S&M loz° HISTORY MUSEU : ~ 9 MAY 2012 PRESENTED GENERAL LIBRARY | Geology Series S)2 THE NATURAL HISTORY MUSEUM VOLUME 56 NUMBER1 29 JUNE 2000 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Geology Series is edited in the Museum’s Department of Palaeontology Keeper of Palaeontology: | Prof S.K. Donovan Editor of Bulletin: Dr M.K. Howarth Assistant Editor: Mr C. Jones Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. 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(Geol.) © The Natural History Museum, 2000 Geology Series ISSN 0968-0462 Vol. 56, No. 1, pp. 1-83 The Natural History Museum Cromwell Road London SW7 5BD Issued 29 June 2000 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Lond. (Geol.) 56(1) Issued 29 June 2000 The history, geology, age and fauna (mainly insects) of Burmese amber, Myanmar CONTENTS 1. A review of the history, geology and age of Burmese amber (Burmite), by V.V. Zherikhin & A.J. ROSS ......cccccscecceeseesseeess 3 2. A list of type and figured specimens of insects and other inclusions in Burmese amber, by A. J. Ross & P.V. York......... 11 3. A preliminary list of arthropod families present in the Burmese amber collection at The Natural History Museum, BOM AO MaDe Ate WIA STULESIN IU ker Aap INOS S vaste sre acto e ane Sees as ssa ds Se eRae ue oe ce ns ada So asc cs Ua oOE Soe sec Suen dation dea saudteeeereoo¥s 21 4. The first fossil prosopistomatid mayfly from Burmese amber (Ephemeroptera; Prosopistomatidae), by DMS TUS IACTUKOV Ge Rarer reece Se sete eee eee oe ee aa kee soak Cee ace ne asc ee T scs Sea nani caobaw nates Pats Sunk coutesedaeetscetusbeesdiveuancensceds 25 5. The most primitive whiteflies (Hemiptera; Aleyrodidae; Bernaeinae subfam. nov.) from the Mesozoic of Asia and Burmese amber, with an overview of Burmese amber hemipterans, by D.E. Shcherbakov .......:..:ccsccecceseeseeseenseeseeseeneenees 29 6. Anew genus and species of Lophioneuridae from Burmese amber (Thripida (=Thysanoptera): Lophioneurina), ON WW, ZAGAT ceca Beer cece eee ace ci ee cae EEE EASE oa SESS Ea cee es CELE EEE EC CE 39 7. Burmapsilocephala cockerelli, anew genus and species of Asiloidea (Diptera) from Burmese amber, [BN Sie) Dol OOTY CITTE (o°2 NU ET 2 ANT (OSHONIS Gi otc ecceiaceceraciceccecaencccoc se Eee ee crc CoS eC CORO CESCOCCER ee EES CCES RCRe EEE aE oe 43 8. Phantom midges (Diptera: Chaoboridae) from Burmese amber, by E. D. Lukashevich ..........:ccccccecccecceeseeneesseeneeeneeeaeenes 47 9. An archaic new genus of Evaniidae (Insecta: Hymenoptera) and implications for the biology of ancestral evanioids, bye PES BaSiDUV UK ASPERGSTIDSYNWVIAG SHULtOnTCc DLT. OULCKC ara wcerces sacaen-cceaane see sanensonceee seen detsreensestees-evesceeenneretores 53 10. Digger Wasps (Hymenoptera, Sphecidae) in Burmese Amber, by A.V. ANtropov ............:csscsscesceesseseceeeeeeeneeaceneeeeeeneeaeeaees 59 11. Electrobisium acutum Cockerell, a cheiridiid pseudoscorpion from Burmese amber, with remarks on the validity ofthe Chemdioidea(Arachniday Chelonethi) by MMU JUGS OM eae .cencs. 2 scence cea cecases ence ene ce-scencenacsenesuee saeeenacensosecesctaseneeeroes 79 si NATURAL HISTQSY 2AUSEUM - 9 MAY 2012 PRESENTED GENERAL LIBRARY we ogeex dee nales 4 ime ied 4 ' a ee gn a ty inc yeaa Pe we Fe —ndliowe | is aiken Minne denies othe cuptt muagerathonar + O nna: vat eaty Star - ciaerenrha : Pid 1s - ae sl - Fsmewite AfkeriEgr shiners asl eee : ; | Esl pines teeta 8 eA Ce a roe te sien ade Shibcies (tects fre Zomp ite pert =" nm’ = ee en | nite fe Lak C De work's. ; =f ae 7 er yee ; j We a ma : e | y ee = 5 = = : : : — 1 ~ _ - a : : ; ; 7 y { . t ¥ _ = , — A Pi _ c ; " ear pt ae i danas? hee é i : ; _ SS 7 >, ; = . - . qT ; ; } ~ : f ne ope 2 ot / i . ? t ~ y *, “J t ie { e - 3 . ca hac meaieelo neers ha ars pn ee ee poly ys ‘ x Bull. nat. Hist. Mus. Lond. (Geol.) 56(1): 3-10 Issued 29 June 2000 A review of the history, geology and age of Burmese amber (Burmite) V.V. ZHERIKHIN Arthropod Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya Str. 123, Moscow 117647, Russia A.J. ROSS Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK SYNOPSIS. An overview of the present state of knowledge of Burmese amber is given based on an exhaustive literature search. Early Chinese literature suggests Burmese amber has been known since the 1“ century AD and was later traded from northern Burma to Yunnan Province. Amber has been recorded from five regions in Burma (Myanmar): the Hukawng Valley, Shwebo District, Thayetmo District, Pakokku District and Pegu District, but only Burmite from the Hukawng Valley in northern Burma has been commercially mined. The first visit by a westerner to the amber mines in the Hukawng Valley was by Captain Hannay in 1836. Subsequent visits by members of the Geological Survey of India showed that the amber-bearing deposits are of Middle Eocene age and consist of shales and sandstones with subordinate limestone and conglomerate horizons. The archaic insects in Burmite and the presence of derived clasts in the amber-bearing sediments suggest that the amber has been reworked and is probably of Upper Cretaceous age. INTRODUCTION Amberis fossilized plant resin. There are many different types which may contain the remains of organisms that were trapped by liquid resin. They provide much palaeontological information, particularly about small terrestrial animals which are rarely preserved in sediments. The inclusions in amber, such as insects, often show very delicate morphological details which can rarely, or never, be seen on other kinds of fossils. For a long time, Baltic amber was the only fossil resin extensively studied palaeontologically while records of inclusions in other resins were scarce and mostly anecdotal. The situation changed during the last decades when systematic studies of organic inclusions in various fossil resins were developed (see Poinar, 1992 for a review). However, many kinds of amber are still poorly investigated. Burmese amber, or Burmite is one of the most mysterious vari- eties of fossil resins in the world. Though known for nearly 2000 years in southeastern Asia and for four centuries in Europe, it is poorly investigated scientifically, first of all because of the location of its deposits in aremote area of Upper Burma which was never easy to reach by a traveller. In this area there were the so-called ‘unadministrated territories’ in colonial times and later, in independ- ent Burma (now Myanmar), this area, situated between the “Golden Triangle’ and the politically unstable Nagas region of India, remains controlled rather by the local clans and insurrectionists than by the central government in Rangoon. Few geologists have visited this area, each for a short time only, and the published data on the amber occurrences are scarce. A comprehensive review of the literature has been undertaken and this paper outlines the present state of know- ledge of Burmese amber. THE HISTORY OF BURMITE 0-1800 AD The most thorough account of the early history of trade in Burmese amber is by Laufer (1906), summarized by Penzer (1922) and © The Natural History Museum, 2000 Fraquet (1987). Laufer published translations of passages referring to amber from ancient Chinese literature. He considers that the Chinese became familiar with Burmese amber in the 1* century AD and that trade in amber between Burma and Yunnan Province be- came established soon after. This was based on the Hou Han shu (Annals of the Later Han Dynasty, 205-265 AD), which mentioned amber in Ai lao, the ancient Shan kingdom in Yunnan which formed in the 1 century AD. Laufer indicated that mentions of amber from Yunnan in the Chinese literature became more frequent during the following centuries. The production of amber in Yung ch’ang in Yunnan is recorded in the Kuang ya (a dictionary from 227—240 AD), the Po wu chi (Records of remarkable objects, from 232-300 AD) and the Pieh lu (a medical book of uncertain age, referred to in a later work of 452-536 AD). Laufer (1906) however, considered that the amber from Yunnan originally came from Burma as Yung ch’ ang is only 180 km from the amber mining area in Burma. Pumpelly (1867) also recorded amber from Yungchang in Yunnan, and also from other localities in Shansi, Shensi, Szchuen, and Kwangtung provinces; so it appears that China does have its own amber deposits. Laufer (1906), however, was sceptical about reports of native Chinese amber and considered that they may refer instead to other resins. It should be noted that European amber was being imported into China at least as early as the 18™ century and that the Chinese had the ability to fake amber from early times (Laufer, 1906). The very large piece of Burmese amber figured by Grimaldi (1996, p. 41) and Ross (1998, p. 14, fig. 33) (Natural History Museum (London), Mineralogy Department, specimen no. 1940,37) was originally purchased in Guangzhou (Canton), China in 1860. D. Grimaldi and A. Shedrinsky (pers. comm.) have confirmed that it is Burmese amber using pyroly- sis-gas chromatography. An analysis of the old Chinese literature together with archaeological data may elucidate the relative impor- tance of Burmese amber in ancient and medieval China but at present this question seems to be unclear. It appears that there are no reviews of records of amber from old Burmese written sources which would be expected. It may be stated that Burmese amber was known in China for a very long time, perhaps since the 1* century AD, and that the export of amber from Burma to Yunnan continued for many centuries. According to Laufer (1906), the first European who men- tioned Burmese amber was the Portuguese Jesuit Father Alvarez 4 Semedo who visited China in 1613. In the English edition of his book on China, published in 1655, he wrote that the amber from Yunnan is ‘digged out of mines, and some times in great pieces: it is redder than our Amber, but not so cleane.’ In 1738 Du Halde also mentioned a red amber from Yunnan (Laufer, 1906). All these early records were quite brief and contained little information about the amber itself. 1800-1885 The first short description of Burmese amber was published by Brewster (1835) who investigated one large piece and pointed out that ‘its general aspect and physical properties seem to differ consid- erably from the ordinary specimens of amber’. In particular, the presence of fissures filled by calcite was noted for the first time. Two years later Pemberton (1837) published an extract from the journal of Captain S.F. Hannay who was the first European to visit the amber mines in 1836 and described them briefly. Hannay men- tioned two amber sites in the Hukong (more correctly, Hukawng) Valley, one called ‘Payen-toung or amber-mine hill’ where the amber is mined and another on the east side of the valley called Kotah-bhum and noted that the latter site ‘is considered sacred by the Singphos, who will not allow the amber to be taken away, although it is of an inferior description’ (Pemberton, 1837: 270). Hannay wrote that the amber is the only subject of barter with the Chinese in this area and its price varies considerably in relation to its quality. He also described the amber mines visited personally by himself as pits of 2—4.5m in depth, dug in a reddish and yellow clay with a smell of coal tar. The only instruments used by miners were ‘a bambu sharpened at one end, and a small wooden shovel’ (Pemberton, 1837: 274), and Hannay pointed out that a low production of amber is the conse- quence of these primitive techniques. He goes on to say ‘the deeper the pits are dug, the finer the amber; and that that kind which is of bright pale yellow, is only got at the depth of forty feet under ground.’ Hannay observed also a number of abandoned pits. Pemberton’s (1837) publication is accompanied with a map showing Hannay’s route. Ten years later another description of the amber mines was published in a collection of posthumous papers by Griffith (1847) who accompanied Captain Hannay to the same area in 1837, one year after Hannay’s first visit. Chapter IV (Griffith, 1847: 60-81) gives a detailed account of their ‘Journey from Upper Assam towards Hookhoom, Ava, and Rangoon’ and all the wildlife encountered. He refers to the Hookhoom Valley rather than Hukong and figures a sketch of the valley. Griffith (1847: 77) described the amber mines ‘situated on a range of low hills, perhaps 150ft [46m] above the plain of Meinkhoon, from which they bear S.W.’ His observations are similar to those of Hannay except Griffith adds more detail. Griffith (1847: 77) describes the pits as ‘square, about four feet [1.2m] in diameter, and of very variable depth; steps, or rather holes are cut in two of the faces of the square by which the workmen ascend and descend.’ Griffith does not mention a restricted sacred area. He supplemented Hannay’s observations by the first short description of the rock sequence; he wrote that ‘The soil throughout the upper portion, and indeed for a depth of 15 to 20 feet [4.6-6m], is red and clayish, and appear to inclose but small pieces of lignite; the remain- der consists of greyish slate clay increasing in density as the pits do in depth: in this occur strata of lignite very imperfectly formed, which gives the grey mineral a slaty fracture, and among this the amber is found’ (Griffith, 1847: 78). The deepest pit seen by Griffith was ‘about 40 feet [12m], and the workmen had then come to water.’ The amber he saw ‘occurred as very small irregular deposits, and in no great abundance’. He believed that the miners ‘have no index to favourable spots, but having once found a good pit they of course dig V.V. ZHERIKHIN AND A.J. ROSS as many as possible as near and close together as they can.’ The passage concerning the amber mines was reproduced later by Noetling (1893a). In 1852 and 1853 Piddington described under the name of Hircine a dark brown fossil resin which Penzer (1922) indicates came from the Pakokku District. This was the first information about the occur- rence of a fossil resin in Burma outside the Hukawng valley. In the Mineralogy Department of The Natural History Museum (London), there is a large rounded piece (no. 54956) which is labelled as coming from Pegu in the Pegu District. It was transferred from the India Museum in 1879. Amber from Pegu has not (as far as the authors are aware) been mentioned in published literature. It is possible that the label is incorrect and that it came from the Hukawng Valley (see below). Watt (1908: 64) indicates that in 1876 there were only a few amber workers in Upper Burma. Ball (1881: 57-58) summarized observ- ations made by Hannay and Griffith and also mentioned unpublished notes by Bayfield about his visit to the amber mines area. He believed that the mines ‘in the valley of the Hukung or Hookoon . . . have been worked with varying success for a very long period and have, it is believed, produced a large quantity of amber.’ Mason & Theobald (1882: 15) say very little about Burmese amber. They mention Griffith’s expedition and publish a quote by Anderson — ‘the amber most valued at Momien is perfectly clear, and of the colour of very dark sherry, and is sold by its weight.’ Up until 1885 the main trade route would still have been from Upper Burma to Yunnan in China. 1885-1947 In 1885 the British deposed the Alaungpaya dynasty and Burma became an annex of India. The commercial interest in Burmese amber increased and the market displaced from China to Mandalay and Rangoon in Burma. The Geological Survey of India sent Dr. Fritz Noetling to investi- gate the amber mining area in northern Burma. The results were published in several papers containing the first detailed description of the amber occurrences (Noetling, 1892, 1893a) and summarized briefly in the annual report of the survey (King, 1893). Some amber samples taken by Noetling were examined by Otto Helm, who assigned Burmese amber from the Hukawng Valley to a new mineral species, called by him Burmite (Helm, 1892, 1893a, 1894, 1902). Noetling (1893a) and Helm (1892) were reprinted as a separate article in Rangoon (Noetling, 1893b, Helm, 1893b), along with a map illustrating the position of the mining area. Noetling (1893a) visited the same part of the Hukawng Valley (he called it Hukong) which was described earlier by Hannay and Griffith (Fig. 1). The mines seen by Noetling were situated south-west of Maingkhwan village (or Maingkwan, called Meinkhwon by Hannay and Meinkhoon by Griffith) at a low hill range, 8 km long, trending north-south which he called Nangotaimaw. The position of the range was fixed at Latitude 26° 15' N, Longitude 96° 30' E; (Maingkhwan lies at 26° 20' N, 96° 36' E). Like Hannay and Griffiths, Noetling could see only quite a small quantity of amber during his visit. He was surprised because at that time the amber trade in Mandalay was already well developed. However, Noetling was informed by Chi- nese traders that the main mines are at the southern end of the range near Lalawng (Lalaung) village; he had no possibility to prove this record but considered this information as very probable. Noetling (1893a: 35) found that the Nangotaimaw range consists ‘of a blue, more or less hard clay, dipping at a high angle (88°) towards west.’ He failed to discover any fossils in the clay but assigned it tentatively to the Lower Miocene because of a great similarity with the blue REVIEW OF BURMESE AMBER Younger Tertiary Eocene Cenomanian Basement Fig. 1 Map of the amber mining area in the Hukawng Valley, northern Myanmar showing the general geology, the main town of Maingkwan (black dot) and the amber mines (white dots) for which co-ordinates are known — 1 Khanjamaw, 2 Lajamaw and 3 Ladummaw. clays of the Chindwin Group. It should be noted, however, that Noetling found a pebble containing an ammonite but could not established its origin; Griesbach (1892) supposed that it was of Cretaceous age and probably originated from the conglomerates of the amber-bearing formation. Noetling (1893a) found that the differ- ent colour of the upper part of the clay deposits, as noted by Griffith (1847), is the result of weathering, and that the amber from this weathered zone is dull and brittle and had a thick oxidized crust. According to Noetling’s (1892, 1893a) observations, the amber is distributed unevenly and forms pockets in the clay, but the position of the pockets cannot be determined before digging. He also noted that the pieces of amber usually have a pebble-like shape and supposed that the amber was transported by water and ‘have under- gone a certain amount of wear and tear before they were deposited at their present resting-place’. The techniques of amber digging described by Noetling was the same as observed by Hannay and Griffith. He was rather sceptical about the economic potential of the amber; in his view Burmese amber could not compete with Baltic amber in the world market because of its ‘colour inferior to that of [Baltic] amber’ and especially because of the presence of fissures filled with calcite. Moreover, at this time the imported Baltic amber already began to replace Burmese amber even in Mandalay where it was known as ‘Indian’ amber and was even cheaper than Burmese amber. Noetling (1893a) noted the presence of insect and wood inclusions in Burmese amber but he did not indicate whether they 5 were found in the samples collected directly in the Hukawng valley or seen in Mandalay. Noetling (1893a) mentioned two more areas, Mantha in the Shwebo District and Yenangyat in the Pakokku District, where some pieces of an amber-like resin were discovered. In Mantha small pieces of a brittle resin were found in abundance in a hard coaly clay of Miocene age. According to Noetling, the resin from Mantha resembles Burmite in appearance but he wrote nothing about the single sample from Yenangyat which was seen by him. He also noted ‘numerous Miocene fossils of marine type’ found in Yenangyat together with the resin. Helm (1893a), who examined the resin samples sent by Noetling, gave a short description of their external features but could not establish whether they are identical with Burmite because the quan- tity of resins was not enough for a chemical investigation. Grimes (1898) also mentioned amber from Yenangyat, which came from thin beds of yellow sandstone within shales. The sandstone beds also contained fragments of fossilized wood. Penzer (1922) indicated that this amber came from Kyun Kyaung stream, 5 km north of Seikkwa (21° 8', 94° 51') and is of Miocene age. Meyer (1894) in connection with the investigation of archaeologi- cal amber materials from Europe published some additional data on the chemistry of Burmite and supposed that this kind of amber may have been exported to Europe in antiquity. Dahms (1901) summa- rized the data on the chemical composition of Burmite in his review of the chemistry of fossil resins; he noted that the results of analysis of two samples differed considerably and supposed that more than a single mineral species may be present in the Burmese material. The Geological Survey of India published yearly production figures of Burmese amber from the Myitkyina District for the years 1898-1940 (Holland, 1905; Holland & Fermor, 1910, 1915; Pascoe, 1921, 1925, 1930; Heron, 1935 and Clegg, 1954). The last report was probably disrupted by the 2™ World War. The figures from the reports are summarized in Table 1. It is evident that the annual production of Burmese amber varied considerably during this period, from 36 kg in 1933 up to 10973 kg in 1906 (there are no data for the years 1931 Table 1 The annual production of Burmese amber in the Myitkyina District, Hukawng Valley, Burma, during the years 1898-1940, from the reports of the Geological Survey of India. Year Quantity Year Quantity Ke Kg 1898 5791 1920 3658 1899 1016 1921 1336 1900 457 1922 183 1901 4928 1923 2433 1902 1524 1924 4536 1903 1880 1925 818 1904 4369 1926 2007 1905 6401 1927 3586 1906 10973 1928 1499 1907 2235 1929 996 1908 2489 1930 107 1909 ISVS 1931 - 1910 3200 1932 584 1911 711 1933 36 1912 1372 1934 203 1913 508 1935 965 1914 660 1936 1626 1915 584 1937 1981 1916 279 1938 _ 1917 3002 1939 304 1918 147 1940 1321 1919 376 Total 82656 Average per year = 1922 kg. 6 and 1938). The total amber production for the 43 years was 82656 kg, and the average annual production 1922 kg. Each report also contains brief information on the occurrences and physical and chemical properties of Burmese amber, based nearly exclusively on the papers by Noetling and Helm. Holland & Fermor (1915, p. 231) in the report for the years 1909-1913 attributed the decrease in production ‘partly to troubles in China with a resultant decreased demand and partly to the rival attractions of rubber plantations and jadeite mining’; there are no attempts to analyse the causes of the fluctuation in amber production in other reports. In 1916 Cockerell published the first paper containing descrip- tions of insect inclusions in Burmese amber, sent to him by R.C.J. Swinhoe of Mandalay. Swinhoe, in Cockerell (1916) considered they were of Miocene age. More arthropod inclusions were sent to Cockerell by Swinhoe in the following years and described in series of papers (listed by Ross & York, this volume). Cockerell (1917a, b, c) noticed that the insects were primitive and suggested that the amber was derived from elsewhere and may be much older, even Upper Cretaceous. The materials were presented after description to the British Museum (Natural History) during the years 1919-1921. A brief review of preliminary palaeoentomological data was pub- lished by Burn (1918) and Cockerell (1920). In 1920 the amber mines area was visited by Dr M. Stuart as he accompanied the Hukawng Valley Railway Survey. The most import- ant new find was a piece of rock containing the foraminiferan Nummulites biarritzensis d’ Archiac of Lower Kirthar (Eocene) age, first reported by Fermor (1922). In Stuart’s (1923a) full account of his geological observations he wrote that during his visit there was no amber being dug and that he could add nothing to Noetling’s observations on the amber mines. He noted that the eastern flank of the hill range, where the amber mines occur, consists of grits and conglomerates belonging to the basal beds of the Tipam Series, which are underlain by the blue amber-bearing clay. Just below the base of the Tipam conglomerates he observed numerous pits dug for flint which occurs as nodules in a thin layer of a chalky limestone. Stuart failed to find this limestone exposed but inspected numerous fragments around the pits and discovered here the single piece containing Nummulites biarritzensis. This find led him to conclude that the lowest exposed amber-bearing clay beds were of Eocene age. Cockerell (1922) in his letter to Nature agreed with Stuart’s conclusion. Cockerell also pointed out that besides the typical red Burmite in the collection sent by Swinhoe there were specimens of a pale yellow resin quite different both in the appearance and in the content of inclusions. In particular, he found in this copal a bee which seems to be identical with the living species Trigona laeviceps Smith. Cockerell concluded that the pale resin is very recent, not older than Pleistocene in age. He wrote that Swinhoe was in doubt on the origin of this resin and that it may have come from China. Cockerell (1922) also indicated that the bee described by him under the name Meliponorytes (?) devicta Cockerell (1921) is enclosed in this pale resin. Cockerell’s letter suggested to Stuart (1923b, 1925) an original hypothesis about the genesis of the Burmese resins. He drew attention to the fact that some modern stingless bees, and in particular T- laeviceps, in southeastern Asia use Dipterocarpus resin and oil for the construction of their nests, producing a substance called dammar. Stuart hypothesized that the pale resin (copal) con- taining bee inclusions may be fossil dammar. He suggested that Burmite may have had a similar origin. The presence of many other kinds of insects in both the copal and amber negates this theory. Other published works from about this time (e.g. Coggin Brown, 1924) summarize what was then known of Burmese amber from earlier publications. However, Cotter (1924) records specimens of brown amber found by Clegg near Mibauk, Thayetmo District (19° V.V. ZHERIKHIN AND A.J. ROSS 27'N, 94° 53' E). These pieces came from stringers of lignite 0.03m thick. Williamson (1932) records three large specimens of Burmese amber, one on display at the Wembley Exhibition of 1924 which belonged to the King of Burma, and two in the possession of Messrs. Liberty. In 1930 the Hukawng valley was visited by Dr H. L. Chhibber. His observations were summarized in the annual report of the Geological Survey of India (Fermor, 1931) and described in more detail in his book on the mineral resources of Burma (Chhibber, 1934). The latter work is the most detailed description of Burmite occurrences avail- able and is summarized below. Chhibber recommended the preparation of a large scale map of all amber workings and to introduce more advanced techniques of mining to improve the amber mining industry. Chhibber published photos of the amber mines for the first time, two of these were reproduced in Grimaldi (1996). After 1947 Chhibber’s (1934) relatively detailed report on the amber mines of the Hukawng Valley was the last one published. In 1947 Burma gained full independance and little has been published since then. Later publications summarized information from earlier works or discussed the taxonomic position of some previously described insects. One later work (missed by most subsequent authors) that made a significant contribution is Sahni & Sastri (1957) who pub- lished observations by Eames on derived limestone clasts from the amber mines which contain the foraminiferan Orbitolina birmanica of Cenomanian age. The most recent review of the geology of Burma (Bender, 1983) contains no new data except for some structural observations based on satellite photographs. In 1989 the country was officially re-named Myanmar. Burmese amber is discussed in nearly every major publication concerning amber (e.g. Schlee & Glockner, 1978; Fraquet, 1987; Poinar, 1992; Grimaldi, 1996; Ross, 1998) but this is based on the previously published literature. Fraquet (1987), Grimaldi (1996) and Ross (1997, 1998) include photographs of specimens of Burmese and Chinese amber. Sometimes it is stated that the amber mines in the Hukawng Valley are now exhausted but there is no information available about their present state (Fraquet, 1987). GEOLOGY OF THE AMBER DEPOSITS IN MYANMAR As can be seen from the historical sketch given above, information about the Burmese amber deposits is limited. The summary of the geological data below is based mainly on Chhibber (1934) and Bender (1983). There are five regions in Myanmar (formerly Burma) where fossil resins have been found: the Hukawng Valley in Myitkyina and Upper Chindwin districts to the north, the Shwebo District in central Myanmar, the Thayetmyo District in the west, the Yenangyat oilfield in Pakokku District, also in the west, and the Pegu District to the south. The samples of fossil resins from Myanmar that have been studied differ in chemical composition and structure (Dahms, 1901; Tschirch & Stock, 1936; Savkevich, 1980) but detailed locality data is absent. The amber from the Hukawng Valley was described under the name of Burmite (Helm, 1892, 1893a) and the resin from the Yenangyat oilfield under the name of Hircine (Piddington, 1852, 1853). Itis not known whether Burmite occurs anywhere outside the Hukawng Valley. The Hukawng Valley is probably the only commer- cial source of amber in Myanmar, and Burmite from there has been REVIEW OF BURMESE AMBER sold in Mandalay and Rangoon since the late 19th century. The specimens with arthropod inclusions kept in Palaeontology Depart- ment of The Natural History Museum (London) probably originated from this area. The Hukawng Basin is a syncline approximately 100 km in diameter (Bender, 1983) (Fig. 1). The sediments dip from 5—20° towards the centre of the basin. Crystalline rocks of the Kumon Ranges form the basement of the basin on the eastern flank, and in the southern and south-western flanks the sediments are bordered by a fault system. Most of the central part of the basin is covered by superficial sediments deposited by the Chindwin River. Three anti- clines occur in the centre of the basin, revealed by satellite images, and an additional anticline is revealed in the western flank of the basin. The largest of the central anticlines is where the amber mines occur. It has a NW-plunging axis with the Eocene amber-bearing rocks exposed in its crest. The eastern limb of the anticline is cut off by a north-south trending fault. In the two smaller central anticlines the crests are formed by limestone of Upper Cretaceous (Cenomanian) age. The amber-bearing deposits were described as clays by many authors, except for Chhibber (1934) who found that they consist mainly of finely bedded dark-blue shales with thin beds of sandstones of various colours (light-blue, pink, grey, ‘salt-and-pepper’), and subordinate limestone and conglomerate beds. Chhibber observed that the beds had been thrown into tightly compressed anticlinal and synclinal folds, so the amber bearing hill range may be structurally more complex than Bender (1983) outlined. Chhibber believed that the limestone fragment with Nummulites biarritzensis collected by Stuart originated from a limestone layer in this formation. The only other fossils recorded by him are some unidentifiable plant impres- sions. The blue colour of the sediments changed to red or yellow in the weathered zone. The sandstones and shales contain very thin lignite seams and the amber is associated with the lignite. Chhibber stated that good quality amber is found mainly at a depth of 10—15m; the amber also occurs higher up but is of inferior quality. The amber pieces may be elliptical, oval or rounded but never irregular or angular in shape. Large blocks of amber are rare but do occur. When a particular layer of sand was reached the diggers stopped mining because they believed that there was no amber beyond it. Chhibber (1934) lists thirteen amber sites in the Hukawng Valley, but unfortunately does not give co-ordinates for most of them. The most active site that he visited was Khanjamaw (26° 15' 50", 96° 33' 37") (Fig. 1). Other sites that he does give co-ordinates for are Ladummaw (26° 11' 19", 96° 28' 4") and Lajamaw (26° 15', 96° 28'). The Nangotaimaw site which was visited by Hannay and Noetling was deserted and Chhibber saw about 200 old pits there. Chhibber observed that the mines were 1.1m square and reached a maximum depth of 14m. The deep pits were lined by bamboo. Water often required bailing out at a depth of 12m. Digging was done with a wooden crowbar tipped with iron and the material was hauled up to the surface in baskets where it was examined for amber. Clearly this is the same technique that was observed by Hannay and Griffith nearly a hundred years before and it is likely that it had been employed for many centuries. The main study on the physical and chemical properties of Burmite was by Helm (1892, 1893a). He noticed that it varied from yellow to brown, with a brown crust. It broke conchoidally and fluoresced blue. This latter character has been mentioned by several authors but Fraquet (1987) states that she never saw this in any specimens that she examined. It is likely that the fluorescence gradually disappears due to oxidation caused by exposure to air; a similar phenomenon was observed by one of us (V.Z.) with Sakhalin amber. Burmite is slightly harder than Baltic amber (Succinite), measuring 2.5—3 on 7 the Moh’s scale. Several authors mention that the amber commonly has thin veins of calcite running through it; however, none of the specimens at the Natural History Museum show this feature. Burmite consists of 80.05% Carbon, 11.5% Hydrogen, 8.43% Oxygen and 0.02% Sulphur and has a specific gravity of 1.030—1.095 (Helm, 1893a). THE AGE OF BURMITE There is considerable confusion in the literature as to the probable age of Burmite. Noetling (1892) was the first to suggest a Lower Miocene age of the amber-bearing deposits of the Hukawng Valley. Cockerell (1917c) considered that the insect fauna in the amber was much older and could be Upper Cretaceous. Stuart (1923a) found the foraminiferan Nummulites biarritzensis in a limestone bed in the amber-bearing strata which was confirmed by Chhibber (1934). This led Stuart to conclude that the beds were of Eocene age, and Cockerell (1922) considered that the insect fauna in the amber was also of this age. Grimaldi etal. (1997) suggested it could be Paleocene or early Eocene based on its physical features. Several authors discussing the Burmite insect fauna stressed that it appears to be too archaic for the Eocene and thus may be of an older age, perhaps even Upper Cretaceous (Zherikhin, 1978; Dlussky, 1996; Rasnitsyn, 1996b; Ross, 1997, 1998). Dlussky (1996) and Rasnitsyn (1996b) considered that the Burmese amber insects could represent a relict fauna that survived into the early Tertiary. Rasnitsyn (1996a) indicates that the fauna is not tropical, which is not what would be expected. Recent studies on the arthropods in the Burmite collection in the Department of Palaeontology of The Natural History Museum, London have revealed five insect families or subfamilies that are not known later than the Cretaceous elsewhere. They are the ant sub- family Sphecomyrminae (species described by Dlussky, 1996; although Grimaldi et al., 1997 expressed doubt about it’s subfamilial position), the wasp subfamily Iscopininae (mentioned by Rasnistsyn, 1996b), the wasp family Serphitidae (figured in Ross, 1997), the thrip family Lophioneuridae (new species described by Zherikhin, this volume) and the whitefly subfamily Bernaeinae (new species described by Shcherbakov, this volume). These taxa are recorded from other ambers of Cretaceous age (see Poinar, 1992; Grimaldi et al., 1997). Of note is that there are no extant subfamilies of ants present, which are very common in Tertiary ambers, although one ant described by Dlussky (1996), Burmomyrma rossi, was placed in incertae subfamiliae but may belong to the Aneuretinae. This evi- dence would suggest that the amber could be of Upper Cretaceous age, but this does not rule out the possibility that the fauna is relict and could still be Tertiary in age. Clearly there is a conflict here in that the presence of Nummulites biarritzensis suggests a Middle Eocene (upper Lutetian—Bartonian) age for the amber-bearing deposits (see Racey, 1995), but the insects suggest an older age for the amber. This could imply that the amber has been reworked. In the literature there is some evidence to support this. Noetling found a pebble containing an ammonite which Griesbach (1892) considered was Cretaceous and came from a conglomerate within the amber-bearing sediments. More impor- tantly, Sahni & Sastri (1957) published observations by Eames who found sub-angular fragments of a hard dark greenish-grey limestone, within a soft orange-cream foraminiferal limestone from the amber mines. He found that the hard limestone fragments contained the foraminiferan Orbitulina birmanica which Sahni & Sastri (1957) considered is of Cenomanian age. Clearly there were Cretaceous Fig. 2 Rounded pebble of Burmese amber with pits caused by impact during transportation. Department of Mineralogy, NHM 54956. Scale bar in millimetres. UU Fig.3 Unpolished surface of a slab of insect-bearing Burmite with pits caused by impact during transportation. Department of Palaeontology, NHM In.19104—6. Scale bar in millimetres. V.V. ZHERIKHIN AND A.J. ROSS deposits being eroded during the Middle Eocene and clasts from them were being deposited in the amber-bearing sediments. The shape of the amber pieces themselves also suggest that they have been reworked. Both Noetling (1892, 1893a) and Chhibber (1934) observed that the amber pieces were rounded like pebbles and not angular, which would suggest that prior to deposition the amber was — 90E 100E 0 500 1000 Kilometers 100E Sea 0 500 1000 Kilometers Low land High land Mountains Figs 4,5 Palaeogeographic maps. 4, for 94 Ma (Cenomanian) showing the position where Orbitulina limestones were being deposited (black dot); 5, for 50Ma (Eocene) showing the position where amber-bearing sediments were being deposited (white dot) and a possible source of derived Cenomanian Orbitulina limestone pebbles (black dot); in both figures the black line in the sea represents the edge of the continental shelf. REVIEW OF BURMESE AMBER already fossilised and underwent transportation that rounded them. The large specimen in the Mineralogy Department of the NHM labelled ‘Pegu’ (no. 54956) (Fig. 2) is dark red, rounded and is covered by pits filled with sediment. These same pits can be seen on the unpolished surfaces of two end slabs (In.19104—6 & In.20150) (Fig. 3) cut from a large insect-bearing block of Burmite in the Department of Palaeontology (NHM). Closer examination shows that some of these pits have conchoidal fracture which develops from a single impact point at the edge of the pit. These pits support the suggestion that the amber pieces have been transported. It’s possible that the “Pegu’ piece is mis-labelled and that it is Burmite from the Hukawng Valley, but further tests are required to confirm this. Thus the available evidence suggests that Burmite is reworked and is probably of Upper Cretaceous age. PALAEOGEOGRAPHY From the evidence currently available there appears to be two periods of deposition related to the amber bearing deposits in the Hukawng Valley. The first was during the Cenomanian when Orbitulina limestones were deposited. These limestones are exposed as small inliers forming the cores of two anticlines within the Hukawng Basin (Bender, 1983). When one of the inliers, lying about 20 km south of the mines, is plotted on PaleoGIS for ArcView (Ross & Scotese, 1997-8), at 94 Ma, the map suggests these sediments were deposited in a sea (Fig. 4) at 11° N, 93° E, about 350 km from the nearest land which lay to the north-east and north-west. If Burmite was forming at this time, or later on in the U. Cretaceous, then it would have probably been produced by trees living on this land mass and could have been deposited on the continental shelf or it may even have been transported further out to sea. Later, by the Middle Eocene, these limestones were uplifted and eroded and pebbles from them were deposited within beds of limestone within the amber-bearing deposits. These younger limestones were depos- ited in a shallow sea as evidenced by the presence of Nummulites. The interbedded shales, sandstones, conglomerates and limestones suggests tectonic instability of the area resulting in repeated trans- gressions and regressions. When plotted on PaleoGIS for ArcView (Ross & Scotese, 1997-8), at 50 Ma (the closest age available), the map suggests these sediments were generally deposited on low- lying land (Fig. 5) at about 12° N, 90° E with higher land lying about 50 km to the north-east. The Cenomanian limestone deposit in Fig. 4 lies very close, to the south-east of this area, and could have been a direct source for derived pebbles. The primary deposits containing the amber are not known but it can be assumed that they were exposed nearby. ACKNOWLEDGEMENTS. Many thanks to Mary Chester-Kadwell for help with finding obscure references, to David Lees for help in producing Figs 1 and 2 from PalaeoGIS for ArcView, to David Grimaldi for refereeing the manuscript and to Phil Crabbe for photography. REFERENCES Ball, V. 1881. 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Die botanischen und chemischen Grundlagen unserer Kenntnisse tiber die Bildung, die Entwicklung und die Zusammensetzung der pflanzlichen Exkrete. 2 (2). Borntraeger, Berlin. 1017-1858. Watt, G. 1908. The commercial products of India. John Murray, London: 1189 pp. Williamson, G.C. 1932. The book of amber. Ernest Benn, London. 268 pp. Zherikhin, V.V. 1978. Razvitie i smena melovykh i kainozoiskikh faunisticheskikh kompleksov (trakheinye i khelitserovye) [Development and changes of the Creta- ceous and Caenozoic faunistic assemblages (Tracheata and Chelicerata)]. Trudy Paleontologicheskogo Instituta, 165. Nauka, Moscow. 198 pp. (in Russian). Bull. nat. Hist. Mus. Lond. (Geol.) 56(1): 11-20 Issued 29 June 2000 A list of type and figured specimens of insects and other inclusions in Burmese amber A. J. ROSS Department of Palaeontology, The Natural History Museum, Cromwell Road, South Kensington, London SW7 SBD P.V. YORK Department of Botany, The Natural History Museum, Cromwell Road, South Kensington, London SW7 5BD SYNOPSIS. All type and/or figured inclusions in Burmese amber are listed. Photographs of 24 holotypes, mostly insects described by Cockerell between 1916 and 1920, are published for the first time. INTRODUCTION Below is a list of all described and/or figured specimens of insects and other inclusions in Burmese amber, with a bibliography of references to where the species have been cited. The specimens are housed in the Department of Palaeontology of The Natural History Museum, London. The list of insects follows the higher classification format used by Ross & Jarzembowski (1993). Within each family, the genera and species are listed in alphabetical order, accompanied by the author, specimen number (with the prefix In.), references (in date order), and finally references to the photo- graphs of 24 previously described holotypes that are published here, in colour, for the first time. Cockerell’s (1916-1921) publications are listed in publication date order; however in Cockerell (1917b) species are mentioned before they were formally established. Some large pieces of amber contain several type specimens and historically each type has been given a separate number; therefore a single piece of amber may have arange of numbers. Where this is the case the range of numbers for the piece of amber is given in brackets after the number for the type. One large block containing type specimens In.19102—23 and In.20149—50 was sliced up into 7 slabs (see Cockerell, 1917d). Slab In.19107—16 was figured in Ross (1998: 13, fig. 32). The 7 slabs (and In.20195, whichhad broken off In.20150) contain a total of 458 inclusions, belonging to many different taxa (see Rasnitsyn & Ross, this volume) which makes the whole block the richest piece of amber ever recorded. Where more than one inclusion occurs inthe same piece of amber but are registered under one number, an additional number is given in brackets. Spahr (1985) lists Anthomyia (s.1.) laminarum Cockerell as com- ing from Burmese amber. This is incorrect, it came from Florissant, Colorado, USA (Cockerell, 1917a). LIST OF TAXA AND SPECIMENS INSECTA Order ZYGENTOMA Family LEPISMATIDAE Allocrotelsa burmiticus (Cockerell, 1917) comb. nov. Holotype, In.19119 (In.19117-22). Cockerell (1917d: 360-361, fig. 2). Flerov et al (1974: 104). Zherikhin (1978: 114). Keilbach (1982: 205). Spahr (1990: 17, © The Natural History Museum, 2000 18). Poinar (1992: 96). Replacement generic name for Lampropholis in Carpenter (1992: 18). Fig. 1. Order EPHEMEROPTERA Family PROSOPISTOMATIDAE Myanmarella rossi Sinitshenkova, this volume. Holotype, In.20173. Order DERMAPTERA Family LABIDURIDAE ?Labidura electrina Cockerell, 1920. Holotype, In.20146. Cockerell (1920b: 212, fig. 1). Zherikhin (1978: 114). Keilbach (1982: 210). Spahr (1992: 18, 79). Poinar (1992: 103). Fig. 2. Order EMBIOPTERA Family ‘BURMITEMBIIDAE’ Burmitembia venosa Cockerell, 1919. Holotype, In.19132. Cockerell (1919a: 194-195, figs 2, 3). Davis (1939: 369-372, figs 1-6). Ross (1956: 76; 1963: 123). Flerov et al (1974: 104). Zherikhin (1978: 114). Keilbach (1982: 210). Spahr (1992: 16, 79). Poinar (1992: 107). Carpenter (1992: 190-191, fig. 122, 1). Szumik (1994: 67). The family name was first used by Zherikhin (1980: 78) but has not been formally established. Fig. 4. Order ISOPTERA Family KALOTERMITIDAE Kalotermes swinhoei (Cockerell, 1916). Holotype, In.19096. Cockerell (1916: 138, fig. 4). Fletcher (1920: 988, pl. 166, fig. 23). Snyder (1925: 157, Table 1). Emerson (1933: 190). Armbruster (1941: 41). Snyder (1949: 370). Williams (1968: 547-551, figs 1— 3). Emerson (1969: 30-31, 34-35). Burnham (1978: 89). Zherikhin (1978: 114). Keilbach (1982: 214). Spahr (1992: 30, 79). Poinar (1992: 104). Nel & Paicheler (1993: 120). Fig. 3. Kalotermes tristis (Cockerell, 1917). Holotype, In.19103 (In. 19102— Soo. Cockerell (1917e: 325, 329, fig. 10). Fletcher (1920: 988, pl. 166, fig. 24). Emerson (1933: 190). Armbruster (1941: 40, 41). Snyder (1949: 370). Williams (1968: 547, 549-551, fig. 4). Emerson (1969: 30-31, 35-36). Burnham (1978: 89). Zherikhin (1978: 114). Keilbach (1982: 214). Spahr (1992: 30, 79). Poinar (1992: 104). Nel & Paicheler (1993: 120). Order HEMIPTERA Family ACHILIDAE ‘Liburnia’ burmitina Cockerell, 1917. Holotype, In.19105 (In.19104— 6). Cockerell (1917e: 329, figs 8, 9). Fletcher (1920: 988, pl. 166, figs 21, 22). Metcalf & Wade (1966: 113). Keilbach (1982: 230). Spahr (1988: 19, 48). Poinar (1992: 125). Transferred from Delphacidae (Araeopidae) by Shcherbakov, this volume. Fig. 7. Family ALEYRODIDAE ‘Aleurodicus’ burmiticus Cockerell, 1919. Holotype, In.19134 (In.19133-4). Cockerell (1919b: 241, 243, fig. 1). Larsson (1978: 71). Zherikhin (1978: 114). Keilbach (1982: 237). Spahr (1988: 29, 48). Shcherbakov, this volume, considers that the Aleurodicidae are a subfamily of Aleyrodidae. Burmoselis evelynae Shcherbakov, this volume. Holotype, In.20193. Family CIXITIDAE Plecophlebus nebulosus Cockerell, 1917. Holotype, In.19094. Cockerell (1917e: 327-328, fig. 7). Fletcher (1920: 987, pl. 165, fig. 12). Fischer (1965: 56). Zherikhin (1978: 114). Botosaneanu (1981: 73-75, 78, figs 1-3). Keilbach (1982: 310). Wichard (1984: 443). Spahr (1988: 22, 48; 1989: 58). Poinar (1992: 160). Carpenter (1992: 258). Shcherbakov, this volume. Family ENICOCEPHALIDAE Disphaerocephalus constrictus Cockerell, 1917. Holotype, In.19112 (In.19107—16). Cockerell (1917d: 361-364, fig. 3). Jeannel (1942: 293-294, fig. 22a). Usinger (1945: 340). Stys (1969: 356-358, 360-361, 363, figs 6-9, photos 4-6). Zherikhin (1978: 114). Keilbach (1982: 227). Spahr (1988: 9, 48). Carpenter (1992: 262). Shcherbakov, this volume. Disphaerocephalus macropterus Cockerell, 1917. Holotype, In.19123(1). Cockerell (1917d: 364). Stys (1969: 352, 359-360). Zherikhin (1978: 114). Keilbach (1982: 227). Spahr (1988: 9, 48). Rasnitsyn (1996a: 19). Shcherbakoy, this volume. Fig. 8. Disphaerocephalus swinhoei (Cockerell, 1917d). Holotype, In.19113 (In.19107-16). Cockerell (1917b: 42; 1917d: 364, fig. 4). Jeannel (1942: 294, fig. 22c). Usinger (1945: 340). Stys (1969, 358-360, figs 6, 10, 11, photos 7, 8). Zherikhin (1978: 114). Keilbach (1982: 227). Spahr (1988: 9, 48). Shcherbakov, this volume. Paenicotechys fossilis (Cockerell, 1916). Holotype, In.19095. Cockerell (1916: 135-136, fig. 1). Fletcher (1920: 988, pl. 166, fig. 20). Jeannel (1942: 294, fig. 22b). Usinger (1945: 340). Bekker-Migdisova (1962: 220, fig. 657; 1991: 307, fig. 657). Stys (1969: 353-355, figs 1-5, photos 1-3). Zherikhin (1978: 114). Keilbach (1982: 226). Spahr (1988: 9, 48). Carpenter (1992: 262). Shcherbakoy, this volume. Order PPOCOPTERA Family PACHYTROCTIDAE ?Psylloneura perantiqua Cockerell, 1919. Holotype, In.19136. A.J. ROSS AND P.V. YORK Cockerell (1919b: 241-242, fig. 2). Smithers (1967: 28). Zherikhin (1978: 114). Keilbach (1982: 221). Spahr (1992: 50, 79). Fig. 5. Family Psyllipsocidae ?Psyllipsocus banksi Cockerell, 1916. Holotype, In.19097. Cockerell (1916: 136-138, figs 2, 3). Fletcher (1920: 987, pl. 165, figs 13, 14). Bekker-Migdisova & Vishniakova (1962: 233, fig. 698; 1991: 327, fig. 698). Smithers (1967: 15). Zherikhin (1978: 114). Keilbach (1982: 217). Spahr (1992: 53, 79). Order THYSANOPTERA Family LOPHIONEURIDAE Burmacypha longicornis Zherikhin, this volume. Holotype, In.20194. Order COLEOPTERA (Cockerell’s type specimens have been examined by P. Hammond (NHM) who considers (pers. comm.) that the generic placement of some species is doubtful. These are indicated by single quotes). Family ANTHICIDAE Eurygenius wickhami Cockerell, 1917. Holotype, In.19110 (In.19107-16). Cockerell (1917e: 324-325, fig. 2). Fletcher (1920: 987, pl. 165, fig. 16). Zherikhin (1978: 114). Spahr (1981: 10). Keilbach (1982: 249). Poinar (1992: 135). Ross (1998: 69, fig. 161 (top left)). Family ?COLYDIIDAE Cryphalites rugosissimus Cockerell, 1917d. Holotype, In.19111 (In.19107-16). Cockerell (1917b: 43, 45; 1917d: 368, fig. 8). Zherikhin (1978: 114). Spahr (1981: 57). Keilbach (1982: 255). Poinar (1992: 152). Carpenter (1992: 316). P. Hammond (pers. comm.) considers that this species does not belong to the Scolytidae (=Ipidae) or Curculionidae, but may belong to the Colydiidae. Fig. 9. Family DERMESTIDAE ‘Dermestes’ larvalis Cockerell, 1917. Holotype, In.19108 (In.19107— 16). Cockerell (1917b: 43, fig. 4; 1917e: 323). Fletcher (1920: 988, pl. 165, fig. 19). Zherikhin (1978: 114). Spahr (1981: 44). Keilbach (1982: 248). Poinar (1992: 143). Fig. 10. Family ELATERIDAE ‘Acmaeodera’ burmitina Cockerell, 1917. Holotype, In.19107 (In.19107—16). Cockerelll (917232353255, 3275.529; fig): Fletcher (9202987. pl. 165, fig. 18). Stys (1969: 357, fig. 6). Zherikhin (1978: 114). Spahr (1981: 14). Keilbach (1982: 248). Poinar (1992: 136). Bellamy (1995: 175-176, fig. 1). Ross (1998: 13, 69, figs 32, 161). ‘Elater’ burmitinus Cockerell, 1917. Holotype, In.19102 (In.19102— 3): Cockerell (1917e: 325, fig. 3). Fletcher (1920: 987, pl. 165, fig. LIST OF INCLUSIONS IN BURMESE AMBER 17). Zherikhin (1978: 114). Spahr (1981: 47). Keilbach (1982: 247). Poinar (1992: 144). Fig. 11. Family RHIPIPHORIDAE Myodites burmiticus Cockerell, 1917. Holotype, In.19092. Cockerell (1917a: 22, fig. 6). Fletcher (1920: 987, pl. 165, fig. 15). Zherikhin (1978: 114). Spahr (1981: 87). Keilbach (1982: 250). Poinar (1992: 151). Fig. 12. Order DIPTERA Family APSILOCEPHALIDAE Burmapsilocephala cockerelli Gaimari & Mostovski, this volume. Holotype, In.20167. Family CECIDOMYIIDAE Winnertzia burmitica (Cockerell, 1917b). Holotype, In.19114 (In.19107-—16). Cockerell (1917b: 41-42, 44, fig. 3). Fletcher (1920: 987, pl. 164, fig. 8). Zherikhin (1978: 114). Keilbach (1982: 346). Spahr (1985: 16, 126). Poinar (1992: 168). Evenhuis (1994: 188). Family CERATOPOGONIDAE ‘Johannsenomyia’ swinhoei Cockerell, 1919. Holotype, In.19133 (In.19133-4). Cockerell (1919b: 243, fig. 3). Zherikhin (1978: 114). Keilbach (1982: 350). Spahr (1985: 19, 126). Szadziewski (1988: 11, 26, 181, 240). Spahr (1989: 16). Poinar (1992: 169). Evenhuis (1994: 253). Borkent (1995: 178). Fig. 13. Family CHAOBORIDAE Chaoburmus breviusculus Lukashevitch, this volume. Holotype, In.20168, paratype In.20168(1). ?Chaoburmus victimaartis Lukashevitch, this volume. Holotype, In.20157 Family EMPIDIDAE Burmitempis halteralis Cockerell, 1917. Holotype, In.19121 (In.19117—22), also In.19106 (In.19104—6). Cockerell (1917d: 367-368, fig. 7; 1917e: 329, fig. 6). Fletcher (1920: 986, pl. 164, fig. 7). Melander (1928: 368). Zherikhin (1978: 114). Chvala (1981: 232). Keilbach (1982: 369). Spahr (1985: 40, 126). Poinar (1992: 180). Carpenter (1992: 444). Evenhuis (1994: 357). Grimaldi & Cumming (1999: 44). Electrocyrtoma burmanicum Cockerell, 1917. Holotype, In.19099. Cockerell (1917a: 22, fig. 5). Fletcher (1920: 986, pl. 164, fig. 6). Melander (1928: 368). Zherikhin (1978: 114). Kovalev (1978a: 77; 1978b: 355). Chvala (1981: 233). Keilbach (1982: 370). Chvala (1983: 57). Spahr (1985: 42, 126). Poinar (1992: 180). Carpenter (1992: 430). Evenhuis (1994: 347). Grimaldi & Cumming (1999: 51). Fig. 14. Family KEROPLATIDAE Burmacrocera petiolata Cockerell, 1917. Holotype, In.19104 (In.19104—6). Cockerell (1917e: 326-327, figs 4, 5). Fletcher (1920: 987, pl. 164, fig. 10). Edwards (1929b: 71). Zherikhin (1978: 114). Keilbach (1982: 331). Spahr (1985: 52, 126). Poinar (1992: 172). Carpenter (1992: 415). Evenhuis (1994: 139). Family MYCETOPHILIDAE ‘Sciara’ burmitina Cockerell, 1917. Holotype, In.19100 (In.19100— i): Cockerell (1917a: 20-21, fig. 3). Fletcher (1920: 987, pl. 165, fig. 11). Zherikhin (1978: 114). Keilbach (1982: 343). Spahr (1985: 108, 127). Poinar (1992: 175). Evenhuis (1994: 173). P. Chandler (pers. comm.) considers that this species does not belong to the Sciaridae, but belongs instead to the mycetophilid subfamily Sciophilinae. Fig. 16. Family PSYCHODIDAE Eophlebotomus connectens Cockerell, 1920. Holotype, In.20147. Cockerell (1920b: 212-214, fig. 2). Edwards (1929a: 424, fig.). Tonnoir (1933: 63, fig. 3a). Fairchild (1955: 183, fig. 5). Hennig (1972: 65, fig. 71). Zherikhin & Sukacheva (1973: 44). Flerov et al (1974: 103). Zherikhin (1978: 114). Keilbach (1982: 359). Spahr (1985: 97, 127). Poinar (1992: 174). Carpenter (1992: 405, fig. 221, 8). Evenhuis (1994: 192). Trichomyia swinhoei Cockerell, 1917. Holotype, In.19101 (In.19100— I): Cockerell (1917a: 21, fig. 4). Fletcher (1920: 987, pl. 164, fig. 9). Hennig (1972: 65). Zherikhin (1978: 114). Keilbach (1982: 358). Spahr (1985: 100, 127). Poinar (1992: 174). Evenhuis (1994: 196). Fig. 15. Family THEREVIDAE Psilocephala electrella Cockerell, 1920. Holotype, In.20148. Cockerell (1920c: 170). Hennig (1967: 3). Zherikhin (1978: 114). Keilbach (1982: 365). Spahr (1985: 117, 127). Evenhuis (1994: 321). Gaimari & Mostovski, this volume. Order HYMENOPTERA Family APIDAE Trigona (Heterotrigona) devicta (Cockerell, 1921). Holotype, In.20702. Cockerell (1921: 544-545, fig. 4; 1922: 714). Kerr (1948: 241, 247). Kerr & Maule (1964: 10-13). Kelner-Pillault (1969: 525). Flerov et al (1974: 104). Zeuner & Manning (1976: 222-224, pl. 2, fig. 9). Kerr & Cunha (1976: 35-36, 38-39, figs 14). Wille (1977: 44-45). Burnham (1978: 123). Moure & Camargo (1978: 564-565). Wille (1979: 243). Spahr (1987: 14, 99). Poinar (1992: 215). This specimen is in copal, not amber, Grimaldi et al (1995: 262). Michener (1990: 126) considers Tetragonula is a junior synonym of Heterotrigona. Trigona (Heterotrigona) iridipennis (Smith, 1854) (extant species). In.43809. Cockerell (1922: 714). Stuart (1923: 84; 1925: 480). Salt (1931: 146). Kelner-Pillault (1969: 525). Flerov et al (1974: 103). Zeuner 14 & Manning (1976: 222-224, pl. 2, fig. 8). Wille (1977: 45). Spahr (1987: 15, 99). This specimen is in copal, not amber, Grimaldi et al (1995: 262). Generic combination from Michener (1990: 126). Family BETHYLIDAE Apenesia electriphila Cockerell, 1917. Holotype, In.19109 (In.19107—16). Cockerell (1917b: 44-45, fig. 6; 1917e: 323). Fletcher (1920: 986, pl. 164, fig. 2). Brues (1933: 169). Zherikhin (1978: 114). Keilbach (1982: 269). Spahr (1987: 16, 99). Bethylitella cylindrella Cockerell, 1917. Holotype, In.19120 (In.19117—22). Cockerell (1917d: 365-367, fig. 6). Brues (1933: 168). Zherikhin (1978: 114). Keilbach (1982: 268). Spahr (1987: 17, 99). Carpen- ten :9922487) shies 19: Epyris atavellus Cockerell, 1920. Holotype, In.20149(1). Cockerell (1920a: 274, 276-277, fig. 2). Brues (1933: 168). Zherikhin (1978: 114). Spahr (1987: 17, 99). Fig. 20. Sclerodermus quadridentatum Cockerell, 1917. Holotype, In.19116 (In.19107—16). Cockerell (1917b: 42-45, fig. 5). Fletcher (1920: 986, pl. 164, fig. 3). Brues (1933: 169). Zherikhin (1978: 114). Keilbach (1982: 269). Spahr (1987: 20, 99). Family EVANIIDAE Mesevania swinhoei Basibuyuk & Rasnistsyn, this volume. Holotype, In.20192 Family FORMICIDAE Burmomyrma rossi Dlussky, 1996. Holotype, In.19125. Dlussky (1996a: 87-88, fig. 2a; 1996b: 453, fig. 2a). Haidomyrmex cerberus Dlussky, 1996. Holotype, In.20182. Dlussky (1996a: 83-87, fig. 1; 1996b: 449-453, fig. 1). Grimaldi, Agosti & Carpenter (1997: 24-25) doubt the placement of this genus in the subfamily Sphecomyrminae. Fig. 21. Undetermined. In.20182(1). Dlussky (1996a: 87, fig. 2b; 1996b: 452, fig. 2b). Family GASTERUPTIIDAE (Aulacidae) Electrofoenus gracilipes Cockerell, 1917. Holotype, In.19117 (In.19117—22). Cockerell (1917d: 364-365, fig. 5). Brues (1933: 157). Zherikhin (1978: 114). Keilbach (1982: 263). Spahr (1987: 15, 99). Carpen- ter (1992: 474, fig. 256, 1). Ayptiogastrites electrinus Cockerell, 1917. Holotype, In.19098. Cockerell (1917a: 19-20, fig. 2). Fletcher (1920: 986, pl. 164, fig. 5). Brues (1933: 157). Zherikhin (1978: 114). Keilbach (1982: 263). Spahr (1987: 16, 99). Carpenter (1992: 474). Protofoenus swinhoei Cockerell, 1917. Holotype, In.19091. Cockerell (1917a: 19, fig. 1). Fletcher (1920: 986, pl. 164, fig. 4). Brues (1933: 157). Zherikhin (1978: 114). Keilbach (1982: 263). Spahr (1987: 16, 99). Carpenter (1992: 474, fig. 256, 1). Fig. 22. Family SERPHITIDAE Serphites sp. In.20190. Rasnitsyn (1996a: 20; 1996b: 9). Ross (1997: 24, fig. 2). A.J. ROSS AND P.V. YORK Family SPHECIDAE Apodolichurus sphaerocephalus Antropov, this volume. Holotype, In.20150. Apodolichurus diaphanus Antropov, this volume. Holotype, In.19123(4). Burmastatus triangularis Antropov, this volume. Holotype, In.20197. Cirrosphex admirabilis Antropov, this volume. Holotype, In.19125(1). Cretampulex gracilis Antropov, this volume. Holotype, In.19123(5). Cretospilomena familiaris Antropov, this volume. Holotype, In.19136(1), 8 paratypes, In.19136(2-9). Mendampulex monilicularis Antropov, this volume. Holotype, In.20711. Prolemistus apiformis Antropov, this volume. Holotype, In.20181. Trigampulex pervetus (Cockerell, 1917). Holotype, In.19093. Cockerell (1917c: 79-80, fig. 1). Fletcher (1920: 986, pl. 164, fig. 1). Zherikhin (1978: 114). Spahr (1987: 87, 99). Poinar (1992: 201). Transferred from Trigonalys (Trigonalidae) by Antropov, this volume. Order LEPIDOPTERA Family MICROPTERYGIDAE Sabatinca perveta (Cockerell, 1919). Holotype, In.19135. Cockerell (1919a: 193-194, figs la—f). Rebel (1936: 165). Kuznetsov (1941: 12, 17, 69, 86). Skalski (1973: 650; 1976: 204, fig. 2; 1977: 7). Whalley (1977: 526; 1978: 74-75, 77). Larsson (1978: 124). Zherikhin (1978: 114). Kristensen & Nielsen (1979: 141-142). Keilbach (1982: 313). Kozlov (1988: 28). Spahr (1989: 33, 72). Poinar (1992: 161). Fig. 6. Order TRICHOPTERA Family HYDROPTILIDAE Burminoptila bemeneha Botosaneanu, 1981. Holotype, In.20180. Botosaneanu (1981: 75-78, figs 4-10). Wichard (1984: 443). Spahr (1989: 53, 72). Poinar (1992: 160). Fig. 17. ARACHNIDA Order ACARINA Family CHEYLETIDAE Cheyletus burmiticus Cockerell, 1917. Holotype, In.19115 (In.19107—16). Cockerell(1917b: 41-42, fig. 2). Petrunkevitch(1955: 97). Zherikhin (1978: 115). Morris (1980: 32). Keilbach (1982: 192). Poinar (1992: 225). Spahr (1993a: 29, 60). Selden (1993: 307). Fig. 23. Order CHELONETHI Family CHEIRIDITIDAE Electrobisium acutum Cockerell, 1917. Holotype, In.19118 (In.19117—22), also In.19123(3). Cockerell (1917d: 360, fig. 1). Schawaller (1978: 3, 17). Zherikhin (1978: 115). Morris (1980: 36). Keilbach (1982: 188). Harvey (1990: 334). Poinar (1992: 220). Spahr (1993a: 18,60). Transferred from the Neobisiidae by Judson (1997: 7, 53). Judson, this volume. Family OLPIIDAE Amblyolpium burmiticum (Cockerell, 1920). Holotype, In.20149. LIST OF INCLUSIONS IN BURMESE AMBER Listed by Spahr (1993: 17, 60). Cockerell (1920a: 274, fig. 1). Petrunkevitch (1955: 82). Schawaller (1978: 3). Zherikhin (1978: 115). Morris (1980: 40). Keilbach (1982: 188). Harvey (1990: 239). Poinar (1992: 220). Spahr (1993a: 17, 60). Transferred from Garypus (Garypidae) by Judson (1997: 12, 53). Fig. 24. Order SCORPIONIDA Unidentified. In.20174. Ross (1998: 36, fig. 100). DIPLOPODA Order POLY XENIDA Family SYNXENIDAE ‘Polyxenus’ burmiticus Cockerell, 1917. Holotype, In.19122 (In.19117-—22). Cockerell (1917b: 40-41, fig. 1). Condé (1954: 75). Zherikhin (1978: 114). Morris (1980: 45). Keilbach (1982: 201). Spahr (1993a: 53, 60). Sergei Golovach (pers. comm.) considers that this species does not belong to Polyxenus or the Polyxenidae. Fig. 18. GASTROPODA Unidentified. In.20194(1). Ross (1998: 22, fig. 59). VERTEBRATA SQUAMATA Unidentified. In.19102—3(1). Ross (1998: 25, fig. 68). Plantae MUSCI Family HYPNODENDRACEAE Hypnodendron sp. V.15536 Dixon (1922: 150, figs a—c; 1927: 96). Jovet-Ast (1967: 107, fig. 50). Spahr (1993b: 27). ACKNOWLEDGEMENTS. Many thanks go to Vladimir Zherikhin, Alexandr Rasnitsyn and colleagues at the Paleontological Institute, Moscow for com- ments, and to Michael Engel (American Museum of Natural History, New York) for checking the Apidae records. REFERENCES Armbruster, L. 1941. Uber Insektenstaaten der Vorwelt. 1. Miociine Randecker Termiten. Archiv fiir Bienenkunde, 22: 3-43. Bekker-Migdisova, E.E. 1962. Otryad Heteroptera. Raznokrylje. In, Rohdendorf, B.B. (ed.) Osnovy Paleontologii: Chlenistonogie. Trakheinye i Khelitserovye. Akademii Nauk, Moscow. Pp. 208-226. 1991. Order Heteroptera. In: Rohdendorf, B.B. (ed.) Fundamentals of Paleontology, Vol. 9, Arthropoda, Tracheata, Chelicerata. Amerind Publishing Co., New Delhi. Pp. 289-317. — & Vishniakova, V.N. 1962. Otryad Psocoptera. Senoedy. In, Rohdendorf, B.B. (ed.) 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Pp. 40-97. & Sukacheva, I.D. 1973. O melovykh naseomonosnykh ‘yantaryakh’ (retiniakh) severa Sibiri. Voprosy Paleontologii Nasekomykh. Nauka, Leningrad. Pp. 3-48. LIST OF INCLUSIONS IN BURMESE AMBER Allocrotelsa burmiticus (Cockerell, 1917d) comb. nov. (Zygentoma: Lepismatidae), Burmese amber, holotype, In.19119. Length of body 2.7mm. ?Labidura electrina Cockerell, 1920b, (Dermaptera: Labiduridae), Burmese amber, holotype, In.20146. Length of head (excluding palps) 0.8mm. Kalotermes swinhoei (Cockerell, 1916), (Isoptera: Kalotermitidae), Burmese amber, holotype, In.19096. Length (including wings) 6.0mm. Burmitembia venosa Cockerell, 1919a, (Embioptera: “Burmitembiidae’ ), Burmese amber, holotype, In.19132. Length 5.0mm. ?Psylloneura perantiqua Cockerell, 1919b, (Psocoptera: Pachytroctidae), Burmese amber, holotype, In.19136. Length of forewing 2.4mm. Sabatinca perveta (Cockerell, 1919a), (Lepidoptera: Micropterygidae), Burmese amber, holotype, In.19135. Length of body 2.5mm. 17 A.J. ROSS AND P.V. YORK Se 10 | 2s oe 11 12 Fig.7 ‘Liburnia’ burmitina Cockerell, 1917e, (Hemiptera: Achilidae), Burmese amber, holotype, In.19105. Length 4.6mm. Fig.8 Disphaerocephalus macropterus Cockerell, 1917d, (Hemiptera: Enicocephalidae), Burmese amber, holotype, In.19123(1). Length of wing 2.6mm. Fig.9 Cryphalites rugosissmus Cockerell, 1917d, (Coleoptera: ?Colydiidae), Burmese amber, holotype, In.19111. Length 2.3mm. Fig. 10 ‘Dermestes’ larvalis Cockerell, 1917e, (Coleoptera: Dermestidae), Burmese amber, holotype, In.19108. Length (excluding hairs) 0.7mm. Fig.11 ‘Elater’ burmitinus Cockerell, 1917d, (Coleoptera: Elateridae), Burmese amber, holotype, In.19102. Length of elytra 7.2mm. Fig. 12 Myodites burmiticus Cockerell, 1917a, (Coleoptera: Rhipiphoridae), Burmese amber, holotype, In.19092. Length 2.9mm. LIST OF INCLUSIONS IN BURMESE AMBER 15 17 Fig. 13 Fig. 14 Fig. 15 Fig. 16 Fig. 17 Fig. 18 ceaiiginciasi 18 Johannsenomyia swinhoei Cockerell, 1919b, (Diptera: Ceratopogonidae), Burmese amber, holotype, In.19133. Length 1.3mm. Electrocyrtona burmanicum Cockerell, 1917a, (Diptera: Empididae), Burmese amber, holotype, In.19099. Length 1.2mm. Trichomyia swinhoei Cockerell, 1917a, (Diptera: Psychodidae), Burmese amber, holotype, In.19101. Length of body 1.6mm. ‘Sciara’ burmitina Cockerell, 1917a, (Diptera: Mycetophilidae), Burmese amber, holotype, In.19100. Length (excluding antennae) 3.4mm. Burminoptila bemeneha Botosaneanu, 1981, (Trichoptera: Hydroptilidae), Burmese amber, holotype, In.20180. Length 3.0mm. Polyxenus burmiticus Cockerell, 1917b, (Diplopoda: Polyxenidae), Burmese amber, holotype, In.19122. Length (excluding hairs) 2.1mm. 20 23 Fig. Fig. Fig. Fig. Fig. Fig. A.J. ROSS AND P.V. YORK 24 Bethylitella cylindrella Cockerell, 1917d, (Hymenoptera: Bethylidae), Burmese amber, holotype, In.19120. Length of thorax 0.9mm. Epyris atavellus Cockerell, 1920a, (Hymenoptera: Bethylidae), Burmese amber, holotype, In.20149(1). Length 2.5mm. Haidomyrmex cerberus Dlussky, 1996, (Hymenoptera: Formicidae), Burmese amber, holotype, In.20182. Length of antenna 1.5mm. Protofoenus swinhoei Cockerell, 1917a, (Hymenoptera: Aulacidae), Burmese amber, holotype, In.19091. Length of wing 3.9mm. Cheyletus burmiticus Cockerell, 1917b, (Acarina: Cheyletidae), Burmese amber, holotype, In.19115. Length 0.7mm. Amblyolpium burmiticum Cockerell, 1920a, (Chelonethi: Garypidae), Burmese amber, holotype, In.20149. Length of chela 0.8mm Bull. nat. Hist. Mus. Lond. (Geol.) 56(1):21—24 Issued 29 June 2000 A preliminary list of arthropod families present in the Burmese amber collection at The Natural History Museum, London A. P. RASNITSYN Arthropod Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya Str. 123, Moscow 117647, Russia A. J. ROSS Department of Palaeontology, The Natural History Museum, Cromwell Road, South Kensington, London SW7 SBD SYNOPSIS. The present account is based on the 117 pieces of Burmese amber in the collections of The Natural History Museum, London. The material was found to be very rich in fossils yielding almost 1200 individual arthropod specimens. INTRODUCTION Below is a list of families of arthropods identified in the pieces of Burmese amber in the Department of Palaeontology of The Natural History Museum, London. The collection was initially identified to Order level by AJR, then APR made a more thorough study with additional identifications by scientists at the Paleontological Insti- tute, Moscow and other institutions. This list provides, for the first time, a detailed account of the Burmese amber collection at the NHM and hopefully will stimulate further research by specialist entomolo- gists and palaeoentomologists. The list follows the format of Ross & Jarzembowski (1993). For each order and family, the identifier(s) is given in brackets, along with the total number of specimens. For each family, the registration numbers of the pieces of amber (pre-fixed by In.) are given. For each piece of amber, the number of individual specimens is given in brackets if there is more than one (abbreviated spm), and if the piece contains a type or figured specimen this is also indicated in brackets (abbreviated T or Frespectively). Specimens In. 19102—23, In.20149- 50 and In.20195 all came from the same block that was sliced up into 7 slabs (see Cockerell, 1917). The 7 slabs contain a total of 458 inclusions, which makes the whole block the richest piece of amber ever recorded. For further information on the type and figured specimens in the collection, see the comprehensive list by Ross & York, this volume. The specimens of copal that Cockerell (1922) recorded are not included because there is uncertainty as to whether they are from Burma. The total number of specimens in the list is 1198. Collembola Order Collembola (APR) 57+1? Family Sminthuridae 32+3? In.19098?, In.19100, In.19123 (4 spm), In.20150? (2 spm), In.20151, In.20186 (10 spm), In.20188 (16 spm) Undetermined 22+1? In.19100, In.19102—3, In.19123 (2 spm), In.19128, In.19132, In.20150 (4+1? spm), In.20151, In.20171 (9 spm), In.20180, In.20183 © The Natural History Museum, 2000 Insecta Order Archaeognatha (APR) 1 Family Machilidae 1 In.20188 Order Zygentoma (APR) 4 Family Lepismatidae 3+1? In. 19104—6, In.19107—16, In.19117—22 (T), In.20149? Order Ephemeroptera (N. D. Sinitshenkova) 1 Family Prosopistomatidae 1 In.20173 (T) Order Blattodea (APR) 58 Family Blattellidae (AJR) 3 In.19102-3, In.20149, In.20172 Family Polyphagidae (AJR) 1 In.20194 Undetermined (mainly nymphs, isolated legs or body parts) 54 In.19102—3 (9 spm), In.19104—6 (2 spm), In.19107—16 (5 spm), In.19117—22, In.19123 (6 spm), In.19128, In.19132 (2 spm), In.19136 (2 spm), In.20149 (9 spm), In.20150 (7 spm), In.20151, In.20152, In.20164, In.20171 (2 spm), In.20174, In.20175, In.20182, In.20192, In.20200 Order Dermaptera 1 Family Labiduridae l In.20146 (T) Order Embioptera 1 Family Burmitembiidae ] In.19132 (T) Order Isoptera (APR) 91? Family Hodotermitidae (K. Krishna) 2 In.19123, In.20160 Family Kalotermitidae 89? In.19096 (2 spm, 1 T), In.19102—3 (11 spm, 1 T), In.19104—6 (3 spm), In.19107—16 (15 spm), In.19117- D5) 22, In.19123 (many, 50? spm), In.20149 (5 spm), In.20174, In.20699 Order Mantodea (P. Vrsansky, APR) 3+1? Undetermined 3+1? In.19100-1 (nymph), In.20150, In.20152 (nymph), In.20162? (ootheca?) Order Orthoptera (APR) 3+2? Superfamily Grylloidea Dit In.20152? (nymph), In.20171 (2 spm) Undetermined De In.19107—16?, In.20161? Order Phasmatodea (APR) 3? Undetermined 3? In.19117—22? (3 spm, eggs) Order Hemiptera (D. E. Shcherbakov) 82 Family Achilidae 28 In.19094 (2 spm), In.19102-3 (8 spm), In.19104—6 (2 spm, | T), In.19117—22 (5 spm), In.19123 (2 spm), In.20149 (4 spm), In.20150 (2 spm), In.20168, In.20186, In.20191 Family Aleyrodidae 3 In. 191334 (T), In.20193 (T), In.20703 Family Aphrophoridae 1b? In. 19094? Family Cixiidae IL In.19094 (2 spm, | T) Superfamily Coccoidea In.19098, In.19102—3, In.19125?, In.19132, In.20155, In.20160, In.20168 (2 spm), In.20172, In.20174, In.20175, In.20186, In.20188, In.20191, In.20193 (2 spm), In.20708 Family Coreidae s.1. In.20186 Family Enicocephalidae 6 In.19095 (T), In.19107—16 (2 spm, 2 T), In.19123 (2 spm, | T), In.19128 Superfamily Fulgoroidea undetermined IS In.19098 (2 spm), In.19130, In.20160, In.20167 (3 Spm) loeZOl7 1, IneZ01725 IineZOl7AS Ine2 Os: In.20191, In.20195 16+1? Family Ochteridae I In.20160 Undetermined 10 In. 19107—16 (5 spm), In.19117—22, In.19125, In.20152, In.20158, In.20185 Order Psocoptera 47 Family Compsocidae l In.20193 Family Pachytroctidae 2 In.19136 (2 spm, | T) Family Psyllipsocidae 4 In. 19097 (T), In.20174, In.20193, In.20706 Undetermined 40 In. 19096, In.19098 (2 spm), In.19102-3, In.19107—16, In.19177—22 (2 spm), In.19136 (18 spm), In.20150, In, 2010S @ spm); In201635 imnr2 02am? Oil. 3» In.20174, In.20175 (2 spm), In.20178 (2 spm), In.20186, In.20187, In.20199, In.20707 A.P. RASHITSYN AND A.J. ROSS Order Thysanoptera (APR) 14 Family Lophioneuridae (V. V. Zherikhin) 1 In.20194 (T) Undetermined 13 In.19133—4, In.19135, In.19098, In.20149, In.20168, In.20172 (8 spm) Order Coleoptera (APR, V. G. Gratshev, N. B. Nikitsky, P. Hammond) 195 Family Aderidae AV? In.20150? Family Anthicidae ] In.19107—16 (T) Superfamily Cantharoidea 1+1? In.19123 (larva), In.20169? Family Carabidae 1 In.20150 (larva) Family Cerambycidae 1 In.20188 Family Chrysomelidae 1 In.20150? (larva) Family Cisidae I? In.19136? Superfamily Cleroidea 1? In.20169? Family Colydiidae 3+3? In.19107—16? (T), In.19117—22?, In.19123 (2 spm), In.20149?, In.20174 Family Cucujidae 4? In.19102—3?, In.19104—6?, In.20150?, In.20174? Family Dermestidae 1+1? In.19107—16 (1+1? spm, 1 T) Family Elateridae 19 In.19102—3 (8 spm, | T), In.19107—16 (5 spm, 1 T), In.19117-—22 (2 spm), In.19123 (2 spm), In.20186, In.20187 Family Eucinetidae 1 In.20186 Family Eucnemidae 2 In.20168 (2 spm) Family Helodidae 12 In.20150? Family Lathridiidae In.19136 Family Melandryidae 1+1? In.19136, In.20706? Family Melyridae 1 In.20160 Family Microsporidae 1 In.19132 Family Mordellidae I In.20160 Family Nitidulidae 1 In.20178 (larva) Family Oedemeridae A In.19123?, In.20190? Family Pselaphidae 6 In.19104—6 (2 spm), In.19123, In.20150, In.20171, In.20207 Family Ptiliidae 3 In.19132, In.19160, In.19107—16 Family Rhipiphoridae i In.19092 (T) LIST OF ARTHROPOD FAMILIES IN BURMESE AMBER Family Salpingidae In.20199? Family Scraptiidae In.19102-3, In.19107—16, In.19117—22, In.19123 (11 spm), In.19126, In.20150 Family Staphylinidae In.19107—16, In.19136 (2 spm), In.20150 (13 spm), In.20152, In.20158, In.20160 (2 spm), In.20169, In.20751, In.20194 Superfamily Staphylinoidea undetermined In.19123, In.20194 Family Throscidae In.20150? Undetermined In.19092 (2 spm), In.19094 (2 spm), In.19098, In.19100-1, In.19102-3 (9 spm), In.19104-6, In.19107—16 (12 spm), In.19117—22 (12 spm), In.19123 (7 spm), In.19131, In.20149 (10 spm), In.20150 (3 spm), In.20151, In.20152 (5 spm), In.20158, In.20160, In.20164, In.20173, In.20176, In.20183 (2 spm), In.20188, In.20189 (3 spm), In.20192, In.20193, In.20194 (4 spm), In.20195, In.20196, In.20200, In.20205, In.20708 Order Diptera (V. A. Blagoderov, M. B. Mostovsky) Family Apsilocephalidae In.20167 (T) Family Cecidomyiidae In. 19096 (3 spm), In.19107—16(T), In.19125, In.20151, In.20160, In.20167, In.20168 (2 spm), In.20173, In.20174, In.20194 (1+1? spm), In.20210 Family Ceratopogonidae In. 191334 (T), In.20168 (5 spm), In.20172, In.20173 (2 spm), In.20190, In.20208, In.20700 Family Chaoboridae (E. D. Lukashevitch) In.20157 (T), In.20168 (2 spm, 2T) Family Chironomidae In.19098 (2 spm), In.19117—22 (2 spm), In.20148?, imie2 0153 (7 spm), In.20168 @ spm), In.20173?, In.20183 (1+2? spm), In.20189, In.20194 (1+1? spm), In.20207, In.20704?, In.20705 (2 spm), In.20707 (4 spm), In.20708 (2 spm) Family Diadocidiidae In.20175 (4 spm), In.20188 Family Dolichopodidae In.20197? Family Empididae In. 19096 (8 spm), In.19099 (3 spm, | T), In.19102—3 (4 spm), In.19104—6 (P), In.19107—16 (3? spm), In.19117— 22 (5 spm, | T), In.19123 (7 spm), In.19136 (2 spm), In.20149?, In.20151, In.20173 (2 spm), In.20178, In.20192 (3 spm), In.20209 Family Keroplatidae In.19104—6 (T), In.20179 Family Limoniidae In.20155 Family Mycetophilidae (P. Chandler) In.19100-1 (T) Superfamily Mycetophiloidea undetermined In.19136, In.20171 (4 spm), In.20175 (2 spm) Family Phoridae In. 19102-3, In.19123, In.20193 Family Psychodidae 22 1? 89 12 25+6? 38+4? 33+2? 23 In.19097?, In.19098, In.19100-1 (2 spm, 1 T), In.19133—-4?, In.20147 (T), In.20155 (2 spm), In.20156, In.20164, In.20167 (5 spm), In.20168 (2 spm), In.20171, In.20172 (2 spm), In.20187 (2 spm), In.20189, In.20198, In.20707 (4 spm), In.20710 (7 spm) Family Rhagionidae 1+1? In.20175, In.20709? Family Scatopsidae | In.20150 Family Sciaridae 5) In. 19096, In.20151, In.20152, In.20188, In.20199 Family Therevidae 1 In.20148 (T) Superfamily Tipuloidea undetermined In.20171 Undetermined 34 In.19097, In.19099, In.19102—3 (2 spm), In.19107— 16 (2 spm), In.19117—22 (3 spm), In.19123 (larva), In.19136, In.20149 (3 spm), In.20150, In.20168, In.20172, In.20173 (2 spm), In.20183, In.20184 (2 spm), In.20188, In.20192 (2 spm), In.20194 (2 spm), In.20199 (2 spm), In.20208 (3 spm), in.20708 (2 spm) Order Hymenoptera (APR) 102 Family Bethylidae 31 In.19107—16 (5 spm, 2 T), In.19117—22 (3 spm, | T), In.19123 (9 spm), In.19136 (2 spm), In.20149 (7 spm, 1 T), In.20150 (5 spm) Superfamily Chalcidoidea 3 In.20208, In. 191334, In.20194 Family Diapriidae Py In.19124? Family Embolemidae 2 In.20151, In.20194 Family Evaniidae 1 In.20192 (T) Family Formicidae 5 In.19125 (T), In.19132, In.20182 (3 spm, 1 T, 1F) Family Gasteruptiidae 6 In.19091 (T), In.19098 (T), In.19117—22 (T), In.19129, In.20169, In.20174 Family Megaspilidae 1 In.20159 Family Mymarommatidae 2 In.20193, In.20208 Family Pelecinidae | In.20174 Family Scelionidae we; In.19092, In.19094, In.19096, In.19102-3 (2 spm), In.19107—16, In.19117—22 (5 spm), In.19136 (3 spm), In.20158 (2 spm), In.20167, In.20183 (2 spm), In.20193, In.20197, In.20706 (2 spm) Family Serphitidae 2 In.20151, In.20190 (F) Family Sphecidae 17+1? In.19093 (T), In.19123 (2 spm, 2 T), In.19125 (T), rm 1OI36r(Orspm, 9) 1)s in 20150 1), In 201647 In.20181 (T), In.20197 (T), In.20711 (T) Family Tiphiidae yi In.20150? (2 spm) Undetermined 4 In.19102-3, In.19104—6, In.20149, In.20208 Order Lepidoptera (APR, G. Robinson) Family Micropterygidae In.19135 (T), In.20167, In.20168, In.20204. Undetermined In.19123, In.20151, In.20172 Order Neuroptera (A. G. Ponomarenko, M. Engel) Family Berothidae In.20193, In.20177, In.20197 Family Osmylidae In.20150 (larva) Superfamily Osmyloidea undetermined In.19107—16? (larva) Undetermined In.19102-3 Order Raphidioptera (A. G. Ponomarenko) Undetermined In.20150 (larva) Order Trichoptera Family Hydroptilidae In.20180 (T) Order Undetermined (APR) In.19102-—3 (3 spm), In. 19104—6, In.19107—16 (2 spm), In.19117—22, In.20149 (2 spm), In.20150 (2 spm), In.20167 (2 spm) Arachnida Order Acarina (A. L. Kartsev, O. L. Makarova) Family Anystidae In.20167 Family Bdellidae In.19128, In.20193 Superfamily Bdelloidea undetermined In.20203? Family Cheyletidae In.19107—16 (T), In.19123 (3 spm), In.20149 Family Erythraeidae In.19102-3, In.19107—16 (2 spm), In.19123 (3 spm), In.19132, In.20149, In.20150 (11 spm), In.20188, In.20193 (5 spm) Family Eupodidae In.20150?, In.20193 (5+1? spm) Family Ixodidae In. 19125 Guvenile) Undetermined In. 19095, In. 19100-1, In. 19102—3 (14 spm), In.19104— 6 (6 spm), In.19107-116 (25 spm), In.19117—22 (8 spm), In.19123 (25 spm), In.19128 (2 spm), In.19132, In.20149 (4 spm), In.20150 (8 spm), In.20151, In.20152 3+1? 13 164 i) A.P. RASHITSYN AND A.J. ROSS (8 spm); In20158, In-20171 GC spm)yin20he: In.20183, In.20186 (3 spm), In.20188, In.20189 (7 spm), In.20699 Order Araneae (K. Y. Eskov) 36 Family Eusparassidae 1 In.20197 (juvenile) Family Myrmectiidae 3 In.20193, In.20197, In.20208 Family Oonopidae 2 In.20210, In.20168 Family Pisauridae (D. Penney) 1 In.19132 Family Tetragnathidae 1? In.20208? Family Theridiidae 1 In.20193 Family Thomisidae 1 In.20174 (juvenile) Undetermined 26 In.19104—6, In.19117—22 (2 spm), In.19136 (3 spm), In.20151, In.20152 (4 spm), In.20154, In.20172 (2 spm), In.20174, In.20178, In.20183, In.20186 (3 spm), In.20189 (2 spm), In.20191 (2 spm), In.20194, In.20700 Order Chelonethi (APR, K. Y. Eskov) 38 Family Cheiridiidae (M. L. I. Judson) 3 In.19117—22 (2 spm, 1| T), In.19123 (F) Family Olpiidae (M. L. I. Judson) | In.20149 (T) Undetermined 34 In.19102—3 (6 spm), In.19107—16 (5 spm), In.19117— 22, In.19123 (11 spm), In.20149 (2 spm), In.20150 (6 spm), In.20171, In.20189, In.20195 Order Scorpionida (APR) Undetermined l In.20174 (F) Diplopoda Order Polyxenida (APR, S. I. Golovach) 60 Family Synxenidae 60 In.19102—3 (4 spm), In.19104—6 (6 spm), In.19117—22 (2 spm, 1 T), In.19123 (24 spm), In.20149 (5 spm), In.20150 (18 spm), In.20169 Order Undetermined (S. I. Golovach) 1? In.19123? REFERENCES Cockerell, T.D.A. 1917. Arthropods in Burmese amber. American Journal of Science, Ser. 4, 44: 360-368. 1922. Fossils in Burmese amber. Nature, 109: 713-714. Ross, A. J. & Jarzembowski, E.A. 1993. Arthropoda (Hexapoda; Insecta). Jn: Benton, M.J. (editor) The fossil record 2: 363-426. Chapman & Hall, London. Bull. nat. Hist. Mus. Lond. (Geol.) 56(1): 25—28 Issued 29 June 2000 The first fossil prosopistomatid mayfly from Burmese amber (Ephemeroptera; Prosopistomatidae) N. D. SINITSHENKOVA Arthropod Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya Str. 123, Moscow 117647, Russia SYNOPSIS. An adult female of Myanmarella rossi, anew genus and species of mayfly is described from Burmese amber. M. rossi belongs to the family Prosopistomatidae and is the first fossil record of this family. INTRODUCTION The Prosopistomatidae are a small, highly specialised and poorly known family of mayflies differing greatly from any other family of the order. Its phylogenetic affinities are uncertain. All living Prosopistomatidae belong to the single genus Prosopistoma Latreille with 15 species distributed widely but discontinuously throughout eastern hemisphere, mostly in the Oriental region (Hubbard, 1979; Pearson & Penridge, 1979; Soldan & Braasch, 1984). The alate stages are known for three species only, P. foliaceum (Fourcroy), P. africanum Gillies (Gillies, 1954, 1956; Fontaine, 1955), and P. pearsonorum Campbell & Hubbard (Campbell & Hubbard, 1998); in addition the wings of an undescribed Malaysian species have been figured (Peters & Campbell,1991). Males are represented by both imagos and subimagos while only subimagos are known for females which supposedly lack the imago stage (Gillies, 1954). Below a fossil from Burmese amber is described and referred to a new genus and species, Myanmarella rossi, which constitutes the first fossil member of the family Prosopistomatidae. Burmese amber originates from the Hukawng Valley in northern Myanmar (formerly Burma). It is probably of Upper Cretaceous age (Zherikhin & Ross, this volume). M. rossi is the only mayfly repres- ented among 840 insect inclusions, which belong to 20 orders, in the Burmese amber collection kept in the Department of Palaeontology at The Natural History Museum, London. SYSTEMATIC DESCRIPTIONS Order EPHEMEROPTERA Latreille, 1810 Superfamily CAENOIDEA Newman, 1853 Family PROSOPISTOMATIDAE Lameere, 1917 Genus MYANMARELLA nov. NAME. After the country of origin, Myanmar. TYPE SPECIES. Myanmarella rossi Sinitshenkova, sp. nov. Prob- ably Upper Cretaceous, Hukawng Valley, Myanmar (Burma). DESCRIPTION. Imago, female. Very small mayfly. Antennae mod- erately widely separated, with strongly inflated base (pedicel tumorous). Forewing 3.5 times as long as hind wing and 2.3 times as long as wide, broadly rounded at apex; RS branched, its hind branch © The Natural History Museum, 2000 strongly approximated to MA,; MP, and IMP not reaching wing base, cubital area with single short intercalary vein, costal area as well as RS and M systems with very weak interrupted longitudinal veins crossing the main ones, anal veins well developed, no less than 2 in number, crossveins few, very weak; hind margin fringed with hairs. Hind wing elongate, with reduced venation, RS branched, crossvenation absent. Legs fully developed, tarsi 5-joined. Cerci longer than body, clearly segmented, probably lacking fringe of long hairs; paracercus absent. Monobasic. REMARKS. Myanmarella differs from living Prosopistoma in hav- ing the imaginal stage in female sex, the antennae longer, the wing venation less reduced, the legs fully developed, with multisegmented tarsi, the cerci long and segmented, the paracercus absent. Myanmarella rossi sp. nov. Figs la—g, 2 HOLOTYPE. NHM Palaeontology Department In.20173, a com- plete mayfly with outstretched wings, situated near the surface of a large piece of amber, in which several other insects are embedded: 3 Diptera (including an isolated wing of Mycetophiloidea situated over the apex of the left mayfly wing), 1 Psocoptera, and | Coleoptera. NAME. After Mr. Andrew J.Ross, Curator of the fossil arthropods in The Natural History Museum, London. MATERIAL AND LOCALITY. Holotype only, NHM Palaeontology Department In.20173, in Burmese amber; Hukawng Valley, Myanmar (Burma); probably Upper Cretaceous (see Zherikhin & Ross, this volume). DESCRIPTION. Adult, female. General colour of body and wings in amber pale yellow. Eyes large, widely separated, ocelli large, round, lateral ones more visible than the front one. Antennae almost 1.5 times longer than width of head. Fore femora slightly longer than tibiae, tibia almost as long as tarsi; midfemora slightly shorter than tibiae, tarsi 1.7 times shorter than tibiae. First tarsal joint is the longest both on fore and middle legs; last joint is the shortest, second and third subequal in length, shorter than fourth. Wings transparent, without markings. Venation of right and left wing slightly different. On both forewings RS forks three times, fork of anterior branch narrow, with subparallel long branches; posterior branch of RS and MA, paired; IMP and MP, not reaching wing base; both CuA and CuP simple, CuA terminating in outer margin before tornus, interca- lary vein in cubital area not reaching CuP, ending at outer margin at the level of tornus; two simple anal veins. On the right forewing (with costal angulation rounded) the base of 26 N.D. SINITSHENKOVA Fig. 1 Myanmarella rossi gen. nov. sp. nov., holotype, In.20173, Burmese amber. a, head, scale 0.01 mm; b, fore leg; c, middle leg; d, right fore wing; e, left fore wing; f, right hind wing; g, left hind wing; scales for 1b—1g 0.1 mm. FOSSIL MAYFLY FROM BURMESE AMBER Fig. 2 Myanmarella rossi gen. nov. sp. nov., holotype, In.20173, Burmese amber. Length of right fore wing 2.25mm. C strongly arched; R and hind branch of anterior RS fork ending in thin fold not reaching outer margin, between hind branches of RS two very weak interrupted longitudinal veins are visible; between MA and IMA two fine folds are visible, the hind one forked, between IMA and MA, one weak longitudinal vein; crossvenation restricted to basal and apical parts of wing. On the left forewing, fore margin is crumpled at the base, in radial area a fine vein forks two times, one near wing base and another slightly proximal from wing midlength, a fine vein forks near the outer wing margin crosses IRS, between IRS and hind branch of RS no veins are visible; IMA weak, near MA a fine interrupted vein; behind CuP a fine curved vein well visible; crossvenation only at basal half of wing (about 10 veinlets). Hind wings pointed at apex, 2.8 (right) to 2.7 (left) times as long as wide, costal angulation widely rounded, longitudinal veins very fine. Venation of hind wings is also different: on the right wing one fork of RS with short intercalary well visible, base of SC not visible, subcostal area wide, R in its apical third attached by one fine crossvein, RS distally forking, MA and MP simple. On the left hind wing apex of SC is not visible, bases of R and M fused, there are several additional fine veins near main longitudi- nal one. Abdomen about twice as long as thorax. Cerci with long segments, the segment is three times as long as wide. MEASUREMENTS. The length of fore wings: right — 2.25 mm, left — 2.15 mm, their width respectively — 0.95 mm and 0.92 mm; the length of hind wings: right — 0.65 mm, left — 0.62 mm. DISCUSSION. The nymphs of living Prosopistomatidae inhabit ex- clusively running waters where they occur mostly on the underside of stones in rapid flows (Gillies, 1954; Peters, 1967; Alouf, 1977; Soldan & Braasch, 1984; Koch, 1988); as a rule they occur in rather small streams though the European Prosopistoma foliaceum is found mainly in large rivers such as the Rhone, Rhein, and Vitava (Lafon, 1952). The winged stages are extremely short-lived, with the life- span possibly less than an hour (Gillies, 1954). Both the nymphs and adults have a very poor perservation potential as fossils. The winged insects are very delicate, and after death their bodies rapidly become dried up and shrivelled so that even freshly collected specimens can not be studied accurately without special care (Gillies, 1954). Thus it is not surprising that no fossil prosopistomatids were known up until now. Myanmarella is placed in this family because of its peculiar wing 4) venation with weak additional longitudinal veins and few crossveins and especially because of its long antennae with the strongly en- larged pedicel. These features are unique synapomorphies of Prosopistomatidae not occurring in any other mayfly family. In living species, the peculiar shape of the antennae occurs in male imagos only (females have this feature less expressed); most prob- ably, this is not sexual dimorphism but a character lost in modern Prosopistoma females because of lacking an imago stage. Myanmarella is clearly less advanced than recent Prosopistoma, as indicated by its less reduced wing venation and especially by the fully developed legs with 5-segmented tarsi. In Prosopistoma, the male imagos have short but functional legs with 3-segmented tarsi while in the females the legs are extremely reduced and non- functional. The long and clearly segmented cerci of Myanmarella are also plesiomorphous in comparison to shorter (even in the male imago) ones of Prosopistoma. On the other hand, Myanmarella is autapomorphous in a complete reduction of the paracercus which is as long as the cerci in Prosopistoma. Thus, Myanmarella represents rather an extinct sister lineage to Prosopistoma than the ancestral stock of the family. The family Prosopistomatidae was placed by different authors either near Baitiscidae (Gillies, 1954), Oligoneuriidae and Tricorythidae (Degrange, 1955; Demoulin, 1955). In the recent phylogenetic system of the order (McCafferty, 1991) itis included in the superfamily Caenoidea and separated from Baetiscoidea. Myanmarella is already a highly advanced genus which does not clarify the disputable phylogenetic relations of the family. It is possible only to say with certainty, that the wing venation of Myanmarella indicates no similarity with that of Baetiscoidea ex- cept for the reduction in number of the anal veins to two. This state occurs in different mayfly lineages and should be of little phylogenetic importance. Biologically Myanmarella was probably similar to Prosopistoma except for having more long-living winged females that reached the imago stage. The functional legs should have allowed them to rest on vegetation; Prosopistoma females cannot rest on a substrate and spend all of their short life in flight. ACKNOWLEDGEMENTS. [| extend my appreciation to Mr. Andrew J. Ross, curator of the fossil arthropods in the Natural History Museum in London for passing me such an exciting specimen of mayfly. I especially thank Prof. William L.Peters, Florida A & M University, Tallahassee, Dr. John. E. Brittain, University of Oslo, for sending me the copies of necessary articles, and Dr. Vladimir V. Zherikhin, Paleontological Institute of the Russian Academy of Sciences, Moscow, Russia, for the criticism of the manuscript. I also thank Mr Peter York (NHM) for photographing the specimen. REFERENCES Alouf, N. J. 1977. Sur la présence du genre Prosopistoma au Liban. Description de P. oronti n. sp. et de P. phoenicium n. sp. (Ephemeroptera). Annals of Limnology, 13 (2): 133-139. Campbell, I.C. & Hubbard, M.D. 1998. A new species of Prosopistoma (Ephemeroptera: Prosopistomatidae) from Australia. Aquatic Insects, 20 (3): 141— 148. Degrange, C. 1955. Sur la morphologie de Prosopistoma foliaceum Fourcroy. Compte Rendus de l’Academie de Science, Paris, 240: 1668-69. Demoulin, G. 1955. A propos des affinités systématiques des Prosopistomatidae (Ephemeroptera). Bulletin et Annales de la Société Royale d’Entomologie de Belgique, 7-8: 209-213. Fontaine, J. 1955. Les formes ailées de Prosopistoma foliaceum Fourcroy (Ephemeroptera). Bulletin de la Société Linnéenne de Lyon, 24: 60-65. Gillies, M. T. 1954. The adult stages of Prosopistoma Latreille (Ephemeroptera), with 28 descriptions of two new species from Africa. Transactions of the Royal Entomologi- cal Society of London, 105: 355-372. 1956. A supplementary note on Prosopistoma Latreille (Ephemeroptera). Pro- ceedings of the Royal Entomological Society of London, 31: 165-166. Hubbard, M. D. 1979. A nomenclatural problem in Ephemeroptera: Prosopistoma or Binoculus? Proceedings of the 2nd International Conference on Ephemeroptera, 1975, Krakow: 73-77. Koch, S. 1988. Mayflies of the Northern Levant (Insecta: Ephemeroptera). Zoology in the Middle East 2, Insecta: 89-112. Lafon, J. 1952. Note sur Prosopistoma foliaceum Fourc. (Ephéméropteres). Bulletin de la Société Zoologique de France, 77: 425-436. McCafferty, W. P. 1991. A phylogenetic classification of the Ephemeroptera (Insecta); a commentary on systematics. Annals of the Entomological Society of America, 84 (4): 343-360. Pearson, R. G. & Penridge, L. K. 1979. First record of Prosopistoma sedlaceki in N.D. SINITSHENKOVA Australia (Ephemeroptera: Prosopistomatidae). Journal of the Australian Entomo- logical Society, 18: 362. Peters, W. L. 1967. New species of Prosopistoma from the Oriental Region (Prosopistomatoidea: Ephemeroptera). Tijdschrift voor Entomologie, 110: 207— 222. & Campbell I. C. 1991. Ephemeroptera (Mayflies). In: The insects of Australia, 2nd edition, 1: 279-293. Melbourne University Press. Poinar, G. 1992. Life in amber. Stanford University Press, Stanford. 350 pp. Rasnitsyn, A. P. 1996. Burmese amber in the Natural History Museum. Inclusion (Wrostek), 23: 19-21. Soldan, T. & Braasch, D. 1984. Two new species of the genus Prosopistoma (Ephemeroptera, Prosopistomatidae) from Vietnam. Acta Entomologica Bohemoslovaca, 81: 370-376. Therikhin, V. V. 1978. Rasvitye 1 smena melovykh 1 kaynozoyskikh faunistitcheskikh kompleksov. Trudy Paleontologitcheskogo Instituta, 165: 198 pp. Bull. nat. Hist. Mus. Lond. (Geol.) 56(1):29-37 Issued 29 June 2000 The most primitive whiteflies (Hemiptera; Aleyrodidae; Bernaeinae subfam. nov.) from the Mesozoic of Asia and Burmese amber, with an overview of Burmese amber hemipterans D.E. SHCHERBAKOV Arthropod Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya Str. 123, Moscow 117647, Russia SYNOPSIS. Relationships, adult morphology, and taxonomic structure of whiteflies are discussed, and their vein nomenclature is corrected. The subfamily Udamoselinae in the broad sense (including Aleurodicinae) is restored; a new subfamily Bernaeinae (family Aleyrodidae) is established comprising most Mesozoic whiteflies. The oldest known whiteflies are described, Juleyrodes gilli gen. et sp. nov. and J. visnyai sp. nov. from the Late Jurassic (and possibly also Early Cretaceous) of Asia. Their nearest relative, Burmoselis evelynae gen. et sp. nov., is from Burmese amber (probably Upper Cretaceous). These genera retain the venation more complete than previously known for whiteflies, confirming that the group descended from Psyllomorpha. Other fossil aleyrodids are listed, as are also the taxa excluded from the group. Burmese amber Hemiptera are reviewed. INTRODUCTION Palaeontological evidence supports the classification of Homoptera proposed by Borner (1904) and developed by Hennig (1969) and Schlee (1969), who group whiteflies with psyllids separating them from aphids plus coccids. The fossil record indicates that these two lineages were separate from the very beginning, so, instead of polyphyletic ‘Sternorrhyncha’, two subordinal names within Hemi- ptera s.l. are used: Aphidinea (including Aphidomorpha and Coccomorpha) and Psyllinea (including Psyllomorpha and Aleyrodomorpha) (Shcherbakov, 1990). In contrast, molecular phylogenies of Hemiptera based on 18S rDNA show the whitefly lineage in a variable position (but never as a sister group to psyllids): as a sister group to aphids plus coccids (after separation of psyllid lineage; Campbell et al., 1994; Dohlen & Moran, 1995a: figs. 6-7); as a sister group to all the other Sternorrhyncha (Dohlen & Moran, 1995: fig. 8); as a member of an unresolved tritomy with psyllids and aphids+coccids (Campbell et al., 1995a: fig. 3); or even as a sister group to psocids when the latter are included in the analysis (Dohlen & Moran, 1995: fig. 5). Such a discrepancy could arise from the fact that 18S rDNA in whiteflies is extremely long, with an unusually high substitution rate (Campbell etal., 1994; Dohlen & Moran, 1995), so at present the morphological and fossil evidence should not be discredited simply in favour of novel molecular techniques. Reduced wing venation of whiteflies has been variously inter- preted (Table 1). The two genera described below have an extra vein (free M) which is not known in other genera, their venation being clearly derivable from that of Protopsyllidiidae (Figs. 1, 3, 5). This fact resolves the vein homology and supports a whitefly origin from primitive Psyllomorpha, as concluded already by Quaintance & Baker (1913: 17): ‘Aleyrodidae .. . form an offshoot from the psyllid stem. This is indicated by the wing venation and by the structure of the mouthparts, legs, and genitalia.” Despite varying tracheal pat- terns in wingpads, the fossil record shows that the Sc in Hemiptera is always fused to R and R1, often except its base (along basal cell) and its apex (developed as a supernumerary free branch of R1 or R stem) © The Natural History Museum, 2000 (Shcherbakov, 1996). The subcostal groove is still traceable along the R+M stem in some protopsyllidiids and whiteflies (Figs. 1, 3, 5). In all extant suborders of Hemiptera the claval veins of the forewing, A1 and A2 (Pcu and Al in more accurate nomenclature) are primi- tively united into a Y-vein (ibid.); in primitive Psyllinea it ends near the apex of the clavus, therefore an oblique anal vein joining the posterior margin of the clavus far from its apex in Bernaeinae and Udamoselinae (see below) should correspond to Al, whereas A2 and A1+A2 are incorporated into this margin. Whitefly wings are usually considered to be uncoupled in flight (e.g. Carver, 1991). Indeed, the hindwing lacks the distal hooks which are present in Cicadinea (=Auchenorrhyncha), psyllids, aphids, and even in vestigial hamulohalteres of male coccids. Nevertheless, it retains a row of about 7—9 strong curved hairs along the proximal C portion (similar to that of psyllids), which aid in keeping the wings together in flight (Quaintance & Baker, 1913: 9). Whiteflies have much higher wingbeat frequencies (143—224 Hz) than other insects with non-fibrillar (synchronous) flight muscles, and the lowest wing loading ever recorded (up to 0.002 g/cm7), i.e. lower than in butter- flies (Wootton & Newman, 1979; Byrne et al., 1988). Such high frequencies are never reached by insects which have fore- and hindwings operating independently in flight. Partial reduction of interalar coupling in whiteflies could be associated with a unique combination of the high wingbeat frequency and low wing loading, both resulting from their miniaturization. Aleyrodidae, commonly known as whiteflies and initially consid- ered a single family, were divided into two subfamilies by Enderlein (1909): Udamoselinae, comprising both Udamoselis Enderlein and Aleurodicus Douglas, and Aleyrodinae. Quaintance & Baker (1913) considered these two genera to be distinct enough to create a third subfamily, Aleurodicinae (leaving Udamoselinae monobasic). Solo- mon (1935) and Sampson (1943) synonymized Aleurodicinae under Udamoselinae on account of the similar head shape and venation (see Table 2). Schlee (1970) doubted this relationship and now Udamoselis is treated as anomen dubium (as its pupal case and some important imaginal characters are unknown, and the unique holotype male is apparently lost), and Aleurodicinae as a valid subfamily 30 Table 1 D.E. SHCHERBAKOV Forewing vein nomenclature in whiteflies according to various authors. The vein symbols used are the usual ones (those from Enderlein and Szelegiewicz are slightly altered for uniformity); cl.f, claval furrow (associated with CuP in Hemiptera); Ax, axillaris (now out of use). Enderlein 1909 C+R1 Rs Quaintance & Baker 1913 C+Sc R1 Borner 1910; Haupt 1934 C R Gomez-Menor 1944 R Schlee 1970 € R Szelegiewicz 1971 C+Sc Rl This paper C (Sc+)R1* * Sc indistinguishably fused to R-R1, not to C (see text). ** Free M developed only in some Bernaeinae. *** Tn fact Pcu (postcubitus = anteriormost anal s.1.; see text). name (Mound & Halsey, 1978). Some authors have elevated whitefly subfamilies to families and Aleyrodidae s./. to superfamily rank, e.g. Schlee (1970). As noted by Solomon (1935) and Sampson (1943), the aleurodicine genera Ceraleurodicus Hempel (= Radialeurodicus Bondar, = Parudamoselis Visnya; synonymy after Mound & Halsey, 1978) and Synaleurodicus Solomon, and especially C. kesselyaki(Visnya, 1941), are strikingly similar to Udamoselis in that the paronychium is very thin and hardly detectable, the 3—7th abdominal segments in males possess upper lateral furrows, C is thickened in the male forewing, and in overall venation pattern (except that A is reduced). The anal vein is developed as in Udamoselis in some other Ceraleurodicus species: C. splendidus Hempel and C. octifer (Bondar) (Bondar, 1923: figs. 4, 6). In other aleurodicine genera the wing veins show gradual reduction up to only R-Rs left (Sampson, 1943), making it senseless to draw the subfamilial boundary between Udamoselis and its nearest relatives. Therefore, despite an incomplete knowledge of the type genus, the name Udamoselinae should be used in the broad sense of Enderlein (1909) and Sampson (1943), i.e. including Aleurodicinae. The abdominal wax plates were recorded in neither Udamoselis nor C. kesselyaki (perhaps they are reduced or over- looked due to abundant wax powdering in the latter). In the nymphal characters, Ceraleurodicus spp. (including C. kesselyaki) are typical Aleurodicinae, except for bearing up to 10 pairs of peripheral intersegmental ridges, at least several of them with tracheal ducts (instead of 3 tracheal ducts, paired thoracic and unpaired caudal, as usual; Sampson & Drews, 1957; Gill, 1990). Such ridges are found elsewhere only in Bondaria Sampson & Drews (imago unknown; Sampson, 1943), and imply a more com- plete complement of spiracles than the usual 4 pairs (2 thoracic and 2 abdominal); this supposedly primitive character merits reexamination. If the nymph of Udamoselis (when discovered) also bears the dorsal ridges, it would be possible to subdivide Udamoselinae into two tribes, a nominate one (including Ceraleurodicus and Bondaria) with ridges, and Aleurodicini with- out them. R1 RS M1 M2 CuA1 S Al + A2 CuP (CyA2 Fig. 1 Cicadellopsis sp. (Protopsyllidiidae), PIN 1255/410, forewing venation; Middle Jurassic; Yenisei River near Krasnoyarsk (Kubekovo locality). M - Cu A=cl.f Ax Rs = M Cu A M - Cu A Ax M — — Cu M - Cu cl.f A Rs - M CuA CuP Rs IME CuA CuP At** Table 2. Udamoselis compared to Alerodicinae and Aleyrodinae in the characters of imago diagnostic at subfamily level. Udamoselis Aleurodicinae Aleyrodinae paronychium (empodium) not visible — spine-like blade-like* abdomin. wax plates: male not describ. 3—5th segs. 3-6th(4th) abdomin. wax plates: female ? 3-6th segs. 34th segs.** forewing C thickened thick./not not CuA present present/lost vestigial/lost forewing CuP present present/lost present forewing A oblique oblique/no longitud?/no vertex conical conical/not not * In Siphoninus Silvestri one claw rather than paronychium is absent (Gomez-Menor, 1944). ** Segmental origin of two wax plates in female Aleyrodinae was variously interpreted: 4th & Sth segments (Bemis, 1904: fig. 42), 3+4th & 5+6th segments (Haupt, 1934; Weber, 1935), or 3rd & 4th (Gill, 1990: fig. 2.13); the latter version is adopted here as consistent with a sexually monomorphic condition in Neomaskellia Quaintance & Baker. The oldest whitefly imagines known so far, Bernaea neocomica (single female) and Heidea cretacica (single male), were described from Lower Cretaceous Lebanese amber by Schlee (1970). He mentioned several symplesiomorphies (head broad; eyes not subdi- vided; median ocellus retained; antennae 8-segmented; paronychium broad; abdominal wax plates absent; male parameres held vertically; ovipositor long, horizontal) and only one doubtful synapomorphy (rostrum reaching beyond hind coxae) of Cretaceous genera relative to Cenozoic whiteflies. Being an orthodox cladist, Schlee created no suprageneric taxon for these two genera. Later, Zherikhin (1980: 51, 80) introduced a family name for them, separating Bernaeidae from Aleyrodidae (all other whiteflies); however, Bernaeidae published in the chronological table without reference to any characters is a nomen nudum (ICZN Article 13e). Living whiteflies are unusual among insects in that their tax- onomy is based on the immatures, and not on imagines which are generally neglected and imperfectly known. Schlee underestimated the structural plasticity of Aleyrodidae; in fact, some bernaeid characters still persist in several extant genera. These data, along with information on new bernaeid taxa from the Late Mesozoic of Asia, Burmese amber (all described below), and Lebanese amber (D. Shcherbakov & D. Azar, pers. comm.), allow the revision of the diagnosis and rank of the group. The diagnostic characters of bernaeids are discussed consecu- tively: 1. Relative head width is variable: the head is generally narrow in Udamoselinae, but sometimes (0).85 as wide as the thorax inAley- rodinae (asinsome species of Aleyrodes Latreille: Gomez-Menor, 1944: fig. 3); the remaining difference could be attributed to the better developed and more globose eyes in bernaeids; two bernaeid genera described below have a narrow head. PRIMITIVE WHITEFLIES OF ASIA AND BURMESE AMBER 2. Degree of separation between the upper and lower eye parts in Aleyrodidae varies from total to almost none (e.g. in Ceraleurodicus splendidus; Bondar, 1923: fig. 4). 3. The median ocellus is retained in Paraleyrodes (Gill, 1990) and possibly also in Aleurodicus destructor (R.J. Gill, pers. comm.); itis either small or lost in some undescribed Lebanese bernaeids. 4. 8-segmented antennae are recorded in Aleurodicus destructor from Thailand (R.J. Gill, pers. comm.); in bernaeids antennae are usually 11-, sometimes 10-, 8-, and even 5(?)-segmented. 5. Paronychium (empodium) is broad, spatulate, longer than claws in Aleurochiton Tullgren (s. s.; Sampson, 1943); microscopic paronychium is invisible in compression fossils, and in the amber bernaeids its shape could vary from leg to leg, depend- ing on preservation (see Schlee, 1970: fig. 16). 6. In modern whiteflies the wax produced by the abdominal plates is distributed over the body with all three pairs of tibiae possessing specialized setation (Navone, 1987; Byrne & Hadley, 1988). In Bernaea the setal combs on the mid and hind tibia (see Schlee, 1970: fig. 16) appear very similar to those of the present-day whiteflies (R.J. Gill, pers. comm.). In various undescribed bernaeids from Lebanese amber the setal rows on tibiae are also somewhat differentiated, and, moreover, the plates themselves are sometimes observable as well. One could assume that the wax-secreting areas on the abdominal venter (perhaps initially not as clear-cut plates) along with complex waxing behaviour were already acquired by the first (Jurassic) aleyrodoids. 7. Parameres are held horizontally in Aleyrodidae, but those of Bemisia Quaintance & Baker turn upward at a rather abrupt angle (R.J. Gill, pers. comm.). 8, 9. Both ovipositor and rostrum are extremely long in Aleurotithius Quaintance & Baker, allowing it to feed and oviposit on its host plant which is covered with very dense, long hairs (Quaintance & Baker, 1914). This long ovipositor, as well as the normal, short whitefly ovipositor (Weber, 1935: figs. 37-38), has changeable orientation: folded upwards at rest, moving posteriorly when in use. An outline of abdominal apex in Bernaea implies that its ‘horizontal’ ovipositor (directed posteriorly) was already capable of moving upwards, its tip fitting just beneath the modified anal tube (Schlee, 1970: figs. 14, 15; the tube homologized with fused 8—10th tergites (after Weber, 1935), but at least in Bernaea the 8th tergite is still free (fig. 15)). The rostrum is of variable length in bernaeids, either just reaching, or extending beyond, the hind coxae (the former condition being quite similar to the typical aleyrodid one). 10. Finally, the forewing venation of bernaeids is even more di- verse than the aleyrodid one: from the most complete version known for Aleyrodomorpha, towards reduction of M and A, then of R1 and CuA (thus only R-Rs and claval furrow left), up to a nearly veinless wing blade in Heidea (see Schlee, 1970: figs. 3, 6, 7). Itis noteworthy that the reduction trend is the same as across living Aleyrodidae, from Udamoselis to typical Aleyrodinae. Hindwing CuA seems to be always developed in bernaeids. A few other characters show somewhat different distribution in bernaeids than in other whiteflies. A coronal suture is retained in most bernaeids (not recorded in Bernaea and Heidea), and lost in aleyrodids. R1 originates before the forewing midlength in bernaeids, and usually beyond it in Aleyrodidae. The basitarsus is usually longer than the distitarsus in bernaeids, and usually subequal to it in aleyrodids. Therefore, neither venation nor body structure (possibly except 31 male genitalia) are reliable in discriminating between bernaeids and typical whiteflies, most characters demonstrating intergradation. Moreover, the earliest member of Udamoselinae from Lebanese amber (D. Shcherbakov & D. Azar, pers. obs.) shows, along with the male genitalia and venation typical of the subfamily, such bernaeid features as wide head, rounded eyes, and long rostrum. No one bernaeid character state could be regarded as apomorphic (including a long rostrum: in Mesozoic homopterans rostra are commonly longer than in their living descendants), so this group is paraphyletic relative to extant whiteflies. Bernaeids are more similar to Udamoselinae, e.g. in the forewing C often thickened, and wax plates on 3—6th abdominal segments in the female. In turn, Udamoselinae are ancestral to Aleyrodinae (Bondar, 1923: fig. 1) rather than constituting its sister group (Campbell et al., 1995b). Instead of separating Bernaeidae from all other whiteflies, it seems reasonable to treat them as a taxon of the same rank as Udamoselinae (=Aleurodicinae) and Aleyrodinae, i.e. as a third subfamily of Aleyrodidae s./., or, alternatively, as one of three families within Aleyrodoidea (the former opinion is accepted herein). SYSTEMATIC DESCRIPTIONS The material described is in the collections of the Department of Palaeontology, The Natural History Museum, London (register num- bers with the prefix In.), and in the Palaeontological Institute, Russian Academy of Sciences, Moscow (register numbers with the prefix PIN). Family ALEYRODIDAE Westwood, 1840 Subfamily BERNAEINAE subfam. nov. [Bernaeidae Zherikhin, 1980: 80, nomen nudum] DIAGNOSIS. Imago. Head usually as wide as thorax, with coronal suture. Eyes entire, rounded. Median ocellus usually retained. Ant- ennae (5)8—11-segmented. Forewing with convex R-RI1 (RI separating before wing midlength), concave Rs, faint concave M, convex CuA, concave claval furrow (CuP), and oblique convex A; C often thickened; M often, and sometimes also CuA or nearly all veins, reduced. Rostrum reaching at least hind coxae. Basitarsus usually longer than distitarsus. Paronychium broad. Wax plates on 3—-6(7?)th abdominal segments in female. Male parameres held vertically. Ovipositor relatively long. COMPOSITION. 4 genera (plus several undescribed ones), Upper Jurassic to Cretaceous (?and Paleogene) of Asia. REMARKS. In two other, extant subfamilies: head narrower than thorax, without coronal suture; eyes subdivided into the dorsal and ventral part (or at least anteriorly emarginate); median ocellus usu- ally absent; antennae 3—7(8)-segmented; forewing lacking M, with R1 origin usually beyond midlength; rostrum usually shorter; basi- and distitarsus usually subequal; paronychium usually blade- or spine-like, or reduced; parameres held rather horizontally; oviposi- tor usually short. In Aleyrodinae, C is never thickened, and wax plates on 34th abdominal segments in female. In Bernaea neocomica the anal vein seems to be developed like in Ceraleurodicus bakeri (Bondar) (Bondar, 1923: fig. 8), i.e. oblique and very close to the forewing base, as could be deduced from Schlee’s (1970) fig. 25. The impression fossils of Bernaeinae are often preserved with the anterodistal segment of the forewings folded back. Oo i) Genus JULEYRODES gen. nov. TYPE SPECIES. Juleyrodes gilli sp. nov.; Upper Jurassic or Lower Cretaceous; Mongolia. NAME. From the Jurassic and the genus Aleyrodes. DESCRIPTION. Forewing widened towards shallowly rounded api- cal margin. Costal margin markedly arched and thickened (especially near base). Rs 3-4 times longer than R stem. Complete M developed basad of R1 origin (about 1/3 of wing length) and joining CuA basally, nearer to CuA distally. Anal vein occupying 1/2 of clavus. Head narrower than thorax. Antennal flagellum slender. OTHER SPECIES. J. visnyai sp. nov., Late Jurassic of Kazakhstan. REMARKS. Distinct in complete M, short R stem, long A, and forewing shape. Figs. 2-5 HOLOTYPE. PIN 4307/231, right forewing (part and counterpart; costal margin incomplete); central Mongolia, Ara-Hangayn aymag, 6 km W of Hotont somon, northern part of Uhaa Mt. (Hotont locality, outcrop 354/7); Upper Jurassic or Lower Cretaceous. Juleyrodes gilli sp. nov. NAME. After Dr R.J. Gill, an expert in living whiteflies. DESCRIPTION. Forewing about twice longer than wide, gradually widening towards an obliquely and shallowly rounded apical margin. Fig.2 Juleyrodes gilli sp. noy., holotype forewing PIN 4307/231, Hotont. R1 CuP Al Fig.3 Juleyrodes gilli sp. nov., holotype (venation). D.E. SHCHERBAKOV Fig. 4 Juleyrodes ?gilli sp. nov., forewing PIN 3965/445, Hutel-Hara. Fig.5 Juleyrodes ?gilli sp. nov. (venation); the wing segment folded back in the impression (arrows) shown in natural position. Costal margin markedly arched and thickened proximally, more so towards base. R stem straight, raised. R1 longitudinal, 2.6 times longer than R stem, originating at 0.3 wing length, convex and subparallel to C proximally, faint distally, ending well beyond Rs midlength. Rs as fine groove, about 4 times as long as R stem. M clearly joining CuA, continued basad as a groove along M+CuA fusion (which is half as long as the R stem), proximally as unpig- mented groove, distally as faint vein nearer to CuA. CuA as faint convex vein, A as fine convex one (turning faint distally). Clavus occupying about 2/3 of wing length; anal vein rather long, 1/2 of clavus (nearly 1/3 of wing length). Forewing dusky, more so along C and R stem, and in clavus, with dark veins (especially C, R, and A). MEASUREMENTS. Forewing: length, 2.1 mm; width, 1.1 mm (holotype). MATERIAL. Specimen PIN 3965/445, left forewing (part and coun- terpart; anterodistal wing segment folded back); eastern Mongolia, East-Gobi aymag, 70 km SW of Saynshand somon, eastern Hara-Hutul Range (Hutel-Hara locality, outcrop 300); Upper Jurassic or Lower Cretaceous, lower Tsagaan Tsav Formation. The specimen differing from the holotype in the slightly more elongate wing could represent either another sex of the same species, or one more, closely related species; more pronounced wing relief and less obvious pigmentation are attributable to somewhat different character of preservation. PRIMITIVE WHITEFLIES OF ASIA AND BURMESE AMBER Figs. 6, 7 HOLOTYPE. PIN 2997/3837, complete female? (part and counter- part); southern Kazakhstan, Karatau Range, Mikhailovka; Upper Jurassic, Karabastau Formation; the only specimen. Juleyrodes visnyai sp. nov. NAME. After Dr A. Visnya, a discoverer of Ceraleurodicus kesselyaki. DESCRIPTION. Forewing with apical margin shallowly rounded. Costal margin strongly arched and thickened proximally, even more so towards base. R stem arched, continued with Rs. R1 oblique, 1.8 times longer than R stem, originating at 0.33 wing length, markedly converging with C, ending before Rs midlength. Rs 3.3 times longer than R stem. M poorly traceable (as a groove along M+CuA and distally nearer to CuA). Forewing dusky, with dark veins. Hindwing more transparent, with dark veins, distinct R1 originating distally. Body squat. Head 3/4 as wide as the mesothorax, with anterior margin produced rounded trapezoidal between eyes. Antennal flagellum slender, multisegmented. Legs untraceable. Abdomen ter- minating in two rounded lateral lobes, and dorsomedian projection (anal tube); beyond the latter only traces of presumed ovipositor are visible (on positive impression). MEASUREMENTS. Forewing length, 3.2 mm; body length (exclud- ing ovipositor), 3.0 mm; antenna length, 0.9 mm; head width, 0.7 mm; mesothorax width, 0.95 mm. REMARKS. Distinct from the type species in the larger size, broader costal area, arched R stem, and shorter, oblique R1. From the same locality, two more whitefly specimens are known, less completely preserved, and smaller in size than J. visnyai holotype (female?). Both have the head narrower than the thorax, and are presumably males. PIN 2997/5071 (body 2.05 mm, forewing 2.2 mm long) is attributable to the genus Juleyrodes on account of its markedly arched and thickened C; despite dissimilar head shape (Fig. 8), it could even turn out to be a male of J. visnyai (however, sexual dimorphism is not widespread in whiteflies, e.g. in Ceraleurodicus kesselyaki, where the male, bearing very long parameres, is larger than female). PIN 2239/532 (body and forewing length 2.3 mm) with less arched C could be identified only as Bernaeinae gen. indet., possibly related to one of two narrow-headed genera described Fig.6 Juleyrodes visnyai sp. nov., holotype, female? PIN 2997/3837 (positive impression), Karatau. 33 Fig.8 Juleyrodes sp., male? PIN 2997/5071, Karatau. herein; it has some eye and genital structures which cannot be interpreted at present, and probably some wax plates (preserved as shaded areas on the abdomen) (Fig. 9). Genus BURMOSELIS gen. nov. TYPE SPECIES. Burmoselis evelynae sp. nov.; probably Upper Cre- taceous; Burmese amber. NAME. From Burma (now Myanmar) and genus Udamoselis. DESCRIPTION. Forewing widened towards obliquely rounded apex. Costal margin weakly arched. Rs about twice as long as the R stem. Nearly complete M starting slightly basad of R1 origin (just before Fig.9 Bernaeinae gen. indet., male? PIN 2239/532, Karatau. wing midlength), nearer to CuA distally. Anal vein occupying 1/3 of clavus. Head narrower than thorax. Antennae 11-segmented, flagellum slender. Basitarsus no longer than distitarsus. Ovipositor long. OTHER SPECIES. None. REMARKS. Similar to Juleyrodes and can be separated from Leba- nese amber genera (including several undescribed ones) in the almost complete M nearer to CuA distally, narrower head and shorter basitarsus. Distinct from Juleyrodes in the shorter M and A, longer R stem, and less arched costal margin. Burmoselis evelynae sp. nov. Figs. 105 11 HOLOTYPE. NHM Palaeontol. Dept., In.20193, complete female near the edge of a large amber piece containing several insects, a spider and a mite (the whitefly near the spider); Burmese amber; probably Upper Cretaceous, Hukawng Valley, Myanmar (Burma) (see Zherikhin & Ross, this volume); the only specimen. NAME. coccids. After Dr Evelyna Danzig, an authority on aleyrodids and DESCRIPTION. Forewing 2.2 times as long as wide, slightly dusky, gradually widening towards obliquely rounded apex. Costal margin weakly and evenly arched; C slightly thickened proximally. R stem nearly straight, raised. R1 almost as long as R stem, originating at 0.44 wing length, straight, faint distally, strongly converging with C. Rs twice as long as the R stem. M traceable from just beyond R1 origin (not joining CuA), distally nearer to CuA. CuA fine convex, slightly bent against R1 origin, probably marking the end of M+CuA fusion (then M+CuA equal to R stem). Clavus occupying 2/3 of wing length; anal vein short, about 1/3 of clavus and 1/5 of wing length. Hindwing 2.4 times as long as wide, rounded apically, transparent; origin of indistinct R1 just beyond wing midlength. Body shrivelled. Head 3/4 as wide as the mesothorax, with anterior margin slightly biconvex and medially emarginate (indication of coronal suture, if not an artifact of preservation). Antenna 11-segmented, apices of flagellomeres marked with rhinaria; | st flagellomere almost as long as three following combined; pedicel large, elongate, obliquely truncate apically. Rostrum invisible, but (judging from the size of a D.E. SHCHERBAKOV — _ JME “ie a — 2 = Woe. “€ Fig. 10 Burmoselis evelynae sp. noy., holotype, female NHM In.20193, Burmese amber. large gas bubble enclosing it) possibly reaching hind coxae. Pronotum preserved almost in vertical plane. Legs moderately long, tibial setae not conspicuous (visible at least on fore tibiae). Basitarsus subequal to distitarsus in hind leg, and slightly shorter than it in other legs. Paronychium presumably small. Pregenital abdomen largely mem- branous (preserved markedly flattened dorsoventrally). Paired dark areas visible on five(?) abdominal segments, possibly representing wax plates. Ovipositor directed posteriorly, projecting beyond anal tube for about 1/2 length of the rest of abdomen, with 4 long lateral bristles on each side. MEASUREMENTS (mm). Forewing length, 1.1, its width, 0.5; hindwing length, 0.95, its width, 0.4; body length (including oviposi- tor), 0.95; mesothorax width, 0.28; antenna length, 0.3; estimated rostrum length, 0.3; fore tibia length, 0.23, fore tarsus length, 0.1; hind tibia length, 0.4, hind tarsus length, 0.18; projecting part of ovipositor, 0.2. OTHER FOSSIL WHITEFLIES Imagines Middle Purbeck of England (Lower Cretaceous: Berriasian): a forewing of Juleyrodes sp. was recently found (D. Shcherbakov & R. Coram, pers. comm.). Lebanese amber (Lower Cretaceous): whiteflies are exceptionally numerous, about 70% of homopterans, and 9% of all insects! (Poinar, 1992): Bernaea neocomica, Heidea cretacica (Schlee, 1970), and several undescribed genera, including the oldest member of Udamoselinae (D. Shcherbakov & D. Azar, pers. obs.). PRIMITIVE WHITEFLIES OF ASIA AND BURMESE AMBER : is . f : i Ah wo m SN: RNY i SHE 8 | ' : WS a) ' ‘ SS SSNS SSS ARS \ 35 ANS . ty, BV “yy 6S S —~——" ‘tl Fig.11 Burmoselis evelynae sp. nov., holotype: (a) habitus, dorsal; (b) antenna, schematized, first segment not shown; (c) profile of abdomen. Burmese amber: “Aleurodicus’ burmiticus Cockerell, 1919, 1 male (Fig. 12). Schlee (1970: 32) doubted its assignment to Aleurodicinae, but the hindwing venation and genitalia confirm the subfamily placement (generic assignment doubtful). - te Fig.12 ‘Aleurodicus’ burmiticus Cockerell, holotype, NHM In.19134, Burmese amber. Baltic amber: ‘Aleyrodes’ aculeatus Menge, 1856, 1 female? (holotype presumably destroyed), not figured, and diagnosed only as ‘similar to living A. chelidonii Latr., except for the end of abdomen being acuminate and bearing two small pointed processes, which was found in both sexes by Burmeister (Entomol. II: 82)’, so generic position is undeterminable. There are 15 aleyrodid specimens in Copenhagen collection (Larsson, 1978), partly studied but not named by Schlee (1970); 2 undescribed specimens in the PIN collection and 3 unstudied specimens (2 imagines, | pupa?) in the NHM (A. Ross, pers. comm.). Whiteflies are also recorded in Mexican and Dominican amber (Poinar, 1992). Nymphs Purbeck and Wealden of England (Lower Cretaceous; Berriasian- Barremian): very small oval nymphs occur regularly (more rarely in Wealden), resembling pupal cases of Aleyrodoidea (Jarzembowski & Coram, 1997: figs. 6-8). Lithographic Limestone of Montsech (Lower Cretaceous; Berriasian): pupal case of ?7Bernaeinae incertae sedis (Whalley & Jarzembowski, 1985: figs 12-13). Eocene (Isle of Wight): pupal case of Aleyrodoidea (Jarzembowski & Ross, 1993: fig. 2). Pliocene (Hessen): Aleurochiton petri Rietschel, 1983, pupal case. Taxa excluded from Aleyrodomorpha Upper Permian: Permaleurodes rotundatum Becker-Migdisova, 1959 (South Siberia) and Aleuronympha bibulla Riek, 1974 (South Africa), 36 both nymphs, and assigned to a separate family Permaleurodidae Becker-Migdisova, 1959 in the Aleyrodoidea. The former genus was regarded by Evans (1963), Hennig (1969) and Schlee (1970) as a doubtful member of Aleyrodomorpha, and by Mound & Halsey (1978) even as acockroach nymph. In fact both these nymphs belong to primitive Homoptera, most probably to Protopsyllidiidae or re- lated group of Psyllinea. Lower Cretaceous (Brazil): Megaleurodes megocellata Hamilton, 1990, ascribed to Aleyrodoidea and tentatively assigned to the Permian family Boreoscytidae. The genus is possibly based on a poorly preserved planthopper, and has nothing in common with boreoscytids (primitive group of Aphidinea). AN OVERVIEW OF BURMESE AMBER HEMIPTERA Of 1200 animal inclusions recorded in Burmese amber from NHM collection by A.P. Rasnitsyn, 75 are Homoptera and 9 Heteroptera, so Hemiptera totalling 84 specimens, or 7%, are in 6th place, after Isoptera and before Diplopoda. (It is the same percentage as for Dominican amber Hemiptera from the collection of Smithsonian Institution, identified by A.P. Rasnitsyn). More than half (47 speci- mens) are Cicadinea (=Auchenorrhyncha), dominated by the extant family Achilidae (Fulgoroidea; 27 specimens). Achilids, feeding on fungi and often corticolous, are numerous (but not dominating) in Baltic amber as well. Burmese amber achilids are represented by both nymphs (including exuvia) and imagines. Only two specimens are identified, both of ‘Liburina’ burmitina Cockerell (erroneously assigned to the Delphacidae genus Liburnia Stal): the holotype, In.19105, and another specimen, In.20150(1). The holotype was re- examined and shows no metatibial spur, and the forewing venation is characteristic of Achilidae; to elucidate the generic position of the species, it should be compared to several extinct genera described from the Cretaceous and Baltic amber. A second fulgoroid family, the extant Cixiidae, is represented by two imagines: Plecophlebus nebulosus Cockerell, holotype In.19094, and one specimen of another genus. Plecophlebus was described in Trichoptera, but later transferred to Fulgoroidea (Botosaneanu, 1981). A third, extinct fulgoroid family is of exceptional interest: 2 imag- ines of different genera belong to the group otherwise occurring in the Aptian of Mongolia and Cretaceous ambers of Taimyr and New Jersey (Shcherbakov, in prep.). 11 more planthopper imagines and nymphs are at present not determinable to family level. Total Fulgoroidea (42 specimens) constitute most of Cicadinea, like in Dominican amber. The only other auchenorrhynchous group deter- mined is Cercopoidea (possibly Aphrophoridae, 1 imago), always rare in ambers. The remaining 4 specimens of Cicadinea are undeter- minable. Other Homoptera (‘Sternorrhyncha’ ) are represented with 19(?+1) coccids (males, females, and possibly nymphs) and 3 whiteflies: “Aleurodicus’ burmiticus Cockerell, 1919 (Udamoselinae s.1.), holotype male In.19134, Burmoselis evelynae gen. et sp. nov. (Bernaeinae), holotype female In.20193a, and an undetermined aleyrodid, In.20703. Coexistence of Bernaeinae and Udamoselinae is otherwise known only in Early Cretaceous Lebanese amber. A further 5 specimens of Homoptera are indeterminable. Six out of nine Heteroptera are Enicocephalidae (such an unusu- ally high proportion is comparable only to that in Lebanese amber): Disphaerocephalus constrictus Cockerell, holotype In.19112; D. macropterus Cockerell, holotype In.19123(1); Electrocephalus swinhoei Cockerell, holotype In.19113; Paenicotechys fossilis D.E. SHCHERBAKOV (Cockerell), holotype In.19095; Enicocephalidae indet. (2 speci- mens). Other families are Coreidae s.]. and Ochteridae, one specimen of each (Yu. A. Popov det.); one more specimen is tentatively determined as a heteropteran. Psyllomorpha are very rare or absent in Cretaceous and Paleogene faunas, and Burmese amber is not an exception. However, two other homopteran groups, leafhoppers (Cicadellidae s./.) and aphids, are surprisingly lacking from the fauna discussed. Aphids are abundant and diverse since the Early Cretaceous, and well represented in Taimyr, Canadian and Baltic ambers; however, they are very rare in Lebanese amber (unpubl. data), virtually absent from Dominican amber (only 2 specimens recorded; Heie & Poinar, 1988; Wegierek, 1998), and now mainly extratropical, so their absence from Burmese amber could be evidence of a tropical paleoclimate. Leafhoppers, likewise abundant since the Early Cretaceous in compression fossil faunas, are (in contrast to aphids) not yet recorded in Cretaceous ambers, well represented in all Cenozoic ambers, and are now diverse on all continents and large islands (including Australia and Madagascar). So their absence is more intriguing and may be due to taphonomical reasons. ACKNOWLEDGEMENTS. | am grateful to Mr A.J. Ross (The Natural His- tory Museum, London) and Dr A.P. Rasnitsyn (Paleontological Institute, Russian Academy of Sciences, Moscow) for an opportunity to study Burmese amber Hemiptera, to Mr D. Azar (Museum National d’ Histoire Naturelle, Paris) for the loan of Lebanese amber whiteflies from his collection, to Dr R.J. Gill (Plant Pest Diagnostics Center, Department of Food and Agriculture, Sacramento) and Dr E.M. Danzig (Zoological Institute, Russian Academy of Sciences, St. Petersburg) for valuable discussion, and to Peter York (NHM) for photography. REFERENCES Becker-Migdisova E.E. 1959. Some new representatives of Sternorrhyncha from the Permian and Mesozoic of the USSR. In, Materialy k ‘Osnovam paleontologii’, 3: 104-116. Paleontologicheskii Institut Akademii Nauk SSSR: Moscow. Bemis F.E. 1904. The aleyrodids, or mealy-winged flies, of California, with references to other American species. Proceedings U. S. National Museum, 27: 471-537. Bondar G. 1923. Aleyrodideos do Brasil. Imprensa Official do Estado: Bahia. Borner C. 1904. Zur Systematik der Hexapoden. Zoologischer Anzeiger, 27: 511-533. — 1910. Die Fliigelgeaderung der Aphidina und Psyllina. Zoologischer Anzeiger, 36 (1): 16-24. Botosaneanu L. 1981. On a false and a genuine caddis-fly from Burmese amber (Insecta: Trichoptera, Homoptera). Bulletin Zoologisch Museum Universiteit van Amsterdam, 8: 73-78. Byrne D.N., Buchmann S.L. & Spangler H.G. 1988. Relationship between wing loading, wingbeat frequency and body mass in homopterous insects. Journal of Experimental Biology, 135: 9-23. & Hadley N.F. 1988. Particulate surface waxes of whiteflies: morphology, composition and waxing behaviour. Physiological Entomology, 13: 267-276. Campbell B.C., Steffen-Campbell J.D. & Gill R.J. 1994. Evolutionary origin of whiteflies (Hemiptera: Sternorrhyncha: Aleyrodidae) inferred from 18S rDNA sequences. /nsect Molecular Biology, 3 (2): 73-88. 5 , Sorensen J.T., & Gill R.J. 1995a. Paraphyly of Homoptera and Auchenorrhyncha inferred from 18S rDNA nucleotide sequences. Systematic Ento- mology, 20 (3): 175-194. , & Gill R.J. 1995b. Origin and radiation of whiteflies: an initial molecular phylogenetic assessment. Jn, Gerling, D. & Mayer, R.T. (editors), Bemisia: 1995 Taxonomy, biology, damage control and management: 29-51. Intercept: Andover. Carver M. 1991. Superfamily Aleyrodoidea. Jn, Naumann, I.D. (editor), The insects of Australia, 2nd ed.: 450-452. Melbourne Univ. Press: Carlton. Cockerell T.D.A. 1919. Insects in Burmese amber. Entomologist, 52: 241-243. Dohlen C.D.von & Moran N.A. 1995. Molecular phylogeny of the Homoptera: a paraphyletic taxon. Journal of Molecular Evolution, 41: 211-223. Enderlein G. 1909. Udamoselis, eine neue Aleurodiden-Gattung. Zoologischer Anzeiger, 34: 230-233. Evans J.W. 1963. The phylogeny of the Homoptera. Annual Review of Entomology, 8: 77-94. PRIMITIVE WHITEFLIES OF ASIA AND BURMESE AMBER Gill R.J. 1990. The morphology of whiteflies. In: D. Gerling (ed.). Whiteflies: their bionomics, pest status and management: 13-46. Intercept: Andover. Gomez-Menor J. 1944. Aleirodidos de interes agricola. Trabajos de Estacion Fitopatologica y Agricola Madrid, Ser. Fitopatologica, 133: 1-36. Hamilton K.G.A. 1990. Homoptera. In, D.A. Grimaldi (ed.). Insects from the Santana Formation, Lower Cretaceous, of Brazil. Bulletin of the American Museum of Natural History, 195: 82-122. Haupt H. 1934. Neues iiber die Homoptera-Aleurodina. (Stellung im System, dussere Anatomie, Biologie, 2 neue Arten.) Deutsche Entomologische Zeitschrift, 1934: 127-141. Heie O. & Poinar G.O., Jr. 1988. Mindazerius dominicanus nov. gen., nov. sp., a fossil aphid (Homoptera, Aphidoidea, Drepanosiphidae) from Dominican amber. Psyche, 95: 153-165. Hennig W. 1969. Die Stammesgeschichte der Insekten. Senckenberg-Buch, 49. W.Kramer: Frankfurt am Main. Jarzembowski E.A. & Coram R. 1997. New fossil insect records from the Purbeck of Dorset and the Wealden of the Weald. Proceedings of the Dorset Natural History and Archaeological Society, 118: 119-124. & Ross A. 1993. Time flies: the geological record of insects. Geology Today, Nov— Dec. 1993: 218-223. Larsson S.G. 1978. Baltic amber — a palaeobiological study. Entomonograph, 1. Scandinavian Science Press: Klampenborg. MengeA. 1856. Lebenszeichen vorweltlicher, im bernstein eingeschlossener thiere. In: Programm, . . . offentlichen Priifung der Schiiler der Petrischule, pp. 1-32. A.W. Kafemann: Danzig. Mound L.A. & Halsey S.H. 1978. Whitefly of the world. A systematic catalogue of the Aleyrodidae (Homoptera) with host plant and natural enemy data. British Museum (Natural History) & J. Wiley: Chichester. Navone P. 1987. Origine, struttura e funzioni di escreti e secreti entomatici di aspetto ceroso distribuiti sul corpo mediante zampe. Anni di Facolti di Sciencia Agraria dell’ Universita di Torino, 14: 237-294. Poinar G.O., Jr. 1992. Life in amber. Stanford University Press: Stanford. Quaintance A.L. & Baker A.C. 1913. Classification of the Aleyrodidae. Part I. Technical Series, Bureau of Entomology, U. S. Department of Agriculture, 27(1): 1— 93) & 1914. Classification of the Aleyrodidae. Part II. Technical Series, Bureau of Entomology, U. S. Department of Agriculture, 27 (2): 95-109. Riek E.F. 1974. An unusual immature insect from the Upper Permian of Natal. Annals of Natal Museum, 22 (1): 271-274. Rietschel S. 1983. Aleurochiton petri n. sp., eine Mottenschildlaus (Homoptera, oy Aleyrodina) aus dem Pliozan von Neu-Isenburg, Hessen. Carolinea, 41: 97-100. Sampson W.W. 1943. A generic synopsis of the hemipterous superfamily Aleyrodoidea. Entomologica Americana, 23 (3): 173-223. & Drews E.A. 1957. Keys to the genera of the Aleyrodinae and notes on certain genera (Homoptera: Aleyrodinae). Annals and Magazine of Natural History, 9: 689— 697. Schlee D. 1969. Sperma—Ubertragung (und andere Merkmale) in ihrer Bedeutung fiir das phylogenetische System der Sternorrhyncha (Insecta, Hemiptera). Phylogenetische Studien an Hemiptera. I. Psylliformes (Psyllina and Aleyrodina) als monophyletische Gruppe. Zeitschrift fiir Morphologie der Tiere, 64: 95-138. 1970. Verwandtschaftsforschung an fossilen und rezenten Aleyrodina (Insecta, Hemiptera). Stuttgarter Beitrage zur Naturkunde, 213: 1-72. Shcherbakoy D.E. 1990. Extinct four-winged ancestors of scale insects (Homoptera: Sternorrhyncha). Proceedings of the 6th International Symposium of Scale Insect Studies, Cracow, II: 23-29. 1996. Origin and evolution of the Auchenorrhyncha as shown by the fossil record. In C.W. Schaefer (ed.). Studies on Hemipteran Phylogeny: 31-45. Entomological Society of America: Lanham (Thomas Say Publications in Entomology: Proceed- ings). Solomon M.E. 1935. On a new genus and two new species of Western Australian Aleyrodidae. Journal and Proceedings of the Royal Society of Western Australia, 21: 75-91. Szelegiewicz H. 1971. Cechy autapomorficzne w budowie skrydel Sternorrhyncha (Hemiptera) i ich znaczenie dla oceny paleozoicznych przedstawicieli tej grupy pluskwiakow. Annales Zoologici, 29 (2): 15—81. [English translation 1976, National Center for Scientific, Technical and Economic Information: Warsaw. ] Visnya A. 1941. A gigantic species of Aleurodidae (Homoptera) from greenhouse-orchideas. Folia Entomologica Hungarica, 6 (1/2): 4-15. Weber H. 1935. Der Bau der Imago der Aleurodinen. Ein Beitrag zur vergleichenden Morphologie des Insektenk6rpers. Zoologica (Stuttgart), 33: 1-71. Wegierek P. 1998. Fossil representatives of the family Greenideidae (Hemiptera, Aphidoidea). First Paleoentomological Conference, 1998, Abstracts: 47. Moscow: Paleontological Institute of the Russian Academy of Sciences. Whalley, P.E.S. & Jarzembowski, E.A. 1985. Fossil insects from the Lithographic Limestone of Montsech (late Jurassic-early Cretaceous), Lérida Province, Spain. Bulletin of the British Museum of Natural History (Geology), 38 (5): 381-412. Wootton R.J. & Newman D.J.S. 1979. Whitefly have the highest contraction frequen- cies yet recorded in non-fibrillar flight muscles. Nature, 280: 402-403. Zherikhin V.V. 1980. Class Insecta. In: Razvitie i smena bespozvonochnyh na rubezhe mezozoya i kainozoya: 40-97. Nauka: Moscow. one ; or 5 fae Mesos fuam's Rercundgpateetilig en eee an eee ae ee tbp—mw.cP & pi Are Cte rhs autora a tienen’ ae ei al >i alae <, rN, >» f aS et, in ; mae 49 de oo Lgl Pe caphoay %, Sh Beth Ch eid “ste cate me ——— oh hepa panes o % oer? ¥ re a nate Weta: Sok ae alah aay ys ere. te hem ; decal rit eau eee ghana bee iN iy ny ‘ 04 4d, car ry Pa 7 Ae etaneas ie a e ae: ae ' oer A pants 8 Genet satire er Li iend : miei wes iA Rg a hear si bia hes bye 1 ec ures tips! Laos) ‘ee fh tS Pe iy ) , ‘oe 8S ; uk fs BAL C74! As Bis § PR eS to a ‘ : ii ian 4 5 rn as 4 ; cf — Se %, 6 ) : 4 5 — =a end i} me —o - roll ; all Del i +4: _ = Ue pe 4, I = ; f > i] 4 fy 1 t = t It t ~ os TKS « é Dei ae re etye ~~— Rae sw vite le chet Wal aes ies Siparse ion ait ot ad reer pre Per Penge dant 1 ated Sisaetisehallnhy i aah Hy aan oa ee << aw aes = ea a ih aig i mare Se oP le —— vol Saal ) Jim ae , « ii aaah A ors) wy * 40? ; cia a eed ® Se 7 > ee oe aonecr ih A! A eR, 5 ee oe by re SF area. = 2 — ‘ i. 7 & @ t 7 A ‘kh - : t 4 ee — an J 7 ; ¥ Sag f ; 5 om 5 cto ae aoe | - i i a f < \ < x D) g ~ f Qo i “ ‘oN Bull. nat. Hist. Mus. Lond. (Geol.) 56(1): 39-41 Issued 29 June 2000 A new genus and species of Lophioneuridae from Burmese amber (Thripida (=Thysanoptera): Lophioneurina) V. V. ZHERIKHIN Arthropod Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya Str. 123, Moscow 117647, Russia Synopsis. Burmacypha longicornis, gen. et sp. nov., is described and placed in the subfamily Lophioneurinae within the family Lophioneuridae (Thysanoptera =Thripida). Burmacypha has unusual wing venation but seems to be related to the Cretaceous genera Undacypha and Jantardachus. It represents a Mesozoic element in the Burmese amber fauna. INTRODUCTION The extinct family Lophioneuridae was originally established by Tillyard (1921) in the order Homoptera; later it was transferred to Psocoptera (Tillyard, 1935) and then united with living Thysanoptera (=Thripida) (Zherikhin, 1980). Lophioneurids are synapomorphous with Thysanoptera s.str. in the short mouth cone formed by stylet-like mandibles and laciniae and elongate labium, the two-segmented tarsi with large eversible pulvilla and small claws, and the loss of outer valvulae of the ovipositor; they are more primitive in retaining the symmetrical mouthparts, antennae inserted below eyes and relatively broad wings with complete venation (Vishniakova, 1981). Lophioneuridae were long-lived and probably distributed world- wide as documented by numerous finds in the Permian of Australia, North America and Asia, as well as in the Mesozoic of Europe, Asia and Australia. Two subfamilies are recognized, Lophioneurinae and Zoropsocinae (Tillyard, 1935). The youngest lophioneurids described up to now are preserved in amber from Taymyr, Siberia (Vishniakova, 1981) which is Santonian in age. A new species from Burmese amber described below represents a new genus. The vein nomenclature below is after Vishniakova (1981). SYSTEMATIC DESCRIPTIONS Order THRIPIDA Suborder LOPHIONEURINA Family LOPHIONEURIDAE Tillyard, 1921 Subfamily LOPHIONEURINAE Tillyard, 1921 Genus BURMACYPAHA nov. TYPE SPECIES. Burmacypha longicornis sp. nov.; Burmese amber, probably Upper Cretaceous, Hukawng Valley, Myanmar (Burma). DiAGNosis. A genus of Lophioneurinae with forewings sclerotized, membrane distinctly cellulate; Sc present, R, M and Cu fused forming long common stem, then R+M and Cu diverge and connect again before apex forming large discal cell. A completely reduced hindwing lacking IR, posterior branch of RS fork ending well behind | wing apex, M perpendicular to R+M stem. Burmacypha longicornis gen. & sp. nov. Figs 1-3 \ |HOLOTYPE. NHM Pal. Dept. In.20194. Inclusion in Burmese amber, © The Natural History Museum, 2000 probably Upper Cretaceous, Hukawng Valley, Myanmar (Burma), (see Zherikhin & Ross, this volume). DIAGNOSIS. mm. Species of Burmacypha with forewing length 0.75 DESCRIPTION. Antennae 7-segmented, long, exerted just before eyes, clearly longer than midlength of forewing; scape and pedicel cylindrical, distinctly longer than broad, flagellum very slender, filiform, without distinct pubescence. Eyes round, convex, coarsely facetted. Clypeal area short. Mouth cone elongate, longer than head, symmetrical. Labrum long, convex in lateral view. Labial palpi long, slender, with first 3 segments subequal, much longer than broad; 4th segment as long as 2nd and 3rd combined, widened, pyriform. Structure of stylets unknown. Pronotum short, broad, raised in side view. Forewing broad, about 2.7 times as long as wide, broadly rounded apically, somewhat sclerotized, with fine, clearly cellulate bare surface, not fringed. Anterior margin nearly straight in proximal part. Forewing venation strong, bare. Sc distinct, very close to anterior margin, reaching somewhat less than 0.3 of wing length. R, M and Cu fused in basal third of wing forming a straight common stem nearly equidistant from fore and hind margin: then this vein divides into two symmetrical stems forming a large, heptagonal discal cell: R+M directed obliquely forwards from origin and Cu directed obliquely backwards from origin. After initial branching, both stems become subparallel to wing margins, and after second branching converge to point of origin of next branch. All branches straight, their points of origin on both stems placed directly opposite each other, about 0.4, 0.6 and 0.75 of wing length. R with 3 branches projecting to anterior margin: IR directed obliquely to anterior margin and base of wing and ending near 0.3 of wing length; R,,, directed sub-perpendicularly to R+M stem and ending about 0.6 of wing length; R,, directed obliquely to anterior margin and apex and ending about 0.85 of wing length. M divided from R+M at point of origin of R,,., directed transversally with two branches, anterior and almost opposite R, ., posterior directed obliquely backwards and joining CuA. Cu stem with two branches; CuA perpendicular to stem and CuP connected with posterior M branch. Anal veins absent. Hindwing membranous, transparent, narrow, about as long as forewing and 4.4 times as long as wide, with anterior margin dis- tinctly sinuate and hind margin convex, lacking fringe, with numerous microtrichiae throughout. Veins weak, colourless. Sc absent. R+M forming long common stem. M simple, originating about 0.6 of wing length, perpendicular to R+M stem, ending at the same level at hind margin. R forming large and short fork apically and ending 40 Fig. 1 Burmacypha longicornis gen. et sp. nov. Holotype, In.20194, Burmese amber. Length of forewing 0.75 mm. ee en eae mbar 2s sun STIS ee een OS COPE TR TAS Fie s SES J» ERE MESSY, Pee z Fig. 2 Burmacypha longicornis gen. et sp. nov. Holotype, In.20194, Burmese amber. symmetrically at either side of wing apex. Cu straight, simple, reaching hind margin about 0.35 of wing length. Abdomen not reaching wing apex, segmentation indistinct. Coxae large, conical. Legs long, slender; femora cylindrical, about as long as tibiae; tarsi about half as long as tibiae, with 1st segment very short, 2nd much longer than broad; large apical pulvilla present; claws very small. V.V. ZHERIKHIN Dimensions: forewing length 0.75 mm, width 0.35 mm, antenna length about 0.55 mm, hind tibia length 0.20 mm. REMARKS. The position of the insect inside the amber piece make the precise measurement of many structures impossible, so that some relations in the above description are indicated as approximate only. DISCUSSION. At first glance, Burmacypha seems to be very unu- sual, especially in respect of the forewing venation; however, similar evolutionary trends occur in some previously described genera, especially in Undacypha Vishniakova (from Unda, Transbaikalia- Lower Cretaceous) and Jantardachus Vishniakova (from Siberian amber- Santonian). The homology of the forewing venation accepted here may be disputable on some points; however, this is the most simple homology, not requiring a hypothesis about the appearance of any ‘new’ veins absent in other lophioneurids. The genus is placed in the subfamily Lophioneurinae because of the following features combined: the wings lack a marginal setose fringe; the veins are bare; Cu base in forewings fused with M+R; and M in the hind wings is simple. Burmacypha is certainly a highly apomorphous genus with many derived characters. However, few of them are synapomorphous with other genera, namely the 7-segmented antennae (shared with other Mesozoic genera of Lophioneurinae), the short and raised pronotum (shared with Undacypha), the terminal segment of the labial palpi enlarged (shared with Jantardachus), and IR in the hind wings lost (shared with both Undacypha and Jantardachus). Much more de- rived characters are unique for Burmacypha: the forewing is broad, sclerotized, with a distinctly cellulate surface; R, M and Cu in the forewing are fused for a long distance; the forewing has a peculiar discal cell closed apically by the transversally turned median vein; the arrangement of the veins around the discal cell remarkably symmetrical; the hindwing has the anterior margin angularly sinuate; M in the hind wing is perpendicular to the R+M common stem; the Ist tarsal segment is very short. The long antennae should also be listed here. Though long antennae are plesiomorphous in Lophioneuridae in general, in this primitive state they are at least 10- segmented. In the majority of Mesozoic genera the number of antennal segments is reduced, and antennae shortened, and the 7- segmented antennae in Burmacypha are almost certainly secondarily long. The long mouth cone may also be apomorphous but it is difficult to compare it in detail with those of Undacypha which is somewhat similar at least superficially. On the other hand, Undacypha and Jantardachus are synapomorphous in the reduction of Sc in the forewings while in Burmacypha Sc is long as in the Palaeozoic lophioneurine genera and the Upper Jurassic genus Karataocypha Vishniakova. The long and slender legs of Burmacypha may be plesiomorphous. A similar state occurs in Undacypha whereas the shortened legs of Jantardachus seem to be apomorphous; unfor- tunately, the leg structure is unknown in other Mesozoic Lophioneurinae. All three lophioneurinae genera known from the Cretaceous, namely Undacypha, Jantardachus and Burmacypha, may be related and perhaps belong to a holophyletic unit. However, they do not seem to be very closely related to each other. Probably, only a small part of the Cretaceous lophioneurid diversity is known, and further finds will illuminate the taxonomic structure of this youngest lineage within the family. The abdomen of the holotype of Burmacypha longicornis is somewhat laterally flattened and transparent, with some dark inter- nal structures visible inside. The internal anatomy of lophioneurids is unknown, and the specimen is very interesting in this respect. Unfortunately, it is difficult to interpret the internal structures NEW LOPHIONEURIDAE FROM BURMESE AMBER 41 Fig.3 Burmacypha longicornis gen. et sp. nov. Reconstruction. accurately. Evidently there appears to be the alimentary tract filled by a dark substance. It seems to be large and forming loop(s) which looks like a large dark mass of unregular shape. Three long appendices originating from this dark mass dorsally may represent either the gastral caecae or the Malpighian tubes. Perhaps some internal genital structures are also visible but I have failed to interprete them. There are no traces of the ovipositor which is well observable in other lophioneurids as well as in the less advanced families of the true thrips; this indicates the specimen is most probably a male. The elongate lophioneurid mouthparts forming a cone indicate that like modern primitive thrips, lophioneurids were adapted to the sucking up of individual small objects such as pollen grains, small soft-bodied animals and insect eggs (Zherikhin, 1980). In Burmacypha the mouth cone is comparatively long assuming a rather advanced trophic specialisation. No lophioneurids have been found in the Cenozoic, and Burmacypha certainly represents a Cretaceous element in the Bur- mese amber fauna. Its zoogeographic relationships are uncertain. In the Early Cretaceous the forms related to Burmacypha were prob- ably distributed very widely if Edgariekia Jell & Duncan from the Lower Cretaceous of Australia is indeed synonymous with Siberian Undacypha as claimed by Ansorge (1996). As stated above Burma- cypha seems to be not closely related to Siberian Jantardachus, the only Late Cretaceous genus known. ACKNOWLEDGEMENTS. [am deeply thankful to Mr. A.J. Ross (NHM) and Dr. A.P. Rasnitsyn (Paleontological Institute, Moscow) for the opportunity to study this extremely interesting fossil. I would also like to thank Mr Peter York (NHM) for the photograph of Fig. 1. REFERENCES Ansorge, J. 1996. Insekten aus dem oberen Lias von Grimmen (Vorpommern, Norddeutschland). Neue Paldontologische Abhandungen, 2: 1-132. Tillyard, R.J. 1921. Two new fossil insect wings in the collection of Mr. John Mitchell from the Upper Permian of Newcastle, N.S. Wales, belonging to the order Hemiptera. Proceedings of the Linnean Society of New South Wales, 46: 413-422. 1935. Upper Permian insects of New South Wales. III. The order Copeognatha. Proceedings of the Linnean Society of New South Wales, 60: 265-279. Vishniakova, V.N. 1981. Novye paleozoyskie i mezozoyskie lophioneuridy (Thripida, Lophioneuridae) [New Palaeozoic and Mesozoic lophioneurids (Thripida, Lophioneuridae). Jn, Vishniakova, V.N., Dlussky, G.M. & Pritykina, L.N. Novye iskopaemye nasekomye s territorii SSSR. Trudy Paleontologicheskogo Instituta, 183: 43-63 (in Russian). Zherikhin, V.V. 1980. Otryad Thripida [The order Thripida]. Jn, Rohdendorf, B.B. & Rasnitsyn, A.P. (editors). Istoricheskoe razvitie klassa nasekomykh. Trudy Paleontologicheskogo Instituta, 175: 69-72 (in Russian). - eo - rf ce oe ae v ; “ab wehasleczncsnaor 2 i Nonsicte * ginecibs wer ofct ie ode roma G8 i: 7 ae ee ee wards ma Spent ams io Gaequadey saat: ign Yor ow —- plane : wy <_he = 5 i. aed hell" acted 7 i : ; ee : a ay iad a) i ‘ish ide qhgr@ ema | _ etna ra +) ae amine ara ome, yo ati ati wT She nyt -" wt whined evicig’t er | Ax Seen bite rohit 9 ati ted ee) | ae cull ausiee, bbe i | Tie ) qpénmpemgnt, 74 Natias Da Viek = PSolias Gent! | sal a ~— , . so aero of wT (NA iit x Ok: stubiande wien bvisguy ot ren Ses oe NS yl ae == Hakcincitie’ mid att Imre ioe 6 t= Me , = = 7 = - a sagan ; — ie Figen 4 ream a Appts ne cre aries mers : . i ag ea ey AVIVFARSTIA is a eet ee ————————— samnidonee nals at ee ee ee mrt re - r 7 ae - - é = a 7 . o A nw & . i * ’ < g - 4 : n se - we * a ab? 5 & 7 ‘ - —_ x aie me ry rn ty 73 : i " : e £ 5 7 ci e Fz -_ Es ay rage = 2 = - a, - a . : - 7 or { > - x ~~ = j ; a D ca , = 7 i 2 : x , i vo ’ . c 7 = { i i ’ iz i i \ \ = eh j ib s > ‘f Bull. nat. Hist. Mus. Lond. (Geol.) 56(1): 43-45 Issued 29 June 2000 Burmapsilocephala cockerelli, a new genus and species of Asiloidea (Diptera) from Burmese amber S.D. GAIMARI Smithsonian Institution, Washington, D.C. 20560 USA M.B. MOSTOVSKI Arthropod Labioratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya Str. 123, Moscow 117647, Russia SYNOPSIS. Psilocephala electrella Cockerell is figured, and another asiloid fly, Burmapsilocephala cockerelli gen. et sp. nov., is described from Burmese amber. The new genus is hypothesized to be close to the extant genus Apsilocephala. The phylogenetic position of Apsilocephala is discussed. INTRODUCTION Psilocephala electrella was described by Cockerell (1920) from Burmese amber based on an inclusion with incomplete wings and body. Through the courtesy of Mr A.J. Ross (Natural History Mu- seum, London) and Prof. A.P. Rasnitsyn (Paleontological Institute, Moscow) the collection of flies in Burmese amber stored in the Natural History Museum, including the holotype of this species (In.20148) (Fig. 1), was available for study. It is difficult to be certain that this species belongs to the genus Psilocephala, because character- istic features are not preserved. Another specimen (In.20167) (Fig. 4) belonging to the superfamily Asiloidea was discovered, with wing venation and leg structures resembling those of the holotype of Psilocephala electrella. However, it differs from the latter in having a slender body and more robust thoracic bristles, except for the dorsocentrals. On balance, it proved to be a representative of a new genus described here. SYSTEMATIC DESCRIPTIONS The following abbreviations are used: For thoracic macrosetae: ppn = postpronotal, np/ = notopleural, sa = supraalar, pa = postalar. For the positions of setal rows in tibiae: v = ventral, pv = posteroventral, av = anteroventral, d= dorsal, pd = posterodorsal, ad = anterodorsal. Order DIPTERA Superfamily ASILOIDEA Family APSILOCEPHALIDAE Genus BURMAPSILOCEPHALA nov. NAME. The name is derived from Burma, and the genus name Apsilocephala. TYPE SPECIES. Burmapsilocephala cockerelli nov. DIAGNOSIS. Scape, pedicel, and first flagellomere subequal in length. Antennal stylus 2.5 times longer than antennal segments 1-3 combined, and slightly thinner than in Apsilocephala. Proboscis © The Natural History Museum, 2000 slightly longer and does not appear to be as fleshy as in Apsilocephala. Postpronotal lobe has a single, strong seta. Mesonotum covered with hairs. One pair of strong scutellar setae present. The foretibial macrosetae have the formula 5y, Spv, Sav, Spd, 5ad. The hindtibia has fewer than 10pd macrosetae. REMARKS. Burmapsilocephala is taxonomically close to the ex- tant genus Apsilocephala Krober, 1914, with only minor morphological differences. In Apsilocephala the pedicel is about half the length of the scape or first flagellomere, which are subequal. The antennal stylus is 1.0 to 1.5 times longer than antennal segments 1-3. The postpronotal lobe has long hairs but no macrosetae. Two pairs of scutellar setae are usually present, although one pair can be stronger than the other. The foretibia of Apsilocephala has only 2— 3pd and 0—1ad, and hindtibia has more than 10pd. Burmapsilocephala differs from Apsilocephala and therevids, in possessing a long stylus and vein R5 ends near the wing tip. Remarkable is the absence of robust dorsocentrals in Burmapsilocephala. Burmapsilocephala cockerelli sp. nov. Figs 2-4 NAME. The name is in the memory of T.D.A. Cockerell. MATERIAL AND LOCALITY. Holotype In.20167, from Burmese amber, Hukawng Valley, Myanmar (Burma); probably Upper Creta- ceous (see Zherikhin and Ross, this volume). DESCRIPTION. Female, the tip of abdomen not discernible. Body and legs slender. Head: Frons narrow, bare. Ocellar triangle nearly flattened. Scape (1), pedicel (2), and first flagellomere (3) subequal in length. Scape and pedicel with short hairs apically. First flagellomere slightly tapered to apex. Stylus apical, 2.5 times longer than segments 1-3 combined. Thorax (Fig. 2): Postpronotal lobe relatively large. Mesonotum highly arched, covered with short hairs. Postalar tubercles distinct. Macrosetal pattern: lppn, 2npl, 1sa, \pa. Scutellum small, trapezi- form; with 1 pair strong macrosetae; covered with long hairs. Pteropleuron bare. Metathorax relatively large. Halter pale with darker knob. Fascicle of long delicate hairs anterior to halter. Wing (Fig. 3) with cell m3 closed and stalked; costal vein termi- nates at wing tip. Legs: Coxae with long hairs. Hind femur without macrosetae. Foretibia with apical ringlet of five macrosetae, and rows of +t Fig. 1 Psilocephala electrella Cockerell, holotype In.20148, Burmese amber. Width of wing 1.5mm macrosetae with formula Sy, Sad, Spd, 5pv, Sav. Midtibia with apical ringlet of 4-5 macrosetae, and rows of macrosetae with formula 3d, Aad, 2p, 3pv, 2pd. Hindtibia with asymmetrical, apical ringlet of macrosetae, and rows of strong and weak macrosetae with formula 3ad, 7 (28)pd, \pv (in distal third), 6av. Metatarsus with rows of strong hairs ventrally (at least in basal half), and with lateral setae, and with 4 (?5) strong apical setae. Tarsal segments 2—5 haired and with apical setae. Empodium bristle-like. Pulvilli and tarsal claws small. Abdomen (Fig. 4): Tergite 1 three times shorter than tergite 2; with lateral fascicles of long hairs. Tergites 3-6 subequal in length. Figs 2,3. Burmapsilocephala cockerelli gen. et sp. nov., holotype In.20167, Burmese amber. 2, head and thorax; 3, wing venation; scale bar 1mm. S.D. GAIMARI AND M.B. MOSTOVSKI Fig.4 Burmapsilocephala cockerelli gen. et sp. nov., holotype In.20167, Burmese amber. Length 5.3mm. Tergite 2 longer than others; tergite 7 slightly shorter than others. Tergites dark with pale, narrow band along lateral and posterior margins. Tergites 1-5 covered sparsely with short hairs; 6-8 covered with short hairs. Tergite 9 smaller and slender than the others, with long and short seta-like hairs. MEASUREMENTS. Body length (without antennae) = 5.3mm, wing length = 2.7mm, wing width = 1.2mm. DISCUSSION. Apsilocephala was originally described as a member of the Therevidae (Kréber, 1914), but this certainly represents an enigmatic genus. Apsilocephala was considered separate from Therevidae by Irwin (1976) and Irwin and Lyneborg (1981), but the position of the genus was not considered further. Apsilocephala was given family status, and included the genera Clesthentia White, 1914 and Clesthentiella Nagatomi, Saigusa, Nagatomi et Lyneborg, 1991 (Nagatomi etal, 1991a). However, the monophyly of the group, or at least its ranking at family-level, was questioned due to absence of definitive synapomorphies (Sinclair et al, 1994). The phylogenetic comments herein do not address the monophyly or internal classi- fication of the family Apsilocephalidae, but only its position within the Diptera. Nagatomi et al (199la, c) hypothesized a close affinity of Apsilocephalidae with the Middle to Late Jurassic family Rhagionempididae based on superficial similarities, but noted that the most important characteristics (e.g. male and female genitalia, condition of empodium) are not known in the rhagionempidids. Nevertheless, Nagatomi and Yang (1998) did synonymize Apsilocephalidae under fossil Rhagionempididae. It seems to be precocious now. Further, Nagatomi et al (199 1a, b, c) and Nagatomi (1992, 1996) hypothesized a sister-group relationship with Empidoidea or with Eremoneura, based again on the presence of surstyli in the male genitalia. Griffiths (1994, 1996), Sinclair et al (1994), Cumming ef al (1995), and Zatwarnicki (1996) rejected this hypothesis based on the observation that the ‘surstyli’ of Apsilocephalidae were erroneously considered homologous to the structures of Empidoidea and Eremoneura. Sinclair et al (1994) and Cumming ef al (1995) retained the genus Apsilocephala within the concept of Therevidae, but as incertae sedis (without further com- ment on Clesthentia or Clesthentiella). Additional consideration by NEW ASILOIDEA FROM BURMESE AMBER Yeates (1994) also maintained this position of Apsilocephala very near to Therevidae, and current morphological (M.E. Irwin, pers. comm.) and molecular (L.L. Yang and B.M. Wiegmann, pers. comm.) studies also suggest that Apsilocephala is very near or possibly within Therevidae. Therefore, the concept of Apsilocephala or the Apsilocephalidae as being near the stem of Eremoneura could be rejected in favour of its original position within or near the Therevidae. However, we keep the families Apsilocephalidae and Therevidae apart and avoid the formal synonymy until the position of fossil Rhagionempididae is clarified. The monophyly of this family as defined by Nagatomi et al (1991la) needs further consideration and definition with synapomorphies, but Burmapsilocephala is certainly closely aligned with Apsilocephala. Following are characteristics of Burmapsilo- cephala shared with Apsilocephala that support this hypothesized affinity and preclude the fossil genus from being placed elsewhere in the Asiloidea or Empidoidea. The elongated, apical, antennal stylus is similar to that in Apsilocephala and unique among therevids, as is the shape of the first flagellomere. The empodium is setiform, not pulvilliform. The scutum has macrosetae (including ppn, npl, and sa), aS does the scutellum. The pteropleuron is bare. The wing venation is identical to that in Apsilocephala, with the costal vein and RS5 terminating at the wing tip. The halter is pale, with a darker knob. Each tibia has an apical ringlet of macrosetae. The lengths of abdominal tergites relative to each other are consistent with that found in Apsilocephala. The extant species Apsilocephala longistyla Krober, 1914, has a distribution restricted to the southwestern U.S.A. (Arizona, Califor- nia, New Mexico, and Utah), and Mexico. Additionally, an undescribed fossil species of Apsilocephala is known from Eocene/ Oligocene Baltic amber (S.D. Gaimari pers. observ.). The other members of Apsilocephalidae are currently known from Tasmania and New Zealand. Information on the biology and immature stages of Apsilocephala is largely unknown, although adults of Apsilocephala longistyla have been collected in emergence traps placed over rock crevices filled with loose, friable soil (W.J. Hanson pers. comm.). Nagatomi et al’s (1991a,c) proposed affinity of Apsilocephalidae with Rhagionempididae suggests a Jurassic origin, yet the earliest known fossils belonging to Apsilocephalidae (herein) are probably Upper Cretaceous in age. This certainly does not exclude Apsilocephalidae from being present in the Jurassic. Therevidae were certainly extant during the Jurassic, with evidence from earliest known fossil therevid, Rhagiophryne bianalis Rohdendorf, 1964, found in number in the Middle—Upper Jurassic deposits in southern Kazakhastan (Mostovski 1998). Asilids and presumable scenopinids and hilarimorphids are also recorded there. 45 ACKNOWLEDGEMENTS. The authors would like to thank Mr A.J. Ross for access to the collection of Diptera in Burmese amber at the Natural History Museum, London, and Prof. A.P. Rasnitsyn for encouraging this work, Drs N.L. Evenhuis, M.E. Irwin, and anonymous referee for reviewing the manu- script, K.C. Holston for comments on an earlier version, and Drs D.K. Yeates and B.M. Wiegmann for comments on the relationships of Apsilocephala. Many thanks to Mr Peter York (NHM) for taking photographs. REFERENCES Cockerell T.D.A. 1920. A therevid fly in Burmese amber. The Entomologist, 53: 169- 70. Cumming J.M., Sinclair B.J. & Wood D.M. 1995. Homology and phylogenetic implications of male genitalia in Diptera — Eremoneura. Entomologica Scandinavica, 26: 120-151. Griffiths G.C.D. 1994. Relationships among the major subgroups of Brachycera (Diptera): a critical review. Canadian Entomologist, 126: 861-80. — 1996. Review of papers on the male genitalia of Diptera by D.M. Wood and associates. Studia dipterologica, 3 (1): 107-123. Irwin M.E. 1976. Morphology of the terminalia and known ovipositing behavior of female Therevidae (Diptera: Asiloidea), with an account of correlated adaptations and comments on phylogenetic relationships. Annals of the Natal Museum, 22: 913- 35). & Lyneborg L. 1981. The genera of Nearctic Therevidae. I/linois Natural History Survey Bulletin, (1980) 32: 188-277. Krober O. 1914. Beitrage zur Kenntnis der Thereviden und Omphraliden. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, (1913) 31: 29-74. Mostovski M. B. 1998. The early stages of the evolution of brachyceran flies (Diptera Brachycera). Unpublished PhD dissertation. Paleontological Institute of the Russian Academy of Sciences, Moscow. 219 pp. (in Russian). Nagatomi A. 1992. Notes on the phylogeny of various taxa of the orthorrhaphous Brachycera (Insecta: Diptera). Zoological Science, 9: 843-57. 1996. An essay on phylogeny of the orthorrhaphous Brachycera (Diptera). Entomologists’ Monthly Magazine, 132: 95-148. , Saigusa T., Nagatomi H. & Lyneborg L. 1991a. Apsilocephalidae, a new family of the orthorrhaphous Brachycera (Insecta, Diptera). Zoological Science, 8: 579-91. . 7 & 1991b. The genitalia of the Apsilocephalidae (Diptera). Japanese Journal of Entomology, 59: 409-23. 7 : & 1991c. The systematic position of the Apsilocephalidae, Rhagionempididae, Protempididae, Hilarimorphidae, Vermileonidae and some gen- era of Bombyliidae (Insecta, Diptera). Zoological Science, 8: 593-607. & Yang, D.. 1998. A review of extinct Mesozoic genera and families of Brachycera (Insecta, Diptera, Orthorrhapha). Entomologists’ Monthly Magazine, 134: 95-192. Sinclair B.J., Cumming J.M. & Wood D.M. 1994 [1993]. Homology and phylogenetic implications of male genitalia in Diptera — Lower Brachycera. Entomologica Scandinavica, 24(4): 407-32. Yeates D.K. 1994. The cladistics and classification of the Bombyliidae (Diptera: Asiloidea). Bulletin of the American Museum of Natural History, 219: 1-191. Zatwarnicki T. 1996. A new reconstruction of the origin of eremoneuran hypopygium and its implications for classification (Insecta: Diptera). Genus, 7: 103-175. Therikhin V.V. & Ross A.J. The history, geology and age of Burmese amber (Burmite). Bulletin of the Natural History Museum, Geology, 56: 3-10. gd) pad . is deena Te sip Coe ie : Geum seasioe “exynce Te ite bien Fee ada rv teiore % Ant a a mi we eee ee <= nies pewet.t 1 inte Gti rant, ~ Bie ae Cforeeet eps sed & Wye a AT a 1 =a = ite “ire Woe tees i i. sain 12 veer Kh by fot Fd : ° ad : ? ha Pilea ba, ‘ )iy_ <= oe Sep a istic a he ary ap a iy ; eS be Ed ' ute «ar =e . 7 i ‘al > we ’ ae _— <<, 's , ste : ae — i i ns ¥ | vis tie Siatirecc er Tit Bull. nat. Hist. Mus. Lond. (Geol.) 56(1): 47—52 Issued 29 June 2000 Phantom midges (Diptera: Chaoboridae) from Burmese amber E.D. LUKASHEVICH Paleontological Institute, Russian Academy of Sciences, Profsouznaya Str. 123, Moscow, 117647 Russia SYNOPSIS. Two new chaoborid species of the extinct genus Chaoburmus gen. nov. are described based on two males and one female from Burmese amber. Diagnostic features of the new genus are approximated eyes, short R3+4 and M1+2 forks, relatively short Sc and A veins, tibial spurs, tarsomere | longer than tarsomere 2, the fifth tarsomere in male simple, undilated, with small simple claws. INTRODUCTION There are many peculiarities of the Burmese amber insect fauna. Some groups (for example, Coleoptera) are numerous and even dominant though usually (in ambers from other localities) they are not so abundant (Rasnitsyn, 1996). The composition of the dipteran families in the Burmese amber assemblage (see Rasnitsyn & Ross, this volume) suggests an unusual ecological situation in the Burmese forest. In order of abundance, the leading families are Empididae (21%), Psychodidae (including Phlebotominae) (17%) and Chironomidae (16%). The dominant position of empidids is recorded otherwise only in the assemblage of Nizhnyaya Agapa locality (lower Upper Cretaceous of Taimyr, North Siberia) where they make up as much as 48% of all the dipterans. From the literature (Larsson, 1978; Zherikhin, 1978; Kulicka et al., 1985) Chironomidae are common in amber, whereas Psychodidae are rare. In the numerically studied collections of fossil resins Chironomidae are usually the most abun- dant: Nizhnyaya Agapa (Taimyr, Cretaceous) — 30%, Yantardakh (Taimyr, Cretaceous) — 71%, Starodubskoye (Sakhalin, Paleocene) — 77%, Baltic amber — 41% (the latter is calculated from the material collected directly at the Kaliningrad amber-mine, K.Y.Eskov, pers. comm.; other published percentages are biased for the reason ex- plained by Larsson (1978: 89): “The Copenhagen collection contains about 900 specimens, approximately 11% of the entire collection, and the museum has been offered at least ten times as many.’). In contrast, numbers of psychodids in the assemblages other than that of Burmese amber is invariably low. Only several psychodid inclu- sions are found in both Cretaceous fossil resin localities from Taimyr, as well as in the Paleocene Sakhalin amber. In Baltic amber the psychodids represent 2.5% (K.Y. Eskov, pers.comm.). Larsson’s figure for the Copenhagen collection is 4%; in the Baltic amber collection of the Museum of the Earth in Warsaw only Nematocera were counted, not all Diptera, but the numbers are significant: Chironomidae — 2592 specimens, Psychodidae — 262. One of the Burmese amber rarities is the find of three specimens of Chaoboridae, or phantom midges. A damaged chaoborid male from Burmese amber was first mentioned many years ago and determined as Chaoborus sp. by Edwards (1923). He wrote: ‘In size and appearance it differs little from the small species at present existing in India’ (p.152). Since then this specimen was mentioned in some reviews (Spahr, 1985; Poinar, 1992). Through the courtesy of Mr Andrew Ross (NHM, London) and Dr. A.P. Rasnitsyn (PIN RAS, Moscow), I had an opportunity to re-examine this inclusion. It does not appear to be amember of Chaoborus Lichtenstein, 1800, because of the presence of approximated eyes, tibial spurs and a comparatively © The Natural History Museum, 2000 short anal vein. The similar, though not conspecific, male and female have also been found in one piece of Burmese amber, showing additional peculiarities in wing venation. All of them are placed ina new genus described below and are housed in the Department of Palaeontology of the Natural History Museum (London). Chaoborids are very rare in amber and usually family participa- tion doesn’t exceed 0.1% of Nematocera (Kulicka et al., 1985). Only ten specimens of Chaoboridae are known from Baltic amber (Upper Eocene—Lower Oligocene), though thousands of dipterans are described from it or determined at least to the family level. Three of these chaoborid fossils were described at the beginning of our century as extinct species of the recent genera Chaoborus (known then as Corethra) and Mochlonyx Loew, 1844 (Meunier, 1902, 1904). Unfortunately, a holotype female of C. ciliata Meunier, 1904 has never been re-examined (and seems to be lost), but two males of M. sepultus Meunier, 1902 were redrawn and redescribed by Hennig (1966). Three others are deposited at the Zoological Muzeum of Copenhagen: one of the specimens was identified, with some reser- vations, as M. sepultus by Hennig, but two remaining specimens from this collection were not determined because of their poor preservation (Larsson, 1978). Three additional specimens are recorded (but not described or determinated even to the genera) from the Museum of the Earth in Warsaw and Gdansk (Kulicka et al., 1985). An additional, hitherto unrecorded specimen from the collec- tion of the Department of Palaeontology of The Natural History Museum (London) has been shown to me by A.Ross; this well- preserved specimen (register number II.18) was determined by meas a member of Chaoborus. Of course, new records are possible, but the general trend that they are quite rare seems to be clear. One more chaoborid, Trichia gracilis Hong, 1981, was described from Chinese Eocene amber (Hong, 1981). This generic name appears to be a junior homonym and was replaced by I[yaiyai (Evenhuis, 1994). Earlier Borkent (1993), in his world catalogue, presumed that this specimen does not belong to Chaoboridae at all: “The lack of a plumose male antenna, the peculiar wing venation and wing shape, the strikingly elongate legs and what appears to be a transverse suture on the scutum all suggest that the species does not belong within the family’, but due to absence of an alternative placement ‘the genus remains an enigmatic member of the Chaoboridae’ (Borkent, 1993: 6). 7: gracilis cannot be assigned to the family because of the above-mentioned characters combined with venation peculiarities which are seen on the published photo (distal shift of all furcations, and posterior M branch aligned to M stem). This specimen belongs to Tipulomorpha beyond doubt, but the proper family identification requires re-examination of the inclu- sion (densely plumose veins are known in Limoniidae, so it may belong to this family). 48 There are some other published records of chaoborid inclusions. Two specimens are mentioned in Nizhnyaya Agapa (Taimyr, Cenomanian, Upper Cretaceous) and one more in the Paleocene Sakhalin amber (Zherikhin, 1978); Evenhuis’ reference (1994) to Zherikhin & Sukatcheva (1973), concerning the fossil chaoborids found in the Cretaceous Siberian resins is erroneous and should be read as Zherikhin, 1978). After a re-examination the discussed specimen from Sakhalin appears to be one of the numerous chironomids, and one specimen, a poorly preserved male from Nizhnyaya Agapa (PIN N 3624/98) is a ceratopogonid. So the only chaoborid from Upper Cretaceous Taimyr resin (Nizhnyaya Agapa) is a female with the first tarsomere as long as the second, described as Taimyborus aequiarticulatus (Lukashevich, 1999). An additional, hitherto unrecorded female has been found in Lower Cretaceous Lebanese amber from D.Azar collection (D.Shcherbakov, pers.comm. ). Chaoboridae are repeatedly mentioned among other Diptera from Saxonian Upper Oligocene-—Lower Miocene amber (Barthel & Hetzer, 1982: with a reference to determination by Schumann; Schumann, 1984; Schumann & Wendt, 1989). It was a preliminary determination, and since that time neither further identification nor quantitative composition of the Bitterfield chaoborid assemblage have been published. Perhaps during detailed re-examination they may appear to belong to the Corethrellidae, the family that until recently was usually included with Chaoboridae. Corethrellidae are recorded from Bitterfield as well as from a variety of Cretaceous and Tertiary resins (however, not yet in Baltic amber): Corethrella cretacea from Lower Cretaceous Lebanese amber, C. prisca and C. miocaenica from Oligocene—Miocene Saxonian amber and C. nudistyla from Oligocene—Miocene Dominican amber, each species being described based on a single male (Borkent & Szadziewski, 1992; Szadziewski et al., 1994; Szadziewski, 1995). The above observations indicate that chaoborid inclusions in fossil resins are as rare as that of Corethrellidae. The case of impression fossils is strikingly opposite: not a single corethrellid impression is found thus far, while thousands of chaoborid impres- sions are collected in numerous Jurassic and Cretaceous localities of Europe and Asia. This contradiction may depend, at least in part, on the habitat preferences by the two midge groups. Extant immature Corethrella are most commonly found in small water bodies, for example, in water accumulated in leaf axils, epiphytic plants and tree hollows, the females of some species are blood-feeders on birds and tree frogs (McKeever, 1986). As a result, adult corethrellids are dendrophilous, in contrast to chaoborids which prefer the herbage for resting. Based on study of numerous samples of resins of differ- ent living conifers Zherikhin & Sukacheva (1989) infer that ‘terrestrial organisms connected with forest vegetation and especially with tree trunks are taphonomically preferred’ (p. 91). In other words, being dendrophilous, Corethrellidae have much higher chance than Chaoboridae of being trapped in resin. Hence, even being a rare group in the source biocenosis, they might even outcompete chaoborids as inclusions in fossil resins. SYSTEMATIC DESCRIPTIONS Family CHAOBORIDAE Edwards, 1912 Genus CHAOBURMUS nov. TYPE SPECIES. Chaoburmus breviusculus sp. nov; probably Late Cretaceous, Burmese amber. NAME. From genus Chaoborus and Burma (now Myanmar). E.D. LUKASHEVICH Table 1 Character states in the genera discussed (apomorphies are in bold, E = as in Eucorethra, C = as in Chaoborus). Character Eucorethra Chaoborus Chaoburmus tars 1/tars2 >1 >1 B—C male tars5 swollen undilated C male claw large, complex small, simple C tibial spurs present absent E eyes approximated well separated E clypeus long short C haltere pedicel with setae without setae C Sc long usually long short R3+4 fork >R3+4 stem >R3+4 stem M1+2 stem E A apex to Rs distal distal or proximal C A apex to m—cu proximal distal Polarity doubtful DIAGNOSIS. Imago. Small densely pubescent midges. Pedicel with- out setae. 13 flagellomeres, last two being the longest. Eyes reniform, approximated. Clypeus length shorter than the head capsule height. Wing transparent. Posterior wing margin setae long and dense. Sc short, ending somewhat distad of RS bifurcation (level with r—m). Appendix veins (RSa and Cua) absent. R3+4 and M1+2 fork short, shorter than their respective stems. Anal vein entering wing margin proximal to m—cu and RS origin level. Haltere without conspicuous setae. Gonocoxite elongate, with long setae; gonostyle bare, without apical seta. Tibiae with apical spurs. Tarsomere | longer than tarso- mere 2, last male tarsomere simple, undilated, claws small and simple. OTHER SPECIES. ?C. victimaartis sp. nov. from Burmese amber. REMARKS. In the ratio of the first to the second tarsomeres, Chaoburmus is similar only to two recent genera, Eucorethra Underwood, 1903 and Chaoborus (character matrix for these genera is given in Table 1; polarity of the characters, except venation, after Saether, 1992), and to all genera described from compression fossils (except those known only from wings: Rhaetomyia Rohdendorf, 1962 and Helokrenia Kalugina, 1985); from all extinct genera dis- tinct in the short anal vein, and from most of those for which the tarsi are known (Astrocorethra Kalugina, 1986, Baleiomyia Kalugina, 1993, Hypsocorethra Kalugina, 1985, Mesocorethra Kalugina, 1993, Praechaoborus Kalugina, 1985) also in their structure, being similar in the simple last tarsomere with small claws and presence of tibial spurs only to Chachotosha Lukashevich, 1996. The latter differs from Chaoburmus in larger size, the apical seta on gonostyle and standard chaoborid venation with long Sc, A, and the R3+4 and M1+2 forks (Lukashevich, 1996). Tarsi are poorly known in Chironomaptera Ping, 1928, presumably collective genus wide- spread in the late Mesozoic of Asia; according to the photographs and drawings of Zhang (1990), in the type species C. gregaria (Grabau, 1923) and in C. vesca Kalugina, 1980 the last tarsomere is simple with small simple claws, and one apical spur is recorded on the fore tibia in the latter species; Chironomaptera is nevertheless distinct from the new genus in the short, rounded or oval gonocoxite and the veins and forks not shortened. Among extant genera, the anal vein is similarly short (though ending far distad of RS origin) and recorded only for Eucorethra, Chaoburmus being distinct from it in the structure of claws and last tarsomere and in the absence of dense haltere pubescence. In these latter features Chaoburmus resembles Chaoborus, but differs from it, besides the anal vein, in the approximated eyes and the presence of tibial spurs (Eucorethra possessing a single spur on each tibia). Neither the short R3+4 and M1+2 forks nor the short Sc are typical PHANTOM MIDGES FROM BURMESE AMBER for Chaoborus and Eucorethra (Saether 1970, 1976; the short Sc terminating at rm level was independently acquired by the only species, recent C. brevisector Edwards, 1930). Additionally, Chaoburmus differs from Eucorethra in the smaller size and unspotted wings. These characters are of specific and not generic level in Chaoborus, and the same could be true of Eucorethra if more than one species were known. As for the extinct genera, a short M1+2 fork often occurred in the Mesozoic, so with some reservation this character can be considered a plesiomorphy. A short R3+4 fork (shorter than the respective stem) is recorded only in Baleiomyia discussoria Kalugina, 1993 (Unda and Daya localities, Transbaikalia, Upper Jurassic-Lower Creta- ceous) and Helokrenia nana Kalugina, 1985 (Kubekovo locality, Siberia, Middle Jurassic; one more, undescribed fossil tentatively 1b Figs la, 1b =Chaoburmus breviusculus Lukashevich, sp.nov., holotype NHM In.20168, male in Burmese amber; views from opposite sides. 49 Fig. 2 Chaoburmus breviusculus Lukashevich, sp.nov., paratype NHM In.20168(1), female in Burmese amber. determined as Helokrenia sp. was found in the Purbeck Beds, England, Lower Cretaceous (Berriasian); Ed. Jarzembowski, pers. comm. ). Besides the short, broad R3+4 and M1+2 forks, Helokrenia and Chaoburmus are similar in their small size and short Sc (ending at r—m level), being distinct in the anal vein length. Though the anal vein wasn’t figured in the description of Helokrenia nana (Kalugina & Kovalev, 1985: 80, fig. 38), it can be seen in the holotype as a long vein ending distal to m—cu as usual. The English specimen possesses a long anal vein as well. The short Sc is characteristic of Jurassic Dixidae (another family of the same superfamily, known since the Jurassic) and the oldest chaoborid, Rhaetomyia and therefore con- sidered a plesiomorphy. Preimaginal stages remain unknown. The combination of other characters is too unusual to assign Chaoburmus to either Eucore- thrinae or Chaoborinae at present state of our knowledge (the balance of possible synapomorphies is in favour of Chaoborinae, see Table 1s Figs 14 NAME. From Latin breviusculus — rather short, alluding to the length of first tarsomere compared to the second. Chaoburmus breviusculus sp. nov. HOLOTYPE. NHM In.20168, inclusion of a well preserved male; Burmese amber; probably, Late Cretaceous (see Zherikhin & Ross, this volume). PARATYPE. NHM In.20168(1)—a well preserved female originat- ing from the same piece of amber, but now separate. DESCRIPTION. Small densely pubescent midges with pale legs. Wing unspotted, three times as long as wide, veins with short macrotrichia. R1 slightly displaced forwards, terminating proximad or level with R3+4 furcation. R3+4 fork 2 (female) —2.5 (male) times shorter than R3+4 stem. Vein r—m aligned with bas M3+4. M1+2 fork 1.5 (female) —2.5 (male) times shorter than M1+2 stem. Anal lobe somewhat reduced. Gonocoxite approximatelly 3 times as long as wide, gonostyle about one-third shorter than gonocoxite. At least fore and middle tibia with one(?) apical spur each. First tarsomere 1.1-1.25 times longer than second one. 50 E.D. LUKASHEVICH Fig.3 Chaoburmus breviusculus Lukashevich, sp.nov., holotype NHM In.20168, male in Burmese amber: total view. Scale = 0.5 mm. Figs 4a-e NHM In.20168, male in Burmese amber; a, genitalia; b, ?fore tarsus (reconstructed, four distal tarsomeres lie separately). c—e, paratype NHM In.20168(1), female in Burmese amber; ¢, wing; d, antenna; e, pair of fore legs; scale bar= 0.5 mm. Chaoburmus breviusculus Lukashevich, sp.nov. a, b, holotype MEASUREMENTS. Male. Antenna length = 0.83 mm, thorax length = 0.63 mm, abdomen length (with genitalia) = 1.55 mm, wing length = 1.0 mm, wing width = 0.34 mm, fore femur = 0.64 mm, fore tibia = 0.62 mm, fore tarsus = 0.2/? 0.16/0.13/0.1/0.07 mm, middle femur = 0.53 mm, middle tibia = 0.53, middle first tarsomere = 0.2, hind femur = 0.65 mm, hind tibia = 0.61 mm, first hind tarsomere = 0.25 mm. Female. Antenna length = no less than 0.5 mm, thorax length = 0.6 mm, abdomen length = 1.25 mm, wing length = 1.1 mm, wing width = 0.35 mm, fore femur = 0.62 mm, fore tibia = 0.6 mm, fore tarsus = 0.2/0.17/0.12/0.1/0.1 mm, middle femur = 0.5 mm, middle tarsus = 0.22/0.17/0.15/0.1/0.1 mm, hind femur = 0.67 mm, hind tibia = 0.62 mm, first hind tarsomere = 0.32 mm. REMARKS. In the ratio of the first to the second tarsomeres and small size C. breviusculus is similar Taimyborus aequiarticulatus Lukashevich, 1999, from Upper Cretaceous Taimyr resin (Nizhnyaya Agapa locality), distinct in venation peculiarities (in the latter Sc, A, R3+4 and M1+2 forks longer), shorter wing macrotrichia, longer clypeus and the presence of tibial spurs. The small size, approximated eyes, R1 displaced forward and reduced anal lobe are the characters of Corethrella Coquillett, 1902, now separated into the family Corethrellidae mainly on account of the larval peculiarities. C.breviusculus differs from this group in the antennal pedicel without setae, two last flagellomeres longest and the tibial spur present on the middle leg. ? Chaoburmus victimaartis sp. nov. Figs 5, 6 NAME. From Latin victima and art — an art victim, alluding to the male posterior part polished away when making a bead. HOLOTYPE. NHM In.20157, inclusion of a well preserved male without distal abdomen and wing parts, legs partly damaged; Bur- mese amber; probably Late Cretaceous. DESCRIPTION. Male. Head broad, pubescent; eyes separated witha little more than greatest pedicel width. Wing unspotted, veins with macrotrichia especially dense on C and RS, forming sparse rows on Sc, M and A. Vein m—cu twice longer than bas M3+4, the latter PHANTOM MIDGES FROM BURMESE AMBER 51 with a pair of apical spurs; only one spur is visible on the fore tibia. First tarsomere 1.4—1.5 times longer than second one. MEASUREMENTS. ‘Total length of inclusion = 2.1 mm; antenna length = 1.2 mm (two last flagellomeres combined = 0.4 mm), pedicel diameter = 0.125 mm, head width = 0.6 mm, width between eyes = 0.16 mm; thorax length = 0.6 mm, thorax width = 0.35 mm, wing width (on level of anal vein apex) = 0.57 mm, fore femur = 0.9 mm, fore tibia =no less than 0.5 mm, fore tarsus = 0.35/0.25/0.2/0.2/ 0.15 mm, middle femur = no less than 0.8 mm, middle tibia = 0.8 mm, middle tarsus = 0.45/0.3/0.2/0.125/0.125 mm, hind tibia = 0.9 mm, three proximal hind tarsomeres = 0.55/0.37/0.3 mm. REMARKS. Distinct from C. breviusculus in the larger size, wider wings, all leg joints dark, ratio of the first tarsomere to the second one, shape of tergites and, possibly, in the number of tibial spurs. Because of the last feature some doubts remain about the generic affinity of this species. ACKNOWLEDGEMENTS. [| am indebted to Mr A.J. Ross (Natural History Museum, London) and Dr A.P. Rasnitsyn (Paleontological Institute, Russian Academy Sciences, Moscow) for a possibility to work with Burmese amber; Fig.5 ?Chaoburmus victimaartis Lukashevich, sp.nov., holotype NHM In.20157, male in Burmese amber. aligned with r—m. Haltere without conspicuous setae: pedicel trans- Dr V.V. Zherikhin for help in bibliographic search and Dr D.E. Shcherbakov lucent, bare; capitulum entirely uniformly dark (presence of very (both Paleontological Institute) for discussions; Dr R.Szadziewski (Univer- short pile cannot be excluded). Parascutellar (postalar) setae present. sity, Gdansk) for critical reading of the manuscript; and Peter York (NHM) for Tergites are subquadrate. All leg joints dark. Middle and hind legs photography. oe A y SS, GF a : \\ Gay Sa, ie SWZ —-—" | NSN = VF Zz bs \\ eX | 7 ke ACS AYR A A KSI SD: % tN OI i. iS ij Me Wh KL THK "iii, oni N\ til a ps —_x ARCHAIC NEW GENUS OF EVANIIDAE 25) Fig.3 Mesevania swinhoei new species, holotype (Scales: 0.5 mm) In.20192, Burmese amber. a, Head, antenna and foreleg; b, Hind tibia, inner face; c, Forewing veins and cells. Cell names. BC: basal cell, DC: discoidal cell, MC: marginal cell, SDC: subdiscoidal cell, SMC: submarginal cell. Rasnitsyn, from the Lower Cretaceous of Mongolia (Rasnitsyn, 1991), is possibly similar to Mesevania in this respect. Although the only known fossil of Praevania is incomplete, it can be distinguished easily from Mesevania in having a long 3rd submarginal cell, forewing vein 2r-rs being closer to vein 2rs-m (Fig. 4b), and in having a much longer ovipositor. Polymerous antenna can also be seen in the Praeaulacidae. The shape of the scape of Mesevania is rather similar to that of aulacids, though much shorter. Mesevania can be differen- tiated from the Praeaulacidae by the short pronotum and by the insertion of the petiole closer to the metanotum. Cretevaniidae differ from Evaniidae (including Mesevania) by the petiole being ex- panded subapically, and by the very different forewing venation with a narrow marginal cell and straight vein Rs which is more or less in line with 1rs-m. Unfortunately, a prominent synapomorphy known for the Evaniidae, the presence of the jugal lobe at the base of both fore- and hindwing, cannot be seen in the specimen described here because of the orientation of the inclusion. However, the massive mesosoma, lack of a conspicuous ‘neck’, presence of a long and slender petiole, high attachment of metasoma to propodeum, small metasoma and relatively short ovipositor, together with its general habitus, best relate this new genus to the Evaniidae. Mesevania swinhoei Basibuyuk & Rasnitsyn, sp. nov Figs 1-3 NAME. _ After R.C.J. Swinhoe, who presented the holotype to The Natural History Museum in 1920. TYPE MATERIAL. Holotype In.20192, Department of Palaeontol- ogy, NHM, London. LOCALITY AND HORIZON. Burmese amber, probably Upper Creta- ceous, Hukawng Valley, Myanmar (Burma) (for details, see Zherikhin & Ross, this volume). 56 DESCRIPTION. Female. Antenna with at least 26 segments; scape stout and short, as long as wide; pedicel as long as scape, much narrower; | st flagellomere longest; first 7 flagellomeres narrow and long, the remainder thickened and short (Fig. 3a). Head with a longitudinal furrow below and another above the toruli, the lower furrow strongly developed and densely pubescent; interantennal carina present; mandible tridentate (Fig. 3a). Forewing 2.4 mm long. Hindwing with 4—5 distal hamuli. Spur formula 1: 2: 2. Fore tibial spur modified; bifurcate at apex with outer tooth long, strongly enlarged at base; fore basitarsus slender (Fig 3a). Hind tibia with a transverse row of short spines apically; spurs short, 0.2 x length of basitarsus. Claws bifurcate (Fig. 3b). DISCUSSION Most of the known fossil evaniids are from relatively more recent geological periods (Burmeister, 1831; Brischke, 1886; Brues, 1933; Poinar, 1992), and are included in extant genera. However, three Mesozoic genera have recently been described and attributed to the H.H. BASIBUYUK ET AL. Evaniidae. Of these, Praevania from the middle Lower Cretaceous of Mongolia (Rasnitsyn, 1991), is putatively the oldest genus in the family but is of uncertain taxonomic placement owing to its poor preservation. The other two are found in New Jersey amber of lower Upper Cretaceous (Turonian) age in eastern USA (Basibuyuk et al., in press). Mesevania is probably also of Upper Cretaceous age and therefore constitutes the fourth known Mesozoic member of this family. The new genus occupies a unique place in the Evaniidae. All other described genera may fall into one group on the basis of the simple hind tibia, reduced forewing venation, elbowed antenna with long scape, and forewing with Rs more-or-less anteriorly curved and with relatively short marginal cell. The new genus has none of these putative synapomorphies but does possess some other synapomorphies of the Evaniidae, viz. the massive mesosoma, short metasoma, absence of a ‘neck’ and presence of a conspicuous petiole. Several other characters displayed by the new genus appear to be apomorphic within the Evanioidea. The possession of a medially short pronotum and reduced hindwing venation are synapomorphies Fig. 4 Forewing venation in selected taxa of Evanioidea. a, Mesevania swinhoei new species (Evaniidae); b, Praevania sculpturata Rasnitsyn, illustration constructed from both left and right wings (Evanioidea incertae cedis); ¢, New species from New Jersey amber (Evaniidae); d, Evania appendigaster (L.) (Evaniidae); e, Prosevania sp. (Evaniidae); f, Praeaulacinae gen. sp. (Paleontological Institute, Moscow no. 4270-1544: Lower Cretaceous) (Praeaulacidae); g, Baissa magna Rasnitsyn (Gasteruptiidae); h, Aulacus longiventris Kieffer (Aulacidae). ARCHAIC NEW GENUS OF EVANITDAE uniting the Cretevaniidae, Gasteruptiidae, Aulacidae and Evaniidae, but excluding the Praeaulacidae, while a short metasoma and the attachment of the metasoma to the propodeum being closer to the metanotum are characters shared by Evaniidae and Cretevaniidae (see above). The most conspicuous distinguishing characters of the new genus, viz. modified hind tibia, multi-segmented antenna with short scape and complete wing venation, are either autapomorphic or plesiomorphic and cannot be used to infer phylogenetic relation- ships (see Figs 3a—c). At the same time, these characters permit us to recognise three grades within the Evaniidae. The first grade is composed by Mesevania and Praevania (if the latter is assigned to the Evaniidae), which are the only taxa retaining 3rs-m crossvein and thus the closed 3rd submarginal cell in their forewing (Figs 4a—b). Likewise, the possession of simple, multi-segmented antennae together with a short scape are plesiomorphic character states for the Evanioidea. These conditions, therefore, suggest that this grade may represent the stem group of the Evaniidae. The transition of the forewing venation from relatively complete to less complete venation, that is, from 10 enclosed cell to 6—7 enclosed cells (Figs 4c—e), and from the polymerous, unmodified antenna to the elbowed, oligomerous type (with 13 or fewer seg- ments), indicates that there are synapomorphic characters uniting the remaining Evaniidae. At this level, the new genera from New Jersey differ from the remaining Cenozoic taxa in retaining several other plesiomorphic forewing venation characters, e.g. large 1st submar- ginal cell, wide basal cell, and more-or-less straight Rs&M, and thus form a grade between Mesevania and the remaining Evaniidae (Figs 4c cf. 4d). The Cenozoic evaniids constitute the third grade, delim- ited by further venational modifications such as the small Ist submarginal cell, large 1st discoidal cell, strong angle between veins Rs and M (Rs & M: Fig. 4a), and vein Rs conspicuously curved towards the anterior wing margin (Figs 4d—e). Relationships among Cenozoic genera are yet to be studied. In addition to Mesevania, conspicuously swollen hind tibiae also occur among the Evanioidea, in the extinct Cretevaniidae (Rasnitsyn, 1975). In the recent reanalysis of Rasnitsyn’s (1988) phylogeny of hymenopteran families, the Cretevanidae appear as the sister group to the Evaniidae (Ronquist et al., 1999). Thus, swollen hind tibiae may be plesiomorphic for the Cretevaniidae + Evaniidae and then subsequently lost in evaniids other than Mesevania. It is also inter- esting in this context to note that the hind tibia is quite expanded distally in male and female gasteruptiids, putatively the sister group of Cretevaniidae + Evaniidae. The function and internal structure of the swollen tibial apex in gasteruptiids, which differs somewhat from that of Mesevania in lacking a plate structure, is currently unknown and warrants detailed investigation. The hind tibial modification of Mesevania is also of particular interest because of its implications for host use by early Evaniidae (and Cretevaniidae). The only hosts of extant evaniids are cockroach oothecae (Brown, 1973), though other extant evanioids have very different biologies: aulacids being koinobiont endoparasitoids of wood-boring sawflies and beetles while gasteruptiids are cleptoparasitoids of solitary bees and masarine wasps (Haack & Wilkinson, 1987; Naumann & Cardale, 1987; Gauld & Bolton, 1988). The swollen hind tibiae of Mesevania are rather similar, at least superficially, to those of the fore leg in female Orussidae, and to a lesser extent, those of the hind legs of Stephanidae. Tibiae swollen in this way, at least in extant taxa, usually indicate the presence of large subgenual organs, which are important in detecting substrate- borne vibrations, and are typically most well developed in females of taxa attacking vibration-producing, xylophagous hosts. Modern evaniid hosts do not produce vibrations (as the oothecae are attacked 3) soon after being laid: Brown, 1973) and so extant evaniids have no need for enlarged subgenual organs. The swollen hind tibia of Mesevania, with its modified plate structure, suggests the presence of a subgenual organ, and it is therefore tempting to speculate that Mesevania may have attacked hosts other than cockroach oothecae. That the hosts of early evanioids were generally different from those of the three extant families is further suggested by the presence of strong transverse sculpture on the mesoscutum in some extinct taxa, e.g. Kotujisca kholbotensis (Kotujellitinae) (Rasnitsyn, 1991) (supposedly closely related to the putative ancestor of gasteruptiids — Gasteruptiinae sensu Rasnitsyn). This mesoscutal sculpture is typical of wasps parasitising endoxylous hosts and perhaps aids the wasp in egress from the pupation site (Eggleton, 1989; Quicke, 1997). 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Phylogeny of the Hymenoptera: A cladistic reanalysis of Rasnitsyn’s (1988) data. Zoologica Scripta, 28: 13-50. Townes, H. 1949. The Nearctic species of Evaniidae (Hymenoptera). Proceedings of the United States National Museum, 99: 525-539. Bull. nat. Hist. Mus. Lond. (Geol.) 56(1): 59-77 Issued 29 June 2000 Digger wasps (Hymenoptera, Sphecidae) in Burmese amber A. V. ANTROPOV Department of Entomology, Zoological Museum of the Moscow Lomonosov State University, Bol’shaya Nikitskaya Street 6, Moscow K-9, 103009, Russia SYNOPSIS. The following new taxa of the family Sphecidae are described from Upper Cretaceous Burmese amber: Burmastatus triangularis gen. et sp. nov. and Cirrosphex admirabilis gen. et sp. nov., are members of the new subfamilies Burmastatinae and Cirrosphecinae; Apodolichurus sphaerocephalus gen. et sp. nov., A. diaphanus sp. nov., Cretampulex gracilis gen. et sp. nov., and Mendampulex monilicularis gen. et sp. noy. are members of the new ampulicine tribes, Apodolichurini, Cretampulicini, and Mendampulicini, respectively; Prolemistus apiformis gen. et sp. nov. and Cretospilomena familiaris gen. et sp. nov., are placed in the tribe Pemphredonini. Trigonalys pervetus Cockerell, 1917, originally described in the family Trigonalyidae, is moved to the new sphecid genus Trigampulex gen. nov., incerta sedis. The principal differences of the evolution of the wing venation between the Upper Cretaceous and recent sphecid wasps are discussed. INTRODUCTION Until now the Apocrita were represented only by the families Aulacidae, Bethylidae, and Trigonalyidae in Upper Cretaceous am- ber from Myanmar (Cockerell, 1917a, 1917b, 1917c, 1917d, 1920) [though the Myanmar Trigonalyidae are now attributed to Sphecidae — see below]. During a brief study of fossil insects in the Department of Palaeontology of The Natural History Museum (London, UK), Prof. A.P.Rasnitsyn (Paleontological Institute of the Russian Acad- emy of Sciences, Moscow, Russia) identified some Burmese amber samples including, among many other arthropods, several sphecid wasps. This material is treated herein. It is necessary to note that the preparation quality, the transpar- ency of the samples and the position of the inclusions inside the samples are highly heterogeneous, and the necessity to maintain the initial state of the samples has considerably limited the oppor- tunities of thorough analysis of the inclusions because of inaccessibility to study some highly important features. Undoubt- edly, this has affected the completeness of the descriptions of the new taxa presented here. Though in recent years some authors have divided Sphecidae into several separate families on the basis of cladistic analysis, I cannot consider for the present that their preliminary results carry complete conviction. Here I follow the traditional view that Sphecidae are an indivisible family of hymenopterous insects, taking into account that it may be a paraphyletic grouping. I have tried to use the majority of the diagnostic characters and terms accordingly the generic revision of the recent sphecid wasps (Bohart & Menke, 1976) to preserve the continuity and comparativeness of the taxa. At the same time, the designation of the scutal lines is given according to the well rea- soned, from my point of view, proposals of Budrys (Budrys & Kazenas, 1992; Budrys, 1993). Specimens were examined under a Zeiss-stereomicroscope in a thick sugar solution rather than immersion oil in order to minimize the possibility of damage. © The Natural History Museum, 2000 SYSTEMATIC DESCRIPTIONS Superfamily APOIDEA Family SPHECIDAE I. New subfamilies of Sphecidae The family Sphecidae was divided into 11 subfamilies in the revi- sion of recent genera (Bohart & Menke, 1976). In recent years some authors have included the genera of the subfamily Larrinae (sensu Bohart & Menke, 1976) in the subfamily Crabroninae (e.g., Menke & Fernandez, 1996). Furthermore, following conception of the indivisible family Sphecidae, the genus Heterogyna Nagy, 1969, as well as the genera Dolichurus Latreille, 1809, Paradolichurus Williams, 1960, Aphelotoma Westwood, 1841, Austrotoma Riek, 1955, Trirogma Westwood, 1841, and Ampulex Jurine, 1807, which were included into separate families in a review of the families of Hymenoptera (Goulet & Huber, 1993), should be attributed to the subfamilies Heterogynainae and Ampulicinae. Fossil sphecid wasps have been already described in the following recent subfamilies: Ampulicinae (Nemkov, 1988), Sphecinae (Cockerell, 1906, 1907; Menke & Rasnitsyn, 1987), Pemphredoninae (Cockerell, 1906; Rohwer, 1909; Evans, 1969, 1973; Sorg, 1986; Budrys, 1993), Crabroninae (Cockerell, 1908, 1909, 1910; Rohwer, 1908; Antropov & Pulawski, 1989, 1993; Prentice & Poinar, 1993; Antropoy, 1995), Bembicinae (Cockerell, 1906, 1922; Rohwer, 1908; Pulawski & Rasnitsyn, 1980, Nemkov, 1990), and Philanthinae (Cockerell, 1906; Rohwer, 1909; Timon-David, 1944). The Lower Cretaceous genera Angarosphex Rasnitsyn, 1975, Baissodes Rasnitsyn, 1975, Oryctobaissodes Rasnitsyn, 1975, Trichobaissodes Rasnitsyn, 1975, and Bestiola Pulawski et A.Rasnitsyn, 1999 have been assigned to the fossil sphecid subfamily Angarosphecinae (Rasnitsyn, Pulawski & Martinez-Delclos, 1999). Furthermore, the subfamilial position has not been ascertained for Archisphex Evans, 1969, Taimyrisphex Evans, 1973, and Cretosphex A.Rasnitsyn, 1975 (Rasnitsyn, 1980). 60 Subfamily BURMASTATINAE nov. TYPE GENUS. Burmastatus gen. nov., here designated. DIAGNOSTIC FEATURES. 1. Inner eye orbits straight, parallel; ocelli simple. 2. Antennae attached to lower part of face; antennal sockets touching clypeal base; female antennae 12-segmented; scapes modified into large triangular plate with elongate ventral part. Clypeus transverse, short. 4. Mandibles without preapical teeth, with obtuse angle externoventrally and obtuse flat inner prominence basally; mandible sockets open; palpal formula 6—4; mouthparts short. 5. Pronotal collar short, flattened dorsally, with sharp transverse keel; pronotal lobes almost touching tegulae. 6. Scutum with adlateral (=parapsidal) lines, reaching at least 2/3 of scutal length; admedian lines not expressed. 7. Mesopleuron with incomplete straight episternal sulcus; omaulus, subomaulus, sternaulus, and acetabular carina absent, scrobal sulcus developed. Metapleuron completely developed. 9. Midtibia with two apical spurs; midcoxae subcontiguous; hindcoxae contiguous; precoxal lobes present; hindfemur apically simple; tarsal claws simple. 10. Propodeum short (shorter dorsally than posteriorly); propodeal dorsal enclosure U-shaped, concave, broader than long; propodeal lateral carinae present; intercoxal carinae absent. 11. Forewings with two submarginal and two discoidal cells; mar- ginal cell very short, with acute apex touching wing margin; recurrent vein I received by submarginal cell I, recurrent vein II received by submarginal cell II; forewing media diverging beyond cu-a. 12. Hindwing jugal lobe narrow, longer than 2/3 of submedial cell length; hindwing media diverging before cu-a; M distinctly longer than r-m and Rs together; Sc and A, , absent. 13. Abdomen without petiole; tergite I with lateral carinae; pygidial plate absent. Oo ge COMPARISONS. Thenew subfamily resembles Ampulicinae in hav- ing developed adlateral lines and two apical midtibial spurs, but differs by the simple tarsal claws and the mandibles with an obtuse externoventral angle. Combination of the last two features with two apical midtibial spurs is characteristic for the subfamilies Astatinae and Xenosphecinae, although these have no adlateral lines. The new subfamily also resembles representatives of Astatinae in having a long jugal lobe (it is short in Xenosphecinae), but differs in the venation of the fore- (particularly by the short marginal cell with its acute apex touching forewing margin) and hindwings (particularly of the medial cell), by the short propodeum, and by lacking the pygidial plate. REMARKS. The form of the pronotum (particularly of its collar, lobes, and lower angles) and of the hindtibial brush shows that the specimen belongs to the Sphecidae, but its main diagnostic features (presence of the adlateral lines, two apical midtibial spurs, obtuse externoventral angle of mandibles, and simple tarsal claws) do not relate it to any recent subfamily of Sphecidae. The considerably specialized fore- and hindwing venation differs from the known fossil genera Angarosphex, Archisphex, Cretosphex, and Taimyrisphex. Such distinct apomorphies as the scape modified into large, vertical, triangular plates, considerably diminished forewing marginal cell with its acute apex touching the wing margin, and the hindwing M being longer than r-m and Rs combined, place this specimen as the representative of a new subfamily. It is necessary to mention, that among recent sphecids the similar form of the hindwing A.V. ANTROPOV medial cell apex is typical for almost all Oxybelini and Crabronini, for some genera of Miscophini (in Saliostethus Brauns, 1897, and Saliostethoides Arnold, 1924 and, to a smaller degree, in Miscophus Jurine, 1807, Miscophoides Brauns, 1897, and Auchenophorus Turner, 1907), and also for some Pemphredonini (as the steps in Ammoplanus Giraud, 1869, and Ammoplanellus Gussakovskij, 1931, and distinctly in Pulverro Pate, 1937). The postfurcal cu-a is found only in the last genus. The reduction of the forewing venation in these groups, which is likely related in the last two tribes to the considerable diminution of the body size, has a very different nature (truncation and reduction or, on the contrary, elongating of the marginal cell; complete or partial reduction of the submarginal cell II, owing to reduction of its outer vein or petiolization; enlarging of the stigma). Consequently, this superficial similarity was, undoubt- edly, acquired independently in each group. Genus BURMASTATUS nov. ETYMOLOGY. The generic name is derived from the toponym Burma and the generic name Astata. The name is masculine. TYPE SPECIES. the only species. Burmastatus triangularis sp. nov., here designated; DIAGNOSIS. _ Inner eye orbits parallel; ocelli simple; antennal sock- ets touching clypeal base; female antennae 12-segmented; scapes modified into large triangular plate; clypeus transverse, short; man- dibles without preapical teeth, with obtuse angle externoventrally and obtuse flat inner prominence basally; mandible sockets open; palpal formula 6-4; mouth parts short. Pronotal collar short, flat- tened dorsally; pronotal lobes almost touching tegulae; scutum with adlateral and admedian lines; mesopleuron with incomplete straight episternal sulcus, not reaching their anterior margin; omaulus, subomaulus, sternaulus, and acetabular carina absent, scrobal sulcus developed; midtibia with two apical spurs; midcoxae subcontiguous; hindcoxae contiguous; precoxal lobes present, short; hindfemur apically simple; tarsal claws simple. Forewings with two submar- ginal and two discoidal cells; marginal cell very short, with acute apex touching wing margin; recurrent vein I received by submar- ginal cell I, recurrent vein II received by submarginal cell II; forewing media diverging beyond cu-a. Hindwing jugal lobe narrow, longer than 2/3 of submedial cell length; hindwing media diverging before cu-a; M distinctly longer than r-m and Rs together. Propodeum shorter dorsally than posteriorly; propodeal dorsal enclosure U- shaped, concave, broader than long; propodeal lateral carinae present; intercoxal carinae absent. Abdomen without petiole; tergite I with lateral carinae; pygidial plate absent. Figs 1-6 ETYMOLOGY. The species name is derived from the Latin mascu- line adjective triangularis, meaning ‘triangular’; the name is a reference to the shape of the female scape. Burmastatus triangularis sp. nov. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.20197; partly preserved female specimen (the following parts are absent: left antennomeres 4-8, the largest part of the left eye, left mandible basally, the largest part of the both left wings from the base, outer side of the left forefemur, left mid- and hindfemora medially, left lateral side of the abdomen, distal parts of the both right wings, and partly the posterior part of the right forewing; left posterolateral side of the abdomen is damaged; distal part of the right forewing considerably and scutellum, postscutellum, and propodeal dorsal side from the left partly deformed) in a polished DIGGER WASPS IN BURMESE AMBER Fig. 1 Burmastatus triangularis gen. et sp. nov., holotype, In.20197, female, Burmese amber. Length 3.2 mm. triangular-round sample of Burmese amber (probably Upper Creta- ceous). DESCRIPTION. Female. Head rounded, weakly flattened; pronotum moderately elongate, with flattened convex collar, prolong lateral ridges, and sharp keel along posterior margin; scutum strongly convex anteriorly, considerably higher than pronotal posterior mar- gin, straight posteriorly; scutellum strongly roundly convex, twice as short as scutum; postscutellum flat, twice as short as scutellum; mesopleuron convex, episternal and scrobal sulci present; metapleuron with broadly separated metapleural pits and weakly expressed metapleural flange; midcoxae with two, hindcoxae with only external carinae; fore trochanters elongate, mid- and hind- trochanters simple; main part of forewing vein placing in proximal half of wing; hindwings with a row of 4 hamuli; propodeal dorsal area with wide enclosure reaching 2/3 of its length from base; propodeal lateral sides outlined dorsally by lateral carinae, ending anteriorly below oval spiracles; propodeal hind side outlined dorsally by curved carinae; abdominal tergite I shorter than wide, with thin medial furrow anteriorly. Mandible ventrally and labrum with very long bristles; pronotum and scutum covered with very short erect and partly semi-adjacent hairs; mesopleuron ventrally covered with hairs, which length equal to hind basitarsus thickness; forecoxae ventrally covered with erect hairs 1.5 times longer than apical antennomere thickness; tibiae with dense short semi-adjacent hairs and with considerably longer and thicker outer spines lengthening towards apices, sparse on fore- and most dense on hindtibiae; fore tarsomeres with dense and short semi- adjacent hairs and sparse apical and longer lateral spines, one of which on I-III tarsomeres longer than following tarsomere, though real rake absent; mid- and hind tarsomeres with dense and short semi-adjacent hairs and with apical spines thinner than those on fore tarsomeres and ventral spines approximately 2 times shorter than those on fore tarsomeres. Body surface weakly sculptured, without distinct punctures, with visible surface dull. Yellowish-red palpi, scape, head ventrally, mesopleuron anteriorly and ventrally, legs completely, and large medial spot on abdominal tergite I. Brown antenna, mandible mainly, pronotum, scutum later- ally and along adlateral lines, scutellum, mesopleuron dorsally and laterally, metapleuron, wing veins, propodeum laterally, and abdomen 61 Figs 2-6 Burmastatus triangularis gen. et sp. nov., holotype, In.20197, female, Burmese amber. 2, Total view laterally. 3, Head, dorsolaterally. 4, Mandibles ventrolaterally. 5, Left antenna laterally; 6, Right wings (a — forewing, b — basal fragment of hindwing). Scale bars = 1 mm. mainly. Black mandible apically, occiput, scutum medially, propodeal dorsal enclosure and hind side, and abdominal tergite III medially. Body length 3.2 mm. The male is unknown. Subfamily CIRROSPHECINAE nov. TYPE GENUS. Cirrosphex gen. nov., here designated. DIAGNOSTIC FEATURES. 1. Antennal sockets covered dorsally by separated frontal prominences. 2. Occipital carina forming a complete circle, almost touching hypostomal carina. 3. Mandible without externoventral angle or notch, with inner preapical tooth, curved forward (slightly prognathous). 4. Palpi not elongate; palpal formula 6-4. 5. Male antenna 13-segmented; flagellomeres (except apical arti- cle) posteriorly with flat triangular prominences, longest on flagellomeres 5-9. 6. Pronotum short, straight posteriorly. 7. Adlateral lines reaching scutal posterior margin. 8. Midcoxae distinctly separated, with dorsolateral ridge. 9. Midtibia with one apical spur. 10. Hindfemur simple apically. 11. Tarsi simple, plantulae absent, tarsal claws probably with a tooth, arolium not shorter than claw. 12. Propodeum not elongate, rounded, without lateral carinae. 13. Forewings with 3 submarginal cells (I and II almost equal in size, II] smallest) and two discoidal cells; both recurrent veins received by submarginal cell IJ; marginal cell large, with acute apex touching wing margin; forewing media diverging beyond cu-a. 14. Hindwing jugal lobe small, rounded; hindwing media diverg- ing before cu-a. 15. Abdomen without separate petiole and with 7 visible seg- ments; tergite I with lateral carina; sternite I with medial ridge; tergite VII sharp. 16. Cerci absent. COMPARISONS. The new subfamily differs from recent sphecid subfamilies, excluding Ampulicinae, by the well developed, long adlateral lines and dentate tarsal claws. Among the representatives of Ampulicinae the new subfamily resembles the fossil Protodolichurus Nemkov, 1988 (Dolichurini) in having separated frontal prominences above the antennal bases, a short pronotal collar, a more or less rounded propodeum, and completely visible abdominal segments. It alsoresembles the recent genera Trirogma Westwood, 1841 (Dolichu- rini) in the hindwing venation and Ampulex Jurine, 1807 (Ampulicini) in having no cerci and a sharply prominent clypeus bearing a medial ridge. Cirrosphecinae subfam. nov. differs from Ampulicinae most notably by the single midtibial spur, very short and low pronotal collar, expressed episternal sulcus, and also by the pectinate flagellum. REMARKS. The form of the pronotum (short collar with a straight posterior margin and distinct pronotal lobes) and of the hindtibial cleaning brush shows that the specimen belongs to the Sphecidae. It is similar to Ampulicinae by the adlateral lines and dentate tarsal claws [the studied specimen has a visible tooth only on the inner claw of the right hindtarsus (the rest of the claws are completely covered by foreign particles or blebs)], which are the diagnostic features for the last, and also by the prognathously curved mandibles, which are usual for the females of Ampulex. As for the form of the mandible, some males of recent Sphecidae often have considerably stronger modified mandibles, and their form may be considered to be a sexual signal. The expressed frontal prominences above the antennal sock- ets also occurs outside the Ampulicinae [for example, both single and separated prominences are usual in some species-groups of the genus Trypoxylon Latreille, 1796 (Crabroninae, Trypoxylini)]. Distinctive features, which are not characteristic for Ampulicinae, are the form of the flagellomeres, the reduction of the 2nd apical midtibial spur, the contiguous hindcoxae, the strongly enlarged submarginal cell Il and diminished submarginal cell III, the deli- cately sculptured propodeum without lateral carinae, and the elongate first abdominal segment bearing a medial ridge on the sternite. Probably, a more detailed study of the specimen will give additional characters substantiating its isolation as a separate subfamily (unfor- tunately, in its present state the sample is open for study only from a certain visual angle on its left side). Genus CIRROSPHEX nov. ETYMOLOGY. The generic name is derived from the Latin noun cirrus, meaning ‘fringe’, and the generic name Sphex. The name is masculine. A.V. ANTROPOV TYPE SPECIES. the only species. Cirrosphex admirabilis sp. nov., here designated; DIAGNOSIS. Clypeal medial lobe sharply prominent, with medial ridge. Lateral ocelli simple. Antennal bases covered dorsally by separate frontal prominences; male antenna 13-segmented; flagellomeres (except apical article) posteriorly with flat triangular prominences, mostly expressed on flagellomeres 5—9. Mandible short, curved forward, with inner preapical tooth. Palpi not elongate; palpal formula 6—4. Occipital carina forming a complete circle, almost touching hypostomal carina. Pronotal lobes almost touching tegulae. Pronotum posteriorly noticeably lower than upper scutal level. Adlateral lines reaching scutal posterior margin; admedial lines contiguous, longer than scutal length half. Mesopleuron with coarse episternal sulcus, scrobal sulcus, and precoxal carina; omaulus and sternaulus absent; intercoxal carina straight. Legs simple; hindcoxae contiguous; tarsal claws with a tooth. Propodeum without lateral carinae, with deep medial furrow posteriorly. Forewing mar- ginal cell long and broad, with acute apex touching wing margin; three submarginal cells, II slightly larger than I, II more than twice as large as widened apically III; both recurrent veins received by submarginal cell II; veins M and RS slightly curved; forewing media diverging slightly beyond cu-a. Jugal lobe small, rounded; hindwing media diverging before cu-a. Abdomen with 7 visible segments, without separate petiole; abdominal sternite I with medial ridge, sternite II uniformly convex, without transverse basal concavity, sternite VII strongly tapered, thorn-shaped; tergite VII sharp, with- out pygidial plate; cerci absent. COMPARISONS. The obvious prevalence of the second submarginal cell over the third is unusual for recent (and known fossil) Sphecidae and is found by way of an exception, for example, in the genera Podalonia Fernald, 1927 (Sphecinae, Ammophilin1), Diploplectron W.Fox, 1893 (Astatinae, Astatini), Tachytella Brauns, 1906 (Crabroninae, Larrini), and Eremiasphecium Kohl, 1897 (Philanthinae, Eremiaspheciini). In all these cases, the second sub- marginal cell does not reach the size of the first submarginal cell and the marginal cell is always apically truncate (Diploplectron) or its apex does not touch the wing margin (the other genera), 1.e., its diminution is associated with a general reduction in size of the distal forewing cells, including the marginal cell. The new genus, however, has a long marginal cell with its apex touching the wing margin, an enlarged second submarginal cell and a partly diminished third submarginal cell. Such reduction of the forewing venation, including only the distal part of the forewing submarginal cells, is presently unknown in recent Sphecidae. Cirrosphex admirabilis sp. nov. Figs 7-10 ETYMOLOGY. The species name admirabilis is derived from the Latin masculine adjective, meaning ‘astonishing’; the name is a reference to the unusual form of the male antennae. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.19125(1); completely preserved male speci- men in a polished, flat, broadly ovate piece of Burmese amber (probably Upper Cretaceous). DESCRIPTION. Male. Malar space short; occiput weakly expressed. Scape not more than twice as long as its maximum thickness; pedicellus almost round; flagellomeres 1-6 quadrate, 7—11 longer than wide, apical flagellomere 2.7 times longer than thick; flagellomeres 1—10 with flattened triangular prominences posteriorly, shortest at flagellomeres 1-4 and 10 and longest at flagellomeres 8— 9. Frontal prominences above antennal sockets small, covering DIGGER WASPS IN BURMESE AMBER “= oe . i Fig. 7 Cirrosphex admirabilis gen. et sp. nov., holotype, In.19125(1), male, Burmese amber. Length 3.5 mm. 10 Figs 8-10 Cirrosphex admirabilis gen. et sp. nov., holotype, In.19125(1), male, Burmese amber. 8, Total view laterally. 9, Left antenna laterally. 10, Left wings. Scale bars = 1 mm. 63 antennal bases dorsally from inside. Pronotal collar short, consider- ably lower than dorsal level of distinctly convex anteriorly scutum. Scutellum considerably roundly convex. Postscutellum short, strongly prominent medially. Mesopleuron convex, weakly sculptured, shiny. Dorsal part of metapleuron higher than 1/3 of its total height. Tarsomeres thin, cylindrical, without plantulae. Apical tarsomeres with small rounded arolium. Propodeal dorsal enclosure flat-con- cave, enclosed laterally with smooth furrows; propodeal hind side with deep medial furrow; propodeal spiracles oval, open. Abdomen somewhat elongate; tergite I from the base with lateral carinae, prolongly carinate; sternite I with medial ridge; tergite VII without pygidial plate, tapered apically; sternite VII strongly tapered, thorn- shaped. Head anteriorly, scutum, scutellum, mesopleura ventrally, postscutellum, tergites V—VI, and sternites II-VI apically with dis- tinct, erect pubescence, equal in length to lateral ocellus diameter or some longer at abdominal apex. Metapleuron, propodeum, and first abdominal segment bare. Foretibia and basitarsus with long, erect, and somewhat curved hairs. Body length 3.5 mm. The female is unknown. II. New tribes of the subfamily Ampulicinae The only fossil ampulucine known so far, Protodolichurus sucinus Nemkov, 1988, was described from the Middle Eocene Baltic amber and was a representative of the tribe Dolichurini. Due to the character- istic features of short pronotum and unmodified abdominal sclerites, it was placed by Nemkov near the hypothetical ancestor of Dolichurini. A second species, Trigonalys pervetus Cockerell, 1917, from the Upper Cretaceous Burmese amber, was erroneously placed in the family Trigonalyidae. The Burmese amber collection at The Natural History Museum also contains four new species belonging to three new genera of Ampulicinae. Subfamily AMPULICINAE 1. Tribe ARPODOLICHURINI nov. TYPE GENUS. Apodolichurus gen. nov., here designated. DIAGNOSTIC FEATURES. 1. Lower frons uniformly convex, without visible prominences. 2. Ocelli simple. 3. Palpi not elongate. 4. Female antennae 12-segmented, long and filiform; pedicel elongate, only slightly shorter than first flagellomere. Pronotum elongate, with deep and wide medial concavity and more or less expressed lateral prolong prominences. Mid- and hindcoxae separated. Midtibia with two apical spurs. Tarsomeres simple, with plantulae. Tarsal claws simple. Forewing venation reduced: marginal cell very short, with anterior part shorter than stigma; enclosed submarginal and discoidal cells absent; submedial cell very short; forewing media diverging far beyond cu-a. 10. Hindwing jugal lobe small, rounded. Medial cell triangular. Hindwing media diverging considerably beyond cu-a. 11. Abdomen with 6 visible segments, without petiole, joined above hindcoxal bases. 12. Cerci absent. es Memon) ESS COMPARISONS. The two representatives of the new tribe differ from other sphecids by the strong specialization of venation of both fore- and hindwings, expressing, firstly, the diminution of the marginal 64 cell and almost complete reduction of the submarginal, discoidal, and subdiscoidal cells, and secondly the proximal removal of only the externoposterior corner of the medial cell. Each of the features taken separately are found in various recent Sphecidae (acute and short marginal cell in Pemphredoninae, Ammoplanini; more or less disappearing submarginal, discoidal, and subdiscoidal cells in Crabroninae, Miscophini; almost triangular hindwing medial cell in Crabroninae, Crabronini). Simultaneous combination of these features is not found in any known recent or fossil Sphecidae. All the more, it is not found in Ampulicinae with their typically complete venation of both wings having minimum specialization. Another important (and unique) difference of the new tribe from all known Ampulicinae is the edentate tarsal claws. Finally, Apodolichurini trib. nov. differs from other Ampulicinae by its ovally elongate propodeum without distinct bend between dorsal and posterior parts and also by the spherical head and strongly elongate antennal pedicel (observable in only one of the species described below, the head was not preserved in the second species). REMARKS. The reduction of the forewing venation is probably related to reduction of body size (maximum length of the discovered species is hardly more than 3.0 mm). At the same time, the character- istic shape of the pronotum, expressed adlateral lines, and two midtibial apical spurs allow the placement of both described species in the subfamily Ampulicinae. Genus APODOLICHURUS nov. ETYMOLOGY. The generic name is derived from the Greek prefix apo-, meaning ‘from’, and the generic name Dolichurus; it is mascu- line. TYPE SPECIES. Apodolichurus sphaerocephalus sp. nov., here des- ignated. DIAGNOSIS. Head spherical; palpi not elongate; antennal sockets not covered by frontal prominences; antennae of female 12-seg- mented, long and filiform, with pedicel almost as long as Ist flagellomere; vertex roundly convex; occipital carina strongly devel- oped, flat-widened ventrolaterally (ventral ends not visible). Pronotum distinctly longer than scutum; pronotal collar approxi- mately as long as scutum, with two lateral ridges. Scutum transverse, with almost complete adlateral and weak admedial lines; scutellum flat, rectangular or trapeziform, transverse; postscutellum short; mesopleuron uniformly convex, with distinct episternal sulcus (other mesopleural sulci and carinae not expressed); intercoxal carina almost straight. Mid- and hindcoxae separate; femora thickened, tibiae and tarsi thin and long; tarsomeres simple, with apical plantulae ventrally (at least at right hindtarsomeres II-IV of the type species because other tarsi are visible only from external side); apical tarsomeres with weak arolium and edentate tarsal claws. Forewing marginal cell very short, with acute apex touching wing margin; submarginal, discoidal, and subdiscoidal cells absent; submedial cell very short, forewing media diverging far beyond cu-a. Hindwing medial cell almost triangular, media diverging beyond cu-a. Propodeum oval-elongate, with weakly developed lateral carinae, not bent between dorsal and hind sides, tubularly elongate apically. Abdominal tergite VI with medial ridge. INCLUDED SPECIES. A. diaphanus sp. nov. Apodolichurus sphaerocephalus sp. nov. and COMPARISONS. Apodolichurus gen. nov. resembles Protodolichurus only in the form of propodeum and some species of Ampulex in the A.V. ANTROPOV Fig.11 Apodolichurus sphaerocephalus gen. et sp. nov., holotype, In.20150, female, Burmese amber. Length 3.1 mm. 13 Figs 12,13 Apodolichurus sphaerocephalus gen. et sp. nov., holotype, In.20150, female, Burmese amber. 12, Total view dorsolaterally. 13, Left wings (a — forewing, b — anterior fragment of hindwing). Scale bars = 1 mm. DIGGER WASPS IN BURMESE AMBER form of pronotum. It differs from both recent and fossil genera of Ampulicinae by the tribal diagnostic features, particularly by the reduced forewing venation, principally different form of the hindwing medial cell, elongate antennal pedicel, and edentate tarsal claws. Apodolichurus sphaerocephalus sp. nov. Figs 11-13 ETYMOLOGY. The species name is derived from the Greek nouns sphaera, meaning ‘sphere’ , and cephalon, meaning ‘head’; the name is a reference to the shape of the female head. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.20150; completely preserved female speci- men in a broad and flat wedge-shaped sample of Burmese amber (probably Upper Cretaceous), polished on one side, and containing numerous inclusions. DESCRIPTION. Female. Head spherical; frons noticeably convex, slightly overgangling over antennal bases, with narrow medial fur- row; eyes small; ocelli small, disposed in almost equilateral triangle; occipital carina laterally flattened and bent outside; antennae long and thin with cylindrical antennomeres. Pronotum with strongly developed, high, and pointed posteriorly prolong lateral ridges; scutum hardly convex, transverse; scutellum rectangular; trochanters (especially fore ones) thin and strongly elongate; femora (especially fore ones) noticeably bulbous ventrally; tibiae and tarsi thin, un- modified, only with short apical spines. Wings with very delicate and almost bare membrane; forewing without trace of Cu; hindwing with a long row of 10 small hamuli. Propodeum elongate, densely and uniformly transversely carinate laterally and posteriorly (invisible dorsally). Abdomen without petiole (first segment detached from propodeum and connected with it by stretched membrane), with Fig. 14 Apodolichurus diaphanus gen. et sp. nov., holotype, In.19123(4), Burmese amber. Length 1.2 mm. 65 tergites almost completely covering sterna laterally; tergite VI sharp, with short medial carina. Body very weakly sculptured: head dull, scutum delicately punc- tate and weakly shiny, abdomen microsculptured and weakly shiny. Pubescence practically absent. Head black; rest parts of body (especially legs) considerably lighter — from brownish-red to amber-red; wing veins whitish- yellow. Body length 3.1 mm. The male is unknown. REMARKS. It is likely that the abdomen of the specimen in its life- time was weakly sclerotized; the sclerites are covered by numerous irregular, prolonged wrinkles. Apodolichurus diaphanus sp. nov. Figs 14-16 ETYMOLOGY. The species name diaphanus is derived from the Latin masculine adjective, meaning ‘translucent’; the name is a reference to the appearance of the body. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.19123(4); partly preserved specimen of un- known sex (the following parts are absent: head, left midtarsus after the basitarsal proximal half, left hindleg after the femoral base, right 16 15 Figs 15,16 Apodolichurus diaphanus gen. et sp. nov., holotype, In.19123(4), Burmese amber. 15, Total view dorsally (scale bar = 1 mm); 16, Position of holotype in amber sample In.19123. 66 foreleg, right hindtarsus after the 3rd tarsomere’s base, left wings, and right hindwing, and apical abdominal segments; right forewing membrane is torn; sclerites of the abdominal segments I-III are separated) lying in the obtuse end (Fig. 16) of the irregularly ovate, flat, and broad wedge-shaped sample of Burmese amber (probably Upper Cretaceous) containing numerous inclusions. DESCRIPTION. Sex unknown (head and abdomen apically absent). Pronotum with weakly prolonged and not angled posteriorly lateral ridges; scutellum trapeziform, narrowed posteriorly; trochanters not elongate; femora moderately thickened; tibiae and tarsi thin, un- modified, only apically with short spines. Forewing with thin trace of Cu. Propodeum uniformly narrowed towards apex; propodeal dorsal enclosure triangular, with thin medial and irregular lateral carinae, distinctly margined by broad transversely carinate concavity; propodeal hind side with thin medial ridge, densely and uniformly transversely carinate laterally. Abdominal sclerites separated (pre- served only a few sclerites lying apart from the thorax). Scutum delicately punctate, with moderately shiny interspaces. Abdominal sclerites almost unsculptured, weakly shiny. Pubescence not expressed. All preserved body fragments almost completely discoloured, translucent, amber-reddish; wing veins completely discoloured. Body length 1.2 mm (without head and abdomen). COMPARISONS. ‘This species differs from A. sphaerocephalus sp. nov. by the considerably weaker lateral prominences of the pronotum, by simple trochanters, noticeably weaker bulbous femora, and by the distinct trace of the forewing Cu. REMARKS. The translucence of the sclerites, absence of the head and the main part of the wings, and also separation of the abdominal sclerites lying separately far from the thorax, indicate that the fossilized specimen was not alive but already dead prior to entrap- ment, with almost or completely destroyed subcuticular structures, including intersegmental ligaments, muscles, and internal organs. 2. Tribe CRETAMPULICINI nov. TYPE GENUS. Cretampulex gen. nov., here designated. DIAGNOSTIC FEATURES. 1. Inner eye orbits parallel; ocelli simple. 2. Antennal sockets touching clypeal base; female antennae 12- segmented. Clypeus uniformly convex, triangular, with medial carina. Lower frons with prominence above each antenna. Occipital carina expressed, at least dorsally. Female mandibles hypognathous, straight, bidentate apically. Palpi considerably elongate (maxillary palpi not shorter than head width). 8. Metasternum obtusely emarginate posteriorly, but not Y-shaped. 9. Hindcoxae separated. 10. Tarsomeres unmodified, without plantulae; tarsal claws with a ventral tooth. 11. Hindwing jugal lobe small, oval. Hindwing medial cell weakly elongate distally, with RS hardly longer than r-m. 12. Abdomen with 6 visible segments, with petiole inserted on level of hindcoxal bases. 13. Cerci absent. Sa SS COMPARISONS. Cretampulicini trib. nov. differs from all tribes of the subfamily Ampulicinae by the apically bidentate mandibles (unknown in Apodolichurini trib. nov. and Trigampulex gen. nov.), and from the recent tribes of the subfamily by the different form of A.V. ANTROPOV the hindwing medial cell (RS hardly longer than r-m). Furthermore, Cretampulicini trib. nov. differs from the tribe Dolichurini by the sharp medial carina of the triangular clypeus, the separated hindcoxae, the absence of plantulae, and by the absence of a transverse concay- ity or carina on the abdominal sternite II. It differs from Ampulicini by the elongate palpi, moderately emarginate but not Y-shaped metasternum, and unmodified tarsi. Cretampulicini trib. nov. differs from Apodolichurini trib. nov. by the pair of lower frontal prominences above the antennal bases, elongate palpi, dentate tarsal claws, com- plete forewing venation, and by the ovally elongate hindwing jugal lobe. It differs from Mendampulicini trib. nov. by the pair of lower frontal prominences above the antennal bases, simple eyes, and by the triangular clypeus with a sharp medial carina. Finally, it differs from Trigampulex gen. nov. by the complete forewing venation (the discoidal cell If completely developed, trace of the forewing Ir absent, recurrent vein I received by the submarginal cell II), elongate propodeum, and by the developed abdominal petiole. It differs from Ampulicini by the marginal cell apex touching the wing margin Genus CRETAMPULEX nov. ETYMOLOGY. The generic name is derived from the Latin mascu- line adjective cretaceus, indicating the age of the specimen, and the generic name Ampulex. It is masculine. TYPE SPECIES. only species. Cretampulex gracilis sp. nov., here designated; the DIAGNOSIS. Clypeus triangular with sharp medial carina; lower frons with a pair of small prominences, hardly covering antennal sockets above; vertex simple; eyes unmodified, with parallel inner orbits; ocelli simple; occipital carina distinct, at least dorsally (ven- tral part of head invisible); mandibles hypognathous, slightly curved apically, with small inner preapical tooth; palpi elongate, maxillary palpi hardly shorter than head width; palpal formula 6-4; antennae long and slender, 12-segmented in female. Pronotum elongate, with pronotal collar concave medially and with sharp lateral ridges; scutum convex (insufficiently visible from above); mesopleuron weakly convex; episternal sulcus thin; omaulus, scrobal sulcus, acetabular carina, subomaulus, and sternaulus not expressed; precoxal carina distinct, slightly continuing on mesopleuron; scutellum flat, short; postscutellum convex, short; metasternum emarginate posteriorly, but not Y-shaped; mid- and hindcoxae separated; legs long and slender, with unmodified tarsomeres; plantulae absent; arolium small, tarsal claws with a tooth. Forewing marginal cell with acute apex touching wing margin, 1.5 times longer than stigma; three submarginal cells: I largest; II trapeziform, narrowed anteriorly (though Ir absent), 1.5 times narrower than I, Ist transverse submar- ginal vein angulately curved; submarginal cell III almost rectangular, 1.5 times narrower but longer than II; two discoidal cells: recurrent vein I received by submarginal cell II, recurrent vein II — by submar- ginal cell III; forewing media diverging beyond cu-a. Hindwing with a row of 7 hamuli; RS short, almost equal to r-m; angle between RS and r-m obtuse; hindwing media diverging at cu-a; jugal lobe elon- gate, approximately equal to 1/3 of submedial cell length. Propodeum long, distinctly bent between dorsal and hind sides; hind part margined by sharp carinae from all sides; lateral carinae not expressed. Abdo- men with thin petiole, inserted on level of hindcoxal bases, and with unmodified segments II-VI; tergite VI conical, without pygidial plate, but with thin medial carina. COMPARISONS. Cretampulex gen. nov. differs from all recent and known fossil genera of Ampulicinae by the apically bidentate man- DIGGER WASPS IN BURMESE AMBER dibles and comparatively small number of hamuli. Among recent genera it resembles Aphelotoma Westwood, 1841 and Austrotoma Riek, 1955 in having the separated lower frontal prominences, differing from them by the medially carinate triangular clypeus, apically bidentate mandibles, elongate palpi, strong lateral ridges of the pronotal collar, absence of the omaulus, sternaulus, tarsal plantulae, and forewing vein Ir, by the narrow forewing submarginal cell III and the relative positions of the recurrent veins, and by the developed hindwing jugal lobe and the abdominal petiole. Among fossil representatives of the subfamily, Cretampulex gen. nov. resembles Protodolichurus in having the separated lower frontal prominences, the medially carinate clypeus, and similar forewing venation, differing from it by the triangular clypeus, longer pronotum with the expressed lateral ridges of its collar, absence of tarsal plantulae and forewing vein Ir, by the narrow submarginal cell II, the form of the hindwing medial cell and confluent cu-a, by the distinct bend between the propodeal dorsal and hind sides, and by the distinct abdominal petiole. Figs 17-20 ETYMOLOGY. The species name is derived from the Latin mascu- line adjective gracilis, meaning ‘slender’; the name is a reference to the shape of the female body. Cretampulex gracilis sp. nov. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.19123(5); completely preserved female speci- men lying in the centre (Fig. 20) of the polished from the both sides, irregularly ovate, flat, and broad wedge-shaped sample of Burmese amber (probably Upper Cretaceous), containing numerous inclusions. DESCRIPTION. Female. Head oval, higher than wide; clypeus trian- gular, with distinct medial carina; frons weakly convex, with narrow medial furrow; separated frontal prominences small, covering antennal sockets above from inside; malar space longer than maxi- mum scape thickness; vertex weakly and uniformly convex; ocelli small, forming an obtuse-angled triangle; oculo-ocellar distance longer than that between lateral ocelli; occipital carina expressed at least dorsally and laterally (ventral part of head invisible); antennal sockets touching clypeal base; mandible rather thin and somewhat widened apically, with weakly curved inside apical and shorter inner preapical teeth; antenna long and slender, with unmodified flagellomeres at least thrice as long as thick, last article rounded apically; palpi distinctly elongate, maxillary palpi not shorter than head width. Pronotum elongate, with medial concavity and distinct, acute-angled dorsoposteriorly lateral ridges; scutum uniformly con- vex (thorax and propodeum invisible dorsally); mesopleuron with distinct episternal sulcus and some concave prolonged lines (prob- ably, their part may be the result of deformation, but not real sulci); legs practically unmodified, without outer tibial spines, but with short ventral bristles on tarsomeres, most distinct on foretarsus. Propodeum elongate, bent between dorsal and hind sides. Abdomen with distinct petiole (it’s detailed structure invisible), with tergites considerably covering sternites laterally; tergite VI sharp-conical, with thin medial carina (prolongly split apically, with right apical part turned up); sting thin and straight. Body sculpture weak, expressed as moderate punctures at frons and scutal visible part, with weakly shiny interspaces; abdomen microstriate, shiny. Clypeus with sparse erect bristles; pronotal lateral ridges, scutum and propodeum at least dorsally, and lower margin of abdominal tergite VI covered with dense erect hairs. Body more or less uniformly darkened (probably, brownish-black); wing veins dark-brown; palpi and tarsi somewhat lighter than body. Body length 3.7 mm. The male is unknown. 67 Fig.17 Cretampulex gracilis gen. et sp. nov., holotype, In.19123(5), female, Burmese amber. Length 3.7 mm. 19 20 Figs 18-20 Cretampulex gracilis gen. et sp. nov., holotype, In.19123(5), female, Burmese amber. 18, Total view ventrolaterally. 19, Wings (a — left forewing, b — right hindwing). 20, Position of holotype in amber sample In.19123. Scale bars = 1 mm. 68 3. Tribe MENDAMPULICINI nov. TYPE GENUS. Mendampulex gen. nov., here designated. DIAGNOSTIC FEATURES. 1. Mandibular sockets open, united with oral fossa. 2. Clypeus uniformly convex, not carinate medially. 3. Lower frons with unpaired prominence. 4. Occipital carina forming a complete circle, considerably sepa- rated from hypostomal carina. 5. Mandible hypognathous, crescent. 6. Palpi considerably elongate. 7. Metasternum emarginate posteriorly, but not Y-shaped. 8. Mid- and hindcoxae separated. 9. Tarsomeres unmodified. 0. Abdominal petiole inserted between and on same level as hindcoxae. COMPARISONS. This new tribe is characterized by the presence of the characters listed above, which are typical to the Ampulicinae. It differs from all other tribes by the long crescent mandibles, compara- tively small eyes, and the occipital carina forming a complete circle (in the studied specimens of Apodolichurini trib. nov. and Cretampulicini trib. nov. the occipital carina is visible only dorsally, and it is unknown for Trigampulex gen. nov. because of the absence of the holotype head). Furthermore, Mendampulicini trib. nov. dif- fers from Dolichurini by the separated hindcoxae and the abdominal petiole inserted between and on the same level as the hindcoxae; from Ampulicini by the clypeus without a medial carina, unpaired frontal prominence, open mandibular sockets, hypognathous mandi- bles, considerably elongate maxillary and labial palpi, thin and unmodified tarsomeres (without ventral pubescence on the penulti- mate tarsomeres and with an arolium on the apical tarsomeres), not Y-shaped metasternum, and by the fore- and hindwing mediae diverging beyond cu-a; from Apodolichurini trib. nov. by the consid- erably flattened head, pronotal collar without expressed lateral ridges, long scutum, and by the dentate tarsal claws; from Cretampulicini trib. nov. by the clypeus lacking a medial carina, unpaired frontal prominence, strongly developed vertex, undeveloped lateral ridges of the pronotal collar, hindwing media diverging beyond cu-a, and by undeveloped hindwing jugal lobe; from Trigampulex gen. nov. by the hindwing media diverging beyond cu-a and by the petiolate abdo- men. REMARKS. Mendampulicini trib. nov. resembles Dolichurini, at least the genus Trirogma, by the main diagnostic features (trapezifom clypeus without medial carina, unpaired frontal prominence, elon- gate palpi, and unmodified tarsi). The crescent mandibles, the flattened head with a complete occipital carina and small eyes, and the hindwing media diverging beyond cu-a testify to the high spe- cialization of the fossil specimen and separate it from other ampulicine tribes. Genus MENDAMPULEX nov. ETYMOLOGY. The generic name is derived from the Latin mascu- line adjective mendax, meaning ‘false’, and the generic name Ampulex. It is masculine. TYPE SPECIES. Mendampulex monilicularis sp. nov., here desig- nated; the only species. DIAGNOSIS. Clypeus trapeziform, with uniformly convex surface and without medial carina; lower frons with obtuse-angulately cari- nate frontal prominence hardly covering dorsally antennal sockets; A.V. ANTROPOV Fig. 21 Mendampulex monilicularis gen. et sp. nov., holotype, In.20711, female, Burmese amber. Length 4.7 mm. vertex considerably elongate; eyes small, strongly convex, with strongly curved inner orbits; ocelli simple; occipital carina distinct, forming a complete circle, separated from hypostomal carina; man- dible hypognathous, long, crescent, with small outer external preapical notch and inner triangular apical tooth; maxillary palpi not shorter than head height, labial palpi also elongate; palpal formula 6—4; antennae long and slender, 12-segmented. Pronotum elongate, with collar slightly broader than long, with medial concavity, and sharply margined laterally; scutum flat, rectangular, slightly longer than pronotal collar, with complete adlateral and hardly expressed admedial lines; mesopleuron convex; episternal sulcus coarse and broad; omaulus, scrobal sulcus, and acetabular carina not expressed; subomaulus and sternaulus absent; precoxal carina strongly devel- oped, not continuing on mesopleuron; scutellum flat, square; postscutellum convex, short; metasternum emarginate posteriorly, not Y-shaped; mid- and hindcoxae separated. Legs long and slender, with unmodified tarsomeres; ventral plantulae absent; arolium small, sharp apically; tarsal claws thin and long, with a tooth near apex. Fore- and hindwing mediae diverging beyond cu-a; hindwing jugal lobe not expressed. Propodeum long, abrupt posteriorly, with U- shaped dorsal enclosure, margined laterally and posteriorly by sharp ridges. Abdominal petiole inserted on level of hindcoxal bases. COMPARISONS. Mendampulex gen. nov. resembles Trirogma, Paradolichurus Williams, 1960, and Dolichurus Latreille, 1809 in having the unpaired, though weaker frontal prominence, but differs from them by the stronger separated antennal sockets, longer pronotum, and by the fore- and hindwing mediae diverging beyond cu-a. Furthermore, Mendampulex gen. nov. differs from the first genus by the long palpi and undeveloped omaulus, scrobal sulcus, and sternaulus; from the second by the undeveloped omaulus and by the tarsi without plantulae; and from the last by the trapeziform clypeus, undeveloped omaulus, scrobal sulcus, sternaulus, acetabu- lar carina, and by the tarsi without plantulae. DIGGER WASPS IN BURMESE AMBER Figs 21-25 ETYMOLOGY. The species name is derived from the Latin neuter noun monile, meaning ‘beads, necklace’, and a diminutive suffix -culum; the name is a reference to the position of the specimen inside the small bead. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.20711; partly preserved female specimen (the following parts are absent: left flagellomeres 1-10 and right flagellomeres 3-4, left foretarsomeres II—V, left midtarsus, both hindlegs after coxae, wings after submedial cell, and abdomen after petiole) in an almost spherically polished bead of Burmese amber (probably Upper Cretaceous). Mendampulex monilicularis sp. nov. DESCRIPTION. Female. Head considerably flattened, with strongly developed vertex; clypeus trapeziform, roundly truncate apically, with obtuse angles laterally; frons flat-convex, ocellar triangle slightly concave; medial ocellus larger than lateral one; eyes small and convex, almost twice as high as distance between inner orbits dorsally; oculo-ocellar distance approximately twice as long as that between lateral ocelli; malar space not longer than lateral ocellar diameter; occipital carina separating from hypostomal farther than 24 Figs 22-24 Mendampulex monilicularis gen. et sp. nov., holotype, In.20711, female, Burmese amber. 22, Head (a — anteriorly, b — posteriorly, c — laterally). 23, Mandibles apically (a — right, frontally, b — left, ventrally, c — left, posteroventrally). 24, Left foretarsomeres III-V (a — dorsally, b — laterally). Scale bars = 1 mm. 69 Fig. 25 Mendampulex monilicularis gen. et sp. nov., holotype, In.20711, female, Burmese amber; total view (a — dorsally, b — ventrally, c — laterally); scale bars = 1 mm. 70 maximum thickness of mandibular base; mandible crescently curved, without inner teeth, with small external preapical notch and triangu- lar inner apical angle; middle flagellomeres 4—5 times as long as thick. Tarsomeres long, slightly broadened apically; apical tarsomere longer than penultimate one; small sharped apically arolium twice shorter than tarsal claw. Propodeum with coarsely margined and radially carinate inside and from outside dorsal enclosure. Clypeal apical margin and external and inner margins of mandible with rows of long erect bristles, those 1.5—2.0 times as long as thickness of mandibular mid part; pro- and mesothorax mainly covered with very short, directed backwards, appressed hairs practi- cally hiding dull and inpunctate surface; precoxal carina posteriorly with longer and also directed backwards dense bristles; tarsomeres I-IV ventrally with rows of dense short bristles and sparser and longer thin ventral (mainly on tarsomere I) and apical spines; apical tarsomere practically bare ventrally. Body mainly dark-brownish, without separate spots; head com- pletely, mandibles basally and apically, apical half of scapes, foretibiae apically, midcoxae, and mesopleura posteriorly black; midfemora and tibiae mainly yellow. Body length 4.7 mm (without abdomen). The male is unknown. III. A new genus of Sphecidae Genus TRIGAMPULEX nov., incertae sedis ETYMOLOGY. The generic name is derived from the generic names Trigonalys and Ampulex. It is masculine. TYPE SPECIES. Trigonalys pervetus Cockerell, 1917, here desig- nated; the only species. DIAGNOSIS. Postscutellum short. Propodeum rounded, without distinctly margined dorsal enclosure and visible lateral carinae. Legs weakly modified: femora shorter than corresponding tibiae; midtibia with two apical spurs and sparse short spines outside; tarsi elongate, with apical tarsomere connected with apex of unmodified penulti- mate tarsomere; plantulae absent; tarsal claws with a tooth. Forewing with three submarginal cells and one discoidal cell (discoidal cell II rudimentary, margined from outside and posteriorly by scarcely visible traces of veins 2m-cu and Cu,). Hindwing medial cell with RS almost perpendicular to R1. Abdomen sessile, petiole undevel- oped. COMPARISONS. Based on the peculiarities of the forewing venation (recurrent vein I received by submarginal cell I and trace of the recurrent vein II received by submarginal cell III; medial cell weakly elongate), T:pervetus may be attributed to either the family Trigonalyidae or to the Sphecidae. The simple trochanters of the mid- and hindlegs, the midtibiae with two apical spurs, the absence of the plantulae at the apices of the unmodified tarsomeres, the tooth claws, the developed cleaning brush of the inner hindtibial spur and hind basitarsus, the weakly developed costal cells of both wings, the distinct trace of vein Ir in the forewings, the outer prolong veins (M-a and Cu.-a of the fore- and RS-a, M-a, Cu-a, and A, -a of the hindwings) ending far from the wing margin, the forewing media diverging distinctly beyond cu-a, the distinctly separated hindwing jugal lobe, and the single row of hamuli suggest a placement outside of Trigonalyidae (where it was originally placed by Cockerell, 1917b), but is suggestive of Sphecidae, perhaps of the subfamily Ampulicinae. Trigampulex pervetus differs from the representatives of Ampulicinae by the distinctly diminished forewing submarginal cell A.V. ANTROPOV Ill, by the reduced outer veins of the forewing discoidal cell II (2m-cu and Cu,), and by the form of the hindwing medial cell (RS almost perpendicular to R1). The last two features also place T:pervetus separately from all Sphecidae, but the inaccessibility for study of many important characteristics of the structure of the head and thorax (first of all, of the mesothorax) does not allow to interpret definitely this species as the representative of a separate subfamily. I therefore place the fossil as incertae sedis near Ampulicinae until new, more perfectly preserved material, is discovered. Trigampulex pervetus (Cockerell, 1917), comb.nov. Figs 26-31 1917 = Trigonalys pervetus Cockerell: 79, fig. 1 (Trigonalidae). HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.19093; partly preserved specimen of un- known sex (the following parts are absent: head, prothorax including forelegs, scutum and scutellum, right wings completely and left wings basally, right midleg from apical half of femur to basitarsus, right hindleg from apical half of femur to tibial middle, and abdomi- nal segments after segment IV; legs and propodeum considerably deformed) in a spindle-shaped polished bead of Burmese amber (probably Upper Cretaceous). DESCRIPTION. Sex unknown (head, prothorax, and abdomen apically lost). Preserved parts of mesopleura strongly deformed, with vague structure; postscutellum (left part preserved) short; legs very slender: femora (especially hind ones) distinctly shorter than corresponding tibiae; midtibiae ventrally in middle and hindtibiae basally obtuse-angled; tarsomeres very long, tarsomere I almost equal to corresponding tibia, apical tarsomere longer than penulti- mate one. Propodeum rounded (though strongly deformed, right up to the cuticular ruptures, those Cockerell has, probably, interpreted as bilobed scutellum). Abdomen, in comparison with wings and thorax, large, sessile; tergite I considerably smaller than each of the rest segments, especially tergite II. Coxae, trochanters, and partly femora covered ventrally with Fig. 26 Trigampulex pervetus (Cockerell), holotype, In.19093, Burmese amber. 20. Total view dorsally. Length 3.4 mm. DIGGER WASPS IN BURMESE AMBER 7A 28 : Figs 27-31 Trigampulex perveius (Cockerell), holotype, In.19093, Burmese amber. 27, total view dorsally; 28, left wings (a — forewing, b — hindwing); 29, Left midleg (a — tibia and tarsus dorsally, b — laterally). 30, Left hindleg (a — tibia and tarsus dorsally, b — laterally). 31, Right coxa, trochanter, and base of femur laterally (a — hindleg, b — midleg). Scale bars = 1 mm. short and dense erect hairs; mid- and hindtibiae with some ventral bristles, the last also with short and dense erect hairs; tarsomeres I— IV with similar hairs, apical tarsomeres bare; propodeum covered with longer, but sparser erect hairs; abdominal sclerites practically bare, visibly transversely microstriate, shiny. Body completely black; wing veins from black to brown; midtibiae reddish basally. Body length 4.0 mm (without head, anterior part of thorax, and abdominal apex), forewing length 3.4 mm. IV. New genera of the subfamily Pemphredoninae The subfamily Pemphredoninae is the most numerous group of sphecids in the fossil record at various ages: the genus Passaloecus Shuckard, 1838 includes Miocene P.scudderi Cockerell 1906, Pfasciatus Rohwer, 1909, and Pmunax Sorg, 1986, Eocene P.microceras Sorg, 1986, and Middle Eocene P.zherichini Budrys, 1993, P.piletskisi Budrys, 1993, and P.electrobius Budrys, 1993; the genus Eoxyloecus Budrys, 1993 — Middle Eocene E.albipalpis Budrys, 1993, E.palionisi Budrys, 1993, E.setipes Budrys, 1993, and E.succinicola Budrys, 1993; the genus Eopinoecus Budrys, 1993 — also Middle Eocene E.samogiticus Budrys, 1993 and E.truncifrons Budrys, 1993. The monotypic Middle Eocene genera Succinoecus Budrys, 1993, Palanga Budrys, 1993, and Eomimesa Budrys, 1993 include S.lituanicus Budrys, 1993, Psucinea Budrys, 1993, and E.rasnitsyni Budrys, 1993. Only three of the described species belong to the Upper Creta- ceous: Lisponema singularis Evans, 1969 (Cedar Lake amber), Pittoecus pauper Evans, 1973, and Cretoecus spinicoxa Budrys, 1993 (last two species from Taimyr amber). The majority of the described genera may be undoubtedly attributed to the subtribe Pemphredonina of the Pemphredonini (sensu Bohart and Menke, 1976), excluding Eomimesa (Psenini, Psenina). The positions of the remaining genera are not as clear. For example, the status of the genus Palanga (lacking a prominent between the antennae clypeal medial lobe, but having a complete occipital carina, ecarinate pronotal collar, long adlateral lines, areolate episternal sulcus, complete forewing venation, and developed pygidial plate) with respect to Spilomena Shuckard, 1838 and Arpactophilus Smith, 1863 is of some doubt. Furthermore, though Palanga has 5- segmented maxillary palpi (judging from the original description, but 6-segmented judging from the personal communication of Dr. David Grimaldi — American Museum of Natural History, New York, USA), these are not shortened as in Spilomenina (sensu Menke, 1989). Similar difficulties exist for the genus Eopinoecus and, to an even 72 greater degree, for Lisponema. If the first really resembles Passaloecus, differing only in the absence of the forewing recurrent vein II (may be a result of specialization, that was not kept by recent representatives of the group), then the truncate forewing marginal cell and the weakly dentate tarsal claws of Lisponema isolate this genus from all Pemphredoninae, excluding extremely specialized Ammoplanini. Unfortunately, the holotype of Lisponemais headless and its mesopleura are not described. Additional investigations are necessary for ascertaining the relationships of Palanga and Lisponema with recent groups of Pemphredoninae. Tribe PEMPHREDONINI 1. Genus PROLEMISTUS nov. ETYMOLOGY. The generic name is derived from the Latin prefix pro-, meaning ‘before’, and the generic name Polemistus. It is masculine. TYPE SPECIES. only species. Prolemistus apiformis sp. nov., here designated; the DIAGNOSIS. Clypeus (partly visible) short; lower frons consider- ably convex, with distinct anteroventral edge covering antennal bases; inner eye orbits almost parallel; ocelli simple, placing in obtuse-angled triangle; occipital carina interrupted, not reaching hypostomal carina; mandible without inner teeth and ventral notch or prominence, with two apical teeth (lower tooth longer than upper one); palpal formula 6—4; female antenna 12-segmented, with all segments longer than thick. Pronotum with short collar, bearing transverse carina; scutum with hardly visible short premedial lines and short traces of adlateral lines; episternal sulcus distinctly ex- pressed (ventral ends invisible); scrobal sulcus distinct; omaulus, subomaulus, acetabular carina, sternaulus, and hypersternaulus absent; precoxal carina developed laterally and ventrally, but not continuing on mesopleuron; midcoxae contiguous; fore- and midtrochanters longer than half of corresponding femora; fore- and midfemora bulbous ventrally, hindfemur triangularly broadened anteriorly in basal 1/3; midtibia bulbous apically, with one apical spur; foretarsal rake and plantulae absent, tarsal claws simple, arolium distinct. Forewing marginal cell long, with acute apex reaching wing outer angle, at least twice as long as not enlarged stigma; two submarginal cells: submarginal cell I hardly shorter than marginal cell, submarginal cell Il approximately thrice as short as marginal cell; two discoidal cells: discoidal cell I hardly longer than stigma, discoidal cell II slightly shorter than submarginal cell I; recurrent vein I received by middle of submarginal cell I, recurrent vein II received by proximal 1/3 of submarginal cell II; forewing media diverging beyond cu-a. Hindwing (hardly visible) with short, rounded jugal lobe, not longer than half of submedial cell; hindwing media diverging before cu-a. Propodeum short, with appressed, but not margined dorsal enclosure, with distinct lateral carinae. First ab- dominal segment with short (shorter that wide) petiole, consisting of sternite I; tergite VI rounded, without pygidial plate or medial carina. COMPARISONS. Single midtibial spur, short abdominal petiole con- sisting of sternite I, and simple tarsal claws allow placement of this specimen into the subfamily Pemphredoninae, and the characteristic forewing venation to the tribe Pemphredonini. Among described fossil pemphredonine genera, Prolemistus gen. nov. resembles Succinoecus in the absence of the hypersternaulus, but differs from it by the apically bidentate mandibles, distinct transverse carinate pronotal collar, simple and thin scrobal and episternal sulci, and by the lack of even a trace of the hypersternaulus. Among recent A.V. ANTROPOV pemphredonine genera Prolemistus gen. nov. most closely resembles Passaloecus and Polemistus Saussure, 1892 by having an uninter- rupted episternal sulcus and by lacking the pygidial plate and outer hindtibial spines. The new genus differs from Passaloecus in the absence of the omaulus, and from Polemistus by the wide head with practically parallel inner eye orbits and the elongate flagellomeres. Prolemistus gen. nov. also differs from both genera by the absence of the hypersternaulus, and from all mentioned genera by the ventrally interrupted occipital carina with the prominent roundly triangular ventral angles, distinctly prominent lower frons overhanging the antennal bases, and by the obtusely broadened hindfemora. Prolemistus apiformis sp. nov. Figs 32-34 ETYMOLOGY. Species name is derived from the Latin masculine adjective apiformis, meaning “bee-shaped’; the name is a reference to the shape of the female body. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.20181; completely preserved female speci- men ina polished irregularly oval sample of Burmese amber (probably Upper Cretaceous). DESCRIPTION. Female. Head considerably widened and thickened; clypeus invisible, but probably short; malar space shorter than forebasitarsal thickness; frons strongly convex, overhanging antennal bases; eyes convex, with inner orbits parallel or slightly converging below; ocelli placing in obtuse-angled triangle, medial ocellus smaller than lateral one; oculo-ocellar distance strongly longer than that between lateral ocelli; occipital carina interrupted ventrally, with prominent lower angles, placing from hypostomal carina at distance, distinctly longer than mandibular base width; mandible widened in middle, with two apical teeth (outer tooth longer than inner one); middle flagellomeres 1.2—1.5 times as long as their maximum thick- ness, without prominences. Pronotal collar with almost right lateral angles, distinctly margined anteriorly and laterally, approximately 1.5 times longer than antennal thickness; scutum flat-convex; ed ‘S oe « “| ae lla: Fig. 32 Prolemistus apiformis gen. et sp. nov, holotype, In.20181, female, Burmese amber. Length 5.1 mm. DIGGER WASPS IN BURMESE AMBER 34 b Figs 33,34 Prolemistus apiformis gen. et sp. nov, holotype, In.20181, female, Burmese amber. 33, Total view (a — dorsolaterally, b — ventrolaterally). 34, Left wings (a — forewing, b — basal fragment of hindwing). Scale bars = 1 mm. scutellum flat; mesopleuron moderately convex; postscutellum vis- ibly convex. Tarsomeres unmodified, without plantulae; apical tarsomere with small arolium. Propodeum short, with dorsal side somewhat concave and shorter than hind side; lateral carinae dis- tinct. Body sculpture weak, hardly visible on head; scutum, mesopleura, and propodeal lateral sides not sculptured, weakly shiny; scutellum dull. Body pubescence weak, very short, mainly appressed with simple light hairs, expressed mainly on temples and abdominal tergites. 73 Body uniformly coloured, black or dark-brown, except somewhat lighter palpi. Body length 5.1 mm. Male unknown. REMARKS. Though the holotype specimen is completely preserved, it is strongly deformed owing to compression. As a result, it appears from some angles that the scutum bears adlateral lines and the left basitarsus is flattened. In fact, however, it concerns only the left side, where the midcoxa, midfemur, mid basitarsus are also, if not more strongly so, compressed laterally and clasped to the flattened abdo- men together with the left hindtarsus. The right mid- and hindtarsomeres are not flattened. 2. Genus CRETOSPILOMENA nov. ETYMOLOGY. The generic name is derived from the Latin mascu- line adjective cretaceus, indicating the age of the specimens, and the generic name Spilomena. It is feminine. TYPE SPECIES. Cretospilomena familiaris sp. nov., here desig- nated, the only species. DIAGNOSIS. Clypeus (partly visible) short, distinctly roundly con- vex (visible in paratypes 3 and 7); frons flat; inner eye orbits almost parallel; lateral ocelli simple (visible in holotype); occipital carina expressed at least dorsally and laterally (visible in holotype and paratypes 4 and 5); mandible without inner teeth and ventral notch or prominence, bidentate apically (visible in paratype 7); palpal for- mula 6-4, palpal segments very short (visible in paratype 3); female antenna 12-segmented, all articles longer than maximum thick. Pronotal collar elongate, with transverse carina; scutum with hardly visible short adlateral lines; mesopleuron with expressed episternal sulcus (visible in holotype and paratype 4); scrobal sulcus, sternaulus, hypersternaulus, omaulus, subomaulus, and acetabular carina not expressed; midcoxae contiguous; legs unmodified, tibiae without outer spines; midtibia with one apical spur; tarsomeres with short apical spines; ventral plantulae and foretarsal rake absent; tarsal claws simple, arolium hardly visible. Forewing marginal cell consid- erably wide, with acute apex touching wing margin, approximately 1.5 times as long as distinctly enlarged stigma; two submarginal cells: submarginal cell I hardly shorter, submarginal cell II approxi- mately thrice shorter than marginal cell; single discoidal cell 1.5 times longer than stigma; recurrent vein received by submarginal cell I; forewing media diverging beyond cu-a. Hindwing media diverging at cu-a, RS practically straight and equal to r-m (visible in paratype 1); hindwing with 5 long hamuli. Propodeal dorsal enclo- sure flattened and weakly margined laterally, bearing a pair of premedial carenulae; propodeum with strong sharp posterolateral thorns, joining posteriorly by transverse carina; propodeal lateral carinae not expressed distinctly. First abdominal segment with elon- gate (slightly longer than wide) petiole, consisting of tergite I and sternite I; tergite VI rounded, with short apicomedial carina (visible in paratype 1), without pygidial plate. COMPARISONS. The single midtibial apical spur, the simple tarsal claws, the mesopleuron without sulci (except the episternal sulcus), the reduced forewing venation, the enlarged stigma, and the short abdominal petiole consisting of tergite I and sternite I allow the placement of Cretospilomena gen. nov. into the tribe Pemphredonini of the subfamily Pemphredoninae. It is difficult to place this genus into any of the subtribes of Pemphredonini that have an unmodified marginal, two submarginal, and single discoidal cells (Menke, 1989), because the specimen combines features of both Stigmina (6-seg- mented maxillary palpi) and Spilomenina (shortened palpal segments 74 and a short abdominal petiole consisting of both tergite I and sternite I). In outward appearance Cretospilomena gen. nov. most closely resembles Microstigmus Docke, 1907 while the forewings resembles those of Spilomena. In the main features of the forewing venation (distinctly enlarged stigma, not reduced marginal, two submarginal, and single discoidal cells) the genus is similar to Lisponema and Eopinoecus among fossil genera, and to Spilomena among recent genera. Lisponema differs from Cretospilomena gen. nov. by the truncate marginal cell with veins 3r and RS-a, by the more strongly developed adlateral lines, deep and prolongly carinate prescutellar furrow, weakly dentate (judging from the original description) tarsal claws, weak posterolateral propodeal angles, and by the non-petiolate abdomen. Eopinoecus, also having the palpal formula 6—4, differs from Cretospilomena gen. nov. by the supraantennal frontal tuber- cle, apically truncate and delimited by lateral teeth medial clypeal lobe, apically tridentate mandibles, distinctly areolate scrobal and episternal sulci bearing a trace of the hypersternaulus, weakly en- larged stigma, hindwing media diverging before cu-a, densely areolate propodeum without posterolateral thorns, and by the flat and short pygidial plate. Both Lisponema and Eopinoecus also differ from Cretospilomena gen. nov. by the following features: the narrow pronotal collar and the group of 6 hamuli on the hindwing anterior margin. Spilomena differs from Cretospilomena gen. nov. by the absence of the occipital carina and sharp posterolateral propodeal thorns, palpal formula 5-4, and the non-petiolate abdomen. Figs 35-45 ETYMOLOGY. Species name is derived from the Latin feminine adjective familiaris, meaning ‘family’; the name is a reference to the probable collective nesting. Cretospilomena familiaris sp. nov. HOLOTYPE. Department of Palaeontology of The Natural History Museum (London), In.19136(1); completely preserved female speci- men (Figs 35, 39), in the narrow part (Fig. 38) of a flat-oviform polished sample of Burmese amber (probably Upper Cretaceous). DESCRIPTION. Female. Head flattened; upper frons weakly con- vex, without expressed medial furrow; inner eye orbits parallel; malar space shorter than antennal thickness; vertex and temples uniformly moderately convex, without angles; mandible unmodi- fied, bidentate apically, outer tooth rather longer than inner one; flagellomeres slightly longer than thick, apical flagellomere dis- tinctly longer than penultimate one. Pronotum distinctly elongate; pronotal collar at least twice as long as scape maximum thickness, with straight transverse carina not reaching pronotal lobes laterally; scutum roundly convex, without visible prolong lines, distinctly arched above level of pronotal collar; prescutellar furrow thick, ecarinate prolongly; scutellum flat, trapeziform; postscutellum equal or hardly longer than scape thickness; mesopleuron uniformly con- vex, with weak, thin, and curved episternal sulcus between pronotal lobe and forecoxal base; legs unmodified, slender, without foretarsal rake and tibial spines; plantulae absent; tarsal claws simple, slightly widened basally; hindwing anterior margin with distal group of 5 rather long and weakly curved hamuli (hindwing bases invisible). Propodeal dorsal enclosure flat, with a pair of straight premedial carinae not reaching its apex, distinctly separated from hind side by straight transverse carina ending laterally with sharp thorns. First abdominal segment narrowed anteriorly forming a petiole consisting of both tergite I and sternite I; tergite VI with distinct medial carina, but without long erect bristles. Head dull, with vague sculpture; scutum uniformly and delicately A.V. ANTROPOV Fig. 35 Cretospilomena familiaris gen. et sp. nov., holotype, In.19136(1), female, Burmese amber. Length 2.3 mm. punctate, with shiny interspaces; abdomen without distinct sculp- ture, weakly shiny. Body pubescence very weak and short, hardly visible. Head and thorax black; mandibles (except dark apices), palpi, antennae, abdomen, and legs from brownish-red to red. Body length 2.3 mm. PARATYPES. Eight females in various states of preservation, all in the same piece of amber as the holotype (In.19136): Paratype 1 (Fig. 40), In.19136 (2). Head anteriorly (including Figs 36,37 Cretospilomena familiaris gen. et sp. nov., females. 36, Paratype 7, In.19136(8), total view dorsolaterally. 37, Paratype 8, In.19136(9), total view laterally. Scale bars = 1 mm. DIGGER WASPS IN BURMESE AMBER 75 Figs 38-45. Cretospilomena familiaris gen. et sp. nov., females. 38, Position of types in Burmese amber sample In.19136. 39, Holotype, In.19136(1), total view laterally. 40, Paratype 1, In.19136(2), (a — total view laterally, b — left forewing, c — fragment of right forewing, d— fragment of right hindwing). 41, Paratype 2, In.19136(3), total view ventrolaterally. 42, Paratype 3, In.19136(4), total view ventrolaterally. 43, Paratype 4, In.19136(4), total view dorsally. 44, Paratype 5, In.19136(6), total view dorsally. 45, Paratype 6, In.19136(7), total view dorsolaterally. Scale bars = 1 mm. 76 frons, eyes, and clypeus) and thorax laterally considerably destroyed; both fore- and right hindwing basally, distal parts of right fore- and hindwings, and also left hindwing completely absent; abdominal tergites I and II flattened laterally. Body length 2.6 mm. Paratype 2 (Fig. 41), In.19136 (3). Left antenna, right hindwing, and right forewing basally absent; left pair of wings and mesopleural structure invisible; legs and abdomen partly covered with foreign particles. Body length 2.3 mm. Paratype 3 (Fig. 42), In.19136 (4). Right pair of wings folded prolongly; structure of thorax laterally and of propodeum invisible; tarsi, mesopleura, and partly right antenna covered with foreign particles. Body length 2.4 mm. Paratype 4 (Fig. 43), In.19136 (5). Following parts visibly de- formed: antennae (articles separated), pronotal anterior part (unusually elongate), and wings (anterior veins curved up to rupture of right stigma, vein between Sc+R and RS, and couplings between wing, tegula, and thorax); only anterior part of right hindwing with a row of 5 hamuli visible; head dorsally covered with foreign particles. Body length 2.7 mm. Paratype 5 (Fig. 44), In. 19136 (6). Mesothorax together with right pair of wings and propodeum dorsally almost completely, prothorax dorsally from right side, and left pair of wings mainly destroyed during polishing the sample; abdominal tergite II with large concavi- ties, mainly anteriorly. Body length 2.5 mm. Paratype 6 (Fig. 45), In. 19136 (7). Both hindwings and forewings distal membranes invisible; thorax laterally including legs hidden under well preserved specimen of Scelionidae; abdominal tergite I and apex covered with foreign particles; head and abdominal tergites I-III strongly concave dorsally. Body length 2.4 mm. Paratype 7 (Fig. 36), In.19136 (8). Mesopleural structure, hind- wings, right forewing distal membrane, and the main part of veins of right forewing invisible; head anteriorly covered with foreign parti- cle; abdominal tergite I apically and tergite II basally destroyed. Body length 2.5 mm. Paratype 8 (Fig. 37), In.19136 (9). Structure of frons, mesopleura, lateral sides of abdomen, and forewings distal membranes invisible; right forewing submarginal cell II absent; only anterior part of left hindwing with a row of 5 hamuli visible; left antenna apically, abdomen ventrally in the middle, and hindfemora partly covered with foreign particles. Body length 2.2 mm. The male is unknown. REMARKS. This is the only known case of the simultaneous preser- vation of a large group of sphecid wasps belonging to one species in a fossil resin. I propose that this fact may suggest the compact nesting of several females in a small biotope, as is observed among some recent xylobiotic species of the genus Spilomena, and was probably typical for C.familiaris sp. nov., if it was not an eusocial mode of life, similar to that of the genera Microstigmus (Matthews, 1968a, 1968b) or Arpactophilus (Matthews & Naumann, 1988). DISCUSSION. The multiplicity of the representatives of the tribe Pemphredonini discovered in fossil resins of various ages, including Upper Cretaceous amber, testifies to the earlier isolation of this specialized group of sphecid wasps and to the establishment of its close association with arboreal plants. A comparative analysis of fossils of various ages demonstrates that the Upper Cretaceous representatives of the tribe already had the characteristic outward appearance, including the wing venation (at least of the forewings), and were characterized by a set of specialized features. These include the apically truncate marginal cell (Lisponema), the strongly convex lower frons covering the antennal bases from above (Prolemistus), the sharp posterolateral thorns of 1 the propodeum (Cretospilomena), the more or less well developed A.V. ANTROPOV teeth of the tarsal claws (Pittoecus, Lisponema), and the shortened 6- segmented maxillary palpi (Cretospilomena). If the last two features are archaic, then the truncate marginal cell, the strongly convex lower frons, and the strongly developed propodeal thorns are not present in both later and recent fossil Pemphredonini. They should be recognized as lost specializations which were spe- cific to genera. The absent or weakly developed prolonged mesopleural sulci (especially the hypersternaulus) and the non- areolate episternal sulcus in the genera Cretoceus, Pittoecus, and Prolemistus, should probably also be considered plesiomorphic. The simultaneous fossilization of many females of C.familiaris sp. nov. is of special interest. I think, this unique case may show the early evolution of sociality in Pemphredonini. CONCLUSIONS Study of the present material suggests that the evolution of the Upper Cretaceous Sphecidae, in contrast to the recent representa- tives of the family, had specific pecularities. As well as such generalized features as the completely developed adlateral lines and two apical midtibial spurs (the shortened hindwing RS in the majority of the studied forms is probably also a generalized fea- ture), they have acquired some apomorphies, that are not present in recent sphecid wasps. The unique structure of the scape and the wing venation of Burmastatus, the pectinate antennae and the ve- nation of the forewing submarginal sector of Cirrosphex, the spherical head, the reduced forewing venation, and the edentate tarsal claws of Apodolichurus, the reduced discoidal cell II of Trigampulex, as well as, probably, the ventrally angulate mandi- bles of Burmastatus, the small eyes, complete occipital carina, and specialized mandibles of Mendampulex, the apically bidentate mandibles of Cretampulex, the strongly convex lower frons of Prolemistus, and the dentate laterally propodeum of Creto- spilomena should be placed into the group of such features. Differences such as general shift of the forewing veins and cells to the wing proximal half and the diminution of the marginal cell in Burmastatus, the diminution of the forewing submarginal cell III with simultaneous enlarging of the marginal cell in Cirrosphex are of interest, because they do not conform to the general directions of the venational reductions that are characteristic for recent representa- tives of the family (truncation or separation of the forewing marginal cell apex with its diminution, reduction of the forewing submarginal sector mainly by the diminution of the submarginal cell II or by junction of the submarginal cells I-II or II-III). It is apparent that the Upper Cretaceous fauna of sphecid wasps was sufficiently variable and differed considerably from the recent representatives of the family by possessing many specialized features, including a different evolutionary development of the wing venation. All mentioned pecularities of the discovered taxa place them far from the hypothetical ancestor of recent Sphecidae. ACKNOWLEDGEMENTS. [am grateful to A.J. Ross and Prof. A.P. Rasnitsyn for giving me an opportunity to study the samples and for useful discussions of some phylogenetic problems, to Prof. Charles D. Michener, Dr. Robert W.Brooks (Snow Entomological Museum, Lawrence, Kansas, USA), and Dr. Gabriel A. R. de Melo (Universidade Federal de Vicosa, Vigosa MG, Brasil) for their valuable consultations on the wing venation of bees and pemphredonine wasps, to Dr. Max Fischer (Naturhistorisches Museum Wien, 2 Zoologishe Abteilung, Austria) for lending me the comparative specimens of Xenosphecinae and to Peter York (NHM) for photography. DIGGER WASPS IN BURMESE AMBER REFERENCES Antropov A.V. 1995. A new species of the genus Trypoxylon Latreille (Hymenoptera, Sphecidae) from Dominican amber. Paleontologicheskiy zhurnal, 1995(1): 125-128. [In Russian] — & Pulawski W.J. 1989. 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Goulet, H. & Huber J.T. (editors).1993. Hymenoptera of the world: an identification guide to families. Agriculture Canada Publication 1894/E. Ottawa, Canada. vii + 668 pp. Matthews R.W. 1968a. Microstigmus comes: sociality in a sphecid wasp. Science Hil (Washington), 160: 787-788. —— 1968b. Nesting biology of the social wasp Microstigmus comes (Hymenoptera: Sphecidae, Pemphredoninae). Psyche, 75: 23-45. & Naumann I.D. 1988. Nesting biology and taxonomy of Arpactophilus mini, a new species of social sphecid (Hymenoptera: Sphecidae) from Northern Australia. Australian Journal of Zoology, 36 (5): 585-597. Menke A.S. 1989. Arpactophilus reassessed, with three bizarre new species from New Guinea (Hymenoptera: Sphecidae: Pemphredoninae). Invertebrate Taxonomy, 2: 737-747. — & Fernandez F.C. 1996. Claces ilustradas para las subfamilias, tribus y géneros de esfécidos neotropicales (Apoidea: Sphecidae). Revista Biologia Tropical, 44 (2): 1-68. — & Rasnitsyn A.P. 1987. Affinities of the fossil wasp, Hoplisidea kohliana Cockerell (Hymenoptera: Sphecidae: Sphecinae). Psyche, 94 (1-2): 35-38. Nemkov P.G. 1988. New genus of digger wasps from Baltic Amber. Paleontologicheskiy zhurnal, 1988(2): 119-121. [In Russian] 1990. A new species of sphecid wasps of the tribe Gorytini from Oligocene deposits of Primorskiy Kray. Paleontologicheskiy zhurnal, 1990(4): 123-125. [In Russian] Prentice M.A. & Poinar G.O. Jr. 1993. Three species of Trypoxylon Latreille from Dominican Amber (Hymenoptera, Sphecidae). Journal of Kansas Entomological Society, 66 (3): 280-291. Pulawski W.J. & Rasnitsyn A.P. 1980. On the taxonomic position of Hoplisus sepultus Cockerell, 1906, from the Lower Oligocene of Colorado (Hymenoptera, Sphecidae). Polskie Pismo Entomologizne, 50: 393-396. Rasnitsyn A.P. 1975. Hymenoptera Apocrita of Mesozoic. Trudy Paleontologicheskogo Instituta, 147: 1-133. [In Russian] 1980. Origin and evolution of hymenopterous insects. Trudy Paleontologicheskogo Instituta, 174: 1-192. [In Russian] , Pulawski W.J. & Martinez-Declos X. 1999. Cretaceous digger wasps of the new genus Bestiola Pulawski and Rasnitsyn (Hymenoptera: Sphecidae: Angarosphecinae). Journal of Hymenoptera Research, 8 (1): 23-34. Rohwer S.A. 1908. A fossil larrid wasp. Bulletin of the American Museum of natural History, 24: 519-520. — 1909. New Hymenoptera from the western United States. Transactions of the American Entomological Society, 35 (1): 99-136. Sorg M. 1986. Grabwespen der Gattung Passaloecus aus fossilen Harzen (Hymenop- tera, Sphecoidea, Pemphredoninae). Passaloecus microceras n. sp., Baltisches Bernstein, oberes Eozan. Passaloecus munax n. sp., Bitterfelder Bernstein, unteres Miozan. Paldontologische Zeitung, 60 (3/4): 277-284. Timon-David J. 1944. Insectes fossiles de |’ Oligocéne inférieur des Camoins (Bassin de Marseille). II. Hyménoptéres. 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(Geol.) 56(1): 79-83 Issued 29 June 2000 Electrobisium acutum Cockerell, a cheiridiid pseudoscorpion from Burmese amber, with remarks on the validity of the Cheiridioidea (Arachnida, Chelonethi) M.L.I. JUDSON Muséum national d’ Histoire naturelle, Laboratoire de Zoologie (Arthropodes), 61 rue de Buffon, F-75231 Paris Cedex 05, France SYNOPSIS. Electrobisium acutum Cockerell is redescribed from a specimen cut from the block of Burmese amber containing the holotype. The presence of strong spines on the carapace and tergites indicates that E. acutum may be closely related to extant South African or Taiwanese species of the genus Cryptocheiridium Chamberlin. Electrobisium and Cryptocheiridium are not synonymized, however, due to insufficient knowledge of E. acutum (the type species of Electrobisium) and problems with the definition of Cryptocheiridium. The superfamily Cheiridioidea, containing the families Cheiridiidae and Pseudochiridiidae, is removed from synonymy with the Garypoidea and regarded as the sister group of the Cheliferoidea. INTRODUCTION Only two pseudoscorpion species have been reported from Burmese amber: Electrobisium acutum Cockerell, 1917 and Garypus burmiticus Cockerell, 1920. The original descriptions (Cockerell, 1917, 1920) allow almost nothing to be said about the systematic position of these species, with the result that they have rarely been considered by subsequent authors. The types were re-examined by Judson (1997), who tentatively assigned Garypus burmiticus to the recent genus Amblyolpium Simon (family Olpiidae) and placed Electrobisium Cockerell in the family Cheiridiidae. The Burmese amber pseudoscorpions in the collections of the Natural History Museum are difficult to study because of the thick- ness of the amber and the abundance of other inclusions. Although the blocks have been cut into slabs, the pseudoscorpions are usually too far from the surface to be examined adequately. However, a small piece containing a specimen of EF. acutum has been cut from one of the slabs. Despite being distorted and partly obscured by debris, this specimen can be examined in sufficient detail to show that it belongs to the Cheiridiidae and that it is probably closely related to Recent species of the genus Cryptocheiridium Chamberlin. The following description is based on this non-type specimen, which was not mentioned by Cockerell. There can be little doubt that it is conspecific with the holotype, since the latter also has carapacial and tergal spines and came from the same block. SYSTEMATIC DESCRIPTIONS Order CHELONETHI Thorell, 1883 Superfamily CHEIRIDIOIDEA Hansen, 1894 DISCUSSION. The superfamily Cheiridioidea has traditionally included the Cheiridiidae, Pseudochiridiidae (sometimes consid- ered a subfamily of Cheiridiidae) and Sternophoridae, although Chamberlin (1931: 234—235) noted that ‘in some ways this group of three rare families is an unnatural one’. In a cladistic analysis of the © The Natural History Museum, 2000 pseudoscorpion families, Harvey (1992) found the Sternophoridae to be the sister group of the Cheliferoidea, while the Cheiridiidae and Pseudochiridiidae formed the monophyletic sister group of the clade Garypidae + Larcidae. This led Harvey to erect the monotypic superfamily Sternophoroidea and synonymize the Cheiridioidea with the Garypoidea. Unfortunately, Harvey’s analysis was complicated by the fact that his unweighted data support a sister-group relationship between the Cheiridioidea and Feaelloidea. Harvey rejected this result and in- stead imposed a sister-group relationship between the Feaelloidea and Chthonioidea by a posteriori weighting of a single character (presence of trichobothria ds). If, as this weighted solution implies, the characters that originally placed the Feaelloidea within the Garypoidea are misleading, should they be regarded as any more reliable for determining the relationships of the Cheiridioidea? There are nine putative synapomorphies uniting the Cheiridioidea (Cheiridiidae and Pseudochiridiidae) with one or more clades of the Elassommatina in Harvey’s preferred cladogram. It could be argued that these represent parallelisms in the case of the Feaelloidea and Garypoidea (s./.), and that it would be even less parsimonious to treat them as such in Cheiridioidea and the remaining Garypoidea. How- ever, I suggest that these characters have been misinterpreted as synapomorphies. Harvey’s (1992) characters 52 (ocular tubercle present) and 85 (arolia longer than claws) can be ignored because the Cheiridioidea show the plesiomorphic states (interpreted as reversals by Harvey). Characters 51 (carapace sub-triangular) and 123 (body ovoid) are evidently correlated: an oval body shape entails a broadening of the carapace (the Feaellidae are exceptional in this respect, due to their unique prosomal modifications). Even if a broad body were accepted as apomorphic, the subjectivity of this character makes it difficult to code in many cases. In Pycnocheiridium Beier, for example, the shape of the body (including carapace and coxae) is similar to that found in most Cheliferoidea. A broadened body is also usually correlated with a broadening of the posterior coxae (character 66), though the Feaellidae and Geogarypidae provide exceptions. The ‘garypoid’ facies of the Pseudogarypidae (presumably part of the ground-plan for Feaelloidea) make the polarity of coxal shape ambiguous. Characters 49 (anterior margin sinuous) and 53 (eyes 80 removed from anterior margin) are also subjective, being correlated with the presence of a ‘cucullus’ (With, 1906). It is often difficult to decide how intermediate forms should be scored for these characters, which again reflect carapace shape. Character 125 (setae curved) is probably more general than Harvey implies. Many Cheliferoidea have strongly curved setae, and truly straight setae are not found in any pseudoscorpions (except when they are secondarily thickened and spine-like). Character 120 (sternite XI enclosing anus) is particularly interest- ing. Chamberlin (1931) and Harvey (1992) believed that the primitive state in pseudoscorpions was to have segment XI divided into a tergite and sternite, which became secondarily fused in some groups. Because this segment is undivided in most families (including the Chthonioidea and Feaelloidea), it is more parsimonious to assume that the reverse is true. Segment XI is only divided into a separate tergite and sternite in Geogarypidae, Garypidae, Cheiridioidea and Cheliferoideae. The ventral displacement of the anal segment (XII) has subsequently led to its becoming surrounded by sternite XI in the Garypidae, Pseudochiridiidae and Cheiridiidae (except in Pycnocheiridiinae and Apocheiridium). The distribution of this character suggests that it has arisen independently in all three families. It therefore appears that most of the characters used by Harvey (1992) to place the Cheiridioidea within the Garypoidea are either miscoded or ambiguous. The traditional arrangement of a sister group relationship between the Cheiridioidea and Cheliferoidea is preferred here. Potential synapomorphies for this clade (Monosphyronida minus Sternophoridae) include the complete fu- sion of the basi- and telotarsi, the presence of dentate to clavate vestitural setae, the loss of the posterior pair of eyes and the loss of reflective tapeta from the anterior pair. An additional synapomorphy uniting these groups may be the presence of a fibrous envelope around the flagellar tunnel of the spermatozoa (Callaini & Dallai, 1994), but no information is available yet about the spermatozoa of Pseudochiridiidae and Sternophoridae. Family CHEIRIDIIDAE Hansen, 1894 Genus ELECTROBISIUM Cockerell, 1917 TYPE SPECIES. designation). Electrobisium acutum Cockerell, 1917 (by original DISTRIBUTION. Burmese amber, Hukawng Valley, Myanmar (Burma); probably Upper Cretaceous (see Zherikhin & Ross, 2000). DIAGNOSIS. Medium-sized Cheiridiidae with strong spines on pos- terior margins of carapace and tergites I-VII. Palp femur apparently longer than carapace and probably angled posteriorly at proximal end. Chela with seven trichobothria on fixed finger and two trichobothria on movable finger; trichobothria est and t near middle of fingers. DISCUSSION. Cockerell’s (1917) assignment of Electrobisium to the Obisiidae (=Neobisiidae) is puzzling because it is not supported by any of the characters mentioned in the original description. Cockerell simply indicated that E. acutum was ‘quite unlike the pseudoscorpions described from Baltic amber, though there is a slight superficial resemblance to Chelifer ehrenbergii’. Chelifer ehrenbergii C. L. Koch & Berendt is a nomen dubium (Harvey, 1991), but it probably belongs in the Cheliferoidea. The fact that Koch & Berendt’s (1854) figure of C. ehrenbergii appears next to that of Obisium rathkii C. L. Koch & Berendt (now Neobisium M.L.I. JUDSON rathkii) — at that time the only fossil neobisiid known — suggests that Cockerell may have confused Chelifer ehrenbergii with Obisium rathkii. Schawaller (1978) noted that the original description and figures of E. acutum were inadequate, but listed Electrobisium in the Neobisiidae, as did Harvey (1991). Electrobisium was transferred to the Cheiridiidae by Judson (1997), without detailed comment. This position is supported by the general appearance of E. acutum, the fusion of the femora and patellae of the legs, and the reduced number of chelal trichobothria. The presence of well developed spines on the posterior margin of the carapace and the anterior tergites (hereafter referred to simply as the ‘dorsal spines’) also suggests that Electrobisium might be related to extant species of the genus Cryptocheiridium Chamberlin, 1931. Unfortunately, it is not clear whether Cryptocheiridium, as cur- rently defined, is a monophyletic group. This genus was erected by Chamberlin (1931) for two species: Cheiridium subtropicum Tullgren (the type species), from South Africa, and C. formosanum Ellingsen, from Taiwan. Chamberlin did not see material of either species and merely distinguished the genus in a key, based on the presence of dorsal spines and the fact that 11 tergites are visible in dorsal view. Beier (1932) provided a more detailed diagnosis of the genus, which has since been enlarged to accommodate extant species from central and eastern Africa, Southeast Asia and Australia (Harvey, 1991), as well as a fossil species from Dominican amber (Schawaller, 1981). None of these species show the strong spines present in C. subtropicum and C. formosanum and their assignment to Cryptocheiridium has mainly been based on the robustness of the palps and the number of trichobothria on the movable finger. The unreliability of the number of trichobothria as a generic character is shown by the existence of an undescribed South African species of Cryptocheiridium which has strong dorsal spines and only a single trichobothrium on the movable finger (pers. obs.). Because this species is evidently closely related to C. subtropicum, it becomes difficult to separate Cryptocheiridium from Neocheiridium Beier. Fig. 1 Electrobisium acutum. Holotype, dorsal view of adult, In.19118, Burmese amber. Length of body 0.9mm. CHEIRIDIID PSEUDOSCORPION FROM BURMESE AMBER 81 Figs 2,3 Electrobisium acutum. In.19123(3), Burmese amber. 2, Dorsal view of adult. Surface granulation only shown in part. Size and position of eyes uncertain; other doubtful or reconstructed parts shown by dotted lines. Femoropatella of left leg II broken; opisthosoma incomplete and slightly foreshortened. Diagonal line represents edge of amber. Scale line 0.5 mm. 3, Chela of right palp, showing trichobothriotaxy (reconstruction, based on left and right chelae; not to scale). The possibility exists, therefore, that the species currently as- signed to Cryptocheiridium represent a heterogeneous assemblage. This would not be important in the present context, were it not for the possibility that even the species with strong dorsal spines might not be closely related. The vestitural setae of C. formosanum are covered by an exudate that gives them a leaf-like appearance, as in some species of Neocheiridium (Mahnert & Aguiar, 1986), whereas those of the South African species lack any covering. C. formosanum is also unusual in having a large tubercle on each side of the carapace (Ellingsen, 1912) anda long flange on the anterolateral margin of the palp coxa (pers. obs.). Given these differences, the possibility that the dorsal spines have arisen more than once has to be considered. This is not difficult to imagine, since they are evidently formed by the elongation of the normal granules found along the posterior margins of the carapace and tergites of most Cheiridiidae. The difference between granules and spines is simply one of degree. This is clearly shown by the ontogeny of the South African Cryptocheiridium, in which the spines are only fully formed in the adult. Because of these doubts concerning the monophyly of Cryptocheiridium, or even of aclade containing the spined forms, it is difficuit to identify relationships between Electrobisium and Recent species. This is compounded by the fact that it is not possible to determine with certainty whether the unusual characters seen in C. formosanumare present or absent in E. acutum. All that can be said for the moment is that E. acutum is more similar to C. formosanum in having a lower number of spines (12-14 per tergite, versus 18—27 in the South African species). It is, of course, possible that E. acutum is notclosely related to either the SouthA frican or the Taiwanese species. The differences in the shape of the palps and their trichobothriotaxy might be significant, but the distortion of the fossils means that caution is required in interpreting these characters. In view of these problems, Cryptocheiridium and Electrobisium are retained as separate genera here. From a practical point of view, it would be unfortunate to have E. acutum as the type species of an extant genus. Electrobisium acutum Cockerell, 1917 Figs 14 1917 = Electrobisium acutum Cockerell: 360, fig. 1. 1978 Electrobisium acutum Cockerell; Schawaller: 3. 1980 ~=Electrobisium acutum Cockerell; Morris: 36. 82 1991 = Electrobisium acutum Cockerell; Harvey: 334. 1992 Electrobisium acutum Cockerell; Poinar: 220. 1997 _ Electrobisium acutum Cockerell; Judson: 7. MATERIALEXAMINED. Holotype: NHM Pal. Dept. In.19118, adult, Burmese amber, Hukawng Valley, Myanmar (Burma) (presented by R. C.J. Swinhoe, Feb. 1919). Non-types: | nymph (protonymph?), in same slab as holotype; | adult, off-cut from block containing holotype, NHM Pal. Dept. In.19123(3). The following description is based on the non-type specimen in the off-cut (BMNH In.19123(3)). Because there is a complete frac- ture just below the pseudoscorpion (previously repaired, probably with Canada balsam), no further preparation was attempted. The study was therefore limited to the dorsal view of the fossil, some parts of which (particularly the carapace) were obscured by debris. A small, poorly preserved mite larva (Parasitengona) is also present in the amber fragment, next to the right chela of the pseudoscorpion. DESCRIPTION. General appearance of fossil as shown in Figures 2— 4. However, this is a caricature of the shape of the living animal: the specimen is distorted, with many parts being unnaturally folded or collapsed. The strong folding of the trochanters and femora of the palps makes them appear thinner than normal. The posterior region of the carapace and the anterior tergites have been constricted laterally, such that the carapace has lost its usual subtriangular shape. Some parts, particularly the carapace, are obscured by a layer of debris lying in the same plane as the specimen. The presence of this layer suggests that the specimen was probably exposed on the surface of the resin for some time, before being covered by a second layer. Part of the posterior end of the body has been lost at the fractured edge of the amber. Colour chestnut brown. Setae small (only a few visible on palps and leg tarsi). Carapace with a deep anterior furrow; presence or absence of posterior pit could not be determined; posterior margin Fig.4 Electrobisium acutum, dorsal view of fossil, In.19123(3), Burmese amber. Length of body 0.7mm. M.L.I. JUDSON with about 10 long, sharp spines; eyes apparently small (observation and figure doubtful). At least ten tergites visible dorsally, but end of body missing; tergites I-VI with 10-12 spines (maximum length about 0.025 mm), similar to those of carapace, tergite VII with only 6 small spines, posterior tergites without spines. Palps attenuate; with strong granulation, giving the margins a toothed appearance; posterior margin of femur strongly produced proximally, but this might be an artefact of folding; patella clavate; hand sub-triangular in outline; trichobothriotaxy as shown in Fig. 3, fixed finger with seven trichobothria, movable finger with two trichobothria. Anterior legs moderately robust, posterior legs elongate; femora fused with patellae; claws normal; arolia apparently small. Ventral surfaces not clearly visible. Measurements (in mm); body length >0.7 (about 0.9 in holotype); carapace length ca. 0.26 mm; palp femur 0.29 x >0.06, patella 0.22 x >0.085, chela length 0.37, hand 0.20 x 0.105, fingers 0.18 long. REMARKS. The holotype specimen (identified by comparison with Cockerell’s figure) is distorted in a similar way to the specimen described above, though the abdomen has a more normal (less rounded) shape (Fig. 1). The contraction of the carapace in the holotype has left the trochanter of leg III visible on both sides in dorsal view, which Cockerell incorrectly drew as continuations of the first tergite. The conspecificity of the two specimens is supported by the presence of dorsal spines in the holotype (these are difficult to see and were overlooked by Cockerell). OTHER PSEUDOSCORPIONS IN BURMESE AMBER Other Burmese amber pseudoscorpions were briefly examined dur- ing a visit to the NHM. Most of these fossils are fragmentary, usually representing parts of palps. It is clear that there is a diverse fauna in the amber, including representatives of the Chthonioidea, Garypoidea, Cheiridioidea and Cheliferoidea. ACKNOWLEDGEMENTS. Fossils in the Palaeontology Department of the Natural History Museum, London were examined through the courtesy of Ms J. G. Darrell and Mr A. J. Ross. Material of extant Cryptocheiridium species was kindly loaned by Mrs L. Brown (Transvaal Museum, Pietermaritzburg) and Dr G. E. E. S6li (Zoologisk Museum, Oslo). Helpful comments on the text were provided by Mark Harvey (Western Australian Museum, Perth). Thanks also go to Mr Peter York (NHM) for photography. REFERENCES Beier, M. 1932. Pseudoscorpionidea II. Subord. C. Cheliferinea. Das Tierreich, 58: 1— 294. Callaini, G. & Dallai, R. 1994. The spermatozoon of pseudoscorpions (Arachnida). Bollettino dell’ Accademia Gioenia di Scienze naturali, 26 [1993]: 35-52. Chamberlin, J. C. 1931. The arachnid order Chelonethida. Stanford University Publi- cations, University Series, (Biol. Sci.) 7 (1): 1-284. Cockerell, T. D. A. 1917. Arthropods in Burmese amber. American Journal of Science, (4) 44: 135-138. 1920. Fossil arthropods in the British Museum — I. Annals and Magazine of Natural History, (9) 5: 273-279. Ellingsen, E. 1912. H. Sauter’s Formosa-Ausbeute. Pseudoscorpions from Formosa. I. Nytt Magasin for Naturvidenskapene, 50: 121-128. Harvey, M.S. 1991. Catalogue of the Pseudoscorpionida. 726 pp. Manchester Univer- sity Press, Manchester. 1992. The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy, 6: 1373-1435. CHEIRIDIID PPEUDOSCORPION FROM BURMESE AMBER Judson, M. L. I. 1997. Catalogue of the pseudoscorpion types (Arachnida: Chelonethi) in the Natural History Museum, London. Occasional Papers on systematic Entomol- ogy, 11: 1-54. Koch, C. L. & Berendt, G. C. 1854. Die in Bernstein befindlichen Crustaceen, Myriapoden, Arachniden und Apteren der Vorwelt. In: Berendt, G. C. Die im Bernstein befindlichen Organischen Reste der Vorwelt Gesammelt in Verbindung mit meheren Bearbeitet und Herausgegeben, 1 (2): 1-124, 17 pls (Nicolai, Berlin). Mahnert, V. & Aguiar, N. O. 1986. Wiederbeschreibung von Neocheiridium corticum (Balzan, 1890) und Beschreibung von zwei neuen Arten der Gattung aus Stidamerika (Pseudoscorpiones, Cheiridiidae). Bulletin de la Société entomologique Suisse, 59: 499-509. Morris, S. F. 1980. Catalogue of the type and figured specimens of fossil Crustacea (excl. Ostracoda), Chelicerata, Myriapoda and Pycnogonida in the British Museum (Natu- ral History). British Museum (Natural History), Publication no. 828, 53 pp, 3 pls. 83 Poinar, G. 1992. Life in amber. 350 pp. Stanford University Press, Palo Alto. Schawaller, W. 1978. Neue Pseudoskorpione aus dem Baltischen Bernstein der Stuttgarter Bernstein Sammlung (Arachnida: Pseudoscorpionidea). Stuttgarter Beitrdge zur Naturkunde, (B) 42: 1-22. 1981. Cheiridiidae in Dominikanischem Bernstein, mit Anmerkungen zur morphologischen Variabilitat (Stuttgarter Bernsteinsammlung: Arachnida, Pseudoscorpionidea). Stuttgarter Beitrdge zur Naturkunde, (B) 75: 1-14. With, C. J. 1906. The Danish expedition to Siam 1899-1900. III. Chelonethi. An account of the Indian false-scorpions together with studies on the anatomy and classification of the order. Det Konigelige Danske Videnskabernes Selskabs Skrifter, (7) 3: 1-214, 1 map, pls I-IV. Therikhin, V. V. & Ross, A. J. 2000. The history, geology and age of Burmite (Burmese amber). Bulletin of the Natural History Museum, London, Geology, 56: 3-10. Volume 46 No. 1 No. 2 Volume 47 No. 1 No. 2 Volume 48 No. 1 No. 2 Volume 49 No. 1 No. 2 Volume 50 No. 1 No. 2 Volume 51 No. 1 No. 2 Bulletin of The Natural History Museum Geology Series Earlier Geology Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. Mid-Cretaceous Ammonites of Nigeria—new amphisbaenians from Kenya—English Wealden Equisetales—Faringdon Sponge Gravel Bryozoa. 1990. Pp. 1-152. 0 565 070274. £45.00 Carboniferous pteridosperm frond Neuropteris heterophylla— Tertiary Ostracoda from Tanzania. 1991. Pp. 153-270. 0565 07028 2. £30.00 Neogene crabs from Brunei, Sabah & Sarawak—New pseudosciurids from the English Late Eocene—Upper Palaeozoic Anomalodesmatan Bivalvia. 1991. 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Pp. 1— 107. £43.40 No. 2 Caradoc brachiopods from the Shan States, Burma (Myanmar)—A review of the stratigraphy and trilobite faunas from the Cambrian Burj Formation in Jordan—The first Palaezoic rhytidosteid: Trucheosaurus major (Woodward, 1909) from the late Permian of Australia, and a reassessment of the Rhytidosteidae (Amphibia, Temnospondyli)—The rhyn- chonellide brachiopod /sopoma Torley and its distribution. 1998. Pp. 109-163. £43.40 .Volume 55 No. 1 Latest Paleocene to earliest Eocene bryozoans from Chatham Island, New Zealand. 1999. Pp. 1-45. £43.40 No. 2 A new stylophoran echinoderm, Juliaecarpus milnerorum, from the late Ordovician Upper Ktaoua Formation of Morocco— Late Cretaceous-early Tertiary echinoids from northern Spain: implications for the Cretaceous-Tertiary extinction event. 1999. Pp 47-137. £43.40 CONTENTS 3 A review of the history, geology and age of Burmese amber (Burmite) V.V. Zherikhin & A.J. Ross 11 A list of type and figured specimens of insects and other inclusions in Burmese amber A. J. Ross & P-V. York 21 Apreliminary list of arthropod families present in the Burmese amber collection at The Natural History Museum, London A. P. Rasnitsyn & A. J. Ross 25 __—iThe first fossil prosopistomatid mayfly from Burmese amber (Ephemeroptera; Prosopistomatidae) D. Sinitshenkova 29 The most primitive whiteflies (Hemiptera; Aleyrodidae; Bernaeinae subfam. nov.) from the Mesozoic of Asia and Burmese amber, with an overview of Burmese amber hemipterans D.E. Shcherbakov 39 Anew genus and species of Lophioneuridae from Burmese amber (Thripida (=Thysanoptera): Lophioneurina), V. V. Zherikhin 43 Burmapsilocephala cockerelli,a new genus and species of Asiloidea (Diptera) from Bur- mese amber S.D..Gaimati.&M.B..Mostovski 47 Phantom midges (Diptera: Chaoboridae) from Burmese amber E. D. Lukashevich 53 An archaic new genus of Evaniidae (Insecta: Hymenoptera) and implications for the biology of ancestral evanioids H.H. Basibuyuk, A.P. Rasnitsyn, M.G. Fitton & D.L.J. Quicke 59 Digger Wasps (Hymenoptera, Sphecidae) in Burmese Amber A.V. Antropov 79 Electrobisium acutum Cockerell, a cheiridiid pseudoscorpion from Burmese amber, with remarks on the validity of the Cheiridioidea (Arachnida, Chelonethi) M.L.I. Judson Bulletin of The Natural History Museum GEOLOGY SERIES Vol. 56, No. 1, June 2000