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An Illustrated Key to the Cyst-Forming Genera and Species of Heteroderidae in the Western Hemisphere with Species Morphometrics and Distribution ~ R

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An Illustrated Key to the Cyst-Forming Genera and Species of Heteroderidae in the Western Hemisphere with Species Morphometrics and Distribution ~ R JOURNAL OF NEMATOLOGY VOLUME 15 JANUARY 1983 NUMBER 1 An Illustrated Key to the Cyst-forming Genera and Species of Heteroderidae in the Western Hemisphere with Species Morphometrics and Distribution ~ R. H. MULVEYz and A. MORGAN GOLDENa Abstract: Diagnoses of the cyst-forming genera of Hcteroderidae (viz., Heterodera, Sarisodera, Globodera, Punctodera, Ca.~todera, and Dolichodera) and distribution and morphometrics of the 34 known cyst species in the Western Hmatisphere are presented along with an illustrated key for the identification of these genera and species. The key is based mainly tm cysts and larvae, attd important morphological and diagnostic features are extensively shown by LM and SEM illustrations. The genus Bidera is placed as a new synonym under the genus Heterodera. Key words: cyst nematodes, taxonomy, identification. Journal of Nematology 15(1):1-59. 1983. Early in 1981 the corn cyst nematode details essential in surveys and related work. (H. zeae Koshy, Swarup & Sethi, 1971 [30]) lnchlded also are generic diagnoses and was found in three corn fields in Kent distribution and morphometric data for County, Maryland, the first known occur- all of tile known species in the Western rence of this cyst species in the United Hemisphere. The genus Bidera is synonym- States (47). Subsequent limited surveys ized with Heterodera. showed the nematode to be on several near- by farms, and preliminary tests indicated MATERIALS AND METHODS field corn, sweet corn, and barley to be Living specimens of several species, in- hosts. Because of the potential threat to cluding Heterodera g[ycines and H. tri[olii, corn production in Maryland and in the were obtained from isolated cultures in the United States posed by this cyst nematode, greenhouse or growtlt chambers at Belts- extensive research--including surveys, host ville, Maryland, while fresh specimens of range and pathogenicity studies, and possi- H. zeae were provided by Dr. L. Krusberg, ble control measures--was planned and ini- University of Maryland. College Park. The tiated. procedures used for weparing these speci- As part ot this overall research effort, mens were essentially the same as those used our report here gives an illustrated key to by Golden and Birchfield (16). Fixed speci- the 6 genera and 34 cyst-forming species mens of the various other species were ob- known to occur in the Western Hemi- tained from the USDA Nematode Collec- sphere. The key is based on both cysts and tion at Behsville. Photomicrographs of larwd characters and provides diagnostic cysts, vulva cones, females, and larvae were made with an automatic 35 mm camera Received for publication 19 April 1982. attached to a compound microscope having tScientific article number A-3308, contribution number 6380 of the Maryland Agricultnral Experiment Station. an interference contrast system; those of in- This research was done in the laboratory of the second fected roots, whole females, and cysts were author and supported in part through a USDA Coopera- tive Agreement with the University of Maryland, Col.lege made with a 8.3 × 10.8 cm sheet-film Park. Thanks are given especially to Dr. Lorin Krusberg, camera attached to a dissecting microscope. Department of Botany, for his assistance and cooperation. Appreciation for technical assistance is extended to In making the scanning electron micro- Donna M. S. Ellington, Support Scientist, and Paula Crow- ley, I.aboratory Technician (in laboratory of second author), graphs, living specimens were fixed In 3% and to Bruce Ingber, Laboratory Technician, Electron glutaraldehyde solution 0.05 M phosphate Microscopy Laboratory, USDA ARS, Beltsville, MD 20705, for SEM technical assistance. We also thank Dr. Roger V. buffer (pH 6.8), dehydrated in a graded Anderson, Biosystematics Research Institute, Canada Agri- series of ethanol, critical-point dried from cuhure, for some of the photomicrographs and SEM micrographs used in tills paper. liquid CO 2, sputter coated with a 20-30 ~Former Chief (retired), Nematology Section, Biosys. tematics Research Institute, Research Branch, Agriculture /tm layer of gold-palladium, and examined Canada, Ottawa, Canada. witll a scanning electron microscope (SEM). 3Nematologist, Nematology Laboratory. Plant Protec- tion Institute, USDA ARS, Beltsvnle, MD 20705. Some specimens were prepared previously The JOURNAL OF NEMATOLOGY for October (14:427-603)was issued 8 December 1982. .... I • • 2 Journal o] Nematology, Volume 15, No. 1, January I983 in Canada by the senior author using the Punctodera chalcoensis Stone, Sosa same methods as Mulvey (39,40). Some Moss 8¢ Mulvey, 1976 distribution data were supplied by the (Fig. 22) USDA Nematode Collection at Beltsville, Described and illustrated by Stone et al. Maryland, while much of the data on distri- (55). bution were based on published reports (5, Type host aud: locality: Maize (Zea mays 18, 35, 39). L.). Chalco area, Valley of Mexico, Mexico SYSTEMATICS State, Mexico. (Figs. 1-164) Distribution: Only Tlaxcala and Mexico States, Mexico. Family: Heteroderidae (Filipjev g: Schuurmans Stekhoven, 1941) Skarbilovich, Punctodera matadorensis Mulvey 1947. & Stone, 1976 Subfamily: Heteroderinae Skarbilovich, (Figs. 23-27) 1947. Diagnosis: After Mulvey g: Stone, 1976 Described and illustrated by Mulvey g~ 02). Stone (42). Femate: Cyst formed after death. Cuticle Type host and locality: Permanent non-annulated. Anus dorsal or on the in- grass. Matador Ranch, Matador, Saskatche- side of the dorsal vulval lip. Vulva terminal. wan, Canada. Second-stage juvenile: Labial disc pres- Distribution: Only Saskatchewan, Can- ent but with low profile and only seen ada. clearly under scanning electron microscope. Cuticle of labial region of even thickness. Punctodera punctata (Thorne, 1928) Esophageal glands fill body width. Mulvey 8~ Stone, 1976 Male: Length up to 1.7 ~'m. Labial disc (Figs. 19-21) present but with low profile, or apparently Described by Thorne (59). so. Tail present or absent. Type host and locality: Wheat (Triti- Type genus: Heterodera Schmidt, 1871. cure aestivum L.) and grasses. Humboldt (See page 50 for generic diagnosis.) area, Saskatchewan, Canada. Distribution: Western Hemisphere-- Punctodera Mulvey 8¢ Stone 1976 Saskatchewan, Labrador, Newfoundland, (Tables 1-4) Sable Island (off Nova Scotia), Manitoba, Diagnosis: After Mulvey & Stone (42). Ontario, Quebec, and British Columbia in Mature female and cyst: Without poste- Canada; Michigan, South Dakota, Texas, rior protuberance; spherical, ovoid, or pear and Arkansas in USA. World--Austria, Bel- shaped. Cuticle with lacelike pattern of gium, England, Holland, Ireland, Norway, ridges and subsurface pattern of puncta- Poland, USSR, West Germany. tions. Vulval slit very short, underbridge Discussion: Thorne's P. punctata speci- absent. Bullae present or absent. Perineal mens (slide 3A) in the USDA Nematode tubercles absent. Circumfenestrate fenestra Collection at Beltsville, Maryland, of white surrounding vulva and circumfenestrate females and cysts provided new informa- fenestra of similar size surrounding anus. tion on vulva and anal fenestra, length of Anus offset toward ventral margin of anal vulva slit, and position of anus. Other in- fenestra. formation concerning larvae (from Second-stage juveniles: Stylet less than Thorne's slide 3B) was taken fi'om Mulvey 30 ixm. Esophageal glands fill body cavity. and Stone (42). Phasmids without lenslike structure in muscle layer. Genus DoIichodera Mulvey & Ebsary, 1980 Male: To 1.5 mm long. Labial disc pres- Diagnosis: After Mulvey ~: Ebsary (44). ent but with low profile. Tail present, less Mature female and cyst: Stylet well de- than half body width. veloped. Body elongate-ovoid, without pos- Type species: Punctodera punctata terior protuberance. Cuticle thin with inter- (Thorne, 1928) Mulvey g: Stone, 1976. rupted, irregular striae, and no regular Heteroderidae Taxonomy, Morphology, SEM: Mulvey and Golden 3 annules. Crystalline layer absent. Vulva slit However, sufficient specimens were not not observed. Underbridge absent, bullae available at that time tot adequate descrip- present. Perineal tubercles absent. Vulva tion. circumtenestrate, anus lacking fenestra. Second-stage la~'oae: Styler 22-24 /xm. l-Ieterodera betulae Hirschmann Esophageal glands occupying entire body & Riggs, 1969 cavity. Phasmids small. Tail 95-120 /zrn (Figs. 65-71; Tables 5-8) long, hyaline area 30% o[ tail length. Type species: DoIichodera fluvialis Mul- Thoroughly described and illustrated by vey 8c Ebsary, 1980. Hirschmann and Riggs (24). Mulvey (39) assigned this species to the cacti group based Dolichodera fluvialis Mulvey 8c Ebsary, 1980 oil having a very short vulva slit. The (Figs. 28-35; Tables 5-8) markedly different morphology of H. betulae, as documented by Golden and Described and illustrated by Mulvey and Raski (18) and also illustrated herein, Ebsary (44). places this species apart from all described Type host and locality: Aquatic grass, Heterodera. Theretore, we have not as- (Spartina pectinata) growing on the bottom signed this species to the genus Cactodera of the Ottawa River (females and cysts or to any of the present groups within the found on the roots by B. A. Ebsary in 1981) genus Heterodera, pending further action near Deschenes, Quebec, Canada. on this by one o[ us (AMG). Distribution: Quebec, Canada. Type host and locality: River birch roots (Betula nigra L.). South bank of Middle Genus Sarisodera Wouts 8¢ Sher, 1971 Fork, White River, 1.75 miles upstream Diagnosis: Emended trom Mulvey
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