The anthomyiid fly Pegomya curticornis (Stein), introduced into Alberta from Europe for the biocontrol of leafy spurge (Euphorbia esula L.), normally induces simple galls on subterranean stems. Here we describe the anatomy of galls collected in Europe that had been induced by P. curticornis on horizontal roots of spurges. Four mature galls were found on the roots of E. virgata Waldst. and Kit. and 10 on the roots of E. esula. Tissues of the root galls were composed of gall parenchyma that had proliferated from feeding sites near the outside edge of secondary xylem. It is assumed that some larvae inadvertently tunnel beyond the base of stems into the roots and that similar galls will form on Alberta spurge

Gassmann, Andre

Shorthouse, Joseph D.

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The Canadian Entomologist  Vol. 126 Ottawa, Canada, March /April 1994 No. 2 GALL INDUCTION BY PEGOMYA CURTICORNIS (STEIN) (DIPTERA: ANTHOMYIIDAE) WITHIN THE ROOTS OF SPURGES EUPHORBZA VIRGATA WALDST. AND KIT. AND E. ESULA L. (EUPHORBIACEAE) JOSEPH D. SHORTHOUSE Department of Biology, Laurentian University, Sudbuy, Ontario, Canada P3E 2C6 and ANDRE GASSMANN CAB International Institute of Biological Control, European Station, CH-2800, Delkmont, Switzerland Abstract The Canadian Entomologist 126: 193-197 (1994) The anthomyiid fly Pegomya curticornis (Stein), introduced into Alberta from Europe for the biocontrol of leafy spurge (Euphorbia esula L.), normally induces simple galls on subterranean stems. Here we describe the anatomy of galls collected in Europe that had been induced by P. curticornis on horizontal roots of spurges. Four mature galls were found on the roots of E. virgata Waldst. and Kit. and 10 on the roots of E. esula. Tissues of the root galls were composed of gall parenchyma that had proliferated from feeding sites near the outside edge of secondary xylem. It is assumed that some larvae inad- vertently tunnel beyond the base of stems into the roots and that similar galls will form on Alberta spurges. Shorthouse, J.D., et A. Gassmann. 1994. Formation de galles sur les racines des euphorbes Euphorbia virgata Waldst. et Kit. et E. esula L. (Euphorbiaceae) par Pegomya curticornis (Stein) (Diptera: Anthomyiidae). The Canadian Entomologist 126: 193-197. Resume L'anthomyie Pegomya curticornis (Stein), impoitte d'Europe et introduite en Alberta pour assurer le contr6le biologique de l'euphorbe Euphorbia esula L., forme normale- ment des galles simples sur la partie souterraine des tiges. Nous dtcrivons ici l'anatomie de galles trouvkes en Europe et caustes par P. curticornis sur les racines horizontales d'euphorbes. Quatre galles mikes ont t t t  trouv&s sur les racines d'E. virgata Waldst. et Kit. et 10 sur les racines d'E. esula. Les tissus des galles radiculaires ttaient compos&s de parenchyme de galle provenant des points d'alimentation situts aux abords de la zone externe du xylkme secondaire. I1 semble que certaines larves s'aventurent au-delh de la base des tiges et creusent des tunnels jusque dans les racines et il faut s'attendre h trouver des galles semblables sur les euphorbes d'Alberta. [Traduit par la rkdaction] Introduction In previous studies (Gassmann and Shorthouse 1990, 1992), we reported on how two species of anthomyiid flies [Pegomya curticornis (Stein) and P. euphorbiae (Kieffer)] induce simple galls on the subterranean stems of several species of spurge Euphorbia (Euphor- biaceae) in Europe. As little is known about the few gall-inducing anthomyiids in the world (Meyer 1987), any information on their biology is of interest. Furthermore, P. curticornis recently has been introduced into Alberta for the biological control of leafy spurge (Euphor- bia esula L.) and, as with all biocontrol programs, additional information on activities of the agents is of value. Here we report on the structural modifications to the roots caused by of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.4039/Ent126193-2Downloaded from https:/www.cambridge.org/core. University of Basel Library, on 10 Jul 2017 at 16:23:54, subject to the Cambridge Core terms194 THE CANADIAN ~ M D I ~ I S T  March /April 1994 Frcs. 1, 2. 1. Habitus of Pegomya curticornis on a horizontal root of Euphorbiae virgata; X 1.5. Arrows indicate the approximate areas from which sections in Figures 2-6 were taken. 2. Cross section of an undamaged hori- zontal root; X78. C, cork; Co, cortex; P, phloem; X, secondary xylem. of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.4039/Ent126193-2Downloaded from https:/www.cambridge.org/core. University of Basel Library, on 10 Jul 2017 at 16:23:54, subject to the Cambridge Core termsvolume 126 THE CANADIAN ENIUMOLOGIR 195 larvae of P. curticornis that had induced galls in horizontal roots instead of subterranean stems. Materials and Methods In the process of collecting over 500 mature galls on the subterranean shoots of various spurges in Hungary, the second author found four mature galls on the roots of E. virgata Waldst. and Kit. More recently, 10 additional root galls were found on E. esula [for the taxon- omy of E. esula group see Crompton et al. (1990) and references therein]. Galls were removed from their hosts in early June; the four from E. virgata and two from E. esula were fixed in Formalin - acetic acid - alcohol (FAA) and sent to the first author for processing. Tissues were dehydrated in a tertiary-butyl alcohol series, embedded in paraffin, sectioned at 8 1J.m with a rotary microtome, and stained with safranin-fast green (Jensen 1962). Sections from unattacked roots were also made from healthy plants and from near the root galls. Flies from the remaining galls from E. esula were reared to adults. Results and Discussion All galls contained a mature larva or pupa. Each gall was approximately 1.5 cm long and 0.8 cm wide (Fig. 1) and found on mature roots exhibiting secondary growth (Fig. 2). The anatomy of roots and four galls from E. virgata was identical with that of the two galls and roots of E. esula; however, all figures discussed here are from E. virgata. In contrast to stems of spurge [which consist of pith, radial files of secondary xylem elements surrounded by secondary phloem, and a cortical parenchyma (Gassmann and Shorthouse 1990)], mature roots have a central core composed of secondary xylem arranged in radiating rows, which collectively occupy about two-thirds of the diameter (Raju 1985). There is no pith and the xylem is surrounded by cambium, phloem, and mechanical tissue which also forms radiating strips. The cortex consists of a few layers of irregular, mostly oval cells. Laticiferous cells occur scattered throughout the cortex (Bakshi and Coupland 1959). In place of the epidermis of stems, the outside margin of roots consists of a thick sheath of cork cells. In all six samples, the larvae had tunnelled (Fig. 3) through the secondary xylem (compared with the pith in the stem) and in some areas had damaged the cambium, phloem, and cortex. Callus proliferated from the damaged areas and was responsible for the irregular root swelling between the gall and vertical stem. Sections through the pupal chamber of mature galls (Fig. 4) showed that masses of gall parenchyma proliferated from the area where the larvae had chewed through the cambium into the cortex. Gall parenchyma had apparently formed as the larvae were feeding, as thick layers completely surrounded the feeding site (Fig. 5), leaving the once central core of secondary xylem to one side. New vascular bundles were formed within the gall parenchyma peripheral to the circular mass of secondary xylem (Fig. 6). Galls induced by P. curticornis on roots of spurge are similar to those induced on stems. Both are simple galls consisting of callus around sites of tunnelling and gall parenchyma around feeding sites, and are without nutritive and mechanical tissues common in the galls of most other insects (Rohfritsch 1992). Although gall insects are highly organ specific and instances of species inducing galls on the wrong organ are rare (Shorthouse and Lalonde 1986), galls of P. curticornis on roots are apparently another such anomaly. It appears that P. curticornis has the ability to manipulate growth of both stems and roots. It is possible that the larvae tunnelled beyond the base of the subterranean shoots into the roots because the shoots on which eggs were laid were too immature to allow larval development. When stems of spurge are galled by P. curticornis, further stem growth and flowering is curtailed (Gassmann and Shorthouse 1990). In root galls, it is not known how much shoot growth is affected; however, tunnelling through secondary xylem and induction of callus- like gall parenchyma is bound to cause some drain on the host plant. Thus, even if some of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.4039/Ent126193-2Downloaded from https:/www.cambridge.org/core. University of Basel Library, on 10 Jul 2017 at 16:23:54, subject to the Cambridge Core terms196 TFE CANADIAN F.NTOMOLOGI~ March /April 1994 FIGS. 3 4 .  Sections through larval tunnel and mature galls inhabited by pupae of Pegomya curticornis. 3. Section of horizontal root between vertical stem and gall showing tunnel (arrow) made by boring larva; X24. 4. Section through mature gall showing proliferation of gall parenchyma along the edge of secondary xylem damaged by feeding larva. Note the layer of dark staining, crushed cells between the secondary xylem and gall parenchyma (arrow); X60. 5. Section through mature gall showing proliferation of gall parenchyma peripheral to the circular zone of secondary xylem. Note the layer of dark staining, crushed cells between secondary xylem and gall parenchyma (arrows); X 16. 6. Section through mature gall showing masses of gall parenchyma peripheral to the core of secondary xylem. Note the presence of vascular bundles (arrows) in the gall parenchyma; X24. C, cork; Co, cortex; GP, gall parenchyma; P, phloem; Pu, pupa; X, xylem. P. curticornis feeding on Canadian spurge induce root instead of stem galls, they will still be responsible for host damage. Acknowledgments We thank V. Michelsen for identifying P. curticornis from the root galls of both E. virgata and E. esula, and the technical staff in Delemont for help with collecting and rearing. Investigations on Pegomya spp. for the biocontrol of leafy spurge were funded by the Alberta Environmental Centre, Vegreville, Alberta, under contracts 82-0830 and 86-0017. We also acknowledge support from an NSERC operating grant and the Laurentian University Research Fund awarded to JDS. of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.4039/Ent126193-2Downloaded from https:/www.cambridge.org/core. University of Basel Library, on 10 Jul 2017 at 16:23:54, subject to the Cambridge Core termsvolume 126 THE CANADIAN ENTOMOLOGIST 197 References Bakshi, T.S., and R.T. Coupland. 1959. An anatomical study of the subterranean organs of Euphorbia esula in relation to its control. Canadian Journal of Botany 37: 613-620. Crompton, C.W., A.E. Stahevitch, and W.A. Wojtas. 1990. Morphometric studies of the Euphorbia esula group (Euphorbiaceae) in North America. Canadian Journal of Botany 68: 1978-1988. Gassmann, A,, and J.D. Shorthouse. 1990. Structural damage and gall induction by Pegomya curticornis and Pegomya euphorbiae (Diptera: Anthomyiidae) within the stems of leafy spurge (Euphorbia X pseudovirgata) (Euphorbiaceae). The Canadian Entomologist 122: 429-439. - 1992. Structural damage and gall induction by Pegomya euphorbiae (Kieffer) (Diptera: Anthomyiidae) w i h n  the stems of cypress spurge (Euphorbia cyparissias L.) (Euphorbiaceae). Phytophaga 4: 61-64. Jensen, W.A. 1962. Botanical Histochemistry. Freeman, San Francisco, CA. 408 pp. Meyer, J. 1987. Plant Galls and Gall Inducers. Gebriider Bomtraeger, Stuttgart, Germany. 291 pp. Raju, M.V.S. 1985. Morphology and anatomy of leafy spurge. pp. 26-71 in Watson, A.K. (Ed.), Leafy Spurge. Monograph Series of the Weed Science Society of America 3: 104 pp. Rohfritsch, 0.1992. Patterns in gall development. pp. 60-86 in Shorthouse, J.D., and 0. Rohfritsch (Eds.), Biology of Insect-Induced Galls. Oxford University Press, New York, NY. 285 pp. Shorthouse, J.D., and R.G. Lalonde. 1986. Formation of flowerhead galls by the Canada thistle gall-fly, Urophora cardui (Diptera: Tephritidae), under cage conditions. The Canadian Entomologist 118: 1199-1203. Date received: 10 June 1993; date accepted: 19 November 1993) of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.4039/Ent126193-2Downloaded from https:/www.cambridge.org/core. 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GALL INDUCTION BY PEGOMYA CURTICORNIS (STEIN) (DIPTERA: ANTHOMYIIDAE) WITHIN THE ROOTS OF SPURGES EUPHORBIA VIRGATA WALDST. AND KIT. AND E. ESULA L. (EUPHORBIACEAE)

GALL INDUCTION BY PEGOMYA CURTICORNIS (STEIN) (DIPTERA: ANTHOMYIIDAE) WITHIN THE ROOTS OF SPURGES EUPHORBIA VIRGATA WALDST. AND KIT. AND E. ESULA L. (EUPHORBIACEAE)

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