Larval morphology of Podallea vasseana Navás and Podallea manselli Aspöck & Aspöck from South Africa (: )

A. Möller1*, L.R. Minter2 & P.A.S. Olivier3 1Centre for Microscopy and Microanalysis, School of Physical and Mineral Sciences, University of Limpopo, Private Bag X1106, Sovenga, 0727 South Africa 2School of Health Sciences, University of Limpopo, Private Bag X1106, Sovenga, 0727 South Africa 3School of Molecular and Life Sciences, University of Limpopo, Private Bag X1106, Sovenga, 0727 South Africa

Morphological descriptions of the egg and the three larval instars of Podallea vasseana Navás are presented. Morphological differences between the larvae of P. vasseana and P. manselli Aspöck & Aspöck are noted, and aspects of their biology are discussed. Key words: Neuroptera, Berothidae, Podallea vasseana, Podallea manselli, egg, larva, morphology, biology.

INTRODUCTION

The order Neuroptera comprises a diverse and four species, and the exarmata group of two ecologically important component of the African species. The genus Podallea occurs in the fauna, especially in arid zones to which they Afrotropical Region, the southwestern part of are particularly well adapted (Mansell & Aspöck, the Arabian Peninsula, Comoro Islands and H. 1990). The neuropteran fauna consists of about Madagascar (Aspöck, U. 1990) with P. vasseana and 6000 species worldwide (Aspöck, H. et al. 2001). P. manselli the only known species from South Members of the family Berothidae (hairy or Africa. beaded lacewings) are distributed throughout Most publications dealing with the morphology much of the tropical and warmer temperate regions of the family Berothidae describe the adults of new of the world. It is a relatively small family comprising species or redescribe existing species. Information less than 100 known species. The subfamily on larval morphology, especially of African Berothinae includes 11 genera that are divided berothids, remains almost unknown (New 1986; into two groups, based mainly on the morphology Minter 1990). The only report on immature stages of the genitalia (Aspöck, U. & Nemeschkal 1998). of African berothids was a description of the eggs Group one consists of the more ‘primitive’ genera, and first instar larva of a Podallea sp. (Minter 1990), Berlekrumyia Aspöck & Aspöck, Lomamyia Banks, which was subsequently described as P. manselli and Lekrugeria Navás, but these do not constitute a (Aspöck, U. & Aspöck, H. 1996). monophyletic group. Group two is mono- Tillyard (1916) was the first to observe oviposition phyletic and contains the genus Berotha Walker and to describe the stalked eggs and the first instar and a group of seven genera arranged in an unre- larvae of the Australian berothids, Spermophorella solved trichotomy: Asadeteva Aspöck & Aspöck, disseminata Tillyard and S. maculatissima Tillyard. the adelphotaxa Nodalla Navás and Stenobiella Gurney (1947) recorded a cluster of 12 eggs laid at Tillyard, and the Podallea group (Podallea Navás, the end of a single thin stalk, and other clusters Isoscelipteron Costa, Quasispermophorella Aspöck & with fewer stalked and unstalked eggs of an un- Aspöck and Spermophorella Tillyard). Adults of the identified species of Lomamyia. He also described 15 species of Podallea have been redescribed and the first and third instar larvae of this species and divided into three species groups, mainly based their association with termites and ants. Toschi on the morphology of their genital structures (1964) described the eggs and first instar larvae of (Aspöck, U. & Aspöck, H. 1996). The vasseana L. latipennis Carpenter. MacLeod (1964) described group comprises nine species, including P. the head capsule and cervix of the third instar vasseana and P.manselli, the pellita group consists of larva of L. flavicornis (Walker), and Johnson &

*To whom correspondence should be addressed. Hagen (1981) discovered that berothid larvae E-mail: [email protected] produce an aggressive allomone to immobilize

African Entomology 14(1): 1–12 (2006) 2 African Entomology Vol. 14, No. 1, 2006 their termite prey. Tauber & Tauber (1968) described in diameter; 16–17 irregular micropyles opening the first complete life cycle within the family when around the edge; surface of micropylar promi- they succeeded in rearing all three larval instars of nence and chorion covered with tubercles inter- L. latipennis. Brushwein (1987) described the life linked by ribs, resulting in a characteristic lacy history of L. hamata (Walker). pattern. Eggburster. Remains attached to embryonic cuticle after larvae have emerged from eggs. Base MATERIAL AND METHODS of eggburster arises from embryonic cuticle at anterior end of head, between antennae and Adult specimens of P. vasseana were collected at above mouthparts; consists of a flat disc, 40 µm in Hans Merensky Nature Reserve (23°43’S 30°40’E) length and 9 µm thick, from which the cutting part near Gravelotte, Limpopo Province and P.manselli arises. Main structure of burster. Long, thin cuticular at Dalmada (23°53’S 29°32’E) 8 km east of Polo- filament which extends upwards and curves kwane, Limpopo Province. posteriorly over dorsal surface of head capsule. Light traps were erected in short grassy open Filament. 107 µm long and 5 µm wide; 16–18 small, woodland at Hans Merensky and short grassy widely spaced teeth occur along outer edge; ante- sparse woodland at Dalmada. Adult berothids riorly, the edges of the filament protrude to form a attracted to the light were captured in small glass pair of lips (labia) on either side of the central ridge vials and kept alive for a few days to obtain eggs that bears the teeth; labia taper and disappear from fertilized females. After oviposition, adults posteriorly, towards apex. A well-developed strut were stored in a freezer before pinning and identi- extends backwards, below and parallel to the fila- fication. ment. This may provide leverage during the Individual eggs were transferred to separate rupturing of the chorion. glass vials to minimize handling of the newly hatched larvae and to prevent cannibalism. After Larval stages hatching, a small piece of filter paper was placed in Measurements. Dimensions of body,head capsule each vial to absorb moisture, and to provide shelter length and width, mesothorax and abdominal for the larva. Larvae were fed 1–2 termites daily. width are given in Table 2. Larvae that died were immediately fixed in 70 % Coloration. Head and mouth parts of first instar ethanol. In addition, 1–3 larvae from each of the pale yellow with grey overlay: darker shading at three instars were preserved in the same way. bases of mouthparts; black eyespots. Head and Measurement of larvae and eggs, and the exami- mouthparts of second instar colourless, except for nation of larval morphology were carried out in black eyespots. Head of third instar dark brown; the laboratory using a Zeiss binocular microscope edges of clypeal area clearly demarcated, forming fitted with an eyepiece micrometer. Light micro- a dark circle; dorsally, fine white lines run along graphs were taken with a Wild M5 Stereo micro- ecdysial cleavage lines of head capsule; eyespots scope. Specimens for scanning electron microscopy obscured by dark coloration of head; mouthparts were prepared following conventional methods light brown. Thorax, legs and abdomen of all three (Cross & Mansell 1978; Rupp 1990; Bozzola & instars very similar in coloration. Thorax pale yellow, Russell 1992). Setal and sensilla notations follow with dark brown markings. Legs colourless to pale that of McIver 1975, 1985; Zacharuk 1980, 1985; yellow. Abdomen with characteristic dark and Zacharuk & Albert 1978. light bands: segments 1,3,5,7,9 reddish-brown; segments 2,4,6,8 yellow with brown markings MORPHOLOGICAL DESCRIPTIONS dorsally; segment 10 colourless. Setae pale. Setation. Illustrated as follows: first instar Podallea vasseana (Navas, 1910) Figs 7–15; second instar Figs 22–24; third instar Figs 28–36. Egg (Figs 1–6) Single stalked; elongate-oval in shape; micropylar First instar larva (Figs 7–21, 41–44) prominence at anterior pole. Stalk flexible, but Body. Average length 1.34 mm (range 1.22– capable of supporting the egg in an upright or 1.44 mm); head dorso-ventrally flattened; thorax vertical plane; measurements of egg given in and abdomen round in cross-section; cuticle of Table 1. Micropylar prominence. Disc shaped; 26 µm thorax and abdomen, except for sclerites, Moller et al.: Larval morphology of Podallea vasseana and Podallea manselli 3

Figs 1–6.Podallea eggs and eggbursters.1–5, Podallea vasseana;6, Podallea manselli.1, hatched egg;2, micropylar prominence; 3, chorion; 4, eggburster attached to egg; 5, egg burster; 6, eggburster attached to embryonic cuticle showing the position of the eggburster between the antennae. Abbreviations: A, antenna; B, burster proper; D, disc; E, eggburster; Ec, embryonic cuticle; Fo, fossa; L, labia; m, micropyle opening; mp, micropylar prominence; r, rib; S, strut; T, teeth; Tu, tubercles. extremely thin and flexible, resembling that of of head capsule length to width = 1.57; sides intersegmental membranes, in having a finely parallel anteriorly and slightly tapered posteriorly; folded surface which allows the abdomen to narrower, membranous cervix attached to its expand during feeding. posterior margin. Dorsally with two longitudinal Head capsule and appendages (Figs 8–9, 16–19). ecdysial cleavage lines originating medial to Head capsule: dorso-ventrally flattened; ratio antennal sockets, extending posteriorly to one

Table 1. Egg measurements (mm) of Podallea vasseana and P.manselli.

P.vasseana P.manselli Measurement Mean (range) n Mean (range) n

Length 0.55 15 0.56 11 (0.50–0.60) (0.55–0.57) Width 0.25 15 0.22 11 (0.25–0.30) (0.20–0.23) Stalk 3.64 13 4.4 17 (3.25–4.5) (3.9–6) 4 African Entomology Vol. 14, No. 1, 2006

n third of the distance from posterior margin of head capsule before splitting into two arms; interior arms join medially and continue to posterior margin of head capsule; exterior arms run postero-laterally, about halfway to this margin. Anteriorly the labrum forms a convex, bilobed

Mean (range) margin to clypeal area, each lobe containing a well-developed trichobothrium. Tentorial pits are

n present at the end of short, longitudinal grooves located at the sides of the clypeal area. Eyespots present, forming smooth, round, slightly convex cuticular lenses (ocular area) at antero-lateral corners of head capsule. Flat, irregular scales cover dorsal and ventral surfaces of head capsule. Antennae three-segmented; scape emerges Mean (range) from a well-defined socket in head capsule, longer than wide, covered with overlapping scales; n pedicel much longer than scape, covered with overlapping scales broader at apex, with two uniporous receptors and two short setae disto- laterally near apex, flexed laterally, thickening just distal to the flexure and tapering again at apex; P.manselli. flagellum shorter and much thinner than pedicel, Mean (range)

and posterior third a little thicker than remainder, annulated along entire length with four short,

n sharp setae and one long, grooved seta apically. Mandibles approximately same length as head capsule, outer margin straight, inner margin curves gently from wide base to middle of mandible tapering gradually towards apex; surface of proxi- Podallea vasseana mal half covered with broadly overlapping scales.

Mean (range) Maxillae. Proximal third very thick, covered in overlapping scales with raised apices, giving inner

n margin a serrated appearance; outer surface follows curvature of inner margin of mandible; inner surface of proximal half curves sharply outwards towards mandible, which it envelops, then tapers gently apically forming the feeding tube; a pair of digitiform sensilla present ventro-laterally, near

Mean (range) middle of maxilla. Labium comprises a postlabium, prelabium and pair of labial palps; postlabium

n broad-based, narrowing anteriorly; prelabium a First instar Second instarsquare structure with small Third instar notch anteriorly between bases of labial palps; labial palps four- segmented, first and third segments short, second and fourth much longer; first two segments P.vasseana P.manselli P.vasseana P.mansellicovered with P.vasseana overlapping P.manselli scales, third and fourth (110–130) (130–200) 230 µm (370–400) (300–360) (350–500) (150–200)(110–130)(130–150) (170–220) (110–130) (110–170) (330–440) (220–260) (330) (200–230) (150–200) (230–260) (260–300) (170–230) (260–370) (1.22–1.44) (1.6–2.6) (3.5–3.75) (3.0–3.9) (3.5–4.75) Mean (range) annulated; fourth segment with ten short apical setae as in third instar. Cervix broader than long; . Measurements of the first, second and third instar larvae of head capsule, cervix and prothorax can telescope into each other. = number of specimens; BL = body length; HCL = head capsule length; HCW = head capsule width; MTW = mesothorax width; AW = abdomen width. Table 2 ment BL 1.34 mm 5AW 2.0 mm 10 125 µm 2.57 mm 6 1 165 µm 3.625 mm 12 2 3.49 mm 1 6 4.10 385 mm µm 7 2 330 µm 6 410 µm 7 HCL 178 µm 6 191 µm 12 375 µm 1 385 µm 2 218 µm 6 242 µm 7 HCWMTW 113 µm 133 µm 6 6 125 µm 143 µm 12 12 160 µm 230 µm 1 1 240 µm 330 µm 2 2 172 µm 281 µm 6 6 198 µm 312 µm 7 7 Measure- n Thorax (Figs 7, 11, 41). Prothorax broadens poste- Moller et al.: Larval morphology of Podallea vasseana and Podallea manselli 5

Figs 7–15. Podallea vasseana, first instar larva. 7, Habitus; 8–9, head capsule: 8, dorsal view; 9, ventral view; 10, leg, posterior (A) and anterior (B) view; 11, thorax, dorsal view; 12, abdominal segment 3, dorsal view; 13, abdominal segment 4, ventral view; 14, abdominal segments 8–10, dorsal view; 15, abdominal segments 8–10, ventral view. Abbreviations: A, antenna; cx, coxa; Cvx, cervix; emp, empodium; fm, femur; Md, mandible; Msth, mesothorax; Mtth, metathorax; Mx, maxilla; OA, ocular area; Plb, postlabium; Plp, labial palps; Prlb, prelabium; Pth, prothorax; Scl, sclerite; St, stipe; tb, tibia; Tr, trichobothrium; tro, trochanter; ts, tarsus. 6 African Entomology Vol. 14, No. 1, 2006

Figs 16–21.Podallea vasseana, first instar larva.16–17, antenna:16, pedicel, distal end showing scales, annulations, two uniporous sensilla and setae; 17, flagellum, distal end showing 4 short setae and a long grooved seta; 18, feeding tube formed by the maxilla and the mandible; 19, digitiform sensilla on maxilla; 20–21, abdominal segment 10: 20, ventral surface showing the dome shaped campaniform sensilla and the overlapping scales; 21, anus. Abbrevia- tions: ao, anal opening; ca, campaniform sensilla; Ds, digitiform sensilla; Md, Mandible; Mx, Maxilla; S, seta; Sc, scales; un, uniporous sensilla. riorly to approximately same width as head sclerites a pair of trichobothria are located on capsule, dorsal sclerites form two triangles, elevated conical bases. covered by indistinct scales; one pair of lateral Legs (Fig. 10). Each consists of an unfused coxa, and one pair of dorsal trichobothria present; trochanter, femur, tibia, tarsus, pretarsus with two one pair of spiracles on pleural membrane be- claws and empodium with bilaminar, elastic pad. tween pro- and mesothorax; meso- and meta- Abdomen (Figs 7, 12–15, 20, 21, 44). Dorsal thorax each have two small, smooth, pointed surfaces of segments 1 to 8 lack large sclerites, but a sclerites dorsally and posteriorly; lateral to these single row of very small sclerites are present on Moller et al.: Larval morphology of Podallea vasseana and Podallea manselli 7 margin between segments and intersegmental Sensilla membranes; spiracles present laterally on segments two to seven; a pair of trichobothria project from First instar (Figs 16, 17, 19, 41–43) and elevated conical bases on segments seven and third instar (Figs 37–40, 45–48). eight; segments nine and ten covered with over- Aporous sensilla. Trichodea are represented by all lapping scales; segment ten tapers posteriorly, medium-sized and long setae on head capsule, bearing anal sucker and anus. thorax, legs and abdomen. First instar trichodea are sharply pointed and grooved, while those of Second instar larva (Figs 22–27) third instar have round tips. Basiconica are present Body. C-shaped; length 2.5 mm; cuticle smooth, on the tip of labial palps. Coeloconica: present very soft and flexible except segments 9 and 10 on intersegmental membranes and abdominal which appear more sclerotic; abdominal segments segments 1–6. Campaniform sensilla: present on without thickened sclerites and scales; segments 9 scape of antenna and abdominal segments 9 + 10. and 10 fused. Trichobothria: present on head capsule, thorax Head capsule and appendages (Figs 22, 25, 26). Head and abdominal segments 7 and 8. Digitiform capsule elongated; mouthparts lying close to- sensilla: occur on the latero-ventral surface of gether, pointing forward; clypeal area extends an- maxilla of first instar and on latero-ventral surface teriorly as far as middle of second antennal of third palpomere of third instar larvae. segment; posterior quarter of capsule flattened Uniporous sensilla: present on scape and pedicel dorso-ventrally; head capsule lacks scales; ap- of antennae and abdominal segments 1–8. pendages of head capsule conspicuously shorter when compared to those of the other instars. Second instar Antennae, mandibles, maxillae and labial palps Aporous sensilla. Styloconica: long setae differing segmentation poorly defined; all bases similar in from the trichodea present in first and third instars, diameter; antennae much longer than maxillae in that the bases are conical rather than socket-like. and mandibles, which occupy the space between the antennae and labial palps; anterior half of Podallea manselli U. Aspöck & H. Aspöck, 1988 mandible tapers sharply; apex of maxilla rather blunt; labial palps four-segmented, tapering This study showed that there are very few gradually to a smooth, rounded apex. morphological differences between the larval Legs (Figs. 25, 27). Project forwards, tarsal claws instars of P. vasseana and P. manselli. Almost all and empodia absent. features of taxonomic importance, such as head capsule, thoracic appendages, mouth parts, abdo- Third instar larva (Figs 28–40, 45–48) men and legs proved to be identical in both Body. Average length 3.49 mm (range 3.0– species. 3.9 mm); head relatively small in comparison with Interspecific differences were found in the that of first instar (ratio of body length to head cap- dimensions of the egg (Table 1) and of the larval sule of first instar = 7.52 and that of third instar = instars (Table 2). P.manselli is larger than P.vasseana 16.0). in all measurable morphological features. In the Head capsule and appendages (Figs 29, 30, 37–40). egg the micropylar prominence of P. vasseana has Head capsule: ratio of head capsule length to between 16–18 micropyle openings while that of width = 1.26. Similar in structure to that of first P. manselli has ten openings. instar except that digitiform sensilla are absent on maxillae and present on the latero-ventral surface Remarks of third palpomere. The original descriptions of both P. vasseana and Thorax (Figs 28, 31) and legs (Fig. 36). Similar to P. manselli were based on the morphology of that of first instar. the adults (Aspöck & Aspöck 1996). Only slight Abdomen (Figs 28, 32–35). Segment 10 with differences, such as those associated with the spinneret dorsally. internal genitalia and the presence or absence of Cocoon. Length: 4 mm; width 2.4 mm, silky scales on the wings of the females, are currently white, spun from spinneret, elongate, oval with used to distinguish and validate the two species. loose strands that anchor it to substrate. The lack of further distinguishing features in adult 8 African Entomology Vol. 14, No. 1, 2006

Figs 22–27. Podallea vasseana, second instar larva. 22–24, dorsal view: 22, mouth parts, head capsule and prothorax; 23, meso- and metathorax, and abdominal segment 1; 24, abdominal segments 8,9+10;25, lateral view. Exuvium of the first instar larva attached to segment 10; 26, mouthparts, lateral view; 27, legs. Abbreviations: A, antenna; Clp, clypeal area; Cvx, cervix; ex, exuvium; Hc, head capsule; Msth, mesothorax; Mtth, metathorax; Plp, labia palps; Pth, prothorax. Moller et al.: Larval morphology of Podallea vasseana and Podallea manselli 9

Figs 28–36. Podallea vasseana, third instar larva. 28, Habitus; 29–30, head capsule and mouthparts: 29, dorsal view; 30, ventral view; 31, thorax, dorsal view; 32–33, abdominal segment 4: 32, dorsal view; 33, ventral view. 34–35, abdominal segments 7–10; 34, dorsal view; 35, ventral view; 36, leg, anterior (A) and posterior (B) view. Abbreviations: A, antenna; Cvx, cervix; cx, coxa; emp, empodium; fm, femur; Md, mandible; Msth, mesothorax; Mtth, metathorax; Mx, maxilla; OA, ocular area; Plb, postlabium; Plp, labia palps; Prlb, prelabium; Pth, Prothorax; St, stipes; Scl, sclerite; tb, tibia; Tr, trichobothria; tro, trochanter; ts, tarsus. 10 African Entomology Vol. 14, No. 1, 2006

Figs 37–40. Podallea vasseana, third instar larva. 37–39, labia palps: 37, digitiform sensilla consisting of three sausage-like structures; 38, digitiform sensilla consisting of two sausage-like structures; 39, apex with basiconica sensilla;40, campaniform and uniporous sensilla on scape of antenna, and campaniform sensilla on mandible.Abbre- viations: ca, campaniform sensilla; Md, mandible; Sca, scape; un, uniporous sensilla. females and the remarkable morphological simi- stalked, laid in groups and remain attached to the larities of the larvae indicate a close phylogenetic stalk until after hatching. First and third instar relationship between these species. larvae are campodeiform, with well-developed legs and move rapidly with the use of an anal BIOLOGY sucker. Second instar larvae are inactive and do not feed. When disturbed they flex and straighten Newly laid eggs are white, with dark bands their bodies spasmodically but are not capable of appearing after 2–3 days. The eggs are single- locomotion. This non-campodeiform, ‘dormant’

Table 3. Duration of egg and larval stages of Podallea vasseana and P.manselli.

P.vasseana P.manselli Batch I Batch II Batch I Batch II Days n Days n Days n Days n

Egg 7 17 6 – 9 21 7–9 30 9 60 1st instar 7–8 17 6 10 9–11 15 6–15 13 2nd instar 3–5 5 4 2 4–7 7 6–11 13 3rd instar 32 1 20 1 59 1 3 3 29 1 66 1 69 1 Cocoon 29 1 14 1 – – Adult 2 1 1 1 – – Complete cycle 80 1 48 1 91 1 – Incomplete cycle Moller et al.: Larval morphology of Podallea vasseana and Podallea manselli 11

Figs 41–48. Podallea vasseana, sensilla and sclerites. 41–44, first instar larva: 41, trichobothrium on the prothoracic sclerite showing the complex socket of the trichoid sensilla, and scales on the sclerites; 42, coeloconica trichoid sensilla on intersegmental membranes of abdomen are small and smooth; 43, trichoid sensilla on abdomen are grooved, tapered to a sharp pointed and lie on a smooth basal membrane – segments 1–6 each bear a pair of uniporous chemosensilla; 44, small sclerites on the margins of the segments and intersegmental membranes. 45–48, third instar larva: 45, trichoid sensilla on the head capsule, thorax and abdomen are grooved and have round tips; 46, coeloconica sensilla on abdominal segments 1–6 are small and smooth; 47, uniporous sensilla on abdominal segments 1–6; 48, trichoid sensilla on head capsule and mouthparts are finely sculptured with a sharp point. Abbrevi- ations: Tr, trichobothria; tri, trichoid sensilla; un, uniporous sensilla; Scl, sclerites. stage may represent a hypermetamorphosis. The specific prey of the two Podallea species was Second instar Podallea larvae rested on the bottom not ascertained in the field. The prey item offered of the vial, whereas Lomamyia larvae attached to both species was Cubitermes sp. themselves to the sides of the vial by the last ab- The development of P. vasseana (Table 3) from dominal segment and hung downwards (Tauber egg to adult took 80 days for one larva from & Tauber 1968). The exuvium of the first instar Batch I, and 48 days for another larva from larva remains attached to the last abdominal seg- Batch II, under uncontrolled conditions. The ment of the second instar larva in both species. development of P. manselli was not completed. 12 African Entomology Vol. 14, No. 1, 2006

Only one larva survived up to third instar, but Africa 8: 67–68. died after 91 days. Of the 60 P. vasseana eggs GURNEY, A.B. 1947. Notes on Dilaridae and Berothidae, with special reference to the immature stages of the obtained, only two larvae were reared to the adult Nearctic genera (Neuroptera). Psyche 54: 145–169. stage. Unfavorable laboratory conditions and lack JOHNSON, J.B. & HAGEN, K.S. 1981. A neuropterous of knowledge of the specific food source could larva uses an allomone to attack termites. Nature 289: have influenced the development times and 506–507. mortality of the larvae. It is possible that Podallea is MACLEOD, E.G. 1964. A comparative morphological study of the head capsule and cervix of larval bivoltine. It is known that Lomamyia hamata Neuroptera (Insecta): family Berothidae. Ph.D. thesis, overwinters as a prepupa (Brushwein 1987) but Harvard University, Cambridge, Massachusetts. this was not observed in this study of Podallea. MANSELL, M.W. & ASPÖCK, H. 1990. Post-symposium Podallea vasseana was collected in late summer neuropterological excursions. In: Mansell, M.W., Aspöck, H. (Eds) Advances in Neuropterology. Proceed- (January and February), while P. manselli was ings of the 3rd International Symposium on Neuropterol- collected in spring (September and October). This ogy, Berg en Dal, Kruger National Park. 287–298. may also have influenced the duration of the Government Printers, Pretoria. larval stages. A more thorough study of the McIVER, S.B. 1975. Structure of cuticular mechano- phenology of adults is required to determine the receptors of . Annual Review of Entomology 20: 381–397. number of generations per year in these species. McIVER, S.B. 1985. Mechanoreception. In: Kerkut, G.A. & Gilbert, L.I. (Eds) Comprehensive Insect Physiology, ACKNOWLEDGEMENT Biochemistry and Pharmacology. 71–132. Pergamon The financial support of the Research and Devel- Press, London. MINTER, L.R. 1990. 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Accepted 20 October 2005