22 JUL197
BULLETIN OF V*
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. XXVIII
BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1974
Printed in England by Unwin Brothers Limited
CONTENTS
ENTOMOLOGY VOLUME XXVIII
No. I. The genus Etiella Zeller (Lepidoptera: Pyralidae): a zoogeographic
and taxonomic study. By P. E. S. WHALLEY i
No. 2. G. B. Buckton's works on Aphidoidea (Hemiptera.) By J. P.
DONCASTER 23
No. 3. A catalogue of the genus-group names of the Zygaenidae (Lepidoptera).
By W. G. TREMEWAN m
No. 4. A catalogue of the family-group and genus-group names of the Gele- chiidae, Holcopogonidae, Lecithoceridae and Symmocidae (Lepidop- tera). By K. SATTLER 153
No. 5. The ant genus Polyrhachis F. Smith in the Ethiopian region (Hymen-
optera: Formicidae). By B. BOLTON 283
No. 6. The higher classification of the Lycaenidae (Lepidoptera) : a tentative
arrangement. By J. N. ELIOT 371
No. 7. A revision of the genera Hieroglyphus Krauss, Parahieroglyphus Carl and Hieroglyphodes Uvarov (Orthoptera: Acridoidea). By J. B. MASON 507
Index to Volume XXVIII 561
INDEX TO NOMENCLATURAL CHANGES IN VOLUME XXVIII
abbreviata, Hieroglyphus daganensis . 531 achaja, Cowania ..... 454 Acupicta . . . 433. 451. 48° (fig-) alsulus, Famegana . 454, 492, 499 (figs)
Amblypodiini . . . 432, 480 (fig.) architecta, Polyrhachis laboriosa . . 308 asomaningi, Polyrhachis 298, 360, 362 (figs)
atrociliata, Polyrhachis schistacea . .318
Famegana . . -453
felici, Polyrhachis lanuginosa . . 335
fernandensis, Polyrhachis decemdentata . 302 flavipes, Polyrhachis decemdentata . . 302 flavofasciella, Etiella . . . 5, 14
florinda, Arhopala .... 399
fracta, Polyrhachis schistacea . .318
furella, Nephopterix .... 5 fuscalis, Nephopterix .... 5
balli, Polyrhachis revoili bequaerti, Polyrhachis . bruta, Polyrhachis militaris bubases, Acupicta .
342 gagatoides, Polyrhachis schistacea . .318
304 gallicola, Polyrhachis cubaensis . . 329
313 gerstaeckeri, Polyrhachis . . . 329
454 gustavi, Polyrhachis decementata . . 302
calabarica, Polyrhachis militaris Candalidini
Catapaecilmatini ... Cerodendra . . .
clariseta, Polyrhachis nigriseta Cleoda . . . coerulea, Titea ... conduensis, Polyrhachis revoili congolensis, Polyrhachis gagates conradti, Polyrhachis lanuginosa Cowania .... crassa, Polyrhachis revoili . critala, Arhopala
. . 313 442, 485 (fig.) PI. 4 ... 432 . .122, 125 . . 305 . 123, 125 .. 454 . . 342 . . 305 . . 335 .. 450 . -342 399, 478 (fig-)
cupreopubescens, Polyrhachis militaris . 313
decellei, Polyrhachis . 301, 360, 362 (fig.) delicatum, Acupicta .... 454 divina, Polyrhachis schistacea . . 318
divinoides, Polyrhachis schistacea . . 318 donatana, Pseudotajuria . . . 454
donisthorpei, Polyrhachis revoili . . 338 drososcia, Etiella grisea 4, 13, Pis i, 6, 10, n,
12 durbanensis, Polyrhachis 327, 357, 359 (figs)
hintza, Zintha holozona, Hypogryphia hortulana, Polyrhachis Hypolycaenini Hypotheclini .
454 5
308 438 433
ifraneella, Staudingeria . . 5
imatongica, Polyrhachis cubaensis . 330
indicus, Hieroglyphus . . 536, 537 (fig.) instabilis, Aphis ..... 29 iperpunctata, Polyrhachis . . .310
kohli, Polyrhachis . . . . • 341
laeta, Polyrhachis lauta . . . • 311 latharis, Polyrhachis 348, 361 (fig.), 365 (fig.) lentiginis, Aphis . 29, 61
lestoni, Polyrhachis 349, 361 (fig.). 366 (fig.) limitis, Polyrachis . . . 350, 359 (fig-) localis, Polyrhachis lauta . 311
Loxurini . • -433
Luciini . • 429
Lycaenesthini • 442
lyrifera, Polyrhachis . • 346
edentula, Aphis . . . . 29, 52
elongata, Hieroglyphus banian . . 541
esarata, Polyrhachis . 303, 360, 363 (figs) Euclimaciopsis . . . . .126
maynei, Polyrhachis . • 346
mayumbensis, Polyrhachis rufipalpis . 317
melanella Etiella . 4, J3
Monalita . *3*. *34
myosticta, Cryptoblabes 5
562
Neoherpa
nigriseta, Polyrhachis
Niphandini .
INDEX
129, 134
• 305
442, 499 (fig-). P1- 4
obsidiana, Polyrhachis gagates . . 305
ochristrigella, Etiellia . . . . 5, 18
ochristrigella, Etiellia . . . 5, 18
Ogyrini . 431, 478 (fig-)
Oxylidini . . 433, 480 (fig.)
penicillata, Aphis .... 29, 61, 75 perpolita, Hieroglyphus . . 515, 516 (fig.) perpolita, Miramia . . . .512
plebeia, Polyrhachis schlueteri . .321
Pseudocyrtomyrma .... 288 Pseudotajuria . . . 451, 483 (fig.)
purpurea, Psychonotis
399, 445. 498 (fig.)
regesa, Polyrachis . 337, 361 (fig.), 364 (fig.) Remelanini .... 437, 483 (fig.) rugulosus, Polyrhachis . . . .318 Rysops. . . 452, 490 (fig.), 499 (fig.)
schoutedeni, Polyrhachis . . . 328
scintilla, Rysops ..... 454 scitivittalis, Etiella 4, 10, Pis i, 5, 10, 12, 14
scrophulariae, Siphonophora . . 29, 81
sincerella, Etiella . . . . 4, 10
spretula, Polyrhachis viscosa . . . 330
ssibangensis, Polyrhachis militaris . . 313
striativentris, Polyrhachis militaris . 313
striolatorugosa, Polyrhachis cubaensis . 325 sublutea, Titea ..... 454
Tarakini . . . . . .427
tenuistriata, Polyrhachis decemdentata . 302 Titea . . . . . . 452
Tomarini . . . . . 439
transiens, Polyrhachis 340, 360 (fig.), 365 (fig.)
ugandensis, Polyrhachis fissa . uniformalis, Catastia
304
5
venustella, Phycita .... 5
volkarti, Polyrhachis 341, 356 (fig.), 357 (fig.)
wilrnsi, Polyrhachis cubaensis . . 325
yangtseella, Epischnia .... 5
Zesiini . . . . . . .431
Zintha 447, 453
THE GENUS ETIELLA ZELLER ' 4JUU' (LEPIDOPTERA : PYRALIDAE) : A ZOOGEOGRAPHIC AND TAXONOMIC STUDY .
P. E. S. WHALLEY
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 28 No. i
LONDON : 1973
THE GENUS ETIELLA ZELLER (LEPIDOPTERAj, PYRALIDAE) : A ZOOGEOGRAPHIC AND
TAXONOMIC STUDY 4 ^1973
BY
Pp. 1-2 1 ; 15 Plates, I Text-figure, j Maps
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 28 No. i
LONDON: 1973
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer Papers was instituted, numbered serially for each Department.
This paper is Vol. 28, No. i of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals.
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Issued 16 March, 1973 Price £2.60
THE GENUS ETIELLA ZELLER (LEPIDOPTERA
PYRALIDAE) : A ZOOGEOGRAPHIC AND
TAXONOMIC STUDY
By P. E. S. WHALLEY
CONTENTS
Page
SYNOPSIS ........... 3
INTRODUCTION .......... 3
ACKNOWLEDGEMENTS ......... 4
ABBREVIATIONS .......... 4
CHECK-LIST OF SPECIES OF Etiella ZELLER ...... 4
SPECIES TRANSFERRED FROM Etiella ZELLER ..... 5
GEOGRAPHICAL DISTRIBUTION ........ 5
PHYLOGENY ........... 7
BIOLOGY ........... 7
Etiella ZELLER, 1839 ......... 8
Key to males .......... 9
Key to females ......... 9
TAXONOMIC SECTION ......... 10
REFERENCES ........... 19
INDEX 20
SYNOPSIS
The genus Etiella Zeller (Lepidoptera, Pyralidae) is redefined and the species in it listed and discussed. Keys to these species, together with maps of their distribution, are given. Four specific synonyms are newly established. The possible phylogeny of the genus is discussed and references to the literature on the biology and control of the Lima-bean Pod-borer, Etiella zinckenella (Treitschke) are given.
INTRODUCTION
THE genus Etiella Zeller (Pyralidae, Phycitinae) contains species whose larvae, where known, feed on the seeds of Leguminous plants. The cosmopolitan species, Etiella zinckenella (Treitschke), popularly known as the Lima-bean Pod-borer, is a serious pest of legumes in many parts of the world. Although the present work is primarily a zoogeographic and taxonomic study of the genus, references to the biology and insecticide control of recent years are also given.
The definition of the genus on page 9 restricts the number of species to seven ; no new species is described here and nine species are transferred to other genera. One of the more difficult problems has been the generic placing of the species removed from the genus. In spite of considerable effort, the placing of these species in this work must be regarded as provisional. They may be more accurately placed when other genera of Phycitinae are revised.
4 P. E. S. WHALLEY
The problems of the Phycitinae and their identification were summarized by Heinrich (1956 : vi), who wrote 'So many misidentifications have been made in the past, even by Lepidopterists of repute, that records in the literature cannot be accepted merely on the authority of the author'. This is as true in Etiella as in the other genera in the subfamily and therefore no previously published records of the genus have been accepted unless they were accompanied by clearly recognizable figures. Although we can be reasonably certain that only E. zinckenella has been found on some continents, each record there still requires critical examination.
Most of the type-specimens of the species described in Etiella have been examined ; in cases where these were not available topotypic material was used. The type- specimens of all the presently valid species in the genus have been examined. Wing measurements given are taken from the apex of the fore wing to the centre of the mesothorax. Wing span is thus approximately twice this figure.
ACKNOWLEDGEMENTS
I am grateful to the following for the loan of specimens: Dr I. F. B. Common, C.S.I.R.O., Canberra; Dr G. Friese, Entomological Institute, Berlin; Dr L. Gozmany, Natural History Museum, Budapest ; Dr J. L. Gressitt, Bernice P. Bishop Museum, Hawaii; Dr G. F. Gross, South Australian Museum, Adelaide; Dr. H. J. Hannemann, Zoological Museum, Berlin; Dr H. Inoue, Fujisawa, Japan, who also presented paratype specimens to the BMNH; Dr E. G. Munroe, Canada Dept. of Agriculture, Ottawa; Dr P. Viette, Natural History Museum, Paris.
I am indebted to my colleagues for their comments and advice and to Mr M. Shaffer for technical assistance. The photographs were mostly taken by the Photographic Section, BMNH, the two stereoscans were taken by the Electron Microscope Unit, BMNH, and a few, as indicated, by the author.
ABBREVIATIONS
AMNH American Museum of Natural History, New York.
ANIC Australian National Insect Collection, C.S.I.R.O., Canberra.
BMNH British Museum (Natural History).
BPBM Bernice P. Bishop Museum, Honolulu.
MNHN Museum National d'Histoire Naturelle, Paris.
SAM South Australian Museum, Adelaide.
TM Termeszettudomanyi Miizeum, Budapest.
CHECK-LIST OF SPECIES OF ETIELLA ZELLER
E. scitivittalis (Walker) sp. rev.
sincerella Meyrick syn. n. E. chrysoporella Meyrick E. grisea grisea Hampson E. grisea drososcia Meyrick stat. n. E. hobsoni (Butler)
melanella Hampson syn. n.
THE GENUS ETIELLA 5
E. walsinghamella Ragonot
flavofasciella Inoue syn. n. E. zinckenella (Treitschke)
anticalis Walker *colonnellus Costa
dymmisalis Walker * 'zinckenella ab. decipiens Staudinger
etiella Treitschke
hastiferella Walker
heraldella Guenee
indicatalis Walker *madagascariensis Saalmiiller *majorellus Costa *rubribasella Hulst
sabulinus Butler
schisticolor Zeller *spartiella Rondani *villosella Hulst E. behrii (Zeller)
subaurella Walker
consociella Walker
ochristrigetta Ragonot syn. n.
SPECIES TRANSFERRED FROM ETIELLA ZELLER
The following species, originally described in Etiella, have been transferred to the genera indicated. All their holotypes have been examined.
Assara albicostalis (Walker), transferred by Roesler, 1965, Inaugural Dissertation
der Universitat des Saarlandes, Saarbriicken. Nephopterix furella (Strand) comb. n. Nephopterix fuscalis (Kenrich) comb. n.
Hypogryphia holozona (Lower) comb. n. Very close to H. rufifasciella Hampson. Staudingeria ifraneella (Lucas) comb. n. The holotype is a female and the
generic position of this species is uncertain. Cryptoblabes myosticta (Hampson) comb. n. Catastia uniformalis (Hampson) comb. n. Phycita venustella (Hampson) comb. n. Epischnia yangtseella (Caradja) comb. n.
GEOGRAPHICAL DISTRIBUTION
Maps 1-7 show the distribution of each species. Except for E. zinckenella the species of the genus are restricted to the Australasian and part of the Oriental regions, with one of these species occurring in Japan.
*Holotype or paratype not examined, topotypic material studied.
6 P. E. S. WHALLEY
A small collection from Monte Bello Is. (West Australia), made in 1952, contained a short series of Etiella specimens. These have proved to include three species (behrii, chry sopor ella, grisea). While the former two are widespread Australian species, the latter is otherwise known only from one specimen on the mainland at Wyndham (Western Australia).
E. scitivittalis is restricted to Australia while E. chrysoporella is known only from Australia and the island of Tanimber in the Arafura Sea. E. grisea is widespread over the Pacific and, while not yet recorded from Java and Sumatra, occurs in Ceylon. Although the genitalia of all the specimens of grisea examined were similar, there is some local variation in pattern and colour. In Ceylon the specimens are pale grey while on Tanimber, the only two specimens examined are much blacker than the other specimens. In spite of wide separation of the populations of this species from the Society Islands to the Marianas, no constant differences in morphology have been found between most of these island populations.
E. hobsoni is widely distributed with, at present, few records from New Guinea and none from Celebes. Specimens from Formosa differ only slightly in pattern from the Australian specimens and, on the few specimens examined, cannot be separated subspecifically.
E. walsinghamella has a similar distribution to hobsoni but is less widely distributed in Australia, while extending through the East Indies right up to Japan. Differ- entiation is again slight over the whole range with some pattern differences but, on the material examined, this is not constant. E. walsinghamella is very distinct in external colour and pattern from E. zinckenella but the rest of the morphology and genitalia in both sexes are similar in these two species. It seems probable that walsinghamella and zinckenella are derived only recently from a common ancestor, from which they have only slightly differentiated.
E. zinckenella is pantropical, but in the present work specimens have not been seen from New Zealand or Hawaii, nor from many of the central Pacific Islands. E. zinckenella is widespread in Nearctic, Neotropical, Ethiopian, Oriental and southern Palaearctic regions and in the northern part of Australia and some Pacific Islands, including Samoa. In spite of being widespread, with much variation in size and colour over the whole range, there is no evidence of local populations differentiating on morphological grounds and it seems likely that its spread has been both rapid and relatively recent and probably assisted by man. Another factor with this species is its own inherent ability for widespread dispersion. Stone (1965 : 16) comments that 'the moths are strong fliers and capable of migrating long distances to reach their host-plant.' Certainly this is amongst the most widespread of any species of moth which has not apparently subspeciated over any part of its range.
E. behrii has been recorded in the literature (e.g., Vesey-Fitzgerald, 1941) from outside the range shown in map 7. Most of these specimens have been re-examined and all have proved to be zinckenella. At present the range of behrii is more restricted than zinckenella although it may well prove to have a similar explosive spread-potential and to become more widespread.
THE GENUS ETIELLA 7
PHYLOGENY
No fossil evidence is available for consideration of the evolution of this genus and only biological and morphological evidence is used in the following discussion. The genus consists of two groups of species with different types of distribution
1. World- wide (one species).
2. Mainly Australasian or Oriental (six species).
If the genus is monophyletic there are two ways of considering its phylogeny.
a. The species in the first group, having spread widely from 'a centre of origin', speciated in the Oriental-Australasian region. This is analagous to a wide- spread species arriving on, for example Hawaii, and then radiating and eventually producing many new species. Cases of this type of peripheral or island speciation are well documented (e.g. Zimmerman, 1970).
b. Conversely, the widespread species arose from a species in the Oriental- Australasian region which then spread rapidly round the tropics.
From morphological studies, the world-wide species (zinckendla) shows more specialized features than some of the species of more restricted distribution. The extreme modification of the costa of the valve in the male and the enormous secondary sac on the bursa of the female can be considered the end points arising from species where these characters are present in a less developed condition. For the alternative argument, that the 'simpler' ones arose by reduction of the characters of the wide-spread species, no supporting evidence has been found in any related genus, which seems otherwise morphologically closer to Etiella, wrhere these speciali- zations do not occur. In one species (scitivittalis) the characters of the genus are present in the least developed form and this species is known at present only from Australia. From this species a series showing gradual development of these charac- ters can be drawn from the species in the genus.
If the genus arose by rapid speciation in the Australasian region of the more widespread species by gradual reduction of the various characters, one must assume that a reduced state is more specialized. As already mentioned, no supporting evidence for this has been found in other genera. For example, the long costal process on the valve of the male is unusual and the evidence suggests that the less specialized condition of the valve is the more general (? primitive) form.
In the absence of other evidence, I consider that the genus is Australasian in origin and that one species has been particularly successful, showing explosive spreading throughout the world between 50° north and 50° south.
The genus Etiella is allied to Pima Hulst, whose larvae also feed on leguminous seeds, but its actual relationship to this and other Phycitid genera will have to wait for further studies on them. Within the genus Etiella a possible phyletic relation can be represented by the morphoseries shown in Text-fig, i.
BIOLOGY
E. zinckenella is a pest of pods of legumes. It has been recorded from 30 species in 21 genera of legumes (Naito, 1961) but few records have been published of hosts other than legumes (e.g., Viktorov, 1938, on water melons). Many accounts of the
8 P. E. S. WHALLEY
biology and life-history of this species have been published and a selection of them is given in the references. Data on the other species in the genus is more limited but all the recorded hosts are species of legumes. The species in the genus appear to have specialised in feeding on the seeds of these plants. For further information on the biology and control measures of E. zinckenella, see Issiki, 1969, (coloured figures of the larvae) ; Kruel, 1963 (occurrence in Germany) ; Naito, 1961 (biology and distribution); Oatman, 1967 (biology in the U.S.A.); Peiu, 1966 (biology in Roumania); Schad, 1943 (biology in France); Stone, 1965 (biology and control measures in the U.S.A.).
Details of host plants are given in the section 'Biology' under each species.
ETIELLA Zeller, 1839
Etiella Zeller, 1839 : 733. Type-species: Phycis zinckenella Treitschke, by monotypy. Rhamphodes Guenee, 1845 : 319. Type-species: Phycis etiella Treitschke, by monotypy. Mella Walker, 1859 : 1017. Type-species: Mella dymnusalis Walker, by monotypy. Alata Walker, 1863 : 108. Type-species: Alata anticella Walker, by monotypy. Arucha Walker, 1863 : 201. Type-species: Arucha indicatalis Walker, by monotypy. Modiana Walker, 1863 : 82. Type-species: Modiana scitivittalis Walker, by monotypy. Ceratamma Butler, 1880 : 689. Type-species: Ceratamma hobsoni Butler, by original desig- nation.
The synonymy given in the Catalogue of the genera of Phycitinae (Whalley, 1970 : 45) has been checked. Although the synonymy remains the same, the genus, which has often been attributed to Zeller, 1846, is here referred to his usage of 1839; (in this work Zeller refers to it as a subgenus of Pempelia Hubner, 1825), this is the date used by Heinrich, 1956.
FIG. i. Suggested relationship of species in Etiella Zeller. (See p. 7)
THE GENUS ETIELLA g
Antennae of male with basal segment enlarged with variable shaped projection on inner margin near base. Shaft with sinus containing long scales. Labial palps very long, usually
2 or more times diameter of eye. Second segment of labial palps grooved to hold aigrette-like maxillary palps. Maxilliary palps of female smaller; third segment of labial palp longer than in male. Fore wing often with ridge of raised scales in antemedian position. Eleven fore wing veins. Hind wing with MZ and MS joined. Eighth segment of male with small hair-tufts Uncus hood-like. Gnathus a simple sharp hook. Valve usually with strongly sclerotized costal process. Aedeagus with strongly sclerotized and spiny vesica with cornuti. Female with bursa elongate, usually with many-spined signum, secondary sac (see PI. 10, figs 66, 68 and 76, indicated by V) usually sclerotized, coming off bursa near junction of ductus bursae. Ductus seminalis arising from various positions on bursa, often from near origin of secondary sac.
KEY TO MALES
Juxta with elongate, pointed and sclerotized arms (PI. 5, figs 26 and 30) . . 2
Juxta blunt-ended, usually with apical hairs on each lobe .... 3 One costal valve process long, sclerotized, other costal process reduced but
visible as sclerotized point. (PI. 5, fig. 30) . . . chrysoporella (p. n) Both valves with very reduced processes, visible only as small, lightly sclerotized
processes (PL 5, fig. 25) scitivittalis (p. 10)
3 (i) Both costal valve processes long, sometimes one slightly shorter than other . 4
One valve process very reduced, less than half length of other (PI. 6, fig.
36) .... ...... grisea (p. n)
4 (3) Juxta lobes long and slender (PI. 8, fig. 55). Valve approximately diamond-
shaped (PL 8, fig. 57) behrii (p. 17)
Juxta lobes not as above, often swollen at apex, valve often elongate and thin 5
5 (4) Fore wings reddish brown, without white costal streak. Broad yellow-orange
median fascia. Hind wings black or dark grey. Genitalia as in PL 7, figs 44-
49 ......... walsinghamella (p. 14)
Fore wings with or without white costal streak. If without streak, fore wings black or dark brown ; if with streak, fore wings varying from pale buff to almost black. Antemedian fascia narrow ........ 6
6 (5) White costal streak. Grey-brown, buff or nearly black fore wings. Very
variable in size. Genitalia as in PL 8, fig. 50 . . . zinckenella (p. 15) White costal streak very indistinct or absent. General colour of fore wings black or dark reddish brown. Hind wings dark. Genitalia as in PL 7, fig. 40 ........... hobsoni (p. 13)
KEY TO FEMALES
1 No sclerotized secondary sac on bursa nor sac attached to bursa by duct (PL 10,
fig. 65) scitivittalis (p. 10)
Secondary sac on bursa or attached by long duct. Secondary sac usually
heavily sclerotized ........... 2
2 (i) Duct of bursa short (PL 10, figs 66, 76) usually wider than long ... 3
Duct of bursa much longer than width ....... 4
3 (2) Ductus seminalis from near middle of bursa. Small sclerotized platelets in
broader first part of ductus seminalis. Secondary sac a sclerotized lobe on bursa (PL 10, fig. 66) ...... chrysoporella (p. n)
Ductus seminalis opening nearer bottom of bursa away from ductus bursae. Spines in ductus seminalis. Secondary sac at end of long duct (PL 10, fig. 76) ........... behrii (p. 17)
4 (2) First part of ductus seminalis with small spines or sclerotized plates . . 5
First part of ductus seminalis without spines or sclerotized plates, duct often
strongly folded ........... 6
P. E. S. WHALLEY
(4) Signum a row of long spines, clearly visible (PI. 14, fig. 96) . . hobsoni (p. 13)
Signum of smaller spines, or signum indistinct (PL 14, fig. 95) . grisea (p. 12)
(4) Ductus bursae heavily sclerotized with long striae. Ostium heavily sclerotized.
Secondary sac clearly without sclerotized spines . walsinghamella (p. 14)
Ductus bursae less heavily sclerotized. Ostium lightly sclerotized. Secondary
sac clearly with sclerotized spines ..... zinckenella (p. 15)
TAXONOMIC SECTION
Etiella scitivittalis (Walker) sp. rev. (PI. i, fig. i; PI. 5, figs 25-29; PI. 10, fig. 65; PI. 12, fig. 83; PI. 14, fig. 93)
Modiana scitivittalis Walker, 1863 : 83. Holotype <J, AUSTRALIA (BMNH) [examined]. Etiella sincerella Meyrick, 1879 : 204. Holotype (J, AUSTRALIA (BMNH) [examined]. Syn. n. Etiella sincerella Meyrick; Ragonot, 1893 : 571. [Etiella zinckenella (Treitschke) sensu Ragonot, 1893 : 572, misidentification.]
(J. Wing, 14-16 mm. Labial palps 3 x diameter of eye. Third segment of labial palp less than J length of second. Fore wing, pattern as in PL i, fig. i, reddish brown behind white costal streak. Orange-brown transverse antemedial fascia. Hind wings smoky grey.
Genitalia <$ (PL 5, figs 25-29). Valves with costal margin thickened, process on costa a single short lobe on each valve. Juxta with two sclerotized lateral lobes, curved and pointed. Aedeagus with large cornuti.
$. Wing, 11-15 mm. Third segment of labial palp longer than in male. Pattern and colour similar.
Genitalia $ (PL 10, fig. 65; PL 12, fig. 83; PL 14, fig. 93). Signum a row of long spines. No sclerotized secondary sac on bursa. Ductus seminalis with minute platelets, broad, arising from near middle of bursa.
DISCUSSION. This is the largest species in the genus and is easily recognized by the size and warm reddish brown colour of the fore wings behind the white costal streak. The costal margin of the valve in the male is heavily sclerotized but with only a small process from the margin. The valve also has a small raised papilla half way along the length just below the costa, this papilla is covered with stout spines. An homologous area in the other species also has the stout spines but is without the raised papillae. The juxta is similar to chrysoporella and the aedeagus is typical of the genus. The female differs from all the others in the genus in the lack of the secondary sac on the bursa. From the position where the ductus bursae enters the bursa there is a trace of the first stages of a secondary sac.
BIOLOGY. No information.
DISTRIBUTION. Map i. Australia, Queensland, New South Wales.
MATERIAL EXAMINED.
Holotype <£ (scitivittalis), AUSTRALIA: Moreton Bay, BM slide no. 13056, in BMNH. Holotype $ (sincerella), AUSTRALIA: Sydney, N.S.W., G.H.R., ix.[i8]78, BM slide no. 13254, in BMNH. 18 specimens, SAM, Adelaide. 5 specimens, BMNH. 8 specimens, ANIC, Canberra.
THE GENUS ETIELLA n
Etiella chrysoporella Meyrick (PI. i, figs 2, 4; PI. 5, figs 30, 31; PI. 10, fig. 66; PI. n, fig. 77; PI. 14, fig. 94)
Etiella chrysoporella Meyrick, 1879 : 206. LECTOTYPE <$, AUSTRALIA (BMNH), here desig- nated [examined].
Etiella chrysoporella Meyrick; Ragonot, 1893 : 576.
tj. Wing, 8-12 mm. Fore wing pattern as in PI. i, fig. 2, brown with golden, iridescence.
Spots on antemedial fascia edged with black. White streaks between veins subterminally.
Usually a white streak from basal area below posterior margin of cell. Costal streak white.
Hind wings pale smoky brown.
Genitalia $ (PI. 5, figs 30, 31). Costal process of valve modified, one process shorter than
other. Juxta with strongly sclerotized and pointed lateral arms. Aedeagus with cornuti
similar to zinckenella.
?. Wings, 8-12 mm. Colour and pattern as male. Third segment of labial palps longer
than in male.
Genitalia $ (PL 10, fig. 66; PL n, fig. 77; PL 14, fig. 94). Ductus bursae short, lightly
sclerotized. Secondary sac on bursa large, heavily spined near junction with bursa. Bursa
with row of long spines. Ductus seminalis, with small sclerotized platelets inside, arising from
near middle of bursa.
DISCUSSION. Specimens from Tanimber do not differ from those from the mainland of Australia. The bright iridescent colours and particularly the more numerous white streaks on the fore wing separate this species from the others in the genus.
BIOLOGY. No information.
DISTRIBUTION. Map 2. Australia, Queensland, South Australia, Northern Territory, Western Australia; Indonesia, Tanimber.
MATERIAL EXAMINED.
Lectotype $, AUSTRALIA: Adelaide, S. Australia, I5.x.[i8]82, Meyrick coll. BM slide no. 13208, in BMNH. 40 specimens, BMNH. 35 specimens, SAM, Adelaide. 8 specimens, ANIC, Canberra.
Etiella grisea Hampson
Etiella grisea Hampson, 1903 : 33.
This species varies from grey to a rather blackish brown colour on the fore wings. It is characterized in the male by the reduction of one of the costal valve processes to a short spine. In the female the spines of the signum, which are characteristic of the other species in the genus, are reduced or absent in grisea . The ductus seminalis in the male is broad and the first part of it from the bursa has small sclerotized platelets inside.
E. grisea is widely distributed from Ceylon to Tahiti. The Ceylon specimens are grey in colour whereas those over the rest of the range tend to be more grey-brown. On Tanimber the only two specimens examined have a much darker fore wing than any of the other specimens and could represent a distinct subspecies; the single
12 P. E. S. WHALLEY
specimen from Monte Bello Island (Western Australia) is also slightly different from the others. The series from Ceylon is constant in pattern and has already been named by Hampson, the remaining specimens are grouped with the species Meyrick described from Tahiti. In spite of wide separation of island populations no differences were found in the morphology of the genitalia, even the Ceylon specimens were similar.
KEY TO THE SUBSPECIES OF E. grisea Hampson
Pale grey fore wing, sometimes with transverse fascia. Ceylon . grisea grisea (p. 12)
Grey or grey-brown fore wing, sometimes with transverse fascia, occasionally with traces
of white costal streak. Australian-Pacific .... grisea drososcia (p. 13)
Etiella grisea grisea Hampson (PI. i, fig. 3; PI. 6, figs 32-35; PI. 10, fig. 67; PI. 14, fig. 95)
Etiella grisea Hampson, 1903 : 33. LECTOTYPE <$, CEYLON (BMNH), here designated
[examined] .
cj. Wing, 10-12 mm. Labial palps 3 x diameter of eye, third segment one quarter length of second. Fore wing pattern as in PL i, fig. 3, pale grey with darker, transverse, antemedial fascia.
Genitalia <$ (PI. 6, figs 32-35). Juxta with elongate lateral lobes slightly clavate at apex. One valve with very short costal process, other valve with long costal process. Aedeagus with spiny, sclerotized vesica with two large sclerotized patches.
$. Wing, 8-5-10 mm. Labial palps with third segment half length of second. Fore wing with slightly less distinct pattern than male.
Genitalia $ (PI. 10, fig. 67; PL 14, fig. 95). Duct of bursa sclerotized, long and strongly ribbed. Sclerotized signum with small spines, small patch of spines on secondary sac of bursa. Ductus seminalis arising from nearer centre of bursa, enlarged at origin with bursa and with small platelets inside.
DISCUSSION. This subspecies is known only from Ceylon. The morphology of the genitalia is similar to grisea drososcia and this latter subspecies has some specimens almost as pale as those from Ceylon, but generally the specimens from Ceylon are separable from the other by their paler grey colour.
BIOLOGY. No information. DISTRIBUTION. Map 3. Ceylon.
MATERIAL EXAMINED.
Lectotype <$, CEYLON, 95.91, BM slide no. 13294, in BMNH. 12 specimens, BMNH.
THE GENUS ETIELLA 13
Etiella grisea drososcia Meyrick stat. n.
(PI. i, figs 5, 6; PI. 6, figs 36-39; PI. 10, fig 68; PI. n, fig. 80; PL 12, figs 85, 87)
Etiella drososcia Meyrick, 1929 : 158. LECTOTYPE <J, TAHITI (BMNH), [examined]. [Etiella zinckenella (Treitschke) sensu Tarns, 1935 : 254, misidentification.]
o*. Wing, 9-14 mm. Head as nominate subspecies. Fore wing, pattern as in PI. i, figs 5, 6, grey or grey-brown with light coloured costal streak. Wings variably marked, often with black spots on indistinct antemedial fascia.
Genitalia $ (PI. 6, figs 36-39). As nominate subspecies.
?. Wing, 11-12-5 rnm. Pattern and colour as male. Third segment of labial palp half length of second.
Genitalia $ (PL 10, fig. 68; PL n, fig. 80; PL 12, figs 85, 87). As nominate subspecies.
DISCUSSION. The two specimens from Tanimber are much blacker than the others and the single male from Prince of Wales Is. (N. Australia) is slightly larger than the other specimens. The only specimen from the mainland of Australia is a female from Wyndham (W. Australia). The pattern and the distinctness of the median fascia are equally variable over the whole of the range of this subspecies. It is probable that this species is widespread over most Pacific islands; so far no specimens have been seen from Java or Sumatra. It is interesting to speculate whether, since this subspecies is known from pods of legumes, it enters into compe- tition with the widespread zinckenella. Will this latter species eventually spread to all the islands and replace grisea or will grisea become commoner? In Ceylon and other places the two species occur together, but there is no information on the detailed ecology of grisea.
BIOLOGY. This subspecies has been bred from the pods of Vigna (Leguminosae) in Fiji and from Crotalaria pods (Leguminosae) in Guam.
DISTRIBUTION. Map 3. Society Is., Tahiti; Cook Is., Rarotonga; Samoa; Fiji; New Hebrides; Solomon Is., St. Cristobal; Australia, Prince of Wales Is., Monte Bello Is., West Australia; Indonesia, Tanimber; New Guinea; Caroline Is., Truk; Mariana Is., Guam.
MATERIAL EXAMINED.
Lectotype c£ : TAHITI: nr. Papeete, iii-iv. 1925 (Cheesman), BM slide no. 13245, in BMNH. 12 specimens, BMNH. 30 specimens, BPBM, Honolulu. 2 specimens, ANIC, Canberra.
Etiella hobsoni (Butler)
(PI. 2, figs 7-9; PI. 7, figs 40-43; PI. 10, figs 69-71; PI. ii, fig. 78; PI. 13, figs 88, 89; PI. 14, figs 96-98)
Ceratamma hobsoni Butler, 1880 : 689. LECTOTYPE <J, FORMOSA (BMNH), here designated
[examined] .
Etiella hobsoni Butler; Ragonot, 1893 : 578. Etiella hobsoni Butler; Shibuya, 1928 : 94. Etiella melanella Hampson & Ragonot, 1901 : 558, Holotypeo*, AUSTRALIA (BMNH), [examined].
Syn. n. Etiella hopsoni, misspelling, Caradja, 1939 : 20.
I4 P. E. S. WHALLEY
<J. Wing, 7-9 mm. Labial palps z\ x diameter of eye, third segment one third length of second. Fore wing, pattern as in PL 2, figs 7-9, grey -black with brown or black transverse fascia. Hind wings smoky grey.
Genitalia $ (PL 7, figs 40-43). Costal process of valve on one side ij x length of other side Juxta lightly sclerotized, apex swollen and hairy. Aedeagus with two large cornuti.
$. Wing, 7-10 mm. Colour and pattern as male.
Genitalia $ (PL 10, figs 69-71 ; PL u, fig. 78; PL 13, figs 88, 89; PL 14, figs 96-98). Ductus bursae heavily sclerotized with prominent longitudinal ribbing. Ductus seminalis with small platelets with very broad opening near centre of bursa.
DISCUSSION. This small species is similar externally to walsinghamella but lacks the prominent yellow-orange fascia of that species and has black, not reddish fore wings. The main variation between specimens of hobsoni is in the colour, or the presence of, the transverse fascia of the fore wing. Some specimens are without this fascia, others have it in a very incomplete form. There is little other mor- phological variation between specimens from as far apart as Formosa and Australia.
BIOLOGY. No information.
DISTRIBUTION. Map 4. Australia, Queensland, Northern Territory, South Australia; New Guinea; Solomon Is.; New Britain; Caroline Is., Truk; Indonesia, Wetar, Timor; Formosa.
MATERIAL EXAMINED.
Lectotype $ (hobsoni), FORMOSA, 80-115, BM slide no. 13247, in BMNH. Holo- type c? (melanella), AUSTRALIA: Adelaide, vii. [i8]gi, BM slide no. 13270, in BMNH. 25 specimens, BMNH. i specimen, SAM, Adelaide. 32 specimens, BPBM, Honolulu.
Etiella walsinghamella Ragonot
(PI. 2, figs 10-12; PI. 7, figs 46-49; PI. 10, figs 72-74; PI. 13, figs 90-92;
PI. 15, figs 99-101)
Etiella walsinghamella Ragonot, 1888 : 27. Holotype $, NEW GUINEA (BMNH) [examined].
Etiella walsinghamella Ragonot; Ragonot, 1893 : 577.
Etiella flavofasciella Inoue, 1959 : 299, Holotype <$, JAPAN (Kyushu University Coll., Japan).
[not examined]. Syn. n.
o*. Wing, 8-12 mm. Labial palps 3^ x diameter of eye, third segment less than one quarter of second. Fore wing, pattern as in PL 2, figs 10-12, brown or reddish brown with yellow- orange fascia. Hind wings smoky grey.
Genitalia $ (PL 7, figs 46-49). Similar to zinckenella. Valve with slightly less slender apex. Aedeagus with cornuti more slender than zinckenella.
$. Wing, 8-1 1 mm. Pattern as male but with fore wings usually with more red scales and hind wings darker, almost black. Narrow median fascia usually present on fore wing.
Genitalia $ (PL 10, figs 72-74; PL 13, figs 90-92; PL 15, figs 99-101). Similar to zinckenella, signum longer in proportion to total length of bursa and ductus bursae usually longer in walsinghamella than zinckenella.
DISCUSSION. Externally the pattern and general appearance of this species separate it from zinckenella but the genitalia of these two species are similar. The
THE GENUS ETIELLA 15
male of walsinghamella has shorter costal processes on the valve and the juxta lobes are blunter at the apex than zinckenella. In the females, the ductus bursae and signum are longer in walsinghamella than zinckenella (not clear in figs 74, 75 due to slight differences in magnification) . E. walsinghamella is closely related to zinckenella and has only diverged slightly from it. No overlap in pattern between these two species has been found but some of the specimens of zinckenella from New Guinea approach walsinghamella in general colour. Within the material of walsinghamella examined, the Australian and Japanese specimens are larger than those from New Guinea, with more red in the fore wing but there is an overlap in these characters and no clear subspecific trend on external characters is shown.
BIOLOGY. No information.
DISTRIBUTION. Map 5. Australia, Queensland; New Guinea; Japan.
MATERIAL EXAMINED.
Holotype <£ (walsinghamella}, NEW GUINEA (specimen lacks abdomen), in BMNH. Paratype $ and $ (flavofasciella) , JAPAN: Orio, Fukuoka Pref., 28.viii.i958 (Kawa- wwra),inBMNH. 8 specimens, BMNH. 8 specimens, SAM, Adelaide, i specimen, BPBM, Honolulu.
Etiella zinckenella (Treitschke)
(PI. 3, figs 13-18; PI. 4, figs 23, 24; PI. 8, figs 50-54, 56; PI. 9, fig. 64; PI. 10, fig. 75; PL n, fig. 79; PI. 15, fig. 102)
Phycis zinckenella Treitschke, 1832 : 201. Lectotype $, SICILY (TM) [examined].
Phycis etiella Treitschke, 1835 : 174 [unnecessary replacement name].
Pempelia Etiella zinckenella (Treitschke); Zeller, 1839 : 179.
Rhamphodes zinckenella (Treitschke); Guenee, 1845 : 319.
Rhamphodes etiella (Treitschke); Guenee, 1846 : 81.
Etiella zinckenella (Treitschke); Zeller, 1846 : 733.
Etiella zinckenella (Treitschke); Heinemann, 1865 : 154.
Chilo colonnellus Costa, [1836] : [243]. Type, ITALY [type-series not traced].
Chilo majorellus Costa, [1836] : [241]. Holotype <$, ITALY [type-series not traced].
Mella dymnusalis Walker, 1859 : 1018. Holotype $, SIERRA LEONE (BMNH) [examined].
Rhamphodes heraldella Guene'e, 1862 : G-72. Holotype $, REUNION (MNHN) [not examined].
Alata anticalis Walker, 1863 : 108. LECTOTYPE $, CHILE (BMNH) here designated
[examined].
Arucha indicatalis Walker, 1863 : 202. Holotype <$, SOUTH AFRICA (BMNH) [examined]. Alata hastiferella Walker, 1866 : 1725. Holotype <$, GRENADA (BMNH) [examined]. Etiella zinckenella ab. decipiens Staudinger, 1870 : 195. Type? EUROPE, [type-series not
traced].
Etiella spartiella Rondani, 1876 : 19 Holotype <$ [ITALY?], [not examined]. Crambus sabulinus Butler, 1879 : 455. Holotype c?, JAPAN (BMNH), [examined]. Etiella madagascariensis Saalmiiller, 1879-80 : 307. Type, MADAGASCAR [type-series not
traced].
Etiella zinckenella (Treitschke); Meyrick, 1879 : 203. Etiella schisticolor Zeller, 1881 : 178. LECTOTYPE $, U.S.A. (BMNH), here designated
[examined].
16 P. E. S. WHALLEY
Etiella zinckenella (Treitschke) ; Meyrick, 1883 : 156.
Etiella villosella Hulst, 1887 : 133. Holotype <?, U.S.A. (AMNH) [not examined].
Etiella zinckenella (Treitschke); Hulst, 1890 : 169.
Etiella rubribasella Hulst, 1890: 170. Holotype <?, U.S.A., (AMNH) [not examined].
Etiella zinckenella (Treitschke); Ragonot, 1893 : 572.
Etiella zinckenella (Treitschke); Hampson, 1896 : 108.
{Etiella scitivittalis (Walker) sensu Hampson, 1896 : 108, misidentification.]
Etiella zinckenella (Treitschke); Oberthiir, 1922 : 333.
Etiella zinckenella (Treitschke) ; Shibuya, 1928 : 93, partim.
Etiella zinckenella (Treitschke); Janse, 1944 : I5-
Etiella zinckenella (Treitschke) ; Heinrich, 1956 : 99.
Etiella zinckenella (Treitschke) ; Commonwealth Institute of Entomology, 1959, Map 105.
Etiella zinckenella (Treitschke) ; Whalley, 1970 : 45.
o*. Wing, 10-15 mm. Labial palps 3 x diameter of eye. Fore wing, pattern as in PI. 3, figs 13-18, variably coloured, red-brown to black-brown. White costal streak usually present. Antemedial fascia orange-brown to orange-red, fascia edged with black on antemedial side, frequently gold iridescence on fascia.
Genitalia $ (PL 8, figs 50-54, 56; PL 9, fig. 64). Costal processes on valves long. Juxta with two broad, elongate, lateral lobes, rounded at apex. Aedeagus with large sclerotized cornuti in vesica and many small spines.
$. Wing, 6-15 mm. Third segment of labial palp longer than male. Colour and pattern similar.
Genitalia $ (PL 10, fig. 75; PL n, fig. 79; PL 15, fig. 102). Ductus bursae long, sclerotized, tapering slightly towards opening with bursa. Secondary sac large, sclerotized, containing spines. Large spiny signum on main bursal sac. Ductus seminalis, without platelets inside, arising near origin of ductus bursae and bursa itself.
DISCUSSION. This species is common in most tropical and many temperate countries, where it is a pest of legumes. Specimens of zinckenella vary considerably in size and, to a lesser extent, in colour, although the pattern is fairly constant. There is a tendency for specimens from North America to be greyer than those from the rest of the world but this is by no means constant. The only other small difference between Old and New World specimens is in the length of the ductus bursae of the female ; this is slightly shorter in the American specimens than in the others, but again this is not constant, some overlap occurring. In Morocco some large dark specimens were included in a series of more normally coloured ones, although no other differences could be found in this series and the genitalia were typical of the species. A series of specimens from Egypt were smaller and more sandy coloured than the more typical red or black-brown. From the rest of the African continent there is a wide range of size and colour variants with series of particularly small specimens from Madagascar. Specimens from the Indian sub- continent, Malaya, China, and Japan were similar to those from the Mediterranean region. In Indonesia some variation was found in wing colour but in New Guinea a distinct form occurs with dark hind wings, instead of the more typical pale smoky brown ones. Over the rest of the range a few specimens were found with this darker hind wing but the New Guinea specimens and some from the Solomon Islands were very dark. No other differences have been found between these specimens of zinckenella, no evidence of seasonal or food-plant forms were found but the data was mostly too fragmentary to examine this aspect. In most specimens the genitalia
THE GENUS ETIELLA 17
were similar, little intra-specific variation was found although certain structures varied slightly. These were the length of the costal processes of the males, the shape and extent of the spines in the vesica and in the females in the extent of the signum. Variation in all these characters was found equally in specimens from all parts of the range of zinckenella.
BIOLOGY. Pest of seeds of legumes throughout the world, recorded from 30 species of 21 genera (Naito, 1961). Detailed accounts of the biology and larvae will be found in Naito (1961), Oatman (1967) and Stone (1968). The eggs are laid in the seed-pod of the legume and the larvae bore into the pods, feeding on the seeds. Figures for the length of life-cycle vary but Stone (1968) gives from 2 months to 9-9 months at 6i°F. The larvae, after feeding on the seeds, bite their way out and pupate in the soil, forming a small cocoon of soil.
Specimens have been examined from all the countries in the following list. Although I have not seen the Costa types (majorellus, colonnellus) I have examined topotypic material and I agree with Zeller (1846 : 751) that these two are just re- descriptions of zinckenella.
DISTRIBUTION. Map 6. All Mediterranean countries and Black Sea area; Southern Germany, Austria. The whole African continent, north and south of Sahara, Tenerife, Sao Thome, Principe, Madagascar, Aldabra, Comoro Is., Seychelles. All countries in the Oriental region; China, Japan, Philippines, Formosa; Hainan, New Guinea, Indonesia, New Caledonia, Solomon Is., Samoa. In Australia from Queensland, Prince of Wales Is. From both North and South America and Canada, Galapagos and most West Indian Islands, Cuba, Puerto Rico.
It has not been recorded from U.K., Northern Europe, Hawaii, New Zealand, and many Pacific Islands.
MATERIAL EXAMINED.
Lectotype $ (zinckenella), SICILY, Treits. 3456, BM slide no. 12121, in TM, Buda- pest. Holotype $ (dymnusalis) , SIERRA LEONE, in BMNH. Lectotype,^, (anticalis) ST. JAGO [Santiago] (Darwin), in BMNH. Holotype $ (indicatalis) , S. AFRICA, in BMNH. Lectotype c? (schisticolor) , CALIFORN., E. schisticolor (Grote), 97, in BMNH. Holotype £ (sabtMnus), JAPAN, in BMNH. Holotype £ (hastiferella] , GRENADA, in BMNH. Over 1000 specimens, mostly BMNH. 25 specimens, BPBM, Honolulu.
Etiella behrii (Zeller)
(PI. 4, figs 19-22; PI. 8, figs 55, 57; PI. 9, figs 58-63; PI. 10, fig. 76; PI. n, figs 81, 82; PI. 12, fig. 84; PI. 15, fig. 103)
Phycis behrii Zeller, 18485 : 883. LECTOTYPE ?. AUSTRALIA (BMNH), here designated
[examined].
Alata consociella Walker, 1866 : 1724. Holotype $, AUSTRALIA (BMNH) [examined]. Alata subaurella Walker, 1866 : 1724. LECTOTYPE £, AUSTRALIA (BMNH), here designated
[examined] .
i8 P. E. S. WHALLEY
Etiella ochristrigella Ragonot, 1888 : 27, LECTOTYPE ?, NEW GUINEA (BMNH), here designated
[examined]. Syn. n.
{Etiella zinckenella sensu auct., nee Zeller, misidentifications.]
Etiella zinckenella (Treitschke) sensu Shibuya, 1928 : 93, partim, misidentification.] {Etiella behrii (Zeller) sensu Evans, 1952 : 181, misidentification.]
o\ Wing, 10-13 mm. Externally similar to zinckenella but often greyer and always with more black spots on the wings. Fore wing, pattern as in PI. 4, figs 19-22.
Genitalia <$ (PI. 8, figs 55, 57; PL 9, figs 58-63). Differing from zinckenella in shape of valve, one costal process noticeably shorter than other (but not as short as in chrysoporella) , juxta with long, slender, rather parallel-sided lobes. Aedeagus with more slender and smaller cornuti than zinckenella.
$. Wing 8-5-12 mm. Similar to female zinckenella, more black spots on fore wing but other- wise difficult to separate externally.
Genitalia $ (PI. 10, fig. 76; PI. u, figs 81, 82; PI. 12, fig. 84; PI. 15, fig. 103). Differing from all other species in the genus in having the secondary sac at the end of a long duct. Neck of ductus bursae short. Ductus seminalis with small spines in first part, opening near base of bursal sac (i.e. away from ductus bursae).
DISCUSSION. This species has long been confused with zinckenella and the two cannot be reliably separated on external characters. In the male behrii, the process on the base of the antennal segment is larger than in zinckenella but both species can be easily separated on genitalic characters. E. behrii is wide spread in Australia and probably occurs in most of Indonesia although I have seen specimens from only a few of the islands. The Malayan and Formosan specimens are slightly darker than the Australian ones but the genitalia are similar. As with all species in the genus the darker colour tends to fade with the increasing age of the specimen. At present the species is known from the mainland of China only from a few specimens from Hong Kong. It will probably prove to be more widespread. Less variation was found in size of specimens of behrii than in zinckenella.
BIOLOGY. E. behrii has been bred from pods of Lucerne and Ground-nuts (both Leguminosae).
DISTRIBUTION. Map 7. Australia, all states; New Guinea; New Hebrides; Malaya; Indonesia, Borneo, Tanimber; Formosa; Hong Kong.
MATERIAL EXAMINED.
Lectotype $ (behrii), AUSTRALIA: Adelaide, BM slide no. 11163, in BMNH. Lectotype $ (ochristrigella), NEW GUINEA: Port Moresby, x. 1887-1. 1888 (Kowald), BM slide no. 13248, in BMNH. Lectotype $, (subaurella) , AUSTRALIA: Sydney, BM slide no. 13240, in BMNH. Holotype $ (consociella) , AUSTRALIA: Moreton Bay, in BMNH. 72 specimens, BMNH. 42 specimens, ANIC, Canberra. 37 specimens, SAM, Adelaide. 25 specimens, BPBM, Honolulu.
THE GENUS ETIELLA ig
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1883. Descriptions of Australian Microlepidoptera. Proc. Linn. Soc. N.S.W. 7 : 148-202.
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sumura. IV. On the distribution of the two species in the world and their host plants.
Kontyu 29 : 39-55.
OBERTHUR, C. 1922. Etudes de Lepidopterologie Comparee, 19 : 402 pp. Rennes. OATMAN, E. R. 1967. An ecological study of the lima-bean borer (Etiella zinckenella Tr.) in
Southern California. Ann. ent. Soc. Am. 60 : 552-555. PEIU, M. 1966. Contributi la studiul biologiei si combateri fluturelui pastailor de Soia
(Etiella zinckenella Tr.) in regiunea lasi. Anal. Inst. cent. Cere. sect. Agric. PL Prot. 3
(1965) : 205-220.
20 P. E. S. WHALLEY
RAGONOT, E. L. 1888. Nouveaux genres et especes de Phycitidae et Galleriidae. 52 pp. Paris.
1893. In Romanoff, N. M. M6moires sur les L6pidopteres. 7 : 658 pp. St. Petersburg.
- 1901. See Hampson and Ragonot.
RONDANI, C. 1876. Papilionaria aliqua Microsoma nuper observata a Prof. Camillo Rondani.
Boll. Soc. ent. Ital. 8 : 19-24. SAALMULLER, M. 1879-80. Neue Lepidopteren aus Madagaskar. Ber. senckerib. naturf.
Ges. 1879-1880 : 258-310. SCHAD, C., & GUIGNAUD, P. 1943- La Pyrale des haricots (Etiella zinckenella Treitschke)
parasite du soja dans le sud-ouest de la France. Annls Epiphyt. (N.S.) 9 : 165-175. SHIBUYA, J. 1928. The systematic study on the Formosan Pyralidae. /. Fac. Agric.
Hokkaido Imp. Univ. 22 : 1—300. STAUDINGER, O. 1870 (1871). Beschreibung neuer Lepidopteren des europaischen Faunen-
gebiets. Berl. ent. Z. 14 : 97-208. STONE, M. W. 1965. Biology and control of the lima-bean pod-borer in Southern California.
Tech. Bull. U.S. Dep. Agric. no. 1321 : 1-46. STRAND, E. 1918. H. Sauter's Formosa- Ausbeute : Pyralididae. Stettin, ent. Ztg 79 : 248-
276.
TAMS, W. H. T. 1935. Heterocera. Insects Samoa 3 (4) : 169-289. TREITSCHKE, F. 1832. Die Schmetterlinge von Europa, 9 (i) : 1-262. Leipzig.
- 1835. Die Schmetterlinge von Europa, 10 (2) : 1-302. Leipzig. VESEY-FITZGERALD, D. 1941. Some Insects of Economic importance in Seychelles. Bull.
ent. Res. 32 : 153-160. VIKTOROV, V. F. 1938. Etiella zinckenella Tr., as a pest of Water melons. Zashch. Rast.
16 : 108-110. WALKER, F. 1859. List of the Specimens of Lepidopterous Insects in the collection of the British
Museum. 19 : 799-1036. London.
1863. List of the Specimens of Lepidopterous Insects in the collection of the British Museum.
27 : 1-286. London.
1866. List of the Specimens of Lepidopterous Insects in the collection of the British Museum.
35 : 1536-2040. London.
WHALLEY, P. E. S. 1970. A synonymic catalogue of the genera of Phycitinae of the world.
Bull. BY. Mus. nat. Hist. (Ent). 25 : 33-72. WOLCOTT, G. N. 1936. 'Insectae Borinquenses', a Revision of 'Insectae Portoricensis'.
/. Agric. Univ. P. Rico 20 : 1-600. ZELLER, P. C. 1839. Versuch einer naturgemassen Eintheilung der Schaben. Isis, Leipzig
167-219.
- 1846. Die knotenhornigen Phyciden. Isis, Leipzig: 729-787.
- i848a. Die Gallerien und nackthornigen Phyciden. Isis, Leipzig: 569-618; 641-691;
721-754-
i848b. Exotische Phyciden. Isis, Leipzig: 857-890.
1881. Columbische Chiloniden, Crambiden und Phyciden. HoraeSoc. ent. Ross. 16 : 154-
256.
ZIMMERMAN, E. C. 1970. Adaptive Radiation in Hawaii with special reference to Insects. Biotropica 2 : 32-38.
INDEX
Alata Walker, 8 Ceratamma Butler, 8
albicostalis Walker, 5 chrysoporella Meyrick, 4, 6, 8, 9, 11
anticalis Walker, 5, 15 colonnellus Costa, 5, 15, 17
Arucha Walker, 8 consociella Walker, 5, 17
Assara Walker, 5 Cryptoblabes Zeller, 5
behrii Zeller, 5, 6, 8, 9, 17 decipiens Staudinger, 5, 15
Catastia Htibner, 5 drososcia Meyrick, 4, 13
THE GENUS ETIELLA
dymnusalis Walker, 5, 15
Epischnia Hiibner, 5
etiella Treitschke, 15
Etiella Zeller, 7, 8
flavofasciella Inoue, 5, 14
furella Strand, 5
fuscalis Kenrick, 5
grisea drososcia Meyrick, 4, 12, 13
grisea grisea Hampson, 4, 6, 8, 9, 10, n, 12
hastiferellus Walker, 5, 15
heraldella Guen6e, 5, 15
hobsoni Butler, 4, 6, 8, 9, 10, 13
holozona Lower, 5
Hypogryphia Ragonot, 5
ifraneella Lucas, 5
indicatalis Walker, 5, 15
madagascariensis Saamiiller, 5, 15
majorellus Costa, 5, 15, 17
melanella Hampson, 4, 13
Mella Walker, 8
Modiana Walker, 8
myosticta Hampson, 5
Nephopterix Hiibner, 5
ochristrigella Ragonot, 5, 18
Pempelia Hiibner, 8
Pima Hulst, 7
Phycita Curtis, 5
Rhamphodes Guen6e, 8
rubribasella Hulst, 5, 16
sabulinus Butler, 5, 15
schisticolor Zeller, 5, 15
scitivittalis Walker, 4, 7, 8, 9, 10
sincerella Meyrick, 4, 10
spartiella Rondani, 5, 15
Staudingeria Ragonot, 5
subaurella Walker, 5, 17
uniformalis Hampson, 5
venustella Hampson, 5
vilosella Hulst, 5, 16
walsinghamella Ragonot, 5, 6, 8, 9, 10, 14
yangtseella Caradja, 5
zinckenella Treitschke, 5, 6, 7, 8, 9, 10, 15, 18
P. E. S. WHALLEY, M.Sc.
Department of Entomology
BRITISH MUSEUM (NATURAL HISTORY)
CROMWELL ROAD
LONDON, SW7 5BD
scitivittalis
MAP i
chrysoporella
MAP 2
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. grisea
MAP 3
MAP 4
I 1 1 1 I I I I
MAP 5
behrii
MAP 7
PLATE i
FIG. i. Etiella scitivittalis Walker, Australia.
FIG. 2. E. chrysoporella Meyrick, Tanimber.
FIG. 3. E. grisea grisea Hampson, Ceylon.
FIG. 4. E. chrysoporella Meyrick, Australia.
FIG. 5. E. grisea drososcia Meyrick, Tanimber.
FIG. 6. E. grisea drososcia Meyrick, Tahiti.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE r
PLATE 2
FIG. 7. Etiella hobsoni Butler, Formosa.
FIG. 8. E. hobsoni Butler, Formosa.
FIG. 9. E. hobsoni Butler, Australia.
FIG. 10. E. walsinghamella Ragonot, Papua.
FIG. ii. E. walsinghamella Ragonot, Papua.
FIG. 12. E. walsinghamella Ragonot, Australia.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 2
8
10
PLATE 3
FIG. 13. Etiella zinckenella Treitschke, Europe.
FIG. 14. E. zinckenella Treitschke, Europe.
FIG. 15. E. zinckenella Treitschke, Madagascar.
FIG. 16. E. zinckenella Treitschke, Madagascar.
FIG. 17. E. zinckenella Treitschke, Papua.
FIG. 1 8. E. zinckenella Treitschke, U.S.A.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 3
• 3mm — I
14
| — 3mm — |
15
16
17
I— 3mm H
18
I — 3mm — I
PLATE 4
FIG. 19. Etiella behrii Zeller, Australia, holotype. FIG. 20. E. behrii Zeller, Australia. FIG. 21. E. behrii Zeller, Australia. FIG. 22. E. behrii Zeller, Malaysia.
FIG. 23. E. zinckenella Treitschke, head dorsal view, showing chaetosema (C). FIG. 24. E. zinckenella Treitschke, head, ventral view, showing pilifers (P) and reduced mandibles (M).
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 4
19
I — 3mm— |
I — 3mm — |
21
| — 3mm— |
22
PLATE 5
FIG. 25. Etiella scitivittalis Walker, Australia.
FIG. 26. E. scitivittalis Walker, details of juxta.
FIG. 27. E. scitivittalis Walker, aedeagus.
FIG. 28. E. scitivittalis Walker, apex of cornutus in aedeagus. Photo P. Whalley.
FIG. 29. E. scitivittalis Walker, cornutus, magnified to show ribbing. Photo P. Whalley.
FIG. 30. E. chrysoporella Meyrick, Australia.
FIG. 31. E. chrysoporella Meyrick, aedeagus.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 5
27
31
PLATE 6
FIG. 32. Etiella grisea grisea Hampson, Ceylon. FIG. 33. E. grisea grisea Hampson, valve. Photo P. Whalley. FIG. 34. E. grisea grisea Hampson, aedeagus. Photo P. Whalley. FIG. 35. E. grisea grisea Hampson, enlargement of apex of aedeagus showing vesica. Photo
P. Whalley.
FIG. 36. E. grisea drososcia Meyrick, Tahiti. FIG. 37. E. grisea drososcia Meyrick, Tahiti, valve. FIG. 38. E. grisea drososcia Meyrick, Tanimber. FIG. 39. E. grisea drososcia Meyrick, Fiji, valve.
Bull. Br. Mm. nat. Hist. (Ent.) 28, i
PLATE 6
36
38
PLATE 7
FIG. 40. Etiella hobsoni Butler, Formosa, holotype.
FIG. 41. E. hobsoni Butler, Formosa, aedeagus.
FIG. 42. E. hobsoni Butler, Australia.
FIG. 43. E. hobsoni Butler, Australia, cornuti in aedeagus.
FIG. 44. E. walsinghamella Ragonot, Papua.
FIG. 45. E walsinghamella Ragonot, aedeagus.
FIG. 46. E. walsinghamella Ragonot, Japan (paratype of flavofasciella Inoue).
FIG. 47. E. walsinghamella Ragonot, Japan, juxta. Photo P. Whalley.
FIG. 48. E. walsinghamella Ragonot, Japan, aedeagus.
FIG. 49. E. walsinghamella Ragonot, enlargement of vesica.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 7
PLATE 8
FIG. 50. Etiella zinckenella Treitschke, Europe
FIG. 51. E. zinckenella Treitschke, apex of aedeagus, showing vesica.
FIG. 52. E. zinckenella Treitschke, aedeagus.
FIG. 53. E. zinckenella Treitschke, apical cornutus in aedeagus.
FIG. 54. E. zinckenella Treitschke, juxta. Photo P. Whalley.
FIG. 55. E. behrii Zeller, juxta.
FIG. 56. E. zinckenella Treitschke, valve.
FIG. 57. E. behrii Zeller, valve. Photo P. Whalley.
Bull. BY. Mus. nat. Hist. (Ent.) 28, i
PLATE 8
56
57
X
PLATE 9
FIG. 58. Etiella behrii Zeller, Australia, (holotype subaurella Walker)
FIG. 59. E. behrii Zeller, aedeagus.
FIG. 60. E. behrii Zeller, cornuti in aedeagus.
FIG. 61. E. behrii Zeller, Formosa.
FIG. 62. E. behrii Zeller, aedeagus.
FIG. 63. E. behrii Zeller, cornuti in aedeagus.
FIG. 64. E. zinckenella Treitschke, cornuti in aedeagus.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 9
A t
PLATE 10
Female genitalia. (Arrows indicate ductus seminalis).
FIG. 65. Etiella scitivittalis Walker, Australia.
FIG. 66. E. chrysoporella Meyrick, Australia, secondary sac (S).
FIG. 67. E. grisea grisea Hampson, Ceylon.
FIG. 68. E. grisea drososcia Meyrick, Tahiti, secondary sac (S).
FIG. 69. E. hobsoni Butler, Formosa.
FIG. 70. E. hobsoni Butler, Australia.
FIG. 71. E. hobsoni Butler, Solomons.
FIG. 72. E. walsinghamella Ragonot, Japan.
FIG. 73. E. walsinghamella Ragonot, Papua.
FIG. 74. E. walsinghamella Ragonot, Australia.
FIG. 75. E. zinckenella Treitschke, Europe.
FIG. 76. E. behrii Zeller, Australia, lectotype, secondary sac (S).
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 10
65
PLATE ii
FIG. 77. Etiella chrysoporella Meyrick, ostium and bursa.
FIG. 78. E. hobsoni Butler, ostium and bursa.
FIG. 79. E. zinckenella Treitschke, ostium.
FIG. 80. E. grisea drososcia Meyrick, ductus seminalis on bursa.
FIG. 81. E. behrii Zeller, Borneo, ostium, bursa and duct of secondary sac.
FIG. 82. E. behrii Zeller, Australia, lectotype, details around ostium.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE ii
PLATE 12
FIG. 83. Etiella scitivittalis Walker, ostium and ductus bursae. FIG. 84. E. behrii Zeller, ostium and ductus bursae. FIG. 85. E. grisea drososcia Meyrick, secondary sac on bursa. FIG. 86. E. grisea grisea Hampson, ostium and ductus bursae. FIG. 87. E. grisea drososcia Meyrick, ostium and ductus bursae.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 12
86 W
87
PLATE 13
FIG. 88. Etiella hobsoni Butler, Solomons, ostium and ductus bursae. FIG. 89. E. hobsoni Butler, Australia, ostium and ductus bursae. FIG. 90. E. walsinghamella Ragonot, Australia, ostium and ductus bursae. FIG. 91. E. walsinghamella Ragonot, Papua, ostium and ductus bursae. FIG. 92. E. walsinghamella Ragonot, Japan, ostium and ductus bursae.
Bull. Br. Mus. not. Hist. (Ent.) 28, i
PLATE 13
PLATE 14
FIG. 93. Etiella scitivittalis Walker, signum in bursa.
FIG. 94. E. chrysoporella Meyrick, signum in bursa.
FIG. 95. E. grisea grisea Hampson, signum in bursa.
FIG. 96. E. hobsoni Butler, Formosa, signum in bursa.
FIG. 97. E. hobsoni Butler, Australia, signum in bursa.
FIG. 98. E. hobsoni Butler, Solomons, signum in bursa.
Bull. BY. Mus. nat. Hist. (Ent.) 28, i
PLATE 14
97
PLATE 15
FIG. 99. Etiella walsinghamella Ragonot, Australia, signum in bursa. FIG. 100. E. walsinghamella Ragonot, Papua, signum in bursa. FIG. 101. E. walsinghamella Ragonot, Japan, signum in bursa. FIG. 102. E. zinckenella Treitschke, Europe, lectotype, signum in bursa. FIG. 103. E. behrii Zeller, Australia, lectotype, signum in bursa.
Bull. Br. Mus. nat. Hist. (Ent.) 28, i
PLATE 15
A LIST OF SUPPLEMENTS TO THE ENTOMOLOGICAL SERIES
OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)
3. WATSON, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20.
4. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September,
I965. £3-25-
5. AHMAD, I. The Leptocorisinae (Heteroptera: Alydidae) of the World. Pp. 156: 475 text-figures. November, 1965. (out of print) £2.15.
6. OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3.
7. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967.
£3.15.
8. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the
world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. £3.50.
9. HEMMING, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972.
10. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. £8.
11. MOUND, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4.
12. WATSON, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5.
13. AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. £5.
14. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: i plate, 331 text-figures. July, 1969. £4.75.
15. ELIOT, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4.
16. GRAHAM, M. W. R.- DE V. The Pteromalidae of North-Western Europe (Hymenoptera : Chalcidoidea). Pp. 908: 686 text-figures. November, 1969.
17. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 68 plates, 15 text-figures. October, 1971. £12.
18. SANDS, W. A. The Soldierless Termites of Africa (Isoptera Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90.
Printed in England by Staples Printers Limited at their Kettering, Northants establishment
2 8 MAR 1973
G. B. BUCKTON'S WORKS ON APHIDOIDEA
/
J. P. DONCASTER
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 28 No. 2
LONDON : 1973
G. B. BUCKTON'S WORKS ON APHIDOIDEA (HEMIPTERA)
BY
JOHN PRIESTMAN DONCASTER
Pp. 23-109; 6 Plates, 54 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol.28 No. 2
LONDON: 1973
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted, in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 28, No. 2 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals.
World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1973
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 6 March, 1973 Price £3.55
G. B. BUCKTON'S WORKS ON APHIDOIDEA (HEMIPTERA)
By J. P. DONCASTER
CONTENTS
Page
SYNOPSIS ........... 25
INTRODUCTION ........... 25
ACKNOWLEDGEMENTS ......... 28
LIST OF BUCKTONIAN SPECIES ........ 28
LIST OF BUCKTONIAN GENERA ........ 30
BUCKTON'S APHID SPECIES ........ 30
LIST OF NON-BUCKTONIAN SPECIES ....... 87
BIBLIOGRAPHY ........... 96
INDEX ............ 103
SYNOPSIS
G. B. Buckton's slide collection, original drawings and notes are used to revise his published work on aphids. Of the 54 aphid species and 2 varieties he described as new, 15 are here accepted as valid, 38 are synonyms (5 newly established) and 3 are nomina dubia. Lectotypes have been designated for 51 species and the valid species redescribed and figured.
INTRODUCTION
SOME twenty-five years after Francis Walker had laid the foundations for our know- ledge of the British aphid fauna, George Bowdler Buckton published his Monograph of the British Aphides, which was to remain the standard work on the subject for the next forty years. Its four volumes, produced by the Ray Society in 1876, 1879, 1881 and 1883, contain descriptions of some 170 British aphids, each illustrated in colour, together with a conspectus of contemporary knowledge of the Aphidoidea as a whole. After Buckton's death in 1905 his collection of some 650 balsam mounts of aphids, together with his original drawings for the coloured plates in the monograph and a quantity of manuscript notes were acquired by the Trustees of the British Museum (Natural History). This collection comprises nearly all the material on which Buckton's descriptions of aphids were based and it is my aim here to use it as the means to revise his taxonomic work in the light of present knowledge.
Buckton was a man of varied interests who devoted most of the earlier part of his career to chemistry and physics. It was after he married and moved to Weycombe, near Haslemere, Surrey, about 1865 that he turned his energies fully to the study of natural history, and entomology in particular. (A sympathetic account of his life is given by W. F. Kirby in his obituary of Buckton (Kirby, 1907).) An injury in childhood left him permanently crippled and seriously curtailed his physical activities.
26 J. P. DONCASTER
Nevertheless, it seems likely that more than half of the aphids described and figured in his monograph he collected himself in Weycombe and its immediate neighbourhood. Many more were sent to him from other parts of Britain by friends and acquaintances whose help he acknowledges in the text. Foremost among them was Francis Walker from whom he received numerous consignments of aphids as well as many of Walker's own balsam mounts. Other notable contributors included Charles Barrett, who sent aphids from East Anglia and South Wales, James Hardy, who sent others from the Cheviot Hills, Sir John Lubbock, who sent subterranean aphids from Beckenham, Kent, and Professor James Trail, who supplied several species from Aberdeen. Also mentioned in the text or in Buckton's manuscript notes are the following correspondents, with the localities from which they sent material: J. Anderson (Chichester, Sussex), Joseph Anderson (Alresford, Hants), Rev. N. Andrews (Southwater, Sussex), G. C. Bignell (Plymouth), Rev. E. N. Bloomfield (Hastings, Sussex), Mr Borrer (Cowfold, Sussex), Hon. J. T. Boscawen (Probus, Cornwall), Mr Brady (Rainham, Essex), T. Brown (Cambridge), Dr Evershed (Shere, Surrey), E. A. Fitch (Maldon, Essex), Mr Foran (Eastbourne, Sussex), Miss Henry (Lurgashall, Sussex), Mr Knaggs (Kentish Town, London), R. McLachlan (Cornwall), E. Newman (Cambridge), James Salter (Basingstoke, Hants), Miss Salvin (Hawksfold, Sussex), Alfred Smee (Carshalton, Surrey), and Frederick Walker (Abingdon, Berks). Buckton also kept up a lively correspondence with Jules Lichtenstein in Montpellier, from whom he received specimens of some two dozen species of aphids from southern France, including balsam mounts made by Lichten- stein and his friend Richter.
In compiling his monograph, Buckton's aim was to include only those aphids which he could examine and sketch while still alive. His method was first to anaesthetize an aphid, then attach its body to a spot of Canada balsam on a slide, spread and attach legs and wings to adjacent spots of balsam, and draw it in outline with the camera lucida. Then, with water-colour, he would fill in the details of its colour, markings, shadows and highlights so as to portray the insect as nearly as possible as it appeared in life. Finally he would measure the wing-span and lengths of body, antenna and siphunculus before adding more balsam and a coverglass to make a permanent preparation. (Buckton, 1876 : i-iii; 1883 : 190-193.)
All Buckton's slides are balsam mounts, the majority made by himself. His collection also included 69 slides made and labelled by Walker (now incorporated with the Walker Collection in the British Museum (Natural History)), and a few, usually with mica covers, acquired from Lichtenstein. Buckton's own slide labels are sketchy and inadequate, with date, locality and hostplant data often wanting. The paired code letters which are attached to nearly every slide are of little help in this respect and seem to relate only to Buckton's system of slide storage. In a few instances, however, where the same code letters appear on a slide and on an original drawing, they may afford some additional evidence of identification.
The sheets bearing the watercolour sketches, used as models for the 134 plates1 of aphids in his monograph, have proved of great value in preparing this revision.
1The published plates total 141, of which two are of fossil aphids only, and five are of aphid parasites and predators. Originals of six aphid plates are missing.
BUCKTON'S WORKS ON APHIDOIDEA 27
Not only are the drawings more accurate, especially in colour characters, than the published lithographs, but the sheets also bear Buckton's own manuscript notes from which his descriptions were compiled, together with records of hostplants, localities, dates, collectors, and other data which often do not appear either on slide labels or in the monograph. Sometimes the pencilled comments, alterations and corrections which he added to these notes have also provided clues to what he was describing. Moreover, his method of working has made it possible, in many cases where data are lacking, to identify his types by comparing his sketches with the mounted aphids. Making allowance for distortion of the specimen from the addition of the coverglass and subsequent shrinkage in balsam, its attitude of body and the positions of the appendages often resemble the sketch so closely that there can be no doubt that one relates to the other. The identity of some of his types has been established by this means alone, and that of many more confirmed by such comparisons. For this reason I have been reluctant in many cases to remount aphids which I believe to be types on account of their similarity in posture to Buckton's sketches. His published measurements, however, are a less reliable check, as Buckton himself points out (1876 : i-ii), 'on account of the unequal foreshortening of the limbs, &c.' when a live insect is measured by means of the camera lucida.
Often the only precise date relating to a specimen is that on the sheet of original sketches, which seems to be the date on which he drew them and not necessarily the date of collection. A date on a slide label, when given, is usually a day or two later than that on the corresponding sketch, a result of his practice of mounting the aphids after he had finished drawing them.
Buckton's monograph represents nearly the whole of his contribution to aphidology. After its publication he described only fifteen further aphid species: eleven from material sent to him from the Indian Region (Buckton, 18890;, 18896, i8gia, 18916, 18930;, 18936, 1893^, 1896, 18990, 18996), and four more from Britain (Buckton, 1886, 1901). In all he published 58 species-group names as new, 15 of which are here accepted as valid, 38 are synonyms of other species, three must be regarded as nomina dubia, one is a nomen nudum and one belongs in the Coccoidea. He also erected nine new genera (excluding those erected for fossil aphids), of which five are currently valid.
Buckton designated no types, but his slides contain specimens of nearly all the species he described, and with the help of his original drawings, manuscript notes and published data it is possible to establish most of his types with reasonable certainty.
Frederick Laing during his curatorship of the aphid collections in the British Museum (Natural History) remounted a number of Buckton's specimens, some of which he marked as types, but he published no type-designations. Theobald also examined many of them and records his opinions in his own work on British aphids (Theobald, 1926, 1927, 1929). Many more of Buckton's specimens have been re- moved from their original mounts, cleared and remounted in gum-chloral mixture by me for the purposes of the present work.
I have been able to designate lectotypes for all but five of the 54 species and two varieties he described as new (disregarding the coccid) as well as locating many of the specimens on which his other descriptions and figures are based. All the types of
28 J. P. DONCASTER
Buckton's British aphid species are in the British Museum (Natural History), London. Of the species he described from the Indian Region, the types of Pemphigus aedificator and Therioaphis maculata are held by the Zoological Survey of India, Calcutta ; and others are in London, except that of Pyrolachnus Pyri, the location of which is unknown to me.
In the following revision, names used for the current identifications (given in bold type) of Buckton's British species conform as far as possible with those in Kloet & Hincks, A Check List of British Insects, 2nd Edition, Part I, 1964.
Numbers in italics refer to slides in the Buckton Collection in the British Museum (Natural History), hereafter abbreviated to BMNH.
'W.' preceding a slide number indicates a Walkerian slide originally in the Buckton Collection.
An asterisk (*) following a slide number indicates that the specimen (s) have been remounted.
ACKNOWLEDGEMENTS
I wish to express my thanks to the many colleagues who have so generously given me material, information, advice and encouragement in the preparation of this revision, notably Mr D. Hille Ris Lambers, Dr F. Leclant, Dr M. D. Leonard, Dr M. Miyazaki, Dr J. Pettersson, Professor A. G. Robinson and Mr H. L. G. Stroyan. I am especially grateful to the Director of the Zoological Survey of India, Calcutta, for lending me type-material of maculata Buckton, to Dr C. I. Carter for identifying many of Buckton's adelgids, and to Dr V. F. Eastop for much practical help. I am indebted to Mrs J. M. Palmer for her technical assistance and to Mr J. V. Brown for the photographs.
LIST OF BUCKTONIAN SPECIES Those with lectotypes are marked with an asterisk (*).
i. Valid Species
*ampullata, Amphorophora *artemisiae, Cryptosiphum *aucupariae, Dysaphis (Pomaphis) *bambusae, Astegopteryx *carnosum, Microlophium *circumflexum, Aulacorthum (Neomyzus) *crithmi, Dysaphis *cupressi, Cinara *immunis, Pemphigus *luteum, Macrosiphum (Sitobion) *muralis, Dactynotus *napaeus, Pemphigus *pilos^tm, Pterocomma Pyri, Pyrolachnus *viciae, Megoura
BUCKTON'S WORKS ON APHIDOIDEA 29
2. Synonyms
*acetosae, Aphis = Aphis acetosae L. * aedificator , Pemphigus = Baizongia pistaciae (L.) *atratus, Chermes = Adelges lands Vallot *bellis, Aphis = Brachycaudus helichrysi (Kaltenbach) *betulae, Chaitophorus = Callipterinella calliptera (Hartig) *betulina, Thelaxes = Glyphina betulae (L.) *carnosa, Endeis = Geoica eragrostidis (Passerini) *castaneae, Callipterus = Myzocallis castanicola Baker *cucurbitae, Aphis — Aphis gossypii Glover *dilineatus, Hyalopterus = Longicaudus trirhodus (Walker) *edentula, Aphis (syn. n.) = Rhopalosiphum insertum (Walker) *fodiens, Schizoneura = Schizoneura ulmi (L.) *formicina, Endeis = Baizongia pistaciae (L.) *fuliginosa, Schizoneura = Schizolachnus pineti (F.) *fuliginosus, Lachnus = Tuberolachnus salignus (Gmelin) var. glauca, Siphonophora rosae = Macrosiphum rosae (L.) *gracilis, Myzus = Metopolophium dirhodum (Walker) *graminis, Rhizobius = Aploneura lentisci (Passerini) *instabilis, Aphis (syn. n.) = Brachycaudus cardui (L.) *lentiginis, Aphis (syn. n.) = Dysaphis (Pomaphis) plantaginea (Passerini) *longipennis, Siphonophora — Metopolophium dirhodum (Walker) *macrocephalus, Lachnus = Cinara pinicola (Kaltenbach) *maculatus, Chaitophorus = Therioaphis trifolii (Monell) *melanocephalus, Hyalopterus = Hayhurstia cucubali (Passerini) *menthae, Siphonophora = Aulacorthum solani (Kaltenbach) *olivata, Siphonophora = Dactynotus cirsii (L.) *opima, Aphis = Brachycaudus cardui (L.) *pedicularis, Aphis = Aphis nasturtii Kaltenbach *pellucida, Endeis = Geoica eragrostidis (Passerini) *penicillata, Aphis (syn. n.) = Aphis grossulariae Kaltenbach *petasitidis, Aphis = Brachycaudus helichrysi (Kaltenbach) *pilosa, Glyphina = Schizolachnus pineti (F.) *polygoni, Siphonophora = Nasonovia ribisnigri (Mosley) *var. rufa, Siphonophora rubi = Macrosiphum funestum (Macchiati) *scrophulariae, Siphonophora (syn. n.) — Cryptomyzus galeopsidis (Kaltenbach) *sisymbrii, Siphonophora = Dactynotus cichorii (Koch) *theaecola, Ceylonia = Toxoptera aurantii (Boyer de Fonscolombe) *viridana, Forda = Forda formicaria Heyden
3. Nomina dubia coccus, Pemphigus formicophilus, Lachnus taxi, Chermes
30 J. P. DONCASTER
4. Nomen nudum cinchonae, Pemphigus
5. Non-aphid jujubae, Rhizobius: in Coccoidea, Margarodidae
LIST OF BUCKTONIAN GENERA (Fossil aphids excluded)
Amphorophora, 1876 : 187; type-species A. ampullata Buckton, 1876.
Brachycolus, 1879 : T4^i type-species Aphis stellariae Hardy, 1850.
Ceylonia, 18910 : 34; type-species C. theaecola Buckton, 1891 (= Aphis aurantii
Boyer de Fonscolombe, 1841.)
Cryptosiphum, 1879 : J44 '• type-species C. artemisiae Buckton, 1879. Megoura, 1876 : 188 ; type-species M. viciae Buckton, 1876. Melanoxanthus, 1879 : 2I '• type-species Aphis salicis Linnaeus, 1758. Oregma, 18936 : 87; type-species 0. bambusae Buckton, 1893. Pterocomma, 1879 : 142 ; type-species P. pilosum Buckton, 1879. Ptychodes, 1881 : 39; type-species Aphis juglandis Goeze, 1778.
Ceylonia is a synonym of Toxoptera Koch, Melanoxanthus of Pterocomma Buckton, Oregma of Astegopteryx Karsch, and Ptychodes of Callaphis Walker.
BUCKTON'S APHID SPECIES
Aphis acetosae Buckton = Aphis acetosae Linnaeus
Aphis acetosae Linnaeus, 1767 : 734.
Aphis acetosae Linnaeus; Fabricius, 1775 : 739.
Aphis acetosae Buckton, 1879 : 80; pi. 62, figs 5, 6, 7. [Synonymized by Borner, 1952 : 77.]
LECTOTYPE here designated: alate viviparous female. Locality not stated. Rumex acetosa. 26.vi.(year ?). (Buckton ?).
Paralectotypes. 2 apterous viviparous females, 5 alate viviparous females, 6 nymphs. Data as lectotype. (426*, 4260*, 4266*, 4260*}.
BIOMETRIC DATA. Lectotype, alata: body length 2-30 mm, antennal flagellum 1-36 mm, ratios of segments III-VI 45 : 25 : 22 : 14 + 35, siphunculus 0-30 mm, cauda 0-16 mm, caudal hairs 15, ultimate rostral segment 0-138 mm, second segment of hind tarsus 0-138 mm, hairs on eighth tergite 7, articular diameter of antennal segment III 31^1, longest hair on ant. seg. Ill I2(z, on hind femur 35(1, on hind tibia 38^, on abdominal tergite III IOJJL, on abd. terg. VIII 18(1, secondary rhinaria on ant. seg. Ill 7 and 8.
Paralectotype, aptera: body length 2-64 mm, ant. flag. 1-27 mm, ratios of segs III-VI 37 : 23 : 21 : 14 -f 32, siph. 0-39 mm, cauda 0-22 mm, caudal hairs 13, ult. rost. seg. 0-146 mm, second seg. hind tarsus 0-139 mm, hairs on eighth tergite 5, artic. diam. ant. seg. Ill 32^, longest hair on ant. seg. Ill i2(jL, on hind femur 25^, on hind tibia 30^, on eighth tergite 2o;z.
Buckton describes this species as new and figures the apterous and alate viviparous
BUCKTON'S WORKS ON APHIDOIDEA 31
females and the nymph from specimens taken, according to his published account, on Rumex acetosa. The manuscript notes accompanying his original sketches for the figures indicate, however, that the aptera he used for figure 5 was taken on thistle, and only the nymph and alata are assigned to Rumex acetosa. There are no slides in the Buckton Collection named acetosae, but one, labelled in Buckton's hand 'A. rumicis on Sorrel', contained the specimens listed above, all of which belong in the Aphis acetosae L. species-complex and include, I believe, the types of at least the alata and nymph which Buckton describes and figures. I have not found a specimen which I can identify with the sketch of the aptera said to have been taken on thistle.
Stroyan (1955 : 309 and 19576 : 354) discusses and compares two forms of Aphis acetosae L. which occur on Rumex acetosa and R. acetosella in Britain. Comparison of Buckton's material with Stroyan's shows a close similarity between both Buckton's apterae and alatae and the form associated with R. acetosa, which is characterized by a nearly solid black dorsal abdominal patch in apterae and transverse segmental dark bands on the abdominal dorsum in alatae. Indeed, the only considerable difference is in the lengths of the hairs on body and appendages, which in Buckton's specimens are on average about half as long as those of Stroyan's, in this respect, as also in having tibial hairs which are longer on the inner side than on the outer, resembling more closely the short haired form on R. acetosella.
Through the kindness of Dr Jan Pettersson of Uppsala I was also able to examine apterae of another form of Aphis acetosae, collected on Rumex crispus in Sweden by Dr F. Ossiannilsson, which showed many similarities with Buckton's specimens. In this case, however, the most obvious agreement was in the shortness of the hairs on body and appendages which came very close to those of Buckton's specimens. The aphids from R. crispus all had the nearly complete sclerotic tergal patch but were smaller in size than either Buckton's or Stroyan's. Buckton's sample, however, is too small to allow clear distinctions to be drawn from such comparisons and for the present I regard his species as one member of the Aphis acetosae L. species-complex.
Pemphigus aedificator Buckton = Baizongia pistaciae (L.)
Aphis pistaciae Linnaeus, 1767 : 737.
Aphis pistaciae Fabricius, 1775 '. 739.
Baizongia pistaciae (F.) Rondani, 1848 : 35.
Pemphigus cornicularius Passerini, 1856 : 261.
Endeis formicina Buckton, 1883 : 91.
Pemphigus aedificator Buckton, 1893^ : 71.
Pemphigella cornicularia (Passerini) Tullgren, 1909 : 171.
PNeorhizobius stramineus del Guercio, 1917 : 249.
Pemphigus pistaciae (L.) Wilson & Vickery, 1918 : 131.
Dasia aedificator (Buckton) van der Goot in Das, 1918 : 144.
[Tycheoides setariae (Passerini) Theobald, 1929 : 183. Misidentification.]
[Pemphigus aedificator Buckton; Takahashi, 1933 : 352. Misidentification.]
Baizongia oestlundi Hottes, 1949 : 86.
Baizongia pistaciae (L.) Davatchi, 1958 : 133.
Lectotype (designated by Doncaster, 1969 : 157) : alate viviparous female
32 J. P. DONCASTER
(fundatrigenia migrans). PAKISTAN, Quetta. Pistacia terebinthus (?) galls. i6.xi. 1890 (Elliot}. (Zoological Survey of India, Calcutta, no. 7282/117.)
Paralectotypes : 20 alate, i apterous viviparous females. Same data as lectotype. (Z.S.I, nos 7283-7383/117. Slide no. 7286/117 presented to BMNH).
I have already dealt elsewhere (Doncaster, 1969) with aedificator Buckton, including descriptions of lectotype and paralectotype. The name was first synony- mized with Baizongia pistaciae (L.) by Davatchi (1958 : 133).
Amphorophora ampullata Buckton (PI. i, fig. 55; Text-figs 1-5)
Amphorophora ampullata Buckton, 1876 : 187; pi. 37, fig. 4.
Rhopalosiphum ampullatum (Buckton) Oestlund, 1887 : 77.
Rhopalosiphum ampullatum (Buckton); van der Goot, 1915 : 142.
Megoura dryopteridis Matsumura, 1918 : 13. [Synonymized by Miyazaki, 1968 : 14.]
Acyrthosiphon (Amphorophora) ampullatum (Buckton) Mordvilko, 1919 : 247.
Amphorophora ampullata Buckton; Mason, 1925 : 10.
Amphorophora ampullata Buckton; Theobald, 1926 : 191.
Amphorophora shidae Shinji, 1933 : 348. [Synonymized by Miyazaki, 1968 : 14.]
Amphorophora ampullata Buckton; Knechtel & Manolache, 1945 ; 484.
Amphorophora ampullata Buckton; Hille Ris Lambers, 1947 : 231.
Amphorophora dryopteridis (Matsumura) Moritsu, 1948 : 83.
Amphorophora ampullata Buckton; Borner, 1952 : 175.
Amphorophora dryopteridis (Matsumura); Paik, 1965 : 73.
Amphorophora ampullata Buckton; Hille Ris Lambers & Basu, 1966 : 14, 15.
Amphorophora ampullata Buckton; Robinson, 1966 : 1253.
Amphorophora ampullata Buckton; Miyazaki, 1968 : 14-17.
Amphorophora ampullata Buckton; Heie, 1969 : 383. [New host record.]
LECTOTYPE here designated: apterous viviparous female. Sussex, Lurgashall. Cystopteris mo ntana in greenhouse. 13. ix. (year ?). (Miss Henry). (37*).
Paralectotypes: 2 apterous viviparous females. Data as lectotype. (37<*>*, 38*.}
Apterous viviparous female. (Plate i, fig. 55; Text-figs 1-5). Colour of macerated specimen: uniformly pale yellowish, except for slight darkening at articulation of antennal segments III and IV, and IV and V; at apices of fore and middle tibiae, and apices of siphunculi. Morphology : body large, broadly oval, 3-28-4-05 mm long. Head smooth, dorsal hairs 52-80(1 long, with blunt apices. Antennal tubercles large, diverging, each ventrally with a prominent hemispherical protuberance bearing one or two hairs. First antennal segment with 9-10 hairs, some spinules on the ventral surface near the base, and some imbrications on the inner surface near the apex. Second segment partly spinulose. Third segment spinulose at the base, remainder smooth, with from 26-39 secondary rhinaria, more or less in line, over nearly its whole length; hairs short, stout, blunt, up to 36^ long. Fourth and fifth segments faintly imbricated, sixth with processus terminalis 4-8 times as long as base (in the only specimen with a complete antenna). Rostrum scarcely reaching hind coxae, ultimate segment rather broad with straight sides tapering towards a blunt apex, o-i6o-o-i74 mm long, about equal in length to second segment of hind tarsus, and with 10-12 non-apical hairs. Femora and tibiae slightly scabrous apically, otherwise smooth. Femoral hairs stout, blunt, the longest reaching about 6o(x. Tibial hairs more acute, more numerous, the longest about 70^. First segments of tarsi on all legs with 3 hairs. Tergum
BUCKTON'S WORKS ON APHIDOIDEA
33
0-2
FIGS 1-5. Amphorophora ampullata Buckton. Lectotype: Fig. i. Head, upper (left) and lower surfaces. Fig. 2. Siphunculus. Fig. 3. Third antennal segment. Fig. 4.
| Cauda. Paratype (370) : Fig. 5. Underside of left antennal tubercle to show protuberance (see text). (Figs 1-4 to same scale.)
34 J- P. DONCASTER
smooth, membranous, sparsely clothed with thick blunt hairs, only about 15-20(0, long on anterior tergites, longer on sixth, seventh and eighth. Eighth tergite with 6-8 hairs, the longest about joyL. Hairs on sternites rather long (iyofx), fine, acute, rather numerous. Siphunculi smooth, except for a few apical striae below the flange, base expanded, basal two-fifths narrow, apical three-fifths evenly swollen, the diameter at the widest part about twice that at the narrowest, 2-3 times as long as the cauda and a little less than a quarter of the body length. Cauda obtuse, 1-7-2 times as long as its basal width, with 18-19 hairs. Subgenital plate with 4 hairs near anterior margin and 10-12 posteriorly.
NOTES. Amphorophora ampullata Buckton sensu latiore has been recorded from western and northern Europe, India, Korea, Japan, and North America. Its food- plants are restricted to ferns, the commonest being species of Athyrium and Dryopteris in the Old World, and of Onoclea in North America.
Hille Ris Lambers and Basu (1966 : 14, 15) distinguish two subspecies of ampullata : bengalensis, based on material from ferns in India, and laingi Mason, 1925, from Onoclea sensibilis, etc., in the U.S.A.
Cryptosiphum artemisiae Buckton (PI. i, fig. 56; Text-figs 6-8)
Aphis gallarum Kaltenbach, 1856 : 236. [Homonym of Aphis gallarum Gmelin, 1790 : 2210.] Aphis artemisiae Passerini, 1860 : 35. [Homonym of Aphis artemisiae Boyer de Fonscolombe,
1841 : 162.]
Cryptosiphum artemisiae Buckton, 1879 : 145; pi. 84, figs 1-4. Pseudolachnus yomogi Shinji, 1922 : 730. [Synonymized by Monzen, 1929 : 48; Takahashi,
1931 : 37, Shinji, 1941 : 623.] Cryptosiphum pseudogallarum Shinji, 1941 : 626-628. [Synonymized by Tao, 1962 : 96.]
LECTOTYPE here designated: apterous viviparous female. Norfolk, Norwich, Brandon. Artemisia vulgaris (galls). 2.viii.(year ?). (Barrett). (42*).
Paralectotypes : 4 apterous viviparous females, I nymph, 5 alate viviparous females, same data as lectotype. (41*, 42a*, 42b*, 43*).
Apterous viviparous female. (Plate i, fig. 56; Text-figs 6-8). Colour of macerated specimen: body pale brown; head, antennae, rostrum, subgenital and anal plates rather darker; legs and cauda darker still. Morphology : body 1-31-1 -66 mm long, broadly oval, about 1-25 times as long as wide. Head short, broad, smooth, frons slightly convex, antennal tubercles absent. Compound eyes rather small, triommatidia scarcely projecting. Cephalic hairs sparse, fine, acute, from 22-32^ long. Antennae short, about 0-25 of the length of the body, of 6 segments, the length ratios of III-VI about 13 : 6 : 6 : 7 -f 6. Antennal hairs sparse, acute, up to about i2(ji long. Primary rhinaria, and the 5-6 accessories around that on VI, heavily fringed. Segments I-III smooth, IV-VI imbricated. Rostrum reaching to or a little beyond second coxae, ultimate segment stiletto-shaped with narrow elongate apex, 0-119-0-134 mm long, about 2-5 times as long as its basal width and from 1-2 to 1-4 times as long as hind tarsus II (0-095 mm). Two of the three normal pairs of apical rostral hairs are set at about one-sixth of its length from the apex, the third pair are displaced almost to the middle of the segment. There are 6 non-apical hairs in the lectotype, 4 or 5 in paratype apterae. Legs short; the femora, which are fused to the trochanters, stout, nearly smooth, but middle and hind femora with a few short rows of spinules on the under surface ; femoral hairs sparse, fine, acute, from 20-24^1 long. Tibial hairs more numerous, some on inner side stouter and spiny, the rest fine, acute, 24-32^1
BUCKTON'S WORKS ON APHIDOIDEA
35
long. Hind tibia about 0-2 of body length. First tarsal chaetotaxy 3, 3, 2. Tergum smooth, with few rather long (32-38^) fine hairs. Siphunculi very small, the opening, scarcely larger than a spiracle, borne on a shallow protuberance. Cauda very short, broad, slightly convex, conforming to the oval outline of the body, with 5 or 6 hairs. Eighth tergite with 6-8 hairs 40-4851 long. Ventral hairs shorter and more numerous, sternites with irregular transverse rows of spinules. Subgenital plate transversely elongate with 5-10 hairs on anterior half and 13-16 along posterior border.
Alate viviparous female (43*). Colour: body pale brown, head, thorax, antennae and legs slightly and evenly darker. Differs from aptera mainly in the narrower and slightly shorter body and relatively longer appendages. The antennae are just over half, and the hind tibiae about one-third of the body length respectively. Antennal segments III-VI are imbricated, and III bears 26-30 circular or oval, rather large secondary rhinaria with wide rims, irregularly arranged over the distal three-quarters of the segment; IV with 2-5 similar rhinaria grouped closely near its apex. Segment III joined to II by a narrow stalk, about half the diameter of the rest of the segment. Wing venation normal aphidine, but in this specimen the media in one fore wing has both its branches forked, an abnormality which Buckton records both in his original sketch of the alate artemisiae and in the published figure (plate 84, fig. 3).
NOTES. Buckton's sample of Cryptosiphum artemisiae includes one alate viviparous female of Color adoa artemisiae (del Guercio), the presence of which could account for the statement in his generic diagnosis of Cryptosiphum (1879 : 144), 'Cauda small, but distinctly seen in the winged forms', and his doubt, expressed two pages later, whether Passerini's artemisiae is the same insect. The originals for his published plate include drawings of both species. The coloured drawings of aptera, nymph and alata used for figs i, 2 and 3 are definitely based on artemisiae Buckton, but the end of an abdomen used for fig. 5 shows a prominent cauda and — in the original but not the reproduction — a distinct siphunculus. This, and the head, antenna and rostrum of fig. 6 would seem to have been taken from the alate Coloradoa. An unpublished sketch of head and antenna clearly belongs to an alate Cryptosiphum.
FIGS 6-8. Cryptosiphum artemisiae Buckton. Lectotype: Fig. 6. Left antenna. Paratype (420.) : Fig. 7. Head, upper (left) and lower surfaces. Fig. 8. Apex of rostrum.
36 J. P. DONCASTER
The sheet of originals is dated August 2nd. If, as I suspect, this is the date when Buckton drew them, the specimens must have been collected some days earlier, i.e. in late July, not 'early August' as published.
Chermes atratus Buckton = Adelges laricis Vallot
Adelges laricis Vallot, 1836 : 72. Chermes coccineus Ratzeburg, 1843 : 202. Chermes strobilobius Kaltenbach, 1843 : 203. Chermes atratus Buckton, 1883 : 39; pi. 120, figs 5, 6. (For full synonymy, see Carter, 1971 : 44.)
LECTOTYPE here designated: alate viviparous female. Surrey, Haslemere. Quercus sp. 2.vi.i87i (?). (Buckton). (159}.
Buckton describes only the alata which he took as a vagrant on oak. There are no specimens named atratus in his collection, but his notes indicate that the alata was collected together with specimens of Thelaxes dryophila. An alate Adelges laricis Vallot, mounted on 759 together with apterous and alate dryophila, agrees reasonably well with Buckton's description, measurements and original drawing, and this I take to be his type of atratus.
Aphis aucupariae Buckton = Dysaphis (Pomaphis) aucupariae (Buckton)
(PI. 2, fig. 57; Text-figs 9-13)
[Aphis sorbi Kaltenbach; Walker, 1850 : 276 partim. Misidentification.]
Aphis aucupariae Buckton, 1879 : 76; pi. 60, figs 3-5.
Anur aphis appelii Borner, 1926 : 225.
Anuraphis aucupariae (Buckton) Theobald, 1927 : 308.
Aphis (Dentatus?) aucupariae Buckton; Mordvilko, 1929 : 52.
Yezabura (Ceruraphis) aucupariae (Buckton) Borner & Schilder, 1932 : 586.
Sappaphis aucupariae (Buckton) Borner, 1952 : 97.
Sappaphis aucupariae (Buckton) ; Stroyan, ig^^a : 20.
Dysaphis (Pomaphis) aucupariae (Buckton) Stroyan, 1963 : 55.
Dysaphis aucupariae (Buckton) ; Shaposhnikov, 1964 : 582.
LECTOTYPE here designated: apterous viviparous female (fundatrigenia) . Sussex, Horsham, Cowfold. Sorbus torminalis. I7.vi.(year ?). (Probably Boner}. (466a*).
Paralectotypes : 7 nymphs, data as lectotype (4666*, 4660*}; 6 larvae, Sussex, Horsham. Sorbus torminalis. i8.v.(year ?). (Probably Boner}. (49*, 490,*, 49b*).
Other material: 3 alate viviparous females, 4 nymphs. Middlesex, Southgate. Sorbus torminalis. 2i.vi.i847. (Walker). (W. 939).
Apterous viviparous female (fundatrigenia). (Plate 2, fig. 57; Text-figs 9-13). Colour of macerated specimen: head, antennae except basal part of segment III, legs except femoral bases, and siphunculi dark to black sclerotic. Sclerotic bands on notum, dorsal sclerites on abdomen, anal and subgenital plates also dark. Morphology: body 2-47 mm long, broadly oval, about 1-4 times as long as broad. Head densely sclerotic, vertex without spinal tubercles,
BUCKTON'S WORKS ON APHIDOIDEA
37
wrinkled, with some scattered spinules, hairs fairly numerous, fine, acute, up to about 6o(jt long. Antennae with all six segments rather coarsely imbricated; hairs acute, the longest on III reaching 38^, slightly longer than articular diameter of segment; processus terminalis 2-6 times as long as base of VI; the flagellum 1-58 mm long, about two-thirds the length of the body; ratios of segments III-VI 48 : 34 : 25 : 14 + 36. Rostrum 0-53 mm long, reaching middle coxae, ultimate segment normal, 0-148 mm long, about twice as long as its basal width, with 4 non-apical hairs, and very slightly shorter than second segment of hind tarsus (0-154 mm). Femora rough and more or less scabrous, mainly on posterior surface, hairs fairly numerous, fine, acute, up to 64^ long. Tibial hairs shorter and stouter, up to 50^ long; hind tibia about half length of body. First tarsal segments with 3, 3, 3 hairs. Abdomen with marginal tubercles on segments II-IV on left side, I-IV on right. Spinal sclerites are present as irregular broken transverse bands on segments I, VII and VIII, and as irregular paired scleroites on the inter- vening segments. One pair only of spinal tubercles is present on segment VIII. Dorsal abdominal hairs are acute, the longest reaching 78-80^.. Eighth tergite with 5 hairs. Siphunculi 0-33 mm long, nearly straight, slightly and evenly tapering from base to apex, imbricated, with a slight annular constriction behind the flange, each 3-3 times as long as its basal width, nearly equal in length to antennal segment IV and just over one-eighth of the body length. Cauda bluntly triangular, 0-12 mm long, about two-thirds as long as its basal width, with 7 hairs. Subgenital plate with 3 hairs on anterior half and 19 irregularly arranged along posterior margin.
NOTES. From Buckton's notes and drawings I conclude that his published description of aucupariae is based on two samples of material, namely (a) six larvae taken probably at Cowfold, near Horsham, Sussex and dated i8th May, and (b)
13
0-2
FIGS 9-13. Dysaphis (Pomaphis) aucupariae (Buckton). Lectotype: Fig. 9. Head, upper (left) and lower surfaces. Fig. 10. Left antenna. Fig. n. Apex of rostrum. Fig. 12. Siphunculus. Fig. 13. Cauda.
38 J. P. DONCASTER
one adult fundatrigenia and seven nymphs taken at Cowfold and dated i7th June. His published account confirms that both larvae and nymphs were found in the same place. Both samples are from Sorbus torminalis; the first (a) was originally mounted on one slide (49) labelled 'Aphis aucuparia' by Buckton, but with no other data, and (b) on 466 labelled 'A. sorbi. Cowfold.' He drew the largest of the larvae in (a) for figure 3, which he regarded as the normal apterous female and described: description and measurements as well as the sketch correspond fairly well with the specimen. The adult fundatrigenia in sample (b) was his model for figure 4, the 'globose' form, which he thought might be the fundatrix. Measurements given in his notes, but not published, support this conclusion. Since the fundatrigenia is the only adult morph in either sample, and is also complete and well preserved, I choose it as lectotype of aucupariae Buckton.
A full account of aucupariae (Buckton) is given by Stroyan (19570 : 20-22).
Oregma bambusae Buckton = Astegopteryx bambusae (Buckton) (PI. 2, fig. 58; Text-figs 14-16)
Oregma bambusae Buckton, 18936 : 87, partim. Oregma lutescens van der Goot, 1917 : 197. Astegopteryx bambusae (Buckton) Doncaster, 1966 : 157.
Lectotype (designated by Doncaster, 1966 : 157) : apterous viviparous female.
FIGS 14-16. Astegopteryx bambusae (Buckton). Paratype 3: Fig. 14. Head, upper (left) and lower surfaces (wax glands shown stippled) . Paratype 26 : Fig. 15. Rostrum. Lectotype: Fig. 16. Left antenna.
BUCKTON'S WORKS ON APHIDOIDEA 39
INDIA, Uttar Pradesh, Dehra Dun. Bambusa arundinacea. Undated. (Cotes).
Paralectotypes : 23 apterous viv. females, i normal larva, i dimorphic larva. Data as lectotype. (52*, 53*, 530*, 54*, 55*, 56*, 532*). (Plate 2, fig. 58; Text- figs 14-16.)
The material on which Buckton bases Oregma bambusae is a mixture of two species, Astegopteryx bambusae (Buckton) and Pseudoregma bambusicola (Takahashi) (see Doncaster, 1966). All the material of true bambusae is in the Buckton Collection, BMNH.
Aphis bellis Buckton = Brachycaudus helichrysi (Kaltenbach) Buckton, 1879 : 98; pi. 69 bis, figs i, 2, 4.
LECTOTYPE here designated : apterous viviparous female. SCOTLAND, Aberdeen. Bellis per ennis. i8.ix.(year ?). (Trail}. (58).
Paralectotypes: 2 alate viviparous females, 2 nymphs. Data as lectotype. (5$).
BIOMETRIC DATA. Lectotype, apterous viviparous female: body length 1-49 mm, antennal flagellum 0-64 mm, ratios of segments III-VI 24 : 15 : 9 : 8 + 24, siphunculus o-n mm, cauda 0-069 mm, caudal hairs ?, ultimate rostral segment 0-104 mm, second segment of hind tarsus 0-089 mm, eighth tergite with 7 hairs, articular diameter of ant. seg. Ill 20(jt, longest hair on ant. seg. Ill IOJJL, on hind femur iSjji, on hind tibia 24^, on abd. terg. VIII 8opi.
Paralectotype, alata: body length 1-84 mm, ant. flag. 0-94 mm, ratios of segs III-VI 41 : 22 : 13 : ii -f 31, secondary rhinaria on III 27, on IV 8, on V o, siph. o-n mm, cauda o-io mm, caudal hairs 5, ult. rost. seg. 0-12 mm, second seg. hind tarsus o-io mm, eighth tergite with 7 hairs, artic. diam. ant. seg. Ill i8pi, longest hair on ant. seg. Ill i2[i, on hind femur 2o\i, on hind tibia 26^, on eighth tergite 4511.
Buckton's slide is in reasonable condition and has not been remounted. Laing has marked it helichrysi Kalt. Theobald (1927 : 285) regarded bellis as a doubtful synonym of helichrysi and subsequent authors have confidently identified it with Kaltenbach's species.
Buckton's original sketches of bellis show the aptera and nymph as dull brownish yellow or brown, which is much more characteristic of helichrysi than the brilliant yellow or yellow-green used to colour those morphs in the plate (at least in copies I have seen). I can find nothing in the mounted aptera which might be construed as the 'vermiform parasites' shown in both Buckton's original and the published figure.
Chaitophorus betulae Buckton = Callipterinella calliptera (Hartig)
Aphis calliptera Hartig, 1841 : 369.
Chaitophorus annulatus Koch, 1854 : 7. [Synonymized by van der Goot, 1912 : 278.]
Chaitophorus betulae Buckton, 1879 : 139; pi. 82, figs i, 2.
[Myzocallis betulae (Buckton) ; Kloet & Hincks, 1945 : 70. Misidentification.]
Calaphis callipterus (Hartig) Borner, 1952 : 58.
Procalaphis callipterus (Hartig) Quednau, 1954 : 23.
Callipterinella calliptera (Hartig) Stroyan in Kloet & Hincks, 1964 : 70.
4o J. P. DONCASTER
LECTOTYPE here designated : ovipara. Essex, Wanstead. Betula. ix. (year?). (Wallace ': ? error for Walker, who collected in Wanstead). (61*).
Paralectotypes : 4 larvae. Data as lectotype. (62).
BIOMETRIC DATA. Lectotype, ovipara: body length 2-48 mm, antennal flagellum 1-07 mm, ratios of segments III-VI 56 : 30 : 22 : 15 -f ?, siphunculus? (incomplete), cauda ? (missing), ultimate rostral segment 0-12 mm, second segment of hind tarsus 0-14 mm, eighth tergite with 12 ? hairs, articular diameter of ant. seg. Ill 35^, longest hair on ant. seg. Ill 23(1, on hind femur 8o[z, on hind tibia ioo[x, on abd. terg. VIII
Buckton describes the apterous viviparous female and the ovipara. His description and figure of the aptera seem likely to be based on the largest of the larvae on 62. Published measurements agree reasonably well with those of this specimen ; further- more, his original sketch, though not the published lithograph, shows larva-like antennae with the third segment indistinctly, or not, divided, and an undifferentiated cauda. His manuscript notes include the comment 'probably this specimen is not quite mature . . .' His description and figure of the ovipara correspond with the ovipara on 61, which has been remounted by Laing and marked Type. I designate this specimen as lectotype of betulae Buckton.
Theobald (1929 : 349-350) quotes Buckton's description and adds his own descrip- tions of ovipara and male; but his specimens (from Betula alba, Boxmoor, Herts, 23.x. 1913) are sexuales of Betulaphis quadrituberculata (Kaltenbach).
Thelaxes betulina Buckton = Glyphina betulae (L.)
Aphis betulae Linnaeus, 1758 : 452.
Vacuna betulae Kaltenbach, 1843 : 177.
Aphis impingens Walker, 1852 : 1042.
Thelaxes betulina Buckton, 1886 : 326; pi. 6, figs 1-6.
Glyphina ? betulae (Linnaeus) Borner, 1952 : 181.
Glyphina betulae (Linnaeus); Stroyan in Kloet & Hincks, 1964 : 84.
LECTOTYPE here designated: apterous viviparous female. Sussex, Hastings, Guestling. Betula. vi. (year ?). (Bloomfield) . (72*).
Paralectotypes: 10 apterous, 3 alate viviparous females, i nymph, i larva. Data as lectotype. (72*, 73, 74).
BIOMETRIC DATA. Lectotype, aptera: body length 1-88 mm, antennal flagellum 0-51 mm, ratios of segments III-V 33 : 12 : 16 + 4, siphuncular diameter 44^, cauda not measurable, caudal hairs 9?, ultimate rostral segment 0-17 mm, second segment of hind tarsus 0-15 mm, eighth tergite with 5 hairs, articular diameter of ant. seg. Ill 3O(x, longest hair on ant. seg. Ill 45[i, on hind femur Sopi, on hind tibia 6o[x, on eighth tergite
Paralectotype alata: body length 1-76 mm, ant. flag. 0-58 mm, ratios segs III-V 39 : 15 : 16 -)- 3, secondary rhinaria on III 6, on IV o, siph. diam. 42^, cauda 0-085 mm, caudal hairs 6, ult. rost. seg. 0-18 mm, second seg. hind tarsus 0-14 mm, eighth tergite with 9 hairs, artic. diam. ant. seg. Ill 20(1, longest hair on ant. seg. Ill ±50jz, on hind femur ±50^, on hind tibia ±70^, °n eighth tergite ±90^ .
Buckton describes the apterous and alate viviparous females and figures two
BUCKTON'S WORKS ON APHIDOIDEA 41
apterae (one 'of a later brood'), a nymph and an alata. I have not succeeded in locating the originals of these figures. (See also Glyphina betulae, p. 88.)
Endeis carnosa Buckton = Geoica eragrostidis (Passerini)
Tychea eragrostidis Passerini, 1860 : 39.
Tychea setariae Passerini, 1860 : 40.
[Tychea setulosa Passerini; Buckton, 1883 : 87. Misidentification.]
Endeis pellucida Buckton, 1883 : 91.
Endeis carnosa Buckton, 1883 : 92; pi. 129, figs 5-8.
[Geoica utricularia sensu auctt. nee Passerini, 1856 : 260. Misidentifications.]
[Geoica squamosa Hart; Theobald, 1929 : 191. Misidentification.]
Geoica discreta Borner, 1952 : 203.
LECTOTYPE here designated: apterous viviparous female. Kent, Beckenham, in ants' nest. ii.i876. (Lubbock). (89*).
BIOMETRIC DATA. Lectotype, aptera: body length 1-78 mm, whole antenna 0-48 mm, ratios of antennal segments I-V u : 9 : 17 19 : 13, ultimate rostral segment 0-19 mm, second segment of hind tarsus 0-12 mm.
Buckton describes and figures the apterous viviparous female. The drawings of the whole insect (fig. 5) both in the lithograph and in the original are very crude, but the detail drawings of the hind end (fig. 6), and the vertex and antenna (fig. 7), relate without question to a specimen of Geoica eragrostidis (Passerini) received with other specimens from Lubbock. In its original mount this specimen was considerably shrunken, and the posterior abdominal segments appeared much as in Buckton's figure 6, with the rectangular anal plate protruding beyond them. In his MS notes and in his published description he interprets the anal plate as the cauda, but the caption to the figure refers to it as the ovipositor. His sketch for fig. 7 shows the flabellate hairs on the vertex, and the 5-jointed antenna in which the length ratios of the segments are about right— i.e. with III the longest — and not as he describes them as being, all 'nearly equal'.
Siphonophora carnosa Buckton — Microlophium carnosum (Buckton) (PI. 3, fig. 59; Text-figs 17-21)
[Aphis urticae Linnaeus, 1758 : 453 ( — Orthezia urticae (L.) in Coccoidea; type in Linnaean Society Collection, London) ; Schrank, 1801 : 106, Kaltenbach, 1843 : 13. Misidentifications.]
[Siphonophora urticae (sensu Schrank non L.) Koch, 1855 : 154.]
[Siphonophora urticae (sensu Schrank non L.) ; Buckton, 1876 : 143.]
Siphonophora carnosa Buckton, 1876 : 144; pi. 20, figs 1-4.
[Macrosiphum urticae (sensu Schrank non L.) Schouteden, 19060 : 241.]
[Acyrthosiphon (Microlophium) urticae urticae (sensu Schrank non L.) Mordvilko, 1914 : 202.]
Amphorophora evansi Theobald, 1923 : 24; 1926 : 193.
Macrosiphum schranki Theobald, 1927 : 403.
Macrosiphum carnosum (Buckton) Lindinger, 1932 : 277.
Macrosiphon carnosus (Buckton); Borner & Schilder, 1932 : 628.
Acyrthosiphon carnosum (Buckton) Hille Ris Lambers, 1933 : 171.
Acyrthosiphon (Microlophium) carnosum (Buckton) Kloet & Hincks, 1945 : 63.
Microlophium evansi (Theobald) Hille Ris Lambers, 1949 : 209, Borner, 1952 : 142, Stroyan in Kloet & Hincks, 1964 : 80.
42 J. P. DONCASTER
LECTOTYPE here designated: apterous viviparous female. 'S. carnosa'. Surrey, Haslemere, Weycombe. Urtica urens (?). i7.vi.(year ?). (Buckton). (go*).
Paralectotypes : i apterous viviparous female, 2 late-stage larvae. Data as lectotype. (go*).
Related material (urticae'}: i apterous, i alate viviparous females, i nymph,
1 larva. No data. (511)', 2 alate viviparous females. 'Nettle. June. W[ey- combe?].' No other data. (512} ; i larva. 'Urtica urens . No other data. (513) ',
2 apterous, i alate viviparous females, i larva. 'Nettle'. Middlesex, Southgate. 20.vii.i847. (Walker). (W. 1034).
Apterous viviparous female. (Plate 3, fig. 59; Text-figs 17-21.) Colour of macerated specimen : uniformly pale except for very slight darkening around antennal joints and at tibial apices.
18
21
FIGS 17-21. Microlophium carnosum (Buckton). Paratype (906): Fig. 17. Head, upper (left) and lower surfaces. Lectotype: Fig. 18. Right antenna (segments V and VI imperfect). Fig. 19. Apex of rostrum. Fig. 20. Cauda. Fig. 21. Siphunculus.
BUCKTON'S WORKS ON APHIDOIDEA 43
Morphology : body 3-4-3-6 mm long, slightly more than twice as long as broad. Head smooth, antennal tubercles large, prominent, diverging, cephalic hairs long, maximally 65-75^, acute or with spear-shaped apices. Antennal segments I-IV smooth, V slightly imbricated, VI normally so; III with 3 small circular secondary rhinaria near the base; antennal hairs sparse, blunt, up to about 5O(z long. Flagellum a little longer than the body; length ratios of segments III-VI about 123 : 87 : 68 : 22 + 96(?). Rostrum scarcely reaching hind coxae, ultimate segment 0-146-0-150 mm long, equal to or slightly shorter than second segment of hind tarsus, with about 10 non-apical hairs. Femora and tibiae smooth, femoral hairs acute, up to 56^ long; tibial hairs similar, but reaching 61-65^ in length, becoming numerous towards tibial apices. First tarsal segments on all legs with 3 hairs. Tergum smooth, hairs sparse, up to 52[z long on anterior segments, reaching 65^. on eighth tergite, which bears 8-9 hairs. Siphunculi 1-25 mm long, straight, expanded at base, with pronounced apical flange, almost completely smooth, with 2-3 rows of reticulations next to the flange. Cauda 0-42-0-43 mm long, about twice as long as its basal width, slightly constricted at about one-third of its length from the base, with 9-10 hairs. Subgenital plate with 2 long and 5 shorter hairs on anterior half, and 16-18 along posterior margin.
NOTES. Buckton originally regarded carnosa as a variety of the large green nettle aphid known to him as Siphonophora urticae. In fact he distinguishes three varieties of this species, two of which, both green, he describes as Variety a and Variety (3 (1876 : 143) and figures on plate 19. His sheet of drawings of carnosa (plate 20), which show a dark, purplish grey aptera, an alata with brown body and green markings, a pink and green nymph and an almost colourless newborn larva, was at first entitled 'Siphonophora urticae No. 2, var y', but he altered the name to carnosa in the belief that it differed specifically from urticae. On the back of the sheet he appends descriptive notes of the four morphs he drew, and which he describes in greater detail in the text (1876 : 144, 145). The reasons he gives for separating carnosa from urticae are that 'urticae is a larger insect, the antennae are disproportion- ately long, the wings are narrower, the thoracic lobes are more pronounced, and the abdomen is spotted laterally.'
The only extant specimens named carnosa by Buckton are two imperfect adult apterae and two larvae, remounted in balsam by Laing. Examples of the other morphs described and figured, except the newborn larva, are, however, included among specimens named urticae, but there is no indication of which, if any, of these were used as models for carnosa and not for urticae. One of the two alatae on 572 shows some similarities to his original for the figure of the alate carnosa, but there is no proof that one relates to the other.
Evidence of host plant associations is equally inconclusive. In the published text carnosa is recorded from 'the stinging nettle, Urtica urens' and urticae from 'the stinging nettle, Urtica dioica . Though Buckton separates the two nettles by their specific names, he apples the same vernacular name to both. His manuscript notes record the hosts of carnosa as stinging nettle and Rubus fruticosus. The notes for urticae were destroyed when he cut out the individual drawings and remounted them in new positions on a fresh sheet. The only slide to bear a specific host identification is 5x3, which contains a single larva and is marked Urtica urens. One other slide is marked 'nettle' ; that named carnosa bears no data other than the name.
Hille Ris Lambers (1933 : 171), following Lindinger (1932), used the name carnosum Buckton to replace urticae Schrank (preoccupied) for the large green nettle
44 J. P. DONCASTER
aphid. But later (H.R.L., 1947 : 204) he applied to this species the name evansi Theobald and used carnosum to replace sibiricum Mordvilko, 1914, a similar but darker and more sclerotic Microlophium with strongly imbricated siphunculi and a narrower cauda with fewer hairs, recorded from Urtica dioica and U. urens in Siberia but on urens only in the Netherlands. He acted in the belief that Buckton's dark coloured aphid described from Urtica urens was sibiricum, but he did so without having had an opportunity to examine Buckton's specimens.
The nearest approach to authenticated specimens of carnosum are the two adult apterae and two larvae named carnosa by Buckton. Though incomplete, they show most of their more important characters reasonably well, but in none can I detect any morphological difference between them and the remainder of Buckton's material named urticae, all of which agrees with the current concept of Microlophium evansi (Theobald).
The evidence, such as it is, suggests that Buckton's first surmise was correct, namely that carnosum is only the dark reddish or purplish colour-form of the large green nettle aphid which occurs commonly during the summer, often mixed with the typical green form. Buckton's diagnostic characters for carnosa (which seem to refer only to the alata, of which no named specimens exist) contain nothing to conflict with this conclusion. Furthermore, although sibiricum has been recorded from Western Europe, Siberia, Japan and North America, it has not, so far as I can discover, been found in Britain.
I can see no alternative, therefore, but to restore the name carnosum Buckton to the large green nettle aphid in place of evansi Theobald, and I select as lectotype the better preserved of Buckton's two adult apterae on go. The name of the other Microlophium thus reverts to sibiricum Mordvilko, 1914.
Callipterus castaneae Buckton = Myzocallis castanicola Baker
Callipterus castaneae Buckton, 1881 : 26; pi. 91, figs 5-9. [Homonym of Callipterus castaneae
Fitch, 1856 : 471.]
Myzocallis castanicola Baker, 1917 : 424. Myzocallis davidsoni Swain, 1918 : i. Myzocallis assimilis Borner, 1940 : 2.
LECTOTYPE here designated: ovipara. Surrey, Haslemere. Castanea saliva. 1 3. xi. (year ?). (Buckton). (98).
Paralectotypes : 5 alate viviparous females, 5 nymphs, i ovipara. Surrey, Haslemere. iv. (year ?). (Buckton). (95, 96, 97.)
BIOMETRIC DATA. Lectotype, ovipara: body length 1-88 mm, antennae incomplete, ratios of segments Ill-base VI 44 : 29 : 24 : 10 + ?, siphunculus o-io mm long, 0-07 mm wide at flange, cauda not measurable, ultimate rostral segment 0-12 mm, second segment of hind tarsus 0-14 mm, articular diameter of ant. seg. Ill 28(z, longest hair on ant. seg. Ill ±20(1., On vertex o-i5|x, on hind femur ±24^, on hind tibia ±5051, on tergite III ±8o(z, on tergite VIII 140^.
Paralectotype, alata: body length 1-48 mm, ant. flag. 1-32 mm, ratios of ant. segs III-VI 65 : 36 : 25 : 13 + 29, secondary rhinaria on III 6, siph. 0-09 mm long, 0-04 mm wide at apex,
BUCKTON'S WORKS ON APHIDOIDEA 45
cauda not measurable, caudal hairs 9, ult. rost. seg. o-n mm, second seg. hind tarsus o-n mm, artic. diam. ant. seg. Ill 22^, longest hair on ant. seg. Ill i2fz, on vertex 30^1, on hind femur 2O(x, on hind tibia 30(1, on tergite III 35^, on tergite VIII 40^.
Buckton describes the alate viviparous female, nymph and ovipara, and, in addition, the 'apterous viviparous female'. Apterae viviparae do not occur in this species, and what he describes is an ovipara, as is shown by his original drawing for figure 5. This according to the legend represents the apterous viviparous female, but the sketch is marked November 12 and was apparently drawn from the ovipara on 98, dated November 13. Both original and published figure show two ova beside the specimen. Since the evidence linking specimen and figure is so strong I choose this ovipara as lectotype.
Pemphigus cinchonae Buckton : nomen nudum.
Pemphigus cinchonae Buckton, 18890 : 6.
Pemphigus cinchona Buckton; Wilson & Vickery, 1918 : 57.
Pemphigus cinchona Buckton; Patch, 1938 : 226, 348.
In an article on Indian insect pests (Rhynchota), Atkinson records having sent to Buckton for identification a sample of insects, thought to be aphids, infesting leaves of cinchona at Sikkim in August, 1888, and quotes Buckton's reply in which he assigns them tentatively to the genus Cerataphis but reserves his opinion on their specific status pending receipt of more material. Immediately following this record is a brief note of another aphid having been sent to Buckton, who named it provision- ally Pemphigus cinchonae, but again deferred describing it until he could study more material. The note adds no further data.
In the Buckton Collection there is one slide (104*), remounted and relabelled by Laing, containing specimens of an unidentified aleyrodid and bearing data which relate it to the 'Cerataphis sp.' in Atkinson's first record. But neither specimens nor other evidence of identity have come to light which can be related to Pemphigus cinchonae.
Siphonophora circumflexa Buckton — Aulacorthum (Neomyzus)
circumflexum (Buckton) (PI. 3, fig. 60; Text-figs 22-26)
Siphonophora circumflexa Buckton, 1876 : 130; pi. 13, figs 1-4.
Macrosiphum circumflexum (Buckton) Schouteden, 19060 : 238.
Myzus vincae Gillette, 1908 : 19.
Siphonophora callae Henrich, 1910 : 26.
Myzus circumflexes (Buckton) Davis, 19140 : 121.
Neomyzus circumflexus (Buckton) van der Goot, 1915 : vii.
Macrosiphum pelargonii var. circumflexa (Buckton) van der Goot, 1915 : 82.
Aulacorthum circumflexum (Buckton) Timberlake, 1924 : 457.
Amphorophora circumflexa (Buckton) Borner & Schilder, 1932 : 624.
Aulacorthum (Neomyzus) circumflexum (Buckton) Hille Ris Lambers, 1947 : 313, 1949 : 198.
LECTOTYPE here designated : apterous viviparous female. Surrey, Haslemere, Weycombe. Sparaxis sp. ii. (year ?). (Buckton). (io6c*).
46
J. P. DONCASTER
Paralectotypes : 5 apterous viviparous females, 2 larvae, i nymph. Data as lectotype. (705, io6a*, io6b*, io6d*).
Apterous viviparous female. (Plate3, fig. 60; Text-figs 22-26). Colour of macerated specimen: body and appendages pale except for the dorsal dark patches on either side of the median line of the thorax, the characteristic irregularly U-shaped patch on the abdomen, and aslight darkening around the articulations of the antennal segments. Morphology: body i -92-2-66 mm long, oval, slightly less than twice as long as broad. Head coarsely spinulose on upper and under sides. Antennal tubercles rather short, their inner surfaces rounded and slightly protruding and bearing one or two short blunt hairs. Vertex with very few hairs, variable in length, the longest reaching 26(ji, blunt or with slightly expanded apices. Antennae coarsely imbricated throughout, the imbrication being mainly confined to the ventral surfaces of all but the sixth segment. Antennal hairs very sparse, blunt or acute, the longest on III only about one-third of the articular diameter of the segment. Processus terminalis nearly four times as long as the base of VI. The flagellum very slightly longer than the body. Length ratios of segments III-VI about 57 : 44 : 39 : 18 + 72. Rostrum reaching to between second and third coxae, ultimate segment with straight sides and rounded apex and with two non-apical hairs, 0-12 mm long,
0-5
FIGS 22-26. Aulacorthum (Neomyzus) circumflexum (Buckton). Lectotype: Fig. 22. Head, upper (left) and lower surfaces. Fig. 23. Left antenna. Fig. 24. Cauda. Fig. 25. Siphunculus (fractured). Paratype (io6a) : Fig. 26. Apex of rostrum.
BUCKTON'S WORKS ON APHIDOIDEA 47
about twice as long as its basal width and about 1-2 times as long as second segment of hind tarsus. Areas of spinulosity, like that on the head but less dense, occur on and around the coxae, in some specimens spreading on to the trochanters and even the bases of the femora. Legs slender, hairs short and sparse, those on the femora rather stout, blunt, up to 2O[i. long; tibial hairs longer, more numerous near the apex, the longest reaching 34^. Hind tibia about two-thirds of the body length. First tarsal segments with 3, 3, 3 hairs. Tergum of abdomen sclerotic, almost smooth, with sparse, very short blunt hairs about lOjj. long. Eighth tergite with 4 longer blunt hairs, the longest 22(z. Siphunculi 0-41-0-57 mm long, straight, apical two-thirds cylindrical, expanded at base, imbricated over whole length with a few apical striae and distinct flange, slightly less than a quarter of the body length, about equal to antennal segment IV, about 10 times as long as their middle diameter. Cauda 0-20-0-26 mm long, finger shaped, slightly constricted in the middle, with 3 pairs of lateral hairs, about 2-25 times as long as its basal width and half as long as the siphunculi. Subgenital plate with 2 hairs on the anterior half and 8 shorter ones along the posterior margin.
NOTES. Buckton's specimens named circumflexa include, in addition to those listed above, two alate viviparous females and two nymphs of Brachycaudus helichrysi (Kaltenbach) (105}. Sketches of head, antenna, rostrum and siphunculi of alate helichrysi occur on the same sheet with the originals of circumflexum, but are named cinerariae and have not been published. The alate circumflexum described and figured was taken, according to Buckton's notes, in a greenhouse at Chichester in May (the notes are dated May 8). This specimen is missing from the Buckton collection.
A full account of circumflexum is given by Hille Ris Lambers (1949 : 198-201).
Pemphigus coccus Buckton : nomen dubium
Pemphigus coccus Buckton, 18896 : 141.
Pemphigus coccus Buckton; Ghulamullah, 1941 : 225.
Pemphigus coccus Buckton; Takahashi, 1966 : 263.
Buckton's brief and inadequate description of this species is based on immature specimens taken from dried galls on Pistacia vera in Afghanistan in 1885. No specimens so named or identifiable with the known data have come to light, nor any other clue to the identity of the species. Ghulamullah and Takahashi both cite the record of coccus Buckton but make no comment. The name must be regarded as a nomen dubium.
Aphis crithmi Buckton = Dysaphis crithmi (Buckton) (PI. 4, fig. 61 ; Text-figs 27-31)
Aphis crithmi Buckton, 1886 : 323; pi. 4, figs 1-6.
Anuraphis crithmi (Buckton) Theobald, 1927 : 405.
[Brachycaudus helichrysi (Kaltenbach); Hille Ris Lambers, 1934 : 32- Misidentification.]
Aphis (Anuraphis) crithmi Buckton; Balachowsky & Cairaschi, 1941 : 99.
PYezabura crithmi (Buckton) Borner, 1952 : 229.
Dysaphis crithmi (Buckton) Stroyan, 1963 : 47.
LECTOTYPE here designated : apterous viviparous female. Devon, Kingsbridge. Crithmum maritimum. vii. 1886 (?). (Bignell). (132*}.
48 J. P. DONCASTER
Paralectotypes : 7 apterous, 2 alate viviparous females, 4 larvae, 3 nymphs. Data as lectotype. (131*, 132*).
Apterous viviparous female. (Plate 4, fig. 61 ; Text-figs 27-31). Colour of macerated speci- men: head, thorax and sclerotic parts of abdomen pale to mid-brown, remainder of abdomen pale to almost colourless. Antennae mid-brown, becoming darker towards apices. Fore and middle legs mid-brown, hind legs darker. Apex of rostrum, siphunculi and anal plate dark brown. Morphology : body 1-88 mm long, oval, not quite twice as long as broad. Head smooth, frons flat, cephalic hairs stout, spiny, up to about 36(z long. Antennae with segments I and II nearly smooth, III-VI imbricated, antennal hairs stout, blunt, the longest about equal to the articular diameter of III. Processus terminalis about 2-8 times as long as base of VI. Flagellum about two-fifths of the body length. Length ratios of segments III-VI about 24 : 12 : 10 : 7 + 21. Rostrum reaching third coxae, ultimate segment 0-14 mm long, rather narrow, elongate, about 1-25 times as long as hind tarsus II (o-n mm), with 2 non-apical hairs. Femora rather stout, with fairly fine, acute hairs, up to 34^ long. Tibial hairs similar, becoming longer towards apex of tibia, reaching about 50(1. Hind tibia about one-third of body length. First tarsal segments with 3, 3, 2 hairs. The abdomen bears a row of paired spinal scleroites from segments I-V, a broken sclerotic transverse band on VI, and continuous transverse bands on VII and VIII. Pleural and marginal scleroites are less conspicuous than the spinals. Abdominal hairs stout, spiny, short (up to 20^) on anterior segments, becoming longer on posterior segments, reaching 72fz on VIII. Rather small marginal tubercles present on segments I-V. Spinal tubercles absent altogether. Siphunculi short (0-134 mm), slightly shorter than apical rostral segment, about 2-75 times their middle diameter and 1-7 times as long as the cauda, slightly tapered and with a few imbrications. Cauda 0-082 mm long, about as long as its basal width, with 5 hairs.
0-5
FIGS 27-31. Dysaphis crithmi (Buckton). Lectotype: Fig. 27. Head, upper (left) and lower surfaces. Fig. 28. Right antenna. Fig. 29. Apex of rostrum. Fig. 30. Siphunculus. Fig. 31. Cauda.
BUCKTON'S WORKS ON APHIDOIDEA 49
NOTES. There were originally two slides made by Buckton of crithmi, both of them in very poor condition with the balsam emulsified and the specimens obscured. Both bore the data 'Aphis crithmi. Samphire.' and one also included the date (July) and 'Plymouth'. In his published description Buckton records crithmi from Crithmum maritimum at Kingsbridge, Devon, and adds that specimens were sent him by G. C. Bignell, who lived at Plymouth. Apart from the omission of date and locality from one of the labels, Buckton's two slides were so similar in all other respects that I regard the specimens they contained as having all belonged to the same sample.
Theobald (1927 : 405) records having examined one of Buckton's slides of crithmi (that containing two alatae, now 131*), but found the specimens so heavily obscured that he could make little of them beyond concluding that they agreed with his concept of Anuraphis. Laing had marked both slides '? helichrysi Kaltenbach' and noted that they should be remounted, but he never did so. Even after remounting into gum-chloral many of the specimens still suffer from shrivelled or collapsed appendages.
Hille Ris Lambers (1934 : 32) places crithmi Buckton as synonym of Brachycaudus helichrysi (Kaltenbach). At that time, when Buckton's two slides would have been in their original state and the specimens scarcely visible, Laing's tentative identifica- tion of them as helichrysi would have seemed reasonable enough.
Neither the original drawings for Buckton's plate of crithmi nor any manuscript notes relating to it have so far come to light.
Stroyan deals fully with crithmi in his revision of the British species of Dysaphis (Stroyan, 1963 : 47-48).
Aphis cucurbitae Buckton = Aphis gossypii Glover Buckton, 1879 : 56; pi. 54, figs 1,2.
LECTOTYPE here designated: alate viviparous female. Surrey, Carshalton. Cucumis melo. 26. ix. (year ?). (Smee). (139).
Paralectotypes : 5 apterous, 4 alate viviparous females, 7 nymphs, i larva. Data as lectotype. (138, 139).
BIOMETRIC DATA. Lectotype alata: body length 1-66 mm, antennal flagellum 1-02 mm, ratios of segments III-VI 31:31: 23 : 14 + 39, secondary rhinaria on III 8, siphunculus 0-19 mm, cauda 0-12 mm, caudal hairs 5, ultimate rostral segment 0-097 rnm, second segment of hind tarsus 0-083 mm, eighth tergite with 2 hairs, articular diameter of ant. seg. Ill 16(1, longest hair on ant. seg. Ill i2jz, on hind femur 2O(i, on hind tibia 30^, on eighth tergite ±25^.
Paralectotype aptera: body length 1-84 mm, ant. flag. 1-08 mm, ratios segs III-VI 34 : 25 : 22 : 13 + 41, siph. 0-29 mm, cauda 0-14 mm, caudal hairs 4, ult. rost. seg. o-io mm, second seg. hind tarsus 0-09 mm, eighth tergite with 2 hairs, artic. diam. ant. seg. Ill 24^1, longest hair on ant. seg. Ill 14^, on hind femur ±30^, on hind tibia 35^, on eighth tergite ? (not measurable).
Both Buckton's slides have been labelled Aphis gossypii Glover by Laing. Theobald (1927 : 141, 145) published this synonymy and subsequent authors have accepted it.
50 J. P. DONCASTER
Lachnus cupressi Buckton = Cinara cupressi (Buckton) (PI. 4, fig. 62 ; Text-figs 32-35)
Lachnus cupressi Buckton, 1881 : 46; pi. 102, figs 1-3. Lachnus juniperinus Mordvilko, 1894 : 134. Lachniella tujae Del Guercio, 1909 : 309.
[Lachnus juniperi (De Geer) ; van der Goot, 1915 : 396. Misidentification.] Dilachnus cupressi (Buckton) Swain, 1921 : 212. Lachnus sabinae Gillette & Palmer, 1924 : 9. Panimerus cupressi (Buckton) Theobald, 1929 : 148. [Panimerus juniperi (De Geer) Theobald, 1929 : 151 partim.] Panimerus tujae (Del Guercio) Theobald, 1929 : 153.
Cinara cupressi (Buckton) Borner & Schilder, 1932 : 570; Braun, 1938 : 480; Hottes & Essig, I72» Szelegiewicz, 1962 : 83.
FIGS 32-35. Cinara (Cupressobium) cupressi (Buckton). Lectotype: Fig. 32. Head, upper (left) and lower surfaces. Fig. 33. Right antennal segments III-VI. Fig. 34. Siphnnculus. Para type (i4oa) : Fig. 35. Apex of rostrum.
BUCKTON'S WORKS ON APHIDOIDEA 51
Cinara tujae (Del Guercio) Braun, 1938 : 480. Neochmosis cupressi (Buckton) Kloet & Hincks, 1945 : 70. Neochmosis tujae (Del Guercio) Kloet & Hincks, 1945 : 70. Cupressobium cupressi (Buckton) Borner, 1952 : 45. Cinara canadensis Hottes & Bradley, 1953 : 86.
LECTOTYPE here designated: apterous viviparous female. Cornwall, Probus. Cupressus sp. i7.xi.i87Q. (Boscawen). (140*).
Paralectotypes : 3 apterous, I alate viviparous females, i nymph. Data as lectotype. (1400,*, I4ob*, 1400*, I4od*, 1406*).
Apterous viviparous female. (Plate 4, fig. 62 ; Text-figs 32-35) . Colour of macerated specimen : head and body more or less uniformly pale brown; antennae with segments I and II pale brown as head, III paler with very slight darkening at apex, IV and V with basal halves pale, apical halves and whole of VI slightly darker. Apical segments of rostrum dark brown. Coxae and trochanters dark brown ; femora pale on basal half, the rest somewhat darker ; tibiae pale with a small dark brown area at the knee and less pronounced darkening at the apices; tarsi brown. Siphuncular cones, cauda, anal and subgenital plates, stigmal plates and muscle-plates brown. Morphology: body 2-70-2-92 mm long, broadly oval, about 1-7 times as long as broad. Head more or less semicircular in outline, clothed with numerous long fine hairs, the longest (frontal) reaching about 145^. Antennae of 6 segments, about one-third as long as the body, with rather numerous long fine hairs ranging maximally from 177-197^ on segment III. II with 9 or 10 hairs, VI with 4-6 hairs confined to the basal third of the segment, processus terminalis with 3 subapical setae; length ratios of segments II I-VI about 36 : 15 : 17 : 14 + 4, secondary rhinari a confined to IV, with i or 2, and V, with i. Rostrum reaching a little beyond third coxae, ultimate segments 0-147 an(^ 0-085 mrn long respectively, slender and tapering, fourth segment with 3 or 4 non-apical hairs. Legs short and stout with numerous long fine hairs reaching about 200^1 on hind tibia, which is about two-fifths of the body length. First tarsal segment with dorsal side much shorter than the basal diameter (23 : 42(0.) . Second tarsal segment 0-24-0-26 mm long, slightly longer than rostral segments 4 and 5 together. Abdominal dorsum membranous with conspicuous muscle-plates, small sclerotic stigmal plates, a transverse row of 4 scleroites on tergite VII and two irregular transverse sclerotic bands on VIII. Siphuncular cones from 0-25- 0-30 mm in diameter, shallow (quite unlike Buckton's exaggerated figure), with numerous hairs in 4-5 whorls. Cauda very broadly triangular with more or less rounded sides, about one-third as long as its basal width.
NOTES. In this instance Buckton's published figures are an improvement on his originals, which are carelessly drawn and crudely coloured. His notes add nothing to the published record.
Hyalopterus dilineatus Buckton = Longicaudus trirhodus (Walker) Buckton, 1879 : 113; pi. 76, figs 1-7.
LECTOTYPE here designated: alate viviparous female. Northumberland, Alnwick. Rosa centifolia var. muscosa ('Moss Rose'), v. (year ?). (Hardy). (147).
Paralectotypes: i larva, 4 nymphs. Data as lectotype. (146).
BIOMETRIC DATA. Lectotype, alata: body length 1-88 mm, antennal flagellum 1-38 mm, ratios of segments III-VI 95:21:21:16 + 20, secondary rhinaria on III 84, siphunculus o-io mm, cauda 0-22 mm, caudal hairs 14, ultimate rostral segment 0-09 mm, second segment of hind tarsus 0-14 mm, eighth tergite with 4 hairs, articular diameter of ant. seg. Ill 32(x, longest hair on ant. seg. Ill 16^, on hind femur 30^, on hind tibia 40^, on eighth tergite ±20^..
52 J. P. DONCASTER
Buckton's specimens named dilineatus are a mixture of Longicaudus trirhodus (Walker) (see above; 146, 147), Macrosiphum rosae (L.) (146) and Myzaphis bucktoni Jacob (3 apt. viv. females on 145*, 1450,*, I45b*). The slide data on 146 and 147 indicate that the trirhodus and rosae are the material sent by Hardy from Alnwick, referred to under Buckton's description of the nymph (1879 : 113). The three apterae of bucktoni, therefore, would seem to belong to the material recorded from Haslemere and Wanstead on Rosa centifolia in July (the slide labels include no locality or date). Wanstead suggests Walker, but none of Walker's extant slides contains bucktoni. A Walker slide (W. 841} in Buckton's collection contains rosarum Kaltenbach which Buckton may have confused with bucktoni.
On his plate of dilineatus Buckton figures the young larva, nymph and alata of trirhodus, and the adult aptera of bucktoni. His original drawing of the adult aptera is a good representation of bucktoni, better than the coloured lithograph, and shows (what the latter does not) that Buckton was uncertain of the true length of the siphunculi, leaving their extremities unfinished and indicated only vaguely by dotted lines. From his manuscript notes, which accurately describe his drawing, it seems that at first he regarded bucktoni as the adult aptera of dilineatus, but in his published account he modifies this view and describes the larva of trirhodus as the adult aptera (I regard the larva on 146 as his probable model), adding a brief description of 'variety a' which, as Jacob (1946 : no) points out, refers to bucktoni (but he omitted to alter the caption to figure 3 accordingly). The descriptions of the nymph, and the alata bred from one of the nymphs sent by Hardy, certainly refer to trirhodus.
Buckton also describes and figures (figs 5, 6, 7) the oviparous female of dilineatus, but there is no specimen of an ovipara so named and the original drawings for these figures are missing. Figures 6 and 7, showing an enlarged antenna and abdominal appendages, suggest Myzaphis rosarum (Kaltenbach), but there is no proof that this is so, and the identity of the ovipara must remain uncertain.
Since three of the four morphs described and figured are all supported by specimens of Longicaudus trirhodus, I place dilineatus Buckton, as other authors have done, as a synonym of trirhodus Walker.
Theobald (1927 : 38) places dilineatus in Longicaudus and redescribes it, believing it to be distinct from trirhodus. However, he quotes Laing as being doubtful whether Buckton's alate female is distinct from trirhodus. Hille Ris Lambers (1934 : 26) considers Theobald's material named dilineatus to be trirhodus Walker. It is unlikely that he saw Buckton's material. Jacob (1946) gives a full account of Myzaphis bucktoni, which he identifies with the description and figure of Buckton's 'variety a' of dilineatus, but without having seen Buckton's specimens. Buckton's three adult apterae of bucktoni agree in all respects with Jacob's diagnosis.
Aphis edentula Buckton = Rhopalosiphum insertum (Walker)
Aphis inserta Walker, 18496: app. xxxix. Lectotype, alate viviparous female, ENGLAND:
Essex (BMNH). Aphis edentula Buckton, 1879 : 39; pi. 48, figs 1-3. Syn. n.
(For full synonymy, see Doncaster, 1961 : 86.)
BUCKTON'S WORKS ON APHIDOIDEA 53
LECTOTYPE here designated : ovipara. Essex, Wanstead. Crataegus monogyna. 7.xi.(year ?). (Walker). (130).
BIOMETRIC DATA. Lectotype, ovipara: body length 1-52 mm, antennal flagellum 0-51 mm, ratios of segments III-V 23 : 10 : 8 -f 22, siphunculus 0-12 mm, cauda not measurable, caudal hairs 5 (?), ultimate rostral segment o-io mm, second segment of hind tarsus o-io mm, eighth tergite with 4 (?) hairs, articular diameter of ant. seg. Ill i6(x, longest hair on ant. ?eg. Ill lo^i, on hind femur I2fi, on hind tibia 25(z, on eighth tergite ±25^.
Buckton describes the apterous viviparous female, the nymph, the alate viviparous female and the ovipara, but he figures only the last three morphs. The specimens, he says, were sent to him by Walker, who collected them from Crataegus at Wanstead in November. There are no slides in Buckton's collection labelled edentula and, from the data available, I can trace no specimens which might have been his models for the nymph and alata. But on slide 130, labelled 'A. crataegi Walk : ovip. female. Nov.' in Buckton's hand, are three oviparae of Rhopalosiphum insertum (Walker), one of which corresponds reasonably well with the original for figure 3 and his measure- ments and description of the oviparous edentula. Since the data on the slide corres- pond with the published data and also with a manuscript note on the sheet of original drawings 'ovip. female on Whitethorn Nov. 7. Walk.', I believe this specimen to be Buckton's type of the ovipara of edentula, which I therefore place as a synonym of insertum Walker.
Theobald (1927 : 213) quotes Buckton's account of edentula in full, adding only that the species is not represented in Buckton's collection.
Borner (1952 : 70) correctly puts edentula as synonym of oxyacanthae Schrank = insertum Walker.
Schizoneura fodiens Buckton = Schizoneura ulmi (L.) Buckton, 1 88 1 : 94; pi. 106, figs 6-12.
LECTOTYPE here designated: apterous viviparous female. Surrey, Haslemere. Ribes nigrum. 15. x. (year ?). (Buckton). (183*).
Paralectotypes : 3 alate viviparous females, 12 nymphs, data as lectotype. (181*, 182, i83a*, 184}.
BIOMETRIC DATA. Lectotype, aptera: body length 1-54 mm, whole antenna 0-32 mm, ratios of segments I-V 13 : n : 28 : 13 : 37, siphuncular diameter 0-07 mm, ultimate rostral segment 0-13 mm, second segment of hind tarsus 0-08 mm, articular diameter of ant. seg. Ill 33^, longest hair on ant. seg. Ill 45^, on hind femur 40^, on hind tibia 50^.
Paratype, alata: body length 1-72 mm, antennal flagellum 0-63 mm, ratios segs III-VI 50 : 12 : 8 : 5 + 3, secondary rhinaria on seg. Ill 18, on IV 2, on V o, on VI o, siph. diam. 0-07 mm, ult. rost. seg. 0-13 mm, second seg. hind tarsus o-io mm, artic. diam. ant. seg. Ill i6[z, longest hair on ant. seg. Ill 42^, on hind femur 44^, on hind tibia 50^.
Buckton describes and figures the apterous viviparous female, nymph and alata, and adds figures of a newborn larva and details of wings and alate antenna. All his specimens named fodiens are Schizoneura ulmi (L.), and I have selected as lectotype the single aptera on 183, the slide data of which correspond exactly with those of his original drawing of that morph (fig. 6), and which was doubtless the specimen
54 J- P. DONCASTER
described and figured. Buckton's published description is incorrect in giving the antennal length as 'three quarters the length of the body' ; his MS notes give the ratio as one-third, and his published measurements (antenna 0-38 mm : body 1-39 mm) are about right when matched with the mounted specimen. The specimen is rather small for an apterous exule of ulmi, and the tarsi are less spinulose than those of other specimens I have examined, but other characters, particularly the wax- gland rosettes with conspicuous central ring, agree well.
Endeis formicina Buckton = Baizongia pistaciae (L.)
Buckton, 1883 : 91; pi. 129, figs i and 3. (For synonymy, see aedificator, p. 31.)
LECTOTYPE here designated: apterous viviparous female. Northumberland, Cheviot. Poa pratensis (? or Carex dioica) roots, v. (year ?). (Hardy}. (189*).
BIOMETRIC DATA. Lectotype, aptera: body length 1-46 mm, whole antenna 0-27 mm, ratios of segments I-V 17:16:16:13:31, ultimate rostral segment 0-13 mm, second segment of hind tarsus o-io mm, primary hairs on subanal plate 8, longest hair on subanal plate eighth tergite with 6 hairs, the longest
Buckton's original slide labelled Endeis formicina (mis-spelled 'formacina') contained 14 specimens belonging to four species: Forda formicaria Heyden, Smynthurodes betae Westwood, Geoica eragrostidis (Passerini) and Baizongia pistaciae (L.). I have remounted these specimens on separate slides (189*, i8ga*-e*) and compared them with Buckton's description and figures of formicina. The specimen that agrees most closely, particularly with the characters shown in the original drawings of whole insect and antenna (used for figs i and 3 on the plate) is the aptera of Baizongia pistaciae (L.). Further confirmation that this was the specimen figured is given by Buckton's MS notes in which the host plant is recorded as Poa pratensis, and the code letters y, which appear on the slide, are also written beside the drawing. The colour characters given in the published description (1883 : 91) agree well with those of the original drawing, but the host plant is published as Carex dioica. The specimens were sent by Hardy, whose name also appears on the original drawing. I therefore choose this specimen as lectotype of formicina Buckton.
Theobald, who recorded (1929 : 193) having seen Buckton's slide, and who also realised that it contained four species, placed formicina as a doubtful synonym of Geoica squamosa Theobald nee Hart ( = eragrostidis Passerini). Borner (1952 : 197) concluded from Buckton's figures that formicina should be assigned to Pemphigus. Stroyan (in Kloet & Hincks, 1964 : 86) correctly placed formicina as synonym of pistaciae (L.).
Lachnus formicophilus Buckton : nomen dubium
Lachnus formicophilus Buckton, 1901 : 257. Lachnus formicophilus Buckton; Donisthorpe, 1902 : 39. Lachnus formicophilus Buckton; Schouteden, 19060 : 201. Lachnus formicophilus Buckton; Donisthorpe, 1927 : 167, partim.
Buckton describes and figures a specimen sent him by Donisthorpe, who took it
BUCKTON'S WORKS ON APHIDOIDEA 55
from a nest of Formica rufa at Oxshott, Surrey. Buckton gives no date, but Donisthorpe (1902) refers to this specimen and gives the date as 1900. Donisthorpe (1927) records collecting formicophilus from nests of F. rufa on 24 April, 1900, at Oxshott, and again on 6 September, 1912, at Weybridge. The Oxshott record doubtless refers to the specimen, now lost, which Buckton described; that from Weybridge is described by Theobald (1929 : 351) as Lachnus (?) formicophilus Buckton and is now in the BMNH collection.
Buckton's description is so vague that it is impossible to determine what he had before him. His roughly-sketched figure shows an aphid with long antennae and legs, rather narrow wings with normal aphidine venation, and an abdomen either shrivelled almost to nothing or absent altogether. The body is said to be small, globular, black, and covered with white flocculent matter, and the expanse of the wings is given as n-o mm. Buckton identifies it as a male.
The unusually large wing span and absence of visible siphunculi and cauda may have led Buckton to place this specimen in Lachnus, but the long antennae (about three-quarters of the length of the wings) must preclude this. The sketch certainly suggests a large aphid, possibly a Callipterine, e.g. Euceraphis punctipennis (Zetterstedt), or, if the wing span were not so large, Phyllaphis fagi (L.), which would accord with the flocculence. But if the specimen were indeed that taken by Donisthorpe on 24 April, the possibility of its being a male must be remote. I find myself in agreement with Schouteden (1906) when he writes: '. . . il me semble un peu exage"re de declarer myrmecophile un Aphide, parce qu'un unique exemplaire, et surtout une forme aile'e! s'en est rencontre" dans un nid de fourmis', and I regard formicophilus Buckton as a nomen dubium.
Donisthorpe's second specimen is an apterous viviparous female of Lachnus (Schizodryobius) longirostris (Borner), or possibly exsiccator (Altum) (= pallipes Hartig), as Borner (1952 : 46) supposed, basing his conclusion, presumably, on Theobald's description and figure.
Lachnus fuliginosus Buckton = Tuberolachnus salignus (Gmelin)
Aphis saligna Gmelin, 1790 : 2209.
Lachnus punctatus Burmeister, 1835 : 93.
Aphis viminalis Boyer de Fonscolombe, 1841 : 184.
{Lachnus longipes Dufour; Buckton, 1881 : 59. Misidentification.]
Lachnus fuliginosus Buckton, 18916 : 40.
Tuberolachnus viminalis (Boyer de Fonscolombe) Mordvilko, 1909 : 374, Das, 1918 : 257.
Lachnus viminalis (Boyer de Fonscolombe); van der Goot in Das, 1918 : 142.
Pterochlorus viminalis (Boyer de Fonscolombe); Swain, 1921 : 211.
Pterochlorus salignus (Gmelin) Theobald, 1929 : 104.
Tuberolachnus salignus (Gmelin) Borner, 1952 : 45.
LECTOTYPE here designated: alate viviparous female. PAKISTAN, Quetta. 1890 ? (Elliot ?). (J97*).
Paralectotypes : 2 apterous viviparous females, i nymph. Data as lectotype. , 198*).
56 J. P. DONCASTER
BIOMETRIC DATA. Lectotype, alata: body length 4-60 mm, antennal flagellum 1-40 mm, ratios of segments III-VI 35 : n : 13 : 12, secondary rhinaria on III 9, on IV 2, on V o, siphuncular diameter 0-12 mm, ultimate rostral segment 0-19 mm, second segment of hind tarsus 0-36 mm, articular diameter of ant. seg. Ill 36(0., longest hair on ant. seg. Ill yojx, on hind femur 8o(ji, on hind tibia 6o(ji.
Paralectotype, aptera: body length 2-84 mm, ant. flag. 0-82 mm, ratios segs III-VI 46 : 20 : 24 : 28, siph. diam. o-n mm, ult. rost. seg. 0-19 mm, second seg. hind tarsus 0-31 mm, artic. diam. ant. seg. Ill 40^, longest hair on ant. seg. Ill 40^, on hind femur 40^, on hind tibia 55^1.
A sample of aphids said to have been taken on apricot, almond and peach trees at Quetta in 1890 was sent by the Indian Museum, Calcutta, to Buckton for identi- fication. Believing he had a new species, he named it Lachnus fuliginosus and described and figured the larva, nymph and alata. There are in the BMNH 15 specimens from the sample sent from Quetta, originally mounted by Buckton on two slides. They include the three morphs Buckton described, as well as several adult apterae, and are a mixture of two species, Tuberolachnus salignus (Gmelin) and Pterochloroides persicae (Cholodkovsky) . The descriptions of nymph and alata, and the three morphs figured (alata, nymph and aptera or larva) all agree with the characters of salignus : only the description of the larva agrees with persicae. These descriptions correspond closely with five specimens originally on one of Buckton's slides (2 apterae, a nymph and an alata of salignus and an aptera of persicae}, named fuliginosus, which were remounted by Laing in 1916 on three separate slides. I regard these specimens as the type-material on which Buckton based fuliginosus. The second of Buckton's original slides (199) contains five apterae and five larvae of persicae only, and is labelled simply 'Lachnus n.s. . . . Quetta', without specific name. For this reason I exclude these specimens from the type-series, although they appear to have formed part of the original sample.
Das (1918 : 258-9, 266-7) records having examined in the Indian Museum part of the same material which had been sent to Buckton, and he found that it contained a mixture of the same two species. He suggests that there may have been some accidental mixing of samples which could account for the inclusion of the Salix- feeding salignus among the persicae taken from Prunus. He realised that most of Buckton's descriptions and all his figures relate to salignus, and he states, moreover, that in the course of his work on elucidating fuliginosus, he not only compared the Calcutta material with Buckton's published account, but also sent for 'the insect from Quetta', from which I infer that he may have had an opportunity to examine at least one of the specimens originally sent to Buckton. This supposition is strengthened by Theobald (1929 : 108) who writes 'Mr Laing has examined the type of Buckton's fuliginosus and finds it is undoubtedly this species [i.e. salignus]. Buckton's slides contain viminalis [= salignus] and persicae, so fuliginosus is really a composite species, but Das selected the type and sank it as a synonym of viminalis.'
Although Theobald's statement implies the existence of a type of fuliginosus, I can find no evidence that a type designation was ever published, or even contem- plated. None of the specimens now in the BMNH bears any type-label or equivalent indication, and although Dr A. P. Kapur, at my request, has kindly searched the
BUCKTON'S WORKS ON APHIDOIDEA 57
collections in the Zoological Survey of India, which hold type-material of some of Buckton's Indian aphid species, fuliginosus seems not to be among them. If in fact one of Buckton's specimens was sent to Das for examination, it would most probably have been the alate female of salignus, mounted by Laing singly on slide J97, rather than the nymph on 198, or the two apterae of salignus and one of persicae on 196. In the belief that this alata is the specimen most likely to have been the subject of Theobald's statement quoted above, I designate it here as lectotype of Lachnus fuliginosus Buckton.
Schizoneura fuliginosa Buckton — Schizolachnus pineti (Fabricius)
Aphis pineti Fabricius, 1781 : 389.
Aphis tomentosa Villers, 1789 : 549.
Schizoneura fulginosa Buckton, 1881 : 96; pi. 107, figs 1-6.
Glyphina pilosa Buckton, 1883 : 16.
Schizolachnus tomentosus (Villers) Mordvilko, 1909 : 375, Swain, 1921 : 212, Theobald, 1929 : 161.
Schizolachnus pineti (F.) Borner, 1952 : 40
LECTOTYPE here designated: alate viviparous female. Surrey, Haslemere, Weycombe. Pinus nigra var. austriaca. Undated. (Buckton). (194).
Paralectotypes : i apterous viviparous female, 2 larvae ('29.x.'), 2 larvae, undated. Data as lectotype. (193, 195}-
BIOMETRIC DATA. Lectotype, alata : body length 2-38 mm, antennal flagellum 0-86 mm, ratios of segments III-VI 49 : 20 : 20 : 18, secondary rhinaria on III 8, siphuncular diameter 0-07 mm, ultimate rostral segment 0-14 mm, second segment of hind tarsus 0-26 mm, articular dia- meter of ant. seg. Ill 24^1, longest hair on ant. seg. Ill 140^, on hind femur 200^, on hind tibia
200[Z.
Paralectotype, aptera: body length 2-28 mm, ant. flag. 0-64 mm, ratios segs III-VI 34 : 16 : 14 : 17, siph. diam. 0-04 mm, ult. rost. seg. 0-14 mm, second seg. hind tarsus 0-27 mm, artic. diam. ant. seg. Ill 32^, longest hair on ant. seg. Ill i2O(ji, on hind femur 140^, on hind tibia
Buckton describes and figures the adult aptera and alata, and also the nymph, a morph which is not included among his specimens. Figure 3, said to be an apterous male, appears to have been drawn from a young larva on 195. His descriptions and figures agree tolerably well with the corresponding specimens.
Siphonophora rosae var. glauca Buckton = Macrosiphum rosae (L.) Buckton, 1876 : 109; pi. 3, figs 1-3, 5-7.
There are no slides named glauca in the Buckton Collection and no indication in the published text of the characters Buckton uses to distinguish glauca from rosae L. His original drawings carry no manuscript notes, but resemble rosae so closely in appearance that I agree with Borner (1952 : 293) in regarding glauca as a synonym of rosae L.
58 J. P. DONCASTER
Myzus gracilis Buckton = Metopolophium dirhodum (Walker)
Buckton, 1876 : 176; pi. 34, figs 4, 5.
(For synonymy, see Doncaster, 1961 : 58.)
LECTOTYPE here designated: alate viviparous female. Surrey, Shottermill. Acer pseudoplatanus. xi. (year ?). (Buckton). (210*).
Paralectotype: alate male. Data as lectotype. (2100.*}.
BIOMETRIC DATA. Lectotype, alata: body length 2-80 mm, antennal flagellum not measur- able (both incomplete), ratios of segments Ill-base VI 73 : 48 : 46 : 22 + ?, secondary rhinaria on III 21, siphunculus 0-41 mm, cauda 0-23 mm, caudal hairs 9, ultimate rostral segment o-n mm, second segment of hind tarsus 0-17 mm, eighth tergite with 4 hairs, articular diameter of ant. seg. Ill 32(jL, longest hair on ant. seg. Ill i2(z, on hind femur 24^, on hind tibia 40^, on eighth tergite 55^.
Buckton describes and figures the alate viviparous female and alate male, which he records as having been taken on sycamore 'in company with Chaitophorus aceris' in November. (In the legend to the plate the male, fig. 4, is mistakenly ascribed to ribis.) Buckton's original slide, named Myzus gracilis, contained the alate female and alate male, which correspond with his text and figures and which I regard as his types, together with four larvae of Periphyllus acericola (Walker). All have been remounted.
Rhizobius graminis Buckton = Aploneura lentisci (Passerini)
Tetraneura lentisci Passerini, 1856 : 264.
Aploneura lentisci (Passerini) Passerini, 1863 : 201.
[Tychea eragrostidis (Passerini ?) Buckton, 1883 : 89, partim. Misidentification.]
Rhizobius poae Buckton, 1883 : 93, nee Thomas, C. A., 1879 : 166.
Rhizobius graminis Buckton, 1883: note below legends to pi. 129.
Tycheoides eragrostidis Schouteden, 19060 : 194.
Neorhizobius poae del Guercio, 1917 : 247.
LECTOTYPE here designated: apterous viviparous female. Northumberland, Cheviot Hills. Poa annua roots. Undated. (Hardy ?). (358}.
Paralectotypes : 7 apterous viviparous females. Data as lectotype. (358).
BIOMETRIC DATA. Lectotype, aptera: body length 1-56 mm, whole antenna 0-25 mm, ratios of segments I-V 14 : 15 : 10 : 10 : 29, ultimate rostral segment 0-085 mm, second segment of hind tarsus 0-080 mm, longest hair on antenna i2(i, on hind femur 14(1, on hind tibia 14^, on eighth tergite 24(1, on cauda 24^.
Buckton first describes this species as Rhizobius poae in the fourth volume of his monograph, but after the text had been printed he discovered that the name had already been published by Cyrus Thomas for an American species in 1879. He therefore concedes priority to poae Thomas and substitutes graminis for his own species in a note added below the figure legends for plate 129.
Buckton's slide (358, still labelled 'poae} contains eight apterous females of Aploneura lentisci (Passerini) and four apterae of a Pemphigus species, all reasonably well preserved. He figures both these species, lentisci in figs 9 and n, and the
BUCKTON'S WORKS ON APHIDOIDEA 59
Pemphigus in 10 and 12. The original sketch for fig. 9 is a careful and accurate drawing of the adult aptera of lentisci, and his published description corresponds closely with it. The sketch of the Pemphigus is rough and shows the ventral aspect only. I therefore place graminis Buckton as a synonym of lentisci Passerini. Buckton's sketch for figure 9 can be matched fairly closely with one of the specimens of lentisci on 358, and this I regard as Buckton's type of graminis.
The mixture of species on Buckton's slide and in his figures of graminis led Theobald (1929 : 213, 262, 265) to synonymize graminis with auriculae Murray (Pemphigus) and also, doubtfully, with lentisci. Records by Theobald (1915 : 151), Willcocks (1922 : 58, 1925 : 122) and Hall (1926 : 47) of graminis Buckton on roots of Gramineae in Egypt all refer to lentisci.
Pemphigus immunis Buckton
[Pemphigus bursarius (L.); Passerini, 1863 : 198, Courchet, 1879 : 49, 93, pi. 5, fig. 4, Buckton, 1881 : pi. 113, figs 6-8, Kessler, 1882: pi. i, figs 2-5, Lichtenstein, 1885: pi. 3, figs i, 2, 1886 : 26, del Guercio, 1900 : 98. Misidentifications.]
Pemphigus immunis Buckton, 1896 : 51.
Pemphigus lichtensteini Tullgren, 1909 : 148.
Pemphigus globulosus Theobald, 1915 : 147.
[Pemphigus napaeus Buckton; Tseng & Tao, 1936 : 168, Takahashi, 1938 : 14. Misidentifica- tions.]
Lectotype (designated by Doncaster, 1969 : 159) : alate viviparous female (fundatrigenia migrans). INDIA, Kashmir, Gilgit Road, Bunji. c. 1400 m. Populus leuphratica. 2.vii.i895. (Alcock). (BMNH 2x7*).
Paralectotypes : i alate viviparous female (fragmentary), 4 nymphs. Data as lectotype. (BMNH 2x7*, 279*). i alate viviparous female, 5 nymphs, 2 larvae (some fragmentary). Data as lectotype. (Zoological Survey of India, Calcutta, 7262/H7*, 7263/H7*).
I have already dealt elsewhere (Doncaster, 1969) with Buckton's Indian material of immunis, including descriptions of lectotype and paralectotypes.
It is interesting to note that when Buckton described immunis as new in 1896 he already had material of this species in his collection and had included in his mono- graph figures of the fundatrix, the antenna of the alate migrant, and the gall (Buckton, 1881 : pi. 113, figs 6-8). The specimens had been sent to him from Montpellier by Lichtenstein who believed them to be bursarius, and some of them that Buckton mounted, including those he drew, are still extant, though not the gall. The aphids comprise a fundatrix, whole specimens and fragments of eight or nine nymphs, and an alate migrant (j8a*, ?8b*, 79). In Buckton's published description of bursarius (1881 : 117), and his figures of it on plate in, based on British material, he is correct in his identifications of specimens and galls, but includes his figures of immunis in a subsequent plate (113) and repeats Lichtenstein's error in ascribing them to bursarius. Buckton's original sketch of the gall of immunis gives a better impression of its colour, texture and position on the twig than the published lithograph.
60 J. P. DONCASTER
Though widely distributed throughout the palaearctic and temperate oriental regions, immunis has not hitherto been recorded in Britain.
Aphis instabilis Buckton = Brachycaudus cardui (L.)
Aphis cardui L., 1758 : 452. (Linnaeus left no aphid types.) Aphis instabilis Buckton, 1879 : 94; pi. 68, figs 1-5. Syn. n.
LECTOTYPE here designated: alate viviparous female. Surrey, Haslemere, Weycombe. Matricaria inodora. i8.vi.(year ?). (Buckton). (220).
Paralectotypes : 3 alate viviparous females, 9 nymphs, i larva. Data as lectotype. (220).
BIOMETRIC DATA. Lectotype, alata: Body length 1-56 mm, antennal flagellum 1-46 mm, ratios of segments III-VI 58 : 38 : 28 : 14 + 46, secondary rhinaria on III 30, siphunculus 0-30 mm, cauda 0-12 mm, caudal hairs 6, ultimate rostral segment 0-19 mm, second segment of hind tarsus 0-14 mm, articular diameter of ant. seg. Ill 26(i, longest hair on ant. seg. Ill i2[z, on hind femur 22 [i, on hind tibia 3O[x, on eighth tergite ±50^.
Buckton describes the apterous viviparous female, two varieties of the nymph, and the alate female of instabilis, which he collected from Pyrethrum inodorum (now Matricaria inodora) and also Epilobium montanum and E. parviflorum. His manu- script notes suggest that material from both Matricaria and Epilobium was collected at Weycombe, but in his published account he mentions having received material from Epilobium from Barrett in Pembroke. Buckton's figures, and especially his original sketches, suggest that instabilis is based on at least two species, but his data are confusing. There are no specimens named instabilis in Buckton's collection. There is, however, a slide (220), unnamed but labelled 'Pyrethrum. June.' in Buckton's hand, which Theobald examined and believed to contain instabilis. It contains alatae, nymphs and a larva of Brachycaudus cardui (L.), correctly identified by Laing who, however, in a note on the slide envelope, doubted whether they were in fact instabilis. Nevertheless, Buckton's drawing of the nymph in figure 3 and that of the larger and darker of the two alatae (figure 4) seem to relate to his MS notes on specimens from Matricaria, dated June 18, in which case the alatae and nymphs of cardui on 220 could supply the models, as Theobald surmised. But the alata in figure 5, said to be newly emerged, is clearly something different. Apart from its pale green colour, its cauda is too big for cardui and, in the original, though not the reproduced figure, marginal tubercles are clearly indicated on the seventh abdominal segment, characters which suggest that it may have been either a vagrant, or taken from among material from Epilobium.
Buckton's drawings of the aptera (perhaps drawn from a larva — the measurements he gives are small for an adult) and 'var. i' of the nymph (figures i and 2) certainly do not look like cardui and could well be of an Epilobium-ieeding aphid. He describes the siphunculi of both morphs as pale, and notes the presence in the aptera of two small tubercles on the antepenultimate abdominal segment: characters which also could be appropriate to such species. But although specimens of some Epilobium- feeders occur in Buckton's collection, they include none that I can relate with
BUCKTON'S WORKS ON APHIDOIDEA 61
any confidence to instabilis. There is one slide ( 163) labelled 'Epilobium. Pembroke. ', containing apterae and alatae of grossulariae Kaltenbach, which I believe to be mater- ial sent by Barrett, but I regard these as the types oipenicillata Buckton for reasons given below (p. 76). Since the only specimens that can clearly be related to instabilis are those from Matricaria, I place instabilis as a synonym of cardui L.
Theobald quotes Buckton's description of instabilis twice, once under Aphis (1927 : 214) and again under Anut -aphis (1927 : 289), but does not recognize it as cardui. Borner (1952 : 231) thinks it likely to be an Epilobium-feeding species but does not identify it.
Rhizobius jujubae Buckton
Buckton, 1883 : 181, 18996 : 277.
Laing (1923 : 247) identifies jujubae Buckton as '. . . a very young and immature species belonging to the Monophlebinae' (Margarodidae) . The single slide of jujubae, labelled simply 'Zizyphusjujuba, India' in Buckton's hand, is in the Coccoidea collections of the BMNH.
Aphis lentiginis Buckton = Dysaphis (Pomaphis) plantaginea (Passerini)
Aphis pyri Hartig, 1841 : 369, nee Boyer de Fonscolombe, 1841 : 189.
[Aphis malifolii sensu auctt. nee Fitch, 1855 : 49. Misidentifications.]
Myzus plantagineus Passerini, 1860 : 31, 35. [No type exists2.]
Myzus mali Ferrari, 18726 : 221.
[Aphis mali F. ; Buckton, 1879 : 45, partim. Misidentification.]
Aphis lentiginis Buckton, 1879 : 59, pi. 55, fig. i. Syn. n.
[Dentatus sorbi (Kaltenbach) van der Goot, 1915 : 177. Misidentification.]
Anuraphis roseus Baker, 1920 : 5.
Dentatus plumbicolor Nevsky, 1929 : 287.
Sappaphis plantaginea (Passerini) Hille Ris Lambers, 1945 : 58, 1948 : 287, Stroyan, 19570 :
passim.
Sappaphis mali (Ferrari) Borner, 1952 : 98. Dysaphis (Pomaphis) plantaginea (Passerini) Stroyan, 1963: passim.
LECTOTYPE here designated: apterous viviparous female (fundatrigenia). Sussex, Horsham, Cowfold. Pyrus communis. 8.vi.(year ?). (Borrer). (373d*}.
Paralectotypes : 3 apterous viviparous females. Data as lectotype. (J7J«*, &*, c*).
BIOMETRIC DATA. Lectotype, aptera: body length 2-42 mm, antennal flagellum 1-89 mm, ratios of segments III-VI 58 : 42 : 27 : 13 -f- 49, siphunculus 0-37 mm, cauda 0-14 mm, caudal hairs 6, ultimate rostral segment 0-15 mm, second segment of hind tarsus 0-14 mm, eighth tergite with 5 hairs, articular diameter of ant. seg. Ill 34^, longest hair on ant. seg. Ill I$\L, on hind femur 34^, on hind tibia 46(1., on eighth tergite 76(x.
Buckton describes and figures the apterous and a late viviparous females of lentiginis, said to have been taken on pear in early June. His manuscript notes on
2See Stroyan, 1957^ : 25.
62 J. P. DONCASTER
the original sketches are dated 8 June. There are no specimens named lentiginis among Buckton's slides, but a slide (373) labelled 'A. Pyraria. Cowfold. On Pear.' contained apterous and alate specimens (now remounted) which agree closely with his description and figures and which I believe to be his types of lentiginis. They are four adult fundatrigeniae of a Dysaphis species, either identical with or closely related to plantaginea (Passerini), and three alatae of Rhopalosiphum insertum (Walker). The four apterae are well preserved and were identified on the original slide by both Laing and Theobald as Anur aphis roseus Baker (= plantaginea Passerini), though neither connected them with lentiginis. Stroyan's (1957) keys for the identification of Dysaphis species bring one to the same conclusion, but doubts arise if one accepts Buckton's record that the aphids were taken on pear, since plantaginea is confined to apple as primary host.
If pear was indeed the host there might be grounds for regarding lentiginis as a 'good' species, since I know of no Dysaphis species resembling lentiginis that has been recorded on pear in England. It would be tempting to put forward the possibility that lentiginis may be the primary host form of gallica Hille Ris Lambers, a Dysaphis species closely similar in micromorphology to plantaginea, the primary host of which is not at present known. But this would seem unlikely since Hille Ris Lambers (X955 : 3°9) failed in attempts to establish autumn migrants of gallica on pear.
Buckton names his species lentiginis on account of two coloured areas on the dorsum surrounding the bases of the siphunculi. He mentions them twice, describing them first as 'conspicuous orange-yellow spots' and later as 'rusty blotches'. In his original sketch of the aptera they are shown as dull coppery red, but in the reproduced figure they are coloured bright yellow. The apterae of some species of Dysaphis do have reddish or brownish areas at the base of the siphunculi which might be matched with the dull red in Buckton's sketch, but I know of none in which these areas are orange or yellow. Buckton's inconsistency on this point may well raise doubts as to the usefulness of these areas taxonomically, at least in the case of lentiginis. Moreoever, Dr F. Leclant tells me (in correspondence) that their presence in other species is variable and that he has noticed them in specimens that are about to moult.
There seems in fact to be no reliable support for the view that lentiginis might be a 'good' species apart from Buckton's host record, which cannot be substantiated. Buckton's host identifications are often unreliable and in this instance, moreover, there is added uncertainty in that he did not collect the material himself, but received it from his friend Borrer at Cowfold. I therefore place lentiginis Buckton as a synonym of plantaginea Passerini.
Siphonophora longipennis Buckton = Metopolophium dirhodum (Walker) Buckton, 1876 : 146; plate 20 bis.
(For synonymy, see Doncaster, 1961 : 58.)
LECTOTYPE here designated: alate viviparous female. Norfolk, Norwich. Poa annua. i.xi.(year ?). (Barrett). (270*).
BUCKTON'S WORKS ON APHIDOIDEA 63
Paralectotypes : i alate male, i nymphal male. Data as lectotype. (270*).
BIOMETRIC DATA. Lectotype, alata: body length 2-70 mm, both antennae from middle of segment III to apex missing, siphunculus 0-38 mm, cauda 0-24 mm, caudal hairs 13, ultimate rostral segment o-n mm, second segment of hind tarsus not measurable (both tarsi malformed), eighth tergite with 7 hairs, longest hair on hind femur 30(1., on hind tibia 35(4.
Buckton describes the apterous and alate viviparous females, the nymph and the alate male but figures only the last three morphs. The only specimens named longipennis by him consist of an alate viviparous female, an alate and a nymphal male of Metopolophium dirhodum (Walker), and an alate viviparous female of Myzus persicae (Sulzer). The slide is labelled 'Poa annua. Nov. i.', which agrees with the heading to the manuscript notes on Buckton's original sketches, where the locality is given as Norwich and the collector Barrett. These notes describe only the morphs figured, of which the alate female and nymph are doubtless taken from those morphs on 270. But although an alate male of dirhodum is present in the sample, Buckton's description and figure of the male longipennis relate to the alate female of persicae. The characteristically porrected antennal tubercles and abdominal markings of persicae, mentioned in his description, are clearly depicted in the original sketch (though less clearly in the reproduction), and the published measurement of siphuncular length agrees better with that specimen than with the male dirhodum.
I choose the alate female as lectotype of longipennis, which name has already been widely accepted as a synonym of dirhodum (Walker).
Theobald (1913 : 118) places longipennis Buckton in Macrosiphum. He gives a fuller description of the apterous viviparous female, from Poa annua in Cumberland, but quotes Buckton's account of the other morphs. He moves longipennis into Myzus in his monograph (Theobald, 1926 : 354). Hille Ris Lambers, who saw Theobald's material of longipennis, but not Buckton's, identified it with dirhodum (H.R.L., 1933 : 175), and subsequent authors have done likewise.
Siphonophora lutea Buckton = Macrosiphum (Sitobion) luteum (Buckton)
(PI. 5, fig. 63; Text-figs 36-40)
Siphonophora lutea Buckton, 1876 : 119; pi. 8, figs 1-4.
Macrosiphum luteum (Buckton) Schouteden, 1901 : 114.
Macrosiphoniella lutea (Buckton) del Guercio, 1911 : 332.
Macrosiphum luteum (Buckton); Theobald, 1913 : 82.
Macrosiphum luteum (Buckton); Laing, 1919 : 273.
Macrosiphoniella lutea (Buckton) ; Theobald, 1926 : 169.
Macrosiphum (Sitobion) luteum (Buckton); Hille Ris Lambers, 1939 : 118.
Macrosiphum luteum (Buckton); Wolcott, 1948 : 155.
Sitobium (Sitobium) luteum (Buckton) Borner, 1952 : 164.
Macrosiphum (Sitobium) luteum (Buckton) Ossiannilsson, 1959 : 494.
Sitobion luteum (Buckton) ; Smith et al., 1963 : 88.
Macrosiphum (Sitobion) luteum (Buckton); Kloet & Hincks, 1964 (I) : 82.
Macrosiphum (Sitobion) luteum (Buckton); Eastop, 1966 : 458.
Macrosiphum(Sitobion) luteum (Buckton); Mamet, 1967 : 63.
Macrosiphum (Sitobion) luteum (Buckton); Ghosh & Raychaudhuri, 1968 : 184.
64
J. P. DONCASTER
LECTOTYPE here designated: alate viviparous female. Orchidaceae. 22.1. (year ?). (Smee). (271*).
Surrey, Carshalton.
Paralectotypes : (2yia*, 2 jib*}.
i apterous viviparous female, i nymph. Data as lectotype.
Alate viviparous female. (Plate 5, fig. 63; Text-figs 36-40). Colour of macerated specimen: the insect shows signs of being teneral and is uniformly pale, with only very slight darkening of lateral abdominal sclerites, muscle-plates, siphunculi and femoral apices. Morphology: body about 2 -5 mm long, slender, nearly three times as long as broad. Head smooth, antennal tubercles distinct but not prominent. Cephalic hairs sparse, up to 2O[i long, with blunt or slightly expanded apices. Antennal segments I and II smooth except for a few scattered spinules on the ventral surface, remaining segments lightly imbricated; III with 12 and 16 circular secondary rhinaria arranged in line over a little more than three-quarters of the segment ; antennal hairs blunt, short, reaching about i6[ji on III, i.e. a little less than half its articular diameter; processus terminalis about 5-5 times as long as base of segment VI ; the whole flagellum about equal to the body length; ratios of segments III to VI 56 : 53 : 47 : 15 + 85. Rostrum about 0-65 mm in length, apical segment bluntly triangular, 0-12 mm long, scarcely longer than
36
37
FIGS 36-40. Macrosiphum (Sitobiori) luteum (Buckton). Lectotype: Fig. 36. Head, upper (left) and lower surfaces. Fig. 37. Antennal segments II I-VI. Fig. 38. Apex of rostrum (penultimate segment fractured). Fig. 39. Siphunculus. Fig. 40. Cauda.
BUCKTON'S WORKS ON APHIDOIDEA 65
second segment of hind tarsus and with six non-apical hairs. Legs long and slender; hind femur about one-third of body length, with sparse, small, spiny hairs reaching i8(i in length; tibial hairs similar, more numerous, up to 3051 long. Fore tarsi missing; first segments of middle and hind tarsi each with three hairs. Three pairs of lateral abdominal sclerites are visible on segments II to V, each more or less oval, furnished with a few spinules and one small papilla; also present on each side are a small antesiphuncular sclerite and a large postsiphuncular sclerite. Paired muscle-plates occur on I-VI. Abdominal tergum smooth, hairs sparse, short — up to i8(j. on the median area — blunt or with slightly expanded apices ; eighth tergite with four hairs, the longest about 32[z. Siphunculi 0-45 mm long, broad at the base which is 3-4 times the narrowest diameter, and tapering towards the slightly flared apex ; reticulated over the apical one-fifth to one-quarter, the remainder with a few imbrications and small groups of spinules. Cauda elongate, 0-25 mm long, slender, a little more than half as long as the siphunculi, with eight hairs.
Apterous viviparous female. This specimen, like the alata, has suffered in remounting, many of the appendages having become detached and some of their extremities lost. It appears to be more mature, with colour characters more pronounced, than the lectotype. Antennal segment III, except for its very base, is dark sclerotic, and the subsequent segments become progressively paler. Also dark sclerotic are the siphunculi and middle and hind femoral apices; the tibiae are paler. The characteristic oval, dark sclerotic patch on the abdominal dorsum between segments I and V shows clearly.
NOTES. Buckton describes the apterous viviparous female, nymph and alata. His manuscript notes give the date 22 January but add nothing further to his published data. In this instance his published figures render the colours and forms of his original sketches reasonably faithfully. His sketch of the alata, presumably drawn from life and perhaps from a specimen more mature than the lectotype, shows the siphunculi black, and the antennae, apices of femora and tibiae, and the tarsi dark.
He indicates what appear to be darkened muscle-plates but shows no lateral abdominal sclerites.
A full account of luteum is given by Hille Ris Lambers (1939 : 118).
Lachnus macrocephalus Buckton = Cinara pinicola (Kaltenbach) Buckton, 1881 : 48; pi. 97, figs i, 2.
LECTOTYPE here designated: alate male. West Sussex, Bramshott. Picea abies. Bred from nymph 26.vii.(year ?). (274).
Paralectotypes : i alate male, i nymph. Data as lectotype. (274).
BIOMETRIC DATA. Lectotype, alate male (abdomen shrivelled): body length 1-62 mm, antennal flagellum 0-96 mm, ratios of segments III-VI 56 : 21 : 22 : 21, siphuncular pore diameter 0-05 mm, basal diameter 0-21 mm, ultimate two rostral segments 0-27 mm, second segment of hind tarsus 0-32 mm, articular diameter of ant. seg. Ill 2Ojx, longest hair on ant. seg. Ill 8o\j., on vertex jO[i, on hind femur go[i, on hind tibia ±15010..
Buckton describes the apterous viviparous female, nymph and alate male, but figures only the two latter morphs under the name macrocephalus. He records (p. 50) that apterae were sent to him from spruce at Walthamstow in June and that he found the same aphid in July at Bramshott, also on spruce. Winged males from the Bramshott sample matured on 26 July. His sheet of sketches contains drawings of an apterous female, ascribed to Walker and dated June 29, which I take to be one of
66 J. P. DONCASTER
the Walthamstow specimens, and also sketches of an alate male and a nymph, described in his MS notes as 'numerous July 20 on the spruce fir, Bramshott'. The sketches of nymph and alata have been used for the figures of macrocephalus (figs i and 2), but the sketch of the aptera is used for figure 3, on the same plate, to illustrate pini L., and again for figure i, plate 5, in his subsequent paper (Buckton, 1886) where he deals with the same species.
Buckton's original slide (274), labelled 'Lachnus macrocephala. Bramshot.' (sic) contains two alate males, a nymph and an ovipara of Cinara pinicola (Kaltenbach) . One of the alate males and the nymph are doubtless the models for the two figures of macrocephalus, but I strongly suspect that the ovipara has been made to play a double role. The close correspondence of its characters and measurements with the published description suggest that it formed the basis for the 'apterous viviparous female' of macrocephalus. At the same time its attitude closely resembles that of the sketch for figure 3, which is drawn on the same sheet with macrocephalus and was perhaps at first accepted by Buckton as that species, but a pencilled note beside it suggests that he changed his mind and later referred it to pini L. This, and its ascription to Walker, lead me to exclude this specimen from the type-series and to choose as lectotype an alate male from the Bramshott sample.
Chaitophorus maculatus Buckton = Therioaphis trifolii (Monell)
(Text-fig. 41)
Calliptevus trifolii Monell, 1882 : 14.
Chaitophorus maculatus Buckton, 18996 : 277.
Callipterus genevei Sanborn, 1904 : 38.
Callipterus trifolii Monell; Davis, 19146 : 17.
[Callipterus ononidis (Kaltenbach) Theobald, 1915 : 134. Misidentification.]
Callipterus trifolii Monell; Das, 1918 : 244.
[Therioaphis ononidis (Kaltenbach) ; Theobald, 1927 : 364.]
[Therioaphis ononidis (Kaltenbach) ; Nevsky, 1929 : 316.]
[Myzocallis ononidis (Kaltenbach); Hottes & Prison, 1931 : 258.]
Myzocallis trifolii (Monell) Gillette & Palmer, 1931 : 892.
Myzocallis trifolii (Monell); Tseng & Tao, 1936 : 161.
Therioaphis collina Borner, 1942 : 273.
Pterocallidium maculatum (Buckton) Borner, 1949 : 49, 1952 : 63.
Pterocallidium lydiae Borner, 1949 : 49, 1952 : 63.
Pterocallidium propinquum Borner, 1949 : 49, 1952 : 63.
Pterocallidium trifolii (Monell) Quednau, 1954 : 35-
Therioaphis maculata (Buckton) ; Dickson et al., 1955 : 93-
Pterocallidium trifolii (Monell); Pintera, 1956 : 121.
Pterocallidium trifolii (Monell); Borner & Heinze, 1957 : 87.
Therioaphis (Pterocallidium) maculata (Buckton) Dickson, 1959 : 63.
Pterocallidium trifolii (Monell) ; Ossiannilsson, 1959 : 400.
Therioaphis trifolii (Monell); Hille Ris Lambers & van den Bosch, 1964 : 36-40.
Therioaphis trifolii (Monell); Richards, 1965 : 96.
LECTOTYPE here designated: apterous viviparous female. INDIA, Rajasthan,
BUCKTON'S WORKS ON APHIDOIDEA 67
Jodhpur, Marwar. Medicago sativa. ¥1.1897. (Collector ?). (Zoological Survey of India, Calcutta, no. 6765/117^).)
Paralectotypes : 3 apterous, 4 alate viviparous females, 3 larvae. Data as lecto- type. (Z.S.I, nos 6765/1*7(0, c), 6766/H7(a-c), 6767/117^-6).)
Apterous viviparous female. (Text-fig. 41). Colour : nearly all traces of pigmentation lost during storage. Morphology: body oval, 1-66-1-83 mm l°ng> about twice as long as broad. Head smooth, antennal tubercles absent, median frontal tubercle prominent, situated between two rather slender, slightly capitate hairs 0-33 mm long; above these a pair of stouter hairs of the same length ; vertex with two stout hairs anteriorly and four shorter ones in a line parallel with the posterior border of the head. Antennal segment I smooth with three fine, acute, very short (14(1) hairs; II smooth with one similar hair; III sparsely spinulose, slightly thickened on basal two-fifths part which bears 6-8 round or transversely oval rhinaria with thick rims. Hairs on III scarcely discernible, apparently acute, 8-iojji long, i.e. up to about half the articular diameter of the segment. No adult specimen among those examined has a complete antenna : that of a last-stage nymph (on same slide as lectotype) has length ratios of segments III-VI 100 : 59^ : 61 : 41^ + 40. Rostrum short, reaching only slightly beyond the fore coxae, apical segment 0-085 mm l°ng. bluntly conical, about two-thirds as long as second segment of hind tarsus. Legs normal, except fore coxae which are very large, nearly i£ times as wide at base as middle and hind coxae. Femoral hairs acute, short (i2-i6[z), tibial hairs acute, longer (35-40^ maximally). First tarsal segments with seven hairs, two dorsal and five ventral. Dorsal body hairs from about 35[z to 62pi long, the majority about 50^, with stout cylindrical stem, apex expanded, fan-shaped in outline. Abdominal tergites with one pair each of spinal, pleural and marginal hairs ; an accessory spinal hair is present on each of tergites I-V, giving seven hairs per tergite; VI has six hairs, VII has five, and VIII has four. Siphunculi short (0-05 mm), smooth, without flange. Cauda 0-15-0-19 mm long, knob oval, with 9-12 hairs.
Alate viviparous female. Colour : all pigment lost except in lateral abdominal sclerites, which are brownish, and the stigma of the fore wing, which shows faint traces of pigmentation.
Morphology: similar to aptera, but dorsal body hairs shorter; of those that are present and measurable, most are about half as long as the corresponding hairs in the aptera. Lateral sclerites present on abdominal segments I I-V, rounded, slightly protuberant, those on II scabrous, wart-like. Antennal segment III with 6-8 secondary rhinaria on basal part, occupying from 0-42 to 0-44 of its total length.
NOTES. The morphological similarities between maculata Buckton and the yellow clover aphid, Therioaphis tnfolii (Monell), are so close that most authors from Davis (1914) onwards have regarded the two species as identical. Dickson (1959), however, found characters by which he could separate populations of yellow clover aphid (YCA) on Trifolium spp. from populations of what had come to be known as spotted alfalfa aphid (SAA) on Medicago spp. in North America, and proposed that the latter aphid should be called maculata Buckton. The two characters which Dickson used to distinguish SAA from YCA were the area of the third antennal segment occupied by secondary rhinaria (less than half in SAA, more than half in YCA), and the presence (in SAA) or absence (in YCA) of dark sclerotic 'dashes' on the underside of the abdomen. Miss L. M. Russell, who had examined some of Buckton's type material from Calcutta, confirmed, in a letter quoted by Dickson (1959), that the sensoriation of antennal segment III in maculata agreed with that of North American SAA, and comparison of the ventral sclerotization showed that the 'dashes', though much bleached from long storage, were present but smaller and narrower than those of North American specimens. In the lectotype and paralectotypes of maculata, which
68
J. P. DONCASTER
FIG. 41 . Therioaphis trifolii (Monell) (maculata Buckton) . Lectotype : whole insect to show dorsal chaetotaxy, etc. (Right fore tibia and tarsus and smaller hairs omitted.)
BUCKTON'S WORKS ON APHIDOIDEA 69
I have examined but which Miss Russell did not see, the sensoriation of antennal segment III agrees with the specimens she did examine and with Dickson's SAA, but in none can I discern any sign of ventral 'dashes', even in alatae. This does not prove their absence, but could be due partly to bleaching and partly to many of the finer cuticular structures being obscured by contained embryos.
Hille Ris Lambers and Van den Bosch (1964) sum up our present knowledge on this subject in the light of information gained from breeding and transfer experiments. They conclude that although Dickson's characters are valid for separating YCA and SAA in North America, where the entire populations of both aphids may each have sprung from single introduced individuals or clones, these differences fall well within the normal limits of variability of trifolii alone in other parts of the world. YCA and SAA are thus merely two varieties of trifolii Monell, under which name maculata Buckton falls as a synonym.
Hyalopterus melanocephalus Buckton = Hayhurstia cucubali (Passerini)
Aphis cucubali Passerini, 1863 : 170, nee Linnaeus, 1746 : 218.
Aphis silenea Ferrari, 1872^ : 72.
Hyalopterus melanocephalus Buckton, 1879 : 116; pi. 77, figs 5-7.
Hyalopterus melanocephalus Buckton; Theobald, 1927 : 30.
Semiaphis cucubali (Passerini) Hille Ris Lambers, 1934 : 25-
Brachycolus melanocephalus (Buckton) Hille Ris Lambers, 1950 : 41.
Hayhurstia cadiva (Walker) Borner, 1952 : 109.
Hayhurstia cucubali (Passerini) Kloet & Hincks, 1964 (I) : 76.
LECTOTYPE here designated: apterous viviparous female. Norfolk, Norwich, Brandon. Silene cucubalus (syn. inflata}. I3.viii.(year ?). (Barrett ?). (283).
Paralectotypes : i apterous viviparous female, 2 nymphs, 2 larvae. Data as lectotype. (283). 5 apterous, 2 alate viviparous females. Surrey, Haslemere, Weycombe. Silene cucubalus. vii.(year ?). (Buckton). (282*).
BIOMETRIC DATA. Lectotype, aptera: body length 1-50 mm, antennal flagellum 0-64 mm, ratios of segments III-VI 25 : n : 12 : n + 21, siphunculus 0-06 mm, cauda 0-14 mm, caudal hairs 7, ultimate rostral segment 0-08 mm, second segment of hind tarsus 0-12 mm, eighth tergite with 5 (?) hairs, articular diameter of ant. seg. Ill i6pL, longest hair on ant. seg. Ill iO(z, on hind femur i6\i, on hind tibia 30(1., on eighth tergite 26^.
Buckton records melanocephalus from Haslemere and Brandon, near Norwich. His manuscript notes indicate that he received the Brandon material first, and took from it an aptera and a nymph as models for figures 5 and 6. These are dated 13 August, and the specimens were probably collected by Barrett. Buckton's description and sketch of the alata are based on specimens he took subsequently at Haslemere. All his material is cucubali Passerini.
Siphonophora menthae Buckton = Aulacorthum solani (Kaltenbach) Buckton, 1876 : 120; pi. 9, figs i, 2.
LECTOTYPE here designated: apterous viviparous female. Surrey, Haslemere. Mentha spicata (syn. viridis). vii.(year ?). (Buckton). (284^}.
yo J. P. DONCASTER
BIOMETRIC DATA. Lectotype, aptera : body length 2-60 mm, antennal flagellum not measurable (both process! incomplete), ratios of segments III-V 30 : 25 : 16 : ?, secondary rhinaria on III 2 and i, siphunculus 0-64 mm, cauda 0-25 mm, caudal hairs 7, ultimate rostral segment 0-15 mm, second segment of hind tarsus 0-13 mm, eighth tergite with 6 hairs, articular diameter of ant. seg. Ill 4Ojj,, longest hair on ant. seg. Ill iojz, on hind femur i8[L, on hind tibia 40(1, on eighth tergite 40^.
Buckton's original slide of menthae contained two alatae and two larvae of Ovatus crataegarius (Walker), an alate Myzus persicae (Sulzer), and an apterous Aulacorthum solani (Kaltenbach), all now remounted. He describes and figures the apterous and alate viviparous females of menthae. Both the figure of the aptera (fig. i) and Buckton's original sketch on which it is based show an aphid of form and colouring typical of Aulacorthum solani, and I have little doubt that the aptera from his slide is his type of the aptera of menthae. There is less certainty about the identity of the alata he described and figured, but there are indications pointing to its being persicae rather than crataegarius. The original sketch shows a predominantly green aphid with black antennae, siphunculi and lateral abdominal sclerites. The antennae are about the right proportionate length for persicae, the cauda is pale, and the siph- unculi are slightly but distinctly clavate. The abdomen, however, is without the dark dorsal patch and transverse bands characteristic of persicae. The published description, however, does mention 'some specimens' with 'disjointed transverse bars on the abdomen'.
Subsequent authors have not unnaturally assumed that when he described menthae Buckton had before him the small pale aphid found on mint that Walker (1850) first described as crataegarius (and later (1852) also as menthae and melissae}. This assumption is supported by Theobald (1926 : 279), who knew that Buckton's slide contained crataegarius and who noted that it also contained an alate persicae and 'an apterous female Myzus sp.' (i.e. solani), but did not associate the latter two with Buckton's description and figures. Hille Ris Lambers was aware that Buckton's type of the apterous menthae was solani, and informed M. D. Leonard, who quoted the information in his paper on the distribution and habits of the mint aphid (Leonard, 1963 : 55). Kloet & Hincks (1964 (I) : 80) list menthae Buckton as a synonym of solani Kaltenbach.
Siphonophora muralis Buckton = Dactynotus muralis (Buckton) (PI. 5, fig. 64; Text-figs 42-45)
Siphonophora muralis Buckton, 1876 : 157; pi. 26, figs 1-4, 7. Macrosiphum muralis (Buckton) Theobald, 1913 : 70, 1926 : 91. Dactynotus muralis (Buckton) Hille Ris Lambers, 1939 : 26. Dactynotus muralis (Buckton); Borner, 1952 : 171. Dactynotus muralis (Buckton) ; Ossiannilsson, 1959 : 503. Dactynotus muralis (Buckton); Heie, 1960 : 194, 206. Dactynotus muralis (Buckton) ; Tashev, 1964 : 163.
LECTOTYPE here designated: apterous viviparous female. Surrey, Haslemere, Weycombe. Mycelis (syn. Lactuca] muralis. 30. vi. (year ?). (Buckton). (2880,*}.
BUCKTON'S WORKS ON APHIDOIDEA
7>
Paralectotypes : 2 alate viviparous females, i nymph, i larva. Data as lectotype. \288b*. c*. d*. 289).
Apterous viviparous female. (Plate 5, fig. 64; Text-figs 42-45). (Description based on type only, an unusually large aptera with alatiform antennal sensoriation.) Colour of macerated specimen: head, rostrum most of antennae, pronotum and siphunculi dark brown. A large area on the mesonotum, the abdominal scleroites, anal and subgenital plates brown; apices of femora and tibiae, and the tarsi, more or less darkened. Remainder of body and appendages, including whole of cauda, pale. Morphology: body elongate-oval, 3-96 mm long, rather more than twice as long as broad. Head smooth with prominent antennal tubercles, dorsal hairs slender, with swollen apices, the longest reaching 55^. Antennal flagellum 0-9 of body length, ratios of segments III-VI 89 : 74 : 69 : 19 + 100; antennal hairs spiny, blunt, up to ^O\L long, not quite equal to articular diameter of third segment, which carries 40 rather small, round, secondary rhinaria distributed over nearly its whole length ; fourth segment with 6 and 7 rhinaria ; processus terminalis five times as long as base of sixth segment. Rostrum reaching to, or only slightly beyond, middle coxae, apical segment short (0-14 mm), blunt, with 8 non-apical hairs. Legs long and slender, hind femur with rather sparse hairs, variable in length, the longest about 48(jt; tibial hairs similar, reaching 50^1. First segments of tarsi with 5 hairs on all legs; second segment of hind tarsus 0-20 mm long. The dorsal abdominal hairs are nearly all carried singly on small scleroites ; the spinal hairs are duplicated on the anterior segments where their maximal length reaches about 45^; small lateral tubercles, each carried on a hair-bearing scleroite, are present on segments II-IV, or II-V; antesiphuncular sclerites are absent. Eighth tergite with 4 hairs, the longest about 70(1. Subgenital plate with 9 hairs along its posterior margin. Siphunculi 1-09 mm long, about one-quarter of the length of the body, cylindrical except for the expanded base, apical one-quarter reticulated, remainder lightly imbricated, flange small. Cauda 0-55 mm long, ensiform, slender, about three times as long as its basal width and half as long as the siphunculi, with 15 hairs.
43
44
0-5
FIGS 42-45. Dactynotus muralis (Buckton). Lectotype: Fig. 42. Head, upper (left) and lower surfaces. Fig. 43. Siphunculus. Fig. 44. Cauda. Paratype, alata (288b) : Fig. 45. Apex of rostrum.
72 J. P. DONCASTER
NOTES. Buckton describes and figures the apterous and alate viviparous females, the nymph, the alate male and the ovipara. The descriptions and figures of the first three morphs relate to muralis; those of the 'male' appear to be based on one of five alate females of Myzus persicae (Sulzer), which are included among Buckton's material of muralis, while the sketch of the 'ovipara' resembles a young larva of muralis present on 28g. His manuscript notes give the date as 30 June.
Pemphigus napaeus Buckton
Buckton, 1896 : 50.
Lectotype (designated by Doncaster, 1969 : 160) : alate viviparous female, fundatrigenia. INDIA, Kashmir, Darkot Pass. c. 3,000 m. Populus sp. galls. (Date of collection and collector not known). (292*).
Paralectotypes : fundatrix, nymph, 6 alate viviparous females. Data as lectotype. (290*, 297*, 2gia*, [29201*], 295*, 294*).
I have already given an account of napaeus Buckton elsewhere (Doncaster, 1969 : 160). All the type material is in the BMNH except one alate fundatrigenia (2920) which is in the collection of Mr D. Hille Ris Lambers of Bennekom, Netherlands.
Siphonophora olivata Buckton — Dactynotus cirsii (Linnaeus)
Aphis cirsii Linnaeus 1758 : 452, Goeze, 1778 : 299, Gmelin, 1790 : 2205.
Aphis serratulae Kaltenbach, 1843 : 25.
[Aphis sonchi Linnaeus; Walker, 1848(3 : 197 partim. Misidentification.]
[Siphonophora cichorii Koch; Buckton, 1876 : 163 partim ? Misidentification.]
Siphonophora olivata Buckton, 1876 : 164; pi. 29, figs 3, 4.
Macrosiphum githargo Theobald, 1926 : 84 ?
Dactynotiis marcatus Hille Ris Lambers, 19310 : 170.
Dactynotus olivatus (Buckton) Hille Ris Lambers, 1933 : 170.
Dactynotus cirsii (L.) Hille Ris Lambers, 1939 : 18.
Dactynotus cirsii (L.); Borner, 1952 : 170.
LECTOTYPE here designated : alate viviparous female. West Sussex, Linchmere. Cirsium vulgare (syn. Carduus lanceolatus). i4.viii.(year ?). (Buckton). (2g8a*}.
Paralectotypes: 2 apterous, i alate viviparous females. Data as lectotype. (2g8b*, c*, d*}.
BIOMETRIC DATA. Lectotype, alata: body length not measurable (that of paralectotype 2g8b is 4-20 mm), antennal flagellum not measurable (both processi incomplete), ratios of seg- ments Ill-base VI 59 : 45 : 39 : 9 + ?, secondary rhinaria on III 74, siphunculus 1-30 mm, cauda 0-68 mm, caudal hairs 27, ultimate rostral segment 0-25 mm, second segment of hind tarsus 0-21 mm, eighth tergite with 6 hairs, articular diameter of ant. seg. Ill 52^, longest hair on ant. seg. Ill 50^, on hind femur 65^, on hind tibia 65(1., on eighth tergite i2o;ji.
Buckton describes and figures the apterous and alate viviparous females, collected on the flower stems of Carduus lanceolatus (now Cirsium vulgare) at Linchmere,
BUCKTON'S WORKS ON APHIDOIDEA 73
Sussex, in mid- August. There were originally two slides named olivata, one (297) containing apterae and larvae of Dactynotiis cirsii (L.) and labelled 'Carduus arvensis. Aberdeen.', and another (298) with two apterous and two alate females, also of cirsii, from Linchmere. Buckton's manuscript notes accompanying his sketches of olivata refer only to the Linchmere sample and give the date 14 August. There is no mention of specimens from Aberdeen. I assume therefore that his description and figures relate only to the specimens on 298. These could provide a model for the alate olivata, of which the characters given are consistent with those of cirsii, but not for the aptera, which Buckton describes and figures as having a black cauda, and the published measurements of which are too small for either of the apterae on 298 (or, for that matter, any of those in the Aberdeen sample). Buckton probably used for his model an aptera of another species which has since been lost.
Theobald (1913 : 79, 1926 : 82), who redescribes the species as Macrosiphum olivatum, also states that the cauda of the aptera is black, but adds the observation (1926 : 84) that the cauda is black in the larva but pale at the base in the adult. He also found one colony in which the cauda in adult apterae was almost entirely black. This suggests that he had encountered a colony of Uromelan aenaeus Hille Ris Lambers and provides a possible clue to the identity of Buckton's aptera. (Buckton's sketch, moreover, may have been made from a larva: the cauda is too small for a typical adult, and this would account for the small dimensions given in his published account.) Borner (1952 : 170, 172) comes to the same conclusion, and includes olivata as a synonym partly of cirsii and parly of aenaeus. Kloet & Hincks (1964 : 82, 83) do likewise.
Aphis opima Buckton = Brachycaudus cardui (L.) Buckton, 1879 : 101; pi. 71, figs 1-4.
LECTOTYPE here designated: apterous viviparous female. Surrey, Haslemere, Weycombe. Cineraria, in greenhouse. Undated. (Buckton). (301).
Paralectotypes : 4 apterous viviparous females, 3 larvae. Data probably as lectotype. (299, 301) . 2 apterous viviparous females, I larva. Data as lectotype, but dated December. (303). 2 apterous, 3 alate viviparous females. Sussex, Chichester. Cineraria, in greenhouse. 3_vi.(year ?). (302).
BIOMETRIC DATA. Lectotype, aptera: body length 2-10 mm, antennal flagellum 1-32 mm, ratios of segments III-VI 49 : 32 : 24 : 13 + 49, siphunculus 0-27 mm, cauda o-n mm, caudal hairs 5 (?), ultimate rostral segment 0-19 mm, second segment of hind tarsus 0-13 mm, eighth tergite with 7 hairs, articular diameter of ant. seg. Ill 28^, longest hair on ant. seg. Ill iO(z, on hind femur 2oy., on hind tibia 50^1, on eighth tergite Sept..
Buckton describes and figures the apterous and alate viviparous females which he took on Cineraria in greenhouses. There are four slides named opima by him (299, 301, 302, 303), containing several apterous and alate Brachycaudus cardui (L.) and a few Myzus persicae (Sulzer). Buckton's description of the aptera of opima includes an account of the later larval stages while the insect is still green, and one of these he illustrates in figure i (though calling it in the caption 'green variety of
74 J. P. DONCASTER
apterous female'). Figure 2 shows the fully adult aptera with its dark pigmentation. These figures and that of the alata (fig. 3) are all consistent with the characters of cardui and can be matched with specimens of cardui on 299, 301 and 302. There is, however, no extant specimen of an early-stage larva which corresponds to the uncoloured sketch used in figure 4.
Buckton records opima from Haslemere, Chichester and Wanstead. His notes and slide labels suggest that the apterae he described and figured came from his own greenhouse at Weycombe (April-September, 3 October, 30 November), the alatae were from Chichester (3 June), and a slide of Walker's, unnamed and labelled 'Cineraria. Walk., No. 2', which contains Myzus persicae, perhaps represents the Wanstead record.
Theobald (1927 : 287) includes opima Buckton in Anur aphis and quotes Buckton's description in full. He mentions Buckton's specimens (on five slides, now numbers 299-505), which Laing had correctly identified as cardui and persicae, and concludes that Buckton's description of opima 'fits cardui perfectly well'. But Theobald mistakenly ascribes Buckton's figures of the immature aptera (fig. i) and the adult alata (fig. 3) of opima to persicae. Even in the reproduced figures the form and proportions of cauda and siphunculi alone would rule this out, while the original sketches show the typical macroscopic characters of cardui even more clearly and leave no room for doubt.
Borner (1952 : 104) places opima as a synonym of cardui, as do Kloet & Hincks (1964 : 75).
Aphis pedicularis Buckton = Aphis nasturtii Kaltenbach
Aphis nasturtii Kaltenbach, 1843 : 76.
Aphis transiens Walker, 18496 : xliv.
Aphis rhamni Koch, 1854 : 119, nee Boyer de Fonscolombe, 1841 : 177.
[Aphis acetosae F. ; Koch, 1855 : 145. Misidentification.]
Aphis pedicularis Buckton, 1879 : 41; pi. 48, figs 4, 5.
Aphis polygoni van der Goot, 1912 : 80, nee Walker, 1848 : 2249.
Aphis abbreviata Patch, 1912 : 170.
Aphis acetosella Theobald, 1918 : 286
[Aphis solanina Passerini; Theobald, 1919 : 161. Misidentification.]
Aphis githaginella Theobald, 1927 : 168.
Aphis neopolygoni Theobald, 1927 : 160.
Aphidula nasturtii (Kaltenbach) Borner, 1952 : 79.
LECTOTYPE here designated: apterous viviparous female. Norfolk, fens. Pedicularis palustris. I4.vii.(year ?). (Collector not stated, perhaps Barrett}. (3120*}.
Paralectotypes : 5 apterous viviparous females. Data as lectotype. (312).
BIOMETRIC DATA. Lectotype, aptera: body length 2-13 mm, antennal flagellum 0-90 mm, ratios of segments III-VI 29 : 21 : 19 : 14 + 29, siphunculus 0-28 mm, cauda 0-20 mm, caudal hairs 7, ultimate rostral segment o-n mm, second segment of hind tarsus o-io mm, eighth tergite with 2 hairs, articular diameter of ant. seg. Ill 22(z, longest hair on ant. seg. Ill 14(1, on hind femur 45^1, on hind tibia 45^, on eighth tergite 45(x.
Buckton describes and figures only the apterous viviparous female and young
BUCKTON'S WORKS ON APHIDOIDEA 75
larva. The single slide, named pedicularis in Buckton's hand, contained six adult apterae of what I take to be Aphis nasturtii Kaltenbach. His original sketch records rather skilfully the rounded shape, yellow-green colour and matt-textured skin characteristic of this aphid.
Endeis pellucida Buckton = Geoica eragrostidis (Passerini)
Buckton, 1883 : 91; pi. 129, figs 2, 4. (For synonymy, see carnosa, p. 41.)
LECTOTYPE here designated: apterous viviparous female. Kent, Beckenham, in ants' nest. ^.1876 (or 7.11.1879). (Lubbock). (89*).
BIOMETRIC DATA. Lectotype, aptera: body length 1-26 mm, whole antenna 0-40 mm, ratios of antennal segments I-IV 8 : 9 : 20 : 13, ultimate rostral segment 0-16 mm, second segment of hind tarsus o-io mm.
The viviparous female is described and Buckton's notes on the sheet of sketches indicate that his specimen was one of those sent by Lubbock from ants' nests. Two dates are written beside the sketch of pellucida: February, 1876, and 7 February, 1879, without indication as to which applies. Of five specimens labelled pellucida by Buckton, that which best fits his description, notes and figures is the larger of two apterous Geoica eragrostidis (Passerini) on 89, and this I believe to be his type. It differs from the type of carnosa (which is also eragrostidis) in having acute instead of flabellate hairs.
Theobald (1929 : 197) refers to what he believed was Buckton's type of pellucida, but he quotes the label of ,377, the two specimens on which, though both eragrostidis, agree less closely with Buckton's data than the one I have chosen on 89. Moreover, the label of 89 and Buckton's sketch of pellucida are both marked '£ No. i'.
None of the specimens named pellucida has antennae 'with five nearly equal joints', as Buckton says in his description, and shows in his sketch and figure 4.
Aphis penicillata Buckton = Aphis grossulariae Kaltenbach
Aphis grossulariae Kaltenbach, 1843 : 67. [No type exists.] Aphis penicillata Buckton, 1879 : 51; pi. 51, figs 5, 6. Syn. n. Aphis penicillata Buckton; Theobald, 1927 : 212. Aphidula grossulariae (Kaltenbach) Borner, 1952 : 78.
LECTOTYPE here designated: apterous viviparous female. Pembroke. Epilo- bium montanum. vii.(year ?). (Collector not stated, perhaps Barrett). (163*}.
Paralectotypes : 4 apterous, 4 alate viviparous females. Data as lectotype. (i63a*, b*, c*).
BIOMETRIC DATA. Lectotype, aptera: body length 1-92 mm, antennal flagellum 0-88 mm, ratios of segments III-VI 26 : 19 : 19 : 14 + 32, siphunculus 0-30 mm, cauda 0-22 mm, caudal hairs 12, ultimate rostral segment 0-16 mm, second segment of hind tarsus o-io mm, eighth tergite with 2 hairs, articular diameter of ant. seg. Ill 2o\L, longest hair on ant. seg. Ill 3051, on hind femur 55(z, on hind tibia 6o\L, on eighth tergite 50^, marginal tubercles present on abd. segs I-IV and VII, I-III and VII.
76 J. P. DONCASTER
Paralectotype alata: body length 2-30 mm, ant. flag. 1-18 mm, ratios segs III-VI 40 : 26 : 24 : !8 + 39, secondary rhinaria on III u, on IV 4, on V i, siph. 0-30 mm, cauda 0-21 mm, caudal hairs 13, ult. rost. seg. 0-14 mm, second seg. hind tarsus o-io mm, eighth tergite with 2 hairs, artic. diam. ant. seg. Ill 2Ofj., longest hair on ant. seg. Ill 32(x, on hind femur 50^, on hind tibia 5O(jt, on eighth tergite 60^, marginal tubercles present on abd. segs I-V and VII, I-III and VII.
Buckton describes the apterous and alate viviparous females from specimens taken at Pembroke in July and subsequently at Haslemere. I believe his types of penicillata to be five apterae and four alatae of Aphis grossulariae Kaltenbach originally mounted on 16 3, which was at first labelled simply ' Epilobium . Pembroke . s/ in Buckton's hand, and later named 'Aphis epilobii (8i)D.C. ^.' on one of Buckton's typewritten labels. (^ occurs on other slides named epilobii.} Buckton's original sketches of penicillata, entitled 'Epilobium. Pembroke. July. \.' show a rather yellowish green aptera with pale yellow-brown appendages, and an alata with black head, thorax, antennae, femora and tibial apices, and a dark green abdomen with clearly defined marginal tubercles, which are mentioned also in the text. These characters are consistent with the mounted specimens, the apterae of which have the unpigmented head, stigmal plates and cauda, as well as marginal tubercles on many of the abdominal segments, characteristic of grossulariae.
Theobald (1927 : 212) quotes Buckton's original description in full and mentions a Buckton slide of specimens he had seen and believed to be penicillata, and of which he adds some details. This slide is 318, which contained seven alatae and three nymphs, but no apterae, of Aphis grossulariae Kaltenbach (now remounted) and had been tentatively named '? penicillata by Laing. The original slide apparently carried no data except the code ^ and another, partly obliterated, which Theobald interpreted as \. y occurs elsewhere only on slides named by Buckton urticaria, which contain a mixture of urticata F. and confusa Walker. The fact that penicillata follows urticaria in the monograph and the two are figured on the same plate may have led Laing to conclude that these specimens were the types of penicillata. I cannot disprove his conclusion, but prefer to regard as Buckton's types the specimens on 163, which carries data that link them with his sketches and also includes adult apterae, which are absent from 318. Buckton recognizes (1879 : 72) that penicillata is distinct from his concept of epilobii Kaltenbach, which he appears to have based on a mixture of epilobiaria Theobald and praeterita Walker, and I think it likely that, having described and figured the specimens on 163 as penicillata, he omitted to alter the name on the label.
Aphis petasitidis Buckton = Brachycaudus helichrysi (Kaltenbach) Buckton, 1879 : 69, pi. 58, figs i, 2.
LECTOTYPE here designated: alate viviparous female. Northumberland, Holy Island. Cynoglossum or Pyrethrum. i.vii.(year ?). (Hardy). (332).
Paralectotypes : 2 apterous, 4 alate viviparous females, 3 larvae. Data as lecto- type. (332). 8 alate viviparous females, 7 nymphs, i larva. East Hertfordshire, Albury. Petasites hybridus. 15. vi. (year ?). (Collector not stated) . (331,333)-
BUCKTON'S WORKS ON APHIDOIDEA 77
BIOMETRIC DATA. Lectotype, alata: body length 1-60 mm, antennal flagellum 1-12 mm, ratios of segments III-VI 45 : 27 : 17 : 12 + 41, secondary rhinaria on III 20, on IV 2, on V o, siphunculus 0-14 mm, cauda 0-09 mm, caudal hairs 6, ultimate rostral segment 0-13 mm, second segment of hind tarsus 0-12 mm, eighth tergite with 7 hairs, articular diameter of ant. seg. Ill i8(z, longest hair on ant. seg. Ill 14(1, on hind femur 2Opt, on hind tibia 3O(i, on eighth tergite 8o[i.
The adult aptera, nymph and alata are described, but only the nymph and alata figured. The hosts are given as Tussilago petasites (now Petasites hybridus) and Cynoglossum officinale, and the localities Albury, Herts, and Holy Island, Northumber- land. Of the three slides labelled petasitidis by Buckton, two (331, 333} contain specimens from the Albury sample, and the third (332) aphids from Holy Island. The label on the last records two hosts, Cynoglossum and Pyrethrum. Of the total of 28 specimens, all but two accidental inclusions are Brachycaudus helichrysi (Kaltenbach).
Buckton records in his notes and in the text that the alata he figured gave birth to two young while under the microscope. An alata in the Holy Island sample shows some similarities in attitude with Buckton's sketch, and the slide includes some young larvae. It seems likely that this is his type of the alate petasitidis and I choose it as lectotype. Though both sketch and published figure show the siphunculi as considerably longer than those of helichrysi, the measurement Buckton gives for siphuncular length is more nearly typical, i.e. 0-17 mm, or about one-eighth of the body length. In the lectotype this proportion is about one-tenth.
Glyphina pilosa Buckton = Schizolachnus pineti (Fabricius)
Buckton, 1883 : 16; pj. 116, figs 1-4. (For synonymy, see fuliginosa, p. 57.)
LECTOTYPE here designated; alate viviparous female. Surrey, Haslemere, Meadfields. Pinus sylvestris. 29.vii.(year ?). (Buckton). (4).
Paralectotypes : i alate, 2 apterous viviparous females, i nymph. Surrey, Haslemere, Weycombe. Pinus sylvestris. 2Q.vii.i874 ? (Buckton). (3). i alate, I apterous viviparous female ('pineti' in Walker's hand). Middlesex, Southgate. Pinus sylvestris. 25. vi. 1847. (Walker). (W.6$3).
BIOMETRIC DATA. Lectotype, alata: body length 1-44 mm, antennal flagellum 0-74 mm, ratios of segments III-VI 43 : 15 : 18 : 17, secondary rhinaria on III 7, siphuncular pore diameter 0-06 mm, ultimate rostral segment 0-12 mm, second segment of hind tarsus 0-25 mm, articular diameter of ant. seg. Ill i8[z, longest hair on ant. seg. Ill ioo[z, on hind femur i8o[i, on hind tibia
Paralectotype aptera: body length 1-76 mm, ant. flag. 0-76 mm, ratios of segs III-VI 43 : 18 : 18 : 18, siph. pore diam. 0-08 mm, ult. rost. seg. 0-14 mm, second seg. hind tarsus 0-29 mm, artic. diam. ant. seg. Ill 34^, longest hair on ant. seg. Ill 140(1., on hind femur 170^, on hind tibia
Buckton describes the apterous and alate viviparous females, but figures the larva and alata. The specimens, his own from Haslemere, and others sent him by Walker from Southgate, were taken from Pinus sylvestris. There are no slides named
7& J. P. DONCASTER
Glyphina pilosa, but two of Buckton's slides, and one of Walker's in Buckton's collection, all named Mindarus abietinus by Buckton, contain specimens of Schizo- lachnus pineti (F.) from Pinus sylvestris at Weycombe, Meadfields (both near Hasle- mere) and Southgate, and are believed to be the types of pilosa.
Buckton evidently thought at first that these specimens were Mindarus abietinus Koch, but changed his opinion when he found that in the fore wings of his alatae the media was apparently unbranched. This ruled out Mindarus and led him to erect pilosa as a new species in Glyphina. In one of his three alatae (that on slide 4} the media is indeed simple, but in that on 3 the branch is discernible though very faint. Walker's alata (W.653) has one fore wing crumpled but in the other the proximal branch of the media cannot be seen. Buckton's original sketch of the alata shows the fore wings each with unbranched media, but alongside is drawn a wing with branched media and the note 'very faint in some specimens'. This wing is not reproduced on plate 116, and in his published description Buckton states unequivocally that the media (he calls it the cubital) is unforked, and believes Walker to be mistaken in supposing — correctly — that the aphid is pineti.
I choose as lectotype the alata on 4, a rather small specimen which agrees better with the published measurements than the others, and is likely to be the one used for figure 2.
Theobald in a footnote (1929 : 81) quotes Laing's opinion, supported by that of Swain (1921 : 212), that Buckton's specimens named Mindarus abietinus are pineti F. and likely to be the types of pilosa. Subsequent authors have accepted this conclusion. Borner (1952 : 40) remarks that individuals of pineti with the media unbranched in one or both fore wings occur as aberrations.
Pterocomma pilosum Buckton
(PI. 6, fig. 65; Text-figs 46-50) Buckton, 1879 : 143; pi. 83, figs 1-5.
(The taxonomy of the genus Pterocomma is so confused that I prefer not to attempt a synonymy of pilosum Buckton. A recent review of the Tribe Pterocommatini Mordvilko is that of Szelegiewicz (1965).)
LECTOTYPE here designated: alate viviparous female. London, Kentish Town. Salix sp. twigs. 28. ix. (year ?). (Knaggs). (342*).
Paralectotypes : 5 larvae, i nymph. Data as lectotype. (343).
Alate viviparous female. (Plate 6, fig. 65; Text-figs 46-50.) (Description based on lectotype only.) Colour of macerated specimen : mostly pale brownish, with the heavily sclerotized parts, e.g. pterothorax, anal plate, knees, rather darker. The dark sclerotic transverse bands shown in the figure are scarcely discernible. Morphology : body large and thick, 3-04 mm long, 1-06 mm broad (I accept Buckton's measurements here, because the specimen has become unnaturally elongated by pressure in remounting). Head with numerous fine, acute hairs up to lyopi long. Antenna! flagellum 1-52 mm, i.e. about half the length of the body, ratios of segments III-VI 60 : 30 : 26 : 16 + 21, antennal hairs rather numerous, fine, acute, long (up to 40^) except on Vlth segment, which has seven hairs up to about gojj. on the base and, on the processus, 4-5
BUCKTON'S WORKS ON APHIDOIDEA
79
much shorter hairs in addition to the 3-4 terminal sensillae. Second antennal segments each with five hairs. Third segments with 37 and 34 rather large, circular secondary rhinaria, mostly on the postero-ventral surface, the fourth with i and 2. Rostrum (detached from specimen; total length not measurable) with ultimate segment broad, tapering only slightly towards apex, 0-20 mm long, slightly longer than second segment of hind tarsus (0-18 mm), with 9 non-apical hairs in two lateral rows. Legs stout with numerous fine hairs, the longest reaching 150^ on the hind femora and 165^1 on the hind tibiae. First tarsal segments on all legs with 5 hairs. Abdomen densely clothed with fine hairs, the longest reaching from lyofj. on tergite III to aoopi on tergite VIII. Marginal tubercles absent. Eighth tergite with 14 hairs. Subgenital plate with 29 hairs. Siphunculi pale, short, 0-22 mm long, about 1-3 times as long as the cauda,
46
50
0-5
FIGS 46-50. Pterocomma pilosum (Buckton). Lectotype: Fig. 46. Head, upper (left) and lower surfaces. Figs 47, 48. Left antenna. Fig. 49. Apex of rostrum. Fig. 50. Siphunculus.
8o J. P. DONCASTER
more or less cylindrical, with small flange. Cauda U-shaped, 0-17 mm long, four-fifths as long as its basal width, with about 20 long (150(1) hairs.
NOTES. Buckton describes the apterous and alate viviparous females and the nymph from specimens taken among colonies of Pterocomma (Melanoxantherium) salicis (L.) feeding on willow twigs. His manuscript notes are dated 28 September, but his published account gives the date as August. The plate contains figures of the three morphs described, but the 'aptera' (fig. i) is drawn to a smaller scale than the other two morphs and gives the impression of being a young larva. Buckton's sketch for fig. i is an accurate drawing of the largest of the larvae on 343, which is in fact larger than the nymph used as the model for fig. 2.
Siphonophora polygoni Buckton = Nasonovia ribisnigri (Mosley)
Aphis lactucae Schrank, 1801 : 120, partim, non L.
Aphis ribisnigri Mosley, 1841 : 684.
Aphis ribicola Kaltenbach, 1843 : 33.
Aphis hieracii Kaltenbach, 1843 : 17, partim; Walker, 18490 : 47.
Siphonophora alliariae Koch, 1855 : 177; Buckton, 1876 : 123.
Siphonophora polygoni Buckton, 1876 : 123; pi. 10, figs 1-3.
[Siphonophora lactucae (L.) ; Buckton, 1876 : 139. Misidentification.]
[Siphonophora cichorii Koch; Buckton, 1876 : 163, partim. Misidentification.]
[? Myzus ribis (L.); Buckton, 1876 : 180, partim. Misidentification.]
Macrosiphum kaltenbachi Schouteden, 19060 : 237.
? Macrosiphum agrostemnium Theobald, 1913 : 146.
Nasonovia ribicola (Kaltenbach) Mordvilko, 1929 : 51, 81.
Submacrosiphum hieracii ssp. teriolanum Hille Ris Lambers, 19316 : 10.
Nasonovia ribisnigri (Mosley) Hille Ris Lambers, 1947 : 316, Borner, 1952 : 136.
LECTOTYPE here designated: alate viviparous female. Surrey, Haslemere, Weycombe. Polygonum persicaria. 27. vi. 1872. (Buckton). (359*).
Paralectotypes : i alate viviparous female, 3 nymphs. Data as lectotype. (559*).
BIOMETRIC DATA. Lectotype, alata: body length 2-04 mm, antennal flagellum 2-56 mm, ratios of segments III-VI 33 : 21 : 18 : 6 -f 52, secondary rhinaria on III 46, on IV 9, on V o, siphunculus 0-44 mm, cauda 0-22 mm, caudal hairs 7, ultimate rostral segment 0-17 mm, second segment of hind tarsus 0-14 mm, eighth tergite with 4 hairs, articular diameter of ant. seg. Ill 3Ojj., longest hair on ant. seg. Ill 35^, on hind femur 3O[i, on hind tibia 45^, on eighth tergite
50(JL.
All the five specimens on Buckton's single slide of polygoni are Nasonovia ribisnigri (Mosley). His original sketches, as well as the published figures, are consistent with the appearance of this species in life and I regard these specimens as his types. Only the alate female and the nymph are described and figured.
Theobald (1926 : 329) places polygoni Buckton in Myzus and paraphrases Buckton's description. He includes also a brief description of the aptera, which Buckton omits. The specimens which Theobald