And Cactodera Cacti with a Review and Key to the Genus Cactodera 1
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Journal of Nematology 22(4):457-480. 1990. © The Society of Nematologists 1990. Descriptions and Comparative Morphology of Cactodera milleri n. sp. (Nematoda: Heteroderidae) and Cactodera cacti with a Review and Key to the Genus Cactodera 1 L. S. O. GRANEY AND G, W. BIRD 2 Abstract: A new species, Cactodera milteri n. sp., is described and illustrated from specimens obtained from roots of common lambsquarter, Chenopodium album L., from Mattawan, Michigan. Cactodera milleri can be differentiated from other Cactodera species by the presence of punctated egg shells and a second-stage juvenile (]2) styler length averaging 21.8 t~m as measured from freshly killed specimens in water mounts. Thirty-four plant species, including 11 weed species, 18 agronomic crop species, and 5 cactus species were tested as potential hosts of C. milleri. The new species reproduced only on Chenopodium album, C. amaranticolor Cofte. Reyn., and C. quinoa Willd. Cactodera cacti (Filipjev & Schuurmans Stekhoven, 1941) Krall & Krall, 1978, a morphologically similar species, is reexamined. A description of the female and additional morphometric and morphologic data of cysts, males, J2, and eggs are provided for several populations of C. cacti. A review of the morpho- metrics of all species of Cactodera and a taxonomic key to the seven species are presented. Key words: Cactodera cacti, Cactodera mitleri n. sp., cactus cyst nematode, Chenopodium album, Che- nopodium amaranticolor, Chenopodium quinoa, common lambsquarter, comparative morphology, cyst nematode, host range, new species, taxonomy. In the fall of 1982, a nematode popu- from Mattawan, Michigan, is described as lation with lemon-shaped cysts was isolated Cactodera milled n. sp. In addition, the re- from soil samples collected from a former productive ability of C. milleri was tested asparagus field in Mattawan, Michigan (4). on 34 plant species, including 11 weed The host of the cysts could not be identi- species, 18 agronomic crop species, and 5 fied at the time of collection because the cactus species: soil samples were taken when the field had A review of previously published mor- just been cultivated and planted to rye. phometrics of cysts, males, J2, and eggs of Preliminary morphological examination of other Cactodera species and additional mor- the cysts, eggs, and second-stage juveniles phometric and morphologic data of fe- (J2) indicated that the population repre- males, cysts, males, J2, and eggs of C. cacti sented a species of Cactodera Krall & Krall, are also presented. 1978, morphologically distinct from all A revised key to the genus Cactodera is species of the genus except C. cacti (Filipjev provided. & Schuurmans-Stekhoven, 1941) Krall & Krall, 1978. The range of measurements MATERIALS AND METHODS of C. cacti (1,2,6,7,9-12,16) overlapped with those of the unidentified Cactodera popu- Field host identification and host range lation. However, an intensive reexamina- The field host of C. miUeri was deter- tion of the morphology and morphomet- mined in the greenhouse by placing in- rics of C. cacti revealed significant fested field soil from the original site in differences, and the Cactodera population flats and periodically observing the roots of the germinated plants for the presence of white females. In addition, 20-cm-d clay Received for publication 3 November 1989. pots were filled with field soil and seeded Michigan Agricultural Experiment Station Journal Arti- with asparagus, Asparagus o~cinalis L. cv. cle 11158. Former Senior Research and Extension Specialist and Mary Washington. As weed seedlings Professor, Department of Entomology, Michigan State Uni- emerged, they were removed. After 3 versity, East Lansing, MI 48824, The authors thank Peter Garrington for the nematode months, the roots of the asparagus plants illustrations. were examined for females. 457 458 Journal of Nematology, Volume 22, No. 4, October 1990 TABLE 1. Plant species tested as hosts of Cactodera miUeri n. sp. Nematode Plant species Common name reproduction Weeds Digitaria sanguinalis (L.) Scop. Large crabgrass Amaranthus retroflexus L. Redroot pigweed Rumex crispus L. Curly dock Chenopodium album L. Lambsquarter + Chenopodium amaranticolor Cofte. Reyn. Lambsquarter + Chenopodium quinoa Willd. Quinoa + Solanum dulcamara L.t Bitter nightshade Polygonum persicaria L. Ladysthumb Polygonum convolvulus L. Wild buckwheat Polygonum pensylvanicum L. Pennsylvania smartweed SteUaria media (L.) Vill.:~ Common chickweed Agronomic crops Phaseolus vulgaris L. cv. Seafarer Dry bean Lycopersicum esculentum Mill. cv. Rutgers Tomato B Glycine max (L.) Merr. cv. Hark Soybean D Zea mats L. cv. Mol7HT Corn Apium graveolens L. cv. F1683 Celery m Mentha arvensis (L.) Hill cv. Black Mitchumz~ Mint w Brassica oleracea L. cv. Golden Acre Cabbage Asparagus ofl~cinalis L. cv. Jersey Giant Asparagus Beta vulgaris L. cv. H20t Sugarbeet m Triticum aestivum L. cv. Hyslopt Wheat Daucus carota L. cv. Gold Pak'{" Carrot Allium cepa L. cv. Krummery Specialt Onion m Beta vulgaris L. cv. Early Wonder Green Topt Table beet Medicago sativum L. cv. Vernal:]: Alfalfa Solanum tuberosum L. cv. Superior§ Potato Trifolium pratense L.:~ Red clover m Lotus corniculatus L.~ Birdsfoot trefoil Vitis vinifera L. cv. Baco Noir§ Grape Cacti Opuntia rufida§ Cinnamon cactus m Opuntia vulgaris Miller§ Irish mittens m Mammillaria elongata DeCandolle§ Golden stars Lobivia silvestrii (Spegazzini) Rowley§ Peanut cactus Schlumbergera truncata (Hayworth) Moran§ Thanksgiving cactus m Seedlings transplanted except as otherwise noted. "{" Seeded directly onto infested soil and thinned to one plant per pot. :1: Seeded directly onto infested soil with no thinning. § Rooted cutting or tuber piece, Thirty-four plant species (Table 1) to be teurized sandy loam soil and a hole, 2.5 cm evaluated as hosts ofC. milleri were directly d x 2.5 cm deep, was made in the center seeded or transplanted as seedlings, rooted of each pot. Twenty-five brown cysts were cuttings, or tuber pieces. Nematode inoc- placed in the hole with water, and a seed- ulum (brown cysts) was obtained from stock ling, rooted cutting, or tuber piece was cultures established on Chenopodium album placed in the hole and secured with soil. L., the field host, and maintained by peri- When plant species were direct seeded, the odic subculturing in the greenhouse at 25 hole containing the cysts was filled with soil _+ 5 C. Four replications of each plant and seeds were spread evenly over the sur- species were prepared as follows: 10-cm-d face and covered lightly with additional soil. clay pots were filled with moistened pas- After 3 months, the soil and root systems Cactodera milleri n. sp. and C. cacti: Graney, Bird 459 were washed vigorously with water to re- served by Adam in 1932, was obtained from move newly formed females and cysts. The the Nematode Collection, Section of Re- soil and root washings were poured onto a cent Invertebrates, Institut Royal des Sci- 600-urn-pore sieve over a 250-tzm-pore ences, Naturelles de Belgique, Bruxelles, sieve. The material from the 250-~m-pore Belgium (I.G. 14.530). sieve was examined microscopically, and The populations of C. cacti used in this the cysts and females were counted. The study are designated as follows: MI (Mich- presence of newly formed females, white igan, USA); FL (Florida, USA); LO (Lot- to yellow in color, or more than 25 brown tum, The Netherlands); GA (Georgia, cysts indicated reproduction of the nema- USA); IT (Italy); WA (Wageningen, The tode. If 25 or fewer brown cysts were re- Netherlands); and TY (type material, covered, the plant was not considered to Maartensdijk, The Netherlands). be a host. Specimen preparation Research populations Light microscopy (LM): Holotype and Nematode specimens of C. milleri were paratype females of C. miUeri and females obtained from greenhouse stock cultures of the MI population of C. cacti were established and maintained by periodic mounted in water under a coverslip on a subculturing on Chenopodium album. thin water agar plate for measurement of Populations of C. cacti were obtained body dimensions. The anterior ends of the from the Department of Horticulture trop- holotype and paratype females of C. milleri ical greenhouse, Michigan State University and females of the MI and LO populations (MSU); University of Florida at Gaines- of C. cacti were prepared by fixing intact ville, courtesy of Dr. R. P. Esser; and Lot- white females in 3% HCHO for 5 minutes tum, The Netherlands, courtesy of Dr. P. and then excising the head and mounting Maas. Stock cultures of the Michigan and it in lactophenol. Cysts of C. milleri and the Lottum populations of C. cacti were main- MI population of C. cacti were prepared for tained on Cereus peruvianus (L.) Miller and measurement of body dimensions as de- Schlumbergera truncata (Hayworth) Moran, scribed for females. Cone tops of C. milleri but the Florida population did not repro- (holotype female and paratype cysts) and duce in the MSU greenhouse. Viable spec- C. cacti (females and cysts of MI and cysts imens of eggs and J2 of the Florida pop- of TY populations) were prepared by the ulation were examined before the rearing methods of Mulvey (10). The female vul- attempt and their data are included in the va-anus distance was measured from cone present study. tops that had been cut to allow flattening In addition to living specimens, perma- of the vulval cone structure and mounted nent slides of eggs and J2 in glycerin of C. in lactophenol, except in the case of the cacti from Italy (slides G-4637, G-4638) and holotype female which remained intact. Georgia, USA (slide G-4664) were ob- The allotype male of C. miUeri was heat tained from the USDA Nematode Collec- relaxed, mounted in water, measured, then tion Nematology Lab, Beltsville, Mary- carefully removed from the water mount, land; and permanent slides of J2 and males processed to glycerin with the Seinhorst in glycerin (slide 5429) were obtained from method (15), and remeasured.