Distribution of Empoasca devastans and its egg parasites in the Indian Union

The cotton jassid Empoasca devastans Distant has been reported as a major pest from all the cotton growing areas of the Indian Union. Its outbreaks have always been a difficult problem for the growers of cotton in the Punjab, and it has often resulted in the complete failure of this crop (Lal, 1941). This jassid also attacks other crops and ornamental plants (Sohi, 1964). A systematic survey of various stages of this jassid was undertaken in the Indian Union by collecting the natural parasites and predators during the period May 1964 to September 1966. We have restricted the survey to the following crops: okra (Abelmoschus esculentus), castor (Ricinus communis), eggplant (Solanum melongena), cotton (Gossypium spp.) and cowpea (Vigna catjang). Five mymarid and one trichogrammatid parasites and four predators were found and recorded (Subba Rao et al., 1965; Subba Rao, 1966), and a new species of egg parasite was described. The material collected from different areas of the survey (Fig. 1) was kept in the laboratory in 15 × 20 cm jars. Any parasites hatching from the material were preserved in 70% alcohol. In addition an assessment of the % parasitism of the eggs of this jassid at Delhi was made for one year from August 1965 to July 1966 on the two seasonal crops, okra and eggplant, and on castor which is available throughout the year. Leaves gathered on the fields of the Institute were dissected in search of the jassid eggs. The visibly parasitised eggs were sorted out and counted.     

Parasitism of orangestriped oakworm (Lepidoptera: Saturniidae) eggs in the urban landscape

Orangestriped oakworm, Anisota senatoria (J. E. Smith), has caused widespread defoliation of oak trees in the urban landscape of southeastern Virginia since 1985. Egg masses were collected from 1988 to 1990 to determine the impact of native egg parasites on A. senatoria populations. The most abundant egg parasite was Aprostocetus new sp. and mean egg mass parasitism was 24.6%. The eupelmid Anastatus hirtus (Ashmead), a new host record, parasitized a mean of 11.7% of A. senatoria egg masses. The encyrtid Ooencytrus sp., a new host record, had a mean egg mass parasitism of 0.09%. Inundative releases of Trichogramma minutum (Riley) in 1989 and 1990 did not increase parasitism rates and mean egg mass parasitism was 2.3%. Parasitism of first generation A. senatoria egg masses was higher compared with second generation. The four egg parasites collected in this study parasitized 30% of A. senatoria egg masses and within egg mass parasitism was 7.9%. These relatively low parasitism rates may partially explain the presence of consistently high A. senatoria populations in southeastern Virginia.     

Egg sac damage and previous egg sac production influence truncated parental investment in the wolf spider,Pardosa milvina

Life history theory predicts that iteroparous animals adaptively partition reproductive effort between current and future reproduction. When rearing costs of current offspring exceed the potential benefits, parental care should be terminated and deferred toward future reproduction. We tested two related predictions that follow from life history theory: (a) parents should be sensitive to offspring viability and withhold parental care if offspring survival probability drops and future reproductive opportunities are likely, and (b) parents should be less sensitive to offspring survival probability when future reproduction is unlikely and maximize parental care late in life. The wolf spider, Pardosa milvina, demonstrates extensive parental care; however, they may also abandon or cannibalize their egg sacs. We tested the effects of egg sac damage and production of a previous egg sac on egg sac abandonment and cannibalism decisions. Among four egg sac groups (1st egg sac intact, 1st egg sac damaged, 2nd egg sac intact, 2nd egg sac damaged), we daily monitored egg sac abandonment and cannibalism and measured differences in egg sac searching, protection, and grooming among removed and damaged egg sacs (N = 116 with 1st egg sac and 88 with 2nd egg sac). Females with first egg sacs abandoned and cannibalized damaged egg sacs significantly more compared to unmanipulated egg sacs; however, females with second egg sacs were insensitive to egg sac damage. Females also spent significantly more time protecting second egg sacs compared to first egg sacs and groomed damaged egg sacs significantly more than undamaged. These results support the general predictions of life history theory that indicate that abandonment and cannibalism should decrease with diminished future reproductive potential and that parents should be less sensitive to indicators of offspring survival probability late in life.     

Costs and Benefits of Maternal Care in a Subsocial Spider,Coelotes terrestris

The maternal social spider Coelotes terrestris demonstrates extended care towards its progeny: the mother guards its egg sac for 3–4 weeks, then stays with its young from the time of their emergence until their dispersal about 1 month later. The present investigation evaluates the adaptiveness of these maternal behaviours by comparing the fitness of females performing them with that of females separated from the egg sac or the spiderlings. By protecting their egg sacs from predation and parasites, and by pursuing this task while supplying the young with food, mothers enhance the survival rate and the development of many of their spiderlings. The costs linked with these activities, estimated by the ability to produce another clutch, appear variable according to the stage in the reproductive cycle. In such terms, the egg sac guarding appears to have a low cost in relation to the care given to the spiderlings.     

Trichogramma egg parasitism ofHelicoverpa armigera on short-duration pigeonpea intercultured with sorghum

The levels of egg parasitism byTrichogramma spp. (Hymenoptera: Trichogrammatidae) on populations ofHelicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) were recorded on sorghum (Sorghum bicolor L.) and on two flushes of flowers on short-duration pigeonpea (Cajanus cajan L.).H. armigera oviposition was concentrated on the early flowering stage of sorghum and the flowering and early podding stages of both flushes of pigeonpea. Parasitism on sorghum increased rapidly as egg density increased and reached a peak of 74.6%. Parasitism on pigeonpea was concentrated onH. armigera eggs laid on the first flush of pigeonpea flowers with a maximum of 69.2%. These high levels of parasitism on pigeonpea coincided with the period of parasite activity on sorghum. The levels of parasitism then declined rapidly and only very low levels were detected on a second flush of flowers. This rapid decline resulted in the overall egg mortality caused byTrichogramma on pigeonpea to be low, with a maximum of 7.8% caused by parasitism, compared to 34.4% on sorghum. The pattern of parasitism suggests that a transfer of parasites occurred from sorghum to pigeonpea. The rapid decline of parasitism on the pigeonpea indicates that parasite populations cannot be sustained on pigeonpea once the influx from sorghum stops. The results are discussed in terms of a possible method of encouraging the transfer of parasites from sorghum to short-duration pigeonpea by producing a more continuous cropping environment.     

Studies on the reproduction of cirripedes. IV. The protein amino-acid composition and gel electrophoresis of the oviducal sac and egg case

The oviducal sac of Pollicipes cornucopia Leach and of Balanus balanoides(L.) and also the egg case of the latter species are almost pure protein: the ash of B. balanoides egg cases contains, however, a considerable quantity of iron. The amino-acid composition of the proteins has been determined; that of the sacs is quite different in the two species. In spite of the different way in which the sacs were obtained it is considered that these differences are specific in origin. This is in accordance with probably specific differences in ovisacases, and the behaviour of the egg lamella of different species towards attack by protease.The behaviour of the proteins of the sac of Pollicipes cornucopia on SDS polyacrylamide gel electrophoresis indicates a variable polypeptide composition according to the ‘age’ of the sac; with increasing age there is a greater degree of polymerization.The results are discussed in relation to the mode of release of ovisacase, its passage across the egg case, activation, and the attack on only one envelope.     

Synchronization of laying by great spotted cuckoos and recognition ability of magpies

Brood parasites rely entirely on the parental care of host species to raise the parasitic nestlings until independence. The reproductive success of avian brood parasites depends on finding host nests at a suitable stage (i.e. during egg laying) for parasitism and weakly defensive (i.e. non‐ejector) hosts. Finding appropriate nests for parasitism may, however, vary depending on ecological conditions, including parasite abundance in the area, which also varies from one year to another and therefore may influence coevolutionary relationships between brood parasites and their hosts. In this scenario, we explored: 1) the degree of laying synchronization between great spotted cuckoos Clamator glandarius and magpies Pica pica during two breeding seasons, which varied in the level of selection pressure due to brood parasitism (i.e. parasitism rate); 2) magpie responses to natural parasitism in the pre‐laying period and successfulness of parasitic eggs laid at this stage; and 3) magpie responses to experimental parasitism performed at different breeding stages. We found that, during the year of higher parasitism rate, there was an increase in the percentage of parasitic eggs laid before magpies started laying. However, the synchronization of laying was poor both years regardless of the differences in the parasitism rate. The ejection rate was significantly higher during the pre‐egg‐laying and the post‐hatching stages than during the laying stage, and hatching success of parasitic eggs laid during the pre‐egg‐laying stage was zero. Thus, non‐synchronized parasitic eggs are wasted and therefore poor synchronization should be penalized by natural selection. We discuss four different hypotheses explaining poor synchronization.     

Behavior and mortality of the southern stink bugNezara viridula in Hawaii

This paper is concerned with some aspects of the ecology of the southern green stink bug, Nezara viridula, in Hawaii. Dispersal studies on adults showed an asymetrical dispersal pattern. The rate of plant to plant to movement by the adults was 3.3–6.7 feet per day. Mortality studies showed that egg predation by ants, Pheidole megancephala, was as high as 80–90 per cent. Egg parasitism by Telenomus basalis was highest during February, 1965, and adult parasitism by Trichopoda pennipes was highest during July, 1964, to January of the following year. The population of N. viridula declined during May, 1964, to May, 1965. This decline occurred simultaneously with the decline in the number of eggs produced per female. It has been speculated that this reduction in egg production was caused by parasitism by T. pennipes.     

Differences in egg parasitism of Chrysophtharta agricola (Chapuis) (Coleoptera: Chrysomelidae) by Enoggera nassaui Girault (Hymenoptera: Pteromalidae) in relation to host and parasitoid origin

Abstract The first instances of egg parasitism of Chrysophtharta agricola , a pest of eucalypt plantations, are recorded. Enoggera nassaui was found parasitising C. agricola egg batches in Tasmania, the Australian Capital Territory (ACT), New South Wales and Victoria: this is the first record of this parasitoid species from Victoria. One instance of Neopolycystus sp. parasitising C. agricola eggs in Victoria was also recorded. Parasitism of egg batches by E. nassaui ranged from 0 to 55% between five geographical populations collected in mainland Australia ( n  = 45), and from 0 to 2% between two populations collected in Tasmania ( n  = 300). For mainland sites at which parasitism was recorded, parasitism rates within sites differed significantly from either population in Tasmania. Reciprocal exposure experiments using one Tasmanian (Florentine Valley) and one parasitised mainland (Picadilly Circus, ACT) population were conducted in the laboratory to examine whether these different parasitism rates were attributable to egg or parasitoid origin. Parasitoids from the ACT parasitised C. agricola eggs of both origins more successfully than parasitoids from Tasmania, with up to 65% wasp emergence compared with 33% from Tasmania. Parasitoid origin significantly affected the number of wasps that emerged from exposed batches, but not the total loss from parasitism.     

Behaviour of colonial and solitary spiders of the theridiid species Anelosimus eximius

Most Anelosimus eximius live in colonies, but a few females emigrate short distances and establish small, individual webs. Field studies were conducted on one colony and nearby smaller webs in order to describe communal activities and division of labour, and to note costs and benefits of remaining in the parental colony and emigrating. Adult and juvenile females repaired the web and captured prey. Adult females, rather than juveniles, cared for egg sacs and fed young. When colonial egg sacs were abundant, females moved from sac to sac. Several colonial females regurgitated food to spiderlings which had recently emerged from a particular sac. In smaller webs inhabited by two or three females, the mother cared for the sac but all females fed the young by regurgitation. Males rarely participated in communal activities. Advantages of colony living include protection from predators, the availability of large prey and, in the event of a female's death, the care of her egg sac and feeding of her young. Cannibalism of the egg sac is a potential cost of coloniality, affecting about 10–20% of colonial sacs. Cannibalism was not observed in the smaller webs. However, the costs of emigrating are very high: most of the solitary females disappeared, leading to interspecific predation on their eggs and young.     

Energy flow from spider eggs through dipteran parasite and hymenopteran hyperparasite populations

Summary The incidence of the parasites in the egg sacs of the spider Argiope pulchella was 100% for Sarcophaga banksi and 25% for Tachinobia repanda. In 1976, 237 spider eggs equivalent to 131 gcal/m² were present. Of these, 212 eggs/m² (=117 gcal) were consumed by S. banksi larvae, leaving 25 spiderlings/m² (=14 gcal) to emerge. The density of S. banksi larvae was l/m² (=87 gcal), of which 0.7/m² S. banksi (=41 gcal) successfully emerged; as few as 0.3 S. banksi/m ² (=25 gcal) were infected by T. repanda; only 14 T. repanda/m² (=8 gcal) successfully emerged. Exploitation efficiency was 89% for S. banksi and 29% for T. repanda. Ecological efficiency was 66% for S. banksi and only 9% for T. repanda. The egg sac area of A. pulchella holds a straight line relationship to the energy content of the eggs; the sacs were grouped into 8 different sizes and each one further into groups containing 1, 2, and 3 S. banksi larvae per sac. Analysis of the sacs at the appropriate time revealed that an S. banksi larva consumed a minimum of 114 eggs (=70 gcal), when present as one of a pair in the smallest sac (0.6 cm² area), and a maximum of 476 eggs (=234 gcal), when present alone in the largest sac (1.3 cm²). Despite this wide difference in food intake, all S. banksi (barring those infected by T. repanda) successfully emerged. The energy content of a pharate pupa, which was 125, 92, and 68 gcal in a sac with 1 cm² area containing 1, 2, or 3 S. banksi, depended on the size of the sac and the number of S. banksi per sac. The corresponding values for the imago were 82, 61, and 45 gcal. The efficiency of S. banksi ranged between 60 and 80% for food conversion and between 35 and 56% for pupation.This work was completed at our Palni Centre     

Fertilization in maize indeterminate gametophyte1 mutant

Summary. Mature embryo sacs of the maize mutant indeterminate gametophyte1 displayed different cellular patterns compared to those of the wild type. About 40% of the ig1 embryo sacs contained three or more synergids and two or more egg cells at the micropylar end. During fertilization in embryo sacs with two synergids, both of them frequently degenerated and were penetrated by two pollen tubes. 75% of the embryo sacs containing three or more synergid cells were penetrated by two or more pollen tubes, although most of them had only one degenerated synergid. Multiple fusions between the sperm cells and eggs frequently occurred in the same embryo sac, which subsequently generated multiple embryos. There were two or more central cells in about 33% of ig1 embryo sacs. The largest central cell was usually adjacent to the egg apparatus and contained two unfused polar nuclei, while those extra central cells located at the chalazal end usually had a single nucleus. Fertilization occurred only between the male gamete and the largest binucleate central cell. The extra central cells eventually degenerated after fertilization.Present address: GI Basic Research Center, Mayo Clinic, Rochester, Minnesota, U.S.A.Correspondence and reprints: State Key Laboratory of Plant Physiology and Biochemistry, College of Biological Science, China Agricultural University, Beijing 100094, Peoples Republic of China.     

Egg rejection and clutch phenotype variation in the plain prinia Prinia inornata

Avian hosts of brood parasites can evolve anti‐parasitic defenses to recognize and reject foreign eggs from their nests. Theory predicts that higher inter‐clutch and lower intra‐clutch variation in egg appearance facilitates hosts to detect parasitic eggs as egg‐rejection mainly depends on the appearance of the egg. Therefore, we predict that egg patterns and rejection rates will differ when hosts face different intensity of cuckoo parasitism. We tested this prediction in two populations of the plain prinia Prinia inornata: Guangxi in mainland China with high diversity and density of cuckoo species, and Taiwan where there is only one breeding cuckoo species, the oriental cuckoo Cuculus optatus. As expected, egg patterns were similar within clutches but different among clutches (polymorphic eggs) in the mainland population, while the island population produced more uniform egg morphs. Furthermore, the mainland population showed a high rate of egg rejection, while the island population exhibited dramatically reduced egg grasp‐rejection ability in the absence of parasitism by the common cuckoo Cuculus canorus. Our study suggests that prinias show lower intra‐clutch consistency in egg colour and lose egg‐rejecting ability under relaxed selection pressure from brood parasitism.     

Isolation of embryo sacs from Dianthus ovules by enzymatic treatments and microdissection

 Mature ovules of Dianthus (Caryophyllaceae) were histologically observed by clearing and serial sectioning to characterize the cells of the embryo sac. The results show that the mature embryo sac was located deep inside the hemitropous ovule due to thick nucellar tissue at the micropylar region. For the isolation of the embryo sacs, ovules were collected from ovaries of flowers 1 day after anthesis, and treated with an enzyme solution for digesting cell walls on a gyratory shaker. After 12 h of enzyme treatment, these ovules were dissected using a glass needle under an inverted microscope to release the embryo sacs. The embryo sacs, characterized by their specific size, were successfully released by these successive treatments. The viability of the embryo sacs was more than 80% as assessed with fluorescein diacetate staining. Fluorescent staining with 4,6-diamidino-2-phenylindole revealed the nuclei of the egg apparatus in the isolated embryo sacs. The procedure for isolating embryo sacs established in this study will offer a new approach to further in vitro studies on fertilization in Dianthus. Received: 20 January 1999 / Revision received: 12 July 1999 / Accepted: 17 August 1999     

Parasites of the spotted stalk borer,Chilo partellus [Lepidoptera: Pyralidae] in South Africa

One egg parasite, 7 larval parasites, 2 pupal parasites and 3 larval hyperparasites were recorded parasitizing the spotted stalk borer,Chilo partellus (Swinhoe) on maize and grain sorghum in South Africa.Trichogrammatoidea lutea Girault [Trichogrammatidae] parasitized eggs ofC. partellus mainly in mid-summer. The larval parasites were active throughout the season with occasional peaks of up to 75% parasitism.Apanteles sesamiae Cameron [Braconidae], proved to be the most abundant larval parasite. It was recorded fromca. 93% of parasitized larvae but its efficiency was reduced by the hyperparasite,Aphanogmus fijiensis (Ferriére) [Ceraphronidae], which reached sometimes up to 100% parasitism on cocoons ofA. sesamiae. The efficiency ofIphiaulax sp. [Braconidae], the 2^nd most abundant larval parasite, was also hindered by the hyperparasite,Eurytoma sp. [Eurytomidae] Pupal parasites were sometimes very abundant reaching up to 100% parasitism without any interference by hyperparasites. The most abundant pupal parasites wereDentichasmias busseolae Heinrich [Ichneumonidae] andPediobius furvus (Gahan) [Eulophidae].      

Size-dependent survivorship in the web-building spiderAgelena limbata

Summary Stage-specific mortality rates and mortality factors for the web-building spiderAgelena limbata, which is suggested to be food-limited, were studied, and the relationship between body size of spiders and survivorship for instar 3 to adults was examined. The mortality rate of the egg sac stage including eggs, deutova (prenymphal stage), and overwintering instar 1 nymphs was low. The low mortality of this stage was partly due to maternal care that reduced the mortality caused by predation and/or abiotic factors. From emergence of instar 1 nymphs from egg sacs to reproduction, the stagespecific mortality rates were almost constant, 32–47%, and the time-specific mortality rates were also constant. These results suggest a Deevey (1947) type II survivorship curve inA. limbata, in contrast to other reports on the wandering or burrowing spiders which suggested type III curves. Important mortality factors for nymphs and adults were parasitism by an ichneumonid wasp and predation by spiders. There were great variations in body size (carapace width) ofA. limbata in the field. Smaller individuals survived at a lower rate to the next stage than larger individuals. This tendency was clearer for the population living under poorer prey availability.A. limbata was unlikely to starve to death in the field because every stage ofA. limbata could survive starvation for a long time in the laboratory, 22–65 days on average. I suggest that the size-dependent survivorship of this spider is associated with vulnerability of smaller individuals to parasitism and predation.     

Hatching from cysts and egg sacs of Heterodera cruciferae and effects of temperature on hatching and development on oilseed rape

Juveniles hatched readily from field cysts and very readily from eggs in egg sacs of Heterodera cruciferae, when exposed to oilseed rape root diffusate. They hatched very poorly, however, from white or brown females from which the above egg sacs had been removed. Some hatching occurred at 8 ^oC but much more occurred at 12 , 16, 20 and 24 ^oC, with most at 16 ^oC. Development of juveniles in roots of oilseed rape occurred throughout the range 8–24 ^oC, and proceeded faster the higher the temperature. The basal development temperature was taken as 5 ^oC and the number of day degrees above this temperature required to reach each stage of development was calculated. From invasion of roots to the hatching of F₁ juveniles required an average of 680 day degrees, but only 210 day degrees were required for the first appearance of egg sacs on adult females. On this basis, two consecutive generations of H, cruciferae would be possible on autumn-sown oilseed rape in southern England, but the second would mature fully only after the crop was harvested. In Scotland, two consecutive generations could also occur but the second would be much less mature by harvest: only about 850 day degrees are available compared to almost 1100 in southern England. In practice, however, overlapping generations probably occur due to flushes of hatching of juveniles (i) at sowing, (ii) when soils warm up after winter and (iii) when the first generation completes its development. The proportion of eggs found in egg sacs was never more than 37% and some field cysts contained about 220 eggs; their egg sacs may, therefore, have contained as many as 150 eggs. Any study of population dynamics or damage assessment will require a quantification of the contribution of eggs in egg sacs to population density. Oilseed rape is direct drilled and may, therefore, be more sensitive to a given population density of the nematode than host crops which are transplanted.     

How can distinct egg polymorphism be maintained in the rufescent prinia (Prinia rufescens)–plaintive cuckoo (Cacomantis merulinus) interaction—a modeling approach

In avian brood parasitism, both the host and the parasite are expected to develop various conflicting adaptations; hosts develop a defense against parasitism, such as an ability to recognize and reject parasitic eggs that look unlike their own, while parasites evolve egg mimicry to counter this host defense. Hosts may further evolve to generate various egg phenotypes that are not mimicked by parasites. Difference in egg phenotype critically affects the successful reproduction of hosts and parasites. Recent studies have shown that clear polymorphism in egg phenotype is observed in several host–parasite interactions, which suggests that egg polymorphism may be a more universal phenomenon than previously thought. We examined the mechanism for maintaining egg polymorphism in the rufescent prinia (Prinia rufescens) that is parasitized by the plaintive cuckoo (Cacomantis merulinus) from a theoretical viewpoint based on a mathematical model. The prinia has four distinct egg phenotypes: immaculate white, immaculate blue, white with spots, and blue with spots. Only two egg phenotypes, white with spots and blue with spots, are found in the cuckoo population. We show that the observed prinia and cuckoo phenotypes cannot be at an equilibrium and that egg polymorphism can be maintained either at stationary equilibrium or with dynamic, frequency oscillations, depending on the mutation rates of the background color and spottiness. Long‐term monitoring of the prinia–cuckoo interaction over a wide geographic range is needed to test the results of the model analyses.     

Natural occurrence of fungal egg parasites of root-knot nematodes, Meloidogyne spp. in organic and integrated vegetable production systems in Spain

A survey was conducted to assess the biodiversity and frequency of infection of fungal egg parasites of Meloidogyne spp. and relate results to soil properties in organic and integrated vegetable production in Spain. Forty sites were sampled at the end of the cropping cycle, 30 under integrated and ten under organic production. Fungal egg parasites were isolated from all organically managed sites and from 73 % sites under integrated production. Species richness and Shannon–Wiener index did not differ between production systems but the percentage of fungal egg parasitism did, as well as soil properties. Percentages of egg parasitism higher than 40 % were found in five and three sites under organic and integrated production, respectively. In all these sites, Pochonia chlamydosporia was present alone or co-occurring with other fungal species. The relative frequency of P. chlamydosporia was positively related to the percentage of parasitism in both production systems.     

REED WARBLER HOSTS FINE‐TUNE THEIR DEFENSES TO TRACK THREE DECADES OF CUCKOO DECLINE

Interactions between avian hosts and brood parasites can provide a model for how animals adapt to a changing world. Reed warbler (Acrocephalus scirpaceus) hosts employ costly defenses to combat parasitism by common cuckoos (Cuculus canorus). During the past three decades cuckoos have declined markedly across England, reducing parasitism at our study site (Wicken Fen) from 24% of reed warbler nests in 1985 to 1% in 2012. Here we show with experiments that host mobbing and egg rejection defenses have tracked this decline in local parasitism risk: the proportion of reed warbler pairs mobbing adult cuckoos (assessed by responses to cuckoo mounts and models) has declined from 90% to 38%, and the proportion rejecting nonmimetic cuckoo eggs (assessed by responses to model eggs) has declined from 61% to 11%. This is despite no change in response to other nest enemies or mimetic model eggs. Individual variation in both defenses is predicted by parasitism risk during the host's egg‐laying period. Furthermore, the response of our study population to temporal variation in parasitism risk can also explain spatial variation in egg rejection behavior in other populations across Europe. We suggest that spatial and temporal variation in parasitism risk has led to the evolution of plasticity in reed warbler defenses.