Cales (Hymenoptera: Chalcidoidea): morphology of an enigmatic taxon with a review of species

Calesinae is a small group of Chalcidoidea (Hymenoptera) that are parasitoids of whiteflies (Hemiptera: Aleyrodidae). One species, Cales noacki Howard, has been introduced from South America into citrus‐growing regions of North America, the Mediterranean and Africa for biological control. The remaining species are found in Australia and New Zealand: a classic Gondwanan disjunction. The subfamily consists of a single genus, Cales, which is currently unplaced within Chalcidoidea. Its taxonomic position has historically been unstable, although most often Cales is associated with Aphelinidae. Here, we present a detailed morphological study of the group with an emphasis on Australian species. Although Cales shares many characteristics with Aphelinidae, especially Coccophaginae and Eretmocerus, more studies of character systems across Chalcidoidea are needed to determine which features may be synapomorphic. Consequently, we leave Cales incertae sedis within Chalcidoidea. We also describe a new species from New Zealand, Cales berryi sp.n. , reared from the whitefly Asterochiton pittospori on lemonwood, Pittosporum eugenioides, and we present a key and review the four known species of Cales.     

Parasitoids of the African wild silkmoth,Gonometa postica (Lepidoptera: Lasiocampidae) in the Mwingi forests,Kenya

Gonometa postica Walker produces silk of high quality, but it is affected by parasitoids attack. A study on the parasitism of G. postica larvae and pupae on host and non‐host plants were undertaken for the first and second generations, corresponding to the long (March–May) and short (October–December) rainy seasons in 2006 at six field sites, three each in the Imba and Mumoni forests of Mwingi, eastern Kenya. All freshly spun cocoons of G. postica were sampled at each site from a total of 100 trees of host plants and other non‐host plants where they have migrated before pupation. The cocoons were kept individually in fine net‐sealed plastic vials to determine percentage parasitism. Two dipterans and four hymenopteran larval–pupal parasitoids were identified from the two forests. The most common parasitoids were Palexorista sp. (Diptera: Tachinidae) and Goryphus sp. (Hymenoptera: Ichneumonidae) with parasitism ranging from 1.8 to 32.7% and 2.2 to 7.5%, respectively. Parasitism varied significantly according to host or non‐host plants, seasons and sites. This study indicates that, of the six parasitoid species recovered, only two had a significant impact in reducing the quality of the cocoons.     

Endoparasitoids of larval Anomis privata (Lepidoptera: Noctuidae), major pest of Hibiscus syriacus (Columniferae: Malvaceae)

Endoparasitoids of Anomis privata larvae include five species in three families of two orders. In this work, two species of Hymenoptera Braconidae (Cotesia sp., Microplitis sp.), one species of Ichneumonidae (Mesochorus vittator) and two species of Diptera Tachinidae (Exorista (Podotachina) sorbillans, Timavia amoena) were investigated. Of the 261 larvae of A. privata examined, 32 had a parasite, so the rate of parasitism was 12.26%. Parasitism by taxon was the highest, at 10.35% (27 individuals), in Cotesia sp. in Hymenoptera Braconidae. Parasitoids of Braconidae and Ichneumonidae were larval parasitoids. A parasitic insect of Tachinidae was a larva–pupal parasitoid. Solitary parasitoids included Microplitis sp. in Braconidae and E. sorbillans in Tachinidae. Gregarious parasitoids included Cotesia sp. in Braconidae, M. vittator in Ichneumonidae and T. amoena in Tachinidae. There was also a multiparasitoid (T. amoena) and two superparasitoids (Cotesia sp., M. vittator). A larva of A. privata sought feed even after it was parasitized every parasitoid investigated in this study, so five species of parasitoids were all koinobiont.     

Mass rearing and augmentative releases of the native parasitoid Chouioia cunea for biological control of the introduced fall webworm Hyphantria cunea in China

The fall webworm, Hyphantria cunea Drury, is an economically important introduced pest in China. A native pupal endoparasitoid, Chouioia cunea Yang (Hymenoptera: Eulophidae, Tetrastichinae) causes considerable mortality of H. cunea pupae in some areas of China. Alternative hosts of C. cunea were tested to find potential substitute hosts for mass rearing the parasitoid. Antheraea pernyi Guerin-Meneville (Lepidoptera: Saturniidae) was chosen as a surrogate host as a single pupa produced an average of 6552 wasps and a maximum of 11,256 per pupa. A mass-rearing technique and successful release program of this parasitoid is described. Parasitism by C. cunea in the areas where it was released averaged 67.74% and was usually over 80%. Other native parasitoids, such as Coccygomimus disparis (Viereck), C. parnasae (Viereck) (Hymenoptera: Ichneumonidae) and Exorita japonica Townes (Diptera: Tachnidae), caused the total average parasitism to exceed 90% on average, reaching a maximum of 96.28% in the release areas. These successful releases indicate that an introduced pest species can also be controlled by the mass-rearing and release of native parasitoids in the country of introduction.     

Ecological and morphological characteristics of endoparasitoids on Elcysma westwoodii (Vollenhoven) (Lepidoptera: Zygaenidae)

Six species of insect endoparasitoids were identified from Elcysma westwoodii, which is the most damaging lepidopteran pest of Prunus yedoensis. From Hymenoptera, two species were identified: a species in Braconidae and Charops striatus in Ichneumonidae. From Diptera, there were four species in Tachinidae: Compsilura concinnata, Exorista sp., Pales sp. and Tachinidae spp. The parasitic ratio was 4.86% (45 of 926 larvae). The hymenopterans were parasitic on 31 individuals of E. westwoodii (68.9%) and the dipterans were parasitic on 14 individuals (31.1%). It was found that parasitoids from the larvae of E. westwoodii were all either endoparasitoids or larval parasitoids. However, Exorista sp. of Tachinidae was found to be either a larval parasitoid or larval-pupal parasitoid. Additionally, all the identified parasitoids were solitary parasitoids, as only one parasite occurred in a larva of E. westwoodii. Because the larva of E. westwoodii eats and molts after it is parasitized, all the parasitoids were identified as koinobionts. There were no big differences in morphological characteristics and life histories between C. striatus and C. concinnata. However, for Exorista sp. and Pales sp., males took 3–5 days longer to emerge from their pupae and had remarkably longer body lengths than females.     

Responses of Chalcidoidea (Hymenoptera) parasitoids to invasion of the cabbage seedpod weevil (Coleoptera: Curculionidae) in western Canada

Invasion of the European weevil, Ceutorhynchus obstrictus (Marsham), was investigated through surveys of its range and population densities in Alberta and Saskatchewan from 2001 to 2005. After it was first reported in southern Alberta, C. obstrictus rapidly expanded its range and abundance. Our more recent surveys indicate that its northward expansion has slowed, but that it has continued to extend its range eastward to southcentral Saskatchewan. The distribution and abundance of parasitoids of C. obstrictus in Alberta and Saskatchewan were investigated from 2003 to 2005 by mass rearing canola pods infested with C. obstrictus larvae. Although weevil populations were not parasitized for several years immediately following its introduction to southern Alberta, a surprisingly diverse assemblage of Chalcidoidea parasitoids, comprising 12 species from four families, were recently reared from weevil-infested canola siliques in Alberta and Saskatchewan. The Chalcidoidea fauna of C. obstrictus include species with both Nearctic and Holarctic distributions, with some species having restricted host ranges and others that are more niche than taxon-specific. These Chalcidoidea species appear to have expanded their host ranges to parasitize C. obstrictus in the region. Most parasitism is attributable to Trichomalus lucidus (Walker), Chlorocytus sp., and Pteromalus sp. (Pteromalidae), and Necremnus tidius (Walker) (Eulophidae). Parasitism levels varied considerably over the three years of this study. From 2003 to 2005 increases in parasitism occurred among all four of the species dominating the parasitoid fauna of C. obstrictus, but greater increases were observed for Chlorocytus sp. and Pteromalus sp. than for T. lucidus. Parasitoid species have sometimes caused substantial levels of host mortality, although current levels are usually less than 15% for all species combined and so are not sufficient to control weevil populations. Implementing a classical biological control program for C. obstrictus by reconstructing its European natural enemy complex is being considered, but it is still uncertain whether parasitism levels by native Chalcidoidea will increase over time since considerable year-to-year variation has been found. Parasitism levels of C. obstrictus should therefore continue to be monitored to assess whether a classical biological control program should be implemented.     

Non-target parasitism of the endemic New Zealand red admiral butterfly (Bassaris gonerilla) by the introduced biological control agent Pteromalus puparum

The New Zealand red admiral butterfly, Bassaris gonerilla (F.) (Lepidoptera: Nymphalidae), has been known as a non-target host for the introduced biological control agent Pteromalus puparum (L.) (Hymenoptera: Pteromalidae) for at least 35 years, but the level of parasitism has never been quantified. Pre-imaginal mortality in B. gonerilla was assessed over the southern summer of 2000/01 at six field sites in the Christchurch area of the South Island, New Zealand. Individual eggs and larvae were identified by tagging the stem of the Urtica ferox Forst.f. plant on which they were found and the fate of these individuals was checked weekly. These data were used to construct a partial life table for B. gonerilla. Egg mortality was very high (95%), with parasitism by an unidentified Telenomus sp. Haliday (Hymenoptera: Scelionidae) causing 57% mortality. Mortality in the larval and pupal stages increased at a constant rate with age and the major mortality factor was disappearance, which was assumed to be a result of predation and dispersal of larvae. The introduced biological control agent P. puparum parasitized 14% of B. gonerilla pupae sampled. However, parasitism by another exotic parasitoid, the self-introduced Echthromorpha intricatoria (F.) (Hymenoptera: Ichneumonidae), was even higher at 26%. A survey of pupal parasitism in three regions of New Zealand (Wellington, Christchurch, and Dunedin) revealed overall parasitism levels of 67% by E. intricatoria and 8% by P. puparum, but due to the difference in emergence times of B. gonerilla and its parasitoids, these are likely to be overestimates of percent parasitism. It is concluded that P. puparum has permanently enhanced mortality in B. gonerilla, but the level of mortality is low relative to egg parasitism by Telenomus sp., larval disappearance mortality, and pupal mortality due to E. intricatoria parasitism. To determine if this level of pupal parasitism has had population effects will require more data and the development of a population model for B. gonerilla.     

A study on the emergence sequence of pupal parasitoids of the pine processionary moth,Thaumetopoea pityocampa

The emergence of pupal parasitoids from collected pupae of Thaumetopoea pityocampa (Lepidoptera: Thaumetopoeidae) was recorded over a 2-year period. The moths and the parasitoid species Villa brunnea (Diptera: Bombylidae), Phryxe caudata (Diptera: Tachinidae), Coelichneumon rudis (Hymenoptera: Ichneumonidae) emerged sequentially. The ecological implications of such parasitism are discussed.     

Sex ratio of Nysius huttoni White (Hemiptera: Lygaeidae) in field and laboratory populations

Abstract

We studied the parasitoids of three insect seed predators of the mast-seeding genus Chionochloa spp. (snow tussocks) at Mt Hutt, New Zealand. Megacraspedus calamogonus (Lepidoptera: Gelechiidae) is parasitised by four species, three Hymenoptera (Zealachertus tortriciphaga [Eulophidae], Diadegma sp. [Ichneumonidae] and Dolichogenidea sp. [Braconidae]) and one Diptera (Uclesiella sp. [Tachinidae]). Diplotoxa similis (Diptera: Chloropidae) is parasitised by Callitula sp. (Hymenoptera: Pteromalidae). Eucalyptodiplosis chionochloae (Diptera: Cecidomyiidae) is parasitised by two Hymenoptera, Gastrancistrus sp. (Pteromalidae) and Zelostemma chionochloae (Platygastridae); all three species have extended diapause. Overall parasitism was 68.5% in M. calamogonus, 1% in D. similis, and 41% in E. chionochloae. Such parasitism in M. calamogonus and E. chionochloae may reduce seed predation in Chionochloa and alter the selective benefit of mast seeding (predator satiation) to the plant. However, seed predation is still high at many sites, so some of the seed predators and parasitoids may be food-limited (bottom-up regulation).     

Survey for parasites ofSesamia calamistis (Lep.: Noctuidae) andEldana saccharina (Lep.: Pyralidae) in southwestern Nigeria

Field surveys were conducted during 1990–92 to document the relative abundance of different species of parasites of the lepidopterous stem borersSesamia calamistis Hampson andEldana saccharina Walker in maize fields in southwestern Nigeria. Species of parasitoids detected on both stem borers included the larvalpupal parasitoidsSturmiopsis parasitica Curran (Diptera: Tachinidae) andBrachymeria feae Masi (Hymenoptera: Chalcididae), and the larval parasitoidDolichogenidea polaszeki Walker (Hymenoptera: Braconidae). The braconidCotesia sesamiae (Cameron) was found attackingS. calamistis. The hyperparasitoidExoristobia dipterae (Risbec) (Hymenoptera: Encyrtidae) was detected on a pupa ofS. parasitica. Parasitic nematodes belonging toMermis sp. and/orHexamermis sp. were found infesting larvae of both stem borers. Overall, larval/pupal parasitization levels at Ibadan were low and ranged from 4.2 to 22.8% forS. calamistis and 1.2 to 13% forE. saccharina. Of the parasites found,S. parasitica was the most common, followed by nematodes. Four hymenopteran egg parasitoids were found attackingS. calamistis: Telenomus busseolae Gahan,T. isis Polaszek (Scelionidae),Lathromeris ovicida Risbec, andTrichogrammatoidea eldanae Viggiani (Trichogrammatidae). Egg parasitization ranged from 13.4 to 41.5%. The only egg parasitoid detected onE. saccharina wasTelenomus applanatus Bin and Johnson, which inflicted only 5% parasitization.     

Bionomic notes on Pachysomoides sp. (Hymenoptera: Ichneumonidae), a parasitoid of the Neotropical social wasp Polistes satan Bequaert (Hymenoptera: Vespidae)

Although most polistine wasp species are found in the Neotropical region, mainly in Brazil, only a very limited number of South American parasitoids or parasites are known to exist. We assessed the frequency of a hymenopterous parasitoid, Pachysomoides sp. (Ichneumonidae, Cryptinae), in the nests of the Brazilian independent‐founding wasp Polistes satan and compared the rates of the parasitization of P. satan by Pachysomoides sp. between the dry (winter) and wet (summer) seasons. Pachysomoides sp. larvae were seen to feed on P. satan pupa and were found in both the upper and lower parts of the host pupal cell (ca. 10 individuals in each host pupal cell). Approximately one‐third of the pupal cells in the P. satan colonies were parasitized in the dry season, whereas there were no parasitized pupal cells in the wet season. Consequently, the rates of parasitization by Pachysomoides sp. were significantly greater during the dry season than during the wet season due to unknown reasons.     

Eucharitid Parasitism of the Neotropical Ant Ectatomma tuberculatum: Parasitoid Co‐occurrence,Seasonal Variation,and Multiparasitism1

Multiparasitism is reported for the first time in Eucharitidae (Hymenoptera: Chalcidoidea). Dilocantha lachaudii, Isomerala coronata, and Kapala sp. were found simultaneously attacking the same population of Ectatomma tuberculatum (Olivier) (Hymenoptera: Formicidae: Ectatomminae) in southern Mexico. Adults of D. lachaudii and I. coronata completed development on the same individual host on three occasions. The genus Kapala is reported for the first time as a parasitoid of E. tuberculatum, and I. coronata is reported in Mexico for the first time.     

Effects of host age on the performance of >Diadromus collaris, a pupal parasitoid of >Plutella xylostella

The ichneumonid >Diadromus collaris(Gravenhorst) (Hymenoptera: Ichneumonidae) is amajor solitary, pupal endoparasitoid of thediamondback moth, >Plutella xylostella(Linnaeus) (Lepidoptera: Plutellidae).Experiments to examine parasitism of the hostpupae of different ages by the parasitoid wereconducted in the laboratory. >Diadromuscollaris preferred host pupae that were in thefirst half of their pupal development. Survivalfrom larva to adult, and size and parasitizingcapacity of the resultant female adultsdecreased dramatically as host pupal ageincreased. When ovipositions were made intohost pupae that were in the last quarter oftheir development, all parasitoids died beforeadult emergence. The performance of >D.collaris, as affected by host pupal age,agrees with the simple diet theory thatpredicts female wasps should select hosts ofhigher nutritional quality for oviposition.     

Recent advances in biological control of important native and invasive forest pests in China

In recent decades, China has suffered severe attacks by both native and invasive forest pests. We have carried out a series of research projects on biological control of these pests. The fall webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae) has been sustainably controlled by an effective gregarious pupal endoparasitoid, Chouioia cunea Yang (Chalcidoidea: Eulophidae), which is native to China, and spraying HcNPV virus against the pest’s larval stage. Pine wilt disease, caused by the pine wood nematode (Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle) (Aphelenchida: Aphelenchidae), is currently the number one pest in China. The strategy for controlling the disease is to manipulate the nematode’s vector, Monochamus alternatus Hope (Coleoptera: Cerambycidae). We discovered that Dastarcus helophoroides (Fairmaire) (Coleoptera: Bothrideridae) is the most important natural enemy in China pine forests. Mass rearing and release techniques were studied and developed. By releasing the parasitoid, 92.6% of the M. alternatus were parasitized in the first year. Meanwhile, three elaterid beetle species were found to prey on the larva of M. alternatus. The red turpentine beetle, Dendroctonus valens (LeConte) (Coleoptera:Scolytidae) was suppressed by a predator, Rhizophagus grandis Gyllenhal (Coleoptera: Rhizophagidae) introduced from Belgium and a total 3334 ha. of pine forests were protected. The oak longhorned beetle, Massicus raddei (Blessig) (Coleoptera: Cerambycidae) is the number one pest in the northeast forests of China, where it damages trunk of oaks, mainly Quercus liaotungensis and Q. mongolicus. An integrated management technique was developed for controlling the longhorned beetle: a special black light was invented for trapping the adults; the parasitoid Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae) was released against young larvae; and the parasitoid Dastarcus helophoroides eggs and/or adults were released when the hosts were mature larvae and/or pupae. By applying the technique for five years in northeastern China oak forests, the oak longhorned beetle has been controlled to a large extent. The emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is native to eastern Asia, including China, and feeding by larvae damages ash trees. Natural enemies of the emerald ash borer were investigated and seven species were found in China, of which Spathius agrili Yang (Hymenoptera: Braconidae), Tetrastichus planipennisi Yang (Hymenoptera: Eulophiae), Sclerodermus pupariae Yang et Yao and Oobius agrili Zhang et Huang (Hymenoptera: Encyrtidae) are predominant and have high potential for biocontrol of the pest. The biology, behavior, ecology and mass rearing techniques of the parasitoids were studied.     

Structure and dynamics of the parasitoid community shared by two herbivore species on different host plants

The structure of the parasitoid community on phytophagous insects can be affected by host plant properties, such as chemical compounds, trichomes, and glandular hairs. To clarify effects of host plants on herbivores and the parasitoid community, I examined the structure and dynamics of the parasitoid community associated with two species of Caloptilia moths (Lepidoptera: Gracillariidae) that feed on different Rhododendron species (Ericaceae) for 3 years in a temperate secondary forest in central Japan. Caloptilia azaleella had overlapping generations in summer and overwintered as larvae on leaves of R. macrosepalum. Caloptilia leucothoes also had overlapping generations in summer, but it did not overwinter on the deciduous shrub R. reticulatum. The parasitoid community of C. azaleella larvae and pupae was composed of 18 species, whereas that of C. leucothoes was composed of seven species. Five species of parasitoids attacked both Caloptilia species. The most abundant parasitoid, Apanteles cf. xanthostigma (Hymenoptera: Braconidae), more frequently attacked C. azaleella than C. leucothoes larvae. In contrast, another abundant parasitoid, Acrysocharoides sp. (Hymenoptera: Eulophidae), more frequently attacked C. leucothoes than C. azaleella larvae. This differential parasitism by the most abundant parasitoid species may be responsible for the differential structure and dynamics of the parasitoid community between the Caloptilia species. The host plant of C. azaleella, R. macrosepalum, more frequently trapped and killed parasitoids (of similar size to Acrysocharoides sp.) on the glandular hairs of leaves than did R. reticulatum. The differential effect of host plants on abundant parasitoids may be related to the differential parasitism by the two abundant parasitoids shared by the herbivore hosts.     

Leaf nutrient levels and the spatio-temporal distributions of <Emphasis Type="Italic">Plutella xylostella</Emphasis> and its larval parasitoids <Emphasis Type="Italic">Diadegma insulare</Emphasis> and <Emphasis Type="Italic">Microplitis plutellae</Emphasis> in canola

Seasonal distribution patterns of the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), and its principal parasitoids Diadegma insulare (Cresson) (Hymenoptera: Ichneumonidae) and Microplitis plutellae (Muesebeck) (Hymenoptera: Braconidae) were investigated over three site-years in commercial fields of canola (Brassica napus L.) in southern Alberta, Canada. The sampling of P. xylostella, D. insulare, and M. plutellae from points arranged in grid patterns, together with the mapping and analysis of their spatial distributions over time, generated a detailed picture of the pattern of crop infestation by the herbivore and its parasitoids. Plutella xylostella exhibited significant aggregations on different scales most often when its host plants were in early flowering. Diadegma insulare adults exhibited significant aggregated distributions during early flowering and distributions subsequently became more uniform as the wasps moved into the crop later in the season. However, M. plutellae distributions were aggregated in mid flowering in only one site-year. The close spatial associations between densities of D. insulare and P. xylostella indicated that host abundance was the main determinant of parasitoid distribution patterns. Spatial distributions of nutrient contents in leaf tissue and their spatial associations with the herbivore and parasitoids were also investigated. Significant spatial associations existed between certain nutrients (e.g. nitrogen, sulfur, and potassium) and P. xylostella distributions. Sulfur exhibited a positive effect on the distributions of D. insulare but not of M. plutellae. We observed similar relationships between nutrients and the distribution of P. xylostella parasitoids as for nutrients and P. xylostella, but these relationships lacked consistency and may be the results of the spatial associations between the parasitoids and their hosts. Aggregated distributions of adults and larvae of P. xylostella hold promise for spatially targeted insecticidal applications as a means for reducing the environmental impact of insecticides on nontarget and beneficial species in canola agroecosystems.     

Natural enemies for the cabbage webworm,Hellula undalis (Fabr.) (Lepidoptera: Pyralidae) in Malaysia

Four species of larval parasitoids were reared from larvae of the cabbage webworm (CWW),Hellula undalis (Fabr.) (Pyralidae: Glaphyriinae) collected from various cruciferous plants and a capparidaceous weed,Cleome rutidosperma (DC). On cabbage, only two species were recorded,viz, Bassus sp. (Braconidae) andTrathala flavoorbitalis (Cam.) (Ichneumonidae). The other parasitoids were braconids,Chelonus sp. andPhanerotoma sp. Egg or pupal parasitoids were not recorded. The parasitoids were not an important mortality factor of CWW on cabbage because they were usually present at the end of the crop season and their numbers were generally low. On the other hand, predator-exclusion experiments indicated that predators were important in determining the density of CWW on cabbage and the within-generation survival in the field. The major predator was the fire-ant,Solenopsis geminata (F.), which foraged on the prepupae and pupae.     

Native parasitoids associated with Tuta absoluta in the tomato production areas of the Spanish Mediterranean Coast

The tomato borer Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) is an invasive pest that produces significant damage to tomato crops in the Mediterranean area. Although several species of predatory bugs are successfully being used for biological control of the pest, little is known about the parasitoids that are able to exploit T. absoluta as a host. With the aim of better understanding parasitoid species richness of T. absoluta along the Mediterranean Spanish Coast, we conducted an extensive survey to determine distribution, host plants and habitats where parasitoids are present. Our results indicated that egg parasitoids are naturally scarce but that the species richness of larval/pupal parasitoids is high and includes 20 different species. Seven of these had not been previously reported as T. absoluta parasitoids. The most frequent parasitoid species recovered were Necremnus sp. nr. artynes (Walker), Stenomesius cf. japonicus (Ashmead) and Neochrysocharis formosa (Westwood) (Hymenoptera: Eulophidae).     

Seasonal changes in seaweed deposition,seaweed fly abundance,and parasitism at the pupal stage along sandy beaches in central Japan

Seasonal relationships among stranded wrack quantity, seaweed fly abundances, and parasitism at the pupal stage were studied along three sandy beaches in central Japan. The seasonal occurrence patterns of puparia of seaweed flies Coelopa frigida and Fucellia spp. generally corresponded to seaweed deposition, which peaked in May–July and October–December. Parasitoids use fly puparia in these seasons. However, the occurrence of seaweed flies and their parasitoids varied among the three sandy beaches and did not correspond to the wrack amounts. These findings suggest that populations of seaweed flies and their parasitoids are seasonally, but not spatially, regulated by bottom‐up processes. The parasitoid assemblage of fly puparia was composed of two Aleochara (Coleoptera: Staphylinidae), two Trichopria (Hymenoptera: Diapriidae), and five pteromalid species (Hymenoptera), but the rate of parasitism was less than 20% and might have had little effect on fly populations.     

Karyotypes of parasitic Hymenoptera: Diversity, evolution and taxonomic significance

Haploid chromosome numbers (n) of parasitic Hymenoptera (= traditional Parasitica + Chrysidoidea) vary from 2 to 23. However, this range can be subdivided into three intervals with n= 14–23 (less derived parasitic wasps, e.g., some Ichneumonidae and Braconidae as well as Gasteruptiidae), 8–13 (many other parasitic Hymenoptera) and 2–7 (Dryinidae, the majority of Chalcidoidea and some advanced Braconidae, e.g. Aphidiinae). The symmetric karyotype with a relatively high chromosome number (n= 14–17) and the prevalence of biarmed chromosomes must be considered as a groundplan feature of parasitic Hymenoptera. Independent reductions of chromosome numbers (n≤ 10–11) occurred in some groups of the superfamily Ichneumonoidea as well as in the common ancestor of the Proctotrupoidea sensu lato, Ceraphronoidea, Cynipoidea and Chalcidoidea. Further multiple decreases in chromosome numbers (n≤ 4–6) took place in some Braconidae, various lineages of the superfamily Chalcidoidea as well as in the family Dryinidae. Two main trends prevailed in the karyotype evolution of parasitic wasps: the reduction of chromosome numbers (mainly due to tandem fusions and less frequently due to centric ones) and karyotypic dissymmetrization (through an increase in size differentiation of chromosomes and/or in the share of acrocentrics in a chromosome set). Although karyotypic features of parasitic Hymenoptera can be used for solving taxonomic problems at various levels, this method is the most effective at the species level.