Foreign exploration for Scirtothrips perseae Nakahara (Thysanoptera: Thripidae) and associated natural enemies on avocado (Persea americana Miller)

Scirtothrips perseae Nakahara was discovered attacking avocados in California, USA, in 1996. Host plant surveys in California indicated that S. perseae has a highly restricted host range with larvae being found only on avocados, while adults were collected from 11 different plant species. As part of a management program for this pest, a “classical” biological control program was initiated and foreign exploration was conducted to delineate the home range of S. perseae, to survey for associated natural enemies and inventory other species of phytophagous thrips on avocados grown in Mexico, Guatemala, Costa Rica, the Dominican Republic, Trinidad, and Brazil. Foreign exploration efforts indicate that S. perseae occurs on avocados grown at high altitudes (>1500 m) from Uruapan in Mexico south to areas around Guatemala City in Guatemala. In Costa Rica, S. perseae is replaced by an undescribed congener as the dominant phytophagous thrips on avocados grown at high altitudes (>1300 m). No species of Scirtothrips were found on avocados in the Dominican Republic, Trinidad, or Brazil. In total, 2136 phytophagous thrips were collected and identified, representing over 47 identified species from at least 19 genera. The significance of these species records is discussed. Of collected material 4% were potential thrips biological control agents. Natural enemies were dominated by six genera of predatory thrips (Aeolothrips, Aleurodothrips, Franklinothrips, Leptothrips, Scolothrips, and Karnyothrips). One genus each of parasitoid (Ceranisus) and predatory mite (Balaustium) were found. Based on the results of our sampling techniques, prospects for the importation of thrips natural enemies for use in a “classical” biological control program in California against S. perseae are not promising.     

Inter- and intraspecific activities of compounds derived from sex pheromone glands of currant borer, Synanthedon tipuliformis (Clerck) (Lepidoptera: Sesiidae)

Gas chromatography and mass spectrometry analyses of crude sex pheromone gland extracts revealed that virgin Synanthedon tipuliformis (Clerck), currant borer (Lepidoptera: Sesiidae) females, produced 6 compounds, structurally related to sex pheromone components of clearwing moths. By comparison of retention times and mass spectra of natural products with corresponding properties of synthetic standards, these compounds were identified as: (2E,13Z)-octadeca-2,13-dien-1-yl acetate (E2,Z13-18:OAc), (3E,13Z)-octadeca-3,13-dien-1-yl acetate (E3,Z13-18:OAc), (13Z)-octadec-13-en-1-yl acetate (Z13-18:OAc), (2E,13Z)-octadeca-2,13-dien-1-ol (E2,Z13-18:OH), (13Z)-octadec-13-en-1-ol (Z13-18:OH) and octadecan-1-ol (18:OH) in the ratio 100:0.7:2.7:3.2:traces:traces. The first 3 compounds were previously known to occur in the sex pheromone gland extracts of currant borers, while the last 3 chemicals are now reported for the first time. Trapping tests carried out in the black currant field revealed that E2,Z13-18:OAc, when tested separately, attracted S. tipuliformis males, while addition of E3,Z13-18:OAc to the main component increased the effectiveness of E2,Z13-18:OAc over seven times. The attractiveness of 6 component lures did not differ significantly from the one of the binary mixture, confirming that E2,Z13-18:OAc and E3,Z13-18:OAc in the ratio100:0.7 are essential sex pheromone components of S. tipuliformis. Trapping tests carried out at the dwelling place of Synanthedon scoliaeformis (Borkhausen) (Lepidoptera: Sesiidae) revealed that, in addition to intraspecific synergistic effect, E3,Z13-18:OAc increased the specificity of the pheromone signal of S. tipuliformis, acting by intraspecific mode as an attraction antagonist against S. scoliaeformis males. By this way, it ensured the specificity of the sex attraction signal of the currant borer. Consequently, both compounds E2,Z13-18:OAc and E3,Z13-18:OAc have to be present in pheromone formulations used for monitoring and/or control of S. tipuliformis to avoid effecting non-target species. Other compounds identified from the sex pheromone gland of S. tipuliformis did not show any significant interspecific activity for males of S. scoliaeformis, however, they provide a basis to achieve specificity of a pheromone signal of S. tipuliformis and could act as attraction antagonists against other clearwing moth species which, like S. tipuliformis, employ E2,Z13-18:OAc as their sex pheromone component.     

Volatile cues used by the parasitoid, Aphytis melinus, for host location: California red scale revisited

Aphytis melinus DeBach (Hymenoptera: Aphelinidae) is a biological control agent of diaspidid scale insects. The parasitoid has a narrow host range but its hosts are polyphagous. We determined the source of volatile cues the wasp uses to locate its few host species when those hosts occur on more than one host plant species. We addressed four questions in regard to the use of volatile cues in host location of California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae): (1) Does A. melinus use volatile cues to assist in host location? (2) Are these cues innately recognized or learned? (3) Are cues produced by female California red scale, or from other sources? (4) Are the cues specific to the host or host plant? These questions were tested through the use of a Y-tube olfactometer. Female A. melinus used volatile cues to orient toward both infested and uninfested host plant material. Wasps learned these cues by associating odors from the host plant with host presence. They had no innate preferences for scale insect or host plant volatile stimuli. Contrary to previous studies, we found no evidence of orientation toward the female-produced sex pheromone of California red scale, nor to volatile cues from the attacked host stage. Wasps given experience with scale insects growing on lemon fruit subsequently oriented toward lemon and orange fruit and leaves. The scale species with which the wasp was given experience did not affect this preference. Wasps given experience with California red scale growing on squash did not orient toward infested lemon fruit. The host ranges of the parasitoid and its hosts are used to explain the adaptive value for the evolution of learned rather than fixed responses to cues used in foraging behavior.     

Blending synthetic pheromones of cerambycid beetles to develop trap lures that simultaneously attract multiple species

We evaluated attraction of cerambycid beetle species to blends of known cerambycid pheromones to determine whether such blends could be used as effective trap lures for detecting and monitoring multiple species simultaneously. Pheromone-baited traps captured 1,358 cerambycid beetles of which 1,101 (81.1%) belonged to three species in the subfamily Cerambycinae: Neoclytus acuminatus (F.), Neoclytus mucronatus (F.), and Xylotrechus colonus (F.). Beetles of these species were significantly attracted to synthetic blends that contained their pheromone components (isomers of 3-hydroxy-2-hexanone, 2,3-hexanediol, or both), despite the presence of pheromone components of different species, including other isomers of 2,3-hexanediol, (E/Z)-6,10-dimethyl-5,9-undecadien-2-yl acetate, and citral. In some cases, attraction was partially inhibited by the pheromone components of heterospecific species, whereas for N. acuminatus, attraction was completely inhibited when blends contained (2R*,3S*)-hexanediol, the racemic mixture of diastereomers of its pheromone, (2S,3S)-hexanediol. Among the remaining beetles captured were three species in the subfamily Lamiinae: Astyleiopus variegatus (Haldeman), Graphisurus fasciatus (Degeer), and Lepturges angulatus (LeConte). All three lamiine species were previously known to be attracted to (E/Z)-6,10-dimethyl-5,9-undecadien-2-yl acetate and were captured in significant numbers by blends containing that compound. Our results suggest that different types of cerambycid pheromones can be combined to create effective multispecies lures for use in surveillance programs that target exotic cerambycid species.     

Molecular and morphometric evidence for separate species of Uncinaria (Nematoda: Ancylostomatidae) in California sea lions and northern fur seals: hypothesis testing supplants verification

California sea lions (Zalophus californianus) and northern fur seals (Callorhinus ursinus) are each believed to host distinct hookworm species (Uncinaria spp.). However, a recent morphometric analysis suggested that a single species parasitizes multiple pinniped hosts, and that the observed differences are host-induced. To explore the systematics of these hookworms and test these competing hypotheses, we obtained nucleotide sequences of nuclear ribosomal DNA (D2/D3 28S, D18/D19 28S, and internal transcribed spacer [ITS] regions) from 20 individual hookworms parasitizing California sea lion and northern fur seal pups where their breeding grounds are sympatric. Five individuals from an allopatric population of California sea lions were also sampled for ITS-1 and D18/D19 28S sequences. The 28S D2/D3 sequences showed no diagnostic differences among hookworms sampled from individual sea lions and fur seals, whereas the 28S D18/D19 sequences had one derived (apomorphic) character demarcating hookworms from northern fur seals. ITS sequences were variable for 7 characters, with 4 derived (apomorphic) states in ITS-1 demarcating hookworms from California sea lions. Multivariate analysis of morphometric data also revealed significant differences between nematodes representing these 2 host-associated lineages. These results indicate that these hookworms represent 2 species that are not distributed indiscriminately between these host species, but instead exhibit host fidelity, evolving independently with each respective host species. This evolutionary approach to analyzing sequence data for species delimitation is contrasted with similarity-based methods that have been applied to numerous diagnostic studies of nematode parasites.     

Determination of the absolute configuration of quercivorol, (1S,4R)-p-menth-2-en-1-ol, an aggregation pheromone of the ambrosia beetle Platypus quercivorus (Coleoptera: Platypodidae)

A pair of enantiomers of trans-p-menth-2-en-1-ol, an aggregation pheromone of Platypus quercivorus, was synthesized from (S)- and (R)-limonene. The retention time of the aggregation pheromone from the insect coincided with that of (1S,4R)-p-menth-2-en-1-ol synthesized from (S)-limonene from GC analyses with a chiral column, enabling the absolute configuration of the aggregation pheromone to be determined as (1S,4R).     

Electroantennogram responses of the three migratory forms of the damson-hop aphid, Phorodon humuli, to aphid pheromones and plant volatiles

Electroantennogram (EAG) responses were recorded from alate fundatrigeniae (spring migrants), gynoparae (the winged female form that produces sexual females) and males, the three migratory forms of the damson-hop aphid, Phorodon humuli (Schrank). EAG responses of gynoparae and males showed typical dose response characteristics to (E)-2-hexenal, (-)-R-carvone, hexanenitrile and (1RS,4aR,7S,7aS)-nepetalactol, the sex pheromone of this species. The 34 plant volatiles elicited broadly similar EAG response profiles in the three migratory forms. Green leaf volatiles produced large responses in all forms; however, the relative order of responsiveness varied. EAG responses to isomers of the monoterpene carvone differed between forms, with males being most, and spring migrants least, responsive. The hop-plant volatile and aphid alarm pheromone, (E)-beta-farnesene, evoked similar EAG responses in all forms. By contrast, males were most responsive to the three sex pheromone components, (-)-(4aS,7S,7aR)-nepetalactol, (+)-(4aS,7S,7aR)-nepetalactone and (1RS,4aR,7S,7aS)-nepetalactol. Males were no more responsive to their own sex pheromone, (1RS,4aR,7S,7aS)-nepetalactol, than to the other aphid sex pheromone components tested. Spring migrants and gynoparae also responded to the three sex pheromone components. This study indicates that migratory forms of P. humuli detect a wide range of volatile compounds, and that they are equally well-adapted for the detection of volatiles associated with host and non-host plants and with other species of aphid.     

蚜虫性信息素

蚜虫属同翅目蚜总科,是重要的农作物害虫。蚜虫长年行孤雌生殖,在中、高纬度深秋时,产生性蚜,以卵越冬。雌性蚜分泌性信息素荆芥内酯(nepetalactone)和荆芥醇(nepetalactol)。该文综述了蚜虫性信息素的结构与组分、性腺的形态结构、性信息素释放及影响因素;性信息素对雌性母和天敌的引诱作用;性信息素在蚜虫种间隔离中的地位和作用以及蚜虫性信息素在蚜虫防治中的应用前景。     

Large-scale preparation of (+)-disparlure,the gypsy moth pheromone,by a practical chemico-enzymatic procedure

The Sharpless asymmetric epoxidation of 8-methyl-2-nonen-1-ol performed on a large scale (over 5 moles) at room temperature gave (2S, 3R)-2, 3-epoxy-8-methyl-1-nonanol with 52%ee. The produced epoxy alcohol of low optical purity was subjected to lipase-catalyzed enatioselective acylation in order to increase the optical purity up to 85%ee. Recrystalyzation of the corresponding 3, 5-dinitrobenzoate gave optically pure epoxy alcohol. (+)-Disparlure, the gypsy moth pheromone, was synthesized in two steps from the thus obtained optically pure epoxy alcohol.     

Perception of volatiles associated with sex and food by different adult forms of the black bean aphid, Aphis fabae

Abstract. Electroantennograms (EAGs) were recorded from adult male and asexual forms (winged and wingless virginoparae and gynoparae) of the black bean aphid, Aphis fabae , during stimulation with two sex pheromone components, (+)-(4a S ,7 S ,7a R )-nepetalactone and (-)-(1 R ,4a S ,7 S ,7a R )-nepetalactol, as well as six plant volatiles, i.e. ( E )-2-hexenal, ( E )-2-hexenol-1, ( Z )-3-hexenol-1, ( Z )-3-hexenyl acetate, hexanal and allyl isothiocyanate. The male antennae are 1000-10,000 times more sensitive to nepetalactol and nepetalactone than to the plant compound ( E )-2-hexenal. Besides this marked difference of EAG peak responses in males, the EAG rise and decay are slower for the sex pheromone components. Males are also much more sensitive to the sex pheromone components than asexual females. This high sensitivity correlates with a predominance of antennal secondary rhinaria, the major sites of pheromone perception in the male. However, it is the primary rhinaria on the antennae of the wingless asexual females that are responsible for pheromone perception. Male antennae are as responsive as the asexual female antennae to the plant volatiles. The specialization of the male for mate location is discussed.     

Responses of male California red scale to sex pheromone isomers

The California red scale, Aonidiella aurantii (Maskell), sex pheromone components were previously identified as two independently attractive norsesquiterpenes. The four possible optical isomers of one and the four possible geometric and optical isomers of the other were synthesized and tested for male California red scale attractiveness in field tests and greenhouse bioassays. The results showed that there was enantiomeric and geometric specificity and only 1 isomer of each component was significantly more active than the others. The active isomers were (3S,6R)-3-methyl-6-isopropenyl-9 decen-1-yl acetate and (3Z,6R)-3-methyl-6-isopropenyl-3,9-decadien-1-yl acetate. The presence of other isomers, including the synthetic analogue 3-methyl-6-isopropylidene-9-decen-1-yl acetate, had no effect on trap catch.     

Biogeography of two species of Symbiodinium (Freudenthal) inhabiting the intertidal sea anemone Anthopleura elegantissima (Brandt)

We have analyzed the genetic profiles of dinoflagellate populations obtained from the Pacific coast sea anemone Anthopleura elegantissima (Brandt) at collection sites from Washington to California. Genetic differences within the symbiont populations of California anemones have been uncovered by restriction length polymorphism (RFLP) analysis of the small subunit (SSU) and large subunit (LSU) ribosomal RNA genes, and by denaturing gradient gel electrophoresis (DGGE) of the internal transcribed spacer region 2 (ITS 2). The existence of two Symbiodinium species is substantiated by sequence analysis of the variable regions V1, V2, and V3 of the SSUrDNA, which also establishes their phylogenetic relatedness to other members of the genus Symbiodinium. Anemones from Washington and Oregon harbor a single dinoflagellate species, for which we propose the name S. muscatinei sp. nov. At these northern locations, S. muscatinei either exists alone or co-occurs with the Chlorella-like green algal symbiont. Our results indicate that S. muscatinei co-occurs with a second dinoflagellate, S. californium, in mixed populations in central and southern California. We suggest that the geographic distribution of these dinoflagellates is related to the temperature cline created by latitude.     

Discrimination of pheromone enantiomers by two pheromone binding proteins from the gypsy moth Lymantria dispar

The gypsy moth, Lymantria dispar, uses (7R, 8S)-cis-2-methyl-7, 8-epoxyoctadecane, (+)-disparlure, as a sex pheromone. The (-) enantiomer of the pheromone is a strong behavioral antagonist. Specialized sensory hairs, sensillae, on the antennae of male moths detect the pheromone. Once the pheromone enters a sensillum, the very abundant pheromone binding protein (PBP) transports the odorant to the sensory neuron. We have expressed the two PBPs found in gypsy moth antennae, PBP1 and PBP2, and we have studied the affinity of these recombinant PBPs for the enantiomers of disparlure. To study pheromone binding under equilibrium conditions, we developed and validated a binding assay. We have addressed the two major problems with hydrophobic ligands in aqueous solution: (1) concentration-dependent adsorption of the ligand on vial surfaces and (2) separation of the protein-bound ligand from the material remaining free in solution. We used this assay to demonstrate for the first time that pheromone binding to PBP is reversible and that the two PBPs from L. dispar differ in their enantiomer binding preference. PBP1 has a higher affinity for the (-) enantiomer, while PBP2 has a higher affinity for the (+) enantiomer. The PBP from the wild silk moth, Antheraea polyphemus (Apol-3) bound the disparlure enantiomers more weakly than either of the L. dispar PBPs, but Apol-3 was also able to discriminate the enantiomers. We have observed extensive aggregation of both L. dispar PBPs and an increase in pheromone binding at high (>2 microM) PBP concentrations. We present a model of disparlure binding to the two PBPs.     

Synthesis of (2S,2'R,3S,4R)-2-(2'-hydroxy-21'-methyidocosanoylamino)-1,3,4-pentadecanetriol,the ceramide sex pheromone of the female hair crab,Erimacrus isenbeckii

The ceramide sex pheromone [(2S,2'R,3S,4R)-2-(2'-hydroxy-21-methyldocosanoylamino)-1,3,4-pentadecanetriol (1)] of the female hair crab (Erimacrus isenbeckii) was synthesized by starting from (S)-serine and 12-bromo-1-dodecanol.     

Application of lipase-catalyzed transformations for the synthesis of insect pheromones and related compounds

This review describes efficient means of preparing optically pure insect pheromones and related compounds via lipase-catalyzed enantioselective reaction on a large scale. (1) A new synthesis of the Japanese beetle pheromone, (R,Z)-(−)-5-(1-decenyl)oxacyclopentan-2-one, established by a combination of two lipase-catalyzed transformation was demonstrated. (2) A chemico-enzymatic procedure for the syntheses of both enantiomers of cupreous chafer beetle pheromone, (R,Z)- and (S,Z)-5-(1-octenyl)oxacyclopentan-2-one, was described. (3) An optical resolution of (±)-2,3-epoxy-8-methyl-1-nonanol, the key intermediate of the synthesis of gypsy moth pheromone, was demonstrated. (4) A practical chemico-enzymatic synthesis of (+)-disparlure in large scale was demonstrated. (5) A facile synthesis of carboxyalkyl acrylate, which is special monomers in the synthesis of the new polymers, by two lipase-catalyzed regioselective reactions was described.     

草地贪夜蛾三个SfruPBPs对本种及同域种劳氏粘虫性信息素及腺体组分的亲和力

[目的]测定草地贪夜蛾Spodoptera frugiperda 3个信息素结合蛋白(pheromone binding protein,PBP)(SfruPBP)对草地贪夜蛾及同域近缘种劳氏粘虫Leucania loreyi性信息素及腺体组分的结合特性,探究草地贪夜蛾这3个SfruPBPs在两种昆虫不同性信息素组分识...     

Cryptic species in the cosmopolitan Bugula neritina complex (Bryozoa,Cheilostomata)

Previous analyses of the mitochondrial gene cytochrome c oxidase subunit 1 (COI) and γ‐proteobacterial endosymbiont diversity have suggested that the marine bryozoan Bugula neritina is a complex of three cryptic species, namely Types S, D and N. Types D and N were previously reported to have restricted distributions along California (western USA) and Delaware and Connecticut (eastern USA), respectively, whereas Type S is considered widespread in tropical, subtropical and temperate regions due to anthropogenic transport. Here, Bayesian species delimitation analysis of a data set composed of two mitochondrial (COI and large ribosomal RNA subunit [16S]) and two nuclear genes (dynein light chain roadblock type‐2 protein [DYN] and voltage‐dependent anion‐selective channel protein [VDAC]) demonstrated that Types S, D and N correspond to three biological species. This finding was significantly supported, in spite of the combinations of priors applied for ancestral population size and root age. Furthermore, COI sequences were used to assess the introduction patterns of the cosmopolitan Type S species. Two COI haplotypes of Type S (S1a and S1d) were found occurring at a global scale. Mantel tests showed correlation between these haplotypes and local sea surface temperature tolerance. Accordingly, the distributions of Type S haplotypes may reflect intraspecific temperature tolerance variation, in addition to the role of introduction vectors. Finally, we show that the Type N may also have been introduced widely, as this species was found for the first time in Central California and north‐eastern Australia.     

草地贪夜蛾4个性信息素结合蛋白基因的克隆及表达模式分析

草地贪夜蛾是世界性的重大害虫,2019年1月入侵我国并迅速扩散到20多个省市。性诱剂是对草地贪夜蛾进行监测和诱杀的有效手段,但是其作用识别机制仍不清楚,限制了高效性诱剂的研发和应用。性信息素结合蛋白(PBPs)在鳞翅目昆虫包括草地贪夜蛾性信息素识别过程中起重要作用。本研究从草地贪夜蛾中克隆了4个编码PBPs的cDNA序列,命名为SfruPBP1-SfruPBP4。4个SfruPBPs均含有完整的开放阅读框,所编码的蛋白具有性信息素结合蛋白的典型特征:N-端有信号肽、具有保守的6个半胱氨酸残基。系统进化分析显示SfruPBPs与斜纹夜蛾和海灰翅夜蛾PBPs的亲缘关系最近,且4个SfruPBPs被聚在不同的进化分支。4个SfruPBPs基因均由3个外显子和2个内含子组成,内含子插入位点高度保守。此外,4个SfruPBPs在草地贪夜蛾基因组上呈串联排列。SfruPBP1、SfruPBP2和SfruPBP3在成虫触角中特异性表达,其中SfruPBP1和SfruPBP2在雄成虫触角中的表达水平显著高于雌虫,而SfruPBP3在雌雄触角中的表达水平无显著差异。SfruPBP4特异性表达于雄成虫腹部。本研究结果为揭示草地贪夜蛾性信息素识别机制奠定了基础。     

Molecular identification of the exotic slug <Emphasis Type="Italic">Arion subfuscus</Emphasis> sensu stricto (Gastropoda: Pulmonata) in California,with comments on the source location of introduced populations

Arion subfuscus sensu lato (s. l.) refers to a cryptic slug species complex that is native to Europe and which, from there, has been introduced into the northeastern U.S. The species complex was detected in California for the first time in 2005 and recently 12 specimens were collected during statewide surveys. The genital morphology of these specimens suggested that only A. subfuscus sensu stricto (s. s.) was present and partial sequences of mitochondrial 16S rDNA (443 bp) showed that they all belonged to a single haplotype of the mitochondrial lineage, S1 sensu Pinceel et al. (Genetica 125: 155–171, 2005). This result was corroborated by a parallel analysis of a 655 bp COI DNA barcode. The 16S rDNA S1 haplotype (S1-03) of the Californian specimens is hitherto known only from the northeastern U.S. Hence, it is likely that populations may have been introduced to California from the northeastern states, rather than directly from the native area of the S1 haplogroup in Europe.