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Identification of Heterorhabditis

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Fundam. appl. Nemawl., 1996,19 (6),585-592 Identification of Heterorhabditis (Nenlatoda : Heterorhabditidae) from California with a new species isolated from the larvae of the ghost moth Hepialis californicus (Lepidoptera : Hepialidae) from the Bodega Bay Natural Reserve S. Patricia STOCK *, Donald STRONG ** and Scott L. GARDNER *** * C.E.PA. \I.E. - National University of La Plata, La Plata, Argentina, and Department ofNematology, University ofCalifornia, Davis, CA 95616-8668, USA, 'k* Bodega Bay lVlarine LaboratDl), University ofCalifomia, Davis, CA 95616-8668, USA and ~,M Harold W. iVlanler Laboratory ofParasitology, W-529 Nebraska Hall, University ofNebraska State Museum, University ofNebraska-Lincoln, Lincoln, NE 68588-0514, USA. Acceptee! for publication 3 November 1995. Summary - Classical taxonorny together with cross-breeding tests and random ampWied polymorphie DNA (RAPD's) were used to detect morphological and genetic variation bet\veen populations of Helerorhabdùis Poinar, 1975 from California. A new species, Helerarhabdùis hepialius sp. n., recovered from ghost moth caterpillars (Hepialis cali/amieus) in Bodega Bay, California, USA is herein described and illustrated. This is the eighth species in the genus Helerorhabdùis and is characterized by the morphology of the spicules, gubernaculum, the female's tail, and ratios E and F of the infective juveniles. Information on its bionomics is provided. Résumé - Identification d'Heterorhabditis (Nematoda : Heterorhabditidae) de Californie dont une nouvelle espèce isolée de larves d'Hepialius californicus (Lepidoptera : Hepialidae) provenant de la réserve naturelle de la baie de Bodega - Les méthodes de taxonomie classique de même que des essais de croisement et l'amplification au hasard de l'ADN polymorphique (RAPD) ont été utilisés pour détecter les différences morphologiques et génétiques entre populations d'Helerorhab­ ditis Poinar, 1975 provenant de Californie. Une nouvelle espèce, Heterarhabdùis hepialius sp. n., présente dans des chenilles d'Hepialis cali/amieus provenant de la baie de Bodega (Californie, États-Unis d'Amérique) est décrite et illustrée. Représentant la huitième espèce du genre, H. hepialius sp. n. est caractérisé par la morphologie des spicules, du gubernaculum et de la queue de la femelle ai.nsi que par les rapports E et F chez les juvéniles infestants. Des données sur sa biologie sont également fournies. Key-words : California, cross-breeding tests, Hepialidae, Hepialis cali/amieus, Heterorhabditidae, Helerorhabdùis hepialus, Lep­ idoptera, Nematoda, RAPD. Entomopathogenic nematodes of the family Hetero­ While sampling populations of pathogens of soil-in­ rhabditidae are considered effective biological control habiting insects in northem and southern California, agents of insect pests (Gaugler & Kaya, 1990). With the several Helerorhabdùis isolates were recovered from in­ increasing commercial use of these nematodes as sect hosts and soil samp1es. Material sent to the senior biological control agents, many new isolates have been author was studied and the presence of a new Helero­ recovered from soil of natural habitats in many parts of rhabdùis species was determined which is herein de­ the world, including Europe (Mracek, 1980; Burman el scribed. al., 1986; Deseo el al., 1988; Vanninen el al., 1989; Horninick & Briscoe, 1990; Griffin el al., 1991), North America (Akhurst & Brooks, 1984; Hara el al., 1991), Materials and methods Australia (Akhurst & Bedding, 1986), and South Amer­ ica (Stock, 1993). However, most of these isolates re­ ISOLATION OF NEMATODES main unidentified at the specific level. Severa1 populations of Hewrorhabdùis were recovered At present, seven species belonging to the genus Hele­ from both soil and insect hosts while conducting a sur­ rorhabditis have been described : H. bacLe1iophora Poi­ vey of soi1-dwelling insects pathogens in northern and nar, 1976; H. megidis Poinar, Jackson & Klein, 1987; H. southern California (Fig. 1). The survey was mainly ori­ zealandica (Poinar, 1990); H. indicus Poinar, Karunakar entated towards the collection ofwhite grubs (Coleopte­ & Hastings, 1992; H. argentinensis Stock, 1993; H. ha­ ran: Scarabeidae), white fringed beetles (Coleoptera: waiiensis Gardner, Stock & Kaya, 1994, and H. brevi­ Curculionidae) and ghost moth caterpillars (Lepidopte­ cal/dis Liu, 1994. ra : Hepialidae). ISSN 1164-5571/96/06 84.001 © Gaulhier-Villars - ORSrOM 585 s. P. Stock et al. Table 1. Isola les of Heterorhabctitis from Calijomia: localion, habùaI, sùe code and associalion wùh insea hoSl are indicaled. LoCltion Habitat Site Code Association \~ith inml hosts Balboa Park park AGA whire grub (San Diego) grassland (Cycio<;iphala pasadane) Bodega Bay inland edge BOD ghost moth (Sonoma) of dunes (H. califomK!4s) Chino HilJs hiIlside CHI white fringed beede (Riverside) managed landsClpe (Graphognaùml sp.) Davis baseball DAV white grub (Yolo) field (C hiTUJ) Hercules golf course HER white grub (Contra Costa) turf (C hina) Moraga golf course AGA white grub (Contra Costa) (turf) (C hiTUJ) Modesto vineyard MOD none round (Stanislaus) Oakmont golf course OAK whÎle grub (Napa) (turf) (C hiTUJ) Temecula golf course TEM none round (Riverside) (turf) Fig. 1. Colleclion sùes of Calijomian Heterorhabditis isolales. 1 : Moraga; 2: Balboa Pm'k; 3: Chino Hills; 4: Davis; 5: dissected from G. rnellonella larvae which were previous­ Hercules; 6: lvIodesto; 7: Oakmom; 8: Temecula; 9: Bodega Iy incubated at 25 oC for 6-7 days post infection with Bay. infective juveniles of the appropriate Heterorhabditis iso­ late. Insect cadavers were dissected in Ringer's solution Nematodes were recovered from both insects and using a stereomicroscope at 50 x magnification. Females soil-samples taken at different localities throughout with immature gonads were selected for the cross­ California (Table 1). breeding tests. Infected insects collected from the field were placed in Lipid agar plates (5 cm diameter) were prepared by White traps until emergence of nematodes. Soil samples the method of Dunphy and Webster (1989) and in­ were baited with Gallenà mellonella larvae as described oculated with the bacterial isolate of the appropriate by Bedding and Akhurst (1975) to test for the presence nematode strain and incubated at 30 oC for 2 days. For of entomopathogenic nematodes. each cross five virgin females and five males of the ap­ In aU cases the recovered nematodes were exposed to propriate isolate were placed on each lipid plate and fresh G. mellonella larvae to coniïrm pathogenicity and incubated at 25 oC; 3 days later an additional five males complete Koch's postulates. Infective juveniles were re­ were added to each plate. covered using White traps and harvested in a cold-room In succesful crosses, progeny were visible after at 10°C. 2-3 days. Controls were always set up for each cross; Endosymbiotic bacteria Photorhabdus luminescens Poi­ virginity/self fertility tests: five virgin females (without nar & Thomas, 1976 were isolated from the intestine of males) were placed onto lipid agar plate; mating test/self surface sterilized infective juveniles following Akhurst cross: five virgin females and five males of the same (1980) and the primary forms ofthe bacteria were main­ isolate were placed on a single lipid agar late and addi­ tained in NBTA plates (Akhurst, 1986). tional five males were added to the plate 3 days la ter. The results of the crosses between different isolates CROSS-BREEDING TESTS USING HETERORHABDiTI5 SPP. were taken as valid only if there were no progeny in the Crosses were carried out on lipid agar plates (Dix et virginity tests and if there were progeny in the self­ al., 1992). Second generation males and females were crosses. 586 Fundam. appl. Nematol. Heterorhabditis hepiaJius sp. n. RAPD ANALYSIS Table 2. Ohgonucleotide primers used in the present study (see Gardner et al., 1994). Methods of randomly amplified polymorphic DNA extraction and RAPD analysis followed Gardner el al. Reference Primer sequence (5' ra 3') (1994). OperonJS random primers (OPA-02, OPA-05, OPA-11, OPA-16, OPB-04) often bases long (Table 2) OPA-02 CAGGCCCTTC were used for aU reaction experiments with an annealing OPA-OS AGGGGTCTTG temperature of 35 oc. Purified DNA from nematode OPA-16 AGCCAGCGAA genome was subjected to PCR reaction. PCR products OPB-04 GGACTGGAGT were resolved by electrophoresis using a 1.7 % horizon­ OPB-ll GTAGACCCGT tal agarose gel, and imaged by staining with ethidium bromide. MORPHO-BIOMETRlCAL STUDIES Nematodes recovered from Galleria mellonella larvae were fixed in T AF and cleared in glycerol. Infective 2645 juveniles and first generation adults of the Helerorhab­ 1605 ditis isolate from Bodega Bay were also recovered from 1198 Hepialis caliJomicus caterpillars. Quantitative measure­ 676 ments were made using a Leitz Ortholux II microscope 517 equipped with an ocular micrometer and Jandel JAVA 398 software with a high resolution video camera. Drawings 222 were made with the aid of a drawing tube. To test the robustness of the morphometric variables, a multivariate analysis (canoninal discriminant analysis) was performed on males and infective juveniles (Stock el al.) 1995). Results CROSS-BREEDfNG TESTS According to the results obtained by the morphologi­ 2645 1605 cal and morphometric observations, the cross-breeding 1198 tests were done only between H. bacleriophora (NC 1 676 strain) and the nine Californian isolates. Crosses were 517 only carried out in Iipid agar plates. 398 From aU the possible combinations, only
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  • Small Ermine Moths Role of Pheromones in Reproductive Isolation and Speciation

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    CHAPTER THIRTEEN Small Ermine Moths Role of Pheromones in Reproductive Isolation and Speciation MARJORIE A. LIÉNARD and CHRISTER LÖFSTEDT INTRODUCTION Role of antagonists as enhancers of reproductive isolation and interspecific interactions THE EVOLUTION TOWARDS SPECIALIZED HOST-PLANT ASSOCIATIONS SUMMARY: AN EMERGING MODEL SYSTEM IN RESEARCH ON THE ROLE OF SEX PHEROMONES IN SPECIATION—TOWARD A NEW SEX PHEROMONES AND OTHER ECOLOGICAL FACTORS “SMALL ERMINE MOTH PROJECT”? INVOLVED IN REPRODUCTIVE ISOLATION Overcoming the system limitations Overview of sex-pheromone composition Possible areas of future study Temporal and behavioral niches contributing to species separation ACKNOWLEDGMENTS PHEROMONE BIOSYNTHESIS AND MODULATION REFERENCES CITED OF BLEND RATIOS MALE PHYSIOLOGICAL AND BEHAVIORAL RESPONSE Detection of pheromone and plant compounds Introduction onomic investigations were based on examination of adult morphological characters (e.g., wing-spot size and color, geni- Small ermine moths belong to the genus Yponomeuta (Ypo- talia) (Martouret 1966), which did not allow conclusive dis- nomeutidae) that comprises about 75 species distributed glob- crimination of all species, leading to recognition of the so- ally but mainly in the Palearctic region (Gershenson and called padellus-species complex (Friese 1960) which later Ulenberg 1998). These moths are a useful model to decipher proved to include five species (Wiegand 1962; Herrebout et al. the process of speciation, in particular the importance of eco- 1975; Povel 1984). logical adaptation driven by host-plant shifts and the utiliza- In the 1970s, “the small ermine moth project” was initiated tion of species-specific pheromone mating-signals as prezy- to include research on many aspects of the small ermine gotic reproductive isolating mechanisms.