a APPENDIX ONE PUPARIA OF BRITISH

This consists of I copy of the revision of the puparia of British Sphuroceridu

(Skidllore 1993), which vas partly based upon the copious quantity of laterial studied during this research progra.me, It delonstrltes the way in which subfossil laterial can assist in lodern iaxonolic studies,

Notes on the of the puparia of British Sphaerocerldae

Peter Skidmore

In the course of studying dipterous material frem archaeological sites in Greenland, Iceland and \'arious parts of the British Isles, the author has examined some is,ooo sphaerocerid puparia. The importance of this in this field of research was noted by Smith (1989), but 1 have found that in total, members of this family, and especially those belonging to the , have massively outnumbered all other Cyc10rrhapha combined. Clearly the identification of this material is of paramount importance in the interpretation of these deposits but the existing literature is inadequate for this purpose and a reappraisal of the tuonomy of sphaerocerid puparia is essential. .

Pitkin (1988) provides references for the puparia of Sphaeroceridae described in the literature, and Okely (1974) gives keys for the identification of those which were then known. Whilst some may have been based upon misidentifications, the main problems were that some important features had been overlooked and that descriptions traditionally included trivial details emanating from the process of pupariation, and of structures common to most cyclorrhaphous puparia. The present paper aims to highlight characters which appear to be of taxonomic importance with a view to producing improved keys.

"Subfossil" puparia such as occur in postglacial deposits are, perhaps surprisingly, of great pot.e:ltial taltonomic value. Traditionally, the identification of emerged adults has been regarded as an essential prerequisite for the production of descriptions of puparia and dipterists ha\'e often discarded unhatched puparia. In "sub fossil" material of course one is forced to rely upon the examination of unhatched for positive identification. The author has found that in deposits he has studied, puparia have occurred in huge numbers, and that between 1 and 10 percent have contained unhatched adults, often in a remarkably good state of preservation. In Iimosinine puparia such vital diagnostic features as male genitalia and pregenital combs, female spermathecae and cerci, and almost all important chaetotactic details may be clearly visible through the puparial integument. Obviously when identifications are based on the adult within the puparium, errors so easily resulting from incorrect correlation of puparia and emerged adults are inapplicable. Dipterists only interested in modem material should also bear this in mind: before throwing out those unhatched puparia, closely examine them (or signs of the unhatched adult inside (see fig. 1~ .

Some of the excavations in which the writer has been involved in this study have yet to be published, even in outline. For those which have, the reader is referred to the following papers covering the respective sites - Baker &t Higham 1982 (Hen Domen, Montgomery), Buckland 1983 and McGovern tl QI 1983 (\\'estern Settlement, Greenland), Greig 1981 (Barrel Latrine, Worcester), Leach 1984 (Taunton) and Sveinbjarnard6ttir 1981 (St.6raborg).

The terminology for morphological features of the puparium follows Skidmore (1985) and nomenclature that of Pitkin (op. clr.).

6

315 1a 1b

"~"":"'.""""",,",:,"'i\~:. 1c

...... ~ .. , ...... I I )

4b

3a 2a 4a

Fl,. 1-4. 1 (s.str.) simi/is, a, puparium (lateral), b, anal spiracles (posterior), c, prospiraeular processes showing two variants. 2a Ischlolepta denticulara, puparium (dorsal) showing details of prospiraeular process and right anal spiracle. 3a, 1. pusflla, puparium (dorsal) showing details of prospiracular process and right anal spiracle. 4 curvipes, a, caudal segment of puparium (dorsal), b, right anal spiracle. (Scales: la, O.6m~j lb, Ie, O.12mmj 2a,3a,4a.4b O.24mm) (Orig.) 7

316 ~ ... ~ ~.. ~ .

. t 5b

.0" :~.-: ...... ~ vf#l,,_- ;'.J ·5a

a

V~,.. b

.... -- ... @ .~ & ~.... ~ 9 8 ·7a 7b F1z. 5-9. 5 HemiosiTUJ bequaerti, a, caudal segment of puparium (dorsal), b, thoracic segments showing detail of prospiracular process. 6 Leplocera (S.SIr.) /ofllinalis, caudal . segment of puparium (ventral) showing detail of left anal spiracle. 7 Opa/imosiTUJ (pappiella) llliputana, a, puparium (ventral), b, anal spiracle in dorsal and frontal view. 8 heteroMura, puparium (dorsal) showing detail of right anal spiracle. 9 (s.str.) Tl4/ilabrls, a, puparium (dorsal) showing detail ofprospiracular process, b, anal spiracle in various aspects. (Scales: O.12mm) (Orig.) 8

317 ...... ~ 10······ ...... 0-

~ .. ~ . .~"" ; ...... ~ ·...... ;··~'t; ~1·:;.r.;·~·:.;,~·~·~.r:·.::r:N.- :, .. ... , --=::::;:.:;~.,-~~~ .. ~:.~:./! .. =£4;~;Q~ ~.,;.;.;y;;,;-~

10b a

10c b 9c ~ 10 11

Fig. 1~11. 10 TelomerlnaJlavipes, a, puparium (ventral) showing details ofprospiracular lobes and anal spiracle, b and c, median and lateral section respectively of ventral ambulatory welt on abdominal segment 4. 11 racovitzal, a, prothoracic and caudal segments of puparium in dorsal and ventral aspect respectively, showing detail of prospiracutar process, b and e, two views of anal spiracle. (Scales: lOa and lla, O.12mm; IOb,e, O.03mm) (Orig.)

9

318 12 13 14

Hg. 12-14. 1l1horacochaeta brachystoma, puparium (dorsal). 13 T. zoster~, a, puparium Qateral), b, same (ventral), showing details of prospiracutar process, c, detail of anal spiracle. 14 (s.SIr) coprintJ, puparium with outline of adult inside, showing details of prospiracutar process and anal spiracle. (Scales: O.12mm) (Orig.)

10

319 Dimensions Q/puraria and adult.!

Apart from the superb works of RoMcek (1982-5) and Marshall & RoMee\:: (1984), which have set the very highest standards in taxonomic work in this family, most workers have omitted biometric details for adults, invoking an argument of extreme variability. Although most sphaerocerids vary in size intraspecifically, there are nevertheless considerable interspecific differences within the family. RoMeek & Marshall not only give the adult size range for each sex, but state the number of specimens examined. In puparial descriptions a single figure has usually been given but the number of specimens studied is rarely stated. The author has found some evidence to suggest that the ratio of adult to puparial length (AlP ratio) is not constant and there may be differences in this ratio between certain genera. Clearly these ratios can only be calculated from a large amount of biometric data. The details for puparia provided in this paper result mainly from material from archaeological excavations. The AlP ratio in the studied lies mainly in the range 0.61-0.78:1. The puparial1ength given by Okely for (s. sIr.) mirabilis appears wildly excessive, especially when compared with the available details for o. (Pappiel/a) lilipwana (Rondani) (see table I). Also the bent form of the puparium of o. mirabilis figured by him is not a specific character but a malformation seen very commonly in many limosinines.

Table 1 Calculated Adult/Puparium Ratios for certain British species of Limosininae. Mean dimensions for adults calculated from the size­ ranges given in RoMcek. Puparial lengths marked with an • from Okely, the remainder calculated from specimens examined by the current author.

Species L. (Adult) mm L. (Puparium) nun AlP ratio

O. lilipulana 1.45 2.28 .64

O. mirabilir 1.34 2.9* .46

P. heleroneura 1.25 1.74 .70

P. moesta 1.50 2.1* .71

P. pullula 1.39 2.0* .70 s. rujilabris 1.64 2.16 .76

T·flavipes 1.41 2.31 .61

T. zosterae 2.34 3.5 .67

frQIIlirac'lImr prQr.(J.r~s.

These 'structures are clearly of great taxonomic significance as has been recognised by previous students of the group. Goddard (1938) an~ Okely rely almost wholly on these in

11

320 their keys to puparia. They range from short stellate or palmate structures to very long dendriform ones adorned with lateral, usually translucent linear papillae (see figs. 14, Ie and 9 for stellate, palmate and elongate dendriform types respectively). In the dendriform type the length of the processes relative to the space separating them provides a useful character (the PIP ratio). The British Limosininae are broadly categorised below on the calculated PIP ratios of their puparia. Whilst the papillae arising from the dendriform types are brittle and prone to loss, their points of attachment remain as distinct alveoli. As noted by Ferm (1987) the pupal spiracles are internal.

Prospiracular process very short (PIP = up to 0.3:1):

Apleromyia c/avivenrris: Chaelopodel/a sCUlellaris: (hinula, ?Iugubris): Herniosina beqlUleni: uplocera (s. sir.) (caenosa,lonlinalis, oldenbcrgi): L. (sg. ) (fu.rcipennis): silvaliea; (fungicola): Pleremis leneslralis: (brachystoma, z.osrerae): Trachyopella (s.slr.) (coprina, lint'ajrons).

Prospiracular process medium to very long (PIP = 0.5-1.5:1):

Coproica (pusio, vagans),' arerrima: Halidayina spinipennis: Opalimosina (s.g Pappiella) lilipurana; O. (s.slr.) (mirabilis); Pullimosina (heteroneura, moesta, pullula); Spelobia (Bi/ronsina) bifrons; S. (s.slr.) (all known puparia including rufilabris); Spinilimosina brevicosrata; (flavipes): Terrilimosina (racovilzai): TrachyopeUa (Nudopel/a) leucoplera.

Anal miracles

The taxonomic importance of these structures has been largely overlooked. Although the clear differences between the forms found in the three has been long recognised, in the Limosininae they appear to have been ignored. It will be clear from the descriptions given below that they provide very promising characters.

Lorera' muscle scars

The grooves along the sides of sphaerocerid puparia have been used as features of taxonomic importance but their utility in this regard is highly dubious as they result from differentials in cuticular tensions during pupariation. The parallel dorsal muscle scars on the caudal segment are similarly unlikely to be of major taxonomic significance.

Ventra' ambulatory wellS

These appear to be remarkably uniform throughout the family, consisting of a median row of 12-20 large rounded spicules with 3-5 rows of minute serrations before and 2-5 rows behind; laterally the large median spicules may diminish in size and the serrations increase so that all spicules become more nearly uniform in size (see fig. IOb,c).

12

321 Key to subfamilies The key in Okely requires some amendment as follows.

1. Anal spiracles on dorsal surface of caudal segment and hence invisible from ventral view (fig. 2a,3a,4a); anal spiracular slits encircling the scar which is central (fig. 4b) or inferior (fig. 2a,3a). Prospiracular process stellate (fig. 3a) or dendriform (fig. Za) ...... •••

- Anal spiracles on posterior surface of caudal segment, so visible from ventral and dorsal . view ...... 2

2. Anal spiracle large, usually barely exserted, the peritreme usually in contact with the posterior face of caudal segment (fig. Ia); anal spiracular slits convergent upon median scar (fig. Ib). Prospiracular processes very short dendriform, stellate or palmate (fig. Ic), never elongate ...... COpromyunae

- Anal spiracles usually on tubercles exserted from posterior surface of caudal segment (figs. 5·14); respiratory' slits peripheral, sear median. Prospiracular horns often very elongate, with lateral papillae (figs. 9a, 10), but sometimes stellate (fig. 13b) or palmate (fig.lla) ...... Umosininae

Copromyzinae

The puparia of this family are surprisingly little known considering the extreme abundance of many of the species and the fact that our largest sphaerocerids belong here. Alloborborus is undescribed in the puparial state and only one (i.e sordidus (Zeit.» is known. It is not yet possible to distinguish the described puparia of CopromylA, Crunwmyia or Lolophila. The anal spiracles may sometimes appear to be slightly exserted (c.f C. pedeslris) but their large size and convergent slits are diagnostic.

Copromyza (s.slr.) similis (fig. I) typifies the known puparia found in this subfamily. Prospiracular process palmate or very short dendriform with 5 to 7 dark lobes. Ventral ambulatory welts usually indiscernible apart from the row of large spicules. Intersegmental margins, dorsally and laterally with a conspicuous series of cuticular knots which, on the sides delimit the pleural seleriles. Anal spiracula,r plates large with very conspicuous convergent slits. Length, puparia 3.60-4.38mm (n=9); adults 3.12-3.78mm (n=9); AlP "" .86. [Material studied, 6 puparia ex cow dung, Bentley Common, Doncaster (England), em. April 1974 and May 1976 (PS); 3 puparia ex: dung Salterforth, Yorks., (England), em. 2 April 1949 (A. Brindle, coIl. Manchester Museum). Many puparia from archaeological excavations at Hen Domen, Montgomery (Wales), Tuquoy, Westray (Orkney) and St6raborg (Iceland) also examined, but these probably include C. (s.slr.) equina.]

Dimensions given in the literature for puparia of other copromyzines are:

Copromyza (s.slr.) eqllina, 3.S-3.8mm (n "" 6, Goddard) C. slercoraria, 3.64mm (n "" I, Goddard) C.jimefaria, 3.3mm (Okely) (S.SIr.) nigra, 4.Smm (Okely) C. nOfahilis, 3.4mm (Goddard, sllb nom. glocioUs)

13

322 C. (Aprerina) pedesrris, 3.5mm (Deeming & Knutson, 1966) . Ferrar (1987) figures the third instar , showing details of anal spiracles and cephalopharyngea1 skeleton, of Borborillus sordidus.

Sphaerocerinae

The described sphaerocerine puparia (i.e pusilla and scabricula, and and monilis) are unlikely to be confused with those of the other subfamilies. The unusual position of the anal spiracles, on the dorsal surface of the caudal segment, with their central or inferior scar and the arrangement of their slits (when visible) are characteristic of this subfamily. Prospiracular processes often of aberrant form (figs. 2a,3a). S. curvipes is the largest species, exceeding 3mm in length, shiny black in colour with small, stellate 8-lobed prospiracular processes. I. scabricula is the smallest at about 1.8mm. Okely separates S. monilis and I. pusilla on the number, colour and shape of prospiracular processes. In I. pusilla the form is of an aberrant stellate type and the lobes (up to 5) are remarkably broad, leaf-like and pale in colour (fig. 3a). Okely's figure of I. scabricula shows that this structure is similar in that species. In stark contrast are the processes seen in S. monilis and I. dt'r:Iiculara (fig. 2a) which are truly dendriform with long stout black spiniform papillae.

The details for the specimens figured here are as follows: Ischioiepra denriculala (fig. 2), Length, puparium 2.20mm; adult 1.85mm tx old cow- . dung, Bentley Common, Doncaster (England) em. May 1976 (PS) I. pusilla (fig. 3), Length, puparium 2.22mm; adult 1.9Smm Victoria Park, Manchester (England), em. 10 June 1924 (L.W. Grensted; Manchester Museum) Sphaerocera curvipes (fig. 4), Length, puparia 3.36-3.8Smm (n ... 7) (Goddard); adults 3.05-4.20mm (n a 9) [Material studied, 7 incomplete puparia from Tuquoy excavation on Westray (Orkney).)

Llmosininae

The puparia of six genera are unknown (i.e Gigalimosina, Kimosina, Opaci/rons, Philocoprella, and Pllncricorpus). Goddard and Okely emphasised the taxonomic significance of the prospirncular processes and the PIP ratios, but unfortunately their descriptions and figures gave little information on the anal spiracles. In all species examined by the author however, and in the better published figures, the slits appear to be peripheral, the median one lying parallel with the edge of the peritreme. Furthermore, the slits are often on raised bosses, separated from each other by deep fissures. In Spelobia these bosses are peculiarly angulate giving a very distinctive uneven hoof-like silhouette to the spiracular plate (fig. 9b).

Most authors have commented upon the whitish or very pale colour of some limosinine puparia. This is particularly the case in Pullimosina (perhaps all species), but is also noted by Okely In Coproic:a pusio and Spelobia (Bifronsina) bi/rons.

In the treatment below dimensions for adults are mostly from Rohlicek.

14

323 Vimmer

Okely and RoMcek figure the puparium of the single species A. c1aviventris. pointing out that the figure in Richards (1930) was drawn from a damaged specimen. Prospiracular processes elongate dendriform (pIP = 0.3: I) with 4 short lobes. Anal spiracles on short, thick tubercles. their slits on low rounded protuberances and apparently peripheral. Ventral ambulatory welts with usual transverse row of large rounded spicules, preceded and succeeded by 3 rows of much smaller ones. Length. puparium (not stated.by Okely); adults (male) 1.49-2.08mm (n ... 171). (female) 1.47-2.S0mm (n = 157).

Genus Duda

RoMcek figures details of all three larval instars and the prospiracular horns and anal spiracles of the mature larva of the single species C. seu/ellaris, but the puparium remains undescribed. Prospiracular horns short dendriform with 9 lobes. Anal spiracles with the typical peripheral alignment of the slits. Length of adults (male) 1.43-1.85mm (n ... 243), (female) 1.5 1-1.87mm (n = 380).

Genus Coproic:a Rondani

Okely figures the puparia of two species (i.e. C. hirtula and pusio), whilst Goddard figures C. vagans. Anal spiracular slits clearly peripheral but the species differ in the length of the prospiracular horns (see below). Also, Okely says that whilst C. hirtula is black. pusio is white. Goddard does not give the colour of C. vagans.

PIP = over I: I. Length 2.1 mm C. pl/sio (= pseudolugubris (Duda» PIP'" c.O.S:1.Length 2.4-2.62mm C. vagans PIP -= c. 0.25:1. Length 1.S-1.75mm C. hirtula.

Ferrar gives spiracular details of C. //lgubris; prospiracular horns long dendriform with 12 close-set lobes and anal spiracular slits again peripheral. PIP not calculable from his figure.

Genus Elachisoma Rondanl

Okely figures E. alerrima stating that it is very like Halidayina spinipennis but smaller. Prospiracular horns extremely long (pIP = c. 1.5: 1) with 7 short pale papillae. Length, puparium 1.6mm (Okely); adults O.6-1.2mm (RoMcek).

Genus Halidayina Duda

According to Okely this differs from the previous one only in the larger size. His figures suggest a very strong resemblance to Spe/obia s. SIT. with a PIP about 1: 1 and anal spiracles similarly exserted. Length, puparium 1.95mm (Okely); adults (male) l.OI-1.28mm (n - 982), (female) 1.03-1.63mm (n ... 1252).

15

324 Genus Hemiosina Rohlicek

H. beqUlleni, the only species, is one of the larger limosinines. Prospiraculai process subpalmate with 6 lobes; anal spiracles rather inflated and situated on broad tubercles (fig.

S). Length, puparia 2.S4-3.Slmm (n I: 14) (Goddard gives 3mm); adults (male) 2.26- 2.72mm (n = 32); (female) 2.22-3.0Smm (n ... 34). [Material studied - 13 puparia, St6raborg farm excavation (Iceland); 1 puparium Buiston Crannog excavation, Kilmarnock: (Scotland). Males were dissected from unhatched puparia from both sites.)

Genus Leptocero Olivier

Four species are known in the puparial state (i.e L. (s. su.) caenosa, jOnJinalis and oldenbergi, and L. (Rochispodo) juscipennis. In L. jOnJinalis (fig. 6), which is a rather large species, prospiracular lobes extremely short, simple (?palmate), with very short lobes; anal spiracles shiny black, abruptly contracted below peritrernes but swollen basally and situated on prominent, widely separated tubercles. Length, puparla 3.19-3.7Smm (n .. IS) (Okely gives 3-3.3mm); adults 2.40-3.4Smm (n = 4). [Material studied - S puparia 4 Bessasta~ir and St6raborg excavations (Iceland); 7 puparia from Tuquoy, Westray (Orkney); 3 puparia Papa Stour (Shetland).]

L. oldenbergi is according to Okely smaller than jOnJinalis with similarly extremely short prospiracular processes. Length 2.8Smm. Ferrar figures L. (Rochispoda) juscipennis, again showing very short prospiracular horns.

Genus Umosina Marquart

L. silvatica, the only species, reaches a rather larger size than any other limosinine. Prospiracular horns very short, subpalmate with 7 Jobes. Goddard says that the anal spiracular discs are flattened and directed inwards. Length, puparla 3.42-3.S2mm (n = S, Goddard); adults (male) 2.86-3.6Imm (n = 144); (female) 3.02-3.73mm (n = 157). [Material examined, 9 incomplete puparia from Tuquoy, Westray (Orkney); 4 from Bessasta~ir (Iceland); 1 from St6raborg (Iceland).]

NB. Gigalimosinaflaviceps is only slightly smaller than silvQrica so should have a large puparium. Length, adults 2.66-3.50mm (n = 236).

Genus Minilimosina Rohlicek

The only species known in the puparial stage is M. (s. SIr.) fungicola, in which the prospiracular processes are very small, palmate, and 3-lobed. Anal spiracles minute, on distinct though short tubercles. Length, puparia 1.68-1.86mm (n .. 3) (Goddard gives 1.8mm); adults (male) l.03-1.3Smm (n = 93); (female) 1.14-1.6Imm (n - 113). [Material studied, 3 puparia from Tuquoy excavation, Westray (Orkney) tentatively referred to this species on the form of the female spermathecae and cerci.l

16

325 Genus Opalimorin.a Rohlicek

Puparia of two species are known although Okely's description of 0 (s. str.) mirabilis is based on a deformed specimen and there is clearly some mistake with the dimensions given. PIP ... c.0.6: I ~th up to 4 elongate lobes. It appears likely that the species is extremely similar to the next one. Length, puparium "2.9mm" (Okely); adults (male) 1.D4-1.45mm (n coo 142); (female) 1.05-1.65mm (n ... 127).

o. (Pappie/la) liliputana (fig. 7) has elongate dendriform prospiracular horns curved backwards, with 6 long pale lobes; PIP = c.' 0.5: I. Anal spiracles noticeably slender triangular in dorsal view, on long, distally narrowed tubercles (very similar to TelomeriTUl flavipes but slits relatively smaller). Length, puparia 2.16-2.40mm (n = 48); adults (male) 1.23-1.58mm (n = 83), (female) 1.34-1.81mm (n = 102). [Material studied, 48 puparia, including unhatched males. showing genital characters perfectly, Tuquoy excavation, Westray (Orkney).] ,

Genus Rondanl

The puparium of P. feneslralis is figured and described by Okely. Prospiracular horns palmate, with up to 3 lobes. Anal spiracles clearly exserted, but more detailed structure indiscernible from his figures. Length, puparium 2.6mm (Okely). Roh~cek regards this as a subgenus of .

Genus Pul/imorina Rohlicek

The puparia of three species have been described and figured (i.e P. hetuoneura. moesta and pul/ula). The writer has also examined puparia of P. heteroneura. Okely says puparia of heteroneura are white, a fact which my experience confirms, whilst in moesta they are pale yellow. Goddard gives the colour of heteroneura as light yellowish brown, but possibly some discolouration may emanate from their environment. In all three species the prospiracular processes are elongate dendriform '(p/P up to 0.6: I). Goddard's figure for heteroneura differs from mine (fig. 8) in the degree of exsertion of the anal spiracles but the disparity is inconclusive. Rohcicek gives electroscan photographs of the anal spiracles of pul/ula; here the slits appear to be on slightly raised prominences and peripherally arranged. Okely details his laboratory rearing of pul/lIla, stressing the parthenogenesis found in this species. Lengths for the three species are as follows. P. heteroneura. puparia. 1.56-1.92mm (n - 75)(Goddard gives 2-2.1mm, n'" 8); adults (male) 0.95-1.44mm (n - 220), (female) 1.04-1.56mm (n ... 252). [Material studied, 7S puparia, some containing male adults, from excavation at Tuquoy, Westray (Orkney).] P. moesta. puparia, 2.lmm (Okely); adults (male) 1.14-1.52mm (n - 131), (female) 1.26-1.87mm (n ... ISS). P. pullula. puparia, 2.0mm (Okely); adults (male) 1.I1-1.23mm (n a 2), (female) 1.07-1.71mm (n .. 294).

17

326 Genus Spelobia Spuler

Puparia of 9 species are known, the most aberrant being S. (Bi/ronsina) bi/rons, the smallest member of the genus and the only one in the subgenus. In this species the puparium is white (Okely). The dimensions for this species are given below.

Puparia of Spelobia S.slr. are exceedingly similar and probably indistinguishable on external characters. In all known puparia the prospiracular horns are elongate dendriform with a PIP ratio about 1: 1 (except S. pseudosefaria at 0.6: 1 (Richards) and parapusio at O.S: 1 (Rohacek); all have conspicuous lateral translucent papillae. Anal spiracles strongly exserted, on conspicuous basal tubercles. Figure 9 shows the puparium of S. rujilabris and attention is drawn to the peculiar shape of the anal spiracles in which the slits are arranged on sharply raised bosses separated by very deep fissures. Spelobia (s.srr.) lureilabris has an identical anal spiracle and the electroscan photograph of S. parapusio in Rohlicek shows the same structure. It is impossible to conclude anything from the figures in Okely and Goddard in this feature but it is most likely that all members of this subgenus will be identical in the fonn of the anal spiracles. Dimensions for members of the genus are as follows.

S. (Bi/ronsina) bi/rons, pupa ria 1.6mm (Okely); adults (male) 1.07-1.26mm (n = 51), (female) 1.l9-1.56mm (n - SI) S. (s.Slr.) clunipes, puparia 2.16-2.55mm (Goddard and Ouly sub nom. crassimana and manicara): adults (male) l.5l-2.5Smm (n = 1007), (female) l.59-2.50mm (n ... 1136) S. lUleilabris, puparia 2.sS(pS)-2.6mm(Okely); adults (male) l.ss-2.0Smm (n - 115), (female) l.59-2.30mm (n = 140)_ [Material studied, 1 puparium "'ith unhatched male adult, Tuquoy excavations, West ray (Orkney).] S. palmara, puparium 2.66mm (Goddard); adults (male) 1.70-2.1Smm (n ... 122), (female) 1.69-2.62mm (n ... 146) S. parapusio. puparia 2.2mm (Okely); adults (male) 1.67-1.8lmm (n = 2). (female) l.s8-2.22mm (n = 259). A parthenogenetic species. S. pseudonivalis, puparia 2.6mm (Okely); adults (male) 1.98-2_78mm (n ... 7), (female) l.S2-2.s8mm (n - 17) S. rujilabris, puparia 1.92-2.40mm (n ... 959); adults (male) 1.50-1.79mm (n = 25), (female) 1.70-2.06mm (n - 21). [Material studied. 959 puparia, many with unhatched male adults, Tuqlloy excavation, West ray (Orkney)]. NB. It is highly likely that some of the hatched pllparia from Tuquoy belonged to S. lUfeilabris although in such localities the present species appears much commoner. Some wings from the same site were more like S. c1unipes in the broad alula so puparia of that species too may have been present. S. la/parum, puparia 2.35mm (Goddard); adults (male) 1.S2-2.10mm (n ... 112); (female) 1.61-2:24mm (n = 215)

Genus Spinilimosina Roh:icek

Ferrar figures the prospiracular horns of S. ruji/rons (Duda), showing them to be elongate dendriform (pIP = 1.5: I).

18

327 Genus Telomerina RoMcek

The puparium of T. flavipcs (fig. 10) is very like that of Opalimosina liliputana but of a rather larger average size and with anal spiracular slits relatively larger. Length, puparia 1.74-2.88mm (n = 1536); adults (male) 1.I5-1.43mm (n - 32), (female) 1.31-1.67mm (n - 54). [Material studied, 1119 puparia from various farmstead sites in the Western Settlement at the head of Godlhaabsfjord (West Greenland); 28 puparia from St6raborg excavation and 13 from that at Bessasta~ir (Iceland); 360 puparia from Pluscarden Priory, Nairn (Scotland); I puparium from Hen Domen, Montgomery (Wales); 6 puparia from Buiston Crannog, Kilmarnock (Scotland); 3 from "Barrel Latrine", Worcester (England); 4 from Taunton excavation, Somerset (England). Unhatched males were dissected from puparia from all of the above sites]

NB. This appears to have been the commonest limosinine eusynanthrope of the Viking settlements in Iceland and probably the only one to reach West Greenland. The species is unknown in Greenland today (Henrikson, 1937) and it presumably shared the demise of the early nordic settlers who took it there from Iceland.

Genus Terrilimosina Rohlicek

The puparium of T. racovitzai is figured here (fig. 11) for the first time and no other species has been described in the literature. Prospiracular processes very short palmate with 3 lobes (fig. lla). Anal spiracles bulbous basally and constricted below peritreme (fig. lla,b,c), shiny black in colour, on short broad tubercles. Length of puparia 2.19-2.68mm (n = 43); adults (male) 1.35"1.79mm (n = 26), (female) 1.63-2.22mm (n = 41). [Material studied, 43 puparia, 3 containing unhatched males, Pluscarden Priory, Nairn (Scotland).]

Genus Thoracocllaeta Duda

Puparia of two British species have been described and figured and are treated here. Confusion between them is unlikely due to their great disparity in size and morphology. T. brachystoma (fig. 12» is a very small species with simple bilobed prospiracular processes (fig. 12b); anal spiracles on rather long processes which are directed outwards distally. T. zosterae is much larger and has been figured by many authors (eg. Richards 1930, Egglishaw 1961, Pitkin 1988, Smith 1989, Belshaw 1989 etc.), being a very familiar and easily recognised inhabitant of littoral wrack beds. The integument is thick and often opaque blackish in colour whilst the long outwardly directed anal spiracular processes and generally flattened form with "crimped margins" give a highly distinctive appearance (fig. l3a,b). Prospiracular processes stellate with up to 12 radially arranged lobes (fig. l3b). Anal spiracular slits peripheral (fig. 13d). Dimensions are as follows.

T. brachysloma. puparia 1.73-2.17mm (n - 17); adults 1.32-1.68mm (n - 3).[Material studied, 17 puparia, some with unhatched adults, collected from decaying seaweed on foreshore at Howbeg, South Uist (Scotland).]

NB. According to Tenorio (in Ferrar, op. cit.) the larvae of the two species dealt with here are very alike but in this one there are 6 prospiracular lobes and in the next one 9.

19

328 T. zosterae, puparia 3.02-4.2mm (n .. 362); adults (male) 1.74-2.40mm (n ... 10); (female) 2.04-2.94mm (n - 5). [Material studied, 27 puparia from rotting seaweed on Howbeg foreshore, South Uist (Scotland); 27 from Tuquoy excavation, Westray (Orkney); 302 from Taunton excavation, Somerset (England); 4 from "Barrel latrine" excavation, Worcester (England); 2 from Bessasta<:lir palace excavation (Iceland).]

NB. Despite the highly distinctive appearance of the puparium of this fly it has been the cause of much confusion in an archaeological context. Belshaw (op. cit.) noted that records of the Urinal fly Teichomyza/Ilsca Macquart from archaeological sites in Britain had proved to belong to T. zosrerae. This discovery has provoked considerable consternation amongst historians. Why was this fly, which is only known to breed in seaweed, so common around human habitations in medieval England ? Had there been a widespread trade in seaweed throughout the country, or has the biology of the fly undergone some major change? Pitkin draws attention to modern inland records of this fly, but these are very rare and probably the result of casual importations.

Genus Trachyope//a Duda

Descriptions of the puparia of three species occur in the literature and one of these is here figured (fig. 14). Okely figures and describes T. (Nudopella) /eucoprera in which the prospiracular horns are elongate dendriform (PIP .. c.0.6: I), with 4 pale lobes. Anal spiracles on very long, broad-based processes, similar to T. zosterae in relative length. Length 2.lmm (Okely).

The other two described puparia belong to TrachyopeUa s.str. Like the previous one they are translucent, pale yellowish brown. In T. Iinea/rons according to Okely the prospiracular processes are very small, palmate with up to 4 lobes, whilst the anal spiracles are strongly exserted on large tubercles. Tenorio (in Ferrar, op. cit.) showed that the larval mouth-hooks are strongly serrate; an unique feature amongst known sphaerocerids and usually regarded as indicating a phytophagous mode of life. Length 1.8mm (Okely).

Assuming the reliability of Okely's figure for the last species, T. coprina differs very strikingly in the absence of anal spiracular tubercles, for here the spiracles are hardly exserted at all (fig: 14). Goddard's figure for this species agrees perfectly with specimens seen by the present writer in this regard, but he states that the prospiracular processes are only 3 lobed. The number clearly varies and often reaches 6 (fig: 14), arranged in a stellate fashion. Length of puparia 1.32-J.74mm (n ... 9313). [Material studied, 9313 puparia, many containing adults (both sexes), Tuquoy excavation, Westray (Orkney).]

Acknowledgements

This work is part of a doctorate generously funded by the Archaeological Operations and Conservation section, Historic Buildings and Monuments, Scottish Development Department, Edinburgh, through the kind offices of Dr. P.C. Buckland of Sheffield University, my tutor in this course of study. It is to them that my greatest debt of gratitude is acknowledged. It is also through Dr. Buckland that material from archaeological excavations in which he has been involved has reached me. Amongst others to whom I offer thanks are Dr. B.R. Pitkin of the Natural History Museum, London, for confirmation of

20

329 . some of my identifications, Dr. J. Sadler of Sheffield University for information on excavation details and to my long-standing friend Colin Johnson of Manchester Museum for the loan of some material froin the collections of that institution.

References

Baker, P. & Hen Domen Montgomery. A Timber Castle on the Higham, R. (1982) English-Welsh Border The Royal Archaeological Institute 1: 67-71

Belshaw, R. (1989) A Note on the Recovery of (Haliday) (Diptera: Sphaeroceridae) from . Archaeological Deposits Circaea 6(1): 39-41

Buckland, P.C. (1983) Norsemen at Nipaitsoq, Greenland. A Palaeoecologica1lnvestigation NOM. Arch. Rev. 16(2): 86-98 :

Egglishaw, H.J. (1961) The Life History of 17lOracochaeta zosterae (Hal.) (Dip!., Sphaeroceridae) Entomologist's mono Mag. 96: 124-128

Ferrar, P. (1987) A Guide to the Breeding Habits and Immature Stages of the Diptera Cyclorrhapha Entol7lonograph 8; 907pp

Greig, J. (1981) The Investigation of a Medieval Barrel·latrine from Worcester Journal ofArchaeological Science 8: 265-282

Henriksen, K.L. (1937) A Revised Index of the of Greenland Mctld. am Gronland 119(10); lllpp

Leach, P. (1984) The Archaeology of Taunton Western Archaeological Trust E:ccavation Mono.r:raph 8; 202pp

McGovern, T.H. et al (1983) A Study of the Faunal and Floral Remains from the Norse farms in the Western Settlement, Greenland Arcric Archal'olngy 20(2): 99-120

Marshall, S.A A Revision of the Genus Telomerina RoMcek & Rohlicek, J. (1984) (Diptera. Sphaeroceridae) Systematic Entomology 9: 127-163

Okely, E.F. (1974) Description of the puparia of twenty-three British species of Sphaeroceridae (Diptera, )

21

330 Trans. R. ent. Soc. Lond. 126(1): 41-56

Pitkin, B.R. (1988) Lesser Dung Flies (Diptera, Sphaeroceridae) Halldbk. Ident. Br. Ins. 10(Se); 175pp

Richards,O.W. (1930) The British Species of Sphaeroceridae (Borboridae, Diptera) Prot'. Zoo/. Soc. Lond. 2(18): 18-348

Roh:icek, J. (1982-5) A monograph and reclassification of the previous genus Limosina Macquart (Diptera, Sphaeroceridae) of Europe Bdtrage zur Entomologie Pt. 1 (1982) 32(1): 195- 282: PI. 2 (1983) 33(1): 3-195: Pt. 3 (1984) 33(2): 203-255; PI. 4 (1985) 35(1): 101-179

Skidmore, P. (1984) The Biology of the of the World Series Ent. 29; 550pp

Smith, K.G.V. (1986) A Manual of . Brit. Mus. (Nat. Hist.); 206pp

Smith, K.G. V. (1989) An Introduction to the Immature Stages of British Flies Handbk. Ident. Br. Ins. 10(14); 280pp

S"einbjarnard6ttir, G. Excavations at St6raborg: a palaeoecological et al (1981) approach Arhtlk hins hlen.lkafornleifaifclags 1980; 17pp

P. Skidmore /69 Carr House Road. Doncaster. South Yorkshire. DN4SDP

22

331 APPENDIX TWO SITE ANALYSES

Introduction. In this section are included the detailed

analyses of all of the major sites discussed in this Thesis. The minor

sites merely appear in brackets here, their content being limited to a

very few specimens and hence not warranting inclusion here. They appear

in the main body of the Thesis. A rigid format is adhered to here in

which the numerical totals relate only to the Minimum Number of

Individuals (MNI) of Dipterous material. Total MNI numbers for samples,

subgroups and total sites are given.

ENGLAND

(1 BRIGG RAFT)

(2 CATTERICK CAMP) 3 LEICESTER CAUSEWAY

Table 40 Diptera of the Leicester Causeway Excavation

Taxon 1.16 I. 23 1.24 1.25 1.26 1.28 1.31 1.42 1.45

405 230 318 343 358 376 432 463 325 221 390 412 240 784 763 "NI

SCitOPSl nDt,t, Oxycerllp.

{ri.tlli. tlnlX 100 3 103

Seplid 'p. 20 20

Ulosinine Ip. 40 40

Thorl(O(h"tl lo.t,r" 2

fT,Jo."in, fllvipes 4 ,

Sphaerocerid ap. 2 2

Agrolyzid ap. 2 2

Drolophi 1id 'P.

332 ?Chloropid SP, 2 2

F.nnia ?sc.llri, 16 7 23

Puparia i ndet, A

B

C 3

D 3

Tohl "NI 118 13 2 - - 60 4 2 2 6 290

Taxon 2, 01 2,03 2,04

510 521 5.0 568 579 604 615 503 509 513 537 570 587 766 2.5 .82 "HI

Sc.tops, nat.t.

?If.yetial. 'P,

CnlaroMY. forMol. 3 8 11

[ristaJi, ten.x 60 60

Seplid 'P, 2 2

",pnroch/.MY' rufiventri.

?/lChiollpt• • P,

Lilosinine SP, 2 3

Tnor.coch.,t. ZOltlrae 2 36 18 57

?T,lOl/lrin. fl.viplJ 5 9 3 17

?Trachyop,IJ. SP, Agrorayzid IP, 2 2 5

fSCltl//'Sp, 28 2 31

Drosophi lid IP,

Acalypterlte SP, 2

?Bot.nophil. fug.x

Antholyiid 'P,

333 f.nni. sr.1lril 2 2 9

?Hydrotae. dentipes

?SpiJogon. sp.

1Huscid sp.

Total "HI- 43 15 I 73 39 2 , I 23 2 3 - 209

Taxon 2.13 2.U 2.15 2.16 2.332.34 3 Ph.?

245 '82 206 207 213 218 231 235 257 703 704 234 342 371 Ft "HI

SCitopse not.t. 9 2 II

lIeg.seJi. !rulipel 2 2

II, !nign

Sepsid .p.

?TephrO(hJi.y, r"liventri. 23 23

He1eo.yZl !Clptio61 II II

?llChioJept. Ip. Lilosinine 'P. 8 9

Thor.eo(h.et. zOlter.e 170 161 333

?feJo.erin' fJ,vipes 3 8 12

HydreJJi'lp.

?SClteJJ'lp. 30 33

?Chloropid sp.

ACllypterate SPI

Antholyi id 'PI f,nni. (lnieu1.ri. f,'(lJ.ri. 68 26

?Hydrot,e, dentJpel

Pupariul C 5 8

334 o 2 4

E 2

F

Total "HI- 2 I 318 210 6 5 2 1 1 552

Total I1NI for Leicester Causeway- 970 NB, Upper line of figures give the Phase categories which relate to lain dating periods thus- 1,16 (50-120); 1,23, 1.24,1.25,1,28, 1.42, 1.45 <120-200); 1,26, 1.31 (250-300): 2,01, 2,04, 2,13, 2,14,

2,15 (pre-1100): 2,03 (1100-1250): 2,16, 2,33, 2,34 (pre-1400): 3 (early post-udievall, Phase not given for Illpl.1 342, 371 and F 1(3664 686), The lower figures ref.r to th. lalple numberl, prefixed F in Appendix

Samples from this site are interpreted below as uncontaminated or as putrid on the basis of the presence or absence of species indicative of cess (see Table 2, page 153), Feature descriptions and dates etc, were provided with the specimens,

Box- Area 1 "ed, RB,

IPh,sl ~t3- ItO- 2001 [Uncontalinated] F318 1553 067 [ROlan w.ll]

Stratiolyid (10xy"" IP, ) larva: 1 Julid: 1 Pol ydenid, I1HI- 1

F343 860 384,1 [ROlan ditch]

c,30 11.11 Julids: 1 Elaterid larval anal seglent

F358 824 370 [ROlan pit]

indet,lp,D 1 pupariul I1Hl- 1

IPh.sI /,tl- 120- tOO} [Uncontalinated]

F432 [ROlan post-hoi.]

c,SO 11111 Julidl

335 F463 1507 670

Cyclorrhapha puparia indeto spoA I, spob I, spoC Ii 3 Julids "NI- 3 F463 1554 675 [Rolan pit]

Drosophilid spo 2 pup.riai 3 small Julids "NI- 2

(PIIISI 1,25- 120- 200) [UncontaMinated] F325 701 305 [Rolan linear feature spread]

Tllor,coch"t, zost,r" 1 pupariul I1NI- 1

(PII,ge 1,26- 25Q- 300) [UncontaMinated 1] F221 315 31 (Rolan pit]

Cyclorrhapha pupariul indetospo Ii Co 10 Julidl "HI- 1 (Ph,gl /,28- /20- 200) (UncontaMinated 1] F390 993 453 [Rolan pit]

Cyclorrhaphan pupariul indetolPoD 1 "NI- I F412 (Rolan pit]

Cyclorrhaphan pupariul indetolPoC Ii I Julid

(Ph,gl /,3/- 25Q- 300) £Uncontuinated]

F240 349 90

Cyclorrhaphan indeto olpoC 1 pupariul "HI- I F240 381 89 [Rolan linear feature] Co 50 Julids

[Phi" 2,13- PrI "OO} [UncontaMinated]

F24S 437 142 [Hedieval pit] ?Ephydrid spoindet, I pupariu. (superficially very like Thor,coch"t, but anal plate luch further

frol bases of posterior spirlcles), "HI- 1

(Ph,sI 20 /1- pre /lOO) [Putrid 1] F482 127 588 [Hedieval pit]

Hydrotl" (Hydrotlloidll) ?dentiplI I puparlul I1NI- I

336 [Plli51 2, /b- pre /J()(JJ [UncontaMinated 1]

F213 341 51 [Medieval ?cess pit]

Cyclorrhaphan indet.sp.C I pupariul "HI- I F218 305 68

Cyclorrhaphan indet.lp.C 3 puparia; I Julid "HI- 3 F218 305 68.3 [Hedieval well]

Cyclorrhapha pupari. indet. sp.e 2, sp.E I "NI- 3 F231 395 86 [Medieval pit]

/fegi5eJi. 1nigrl I pupariuI; 2 Julids "NI- I F23s 383 850 [Medieval rubbish pit]

HydreJ/i.sp. 1 pupuiuI; I Julid; 2 Polydeslids; I A,j,uJ. "NI- I F2s7 (476) [201]

indeterlinat. concretion

F257 '91 204 [Hedieval well]

Cyclorrhaphan indet.5p.O I pupariul "HI- I

[PlliSI 3- POlt 'iflY-ledil'l.lJ [Putrid] F234 374 72 [Post-ledievaI pit]

Tllori,o,lIl1t. zOlter., I pupariul HNI- I

£1'11.51 1- not .t.tedJ [Uncontninated?l F342 845 37g [Not Ita ted]

Cyclorrhaphan puparia indet.lp.O 2 "NI- 2 F371 908 396 [Not stated]

Cyclorrhaphan indet.lp.O 1 pupariul "NI- I Subtotal for Box Arll I "edt II pha5ls; 24 lalplll; 185 Iplcilens

Box Aria I Priority* F106, F207, F230, F376, F'OS

£Ph161 /, /6- 5(J- UOJ [Putridl

F40s Area IRS-

337 F405 1753 775

fanni' '",/"is6 puparia, Sphaerocerid sp, 2 puparii; I Julid I1NI: 8

F405 1753 775,3

fanni' 'sci/iri, 10 puparia, pupariul sp,D I, 2 Julids IiNI- II F405 1753 775,4 [RoMan well]

[ristilis ten,x, c,IOO larvae, including at lent 2 prepupae "NI- 100 [PhasI 1,23- 120- 2001 [Putrid]

F376 Area I RS-

F376 (1038) [508] [Rolan pit]

Indeterainate, totally lineralised concretions but one possibly I f,nni, pupariul Ind another I

different Cyciorrhaphoul pupariu" In I third tube under sale coding ',60 puparil apparently of

Sepsidae and Lilosininae I1NI: 62

Bag F230 RS Are. 1-

{Ph,sl 1,23 /2Q- tOOl [Putrid]

F230 1627 782

fanni' 'p, S puparia F230 1627 782

[ri,t,ii. 'p, 3 calcified larvae

F230 1627 782 Pupariu, 'p,D I, ?Chloropid puparia 2, 2 Julids, I Polyde'lid "NI- 3

F230 1228 S7S [Rolan weill

flnni, 'p, 2 puparia I1NI- 2

Bag F206 "ED Frol Flots-

[Ph", 2, /5 pr, 11001 [Putrid]

F206 757 747

Thor"o,ha,ta lo.t,r" 132 puparia, I lineralised adult showed the very strong lid-tibial dorlal

bristles: 10 lulid and Polyde'lid frlgMentl I1NI- 132

338 F206 757 765

Acalyp1era1e sp,indet, I pupariul

F206 760 737

Cyclorrhaphan pupariul indet,lp,E I: Thor.coch,eta zo,terae 16 puparia "Nl- 17 F206 895 761

f.nnillp, I pupariuI: puparia indet,sp,C I, 'p,E I, Ephydrid 'p, c,30 puparia closely reseabling

Thoraeoehaeta zoster,e "NI- 33 F206 895 761 Se,topsl not.t, 9 pupae: Lilosinine 'p, 8 puparia fused into concretion "NI- 17 F206 1238 706

? TephrocllJ,.y. ruliventril 18 puparia, lfegageJj, !rufiplJ 2 puparia, 1Telo.erinl II,viplJ 3

puparia, Fanni, BP, c,30 puparia: 6 JuIid frlglents: 4 woodlouse headl "HI- 53 F206 1238 766

!fanni, ICilaril, 2 heads of adults, IIle and feule "NI- 2

F206 1238 766

flnni, 'P, 18 puparia: ?Por(l1/io Ie,ber, I tergal fraglent, I head fragment: Blapllpol I head

fraglent "HI- 18 F206 1238 766

Heleolyzid 'p, 5 puplria F206 1238 766),

fannia 'P, 17 puparil: I JuIid; !Porce//io Ic,be~ I head "NI- 17

F206 1606 740 7Chloropid BP, I pupariuI, pupariul 'p,D 2, Antholyiid BP, I pupariul "NI- 3

F206 1610 745 [Medieval weill

Thoracochaet, zOlterae, c,22 puparia "HI- 22 Bag * AI 1991 AREA I "ED, F207 frol FLOTS (Phlll 2,16 prl 1400J [Putrid]

339 F207 238 32

Scatopse notata, I pupa, Thoracochaeh zosteru, 4 puparia, rreio.erin. fi,vipes, 1 pupariul

"HIs 6

F207 252 35 1 siall lulid,

Tlloracocllaei. loster.e, 8 puparia, TTeIoleriRi flavipes, 2 puparia, fanni. sp, I pupariul

1 sull Julid

F207 257 53

LilOiinine sp, 1 pupariuI, HeJeo.yz. 'e.ptios., 10 puparil (I II): 1 Julid "HI- 11

F207 257 53,2,

Tlloracocllili. losterae, 3 puparil, fannil sp" I pupariulj 1 AgJenul brunneus: 3 linute Julids

"NI. 4

F207 287 41 c,6 siall Julids

ScaiopSl noi.i., I pup., pupariul indet,lp, F I, HeJeo.yza 'e'ptios" I pupariuI, Tlloracoelllli,

loster", 105 puparh, rreJollrin, fJ.vipes, 3 puparia, fanni. sp" 13 pupariai 6 slill Julids

"HI· 124 F207 287 41

TllorlCoeh,et, lo.ieraf, 1 pupariuI, f.nni'lp" I pupariuli 1 11111 lulidi Ar"diJJidiu. vulgare,

1 tergll frlg.ent F207 288 120

Thoraeoehlltl zosierll, c,30 puparia, f,nnil iP" 1 pupariul "NI· 31

F207 288 120

flnni. sp" 5 pupariai woodlouse IP" 1 tergal fragMent "NI- 5

F207 41 7 125

Thoracochaei, lo,ierll, 7 puparia, Fannil IP, I I pupariul "NI· 8

F207 420 126

Fanni. sp" 2 puparh

F207 47S(or 495) 173 [Medieval pit]

340 TlIor,cocllaet, zosterae, 5 puparia, ? TeJolerin, fJ,vipes, 2 puparh, Fanni, IP" 1 pupariull

1 sull Julid

Subtotal for Box Arel 1 Priority- 5 phases; 34 samples; 764 5peci~en5 AREA 2 Box Al 1991 AREA 2 PRIORITY * F521, 540, 579 Bag * FS21 AREA 2 MED F521 frol FLOTS [PhiSl 2,01 pre IIOO} [Putrid]

F521 2072 107 I indet,cllcified larval fraglent

FS21 2073 110 ?Dr050philid IP, I puplriul(ledian segMents only) MNla I

FS21 2084 108 ?Lilo,inine IP, I pupariuII(without end leglent)

F521 2138 118 indet,organic frag.ent

FS21 2140236

? Te/o.erin, fJ,ripls, 5 pupari., Drolophilid IP, I pupariuI, F.nnil'p, 2 puparia I'INla 8

F521 2219 193 , indet,cuticular fragMents

, F521 2219 193

1SpiJogonl IP, I pupar iUI, ?Hydrotae. dentipe., I pupariul indet: , i ndet cuticular fragMents I'INla 2

F521 2365 221

7S"teJJ'lp, 28 puparia (Iuperficially lik. T,zolter" but len flaUened, Inal seglent longer and

anal Itglents different), ?Bot,nophi/' fU9'X, I puplriul I'INI- 29

F521 2267 220

341 ?sepsid sp, 2 puparia, Finnia sp, 1 pupariul

F521 2365 221

1 ?"u5cid pup.riua, I pupariul indet,

Bag * F579 AREA 2 KED NB,

(PII.se 2. 01 prl 1I00J [Putr idl

F579 2500 289

TllorlCocll.eta 10sterae, I pupariuI, F.nni,sp, 17 puparia: I sllall Brlconid cocoon, I ike AplntlJIS

KNI:· 18 F579 2500 289

F.nnil sp, 1 pupariul

F579 2501 288

TlIor,cocnlet.lo.terll, 1 pupiriulll, ?Trachyope1la sp, I pupariuI, 1TephrocMilY. rufivlntris,

pupariuI, '.nni'sp, I pupariul "NI- 4

F579 2502 291

Fannia sp, I pupariul "NI: 1

F579 2504 292

1 indet,lineralised fragaent

F579 2510 298

"useid sp, I pupariuaCa cast in I calcareous nodule) "NI- I F579 2510 29B CF

[rist,Ji. ten,x, c,30 calcified larvae, at lellt 3 of ihe. in prepupal stage

nNI- 30

F579 2512 324

?CMorolY' lor.oll, 4 calcified larval, fT,phroclllllY. rufiventriJ, 1 pupariua nNI- 5 F579 2514 325

en/oro.y. lor,oJ~ I larva "NI- 1

342 F579 2536 299

Chloro.y. for'Ds~ 3 larvae

F579 2536 299 [plastic box]

[rist.lis ten.x, e,25 calcified larvae, sOlie in early prepupal stage but no sign of "hill" though

lany with pupal hornl indicated by sioothed elevations "NI- 2S

F579 2536 299 ["edifyal pit]

Cyciorrhaphoul pupariul indetlsp,D I; 2 Ichneulon or sawfly cocoons "NI- I Bag F540 * AREA 2 "ED 2

[Pllase 2,01 prl IIOOJ [Putrid]

F5'0 2137 103

fTelollgrin. fl.vipes, 2 puparia

F540 2143 167 Stratiolyiid 'P, 3 ,.all larvae, 1lelolerina flavipe&, 2 puparil "HI- 5

F540 2144 106

Lilosinine SP, 2 puparil "NI- 2

F540 2166 169

fTllollrin, fllVipll, 2 puparia, f.nni, SP, 2 puparh "NI- 4

FS40 2230 168 ["edieval pit/wllll

fTeJollrin, fllVipel, 3 puparia, flnni' SP, 7 larva Subtotal for Area 2- 4 phales; 30 lalples; 164 specilens

BOX AI 1991 AREA 2 "ED AREA 3 "ED' RO"AN

Bag ROlan Area 3

[Phasl ?i not ,t.ted} [Putrid?]

F--- 3664 686

f.nni.lp, (not F. ,,,Ilfi,) 1 puparhl end segMent "HI- 1

[Ph,sll,j] 120- 300} [Uncontalinated]

F784 3908 791 [Rolln pi U

343 ?Agrolyzid ap, 2 puparia(fraglents only)

(PIlaSI /, j5 250- 300l [Putrid]

F763 3912 758 [ROlan gravel working

Se.topse notlt., I pupa, ? TIJoII,inl (J.VipIS, • puparil, Tlloneoell.et, zo.t,r." I pupariull "HI- 6 F766 3904 748 [ROlan pit]

1 TeJolerinl (J,ripes, 3 puparia

Bag Area 2 Hed F503

(PhISI 2, OJ p" IIOOl [Uneontuinatedl

F503 2005 1 ["edievil pit 1Ephydrid .p, 2 puparil, Agrolyzid IP, 2 puparil

Bag Are. 2 Hed FS09

(PhlSl 2,01 p" W)O) [Putrid ?l

FS09 2023 5 '"edieval pit]

TilChioieptllp, 1 pupariul

Bag Are. 2 Hed FS10

[PhISI 2,0/ p" /lOOl [Uncontalinated]

F510 2029 36 [Hediev.l lined pit]

?Ephydrid IP, I pupariul (al before)

Bag Are. 2 Hed, "1nsectl frol f10ts" FSI3

[PhlSl 1, OJ p" 1I00} [Putrid]

F513 203. 10

Antholyi!d IP, I pupariul in perfect cond! tion though nerged, cOlpltte wi th III necessary dilgnostic lirvil characters: the only luch speeilen and probably I lodern conta.inant, "HI- I

FSI3 20S8 55

Thorlcochletl lOltlrl~ • puparil, Agrolyzid Ip. 1 pupariul "HI- 5

344 FS13 2088 93 FF [Medieval pit

fannia 'Clilril, 2 puparia) F513 2088 93

Thorlcochaetll0stera" 13 puparia(1 el), Agrolyzid IP, 1 pupariul HNI- 14 Bag Area 2 Hed FS37

[Phase 2.01 prl /I(J(JJ [Putrid] FS37 'BUG 11 [Hedieval pit

flnni, ffLliaril, I pupuiull Bag Area 2 Hed F568

[PhISI 2. Q/ pr, /lOOJ [Putrid?] FS68 2559 308 [Medievil pittwelll

I totally lineralized concretion, possibly centred on I f,nni, puplriul Bag Area 2 Med F570

[PhaSi 2.01 pre UOOJ £Uncontalinlted]

F570 2481 280 ACllypterate sp. indet. 1 fossilized pupariul FS70 2481 280.2 [Medieval pit] Antholyiid ap. 1 pupariul (in good condition) Bag Area 2 Hed F587

[Phasl 2.01 pr' W)OJ [Putrid]

FS87 2460 262 [Medieval pit]

1ThoriCochllta lo.ter", I pupariul

Bag Area 2 ~ed F604

[Phasl 2.0/ prl II00J [Putrid] F604 2615 368 [Medieval pit]

Sc,tOP" notlt~ 2 pupae, Thor,coch'ltl zOJtlr'~ 35 puparia (plus one totally lineralised larvl),

Fanni, 'elJaril, I pupariuI, I lineralized larva HNI- 38

345 Bag Area 3 Hed

[Pllas~ 2,03 /lOQ- 12501 [Uncontalinated]

F615 2657 413 [Structure ledieval 1]

?Kay,tiola sp, 1 pupariuI, ?Agrolyzid sp, I pupariul "NI- 2

(Pllase 2.33 pr' UOOl [Putrid]

F703 3345 619

/scllio/lpta sp, I pupariuI(linute one), ? T,lo.nina flaripes, I pupariul "NI- 2

F703 3507 608

Ephydrid sp, I pupariul (IS Ibove)

F703 3556 620 [Medieval pit]

Sepsid sp, I pupariul, ? rhor,cochaeta zost,r", I pupariuI(lineralized) "HI- 2

[Phase 2,34 pr' UOOl [Uncontalinated]

F704 3509 611 [Medievil pit]

?Ephydrid sp, 2 puparia

F726 3574 722 [Medievil pit]

7Ephydrid IP, 5 puparil (II Ibove)

Subtotal for Box AI 1991- 9 phasel: 24 Illplel: 98 specilenl

Total laterill- 28 phlsel: 1I2111plel: 1211 Ipecilenl

(4 LINDOW MAN II) 5 LINDOW MAN III

a) L"3, 0-10

£i.onill, lit, 6 larval head capsules, P,dicil •• l.t, 3 larval head capsule, StiJpon ,ublunatu., head of Idult: 1110 nUlerOUI fraglilents of Pl.teu•• ri. dileO/Of, !Cypllon etc" Ii tes etc,

"NI- 10 b) LH3 , 100-1 lOCI,

£i.oni. " lit. 2 hrval head capsulel, Pedid. I, lat, I larval head caplule: Iho nUlerous

346 frageents of Pl.teullri. but also with I elaterid larval end-seglent

"HI= 3

c) L"3, 110-1 IS,

li.oni'l, lat, I larval head capsule, Orthocladiine lidge, I adult thoracic fragMent

StiJpon lublun,tuI, I adult head capsule and 2 thoracesi also IfYrliCl ?ruginodil, I head capsule Ind

1 thorlcic fragaent, S(oJopo.tethullp, I pronotal frag.ent, Cicadellid 'p, indet, 1 adult facial luk

and clypeus "NI- 4

d) L"3 1I5-130CI,

li.oni. I, lat, 18 larval head capsules; also Hebrus ruficeps, 3 inCOMplete adults, and nUMerous

fragMents of Coleoptera "NI- 18

e) L"3, 140-150CI,E sect,

fPrionOClrllp, I pupal respiratory horn and 2 pupal tergal fragMents, U.oni, I, lat, IS larval

head capsules, Li.nophiJ, I, lat, 18 pupal tergiies, Tipulid IP, 12 pupal end seglenh(S lales •

fenln), Orthocladiine lidge, 1 adult thoracic fragunt, Oilophul febriJis, I adult fellie hnd

capsule,(Polt-occipital lobes luch hrger than in O,fIlOrltUS), Also Hebrul ruffceps, 5 inco.plete

adul ts for.iCl sp, I adult head capsule(worhr), ?AI Yliine wup, 1 adult head capsule aho I

cocoon and fraglents of lany adults of PJ.teu.,ril, "HI- 35

f) L"3,Frag,3, Skin and bonel frol elevator,

li.oni'l, lat, 1 larval head capsule, Aho a few indet, fibrous and nl11 Coleopteroul frag.entl,

"HI- I

g) L"3 AE Body laMple,

li.oni, I, lat, IS larval head capsules, Pedic i' I, lat, 2 larval head capsules, Dolichopodid 'p, I,

I adult head frag.ent, sp,b I adult head frlglent, IP,C I lall genitll leglent lost reseabling I

Rh'phiuI'P, but cannot b. related to any of thl currently known British species, Aho [Hebrus

ruficeps, 1 incoaplete adult and' nUlerOUI Coieopterous fragllnts- Pl,teu.arilltc, 1 "HI- 20 h) L"3,39,150B,Feb,B7, Frol start onl lOSS,

347 li,oni, '. ht. I hrval head capsule, Stilpon lublun.tul. I adult head capsule and 1 thorax. Also

IfYflic, fruginodis. I head of adult (worker) "HI· 2

i) l~3.lW/B1.30.3.88.

liloni,s. ht. SA hrval head capsule., P,dici'l. lat. 1 larval head capsule, Tipulid sp. indet. 2

lale pupal end segr.ents(as in sIMple e), ?Phryxi sp. I fragMent of adult head. This consists .erely

of the vibrissal angle with the facial ridges and the occipital dilation. In chaetotactic and

other details it very closely reseMbles lelbers of this genus but could belong to a related one).

Also lepidopteron sp. indet., 1 incolplete Hebrus rufi"pJ, 7 inCOMplete adul t Aphrodes Ip. I

fragaent of adult head capsul. ?Ps,"otlttix .p. I fragMent of adult head capsule FOffiCi ?Jlllni.

I adult head capsule (worker) luius '!niger, I adult head Clpsule (worked, l1y,,]'(. ?ruginodis. I

adult held capsule

of PI.tluI"i, and Ilny fragllnts of thil and other Coleoptera .tc. "HI- 88

j) l"3.lW/BI

Stygnocoril Ip. 1 head clpsul. Ind 1 pronotal fraglent Vlop. reti(ul.t,. I cOlplete for.wing]

Total saaples 9 Total Dipterous Taxa 11 Total "NI- 196

6 MINISTRY (LONDON)

Apart from the first one listed below, the samples were in extremely poor condition similar to those from the Leicester Causeway.

No data were provided by the archaeologist concerned.

86 (3998) (355)

Hello.yz, '"ptio" 6 puparil (2 unh., but not Illes), The,i" putri' 3 puparil( .. ),

Th"r"",h"t, zOlt,r" 975 puparia (c. 51 unhatched; in very good conditon, the ventral aabulatory

W.ltl clearly identical !lith confined llterial of the Ipecin), f.nni, .p. 8 puparil (totally fossil1 zed)

Sto."XYJ CI},Urln, 2 puparia (II)

348 86 (3998) (364)

Heleolyzl ?aptiosa 5 puparia (unh,), Tnoraeoenalta lostera, c,140 puparil (extre.ely poor

condition, at least SO per cent unhatched and fossilised just like the leicester laterial),

fannil sp, 20 larvae/puparia (unh,), Ifllseinl 1stabulanl 1 pupariul (unh,) "NI=- 166

86 (3998) (363)

Heleo.yzl ?eaptiola 1 pupariul (unh"subfos5ilised), Thorlcochlltl zosterl' c,350 puparia (allost

III unh, ,subfossilised), flnni. sp, 68 puparia (unh, ,Ill fossilised), Sto.oxys cllcitrln. 6

pup, (unh" ainerllised but c'ph,skel,lounted) "NI- 425 Total 5iaples 3 Totll Tlxa 6 Total "NI= 1585

7 NEWARK DITCH

Tipllia sp, 1 pupil fragMent, Ptychoptlrl,lbi.,nl 1 nle genihl segllnt (colplete), p, .inut,

1 lale genital leglent (coMplet.): 5 adult headl, 22 dorsul' and 3 abdolinal tergites, Psychodid sp,

3 pupae

SpnaeroClrl cur vip" II puparia (el) Ind I adult thorax, /schiollpt. 1pIISiJJI 19 puparia(3 unh)

Copro,yz, IP, 4S puparia (2 unh"clearly showing hind tibial spurs), Opa/i.a.in. JiJiputlnl 14

puparil (2 unh,lal.s in good condition), lilosinine sp, I 9 puparil(el), lilosinine sp,2 1 pup.riu.

(211, long), Agrolyzid sp, 3 pup.ria (el) (long dorsal postspiracular processes), Calliphor./lucili. sp, I puparial Inll spirlcul.r plate, Total taxa 14 Total "NI- 145

(8 OWSLEBURY>

(9 OXFORD: St.Budoc's Church)

(10 PLYr-DUTIO

11 RUDsrON WELL

RIUQO, 660118

Bibionid sp, I .1&11 larval head-caplule, 1SpI}obillp, 2 wings (bldly bleached), lilosinine IP, I

349 4 puparia, Lilosinine sp,2 3 puparia, Piophilid sp, I pupariul, ?Antholyzid sp.1 1 pupariuI; sp,2 1

pupariul (fraglent only) "HI= 9 660112

TipuJI sp. 1 pupal fraglent, Psychodid sp, c, SO pupae (Buckland, lc.), SyJ'IieoJI

cinctllllnutraJis ling, Seltopsl notit, 1 pupa (with larYil skin), lfegislJil1 !rulipll type· 1 pupariuI, The.irl sp. I wing, Thorleochllt, losterll 2 puparia, ?Antholyzid IP, 6 puparia, !Oeli, sp, 1 pupariul (rather like O,flbrieU) , Coenosiine sp. 1 pupuiul (51a11 sp. with fluted cuticle),

Acalypterltes Ind Antholyiids indet, 20 puparil "NI= 8S

Total taxa c.21 Total KNI= 94

12 SHIRLEY POOL

Bottol

Psychodid sp. 22 pupal wing-coverl, with tergal seglentl and 1 facial .ask, Chironolid Sp,l (large species) 5 adult thoraces, Chironolid sp,b (Iediu. species) 2 adult thorac.s, Chirono.id sp,c(siall species) 2 adult thorlces, Bibionid sp, 1 feili. Idult head, Tltlnoelr, .p, 1 fraglent of Idult head

?Chloropid 'p, 1 Idult thorax. [Also Corixids nUMerOUI fragMenh; /(icrO'llli, IP, IIny fragaents;

Saldid sp. frag.ent of I large spec in; Aphrophofl sp. several wings and body frlgMents; IflgillJul notuJ, .any wings ttc. !Pnryg,nl' sp. I fraglentl of Idult; !Forliel sp, 1 head capsules of I llrge

Ipecies: /(yrli" 'P. several adult heads; also 10le fraglents of Pl,teu.lfis. J /(NI- 3i

10- SCI

[6,rri'sp. levtrll frlQunh of htlielytrl; Apnrophofl.p, wing fragMenh, Phihenus spu.uiul wing frlglents; 1150 10le cOMpltte Oribatid lites, perhaps lodern contllinants)

0-20cI

Tipul id Ip. I frlgillent of larval head-capsule, [Aho Lepidopterous/Trichopterous sp.. several indeterlinlte larvil frlglents including part of head capsule which appears to be lepidopteroulJ.

"NI- 1 Total li.ples 3 Total Dipterous Taxa 7 Total "NI- 35

350 13 SOUTHAMPTON PIT)

14 TAUNTON

P" 19 (77) (120) (Tube 1)

Psychodid SP, 1 pupa, lScltopSi nol.e. 1 pupil tergite, Copro.yz. SP, 1 pupariuICu), Ischiolepta

'P, 7 pupuia, ThorICochiet. ZO.t,fdl 10 puparia, SplJobi. sp,l 26 pupariaCu)(long porrect prosp, horns), ?Speiobi. sp ,2 I pupariul(linute but lIith sili In prosp, horns), ",iolerin. fJ''1ipl.

44 puparia C7 unh), lilosinine SP, 2 puparil (tI) (very SMall speCies), ?/t.diz. gl.brl I pupariu,

"NIIl 94

(Tube 2)

TII,.i" putri.INe.opod. nitidui. 3 puparia

(Tube 3)

Sc.top., not.l. 13 pupae (el) (lacking larval skins), Lilosinine SP, 1 puparia (e~) "NIII 14

(Tube 4)

Sciarid 'P, lIing, ?SCltoPII not.i. I pupil fraglent, TIII.i" putri,INe,opod. nitidul. 2 pupuia, Sep'i'lp,C (sililler SP,) 1 pupariuI, ?SphllroClr.lp, 3 puparia, Lilosinine 'P, 8 puparia

(el) (very short prosp,horns), Tllor.cocll.,t. zOlt,r., 275 puparia ('0 unh"but adult features only relotely discenible in 6) "NIII 291

Clube Sl

!Tllor.cocll.,t. zo,ter., 5 puparia(e.), Lilolinine SP, 6 puparil(el), Antholyiid/Sclthophlgid SP, I pupariuI(eI), Hydrot.,. dlntiplI I pupariuICel), lfusCi dOl,.tiCi I pupariuI(II), Sto,axYI

"icitrln, 5 pup.ria(n) "HIli 19

(Tube 6)

Heleolyzid IP, 3 pup.ria(ea), Copra'Yl. IP, I pupariuI(el), ~"Clthopfl'9' SP, 1 pupariuI(u),

?Hydrot," SP, I pupuill fragunt "NIII 6

Total I •• plel' Totll dipteroul TaXI 20 Total "NIII 427

351 15 THORNE MOOR TRACKWAY

Thorne "oor Tree 1 Botto.

Pedicil rivos. IS large head capsules(unifor. size>: 2 udiu ••ized. [Also elaterid 'p, 1 larval

end-seglent and Ichneulonid 'p, I large cocoon (?Blnchus sp,) "NI- 17

2 Thorne "oor Tree I Top. Iroot and hole dregs l

Tipulid IP" 3 pupal end-Iegllnts, Psychodid sP,. 1 wing, Chironolid sp, 2 larval head-capsules,

Ceratopogonid sp, 1 pupal thorax very like that of Sph,ero,iu 15 figured in Slith (1989>: possibly

P,lpo,yi,. or closely related genul, 3 adua thoraces, K.yttiol. 'P, I pupariuI, ?E.pi. 'p, 1 alMost cOlpleh adult thorax. !8rachyop, 'p, I adult thorax, Pllloni, !err,ng I wing fragMent (COital cell only but extreaely like this species>, [Also a lUI of Coleopteroul, Helipterous and Hyltnopterou5 fraglents; al50 Oribatids- in excellent preservation] "NI- 14

3 Thorne Hoor Trackway Cranberry horizon

Pedici, riros. II larval hnd capsules U very large. 6lediul. I very .1111), Leptoce,in! sp, I puparial end segllnt, ?Antholyzid IP, 4 pupuia(el>j CordilurllPogonot, llircu. I pupariuI(el>.

?SpiJogon, sp. I pupariuI(el) "HI- 18

4 Thorne Hoor Trackway over Fen peat

Pedici, riroll 4 large larval head-capsules. Antholyiid sp, I pupuial end-Iegaent of large spec in

?SpiJogon,IColno.i, sp, 2 puparia(el), About size of Phloni, .ubvlnt, but end-seglent concentrically folded and longitudinally fluted, "NI- 7

5 Thorne Plat for. C 14 Birch Bark

P,dicil rivoll 2 larvil head caplulll, I of thea large "NI- 2

TDtal ,alplel 5 TDtal Dipterous TIXI 13 Total HNl- 58

352 16 WRCESTER BARREL LATRINE

S(atopse notlta I pupa(el)(lacking larval skin), 1Heleorayzid sp, I pupariuICel), Sepsids

puparia C6reig Ie,), Leptoelr. 'P, 7 puparia, Thoracoch"ta zost,rae 19 pupariaCel), LiloBinine Bp,1

7 puparia(unh,but illature), Lilosinine 5p,2 2 pupariaCera), Huscina 'st.bu/ans 1 puparial end- segment Total taxa 8 Total "NIs 40C+)

(178, 17b YORK)

WALES

18 HEN DOMEN

12e

Coenosiine BP, 1 pupariuI(el), Hydrot.,. dentJpI' 2 puparia, [Also nUlerOUI fragMent, of .illipedel

"Nls 3 135

!The.ill 'p, I pupariul(u), Sph.lfoelrl curvipu 1 adult frlglenti 10 puparh(u), Copro.yz. sp,

36 puparia(era), $pl/obi.lp, 4 pupariul(J unh), Lilosinine sp, indet, 1 pupariul (near Teiollrinl but

luller), /Jus" dOllsti" 8 puparia, Stoloxy, CI/eitran. 1 pupariul, "NI- 58

Total Taxa 9 Total "NI- 61

SCOTLAND 19 BUISTON CRANNOG

Descriptions of phlse, contexts and soils frol Kenward It 11,(1994)

PHASE I Group 2 [Wood litter on pri.ary .oundl

Context liS [Wood litter]

Cltirono,u. !plu.ofus, 1 JarvI! head-capsule, , adult thoraces; Hi/.ra chori", I ring,' IJrosophi/,

'P, 1 wing (very like IJ, .nd./usi".), [Also "ayfly sp" I lubillginal wing; Hyaenoptera Parllitica,

C, 10 fragraents.] "NI- 7

353 [Deposit Fine lilt with and large cut wood rnains] Interpretation The larval chironolid

shows this was an aquatic sedi.ent, Hili" and uyfliu also aquatic in hllture ltages, The

OrosoplliJi was adventitious, Exophilous cOI.unity with no evidence of hUliln presence Context 119 [Wood itter]

Tipulid BP" I pupal end-segllnt: Cflirono.ul ,'pJUJOSUJ, I adult thorax, [Also corixid and Trichopteron fragMents, MNI- 2 [Deposit Coarse organic IIhrill, with luch wood (cut-larks evident)] Interpretation Probabl y seli-aquatic, lake-Iargin habitat, Adults of aquatic insects present but no larvae, The tipulid pup I could be PrionoClrl, which can be found ..ongst losses etc, in shallow water, Exophilous cOllunity with no lign of hUlan presence, PHASE I I1NI- ~ PHASE II Group 3 [Sub.ergence layer] Context 110 [Sedilent over upperlost turf layer of Phase II [Corixid IP, I wing and 2 abdOMens] [Deposit Dark brown lilty organic laterill] Interpretation Probably In Iquatic habitlt but sinc.

Corixids are active fli.rs inconclusive without further evidence,

PHASE III Group S [Oulping over priMary loundl Context 108 [Slall spread of peat]

Bibionid sp, I thorax: 1 Antholyiid spp" I puplriul, I adult held: Icalypterate IP, I adult head, (Also fraglents of Heliptera Heteroptera and Paralitica Hy.enopteraJ "NI- 3

[Depolit Highly organic, with luch wood (cut-larks evident) and IOU charcoal) Interpretation

Wholly inconclulive, the only Dipteron being an adult Bibionid of which the larvae ar. hUlicolous but the adults very Ictive flierl, Ablenc. of Dipteroul i.latures re.arkable if thil WII a lite of active organiC deco,position,

PHASE III 6roup 7 [Early inhabitltion to the louth-east: occupation level IJ

Context S~ ["051 litter]

354 Sepsid SPol I pupal end-seglent; Hydrl/Ji,sp, I pupariuI; 'I1ntllo.yz.sp, 5 puparia, one which contained relains of unhatched adult also contained a Pterolalid parasite, NNI- 7 [Deposit "oss-peat with .Iall pieces of wood and a little sand] Interpretation Two dipterous taxa breed in grasses, lainly wetland speCies, and pupariate in the substrate, Sepsids breed in rotting plant latter, including lasses of algae, and in dung, Awaterlide habitat would be inferred but there is no unequivocal indication of hUlan presence, An exophilou! cOllunity, The presence of an adult fly and a Pteroulid parasite inside a pupariul is inexplicable; they were both facing forwards inside the pupariul 10 had not entered after elergence, Often insects seek Ihelter inside tlpty puparia, but thil had clearly not been the case here

PHASE III Group 8 [Early inhabitation to the louth-ealt; occupation level 2)

Context 50 [Fill over stone-built hearthl

/fuI" do.estic., 3 puparia [Aho Corixid sp, I thorax; Delphacid sp" I abdolinal lection containing lale genitalia; Trichopteron sp" I thoracic fraglent; Dry.ul sp" I head] "HI- 3 [Deposit Brown organic soil with little clay and sOle sand, Wood lacrofossil! (birch?) and charred/burnt bone were present] Interpretation Whillt there is Itrong .vidence of an hygrophilous assemblage, the pr.sence of /fuse, puparia show there wal a positive dOlestic refuse content Context 55 [Clay floor abutting hearth]

/fero'Yl. sp, I pupuiuI; /fuSCl do.estiCl, I pupniuI(II,); NlJin. ,.,let.. I pupariuI; f SpiJogon. sp" • puparia [Also I head of Cicadellid adult] "HI- 7 [Deposit Activi loil with live worls present, "oderat. brown li1ty deposit with wood «25 II), 10le

Itones 0300 I.), and bone frag.enh)] Interpretation The only il.. tures were the /fuse, puparia, showing that there IIlI a dOlestic refuse content, The other taxa are non-Iynanthropic hygrophiloul

.xophiltl Context 56 [TWig-layer in floor alsociated with hearth]

ChJoro.ri. lor.oll, I hrval skin (11101t cOlpleh); elpid 'p, I pupil tergal fragllenti

Acalyphratl IP" 5 puparia; S"tnoph'9,IBot.nophiJ. sp, I puparial end-segltnt; "usca dOlesticl, 5

355 puparia(el) HNI. 13

[Deposit A detri hI peat] Interpretation In that the do.inant taxon was /fu,CI, thil contained a

Itrong dOlestic refuse content, possibly, as suggested by Kenward It 'l, containing rotting hay-like

laterial, Chloro.yi. would breed in such laterial or in dung

Context 209

/fusCi dOllsiiCl, 14 puparia(tI,)

[Not included in Kenward It 'I" 19941 Interpretation Endophiioul eusynanthropic co •• unity

cOMprising /fU'CI only,

Context 233 [Fill of Itone-built hearth 51]

Prionoce" turcic. 1 pupa (ule); Tipul, (', vernalis, 2 pupae (fuales); Pi/lfi, dis(icoJJi., 1

adult head (identical to thil species): Eri(Jpt~rlsp, I pupal wing sheath; chironoMid sp. 2 larval heads-capsules; Iscflioltpt, !pu.iJl" I thorax; Copro.Y1' sp, 1 adult head: HydfllJi. sp, 1 pupariuM containing alYliine parasite; /flro.y:, 'p, I pupariuM; HUSLI dOllS tiLl, 2' pupariaCeM,) [Also'

Cono"lu. Inceps, 'forewings: I l,vesllJ, sp, 1 forewing: Psyll' ,Ini, forewing; 1Blltis 'P., 2 thoracel; trichopteron IP, 1 wing fraglent: , Phryg.ne, ?obsollt., , adult heads: parasitiC hYMenopterous, 6 .dult heads (incl. Alysiinel and Diapriid)] "HI- 3S

[Depolit Brown organic lilt with blue-grey inclulions, large •• ountl of plant l.crof05Iill, charcoal

Ind wood] Interpretation Th. I5subhge, being dOMinated by /fU.CI, supports the second of the luggestionl in Kenward It .1., (op. (it,) thlt this Wil Ian occupation deposit to which aquatic insectl had been added by flooding or redeposition of waterlain sediMent- The HOMpoteronl Cono.t/us and IIvtsell, inflr rushes (Jun(uJ spp,) Ind grnses respectively, whiht alders wert Ilso preunt

(ie, P'rll, .Inil, AlYliines Ind Diapriidl Irt parasitic on Dipterous larvae and puparh

Context 23' [Sand below hearth]

l f/f,yetiol'lp" I Ifllse pupariu. : Cyclorrhaphan 'p, 2 puparial front upper Upl only (/fusCilize or larger) [Aho ?Blltig 'p" 1 thorax; Corixid sp" several frag.ents: Trichopteron 'p" I thoracic fragMent: parlli tic HYMenopteron, I head; HYfliCi 'P" I head] "HI- 3

[Deposit -Dark brown peat with fraglltnts of bone evident I Interpretation Too Ii ttl. dipteroul

356 laterial but probably II suggested by Kenward It .1.(1994). An exophiloul, non-synanthropic

Isserablage

Context 235 [Clay below hearth]

Priono",. tureic., 1 adult head, 1 pupal end segllent; Tipulid sp. (?Prionoelr'), 5 larval heads­

capsules(various lizes); Ulosinine sp. I adult head; Ephydrid sp., I pupariul containing Alysiine

parasite; £i.nelli, sp. 1 adult head: Acalypterate sp" 5 pupal fragments [Allo ?Silti, sp" several

thoraces; /'Iyst,cUes sp. I thorax; Trichopteron sp, I adult head; £l.sluch,I£J.s.ostethus, I

Icutelhr fragMent; Saldid 'p" I thorax: Philaenu, 'pu."iu.. sevenl thoracic and wing fraglents:

I thorax: /dioeeru, IP" I wing: lireSlll• • p.. 1 wing: Psyll id Ip .. I wing; several head capsules of Proctotrupoids, Braconids and Pterolalids; For,i" Je •• ni, 1 head] "NI-" [Deposit Brown organic Ii It wi th blue-grey inc Ius ions, large lIountl of plant IIcrofossill,

charcoal and wood] Interpretation A rich hygrophiioul non-synanthropic exophilou. assemblage

typical of a waterside habitat with wet Iud and lilt ltc, 6ranland with birch and carrhnds, No

Ividenc. of Iny hUlan presence

PHASE III 6roup 9 [Early inhabitation to .outh east; occupation level 3]

Context .8 [Clay floor abutting hearth; over context 60]

Hlllitopot.IChr"opf 'P. 1 pupal tergal fragaentj toxo"" 'p, 1 pupariuM: !Antllo.yr. IP, 1 pupariul: ?Agrolyzid sp. 1 pupariuI; Hydrllli. IP, I pupariuI: /'Iero.,I. 'p" 1 pupariuI; /'Ius"

do.esU", 3 puparia: '~'piJogon. IP, 1 pupariua; Helin. freet., 1 pupariuI: Ph,oni' '.ngelie" puparill frag.ent [Allo Corixid SP, 1 pronotul: /dioceruf 'P, 1 forewing: /'Iyr,i" !5P, 1 head] "NI- 12

[Deposit Chy floor hyer with uh, thin fibrous .eall and burnt bone fragMents Interpretation A predolinantly exophiioul hygrophiioul Isselbllge with indicatorl of decaying lodden grllses, organiC

Iud Ind drier hUIUI Ind folilge of Silicicele, but with Huse. Ihowing a presence of dOMfstic refuse

Context 71 [Wood chip. Ind twigs]

£rioptlf' Jut" v, tllnionot., 1 wing fragMent; SCltoPII notltl, 3 pupal fngaentl "HI- 4

357 [Deposit (n above)] Interpretation Inconclusive though SCitopse luggests lOll! declying organic

utter

Context 207 [Peat and twigs- I floor laierill 1]

Li.osin•• iJv.Ue., 'puparia: ?Tr.ehyopIJJ. sp, 1 pupariuI; /fuI" dO'~ltj", 5 puparia [Also

PhiJ,enus spu,uius. 1 forewing; COnDIe/U' .nceps. 5 wings; IlVfSfJJ. sp, 1 forewing; P,yJ}.

ef, pu/ehr., 1 adult head; Pteroulid sp, 2 heads; Ceraphronid sp, 1 head; Diapri id sp, 1 head] PlNI- 10

(Deposit Dark brown to black hUlic lIaterial with luch wood, I little bone and charcoll Ind 101e

hazel nuts] Interpretation The dOlinanc. of /fUSei and £j.osin. Itrongly luggest this was a floor but there was I Itrong presence of Ho.opterl Iisociited with grllses and rushes, poslibly brought in with hay Context 213 (Redeposited pelt extending westward frol hearth]

?Spilogona IP, 1 pupariul [Also FOrliCi /11I'nr~ 1 head] "MI- 1 [Deposit Detrital organic later ill] Interpretation Probably Iud or silt deposit with no lign of

hunn i nfluenc. Context 215 [Wood chipi and twigs",Wal it I floor depolit?]

/fuse. dOIl.tiCl, " pupariaC.I) "NI- 11

[DepOSit Wood frlg.entl, chlrcoll, leeds and hazel nutl were evident in thil I •• ple]

Interpretation Th. presence of /fusc. Ilone shon thlt this was an endophilous .ulynanthropic Isselbllge, Al.olt certainly a floor deposit Context 223 [Underlying helrth 32",Was it I floor depolit?]

/fUI(, do.estiCl, 1 pupariuICII,) "NI- I [Deposit This lilty .upl. contained wood Ind very fragaented bone IIterial] Interpretation The prelenc. of only I lingl. /fu.e.luggelts thil wal In Indophilous lusynanthropic ISle.bllge, probably I floor

Context 22. [Kixed dump of 1051, peat and twigs",Wal it I floor lurface?]

IAntho.yz. 'p, 1 pupariuICel,l; /fu." dOllltiCl, 1 puplriulClI,l PlNI- 2

358 [Deposit Woody peat] Interpretation NlJs" indicates I presence of domestic refuse, whilst

Ant!lo,yzi would infer presence of I species of grass (eg, fllrlg.l'tll, flll/.N's, Oactyli. etc,),

Possibly frol a floor surface

Context 227 [Mixed dUMp of lOSS, peat and twigs" ,Was it a floor lurface?]

H""topot.IChrysops sp, I pupal fragMent; 1:'''p,/obi.lp, I pupariuICel); fTrlChyop,J/. lineafrons, 2

puparia(et); '£i.osin. lilv.ti", 1 pupar iUICn):: t Antho.yzi sp, 1 pupariuI; NlJs(, (/ollltia, 643

pupariaCea, )" 2 adul t thoraces [Also Corixid sp, I thorax; Conoll/us Inelp5, 8 forewings;

Strophingi. lficlI, I forewing; "yraica 'P, I head; 2 Alysiinu and 1 HYlenopterous cocoon) "NI- 649 [Deposit Detri hi peat with luch wood and SOli hazel nutl] Interpretation The huge breeding

population of Nusc. and virtual absence of III else indicltes In endophilous eusynanthropic asseMblage such IS one would expect frOI I fouled dOlntic floor, Tr.chyop,J/. Ind Li.o,in. could share such a habitat, whilst the Tabanid pupa could have been brought in with peat or rush litter along with the Corixid and Conol/,/u, .ne,p5, a /un,u~feeding HOlopteron, AlYliine Braconidl parllitill Dipterous IIrvae, whilst Strophingi. ill Plyllid bug associated with Eriel

Phlse III "NI- 794

PHASE IV 6roup " [Occupation to the north, First hearth]

Context 327 [Ash and burnt bone]

TrichoCl" sp, 2 pupal wing-sheathl; Di/ophus sp, I thoracic frlgllnt: SCitOps, no t.tl, U pupae( II); f,il. (Chll"p,i/.) 'p, I pupariuI: fT,phro,hlilY. 'p, I pupariuI; 18p,/obi. sp, 1 pupariuICII); rrrlcllyop,lI. linnfronl, I pupariuICel); fLz'.olin. sjJv.ti", I pupariuI(el); Ephydrid

IP, I pupariuI; Calliphorid IP, I puparhl .nd Itg.ent; Acalyptente 'p, I pupariuI; Euu.syphorl

?CYInic%r, I pupariua; Neolyi. cornicin., I pupariuI; NIJIC. dOleltiel, , pupar!a(el), 1 adul t thoracic fragMent; N,IopIl19IJS ovinus, I head [Also CDnoll/IJS .n"pl, 2 wings: Ichneulonid IP, I head; Elaterid IP, I larval end-legMent] "NI- 31

[DepolH Dark brown organic IIUer wi th lIall ..ounh of stones] Interpretation A COMplex of

IIteriall including rUlhes, ulb.ls (or COlPOSittl) conhlinated by decaying vegetablt utter, co.-

359 dung and perhaps sheep-dung, Sheep or fleeces indicated by HeJophagus ovinus, The low count for

HusCi do,estiCi but higher nUlbers of Se.topse not.t. and rr.ehyopeJi. (collpared with F312) suggests that although ferlenting organic latter existed in both 5a.ples, the laterials were different, This lalple possibly consisted Ilinly of ungulate dung whilst F312 lay have been .ainly of hUlan origin,

PHASE IV Group 15 [Second hearth] Context 12 [Spread of ash, part of complex around hearth]

Tipulid IP, I pupal fraglent; ?P.ii. (Clllllep.iJ.J SPol 2 puparia; HeJeo.yZl ?borelli., pupariulj Ephydrid sp, I pupariuI: !fusCi dOIl.ticl, II puparia (10 II) [Also Cono,lius .nceps, 2 wings: Iive"Ji. sp, I wing: Proctotrupid sp, I thorax; HYrlie.sp, I head and ali trunk 1

MNIli 1615

[Deposit Brown sandy soil with charred bone fragllents Earlier described II "hearth and food (dolestie) debris in early roundhouse, with ash, charcoal and burnt bone] Interpretation AcOlplex of sodden !un"/I and gran liUer with ulbell (or co.posites), rich hUIU5 and ferlenting organiC laterial including faecal and anilal protein cOlponents With Hu,c. predolinating the latrix shows a strong endophilous eusynanthropic tlllllent and hence the deposit us lost likely a fouled floor deposit, Non-synanthropic elelents probably introduced with the floor litter laterial or, as Kenward

,t .1,(1994) suggest, subsequent strays frol wetland habitats adjacent to, or luperillposed upon, the lite, The conclusion in Kenward " II. (op. cit. p,34) is supported by the Dipterous IIterhl, except that the 'House fauna", which includes HII.el and H,Jeo,yz~ greatly do.inates the Diptera,

PHASE IV 6roup 16 [Associated with upper.cst hearth]

Context 312 [Thin co.pacted Ipread of Itr.w]

Se.topsi notlt~ 10 pupae(laeking larval skinl but one pupa contained lall genitalia 10 identity verified); li.osin•• ilv.tiCl, I pupariuI; rrlCllyopIlJ. 'coprin. I pupariu.; r,lollrin. fllrip,., I pupariuI; ',il. (Chllllp.iI.J sp, I pupariuI: HusCI dOll,tiCl, 553 puplI'ia(S45 el): Hllopll'gu, ovinu, I head Ind thorax; C.Jiipllo" ap, I puplrial end leglent [Aho Antlloeoris sp, I head; Deitocephllid

360 Sp, 1 wing; ConoleJus In(ep~ 4 wings; Lepidopteran sp, 1 larval skinl "NI- 569

[Deposit Dark brown organic laterial Earlier report descibes thil IS Ilayer of cOllplcted Itraw(f reeds)· straw floor litter in early roundhouse] Interpretation A deposit totally dOlinated by ferMenting organic uterial wi th at least a trace of ani .. l protein which suggested fouled floor­ laterial and(or) stable lanure; predolinantly In endophilous, eusynanthropic aS5elbiage, Subsidiary contenh inc luded grasl and rush-Ii tter, sOle herbaceous (ulbell or cOlposi tes) and sOle carrion, Sheep indicated by presence of the Sheep Ked HeJopnagus orinu~ Context 316 [Extensive llyer of brush, under 312)

fHayetioJ. IP, 16 "falle puparia"j SCitopse not.t., 1 pupa; /schioJept. sp, 3 puparia(III);

'ii.osin. li/v.ti", 1 pupariuI(el); HusCi do.estiCl, 2 puparia(\ II) "HI- 23 [Deposit "A woody peat' Earlier report describes this IS a layer of brash in thick bundles-' sub­ floor twig litter in early ('4th century) roundhousel, Interpretation Possibly predolinantly grlll­ litter, strongly inferred by the do.inant taxon, but contalinated by faecal cOlponent and including ferllenting organic latter frol house or stable floor;

Context 341

ScatopSl notata, 10 pupae(lIith hrval skins); /.,MoJept. !pulilJ., 9 puparia(one containing

Alysiine parnite), I adult head; ; TeJo.erin. fJ.vipel, 5 puparia; !SpeJobi. sp, 6 puparia

(incolplete); Tr""yopeJi. 'coprin., 34 puplrh(30 n)j !AntholYz. sp, 2 puparia; Acalypterate sp, pupariuI; HllsCI do,estiCl, 8 puparia(n) (Also 1 Deltocephalid wing) HNI. 75 [Deposit Dark brolln peat lIith luch wood (large .Iounts of which had never been cut), hazel nuts and 10le charcoal, Siall .Iounts of quartz grains, slndstone Ind fine gravel lIere present'

Earlier report .tatel it 1111 floor of early roundhouse) Interpretation AcOlplex of grlss and /uncuJ litter dOlinated by In ISseMblage including eusynanthropic endophiln which bred in ferlenting orglnic I.tter and dung, including IOle ani.al protein accullulated in I dark .itultion

The conclusion in Kenward It .1,(1994) that thil WII In occupation lurface is supported by the dipterous Ividenc., but not. 'that the 62 li.o.in•• iJv.ti(. pupl1'il were not frol this supl. but froll 402

361 Phase IV Total "NI- 714

PHASE V Group 21 [Consolidation)

Context 39 [CoMpressed reeds or straw", II there a human influence?]

lfusCi do.estiCl, I pupariuI(ea,) "HI- I

[Deposit The unhulified peat contained no charcoal or wood) Interpretation The only Dipteron

recovered, IflJs" do.estic., shoWl irrefutable evidence of hUlin occupation as it is eusynanthropic

Context 41 [Dark fibrous roots; redeposited turf, to 20cI thick, extent about two square

letrel",ls there hUlan influence?]

Liloniid 'p, I adult head: 1Li.osin, Si/VltiC', I pupariuI; lfusCi do.estic., I pupariuI:

lfyd.e.IHyospi/, sp, I pupariul [Also Conole/us ,nceps. I wing; Trioll furticlI, I wing)

"NI- 4

[Deposit Peat, rich in 1051 and containing hazel nuts and slall bone frlgMentl]

Interpretation HusCi again gives clear evidence of hUlln presence Ind lfyd",llfyospil, and £i,olin,

suggest I hecal cOlponent Could be sUe of reduced hUlln activity II suggested by Kenward It II. ( op. "t. ),

Context 45 [,.11 there I hUlan influence?)

frionoclr.ITJpul., 2 larval head-capsules, 3 pupal fraglents, I adult abdolinal sternite:

Oilophu'lp. I head and I thorax (of fellle); Rhlg'OC/11i coolii, I pupa (containing adult lalt);

ACllypterlte spP. 4 puparia

,Ini, I forewing; E hterid IP, I larval Inal Itglllntl "NI- II

[Deposit A detri hi peat with 101e large wood fragllnts] Interpretation A hUlicolous, non-

Iynlnthropic lSulbhge frione"" would indicate a wet depositional lite but the specillenl could equally have belonged to one of the hUlicolous TlpU/' Ipecies, Aphrophor' indicates proxility of

Ainul glutinoll or a S,lix specitl but, being an adult, it could be adventitious, No Dipteroul evidence of any hUlan presence

Context 308 [Source unknown]

Tipulid IP, I pupal hrgd fragllent; £ristllis IfbustorlJ.I,busivus, I head Ind thorax; Scathophagid

362 or Bot.nophiJi 'p, 1 pupariul(u,): ?Spilogonl 'p, 2 pupilriil(u), [Also Cono,eJII' sp, 1 wing]

HNI= 5 [Deposit This dark brown peat contained large quantities of wood, sOle with cut-.arks evident· Earlier report describes this as deep organic deposit (lOSS, twigs, reeds, branches, peat)-organic

dump] Interpretation Organic Iud wi th sOle hUIU5, 'uncus li thr and a possible traci of hecal

laterial (if Sc.tnopn.gl was present, but this uncertain) Otherwise I non-synanthropic Isse.bllge Context .02 [Source of later tal unknown]

Prionoeer,ITipuJ. sp, 1 larval head-capsule); Ceratopogonid sp, 1 pupl(front half only); SCitOpsI

"Dtiti, 1 pupa: ?11.Yltioli sp, 2 Ihlse puparia'; Sepsid sp, 1 pupariuI; Copro.yz. sp, 1 pupariul

(containing lale genitllil); li.osini .ilv.tiei, 62 puparil(61 II); Sphaerocerid sp, 7 puparia (very

like TeJo.erin. fJlvipfl and Herniosi"i bequ"rW; OrDsopltili '.uDobSLur., 1 pupariuI(u):

1SCltltophigi 'p, 3 puparia(eI): HydrDt.,. !IJbipunct. 3 puparia(ea); ?SpiJogO"' sp, 1 puparial end

segMent [Also FOrMiCi ?J1•• nJ~ 3 heads: Proctotrupid sp, 1 thorax] HNI- 84

[Deposit Dark brown peat with luch wood (solie cut) Ind 10le Ion' Earlitr report describes thil deposit IS a dUlp of lediuI- dirk brown fibrous peat with lOSS and wood-chips- a levelling or consolidating dUlp] Interpretation Traces of hUlus and grass-litter but dOlinated by dung and (or) fouled, sodden plant litter, which had probably accululated in a dark place- possibly I stable, UngUlate dung is inferred with at lost very low ferlentltion, the therl.l levels probably below threshold for Husc. dOll,tiCl, which wn replacld by li,o,in, si/v,tiel IS the IOlt nUlerous speciel

At lost I leli-synanthropic asselblage,

Context 310 [Sourc. of I.terial uncertainl

Tipulid .p, I pupal fngunt: Scatopsid sp, I pupal end segllnt: H.Il11topot.ICnrysops sp, I pupil anal segllenti £"pisIRhl.pllo.yil IP, 1 pupal abdolinal section: ?Antholyiid sp, 2 heads (with liny orbitals and strong crossed frontals)j Hllin. ,v,ct., I pupariul [Also heads of Lyguid bugs and HYrJiCi sp,l tlNI- 7 [Deposit Very dark brown fibrous peat with sull pieces of cut wood] Interpretation a lixed waterside Ind drier hUIUI non-synlnthropic Isselblage with absolutely no evidence of hUlln presence

363 Phase V Total "NI- 112

PHASE VIII [Not statedl

Context F2 [Not Ita ted]

Prionoee"lTipuJi sp, 6 larval head-capsules: Scatopsid sp, 2 pupae(no larval skins>: Chioro.yi.

(Ofl~'S', 1 larval head-capsule; Ceroxy. urtiCle, 7 puparia: P,iJ, (Ch ••• ep,iJ,) sp, 62puparil;

#eJe~'''Yl' 'serflt., 19 puparil; ?Leptoee" sp, .2 puparia; Hernio5in. be

TeJollerin. f J.vip", 88 puparh; ?Piophi I id sp, 1 pupariuI; ?Ephydrid sp, 60 puparia; ?Drosophi lid

sp, 1 pupariuI; Antholyi id sp, I pupariuI; PI/Ioni. ',5tr, I pupariuI; lfe/opl/aglJs OVinIJ5,

I thorax; Calliphorid, 1 puparial end-segment [Also 3 Ory,u, sp, 3 thoraces; IfYfliCi 'p, I head] MNI- 307 [Deposit No infor.ationl Interpretation Probably the lost cOlplex of the Buiston Crannog sl'ples, this wa, dOlin.ted by two cO.lunities inferring different origins, There was a vlry rich synanthropic assemblage in which the lost abundant specie, was thl therlophiloul, endophiioul necrophage,

Tllo.erin. fI,vipe" Together with #erniosin. bequurti and #el,ol1' thesl inftr a dark breeding

environllnt, such 15 would be ideal for /fIJS" do.estiCl, Yet that species wn wholly absent! Ceroxy.

urtical, the Ephydrids and !Lepio"" IP, teltify to a wet, though ltill foul environaent luch II

occurs in the organiC Iud around aanure heaps or cen-pits, PI/.oni. 11 priaarily hUlicolous and uy

have entered the Ivstel initial IV alongst litter, or lubsequently during the hUlifying proceSl, A

possibl, scenario for thil deposit wal thlt it WII a floor deposit which had been dUMped into a

larshv Ire., The curioul absence of /fuse, however, in view of its abundance in other horizons, lav suggest that it could have died out locally before thil laterial was deposited

Phase VIII Total "NI- 307

PHASE? [No inforlationl

Context 005 [No inforl.tion]

Oilophu. 'P, 2 I.le thoraces "NI- 2 [Deposit No inforlation] Interpretation Inconclusive line. OiJopl/u. IIln an very activ. fliers P,at abov, Irch [No inforlation]

364 Liloniid 'p, 13 larval head-capsules; Dolichopodid sp, I pupal fragMent; loxo"r. BP, 2 puparia:

Sepsid sp, 1 pupariuI; Copro.yu sp, 1 pupariull, 1 lIing; rSpiJogon. sp, 8 puparia [Also Ory.us sp, 3

pronob: 2 Conolt,Jus tln"ps, 2 forewings; "v,s,ll,sp, I forewing

Elaterid sp, 3S larval end-segments] [Deposit No inforMation] Interpretation Could be a natural waterside alselblage, there being no

unequivocal indication of hUlan presenc., KNI- 27

TOTAL KNI: 1979 Taxonolic List of Diptera frol Buiston Crannog

Priono(lr, turcic, 1(233); 4(235) KNI- 5

Priono(lrilTipul.sp, 2(45); I( 308); 1(402); 6(2) I1NI- 10

Tipul' ?vernlJiI 2(233) KNI- 2

Tipulid 5P, 1(119); 1(12) KNI- 2

PiJui. dilCi(ollil 1(233) KNI-

frioptlrl Jute. t'lnionot. 1(71) KNI-

frioptlrl BP, 1(233) KNI-

Lilloniid sp, lUI); 5(POA) KNI- 6

Ceratopogonid BP, 1(402) KNI-

Chi ronoltu. ?pl UIIOSUI 4( 115): 1(118) HNI- 5

Chi ronolid BP, 2(233) I1NI- 2 Tricho(,,, 'p, 1(327) I1NI-

DiJophullp, 1<327>; 1(45); 2(005) HNI· 4

Bibionid 'P, I (J OS) "NI-

SCltopSl not.t. 3(71): 14(327); 10(312); 10(341); 1(402); 2(2) KNI- 40

Rhlg.o(I", cooii 1(4S) KNI·

?""yetiol, 'P, 1(234); 16(316); 1(402) KNI- 18

ChJor".yi. 10rloll 1(56): 1(2) KNI- 2

Hlllltopot,IChry.op. sp, )(AS) ; 1(227) "NI· 2

365 Ellpid SP, 1(56) MNI­

Hilara ehorie, 1(115) MNI­

Dolichopodid SP, I(POA) I1NI· fristdJis .busivlls/,rblJltoru. 1(308) I1NI­

Ceroxy. urtical 7(2) tlNI- 7

LoxoCl" SP, I (A8); 2(POA) I1HI- 3

Psil,(.g, Challepsil,) SP, 1(327); 2(12); 1(312); 62(2) I1NI- 66

Sepsid sp. 1(59); I(02); J(POA) tlNI- 3

",phroeh) ••y, 'P, 1(327) I1NI­

Hlllo.yz, !borllli' J( 12) I1NI·

H, 'serflt,/eaptios. 19(2) I1NI· 19

Copro.yz, SP, 1(233); 1(402); I(POA) /'INI· 3 i.ehioJept. ?pusiJJ, 1(233); 3(316); 9(41) tlNI· 13

Llptoelr, SP, 42(2) MNI- 42 £j,olin, .iJv,tiCi 4(207); 1(227); 1(327); 1(312); 1(316); l(Al); 62(402) MN[- 71

Herniosin, bequilrti 13(2) I1HI· 13

fSpIJO/Jil Sp, I (227); 1(327); 6(341) I1NI- 8 'rachyoplJJ, !eoprin, 1<207>; 1(312); 34(341) I1NI- 36 r. 'linel/rons 2(227); 1(327) I1NI- 3 ',lo"rin, (l,vipl' 1(312); 5(341); 88(2) I1NI- 94

f',Jo.erin,/filfnio.in, 7(402) tlNI· 7

Ulosinine ap, 1(235) I1NI-

Piophil id 'P, 1(2) I1HI-

!Antho.yz. SP. 5(59); 1(48); 1(224); 1(227); 2(341) I1NI- 10

Agrollyzid sP. 1(48) I1NI- 48

HydrlJji' Ip. 1(59); 1(233); 1(48) I1NI- 3

£j,neJii' 'p. 1(235) I1NI-

366 Ephydrid sp, 1(2351: 1(327): I( 12): 60(2) "NIII 63

Drosophila sp. 1<1 IS): 1(402- !suDo/)seuri): 1(2) I1NIII 3

/fero.yza sp. J(551: 1(233): 1(48) I1NIII 3

Acalypterate sp. 1(108): 5(56): 5(235); 1(341): 4(4 species- 45) I1NI- 16

S"thophag'sp. 3(402) : I1NIII 3

SCithoph'9.IBot,nophiJ, Ip. J( 56): 1(308) I1NI- 2

Antholyi i d !lp. 1(108): 1(2) I1NI- 2

Hydrotae, !aJbipunet, 3(402) I1NI- 3

EudasypMfI ?'YanicoJo!' 1(327) I1NI-

Neo,yi, eorniein. I( 327) I1HI-

/fuse, dOllstiCi 3(50): 1(55); 5(56); 14(209): 24(233): 3(48): 5(207): 11(215): 1(223): 1(224);

643(227); 4(327); I J( 12): 553(312): 2(316); 8(341): 1(39): 1(41) I1NI-1291

?SpiJogon, sp. 4(55); 1(48); 1(213): 2(308): 1(402): 8(POA) IiNI- 17

/fydae'l/fyospi Ja Ip. I W ) I1NI-

Helin"v,et, 1(55): 1(48) I1NI- 2

PI/,oni, ',ngeJi,,, J( '8) IiNI-

P. ferl'ln. 1(45) I1NI- p. (1.lh.) Ip. 1(2) MNI-

/f,Joph,gu.ovinu. 1(327): 1(312): 1(2) MNI- 3

C,Jllpllori !lp. 1(312) MNI-

Calliphorid sp. 1(327): 1(2) I1NI- 2

Cyclorrhaphan sp. 1(234) MNI-

Total Dipteroul Taxa Tohl "NI-1981

(20 DUN VULAN)

367 21 LOCH DRUIDIBEG (SOUTH UIST) [Organic silts from islet in lake]

I 0-2cI.

Tipul. ?oler.".lp.ludol. 2 pupal fragMents, I larval head-capsule; I.i.oni. ftrivitt.t. I pup.1

~ing-(ov",' !l.iMOni. I. lat. sp. I larval head-clpsule (very lIall); I.i.nopnil. ?eigeni nUMerous

pupal fragMents, including pupal horns, pupal flcial-shield containing head of Idult; Psychodid sp.

I pupal skin, I pupal wing-cover; Chironolid sp. , larval head-c8psults, pupal wing-coverl, leveral adult thoraces and wings (at least two different species); Bioio ?nigriventril2 wings; Iciarid sp. 1 wing; Tricllin. ,?cI.vipes 1 wing; Elpid sp.l (7 Tricllin.) 3 pupal frigMents (with short tergal spines)i npid sp. 28 pupal fragaents; npid sp. 2b pupal fragllents; Hilar. clloric. I wing; Rh.plliu. sp. 2 inCOMplete pupae, I containing Interior half of adul t nle; C.llpsicnellus ?Joripes I wing;

I.onchopterl lute, 1 wing (1I1e), 1 ldult head; l1egase/i, sp. 2 puparia(tI), 1 wing; Coelop. 'P. I

Idult head; Hydrelli. sp. 14 puparia(u); Sc.teJi. lubgutt.t. 1 wing [Also Plocopteran IP, I head capsule; Anthocorid IP, 3 adult heads; Corixid sp, indet, I wing; Deltocephalid sp. I wing: /.ivi, luncoru. 1 wing; PsyJl.lp. 10 wings, 1 Idult head; ruoeroiachnus vi.in.JiJ 1 wing; lepidopteroul

IPP, 5 larval head-cIpsul ..; U.neplliJus !vitt.tu. 1 wing; l1ystac ides azurea cOlplete gen1 h I capsule of 1lI1e adulti Tinodes .aeneri 2 wing fragllentsi COMplete adult nit genital capsule;

Coleophroul spp. 19 larval head-capsules (It least two different species); AspiJot. sp, I wing; leveral 'Iall Alysiine Idult heidi, pOllibly of saRe species,l HNJ· 42 b 2-6cI

U.nophiJ, ?eiglni lany pupil fragll!nh; Liloni. ?trivitt.t. 2 pupaei Liaoniid sp, 1 larval held-clpsule (closer to L, .odlli. than IIcrOltig,,); ?l1olophilus 'p, I Idult head; Chironolid sp, larval head-clpsule: Bibio fjolllnni./nigriventri. I wing, I ferille Idult head; Eapid IP,

C? rrkflinllp,) 2 pupae(with short-spined tergll urgins); Elpid sp, 2 1 pupal fragllent; hpid sp. 3

(!Hil.,. 'p,) " incolplete pupae (with long tlrgal hair-fringe); Oolichopodid sp, I incolplete pupa containing adult; COllap. frigid. 3 wings; 610.Yl1 b.ll(horskii I wing; HydreJJi. 'p, 2 puparh(tI)i Hy.din, ?gultlt. I wing: C.JJiph",. IP, 1 wing frag.ent; l.phne "udll. 1 pupariul containing coaplet. Ille genihlia: Antholyiid IP, 1 pupariul(lI) (Iililar ones in other horizons);

368 Dipteroul sp, indet, I adult thoracic fraglent [Also uyfly IP, 1 nYMphal wing-cover; ~"'JduJ. 'P, I

wing; Corixid sp, I forewing: PIliJ.enus spu,.rilll I wing fragllent (clavus): Psyll. sp, " wings:

Ify"i" sp, I wing: TuberoJlCnnlJ' vi,in.Jis 2 forewings: I hindlllng: Lepidopterous sp, 3 pupal

fragMents; Tinodes r.eneri 1 wing fragMent] [Seell to have no record of nUlbers of 101e taxa)

c 6-IOcI,

Tipu/. foJerICII/p.JlJdos. I pupal fragaent; Li.oni. sp, 4 larval head-capsules; li.nophiJ.

',eigeni nny pupal fraglents; Chironolid sp, (as in a) several larval head-capsules, 1 larva found cOlplete and in perfect condition as in lift- presuubly lodern contuinant; Bibionid sp, I ule adult head: Elpid sp,l (fTrichin. 5p,) 1 pupal fragsent containing adult: Elpid sp, 2b 1 incoMpleh pupa: He.erodrollliine sp, 2 pupal abdolinal segMents: Dolichopodid Ip,1 I pupa; /leg.seli. sp, 2 wings; PJ.tycl/eirIJs cJype.tus I, ht, I adult tergal fragllent; S"t,ll. IP, 1 pupal end segMent;

CIte., sp, I adult head: Acalyphrat. 'p, C1Chloropid or 0polyzid) 1 pupariuI; Ph.oni. finCln, 2 wings: 1Musc id sp, I egg: Calyptrate sp, pupariulIl containing Ichneulonid cocoon [Allo

Psocophran sp, I adult head; Antnocori. sp, adult head: PIliJ,enlJs ,pullrius 2 wing5(7 large dark fori): Apl/rodes s, lat, sp, 1 wing; Deltocephalid sp, (as in a) I wing: ?Oe/pl/acode, IP, 1 wing:

Delphacid sp, I wing (venation obscure): Livi. Juncoru. I wing: Psy/l.lp, several wings: 2 adult heads; 'rioz. sp, I wing: Lepidopterous spp, 1 pupal fragllent, several larval head-capsules;

Li,nephiJus sp, 2 wings; Coleopteroul IPP, (sue ho .. in a) nny larval head-capsules; Parasitic. spp, lany adult headl of AlYliinae, Pterolalidae etc,] HNI- 23 d 10-UCI,

Li,nophiJ, ?"igeni I pupal head shield conhining adult head intact, nUlerous pupal tergal fraglentl and 10 end-seglenti (6 nle,' feule): Tipulid sp, 6 larval head-capsules; Bibio

?nigrivlntri. I wing; Chironolid 'pp, nny larval hud-capsules, I adult thorax: Elpid sp,l

(?Trichinlsp,) I pupa: Elpid Ip,2a I pupa: EMpid sp,2b I pupa (hIS long respiratory prOCI5Ie5 frOI utlthorlcic spiracle); Dolichopodid sp, I I pupa: /legaslli. IP, I wing; TI/"i" "'lCllii 1 wing;

Copro.yzll, hto,sp, I wing (about 2111, long): HydrtIJi'lp, 1 pupariuI(.II); ?Ephydrid sp, 1 thorax

369 about size of PMJygrid but with single line of acrostichah which end at level of penultiute

postsutural (3 postsutural dorsocentrals>: Neo,yid cornicind 2 adult thoracic fragllents- incl. right side of scuteilul and left side of presutural and postsutural areas showing bristle alveoli clearly,

1 wing: Dipterous SP, indet, 1 IduU head frlgllnt [Also Anthocori. sp, 2 aduU pronota: ?Sigar,

dorSlJi, t forewing: Delphacid sp, 10 wings (venation obscure, sOle wi th dark diagonal suffusion);

Livi' juncoru, 1 wing; P,ylJ,sp, , wings; 2 adult heads (closest to P,brunneiplnni, or 'IJ,covita>:

Lepidopterous spp, 13 larval head-capsules, pupal fraglilents; Li,nlphiJlJl ffJlvicornil 2 wing fragMents: IfYrliCl sp, I right forewing, tlNI- 32 e "-l7cl

TipuJa ?oJlfiCla/pi/udell several pupal fragmenh: !Lilloni, sp, I larval head capsule: Li,nophiJ,

?lIigeni, nUlerous ,pupal fragMents: Psychodid SP, 2 pupal wing-covers: Chironolid SPP, (at lent 2 species), few hrval head-capsules, 3 thoraces; Bibio SP, 1 inco.plete larval skin: Elpid sp,l

('!Trichina SP,) &everal pupae: Elpid sp ,2a: I pupa: Helerodroliine sp, pupal tergi tes: Dolichopodid

SP, pupal abdolinal frag.ent: !SClte/Ji sp, pupariul: ?Nealyi, 'Clrnicin, I fragaent of adul t head: Phaoni, finClni 2 wings; t1uscid sp. egg (as in 6-10; 01) [Also Corixid sp, 1 fenle ovipositor and thoracic fragMent: PhiJaenus .pullriu;, nny adult heads and wings: AphrCldlJl,lat,sp.

1 head and 3 wings: ?fuICl/i, IP, I wing: Delphacid SP, 2 heads, nny thoracic and abdolinal fragMl!nts (1 including IIle genihlia as in another horizon(?), alio nUlerous wings: Ply111lp, 2 wings: TrioZi IP, 1 wing: Aphid IP, 2 fragMentl: Lepidopterous IPP, few larva} head-capsules; IIny pupal fragMents, including 2 wing-covers: parasitic HYMenopteron: thorax: IfYrli" 'p, I adult head:

Coleopteron 'P. 2 larval head capsule (Iinuate c!ypeul)]

17-20cl,

Li.nophiJI ?eiglnJ: Ievera! pupal fragMents: Chironolid 'p, 1 larval head-capsule: Bioi" 'p. 1 wing; 1Tric/lin, 'P. I pupa: Cyclorrhaphan 'P. I wing [Allo Corixid 'P, 1 wing: PhiJaenul .pu.ariut,

I adult held and 1 wing: DeUocephllid 'p, 3 wings; Delphacid 'p. nUlerou, wings: Lilli, iuncoru. I ring,' PsyJJ, ap. 2 wingl: Platygasterid 'P. I wing: Coleopteron sp. 1 larval head capsule (Iinuat. clypeul)] tlNI- 9

370 g ~-24cI

Li,nophii, ?Jeigeni nUllerous pupal fragMents: Sepsid sp, I puparial end-,eglent: Hydre/Ji, sp, I

pupariul containing relains of Alysiine parasite: Cyclorrhaphan sp,(either Acalypterate or Anthollyiid

wi th stellate anal spi rae les): 2 pupar ial end-segunh [Also DeJph.codes sp, uny wings and heads;

Psyi/' sp, I wing3 "HI- 9

Total samples 7 Total Dipterous Taxa 41 Total "NI- 215

22 LOCH OLABHAT (North Uist)

1 a Context 563 Eilean Domhnuille a Spionnaidh, C,623, ED,89 : 9:1,

Tipulid sp, 3 puparial respiratory horns (fragMents only): SCitopse nat,t" 4 pupal end seglents:

Lillosinine spp, 3 pupal fragMents (different species): SCithaph,g' sp, 2 puparia (fragments only):

1Nupedi'l I'lflg/' 'p, I pupariul (incollplete); £udasyphor' cy,ne/J, I puparial end IegMent; Neo'ri'

cornicin, I puparial end-segllent ~NI- U

b Context 1008, Underwater excavation Salple 8, C,I008, ED,89,' 1989

?Plychodid sp, 1 pupal end-segllent; Oryg" /UctUOfU' I puparhl end-seglent; lepto",,? fontin,/i,

2 puparh(u)j ThorlCochlltl zoster" 11 puparh(lI)j li.osinl Ii/rill" 17 puparia(IS II);

? Te/o.erin. fJ,rip" I puparlut(u): ?TrIC"yopelJl sp, 3 puparia(unh)j Hydrotll' tlentip" 2 puparia(tI) "HI- 38

Total lalples 2 Total dipterous Taxa 17 Total "NI- 52

(23 PAPA STOUR. Shetland)

24 PLUSCARDEN PRIORY (Moray)

Tricl/oel,. sp, 1 larval head capsule: Plychodid sp, 6 pupal fragllents: Phorid 'p, I pupariul (ea):

T,rriJi.o.In, rlCoritzli 403 puparil (5 unh), 3 wings, 18 adult heads, 19 thoraces:

HI/IO'YZI 'c,ption/Blfflt, 14 puparil(u) (Slaller with non-spiculate end-seglltnt)j ?ScoJioelnt,.

IPI 5 pupuh(lI) (larger with Iplculate end-segment): flnni, "CI/lfl. 8 puparia(u): 'Hllin,

371 rlVersio, 3 puparia(n) "NI- 3

Total Taxa 8 Total HNI=436

25 SKARA BRAE (Mainland, Orkney>

Tipul. 'p, 12 larval head-capsules of various instin, probably III sal' species; Bibionid sp, 7 feule adult head (Oilophus or 8ibio); Ifeleo,yz. boreali, 1 pupariul with Ille genitalia; ? Spelobi. sp, 6 pupariaCu); ?AlJopiophil, yulgaris 5 puparia (one with lale genitalia): Drosophilid sp, 1 puparial fragMent; C,lJiphori ur.lensi, 1 pupariul(81); ~"c'thoph.g, sp, 1 pupariuI(u); Antholyiid sp, 1 pupariuI(u); 1'1I.0Tti, 1 iTt(ln. 1 pupariul "NI- I

Total taxI 10 Total KNI- 28

26 SOUTH LOCH BOISDALE (South Uiat)

I 90-95cI (Dark brown poorly hUlified peat with twigs and wood traglents]

Tipula/l'rionoClrllp, I hrval head-capsule; I'edici. ?riYoII 3 larval head-capsule fraglllents;

£i.nopniJl sp, 2 tlrgal fragMents, 3 felale end-segMents; Cordilurl sp, I puparial end-segMent (Also

Dry.us sp, 1 adult head; O,lpn,codfl sp. 19 wings; COTto.,lu, ,TtC,PS 8 wings: Hyunopteroul spp, ('Il.gynod'f and Belytids) 5 adult heads] KNIll 7 b 110-115el ["ediul brown poorly hUlified fibrous peat]

lj.nophil, sp,Cas before) 3 pupal end-segllnts and 2 tergal fraglents; Cyelorrhaphan sp. I large puparial spiracle (Also Cono"lu, ,n"p' 2 wings; £iyi, juncoru. I wing; Lepidopterous sp. indet, fraglent of a larval head-capsul. Clarva would have been about I inch long) "NI-'

C 140-145cl ["ediul brown poorly hUlified peat with large wood fraglentl and twigs)

?li.oni. (Oicr.no.y,) ?od"t. 90 larval head-capsuln: Pediei, riYon 5 larval head-capsules; pupal prolpiracular lobe: Li.Ttophil, I. lat.lp. 3' Ill. and 6 fellh pupal end-Iegllnts; 22 pupal fragrlllnts: 1 pupal wing-f lap; Tipull/frionoce" Ip 1 larval head-capsule; ?Ceratopogonid sp. indet, 2 thoraces; Dilopllu,sp, l adult thoraces, 2 feili. adult heads; Opo.yz. 9'rlin,tioni.

(or petreiJ I wing; Acalypterate sPP. indet. 2 puparial end-Iegfllents (both different); Coenosiine sp,

372 2 puparill end-seglents [AI so Ory.us sp. 2 adult heads and S pronota; 1AphropMra .lni fragMent of

adult; Cixiu5 !nerrOSU5 I wing tip; Oelphacodes sp. indet. lOS wings; Cono.elus .ncep.8 wings;

HYlenopterous sPP. Cat least 6 species) 10 adult heads; fOrliCi ?Je.,ni I adult head; .'lfy"i" sp.

adult thoracic fragMent; Lepidopterous sp. indet. I larval head-capsule] MNI= 97

d I~S-ISOcl ["ediul brown poorly hUlified peat with large wood fragMenis and twigs]

?Tipuil/Priono"rI sp. 3 larval head-capsules; Pedici. rivos. 3 larval head-capsules; Li.nophiJ,

sp. 8 lale and 6 felale pupal end-seglents; 1 pupal wing-flap fragatnt; 7Ceratopogonid sp. 2 adult

thoracn; CordiJuflsp. 2 puparia [Allo Ory.ul sp. 2 adult heads and thoracic fragMents;

S,Idul'lp. adult pronotul: Cono.elu. ,ncep' 5 wings; ?Oelphlcadl' sp. 100 wingl (Iilple venation);

Strophingi. Ifi", 2 wings; paraSitic HYlenoptera 6 adult headl Cat least 3 diferent species);

Chalcid Ip. indet. 1 adult head; for.iCl I".ni 3 adult heads] MNla 24

e ISO-ISSei ["ediul brown poorly hUlified peat with large wood fragMentl and twigsl

Tipul,/Priono"" Ip. I larvill head-capsule and pupal fragMents: £i.oni, (Oi",no.y.J ?odl5t, I

larval head-capsule; Pedici. riroll 17 larval head capsulu (probably full range of inshrs);

Li.nophiJ.sp. 90 pupal tergitll, 4 pupal wing-covers, I pupal facial luk, 23 pUPil I end IegMents

(incl. IS lilIes); Psychodid sp. indet. 3 pupal wing-covers [Also Ory.ul sp. 3 fragMentl of adults;

Pllililnus spullrius 1 wing; ?Thllnotettix sp. I wing; Cixiu.lp. 1 adult clypeul; 1 wing frag.ent

(base darkened): OeJphlCod" .p. 180 wing!l (veins liaple, all dark, non-tuberculate): ?OelphlCodes sp. 20 wings (vlins Ilightly tuberculate and lorl furcated towards apex; Cona.elus ,n,ep' 2 wing!l of brachypters, 1 IiIcropier wing; '0""1 III~ni 2 adult heads) MNIa ~9

Total lalplel 5 Total Dipterous Taxa 13 Total MNIa 153

27 TIlQUOY a TQ 8011.

C,loblt, petroneJJ., J pupariuM: Orr;" luctUOSUI, I pupariuI; SpIIODi. 'rufiJ~bri .. 1 pupariuI:

?HydreJJi. sp., 2 pup.ria; ?Calliphorid sp .. 1 pupariuI: Cyclorrhaphan Ip. 1 pupariul "NI. 7 b TQ 803J.

373 Sc.topse notata, 1 pupa; ?lfal,eo.y;, scio.yzin" 2 adult heads; Copro.yz, Iquin./fi.ilis, 2

wings,2 puparia; Op,Ji.osin, JiJiput'TII, I pupariuI; ~p,/obi' ?rufiJ,bri., 8 puparia; TrlehyopIU,

eoprina, 3 puparia

c TQ 803- 88 Pitt I

Bibionid SP, 2 larval head-capsules; 'SCItopsci'rI vivid" 1 adult head and I wing: Se,topSl

no t.t., 1 larYi and 6 pupae: Copro.yz. ,quin./si.iUs, 2 wingl,1 head, 64 puparia; "hor"ochlll,

zoster." 13 puparia; TrlehyoplJi. coprina, 201 puparia; Op,Ji.osin. JiJiput.n., 9 puparia; Spl/obi,

?rufi/.bri., 332 puparia; "Li.osin, IP, vi' 2 puparia: PulJi.osin, hlterofllUrI, U puparia; Lilosinine

SP, indet, 4 adult headi: Cyclorrhaphan 'P, 1 pupariul "NI- 649 d TQ 804

H""topot,/Chrysops SP, 1 pupal fragMent; Copro.yz, equin,/si.iUs 14 puparia; ThorlCoeh"t,

zosterll, 6 pupariaj Trlehyopell' coprin" 5 pupariaj Spllobi. ?rufi/.ori" 15 pupari.

"HI- 41

I TQ 806 (HBIJ,Sadler Iuggests horizons 806 and 803 synchronous)

Tipul' BP, I larvil head-capsule: P,diei, (Tricypllon,) i",eul,t" 2 adult heads: ?Bibionid sp. adult head: ?SCltoplCi'rI 'livid" 1 wing: SCltops, not.t., 15 puple: ~'y.pycnus dlSoutterri, 1 wing

Lonchopt,,, lute., I wing of feuh; C,lob.t. !pltronlll" 1 pupariuI; Coelop' !frigid., I thorax;

H,teroch,iJ, buc"t" 2 puparia; SphllroCl" curviplI, 10 puparia, 3 adult heads: COpfO.YZ'

,quin./,i.iii" 137 puparla, 27 wings, 10 heads, 2 thoraces; Op,cUrons hll.id" I wing; fThorlCochllt, zoster", 7 puparia; KiniJi.o.in, ?fllngieol., 3 puparia; Op,li.osin, JiJiput,n" 38 puparia, 3 wings: Spllobi. 'rufi/,bri,,· 304 puparia: PuJli,osin, 1I1teronlllfi, 45 puparia:

TrlChyoplJi, coprin., 8351 pupui.: Sphurocerid SPP, I pupariuI, 6 heads, 3 thoraces: ?SCltllJ,Sp. 2 puparia: Li,nlUi, qu,d"t" I wing; Scapto.yz, gra,inu, (or Drosophil'lp,) I pupariuI: Acalypterate

SP, I pupariuI; Cyclorrhaphan SPP, 3 puparia; SCltllophlg' ?fllrc,tl, 3 wings: Antholyiid SP, 2 puparia; Coenosi"ollieu/I, 1 wing: Helin. ,vlCtl, 1 pupariuI, 1 adult Icutellull (this 'pecies?)

"HI- 8993

374 '---- -

TQ 808

Tipu/, o/er'L"Ii/p,/udo", 1 pupa: Chi ronolids, 5 adul t thoraces: Psychodid IP, 1 pupal f rlgllent:

SCitopse not,t, 3 pupae: Neutoceran sp, 1 hrval head-capsule: Copro.yz, ,quin,/siliJis 58

puparia(u), 10 wings, 14 heads, 5 thoraces, (NB, 1 of thoraces positively identifiable II ,quin"

the lesopleufon being in perfect condition,): i.ill~sina li/vatiCi 2 wings, , puparia: Spelobi'

c/unipes, 3 wings: ~~ !rufi/lbri. 193 puparia: TriellyopeiJ. coprin. 588 puparia (573 II):

0plJi,o,i", JiJiputl". 1 wing (NB, Tthere I.y have been 10lle badly fragMented puparia uongst the

total given for Sp,/obi, above): Pulli,osin, Iteteroneurl 5 puparia: Lilosinine spp, indet, 3 heads, 1

thorax: 1S"toplli/, sp, 1 pupariul: Cyclofrhaphan spp, 2 puparia (different speCies) [Also heads of 3

species parasitiC HYlenoptera] "NI- 901 g TQ 810

?/.i.oni, 'P, 2 larval head-capsules: f,diei, fTricypllon,) !ill'L"u/at, 4 !lrnl head-capsules:

Li.nophi/, sp, (pupal fraglllntl of a nle and feull); [riopter, ?Jut" v, t.enionot, 2 hrval head­ capsules, 1 pupal fraglltnt; Ceratopogonid liP, thorax: Bibio nigriventri. 2 wings:

£yeoriell,18,,(/y.i. sp, 1 wing: SCitopSl IIot,t, 1 pupa; ?Syrphid BP, 1 fragraent of pupa; £,ptoelf'

fontinaJis 3 puparia, 1 adult head, 2 thoraces, and 1 wing; Copr",yz, sp, 2 puparia, 5 adult helds and 2 .ings: (,.i,iJi. 2 thoraces; !£i''''in, Ii/y,tiel 5 puparia: Tllorl(o,I1"I, zOlterll pupariul: TrlCl1yop,JJ, coprin, IS puparia; ~p,/obi' !rufilabri. 9 puparia; Pulli.osin, l1,tlfon,ufI,

, puparia: Lilosinines sPP. indet, 5 adult heads: Sphaerocerid SP, indet, 6 puparia: SCitophi/' SP, 13 puparia: fSClpto.yz, grllinu. 2 puparia; ?Tl1ricop. IP, 1 pupariuI: Helina 'P, 2 puparia:

Cyclorrhaphan spp. 6 puparia [Also head of Trichopterous adult] "HI- 105 h TQ 813

Tipulid SP, 1 larval head-capsule: Copro.yz, Iquina/li.iJiI I pupariuMl Sp,iobi' 'rufi/.bril, 5 puparia: • £i,o,ill' SP,' 2 adult heads, 2 thoraell: Agrolyzid SP, 8 puparia: COlnosia SP, 1 pupariul

[Also heads of paralitic HYlenoptlra- 8 Pterol.lids, 1 Jchneulonid, 2 Ceraphronidl, and indet, thorax] MNI- 18

375 TQ 8U

li.oni, IP, 5 larval head-caplules; fldul, aricypflon,) 1".CIII,t" 2 larval hlld-capluln, 3

wings, 3 Idul t hndl; [rfoptu. r, t.MiMOt., 5 larval hlld caplules: Chi ronolldl, 3 thoncn; DIDIO

nigrirentri', , thorax, 2 wings; Oilophu. (I"rillll'"o"tu., 5 adult hlldl, I thorlx: Copro.yr,

,quln.I.I,iJis, I wing, I hud; LlptocI" fontin.Ji" I thorax; £i.o.ln•• ilr.tu., I wing;

TflClIyopIIJ. (oprin., 17 puparil{fll; ~'pllobil !rufil,"ri" 2 puparil(u); fuJJi,o.lnl IIItlronlufl, 2

puparia; ·Li.osin,sp,· 3 thoraces; Sphlfrocerid IP, I pupuiuI; $(,thoph'9' 'P, I puparlul; 'uc,IJi,

!llIcorll', 2 Idult 1111 hlldl

TQ 815

Chlronolida, 8 thoracni S"top" not.t., I pupa; topro.y:, Iquln./.i.iJi., 2 wings; TflCllyoplJJ,

caprin., I pupariuI; • Li,osin, IP,' I adul t thonx; Ephydrld IP, I pupariuI; Sc,pto.yr, 9rlllnu" I

pupariuI; SCithoph'9' !fllrClt., 1 wing; Cyclorrhaphln IP, I pupariUl

ftNI- J7] k TQ 816

Tipulid SP, 2 larval hud capsules; [rioptl" Jute. " t.,nionot., I pUpl, 3 wingl, thorax:

SCitOPSI not.,., 2 larval Ikinl; SpIJobi, fru(iJ.fJri., 13 pup.,ia "NI- 22l

TQ 817

Tipulid IP, 16 larval head-capsulfli f,diei. fTricypllon.J !i""uJ.t. 3 Ilrval hlldl, J adult head; Chirono.ids, • thoncfl; Centopogonid IP, I thoraxi !Se,topl(i,,, virid" I wing: £ycorilJJ,/ lridy.i. SP, I wing: 6ibio nigrir,ntri., 1111 wing; S(ltop" not.t" I PUPI; (opro.yz.

,quin.I,i,iJis, II puparia, 1 wino, • heads, thorax: Llptoel" (ontin.lil, 1 ling: TflCllyopIJJ. caprin., 132 puparh: fulli.osin. ",t"on,u", 5 puparia(fI)j Spllobi, frulil.bri., 63 pupuil(tl);

Se.pto.yr. grllinu., 2 puparil, I wing: HydflJJi. f.odllt" 2 thoracllj S(lthoph'9' 'P, , pupariuI:

(olno.i.lp, 3 pupariaj HIJin. ,r"t" I pupariuI; Hllin'lp, 1 pupariuli Cyclorrhaphan IP, 2 puparia

[Also 3 adult heads of parasitiC Hy.enoptera, incl , I Ptero.alidl "NI- 260

Total II.plel 12 Totll Dipteroul Taxa 55+ Total "HI- II, JOJ

376 ICELAND 26 BESSASTABIR

Bessastabir 10 (pre 1226)

Se,topsi notit, S larval skins, S pupal fraglents; Heleo.yz. lerr.t. 190 puplria(el);

?AJJopiophila vulgaris I pupariuI; Copro.yz. sp, I wing; Lepto"" ?fontinali. 6 puparh(ea);

?TlIoracochieta losteril 2 puparil(e.); £i.osin. Sib'iUCI 2 puparia; T,lo.erin, fJavipu 226 puparia; Se, thophaga sp, 3 puparh(el); (Keloph.gus evinus ~ I1NI- 439

Bessastabir II (pre 1485)

TriehoCl" sp, 3 wings, I pupal fragltnt, 4 larval head-capsules; Psychodid sp, I pupil fngllnt;

Sc.tepse not.t, 3 wings, 56 pupae, 7 larval skins, 1 larval head-capsule: Scilrid sp. (llrge species having estillted wing length of C,511) 1 wing fragment (basal third only); Oolichopul plu.ipe. 1 pupal fachl luk: KegISlJi. cf, rufipes I pupariuI(u); Heleo.yz, serrat, 2 adult heads, 14 wings,

94 puparia: Copro.yz.sp, 4 wing, S adult heads, 2 pup.ria: Lepto"" !fontinaJis 3 wings, 2 puparhCu): Thor,cechilt, zoster" 1 pupariuI: ?!i.osin, .i/v.ti" 3 puparh: ?Herniosini bequilrti 4 wings: Teloltrin, "nipe. 229 pupariaj ?AJJopiophiI. vulgui. I pupariuI:

Acalyptente sp, indet, 2 puparhCu): Sc,thopha!J' c,/ida I adult head: $, CI/ida/litorea 4 wings:

$, furCita/.tercoflri, 13 puparhCea): ?Nupedi. sp, 1 wing: (Kelopha9us evinus 9) [Also

3 heads of adult parasitiC HYlenoptera including I Ichneulonidl I1NI- 429

Bessasta6i, U

(Keloph'!Jus ovinus I)

Bessash6i r 15

SCitOPSl not,t. I pupa: (KeloplJ'fJuS evinus I)

Bessastabi r 17

HllIO.YI' lerflt, 2 wings, I pupariulj Copre.yza sp, I puparill fraglent: r,lo.,rin, fJ,vipes 3 puparh: SCithophag, ClJid,/litorli I wing [Aho I Trichopteron wing-frag.enU

377 Bessistabir 18 SYI,

Telo,erina fJaripes 1 pupariull I1NI- 1

Bessastabir 19 SYI,

Triel/ocera sp, 1 wing-fragllent; Scatops, notaia 1 pupa (in larval skin); Heleo.yz ••errat. I adult abdolinal fragMent, 2 wings, 1 pupariuaC .. ); Herniosin. bequaerti 2 puparia; Tel0.erin.

fl''1ipes 10 puparia; SCltl/opl/.g. sp, 1 wing, 1 pupariul [Also 1 head of an adult AIYliine Braconid wasp and 1 Linyphid spider cephalothoraxl I1NI- 16

Total la.ples 7 Total Dipterous Taxa 18 Total I1NI- 891

(29 EYNHYRNINGUR)

30 FINNBOGASTADIR

A '4901 Sv I' [Specilens 111 fraglentary Ind strongly bleachedl

/feg.seJiI IP, .indet, I pupariulI(eI\); Heleo,yz. boreali, 348 pupariaCu)j SClt,JJ. sp, 1 wing:

SCithoph.g. sp, 2 puparia(el); I wing: ?Pegopllt. intir" I pupariuICu) I1NI- 354

B '4901-39 21/7/90 Road cut profile Col, SU 9

Hel,o,yz. bore. Ii. 21 puparil(u); H. "frata I pupariuICel)j fAJJopiophiJ. 1 puparial fragMentCe.): C.lliphor.lp, I larvil Ikin [There wer, two lodern contaMinantl in this lallple, nalely a perfect lale Sciarid and a felale Psychodidl I1NI- 24 C '4901-30 Road cut Profile C12 la'ple'

Heleo,yz. borelli. 7 puparilCel) (length 3,0- 5,4.1): fCoprolyzl sp, I puparial end-Ieglent possibly belonging to this genusj AJJopiophiJ,sp, I puplriul(u)j fDot.ntJphil. fuga,,/( sp, pupariuI(u) I1NI· 10

D '4901-3921/7/901 Road cut profil. Col,Salple SUII"

HeleolYz, borel/f.lp, 7 puparia(e.); AJJopiophil. sp, 1 puparial fragMent(u)

I1NI· 8

378 E "4901-39SUll"

HeJeo,yz. bore.lis 51 badly fragmented puparii!Cel) (lastly SlaU but three decidedly larger, possibly belonging to H. ,erfit,),' AJJopiophiJ. sp, 4 puparia(u); Sc.thoph.g. 'p, 1 pupariuI(u)

MNI= 56 F "'901-39 Road cut(12 SAK) 7,12,90"

HeJeo.yz. bore. lis 163 puparil (141 ell) (length 3,42- 5.4111); AJJopiopflil.lp, 10 puparh(n)

(length 3,42- 3,90 •• ); Sc.thopha9'sP, 9 puparia(7 ell, 2 unh.with reflains of fela1e terllinalh) "NI· 182 6 R,C,Prof,col Sal SU 14 21/7/90"

Heleo,yz. bore, Ii, 307 puparia(279 n) (reurkably constant in lize and including no large specilens; presuJlably III borealis),' ?Copro,yza sp. I pupariuI(unh); AJJopiophiJ. sp. 45 puparh(41 efl, 4 unh); Sc.thophag. sp. 6 puparia(e.) KNI- 359

H "49011-39 21/7/90 Raid cut prof.coluln Salple SU 14" [3 tubes- a,b,c]

HlleolYz. bore,/is puparia IS follon- .) 338(302 nl, b) 393(321 tI), c) 324(263 n) • 1055(886

II) (another elpty pupariul 6.0 .. long us perhaps H. serrat,); AJJopiophi/. sp. puparh 15 follows­

.) 23(21 el), b) 22(21 el. 1 unhitched one, on the Itructure of the felale genital seg.ents looks lore lik •• SCiptolYZ, sp.), c) 21

8(eM), b) 2<1 n), c) 11(11). 21<20 II): "egop/,t. infir" 1 pupariuI(III) MNI. 1144

Total Illplel 8 Total Dipterous Taxa 10 Total "NI- 2136

31 GJOGuR

A '4901-7 6jogur SU 38 J,S, 1

HeJeo.yu borliJis 1 pupuiu.

8 '4091-7 6jogur SU 40 J,S,'

Hlleo,yz.Dorl,li, 15 puparil(14 .1): H. '"rr,t. 2 puparial end-seglents probably this species, being luch llrger than previous 15 specilens; ?C.//lphor, sp, fragMents of 1 pupariul

"NI. 18

379 C "901-7 6jogur SU '7/48 1,5,' [bleached and in poor Itat. of preservation

Heieo.yz. borellis 12 puparh(u); Allopiophil,sp, 66 pupariaCu); 1Ephydrid sp, indet, \ puparial end-seglent; SCithophaga sp, 8 puparia(e~) HNI- 87 o "90\-7 6jogur SU 60- 61 S,S,' [bleached, frag.ented and in poor preservation]

Heieo,yzi borllJi, 6 puparh(n); AlJopiophiJ. sp, 1 puparhl fragMent; SCithoph'ga .p, 1 puparial end seglent HNI- 8

E "901-7 6iogur 5U 67 J,5,'

AlJopiophiialp, I pupariuICea); ?Calliphora sp, 1 puparial anterodornl cap "NI" 2

Total 5a.ples 5 Total Dipterous Taxa 6 Total "NI- 116

(32 HOLT>

(33 6SABAKKI>

(34 PAPEY>

35 REYKHOLT

[Puuge]

Sy.,4 9207 60-80 7123 30-70 28,2,88

lfeglSeli, cf, rufipe. I pupariuI(ea); (lfeioph.gulovinul 19) I1NI- 20

Sy., 14 28,2,88 x 207,60-80, Y123,5-70

Prionocera turcica I felale pupal end-segMent, 8 larval head-capsules of uniforl lize(this specin?); Rhl.pho.yia !si.plex 2 pupae; lfegaseJi. cf, nig" 3 puparil(el>; ", cf, rulipe. pupariul(u); Hello.yz. ?boreali' 11 puparia(IO .1); .fAJJopiophila vuiglri' • puparia(n); rPhilygri. vittiplnni. 1 pupariul; OrosophiialSClpto.yz,sp, 1 pupariul; SCithop/tag'

(urCita/stlrcoriria 1 wing, • puparia(tII)j Bot.nophiia ffugl,l( • puparia(el); Inthollyiid sp, pupariuI(u); .'Spilogon,sp, 2 pupariaCe.) [Also Saldid sp, 1 helielytron fragMent; Trichopteron sp, 3 larval Clsel containing larval IcIerites, 1 adult wing frlgMent] I1NI- 38

Sy., 15

380 Tipul. confus.lrufina I larval head capsule (l. 51l1li long); Hello,yz. !boruJi. 1 pupariuI(ea);

?Spilogon. sp, 1 puparial end-seg.ent; (/fllophagus ovinu. J.I MNI- 3

SY·,32

"ipula sp, 1 pupal fragMent; /fegaseJi. cf, rufiplI 1 pupal end-segMent; OoJichopu •.?plu,ipes pupal fachl usk; Heleo.yz. 'bofl.li. 4 puparia; ?AIJopiophila vulgaris 22 pupariaCea);

(HelophaguJ ovinus 1) "HI- 31

SY·,33

/felophaguJ ovinus 5 adul t heads, 8 thoraces HNI- 8

[Rool 2]

Sy.,19

/fegaseJi. cf, rufip" 1 pupariul(ea); Sphaerocerid 'P, I pupariuICII); fAJJopiophil. vulgaris pupariul(el); fSpilogon. sp, 1 pupariul(n); (/felophagul Dvinus 11)) "HI- ( Sy.,20

SCllop" notlt, 1 pupa with larval skin(e.); /fegaseli. cf, ruNpe. 33 puparia(tI); Heleo.Yl' bore.li. 7 puparia<1 1I1e unh, ) MNI- ~l Sy.,21

/feloph.guJ ovinuJ 43 [Aho Pediculu.sp, uny fraglents]

Sy., 22

Tric/loClfl 'p, 15 pupal fachl usks; Scatop" notlt, 1 pupa, 3 larval head-clplulel, 5 pupal

Intennal Iheathl; "biD po,on., 2 larval skins C1 allost co.pllte); Oo/ichopu, ?plu,ipII I pupal facial IIsk; /feglleJi. c1, rufipII 26 pUpal'il(II); Copro,yz.lp, 5 wings; lilolinint IP, pupariul(u); !Allopiophi I. vulglfi. I pupariuICu); /feloph.guJ DvinuJ 50 CAho Trichopteron BP,

2 Idult heads, The Balpl. 1150 contained I las I of Coleopterous fragraents lixed with hairs and feathers] MNI- 102

Sy.,30

/feglSeJi. ct, nigr. I pupariuI; Heleo.Yl' !borliJi. 19 pupar!a(era); Sphaerocerid 'P, 1 pupll'iuI;

Sc.lhoph.g.sp, 2 puplril; (/feloph'glJf ovinul 2) "HI- 23

381 [Rool 1] SYI,36

Megasel it cf, rufipe. 2 pupar ial end-segments: /felopl/agus ovinus I adult thorax [Aho

Lepidopteron sp, 1 larval head-capsule] MNI= 3 [RooM 1] SYI, ? x 208, 60-209, Y 123,18-57

/Jelopl/aguJ ovinus • adult heads, 9 thoraces MNI- 9 (R98l)

f/Jelopl/agulovinus 210) MNI- 210

Total lalples 11 Total Dipterous Taxa c 21 Total MHI- 554

36 ST6RABORG

1981 Hus I. Pit [Icelandic Hus- rool]

OiJophus 'P, 2 feull adult heads: T /Jayetiol, sp, 8 pupuit: Heleo.Y11 .errit, I pupariul:

?Copro.YIl sp, 2 puparh: Scatl/opl/aga sp, 4 puparia: Fanni, sp, 1 pupariulI(ea); ?:;'piJogon.sp,

pupariul(u) [Also 1 Lepidopterous larval head capsule and lany Coleopterous frilgments- ',(I/inus,

?PhiJonthu., Hypnoides, OtiorrhynchuJ etc,] MNI- 19

2 198. Under passage HOS T 30

H,leo.yz. serrlt, 9 puparia

3 1984 Hus 31/7

Schrid IP, indet, 1 .ing: HyrfrlJ/i. 5P, 1 pupariul(III): Orosopllil. 'P, 1 pupariuI(u):

SCithophlgl 'p, 1 pupariuI(ea); ?SpiJogonl sp, I pupariul(n) MHI- 5 • (12) 13-17 HUI

Heleo'Y11 .errltl I pupariuI: Hydrillia IP, I pupariul [AIlo fraglent of large wing, 1

MNI- 2

5 20/8 Layer below west pari of Hut 33

Antholyiid sp, indet, I pupariuI(el) [Also Ichneulonid adult head,] "NI- 1

6 85 Hus 3' Floor

H,leo,yzi serrlt, 2 pupari.,' /J,lophagIJI Dvinul I pupariul MHI- 3

382 7 1981 Hus 14 under floor

.'lfayetiolalip, 120 puparia: Oilophus 'p, S adult heads and 2 thoraces: Hello"Yli "rflt. 3 puparia:

Sphaerocerid spp, 7 puparia (at least 3 species, 1 very nall: hrger ones possibly Copro.yz. Ip,)j

HydrelJi.lp, 3 puparil: SCithoph.ga IP, 10 puparia: F.nnia sp, 2 puparia: 1 Tflri(ops sp, I pupariuI:

1SpiJogon. sp, I pupariura [Also I Ichneuraon cocoon, 1 ?sawfly cocoon, I very Slall Lepidopteroul

larval head-capsule and 5 different parisi tic HYlenopterous adult heads, I

"HI- 152

8 HUI 18 Floor

Ifayetiola sp, 4 puparia: $(ltOPSI notat. 1 pupaCera): HelloMYz, serr.t. 3 wings: 31 puparill

?Copro,yz.lip, I pupariuraCu): Herniosina DlqlJaerii 8 puparia(e.) "HI-.S

9 1981 F Trench S,Profile ColUMn '0-45cI, 3,825 Kg, 12,9,81

Heleo.yz. serflt. 96 pupariaj Copro.yz.sp, 2 puparia; ?C.lliphora sp, • puparia; ?Hydrot.e.

dentipes I pupariulI(u) IINI- 103

10 Pit under HUI I.

SCitopSl notat. 6 puple with larval skinsCu): Hlleo'Yll serrati 250 puparil: Telolerin. fl.viplI

9 puparia(ell): Herniosin. Dequaerti 1 pupariul IINI- 266

11 1981 F, Trench W,Profile Colulln 35-40CI,

HIIIO'Yll .,rrlt. 3 puparia(fII); ?Tflllifl dllpfi I pupariul: Lilosinine liP, indet, 1 pupariul:

Fanni. lip, I pupariul "NI- 6

12 F Trench S,Pofile 10-ISCI,

Hello,yz. "rflt. 2 puparia: ?Thllirl d•• pfi 2 puparial end seglents: Acalypierate sp. 3 pupariaj

Sc.thoph'g' sp, 2 puparia: Cyclorrhaphan spp, 3 pupari.l fragMenis "HI- 12

13 1981 F Trench/W,Profile Coluln 40-45cI. 3,49kg,

Hlleo.yz. ,err.t. 1 pupariuI: 1Sc.thopfl.g, 'p, 1 pupariul IINI- 2

14 1981 F Trench

Chironolid Ip,indet, I adult thorax: Hello.yz, "rr,t, 26 puparia: 1 adult thoraxj S('th~h,g,

IP, 2 pupari.j Intholyiid sp, 2 puparia "NI- 31

383 15 6 Trench NProfile 65-70'1, SCitoPSI notat, 2 pupal end-segunts; Heilo'YI' "rr,t, 9 puparia; S"thophag, sp, 5 puparia: Tlio.erin, fJ,vipes 6 puparia; ?Agrolyzid sp, no, 7 I pupariuI; Calliphorid/Scathophagid sp, no, 6

pupariuI; Cyclorrhaphan sp,no,S I pupariul tlNI- 2S 16 Drain under H14: Upper Layer

Trichocl" ,.cuJipennl. 9 wings, 6 adult thoraces and 2 heads, 2 pupal wing-sheaths; OiJophul sp,

I adult thorax; Sc,tOPSI not,t, 34 pupae(el), lostly in larval skins, 9 adult thoraces, 9 larval

head-capsules; Hello.YI' serr,t, 201 puparia, 17 wings, 7 adult heads; Copro,yz, sp, 6 wings;

Herniosin, bequllrti "puparia; Teloflerina fl,vip" 48 puparia; '~'peloDj, 'p, I wing; ?Sc'pto.yu

'p, I puparial end-seglent; /feloph'gus ovinul 2 adult heads and 4 thoraces [Also 4 Otlorhynchu.

!nodosus and I Lepidopterous larval head-capsule] tlNI- 313 17 F Trench S Profile coluln 25-30CI, 12,8,81

Heleo.yz, ?borelli. I pupariul(tI) (very nail one); ?Copro,yz'IP, 9 pupariaCtI) tlNI- 10

18 H~5 18 Floor E-W

Se,topsi not,t, 3 pupae with larval skins; Hello,yz, 'p, 84 puparia el, Further 9 unh,compriled 2

IIrr,t, and 2 borlllis: Telolfrin, fl,vipeg 3 puparia; Lilosinine sp, I pupariuMCelt) tlNI- 91

19 H~5 14 Pit

Heleo'Y:1 sp, I pupariul tlNI- I 20 F Trench Profile Coluln 35-40cI W7,

Heleo.yzi sp, 6 puparil(el); Copro,yz, sp, 3 puparia(2 II); L,ptoCl" fontinlJi. 12 pupariaC7 II)

tlNI- 21

21 1981 F Trench S Profile 25-30CI,

Se,topsi not,t, I larva; Hello.,z'lp, I pupariuI(el)j !CoprO'rZIIP, 33 puparia; Lilosinine

5PP, ( ?lfiniJi,osin, and nrIChyopeJJ,) 1 puplri'(11.1/ Ailopiophil, !vullgri. I pupariuI(II);

Sc,thoph,g. IP, I pupariuI(el) tlNI- 39

384 22 STORABOR6 C

Tri,ho"" ••culiplnnis 2 wings and possibly pupal fragMents; HIllo.y;. sp, 8 puparia(ell)i 2 wings

(prob, serr,t,)i Lilllosinine spp, (flfiniJi.osinland T"chyoplJJ,) 2 puparia "HI- 11

23 STORABOR6 E

SCitopSl notltl 1 pupa with larval skin; Hlilo.yz,sP, 9 pupariiO unh, tellle); ?lfiniJi.osin, sp,

10 pupariaCea)j AJJopiophii. ?'1I1/garis 1 pupariuICea); 1SpiJogon • • p. 1 pupariuI(eM); /fliophlgllS o'linus I adul t head "HI- 23

24 STORABOR8 F

SCitopsI notltl 3 pupae, I with larval skin: Hillo.y!. 'p, I wing; 89 puparil(83 lI)(of 6 unh. lales, one 3, hi, long): Herniosin, biquilfti 22 puparia<18 II); :J(,thoph'9.IP. 3 puparil(2 with perispiracular plpillae Ihorter, lore uniforl in lize; other hal the. larger Ind outerlost large5t)

"HI- 117

25 STORABOR8 K

Hlllo.yz. sp, 13 pupariaee.): Copro.yz. Ip. 1 wing: Sphaeroc.rid 'p, 2 puparia(.I) "HI- 16

26 F TRENCH(W) 0-5 STO

HIJIO'Yli sp, U puparia(63 n; I unh, tellle); SCithoph.g. 'p, I puparhl end-legMent: ?SpiJogonl

'p, I puparial end-seglent (anll spiracles lining but could be I SpiJogon, sp,) "HI- 66

27 Chlrcoal J Pit uner H~. 14

Hlilo.yz, sp, 2 pupariaee.); ?Leptoelr.lp. I puplriul(el); ?~piJogon. Ip. 1 puparial end-seg.ent

"HI- 4 28 STO I

?IfIYltioJ.sp, 8 puparia: Sc,topJI not.t. 5 pupae; H,l,o.yz, sp. 23 puparh(21 ert):?Copro,yz.lp.

2 puparia(efll): ?tlptoClr. fontin.Ji. 3 puparia(el)i AJJopiophiJ.lp, 4 puparia(el)i ?? /flonllrl Ip.

I pupariuI(el): Antholyiid/Scathophagid IP, I pupariutCell) "NI- 47

29 STO 2

Triel'lo(l" "c"Jipinni. I pupil wing-cover; /faYltioi, sp, 8 puparia; HeilOlrl11p. 3 pupal'ia(era);

385 Herniosin, bequllrti 5 puparia(u); Antholyiid .p, indet, 2 puparia(u); /(elopll.gu.ovinu. 1 adult

head; tel/Uphor, sp, indet, 1 puparial frag.ent I1NIa 21 30 F 1RENCH(U) STO 95-100

HIIIO.Yll sp, 11 puparia(u); 3 adult heads and 2 thoraces; Te/ollrin, fllVipt. 1 pupariuI( til):

?Hyd"lIi, gri"ol, I pup"iu.(II) ifNI- 13 31 1982 Hus 19

Trichoeer'"t"uJipennis 1 wing and 1 adult head; Seltopl' not,t, 1 pupa(el); Hello,yz'lerflt,

1I puparia(37 II; 1 of unhatched ones a 1I1e); Copro.yz, sp, 'puparh(III); Hlrniosin, Dlqu,erti 2S o puparia(u); T,lollrin, fJ''1iplI 55 puparhCea); St",thoph'g'sp, 2 puparia(u); Ifeloph'gu, ovi nUl

1 adult head £Also hind leg of 6yrinulsp,) I1NI- 130 32 Drain under Hus 17

Triel/oeefl .8CuJipenni. 'pupal fragllenb; 13 adult heads, 20 thoraces and" wings; OiJophu. sp,

1 adult head(lale); Sc,tople not,t, 'pupae (with larval skins), 7 larval head-capsule., 3 adult thoraces; He/eo,yzi Ilrr,t, 50 puparia; 28 adult heads, 19 thoraces, 38 wings and 3 basal abdolinal

tergite. (2 of these darker in colour when in spirit but yellowish when dry, 50 probably all this

species,); Copro.Y11 s,ltr,sp, 1 pupariuICe.), 2 wings (not frol sale speCiMen) and 2 thoraces

Cru,o.yi, nUid, 1 thorax, showing conspicuoully .trhted lesopleurae: H,rniosin, bequllrti 5

puparia(e.), , wings; Lilosinine sp, 31 puparia(e.), 3 adult heads $e,thoph,g, 'p, 2 puparia(e.);

lfe/op/7,gul ovinul c. 10 adult hnds and thoraces CAho c,20 Pediculus] I1NI- 126

33 Drain under Hus 17

1f,}oplllgUI ovinul I pupariuI; 3 adul h 34 F Trench 20-25cI S1 13-81

!/f,yetio/, sp, I pupariu.; !Copro'11' sp, I pupariuI(e.); !HydflJJi, grilee}, I pupariuICII);

If,lophigul Dvinul 1 adult thorax HNI- ,

35 H~s 18 Pit

Sc,tep" not,t, 1 pupa(el>; Hll, borl,li. 8 puparia(1 unh,); /(1/, ovinus 2 Idult heads, "NI- II

386 36 Hus 18 Floor

H,/,OlYl. sp, 78 puparia(el)(lize range 3,9- 6,611; naller ones probably including H,borl,lis);

Kerni"sin. beqllaerti 2 puparia( I perfect unh, nle); fe/ollrin. fI.vipls 7 puparia(ell HNI- 87

37 1982 Hus 7 Drain

rrkhoel,. 'dculipenni' I pupal wing-cover; SCitopse not.t. I pupal trag.ent; Heleo.Yl.lp, 16

puparial end-segments(4 of these larger than other. 50 presumably H,slrr.t~ rel.inder probably

including H,Dore.IiJ,); ?eOprO.YId IP, I puparial end-uglent; Te/ollrin, fl''1ip" 16 puparia(ul;

rrracllyopeJJ,sp, 1 pupariuI(ul; Sc.thopll.g.lp, 2 puparial end segllents HNI- 38

38 STORABOR6 (1982) 3

CoprO.YZII, sb, sp, 1 wing; ?Htrniosin. bequaerti I puparill end-segunt "NI- 2

Total salples 38 Total Dipterous Taxa 35 Total "NI- 1875

GREENLAND

36 QEQERTASUSSUK

Qt 87 U

NlollI'i. pro.inln. 7 puparia(ea); Hlleo.Y1. boreali. 2S puparia(eal (all large, C, 5, 7111);

AJJopiophil' '111/'gll'i. 11 puparia(em); LlJiopiopllil, pi/Oil S puparia(ea); Pllol'li. terrll-nO'l1l 7 puparia(ul; Hydrot"•• nxi. 2 puparia(u); PII.oni.lp, 2 puparia(ell) "NI- 59

2 Qt 87 B6

Hello'Y1. borlili. 51 puparh(ea); AJJopiopllil' '1l1lgari. (9 puparia(u); LlJiopiophil. pilon 38 puparh(u); PIIol'li' tlffil-novae40 puparia(ul; Hydrot.e. ,nxi, 6 puparia(u) PlNI- 18(

3 Qt 87 89

Neoleri, prolinen. U pupariaCl3 nl; HIIIOlY1. borllU. (0 puparia(II); IU/opiophil. '1!JlglI'i.

(9 puparia(J unh); lasiDpiopllil. pUDI. 70 puparia(1 unh); S(ltllophlg•• p, 1 pupariul(u); Pllor.i. tlrrll-nO'lae 123 puparia(2 unh,); !fllClJJi, sp, 38 puparia(el); !fIlCllJi. 'p, 32 puparia(31 tI);

Cyclorrhaphan 'p, I pupariuI('.) PlNI- 368

387 4 Qt 87 B 19/19 2 6reenland

Sciarid sp. I adult head; Neoleri. pra.inenJ 63 puparh(e.), IIIIIOIIYI bo,.,,/i. 323 puparhCu:

length 3.6-5.811), c.12 head and thoracic fngllent5; AJJopiopllil, vulgariJ 229 puparia(n. 3.12-

5,4 •• ): LuiopiophiJ. pilon 108 puparia(eI. length 4.5-5.1111), ?Fu(lJiia/ ?Paregle) 126 puparh(2

unh); Hydrotaea Inxi. I pupariuI(e.)j SpiJogon.lip. 1 pupariul (an. lip. 0.05.1): Phorllil ter,.,,­

nov., 21 pupariaCe.), c. 2 tergal fraglents [Also parasitiC HYlenoptera 2 fr.glents; Ply/I'lip. 3

wings tlNI" 876

5 Qt 87 824

Neol"i. pro.intnJ 3 puparh(n); Heleo'Yl' borealis 6 puparia(2 unh. 3.6-5.6111); Ailopiophi I.

vulgaris 6 puparia(1 unh): £lJiopiophil, pilo51 I pupariuI(ea): Scathophagllp. I pupariul\(unh),

?Pareg/e sp. 8 puparia(unh): Phofli, terrae-novae 24 puparia(4 unh) "NI- 49

6 Qt 87 CIO

Nlo/"i, prolinlnl 3 puparia(u): HI/IOIIYZ' borldJi, I pupariuICe.): AJJopiophiJ, vu/gari. 19 puparia(n): ?Fucellia sp, 2 puparh(eI): Hydrotald ,nxil 1 pupariuIICeI); Pho,.,i, terrae-nov" 26 puparia(e.) [Also Lepidopteran 'p, I pupal end-segMent] tlNI- 62

7 Qt 87 CU

Neo/"i, prollinenl 170 puparh(u)j 2 Heleolyzid thoracic fragunh: HlleollYl. boreaJi. 22 puparh(u; 3,9-4.5 .. ); AJJopiophiJ, vulg.ri, 126 puparh(ea): SCitliJ, "tagn,li, l pupariuI(u)j

?Pareg/I sp. 21 puparia(e.)j ?FuClJii.sp. 7 puparh(II); Hydrotlll.nxi. 1 pupariul(III); Spilegon,

.p. 2 puparia(!.): Phorlli. tlrrie-novil 34 puparia('I) tlNI- 386

8 Qt87CI9

lIeleo'Yll boreaJiJ I pupariuICunh, 3,6111); Ailopiophil, vulgariJ 62 puparhCl unh)i £,siopiophiJa piloll 2 puparil(u); ?FuClJ/i, sp. 2' puparh(2 unh); Deli, fabricii I puparluICn): laphne sp.

1 pupariuICel), ?Pareg/e Ip. 2 pupariaCn): Spiiogon'lp, 4 pupariaCu): "uscid BP, I pupariuICIII):

Phorlli, terr,e-nov" II puparia(lI) [Also Lepidopteran sp. 1 pupal end-segMent] tlNI" lOa

9 Qt 87 C20

Neo/eri. prollinenJ 4 puparhCeJI); HI/lo,yz, bOrl.Ji, 17 pupuia(u)i AJiopiophiJ. vulglfj, 56

388 puparh(n); Luiopiophil. pilou 5 puparia(u); SCitopftil,sp, 1 pupariull(u)j l.phn, SP, 1

pupariul(u); ?FuceIJi. sp, 9 puparia(u); ?Paregle sp, I pupariuI(ell): ~'pjJogon' SP, 7 pupaTh(eII);

?Tachinid sp, I pupariuI(u); Pho"i. terrie-nov" • puparia(u) "NIII 106

Total samples 9 Total Dipterous taxa 11 Total "NI- 2198

NORSE WESTERN SETJl.EMENT

37 EGALUGIALI K: HEAD OF ITIVDLEQ. 15 kll NW OF SANDNES (v. (5)

Y4S "Fl

HeJeollYz. borelli. 2 puparia(tII)

2 US "F2

HeJeollYz. borelli. 3 pupuia(lII) [Also Alysiine Braconid IP, I end-ngrltnt] "NI- 3

3 Y'S "F3

HeleollYu borilli. 2 puparia(tII): Deli. f,briefi 1 puparial end-seglllnt "Nl- 3

4 Y'S "F'

HeleollYz, borilli. 11 puparh(n): felollrin. fJ.vip" 1 pupariuI(u): Deli. f.brkjj 3 puparial

end-seglents,' Hydrotae ••nxi. I pupariull(el) [Also Hyaenoptera Parui tiCI SP, I I Idul t thorax

(larger Ipecies without notauli): Parasitica 'p,2 1 adult thorax (51la11er Ipecies with distinct notau1i)] "NI- 16

5 Y4S "FS

H,leoIlYz, bor"li. 10 pupafia(2 unh) (sial lest about 3.1, long); H. 'serrata; I pupariul (incolplete, ell) (would have leasured about 6111, when cOlplete: dark reddish rather thin dirty ochreoul Ind with very strong trlnsverse ridges Ilong each intersegMental boundary and IcrOS5 Middle of elch leglent- just II in typicil speeillens of this Ipecies) "NI- II

6 Y4S "F6

Si'lIliuIISP, I Idult fellie head: H,l,oIlYz. bor"li, 26 puparia(3 unh); D,li. flbriejj 2 puparial end-seglents "NI- 29

Totll la.plel 6 Totll dipterous Taxa 5 Total "NI- 6'

389 38 NIAQuSSAT (V.48)

NA I, 77 Y48 • GREENLAND

He/eo.yZl borelli. II puparil(9 n); Telo.erin. f/a'lipes 2 puparh(u): ?AJJopiopllil, vulgaris

44 puparia(el); C,JJiphori ura/ensis I pupariuI(unh); $cathopha9' fured. 2 puparia(u); Deli,

f,bricii 12 puparia(u); Botanophil, .'b,taru. 47 puparia(43 II); /fe/oph.gu,o'linus I puparial

fragMent MNI- 120

2 NA V,S -1-77 C9 I'C SAMPLE

He/eo.yz, bore.li. I wing, I pupariuI(II); fe/o.erin. f/.vipes 1 adult head; ?AJJopiophi/,

'1ulagifi. 5 puparia(u); £asiopiophi/, pilo., I pupariuI(el); ?C,JJiphori ura/Msi. I

pupariuI(II); Se,thophag' litorea I adult thorax; S, furCita 6 puparh(u); Deli, f,bricii

pupariuI(el) [Also Psyl/' sp, 2 wings (2,7 •• long): ?AlYliine sp, 2 adult thoraces]

"NI- 17 3 NA V4S -1-77 C9 20-30c.

Neo/,ri. pro.in,ns I puparhl end-seglllnt; H,/o.yz, borlliis 157 puparia(153 II): fe/olerin,

fI''1ipes 76 puparia(74 III): SCithophaga furcat, 2 puparia(II): Deli, fabridi 4 puparial end­

segment.: 'Pego.y.sp, I pupariuI(el) (posterodorsal cornu with linear window): 8otanophil' fbet.ru.

I pupariuI(el); Hydrotaea ,nxi, 2 puparia(ell) [Also Nysiu. groenl,ndieus 4 heads, 3 thoracic

fragMents] "NI- 244

3b NA-I-771 C9 30- 40c.

Neo/lfi, pro.inen. 3 puparil(e.); H,/,o.yz, bOflili. 97 puparh(95 til); ',/ollrin, fI''Iipe. 9 puparia(II): 8otlnophi/, ?b,t,ru. II Puparia(u): Deli, f,brieii 2 puparil(u): SpiJo9on'IP. pupariuI(el) MNI- 123

4 NA-I-771 C9 60-70cI

Si.uliu.lp, I adult thorax; Heleo.yz, Doreali. 341 puparh(32S eI); Keo/eri. pro.inens 2 puparh(II): Te/ollrin, fI''Iipfl 174 puparia(167 eI): SCithoph'9a furCitl 6 puparia(III);

8otanophiJ. ,'Ibetlfu, 7 puparil(u): B,profug' I pupariuI(II); Deli, f,brieii 5 puparia(e.);

D, 'p/,tUfi 2 puparia(u); Hydrot", ,nxi, 4 puparia(3 II); Phaoni, ?p,/JidisqUIlI I pupariuI(u)

390 [Also Nysius groenJ.ndicus 1 adult head; Oeltocephalid sp, 1 adult head; PsyiJ, sp, I adult thorax] HNI- 54

Ab NA-I-771 C9 90-100cl

Trichoclr.sp, I larval head capsule; Neo/eri' pro,inens 3 pupariaCII); Heleo.yz, borllli, 251 pupariaC244 ea" In one containing parasitiC HYlenopteron), 2 adult heads; TeloMerin. fJ,ripes 130 pupariaCI20 el); Agrolyzid sp, I pupariuICel); Sc.thoph,g. furc,t. 5 pupariaCell); PegolY.sp, pupariuICunh, contained cocoon of paraSite but only very few fr'gMents of an adult WISP- evidently decollposed in situ); Deli, f.bririi 2 pupariaCeI); Hydrot.l ••nxi. I pupariulI(eI); l'IeJoph.gu, orinus 1 pupariuICell) HNI- 396 Ac NA-I-771 C9 120-130cl

Tipul ••rctiel I lile pupal end-seglltnt; Heleo.yz. bore. lis IS pup.riaCn); relomerin. fl.ripes

22 puparia(21 II); PhiJygra ?rittipenni. I pupariuI(unh,); SelteJJ. ?st'gnali' I pupariulI(II);

SCithoph'g' furClt" 2 puparhCn); Bat.nophiJ, fbetlrul I pupariuIICu); ?Lllio." 'p, pupariuICunh,); Deli. f.bricii 9 pup.riaC .. ); 0, fp/.turl 2 pupariaCII); SpiJogon. sp, pup.riuICe.) [Also I Iiall .icrolepidopteroul pupal end-legMent] HNI- 56

Ad NA-I-771 C9 127,S-130c.

#eo/eri. ,'pro.inln. I adult thorax (but lesopleurae with extrllllly finl alveo!il; Heleo.yz. borllJi, '9 pupariaCel); r,/o.erin. fJ,ripes I pupariuI(unh.): fAJJopiophiJ. vlI/glril 20 puparhC 18 II); SClteiJ. "t.gn.lis 2 pupariaCIII); Sc.tophiJ. crilJrat. I pupariul(u); Selthoph'g' furCii. 2 puparia(e.): Pego'Y.lp; (polterodor .. l cornu with hrge fenestra, see Plat. 27 fig, ., 2 puparia(J unh,); Bot,nophil. !lJetlfu, 'pupariaCn); .?Lilia.,. sp. II pupariaCtII); l'phn, 'p, I pupar i UI (II);

Deli, f.bricii 16 puparh(ell; but one of these contained I Bot.nophiJ. pupariu. facing laMe direction, It is not clear how this happened); Ph.oni. ?p,liidisqu,., I pupariuI(n)

[Also 8 indet,leglented pupaeCI unh,); 2 end-seg.ents of Lepidopterous pupaeCone ending in two spines, other in I group of long Itiff hairs) HHI- 112 4d NA-I-771 C9 130-140el

391 Ne,,/,ri, pro,inens 2 puparia(ell); He/lolYl' bore. lis 10 puparia(9 ell: T,/"",rin, f/lVipls 8

puparia(u)j Allopiophi/, vu/gifis I pupariulI(efI); Scate/J, ?st.gn./is 1 pupariulI(u)j SCitophi/.

rribrat. 1 pupariuMCu)j SCithoph.g. furc.r, 3 pupariaCerrll/ ?FuCllli'lp. 1 pupariul(u)j

?L,sio",,' 5p. 3 puparil(e.>: Deli, I.bricii 13 puparil(el): SpiJogon. sp. Clinute an. sp.) 1 pupariul(el) [Also 3 end-seglents of licrolepidopterous pupie (ending in stiff hairs): 1Noctuid sp.

1 large larval head-capsule: 1 Ichneulonid or sawfly 1 cocoon (IS in sOle other salples- very pupariul- like») I1NI- 44 S NA-I-771 C9 140-8a5e

Sillllliul sp. 2 adult thoracn,' HI/eo'YI' bore.li. \I wings, 6 adult heads, 12 puparhOO el):

Neo/eri' prolinens 1 pupariul(u): Te/ollrin. fI,vipes 16 puparia(\5 lit): ?AJlopophi/, vuJg.ri. 3 wings; '7 puparia"O ul; SClthoph'g' furClt, 2 wings: 28 pupariaC25 tI): Dlli, f.Dricii 12 puparia( 11 eI): D. ?p/'tllrl 2 pupariaCu): PI90IY'lp. 'puparhCel): !SpiJogon. sp. 1 puparial Ind­ seg.entCea) (ainute Inal spiracles)j 1Huscid sp. 1 wing [Also Tlchneulonid sp. 2 cocoons] I1HI- 126

6 NA-I-77 CIO 67. SCI

He/IClIYz, borl,/i. 92 puparil(91 ea) (NB.SoI' of 'Iallest speciaens hardly larger than largest

T,Jo.erin,),' TeJollrin, f/''Iipes 34 puparia(33 ell): Hydrot.e, .nxi, 1 pupariulCn)

HNI- 127 7 NA-I.771 CIO 117.5ea

He/eolYz, borlln. 19 puparh(ea)j TI/olerin, fJlViplI 1 pupariul(ea): Bot,nophi/. ?profug. 2 pupariaCe.>j Hydrot.e, ,nxi, 1 pupariuI(el) HNI- 23

8 NA-I-771 120 CI

HI/IOIYI' borl,/i. 166 puparia(162 II) (NB. lize range confir.ed by lurstyli 3.12- '.9S •• ):

T,/oMlrin, f/''Iip,. 5' pupariaC53 III): SCithoph'g' (IIrCit. 1 pupariuI(e.): ?PegolY'lp. 1 pupariul(eI): /fe/oph.gus ovinus I pupariuI(u) HNI- 223

9 NA-I-771 09 112.5-115c.

NeoJeri' ?prollinlns 1 pupariulI(n)j HI/IOIIYZ' borelli. 2 pupariaO ell): Te/e.erin, fJ''Iipes 2 puparia(u); ?Pego'Y.lp. 2 puparia(eI)j 8ot.nophi/. 'IlJltlrUM 8 puparh(ea)j D,li. flDricii 31

392 puparia(u): 0, ?pl,tura 1 pupariuM(n): !£uioll' 'P, 1 PUpuiuICel) IAlso lchneullonid/sawfly sp, cocoon (very pupariu~-like: as above)] tlNI- 48

10 NA-I-77 D9 125c.,

Heleo.YI' oorelli, 22 puparia(21 n): relonrin. fJ.vipes 1 pupariul(u): Allopiophil' vulgaris

7 pupariaCe.): eot.nophil, !bet.ru. 1 pupariua(u): Deli, f.orieii 1 pupariulCu) [Also Ichneulonl sawfly sp I cocoon (pupariul-like as above)] HNI- 32

II NA-I-77 D9 127,S-130c.

Heleo.yz, bor"li, 1 pupariuICII): SCitella 'st'gn.lis 27 puparia(2S tI): SCitophil. cribrlt, 48 pupariaCu): !Pege.y,sp, I pupariuIlCu): Bot.nophiJ, .'betarull 5 puparia(tI): ?£lIioll" sp, 13 puparia(u); t.phne sp, I pupariuI(eM); Deli. f.brieii 18 puparia(n): 0, .'platul'l 2 puparia(II):

Spilogon. sp, I pupariuICe.),(ln,sp,O,07~.) [Also Deltocephalid 'p, I adult head: PsyJJ, 'p, I adult head: Lepidopterous 'p, 4 large larval landibles, HYlenoptera parasitica sp, I letasternuM:

Ichneulonid sp, 1 large black petiole] tlNI- 117

12 NA-I-77 D9 130cI ·Wet above underground"

Neollfi' !inseript. 3 pupariaCII): Heleo.yz, boreali, 26 puparia(23 II" one unhatched one with ellpty cocoon of HYlenopterous parasite); AJJopiophila vulgaris 5 puparlaCea): SClteiJ. ',tagn.lis 6 puparh(J unh,); ~'c'tophil' cribrlt. 5 puparhCta): Calliphora IIfllenli, I pupariulI(unh,but inCOMplete): Scathopha9' furcat. I pup,riu.(el): !Pego.YIIP, I pupariuI(II): eot.nophill !oetll'ull

'2 puparia(lJ II): ?Llliolll'IP, 40 puparia(J containing fragments of parasite); ffuCllli.sp, 3 pupariaCtI): Deli, f,orieii 73 puparhC72 II): 0, !pJ,tUI'l 1 puparluICfII); ,0. .p. 1 pupuiuletlJ

I1NI- 208

13 NA-I-77 09 130-140cI ·old ground lurface·

Neoleri, !pro.inen, I pupariuI(III); HeleollYI. borealis 5 puparla(3 III" an unhatched one with cocoon containing adult parasitic HYMenopteron): ScatophiJ. cribl'lt. 35 puparia(28 u): SCltljJ,

?ttlgn.Ji. 24 puparia(21 eM" an unhatched one containing puuite): $(lthophlgl furcat, 5 puparia(4 ell" an unhatched one containi ng cocoon of parui tic HYlnopteron): BotanophiJ, ?profug. , pupariae,.>; ?Lllio ••'lp, 12 puparia(lI tI" In unhachd one containing 1I1e lurstyli); OeJiI

393 f.fJricii 10 puparia(ea) [Also Otiorrhyncl1us 1."ticlls I adult head] MNI= 96 Total saMples 18 Total Dipterous Taxa 28 Total MNIa 2661

39 NIPAATSOQ (V. 54)

Ni V54 77-2 555 lORD FRA ONRADE ONVRIN6 HOSENS

HeleolYza borealis 9 puparia(S ell): r,lo.erin, fJ,vipes 27 puparia(2S eI): C,JJiphofl IIfllensi.

I pupariuICea> . HNIa 37 2 Ni V54 77 RUM III 1977

AedlS sp, 1 adult hud: Heleo.yza borelli, 2 puparial end-seglents; relollrin, fl,vip" 441 pupariaC431 el) HHI- 444

3 Hi YS4 77-2 RUH IV Dipt,

Heleo,yz. bor"li. 26 puparh(u): fScoJiocentfi fr.tern. 3 puparhCn); felollrin. fl,viplI 2 pupariaCel) HNI- 31 4 Hi V5. 77-2 RUM VI

HeleolYz, Dorelli. 67 pupariaCel): ~ '.errlt, 20 puparia(17 el"lostly darker, larger and lore

Itrongly tranlveru-ridged than previous): fScoJiocentrl ffltern, 103 puparh(J02 el): r,loMerin, fl,vipes 5 puparil(4 unh,): Cyno.y. lortuorll. I adult head fragll\ent: ""orMil terril-nova, 2 puparial fragMent; Scathopl/ap'lp, I pupariuI(tI): Deli, flbricii 2 puparia(ea): f:;i'pilopon.lp, I pupariuICel) [Also Ichneulonid sp, 1 cocoon] "HI- 202 HB, Dissection of the unhatched specimenl showed that 2 Heleolyzid taxa were present, nalely

Heleo,yz, bore.li. and ,'IScoJiocentra ffltern., Puparia of the two differed 15 follows:-

a generally with lore or less conspicuous transverse ridges Ie ross leglents, albulatory welts

lesl conspicuously Ipiculate ventrally and posterior spiracles sialler (0,12-0, lSI. in

vertiCil dialeter) Heleo.yz. borealis • without transverse ridges across seglents: weltl lore distinctly spiCUlate, sOleti.es vaguely protuberlnt, POlterior Ipirlcles llrger (vertical dia.tter 0,16-0,201.)

394 ?Sreliocentr. trltern.

On this basis I 5allple recounted gave 40 !SccJicClntr. and 26 HeJeo.yzl

5 Ni VS4 77-2 RUK VI

Aedes 'p, I adult thorax: Heleo.yz. borellis 2 puparia(ea): TIJollr1'n, fl.ripu 370 puparh

(354 el) "NI- 373

Total 5aMpies 5 Total taxa 10 Total KNI- 1087

'0 SANDNES (V. 51)

FELT I (21 5alples) (NB, Danish Felt- Trench)

AU I

1070 Unit P8 Level 5 v 50 (Twig)

Si.uliul.p, 3 thoraces: Hel10.yz, borllii. 27 puparia(26 l!I)j !Copro.yz. 'p, I pupariuI(el):

r,lollrin. fl.riplI,' 1 pupariuI(II)j $(lthophiga 'p, 2 wing fraglents: DeJi.lp, 1 pupariuI(el)j

?Bot.nL7phiJ.sp, 3 puparia(lI)j (lflloph.gu.orinu. 3 fragments of adulh) [Allo N.bi.

fJ,volltirgin,tu. I adult pronotulJ KNI- 40

1158 T5 54-59cI (Twig)

HllI0'Yl. borlili. II puparia(10 lI)j D,li. t,bri,,'j 3 puparia(ea): !fuClJli. 'p, I pupariuI(ea) "NI- IS

1204 Unit TIO Level SU 65 (Fire place)

forcipo.yi. 'p, 3 thoraces: Hlllo.yz, borliJi. 3 adult heads: I wing: 325 puparia(295 I!I):

?AJJopioplliJ. rulgari./ 1 pupariul(el)j SCithoph.g. turClt. 1 adult head fragltnt, I pupariuI(II)j

S, litorll I adult thorax: Deli. t.brjrU 6 puparia(l!I)j 80tanophiJ. ,'bltaru. I pupariul(lI)j

8. pretuga I pupariuI(n»i (/feJopl/lgu. Dvinus 2 fragaentl of adul h) (Aho Lepidopteron IP, pupal end-segllent: !Phi/enthul sp, I adult pronotul) "NI- 341

1277 Unit P8 Level SU 64 (Wood)

Deli. ?pJ.tUri I puparhl end segraent (AU I 397] KNI- I

(No AU II)

395 AU III

673 Unit QIO Level SU47 (Charcoal)

Si,uliu,sp, 2 adult heads and 26 thoraces; ?8rldy.j. sp, I wing; fieleo.Y1l bore,li, 4 pupariaO unh); fu{e11i.sp, I pupariuI(u); Bot.nophiJ. sp, 1 pupariuI(u) (KeJoph,gu, o~inu. 1 adult fragMent) "NI- 34

856 Unit T9 Level 75-80cI (TURF)

SClthoph'g' fur,,'. 2 larval anal regions: Bot,nophil1 sp, 1 larval skin [Also lepidopteron sp, 2 fragMents of 2 large pupae; parasitic Hymenopteron; 1 adult thorax) "MI- 3

1013 Unit P9 Level SU 19 (TURF)

Cyclorrhaphan sp, 2 puparial fragMents(unidentiable) [Also Hymenopteron IP, 1 cocoon fragment] "NI- 1

1052 Unit Q8 SU 19

fieJeolyza borelJi. 2 puparhCu); Te/o.erin. fJ,vipeJ 1 pupariuICu); fPhyto,yz'IP, 1 pupariuI;

Se.thoph.g. fure,t, 50 puparia(43 el" 3 unhatched ones containing IchneuMonid adults); 8otanoplli/,

?"etaru. 7 puparia(el) [AU III 97] "NI- 61

AU IV

302 Unit QIO Level 20-25cI

lIe/eo.yz, borfiJi. 3 wings, 5 puparia(u); H. ?serr,t, 1 pupuiuICunh): S"te11, f,t,gn,li. 2 puparhCu); Oeli, f,brieU 2 pupariaCul "'NI- 10

6331 R8 80-85C1

Si,uiiu,sp, 6 adult thomes; lIeleo.Yll bore, Ii, 3 wings, U puparia(39 ell; ',/oJerina fJ,viplf

3 pupariaCu); $Clte11, ,.t.gn,Ji. 2 puparia(,.); S"tophii, fcribr,t. I puparhl frlgllent; Pho"i, t,rr,e-nov,e I puplfi.i frag.ent,' S(.thopn,g. furClt.: 3 pupariaCell)

Delia hbricii 2 pupariaCu); flJelJJi.lp, 1 pupariuI; Bot.nophiJ. ?profug. 2 pupariaCIII)(sp,06)

(Keloph,gul ovinu. I Idult fragMent) [Also ?psychid IP, I pupa inside transparent cocoon- has anterior appendages or wing-covers; 1trichopteron sp, I wing; Pedi(u/ul sp, 5 speciMenl]

"NI- U

396 736 15/8 65-70cI (Peat)

Heieolrz• bore.iis 2 puparia(u)j Teio.erin. fi''1ipe. 1 pupariul(n}: Sc.thoph.g. furcat. 1 pupariuI(n): PegolY. sp. A pupariulI(u}: BotanophiJ. ?profuga 3 puparh(el): D,li. fabrjcij 1 pupariuI(el) [Also Lepidopteron sp. 1 pupal end segMent and sOle tergal frag.ents (as in saMple 856

KNK A 79: not at all spiny)] HNl= 12

783 Unit R8 Level SU 15 [should this be 763 1) (Hay)

Heie~llYz' borlili. 2 puparh(u)j Dlii, f,bricii 2 puparia(ea); Zaphnlsp. 1 puparhl end segMent

HNl- 5

10'1 Unit 128 Level SU 56

HliIOlYZ' borelli. I wing, 13 puparia(el): Scatell. 'p,tlgnaii, 6 puparia( .. ): PhytOlYZ'lp. 2 pupariaO el): SCithoph.g. furCit. 3 puparia(u} [AU lY liS) HNl- 2A

AU Y

350 Unit 59 Level 25-30 (Twig)

Heilol/Yz, borlili. 24 puparia(e.)j Tliolltrin. fi''1iplJ 3 puparia(u): $Cltophil. ?cribfltl 9 puparia(II): SCithophag. furc.t.,' 2 puparia(n): D,ii. flbricii 1 pupariuI(el}: Botlnophil. 'P. pupar h I end-seg.ent: SpiJ ogonl cf,.urd. 1 pupar iUI(ea) HNI- " 367 QIO Level 30-35cI (Twig)

HeilolYz, borllli. 13 puparia02 II): H, !serr.t. 2 puparia(1 II): Nlo/eri. prolinens I pupariuI:

Teiolterin. fi''1iplI 1 pupariul HNl- 17

3691 R91 level 30-35 (Twig)

/JegiSlli. ap. I pupariul(el); Hlilo,y:, borelli. 6 pupar1a(lIl: Tliollrin. fi.riplI 2 puparia(u)j SCitopllii. 'cribrit. 12 puparia(u)j SCithopll'gl furcata 1 pup.,i.l end-segMent;

D,li. f.bricii 3 puparia(u}j FU(llJii Ip. 1 puparial end-segaent [Also parasitic HYllnopteron sp.

I thorax(siall speCies)] HNI- 26

372 Unit 510 Level 30-35cI (Twig) (All speCiMens badly bleached in this laMplel

HliIOlYZ' borllli. 5 puparia(4 II): Neoilril prolinln, 1 pupariul(ul: T,io",in. fi''1ipls 2 puparia(lII)j S"tlli. "I,gn.Ji, 33 puparia(32 el): Bot.noplliillp. 2 puparh(III); DeJi. f.DricU.

397 puparia(3 ell) I1NI- 47

373 Unit 310 Level 30-35cl (Twig)

Heleo"yz. borelli, 15 puparia(era); SCltell. ?stagnali' 18 puparii(15 II); SCltophil. ?cribr.t. 3

puparh(el); Sc.thoph.g. (urCit. I puparh(unh); 11.phn, sp, I pupariuI(u); Deli. f.bricU 5

puparia( I unh); BotanophiJ. 'P, I pupariuI(el); ?Fuc,lJi. sp, I pupariul(el) "MI- 45

3741 101 Level 30-35cI (Twig)

Heleo.yZl Dore.lis 7 puparia(u); Deli. I'Dricii 1 pupariul(u) MNI- 8

399 Unit Q10 Level 25cI (Twig)

Heleo.yz. borelli. 13 puparia(u); SeatelJ. ?st.gn.lis I pupariuI(n); Philygrl vittipenni' I

pupariul(n); SCltllophag. lurelt. I pupariuI(u); (uel1li. sp, 1 pupariuI(era); Spilogon. cf, surd.

I pupariuI(el); S.sp,b 2 puparia(n) I1NI- 20

540 T9 45-50cI (Turf)

Heleo,yz.Dor"lis • puparia(u); r,lolerin. fJ.vipes: 33 puparia(31 fll); SCltoplliJ. ?cribrat, 45 puparh(44 el); $"te1l. ?st.gn.lis I pupariul(unhl; Agrolyzid sp, 1 pupariul(ell); $Clthopflag'

lureat. 20 puparia(l8 til); Pego,y.sp, 3 puparia(em): Bot.nophl, bet.ru, 2 puparia(II); 8. ?profug.

1 pupariuI(ell); D,li. I.briejj 11 puparia(el); D. ?pl.turl 1 pupariul(III),

Spilogon.sp,b 2 puparial end segMents tlNI- 124

591 Unit 59 Level 35-40cl (Twig)

SiluJiu. sp, 2 thoraces; Hel,o.yz. oor"Ji, 25 pupar11(ell; "JoJlrin. fJ.vip" 4 puparia(J unh)

?AJJopiophiJ. vu/glri. 2 wings: SCitophiJ. ?aiont. 2S puparia(22 II); Agrolyzid IP, pupariuI(el); Se,thophag' lure.t, I pupariuI(el); 8ot.nopllil. o,t.ru, 12 puparia(era); O,li.

I.briejj 5 puparia(4 II), 4 wings; ?£asio". I puparial end ugunt [AU Y 403]

I1NI- 77 AU YI

029 Unit R6 level SU 3 (Grey land layer)

Hel,o.yz. oor"Ii, 16 puparil(el); ',Jollerin. fJ.vip" 1 pupariuI(II); D,li. f.orjejj I pupariul(fI); Bot.nophiJ. !profllg' 1 puplriul(II); (1/"l1i. liP, 2 pup.rill end-segMentl(lI) [AU YI

398 21] HNI- 21

1AU 1287/R8/SU 60

HIIIOIIYl. bor"li. 1 adult head, 34 puparia(31 eI)j ?IUiopiophil' I'ulglfil 2 puparia(eM)j

SClteJl, ?st.gn.lis I pupariuI(eI)j SCithoph.g. fUI'Clt. 10 puparia(9 eI),' Deli. flbrieii 15

puparia(14 el., unhatched one containing parasitic HYlenopteron); D. ?pl.tur. 8 puparia(6 el)i

Bot.nophiJ. ?profug. 2 pupuia(eI)j Thino5C1tl1li. qUldri'l1to51 1 pupariuI(ea)j 'tl5iOll'lp.

pupariuI(el) HNI- 74

Total HNICFelt I)- 1107)

FELT II [3 Illplel]

044 MIS 'Profile Cleaning'

HI1I10'YI. Dor,.li, 6 pupuil(n); SCltIJJ. t,t,gn.li, 1 pupariulII(lI) [Aho Lepidopteron 'P.

pupal end-seglent with pale, fine Ihori spines in transverse bands)] "NI. 7

877 Felt II Gr.fi B

HIIIO'YIl bor"Ji, 12 puparil( 10 II) [Also Lepidopteron sp. 2 pupal end segaents( sue u type in

a5 in 'Profile Cleaning' salple») "NI. 12

1046 FELT II 6R0FT 8 LAG 7

Hello'YIl borelli. 133 puparia(IlS II); H, '"rflt. I pupariuI(el)j ?AIJopiophiJ. I'ulgll'i'

pupariuICeI); SCltlJJ. ,.t,gn,ii. I puparill end-segMent; $Cltnopll.g. IlIl'elt. I adult head

fragMent, I larval skin fragMent, S pupariaCu); S. litorll 3 1I1e adult hypopygia with strong

suntyli: llphnlsp. I puparill frag.ent; (/1,loph.gulol'inu. I adult fragMent) CAlso PSyJJllp.

thorax: Ichneulonid 'P. 2 thoracel) "HI- 147

12381 U 1 ?

#,I'O.Yll borelli. 2 adult heads 7 wings, 568 puparh(523 II); H, 's"rlt. 1 pupariulI; C./Jiphofl

IIflllnsi. 3 puparia(u)j D,li. ,.orieii 1 pupariuI(u)j D. tpJ,tufi 1 pupariuICellU

"HI· 574

Total HNI(Fllt II). 740

399 FELT III [3 salples]

236 Unit DI level 30-35cl (Possible dung/stable waste)

Heleo.yu bOfeilis II puparia(u)j :'c,tell, !stagn.lis 3 puparia(u)j SCithoph'gl lurCitl

pupariul(el) "NI- IS 237 + 240 Unit CI Felt III 32-35cI (Possible dung/ltable waste)

Tipul, "ctiel I fragraent of pupa: Heleo.yz, DorliJi. 9 puparia(u): Telolerin. fJ,vipe. 8

puparh(u): Sc,tell,? st.gn"i. 5 puparh(u); Deli, "Dricii 5 puparia(4 II); 8ot.nophil,

!Dltar/II I pupariulI(II); 8, !profug, 'puparia(el) "NI- 33 500 Unit DI Level 20-25cI Felt III

Hlleo.yz, bore,li. 12 puparh(n)j SCltlloph'9' furClt, 2 puparil(el); Deli, f,brieU 2

puparh(n); 8ot,nophil, fprolug' 2 puparia(el) HNI- 18

Total HNI(Felt III). 66

FELT V [I salple]

SARQAQ 5101298

Oryg., Juetuo5uI I pupariuI(e.) "NIII I

Total "NI(Felt V)II 1 I FELT VI [8 salples]

989 Felt VI COL,SA". 14, 64-68cI

Heleo.yz, bore,li. 4 puparia(el): Sc,thopll'9' (ure,t,3 puparia(e.); Deli, I,bricii 3 puparia(el)

[Also ?"icrolepidopteron sp, 1 pupil end leglent; lepidopteron IP, 1 end-legMent (Ipinole on II in other salples); parasitic HYlenopteron .p, 1 thorax (Sp, I)] "NI- 10

991 Felt VI COl,SA", 68-73cI,

f/lilygri' vittiplnni. 1 puplriul(el); 8ot.nophiJ. !profug.3 puparhl end-segllnts; Deli, Ilbricii

5 puparia(4 II); ?Pllioni, p,llidi.qu." 1 puparill end-legMent [Aho lepidopteron sp, 12 pupal end-seglents (larger, Ipiny onel, al before); Ichneulonid sp, 2 cocoons; HYlenopteron sp. 1 thorax (Sp.S)] "NI- 10

4-00 991 Felt VI Col. Sam. no. 15

Si,uliu, sp, I thorax: HeJeo,yzl borealis 1 wing: Bot.nophii, roet,ru.; 1 puparium(n)*

[Also PsyJJ, sp, 15 wings] "NI- 3 992 Level 33-36cl. Felt vi

BotanophiJ. ap, 1 puparial end leglent (as lalple 17 Ibove) "NI- 1 994 Felt VI Level 39-42cl CoI,Sal,

(I indet. cuticular fraglent, possibly frol I puplriull

995 colutn Salple nO,3 Felt VI depth 24-27cI

Si,uliu,sp, II thoraces; Delil f.oriejj 1 pupariut(ul [Also insect Ip. I coaplete genital

capsulel HNI- 12 998 Felt VI Col. SA". 10 046-52cI

HeJeo.yz. Dorllli, I pupariul(ul; Delia fabricii 6 puparia(lI) [Also Lepidopteron sp: I pupal

end seglent] "NI- 7 S.A.R.P, COL SAMPLE 9, FELT VI 42-46 (SAND)

OrYf111 Juctuosu. I adult thorlX: Deli, f,oric ii 1 puparial end-segllnt MNI- 2 S,A,R,P, COLUMN SAMPLE 12 FELT VI 55-60CI,

HeJeo,yzl Dorellis 5 puparia(ea); SCithophlf11 furcat. 5 puparia(ul; geli. f.bricjj 7 puparia(6

era): BotanophiJ, rprofug. 2 puparia(e.) "NI- 19 Totll MNI(Felt VI)- 64 FELT VII [3 Illplel] 981 Felt VII 38-39 (Natural)

[Single leaves of Betull n.n'lnd ?Sllix herDIC"]

984 Unit Felt vii Level 33-38cI (Natural)

Si.uliuISp, 2 thoraces HHI- 2

'8S Felt VII12&,S.83 (NlturI})

Si,uliuIIP, I ldult thorax [Also pUlli tic HYlenopteron; 1 thorax; Quediullp, I Idul t hlld;

(aho I Illf of ?BltuJ. n.n,,)] HHI- I

4-01 Total "NICFel~ VII)- 3

?FELT

36~/R91/Level 30-35cl [PMJonthus SP, 1 adult head]

1070 Unit P8 Level SU 50

' SP, 2 wing-fragMentl tlNI- 1

Total tlNICFelt 1)- I

Total la.plel '0 Total Dipterous Taxa 33 ·Total tlNI- 1986

41 TUMERALIK (V. 35)

la 1981 V35 TPI 35cI

Si.uliu,sp, 2 adult thoraces: He1eolYZIoore.Ji. 231 pupariaC21S n: unhatched ones cOlprised S

nles, 10 fellies and one containing cocoon of HYlI!nopterous pausite): T,lc.erin. fl.vipe. 2

puparia(n): SCithophag. fUf(.t. 6 puparia(5 II): Deli. f.iJricii 1 pupariul(lI) tlNI- 242

2 6REENLAND '81 V3S TPI 35cI

HI1eolyz.oor"li. 752 puparh044 II: 8 unhatched onu queried as H, "frlt.): fl/olefin. fl. vip,.

32 puparia(20 II): SCithophagl furc.t. 10 pupariaC7 tI): Bot.nophil, ?olt.ru. 1 pupariul(el)

[Also ?Ichneulonid IP, I cocoon(incolplete)] tlNI- 795

3 V35 TPI 4Sci

Silluliu, SP, I adult head and 3 thoraces,' HeJlo,yzl DorliJi. 31 puparil(e.>: H. ',errlt. puparial end-segment (Iuch larger than previous): Te10lerin, fl.viplI I pupariulI(unh) [Aho

Alysiine SP, 2 'anal seglents'(Sp, I): indet 'P, pupal fragments(Sp ,2): Coleopterous ap, 2 larval clypeolabrum] tlNI- 36

~ TPI 135cI

He/eelYz, berll/i, 7 puparia(u); Neo/tri' pro.inln. 1 puparial end-segMent: TI/ollerin. fl.viplI

1 pupariuI: ~"(ltl/ll 'st.gn.lis I wing, I pupariuICel): SCithoph.g. fUfClt. 2 puparia(III): Fucil/i,

'P, 31 puparia(lI)j Bet.nophiJ. !profug. SP, I pupariulCII) tlNI- U

Total I •• ple. 4 Total Dipterous Taxa 6 Total tlNI- 1117

4-02 42 LAKE 5 km NNE OF SANDNES, 2km E OF TUMERALIK (V. 36)

V36 TPI 35ell

SillUJiUI SP, I adul t head tlNl- II

Total samples I Total Dipterous Taxa I Total tlNI- 1 For Western Settle.ent sites; T018l salples 66 Total Taxa 38 Total tlNI- 6916

403 APPENDIX THREE THE MUSCIDAE OF NORTH ATLANTIC

Table 41 The Muscidae of the Canadian Arctic Archipelago, Greenland

and the coastal areas of north-western Quebec and Labrador

Species 2 3 • 5 6 7 8 9 10

Rheinll.Nltiin.e

IJu,a'na il'(ida (Huri,) x x H

IJ.• t.bul.n, (f.iiln) x H

Auiiin"

Thricops .lbibu.Ii, (Ze1terstedt) x H

T. coquiiltti (~alloch) x

T. innocuu. (ZeUerstedU X lC H

T..eptentrion.lis (Stein) X lC

T. spini;er (Stein) lC lC

T. .ubrostra tu. (Zetters tedU lC H

OrYlei, ,ipicoJ. CRondani) lC H

O. c.ntabrigilnsi' (Hucke1t> lC

O.groenJ.ndic, CLundbeck) x X lC x x AH

O. neobore.Ii, (Snyder) lC x AH

O.pri/)iJoffensi. (I1alloch) x x X lC lC

O. qu.drillto., ("alloch) X lC

O."gni, (HolMgren) X X X X lC X lC AH

O. setibui. CHuckett> x X lC

Hydrot., ••nxi, (Zetterltedt) X lC X x H

H. 'flipl' CFalUnH-occuJt.l) X X H

H,bi"c"l.t. ("eigen) x H

H, crilt.t. "alloch lC

404 Species 2 3 4 5 6 7 8 9 10

H, dentipes (Fabricius) )( H

H, floccos. (Macquart)(=lrlipes, auctt,) x H

IflJscinae

Ifese.brina /atrei/Ji R,-D, x H

IflJsc. dOlestica Linnaeus x )( H

lfydaeinBl

Ifydaea ,(finis (Meade)(=disci,ana) x H

If, tlJrtiv. Stein )(

If, nlJbill Stein x H

If,obscurlJJ. Malloch (=bengtssoTlj) x H

If. orichi/cil (Zetterstedt) )( x H

If.pI/pali. Stein x )( H

Ifyospil. ,editabund, (Fabricius' x H

Hebecne" vIsplrtin. (Fallen) x H

6rapholY' "cul.t, (Scopoli) x x )( H

Co,nosiin"

SpiJogon. Icuticorni. (Malloch) x H

S•• ,nea HUcKltt )( )( H

S•• erl' (Fallen) x H

S.lestu"ilJ' Huckett )( )( A

S•• lbisqu"a Ri ngdahl x x AH

S•• lbineplnnnis Huckett )( )( )( )( AH

S•• j.qvi.tii (Hollgren) x )( )( )( )( x )( AH

S•• lpic.

S.• ltico/. (Malloch) )( )(

5•• r(ti(0/, (Zetterstedt) )( x A

405 Species 2 3 4 5 6 7 8 9 10

S,.renos. (Ringdahl) x H

S, b.itk. (Ringdahl) x H

S,bifi.bri.t. Huckett x A

S, calcari. Huckett x

S,CoMpa,t. Huckett x

5, eonUgu. Huekett x x x A

5, contr.difrons (Zettentedt) x x x H

S,defJor.t. (Holmgren) x x x x x x x A

S,denud.t. (Hol.gren) x x x x x x AH

S,dors.t. (Zetteritedt) x x x x x x x x x x AH

S, IIpeJioguter Huckett x x A

S,extens. (Malloch) )( )( x It X A

5, f.ti•• Huckett x A

S, forU,uJ. HuckeU x x

S, genu.Ji. Huckett x

S,gl.ue. (Stein) x H

S,lIuJer./i. HuckeU )( x x x A

5, lIypopygi.Ji. Huckett x

5, in.t.n. HucktU x

S, J.tipenni. HuckiU x

S, Jeu(ogas tel' (Zetters tedt) x H

S, Ji.nophorin. (Stein) x

S,1I1.1I1i (Ringdahl> x x x x H

S,.eg.,to•• (Sohel.n) x x x x x AH

S, .el.nosoll Huckett x x x x x x x x )( A

5, .icln. (Ringdahl) x x x x H

406 Species 2 3 4 5 6 7 8 9 10

$,.inic.lyptr.t. Huckett x A

$, .on.c.n tn. Co Iiin x x x x x x x A

$,n.rin' (Walker) x

$, nigrivlntril (Zettentedt) x H

$, nititii(lud. Schnabl (. i.it.trix, l1all, ) x x x AH

$, norvlgiCi Ri ngdahl x x H

$,novl.,.cul.t. (Zetterstedt) x x H

S,nut.k, Huckett x x A

$,oDscurl (Malloch) x x x x A

$,oDs(uripenni' (Stein) x x H

$,oosolltl (Malloch) x x x x x x x x AH

5,OPIC' Schnabl x x x x AH

$,pICifiCi (Meigen) (a v.nl, Zetterstedt) x H

$, plf,.bul.t. Huckett x

$,prinClp' Huck.tt x A

$,projICtl Huck.tt X x A

5,PSlUdodisPIf (Frey) x x x AH

$,pu.ill, Huck.U x x

$, roDust. HuckeU x x x A

$, rostfltl (Ringdahl> x x AH

$,lInctip.uJi ("ilHoch) x x x x x x x x x AH

$,II.i910DOII (Ringdahl) x x x x H

$, sitinervi. HuckeU x x

$,susplcfl (Mililoch) x A

$, tornln.i. (Ringdilhl> x x x x x x AH

$, tri.nguJiflf. (Zetterltedt) x x x x H

4-07 Species 2 3 4 5 6 7 8 9 10

5. trigonit' (ZeUerstedt) x H

5. trigonif,r, (Zettentedt) x x x

S. tri /ine, t, Huckett x x

5. tund", (Schnabl) x x x x x x x AH

5. turbidipennis Huckett x x x A

5,ZiitZeYi Schnabl (. liti II,ini) )( x )( )( x x x x )( AH

5, zetterstedtii (Ringdahl> x x H

Lispoides ,equitrons (Stein) )(

Li,nophorl groenJlndi(' "aBoch x

L. nigripes (R.-D,) x x x H

L. rotund.ti Collin )(

L,.inu,t, Collin x H

L, uniset, Stein )( x x H

Lisp' (lnlden.i. Snyder x

L. cotidi,n, Snyder' x

L. t,ntl(uJlt, (DI Seer) x H

L, u/iginoSl Fal Un x H Pseudocolnosia brlvicauda Huckltt x

P,.o/it"il

Ciriee, ,I" ("ligen) x H C. breritlfsi. ("alloch) x

C,lfythro"" R, -0, x H C. vari,n. ("alloch) x

5choeno,yz, (hry,o.to •• LOlw x S, dors,li. Loell x x

S, litor,ll, (fall'n) x x H

408 Species 2 3 4 5 6 7 8 9 10

Ifacrorchis alone (Walker) x

Coenelia apiCit. HuckeU x x x x

C, anthracina Malloch x

C, .tritibi. Ringdahl x x H

C, Cln.densi. Curran x

~co.itl (Huck.tt) x x H

C, confor .. Huckett x

C, conflict. Huckett x

C,de,or./i. Huckett x

C, ,inor Huckett x x

C,lollicull (Fall'n) x H

C,lorri.oni Malloch x

C,nigrescln. Stein x x

~octopun'tat' (Zetterltedt) x x x H

C, oregonlnsis Malloch x

C, t,rslt. Huckeii x x

C, trisetl Stein x H

C, tendiplI Huc;ett x x H

C, verrilli Coll in x x H

Ph.oniinll

Helin ••nnos. (Zetterstedt) x H

H. DiS.Ji, (ZethntedU x H

H,einerell. (van der Wulp) x H

H.lvICt. (Harril) (-/iltificl, R,-D, ) x x H

H. fulvi.qu ... (ZettenhdU x x H

H. lute/squlla (ZetterstedU x x x H

4-09 Species 2 3 4 5 6 7 8 9 10

H,.aeuJipenni. (Zetterstedt) x H

H..arguer ita Snyder x

H, nigriblSis (Walker) x

H,obseurlta (Heigen) x H

H, reversio (Harris) (=duplkata, "g,) x H

H. fofhi Ringdahl (•• ubvitt,tl Seguy) x x H

H.spinosa (Walker) x

H,squ,iens (Zetterstedt) x H

Lopnoseeies fren.tus (Hoillgren) x x x x H

L•• ini .. ("alloch) x

fh,oni. ,troeitrfll1alloch x

P,ltroey,ne, Ringdahl x H

f./Jidentlt' Ringdahl x x x A

P. consobrin, (ZetterstedU x H

P.lrrang (l1eigen) x H

P,lugubri' (l1eigen) (.,orio, ZeUerstedU x H

P•• ontico/, Malloch x

p, p,J/i di.quII' (Zetters tedU x x x AH

P. protublrln. tlalloch x

P.fugi, (Walker) x x

f.serv' (l1eigen) x H

P•• ubfulCinervig (Zetterstedt) x x H

168 Species 78 81 II' 26 38 20 7 7 12 31 Key to Table 'A: This list gives the species recorded in Huckett (1965) frol the illands of the Canadian Arctic Archipelago, Boothia Peninsula, north coastal areas of Quebec and coastal areas of Labrador (all north of 10' C, July lean isother.), 1 tlelville and Boothia Peninsula: 2 Quebec(Fort

410 Chimo, Payne Bay, Sugluk and Nottinghaa Island,: 3 Labrador coast frol Cartwright to northernMost

cape; • Southalpton Island(Coral Harbour); 5 Victoria Island; 6 Baffin Island; 7 Cornwallis Island;

8 Prince Patrick Island: 9 EllesMere Island 10 6reenland, (H indicates Holarctic species; A indicates Arctic species, Casual records of no zoogeographical significance are bracketed,)

Table 42; The Muscidae of the Atlantic seaboard and Islands of the

Palaearctic

Species a, b, c, d, e, f, g, h, i, j, k, 1, I, n, 0, p: , 1. 2, 3, ',5,6,7,8, Achanthipterinae; (one palaearctic species only)

A'ha~thiptt!ra rohrellifor.js (R,-D) • •• , • ,x, .x,x, .x.x,x,x,x, Rheinwardtiinae;

Ifllscina llvida (Harris) , ,x,x,x,x,x,x, .x, x, ,X,X,X,X,X,.X, , , H If, pascuoru. (He i gen) · , , . . ,x.x, · • , ,x,x, x,x, , . , , , , , H If,prol.pSl (Harris) , •x,x,x, •x, x, x, x, x• ,x,x, x, X, x,, x, , , , , , , • H If.staDulln. (Fallen) .x. ,x,x,x,x,x, ,x, x, x, x, x, x, x, x, , , , , , ,x. • • H Synthesio.yil nudillil (Y,d,Wulp) , · •x. x. x. • . • • • • • . • • •• • • • · , • . • H Azeliinae

AZllil Iterrill (Heigen) .x • •x. x. x, X. K, X, X, x, •x, • • • · • • • • · A. "lipe. (Haliday) .x. •x. x, x, x, x. x, X, x, •x, X, x, x, , , H • • • · • A.gibbe" (Heigen) , •x, .x,x . , x. ,X, X,.X,X, • • • • · • • · · • H A•• onodactyll Loew · • · • • · • • · • • • .X. • • •• • • • · · • , • A,nlDulosl R,-O, (a,acqulrti Staeger) , • , .. .1.x, .X, x,x, x, x, x,x, x" x, ,X,X, , A. plrv. Rondani · . , • • t ,X, • f ,x, . . " · ... , · , ,

A. irigoniCi Hennig • , I , • , • • • .)(, • X ,x, •• it · . , .. , A. triquetra (Wiedelann) · .... . ,x. ,x, x, x, x, X, x, X, "x, , , .x .. H

. A, Ilttlrltldti Rondani .,',. "x, ,x,x,x,x,x,x,x,x"x,x.x. ,X". , H

411 Species ., b, c, d, I, f 19, h, i , j, k,l,., n, 0, p, ,1,2,3,',5,6,7,8,

'hrieops leu/lipes (Zetierstedt) t •••• I ,X, ,x, ,X, ,x, ,X,X, , • I I • , • , I

T. I/bibasllis (Zetterstdt) I • I ••• , x, ,x, X" • •• I • , I • H

II ••••••• I ',beekeri (Pokorny) • ••• I • ,x, ,x,

• I •• , , ,x, H " coquiJJlti (Malloch) · .... " ..

T. eunet.ns ("eigen) (=hirsutu/a, Zett.), I ••• ,X,X, ,X,X,X, ,X, X, X, X" x, x, , , ,x, , ,

T.diaphanu. (Wiedelann) ,x,x, ,X, x, X, x, X, x, x, x,, x, • • , •• I , H

T. 10'l10/ltU5 (Zltterstedt> • • I ••• , , ,X, I •••• x,x,. • •• I •• I I

r. lure,tu. (Stein) , , . ,I, , , ,x,x, • · ...... H T.gln.rul

T.llirtu/. (Zetterstedt) (·subrostrltu., Zett,) ,x, ,X,X,. I H · ... . , .. • •••• t •

T.innoeul (Zetterstedt) , , , , , , , , ,x, , , , , ,X, X"X, , , , , , , , H T.1i'lidi'lintril (ZetterstedU , • • • , • .', , • • x, x.. • · , • • • H .X,X,X,X, ,X,X,X, X,X,)(, , T. /ongiplI (Zetterltedt) · I · • , .x,x, .x.x, . , T. nigrilron. (Robineau-DlsYoidy) , , • • • • X,X. •x. X,X • ,X,X.X,X .. · • . , , · . , ',nigritlJJu. (Zetterstedt) · . · • • .X,X, .x.x,x, .x, ,x,x .. x,x, • • , • ,. ro.tfltu. (Meade) , , , , , , ,X, ,x, x, . .X,X,X,X,,)(,X, , ,X,X,

T.,e,ieintreu.

, Ory,ei, I/piea/, (Rondani) , , , , , ., , I , I II , , I , , H · , " • . . 0, (Rondanil , , , , , ,x, , brullli, I ,x, , ,x, , • " X, I X, , . , , , 0, I.scieu/,tl (Stein) . . , , , , ,', , , . , , , , , , , , , , , ,

, ,X, O,n.'lt, (Fall'n) • I • • , ,X,X, , X, X, X, X, x, ,X, x, x, , , , , ,

0, tIt" ("eigen) , , , , • , • , , • . . ,x, ,x, X" , , . , , , , ,

0, 'IiClnl (Harril) , , , , • , ,X, .X,X, . ,X, x, X, X, , x, •x, , , • , •

412 Species a, b, c, d, e, f, g, h, i, j, k, 1,., n, 0, p; , I. 2, 3, ',5,6,7,8, Hydrotaea ,enlscens (Wd.) tintr.) · , , • , .X,x. ,x, , , ,X,X,X,x" , , , , . , , . H H.IJbipuncta (Zettertstedt) , , , , , , ,x, ,x, x. , X, x, X, X, X, , , ,X, , , . , , H,anxia (Zetterstedt) · , , , , , , , , , . , .x, x" . . , , , , H H, ar.ipes (Fall'n) (·oault.. I1g. ) • x, X, , X, ,X,X,X,X,x • ,x,x. x.x, x, ,x, , X, , ,x, , . H

, H, ,trisqu," Ri ngdahl • ••• I ••• . . x., . ,x, , ,

H, /Jasdlni Coli in • t •• , I •• ,x, x, . ,x, • t It , • ,x, , , H

, , Nea) (H, crist, tl l1alloch Casual • •• I •• , , , , •• t • II • t , I , i,

H, /Jorussill Stein , ,x, ,x. , • ,x.x.x. X,, , , , , • , , ,

H, Clpensi. (Wiedeunn) ,x.x, , , ,X,x, ,X, , , x,x, , ,x, , , , , , , , , , H

H, cinerea Robineau-DesYoidy ,x, , ,X,x, , x, , , X, , ,x, II , , , , , , , ,

H,cyrtoneurin, (Zetterstedt> , , ,x,x, ,X,x, ,X,X,X,X,x" , , , , I , , ,

H, dentipt. (Fabricius) ,x, , ,x, X, X, x, X, x, X, x, x, x, X, x, x. ,x, x. x, X, x, x, H

H, diaboJus (Harr is) , • I •• ,x,X, ,X.X.X,X,X.X,X.X"x, ,x.,., H

H, fJoccosl (Hacquart) ,x, , , , ,x.x, ,X,X,X,X,X,X,X,x" t I ••• , , I H

H, gJa/JricuJa (FalUn) ,x. .X,x. ,x, . ,x.x,x,x. x,, I ••• I •••

H, hirticep. (FalI6n) , • I •• ,X,x, , , , .. . ,x. II I ••• , ••• H, ignav, (Harris) ,X,X,X,X,X,X,X,X,X,X,lC,X,X,X,X,X" . , ...... H H, irritan. (Fa1l6n) , ?, x, x, x, x, x, x, X. x, x. x. x•• x. x. x, x.

, , ,x, H, Jundbeeki Kiehillen · ...... ,x, . II , ••• , ••• H,lIridion.Ji. Portchinsky · .... ,x.x, ,x. .x. x.x,x,x, · ....

H, "teorica (Linnaeus) I •••• ,x,x, ,X,lC,X,X,X,x,x,x" , ,x, • • , t' , H

H, .iJitari. (l1eigen) ,x,x, ,x,x,x,x,x" , ,x, H · .. , .... • I •••

H, nidicol' Malloch • • , , I I •• ,x, , •. x, , • " •••• I ••• H H, p.J.,striCi (l1eigen) , ..... ,x, ,x, ,X,X,X,X,x, " , , , · .... 7 H, p.ndeJJei ,x,x, ,x, , , Stein • •••• ,x,x,x,x,' • • ? I • , • I H, p,rv. l1eade · , ... , ,x, ,x, , , ,x,x,x, " . , .. , ...

'13 Species .iI.b,c. d.e. fog, h, i. Lk,l .• , n,o. pool, 2, 3,',5.6,7.8,

H. pe//ucen! Portchinsk y • XI I I • I X, XI • XI • X, X. XI XI XI I. I • t • I I • I

•••• I ••• II •• I , • I , , H.penieill.t. (Rondani> • t ••• • ? x,

H.pilipes Stein • • I • I •• )(, ,x, •• ,x, ,x,x .. I • t t • I • I H

H.pilitibil Stein • I I • , •• I I X. I ••• I x, XI. I • I •• I I I H

H. ringdahli Stein • • I •••• , ••••• ,x,x. x,, , • I • I • , I H

H. scallbus (Zetterstedt) · , , .... , .... .• ,x. XI I , • I ••• , I H

H. sililis Heade • • I •• • ,x, .x,x, , ,X,X,X,X"X, • I • I •••

H. tubercuJ.t. Rondani , , . ,x, x, , x, I , • x. X, X•• 1 I • I •••• I H

H. ve/utin. Robineau-Desyoidy ,x, , . , ,x, x, ,x, ,x, x, x, X, x, x, , , I • I ••••

Pot"i. Jitt,1rllis Robineau-DesYoidy I • , • I x, I x, • x, x, X. X, X, X,. • • • • • • I • H

P. setifllur (Stein) I • f , ••• , ,x, • , • , ,X. I. • I ••• , • I

"uscinae

Hesellbrinl interledil Zetterstedt I • , •• I I • 1 I ••• I ,)(, II •• I •• I ••

X, X, H•• eridiln. (Linnaeus) , . , I • I XI XI ,x, x, x, x, x, x,, x, · ......

K..ystICII (Linnaeus) • X, XI ••• x. ,x. x,x, x., ......

K. resp/enden. Wahlberg ••• X, II I • I •••• I • , I •• · .)(. ....

PoJietes do.ilor (Harris) , , , .x, ,X, x, ,x, X, x, X, X, X,X, x,, x, • t •••••

P. hir ticrUfI Heade • • I •• .x. x•...••.•• • •• I • H

P.lardari' (Fabricius) ,X,X, ,X, X, x, X, x, x, x, x.. x, ,x, • •• I • .....

P."ior (Ringdahl) •••• I •• X•• 1 • • ••••• I

P. IIridionlli. Peris • Llorente ,x, , , ,x, x,x, I ••• I • I • II ••••••••

P. nigroli.b.t, (Bonsdorffl •• I X, X, t · ...... • ••• I •••

P, steinii (Ringdahl> f • I • , , , • ,x, , • ,x,x,x,x .. • ••• , • I •

Husel ,utu.nllis De 6eer I ••• ,X,X,X,X,X,X,X,X,X,X,X,X,. • t • t • , I •

H. biset, Hough ,x,x,x, , I , • t • , •• , •• I' •• I •••• I

H, dOll5ti" Linnaeul ,X,X,X,X,X,X,X,X,X,X,X,X,X,X,X,X"X,X,X, ,x,x, , , H

414 Speciu •. b, c, d. e. f. g. h. i, L k.l.l. n. 0, p, , 1.2,3.'. S. 6, 7.8,

If, Jarvipara Portchinsky , , , , ,X, X,x, , , ... , , , II I •• t • , I •

If, osiris Wiednann ,x, X, X, X, x, X, x, ,x, , ,x, x, ,x, II •••••• t •

If,sorbens Wiedelann ,x,x,x,x,x.x, , • I I •• • I' t •• I ••• I

If, tellpestiv, Fallen .x,x, . ,x.x.x.x .. ,X,X.X,XIX. It ••••••••

If, vitripennis IIfigen ,x,x, , ,X, x,x, , I • , , I •• •••••••• I I

X, X, X, X, II •• I ••••• Iforelli•• enescens Robineau-DesYoidy • •• I ,x, ,x, x, x, x, x,

I ••• I •• I ••••• I If, lfelL'Si Sannoff • I •• I ,x, • • • I •

If,hortorul (Fallen) ,x, x, ,x, x. x, x. x, x. X, x" x. x, x. x,

If,pod.gric. .)(,x ... ,x, ,x, ,x.x ... H (Loelf) • • I •

If,.i.plex (Lon) ? , .x.x.x.x,X. ,x.x.x,x.x .. x, ,x,x, '" · ... Neolyi. cornicjcn. (Fabricius) ,x, x,x, ,x.x,x, ,X,X,X,X,X,X,X,X"X,X,X,X, , ... H N, viriaescens (Robineau-De5Yoidy) · .... ,X,X, ,x,x,x,x,x,x,x,x"x,

PYfllli. riP','/( (Harris) ,x.x, ,X,X, ,x,x,x,1 .. , •••••• I I

P, '1ivia. Robinuu-Desvoidy ,x, , , , ,x,x, , , ,x,x,x,x,1,x" , , , I ••• I ? £ua.syphor. cy.nell. (lleigen) · ... ,X,X,x, ,X,X,X,X,X,x, ,X"x, ,x, . , ... £. Cf.nicolor (ZeUerltedU • •••• ,x,x, ,x,1, ,X,X,X,X,X"X, , , , , , , , (H) £,zilini (Hennig) . . ,x. , . , • ,x, ,x, x" , •••••• I

D,syphor ••lbof.sci.t. (llacquart) ,x,x,x, ,X,X,X,x, • , ,X,x, • • I •••• f ••••

0, penic i /J. I. (Egger) ,x, , , , ,X,x, , , ,X, ,x, , I , •• I •• I •••

D.pr.torul (lleigen) ,x, ,x,X,X, , , ,x, ,x, • I " •••••• t ,

StoMoxyinae

Sto.oxys "Jcitrln. (Linnaeul) ,x, x, X, X, x, X, X, x, X, X, x, X, X, x, x, x,, , , , I •• I • H

Hlelltobi. irrit.n. (Linnaeus) ,x, , ,X, ,X,X, ,X, ,X,X,X,X,X, II , ,x, (H) H,,"tobol(l .tilul.ns (lleigen) ..... ,X,X, ,x,x,x,x,x,x,x,x .. x, ,x,X, . , .. Atherigoninae

Atl/erigon. hUlerlJi. (Wiedelann) f ,x .. , ...... , ......

415 Spec its , a, b, c, d. e, f. g. h, i, j, k, I, I. n, 0, p. , 1,2,3,4,5,6,7,8,

A, naqrii Steyskal . . x... t , •••• 1 •• t t ••••• , ••••

A, orient,Jig Schiner • I X. I • I •••• I • I •••••••• I • I ••

A,pull. (Wiedelann) ,x, . , .. , •••.. I ••••• t •• t •• t I

A,socc.ta Rondani ,x, , , , t •• I t I , , , • • II , •• I ••••

A, 'I.ri. Uleigen) •x.x.x, x. , x,x • • • II ••• t ••• I

A, forti Deeling ,x .. , .... I • I •• I ••••• I • , • I I •

"ydaeinae

Ifyd. II .(fini. Meade (-di"illn. "all,), , t I • I •• ,x, XI •• )(, X.X, x...... H

, , , , , ,x, ,x, x, " x, x, X, X, , , , , , , , , If, .ndJl. "eigen . "

If, .nJ(ul. (Zetterstedt> , , , , , , ,x.x, , ,x, x, x, X, , X, , , , , , , I

If. corni (Scopoli) (aJCutIJ/ari. R. -0, ) , , I I I , , X, ,X, X, x, X, x, x, X, x, ,X, I , , , , , , ?

If, desert. (Zetterltedt) , , , , , , ,x, ,X, , , , , ,x, x,. x. , , , I , , ,

If,IJecta (Zetterstedt) , , , , , ,X,X, ,x, x, , ,x.x, . x,. , , , , . . . , H

If, hUMeri I i. R, -D. (a tinct., Zett, ) , • . • , ,x,x, ,x,x, ,x, x, X, x, X, ,x, , , , , , , , If, J.teriti. (Rondani> (-riedeJi Stein) , , , • , , ,x,

, x, If, ••culirentri. (Zetterltedt) · ... , ... ,x, ,x, I •••••• t • I

I , ,x, If, .iean, (Ringdahl) · ... , .. , .... I' • , I , • , • I , , If, nebuloSl (Stein) · , .... • x, ,x . ,x,x,x, 'I I • I •••••

If, nitidirentri, (Ringdahl> I •••••• I ••• I , • ,x, II ...... If, oblCurlJ/. Malloch · ...... ,x, ,x, x.. H

If.orielli/ell (ZttterstedU I •• I I ••••• , , ,x, X" ••• , •• I • H If, orthon,r" ("acquart) (-detritl, Zett,), .... ,x, ,x. x, , ,x, x,x, x, ,x, , , , , , , ,

If.pl/pali. Stein • •• I ••••• , •• , , ,x, X" , , , , ,x, , , H

If, rufinerri, (Pokorny) • ••••• I t I , • I ,I, • •• 1 •• , I •• t • H

If, "ti/flur Ringdahl ,X, ,x,x, , · ..... ,x,x,x,x" • ••• I •• ,

If•• ootrYlni Ri ngdahl I ••• , • I •• I • I . . ,x,x,. ••• I I ••• H

416 Species I, b,c, d,e, f,g, h, i. j, k,l, I, n,o, p, ,1,2,3,',5,6,7,8,

H H,urbana (l1eigen) , •••• I ,X,X,X,X,X,X,X,X,X,X"X,X,X, , ....

Hyospila .lpin, Hendel ,a, ,a, .. I' ••••• I I •

H,bi.,cul.t. ("acq,) (=hennigi Greg,' POY,), , ,x, ,x, x, , ,x,x,x, x,, • I ••• I

H,ledit'Dund, (Fabricius) - ,x,x,x, ,x,x,x, ,x,x, ,x,x,x,x,x .. x,x,x,x,x,x, , , H

Hebecnell .ntl/rlCin, Stein · . . x. . . . . I • , • • ••• I , I •

H, (UIOS. (l1eigen) ,x,x.x,x,x,x,x, ,)(. X, • ,x. ,x, x,, • , •••• I •

H, nigrl (R, -0,) (- vesper tin" auc tt, ) • • ,X, .x,x,x, ,x. X, x, X, x, x, x, x.. , I ••• t • I H

H, nigrjeolor (Fa1l6n) I •• I • • • X, ,x, x, x, x, x, x, x, X, ,x, , , , , , , , H

H, u.bratj" (l1eigen) • •••• ,x, x, ,x, x, x, X, x, x, x, x, ,x. x, X, ,x, , , , H

H, vesper tin,

Coenosiinae

Spilogon ••ere, (Fall'n) , , , , , ,X,X, ,x, X, ,X,X,X,X, X, ,x, ...... , H

S. acrostich.Jis (Stein) I ••• I ••• , , • . ,I, ,x, •••• I I • I • I

5, .lbisquil. (Ringdahl) • • , t I •• I • , • f •• ,x, " , I I • I I • I AH

5, .l.qvistii (Hollgren) ,x, , , , , AH • •••• I , , .... S•• lpi" (Zetterstedt> · ..... ,I, ,x, , , ,I, ,x, x, ,x, , , , ,X.X, , H

$,.rlnosl (Ringdahl) • • I , ••• I •• , •• I ,X,X .. I ••• I I •• H

5, .trisqullul. Hennig • • , •••• I ••• f , •• x. x.. • • I ••• AH

$, Dllti" (Ringdahl> , , x, , , ,X,X,X,X" ,x, , , H ...... • •• I

$, billrl,t, (Shin) I •••• t •• , x, I • X,X,X, ••• • •• , • t ••

S,brunneifrons Ringdahl (-dors.t. Rgd,), ••••• , , , , . , , , ,x, x,, · ... , .. ,

S. brunneisquall' (Zetterstedt) , , , , , , ,I, ,x, x, , ,X,X .. X, , , , , , , ,

~e'rboneJJI (Zetterstedt) , , , , • ,x,x, , , , • ,x, , x,x .. , , • , , , , ,

$, contrlCtifrons (Zettentedt) , , , , , , ,x. ,x, x, , ,x, X,X, x, ,x, , ,x, , , H

$, dlnigrat. (l1eigen) , , , , , ,x.x, ,x.x, , ,X,X,X, X,, X,X,X, , , , , ,

~denudlt. (Hollgren) , , , , , , , , , , , , , , , , , , , , , ,x, AH • "

417 Spec in a,b,c, d,e, f,g, h, i, j,k,l,., n,o, p" 1, 2,3, ',5,6,7,8,

5, depressiuscuJ. (Zetterstedt) • ••• , • . I, .X. I ••• ,X, X'I • •. •• x, ••

$,deprls,uJ. (Zetterstedt) .x, x,, I I I •••• I

,x, x, X, x, X, , , , 5, disp.r (FalUn) • •• I • ,x,x, • I , •••

5,dorl"1 (Zetterstedt) · , .. , ... , , , •• , • XI X,. , , , , , , ,X, AH

X, X, ,X, • I ••• 5,fllJIni Pont • I •••• I • ,x, ,x,?, x, x, x, x,

,1, $,grillo/I (Collin) · ...... ,x, . I' ••• I ••• I

$. Kroglruli (R i ngdahl ) ,x, I' •• I ••• I I

$, J.pponic. (Ringdahl> • •••••• t •••••• ,x, II ••••• t I •

H $.JIU(091Itlr (Zetterstedt) · ...... • • I • ,x, .. • •• I I I I •

I It. , ••• ,x, ,x, x,, $, litorll (Fall'n) (- Jongipil Rgd, ) .... • •• I • I , ,

5.Iagni(ludl Ringdahl I •• I •••••••••• ,X, t I · ......

$. '1llilli (Ringdahl> ,X. X,. • ••• I , • I H

$,lIrginiflr' Hennig (·,uginIJi" FIn, ), •• I , • , , , , , I X.X, 1.. I • I •• I • ,

$,Iuin. (Collin) t ••••• ,x, ,x,X, ,X,x,X, , "x, , , •••• t

$,II.dei (Schnabl) , , , , , , ,x, ,x, x, , , x, ,x, x, ,x, , x, , , , , ,

$,IlgI5tO" (Bohelan) , , , , , , , , , , , , , , ,x, , , , , ,x,x,x, AH " $,Iklns (Ringdahl> , , , , , , , , , , , , , , ,X, X,, , , , , ,X, , , H

$, nigrivlntri. (Zettersiedt) , , , , , , , , , , , , , , ,X, I" , , , , , , , , H

$, nitidiCiudl (Schnabl> (-depress,) , , , , , , , , , • • , • . ,x,x .. • , , , , , , , AH $, norvlgiCi (Ringdahl> , , , , , , , , , , , , , , ,x, x" , , , , , , , , H

$,novI.,,(uJ.t. (Zetterltedt) , , , , , , , , , ,x, , , , , H • • • • • " • • • •

$,oD,(uriplnnis (Stein) • I ••• I ••• t •••• ,x, x,, H

$,op.(. (Schnabl) , , , , , ,X,X .. ,x, , , AH • , • I •••• . • • I ,

$,p.(ili" (Meigen) (-vln., ZeU,) • ••••••• ,x, • ,x, x, x, x, x,, • ••• ,x, , . H

$.placid, (Huckett) (-gl,ucelJ., Rgd,) • •••••••• I •• I • ,x, II •• I • I • I • H

S,plludodispar (Frey) , ••••• , •• , I ••• ,x, ••••••• I • • AH

418 Spec ies ., b, c, d, " f, 9, h, i, j, k, 1,., n, 0, p: , 1. 2, 3, 4,5,6,7,8, H S,puberui. (Ringdahl) • •••• I ••• I •••• ,x, x,, • I I , ••••

S,pusill. (Huckett) (It.subfuscisqu,", Rgd,), , , , , , , , , , , , ,x, " , , , , , , , , H

S,PY911,e, Ringdahl , , , , , , , , , , , , •.. x. . •...•• ,

S, quinqueline,t. (Zetterstedt) • ••••••• t •• I •• ,x. X,. • ••.••• • AH

S,lIndipauli ("alloch) • •• I • I • , , . ,x. x,. • •• , • , • • AH $, "utui,t, (Schnabl> · ...... x, ,x,x,x, ... " . , .... H S"e,igioboSi (Ringdahl) • • I , • I •• · , . ... .x,x .. • •• I I •••

, •• , I • I • $, seple.not.I, (Zetterstedt) • •• f •••• ,x, ,x, x,.

$,lIplenlrion.Ii, (Ringdahl> , •• I • I •• , .X, t I • • I I . · ......

$, seligerl (Stein) • •• I • I • I •x. x, , ,x, XI )(. X,. I • I I •• I • H

S,SltuJOS' (Ringdahl) • •••••• , •••• I • ,x, x,, • •••••• I

$,.j~stedti (Ringdahl) I •• I • , , , ,x. x,, , · ... . • ••• t •

S, lolilari.n. (Collin) • I , , • ,x,x, ,x, x, , ,x, • , • ,x,x, • •••• t

• ,X,X,t •• S,sororcuJ. (Zetterstedt) • , •• I • , I · .... · . , ...

$,Iurd. (Zetterstedt) I ••••• ,x. , x. x, ,x, ,x. X•• X•• x. x, I I • I •• H

$, tenuis Hennig • ••• I •• I ••• fl' .x.

S, tornensi. (Ringdahl) ,x. •• AH · ...... · ... • r I I I I

$, tri.nguliflrl (Zetterltedt) , x. ,X,)C1t , • I •• , H • •••• t •• • t •• , x.

,x, I • S, tri,nguiiger, (Zetterltedt) • • • • t • · ... XI.X, . , .....

$, trigon.', (ZetierstedU t • t •••• I •••••• .x.x .. · , ...... H

S, triJine,t, (Huckett) • • I ••••••••• t • • I X,. I ••••••• H

S, tundr" (Schnlbl) • ••••••• , •• , I , ,x.x .. • •••• I • I AH

tundri(l (Schnabll S, • ••••••• t •••• , ,x. x•• • ••••• , , AH

S, vlrslviensi. Schnabl , , , . x, x, ,x, It H · ...... • •• t •• .? • $, veterri .. (ZeUerstedtl • ••••••• ,x, , , . x. x.x, x.. · ... , ... Vi/Jeneuri, IIstUU' (Villeneuve) · ..... ,x, .x,x.x.x.x.x, , ,.X, ......

419 Species ., b, C, d, t, f, g, h, i. j, k, I, I, n, 0, p; , l. 2, 3, " 5, 6, 7,8, 6rlpho.y. laeuJata (Scopoli) · , ... ,X,X, ,X,X, ,x,x,x,x,x"x,x,x,x, ,x, , . H

,x, x•• , ,X.X, X•• XI •• I • t , , H 6, v•• inor Robineau-Desvoidy • t I •• • I X,

NloJi,nophor. lar ili.a (Roder) · .... , ?, x, ,x, • • x,x, •••••• , • I • , I

N. virgo (Villeneuve) , , , .. . ,x, ,x, • . ,x, I • It I •••• f I I

Li.nophorl "'cKeri (Stein) I ,X, • , •••• I ••• • II ••• I •• I ,

L."ipunctlti (Stein) ,x,x,x, . ,x. , .... , , • I II •• I • I I I •

L. Clisi. (Villeneuve) ,x,x, ,x, , • II •• t • I I I • · .... • I • ,

L.,xUti (Kowarz) , , , I • • ,x. ,X, t .X,X. I ,XliX ••• I • t I •

L, fJlvitlflis Stein • ,x, I •••••• , I It •• I •••• I

L,Iaeulos. ("eigen) I •••• ,X,x, · x. x, , , x, ,x, x" ,x, , •• I ••

L, nigriplI (Robineau-Desvoidy) · .... · ,x, ,x, , ,x, ,X,x, X,, X, , • , •• I • H

L, nitiditllorax (Stein) • ,x, •• I • I •• , • , • I • II •••••• I •

L,o"scurisquIMI Stein · ,x ...... • •••••• I ., • , • I • , • ,

L. o"sign. t. Rondani ,X,x, , ,X,X,x, • •• I I ••• " I •••• I ••

L, olY'pill Lyneborg .X •• I • ,X,x, x,x, x, , ,x,x,

L,plndell,i S~guy (-rondanii Hennig) · .... ,X,x, · ... , , ,x, XII • I • I ,X, • ,

,x,X, ,x, II L,pollinifron. Stein · , , .. · ... .X, • I ••••••

L. pulchr i (IpS (Loell) . 1. •• I I I ••• I II I ••• , • , •

l. qUI tlrn. (Lon) I ,X, • , • I •• I • I • I I I II , ,

L, ripari. (FalUn) ,X,x,x, , ,X,x, ,X,X,X,X,x,X,X,X" , ,X, ,x,X, , ,

L, ruffIan. (Strobl) ,x, .•. ,x, . • • • • • I , , II I •• I ••••

L•• crupulosl (Zetterstedt) ... , , I ,X, ,x, • , • ,x,x. X, ••• ,x, , •• ,

L•• etinerv. Schnabl , ,? I • ,X,X,X, ••• , •• I • II •• I , • I , ,

L•• inuit. Collin •• , •• I r I •• I ••• , • x... , .. x.x, .. 6

(AI L. tigrinl Shin) I • I •• ,x,X, ,x,X, I XIX. X,X, , ••• • I I , , I

L, tri.nguJ. (Fll1~n) , , , , , ,X,x, ,X,x, ,x,x,x,x,x"x,x, ,x, , , , ,

420 Spec in a, b, c. d, e, f, g, h, i, j, k, 1,', n, 0, p: • I. 2, 3. ',5,6,7,8 ,

£, uniset, Stein I •• 1 t I • •• )(, I , •• ,x,x ... ,x ... I • I

,x ...•• I I • I • , ••••• 1 I •• , , •••

£, ,ssi.ilis Wiedennn ,x ... ,X, ,x .... I • I I •• 1 • , • I •• I I

£, 'lesi. "eigen t I •• • ,x,x, .x .. ,x.x,X.X, " I •••• , ••

· . x.... )(,X. I , ••••• I II I •• , • t I •

£"onsanguinea Loe. · ... .. x,x. ,x .. ,X.X, ,x, II • I •• ,x ...

£, f lal'i, incta Loell , •• I • • X.X •• I • I • I I • t. , I ••••••

£, frigid, Erichson • • , • I • I ••• I I , , ?I • II • I • f , ••• H

£,halophila Becker · .1. , , ••••••• I I ••••• I •••

7 L,hydro.yzina Fallen I ••••• , °1 ••• I X.X,X. I X••••• , ••••

£, litore, FalUn , •••••• •• )(,X •• X,X.X,)(. II , ••• I •• I

£, loe~i Ringdahl .x.x, ... x.x. ,x ... X,X.X.X. II •••• I •••

L, longi,oJJis "eigen I , , I • ,X, •• I • I .X .•••••••• I •••

L, "rin. Becker • • XI , •• ,X, ••••• I • f • I ••••••••

£, .elaieuel Lon ,X, •••• x,x, . , ••• , .. " I ••••• , ,

£, nan. "acquart ,X,X,X,X,X.XIX, ,XI' , ,X,X •• If ••••••••

L. nil'aJi. Wiedennn .x, X, • ,x, X, •• t , , • I ••• , , , , ••• , •

£, pectinipe. Becker ,X,X, • I • X, •• I •••• , •••• I ••••••

L. persi(, Becker ,x ... I •••••••• I I •• t • , • , , , , •

i, pyglll,e, Fal Un ,X, X, , , I X, X, I X, X, X, X, X, X, )(, XII , , , I •••• H

L,quaerlns Villeneuve •••• •. x, •••• I • I , • II •••• I , I I

i. rigid, Becker ,X. I •••••• I , I ••• , ••• I ••••••

£, ICIJirl'S ,.roce.na Canz.' "en, .x, .. fl. I I • t I • If. I •• I ••• I , ,

i, tentacul.t. (De Geer) ,)(,)(,)(, ,X, X,)(, ,X, X, I X, X, X, X, XII ,X, , , , , , , H

I , , •• • ,x, ,x, . ,XIX,X,X,XIIX, •• , • I •• H

Li.nospil. ,/Difron. (Z.tterlted1) , • I I • ,)(,)(, ,X, , ,X,)(, X, X, XII X, X, , I , , • , H

421 Spec in I, b,c, d, e, fog,h, i, j,k,l,l, n,o, p;.1.2. 3.'. 5,6,7,8,

,X, X, , ,X,X,X, X., X.X, IX ••• I I Pseudocoenosi, ,bnorlis Stein • , •• I •••

• ••••• I H p, soJit.ri. (ZeUerstedt) I •••• ,X.X,X. .X.X.X. x., X. e.rice, .11. ("eigen) , , , ,1,1, ,x. X, ,?, ,x. X. x,, X. ••••• t t H

C, 6istri.t. Stein · ,x, ••••••• , II •• I ••• I •

C, br.chi.Jis (Rondanil ,x .. ,X.X. ,X, , ,X,X, • • II •••• I •••

~Irythroclr' Robineau-DesYoidy · , ... .. x. ,x, X. X. x, x, X. x, x,, x, I It. , , I H

C, (,leul,t, (Collin) t • I I •••• ,x •••• ,x, • It ••••••••

C, (CoIl in) I • I I , • 1 I • t • I •• I (uscitilJi. • I I • ...... x.

C, .ikii (Strobl) ,X,X,X, ,)I,X,X. , , ••••• • I' • ••• I , ••

C, . • ,x, ,x, , , ,X.X, p,JJ ip,!pi. (Zetterstedt) · ... . x., • • I • I •• ,

C, rubricornis (Zetterstedt) • I •••• .x. ,x, X, " • ••• , I I ,

C,.puri, (Zettentedt> , , , , ,x, ,x, ,x,x. x., • , •• I • I •

C, ungul,t, (Rondani) • X, ,x.x, . , , , I • I • , It I ••• I •••

~ vern. (Fabricius) , , , , , ,x. X, ,x, X. ,x, X, x, x. X. ,x, •• I •• I • H

Orchisi. cOlt.t. ("eigen) .x .. x.x, .X,X, .X. I • It I , ••••••

Sp.noch;llt, dorSi! i, (Roser) , , , I ,X. ,X. x. X. x, x, x, •• 1 ••••• , • I

Schoeno.yz, litorl!l. (Fall'n) , X. , , , ,X.X, ,x, X. X, X, X, x, x, x,, x. x, X, X, x, , , , H

(S, JitoflJJ. I',IIjor Tiensuu · , . • XI • I ••• • II • I • I I • I .)

/facrorchisledit,t. (Fall'n) , , , , , , ,X, ,X,X, ,X,X,X,X,X"X, •••• I I •

Oexiop.i. fJ,vipes Stein · .)( . • • • • I , I' , , , •••• ,

0, I"ustri. Karl I I • I I ••• ,x,x, , ,x,x, I It , •• f •• I •

O,/,cteipenni, (Zetterltedt) , ... , ,x.x. ,X.X.X.X.X.X.X.X"X.X,x, ,x, , , ,

0, Jitor,Ji, (Zetterlbdt) • ••• , t ,x, ,X, , ,x,x,x,x, " , , .1, , , , ,

0, .inut,Ji.

0, ruficorni. ("acquart) I ••••• ,x. ,X,X, ,X.X.X,x,x" • I • I • , ••

COlnosi, ICulin.t. Strobl • •• , •••• I • , . ,x, ,x,x,. • •• I , , • ,

422 Spec in a, b, cI d, e,f I g, h, i I j I k, 1I.' n, 0, p: , ',2,3, ',5,6,7,8,

,x,x, ,x,x, ,X,X,X,X, .. , C, ,gro.yzin. (FalUn) • • I •• · . ...

C, .lbicornis Heigen • I •••• ,x, ,x,x, , ,x,x,x,x" • ? • • I • I I

C. "bulins Heigen , ,x, , , ,x,x,x,x,x, •• I •••••• ,

, x, X, , ,X, x, x, I' •••• , ••• C, .ntennat. (ZeUerstedt) • •• I • ,XIX.

C•• t,. "eigen • ••• f .X,X, ,X .• XIX,X,XIX. , ••••••••• C•• tritibi. Ringdahl · .... , , , , ,x, " • , , , , • , , H

C, • ttenll't. Stein ,x, x,x, . ,x, , , , •• " ••••• It'

I • I • ~oilinleJl. (Zetterstedt) • , •• I • ?, • ,x, .x, x,x, ,X, XliX ••• ,

C.oi'litt.t. Stein · ,x, . I • I • , , I •••• • It •• I • t •••

C. ofl'lisqllil' Fonseca • • I • I • I • .x, I • ••••• • • I •• , C, Cillpistri. (R, -0,) (·stxnot.ti "g, ) . ,x. ,x,x, ,x, ,x,x, x,, ,x, · ..... H

C, cinguJ ipes (Zetterstedt) ,x, '1 ••••••••

C, co.it. Huckett • •• t ••••••• , I • ,x, ... I ••••• I H

~de.lb.tl (Zetterltedtl .x, x,. I •••••••

C, distinguens Collin • , ••• I , • ,x, •• ,x,x, .... , , •. , •.

C, dubios. Hennig · . , ... ,x, ,x,x .. t X•• • I •• , ,

C. f"oraJiJ (Robineau-Dnvoidy) ,x, ,X,X,X,X,X,X,X,X"X,X,

C, fl''IicorniJ (Flll'n) • •••• I , , , . . x. ,x, II • I I •• I , •

C, fl''Ii•• n. (Zetterstedt) , , , . ? ,X,X, ,X,X,X,X, II , , • If' I •

C, fflyi Tiensuu , , , , x, I • I ••••••• I II • , • , I • I •

C, frlyi ft,kandid. Frey · . , , x, •• I • , • I I II ••• I •••• C, genu,J i. Rondani • I •• t • f.x, I •••••• I .t .. f • I I I •

C,grlcili. Stein , ..... ,a, t •••••• ••••• , •• I I •

C.grlCili'lentri' Ringdahl • • I , • . ,x. , •••• I • ,X ••• I ••••••

C, hisp.niensil Hennig · , ... ,x, • . , •• I •• • II , , , , .... C, hu.ili. "eigen ,x, x, x, x, • X. x, .X, ,x,x,x,x,x,x .. 7, ••••••• H

4-23 Species a, b, c, d, e, f, g, h, i, j, k, 1.1, n, 0, p: . 1. 2, 3, " 5, 6,7,8,

C, infantul, Rondani I I I •• I • XI XI I ,XI I I I' I • , I • I ••

,x,x, ,X,X, ,XIX.X,X,XIIX. t ••• , •• C, interledia (Fall'n) • • , • I

C. lyneborgi Pont · , ... ,x .. • •••• I • I I' • , I ••• I •

, ,X,X, ,X,)(,X,X,X,X,X,X"X,X, , , , , , , C, .eans "eigen I •••

C.• ixta Schnabl ,x, • , I XI X•• , , • I ••• , I I ••• I • , I ,

C•• ollicul, (Fall'n) ,X,X,X,X,)(,X,X,)(,)(,X,XIIX,X, ,X, , , " H

, I • II • , I •• I • I C. nevadensis Lyneborg ,x, , , ,x, • • •••• I

C. nigridigita Rondani · .... ,x,1, .. x...... I I ••

~ob5curllla (Rondani) I •••• •• x, .••• . ,x. , " • •••• t • t

C,octopunctata (Zetterstedt) . ,x, X. I , •••••• I H

C. DC t05ign, ta Rondani I •••• , ,x, , , • • I' •• t •• I • I

C.p,ludi' Tiensuu · .. , .... , x, ,x, • •• 1 I ••• , It'

C, plteJi iger, Rondani · ..•• . ,x .••. , . t I • I. , •••• I t I

,X, X, X, X, X, X, ,X, C,pedeJi, (FalUn) • I I • I , ,x, x, x, x, • I •• I I

C,perpusiJi, "eigen , ... , . ,)(. ,x, ... )(.)( .... )(1 • I •• , ••

,X,)(, C, praetext, t, Pandell' · , , , f • II I ......

C, pudoroll Co 11 in • • It' •• I ,x, , , ,x, , ,X" x, • I , I •••

C, pu/ic.,i, (ZetterstedU • I •• ,x, I I . . x,x, x.. , , , , , , H

C,pu.iJ, (FalUn) , ?, I ••• ,X, ,X,X,X,X,X,X,X,X"X,X,)(,X, ,x, , , H

C,pyglaea (Zetterltedt) , .x, ,x, , . x. x. x,x, .... • • I •• ......

C, rufip,lpis Heigen · , ... ,X, X, X, X, X, ,X, X, X, X, X" X, x, , ••• I •

C, slx.acul,t. Heigen I •••• .x,x ... ,)(, , .. I II • , t • I • I •

C,sexpustu),ta Rondani • •••• •• 1, • I • I ••• • II •••• I •••

C, Wood ,x, .tig.,tiCi • •••• , , . • •••• • I ••••• I I ••

C, .trigiplI Stein ,X,)(, • • • • • • • • It , • I • ••• t . , . , . C, testae" (R, -D,) (- tri(o)of, Z.tt,) ,X, , ,X,X,X,X,X,X,lC, ,lC,lC,)(,X,)("X,X,lC, .....

424 Species l,b,c, d,e, f,g, h, i, j,k, 1,., n,o, p:, 1.2,3," 5,6, 7, 8, H C, tigrin, (Fabricius) ,x, , , ,x,x,x, ,x,x,x.x,x.x,x,x"x,x,x,x, , , , ,

,x, ,x, ,x,x,x,x,x, " , ••• I I •• C, trilineell. (Zetterstedt> • • I • , •

,x, ,x,x, I • ,XIX,X" H C. verr.lI i CoIl in • I •••• ......

C, vibri5slt. Collin I • I I • I ,x,x,x ...... " I • I •• I I I

Phaoniinae

Brontle, 'felophil, (Brauer' Berg,) ,x,x, , , . ,x, ••• t •• I • I. I ••• I • I •

B,genuruf, (Pandelle) ,x, • ,X.X. I •••• t , ••• I •••••••

B, hu.ilis (Zetterstedt) I ••••• ,x, ,x, x, ,x, x, x, x, II ••••• I • I

B,po/ystig.. ("eigen) ,x, . t •• I I •••• I • I I ••••

8, tonotrui (Wiedennn) ,x,x.x. . . , .... I • I , • I •••• I

Dichllto.yi, doubleti (PandelU) I ••• I • ,x, , , , It I , I I I I , ,

Helin. ,Molin.lis (Zetterstedt) · , .... ,x, ,x, ,x,x, If ••• I ••• t H.,l/ot,JlI ("eigen) , ,x, ,x, x, ,x, x,x,x, .. , , , , , , , ,

, , , H, ./.eriensiJ (Strobl) , ,x, , , , x, , • , , " • . . . • . H..nnos, (Zetterstedt) · . • , , ,x,x, ,x, ,x, ,x, X,X, X" , , , , ,x, , • H H,.rctat, Collin ,x, , , , ,X, , , , , , , , , , " H,.t/,nti(,

H.bl/51(i (Seguy) , , , , , , ,x, , , , , , , , , , , , , , , , • " H. bolle/linni (Ringdahl) ,x, x,, H

H, , ,x,x, ,X, , calcll t. (Rondani) . . . . ,x,x,x,x, " ,x, , I ••••

H. cl/CI,tifor.i. (Schnabl) (-piuepi/oSI) , x, , ,x,x,x, ,x,x,x, ,x,x,x, ., • , ••• I ••

H,ce/s. (Harris) , , , , , , ,X, ,x,x, ,X, x,x,x, X,, , , , , , , , ,

H. cili.t, Karl , , , , , , , , , , , , , ,X, , , , , , , • " , ,

H,cililtO(O.tl (Zettentedt) , , , , , ,x,x, ,X, ,x, x, ,X,X, X,, , , , , • , , , H, cilip" (Schnlbl) , , , , , ,X, , ,x, , , , , .x.x .. , , . . , , , ,

4-25 Species ., b, c, d, e, f, g, h, i, j, k, I,., n, 0, p:, 1,2,3,4.5,6,7,8,

•• I X, •• I • XI • X.X. XI •••••••• t H H,einere//. (van der Wulp) • I ••

H,elara (Heigenl • X,X,X, • ,X,X, . , , , t I , • II •• I •••••

, , , , , , H,eon(olor (Czerny) • • I I • · ,x, • ? . x......

H, (onlini. (Fall'n) ,X,X, • X,X, ,X.X,X,X,X ... X, . ,x . • •• 1H

H, consi,i Jig (Fallen) · ..... ,x, ,x, x, ,X, X.,X, • ,x, ••• I

.. , ,x,x .. ,X. ,x, .... H H, cothurn.t. (Rondani> • I •••• • ? ,x, x.

H, crinit. Collin · ...... ,x, . , , ...... •. I •• , •

,x, x, • t " ••• I I ••• H, czernyi Lyneborg I • I • ...... ,

H, daicie. (Walker) ,x,x, •.. , .....

, x, • • • I • I I , • II • I •• f • , • H, deleta (Stein • I •••

H,depun(t. (Fall'n) · ... , ,x. X.)C. X. x. X, x, X. x, x. x.. x. , I ••• H, Ivecta (Harril) ,x,X.X, , ,x.x. ,X,X,X,X,X.X.X,X .. X.X.X, ,x, , , , H

H, fl.visqu.,. (Zetterltedt) • ••••• t •• I •• I , ,x. X" • I • , t , •• H H, fr.ter(ul. (Zetterstedt) · .... • ,x, ,X,X.X, , .X,X,X .. X, .....

H,i.pun(t. (Fallen) ,x.x, ,x,x, ,x.x.x.x,x"x.x,x • • • , I •

H, interlldi. (Villeneuve) • ••• I • ,x, ,x, •• X. ,XI • " ••• , • I •• H, Jasiophtllal.a (Hacquart) · .... ,x.x. ,x,x, ,x.x,x, , ,. H, Jatitlf.i. Ringdahl ,x.x. ,X.X, ,X.X.X,X.X" · .... • ••• I I •• H, Jaxifron. (Zetterstedtl ,I.X.X, X., · ...... H H, Jongi(orni. (Zetterstedt) H · , . , . , ... , .. . . ,x. x.. • ••• I •••

H, Jundbl.di Tiensuu · .. x....•. I ••••• I II I ••••• I • H, Juteisqu," (Zettentedt) · ...... , ,x, x...... H

H, .acuJipenni. (Zetters tedtl , , , I I ,X,X, ,X,X, I ,X,X,X,X"X,X • . . . , .. H H, .oedl ingln.i. (Schnab 1) . ,x, .... ,1, ... , ..

H, nev.O,nnoSi Lyneborg , , , , , ,x . • • , •••• , I II I ••• I • I •

H, nlvadensi. Lyneborg ,x. ,x. • ••• , •• I •••••• I ••••

426 Species a,b,c, d,e, f,g, h, i, Lk,l,., n,o, Pi ,1,2, 3, 4,5,6,7,8,

H.obs(ur.t. (Heigen) ,x,x, ,x,x, ,x,x,x,x,x"x,x, t ••••• H

H, oDscur.toides (Schnabl) • . . ,x, . , •• ,x, , . " I I I •• t • I

, , , I • '1 I ••• I , •• H.obscurisqu.M. (Stein) · ,x. • I ••

H. obtusipennis (Fallen) ,x, , x, , , I' I , t • , I • ,

H,pandeiiei (Villeneuve) I ••• , • ,x, • t I • I I ,x, II •• I , It' •

H,pendui. (PandelUl I • t ••• ,x, • • I I •• I I .t I ••• I I • t

H,perlxigu. (PandellU I • I •• · .x. · , , I •••• I •• , , • t I • I

H,pertus. (Heigen) ,X,X, ,X, x, ,X,X, X,X. I' , • , •• t ••

H.poJiinoSi (Stein) , , , , , x, , , , , , , , . I · . • " • • · • · · H.protuberans

H,quadru. (Fabricius) .X. X. , , •• X•• X, X.X, X.X, X,X. x .. X. X. ,X. I I ••

H, rlver,io (Harris) • X,X,X. ,x,X.X.X.X,X.X.X.X,X.X.X .. X,X,X.X,X, ••• H

H, ,ubvittat. (Seguy) I .X,X, ,XIX, I .X,X.X.X"X.X. . • • H

Hennig I ,X, ••• I I ••• H, rufitibi.Jis . , , It ••••••• I

H, setiventris Ringdahl , .. • X.X, ,X,X,X,X.X,X,X. II , •• , ••••

H,sex,,(ui.ta (Preyssler) . . x. • X.X,X.X • ,x. , ,X.X.X,X. ••••• t •••• H

H, ,i.u/atrix (Seguy) , , , , , ,P. , , , , I , , , , , , , , , · " • •

H. spinicosta (Zetterstedt) , , , , , , ,X.X .. I I , , H • • • • • • • • • . I •

H, squa/ens (Zethrstedt) , I I I I I ,X.X .. , , , , • • • • • • • • I • • • H H, tetrlstigll ("eigen) (.fJ'griplI,Rd,), ,x. , X. ,X,x,x,x,1, .. . , · .....

H. trivittltl (Zett,) (.,tripes, "eade) f •• I • , .X, ,x, • ,X, X, X, X, x,, ,x, · .....

H. veter.n. (Zetterltedt) • I • • • , • • I • • • • • • XI X,

4.27 Species a,b,c, d,e, f,g, h, i, j,k,l,., n,o, p; .1.2. 3, tS,6, 7. 8,

H, rieinl (Czerny) • •• I • • ,x, .X,X, • ,X, ,X, X,I •• I •••• I

H, ri/issi•• Hennig · . ,X, I ••• • ••• I .' • • II • , , I • I I I

H, riJiositusis (PandelW · .... . ,x, • •• I I • I • II I • I •• I • t

H, roekerothi Lyneborg ,x, . • • I •• , • • It ••••••••

Lop"os"Jls cinereirentris (Zett. ) ••••• "x, ,X,X, •• ,X,X,X"X", '" •• H

L, frenatus (Hol.gren) I ••• I ••••••••• . x. X, I ••••• I •• H

, I •••••• L. Mans (Zetterstedt) • I •• , I I I • I ••• , ,X. X. • H

, , • ,X. x.. x, ... I • I • L, .utatus (Fallb) • •• I • , .. •x.x.

Ph.onia IIneirentris

, , ,X, p, .JpicOJI (ZetterstedU • • I • t • ,a, ,a. x, •• , . , •• , .

P•• libi lis (l1eigen) • •• I •• ,x, .. ,x. ,x, ... I ••••••

P,lIicuJ. Villeneuve .. x. • • I •••• • I •••• I ••••

P•• ngelical (Scopoli) (-bas.lis, Zeit.). ,x, x, ,x, X, X, X, x, x, X, x,, x, x. x,

P.lnguJicorniJ (Zetterstedt) · ...... , , ,x. ,x,x,x,x" • I 1 , • , ••

P, .pic.lis Stein • • I ••••• ,x, ,x, ,x, . t I ••• t ••• H

P. I tr i"ps (Lon) • •• , • • ,X, ,x, • ,X,X,X,X, .. , ..... , . P•• troq.nea Ringdahl · ...... , ,x ...... H

p. bi tinc tl (Rondan i) • I •••• ,x, ,x, .• ,x, • • It •• I •• I • ,

P. bO/ltieo}. (Rondanil , , , , , , ,x, , , , , , , , , , , , , " , , , , P. candic.ns (PandelW , , , , , , ,x, , , , , , , , , " , , , , , , , , P.c.nesclns Stein , , , , , ,x, ,x, , , , , ,x, , " , , , , , , , , P, c".fin.t. (PandelW , , ... . ,x. · ..... II •• I , • , ••

P,cinctl (Zetterstedt) • ••• •. x,x, ,x, .. X.X.X,X, I ••••••••• P,c%rlni Collin ,x, • ••• I • I. t • , , ,

P. consobrinl (Zettentedt) • • I ••••• , x, , , , x, .x. X, .x, , ,x, .. , . H

P, Hennig czernyi · ...... • • ,X, ,x, , .... , , ...

428 Species I. b. c. d, e, f. 9, h, 1, L k, I,., n, 0, p; , 1,2,3,',5,6,7,8,

P, emlnl (~eigen) , , , , , ,1.x, .x,x.x,x,x,x,x,x"x,x,x, ,X,X, , , H

,X,X,X, ,X,X,X,X" • •• I • , I • P,errone. (Schnabl) • •• f • , ,x,

P,exoleta (Heigen) • , • , • I ,x, ,x,x, , ,X,X,X, " • •• f ••• t

P, (.JJeni "ichelsen · ...... x. . X, XI • XI XI X. X. x,. , . . I • • • •

P. (ug.x Tiensuu • •• I •••• I ••• , • ,x, ..... I ••••

P. (US" ("eade) • • t • I ••• .x ... x, It ••••••••

X. • X. P. (us"ta (FalUn) · ..... • X, •x, x, x, x• • t ••••••

p, gobertii (~ik) •••.•. • x, ,x,? ,X, ,X.XIX. 1 ••••••••

P.grl(iJif Stein • , , •••• • ,X, I .1.x,x,x,x .. t , • , ••••

P.grandaer. (Zetterstedt) • • I • I • , , , , , •. . ,x, .1 • I • I ••••

It , •• I •••• P,h./ter.t. (Stein) • • , •• I ,X, ,X,X. ,X.X,X.X,

P. hennigi Lyneborg · ... , . x...... • • II , . , .... ,

,X, P.hybrida (Schnab1) I • I • I , ,x, I ••• , • x,, • •• I • , •• H

, , , P,in,.na (Wiedennn) • •••• I ,X. ,x, X, x, x. x. x, X. X, ,X, X. X, X, x,

P,/lfo5,hersk jj (Schnabl> • •••••• •• X, I • 11, ,x, " •• I I • , ••

, , , , , , • I P. korlfzii (Schnabl> · ..... ,X ••• • ••••• , I

P.I.et. (Flll~n) ,x, ,x, ,X,X,X,X" • • •• I • , . . , ....

P, latipalpi. Schnabl • •• I , ••• ,x, , , , , , ,x,, )(, • I • I • I • P. lon9i,orni. Stein , , , ,x.X, x,, · ...... P,lugubriJ Oieigen) ,x, ,x, , , • •• I I I . .. x. x...... ,)(, P,lIgni,ornil (Zetterstedt) • ••• I I ,x, ,x,1, x, )(,X, )(., • ?t I •• I p, IIdi ter ranll Henni g ,x .... x.x.x. ,x ... ,X. I •••••• . . . . .

P,IIigeni Pont • , , •• t ,X. ,x.T.x. ,X.X,X,X, •• , • • • • I •

P, .y.ti" 01eigen) • • • • t • , • • XI X, • x, x. x, X, x...... P. nigrisqu.II Stein · ,x ...... II ••••••••

P, nYMphaliruM (Robineau-Desvoidy) • ..•• •• x, ,x .•• t ,X,X. I ••••• , •••

429 Species a, b,c, d,e, f,g, h, i, j,k, l,ra, n,o, P:, 1,2,3, (,5,6, 7,8,

P,piJJidi (Fabricius) , , , ,X, ,x,x, ,X,X,X,X,X,X,X,X"X, , , , , , , ,

P,p,JJidisqu", (Zetterstedt) I • I ••• ,x. X, •• , , , •••• H

P,pi/pata (Stein) , , , , ,X, ,X, ,X,X,X,X,X,X,X,X"X, , , x, , , , ,

,X, ,X, X, , , , , , , , , p, perdit' 01eigen) , , , , , , ,x,x, x,x, "

P, prltensis (Robineau-Desvoidy) , , , , , , ,x, ,x, , ,x,x,x,1,x,. , , , , , , , , H

••••• • II • I I •••• p, profugax (Pandel16) • •• I •• ,x, t

P,puill" (Czerny) · .... I? • I , I •• •••••••••••

P,pura (Loew) I •••• ,X.X, I ••••• , • It • I • , •••• p, /ufipa/pis ("aquart) ,X,X, ,X,X,X.X.X,X,X, " , , , ..... , p, rufjyentr if (Scopoli> • , , • , ,X.X, ,x, X, X, x, x. X, X, x, ,x, , x. , .' , , P,.cute//ata (Zetterstedt) , , , , ,1, x, , ,x, . , , , ,X, X" , , , , , , , , P,serYI ("eigen) , , , , , ,x,x, ,x,x, ,X, x, x, x, x,, , , , x, , , , . H p,.iebeck i Schnabl · .... . •x, ,x, • , t X, ,X, X, • ••• I I • , • p, sordid/squIIII Stein .X,X, I • , •• I • I • I •••••• I I I ••••

, , , p, steinii (Strobl> · .... , .. ,x,x, x,, •••• I •

,x, , ,X, • I I t p, subfusc ineryi. (ZetterstedU • •• I , , , . x,, , , , x. H p, subvent, (Harris) , , , , x, ,x, X, x, x. X, X, X, x, X. X, x, ,X, ......

p, SUI( iCl Ringdahl , X, II , • I ••••• • •• I • t . . , . , .

P, tiefii (Schnabl> , ••• I . ,x. , , , • f • ,x, II I • , I • , ••

p, trillcuJlta (Bouch.) .X.X,X.)(, ,x.x, ,x, • ,X.X,X.X, " , , , • t I I • p, #uguriorul (Scopoli) ,x, , , ,x,x,x, ,X,X,X,X,X,X.X,X"X, ,X, , , , , ,

p, Yalid, (Harril) • • I •• , x. X, ,x. X, x, x. x. x. X, x, , 7, I ••••••

P, yillan, Robineau-Desvoidy . . , .. , ,x, ,X, x, , ,x, •x, x, , X......

P,*anlbergi Ringdahl I I •••• , • , , , , , , ,x ..• •• , •• I ••

P,lugllayeria, (Schnabl> ,x. ,x, .x. ,x. x...... • I • t H Key to Tabl. 42: ., "orocco: b, Canaries: c, nadeirai d, Azorel: " Portugali f, Spain: 9, France:

430 h, Channel Isles; i, Britain; i, Ireland; k, Belgium; I, Netherlands; ., Ger~any; n, Den.ark; 0,

Sweden: p, Norway, I Inner Hebrides: 2 Outer Hebrides; 3 Orkney: 4 Shetland; 5 Faroes; 6 Iceland; 7

Jan Mayen; 8 Spitzbergen, (Species larked Halso occur in the Nearctic, whilst A indicates an Arctic species, Brackets indicate casual importations of no zoogeographical significance,)

431 7 CONCLUSIONS

This study has investigated aspects of the Quaternary history of the Dipterous fauna of the North Atlantic region and has combined two broad approaches, both requiring an ecological base. The taxonomic path can only commence with the oldest specimens available for study, but the zoogeographical route allows extrapolation to earlier times.

In the course of this research programme over 35,000 Dipterous specimens belonging to 45 families have been identified from 43 sites in West Greenland, Iceland and the United Kingdom. These sites cover a spread of dates over the last 5000 years. The great durability of

Cyclorrhaphous puparia 1s demonstrated by the fact that these comprised about 87 percent of all Dipterous specimens examined.

Since most of the area concerned in this study was ice-covered durin~ the last glaciation, a state of TBbula rasa prevailed throughout as the ice-sheets retreated. These newly exposed areas were then available for biotic colonisation. The period discussed in this Thesis commences with the retreat of the ice from this area.

In terms of the site-studies discussed here, most have been from human occupation locations of post-Bronze Age date, carried out during archaeological excavation work. Consequently none relate to a Tabula rasa situation, and many excavations of immediate post-glaCial depOSits need to be carried out before the early stages of insect colonisation of the Tabula rasa can be adequately understood. However, recent work in the High Arctic is valuable in providing models. Thus, it has been shown that the pioneer Dipterous colonisers of newly exposed areas there are ones whose larvae are aquatic, notabl y and

432 muscid predat ors belonglng to the genus Spilogona tfvlcAlpine 1965b). It

is further widely recognised that as the July mean decreases to a level

inadequate for life-processes, which in this context is very crudely

correlated with increasing latitude, the proportion of hygrophilous

members of the invertebrate fauna also increases

could therefore be reasonably postulated that these groups might have

formed the vanguard of insect colonists following the northward retreat

of the ice-sheets across the Holarctic at the close of the last glaciation. Subfossil evidence is awaited from relevant site-studies in

temperate regions to test this hypothesis.

Whilst the main emphasis of this Thesis is concentrated upon the interpretation at 6ubfossil Dipterous assemblages, it is for the reasons outlined above that a strong supporting aspect has been an appraisal of the biogeography of Diptera in the North Atlantic region, and in this connection a publication is in preparation on the Dlptera of the Western Isles of Scotland as part of a wider, North Atlantic, coverage.

A knowledge of the present distributional range of Diptera may hel p in the creat 10n of models of probable origins of the faunas of part icular regions through the occurrence of faunal harmonies. This

Thesis includes distribution tables of the Muscidae of the North

Atlantic re~ion, including the European Atlantic seaboard countries from Spain to Spitsbergen, westwards to the eastern Nearctic from

Newfoundland to Ellesmere Island. Also an updated annotated Check List of t he Muscidae of Greenland has been incorporated 1n response to particular interest in that island. " Arctic species" are defined as those species known mainly, or solely. from areas north of the 10

433 degree Centigrade July mean Isotherm, which is now widely recognised as marking the southern boundary of the Low Arct ic. On the basis of the

Muscid genus Spil ogona, it is evident that the Greenlandic fauna is of

Nearctic origin. There were probably two main routes of colonisation, namely a northern one via Ellesmere Island across the Robeson and

Kennedy channels, and a southern one. in aerial planktonic drift, across the Davis Strait from the Labrador coast. The Icelandic

Spilogona fauna. on the other hand. is of European origin and a surface-rafting mechanism of colonisation appears perfectly feasible, assuming transportation of larvae or unhatched puparia. On the basis of

Arctic elements in the Muscid fauna, Iceland has a higher proportion than any other European countrYi a conclusion which could be predicted geographically. but which is at variance with the traditional view 8S expressed by Nielsen et a1. (1954). On the other hand. and again contrary to popular belief. Scotland has no Arctic elements.

Prompted by the interest in the High Arctic insect fauna which was generated during the preparation of this Thesis, an intensive pitfall and water trap survey was undertaken in north-western Ellesmere Island by Capt. R. Godfrey during the summer of 1994 and a ,joint report is in preparation. Ttlis was part of the first ecological survey ever carried out in this pristine. uninhabited High Arctic location. and it will supplement the exhaustive work already completed around Lake Hazen, some 150 miles to the north-east by successive teams from the Canadian government. Although Capt. Godfrey's 10ca11 t y is clearly a rich area well advanced beyond the raw Tabula rasa state, the 1994 study spectacularly demonstrates that Chironomidae and Spilogona still massively dominate the fauna.

434 Previous work on the "Cow-dung Community" (Skidmore 1991> and an

interest in Arctic , prompted a fruitless literature search for

information on possible insect coprophages in Arctic environments.

Capt. Godfrey undertook an investigation of Musk Ox and Caribou dung in

his survey area on Ellesmere Island during 1994. Despite prolonged

searches through copious quantities of this material, he found no

invertebrates whatsoever, and he concluded that breakdown was achieved

purely through weathering processes. Is herbivore dung then just

another of the many niches referred to by Downes (1964) which are

unoccupied in the Arctic environment, and if so, what inhibitory

processes are operative? Also, was there a period after the retreat of

ice-sheets much further south, throughout the Holarctic, when no

coprophagous faunas existed there either? If so, how long was it before

such taxa as Aphodius appeared in these areas?

In the High Arctic the endophllous Dipterous necrophages are

Piophilidae and the Calliphorids Phormia terrae-novae and

P. (=Boreellus) atriceps; Calliphora species do not occur so far north.

Interestingly, the multitudes of Calliphorid puparia found in the

Covenden (Shropshire) mammoth were all P. terrae-novae (pers. comm. Y. Z.

Erz in

Since a major part of this study involved the identification of

fragmented Dipterous adults and immature stages. the inadequacies of

the available literature for this work were inevitably revealed. An

important section of the Thesis highlights morphological details which are relevant in subfossil, as distinct from modern, material. Regarding immature stages two examples will suffice.

1 Tipuloid larval head-capsules are otten very numerous in site-

435 samples, but their identification is impossible with existing

11 terat ure. This is because the extensive taxonomic publ1cat ions on

Tipuloid larvae focus on the morphology of the posterior spiracular and

anal regions. Extremely few Tipuloid larval head-capsules have been

figured. For the purposes of this Thesis a limited study of these

structures, in a range of Tipuloid genera, was undertaken. This

revealed an astonishing range of forms which wholly justified the

traditional view of continental authors that the Tipulidae as

understood by British workers comprises a number of distinct families.

The limited study undertaken here sufficed to show that even in the

genus Tipula the larval head-capsule exhibits a diversity which points

to a great potential for taxonomic study.

2 Initial attempts to identify Sphaerocerid puparia also proved the

inadequacy of extant literature and precipitated a re-appraisal of the

group, the results of which were published (Skidmore 1992).

Traditionally, puparial descriptions were based on specific identities

obtained from emerged adults. Work on subfossil puparia of necessity

emphasised a preferred option, namely that identifications should be

founded on unernerged adults, thus eradicat ing the constant danger of

incorrect correlat ion of puparia with hatched imagines. The present study has shown that in large assemblages of subfossll puparia, up to ten percent contained dead adults at the point of emergence. In specimens showing a good state of preservation, specific identification on male genitalic and other characters can be achieved.

Whilst there is an extensive literature on Dipterous puparia, there remains a great need for comparative morphological studies across a wide range of tamilies. Surprisingly for instance, a comprehensive

436 survey of the papillar arrangement of the "end-segments" of

Cyclorrhaphous puparia has not been produced. An in1 tial examination

for this work suggests that the characteristic arrangement found in

many Calyptrates (eg. Scathophaga, Calliphorinae and many )

appears to be very rare in the Acalyptrates. In some families unique

patterns occur (eg. Piophil1dae). The great range of puparial form

found in the is of particular interest and further work may

demonstrate a need to re-evaluate familial concepts in the superfamily

Drosophlloidea. to which the Ephydridae belongs. The Anthomyiidae is

the most problematical Calyptrate family in terms of puparial taxonomy.

A vast amount of work needs to be done here before puparia can be identified with any degree of certainty since so many taxa remain unknown in this stage of development. Also. the biological and ecological requirements of very few species are well-known. In the present connection. advances in our knowledge of Anthomyiid pupada are particularly important since they are often abundant in subfossil assembl ages.

During the stUdy of the \\iestern Settlement sites 1n Greenland, many interesting questions arose from the analysis of Dipterous material in the middens. The analysis of puparia from these features exposed a remarkable scarcity of heliophilous necrophages compared with the Palaeoeskimo site at Qeqertasussuk studied by Becher and Fredskild

(1993). This inferred not only an absence of decaying meat on the Norse middens. but of bone-marrow too. This prompted Dr.T. H. McGovern to re­ assess his data on bone fragmentation levels in the Western Settlement middens. He iound that these levels were in fact exceptionally high. strongly 6u~gesting that the inhabitants habitually extracted the

437 marrow. This discovery in turn led to a re-appraisal of the annual

round of activity on the Norse farms which revealed problems of fat

deficiency in the human diet.

It is possible that the unmistakable puparia of the Anthomyiid fly

Delia fabrtcii in Norse farm deposits in the Western Settlement reflected pasture quality. This being so, the scarcity in the Nipaatsoq samples may be highly Significant. Analysis of the far more copious samples from the nearby farm of Gus collected during 1995 may shed more light on this question. Also, the pitfall and water trap survey carried out around Gus by Phil Buckland at the same time should provide an invaluable modern insect fauna against which the subfossil assemblage can be contrasted.

In the Western Settlement middens, the occurrence of Telomerina

flavipes and HeleomyzB borealis. and absence of Piophilids and

Calliphorids was clearly indicative of discarded floor material from the farm living quarters. In that situation TelomerinB flBvipes was a highly thermophilous t roglodyt ic eusynanthrope, and in the analysis of puparia from the Nipaatsoq bedroom it helped to paint a picture of a macabre end to the human occupation of the farm. The room temperature dropped and the carrion presence, as indicated by troglodytic necrophages, rose. Lat er, it appears that sLln-loving necrophages moved in, perhaps as a result of roof collapse.

Another curioLls feature of the Western Settlement middens was the discovery of a predominance of Fucellia puparia, indicating a presence of seaweed, at a farm site which was most distant from the shore.

Elsewhere there was no Dipterous evidence for the utilization of seaweed on any of these Norse farms, despite their close proximity to

438 the coast.

It would be extremely interesting if excavated and modern pitfall

and water trap material could be obtained from the Eastern Settlement,

as possible differences between the faunas of the Eastern and Western

Settlement areas could be predicted.

It has been clearly demonst rat ed t hat at least two species of

Diptera

eusynanthropic associates of the Norsemen in the Western Settlement and

that their demise coincided with that of the settlement. That they had

been transported thence from Iceland appears to be beyond doubt.

The studies of the Norse domestic sites in the Western Settlement

and Iceland show that that by far the commonest household flies

belonged to the genus Heleomyza (serrata and borealis). The true

" ", Musca domestica was clearly absent from these islands, and

during this stUdy no specimens from anywhere north of the Scottish

Lowlands were see. It is therefore suggest ed that the "Viking Housefly" was one or other of these Heleomyza species, dependant on the area concerned. H. serrata was replaced in the colder parts of northern

Iceland and in Greenland by H. borealis. M. domestica was thought to have originated from the eastern Mediterranean. It is markedly thermophilous in northern Temperate regions and 1s probably merely a casual import through commerce further north (Skidmore 1985).

Ident it icat ion of fragments of adult D1ptera relies heavily on comparison with modern material. Very seldom are the "key characters" used in the literature available in sub fossils, and this study has demonstrated the vital need to re-examine those more durable parts of the adult Dipteran which persist 1n subfossil states. Thus, head and

439 thoracic fragments occur commonly, whilst legs are hardly ever found except in unhatched puparia. Although bristles have usually been lost, their alveoli remain as conspicuous clear dots, di ff erent thicknesses of brist les being indicated by a range of alveolar diameters. So, chaetotactic and other details of heads and thoraces can be extremely useful. A study of modern Bibionid adult heads, which are common in some terrestrial habitats, including samples from the Falkland Islands where the family appears to be extinct, showed considerable differences which have not been highlighted in the Ii terature. Similarly, in the

Tipulidae, the adult frons was found to possess hitherto apparently unrecognised features whereby Prionocera can be clearly distinguished from Tipula. The morphology section of this Thesis discusses these mat ters more fully, not ing limitat ions in our current knowledge and pointing out research areas of great potential taxonomic value.

In interpretive work based on subfossil material, a cautionary tale is offered by ThorBcochaeta zosterae. Today this is regarded as an exclusively coastal fly where it breeds only in decaying seaweed.

However, from Roman times until about the eighteenth century it was apparent ly an ubi qui tous member of the cess-fauna, as clearl y demonstrated by its great abundance in many archaeological excavations throughout the United Kingdom and on the Continent. The correct identity has been established beyond doubt on male genitalic features of unhatched adults. Formerly specimens from such deposits were misidentified as the Urinal fly Teichomyza fusca, the puparia of which have some superficial resemblance to those of T. zosterBe

1989). The reason for the sudden disappearance of this fly from inland sites, and its modern restriction in Britain to accumulations of

440 seaweed, remains unexplained.

This research programme has sought to demonstrate that Diptera can

provide a most valuable tool in the interpretat ion of past

environments, and undoubtedly Diptera will prove to be of increasing

importance 1n archaeological and post-glacial studies. Indeed, it is

recommended that all future excavation projects into such deposits

should include an investigation of the Dipterous remains. Althou8h the

number of Dipterous taxa from a given excavation rarely approaches that

of the Coleoptera, the flies usually testify to different aspects of

the biotope. They can be particularly useful in providing windows into

the human li ving conditions in a way very seldom possi ble with other

groups of Invertebrates. On the other hand, the Coleoptera are

generally far superior to Diptera in helping paint a broad canvas of

the wider vegetational environment. This is because the phytophagous

Dipterous families which include highly host-specifiC taxa (eg.

Cecidomyiidae, Tephr1tidae and ) rarely appear in subfossll

assemblages. In one respect the Diptera excel however; since with

Diptera one normally deals with the immature stages, the subfossil matrix is usually that in which the live insects were breeding. A knowledge ot the ecological requirements of the immature Diptera provides a clear statement on the nature of the matrix.

An unexpected dividend from this research has been the realisation of the great value of studies of 6ubfossil material in the taxonomy of the immature stages of modern Diptera.

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469 ADDITIONS AND CORRECTIONS p.5 Insert- 10 Adaitlons and Corrections 470 p.e Insert under Text f1gures- 19 Map of North Atlantic area showin~ various 1s1anos etc. referred to 1n the Zoogeographlcal section. 314 p. 15, 20 Thoracochaeta zosterae and MUSC3 domestiC3 were also found 1n prOfusion in Viking deposits in York

tm. mid t emur: ~c. genital c3psule; h. humer·]l callus: p. prosternum.

INltn prosternal br1stles: or. prOboscis: 55. Burst."ll: vb. v1brls53; w.

Wln~s.

p.40 Minute pup&l r~soiratory horns are pres~nt in Gal11cnorld~e

t.,ers. co:nlTJ. 't. Z. Erzin~l1o~lu).

pp.40, 143 In Cdl11ohor.:J the darkenea rod between the mouth-ho(,)KS 1S

not.:m aCcessorv ':>("31 scler1te. It ii a normally translucer.t rld~e 0"

t.he root of the oral cavity which forms 3tt3chment tor th\! mouth­

c10sin~ muscle. ana 1s orobably present in all c~clorrn"¢hous 1l3rv"~.

it 13 however sclerot1sed d1stallv. and henc~ very cons91cuous. In

';.JJ.llPnor.J ~"ers. :omm. Y. L Er::ln~l1cglu'.

p.ti5 dro3chead \L~cO) recora~ J~1cnocer.J l~rvae frem ~ hum,," coro~ •.

p. II Lar~3e ot ~catoDs~ notat~ h~ve oe~n touna 1n ~ound~nce in ~ hum~n

.;~rps.a tpres. comm. I. Z. t:r=1n<;11c~lu).

p. 14~. 1 i4 l'tlorm1..3 r.err.?tt?-nov.Je PUPllr1.., wer.~ ,)o'Jndllnt 1n th" ";'or.dO·J~r

IO-lmmotn. ana It t'.:lS oeen r4;,:lro:d t rom unnti'-sOolko,j soLl b~n«3th cr,1C1-:i:n-

·PI} runs in Siberia

Y. Z. Erzinc;lioglu).

pp.195, 196 Dr. Y.Z. Erzln~l1oglu tells me that he knows of no evidence

that Calliphora can breed in dung. as stated by Smith (le.).