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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 Myrmecaelurus trigrammus (Pallas, 1771) (, 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 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. 12, 13). On the base of mandibles campaniform sensilla are found (Fig. 12).

FIGURES 1–6. Third instar larva of Myrmecaelurus trigrammus. 1. Dorsal view. 2. Ventral view. 3. The head, dorsal view. 4. The head, ventral view. 5–6. Tip of the abdomen with digging bristles. 5. Dorsal view. 6. Ventral view. Scale bars 1 mm.

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FIGURES 7–14. SEM micrographs of the head of third instar larva of M. trigrammus. 7. Frontal margin of the head with longitudinally grooved bristles and plumose hairs, dorsal view. 8. Bristles on the ventral surface of the head capsule. 9. Right antenna and the eye tubercle, dorsal view. 10. Tip of the antenna with sensilla basiconica on the last flagellomere. 11. Eye tubercle with six stemmata, dorsal view. 12. The base of the mandible: campaniform sensilla (an arrow), plumose hairs and bristles. 13. Plumose hairs and longitudinally grooved bristles between the eye tubercle and the mandible. 14. Detail of the plumose hairs close to the base of the mandible. Abbreviations used in these and subsequent figures: a—antenna; c—coxa; cl—claw; cs—campaniform sensillum; db— digging bristle; ds—digitiform sensillum; et—eye tubercle; f—femur; md—mandible; mx—maxilla; ph—plumose hair; sb— sensilla basiconica; sc—sensillum chaeticum; sco—sensillum coeloconicum; st—sensilla trichodea; T8—tergite 8; T9—tergite 9; ta—tarsus; tr—trochanter; tt—tibio-tarsus; To2—tooth 2; To3—tooth 3.

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FIGURES 15–22. SEM micrographs of third instar larva of M. trigrammus (continued). 15. Distal part of left mandible with a screw-like tip of second and third tooth. 16. Second and third tooth with a polyhedral surface of the tips. 17. Two sensilla coeloconica on the polyhedral tip surface. 18. Sensillum coeloconicum on the tip of the second tooth.19. Position of digitiform sensillum on the left maxilla. Note short denticles (asterisk) on the mandible. 20. Magnified digitiform sensillum. 21. Labial palp with two groups of sensilla basiconica (arrows). 22. Sensilla basiconica on the tip of the labial palp.

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FIGURES 23–30. SEM micrographs of third instar larva of M. trigrammus (continued). 23. Sensilla basiconica on the median part of the last segment of labial palp. 24. Fine granular sculpture on the pronotum. 25. Lateral tufts of hairs on ventral side of 6th abdominal segment. 26. Longitudinally grooved bristles (sensilla chaetica) on femur of the hind leg. 27. Hind leg. 28. Tarsus of fore leg. 29. Two sensilla trichodea between the claws of fore leg. 30. Bulge with four digging bristles on the tip of the abdomen.

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Length of mandibles equal to head capsule width. Mandibles with three teeth (Figs. 3, 4), thereof tooth 2 is the longest and in contact with tooth 3. Tooth 3 is the shortest one. Distance between tooth 1 and 3 only slightly longer than greatest mandibular width. Shape of teeth in M. trigrammus is unique among antlions studied so far by means of electron microscopy (Eisenbeis & Wichard 1987; Nicoli Aldini 2007; Satar et al. 2007; Cesaroni et al. 2010; Pantaleoni et al. 2010; Devetak et al. 2010a,b). Tips of all three teeth are not smoothly pointed like in other antlion species, but they are screw-like with a polyhedral surface (Figs. 15–17). There are no bristles between tooth 2 and 3, which is characteristic of Myrmecaelurus larvae (Fig. 4). Mandible typically with 8 to 9 (rarely 7) black long bristles (sensilla chaetica) between mandibular base and tooth 1, and 2 to 4 bristles between tooth 1 and 2 (Figs. 3, 4). There are no bristles between the third tooth and pointed end of mandible. Bristles on exterior margin of mandible (Fig. 3) extend from the proximal end of jaws to the base of pointed end of mandible and most of them are significantly longer than tooth 3. Mandibles and maxillae bear a great number of sensilla coeloconica (Figs. 17, 18). The sensilla are set on the floor of relatively shallow depressions in the cuticle and are scattered in the whole length of the jaws including mandibular teeth. On the distal end of the maxilla a single digitiform sensillum occurs (Figs. 19, 20). On distal mandibular surface a few denticles occur (Fig. 19). Labial palps are four-segmented. On their distal segment two groups of sensilla basiconica occur, one group on the tip, the other subterminally (Fig. 21, 22). Subterminal group of sensilla basiconica is sunk into a deep depression (Fig. 23). On prothorax, dorsally, two dark parallel lines occur, and on each lateral side of mesothorax and metathorax there is a large dark brown spot (Fig. 1). Prothorax has fine granular sculpture with longitudinally grooved bristles and plumose hairs scattered on its dorsal side (Fig. 24). Paired segmental tufts of long, longitudinally grooved bristles laterally fringe the meso- and metathorax and abdominal segments (Figs. 1, 2, 25). Legs with longitudinally grooved bristles (Fig. 26). In the hind legs tibia and tarsus fused forming tibio-tarsus (Fig. 27). The tip of the tarsus of all legs has two claws (Fig. 28). Between the claws two short sensilla trichodea occur (Fig. 29). Hind legs without dark spots. Tergite 8 with 2 pairs of short stout bristles (Figs. 5, 31). Sternite 8 with a number of short digging bristles (Fig. 6). Tergite 9 with two prominent bulges distally, each of them bearing four short black digging bristles (Figs. 5, 6, 30, 31). Besides them, two rows of digging bristles, distal row with three bristles on each side of the bulge, and proximal row containing 8 bristles on each lateral side of tergite 9 (Fig. 31).

FIGURE 31. Tip of the abdomen: 8th and 9th tergite with digging bristles. Legend: 1–8: proximal row of digging bristles; I–III: distal row of digging bristles; a–d: a group of digging bristles on the bulge.

Behaviour of second and third instar larvae

On the surface of its natural substrate, the larva moves forward or backward. Taken out of its typical habitat, it

LARVAL MORPHOLOGY OF MYRMECAELURUS TRIGRAMMUS Zootaxa 3641 (4) © 2013 Magnolia Press · 497 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. starts moving forward rapidly. The larva moves backwards when it buries itself in the sand or when it comes across an obstacle. The larva of Myrmecaelurus is a facultative pit-builder, using both mechanisms of prey capture, namely it usually builds a pit, but occasionally waits for the prey without constructing a pit. In presence of prey in the pit, sand tossing, grasping and submersion were recorded. Most of the behavioural traits are related to pit- builders.

Discussion

The larval morphology of the Western Palearctic antlion taxa has been described by means of scanning electron microscopy only for a few species: formicarius Linnaeus (Eisenbeis & Wichard 1987), Myrmeleon inconspicuus Rambur (Nicoli Aldini 2007), Myrmeleon bore (Tjeder) (Nicoli Aldini 2007), Myrmeleon mariaemathildae Pantaleoni, Cesaroni & Nicoli Aldini (Pantaleoni et al. 2010), Myrmeleon yemenicus Hölzel (Devetak et al. 2010b), tetragrammicus (Fabricius) (Satar et al. 2007), Neuroleon microstenus (McLachlan) (Devetak et al. 2010a), Gymnocnemia variegata (Schneider) (Cesaroni et al. 2010) and Megistopus flavicornis (Rossi) (Cesaroni et al. 2010). Existing morphological descriptions of Myrmecaelurus trigrammus larvae, limited to old scattered publications—with the exception of Willmann’s description in 1977 and Badano’s review of European antlion larvae (2012)—are generally incomplete. Larvae of M. trigrammus have non-prominent eye tubercles and numerous bristles on the jaws, which are both characteristic for pit-builders, like Myrmeleon species (e.g. Eisenbeis & Wichard 1987; Nicoli Aldini 2007). Each eye tubercle in M. trigrammus larvae bears seven stemmata. Thus, Mirmoayedi’s statement that larval eye in this species is composed of eight stemmata (Mirmoayedi 2008) is not correct. In the eyes of all larval antlion species studied so far (see Jockusch 1967; Eisenbeis & Wichard 1987; Nicoli Aldini 2007; Satar et al. 2007; Cesaroni et al. 2010; Pantaleoni et al. 2010; Devetak et al. 2010a,b) an eye tubercle bears seven stemmata. Six of them are on the dorsal anterior part of the tubercle, and the seventh one oriented ventrally is a rudiment (e.g. Jockusch 1967; Lipovšek Delakorda et al. 2009). Screw-like tips of the mandibular teeth probably serve to anchor the jaws firmly in the body of a prey. On the abdominal tip of M. trigrammus two groups of digging bristles are arranged in the same way as in Distoleon tetragrammicus (Satar et al. 2006) and in a similar pattern as in Neuroleon species (Steffan 1975; Devetak et al. 2010a). Out of nine basic sensillum types that have been described in larval antlions (Lipovšek Delakorda et al. 2009 and this paper), all except the dolichasters have been found in M. trigrammus larvae. Among mechanosensitive receptors (see Doflein 1916; Römer 2003), four types of bristles and hairs were noted, namely longitudinally grooved bristles—sensilla chaetica, hairs—sensilla trichodea, plumose hairs and digging bristles. Tufts of the mechanoreceptive bristles, sensilla chaetica, positioned on the thorax, play a certain role in detection of substrate vibrations produced by prey moving on sand surface (Le Faucheux 1972). Two types of chemoreceptors—sensilla basiconica and sensilla coeloconica—occurring also in four other antlion species (Lipovšek Delakorda et al. 2009; Devetak et al. 2010a,b) were recorded in M. trigrammus. A unique feature is the finding of two groups of sensilla basiconica occuring on labial palps of M. trigrammus, one terminal and one subterminal. Both receptors—sensilla basiconica and sensilla coeloconica—play a role in detection of chemical substances important in feeding and sensing the ’s environment (Zacharuk 1985; Galizia 2008; Glendinning 2008). Koch (1983) described chemoreceptor sensilla on the mandibles of Euroleon and Satar et al. (2006) mentioned antennal sensilla in Distoleon. These receptors play a role in antlions during catching behaviour when prey is grasped and fixed with mandibles and tip of mandibles and teeth penetrate a prey. It is suggested that antlions estimate taste of prey with these chemoreceptors (Glendinning 2008). Campaniform sensilla, found in M. trigrammus larvae on the mandibles, were noted in other antlion species on the legs and on the distal parts of the abdomen (Devetak et al. 2010a). It is presumed that the abdominal campaniform sensilla in N. microstenus play a role in the control of digging in sand (Devetak et al. 2010a), whilst those on the legs have a proprioceptive role (Römer 2003). A special type of cuticular receptor, the digitiform sensillum, is reported here for antlion larvae for the second time (Lipovšek Delakorda et al. 2009) and figured for these for the first time. In Neuroptera, the digitiform

498 · 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. sensilla were noted first in larvae of the family Dilaridae (Minter 1992). The functional significance of that receptor type is unclear (e.g. Giglio et al. 2003). In elaterid larvae the sensilla have been suggested to function as mechanoreceptors, responding to contact and to substrate vibrations (Zacharuk et al. 1977; Zacharuk 1985). These presumed vibration receptors provide contact with the walls of the larval tunnel in the soil, thanks to their position on the labial and maxillary palps. Later, studies on the mouthparts of scarabaeid larvae revealed the putative function as hygro-/thermoreceptive organ (Eilers et al. 2012). Their role in antlion larvae is not clear and should be confirmed electrophysiologically. Myrmecaelurus larvae are predators, using both mechanisms of prey capture. Elimelech & Pinshow (2008) estimated costs in Israel Myrmecaelurus sp. by measuring resting and activity metabolic rates and determining the duration of pit maintenance at various encounter rates with prey. Benefits were estimated from the energy gained per prey item captured at different encounter rates. The proportion of antlions that constructed pits was higher when they were in intermediate body condition than when in good or in poor body condition (Elimelech & Pinshow 2008). Ambushing without a pit may serve as a default when physiological constraints limit the larvae’s ability to invest in pit construction and maintenance, or when larvae are sated (Elimelech & Pinshow 2008). In conclusion, Myrmecaelurus larvae are facultative pit-builders, performing both characters of pit-builders and non-pit-builders. Non-prominent eye tubercles and numerous mandibular bristles are morphological traits of pit-building antlions. Pit construction, sand tossing, grasping and submersion are behavioural traits of pit-building antlions. Forward movement on sand surface and waiting for prey without constructing a pit are two non-pit- building behavioural traits.

Acknowledgements

We thank Dr. Davide Badano (Sassari) and an anonymous reviewer for critical comments on an earlier version of the manuscript. This project was supported by the Slovenian Research Agency within the Biodiversity Research Programme (Grant No. P1-0078).

References

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