Morphology and chaetotaxy of the first instar larva of ...

Zootaxa 2897: 18–26 (2011) www.mapress.com / zootaxa/ Copyright © 2011 · Magnolia Press

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

Morphology and chaetotaxy of the first instar larva of the scorpionfly Sinopanorpa tincta (Mecoptera: Panorpidae) HONG-MIN CHEN1 & BAO-ZHEN HUA1,2 1 Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Education; Entomological Museum, Northwest A & F University, Yangling, Shaanxi 712100, China. E-mail: [email protected], [email protected] 2 Corresponding author

Abstract The morphology and chaetotaxy of the first instar larva of the scorpionfly Sinopanorpa tincta (Navás, 1931) are described and illustrated for the first time with the aid of light microscopy and scanning electron microscopy, with particular attention to the mouthparts and their sensilla. A comparison is made with the larvae of other known Mecoptera, especially Panorpa and Neopanorpa in Panorpidae. Key words: Sinopanorpa, Panorpidae, larva, chaetotaxy, sensillum, mouthparts

Introduction Sinopanorpa Cai & Hua was established with Panorpa tincta Navás, 1931 as its type species and presently also includes S. digitiformis Huang & Hua and S. nangongshana Cai & Hua (Cai et al. 2008). The genus belongs to the most speciose family of Mecoptera, Panorpidae, and can be recognized by the following characters: wing membrane deeply yellow with sooty brown markings; the end of vein 1A extending beyond the origin of vein Rs; vein R2 of wings generally 3-branched; tergum VI of male without anal horn, abdominal segment VII much thinner at the basal 1/3 than the distal 2/3; gonostylus of male genitalia long and slender with a well-developed basal lobe, ventral parameres simple. Although the larval stages of S. tincta were briefly summarized by Cai et al. (2008), a detailed description has not been presented until now. Sinopanorpa tincta (Navás, 1931) is widespread in the Qinling Mountains (Cheng 1957, Chou et al. 1981). During our recent investigation, the first instar larvae of S. tincta were successfully obtained through rearing. Herein their morphology and chaetotaxy are described and illustrated by means of light microscopy and scanning electron microscopy. Chaetotaxy nomenclature primarily follows Hinton (1946) and Cai & Hua (2009a).

Material and methods Adults of S. tincta were captured at Mount Taibaishan (33°47 ′ N, 107°38′ E, 1250 m), Shaanxi Province, from July to August, 2007. The live adults were reared in screen-wired cages (40 × 60 × 60 cm) and provided live potted plants for resting and wet absorbent cotton pads for drinking and maintaining humidity. Adults were provided freshly killed insects as food items, including katydids, grasshoppers, caterpillars and stick insects. Eggs were collected from about thirty female adults and incubated at room temperature in plastic containers layered with dampened absorbent cotton pads. First instar larvae were fixed and observed in this study. The larvae were preserved in 75% alcohol after immersion in 60°C water to straighten and expand the body for light microscopy observation. In order to examine chaetotaxy, the larvae were cut longitudinally along the midventral line, macerated in cold 4% NaOH for 24 hours, rinsed with water, and preserved in 75% alcohol mixed with a few drops of glycerol. The integument was temporarily mounted on a slide to observe the chaetotaxy, which was

18

Accepted by E. Bernard: 2 May 2011; published: 30 May 2011

illustrated with the aid of a camera lucida attached to a Nikon Stereoscopic Zoom Microscope SMZ 1500. The draft drawings were improved with Adobe Illustrator CS3. Larvae were fixed in Carnoy’s fixative (ethyl alcohol:glacial acetic acid = 3:1 v/v) for 12 hours for scanning electron microscopy (SEM) observation. After dehydration in a graded ethanol series, the samples were transferred to isoamyl acetate twice for 30 min, critical point-dried in liquid CO2, sputter-coated with gold, and observed in a JEOL JSM-6360LV scanning electron microscope. The voucher specimens are preserved in 75% alcohol and deposited in the Entomological Museum, Northwest A & F University, China (NWAU).

Results Gross morphology of the first instar larva A newly eclosed larva is 3.6–5.3 mm long and eruciform with a cylindrical body (Fig. 1). The head is hypognathous, with the mandibulate mouthparts directed downwards. The whole body is grayish white, except that the distal regions of the mandibles are reddish brown and the eyes are carmine. Several hours after hatching, the entire head becomes dark yellowish brown. The thorax possesses 3 pairs of thoracic legs, and the abdomen bears 8 pairs of ventral prolegs, one pair on each of the first eight abdominal segments. Abdominal segments I–IX each bear a pair of erect subdorsal annulated processes; these processes are much longer on segments VIII–X than on the preceding segments. Segment X possesses only one long mid-dorsal annulated process. The telson is fleshy and terminates in a protrusible sucker composed of four petaloid structures, which are embellished with more minute setae. The sucker aids movement of the larva. The larval respiratory system is peripneustic, with one pair of prothoracic spiracles and 8 pairs of abdominal spiracles, which occur on the first eight abdominal segments.

FIGURE 1. First instar larva of Sinopanorpa tincta, inset showing the left compound eye. Scale bar = 0.5 mm.

Thoracic legs are 5-segmented (Fig. 4A). The basal coxa is fleshy and the thickest segment. The femur is the longest, with 7 or 8 setae on the lateral side. The tibia is two-thirds the length of the femur, with dense microtrichia laterally on the distal half. A conical process is located on the mesial side of the base of the tibia. The tarsus is transversely wrinkled subdistally and the distal pretarsus tapers towards the apex, covered with numerous microtrichia. The prolegs are transversely wrinkled in the subdistal region, taper towards the apex, and bear numerous microtrichia on the anterior side (Fig. 4C).

Morphology and chaetotaxy of the head The head is well sclerotized, with an average width of 630 μm. The inverted Y-shaped ecdysial line is formed by the coronal and frontal sutures, with the stem medially extending from the occipital foramen and the lateral arms

MORPHOLOGY & CHAETOTAXY OF LARVAL SINOPANORPA TINCTA

Zootaxa 2897 © 2011 Magnolia Press ·

19

diverging downward to the frontoclypeal sulcus (Fig. 2A). The front of the head (or frontoclypeal area) is divided by the frontoclypeal sulcus into the frons above and the clypeus below. The egg burster, a sharp triangular projection that ruptures the eggshell during hatching and then disappears during the first molt, is situated medially close to the distal part of the frons (Fig. 2A). The anterior tentorial pits are located at the lateroventral margin of the intersection of frontal suture and frontoclypeal sulcus. The posterior tentorial pits lie ventrally near the occipital sulcus. A pair of compound eyes, which are carmine with a somewhat paler margin, are situated dorsolaterally between the vertex and genae (Fig. 2B). The hemispherical eyes are composed of about 40 suborbicular facets (Fig. 2D). A pair of three-segmented antennae are located at the medio-anterior region of the compound eyes (Fig. 2B). The basal scape is short and broad, arising from a membranous fossa strutted by an antennal sclerite. The pedicel is thick and subconical, nearly 4× the length of the scape. The membranous apical part of the pedicel bears approximately 32 cup-like depressions which are arranged in two opposed groups on the inner and ventral sides (Fig. 2E). Three rounded smaller sensilla are present among the depressions. The flagellum is slender and conical, slightly shorter than the pedicel (Fig. 2B). Four sensilla basiconica are located on the apical surface of the flagellum. The clypeus is trapezoidal in shape, and subdivided into a basal sclerotized postclypeus and an apical membranous anteclypeus. These two parts have no definite delimitation, which is indicated by a transverse row of four setae.

FIGURE 2. Head of first instar larva of S. tincta, SEM. (A) Dorsal view; (B) Lateral view; (C) Ventral view, with the flagellum removed during preparation; (D) Compound eye; (E) Antenna, ventral view. Abbreviations: Ant, antenna; C, clypeal setae; Cas, cardo-stipes; CS, coronal suture; E, compound eye; EB, egg burster; FCS, frontoclypeal sulcus; F, frontal setae; Fl, flagellum; Fr, frons; FS, frontal suture; Gl, galea; L, lateral setae; Lm, labrum; LP, labial palp; Md, mandible; MP, maxillary palp; O, ocular setae; Pe, pedicel; Pm: postmentum; Prm, prementum; Sc, scape; SO, subocular setae; V, vertical setae; Vx, vertex. Scale bars: A–C = 100 μm; D and E= 50 μm.

20 · Zootaxa 2897 © 2011 Magnolia Press

CHEN & HUA

Thirteen pairs of primary setae are present on the cranium (Fig. 2A–C). Two pairs of clypeal setae are located between the anteclypeus and postclypeus, with the outer pair (C2) about three-quarters the length of the inner pair (C1). Two pairs of frontal setae (F1, F2) are situated in the frontal area along the frontal suture. F1 is near the frontoclypeal sulcus, and slightly longer than F2. The ocular setae (O1, O2) are respectively located at the dorsal and ventral margins of the compound eye; O2 is slightly shorter than O1. Ventral to the compound eyes are two pairs of subocular setae (SO1, SO2). SO1 almost equals to O2 in length and is about twice as long as SO2. Posterior to the eyes are two pairs of lateral setae (L1, L2), with L1 much longer than L2. Located on the vertex are three pairs of vertical setae (V1–V3) arranged in a triangle, with V1 much longer than V3. Most of the setae on the larval head appear to be setiform.

Morphology and sensilla of the mouthparts The labrum is trapezoid and slightly notched midventrally, with two pairs of long setae (Lr1, Lr2) located along its distal margin (Fig. 3A). The epipharynx is membranous, carpeted with rows of appressed microtrichia marginally and inwardly-curved spines medially (Fig. 3A). Three pairs of sensilla basiconica are present on the distal area of the epipharynx. A triangular sclerite, connecting the labrum with the clypeus, is set with a pair of sensilla basiconica and is accompanied by a row of spines along the lateral sides, respectively (Fig. 3A). The mandible is typically dicondylic, with the anterior ginglymus (g) and the posterior condyle (c) (Fig. 3B, C). The former is a cavity that articulates with a convex process of the clypeus near the midventral part of the antenna and the latter is a rounded head adapted to fit into a socket at the lower end of the gena. The strongly sclerotized mandible curves inwards and tapers toward the apex, forming an acuminate triangular tooth apically (Fig. 3B, C). The incisor region bears a median and two basal cusps. The molar region bears a large patch of numerous cone-shaped molar teeth, which are bordered on each side by a row of stiff spines. The molar teeth are sharp in newly hatched larvae, but become worn down with use (Fig. 3C). At the base of the mandible is mesially a bundle of long branched microtrichia (Fig. 3B). Two long and one short seta are present on the outer surface (Fig. 2B). The stipes of the maxilla appears to fuse with the cardo, resulting into a cardo-stipes, with two long and one short seta (Figs. 2C, 3D). The maxillary palp is three-segmented, arising from a thick palpifer, which bears a long seta on the ventral side. The short proximal palpomere bears several spinules apically on the dorsal side (Fig. 3E). The second palpomere has two apical setae (Fig. 3D). The distal palpomere is the longest, bearing eleven sensilla basiconica at the apex. The galea has six sensilla basiconica apically among dense microtrichia (Fig. 3E). Located at the central part of the dorsal side of the galae are a prominent palm-like sensillum and numerous microplates (Fig. 3F). The unsclerotized lacinia, resembling a receptacle attached to the galea, is covered with numerous sensilla and microtrichia (Fig. 3E). The labium is vestigial, greatly reduced in size, with ligula absent. Most parts of the labium are retracted into the capsule; only a pair of 2-segmented labial palps are prominently visible from the basal, mesially separated prementum between the cardo-stipes bases of the paired maxillae (Figs. 2C, 3G). Located at the base between the divided prementum is the opening of the salivary duct (Fig. 3G). Two pairs of minute setae are located on the postmentum (Fig. 2C). The prementum and the proximal palpomere bear distally a tuft of long hairs on the inner side, respectively (Fig. 3H). The distal palpomere is conical and approximately three times as long as the proximal segment, possessing ten sensilla basiconica on the apical side.

Chaetotaxy of the trunk Sclerotized pinacula, from which various setae arise, are regularly distributed on the trunk of the first instar larva. The chaetotaxy of the trunk is illustrated in Figure 5. T1. The prothorax (T1) bears a broad subrectangular prothoracic shield, extending from near the middorsal line downward almost to the lateral pinaculum. The prothoracic spiracle is situated at the posterolateral corner of the shield, and contains 12–15 apertures and an atrial orifice at the center (Fig. 4B). Along the anterior edge of the shield are three punctures and three long setae (XD1, XD2 and SD2), which are 104, 140 and 143 μm long, respectively. Along the posterior edge are two long setae (D1, SD1), one minute seta (MSD1), and one puncture. D1 is about two-thirds the length of SD1. D2 is almost at the same level with D1. They are all clavate except the spiculate



21

MSD1. The lateral seta (L1) is clavate on a lateral pinaculum anteroventral to the spiracle, and slightly shorter than XD2 or SD2. Four subventral setae (SV1–SV4) are situated on a crescent pinaculum dorsal to the coxal cavity, and are 138.5, 94.0, 39.8 and 26.7 μm long, respectively. Only SV2 is slightly expanded at the apex. Further ventrad to the prothoracic leg and near the midventral line is a short ventral seta (V1). Four minute setae are distinctly anterior to V1.

FIGURE 3. Mouthparts of first instar larva of S. tincta, SEM. (A) Epipharynx; (B) Right mandible, front view; (C) Left mandible, caudal view; (D) Maxillae and labium, caudo-ventral view; (E) Right maxilla with a labial palp, front view; (F) Magnification of the basal part of the galea and lacinia, front view, showing sensilla; (G) Labium, caudal view; (H) Left labial palp, front view. Abbreviations: c, condyle of mandible; Cas, cardo-stipes; g, ginglymus of mandible; Gl, galea; IC, incisor cusp; La, lacinia; LP, labial palp; Lr, labral setae; MP, maxillary palp; MR, molar region; MT, molar teeth; Pf, palpifer; Prm, prementum; PS, palm-like sensillum; SB, sensillum basiconica; SD, salivary duct. Scale bars: A–D = 50 μm; E, G and H = 20 μm; F = 10 μm.


CHEN & HUA

FIGURE 4. First instar larva of S. tincta, SEM. (A) Thoracic legs, lateral view; (B) Prothoracic spiracle; (C) Ventral prolegs of abdominal segments VII and VIII (A7, A8), lateral view; (D) First abdominal spiracle. Abbreviations: AO, atrial orifice; Ap, aperture; Cx, coxa; Fm, femur; Pts, pretarsus; T, thorax; Tb, tibia; Ts, tarsus. Scale bars: A = 200 μm; B and D = 10 μm; C = 50 μm.

T2–T3. The meso- (T2) and metathorax (T3) are similar in chaetotaxy. Spiracles are absent. On the dorsal pinaculum are four clavate setae (D1, D2, SD1 and MSD1). D1 is about two-thirds as long as SD1, which measures 124 μm. D2 is as long as MSD1. A minute seta (MD1) is placed at the anterior edge of the dorsal pinaculum. SD2 is also clavate, slightly longer than SD1. Located on the lateral pinacula are two lateral setae (L1, L2), each being accompanied by a short nail-like seta. L1 is clavate and slightly shorter than the setiform L2. Four subventral setae (SV1–SV4) are situated on two detached pinacula, with the clavate SV2 two-thirds the length of spiculate SV1. One ventral seta (V1) and two minute setae (MV1, MV2) are located ventrally to the coxal cavity. A1–A7. The first seven abdominal segments (A1–A7) are similar in morphology and chaetotaxy, each with a pair of spiracles and a pair of ventral prolegs (Figs. 1, 4C). The first abdominal spiracle is composed of 8–10 apertures (Fig. 4D) and is larger than the remaining, which have only 6–8 apertures. The suberect annulated processes on the first seven abdominal segments measure 199 μm. Located on the dorsal pinaculum are four clavate setae (D1, D2, SD1 and MSD1). D1 is twice as long as D2 and about two-thirds the length of SD1. MD1 lies at the anterior edge of the dorsal pinaculum. SD2 on the subdorsal pinaculum is evidently longer than SD1. Posteriorly to the spiracle, a long clavate lateral seta (L1) is located on the same pinaculum with two short setae (ML1, ML3). L2 appears on the lower lateral pinaculum with ML2. L1 measures 1.5× the length of L2. Located on the two subventral pinacula are four setiform setae (SV1, SV2, MSV1 and SV3). SV1 is 172 μm, about twice the length of SV2. Paired ventral setae (V1, V2) are located anteriorly to the proleg. A8. Abdominal segment VIII (A8) resembles the first seven segments by having a pair of spiracles and a pair of prolegs (Fig. 1). Three short clavate setae on the basal part of the fleshy protuberance average 17.9 μm in length.



23

Anterior to the protuberance are two clavate setae (D2, SD1) and one nail-like seta (MSD1). MD1 is situated at the anterior part of the dorsal pinaculum. The setae between the spiracle and the midventral line are arranged in the same way as the proceeding segments. L1 is nearly 1.6× the length of L2. SV1 is 165.4 μm long, corresponding to 1.5× the length of SV2.

FIGURE 5. Chaetotaxy of the trunk of S. tincta, first instar larva. D, dorsal setae; L, lateral setae; MD, minute dorsal seta; ML, minute lateral setae; MSD, minute subdorsal setae; MSV, minute subventral seta; MV, minute ventral setae; SD, subdorsal setae; sp, spiracle; SV, subventral setae; V, ventral setae; XD, prothoracic setae. Scale bar = 0.1 mm.

A9. Abdominal segment IX (A9) bears the longest paired annulated processes (Fig. 1). Spiracles and prolegs are lacking. Three setae on the basal protuberance are arranged similarly to that on A8. Four setae (D2, SD1, SD2


CHEN & HUA

and MSD1) are present on the dorsal pinaculum. SD2 is slightly longer than others. MD1 is located at the anterior part of the dorsal pinaculum. L1 with a short seta lies on the lateral pinaculum. SV1 is nearly as long as L1 and much longer than SV2. V1 is accompanied by a minute seta and 89.1 μm long, similar to V2. The setae on this segment are generally spiculate, with D2 and SD1 not so sharp at the apices. A10. Abdominal segment X (A10) possesses only an annulated process from a middorsal protuberance. Spiracles and prolegs are also absent. The dorsum of A10 resembles a palm-like structure, with one long and two short setae on the dorsal region and four digital processes each terminating in a short seta on the lateral region. Three short clavate setae (D2, SD1 and SD2) are not in a straight line, and almost in the same length. Ventrally, one short seta is present on a gourd-shaped pinaculum and three short setae are present on a narrow pinaculum.

Discussion Larval morphology and chaetotaxy have been documented in several families of Mecoptera. The larva of Nannochristdae is aquatic, elongated and campodeiform, bearing ramous setae on the head and a pair of hooks and papillae on the last segment (Pilgrim 1972). The larvae of Boreidae and Panorpodidae are both scarabaeiform, with the thoracic legs terminating in an elongated cylindrical projection (Peterson 1951, Cooper 1974, Penny 1977, Russell 1982), although the larva of Panorpodidae is eyeless (Byers, 1997). The larva of Bittacidae is eruciform and bears long clavate setae on the head and long fleshy protuberances on the trunk, with two or three branches from which the setae arise (Currie 1932, Setty 1940, Tan & Hua 2008, 2009). In the species-rich family Panorpidae, the larvae of several species in Panorpa and Neopanorpa have been described (Steiner 1930, Yie 1951, Byers 1963, Mampe & Neunzig 1965, Boese 1973, Cai & Hua 2009a, 2009b, Hua & Cai 2009). The larvae are eruciform, with a pair of prolegs on each of the first eight abdominal segments. They bear a pair of annulated processes on abdominal segments I–IX and only a single median annulated process on A10. The first instar larva of S. tincta has not been reported previously for its detailed morphology and chaetotaxy. Although the compound eyes of mecopteran larvae do not increase in number during their development (Paulus 1989), their ommatidial number varies from species to species. The compound eye in first instar larvae is comprised of 20 ommatidia in Panorpa debilis Westwood (Felt 1896), 28 in P. klugi MacLachlan (Miyaké 1912), and 25 or more in P. nuptialis Gerstaecker (Byers 1963) and P. qinlingensis Chou & Ran (Cai & Hua 2009a). In the fourth instar larvae, there are 30–35 facets in P. communis Linnaeus (Steiner 1930) and 24–30 in other Panorpa spp. (Boese 1973). Based on our investigation, the compound eye in the first instar larva of S. tincta is comprised of approximately 40 ommatidia, noticeably greater than that in Panorpa. The epipharynx, maxillary galeae and palps, and labial palps of S. tincta all possess sensilla basiconica, which are assumed to be gustatory chemosensilla and probably allow for a greater ability to reject or accept food items (Zacharuk & Shields 1991). In general, the larvae of Panorpidae carry sensilla basiconica at the apices of the maxillary and labial palps. However, the sensilla basiconica on the maxillary and labial palps are 11 and 10, respectively, in S. tincta, in contrast to 6-8 and 6 in Panorpa (Yie 1951, Cai & Hua 2009a), and 10 and 10 in Neopanorpa (Yie 1951). Sinopanorpa tincta can be readily distinguished from Neopanorpa by the following characters of larval chaetotaxy: (i) the primary setae are long and slender with the distal part slightly expanded in S. tincta, but are very short and gradually expanded from the base to the apex in Neopanorpa; (ii) dense microtrichia are absent along the posterior part of A10 in S. tincta, but present in Neopanorpa; (iii) cranial C1 is almost as long as C2, and SO1 is slightly shorter than O1 in S. tincta, but C1 is only one-third the length of C2 and SO1 is much longer than O1 in Neopanorpa; and (iv) prothoracic SD1 is about 1.5× the length of D1 in S. tincta, but is nearly as long as D1 in Neopanorpa (Yie 1951; Cai & Hua 2009b). Larval chaetotaxy is more diverse in Panorpa (Steiner 1930, Mampe & Neunzig 1965, Yie 1951, Boese 1973, Cai & Hua 2009a), likely reflecting its paraphyly (Willmann 1989, Misof et al. 2000). However, S. tincta can be differentiated from Panorpa in the first instar larva as follows: (i) mesothoracic MV1 and MV2 are situated more posteriorly and closer to V1 in S. tincta; (ii) S. tincta has four setae on the dorsal pinaculum of A8 rather than three as in P. ochraceocauda Issiki and P. longiramina Issiki & Cheng (Yie 1951); and (iii) S. tincta bears no seta anterior to the ventral setae on A9 (one seta in Taiwanese Panorpa (Yie 1951)). All these studies suggest that larvae can provide useful characters for taxonomic and systematic study in Panorpidae. MORPHOLOGY & CHAETOTAXY OF LARVAL SINOPANORPA TINCTA


25

Acknowledgements We thank Tao Li for assistance in rearing the larvae. Dr. James Dunford and Dr. Ernest C. Bernard kindly reviewed the manuscript and helped improve the language. This research was financially supported by the National Natural Science Foundation of China (grant no. 30670255).

References Boese, A.E. (1973) Descriptions of larvae and key to fourth instars of North American Panorpa (Mecoptera: Panorpidae). University of Kansas Science Bulletin, 50, 163–186. Byers, G.W. (1963) The life history of Panorpa nuptialis (Mecoptera: Panorpidae). Annals of the Entomological Society of America, 56, 142–149. Byers, G.W. (1997) Biology of Brachypanorpa (Mecoptera: Panorpodidae). Journal of the Kansas Entomological Society, 70, 313–322. Cai, L.J. & Hua, B.Z. (2009a) Morphology of the immature stages of Panorpa qinlingensis (Mecoptera: Panorpidae) with notes on its biology. Entomologica Fennica, 20, 215–224. Cai, L.J. & Hua, B.Z. (2009b) A new Neopanorpa (Mecoptera: Panorpidae) from China with notes on its biology. Deutsche Entomologische Zeitschrift, 56, 93–99. Cai, L.J., Huang, P.Y. & Hua, B.Z. (2008) Sinopanorpa, a new genus of Panorpidae (Mecoptera) from the Oriental China with descriptions of two new species. Zootaxa, 1941, 43–54. Cheng, F.Y. (1957) Revision of the Chinese Mecoptera. Bulletin of the Museum of Comparative Zoology, 116, 1–118. Chou, I., Ran, R.B. & Wang, S.M. (1981) Study on the classification of the Chinese Mecoptera (I, II). Entomotaxonomia, 3, 1– 22. Cooper, K.W. (1974) Sexual biology, chromosomes, development, life histories and parasites of Boreus, especially of B. notoperates. A southern California Boreus. II. (Mecoptera: Boreidae). Psyche, 81, 84–120. Currie, G.A. (1932) Some notes on the biology and morphology of the immature stages of Harpobittacus tillyardi (Order Mecoptera). Proceedings of the Linnean Society of New South Wales, 57, 116–122. Felt, E.P. (1896) The scorpion flies. Tenth Report of the State Entomologist on the Injurious and Other Insects of the State of New York, 10, 463–480. Hinton, H.E. (1946) On the homology and nomenclature of the setae of lepidopterous larvae, with some notes on the phylogeny of the Lepidoptera. Transactions of the Royal Entomological Society of London, 97, 1–37. Hua, B.Z. & Cai, L.J. (2009) A new species of the genus Panorpa (Mecoptera: Panorpidae) from China with notes on its biology. Journal of Natural History, 43, 545–552. Mampe, C.D. & Neunzig, H.H. (1965) Larval descriptions of two species of Panorpa (Mecoptera: Panorpidae), with notes on their biology. Annals of the Entomological Society of America, 58, 843–849. Misof, B., Erpenbeck, D. & Sauer, K.P. (2000) Mitochondrial gene fragments suggest paraphyly of the genus Panorpa (Mecoptera, Panorpidae). Molecular Phylogenetics and Evolution, 17, 76–84. Miyaké, T. (1912) The life-history of Panorpa klugi MacLachlan. Journal of the College of Agriculture, Imperial University of Tokyo, 4, 117–139. Paulus H.F. (1989) Das Homologisieren in der Feinstrukturforschung: Das Bolwig-Organ der höheren Dipteren und seine Homologisierung mit Stemmata und Ommatidien eines ursprünglichen Fazettenauges der Mandibulata. Zoologische Beitrage (N.F.), 32, 437–478. Penny, N.D. (1977) A systematic study of the family Boreidae (Mecoptera). University of Kansas Science Bulletin, 51, 141– 217. Peterson, A. (1951) Larvae of Insects. Part II. Published by the author. Columbus, Ohio, USA. Pilgrim, R.L.C. (1972) The aquatic larva and the pupa of Choristella philpotti Tillyard, 1917 (Mecoptera: Nannochoristidae). Pacific Insects, 14, 151–168. Russell, L.K. (1982) The life history of Caurinus dectes Russell, with a description of the immature stages (Mecoptera: Boreidae). Insect Systematics & Evolution, 13, 225–235. Setty, L.R. (1940) Biology and morphology of some North American Bittacidae (Order Mecoptera). American Midland Naturalist, 23, 257–353. Steiner, P. (1930) Studien an Panorpa communis L. I. zur Biologie. II zur Morphologie und postembryonalen Entwicklung des Kopfskeletts. Zeitschrift für Morphologie und Ökologie der Tiere, 17, 1–67. Tan, J.L. & Hua, B.Z. (2008) Morphology of immature stages of Bittacus choui (Mecoptera: Bittacidae) with notes on its biology. Journal of Natural History, 42, 2127–2142. Tan, J.L. & Hua, B.Z. (2009) Description of the immature stages of Bittacus plaus Cheng (Mecoptera: Bittacidae) with notes on its biology. Proceedings of the Entomological Society of Washington, 111, 111–121. Willmann, R. (1989) Evolution und phylogenetisches System der Mecoptera (Insecta: Holometabola). Abhandlungen der Senkenbergischen Naturforschenden Gesellschaft, 544, 1–153. Yie, S.T. (1951) The biology of Formosan Panorpidae and morphology of eleven species of their immature stages. Memoirs of the College of Agriculture, National Taiwan University, 2, 1–111. Zacharuk, R.Y. & Shields, V.D. (1991) Sensilla of immature insects. Annual Review of Entomology, 36, 331–354.


CHEN & HUA