Courtship behavior in the Aphytis lingnanensis group, its potential usefulness in taxonomy, and a review of sexual behavior in the
      parasitic Hymenoptera (Chalcidoidea: Aphelinidae)

Gordh, G; DeBach, P

Courtship behavior in the Aphytis lingnanensis group, its potential usefulness in taxonomy, and a review of sexual behavior in the parasitic Hymenoptera (Chalcidoidea: Aphelinidae)

Authors

Gordon Gordh
Paul DeBach

Authors Affiliations

Gordon Gordh was Assistant Professor and Assistant Entomologist, Division of Biological Control, Department of Entomology, Riverside; Paul DeBach was Professor of Biological Control, Department of Entomology, Riverside.

Publication Information

Hilgardia 46(2):37-75. DOI:10.3733/hilg.v46n02p037. March 1978.

PDF of full article, Cite this article

Abstract

The aphelinid genus Aphytis contains the most important natural control agents of numerous armored scale insect pests. The genus consists of several species groups, including the lingnanensis group. This group has several sibling or near-sibling species which are exceptionally useful in biological control.

Accurate identification of these parasites is supremely important in biological control, but this is often difficult when sibling or cryptic species are involved, as morphological characters used to discriminate taxa are absent. Therefore, studies were conducted to determine the feasibility of using behavioral characters for the separation of such species in the lingnanensis group.

Qualitative analysis of courtship behavior revealed that it is possible to separate A. lingnanensis from A. melinus DeBach, A. holoxanthus DeBach, A. fisheri DeBach, and a semispecies of A. melinus, on the basis of overt male behavior and female response to male behavior.

Quantitative analysis of 16 cultures in the lingnanensis group showed that it is possible to identify different species by using precoital courtship duration, postcoital mount duration, precoital wing vibration pulse frequency, postcoital wing vibration pulse frequency, number of contacts between the sexes, and number of male courtship attempts, as characters. In some instances, the parameters tested are useful in grouping allopatric populations of a species.

Some antennal ablation experiments were done and antennae were chemically treated to determine their role in courtship. These studies indicated that antennae are essential for successful copulation.

Scanning electron microscope studies revealed a mesothoracic

Literature Cited

Abbott C. E. How Megarhyssa deposits her eggs. J. N.Y. Entomol. Soc. 1934. 42:127-33.

Abdelrahman I. Growth, development and innate capacity for increase in Aphytis chrysomphali Mercet and A. melinus DeBach, parasites of California red scale, Aonidiella aurantii (Mask.), in relation to temperature. Austr. J. Zool. 1974. 22:213-30. DOI: 10.1071/ZO9740213 [CrossRef]

Adriaanse A. Ammophila campestris Latr. und Ammophila adriaansei Wilcke. Ein beitrag zur vergleichenden Verhaltenforschung. Behavior. 1947. 1:1-34. DOI: 10.1163/156853948X00010 [CrossRef]

Alam S. M. The biology of Metaphycus taxi Alam (Encyrtidae: Hymenoptera) in the constant temperature room, with notes on the anatomy of its pre-imaginal stages. Indian J. Entomol. 1957. 19:231-40.

Alexander R. D. The role of behavioral study in cricket classification. Syst. Zool. 1962. 11:53-72. DOI: 10.2307/2411453 [CrossRef]

Alexander R. D. The evolution of mating behavior in arthropods. Symp. R. Entomol. Soc. London. 1963. 2:78-94.

Allen W. W. The biology of Apanteles medicaginis Muesebeck (Hymenoptera: Braconidae). Hilgardia. 1958. 27:515-41. DOI: 10.3733/hilg.v27n18p515 [CrossRef]

Altson A. M. The life-history and habits of two parasites of blowflies. Proc. Zool. Soc. Lond. 1920. 15:195-243.

Arambourg Y. Chelonus eleaphilus (Hym. Braconidae) parasite de Prays oleae (Lep. Hyponomeutidae) élevage, charactéristiques et biologiques. Ann. Soc. Entomol. Fr. (N.S.). 1968. 4:385-411.

Askew R. R. Eupelmus urozonus Dalman (Hym., Chalcidoidea) as a parasite in cynipid oak galls. Entomol. 1961. 94:196-201.

Askew R. R. Considerations on speciation in Chalcidoidea (Hymenoptera). Evolution. 1968. 22:642-45. DOI: 10.2307/2406886 [CrossRef]

Assem J. van den. Reproductive behaviour of Pseudeucolia bochei I. A description of courtship behaviour. Neth. J. Zool. 1969. 19:641-48. DOI: 10.1163/002829669X00080 [CrossRef]

Assem J. van den. Courtship and mating in Lariophagus distinguendus (Forst.) (Hymenoptera, Pteromalidae). Neth. J. Zool. 1970. 20:329-52. DOI: 10.1163/002829670X00150 [CrossRef]

Assem J. van den. Maagd of geen maagd? Een markwaardige categorie wijfjes bÿ pteromalide Lariophagus distinguendus Kurdj. Entomol. Berich. 1974. 34:123-25.

Assem J. van den. Temporal patterning of courtship behaviour in some parasitic Hymenoptera, with special reference to Melittobia acasta. J. Entomol. (A). 1975. 50:137-46.

Assem J. van den, Povel G. D. E. Courtship behaviour of some Muscidifurax species (Hym., Pteromalidae); A possible example of a recently evolved ethological isolating mechanism. Neth. J. Zool. 1973. 23:465-87. DOI: 10.1163/002829673X00049 [CrossRef]

Assem J. van den, Visser J. Aspects of sexual receptivity in female Nasonia vitripennis (Hym., Pteromalidae). Biol. Behav. 1976. 1:37-56.

Avidov Z., Rössler Y., Rosen D. Studies on an Israel strain of Anagrus pseudococci (Girault) (Hym. Encyrtidae). II. Some biological aspects. Entomophaga. 1967. 12:111-18. DOI: 10.1007/BF02370607 [CrossRef]

Bakkendorf P. Biological investigations on some Danish Hymenopterous egg-parasites, with taxonomic remarks and descriptions of new species. Entomol. Meddelelser. 1934. 19:1-134.

Balfour-Browne F. On the life-history of Melittobia acasta Walker; a chalcidoid parasite of bees and wasps. Parasitology. 1922. 14:349-70. DOI: 10.1017/S003118200001026X [CrossRef]

Bar D., Gerling D. Biological studies of Pteroptrix smithi (Hymenoptera: Aphelinidae). Entomophaga. 1971. 16:19-36. DOI: 10.1007/BF02370686 [CrossRef]

Barrass R. The courtship behavior of Mormoniella vitripennis Walk. (Hymenoptera, Pteromalidae). Behavior. 1960a. 15:185-209. DOI: 10.1163/156853960X00223 [CrossRef]

Barrass R. The effect of age on the performance of an innate behaviour pattern in Mormoniella vitripennis Walk. (Hymenoptera, Pteromalidae). Behaviour. 1960b. 15:210-18. DOI: 10.1163/156853960X00232 [CrossRef]

Barrass R. A quantitative study of the behaviour of the male Mormoniella vitripennis (Walker) (Hymenoptera, Pteromalidae) towards two constant stimulus-situations. Behaviour. 1961. 18:288-311. DOI: 10.1163/156853961X00178 [CrossRef]

Barrass R. Preening and abdomen dipping by the male Mormoniella vitripennis after courtship. Behaviour. 1969. 35:304-312. DOI: 10.1163/156853969X00279 [CrossRef]

Beling I. von. Zur Biologie und Zucht der Schulpfwespe Angitia armillata Gr. (Hymenopt. Ichneum. Ophion.). Arb. Biol. Reichsanstalt Land u. Forstwirt. Berlin-Dahlem. 1933. 20:237-44.

Bennett F. D. Parasites of Ancylostomia stercorea (Zell.) (Pyralidae, Lepidoptera) a pod borer attacking pigeon pea in Trinidad. Bull. Entomol. Res. 1960. 50:737-57. DOI: 10.1017/S0007485300054778 [CrossRef]

Broodryk S. W., Doutt R. L. Studies of two parasites of olive scale, Parlatoria olea (Colvée). II. The biology of Coccophagoides utilis Doutt (Hymenoptera: Aphelinidae). Hilgardia. 1966. 37:233-54. DOI: 10.3733/hilg.v37n09p233 [CrossRef]

Buckell E. R. Notes on the life-history and habits of Melittobia chalybii Ashmead (Chalcidoidea: Elachertidae). Pan-Pac. Entomol. 1928. 5:14-22.

Burkhardt F. Eine neue Chalcididae der Gattung Dibrachys. Zentralblatt f. Bakteriologie, Parasitenkunde und Infektionskrankheiten. Abt. II. 1916. 46:502-04.

Campbell B. Sexual Selection and the Descent of Man, 1871-1971. 1972. Chicago: Aldine Publ. Co. 378p.

Cherian M. C., Narayanaswami P. S. The biology of Microbracon chilonis Viereck—a larval parasite of Chilo zonellus Swin. Ind. J. Entomol. 1942. 4:5-7.

Chrystal R. N. Studies of the Sirex parasites. Oxford Forestry Mem. 1930. 11:63

Clancy D. W. The biology of Tetracnemus pretiosus Timberlake. Univ. Calif. Publ. Entomol. 1934. 6:231-48.

Clarke C. A., Sheppard P. M. Offspring from double matings in swallowtail butterflies. Entomologist. 1962. 95:199-203. 1 fig., 1 tab

Clausen C. P. The biology of Schizaspidia tenuicornis Ashm., a eucharid parasite of Camponotus. Ann. Entomol. Soc. Amer. 1923. 16:195-217. 13 figs

Cousin G. Étude biologie d’un Chalcidien: Mormoniella vitripennis Walk. 1933. 67: Paris: Bull. Biol. p. 371-400.

Cox J. A. Ascogaster carpocapsae Viereck, an important larval parasite of the codling moth and the oriental fruit moth. Tech. Bull. N.Y. State Agr. Exp. Sta. 1932. 188:

Crandell H. A. The biology of Pachycrepoideus dubius Ashmead (Hymenoptera), a pteromalid parasite of Piophila casei Linne (Diptera). Ann. Entomol. Soc. Amer. 1939. 32:632-54.

Dahms E. C. The courtship behavior of Melittobia australica Girault, 1912, (Hymenoptera: Eulophidae). Mem. Qdlns. Mus. 1973. 16:411-14.

Darwin C. The Descent of Man, and Selection in Relation to Sex. 1871. London: J. Murray. DOI: 10.1037/12293-000 [CrossRef]

Delanoue P., Arambourg Y. Contribution a l’étude en laboratoire d’Eupelmus urozonus Dalm. (Hym. Chalcidoidea Eupelmidae). Ann. Soc. Entomol. Fr. (N.S.). 1965. 1:817-42.

Delanoue P., Arambourg Y. Contribution a l’étude en laboratoire de Pnigalio mediterraneaus (Hym. Chalcidoidea, Eulophidae). Ann. Soc. Entomol. Fr. (N.S.). 1967. 3:909-27.

Delanoue P., Arambourg Y. Contribution a l’étude en laboratoire d’Eurytoma martellii (Hym., Chalcidoidea, Eurytomidae). Ann. Soc. Entomol. Fr. (N.S.). 1969. 5:811-24.

Dobzhansky T. Genetics and the Origin of Species. 1951. New York: Columbia Univ. Pr. 364p.

Ducke A. Über Phylogenie und Klassifieation der sozialen Vespiden. Zool. Jb. Syst. 1914. 36:303-30.

Emerson A. E. Termite nests—a study of the phylogeny of behavior. Ecol. Monog. 1938. 8:249-84. DOI: 10.2307/1943251 [CrossRef]

Evans H. E. Comparative ethology and the systematics of spider wasps. Syst. Zool. 1953. 2:155-72. DOI: 10.2307/2411559 [CrossRef]

Evans H. E. The Comparative Ethology and Evolution of the Sand Wasps. 1966. Cambridge: Harvard Univ. Pr. 526p.

Fink D. E. The biology of Macrocentrus ancylivora Rohwer, an important parasite of the strawberry leaf roller (Ancylis comptana Froehl.). J. Agr. Res. 1926. 32:1121-34.

Finney G. L., Flanders S. E., Smith H. S. Mass culture of Macrocentrus ancylivorus and its host, the potato tuber moth. Hilgardia. 1947. 17:437-83. DOI: 10.3733/hilg.v17n13p437 [CrossRef]

Fisher J. P. The biology and taxonomy of some chalcidoid parasites (Hymenoptera) of stem-living larvae of Apion (Coleoptera: Curculionidae). Trans. R. Entomol. Soc. Lond. 1970. 122:293-322. DOI: 10.1111/j.1365-2311.1970.tb00525.x [CrossRef]

Fisher R. C. Life history and ecology of Horogenes chrysostictos Gmelin (Hymenoptera, Ichneumonidae), a parasite of Ephestia sericarium Scott (Lepidoptera, Phycitidae). Can. J. Zool. 1959. 37:429-46. DOI: 10.1139/z59-051 [CrossRef]

Flanders S. E. The role of the spermatophore in the mass propagation of Macrocentrus ancylivorus. J. Econ. Entomol. 1945. 38:323-27.

Foster W. A. Co-operation by male protection of ovipositing female in the Diptera. Nature. 1967. 214:1035-36. DOI: 10.1038/2141035a0 [CrossRef]

Gerling D. Studies with whitefly parasites of Southern California I. Encarsia pergandiella Howard (Hymenoptera: Aphelinidae). Can. Entomol. 1966. 98:707-24. DOI: 10.4039/Ent98707-7 [CrossRef]

Girault A. A., Saunders G. E. The chalcidoid parasites of the common house or typhoid fly (Musca domestica Linn.) and its allies. Psyche. 1910. 17:9-28. DOI: 10.1155/1910/17925 [CrossRef]

Given B. B. Evolutionary trends in the Thynninae (Hymenoptera; Tiphiidae) with special reference to the feeding habits of Australian species. Trans. R. Entomol. Soc. Lond. 1954. 105:1-10. DOI: 10.1111/j.1365-2311.1954.tb00762.x [CrossRef]

Goodpasture C. Comparative courtship behavior and karyology in Monodontomerus (Hymenoptera: Torymidae). Ann. Entomol. Soc. Amer. 1975. 68:391-97.

Gordh G. Biological investigations on Comperia merceti (Compere), an encyrtid parasite of the cockroach Supella longipalpa (Serville). J. Entomol. (A). 1972. 47:115-23.

Gordh G. The comparative external morphology and systematics of the Neotropical parasitic fig wasp genus Idarnes (Hymenoptera: Torymidae). Univ. Kansas Sci. Bul. 1975. 50:389-455.

Gordh G. Goniozus gallicola Fouts, a parasite of moth larvae with notes on other bethylids (Hymenoptera: Bethylidae; Lepidoptera: Gelechiidae) 1976. p.27. U. S. Dept. Agr. Tech. Bul. 1524

Gordh G., Lacey L. Biological and behavioral observations of Plagiomerus diaspidis Crawford, a primary internal parasite of the armored scale insect Diaspis echinocacti (Bouché) (Hymenoptera: Encyrtidae; Homoptera: Diaspididae). Proc. Entomol. Soc. Wash. 1976. 78:132-44.

Gordh G., Evans H. E. A new species of Goniozus imported into California from Ethiopia for the biological control of pink bollworm and some notes on the taxonomic status of Parasierola and Goniozus (Hymenoptera: Bethylidae). Proc. Entomol. Soc. Wash. 1976. 78(4):479-89.

Griswold G. H. On the bionomics of a primary parasite and of two hyperparasites of the Geranium aphid. Ann. Entomol. Soc. Amer. 1929. 22:438-57.

Grosch D. S. The importance of antennae in mating reaction of male Habrobracon. J. Comp. Psys. Psych. 1947. 40:23-39. DOI: 10.1037/h0062951 [CrossRef]

Grosch D. S. Dwarfism and differential mortality in Habrobracon. J. Exp. Zool. 1948. 107:289-313. DOI: 10.1002/jez.1401070207 [CrossRef]

Hafez M. The biology and life-history of Apanteles ruficrus Hal. (Hymenoptera-Braconidae). Bul. Soc. Fouad I^er Entomol. 1947. 31:225-49.

Hanna A. D. The male and female genitalia and the biology of Euchalcidia caryobori Hanna. (Hymenoptera: Chalcidinae). Trans. R. Entomol. Soc. Lond. 1934. 82:107-36. DOI: 10.1111/j.1365-2311.1934.tb00030.x [CrossRef]

Haramoto F. H. Observations on the biology of Opius oophilus Fullaway (Braconidae-Hymenoptera). Proc. 8th Pac. Sci. Cong. Manila. 1957. 3:1275-89.

Hase A. von. Beitrage zur morphologischen und biologischen Kenntnis der Schulpfwespe Lariophagus distinguendus (Forst.) 1919. pp.402-32. Kurdj. Gesell. natursfor. freunde, Berlin

Hase A. von. Neue Beobachtungen uber die Mannchen und Weibchen der Schulpfwespe Nemeritis canescens (Hymenoptera: Ichneumonidae) Arb. morph. tax. Ent. Berlin-Dahlem. 1937. 4:47-61.

Heatwole H., Davis D. M., Wenner A. M. The behavior of Megarhyssa, a genus of parasitic hymenopterans (Ichneumonidae: Ephialtinae). Zeitschr. f. Tierpsychol. 1963. 19:652-64. DOI: 10.1111/j.1439-0310.1962.tb00796.x [CrossRef]

Heatwole H., Davis D. M., Wenner A. M. Detection of mates and hosts by parasitic insects of the genus Megarhyssa (Hymenoptera: Ichneumonidae). Amer. Midl. Nat. 1964. 71:374-81. DOI: 10.2307/2423295 [CrossRef]

Heinroth O. Beiträge zur Biologie, namentlich Ethologie und Psychologie der Anatiden. 1911. Berlin: Verh. V. Internat. Ornith. Kong. p. 589-702. 1910

Hobbs G. A., Krunic M. D. Comparative behavior of three chalcidoid (Hymenoptera) parasites of the alfalfa leaf-cutter bee, Megachile rotundata, in the laboratory. Can. Entomol. 1971. 103:674-85. DOI: 10.4039/Ent103674-5 [CrossRef]

Hunter-Jones P. Fertilization of eggs of the desert locust by spermatozoa from successive copulations. Nature. 1960. 185:336 DOI: 10.1038/185336a0 [CrossRef]

Hussey N. W. The life-histories of Megastigmus spermotrophus Wachtl (Hym. Chalcidoidea) and its principal parasite, with descriptions of the developmental stages. Trans. R. Entomol. Soc. Lond. 1955. 106:133-51. DOI: 10.1111/j.1365-2311.1955.tb01266.x [CrossRef]

Huxley J. Darwin’s theory of sexual selection and the data subsumed by it, in the light of recent research. Amer. Nat. 1938. 72:416-33. DOI: 10.1086/280795 [CrossRef]

Jackson D. The biology of Dinocampus (Perilitus) rutilus Nees, a braconid parasite of Sitona lineata L.—Part I. Proc. Zool. Soc. Lond. 1928. 40:597-630.

Jackson D. Observations on the life-history of Mestocharis bimacularis (Dalman) (Hym. Eulophidae), a parasitoid of the eggs of Dytiscidae. Opusc. Entomol. 1964. 29:81-97.

Jackson D. Observations on the biology of Caraphractus cinctus Walker (Hymenoptera: Mymaridae), a parasitoid of the eggs of Dytiscidae (Coleoptera). iii. The adult life and sex ratio. Trans. R. Entomol. Soc. Lond. 1966. 118:23-49. DOI: 10.1111/j.1365-2311.1966.tb00833.x [CrossRef]

Jenni W. Beitrag zur Morphologie und Biologie der Cynipide Pseudeucoila bochei Weld, eines Larvenparasiten von Drosophila melanogaster Meig. Acta Zool. 1951. 32:177-254. DOI: 10.1111/j.1463-6395.1951.tb00468.x [CrossRef]

Johnson C. Breeding behavior and oviposition in Calopteryx maculatum (Beauvais) (Odonata: Calopterygidae). Amer. Midl. Nat. 1962. 68:242-47. DOI: 10.2307/2422650 [CrossRef]

Jones E. P. The egg parasites of the cotton bollworm, Heliothis armigera Hubn. (obsoleta Fabr.) in southern Rhodesia 1937. pp.37-105. Mazoe Citrus Exp. Sta. Ann. Rept. 1936

Juillet J. A. Morphology of immature stages, life-history, and behavior of three hymenopterous parasites of the European pine shoot moth, Rhyacionia buoliana (Schiff.) (Lepidoptera: Olethreutidae). Can. Entomol. 1959. 91:709-19. DOI: 10.4039/Ent91709-11 [CrossRef]

Kamal M. Biological studies of some hymenopterous parasites of aphidophagous parasites of aphidophagous Syriphidae. Tech. Sci. Serv. Entomol. Sect. Min. Agr. Egypt Bull. 1939. 207:109

Kashef A. Étude biologique du Stegobium paniceum L. (Col. Anobiidae) et de son parasite: Lariophagus distinguendus Först. Ann. Soc. Entomol. Fr. 1955. 124:1-88.

Khasimuddin S., DeBach P. Mating behavior and evidence of a male sex pheromone in species of the genus Aphytis. Ann. Entomol. Soc. Amer. 1975. 68(5):893-893.

Kugler J., Wollberg Z. Biology of Agrothereutes tunetanus Haber. (Hym. Ichneumonidae) an ectoparasite of Orgyia dubia Tausch. (Lep. Lymantriidae). Entomophaga. 1967. 12:363-79. DOI: 10.1007/BF02376923 [CrossRef]

Linsley E. G., McSwain J. W., Smith R. F. The bionomics of Diadasia consociata Timberlake and some biological relationships of emphorine and anthophorine bees (Hymenoptera, Anthophoridae). Univ. Calif. Publ. Entomol. 1952. 9:267-90.

MackAuer M. Sexual behavior of and hybridization between three species of Aphidius Nees parasitic on the pea aphid (Hymenoptera: Aphidiidae). Proc. Entomol. Soc. Wash. 1969. 71:339-51.

Martin C. H. Biological studies of two hymenopterous parasites of aquatic insect eggs. Entomol. Amer. 1927. 8:105-56. (n.s.)

Mayr E. Animal Species and Evolution. 1963. Cambridge: Belknap Press, Harvard Univ. Press. 797p.

Michelsen A. Observation on the sexual behaviour of some longicorn beetles, subfamily Lepturinae (Coleoptera, Cerambycidae). Behaviour. 1963. 22:152 66 DOI: 10.1163/156853963X00338 [CrossRef]

Michener C. D. The Social Behavior of the Bees. 1974. Harvard Univ. Press, Cambridge: A Comparative Study. Belknap Press. 404p.

Miller M. C. Emergence and mating of Tetrastichus incertus, a parasite of the alfalfa weevil. J. Econ. Entomol. 1966. 59:1532-33.

Moran V. C., Brothers D. J., Case J. J. Observations on the biology of Tetrastichus flavigaster Brothers and Moran (Hymenoptera: Eulophidae), parasitic on psyllid nymphs (Homoptera). Trans. R. Entomol. Soc. Lond. 1969. 121:41-59. DOI: 10.1111/j.1365-2311.1969.tb00516.x [CrossRef]

Morris K. R. S., Cameron E. The biology of Microplectron fuscipennis Zett. (Chalcid.), a parasite of the pine sawfly (Diprion sertifer, Geoff.). Bul. Entomol. Res. 1935. 26:407-18. DOI: 10.1017/S0007485300036737 [CrossRef]

Muesebeck C. F. W. Monodontomerus aureus Walker, both a primary and a secondary parasite of the browntail moth and the gypsy moth. J. Agr. Res. 1931. 43:445-60.

Mürr L. Über den Geruchsinn der Mehlmottenschulpfwespe Habrobracon juglandis Ashmead. A. vergl. Physiol. 1930. 11:210-70. DOI: 10.1007/BF00338869 [CrossRef]

Myers J. G. Further notes on Alysia manducator and other parasites (Hym.) of muscoid flies. Bull. Entomol. Res. 1929. 19:357-60. DOI: 10.1017/S0007485300020721 [CrossRef]

Nielsen E. T. Sur les habitudes des hyménoptères aculéates solitaires. I. (Bethylidae, Scoliidae, Cleptidae, Psammocharidae). Entomol. Meddelelser. 1932. 18:1-57.

Noble N. S. Studies of Habrocytus cerealellae (Ashmead), a pteromalid parasite of the angoumois grain moth, Sitotroga cerealella (Oliver). Univ. Calif. Publ. Entomol. 1932. 5:311-54.

Noble N. S. The citrus gall wasp (Eurytoma fellis Girault). Dept. Agr. N.S.W. Sci. Bull. 1936. 53:1-41.

Noble N. S. Euplectris agaristae Craw. a parasite of the grape vine moth (Phalaenoides glycine Lew.). Dept. Agr. N.S.W. Sci. Bull. 1938a. 63:5-27.

Noble N. S. Epimegastigmus (Megastigmus) brevivalvus Girault: A parasite of the citrus gall wasp (Eurytoma fellis Girault); with notes on several other species of hymenopterous gall inhabitants. Dept. Agr. N.S.W. Sci. Bull. 1938b. 65:5-46.

Noble N. S. A new species of Megastigmus parasitic on Tepperella trilineata Cam., a wasp causing galling of the flower buds of Acacia decurrens. Proc. Linn. Soc. N.S.W. 1939. 64:266-78.

Parker G. A. Sperm competition and its evolutionary consequences in the insects. Biol. Rev. 1970a. 45:528-68. DOI: 10.1111/j.1469-185X.1970.tb01176.x [CrossRef]

Parker G. A. Sperm competition and its evolutionary effect on copula duration in the fly Scatophaga stercoraria. J. Ins. Physiol. 1970b. 16:1301-28. DOI: 10.1016/0022-1910(70)90131-9 [CrossRef]

Parker G. A. Courtship persistence and female-guarding as male time investment strategies. Behaviour. 1974. 48:157-84. DOI: 10.1163/156853974X00327 [CrossRef]

Parker H. L. Macrocentrus gifuensis Ashmead, a polyembryonic braconid parasite in the European corn borer. U.S.D.A. Tech. Bull. 1931. 230:1-63.

Parker H. L., Smith H. D. Eulophus viridulus Thoms., a parasite of Pyrausta nubilalis Hubn. Ann. Entomol. Soc. Amer. 1933. 26:21-36.

Parker H. L., Thompson W. R. Contribution à la Biologie des chalcidiens entomophages. Ann. Entomol. Soc. Fr. 1928. 97:423-65.

Petrunkevitch A. The value of instinct as a taxonomic character in spiders. Biol. Bull. 1926. 50:427-32. DOI: 10.2307/1536614 [CrossRef]

Pinkus H. The life history and habits of Spalangia muscidarum Richardson, a parasite of the stable fly. Psyche. 1913. 20:148-58. DOI: 10.1155/1913/56729 [CrossRef]

Pinto J. D. A synopsis of the bionomics of Phodaga alticeps (Coleoptera: Meloidae) with special reference to sexual behavior. Can. Entomol. 1972a. 104:577-95. DOI: 10.4039/Ent104577-4 [CrossRef]

Pinto J. D. The sexual behavior of Meloe (Meloe) strigulosus Mannerheim. J. Kans. Entomol. Soc. 1972b. 45::128-35.

Plath O. E. Bumble Bees and Their Ways 1934. p.210. 2 pl. New York, MacMillan Co

Quednau F. W. Notes on mating behavior and oviposition of Chrysocharis laricinellae (Hymenoptera: Eulophidae), a parasite of the larch casebearer (Coleophora laricella). Can. Entomol. 1967. 99:326-31. DOI: 10.4039/Ent99326-3 [CrossRef]

Quezada J. R., DeBach P., Rosen D. Biological and taxonomic studies of Signiphora borinquensis, new species, (Hymenoptera: Signiphoridae), a primary parasite of diaspine scales. Hilgardia. 1973. 41:543-604. DOI: 10.3733/hilg.v41n18p543 [CrossRef]

Rao S. V., DeBach P. Experimental studies on hybridization and sexual isolation between some Aphytis species (Hymenoptera: Aphelinidae). I. Experimental hybridization and an interpretation of evolutionary relationships among the species. Hilgardia. 1969a. 39:515-53. DOI: 10.3733/hilg.v39n19p515 [CrossRef]

Rao S. V., DeBach P. Experimental studies on hybridization and sexual isolation between some Aphytis species (Hymenoptera: Aphelinidae). II. Experiments on sexual isolation. Hilgardia. 1969b. 39:555-67. DOI: 10.3733/hilg.v39n19p555 [CrossRef]

Rao S. V., DeBach P. Experimental studies on hybridization and sexual isolation between some Aphytis species (Hymenoptera: Aphelinidae). iii. The significance of reproductive isolation between interspecific hybrids and parental species. Evolution. 1969c. 23:525-33. DOI: 10.2307/2406850 [CrossRef]

Richard G., Barton Browne L. Sequential analysis and regulation of insect reproductive behavior. Experimental Analysis of Insect Behaviour. 1974. New York: Springer-Verlag. 366p.

Rosen D., DeBach P. Systematics, morphology and biological control. Entomophaga. 1973. 18:215-22. DOI: 10.1007/BF02371468 [CrossRef]

Safavi M. Le comportment sexual chez les Asolcus Nakagawa (Hym., Proctotrupoidea, Scelionidae). Rev. Path. veg. d’Entomol. agr. Fr. 1963. 42:127-34.

Schell S. C. The biology of Hadronotus ajax Girault (Hymenoptera-Scelionidae), a parasite in the eggs of squash-bug (Anasa tristis DeGeer). Ann. Entomol. Soc. Amer. 1943. 36:625-35.

Schlager G. Sperm precedence in the fertilization of eggs in Tribolium castaneum. Ann. Entomol. Soc. Amer. 1960. 53:557-60.

Schlinger E. I., Hall J. C. The biology, behavior and morphology of Praon palitans Muesebeck, an internal parasite of the spotted alfalfa aphid, Therioaphis maculata (Buckton) (Hymenoptera: Braconidae, Aphidiinae). Ann. Entomol. Soc. Amer. 1960. 53:144-60.

Schlinger E. I., Hall J. C. The biology, behavior and morphology of Trioxys (Trioxys) utilis, an internal parasite of the spotted alfalfa aphid, Therioaphis maculata (Hymenoptera: Braconidae, Aphidiidae). Ann. Entomol. Soc. Amer. 1961. 54:34-45.

Seitner M., Nötzl P. Pityophthorus henscheli Seitner und sein Parasite, Cosmophorus henscheli Ruschka. Zeitsch. Angew. Entomol., Berlin. 1925. 11:187-96. DOI: 10.1111/j.1439-0418.1925.tb00001.x [CrossRef]

Sekhar P. S. Mating, oviposition, and discrimination of hosts by Aphidius testaceipes (Cresson) and Praon aguti Smith, primary parasites of aphids. Ann. Entomol. Soc. Amer. 1957. 50:370-75.

Selander R. K., Campbell B. Sexual selection and dimorphism in birds. Sexual Selection and the Descent of Man. 1871-1971. 1972. Chicago: Aldine-Atherton. p. 180-230.

Simmonds F. J. Observations on the biology and mass-breeding of Spalangia drosophilae Ashm. (Hymenoptera: Spalangiidae), a parasite of the frit-fly Oscinella frit (L.). Bull. Entomol. Res. 1953. 44:773-78. DOI: 10.1017/S0007485300024718 [CrossRef]

Smith L. M. Macrorileya oecanthi Ashm. a hymenopterous egg parasite of tree crickets. Univ. Calif. Publ. Entomol. 1930. 5:165-72.

Speith H. T. Mating behavior within the genus Drosophila (Diptera). Bull. Amer. Mus. Nat. Hist. 1952. 99:399-474.

Spencer H. Biology of the parasites and hyperparasites of aphids. Ann. Entomol. Soc. Amer. 1926. 19:119-57.

Thornhill R. Sexual selection and paternal investment in insects. Amer. Nat. 1976. 110:153-63. DOI: 10.1086/283055 [CrossRef]

Thorpe W. H., Huxley J. Ecology and the future of systematics. The New Systematics. 1940. Oxford Univ. Pr. p. 341-364.

Thorpe W. H. The modern concept of instinctive behavior. Bull. Anim. Behav. 1948. 2:7-12.

Torchio P. F. A chalcid wasp parasite of the alfalfa leafcutting bee. Utah Farm and Home Science. 1963. 24:70-71.

Trivers R. L., Campbell B. Parental investment and sexual selection. Sexual Selection and the Descent of Man, 1871-1971. 1972. Chicago: Aldine-Atherton. p. 136-79.

Vité J. P., Pitman G. B. Bark beetle aggregation: effects of feeding on the release of pheromones in Dendroctonus and Ips. Nature. 1968. 218:169-70. DOI: 10.1038/218169a0 [CrossRef]

Vité J. P., Pitman G. B. Aggregation behaviour of Dendroctonus brevicornis in response to synthetic pheromones. J. Ins. Physiol. 1969a. 15:1617-22. DOI: 10.1016/0022-1910(69)90182-6 [CrossRef]

Vité J. P., Pitman G. B. Insect and host odors in the aggregation of the western pine beetle. Can. Entomol. 1969b. 101:113-17. DOI: 10.4039/Ent101113-2 [CrossRef]

Voukassovitch P. Sur la biologie de Goniozus claripennis Först. parasite d’Oenophthira pilleriana Schiff. (Hymenoptera). Soc. d’Hist. natur. de Toulouse Bull. 1924. 52:225-46.

Waloff N. Biology of three species of Leiophron (Hymenoptera: Braconidae, Euphorinae) parasitic on Myridae on broom. Trans. R. Entomol. Soc. Lond. 1967. 119:187-213. DOI: 10.1111/j.1365-2311.1967.tb00509.x [CrossRef]

Whiting P. N. Reproductive reactions of sex mosaics of a parasitic wasp, Habrobracon juglandis. J. Comp. Psychol. 1932. 14:345-63. DOI: 10.1037/h0071790 [CrossRef]

Wigglesworth V. B. The Principles of Insect Physiology. 1950. 4th Ed. London: Methuen and Co. 544p.

Wilkes A. Sperm transfer and utilization by the arrhenotokous wasp Dahlbominus fuscipennis (Zett.) (Hymenoptera: Eulophidae). Can. Entomol. 1965. 97:647-57. DOI: 10.4039/Ent97647-6 [CrossRef]

Willard H. F. Parasites of the pink bollworm in Hawaii. U.S.D.A. Tech. Bull. 1927. 19:1-15.

Williams J. R. The biology of Physcus seminotus Silv. and P. subflavus Annecke and Insley (Aphelinidae), parasites of the sugar-cane scale insect Aulacaspis tegaleusis (Zhnt.) (Diaspididae). Bull. Entomol. Res. 1972. 61:463-84. DOI: 10.1017/S0007485300047337 [CrossRef]

Wilson F. Adult reproductive behavior in Asolcus basalis (Hymenoptera: Scelionidae). Aust. J. Zool. 1961. 9:737-51. DOI: 10.1071/ZO9610739 [CrossRef]

Wilson F., Woolcock L. T. Environmental determination of sex in a parthenogenic parasite. Nature. 1960. 186:99-100. DOI: 10.1038/186099a0 [CrossRef]

Also in this issue:

Land grant model: Help for the new Russia

A study asks: Are California's farmers headed toward sustainable agriculture?

Commodity advertising pays… or does it? What it takes to keep those raisins dancing

Late-season nitrogen may be efficient way to increase winter wheat protein

Competitive with soybean: White lupin could be new high-protein seed and forage for California

In California's municipalities, saving native oaks calls for planning

Apple rootstocks evaluated for California

To control postharvest decay and insects, moist heat treatments of strawberries are studied

Resistant cultivars, fungicides combat downy mildew of spinach

New moisture meter could curb overdrying of walnuts

Management strategies outlined: Research reveals pattern of cucurbit virus spread