Diptera: Chloropidae) — an Inquiline in Galls Formed by Lipara Flies on Common Reed

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Morphology of preimaginal stages of Cryptonevra flavitarsis (Diptera: Chloropidae) — an inquiline in galls formed by Lipara flies on common reed

Maria Grochowska

Grochowska, M. 2008: Morphology ofpreimaginal stages of Cryptonevraflavi— tarsis (Diptera: Chloropidae) — an inquiline in galls formed by Lipara flies on

common reed. — Entomol. Fennica 19: 18—24.

The paper presents a complete description ofthe morphology ofthe preimaginal stages of Cryptonevraflavitarsis (Meigen, 1830), an inquiline in galls ofLipara flies on common reed (Phragmites australis). All stages are described for the first time. In descriptions ofthe larval stages, particular attention is given to the diag- nostic characters: facial mask, cephaloskeleton, anterior and posterior spiracles, and locomotory structures. The paper forms a basis for future descriptions ofthe yet unknown life cycle of this fly.

M. Grochowska, Department ofZoology, Maria Curie—Skiodowska University, Akademicka 19, 20—033 Lablin, Poland; E—mail.‘ [email protected] Received 4 August 2006, accepted 20 April 2007

1. Introduction (Giraud 1863, Duda 1933, Hennig 1943, Nart— shuk 1962). The overwintering stage is the pupa. The genus Cryptonevra Lioy, 1864, contains 12 Adults can be seen flying in reed beds from late species (Nartshuk 1984). Four of them: C. flavi— April to August (Blair 1932, 1944, Erdos 1957, tarsis (Meigen, 1830), C. diadema (Meigen, Wendt 1968, Nartshuk 1969, Waitzbauer 1969, 1830), C. consimilis Collin, 1932 and C. nigri— Waitzbauer et al. 1972, Pokomy & Skuhravy tarSiS (Duda, 1933), are inquilines in galls formed 198 1). C.flavitarsis is distributed from Great Bri- by Lipara and Platycephala flies on Phragmites tain in the west to the Angara subregion in West- australis (Cav.) Trin. (Kramer 1917, Collin 1932, ern Siberia and Mongolia in the east. The north- Duda 1933, Hennig 1935, 1943, Blair 1932, ern limit of its range reaches as far as Finland, 1944, Erdos 1957, Wendt 1968, Nartshuk 1969, while the southern limit extends from northern 1972, 1996, Waitzbauer 1969, Waitzbauer et al. Africa (Morocco) to Armenia, Azerbaijan, to 1972, Beschovski 1984, De Bruyn 1985, Ferrar China (Beschovski 1984, Nartshuk 1984). 1987). C. flavitarsis is the most common ofthese The developmental stages of C. flavitarsis four (Pokomy & Skuhravy 1981). Larvae of C. comprise the egg, three larval stages, the pupa and flavitarsis develop among leaves in the apical part the imago. Of these, the imago has been well de- of a gall or in the larval chamber next to larvae of scribed (Meigen 1830). Habitus drawings of the Lipara and Platycephala. C. flavitarsis is also puparium are also available (Waitzbauer 1969). found among normal and decaying leaves in the Data on the other stages are lacking. apical parts of reed stems not inhabited by other The aim of the present study was to describe insect species and inside stems ofTriticum repens the morphology of all preimaginal stages of C. ENTOMOL. FENNICA Vol. 19 0 Preimaginal stages ofC. flavitarsis 19 flavitarsis, thus paving the way to describing the species’ life-cycle and its impact on common reed stems.

2. Materials and methods

The materials included all stages of C. flavitarsis obtained from the apical part of common reed stems. The stems were collected in reed beds in and around Lublin (central-eastern Poland) in the mm years 2002—05. Fresh stems were randomly col- 0.25 lected in May and June, and stems bearing visible galls in the apical part without inflorescences mm were collected outside of this period. The stems 0.25 were subsequently sectioned in the laboratory un- der a stereoscopic microscope. The sectioning of Fig. 1. C. flavitarsis, egg. — a. Dorsal view. — b. Lateral galls yielded all larval stages and pupae. Eggs view. — c. Anterior section. — d. Posterior section. were obtained from the upper surface of leaf blades and from sectioned abdomens of females. First-instar larvae were reared from eggs col- forming squares denser and darker at both ends, lected in the field or laid by females in the labora- more regular and darker at anterior than posterior tory. Imagines were obtained by rearing larvae pole. Chorionic surface covered with micro- and pupae in Petri dishes lined with filter paper. scopic closely-spaced processes making it look Overall, the study material comprised: 30 like a nap. eggs, 15 first-instar larvae, 10 second-instar lar- First—instar larva (Fig. 2). 1.05—1.30 mm in vae, 20 third-instar larvae, 15 pupae, and 20 length, 0.18—0.22 mm at widest diameter. Body imagines. white, slender, oval in cross-section (Fig. 2a). Fa- The specimens chosen for detailed study were cial mask (Fig. 2b) with a pair oftwo-segment an- macerated in 10% KOH for 24 hours at room tem- tenna, maxillary and frontal palpi. Basal segment perature, washed in distilled water, cleared in of antenna ring-like, slightly smaller in diameter chloral hydrate and chloral phenol, and preserved than apical segment. Apical segment dome- in glycerol. Drawings were based on the micro- shaped. Maxillary palpi with numerous sensilla. scopic preparations. Frontal palpi with two sensilla over mouthhooks. Imagines were determined according to the Oral ridges straight, not branching, arranged key by Nartshuk et al. (1970) and comparedto the symmetrically on both sides of mouth opening first description of the species (Meigen 1830). (Fig. 2b). Cephaloskeleton (Fig. 2c) of light am- The descriptions oflarvae follow the terminology ber colour, 0.25 mm in length, with 0.06 mm-long of Courtney et al. (2000). Parts ofthe larval body mouthhooks. Mouthhooks long, narrow (Fig. not defined in Courtney et al. (2000) are named 2d). Apical tooth hook-like bending downwards, according to Nye (1958) and Nartshuk (1987). no oral teeth. Basal tooth poorly distinguishable. Base of mouthhook long and narrow. Additional sclerite shaped like wide plate adjoining external 3. Description of preimaginal stages surface of mouthhook at right angle (Fig. 2c, d). Intermediate sclerite closely connected with Egg (Fig. 1). 0.90—1.00 m long, 0.23—0.25 mm basal sclerite. Basal sclerite divided into ventral wide, mat, white, oblong, slightly rounded at both and dorsal cornua. Ventral cornu longer and ends (Fig. 1 a). Length to width ratio 4: 1. Dorsally wider than dorsal cornu. Both cornua poorly flat, ventrally arch-like (Fig. 1b). Surface covered sclerotised, mostly membraneous. Dorsal cornu with longitudinal and transverse ribs (Fig. 1 c, d) narrowing posteriorly, continuous with dorsal 20 Grochowska ' ENTOMOL. FENNICA Vol. 19

mm dorsal bridge dorsa cornu 0.50

Fig. 2. C. flavitarsis, larva. additional sclerit first-stage apical tooth — a. Ventral view.

— b. Facial mask.

— c. Cephaloskeleton.

— d. Mouthhook ventral

view. — e. Posterior spiracle. — f. Posterior segment ventral view. kw — g. Spinules on ventral intersegmental regions of seventh and eighth h g 0mm abdominal segments. 0.025 mm — h. Anal spinules.

bridge anteriorly. Hypopharynx with prominent mm in length, including 0.08 mm long mouth- longitudinal ventral cibarial ridges extending into hooks. (Fig. 3c). 4—5 oral teeth behind apical lumen of cibarial pump. Posterior spiracles on tooth (Fig. 3d, e). Basal tooth wide and short, con- stigmatophores, with two spiracular openings tinuous with long and narrow base ofmouthhook. and two pairs of branching interstigmal hairs Intermediate sclerite long and narrow, connected (Fig. 2e, f). Spiracular trunk long (0.04—0.05 mm) with basal sclerite (Fig. 3c). Basal sclerite distally and thin (0.0025 mm). Distance between spira- divided into ventral and dorsal comua. Ventral cles 0.03 mm. Spinules present at segment bor- cornu longer and wider than dorsal comu. Dorsal ders ventrally and around anal plate (Fig. 2g, h). cornu narrowing posteriorly, continuous with Second—instar larva (Fig. 3). 1.68—3.04 m dorsal bridge anteriorly. Rod-shaped parastomal long, 0.28—0.55 mm at widest diameter. Body bar projecting anteriorly from basal sclerite. white, long, slender, oval in cross-section, poste- Hypopharynx with prominent longitudinal ven- rior end slightly rounded (Fig. 3a). Facial mask tral cibarial ridges extending into lumen of with pair of antennae, maxillary and frontal palpi cibarial pump. Anterior spiracle with 5—7 lobes (Fig. 3b). Basal segment ofantenna ring-like, api- (the number variable between individuals and be- cal segment dome-shaped, narrower at base, lo- tween left and right spiracles) (Fig. 3f). Posterior cated inside basal segment. Maxillary palpi with spiracle on stigmatophores with three oval numerous sensilla. Frontal palpi with two sensilla spiracular openings (Fig. 3g, h). Spiracular plate over mouthhooks. Anterior and prefrontal pa- quadrilateral with widely rounded comers. Ecdy- pillae present. Numerous oral ridges on both sial scar well visible. Spiracular hair short, sides of mouth opening and over mouthhooks. branching (Fig. 3g). Spiracular trunk 0.06 mm Cephaloskeleton partly sclerotised, 0.35—0.36 long, 0.008 mm wide. Distance betweenposterior ENTOMOL. FENNICA Vol. 19 - Preimaginal stages ofC. flavitarsis 21

anterior papilla prefrontal papilla

, antnna

palpus

0.025 mm Fig. 3. C. flavitarsis se- b cond-stage larva. dorsal bridge dorsa cornu — a. Lateral view.

— b. Facial mask.

— c. Cephaloskeleton. — cl. Mouthhook, lateral view. — e. Mouthhook, ventro-lateral view.

— f. Anterior spiracle. — g. Posterior spiracle. — h. Posterior segment, ventral view. — i. Spinu- les on ventral interseg- 042% pg mental regions of seventh and eighth abdominal segments.

— k. on ventral Spinules a... amt/oi, 0.10 mm <. v k , w, a intersegmental regions of v:)(\ n90 “ > of second and third ab- A A «v ”:29/:4?an 3‘”: ”n o\ a " “Mafia/'6 dominal segments. ,_// .L — |. Anal spinules.

spiracles 0.06 mm. Small spinules present at seg- mouthhook. Intermediate sclerite elongated, ment borders ventrally and around anal opening trianuglar, connected with basal sclerite. Basal (Fig. 3 i, k, l). sclerite distally divided into dorsal and ventral Third-instar larva (Fig. 4). 40075.12 mm comua. Ventral comu longer and wider than dor- long, 06441.12 mm at widest diameter. Body sal comu. Dorsal comu narrowing posteriorly, milky white, long, slender, oval in cross-section, continuous with dorsal bridge anteriorly. Rod- posterior end slightly rounded (Fig. 4a). Facial shaped parastomal bar projecting anteriorly from mask with pair of two-segment antennae and basal sclerite. Hypopharynx with prominent lon- maxillary palpi (Fig. 4b). Basal segment of an- gitudinal ventral cibarial ridges extending into lu- tenna ring-like, apical segment dome-shaped, men ofcibarial pump. Anterior spiracle with 449 narrowed at the base, located inside basal seg- lobes (the number variable between individuals ment. Maxillary palpus with numerous sensilla and between left and right spiracles) (Fig. 4e). (Fig. 4b). Pair of frontal palpi over mouthhooks, Posterior spiracle on stigmatophores with three each with 2 large sensilla. Anterior and prefrontal oval spiracular openings (Fig. 4f). Ecdysial scar papilla present. Numerous oral ridges on both well visible. Spiracular plate quadrilateral with sides of mouth opening and over mouthhooks. widely rounded comers. Spiracular hair branch- Cephaloskeleton partly sclerotised, 0.614068 ing (Fig. 4f). Spiracular trunk 0.08 mm long and mm in length, including 00740.08 mm long 0.05 mm wide. Distance between spiracles 0.05 mouthhooks (Fig. 4c). Numerous, very small oral mm (Fig. 4g). Large, plate-like spinules in teeth behind apical tooth along lateral surfaces of closely-spaced rows on lateral surfaces along an- mouthhook. Oral teeth reaching as far as basal terior border of first thoracic segment (Fig. 4h), tooth (Fig. 4d). Basal tooth massive, in middle of much smaller and less numerous spinules on ven- mouthhook, continuous with broad base of tral surface of same segment (Fig. 4i). Small spi- 22 Grochowska - ENTOMOL. FENNICA Vol. 19

. anterior .. . papilla antenna prefrontalpipilla A frontal maxillary

, palpus _ palpus

Fig. 4. C. flavitarsis,

E larva. m) ‘ third-stage oral i -\ ridges — lg a. Lateral view. b c: dorsal cornu

E — E b. Facial mask. G

— "2 ventral cornu c. o intermediate sclerite Cephaloskeleton.

— a d. Mouthhook, ventro—

lateral view. — e. Ante- ventral cibarial ridges — basal sclerite rior f. Poste- basal tooth spiracle. tOOth ror — Poste- apica oral tooth spiracle. g. rior segment, ventral view. — h. Spinules on Ia- teral anterior part of first thoracic segment. — i. Spinules on ventral anterior part of first thoracic segment. — k. Spi- nules on ventral intersegmental regions of second and third abdominal segments. — |. Spinules on ventral intersegmental regions of seventh and eighth abdominal segments. — m. Anal spinules.

nules at segment borders ventrally and around which are the Chloropidae with 26 species anal opening (Fig. 4k, 1, m). (Nartshuk 1996). Accordingly, common reed has Puparium (Fig.5). 2.764.44 mm long, 0.667 been considered a model plant for studies of 1.14 mm at widest diameter, oblong, straw-col- phytophage-plant relationships (Skuhravy 1981, oured to brown. Narrowed anteriorly, flattened Nartshuk 1996). Such relationships are important dorsoventrally, posterior end broad, slightly for the functioning ofmany ecosystems. Gaining rounded (Fig. 5a). Anterior spiracles present on a comprehensive understanding of the phyto- both sides of anterior border of puparium (Fig. phageiplant relationship is not possible without 5b, a). Posterior spiracles strongly sclerotised, on understanding the morphology and biology ofthe stigmatophores (Fig. 5c, d). Numerous folds of phytophages. cuticle in anterior part dorsally and all along ven- The present paper is a rare detailed account of tral surface (Fig. 5a, d). the morphology of one of many species inhabit- ing stems of common reed. Most available data pertain to with the morphology of Lipara flies 4. Discussion (Waitzbauer 1969, Chvala er a]. 1974, Grochow- ska 2006aa3, 2007), with few papers describing Common reed is among the plants supporting the the genus Cryptonevra (Hennig 1943, Waitz- richest phytophage faunas in the Palaearctic bauer 1969). Known descriptions include the (Emelj anov 1967), with about 200 consumer spe- third-instar larva and pupa of C. nigritarsis (Hen- cies belonging to seven orders of Insecta and one nig 1943, Waitzbauer 1969) and a drawing ofthe family of Acarina. The order of Diptera is repre- pupa of C. flavitarsis (Waitzbauer 1969). The sented by eight families, the most numerous of third-instar larva of C. flavitarsis differs from the ENTOMOL. FENNICA Vol. 19 ° Preimaginal stages ofC. flavitarsis 23

0.025 mm

Fig. 5. C. flavitarsis, puparium. — a. Dorsal view.

— b. Anterior spiracle. — c. Posterior spiracle. — d. Posterior segment, ventral view. larvae of C. nigritarsis described by Hennig phages. They are found in galls of Lipara and (1943) in several characters. The former is larger, Platycephala flies together with larvae of C. dia- with the pseudocephalon not divided into lobes, dema, C. nigritarsis and Incertella zuercheri. two-segment antennae and mouthhooks with There have also been very rare records of these conspicuous, closely spaced, small teeth. The larvae found among decaying leaves ofthe apical corresponding larva of C. nigritarsis has a part of stems not inhabited by other species ofin- mouth-hook without teeth and three-segment an- sects. tennae. The structure ofspiracles is also different. The anterior spiracles of the C. flavitarsis larva Acknowledgements. I would like to thank Dr Lech Le- Chowski for his and comments on this have 479 lobes, and the spiracular plate of poste- suggestions paper. rior spiracles is quadrilateral, while the larva of C. nigritarsis has anterior spiracles with 475 lobes References and the spiracular plate of posterior spiracles is oval. The puparium of C. flavitarsis, previously Beschovski, V. 1984: A Review of En- known only from drawings (Waitzbauer 1969), is Zoogeographic demic Palearctic Genera of Chloropidae (Diptera) in

described for the first time in this and illus- 7 paper View of Their Origin and Formation. Acta 2001. trated with original drawings. In his description Bulgarica 21: 3726. of the puparium of C. nigritarsis, Hennig (1943) Blair, K. G. 1932: Some notes on the galls ofLipara lucens merely stated that the puparium bore characters Mg. (Dipt.). 7 Entomologist’s Monthly Magazine 68: 10713. of the third-instar larva. As a result, his descrip- Blair, K. G. 1944: Some galls ofLipara lucens Mg. (Dipte- tion can be to the above hardly compared descrip- 7 ra, Chloropidae). Entomologist’s Monthly Maga- tion of the puparium of C. flavitarsis. zine 80: 1897190. Larvae of C. are J J. H. & V. 1974: flavitarsis typical sapro- Chvala, M., Doskoéil, ., Mook, Pokorny, 24 Grochowska ° ENTOMOL. FENNICA Vol. 19

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