SIM. BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 35 BRITISH MUSEUM (NATURAL HISTORY) LONDON : 1977 ISSN 0524-643 1 Printed in England by Unwin Brothers Limited CONTENTS ENTOMOLOGY VOLUME 35 No. i. A review of Cinara subgenus Cinarella (Hemiptera : Aphididae). By V. F. EASTOP i No. 2. American leaf -litter Thysanoptera of the genera Erkosothrips, Eurythrips and Terthrothrips (Phlaeothripidae : Phlaeothripinae). By L. A. MOUND 25 No. 3. A revision of the families Synneuridae and Canthyloscelidae (Diptera). By A. M. HUTSON 65 No. 4. A taxonomic revision of the genus Callipielus Butler (Lepidoptera : Hepialidae). By G. S. ROBINSON 101 A REVIEW OF CINARA SUBGENUS (HEMIPTERA : APHIDIDAE) V. F. EASTOP BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 35 No. i LONDON : 1976 A REVIEW OF CINARA SUBGENUS CINAREL (HEMIPTERA : APHIDIDAE) BY VICTOR FRANK EASTOP Pp. 1-23; 14 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 35 No. i LONDON : 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in Jive 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. 35 No. I of the Entomology 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.) ISSN 0524-6431 Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 25 November, 1976 Price 3-25 A REVIEW OF CINARA SUBGENUS CINARELLA (HEMIPTERA : APHIDIDAE) By V. F. EASTOP CONTENTS Page SYNOPSIS ........... 3 INTRODUCTION .......... 3 KEY TO THE APTERAE viviPARAE OF Cinarella ..... 4 THE SPECIES OF Cinarella ......... 7 HOST PLANTS OF Cinarella ........ 17 BlOMETRIC DATA FOR THE APTERAE VIVIPARAE OF Cinarella . . . 1 8 DISCRIMINANT PROPORTIONS FOR THE APTERAE VIVIPARAE OF Cinarella . 20 ACKNOWLEDGEMENTS ......... 21 REFERENCES ........... 21 INDEX ............ 23 SYNOPSIS A key is given for the separation of the viviparous morphs of the species of Cinarella and a few other similar species of Cinara. References are given to the original descriptions, principal redescriptions, and synonyms of each species. Data on host plants and geographical distribution are summarized and biometric data are tabulated. Where possible, references are given to accounts of the biology of each species. INTRODUCTION THE purpose of this paper is to provide a key for the identification of the species of Cinarella Hille Ris Lambers, 1948 and a lead in to the literature concerning them. Members of Cinarella are known from all continents except Australasia and the Antarctic, feed on species of Pinus of economic importance and are frequently misidentified. The recent outbreaks in Brazil of Cinara maritimae, originally described from France, and of C. piniformosana, originally described from Taiwan, indicate the need for a revision of this group of species. Both C. maritimae and C. piniformosana have been misidentified with C. pinea recently and C. pinea has also been known as C. pineti and C. pini. Hille Ris Lambers (1948 : 275) proposed Cinarella as a new name for Borner's (1939 : 76) concept of Cinara Curtis, 1835, and used it as a subgenus of Cinara to include both Lachnus pineus Mordvilko as type and his new species Cinara excelsae. References to the original descriptions and type citations of these genus-group names are to be found in Eastop (1972 : 104-106) as are those of Cinarella Borner, 1949 nee Hille Ris Lambers, 1948, its replacement name and the other subgenera and synonyms of Cinara. Cinarella Hille Ris Lambers, 1948, constitutes one of the better defined species-groups of the genus Cinara, being characterized by the 4 V. F. EASTOP absence of a mesosternal tubercle in the apterae viviparae (present in the summer apterae of Cinara s. str.) and the elongate first tarsal segments of all morphs: see Clnara piniformosanus (Text-figs 7-11, p. 15). The species of Cinarella usually bear rather thick body hairs and the apterae tend to have stalked eyes and their third and fourth antennal segments rarely bear rhinaria (often present in the apterae of Cinara s. str.). However, alatiform apterae of Cinarella may bear rhinaria on the third and fourth antennal segments. As some species of Cinara s. str. such as laricis have rather elongate first tarsal segments it can be difficult to assign alatae and Cinarella is not regarded as more than a subgenus of Cinara Curtis. Cinarella species live only on Pinus and with few exceptions (see p. 17) are confined to subsections Sylvestres and Australes of the section Pinus (= Diploxylon). Cinarella are widely distributed; to judge from their host plants, five species originated in the Palaearctic region, three from the Oriental region and two from the Nearctic region. Several of these species have been distributed more widely by commerce. Information about the distribution of Pinus has mostly been obtained from Critchfield & Little (1966), Little & Critchfield (1969) and Mirov (1967). References to the principal accounts of Cinara were given by East op (1972 : 104). Fossel (1970 : 129-190; 1972 : 125-144, 145-156, 185-191) has given accounts of the Austrian fauna. Ehrhardt & Kloft (1962 : 544-546) give details of a method for rearing Cinara species in. the laboratory. Existing accounts of Cinara deal only with fauna of parts of continents and one of the purposes of the present paper is to provide a key for recognizing species wherever their origin. The alatae viviparae of several species are unknown but as the alatae of Cinara are usually similar to their apterae, the following key can be used for alatae as well as for apterae if the rhinarial characters are ignored. In general rostral characters are the same in alatae as in apterae. As alatae with long hind tibia bearing short hairs are rare, the ratio of these measurements for alatae is the lower half of that ratio for apterae. Similarly the ratio of tarsal segments 2 : i for alatae tends to be at the upper end of that ratio for apterae. KEY TO THE APTERAE VIVIPARAE OF Cinarella Third antennal segment 3-0-3-7 times as long as the maximum diameter of the siphuncular cones and 12-14 times as long as the longest hair borne on the third antennal segment. Hind tibiae about 30-35 times as long as the longest hair borne on them. Fifth antennal segment 0-75-0-90 times as long as the sixth. Fourth rostral segment 2-9-3-0 times as long as the fifth segment and 0-9 to equal in length to the first segment of the hind tarsus. Second segment of hind tarsus 2-0-2-2 times as long as the fourth rostral segment and 1-9-2-1 times as long as the ventral length of the first segment of the hind tarsus. Anterior abdominal segments without scleroites. A rather short-haired species, hairs on third abdominal tergite up to only about 3~35 A< m l n g an d longest hairs on hind tibiae about 60 jum long. lachnirostris (p. 9) Third antennal segment 0-8-2-7 times as long as the diameter of the siphuncular cones and 3-5-13-0 times as long as the longest hair borne on the third antennal segment and if more than 10 times as long then the fourth A REVIEW OF CINARA SUBGENUS CIN A RELLA 5 rostral segment is i -2-1 -8 times as long as the first segment of the hind tarsus. Hind tibiae either 9-28 or 45-50 times as long as the longest hair borne on them. Fifth antennal segment 1-0-1-7 times as long as the sixth. Fourth rostral segment 1-5-2-6 times as long as the fifth rostral segment 2 2 (i) A rather short-haired species (Text-fig. 2), the longest hairs on the third abdominal tergite about 35 /um long. The hind tibiae are 3-7-4-2 mm long and 45-50 times as long as the longest hair borne on them. Third antennal segment (Text-fig, i) about 12-13 times as long as the longest hair borne on it. Processus terminalis often bearing 5 subapical hairs. formosana (p. 8) The longest hairs on the third abdominal tergite usually 70-200 ,um long, if only 35-65 /urn long (schimitischeki) then hind tibiae 2-2-3-0 mm long and 15-27 times as long as the longest hair borne on them. Third antennal segment usually 3-2-9-3 times as long as the longest hair borne on it but 9-4-12-0 times as long in schimitsheki (see previous character). Most species with only 4 subapical hairs on the processus terminalis but several species 5 or 6 subapical hairs ......'.... 3 3 (2) Longest hairs on third abdominal tergite 35-100 /urn long and evidently shorter than the longest hairs, 100-150 pm, borne on the hind tibiae. Hairs on abdominal tergites 1-6 arising from small dark scleroites about 20-25 /* m in diameter (Text-figs 12, 13). Third antennal segment 5-9-12-0 times as long as the longest hair borne on it. Hind tibiae dark, mostly black but slightly paler at the proximal one-third and 15-27 times as long as the longest hairs borne on them. Second segment of hind tarsus 0-9-1-2 times as long as the fourth rostral segment which is i -4-1 -8 times as long as the first segment of the hind tarsus. Eighth abdominal tergite bearing 27-60 hairs on a transverse dark band which is almost or completely broken in the mid dorsal line ........ schimitscheki (p. 14) Longest hairs on third abdominal tergites 70-210 fim long and if less than no ^m long then usually with some arising from larger scleroites with a diameter of 50-77 pm or more. Third antennal segment 3-2-9-3 times as long as the longest hair borne on it and if more than 5-5 times as long then either the second segment of the hind tarsus is i -5-1 -8 times as long as the fourth rostral segment or the fourth rostral segment is 0-8-1-2 times as long as the first segment of the hind tarsus or the hairs on the third abdominal tergites are up to 130-180 ^am long and evidently longer than those of up to 70-130 (im long on the hind tibiae. Eighth abdominal tergite bearing 13-28 hairs. ........... 4 4 (3) Fourth rostral segment 310-360 /zm long and 2-4-2-6 times as long as the fifth rostral segment and i -5-1 -8 times as long as the first segment of the hind tarsus. Second segment of hind tarsus 0-9-1-1 times as long as the fourth rostral segment. Hind tibiae 9-11 times as long as the longest hair borne on them. Scleroites small and pigmented bar on eighth abdominal tergite broken medianly. ....... atrotibialis (p. 7) Fourth rostral segment 150-300 /um long, 1-7-2-4 times as long as the fifth segment and 0-7-1 -7 times as long as the first segment of the hind tarsus, but if more than 1-5 times (maritimae] then hind tibiae 1626 times as long as the longest hair borne on them. Second segment of hind tarsus 1-1-2-4 times as long as the fourth rostral segment. Hind tibiae 11-28 times as long as the longest hair borne on them. ....... 5 5 (4) Shorter-haired species: longest hairs on abdominal tergites 2-5 only about 70-80 ^m long and when arising from scleroites these have a greatest diameter of less than 75 fim. Third antennal segment 6-15 times as long as the longest hairs borne on it. Hind tibiae with a pale area at about the basal one-quarter. 6 V. F. EASTOP Second segment of hind tarsus only 310-370 //m long yet 1-5-1-8 times as long as the fourth rostral segment (190-230 fj,m) .... 6 Longer-haired species: longest hairs on abdominal tergites 2-5 from 80-200 /am long and if less than 120 /*m then either some arise from scleroites with a maximum diameter of 75-180 /urn (pinea), or the hind tibiae are black (piniformosana) . Third antennal segment 3 -5-7-5 times as long as the longest hair borne on it. Second segment of hind tarsus 1-1-2-4 times as long as the fourth rostral segment but when 1-5-1 -8 times, the second segment of the hind tarsi usually exceeds 370 fim. in length except in pergandei in which the third antennal segment is only 3 -5-4-5 times as long as the longest hair borne on it. 7 6 (5) Second segment of hind tarsus i -5-1 -9 times as long as the first segment. Fourth rostral segment i -o-i -3 times as long as the first segment of the hind tarsus. Third antennal segment 6-10 times as long as the longest hair borne on it. Diameter of siphuncular cone 4-0-5-5 times that of its porus. Processus terminalis bearing 4 or 5 subapical setae. Some of the dorsal abdominal hairs arising from scleroites with a maximum diameter of up to 65-75 pm. . piniphila (p. 14) Second segment of hind tarsus 2-1-2-6 times as long as the first segment. Fourth rostral segment i -41 -6 times as long as the first segment of the hind tarsus. Third antennal segment 10-15 times as long as the longest hair borne on it. Diameter of siphuncular cone only 2-3-3-5 that of its porus. Processus terminalis bearing 5-7 subapical setae. Scleroites not evident. hyperophila (p. 8) 7 (5) Largest scleroites on the second to fifth abdominal tergites with a greatest diameter of 75-300 ^wrn. Second antennal segments bearing 5-10 hairs and base of sixth antennal segment bearing 2-8 hairs. Processus terminalis usually bearing 4, exceptionally 3 or 5 subapical setae. Second segment of hind tarsus i -3-2-2 times as long as the fourth rostral segment which is often 0-8-1-2 but sometimes 1-2-1-4 times as long as the first segment of the hind tarsus. Hind tibiae usually 11-15 but sometimes 15-20 times as long as the longest hair borne on them. ........ 8 Largest scleroites on abdominal tergites 2-5 with a maximum diameter of 25-75 /um. Second antennal segments bearing 6-14 hairs and base of sixth antennal segment bearing 6-12 hairs. Species with 4, 5 or 6 subapical setae on the processus terminalis. Second segment of hind tarsus 0-9-1-6 times as long as the fourth rostral segment which is i -21 -8 times as long as the first segment of the hind tarsus. Hind tibiae usually 15-26 but sometimes only 12-15 times as long as the longest hair borne on them. . . 10 8 (7) Fourth rostral segment i -2-1 -4 times as long as first segment of hind tarsus and 2-0-2-4 times as long as the fifth rostral segment. Second segment of hind tarsus 1-3-1-6 times as long as the fourth rostral segment and 1-6-2-1 times as long as the first segment of the hind tarsus. Many of the hairs on the seventh abdominal tergite arising from a fenestrated transverse band. First tarsal segments without dorsal hairs. .... neubergi (p. n) Fourth rostral segment 0-7-1-2 times as long as the first segment of the hind tarsus and 1-6-2-3 times as long as the fifth rostral segment. Second segment of hind tarsus 1-4-2-2 times as long as the fourth rostral segment and i -4-1 -9 times as long as the first segment of the hind tarsus. Hairs of the seventh abdominal tergite usually arising from individual large scleroites which only rarely anastomose to form a much fenestrated transverse dark band. ............ 9 9 (8) First tarsal segments usually bearing i or 2 dorsal hairs. Tibiae usually pale, sometimes dusky over the distal one-third. Most hairs on tergites 1-7 arising from more similarly sized and often transversely elongate scleroites A REVIEW OF CINARA SUBGENUS CINA RELLA 7 (Text-fig. 6) of from 75 x 40 //m to 180-50 fim diameter with few if any scleroites of about 14 /um diameter. Fifth abdominal tergite bearing 28-49 hairs between the siphunculi. Transverse dark band on eighth abdominal tergite often complete, broken in the mid line in about one-third of the specimens studied. ........ pinea (p. 12) First tarsal segments without dorsal hairs. Tibiae usually black, sometimes the very bases of the fore and mid tibiae and the basal third of the hind tibia are paler. Hairs on abdominal tergites 1-7 arising from roundish scleroites of very different sizes, most of the scleroites are little more than pigmented hair bases of 10-15 i" m diameter while others on each segment have a diameter of 75-100 ftm. Fifth abdominal tergite bearing 50110 hairs between the siphunculi. Transverse dark band on eighth abdominal tergite usually broken in the mid line, the two parts being linked by a narrow band of pigmentation occasionally. ...... pergandei (p. 12) 10 (7) Second segment of hind tarsus (Text-fig. 1 1) i -7-2-4 times as long as the fourth rostral segment (Text-fig. 10) which is 150-205 //m long and 0-8-1-2 times as long as the first segment of the hind tarsus. The tibiae are black. Eighth abdominal tergite bearing a broad transverse band from which the hairs arise (Text-fig. 7). Hairs on the seventh abdominal tergite arising from large scleroites which in heavily pigmented specimens anastomose to form a fenestrated transverse band. ..... piniformosana (p. 13) Second segment of hind tarsus 1-1-1-6 times as long as the fourth rostral segment which is 250-300 fim long and 1-2-1-7 times as long as the first segment of the hind tarsus. Hind tibia with a pale area at about the basal one-third. Fifth abdominal tergite bearing 30-60 hairs between the siphunculi (Text-fig. 4). Eighth abdominal tergite bearing a dark transverse band which is sometimes broken medianly. ..... n 11 (10) Hind tibiae 16-27 times as long as the longest hair (70-130 ^m) borne on them. Third antennal segment (Text-fig. 3) 4-0-7-4 times as long as the longest hair borne on it and 1-5-2-0 times as long as the fourth rostral segment. Second antennal segment bearing 7-1 1 hairs, base of sixth antennal segment bearing 7-12 hairs and processus terminalis bearing 4 or 5, rarely 3 or 6, subapical setae. First tarsal segments with or without dorsal hairs. Seventh and eighth abdominal tergites both bearing dark unbroken transverse bands, that on the seventh tergite being fenestrated in lightly pigmented specimens. ......... maritimae (p. 10) Hind tibiae 12-15 times as long as the longest hair (150-175 /urn) borne on them. Third antennal segment 3-5-5-0 times as long as the longest hair borne on it and usually 2-0-2-4 times as long as the fourth rostral segment. Second antennal segment bearing 6-9 hairs, base of sixth antennal segment bearing 6-7 hairs and processus terminalis bearing 4, rarely 3, subapical setae. First tarsal segments without dorsal hairs. Pigmented band of eighth abdominal segment broken in the mid dorsal line and seventh abdominal segment with only scleroites. .... watsoni (p. 17) THE SPECIES OF CINARELLA Cinara atrotibialis David & Rajasingh Cinara atrotibialis David & Rajasingh, 1968 : 103-105. [Type, Madras Christian College; INDIA: Assam, Shillong, Pinus sp., 19^.1967.] Cinara sp., Robinson, 1972 : 604. Cinara khasyae Robinson, 1972 : 1925-1927. 8 V. F. EASTOP SPECIMENS STUDIED. THAILAND: Mt Suthep, Pinus khasya, 26.111.1970, 2 apterae (paratypes of khasyae). INDIA: Assam, Shillong, Pinus insularis, 10.1.1960, 2 immature specimens (A. K. G [hosh].); Himachal Pradesh, Simla, Pinus sp., 1.1970, immature specimen (L. K. Ghosh). PHILIPPINE Is.: Baguio Mt Prov., Pinus insularis, 24^1.1969, i apt era ( V. J. Calilung) . HOST-PLANTS. Pinus insularis ( khasya). DISTRIBUTION. India (Assam, Himachal Pradesh), Thailand, Philippine Is. NOTES. The single specimen seen from the Philippine Is. had a body length of only 2-7 mm but the fourth rostral segment was. 264 jum long. The relationship between body length, fourth rostral segment length and temperature during development in aphids is complex but it is possible that the Philippine specimen represents a distinct subspecies and that khasyae is also subspecifically distinct. Pinus insularis consists of geographically isolated populations, those on the Asian mainland being known as P. khasya and those in the Philippine Is. as P. insularis. There is unlikely to be much gene flow between the aphid populations of the Himalayas, Thai mountains and Philippine Is. Cinara formosana (Takahashi) (Text-figs i, 2) Dilachnus sp., Takahashi, 1923 : 46. Dilachnus formosanus Takahashi, 1924 : 73-74. [Type, Taiwan Agricultural Research Institute; TAIWAN: 'Formosa', Taihoko, Shinten, Pinus sp., 13. x. 1923, attacking the shoot.] [Panimerus piniformosanus (Takahashi) Takahashi, 1928 : 28 nee 1921. Misidentification.] Neochmosis formosanus (Takahashi) Takahashi, 1930 : 325. Cinara formosana (Takahashi) Takahashi, 1931 : 23; 1936 : 493; Tseng & Tao, 1936 : 123; Tao, 1962 : 44-45; Higuchi & Miyazaki, 1969 : 31; Inouye, 1970 : 67-68. Panimerus kiangsiensis Lou, 1935 : 37-42. [Types unknown; CHINA: Kiang-si, Lou-chan, 25. iv. 1930, numerous alatae, Yian, Wei-i.] SPECIMEN STUDIED. CHINA: Fukien, i aptera, M. S. Yang coll. (no. 5319) (R. Takahashi). HOST-PLANTS. Recorded from Pinus thunbergii and P. densiflora and more rarely from P. luchuensis and P. massoniana of the subsection Sylvestres. DISTRIBUTION. Seen from China, originally described from Taiwan (Formosa) and also recorded from Japan. Cinara hyperophila (Koch) Lachnus hyperophilus Koch, 1855 : 232-234. [Types unknown AUSTRIA: Konigswiessen, Pinus sylvestris, May, June, October, November.] Lachnus pineus var. hyperophilus Koch, Mordvilko, 1894/5 : 102. Cinara (Subcinara) hyperophila (Koch) Pasek, 1954 : I 39~ I 4o; Heinze, 1962 : 156, 157. Cinara hyperophila (Koch) Szelegiewicz, 1962 : 82; Pintera 1966 : 289, 290. A REVIEW OF CINARA SUBGENUS CINA RELLA g - Ife FIGS 1,2. Cinara formosana, aptera vivipara. i, second and third antennal segments, x 100. 2, fourth and fifth abdominal tergites, X 40. SPECIMENS STUDIED. POLAND: Otwock, Pinus silvestris, 4.vii.i887, 2 juveniles, A. Mordvilko coll. (H, Szelegiewicz). HOST-PLANT. Pinus sylvestris. DISTRIBUTION. Austria, Czechoslovakia and Poland. The record from Sweden appears to have been based on a fundatrix of Cinara pini (L.). NOTES. A little known central European species of which no adults have been seen by the present author. It was included in the key from the data provided by Pintera (1966 : 290). Koch (1855 : 2 34) found numerous males in April and May on the small trees from which he had collected oviparae in September and October. This suggests that hyperophila was not adapted to conditions at Konigswiessen. As hyperophila survives at not much higher latitudes in Czechoslovakia and Poland the small trees Koch studied in Austria may have been growing in a sheltered position and in receipt of supplementary light and heat during the winter. Cinara lachnirostris Hille Ris Lambers Cinara lachnirostris Hille Ris Lambers, 1966 : 201-203. [Type, Hille Ris Lambers collection, Bennekom, Netherlands; data as for specimen studied.] io V. F. EASTOP SPECIMEN STUDIED. PAKISTAN: Murree, 2250111, Pinus ? excelsa, 3. 11.1964, i aptera (D. Hille Ris Lumbers) . HOST-PLANT. Probably Pinus wallichiana (= excelsa Wallich). DISTRIBUTION. Pakistan (Murree, 2250 m). NOTES. Probably not really a Cinarella but the first tarsal segments are long enough to take it to Cinarella in some keys. Cinara maritimae (Dufour) (Text-figs 3, 4) Aphis pini subsp. maritimae Dufour, 1833 : 243-245. [Types unknown; presumably FRANCE: 'du pin maritime'.] [Cinara pineti (F.) Blanchard, 1939 : 869. Misidentification.] Cinara excelsae Hille Ris Lambers, 1948 : 273-274. [Types Hille Ris Lambers collection; 'PALESTINE': Beth-Hakerem, Pinus excelsa, 29.^.1946; ISRAEL: Jerusalem, 26. iv. 1946.] Bodenheimer & Swirsky, 1957 : 245. Cinara (Cinarella) excelsae Hille Ris Lambers; Qanakcioglu, 1966 : 138. Cinara maritimae (Dufour) Eastop & Hille Ris Lambers, 1976 : 135. SPECIMENS STUDIED. CYPRUS: Nicosia, pine, ig.iii.ig68, 5 apterae, G. P. Georghiou coll. (R. C. Dickson). FRANCE: Massif de 1'Esterelle, Pinus ? maritima, 3i.viii.i959, 2 apterae, i alata (D. Hille Ris Lambers); Brittany, La Trinite sur Mer, P. ? maritima, 25.vii- T-5.viii.i968, 6 apt., i al. (V.F.E.). MOROCCO: Rabat, P. maritima, 1.1928, 6 apt., 1 al. (/. Mimeur) F. V. Theobald colln. ISRAEL: Ein-Hsishofeth, Pinus sp., 2 apt.; Jerusalem, 23.iv.i946, 2 apt. (E. Swirski). TURKEY: Aydin-Mazon ormani, 650 m, 18. 11.1964, 2 apt. (H. ^anakfioglu). BRAZIL: S. P., Campinas, suction trap, iv-vi.i968, 5 al. (C. L. Costa); S.P., Pinus sp., 2i.v.i969, 9 apt. 3 al. (C.L.C.); S.P., Piricicaba, Pinus elliottii, 25-26. vi. 1968, 5 apt. 6 al.; 2i.v.i969, 5 apt. 5 al. (C.L.C.); P. caribaea bahamensis, 25.viii.7i, n apt. 2 al., A. Zamith coll. (F. Mariconi); R. S., Sanata Maria, Pinus sp., 7 apt. 2 al., (D. Link); R.S., Sao Sepe, P. elliottii, 22.xi.72, 4 apt. i al. (V.F.E.). CHILE: Conception, 28.1.1961, 2 al. (D. Hille Ris Lambers). HOST-PLANTS. Seen from members of the Pinus halepensis (= maritima Lamarck) /pinaster (= maritima Poiret) group from which it was originally described and recorded from P. thunbergii and possibly P. densiftora also of the subsection Silvestres. Specimens have also been seen from P. caribaea and P. elliottii of the subsection Australes and are also recorded from introduced P. canariensis (subsection Canariense) and P. radiata (as insignis, subsection Oocarpae) in Israel (Swirski, 1963 : i). DISTRIBUTION. Seen from Cyprus, France, Morocco, Israel, Turkey, Brazil and Chile and probably also occurring in Argentina (Blanchard, 1939 : 869, as pineti F.). NOTES. Although described more than 140 years ago, widespread in the A REVIEW OF CINARA SUBGENUS CINA RELLA FIGS 3, 4. Cinara maritimae, aptera vivipara. 3, second and third antennal segments x 100. 4, fourth and fifth abdominal tergites, X 70. Mediterranean region, extending as far north as Brittany on pines planted for coastal protection and also occurring in South America, C. maritimae is rarely mentioned in entomological literature. C. maritimae has probably been confused with C. pinea which is very common on Pinus sylvestris in Europe but in which the fourth rostral segment is only 0-8-1-2 times as long as the second segment of the hind tarsus (1-3-1 -7 in maritimae). Cinara neubergi (Arnhart) Lachnus neubergi Arnhart, 1930 : 392-398. [Types unknown; AUSTRIA: Neuberg a. d. Miirz, July 1927/9, twigs of Pinus montana.] Cinara neubergi (Arnhart) Heinze, 1962 : 158; Szelegiewicz, 1962 : 69, 86; Pintera, 1966 : 286-287; Fossel, 1970 : 164, 170, 176, 182. SPECIMENS STUDIED. ITALY: Valle Campo, 2000 m, Pinus mugo pumilio, 2.ix.i975, 2 apterae (A. Binazzi). SWITZERLAND: Grimmels, 2100 m, P. mugo, 4.viii.i956, i apt., W. Eglin coll. (D. Hille Ris Lambers). HOST-PLANT. Pinus mugo (= montana Miller nee Lamarck). 12 V. F. EASTOP DISTRIBUTION. Originally described from the Austrian Alps, seen from Italy and Switzerland and also recorded from Bulgaria, Germany and U.S.S.R. (Ukraine). NOTES. Braun (1930 : 485) thought that neubergi was a variety of C. pinea. Cinara pergandei (Wilson) Lachniella pergandei Wilson, 1919 : 46-47. [Types, USNM; U.S.A.: Va, Pinus inops, June, 1903 & 1905, T. Pergande coll.] Cinara longispinosa Tissot, 1932 : 4-5. [Type, Fla agric. expt. Stn; U.S.A.: Fla, Gainsville, Pinus taeda, 3.viii.i929, A. N. Tissot coll.] Cinara pergandei (Wilson) Palmer, 1945 : 451-452; Pepper & Tissot, 1973 : 75-83. SPECIMENS STUDIED. CANADA: Manitoba, Winnipeg, glasshouse, 2i.iii.ig62, 6 apt., Pinus banksiana, greenhouse, 2.x. 1961, i al. (G. A. Bradley). U.S.A.: Fla, Gainsville, Pinus glabra, u.xii.i964, i apt. (J. 0. Pepper & A. N. Tissot); Iowa, Ames, jack pine, 22.vii.i924, 5 juveniles (F. C. Hottes); N.C., Toxawdry, Pinus echinata, 27.viii.i957, i apt. (W. R. Richards}. HOST-PLANTS. Originally described from Pinus inops (== virginiana) and seen from P. banksiana also of the subsection Contortae. C. longispina was originally described from P. taeda, has been seen from P. echinata and P. glabra and has also been recorded from P. clausa and P. rigida all of the subsection Australes. There is also a single record from P. tanyosho (= densiflora] of the subsection Sylvestres. NOTES. Bradley (1951 : 334) gives an account of the biology and Bradley & Hincks (1968 : 42, 45) give accounts of parasites and attendant ants. Cinara pinea (Mordvilko) (Text-figs 5, 6) Lachnus pineus Mordvilko, 1894/5 : 75~77> %> 82, 94, 100; 1895 : IO2 . 126-130. Lachniella inoptis Wilson, 1919 : 18-19. Cinara pinea (Mordvilko) Borner, 1932 : 569 and many later authors. [See Eastop, 1972 : 156-160; Fossel, 1970 : 164, 170, 176, 182; 1972 : 125-144; Pepper & Tissot, 1973 : 84-93-] Cinara diver siseta Borner, 1952 : 41, 241. Cinara kaltenbachi Hottes, 1954 : 170, footnote. SPECIMENS STUDIED. 76 apterae viviparae, 40 alatae viviparae, see Eastop, 1972 : 157-159 for details. HOST-PLANTS. Pinus sylvestris and occasionally other Pinus spp. (see Eastop, 1972 : 159 for details). DISTRIBUTION. Europe, from Britain to U.S.S.R. (Caucasus) and Sweden to Italy (see Eastop, 1972 : 161 for details) and introduced to North America. Japanese and other Oriental records may apply to other species, mostly C. piniformosana. A REVIEW OF CINARA SUBGENUS CINA RELLA FIGS 5, 6. Cinara pinea, right siphunculus and part of fifth abdominal tergite, x 87. 5, alata. 6, aptera. NOTES. Fossel (1972 : 125-144) describes two forms of C. pinea from Austria which may be another expression of the polymorphism found in British alatae (Eastop, 1972 : 159-160). Scheurer (1971) found that C. pinea was little affected by ants, unlike C. pini and C. escherichi ( nuda) which were dependant on them. Pechhacker (1974 : 42-45) has discussed the effect of pest control operations in forests on populations of Cinara pinea and other aphids and ants. The dorsal abdominal scleroites tend to be smaller in the alatae than in the apterae (Text-figs 5,6). Cinara piniformosana (Takahashi) (Text-figs 7-11) [Lachnus pini (L.) Maki, 1915 : 35. Misidentification.J Dilachnus sp., Takahashi, 1921 : 82. 14 V. F. EASTOP Dilachnus piniformosana Takahashi, 1923 : 47; Hori, 1929 : 58. Neochmosis piniformosanus (Takahashi) Takahashi, 1930 : 325. [Cinara pineti (Koch) Takahashi, 1931 : 22-23; Tao, 1962 : 35-36; Paik, 1965 : 17-18. Misidentification . ] [Cinara formosana (Takahashi) Inouye, 1939 : 141-142; 1956 : 215-216; Moritsu, 1956 : 338. Misidentification.] Cinara piniformosana (Takahashi) Inouye, 1970 : 63-65. SPECIMENS STUDIED. JAPAN: Honshu, Kyoto, Pinus sp., 6.^.1964, 3 apterae (H. Takada). BRAZIL: S.P., Campinas, water traps, v-vi, 1968, i al. (C. L. Costa); S.P., Suzano, Pinus densiflora -f- thunbergii, 27.viii.i97i, 3 apt., Pinus sp., x.i97i, 5 apt., 3 al., S.P., Carlos, Pinus sp., ? 1971, 9 apt. (F. Mariconi). HOST-PLANTS. Usual host Pinus densiflora and also seen from P. thunbergii and recorded from P. sylvestris, P. luchuensis and P. massoniana, all of the subsection Sylvestres. DISTRIBUTION. Seen from Japan and Brazil, originally described from Taiwan (Formosa) and also recorded from Korea. BIOLOGY. Attended by various ants and overwintering as eggs in Japan where the males are alate (Inouye, 1970 : 64). NOTES. Paik (1972 : 146-147) illustrates a species with very short first tarsal segments, a very hairy fourth rostral segment and a short fifth rostral segment under the name C. piniformosana (Takahashi, 1929) which does not seem to be either C. piniformosana (Takahashi, 1923) or C. formosana (Takahashi, 1924). Cinara piniphila (Ratzeburg) Aphis piniphila Ratzeburg, 1844 : 219-220, pi. 2, fig. 5. [Types unknown; locality not stated, presumably GERMANY : needles and young twigs of Pinus sylvestris.] Lachnus pineus var. curtipilosa Mordvilko, 1895 : I 3- I 32. [Types unknown; POLAND: Kielce, June, 1893.] Eulachnus mingazzinii del Guercio, 1909 : 326-329. [Types unknown; ITALY: Florence, Pinus sylvestris. .] Cinara piniphila (Ratzeburg) Borner, 1952 : 41; Szelegiewicz, 1962 : 86-87; Pintera, 1966 : 288- 289. SPECIMEN STUDIED. GERMANY: Vogelherd Wald, Pinus sylvestris, ii.v.i936, i aptera (K. Heinze). HOST-PLANT. Pinus sylvestris. DISTRIBUTION. Seen and originally described from Germany and subsequently recorded from Austria, Czechoslovakia, Italy and Poland. Cinara schimitscheki Borner (Text-figs 12-14) Cinara schimitscheki Borner, 1940 : i. [Types, Deutsches Entomologisches Institut, Ebers- walde ; GERMANY : Naumburg, twig ends of Pinus austriaca.'] Tremblay & Micieli de Biase, 1970 : 210-213 and many others, see Eastop, 1972 : 164. A REVIEW OF CINARA SUBGENUS CINA RELLA FIGS 7-11. Cinara piniformosana, aptera vivipara. 7, whole insect, x 20. 8, antenna, x 55- 9. sixth antennal segment, x no. 10, rostrum, apex of second to fifth segment, X no. n, hind tarsus, X no. i6 V. F. EASTOP Cinara kosarowi Tashev, 1962 : 207-210. [Types, University of Sofia; BULGARIA: Pinus leucodermis , g . vii . 1 960] . SPECIMENS STUDIED. In addition to those listed by Eastop (1972 : 165). GREAT BRITAIN: England, Surrey, Kew Gardens, Pinus nigra, 30. 1.1965, 4 juveniles; 10. vii. 1971, i apt.; I7.vi.i972, 2 apt. (V.F.E.). ITALY: Campini near Florence, Pinus pinea, 2i.v.i973, numerous juveniles (4. Binazzi}. 12 13 FIGS 12-14. Cinara schimitsheki, aptera vivipara. 12, part of right side of fifth abdominal tergite, x 87. 13, enlargement of part of 12, x 450. 14, fourth and fifth rostral segments, x 175- A REVIEW OF CINARA SUBGENUS CINA RELLA 17 HOST-PLANTS. Usually on Pinus nigra including several of its varieties and also recorded from P. leucodermis and P. mugho subsp. mughus of the subsection Sylvestres. A large sample of immature specimens has been seen from P. pinea of the subsection Pinea. DISTRIBUTION. Europe, England to the U.S.S.R. (Crimea) and Turkey, but not Baltic or Scandinavian, see Eastop, 1972 : 165. BIOLOGY. Lives singly on the young shoots during May and June but not evident after early July when it is said by Pasek (1954 : 175) to go under the bark of older branches. Cinara watsoni Tissot Cinara watsoni Tissot, 1939 : 43; Pepper & Tissot, 1973 : 132-140. SPECIMENS STUDIED. U.S.A.: Fla, Gainsville, Pinus taeda, 5.^.1933, i aptera (paratype) (4. N. Tissot); 7.111.1965, i al. (J. 0. Pepper); Pennsylvania, Philipsburg, Blk Mash Dam, Pinus rigida, i.viii.i954, 4 apt. (J.O.P.); Scotia, P. rigida, I5.viii.i964, i al. (J.O.P. & A.N.T.). HOST-PLANTS. Seen from Pinus rigida and P. taeda and also recorded from P. echinata, elliotti, glabra, palustris and serotina, all of the subsection Australes. DISTRIBUTION. U.S.A., widespread over about the eastern third, see Pepper & Tissot, 1973 : 139. NOTES. Brooks & Warren (1964 : 310-316) give an account of the biology of C. watsoni and Werner & Clark (1969 : 436-437) of control measures. HOST PLANTS OF CINARELLA Pinus L. Section Strobus (= subg. Haploxylon or section A. Haploxylon of Shaw, 1914). Subsection Strobi (= group Flexiles of Shaw). P. peuce/wallichiana group including excelsa and griffithii. Cinara lachnirostris Section 4 Pinea Subsection Canariense P. canariensis Cinara maritimae Subsection Pineae P. pinea Cinara schimitscheki Section 5 Pinus (= subg. Diploxylon or section B. Diploxylon of Shaw or subg. Eupitys of Rehder, 1949) Subsection Sylvestres (= Laricones of Shaw) i8 V. F. EASTOP TABLE Biometric data for the Host plant Number of specimens Body length mm Diameter of sipbuncular cone Diameter of scleroites on abdominal tergites 2-5 Length in pirn of antennal segments III IV lachnirostris wallichiana i 2-4-3-2 140-180 absent 490-570 170-210 hyperophila sylvestris o not evident formosana thunbergii i 4-0-5-5 680-750 10-15 (60) 720-760 300-320 schimitscheki nigra 12 3-3-5-1 440-720 20-25 630-860 270-350 fiiniphila sylvestris 2 3-9-4-5 490-530 10-75 460-590 180-250 neubergi mugho 3 4-2-4-3 570-670 100-150 540-650 250-300 pinea sylvestris 76 3-1-5-1 270-700 75-i8o 500-740 220-360 pergandei Pinus spp. 7 2-9-4-5 310-570 100-310 490-620 190-230 piniformosana thunbergii 35 2-4-4-1 300-550 50-75 440660 170-260 utrotibialis insularis 3 2-8-4-3 350-490 45-65 510-700 230-320 tnaritimae pinaster 37 2-5-4-2 330-600 10-45 (-60) 390-620 160-270 watsoni rigida 4 2-7-4-3 380-480 60-70 440-630 210-330 length of longest hair on third hind abdominal antennal tibia tergites segment 3 Number of secondary rhinaria on antennal segments III IV lachnirostris 40-45 55-65 30-40 70-75 o(-3) o(-3) (-1) hyperophila 50 80 formosana 55-65 75-85 30-40 i 30-140 o O-I 1-2 schimitscheki 70-120 100-150 35-100 95-130 o o (o )i( 2) piniphila 60-80 90-120 70-80 70-90 o O neubergi 110-130 170-200 140-170 50-180 o o 13 pinea 90-210 120-230 95-210 20-230 o(-4) 0(-2) 0(-2) pergandei 110-170 150-190 i 20-200 20-200 o o-3 O-2 piniformosana 90-140 90-160 80-160 IO-2IO 0<-2) o O utrotibialis 90-140 170-200 110-130 00-170 o O maritimae 60-120 70-130 120-180 2O-I8O (0)1 2( 6) (O )l 2( 4) 0-2 watsoni 110-150 150-175 130-150 2O-I8O o o(-D I P. densiflora, luchuensis, massoniana, thunbergii group Cinara formosana, C. piniformosana and possibly C. maritimae P. halepensis (= maritima Lamarck), pinaster (= maritima Poiret) group Cinara maritimae P. insularis (including kesiya = khasya) Cinara atrotibialis P. leucodermis & nigra (= austriaca) including caramatica (= pallasiana) Cinara schimitsheki P. luchuensis, massoniana see densiflora P. mugo (= montana Miller nee Lamarck) Cinara neubergi P. nigra see leucodermis. P. maritima see halepensis & pinaster A REVIEW OF CINARA SUBGENUS CINA RELLA aperae viviparae of Cinarella Length in urn of antennal segments V VT Length of Length of rostral hind tarsal segments in |zm segments in pirn Hind tibiae V 4 5 i 2 length in colour mm 230-260 2OO-2IO + 75-IOO 170-180 55-65 80-190 360-370 1-9-2-0 paler on basal third 190210 105 20-150 310-370 paler on basal third 280-300 170-180 + 55-65 270-280 130-140 20-230 380-390 3-7-4-2 paler on basal half, 300-380 130-200 + 50-70 290-360 130-170 70-220 320-380 2-2-3-0 black 280-330 180-210 + 45-65 220-230 100 no 90-210 350-360 2-0-2-7 paler on basal third 300-340 I7O-2IO + 5O-65 280-300 125-145 20-240 390-450 2-3-2-7 paler on basal third 280-400 140-230 + 40-75 210-290 110-170 20-330 350-530 1-8-3-4 pale or spotted 260-360 175-220 + 50-65 150-230 70-110 80-230 300-370 2-0-2-8 dark, paler basally 230-320 130-210 + 50-75 150-210 85-105 60-215 320-430 1-7-2-5 black 280-410 150-240 + 60-80 310-360 125-140 80-210 290-360 1-7-2-0 dark 210-350 140-210 + 40-70 250-300 110-140 60-220 320-420 1-5-2-5 dark, paler basally 280-350 160-190 + 35-55 200-280 110-150 70-210 290-350 1-9-2-5 dark, paler basally Number of hairs on first tarsal segment dor sally II antennal segments VI processus Base terminalis accessory hairs on ultimate rostral segment abdominal 3 tergites 8 sub-genital plate subapically 9 10 5-8 5-6 19 12 37 9 5-7 7-8 7-8 5 6 24 13 56 o 9-13 6-9 (3-) 4 6 26-73 27-60 50-62 I 10-13 7-1 1 4 4-6 29-63 22 44 7-8 5-7 4 4-5 18-28 I5-l8 36 1-2 5-9 2-8 (3-)4(-5) 4-6 18-49 13-26 22-52 O 6-10 4-8 4 4 50-110 15-28 40-55 7-14 6-10 4 (-5) 4-7 30-50 14-25 22-50 6-8 7-8 4 4-6 25-26 I5-I6 37 0-2 7-1 1 7-12 (3-) 4-6 4-7 30-60 15-28 26-41 6-9 6-7 (3-) 4 4(-6) 50-60 13-19 45-55 P. pinaster see halepensis P. resinosa Cinara pinea P. sylvestris including hamata (= sosnovskyi) Cinara hyperophila, C. pinea, C. piniformosana, C. piniphila P. thunbergii see densiflora Subsection Australes P. caribaea Cinara maritimae P. clausa Cinara pergandei P. echinata, glabra, taeda Cinara pergandei, C. watsoni 20 V. F. EASTOP be en 4) g M MOOO ONI^M ^t^fOOO MOO en tn T3 rt e -*- 1 11 1 1 1 1 1 1 1 1 1 1 1 O\wvO'OOO'O Tj-iot^>OtxvO 8^ .00 >OrON n5 OMO'^-O'* > NOONO'<-rO 2 J* o rONNNNNNNNNMN 1 1 1 1 1 1 1 1 1 1 1 <^oo o^o o>oooo r>Tj-O !> 1 1 > ? 1 1 1 1 1 1 I 1 I 1 1O NfOWNMMONOl-l < O o g 5 l?r 8 fa S ^ g J3 1 1 m oo t^oo o o >o ro w >o o rn T^-NMMNI-INMNM 1 1 1 1 1 1 1 1 1 1 1 M t-^i/IOONNwioOv^ON ^ XI ^13 N oO>Ol^NOi-irOl^OTl- g a ^3^ g ? 3 ** fl SP o o S) fO NNNNPOPOPO'-INN 1 1 I 1 1 1 I 1 1 1 a\ oc^oooa^fovoioioo 5) i f ^ 2 8 N WwNMMWNi-it-iM 1 1 .fa -^ g rt - X C Tj- ro^O^^T^P^P 9 ^ |>1 fi HI MIHIHlOVO^'Ot^tVIO 1 1 1 1 1 T 1 1 1 1 1 N NO\Of>N>o-OO ^rofirnrt-'l-ro H A -t-> u I- -2 ^,4) t^t^M ION H w ^r^io-^-o 0) S-s c S S o g o M CJ T N liliililll O lOO^OOOOOOO OO^MOO M T3 & 1 '03 rowQi-iOOOOOt-iOi-i to >tt 2 Q ^ a C > M S f> <* m H i|* IHit iiiiliiiiiil iliuluiii! A REVIEW OF CINARA SUBGENUS CINARELLA 21 P. elliotti Cinara maritimae, C. watsoni P. pungens, rigida, serotina Cinara watsoni P. taeda see echinata & glabra Subsection Contortae ( = Insignes of Shaw in part) P. banksiana Cinara pergandei, C. watsoni P. virginiana (= inops) Cinara pergandei, C. pinea Subsection Oocarpae (= Insignes of Shaw in part) P. radiata ( = insignis) Cinara maritimae ACKNOWLEDGEMENTS Most of the specimens studied in the course of this work are in the collection of the British Museum (Natural History). Thanks are due to the collectors indicated with collection data. Drs D. Hille Ris Lambers, F. Ossiannilsson, J. O. Pepper, J. Petterson & H. L. G. Stroyan have also kindly made other relevant material available. Mr J. Lewis has provided information about species of Pinus and the literature concerning them. Text-figs 5, 6 and 12-14 are the work of Mrs J. Palmer and 7-11 of Mr A. Sutton. REFERENCES All but the more recent references may be traced in the bibliographies of aphid literature published by Sharma (1969-72), Smith (1972) and Szelegiewicz (1969). Most of them are in the bibliography of Eastop's (1972 : 176-184) account of British Cinara and are not repeated here. BROOKS, H. R. L. & WARREN, L. O. 1964. Biology of a pine bark aphid, Cinara watsoni, and its response to temperature. /. Kans. ent. Soc. 37 : 310-316. CRITCHFIELD, W. B. & LITTLE, E. L. 1966. Geographic distribution of the pines of the world. Misc. Publs U.S. Dep. Agric. no. 991 : v + 97 pp. DAVID, S. K. & RAJASINGH, S. G. 1968. A new species of Cinara Curtis (Hemiptera: Aphidoidea) from eastern India. Orient. Insects 2 : 103-105. EASTOP, V. F. 1972. A taxonomic review of the species of Cinara Curtis occurring in Britain (Hemiptera: Aphididae). Bull. Br. Mus. nat. Hist. (Ent.) 27 : 104-186. & HILLE Ris LAMBERS, D. 1976. A survey of the Aphididae of the world. 586 pp. The Hague. EHRHARDT, P. & KLOFT, W. 1962. A simple method of rearing Lachninae for investigation in the laboratory. Trans. Int. Congr. Ent. 11(2) : 544-546. FOSSEL, A. 1970. Anleitung zur Determination einiger in Mitteleuropa verbreiten Vertreter des Genus Cinara Curt. (Aphidoidea, Lachnidae). Waldhygiene 8 : 129-190. 19720. Die Populations dichte einiger Honigtauerzeuger und ihre Abhangigkeit von der Betreuung durch Ameisen. Waldhygiene 9 : 185-191. 22 V. F. EASTOP 19726. Vergleichende Messungen an Cinara (C.) pinea (Mordv.) (Homoptera, Aphidina). Mitt, naturw. Ver. Steierm. 102 : 125-144. HILLE Ris LAMBERS, D. 1966. New and little known aphids from Pakistan (Homoptera, Aphididae). Tijdschr. Ent. 109 : 193-220, pi. HOTTES, F. C. 1954. Proposed addition to the 'Official List of specific names in Zoology' of the specific name 'pini' Linnaeus, 1758, as published in the binominal combination 'Aphis pini' and as interpreted by DeGeer (1773) (Class Insecta, Order Hemiptera). Bull. zool. Nom. 9 : 166-183, 184-187, 188-190. KEARBY, W. H. & BLISS, M. 1969. Field evaluation of three granular systemic insecticides for control of the aphids Eulachnus agilis and Cinara pinea on Scotch pine. /. econ. Ent. 62 : 60-62. LITTLE, E. L. & CRITCHFIELD, W. B. 1969. Subdivisions of the genus Pinus (Pines). Misc. Publs U.S. Dep. Agric. no. 1144 : 51 pp. Lou, HOAI-PAO. 1935. Recherches sur la faune aphidologique de la Chine. 124 pp, 6 figs. Lyon. MIROV, N. T. 1967. The genus Pinus. viii -+- 602 pp. New York. PAIK, W. H. 1972. Illustrated encyclopaedia of fauna & flora of Korea. 13, Insecta (V) Aphididae. 751 pp. including 80 pis. PALMER, M. A. 1945. Supplementary notes on ten described species of Lachnini (Aphidae). Ann. ent. soc. Am. 38 : 447-453. PECHHACKER, R. H. 1974. Uber die Wirkungen chemischer Forstchadlings-bekampfungen aus der Luft auf Honigtau-Erzeuger und Ameisen. Anz. Schddlingsk. 47 : 42-45. PEPPER, J. O. & TISSOT, A. H. 1973. Pine-feeding species of Cinara in the eastern United States (Homoptera: Aphididae). Fla agric. exp. Stn Monogr. Ser. 3 : 160 pp. + 6 pis. RATZEBURG, J. T. C. 1844. Die Forst-Insecten. 3 : 314 pp. + 16 pis (Aphis pp. 205-223). ROBINSON, A. G. i972a. Annotated list of aphids (Homoptera: Aphididae) collected in Thailand, with description of a new genus and species. Can. Ent. 104 : 603-608. 19726. New species of aphids (Homoptera: Aphididae) from Thailand. Can. Ent. 104 : 1925-1929. SCHEURER, S. 1971. Einfluss der Ameisen und der natiirlich Feinde auf einige an Pinus silvestris L. lebende Cinarinen in der Diiboner-Heide (DDR). Polskie Pismo ent. 41 : 197 : 229. SHARMA, M. L. 1969-1972. Bibliography of Aphidoidea. 1, 293 pp. (1969), 2, 221 pp. (1971), 3, 207 pp. (1972). Quebec. SMITH, C. F. 1972. Bibliography of the Aphididae of the world. Tech. Bull. N. Carol, agric. Exp. Stn no. 216 : 717 pp. SWIRSKY, E. 1963. Notes on plant lice (Aphidoidea) of Israel. Israel}, agric. Res. 13 : 9-23. SZELEGIEWICZ, H. 1969. Mszyce-Szkodniki roslin wektory chorob wirusowych i producenci spadzi - bibliografia. Kom. ochr. Rosl. Polsiej Akad. Nauk. Warsaw, 250 pp. TAKAHASHI, R. 1923. Aphididae of Formosa. Part 2. Rep. Govt Res. Inst. Dep. Agric. Formosa 4 : 1-173 + 9 P^ s + 3 PP- index. 1930. Some Aphididae of Loochoo. Trans, nat. Hist. Soc. Formosa 20 : 317-327. 1936. Some Aphididae from south China and Hainan (Homoptera), I. Lingnan Sci. J. 15 : 595-606. TAO, C. C. 1962. Aphid fauna of China, Lachninae. [In Chinese.] Sci. Yb. Taiwan Mus. 5 : 33-82. TISSOT, A. H. 1932. Six new aphids from Florida with descriptions of the sexual forms of another species. Fla Ent. 16 : 1-13. TREMBLAY, E. & MICIELI DE BIASE, L. 1970. Notulae Aphidologae. II. Notizie sugli Afidi del Pinus nigra Arn. Boll. Lab. Ent. agr. Filippo Silvestri 28 : 204-223. WERNER, R. A. & CLARKE, E. W. 1969. Absorption, translocation, and distribution of phorate in loblolly pine seedlings. /. econ. Ent. 62 : 436-437. WILSON, H. P. 1919. Some new lachnids of the genus Lachniella (Homoptera-Hemiptera). Can. Ent. 51 : 18-22, 41-47. A REVIEW OF CINARA SUBGENUS CINA RELLA INDEX Synonyms are in italics; principal page references are in bold. atrotibialis, 5, 7-8, 18, 20 Cinara, 3 Cinarella Borner, 3 Cinarella Hille Ris Lambers, 3, 4 curtipilosa, 14 diver siseta, 12 escherichi, 12 excelsae, 10 formosana, 5, 8, 9, 14, 18, 20 hyperophila, 6, 8-9, 18-20 inoptis, 12 haltenbachi, 12 khasyae, 7 kiangsiensis , 8 kosarowi, 16 lachnirostris, 4, 9-10, 17, 18, 20 laricis, 4 longispinosa, 12 maritimae, 3, 7, 10-11, 17-21 mingazzinii, 14 neubergi, 6, 11-12, 18, 20 mula, 12 pergandei, 7, 12, 19-21 pinea, 3, 7, n, 12-13, 18-21 pineti, 3, 10 pini, 3, 9, 13 piniformosana, 3, 4, 7, 8, 12, 13-14, 15, 18-20 piniphila, 6, 14, 19, 20 schimitscheki, 5, 14, 16-17, 20 watsoni, 7, 17, 18-21 V. F. EASTOP, D.Sc. Department of Entomology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD ENTOMOLOGY SUPPLEMENTS 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, 1965- 3-25. 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. 19- 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. 19. CROSSKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. 6.50. 20. VON HAYEK, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. 12.30. 21. CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. 9-55- PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND AMERICAN LEAF-LITTER THYSANOPTERA OF THE GENERA ERKOSOTHRIPS, EURYTHRIPS AND TERTHROTHRIPS (PHLAEOTHRIPIDAE : PHLAEOTHRIPINAE) L. A. MOUND BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 35 No. 2 LONDON : 1976 AMERICAN LEAF-LITTER THYSANOPTERA OF THE GENERA ERKOSOTHRIPS, EURYTHRIPS AND TERTHROTHRIPS (PHLAEOTHRIPIDAE : PHLAEOTHRIPINAE) BY LAURENCE ALFRED MOUND Pp. 25-64; 50 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 35 No. 2 LONDON : 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Scientific 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. 35 No. 2 of the Entomology 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.) ISSN 0524-6431 Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 25 November, 1976 Price 4-75 AMERICAN LEAF-LITTER THYSANOPTERA OF THE GENERA ERKOSOTHRIPS, EURYTHRIPS AND TERTHROTHRIPS (PHLAEOTHRIPIDAE : PHLAEOTHRIPINAE) By L. A. MOUND CONTENTS Page SYNOPSIS ........... 27 INTRODUCTION ........... 27 ECOLOGY AND EVOLUTION IN LEAF-LITTER THYSANOPTERA ... 28 GENERIC RELATIONSHIPS ......... 29 GENERA EXCLUDED FROM THE Eurythrips COMPLEX .... 29 ACKNOWLEDGEMENTS ......... 30 CHECK LIST OF THE SPECIES DISCUSSED IN THIS PAPER . . . . 31 Erkosothrips STANNARD ......... 33 Key to species .......... 33 Eurythrips HINDS .......... 37 Key to species ........... 40 Terthrothrips KARNY. ......... 62 REFERENCES ........... 63 INDEX ............ 64 SYNOPSIS Keys are provided to the eight species of Erkosothrips (in which two new specific synonyms and two new combinations are established) and 38 species of Eurythrips (in which 15 new specific synonyms and six new combinations are established). A check list is given and includes 27 species of Terthrothrips. These genera are restricted to North and South America. Reasons are presented for accepting the validity of the two African genera Porcothrips and Zuluiella. INTRODUCTION THE Thysanoptera which live in leaf litter provide a series of complex taxonomic problems. Many species are widespread in this habitat, a few display clinal variation in structure (Mound, 19720.), others display allometric patterns of growth which are sometimes sex linked (Mound, 19746), yet others display confusing patterns of variation which do not at present have any explanation (Mound, 19726). It is not only at the species level that this variation is difficult to interpret, the genera of leaf-litter thrips are frequently ill-defined assemblages of more or less similar forms. Unfortunately most of the nominal species have been based on only one sample, or one morph or even one specimen. Despite the fact that there are 54 specific names available in the genus Eurythrips few of these species have ever 28 L. A. MOUND been illustrated or compared one to another, and there is little discussion in the literature of intraspecific variation. Before the genera can be better denned it is essential to re-examine the variation within and between the known species, and this is the limited objective of the present paper. The only published key to the species of Enrythrips is limited to those species which have been found in Illinois (Stannard, 1968). Most of the species in this genus were described by J. D. Hood whose collection was not generally available to other workers until after it was purchased by the Smithsonian Institute in 1964. The specific criteria used by Hood can now be shown to be periodically inconsistent, and some of his descriptions are inaccurate. Moreover the type-series of several species described by J. R. Watson included mixtures of more than one species. These problems led to misidentifications by other workers with the result that 16 of the 54 specific names in Eurythrips are placed in synonymy in the present study. Unfortunately the number of samples available is still inadequate in most species and further synonymy probably awaits discovery. Leaf-litter thrips appear to be restricted to the eastern half of the United States, few specimens having been seen from west of longitude 95. This probably represents a true faunal difference associated with differing soil conditions. However, even within the eastern U.S.A. most Thysanoptera have been collected close to the well- established areas of entomological work, Illinois, Washington D.C., Cornell and Gainesville. In the Neotropics most specimens have been collected in Brasil (Santa Catarina State), Panama and Trinidad. No specimens have been studied from western South America although there are a number of undescribed species from Columbia in the Senckenberg Museum, Frankfurt. ECOLOGY AND EVOLUTION IN LEAF-LITTER THYSANOPTERA The complexities of structural variation. - in leaf-litter thrips are probably determined by the habitat. Allometric growth patterns, as well as the production of winged or wingless morphs, are probably controlled by the amount of food ingested by developing larvae. However, the quantity and quality of food available will vary with the season and weather conditions because these thrips feed on fungal hyphae or the breakdown products of hyphal decay. The capacity to react to these fluctuations in the environment, both by varying the body structure and by varying the size of populations, is an important biological characteristic which is itself inherited by many species. Moreover, because of the relative uniformity of the leaf-litter habitat populations are extensive as well as locally dense, and the infrequency of winged individuals will result in a low rate of gene flow through these extensive populations (Mound & O'Neill, 1974). Localized variations within these populations will therefore not be eradicated quickly resulting in the observable pattern of variation within and between localities. Taxonomists have traditionally treated this type of variation in a simplistic manner, by naming as a new species each sample which differs structurally from other samples, be this one or many individuals. This approach does not have a valid philosophical basis AMERICAN LEAF-LITTER PHLAEOTHRIPIXAE 29 within the known and presumed biology of these insects as outlined above. The leaf-litter Thysanoptera appear to include a number of good examples of reticulate evolution, in which one or more characters have been suppressed genetically in any one line, that is not expressed phenotypically although still available within the genetic framework, but have reappeared in that line at a later stage. Dollos law appears to be quite inapplicable to this group of insects. GENERIC RELATIONSHIPS The genera related to Eurythrips are very difficult to assess. Hood (1954 : 25) refers to the problem of denning genera in this group and states that there are 'so many puzzling cross-resemblances that one wonders whether we have not in effect cut across the grain, rather than with it'. In fact Hood appears to have placed species in different genera in an arbitrary fashion and there are probably further specific synonymies to be found in which the species were described originally in different genera. Any future analysis of this group must be based on Hood's collection, including not only the types and long series of identified specimens but also the many hundred mounted specimens which Hood left unlabelled and un worked. The genera to be studied include Copiothrips, Phragmo- thrips, Pleurothrips and Orthothrips none of which can be distinguished satisfactorily from the three genera discussed in this paper. GENERA EXCLUDED FROM THE EURYTHRIPS COMPLEX The ten Neotropical species described in Porcothrips are here all transferred to Erkosothrips , Eurythrips or Terthrothrips. The genus Porcothrips Priesner (1951) is here regarded as a valid genus for one West African species, P. liberiensis (Priesner), which is known from a single specimen. Further material is needed of this species to determine the condition of the prothoracic sternites. The head is almost completely reticulate (Text-fig, i) but the setae on the basal antennal segments are all acute. The epimeral sutures are probably complete and both pairs of setae on the anterior margin of the pronotum are very small. The meso- notal mid-lateral setae are 50 /urn long but the pterothoracic ventrolateral setae are short and acute. The metanotum and pelta are reticulate, and the lateral areas of the abdominal tergites sculptured (Text-fig. 2). The sternal accessory setae are long and stout, there are at least eight setae on sternite eight on which segment the median marginal setae are exceptionally stout. The fore tibia has a fringe of long setae on the inner margin. The genus Zuluiella Jacot-Guillarmod (1939) was synonymized with Eurythrips by Stannard (1968). However, the type-species Z. distincta Jacot-Guillarmod from South Africa can be distinguished from all Eurythrips species by the presence of a maxillary bridge (Text-fig. 4), the well developed pronotal anteromarginal setae, and also by the large median setae on the tergites (Text-fig. 5). For these reasons Zuluiella must at present be accepted as a valid genus. The only other L. A. MOUND FIGS 1-7. Heads and right hand side of tergite IV. 1,2, Porcothrips liberiensis; 3, Erkoso- thrips subcalvus; 4, 5, Zuluiella distincta; 6, Eurythrips ampliventralis (cornutus holotype) ; 7, Eurythrips virginianus. species placed in this genus is Zuluiella antennata Jacot-Guillarmod (1940). This species was placed in Malacothrips by Hartwig (1952), and this seems to be a reasonable generic placement in view of the well developed pronotal anteromarginal setae and the very weakly developed praepectal plates. ACKNOWLEDGEMENTS This study was made possible firstly by the bequest to the British Museum (Natural History) (later abbreviated to BMNH in the text) of the collections of Floyd Andre including long series of North American Eurythrips (Mound, 1974^), and secondly by the courtesy of Kellie O'Neill and her husband Dr B. D. Burks in providing the author with the opportunity to work in Washington on the Hood collection for four weeks during 1975. The author is grateful to the following for AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 31 the loan of specimens in their custody (the abbreviations used later in the text are given in parentheses): Dr Paul Arnaud, California Academy of Sciences, San Francisco (CAS); Dr H. A. Denmark, Florida State Arthropod Collection, Gaines- ville (FSAC); Miss Kellie O'Neill, United States Department of Agriculture, c/o U.S. National Museum of Natural History, Washington (USNM); Dr L. J. Stannard, Illinois Natural History Survey, Urbana (INHS); Dr R. zur Strassen, Senckenberg Museum, Frankfurt-am-Main (SMF). The text-figures were prepared by my colleague Mrs J. M. Palmer. CHECK LIST OF THE SPECIES DISCUSSED IN THIS PAPER ERKOSOTHRIPS Stannard, 1955 bucca Hood, 1957 caelatoris (Hood, 1954) comb. n. claviger (Hood, 1941) floridensis Stannard, 1955 interior Stannard, 1955 reticulatus (Watson, 1934) sculpturatus (Hood, 1936) syn. n. silvarum (Hood, 1941) syn. n. silvaticus Hood, 1957 subcalvus (Hood, 1954) comb. n. EURYTHRIPS Hinds, 1902 alarius Hood, 1957 ampliventralis Hinds, 1902 amplus Hood, 1934 syn. n. conjunctus Hood, 1934 syn. n. connatus Hood, 1938 syn. n. cornutus Moulton, 1929 syn. n. disjunctus Hood, 1941 syn. n. batesi (Watson, 1935) bifasciatus (Hood, 1954) comb. n. bisetosus (Hood, 1954) comb. n. citricollis Hood, 1941 citricornis (Hood, 1954) comb. n. occipitalis Hood, 1957 syn. n. conformis Hood, 1957 costalimai Hood, 1950 cruralis Hood, 1957 dissirnilis Hood, 1938 elongatus Hood, 1957 forticauda (Hood, 1954) comb. n. forticornis Hood, 1939 fuscipennis Moulton, 1929 genarum Hood, 1957 constricttis Stannard, 1958 syn. n. hemimeres Hood, 1957 hindsi Morgan, 1913 eddeyi (Watson, 1935) s y n - n hookae Hood, 1933 longilabris Watson, 1921 hard Hood, 1925 syn. n. 32 L. A. MOUND modestus (Bagnall, 1917) cinctus Hood, 1927 syn. n. varius Moulton, 1929 syn. n. xanthozonus Hood, 1957 syn. n. musivi Hood, 1957 nigriceps Hood, 1957 nigricornis Hood, 1960 osborni Hinds, 1902 macrops Hood, 1925 syn. n. setiger Stannard, 1958 syn. n. peccans Hood, 1957 pettiti Hood, 1941 pusillus Hood, 1957 setosus Hood, 1950 simplex (Hood, 1954) comb. n. striolatus Hood, 1957 subflavus Hood, 1950 tar salts Hood, 1925 montanus (Watson, 1933) morulops Hood, 1950 syn. n. trifasciatus (Hood, 1954) comb. n. tristis Hood, 1941 utnbrisetis Hood, 1933 virginianus Hood, 1952 watsoni Hood, 1941 TERTHROTHRIPS Karny, 1925 balteatus Hood, 1957 : I 5 2 bicinctus Hood, 1954 : 34 brunneus Hood, 1957 : 148-149 bucculentus Hood, 1957 : I 45~ I 46 bullifer Hood, 1957 : I 49~ I 5 carens Hood, 1957 : 147-148 clavivestis Hood, 1935 : 195-199 cochlearius Hood, 1954 : 27-28 consobrinus Hood, 1954 : 2 5~ 2 6 crassus Hood, 1954 : 28-29 fuscatus Hood, 1954 : 2 9~3 gracilicornis (Hood, 1925 : 59) gracilis Hood, 1954 : 2 9 hebes Hood, 1957 : I 5 2 ~ I 53 impolitus Hood, 1957 : I 53~ I 54 irretitus Hood, 1957 : I 5- I 5 I longulus Hood, 1954 : 26-27 luteolus Hood, 1957 : I 5 I-I 5 2 magnicauda Stannard, 1955 : 174 minor Hood, 1954 : 2 ^ peltatus Hood, 1957 : 146-147 percultus Hood, 1957 : I 54~ I 55 sanguinolentus (Bergroth, 1896 : 66) serratus Hood, 1954 : 3 I- 3 2 striaticeps Hood, 1954 : 26 unicinctus Hood, 1954 : 30-3 1 viduus Hood, 1957 : J 47 AMERICAN LEAF-LITTER PHL AEOTHRIPIN A.E 33 ERKOSOTHRIPS Stannard Erkosothrips Stannard, 1955 : 81-83. Type-species: E. interior Stannard, by original designation. This genus was erected for a group of species in which the head is conspicuously reticulate and some of the setae on the basal antennal segments have dilated apices (Text-figs 3, 34). Unfortunately, neither on these characters nor on any others can Erkosothrips be satisfactorily distinguished from either Eurythrips or Copiothrips. The Neotropical species described in Porcothrips are transferred in this paper to Eurythrips, Erkosothrips or Terthrothrips, but the species in Copiothrips have yet to be re-examined. Eurythrips virginianus has the dorsal surface of the head partially sculptured and the setae on antennal segments II and III are broad at the apex (Text-figs 26, 32). Typical Erkosothrips have the postocular setae rather short (Text-fig. 3), but in caelatoris these setae are much longer than in the type-species. Moreover the head of caelatoris is weakly sculptured and rather similar to Eurythrips trifasciatus (Text-fig. 23). In Erkosothrips bucca the post- ocular setae are nearly straight, not curved, although the head is densely reticulate; however, the antennal setae do not have expanded apices. Erkosothrips, as defined by Stannard (1957) is probably polyphyletic, the sculptured dorsal surface to the head having evolved independently in both the North American Eurythrips line as well as the South American Terthrothrips line. Macropterae are known for only two species in the genus, caelatoris and reticulatus. KEY TO SPECIES (* species not studied) Ventrolateral setae on metathorax acute at apex ....... 2 Ventrolateral setae on metathorax expanded at apex ...... 3 Antennal segments II-III with one or more pairs of expanded setae; segments IV-V with slender pedicel about twice as long as wide; <$ with narrow glandular area near anterior margin of sternite VIII [Florida to New York and Iowa] reticulatus (p. 35) Antennal segments II-III with all setae acute at apex; segments IV-V with pedicels scarcely as long as wide; <$ glandular area on sternite VIII circular with long accessory setae arising laterally [Brazil] ...... bucca (p. 34) Median dorsal setae of head usually expanded at apex ...... 4 Median dorsal setae of head acute at apex (Text-fig. 3) . . . . . . 6 Abdomen largely yellow; lateral setae near pelta expanded apically [Michigan, Illinois, Missouri] interior (p. 35) Abdomen brown; lateral setae near pelta acute at apex ..... 5 Postocular setae placed in line with lateral margin of eye; $ with narrow glandular area at anterior margin of sternite VIII; $ not known [Florida] . *floridensis (p. 35) Postocular setae placed mesad of lateral margin of eye; <$ with glandular area occupying most of sternite VIII; $ with setae B : and B 2 on tergite IX expanded at apex [New York; Michigan] claviger (p. 34) Major setae exceptionally short, pronotal posteroangular setae less than one-fifth as long as median length of pronotum; antennal segment IV with 2 sense cones (Text-fig. 34); setae B l and B z on tergite IX expanded at apex [Brazil]. subcalvus (p. 36) 34 L- A. MOUND Major setae normal, pronotal posteroangular setae more than one-third as long as median length of pronotum; antennal segment IV with 4 sense cones; setae 5 t on tergite IX expanded at apex, but B z acute ....... 7 7 Body largely yellowish brown, tube dark brown ; median area of tergites polygonally reticulate [Brazil] .......... silvaticus (p. 36) Body brown, legs yellowish brown ; median area of tergites with transversely elongate reticulations [Brazil] ......... caelatoris (p. 34) Erkosothrips bucca Hood Erkosothrips bucca Hood, 1957 : 167. Holotype $, BRAZIL (USNM) [examined]. This micropterous species is known only from the type-series of six females and three males. SPECIMENS STUDIED. BRAZIL: Santa Caterina State, Nova Teutonia, holotype $ with 3 $, 2 ^paratypes under fallen leaves, iv. 1954 (F. Plaumann) (USNM). Erkosothrips caelatoris (Hood) comb. n. Porcothrips (?) caelatoris Hood, 1954 : 34. Holotype $, BRAZIL (USNM) [examined]. This species is intermediate between Erkosothrips, Copiothrips and Eurythrips. Although described from a single micropterous female there is a series of specimens in the Hood collection including macropterous females as welj. as micropterae of both sexes. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ microptera, viii. 1952 (F. Plaumann), also 40 specimens (^ $ and 9 mac.) from same locality on various dates 1953-4 (USNM). Erkosothrips claviger (Hood) Eurythrips claviger Hood, 1941 : 166-169. Holotype $, U.S.A.: New York (USNM) [examined]. Erkosothrips claviger (Hood) Stannard, 1955 : 83-84. This micropterous species was described from six females and 15 males collected in New York, but Stannard records a further nine specimens from Michigan and suggests the species is northern in its distribution. SPECIMENS STUDIED. U.S.A.: New York, Oswegatchie, holotype $ on dead grass and debris, 26. viii. 1940 (J. D. Hood), together with 3 $, 12 ^ paratypes bearing similar data (USNM). AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 35 Erkosothrips floridensis Stannard Erkosothrips floridensis Stannard, 1955 : 84. Holotype $, U.S.A. : Florida (INHS) [not examined]. This species was described from two males collected in leaf mould in the Ever- glades National Park, Florida. These specimens have not been studied and the species is placed in the key above from the original description and illustration. The female is unknown, and the possibility that the two original males are aberrant specimens of reticulatus needs further investigation. Erkosothrips interior Stannard Erkosothrips interior Stannard, 1955 : 84-85. Holotype $, U.S.A.: Illinois (INHS) [not examined]. This species was described from 17 $, 10 $ collected in Illinois and one female collected in Michigan. Subsequently Stannard (1968 : 409) recorded the species from Missouri. Although the abdomen is yellow, interior is similar in structure to both claviger and floridensis. SPECIMENS STUDIED. U.S.A.: Illinois, Allerton, i $, i ^ paratypes in woodland litter, 5.^.1940; I., Monticello, i $ in woodland cover, 28. ix. 1938 (Farrer) (USNM). Erkosothrips reticulatus (Watson) Glyptothrips reticulatus Watson, 1934:45-46. LECTOTYPE Q, U.S.A.: Florida (FSAC), here designated [examined]. Eurythrips sculpturatus Hood, 1936 : 5-9. Holotype $, U.S.A.: Louisiana (USNM) [examined]. Syn. n. Eurythrips silvarum Hood, 1941 : 163-166. Holotype $, U.S.A.: New York (USNM) [examined]. Syn. n. Erkosothrips reticulatus (Watson) Stannard, 1955 : 85; 1968 : 409-410. Stannard (1955) distinguished sculpturatus from silvarum on the degree of swelling of the cheeks and the position of the postocular setae, although subsequently he stated (1968 : 410) that these characters could not be used to distinguish between these species satisfactorily. Usually the silvarum head shape is found in apterae and the sculpturatus head shape in micropterae, and both of these forms are repre- sented in the syntype series of reticulatus in the Hood collection. Stannard did not place reticulatus in his key to species (1955) because the 'holotype' referred to by Hood (1941 : 163) was not available for study and Watson's original series evidently contained more than one species. However, later, Stannard (1968 : 410) designated a lectotype for reticulatus, although as neither the locality (Gainesville) nor the collector (Bates) of this specimen are mentioned in the original description this designation is invalid. The 'holotype' referred to by Hood apparently remained in Hood's collection until 1975 when it was studied by the present author in Washington. This specimen, which is here designated lectotype, has the head 36 L. A. MOUND shaped like sculpturatus with the postocular setae arising mesad of the external eye margin. However the cheek setae are not expanded apically, the B x setae on the ninth abdominal tergite are abruptly rounded, not broadly expanded apically, and the specimen is apterous, not micropterous. The lack of correlation between these various characters, both in this specimen and in several of the others listed below, has led the present author to regard all the material as conspecific. The different forms have been collected together both in Florida and Virginia. Stannard (1968) refers to a macropterous female from North Carolina. MATERIAL STUDIED. U.S.A.: Florida [Levy County], Gulf Hammock, lectotype 9 of reticulatus, 'Type 9' on slide with I <$ from dead leaves of sweet gum, oak etc. on ground (A. N. Tissot) (FSAC); F., Petersburg, 'type ^' of reticulatus, dead leaves, mostly oak, 3.ix.i933 (/. W. Hea] (FSAC); F., Gainesville, i 9 paralectotype of reticulatus, 17.1.1932; F., Hawthorne, i 9 paralectotype of reticulatus, 1.^.1932 (USNM); F., Mariana, i $, i ^ in moss, 3.1^.1933 (BMNH); Louisiana, Tallulah, holotype 9 of sculpturatus with 2 9, 2 ^ paratypes in humus, 11^.1934 (]. W. Folsom) (USNM); Georgia, Abbeville, 3 <$, iii. 1950 (USNM); Virginia, Falls Church, 6 $, 2 <$ in moss, 1943-45 (F. Andre}; V., Mt Vernon, 4 9, i ^ in sedge, 1940-44 (F. Andre] (BMNH); V., Lexington, i $, 2 ^, xi. 1940 (USNM); Maryland, Libertytown, i $, i ^ in moss,' 1944 (F. Andre] (BMNH); Iowa, White, i 9, 2 ^ in moss, I2.xii.i935 (F. Andre); L, Page County, i 9 m dead leaves, xii. 1935 (F. Andre) (BMNH); New York, Enfield Glen State Park, holotype 9 of silvarum on fallen pine needles, 15. ix. 1940 (F. R. Nevin) (USNM, with 30 9, 17 c? from type-locality, i 9 paratype BMNH). Erkosothrips silvaticus Hood Erkosothrips silvaticus Hood, 1957 : 166-167. Holotype 9, BRAZIL (USNM) [examined]. This micropterous species is known only from the type-series of 23 females and six males. This species resembles several species placed at present in Copiothrips. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype 9 under fallen leaves, vi. 1954 (F. Plaumann] (USNM); 20 9, 4 c? paratypes with similar data except for collection dates (USNM). Erkosothrips subcalvus (Hood) comb. n. (Text-figs 3, 34) Porcothrips subcalvus Hood, 1954 : 33~34- Holotype 9, BRAZIL (USNM) [examined]. Although this micropterous species was described in Porcothrips it is similar to several species in Copiothrips and Erkosothrips, particularly to E. reticulatus. The ventrolateral setae on the metathorax have expanded apices although they are scarcely 25 /Am long. The original series comprised seven females. AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 37 SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ with 5 $ paratypes, viii. 1952 (F. Plaumanri) (USNM); also about 70 specimens of both sexes with similar data to holotype, collected 1953-55 (USNM; i $ in BMNH). EURYTHRIPS Hinds Eurythrips Hinds, 1902 : 202. Type-species: E. ampliventralis Hinds, by original designation. The genus Eurythrips has been denned by Stannard (1957) although at present it cannot be satisfactorily distinguished from Erkosothrips, Terthrothrips, Copiothrips, Orthothrips, Phragmothrips and Pleurothrips. Moreover most of the Neotropical species described in Porcothrips are here placed in Eurythrips. None of the species in Eurythrips has reticulation on the head close to the postocular setae, although in a few species the median area of the head between these setae is weakly reticulate (Text-fig. 23). In contrast, the species in Erkosothrips have the dorsal surface of the head entirely reticulate, and moreover, the postocular setae are usually short (Text-fig. 3) and some of the setae on the basal antennal segments have blunt or expanded apices. All the species in this complex of genera from the New World have large praepectal plates on the ventral surface of the prothorax, and the pronotal anteromarginal setae are always short although the anteroangulars are usually long (Text-figs 27, 28). Certain species of Malacothrips are very similar to Eurythrips species, notably Malacothrips roycei Hood from Florida and M. antennatus (Jacot-Guillarmod) from South Africa. However, both of these species have well developed pronotal anteromarginal setae and the praepectal plates are either absent or weakly developed. The species of Eurythrips exhibit a variety of patterns of structural variation and also of geographical distribution. As interpreted here, ampliventralis is a widespread species extending from Iowa in the north, through Panama and Trinidad to southern Brazil. No structural variation has been observed to be correlated with geographical distribution in ampliventralis although four nominal species from the Neotropics, each with a foretarsal tooth, may be geographical variants of this species. However, at several sites throughout its range ampli- ventralis has been found to vary in the width of the pedicel of the terminal antennal segments (Text-figs 35, 36), and moreover the number of antennal sense cones is not constant. Another species with an extensive range is batesi. This is found from Maryland to Mexico and Trinidad, but is relatively constant in structure. In contrast modestus is a widespread and variable species from the Carribean area to Brazil, and it is possible that nigricornis is the same species. Moreover longilabris from Texas and Florida is also closely related, although this species is itself probably only an extreme southern variant of the North American species hindsi. This type of 'Rassenkreise' is also known in the leaf-litter genus Allothrips in North America (Mound, 19720) . Unfortunately half of the species of Eurythrips are still known from single samples or even single specimens. Many of these will probably be placed in synonymy when their variation is worked out. L. A. MOUND 8 FIGS 8-17. Heads of Eurythrips species. 8-10, ampliventralis (8, disjunctus paratype; 9, cornutus paratype); n, bisetosus; 12, citricornis; 13, dissimilis; 14, elongatus; 15, forti- cauda; 16, hemimeres; 17, hindsi. AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 39 FIGS 18-26. Heads of Eurythrips species. 18, hookae; 19, longilabris; 20, modestus (cinctus holotype); 21, osborni; 22, tarsalis; 23, trifasciatus; 24, simplex; 25, tristis; 26, virginianus. 4 o L. A. MOUND Most of the characters used in the key to species below are relatively simple. However, the variation in a character is in most species directly related to the number of samples studied, and so the key will be least effective in areas from which samples are at present not available, e.g. South Carolina, Georgia and Arkansas. The form of the setal apices which is used in the key refers to specimens mounted in Balsam (Mound & Pitkin, 1972). Specimens mounted in chloral hydrate and gum arabic mountants frequently have the setal apices deformed. These mountants can cause a weakly expanded setal apex to collapse and appear perfectly normal but acute. The author has used the botanical term 'acuminate' to indicate an acute seta with the apex drawn out evenly to a long fine point. The term 'pointed' which has frequently been used previously covers a variety of setal structure. The author distinguishes as 'softly pointed' or 'blunt' those setal apices which, when examined in Canada Balsam with phase contrast microscopy, can be seen to be translucent and minutely flattened at the apex. The most common species with setae of this type is ampliventralis in which the B setae on the ninth tergite are 'pointed' (teste Stannard, 1968 : 414) but are certainly not 'acuminate' in the sense of couplet 8 of the key below. KEY TO SPECIES Head reticulate between the postocular setae (Text-fig. 23) ; pelta with distinct lateral wings .......... .2 Head not reticulate between the postocular setae, rarely with transverse lines of sculpture (Text-figs 8-17); pelta usually different ...... 5 Pronotal anteroangular setae about 15 ,um long, less than 0-25 as long as postero- angular setae; antennal segment IV with four sense cones (Text-fig. 29). Dorsal surface of head with equiangular polygons (Text-fig. 23) ; $ glandular area occupies most of sternite VIII [Brazil] .... trifasciatus (p. Go) Pronotal anteroangular setae almost as long as posteroangular setae, antennal segment IV with three sense cones ........ 3 Setae B l on tergite IX scarcely wider at apex than B 2 , softly pointed; head with transversely elongate polygons medially [Brazil] .... striolatus (p. 59} Setae B v on tergite IX sharply expanded at apex, B 2 acute at apex; head with equiangular reticulations .......... 4 Head dark brown, darker than pronotum [Brazil] .... musivi (p. 56) Head yellowish brown, lighter than pronotum [Brazil] . . bifasciatus (p. 48) Head exceptionally long, more than 1-8 times as long as width across eyes (Text- fig. 14). Head, metathorax, abdominal segments VIII-X, antennal segments I-II and VI-VIII brown, rest of body and legs yellow; $ without glandular area on sternite VIII [Brazil] elongatus (p. 51) Head variable in shape, always less than 1-7 times as long as width across eyes . 6 Antennal segments II-III bearing one or more setae with expanded apices (Text- figs 32, 33)- $ sternite VIII with narrow glandular area at anterior margin, sometimes with one or more accessory setae between glandular area and posteromarginal setae (Text-fig. 44); setae B l and B 2 on tergite IX expanded at apex ... 7 Antennal segments II-III with all setae acute ....... 8 Pterothoracic ventrolateral setae with expanded apices; dorsal setae on antennal AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 41 segment II strongly expanded, segment IV with four sense cones (Text-fig. 33) ; pronotal epimeral sutures usually complete; pelta and median area of tergites very weakly sculptured [Brazil] ...... forticauda (p. 51) - Pterothoracic ventrolateral setae small with acute apices; dorsal seta on antennal segment II blunt, segment IV with three sense cones (Text-fig. 32); pronotal epimeral sutures not complete; pelta and median area of tergites sculptured [eastern U.S.A.] ......... virginianus (p. 61) 8 Setae B l on tergite IX acuminate at apex; dorsolateral setae on tergite VII fre- quently acuminate ........... 9 - Setae B^ on tergite IX never acuminate, softly pointed with apex translucent to blunt or expanded; dorsal posterolateral setae on tergite VII blunt or expanded at apex ............. 21 9 Body, antennae and legs clear yellow, head, tube and antennal segments VII-VIII weakly shaded greyish brown [Brazil] ...... subflavus (p. 59) - Body colour different, more extensively brown . . . . . . . 10 10 Abdominal segment II yellow in contrast to segments VII-IX which are brown . n - Abdominal segments more or less uniformly brown, segments IV-VI sometimes paler than II-III and VII-IX . 13 11 Pronotum and abdominal segments I-VI, also legs, yellow; head and tube brown, pterothorax light brown, antennae brown, segment II paler distally. Pronotal epimeral sutures not complete; antennal segment III with three sense cones [Florida] .......... citricollis (p. 49) - Pronotum and abdominal segments IV-IX brown . . . . . . 12 12 Antennal segment III with three sense cones; hind femora yellow [Brazil]. nigricornis (p. 56) - Antennal segment III with two sense cones; hind femora brown [West Indies, Panama, Brazil] modestus (p. 55) 13 Abdominal segments IV-VI I yellow with brown shading medially; antennae brown except pedicel of III; tube yellow in distal half; antennal segments III and IV with three sense cones; setae B on tergite IX acuminate and longer than tube, on tergite VII expanded at apex; postocular and pronotal setae expanded at apex, pretarsal tooth less than 0-25 times as long as tarsal width [Panama] hookae (p. 54) - Not this combination of characters ..... 14 14 Fore tarsal tooth small, less than 0-25 times as long as tarsal width in both sexes; pronotal anteroangular setae usually 0-5 times as long as posteroangular setae . 15 - Fore tarsal tooth large in both sexes, usually more than 0-5 times as long as tarsal width; anteroangular setae frequently less than 0-25 times as long as postero- angular setae ....... 18 15 Postocular setae clearly expanded at apex. Head with median dorsal setae and postocellar setae usually small but sometimes elongate (Text-fig. 25) ; $ glandular area occupies most of sternite VIII [eastern U.S.A.] tristis (p. 60) - Post ocular setae acuminate or blunt . . ..... 16 1 6 Antennal segment IV with four sense cones [Brazil] . . pusillus (p. 58) - Antennal segment IV with three sense cones . 1 7 17 Antennal segments IV-VI sub-glucose, sharply constricted to basal neck and often irregular in profile, segment IV less than i -5 times as long as wide (Text-figs 37, 38) [eastern U.S.A.] .... Mndsi (p. 53) - Antennal segments not sharply constricted at base, segment IV more than i -5 times as long as wide (Text-fig. 41) [Florida, Texas] . longilabris (p. 55) 1 8 Postocellar setae usually longer than ocellar triangle (Text-fig. 22). Antennal segment III with two or three sense cones, IV with three sense cones 42 L. A. MOUND (Text-figs 30, 31); pronotal epimeral sutures usually not complete [eastern U.S.A., Brazil] tarsalis (p. 59) Postocellar setae shorter than ocellar triangle . . . . . . . 19 19 Pronotal epimeral sutures not complete. Antennal segment IV with four sense cones [Brazil] . . . setosus (p. 58) Pronotal epimeral sutures complete ......... 20 20 Tube dark brown; setae B l on tergite IX scarcely 0-9 times as long as tube; antennal segment IV with three sense cones (Text-fig. 40) ; , 3 < micropterae in Hood's box of ampliventralis (USNM); California, 3 $, i c? (F. Andre) (BMNH); New Mexico, Espanola, i $, iii. 1939 (BMNH). MEXICO: Tampa, i $, vi. 1943 (BMNH). PANAMA: Barro Colorado L, holotype $ macroptera of conjunctus with i $ macroptera, 2 ?, 5 <$ micropterae (all paratypes) (USNM, i $ BMNH). CUBA: Baragua, holotype $ macroptera of cornutus, iv. 1927 (CAS); $ paratype macroptera with identical data (BMNH). TRINIDAD: St Joseph, 48 L. A. MOUND holotype $ macroptera of amplus, iii. 1918 (C. B. Williams] (USNM); Curepe, 3 $ micropterae, xi. 1970 (L. A. Mound) (BMNH). BRAZIL: Santa Catarina State, 3 $ macropterae, v. 1949 (F. Plaumann] (USNM). Eurythrips batesi (Watson) (Text-figs 28, 47) Glyptothrips batesi Watson, 1935 : 56-57. LECTOTYPE $>, U.S.A.: Florida (FSAC), here designated [examined]. Eurythrips batesi (Watson) Hood, 1941 : 163. This species was described from 'several hundred apterous [sic] females and a single winged one, and many males'. Hood (1941 : 163) in a footnote referred to a slide which he had labelled 'holotype', and it is this specimen which is here designated lectotype. Stannard (1968 : 416) was unable to locate this specimen in the Watson collection because, unknown to him, it had remained on loan in the Hood collection where the present author studied it in 1975. Watson included specimens in his type-series from Florida, Georgia and Tennessee, but the original series comprised more than one species. Stannard records batesi from most of the states between southern Illinois, Maryland, Florida and Texas, and in addition records a possible variant from New Mexico, Mexico and Jamaica. As indicated below the present author has also collected batesi in Trinidad. In contrast to Watson's description neither Stannard nor the present author have seen macropte- rae, and the males and females are micropterae not apterae. SPECIMENS STUDIED. U.S.A.: Florida, Alachua Cty, Devils Mill Hopper, lectotype $ from dead leaves, 24.xii.i933 (J.R.W.-B.F.) (FSAC); F., Quincy, i , x. 1930; F., Gainesville, i $, i. 1932; F., Ocala Forest, 3 $, xi. 1933; F., Jena, i <, ii. 1934, all apparently from Watson's syntype-series (USNM); F., Alachua Cty, 5 , 3 , ix. 1944; V., Arlington, 4 $>, i $, x. 1939 (BMNH); V., Natural Bridge, 6 , 7 <$, xi. 1940 (USNM); Maryland, Beltsville, 2 $, 2 < in Sphagnum, i. 1945 (BMNH); Delaware, Newark, 2 $, 2 <, iv. 1951 (BMNH). MEXICO: i $ with no data, ex Floyd Andre (BMNH). TRINIDAD: various localities near Curepe and Tunapuna, 45 $, 30 $ in leaf litter, 1970-72 (L. A. Mound 6- B. R. Pitkin) (BMNH). Eurythrips bifasciatus (Hood) comb. n. Porcothrips (?) bifasciatus Hood, 1954 : 37-38. Holotype $, BRAZIL (USNM) [examined]. This species is similar to Eurythrips musivi. It is known only from a single specimen. AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 49 SPECIMEN STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ macroptera, viii. 1952 (F. Plaumann) (USNM). Eurythrips bisetosus (Hood) comb. n. (Text-fig, n) Porcothrips (?) bisetosus Hood, 1954 ' 35~3 6 - Holotype $, BRAZIL (USNM) [examined]. This species was described from eight males and eight females, all micropterae. Although described in Porcothrips these specimens are similar to the micropterae of Eurythrips hemimeres. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ microptera with 7 <$, 6 $ paratypes, viii. 1952 (F. Plaumann), also approximately 60 specimens of both sexes with similar data collected 1953-54 (USNM). Eurythrips citricollis Hood Eurythrips citricollis Hood, 1941 : 240-243. LECTOTYPE $ macroptera, U.S.A.: Florida (USNM), here designated [examined]. The type-series of this species comprised three rriacropterous and eleven micropterous females. Although a 'paratype' was referred to in the original description no holotype was designated. The lectotype designated here is a female macroptera labelled 'holotype' by Hood. The male is still unknown. SPECIMENS STUDIED. U.S.A.: Florida, Winter Park, lectotype $ macroptera under fallen pine needles, n.iii.i94i (M. J. West/all), with 2 $ macropterae, 10 $ micropterae bearing identical data (USNM). Eurythrips citricornis (Hood) comb. n. (Text-figs 12, 43) Porcothrips (?) citricornis Hood, 1954 : 36-37. Holotype $, BRAZIL (USNM) [examined]. Eurythrips occipitalis Hood, 1957 : 139-140. Holotype $, BRAZIL (USNM) [examined]. Syn. n. Hood described each of these two species from two micropterous females. The head is partially sculptured (Text-fig. 12), but the setae on the basal antennal segments are acute, and unlike several Erkosothrips and Copiothrips species the pterothoracic ventrolateral setae are also acute. Hood stated that occipitalis was 'easily distinguished by the large, knobbed, occipital setae which are almost equal to the postoculars'. This, however, is true only of the holotype because one of the 50 L. A. MOUND occipital setae on the paratype is small and blunt. The occipital setae of Eurythrips tristis are also shown below to be variable. The specimens of citricornis listed below all have the occipital setae short and acute, and moreover, the colour of the tibiae and the sculpture of the head and pelta are variable. The type- specimens of occipitalis are therefore interpreted here as representing a dark form of citricornis. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ of citricornis with i $ paratype micropterae, viii. 1952 (F. Plaumann); 12 9, i $ with similar data 1953-55; holotype $ microptera of occipitalis with similar data, vii. 1955 and i $ paratype, viii. 1955 (USNM). Eurythrips conformis Hood Eurythrips conformis Hood, 1957 : 136-137. Holotype ^, BRAZIL (USNM) [examined]. This species was described from nine micropterous females. It appears to be closely related to costalimai which is known from a single macropterous female. The variation in the group of species closely related to ampliventralis requires further study. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype 9 microptera from fallen leaves, ix. 1954 (F. Plaumann), with 6 $ micropterous paratypes bearing similar data but collected between vii. 1953 and x. 1955 (USNM). Eurythrips costalimai Hood Eurythrips costalimai Hood, 1950 : 99-102. Holotype $, BRAZIL (USNM) [examined]. This species is very similar to conformis, differing mainly in the size of the fore tarsal tooth and the shorter setae on the abdominal tergites. The importance of these differences is difficult to assess because conformis is known only from micropterae and costalimai from a single macroptera. Both characters are known to be related to the degree of wing development in certain groups of Thysanoptera. The unique holotype of fuscipennis also keys out with costalimai and these two nominal species require further study and comparison with other members of the ampliventralis-group. SPECIMEN STUDIED. BRAZIL: Jacarepagua [Rio de Janeiro], holotype $ macroptera on Andropogon condensatus, 18^.1948 (Hood) (USNM). Eurythrips cruralis Hood Eurythrips cruralis Hood, 1957 : 140. Holotype 9, BRAZIL (USNM) [examined]. The three female micropterae on which this species was based can be distinguished AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 51 from the other members of the ampliventralis-group by the unusual colour of the legs and tube. SPECIMEN STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ microptera from fallen leaves, viii. 1952 (F. Plaumann), with I $ paratype bearing similar data except viii. 1955 (USNM). Eurythrips dissimilis Hood (Text-figs 13, 40, 49) Eurythrips dissimilis Hood, 1938 : 365-366. Holotype $, U.S.A.: Florida (USNM) [examined]. This species was described originally from one micropterous female. Both dissimilis and forticornis appear to be related to the hindsi-group but they are unusual in having the epimeral sutures complete. SPECIMENS STUDIED. U.S.A.: Florida, Sebring, holotype $ microptera swept from grass, 7.^.1938 (J. W. H. Rehn) (USNM); Texas, Palacios, 26 $, 20 $ on grass, i.iv.i939 (USNM); Virginia, Mt Vernon, I < from Andropogon, ii. 1940 (F. Andre) (BMNH); New Jersey, Cape May, i $, x. 1949 (USNM). MEXICO: Axtla, i 9, v. 1939 (USNM). CUBA: Ceiba Mocha, Matunzas, i <$, vii. 1940 (USNM). Eurythrips elongatus Hood (Text-figs 14, 42) Eurythrips elongatus Hood, 1957 : 140-141. Holotype $, BRAZIL (USNM) [examined]. This species was described from two females and one male, all micropterae. These differ from other members of the genus in having a relatively long, slender head. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ microptera under fallen leaves, vii. 1953 (F. Plaumann), also i $, i $ micropterae with similar data except ii. 1954 (USNM). Eurythrips forticauda (Hood) comb. n. (Text-figs 15, 33, 44) Porcothrips (?) forticauda Hood, 1954 : 34-35. Holotype $, BRAZIL (USNM) [examined]. This is one of the species which appears to be intermediate between Eurythrips and Erkosothrips. The head is weakly reticulate but the setae on the basal 52 L. A. MOUND antennal segments and the pterothoracic ventrolateral setae have expanded apices (Text-fig. 43). Moreover the meso- and metathoracic femora also bear a single stout seta with an expanded apex as in osborni. The type-series consisted of 26 female and seven male micropterae. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ microptera with 22 $, 6 <$ paratype micropterae, vii. 1952 (F. Plaumanri), also more than 70 further specimens with similar data, 1953-54 (USNM, i $>, i < BMNH). Eurythrips forticornis Hood Eurythrips forticornis Hood, 1939 : 593-594. Holotype $, U.S.A. : Alabama (USNM) [examined] . This species is similar to dissimilis in the form of the head and smooth pronotum. However, the setae on the ninth abdominal tergite are longer, and the tube coloured differently. The species was based on a single female. SPECIMENS STUDIED. U.S.A.: Alabama, Mobile, holotype 9 microptera on dead grass, I7.xii.i938 (Hood); Virginia, New Market, I $ microptera on grass, 5.1.1941 (L. C. Pettit] (USNM). Eurythrips fuscipennis Moulton Eurythrips fuscipennis Moulton, 1929 : 65-66. Holotype $, CUBA (CAS) [examined]. This species is referred to under costalimai with which species it needs to be further compared. SPECIMEN STUDIED. CUBA: Baragua, holotype $ macroptera from unknown host plant, 15.^.1927 (L. C. Scaramuzza) (CAS). Eurythrips genarutn Hood (Text-fig. 46) Eurythrips genarum Hood, 1957 : 54-55- Holotype $, U.S.A.: Virginia (USNM) [examined]. Eurythrips constrictus Stannard, 1958 1272-273. Holotype $, U.S.A.: Kentucky (INHS) [not examined]. Syn. n. The original description of this species clearly states 'two sense cones on outer surface of segment three of antennae'. However, this statement is not true of the holotype, nor of the five female and one male paratypes which have been studied. Antennal segment III of all the specimens listed below bears only two sense cones, one on the inner and one on the outer surface. As a result constrictus cannot be distinguished from genarum. This species is very similar to ampliventralis AMERICAN LEAF-LITTER PHLAEOTHRIPINAE 53 and can only be distinguished by means of the poor characters indicated in the key. The base of antennal segment VIII is moderately thickened and well within the range of variation of ampliventralis , and the fore tarsal tooth is not apparent. It seems likely that genarum will eventually be shown to be a local variant of ampliventralis. The type-series of genarum consisted of eight female and one male micropterae from Virginia, and the type-series of constrictus comprised one female with six male micropterae from Kentucky and 21 female with 15 male micropterae from Illinois. SPECIMENS STUDIED. U.S.A.: Virginia, Fairfax Cty, holotype $ microptera of genarum on Andropogon with 5 $, i < paratypes, 5.1.1941 (L. C. Pettit) (USNM); V., Mt Vernon, i $, i < on Andropogon, iQ.xii.i944 (F. Andre); V., Falls Church, 2 <$ on sedges, 31. x. 1943 (F. Andre] (BMNH); Illinois, Zenith, i <$ determined as constrictus by L. J. Stannard, i8.iii.i966 (INKS). Eurythrips hemimeres Hood (Text-fig. 16) Eurythrips hemimeres Hood, 1957 : 142. Holotype $, BRAZIL (USNM) [examined]. Judging from the type-series of 155 specimens this was the most abundant of the many species of Eurythrips collected by F. Plaumann in southern Brazil. It appears to be a member of the batesi-group of species, and has expanded apices to the setae on the ninth tergite but a large and complete glandular area on sternite eight of the male. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype $ macroptera from fallen leaves, iv. 1954 (F. Plaumann), also 149 specimens including $ <$ micropterae with similar data except collection dates, viii. 1952 to x. 1955 (USNM, i $, i J micropterae BMNH). Eurythrips hindsi Morgan (Text-figs 17, 37, 38) Eurythrips hindsi Morgan, 1913 : 27-28. Holotype $, U.S.A.: Tennessee (USNM) [examined]. Glyptothrips eddeyi Watson, 1935 : 57-58. LECTOTYPE $, U.S.A. : South Carolina (FSAC), here designated [examined]. Syn. n. This species was described from a single micropterous female, although specimens have now been studied from most of the states between Illinois, Maryland, South Carolina and Arkansas. Over most of this area specimens of hindsi have the median antennal segments exceptionally short and broad (Text-fig. 37), but some specimens from Tennessee have longer antennae (Text-fig. 38). The species known as longilabris (=harti) is closely related and may be merely a southern variant, 54 L- A. MOUND but this species itself appears to grade into the modestus-group from the Caribbean. The hindsi-group also includes tristis and tarsalis. The postocellar setae are usually short in both hindsi and tristis, although these setae are almost always elongate in tarsalis. The postocular setae are acuminate in hindsi, expanded apically in tristis but softly pointed in tarsalis. Moreover these setae are acuminate in the longilabris holotype but softly pointed in the harti holotype although these two are treated here as one species. The syntypes of eddeyi, labelled paratypes by Watson, have been found to contain at least seven species. A lectotype is selected and listed below under specimens studied for the purposes of synonymy. The other syntypes are as follows. ampliventralis : i <$, broom sedge tops, Florida, Gainesville, ii. 1934 (USNM); i J, cabbage, no locality, ii. 1931 (FSAC). osborni: i <$, i 9, swamp grass, South Carolina, Clemson Coll., ii. 1931 (FSAC). pettiti: i $, pine needles, North Carolina, Ashville, x. 1934 (FSAC). tristis: i 9, broom sedge, South Carolina, Clemson Coll., ii. 1931 (FSAC). watsoni: 2 $, pine needles, North Carolina, Ashville, x. 1934 (USNM). Malacothrips roycei: 4 ^, i 9> swamp grass, South Carolina, Clemson Coll., ii. 1931 (FSAC). SPECIMENS STUDIED. U.S.A.: Tennessee, Clarkesville, holotype 9 microptera of hindsi in stool of broom sedge, ly.x.igio (W. E. Hinds) (USNM); T., Hamilton Cty, 5 <$, 7 9 micropterae, xi. i939-ii. 1941 (USNM, i 9 macroptera with similar data BMNH); Alabama, Escambia Cty, 5 9 micropterae, i 9 macroptera, 1936-7 (USNM); Arkansas, no locality, 5 9 micropterae in dead leaves (?F. Andre) (BMNH); Mississippi, Lincoln Cty, i 9, iii. 1938; M., McNeill, i ?, 3 5 $ micropterae, vii. 1933 (USNM). Eurythrips peccans Hood (Text-fig. 27) Eurythrips peccans Hood, 1957 : 135. Holotype $, BRAZIL (USNM) [examined]. This species was based on 19 female and one male micropterae. These resemble the macropterous female described as alarius but differ in colour and the number of sense cones on the fourth antennal segment. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype 9 microptera with 16 9, i $ paratype micropterae from fallen leaves, v. 1953-x. 1955 (F. Plaumann) (USNM). 5 8 L. A. MOUND Eurythrips pettiti Hood Eurythrips pettiti Hood, 1941 : 203-206. Holotype $, U.S.A.: Virginia (USNM) [examined]. This species may be distinguished from the other North American members of the genus by the expanded apices of the major setae on the ninth abdominal tergite and the yellow colour of the first two abdominal segments. The type-series comprised three micropterae, two females and one male, but Stannard (1968 : 420) refers to female macropterae and records the species from Illinois, Michigan, Pennsylvania, Missouri, Arkansas and North Carolina. SPECIMENS STUDIED. U.S.A.: Virginia, New Market, holotype 9 microptera on grasses with I $, I , 16 $ micropterae (F. Andre) (BMNH); V., Fairfax Cty, 14 $, i $ micropterae; V., Luray, i $ macroptera; V., Natural Bridge, 5 $ macropterae, 7 micropterae (USNM); New York; Hammondsport, 7 $, 14 <; N.Y., Oswegatchie, i $ macroptera; N.Y., St Lawrence, 4 $, i $ (USNM); Iowa, Ottumwa, 22 $, 14 $ (F. Andre) (BMNH); North Dakota, Northwood, 2 $ (USNM); Tennessee, Gatlinburg, i $, i ^ 'paratypes' of montanus from dead leaves on ground, I5.viii.ig32 (J.R.W.) (USNM); Texas, Blessing, i <$, iv. 1939 (USNM); Georgia, Metter, 2 $, iii. 1950 (USNM); North Carolina, Bent Creek, i $ bearing manuscript name, 17.^.1935 (Jacot) (USNM); Florida, Clearwater, 2 ?, i & xii. 1937 (USNM). BRAZIL: Sao Paulo State, Salesopolis, Boracea, holotype $ macroptera of morulops with 10 $, 12 <$ paratype micropterae bearing similar data, 6.vi.i948 (Hood) (USNM, i $ BMNH); Para State, Belem and Santarem, 2 ?, i micropterae, 1951 (USNM). Eurythrips trifasciatus (Hood) comb. n. (Text-figs 23, 29) Porcothrips (?) trifasciatus Hood, 1954 : 38-39. Holotype $, BRAZIL (USNM) [examined]. The head is weakly sculptured in this species, as in striolatus, but the antennae are strongly sculptured (Text-fig. 29). The major dorsal setae on the ninth abdominal tergite are softly pointed, not acuminate. The species was based originally on a single female microptera. SPECIMENS STUDIED. BRAZIL: Santa Catarina State, Nova Teutonia, holotype ? microptera, viii. 1952 (F. Plaumann); also 70 further specimens, including $ macropterae and 9 <$ micropterae, with similar data except dates of collection (USNM). Eurythrips tristis Hood (Text-figs 25, 39) Eurythrips tristis Hood, 1941 : 157-160. Holotype?, U.S.A.: New York (USNM) [examined]. The type-series of tristis comprised three female macropterae and nine female with four male 'apterae'. However, these apterae have very small wing lobes and should be referred to as micropterae despite the extreme reduction of the ocelli. This species is not included by Stannard (1968) in his key to the species AMERICAN LEAF-LITTER PHLAEOTHRIPIN AE 61 from Illinois, but specimens of tristis come down to 'osborni' in that key and specimens listed below from Illinois, Kentucky and Michigan were labelled as osborni by Dr Stannard. (E. setiger Stannard is given above as a synonym of osborni Hinds.) The record of osborni from Georgia (Beckham et al. 1971) is probably also a misidentification of tristis, although the two females listed below from Georgia have longer antennae than specimens from Virginia and moreover the apices of the dorsal setae on tergite nine are not quite acuminate. Several females from Virginia, both macropterae and micropterae, have either or both of the postocellar and median dorsal (occipital) setae on the head elongate. These specimens emphasise the relationship between tristis, tarsalis and hindsi. SPECIMENS STUDIED. U.S.A.: New York, Oswegatchie, holotype $ macroptera with 2 $ macropterae, 5 $, 4 <$ micropterae (paratypes) in dead grass, g.ix.ig/j-O (Hood}; N.Y., Apulia Station, I $ paratype microptera, n.x.i94o; N.Y., Warrensburg, 3 $ paratype micropterae, 12. x. 1940 (USNM); Illinois, Marshall, i <, v. 1949; I., Monticello, i 9, v. 1940 (INKS); Kentucky, Mammoth Cave, i ^, iv. 1950 (INKS); Michigan, George Res., i $, i. 1950 (INKS); Virginia, Falls Church, 16 9, 10 ^ micropterae, 1944-45 (F. Andre); V., Four Mile Run, i $ macroptera (F. Andre] (BMNH); V., Natural Bridge, I (CNC, Ottawa); Washington, Mt Rainier, Eagle Park, I9.vii.i922 (A. L. Melander], i $ (DEFW, St Paul); Colorado, Nederland, 300', 5.vii.i96i (/. G. Chillcott), i ). MATERIAL EXAMINED. Holotype <$ (dry-pinned), U.S.A.: California, Amador Co., Pioneer, 8. v. 1961 (0. W. Richards) (BMNH). Paratypes. U.S.A.: same data as holotype, i J, I $ (slide mounted). A further single female is very much larger than the type-series (wing-length 6-0 mm). This specimen also has unequal hind tibial spurs and a much longer head and so may be a second species. Its data are: U.S.A.: Oregon, Humbug Mts, Curry Co., 2o.vi.i939 (T. Aitken) (B. Brookman coll., in USNM, Washington (per E. F. Cook)). CANTHYLOSCELIDAE Rodendorf Corynoscelinae Enderlein, 1912 : 264. Type-genus: Corynoscelis Boheman, 1858 [junior homonym of Corynoscelis Burmeister, 1847]. Corynoscelidae Enderlein; Enderlein, 1936 : 56. Canthyloscelididae Rodendorf, 1951 : 64. Type-genus: Canthyloscelis Edwards, 1922. Corynoscelinae [Enderlein]; Toilet, 1959 : 144. [Proposed as new subfamily.] Hyperoscelidae Hardy & Nagatomi, 1960 : 263. Type-genus: Hyperoscelis Hardy & Nagatomi [replacement name for Corynoscelis Boheman]. Syn. n. Hyperoscelididae Hardy & Nagatomi; Rodendorf, 1964 : 16. NOMENCLATURE. The International Code of Zoological Nomenclature (1964), Article 23d(i), states that the oldest valid family-group name must be used. The homonymy of Corynoscelis Boheman makes the family name based on it invalid (Article 39). If the two families considered in this paper are regarded as one composite family, the valid family-group name would be Synneuridae Enderlein, 1936, as this name has priority. Where they are regarded as two separate families, as in the present work, the name Canthyloscelidae, proposed as Canthyloscelididae by Rodendorf (1951), predates Hyperoscelidae Hardy & Nagatomi (1960), and so is the valid name for the restricted family including Hyperoscelis and Canthyloscelis. Thus the family name Hyperoscelidae must be changed: for Canthyloscelidae if the family is restricted to Canthyloscelis and Hyperoscelis, or Synneuridae if Synneuron and Exiliscelis are included. I regard it as both unnecessary and undesirable to change the family-group name yet again for a group that, though small, has already been widely discussed outside the realm of basic taxonomy: in fields such as the evolution of Diptera (Rodendorf, 1964; Mamaev, 1968), zoogeography (Hennig, 1960; 1965), anatomy, ecology, physiology and development (Rodendorf, 1951; Hennig, 1954; Krivosheina & Mamaev, 1967; Mamaev & Semenova, 1969; Krivosheina, 1969), as well as important general faunal lists (e.g. Cook, 1965; 1967; Martinovsky, 1972) and key 78 A. M. HUTSON works (e.g. Seguy, 1951; Brues, Melander & Carpenter, 1954; Bei-Bienko, 1969; Colless & McAlpine, 1970; Hennig, 1973). It is to be hoped that this latest change 'in the interests of stability' is the last of such changes. DIAGNOSIS. As in key to families (p. 70). DISTRIBUTION. South America, New Zealand and Palaearctic. KEY TO THE GENERA i Median fork complete. Claws simple. Antennae only about two-thirds length of thorax. (Palaearctic) . . . . . . HYPEROSCEL1S (p. 78) Median fork with M 2 interrupted at base. Claws with a comb-like or toothed basal enlargement. Antennae as long as head and thorax together. (Neotropical and New Zealand) CANTHYLOSCELIS (p. 83) HYPEROSCELIS Hardy & Nagatomi (Text-figs 13-19) Corynoscelis Boheman, 1858 : 56. Type-species: Corynoscelis eximia Boheman, 1858, by monotypy. [Preoccupied by Corynoscelis Burmeister, 1847.] Spiloptera Zetterstedt, 1860 : 6487. Type-species: Spiloptera arctica Zetterstedt, 1860, by monotypy. [Proposed in synonymy in footnote.] ? Eucorynoscelis Rodendorf, 1951 : 65. Type-species: Corynoscelis eximia Boheman, 1858, by monotypy. Hyperoscelis Hardy & Nagatomi, 1960 : 264. [Replacement name for Corynoscelis Boheman.] NOMENCLATURE. Zetterstedt's footnote (1860) stating that he had prepared a description of Corynoscelis under the name Spiloptera produces a problem of nomenclature. Jerdon (1862) applied the name Spiloptera to a genus of birds and Gates (1889), considering Spiloptera Jerdon a junior homonym of Spiloptera Zetterstedt, proposed the name Elachura as a replacement name. Elachura is currently regarded as a junior synonym of Spelaeornis David & Oustalet, 1877 (Peters, 1964). Hardy & Nagatomi (1960), realising the homonymy of Corynoscelis Boheman and Corynoscelis Burmeister, proposed the replacement name Hyperoscelis for Boheman's genus. At that time their action was perfectly correct, but since then the International Code of Zoological Nomenclature (ICZN, 1964) has been altered so that Article nd now states that a name proposed in synonymy is to be regarded as available if it has been used as a primary homonym. Such is the case with Spiloptera Zetterstedt, which is therefore the oldest available name for this genus. However, since Spiloptera Zetterstedt has never been referred to in entomological literature since 1860, I am regarding it as a nomen oblitum and intend to apply to the International Commission on Zoological Nomenclature to ratify this. The name Eucorynoscelis also predates Hyperoscelis, but I am not certain of its validity. It appears in the legend of a figure in Rodendorf (1951) in the combination Eucorynoscelis eximia Boheman. It is not mentioned in the text. SYNNEURIDAE AND CANTH YLOSCELIDAE 79 I have not found the name anywhere else. If this is its only mention then it is an obvious synonym of Corynoscelis and Hyperoscelis (same type-species by monotypy), but since it may be described elsewhere (it is included in his family Canthyloscelidae), I have applied it to this genus with reservations. DIAGNOSIS. Three ocelli present, median one reduced. Antennae i6-segmented, loosely jointed. Eyes narrowly separated above and below antennae. Gena small, mouth margin very close to bottom of eyes. First segment of palpi very small, second rounded, third and fourth elongate. Minute single spurs on fore and mid tibiae. Tarsal claws simple. Empodium broadly rounded and fringed with hairs. Hind femur swollen. Hind tibia slender and curved to fit femur. Fore and mid legs yellow, hind legs variegated. Wings with posterior veins well pigmented. A short vein R t present and usually ending in costa, otherwise incomplete and ending free. Vein r-m represented by a long fusion. Median fork complete. Cu z very weakly present. Anal vein reaching margin, but weak towards wing base. Basal abscissa of M 3+4 in line with short m-cu. Seven unmodified pregenital segments. with tergite 8 almost atrophied. $ with 2-segmented ovipositor and one simple spermatheca. DISTRIBUTION. Palaearctic. DISCUSSION. Boheman (1858) first described Corynoscelis eximia from material sent by Holmgren from Swedish Lapland. In the same year Loew (1858) described and figured the same specimen (s). Presumably because he knew that Boheman was describing it, Loew did not give it a name. Zetterstedt (1860) described it using Boheman's name, but in a footnote stated that he too had already seen Holmgren's specimen and had prepared his description of it under the name Spiloptera arctica. Mik (1886), still discussing the same material, though probably not from first hand experience, noted its superficial similarity to certain Empidid genera (Oedalea and Hybos) and confirmed this similarity to the latter genus when he (Mik, 1900) obtained a specimen from Rumania which he considered to be the same species (but see p. 83). Lundstrom (1910) described what he thought was the female of eximia, but the sex and identity of these specimens are in doubt (see p. 82). Dahl (1911) described a larviform female insect as the female of Corynoscelis. The specimen had been found on an empty lepidopterous pupa and, assuming it to be parasitic, Dahl discussed its relationship with fleas, etc., as well as Scatopsidae and Phoridae. Bergroth (1912) was quick to point out Lundstrom's (1910) description attributed to the female of eximia as a normally full-winged fly, but agreed with Dahl that his specimen belonged near Corynoscelis and Scatopse in the Bibionidae. Enderlein (1912), without seeing the specimen himself, considered Dahl's specimen as more Mycetophilid-like and erected a new genus and species for it, Dahlica larviformis. In 1936 Enderlein put it in a separate subfamily of Mycetophilidae. There it remained until Bartoova & Duskova (1958) described a second species, Dahlica hirta, from Czechoslovakia. Their good description and figures enabled Stys (1960) to re-examine the affinities of Dahlica and he realized that both species were moths and probably Psychidae. Dahlica larviformis Enderlein is now regarded as a synonym of Solenobia triquetrella (Hiibner, 1812) (Lepidoptera: Psychidae) (vide Dierl, 1968). Meanwhile, Frey (1916) had recorded another specimen from Finland, after 8o A. M. HUTSON which no further specimens of the genus are known to me until Hardy & Nagatomi (1960) described a second species, insignis, from a single male from Japan. Mamaev & Krivosheina (1969) reared large numbers of Hyperoscelis of two species from larvae found in rotting wood. By examining type-material of eximia they were correctly able to decide which species was undescribed. The undescribed species, veternosa, was described from 21 males and the sexes of the material of eximia were not noted, so it was not until Martinovsky (1972) that a true female Hyperoscelis was adequately described. This is the most recent published record of Hyperoscelis known to me. Of the three described species I can only recognise two as valid, and these are separable by the following key. KEY TO SPECIES OF Hyperoscelis i Length of R 5 equal to distance between the base of M 1+z and the fork of R i+5 . Basal part of M not parallel with R lt unsclerotized, without macrotrichia ; macrotrichia absent between these two veins. Tergite 8 of male with lateral processes and a deep median emargination, claspers as Text-fig. 17, sternite 9 with marked lateral extensions (Text-figs 13, 15-17). Female ovipositor as Text-fig. 19 eximia Boheman (p. 80) - R 5 shorter than distance between base of M 1+2 and the fork of R t+5 . Basal part of M before the r-m fusion parallel with R lt heavily sclerotized and covered with macrotrichia; macrotrichia present on wing membrane between these two veins. Tergite 8 of male smoothly rounded with a shallow median emargination, claspers as Text-fig. 18, sternite 9 without marked lateral extensions (Text-fig. 14). veternosa Mamaev & Krivosheina (p. 83) Hyperoscelis eximia (Boheman) (Text-figs 13, 15-17, 19) Corynoscelis eximia Boheman, 1858 : 56. Syntypes 2 <$, i $, SWEDEN: Tarna [6543' N, I5i7' E], Laxfjellet, 15-18. vii. (A. E. Holmgren) (NR, Stockholm) [examined]. Hyperoscelis insignis Hardy & Nagatomi, 1960 : 265. Holotype , JAPAN: Hataganaru (Tazima), 26^.1955 (E. Fujita) (KUF, Fukuoka) [examined]. Syn. n. DIAGNOSIS. As in key to species. Some authors (Hardy & Nagatomi, 1960 and Martinovsky, 1972) have overlooked the small first palpal segment and considered the palps 3-segmented. All specimens I have seen have 4-segmented palps. MATERIAL EXAMINED. 2 <$, i $ syntypes of eximia, SWEDEN (details in synonymy). Holotype <$ of insignis, JAPAN (details in synonymy). CZECHOSLOVAKIA: Moravia, Bedrichov bei Rymarov (Bezirk Sumperk), 500 m, i8.vi.i970 (/. Martinovsky}, i $ (coll. J. Martinovsky, Olomouc); Moravia, Josefova u Branne, 20. vi. 1972 (/. Martinovsky}, i $ (BMNH). DISCUSSION. Boheman's specimens are in good condition. No holotype was originally recorded and no lectotype has been designated since. As there is no SYNNEURIDAE AND CANTH YLOSCELID AE 81 suggestion that the series consists of more than one species, I have not selected a lectotype. The genitalia of one of the males have been cleared and are in a microvial on the pin, and are figured here (Text-figs 13, 15-17). Mik (1900) recorded a specimen of this species from Rumania, but his specimen proves to be veternosa (see under that species). 13 15 19 FIGS 1319. Hyperoscelis species. 13, eximia, genitalia, ventral view; 14, veternosa, genitalia, ventral view; 15-17, eximia showing (15), <$ sperm pump, lateral view, (16)^ sperm pump, ventral view, (17) outline of lateral view of clasper; 18, veternosa, outline of lateral view of $ clasper; 19, eximia, $ ovipositor, ventral (left) and dorsal (right) views. 82 A. M. HUTSON Lundstrom (1910) and Frey (1916) recorded three specimens (as $) from three localities in Finland. Two of these specimens have been attacked by pest beetle and all that remains are three wing fragments and one antenna. One of the wing fragments shows the base of a strong vein R^ which, estimated from the position of the medial veins, may be in the relative position of eximia rather than veternosa. Lundstrom's two specimens were from Ruovesi [6i59'N, 2405'E] and Tuovilanlaks [c.63N, c.28E]. There is a bare celluloid strip on the pin of the Ruovesi specimen; whatever was mounted on it is lost except for a small piece of tissue which could be eggs. If it is eggs then the specimen must have been female, otherwise there is no way of sexing the original specimen. A separate mount of one antenna is part of this specimen. The Tuovilanlaks specimen is the specimen that Hardy & Nagatomi (1960) used to compare with their specimen from Japan and is a male of veternosa, misleading them into describing their specimen as a new species. Of Frey's specimen from Kangasala [6i3o'N, 24oo'E] there is no trace on the pin. One, or two, of the wing fragments mentioned above belong to this specimen, but it remains unidentifiable. The Tuovilanlaks specimen and the remains of the other two specimens are in the University Museum, Helsinki, Finland. Rodendorf (1951), in his discussion on the locomotory organs of Diptera, figures the wing of what he calls Eucorynoscelis eximia Boheman from Denmark. The status of the name Eucorynoscelis has already been discussed and I know of no specimen of this family from Denmark. Hardy & Nagatomi (1960) described H. insignis from a single $. In structural details it does not differ from eximia, although there are quite distinct differences in the colour of the thorax and abdomen. In all other specimens of the genus examined in the present study the only yellow visible on these parts was around the anterior spiracle, otherwise the thorax and abdomen are dark reddish brown to black. In the Japanese specimen, most of the pronotum, the area of the anterior spiracle and the scutellum, particularly medially, are all yellow and the dorsum is yellow with three black stripes. The abdomen has yellow spots along the mid-dorsal line and the sternites are completely yellow. Despite these colour differences I have no hesitation in synonymizing insignis with eximia. Mamaev & Krivosheina (1969) recorded eximia from the Maritime Province on the east coast of the U.S.S.R. Their specimens were bred from various types of rotting forest timber, mainly elm. Other series of larvae that were probably of this species were found in spruce wood much damaged by brown rot. These were from the Moscow and Yaroslavl Regions of the U.S.S.R. This material is in the IAEME, Moscow. Martinovsky (1972) was the first to adequately describe and figure the female from his first specimen, which was found in a stand of Rubus idaeus L. in a mixed wood. His first specimen is in his private collection and he has kindly donated a second specimen to the BMNH. DISTRIBUTION. Sweden, Czechoslovakia, U.S.S.R. (Maritime Province), Japan. Also probably in Finland and the Yaroslavl and Moscow Regions of U.S.S.R. SYNNEURIDAE AND CANTH YLOSCELIDAE 83 Hyperoscelis veternosa Mamaev & Krivosheina (Text-figs 14, 19) Hyperoscelis veternosa Mamaev & Krivosheina, 1969 : 936. Holotype <$, U.S.S.R.: Ukraine, Rakhov, 17. vi. 1966 (B. Mamaev) (IAEME, Moscow) [not examined]. DIAGNOSIS. As in key to species. MATERIAL EXAMINED. U.S.S.R.: same data as holotype, 2 <$ (paratypes) (BMNH). RUMANIA: Bucharest, 1898 (J. Mik), i <$ (NM, Vienna). FINLAND: Tuovilanlaks, 2.vi.i865, i c? (ZMU, Helsinki). DISCUSSION. The type-material of 21 males was reared from larvae in spruce wood affected by brown rot. In recording his specimen from Rumania, Mik (1900) noted that vein R (by current terminology) is rudimentary and does not reach the costa. As this is a feature of veternosa, the specimen was re-examined and found to be < veternosa. While the identity of Lundstrom's Ruovesi specimen and Prey's Kangasala specimen remains in doubt as the specimens are destroyed (see above under eximia) , Lundstrom's other specimen, from Tuovilanlaks, was described and figured by Hardy & Nagatomi (1960) as eximia. The specimen is mounted on a slide and is a ( veternosa. DISTRIBUTION. Finland, Rumania, U.S.S.R. (Ukraine: Transcarpathian Mts). CANTHYLOSCELIS Edwards (Text-figs 20-25) Canthyloscelis Edwards, 1922 : 268. Type-species: Canthyloscelis antennata Edwards, 1922, by original designation. DIAGNOSIS. Two large lateral ocelli, small median ocellus present (subgenus Araucoscelis) or absent (subgenus Canthyloscelis). Antennae i6-segmented, simple to distinctly pectinate, at least as long as head and thorax combined. Palpi 4-segmented, long and slender. Tibial comb well developed (Araucoscelis) or poorly developed (Canthyloscelis). Tarsal claws with a large basal lobe bearing a number of small teeth. Empodium large and fleshy. Hind femur greatly swollen, often with a row of small pegs on the ventral surface. Hind tibia slender and curved to fit tightly against femur. Wings with posterior veins almost as heavily pigmented as anterior veins. A short R t present and ending in costa. r-m represented by a long fusion. M z interrupted at base. Males with only 2 or 4 (Canthyloscelis) or 6 (Araucoscelis) unmodified pregenital segments. Ovipositor 2-segmented. One spermatheca, strikingly modified in Canthyloscelis (Text-fig. 25) or simple in Araucoscelis. DISCUSSION. Edwards (1922) described Canthyloscelis for three species in New Zealand. In 1930, he described a second subgenus (Araucoscelis} for two species from South America, to which he later (1934) added a third species, also from South America. In describing a fourth South American species, Toilet (1959) separated the South American species into two subgenera on the basis of the pectinate 84 A. M. HUTSON antennae of the male of one species. A fourth New Zealand species is described here. The sharp distinctions between the New Zealand and South American species-groups are clear; these could well be regarded as separate genera, but it is convenient to leave them as congeners for the time being, thereby expressing the opinion that they are sister-groups and that together they are the sister-group of Hyperoscelis (as proposed by Hennig, 1960). I would also agree with Hennig (op. cit.) that the South American species do not merit separation into two subgenera purely on the basis of whether or not the male antennae are pectinate, a patently derived character. Certainly the South American species do not differ in any fundamental character, and with the species synonymy given here such a division serves no useful purpose. DISTRIBUTION. New Zealand and South America. KEY TO SUBGENERA AND SPECIES OF Canthyloscelis Median ocellus absent. Eyes narrowly separated above antennae. Flagellar segments with distinct necks. Gena well developed, so lower margin of eyes well separated from mouth margin. Meron well developed. Hind femur swollen in apical two-thirds. Wing vein R 5 not thickened towards tip. Basal abscissa of vein M 3+i almost vertical, joining Cu before the posterior fork. Two or 4 unmodified pregenital segments in male. Female with single complex spermatheca. (New Zealand) (Subgenus CANTHYLOSCELIS) 2 Small median ocellus present. Eyes broadly in contact above antennae. Mouth margin very close to bottom of eyes. Meron somewhat reduced. Hind femur swollen from near base. Wing vein R 5 thickened beyond fork with R t . Basal abscissa of M 3+4 almost vertical and in line with short m-cu. Six unmodified pregenital segments in male. Female with single simple spermatheca. (South America) (Subgenus ARA UCOSCELIS) 5 Wings with dark mark near apex. Pleurae at least somewhat darkened. Eyes of . (All paratypes; all in BMNH.) DISTRIBUTION. Border area of Argentina and S. Chile, near L. Nahuel Huapi. Canthyloscelis (Araucoscelis) pictipennis Edwards Canthyloscelis (Araucoscelis) pictipennis Edwards, 1930 : 92. Holotype $, ARGENTINA: Terr. Rio Negro, Lake Frias, 3-xii.i926 (F. & M. Edwards) (BMNH) [examined]. Canthyloscelis apicata Edwards, 1934 : 186. 4 syntypes , CHILE: Comudes, 16-17.^.1902 (S. Schonemann) (i in BMNH) [examined]. Syn. n. Canthyloscelis (Chiliscelis) valdiviana Toilet, 1959 : 147. Holotype , CHILE: Valdivia prov., nr L. Panguipulli, Shoshnenco, 200 m, 4-6.111.1955 (L. E. Pena) (IRSNB, Brussels) [examined]. Syn. n. DIAGNOSIS. Flagellar segments of male slightly serrate, loosely jointed but without obvious necks. Palps pale yellow. First hind tarsal segment about 1-5 times as long as second. Wings with dark subapical band and slightly darkened costal cell. Posterior veins darkened, as usual. Head (usually), thorax and abdomen dark reddish brown, thorax unstriped. Legs as pectinata, except that hind coxae dark in the Casa Pangue g. Sternite 7 of male greatly enlarged, bowl-shaped with genitalia directed dorsally. Tergite 9 with pair of broad, straight median processes. Claspers broad, triangular, with small inner lateral processes. The processes below this (? dististyles) short, straight and blunt, slightly bulbous-tipped. MATERIAL EXAMINED. Holotype $ of pictipennis, ARGENTINA: (details in synonymy), i $ syntype of apicata, CHILE (details in synonymy). Holotype <$ of valdiviana, CHILE (details in synonymy). CHILE: Llanquihue, Casa Pangue, xii. 1926 (R. 6- E. Shannon), i <$ (USNM, Washington); Chiloe Island, Dalcahue, 10-12.^.1957 (L. E. Pena), i $ (allotype of valdiviana) (IRSNB, Brussels). SYNNEURIDAE AND CANTH YLOSCELIDAE 87 DISCUSSION. Edwards (1934) recorded a specimen as the male of pictipennis, which he had described from a single female. I have examined this specimen and find it the same species as the single male syntype of apicata that is in the BMNH. C. apicata was described from four male syntypes, the other three are stated to be in the Zoological Museum of Berlin (now the Zoologisches Museum fur Naturkunde der Humboldt-Universitat) and have not been examined in the present study. C. valdiviana was described from several males and two females, and the male holotype and female allotype were examined in the present study. I can see no difference at the species level between the holotype of valdiviana and the BMNH type of apicata and so am confident in synonymising these two species. Thus apicata, the male attributed to pictipennis by Edwards (1934), and valdiviana are all the same species. The two females I have seen are the type of pictipennis and the allotype of valdiviana. Slight differences in the structure of the ovipositor are detectable, but are not enough to regard the two as distinct species and colour differences are no more than the variety shown by the males. The locality of the type of pictipennis, although actually in Argentina, is closer to the locality of the males described as valdiviana and pictipennis, than are the females described as valdiviana. I therefore regard all these as a single species, i.e. apicata and valdiviana are synonyms of pictipennis. DISTRIBUTION. Chile and the border of Argentina between 37 and 43S (I have not been able to trace Comudes, the type-locality of apicata}. Subgenus CANTHYLOSCELIS Edwards Canthyloscelis Edwards, 1922 : 268. DIAGNOSIS. As in key on p. 84. DISCUSSION. Four species are recognized in this subgenus. They could be separated into two species-groups, namely the antennata-gronp (antennata, nigricoxa, balaena) and the claripennis-group (claripennis) , on the characters in the first half of couplet 2 of the key on p. 84. DISTRIBUTION. New Zealand. Canthyloscelis (Canthyloscelis) antennata Edwards (Text-figs 20, 24-25) Canthyloscelis antennata Edwards, 1922 : 268. Holotype <$, NEW ZEALAND: Wainuiomata, in forest, I4.xii.ig20 (G. V. Hudson] (BMNH) [examined]. DIAGNOSIS. Eyes of male almost touching above antennae. Dorsum of thorax with pale ground colour and three more or less distinct stripes. Pleura somewhat darkened. Wings with dark subapical band. Wings with microtrichia restricted to marginal areas on upper surface, lower surface lacking microtrichia in costal, R lt basal and most of anal cells and extreme bases of other cells. All coxae pale yellowish. Hind basitarsus about twice as long 88 A. M. HUTSON as second tarsal segment. Male abdomen (Text-fig. 20) with two unmodified pregenital segments. Tergite nine of male with a pair of long fine points. MATERIAL EXAMINED. Holotype , NEW ZEALAND (details in synonymy). NEW ZEALAND: same data as holotype, 2 <$ (paratypes); no locality (G. V. Hudson), 1 ^, i $ (paratypes); Ohakune, 20. xi. 1919 (T. R. Harris), i ^; Ohakune, xi. 1922 (T. R. Harris), i <, i $>; Ohakune, 1-9.^.1923 (T. R. Harris), i <$, i $; Ohakune, 2060', i.iii.igig (T. R. Harris], 2 $; Ohakune, 2060', 9.^.1920 (T. R. Harris), 2 $; no locality, 1928 (G. V. Hudson), 3 & i $. (All in BMNH.) DISTRIBUTION. New Zealand. \ Canthyloscelis (Canthyloscelis) nigricoxa Edwards (Text-fig. 21) Canthyloscelis nigricoxa Edwards, 1922 : 269. Holotype <$, NEW ZEALAND (G. V. Hudson) (BMNH) [examined]. DIAGNOSIS. Eyes almost touching above antennae. Dorsum of thorax uniformly reddish. Pleurae somewhat darkened. Wings with dark subapical band. Wings with microtrichia restricted to marginal areas on upper surface and lower surface lacking microtrichia on costal, RL basal cells and basal areas of cells R 5 and M z , and restricted to outer parts in cells R v M 3+t , Cu and anal. Hind coxae shining black, others yellowish. Hind basitarsus about equal in length to second tarsal segment. Male abdomen (Text-fig. 21) with two unmodified pregenital segments. Tergite 9 of male simple. MATERIAL EXAMINED. Holotype <$, NEW ZEALAND (details in synonymy). DISTRIBUTION. New Zealand. Canthyloscelis (Canthyloscelis) balaena sp. n. (Text-fig. 22) DIAGNOSIS. Eyes narrowly separated above antennae. Dorsum of thorax uniformly light brown. Pleurae similarly coloured, becoming darker towards ventral parts. Wings with dark subapical band. Wings with microtrichia widespread on dorsal and ventral surfaces. Wings lacking microtrichia in most of costal, Cu, anal and basal cells and extreme bases of cells R and M 2 . R 1 ending beyond the level of the beginning of vein M 2 . All coxae pale yellowish. Hind basitarsus more than twice length of second tarsal segment. Male abdomen (Text-fig. 22) with two unmodified pregenital segments (the second short and narrow). Tergite 3 folded, with the fold running across the abdomen, the lateral corners forming anteriorly directed processes, apical median area strongly emarginated. Tergite 4 weakly developed and medially emarginated. Tergite 5 so heavily emarginated that it appears as two lateral semicircular plates. Tergite 6 more distorted to appear as two crescent-shaped plates joined at their antero-dorsal ends. Tergites 7 and 8 simple. Tergite 9 with short broad points separated by a shallow evenly curved emargination. Claspers very small and rounded. SYNNEURIDAE AND CANTHYLOSCELIDAE 89 MATERIAL EXAMINED. Holotype <$, NEW ZEALAND: 1928 (G. V. Hudson), no. i36q (BMNH). DISCUSSION. The specimen was in the series of antennata, but from the structure of the abdomen I would regard it as closer to nigricoxa. The structure of the abdomen and the extensive microtrichia of the wings readily distinguish this species from the others of the antennata-group. DISTRIBUTION. New Zealand. Canthyloscelis (Canthyloscelis) claripennis Edwards (Text-fig. 23) Canthyloscelis claripennis Edwards, 1922 : 268. Holotype $, NEW ZEALAND (G. V. Hudson) (BMNH) [examined]. DIAGNOSIS. Eyes of male distinctly separated above antennae. Dorsum of thorax with pale ground colour and three vague, dull brown, (almost) contiguous stripes. Pleurae uniformly yellowish. Wings completely clear. Wings with microtrichia extensive on upper as well as lower surface of wing, covering entire wing except for extreme base. All coxae pale yellowish. Hind basitarsus about twice as long as second tarsal segment. Male abdomen (Text-fig. 23) with four unmodified pregenital segments. Tergite 9 with a broad simple flange. MATERIAL EXAMINED. Holotype <$, NEW ZEALAND (details in synonymy). NEW ZEALAND: Ohakune, i. 1920 (T. R. Harris), i ; Ohakune, i5.xii.i922- 15.1.1923 (T. R. Harris), i 9; Ohakune, v. 1922 (J. W. Campbell), i $. (All in BMNH.) DISCUSSION. The second <$ is the specimen that Edwards (1930) mentions as a fourth New Zealand species of Canthyloscelis. DISTRIBUTION. New Zealand. CHECKLIST AND PROPOSED CLASSIFICATION OF SYNNEURIDAE AND CANTHYLOSCELIDAE Family SYNNEURIDAE Enderlein, 1936 Genus SYNNEURON Lundstrom, 1910 annulipes Lundstrom, 1910 decipiens sp. n. silvestre Mamaev & Krivosheina, 1969 Genus EXILISCELIS gen. n. californiensis sp. n. Family CANTHYLOSCELIDAE Rodendorf, 1951 Corynoscelidae Enderlein, 1912 Hyperoscelidae Hardy & Nagatomi, 1960 9 o A. M. HUTSON Genus HYPEROSCELIS Hardy & Nagatomi, 1960 Corynoscelis Boheman, 1858 Spiloptera Zetterstedt, 1860 ? Eucorynoscelis Rodendorf, 1951 exitnia Boheman, 1858 insignis Hardy & Nagatomi, 1960 syn. n. veternosa Mamaev & Krivosheina, 1969 Genus CANTHYLOSCELIS Edwards, 1922 Subgenus ARAUCOSCELIS Edwards, 1930 Chiliscelis Toilet, 1959 pectinata Edwards, 1930 pictipennis Edwards, 1930 apicata Edwards, 1934 s y n - n - valdiviana Toilet, 1959 syn. n. Subgenus CANTHYLOSCELIS Edwards, 1922 antennata Edwards, 1922 nigricoxa Edwards, 1922 balaena sp. n. claripennis Edwards, 1922 SYSTEMATICS Synneuridae and Canthyloscelidae, as treated here, consist of four very well differentiated small genera each of which has a limited distribution within the wide distribution of the group as a whole: Synneuron with three species and a Holarctic distribution, Exiliscelis with one (possibly two) Nearctic species, Hyperoscelis with two Palaearctic species and Canthyloscelis with two well marked subgenera, one with two species in the Neotropical region and one with four species in New Zealand. This is strong evidence of a relict group. They are undoubtedly related to the Scatopsidae, which is a relatively large family with a worldwide distribution and about 200 species. Some Scatopsidae, particularly Coboldia fuscipes (Meigen), but also such species as Scatopse notata (Linnaeus) and Holoplagia guamensis Johannsen, have been able to occupy niches over a wide distribution, both naturally and with the aid of man. The Scatopsidae, Synneuridae and Canthyloscelidae are clearly derived from the same stock, the Scatopsidae being the present day successful and versatile lineage. The evolution of this group and the relationships of one successful group to four relict groups is of interest. In preparing the description of Exiliscelis, the Synneuridae and Canthyloscelidae were compared with a variety of Scatopsidae, particularly Scatopse notata (Linnaeus), Aspistes berolinensis (Meigen), Anapausis soluta (Loew), Psectrosciara africana Cook and Ectaetia clavipes (Loew). From this it was apparent that the Scatopsidae and Synneuron were almost as closely related to each other as Hyperoscelis is to Canthyloscelis. Exiliscelis appeared to be the most primitive genus and was more closely related to Synneuron than to Hyperoscelis. A systematic analysis of about 30 characters was prepared, dividing the various states of each character into plesiomorphic (primitive) or apomorphic (derived) condition (Hennig, 19666) based on the assumption that the whole group belongs to the Mycetophiloid complex of Nematocera. These characters are SYNNEURIDAE AND CANTHYLOSCELIDAE 91 o oo H ON 5 X X X X X O O o o o o o O o o o o o o o o o o o o o o o o X X o o o M CO X X o o o 1 10 . . a! H X X X o 'C n) ON X X X o o 0) 2 ^ *^ X X X o o o tn OH _^ a) * M X X X o o o E .c _g ^ X X X o o o Ifl ^ CO X X o X o 1 4_) cd C/5 O ^-^ u M o X X X o o 3 ' c C/3 IH 10 g o X o X H -4-" 'a; o (/) O O o^l^ u o "rt o CO 3 a >. (fl Exilisce & M S'S^^ t: o < o g ISI (/> u cj y. 92 A. M. HUTSON listed in the Appendix, giving the state of each character exhibited by the Scatopsidae, followed by the alternative state in square brackets. Where the state is considered to be apomorphic it is given in italics. These can be compared with the descriptions given earlier in this paper. As might be expected in a group containing 200 species, Scatopsidae shows much more variation than in any of the other genera. This may have lead to confusion in the selection of the apomorphic state of characters, but it is to be hoped that mistakes of this type have not distorted the overall picture of relationships. Table i shows a character matrix for the 32 characters examined. A representation of the probable phylogenetic relationships based on these data is given in Text-fig. 26 and this agrees with the opinion on relationships arrived at by traditional means. By both methods the Scatopsidae are only a part of the Synneurid complex and whether the Synneuridae can continue to be maintained as a separate family must be questioned. Further evidence may exist in the fossil record. Rodendorf (1946) described Protoscatopse jurassica from Jurassic material from Kara Tau, Kazakhstan, U.S.S.R. He puts it in a separate family, the Protoscatopsidae. It has not been examined in the present study, but it is obviously a specimen that should be re-examined, since Rodendorf's figures suggest some interesting characters, such as the presence of R A Scatopsidae Synneuron Exiliscelis Hyperoscelis Canthyloscelis (Araucoscelis) Canthyloscelis (Canthyloscelis ) FIG. 26. Phylogenetic relationships of Scatopsidae, Synneuridae and Canthyloscelidae. Numbers refer to the characters listed in Appendix i and tabulated in Table i. = apomorphy, O = plesiomorphy. SYNNEURIDAE AND CANTH YLOSCELID AE 93 ABLE 2 oscelis, Scatopse, Mycetophilidae and Cecidomyiidae. Mycetophilidae Cecidomyiidae Distinct head capsule Head capsule very small, but distinct Mouth parts well developed Mouthparts very reduced Body of twelve segments Body of twelve segments Integument soft, colourless Integument soft, thin and and usually thin colourless y Body (usually) subcylindrical Body usually slightly flattenei dorso-ventrally Spiracles sessile Spiracles not or only slightly projecting from body Spiracle i larger than others All spiracles same size en "o TO IH "B< en HH O en IH 'TO OH ON 4n $ M 'On en en IH ' oo Last pair of spiracles on yth Last pair of spiracles on abdominal segment penultimate segment :t Soft terminal lobes sometimes Soft terminal lobes sometimes +-> C CD en CD IH OH -l-> C CD en OH H -g >, 13 a % T3 arval characters of Cant 1 Distinct head capsule CD OH CD en S OH 3 Body of twelve segments Integument tough, strong] discoloured with strong hairs and granulations Body flattened dorso-vent Spiracles on short tubes *H CD -(-> ai en 13 IH 'rt 4-> 3 CO C O W) CO SH 'o 'On ON Last pair of spiracles on posterior border of penultimate segment Terminal armature of two processes on two isolate plates ! d "o IH n OH en C TO 1 o CH rO o en C IH o C! rM a O UH T* ft Comparisoi Canthyloscelis Head capsule absent O SH O OH O I-, "TO IH TO CD CD bo CD en CD M-l O Integument soft, thin colourless Body subcylindrical Spiracles sessile All spiracles same size structure en 'o TO "On en yellowish or brownish with bold, paler markings (not well-defined in C. chiliensis) forming a distinctive pattern (PI. i, figs 1-8); subterminal line of transverse, pale streaks always well-defined. Costa straight, slightly convex towards apex ; termen curving evenly and continuously with posterior margin. Venation as in Text-fig. 4: Sc unbranched, R z and R 3 stalked from two-thirds to four-fifths, M complete, forked, arising from CuA, basal half of CuP present in fore wing. Hind wing orange to greyish brown, basally reddish or yellowish in several species, in some species with pale subterminal spots; underside of hind wing with dense hair in subcostal field. GENITALIA <. Eighth sternite present, small, relatively heavily sclerotized, showing tendency towards separation into pair of lateral sclerites, but absent in C. izquierdoi. Saccus semicircular, heavily sclerotized. Ventral lobe from vinculum overlying saccus, heavily sclerotized. Vinculum stout, U-shaped. Valve stout, kidney-shaped, triangular, or narrow with inturned tip, heavily sclerotized and densely setose. Base of juxta pinched to form tube (but tube not closed caudally in C. argentata). Tegumen and caudal region of juxta completely fused, forming large, globose intromittent organ; juxta with weakly sclerotized ventral crease or medial emargination (the latter particularly conspicuous in C. izquierdoi), with or without lateral digitate processes. Tegumen extended caudally as a sideways T-shaped sclerotization, ventral branch of 'T' produced into heavily sclerotized digitate process in several species. Pair of socii [see 'Morphology'] adjacent to dorsal margin of tegumeri, with or without minute, setose mediocaudal sclerite. Uncus, if present, a more or less heavily sclerotized fold in caudal abdominal membrane, Y-shaped or slit-like. Aedeagus membranous, scobinate at base, consisting of infolded ductus ejaculatorius; base of ductus ejaculatorius distal to anus with pair of shallow, bulbous lateral swellings and dorsomedial swelling, base of dorsomedial swelling lightly sclerotized. GENITALIA ^. Eighth segment complete, with fully developed spiracle, eighth sternite small. Ninth sternite heavily sclerotized, tapered posteriorly, with doubly peaked apex. Papillae anales large, triangular or quadrangular. Apophyses absent. Ductus bursae with fine transverse constrictions; bursa copulatrix without signum. KEY TO MALES 1 Antennal segments bipectinate (PI. 6, fig. 33) ..... salasi (p. 117) Antennal segments angular (PI. 6, figs 36-32, 34, 35) . . . . . 2 2 Valve narrow and elongate with inturned tip (PI. 3, fig. 17) . . argentata (p. 118) - Valve broad and short, triangular or kidney-shaped, tip not turned inwards . . 3 3 Wing expanse less than 50 mm, fore wing with black streak on posterior margin (PI. i, fig. 7); juxta with deep medial emargination (PI. 3, fig. 16) izquierdoi (p. 117) - Wing expanse greater than 50 mm, fore wing without black streak on posterior margin; juxta without deep medial emargination or only with weakly sclerotized medial line ............. 4 4 Ventral lobe of vinculum with pair of conical digitate processes towards anterior margin but without transverse ridge ........ 5 Ventral lobe of vinculum without conical processes but with transverse ridge . . 6 5 Fore wing with bold silvery white markings (PI. i, fig. i); socii separate arenosus (p. 112) - Fore wing without bold silvery white markings (PI. i, fig. 2); socii fused together chiliensis (p. 113) U2 G.S.ROBINSON 6 Fore wing with conspicuous bold white markings (PI. i, fig. 3) ; digitate process from ventral arm of tegumen claw-shaped (PI. 2, fig. 12) . . . . digitata (p. 114) Fore wing without conspicuous white markings; digitate process from ventral arm of tegumen short and tooth-like or stout and blunt (PI. 2, fig. 13, PI. 3, fig. 14) . 7 7 Base of hind wing dull yellowish brown, fore wing without narrow silvery subterminal line (PI. i, fig. 4); eighth sternite constricted medially, digitate process from ventral arm of tegumen stout and blunt (PI. 2, fig. 13) . . brunnescens (p. 115) Base of hind wing pinkish, fore wing with silvery subterminal line (PI. i, fig. 5); eighth sternite not constricted medially, digitate process from ventral arm of tegumen short, tooth-like (PI. 3, fig. 14) . . . . . castilloi (p. 116) KEY TO FEMALES Females of chiliensis, digitata, brunnescens, castilloi, salasi and argentata are unknown. 1 Epiphysis one-quarter length of tibia; antennal segments barrel-shaped (PI. 6, fig. 36) .......... sp. indet. (p. 119) Epiphysis one-half length of tibia; antennal segments angular .... 2 2 Papillae anales triangular, overlapping ninth sternite which has pointed tips (PI. 5, fig. 28) .......... arenosus (p. 112) Papillae anales quadrangular, not overlapping ninth sternite which has rounded tips (PI. 5, fig. 29) ........ izquierdoi (p. 117) DESCRIPTIONS OF SPECIES Callipielus arenosus Butler (PL i, fig. i; PI. 2, fig. 10; PI. 3, fig. 18; PI. 5, figs 26, 28; PL 6, fig. 30) Callipielus arenosus Butler, 1882, Trans, ent. Soc. Lond. 1882 : 24, pi. i, fig. 6. LECTOTYPE , CHILE: [Valdivia (Edmonds}'] (genitalia slide no. 12348; BMNH), here designated [examined]. Callipielus arenosus Butler; Viette, 19500! : 52, fig. I [redescription]. Callipielus leukogramma Bryk, 1945, Ark. Zool. 36A(3) : 26, pi. 2, figs 15, 16. Holotype <$, ARGENTINA: Lago Nahuel Huapi, Llau Llau Peninsula, N. of Puerto Nuevo, 770-780 m, x.i933-iii.i934 (Lundblad) (genitalia slide no. 8-02; NR, Stockholm) [examined]. Syn. n. $. 51-61 mm. Antenna with 46-49 angular segments (PI. 6, fig. 30). Head buff -brown, labial palpus with darker lateral scales. Thorax and tegula buff-brown. Fore wing dark brown and ochreous brown with bold silvery white markings picked out with dark brown margins. Hind wing greyish brown with paler, diffuse subterminal maculae; base of hind wing and anal angle yellow. Legs light buff-brown, darker and greyish above. Epiphysis one-half length of tibia. ? 7> 75 nim. Antenna with 43 angular segments (one specimen only examined) . Similarly patterned to <$. GENITALIA ^ (PI. 2, fig. 10; PL 3 fig. 18; PI. 5, fig. 26). Eighth sternite 0-9-1-6 x 0-3 mm. Ventral lobe of vinculum with pair of densely sclerotized, short, conical, digitate processes towards anterior margin. Saccus one-half to two-thirds as wide as vinculum. Valve blunt, triangular, heavily sclerotized and setose, with well-defined dark transverse ridge across outer surface. Sclerotized region of diaphragma between valves quadrate, anterior corners extended laterally. Juxta completely fused with tegumen, sclerotization extended to caudal lip of intromittent organ; posterior lateral processes of juxta semicircular, shallow, with saw-toothed edge. Ventral arm of tegumen "I" very narrow, elongate. Socii large, setose. Small mediocaudal sclerite present or absent. Uncus a crescentic, heavily sclerotized pocket (distinctly Y-shaped in genitalia slide 8-02). GENITALIA $ (PL 5, fig. 28). Ninth sternite large (1-80 x 1-30, 1-60 x 0-95 mm). Papillae anales triangular, overlapping ninth sternite. REVISION OF CALLIPIELUS BUTLER 113 REMARKS. C. arenosus is closely allied to C. chiliensis but the latter species has a very different wing pattern, lacking the bold silvery white markings of arenosus. The male genitalia of the two species are very similar but the ventral arms of the tegumen T' are shorter and broader in chiliensis than in arenosus; in arenosus the socii are not fused medially and the uncus does not form a deep and rather square pocket, longer than wide, as in chiliensis. The type-series of C. leukogramma and a single male of arenosus, all in NR, Stockholm, carry labels reading Tatagonien 1932-34 Lago Nahuel Huapi Exp. Ljungner'. Full locality data as cited for the holotype (above) are taken from Bryk (1945). DISTRIBUTION. Chile, Argentina. MATERIAL EXAMINED. ARGENTINA: i $, 'taken laying on wet turf near a stream in open rega', 24.xii.i93i (Reynolds) (genitalia slide no. 12349; BMNH); 2 <, i $ (including allotype and paratype of leukogramma), Lago Nahuel Huapi, Llau Llau Peninsula, N. of Puerto Nuevo, 770-780 m, x. 1933-1^.1934 (Lundblad) (genitalia slide nos S-OL, 8-03; NR, Stockholm). CHILE: i $, Tierra del Fuego, Rio McLelland, 24.xii.i9O4 (Crawshay] (genitalia slide no. 12351; BMNH). Callipielus chiliensis Viette (PI. i, fig. 2; PL 2, fig. ii ; PL 3, fig. 19) Callipielus chiliensis Viette, 1950, Zool. Meded., Leiden 31 : 74, figs 16, 17, 18. Holotype , CHILE: Valdivia, 1.1938 (Andreas 6- de Graag) (genitalia slide no. 2187 [Viette]; RNH, Leiden) [not examined]. [Callipielus arenosus Butler, 1882 : 24 (partim i only). Misidentification.] (J. 75 mm. Antenna with angular segments (number unknown). Head greyish, labial palpus darker, brownish. Thorax and tegula buff-brown. Fore wing yellowish brown, darker at costa, with silvery white medial streak close to posterior margin; dark brown diffuse subterminal line, reniform stigma and small streak between reniform and posterior margin; line of small, yellowish spots at termen. Hind wing ochreous brown with subterminal and terminal lines of minute, paler spots; base of hind wing and anal angle yellowish. Legs light buff-brown, darker and greyish above. Epiphysis one-half length of tibia. $. Unknown. GENITALIA g (PL 2, fig. n; PI. 3, fig. 19). Eighth sternite 1-3 x 0-5 mm, trapezoidal, anterior margin 0-8 mm long. Ventral lobe of vinculum with pair of densely sclerotized, short, conical, digitate processes towards anterior margin. Saccus one-half as wide as vinculum. Valve blunt, triangular, heavily sclerotized and setose, with anterior depression in ventral surface, caudal margin of depression a densely sclerotized ridge. Sclerotized region of diaphragma between valves trapezoidal. Juxta completely fused with tegumen, sclerotization extended to caudal lip of intromittent organ; posterior lateral processes of juxta semicircular, shallow, with saw-toothed edge. Ventral arm of tegumen 'T' broad, rather blunt. Socii large, setose, only lightly sclerotized, fused medially. Mediocaudal sclerite absent. Uncus small, a shallow V-shape, with small invaginated membranous pocket which is longer than wide. REMARKS. C. chiliensis is closely allied to C. arenosus but lacks the bold silvery white markings of the latter species. For details of differences in the male genitalia ii4 G. S. ROBINSON see 'Remarks' for arenosus. C. digitata differs from chiliensis in that the juxta of digitata has a small medial emargination and is without posterior lateral processes and the bases of the ventral arms of the tegumen are evaginated to form a stout, dark, claw-like digitate process. C. digitata has much darker fore wings than chiliensis. DISTRIBUTION. Chile. MATERIAL EXAMINED. CHILE: i <, (Edmonds] (genitalia slide no. 12350; BMNH) (paralectotype of arenosus). Callipielus digitata sp. n. (PL i, fig. 3; PL 2, fig. 12; PL 4, fig. 20; PL 6, figs 31, 38) [Callipielus arenosus Butler; Pfitzner & Gaede, 1937 : I2 98, pi. 185, fig. e (partim i only). Misidentification.] (J. 69 mm. Antenna with angular segments (PI. 6, fig. 31) (number unknown). Head whitish, scales tipped with dark brown but completely brown adjacent to eyes, labial palpus greyish brown. Thorax and tegula dark grey-brown anteriorly, metathorax pale yellowish brown. Fore wing dark chocolate-brown patterned with cream and light ochre, distinctive cream triangle at end of cell and pair of basal posterior comma-shaped marks. Hind wing light greyish brown without light subterminal spots, veins darker; base of wing and anal angle dull yellowish with slight pink tint. Legs light greyish brown, slightly darker above. Epiphysis one-half length of tibia (PI. 6, fig. 38). $. Unknown. GENITALIA^ (PI. 2, fig. 12; PI. 4, fig. 20). Eighth sternite small, 0-85 x 0-20 mm, distinctly narrowed medially. Ventral lobe of vinculum with densely sclerotized transverse ridge a little beyond one-half anteriorly. Saccus one-half as wide as vinculum. Valve blunt, triangular, densely sclerotized and setose, with shallow depression in outer surface, posterior corner swollen to form slight hump extending one-third length of posterior margin, hump polished, without setae. Lightly sclerotized region of diaphragma between valves apparently ovate. Juxta completely fused with tegumen, relatively lightly sclerotized but darker posteriorly, sclerotization extended to caudal lip of intromittent organ but with small medial emargination and weakly sclerotized medial line posteriorly; posterior lateral processes of juxta absent. Ventral arm of tegumen "I" narrow, elongate, but weakly sclerotized and ill-defined; base of ventral arm with densely sclerotized claw-shaped digitate process. Socii large, setose, only weakly sclerotized. Small mediocaudal sclerite possibly present as faint sclerotization. Uncus small, forming shallow, triangular, membranous pocket in sclerotized surround. REMARKS. C. digitata is closely allied to C. brunnescens but the latter species lacks the cream markings of digitata. In brunnescens the process from the base of the ventral arm of the tegumen T' is blunt, the socii are densely sclerotized anteriorly and the uncus is represented by a medially divided, narrow, transverse, sclerotized crescent supporting the anterior edge of a shallow but broad, trapezoidal, membranous pocket. The specimen described here appears to be that illustrated by Pfitzner & Gaede (1937 :pl. 185, fig. e) as C. arenosus although, if so, the illustration is somewhat imaginative. The 'beautiful series' of 'arenosus' recorded by Pfitzner REVISION OF CALLIPIELUS BUTLER 115 & Gaede from the Berlin Museum is now represented only by the specimens described here as digitata and brunnescens. The description of 'arenosus' by Pfitzner & Gaede (1937 : 1298) fits true arenosus (above) rather than their illustrated specimen. The description and locality (Valdivia) may be based on Butler's (1882 : 24) original description of arenosus. DISTRIBUTION. Chile. MATERIAL EXAMINED. Holotype <, CHILE (genitalia slide no. H-oi; MNHU, Berlin). Callipielus brunnescens sp. n. (PL i, fig. 4; PI. 2, fig. 13; PL 4, ng. 21; PL 6, fig. 32) [Callipielus arenosus Butler; Pfitzner & Gaede, 1937 : 1298 (partim i ^ only). Misidentifi- cation.] (J. 75mm. Antenna with 63 angular segments (PI. 6, fig. 32). Head light greyish brown, scales lighter basally, labial palpus dull brown. Thorax and tegula dark greyish brown, only a little paler posteriorly. Fore wing dull purplish brown with sandy tint to central field, few very small whitish spots ringed with darker brown, light sandy brown markings in anterior half of wing and on costa. Hind wing pale dull brown, termen light sandy yellow, darker spots between veins at termen; base of hind wing and anal angle yellowish brown; fringe of hind wing greyish brown. Legs dull purplish brown, slightly darker above. Epiphysis one-half length of tibia. $. Unknown. GENITALIA $ (PI. 2, fig. 13; PI. 4, fig. 21). Eighth sternite small, 1-20 x 0-25 mm, distinctly narrowed medially. Ventral lobe of vinculum with densely sclerotized transverse ridge a little beyond one-half anteriorly. Saccus one-half as wide as vinculum. Valve blunt, triangular, densely sclerotized and setose, with shallow depression in outer surface, posterior corner swollen to form a marked hump extending one-quarter length of posterior margin, this hump polished, without setae. Lightly sclerotized region of diaphragma between valves quadrate. Juxta completely fused with tegumen, relatively lightly sclerotized but darker posteriorly, sclerotization extended to caudal lip of intromittent organ but with broad, shallow medial emargination and weakly sclerotized medial line posteriorly; posterior lateral processes of juxta represented by heavily sclerotized lateral humps. Ventral arm of tegumen T' fairly broad and elongate, heavily sclerotized; base of ventral arm with densely sclerotized thumb-shaped digitate process. Socii large, setose, heavily sclerotized anteriorly. Mediocaudal sclerite absent. Uncus a medially-divided, narrow, sclerotized transverse crescent supporting anterior edge of shallow but broad trapezoidal membranous pocket. Membrane between socii and uncus with two pairs of groups of small, scattered setae. REMARKS. C. brunnescens is closely allied to C. digitata but is more dull-looking and has darker hind wings. For differences in the male genitalia see 'Remarks' for digitata. C. brunnescens differs from C. castilloi in that the latter has a pinkish tint at the base of the hind wing, a silvery white interrupted subterminal line in the fore wing and a shorter, blunter ventral arm of the tegumen 'T', the basal process of which is also shorter and blunter. The eighth sternite of castilloi is not constricted medially, the mediocaudal sclerite is present and the uncus is a broad, sclerotized 'U' lying at the base of a shallow pouch. n6 G. S. ROBINSON DISTRIBUTION. Chile. MATERIAL EXAMINED. Holotype $, CHILE: (Lossberg) (genitalia slide no. H-02; MNHU, Berlin). Callipielus castilloi sp. n. (PI. i, fig. 5; PI. 3, fig. 14; PI. 4, fig. 22) (J. 66 mm. Antenna with angular segments (number unknown). Head dark brown, scales of vertex whitish basally; labial palpus dark brown. Thorax and tegula dark brown, lateral mesothoracic hair paler, with a pinkish tint. Fore wing dark, dull brown with diffuse olivaceous spots beyond cell and towards base and posterior margin; subterminal line whitish, broken. Hind wing dark greyish brown without terminal or subterminal spots ; base of hind wing and anal angle yellowish with distinct rose-pink tint at base. Legs dark brown, paler dull brown above. Epiphysis one-half length of tibia. $. Unknown. GENITALIA ^ (PI. 3, fig. 14; PL 4, fig. 22). Eighth sternite 1-3 x 0-2 mm. Ventral lobe of vinculum with densely sclerotized transverse ridge a little beyond one-half anteriorly. Saccus one-half as wide as vinculum. Valve blunt, triangular, densely sclerotized and setose, with shallow depression in outer surface, posterior corner swollen to form pronounced, angular hump extending one-quarter length of posterior margin, this hump polished, without setae. Region of diaphragma between valves hardly sclerotized. Juxta completely fused with tegumen, sclerotization fairly even and extended almost to lip of intromittent organ but with marked medial emargination and weakly sclerotized medial line; posterior lateral processes of juxta absent. Ventral arm of tegumen "I" short and broad, base with short, densely sclerotized digitate process. Socii well-sclerotized, setose. Small mediocaudal sclerite present. Uncus broad, U-shaped, in shallow depression in caudal membrane. REMARKS. C. castilloi is very closely allied to C. brunnescens but has a pinkish base to the hind wing (see 'Remarks' for brunnescens). Of the three species in which a digitate process is formed from the ventral arm of the caudal T'-shaped extremity of the tegumen, castilloi has the shortest process; it has the most angular development of the protuberance on the posterior corner of the valve and exhibits the largest posterior emargination of the juxta of the three species. In C. salasi the juxta has posterior lateral processes, the ventral arm of the tegumen T' is reduced and the basal process is represented by a ridge; the ventral lobe of the vinculum has no transverse ridge. The holotype of this species has suffered partial bleaching to a light creamy brown colour either as a result of natural homoeosis or, more probably, chemical action. The apex of each wing is affected, bleaching of the left fore wing extending over most of the wing with the exception of the base of the costa. DISTRIBUTION. Chile. MATERIAL EXAMINED. Holotype ^, CHILE: Cautin, Temuco, Carillanca Experimental Station, 3.vii.i975 (Salas) (genitalia slide no. 13147; BMNH). REVISION OF CALLIPIELUS BUTLER n ? Callipielus salasi sp. n. (PL i, fig. 6; PI. 3, fig. 15; PI. 4, fig. 23; PI. 6, figs 33, 37) v$ v""Uv. 4 ^ : ;wv v 3 $?$?> 22 24 .^,, r 25 \ PLATE 5 Aedeagus of male FIG. 26. Callipielus arenosus Butler [holotype of leukogramma], Argentina. Female genitalia FIG. 27. Callipielus sp. indet., Chile. FIG. 28. C. arenosus Butler [paratype of leukogramma], Argentina. FIG. 29. C. izquierdoi (Ureta), allotype, Chile. Bull. Br. Mus. nat. Hist. (Ent.) 35, 4 PLATE 5 27 \ PLATE 6 Antennal segments FIG. 30. Callipielus arenosus Butler, g lectotype, Chile. FIG. 31. C. digitata sp. n., <$ holotype, Chile. FIG. 32. C. brunnescens sp. n., <$ holotype, Chile. FIG. 33. C. salasi sp. n., paratype, Chile. FIG. 34. C. izquierdoi (Ureta), g, Chile. FIG. 35. C. argentata Ureta, <$ paratype, Chile. FIG. 36. Callipielus sp. indet., $, Chile. Labium and labial palpus FIG. 37. C. salasi sp. n., paratype, Chile. Fore leg FIG. 38. C. digitata sp. n., /$ holotype, Chile. Bull. Br. Mus. nat. Hist. (Ent.) 35, 4 PLATE 6 37 " ENTOMOLOGY SUPPLEMENTS 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, 1965- 3-25- 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-i5. 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. 3.40. 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. 19. CROSSKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. 6.50. 20. VON HAYEK, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. 12.30. 21. CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. 9-55- PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND