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The Prothoracic Gland of the Chrysopidae (Neuropteroidea: Planipennia)

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The Prothoracic Gland of the Chrysopidae (Neuropteroidea: Planipennia) Proceedings of the 4th ECE/XIIJ. SIEEC, Godo/lo 1991 into a numerical phenetic species. For this operation a personal computer. The Prothoracic Gland of the Chrysopidae (Neuropteroidea: Planipennia) The prothoracic gland J three suprageneric taxa she R. Giisten and K. Dettner most significant of these w two lobes in all 18 species , Italochrysa (Belonopterygi Another difference shown Key words: green lacewings, defensive secretions, SEM, GC/MS, evolution, phylogeny partly concealed by the pro sa and much more ventrall; Introduction The reservoirs of the ir shape, differ considerably i Some species of green lacewings (Planipennia: Chrysopidae) produce a distinc- of Chrysopa, they nearly J tive scent during molestation, usually described as allylic or faecal-like. As the smaller (relative to body source of the secretion causing the odour, paired prothoracic glands were deter- Cunctochrysa and Chrysopa mined (McDunnough 1909), but later investigations were restricted to a detailed morphological and anatomical description of the glands in Chrysopa per/a by Sulc (1914). An analysis of secretion chemistry in Chrysopa oculata by Blum et al. (1973) revealed the presence of skatole (3-methylindole) as a gland content, which is responsible for the odour. Up to now, all references to the prothoracic glands assume their existence in the strong-smelling species only; a statement by Sulc (1914, p. 3) that they also occur in Chrysoperla carnea, which is inodorous, was obviously overlooked. In order to record the actual distribution of these glands within the family, we examined 20 European chrysopid species for the occurrence of these organs. Besides 18 species of the tribe Chrysopini (Chrysopinae), Italochrysa italica (Chry- sopinae: Belonopterygini) and Nothochrysa fulviceps (Nothochrysinae) were also included in the study. We present the results of investigations of gland morphology and secretion chemistry, and we discuss the probable function of the prothoracic secretion, the morphological, chemical and functional evolution of the gland, and its possible significance for phylogeny and systematics of the family. Methods Fig. 1. Position of gland open Nothochrysa fulviceps (top), ltt Morphological investigations of the prothoracic gland were carried out by species are not drawn to scale. 1 scanning electron microscopy (SEM) of macerated halves of prothoraces, showing - pleural sclerites the gland reservoir as well as properties of the reservoir opening and fine structure of glandular units. For chemical analysis, we used a gas chromatography/mass spectrometry coup- ling (GC/MS), which provides separation of secretion compounds and gives spe- cific mass spectra which help identify individual substances. Morphological and chemical characters of the prothoracic gland evaluated in this study were entered 60 th ECE/XIJL SIEEC, Godol/o 1991 into a numerical phenetic analysis, creating a phenetic tree of the investigated species. For this operation, we used the program NTSYS-pc 1.50 (Rohlf 1988) on a personal computer. e Chrysopidae ipennia) Results The prothoracic gland proved to be present in all the 20 species studied. The three suprageneric taxa showed a number of differences in gland morphology. The tner most significant of these was the shape of the reservoir (Fig. 1): it is divided into two lobes in all 18 species of Chrysopini, into three less clearly separated lobes in Italochrysa (Belonopterygini), but undivided in Nothochrysa (Nothochrysinae). Another difference shown in Fig. 1 is the position of the gland opening, which is GC/MS, evolution, phylogeny partly concealed by the pronotum in Chrysopini, situated just below it in Jtalochry- sa and much more ventrally so in Nothochrysa. The reservoirs of the investigated species of Chrysopini, while very similar in shape, differ considerably in size. In Peyerimhoffina, Chrysoperla and most species '.hrysopidae) produce a distinc- of Chrysopa, they nearly fill out the prothorax, while they are nearly 4 times allylic or faecal-like. As the smaller (relative to body size) in the species of Nineta. Chrysopidia, Mallada, >rothoracic glands were deter- Cunctochrysa and Chrysopa viridana are intermediate. 1s were restricted to a detailed ands in Chrysopa per/a by Sulc 'a oculata by Blum et al. (1973) as a gland content, which is Is assume their existence in the 1 14, p. 3) that they also occur in ly overlooked. ;e glands within the family, we occurrence of these organs. nae), Italochrysa italica (Chry- 1s (Nothochrysinae) were also tigations of gland morphology le function of the prothoracic 1al evolution of the gland, and cs of the family. Fig. I. Position of gland opening slit and reservoir shape in three species of Chrysopidae: Nothochrysa fulviceps (top), Italochrysa ita/ica (middle), Chrysopa per/a (bottom). (The three :c gland were carried out by species are not drawn to scale. Abbreviations: gr - glandular reservoir; go - gland opening; ps 1alves of prothoraces, showing - pleural sclerites · oir opening and fine structure 1phy/mass spectrometry coup- on compounds and gives spe- 1bstances. Morphological and ted in this study were entered 61 The chitinous elements of the glandular units - functional units of a gland cell substances from an exocrim and accessory cells, which create a cuticular ductule - are observable in macerated already been shown in Opilic SEM-preparations (for detailed discussion of glandular units, see Noirot and substances are contained in ti Quennedey 1974). We found that in Chrysopidae, glandular units associated with for defence is found in Nineta the prothoracic gland reservoir are extremely similar to dermal glandular units that gut contents are nearly a distributed over the integument. Interspecific differences concerning shape and tion by all inodorous specie fine structure of glandular units could be shown only in the length of the cuticular Alkenes represent ideal solv duct. In this trait, species of Chrysopa (except Ch. viridana) are most divergent, have little or no repellent or t having ducts of about 80 µm length, compared with 10-30 µm in other species. this hypothetical initial stage Secretion chemistry was analyzed in 13 species from 5 genera of the tribe more and more enhanced du Chrysopini. The 30 substances found can be arranged in 7 groups according to their corresponding with a stepwii chemical structure. A fraction of alkenes is found in all species and, remarkably, its kinds of multicomponent sec composition is very similar in all. The main compound of this fraction is (Z)-4-tri- predators as target organism decene. The other compound groups show various chemical compositions and rected against bats. As the pr< might be derived from quite different biogenetic pathways. They are usually spe- Chrysopidae, it can be used : cific for certain genera, e.g. terpenoids (Chrysopidia), octanoic acid (Chrysoperla), this can be done on different long-chain hydrocarbons (Chrysoperla and Mallada), amides (Chrysopa) apd ska- The subfamilies Nothoch tole (all Chrysopa except Ch. viridana). A special case is observed in Nineta, where gated tribes of Chrysopinae ( no substances other than the alkene fraction are present. ral differences in overall gli examination of this charact( generic taxa within the famil' Discussion sing for the Ankylopterygini, · ni (which are probably not rr The idea that the strong-smelling secretion of Chrysopa species has defensive after the extensive investigati function is· quite straightforward and was first proposed by Melander and Brues (1906) and adopted by most later authors. In view of the great olfactory sensitivity of mammals to skatole (Laffort 1963), bats (Chiroptera) seem to be likely target organisms for the defence secretion, as they are potentially important predators of the night-flying Chrysopidae. Blum et al. (1973), however, found evidence for a repellent effect not only against mammals (mice) but also against arthropods (ants). For a number of reasons, we believe that the prothoracic secretion in the inodorous species serves as a defensive allomone as well, rather than representing some kind of pheromone. Firstly, it appears that these species, just as the odorous ones, discharge their gland contents when molested, even though this is difficult to observe in most cases. Also, it is unlikely that this secretion represents a phero- mone playing a role in sexual interactions, as there is no sexual dimorphism and as there are already other pheromone glands known in these insects (male abdominal glands; Wattebled and Canard 1981 ). Aggregation or alarm pheromones are not to be expected in the solitary green lacewings. The glands of Nineta show the most primitive situation among Chrysopini chemically as well as morphologically. If they are regarded as a model for the initial stage in the evolution of the pro thoracic gland, it must be asked which could be the defensive value of the alkene-containing secretion. A possible hypothesis is that these hydrocarbons act as a solvent and spreading agent for gut contents which are exuded in defence. Discharging faeces or gut contents is the simplest way of chemi- Fig. 2. Dendrograms showing phe cal defence in arthropods (Dettner 1989). The effect of repellent substances con- side: based on multilocus electro tained in gut contents can be both more potent and more prolonged if solvent based on chemical and morpholog 62 :::tional units of a gland cell substances from an exocrine gland are added. A comparable phenomenon has re observable in macerated already been shown in Opiliones (Eisner et al. 1978); in these, however, the active lar units, see Noirot and substances are contained in the glandular secretion. The idea that a mixing of fluids iular units associated with for defence is found in Nineta and other chrysopids is supported by the observation to dermal glandular units that gut contents are nearly always discharged together with the prothoracic secre- ces concerning shape and tion by all inodorous species, and also, less obviously, by the odorous species. the length of the cuticular Alkenes represent ideal solvents for active substances in defensive mixtures, but dana) are most divergent, have little or no repellent or toxic effect on their own (Dettner 1991 ).
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