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Larval Morphology of the Antlion Myrmecaelurus Trigrammus (Pallas, 1771) (Neuroptera, Myrmeleontidae), with Notes on Larval Biology

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Larval Morphology of the Antlion Myrmecaelurus Trigrammus (Pallas, 1771) (Neuroptera, Myrmeleontidae), with Notes on Larval Biology TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Zootaxa 3641 (4): 491–500 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3641.4.14 http://zoobank.org/urn:lsid:zoobank.org:pub:B04E6A99-02A2-42EB-A8F5-9665CBB89DF1 Larval morphology of the antlion Myrmecaelurus trigrammus (Pallas, 1771) (Neuroptera, Myrmeleontidae), with notes on larval biology DUŠAN DEVETAK1, VESNA KLOKOČOVNIK1, SAŠKA LIPOVŠEK1,2, ELISABETH BOCK3 & GERD LEITINGER3 1Department of Biology, FNM, University of Maribor, Koroška 160, 2000 Maribor, Slovenia. E-mail: [email protected] 2Medical Faculty, University of Maribor, Slomškov trg 15, 2000 Maribor, Slovenia 3Institute of Cell Biology, Histology and Embryology, Medical University Graz, Harrachgasse 21, 8010 Graz, Austria Abstract Morphology and behaviour of third instar larvae of the Holomediterranean antlion species Myrmecaelurus trigrammus (Pallas) are described. Larvae are facultative pit-builders, they either ambush their prey at the surface, or dig pitfall traps that prey fall in to. Dark brown spots on dorsal and ventral sides of the head and on dorsal side of the thorax are charac- teristic of the larvae. Eye tubercles are not prominent. Jaws are equipped with long bristles, campaniform sensilla, sensilla coeloconica, and digitiform sensilla. A unique feature is the shape of the tips of all three teeth that is screw-like with a polyhedral surface. The body surface is covered with longitudinally grooved bristles and plumose hairs. On the tip of the antennae and on terminal and subterminal parts of labial palps sensilla basiconica occur. On the 9th abdominal segment there are two bulges, each of them bearing four digging bristles. Non-prominent eye tubercles and numerous mandibular bristles are morphological traits of pit-builders. Most of the behavioural traits are related to pit builders, whereas forward movement, waiting for prey without a pit and frequent changing of ambush location are traits of non-pit builders. Key words: Sensilla coeloconica, sensilla basiconica, campaniform sensilla, digitiform sensilla, facultative pit-building antlion Introduction Antlions (Myrmeleontidae) are the largest family of Neuroptera, with about 2000 described species occurring on all continents and most large islands of the world (Stange 2004). Larval antlions have adopted a variety of predation strategies (Mansell 1996, 1999). Only a few antlion species build pitfall traps, and this is considered to be the most specialized strategy for capturing prey (Mansell 1996, 1999; Gepp 2010). The antlion genus Myrmecaelurus Costa includes 50 species and is confined to central and southern Europe, North Africa, and large parts of Asia (Aspöck et al. 2001; Stange 2004). The only European species, Myrmecaelurus trigrammus (Pallas, 1771), has a Holomediterranean distibutional pattern (Aspöck et al. 2001). The oldest description of a M. trigrammus larva dates back to the second half of the 19th century when Redtenbacher (1883, 1884a,b) presented its morphology. In three further descriptions only a few details concerning larval morphology were presented (Doflein 1921; Steffan 1975; Mirmoayedi 2008). Solid depictions of third instar larvae of M. trigrammus were provided later (Willmann 1977; Gepp 2010; Krivokhatsky 2011; Badano 2012), thereof two excellent descriptions based on antlions from the Dodecanese Islands (Willmann 1977) and from Italy, Romania and Turkey (Badano 2012). Notes on behaviour of the larvae are given by Doflein (1921) and Popov (1984). The larva of M. trigrammus is unique among antlions. Its behaviour has been studied in detail because it combines two prey-capture methods; it either ambushes its prey at the surface, or digs pitfall traps that prey fall in to (Doflein 1921; Popov 1984). The use of one capture method or the other depends on a combination of the influences of past net energy gain and the antlion’s most recent change in encounter rate with prey, as it was demonstrated for an unidentified Israeli Myrmecaelurus species (Elimelech & Pinshow 2008). Thus, the larva of Accepted by A. Contreras-Ramos: 21 Mar. 2013; published: 26 Apr. 2013 491 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. the genus Myrmecaelurus occupies an intermediate position between pit-building and non-pit-building antlion larvae. Prior to the present study, no fine structure of body surface of Myrmecaelurus larvae using scanning electron microscopy was documented. It was hypothesised that the larvae will be characterized by traits of both groups of antlions using different capture methods, i.e. pit-builders and non-pit-builders. Material and methods Macedonia: Jasen Reserve: Vlaka near Gorna Belica, 470 m, N 41° 40.99' E 21° 14.3', 6. VII. 2011, 11 third instar larvae. The larvae were found in an open grassland area with single bushes or trees of Quercus trojana, Paliurus spina-christi, Juniperus oxycedrus, Prunus spinosa and Quercus frainetto. Seven larvae are preserved in the first author’s collection, and four larvae are deposited in the Laboratory for Electron Microscopy of the Medical University Graz. Larvae were placed in plastic cups (diameter 10 cm, height 7 cm) or Petri dishes (diameter 9 cm) filled with sand brought from the original collecting site. The larvae were kept in the laboratory at a room temperature of 25±1°C. Larvae were fed with the ants Lasius niger (Linnaeus) and Lasius fuliginosus (Latreille). Feeding took place every day and behaviour of the larvae was recorded with a Sony DCR-HC32E digital video camera. Measurements were conducted on alcohol-preserved third instar larvae using an ocular micrometer. We measured body length (including the mandibles), body width, head length (without mandibles), head width and mandible length. For morphometry see detailed description in Nicoli Aldini (2007). The three teeth on the inside of the jaw are numbered 1–3, starting from the tooth nearest to the base of the mandible (see Doflein 1916; Lucas & Stange 1981). Body colouration was determined using Munsell Soil Color Charts (Soil Survey Division Staff 2000). Larvae were photographed under a stereoscopic zoom microscope Nikon SMZ800 with a mounted digital camera Nikon DS-Fi1, and processed with NIS-Elements F 3.0 software. Digital images captured at different focal planes were assembled using the application Helicon Focus 4.62 Lite. Larval morphology was studied by means of scanning electron microscope (SEM) as well as stereoscopic microscopy. From the field, bodies of larvae were covered with tiny sand particles, so specimens were cleaned using an ultrasound cleaner Sanorex TK 52. Antlions used for electron microscopy were fixed in a mixture of 2 % paraformaldehyde and 2.5 % glutaraldehyde, dehydrated, critical point dried, sputter-coated with gold and palladium, and examined with a digital SEM Zeiss DSM 950. Results Description of the larva Only the third instar larva was studied comprehensively. This is because one can compare it with existing descriptions of third instars of other antlion species. Body colouration (Figs. 1–6): ventral side reddish yellow (Munsell 2000: 7.5YR 8/6), dorsal side pink (7.5YR 7/4) to reddish yellow (7.5YR 8/6). Mandibles (except median margin of mandibles) and distal parts of mandibular teeth brown (7.5YR 4/4). Median margin of mandibles reddish yellow (7.5YR 7/6), very pale brown (10YR 7/4) to light brownish grey (10YR 6/2). Spots on dorsal side of head very dark brown (10YR 2/2) or very dark greyish brown (10YR 3/2) to dark greyish brown (10YR 4/2). Spots on ventral side of head very dark greyish brown (10YR 3/2) to dark brown (10YR 3/3). Pronotum reddish yellow (7.5YR 7/6). Bristles black (10YR 2/1) to very dark greyish brown (10YR 3/2). Dark spots on dorsal part of mesothorax and metathorax brown (10YR 4/3). On ventral part of thorax and abdomen there are no spots. Size: body length (including mandibles) 11.7–15.4 mm, body width (widest part) 4.9–5.3 mm, head capsule length (without mandibles) 1.9–2.5 mm, head capsule width 1.8–2.4 mm, mandible length 1.8–2.4 mm. Head heart-shaped (Figs. 3,4), a little longer than wide. Anterior margin of the head concave (Fig. 7), bearing numerous bristles and hairs. Dorsal side of head capsule dominated by large dark brown spots interrupted by an anchor- or Y-shaped, light reddish yellow or pale brown pattern (Figs. 1, 3). Ventral side of head with characteristic 492 · Zootaxa 3641 (4) © 2013 Magnolia Press DEVETAK ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. darker colouration with a light median line (Fig. 4). Head surface covered with numerous longitudinally grooved bristles (sensilla chaetica) and plumose hairs (Figs. 7, 8, 13, 14). Antenna (Fig. 9) brown, about 2 times longer than maximal width of mandible. The number of flagellar segments varies from 13 to 14. On the distal part of the last flagellomere a group of three sensilla basiconica are found (Fig. 10). Eye tubercles (Figs. 9, 11) not prominent, light yellow to light brown with seven dark brown stemmata. Colour of stemmata may depend up on light adaptation state. Six stemmata occur on dorsal part of eye tubercle (Fig. 11), and seventh stemma is positioned on the tubercle ventrally. Whilst six dorsal stemmata are equally-sized and close to each other, the seventh one is smaller. On eye tubercles several longitudinally grooved bristles are found. Near the base of mandibles, close to eye tubercle, bristles (sensilla chaetica) and plumose hairs occur (Fig.
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