Morphological and Biological Parameters of the Knapweed Nematode, Subanguina Picridis 1
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154 Journal of Nematology, Volume 18, No. 2, April 1986 weeds. 43. Acroptilon (Centaurea) repens (L.) DC. Ca- repens (L.) DC., Russian Knapweed (Compositae). Pp. nadian Journal of Plant Science 60:993-1004. 105-110 in J. S. Kelleher and M. A. Hulme, eds. 19. Watson, A. K. 1986. Host range of, and plant Biological control programmes against insects and reaction to, Subanguina picridis. Journal of Nematol- weeds in Canada 1969-1980. Slough: Common- ogy 18:112-120. wealth Agriculture Bureaux. 20. Watson, A. K. 1986. Morphological and bio- 22. Watson, A. K., and J. D. Shorthouse. 1979. logical parameters of the knapweed nematode, Suban- Gall formation on Cirsium arvense by Ditylenchus dip- guina _picridis.Journal of Nematology 18:154-158. saci. Journal of Nematology 11:16-22. 21. Watson, A. K., and P. Harris. 1984. Acroptilon Journal of Nematology 18(2):154-158. 1986. © The Society of Nematologists 1986. Morphological and Biological Parameters of the Knapweed Nematode, Subanguina picridis 1 A. K. WATSON 2 Abstract: Specimens of the knapweed nematode Subanguina picridis (Kirjanova) Brzeski obtained from different host plants were highly variable in measurement and structure. This variability refutes the validity of six Subanguina species attacking plants in the Asteraceae. Key words: Acroptilon repens (Russian knapweed), host specificity, Subanguina picridis (knapweed nematode), taxonomy, variability. The gall forming nematode Subanguina MATERIALS AND METHODS picridis (Kirjanova, 1944) Brzeski, 1981, Galls were obtained from A. repens, Cen- syn.: Paranguina picridis (Kirjanova, 1944) taurea diffusa Lam., Cynara sco$ymus L., and Kirjanova & Ivanova, 1968, has been eval- Onopordum acanthium L. plants inoculated uated in Canada as a potential biological with S. picridis (12). Galls were collected control agent of Russian knapweed, Acrop- 20-40 days after their appearance and were tilon repens (L.) DC., syn.: Centaurea repens dissected and immersed in aerated distilled L. The nematode has a restricted host water for 1 hour. Nematodes were re- range and damages its host. It has been moved from the water with a pipet and released in Canada for experimental con- concentrated in a small volume of distilled trol tests of Russian knapweed (12-14). water. They were then gently heat killed, Russian workers reported that S. picridis fixed in TAF (4), and processed to pure was specific to A. repens (5,6). However, glycerin using Seinhorst's (9) glycerol-eth- host tests have demonstrated that the host anol method. Measurements ofstylet length range of S. picridis is not restricted to Rus- of living nematodes from A. repens and C. sian knapweed but that some members of scolymus galls were also made. Data were the Cynareae tribe are also hosts (12). analyzed by an analysis of variance with a Therefore, S. picridis specimens from galls one-way classification (10). on the different hosts were examined to In addition, nematodes from A. repens, determine if different host plants affected C. diffusa, C. scolymus, and Cirsium flodmanii nematode morphology and anatomy. (Rydb.) Arthur, collected and fixed simi- larly, were prepared for the scanning elec- tron microscopy. Fixed specimens were de- Received for publication 26 February !985. hydrated in a graded ethanol series, then Research conducted at Agriculture Canada, ResearchSta- transferred to amyl acetate, and critical tion, Regina, Saskatchewan,Canada S4P 3A2. The work was supported by the British Columbia Cattleman'sAssociation, point dried using freon. Specimens were the Universityof Saskatchewan (Hantleman Scholarship), and individually placed on the adhesive surface Agriculture Canada, Regina Research Station. Department of Plant Science, MacdonaldCollege of McGill of stubs with their heads free and coated University, 21,111 Lakeshore Rd., Ste.-Anne-de-Bellevue, with carbon and gold. They were exam- Quebec H9X 1CO, Canada. ined and photographed in a Cambridge I thank P. Harris for reviewing the manuscript, G. Bray- brook for preparation of the SEM specimens, and A. Virly scanning electron microscope, using 20-kV for preparation of photographic plates. electron gun potential. Subanguina picridis: Watson 155 ~~o "~ ..~ " ~ ~ ~ O. v v .9 0 ~ ~v, q9 .9 = ~ o ~ + 0 X ~a ~ a o e ~2 ~ ~ 0 0 0 @.-- < fl II fl II II I .................................... 156 Journal of Nematology, Volume I8, No. 2, April 1986 TABLE2. Dimensions of eggs from gall cavities and from within the uterus of females of four populations of Subanguina picridis on different host plants (with mean and range). Population n Length (ttm) Width (/*m) Gall cavity Acroptilon repens 10 71.6 b 53.1 a (56.3-85.5) (42.8-60.3) Onopordum acanthium 10 79.7 ab 44.8 b (67.5-94.5) (38.3-54.0) Cynara scolymus 10 81.9 a 36.5 c (72.0-96.8) (33.8-45.0) Centaurea diffusa 10 73.6 ab 39.9 c (60.8-87.8) (27,0-45.0) Uterus Acroptilon repens 10 70.9 b 39.4 a (60.8-76.5) (31.5-45.0) Onopordum acanthium 10 78.1 a 36.5 ab (69.8-83.3) (31.5-40.5) Cynara scolymus 5 84.6 a 35.1 b (76.5-101.3) (31.5-38.3) Centaurea diffusa No eggs observed Means in the same column'sharingthe same letter do not differsignificantly at P -< 0.05 accordingto Duncan's multiple- range test. RESULTS' males from galls on A. repens were also wid- Dimensions of S. picridis from different er than those in females from galls on C. hosts varied considerably (Tables 1, 2). scolymus. Length of females from A. repens was (P -- The number of lip annuli on specimens 0.05) shorter than specimens from 0. acan- ofS. picridis from A. repens galls varied from thium and C. diffusa but not different from three to five, whereas specimens from galls C. scolymus (Table 1). Significant (P -< 0.05) on the other plants had four or five lip differences occurred among the popula- annuli (Figs. 1-4). Such differences have tions in the values of a, b, and c. Males been used to separate species of Suban- from C. diffusa galls differed (P-< 0.05) guina, but populations of S. picridis exhib- from males from galls on other host plants ited similar variability when reared on dif- in the measurements of L, a, b, and c, and ferent hosts. spicules of males from galls on C. scolymus were shorter (P < 0.05) than spicules of DISCUSSION males from other plants. Prior to Brzeski's (2) taxonomic treat- The stylets in some fixed specimens were ment of Anguinidae, Paranguina con- obscure, so comparisons of stylets among tained six species (10). Species were distin- populations could not be made. Most of guished by host range and morphology (6). the visible stylets in preserved specimens Morphological characters used were the were 8-10 /zm long. However, stylets of number of head annuli, position of excre- living adults from A. repens and C. scolymus tory pore, width of stylet base, stylet length, galls were 12/~m long. spicule and egg dimensions, and cauda Dimensions of eggs from galls on differ- morphology. For those Subanguina species ent hosts also varied. Eggs from a gall on that parasitize members of the Cynareae A. repens were shorter (P < 0.05) than those tribe of the Asteraceae family, most di- from a C. scolymus gall and wider than those mensions of the different species overlap from O. acanthium, C. scolymus, or C. diffusa (6,8). The report by Ahmed et al. (1) of galls (Table 2). Eggs within the uterus of Anguina microlaenae (Fawcett, 1938) Stei- females from A. repens galls were shorter ner, 1940 causing leaf and stem galls on (P - 0.05) than eggs in females from O. Centaurea rigida Wiild. and Cousinia steno- acanthium and C. scolymus galls. Eggs in fe- cephala Boiss., both of the Cynareae tribe Subanguina picridis: Watson 157 Fins. 1-4. En face views of Subanguina picridis from four different host plants. Bar represents 1 ttm. Numbers indicate the number of lip annuli. 1) Adult from a gall on Acroptilon repens. 2)Juvenile from a gall on Centaurea diffusa. 3) Adult from a gall on C)mara scolymus. 4) Adult from a gall on Cirsiumflodmanii. of the Asteraceae family, in Iraq is prob- anguina that form leaf and stem galls on ably incorrect because the nematode was plants of the Cynareae tribe are a single identified using a key that did not include species. Possibly host races have developed Paranguina species. on the various host plants in the Soviet Because a species is a dynamic group of Union. Sturhan (11) suggested that the individuals, variability within natural pop- probability of new distinct races forming ulations is to be expected. Populations that under natural conditions is low and that are morphologically different are not nec- races may develop in geographical isola- essarily taxonomically different (3). In this tion or may evolve in ecological or phys- study, differences were shown to exist iological isolation within a host plant. among populations of S. picridis in galls The species of Subanguina on the Cy- from different host plants. The degree of nareae host plants are not geographically variation in morphological characters of S. isolated. They are all found in the same picridis suggests that the six species of Sub- area of the southern Soviet Union (O. V. 158 Journal of Nematology, Volume 18, No. 2, April 1986 Kovalev, pers. comm.). Subanguina picridis culture, Fisheries & Food. London: Her Majesty's Sta- is widespread in the river valleys of Ta- tionary Office. 5. Ivanova, T. S. 1966. Biological control of dzhikistan, and four of the other species of mountain bluet (Acroptilon picris C.A.M.). (In Russian.) Subanguina on Cynareae host plants occur Izvestiya Academii Nauk Tadzhikskol SSR 9:51-63.