LIBRARY OF TilE

JOHNS HOPKINS UNIVERSITY

COLLECTION OF THE

WILLIAM H . WELCH MEDICAL LIBRARY

Anatomical drawings of biting flies

Boris Jobling

Boris Jobling just before his retirement in 1958.

Anatotnical drawings of biting flies

Boris Jobling

Prepared for publication by David J. Lewis Department of Entomology

British Museum (Natural History) produced in association with the Wellcome Trust

Contents

Introduction 7

The artist 9

The Jobling collection 10

The species illustrated here 11

Abbreviations 12

The drawings

The sandfly- Phlebotomus papatasi 17 (Figures 1-113)

The mosquito - Aedes aegypti 49 (Figures 114-195)

The blackfly - Simulium 69 (Figures 196---239)

The deerfly - Chrysops caecutiens 81 (Figures 240-295)

The stablefly - Stomoxys calcitrans 101 (Figures 296-356) References 117

Acknowledgements

Sincere thanks are due to the following. Dr A. J. Duggan facilitated Jobling's later work, conserved his collection, and made the drawings available for publication. Dr L. A. Mound, Keeper of Entomology, British Museum (Natural History), provided working facilities for D.J.L., supported the proposal to publish this work, and provided editorial expertise. Mr Jobling and his son Mr Edwin Jobling gave much information, and Dr L. G. Goodwin, C.M.G., F.R.S., Honorary Director of the Wellcome Museum in 1984, provided facilities there. Dr R. P. Lane made many helpful suggestions. Mr Jobling, while in Egypt, received much help from the late Dr H . Hoogstraal, American Naval Research Unit No. 3, Mr H. Lomas, Wellcome Foundation Office in Cairo, and Mrs Lomas.

Introduction

The late Boris Jobling was employed by the Wellcome Organization and made numerous insect and other drawings of high scientific and artistic quality. Of these, 356 were not published, and are reproduced here to make them widely available and to draw attention to the whole Jobling collection of specimens and drawings in the Wellcome Museum of Medical Science, London, which is now administered by the Wellcome Trust (1985).

Failing eye-sight prevented Jobling from completing a text for the drawings, and in 1981 Dr A. ]. Duggan, then Director of the Museum, consulted me about them . I offered to prepare them for publication and provide an introductory text. The drawings alone, with occasional reference to Richards & Davies (1977) or similar text-books on entomology, serve as an anatomical atlas of these representative species of biting flies. Specialised publications are cited for further information.

The drawings are useful for teaching and illustrate aspects of anatomy, histology, physiology, taxonomy and medical entomology. Regarding the last, some of the sensilla, for instance, guide a fly to its host, which may be a reservoir of disease. The structure of the mouth parts determines the nature and depth of a wound, and thus influences the uptake and subsequent deposition of pathogenic organisms. The cibarium and pharynx are part of the parasites' route into and out of a fly. Saliva, secreted by glands, is essential for the feeding process and for transmission of some parasites, and often causes irritation to man. The crop of sandflies stores sugar which is necessary for the development of Leishmania. In the alimentary canal some parasites are greatly influenced by the nature of the epithelium, and by the peritrophic membrane (Gemetchu, 1974; Le Berre, 1967) which prevents many from developing. In some sandflies an abundant abdominal fat-body is an indicator of autogeny. The female reproductive organs are used in the basic estimation of physiolo-gical age in relation to disease.

One may well ask what is the place of art in a scientific work. Lapage (1961) gives the answer and points out, inter alia, that some objects depicted already have an element of beauty, that scientific preparation precedes the artist's work, and that he must exercise his talent without sacrificing accuracy. Jobling has achieved a vivid realism which calls attention to the often obscure objects portrayed.

Digitization

I rst became aware of Joblings drawings several years ago during a seminar at Johns Hopkins University in Baltimore, Maryland. Despite having worked in the medical entomology eld for a number of years, I had not previously encountered an adequate presentation of the anatomical features of hematophagous insects. Indeed, it was a common source of frustration in the lab when we would seek to identify an internal anatomical structure, most often in the context of a marked transgenic mosquito strain, that we would ultimately be resigned to make inference from the homologous features of a model insect whose anatomy was thoroughly described and readily available. Generally, this meant making comparisons with the fruit y Drosophila melanogaster, which was inadequate not only because it is not hematophagous, but also because its most recent ancestor shared with mosquitoes is estimated to have lived more than 250 mya.

It was therefore with great excitement that I obtained a copy of this work from The William H. Welch medical library to use during the preparation of my doctoral dissertation. Joblings illustrations proved so useful that I sought to acquire my own copy of the book, only to nd out that it was long out of print. Retaining copies of these beautiful sketches for my own reference was suciently important that I decided to digitize the entire work myself manually. This electronic document, representing more than 80 hours of work, is the result of that process.

Each page of the original book, including the front and back covers, was scanned at 300 dpi as an 8-bit grayscale TIFF. The content in each of these 127 images was individually aligned and centered on an 8.5 x 11 inch page, taking care to retain the scale of the gures on each page, although the original page size was A4. The text on each page was recognized electronically and made searachable. The light gray background noise of the scanned images was removed manually during processing, except in cases where it would aect the texture of the drawing. This resulted in an interesting eect on the processed drawings that I like to think slightly enhances their three-dimensional appearance.

It is my hope that converting these drawings to a modern, digital format will aid in their dissemination among researchers old and new as well as anyone who would not otherwise have had the opportunity to experience such truly extraordinary works of art.

John R. Clayton October 2012

Boris Jobling at work in 1955.

The artist

Boris Jobling's grandfather John was born about 1834, served in the Royal Navy, and became a railway engineer near Newcastle. He was a Chartist, and the unpopularity of the movement led him to emigrate to Russia. He married in 1865 and his son Edwin John became a mechanical engineer and married Agripina, a Russian who became British by marriage. Boris was born at Gorbatka, 260 km east of Moscow, . on 26 December 1893. He studied science and art at the Rial secondary school in the neighbouring town of Murom, and from 1915 to 1919 entomology and protozoology at the Shaniavsky University, Moscow. He then taught natural history at his old school and became an assistant at the Freshwater Biological Station on the River Oka. He was imprisoned briefly in the aftermath of the revolution, and in 1920 he and the late C. A. Hoare (Garnham, 1984) were repatriated. Boris crossed the Russian frontier into Finland and travelled to the port of Hanko where he met Hoare, and Ada Braithwaite whom he was to marry in 1925. They sailed in the S.S. Dongola which was sent by the British Government to fetch refugees, and reached England in March. Boris lived in the Mitcham Institute for Refugees and later in a rooming house in Charleville Road, Kensington. He died at Guildford on 26 April 1986 and is survived by his wife and a son and daughter.

Jobling worked as a private artist for Hoare, and his skill was noticed by the late C. M. Wenyon who was preparing his great work on protozoology. Jobling was taken on as a temporary artist in 1921 by the Wellcome Bureau of Scientific Research which had been founded in 1913 (Duggan, 1981), and in 1923 was appointed to the staff as entomologist by Sir Henry Wellcome. The Bureau was a unique etablishment, a scientific institute devoted to fundamental research on problems connected with tropical medicine and parasitology (Goodwin & Bruce-Chwatt, 1986). Jobling worked at Wellcome's Entomological Research Station at Wisley, later at Claremont, Esher, and during the Second World War moved to the Wellcome Research Institution in Euston Road, London. In the 1950s he visited Egypt to study Phlebotomus papatasi. He retired from the service of the Wellcome Foundation Limited on 31 December 1958 and became a Fellow of the Institute of Biology in 1966.

He published 23 papers between 1925 and 1940, 13 between 1947 and 1959, and one in 1976. Most were on Streblidae (parasites of bats), but other subjects were Phlebotomus (1976), Culicidae (1935a, 1936, 1938b), Culicoides (1928a, 1953), Muscidae (1935b), Glossinidae (1933), Hippoboscidae (1926), Nycteribiidae (1928b), the tick Ornithodoros (1925) and supports for cover-glasses (1938a). The tick paper and the first one on Culicoides were described respectively as a 'well illustrated paper' and a 'finely illustrated anatomical study' by Alcock (1925, 1928). Buxton (1955: 51), in his book on the tsetse flies, used Jobling's 'detailed and beautifully illustrated work' on the structure of the feeding apparatus.

Jobling contributed illustrations of Protozoa for several publications. Wenyon (1926) referred to his 'knowledge of biology combined with his artistic skill' and many original drawings. Hoare (1949) thanked his colleague for coloured plates, and (1972) 'was fortunate in having the assistance of my friend Mr B. Jobling, F.l. Biol., whose artistic skill and knowledge of the subject ensured faithful execution of the original illustrations and the reproduction of numerous figures from other sources'. Jobling painted malaria parasites for James (1929: 273) and James, Nicol & Shute (1938: pl. 11), and for plate 2 of Shute & Maryon (1960) which was copied for Wilcocks & Manson-Bahr (1972: 959). He also illustrated Orthoptera for B. P. Uvarov.

Goodwin & Bruce-Chwatt (1986) described Jobling as 'one of the very finest entomological artists in the classical style'.

9

The J obling collection This is in the Wellcome Museum of Medical Science, 183 Euston Road, London NW1 2BP.

General morphology of the head in Diptera There are 87 drawings, in nine groups, and 55 typed pages of unpublished text on sutures and the tentorium, and on mouth parts. The drawings include the antennae of Pangonia and Tabanus. The term sialophore is proposed for the "hypopharynx" of Diptera, which Jobling regards as the modified dorsal wall of the labium.

Drawings of the morphology of biting flies These comprise many already published, such as 57 in his 1928 paper on the head of Culicoides, and the originals of those reproduced here.

Microscope slides These include thousands of sections used for the drawings.

Protozoa

There are hundreds of drawings, including many for Wenyon's work (1926), the plates of which were lost in a fire.

The species illustrated here

The drawings were made without a camera lucida and are based on numerous measurements. They are reproduced here without reduction, as originally intended. In many of them cytological detail is shown by the difficult use of dots as described by Cannon (1936). No less than 474 parts of the insects are labelled. For each species the drawings, or groups of them, were numbered, and those in a group lettered A, B etc. All are now numbered consecutively, but the groups are kept and indicated by their sets of captions. The scale lines are all in millimetres.

For all species the descriptions of anatomy by Gordon & Lavoipierre (1962) and external anatomy by Colless & McAlpine (1970), McAlpine (1981) and Matsuda (1965, 1970, 1976) are useful. C. T. Lewis (1972) and Mciver (1982) discussed sensilla, Downes (1971) and Snodgrass (1944) reviewed trophic structure, and Detinova (1962) described age-grouping. The species illustrated here repre-sent a wide range of Diptera and of feeding mechanisms. Phlebotomus, Aedes and Simulium are in the Nematocera, Chrysops is in the Brachycera, and Stomoxys in the Cyclorrhapha. Phlebotomus, Simulium and Chrysops are pool feeders, Aedes is a capillary feeder, and Stomoxys a rasping pool feeder.

Phlebotomus papatasi (Scopoli) (Psychodidae; Figs 1-113) is an important vector of Oriental sore (cutaneous leishmaniasis), and an example of the genus which includes vectors of several forms of Leishmania. Sandfly anatomy was described by Davis (1967; some terminology not widely acepted), Just (1973), authors mentioned by them, Jobling (1976) and Rudin & Hecker (1982), and external features by Abonnenc (1972), Forattini (1973), D. J. Lewis (1973, 1975, 1982), Perfil'ev (1968), Theodor (1958) and others. Magnarelli, Modi & Tesh (1984) studied ovarian changes, and Killick-Kendrick (1979) discussed leishmania! development in relation to structure of the gut.

Aedes aegypti (L.) (Culicidae; Figs 114-195) is the well known vector of urban yellow fever, and a major vector of dengue. Christophers (1960) wrote a comprehensive book on the species, Clements (1963) dealt with features of histology and physiology, and Harbach & Knight (1980) described the external anatomy of mosquitoes.

Simulium (Wilhelmia) sp. (Simuliidae; Figs 196--239) may comprise more than one species. The figures, originally labelled S. equinum L., were drawn before it was known that the related S. lineatum Meigen differs from it in having a sub-ovoid spermatheca (Crosskey, 1981: 147) very like that shown in Figs 214, 229 and 235. The head of a Wilhelmia was studied by Wenk (1962). One of these species bites horses in Britain (Edwards, Oldroyd & Smart, 1939), and S. lineatum attacks cattle in Europe. The simuliids or black-flies include notorious man-biters in many countries, and vectors of Onchocerca volvulus Leuckart, the cause of river blindness (Crosskey, in press; Laird, 1981) in Africa and tropical America.

Chrysops caecutiens L. (Tabanidae; Figs 240--295) is a common European man-biting species (Edwards, Oldroyd & Smart, 1939) and is related to vectors of human loiasis in Africa (Oldroyd, 1957; Service, 1980). The biting process in Chrysops has been described by Gordon & Crewe (1953), and aspects of anatomy by Lavoipierre (1958).

Stomoxys calcitrans L. (Muscidae; Figs 298-356), the stable fly, is a widespread biting pest of man and animals (Greenberg, 1971; Service, 1980). Its subfamily has been reviewed by Zumpt (1973), and its peritrophic membrane studied by Lehane (1976).

11

Abbreviations

Notes Sial: this is the hypopharynx (of Diptera) of most authors . Ven: used here for the abdominal part of the mid-gut 9 stn: sternite 9 of male P. papatasi. The coxite arises, not from this, but from the phallobase, according to Just

(1973). 10 stn: sternite 10 of the same. This is sternite 9, according to Just (1973). Various: in some cases other authors have used different names.

a antenna cb. lm. tr cibario-labral trachea abd. gng abdominal ganglia cb. pmp cibarial pump abd. nv abdominal nerve cd car do abd. spr abdominal spiracles cer cerci ac. gl accessory glands cg. eel corneagen cell adv adventitia (pericardium) cho chorion aed aedeagus cir. m circular muscles aed. p aedeagal pouch cl clypeus af antennifer cl . ap clypeal apodeme air. s air sac de. chn collecting channel al. m alary muscles elf cleft amp ampulla clge. sut clypeogenal suture an anus cl . nv clypeal nerve ant. ar. t anterior arm of tentorium clsp claspette ant-I. t antero-lateral trachea cl. sut clypeal suture ant. thor. spr anterior thoracic spiracle clx calyx ant. t . p anterior tentorial pit cmd. e compound eye ant. w anterior wall em. ovd common oviduct a. nv antenna! nerve cmp. snl campaniform sensilla a or aorta em. sl. d common salivary duct aor. sin aortic sinus comp. m compressor muscle ap apodeme en canal api. eel apical cell cnn. stk connecting stalk api . snl apical sensilla con connective a. pul. org. antenna! pulsatory organ cor corneal lens ar arista cox coxite asc ascoids cp. cl cap cell as. nur association neurone cr crop a. tr antenna! trachea ern cornua a. ves antenna! blood vessel crp . al corpus allatum ax . aps axial apophysis crp. card corpus cardiacum ax. fb axial fibre of sense cell crp . card. nv corpus cardiacum nerve axn axon cv. con cervical connective

cv. nv cervical nerve bld . eel blood cells or haemocytes cv . sc cervical sclerite b . mem basement membrane ex. nv coxal nerve br brain ex. 1, 2, 3 coxa of fore, middle and hind legs bs . pl basal plate bs. snl basiconic sensilla d duct buc bucca d. ac. gl duct of accessory gland burs bursa copulatrix d. cr duct of crop

dr. ar. t dorsal arm of tentorium car cardia dr. arh sg dorsal arch of sigma cb cibarium dr. com dorsal commissure

12

ABBREVIATIONS

dr. c. trn dorsal cephalic trunk hist. m histolysed muscles dr. dphr dorsal diaphragm hl. tr haltere trachea dr. dv dorsal diverticulum hphy hypopharynx dr. long. tg . m dorsal longitudinal tergal muscle ht heart dr. mes. nv dorsal mesothoracic nerve hy hyoid dr. met. nv dorsal metathoracic nerve hyp . gng hypocerebral ganglion dr. nv dorsal nerve dr. prt. nv dorsal prothoracic nerve i. eel interstitial cells dr. sin dorsal sinus il ileum dr. trn dorsal trunk in. pre inner process dr. w . lm dorsal wall of labrum ins insula

int intima ej. d ejaculatory duct in. tbr inner tuberosity of mandible en. eel enveloping cell in. to inner tooth epi epithelium intrb . chn interbifid channel epi. eel epithelial cell in. w inner wall epid epidermis ir . pg. eel iris pigmented cells eps episternum e. rig epistomal ridge J. 0 Johnston organ e. sut epistomal suture

f frons kl keel

fa) falciform process 1. abd. trn lateral abdominal trunk fat . b fat body larv. m larval muscles f-cl fronto-clypeus lb labium fd. en food canal lb. g labial gutter fgl. sut frontogenal suture lbl Ia bellum f. gng frontal ganglion lbl. cls labellar canaliculus f. gng. con frontal ganglion connective lbl. frc labellar furca fl filament lbl. gl labellar glands fie fulcrum fig flagellum lbl. pre interlabellar process lb. mem labial membrane fol follicle lb. nv labial nerve fol. epi follicular epithelium lb. pis . org labial pulsatory organ frc furca lb. tr labial trachea frk fork lc lacinia fun funicle le lever ga galea lg. nv leg nerve ge. rd genal rod lg. tr leg trachea gen. ch genital chamber or atrium lig ligament gen . fl genital filament 1. lob lateral lobe of tenth abdominal gen. frc genital furca segment gli glia 1. long. tg. m lateral longitudinal tergal muscle gls glossa lm labrum gl. vs . sem gland of vesicular seminalis lm. nv labral nerve

lm. tr labral trachea gon gonopore Ink premento-labellar and mandibulo-grm germarium

grm. eel germ cells genal links gs cae gastric caecum log. 8 stn lobe of eighth sternite gs. nv gastric nerve long. dr. m longitudinal dorsal muscle gu gula long. m longitudinal muscle gus. snl gustatory sensilla long. stn. m longitudinal sternal muscle

lor. ar loral arm h haematin 1. ovd lateral oviduct hin-g hind-gut lp lip

13

ABBREVIATIONS

1. pr lateral process of labial gutter 33 salivary pump muscle 1. thor. trn lateral thoracic trunk 34 levator of head lv laterovertex 35 compressor muscle of sac of 1. w. lm lateral wall of labrum pompetta

36 protractor of genital filament m:- muscles 37 depressor of ninth abdominal

m. tro muscle of trochanter 1 levator of scape, in some abductor of segment 38 levator of tenth abdominal segment

same 39 adductor of style 2 depressor of scape, in some 40 abductor of style abductor 41 rotator of style 3 retractor of scape; in some may be 42 levator of coxite abductor 43 abductor of coxite

4 extensor of pedicel 44 adductor of coxite 5 rotator of pedicel 45 abductor of claspette, in some 6 flexor of pedicel depressor 7 tormo-labral muscle 46 adductor of claspette, in some 8 anterior labral muscle levator 9 adductor of mandible 47 depressor of aedeagus, in some 10 abductor of mandible retractor 11 tentorial adductor of mandible 48 levator of aedeagus 12 tentorio-cardinal muscle or

protractor of maxilla 49 depressor of claspette 50-53 muscles of tenth abdominal segment

13 tentorio-stipital muscle or retractor of stipes

Mal. tu Malpighian tubes 14 cranial flexor of lacinia or in some lacinia retractor of same md mandible

15 geno-stipital muscles or diagonal md. end mandibular condyle adductor of stipes md. max. tr mandibulo-maxillary trachea

16 stipito-palpal muscle or levator of md. nv mandibular nerve maxillary palpus; in some may be mdv mediovertex abductor of same or retractor of mem membrane maxilla mes. gng mesothoracic ganglion

17 depressor of maxillary palpus, in met. gng metathoracic ganglion some abductor mid . cr. sut midcranial suture

18 levator of second palpal segment mid-g mid-gut 19 depressor of second palpal segment mit mitosis 20 retractor of prementum, or labium msn . inv mesenteric invagination 21 abductor of first segment of labellum mt mentum 22 adductor of first segment of mtr. nur motor neuron

labellum, in some flexor mx maxilla 23 abductor of second segment of mx. gl maxillary glands

labellum, in some extensor mx. lob maxillary lobe 24 anterior cibarial dilator mx. nv maxillary nerve 25 posterior cibarial dilator mx. pip maxillary palpus 26 retractor of cibarial pump 27 protractor of cibarial pump, in some n nucleus

depressor n. epi . eel nucleus of epithelial cell 28 sphincter buccalis nephr nephrocytes of epicardial cells 29 dilator buccalis nlm neurilemma 30 precerebral dilator of pharyngeal nr. eel nurse cells

pump ns. eel neurosecretory cells 31 postcerebral dorsal dilator of nv nerve

pharyngeal pump nv. cr nerve to crop 32 posterior lateral dilator of nv. hi nerve to haltere

pharyngeal pump nv. lam neural lamella

14

ABBREVIATIONS

nv. sl. d nerve to salivary duct pof. sut postfrontal suture nv. wng nerve to wing ppd postpedicel

pre process 0 ocelli pre . w. lm preoral wall of labrum oc occiput prg proctiger oc. end occipital condyle prit. mem peritrophic membrane oc. for occipital foramen prmt prementum ocl. m occlusor muscle prn pronotum oclr. sc ocular sclerite prnv perineurium oclr. sut ocular suture prot protuberance oclr. tr ocular trachea prp periopticon or lamina ganglionaris oc. sut occipital suture prs prescutum oe oesophagus pr. sh peritoneal sheath oen oenocytes prst prestomium olf. p olfactory pits prst. sc prestomal sclerite olf. snl olfactory sensilla prst. te prestomal teeth ooc oocyte prt. gng prothoracic ganglion oog oogonium pst. ar. t posterior arm of tentorium oos oosome pst-1. trn postero-lateral trunk op openings of ducts pstn postnotum or postscutellum op. ac. gl opening of accessory gland pstr pseudo trachea op. sp . d opening of spermathecal duct pstr. rng pseudotracheal rings opt. lob optic lobe pstr. slt pseudotracheal slit opt. nv optic nerve pstr. te pseudotracheal teeth ost ostia pst. t. p poterior tentorial pit ost. vl ostial valve pst. w posterior wall out. chia outer chiasma pt ptilinum out. pre outer process of mandible ptl. blad petiolate blades out. w outer wall ptr peritreme ov ovary pt. sut ptilinal suture ovl ovariole pv postvertex ovl. ped ovariole pedicel pyl pylorus ovl. sh ovariole sheath pyl. sph pyloric sphincter muscle ov. sh ovarian sheath

r ring ped pedicel rad. str radial striation peg peg rb rabdom pen penniculus rd rod pg. eel pigment cell rec. nv recurrent nerve pge postgena rect rectum pge. brg postgenal bridge rect. p rectal papilla pgen. p postgenital plate ref refractive spheres pg. gr pigment granules reg. eel regenerative cells pgl paraglossa rem. fol. epi remainder of follicular epithelium ph pharynx res reservoir ph. pmp pharyngeal pump ret. eel retinal or sense cells phr phragma ret. pg eel retinal pigment cells plf palpifer rig ridge pls. mem pulsatory membrane rst. mem rostrum membrane pls. or pulsatory organ pm pompetta sarc sarcosomes pmt postmen tum sc sclerite poe pas tocci pi tal sci scolopophores poe. sut postoccipital suture sci. org scolopophorous organ poe. rig postoccipital ridge sco scolops

15

ABBREVIATIONS

scp scraper tn. 11 tendon of tentorial adductor of sctl scutellum mandible sctl. pis . org. scutellar pulsatory organ tor torma scut scutum tr trachea si siphon trh tracheoles sial sialophore (hypopharynx) tr. 1, 2, 3 tracheae of fore, middle and hind sl. en salivary canal legs sl. d salivary duct tro. 1, 2, 3 trochanters of fore, middle and hind sl. gl salivary glands legs sl. pmp salivary pump tun. pr tunica propria smt submentum tu. sh tubular sheath snl sensilla sns. eel sense cells v vertex sns. or sense organ vas. de vas deferens sns. p sensory pits vas . ef vas eferens so sockets of sensilla vbr vibrissae sam. eel somatic cells ven ventriculus sp spermatheca vii villarium spat spatulate rods vl valve spd spermatids vnt. arh. sg ventral arch of sigma sp. d spermathecal duct vnt. c. trn ventral cephalic trunk spg spermatogonia vnt. com ventral commissure sp. gl spermathecal glands vnt. dphr ventral diaphragm sph. m sphincter muscle vnt. nv ventral nerve spi spicules vnt. nv. crd ventral nerve cord spm spermatozoa VS vesicle s. pm sac of pompetta vs. sem vesicular seminalis spr spiracles vst vestige spr. atr spiracular atrium spr. tr spiracular trachea w. aor wall of aorta spt spermatocytes wng. tr wing trachea sr. blad serrated blades SIC sarcostyles or myofibrils y yolk sr. rig serrated ridges st stipes 2. abd. gng second abdominal ganglion stl style 6. abd. gng sixth abdominal ganglion stom. inv stomodaeal invagination 7 & 8. abd. sub. gng suboesophageal ganglion gng fused seventh and eighth abdominal sus . sc suspensory sclerite ganglion

10 abd. s tenth abdominal segment t tentorium 1. br protocerebrum tb tibia 2. br deu tocerebrum t. brg tentorial bridge 3. br tri tocerebrum teg. nv tegumental nerve 1. nv first lateral nerve ten. taenidium 2. nv second lateral nerve tes testis 1. s. lb. plp first segment of labial palpus tes. sh testicular sheath 2. s. lb. plp second segment of labial palpus tg-stn . m tergo-sternal muscle 1. s. lbl first segment of labellum th labial theca 2. s. lbl second segment of labellum thor. gng thoracic ganglion 1. s. max. plp first segment of maxillary palpus tn. 9 tendon of adductor muscle of 1-10 stn sternites

mandible 1-10 tg tergites tn . 10 tendon of abductor muscle of

mandible

16

The Sandfly

Phlebotomus papatasi

17

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part of head of cf; 13, spiculate scales from pharyngeal wall.

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Figs 20--26 Phlebotomus papatasi, ~. 20, dorsal view of right maxilla; 21, distal part of lacinia; 22, dorsal view of labium and part of labial gutter; 23--25, transverse sections of second and first segments of labella and middle

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29

e.rig

24, I ' I '

' I

\ \

25 \

\ sl.pmp ,

I

' 20 lb .nv

21 pdc-"

,

4 -- --5-----

2', 27,

' ' ant.t.p..._ '

' ' ' ....

11-- .. .....

'

18

I

ls.mx.plp

br 31 30

29

' \ sub.gng

' \ ' cm.sl.d sJ.gJ

33 27 I I I prt.gng

j.o .2 3

br

28

9 /

/ /

--

-- 32 ..... ~

'

' pst.t.p I

' I ' '20 ' I

'I 15 cd

Figs 27-29 Phlebotomus papatasi, 9. 27, longitudinal section of middle part of head and anterior part of thorax; 28, same of right half of head; 29, muscles of posterior part of cibarial pump.

24

tor',

31 I

sl.pmp scp,

'

26 I crp.card .nv- --

_ -rec.nv

vnt.c.trn

- - fat.b-- ... ,. t

3br" .1 32 33

rec.nv a or

lbr vnt.c.trn \ 31 .05

Figs 30-35 Phlebotomus papatasi, ~; transverse sections. 30, basal parts of labrum, mandibles, sialophore, laciniae and labium; 31, 32, anterior and middle parts of head; 33, 34, posterior parts of pharyngeal pump and

head; 35, oesophagus, corpus cardiacum, hypocerebral ganglion and aorta.

25

N 0\

lm ' ,,

\''

'

sial .. ~ v

24 /

/ /

' ' pmt

cl.nv ' '

\ cm.sl.d

f.gng

' '

' ' '

\ ' \ '

mx.nv ' md.nv

Fig. 36 Phlebotomus papatasi, 2; nervous system of right half of anterior part of head.

29 28

rec.nv

..,..30

ph.pmp

' 3br \ , .

\ nv.sl.d

c

cbpmp_

\ \

\ sl.pmp

\ \

I I I

\ sl.gl 1 ' ph.pmp thor .gng

' sub.gng

cm.sl.d

pstn I

' tro.2

cr

,

Mal.tu /

hin-g f

I

/ I OV I

6abd.gng

7+8abd .gng

Fig. 37 Phlebotomus papatasi,

mid-g

ven

ph.pmp ~,

I

I \

I I

I

I

,

I

,

30

- oe

cr

\ Mal.tu

Fig. 38 Phlebotomus papatasi, ~; alimentary canal.

28

44

' oe

long.m I

41

car aor ' w.aor I '

' \

d.cr 42

c1r'.m- ..... -45

.8

ven

reg.ce/ I

cir.m ' '

b.mem-

''long.m cir.m

I I

pr.sh prit.mem

" " /

43

46

47

\ prit.mem

Figs 39-47 Phlebotomus papatasi, s>; histology of crop and mid-gut. 39, transverse section of duct of crop; 40, longitudinal section of empty crop; 41, surface view of stretched wall of same showing thick circular muscles and thin longitudinal ones; 42, longitudinal section of oesophagus; 43, transverse section of thoracic part of mid-gut; 44, surface view of wall of same showing disposition of longitudinal and circular muscles; 45, longitudinal section of abdominal part of mid-gut; 46, 47, the same a few, and about 16 hours, after feeding.

29

48

50 l~ng.m

.05

-.... msn.inv

'fong.m

pr.sh

52

j --

I I

vii

I I

Mal.tu

49

51

/ ,pg.gr

53

b.mem"

Figs 48--53 Phlebotomus papatasi, S?; histology of hind-gut and of Malpighian tubes. 48, posterior part of mid-gut and anterior part of hind-gut; 49, connection of Malpighian tube with mid-gut and hind-gut and continuation of villarium into Malpighian tube; 50, 51 , transverse and longitudinal sections of ileum; 52, 53,

longitudinal and transverse sections of middle part of Malpighian tube.

30

Stg \ ht

/ I

6tg

/ /

6stn oen ,

I long.stn.m

7+8abd.gng

/ob.B.stn

54

op.sp.d

55

9stn

\ 7stn

/ /

.1

' ' ' lob.B.stn

prg

Figs 54, 55 Phlebotomus papatasi, S? . 54, longitudinal section of posterior end of abdomen; 55, transverse section of ninth abdominal segment.

31

a.nv

md.nv

mx.nv"

.1

'

57 crp.card .nv rec.nv

Figs 56, 57 Phlebotomus papatasi,

.. mtr.nur

mes.gng'

58

e>--------- b.mem cm.sl.d

.1

60 62

Figs 58-62 Phlebotomus papatasi, 2. 58, nervous system of thorax in lateral view; 59, section of prothoracic ganglion; 60, salivary glands; 61, 62, sections of right and left glands.

63 .02

nv./am, '

ns.cel

bid. sin

' ' rec.nv /

hyp.gng"

phpmp

66

64

d.cr"""'

crp.card \ \

" rec.nv"

mid-g ... 65

.05

crp.a/

mid-g

Figs 63--66 Phlebotomus papatasi, 9. 63, neurosecretory cells; 64, cells of corpus cardiacum; 65, diagram of dorsal view of stomatogastric nervous system and associated endocrine glands; 66, right half of same showing

histology of glands and their connection with brain and stomatogastric nervous system.

34

2stn 67 long.stn.m

,.

cv.mem

Figs 67-69 Phlebotomus papatasi,

I fat.b 70

72

I 1./ong.tg.m

.02

75

Figs 70-75 Phlebotomus papatasi, S?. 70, transverse section through dorsal half of third abdominal segment; 71, nephrocytes; 72, part of heart showing its internal structure and attachment to its wall of alary muscles; 73, blood cells, one of them attached to histolysed larval muscle; 74, 75, nephrocytes of unfed female and after

oviposition.

36

a

I I

I oc/r.sc

pdc

77

rec.nv--

ph pmp--

.1

cor

irpg.ce/ __

ret. pg.cel-b.mem,

prp-

80

lbr

76

11 v 78 79

,cg.ce/

--~----11>. .. , - . .

-

----~} ,%~r[)J

-ret.cel

.05

out.chia

Figs 76-80 Phlebotomus papatasi, S? . 76, transverse section of anterior part of head showing antenna! pulsatory organ which consists of vesicle and two muscular pulsatory membranes; 77, longitudinal section of first antenna! segment; 78, transverse section of tubular part of pulsatory membrane near vesicle; 79, pulsatory

organ of scutellum; 80, ommatidium of light-adapted compound eye.

37

" cb-lm.tr'

81

Figs 81-84 Phlebotomus papatasi, 2. 81, tracheal system of head in dorsal view; 82, dorsal view of origin of dorsal and ventral cephalic tracheal trunks from lateral thoracic trunks from near anterior thoracic spiracles;

83, 84, anterior and posterior thoracic spiracles.

.3

85

/.thor. trn

l.abd.trn /

/

86

.04

Figs 85, 86 Phlebotomus papatasi, S?. 85, tracheal system of right half of thorax and of abdomen; 86, abdominal spiracle.

39

, lig

.3

ov/...-''

...

'ac.g/

lob.B.stn op.sp.d ' 1 op.ac.g

Fig. 87 Phlebotomus papatasi, Cj?; reproductive organs, tracheal system not shown.

40

lig ' sp.gl

91

b.mem ~

...

. oog m

.05 92

nr.cel \ \ ovl

' \ \

\ fol ,,

90 89

Figs 88--93 Phlebotomus papatasi, ~; histology of internal reproductive organs. 88, 89, longitudinal sections of spermatheca and ovary; 90, ovariole with one developing follicle containing seven nurse cells and one oocyte, and in germarium five dividing oogonia; 91, end of calyx and beginning of lateral oviduct; 92, section of wall of

accessory gland; 93, musculature of ovarian sheath.

41

~

7stn

94

8stn

95

" /

8tg ,

/ob:8.stn

.1

\Otg

op.sp .d

9tg ...

cer

Figs 94--96 Phlebotomus papatasi,

I fl ,

35' ....

.....

100

97

vas.def /

.1

101

98

gon

/ I

99

ej.d ~~

,. ,.

,.

s.pm,

tu.sh-

gen.{/ "

.05

rd

.03

102

,

/

35

Figs 97-102 Phlebotomus papatasi, cf' . 97, reproductive organs; 98, section of pompetta showing inner sac and opening of ejaculatory duct; 99, horizontal section of same showing compressor muscles of sac and protractors

of pompetta and genital filaments; 100-102, genital filaments.

spm

api.cel ' '

mi

103

spd I ej.d,

' ' '

104

.05

- pr.sh

b.mem

m,

pr.sh, , epi"

b.mem

105

Figs 103-105 Phlebotomus papatasi, d; histology of testis and vesicula seminalis. 103, longitudinal section of testis; 104, longitudinal section of vesicula seminalis and its glands, each extending from vas deferens, and

latter with narrow canal full of spermatozoa; 105, transverse section of ejaculatory duct.

44

7tg /

.1

c/sp

-1 Ostn

9tg~

108

.05 109

107

Figs 106-110 Phlebotomus papatasi, cf; genitalia. 106, end of abdomen after torsion; 107, ventral view of ninth abdominal segment; 108, tenth abdominal segment; 109, longitudinal section of tactile sensillum of style; 110,

penniculus.

45

I I

8tg

aed

39 '

.05

41 stl

.2

' 46 aed--

113

50

Figs 111-113 Phlebotomus papatasi, d'; musculature of genitalia . 111, inner view of right half of ninth and tenth abdominal segments and of right coxite; 112, transverse section of tenth abdominal segment; 113, same

through basal parts of coxites.

46

The Mosquito

Aedes aegypti

47

--- th

_./bl 114

_ mid-cr.sut

mx.plp

I I

.3

c/ge .sut ,

pge --

clge.sut '

115

.3

lc 116

cv.sc oe ~ _

117

Figs 114-117 Aedes aegypti, S?. 114, anterior view of head and proboscis; 115, lateral view of anterior part of head showing connection of anterior arm of tentorium with ciypeus, position of torma, and connect!on of cibarial pump by ciypeal apodeme with clypeus; 116, lateral view of head, left compound eye removed to show position of pharyngeal pump; 117, middle part of ventral wall of head showing internal position of

stipites, occipital foramen and fused postgenae of which anterior parts form postgenal bridge.

33

118

--- -~?J-- op I { '

.1

---- sl.pmp

''cm.sl.d

sial---

.5

ph.pmp---

122

.05

- lm

l.w.lm--

' dr.w./m

lm

.1

cl

cl.ap- __ _

119

121 buc

Figs 118-122 Aedes aegypti, ~. 118, posterior and anterior parts of sialophore; 119, lateral view of clypeus, cibarial pump, pharyngeal pump, sialophore and labrum; 120, ventral view of anterior part of labrum; 121, dorsal view of cibarial pump and posterior part of labrum, showing attachment of clypeal apodemes to lateral margins of cibarial pump, and sense organs of anterior wall of latter; 122, scale-like processes of posterior

parts of walls of pharyngeal pump.

lbl.prc

125

dr.w.lm 21 ,,,

123

.05

. 1 126

cl .05

.1

7

I s.mx.plp

fb.nv

.nv 124 ---lb.tr

128 127

Figs 123-128 Aedes aegypti, 2. 123, anterior part of prementum and two-segmented Iabella, left Ia bellum showing ventral wall; 124, internal structure of same, showing three pairs of muscles and their attachment to segments of Iabella, labial nerves and tracheae; 125, transverse section of first segment of Iabella; 126, S3me of anterior part of proboscis; 127, same of middle part of proboscis showing position of syntrophium (combina-

tion of mouth parts) in labial gutter; 128, same of anterior part of clypeus and base of mouth parts. '

sub.gng .... ph pmp,.,

sns. org f \ /

~

- ... ge ----16

130

.1 a or

132

Figs 129-133 Aedes aegypti, 9. 129, transverse section of anterior part of head; 130, same of posteriL1r end of cibarial pump and of anterior part of bucca, showing origin of protractor muscles of cibarial pump on tentorium and their insertion together with retractor muscles of the same on cornua; also showing origin of salivary pump muscles on loral arms; 131, sense organ (?) of posterior part of antenna! pulsatory organ; 132, section of posterior part of pharyngeal pump and its circular sphincter muscle; 133, section of posterior part of head showing position of pharyngeal pump and insertion on its plates of postcerebral dorsal dilators and

posterior lateral dilators.

31 f

cl .nv f.gng I '\, 29 ... _ ' m.nv'

fd.cn \

' em~ sl.d cv.con

th /b.nv sl.pmp I

/b.g .3

Fig. 134 Aedes aegypti, S?; right half of head showing sucking apparatus comprising food canal of labrum, cibarial and pharyngeal pumps, and their muscles .

52

bs.snl

.05

137

2

' 27 .05 st- ,

12 .. ,

136

135

Figs 135-137 Aedes aegypti, S?. 135, distal end of mandible in dorso-lateral view; 136, musculature of mandible and maxilla; small drawing shows articulation of mandible by mandibulo-genallink; 137, distal end of lacinia.

53

scut '

long .dr .m I scut/

. 1

fat .b ht

Mal.tu

mid-g I

I

Fig. 138 Aedes aegypti,

' sf.pmp

ten

141 139

sl.d ,...

b.mem

.05

Figs 139-143 Aedes aegypti, 2. 139, salivary gland immediately after emergence of mosquito from pupa; 140, salivary glands of two-day-old mosquito, and longitudinal and transverse sections of common salivary canal; 141, section of middle gland showing intracellular duct and its supporting disc; 142, 143, longitudinal and

transverse sections of lateral gland.

int

' '

.2

I I '

c:r ~-

.05 .1

. I

I ;

I hin.g ~ _

...........

\

' \ \

\ \

\ \ \

----... -- ... ___ ...._-

144

\ \

\ \

\

146

mid-g ' I

I

-------

n.epi .eel

.05

an

Figs 144-147 Aedes aegypti, S!. 144, alimentary canal; 145, diagram of dorsal diverticulum showing thick circular and thin longitudinal muscles and very thin longitudinal folds in wall; 146, wall of ventral diverticu-

lum or crop; 147, section of walls of oesophagus and crop.

56

oe

pr.sh I

I I

c1r.m

148

long.m'

149

152

,

cr

...

'

'

' ' ~il

' epi

reg.cel

d.cr' .1

150

____ .vii

et>i

153

,

' ' ,

pr.sh cir m

151

Figs 148-153 Aedes aegypti, ~; histology of mid-gut. 148, median longitudinal section of anterior part; 1491 longitudinal section of abdominal part, or ventriculus, of unfed mosquito; 150, section of narrow thoracic part fixed during feeding; 151, same of posterior part of ventriculus; 152, middle part of ventriculus of fully fed female, showing strongly flattened epithelium and villarium; 153, same 24 hours after feeding, showing

peritrophic membrane.

ven

epi '

154

b.mem _.,

158

159

~ long.m

-- -~pyl.sph

b.mem j

Mal.tu ---

.05 ___ /ong.m

- -------. hin.g-------

;'

, epi

155

.05

epi .eel

long.m

pr.sh ,

,

/ /

long.'m

cir.m

157

rad.str --

.05

int----

160

Figs 154-161 Aedes aegypti, ; histology of hind-gut and Malpighian tubes . 154, posterior part of mid-gut and anterior part of ileum; 155, same from a different angle, showing origin of Malpighian tube from mid-gut and the same structure of their epithelial cells; 156, 157, longitudinal and transverse sections of middle part of Malpighian tube of unfed female; 158, longitudinal section of colon and its short narrow posterior part, in both of which epithelial cells lack spicules; 159, longitudinal section of thick part of colon (see Fig. 171); 150, 161,

transverse and longitudinal sections of posterior part of rectum.

.05

aus

'tg -stn .m

~I. thor .trn 163

cir.m '

b.mem'

.5

164

.5 - -tro.l

162

Figs 162-164 Aedes aegypti, 2. 162, slightly oblique transverse section of anterior part of thorax, showing position of salivary glands, oesophagus and corpora alata; 163, corpora alata on sides of aorta; 164, transverse

section of oesophagus .

59

0'-0

suH.gng

. crp.card crp.al

.r

oe

/

.2

Fig. 165 Aedes aegypti, 9; stomatogastric nervou s system and endocrine organs.

a or

13/

.3

I 6stn

166

.03 dr.dphr

169

'

', vntdphr

vnt.nv.ct d

\ \

\

.05

al.m

.05

167

168

Figs 166-170 Aedes aegypti, S? . 166, transverse section of sixth abdominal segment showing position of heart and attachment of its alary muscles to intersegmental membrane, etc.; 167, part of heart with its cavity exposed to show ostia and ostial valve, dorsal diaphragm and nephrocyte among muscles and elastic tissue; 168, section of ventral diaphragm; 169, four haemocytes from surface of mid-gut and three from ovaries; 170,

oenocytes in abdominal fat body.

~ N

I

\ \

\

'

7 +8 abd .gng ' 7stn fat.b

.2

8tg .

p'gen .p

J?;f oen 8stn

Fig . 171 Aedes aegypti, 2; longitudinal section of posterior end of abdomen.

pr.sh"

173

., .,

/

.3

174

cm.ovd--

ac.gl --pgen. P

'

tr

'.l.ovd

8 abd.gng

.05

rect 175

.3 I Ostn. __

gen.ch/

177

Figs 172-177 Aedes aegypti. 2. 172, reproductive organs of eight-hour-old female; 173, transverse section of accessory or collaterial gland; 174, longitudinal section of duct of same; 175, longitudinal section of distal end

of bursa copulatrix; 176, posterior end of abdomen; 177, ventral view of same.

.05

'' ::-- -fl '\

. 1

cho ns.cef. __

~-ov.sh

ooc --

179 fol.epi ----

y ..

180

Figs 178-180 Aedes aegypti, 2. 178, surface view of ovarian sheath, showing its branching and anastomozing muscles; 179, part of ovary of just-emerged female; 180, ovariole with germarium and three follicles .

64

.05 I

I I

I I

long.m I

I

181

cm.ovd

182

__ .sp.gl

.03

d.ac.gl

gon 184

.05

Figs 181-185 Aedes aegypti, S?. 181, section of spermatheca and its duct; 182, spermathecal gland; 183, transverse section of spermathecal ducts, surrounded by strong longitudinal muscles; 184, longitudinal median section of posterior end of abdomen, showing migration of spermatozoa from bursa copulatrix through spermathecal ducts into spermathecae after copulation; 185, transverse section of common oviduct,

showing sphincter muscles and two dilator muscles arising on eighth sternite.

186

188

aed .. ---

\

pr.sh

int

',

'

.05

' ',m

m

vs.s

spm --

-b'.mem

189

ac.gl

.1

190

187

\ fat.b

Figs 186--190 Aedes aegypti, d. 186, reproductive organs showing two testes, one with fat body removed to show division into apartments; 187, transverse section of accessory glands and vesicula seminalis, its lumen filled with spermatozoa; 188, longitudinal section of wall of vesicula seminalis; 189, same of accessory glands;

190: same of vesicula seminalis and accessory gland, showing opening of latter.

tes .sh

fat .b-

.1

m t----

\. fl ----- -, ,, grm .eel

spg '

spt

spm

spd

Fig. 191 Aedes aegypti, d; longitudinal section of testis showing its division into apartments or cysts.

192

~I sp stl '

.3

194

, ej.d

.05

Figs 192-195 Aedes aegypti, cf. 192, lateral view of genitalia of Aedes sp., left coxite removed; 193, ventral view of same of A. aegypti; 194, parts of tenth abdominal segment; 195, right half of aedeagus showing its

connection with membrane and ejaculatory duct.

68

The Blackfly

Simulium

69

. 1

0

197

Figs 196, 197 Simulium (Wi/he/mia) sp ., 2. 196, anterior view of head; 197, tentorium of same.

70

198

.2

.05

.15

202

Figs 198-204 Simulium (Wilhelmia) sp ., S?; mouth parts. 198, labrum; 199, left mandible; 200, right maxilla; 201, lateral view of anterior end of labrum showing position of scraper; 202, right half of labium showing labial

muscles; 203, ventral view of labium; 204, dorsal view of sialophore, pharynx and oesophageal pump.

205 . 1 206

.1

209

st. pmp .05

.05

.1

Figs 20~211 Simulium (Wilhelmia) sp., 2. 205, transverse section through first segment of labella; 206, same through middle part of proboscis; 207, same through oesophageal pump; 208, same through anterior part of salivary pump; 209, same through anterior part of pharynx and posterior part of salivary pump; 210, section of

salivary glands; 211, same of reservoir of .salivary glands.

72

.2

Figs 212-213 Simulium (Wilhelmia) sp ., 2; sections of head showing internal structure and musculature. 212, longitudinal, through middle part; 213, oblique transverse.

73

~

1.0

Fig. 214 Simulium (Wilhelmia) sp ., 2; right half of body showing natural position of internal organs .

Fig. 215 Simulium (Wi/helmia) sp., 2; alimentary system.

75

card . gng

216 217

crp. a/

218

Figs 216--219 Simulium (Wilhelmia) sp.,

. 1 220

222

.05

.1 221

.05

223 224

Figs 220-224 Simulium (Wilhelmia) sp., S?; histology of alimentary system. 220, longitudinal section of anterior part of mid-gut; 221 , 222, transverse and longitudinal sections of wall of mid-gut; 223, 224, longitudinal and

transverse sections of hind-gut (ileum).

77

.1

.1 226

225

.1

9tg

Figs 225--228 Simulium (Wilhelmia) sp.,

230

.03

.05

232

.05

Figs 229-234 Simulium (Wilhelmia) sp., S?; reproductive organs. 229, general structure; 230, transverse section of duct of accessory gland; 231, section of spermathecal duct; 232, section of accessory gland; 233, longitudinal section of spermatheca; 234, ovariole showing its attachment to muscular sheath, and different stages of

development of follicles.

235

238

: .: .. ~stl

b.mem !.~--cir. m

.05

236

Figs 235-239 Simulium (Wilhelmia) sp., d; reproductive organs. 235, general structure; 236, longitudinal section of testis; 237, transverse section of vasa differentia and vesicula seminalis; 238, section of wall of vesicula seminalis; 239, median longitudinal section of posterior end of abdomen, showing genital opening,

structure of ductus ejaculatorius, etc.

The Deerfly

Chrysops caecutiens

81

00 N

pv

1.0 I ~ l

oc

lm

' 240 241 fbi

242

Figs 240-242 Chrysops caecutiens,

--{\

v

248

Figs 243--248 Chrysops caecutiens, 5?; mouth parts . 243, labrum and its distal end; 244, left mandible; 245, right maxilla in ventral view; 246, sialophore and cibarial and pharyngeal pumps; 247, labium in dorsal view; 248,

collecting channel and pseudotrachaea of left labellum.

249

250

I .05

251

Figs 249-253 Chrysops caecutiens , S?; internal structure of labium. 249, labial glands; 250, transverse sections of middle part of Iabella and pseudotracheae; 251, transverse section of middle part of labium showing position of other mouth parts; 252, sensilla coeloconica of inner wall of Iabella; 253, sensilla trichodea of outer wall of

Iabella .

. 1

.05

. 1

Figs 254--259 Chrysops caecutiens, S?; transverse sections. 254, most anterior part of head; 255, posterior part of pharyngeal pump; 256, posterior part of head showing oesophageal sphincter (circular muscles) etc.; 257,

oesophagus; 258, duct of crop; 259, crop.

cb.

lm

sial

ph. pmp

mid-g

an

Fig. 260 Chrysops caecutiens, Cj2; alimentary canal and salivary glands.

86

' '

263

262

.05

264

Figs 261-264 Chrysops caecutiens,

.01

265

Fig. 265 Chrysops caecutiens, 2; longitudinal sections of anterior part of mid-gut and stomodaeal invagination.

thor. gn9 '\.0

_vnt. dphr

.5

.05

Figs 267-270 Chrysops caecutiens, ~; histology of narrow thoracic, and abdominal, parts of mid-gut. 267, transverse section of middle of abdomen showing Malpighian tubes, heart, ventral diaphragm, ovary and parietal and visceral fat-body; 268, same of narrow thoracic part of mid-gut; 269, 270, longitudinal sections of

wall of ventriculus showing discharge of secretory globules by epithelial cells.

90

.05

272

273

ven

Figs 271-274 Chrysops caecutiens,

Fig. 275 Chrysops caecutiens, 2; heart and respiratory system. 92

nephr

Figs 276, 277 Chrysops caecutiens, c;?; histology of heart and ventral diaphragm. 276, transverse section of heart; 277, abdominal part of ventral diaphragm.

f

cbl. pmp

cl

1.0

lm th

279

278

Figs 278, 279 Chrysops caecutiens. ~; 278, internal structure of head; 279, nervous system.

94

280

Figs 280--282 Chrysops caecutiens, 2; histology of anterior part of ventriculus and corpus allatum, transverse sections. 280, anterior part of thorax; 281, caecum of anterior part of ventriculus; 282, corpus allatum, corpus

cardiacum and aorta.

95

284

.1

Figs 283-286 Chrysops caecutiens, 9. 283, histolysis of larval muscles by haemocytes; 284, oenocytes; 285, 286, parietal and visceral fat-body.

\0 '-1

Fig. 287 Chrysops caecutiens, S?; longitudinal section of posterior part of abdomen.

fig 1.0

Fig. 288 Chrysops caecutiens, 2; reproductive organs.

.05

289

290

292

.1

cir. m

293

Figs 289-295 Chrysops caecutiens, 2; histology of reproductive organs. 289, transverse section of accessory glands; 290, same of posterior part of abdomen; 291, ovariole; 292, transverse section of wall of spermatheca showing unicellular glands and their intracellular ducts; 293, 294, same of glandular and posterior parts of

spermathecal duct; 295, same of accessory gland duct .

The Stablefly

Stomoxys calcitrans

101

pt. sut-

f-

Fig. 296 Sto111oxys calcitrans, 2; anterior view of head . 102

Fig . 297 Stomoxys calcitrans, 2; internal stru cture of head and proboscis.

298

ern

.1 pre.w.lm

304

Figs 298-304 Stomoxys calcitrans, S?; mouth parts. 298, distal end of sialophore; 299, distal end of labrum; 300, proboscis; 301, 302, transverse sections through anterior and middle parts of proboscis; 303, ventral view of

Iabella, left half showing internal structure; 304, transverse section through most posterior part of same.

mal. t

Fig. 305 Stomoxys ca/citrans, 2; dorsal view of dissected fly showing position of internal organs.

105

Fig. 306 Stomoxys calcitrans, S?; alimentary canal.

106

OL, int

307 308 309

.1

311

. 1

312

Figs 307-313 Stomoxys calcitrans, 9; structure of salivary glands, oesophagus and crop. 307, transverse section through duct of salivary glands; 308, 309, transverse sections through anterior and posterior parts of salivary gland; 310, 311, transverse sections through most anterior and cerebral parts of oesophagus; 312, histological

structure of wall of crop; 313, transverse section through crop.

107

].02 .02 314 16

315 .1

.05

321

.03 317 .1

vii .03

cir. m pr. sh

320 b.mem

319 reg. eel 322

Figs 314-322 Stomoxys calcitrans,

.2

323

. 1

325

.05

int

long. m 326 327

Figs 323-327 Stomoxys calcitrans, ~; structure of anterior and posterior intestine. 323, rectal valve; 324, rectal papilla; 325, 326, transverse sections of wall of ileum and posterior part of rectum; 327, longitudinal section of

posterior part of abdomen. 109

prit. m

if

328

.03 .1

vii 331 ' ' ' \

'

.02

330 332

Figs 328-332 Stomoxys calcitrans,

mal. tu .1

333

.03

336

long. m 334

.03

.02

337

Figs 333-337 Stomoxys calcitrans, S?; structure of heart. 333, transverse section through posterior part of heart; 334, part of heart showing ostial valves and alary muscles with nephrocytes or pericardia! cells; 335,

pericardia! cells; 336, haemocytes; 337, fat-body with two oenocytes.

111

Fig. 338 Sto111oxys cnlcitrnns,

339

.03

340

Figs 339, 340 Stomoxys calcitrans,

1.0

epi

sph. m 342

m 346

344 343

Figs 341-346 Stomoxys calcitrans, C(; reproductive organs . 341, general structure; 342, transverse section of accessory glands; 343, 344, same of anterior and posterior parts of its duct; 345, 346, transverse sections of

spermathecae and their ducts.

114

348

.1

.5

349 rem. fol. epi

.03

350

Figs 347-350 Stomoxys calcitrans, 2; histology of ovary. 347, two ovarioles, one containing remainder of disintegrated epithelium of first follicle after oocyte has left follicular tube; 348, transverse section of lateral oviduct; 349, surface view of ovarian sheath showing anastomosing muscles; 350, posterior end of egg

showing oosome.

.02 tes. sh

355

Figs 351-356 Stomoxys calcitrans, d'; reproductive organs. 351, general structure; 352, 353, transverse sections of vesicula seminalis and vas deferens; 354, same of posterior part of ejaculatory duct; 355, longitudinal

section of testis; 356, wall of same.

116

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QX 505 J62A 1987 JoBLINGJ BoRIS~ 1893-1986 ANATOMICAL DRAWINGS OF

BITING FLIES

. MIG 3 1 ua I ...)

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