Anti-predator defence mechanisms in sawfly larvae of Arge (Hymenoptera, Argidae)

Larvae of the sawfly Arge (Hymenoptera, Argidae) are exposed to predators such as ants. Their defence mechanisms, which have been almost unstudied, were investigated by behavioural observations coupled to a morphological approach and by testing the bioactivity of several body parts. Arge larvae raised their abdomen when contacted by Myrmica rubra workers. The ants rarely bit a larva and generally retreated immediately, sometimes without contacting it. Most of those few ants that bit a larva then showed an uncoordinated walk. Crude hemolymph from a common species, A. pagana, was a feeding deterrent towards ants. Hemolymph extracts remained active up to a concentration of 0.8 microg DW extract per microlitre solution, and were more active than integument and gut extracts. We also observed ants paralysed by extracts, especially from the gut. It is likely that this toxicity is due to a polypeptide, lophyrotomin, which is known to occur in A. pullata. Six or seven non-eversible ventro-abdominal glands occurred in all species studied (A. fuscipes, A. nigripes, A. ochropus, A. pagana, A. pullata, A. ustulata). These glands contain volatiles. We consider both types of chemicals to be important in defence, and we propose that the paralysing effect is a common feature among Arge species.     

平颜三节叶蜂属一新种暨中国种类检索表(膜翅目:三节叶蜂科)

记述采自浙江的膜翅目三节叶蜂科平颜三节叶蜂属1新种:短顶平颜三节叶蜂Aprosthema brevivertexis Wei sp.nov..该种与同域分布的天目平颜三节叶蜂A.tianmunicum Wei& Wen,2000近似,但前翅翅痣下方具明显烟褐色横斑,1Rs显著长于2Rs室,Rs脉第2段1.3倍于Rs脉第3段长;单眼后区宽长比等于2.2,后部不高于单眼面;颚眼距稍窄于单眼直径;中胸背板前叶无中纵沟,具细纵脊;背面观锯鞘互相远离,侧面观锯鞘端部急尖等,与后者不同.     

平颜三节叶蜂属一新种暨中国种类检索表(膜翅目:三节叶蜂科)

记述采自浙江的膜翅目三节叶蜂科平颜三节叶蜂属1新种:短顶平颜三节叶蜂Aprosthema brevivertexis Wei sp.nov..该种与同域分布的天目平颜三节叶蜂A.tianmunicum Wei& Wen,2000近似,但前翅翅痣下方具明显烟褐色横斑,1Rs显著长于2Rs室,Rs脉第2段1.3倍于Rs脉第3段长;单眼后区宽长比等于2.2,后部不高于单眼面;颚眼距稍窄于单眼直径;中胸背板前叶无中纵沟,具细纵脊;背面观锯鞘互相远离,侧面观锯鞘端部急尖等,与后者不同.     

中国刺背三节叶蜂属一新种暨分种检索表（膜翅目：三节叶蜂科）

记述中国浙江丽水市九龙国家湿地公园分布的刺背三节叶蜂属1新种:丽水刺背三节叶蜂Spinarge lishui Liu,Li&Wei sp.nov..给出了刺背三节叶蜂属世界分种团和中国种类检索表.     

中国刺背三节叶蜂属一新种暨分种检索表(膜翅目:三节叶蜂科)

记述中国浙江丽水市九龙国家湿地公园分布的刺背三节叶蜂属1新种:丽水刺背三节叶蜂Spinarge lishui Liu,Li&Wei sp.nov..给出了刺背三节叶蜂属世界分种团和中国种类检索表.     

Host plant adaptations in myrtaceous-feeding Pergid sawflies: essential oils and the morphology and behaviour of Pergagrapta larvae (Hymenoptera, Symphyta, Pergidae)

Australian pergine sawflies typically feed on eucalypts and other closely-related Myrtaceae, which are known for their high content of essential oils. We describe a novel morphological adaptation of the inner mandibular surface of larval stage Pergagrapta species, which feed on leaves of Melaleuca quinquenervia (Myrtaceae). This 'scopa mandibularis' forms an extensive mesh of setaceous papillae on what is usually the grinding surface of the mandible. Behavioural, chemical and morphological investigations of the sawfly- Melaleuca system suggest that the scopa may function in the physical separation of toxic leaf oils from the diet. The oils are stored in a pharyngeal diverticulum from where they are ejected under two circumstances. Oil from the diverticulum is voided prior to feeding and/or during feeding at night, which indicates a mechanism to eliminate host-associated oils. Larvae rest in close aggregations during the day, when they retain a full diverticulum, but the oils may be emitted for defensive purposes when larvae are disturbed. Chemical evidence suggests that 1,8-cineole, the major component in the M. quinquenervia leaves, is selectively metabolized to a more soluble hydroxycineole. We postulate that the separation and regurgitation of oils is not only a defence mechanism against predators, as usually stated, but also a mechanism by which pergid larvae eliminate oils from their diet, to reduce the toxicity of their food plants.     

Mandibular allometry and male dimorphism in a group of obligately mycophagous beetles (Insecta: Coleoptera: Staphylinidae: Oxyporinae)

Analyses of seven species of the obligately mycophagous staphylinid beetle genus Oxyporus indicate that five species display male mandibular allometry. Among species with dimorphic separations, two general allometric patterns were observed: (1) change in mandibular length at a hypothesized switch point is discontinuous rather than continuous, and (2) the linear slope of mandibular length versus elytral length changes at the hypothesized switch point. The male dimorphisms are likely related to male mating strategies where minor males increase the chances to encounter females by maintaining maneuverability, while major males do the same through intraspecific combat. Male dimorphisms may also be correlated with patterns of fungal usage.     

Characterisation of digestive protease in the rose sawfly,Arge rosae Linnaeus (Hymenoptera: Argidae)

Insect midgut proteases are excellent targets for insecticidal agents such as protease inhibitors. These inhibitors are used for producing transgenic plants, resistant to pests. For achieving this goal, it is necessary to find the nature of specific proteases and their properties for adopting possible pest management procedure. Therefore, characterisation of the enzymes in the gut of the rose sawfly, Arge rosae (Hymenoptera: Argidae), responsible for proteolysis, was performed using a range of synthetic substrates and specific inhibitors. The optimum conditions for general proteases and trypsin were achieved at pH 10. The highest activity for general proteases was obtained at a temperature of 45°C. The use of specific inhibitors and SDS-PAGE (sodium dodecyl sulfate polyacrylamide gel electrophoresis) provided evidence to suggest that most of the proteases belonged to the serine group because of high inhibitory effect of phenyl methane sulfonyl fluoride on total proteolytic activity. Also, inhibition assays and zymogram analysis showed that metalloproteases are present in A. rosae digestive system. These results indicated that A. rosae larvae mainly used serine proteases for protein digestion, with chymotrypsin as the dominant form. The kinetic parameters of trypsin-like proteases using N-benzoyl-dl-arg-p-nitroanilide as substrate indicated that the K _m and V _max values of trypsin in the gut of the fifth instar larvae were 730 ± 17.3 μM and 456 ± 13.85 nmol min^?1 mg^?1 protein, respectively.     

Maternal care in the red-headed spruce web-spinning sawfly,Cephalcia isshikii (Hymenoptera: Pamphiliidae)

We describe oviposition and maternal behavior in the sawfly Cephalcia isshikiiand examine the adaptive significance of this behavior. Females deposited eggs in a single but loose cluster on needles of terminal twigs of spruces, Piceaspp., and remained with the eggs usually on the underside of the twig facing toward the tip. The female attended her eggs until death without taking food but did not follow the first-instar larvae that moved from natal needles even if she survived until then. When the female was disturbed, she usually moved toward the source and attempted to bite it. Though at much lower frequencies, this aggressive behavior was also observed in gravid females and even in males. Field observations and female removal experiments indicated that the female enhanced the survival of the eggs through the reduction of arthropod prédation.     

Genetic diversity and complementary sex determination (CSD) in Dolerus aeneus (Hymenoptera, Symphyta): implications for the conservation of an ecologically-important sawfly

Graminivorous sawfly numbers have declined steadily over recent years as a consequence of agricultural intensification. In spite of these declines and the importance of sawflies (Hymenoptera, Symphyta) as a food source for threatened populations of farmland birds, sawfly conservation studies have been restricted to census-based research. This is largely due to certain aspects of the sawfly life-cycle which make adequate sampling of these insects difficult. However, genetic research provides a valuable insight into population structure that cannot be obtained by traditional ecological means, and one which is essential for the development of conservation management strategies. In this study, we investigate the population genetic structure of the graminivorous sawfly Dolerus aeneus, and conduct a preliminary study to determine whether complementary sex determination (CSD), which can produce sterile diploid males under inbreeding conditions, operates in this species. Our research suggests that fragmentation of sawfly habitat as a result of agri-intensification has not yet acted to isolate D. aeneus populations, although some genetic effects are apparent (inbreeding and low diversity compared with other solitary Hymenoptera). In addition, diploid males were detected which may have compromised fertility and could indicate that CSD is operative in D. aeneus. This study highlights the need for further genetic research in sawflies to assess population structure on a UK-wide scale and to assess the prevalence of diploid males in key species. We discuss our findings in wider context of the genetics of Hymenoptera and the conservation and management of farmland biodiversity.     

Chemical detoxification vs mechanical removal of host plant toxins in Eucalyptus feeding sawfly larvae (Hymenoptera: Pergidae)

The essential oils that characterize the eucalypts and related Myrtaceae pose a challenge for herbivores. Phytophagous insects that feed on oil-rich Myrtaceae have developed specific mechanisms to deal with these oils, some of which are notoriously toxic (e.g. 1,8-cineole). Some of the eight Australian subfamilies in the sawfly family Pergidae are associated exclusively with Eucalyptus and Melaleuca species that often have high concentrations of essential oils. Unexpectedly, the Perginae and Pterygophorinae use different mechanisms to deal with the same toxic components in their respective host plants. Larvae of the Perginae have the inner surface of their mandibles equipped with soft brush-like structures that are unique among phytophagous insects in general. The proposed role of these ancillary mandibular structures in separating leaf oils from nutritive plant matter could be confirmed in experiments with larvae of two pergine species. The oil sequestration is, however, incomplete and chemical gut content analyses by gas-chromatography (GC) revealed that 1,8-cineole does enter the midgut and is metabolised to hydroxycineole. Although the related Pterygophorinae also feed mainly on oil-rich Myrtaceae, they do not sequester the oil and lack morphological structures on their mandibles. Chemical analysis of the gut content of two pterygophorine species showed that they rely solely on chemical detoxification of the relevant plant compounds, with GC demonstrating that the 1,8-cineole is removed far more rapidly and completely than in the pergine species.     

Metabolic dynamics across prolonged diapause development in larvae of the sawfly,Cephalcia chuxiongica (Hymenoptera: Pamphiliidae)

Nutrient metabolism is crucial for the survival of insects through the diapause. However, little is known about the metabolic mechanism of prolonged diapause. The sawfly, Cephalcia chuxiongica (Hymenoptera: Pamphiliidae), is a notorious defoliator of pine trees in southwest China. One of the distinguishing biological characteristics of this pest is the prolonged diapause of about 1.5 years. In this study, the body lipid, carbohydrate (total body sugar, glycogen, trehalose, and glucose), protein, and glycerol contents were measured in diapausing larvae of C. chuxiongica. The results showed that the changes of biochemical composition in C. chuxiongica are associated with the diapause initiation, maintenance, and termination phases. During the initiation phase, trehalose, glucose, and glycerol increased significantly, but glycogen decreased sharply. In general, the lipid, carbohydrate, and glycerol levels decreased gradually across the maintenance phase. At termination phase, the contents of glycogen and lipid persistently decreased, while an increase of trehalose, glucose, and glycerol contents were detected. The protein level was significantly higher at maintenance phase than at initiation and termination phases. It was also found that elevation of trehalose, glucose, and glycerol contents occurred in winter. These implies that the metabolites with altered levels in diapausing larvae of C. chuxiongica are responsible for maintaining a prolonged development and overwintering.     

Gregariousness,field distribution and defence in the sawfly larvae Croesus varus and C. septentrionalis (Hymenoptera,Tenthredinidae)

Summary The larvae of Croesus varus are cryptic and those of C. septentrionalis aposematic. In both species, females lay eggs in groups, but the groups formed by young larvae are smaller in C. varus than in C. septentrionalis. In both species, group size decreases during successive larval instars, and the cryptic species becomes solitary, while the aposematic species remains gregarious during all larval instars. On Alnus shrubs in the field, the distribution of the groups ofC. septentrionalis larvae is more aggregative. They are found mainly on mediumsized shrubs (1.5–2.5 m high), whereas groups of C. varus larvae are also found on larger shrubs. Moreover, during the 2 years of observation, C. septentrionalis was present for a shorter time than C. varus, even when the former was much more abundant than the latter. Thus, conspicuousness of the aposematic species should be amplified by this spatial and temporal aggregation. The efficiency of both defensive strategies is compared and discussed, taking into account these particular field distributions, predation and some parasitism factors, and the chemical defences of both species.     

Superhydrophobic cuticle with a "pinning effect" in the larvae of the iris sawfly, Rhadinoceraea micans (Hymenoptera, Tenthredinidae)

The integument of some sawfly larvae can be easily damaged causing haemolymph bleeding (easy bleeding phenomenon). In the present study, we investigated intact cuticle, cuticle without wax coverage and cuticle replicas of sawfly larvae by using cryo-scanning electron microscopy and contact angle (CA) measurements. The easy bleeder Rhadinoceraea micans was compared to the non-easy bleeder Nematus pavidus (both Tenthredinidae), and the examination showed that the surface structures of R. micans were hierarchically arranged, whereas the cuticle surface of N. pavidus appeared much smoother. The intact cuticle proved to be superhydrophobic in R. micans (CAs: 156° dorsally, 120° ventrally), but not in N. pavidus (CAs: 67° dorsally, 47° ventrally). The wettability of R. micans increased significantly after the waxes had been peeled off. Replicas of the intact cuticle surface were hardly wettable in both species. It is suggested that wax crystals in combination with cuticle microsculptures are the most important features for rendering the integument of the easy bleeder species superhydrophobic. The wax-free tips of the cuticle microsculptures in R. micans are sites where haemolymph droplets can adhere. The integument surface of easy bleeders is assumed to be adapted to their particular defence strategy, simultaneously contributing to the maintenance and "reuse" of released haemolymph droplets on the superhydrophobic surface due to this "pinning effect".     

Economy of food utilization in a gregarious sawfly,Arge nigrinodosa Motschulsky (Hymenoptera: Argidae): Relative abundance of food in the natural habitat and behavioral pattern of larval feeding

Summary Relative abundance of natural food resource, leaves of wild rose, ofArge migrinodosa was evaluated by comparing the food quantity which the larvae composing a colony utilized throughout the larval stage with that which is available in the habitat. In this connection, behavioral pattern of larval feeding was also observed under natural conditions, and its ecological significance was discussed. It was quantitatively clarified that the larvae faced the severe shortage of food and had to disperse to other shoots with the expense of probably high risk. Under pressures of such insufficient supply of food, the larvae follow the most efficient pattern of feeding behavior that they move to the tip of shoot at first and feed regularly downward thereafter. Though individual larvae behaved completely independently of each other, the aggregation appeared to be formed and maintained because all the larvae from the same egg mass follow this fixed feeding pattern.