
Currently known as Cacoides mungo (Needham). 
murcia Lieftinck (Idionyx), 1971b: 6, 12—14, fig. 3—4. Holotype & and first descri- 
bed 9. Central Sumbawa I. (Lesser Sunda Is.), Semongkat-atas, 21.1V.1961 (&) 
and Semongkat, 300 m, 27.1V.1961 (9 ), P. Jauffret & R. Pujol, ex coll. A. Heymer. 
— Corduliidae. 
mutans Lieftinck [subsp. of Nesoxenia mysis (Selys) }, 1942: 462, 465—466, pl. 24 
fig. 18, pl. 41 fig. 163. Holotype § and first described 9. W. New Guinea, Mac- 
Cluer Gulf, northcoast of Bombarai Peninsula, Babo, 22-23.VIII.1941, E. Lundqvist 
(Negumij Exped.). — Libellulidae. 
Probably a distinct species. 
mutata Lieftinck (Oligoaeschna), 1940b: 381—383, 384, fig. 18. Holotype g'. E. Bor- 
neo, Kutai, Samarinda, 1.1939, native coll, M.E. Walsh ded. First described 9. 
E. Borneo, Kutai, Sangkulirang distr., Kariorang, 21.11.1937, J. W. Q. de Quarles. — 
Aeshnidae. 
myzomela Lieftinck (Palaiargia), 1957: 51—53, fig. 11—12, pl. 2 fig. 1. Holotype 
9. West Central New Guinea, Wissel Lakes area, Kamo Valley, Ituda, 1500 m, 
12.VIII.1955, J. L. Gressitt. — Coenagrionidae. 
naias Lieftinck (Libellago), 1932a: 4—9. Holotype g' and first described 9. W. 
Flores I. (Lesser Sunda Is.), Wai Radjang, along small streams, 16-18.X1.1929, J.K. 
de Jong. — Chlorocyphidae. 
neotropica Belle (Phyllocycla), 1970: 97—100, fig. 147, 150—153, pl. 13b. Holotype 
d'. Surinam, Brownsweg (road near Makambi creek), 11.VI.1961, D. C. Geijskes. 
— Gomphidae. 
nereis Lieftinck [subsp. of Agrionoptera insignis (Ramb.)}, 1948b: 292—294, fig. 7. 
Holotype g and first described 9. Engano I. (off S.W. Sumatra), Meok, 24.V.1936, 
J. K. de Jong et al. — Libellulidae. 
nereis Lieftinck {subsp. of Coeliccia membranipes (Ramb.)], 1948b: 287—288, fig. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 103 


3—4. Holotype g and first described 9. Engano I. (off S. W. Sumatra), Kiojoh, 
2.VI.1936 (g') and 14.VI.1936 (2), J. K. de Jong et al. — Platycnemididae. 

[nigra Selys in Selys & Hagen [Sapho (mnais)}, 1854: 65. Holotype &. “Japon”; 
destroyed (not in ML). — Calopterygidae. 

An enigmatic specimen of unknown identity, figuring under the above names, 

though classified in a note sub Mais strigata Hagen. } 

nigra Foerster 77 Laidlaw (var. of Disparoneura verticalis humeralis Selys), 1907: 
12—13. Lectotype &. Malay Peninsula, labelled by Förster: Camp Jor, Wasserscheide 
Pahang-Perak, 2000 Fuss, Albert Grubauer/Disparoneura verticalis Rasse humeralis 
Selys var. nigra F.; and: figured by C. H. Kennedy Aug.—Sept. 1924 (print). — 
Protoneuridae. 

A synonym of Prodasineura verticalis (Selys). 

nigriventris Lieftinck (Idiocnemis), 1937a: 8—9, fig. 2. Lectotype 4 and first descri- 
bed 9. N. New Guinea, Humboldt Bay area, Hollandia, XI—XII.1931 (&) and 
1.1932 (9), W. Stüber. — Platycnemididae. 

nigrofasciata Lieftinck (Notoneura), 1932b: 523, 537538, fig. 32—34. Lectotype 
3 and allotype 9 (in cop.). N. New Guinea, Humboldt Bay area, Hollandia, 40 
km S., 300 m, XI—XII.1931, W. Stüber. — Protoneuridae. 

nigrofasciatum Lieftinck (Pseudagrion), 1934a: 6—8, fig. 1—2. Lectotype 4. Java 
Orient., Hekmleijer} (written label). — Coenagrionidae. 

nubecula Lieftinck (Rhinocypha), 1948a: 266—270, fig. 16. Lectotype & and first 
described 9. N. Sumatra, Atjeh, Laut Tawar (Lake Takingeun), north shore, 1210 
m, 19.VII.1941, F. C. van Wagensveld. — Chlorocyphidae. 

nymphaeae Lieftinck (Aethriamanta), 1949a: 233—234, fig. 287. Holotype & and 
first described 9 . N. New Guinea, Humboldt Bay area, Lake Sentani, W. of Hollandia, 
waterlily ponds, 13.1.1933 (3) and 18.1.1933 (9), W. Stüber. — Libellulidae. 


obiensis Lieftinck (Argiolestes), 1956b: 66, 75—77, fig. 3—4. Holotype 8 and first 
described 9. Ohi I. (N. Moluccas), N.W. Obi, Obi lake, kali Telaga, 100—160 m, 
6.VIII.1953, A. M. R. Wegner et al. — Megapodagrionidae. 

obiensis Lieftinck (Palaiargia), 1957: 53—54, fig. 13. Holotype 4 (immature). 
Obi I. (N. Moluccas), N.W. coast, Laiwui, 1.X.1953, A. M. R. Wegner et al. — 
Coenagrionidae. 

obliterata Lieftinck (Idiocnemis), 1932b: 499—501, fig. 5. Lectotype g' and first 
described 9, N. New Guinea, Humboldt Bay area, Hollandia, IX.1930, W. Stüber. 
— Platycnemididae. 

obscura Fraser (Neurothemis), 1926: 472. Lectotype 4. Karimondjawa Is. (Java 
Sea), X.1920, H. C. Delsman. — Libellulidae. 

Currently placed as a subspecies of Newrothemis terminata Ris. 

obscurum Lieftinck [subsp. of Pseudagrion pilidorsum (Brauer) |, 1936a: 128, 131— 
132, fig. 12. Lectotype g'. Nias I, Amadraja, 10.IX.1931, J. C. van der Meer 
Mohr. — Coenagrionidae. 

obtusum Albarda (Zyxomma), 1881: 1, pl. 1 fig. 1—2. Lectotype 4 and first 
described @. Central Sumatra, labelled: ‘‘Soepayang, 24/4 77” (&) and “tot 24/4 
77° (2) Sumatra Exp., both labelled: Zyxomma obtusum Hag., in H. Albarda’s 
writing, ex coll. H. Albarda. — Libellulidae. 

oceanis Lieftinck (subsp. of Rhinocypha angusta Selys), 1947: 220—224, fig. 4—5, 


104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


pl. 2 fig. 3. Holotype J and first described 9. Engano I. (off S.W. Sumatra), 
Buah-buah, ult. V—VII.1936, J. K. de Jong et al. — Chlorocyphidae. 

oceanis Lieftinck [subsp. of Neurothemis ramburi (Brauer) }, 1948b: 297— 298. Holo- 
type g and first described 9. Engano I. (off S.W. Sumatra), Buah-buah, 5.VI. 
1936, J. K. de Jong et al. — Libellulidae. 

olthofi Lieftinck (Teinobasis), 1949a: 169—171, fig. 218. Holotype g'. N. Central 
New Guinea, Idenburg River, Bernhard Camp, 50 m, 12.1V.1939, J. Olthof. — 
Coenagrionidae. 

olympica Lieftinck (Hemicordulia), 1942: 549—555, pl. 36 fig. 114—115, pl. 37 
fig. 116—117. Holotype ¢ and first described 9. Central N. New Guinea, Snow 
Mountain (Nassau) Range, Lake Habbema, 3225 m, 21.VIII.1938, L. J. Toxopeus 
(3rd Archbold Exped.). — Corduliidae. 

oppositum Lieftinck (Papuagrion), 1949a: 146—148, fig. 190, 206. Holotype & 
and first described 9. N. New Guinea, Humboldt Bay area, southern Bewani 
Hills, 300 m, Parfi, 20.V1.1939, W. Stüber. — Coenagrionidae. 

optata Hagen #7 Selys (Argia), 1865: 390 (18 sep.). Holotype g. Obi I. (N. Mo- 
luccas), labelled “Ins. Obi, Bernstein” and “Optata Hag.” in H. A. Hagen’s hand- 
writing. — Coenagrionidae. 

First described 9. Obi I., northwest coast, Laiwui, 18.X.1953, A. M. R. Wegner 
(see Lieftinck, 1957: 58—59). 
Now Palaiargia optata (Hagen zn Selys). 

oreadum Lieftinck (Oreagrion), 1949a: 206—211, fig. 249—251, 260—263, 264— 
266, 269—270. Holotype g' and first described @. Central W. New Guinea, Snow 
Mountains (Nassau) Range, Lake Habbema, 3250 m, 1.VIII.1938, L. J. Toxopeus 
(3rd Archbold Exped.). — Coenagrionidae. Plate 3. 

oreophila Lieftinck (Huonia), 1935c: 271, 274—277, fig. 33 (not 35), 36B, 37. 
Holotype g and first described ©. N. New Guinea, Humboldt Bay area, southern 
Cycloop Mts., 1000—1200 m, VIII.1932, W. Stiiber. — Libellulidae. 

othello Lieftinck (Agyrtacantha), 1942: 571—572, pl. 38 fig. 129—131. Holotype 
9. Central N. New Guinea, Idenburg River, Bernhard Camp, 50 m, 13.1V.1939, 
L. J. Toxopeus (3rd Archbold Exped.). — Aeshnidae. 


palawana Lieftinck (Coeliccia), 1940b: 358—359, fig. 13. Holotype g'. Philippine 
Is.: Palawan I., Alfonso XIII, 6.111.1935, K. Kuwasima. — Platycnemididae. 

palawana Lieftinck {subsp. of Copera vittata (Selys) }, 1940c: 290, 303—304, fig. le, 
pl. 10 fig. 20—22, pl. 11 fig. 8, pl. 12 fig. 9, pl. 13 fig. 7, pl. 14 fig. 7. Holotype 
3 and first described 9. Philippine Is.: Palawan I., Alfonso XIII, 6.V.1935, K. 
Kuwasima, — Platycnemididae. 

pallidifrons Ris (Rhinocypha), 1927: 9—10, fig. 3. Holotype 9. W. Central Suma- 
tra, Padang Highlands, Ophir distr., Tanangtalu, 1000 m, V.1915, E. Jacobson. 
Ex SMF no. 1389. — Chlorocyphidae. 

pan Laidlaw (Drepanosticta), 1931 (&). First described 9 (juv.). Malay Peninsula, 
Perak, Cameron’s Highlands, Jor River, 600 m, 13.111.1963, M. A. Lieftinck. See 
Lieftinck, 1965a: 174—176 (& 2). — Platystictidae. 

pandanicola Lieftinck (Amphicnemis), 1953f: 388—392, fig. 2. Holotype { and 
first described Q. S. Borneo, Sampit distr., 0—50 m, near Sampit, about 50 km 
inland, 20.VII.1953, M. A. Lieftinck. — Coenagrionidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 105 


pandanicolum Lieftinck (Papuagrion), 1949a: 153—155, fig. 188—189, 195. Holo- 
type d and first described 9. Central W. New Guinea, Snow Mountain (Nas- 
sau) Range, Moss-forest N. of Lake Habbema, 2700 m, 23.VIII.1938 (g') and 
2850 m, 28.VIII.1938 (2), L. J. Toxopeus (3rd Archbold Exped.). — Coenagrioni- 
dae. 

panybeus Hagen (Anax), 1867 (&). First described 9 . N. Celebes, Tondano, VII.1933, 
C. van Braekel. See Lieftinck, 1942: 597—599. — Aeshnidae. 

papuensis Lieftinck [subsp. of Neurothemis ramburi (Brauer)], 1942: 482—484. 
Holotype g' and first described 9. N. New Guinea, Humboldt Bay area, S. Cy- 
cloop Mts., near Lake Sentani, 150 m, 18.1V.1939, J. Olthof (&), and Hollandia, 
11.1931, W. Stüber (9). — Libellulidae. 

papuensis Lieftinck (subsp. of Tetrathemis irregularis Brauer), 1942: 451—453, pl. 
23 fig. 7, 11. Lectotype 4 and first described 9. N. New Guinea, Humboldt Bay 
area, Hollandia, XI—XII.1930 (&) and IV—VI.1931 (9), W. Stüber. — Libel- 
lulidae. 

paruensis Geijskes (Micrathyria), 1963: 74—78, fig. 16. Holotype @ and first 
described 9. N. Brazil, affluent creek of Upper Paru River, near Apisiké, 19.IV. 
1952 (g') and 20.1V.1952 (2), D. C. Geijskes. — Libellulidae. 

pasiphae Lieftinck (Gynacantha), 1948c: 420—421, 424, 426—427, fig. 1. Holotype 
d. N. Halmahera I. (N. Moluccas), labelled “Halmahera [VII—VIII.1861, H. 
A.} Bernstein”, and “Gynacantha Maclachlani g'” in R. Martin's handwriting. 
First described 9. N. Halmahera I., Tobelo, VI—VII. 1931, M. J. van Diejen. — 
Aeshnidae. 

pavo Lieftinck (subsp. of Neurobasis kaupi Brauer), 1955a: 158—160, pl. 3 fig. 2. 
Holotype g and first described 9. S. Celebes, Mt. Lompobatanig (Pic of Bon- 
thain), Borong Rapoa, 700 m, 21.VIII.1949, A. Diakonoff. — Calopterygidae. 

pectitus Lieftinck (Argiolestes), 1949a: 53—54, fig. 28, 37; 110—116, fig. 47, 
53—63. Holotype g and first described 9. Central N. New Guinea, Idenburg 
River basin, Bivouac 8 km S.W. of Bernhard Camp, 1150 m, 6.1.1939 (g'), and 
Rattan Camp, 1150 m. 9.11.1939 (2), L. J. Toxopeus et al. (3rd Archbold 
Exped.). — Megapodagrionidae. 

pelecotomum Lieftinck (Pseudagrion), 1932b: 567, 577-578, fig. 57, 59. Lectotype 
d. N. New Guinea, Humboldt Bay area, Lake Sentani, I—II.1931, W. Stüber. 
First recorded 9. N. Central New Guinea, Idenburg River, Bernhard Camp, 
50 m, 12—18.X.1938, J. Olthof (4 9 in cop.). See Lieftinck, 1949a. — Coen- 
agrionidae. 

penelope Ris (Gynacantha), 1915b (3). First. described 9. Central Celebes, Luwu 
distr., Palopo, 15.111.1941, “at house a.m.”, L. L. A. Maurenbrecher. See Lieftinck, 
1948c: 418—419. — Aeshnidae. 

perameles Lieftinck (Papuagrion), 1949a: 159—160, fig. 201. Holotype 9. S. New 
Guinea, Lorentz River, Sabang, 5.VII.1907 (Zuid N. Guinea Exped.), determined 
by H. W. van der Weele as Argiolestes spec. — Coenagrionidae. 

First described by Lieftinck (1935c: 247) sub P. magnanimum (Selys), but later 

recognized as a distinct species. 

perfuscatum Lieftinck (Pseudagrion), 1937b: 94—97, fig. 21. Holotype g and first 
described 9. W. Borneo, Singkawang, Bengkajang, 5—16.X1.1931, L. Coomans de 
Ruiter. — Coenagrionidae. 


106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


perimecosoma Lieftinck (Palaiargia), 1957: 44—50, fig. 1—2, 4—10 & pl. 3 fig. 1. 
Holotype g and first described 9. Halmahera I. (N. Moluccas), central Halma- 
hera, Mumar River, 200—300 m, 25.IX.1951, Sundanese collectors. — Coen- 
agrionidae. 

perplexus Lieftinck (subsp. of Onychogomphus geometricus Selys), 1935b: 20—21, 
fig. 2. Holotype 4 (ex alcohol). N.E. Sumatra, Eastcoast prov., Deli, labelled 
“Sumatra S.O.K. Medan, 20 m, 1917, J. B. Corporaal”. — Gomphidae. 

The specimen now lacks its terminal abdominal segments. 

persephone Lieftinck (Diplacina), 1933d: 42, 47—49, fig. 28. Lectotype g' and first 
described 9. N. New Guinea, Humboldt Bay area, Hollandia, IV—VI.1931 (4), 
and same area, Nettar, XII.1932 (2), W. Stüber. — Libellulidae. 

pertinax Lieftinck (Lestes), 1932b: 493—495, fig. 1. Lectotype J and allotype 9 
(in cop.). N. New Guinea, Humboldt Bay area, Hollandia, III.1931, W. Stüber. — 
Lestidae. 

pesechem Lieftinck (Papuagrion), 1949a: 151—152, fig. 178—181, 193. Holotype 
d and first described 9. Central N. New Guinea, Sahuweri River Valley, Sigi 
Camp, 1350 m, 19.11.1939, L. J. Toxopeus (3rd Archbold Exped.). — Coenagrion- 
idae. 

Now P. pesechem pesechem Lieft. 

phaeomeria Lieftinck (Gynacantha), 1960a: 253—254, fig. 13a, b. Holotype {. 
Thailand-Tenasserim frontier area, Kwae Noi River valley, S. of Three Pagodas 
Pass, Niki, 150 m, 30.V.1946, J. E. Jonkers. — Aeshnidae. 

phaeoneura Lieftinck (Tramea), 1953e: 268—269. Holotype g and first described 
9. N. Borneo, Dent Province, Mt. Marapok, collector G. (printed label). The more 
precise locality is the one given for Linaeschna polli Martin, on. p. 107 of this paper. 
— Libellulidae. 

Now Trapezostigma phaeoneura (Lieftinck). 

phalantus Lieftinck (Macrogomphus), 1935a: 201—203, fig. 10—12. Holotype ¢ 
and first described 9. W. Borneo, Singkawang, swampy forest near Bakuan, low 
country, 15.IX.1932, L. Coomans de Ruiter. Gomphidae. 

phantasma Lieftinck (Rhinocypha), 1935a: 175—176. Lectotype g and first 
described ©. Buton I. (off S.E. Celebes), Dwaalbaai, 19.1.1917, native coll, ex 
coll. J. Lindemans & D. C. Geijskes. — Chlorocyphidae. 

phasma Lieftinck (Selysioneura), 1932b: 542, 547—549, fig. 42—43. Lectotype 4 
and first described 9. N. New Guinea, Humboldt Bay area, Cycloop Mts., ca. 
500 m, IV—VI.1931, W. Stüber. — Isostictidae. 

philippa Ris (Idionyx), 1912c (2 Mindoro). First described ¢. Philippine Is., Min- 
danao I., Misamis Or., Bal-ason, 2.1V.1960, H. Torrevillas; and & 9, Leyte I, S. 
Leyte, Anahawan, Mainit Spring, 31.V.1970, C. Plateros. See Lieftinck (1971b: 
7—11, fig. 1—2). — Corduliidae. 

phoenicura Ris (Planiplax), 1912b (3). First described 9. Surinam, Paramaribo, pond 
on shell ridge of Charlesburg, 6.VI.1957 (3 2), J. Belle. See Geijskes, 1964: 41— 
42, fig. 20. — Libellulidae. 

phoenissa Ris (Caconeura), 1929: 143—144, fig. 2. Holotype g' (indicated as Type 
by Ris). Buru I. (S. Moluccas), Nal Besi, 26.IV.1921, L. J. Toxopeus. — Proto- 
neuridae. 

Currently placed in Notoneura Tillyard. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 107 


phryne Lieftinck (subsp. of Diplacina phoebe Ris), 1953d: 158, 182, fig. 9. Holo- 
type g'. Ambon I. (S. Moluccas), 27.VII.1948, M. A. Lieftinck. First described Q. 
Ambon I., Sungei Waitumu, ca. 30 m, 23.IV.1941, Sundanese collector. — Libel- 
lulidae. 
pijpersi Belle (Progomphus), 1966a: 11—14, fig. 23—28. Holotype (in frag- 
ments) and first described 9. Surinam, Wilhelmina Mountain Range (Camp 3), 
5.VIII.1963, H. Pijpers. — Gomphidae. 
plagiatus Lieftinck (Burmagomphus), 1964b: 16, 29—33, fig. 36—41. Holotype gd. 
Malay Peninsula, Johore Bahru, Kp. Tinggi Mawai Rd., Sungai Mupor, 17.IV. 
1963, J. I. Furtado. Immature 9, from same loc. and date, M. A. Lieftinck. — 
Gomphidae. 
plagioxantha Lieftinck (Notoneura), 1932b: 523, 527, 538—540, fig. 3537. Lecto- 
type J and first described 9. N. New Guinea, Humboldt Bay area, Hollandia, 
111.1931, W. Stüber. — Protoneuridae. 
platystyla Lieftinck (Amphicnemis), 1953e: 246—247, fig. 6a—b. Holotype {. 
S.E. Borneo, Kandangan distr., Ampah, 0—20 m, 29.IV.1948, Liem Swie Liong. 
— Coenagrionidae. 
platyura Lieftinck (Oligoaeschna), 1940b: 378—380, 383, fig. 16—17. Holotype dg 
and first described 9. E. Borneo, N. Kutai, Sangkulirang distr., Pelawan Besar, 
V.1937 (8) and Batau Besi, V.1937 (2), M. E. Walsh. — Aeshnidae. 
podolestoides Laidlaw (Devadatta), 1934: 101—103, fig. 1. Holotype & and first 
described 9. W. Borneo, Singkawang, Gunung Poteng, 550 m, 21.VIII.1932, 
L. Coomans de Ruiter. — Amphipterygidae. 
Now D. podolestoides podolestoides Laidlaw. 
polli Martin (Linaeschna), 1909a: 136—137, fig. 133—134, pl. III fig. 9. Holotype 
d'. {N.W. Borneo, Sabah-Sarawak boundary, Brunei Bay area}, with printed label 
“N. Borneo, Dent Province, Mt. Marapok, coll. G.”, and labelled by R. Martin 
“Linaeschna Polli m.”, and “Type unique”. — Aeshnidae. 
Type-species of Linaeschna Martin (op. cit). 
pollux Lieftinck (Onychogomphus), 1941b: 247—253, pl. 13 fig. 7, pl. 14 fig. 4—6, 
pl. 15 fig. 1—4. Holotype &. S. Sumatra, Lampong distr., Wai Tebu (river) near 
Talang Padang, ca. 400 m, 29.III.1940, M. A. Lieftinck. — Gomphidae. 
polyhymnia Lieftinck (Macromia), 1929a: 66, 97—99, fig. 18a-c. Holotype J.N. W. 
Sumatra, Padangsche Bovenlanden [Padang Highlands}, Fort de Kock, 920 m, 
VIII.1924, E. Jacobson. — Corduliidae. 
pontica Selys (race of Libellula fulva Muell.), 1887: 12. Lectotype & and first described 
9. Iran, both labelled “Mesopotamia Malatia”, and on the reverse side “L. fulva 
ras australis’ (G') and “L. fulva Mull. v. pontica 9” (2) in H. Albarda’s writing, 
coll. Albarda acq. 1892 (print). — Libellulidae. 
praecellens Lieftinck (Lestes), 1937b: 59—62, fig. 2—3. Holotype 4 and first 
described 9. S. Java, Tjidamar near Tjidaun, ca. 100 m, forest pool near Sempur- 
tjondong, 5.XI.1935, M. Bartels Jr. — Lestidae. 
praetermissum Lieftinck (Ceriagrion), 1929b: 115—117, fig. 9. Holotype 4 and 
first described ©. Java, M. C. Piepers. — Coenagrionidae. 
See Asahina (1967: 308—310, fig. 189—195). 
praevia St. Quentin (Gomphoides), 1967 (3). First described ©. Brazil, Santa Catarina 


108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


(Nova Teutonia), 12.11.1949, coll. unknown. See Belle, 1970: 124—126, fig. 199, 
200. — Gomphidae. 

praevius Lieftinck (Lestes), 1940b: 344—347, fig. 3. Holotype & and first described 
©. E. Borneo, N. Kutai, Sangkulirang distr., Maluwi, IV.1937, M. E. Walsh. 
— Lestidae. 

propinqua Lieftinck (subsp. of Tramea euryale Selys), 1942: 539—541, pl. 33 fig. 
90—91, pl. 34 fig. 98. Holotype g' and allotype 9 (in cop.). N. New Guinea, 
Humboldt Bay area, near Lake Sentani, 300 m, 11.1933, W. Stüber. — Libellulidae. 

Currently classified as a subspecies of Trapezostigma transmarina (Brauer). 
prothoracale Lieftinck (Papuagrion), 1935c: 238, 242—245, fig. 16—18. Holotype 
d and first described. 9. N. New Guinea, Humboldt Bay area, Agaffo, near Lake 
Sentani, sago forest swamp, 27.XI1.1932, W. Stüber. — Coenagrionidae. 
prothoracica Selys (Telebasis), 1877 (3). First described 9. “Waigeoe Bernstein” 
(print) [Waigeu I., off N. W. New Guinea, III-V.1863]}. See Lieftinck, 1938: 
116—117. — Coenagrionidae. 
Now Teinobasis prothoracica (Selys). 

pruinescens Lieftinck (Idiocnemis), 1949a: 96— 100, fig. 105, 134. Holotype g and 
allotype ® (in cop.). Papua [S.E. New Guinea], Central Prov, Haveri, 700 m, 
13.VIII.1893, Lamberto Loria. Ex Museo Civico di Stor. Nat, Genova. — 
Platycnemididae. 

pruinosa Selys (Mnais), 1853: 20—21. Lectotype & (adult). With printed label “v. 
Siebold, Japan”, a green museum-label “cotype’’, and a white museum’s identification 
label “Mnais pruinosa Sel., det. Selys’. With additional label in Dr. S. Asahina’s 
writing: “Mnais strigata & f. pruinosa Selys, det. Asahina 1968. This is one of the 
original series of pruinosa”. — Calopterygidae. 

Type-species of Mnais Selys. 

puella Lieftinck (Brachygonia), 1937b: 105—107, fig. 25. Holotype g and first 
described ®. W. Billiton I., Seru, 2.1V.1936, F. J. Kuiper. — Libellulidae. 

pugnax Lieftinck (Anax), 1942: 583, 587, 604— 606, pl. 38 fig. 134, pl. 39 fig. 138, 
141, pl. 41 fig. 156. Holotype g and first described 9. Central N. New Guinea, 
Sahuweri River basin, Araucaria Camp, 800 m, 21.VIII.1939 (g') and 24.111.1939 
(2), L. J. Toxopeus (3rd Archbold Exped.). — Aeshnidae. 

[pulverulans Selys (Race? de Agriocnemis incisa Hagen) [= A. femina (Brauer) }, 
1877: 150—151. Lectotype g', destroyed. “Celebes. Mus. de Leyde et coll. Selys”. 
Not in coll. Selys (IRSN). — Coenagrionidae. 

Currently considered synonymous with A. femina (Brauer), but probably subspeci- 
fically distinct. }. 

punctata Selys (Petalia), 1854 (3). First described ©. Chili, “Felder Chile” (written, 
white disk), and two drawer labels “Puncta Hagen” and “Petalia Hagen”; with 
additional red Allotype label by M. A. L. — Neopetaliidae. Plate 7. 

See de Selys, 1869b: 204. 
Now Neopetalia punctata (Selys). 
Type-species of Petalia Selys (op. cit., nom. praeocc.). 

pusilla Lieftinck (subsp. of Rhinocypha pagenstecheri Foerster), 1953c: 136—137. 
Holotype g', Damar I. (Lesser Sunda Is.), labelled „Insel Dammer’’, collector un- 
known, ex coll. F. Förster. — Chlorocyphidae. 

pyroprocta Lieftinck (Notoneura), 1960b: 122—126, fig. 7—12. Holotype g' and 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 109 


first described ©. Misool I. (off W. New Guinea), environs of Fakal, 0—50 m, 
30.IX.1948, M. A. Lieftinck. — Protoneuridae. 


quadristigma Lieftinck (Prodasineura), 1951a: 80—83, fig. 3, 4, 7, 8, 14—16. 
Holotype g' and first described 9. S. E. Borneo, Kandangan distr., Ampah, 0—20 
m, IV-V.1948, Liem Swie Liong. — Protoneuridae. 


raineyi Williamson (Telagrion), 1915 (dg). First described 9. Trinidad, Brasso, 
8.11.1932, G. Belmontes. See Geijskes, 1932: 248— 249. — Coenagrionidae. 

ranatra Lieftinck (Selysioneura), 1949a: 78—80, fig. 82—83. Holotype ¢ and first 
described 9. Central W. New Guinea, Sahuweri River valley, Araucaria Camp, 800 
m, 2.1V.1939 (g') and 21111939 (2), L. J. Toxopeus & J. Olthof (3rd Archbold 
Exped.). — Isostictidae. 

ranauense Schmidt [subsp. of Pseudagrion pruinosum (Burm.) }, 1934: 347—348, fig. 
47b (8 $ Sumatra). Lectotype g'. S. W. Sumatra, Benkulen, Lake Ranau, 1.1929 
(Deutsch. limnol. Sunda-Exped. 1928—29). — Coenagrionidae. 

Selected by E. Schmidt and in 1935 presented to author. A syntype Z is in the 

Brit. Mus. (N.H.) collection, see Kimmins, 1970: 191. 

rangifera Lieftinck (Notoneura), 1949a: 70—71, fig. 77—79, 91—92. Holotype 
d' and allotype ® (in cop.). S. New Guinea, Digul River territ., Ederat near Aja 
Maru, 11.1939, J. M. van Ravenswaay Claasen. — Protoneuridae. 

rangifera Lieftinck (Selysioneura), 1959a: 290—291, 294, fig. 6, 25—27. Holotype 
d'. Normanby I. (d'Entrecasteaux Archip.), Mt. Pabinama, 3 mi. E.N.E. of Cape 
Prevost, 820 m, 4.V.1956, L. J. Brass (Sth Archbold Exped.) — Isostictidae. 

rappardi Lieftinck (Onychogomphus), 1937b: 110—113, fig. 27—28. Holotype &. 
S. W. Sumatra, Benkulen prov, Redjang distr., Pagar Gunung, 550 m, “open stream”, 
12.XI1.1936, F. W. Rappard. — Gomphidae. 

raymondi Lieftinck (Phyllothemis), 1950a: 643—645, fig. 7—9. Holotype 4. N. E. 
Sumatra, Deli, Serbalawan, Dolok Ilir Estate, 200 m, 17.V.1948, R. Straatman. — 
Libellulidae. 

rectangulare Lieftinck (Papuagrion), 1937a: 27, 31—33, fig. 21. Holotype 4 and 
first described 9 . N. New Guinea, Humboldt Bay area, southern slope of Mt. Cycloop, 
1000 m, 23.IX.1934 ( g') and 17-23.1X.1935 (9), W. Stüber. — Coenagrionidae. 

resecta Lieftinck (Coeliccia), 1953e: 236—239, fig. 3. Holotype <. S.E. Borneo, 
Kutai, Balikpapan, Sungai Mentawir, sea-level, 6.X.1950, A. M. R. Wegner. — 
Platycnemididae. 

reticulata Fraser (Gynacantha), 1926: 478—479. Holotype 8. Kei Is., labelled “Kei 
Eiland, Gg. Daab, no. 142, 14.1V.1922, H. C. Siebers’’, with Fraser’s identification 
and “Type” written on envelope. — Aeshnidae. 

Syn. of Agyrtacantha dirupta (Karsch), see Lieftinck, 1930b: 166. 

retrograda Lieftinck (subsp. of Rhinocypha tincta Rambur), 1938: 49—66, fig. 1, 
4B-D, I, 5D. Holotype 8 and first described 9. W. New Guinea, Berau Peninsula 
(Vogelkop), N.E. coast, Manokwari, 23.V.1903 (g') and 5.VIII.1903 (2) (Nieuw 
Guinea Exped.), & “R. tincta Rb.”, det. R. Martin 1907, Q “R. tincta Rb.”, det. 
H. W. van der Weele. — Chlorocyphidae. 

thaphia Lieftinck (Selysioneura), 1959a: 288—290, 293, fig. 2, 22—24. Holotype 
d'. Normanby I. (d’Entrecasteaux Archip.), Mt. Pabinama, 3 mi. E.N.E. of Cape 


> 


110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Prevost, 820 m, 10.V.1956, L. J. Brass (5th Archbold Exped.). — Isostictidae. 
Plate 1. 

rheophila Lieftinck (Huonia), 1935c: 271—274, fig. 34—35 (not 33!), 36A. Lecto- 
type d and first described 9. N. Halmahera I. (N. Moluccas), Tobelo, VII-VIIT. 
1931, M. J. van Diejen. — Libellulidae. 

rhodosoma Lieftinck (Ischnura), 1959b: 220—222, fig. 1, 4. Holotype g'. W. New 
Guinea, Berau Peninsula (Vogelkop), Arfak Mts., Anggi Gita (lake), ca. 1800 m, 
15-25.X.1948, A. J. Kostermans. — Coenagrionidae. Plate 2. 

richardi Lieftinck (Lanthanusa), 1942: 494—497, pl. 27 fig. 40—41, 44. Holotype 
dg and first described 9. Central N. New Guinea, Sahuweri River valley, Mist 
Camp, 1800 m, 14.1.1939 (g') and 9.1.1939 (2), L. J. Toxopeus (3rd Archbold 
Exped.). — Libellulidae. 

[risi van der Weele (Lestes), 1909: 23—24, fig. 1—2. Holotype (presumably lost; 
see Lieftinck, 1960c: 151—154, pl. & fig., with full information). First described 
9 (immature), S. New Guinea, labelled ““Merauke” (pencil), “Lestes 9 nova spec.!” 
(red ink, F. Ris’ handwriting), “Lestes risi Weele, type” (H. W. van der Weele's 
writing). — Lestidae. 

The lost (and unique) G' is here considered to be the holotype, as it would seem 
unwise to designate the 9 as the lectotype of L. risi Weele. 

Currently placed in the subgenus Indolestes Fraser, which has priority over Tricho- 
lestes Fraser, 1951 (nom. praeocc.) and Esaulestes Fraser, 1957, two generic names 
proposed subsequently for this aberrant species; see, however, Lieftinck, 1960c 
(op. cit.) }. 

robusta Fraser (Drepanosticta), 1926: 491—492. Lectotype 9. Kei Islands, Gn. 
Daab, 1922, no. 149, H. C. Siebers, “Drepanosticta robusta @ Type”, in Fraser's 
writing. One very immature g', same locality, no. 89, “Drepanosticta robusta sp.n.”, 
also in Fraser's hand, but with additional red label “Cotype”. — Platystictidae. 

rosenbergi Brauer (Tramea), 1866 (&). First described 9. Sula Is. (S. Moluccas), 
Sula Mangole, Lampau, sea-level, 1X.1939, S. Bloembergen. See Lieftinck, 1942: 
523—524, 531—532, pl. 34 fig. 100. — Libellulidae. 

Now Trapezostigma rosenbergi (Brauer). 

rufipedum Lieftinck (Papuagrion), 1937a: 26, 35—37, fig. 23—24. Holotype d'. 
N. New Guinea, Humboldt Bay area, southern slope of Mt. Cycloop, 1000 m, 
19.IX.1935, W. Stüber. — Coenagrionidae. 

rufithorax Selys (Telebasis), 1877: 122—123. Lectotype g' and first described Q. 
Obi I. (N. Moluccas), labelled “Bernstein ins. Obi” (written, white disk), © 
with additional white label “3 Hagen” (written), under drawer label Telebasis 
rufithorax Selys. Selected by Lieftinck, 1947. — Coenagrionidae. 

Currently placed in Teznobasis. 

rufostigma Selys (Platysticta), 1886b (JS). First described 9. W. Borneo, env. of 
Singkawang, forest-brook near Andjungan, Mampawa Rd., 19.111.1932, L. Coomans 
de Ruiter. See Lieftinck, 1933a: 295—296 (4 £). — Platystictidae. 

Now Drepanosticta rufostigma (Selys). 

sagitta Lieftinck (subsp. of Rhinocypha tincta Ramb.), 1938: 66—67, fig. 1, 4 E-H, 
Q. Holotype & and first described 9. W. New Guinea, Berau Peninsula (Vogelkop, 
southern interior), Vengatap near Aja Maru, 500 m, XII.1937 (g') and id., Jenau, 
350 m, XII.1937 (2), J. M. van Ravenswaay Claasen. — Chlorocyphidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 111 


sagittiferum Lieftinck (Aciagrion), 1949a: 200—201, fig. 243. Lectotype g'. Tanim- 
bar (Timorlaut) Is., Jamdena I., IV.1938, P. Buwalda. — Coenagrionidae. Plate 2. 

First described 9. E. Sumba (Lesser Sunda Is.), Rende Wai, 100 m, 14.VI.1949 
(Sumba Exped. 1949, NMB, XII B 39). 

Type-species of Austroallagma Lieftinck, 1953c: 157—163, fig. 

salomonis Lieftinck (Agriocnemis), 1949b: 349—352, fig. 9. Lectotype g'. Solomon 
Is., Ganonga I. (south of Vella Lavella I.), Koreovuka, 2.X.1936, R. A. Lever. — 
Coenagrionidae. 

saltator Lieftinck (Argiolestes), 1956b: 65, 105—106, fig. 42—43. Holotype {. 
Papua, Maneau Range, N. slopes of Mt. Dayman, 1550 m, 9.VII.1953, G. M. Tate 
(4th Archbold Exped.). — Megapodagrionidae. 

saltuarius Lieftinck (Argiolestes), 1956b: 71, 101—102, fig. 37—38. Holotype &. 
Papua, Port Moresby, 40 km inland, hill forest, Rona Falls, 275 m, 17.11.1936, coll. 
unknown. — Megapodagrionidae. 

salva Ris (Lyriothemis), 1927: 37—38, fig. 24. Holotype G. Central Sumatra, 
Kerintji, Pantjuran Gading (Barisan Range), 1000 m, IX.1915, Edw. Jacobsen. Ex 
SMF no. 17628. — Libellulidae. 

schmidtianum Lieftinck (Pseudagrion), 1936a: 122—124, 127, fig. 8—10. Lectotype 
d'. S. Timor I., Amarasi, X11.1931-1.1932, Ed. Handschin. — Coenagrionidae. 

scintillans Lieftinck (Teinobasis), 1932b: 580—582, fig. 61. Lectotype g and first 
described 9. N. New Guinea, Humboldt Bay area, Hollandia, XI-XII.1930 (g') 
and VI-VII.1930 (2), W. Stüber. — Coenagrionidae. 

selysi Foerster (Argiolestes), 1899a ( g'). First described 9. N. New Guinea, Humboldt 
Bay area, Cycloop Mts, 1000 m, VI-X.1932, W. Stüber. See Lieftinck, 1935c: 
208—211. — Megapodagrionidae. 

Now Podopteryx selysi (Foerster). 

[semitincta Selys (Rhinocypha), 1869a: 664—665 (20—21 sep.). The original speci- 
mens from ‘Gilolo” (Halmahera I., N. Moluccas) in the Leiden museum are no more 
in existence. Females examined i Hagen and de Selys are still represented in that 
collection from S. Halmahera, Batjan I. and Morotai I., leg. Bernstein. — Chlorocy- 
phidae. } 

Specimens of either sex, labelled by de Selys himself and originating from Elbor, 
Gilolo and Batjan (N. Moluccas), are in de Selys’ collection (IRSN), so that any 
selection of a lectotype should, I think, be made from one of these. See also Lieftinck, 
1938: 59—63, fig. 

Currently placed as a subspecies of R. tincta Rambur. 

serena Lieftinck (Teinobasis), 1932b: 582—585, fig. 62. Lectotype and first des- 
cribed 9. N. New Guinea, Humboldt Bay area, Hollandia, XI-XII.1930, W. Stüber. 
— Coenagrionidae. 

Now T. serena serena Lieft. (1949a). 

sericea Selys (Cordulia), 1871 (3). First described ©. Surinam, Coppename River near 
Kroetoe Mt., bush creek, 23.XI.1943, D. C. Geijskes. See Geijskes, 1970: 16—18, 
fig. 3—4. — Corduliidae. 

Now Paracordulia sericea (Selys). 

siebersi Fraser (Drepanosticta), 1926: 490—491, fig. 6a. Holotype & and “allotype 
2” (mutilated and in poor condition). E. Java, Mts. Tengger, 5000 ft., without 


112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


further particulars. Identification in F. C. Fraser's handwriting. — Platystictidae. 
Bona species. 
sieboldi Selys (Cordulegaster), 1854: 107. Lectotype ® (in fragments: most of thorax 
and abdomen missing). Japan, labelled “Burger, Japan” (written, white disk), under 
drawer label “Sieboldii Selys” (written). Since de Selys stated “Japon, Leide et Coll. 
Selys”, this example was selected lectotype by M.A.L. in 1958. — Cordulegasteridae. 
Currently known as Anotogaster sieboldi (Selys). 
silaceum Lieftinck (Pseudagrion), 1932b: 568—570, fig. 54. Lectotype g and first 
described 9. N. New Guinea, Humboldt Bay area, Hollandia, IX.1930 (g') and 
11.1931 (2), W. Stüber. — Coenagrionidae. 
silvarum Lieftinck (Orthetrum), 1934c: 267—269, fig. 3. Lectotype &. W. Java, 
Mt. Pangrango-Gedeh, forest above Tjisarua Est., 1300 m, 17.VII.1932, M. A. Lief- 
tinck. — Libellulidae. Plate 6. 
silvicola Lieftinck (Notoneura), 1949a: 72, fig. 71—73, 93. Holotype g'. S.W. New 
Guinea, Etna Bay, sea-level, 24.X1.1939, H. Boschma (Le Roux Exped.). — 
Protoneuridae. 
silvicola Lieftinck (Huonia), 1942: 489—492, fig. 1. Holotype g'. W. New Guinea, 
Berau Peninsula (Vogelkop), Sungai Anakasi (westcoast of the “neck” of Vogelkop, 
14.V.1941, E. Lundqvist (Negumij Exped.). — Libellulidae. 
First described ©. Misool I. (off W. New Guinea), Fakal, ca. 150 m, 5.X.1948, 
M. A. Lieftinck. See Lieftinck, 1953d: 192, fig. 11. 
simalura Ris (subsp. of Brachydiplax chalybea Brauer), 1919: 1123 (&'). First described 
9. Engano I. (off S.W. Sumatra), Kiojoh, 21.V—12.VIL.1936, J. K. de Jong. See 
Lieftinck, 1948b: 295— 296. (Probably a distinct subspecies.) — Libellulidae. 
simalurum Lieftinck [ subsp. of Pseudagrion pilidorsum (Brauer) }, 1948b: 289— 290, 
fig. 5. Lectotype 4. Simalur I. (off W. Sumatra), Sinabang, 1.1913, E. Jacobson, 
coll. no. 3327. Described by Ris (1915a: 12—13) sub P. pilidorsum Brauer. — 
Coenagrionidae. 
simplex Lieftinck (subsp. of Mesogomphus reinwardti Hagen & Selys), 1934b: 20, 
23—27, fig. 3b, 4. Lectotype g'. Sumatra mer., Benkulen, Banding Agung [Lake 
Ranau}, 27.X.1929, “at lamp”, C. G. G. J. van Steenis. — Gomphidae. 
Now Paragomphus simplex (Lieftinck), but possibly synonymous with P. capricor- 
nis (Foerster), from Malaya. 
simplex Lieftinck (Argiolestes), 1949a: 51—53, fig. 27, 38. Holotype g' and first 
described ®. Central N. New Guinea, Idenburg River basin, above Bernhard Camp, 
700—750 m, 19.111.1939, L. J. Toxopeus & J. Olthof (3rd Archbold Exped.). — 
Megapodagrionidae. 
simulata Belle (Aphylla), 1964a: 26—32, fig. 7—10; 34, fig. 12—14 (larva, sup- 
position). Holotype ¢ and first described @. Surinam, upper Coropina River, Dauw- 
dropkamp, 14.X.1956 (g') and Upper Para River, 4.1.1959 (Q@), J. Belle. — Gom- 
phidae. 
simulatrix Lieftinck (Hylaeargia), 1949a: 136—139, fig. 146, 160, 163—167, 168— 
170. Holotype g and first described 9. Central N. New Guinea, Sahuweri River 
basin, Sigi Camp, 1500 m, 15—28.11.1939, L. J. Toxopeus (3rd Archbold Exped.). 
— Coenagrionidae. 
Type-species of Hylaeargia Lieftinck (op. cit.). 
sinense Asahina (Ceriagrion), 1967: 311—313, 326, fig. 203—211. Holotype g'. S.E. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 113 


China, West Tien-Mu-Shan, 9.VI.1935, H. Hone. — Coenagrionidae. 
sinuatipennis Lieftinck (Arrhenocnemis), 1933d: 4—10, fig. 1—5. Holotype g'. N. 
New Guinea, Humboldt Bay area, upper course of Korimé River, Ajiep, ca. 10 km 
S.W. of Lake Sentani, 165 m, XI—XII.1931, “Leib schwarz, Punkte grün. Brust blau, 
oben mit 2 grünen Streifen. Buckel schwarz” (living colours, W.S.), W. Stüber. First 
described 9. N. New Guinea, Mamberamo River valley, Van Rees Hills, Batavia 
rapids (low country), HI—IV.1940, J. P. K. van Eechoud. — Megapodagrionidae. 
Type species of Arrhenocnemis Lieftinck (op. cit.). 
Originally described in Platycnemididae but later transferred to Megapodagrionidae. 
Family status still questionable. 
sinuatus Fraser (Burmagomphus), 1933 (9). First described &. Ceylon, C.P., Hara- 
gama, stream near Mailapitiya, 16—18.1X.1938, and banks of Mahaweliganga, 18.IX. 
1938, M. A. Lieftinck. See Lieftinck, 1940a: 104—110, fig. 7—9, pl. 1 fig. 3 (incl. 
larva). — Gomphidae. 
Now B. pyramidalis sinuatus Fraser. 
smedleyi Laidlaw (subsp. of Amphicnemis louisae Laidlaw), 1926: 232, fig. 3. Lecto- 
type d and first described 9. Mentawei Archip., Siberut I., IX.1924 (g'), and 
16.IX.1924 (9), C. Boden Kloss & N. Smedley, ex coll. F. F. Laidlaw. — Coen- 
agrionidae. 
Now considered bona species. 
snellemanni Albarda #7 Selys (Micromerus), 1879: 398—399 (52—53 sep.). Lecto- 
type dg. Central Sumatra, with written labels in Snelleman’s hand: “Sumatra Exped. 


van] Soengfei} aboe nfaar} Moearfa} Lab.[oe} 3—13/10.1877” — “Micromerus 
Snellemanni Albarda g'”’, in Albarda’s writing. Ex coll. H. Albarda. — Chlorocy- 
phidae. 


First described 9. N. Sumatra, Atjeh, Pendeng, 400 m, 11.1937, A. Hoogerwerf 
(BM), see Laidlaw, 1950, Trans. R. Ent. Soc. Lond. 101: 276—277. 
Currently known as Libellago (Melanocypha) s. snellemanni (Albarda). 
See also Micromerus sumatranus Alb. and Vestalis lugens Alb. 
snelleni Selys (Rhyothemis), 1878b: 293, 299 (& Celebes). Lectotype 4. N. Celebes, 
labelled “Rosenberg, Celebes Ayer Pann” (print & written), “28 Hag. 65” (written 
on yellow), “Snelleni Sélys” (written drawer-label). — Libellulidae. 
For collector and locality notes of this and other species taken by Von Rosenberg, 
see Lieftinck (1942: 599). 
Currently classified as a subspecies of Rhyothemis phyllis (Sulzer). 
sollaarti Lieftinck (Brachydiplax), 1953e: 262—264, fig. 11 a-b. Holotype &. S. 
Sumatra, Lampong distr., Tandjong Karang, Bergen Estate, 50 m, 26.XII.1952, A. 
Sollaart. — Libellulidae. 
spatulifera Lieftinck (Drepanosticta), 1929b: 112—114, fig. 4—6. Holotype g and 
first described @. Java occ., prov. Banjumas, G. Slamet, Baturraden, 2500 ft., 15.IV. 
1928, F. C. Drescher. — Platystictidae. 
spinicaudum Lieftinck (Papuagrion), 1937a: 27, 29—31, fig. 19. Holotype g'. N. 
New Guinea, Humboldt Bay area, southern slope of Mt. Cycloop, 1000 m, 5.IV. 
1935, W. Stüber. — Coenagrionidae. 
sponsus Lieftinck (Argiolestes), 1956b: 70, 86—87, fig. 18—19. Holotype & and 
first described 9. Central N. New Guinea, Idenburg River basin, Hill-forest Camp, 
between Araucaria Camp and Bernhard Camp B, ca. 700 m, 29.111.1939, L. J. 


114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Toxopeus & J. Olthof (3rd Archbold Exped.). — Megapodagrionidae. 

starreanum Lieftinck (Pseudagrion), 1949a: 181, fig. 173—174, 212. Lectotype ¢. 
W. New Guinea, Kowiai distr., Kaimana (southcoast), 17.XI.1941, J. J. van der 
Starre. — Coenagrionidae. 

starrei Lieftinck (Pornothemis), 1948a: 270—274, fig. 17. Holotype g'. E. Sumatra, 
Palembang, “on board steamer on Musi River, near Palembang”, 25.11.1938, J. J. van 
der Starre. — Libellulidae. 

stenomantis Lieftinck (Selysioneura), 1932b: 543, 549—551, fig. 4446. Lectotype 
d' and first described 9. N. New Guinea, Humboldt Bay area, Hollandia, IV—VI. 
1931, “nur auf Baume’, W. Stüber. — Isostictidae. 

stenoptera Lieftinck (Gynacantha), 1934c: 265—266, fig. 1. Holotype g'. With old 
locality label: “Auctie vlan] Eyndhfoven} Java”, in H. Albarda’s handwriting. — 
— Aeshnidae. 

stephanodera Lieftinck (Asthenocnemis), 1949a: 90—91, fig. 96, 98, 100, 108—110. 
Holotype @. “Nelle Guinée v. 5 cornes thorax” (red label in R. Martin's hand- 
writing), ex Mus. Paris. — Platycnemididae. 

N.B. — Erroneously labelled as from New Guinea! The specimen undoubtedly 
originates from the Philippine Islands, which is evident from a series of both sexes that 
have been taken in Palawan I. (P.I.) by members of the Danish “Noona Dan” 
Expedition (September 1961), the ¢ being still undescribed. 

Type-species of Asthenocnemis Lieft. (op. cit.). 

Family status doubtful. 

stigmatizans Lieftinck (Teinobasis), 1938: 109—112, 113, fig. 42, 44. Holotype & 
and first described 9. N. New Guinea, Bewani-Wari Hills, Sawia, 200 m, X—XI. 
1936, W. Stüber. — Coenagrionidae. 

stillacruoris Lieftinck (Paramecocnemis), 1956a: 254—258, fig. 5—7 (as P. stilla- 
cruoris). Holotype g'. N.E. New Guinea, Upper Jimmi Valley, north slopes of 
Sepik-Waghi Divide (W. of Mt. Wilhelm), Tsenga, 1200 m, 13.VII.1955, J. L. 
Gressitt. — Platycnemididae. 

[strigata Hagen in Selys (Mnais), 1853 (& Q Japon). First described 9. Japan; des- 
troyed? (not in ML). — Calopterygidae. 

Redescribed and fore wing figured by de Selys in the Monographie (“... une 
femelle du Musée de Leyde”, 1854: 64—65, pl. 3 fig. 1).} 

strumidens Lieftinck (Idiocnemis), 1958: 261—263, 282, fig. 12—16; 290—292, 
fig. 69, 74—76. Holotype & and first described 9. W. New Guinea, western ex- 
tremity, Sorong, grassy trail near Malano, 30.IX.1948, M. A. Lieftinck. — Platy- 
cnemididae. 

stueberi Lieftinck (Anacordulia), 1938: 127—128, fig. 52 (as A. stiberi). Holotype 
Q. N. New Guinea, Humboldt Bay area, Hollandia, 19.XII.1930. W. Stüber. — 
Corduliidae. 

First discussed and figured &. N. New Guinea, Bewani Hills, Wembi, 270 m, 
18.X11.1937, W. Stüber. 

Currently considered as a synonym of Metaphya tillyardi Ris, 1913, and Anacor- 
dulia maccullochi Tillyard, 1926 (see Lieftinck, 1961b: 420—423, fig. 4—5, d'). 

Now placed in Metaphya Laidlaw. 

stueberi Lieftinck (Papuargia), 1938: 101—105, fig. 36a, 37a, 38—39 (as P. stüberi). 
Holotype g and first described 9. N. New Guinea, southern Bewani Hills, upper 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 115 


course of Tami River, 5 days’ journey S.E. of Hollandia, 250 m, Fumb (Keerom?) 
River, 26.1V.1937, W. Stüber. — Coenagrionidae. 
Type-species of Papuargia Lieftinck (op. cit). 
stueberi Lieftinck (Ischnura), 1932: 594—596, fig. 66 (as I. stüberi). Lectotype & 
and first described 9. N. New Guinea, Humboldt Bay area, Hollandia, 111.1931, W. 
Stüber. — Coenagrionidae. 
suavis Lieftinck (Teinobasis), 1953e: 244—246, fig. 5. Holotype 4 and first described 
©. S.E. Borneo, Kandangan distr., Ampah, 0—20 m, 7 km N.W, IV—V.1948 (d') 
and same area, Ranamun, 21.V.1948, Liem Swie Liong. — Coenagrionidae. 
subornatus Lieftinck (subsp. of Argiolestes ornatus Selys), 1935c: 218, 224—228, 
fig. 8b-d, 10. Lectotype and allotype @ (in cop.). N. New Guinea, Humboldt Bay 
area, Kressi, 120 km south, 300 m, 1.1932, W. Stüber. — Megapodagrionidae. 
Currently considered a distinct species (see Lieftinck, 1949a: 46, and 1956b: 88— 
89, fig. 23). 
suensoni Lieftinck (Gomphus), 1939b: 285, 294—297, fig. 11—12. Holotype ¢ and 
first described 9. Central E. China, South Shensi, 13.VII.1936, E. Suenson. — 
Gomphidae. 
Now transferred to genus Sznogomphus May. 
sufficiens Lieftinck (subsp. of Idiocnemis zebrina (Lieft.), 1958: 274, fig. 38—42. 
Holotype & and first described ©. Fergusson I. (d’Entrecasteaux Archip.), mountains 
between Agamoia and Ailuluai, S.-coast, 900 m, 14.VI.1956, L. J. Brass (Sth Arch- 
bold Exped.). — Platycnemididae. 
sufficiens Lieftinck (Lanthanusa), 1955b: 157—160, fig. 1—4. Holotype g and first 
described 9. Goodenough I. (d’Entrecasteaux Archip.), 1600 m Camp, east slopes, 
11—12.X.1953, K. M. Wynn & L. J. Brass (4th Archbold Exped.). — Libellulidae. 
sufficiens Lieftinck (Rhyacocnemis), 1956a: 250—254, fig. 1—4. Holotype &. 
Goodenough I. (d'Entrecasteaux Archip.), 1600 m Camp, east slopes, 12.X.1953, 
K. M. Wynn & L. J. Brass (4th Archbold Exped.). — Platycnemididae. 
Type-species of Rhyacocnemis Lieftinck (op. cit.). 
sumatrana Lieftinck (Oligoaeschna), 1953e: 253—255, fig. 8. Holotype g'. W. Su- 
matra, Mt. Kerintji (Peak of Indrapura), Kaju Aro Est., 1600 m, 30.X.1952, R. 
Straatman. — Aeshnidae. 
sumatranus Albarda #7 Selys (Micromerus), 1879: 397—398 (51—52 sep.). Lecto- 
type g and first described 9. Central Sumatra, with written labels in Snelleman’s 
hand: “Sumatra Exped./Silago tot 12/7.77” (4), and “Moeara laboe tot 28.11.77” 
(2); with “Micromerus sumatranus Albarda” in Selys’ writing. Ex coll. H. Al- 
barda. — Chlorocyphidae. 
Now Libellago sumatrana (Albarda). 
See also Micromerus snellemanni Alb. and Vestalis lugens Alb. 
sundana Lieftinck (subsp. of Epophthalmia vittata Burm.), 1931: 30, 38, 45, 61—64, 
pl. 1 fig. 3, fig. 1, 8, 16. Holotype ¢ and first described 9, W. Java, Bogor (Bui- 
tenzorg), 250 m, Botanic garden, 21.III and 3.X.1930, respectively, M. A. Lieftinck. 
— Corduliidae. 
sundana Krueger (Platysticta), 1898 (3). First described 9. West, Central and East 
Java, various locs., no dates selected. See Lieftinck, 1934d: 470—471. — Platysticti- 
dae. 
Now Drepanosticta sundana (Krueger). 


116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


[superba Selys (Telebasis), 1877: 114—115. Originally described after males and females 
from “Celèbes, Menado (Musée de Leyde); Moluques, par M. Lorquin (Coll. Sélys)”’. 
The name is Hagen’s (see de Selys: “Agrion superbum Hag., Mus. de Leyde”, op. 
cit.: 114), but was validated by de Selys. At present old-time examples of T. superba 
in the Leiden museum are labelled “Halmahera [leg.} Bernstein”, sub nom. Agrion 
nobile Hagen, in H. A. Hagen’s handwriting, the individuals from Celebes being 
probably destroyed. Hence one of the specimens labelled “Moluques, Lorquin”, in de 
Selys’ collection (IRSN) should be designated lectotype. — Coenagrionidae. 

Currently placed in Teznobasts.} 

superstes Foerster (Bironides), 1903 (G'). First described 9. E. Papua, Peria Creek, 
Kwagira River, 50 m (Camp 7, 4th Archbold Exped.), 25.VIII.1953, G. M. Tate. 
See Lieftinck, 1963c: 758—759. — Libellulidae. 

surinamensis Geijskes (Micrathyria), 1963: 71—74, fig. 15. Holotype g'. Surinam, 
near Sectie O, 11.V.1949, D. C. Geijskes. First described 9. Surinam, Sipaliwini 
savanna, pool in forest near airstrip, 12.11.1961, D. C. Geijskes. — Libellulidae. 

sylvia Lieftinck (Procordulia), 1935c: 295, 298—300, fig. 50. Holotype g and first 
described 9 (immature). N. New Guinea, Humboldt Bay area, Cycloop Mts., ca. 
1000 m, 12.IX.1932 (g') and 19.V.1932 (2), W. Stüber. — Corduliidae. 


tachyerges Lieftinck (Mesogomphus), 1934b: 19, 30—33, fig. 3, 6. Lectotype g'. E. 
Sumba I. (Lesser Sunda Is.), Kananggar, 700 m, Waidjelo River, V.1925, K. W. 
Dammerman. — Gomphidae. Plate 5. 

First described 9. W. Sumba I, Waimangura, 436 m, 19.VIII.1949 (Sumba 
Exped. 1949, NMB), see Lieftinck, 1953c. 
Currently known as Paragomphus tachyerges (Lieft.). 

tanysiptera Lieftinck (Palaiargia), 1953e: 239—240, fig. 4. Holotype g'.Halmahera I. 
(N. Moluccas), Mt. Sembilan, 600 m, 4.X.1951, Sundanese collectors. — Coen- 
agrionidae. 

tarafia Lieftinck [subsp. of Nesoxenia mysis (Selys) }, 1942: 462, 464465, pl. 24 fig. 
17, pl. 41 fig. 164. Lectotype g and first described 9. N. New Guinea, ca. 150 km 
W. of Humboldt Bay (Hollandia), Dempta Hills, Tarafia, 600 m, 10.V.1935, W. 
Stüber. — Libellulidae. 

Probably bona species. 

tenebricosa Lieftinck (Prodasineura), 1937b: 82—84, fig. 15. Holotype & and first 
described 9. W. Borneo, Singkawang, forest brook near Bengkajang, 15.VI.1933 
(g') and same area, Raja river near Montrado, 22.X.1931 (9), L. Coomans de 
Ruiter. — Protoneuridae. 

tenella Lieftinck (Drepanosticta), 1935b: 7, fig. 1. Lectotype & and first described 9. 
S. Sumatra, S. Lampong prov., Gisting and environs, foot of Mt. Tanggamus, 4—600 
m, 25—30.XII.1934, M. A. Lieftinck. — Platystictidae. 

tenella Lieftinck (Archibasis), 1949a: 184, 186, 188, fig. 235—236. Holotype &. 
W. Borneo, Singkawang, forest-brook near Bengkajang, 15.VI.1933, L. Coomans de 
Ruiter. First described 9. E. Borneo, Kutai, Sangkulirang, VI.1937, M. E. Walsh. 
— Coenagrionidae. 

tenera Lieftinck (Hemicordulia), 1930b: 157—159, fig. 22. Holotype 3. Central 
Java, prov. Banjumas, G. Slamat, Baturraden, 760 m, 14.11.1929, native coll., F. C. 
Drescher. — Corduliidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 117 


tenuispinus Lieftinck (Argiolestes), 1938: 73—75, fig. 9, 10a, 11. Holotype g' and 
first described ©. S.E. New Guinea, Astrolabe Range, Moroka, 1300 m, IX.1893 
(8) and 27.VII.1893 (9), Lamberto Loria, ex MCG. — Megapodagrionidae. 

terpsichore Foerster (Macromia), 1900 (&'). First described 9. N. New Guinea, 
Humboldt Bay area, Hollandia, V.1930, W. Stiiber. See Lieftinck, 1942: 562—563. 
— Corduliidae. 

teuchestes Lieftinck (Bironides), 1933d: 30, 33, fig. 18, 19—22. Lectotype g and 
first described 9. N. New Guinea, Humboldt Bay area, Hollandia, IV—VI.1931 
(g'), and same area, 400 m, IX.1930 (9), W. Stüber. — Libellulidae. Plate 6. 

thais Lieftinck (Huonia), 1953d: 189, 208—216, fig. 10C, 19—23. Holotype & and 
first described 9. Misool I. (off W. New Guinea), Kasim River, 100—150 m, 
12.1X.1948 (g') and 17.IX.1948 (9), M. A. Lieftinck. — Libellulidae. 

thalassina Lieftinck (Notoneura), 1949a: 74—76, fig. 94—95. Holotype & and 
first described 9. S. New Guinea, Digul River territ., Ederat near Aja Maru, 11.1939, 
J. M. van Ravenswaay Claasen. — Protoneuridae. 

thalia Lieftinck (Selysioneura), 1953b: 642—646, fig. 1, 5, 9, 12. Holotype g' and 
first described 9. Halmahera I. (N. Moluccas), Mumar River, 200—300 m, 25.IX. 
1951, Sundanese collectors. — Isostictidae. Plate 1. 

thisbe Lieftinck (Huonia), 1953d: 188, 200—203, fig. 16, 19, 20. Holotype &. Misool 
I. (off W. New Guinea), Fakal, ca. 150 m, 23.1X.1948, M. A. Lieftinck. — Libel- 
lulidae. 

tillyardi Lieftinck (subsp. of Tramea loewi Brauer), 1942: 527530, pl. 33 fig. 97, 
pl. 34 fig. 103. Holotype & and first described ©. N.E. Australia, N. Queensland, 
Redlynch, 4.IX.1938 (g') and Kuranda, 12.VIII.1938 (2), R. G. Wind. — Libel- 
lulidae. 

Falls within the range of variation of Trapezostigma loewt (Brauer), see Lieftinck, 

1962: 91. Hence synonymous with that species. 

titschacki Schmidt (Gomphoides), 1952 (g'). First described Q. Perú, Rio Rondos 
(Huallaga River area), X.1962—11.1963 (& 9), L. Gómez Alonso. See Belle, 1970: 
100—102, fig. 154—155. — Gomphidae. 

Now Phyllocycla titschacki (Schmidt). 

tonsillare Lieftinck (Aciagrion), 1937a: 49—51, fig. 34—35. Lectotype { and allo- 
type $ (in cop.). N. New Guinea, Humboldt Bay area, upper course of Korimé 
River, Mameda (W. of Lake Sentani), 200—300 m, XII.1934. W. Stüber. — Coen- 
agrionidae. 

toxopei Lieftinck (Hemicordulia), 1926: 281—284, fig. 1—4. Holotype 9. Buru I. 
(S. Moluccas), Leksula, 25.XII.1921, L. J. Toxopeus. — First described 3. Buru I, 
Kunturun, 9.VII.1921, same collector (Lieftinck, 1930a: 314—315, fig. 3). — Cor- 
duliidae. 

tristis Lieftinck (Argiolestes), 1935c: 218, 229—231, fig. 13. Lectotype ¢ and allo- 
type ® (in cop.). N. New Guinea, Humboldt Bay area, Cycloop Mts., southern 
slopes, 1000 m, 26.IV.1932, W. Stüber. — Megapodagrionidae. 

truncatum Lieftinck (Xiphiagrion), 1949a: 197—199, fig. 238. Holotype & and first 
described 9. N. New Guinea, Humboldt Bay area, southern Bewani Hills, upper 
reaches of Tami River, 300—500 m, Sawia, X—X1.1936, W. Stüber. — Coenagrion- 
idae. 

tumidula Lieftinck (Agyrtacantha), 1937a: 63—65, fig. 39c, 40. Lectotype & and 


118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


first described 9. N. New Guinea, Humboldt Bay area, southern slope of Mt. ey 
cloop, 700 m, IX.1933, W. Stüber. — Aeshnidae. 
umbratilis Lieftinck (Selysioneura), 1932b: 543, 551, 602, fig. 47. Lectotype 9. 


N. New Guinea, Humboldt Bay area, Hollandia, IV—VI.1931, W. Stüber. First _ 


described &, same area, Hollandia, 1.11.1932, W. Stüber (see Lieftinck, 1949a: 81— 
82, fig. 84). — Isostictidae. 

undulatus Needham (Gomphoides), 1944 (&). First described 9. Surinam, Coropina 
Creek, upper part, near Zanderij, 30.1V.1951, D. C. Geijskes. See Geijskes, 1964: 
40—41, fig. 18—19. — Gomphidae. 

uropetala Lieftinck (Oligoaeschna), 1968a: 149, 152, 171—174, fig. 6—7. Holotype 
d and first described 9. N.E. Sumatra, Deli, Laut Tador, 90 m, 10.1V.1949 (8) 
and 12.V.1948 (9), R. Straatman. — Aeshnidae. 

vanderweelei Martin (Heliaeschna), 1907b: 222—223 (sub H. van der Weelei). 
Holotype g'. With erroneous locality label: “Sala Bendo, Liberia Buttikofer’, written 
on white disk, “Heliaeschna Van der Weelei Martin &” in R. Martin's writing, and 
“limpida Sél” in pencil (unknown hand). Ex Mus. Paris. — Aeshnidae. 

Syn. of Heliaeschna simplicia Karsch. See also Lieftinck, 1940b: 389. 

venatrix Lieftinck (Plattycantha), 1937a: 73—75, fig. 43, 45. Lectotype g'. N. New 
Guinea, Humboldt Bay area, southern slopes of Mt. Cycloop, 1000 m, IX.1932, W. 
Stiiber. — Aeshnidae. 

venilia Lieftinck (Selysioneura), 1953b: 651—654, 658, fig. 4, 14. Holotype 9. W. 
New Guinea, Berau Peninsula (Vogelkop), Beraur distr., Klamono Oilfields, 40 km 
inland, 40—50 m, 23—24.VIII.1948, M. A. Lieftinck. — Isostictidae. Plate 1. 

venusta Lieftinck (Oligoaeschna), 1968a: 147, 168, fig. 6 & pl. 13. Holotype d'. 
E. Borneo, Kutai, Tabang, Bengen River, 125 m, 9.IX.1956, A. M. R. Wegner. — 
Aeshnidae. 

viduata Lieftinck (subsp. of Ischnura aurora Brauer), 1949a: 222—223, fig. 275. 
Holotype 4 and first described ©. Central W. New Guinea, Baliem River valley, 
environs of Baliem R. Camp, 1700 m, 15.X1.1938, L. J. Toxopeus (3rd Archbold 
Exped.). — Coenagrionidae. Plate 2. 

viduatum Lieftinck [subsp. of Orthetrum sabina (Drury) }, 1942: 475—478, pl. 25 fig. 
33. Holotype & and allotype ® (in cop.). Central W. New Guinea, Baliem River 
valley, 1600 m, 11.X11.1938, L. J. Toxopeus (3rd Archbold Exped.). — Libellulidae. 

viola Lieftinck (Archibasis), 1948 (nom. nov., sine descr.); 1949a: 184, 186, 188— 
189, fig. 230, 234. Holotype 4 and first described 9. Karimondjawa Is. (Java Sea), 
Kali Todjero, P. Karimon, 28.X1.1930, M. A. Lieftinck. — Coenagrionidae. 

virgula Lieftinck (Selysioneura), 1959a: 282—284, fig. 1, 10, 19—21. Holotype {. 
Woodlark I. (Murua I.), Kulumadau, 10.11.1957, W. W. Brandt. — Isostictidae. 
Plate 1. 


walshae Lieftinck (Acrogomphus, sub walshz), 1935a: 203—205, fig. 13—15 (1937b: 
118, footnote, nom. emend.). Holotype g'. S.W. Sumatra, Benkulen distr., Muara 
Tenam, ca. 250 m, 16—23.VI.1925 (rect. 1935), M. E. Walsh. First described 9. 
W. Java, Djampang Tengah (Bodjonglopang), Tjitalahab Estate, 650 m, 30.1V.1940, 
taken in cop., Mrs. C. Rosier. — Gomphidae. Plate 4. 

wayana Geijskes (Staurophlebia), 1959: 151, 164—168, fig. 74, pl. 7b. Holotype & 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 119 


and first described 9. Surinam, Loë Creek, Litani River, 20.VIII.1939, D. C. Geijs- 
kes. — Aeshnidae. 

wijaya Lieftinck (Microgomphus), 1940a: 98—104, fig. 4—6 & pl. 1 fig. 4 (exuviae). 
Holotype & (freshly emerged, with exuviae) and first described Q (immature). Cey- 
lon, Central Prov., Haragama, tributary of Mahaweliganga River, near Mailapitiya, 
500 m, 17—18.IX.1938, M. A. Lieftinck. Holotype & preserved with exuviae in 
alcohol. — Gomphidae. 

williamsoni Ris (Dythemis), 1919 (& ). First described 9. Surinam, Upper Coropina 
Creek near Zanderij, 1.11.1958, J. Belle. See Geijskes, 1964: 45—47, fig. 17. — 
Libellulidae. 

williamsoni Laidlaw (Leptogomphus), 1912 (3). First described 9. E. Borneo, Kutai, 
Sangkulirang distr., Dagaunan, VI.1937, M. E. Walsh. See Lieftinck, 1948a: 247— 
249, pl. 8 and fig. 10. — Gomphidae. 

xanthe Ris (Rhinocypha), 1927: 7—9, fig. 2. Lectotype g and first described 9. 
N. Central Sumatra, foot of Kerintji Peak, Sungai Kumbang, 1400 m, VIII.1915, E. 
Jacobson, & with coll. no. 88/09 (or 60/88). — Chlorocyphidae. 

xanthe Lieftinck (Notoneura), 1938: 93, fig. 27—28. Holotype g. Salawati I. (off 
western extremity of New Guinea), [H. A.} Bernstein leg. — Protoneuridae. 

xanthocyane Lieftinck (Oreagrion), 1949a: 216—220, fig. 246, 252—253, 256—257, 
268, 273—274. Holotype g' and first described 9. Central N. New Guinea, Snow 
Mountain (Nassau) Range, peat bog swamps in moss forest, N. of Lake Habbema, 
3000 m, 24.VIII.1938, L. J. Toxopeus (3rd Archbold Exped.). — Coenagrionidae. 
Plate 3. 

xanthocyanus Selys (Micromerus), 1869a: 666 (22 sep.). Lectotype 3. [N. Celebes}, 
labelled “Rosenberg, Limbotto, Celebes’’ (written), and ‘“Micromerus xanthocyanus 
Selys’, in de Selys’ handwriting. For particulars concerning the collector, his locali- 
ties and dates of capture, see Lieftinck, 1942: 599. 

Now Libellago xanthocyana (Selys). 


zebrina Lieftinck (Idiocnemis), 1958: 272—274, 283, fig. 33— 37. Holotype g' and 
first described ©. Papua, North-Eastern Prov., Cape Vogel Peninsula, Menapi, up 
to 150 m, 9.IV.1953, G. M. Tate (4th Archbold Exped.). — Platycnemididae. 


120 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


IV. ANNOTATED LIST OF TYPE-SPECIMENS IN FOREIGN 
INSTITUTIONAL COLLECTIONS DESCRIBED BY DUTCH AUTHORS 


N.B. — The following symbols (bracketed in the next list) have been used to indicate 
the whereabouts of type-specimens of species-group taxa belonging to this category: 


AMNH 
BISH 
BM 
CAS 
CNHM 
CUI 
IRSN 
KU 
MAT 
MBUD 
MC 


MKB 
ML 
MNB 
MP 
NMB 
NRS 
ROMT 
RSM 
SMF 
UMMZ 
USNM 


ZIUL 
ZMA 
ZMH 


American Museum of Natural History, New York 

Bernice P. Bishop Museum, Honolulu 

British Museum (Natural History), London 

Science Museum, California Academy of Sciences, San Francisco 
Chicago Natural History Museum, Chicago 

Department of Entomology, Cornell University, Ithaca 

Institut Royal des Sciences Naturelles, Bruxelles 

Entomological Laboratory, Kyushu University, Fukuoka 

Koninklijk Museum voor Midden-Afrika, Tervuren 
Természettudomanyi Múzeum Allattára, Budapest 

Zoologisk Institut & Museum (Universitetets Zoologiske Museum), Copen- 
hagen 

Zoologisches Forschungsinstitut u. Museum Alexander Koenig, Bonn 
Rijksmuseum van Natuurlijke Historie, Leiden 

Museum fiir Naturkunde an der Humboldt-Universitat, Berlin 
Muséum National d’Histoire Naturelle, Paris 

Naturhistorisches Museum, Basel 

Naturhistoriska Riksmuseet, Stockholm 

Royal Ontario Museum, Toronto 

Royal Scottish Museum, Edinburgh 

Natur-Museum Senckenberg, Frankfurt a.M. 

Museum of Zoology, University of Michigan, Ann Arbor 

United States National Museum (now National Museum of Natural History, 
Washington, D.C.) 

Zoological Institute, University, Lund 

Instituut voor taxonomische Zoölogie, Amsterdam 

Zoologisches Staatsinstitut und Zoologisches Museum, Hamburg 


aconita Lieftinck (Trithemis), 1969c: 40— 44, fig. 7C. Holotype &. Congo, Katanga, 
Lower Luapula Region, Kisamamba, S. of Kasenga, Mululushi River, at bridge on 
Kasenga-Kialwe road, 950 m, 3.1V.1961, J. J. Symoens, no. 8504 (MAT). — Libel- 


lulidae. 


aequalis Lieftinck (Trithemis), 1969c: 44—45, fig. 7A. Holotype 4 and first described 
Q.N.E. Zambia, Lake Bangweulu Region, 2 km from Ndoba, 1150 m, 30.XII.1961 
(g') and Mundubi, W.-shore of Lake Chifunabuli, 1140 m, 30.XII.1961 (9), 
J. J. Symoens, nos. 9134 (g') and 9149 (2) (MAT). — Libellulidae. 

aerides Lieftinck (Teinobasis), 1962: 20, 34— 36, fig. 10a-c. Holotype g'. E. Caroline 
Is. (Micronesia), Ponape I., Mt. Pairot, 660 m, IV—IX.1950, Adams (USNM, 
65142). — Coenagrionidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 121 


alcathoe Lieftinck (Gynacantha), 1961a: 145—148, fig. 44, 51, 52. Holotype gd. 
Philippine Is., Mindanao I., Davao Prov., Tagum, Madaum, sea-level, 10.X.1946, 
F. G. Werner (CNHM). — Aeshnidae. 

anderi Lieftinck (Anisogomphus), 1948d: 59—63, fig. 1—2. Holotype g'. China, 
prov. Hunan, Tien-Chao-Shan, ca. 1200 m, 31.VIII.1938, G. Österlin (ZIUL). — 
Gomphidae. 

annulosum Lieftinck (Ceriagrion), 1934a: 11—12, fig. 4. Holotype g'. Java, with 
pin-label: Java Fr[uhstorfer} 61 (yellow square), and Pseudagrion annulosum Selys, 
n.sp Java (written drawer label, both in de Selys’ writing) (IRSN). — Coenagrion- 
idae. 

First described ©. S.W. Java, Sempurtjondong, Tjidaun, 8.X1.1935, M. Bartels Jr. 

(see Asahina, 1967: 263—265, 323, fig. 13). 

arboreus Lieftinck (Burmagomphus), 1940a: 111, 112. Lectotype g'. Burma, R. A. 
Earnshaw; with several identification labels (UMMZ). Selected by Lieftinck, 1964. 
— Gomphidae. 

Nom. nov. pro Burmagomphus vermiculatus (rect. vermicularis!) Williamson, nec 

R. Martin, from Burma, i.e. the above specimen (see Williamson, 1907, Proc. U.S. 
Nat. Mus. 33: 301—303, fig.). Also for Burmagomphus williamsoni Fraser, 1926, 
nec B. vermicularis williamsoni Foerster, 1914. For further particulars, see Lieftinck, 
1940a: 111, and 1964b: 15, 17—22, fig. 

ariel Lieftinck (Teinobasis), 1962: 20, 31—34, fig. 8—9. Holotype g and first 
described 9. E. Caroline Is. (Micronesia), Ponape I., Agric. Exper. Sta, VI—IX. 
1950, Adams ( &£, USNM 65141). No selection of allotypic 9. — Coenagrionidae. 

arsinoe Lieftinck (Diplacina), 1953d: 156, 171—173, fig. 5—6. Holotype 4. Papua, 
Kokoda, 1200 ft, VIII.1933, L. E. Cheesman (BM). See Kimmins, 1968: 281. — 
—Libellulidae. 

aruanus Lieftinck (Lestes sg. Indolestes), 1951b: 8—9, fig. 4. Holotype g' (head 
wanting). Aru Is., L. albicauda McLachlan, det. R. MacLachlan (BM). See Kimmins, 
1970: 194. — Lestidae. 

asahinai Lieftinck (Macromidia), 1971b: 21. Nom. nov. pro Idionyx philippa, Asahina, 
1968 ( Palawan), nec Ris, 1912c (® Mindoro, P.I.). Holotype g'. Palawan I, 
5—8 mi. E. of Tarumpitao, Pf., 28.V.1958, H.E.M. (see Asahina, 1968: 357—358, 
fig. 5—10, pl. 2 fig. 11) (BISH). — Corduliidae. 

atrocyana Lieftinck (Notoneura), 1960b: 117—120, fig. 1—5. Holotype 4 and first 
described @. Waigeu I. (off N.W. New Guinea), Camp Nok (Mt. Buffelhoorn), 
2500 ft, IV.1938, L. E. Cheesman (BM). See Kimmins, 1970: 176. — Protoneuridae. 

atropha Lieftinck (Vestalis), 1965b: 337, 351—353, fig. 4, 8. Holotype g'. N.W. 
Borneo, Sarawak, Mt. Dulit Trail, 10.VIII.1932/Primitive forest/Oxford Univ. Exped. 
(BM). See Kimmins, 1969: 304. — Calopterygidae. 

aurantipes Lieftinck (Platycnemis), 1965c: 248— 250, fig. 10. Holotype { and first 
described 9. Madagascar, Prov. Tamatave, Périnet, 25.1X.1958, F. Keiser et al. 
(NMB). — Platycnemididae. 

austrosundanum Lieftinck (Orthetrum), 1953c: 208—210, fig. 72. Holotype g' and 
first described 9. Sumba I. (Lesser Sunda Is.), Waimangura, ca. 430 m, Wulu 
Mano, 20.VIII.1949 (Sumba Exped. 1949, NMB XII C 302). — Libellulidae. 


bellax Lieftinck (Lestes sg. Indolestes), 1953c: 145—148, fig. 4—5. Holotype & 


122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


and first described 9. Sumba I. (Lesser Sunda Is.), E. Sumba, Rende Wai, 100 m, 
11—16.VI.1949 (Sumba Exped. 1949, NMB, XII B 37). — Lestidae. 

berlandi Lieftinck (Drepanosticta), 1939a: 145—147, fig. 1. Holotype g'. Lombok I. 
(Lesser Sunda Is.), Sapit, 2000 ft, IV.1896, H. Fruhstorfer (MP). — Platystictidae. 

berlandi Lieftinck (Macromia), 1941a: 94— 098, fig. 1—2. Holotype g'. Indochina, 
labelled: “Tonkin, monts Mauson, Avril-Mai, 2—3.000, H. Frihstorfer’’, and “M. 
borneensis”, by R. Martin (MP). See also Martin, 1907a: 69, fig. A (base of hind 
wing d holotype, sub M. borneensis Krüger, misidentified). — Corduliidae. 

bigemmata Lieftinck (Elattoneura), 1971a: 194—198, fig. 6. Holotype g'. Ceylon, 
Western Prov., Labugama, 24 mi. E.S.E. of Colombo, loc. 17, 9.III.1962, A. Perera, 
Lund Univ. Ceylon Exped. 1962 (ZIUL). — Protoneuridae. 

bivittata Lieftinck (Protosticta), 1939a: 151—154, fig. 4. Holotype g'. Celebes, label- 
led “Célébes/Protosticta simplicinervis?? nov. sp.”, in R. Martin's handwriting (MP). 
First described 9. S. Celebes, Bonthain, C. Ribbe, 1884 (ML). 

blandulus Lieftinck (Heliogomphus), 1929b: 123—125, fig. 14—16. Holotype &. 
Equat. [W.] Borneo, basin of Kapuas River, “Sumpfwald am Bika Fluss”, 4.11.1925, 
leg. Prof. H. Winkler (ZMH). — Gomphidae. 

boharti Lieftinck (Tapeinothemis), 1950a: 638—642, fig. 5—6. Holotype 9. Solomon 
Is., Little Florida I., 15-31.111.1945, G. E. Bohart (USNM). — Libellulidae. 

Type-species of Tapeinothemis Lieftinck (op. cit.). 

brincki Lieftinck (Drepanosticta), 1917a: 191—192, fig. 2a-c. Holotype g' and first 
described ® (subadult). Ceylon, Prov. Sabaragamuwa, Deerwood, Kuruwita, 6 mi. 
N.N.W. of Ratnapura, loc. 90 III, 21.11.1962, in ravine, Lund Univ. Ceylon Exped. 
1962 (ZIUL). — Platystictidae. 

burmana Lieftinck (Gynacantha), 1960a: 248—251, fig. 11a-b. Holotype g'. Burma, 
labelled in de Selys’ writing: “Palone, 16.IX.89” and “Gynac. bayadera S. g'”’, with 
additional labels from R. Martin (IRSN). — Aeshnidae. 


caesarea Lieftinck (Gomphidia), 1929b: 139—143, fig. 31—32. Holotype g'. Central 
W. Borneo, Lebang Hara, 25.XI-5.XI1.1924, Prof. H. Winkler leg. (ZMH). — 
Gomphidae. 

calliope Lieftinck (Gynacantha), 1953e: 260—262, fig. 10. Holotype g'. Waigeu I, 
off N. W. New Guinea, Camp Nok, 2500 ft, IV.1938, L. E. Cheesman (BM). 
See Kimmins, 1969: 301. — Aeshnidae. 

callirrhoe Lieftinck (Diplacina), 1953d: 156, 169—171, fig. 5—6. Holotype d'. 
W. New Guinea, Berau Peninsula (Vogelkop), Karoon, labelled “N. Guinée La- 
glaize” and “Diplacina smaragdina S. g'” (lilac labels in de Selys’ handwriting) and 
additional pin-labels “D. smaragdina det. F. Ris” (IRSN). — Libellulidae. 

N.B. — The 4 and © in de Selys’ collection, with the same locality labels and 
selected by F. Ris and myself as the types of true D. smaragdina Selys, are lectotype 
and allotype, respectively, of that species; these also are deposited in the Brussels 
museum (see lieftinck, op cit.: 173—174). 

cardinalis Lieftinck (Agrionoptera), 1962: 72, 74, fig. 20b, 21f, 22g, h. Holotype 
d' and first described 9. W. Caroline Is. (Micronesia), Palau Is., Babelthuap I., 
Ulimang, XII. 1947, Dybas (& USNM 65145, 9 USNM). — Libellulidae. 

carolinensis Lieftinck (Teinobasis), 1962: 20, 23—24, fig. 5, 6, 10. Holotype g' and 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 123 


first described 9. E. Caroline Is. (Micronesia), Truk Is., Wena (Moen) I., Nantuka 
area (Mts. Chukumong and Tonaachau), 21.111.1949, Langford (4 USNM 65137) 
and 25.III.1949, same coll. (@ USNM). — Coenagrionidae. 

castor Lieftinck (Onychogomphus), 1941b: 246—247, pl. 16 fig. 6, pl. 14 fig. 1—3. 
Holotype &. Malay Peninsula, Kelantan, Heyne vdt. 1903, ex coll. F. Förster 
(UMMZ). — Gomphidae. 

ceylonicum Lieftinck (Mortonagrion), 1971a: 198—199, fig. 8. Holotype g'. Ceylon, 
Northwest Prov., Kadaimparu, 15 mi. N. of Negombo, loc. 36, 3.1.1962, Lund Univ. 
Ceylon Exped. 1962 (ZIUL). — Coenagrionidae. 

chrysochlora Lieftinck (Hemicordulia), 1953c: 188—191, fig. 54—57. Holotype {. 
Sumba I. (Lesser Sunda Is.), Central Sumba, Langgaliru, 400—600 m, 4-15.X.1949 
(Sumba Exped. 1949, NMB, XII C 230). — First described 9. W. Sumba I. 
Waimangura, 436 m, 12-26.VIII.1949 (id., same coll.-no.). — Corduliidae. 

coartans Lieftinck (Argiolestes), 1956b: 69, 81—84, fig. 10, 14—15. Holotype { 
and first described 9. Waigeu I., off N.W. New Guinea, Camp Nok, 2500 ft., 
IV.1938, L. E. Cheesman (BM). See Kimmins, 1970: 198. — Megapodagrionidae. 

commutata Lieftinck (Notoneura), 1938: 90—91, fig. 23. Holotype & and first 
described 9. Bismarck Archip., New Britain, Herbertshoh, 1896 (&) and Ralum 
(2), F. Dahl (SMF); selected by Lieftinck, 1949b: 335. — Protoneuridae. 

Nom. nov. pro Caconeura exul: Ris, 1900 (3 $ New Britain, Herbertshöh), nec 

Alloneura exul Selys, 1886b. 

cyrene Lieftinck (Synthemis), 1953a: 71, 74, 78—81, fig. 5—9. Holotype { and first 
described 9. N.E. New Guinea, Saiko, 5500—6000 ft, Buba River (Upper Waria 
River), IX-X.1936, F. Shaw-Mayer (BM). — Corduliidae. 

See Kimmins, 1968: 296. 


dajakanus Lieftinck (Lestes sg. Indolestes), 1948e: 3—6, fig. 2—3. Holotype &. 
N.W. Borneo, Sarawak Distr., E. Mjöberg (NRS). — Lestidae. 

decorata Lieftinck (Oligoaeschna), 1968a: 146, 158—159, pl. 12 & tfig. 2. Holotype 9. 
Assam, Shillong, 5000 ft., 5.V.1924, Crinoline Falls, T.B. Fletcher, ex coll. F. C. 
Fraser (BM). — Aeshnidae. 

deminuta Lieftinck (Diplacodes), 1969c: 32—35, fig. 5a-d. Holotype g and first 
described 9. N.E. Zambia, Lake Bangweulu Region, Samfya, 1160 m, 13.1.1962 
(3) and id., “dembo” of Kasamba River, 1160 m, 17.1V.1963 (9 ), J. J. Symoens, 
no. 9960 (¢) and 10296 (9) (MAT). — Libellulidae. 

dendrolagina Lieftinck (Drepanosticta), 1938: 83—84, 87, fig. 19c, 20. Lectotype SG 
and first described 9. N.E. New Guinea, Astrolabe Bay, Sattelberg (Gegagalu), 900 
m, Carl Wahnes 1899, various dates, ex coll. F. Foerster (UMMZ). — Platystictidae. 

dentiplaga Lieftinck (subsp. of Rhinocypha tincta Ramb.), 1938: 69—70, fig. 1, 
3A-C, 4N, 5F. Holotype g and first described 9. N.E. New Guinea, Huongolf, 
Hänishafen, Taimi Inseln, III.1900, Carl Wahnes (UMMZ). — Chlorocyphidae. 

dione Lieftinck (Macromia), 1971b: 26—28, fig. 8—9. Holotype g'. N.E. Sumatra, 
Deli/6594, L. Martin, with label ‘“Macromia? westwoodi Selys" (unknown writing) 
(MNB). — Corduliidae. 

discrepans Lieftinck (Aethiothemis), 1969c: 26—28, fig. 4. Holotype g' and first 
described 9. Congo, Katanga, Lower Luapula Region, Kabiashia, Luanza River, 
1000 m, 23.1V.1966, F. Malaisse, no. 4155 (MAT). — Libellulidae. 


124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


dolabrata Lieftinck (Teinobasis), 1938: 114—116, fig. 47. Holotype & and allotype 
® (in cop.). N.E. New Guinea, Huongolf, Hänischhafen, Taimi Ins., III.1900, Carl 
Wahnes; nos. 2239 and 2240 of F. Förster's collection (UMMZ). — Coenagrionidae. 

donaldi Lieftinck (Lanthanusa), 1955b: 161—164, fig. 5—6. Holotype 8 and first 
described 9. N.E. New Guinea, Saiko, 5500—6000 ft., Buba River (Upper Waria 
River), IX-X.1936, D. Shaw-Mayer (BM). See Kimmins, 1968: 284. — Libellulidae. 

ducatrix Lieftinck (Devadatta), 1969a: 205—207, fig. 1—2. Holotype 4. Tonkin, 
Than Moi, VI-VII, H. Fruhstorfer, with yellow label “Heptaneura natator Martin 
Type” (nom. nud.), in R. Martin’s handwriting (MP). — Amphipterygidae. 


eduardi Lieftinck (Hemicordulia), 1953c: 185—188, fig. 52—53. Holotype dg. W. 
Sumba I. (Lesser Sunda Is.), Rara, 340 m, near Korokangali, 7.VIII.1949, A. M. R. 
Wegner (Sumba Exped. 1949, NMB XII C 229). — Corduliidae. 

elisabethae Lieftinck (Nesolestes), 1965c: 243— 244, fig. 7. Holotype &. Madagascar, 
Prov. Diégo-Suarez, Montagne d’Ambre, 24.V.1958, F. Keiser et al. (NMB). — 
Megapodagrionidae. 

elisabethae Lieftinck (Zygonyx), 1963b: 56—58, fig. 4—6. Holotype g and first 
described 9. Madagascar, Prov. Diégo-Suarez, Montagne d’Ambre, 21.V.1958 (g') 
and 23.V.1958 (9), F. Keiser et al. (NMB). — Libellulidae. 

ephippiatus Lieftinck (Argiolestes), 1956b: 72, 97— 98, fig. 35— 36. Holotype &. 
N.E. New Guinea, Kokoda, 1200 ft., IX.1933, L. E. Cheesman (BM). See Kimmins, 
1970: 197. — Megapodagrionidae. 

euphrosyne Lieftinck (Macromia), 1952a: 439, 450—454, fig. 3, 7. Holotype gd 
and first described 9. Waigeu I. (off N.W. New Guinea), Camp Nok (Mt. Buffel- 
hoorn), 2500 ft., IV.1938, L. E. Cheesman (BM). — Corduliidae. 

evelynae Lieftinck (Notoneura), 1960b: 120—122, fig. 6. Holotype g'. Waigeu I. 
(off N.W. New Guinea), Camp Nok (Mt. Buffelhoorn), 2500 ft, IV.1938, L. E. 
Cheesman (BM). See Kimmins, 1970: 177. — Protoneuridae. 

evelynae Lieftinck (Synthemis), 1953a: 71, 73, 81—84, fig. 10—11. Holotype & 
(abdominal segments 6— 10 and apps. missing) and first described ©. Papua, Mafulu, 
4000 ft., XII.1933, L. E. Cheesman (BM). See Kimmins, 1968: 296. — Corduliidae. 

exoleta Lieftinck (Coeliccia), 1961a: 140—141, fig. 49. Holotype &. Philippine 
Is, Mindanao I, Davao Prov, E-slope of Mt. McKinley, 2500 ft, 7.IX.1946, 
F. G. Werner (CNHM). — Platycnemididae. 


fickei Foerster (Protolestes), 1899b (9). First described 4. N. Madagascar, Prov. 
Diégo-Suarez, Montagne d’Ambre, 24.V.1958, F. Keiser et al. (NMB & ML). See 
Lieftinck, 1965c: 244— 246, fig. 9. — Megapodagrionidae. 

floresiana Lieftinck (Drepanosticta), 1939a: 147—149, fig. 2. Holotype & (head and 
prothorax missing). Flores I., labelled “Flores” in R. Martin's handwriting (MP). 
First described 9, same label (ML). — Platystictidae. 

fontinalis Lieftinck (Drepanosticta), 1937b: 64—67, fig. 5—6. Holotype g'. Malay 
Peninsula. Two labels in F. Förster’s handwriting: “Kelantan Ost Malacca Heine 
vdt. 1903/Platysticta quadrata kelantana n. rasse Forster” [nom. in litt.} (UMMZ). 
First described 9. Malay Peninsula, Selangor, Templer Park, 12 mi. from Kuala 
Lumpur, forest seepage near Sungai Chui Tinggi, 21—23.111.1963 (3 2), M. A. 
Lieftinck (ML). See Lieftinck, 1965a: 181—184. — Platystictidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 125 


tortis Lieftinck (Teinobasis), 1962: 20, 26—29, fig. 7. Holotype 8 and first des- 
cribed 9. E. Caroline Is. (Micronesia), Ponape I., Mt. Nahnalaud, VI-IX.1950, 
Adams (USNM 65139). — Coenagrionidae. 
fraseri Lieftinck (Drepanosticta), 1955e: 70—72, fig. 1. Lectotype g'. Ceylon, Central 
Prov., Kandy, 2.X1.1953, F. Keiser et al. (NMB). — Platystictidae. 
fraterna Lieftinck (Notoneura), 1933c: 410—413, fig. 1. Lectotype 4 and first 
described 9. N. Australia, N.T., Z.-Lagoon, 19.1V.1931, Ed. Handschin (NMB). — 
Protoneuridae. 
frontalis Lieftinck (Eusynthemis), 1949b: 359—361, fig. 12. Holotype 9 (immature). 
Solomon Is., Guadalcanal I., XII. 1921. J. A. Kusche (CAS). — Corduliidae. 
fruhstorferi Lieftinck (subsp. of Onychogomphus modestus Selys), 1934b: 34—36, 
fig. 7. Holotype &. With labels “Java Fr [uhstorfer} g'” and “Onychogomphus 
Fruhstorferi § Java”, in de Selys’ handwriting (IRSN). — Gomphidae. Plate 4. 
First discussed 9. S. Sumatra, Lampong distr., foot of Mt. Tanggamus, Giesting, 
500 m, 7.VII.1934, L. J. Toxopeus (ML). 
Now considered a distinct species (see Lieftinck, 1948e: 23). 


gamblesi Lieftinck (Lokia), 1969c: 16—18, fig. 2. Holotype &. N.E. Zambia, Middle 
Luapula Region, Kisongo, swampy ‘“dembo” of Loshi River, 1190 m, 24.X11.1962, 
J. J. Symoens, no. 9913 (MAT). — Libellulidae. 

gracilis Belle (Epigomphus), 1970: 12—15, fig. 20—24. Holotype 2. Brazil, Mirituba, 
Tapájos, Amazon, IV.1921, A. H. Fassl, cat. no. 15122 (SMF). — Gomphidae. 

guamensis Lieftinck [subsp. of Agrionoptera insignis (Ramb.)}, 1962: 71, 73, fig. 
22c-d. Holotype & and first described 9. S. Mariana Is. (Micronesia), Guam I., 
V.1945, Bohart & Gressitt (J USNM 65777, 9 USNM). — Libellulidae. 

gurneyi Lieftinck (Papuagrion), 1949b: 341— 343, fig. 5. Holotype 4. Solomon Is. 
Bougainville I., 6.VIII-16.XI.1944 (no precise date), A. B. Gurney (USNM). 
— Coenagrionidae. 


handschini Lieftinck (Isosticta), 1933c: 417—420, fig. 3—4. Lectotype g' and first 
described 9 (juv.). N. Australia, N.T., Burnside, 21.1V.1932 (g') and Kadarri, 
20.1V.1931 (2), Ed. Handschin (NMB). — Isostictidae. 


icterica Lieftinck (Macromia), 1929a. 64, 84—86, fig. 11—12. Holotype g'. China, 
Canton (no further particulars), ex coll. K. J. Morton (RSM). — Corduliidae. 
ierea Geijskes (Argia), 1932: 244—248, fig. 6—9. Lectotype d'. Trinidad I., Sangre 
Grande, 16.1.1930, G. Belmontes (ROMT). — Coenagrionidae. 
Synonymous with Argia insipida Hagen in Selys. 
imitans Lieftinck (Calicnemia), 1948e: 12—15, fig. 7. Holotype 4 and first described 
®. S. Burma, Tenasserim, Malvedaung, 30 km S. of Ye, 300 m, 17.X1.1934, R. 
Malaise (NRS). — Platycnemididae. 
inaequidens Lieftinck (Idiocnemis), 1932b: 503—504, fig. 7—8. Holotype 4. N. 
Guinea Bird 1900, Stephansort Astrolabe Bai (printed label), ”Idiocnemis bident.” 
(F. Forster's handwriting) (MBUD). — Platycnemididae. 
First described 9. N.E. New Guinea, Bongu, Urwald Astrolabebai, Wahnes leg., 
Sommer 1899 (F. Forster's handwriting) (MBUD; ML). 
inaequistigma Fraser (Millotagrion), 1953 (6). First described 9. E. Madagascar, 


126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Prov. Tamatave, Moramanga, 9 km south, 22.X11.1957, F. Keiser et al. (NMB). See 
Lieftinck, 1965c: 251—253, fig. 11. — Coenagrionidae. 
Type-species of Millotagrion Fraser (op. cit.). 

incisurum Lieftinck (Pseudagrion), 1949b: 336—338, fig. 2. Holotype g'. Solomon 
Is., Guadalcanal I., XII.1920-1.1921, J. A. Kusche (CAS). — Coenagrionidae. 

inconspicua Lieftinck (Drepanosticta), 1938: 85—86, 87, fig. 21. Holotype g'. Waigeu 
I., off N.W. New Guinea, labelled ’’Waigiu”, probably ex Staudinger (SMF). — 
Platystictidae. 

ixias Lieftinck [subsp. of Rhyothemis phyllis (Sulz.) }, 1953c: 221—223. Holotype ¢ 
and first described ©. Sumba I. (Lesser Sunda Is.), E. Sumba, Melolo, sea-level, 
23.V.-8.VI.1949 (Sumba Exped. 1949, NMB, XII C 342). — Libellulidae. 

johnseni Lieftinck (Burmagomphus), 1966a: 193—197, fig. 1—5. Holotype g'. Thai- 
land, N.W., Kwae Noi area, Ban Kao, 13-18.X1.1961, P. Johnsen, no. 1286 (MC). 
— Gomphidae. 

junghuhni Lieftinck (Megalogomphus), 1934c: 266—267, fig. 2. Holotype 9. With 
two labels: “Java, Heyne”, and “Heterogomphus près cochinchinensis Selys 9”, 
the latter in de Selys’ handwriting (IRSN). — Gomphidae. 


keiseri Lieftinck (subsp. of Macrogomphus annulatus Selys), 1955e: 81—83, fig. 4. 
Holotype g. Ceylon, Central Prov., Weragamtota, 14.1X.1953, “am Licht”, F. 
Keiser et al. (NMB). — Gomphidae. 

khasiana Lieftinck (Oligoaeschna), 1968a: 146, 156—158, fig. 2—3. Holotype g'. 
Assam, Khasia Hills, in R. MacLachlan’s writing, ex coll. MacLachlan (BM). — 
Aeshnidae. 

kimminsi Lieftinck (Idiocnemis), 1958: 270—272, 284, fig. 56—60. Holotype ¢ 
and first described 9. New Britain (Bismarck Archip.), A. Willey, Reg. Mar. 1, 
1898, nos. 98 & 73, both with label “Idiocnemis nov. spec.”, in F. Ris’ handwriting 
(BM). See Kimmins, 1970: 180. — Platycnemididae. 

kimminsi Lieftinck (Synthemis), 1953a: 71, 73, 75—78, fig. 1—4. Holotype g and 
first described 9. N.E. New Guinea, Saiko, 5500—6000 ft, Buba River (Upper 
Waria River), 24.X.1936, F. Shaw-Mayer (BM). See Kimmins, 1968: 298. — 
Corduliidae. 

kirbyi Foerster (Wahnesia), 1900: 105 (no descr., hab. Bongu, N.E. New Guinea). 

See kirbyi Lieftinck, next item. 

kirbyi Lieftinck (Argiolestes), 1935c: 217, 228—229, fig. 11—12. Lectotype g. N. 
Guinea Biró 1899 Sattelberg, Huon Golf, 22.11.1899 (printed labels), Wahnesia 
Kirbyi, in Dr. Pongracz’ handwriting (MBUD). — Megapodagrionidae. 

Although no specific description of A. kirbyz was given by Förster, the name was 
validly proposed and should be attributed to Förster, not Lieftinck. See Förster, 1900: 
105 (Wahnesia n.g. with types W. Kirbyi n.sp. and montivagans n.sp., in MBUD, 
both nom. nud.; and Lieftinck (1938: 78). 

Correctly classified as A. kirbyz (Foerster). 


laidlawi Lieftinck (Burmargiolestes), 1960a: 236—237. Type series (3 g 1 Q from 

N. India, Darjeeling distr., and 2 & 1 9 from Gopaldhara, same distr.) ex Indian 
Museum, Calcutta (?), possibly destroyed. 

Nom. nov. pro Argiolestes melanothorax: Laidlaw (1917: 323—325, pl. 15 fig. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 127 


1, & Darjeeling), nec A. melanothorax Selys, 1891a (Burma). 
Type-species of Burmargiolestes Kennedy, 1925, who founded the genus on the 
Indian species (see Lieftinck, op. cit.). 
leonorae Lieftinck (Idiocnemis), 1949a: 95—96, fig. 106—107. Holotype g'. NE. 
New Guinea, Huon Gulf, “Bidung [or Bulung?} gebiet, Ogeramnang, 1500—2000 
m, 1914” (F. Forster's handwriting), No. 2499 of the Förster collection (UMMZ). 
—Platycnemididae. 
leonorae Lieftinck (Teinobasis), 1937b: 97—99, fig. 22. Holotype g'. Malay Peninsula, 
Penang I., Staudinger vend., ex coll. F. Förster (UMMZ). — Coenagrionidae. 
Synonymous with T. rajah Laidlaw, 1912. 
liberata Lieftinck (Tramea), 1949b: 371—372. Holotype &. Solomon Is., Guadalcanal 
I, XII.1920, J. A. Kusche (CAS). — Libellulidae. 
Now Trapezostigma liberata (Lieftinck). 
malaisei Lieftinck (Rhipidolestes), 1948e: 7—9, fig. 4—5. Holotype g'. N.E. Burma, 
Kambaiti, 2000 m, 28.V.1934, R. Malaise (NRS). — Megapodagrionidae. 
manicaria Williamson (Metaleptobasts), 1915 (g'). First described 9. Trinidad, Sangre 
Grande, 5.111.1930, G. Belmontes (ROMT). See Geijskes, 1932: 260—262, fig. 
17—28. — Coenagrionidae. 
marsyas Lieftinck (Drepanosticta), 1965a: 173, 178—180, fig. 3—7. Holotype {. 
Malay Peninsula, Pahang, Cameron’s Highlands, 4—5000 ft, 15.VI.1935, H. M. 
Pendlebury 1937—472 (ex F. M. S. Mus.), with red type-disk and “Drepanosticta 
marsyas g' Type” {nom. in litt.} in F. F. Laidlaw’s handwriting (BM). See Kimmins, 
1970: 174 (the month of capture should be June, not April). — Platystictidae. 
martini Ris (Argiocnemis), 1900 (8). First described 9 (imperfect). Admiralitäts 
Inseln [Admiralty Is.}, N.-Küste, Seeadler Hafen, Papitalei, No. 140 der Hamburg 
Südsee Expedition, 18-20.X.1908, G. Duncker (ZMH). See Lieftinck, 1949b: 347. 
— Coenagrionidae. 
Now Mortonagrion martini (Ris). 
mauritia Williamson (Metaleptobasis), 1915 (3). First described 9. Trinidad, Sangre 
Grande, 26.1.1930, G. Belmontes (ROMT). See Geijskes, 1932: 258—260, fig. 
15—16. — Coenagrionidae. 
melidora Lieftinck (Palaiargia), 1953e: 241—244, fig. 4. Holotype 4 and first des- 
cribed 9. Waigeu I. (off N.W. New Guinea), Camp Nok, 2500 ft, III-IV.1938, 
L. E. Cheesman. Holotype & with collector's note: “lateral stripe of thorax pale 
gr(een), dorsally blue, mac(ulae) of abdomen blue” (BM). See also Kimmins, 1970: 
188. — Coenagrionidae. 
merina Lieftinck (Agriocnemis), 1965c: 254—255, fig. 12. Holotype 4. Madagascar, 
Prov. Tamatave, Fampanambo, 19.X1.1958, F. Keiser et al. (NMB). — Coenagrion- 
idae. 
microstigma Lieftinck (Argiolestes), 1956b: 72, 99— 101, fig. 40—41. Holotype g'. 
Papua, Mafulu, 4000 ft, 1.1934, L. E. Cheesman (BM). See Kimmins, 1970: 197. — 
Megapodagrionidae. 
mima Lieftinck (Libellago), 1932a: 2—4, fig. 1. Holotype g'. Borneo, with labels 
“63/mimus Borneo” (yellow) “Micr. mimus S. ¢° Borneo” (white), the last two in 
de Selys’ writing; L. mima Lieft., holotype, det. M.A.L. (IRSN). — Chlorocyphidae. 
A synonym of L. semiopaca (Selys). 


128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


montivagans Foerster (Wahnesia), 1900: 105 (no descr., hab. Bongu, N. E. New 
Guinea). 
See next item (same species). 
montivagans Lieftinck (Argiolestes), 1935c: 204, 214—217, 233—234 (sub A. 
sidonia Martin). Lectotype & and first described 9. N. Guinea Bird 1899 Simbang 
Huon Golf, 18.11.1899 (print), Wahnesia montivagans, in Dr. Pongracz’ writing 
(MBUD). 
See explanation sub A. kirby? Lieft. (huj. op.). 
Currently classified as A. montivagans (Foerster). 


nasiterna Lieftinck (Palaiargia), 1938: 99—100, fig. 34—35. Holotype g'. Waigeu 
I., off N.W. New Guinea, ex Staudinger (SMF). — Coenagrionidae. 

nigrolutea Lieftinck (Teinobasis), 1962: 19, 29—31, fig. 8. Holotype ¢ and first 
described @. E. Caroline Is. (Micronesia), Ponape I., S. of Nanpohnmal, 17.1.1953, 
Clarke (& USNM 65140). — Coenagrionidae. 


occipitalis Belle (Epigomphus), 1970: 15—18, fig. 27—36. Holotype g and first 
described 9. Perú, Mishuyacu, Iquitos, Amazon, 16.VII.1930 (4) and 26.111.1930 
(2), Klug, cat. no. 15116 and 15118, respectively (SMF). — Gomphidae. 

ochrostomus Lieftinck (Argiolestes), 1949a: 48—49, fig. 14—16, 25. Holotype & 
and first described 9. “Waigiu [Waigeu I., off N.W. New Guinea], Staudinger 
vend., ex coll. F. Ris (SMF). — Megapodagrionidae. 

olivaceus Lieftinck (Heliogomphus), 1961a: 143—145, fig. 50. Holotype g'. Philip- 
pine Is., Palawan I. group, Busuanga I. (Calamianes), Dimaniang, near sea level, 
III.1947, H. Hoogstraal (CNHM). — Gomphidae. 

orchestra Lieftinck (Idionyx), 1953c: 193—196, fig. 65—67. Holotype g and first 
described 9. Sumba I. (Lesser Sunda Is.), W. Sumba, Waimangura, 436 m, 12-26. 
VIII.1949 (Sumba Exped. 1949, NMB XII C 241). — Corduliidae. 

orestes Lieftinck (Gomphus), 1939b: 285, 287— 290, fig. 5— 6. Holotype g and first 
described 9. E. China, Fu Kien, near Kwa Tun, 2300 m, 8.VI.1938 (g') and 
19.V.1938 (2), J. Klapperich (MKB). — Gomphidae. 

Transferred to genus Sinogomphus May. 

oryzae Lieftinck (subsp. of Ischnura aurora Brauer), 1962: 44, fig. 13. Holotype & 
and first described 9. Ryukyu Is., Okinawa I., Chizuka, VII. 1945, Bohart & 
Harnage (& USNM 65143, 9 USNM). — Coenagrionidae. 


palauense Lieftinck (Pseudagrion), 1962: 36—38, fig. 10. Holotype g'. W. Caroline 
Is. (Micronesia), Palau Is., Babelthuap I., Melekeiok-Ngardok Lake, 24.11.1936, 
Esaki (KU). — Coenagrionidae. 

palauensis Lieftinck (Drepanosticta), 1962: 14—16, fig. 3. Holotype g' and allotype 
® (in cop.). W. Caroline Is. (Micronesia), Palau Is., Babelthuap, jungle 2 mi. 
N.W. of Ngiwal, 21.V.1957, Sabrosky (USNM 65135). — Platystictidae. 

palauensis Lieftinck (Teinobasis), 1962: 19, 21, fig. 4—5. Holotype g'. W. Caroline 
Is. (Micronesia), Palau Is., Koror I., 15-24.111.1948, Maehler (USNM 65136). 
— Coenagrionidae. 

palawana Lieftinck (Prodasineura), 1948a: 232—234, fig. 4. Holotype g'. Palawan 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 129 


I., labelled “Palawan, Fruhst(orfer)”, in de Selys’ handwriting (IRSN). — Proto- 
neuridae. 
pallida Belle (Phyllocycla), 1970: 109—111, fig. 168—170. Holotype d'. Brazil, 
Santa Catarina (Nova Teutonia), 4.XII.1949; ex coll. K. H. Buchholz (MKB). — 
Gomphidae. 
pechumani Belle (Epigomphus), 1970: 19—20, fig. 3741. Holotype g'. Colombia 
(CUI). — Gomphidae. 
peleus Lieftinck (Gomphus), 1939b: 285, 291—294, fig. 9—10. Holotype &. E. 
China, Fu Kien, near Kwa Tun, 2300 m, 31.V.1938, J. Klapperich (MKB). — 
Gomphidae. 
A topotypical 4 paratype was taken I.VI.1938 by the same collector (ML). 
Now placed as Sinogomphus peleus (Lieftinck). 
petalura Lieftinck (Oligoaeschna), 1968a: 150, 175—176, fig. 8. Holotype g'. Hainan 
I, Mt. Wuchi, 24.V.1903, 1911—288, Jagoria sp., det. D. E. Kimmins (BM). — 
Aeshnidae. 
philippa Lieftinck (Drepanosticta), 1961a: 132—133, fig. 45B, 46A-C. Holotype gd 
and first described @. Philippine Is., Luzon, Abra prov., Massisiat, creek in mountain 
gully, 21.V.1946, H. Hoogstraal (CNHM). — Platystictidae. 
ponapensis Lieftinck (Teinobasis), 1962: 20, 24—26, fig. 7. Holotype g'. E. Caroline 
Is. (Micronesia), Ponape I., Mt. Ngihneni, 730 m, VI-IX.1950, Adams (USNM 
65138). — Coenagrionidae. 
prometheus Lieftinck (Gomphus), 1939b: 278—281, fig. 1—2. Holotype 8 and 
first described 9. E. China, Fu Kien, near Kwa Tun, 2300 m, 5.VI.1938 (4) and 
25.VI.1938 (2), J. Klapperich (MKB). — Gomphidae. 
A topotypical & paratype was taken the same day as the holotype (ML). 
Type-species of Fukienogomphus H. F. Chao. 
proselytus Lieftinck (Protolestes), 1965c: 246. Lectotype &. E. Madagascar, Ambodi- 
rafia, XI, Olsufieff. — Megapodagrionidae. 

This is one of the specimens described and figured by the late Erich Schmidt as 
Protolestes Fickei Foerster, which is a different species. See Schmidt, 1951: 147—150, 
fig. 22a, 23c, 24, 26a-b; and Lieftinck, op. cit., who proposed the new name. 

The lectotype (ex coll. E. Schmidt) is at present in Dr. S. Asahina’s collection 
(Tokyo). 

prothoracalis Lieftinck (Argiolestes), 1956b: 72, 98—99, fig. 33—34. Holotype ¢. 
N.E. New Guinea, Kokoda, 1200 ft, VI.1933, L. E. Cheesman (BM). See Kimmins, 
1970: 197. (The original locality is wrongly given as in “Northern Papua”). — 
Megapodagrionidae. 

pulverulans Lieftinck (Nesolestes), 1965c: 239240, fig. 3—5. Lectotype ¢ and first 
described 9. Madagascar, Prov. Fianarantsoa, Ranomafana, 22.1.1958, F. Keiser et al. 
NMB). — Megapodagrionidae. 

pusilla Lieftinck (subsp. of Agrionoptera sanguinolenta Lieft.), 1962: 71, 76—77, fig. 
20c, 21g, 22k-l. Holotype 4 and first described 9. Caroline Is. (Micronesia), Truk 
I., Wena I. (Moen), Nantuka (Civil Adm. Area), 31.1.-27.1V.1949, Potts (& USNM 
65147, 9 USNM). — Libellulidae. 

pylades Lieftinck (Gomphus), 1939b: 285—287, fig. 3—4. Holotype g'. E. China, 
Fu Kien, near Kwa Tun, 2300 m, 24.V.1938, J. Klapperich (MKB). — Gomphidae. 

Transferred to Sinogomphus May. 


130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


pytho Lieftinck (Drepanosticta), 1937b: 68—70, fig. 7. Holotype g'. W. Sumatra, 
Padang 1913 (?Rolle vend.), labelled on paper triangle “Platysticta Grosskopf, W. 
Sumatra, Padang’, in F. Forster's handwriting (UMMZ, coll. no. 1252). — 
Platystictidae. 


radama Lieftinck (Nesolestes), 1965c: 241—243, fig. 8. Holotype g and first des- 
cribed ©. Madagascar, Prov. Tananarive, Moramanga, 9.X.1958 (g') and Prov. 
Tamatave, Périnet, 4.XI1.1957 (© ), F. Keiser et al. (NMB). — Megapodagrionidae. 

ramajana Lieftinck (Disparoneura), 1971a: 194, fig. 3—4. Holotype g (immature). 
Ceylon, Central Prov., Horton Plains, 7000 ft, 12 mi. S.S.E. of Nuwara Eliya, loc. 
163, 19.111.1962, Lund Univ. Ceylon Exped. 1962 (ZIUL). — Protoneuridae. 


sanguinolenta Lieftinck (Agrionoptera), 1962: 71, 74—75, fig. 20c-d, 21g-h, 22i-l. 
Holotype & and first described 9. E. Caroline Is. (Micronesia), Ponape I., Mt. 
Paipalap, 240 m, VI-IX.1950 (g') and Agric. Exper. Sta., VI-IX.1950 (© ), Adams 
(& USNM 65146, 9 USNM). Libellulidae. 

Now A. sanguinolenta sanguinolenta Lieft. 

sanguinolenta Lieftinck (Rhinocypha), 1961a: 125—128, fig. 44a. Holotype g'. S. 
Philippine Is., Mindanao I., Davao Prov., E.-slope of Mt. McKinley, 2500 ft., 7.IX. 
1946, F. G. Werner (CNHM). — Chlorocyphidae. 

schroederi Belle (Cyanogomphus), 1970: 27—29, fig. 48—51. Holotype g'. Brazil, 
Tapajos, Amazon, V.1920, A. H. Fassl, coll. no. 15235 (SMF). — Gomphidae. 

septima Martin (Macromia), 1904 ( 9 ). First described &. Java, Fr[uhstorfer} [18 }93 
(IRSN). See Lieftinck, 1929a: 67, 100—101, fig. 19—20. — Corduliidae. 

simonae Lieftinck (Tanymecosticta), 1969a: 208—210, fig. 3—5. Holotype g'. Kei Is., 
labelled “Protosticta? Platysticta auriculata? Key” (pink label in R. Martin’s hand- 
writing) (MP). — Isostictidae. 

sinhalensis Lieftinck (Drepanosticta), 1971a: 192— 193, fig. 1. Holotype g'. Ceylon, 
Prov. Sabaragamuwa, Deerwood, Kuruwita, N.N.W. of Ratnapura, loc. 90III, 18- 
21.11.1962, Lund Univ. Ceylon Exped. 1962 (ZIUL). — Platystictidae. 

solitaris Lieftinck (?Anisogomphus), 1971a: 199—203, fig. 5 & 9. Holotype & (im- 
mature, with exuviae). Ceylon, Central Prov., Rambukpath Oya, 10 mi. N.W. of 
Hatton, loc. 153, 18.11.1962, Lund Univ. Ceylon Exped. 1962 (ZIUL). — Gom- 
phidae. 

sophrosyne Lieftinck (Macromia), 1952a: 442, 449— 450, fig. 1, 8. Holotype &. 
Waigeu I. (off N.W. New Guinea), Camp Nok (Mt. Buffelhoorn), 2500 ft., 
IV.1938, L. E. Cheesman (BM). — Corduliidae. 

sutteri Lieftinck (Lestes sg. Indolestes), 1953c: 142—145, fig. 3, 6. Holotype & 
and first described 9. Sumba I. (Lesser Sunda Is.), Central Sumba, Langgaliru. 
400—600 m, 4-15.X.1949 (Sumba Exped. 1949, NMB XIIB 38). — Lestidae. 

symoensi Lieftinck (Phyllogomphus), 1969c: 11—13, fig. 1. Holotype 9. Congo, 
Katanga, Kafubu Region, Lubumbashi (Elisabethville), 1300 m, 28.XII.1960, J. J. 
Symoens, no. 8026 (MAT). — Gomphidae. 


telamon Lieftinck (Gomphus), 1939b: 285, 290—291, fig. 7—8. Lectotype g'. E. 
China, Fu Kien, near Kwa Tun, 2300 m, 30.V.1938, J. Klapperich (MKB). — 
Gomphidae. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 131 


A topotypical & paratype was taken the same day (ML). 
Transferred to genus S‘nogomphus May. 
thalia Lieftinck (Macromia), 1929a: 67, 103—104, fig. 21. Holotype 8. Assam, 
Khasia Hills; ex coll. F. F. Laidlaw (BM). See Kimmins, 1968: 300. — Corduliidae. 
thelmae Lieftinck (Ischnura), 1966b: 92—96, fig. 1. Holotype g and first described 
@. South Pacific: Rapa I. (Oparu), Rapa Mati Bay, 23.X.1963 (¢, USNM, reg. 
no. 68921) and Rapa Haurei, 3.XII.1963 (9, USNM), J. F. Gates Clarke & Mrs. 
Thelma M. Clarke. — Coenagrionidae. 
thelyphonus Lieftinck (Microgomphus), 1929b: 125—128, 130, fig. 17—20. Holotype 
d' and first described ©. Java mer., H. Fruhstorfer, labelled ‘““Microg. race de chelifer, 
à étudier” (4) and “Microgomphus Fruhstorferi S”, in de Selys’ handwriting. 
Types selected by M.A.L. in description and labelled accordingly, from a total of 
2 g and 1 9 (IRSN). — Gomphidae. Plate 4. 
Currently known as Microgomphus chelifer thelyphonus Lieft. 
thisbe Lieftinck [ subsp. of Rhyothemis regia (Brauer) }, 1953c: 223—226. Holotype gd 
and first described 9. Sumba I. (Lesser Sunda Is.), E. Sumba, Rende Wai, 100 m, 
11-16.VI.1949 (Sumba Exped. 1949, NMB, XII C 346). — Libellulidae. 
timorana Lieftinck (subsp. of Rhinocypha pagenstecheri Foerster), 1936a: 111—114, 
fig. 3. Lectotype & and first described @. S. Timor I. (Lesser Sunda Is.), Kupang, 
XI1.1931-1.1932, Ed. Handschin (NMB). — Chlorocyphidae. 
tincta Rambur (Rhinocypha), 1842 (&). First described 9. Waigeou (lavender-coloured 
label in de Selys’ writing) [ Waigeu I., off N.W. New Guinea} (IRSN). See Lieftinck, 
1938: 63—64, fig. 1. — Chlorocyphidae. 
Now R. tincta tincta (Ramb.). 
tonkinicus Fraser (Onychogomphus), 1926 (g'). First described ©. Tonkin, Than-Moi, 
VI-VII, H. Fruhstorfer (print), with two labels “Heterogomphus naninus Foerst. ©”, 
and “Onychogomphus naninus Foerster @ Type”, both in F. Förster's handwriting 
(UMMZ). 
Now Phaenandrogomphus tonkinicus (Fraser), see Lieftinck, 1969a: 210—214, 
fig. 13—14. 
trimaculata Lieftinck (Drepanosticta), 1939a: 149—151, fig. 3. Holotype g'. Philip- 
pine Is., Luzon, Bilucao, 1876, L. Laglaize, with label in R. Martin’s handwriting 
“Platysticta, près bicornuta ou bicornuta” (MP). — Platystictidae. 
turconu Selys (Rhinocypha), 1891b: 215—216 (J'). First described 9. Philippine 
Is, Mindanao I., Davao Prov., east slope of Mt. McKinley, 3000 ft, 22.1X.1946 
(32), F. G. Werner (CNHM). See Lieftinck, 1961a: 122—124. — Chlorocyphidae. 


walkeri Geijskes (Oligoclada), 1931: 213—214. Holotype g. Trinidad I. Sangre 
Grande, 26.111.1930, G. Belmontes (ROMT). — Libellulidae. 

werneri Lieftinck (Coeliccia), 1961a: 137—138, fig. 48. Holotype 4. Philippine Is. 
Palawan I., Iwahig, mountains W. of Lapulapu, 2—3000 ft., 1-2.111.1947, F. G. 
Werner (CNHM). — Platycnemididae. 


yapensis Lieftinck [subsp. of Agrionoptera insignis (Ramb.)}, 1962: 71, 72—73, fig. 
20a, 21e, 22a-b. Holotype & and first described 9. W. Caroline Is. (Micronesia), 
Yap I, Ruul district, VII-VIII.1950, Goss (4 USNM 65144, 9 USNM). — 
Libellulidae. 


132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


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M 


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134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


—. 1934. Notes on some Oriental dragonflies (Odonata), with description of a new species. 
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——, 1929a. A revision of the known Malaysian dragonflies of the genus Macromia Rambur, 
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—, 1929b. Contributions to the dragonfly fauna of the Sondaic Area. I. — Tijdschr. Ent. 72: 
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——, 1930b. Contributions to the dragonfly-fauna of the Dutch East Indies. II. — Treubia 12: 
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——., 1931. A revision of the genus Epophthalmia Burm. (Odon., Corduliinae). — Treubia 13: 

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, 1932a. Notes on the genus Libellago Selys, with descriptions of two new species (Odon.). 
— Konowia 11: 1—11, 1 fig. 

——, 1932b. The dragonflies (Odonata) of New Guinea and neighbouring islands. Pt. 1. 
Descriptions of new genera and species of the families Lestidae and Agrionidae. — Nova 
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, 1932c. Two new species of Odonata from Java. — Stylops 1: 248—253, 3 fig. 

— _, 1933a. A synopsis of the Bornean Platystictinae, with descriptions of three new species 

(Odon.). — Konowia 11 (1932): 281—297, 4 fig. (Issued March 15, 1933). 

———, 1933b. A new genus and species of Libellulinae from Western Borneo (Odonata). — 
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—, 1933c. Odonaten aus Nordaustralien. Prof. Dr. E. Handschin Studienreise auf den Sun- 
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——, 1933d. The dragonflies (Odonata) of New Guinea and neighbouring islands. Pt. II. 
Descriptions of a new genus and species of Platycneminae (Agrionidae) and of new 
Libellulidae. — Nova Guinea 17 Zool. 1: 1—67, 35 fig. 


—., 1934a. Descriptions of five new species of Agrionidae from Java (Odonata). — Stylops 
3: 6—17, 7 fig. 

——, 1934b. Notes on a few Gomphidae from the Indo-Australian Archipelago, with descriptions 
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, 1934c. New species and races of Javanese Odonata. — Stylops 3: 265—270, 3 fig. 
——, 1934d. Notes on the genus Drepanosticta Laid., with descriptions of the larva and of 
new Malaysian species (Odon., Zygoptera). — Treubia 14: 463—476, 5 fig., pl. 9 & 10. 

——, 1934e. An annotated list of the Odonata of Java, with notes on their distribution, habits 

and life-history. — Treubia 14: 377—462, 1 map. 

——, 1935a. New and little known Odonata of the Oriental and Australian regions. — Treubia 

15 PO UI (aes, 
——, 1935b. A synopsis of the Odonata (dragonflies) of Sumatra. — Misc. Zool. Sum. (Medan) 
92—93: 1—23. 

——, 1935c. The dragonflies (Odonata) of New Guinea and neighbouring islands. Pt. III. 
Descriptions of new and little known species of the families Megapodagrionidae, Agrion- 
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Synthemis and Procordulia). — Nova Guinea, 17 Zool. 2: 203—300, 50 fig. 

, 1936a. Die Odonaten der Kleinen Sunda-Inseln, Prof. Dr. E. Handschin Studienreise auf 
den Sundainseln und in Nordaustralien 1930—32. — Rev. Suisse Zool. 43: 99—160, 
19 fig. 

, 1936b. A new genus and species of Libellulidae from Celebes (Odon.). — Treubia 15: 
399— 403, 2 fig. 


vo 


v 


vu 


M 


v 


vo 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 135 


1936c. A synonymic note on Celebothemis delecollei Ris (Odon.). — Treubia 15: 403. 
1937a. The dragonflies (Odonata) of New Guinea and neighbouring islands. Pt. IV. 
Descriptions of new and little known species of the families Agrionidae (sens. lat.), 
Libellulidae and Aeshnidae (genera Idiocnemis, Notoneura, Papuagrion, Teinobasis, 
Aciagrion, Bironides, Agyrtacantha, Plattycantha, and Oreaeschna). — Nova Guinea, new 
ser. 1: 1—82, 47 fig. 

1937b. Descriptions and records of South-east Asiatic Odonata. — Treubia 16: 55—119, 
32 fig. 

1938. The dragonflies (Odonata) of New Guinea and neighbouring islands. Pt. V. 
Descriptions of new and little known species of the families Libellaginidae, Megapod- 
agrionidae, Agrionidae (sens. lat.) and Libellulidae (genera Rhinocypha, Argiolestes, 
Drepanosticta, Notoneura, Palaiargia, Papuagrion, Teinobasis, Nannophlebia, Synthemis 
and Anacordulia). — Nova Guinea new ser. 2: 47—128, 52 fig. 

1939a. Sur quatre espèces nouvelles de Platystictidae de l'Inde orientale (Odon). — 
Revue francaise d’Ent. 6: 144—154, 4 fig. 

1939b. Six new species of Gomphus from China (Odon.). — Temminckia 4: 277—297, 
12 fig. 

1940a. On some Odonata collected in Ceylon, with descriptions of new species and larvae. 
— Ceylon J. Sci. (B) 22: 79—117, 11 fig., 1 pl. 

1940b. Descriptions and records of South-east Asiatic Odonata (II). — Treubia 17: 337— 
390, 18 fig. 

1940c. Revisional notes on some species of Copera Kirby. With notes on habits and larvae 
(Odon., Platycneminidae). — Treubia 17: 281—306, 5 fig, 5 pl. 

1941a. Une espèce nouvelle de Macromia du Tonkin (Odon.). — Revue francaise d’Ent. 
8: 94—98, 2 fig. 

1941b. Studies on Oriental Gomphidae (Odon.) with descriptions of new or interesting 
larvae. — Treubia 18: 233—253, 7 pl. 

1942. The dragonflies (Odonata) of New Guinea and neighbouring islands. Pt. VI. 
Results of the Third Archbold Expedition 1938—39 and of the Le Roux Expedition 1939 
to Netherlands New Guinea (I. Anisoptera). — Treubia 18: 441—608, 2 fig., 19 pl. 
1947. Two interesting new insular Rhinocypha from Malaysia (Odon.). — Tijdschr. Ent. 
88 (1945) 1947: 215—224, 5 fig., 2 pl. 

1948a. Descriptions and records of South-east Asiatic Odonata. — Treubia 19: 221—278, 
19 fig. 

1948b. The Odonata of Engano, with a survey of the dragonfly-fauna of the West Sumatran 
chain of islands. — Treubia 19: 279—304, 8 fig., 1 pl. 

1948c. Some species of Gynacantha from Celebes and the Moluccas, with a key and 
description of two new species (Odonata). — Treubia 19: 417—428. 

1948d. A new species of Anisogomphus Selys, with notes on A. m-flavum (Selys) 
(Odon.). — Opusc. Entom. (Lund): 59—63, 3 fig. 

1948e. Entomological results from the Swedish Expedition 1934 to Burma and British 
India. Odonata. — Arkiv f. Zool. 41A, 10: 1—23, 8 fig., 1 map. 

1949a. The dragonflies (Odonata) of New Guinea and neighbouring islands. Pt. VII. 
Results of the Third Archbold Expedition 1938—39 and of the Le Roux Expedition 1939 
to Netherlands New Guinea (II. Zygoptera). — Nova Guinea new ser. 5: 1—271, 355 
fig., 1 map. 

1949b. Synopsis of the Odonate fauna of the Bismarck Archipelago and the Solomon 
Islands. —Treubia 20: 319—374, 12 fig., 1 map. 

1950a. Additions to the Odonate fauna of South East Asia, with description of two new 
genera and three new species. — Treubia 20: 631—645, 9 fig. 

1950b. Further studies on Southeast Asiatic species of Macromia Rambur, with notes 
on their ecology, habits and life history and with descriptions of larvae and two new 
species (Odon., Epophthalmiinae). — Treubia 20: 657—716, 61 fig. 

1950c. Two new species of. Podolestes Selys from Borneo, with a key for the identification 
of the known species (Odon., Megapodagriidae). — Zool. Meded. Leiden 31: 39—47, 
4 fig. 

1951a. Notes on Malaysian Prodasineura with descriptions of two new species from 


136 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Borneo and a key to the blue-coloured species (Odonata). — Idea (Bogor) 8: 74—83, 
16 fig. 

1951b. Results of the Archbold Expeditions. No. 64. Odonata of the 1948 Archbold Cape 
York Expedition, with a list of the dragonflies from the Peninsula. — Amer. Mus. 
Novit. 1488: 1—46, 14 fig., 1 map. 

1951c. The identity of some Malaysian species of Ceriagrion, with descriptions of two 
new species (Odonata). — Treubia 21: 183—197, 4 fig. 

1952a. On the Papuasian representatives of the genus Macromia Rambur, with descriptions 
of five new species and some larval forms (Odon.). — Treubia 21: 437—468, 22 fig. 
1952b. Description of a new species of Lestes from South Australia. — Entom. Ber. 14: 
126—128, 2 fig. 

1953a. Revision of the Australasian species of Synthemis Selys (Odon., Corduliidae). 
With descriptions of four new species and a key to their identification. (O.S.R. Public. 
no. 38). — Idea (Bogor) 9: 70—88, 14 fig. 

1953b. The larval characters of the Protoneuridae (Odon.), with special reference to the 
genus Selysioneura Foerster, and with notes on other Indo-Australian genera. — Treubia 
21: 641684, 48 fig. 

1953c. The Odonata of the island Sumba, with a survey of the dragonfly fauna of the 
Lesser Sunda Islands. — Verh. Naturf. Ges. Basel 64: 118—228, 72 fig., 1 map. 
1953d. Revisional notes on the genera Diplacina Brauer and Huonia Foerster (Odon.). 
— Treubia 22: 153—216, 23 fig. 

1953e. Additions to the Odonate fauna of the Indo-Australian Archipelago. — Treubia 
22: 233269, 11 fig. 

1953f. New dragonflies (Odonata) from Borneo, with notes on their habits and larvae. — 
Treubia 22: 381—406, 7 fig. 

1954. Handlist of Malaysian Odonata. A catalogue of the dragonflies of the Malay Penin- 
sula, Sumatra, Java and Borneo, including the adjacent small islands. — Treubia 22 
Suppl. xiii + 202 pp., 1 map. 

1955a. Notes on the Australasian species of Nezrobasis Selys (Odonata, Agriidae). — 
Nova Guinea, new ser. 6: 155—166, 6 fig., pl. III, 1 map. 

1955b. Two new species of Lanthanusa Ris, from the high mountains of New Guinea 
(Odonata). — Zool. Meded. Leiden 33: 157—164, 9 fig. 

1955c. Further inquiries into the Old World species of Macromia Rambur (Odonata). — 
Zool. Meded. Leiden 33: 251—277, 28 fig. 

1955d. Notes on species of Nannophlebia Selys from the Moluccas and New Guinea 
(Odonata). — Zool. Meded. Leiden 33: 301—318, 14 fig. 

1955e. Synopsis of the dragonflies (Odonata) of Ceylon. — Zool. Meded. Leiden 34: 
67—87, 4 fig. 

1956a. Two new Platycnemididae (Odonata) from the Papuan Region. — New Guinea 
new ser. 7: 249—258, 7 fig. 

1956b. Revision of the genus Argiolestes Selys (Odonata) in New Guinea and the Moluc- 
cas, with notes on the larval forms of the family Megapodagrionidae. — Nova Guinea 
new ser. 7: 59—121, 76 fig. 

1957. Notes on some Argiine dragonflies (Odonata) with special reference to the genus 
Palaiargia Foerster, and with descriptions of new species and larval forms. — Nova 
Guinea new ser. 8: 41—80, 35 fig. & pl. II-V. 

1958. A review of the genus Idiocnemis Selys in the Papuan Region, with notes on some 


larval forms of the Platycnemididae (Odonata). —- Nova Guinea new ser. 9: 253—292, 
76 fig. 

1959a. New and little known Isostictine dragonflies from the Papuan Region (Odonata, 
Protoneuridae). — Nova Guinea new ser. 10: 279—302, 46 fig. 


1959b. On the New Guinea species of Ischnura Charpentier and Oreagrion Ris, with 
special reference to the larval forms and notes on the species of adjacent regions (Odonata, 
Coenagrionidae). — Nova Guinea new ser. 10: 213—240, 29 fig. 

1960a. On the identity of some little known Southeast Asiatic Odonata in European 
museums described by E. de Selys Longchamps, with descriptions of new species. — Mem. 
Soc. Ent. Ital. 38: 229—256, 13 fig. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 137 


———, 1960b. Three new species of Notoneura Tillyard from western New Guinea (Odonata, 
Protoneuridae). — Nova Guinea, Zool. 7: 117—126, 12 fig. 

——, 1960c. Considerations on the genus Lestes Leach, with notes on the classification and 
descriptions of new Indo-Australian species and larval forms (Odonata, Lestidae). — 
Nova Guinea, Zool. 8: 127—171, 51 fig., pl. IV. 

——, 1961a. New and interesting Odonata from the Philippines. Philippine Zoological Ex- 
pedition 1946-47. — Fieldiana (Chicago) Zool. 42: 119—149, 52 fig. 

———, 1961b. Notes on the affinity and nomenclature of some Old World Corduliidae (Odonata). 
— Proc. Kon. Ned. Akad. Wet. (Amsterdam) Zool. ser. C, 64 no. 3 (1960): 410—423, 
4 fig., 1 map. 

———, 1962. Odonata in Insects of Micronesia (Honolulu) 5: 1—95, 30 fig., 1 map. 

—, 1963a. Contributions to the Odonate fauna of the Solomon Islands, with notes on 
zygopterous larvae. — Nova Guinea, Zool. 21: 523—542, 24 fig., pl. 26. 

——, 1963b. The type of Libellula hova Rambur, 1842, with notes on the other species of 
Zygonyx Selys from Madagascar (Odonata). — Verh. Naturf. Ges. Basel 74: 53—61, 
8 fig. 

——, 1963c. New species and records of Libellulidae from the Papuan Region (Odonata). — 
Nova Guinea, Zool. 25: 751—780, 39 fig. 

—, 1964a. Synonymic notes on East Asiatic Gomphidae with descriptions of two new species 
(Odonata). — Zool. Meded. Leiden 39, feestbundel H. Boschma, 17 Jan. 1964: 89—110, 
22 fig. 

—, 1964b. Some Gomphidae and their larvae, chiefly from the Malay Peninsula (Odonata). 

— Zool. Verhand. Leiden 69: 1—38, 47 fig. 

, 1965a. Some Odonata of the genus Drepanosticta Laidlaw, chiefly from the Malay Penin- 

sula (Platystictidae). — Zool. Meded. Leiden 40: 171—186, 15 fig. 

, 1965b. The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopte- 

rygidae). — Tijdschr. Ent. 108: 325—364, 13 fig. 
—, 1965c. Notes on Odonata of Madagascar, with special reference to the Zygoptera and 
with comparative notes on other faunal regions. — Verh. Naturf. Ges. Basel 76: 
229—256, 12 fig., 2 tab. 
——, 1966a. A new species of Burmagomphus Williamson, from Thailand (Odonata). — 
Entom. Meddel. Copenhagen 34: 193—197, 5 fig. . 
——., 1966b. Some Odonata of Rapa Island, with descriptions of three Polynesian species of 
Ischnura Charpentier. — Tijdschr. Ent. 109: 89—102, 3 fig. 
——., 1966c. A survey of the dragonfly fauna of Morocco (Odonata). — Bull. Inst. r. Sci. nat. 
Belg. 42: 1—63, 32 fig., 1 map. 

, 1968a. A review of the genus Oligoaeschna Selys in Southeast Asia. — Tijdschr. Ent. 111: 
137—186, 11 fig., pl. 12—13. 

, 1968b. The Odonata of Rennell Island, Solomon Archipelago. — The Nat. Hist. of Rennell 
Island, BSI. 5: 67—74, 1 fig. 

————, 1969a. Two new Odonata from Southeast Asia, with comments on previously described 
species. — Dtsch. Ent. Z. N.F. 16: 205—215, 14 fig. 

——, 1969b. Over de oecologie van libellelarven. — De Levende Natuur 72: 103—109, 3 fig. 

, 1969c. Odonata Anisoptera. In Hydrobiological Survey of the Lake Bangweulu Luapula 

River basin. 14. Brussels, 64 pp., 10 fig., 1 map. 

——, 1971a. Odonata from Ceylon. Report No. 4 in Rep. Lund Univ. Ceylon Expedition 1962, 

1. — Entomologica Scandinavica (Lund) Suppl. 1: 189—208, 10 fig. (Publ. Munksgaard, 

Copenhagen). 

1971b. Studies in Oriental Corduliidae (Odonata) I. — Tijdschr. Ent. 114: 1—63, 50 fig. 

Meciachlan: R. 1898. On two species of Calopterygidae from the island of Lombock, with varietal 
notes. — Ent. Mon. Mag. (2) 9: 272—274. 

Martin, R. 1904. Liste des Névroptères de l’Indo-Chine. — I» Mission Pavie, Zool.: 204—221. 

, 1907a. Cat. Coll. Selys Longchamps. Cordulines 17: 1—94, fig. 1—99, col. pl. III. 

———, 1907b. Deux espèces nouvelles du genre Heliaeschna (Aeschnidae). — Notes Leyden Mus. 
28: 221—223. 

——., 1909a. Cat. Coll. Selys Longchamps. Aeschnines 19: 85—156, fig. 78—156, col. pl. 
III-IV. 


138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


, 1909b. Cat. Coll. Selys Longchamps. Aeschnines 20: 157—223, fig. 157—219, col. pl. 
V—VI. 
Navas, L. 1932. Névroptères et insectes voisins. Chine et Pays environnants. III. — Notes d’Ent. 
Chinoise (Chang-Hai) 8: 1—11, fig. 18—21. 
Needham, J. G. 1940. Studies on neotropical Gomphine dragonflies (Odonata). — Trans. Amer. 
Ent. Soc. 65: 363—394, pl. 20—22. 
, 1944. Further studies on neotropical Gomphine dragonflies. — Trans. Amer. Ent. 
Soc. 69: 171—224, pl. 14—16. 
Rambur, P. 1842. Histoire naturelle des insectes. Névroptères. Paris. xvii + 534 pp., 12 pl. 
Ris, F. 1898. Neue Libellen vom Bismarck Archipel. — Entom. Nachr. 24: 321—327. 
, 1900. Libellen vom Bismarck-Archipel gesammelt durch Prof. Friedr. Dahl. — Arch. f. 
Naturgesch. 66: 175—204, pl. 9—10. 
, 1910. Cat. Coll. Selys Longchamps. Libellulinen 11: 245—384, fig. 153—232, col. pl. II. 
, 1912a. Über Odonaten von Java und Krakatau gesammelt von Edward Jacobson. — 
Tijdschr. Ent. 55: 157—183, pl. 6—8. 
, 1912b. Cat. Coll. Selys Longchamps. Libellulinen 14: 701—836, fig. 408—491, col. pl. VI. 
, 1912c. Neue Libellen von Formosa, Südchina, Tonkin und den Philippinen. — Suppl. 
Ent. 1: 44-85, 19 fig., pl. 3—5. 
, 1913. Die Odonata von Dr. H. A. Lorentz’ Expedition nach südwest Neu-Guinea 1909 
und einige Odonata von Waigeu. — Nova Guinea 9, Zool. 3: 471—511, 28 fig. 
, 1915a. Fauna simalurensis. Odonata. — Tijdschr. Ent. 58: 5—21, 5 fig. 
, 1915b. Neuer Beitrag zur Kenntnis der Odonaten-Fauna der Neu-Guinea Region. Nova 
Guinea 13, Zool. 2: 81—131, 36 fig. 
, 1919. Cat. Coll. Selys Longchamps. Libelluninen 162: 1043—1278, fig. 605—692. 
, 1927. Odonaten von Sumatra, gesammelt von Edward Jacobson. — Zool. Meded. Leiden 
10: 1—49, 24 fig. 
, 1929. Fauna Buruana. Odonata gesammelt von L. J. Toxopeus auf Buru, 1921—1922, 
nebst einigen Odonaten von Amboina. (2. Teil, Zygoptera). — Treubia 7 Suppl.: 139—147, 
6 fig. 
St. Quentin, D. 1967. Die Gattung Gomphoides Selys (Ordnung Odonata) und ihre Verwandten 
in der neotropischen Region. — Beitr. Neotrop. Fauna (Stuttgart), 5: 132—152, 8 fig. 
Schmidt, Erich. 1934. Odonata der Deutschen Limnologischen Sunda-Expedition. — Archiv f. 
Hydrob., Suppl. 13 Tropische Binnengewässer 5: 316—397, 93 fig., pl. 14—17. 
, 1951. The Odonata of Madagascar, Zygoptera (transl. by F. C. Fraser). — Mém. Inst. 
Sci. Madagascar (A) 6: 115—283, 89 fig. 
, 1952. Odonata nebst Bemerkungen tiber die Anomisma und Chalcopteryx des Amazonas- 
Gebietes. Iv Beitr. Fauna Perus (Jena) 3: 207—256, 13 fig., 4 pl. 
Selys Longchamps, E. de. 1853. Synopsis des Caloptérygines. — Bull. Acad. Belg. 20 Annexe: 1—75. 
, 1854. Synopsis des Gomphines. — Bull. Acad. Belg. 21 (2): 23—112 (3—93 sep.). 
——, 1862. Synopsis des Agrionines, 3™° Légion: Podagrion. — Bull. Acad. Belg. (2) 14: 
5—55 (3—42 sep.). 
—, 1863. Synopsis des Agrionines, de Légion: Platycnemis. — Bull. Acad. Belg. (2) 16: 
147—176 (3—32 sep.). 
—, 1865. Synopsis des Agrionines, 5"° Légion: le grand genre Argia. — Bull. Acad. Belg. 
(2) 20: 375—417 (3—45 sep.). 
———, 1869a. Secondes Additions au Synopsis des Caloptérygines. — Bull. Acad. Belg. (2) 27: 
645—680 (1—36 sep.). 
——, 1869b. Secondes Additions au Synopsis des Gomphines. — Bull. Acad. Belg. (2) 28: 
168—208 (5—45 sep.). 
——, 1871. Synopsis des Cordulines. — Bull. Acad. Belg. (2) 31: 238—565 (5—128). 
——, 1876. Synopsis des Agrionines (suite) 5™° Légion: le grand genre Agrion. — Bull. Acad. 
Belg. (2) 41—42: 247— 322, 496—501, 539, 1233—1309 (3—199 sep.). 
——, 1877. Synopsis des Agrionines (suite et fin) 5™° Légion: Agrion. — Bull. Acad. Belg. 
(2) 43: 97—159 (3—65 sep.). 
———, 1878a. Quatriémes Additions au Synopsis des Gomphines. — Bull. Acad. Belg. (2) 46: 
408—471, 658—698 (3—106 sep.). 
——, 1878b. Odonates de la région de la Nouvelle Guinée. — Mitteil. Kônigl. Zool. Mus. 
Dresden 3: 289—323. 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 139 


——, 1879. Quatriémes Additions au Synopsis des Caloptérygines. — Bull. Acad. Belg. (2) 47: 
349—409 (3—63 sep.). 

——, 1885. Programme d'une Revision des Agrionines. — C.R. Soc. Ent. Belg. (3) 66, Ann. 
Soc. Ent. Belg. 29: CXLI—CXLVIII. 

—, 1886a. Odonates nouveaux de Pékin. — C.R. Soc. Ent. Belg. (3) 78, Ann. Soc. Ent. 
Belg, 30: CLXXVIII—CLX XXV. 

———, 1886b. Revision du Synopsis des Agrionines. — Mém. cour. Acad. Belg. 38 (4): iv + 
233 pp. 

——, 1887. Odonates de l’Asie Mineure, etc. — Ann. Soc. ent. Belg. 31: 1—49. 

——, 1889. Odonates de Sumatra, comprenant les espèces recueillies à Pulo Nias par M. le Dr. 
E. Modigliani. — Ann. Mus. civ. Genova 7 (27): 444484. 

——, 1891a. Viaggio di Leonardo Fea in Birmania e regioni vicine. 32. Odonates. — Ann. Mus. 
civ. Genova 10 (30): 433—518. 

——, 1891b. Additions aux Odonates des Philippines. — Anal. Soc. Espan. Hist. Nat. 20: 
209—218. 

Selys Longchamps, E. de & H. A. Hagen, 1854. Monographie des Caloptérygines. — Mém. Soc. 
Sci. Liege 9, XI+-291 pp., 14 pl. 

, 1858. Monographie des Gomphines. — Mem. Soc. Sci. Liege 11, VIII+p. 257—720, 
23 pl. 

Tillyard, R. J. 1913. On some new and rare Australian Agrionidae (Odonata). — Proc. Linn. 
Soc. N.S. Wales (1912) 37: 403—479, pl. 44—49. 

, 1926. On a collection of Papuan dragonflies (Odonata) made by the late Mr. Allan 
R. McCulloch in 1922-3, with descriptions of new species. — Rec. Austral. Mus. 15: 
157—166, 6 fig. 

Watson, J. A. L. 1967. An analysis of Trapezostigma eurybia (Selys, 1878) and related Indo- 

Australian species (Odonata, Libellulidae). — Nova Guinea, Zool. 36: 377—400, 49 fig. 

Weele, H. W. van der. 1909. Neuropteroidea. — Nova Guinea 9, Zool. 1: 19—25, fig. 1—2. 

Williamson, E. B. 1915. Notes on neotropical dragonflies, or Odonata. — Proc. U.S. Nat. Mus. 
48: 601—638, pl. 38—44. 

, 1916. A new Cyanogomphus (Odonata). — Ent. News 27: 167—172, pl. 8—9. 


ADDENDA TO PART IV 


argentea Ris (Argyrothemis), 1911 (¢). First described 9. French Guyana, Massikiri, 
Oyapock, 16.X1.1969, Mission Balachowsky et al. (MP). See Geijskes, 1971: 668— 
669, fig. 2 A-B. — Libellulidae. K 

carminita Calvert ( Telebasis), 1909 (3). First described ©. French Guyana, Savanne 
à Graminées, 30 km de Kourou vers Trakombo, 21.X.1969 (g 9), Mission Bala- 
chowsky et al. (MP). See Geijskes, 1971: 660—663, fig. 1 F. — Coenagrionidae. 

longitudinalis Ris (Anatya), 1919 (3). First described @. French Guyana, Saut- 
Maripa, 25—27.XI.1969, at light (& ©), Mission Balachowsky et al. (MP). See 
Geijskes, 1971: 672—673, fig. 3 A-B. — Libellulidae. 

Now Erythrodiplax longitudinalis (Ris). 


REFERENCES 


Calvert, P. P., 1909. Contributions to a knowledge of the Odonata of the neotropical region, ex- 
clusive of Mexico and Central America. — Ann. Carnegie Mus. 6: 73—280, 9 pl. 

Geijskes, D. C., 1971. List of Odonata known from French Guyana, mainly based on a collection 
brought together by the Mission of the “Museum National d'Histoire Naturelle”, Paris. 
— Ann. Soc. ent. Fr. (N.S.) 7: 655—677, 3 fig. 

Ris, F., 1911. Cat. Coll. Selys Longchamps. Libellulinen 12: 385—528, fig. 233—317. 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 PLATE 1 


TEE 


thalia 


venilia 


arboricola 


rhaphia 


virgula 


leptalea 


Genera Selysioneura and Tanymecosticta 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 PLATE 2 


viduata & 


U ee 


rhodosoma & 


acuticauda 4 9 


sagittiferum & 2 


Genera Archboldargia, Ischnura and Austroallagma 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 PLATE 3 


armeniacum 6 2 


Genus Oreagrion 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 


PLATE 4 


drescheri 


thelyphonus 


inscriptus 


fruhstorferi 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


Heliogomphus, Onychogomphus and Acrogomphus 


, 


Genera Burmagomphus 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 PLATE 5 


acuta D 
magnificus 


dictatrix 


Genera Chlorogomphus, Paragomphus, Plattycantha and Oreaeschna 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 


PLATE 6 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 


1 


| 


un À 


N 


[; 


an 


( 
gomphoides 


da 


mn Mm 
Cnn 


silvarum 


teuchestes 


arborophila 


alexia 


arethusa 


buruensis 


Bironides, Microtrigonia, Orthetrum and Huonia 


3 


Genera Nannophlebia 


Specimens of Odonata 


Catalogue of type 


M. A. LIEFTINCK : 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 2, 1971 PLATE 7 


Photos Ch. Hoorn Jr. 
Above: Nannophyopsis chalcosoma Lieftinck, &: below: Neopetalia punctata (Selys), 
first described Q 


M. A. LIEFTINCK : Catalogue of type-specimens of Odonata 


No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.FI. 45.—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.F. 70.—. 


No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Ditsh & Uvarov 
and Xenocatantops Ditsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. 
D.FI. 25.—. Ss 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. 
DFI. 75.—. 


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DESCRIPTIONS D’ATTACIDAE (=SATURNIIDAE) ') NOUVEAUX 
D’AMERIQUE CENTRALE ET DU SUD (LEPIDOPTERA) 


par 


CLAUDE LEMAIRE 


Paris, France 


SYNOPSIS 


L'auteur décrit 17 espèces et 7 sous-espèces nouvelles d’ Attacidae (= Saturniidae) d'Amérique 
centrale et du Sud. Ces taxa se répartissent dans les quatre sous-familles représentées sur le conti- 
nent américain: Attacinae (5 espèces, 2 sous-espéces), Hemileucinae (5 espèces, 1 sous-espéce), 
Arsenurinae (1 sous-espéce), Dryocampinae (7 espéces, 3 sous-espéces). La note contient également 
6 synonymies nouvelles et la désignation de 2 lectotypes. 

Le matériel typique appartient au British Museum (Nat. Hist.) et aux collections de M. M. 
LicHy, DARGE et LEMAIRE; les holotypes de ces trois derniéres collections seront déposés au 
Muséum national d'Histoire naturelle, Paris. 


ATTACINAE 


Copaxa (Copaxa) escalantei spec. nov. 
Fig. 1, nr. 1—2; Pl. 1 Fig. 1 


Holotype: 1 g', Mexique, Puebla, San Juan Apulco, VI.1960 (Dr. T. Escalante) 
(genitalia &, prép. C. Lemaire n° 2298) (coll. C. Lemaire); allotype: 1 9, X.1963; 
paratypes: 4 &, V1.1960, X.1963, tous mêmes localité, récolteur et collection. 

d. — Envergure: 87—112 mm; longueur des ailes antérieures: 50—61 mm. 

Antennes jaune paille: 30 articles. Téte, prothorax et tarses gris brun violacé, le reste 
du corps jaune. Couleur fondamentale des quatre ailes jaune vif sur la face dorsale, 
jaune terne mêlé de grisâtre ou de brunâtre sur la face ventrale. Ornementation habi- 
tuelle des espèces du genre, caractérisée ici par la netteté avec laquelle se détachent, sur 
le dessus des ailes antérieures, en noirâtre, brun noir ou violacé, les nervures, la rayure 
externe, très large et s'interrompant sous l’apex au niveau de R 5, et la bande submargi- 
nale. Le nombre de taches hyalines varie de trois à cinq sur les ailes antérieures, de deux à 
quatre sur les ailes postérieures. Un semis d’écailles noirâtres sur les bords distal et 
subcostal de la cellule. 

Q . — Envergure: 127—132 mm; longueur des ailes antérieures: 70 mm. 

Diffère du mâle, outre la forme plus allongée des ailes antérieures, par la coloration 
fondamentale beige brun du corps et des quatre ailes, avec, sur ces dernières, des zones 
brun orangé et violacées. Six ou sept taches hyalines sur les ailes antérieures, six sur les 
ailes postérieures. 


1) Par application de l'article 23 (a) (ii) du Code international de Nomenclature zoologique et de 
l'opinion 450, publiée le 8 mars 1957, de la Commission internationale de Nomenclature zoolo- 
gique qui, en vertu de ses pleins pouvoirs a placé Attacidae Burmeister, 1878, sur la liste of- 
ficielle des noms du groupe-famille en Zoologie, sous le n° 142. Décision regrettable, rendant 
inutilisable Saturniidae Boisduval, 1837, en dépit de sa priorité et allant à l'encontre de l'usage 
consacré par la majorité des auteurs. A noter qu’Attacidae avait été utilisé par Duponchel dès 
1844! 


142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


Espéce trés proche de C. (C.) expandens Walker (= trimacula W. Rothschild, syn. 
nov.?)) avec qui elle a été confondue, notamment par Draudt (1929: 724, pl. 105, a g') 
et par C. C. Hoffmann (1942: 236, n° 1415). En différe par le nombre supérieur des 
taches hyalines ornant les exemplaires des deux sexes (au plus trois sur les ailes antérieu- 
res et une seule sur les ailes postérieures chez C. (C.) expandens), par la coloration 
fondamentale jaune trés constante des exemplaires males (varie du jaune au brun foncé 
et au brun rougeâtre ou orangé chez C. (C.) expandens) dont l'apex est en général plus 
aigu et saillant. Antennes sensiblement plus courtes (30 articles contre 34 à 36). L'ar- 
mure génitale mâle (Fig. 1 nr. 3—4) est nettement distincte de celle de C. (C.) expandens 
par la forme de l’aedeagus et surtout par la structure des larges processus sclérifiés ou 
harpes qui émanent des valves sur la face interne de la costa. 

Je dédie avec plaisir cette espèce inédite à l’éminent entomologiste mexicain, M. le 
Docteur T. Escalante, à qui je suis redevable du matériel typique. 


Copaxa (Copaxa) andensis spec. nov. 
Fig. 2 nr. 1—2; PI. 1 Fig. 2 


Holotype: 1 4, Colombie, Valle Cauca, Los Andes, 1700 m, 25.X11.1966 (L. et L. 
Denhez) (genitalia g', prép. C. Lemaire n° 1854) (coll. C. Lemaire); paratypes: 2 d', 
même localité, 21.11.1966, 2 &, Valle Cauca, route de Cali à Buenaventura, km 18, 1800 
m, 28.X, 9.XII.1970, tous mêmes récolteurs et collection. 

d'. — Envergure: 88—104 mm; longueur des ailes antérieures: 51—58 mm. 

Antennes jaune paille: 31 articles. Tête et prothorax noirätres, avec fréquemment une 
touffe de poils jaune entre les scapes. Tarses roses, le reste du corps brun foncé, plus 
ou moins orangé. Couleur fondamentale de la face dorsale des quatre ailes beige brun 
foncé, avec généralement des zones plus orangées, de la face ventrale gris beige violacé. 
Les rayures brun noir et la bande submarginale sont assez faiblement marquées mais 
les fascies antémédianes sont généralement nettes. Les taches hyalines sont liserées, sur 
la face dorsale, de noir, puis de jaune; il en existe de trois à cinq sur les ailes antérieures 
et, sur les ailes postérieures, un seule, avec parfois un petit point accessoire. 

Espèce voisine de C. (C.) multifenestrata (Herrich-Schäffer) avec laquelle elle coexiste 
dans la localité typique. En diffère par la forme de l’apex qui est arrondi, par l’inter- 
ruption de la rayure externe des ailes antérieures entre M 1 et M 5, atteignant rarement 
celle-ci et par la réduction du nombre des fenêtres hyalines (il en existe généralement 
plusieurs sur les ailes postérieures de C. (C.) multifenestrata). Les femelles que nous 
rapportons avec un léger doute à cette espèce présentent le même ensemble de caractères. 
L’armure génitale mâle (Fig. 2) est distincte de celle de C. (C.) multifenestrata par sa 
structure bien plus robuste, avec l’aedeagus nettement plus long et surtout par la forme 
des harpes qui, au lieu d’être réduites à de petites formations subconiques (Fig. 2, n° 3), 
se présentent ici comme de longs appendices très sclérifiés, légèrement épineux. Cette 
espèce nouvelle est également connue de l’Equateur, rio Negro, du Pérou, Carabaya, de 
la Bolivie, Cochabamba, où elle vole à des altitudes variant entre 1500 et 2000 mètres. 


2) C. (C.) trimacula a été décrit par W. Rothschild (1895:40) d'après deux exemplaires d’Ameri- 
que centrale, sans autre précision de localité, qui sont actuellement conservés par le British 
Museum (Nat. Hist.). Nous désignons comme lectotype celui de ces deux spécimens dont 
l’armure génitale mâle, examinée par nous, porte le n° de préparation: Saturniidae, n° 120, 
on slide. L'holotype & de C. (C.) expandens est originaire du Venezuela. 


C. LEMAIRE: Attacidae nouveaux 143 


Copaxa (Copaxa) denhezi spec. nov. 
Fig 83 Pi) Bigs 3 


Holotype: 1 g', Colombie, Valle Cauca, Anchicaya, 1000/1400 m, 6.VIII.1968 
(genitalia g', prép. C. Lemaire n° 1840) (L. et L. Denhez) (coll. C. Lemaire). 

d. — Envergure 97 mm; longueur des ailes antérieures: 55 mm. 

Antennes jaune orangé: 30 articles. Front brun foncé, prothorax gris violacé, le reste 
du corps beige brun. Ailes brunes sur les deux faces, avec de légéres éclaircies sur le des- 
sus des ailes antérieures et des zones gris violacé sur la face ventrale. Ornementation 
habituelle des espèces du genre. La rayure externe des ailes antérieures s'interrompt à 
l'intersection de R 5 et du bord externe, les bandes antémédianes sont nettes, les bandes 
submarginales, sur la face dorsale, étant au contraire très estompées. Les taches hyalines 
sont finement liserées de brun noir, puis de jaune, sur les deux faces; elles sont au 
nombre de six sur les ailes antérieures, de quatre sur les ailes postérieures. 

Difficilement séparable de C. (C.) multifenestrata; ne diffère des grands exemplaires 
sombres de cette espèce que par la forme un peu plus falquée des ailes antérieures, avec 
apex plus aigu et par la position de la rayure externe de ces ailes qui, sur la face 
dorsale, atteint, au niveau de R 5, le bord externe dont elle reste toujours assez éloignée 
chez C. (C.) multifenestrata. L'armure génitale mâle (Fig. 2 nr. 3), bien plus robuste que 
chez ce dernier, en diffère par la structure de l’aedeagus et surtout des harpes, beaucoup 
plus développées et rappelant davantage celles de C. (C.) escalantei (Fig. 1 nr. 1—2). Les 
lobes de l’anellus sont beaucoup plus courts que chez celui-ci et plus fortement asymé- 
triques. 

Espèce dédiée à mon excellent correspondant M. L. Denhez à qui je suis redevable de 


holotype. 


Copaxa (Copaxa) canella miranda subsp. nov. 
PINS ord PIN EI on 


Holotype: 1 g', Bolivie, Cochabamba, Carrasco, Siberia, 1650 m, 1.1964, (coll. C. 
Lemaire); allotype: 1 9, 26.XII.1962; paratypes: 19 g, XII.1962, I, X, X11.1963, 
1.1964, 7 9, X, X11.1963, 1.1964, tous mêmes localité et collection. 

d'. — Envergure: 95—126 mm; longueur des ailes antérieures: 50—65 mm. 

9. — Envergure: 112—124 mm; longueur des ailes antérieures: 62—68 mm. 

Cette sous-espèce diffère de la sous-espèce nominative par l’envergure moyenne, très 
supérieure, des exemplaires des deux sexes, celle de C. (C.) c. canella n’excédant guère 
105 mm, ainsi que par son ornementation beaucoup plus marquée et contrastante, sur- 
tout celle du mâle, notamment sur les ailes antérieures où les nervures sont très forte- 
ment soulignées de noirâtre et la rayure médiane onduleuse toujours très nette. La 
coloration fondamentale des exemplaires mâles, brune chez C. (C.) c. canella, est ici 
beige terne, avec des reflets orangés et un très abondant semis d’écailles noirâtres, ou 
orange vif; elle rappelle alors celle de C. (C.) joinvillea Schaus (mais ou l’apex des 
ailes antérieures est moins saillant) et surtout de C. (C.) simson Maassen à qui les 
plus grands exemplaires ressemblent étonnamment (mais dont l’armure génitale est 
différente). La coloration dominante de la femelle est jaune vif, avec des semis d'écailles 


noirâtres et les bandes submarginales rouge vineux, comme chez C. (C.) c. flavina 
Draudt. 


144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


Nous connaissons cette sous-espèce nouvelle, outre la localité typique, de diverses 
stations de la Bolivie, Cochabamba (El Limbo, 2000/2200 m, Alto Palmar, 1000 m) 
et du Pérou, Junin (La Merced) et Pasco (Oxapampa). 


Copaxa (Sagana) herbuloti spec. nov. 
Fig. 4; Pl. 3 Fig. 6 


Holotype: 1 g', Pérou, Piura, Huancabamba genitalia &, (prép. C. Lemaire n° 1820) 
(coll. C. Lemaire). 

d'. — Envergure: 70 mm; longueur des ailes antérieures: 40 mm. 

Antennes jaune noirâtre: 25 articles. Front brun rougeâtre, dessus du thorax semé 
de longues écailles jaunes, abdomen brun orangé sur la face dorsale, rougeâtre sur la 
face ventrale, tarses roses. Ailes antérieures très falquées, apex arrondi, bord externe 
fortement concave, tornus droit. Couleur fondamentale des ailes antérieures 
brune avec un abondant semis d'écailles jaunes; aire externe brun noiràtre. Tache pré- 
apicale noire, liserée distalement de blanc et suivie en dehors d'une tache rose. Rayures 
brunes, extrêmement vagues, l'externe nettement préapicale et tangente à la grande 
fenêtre disco-cellulaire qui est liserée de brun noir. 

Zone subcostale et basale des ailes postérieures rose, aire médiane brune, semée 
d'écailles jaunes; fenétre disco-cellulaire liserée de noir, puis de jaune. Aire externe 
orangée. Petites taches submarginales brunes. Pas de rayures nettes. 

Face ventrale rougeätre sur l'aire baso-médiane et la région apicale, brun jaune sur 
l'aile externe qui est précédée, sur les quatre ailes, d'une bande violacée. 

Par la structure de l’armure génitale mâle (Fig. 4) et par la position de la deuxième 
médiane des ailes antérieures qui émane de l'angle antérieur de la cellule, cette espèce 
entre dans le sous-genre Sagana Walker auprès de C. (S.) sapatoza (Westwood). Dif- 
fère de ce dernier, notamment par la forme des ailes antérieures, plus falquées, la colora- 
tion des quatre ailes, l’effacement des rayures et la forme des fenêtres disco-cellulaires; en 
revanche les armures génitales mâles des deux espèces sont très voisines. 

Avec grand plaisir je dédie cette espèce nouvelle à mon excellent ami Claude Herbulot. 


Rothschildia paucidentata spec. nov. 
Fig. 5; Pl. 4 Fig. 7 


Holotype: 1 &, Venezuela, Aragua, Colonia Tovar, 1700 m, 29.V.1963 (W. Gatz) 
(coll. C. Lemaire); allotype: 1 Q, sans date de capture; paratypes: 3 g', V, VII.1963, 
tous méme récolteurs, localités, et collection. 

d'. — Envergure: 108—118 mm; longueur des ailes antérieures: 59—63 mm. 

Antennes jaune orangé: 30 articles. Front brun, larges colliers pro- et métathoraciques 
blancs, thorax et face dorsale de l’abdomen acajou plus ou moins rougeätre. Touffe 
anale blanche, dessous du corps brun avec une touffe de poils rougeâtres sur la région 
antérieure du thorax et deux lignes stigmatales et substigmatales brunes, liserées de 
blanc. Face antérieure des tarses brune, postérieure blanche. 

Ailes antérieures allongées et falquées. Couleur fondamentale acajou sur la face 
dorsale, brun foncé sur la face ventrale. Ornementation habituelle des espèces du genre. 
Rayures externes blanches, liserées distalement de brun rouge, celles du dessus des 


C. LEMAIRE: Attacidae nouveaux 145 


Armure genitale 4 des Attacidae. Fig. 1—1, Copaxa (Copaxa) escalantei sp.n., vue ventralement, 
pénis détaché; 2, id., pénis vu latéralement; 3, C. (C.) expandens Walker, pénis vu latéralement; 
4, id., harpes. Fig. 2. — 1, Copaxa (Copaxa) andensis sp.n., vue ventralement, pénis détaché; 2, id., 
pénis vu latéralement; 3, C. (C.) multifenestrata (Herrich-Schäffer), harpes. Fig. 3. — 1, Copaxa 
(Copaxa) denhezi sp.n., vee ventralement, pénis détaché; 2, id., pénis vu latéralement. Fig. 4. — 
1, Copaxa (Sagana) herbuloti sp.n., vue ventralement, pénis détaché; 2, id., pénis vu latéralement. 
Fig. 5. — 1, Rothschildia paucidentata sp.n., vue ventralement, pénis détaché; 2, id., pénis vu 
latéralement. L’échelle figurée représente 1 mm 


146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


ailes antérieures droites, dirigées obliquement vers la base entre la costa et le bord 
interne, avec une seule dent bien indiquée entre Cu 1b et Cu 1a. Bandes submarginales 
violacées et semées d'écailles noires sur les deux faces, vagues dorsalement, nettes 
ventralement. Taches marginales des ailes postérieures rouges entre la costa et Cu 1b, 
noires ensuite. Trait préapical blanc des ailes antérieures droit entre R5 et M1, et se 
prolongeant vers la base sur M 1. Fenêtres disco-cellulaires triangulaires, relativement 
petites, présentant sur les ailes antérieures un angle rentrant très net. 

Q. — Envergure: 118 mm; longueur des ailes antérieures: 66 mm. 

Ailes beaucoup moins falquées que le mâle, même coloration. Fenêtres disco-cellu- 
laires plus grandes, dépourvues ou presque d’angles rentrants sur les ailes antérieures, à 
bord proximal droit ou convexe sur les ailes postérieures. Rayures externes des ailes 
antérieures pas ou peu obliques, un peu plus sinueuses que chez les exemplaires mâles. 

Espèce voisine de R. lebean (Guérin-Méneville) dont elle diffère par des caractères 
externes constants: coloration plus sombre et, semble-t-il, très peu variable (à l'inverse 
de chez R. /. lebeau), rayures externes des ailes antérieures beaucoup plus droites et 
différemment orientées, surtout celles des exemplaires mâles, avec un trait proximal 
blanc nettement plus étroit et une absence presque totale de dents. Les fenêtres disco- 
cellulaires sont beaucoup plus petites, celles du mâle n’atteignant pas la rayure externe 
des ailes postérieures, avec des liserés blancs, puis noirs, bien plus fins, parfois imper- 
ceptibles. Forme de ces fenêtres également très différente puisque chez R. lebeau, leur 
bord interne est, sur les ailes antérieures, droit ou convexe et dépourvu d’angle rentrant 
et, sur les ailes postérieures, convexe.*) 

Les armures génitales mâles (Fig. 5) des deux espèces sont identiques mais le fait ne 
présente pas de signification particulière, étant assez fréquent dans le genre Rothschildia: 
ainsi l’armure de R. jorulla (Westwood) ne diffère-t-elle pas non plus de celle de 
R. lebean. 


Rothschildia lebeau yucatana subsp. nov. 
PI. 4 Fig. 8 


Holotype: 1 &, Mexique, Quintana Roo, X-can, VI.1969 (E. C. Welling) (coll. C. 
Lemaire); allotype: 1 9, même localité, 21.V.1969; paratypes: 13 g', 2 ®, même 
localité, VI.1964, V, VI, VIII.1967, V, IX.1968, V, VI.1969; 6 8, Yucatan, Chaksikin, 
VI.1964, 1 G', Yucatan, Umän, VI. 1966, 2 6, Yucatan, Chichen Itza, VI.1956, tous 
même récolteur et collection. 

d'. — Envergure: 83—103 mm; longueur des ailes antérieures: 48—60 mm. 

9. — Envergure: 95— 98 mm; longueur des ailes antérieures: 57—61 mm. 

Cette sous-espèce qui paraît propre à la péninsule du Yucatan, présente les mêmes 
variations de coloration que la sous-espèce nominative, mais, par son envergure moyenne 
très inférieure à celle-ci (dont certains exemplaires dépassent 120 mm), ainsi que par 
la présence d’un liseré marginal blanc, constant sur la face dorsale des ailes antérieures, 
très fréquent sur les ailes postérieures, elle rappelle surtout R. lebeau forbesi Benjamin. 
La rayure externe des ailes antérieures est plus large que chez ce dernier et en général 
plus fortement dentée, moins défléchie vers la base sous la costa et plus régulièrement 
convexe. Les liserés blancs des fenêtres disco-cellulaires sont généralement plus nets. 
Mais ce qui caractérise le mieux cette sous-espèce inédite est la forme très peu falquée 


3) Pour comparaison, voir R. lebeau yucatana subsp. nov., Pl. 4 Fig. 8. 


C. LEMAIRE: Attacidae nouveaux 147 


des ailes antérieures des exemplaires mâles, avec l’apex à peine saillant et le bord 
externe presque droit, comme chez la femelle et au contraire des autres sous-espéces où 
le mâle a toujours les ailes antérieures beaucoup plus falquées. 


HEMILEUCINAE 


Dirphia lichyi spec. nov. 
Fig. 6; Pl. 5 Fig. 9—10 


Holotype: 1 g', Venezuela, Aragua, Rio Choroni, Puerto Colombia, ex larva, 7.1. 
1945 (R. Lichy) (sa coll.); allotype: 1 9, 13.11.1945; paratypes: 22 4, 13 9, tous 
même localité, ex larva, 27.1./7.111.1945, 1 g', Carabobo, vallée du Rio Borburata, 250 m, 
1.X11.1956/31.111.1957, 1 &, même localité, 670 m, 10.V.1950, tous mêmes récolteur et 
collection. 

d. — Envergure: 76—91 mm; longueur des ailes antérieures: 43—48 mm. 

Antennes jaune orangé: 43 articles. Tête, dessus du thorax et tarses bruns, abdomen 
jaune orangé, annelé dorsalement de noir. Aires interne et externe des ailes antérieures gris 
clair, aire médiane brune, rayure externe gris clair, liserée distalement de brun, bande 
submarginale brune, bordée distalement de gris clair, très striée et ne se détachant nette- 
ment qu’entre Cu 1b et M3, d'une part, M1 et la costa, d'autre part. Le signe en Y 
couché qui représente la tache disco-cellulaire chez les Dirphia du groupe, D. tarquinia 
(Cramer) est ici très fin et sa branche proximale n’atteint généralement pas l’aire interne; 
sa branche subcostale, plus courte, renferme un trés léger trait brun jaune. 

Ailes postérieures orange terne, plus ou moins ombrées de noirätre sur l’aire externe, 
ornées d’une très vague rayure externe noirâtre, convexe et d'une tache disco-cellulaire 
noire, en forme de virgule. Face ventrale uniformément beige orangé, sans aucun or- 
nement. 

Q. — Envergure: 105— 120 mm; longueur des ailes antérieures: 58—65 mm. 

Différe essentiellement du mâle par sa taille très supérieure, son aspect plus massif et 
par la coloration des ailes antérieures dont l’aire baso-médiane est entièrement brune et 
laire externe, plus étroite, brun clair, occupée dans toute sa partie proximale par la large 
bande submarginale gris rosé. Tache disco-cellullaire des ailes antérieures plus allongée, 
avec le petit trait brun jaune de la branche subcostale très vague ou absent; celle des 
ailes postérieures plus large. 

Espèce très proche de D. tarquinia et ayant la même envergure, très supérieure à 
celle des autres Dirphia du même groupe. Elle en différe, le mâle par le développement 
de l’aire médiane, qui, sur la face dorsale des ailes antérieures, au lieu d'être réduite à 
un simple losange, parfois très petit, occupe tout le milieu de l’aile et est aussi large 
sur le bord interne que sous la costa, la femelle par la disparition totale de la zone claire 
correspondant 4 l’aire interne des mêmes ailes. Les deux espèces coexistent dans une 
des localités typiques (Rio Borburata). Leurs armures génitales (Fig. 6) sont similaires 
mais le fait est courant entre espéces voisines du genre Dirphia. 

La femelle de D. /ichyi a été décrite par Bouvier (1924: 384) comme «variation extrême 
de tarquinia», en même temps qu'un mâle appartenant, lui, à cette dernière espèce. 
Schüssler (1934: 412), jugeant à tort ces deux exemplaires, sans les avoir vus, conspécifi- 
ques, les a nommés Hyperdirphia tarquinia forme bouvieri, nom de rang infra-subspécifi- 
que, à invalider par application de l’article 45 (d) (iii) du Code international de Nomen- 
clature zoologique. Afin d’éviter toute confusion, nous désignons comme lectotype de 


148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


cette «forme» le syntype male du Venezuela, Bané (M. Grisol), conservé par le Muséum 
de Paris. D. tarquinia forme bouvieri devient ainsi un synonyme pur et simple de D. 
tarquinia. 

Il était juste de dédier cette nouvelle espèce 4 M. René Lichy qui, en réalisant 
son élevage, a permis de la séparer avec certitude de D. tarquinia. 


Dirphia crassifurca spec. nov. 
Fig. 7; PI. 6 Fig. 11—12 


Holotype: 1 g', Venezuela, Mérida, Valle la Mucuy, 2300 m, 7.IX.1956 (R. Lichy) 
(sa coll.); allotype: 1 Q, 8.IX.1956; paratypes: 13 g', 1 9, 1/11.IX.1956, 17.VI.1966, 
tous mémes localité, récolteur et collection. 

d. — Envergure: 66—74 mm; longueur des ailes antérieures: 35—40 mm. 

Antennes jaune paille: 38 articles. Palpes labiaux brun noir, front, prothorax et dessus 
du corps orangés, dessus du thorax brun foncé, de l’abdomen orange, annelé de noir; 
tarses bruns. Aire baso-médiane des ailes antérieures gris clair, assez étroite, aire médiane 
brun foncé, aire externe brun clair mais presque entièrement recouverte par les stries, 
alternativement gris clair et brun foncé, de la bande submarginale. Rayures gris clair, 
externe dirigée très obliquement en dehors de la costa vers le bord interne, l’interne 
droite. Tache disco-cellulaire en Y couché a branches très larges, mais courtes, la distale 
fréquemment tronque, la subcostale renfermant un très léger trait jaune clair. 

Ailes postérieures orange s’obscurcissant vers le bord externe; rayure externe noirâtre, 
très estompée. Petit trait disco-cellulaire noir. Face ventrale orangée avec, sur chaque 
aile, une rayure externe blanche très estompée, celle des ailes postérieures droite. Un 
vague trait discocellulaire sur ces dernières. 

9. — Envergure: 88—91 mm; longueur des ailes antérieures: 47—49 mm. 

Aire interne des ailes antérieures gris violacé, aire médiane brune, aire externe brune 
également mais occupée, dans sa partie proximale, par la bande submarginale beige 
rosé, avec quelques stries brun noir. Ailes postérieures gris noiràtre avec un reflet orangé 
vers la base. Vague tache disco-cellulaire blanchâtre, teintée d’orangé. Face ventrale brun 
terne, rayures externes blanches, celles des ailes postérieures bien marquées; la tache 
disco-cellulaire de la face dorsale transparaît nettement. 

Armure génitale mâle (Fig. 7) beaucoup plus grêle que celle de D. tarquinia, mais 
sans différence de structure notable. 

Cette espèce dont l’envergure, chez les exemplaires des deux sexes, est bien moindre 
qui celle de D. tarquinia et D. lichyi, est essentiellement caractérisée par la largeur des 
branches de la tache disco-cellulaire des ailes antérieures, très supérieure à celle observée 
chez toutes les espèces du méme groupe. 


Cerodirphia marahuaca spec. nov. 
Kiens ME la nl 


Holotype: 1 8, Venezuela, Amazonas, base du Cerro Marahuaca, Cafio Tchari, 
7.V.1950 (R. Lichy) (genitalia &, prép. C. Lemaire n° 2324) (coll. R. Lichy); para- 
type: 1 g', mêmes localité, date, récolteur et collection. 

d'. — Envergure: 70—75 mm; longueur des ailes antérieures: 37—40 mm. 

Antennes jaune paille: 43 articles. Palpes labiaux, front, thorax et tarses orangés; 


C. LEMAIRE: Attacidae nouveaux 149 


abdomen noir, annelé de blanc sur les faces dorsale et latérales, beige orangé sur la face 
ventrale, touffe anale orangée. 

Couleur fondamentale de la face dorsale des quatre ailes rose, avec une légère nuance 
orangée, notamment sur la base des ailes postérieures. Rayures externes noiratres, celles 
des ailes postérieures très estompées ou réduites 4 un trait subcostal. Tache disco-cellu- 
laire des ailes antérieures, en forme d’Y couché, relativement petite, avec la branche 
inférieure très réduite, celle des ailes postérieures absente. Face ventrale beige rosé, 
très pâle avec, pour seule ornementation, les rayures externes noiratres. Costa des ailes 
postérieures blanches, bordées de noir, sur les deux faces, franges concolores. 

Cette espéce différe de C. speciosa (Cramer) par le rose beaucoup plus vif de la 
coloration de la face dorsale des quatre ailes, où les nervures ne se détachent pas en 
noirâtre, et par son envergure très supérieure, par laquelle elle se situe auprès de C. mota 
(Druce). Mais, chez ce dernier, les nervures sont soulignées de jaune d'or sur la face 
dorsale, les franges sont blanches et il n'existe aucune trace de rayure externe sur le 
dessus des ailes postérieures. 

L'armure génitale mâle (Fig. 8) est très différente de celles de C. speciosa et C. mota: 
elle est essentiellement caractérisée par la structure de l'uncus dont l'extrémité postérieure 
est simple (elle présente deux pointes chez C. speciosa) et la face dorsale surmontée 
d’un rostre n’existant chez aucune des espèces voisines. Le huitième segment est dépourvu 
de sclérification différenciée. 


Cerodirphia avenata araguensis subsp. nov. 
Fig. 9; PL 7 Fig. 15 


Holotype: 1 g', Venezuela, Aragua, rio Choroni, 13.VI.1970 (Mme A. Gadou) (geni- 
talia 4, prép. C. Lemaire n° 2105) (coll. C. Lemaire); allotype: 1 ®, Aragua, route de 
Maracay à Choroni, km 25, 1600 m, 15.IX.1945 (R. Lichy) (sa coll.); paratypes: 
29 g, VIII.1940, VI.1942, IX.1943, IX.1945, VII.1948, IX.1950, VIII.1954, VIII. 
1966, mémes localité, récolteur et collection. 

d'. — Envergure: 70—89 mm; longueur des ailes antérieures: 36—47 mm. 

Antennes jaune paille: 37 articles. Palpes labiaux, front, thorax et pattes brun orangé; 
abdomen noir, annelé de blanc, sur les faces dorsale et latérales, brun foncé ou brun 
orangé sur la face ventrale, touffe anale orangée. 

Face dorsale beige brun ou brun roux plus ou moins orangé, avec une rayure externe 
noire sur chaque aile, celles des ailes antérieures droites et très largement préapicales, 
celles des ailes postérieures convexes. Tache disco-cellulaire des ailes antérieures blanche, 
en forme d’Y couché, dont la branche subcostale renferme un long trait noir, strié de 
blanc, celle des ailes postérieures blanche, en forme de virgule, souvent très développée. 
Dessous des ailes de la même teinte que la face dorsale, même ornementation mais la 
tache disco-cellulaire des ailes antérieures fait défaut ou transparaît faiblement. Costa des 
ailes postérieures blanche sur les deux faces, franges concolores. 

9. — Envergure: 102 mm; longueur des ailes antérieures: 54 mm. 

Couleur fondamentale brun roux orangé sur les deux faces. Même ornementation que 
le mâle. 

Draudt (1930: 769, pl. 116, c 9) a décrit C. avenata de Paname, Chirique, comme 
„forme” de C. speciosa (Cramer) alors qu'il s'agit en fait d'une espèce distincte, 
caractérisée par la teinte, d'un rose très différent, des quatre ailes, par l'effacement des 


150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


Armure génitale 4 des Attacidae. Fig. 6. — 1,Dirphia lichyi sp.n., vue ventralement, pénis détaché; 
2, id., pénis vu latéralement. Fig. 7. — 1, Dirphia crassifurca sp.n., vue ventralement, pénis 
détaché; 2, id., pénis vu latéralement. Fig. 8. — 1, Cerodirphia marahuaca sp.n., vue ventralement, 
pénis détaché; 2, id, même armure vue latéralement: 3, id., pénis vu latéralement; 4, id., sternite 8. 
Fig. 9. — 1, Cerodirphia avenata araguensis subsp.n., vue ventralement, pénis détaché; 2, id., pénis 
vu latéralement; 3, id., sternite 8; 4, id., tergite 8. Fig. 10. — 1, Kentroleuca spitzi sp.n., vue 
ventralement, pénis détaché; 2, id., pénis vu latéralement. L'échelle figurée représente 1 mm 


C. LEMAIRE: Attacidae nouveaux 151 


nervures qui se détachent toujours en noir sur le fond chez C. speciosa et surtout par la 
présence, sur les deux faces des ailes postérieures d'une tache disco-cellulaire blanche. 
L’armure génitale mâle (Fig. 9) diffère de celle de C. speciosa, par la stucture de l’uncus, 
plus large et par l'allongement de la sclérification du huitième tergite. C. avenata 
araguensis est caractérisé par l'envergure moyenne des exemplaires, très supérieure à 
celle de la sous-espèce nominative, par sa coloration dominante brune, plus ou moins 
orangée, jamais rose, et enfin par la netteté des rayures sur les deux faces. 


Kentroleuca spitzi spec. nov. 
Ets @10 PERRE els 


Holotype: 1 9, Brésil, Goiás, Leopoldo de Bulhoes, X.1936 (R. Spitz) (coll. R. 
Spitz < Tring Mus. < British Museum, Nat. Hist.); allotype: 1 4, Brésil, District 
Fédéral, Brasilia, Brasilia Country Club, 19.X.1968 (N. Tangerini) (coll. C. Lemaire); 
paratypes: 2 9, X.1968, X.1969, mêmes localité, récolteur et collection. 

d'. — Envergure: 44—51 mm; longueur des ailes antérieures: 23—27 mm. 

Antennes bipectinées, noires: 54 articles. Palpes labiaux brunâtres, tête et dessus du 
thorax gris jaune, face dorsale de l'abdomen jaune orangé, dessus du corps gris clair, 
taches stigmatales blanches dans de larges stries latérales noires, tarses noirs. 

Apex des ailes antérieures arrondi, obtus, bord externe convexe, tornus très large et 
arrondi. Couleur fondamentale noire sur laquelle se détachent, en jaune, les nervures, 
en blanc, les rayures (l'externe droite), les franges et deux larges stries dont la première 
traverse longitudinalement la cellule et la seconde relie la base au bord externe, entre 
la nervure anale et Cu 1b. 

Ailes postérieures noires, un peu plus sombres sur l'aire baso-médiane, nervures jaunes, 
franges blanches. Face ventrale noire, nervures jaunes, rayures externes blanches, 
nettes sur les quatre ailes; sur les ailes antérieures, mêmes stries blanches que sur la face 
dorsale, mais très estompées. 

9. — Envergure: 58—70 mm; longueur des ailes antérieures: 30—35 mm. 

Ne diffère du mâle que par la présence d’assez vagues rayures externes noiratres ou 
blanchâtres sur le dessus des ailes postérieures. Franges de ces ailes grises. 

Diffère de K. lineosa (Walker) et de K. albilinea (Schaus) par la forme plus arrondie 
des ailes antérieures et, de plus, du premier par l'absence de rayure externe sur la 
face dorsale des ailes postérieures, au moins chez les exemplaires mâles, du second par 
la teinte noire, chez les exemplaires des deux sexes, de l’aire baso-médiane de ces ailes. 
Aspect général plus proche de K. dunkinfieldi (Schaus), mais, chez ce dernier, les deux 
fascies longitudinales blanches des ailes antérieures font défaut, les rayures externes de 
ces ailes sont fortement convexes, celles des ailes postérieures nettes, et les antennes sont 
jaune orangés. 

Armure génitale mâle (Fig. 10) plus grêle que celles des trois espèces précitées; au 
contraire de chez elles, la vesica est dépourvue de cornutus. 


Travassosula mulierata spec. nov. 
Fist Dl7, Fig) 16 


Holotype: 1 &, Pérou, Junin, Chanchamayo, 700 m, VI.1960 (coll. C. Lemaire) ; para- 
type: 1 g', Pérou, Carabaya, rio Huacamayo, La Union, 2000 ft, XII-1904 (saison hu- 


152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


mide) (G. Ockenden) (Tring Mus. = British Museum, Nat. Hist.). 
d'. — Envergure: 48—52 mm; longueur des ailes antérieures: 27—29 mm. 
Antennes noiratres: 32 articles; la longueur des pectinations apicales diminue pro- 
gressivement sur les 12 derniers articles où seules les pectinations basales subsistent. 
Palpes labiaux, tête, thorax bruns, abdomen rouge dorsalement, brunâtre latéralement 
et sur la région anale, rougeâtre ventralement; tarses noirs, avec des écailles violacées. 
Face dorsale des quatre ailes brun terne. Ailes antérieures très allongées, bord externe 
convexe, avec une petite dent subapicale, tornus arrondi. Ailes postérieures présentant vers 
le milieu du bord costal une profonde incision, suivie d'une dent et d'une légère con- 
cavité. Ornementation, sur les deux faces, semblable à celle de T. subfumata (Schaus). 
Le mâle de 7. mulierata ressemble étonnamment à la femelle de T. subfumata et 
diffère donc, par les mêmes caractères externes qu'elle (envergure supérieure, forme 
des quatre ailes, avec l’incision beaucoup plus profonde du bord costal des ailes 
postérieures) du mâle de cette dernière espèce. Armure génitale (Fig. 11) plus robuste 
et structure des valves différente, avec l'extrémité de la costa très sclérifiée, en forme de 
pointe, et le bord interne du sacculus moins renflé. Bulbus ejaculatorius plus long. 


ARSENURINAE 


Copiopteryx semiramis gadouorum subsp. nov. 
PI. 8 Fig. 17 


Holotype: 1 g', Venezuela, Bolivar, El Pao, 450 m, VII-1968 (M. et M®° A. Gadou) 
(coll. C. Lemaire); allotype: 1 9, V.1963; paratypes: 5 g', 1 9, V, VI.1963, VI.1967, 


tous mêmes localité, récolteurs et collection. 


d'. — Envergure à l’apex: 79— 90 mm; longueur des ailes antérieures: 49—53 mm; 
longueur des ailes postérieures: 117—130 mm. 
2. — Envergure: 94—95 mm; longueur des ailes antérieures: 55 m; longueur des 


ailes postérieures: 109 mm. 

Sous-espèce caractérisée par la taille moyenne des exemplaires des deux sexes, très 
nettement inférieure à celle de la sous-espèce nominative: ainsi, chez cette dernière 
qui est connue du Surinam, des Guyane française et ex-anglaise et du Nord du Vene- 
zuela, |’ envergure du mâle se situe entre 105 et 110 mm, de la femelle entre 100 et 125 
mm et la longueur des ailes postérieures atteint et dépasse 160 mm chez le mâle (145 
mm sur la figure de Cramer), 140 mm chez la femelle. C. s. gadouorum se situe donc, 
par la taille, auprès de C. s. montei Gagarin (= C. s. travassosi May) que Travassos 
(1934:412) considérait comme une espèce valable, mais en diffère par un certain nom- 
bre de caractères, particulièrement évidents chez les exemplaires mâles: aspect général 
plus gracile, coloration plus claire, bord externe des ailes antérieures moins oblique 
entre R 1 et l'angle interne, fenêtres disco-cellulaires de ces ailes beaucoup plus étroites, 
queues proportionnellement un peu plus longues mais surtout sensiblement plus grêles 
(largeur 2 mm, 5 contre 3 mm, 5 environ, à mi-distance de la base de l'aile et de 
son extrémité postérieure). Ces caractères sont moins prononcés chez les exemplaires 
femelles, sauf en ce qui concerne la longueur des ailes postérieures et la coloration. 

C'est avec le plus grand plaisir que je dédie cette nouvelle sous-espèce à M. et à Mm 
A. Gadou qui ont personnellement récolté la totalité du matériel typique. 


C. LEMAIRE: Attacidae nouveaux 153 


DRYOCAMPINAE (= ADELOCEPHALINAE)*) 


Eacles imperialis quintanensis subsp. nov. 
Pl. 9 Fig. 18—19 


Holotype: 1 g', Mexique, Quinta-Roo, X-Can, 12.V.1969 (E. C. Welling) (coll. C. 
Lemaire); allotype: 1 9, même date; paratypes: 44 G', 6 9, VI.1967, V.1968, V, VI. 
1969, tous mémes localité, récolteur et collection. 

d'. — Envergure: 89—106 mm; longueur des ailes antérieures: 45—55 mm. 

©. — Envergure: 113—121 mm; longueur des ailes antérieures: 58—65 mm. 

Sous-espèce dimorphique: sur 45 exemplaires mâles, 23 présentent une coloration 
fondamentale jaune, avec une ornementation et des zones brun violacé, tandis que, chez 
les 22 autres, le jaune est partout remplacé par une teinte gris violacé, rarement brun 
rougeâtre, très uniforme et beaucoup plus terne que chez E. imperialis nobilis Neumoe- 
gen. Le méme dimorphisme existe chez les exemplaires femelles, avec semble-t-il, prédo- 
minance des spécimens jaunes (6 sur 7). 

Sur les ailes antérieures les rayures interne et externe sont toujours séparées par une 
large zone médiane alors qu'elles fusionnent au dessus du bord interne chez la plupart 
des exemplaires mâles des sous-espèces néarctiques: E. 7. imperialis (Drury), E. 7. pini 
Michener, E. 7. nobilis Neumoegen, E. 7. oslari W. Rothschild. Les ailes antérieures des 
mâles sont plus allongées et falquées que chez ces quatre sous-espèces, les rayures plus 
étroites sur les quatre ailes; la tache ocellaire subcostale des ailes antérieures est toujours 
très éloignée de la rayure interne au lieu d'être presque constamment recouverte par cette 
dernière. 

Sur ces divers points les mâles d’E. 7. guintanensis, mis à part E. 7. tucumana W. Roth- 
schild, 1907, dont les rayures, surtout l'externe des ailes postérieures sont beaucoup plus 
larges, se rapprochent de ceux des sous-espéces sud-américaines (mais aucune de celles-ci 
n'est dimorphique): E. i. magnifica Walker, 1855, E. i. cacicus (Boisduval), 1868, E. 7. 
opaca Burmeister, 1878, E. 7. approximans Bouvier, 1923. Comme chez ces sous-espèces, 
le dessus du thorax de la femelle, du moins de celle appartenant à la morphe claire, est 
entièrement jaune, à l'exception d’une tache mésothoracique violette; en revanche les 
taches ocellaires sont ici beaucoup plus fortes chez les exemplaires des deux sexes et 
elles présentent, surtout chez les individus à fond jaune, un centre clair beaucoup plus 
contrastant. De plus, sur les ailes antérieures, ces taches sont toujours plus rapprochées, 
en contact ou presque, au lieu d'être le plus souvent isolées. 

Les mêmes caractères séparent la sous-espèce nouvelle d’E. 7. decoris W. Rothschild, 
décrit du Guatémala mais également connu du Mexique (Etats de Mexico, Oaxaca, Vera- 
cruz et Chiapas) qui n'est pas dimorphique, dont l’envergure moyenne, surtout celle du 
mâle est très supérieure et dont la tache sous-costale des ailes antérieures, chez les exem- 
plaires de ce sexe, est plus petite et se perd fréquemment dans la rayure interne. 


*) La désignation de cette sous-famille pose un problème de Nomenclature assez délicat. Le genre 
Ceratocampa Harris, 1833, basé sur deux espéces qui sont en fait les types respectifs des genres 
Citheronia Hübner, 1819 et Eacles Hübner, 1819, n'est pas valable et il en est de même de 
Ceritocampidae Harris, 1841, nom dérivé du groupe famille, par application de l'article 11 (e) 
du Code international de Nomenclature zoologique. Le plus ancien nom utilisable du groupe 
famille devient donc Dryocampini, nom de tribu, proposé par Grote et Robinson en 1866, si, 
comme nous l’estimons, le genre Dryocampa Harris, 1833, est valable. Si au contraire on pense que 
Dryocampa n'est qu'un synonyme ou un sous-genre d’Anisota Hübner, 1820, la famille doit se 
nommer Adelocephalinae Boisduval, 1868 (= Citheroniinae Neumoegen et Dyar, 1894 = 
Sphingicampinae Packard, 1901 — Syssphinginae, Packard, 1905). 


154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


Eacles imperialis anchicayensis subsp. nov. 
Pl. 10 Fig. 20 


Holotype: 1 4, Colombie, Valle Cauca, Anchicaya, 1000/1400 m, 3.X1.1964 (L. et L. 
Denhez) (coll. C. Lemaire); allotype: 1 9, 17.XII.1964; paratypes: 23 g', 10 9, VIII. 
1963, I, VIII à XII.1964, I, VII, VIII, X.1965, VIII.1967, VI.1968, tous mêmes localité, 
récolteurs et collection. 

d. — Envergure: 110—131 mm; longueur des ailes antérieures: 55—65 mm. 

9. — Envergure: 145— 164 mm; longueur des ailes antérieures: 79— 85 mm. 

Cette sous-espèce, non dimorphique, présente un certain nombre de caractères communs 
avec celles déjà connues de |’Amérique centrale et du Sud, tels la forme falquée des ailes 
antérieures du mâle, la coloration jaune, à l'exception d'une tache mésothoracique violette, 
du thorax de la femelle, la position relative des rayures sur la face dorsale des ailes 
antérieures, le diamètre des taches ocellaires chez les exemplaires des deux sexes. De 
toutes ces sous-espèces, d’ailleurs décrites de régions géographiques toutes autres, 
Guatemala et Mexique (E. 7. decoris, et E. 7. quintanensis), Est et Sud-Est du Brésil 
(E. i. magnifica et E. 7. cacicus), Argentine (E. 7. opaca et E. i. tucumana) et Venezuela 
central (E.7. approximans), elle diffère par son envergure beaucoup plus grande, par 
la forme plus allongée et falquée des ailes antérieures du mâle avec l'apex très aigu et 
le bord externe plus fortement concave et enfin par la vivacité de la coloration fondamen- 
tale jaune, contrastant beaucoup avec les zones sombres violet foncé qui sont très intensé- 
ment marquées chez les exemplaires miles. 


Citheronia lichyi spec. nov. 
Fig. 12; Pl. 10 Fig. 21 


Holotype: 1 g', Venezuela, Amazonas, rio Temi, Yavita, 128 m, 9.IX.1947 (R. Lichy) 
(genitalia g', prép. C. Lemaire n° 2267) (coll. R. Lichy); paratype: 1 g', mêmes loca- 
lité, récolteur et collection. 

d'. — Envergure: 69—73 mm; longueur des ailes antérieures: 35—38 mm. 

Antennes jaune orangé: 43 articles dont 15 quadripectinés, suivis de 26 simples. Front, 
tibia et tarses orangés, le reste du corps jaune safran. Ailes antérieures très allongées, 
apex aigu, bord externe convexe, tornus très arrondi et obtus. Couleur fondamentale jaune 
safran; aires interne et externe gris-ardoise sur lesquelles les nervures sont finement 
teintées d'orangé. Bande submarginale orangée, diffuse. Petit point disco-cellulaire 
orangé à centre noirâtre, quelques vagues taches orangées sur l'aire médiane, la plus 
nette entre À et Cu 1b. 

Ailes postérieures jaune clair, passant au jaune safran vers le bord anal, avec seulement 
un point disco-cellulaire rose. Face ventrale semblable à la face dorsale mais l’aire interne 
des ailes antérieures est de la même teinte jaune safran que le fond. 

L'armure génitale (Fig. 12), quoique plus grêle, n'est distincte de celle de C. brissoti 
(Boisduval) que par la structure de l’aedeagus dont le bord ventral est droit au lieu 
d'être fortement concave. 

Cette espèce diffère, par son ornementation, presque aussi réduite que celle de 
C. sepulcralis Grote et Robinson, de toutes les autres espèces du genre. Ainsi, font 
totalement défaut, sur l’abdomen, les anneaux dorsaux polychromes et les taches stigma- 
tales, sur les ailes antérieures, la série de larges taches postmédianes claires, si caractéristi- 


C. LEMAIRE: Attacidae nouveaux 155 


Armure génitale 4 des Attacidae. Fig. 11. — 1, Travassosula mulierata Sp.n., vue ventralement, 
pénis détaché; 2, id., pénis vu latéralement. Fig. 12. — 1, Citheronia lichyi Sp.n., vue ventralement, 
pénis détaché; 2, id., pénis vu latéralement. Fig. 13. — 1, Citheronia andina sp.n., vue ventralement, 
pénis détaché; 2, id., pénis vu latéralement. Fig. 14. — 1, Bouvierina gadonae sp.n., vue ventrale- 


ment, pénis detaché; 2, id. pénis vu ventralement. L'échelle figurée représente 1 mm 


156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


ques de la plupart des Cifheronia, sur les ailes postérieures, les rayures interne et externe. 
C'est avec le plus grand plaisir que je dédie 4 M. René Lichy cette nouvelle espèce, 
certainement une des plus intéressantes de toutes celles qu'il ait récoltées au Venezuela. 


Citheronia andina spec. nov. 
le 135 Tk IO) Joes 22 


Holotype: 1 g', Pérou, Junin, La Merced, 700 m, IX.1963 (G. et L. W. Harris) 
(genitalia g', prép. C. Lemaire n° 1536) (coll. C. Lemaire); paratype: 1 &, Pérou, 
Huallaga-Tai, même collection. 

d'. — Envergure: 101—106 mm; longueur des ailes antérieures: 51—55 mm. 

Antennes jaune orangé: 39 articles dont 15 quadripectinés, suivis de 22 simples. Front 
rouge orangé, une touffe de poils jaunes à la base du pédicelle, thorax rouge orangé, 
avec des touffes de poils jaunes sur le prothorax, le métathorax, la base des ailes an- 
térieures et la face ventrale. Abdomen rouge orangé, annelé de jaune, sur la face dorsale, 
jaune sur la face ventrale; stigmates rouge orangé. Tarses de la même teinte, semés de 
poils jaunes. 

Ailes antérieures très allongées, apex légèrement arrondi, bord externe un peu con- 
vexe. Couleur fondamentale gris ardoise, nervures peu contrastantes, sauf sur le bord 
externe où elles prennent une teinte orange plus marquée. Ornementation, sur les deux 
faces, semblable à celle de la plupart des espèces du genre. Taches postmédianes du 
dessus des ailes antérieures jaune pâle, certaines pouvant être semées d'écailles orange; 
les trois premières, la cinquième, la sixième et la dernière en partant de la costa sont les 
plus larges, la quatrième est parfois absente, la septième très petite. Tache basale jaune 
clair. Autres taches et bande submarginale orange terne. Ailes postérieures jaune pâle, 
avec une large zone basale rouge contenant le plus souvent la tache discocellulaire de la 
même teinte et pouvant même fusionner avec la rayure externe; rouge également, celle-ci 
s'élargit sensiblement vers le bord anal. Bord externe des ailes postérieures convexe. 

Espèce très voisine de C. equatorialis, C. aroa, et C. bellavista*). Diffère nettement des 
deux dernières par la forme des ailes antérieures, plus larges, avec l'apex bien moins 
aigu, le bord externe légérement convexe au lieu d'être absolument droit, le tornus plus 
arrondi. Le bord externe des ailes postérieures est plus franchement convexe, que chez 
C. aroa surtout, et, sur la face dorsale de ces ailes, la zone basale rouge est en général 
plus étendue. 

Beaucoup plus difficilement séparable par les caractères externes de C. egwatorialis, 
si ce n'est que, chez ce dernier, d'après les exemplaires examinés, les deux arceaux 
formés par la bande submarginale, sur la face dorsale des ailes antérieures, entre Cu 1b et 
M 3, sont beaucoup plus larges et entièrement recouverts d’écailles orange au lieu de 
présenter sur leur bord proximal une profonde concavité gris ardoise. 

Les armures génitales des quatre espèces présentent d'excellents caractères qui peuvent 
être ainsi résumés: 

1. Bord interne du sacculus très fortement recourbé vers l'avant, dans le prolongement 


1) L'examen du matériel typique nous a permis d'établir que C. consobrina W. Rothschild, 1907, C. 
mogya Schaus, 1920 et C. mexicana var. brasiliensis Bouvier, 1927, sont des synonymes récents 
de C. aroa Schaus, 1896, syn. nov. En revanche, C. mexicana var. equatorialis Bouvier, 1927 et 
C. mexicana forme bellavista Draudt, 1929 sont des espèces valables. 


C. LEMAIRE: Attacidae nouveaux 157 


du crochet apical, bord postérieur du juxta denté, processus ventral de l’aedeagus rela- 
ementsorelenmn nn SUE GEN 
— Bord interne du sacculus sanken bord posterienr A Amen non denté, aedeagus tres 
robuste à fort processus ventral . . . . OD 
2. Sacculus aussi long que la costa, à crochet afstel dorer res Procent ventral 
de l’aedeagus relativement court, à extrémité apicale droite. . . C. egwatorialis 
— Sacculus nettement plus court que la costa à crochet apical lisse. Processus ventral de 
l’aedeagus très fort, à extrémité apicale aigüe . . . BURN) 
3. Gnathos très étroit. Bord dorsal du processus ventral de Masern muni d’une dent 
tres \protminente. … … . Aa Clare 
— Gnathos plus large. Bord local du processus ai AL l'aedeagus régulièrement 
convexe ou très légérement INCIS (Gore TS) ee Rae andina 
Nous avons reçu C. andina, non seulement de la ocio ars mais également de 
l’Equateur, Oriente et de la Bolivie, Cochabamba. 


Bouvierina gadouae spec. nov. 
Fig. 14; PI. 11 Fig. 23 


Holotype: 1 g', Venezuela, District Fédéral, Caracas (Mme A. Gadou) (genitalia &, 
prép. C. Lemaire n° 1579) (coll. C. Lemaire); allotype: 1 9, Venezuela, Miranda, 
Los Teques (même coll.); paratypes: 34, District Fédéral, Antimano, 900 m, VIII, IX. 
1934, 1 @, Aragua, plage de Puerto Colombia, 15.VIII.1944, 1 9, Caracas Sud, 880 
m, 1.VI.1937 (R. Lichy) (sa coll.), 2 9, Los Teques (coll. C. Lemaire). 

d. — Envergure: 56—62 mm; longueur des ailes antérieures: 28—31 mm. 

Antennes jaune orangé: 43 articles dont 19 quadripectinés, suivis de 22 simples. 
Front et prothorax beige jaunâtre, dessus du thorax de la même teinte, passant au gris 
violacé sur les tegulae; abdomen beige orangé, ombré de grisâtre, sur la face dorsale, gris 
violacé sur la face ventrale. Tarses gris violacé. 

Ailes antérieures peu allongées, apex assez arrondi, bord externe convexe, tornus 
large. Couleur fondamentale beige terne, avec de nombreuses mouchetures gris violacé; 
aires interne et externe gris violacé. Rayure interne très vague, externe plus ou moins 
marquée, nettement préapicale (4 mm), droite ou un peu convexe. Taches apicale et 
subcostale blanches pouvant fusionner. 

Face ventrale gris beige terne, mouchetée de noirâtre; une zone rose vers la base des 
ailes antérieures. Seuls ornements: sur celles-ci, une rayure externe noirâtre ou brunâtre 
et une forte tache disco-cellulaire noire, sur les ailes postérieures, une rayure externe très 
vague. 

Q. — Envergure: 63—78 mm; longueur des ailes antérieures: 30—40 mm. 

Antennes simples. Même coloration que le mâle, avec toutefois le dessus de l'abdomen 
plus orangé, l'aire médiane des ailes antérieures plus claire, les taches subcostales et 
disco-cellulaires blanches de ces ailes remplacées par un halo noirâtre et enfin la zone 
rose de la face ventrale très atténuée. 

Présence, comme chez B. smithi (Druce, 1904) (= andrea Dognin, 1912, syn. nov.) 
de nombreuses mouchetures noirâtres sur la face dorsale des ailes antérieures, mais 
coloration toute autre, de même que la position de la rayure externe de ces ailes, apicale 
chez B. smithi, très éloignée de l’apex chez l’espece nouvelle qui rappelle à cet égard, 
ainsi que par son aspect général, teinte mise à part, B. guadrilineata (Grote et Robinson). 


158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


Armure génitale male (Fig. 14) très différente de celles de B. smithi et B. quadrili- 
neata, notamment par la structure de l’uncus, des valves et de l’aedeagus. 

Je dédie avec plaisir cette espèce inédite 4 Mme A. Gadou 4 qui je suis redevable 
d'une partie du matériel typique. 


Bouvierina ocellata jasonoides subsp. nov. 
Fig. 15; Pl. 11 Fig. 24 


Holotype: 1 g', Colombie, Valle Cauca, Calima, 1800 m, V.1970 (L. et L. Denhez) 
(genitalia g', prép. C. Lemaire n° 1579) (coll. C. Lemaire); paratypes: 3 g', IV.1967, 
X11.1968, 11.1969, mêmes localité, récolteurs et collection. 

d'. — Envergure: 76—85 mm; longueur des ailes antérieures: 42— 46 mm. 

Antennes jaune orangé: 42 articles dont 20 quadripectinés, suivis de 20 simples. Tête 
et dessus du corps orange vif, tegulae violettes, face ventrale de l'abdomen beige violacé, 
tarses violets. 

Ailes antérieures triangulaires: apex très aigu, bord externe droit, très oblique. Couleur 
fondamentale orange vif, passant au violet sur les aires interne et externe, rayures droites, 
violettes. Point disco-cellulaire blanc, pouvant être surmonté d'un minuscule point sub- 
costal de la même teinte. Bord externe des ailes postérieures généralement lobé vers 
langle anal. Coloration dominante de ces ailes orange, passant au rouge sur le bord anal; 
vague rayure externe rouge, droite ou légérement convexe reliant la costa, un peu avant 
apex, au début du dernier tiers du bord anal. 

Face ventrale jaune orangé sur l'aire interne des ailes antérieures, gris violacé sur 
laire externe, de cette dernière teinte, mais plus pâle, sur les ailes postérieures. Forte 
tache disco-cellulaire des ailes antérieures noire à centre blanc. Rayures violettes, celles 
des ailes postérieures incomplètes. 

Par la structure de l’armure génitale mâle (Fig. 15) se situe très pres de B. ocellata 
bidens (W. Rothschild) ; toutefois, d'après le matériel examiné, le gnathos est plus large, 
les harpes, très fines, sont sensiblement plus longues et le bord interne du sacculus 
présente, près de l’extrémité apicale, un renflement qui est généralement absent ou imper- 
ceptible chez B. o. bidens. Diffère surtout de ce dernier par des caractères externes con- 
stants: apex des ailes antérieures plus aigu, coloration beaucoup plus vive de la face 
dorsale de ces ailes, dont le ton orange tend davantage vers le jaune, avec les zones violet- 
tes plus contrastantes. Même différences de teinte mais plus atténuées, sur la face ventrale. 
Bord externe des ailes postérieures lobé et non réguliérement convexe. D’aspect et bien 
que les deux espèces appartiennent 4 deux genres très différents, B. 0. jasonoides res- 
semble surtout étonnamment a Adeloneivaia jason (Boisduval), tant par la forme des 
ailes que par la teinte et l’ornementation, avec pour seul caractère distinctif la coloration 
violette des tegulae. Il n'est pas exclu qu'il s'agisse d'une bonne espèce. 


Scolesa nebulosa spec. nov. 
Fig 168 IDE ail lee, 25) 


Holotype: 1 &, Brésil, Pernambuco (genitalia g', prép. C. Lemaire n° 2399) (coll. 
Frank Johnson = British Museum, Nat. Hist.). 

d'. — Envergure: 52 mm; longueur des ailes antérieures: 26 mm. 

Antennes jaune orangé: 32 articles dont 15 quadripectinés, suivis de 15 simples. 


C. LEMAIRE: Attacidae nouveaux 159 


Front beige, thorax gris violacé, avec une ligne médiane beige sur le métathorax, abdomen 
beige orangé sur les faces dorsale et latérales, gris violacé sur la face ventrale; tarses 
gris violacé. 

Ailes antérieures trés allongées, bord externe convexe, tornus large et arrondi. Couleur 
fondamentale gris violacé, avec des mouchetures noiratres, plus pâle sur les aires interne 
et externe. Pas de rayures. Vague point disco-cellulaire noir. Ailes postérieures gris 
violacé terne, passant au beige vers le bord anal, sans aucune ornementation. Face ven- 
trale uniformément gris violacé, sauf une zone beige au dessus du bord interne des 
ailes antérieures et, sur ces ailes, quelques stries et un point disco-cellulaire noirâtres. 

Aspect général de S. hypoxantha (W. Rothschild), mais absence de rayures et tache dis- 
cocellulaire des ailes antérieures plus petite. Armure génitale mâle (Fig. 16) très dif- 
férente (harpes, processus émanant de la base de la valve gauche et bulbus ejaculatorius 
beaucoup plus courts) et rappelant davantage celle de S. totoma (Schaus), espèce dont 
la coloration et l’ornementation sont toutes autres. La vesica est garnie de trois cornuti 
dont un très allongé: il n'en existe que deux chez S. hypoxantha et S. totoma. 


Ptiloscola descimoni spec. nov. 
Fig M7 IPS 10226 


Holotype: 1 g', Equateur, Occidente, Santo Domingo, 600 m, 17.VII.1968 (S. E. 
Velastegui) (genitalia 4, prép. C. Lemaire n° 1719) (coll. C. Lemaire). 

d'. — Envergure: 55 m; longueur des ailes antérieures: 30 mm. 

Antennes jaune orangé. Tête et thorax beige violacé, avec sur le thorax une ligne 
médiane jaune. Abdomen jaune, finement annelé de noirâtre, sur la face dorsale, brun 
violacé sur la face ventrale. Tarses rose violacé. Dessus des ailes antérieures gris violacé, 
plus clair sur l’aire externe entre le bord interne et Cu 1a. Tache disco-cellulaire réduite 
à un simple trait. Rayures brunes, onduleuses. Bord externe des ailes postérieures 
légérement convexe. Aire baso-médiane de ces ailes brune, aire externe gris violacé. 
Face ventrale gris violacé, assez terne, rayures brunes. 

Espèce ne différant, semble-t-il par aucun caractère externe de P. Jilacina (Schaus, 
1900) (= affinis W. Rothschild, 1907). Armures génitales mâles très différentes: chez 
P. descimoni (Fig. 17), les deux pointes situées latéralement sur le bord dorsal de 
l’uncus sont beaucoup plus longues, le long processus qui émane de la face interne de 
chacune des valves est plus robuste et plus fortement sclérifié, il n'existe pas d’épines à 
la base de ce processus, le lobe gauche du juxta en est également dépourvu et enfin 
l’aedeagus présente latéralement un éperon n’existant chez aucune autre espèce du genre. 

Avec le plus vif plaisir, je dédie cette espèce inédite à mon ami Henri Descimon à 
qui je suis redevable de beaucoup de matériel de l’Equateur, provenant principalement 
de ses récoltes personnelles. 


Ptiloscola dargei spec. nov. 
Bio 18 PL 12 Fis.27 


Holotype: 1 g', Nicaragua, route de Managua à Rivas, km 14,5, 6.IX.1969 (M. Darge) 
(sa coll.); allotype: 1 9, 4.X.1969; paratypes: 12 4, 3 9, VI, IX.1969, tous mêmes 
localité et récolteur, 3 &, Mexique, Chiapas, volcan Tacana, San Jeronimo, 450 m, 
IX.1970 (E. C. Welling), 1 g', Honduras britannique, Stann Creek District, Midd- 


160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


Armure génitale & des Attacidae. Fig. 15. — 1, Bouvierina ocellata jasonoides subsp.n., vue 

ventralement, pénis détaché; 2, id., pénis vu ventralement. Fig. 16. — 1, Scolesa nebulosa sp.n., 

vue ventralement, pénis détaché; 2, id., pénis vu latéralement. Fig. 17. — 1, Piiloscola descimoni 

sp.n., vue ventralement, pénis détaché; 2, id., pénis vu ventralement. Fig. 18. — 1, Priloscola dargei 

sp.n., vue ventralement, pénis détaché; 2, id., pénis vu ventralement. Fig. 19. — 1, Ptiloscola wel- 

lingi sp.n., vue ventralement, pénis détaché; 2, id., pénis vu ventralement. L'échelle figurée repré- 
sente 1 mm 


C. LEMAIRE: Attacidae nouveaux 161 


lesex, 3.VII.1964 (même récolteur) (tous coll. C. Lemaire, sauf 9 paratypes G', 2 
paratypes ® de Managua, #7 coll. M. Darge). 

d'. — Envergure: 48—52 mm; longueur des ailes antérieures: 25—27 mm. 

Antennes jaune orangé: 34 articles dont 13 quadripectinés, suivis de 19 simples. 
Front brun, dessus du thorax violet, avec une ligne médiane jaune de largeur variable, 
parfois absente. Face dorsale de l’abdomen jaune, annelée de noirâtre, dessous du corps 
brun violacé, tarses roses. Ailes antérieures brun violacé, plus ou moins foncé, plus 
sombres sur laire médiane et souvent sur l'aire interne. Rayures brunes, légérement 
onduleuses, tache disco-cellulaire réduite 4 un simple trait où à un point. Bord externe 
des ailes postérieures convexe, aire baso-médiane de ces ailes brune, aire externe brun 
violacé. Face ventrale des quatre ailes de la même teinte mais passant fréquemment au 
noirâtre sur l'aire baso-médiane, rayures brun noir. 

®. — Envergure: 66 mm; longueur des ailes antérieures: 35 mm. 

Antennes simples, jaune foncé. Tête et dessus du thorax bruns, avec une très étroite 
ligne médiane jaune sur le thorax. Ailes beaucoup plus larges que celles du mâle, même 
ornementation; coloration dominante brune sur les deux faces. 

Diffère surtout par la structure de l’armure génitale mâle (Fig. 18) de P. descimoni 
et P. lilacina. Les valves sont plus courtes que chez ces deux espèces, le processus 
émanant de la face interne de chacune d'elles est moins développé et moins sclérifié que 
chez P. descimoni et les épines qui se dressent à la base de ces processus, comme chez 
P. lilacina, sont ici beaucoup plus courtes. Uncus semblable à celui de P. Jilacina, juxta 
et aedeagus également, mais ce dernier beaucoup moins robuste. 

Je suis heureux de pouvoir dédier cette espéce nouvelle 4 M. Michel Darge qui a 
récolté au Nicaragua la plupart du matériel typique et a bien voulu m'en confier l’étude. 


Ptiloscola wellingi spec. nov. 
Fig: Pl. 12). Bic. 28 


Holotype: 1 &, Mexique, Quintana-Roo, X-Can, 18.V.1969 (E. C. Welling) (coll. 
C. Lemaire); allotype: 1 9, 20.V.1969; paratypes: 10 g', 2 2, même localité, VI,XI. 
1967, V.1969, 5 g', Yucatan, Chaksikin, Uman, VI, XI.1967, V.1969, tous mêmes 
récolteur et collection. 

d. — Envergure: 37—46 mm; longueur des ailes antérieures: 20—23 mm. 

Antennes jaune clair: 41 articles dont 18 quadripectinés, suivis de 21 simples. Téte 
et thorax de coloration extrêmement variable, du brun au beige clair et au beige orangé, 
tegulae fréquemment gris violacé; abdomen jaune ou beige orangé presque toujours 
annelé de noir, beige violacé, plus ou moins foncé sur la face dorsale; tarses rose 
violacé. 

Coloration du dessus des quatre ailes également trés variable mais toujours claire, 
beige, beige orangé, gris violacé. Ornementation habituelle des espéces du genre: rayures 
noiratres, plus ou moins violacées, un peu lunulaires, surtout sur les ailes antérieures, 
celles des ailes postérieures s’estompant fréquemment avant le bord anal. Tache disco- 
cellulaire des ailes antérieures généralement blanche, pouvant être accompagnée d'une 
petite tache annexe de la même teinte, entre M 3 et M 2, ou parfois réduite à un point 
ou à un trait noirätre. Bord externe des ailes postérieures convexe. Face ventrale de la 
même teinte que la face dorsale. Rayures noirâtre violacé, celles des ailes antérieures 
pouvant être partiellement effacées. 


162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


Q. — Envergure: 50—54 mm; longueur des ailes antérieures: 26—28 mm. 

Antennes simples. Coloration du corps présentant vraisemblablement les mémes va- 
riations que chez les exemplaires males. Ailes beaucoup plus larges, à fond brun clair ou 
beige orangé chez les spécimens examinés. Tache disco-cellulaire des ailes antérieures 
réduite 4 un simple trait ou halo. 

Espèce voisine de P. surrotunda (Dyar). En diffère par son envergure sensiblement 
inférieure et par sa coloration beaucoup plus claire. L’armure génitale mâle (Fig. 19) 
est beaucoup plus gréle, les longs processus émanant de la face interne des valves ne 
sont pas droits mais recourbés vers l’apex, l’extrémité postérieure du lobe droit du juxta 
est moins sclérifiée et moins abondamment garnie d’épines et surtout le lobe gauche de la 
même pièce est dépourvu de la touffe d’épines très sclérifiée qui existe chez P. surrotunda. 

Cette espèce inédite est dédiée à mon excellent correspondant mexicain, M. E. C. Wel- 
ling à qui je suis notamment redevable de la très belle série typique. 


AUTEURS CITES 


Bouvier, E. L., 1924. Sur les Saturniens Hémileucides du groupe de Dirphia targuinia Cram. (Hy- 

perdirphia Packard, 1903). — Ann. Soc. ent. France, 93, pp. 381—389, 1 fig. phot. 

, 1936. Etude des Saturnioides Normaux. Famille des Saturniidés. — Mém. Mus. nat. Hist. 

nat., (N.S.) 3 (fasc. unique), 354 p., 82 fig., 12 pl. phot. h.-t. 

Draudt, M., 1929—1932. 12 Familie: Saturnidae [sic], in A. Seitz, Die Gross-Schmetterlinge der 
Erde, vol. 6 (Die Amerikanischen Spinner und Schwärmer), pp. 713—827, pl. col. ht. 
101—137 et 142. A. Kernen, Stuttgart. 

Hoffmann, C. C., 1942. Catalogo Systematico y Zoogeografico de los lepidopteros mexicanos. Ter- 
cera parte: Sphingoidea y Saturnioidea. — An. Inst. Biol., Mexico, 13, pp. 213—256. 

Rothschild, W., 1895. Notes on Saturniidae: with a preliminary revision of the family down to the 
genus Automeris, and descriptions of some new species. — Nov. Zool., 2, pp. 35—51. 

Schüssler, H., 1934. — Saturniidae: 2. Subfam. Saturniinae II et 3. Subfam. Ludiinae I., iz E. 
Strand, Lepidopterorum Catalogus, Pars 58, pp. 325—484. W. Junk, Berlin. 

Travassos, L., 1934. Copiopteryx montei Gagarin, 1934 (Lep. Saturniidae). — Rev. de Ent., Rio de 
Janeiro, 4 (3), p. 412. 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 1 


Pl. 1, Fig. 1, Copaxa (Copaxa) escalantei sp.n., holotype &. Fig. 2, Copaxa (Copaxa) andensis 
sp.n., holotype & 


C. LEMAIRE: Attacidae nouveaux 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 2 


PI. 


2, Fig. 3, Copaxa (Copaxa) denhezi sp.n., holotype 4. Fig. 4, Copaxa (Copaxa) canella miranda 
subsp. n., holotype & 


C. LEMAIRE: Aftacidae nouveaux 


rn 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 3 


Pl. 3, Fig. 5, Copaxa (Copaxa) canella miranda subsp. n., paratype 9. Fig. 6, Copaxa (Sagana) 
herbuloti sp. n., holotype & 


C. LEMAIRE: Attacidae nouveaux 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 4 


Pl. 4, Fig. 7, Rothschildia paucidentata sp. n., paratype ¢. Fig. 8, Rothschildia lebeau yucatana 
subsp. n., holotype & 


C. LEMAIRE: Aftacidae nouveaux 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 


PI. 5, Fig. 9, Dirphia lichyi sp.n., holotype &. Fig. 10, id., allotype 9 


C. LEMAIRE: Attacidae nouveaux 


‘TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 6 


11 


12 


13 


Pl. 6, Fig. 11, Dirphia crassifurca sp. n., holotype 4. Fig. 12, id., allotype 2. Fig. 13, Kentroleuca 


spitzi sp. n., holotype ® 


C. LEMAIRE : Attacidae nouveaux 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 7 


14 


15 


16 


Pl. 7, Fig. 14, Cerodirphia marahuaca Sp. n., paratype 4. Fig. 15, Cerodirphia avenata araguensis 
subsp. n., paratype 4. Fig. 16. Travassosula mulierata sp. n., paratype & 


C. LEMAIRE: Attacidae nouveaux 


a eae 


9 od&ojoy “u ‘dsqns wnvonopvd Stuvi was x(uardordor “Li ‘314 ‘8 ‘Id 


PLATE 8 
a 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 9 


18 


19 


Pl. 9, Fig. 18, Eacles imperialis quintanensis subsp. n., holotype &. Fig. 19, id. paratype & 
(morphe jaune) 


C. LEMAIRE: Attacidae nouveaux 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 10 


20 


21 


22 


PI. 10, Fig. 20, Eacles imperialis anchicayensis subsp. n., holotype &. Fig. 21, Citheronia lichyi 
Sp. n., paratype &. Fig. 22, Citheronia andina sp. n., & 


C. LEMAIRE: Attacidae nouveaux 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 11 


23 


24 


Pl 11, Fig 23, Bouvierina gadouae sp. n., holotype &. Fig. 24, Bouvterina ocellata jasonoides 
subsp. n., paratype &. Fig. 25, Scolesa nebulosa sp. n., holotype 4 


C. LEMAIRE: Aftacidae nouveaux 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 3, 1971 PLATE 


26 


Pl. 12, Fig. 26, Ptiloscola descimoni sp. n., holotype &. Fig. 27, Ptiloscola dargei sp. n., holotype 
6. Fig. 28, Priloscola welling: sp. n., holotype & 


C. LEMAIRE: Attacidae nouveaux 


No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.FI. 45.—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.Fl. 70.—. 


No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Ditsh & Uvarov 
and Xenocatantops Ditsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. 
D.Fl. 25.—. 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. 
D.FL 75.—. 


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F. S. LukoscHus & F. M. DRIESSEN. — Amorphacarus parvisetosus spec. nov. (Myobii- 
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AMORPHACARUS PARVISETOSUS SPEC. NOV. (MYOBIIDAE, 
TROMBIDIFORMES), FROM NEOMYS FODIENS PENNANT 
(SORICIDAE) 
by 
F. S. LUKOSCHUS & F. M. DRIESSEN!) 


Zoological Institute, Catholic University, Nijmegen 
ABSTRACT 


An undescribed species of Amorphacarus (Myobiidae), has been found in the head and shoulder 
region of the water shrew, Neomys fodiens Penn. (Soricidae). Remarks on the genus Amorphacarus 
are appended. 


On investigating a number of Neomys fodiens Pennant, caught near Nijmegen, we 
found in the head and shoulder regions a number of mites of the family Myobiidae 
which deviate considerably from species so far described. 

The following characteristics: legs of the first pair unequal, one being dwarfed, 
segment I longer than broad, segment II much broader than long with ventral tubercle, 
tarsus II with two claws, most of the hairs with very fine spatulate points, some of the 
dorsal setae slightly inflated, very long and double and coiled penis, all point to the 
genus Amorphacarus Ewing, 1938. The species of this genus are skin parasites of 
Soricidae (Poppe, 1896; Jameson, 1948; Ewing, 1938; Radford, 1949; Dubinin, 1957; 
Jameson 1971). 


Amorphacarus parvisetosus spec. nov. 


Female (holotype). — Body broader than in other species of the genus. Length 
including gnathosoma 570 u, in 17 paratypes measured 9 504 u (431—571), width 
356 u, in the paratypes 9 340 u (263—364). 

Dorsum (Fig. 1). Body transversely striated with exception of the vulvar and genital 
regions. Pores of dorsal glands (P) between vertical internal and vertical external (v 7 
and v e). Three pairs of long filiform setae, very finely striated lengthwise v e 
(100 u), scapular external (sc e) 200 u, lateral 1 (/ 1) 220 y. Vertical internal (v 7), 
scapular internal (sc z) and the file of dorsal setae (d1—d4) short, cored (Howell & 
Elzinga, 1962) with fine spatulate point (detail d 1). Lateral setae 4 (/ 4) short, 
setiform. Laterals 2 and dorsals 5 lacking. Genital opening terminally with five 
pairs of genital setae (gI—g5), g 4 and g 5 functionally arranged to copulatory 
complex. Vulva (Vw) surrounded by two weakly sclerotized vulvar valves with genital 
hooks (4 3), which usually cover opening of bursa copulatrix. Anal internal (a 7) and 
anal external (4 e) setae short, almost spine-like (Fig. 2). 

Venter (Fig. 3). — Ventral side delicately striated. Back of coxal field I reshaped 
to tubercles standing caudally, more strongly sclerotized. Setae of coxal regions setiform. 
Present are cx I 1, 2, 4, cx II 1, 3, 4, cx III 1, cx IV 1. Lateral setae 5 (/ 5) of about 
330 u. 


1) Present address: Nederlands Instituut voor Zuivel-Onderzoek, Ede, Netherlands. 


163 


164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 1—8. Amorphacarus parvisetosus spec. nov., female (holotype), 1, female dosum; 2, vulva 
region; 3, female venter; 4, broad lateral seta 1 of a paratype; 5, gnathosoma dorsal; 6, broad lateral 
seta 5 of a paratype; 7, leg I dorsum; 8, leg I venter 


S. LUKOSCHUS & M. DRIESSEN: Amorphacarus parvisetosus 165 


Gnathosoma (Fig. 5). — Gnathosoma specialized, with stinging and sucking appara- 
tus. Chelicerae (Ch) with barbed hooks on the outside. Pedipalps and hypostome (Hyp) 
closing mouth opening on lateral and ventral side. Pedipalps of two movable segments, 
ventral long claw-shaped tibia (77), surrounding tarsus (Ta), both segments carrying 
one hair each. Gnathosoma with two ventral hairs, rostral anterior (ra) and rostral 
posterior (rp). Dorsal to basis of pedipalps rostral dorsal setae (rd), directed towards 
the mouth opening. Stigmata (sg) dorsal at the back of gnathosoma, no stigmal hairs. 

Legs, ventro-lateral. All legs with five free segments (trochanter (Tr), femur (Fe), 
genu (G), tibia (77), tarsus (Ta) and two claws (K/). Legs II—IV with pretarsus 
(Pir), claws situated dorsally (Fig. 14, 16, 18). 

Legs on the first pair (Fig. 7, 8) inequal, eight paratypes measured have a longer right 
leg, nine a longer left one. Proportion great leg/small leg, 9 1.163 (1.06—1.23). Tarsi 
with very small claws (K/), ventro-terminally inserted, without noticeable pretarsus. 
Claws hair-like in adults, but well developed in subadult stages (Fig. 26). When 
comparing species of Myobiidae one comes to the conclusion that claws are present 
instead of setae. There is no species with more than six setae and three solenidia on 
tarsus I. As in A. parvisetosus there are six well-shaped setae and three solenidia 
(w 1-3), the other two very little spine-like forms must be reduced claws. Except 
long spatulate p d all tarsal setae are setiform. Genu postero-lateral with large clasping 
tubercle. We are not quite sure, whether a very little solenidion sigma is present dorso- 
mediad near anterior border of genu. Femur with broad tubercle, movable enlarged 
hair (pv) and dorsally a long and thick, lengthwise striated postero-dorsal seta (pd), 
characteristic for many myobiids, and a short 72 d, also striated. 

Legs II with strong claws of unequal size, dorsally on the pretarsus (Fig. 14, 15). 
Solenidia omega 1 postero-lateral and omega 2 antero-dorsal. The medio-dorsal hair 
of the tarsus is long and setiform in the female (Fig. 1), stout, striated and spatulate 
at the point in the male (Fig. 14). Ventral hairs seti to filiform and non striated. 
Solenidion sigma distinct dorso-mediad on genu. 

Legs III and IV (Fig. 16—19) with claws of very different size. A part of ventral 
setae of tarsus, tibia and genu are enlarged and striated lengthwise. 

Male (allotype). — Body broader than in other species. Length, including gnatho- 
soma, 319 u, in 14 paratypes measured 9 328 u (314—347); width 180 y, in the 
paratypes @ 194 u (179—224). 

Dorsum (Fig. 9). — Setae ve, sce and /1 long, filiform. Setae vi, sci and /4 
very short. The file of dorsals 1—5 is lacking. Genital opening (Fig. 11) at level of 
legs III. Genital orifice surrounded by four shields: one short and broad with two 
pairs of setae, two little shields with one seta each, and a middle scutcheon-shaped 
sclerotization with two large hairs forming a sledge for the penis, a pair of broad, 
barbed striated hairs and a pair of little setiform ones. The penis begins (also in all 
paratypes) median above legs IV, bends backwards to the left and then continues with 
two loopings towards the genital opening. 

Venter (Fig. 10). Gnathosoma and hairs in coxal regions as in female. 

Leg I (Fig. 12, 13) different from that in female: setae p d of tarsi remarkably longer 
than in female, setae m d of genu short, stout and striated lengthwise. Seven paratypes 
measured have a longer right leg and seven other a longer left leg, proportion great 
leg/small leg o 1.23 (1.14—1.31). 


166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


200p Hi 1 1 Jsgy 
9 — 10 ll — 19 


Fig. 9—19. Amorphacarus parvisetosus spec. nov, male (allotype). 9, male dorsum; 10, male 
venter; 11, genital region; 12, leg I dorsum; 13, leg I venter; 14, tarsus II dorsum; 15, tarsus II 
venter; 16, tarsus III dorsum; 17, tarsus III venter; 18, tarsus IV dorsum; 19, tarsus IV venter 


S. LukoscHus & M. DRIESSEN : Amorphacarus parvisetosus 167 


Chaetotaxy 
legs I II III IV 
tarsus 6 + (3) 6 + (2) 6 6 solenidia in () 
claws 2 2 2 2 
tibia 6 6 6 6 
genu 8 + (1?) 6 + (1) 6 6 
femur 5 5 2 2 
trochanter 3 2 3 3 


IMMATURE DEVELOPMENTAL STAGES 


As hitherto neither the larval nor the nymphal stages of Amorphacarus species have 
been described nor drawn, nor their chaetotaxy, we present detailed figures for A. par- 
viselosus. 

Eggs. — Fully developed eggs within eight females (Fig. 3), length 9 89 u (83—95). 
Eggs attached lice-like to basis of hairs. 

Common to all developmental stages are legs with four free segments. Presence of 
solenidion sigma on genu II indicates fusion of genu and femur. Legs I with two big 
claws ventrally without pretarsus. Tarsus I with one dorsal and one ventral big clasping 
tubercle. Legs II—IV with pretarsus and one claw, only leg II in the tritonymph with 
two claws. Gnathosoma long, cylindrical and split at the end. Dorsal pore (P) situated 
far above vertical external. Body transversely finely striated, with exception of ventral 
side of legs II, where cross-striation bulges caudally. 

Larva. — Length including legs I, 210 u, width 110 y. 

Dorsum (Fig. 21) demonstrates the primitive phylogenetical pattern present in all 
genera from insectivores of the third evolutionary branch of Myobiidae (Dusbabek, 
1969). Present are ve, sci, sce, / 1, 1 3,15, d 1,4 2, d 3, d 5. The hairs are blade-like, 
barbed and striated similar to the description of dorsals 1—3 of females from A. 
elongatus, called “schuppenförmig, zweigliedrig” by Poppe (1896). Vulva medio-sub- 
terminal. 

Venter (Fig. 20). Present are only ex I 1. Legs I unequal (quotient 1.18). Femur I 
with remarkably long, setiform p d. 

Protonymph. — Length including gnathosoma 219 y, width 163 u. Dorsum (Fig. 23). 
Less setae than in the larva, hairs relatively small and only partly slightly barbed. Present 
are ve, sci, sce, 11 and / 4. Like in protonymphs of genera of third evolutionary branch 
the file of dorsal setae and / 3 are lacking; p d seta on femur I stout, conical, striated. 
Trochanteres II—IV without setae. 

Venter (Fig. 22). Added are peg-shaped and striated second seta in coxal field I 
and setiform first setae in coxal fields II and III. 

Deutonymph. — Length including gnathosoma 255 y, width 175 u. 

Dorsum (Fig. 25). Chaetotaxy as in the protonymph. Trochanteres I—III with one 
seta each. Legs I unequal (quotient 1.10). 

Venter (Fig. 24). Setation as in the protonymph, additional cx IV 1. 

Tritonymph. — Length including gnathosoma o 354  (275—431), width 0 223 u 
(oh) 

Dorsum (Fig. 29). Setation like deutonymph, additional dorsal 5, vertical internal and 
trochanter seta on leg IV. 


168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 20—27. Amorphacarus parvisetosus spec. nov., 20, larva venter; 21, larva dorsum; 22, pro- 
tonymph venter; 23, protonymph dorsum; 24, deutonymph venter; 25, deutonymph dorsum; 26, 
tritonymph leg I venter; 27, tritonymph leg I dorsum | 


S. LukoscHUs & M. DRIESSEN : Amorphacarus parvisetosus 169 


Venter (Fig. 28). Additional to setation of deutonymph are rostral posterior, enlarged 
cx I 3 and setiform cx II 3. Legs I unequal (quotient 1.10). Tarsus I ventrally (Fig. 26) 
with two big claws and a clasping tubercle. Enlarged hairs on tibia and genu-femur 
with probably clasping function. Tarsus I dorsal (Fig. 27) with large clasping tubercle, 
pd of femur I short and stout. 


Chaetotaxy of subadult stages 


larva protonymph deutonymph tritonymph 
legs I II III Ih EE B IV: I I II IV I I III IV 
claws 2 1 1 2 1 1 1 2 1 1 1 2 1 1 1 
tarsus 2 6 6 4 6 6 6 5 6 6. 6 6 6 6 6 
solenidia 1 2 — 2 2 — — 2 2 — — 2 2 — — 
tibia 5 5 4 Gee wo) Aard GG. Bo a CNG? = oe 
genu-femur 2. 2.0 — D Bi Bo SIA we 2002 Sal MR 
trochanter — | — | Se te — Sil 


Type host: Neomys fodiens Pennant. 

Type locality: Nijmegen, the Netherlands, 15.1X.1967, leg. F. Lukoschus. On the 
same host and locality: 24.IX.1964, 12.V.1967, 12.IX.1967. Isle of Texel, The Nether- 
lands, 26.IX.1968. Gmünd, Austria, 16.VII.1966, 16.X.1966, Wien, Austria, 1967 
(hosts preserved in alcohol, in Naturhistorisches Museum Wien, by courtesy of 
Dr. K. Bauer). 

Miss M. Kolebinova, Zoological Institute, Academy of Sciences, Sofia, found the 
species on the same host near Witescha, Bulgaria, 23.VIII.1966. Probably the mite is 
specific for the host throughout its range. 

Pathogeny. — The mites were found at the base of the hairs in head and shoulder 
parts. Larvae and nymphs prove to be cell-fluid suckers, only adults are found with 
blood in the intestines. In 4 out of 41 adults one of the legs IV shows stump-shaped 
regeneration, that points to a lesion caused by self-defence of the host. 


IDENTIFICATION OF THE SPECIES 


Especially the size of the dorsal setae 2 and 3 are important, as well as the presence 
of dorsals 5. 


Table I. Characteristics of females of Amorphacarus species. Data in u eX = barbed hairs. 
dates taken from d 2 d 3 12 d 4 13 d 5 
A. elongatus description + 65 X 70X — 28 X 27 X 27X 


specimens from 
the type host 

A. hengererorum description + 41X 42X 31X 17 20 — 
holotype 

À. parvisetosus types 5 4 — 6 6 — 


170 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


34 


38 


Fig. 28—29. Amorphacarus parvisetosus spec. nov. 28, tritonymph venter; 29, tritonymph dorsum; 
30—38. Amorphacarus elongatus (Poppe, 1896), 30, leg I venter; 31, leg I dorsum; 32, tarsus II 
venter; 33, tarsus II dorsum; 34, tarsus III venter; 35, tarsus III dorsum; 37, tarsus IV dorsum; 
38, dorsal seta 1; 36, Amorphacarus hengererorum Jameson, 1948 (holotype), tarsus IV dorsum 


S. LUKOSCHUS & M. DRIESSEN : Amorphacarus parvisetosus 171 
REMARKS ON THE CHARACTERISTICS OF Amorphacarus 


Genus Amorphacarus has been erected by Ewing (1938) for A. elongatus Poppe, 1896, 
and was considered monotypic at that time. The description was valid only for females. 
Males were not available for study. The following characteristics were given: 

“Legs of the first pair unequal, one being dwarfed; segment I longer than broad, 
inner margin concave; segment II much broader than long, with dorsal tubercle; seg- 
ment III distinct from IV, bearing large clasping tubercle; segment IV broader than 
long, rounded distally. Capitulum asymmetrical. Tarsus II with two claws; tarsi III and 
IV each with one claw. Some of dorsal setae inflated or foliaceous.” 

Jameson, describing the second species, A. hengererorum (1948), gives the following 
characteristics (1955): Leg I apparently of three segments, the fourth sometimes con- 
cealed beneath the third; one pair of terminal claws; the first pair of legs markedly 
unequal in size. Leg II with a pair of claws; legs III and IV with a single claw. Dorsal 
setae slightly inflated, non-striated. Body shape rather slender. Penis very long and 
double coiled. From shrews (Soricidae).” 

Radford (1948, 1949): "Capitulum and leg I asymmetrical; tarsus II with two claws 
in the female, one in the male; tarsi III and IV with one claw. Leg I with three or four 
segments without tarsal claws’. 

The characteristics mentioned above for A. parvisetosus make extension of the diagnosis 
of the genus Amorphacarus necessary. 

In view of the differences given by the various authors, it appears worth while to 
look for more details in A. elongatus with more modern optical tools. 

We are able to state, that: 

1) Leg I of A.elongatus from the type-host Sorex araneus (Fig. 30, 31) has five seg- 
ments and a pair of very little claws. Though the last three segments are compressed, 
their existence is furthermore demonstrated by the complete chaetotaxy, as in the more 
primitive genera Protomyobia Ewing, 1938, Eadiea Jameson, 1948, Pteracarus Jame- 
son & Chow, 1952. Compare also Jameson (1955), fig. 2, Eadiea desmanae Lukoschus 
1969 and Eadiea multisetosa Lukoschus & Driessen (1969). 

2) The legs II-IV (Fig. 32—35, 37) have two claws. These are very difficult to dis- 
tinguish in the legs III and IV, the posterior claw being even smaller than in the species 
described above and in Archemyobia inexpectata Jameson, 1955. 

3) All the body hairs show slight spatulate points, as already reported by Poppe 
(1896): “Eine besonders langgestreckte Art, deren Borstenbesatz der Dorsalseite des 
@ dadurch ausgezeichnet ist, dass sämtliche Borsten an ihrem distalen Ende abgestumpft 
sind” (Fig. 38). 

4) In the type-specimens of A. hengererorum Jameson, 1948, the body hairs have also 
a slightly spatulate point. In the female (holotype) leg IV has also a small second 
claw, very difficult to distinguish (Fig. 36). 

New generic characteristics proposed are as follows: Leg I with five segments. Seg- 
ments III-V compressed. First pair of legs asymmetrical in size. Body setae with slightly 
spatulate points. Penis very long and double-coiled. Claws very variable. Legs I with two 
very small or without claws. On leg II the claws are subequal-unequal or there is only 
one claw. Legs III and IV with very unequal claws or with only one claw. From shrews 
(Soricidae). Genotype Amorphacarus elongatus Poppe, 1896 (syn. Myobia elongata Pop- 
pe 1896): 


172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


When regarding the very long and double-coiled penis, the setae with slightly spatu- 
late points, and the asymmetrical size of legs I as most important characteristics, and not 
the number and the size of the claws, it may be possible to include into the genus 
Amorphacarus species from Soricidae in America and Asia, to be described in the near 
future by E. W. Jameson, jr., Davis, California, as well as Blarinobia species. 

Types. — Holotype 9 and allotype g': Rijksmuseum van Natuurlijke Historie, Lei- 
den. Paratypes 9 and ¢: British Museum (Natural History), Londen; U.S. National 
Museum, Washington; Zoologische Staatssammlung, München; Department of Zoology, 
University of California, Davis (U.S.A.); Zoological Institute, Academy of Sciences, 
Leningrad; Muséum National d'Histoire Naturelle, Paris; Instituut voor Tropische Ge- 
neeskunde Prins Leopold, Antwerpen; Bulgarische Akademie der Wissenschaften, Zoo- 
logisches Institut, Sofia; Zoölogisch Laboratorium der Katholieke Universiteit, Nijme- 
gen’). 

The institutes mentioned above will also be provided with specimens of A. elongatus 
(Poppe, 1896) from the type-host Sorex araneus, with well visible claws on the tarsi III 
and IV. 

Acknowledgement. — We are indebted to Prof. Dr. E. W. Jameson, Davis, U.S.A., 
for his discussions and for reading the manscript. 


REFERENCES 


Dubinin, V. B., 1957. A new classification of the mites of the superfamilies Cheyletoidea W. Dub. 
and Demodicoidea W. Dub. (Acariformes, Trombidiformes). Parasitol. Rev. Zool. Inst. 
Acad. Sci. U.S.S.R. 17: 71—136. 

Ewing, H. E., 1938. North American mites of the subfamily Myobiidae, new subfamily (Arachnida). 
Proc. Ent. Soc. Wash. 40: 180—197. 

Howell, J. F. & J. Elzinga, 1962. A new Radfordia (Acarina: Myobiidae) from the Kangaroo rat 
and a key to the known species. Ann. Ent. Soc. Amer. 55: 547—555. 

Jameson, E. W., 1948. Myobiid mites (Acarina : Myobiidae) from shrews (Mammalia : Soricidae) 

of eastern North America. Journ. Parasitol. 34: 336-—342. 

, 1955. A summary of the genera of Myobiidae (Acarina). Journ. Parasitol. 41 : 407—416. 

Jameson, E. W. & C. Y. Chow, 1952. Pteracarus, a new genus of myobiid mites (Acarina : Myo- 
biidae) from bats (Mammalia : Chiroptera). Journ. Parasitol. 38: 218—221. 

Lukoschus, F. S., 1969. Eadiea desmanae spec. nov. (Acarina : Myobiidae) von Galemys pyrenaicus. 
Acarologia 11 (3): 575—584. 

Lukoschus, F. S. & F. M. Driessen, 1969. Eadiea multisetosa spec. nov. (Myobiidae : Thrombidi- 
formes) von Crocidura russula. Zool. Anz. 182: 383—390. 

Poppe, S. A., 1896. Beitrag zur Kenntnis der Gattung Myobia v. Heyden. Zool. Anz. 19: 337—349. 

Radford, Ch. D., 1948. A revision of the fur mites Myobiidae (Acarina). Bull. Mus. Nat. Hist. 
Paris, 2e sér. 20: 458—464. 

—, 1949. A revision of the fur mites Myobiidae (Acarina) (suite). Bull. Mus. Nat. Hist. Paris 
2e sér., 21: 91—97. 


1) The coll. nos. received are: 


A. parvisetosus A. elongatus 
Leiden P 1208-8 P 1210 
Paris Sey JJ ADD J) Ue 55] 2 
Hamburg A 43 | 69 


Miinchen P 495 if 1-2 P 494 / 1 


LIFE CYCLE OF MELESODECTES AURICULARIS 
FAIN & LUKOSCHUS (GLYCYPHAGIDAE, 
SARCOPTIFORMES) 


by 
F. S. LUKOSCHUS, A. W. A. M. DE COCK 


Zoological Institute, Catholic University, Nijmegen, Netherlands 
and 


A. FAIN 
Institut de Médecine Tropicale Prince Léopold, Antwerpen, Belgium 


ABSTRACT 


Rearing of hypopi of Melesodectes auricularis Fain & Lukoschus, 1968, was successful. Mor- 
phology suggests relationship to Hypoderidae and Saproglyphidae. 


INTRODUCTION 


In a former paper Fain and Lukoschus (1968) described hypopus and tritonymph of 
this species found in the concha of European badgers. Hypopi lacking attaching organs 
on idiosoma (sucker plates or claspers) or on tibiae of forelegs show similarities to 
hypopi of the family Hypoderidae, but they have more primitive legs. 

In the meantime we succeeded in rearing hypopi from badgers killed by trucks in 
the spring. We are very obliged to the Rijksinstituut voor Veldbiologisch Onderzoek 
(RIVON) at Zeist and to Dr. Franz Krapp, Zoologisch vergl.-anatomisches Institut 
Freiburg (Switzerland) for forwarding us parasitized conchae of badgers. 


METHODS AND RESULTS OF REARING EXPERIMENTS 


Original material was obtained from the ears of European badgers (Meles meles): 
28.111.1969 near Grimentz (Kanton Wallis), Switzerland, 17.1V.1969 near Freiburg, 
Switzerland, 20.1V.1969 near Vierlingsbeek (Prov. Limburg), the Netherlands. They 
were lactating females. A starved suckling 20.1V.1969 near Vierlingsbeek. 

The conchae sent to us were partly scraped out, other parts containing hypopi have 
been divided into pieces with: 

1) only epidermis, with ear-wax; 2) epidermis with fatty connective tissues; 3) concha, 
fatty tissues and muscular tissues. 

The ear pieces were placed into glass tubes plugged by cotton wool and kept in dark- 
ness at room temperature and at constant relative humidity of: a) 75%, b) 85%, 
c) variable humidity by wetting of the cotton stopper twice a day. 

A similar technique has been utilized with success by Fain (1968, 1969b and 1969c) 
to obtain the development of the hypopi of several other species of Glycyphagidae. 

Hypopi and ear-wax scraped off were placed in glass tubes with: 

4) yeast; 5) evaporated milk; 6) oatmeal and yeast; 7) pollard and yeast; 8) skin 
scales, yeast, evaporated milk; 9) skin scales, yeast, oatmeal and pollard. 

The tubes were kept under the humidity conditions mentioned above. In all rearing 


173 


174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


ER 
N 
VE 


3) 
a 


È 


Fig. 1, 2 — Melesodectes auricularis Fain & Lukoschus, 1968. 1) female venter, 2) bursa copulatrix. 
al-a5 anal setae, ga genital anterior, g 72 genital median, gp genital posterior, s lateral setae, sh 
subhumeral 


LUKOSCHUS, DE COCK & FAIN: Life cycle of Melesodectes 175 


media and humidity conditions partial rise of tritonymphs was observed, but development 
to adults only in the tubes with ear parts, further development to larvae and protonymphs 
only in tubes with fatty tissues. 

At 85% relative humidity mould cultures were developing, but mites were not ob- 
served to feed on them. Best results were obtained with 3) concha + fatty tissues + 
muscular tissues at variable humidity. 

In this rearing experiment one of us (F.L.) noted as shortest developmental intervals: 
17.1V.1969 death of badger, shipment of ears; 20.IV. arrival at Nijmegen, tritonymphs 
present, starting of rearing experiments; 21.IV. first adults; 22.IV. first copulations; 
24.IV. first eggs; 26.IV. first larvae; 29.IV. first protonymphs; 4.V. all mites dead 
without forming of hypopi; large numbers of larvae developed to only few protonymphs. 

Tests of reproduction capacity by placing newly hatched adult couples into rearing 
mixtures 8 and 9 proved to be ineffective, only three of 15 couples laid up to four eggs 
CENT) 

Function of thickened legs II of male could be observed in copulating pairs. Tibiae 
IV of female are grasped by the clasper formed by apophysis on femur and tibia of male 
leg II. Frequently but not always, also tibia or tarsus of leg III of female is grasped by 
male tarsus II bent backwards. 

Transfer of larvae to rearing media 5—9 did not result in rise of protonymphs. 

On the suckling badger, the mother of which had parasitized ears, no hypopi or other 
developmental stages of this species were found in the ears nor between the hairs of the 
fur. On the ventral side of the abdomen and on the inner side of femora of hind legs 
small injuries of the epidermis were observed. 

In skinning with the aid of a dissecting microscope in these parts and ventrally on 
thorax 16 hypopi were found in the connective tissue, partly in the muscular tissue, 
partly in the tissues attached to the skin. 


SYSTEMATICAL POSITION OF THE GENUS Melesodectes AND OF 
THE SUBFAMILY MELESODECTINAE 


Until adults of the genus Melesodectes were investigated by us, we ranged it among 
the Glycyphagidae. From the discovery of both adult forms and larvae it now appears 
that this genus also shows some characteristics known from Saproglyphidae. 

Zachvatkin (1941) has divided Tyroglyphoidea (= actually Acaroidea) in three 
families, of which characteristic data are summarized below: 

1) Tyroglyphidae (= actually Acaridae): Tarsal claws sessile and connected with 
tarsus by two sclerites. Idiosoma with distinct sejugal furrow. Cuticle smooth, small and 
colourless. Hairs of hysterosoma neither pectinated nor leaf-shaped. Female without 
epigynium. Male with well developed adanal suckers and two suckers on tarsi IV. Genitai 
suckers well developed, digitiform. Organ of Claparède (’’Bruststiele’’) well developed 
in larva. 

2) Saproglyphidae: Tarsal claws small, ambulacrum situated on top of a long pretarsus 
and not connected with tarsus by two sclerites. Idiosoma with distinct sejugal furrow. 
Cuticle smooth or distinctly striated, membranous. Idiosomal hairs smooth and piliform. 
Female with or without epigynium. Male without adanal or tarsal suckers (except in the 
genus Pontoppidania Oudemans where these suckers are present). Genital suckers ge- 
nerally disciform. Hairs v e are lacking. Organ of Claparéde well developed. 


176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 3, 4. — Melesodectes auricularis Fain & Lukoschus, 1968. 3) male, venter, 4) supracoxal seta 


3) Glycyphagidae: Tarsal claws as in Saproglyphidae. Sejugal furrow usually lacking. 
Cuticle thick, coloured and often with patterns or verrucous. Idiosomal hairs variable, 
rarely piliform. Male without adanal or tarsal suckers. Epigynium generally present in 
female. Genital suckers usually present, but slightly developed (disciform, rarely 
digitiform). Hairs ve generally present. Organ of Claparède reduced. 

The genus Melesodectes shows claws similar to those in Saproglyphidae and Glycypha- 
gidae. The sejugal furrow is present, but is only slightly characterized and may even quasi 
disappear in swollen specimens. Cuticle slender, colourless without patterns nor verru- 
cous; however, on certain parts of the body of the less swollen specimens, some very 
narrow irregular ridges can be observed. Hairs of idiosoma are piliform, carrying short 
and only few barbules. The female shows a well shaped epigynium. The male is 


LUKOSCHUS, DE Cock & FAIN : Life cycle of Melesodectes 177 


Fig. 5. — Melesodectes auricularis Fain & Lukoschus, 1968, larva, venter 


devoid of adanal or tarsal suckers. The genital suckers are very short, almost disciform. 
In the larva the organ of Claparéde is well developed. Finally the tarsi appeared to 
be long and narrow, particularly in the female. 

Some characteristics, such as the presence of the sejugal furrow, the weak and smooth 
structure of the cuticle, the well developed organ of Claparéde, bring this genus close 
to the family Saproglyphidae. However, this genus does not fully fit into this 
family, because of the presence of the hairs ve, the slight development of the sejugal 
furrow, the distinct elongation of the tarsi, and the branched shape of the supracoxal 
hair. In view of these characteristics it may rather be ranged among the Glycyphagidae. 
Actually, it forms a different group, intermediate between these two families. 

With respect to the hypopus, it cannot be ranged among any of the three families of 
Acaroidea, because of its deviating character. 

Hence our supposition may be justified to maintain this genus in an independent 
subfamily (Melesodectinae) and to range it provisionally within the family Glycy- 
phagidae. 

Female: Idiosoma egg-shaped, length except gnathosoma average of 24 specimens 
measured 564 n (464—664), width 9 400 u (291—455). Cuticula smooth, without 
epidermal structures or coloured pattern, with some irregular striations or wrinkles. 

Venter (Fig. 1): Epimera I fused in Y-chape, epimera VI fused with epimerites 
III, forming closed coxal fields. Genital opening between epimera II and III; epigynium 


178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 6. — Melesodectes auricularis Fain & Lukoschus, 1968 protonymph venter 


sickle-shaped and well sclerotized. Two vulvar valves without genital apodemes. Disc- 
shaped short genital suckers beneath vulvar valves. Anal opening ventral subterminal 
(129 x long). Bursa copulatrix (Fig. 2) opens terminal in a more sclerotized chamber, 
protuberant in part of the females. Bursa 150 y long and relatively very wide, spermatheca 
soft and only somewhat larger than bursa. Two cup-like appendages near mouth of 
spermatheca. Ventral hairs setiform: ga (40 u), gm (40 u), gp (54 u), al (30 u), 
a2 (45 u),a3 (29 u), a4 (25 u), a5 (24 u). Dorsal setae translocated to venter, pecti- 
nated GA (SS fH) SCO): 


LukoscHus, DE Cock & FAIN : Life cycle of Melesodectes 179 


Gnathosoma: Pedipalpal coxae with two pairs of setae, lateral of them pectinate. 
Pedipalps with two segments, proximal segment with two setae, distal with one seta, 
one solenidion and button-like proximal protuberances. Chelicerae heavily dentated 
with little mandibular spine and four conical spurs. 

Legs with five free segments, all tarsi elongated. Single little claw on lengthened 
pretarsus. Chaetotaxy of legs: tarsi 12-12-10-10, tibiae 2-2-1-1, genua 2-2-1-0, femora 
1-1-0-1, trochantera 1-1-1-0, coxal fields 1-0-1-0. 

Solenidiotaxy: Tarsi 3-1-0-0, tibiae 1-1-1-1, genua 2-1-1-0, famulus present on tarsus 
I (Fig. 17,18). 

Dorsum (Fig. 8): Sejugal furrow indistinct and only in median part. All dorsal setae 
with exception of ve (25 u) pectinate. v7 (90 u) in front of smooth ve. Supracoxal 
setae (Fig. 11) branched and forked with small organ of Grandjean, sce (225 u) four 
timesmas long as scz2 (57. u), 21 (60) u), 42 (29 u), 43 (385 u), 24 (140 
u). a (255 m)» Bil (65 u), 2, (50 mE 15 (33 u), 14 (280 1) h (198 u). Pore 
of oil gland medially to lateral 3. 

In gravid females up to five eggs with well developed shells are observed. In freshly 
deposited eggs the shell has a honeycomb pattern, when observed under the dissecting 
microscope, however, after mounting in Hoyer's mixture no shell structures are 
visible. In far embryonated eggs two small chitineous cup-like structures are present. 

Male: Idiosoma length average of 20 specimens measured 406 u (360-450), width 9 
274 u (252—302). General shape unlike female, because of distinctly brown coloured 
legs. Legs II strongly thickened, forming copulatory clasping organs. 

Venter (Fig. 3): Epimera I long Y-shaped, epimera II and epimerites II forming 
almost closed coxal fields II, epimera III and IV fused. Genital opening behind legs 
IV, penis short. Two pairs of small disc-shaped genital suckers between very g p, g m 
33 4, ga 10 y. Anus ventro-terminal, with only three pairs of anal setae. Gnathosoma 
as in female. 

Legs thicker, tarsi shorter than in female, pretarsi longer, stalk much broader than 
in female (Fig. 17, 20). Legs II strongly thickened and specialized for clasping organs. 
Large two-pointed femoral apophysis and tibial spur-shaped pincers, supported by 
posterolaterally notched tarsus. 

Chaetotaxy: Tarsi 8-8-6-6, tibiae 2-2-1-1, genua 2-2-1-0, femora 1-1-0-1, tochantera 
1-1-1-0, coxal fields 1-0-1-0. 

Solenidiotaxy: Tarsi 3-1-0-0, tibiae 1-1-1-1, genua 2-1-1-0. Famulus present on tarsus 
I(Eig 195 20). 

Dorsum as in female. Setae insignificantly smaller than in female. 

Larva: Idiosoma egg-shaped, length average of 14 specimens 208 u (164—306), 
width @ 159 u (99—216). Cuticula in freshly emerged specimens mammilated and 
irregularly striated, in fullgrown larvae smooth like in adults. 

Venter (Fig. 5): Epimera I fused in V-shape, epimera II and III free. Organ of 
Claparède 28 u long with long well sclerotized and coloured basal part, soft uncoloured 
ring and coloured end. Anus ventral subterminal without anal setae. Gnathosoma as in 
adults. 

Legs almost like in female. Chaetotaxy of legs: Tarsi 12-12-10, tibiae 2-2-1, genua 
2-2-1, femora 1-1-0, trochantera 0-0-0, coxal fields 1-0-1. 

Solenidiotaxy: Tarsi 1-1-0, tibiae 1-1-1, genua 2-1-1. Famulus on tarsus I remarkably 
longer than in adults (Fig. 12, 13). 


180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


10 11 


\ \ 
\ 
N 


Fig. 7—11. — Melesodectes auricularis Fain & Lukoschus, 1968. (7) larva, dorsum, (8) female, 

dorsum, supracoxal setae of larva (9), protonymph, (10) and female (11). sc supracoxal setae, 

d 1-5 dorsal setae, h humeral seta, / 1-5 lateral setae, ve vertical external setae, v7 vertical internal 
setae 


Dorsum (Fig. 7): Idiosomal setae (lacking are 4 4 and / 4) with exception of ve 
and d 2 pectinate and with the same length relations as in adults. Supracoxal setae (Fig. 9) 
branched and forked, Grandjean organ palmate (more distinct than in adults). 

Protonymph: Length of idiosoma average of 14 specimens measured 357 u (302— 
473), width @ 253 u (198—338). 

Venter (Fig. 6): Epimerae I fused in V- or very short Y-shape, epimerae II-IV free. 
Anus ventral subterminal with three pairs of anal setae. One pair of disc-shaped genital 
suckers and genital median setae between legs IV. Gnathosoma as in adults. 

Chaetotaxy of legs: Tarsi 12-12-10-10, tibiae 2-2-1-0, genua 2-2-1-0, femora 1-1-0-0, 
trochantera 0-0-0-0, coxal fields 1-0-1-0. 


LukoscHus, DE Cock & Fain: Life cycle of Melesodectes 181 


Solenidiotaxy: Tarsi 2-1-0-0, tibiae 1-1-0-0, genua 2-1-1-0. Famulus present on tarsus 
I (Fig. 14, 15). 

Dorsum like tritonymph, only length of setae somewhat shorter, supracoxal setae 
(Fig. 10) less branched. 


DEPOSITION OF SPECIMENS 


Adults and developmental stages have been deposited in: 
Rijksmuseum van Natuurlijke Historie; 
British Museum (Natural History) ; 
Rocky Mountain Laboratory, Hamilton, Montana; 
Field Museum of Natural History, Chicago; 
Department of Zoology, University of Massachusetts, Amherst, Mass. ; 
Zoologisches Staatsinstitut und Zoologisches Museum, Hamburg; coll. nr. A22/71 
U.S. National Museum, Washington; 
Institute of Parasitology, Academy of Sciences, Praha; 
Zoological Institute, Academy of Sciences, Leningrad; 
Institut de Médecine Tropicale Prince Léopold, Antwerpen; 
Zoölogisch Laboratorium, Katholieke Universiteit, Nijmegen. 


DISCUSSION 


Morphological characteristics of hypopi, adults and larvae suggest relations to the 
family Hypoderidae (subcutaneous parasites of birds and rodents) and also to the family 
Saproglyphidae. 

Genera of the subfamily Hypodectinae, parasites of birds, have a remarkable biology. 
Developmental cycles have been observed in Hypodectes propus Nitzsch, 1861, subdermal 
parasites of pigeons, by Fain and Bafort (1966). In this species free small hypopi are 
found in nests of pigeons during breeding time and also under the skin in nestlings. 
Within the tissue of birds these hypopi extend for 7—10 times. They rest within the 
tissues until the full-grown pigeon sits on eggs. Then the large tissue hypopi are elimina- 
ted by the bird and give direct development to adults. From the numerous eggs hatch 
free hypopi. Larval and protonymphal stages remain rudimentary within the egg 
shell. Development is shortened by lack of free living larvae, protonymphs and 
tritionymphs and in time development from large tissue hypopi — adults — reproduction 
— free hypopi to a short breeding period. Most absorption of food seems to be within 
tissue hypopus. There are no observations on feeding of the adults. Free hypopi get 
through little resistant epidermis of nestlings (Fain, 1967). Development is suggested 
to be affected by hormonal conditions of host while breeding. 

Of the subfamily Muridectinae, subdermal parasites of rodents, neither adults nor 
developmental cycles have been observed (Fain, 1968, 1969). 

Although in our rearing experiments the whole cycle, including forming of hypopi 
ex protonymphs, could not be observed, results may be compared. Development from 
hypopi to protonymph occurs within a very short time (shortest observed time: 12 days, 
in laboratory conditions). As lactation period in Meles meles lasts 12—14 weeks, there 
will be sufficient time for infection of nestlings, even if the development may be distinctly 
longer under field conditions. 


182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 12—20. — Melesodectes auricularis Fain & Lukoschus, 1968, leg I of larva dorsally (12). 
ventrally (13), of protonymph (14, 15), of hypope (16), female (17, 18) and male (19, 20) 


Contrary to Hypodectinae, feeding of free living stages is necessary. There has been 
no development nor egg production without suitable food. Low numbers of developed 
protonymphs may be caused by shortage of food or decay. 

It is interesting to mention that Fain (1969c, p. 758), working with hypopi of 
Labidophorus talpae and Orycteroxenus dispar from the mole (Talpa europaea), succeed- 
ed in obtaining the development of hypopi into tritonymphs, but was unable to rear 


LukoscHus, DE Cock & FAIN: Life cycle of Melesodectes 183 


adults from the latter. It seems that here also the nymphs were unable to develop into 
adult stage in the absence of suitable food. In two other experiments this author (1968b 
and 1969b), working in Central Africa with the same technique, was able to abtain the 
development of endofollicular hypopi of the genera Lophuromyopus and Rodentopus 
into tritonymphs and adults. Free moving adults were obtained after a delay of 6 days. 
No food was provided during the experiment. 

These experiments show that the development into the adult stage is variable according 
to the species involved. Some may require food, while other do not. 

Failure of rearing mites on defined rearing media, fitted for most species of nest- 
inhabitant mites, suggests that Me/esodectes is not a free-living nest-inhabitant, hypopi of 
which are phoretic on hosts for reaching new habitats, like Xenoryctes krameri, but a 
specialized monophageous parasite. 

Successful rearing only on epidermis and fat tissue of the host suggests that in field 
conditions free living stages also feed on debris of host tissues present within the nests 
only during lactation period. 

Observation of injuries and penetrated hypopi subcutaneous in a nestling shows close 
relations to Hypoderinae. The strong knife-like claws of hypopi (also typical in many 
Hypodectinae genera) may enable the penetration of the epidermis of nestling. 

Presence of chitineous cup-like structures within the embryonated egg shows striking 
similarities to those in some families of host specific cutaneous mites like Myialges 
anchora (Oudemans, 1935), Myialges macdonaldi (Evans, Fain & Bafort, 1963) and 
Gliricoptes betulinus (Kok, Lukoschus & Fain, in press). 


REFERENCES 


Evans, G. O., A. Fain & J. Bafort, 1963. Découverte du cycle évolutif du genre Myialges avec 
description d'une espèce nouvelle (Myialgidae : Sarcoptiformes). — Bull. Ann. Soc. Roy. 
Ent. Belgique, 99 (34): 486—500. 

Fain, A., 1965. A review of the family Epidermoptidae Trouessart parasitic on the skin of birds 
(Acarina : Sarcoptiformes). — Verh. Kon. Vlaamse Acad. Wetensch., 27, nr. 84 (I-II): 
1—176, 1—144 (185 figs.). 

——, 1967. Les hypopes parasites des tissus cellulaires des oiseaux (Hypodectidae : Sarcopti- 
formes). — Bull. Inst. Roy. Sci. nat. Belgique, 43 (4): 1—139. 

—, 1968a. Un hypope de la famille Hypoderidae Murray, 1877, vivant sous la peau d'un 
rongeur (Hypoderidae : Sarcoptiformes). — Acarologia, 10 (1): 111—115. 

—., 1968b. Acariens nidicoles et détriticoles en Afrique au Sud du Sahara. IV. Découverte du 
cycle évolutif du genre Lophuromyopus Fain, 1965. — Rev. Zool. Bot. Afr. 78: 161—174. 

—, 1969a. Les deutonymphes hypopiales vivant en association phorétique sur les mammifères 
(Acarina : Sarcoptiformes). — Bull. Inst. Roy. Sci. nat. Belgique, 45 (33): 1—262. 

——., 1969b. Acariens nidicoles et détriticoles en Afrique au Sud du Sahara. V. Description du 
cycle évolutif de deux espéces du genre Rodentopus Fain. — Acarologia, 11: 304—316. 

——., 1969c. Morphologie et cycle évolutif des Glycyphagidae commensaux de la Taupe, Talpa 
europaea. — Acarologia, 11: 750—795. 

Fain, A. & J. Bafort, 1966. Les hypopes parasitant les tissus cellulaires des pigeons sont les deuto- 


nymphes d’un acarien libre et pas celles d’un acarien plumicole. — Rev. Zool. Bot. Afr., 74: 
313—316. 

Fain, A. & F. Lukoschus, 1968. Une nouvelle deutonymphe hétéromorphe (hypope) parasite du 
Blaireau (Meles meles) en Hollande (Acarina: Sarcoptiformes). — Rev. Zool. Bot. Afr., 
78: 175—182. 

Kok, N. J. J., F. S. Lukoschus & A. Fain, 1971. Four new species of the genus Gliricoptes Law- 
rence, 1956 from European rodents. — Acta Zool. Pathol. Antwerpiensia (in press). 


Oudemans, C. A., 1935. Description du Myialges anchora Sergent & Trouessart, 1907 (Acarien). 
— Ann. Parasitol., 13 (1): 5—11. 


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FOUR NEW SPECIES OF THE GENUS PSORERGATES TYRELL 
FROM EUROPEAN HOSTS (ACARINA, PSORERGATIDAE) 
by 
F. S. LUKOSCHUS, A. W. A. M. DE COCK & F. M. DRIESSEN!) 


Zoological Institute, Catholic University of Nijmegen 


ABSTRACT 


Four new species of the genus Psorergates, itch mites of small mammals, are described. Systema- 
tic positions are discussed. The new species Psorergatus muscardinus, baueri, neerlandicus and 
quercinus are parasites of Muscardinus avellanarius (Germany), Neomys fodiens (Austria), Microtus 
oeconomus (the Netherlands) and Eliomys quercinus (Spain), respectively. 


INTRODUCTION 


This study is a continuation of a series of investigations on Psorergatidae, itch mites 
of rodents, carnivores and insectivores from Europe (Fain, Lukoschus & Hallmann, 
1966; Lukoschus, Fain & Beaujean, 1967; Lukoschus, 1967, 1968a, b, 1969; Fain & 
Lukoschus, 1948; de Cock, Lukoschus & Ariani, 1971) and Canada (Kok, Lukoschus 
& Clulow, 1970, 1971). 

When systematically investigating European hosts, one of us (F.L.) succeeded in 
discovering new species of the genus Psorergates which are described below. 

The species have been collected from alcohol preserved hosts in Naturhistorisches 
Museum, Wien (Dr. K. Bauer), Centro Pirenaico de Biologia Experimental in Jaca, 
Spain (Prof. Dr. E. Balcells R.), from dor-mice kindly sent to us for observations by 
Dr. O. Henze, Institut für angewandte Zoologie, München, and from freshly trapped 
voles. We highly appreciate the kind co-operation of our colleagues. 


Psorergates muscardinus spec. nov. 
(Figs. 1—13) 


Female (holotype). — Shape and body as in other species of the genus 
Psorergates. Length including gnathosoma 178 u, average in 18 paratypes measured 
168 u (145—186), width 145 u, average in the paratypes 139 u (124—152). 

Venter (Fig. 1). Cuticle soft. Ventral setae (vs) 10 u (8—12), distance between 
ventral setae 12 y (7—14). Genital opening (vu) 18 u (15—19) long, lying between 
two adanal lobes, each of which carries a pair of terminal setae of about 81 mu (69—88). 
Epimerae I bent abroad. Epimerae II—IV beneath body surface. 

Legs inserted ventro-laterally. Articulation between epimera and trochanters strongly 
sclerotized. All trochanters with a long and strongly sclerotized ventral spur that is 
pointing to the anterior part of the body. Femora of all legs with a large prominent 
ventral spur and a pair of postero-lateral setae. Length of these setae 23—27 u on femora 


1) Present address: Nederlands Instituut voor Zuivel Onderzoek, Ede. 
185 


186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, 


N 


i 


Fig. 1—6. Psorergates muscardinus spec. nov. female. 1, hol 
tibia, ventral; 3, leg I, tarsus and tibia, dorsal; 4, holotype, d 
6, gnathosomal seta 


AFL. 4, 1971 


otype, venter; 2, leg I, tarsus and 
orsum; 5, chelicera in lateral view; 


F. S. LukoscHus, A. W. A. M. DE Cock & F. M. DRIESSEN: Four species of Psorergates 187 


I—IV. Apical seta of each pair almost twice as long as basal one. Genua with postero- 
lateral seta, 5 u on genua I—III and 17 u (17—29) on genu IV. Tibiae with short club- 
like spines antero-ventrally (not present on leg IV) and a longer dorso-medial seta. Tarsi 
with long dorso-posterior seta (dp), shorter dorso-anterior seta (da) and latero-ventral 
club-like spine (Fig. 2). Postero-dorsal seta not present on tarsi IV. Tarsi I and II 
dorsally with two solenidia (Fig. 3), dorso-medial solenidion bulbous, antero-lateral one 
lying inside a duplication of the epidermis. Two single-pointed claws and a bilobed 
empodium inserted ventrally at the end of the tarsi. 

Dorsum (Fig. 4). Dorsal shield well sclerotized and distinctly punctured. Length of 
shield 99 u, average in paratypes 101 u (96—106), width 78 u, average in paratypes 
80 u (74-83). Three pairs of lateral setae present on the shield, second pair about 8 y 
(8—9) long. Antero-medially of the first pair of lateral shield setae a pair of minute 
setae present, 11 u (10—12), at some distance of the margin of the shield. Soft parts 
of the dorsal side striated. 

Gnathosoma and mouth parts like in other Psorergates species. Gnathosomal setae 
(Fig. 6) 5,5 u (5—7) long, bilobed with slightly incised borders. Subgnathosomal 
setae (sg) short, pharyngeal bulb (pb) oval with a gland duct to mouth opening. On 
the palpal tibia two dorsal setae present, the anterior one very small, length of posterior 
seta 12 u (12—14). Further present on the palpal tibia a small soft lateral seta and a 
spur on the dorsal end. Palpal tarsus with a bi-pointed and a three-pointed claw. 
Chelicerae with stinging bristles and dentated digitus fixus. The digitus fixus from a 
squashed paratype in lateral view is given in Fig. 5. 

Male (allotype). — Shape and body like in other Psorergates species. Length including 
gnathosoma 154 u, average in 15 paratypes measured 144 „ (131—154), width 138 u, 
average in the paratypes 125 u (115—138). 

Venter (Fig. 8). Similar to that in female, but only with two terminal setae of 34 y 
(32—38) on a sclerotized medial tubercle. Legs as in female, but all setae somewhat 
shorter: ventral setae 8 u (6—10), distance between ventral setae 14 u (13—20), 
trochanteral setae 8 u (7—9), femoral setae I—III 15 x (14—16), femoral setae IV 
20 u (17—22), genual setae I—III 4 y, genual setae IV 16 u (13—18). 

Dorsum (Fig. 7). Genital opening near medio-anterior border of shield. Two pairs 
of minute genital setae beside the genital opening, posterior setae 8 u (8—9) apart, 
anterior setae 5 m (3—5) apart. Length of the very weakly sclerotized dorsal shield 
99 y average in the paratypes 89 u (79—99), width 74 u, average in the paratypes 
68 u (63—74). Single-pointed penis of 29 u (29—34), penis envelope 22 u (18—24), 
on dorsal side shorter. Striation pattern of the soft dorsal parts different from female. 

Gnathosoma and mouth parts built as in female. Posterior palpal tibial setae 5 u 
(3—5) and gnathosomal setae 3 u (3—5). 

Immature stages. — Egg with thin shell, almost globular. Dimensions of five 
measured eggs: 92 X 87 u, 90 X 85 u, 120 X 108 u, 110 X 108 u, 103 X 101 u. 

Larva (Fig. 9). Disc-shaped with three pairs of two-segmented legs. Average length 
in 11 specimens measured 106 u (97—120), average width 87 u (78—99). Cuticle 
soft, with indistinct dorsal striation. Trochanters with well developed spurs, small 
indistinct epimerae, segments, femur to tarsus fused, forming a flattened unit with two 
three-pointed claws. Gnathosoma (Fig. 10) almost as large as in adults, but setae on 
gnathosoma (Fig. 11) somewhat smaller, subgnathosomal setae longer than in adults. 

Protonymph (Fig. 12). Like larva, but four pairs of two-segmented legs. Fused 


188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 7—8. Psorergates muscardinus spec. nov. male. 


7, allotype, dorsum; 8, allotype, venter 


189 


F. S. LukoscHus, A. W. A. M. DE Cock & F. M. DRIESSEN: Four species of Psorergates 


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‘IL ‘fessop ‘eAre] ewosoygeug ‘QT ‘IoJUoA FATE] “6 ‘sase}s jezuowdojoaop ‘Aou ‘ads smurpwssuwm saySsasosq ‘CI—G “BIg 


190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


segment with an apophysis that may be lacking on leg IV. Solenidia (so) present on 
legs I and II. Setae on gnathosoma longer than in larva, exept subgnathosomal setae. 
Average length in 6 specimens measured 127 u (115—143), average width 108 u 
(97—122). Posterior palpal tibial setae longer than in larva. 

Deutonymph (Fig. 13). Like protonymph, but larger. Spur on the fused segment 
bigger than in protonymph and never lacking on leg IV. Two solenidia present on leg I, 
one solenidion on leg II. Gnathosomal setae longer than in protonymph, as well as 
posterior palpal tibial setae (4—5 u), but shorter than in adults. Average length in 
10 specimens measured 161 u (149—184), width 138 u (124—156). 


Type host. — Myscardinus avellanarius Linnaeus. 

Type locality. — München, Germany, 18.V.1969 and 8.VII.1969. 

Pathology. — The mites live within the epidermis of the ear concha, causing hyper- 
ceratosis. 


Deposition of types. — Holotype and allotype: Zoologisches Institut und Zoologi- 
sches Museum, Hamburg. 

Paratypes 9 and g': U.S. National Museum, Washington; British Museum (Natural 
History), London, coll.nr. 1971/158-9; Rijksmuseum van Natuurlijke Historie, Leiden, 
coll.nr. P 1213-4; Naturhistorisches Museum, Wien; Muséum National d'Histoire 
Naturelle, Paris, coll.nr. 11.1. 13-14; Acarology Laboratory, Columbus, Ohio; Field 
Museum of Natural History, Chicago; Institut de Médecine Tropicale Prince Léopold, 
Antwerpen; Department of Zoology, Catholic University, Nijmegen. 


Psorergates baueri!) spec. nov. 
(Figs. 14—19) 


Female (holotype). — Shape and body build as in other species of the genus Psorer- 
gates from insectivora (Lukoschus 1968a,b). Length including gnathosoma 117 u, 
average in 11 paratypes measured 114 u (99—129), width 92 u, average in the para- 
types 1/4 (781010); 

Venter (Fig. 14). Cuticula soft. In the middle of the venter two setae of about 4 u 
(4—6), and 6 w (5—10) apart. Epimera II—IV and anterior part of epimera I 
lying beneath the body surface. Posterior part of epimera I bent outwards and circularly 
closed. Genital opening 10 u (8—11) long, lying between two adanal lobes, each of 
which carries a pair of terminal setae of 54 u (48—58). 

Legs inserted ventro-laterally. Articulation between epimera and trochanters strongly 
chitinized. All trochanters with a strongly sclerotized spur that is pointing to the 
anterior part of the body, and a small spur directed towards the large femoral spur. 
Distally to the strong spur a trochanteral seta of 5 u (5—8). Femora of all legs with a 
large ventral spur. Femora I—III with a pair of postero-lateral setae of 24 u (20—27), 
of which the basal is always shorter than the apical. Femur IV with only one seta: 24 u 
(20—27). Postero-lateral setae on genua I—III 1 u, on genu IV 36 u (33—45). Tibia 
with short antero-ventral spine (lacking on leg IV) and a longer dorso-medial seta. Tarsi 
with postero-dorsal (dp) seta of 16 u (14—18), antero-dorsal (da) seta of 2—3 y and 
a ventral spine. Postero-dorsal seta not present on leg IV. Two solenidia on the dorsal 


1) The species is named in honour of Dr. K. Bauer, Naturhistorisches Museum, Wien (Austria). 


F. S. LukoscHhus, A. W. A. M. DE Cock & F. M. DRIESSEN: Four species of Psorergates 191 


Fig. 14—19. Psorergatus baueri spec. nov., female. 14, holotype, venter; 15, holotype, dorsum; 
16, leg I, femur-tarsus, dorsal; 17, chelicera in lateral view; 18, palpal claw in lateral view; 
19, gnathosomal seta 


192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


side of tarsi I and II (Fig. 16), dorso-medial solenidion bulbous. antero-lateral within a 
duplication of the epidermis. Two two-pointed claws and a bi-lobed empodium inserted 
ventrally at the end of the tarsi. 

Dorsum (Fig. 15). Dorsal shield well sclerotized and distinctly punctured. Soft parts 
of dorsum striated. Length of shield 77 u, average in the paratypes 77 u (67—81), 
width 58 u, average in paratypes 62 u (53—68). Three pairs of lateral setae, distinctly 
distant from the margin of the shield. Second pair 4 u (3—5) long. Antero-medially of 
first pair of lateral setae a pair of minute setae present. 

Gnathosoma and mouth parts like in other Psorergates species. Gnathosomal setae 
bilobed, 7 u (5—8), short lobe with relatively deeply incised border (Fig. 19). Posterior 
dorsal seta on palpal tibia very short (2 4). Further present on palpal tibia: a very small 
anterior seta, a lateral seta, and a spur on the dorsal end. Palpal tarsus with two claws 
(Fig. 18) and one spine. Chelicerae with stinging bristles and dorsally dentated digitus 
fis (Big): 

Male unknown. 

Immature stages. — Egg: thin-shelled, 69 X 58 u, 78 X 69 u, 71 X 60 y. 

Larva: Like in other Psorergates species, 76 X 62 u. 

Protonymph: Not found. 

Deutonymph: Like in other Psorergates species, 120 X 99 u, without spurs on fused 
leg segments. 

Type host. — Neomys fodiens (Pennant). 

Type locality. — St. Michael, Salzburg, Austria, 9.VII.1967. 

Pathology. — The mites were found between the tendons of the forelegs within the 
upper layers of the epidermis, causing hyperceratosis. 

Deposition of types. — Holotype ® at Wien; paratypes at Leiden P 1218-9, Ham- 
burg, Washington, Antwerpen, Nijmegen, London (1971/166), Paris (11 J 15). 


Psorergates neerlandicus spec. nov. 
(Figs. 20—25) 


Female (holotype). — Shape and body build as in other Psorergates species from the 
apodemi-group (Lukoschus, Fain & Beaujean, 1967). Length, including gnathosoma, 
132 u, the same as in two paratypes, width behind second pair of legs 115 u, like in 
paratypes. 

Venter (Fig. 20). Venter with soft cuticula. In the middle two ventral setae of 7 u 
long, 9 y apart. Genital opening 10 y long, lying beween trochanteral spurs IV, flanked 
by two adanal lobes, each of which carries a pair of terminal setae of about 79 u long. 
Chitineous ducts from genital opening indistinct. Anus not present. Posterior part of 
epimera I half circular, bent outwards. This crescent-shaped part standing out from 
the venter. Anterior part of epimera slightly, articulation with trochanters strongly 
chitinized. Epimera II—IV shaped as an oblong obtuse-angled triangle beneath surface 
of venter. Inwards-pointing parts slightly, articulation with trochanters, strongly 
sclerotized. 

Legs inserted ventro-laterally. Trochanters with strong, extending ventral spur, pointing 
to the anterior part of the body, and a seta (9 u) distally of this spur. Basal part of 
trochanters strongly chitinized with a small spur towards the femoral spur. Femora of 
all legs with a pair of postero-lateral setae and a large ventral spur. Basal seta of pairs 


F. S. LugoscHus, A. W. A. M. DE Cock & F. M. DRIESSEN: Four species of Psorergates 193 


Fig. 20—22. Psorergates neerlandicus spec. nov. female. 20, holotype, venter; 21, leg I, tarsus, 
dorsal; 22, holotype, dorsum 


194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


always little shorter than apical. Apical setae 19 y long on femora I—III, 24 y on 
femur IV. A seta present upon a small sclerotized elevation on the genua, with the 
length of 16 u on genu IV and of 5 y on genua I—III. Short tibia with an antero- 
ventral spine (lacking on tibia IV) and a longer dorso-medial seta. Tarsi with antero- 
ventral spine, postero-dorsal seta and antero-dorsal seta of about the same length. 
Postero-dorsal setae not present on tarsus IV. Tarsi I and II with a big, club-shaped 
solenidion and a smaller solenidion within a duplication of the epidermis (Fig. 21). 


Fig. 23—25. Psorergates neerlandicus spec. nov. male. 23, allotype, venter; 24, allotype, dorsum; 
25 penis, enlarged 


F. S. Lukoscuus, A. W. A. M. DE Cock & F. M. DRIESSEN: Four species of Psorergates 195 


Both strong single-pointed claws and bilobed empodium inserted ventrally at the end 
of the tarsus. 

Dorsum (Fig. 22). Dorsal shield well sclerotized and punctured up to 2 u from 
the shield border. Distinctly marked off from the softly chitinized, striated dorsal side. 
Length of shield 77 u, width 77 u. Three pairs of lateral setae of 5 u, standing at the 
border of the shield. Median setae very small, nearly point-shaped, situated 8 4 from 
shield margin. 

Gnathosoma and mouth parts like in other Psorergates species. Gnathosomal setae 
4 u long, two-segmented, with unindented border. Middle part of gnathosoma epistoma- 
like, covering mouth opening and posterior part of the chelicerae. Palpal tibia with two 
dorsal setae: posterior one 12 u long and well-developed up to the end (seems to be 
broken), anterior seta very short (1 u). On lateral side of the palp a soft seta. Dorsal 
end of palpal tibia with strongly chitinized spur. Palpal tarsus inserted ventro-medially. 
Chelicerae consisting of dentated digitus fixus with saw-function and stinging bristles. 

Male (allotype). — Shape and body build like in other Psorergates species from 
apodemi-group. Length including gnathosoma 110 u, average in 4 paratypes 109 u 
(104—119), width 94 y, average in the paratypes 94 u (82—101). 

Venter (Fig. 23). Terminal setae 62—67 u long, standing on a small sclerotized 
median tubercle. Setae on ventral side barely shorter than in female: ventral setae 5—6 
u, distance between ventral setae 7—10 y, trochanteral setae 7—9 y, femoral setae 
I-II 16—20 u, femoral setae IV 19—20 u, genual setae I—III 2 u and genual setae 
IV 12—14 u. 

Dorsum (Fig. 24). Similar to female, but antero-medially on dorsal shield an oval 
genital opening with two pairs of minute setae. Anterior pair of genital setae 18—19 u 
apart, posterior pair 7 „ apart. Average length of shield 73 u (70—78), width 70 y 
(67—73). Lateral shield setae 4 u long. Average length of penis 26 y (24—30 u), 
average length of penis envelope 22 u (20—23) (Fig. 25). Penis envelope shortened 
on dorsal side. 

Gnathosoma as in female. Posterior tibial setae of palp somewhat shorter: average 
10. m (GS), 

Type host. — Microtus oeconomus (Pallas). 

Type locality. — Texel Island, the Netherlands, 26.1X.1968. 

Pathology. — The mites were found within the epidermis on the venter of the abdo- 
men, causing hyperceratosis and papillomitosis. 

Deposition of types. — Holotype and allotype at Leiden P 1220-1. Paratypes: Ant- 
werpen, Washington, Nijmegen. 


Psorergates quercinus Spec. nov. 
(Figs. 26—33) 


Female (holotype). — Shape and body build as in other species of the genus 
Psorergates. Length including gnathosoma 175 u, averages in 10 paratypes measured 
169 u (157—179), width 125 u, average in paratypes 143 u (118—151). 

Venter (Fig. 26). Cuticle soft. Ventral setae 11 u long, 16 u (14—18) apart. An- 
terior part of epimerae I and epimerae II—IV beneath body surface. Posterior part of 
epimerae I standing out from venter. Genital opening more chitinized than in other 


196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 26—30. Psorergates quercinus spec. nov. female. 26, holotype, venter; 27, holotype, dorsum; 
28, larva, leg I, ventral; 29 protonymph, leg I, ventral; 30, deutonymph, leg I, ventral 


F. S. LukoscHus, A. W. A. M. DE Cock & F. M. DRIESSEN: Four species of Psorergates 197 


species, 15 u long, lying in front of two adanal lobes, each of which carries a pair of 
terminal setae of about 80 u (72—84). 

Legs inserted ventro-laterally. Trochanteral spurs peaked. Base of trochanters strongly 
chitinized. Trochanteral setae 12—16 u. Femora with a large prominent spur and two 
setae. Apical one of these setae slightly longer than basal: 18—22 y long on femur 
I-II, 22—24 y on femur IV. Genua with a seta on a small elevation: 5 u on genua 
I—III, 16—18 , on genu IV. Tibiae with a thin antero-ventral spine (lacking on tibia 
IV) and a longer dorso-medial seta. Further present on tarsi an antero-ventral spine, a 
postero-dorsal seta (10 „) and an antero-dorsal seta (11 u). Postero-dorsal seta lacking 
on tarsus IV. Bilobed empodium and two single-pointed claws inserted ventrally at end 
of tarsus. Tarsi I and II with a big solenidion and a minute one within a duplication 
of the epidermis. 

Dorsum (Fig. 27). Dorsal shield well sclerotized and distinctly punctured almost up 
to the margin. Anterior border slightly concave. Average length 112 u (109—118), 
average width 85 u (76—87). Shield setae standing on small unpunctured elevations, 
distinctly at some distance from shield border. Lateral setae 7—8 u long. Minute median 
setae 16—18 u from shield margin. 

Gnathosoma and mouth parts as in other Psorergates species. Gnathosomal seta 5 u 
long, at the end manifold incised. Palpal tibia dorsally with two setae of which the 
posterior one is 17—19 n long and well-developed up to the end (seems to be broken). 
Anterior seta on palpal tibia very short; 2 u. On lateral sides of the palpi a soft seta. 
Palpal tarsus inserted ventro-medially. Chelicerae with dentated digitus fixus and 
stinging bristles. 

Male (allotype). — Shape and body build as in other Psorergates species. Length 
including gnathosoma 150 u, average in 8 paratypes 142 u (129—154), width 130 u, 
in paratypes 123 u (109—134). 

Venter (Fig. 31), similar to female, but only two terminal setae (46—49 u) on a 
long sclerotized median tubercle. Legs as in female, but all setae shorter; ventral setae 
7 u, distance between ventral setae 16—19 y, trochanteral setae 6—9 y, femoral setae 
III 12—14 u, femoral setae IV 13—16 u, genual setae I—III 2—4 y and genual 
setae IV 8—13 u. 

Dorsum (Fig. 32). Dorsal shields well sclerotized and distinctly punctured. Behind 
the first pair of lateral setae shield bent inwards in a typical way, similar in all para- 
types. Second lateral shield seta 5 x long. Soft parts of dorsum striated. Two pairs of 
minute genital setae on small unpigmented tubercles, anterior genital setae 24—25 à 
apart, posterior setae 19—20 u apart. Length of single-pointed penis 46—52 y, penis 
envelope 28—31 u. Penis envelope deeply incised dorsally. 

Gnathosoma as in female, palpal tibial setae (6—8 y) and gnathosomal setae (4 uu) 
shorter. 

Immature stages. — Egg. Almost globular, thin-shelled. Dimensions of 6 eggs 
measured: 91 X 85 u, 94 X 85 1, 91 X 85 u, 85 X 79 u 108 X 105 u, 105 X 105 u. 

Larva. Disc-shaped with three pairs of two-segmented legs. Dimensions of three 
specimens measured: 96 X 77 u, 91 X 88 u, 116 X 99 y. Cuticle soft with indistinct 
dorsal striation. Trochanters with well-developed spurs, small, indistinct epimera, 
segments femur to tarsus fused, forming a flattened unit with two three-pointed claws. 
On this unit further present: on leg I (Fig. 28) one solenidion and a small spur, on leg 


198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


Fig. 31—33. Psorergatus quercin 


us spec. nov., male. 31, allotype, venter; 32, allotype, dorsum; 
33, penis, enlarged 


F. S. LukoscHus, A. W. A. M. DE Cock & F. M. DRIESSEN: Four species of Psorergates 199 


II one solenidion and an indistinct spur. Leg HI without spur or solenidion. Gnathosoma 
almost as large as in adults, but setae on gnathosoma somewhat shorter. 

Protonymph. Like larva, but four pairs of two-segmented legs. Spur on fused seg- 
ment better developed and present on all legs. Solenidia present on leg I and Il. 
Setae on gnathosoma longer than in larva, subgnathosomal setae somewhat shorter. 
Posterior palpal tibial setae 4 u long. Average length in 4 specimens measured 131 u 
(119—142), average width 117 u (108—128). 

Deutonymph. Like protonymph but larger. Spur on fused segment stronger than in 
protonymph. Posterior palpal tibial setae longer than in protonymph (6,5 u). Average 
length in 5 specimens measured 173 u (148—187), average width 148 u (133—153). 


Type host. — Eliomys quercinus (Linnaeus). 

Type- locality. — Madrid, Spain, 16.VII.1967. 

Pathology. — The mites live within the epidermis of the concha of the ears, causing 
hyperceratosis. 


Deposition of types. — Holotype and allotype: Departamento de Zoologia del Suelo y 
Entomologia aplicada, Madrid. Paratypes: Hamburg, Washington, Leiden (P 1215-7), 
London (1971/156-7), Wien, Paris (11 J 11-12), Ohio, Chicago, Antwerpen, Nij- 
megen. 


COMPARISON WITH RELATED SPECIES 


On account of their characteristics one cannot incorporate Psorergates muscardinus 
and Psorergates quercinus within the apodemi-, muricola- ot dissimilis-group. They 
show closer relationship to Psorergates eliomydis. Comparison of these three species is 
only possible for females. P. muscardinus is distinct from P. eliomydis by its bigger size, 
shape of the genital opening and bilobed gnathosomal setae. The epimerae I are less 
bent and narrower at the anterior end in P. muscardinus. The postero-dorsal seta (dp) 
on the tarsi is only slightly longer than the antero-dorsal seta (da) in P. eliomydis, 
but more than twice as long in P. muscardinus. Further differences are to be found in 
the length of ventral setae, genual setae, lateral shield setae, posterior palpal tibial setae 
and gnathosomal setae. 

P. quercinus differs mainly from P. eliomydis by its bigger size and bilobed gnatho- 
somal setae. The anterior end of the epimera I is narrow and lying beneath the body 
surface, as contrasted with P. eliomydis. There also are differences in length of the 
ventral setae, femoral setae I—III and gnathosomal setae. 

Females of P. muscardinus and P. quercinus have the same size, but differ in shape 
of the gnathosomal setae, genital opening and anterior part of the epimera I. The 
dorsal shield of P. gwercinus is more oblong than in P. muscardinus. The median setae 
are farther removed from the shield margin in P. quercinus and further differences are 
present in the length of the femur setae I—III, posterior palpal tibial setae, and the 
distance between the ventral setae. The males are more different. The typical shape of 
the dorsal shield of P. gwercinus males is an important distinctive character. Also 
remarkable is the difference in length of penis and penis envelope. The genital setae 
are much wider apart in P. gwercinus than in P. muscardinus. Further differences are 
present in the length of the terminal setae, setae of genu IV, shield setae and posterior 
palpal tibial setae. 

Psorergates neerlandicus shows the diagnostic characters of the apodemi-group; the 


200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 4, 1971 


palpal tibial setae are thick (and seem to be broken at the end). In the males the 
anterior genital setae are more than twice as far apart as the posterior ones. There are 
very few differences between this species and P. microti. Females only differ slightly in 
size and length of some setae. P. neerlandicus females have longer terminal, ventral, 
femoral and gnathosomal setae. Males of P. neerlandicus have obviously longer terminal 
and shorter genual setae I—III than males of P. microti. Further there are minor dif- 
ferences present in the lengths of femoral setae I—III and penis envelope. 

P. baueri from the host Neomys fodiens, belongs to the ‘‘insectivora’’-group, because 
the dorso-anterior setae of the tarsi are short, the lateral shield setae are distinctly remote 
from the shield margin and the ends of the gnathosomal setae are incised several 
times. The relationship to P. sorici and P. cinereus appears from the bi-pointed claws and 
the presence of only one seta on femur IV. 

P. cinereus is much bigger than P. baueri and there are obvious differences in the 
times. The relationship to P. sorici and P. cinereus appears from the bi-pointed claws and 
gnathosomal setae. There is less difference between P. sorici and P. baueri: P. baueri has 
shorter femoral setae IV, trochanteral setae and gnathosomal setae, but the ventral setae 
are slightly longer. 


REFERENCES 


Cock, A. W. A. M. de, F. S. Lukoschus & A. P. Ariani, 1971. Psorergates etruscus spec. nov. 
(Acari: Psorergatidae), a new itch mite from Suncus etruscus (Soricidae). Ann. Ist. Mus. 
Zool. Napoli (in press). 

Fain, A., F. Lukoschus & P. Hallmann, 1966. Le genre Psorergates chez les muridés. Description 
de trois espéces nouvelles (Psorergatidae: Trombidiformes). Acarologia 8: 251—274. 

Fain, A. & F. Lukoschus, 1968. Psorergates (Psorobia) foinae sp. n. acarien producteur de gale 
chez la fouine en Belgique. Bull Inst. r. Sci. nat. Belg. 44: 1—6. 

Kok, N. J. J., F. S. Lukoschus & F. V. Clulow, 1970. Psorobia castoris spec. nov. (Acarina: 
Psorergatidae), a new itch mite from the beaver, Castor canadensis. Can. J. Zool. 48: 
1419—1423. 

Kok, N. J. J., F. S. Lukoschus & F. V. Clulow, 1971. Three new itch mites from Canadian small 
mammals (Acarina: Psorergatidae). Can. J. Zool. (in press). 

Lukoschus, F., A. Fain & M. M. J. Beaujean, 1967. Beschreibung neuer Psorergates-Arten 
(Psorergatidae: Trombidiformes). Tijdschr. Ent. 110: 133—181. 

Lukoschus, F. S., 1967. Krätzmilben an spanischen Kleinsäugern (Psorergatidae: Trombidiformes). 
Rev. Iber. Parasitol. 27: 203—228. 

Lukoschus, F. S., 1968a. Neue Krätzmilben von einheimischen Insektivoren (Psorergatidae: 
Trombidiformes). Tijdschr. Ent. 111: 75—88. 

Lukoschus, F. S., 1968b. Psorergates desmanae sp. nov., eine neue Krätzmilbe von Galemys 
pyrenaicus (Psorergatidae: Trombidiformes). Bull. Mus. nat. Hist. nat. 2, sér. 40: 
125—131. 

Lukoschus, F. S., 1969. Psorergates (Psorobia) mustelae spec. nov. Eine neue Krätzmilbe von 
Mustela nivalis L. (Acarina: Psorergatidae). Zool. Anz. 183: 110—118. 


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No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.FI. 45.—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.FI. 70.—. 


No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Ditsh & Uvarov 
and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. 
D.Fl. 25.—. 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. 
D.Fl. 75.—. 


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THE GENUS BUATHRA CAMERON IN EUROPE ') 
(HYMENOPTERA, ICHNEUMONIDAE) 
by 
G. VAN ROSSEM 


Plant Protection Service, Wageningen 


ABSTRACT 


Three European species of Buathra Cameron, 1903, can be distinguished. Designations of 
lectotypes are presented for Ichneumon laborator Thunberg, 1822, Cryptus curvicauda Thomson, 
1896, and Cryptus divisorius Tschek, 1872. Cryptus medius Szépligeti, 1916, does not belong to 
Buathra (Townes, 1970). The paper is illustrated by five figures. 


ACKNOWLEDGEMENTS 


For the loan of specimens I am grateful to: W. J. Pulawski, Muzeum Zoologiczne, 
Wroclaw (ZI); D. S. Peters, Natur-Museum Senckenberg, Frankfurt (SM); M. Fischer, 
Naturhistorisches Museum, Vienna (NMW); B. Petersen, Universitetets Zoologiske 
Museum, Copenhagen (MC); S. Jonsson, Uppsala Universiteit (UU); H. Andersson, 
Department of Entomology, Lund University (ML); Rijksmuseum van Natuurlijke 
Historie, Leiden (RMNH); C. A. W. Jeekel, Zoölogisch Museum, Entomologische 
Afdeling, Amsterdam (MA); K. W. R. Zwart, Laboratorium voor Entomologie, Wage- 
ningen (ELW); and the private collectors, J. G. Betrem, Deventer; J. Glowacki, Poland; 
E. Haeselbarth, München, and H. Priesner, Linz. 

I am much indebted to my colleagues A. Noordijk, for drawings and D. L. J. 
Dijkstra, for advice on arithmetical problems. I am grateful to C. G. Johnson, Rotham- 
sted Experimental Station, for critical remarks and corrections. 


METHODS AND ABBREVIATIONS 


External measurements were taken with an ocular micrometer (1 cm at 10 x) on 
a Zeiss stereo microscope. Absolute measurements were taken at 10 x enlargement. 
The length of the ovipositor was measured from the apex of the gaster. Most relative 
measurements were taken at 40 X. For length: breadth relations I use the term index. 
As differences in indices are often very small, students are advised to take measurements 
carefully. For terminology I follow Richards (1956) and Townes (1969). 


OOL = distance from the outer edge of a lateral or posterior ocellus to the compound 
eye (ocular-ocellar line). 
POL = distance between the inner edges of the two lateral ocelli (postocellar line). 


1) For keys to the genera of Cryptina see Van Rossem (1969b) and Townes (1970). 
201 


202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 5, 1971 
INTRODUCTION 


The genus Bwathra was proposed by Cameron (1903). A full description was given 
by Townes (1970). Separation of Buathra from Meringopus is rather difficult and 
arbitrary as it is based on the variable position of the axillary vein in the hind wing. 
The establishment of Bzathra is thus controversial and in my opinion Buathra could 
be merged into Meringopus. An example of a species in an intermediate position is 
Meringopus reverendus Van Rossem, 1969a (Townes, 1970: 193). 

Townes (1970) placed Cryptus medius Szépligeti, 1916, in Buathra. This cannot 
be valid because Szépligeti’s lectotype does not have the dorsal tentorial pits. I consider 
C. medius to be a species of Cryptus (Van Rossem, 1969b). 

Assessment of western European Bwathra material shows that three taxa can be 
distinguished. One of these is B. divisoria, an alpine species, which presents no problems. 
The two others, in older literature referred to as “Cryptus tarsoleucus’ and “ Cryptus 
laborator”, were difficult to separate and at first offered no satisfactory evidence for 
considering them as independent species (Habermehl, 1918, 1920; Roman, 1903, 1910). 
However, after prolonged study I found a character that separates at least the females 
of the two above mentioned taxa with reasonable certainty. Whether males can be 
matched is uncertain. Consequently the lectotype of Thunberg’s (1822) Ichneumon 
laborator cannot contribute much to our knowledge. It is clear from his text that Roman 
(1912) did not take Thunberg’s species for the same as “Cryptus tarsoleucus’ of authors, 
thus we could for the sake of stability best follow Roman and continue to use the name 
“laborator” for the other taxon. Roman (1912) placed Cryptus fulvipes “Kriechbaumer 
ap. Magr.” in the synonymy of Cryptus laborator Thunberg. From Magretti's (1884) 
original description it is clear that also in this case there is question of male type 
material, which makes certain identification impossible. It should be noted, that Kriech- 
baumer never described a Cryptus fulvipes and consequently Magretti is the author. As 
the name fw/vipes has no priority, we can best leave the question for what it is. 


Key to the species 


Females 


1. Gena slightly concave just behind the lower articulation of mandible (Fig. 1). 
Ovipositor about the length of hind tibia. . . . . . B. divisoria (Tschek) 
— Gena not concave. Ovipositor conspicuously longer than hind tibia. . . . . 2 
2. Postanellus : ovipositor tip (distance between nodus and apex, Fig. 2) > 1.0. 
Ovipositor somewhat upcurved, nodus rather strong (Fig. 3). Anterior propodeal 


carina present. Index of hind femur > 5.6. . . . . B. tarsoleuca (Schrank) 

— Postanellus : ovipositor tip (Fig. 2) < 1.0. Ovipositor straight, nodus rather weak 

(Fig. 4). Anterior propodeal carina absent. . . . . B. laborator (Thunberg) 
Males 


1. The following characters in combination: gena concave just behind the lower 
articulation of mandible (Fig. 1). Propodeal spiracle almost round. Both propodeal 
carinae well developed, posterior one sublaterally rather strongly dentated . 

B. divisoria (Tschek) 


G. VAN ROSSEM: Genus Buathra 203 


u @haracterse not combinednastaboven. mr, ET NE sia) Se fy deel 2 
2. Anterior propodeal carina present. . . . . . . . B. tarsoleuca (Schrank) 
— Anterior propodeal carina wholly or partly absent . . B. laboraior (Thunberg) 


Remark : this distinction between the males of B. tarsoleuca and B. laborator is 
unreliable. 


Buathra tarsoleuca (Schrank, 1781) 


Ichneumon tarsoleucus von Paula Schrank, 1781, Enumeratio insectorum austriae indigenorum: 
359 (type lost: nomen dubium), &. 
Cryptus curvicauda Thomson, 1896, Opusc. Ent. 21: 2350, Q. 


The type material of Ichneumon tarsoleucus Schrank, all males, is lost. Irrespective 
of the fact that males remain undeterminable, it is not possible to decide on the identity 
of Schrank's material from the short description. Thus Ichneumon tarsoreucus is a 
nomen dubium. The interpretation of “Cryptus tarsoleucus’ of authors goes back to 
Gravenhorst (1829: 447) and his first reviser, Taschenberg (1865: 71). A specimen 
that is probably an original of Gravenhorst is still in the collection at Wroclaw, but it 
is rather aberrant from the general conception of authors, nor is this specimen mentioned 
by Gravenhorst or Taschenberg; therefore I have not selected it as the neotype but have 
marked it with an orange label stating: “Cryptus tarsoleucus (Schrank) 3rd specimen 
in Gravenhorst coll.” 

Some important characters of the female. — Apical part of ovipositor between nodus 
and tip (of upper valve) (Fig. 3) shorter than postanellus. Index: > 1.0. Anterior 
propodeal carina mostly present, at least medially. Gaster with a tendency towards a 
deep reddish brown or purple reddish tone. Ovipositor nearly always upcurved, nodus 
strong (Fig. 3). 

Male. — I have not succeeded in associating the proper male with the tarsoleuca 
female. The character given in the key (presence of anterior propodeal carina) is un- 
reliable. 

With respect to the type material of Cryptus curvicauda Thomson, H. Andersson, 
at Lund, reported as follows: There are no specimens under this name in Thomson’s 
collection. All the other species mentioned in Opzsc. Ent. 21 are represented in the 
collection and also another species labelled “bellitarsis”. This “bellitarsis’" is represented 
by six females, labelled “OG” = Sweden, Östergôtland (the type area of curvicanda). 
I think that the females from Östergôtland labelled as bellitarsis must be the syntypes 
of curvicauda and that Thomson first intended to use that name and forgot to change 
the label. 

Characteristics of the lectotype of Cryptus curvicauda. Female, 16.0 mm. Front wing 
11.8 mm. Labels: a small white tag “OG”, a modern green label “1968/11”; a red 
rimmed box label “belletarsis m.” in ink. Index of apical part of ovipositor and post- 
anellus, 1.2. Anterior propodeal carina indicated mediaily. Gaster reddish brown. 
Ovipositor curved, nodus strong. Lectotype labelled accordingly and identified as B. 
tarsoleuca. Two other specimens were labelled paralectotype. 

Occurrence. — The female is likely to occur on peat-moors, perhaps in the vicinity 
of fens. 

Material examined. — France: &, La Grave, 1500 m, leg. Becker (coll. Dittrich) 
(ZI). Italy: g', Bolzano, VI.1913, leg. Smits van Burgst (ELW). Netherlands: 9? 


204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 5, 1971 


Gieten, 17.VI.1950 (coll. Betrem); 9, Dwingelo (Dr.), Davidsplassen, 27—31.VII. 
1963, leg. Ph. Pronk (RMNH); 2, Terlet (Gld), 19.IX.1966, leg. R. T. Simon 
Thomas (PD); 9, Leuvenum, VII.1926, leg. H. J. Klaassen (MA); 9, Winterswijk, 
29.VI.1921, moeras bij de Vlijt (?), coll. Koornneef (ELW); 2, Oisterwijk, 1.VIII. 
1921, op Peucedanum palustre, coll. Koornneef (ELW); g, den Dolder, 21.V.1931, 
op Anthriscus, leg. Bouwman, coll. Koornneef (ELW). Poland: 9, Orlach (near 
Wroclaw ?), 13.VII.1880, coll. Dittrich (ZI); 9, Myszyniec, 1949 (coll. J. Glowacki). 
Sweden: 3 9, Östergôtland, coll. Thomson (lectotype and paralectotypes of Cryptus 
curvicauda) (ML). 


Buathra laborator (Thunberg, 1822) 


Ichneumon laborator Thunberg, 1822, Zap. imp. Akad. Nauk 8: 273, 4. 

Cryptus fabricii Schigdte, 1857, in Rink, Gronland geographisk og statistisk beskrevet 2 (3): 
62 Q 6: 

Cryptus fulvipes Magretti, 1884, Bull. Soc. Ent. Ital. 16: 99, 4. 


Characteristics of the lectotype of Ichneumon laborator. Male, 12.9 mm. Front wing 
8.5 mm. Labels: one label in ink with Roman’s handwriting “Cryptus laborator Thbg” 
(Roman, 1912). With the specimen is an old box label in ink (Thunberg’s hand- 
writing?) “laborator T.”, in the left bottom corner is written “Sv” (Suecia?). Lectotype 
hereby designated. Head black. Antennae with tyloidae on segments 18—26. Frons rather 
concave, with a distinct pit dorsolaterally of each antennal socket. Thorax black. Anterior 
propodeal carina mesally indicated. Posterior propodeal carina complete, sublaterally 
dentated. Index of propodeal spiracle, 2.5. Wings slightly infuscate. Nervulus somewhat 
antefurcal. Axillary vein (3A) in the hind wing convergent to inner margin. Index of 
hind femur, 6.0. All femora yellow red. Gaster slender, with first tergite black, polished. 
The following tergites alutaceous, with adpressed hairs, blackish with a ferruginous 
undertone. The hind margin of tergite 2 fulvous. Apex of clasper truncate’). 

I regret to say that I have been unable to identify Thunberg’s specimen with any 
certainty. I think that at present it is best to follow Roman (1912) and thus take the 
name “laborator” for the “other” taxon. The female is easily separated from the 
tarsoleuca female (compare the key). 

With respect to Cryptus fabricii Schigdte I can make the following observations. 
There are two syntypes in the Universitetets Zoologiske Museum at Copenhagen (a 
male and a female). A lectotype has not been selected up to now. As far as I can 
judge the male closely agrees with Dutch males of Bwathra laborator. Through the 
kindness of Dr. B. Petersen I was able to study a female from Greenland. This 
specimen falls within the range of ordinary European material of B. laborator. I see no 
special reason to distinguish a subspecies for material from Greenland (Townes & 
Townes, 1962). Apart from B. laborator laborator and B. laborator fabricii, Townes & 
Townes (1962) recognized three other subspecies. A note on Cryptus fulvipes Magretti 
was given in the introduction. 

Biology. — Buathra laborator is a parasite of Lepidoptera. The Laboratorium voor 
Entomologie at Wageningen (ELW) has a Q bred from a pupa of either a Noctuid or 
a Notodontid, found in a clump of grass at Wageningen. The cocoon of the wasp 


') I believe this character to be unreliable. 


G. VAN RossEM: Genus Buathra 205 


measures 17 mm and is almost black in colour with a conspicuous greyish band in the 
middle. The aperture of escape lies immediately dorsad (or ventrad) of the pole. 
K. W. R. Zwart kindly informed me that he was able to breed this species on larvae of 
Galleria mellonella (Linné, 1758) which had just formed cocoons, thus suggesting a 
pupal parasite. Zwart found the larva to be ectoparasitic, as should be expected. In the 
laboratory also a cocoon of a S wasp was obtained on Galleria mellonella. The 
colour of this cocoon is lighter than of the other specimen, but it also shows the central 
band (length: 14 mm). 

Material examined. — Austria: 9, Oberweiden, Marchfeld, A.i., leg. Hammer (coll. 
Priesner); 9, Weidling, Ai. Wien, Mader (coll. Priesner). Denmark: 9 g', Gron- 
land (syntypes of Cryptus fabricii Schiodte) (MC); 9, Gronland, leg. Vahl (Mus. 
Westerm.) as C. glabrator Zetter. (MC). England: § 9, England (no data) 1949 
(coll. Betrem). France: 9, BA, la Foux d’Allos, 1700 m, 29.VII.1964 (leg. M. C. 
and G. Kruseman) (MA), Germany: g', Ransern(?), 15.V.1890 (coll. Dittrich) 
(ZI); 9, illegible, 4.VIII.1905 (coll. Dittrich) (ZI). Netherlands; ©, Ellecom, 
5.IX.1922 (coll. Koornneef) (ELW); 9, 20.VI.1940 (coll. Koornneef) (ELW); 
d', Groesbeek, 2.VI.1898, leg. De Vos (coll. Oudemans) (MA); 2 9, Wa- 
geningen, 19.VI.1963 & 21.III.1965 (ELW); 3 9, Wageningen, resp. mother 
& 2 daughters & 1 & (son) (leg. Zwart) (ELW); ®, Neerijnen (Gld.), 15—30.VI. 
1950 (leg. C. de Jong) (RMNH); the following females from Rhenen (coll. Koorn- 
neef) (ELW) 29.V.1919; 16 & 18.VIII.1930; 12.VI.1932; 6 & 18.VIII.1933; 26.VII. 
OSD ENIS TO VALI 3550 135V.19363,17, 2648 27.V11.1936; 30.VII.19367(2 sp.); 
SV 936, 22spr): 11,1 16° & 25. VIll1936; 24 & 25.919375; TNO 2 8: 
Rhenen, 18.V.1934 (ELW); ®, Hilversum, 17.VI.1952 (coll. den Hoed) (ELW). 
Poland: 9, Nimptsch (south of Wroclaw) (coll. Dittrich) (ZI); 2 9, Podkowa 
LeSna ad Warszawa, 14.VI.1953 (4?) and 1 9, 26.VII.1953 (coll. J. Glowacki). 
Sweden: g', lectotype of Ichneumon laborator Thunberg (UU). 


Buathra divisoria (Tschek, 1872) 


Cryptus divisorius Tschek, 1872, Verh. zool.-bot. Ges. Wien 22: 235, @ 4. 


The type series (2 9 & 1 g') of Cryptus divisorius is extant and kept in the Vienna 
Museum. The species should be placed in the genus Bwathra. 

Characteristics of the lectotype of Cryptus divisorius. Female, 8.4 mm. Front wing 
6.9 mm long. Labels: a white tag “1/10 866” in ink (Tschek’s writing); a printed 
label “Tschek 1872 Piesting”; “divisorius Tsch. det. Habermehl.” Lectotype hereby 
designated. 

Head black. Labial and maxillary palpi brown. Mandibulae fuscous. Clypeus strongly 
convex, polished, with some scattered punctures. Face alutaceous, closely and finely 
punctured, with adpressed, silvery hairs. Facial convexity shown. Lower half of frons 
concave, polished, with some transverse ridges and conspicuous tentorial pits dorso- 
laterad of antennal sockets. Upper frons somewhat rugose. OOL-region alutaceous. 
Left antenna missing. Right antenna missing beyond 6th segment. Postanellus slender, 
index 6.6. OOL : diameter posterior ocellus, 1 : 1. OOL : POL, 3 : 4. Vertex somewhat 
coarse, with transverse wrinkling. Temple and gena polished towards outer orbit of 
compound eye, with scattered punctures, Toward genal carina with rather conspicuous 


206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 5, 1971 


wrinkling. Temple and gena with short grey bristles. Lower part of gena directly 
beyond mandibular base with a concavity (compare Fig. 1). This character is rather 
difficult to observe when the head is pressed against pleuron 1. Malar space coriaceous, 
not quite 1.5 the mandibular breadth. Inner and outer orbits and spot on eye margin 
at vertex narrowly marked yellow. 

Thorax black. Epomia strong. Mesoscutum polished, laterally with regular punctures, 
mesally somewhat coarse. Prescutal sutures well developed. Notum 1, episternum 2 and 
propodeum coarsely sculptured. Propodeum with anterior transverse carina present but 
weak. Posterior transverse carina well developed, sublaterally rather strongly dentated. 
Propodeal spiracles relatively small, index 2. Wings subhyaline. Nervulus (Cu-a) slight- 
ly antefurcal. Axillary vein (3A) in hind wing convergent to inner margin. Legs with 
front coxae, trochanters, femora (for the greater part), middle coxae, trochanters and 
femora (for the greater part) fuscous-ferruginous. Hind coxae and trochanters black, 
femora orange. Front tibia somewhat inflated, brownish. Tibiae and tarsi of other legs 
brown. 

Gaster with first segment black. Other tergites finely coriaceous, dark brown. Ovi- 
positor 2.8 mm. Ovipositor: hind tibia, 62 : 65 (Fig. 5). 


Fig. la, &, Buathra divisoria (Tschek), gena concave, Villnöss, 1965, leg. Van Rossem. 
Fig. 1b, the same, schematically 


Description of the male paralectotype of Cryptus divisorius. Length 9.0 mm. Front 
wing 6.3 mm long. Labels: a small green tag, a white printed label “Tschek 1872 
Piesting”, a box label “divisorius Kriech” (in purple ink), a white tag “divisorius 
Tsch. det. Habermehl”. 

Head black. Right antenna missing beyond 12th segment, left antenna missing beyond 
5th segment. Deep tentorial pits present dorsolaterad of antennal sockets. Gena concave 
directly behind the mandibular base (Fig. 1). Yellow markings on: mandibles, clypeus, 
inner orbits with a continuation on malar space, outer orbits. Spot on eye margin in 
OOL-region. 

Thorax black. Both propodeal transverse carinae well developed, the posterior one 
sublaterally dentated. Propodeal spiracle almost circular, index 1.3. Axillary vein (3A) 


G. VAN ROSSEM: Genus Buathra 207 


of hind wing convergent to inner hind margin. Legs with 2nd (partly), 3rd and 4th 
segment of hind tarsi white. Gaster with a ferruginous undertone. 

Material examined: Austria: 2 9 & 1 G', Piesting, leg. Tschek, the type series of 
C. divisorius (NMW); 1 gd, Obere Regalm, Kaisergebirge, Tirol, 1300 m, 9.VIII. 
1959 (coll. Haeselbarth); 1 g', Vorderkeiserfelden, Waisergebirge, Tirol, 1500 m, 
1.VII.1956 (coll. Haeselbarth). Italia: 9, Funes (Villnòss) (prov. Bolzano) 1100— 
1400 m, 13—30.V.1965, leg. G. van Rossem (PI. Prot. Serv.). Fürstentum Liechten- 
stein: ©, Triesenberg, 1450 m, 1.IX.1969 (coll. Haeselbarth). 


REFERENCES 


Cameron, P., 1903. — Descriptions of twelve new Genera and Species of Ichneumonidae (Here- 
siarchini and Amblypygi) and three Species of Ampulex from the Khasia Hills, India. — 
Trans. Ent. Soc. Lond. 1903: 219—238. 

Gravenhorst, J. L. C., 1829. — Ichneumonologia Europaea 2 (2), genus V; Cryptus. Vratislaviae. 

Habermehl, H., 1918. — Beiträge zur Kenntnis der palaearktischen Ichneumonidenfauna. — Z. 

wiss. InsektBiol. 14: 145—152. 

1920. — Beitrige zur Kenntnis der palaearktischen Ichneumonidenfauna. — Neue Beitr. 

syst. Insektenk. 1: 63—69. 

Magretti, Paolo, 1884. — Nota d'Imenotteri raccolti dal Signor Ferdinando Piccioli, nei dintorni 
di Firenze. Colla descrizione di alcune nuove specie e di un genere nuovo (V. Tav. Ila). 
— Boll. Soc. Ent. Ital. 16: 97—122. 

Richards, O. W., 1956. — Hymenoptera. Introduction and keys to families. — Handb. ident. 
British Insects 6 (I): 1—94. 

Roman, A., 1903. — Om nagra Cryptus-Arter, särskildt C. curvicauda Thomson. — Ent. Tidskr. 
24: 77-78. 

——, 1910. — Notizen zur Schlupfwespensammlung des schwedischen Reichsmuseums. — Ent. 

Tidskr. 31: 109—196. 

, 1912. — Die Ichneumonidentypen C. P. Thunbergs. — Zool. Bidr. Upps. 1: 229—293. 

Rossem, G. van, 1969a. — A study of the genus Meringopus Foerster in Europe and of some 
related species from Asia (Hymenoptera, Cryptinae). — Tijdschr. Ent. 112: 165—196. 

——, 1969b. — A revision of the genus Cryptus Fabricius s.str. in the western Palearctic region, 
with keys to genera of Cryptina and species of Cryptus (Hymenoptera, Ichneumonidae). 
— Tijdschr. Ent. 112: 299—374. 

Schigdte, J. C., 1857. — Udsigt over Gronlands land-, ferskvands- og strandbreds-arthropoder. In: 
Rink, H. J., Gronland, geographisk og statistisk beskrevet 2 (3): 62. 

Schmiedeknecht, O., 1931. — Opuscula Ichneumonologica. Suppl. Fasc. 10: 13—77. 

Schrank, F. von Paula, 1781. — Enumeratio insectorum austriae indigenorum. Augustae Vindeli- 
corum. Ichneumonidae: 344—379. 

Szépligeti, V., 1916. — Ichneumoniden aus der Sammlung des Ungarischen National-Museums. — 
Ann. Mus. Nat. Hung. 14: 225—380. 

Taschenberg, E. L., 1865. — Die Schlupfwespenfamilie Cryptides (Gen. V. Cryptus Gr.) mit 
besonderer Beriicksichtigung der deutschen Arten. — Z. ges. Naturw. 25: 1—142. 

Thomson, C. G., 1896. — Opusc. Ent. 21: 2348—2354. 

Thunberg, C. P., 1822. Ichneumonidea, Pars I. — Zap. imp. Akad. Nauk 8: 273. 

Townes, H. & M. Townes, 1962. — Ichneumon Flies of America north of Mexico: 3. Subfamily 
Gelinae, Tribe Mesostenini. — Bull. U. S. Nat. Mus. 216 (3): 1—602. 

Townes, H., 1969. — The genera of Ichneumonidae, part 1. — Mem. Amer. Ent. Inst. 11: 1—300. 

, 1970. — The genera of Ichneumonidae, part 2. — Mem. Amer. Ent. Inst. 12: 1—537. 

Tschek, C., 1872. — Ueber einige Cryptoiden meist aus der Osterreichischen Fauna. — Verh. zool.- 
bot. Ges. Wien 22: 231—252. 


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CES 


Fig. 2, 3, 4 and 5, ovipositors of Buathra: 2, distance between nodus and apex of ovipositor tip; 
3, B. tarsoleuca, Myszyniec, 1949, leg. Glowacki; 4, B. luborator, Rhenen, 30.vii.1936, leg. Koorn- 
neef; 5, B. divisoria, paralectotype 


G. VAN RosseM : Genus Buathra 


A NEW SPECIES OF CRYPTUS FROM THE 
CANARY ISLANDS (HYMENOPTERA, CRYPTINAE) 
by 
G. VAN ROSSEM 


Plant Protection Service, Wageningen 
Cryptus verutus spec. nov..!) 


Characteristics of the holotype. Female, 9.2 mm. Front wing, 7.5 mm. Labels: a white 
tag “Guimar”, 31.V.(19)65 — Tenerife, leg. I. Klimesch (written). Holotype, labelled 
accordingly. The specimen belongs to the collection of Prof. Dr. H. Priesner at Linz 
(Austria). 

Head black with slight yellow lining of inner and outer orbits. Clypeus strongly 
convex, polished with some scattered punctures from which relatively long, erect bristles 
arise. Facial convexity weak. Face regularly and closely punctured, with fine alutaceous 
microsculpture. Malar space : breadth mandibular base, 1 : 1 (broad). Scapi broadly pear- 
shaped. Postanellus slender, index 6.0. Antennal scrobes weakly concave, transversely 
wrinkled. Frons rugosely wrinkled. OOL: diameter posterior ocellus, 1 : 1. OOL narrow, 
alutaceous. Ocelli relatively large. Vertex with close punctures and somewhat transversely 
wrinkled. Temple and gena polished, finely and widely punctured with conspicuous, 
suberect hairs. 

Thorax black, only a yellow spot on notum 1 at dorsal apex of epomia, all over 
roughly sculptured, except mesoscutum and scutellum 2 which are polished with more 
widely spaced rough punctures. Entire thorax with erect to suberect strong bristles. Epomia 
strong. Prescutal sutures long, reaching wing base, with rough transverse wrinkles. 
Speculum large. Wings somewhat infuscate. Areola with strongly converging sides, 
closely resembling that in C. immitis Tschek. In the hind wing, medial cell long, with 
sides (M + Cu-1 and 1A) parallel. Nervellus intercepted far below the middle, the 
angle with M + Cu-1 right. Cu-1 reaching wing margin. Axillus gradually converging 
towards inner hind margin of hind wing. Propodeum with anterior transverse carina 
obsolescent; posterior transverse carina well developed in the sublateral region, there 
forming a rather strong ridge. Propodeal spiracle small, elliptic. Legs with all coxae and 
most of the trochanters black. All femora orange; indices: 4.5 (1); 5.5(2); 5.7 (3). 

Gaster with petiole slender. Petiole (measured from dorsal edge of orifice to 
spiracle): postpetiole, 5 : 2. Petiole black, rest of gaster orange. All tergites sub- 
polished, with vague indication of microsculpture. Ovipositor slightly curved, ex- 
ceptionally long, 1.0 length of front wing, with a long tapering tip. 

Cryptus verutus hierroensis subsp. nov. 
A specimen from the island of Hierro (Canaries) agrees reasonably with the specimen 


1) Verutus = armed with a spear. 
209 


210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 5, 1971 


described above. Some characters rather suggest that it belongs to a geographically 
isolated population. 

The following description relates to the holotype, one of two specimens under the 
label Cryptus hellenicus in Schmiedeknecht’s material in the Berlin museum (compare 
my paper on Meringopus, 1969a). 

Characteristics of the holotype. Female, 9.5 mm. Front wing, 7.5 mm. Labels: a light 
blue tag Hierro (Canar. Ins.) Hinz V. (printed); 9.IV.(18)98 (written). A large 
white label with Schmiedeknecht’s writing: “Cryptus n. sp. dicht bei C. hellenicus 
Schmied’. Holotype, labelled accordingly. 

Head black with slight yellow lining of inner and outer orbits. Clypeus strongly 
convex, polished, with some scattered punctures. Hairs spoilt, but relatively long. Facial 
convexity weak. Face regularly and closely punctured, with fine alutaceous microsculpture. 
Malar space: breadth mandibular base, 6:5. Scapi broadly pear-shaped. Postanellus 
slender, index 7.4. Antennal scrobes weakly concave, lower part polished. Frons 
rugosely wrinkled. OOL: diameter posterior ocellus, 1 : 1. OOL narrow. Ocelli relatively 
large. Vertex closely punctate, somewhat transversely wrinkled. Temple and gena 
polished, finely and widely punctured, with long hairs (spoilt). 

Thorax black, only a yellow spot at dorsal apex of epomia; all over roughly sculptured, 
except mesoscutum and scutellum 2, which are polished, with more widely spaced rough 
| punctures. Entire thorax with erect to suberect strong bristles. Epomia indicated. 
Prescutal sutures long, reaching beyond base of fore wing, with rough transverse 
wrinkles. Wings somewhat infuscate. Areola with strongly converging sides. In the 
hind wing, medial cell long, with sides (M + Cu-1 and 1 A) parallel. Nervellus 
intercepted far below the middle; the angle with M + Cu-1 right. Cu-1 reaching wing 
margin. Axillus converging towards hind margin. Propodeum with anterior transverse 
carina obsolescent; posterior transverse carina present, bluntly and weakly dentate sub- 
laterally. Propodeal spiracle small, elliptic. Legs with all coxae and trochanters black. All 
femora orange; indices: 4.6 (1); 5.6 (2); 5.2 (3). 

Gaster. An important difference with the nominate form lies in the relation between 
the lengths of petiole and postpetiole. Petiole (measured from dorsal edge of orifice 
to spiracle) : postpetiole, 4.5: 2. Petiole definitely shorter and stouter than in the 
nominate form, black. Rest of gaster orange. All tergites finely alutaceous. Ovipositor, 
0.96 length of front wing, slightly curved. Tip resembling that in the nominate form. 

Male. — Unknown. 


Discussion. — Cryptus verutus shows relationship towards Cryptus immitis Tschek. 
There are nevertheless some important differential characters; a. the ovipositor longer, 
0.96—1.0 length of front wing; 0.63—0.73 in immitis, b. only anterior transverse 
carina present; in immitis both carinae in most specimens strong, with indication of 
area superomedia; c. tergites 2 and 3 almost polished to subpolished; in zmmitis rather 
conspicuously granulated; d. antennae without white band; in zmmitis with a white band. 

In my key (Van Rossem, 1969b : 310) to the females of Cryptus, C. verutus should 
be placed in item 11 as follows: 


11. Ovipositor longer than 0.80 length of front wing. Tergites 2 and 3 subpolished or 
with very fine microsculpture. Only anterior carina of propodeum present . 
C. verutus sp. n. 


G. VAN ROSSEM: A new Cryptus 211 


— Ovipositor shorter than 0.80 length of front wing. Tergites 2 and 3 conspicuously 
alutaceous or finely granulated. Usually both propodeal carinae well indicated . 


11a & b (as in my key) 


ACKNOWLEDGEMENTS 


For the loan of specimens I am indebted to Prof. Dr. H. Priesner (Linz, Austria) 
and Dr. E. Königsmann, Zoologisches Museum (Berlin, DDR). 


REFERENCES 
Rossem, G. van, 1969a. — A study of the genus Meringopus Foerster in Europe and of some 
related species from Asia (Hymenoptera, Cryptinae). — Tijdschr. Ent. 112: 165—196. 


, 1969b. — A revision of the genus Cryptus Fabricius s.str. in the Western Palearctic Region 


with keys to genera of Cryptinae and species of Cryptus (Hym., Ichneumonidae). — 
Tijdschr. Ent. 112: 299—374. 


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ADDITIONAL NOTES ON THE GENUS TRYCHOSIS 


FOERSTER IN EUROPE (HYMENOPTERA, CR YPTINAE) 
by i 
G. VAN ROSSEM 


Plant Protection Service, Wageningen 


Trychosis priesneri spec. nov. 


Characteristics of the holotype of Trychosts priesneri. — Female, 9.3 mm. Front wing 
6.7 mm. Labels: a white tag “Winden, Bgld. [Burgenland] 12.VII.1959 H. Priesner”. 
Holotype, labelled accordingly. The specimen is in the collection of Prof. Dr. H. 
Priesner, Linz, Austria. 

Head black, roughly sculptured, relatively small. Thorax black. Mesoscutum strikingly 
high, giving the thorax a hunchbacked appearance. Vertical carina on anterior part of 
epicnemium developed into a rounded tooth-like structure. Gaster with postpetiole and 
tergites 2,3 and 4 (in part), orange. Apex fuscous. Ovipositor, 0.17 length of front 
wing, relatively short. 15 other specimens, including 3 males, were labelled paratype. 


Description of the female. — Front wing 6.0—7.5 mm. Head relatively small, black, 
palpi brown. Closely and roughly punctured; with conspicuous, erect silvery hairs. 
Clypeus strongly convex. Antennae brown, ventrally reddish, index of postanellus 
3.5—4.1. Antennal scrobes scarcely indicated. Frons roughly punctured. 

Thorax black, with erect silvery hairs. Pronotum closely and roughly punctured with 
strong crista. Mesoscutum strongly rising above the level of the ocelli, giving the 
specimen a hunchbacked appearance, more widely punctured and polished between. 
Scutellum rather convex, with strong punctures. Vertical carina on anterior part of 
epicnemium developed into a rounded tooth-like structure as in T. tréstator, Episternum 
2 with rough punctures almost touching, interspaces shiny. Propodeum large, strongly 
declivitous towards propodeal orifice, closely sculptured; with anterior transverse carina 
in most specimens complete; posterior transverse carina sublaterally present. Wings 
somewhat fuscous. Nervellus intercepted above the middle. Coxae fuscous-red to black. 
Femora rather variable in colour between fuscous-yellow and black (hind femora). 
Tarsi between fuscous-yellow and brown. Femora rather stout; index front femur, 
3.4—4.0; hind femur, 4.6—5.3. 

Gaster with postpetiole, tergites 2, 3 and 4 (in part) orange, polished, with minute 
puntures from which adpressed to suberect microsetae arise. Apex fuscous. Ovipositor 
strikingly short, 0.16—0.17 length of front wing. 


Male. — For description three males were available. The colour pattern of these 
closely resembles that of the female. The most characteristic feature of the female, viz. 
the hunchbacked thorax, also occurs in the male and makes it rather easily recognizable. 

Head black, rather small, flat in the antero-posterior line, closely and roughly sculp- 
tured, with erect silvery hairs. Clypeus convex. Antennal scrobes little pronounced. 
Tyloids on antennal segments 12—17. 

213 


214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 5, 1971 


Fig. 1. Trychosis priesneri, 9, Konya Asiat. Türk.) (coll. Priesner), X 6.5 - 


Thorax black, mesoscutum strongly rising above the level of the ocelli. Pronotum, 
mesoscutum and episternum 2 with strong punctures, interstices shiny. Vertical carina 
on anterior part of epicnemium developed into a rounded tooth-like structure. Scutellar 
fovea narrow and deep. Scutellum 2 strongly convex. Propodeum roughly sculptured. 
Anterior transverse carina complete; posterior interrupted medially. Coxae fuscous-red 
to black. Front and middle femora yellowish-red; front and middle tibiae and tarsi 
yellow, dorsally with a rather conspicuous streak of white. Hind legs fuscous, including 
the tarsi. 

Gaster with postpetiole, tergites 2, 3 and 4 (in part) orange, subpolished, with 
fine punctures, implantations of adpressed microsetae. Apex fuscous. 

In my key (Van Rossem, 1966) the females of Trychosis priesneri are inserted as 
follows: 


2. Vertical carina on anterior i of ae developed into a rounded tooth-like 


SEEUCEUTE MEME: i & mme 
3. Tergite 2 subpolished, ath very dese setae ani Focale, Toe not con- 
spicuously rising above the level of the ocelli . . T. tristator and T. glabricula) 
— Tergite 2 polished, setae and microsculpture more widely placed. Thorax hunch- 
backedy(Figt ya tim eo yo 200.00. Mebo eee 
Separation of the males: 
1. Vertical carina on anterior part of epicnemium with a strong, tooth-like structure . . 2 
2. Front and middle tibiae with a rather a white streak. Hind leg fuscous. 
Thorax strongly hunchbacked . . . . art ENT 027 
— Front and middle tibiae without a white cingolo foi tarsi with a white band. 
Thorax notrhunchbackedens esta un. warten nt, bemeten AREN 


Distribution. — The species seems to be widely distributed in the central and southern 
regions of Europe. The greater part of the material which I studied was kindly placed at 
my disposal by Professor Dr. H. Priesner, Linz (Austria), who told me that he had 
known this species since 1959. I recognized it when studying Palearctic Trychosis 
material sent to me by Dr. Henry Townes. I am glad to name this species after Dr. 
Hermann Priesner. 


1) T. glabricula is possibly only a small form of T. ¢ristator. 


G. VAN ROSSEM: Notes on the genus Trychosis 215 


Material examined. — Austria: 9, Winden, Bgld., 12.VII.1959, leg. H. Priesner 
(coll. Priesner) (the holotype); ®, Winden, Bgld., 12.VII.1959, leg. H. Priesner; 
@, Winden, Bgld., 9.VII.1964, leg. H. Priesner; g', Winden, Bgld., 4—7.VII.1959, 
lern Eisspriesner; O° Winden, Beld: 12.NILE1959, leg. H. Priesner; 9, Winden, 
10.VII.1962, leg. H. Priesner; 9, Umgebung Wien, 10.VI.1958, leg. K. Kusdas (coll. 
Priesner); 9, Neusiedlersee, Umg., Bgld., 11.VII.1960, leg. Kusdas (coll. Priesner) ; 
Q, Neusiedlersee, Umg., Bgld., 14.VII.1961, leg. Kusdas (coll. Priesner). France: à, 
Carpentras (Vaucl.), 24—27.V.1952, leg. Verhoeff (coll. PD)*); g', Carpentras 
(Vaucl.), 1—3.V111.1953, leg. Verhoeff (coll. PD)*). Grecce: 9, Sitia (Kreta), 20.V. 
1963, leg. Kusdas (coll. Priesner). Switzerland: 9, Wallis (ex coll. Berlin) (coll. 
Townes). Turkey: 9, Konya, Asiat. Türk, 26.V.1965, leg. J. Schmidt (coll. Priesner). 
Yugoslavia: 9, Hvar (Dalmatia), 10.VI.1962, leg. Gusenleitner (coll. Priesner). Label 
illegible: 9, Ostrów Mar (?), 26.VII.1958, coll. Glowacki. 


Trychosis gradaria (Tschek, 1870) 


Cryptus gradarius Tschek, 1870, Verh. zool.-bot. Ges. Wien 20: 151—152. 
Goniocryptus punctatus Szépligeti, 1916, Ann. Mus. Nat. Hung. 14: 250. Syn. nov. 
Trychosis gradaria: Van Rossem, 1966, Zool. Verh. 79: 16. 


The specimen of Gontocryptus punctatus in Szépligeti’s collection at Budapest is a 
female of Trychosis gradaria (Tschek, 1870), which thus appears to be a senior synonym 
of punctatus. As the author mentions only one specimen, this is the holotype and I have 
labelled it as such. 

Characteristics of the holotype of Gontocryptus punctatus. Female, 12.4 mm. Front 
wing 7.5 mm. Labels: P. Maróth (= Pilis-Maróth) Szépligeti; a red tag: a label with 
Szépligeti’s writing “Goniocryptus 18. punctatus n.sp.” (18 is the number, not meaning 
18-punctatus). Index of front femur, 3.3. Index of radial cell, 4.1. Ovipositor 1.8 mm. 


REFERENCES 


Rossem, G. van, 1966. — A study of the genus Trychosis Foerster in Europe (Hymenoptera, Ich- 
neumonidae, Cryptinae). — Zool. Verh., Leiden 79: 1—40. 

Szépligeti, V., 1916. — Ichneumoniden aus der Sammlung des Ungarischen National-Museums. 
— Ann. Mus. Nat. Hung. 14: 225—380. 


*) These specimens were placed under T. #ristator in my paper of 1966. 


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No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.F. 45.—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.Fl. 70.—. 


No, 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov 
and Xenocataniops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. 
D.FI. 25.—. 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. 
D.FI. 100.—. 


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B. THEOWALD. — Die Tipuliden der Benelux-Länder (Diptera, Tipulidae), p. 217— 
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DIE TIPULIDEN DER BENELUX—LANDER 
(DIPTERA, TIPULIDAE) 


von 


B. THEOWALD!) 


Zoölogisch Museum, Amsterdam 


ZUSAMMENFASSUNG 


Seit Goetghebuer & Tonnoir (1920—1921) und De Meijere (1939) sind keine Listen mehr 
veröffentlicht worden über die gesammte Tipuliden-Welt eines der Benelux-Länder. Nach einer 
kurzen Beschreibung der Landschaft geben wir eine Liste der Tipulinae, Cylindrotominae und 
Limoniinae. In dieser Liste wird erstens verzeichnet in welchem der Benelux-Länder die Arten 
vorkommen, zweitens in welchen Landschaften sie bisher gesammelt worden sind. Folgen noch 
Bemerkungen über Arten, die nur wenig gefunden sind. Hier auch die Beschreibung von Limno- 
phila (Limnophila) arnoudi spec. nov. 


KURZE BESCHREIBUNG DER LANDSCHAFT UND WICHTIGSTE SAMMELGEBIETE 


„Die Gegend... ist der Erzeugung von Bachmiicken 
besonders giinstig. Viele Siimpfe...., die nicht unbe- 
deutenden, hin und wieder Sumpfungen, Gräben, feuchte 
Wiesen einschliessenden Waldungen,.... die Ufer der 


langsam fliessende (Flüsse) sind der Entstehung dieser 
Thiere besonders beförderlich. Man wird daher den 
Reichthum unsrer Gegend an Arten leicht erklärbar 
finden.” 


Schummel, 1813: Vorrede. 

Dasselbe können wir auch sagen von den Benelux-Ländern: Belgien, Holland und 
Luxemburg, wo auf etwa 65.000 qkm jetzt 100 Tipulinae, 4 Cylindrotominae und 188 
Limoniinae bekannt sind. Zum Vergleich: Finnland, 337.000 qkm, 105 Tipulinae 
(Mannheims, 1967); Italien, 301.000 qkm, 130 Tipulinae (Mannheims & Theowald, 
1959); Schweden, 449.000 qkm, 117 Tipulinae, 4 Cylindrotominae und 202 Limoniinae 
(Tjeder, 1955). 

Die Benelux-Lander zeigen auf einem kleinen Gebiet eine grofie Abwechslung durch 
Landschaften (siehe Karte). An der Küste entlang finden wir Sanddünen (1), trocken 
und warm, mit Diinenkesseln, die meist etwas feuchter sind. 

Hinter den Diinen (2) gibt es feuchte Gebiete, meist unter dem Meeresspiegel, von 
Diinen und Deiche gegen das Wasser geschiitzt. Es ist das Gebiet von Wiesen und 
Flachmooren, auch von Tongebieten mit Landwirtschaft und von vereinzelten kleinen 
Waldern mit Alnus, Salix und Populus. Im Norden dieses Gebietes sind auch salzige 
Küstenwiesen. 

Der östliche Teil von Holland und der anschließende Teil von Belgien (3) haben 
trocknere Sand- und Tonboden mit Wiesen und Landwirtschaft, aber auch mit Wäldern 
von Betula, Quercus, Carpinus und Fagus. Hier gibt es auch Heiden und Hochmoore 
oder Restante davon. Das äußerste Osten dieses Gebietes, anschließend an Deutschland, 
hat etwas mehr Landklima. 


1) Anschrift des Verfassers: Instituut voor Taxonomische Zoologie (Zoologisch Museum), afd. 
Entomologie, Zeeburgerdijk 21, Amsterdam-O. 
PANG) 


218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 1971 


Nordsee 


D 
< 
= 
= 
O 
u) 
. 
9 
o) 
a 


Karte 1. Die Benelux-Länder. Einteilung in Landschaften (schwere Linie) und die wichtigsten 
Stellen wo gesammelt worden ist (schraffiert). Erläuterung im Text 


B. THEOWALD: Die Tipuliden der Benelux-Länder 219 


Das Hügelland von Südlimburg und den Ardennen (4) steigt von 100 bis zu 
600 m hinauf. Zum Teil wird dieses Gebiet benutzt für Wiesen und Landwirtschaft. 
Hier finden wir aber auch größere Wälder mit Nadel- und Laubholz. Die südlichen 
Täler der Ardennen sind auffallend wärmer als der Rest der Benelux-Länder und dort 
finden wir Pflanzen und Tiere eines viel südlicheren Areals. 

Die Hochmoore der Hautes-Fagnes (5) auf ungefähr 900 m sind kalt und haben 
mehrere subalpine Pflanzen- und Tierarten. 

Diese große Abwechslung von feuchteren und trockneren, kälteren und wärmeren, 
flachen und hügeligen Gebieten mit allerhand Bodenarten und Vegetationen macht es 
verständlich daß in den Benelux-Ländern viele europäische Tipuliden-Arten einheimisch 
sind. 

Obwohl seit etwa 1850 in vielen Gebieten der Benelux-Länder nach Tipuliden ge- 
sucht worden ist, können wir noch wenig sagen über ihre genaue Verbreitung. In 
einigen Gebieten ist mehrere Jahre intensiv gesammelt worden, in andern nur dann 
und wann einige Tage. Infolge dessen sind oft seltenere Tipuliden-Arten nur bekannt 
von Stellen wo intensiv geforscht wurde, z.B. von Amsterdam (!), wo De Meijere von 
1890 bis 1940 sammelte. Dergleiche Arten werden aber gewiß auch wohl an weniger 
städtischen Stellen vorkommen. 

An den nachfolgenden Stellen ist intensiv gesammelt worden (Karte): 

a. das Gebiet zwischen Haag und Leiden: Van der Wulp, 1850—1900; 

b. die Dünen westlich von Haarlem: De Meijere, 1890—1940; 

c. Amsterdam und die Gebiete südlich dieser Stadt: De Meijere, 1890—1940; 

d. die Gebiete südöstlich von Amsterdam: Theowald, 1950—1960; 

e. die Umgebung von Venlo: Van den Brandt, 1860—1900; 

f. Gent und Umgebung: Goetghebuer, 1915—1925; 

g. Brüssel und Umgebung: Tonnoir, etwa 1920; 

h. Südlimburg: Br. Arnoud, 1950—1965; 

j. die Hochmoore der Hautes-Fagnes und Umgebung: Collart und andere vom Museum 

Brüssel, 1950— 1955; 

k. das ganze Ardennen-Gebiet: De Laever, 1954—1970. 

Im Vergleich mit vielen andern europäischen Ländern sind die Benelux-Länder 
eingehend auf Tipuliden untersucht worden. Es wird aber auch hier noch lange dauern 
bis wir von allen Arten die Verbreitung bis in Einzelheiten kennen. 


ARTENLISTE 
Verzeichnis der Abkürzungen: 
Land: H, B oder L: die Art kommt vor in Holland, Belgien oder Luxemburg; 
—: die Art ist im diesbezüglichen Lande nicht nachgewiesen. 
1, Küstendünen, 
2, westliche feuchtere Gebiete, 
3, zentrale und östliche trocknere Gebiete, 
4, Hügelland von Südlimburg und Ardennen, 
5, kältere Hochmoore der Hautes-Fagnes und Umgebung. 
— nicht in dem Gebiet nachgewiesen. 
. ist in diesem Gebiet nur an vereinzelten Stellen nachgewiesen worden, 
+ ist in diesem Gebiet an mehreren Stellen nachgewiesen (mehr als 5 Stellen). 
* vor den Artnamen siehe unter: Bemerkungen über einzelne Arten. 


220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 


Tipulinae 


Dolichopeza Curtis, 1825 
albipes (Ström, 1768) 
Ctenophora Meigen, 1818 (= Flabellifera Meigen, 1800) 
Subg. Ctenophora Meigen, 1818 
elegans Meigen, 1818 
festiva Meigen, 1804 
flaveolata (Fabricius, 1794) 
*guttata Wiedemann in Meigen, 1818 
“ornata Wiedemann in Meigen, 1818 
pectinicornis (Linnaeus, 1758) 
Subg. Tanyptera Latreille, 1804 
atrata (Linnaeus, 1758) 
nigricornis (Meigen, 1818) 
Subg. Dictenidia Brullé, 1833 
bimaculata (Linnaeus, 1761) 
Nephrotoma Meigen, 1818 (= Pales Meigen, 1800 
= Pachyrhina Macquart, 1834) 
aculeata (Loew, 1871) 
“analis (Schummel, 1833) 
appendiculata (Pierre, 1919) (= Tipula maculata 
Meigen, 1804, nec Tipula maculata Linnaeus, 1758 
= Tipula maculosa Meigen, 1758 = Tipula macu- 
losa Gmelin ir Linnaeus, 1790) 
cornicina (Linnaeus, 1758) 
crocata (Linnaeus, 1758) 
dorsalis (Fabricius, 1781) 
“flavescens (Linnaeus, 1758) (= lineata Scopoli, 
1763) 
flavipalpis (Meigen, 1830) 
guestfalica (Westhoff, 1880) 
*lindneri (Mannheims, 1951) 
lunnlicornis (Schummel, 1833) 
pratensis (Linnaeus, 1758) 
*quadrifaria (Meigen, 1804) 
*quadristriata (Schummel, 1833) 
*scalaris (Wiedemann in Meigen, 1818) 
“scurra (Meigen, 1818) 
*submaculosa Edwards, 1928 
Prionocera Loew, 1844 (= Stygeropsis Loew, 1863) 
*pubescens Loew, 1844 
subserricornis Zetterstedt, 1851 
turcica (Fabricius, 1781) 
*Tipula Linnaeus, 1758 
Subg. Tipula Linnaeus, 1758 
czizeki De Jong, 1925 
“oleracea Linnaeus, 1758 
*paludosa Meigen, 1818 
Subg. Yamatotipula Matsumura, 1916 
caesia Schummel, 1833 
*coerulescens Lackschewitz, 1923 
couckei Tonnoir, 1921 
lateralis Meigen, 1804 
marginata Meigen, 1818 


Land 

H B L 
gi B — 
Ri BD — 
El D — 
et D — 
Hit 
gi 3 — 
gi Ba 
Ei B 
H B — 
H B — 
Ri B — 
EB IL, 
= 
H B — 
IE 13 = 
iat B = 
He Besse 
Ri 18) — 
EL B — 
Hh Be 
Rig — 
El Br 
EB 

EIB 
Ent 
Jal 18 IL 
El D = 
BL — 
EME 
ELI — 
Fy ees 
jel 13) 
Ri == 
A B — 
ll B — 
FB 
ET 
Èl B — 


1971 


1 


| Sr 


ap | 


| a | 


Landschaft 
D 3 À 
+ 
Ae 4 
ale) 
+ + 
ne 
Lia 
de, fee 
244 
ze 
a 
un 
EO - 
Lia 
Li. 
er 
ae 
+ 
di: 
di 
Liu 
= = 
> + + 
= + + 
+ 
pod 
+ + 
Li + oe 
Aah ETS 
++ 
Li 


B. THEOWALD: Die Tipuliden der Benelux-Länder 


montium Egger, 1863 (= pseudolateralis Tonnoir 
in Goetghebuer & Tonnoir, 1921) 
*pruinosa Wiedemann, 1817 
guadrivittata Staeger, 1840 
solstitialis Westhoff, 1881 (= pierre: Tonnoir in 
Goetghebuer & Tonnoir, 1921) 
Subg. Acutipula Alexander, 1924 
fulvipennis Degeer, 1776 
luna Westhoff, 1881 
maxima Poda, 1761 
vittata Meigen, 1804 
Subg. Platytipula Matsumura, 1916 
luteipennis Meigen, 1830 
melanoceros Schummel, 1833 
Subg. Nigrotipula Hutson & Vane-Wright, 1969 (= 
Anomaloptera Lioy, 1863, nec Perris, 1843) 
nigra Linnaeus, 1758 
Subg. Schummelia Edwards, 1931 
variicornis (Schummel, 1833) 
*yerburyi Edwards, 1924 
*zernyi Mannheims, 1952 
zonaria Goetghebuer in Goetghebuer & Tonnoir, 
1921 
Subg. Vestiplex Bezzi, 1924 
*excisa Schummel, 1833 
hortorum Linnaeus, 1758 (in: Goetghebuer & Ton- 
noir, 1921 als nubeculosa Meigen) 
*nubeculosa Meigen, 1804 (in: Goetghebuer & Ton- 
noir, 1921 als rubripes Schummel) 
*scripta Meigen, 1830 
Subg. Lunatipula Edwards, 1931 
“alpina Loew, 1873 
brunneinervis Pierre, 1921 
bullata Loew, 1873 
cava Riedel, 1913 
dilatata Schummel, 1833 
*fascipennis Meigen, 1818 
“helvola Loew, 1873 
“limitata Schummel, 1833 
“livida Van der Wulp, 1858 
“longidens Strobl, 1909 
“lunata Linnaeus, 1758 
*magnicauda Strobl, 1895 
peliostigma Schummel, 1833 
*pustulata Pierre, 1920 
selene Meigen, 1830 
vernalis Meigen, 1804 
Subg. Mediotipula Pierre, 1924 
“bidens Bergroth, 1888 
*stigmatella Schummel, 1833 
Subg. Savishenkia Mannheims, 1962 
*alpium Bergroth, 1888 
*benesignata Mannheims, 1954 
“limbata Zetterstedt, 1838 


221 

Land Landschaft 
FIEBER ew LD 
lol, de) — — du. A 
HBL | + + + + 
ER Ze en EN SA 
EB ue ire 
H B — Jd le 
pi E + + + + 
HUE — +++ 
H B — + + + + 
END ee 
Jal 38}. — — + = 
HB $+ + 
ar = + + + + 
jet 3 = — + . — 
= pu AREE di 
MB oe la Ale ae 
Fie DE Sr ely es 
HB — iv te dE 
ist i I Ste I 
H BL /++++ + 
SS Bie Sage ae DST Er 
Be ERA = 
e BERN, EE DA NEE 
Ey ay de po dey ce 
Et ey — N ng 
at ee A 
Et IB = aS) dE — 
AE ek BE > am 
Ei Bel a se — 
A Re den TR) di LU EE 
Hy Ba rar ar 

B — . — 
Hi By — — 

B pe 
iets} = SEEN a 
HB fet Hohn 
rite hen CR ES 
eA en KAREL I go 
HB REA 
Ba Sion Ig 
ER A Beery eee TEE 


222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 


grisescens Zetterstedt, 1851 (= macrocera Zetterstedt, 
1851, nec Say, 1823) 
marmorata Meigen, 1818 
obsoleta Meigen, 1818 
pagana Meigen, 1818 
rufina Meigen, 1818 
*signata Staeger, 1840 
staegeri Nielsen, 1922 
subnodicornis Zetterstedt, 1838 
*subvafra Lackschewitz, 1935 
Subg. Prerelachisus Rondani, 1842 
hortensis Meigen, 1818 
irrorata Macquart, 1826 
*meigeni Mannheims, 1966 (= hortulana Meigen, 
1818, nec Linnaeus, 1758) 
neurotica Mannheims, 1966 (= nervosa Meigen, 
1818, nec Schrank, 1803) 
pabulina Meigen, 1818 
pseudoirrorata Goetghebuer in Goetghebuer & Ton- 
noir, 1921 
*pseudovariipennis Czizek, 1912 
truncorum Meigen, 1830 
varipennis Meigen, 1818 
*winthemi Lackschewitz, 1932 
Subg. Odonatisca Savtshenko, 1956 
juncea Meigen, 1818 
Subg. Dendrotipula Savtshenko, 1964 
*flavolineata Meigen, 1804 
Subg. Beringotipula Savtshenko, 1961 
unca Wiedemann, 1817 


Cylindrotominae 


Cylindrotoma Macquart, 1834 
distinctissima (Wiedemann in Meigen, 1818) 
Diogma Edwards, 1938 
glabrata (Meigen, 1818) 
Phalacrocera Schiner, 1863 
replicata (Linnaeus, 1758) 
Triogma Schiner, 1863 
trisulcata (Schummel, 1829) 


Limoniinae 


Limonia Meigen, 1803 
Subg. Metalimnobia Matsumura, 1911 
bifasciata (Schrank, 1781) 
“quadrimaculata (Linnaeus, 1761) 
guadrinotata (Meigen, 1818) 
“zetterstedti Tjeder, 1968 (= elegans Zetterstedt, 
1838, nec Wiedemann, 1830) 
Subg. Limonia Meigen, 1803 
dilutior Edwards, 1921 
“flavipes (Fabricius, 1787) 
hercegovinae Strobl, 1898 
macrostigma (Schummel, 1829) 


a 
ee) 


TE EE El EEEN) 


| 


| xy ee | 


Ge oe or 


EE La NE 


LL XL 
WW 


Gh HD 
w | ew | 


Land 


œ © © © © id 


Wu 


oJ 


L 


Is. 


1971 


1 


sr | 


| 


dee 


Landschaft 
DES 4 
+ + 
each 
a 
SE 
heed el 
+ + 
by 
zake: 
+ + + 
AL 
+ + + 
FRE 
+ + 
a 
+ + + 
HUE + 
+ sam 
Lei 
+ + 


ia 


a= ial 


B. THEOWALD: Die Tipuliden der Benelux-Länder 223 


IL 


An 


Landschaft 
2 4 


E 
= 


*maculicosta (Coquillet, 1905) 
nigropunctata (Schummel, 1829) 
nubeculosa (Meigen, 1818) 
“stigma (Meigen, 1818) 
“sylvicola (Schummel, 1829) 
*taurica (Strobl, 1894) 
*tripunctata (Fabricius, 1781) 
*trivittata (Schummel, 1829) 
Subg. Neolimonia Alexander, 1964 
*dumetorum (Meigen, 1818) 
Subg. Dicranomyia Stephens, 1829 
*aquosa Verrall, 1886 
autumnalis (Staeger, 1840) 
chorea (Meigen, 1818) 
*danica Kuntze, 1919 
didyma (Meigen, 1818) 
“distendens Lundström, 1912 
“frontalis (Staeger, 1840) (= ostensackeni Westhoff, 
1882). 
“fusca (Meigen, 1818) (= pilipennis Egger, 1863) 
“lucida De Meijere, 1919 
luteipennis Goetghebuer in Goetghebuer & Tonnoir, 
1920 
*melleicauda complicata De Meijere, 1919 
*mitis (Meigen, 1830) 
*modesta (Meigen, 1818) 
*omissinervis De Meijere, 1919 
*ornata (Meigen, 1818) 
sera (Walker, 1848) 
“sericata (Meigen, 1830) 
*stigmatica (Meigen, 1830) 
*tristis (Schummel, 1829) 
*ventralis (Schummel, 1829) 
Subg. Melanolimonia Alexander, 1965 
morio (Fabricius, 1787) 
Subg. Achyrolimonia Alexander, 1965 
*decemmaculata (Loew, 1873) 
Subg. Atypophthalmus Brunetti, 1911 
*inusta (Meigen, 1818) 
Subg. Rhipidia Meigen, 1818 
*ctenophora (Loew, 1871) 
duplicata (Doane, 1900) (= maculata Meigen, 1818, 
nec Meigen, 1804) 
*yniseriata Schiner, 1864 
Elliptera Schiner, 1863 
*omissa Egger, 1863 — M piani 
Dicranoptycha Osten-Sacken, 1859 
“fuscescens (Schummel, 1829) (= cinerascens sensu) 
De Meijere et Goetghebuer & Tonnoir, nec Meigen, 
1818) Ei Bi SEEN SUN Se 
Taphrophila Rondani, 1856 (= Antocha Osten-Sacken, 
1859) 
vitripennis (Meigen, 1830) EB MM 
Thaumastoptera Mik, 1866 
calceata Mik, 1866 Bia “a ia LE er 


i 
a 


TETE 


oolite Mees MeMo: Me: Mee: N=; 
| 
| 
I 
- ++] 


E. 


co 


io] 
> 
L 


vel el » 
| 
+4 
le 


oel fool fae) Jaffa (ue Yao) fax] | 
Bow 
| 
| 
| - 
ae 
=] 
==] 


foo) facflacfac} tao} fact | 


aal 
deal 
| 


wouwvel elen tx 


mom mul 
lov) 
| 
_ 
- 
| 


| 
œ 
| 
| 
| 
| 
| 


EE & 
[vo] 
| 
GE 
TE 
a 
Est 
Ei: 


224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 


Helius Lepeletier & Serville iz Latreille et al., 1828 
(= Megarhina Lepeletier & Serville im Latreille 
et al., 1828 — Rhamphidia Meigen, 1830) 
flavus Walker, 1856 
longirostris (Wiedemann in Meigen, 1818) 


Ula Haliday, 1833 
*mollissima Haliday, 1833 (= crassicauda Agrell, 
1945) 
“sylvatica (Meigen, 1818) (= macroptera (Macquart, 
1826) ) 
Pedicia Latreille, 1809 
Subg. Pedicia Latreille, 1809 
rivosa (Linnaeus, 1758) 
Subg. Crunobia Kolenati, 1860 
“littoralis (Meigen, 1818) (= inconstans Osten-Sacken, 
1859, sensu Goetghebuer & Tonnoir, 1921) 
*straminea (Meigen, 1818) (= schineri (Kolenati, 
1860) ) 
Subg. Tricyphona Zetterstedt, 1837 
immaculata (Meigen, 1818) 
*schummeli Edwards, 1921 
*unicolor (Schummel, 1829) 
Subg. Ludicia Hutson & Vane-Wright, 1969 
“claripennis (Verrall, 1888) 
*]ucidipennis (Edwards, 1921) 
Subg. Amalopis Haliday, 1856 
occulta (Meigen, 1830) (= gmundensis Egger, 1863) 
Dicranota Zetterstedt, 1838 
Subgenus Dicranota Zetterstedt, 1838 
bimaculata (Schummel, 1829) 
Subg. Paradicranota Alexander, 1934 
brevitarsis Bergroth, 1890 
gracilipes Wahlgren, 1905 
*pavida (Haliday, 1833) (=Jongitarsis Bergroth, 1891) 
“subtilis Loew, 1871 


Dactylolabis Osten-Sacken, 1859 
*sexmaculata Macquart, 1846 
transversa (Meigen, 1818) (= gracilipes Loew, 1869) 

Epiphragma Osten-Sacken, 1859 
“ocellaris (Linnaeus), 1761 

Austrolimnophila Alexander, 1920 
“ochracea (Meigen, 1818) 

Pseudolimnophila Alexander, 1919 
*lucorum (Meigen, 1818) 
sepinm (Verrall, 1886) 

Limnophila Macquart, 1834 

Subg. Estonia Van der Wulp, 1874 
barbipes (Meigen, 1818) 

Subg. Limnophila Macquart, 1834 
“angustipennis (Meigen, 1818) 
*arnoudi spec. nov. 
pictipennis (Meigen, 1818) 
punctata Schrank, 1781 


Land 

H B 
Hi 
H B 

H B 

B 

H B 

igi 18} 

H B 

jet 18 

H B 

H B 

HB 

— B 
— B 
H B 

— B 
— B 
H B 

H B 

— B 
— B 
H B 

H B 

EB 

EIB 

H B 

H B 

H B 

H B 

H B 


6, 


1971 


1 


+ | 


Landschaft 
2 3 4 
rd 
+ + + 
— + + 
— — + 
+ + + 
— — + 
— + 
nce 
++ + 
+++ 
st 
+++ 
+ + — 
+ + 
EL 
+ + 


at | 


+++ + 


+ 


B. THEOWALD : Die Tipuliden der Benelux-Länder 


Subg. Idioptera Macquart, 1834 
fasciata (Linnaeus, 1767) 
*pulchella (Meigen, 1830) 
Subg. Eloeophila Rondani, 1856 
maculata (Meigen, 1804) (= marmorata (Meigen, 
1818) ) 
*mundata Loew, 1871 
*submarmorata Verrall, 1887 
*trimaculata (Zetterstedt, 1838) (in: Goetghebuer & 
Tonnoir, 1921, als Idioptera trimaculata Zett.) 
*verralli Bergroth, 1912 (= dalei Edwards in De 
Meijere, 1921) 
Subg. Philidorea Bigot, 1854 
*abdominalis Staeger, 1840 
“aperta Verrall, 1887 
“bicolor (Meigen, 1818) 
dispar (Meigen, 1818) 
ferruginea (Meigen, 1818) 
fulvonervosa (Schummel, 1829) (= lineonella Ver- 
rall, 1887) 
*glabricula (Meigen, 1830) 
“heterogyna Bergroth, 1913 
lineola (Meigen, 1818) 
*meigeni Verrall, 1887 
*phaeostigma (Schummel, 1829) 
*squalens Zetterstedt, 1838 
Subg. Brachylimnophila Alexander, 1966 
adjuncta (Walker, 1848) (= dimidiata De Meijere, 
1918) 
*nemoralis (Meigen, 1818) 
Subg. Neolimnomyia Séguy, 1937 
*batava Edwards, 1938 (= leucophaea De Meijere, 
1921, nec Meigen, 1818) 
filata Walker, 1856 
Subg. Adelphomyia Bergroth, 1890 
“punctum (Meigen, 1818) 
Pilaria Sintenis, 1889 
discicollis (Meigen, 1818) 
fuscipennis (Meigen, 1818) 
*platyptera (Macquart, 1826) 
scutellata (Staeger, 1840) (= subtincta Zetterstedt, 
1851) 
Hexatoma Latreille, 1809 
“bicolor (Meigen, 1818) 
*gaedii (Meigen, 1830) (= saxonum Loew, 1867) 
*nubeculosa Burmeister, 1829 
“vittata (Meigen, 1830) 
Paradelphomyia Alexander, 1936 
Subg. Oxyrhiza De Meijere, 1946 
*nielseni Kuntze, 1919 
selenis (Haliday, 1833) (in: Goetghebuer & Tonnoir: 
Adelphomyia selenis Hal.) 


Gonomyia Meigen, 1818 
Subg. Ellipteroides Becker, 1907 


TT XE 


EAU EEK sa fac} tao} foe) | fact aol Goch | EEE EEE Erle fac! fac} [| fa 


AR | tach tact dad ao) ar) 


EG 


225 
Land Landschaft 
Bl D ASS 
B mee Wee 
B b — + 
B = 4 
ee 
Br er: 
B= N eme 
B — ae ale = 
Bi Lette dI ee 
B — dd Ore 
B — Le 
B — + 4 4 4 
19, = ie no He 
Be Sans A 
BU heem) eN 
Bee = 5 { 
o he a 
B — — ale 
Bi = sees sine 
Bye eee ee 
Bf et eee 
BI E oe ye 
B — EE 
(ae Ne CARRE 
B — SS CN 
B — — + — — 
geen gen p 
A, pad 
Bae 
Bi ar == 


226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 


*alboscutellata Von Roser, 1840 
lateralis Macquart, 1835 
Subg. Lipophleps Bergroth, 1915 
abbreviata Loew, 1873 
Subg. Gonomyia Meigen, 1818 
*bifida Tonnoir in Goetghebuer & Tonnoir, 1920 
dentata De Meijere, 1920 (= incisurata Tonnoir in 
Goetghebuer & Tonnoir, 1920) 
*lucidula De Meijere, 1920 
“recta Tonnoir in Goetghebuer & Tonnoir, 1920 
simplex Tonnoir in Goetghebuer & Tonnoir, 1920 
tenella (Meigen, 1818) 
Neolimnophila Alexander, 1920 
carteri Tonnoir in Goetghebuer & Tonnoir, 1920 
placida (Meigen, 1830) 
Crypteria Bergroth, 1913 
limnophiloides Bergroth, 1913 
Lipsothrix Loew, 1873 
“errans (Walker, 1848) 
*remota (Walker, 1848) (in: Goetghebuer & Tonnoir 
als Limnophila clara Tonnoir) 
Gnophomyia Osten-Sacken, 1859 
lugubris (Zetterstedt, 1838) (= viridipennis Gim- 
merthal, 1847 = tripudians Bergroth, 1891) 
Rhabdomastix Skuse, 1890 
Subg. Secandaga Alexander, 1911 
laeta (Loew, 1873) 
Cheilotrichia Rossi, 1848 
Subg. Cheilotrichia Rossi, 1848 
imbuta Meigen, 1818 
Subg. Platytoma Lioy, 1863 
cinerascens (Meigen, 1818) (= nubila Schummel, 
1829) 
Subg. Gonempeda Alexander, 1924 
flava (Schummel, 1829) 
Erioptera Meigen, 1803 
Subg. Erioptera Meigen, 1803 
*divisa (Walker, 1848) (= macrophthalma Loew, 
1871) 
fuscipennis Meigen, 1818 
“fusculenta Edwards, 1938 
gemina Tjeder, 1967 (flavescens Linnaeus sensu 
Meigen, 1818, nec Linnaeus, 1758) 
griseipennis Meigen, 1838 
“limbata Loew, 1873 
“lutea Meigen, 1818 
*meijeri Edwards, 1921 
*minor De Meijere, 1920 
nielseni De Meijere, 1921 
*nigripalpis Goetghebuer in Goetghebuer & Tonnoir, 
1920 
squalida Loew, 1871 
trivialis Meigen, 1818 
Subg. Trimicra Osten-Sacken, 1861 
pilipes (Fabricius, 1887) 


TE EEL 
wo 


fol fool fact lac) || faci fark fan) fado arl 


re a | 


Land 


B 
B 
B 


ier) 


IL 


1971 


Landschaft 
2 3 4 
ae 
= + 
JE 
+ + 
en 
+ + 
ni 
Li 
LE 
dee 
we 
+ + 


5 


B. THEOWALD: Die Tipuliden der Benelux-Länder 


Subg. Symplecta Meigen, 1830 
hybrida (Meigen, 1804) (= punctipennis (Meigen, 
1818) ) 
Subg. Psiloconopa Zetterstedt, 1830 (= Acyphona 
Osten-Sacken, 1869). 
areolata Siebke, 1872 
maculata (Meigen, 1804) 

“occoecata Edwards, 1936 
stictica (Meigen, 1818) 
vicina (Tonnoir in Goetghebuer & Tonnoir, 1920) 

Subg. Arctoconopa Alexander, 1955 
*melampodia (Loew, 1873) 
Chionea Dalman, 1816 

“lutescens Lundström, 1907 

Ormosia Rondani, 1856 
Subg. Ormosia Rondani, 1856 

“albitibia Edwards, 1921 

“bicornis (De Meijere, 1920) 

“clavata (Tonnoir iz Goetghebuer & Tonnoir, 1920) 
*depilata Edwards, 1938 (—hederae Curtis sensu 

De Meijere, 1920, nec Curtis, 1835) 

hederae (Curtis, 1835) (= uncinata De Meijere, 1920) 
lineata (Meigen, 1904) 
murina (Goetghebuer in Goetghebuer & Tonnoir, 

1920) 
nodulosa (Macquart, 1826 sensu De Meijere, 1920) 
pseudosimilis Lundström, 1906 
*staegerina Alexander, 1953 (= similis Schummel 
sensu Staeger, 1840, nec Schummel, 1829) 
Subg. Rhypholophus Kolenati, 1860 
bifurcata Goetghebuer in Goetghebuer & Tonnoir, 
1920 

“bivittata Loew, 1873 

“haemorrhoidalis Zetterstedt, 1838 
varia (Meigen, 1818) 

Subg. Scleroprocta Edwards, 1938 

*pentagonalis (Loew, 1873) (= sororcula Zetterstedt 
sensu auct. nec Zetterstedt, 1851 — fascipennis 
sensu auct. nec Zetterstedt, 1838) 

*sororcula (Zetterstedt, 1851) (—danica Nielsen, 1925) 

Molophilus Curtis, 1833 

Subg. Molophilus Curtis, 1833 

*appendiculatus (Staeger, 1840) (= armatus De Meij- 
ere, 1920) 

“ater (Meigen, 1804) 

*bifidus Goetghebuer in Goetghebuer & Tonnoir, 1920 

*bibamatus De Meijere, 1918 
cinereifrons De Meijere, 1920 
corniger De Meijere, 1920 
“curvatus Tonnoir in Goetghebuer & Tonnoir, 1920 
*czizeki Lackschewitz, 1939 (1940) 

*flavus Goetghebuer in Goetghebuer & Tonnoir, 1920 
griseus (Meigen, 1818) (= bifilatus Verrall, 1886) 
medins De Meijere, 1918 
niger Goetghebuer in Goetghebuer & Tonnoir, 1920 


Land Landschaft 
H BL RE 
Hy B rose 
H B Sb eee 
EB Ir 
H B lain Wiel 
lel 13} ss ai 

B == 
H B = 

B ES 
H B 

B 
EIB =e ate 
H B + È 
Hy B aa Uy aire 
EB ee 
Hen Fate az 
H B — 

B un 

B 
Te = = Br 
H B ot 
HB ASE 
H B — — 
TT EE a 
HE stain 
H B 4 
H B — — 
H B ee 
H B MEET 
H B — . 
H B — — 
ne 5) BE QE 
H B 
FNB Sai 
H B STE 
eB = i 


228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 1971 


Land Landschaft 
H B L N NS 
obscurus (Meigen, 1818) H B — . ++ + 
“occultus De Meijere, 1818 (= falciger Goetghebuer 
in Goetghebuer & Tonnoir, 1920) H B — — — . + 
ochraceus (Meigen, 1818) (= appendiculatus Ver- 
rall, sensu De Meijere nec Staeger, 1840) H B — 
ochrescens Edwards, 1938 (= ochraceus De Meijere — + + — 
et Goetghebuer & Tonnoir, nec Meigen, 1818) H B — h == 
pleuralis De Meijere, 1920 H B — — + — 
propinquus Egger, 1863 (= gladius De Meijere, 
1920) H B — + + 
serpentiger Edwards, 1938 (=propinguus De Meijere 
et Goetghebuer & Tonnoir, nec Egger, 1863) H B — — 5 a 
undulatus Tonnoir in Goetghebuer & Tonnoir, 1920 B al as 
Tasiocera Skuse, 1890 
Subg. Dasymolophilus Goetghebuer in Goetghebuer & 
Tonnoir, 1920 
murina (Meigen, 1818) H B — . + 


BEMERKUNGEN UBER EINZELNE GATTUNGEN UND ARTEN 


Ctenophora guttata Wiedemann 
Holland: nur zwei Exemplare bekannt (Delft, Friedhof Jaffa, V.1866, Piaget; und 
Breda, V. (etwa 1900, Heylaerts). 
C. ornata Wiedemann 
Holland: Umgebung Arnheim. 
Erstnachweis für Belgien: 2 G Buzenol, 15.VI.1964, De Laever. 
Nephrotoma analis (Schummel) 
Erstnachweis für Luxemburg: Mondorf und Bech-Kleinmacher, Juni 1970, Theowald. 
N. flavescens (Linnaeus) 
Erstnachweis für Luxemburg: Mondorf und Bech-Kleinmacher, Juni 1970, Theowald. 
N. lindneri (Mannheims) 
Erstnachweis für Belgien: 1 ® La Panne, 17.V.1932, Goetghebuer. Das Exemplar 
steckte in der Sammlung Goetghebuer (Brüssel) unter pratensis (Linnaeus). 
N. quadrifarta (Meigen) 
Erstnachweis für Luxemburg: 8 &, 3 9 Mondorf, 29.V1.1970, Theowald. 
N. quadristriata (Schummel) 
Erstnachweis für Belgien: 1 & Coxyde-Bains, 14.VI. 
N. scalaris (Wiedemann) 
Alle von Goetghebuer & Tonnoir als scalaris aufgeführten Exemplare sind flavipalpis 
(Meigen). 
N. scurra (Meigen) 
Erstnachweis für Luxemburg: 1 9 Mondorf, 29.VI.1970, Theowald. 
N. submaculosa Edwards 
De Laever & Mannheims (1958) führten diese Art erstmalig auf für Belgien 
(Sclessin, Liége, Bois de la Vecquée). Ich sah noch Exemplare von Archennes, 
Coxyde-Bains, Coxyde-St. Idesbald, Knocke und La Gileppe. Es ist auch in Belgien 
wohl eine viel vorkommende Art. 
Prionocera pubescens Loew 
De Laever & Mannheims (1958) führten diese Art erstmalig auf für Belgien (Vence). 


B. THEOWALD: Die Tipuliden der Benelux-Länder 229 


Im Museum Brüssel sah ich noch 2 & Postel, 11.VI.1923, G. Severin. 
Tipula Linnaeus 
Wir folgen der neuen Einteilung in Untergattungen vorgeschlagen von Savtshenko 
(1961), fügen aber die Untergattungen Péerelachisus und Oreomyza zusammen 
(Alexander, 1965; Theowald, 1957a). 
T. (Tipula) oleracea Linnaeus 
Goetghebuer & Tonnoir (1921) stellen palwdosa Meigen als Synonym zu oleracea 
Linnaeus. In der Sammlung Brüssel stehen sie aber als zwei gute Arten aufgestellt. 
T. (Tipula) paludosa Meigen 
Erstnachweis für Belgien: wie zu erwarten war, sah ich diese Art von vielen Fund- 
orten. 
T. (Yamatotipula) coerulescens Lackschewitz 
Erstnachweis fiir Belgien: Hautes Fagnes, Sourbrodt, 24—26.V.1957, De Laever. 
T. (Yamatotipula) pruinosa Wiedemann 
Erstnachweis für Luxemburg: 1 & Mondorf, 28.VI.1970, Theowald. 
T. (Schummelia) yerburyi Edwards 
Erstnachweis fiir Belgien: Mirwart. 
Ich sah mehr als 20 Exemplare dieser Art aus Belgien. Sie wurden 1950—1955 ge- 
sammelt in der Umgegend von Mirwart. 
T. (Schummelia) zernyi Mannheims 
Erstnachweis für Belgien: 1 4 Buzenol, Vallée de Claireau, 31.VII.1919. A. Tonnoir. 
Dieses Exemplar stand in der Sammlung Goetghebuer, Brüssel, unter dem Namen 
yerburyi Edwards. 
T. (Vestiplex) excisa Schummel 
Erstnachweis für Holland: 1 © Kerkrade, Amseldal, 1.1X.1965, Br. Arnoud. Diese 
boreo-alpine Art war bisher nur aus den Alpen und aus Lappland bekannt. Sie wurde 
in Holland erbeutet in einem ziemlich kalten und feuchten Tal auf 150 m. Unter 
den mehr als 4000 Tipuliden die ich von den Ardennen und den Hautes Fagnes 
sah, war diese Art nicht vertreten. 
Schummel hat diese Art beschrieben von „Um Breslau”, d.h. auf ungefähr derselben 
Höhe. Theowald & Mannheims (1962) nahmen an, daß es sich bei der Fundortangabe 
Schummels wahrscheinlich um einen Irrtum handelt. Der Fund in Holland weist aber 
darauf hin, daß es vielleicht doch Restpopulationen dieser Art im europäischen Flach- 
land gibt. 
T. (Vestiplex) nubeculosa Meigen 
Erstnachweis für Luxemburg: 1 © Mondorf, 27.VI.1970, Theowald. 
T. (Vestiplex) scripta Meigen 
Erstnachweis für Luxemburg: 4 g', 4 9 Mondorf, 27.VI.1970, Theowald. 
T. (Lunatipula) alpina Loew 
Erstnachweis für Belgien: 2 & Torgny, 29.VI.1958, De Laever. 
T. (Lunatipula) fascipennis Meigen 
Erstnachweis für Luxemburg: 1 g', 2 2 Bech-Kleinmacher, 30.VI.1970, Theowald. 
T. (Lunatipula) helvola Loew 
Erstnachweis für Belgien: 2 g', 1 9 Han-sur-Lesse, 17.VII.1957, De Laever. 
T. (Lunatipula) limitata Schüummel 
Erstnachweis fiir Belgien: Mirwart (in Museum Briissel), Bihain und Wavreille (De 
Laever). 


230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 1971 


T. (Lunatipula) livida Van der Wulp 
Erstnachweis für Luxemburg: 3 &, 3 9 Mondorf, 27—29.V1.1970, Theowald. 

T. (Lunatipula) longidens Strobl 
Erstnachweis für Belgien: 1 & Wavreille, 9.VI.1958, De Laever. 

T. (Lunatipula) lunata Linnaeus 
Erstnachweis für Luxemburg: 7 &, 6 ® Mondorf, 27—29.VI.1970, Theowald. 

T. (Lunatipula) magnicauda Strobl 
Erstnachweis für Belgien: 1 & Han-sur-Lesse, 28.1V.1957, De Laever; 1 g' Wavreil- 
le, Belvaux, 11.V.1958, De Laever. 

T. (Lunatipula) pustulata Pierre 
Erstnachweis für Belgien: 1 g° Bihain, 22.IX.1962, De Laever. 

T. (Mediotipula) bidens Bergroth 
De Laever & Mannheims (1958) führten diese Art erstmalig auf für Belgien (Han- 
sur-Lesse). Ich sah weitere Exemplare von Spy-Bois, Onoz-Spy und Hastière. 

T. (Mediotipula) stigmatella Schummel 
Erstnachweis fiir Belgien. 
Im Museum Briissel steht ein Exemplar dieser Art ohne Fundort aus der Sammlung 
E. Candéze. Nach Horn & Kahle (1935) erhielt das Museum Briissel im Jahre 1899 
die Sammlung Candèze: „Elaterid., Lamellicorn., Lucanid., Longicorn. u. belg. Dipt.” 
Demnach stammt das Exemplar wohl aus Belgien und wahrscheinlich aus den Ar- 
dennen. 

T. (Savtshenkia) alpium Bergroth 
De Laever & Mannheims (1958) stellten erstmalig das Vorkommen dieser Art in 
Belgien fest (Membach, Lac-de-la-Gileppe). In Collection de la Belgique (Briissel) 
sah ich 5 als obsoleta Meigen bestimmte Stiicke von Hockai, Herzogenwald und 
Baraque Michel. In der Sammlung Goetghebuer sah ich noch ein als marmorata 
Meigen bestimmtes Exemplar von St. Hubert. 

T. (Savtshenkia) benesignata Mannheims 
Erstnachweis für Belgien: 1 g Hautes Fagnes, Duzo-Moupas, 10.IX.1948, R. Tollet. 

T. (Savtshenkia) limbata Zetterstedt 
Erstnachweis für Belgien: Bullange, 1.VIII—21.1X.1923, G. Severin. 

T. (Savtshenkia) signata Staeger 
Die in Goetghebuer & Tonnoir (1921) aufgeführten Exemplare sind alle staegeri 
Nielsen. 
Erstnachweis für Belgien: 1 9 Buzenol, 6.X.1962, De Laever. 
Theowald (1957b) führte diese Art erstmalig auch für Holland (Noorbeek). Seitdem 
wurde sie auch bekannt von Amsterdam. 

T. (Savtshenkia) subvafra Lackschewitz 
Erstnachweis für Belgien: 1 & Hockai, 28.1X.1950, J. Cooreman; 1 G', 4 2 Bihain, 
30.X.1965, De Laever. 

T. (Pterelachisus) meigeni Mannheims (= hortulana Meigen, nec Linnaeus) 
Fast alle in Goetghebuer & Tonnoir (1921) aufgeführten Exemplare von hortulana 
sind pseudovartipennis Czizek. Ich sah aber meigeni (= hortulana) von einer Reihe 
von Fundorten über ganz Belgien. 

T. (Pterelachisus) psendovariipennis Czizek. 
Siehe Bemerkung unter megen? Mannheims. 


B. THEOWALD: Die Tipuliden der Benelux-Länder 231 


T. (Pterelachisus) winthemi Lackschewitz 
Erstnachweis für Belgien: 1 4 Baillonville, Basse Adam, 9.V.1967, De Laever. 
T. (Dendrotipula) flavolineata Meigen 
Erstnachweis für Belgien: 1 © Mondorf, 27.VI.1970, Theowald. 
Limonia (Metalimnobia) quadrimaculata (Linnaeus) 
Erstnachweis für Holland: 1 & Kotten, 2.VII.1952, Theowald; 1 ® Noorbeek, 
27.V.1953, Br. Arnoud. 
Erstnachweis für Belgien: Forêt de Soignes, Bois de Ghlin, Mirwart, Onoz, Floren- 
ville. 
L. (Metalimnobia) zetterstedti Tjeder, 1968. 
Erstnachweis für Belgien: Han-sur-Lesse, Losheimergraben, Membach, St. Hubert, 
Vestreux und Cedrege, alle De Laever (nach Mannheims, in litt.). 
L. (Limonia) flavipes (Fabricius) 
Erstnachweis für Luxemburg: 10 &, 21 9 Mondorf, 27—30.VI.1970, Theowald. 
L. (Limonia) maculicosta (Coquillet, 1905) 
Erstnachweis für Belgien: 1 & Torgny, 17.VII.1934, De Laever (nach Mannheims, 
in litt). 
L. (Limonia) stigma (Meigen) 
Goetghebuer & Tonnoir kennen diese Art in Belgien nur von Virton. Ich sah noch 
Exemplare von Mirwart und Malmedy. 
L. (Limonia) sylvicola (Schummel) 
In Belgien noch immer nur bekannt von Virton. 
L. (Limonia) taurica (Strobl) 
Erstnachweis für Belgien: 1 9 Seraing s. Meuse, 12.V1.1955, De Laever (nach Mann- 
heims, in litt.). 
L. (Limonia) tripunctata (Fabricius) 
Erstnachweis fiir Luxemburg: Mondorf und Bech-Kleinmacher, Juni 1970, Theowald. 
L. (Limonia) trivittata (Schummel) 
War in Belgien nur bekannt von Genval. Ich sah weitere Exemplare von Pepinster, 
Forêt de Soignes, Huccorgne und Bossière. 
Erstnachweis fiir Luxemburg: Mondorf und Bech-Kleinmacher, Juni 1970, Theowald. 
L. (Neolimonia) dumetorum (Meigen) 
Erstnachweis für Luxemburg: 1 & Mondorf, 27.VI.1970, Theowald. 
L. (Dicranomyia) aquosa Verrall 
In Belgien noch immer nur bekannt von Denée. 
Leef Dicranomyia ) danica Kuntze 
De Meijere (1928) führt diese Art auf für Amsterdam. Bis heute sind ín Holland 
keine weiteren Fundorte bekannt. | 
L. (Dicranomyia) distendens Lundstròm 
In Belgien noch immer nur bekannt von Hockai. 
L. (Dicranomyia) frontalis (Staeger) 
In Belgien noch immer nur bekannt von Heide. 
L. (Dicranomyia) fusca (Meigen) 
Diese Art war in Holland nur bekannt von Ootmarsum. Ich sah weitere Exemplare 
von Herkenbosch, Eygelshoven, Bunde und Ravensbos. 
L. (Dicranomyia) lucida De Meijere 
Diese Art war in Holland nur bekannt von Bunde. Ich sah weitere Exemplare von 
Heerlen, Kerkrade, Eygelshoven und Eys-Wittem. 


232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 1971 


L. (Dicranomyia) melleicauda complicata De Meijere 
Nach Mannheims (in litt.) ist complicata De Meijere, 1919, eine Unterart der neark- 
tischen mellezcauda Alexander, 1917. 
Die Unterart wurde beschrieben nach Exemplaren von Diemen. Bis heute sind in 
Holland keine weiteren Fundorte bekannt. 
L. (Dicranomyta) mitis (Meigen) 
Diese Art war in Belgien nur bekannt von Hockai. Ich sah Exemplare von mehr als 
15 weiteren Fundorten. 
Erstnachweis für Luxemburg: 4 &, 1 © Mondorf, 27.VII.1970, Theowald. 
L. (Dicranomyia) modesta (Meigen) 
Erstnachweis fiir Luxemburg: Mondorf und Bech-Kleinmacher, Juni 1970, Theowald. 
L. (Dicranomyta) omissinervis De Meijere 
Die Art wurde beschrieben nach einem Männchen von Zwammerdam. Ich sah weitere 
Exemplare von Holland: Brabantse Biesbosch 24, 6 9, 15—18.VI.1970 M. Peer- 
deman. 
L. (Dicranomyia) ornata (Meigen) 
Diese Art war in Holland bekannt von Houthem. Ich sah ein weiteres Exemplar 
von Gulpen. 
Erstnachweis für Belgien: 1 4 Houyet, 14.VII. 
L. (Dicranomyia) sericata (Meigen) 
In Belgien war diese Art nur bekannt von Falaën. Ich sah 1 & und 7 © von Sy. 
L. (Dicranomyia) stigmatica (Meigen) 
Goetghebuer & Tonnoir führen diese Art auf von Melle und Linkebeek, fügen aber 
hinzu: „Cette détermination n'est pas tout-a-fait sire, nos captures ne comportant que 
des 9 @”. Ich habe keine weiteren Exemplare dieser Art gesehen. 
L. (Dicranomyia) tristis (Schummel) 
In Belgien war diese Art nur bekannt von Virton. Ich sah keine weitere Exemplare. 
L. (Dicranomyta) ventralis (Schummel) 
In Belgien war diese Art nur bekannt von Falaën. Ich sah noch 1 9 von Postel. 
L. (Achyrolimonia) decemmaculata (Loew) 
Erstnachweis für Holland: 1 9 Neerijnen, 12.V.1953, C. de Jong; 1 © Amersfoort 
17.V1.1961, Theowald. 
Erstnachweis fiir Belgien: Ben-Ahin, Forét de Soignes, Reinhardstein. 
L. (Atypophthalmus) inusta (Meigen) 
Erstnachweis für Belgien: 1 & Falmignoul (Ravin du Colibri), 9.VII.1943, Collart. 
L. (Rhipidia) ctenophora (Loew) 
De Meijere (1921) kennt von dieser seltenen Art 1 4 aus Bussum. Sie wurde später 
in Holland nicht wieder gefunden. 
L. (Rhipidia) uniseriata Schiner 
De Meijere (1919) kennt 1 9 von Bodegraven. Bis heute sind von Holland keine 
weiteren Exemplare bekannt. 
Elliptera omissa Egger 
Erstnachweis für Belgien: Eprouve (grottes), Visé, Bomal. 
Dicranoptycha fuscescens (Schummel) 
De Meijere (1919) kennt 1 9 dieser Art aus Holland (Zutphen). Ich sah noch 1 & 
Schin-op-Geul, 1.V.1961, Gravestein. 
In Belgien ist diese Art nur bekannt von Virton. 


B. THEOWALD: Die Tipuliden der Benelux-Länder 233 


Ula mollissima Haliday 
Erstnachweis für Holland: 1 & Amsterdam, 23.IX.1960, J. Lourens; 1 & Amsterdam 
28.X.1960, K. J. N.; 1 9 Amsterdam, 13.VI.1962; 1 g', 2 Q Berg en Terblijt, 
Mussenputgroeve, 18.1.1962, Exc. Zool. Museum. 
Erstnachweis für Belgien: Hautes Fagnes, Duzo-Moupas; Forêt de Soignes; Mirwart; 
Bois de Cerfontaine. 
U. sylvatica (Meigen) 
Erstnachweis für Luxemburg: 1 9 Bech-Kleinmacher, 30.VI.1970, Theowald. 
Pedicia (Crunobia) littoralis (Meigen) 
De Meijere (1921) kannte diese Art aus Holland nur von Bunde. Ich sah weitere 
Exemplare von Epen, Mechelen, Holset und Vaals. 
P. (Crunobia) straminea (Meigen) 
Erstnachweis für Holland: 2 & Wageningen, IX, Coll. Geijskes. 
In Belgien war diese Art bekannt von Hockai. Ich sah weitere Exemplare von 
Francorchamps, Mirwart und Malmedy. 
P. (Trichyphona) schummeli Edwards 
De Meijere kennt diese Art aus Holland von Winterswyk. Das Beleg-Exemplar ist 
aber nicht mehr in seiner Sammlung vorhanden. Ich fand aber unter den unbestimm- 
ten Stücken noch 1 & Vlodrop (Limb.), 7.VI.1903, De Meyere. 
Ertsnachweis für Belgien: Hautes Fagnes, Chôdires; H.F., Duzo-Moupas; H.F., Nes- 
seloo; H.F., Baraque Michel; Herzogenwald. 
P. (Tricyphona) unicolor (Schummel) 
In Belgien war diese Art bekannt von Hockai. Ich sah weitere Exemplare von Hautes 
Fagnes, Longloup und H.F. Les Wez. 
P. (Ludicia) claripennis (Verrall) 
Erstnachweis für Holland: 1 & Holset, 13.VIII.1955, Br. Arnoud. 
P. (Ludicia) lucidipennis (Edwards) 
Erstnachweis für Belgien: 1 & Jemelle, 6.VIII. 
Dicranota (Paradicranota) brevitarsis Bergroth 
Erstnachweis für Belgien: 1 & Ben-Ahin, 18.VI; 1 9 Lesterny, 14.V. 
D. (Paradicranota) pavida (Haliday) 
In Holland war diese Art nur bekannt von Bunde. Ich sah weitere Exemplare von 
Terziet und Holset. 
Erstnachweis für Belgien: Cerfontaine, Mirwart, Forêt de Soignes. 
D. (Paradicranota) subtilis Loew 
In Holland war diese Art bekannt von Bunde. Ich sah weitere Exemplare von Heer- 
len und Eygelshoven. 
Erstnachweis fiir Belgien: Bevercé, Bois de Dave, Olloy de Berceau, Virton. 
Dactylolabis sexmaculata Macquart 
Erstnachweis für Belgien: Furfooz, Mazy, Herzogenwald, Mazy-Onoz, Thon. 
Epiphragma ocellaris (Linnaeus) 
Erstnachweis für Luxemburg: 2 g', 1 @ Mondorf, 27—29.VI.1970, Theowald. 
Austrolimnophila ochracea (Meigen) 
Erstnachweis für Luxemburg: 4 9 Mondorf, 29.V1.1970, Theowald. 
Pseudolimnophila lucorum (Meigen) 
Diese Art war in Belgien nur bekannt von dem Forét de Soignes. Ich sah weitere 
Exemplare von Poix St. Hubert, Falaén, Houvet, Mirwart und Malmedy. 


234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 1971 


Limnophila (Limnophila) angustipennis (Meigen) 
Erstnachweis für Belgien: 1 9 Lamerteau, 2.VIII. 


Limnophila (Limnophila) arnoudi spec. nov. (Abb. 1) 


Diese Art ist Limnophila (Limnophila) punctata (Schrank, 1781) tauschend ähnlich. 
Zwischen den Weibchen beider Arten finde ich keine Unterschiede. Vielleicht sind die 
Beine bei den Weibchen von arnoudi etwas robuster gebaut. Bei den Männchen gibt es 
nur deutliche Unterschiede im Bau des Hypopygs. 


Abbildung 1. 9. Tergit und Basistylusanhänge. a, Limnophila punctata (Schrank); b, Limnophila 
arnoudi spec. nov. 


Das 9. Tergit ist bei arnoudi in der Mitte fast nicht ausgeschnitten, bei punctata 
deutlich ausgeschnitten mit zwei an der Spitze geschwärzten Hinterrandfortsätzen. Die 
äußeren Basistylusanhänge sind bei arnoud? stumpf und am Ende stark geschwärzt, bei 
punctata spitz und nur am äußersten Ende der Spitze schwarz. Die inneren Basi- 
stylusanhänge tragen bei arnoudt einen Büschel von etwa acht langen schwachgebogenen 
Haaren, bei punctata sind immer nur vier kurze starkgebogene Haare vorhanden. 

Holotypus: & Holland, Streythagerbeek, 18.V.1963, Br. Arnoud leg. 

Paratypen: Holland, 1 g', dieselben Data wie der Holotypus; Eygelshoven, 2 g', 20.V. 
1966; 1 d', 26.V.1967; 1 &, 19.V.1968; 1 dg, 31.V.1968; Noorbeek, 1 d, 27.V. 
1958, alle Br. Arnoud leg. Holland, Bunde, 4 &, 26.V.1951, Theowald leg. Belgien, 
Tervueren, 1 4 (ohne weitere Data). 

Geografische Verbreitung: Ich sah Exemplare aus Holland (Südlimburg) und 
Belgien (Umgebung Brüssel). Mannheims (in litt.) sah Exemplare aus dem Sieben- 
gebirge (südöstlich von Bonn) und Savtshenko (in litt.) fand sie in den Karpaten. 
L. (Idioptera) pulchella (Meigen) 

Diese Art war in Belgien nur bekannt von Baraque Michel. Ich sah weitere Exem- 

plare von Hockai und Rymenam. 

L. (Eloeophila) mundata Loew 
In Belgien bekannt von Virton. Ich sah keine weiteren Exemplare dieser Art. 


B. THEOWALD: Die Tipuliden der Benelux-Länder 235 


L. (Eloeophila) submarmorata Verrall. 
Erstnachweis fiir Holland: Eygelshoven, Mechelen, Heerlen, Noorbeek. 
L. (Eloeophila) trimaculata (Zetterstedt) 
Erstnachweis fiir Holland: 2 &, 1 9 Wageningen, 30.1V.1936, Geyskes. 
In Belgien war diese Art bekannt von Hockai. Ich sah weitere Exemplare von Mal- 
médy und Nonceveux. 
L. (Eloeophila) verralli Bergroth 
In Holland war diese Art bekannt von Amersfoort. Ich sah keine weiteren Exemplare 
dieser Art. 
L. (Philidorea) abdominalis Staeger 
In Belgien war diese Art bekannt von Eecloo. Ich sah keine weiteren Exemplare 
dieser Art. 
L. (Philidorea) aperta Verrall 
Erstnachweis für Belgien: 1 Q Mirwart, 6.VII. 
L. (Philidorea) bicolor (Meigen) 
In Belgien ist diese Art nur bekannt von Hockai. 
L. (Philidorea) glabricula (Meigen) 
Erstnachweis für Holland: 1 & Eygelshoven, 21.V11.1954, Br. Arnoud. 
L. (Philidorea) heterogyna Bergroth 
Erstnachweis für Belgien: 1 9 Postel, VII. 
L. (Philidorea) meigeni Verrall 
Diese Art wurde von De Meijere mit phaeostigma (Schummel) verwirrt. 
Erstnachweis für Holland: Hilversum, Vlodrop, Amersfoort, Duurswoude. 
L. (Philidorea) phaeostigma (Schummel) 
In Holland nur bekannt von Brunssum und Bennekom. Die Angaben von De Meijere 
sind meigeni Verrall. 
Erstnachweis für Belgien: Hockai; Bihain; Samrée; Haute Fagne, Noir Flohay; H.F., 
Duzo-Moupas; H.F., Baraque Michel; H.F., Fraineux; Calmpthout. 
L. (Philidorea) squalens Zetterstedt 
Erstnachweis fiir Belgien: Haute Fagne, Chodires ; H.F., Hoegne; H.F., Duzo-Mou- 
pas; H.F., Baraque Michel; Hockai, Bihain, Postel, Mirwart. 
L. (Brachylimnophila) nemoralis (Meigen) 
Erstnachweis für Luxemburg: 2 & Mondorf, 29.VI.1970, Theowald. 
L. (Neolimnomyia) batava Edwards 
Erstnachweis für Luxemburg: 1 4 Mondorf, 27.VI.1970, Theowald. 
L. (Adelphomyia) punctum (Meigen) 
Diese Art war in Holland nur von Denekamp bekannt. Ich sah weitere Exemplare 
von Heerlen, Strijthagen und Eygelshoven. 
Pilaria platyptera (Macquart) 
Erstnachweis für Belgien: Bois de Couvin, Chimay, Oignies, Cerfontaine, Furfooz. 
Hexatoma bicolor (Meigen) 
Diese Art war in Belgien bekannt von Hockai. Ich sah weitere Exemplare von Mal- 
medy, Bois de Couvin und Mirwart. 
H. gaedi (Meigen) 
Van den Brandt erbeutete vor etwa hundert Jahren einmal ein Weibchen in der Nahe 
von Venlo. Diese Art wurde später nie wieder gefunden. 


236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 1971 


H. nubeculosa Burmeister 
Van der Wulp nennt ein Weibchen aus Holland. Das Exemplar ist nicht mehr vor- 
handen und die Art ist nie wieder gefunden worden. 
H. vittata (Meigen) 
Erstnachweis für Belgien: 1 ® Boma, 24.VI. 
Paradelphomyia (Oxyrhiza) nielseni Kuntze 
Erstnachweis für Holland: 1 & Baarn, 26.IX.1905, De Meijere. 
Gonomyia (Ellipteroides) alboscutellata Von Roser 
In Holland ist diese Art seit De Meijere (1916) nur von Bunde bekannt. 
Erstnachweis fiir Belgien: Houyet und Malmedy. 
G. (Gonomyia) bifida Tonnoir 
Diese Art wurde beschrieben nach Exemplaren von Virton. Bis heute sind keine 
neuen Fundorte in Belgien bekannt. 
G. (Gonomyia) lucidula De Meijere 
Erstnachweis für Luxemburg: 1 9 Bech-Kleinmacher, 30.VI.1970, Theowald. 
G. (Gonomyia) recta Tonnoir 
Diese Art wurde beschrieben nach Exemplaren von Virton und ist bisher in Belgien 
nur von diesem Fundort bekannt. 
Lipsothrix errans (Walker) 
Erstnachweis fiir Belgien: Bihain, Mirwart, Forét de Soignes. 
L. remota (Walker) 
Goetghebuer & Tonnoir (1921) kennen diese Art als Limnophila clara Tonnoir von 
Buzenol. Ich sah weitere Exemplare von: Mirwart, Lesterny, Grotte de Rochefort, 
Malmedy. 
Erioptera (Erioptera) divisa (Walker) 
In Holland war diese Art bekannt von Epen. Ich sah weitere Exemplare von Vaals, 
Kerkrade, Heerlen, Eygelshoven und Holset. 
In Belgien ist diese Art nur bekannt von Virton. 
E. (Erioptera) fusculenta Edwards 
Erstnachweis für Holland: De Bilt, Heerlen, Wassenaar, Zwammerdam, Mechelen, 
Kerkrade, Ravensbos, Vijlen, Bodegraven, Zunderdorp, Nieuwersluis, Epen. 
E. (Erioptera) limbata Loew 
In Belgien ist diese Art nur bekannt von Uccle. 
E. (Erioptera) lutea Meigen 
Erstnachweis für Luxemburg: 1 & Bech-Kleinmacher, 30.VI.1970, Theowald. 
E. (Erioptera) meijeri Edwards 
In Holland ist diese Art nur bekannt von Leimuiden. 
E. (Erioptera) minor De Meijere 
In Holland ist diese Art nur bekannt von Nieuwersluis. 
E. (Erioptera) nigripalpis Goetghebuer 
In Belgien ist diese Art nur bekannt von Forét de Soignes. 
E. (Psiloconopa) occoecata Edwards 
Erstnachweis für Luxemburg: 1 & Bech-Kleinmacher, 30.VI.1970, Theowald. 
E. (Arctoconopa) melampodia (Loew) 
Erstnachweis fiir Belgien: Mazy, Mazy-Onoz, Onoz-Spy. 
Chionea lutescens Lundstrom 
In Belgien ist diese Art nur bekannt von Francorchamps. 


B. THEOWALD: Die Tipuliden der Benelux-Länder 237 


Ormosia (Ormosia) albitibia Edwards 
Erstnachweis für Belgien: 1 & Petit-Lanaye (Grotte carrière), 8.IX. 
O. (Ormosia) bicornis (De Meijere) 
In Holland nur bekannt von Linschoten, in Belgien nur von Virton. 
O. (Ormosia) clavata (Tonnoir) 
Diese Art war in Belgien nur bekannt von Virton. Ich sah noch 1 £ von Mirwart. 
O. (Ormosia) depilata Edwards 
Erstnachweis für Belgien: Lesterny, Oignies, Bois de Couvin, Chimay. 
O. (Ormosia) staegerina Alexander 
Erstnachweis für Belgien: Ethe, Buzenol. 
O. (Rhypholophus) bivittata Loew 
Diese Art ist aus Holland nur bekannt von Diemen. 
O. (Rhypholophus) haemorrhoidalis Zetterstedt 
Erstnachweis für Holland: 1 & Heerlen, 24.1X.1964, Br. Arnoud. 
O. (Scleroprocta) pentagonalis (Loew) 
Diese Art ist in Holland nur bekannt von Bunde. 
O. (Scleroprocta) sororcula (Zetterstedt) 
Erstnachweis für Holland: 1 & Eygelshoven, 19.V.1958, Br. Arnoud. 
O. (Molophilus) appendiculatus (Staeger) 
Erstnachweis für Luxemburg: 2 & Mondorf, 28.VI.1970, Theowald. 
O. (Molophilus) ater (Meigen) 
Erstnachweis für Holland: 1 & De Lutte, 15.VI.1926, De Meijere. 
O. (Molophilus) bifidus Goetghebuer 
Erstnachweis fiir Holland: Heerlen, Mechelen, Eygelshoven. 
Erstnachweis für Luxemburg: 3 &, 2 © Bech-Kleinmacher, 30.VI.1970, Theowald. 
O. (Molophilus) bihamatus De Meijere 
Diese Art ist in Belgien nur bekannt von Overmeire. 
O. (Molophilus) curvatus Tonnoir 
Erstnachweis für Holland: 1 & Eygelshoven, 16.VII, 1954, Br. Arnoud. 
In Belgien war diese Art bekannt von Forêt de Soignes. Ich sah weitere Exemplare 
von Mirwart und Lesterny. 
O. (Molophilus) czizeki Lackschewitz 
Erstnachweis für Belgien: 2 ¢, 2 © Lesterny, 14.V. 
O. (Molophilus) flavus Goetghebuer 
In Holland ist diese Art nur bekannt von Bunde. 
In Belgien war diese Art bekannt von Hockai. Ich sah weitere Exemplare von 
Francorchamps. 
O. (Molophilus) occultus De Meijere 
In Belgien war diese Art nur bekannt von Hockai. Ich sah weitere Exemplare von 
Haute Fagne (Nesselo-Roer). 


LITERATURVERZEICHNIS 


Alexander, C. P., 1965. — Tipulidae in A catalog of the Diptera of America north of Mexico. — 
Agriculture Handbook 276: 16—90. U. S. Department of Agriculture. 

Goetghebuer, M. & A. Tonnoir, 1920—1921. — Catalogue raisonné des Tipulidae de Belgique. — 
Bul. Soc. Ent. Belg. 2: 104—112, 131—147; 3: 47—58, 105—125. 

Horn, W. & I. Kahle, 1935. — Uber entomologische Sammlungen, Entomologen und Entomo- 
Museologie. Ent. Beih. 2: 1—160. 


238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 6, 1971 


Laever, E. de & B. Mannheims, 1958. — Tipulides nouveaux pour la Belgique. — Bull. Ann. Soc. 
Roy. Ent. Belg. 94: 230—231. 

Meijere, J. C. H. de, 1919—1921. — Studien über palaearktische, vorwiegend holländische, Lim- 
nobiiden, insbesondere über ihre Kopulationsorgane. — Tijdschr. Ent. 62: 52—97; 63: 
46—86; 64: 54—118. 

—, 1928. — Vierde Supplement op de Nieuwe Naamlijst van Nederlandsche Diptera. — 

Tijdschr. Ent. 71: 11—83. 

, 1939. — Naamlijst van Nederlandsche Diptera. — Tijdschr. Ent. 82: 137—174. 

Mannheims, B., 1967. — Liste der Tipuliden Ostfennoskandiens. — Not, Ent. 47: 77—80. 

Mannheims, B. & B. Theowald, 1959. — Die Tipuliden Italiens. — Mem. Soc. Ent. Ital. 38: 


15—54. 

Savtshenko, E. N., 1961. — Tipulidae, Genus Tipula L. — Fauna SSSR, Nov. Ser. 79, Diptera 2: 
1—487. 

Schummel, T. E., 1833. — Versuch einer genauen Beschreibung der in Schlesien einheimischen 
Arten der Gattung Tipula Meigen. — Beitr. Ent., Besonders in Bezug auf Schlesien, 3: 
9—128. 

Theowald, B., 1957a. — Die Entwicklungsstadien der Tipuliden (Diptera, Nematocera), insbe- 
sondere der West-Palaearktischen Arten. — Tijdschr. Ent. 100: 195—308. 

, 1957b. De Nederlandse Langpootmuggen (Tipulidae). — Wetensch. Med. K.N.N.V. 

24: 1—28. 


Theowald, B & B. Mannheims, 1962. — Die Arten der Tipula (Vestiplex) excisa-Gruppe in der 
Paläarktis. — Bonn. Zool. Beitr. 4: 360—402. 

Tjeder, B., 1955. — Catalogus Insectorum Sueciae, XIV, Diptera: Fam. Tipulidae. — Opusc. Ent. 
(2) 20: 229—247. 


Le Wy 
sy 


f i Li 
SIMONI 


No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.FI. 45—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Biadrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae)). 350 pp. en 98 pl. D.Fl. 70.—. 


No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov 
and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. 
DFI. 25.—. 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. 
D.Fl. 100.—. 


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J. KRIKKEN. — New species of the Papuan genus Tafaia Valck Lucassen (Coleoptera : 
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NEW SPECIES OF THE PAPUAN GENUS TAFAIA 
VALCK LUCASSEN (COLEOPTERA: CETONIIDAE) 


by 
J. KRIKKEN 


Rijksmuseum van Natuurlijke Historie, Leiden 


With 53 text-figures and 1 plate 


ABSTRACT 


Four new species of the genus Tafaia Valck Luc. are described and figured, viz. T. brunnea, 
T. difficilis, T. bacchusi and T. signifer, all from New Guinea. These are keyed along with the 
only species hitherto known and a form not described herein. A key to some genera of the Loma- 
piera alliance serves to distinguish Tafaza from allied genera. 


A few decennia ago Valck Lucassen (1939: 141) proposed the genus Tafaia for the 
accommodation of a singular new cetoniine beetle of the Lomaptera alliance, collected 
in eastern New Guinea. Valck Lucassen’s paper remained the only original record of the 
genus. In that same year, however, two expeditions exploring unknown portions of 
western New Guinea had already secured other forms referable to Tafaia. I noticed 
these beetles when looking through the Leiden museum’s cetoniid accessions. Recently 
the British Museum (Natural History) placed further Tafaia-like beetles at my disposal, 
while correspondence with G. Ruter revealed that more captures are due to be reported. 

Some obstacles in classifying Tafaia species and their relatives cannot be suppressed. 
The type of T. viridiaenea Valck Luc. is a female, and a comparison with the females 
described hereafter showed them all to be quite similar. The males, however, are variably 
different, and judged from one case a pronounced sexual dimorphism may be expected 
in others. In a second case a male could not be associated with a female from the same 
region, though their conspecificity seems likely. Anyway, despite a certain heterogeneity, 
the available males and females all run fairly easily down to Tafaia in Valck Lucassen’s 
key (1961: 4—5) to the genera of the Lomaptera alliance. In this connexion it may 
be noticed that, although Valck Lucassen has produced a very useful key, a clear-cut 
classification has not been gained so far, owing to the existence of poorly known 
intergeneric and otherwise problematic forms. Valck Lucassen himself was well aware 
of this, as appears from the notes accompanying his key. To avoid any misunder- 
standing, the generic limitations of Tafaia are reconsidered below, in a key to some 
genera of the Lomaptera alliance (= Lomapterides sensu Schenkling, 1921: 110—128). 
From this key the close affinities of Tafaia and Digenethle Thomson are evident; they 
may eventually turn out to be at most subgenerally distinct. 

For allowing the study of the T. viridiaenea type and for submitting more material, 
I am indebted to Messrs R. D. Pope and M. E. Bacchus of the British Museum (Natural 
History), London. Mr. G. Ruter, correspondent of the Muséum National d'Histoire 
Naturelle, Paris, kindly allowed the inclusion of information on two Tafaia-like beetles 
while a paper of his dealing with these insects is forthcoming. 


239 


240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


Fig. 1—10. Comparison of genera of the Lomaptera alliance, outlines. 1, Lomaptera macrophylla 

Gestro, 4, Wissel Lakes area, head; 2, Agestrata punctatostriata Lansberge, 4, Sangi & Talaud, 

clypeus; 3—6, Microlomaptera aenea Kraats, 6, Aru Is.; 7—10, Digenethle caelata Thomson, 4, 

Sattelberg; 3, 7, head; 4, 8, pronotum; 5, 9, left elytron; 6, 10, pygidial outline. 1 — 3, 7, full-face, 
4, 5, 8, 9, dorsal, 6, 10, dextro-lateral view. Scale-lines = 1 mm 


KEY TO SOME GENERA OF THE Lomaptera ALLIANCE 


1. Anterior border of clypeus emerginate, lateral lobes simply rounded (Figs. 3, 7, etc.). Scutellum 
very distinct (Figs. 5, 9, etc.). Pygidial shape simple, more or less convex, with simply arcuate 
outline. (ventrally view) iii) se). sewer eerden vends CAPES 

— Anterior border of clypeus deeply incised (Fig. 1), or shallowly emarginate with small, pointed 
lateral protrusions (the oriental genus Agestrata Eschscholtz, Fig. 2), in the latter case body- 
dimensions excessive. Scutellum indistinct, or even entirely invisible. Pygidium frequently 


J. KRIKKEN: New species of Tafaia 241 


modified. Parameral structure differing according to genus. — E and S Asia, Australia, and 
neighbouring islands, hundreds of species . . . 0 a ecightusenera 
. Pygidium shallowly convex, without distinct dorso- Senta] pension (a Fig. 6 with 10). 
Phallus relatively complex, with transverse dorso-basal parameral cavity; flaps between para- 
meres (Fig. 11) small. Dorsal outline of pronotum as in Fig. 4, at least with noticeable antero- 
lateral angles. Clypeal surface convex medially. Small forms, total length not exceeding 20 mm. 


— Aru Islands and New Guinea, two described species. . . . . Microlomaptera Kraatz 
Pygidium strongly convex, outline as in Fig. 10, 23, etc. Phallus relatively simple. Dorsal out- 
line of pronotum with obsolete antero-lateral angles (Fig. 8, etc.) . . . . . . 5 6 


. Clypeal surface concave, consequently margins raised, more or less distinctly carini Flaps 
between parameres small or absent (Fig. 13 and 14). Superior terminal spur of hind tibiae 
spatulate (Fig. 20) (in the known females). If elytral disc striolate, pattern of striae not 
fully obsolete. — New Guinea, at least six species. . . heren te lajara Nalck luc. 
Clypeo-frontal surface somewhat convex. Flaps between res (Fig. 12) large. Elytra non- 
striate, non-costate, densely, braidedly striolate throughout discal striolae transverse or oblique. 
— New Guinea, four described species . . . . . . . . . . . Digenethle Thomson 


KEY TO THE SPECIES OF Tafaia 


. Forms with tapering mesosternal process (Fig. 22, 49), its length exceeding middle coxal width. 
Postero-lateral angle of hind coxae 21, etc.) distinct owing to concavely curvilinar 
posterior edge . . EN METRI NAT RD EAT en (elec NEZ 
Mesosternal process (Eis 28) oan Do el angle of hind coxae (Fig. 27) obtuse. 
Pronotum bicolorous. Sides of distal sternites with sparse primary and dense secondary punctation 
. Pygidium protruding, but short, largely covered by elytra. Antennal club (Fig. 47) modified in 
the males (at least as long as the remaining segments combined). Anal sternite very short (e.g. 
as in Fig. 50); pygidial length (ventral view) at least twice the median length of this sternite; 
borders of anal sternite not reaching ventro-dorsal transition. Strial pattern of elytra distinct. 
Elytral and pronotal colours not different. For tibiae of males with apical denticle only . 6 
Pygidium distinctly visible from above. Lamellae of antennal club (compare Fig. 37 with 47) 
not enlarged. If anal sternite relatively short, pronotal and elytral colours different, and ventrum 
(particularly anteanal sternite) only very sparsely setose . . . A APRES 
. Abdominal sternites, particularly anteanal one, with numerous Brown A Det concolorous; 
colour of appendages may differ from that of body, which is brownish or greenish. Larger 
forms, with spatulate superior terminal spur on the hind tibiae of the only known female sex . 4 
Abdominal sternites glabrous or nearly so, punctation sparse. Dorsum distinctly bicolorous; 
head and pronotum yellow-brown, elytra dark brown, etc. Lateral borders of pronotum raised. 
Elytra non-striate. Clypeal surface with crowded punctulation (Fig. 31a). Length 22.5—27 mm. 
— E New Guinea. . . De Fel ay kbacchusı,sp. Nox. 
. Clypeal surface distinctly ee vents) ml fee Gi dai sternites with numerous 
distinct seta-bearing primary punctures. Colour of entire dorsum greenish bronze. Spatulate spur 
of hind tibiae wide, width subequal to length of second tarsal segment, hind tibiae orange, apex 
black. Habitus PI. 1 Fig. 1. Length 30 mm. — E New Guinea: Mt. Tafa . viridiaenea Valck Luc. 
Cephalic surface with many primary punctures separated by more than their own diameters. 
Lateral surface of abdomen with dense transverse striolation, primary punctures obsolete, though 
setae present. Colour of entire dorsum brown. Spatulate spur of hind tibiae (Fig. 20) 
relatively narrow. All tibiae blackish. ne 30—31.5 mm. — W New Guinea: Central 
Range... . +. brunnea sp. nov. 
. Middle tibiae with Si spine at don Go ds from tibial ase! Serially arranged primary 
punctures on elytral disc very distinct. Single known female with spatulate superior spur in hind 
tibiae. Elytral apex striolate. Length 31 mm. — W New Guinea: Central Range . : 
difficilis sp. nov. 
Middle Ehe ne el Keane, Ceres de at punctures on elytral disc indistinct. 
Superior spur of hind tibiae tapering. Elytral apex non- striolate. Length 26 mm. — W New 


242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


Fig. 11—14. Parameres, full-face view. 11, Microlomaptera aenea Kr., Aru Is.; 12, Digenethle caelata 
Th., Sattelberg; 13, Tafaia sp. nr difficilis, Moss Forest Camp; 14, Tafata bacchusi sp. nov., Mt. 
Kaindi, with laevo-lateral view. Scale-lines — 1 mm, 12—14 same scale 


Guinea: Central Range. . . . . . male of difficilis or sp. ined. 
6. Dorsum green. Head glabrous, dies ral concave. Antescutellar lobe truncate. Lamellae 
of antenna in male about as long as remaining segments combined. Length 22 mm. — E New 
Guinea: Mt. Jimmi. . . . sp. nov. Ruter ms. 
— Dorsum brown. Clypeo- frontal cures iosa dpi mais ook raised (Fig. 42, 51). 
Antescutellar lobe (Fig. 43) conspicuously notched. Antennal lamellae of male (Fig. 47) much 
longer than remaining segments combined. Length 23—24 mm. — E New Guinea: Morobe 
Districten aan oe et D RI NA a en e TAR yo GIRAA? Go DOV. 


Tafaia brunnea sp. nov. 
Fig. 15—23, PI. 1 Fig. 2 


Female (holotype). — Approximate length 30, width 16, height 11 mm. Brown, with 
tinge of red, weakly shiny, most of integument shagreened; tips, margins, sutures of 


J. KRIKKEN: New species of Tafaia 243 


several elements more or less infuscated, tibiae and tarsi blackish; pilosity brown. 
Habitus, Pl. 1 Fig. 2. 

Cephalic contours, Fig. 15. Clypeus shallowly concave, hence margins raised; cephalic 
primary punctation moderately dense, punctures approximately isodiametric, moderately 
impressed; punctural diameters on clypeus 0.05—0.15 mm, their densities ca. 20/sq. mm; 
interspaces with close secondary punctation, punctures fine, shallow. Eye-canthi with 
about 5 long setae. Maximum length of head 4.50, maximum width 4.95 mm; ratio 
I/w 0.90. 

Pronotal contours, Fig. 16; surface evenly convex, lateral margins not raised. Pronotal 
punctation double; discal surface with scattered, sparse primary punctation, densities 
increasing laterad, passing into marginal striolation; diameters of postero-median 
punctures 0.05—0.10 mm, their densities ca. 10/sq. mm; diameters of sublateral 
punctures 0.10—0.15, their densities 10—15/sq. mm; entire pronotal surface with 
secondary punctation like on the head. Median length of pronotum 9.6, maximum width 
12.7 mm; ratio 1/w 0.75. Scutellum (Fig. 17) raised medially, with numerous extremely 
fine punctures, and several seta-bearing punctures in front. 

Elytral contours, Fig. 17; juxtasutural costa of elytron distinct behind, obsolete in 
front; strial pattern indistinct; elytral base sparsely punctate; disc with numerous in- 
fuscated punctures, each surrounded by an arcuate striola; separate elements obsolescent 
caudad and laterad, passing into braided striolation. Secondary punctation dense, 
punctures extremely fine. Distance between elytral and scutellar apices 14.5 mm; 
maximum length of left elytron 19.5, maximum width of elytra combined 14.5 mm, 
their ratio 1.34. 

Latero-ventral surface of prothorax densely striolate, with numerous semierect setae; 
antero-median portion of prosternum distinctly raised, with long brown setae in front. 
Mesosternal process (Fig. 22) well-developed, dorsally with median ridge, ventrally 
weakly transversely convex. Mesepimera and mesosternum densely striolate, with numerous 
semierect setae. Metasternal disc shallowly sulcate medially, shiny, with several fine 
punctures; metasternal wings, metepisterna and greater part of hind coxal surface 
densely transversely striolate, with numerous semierect setae. Metepimera shiny, with 
small as well as extremely fine punctures. Postero-lateral angles of hind coxae (Fig. 21) 
distinct. Visible abdominal sternites 1—4 medially with sparse hemipunctures (i.e. 
punctures with about half of their borders obsolete) bearing fine semierect setae; 
laterally these sternites are densely punctulate-striolate, similarly setose; sternite 5 with 
numerous hemipunctures in the middle, punctures with coarse, semierect setae; laterally 
this sternite is transversely striolate, with similar setae, middle of sternites 1—6 with 
vague, fine secondary punctation; anal sternite finely, transversely striolate, glabrous. 
Pygidium (Fig. 23) with arcuate apical outline (seen from above); dorsal and ventral 
surface separated by transversely carinate apex; anal edge indistinctly marginate, margin 
widened medially; pygidial sculpture consisting of approximately concentric striolae; 
many fine setae are noticeable. 

Fore tibia (Fig. 18) with 3 denticles and well-developed terminal spur; both sides 
with many seta-bearing hemipunctures. Inferior side of fore femur with setae, striolae 
and punctures. Middle and hind tibiae (Fig. 19) with scattered medium-sized hemi- 
punctures, bearing semierect setae; outer surface of middle and hind tibiae lacking 
elevations; superior terminal spur of hind tibiae (Fig. 20) spatulate, longer than simply 
acuminate inferior one; spurs of middle tibiae both acuminate, inferior slightly longer. 


244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


Fig. 15—23. Tafaia brunnea sp. nov., 2 holotype, Top Camp. 15, left half of elytron, inset with 

details; 16, pronotum, 16a, enlarged details of pronotal disc (point a); 17, left elytron, scutellum; 

18, right fore tibia, superior side; 19, right hind leg; 20, apex right hind tibia; 21, lateral part 

of hind coxa; 22, mesosternal process, laevo-lateral; 23, pygidium 15, 18—19, full-face, 16—17, 
dorsal, 20, dextro-lateral, 21, 23, ventral view. Scale-lines = 1 mm 


Inferior side of middle and hind femora sparsely striolate-punctate, with semierect setae, 
mixed with many irregularly set punctules. 


Variation. — Length 30—31.5 mm. No taxonomically noteworthy differences among 
the few females at hand. 
Diagnostic remarks. — Tafaia brunnea may be identifiable from the sculptural 


features of many body-parts and the following combination of characters. Dorsum 
brown, concolorous, tibiae entirely blackish. Mesosternal process well-developed, 
tapering. Postero-lateral angle of hind coxae distinct. Spatulate spur of hind tibiae 
(in the female) relatively narrow. Pronotal borders immarginate. Elytral striation 
obsolete. 


J. KRIKKEN: New species of Tafaia 245 


Material examined. — 3 females (Leiden museum). 

Holotype, western New Guinea, labelled as follows: “Ned. Ind.-American/New 
Guinea Exped./Top Camp 2100 m/8.11.1939 L. J. Toxopeus”. One paratype with the 
same data, except collected 7.11.1939; another from the Wissel Lakes area: Digitara, 
ult.X1.1939, leg. J. Hoeka (K.N.A.G.-Le Roux expedition). 


Tafaia difficilis sp. nov. 
Fig. 24—29, Pl. 1 Fig. 3 


Female (holotype). — Approximate length 31, width 17, height 11.5 mm. Dark 
brown, more or less shiny, pronotum very shiny; lateral portions of thoracic segments 
(including pronotum), as well as hind coxae and sides of pygidium yellow-brown; tips, 
margins, sutures of several elements more or less infuscated, tibiae blackish; pilosity 
brown; microsculpture indistinct on head and pronotum (magnification 50). Habitus, 
PMR 193: 

Cephalic contours, Fig. 24. Clypeus concave, antero-lateral borders distinctly raised; 
clypeo-frontal surface with vague secondary punctation and rugulation; primary punctures 
closely set anteriorly, approximately isodiametric, shallow, ill-defined, their diameters 
ca. 0.1 mm, densities in the middle ca 30/sq. mm. Surface of vertex with vague primary 
and secondary punctures. Eye-canthi with slightly over 10 erect setae. Maximum length 
of head 4.75, maximum width 5.35 mm; ratio 1/w 0.89. 

Pronotal contours and disposition of colours, Fig. 25; surface evenly convex; sides 
shallowly marginate except near postero-lateral angles; lateral zones yellow-brown with 
an isolated patch of dark brown, discal surface dark brown, extremely shiny. Pronotal 
primary punctures shallow but distinct, approximately isodiametric, their diameters in- 
creasing laterad, lateral punctures with obsolete hind borders and some of them con- 
fluent; primary punctures on basal lobe small and sparse; densities of primary punctures 
on disc about 40 or less/sq. mm, their diameters less than 0.05 mm, increasing laterad 
to over twice that size for the punctures along the border; secondary punctation dense, 
covering entire pronotum, punctures extremely fine. Median length of pronotum 9.1, 
maximum width 12.4 mm; ratio l/w 0.73. Scutellum (Fig. 26) punctate-regulate in 
front, with numerous semierect setae; posterior surface with exceedingly fine punctation. 

Elytral contours, Fig. 26; juxtasutural costa distinct behind, obsolete in front; striae 
represented by 6 series of well-defined umbilicate punctures, their diameters varying, 
amounting to nearly 0.5 mm, distances varying as well; distal portions of elytron trans- 
versely striolate; remaining surface with fine, sparse, scattered primary punctures, a 
secondary punctation like on the pronotum, though less dense, and a microsculpture. 
Distance between elytral and scutellar apices 16.8 mm; maximum length of left elytron 
19.8, maximum width of elytra combined 14.5 mm, their ratio 1.38. 

Legs and ventral side of body with minute secondary punctures, partly obliterated 
or replaced by other sculptures. Coxae and ventro-lateral portions of thorax with seta- 
bearing striolae, metepimera with seta-bearing punctures. Yellow-brown are: proepimeral 
concavities, humeral extremity of mesepimera, metasternal wings, metepisterna, and 
metepimera (margins infuscated). Raised antero-median part of prosternum longitudin- 
ally sulcate. Mesosternal process (Fig. 28) short. Postero-lateral angle of hind coxae 
(Fig. 27) rounded. Visible abdominal sternites striolate-setose, 2—4 with many sparsely 
set, small hemipunctures bearing semierect setae; anteanal sternite with coarser, large, 


246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


Fig. 24—29. Tafaia difficilis sp. nov. ® holotype, Moss Forest Camp. 24, left half of head, 

inset with details; 25, pronotum, 25a, enlarged details of pronotal disc (point a); 26, left elytron, 

scutellum; 27, lateral part of hind coxae; 28, mesosternal process, laevo-lateral; 29, pygidium. 24, 
full-face, 25—26, dorsal, 27, 29, ventral view. Scale-lines = 1 mm 


seta-bearing hemipunctures, their diameters ca 0.1 mm, density in the middle ca 7/sq. 
mm; anal sternite transversely striolate with some inconspicuous punctures lacking setae. 
Pygidium (Fig. 29) with arcuate apical outline (seen from above), extreme apex lightly 
emarginate; dorsal and ventral sides distinctly separated, ventral side with median 
longitudinal callosity, but apex slightly impressed; sculpture consisting of approximately 
concentric striolae, obliterated on ventral surface; setae sparse, fine, inconspicuous. 

Fore tibia with 3 denticles and well-developed terminal spur, like in T. brunnea; 
superior side with numerous scattered, rather large seta-bearing hemipunctures, in- 
ferior side with several smaller seta-bearing punctures. Fore femur with many small 
seta-bearing punctures above, larger seta-bearing hemipunctures beneath. Middle and 
hind tibiae with scattered medium-sized hemipunctures bearing coarse semierect setae; 
external side of middle tibiae with spine-like projection slightly behind the middle; 
superior terminal spur of hind tibiae spatulate, longer than simply acuminate inferior 
one; spurs of middle tibiae of equal length, acuminate; middle tibiae with many, hind 
tibiae with few seta-bearing hemipunctures, their dimensions variable. Inferior side of 
middle and hind femora with scattered hemipunctures bearing semierect setae, less 
sparse on the middle ones. 


J. KRIKKEN: New species of Tafaia 247 


Diagnostic remarks. — Tafaia difficilis may be identifiable from the following 
combination of characters. Mesosternal process short. Postero-lateral angle of hind coxae 
obsolescent. Pronotum with yellow-brown lateral margins enclosing a dark patch halfway 
pronotal length. Sculptural features of dorsum distinctive as well. 

Material examined. — Holotype, western New Guinea, labelled thus: “Neth. Ind.- 
Amer. New/Guinea Exped. 2600—/2800 m. Moss Forest/ Camp 9.X.—5.X1.1938/L. J. 
Toxopeus leg.” (Leiden). 

Note. — See at the end of the following description. 


Tafaia sp. 
Fig. 13, Pl. 1 Fig. 4 


Male. — Approximate length 26, width 12, height 9 mm. Dark brown, largely shiny; 
lateral portions of thoracic segments (including pronotum) as well as hind coxae and 
most of pygidium yellow brown; tips, margins, sutures more or less infuscated; pilosity 
brown; most of integument with microsculpture (magnification X 50). Habitus, Pl. 1 
Fig. 4. 

Cephalic contours largely similar to those of T. difficilis (preceding description). 
Clypeus concave, lateral borders distinctly raised; clypeo-frontal surface with vague 
secondary punctation and rugulation; primary punctures closely set anteriorly, ap- 
proximately isodiametric, very shallow and ill-defined. Vertex with more distinct primary 
punctures and exceedingly fine secondary ones. Eye-canthi with slightly more than 
10 erect setae. Maximum length of head 4.65, maximum width 4.70 mm; ratio 1/w 1.00. 

Pronotal contours and disposition of colours largely similar to those of T. difficilis; 
surface evenly convex; sides shallowly marginate except near postero-lateral angles. 
Pronotal primary punctures approximately isodiametric, shallow but distinct, their hind 
borders more or less obsolescent; primary punctures of basal lobe fine; densities of 
discal primary punctures ca 25—30/sq. mm, their diameters ca 0.05 mm; extremely 
fine secondary punctation covering entire pronotum. Maximum length of pronotum 
8.15, maximum width 10.2 mm; ratio 1/w 0.81. Scutellum (shaped like in T. difficilis) 
punctate in front, with some setae; apical surface with exceedingly fine punctures. 

Elytral contours and disposition of striae largely similar to those of T. difficilis; 
juxtasutural costa distinct behind, obsolete in front; striae represented by several irregular 
series of ill-defined impressions, basally with traces of genuine punctures in their centres, 
distally these impressions are irregularly confluent; surface on and around apical 
umbone non-striolate, slightly uneven; entire surface with secondary punctation like on 
pronotum. Distance between elytral and scutellar apices 11.6 mm; maximum length of 
left elytron 15.9, maximum width of elytra combined 12.3 mm, their ratio 1.30. 

Legs and ventral side of body with minute secondary punctures. Ventro-lateral 
portions of prothorax striolate, partly setose, posterior concavity yellow-brown, other 
thoracic elements also yellow-brown, as in T. difficilis. Raised antero-median part of 
prosternum longitudinally sulcate. Mesosternal process short. Mesepimera proximally 
with seta-bearing striolae, elsewhere, on metepisterna, metasternal wings, and hind 
coxae, with sparse, small, seta-bearing hemipunctures, some of them transversely con- 
fluent. Postero-lateral angle of hind coxae shortly rounded. Abdominal sternites with 
very sparsely set, very small hemipunctures, partly bearing fine semierect setae; anal 


248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


segment with fine subtransverse striolae laterally. Superior surface of pygidium with 
similar fine striolae, inferior side with sparse, small punctures; anal border marginate. 

Fore tibia with only two denticles, terminal spur well-developed; both sides with a 
number of small hemipunctures, a few of them bearing fine setae. Inferior side of 
middle and hind tibiae with scattered hemipunctures bearing semierect setae, denser on 
the middle ones, external projections lacking, spurs of middle and hind tibiae simply 
tapering, superior spurs longer than inferiors. Inferior side of fore and middle femora 
sparsely provided with small punctures, some of them bearing setae; hind femora almost 
entirely smooth and glabrous beneath. 

Parameres, Fig. 13. 

Material examined. — One specimen (Leiden), western New Guinea, labelled thus: 
“Neth. Ind. - Amer. New/Guinea Exped. 2600—/2800 m. Moss Forest Camp 9.X.— 
5.X1.1938/L. J. Toxopeus leg”. 

Note. — This specimen may represent the male sex of T. difficilis sp. nov., since 
the features mentioned in the key to the species (couplet 5) may be only sexual. As I am 
not certain of its identity, I am describing this male separately, without naming it. 


Tafaia bacchusi sp. nov. 
Fig. 14, 30—41, Pl. 1 Fig. 5 


Male (holotype). — Approximate length 22.5, width 16, height 7 mm. Cupreous 
yellow-brown: head, thorax, scutellum, pygidium, all weakly shiny; elytra, legs, antennae, 
abdominal sternites much darker brown, moderately shiny; most parts with a faint tinge 
of green; tips, margins, sutures of several parts more or less infuscated; entire integument 
more or less shagreened; pilosity brown. Habitus, Pl. 1 Fig. 5. 

Cephalic contours, Fig. 30. Clypeus shallowly concave, margins raised, rather abruptly 
declivous laterally; cephalic surface slightly uneven, almost entirely crowded with minute 
punctures; clypeus with many vague puncture-like impressions; surface adjacent to eyes 
as well as eye-canthi with a number of distinct fine punctures, crowded punctulation 
absent there. Maximum length of head 3.65, maximum width 4.15 mm; ratio 1/w 0.89. 

Pronotal contours, Fig. 31; surface evenly convex, sides marginate, except near 
shortly rounded postero-lateral angles. Pronotal sculpture like that of the head, minute 
punctures not crowded though closely set, more than 100/0.25 sq. mm; disc with small 
superficial impressions, their diameters not exceeding 0.1 mm; impressions of lateral 
surface arcuate, margins almost striolate. Maximum length of pronotum 7.3, maximum 
width 8.9 mm; ratio 1/w 0.82. Scutellum (Fig. 32) almost smooth. 

Elytral contours, Fig. 32; juxtasutural costa of elytron distinct behind, obsolete in 
front; disc with several more or less arcuate punctures which distally pass into striolae, 
particularly around apical umbone; entire surface covered with closely set, minute 
punctures. Distance between elytral and scutellar apices 10.2 mm; maximum length of 
left elytron 13.5, maximum width of elytra combined 10.9 mm, their ratio 1.24. 

Outline of antenna and mouthparts, Fig. 37—41. Femora, tibiae, and ventral side of 
body with cover of minute secondary punctures; ventro-lateral portions of thorax with 
distinct though in places very superficial striolation. Antero-median portion of proster- 
num distinctly raised, with long brown setae in front. Mesosternal process (Fig. 35) 
well-developed; parts of ventral elements of mesothorax with erect setae. Metasternum 
with hemipunctures passing into striolae laterad, many bearing subappressed to erect 


J. KRIKKEN : New species of Tafaia 249 


Fig. 30—41. Tafaia bacchusi sp. nov. 6 paratype, Mt. Kaindi. 30, left half of head, inset with 

details; 31, pronotum, 31a, enlarged details of pronotal disc (point a); 32, left elytron, scutellum; 

33, right fore tibia, superior side; 34, right hind leg; 35, mesosternal process, laevo-lateral; 36, 

pygidium; 37, right antenna; 38—41, mouth-parts, 38, right mandible, 39, right maxilla, 40, labrum, 

41, mentum and labium. 30, 33—34, full-face, 31—32, dorsal, 36—41, ventral view. Scale-lines 
= 1 mm, except 3la = 0.3 mm 


fine, very inconspicuous setae; disc with secondary punctation only. First visible ab- 
dominal sternite striolate, lateral portions of following very vaguely striolate, medial 
surface at most slightly wrinkled. Pygidium (Fig. 36) with arcuate apical outline (seen 
from above), dorsal and ventral surface separated; sculpture consisting of approximately 
concentric striolae, obliterated on ventral side. 

Fore tibia (Fig. 33) with 3 denticles and well-developed terminal spur; both sides of 


250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


tibia scatteredly hemipunctate, some punctures of inferior side seta-bearing, internal 
side with distinct series of seta-bearing punctures. Fore femur with striolae behind, 
anterior margin with seta-bearing punctures. Middle and hind tibiae (Fig. 34) with 
scattered, medium-sized hemipunctures, most of them bearing a fine short seta; external 
side of middle and hind tibiae lacking elevations; terminal spurs all acuminate, of equal 
length in the middle legs, superior one distinctly longer in the hind legs. Inferior side 
of middle and hind femora almost glabrous, faintly striolate, hemipunctate behind, 
primary sculpture of hind femora vague. 

Parameres, Fig. 14. 

Variation. — Length 22.5—27 mm. No taxonomically noteworthy variation. I have 
examined the parameres of the six males available and they are all remarkably alike. 

Diagnostic remarks. — Tafaia bacchusi may be identifiable from its small size, sculp- 
tural features, its colours and their disposition, and the following combination of 
characters. Abdominal sternites in the males not impressed medially, sparsely punctate 
and glabrous, or nearly so. Mesosternal process well-developed, tapering. Postero-lateral 
angle of hind coxae distinct. Phallus lacking parameral flaps. Lateral borders of prono- 
tum raised. Elytral striation obsolete. Anal sternite short (at least in the male). 

Material examined. — Six males, five in the British Museum (Natural History), one 
in Leiden. Holotype, eastern New Guinea, labelled thus: “Neth. Guinea:/Morobe Dist./ 
Mt. Kainde 8,000 ft./22.IX.1964”, “Stn. No. 20.”, “M.E. Bacchus./B. M. 1965—120”. 
Three paratypes with the same data; two paratypes from Mt. Hagen, one leg. F. S. 
Mayer, the other collected at 8,000 ft in 1949. 

Note. — G. Ruter is going to publish a description of a male from Wapenamanda 
closely allied to this species. From the ms. description and the drawings supplied to me 
some differences could be abstracted, but I am unable to place it satisfactorily and to 
insert it in my key. 


Tafaia signifer sp. nov. 
Fig. 42—53, Pl. 1 Fig. 6 


Male (holotype). — Approximate length 24, width 11.5, height 9 mm. Brown, 
shiny; frons, scutellum, parts of ventral side, pygidium and legs blackish; tips, margins, 
sutures of several elements more or less infuscated; pilosity yellow to orange. Habitus, 
PI. 1 Fig. 6; legs very slender. 

Cephalic contours and disposition of colours, Fig. 42. Clypeus distinctly concave, 
margins strongly raised, particularly along anterior emargination, clypeal surface with 
close primary punctation, punctures deep, well-defined, more or less isodiametric, many 
bearing erect setae; punctural diameters 0.05—0.15 mm, their densities 50—60/sq. mm; 
secondary punctures vague, scattered, minute. Punctation of blackish posterior part of 
head sparse in the middle, close beside eyes; these punctures medium-sized, with fine 
setae; secondary punctation indistinct. Eye-canthi with a number of fine seta-bearing 
punctures. Maximum length of head 4.2, maximum width 4.2; ratio I/w 1.00. 

Pronotal contours, Fig. 43; surface shallowly, evenly convex; lateral margin broadly 
thickened, except behind; basal lobe extensive, its apex distinctly notched. Medial surface 
of pronotum sparsely punctate, punctures distinct, approximately isodiametric, their dia- 
meters on disc 0.05—0.10 mm, densities 10—20/sq. mm; lateral punctation dense, 
more or less transverse, particularly in front, passing into poorly pronounced marginal 


J. KRIKKEN : New species of Tafaia 251 


Fig. 42—53, Tafaia signifer sp. nov.; 42—50, & holotype, Mt. Kaindi; 51—53, © allotype, Bulolo. 

42, left half of head, inset with details (setosity indistinct from above); 43, left half of pronotum, 

inset a with enlarged details of centre (point a) of pronotum; 44, left elytron, scutellum; 45, right 

fore tibiae, superior side (no details); 46, left hind leg; 47, left antenna; 48, outside of 3rd lamella; 

49, mesosternal process, with indication of colour limit, dextro-lateral view. 51, left half of head; 

52, right fore tibia, superior side; 53, outside of 3rd antennal lamella. 42, 45, 46, 51, 52, full-face, 
43—44, dorsal view. Scale-lines — 1 mm, except inset 43a — 0.3 mm 


252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


striolation; density sublateral punctures 20—30/sq. mm; secondary punctation dense. 
Maximum length of pronotum 8.3, maximum width 9.9 mm; ratio I/w 0.83. Scutellum 
(Fig. 44) largely covered by pronotal lobe, scutellar apex acute, surface with sparse, 
minute punctures. 

Elytral contours, Fig. 44; juxtasutural costa distinct behind, obsolete in front; juxta- 
sutural interstria separated from adjacent surface by longitudinal striola; disc with three 
shallow costula, 2nd obsolete behind, the others extending to distal surface; costulae 
separated by irregularly arcuate striolae, some of them associated with punctules; lateral 
and distal surface densely transversely striolate, striolation interrupted by aforesaid 
costulae; humeral surface non-striolate; apical umbone obsolete; secondary punctation 
scarcely noticeable (magnification 50). Distance between elytral and scutellar apices 
9.6 mm; maximum length of left elytron 14.0, maximum width of elytra combined 10.9 
mm, their ratio 1.29. 

Segments of antennal club (Fig. 47) exceedingly long, about twice the length of the 
remaining segments together; exposed inward side of lamella 1 with track of semierect 
setae, their length decreasing apicad, outward side of lamella 3 with striola paralleling 
superior edge (Fig. 48). Antero-median portion of prosternum distinctly raised, with 
very long setae. Latero-ventral surface of prothorax striolate, setose, except for yellow- 
brown margin; proepimeral cavity deep. Mesosternal process (Fig. 49) large, somewhat 
prow-shaped; base blackish, tip brown. Yellow mesepimeron striolate-punctate, finely 
setose, laterally with few punctules only. Metasternal disc with sparse hemipunctules 
bearing fine setae; secondary punctation distinct, though punctures minute, sparse, 
scattered. Remainder of pectus, and hind coxal surface striolate, setose. Postero-lateral 
angle of hind coxa (Fig. 46) very distinctly produced. Black-brown venter with 
shallowly impressed midline, sternites laterally with arcuate, more or less confluent, 
seta-bearing striolae; medial surface with sparse, partly seta-bearing hemipunctures, 
setae very fine; posterior margin of anteanal sternite with numerous hemipunctures 
bearing long, coarse setae; secondary punctures as on metasternal disc, though less 
distinct; visible part of anal sternite (Fig. 50) remarkably narrow. Pygidium (Fig. 50) 
strongly convex, with distinctly separated dorsal and ventral sides; apex protruding 
only slightly beyond elytral apex; anal border distinctly marginate; pygidial surface 
with dense, approximately concentric striolation; sides of ventral surface yellow-brown. 

Fore tibia (Fig. 45) with acute apical denticle only, terminal spur well-developed; 
superior side of tibia finely striolate, several striolae associated with punctule bearing 
fine seta; inferior side with numerous seta-bearing punctules; colour of this male’s fore 
tibia externally blackish, internally brown, colours of other tibiae similar. Tarsal segments 
brown, infuscated distally; claws large, simply sickle-shaped. Fore femur superiorly 
closely hemipunctate in front, nearly all punctures bearing a semierect, rather long seta; 
inferior side striolate-setose. Middle and hind tibiae (Fig. 46) lacking external 
elevations; internal costa with a series of densely, obliquely set, long setae, remaining 
surface with numerous small punctures bearing small bristles; terminal spurs of middle 
and hind tibiae acuminate, those of middle tibiae about equal-sized, superior one on hind 
tibiae slightly longer than inferior; tarsal segments of middle and hind tibiae slightly 
compressed. Inferior side of middle and hind femora striolate, setose, setae long, 
semiappressed. 

Parameres asymmetric, apparently mis-shapen; two separated flaps present. 

Female (allotype). — Approximate length 23, width 12, height 8.5 mm. Brown, 


J. KRIKKEN: New species of Tafaia 253 


shiny; frons, parts of pectus blackish; sternites dark brown; tips, margins, sutures, 
lateral spots on pronotum more or less infuscated; colour generally lighter than in male 
holotype; pilosity yellowish. Habitus much like male. 

Cephalic contours, Fig. 51. Clypeus distinctly concave, margins only shallowly raised 
along anterior margination; diameters of punctures 0.05—0.15 mm, densities ca. 15 
and less/0.25 sq. mm. Maximum length of head 4.2, maximum width 4.0 mm; ratio 
1/w 1.05. 

Pronotum and scutellum largely similar to those of male; diameters of punctures on 
pronotal disc 0.05—0.10, their densities 10—20/sq. mm; densities sublaterally 20— 
30/sq. mm. Maximum length of pronotum 8,3, maximum width 10.3 mm; ratio 1/w 0.81. 

Elytra largely similar to those of male, differences as follows; juxtasutural costa less 
distinctly raised behind; discal interstriae more equally developed, extending onto distal 
surface without becoming obliterated by dense striolation, which is restricted to lateral 
and apical parts. Distance between elytral and scutellar apices 11.6 mm; maximum 
length of left elytron 14.6, maximum width of elytra combined 11.8 mm, their ratio 
1.21. 

Segments of antennal club (Fig. 53) large, but length not exceeding that of remaining 
segments together. Lateral parts of pectus and hind coxae largely brown. Mesosternal 
process tapering, lateral ridges indistinct. Abdominal sternites not impressed medially, 
lateral sculpture of visible sternites 1—4 extending onto medial surface; anteanal 
sternite with numerous seta-bearing hemipunctures, their densities ca 12/sq. mm. Pygidial 
colour largely yellow-brown, dorso-basal surface dark-brown. 

Fore tibia (Fig. 52) 3-denticulate, broad; superior side with partly seta-bearing 
hemipunctures; denticular side somewhat infuscated. Setae fringing internal costa of 
middle and hind tibiae less numerous (ca 30) than in male; remainder of these tibiae 
with many seta-bearing hemipunctures; superior spur of hind tibiae narrowly spatulate. 
Inferior side of middle and hind femora with partly confluent seta-bearing hemi- 
punctures. 

Sexual dimorphism and variation. — Since only one specimen of either sex is avail- 
able, these aspects cannot be distinguished with certainty. Most conspicuous are sexual 
differences in the structure of antenna and fore tibia; the male has the abdominal 
sternites impressed along the midline. 

Diagnostic remarks. — Tafaia signifer may be identifiable from the following 
combination of characters. Basal lobe of pronotum distinctly notched in front of scutel- 
lum, of which only a small portion is visible. Clypeo-frontal surface setose. Visible 
portion of anal sternite very narrow, its posterior border not reaching ventro-dorsal 
transition. Only apex of pygidium visible from above. Elytral disc with shallow longi- 
tudinal costulae, apico-sutural angle of elytra distinct. Colour predominantly brown. 
Antennal lamellae of male about twice the length of the remaining segments together. 
Fore tibiae of male slender, with apical denticle only, those of female broad, 3-denti- 
culate. Clypeus of male with strongly raised margins. Mesosternal process well-developed, 
tapering. Postero-lateral angle of hind tibiae produced. Lateral borders of pronotum 
distinctly raised. 

Material examined. — One male, one female, Department of Agriculture, Port 
Moresby, to be deposited in the British Museum (Natural History). Holotype, eastern 
New Guinea, with locality-label reading: “In flight./Mt. Kaindi/ Wau, M. Dist./ 
18.1.1970 F. R. Wylie”. Allotype from the same region: Morobe Dist.: Bulolo, 25.X1. 
1968, leg. F. R. Wylie, taken from “gauze screen house”. 


254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 


Approximate location altitude of places mentioned above!) 


Bulolo 7 12 § 146° 39’ E 600 m 
Digitara 3° 41’ 136° 28 1800—2200 
Hagen, Mt. 5 45 144° 05° 2400 
Kaindi, Mt. 7 DI 146° 43' 2400 
Moss Forest Camp 4° 00° 138° 43’ 2650—2800 
Tafa, Mt. 9° 38 1 OF 1800 
Top Camp 3° 30’ 139° 02’ 2100 
Wapenamanda 5° 40' WSS 55 1750— 2700 


1) To be located: Mt. Jimmi (E New Guinea, between Baiyer River and Jimmi River, high 
altitude, collections made by Jolivet, 1969). 


REFERENCES 


Schenkling, S., 1921. Scarabaeidae: Cetoninae. — Coleopterorum Catalogus 72, 431 pp. 

Valck Lucassen, F. T., 1939. Description of a new genus and a new species of Lomapterides 
(Coleoptera: Cetoniidae). — Ent. Ber., Amsterdam 10: 141—143, fig. a—d. 

, 1961. Monographie du genre Lomaptera Gory & Percheron (Coleoptera, Cetonidae). 
[D. L. Uyttenboogaart & C. de Jong, eds.}. — Amsterdam: Ned. Entom. Vereeniging, 
IV + 299 pp. fig. 1—739, 1—28, 1 map. 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 7, 1971 PLAAT 1 


Plate 1. Figs. 1—6. General appearance of Tafaia species. 1, T. viridiaenea Valck Luc., 2 holo- 

type, Mt. Tafa, length 30 mm; 2, T. brunnea sp. nov. ® holotype, Top Camp, length 30 mm; 

3, T. difficilis sp. nov. @ holotype, Moss Forest Camp, length 31 mm; 4, Tafaia sp. nr. difficilis, 

4, Moss Forest Camp, length 26 mm; 5, T. bacchusi sp. nov, 6, holotype, Mt. Kaindi, length 
22.5 mm; 6, T. signifer sp. nov. & holotype, Mt. Kaindi, !ength 24 mm 


J. KRIKKEN : New species of Tafaia 


ON EN 


No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.FI. 45.—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae)). 350 pp. en 98 pl. D.Fl. 70.—. 


No. 4. F. Willemse, — Preliminary revision of the genera Stenocatantups Dirsh & Uvarov 
and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. 
D.Fl. 25.—. 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. 
D.FI. 100.—. 


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LIFE HISTORIES OF SPECIES IN THE PARDOSA PULLATA 


GROUP, A STUDY OF TEN POPULATIONS IN THE 
NETHERLANDS 
(ARANEAE, LYCOSIDAE) 


by 


J. DEN HOLLANDER 
Department of Biology, Free University, Amsterdam 


ABSTRACT 


The phenology of several populations of the Pardosa pullata group — P. prativaga (L. Koch), 
P. prativaga var. fulvipes (Collett) and P. pullata (Clerck) — in the Netherlands was studied in 
1969 and 1970. The results are discussed in the context of the habitats of the respective species, 
the classification of the P. pullata group, as well as in the context of the possibility of hybridization 
in mixed populations. 


CONTENTS 


I. Introduction À 
II. Materials and methods à 
III. Results 


IV. 


Va 
WIE 
VII. 


1. Changes in spa bre oi structure à duc the year . 
A. General differences between the subgroups . 


a. Adults 
b. Juveniles 


B. Population differences within ae sram. 


a. subgroup prativaga . 
b. subgroup fulvipes 
c. subgroup pullata . È 
C. The number of eggs per cocoon . 
D. Conclusions 
2. Growth of juvenile spiders 
A. subgroup prativaga . 
B. subgroup pullata 
C. subgroup fulvipes 
D. Conclusions 


3. The habitat of the he population fi 


A. Description 

B. Conclusions i 5 
General conclusions and discussion : 
Postcript 
Summary 
RE ledoements 
References . 


255 


256 
256 
25 
257 
257 
257 
258 
258 
258 
263 
263 
264 
265 
266 
266 
268 
268 
269 
270 
270 
273 
275) 
279 
279 
280 
280 


256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 
I. INTRODUCTION 


In the Pardosa pullata group the species Pardosa prativaga (L. Koch, 1870) (in- 
cluding P. prativaga var. fulvipes (Collett, 1875)) and P. pullata (Clerck, 1757) are 
distinguished. The species differ in the annulation and spinosity of the legs to a fairly 
large extent (Wiebes, 1959). However, the general morphology suggests a very close 
relationship. Comparison of several populations showed that within either species mor- 
phological differences exist between samples from pure and those from mixed popu- 
lations. These data and the observation of specimens intermediate between P. pullata 
and P. prativaga in the field suggest that hybridization may occur (Locket & Millidge, 
1951; Den Hollander, 1970). 

In this paper the phenology of the species mentioned above is dealt with. The results 
are discussed in the context of the habitats of the species, the systematic relationships 
and the possibility of hybridization between the species. Studies concerning physiological 
and ethological barriers, as well as niche differentiation are in progress. 

The life-history of the species of the P. pullata group is of the common Pardosa type 
(Wiebes, 1960; Vlijm, Kessler & Richter, 1963; Vlijm & Kessler-Geschiere, 1967). In 
early spring only juvenile spiders occur. Within a few weeks after the appearance 
of the first adults the numbers of males and females without cocoon reach a maximum. 
Probably during this period mating occurs (Tretzel, 1954; Wiebes, 1960). The first 
females with cocoon appear in the same period. Pulli emerge from the cocoons after 
about 4 weeks; they stay 1—3 days upon the abdomen of the female. 

In the beginning of June the numbers of adult males decrease. The number of 
females decreases in late summer. Simultaneously the numbers of juveniles increase. 
These juveniles overwinter. 


II. MATERIAL AND METHODS 


The P. pullata group is considered separated into three subgroups: the pullata, 
prativaga and fulvipes subgroups (Den Hollander, 1970). 

Ten populations in the Netherlands (Table 1) were sampled with a frequency of 
about once per ten days, from April to October, in 1969 and 1970. Each sample was 
obtained by collecting any visible spider by hand during a 20 min. period. Thus catch 
size will depend on such factors as the structure of the habitat, the developmental stage 
of the spider and the weather conditions. 

Measurements of the cephalothorax length (i.e. the distance between the posterior 
lobes of the cephalothorax and the anterior margin of the posterior median eyes, cf. Den 
Hollander, 1970), were used to study the succession in time of the larval instars. 
Measurements in micrometer units (1 MU = 0.042) were obtained with a Reichert 
Stereomicroscope at a magnification of 6.3 X 4 (accuracy 2.3%). When taking the 
measurements, the anterior margin of the eyes and the posterior lobes of the cephalo- 
thorax were brought in focus simultaneously. 

The period of carrying pulli is short in relation to the period between the successive 
samples, so that the chances of catching females carrying pulli are small. Consequently, 
only few of them were caught. Therefore data on this stage are not included. 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 257 


Ill. RESULTS 


1. Changes in population structure during the year 
A. General differences between the subgroups 


Table II presents mean numbers per sample of juveniles, adult males and females 
(the latter without cocoon, indicated as 9 9, or with cocoon, indicated as 9 Qc), for 
the three subgroups separately. 


a. Adults 

Comparison of the phenological characteristics of the three subgroups gives the 
following results. Differences occurred as to the period of the final moult in the three 
subgroups (pullata: 4''/4”; fulvipes: 5’; prativaga: 5”; 4’, 4”, 4" stand for first, 
second and third decades of April, respectively). In each subgroup adult males were 
found about one week earlier than females. The highest numbers of males 
occurred in May. However, in the subgroup pullata males were also present in April 
and, in smaller numbers, in June and July. High numbers of males occurred in the 
fulvipes subgroup in both May and June. Males of the pratzvaga subgroup generally 
were found only during May. 

The numbers of females without cocoon (9 ® ) were highest in May in the sub- 
groups pullata and prativaga. In the fulvipes subgroup large numbers of 9 ® occurred 
in May, but also in June and July. Small numbers of 2 ® occurred in June in the 
subgroups pullata and prativaga, and in August and September in the subgroup fulvipes. 
Thus in May the numbers of both ¢ 4 and 9 ® were highest. This indicates that the 
period of copulation then occurs. With respect to this period rather slight differences 
existed between the three subgroups (pullata: 5'/!"; fulvipes: 5” |"; prativaga: 5"). 


Table I. The localities of the studied populations of the P. pullata group 


Locality Code Species 

Vogelenzang, Heemstede VZ P. prativaga 

Groene Punt, Oostvoorne GP P. prativaga 

Lange Pad, Oostvoorne LP P. prativaga 

Bosweitje, Rockanje BW P. prativaga, P. pullata 
Amstelveense Poel, Bovenkerk AP P. pullata 

Hollandse Rading, Hilversum HR P. pullata 

Arnica-weitje, Schiermonnikoog Sch P. pullata 

de Eese, Steenwijk E P. prativaga var. fulvipes 
de Woldberg, Steenwijk S P. prativaga var. fulvipes 


Females carrying cocoons (9 Qc) appeared in the middle of May (5”) in the 
pullata subgroup. In the subgroups prativaga and fulvipes they appeared in the begin- 
ning of June (6’). Thus the egg maturing period (i.e. the period between the appearance 
of the first @ © and that of the first 9 Qc) was different in the three subgroups 
(prativaga: 10—20 days; fulvipes and pullata: 20—30 days). 

In all three subgroups the numbers of @ Qc were highest in June. However, in the 
subgroup pullata 9 Qc occurred already in May and in the fulvipes subgroup large 
numbers of © Qc were found not only in June, but also in July and August. In the 
prativaga subgroup they hardly occurred outside June. After the periods of peak num- 


258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


bers only small numbers of 9 Qc were found in July (prativaga), in July and August 
(pullata) and in September (fulvipes). 

The first pulli emerged from the cocoons in the period between the middle of June 
and the beginning of July. Again some differences existed between the subgroups 
(pullata: 6/7"; prativaga and fulvipes: 6''’/7’). Thus in the subgroups prativaga and 
fulvipes females carry their egg sacs during a period of 20—30 days. The length of 
this period seems to be longer in the subgroup pwllata (30—40 days). 

In the subgroup prativaga adult females hardly occurred after the emergence of the 
pulli. This means that females of this subgroup produced a cocoon only once. Probably 
they die after the emergence of the pulli. Adult females did occur in the subgroups 
pullata and fulvipes in that period. However, in the pullata subgroup their numbers 
were low, as compared with those in May and June. The fulvipes females occurred in 
comparable numbers both before and after the emergence of the pulli. For the sub- 
groups pullata and fulvipes more information comes from comparing the numbers of 
females carrying cocoons with those without cocoons (@ ®c/9 9). In the subgroup 
pullata maxima in this ratio occurred in the beginning of June (6’) and in the middle 
of July (7”), respectively. In the fulvipes subgroup the first peak occurred in the 
middle of June (6”), the second one around the end of July (7”’/8’). It may be 
assumed that the @ Qc found after the emergence of the first pulli are females which 
had constructed a cocoon for the second time. Indeed the sampled cocoons in the period 
just after the emergence of the first pulli were very fresh. Thus the majority of the 
females in the pzllafa subgroup produced a cocoon only once, as was the case in the 
prativaga subgroup. A smaller part produced a second cocoon (small numbers in July). 
In the fulvipes subgroup large numbers of females with a cocoon could be caught in the 
recpective periods. This indicates that in this subgroup production of a second cocoon 
is the rule. The small numbers of ® Qc occurring after the indicated periods in the 
respective subgroups, viz. in July for prativaga, in August for pullata and in September 
for fulvipes, probably concerned females which accidentally had lost their cocoon before 
the pulli emerged and afterwards produced a new one. 


b. Juveniles 

Juvenile spiders also occurred at different times during the year in the three sub- 
groups. Generally their numbers decreased strongly during the period of the final 
moult. In the subgroups pullata and prativaga juveniles were nearly absent in the 
samples after the appearance of the first @ Qc. In the fulvipes subgroup, on the other 
hand, large numbers of juveniles occurred throughout the whole summer. 

In the subgroups pullata and fulvipes juveniles occurred in large numbers in autumn, 
as well as in spring. However, prativaga juveniles scarcely could be found in the 
autumn samples. This may be caused by the specific behaviour of these juveniles in 
relation to habitat (see below). 


B. Population differences within the subgroups 


Table III presents the numbers of juveniles, adult males and females (the last 
without or with cocoon, indicated with @ @ and @ Qc, respectively) per sample. 


a. Subgroup prativaga 
The phenological data of the different populations of the prativaga subgroup were 


259 


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J. DEN HOLLANDER: Life-histories of Pardosa pullata group 263 


very similar. Small differences were found between the successive years. In 1970 9 Qc 
occurred about 10 days earlier (5”’) than in 1969 (6’). In all the populations of the 
prativaga subgroup more adult females occurred after the emergence of the first pulli 
from the cocoons (July) in 1970 than in 1969. In the former year probably a relatively 
small proportion of these females produced a second cocoon. 


b. Subgroup fulvipes 
The phenological data are rather similar in the different fulvipes populations. In 
population E adults occurred somewhat earlier (4”’) than in population S (5’). In the 
former females produced their cocoon about ten days ahead of population S (5’’’ and 
6’, respectively). These differences existed in both years although they were less 
conspicuous in 1970 than in 1969. 


c. Subgroup pullata 

The phenological data of the populations of the pullata subgroup were rather dif- 
ferent from each other, as compared with those of the subgroups prativaga and fulvipes. 
This was especially true in 1970. The differences concern the time of the final moult, 
the length of the egg maturing period, the time of cocoon construction and the length 
of the period when males occur in the populations. In 1969 the phenological data of the 
populathions HR, BW and Sch were similar. In population AP both males and females 
became adult 10—20 days later (4”’ and 5’, respectively) than in the other populations 
(4”). In all populations 9 Pe appeared in the same period (5”). Thus the length of 
the egg maturing period varied from 10 (AP) to 30 days (HR, BW, Sch). The 
number of males decreased strongly in the beginning of June (6’) in the respective 
populations. This means that the length of the period that males occurred in large 
numbers in the populations varied from 40 (AP) to 50 days (HR, BW, Sch). 

In 1970 the phenological data of population BW were comparable with those in 
1969. However, in the other populations the first adults appeared 10—30 days later 
than in the former one (Sch: 10 days; AP: 20 days; HR: 30 days). Obviously the time 
of the final moult in 1970 occurred 0—30 days later than that in 1969 in the respective 
days (BW: 0 days; AP: 0—10 days; Sch: 10 days; HR: 30 days). 

Again in 1970 the numbers of males decreased strongly in the same period as in 1969 
in all populations (6’). As males became adult rather late in some populations, the 
length of the period that males occurred in the populations varied in 1970 from 20—50 
days (HR: 20 days; AP: 30 days; Sch: 40 days, BW: 50 days). 

As opposed to females in population BW, those in the populations HR, Sch and AP 
constructed their cocoons in 1970 ten days later than in 1969 (5’’ and 5’’’, respectively). 
Thus the length of the egg maturing period varied in 1970 from 10—30 days (HR: 
10 days; BK: 20 days; Sch: 30 days; BW: 30 days). 

In 1970, as compared with 1969, the numbers of 9 Qc were larger in the July and 
August samples. This was especially clear in the population HR. As in the prativaga 
subgroup, more females probably produced a second cocoon in the former year. 

From the above data the following conclusions can be drawn. The time of the final 
moult is variable within the pullata subgroup. Variations occur between different 
populations in one year, as well as in one population between successive years. However, 
construction of cocoons and disappearance of males from the populations occur about 
in the same period in all populations in the two successive years. Thus especially the 


264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


length of the egg maturing period (10—30 days) as well as the length of the period 
that males occur in the populations (20—50 days) varies. The later the final moult 
occurs in spring the shorter both periods are. In this respect the subgroup pullata differs 
clearly from the subgroups prativaga and fulvipes. The final moult in the latter two 
subgroups normally occurs in late spring (5’, 5”). The length of the egg maturing 
period in these subgroups (10—20, and 20—30 days, respectively) equals that of the 
period in those pullata populations, in which the final moult also occurs in late spring. 
The same holds for the period that males occur in the populations in large numbers 
(prativaga: 20 days; fulvipes: 20— 40 days; pullata: 20—50 days). 


C. The number of eggs per cocoon 


Table IV gives the mean number of eggs per cocoon for the respective populations 


Table IV. The mean number of eggs per cocoon in P. pullata. The numbers are given per 
population (at different times during the breeding season) for 1969 and 1970, respectively 
(a: subgroup prativaga; b: subgroup pullata;, c: subgroup fulvipes). 


a: subgroup prativaga 


population data number of cocoons variation mean number of eggs 
VZ 4/6; 15/6-16/6 32, 12 31-72, 27-50 52, 42 
BU 2/6-26/6; 28/5- 5/6 33, 24 32-72, 25-70 52, 47 
LP 19/6-26/6; 5/6-30/6 20, 16 (32-63) 49, 45 
GP 2/6- 9/6; 5/6 10, 13 29-52, 40-70 46, 48 


b: subgroup pullata 


HR 4/6; 26/5 205 18 22-39, 22-40 31, 29 
29/7 18 8-27 18 

BW 23/5- 2/6; 22/5 335122 23-56, 30-54 37, 39 
28/7; 30/6 Mo 18 10-40, 12-39 26, 27 

Sch 5/6; 19/6 B89 16 21-44, 14-39 30, 26 
23/7-30/7 18 7-33 19 

AP 22/5- 5/6; 26/5 26, 20 22-62, 30-54 44, 41 


c: subgroup fulvipes 


S 6/6-13/6; 16/6 26, 36 20-47, 15-49 SAS Sil 
Weg 19/7 23530 8-29, 9-48 100522 

24/8 10 4-16 11 

E 6/6-13/6; 16/6 26, 25 24-54, 19-45 38 Si 
1/7 DARA 10-25, 7-27 Vo We 


1/8; 24/8 10 7-17 12 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 265 


and the successive years. The egg sacs were taken from samples in June, July and 
August. 

Obviously the largest numbers of eggs per cocoon occurred in the subgroup prativaga, 
the smallest numbers were found in the fulvipes subgroup in the cocoons sampled in 
August. The numbers of eggs in the cocoons of the pullata subgroup (sampled in 
June and July) were slightly higher than in the fulvipes subgroup. The numbers in 
the July egg sacs were about 30—40% lower than those in the June egg sacs. Again 
30—40% less eggs were found in the August cocoons than in the July cocoons. Thus 
it seems likely that in those populations of the pwllata subgroup in which cocoons can 
be found during July, these cocoons represent second cocoons. In the same way, cocoons 
found in the populations of the fu/vipes subgroup during July and August represent 
second and third cocoons, respectively. 

The mean number of eggs per cocoon differs only little between the two populations 
of the subgroup fulvipes. The differences between the four populations of the sub- 
group prativaga were also small, although some differences existed between the numbers 
of eggs per cocoon in the successive years, particularly in the population VZ. Rather 
large differences occurred between the populations of the pwllata subgroup. The egg 
sacs in the populations BW and AP contained a larger (30%) number of eggs than 
those in the populations HR and Sch. 

Although the smallest numbers of eggs per cocoon occurred in the f#lvipes sub- 
group, these females produced the largest numbers of offspring. All females produced 
two egg batches and 40% produced a third one. Thus the total number of eggs pro- 
duced by 100 females amounts to about 5500. In the prativaga subgroup the females 
produced a cocoon only once and the total number of eggs produced by 100 females 
thus amounts to about 4700. Only 40% of the females in the pullata subgroup produced 
a second cocoon. The total number of eggs produced by 100 females in this subgroup 
thus amounts to about 4400, the smallest number as compared with that in the sub- 
groups prativaga and fulvipes. 

Concerning these figures, however, it must be taken into account that not all eggs 
in an egg batch develop to pulli, because parasites may destroy some of the eggs in a 
cocoon (Kessler, personal communication). 


D. Conclusions 

The final moult occurs earlier in the pullata subgroup (April) than in the subgroups 
prativaga and fulvipes (May). However, the time of the final moult is variable within 
the pullata subgroup both between populations and successive years. On the other hand 
cocoon production, as well as the disappearance of males from the populations, occur 
in the three subgroups about in the same period (the second half of May and 
the beginning of June, respectively). As a consequence the length of the egg maturing 
period, as well as the length of the period that males occur in the populations, is longer 
and more variable in the pullata subgroup than in the subgroups prativaga and fulvipes. 
The mean number of eggs per cocoon is largest in the prativaga subgroup and smallest 
in the fulvipes subgroup. This number decreases in the successive cocoons which are 
produced during the breeding season. Rather small differences exist between the suc- 
cessive years. The differences between the populations in the pullata subgroup are 
larger than those in the other subgroups. 

On the basis of the occurrence of adult females with and without cocoons, as well as 


266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


the numbers of eggs per cocoon at different times during the breeding season, it is 
concluded that females in the prativaga subgroup produce a cocoon only once. Females 
in the pullata subgroup would produce once or twice a cocoon (variable per population 
and per year). Females in the fz/vipes subgroup normally produce an egg batch two or 
three times in succession. 


2. Growth of juvenile spiders 

In the previous section the assumption was made that females in the subgroup of 
prativaga generally produced a cocoon only once, though in 1970 a minority of the 
females may have produced a second one. In general a minor part of the females in the 
pullata subgroup produced a cocoon twice, whereas in the subgroup fz/vipes all females 
normally produced two cocoons. To obtain more evidence on this point the cephalo- 
thorax length of the juveniles from the autumn and spring samples was measured. 
Graph 1 represents the frequency distributions of the cephalothorax length of juveniles 
in the respective samples. 

It is assumed that every part of a polymodal frequency distribution (when it occurs 
consistently both in autumn and in spring) will represent juveniles emerging from 
cocoons produced in different periods of the breeding season. The number of larval 
instars of the concerned species is unknown. This number is rather variable even at the 
species level and seems to be dependent on environmental factors (Browning, 1941; 
Levy, 1970). 

Generally, spiders moult three times during the larval stage and five to ten times 
during the nymphal stage (Holm, 1940; Juberthie, 1955; Vachon, 1957). In this 
study the successive larval instars as defined by cephalothorax lengths are presented by 
a graph indicating the number of moults, which will occur before the adult stage is 
reached (e.g., adults: stage A; subadults: stage A-1 etc.). Subadults in the three sub- 
groups clearly show sexual dimorphism. As the spiders were collected by hand only 
juveniles of the later larval instars occurred in the samples. 

Graph 1 shows that the cephalothorax lengths of juveniles in the corresponding stages 
are about the same in the three subgroups (stage A-1: 55 MU; stage A-2: 47 MU; stage 
A-3: 40 MU; stage A-4: 33 MU). Increase in cephalothorax length in the successive 
instars thus amounted to 7—8 MU. Pulli emerging from the cocoons showed cephalo- 
thorax lengths of about 20 MU. Thus it is plausible that the cephalothorax length of 
stage A-5 juveniles (which were not represented in the samples) amounts to about 
26 MU and that pulli emerging from the cocoons correspond to stage A-6 juveniles 
(20 MU). In all the three subgroups pulli emerging from the cocoons will moult six 
times before the adult stage is reached. 


A. Subgroup prativaga 

In general juveniles in this subgroup show unimodal cephalothorax lengths in all 
samples (Graph 1a). This means that these juveniles will have emerged from the 
cocoons in the same period. In the autumn the juveniles have grown to stage A-2, i.e., 
four moults occurred after the emergence from the cocoons. These juveniles overwinter. 
Next spring they moult to subadults and afterwards the final moult occurs. 

Especially in the autumn of 1970 small juveniles occurred in the samples from the 
populations BW and LP. These juveniles may have emerged from second cocoons 
produced by some females in these populations (see above). 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 267 


Pulli emerged from the first cocoons in the beginning of July and from the second 
ones in the beginning of August. During September the juveniles from the first cocoons 
moult to stage A-2, those from the second cocoons to stage A-3. Thus four moults after 
emergence from the cocoons take 2—3 months, while three moults take 1—2 months. 


B. Subgroup pullata 


In general the frequency distributions of the cephalothorax length are bimodal 
(Graph 1b). The respective length classes ín the autumn and spring samples represent 
stage A-2 and stage A-1 (subadult) juveniles. The bimodality of the distributions is 
very clear in the populations BW and Sch in 1969 and 1970. This suggests that in 
these populations the juveniles which occurred in autumn emerged from cocoons in 
different periods of the breeding season (stage A-1 from first cocoons in June, stage 
A-2 from second cocoons in July). After overwintering stage A-1 juveniles moulted to 
the adult stage, stage A-2 juveniles moulted to the subadult stage and adult stage, 
successively. 

However, differences exist between the respective populations and between the succes- 
sive years. In spring 1969 the juveniles of population HR showed uniform cephalo- 
thorax lengths (subadult). The same holds for juveniles collected in autumn 1969 and 
in spring 1970. It may be concluded that in this population, in both the breeding seasons 
1968 and 1969, females produced a cocoon only once. On the other hand, in the 
autumn of 1970 juveniles of two length classes occurred in the samples. Therefore, in 
this breeding season at least part of the females evidently produced a second cocoon. The 
same, though to a lesser extent, is true for the population AP. 

The last moult of juveniles before overwintering occurs during September in the 
pullata subgroup. Dependent on emergence from first (the middle of June) or second 
(the end of July) cocoons this moult represents the fifth or fourth moult after the 
emergence of pulli, respectively. Thus the developmental time from pulli (probably 
stage A-6) to subadult (stage A-1) takes 3 months, that from pulli (A-6) to stage A-2 
juveniles 1—2 months. This means that the duration of the successive larval instars of 
juveniles in the pzllata subgroup is shorter than that of juveniles in the subgroup 
prativaga. 


C. Subgroup fulvipes 


The growth of juveniles in the fulvipes subgroup is rather more complex. First, 
juveniles occur during the whole breeding season and secondly, the frequency 
distributions of cephalothorax lengths are polymodal (Fig. 1c). In late summer and 
autumn (August-October) three length classes could be clearly distinguished. Most 
juveniles occurred in the class of the largest cephalothorax lengths (stage A-1, subadult), 
but also stage A-2 and stage A-3 occurred. In spring juveniles of the same three length 
classes occurred. In summer (June-August) juveniles of various cephalothorax lengths 
were found but they could be classed mainly in stage A-2 and stage A-3. 

It is suggested that this rather complex picture may result from the fact that females 
in the fulvipes subgroup produce two or three cocoons successively. After the breeding 
season stage A-3 juveniles appeared in the samples at the end of August. These juveniles 
will have emerged from first cocoons in the beginning of July. They moult to stage A-2 
juveniles in September and in this stage they overwinter. In spring these juveniles moult 
to subadult (stage A-1) and then to adult. The life history of these juveniles thus 


268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


closely corresponds to that of the prativaga subgroup. The developmental time from the 
pulli stage (stage A-6) to the fourth moult afterwards (stage A-2) also amounts to 
2—3 months. 

However, at the same time stage A-3 juveniles still occurred in September. These 
juveniles will have emerged from second cocoons around the end of July. They over- 
winter in stage A-3 and moult to stage A-2 next spring. In this stage they are found 
during the breeding season. In August moulting to subadult (stage A-1) occurs, then 
they overwinter for a second time. Next spring the final moult occurs. The develop- 
mental time of these juveniles from pulli to the third moult afterwards (stage A-3) 
amounts to 1—2 months, as was the case in the prativaga subgroup. 

During May stage A-3 juveniles again appeared in the samples. These juveniles will 
have emerged from third cocoons (in September). They may have overwintered as 
stage A-4 juveniles and moulted to stage A-3 in spring. During July and August these 
juveniles moult once (stage A-2) and again to become subadult (stage A-1). After 
overwintering the final moult occurs next spring. As a result of this specific life history 
of pulli emerging from second and third cocoons juveniles occur in the populations 
throughout the breeding season. 

Concerning these juveniles an interesting phenomenon was found in the population 
E during the breeding season 1970. In June 1970 rather large numbers of subadults 
(stage A-1) were found which showed the abnormality of excessively large cephalo- 
thorax lengths. These lengths even exceeded that of adults. These anomalies may be 
caused by disturbancy of endocrine processes. A period of low temperatures seems to be 
essential in controlling the endocrine processes which regulate the final moult in spiders 
(unpublished results from the department). Probably the chain of these processes was 
disturbed by the exceptionally warm and sunny weather during October 1969 (about 
27° C and 5% above normal, respectively). 

The fact, that fulvipes juveniles which emerge from second and third cocoons do not 
moult after the first overwintering, suggests that the developmental stage during which 
the period of low temperatures occurs is important. The above data show that only when 
overwintering occurs in the last two instars (stage A-2 and stage A-1) before the adult 
stage, the final moult takes place next spring. 


D. Conclusions 


A comparison of the growth of juveniles in the three subgroups gives the following 
results. The number of larval instars after the emergence of pulli from the cocoons 
appears to be the same in the three subgroups. However, the duration of the successive 
larval instars (before overwintering) is shorter in the subgroup pullata than in both 
the subgroups prativaga and fulvipes. In all three subgroups the last moult before over- 
wintering occurs in September, the first moult after overwintering in April (Graph 2). 
The stage in which instars overwinter differs between the subgroups. In the subgroup 
prativaga juveniles overwinter in the instar before the subadult stage (stage A-2). Juven- 
iles in the pullata subgroup overwinter in stage A-2 as well as in the subadult stage (stage 
A-1). Juveniles in the subgroup fz/vipes overwinter in the last four instars (stage A-4, 
A-3, A-2 and A-1). These differences can be understood as a result of differences in 
the duration of the successive larval instars as well as in the time of emergence from 
the cocoons (first, second or third cocoons). The measurements of cephalothorax length 
of juveniles confirm the conclusions stated in the previous section. Females in the sub- 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 269 


Population VZ 


numbers 
15 
9- II 
70 10 
- Population BW 5 
9-IT 0 
4-0 70 
Population LP Population GP 
BL a val 
3 70 sm 
u 
4E ak 
[n] 
Salk 
7 B ub 
69 
Jen 
9-T e li mm 
5 4- II | 
Gela 5 
69 È 
4- II 
B 
5- II 
CI = 
"u fe n “i 
1 ’ Ù 1 Ù n ! a 
30 40 50 60 30 40 50 60 ay 40 50 60 30 40 50 60 
m.u. 


Graph la. The frequency distributions of the cephalothorax length of juveniles during the year. 
Subgroup prativaga 


group prativaga normally produce a cocoon only once. Those in the pullata subgroup 
produce a cocoon once or twice dependent on the population and the year. In the sub- 


group fulvipes females normally produce two cocoons and part of the females produce a 
third one. 


3. The habitats of the studied populations 


A. Description 


The general features of the habitats of Pardosa species have been described e.g. 
by Dahl (1908), Locket & Millidge (1951) and Wiebes (1959). According to 
these authors P. prativaga occurs in fields and marshy soils with a rather dense growth 
of high grasses and other plants. P. pullata would occur in the same localities but also 


270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


15 Population HR 
Population BW 


5 da i 
9-IT 
Population Sch 205 0 
È 
moek opulation AP 8-7 
So 
8 - IT 2 Da 
5 


9-I 


ul 
al 
“al 
A 
© 


Pri 


EB 
30 40 50 60 30 40 50 60 30 40 50 60 30 40 50_ 60 


Graph 1b. The frequency distributions of the cephalothorax length of juveniles during the year. 
Subgroup pullata 


in more dry places. A densly structured low vegetation, e.g. a moss carpet, will mostly 
be present. P. prativaga fulvipes, as far as we know for the Netherlands, is restricted to 
fens, grown with Sphagnum. 

It is evident that the structure of the vegetation influences the temperature in this 
vegetation. Temperature is important for the life history of spiders, especially for the 
time of the final moult, the length of the egg maturing period and the development of 
juveniles (see above, and Jones, 1941). Temperature in the places where the spiders are 
found depends on weather conditions, exposure of the habitat and the structure of 
the vegetation. Observations concerning these features have been made during the 
sampling periods in the populations of the P. pullata group (Table 1, on p. 257). 

The populations LP, BW and GP occur in marshy valleys on the border of calcareous 
dunes. The field LP (prativaga) is sheltered by high shrubs (Salix, Hippophae and 
Betula) interspersed with Calamagrostis and Phragmites. The field GP (prativaga) is 
sheltered only partly by the surrounding shrubs (Salix, Hippophae, Phragmites, Calam- 
agrostis). The vegetation of both fields mainly consists of Salix, Hydrocotyle, Carex and 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 271 


numbers 
15 Population S 
10 
5 10-I 
69 
0 
9-II 
Population E 
10-I 
69 9-IIT 8- III 
a 70 
BS 


= 


9-I 

a ea 8-H | E 
8-II | 

| 


6-Ie Ebummsse 6-1 = Ff Je le 
5- II 5-I 
of aad À PR 


4-0 5-II 4-I 


È 
ba 
r 


a 
N 


RE 
le 


a 1 ! 1 ‘ ‘ 4 
30 40 50 60 30 40 50 60 30 40 50 60 30 40 50 60 
m.U. 


Graph 1c. The frequency distributions of the cephalothorax length of juveniles during the year. 
Subgroup fulvipes 


Parnassia. This vegetation is mown yearly. Thus the habitats of both populations con- 
sist of a high (more than 70 cm) loosely structured vegetation surrounding a low 
(10—20 cm) densely grown over habitat. The spiders occurred especially in the 
transitional area. In spring juveniles were found mainly on the mown field. In autumn 
both adults and juveniles occurred in the surrounding shrubs. 

The field BW (prativaga, pullata) also is yearly mown. It is surrounded by trees and 
shrubs (Betula). The vegetation consists of Lysimachia, Hydrocotyle, Carex and moss 
(Polytrichum). In spring this vegetation is rather low (10 cm) and interspersed with 
bare ground. In summer the Lysimachia has grown up to 40 cm, and thus a loosely 
structured vegetation occurs. The spiders were found all over the mown field. 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


272 


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J. DEN HOLLANDER: Life-histories of Pardosa pullata group 273 


The above mentioned habitats are very wet during spring and autumn. The population 
VZ (prativaga) occurs in a rather dry area throughout the year. This area is situated on 
the border of dunes and arable land and is surrounded by trees (Fagus, Quercus, Popu- 
lus). The vegetation consists of Carex, Calamagrostis, Urtica and Cirsium. In spring 
juveniles mainly occurred on the flattened grass. Adults and, in autumn, juveniles could 
be found in the loosely structured Urtica and Cirsium vegetation (height up to 40 
cm). 

The population Sch (pwllata) occurs in a marshy dune meadow sheltered by shrubs 
and trees (Alnus). This yearly mown field is densely grown with Hydrocotyle, Carex, 
Festuca and moss carpets of Polytrichum and Sphagnum (15 cm). Especially in spring, 
autumn and winter this area is very wet; the spiders occurred throughout the area. 

The population HR (pullata) occurs in a rather dry area surrounded by trees (Pinus). 
The area is densely grown with a moss carpet (Polytrichum) (10 cm) interspersed with 
tussocks of Festuca and Calluna. 

The population AP (pullata) occurs in a yearly mown marshy area grown over 
with a moss carpet (Polytrichum, Sphagnum: 10 cm), Carex, Iris, Hydrocotyle and 
Juncus (30 cm). The area is very wet throughout the year. It is rather exposed in that 
it is not surrounded by trees and shrubs. As in the populations Sch and HR, both 
juveniles and adults occurred throughout the area. 

Both the populations E and S (fulvipes) occur in fens sheltered by trees (Pinus, 
Larix and Betula). The vegetation of both areas consists of a moss carpet (Sphagnum, 
10 cm) interspersed with tussocks of Eriophorum and Erica. The field in which 
population E occurs is more dry with more and larger tussocks of Eriophorum (40 cm) 
than the field of population S. The spiders occurred throughout the field both in the 
moss carpet and in the tussocks of Eriophorum. 


B. Conclusions 


These data show that in general the habitats of the populations of the P. pullata group 
are wet fields sheltered by trees and shrubs. However, differences in soil moisture occur. 
The fulvipes populations (E and S) are found in the wettest habitats. Differences 
between the species exist in the structure of the habitat and in the behaviour of the 
spiders in the vegetation. 

Populations of the prativaga subgroup (LP, BW, GP, VZ) occur in loosely structured, 
rather high vegetation with transitional areas. Adults remain low in the vegetation and 
motionless at low temperatures (ground temperature less than 15° C). With sunny, warm 
weather they walk through and over the plants. Juveniles (stage A-2 and stage A-1) 
show high locomotory activity in spring throughout the still undifferentiated vegetation. 
In autumn they (stage A-2) stay mainly in the dry layers of litter. 

Populations of the pullata subgroup (BW, Sch, AP, HR) occur in densely structured, 
rather low vegetations, generally also including moss carpets. At low temperatures (less 
than 10° C) the spiders remain motionless in small spaces inside the vegetation. During 
warm, sunny weather they leave these spaces and walk upon the vegetation layer. 
Differences in behaviour in relation to the habitat are slight between juveniles (both in 
spring and in autumn) and adults. 

The occurrence of populations of the f#/vipes subgroup (E an S) is restricted to 
fens grown with a Sphagnum carpet. However, they occur in the densely structured 
moss carpet as well as in the loose tussocks of Eriophorum. The behaviour of the 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


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J. DEN HOLLANDER: Life-histories of Pardosa pullata group 275 


spiders of this subgroup in relation to habitat structure is comparable with that of 
the pullata subgroup (moss carpet) as well as with that of the prativaga subgroup 
(Eriophorum tussocks). However, under unfavourable weather conditions adults and 
especially juveniles show higher locomotory activity throughout the year than both 
pullata and prativaga. 


IV. GENERAL CONCLUSIONS AND DISCUSSION 


The results with respect to the phenology of the species of the P. pullata group are 
summarized in Table V. These results agree very well with the data given by Dahl 
(1908), Palmgren (1939), Wiebes (1959) and Vlijm & Kessler-Geschiere (1967). 
Generally the final moult occurs ín spring (April, May) in the three subgroups of the 
P. pullata group. During May the numbers of males and females are highest, indicating 
that the period of copulation then occurs. At the end of May and the beginning of June 
the females produce their cocoons. In the same period the males disappear from the 
populations. Pulli emerge from the cocoons at the end of June and the beginning of 
July. They moult six times before the adult stage is reached. Between every moult, until 
the subadult stage is reached, the juveniles grow about 0.3 mm in cephalothorax length 
from 0.84 mm in the pulli stage to 2.31 mm in the subadult stage. The cephalothorax 
length of adult males and females is different in the three subgroups (Den Hollander, 
1970). 

Some differences in phenology exist between the three subgroups of the P. pullata 
group (Table V). These differences mainly concern the period in which the final moult 
occurs and the growth of juveniles in relation to the period in which they emerge from 
the cocoons. In general both males and females of the pullata subgroup moult to adult 
earlier in spring (about 20 days) than those of the subgroup fulvipes; in the subgroup 
prativaga the final moult occurs about 10 days later than in the subgroup fulvipes. 
However, cocoon production occurs at about the same time in the three subgroups 
(second half of May). Thus the egg maturing period, measured from the moment in 
which the first adult females can be found till the moment in which the first females 
produce a cocoon, is longer in the pullata subgroup than in both other subgroups. 

Even when the final moult occurs later, as happens in some populations in the pullata 
subgroup in some years, the females produce their cocoons at about the same time. Thus 
dependent on the period in which the final moult occurs the length of the egg maturing 
period varies in the pullata subgroup from 10 to 30 days (prativaga: 10—20, fulvipes: 
20—30 days). The cocoon carrying period is longer in the subgroup pullata than in 
both other subgroups (30—40 days and 20—30 days, respectively). This means that 
the period between the appearance of the first adults and the emergence of the first 
pulli from the cocoons (i.e. one reproduction cycle) is different in the three subgroups. 
Generally this period is shortest in the prativaga subgroup (50—60 days). In the sub- 
group fulvipes this period lasts 60—70 days. The reproduction cycle is short (45 days) 
or long (75 days) in the subgroup pullata, dependent on the time of the final moult 
(the later the final moult occurs, the shorter this period lasts). Adult males disappear 
from the populations in the beginning of June in the three subgroups. Because sub- 
adult males moult to adult earlier in the pullata subgroup than in the other sugroups; 
the length of the period that males occur in the populations, is longest in this sub- 
group, but is variable in relation to the time of the final moult. In the prativaga 


276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


subgroup this period normally is shorter than that in both other subgroups. Richter c.s. 
(1971) also found that, under laboratory conditions, the reproduction cycle of P. prati- 
vaga was shorter than that of P. pullata. Because the number of eggs in cocoons of 
prativaga is larger than in pwllata, P. prativaga produces a larger number of offspring 
in a shorter period than P. pullata. Richter c.s. (1971) discussed these results in the 
context of the abundance of the habitats and the use of the habitats by the two species. 
P. prativaga should occur in rather specific habitats and this species might make a 
differential use of its habitat. This means that P. prativaga may be considered as a 
stenotopic species, P. pullata on the other hand would be an eurytopic species. However, 
P. prativaga var. fulvipes can be found only in fens with a Sphagnum carpet (Dahl, 
1908; Holm & Kronestedt, 1970). Thus it may be considered a very stenotopic species. 
Nevertheless, the first reproduction cycle is longer than that in the subgroup prativaga 
and the egg sacs contain considerably less eggs. The length of its reproduction cycle may 
even be longer than that in the subgroup pullata and the number of eggs per cocoon is 
slightly lower than that in the pullata subgroup. Moreover, the mentioned laboratory 
experiments have not shown the variable length of the reproduction cycle in the pullata 
subgroup which sometimes even may be as short as that in the subgroup prativaga. 
However, in this context the microclimatic conditions of the habitat used by the 
respective species may be important. 

Generally, the populations of the P. pullata group occur in two types of habitats. These 
habitats are characterized by a loosely structured, rather high (50 cm) vegetation on 
the one hand (prativaga) and a densely structured, rather low (15 cm) vegetation on 
the other hand (pullata, fulvipes). As a consequence of these different structures the 
characteristics of the microclimate vary between these habitats (Geiger, 1961). In the 
high vegetation a rather stable microclimate occurs. This microclimate is stabilized 
with regard to macroclimate at slightly lower temperatures and higher humidities. 
The microclimate in the low vegetation is very unstable. At sunny, calm weather both 
temperature and humidity of the microclimate are higher than those of the macroclimate. 
However, these conditions change quickly when wind velocity increases and the weather 
is clouded. Especially in spring the microclimate in the low vegetation is unstable 
because weather conditions vary in that time to a large extent. Thus, in the high 
vegetation a rather moderate stabilized microclimate occurs. On the contrary, in the low 
vegetation extreme conditions occur, dependent on the weather. 

These microclimatological features may relate to the phenological characteristics of the 
species. The instability of the microclimate in the habitat of the populations of the 
subgroup pullata (low vegetation) may be related to the variations in the period in 
which the final moult occurs, both between populations and between the successive years. 
Inter-population variation of phenological data was larger in 1970 than in 1969. In 
addition, in most populations the final moult occurred later in 1970 than in 1969. 
Indeed, weather during the spring was very different in these years. In April 1970 
there was 26% less sunshine than in 1969. Probably, the differences in the period of 
the final moult between the respective populations of the pullata subgroup are caused 
by differences in the structure of their habitats. 

The microclimate in the habitat of the populations of the subgroup prativaga (high 
vegetation) is rather stable. Indeed, the final moult occurred very synchronized in the 
respective populations and in the same period in the successive years. The habitat of the 
populations of the fz/vipes subgroup is to some extent intermediate between that of the 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 277 


prativaga subgroup and that of the pullata subgroup. Both high and low vegetations 
occur in this habitat. Although the number of populations studied is too small for a 
clear conclusion, the final moult in the fulvipes subgroup seems to be less synchronized 
than that in the prativaga subgroup, but more synchronized than that in the pwllata 
subgroup. 

As most phenological characteristics relate to the period in which the final moult 
occurs, they are influenced by the microclimatological characteristics. Thus it seems 
plausible that variations in phenological characters of the species in the P. pullata group 
are related to the type of habitat in which they occur. 

In this context the growth of juveniles is important. In the prativaga subgroup pulli 
emerge from the cocoons in the beginning of July only. They grow and overwinter as 
stage A-2 juveniles. Next spring they moult to subadult and adult (Graph 2a). 

In the pullata subgroup pulli emerge from the cocoons in the second half of June 
(first cocoon) and in the second half of July (second cocoon), these juveniles over- 
winter as stage A-1 juveniles (first cocoon) and stage A-2 juveniles (second cocoon). 
(Graph 2b). 

In the fulvipes subgroup females produce three cocoons in succession. Pulli emerge 
from these cocoons in the beginning of July (first cocoon), the beginning of August 
(second cocoon) and in the beginning of September (third cocoon), respectively. The 
juveniles which have emerged from the first cocoons grow to the adult stage as 
juveniles in the subgroup prativaga do. Those which have emerged from the second and 
third cocoons, however, overwinter twice before the final moult occurs. The first 
overwintering occurs during stage A-3 and stage A-4 for juveniles emerged from the 
second and third cocoons, respectively. The second overwintering takes place during 
stage A-1 (subadult) for both categories of juveniles (Graph 2c). 

These results show that only juveniles which overwinter during the last two instars 
before the adult stage moult to adults next spring. Laboratory experiments (unpublished 
results) have shown that a period of low temperatures is necessary to induce the final 
moult. The present results show that such an induction only can occur during the last 
two instars before the final moult. Probably endocrine processes have to be started by 
a period of low temperatures during this developmental stage (Eckert, 1967; Streble, 
1966). When these processes have been started the juveniles moult once or twice in 
succession to become adult during spring. In the other case they moult only once and 
occur in this stage in the populations during the summer. However, juveniles in the 
pullata subgroup normally moult to adult earlier in spring than juveniles in both other 
subgroups. Probably an external factor induces the time of the final moult. Temperature 
may be this factor. If this is true, then it may be concluded that the final moult in the 
pullata subgroup is already induced at lower ne than in the subgroup prativaga 
and fulvipes. 

The numbers of eggs produced per female are rather similar in the three subgroups 
(prativaga: 47; pullata: 44, fulvipes: 55). However, this number is produced in the 
prativaga subgroup in one egg batch, whereas in the pullafa subgroup this number is 
produced partly in two egg batches and in the fu/vipes subgroup in three egg batches. 
This phenomenon may represent a mechanism of spreading of the risk (Den Boer, 
1968). The habitat of P. prativaga var. fulvipes is very unstable, that of P. prativaga 
is rather stable. Thus P. prativaga var. fulvipes produces the total number of eggs in 


278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


three periods, P. prativaga in one period. The stability of the habitat of P. pullata is 
about intermediate: the total number of eggs is produced in one or two periods. 

Using the classification of Southwood (1962) and Johnson (1969), Richter (1970) 
mentioned the habitat of P. pullata as abundant with intermediate stability. The habitat 
of P. prativaga would be rare and unstable. The same seems to be true for P. prativaga 
fulvipes to a larger extent. However, no clear relations seem to exist between the total 
number of eggs produced per female, in one, two or three egg batches, and the 
abundance and stability of the habitat. 

The above mentioned results also may be discussed in the context of the classification 
of the species in the P. pullata group. Dahl (1908), Simon (1932), Wiebes (1959), 
Tongiorgi (1966) and Holm & Kronestedt (1970) describe various species and sub- 
species within the P. pzllata group. This division is mainly based on small differences 
in the structure of the external genital organs, the annulation and spinosity of the legs 
and the size and colour of the specimens. However, Den Hollander (1970) has shown 
that some of these characters vary to a rather large extent, both within and between 
populations of the same species (P. pullata, P. prativaga and P. prativaga var. fulvipes). 
He also suggests the occurrence of hybridization in mixed populations of P. pullata and 
P. prativaga. Indeed, under laboratory conditions, intermediate specimens have been 
obtained from males of P. prativaga and females of P. pullata (Den Hollander, in 
preparation). Thus the distinction between the various species and subspecies of the 
P. pullata group seems to be rather doubtful. Indeed, the phenological differences 
between the species studied by me are rather small. Rather large variations could 
be shown between populations of the same species (P. pullata). Probably the 
phenological differences as well as the variation of phenological characteristics may 
relate to specific characteristics of the habitats used by the respective species. 

On the other hand, various species of the P. pullata group occur in mixed populations. 
In France mixed populations of P. pullata and P. prativaga, P. pullata and P. femoralis, 
and of P. prativaga and P. femoralis have been observed. In the Netherlands mixed 
populations of P. prativaga and P. pullata, and of P. pullata and P. prativaga var. fulvipes 
occur (Den Hollander, 1970, and unpublished results). Thus the differences between 
the species may also result from a differential use of the habitat by the respective 
species. The evolution of the P. pullata group seems to be such that various species have 
originated which at present can be mainly separated only by their ecological 
characteristics. The development of morphological, ethological and physiological dif- 
ferences seems to be in progress (ecological speciation, Grant, 1963). 

It should, however, also be mentioned that populations of the P. pullata group may 
have been isolated geographically from each other, e.g. during ice-ages. During this time 
of isolation characters may have been acquired which promote or guarantee reproductive 
isolation when the external barriers break down (Mayr, 1942). In the case of the 
P. pullata group, the development of these characters may be, as yet, incomplete. 

Whatever the mechanisms of speciation (ecological or geographical) may have been, 
the various species have been adapted to special types of habitat. As a consequence, 
isolating characters have been evolved. 

Further studies concerning these problems are necessary for further evaluation of 
the relations between the species of the P. pullata group. 

The occurrence of hybridization between closely related species in mixed populations 
may be prevented by several mechanisms (Mayr, 1969). Mating between males and 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 279 


females of different species. may be prevented by seasonal or habitat isolation, ethological 
isolation and mechanical isolation. The results of the present study deal with the 
occurrence of seasonal isolation in mixed populations of P. prativaga and P. pullata. 
Preliminary laboratory experiments have shown that ethological and mechanical barriers 
do not exist between males of P. prativaga and females of P. pullata. Ethological 
barriers seem to prevent mating of males of P. pullata with females of P. prativaga 
(cf. Den Hollander, in preparation). 

The results of this phenological study have shown that males of P. pullata and 
receptive females of P. prativaga normally occur together in mixed populations. The 
reversed combination of males and females does not occur. Thus in mixed populations 
the species P. prativaga is isolated from the species P. pullata, partly be seasonal barriers 
and partly by ethological barriers. However, the seasonal barriers can be broken down 
by cold weather during spring. Then the final moult in P. pullata is postponed and 
males of P. prativaga occur together in the populations with receptive females of 
P. pullata. Generally, the two species use a different type of habitat. Indeed, even in 
mixed populations they seem to use different parts of the habitat. However, this is 
especially true for females carrying cocoons, while females without cocoons, ie, 
receptive females, as well as males seem to use the habitat less differentially (Den Hol- 
lander & Lof, in preparation). These results show that habitat isolation, seasonal 
isolation, as well as ethological isolation, exist between the species concerned but that 
each of these barriers seems to be incomplete. Thus hybridization in mixed populations 
between males of P. prativaga and females of P. pullata may occur when, because of the 
weather conditions in spring, the final moult in P. pullata is postponed till the begin- 
ning of May. 


Postscript. According to Mr. Kronestedt who kindly studied a number of 
specimens of each of the investigated populations, the form P. prativaga var. fulvipes 
(Collett, 1875) correctly should be named P. sphagnicola (Dahl, 1908). (Holm & 
Kronestedt, 1970). 


V. SUMMARY 


1. Ten populations of the Pardosa pullata group (P. prativaga: 4; P. prativaga var. 
fulvipes: 2; P. pullata: 4) were sampled once every ten days from April till September 
in 1969 and 1970. 

2. The numbers of juveniles, adult males and females (the last with and without 
cocoon) in every sample were established. The length of the cephalothorax of the 
juveniles in every sample was measured. On the basis of these data the phenology of 
the species of the P. pullata group was studied. 

3. Generally the final moult occurs earlier in spring in P. pullata, than in both P. 
prativaga and P. prativaga var. fulvipes. However, the time of the final moult varies in 
P. pullata per population and per year. 

4. First cocoon production occurs in about the same period in the studied populations 
in both years. 

5. In P. prativaga females normally produce a cocoon only once; in P. pullata they 
produce a cocoon once or twice, and in P. prativaga var. fulvipes the females produce 
two or three cocoons in succession. 


280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


6. The development of juveniles from pulli to the adult stage is somewhat different 
in the different species. 

7. The numbers of eggs per cocoon in P. prativaga are larger than those in both 
P. pullata and P. prativaga var. fulvipes. When females produce two or more cocoons in 
succession, the number of eggs per cocoon is about 40% lower in each following egg 
batch. 

8. The differences in the period in which the final moult occurs in populations of 
the P. pullata group are discussed in the context of microclimatological characteristics 
of the habitat. 

9. As yet, no clear relationships seem to exist between the total number of eggs 
produced per female, in one, two or three egg batches, and the abundance and stability 
of the habitats. 

10. The phenological data have been discussed in the context of the classification of 
the P. pullata group as well as in the context of the possibility of the occurrence of 
hybridization in mixed populations. 


VI. ACKNOWLEDGEMENTS 
I am deeply indebted to Prof. Dr. L. Vlijm for his valuable criticism during the 


investigation and the preparation of the manuscript, to Mr. G. W. H. van den Berg 
for drawing the graphs and to Miss M. J. Diesbergen for typing the manuscript. 


VII. REFERENCES 


Browning, H. C., 1940. — The relation of instar length to the external and internal environment 
in Tegenaria atrica (Arachnida). — Proc. Zool. Soc. London 110: 303—317. 
Boer, P. J. den, 1968. — Spreading of risk and stabilization of animal numbers. — Acta Biotheore- 


tica 18: 165—194. 

Dahl, F., 1908. — Die Lycosiden oder Wolfspinnen Deutschlands und ihre Stellung im Haushalte 
der Natur. — Nova Acta Acad. Caes. Leop.-Carol. 88: 177—558. 

Eckert, M., 1967. — Experimentelle Untersuchungen zur Häutungsphysiologie bei Spinnen. — Zool. 
Jb. Physiol. 73: 49—101. 

Geiger, R., 1961. — Das Klima der bodennahe Luftschicht. — Die Wissenschaft 78: 1—646. 
Vieweg, Braunschweig. 

Grant, V., 1963. — The origin of adaptations. — New York: Columbia University Press. 

Hollander, J. den, 1970. — Morphological variation in Pardosa prativaga L. Koch, 1870, P. grati- 
vaga var. fulvipes Collett, 1875, and P. pullata Clerck, 1757 (Araneae, Lycosidae). — 
Tijdschr. Entomol. 113: 273—288. 

—., Hybridization experiments in the Pardosa pullata group (Araneae, Lycosidae). — In 
preparation. 

Hollander, J. den & H. Lof. — Differential use of habitat in a mixed population of Pardosa 
prativaga and P. pullata. — In preparation. 

Holm, A., 1940. — Entwicklung und Entwicklungsbiologie der Spinnen. — Zool. Bidrag Uppsala 
29: 1—214. 

Holm, A. & T. Kronestedt, 1970. — A taxonomic study of the wolfspiders of the Pardosa pullata- 
group (Araneae, Lycosidae). — Acta ent. bohemoslov. 67: 408—428. 

Johnson, C. G., 1969. — Migration and dispersal of insects by flight. — London: Methuen. 

Jones, S. E., 1941. — Influence of temperature and humidity on the life history of the spider 
Agelena naevia (Agelenidae). — Ann. Ent. Soc. Am. 34: 557—571. 

Juberthie, C., 1955. — Sur la croissance post-embryonaire des Aranéides. — Bull. Soc. Hist. Nat. 
Toulouse 90: 83. 


J. DEN HOLLANDER: Life-histories of Pardosa pullata group 281 


Levy, G., 1970. — The life cycle of Thomisus onustus (Thomisidae: Araneae) and outlines for 
the classification of the life histories in spiders. — J. Zool. 160: 523—536. 

Locket, G. H. & A. F. Millidge, 1951. — British spiders, vol. 1. — London. 

Mayr, E., 1942. — Systematics and the origin of species. — Columbia University Press, New York. 

, 1969. — Animal species and evolution. — Harvard University Press, Cambridge. 

Palmgren, P., 1939. — Die Spinnenfauna Finnlands. I. Lycosidae. — Acta Zoologica Fennica 25: 
5— 86. 

Richter, C. J. J., 1970. — Aerial dispersal in relation to habitat in eight wolf spider species (Par- 
dosa, Aranaea, Lycosidae). — Oecologia (Berl.) 5: 200—214. 

jRichter, C. J. J., J. den Hollander & L. Vlijm, 1971. — Differences in breeding and mobility 
between Pardosa pullata (Clerck) and Pardosa prativaga (L. Koch) (Lycosidae, Araneae) 
in relation to habitat. — Oecologia 6: 318—327. 

Simon, E., 1932. — Les Arachnides de France 6 (4): 1047—1141. — Paris. 


Southwood, T. R. E., 1962. — Migration of terrestrial arthropods in relation to habitat. — Biol. 
Rev. 37: 171—214. 
Streble, H., 1966. — Untersuchungen über das hormonale System der Spinnentiere (Chelicerata) 


unter besonderer Berücksichtigung des „endokrinen Gewebes” der Spinnen (Aranea). — 
Zool. Jb. Physiol. 72: 157—234. 

Tongiorgi, P., 1966. — Italian Wolfspiders of the genus Pardosa (Araneae: Lycosidae). — Bull. 
Mus. Comp. Zool. 134: 275—334. 

Tretzel, E., 1954. — Reife und Fortpflanzungszeit bei Spinnen. — Z. Morph. Okol. Tiere 42: 


634— 691. 
Vachon, M., 1957. — Contribution à l'étude du développement post-embryonaire des araignées. I. 
Généralités et nomenclature des stades. — Bull. Soc. Zool. France 82: 337—354. 


Vlijm, L. & A. M. Kessler-Geschiere, 1967. — The phenology and habitat of Pardosa monticola, 
P. nigriceps and P. pullata (Araneae, Lycosidae). — J. Anim. Ecol. 36: 31—56. 

Vlijm, L., A. Kessler & C. J. J. Richter, 1963. — The life history of Pardosa amentata (Cl.) 
(Araneae, Lycosidae). — Ent. Ber. 23: 75—80. 

Wiebes, J. T., 1959. — The Lycosidae and Pisauridae (Araneae) of the Netherlands. — Zool. 
Verh. 42: 1—78. 

———, 1960. — De Wolfspinnen van Meijendel. — Ent. Ber. 20: 59—62, 69—74, 83—89. 


| ty MT 


rie deal 
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MES un 


REGISTER VAN DEEL 114 


* Een sterretje duidt een naam aan nieuw voor de wetenschap. 

* An asterisk denotes a name new to science. 
** Twee sterretjes duiden de naam aan van een voor de Nederlandse fauna nieuwe soort. 
** Two asterisks denote the name of a species new to the Netherlands fauna. 


Uit dit register zijn weggelaten de namen van Odonata, voorkomende in: ,,A catalogue of the 
type-specimens of Odonata...’ etc., pp. 65-139 van dit deel. 


PAGA TI Eee cia ns en hea a ow ise ue 283 Etymenopterave: NN 785 
Ardon laan We nee en a 1283 Bepidopterar. Sr MR RR 2 85) 
Coleoptera en ale een 283 Mammal e E 25 6 
Diptera TE) Odonata stereo iten 236 
Plantes: o acy te ae) RAC Leur eu 280 
ACARI Pardosa 256 etc. Atypophthalmus 223 
prativaga 256 etc. Austrolimnophila 224, 233 
Amorphacarus 163, 171, 172 pullata 256 etc. autumnalis 223 
anchora 183 sphagnicola 279 barbipes 224 
Archemyobia 171 batava 225, 235 
auricularis 172 COLEOPTERA benesignata 221 
*baueri 190, 200 Beringotipula 222 
betulinus 183 *bacchusi 241, 248 bicolor 225, 235 
Blarinobia 172 *brunnea 241, 242 bicornis 227, 237 
cinereus 200 *difficilis 241, 245, 248 bidens 221, 230 
desmanae 171 Digenethle 239, 241 bifasciata 222 
dispar 182 Microlomaptera 241 bifida (Gonomyia) 226, 236 
Eadiea 171 *signifer 241, 250 **bifidus (Molophilus) 227, 
eliomydis 199 Tafaia 239, 241 {237 
elongatus 167, 169, 171, 172 viridiaenea 239 bifurcata 227 
Gliricoptes 183 bihamatus 227, 237 
hengererorum 169, 171 DIPTERA bimaculata (Ctenophora) 220 
Hypodectes 181 bimaculata (Pedicia) 224 
inexpectata 171 abdominalis 225, 235 bivittata 227, 237 
krameri 183 Achyrolimonia 223 Brachylimnophila 225 
Laphuromyopus 183 aculeata 220 brevitarsis 224 
macdonaldi 183 Acutipula 221 brunneinervis 221 
Melesodectes 173, 175, 183 Acyphona 227 bullata 221 
microti 200 Adelphomyia 225 caesia 220 
multisetosa 171 adjuncta 225 calceata 223 
*muscardinus 183, 199 albipes 220 carteri 226 
Myialges 183 albitibia 227, 237 cava 221 
*neerlandicus 192, 199 alboscutellata 226, 236 Cheilotrichia 226 
Orycteroxenus 182 alpina 221, 229 Chionea 227 
*parvisetosus 163, 169, 171 alpium 221, 230 chorea 223 et 
Pontoppidania 175 Amalopis 224 cinerascens (Cheilotrichia) 
propus 181 analis 220, 228 [226 
Protomyobia 171 angustipennis 224, 234 cinerascens (Dicranoptycha) 
Psorergates 185 anomaloptera 221 [2 
Pteracarus 171 aperta 225, 235 cinereifrons 227 
*quercinus 195, 199 appendiculata (Nephrotoma) clara 226 
Rodentopus 183 [220 **claripennis 224, 233 
sorici 200 appendiculatus (Molophilus) clavata 227, 237 
talpae 182 [227, 228, 237 coerulescens 220, 229 
Xenoryctes 183 aquosa 223 ssp. complicata 223, 232 
Arctoconopa 227 cornicina 220 
ARANEIDAE areolata 227 corniger 227 
armatus 227 couckei 220 
femoralis 278 *arnoudi 224, 234 crassicauda 224 
var. fulvipes (P. prativaga) **ater 227, 237 crocata 220 
[256 etc. atrata 220 Crunobia 224 


283 


284 


Crypteria 226 

Ctenophora 220 

ctenophora 223, 232 
**curvatus 227, 237 

Cylindrotoma 222 

czizeki (Molophilus) 227, 


[237 


czizeki (Prionocera) 220 
dalei 225, 235 
danica (Limonia) 223, 231 
danica (Ormosia) 227 
Dasymolophilus 228 

**decemmaculata 223, 232 
Dendrotipula 222 
dentata 226 
depilata 227, 237 
Dicranomyia 223 
Dicranoptycha 223 
Dicranota 224, 233 
Dictenidia 220 
didyma 223 
dilatata 221 
dilutior 222 
dimidiata 225 
Diogma 222 
discicollis 225 
dispar 225 
distendens 223, 231 
distinctissima 222 
divisa 226, 236 
Dolichopeza 220 
dorsalis 220 
dumetorum 223 
duplicata 223 
elegans 220 
Elliptera 223 
Ellipteroides 225 
Eloephila 225 
Epiphragma 224, 233 
Erioptera 226 
errans 226, 236 
Eutonia 224 

**excisa 221, 229 
falciger 228 
fasciata 225 
fascipennis (Ormosia) 227 


fascipennis (Tipula) 221, 229 


ferruginea 225 

festiva 220 

filata 225 

Flabellifera 220 

flava (Rhabdomastix) 226 
flaveolata 220 


flavescens (Erioptera) 226 
flavescens (Nephrotoma) 220, 
[228 


flavipalpis 220 
flavipes 222, 231 
flavolineata 222, 231 
flavus (Helius) 224 


flavus (Molophilus) 227, 237 


frontalis 223, 231 
fulvipennis 221 
fulvonervosa 225 
fusca 223, 231 


fuscescens 223, 232 
fuscipennis (Erioptera) 226 
fuscipennis (Pilaria) 225 
**fusculenta 226, 236 
gaedii 225, 235 
gemina 226 
glabrata 222 
**glabricula 225, 235 
gladius 228 
gmundensis 224 
Gnophomyia 226 
Gonempeda 226 
Gonomyia 225, 226 
gracilipes 224 
griseipennis 226 
grisescens 222 
griseus 227 
guestfalica 220 
guttata 220, 228 
**haemorrhoidalis 227, 237 
hederae 227 
Helius 224 
helvola 221, 229 
hercegovinae 222 
heterogyna 225, 235 
Hexatoma 225 
hortensis 222 
hortorum 221 
hybrida 227 
Idioptera 225 
imbuta 226 
immaculata 224 
incisurata 226 
inconstans 224 
inusta 223, 232 
irrorata 222 
juncea 222 
laeta 226 
lateralis (Gonomyia) 226 
lateralis (Prionocera) 220 
leucophaea 225 
limbata (Erioptera) 226, 236 
limbata (Tipula) 221, 230 
limitata 221, 229 
Limnophila 224, 234 
limnophiloides 226 
Limonia 222 
lindneri 220, 228 
lineata 227 
lineonella 225 
Lipsothrix 226 
littoralis 224, 233 
livida 221, 230 
longidens 221, 230 
longirostris 224 
longitarsis 224 
lucida 223, 231 
lucidipennis 224, 233 
lucidula 226, 236 
lucorum 224, 233 
Ludicia 224 
lugubris 226 
luna 221 
lunata 221, 230 
Lunatipula 221 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


lunulicornis 220 

lutea 226, 236 

luteipennis (Limonia) 223 
luteipennis (Prionocera) 221 
lutescens 227, 236 
macrocera 222 
macrophthalma 226 
macroptera 224 
macrostigma 222 

maculata (Gonomyia) 227 
maculata (Limnophila) 225 
maculicosta 223, 231 
magnicauda 221, 230 
marginata 220 

marmorata 225 

maxima 221 

Mediotipula 221 

medius 227 

Megarhina 224 


**meigeni (Limnophila) 225, 


[235 


**meigeni (Tipula) 222, 230 


meijeri 226, 236 
Melampodia 227, 236 
melanoceros 221 
Melanolimonia 223 
melleicauda 223, 232 
Metalimnobia 222 
minor 226, 236 

mitis 223, 232 
modesta 223, 232 


** mollissima 224, 233 


Molophilus 227 
montium 221 

morio 223 

mundata 225, 234 
murina (Ormosia) 227 
murina (Taeiocera) 228 
nemoralis 225, 235 
Neolimnomyia 225 
Neolimnophila 226 
Neolimonia 223 
Nephrotoma 220, 228 
neurotica 222 

nielseni (Erioptera) 226 


**nielseni (Paradelphomyia) 


[225, 236 
niger 227 
nigra 221 
nigricornis 220 
nigripalpis 226, 236 
nigropunctata 223 
Nigrotipula 221 
nodulosa 227 
nubeculosa (Limonia) 223 
nubeculosa (Hexatoma) 225, 
[236 
nubeculosa (Tipula) 221, 229 
nubila 226 
obscurus 228 
obsoleta 222 
occoecata 227, 236 
occulta 224 
occultus 228, 237 
ocellaris 224, 233 


ochracea 224, 233 
ochraceus 228 
ochrescens 228 
Odonatisca 222 
oleracea 220, 229 
omissa 223, 232 
omissinervis 223, 232 
Ormosia 227 


ornata (Ctenophora) 220, 228 
ornata (Limonia) 223, 232 


Oxyrhiza 225 
pabulina 222 
Pachyrhina 220 
pagana 222 

Pales 220 

paludosa 220, 229 
Paradelphomyia 225 
Paradicranota 224 
pavida 224, 233 
pectinicornis 220 
Pedicia 224, 233 
peliostigma 221 
pentagonalis 227, 237 
phaeostigma 225, 235 
Phalacrocera 222 
Philidorea 225 
pictipennis 224 
pierrei 221 

Pilaria 225 

pilipes 226 

placida 226 
platyptera 225, 235 
Platytipula 221 
Platytoma 226 
pleuralis 228 
pratensis 220 
Prionocera 220, 228 
propinquus 228 
pruinosa 221, 229 
pseudoirrorata 222 
pseudolateralis 221 


Pseudolimnophila 224, 233 


pseudosimilis 227 


pseudovariipennis 222, 230 


Psiloconopa 227 
Pterelachisus 222 
pubescens 220, 228 
pulchella 225, 234 
punctata 224 
punctipennis 227 
punctum 225, 235 
pustulata 221, 230 
quadrifaria 220, 228 


**quadrimaculata 222, 231 


quadrinotata 222 
quadristriata 220, 228 
quadrivittata 221 
recta 226, 236 
remota 226, 236 
replicata 222 
Rhabdomastix 226 
Rhamphidia 224 
Rhipidia 223 
Rhypholophus 227 
rivosa 224 


REGISTER 


rubripes 221 
rufina 222 
Savtshenkia 221 
saxonum 225 
scalaris 220, 228 
schineri 224 
schummeli 224, 233 
Schummelia 221 
Scleroprocta 227 
scripta 221, 229 
scurra 220, 228 
scutellata 225 
Secandaga 226 
selene 221 
selenis 225 
sepium 224 
sera 223 
sericata 223, 232 
serpentiger 228 
sexmaculata 224, 233 
signata 222, 230 
similis 227 
simplex 226 
solstitialis 221 
**sororcula 227, 237 
squalens 225, 235 
squalida 226 
staegeri 222 
staegerina 227, 237 
stictica 227 
stigma 223, 231 
stigmatella 221, 230 
stigmatica 223, 232 
**straminea 224, 233 
Stygeropsis 220 
submaculosa 220, 228 


**submarmorata 225, 235 


subnodicornis 222 
subserricornis 220 
subtilis 224 
subtincta 225 
subvafra 222, 230 
sylvatica 224, 233 
sylvicola 223, 231 
Symplecta 227 
Tanyptera 220 
Taphrophila 223 
Tasiocera 228 
taurica 223, 231 
tenella 226 
Thaumastoptera 223 
Tipula 220, 229 
transversa 224 
Tricyphona 224 
trimaculata 225, 235 
Trimicra 226 
Triogma 222 
tripudians 226 
tripunctata 223, 231 
tristis 223, 232 
trisulcata 222 
trivialis 226 
trivittata 223 
truncorum 222 
turcica 220 


Ula 224, 233 
unca 222 
uncinata 227 
undulatus 228 
unicolor 224, 233 
uniseriata 223, 232 
varia 227 
variicornis 221 
varipennis 222 
ventralis 223, 232 
vernalis 221 
verralli 225, 235 
Vestiplex 221 
vicina 227 
viridipennis 226 
vitripennis 223 


285 


vittata (Hexatoma) 225, 236 


vittata (Tipula) 221 
winthemi 222, 231 
Yamatotipula 220, 229 
yerburi 221, 229 
zernyi 221, 229 
zetterstedti 222, 231 
zonaria 221 


HYMENOPTERA 


bellitarsis 203 
Buathra 201, 202 
Cryptus 202, 209 
curvicauda 203 
divisoria 202, 205 
fabricii 204 
fulvipes 202 
glabricula 214 
gradaria(us) 215 
hellenicus 209 
Ichneumon 202 
immitis 209 
laborator 202, 204 
medius 202 
Meringopus 202 
*priesneri 213 
punctatus 215 
reverendus 202 
tarsoleuca 202, 203 
tristator 213 
Trychosis 213 
*verutus 209 


LEPIDOPTERA 
Adeloneivaia 158 


affinis 159 
albilinea 151 


*ssp. anchicayensis (E. im- 


[perialis) 154 


*andensis 142 
andina 156 
andrea 157 
Anisota 153 nota 


ssp. approximans (E. im- 


[perialis) 153, 154 


**ssp. araguensis (C. avenata) 


[149 


286 


aroa 156 
avenata 149 
bellavista 156 
ssp. bidens (B. ocellata) 158 
f. bouvieri (D. tarquinia) 147 
Bouvierina 157, 158 
ssp. brasiliensis (C. mexicana) 
{156 nota 
brissoti 154 
ssp. cacicus (E. imperialis) 
[153, 154 
canella 143 
Ceratocampa 153 nota 
Cerodirphia 148 
Citheronia 153 nota, 154, 156 
consobrina 156 nota 
Copaxa 141-144 
Copiopteryx 152 
*crassifurca 148 
*dargei 159 
ssp. decoris (E. imperalis) 
[153, 154 
*denhezi 143 
*descimoni 159, 161 
Dirphia 147, 148 
Dryocampa 153 nota 
dunkinfieldi 151 
Eacles 153 
equatorialis 156 
*escalantei 141 143 
expandens 142 
ssp. flavina (Cop. canella) 
{143 
*gadouae 157 
*ssp. gadouorum (C. 
[semiramis) 152 
Galleria 205 
*herbuloti 144 
hypoxantha 159 
imperialis 153 
jason 158 
*ssp. jasonoides (B. ocellata) 
[158 
joinvillea 143 
Kentroleuca 151 
lebeau 146 
*lichyi (Citheronia) 154 
*lichyi (Dirphia) 147, 148 
lilacina 159 
lineosa 151 
ssp. magnifica (E. imperialis) 
[153, 154 
*marahuaca 148 
mellonella 205 
mexicana 156 nota 
*ssp. miranda (Cop. canella) 
{143 
mogya 156 nota 
ssp. montei (C. semiramis) 
[152 
mota 149 
*mulierata 151 
multifenestrata 142, 143 
“nebulosa 158 


ssp. nobilis (E. imperialis) 
[153 
ocellata 158 
ssp. opaca (E. imperialis) 
[153, 154 
ssp. oslari (E. imperialis) 153 
*paucidentata 144 
ssp. pini (E. imperialis) 153 
Ptiloscola 159, 161 
quadrilineata 157 
*ssp. quintanensis (E. imperia- 
[lis) 153, 154 
Rothschildia 144 
sapatoza 144 
Scolesa 158 
semiramis 152 
sepulcralis 154 
simson 143 
smithi 157 
speciosa 149 
*spitzi 151 
subfumata 152 
surrotunda 162 
tarquinia 147, 148 
totoma 159 
ssp. travassosi (C. semiramis) 
[152 
Travassosula 151 
trimacula 142 
ssp. tucumana (E. imperialis) 
[153, 154 
wellingi 161 
*ssp. yucatana (R. lebeau) 146 


MAMMALIA 


Eliomys quercinus 185, 199 

Labidophorus 182 

Meles meles 173, 181 

Microtus oeconomus 195 

Muscardinus avellanarius 190 

Neomys fodiens 163, 169, 
[192, 200 

Sorex araneus 171, 172 

Talpa europaea 182 


ODONATA 


amymone 31, 32 
arachnomima 28, 29 
artemis 14 
*asachinai 18, 21 
*astarte 31, 42 
atrovirens 18 
australiae 14 

ssp. borneensis (J. dohrni) 4 
callisto 23, 29 
*campioni 48, 55 
celaeno 31, 35 
chalciope 32, 33 
Chlorogomphus 21 
cincta 29 

claudia 8 

corycia 29 

cydippe 29 


TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 114, AFL. 8, 1971 


Didymops 1 

*dione 26, 29 
dohrni 4 
donaldi 14, 16, 18 
Epophthalmia 1, 4 
erato 29 
ssp. erratica (genialis) 18, 20 
euphrosyne 31 
eurynome 31, 33 
euterpe 29 
fenella 53, 54, 57 
flexicauda 55 
fulva 18 
ssp. fumata (moorei) 29 
genialis 18, 19 
gerstaeckeri 22, 29 
Hemicordulia 14 
hermione 31 
Idionyx 2, 3, 4, 17 
Idiophya 2 
Indomacromia 16 
jucunda 29 
lachesis 31, 38 
laidlewi 4 nota 
macromia 1, 4, 5, 6 
Macromidia 1, 2, 3, 4, 5, 16 
magnificus 21 
melpomene 31, 41 
Metaphya 5 
miranda 47, 54, 55 
mnemosyne 29 
montaquei 51, 55 
montana 4, 7, 14, 16 
moorei 29 
numfordi 14 
murcia 4, 6, 12 
nilgiriensis 2, 3 
optata 4 nota 
orchestra 4, 7 
philippa 3, 4, 7, 8, 18 
Phyllomacromia 2 
polyhymnia 29 
Procordulia 14 
rapida 18 
salva 3, 8 
samal 18 
sambawana 14 
septima 29, 30 nota 
shanensis 18 
sophrosyne 31 
splendens 30 nota 
Synthemis 2, 45 
terpsichore 31, 32, 35 
viridescens 31, 41 
westwoodi 29 
yolanda 4, 7, 14 


PLANTAE 


Alnus 217 

Betula 217, 271, 273 
Calamagrostis 271, 273 
Calluna 273 

Carex 271, 272, 273 
Carpinus 217 


Cirsium 273 

Erica 273 

Eriophorum 273, 275 
Fagus 217, 273 

Festuca 273 

Hippophae 271 
Hydrocotyle 271, 272, 273 


REGISTER 


Iris 273 

Juncus 273 
Larix 273 
Lysimachia 272 
Parnassia 272 
Pinus 273 


Polytrichum 272, 273 
Populus 217, 273 
Quercus 217, 273 
Salix 217, 271 
Sphagnum 271, 273 
Urtica 273 


287 


No, 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende 
gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.F, 45.—. 


No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), 
(The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.Fl. 70.—. 


No. 4. F. Willemse, — Preliminary revision of the genera Stenocatantcps Dirsh & Uvarov 
and Xenocatantops Ditsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. 
D.FI. 25.—. 


No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 PP. 
D.FI. 100.—. 


ENTOMOLOGISCHE BERICHTEN 


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TRICHOPTERORUM CATALOGUS 
by 
F. C. J. FISCHER 
The Catalogue contains all species of recent and fossil Trichoptera of the world with their 


synonymy and distribution. The complete Catalogue will consist of 15 parts. 
Price of separate parts (in parentheses, copies printed on one side of the pages): 


Vol. 1, 1960, Necrotauliidae, Prosepididontidae, Rhyacophilidae, 168 pp., D.F. 39.— (D.FI. 41.50) 
Vol. 2, 1961, Philopotamidae, Hydroptilidae, Stenopsychidae, 189 pp., D.FI. 39.— (D.Fl. 41.50) 
Vol. 3, 1962, Polycentropodidae, Psychomyiidae, 236 pp, D.Fl. 45. — (D.FI. 47.50) 


Vol. 5, 1964, Phryganeidae, Limnocentropodidae, Molannidae, 214 pp., D.Fl. 45 — (D.Fl. 47.50) 
Vol. 6, 1965, Calamoceratidae, Odontoceridae, Philorheithridae, Leptoceridae I, 242 pp., D.FI. 
[oye (MD Se) 
Vol. 7, 1966, Leptoceridae II, 163 pp., D.Fl. 42.— (D.FI. 45.50) 
Vol. 8, 1967, Goeridae, Limnephilidae, pars 1, 263 pp., D.Fl. 55.— (D.Fl. 59.50) 
Vol. 9, 1968, Limnephilidae, pars 2, 363 pp., D.Fl. 90— (D.Fl. 96.—) 
Vol. 10, 1969, Limnephilidae, pars 3, 332 pp., D.Fl 85.— (D.Fl. 92.—) 
Vol. 11, 1970, Sericostomatidae and other families, 316 pp., D.Fl. 95.— (D.Fl. 102.50) 
Vol. 12, 1971, Supplement to Vols. 1 and 2, 311 pp., D.FI. 98— (D.Fl. 106.—) 
Vol. 13 in press 


1 
2 
3 
Vol. 4, 1963, Hydropsychidae, Arctopsychidae, 225 pp., D.FI. 45.— (D.Fl. 47.50) 
5 
6 


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