Postembryonic development of the cervicothoracic skeleton of Euborellia annulipes (Lucas) (Dermaptera: Labiduridae)

The nymphal and adult cervicothoracic skeleton of Euborellia annulipes (Lucas) is described and discussed.     

The vitellogenin of the ring-legged earwig,Euborellia annulipes (Lucas) (Insecta: Dermaptera)

Using Polyacrylamide gel electrophoresis, labelling procedures and immunological methods, an extraovarian synthesis of vitellogenin has been demonstrated in Euborellia annulipes. Electropherograms of the haemolymph of 1-wk-old mated females show an extra protein fraction on the second day after mating; the concentration of this fraction in the haemolymph falls from the fifth day after mating. The fatbody incorporates [³H]leucine into the proteins synthesised by it during the period of oocyte formation. The proteins extracted from the fatbody and ovary of females of the first reproductive cycle show a high level of radioactivity 3 days after mating, suggesting a simultaneous release of proteins from the fatbody and uptake by the oocytes. On the other hand, TCA-precipitable fractions of the ovary, obtained from females on the pre-ovipositional day, record a high [³H]activity but similar fractions from the fatbody only show minimum radioactivity. Antibodies prepared against the antigens obtained from the crude yolk extract reacted with the proteins from the haemolymph and also with the proteins from fatbody extracts of females of the first reproductive cycle.     

Postembryonic development of the cervicothoracic muscles of Euborellia annulipes (Lucas) (Dermaptera: Labiduridae)

The cervicothoracic muscles of nymphal and adult Euborellia annulipes (Lucas) are described, the former for the first time. In the adult, eight new muscles are identified, while the nymphs possess a further seven muscles that disappear at maturation or before. Otherwise the same muscles occur in nymphs as in adults, though some nymphal muscles are less clearly separated from one another than their adult homologs. The attachment sites of certain muscles show a number of slight differences between nymphs and adults. The work emphasizes the necessity of taking the immature musculature into account in assessing the muscular pattern represented in an insect order.     

Paternity and egg cannibalism in the ringlegged earwig Euborellia annulipes (Dermaptera: Anisolabididae)

Cannibalism is a common occurrence in nature, and many cannibals prey on relatively small and defenseless life stages, such as eggs or young juveniles. Such behavior provides many benefits to the cannibal, but cannibalistic individuals also face risks, including the cost of decreasing their inclusive fitness by eating close relatives such as siblings or offspring. This risk can be mitigated if cannibals can recognize and avoid preying on kin. Here, we tested whether male ringlegged earwigs Euborellia annulipes avoid cannibalizing eggs that they had sired. In this species, females care for their own eggs, but males provide no care and frequently prey upon eggs. We found that when presented with an unguarded clutch of eggs, male earwigs nearly always cannibalized some eggs, but that the proportion of eggs eaten was smaller if the male had sired the clutch. This suggests that males can distinguish between their own offspring and unrelated offspring and that they avoid harming their kin.     

Identification of 10 microsatellite loci in the earwig Labidura riparia (Dermaptera,Labiduridae)

Ten microsatellite loci were isolated from the earwig Labidura riparia (Pallas, 1773). The polymorphism of the loci was assessed in 24 individuals from one population. The number of alleles ranged from four to 11 alleles and observed and expected heterozygosities from 0.250 to 0.833 and from 0.551 to 0.861, respectively.     

Effects of insecticide sublethal concentrations on the ringlegged earwig, Euborellia annulipes (Lucas)

Integrated pest control of the sugarcane stem borer (SSB), Chila infuscatellus, recommends the use of a voracious predator as the ring legged earwig (RE), Euborellia annulipes (Lucas) and insecticide application. This study was conducted to determine sublethal effects of two recommended pyrethroid insecticides, cypermethrin and deltamethrin, on predation, the number of eggs/batch, % egg hatch and the developmental period of the offspring of RE adults. Sublethal concentrations of cypermethrin (10% EC) and deltamethrin (3% EC) at LC1 and LC10 were topically applied with a micro-applicator on tested individuals with direct and indirect methods. They were applied directly once on adults of RE or on ten 3rd instar larvae of SSB which were then fed to RE adults. Surviving adults were further investigated. Insecticide-treated RE adults with LC1 and LC10 of cypermethrin showed significantly slower onset of predation averaged 14.20 and 18.35 h, lower number of preys killed, averaged 4.8 and 4.2 larvae/d, respectively, as compared to those of untreated ones. The corresponding figures of treated adults with deltamethrin were 30.5 and 42.2 h, 4.6 and 3.8 larvae/d respectively. Similar results were obtained on RE adults that fed on insecticide-treated 3rd instar SSB larvae with the same concentrations of insecticides. The only different result was recorded on the number of preys killed by indirectly treated adults with LC1 of cypermethrin and deltamethrin averaged 9.6 and 8.8 larvae/d, respectively, which was not significantly different as compared to those of untreated ones. Insecticide-treated RE adults with LC10 of cypermethrin and deltamethrin showed significantly lower number of eggs/batch averaged 31.4 and 27.4 eggs, lower % egg hatch averaged 29.8 and 31.7%, prolonged developmental period of the offspring averaged 91.8 and 113.0 d, respectively, as compared to those of untreated ones. The corresponding figures of those from RE adults that fed on insecticide-treated 3rd instar SSB larvae were 30.7 and 37.5 eggs, 36.1 and 32.8%, 95.6 and 113.7 d, respectively, as compared to those of untreated ones. This study concluded that sublethal concentrations of cypermethrin and deltamethrin at LC1 and LC10 which did not kill RE adults reduced their predation, lowered the number of eggs/batch and % egg hatch, and prolonged the developmental period of their offspring. It is suggested that RE may not be totally safe for combination use with cypermethrin and deltamethrin in controlling SSB.     

Temperature-dependent functional response of Euborellia annulipes (Dermaptera: Anisolabididae) preying on Plutella xylostella (Lepidoptera: Plutellidae) larvae

Temperature mediates trophic interactions, including relationships between insect pests and predators, and functional response studies are often used to determine the suitability of predators as biocontrol agents. We investigated the effects of temperature on the functional response of Euborellia annulipes (Lucas) (Dermaptera: Anisolabididae) preying on Plutella xylostella (L.) (Lepidoptera: Plutellidae) larvae. Predation rate, type of functional response, attack rate (a'), handling time (T_h), and maximum predation rate (T/T_h) of the predator were estimated using seven prey densities and three thermal conditions. The functional response of E. annulipes to P. xylostella was temperature-dependent, type III under the lower temperatur (18°C and 25 °C) , and type II at 32 °C. We observed increasing values of a’ in 25 °C and 32 °C, decreasing values of Th and highest T/T_h as the thermal condition increased. Our findings suggest that E. annulipes could be effective to control P. xylostella under different thermal conditions, however its predation behavior changes according to temperature variation.     

Quantification of cockroach allatostatin-like peptide and its myotropic effects in males of the earwig Euborellia annulipes

A monoclonal antibody to allatostatin I of the cockroach Diploptera punctata was used to establish a competitive enzyme‐linked immunosorbent assay for quantification of allatostatin‐like peptides in the hindgut of the adult male earwig, Euborellia annulipes. Hindguts of 0‐day males contained significantly more allatostatin‐positive material than those of 8‐day males fed on catfood. However, males starved for the first 8 days of adult life had significantly higher levels of allatostatin‐positive material than those of either 0‐day or of 8‐day fed males. Hindguts from 0‐day old males exhibited lower spontaneous motility in vitro than those from 8‐day males. Hindguts from males at both ages responded to allostatin with reversible, dosage‐dependent decreases in hindgut motility, and responded to proctolin with reversible, dosage‐dependent increases in hindgut motility. When both allatostatin and proctolin were applied to hindgut preparations simultaneously and in equal concentrations, the response varied with the stage of the male. Starvation enhanced hindgut motility and abolished the response to allatostatin, but not to proctolin. These results indicate the presence of material similar to cockroach allatostatins in male earwigs, and that the levels change with age and physiological stage. Furthermore, such peptides may indeed be regulatory neuropeptides and could modulate hindgut contraction. There was an increase in sensitivity to exogenous allatostatin in the hindgut during development from day 0 to day 8 in feeding males, but a loss in sensitivity in response to starvation; sensitivity to exogenous proctolin also increased with age, but such responsiveness was not diminished by starvation.     

FMRFamide-like material in the earwig, Euborellia annulipes, and its functional significance.

The neurosecretory system of the earwig, Euborellia annulipes, contained material similar to that of FMRFamide, as shown by immunocytochemistry. Within the brain were two pairs of darkly staining perikarya in the medial protocerebrum, and up to four pairs of immunoreactive cells in the lateral protocerebrum. The corpora allata appeared immunoreactive in 10-day females, but not in 2-day-old adults. Additionally, immunoreactive material was detected in midgut endocrine cells of both 2- and 10-day-old females. FMRFamide at 1 to 100 nM did not inhibit juvenile hormone production by earwig corpora allata in vitro. This was true of glands of low activity from 2-day cat food-fed or starved virgin females, 10-day starved females, and those of relatively high activity from 10-day-old, cat food-fed females. In contrast, FMRFamide at 50 and 100 (but not at 1) nM stimulated gut motility in vitro in distended guts from 2-day fed females. Preparations from starved females and those from 10-day fed females (in which feeding behavior is on the decline) did not respond to exogenous FMRFamide with enhanced rates of contraction. Lastly, preparations from females starved for 7 days and subsequently fed for 3 days responded to 10 nM FMRFamide with increases in gut motility.     

Right-handed penises of the earwig Labidura riparia (Insecta, Dermaptera, Labiduridae): evolutionary relationships between structural and behavioral asymmetries

The number of penises vary in the insect suborder Forficulina (order Dermaptera; earwigs). Males of the families Diplatyidae, Pigidicranidae, Anisolabididae, Apachyidae, and Labiduridae have two penises (right and left), while those of the Spongipohridae, Chelisochidae, and Forficulidae have a single penis. The proposed phylogenetic relationships among these families suggest that the single‐penis families evolved from an ancestor possessing two penises. To date, examinations of double‐penis earwig species have found that only a single penis is used per single copulation. These diversities in structural and behavioral aspects of genitalia raises the following intriguing questions: How are the two penises used? Why did a penis degenerate in several earwig families, and which one was lost? To address these questions, structural and behavioral asymmetries were examined in detail for a representative species Labidura riparia (Labiduridae). Although there was no detectable morphological differentiation between the right and left penises, male L. riparia predominantly used the right one for insemination. This significant “right‐handedness” developed without any experience of mating and was also manifested in the resting postures of the two penises when not engaged in copulation. However, surgical ablation of the right penis did not influence the insemination capacity of males. In wild‐caught males, only about 10% were left‐handed; within this group, abnormalities were frequently observed in the right penis. These lines of evidence indicate that the left penis is merely a spare intromittent organ, which most L. riparia males are likely never to use. Additional observations of five species of single‐penis families revealed that the left penis degenerated in the common ancestor of this group. Considering the proposed sister relationship between the Labiduridae and the single‐penis families, it is possible that such behavioral asymmetries in penis' use, as observed in L. riparia, are parental to the evolutionary degeneration of the infrequently used left penis. J. Morphol., 2006. © 2006 Wiley‐Liss, Inc.     

Possible atavisms of genitalia in two species of earwig (Dermaptera), Proreus simulans (Chelisochidae) and Euborellia plebeja (Anisolabididae)

Male and female genitalia generally show a rapid evolutionary rate, which raises the problems related to homologization and the determination of the polarities of evolutionary changes. In earwigs (Dermaptera), multiple or branched female sperm-storage organs (spermathecae) have been reported for members of the Karschiellidae, Pygidicranidae, and Diplatyidae, collectively termed the “basal” Dermaptera. Whether the complicated spermathecae represent a plesiomorphy or an apomorphy has not been resolved. Here I report the occurrence of multiple or branched spermathecae in gamma-irradiated samples of two earwig species, Euborellia plebeja (Dohrn, 1863) (Anisolabididae) and Proreus simulans (Stål, 1860) (Chelisochidae), which belong to the “higher” Dermaptera (Apachyidae, Labiduridae, Anisolabididae, Spongiphoridae, Chelisochidae, and Forficulidae). Females belonging to the higher Dermaptera normally have a single-unbranched spermatheca. I discuss examples of possible atavisms in relation to the evolutionary pathways of spermathecal morphology. Possible atavisms in the number of male organs for sperm transfer (virgae) are also reported.     

Predation potential of the earwig Euborellia annulipes on fruit fly larvae and trophic interactions with the parasitoid Diachasmimorpha longicaudata

The earwig Euborellia annulipes (Lucas) (Dermaptera: Anisolabididae), a generalist predator, has been observed in fruits infested with fruit fly larvae, which are frequently parasitized by parasitoid wasps. Neither the capacity of earwigs to predate on fruit flies nor intraguild interactions between earwigs and fruit fly parasitoids have been investigated. Here, we studied in laboratory conditions the predation on the fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) by the earwig E. annulipes, and whether parasitism of fruit fly larvae by the parasitoid wasp Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) influences predation by the earwig. We evaluated the predation capacity, functional response and prey preference of E. annulipes for parasitized and non-parasitized fruit fly larvae in choice and no-choice tests. We found that earwigs prey on second- and third-instar larvae and pupae of C. capitata and consumed larger numbers of second-instar larvae, followed by third-instar larvae and pupae. Females prey on larger numbers of fruit flies than did males, regardless of the prey developmental stage, but both sexes exhibited a type II functional response. Interestingly, males killed but did not consume fruit fly larvae more than did females. In no-choice tests, earwig females consumed equal numbers of parasitized and non-parasitized fruit fly larvae. However, in choice tests, the females avoided feeding on parasitized larvae. Subsequent tests with hexane-washed parasitized and non-parasitized larvae showed that putative chemical markings left on fruit flies by parasitoids did not drive the earwig preference towards non-parasitized larvae. These findings suggest that E. annulipes is a potential biological control agent for C. capitata, and that, because the earwig avoids consuming larvae parasitized by D. longicaudata, a combination of the two natural enemies could have an additive effect on pest mortality.     

The mutual affinities of the major earwig taxa(Insecta, Dermaptera)

Giles ' (1974) claims that the Hemimerina have closer affinities with the Forficulina-Arixenüna complex than with any other orthopteroid order is confirmed by detailed studies of his own data. The Arixenüna are shown to have close affinities with the Labüdae within the Forficulina. The Forficulina and the Hemimerina have more distant affinities than most previous authors have suggested. The Forficulina is composed of two sister groups, which are each given infra-ordinal status, namely the Pygidicranidea for the pygidicranid earwigs and the Forficulidea for the rest. The Forficulidea is seen to be composed of two mutual sister groups at super-family level, namely the Labioidea for the Carcinophoridae, Labüdae and the Arixenüdae and the Forficuloidea for the Labiduridae (sensu stricto), the Chelisochidae and the Forficulidae. Within the Labioidea, the Labüdae and the Arixenüdae are seen as possible sister groups. The classification of the lower taxonomic levels along Hennigian lines is a matter for further investigation.     

Maternal behavior and clutch manipulation in the ring-legged earwig (Dermaptera: Carcinophoridae)

The expression and maintenance of maternal behavior in the earwig,Euborellia annulipes, was examined through manipulation of clutch size, age, and species and through observations of interactions between brooding females. Females underwent discrete gonadotrophic cycles culminating in oviposition of first clutches that were highly variable in size. Neither the head capsule width nor the age of the mother was correlated with clutch size. Maternal care extended through embryogenesis and for the week following hatching. Clutch removal significantly shortened the interclutch interval, indicating that the presence of brood inhibited the onset of the second gonadotrophic cycle. Brooding females readily accepted replacement clutches of the same age. Thus, mothers did not appear to distinguish their own eggs from those of other females. Experimental doubling of clutch size did not significantly reduce the proportion hatching or fledging. In contrast, reducing clutch size diminished the percentage successfully fledging. Manipulation of clutch age resulted in reduced hatching/fledging success. Placing two females, each with newly laid clutches, in the same cage usually resulted in egg transfer from the nest of one female to that of the other within 12 h. Nests of females with larger forceps were significantly more likely to contain both clutches. When mothers with first clutches were paired with mothers with third clutches, eggs were more likely to be transferred to the nest of the older female.E. annulipes females with newly laid clutches appeared to accept as replacement clutches eggs of the earwigDoru taeniatum. Alien clutches were maintained for the typical duration of embryogenesis; however, noD. taeniatum hatchlings were observed.     

Zur biologie und morphologie von Oecobius annulipes lucas (Araneae,Oecobiidae)

The biology of various species of the cribellate family Oecobiidae has been known only fragmentarily. Therefore observations on web construction, catching and mating behaviour of Oecobius annulipes are made. They are paralleled by morphological investigations on the mouth parts and the spinning apparatus. The results are in closest conformity with the corresponding characters of the ecribellate spider Uroctea durandi. A close phylogenetic relationship of both families is assumed.     

Ultrastructural changes in the corpus allatum after azadirachtin and 20-hydroxyecdysone treatment in adult females of Labidura riparia (Dermaptera)

In previous reports, we have shown that the injection of azadirachtin (AZA) as well as 20-hydroxyecdysone (20E) into vitellogenic females of Labidura riparia induces inhibition of vitellogenin synthesis and ovarian development. Juvenile hormone (JH) treatment rescues vitellogenin synthesis and ovarian growth (Sayah et al., 1995, 1996). In this work, we have studied ultrastructural changes of corpus allatum (CA) after injection of 200, 400, and 600 ng of 20E or 1, 3, and 5 microg of AZA. CA cells exhibit signs of inactivity in both AZA and females treated with 20E at doses of 3 microg and 400 ng, respectively. Conspicuous cytological effects consisting of multivesicular bodies with dense contents, abnormally large intercellular spaces comprising myelinic structures, and rare smooth endoplasmic reticula occurred in cytoplasm of CA glandular cells in both experimental females. However, the CA ultrastructure of females injected with 20E differs from CA cells of females injected with AZA in having a cytoplasm containing numerous electron-lucent intracellular areas and marked glycogen zones. They also differ in having abundant microtubules and well-developed junctional membranes. At a dose of 600 ng of 20E or 5 microg of AZA, the intensity of the cytotoxic effects is more apparent. CA cells display pycnotic nuclei, spherical mitochondria, large multivesicular bodies, and vacuolization of the cytoplasm. These results are discussed and compared with observations made on other insect species.     

Artificial diet for continuous maintenance of the maritima earwig Anisolabis maritima (Bonelli) (Dermaptera: Carcinophoridae)

Anisolabis maritima is an important predator for the eggs of the red palm weevil Rhynchophorous ferrugineus. It could be successfully reared in the laboratory, on an artificial diet composed of dry kidney beans, Brewers yeast, chicken, egg yolk, agar, ascorbic acid, mould inhibitors, vitamin B₁₂, folic acid and riboflavin.     

Ultra morphology of sensilla on the head and mouth parts of the earwig Anisolabis martima (Bonelli) (Dermaptera: Carcinophoridae)

The earwig Anisolabis maritima is a cosmopolitan insect with a worldwide distribution, omnivorous in feeding, eating the eggs of different insects. Recently recorded in Saudi Arabia as a beneficial predator on eggs and newly hatched larvae of the red palm weevil Rhyncophrous ferrugineus, scanning electron microscopy (SEM) observations revealed the presence of different kinds of receptor sensillae occurring on the head and mouth parts of A. maritime. The sensillae are the same, but differ in numbers on both sexes. The head surface bears three subtypes of trichoid sensillae, one type of ceoloconica. Labrum, labium, labial palps and maxillary palps, possess different kinds of sensillae. Three subtypes of trichoid have been identified on the labrum. The labium contained one type of tricoid and basiconic sensillae. There are great numbers and numerous sensillae cheatica differing in length distributed on the maxillary and the labial palps on all segments between other stutter spines. The last terminal segment of maxillary or labial palps bears the greatest number of the sensillae cheatica. Adult A. maritima may be introduced in the future as a beneficial predator of eggs of R. ferrugineus as a tool for integrated pest management.