Heterodera sacchari




Rev 10/20/2023

Sugarcane Cyst Nematode Classification Hosts
Morphology and Anatomy Life Cycle
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Heterodera sacchari Luc & Merny, 1963

One of the species of the genus that comprise the "Sacchari group" of Heterodera spp.  Species of the groupes have lemon-shaped cysts, are ambifenestrate, and have finger-like bullae projecting from the underbridge of the vulval cone (Tanha Maafi et al., 2007).

According to our present knowledge, H. sacchari is definitely present in tropical Africa, H. goldeni is found in the middle east and H. leuceilyma occurs in the subtropical south-east of North America. The species identity of H. sacchari records from other Asian countries, as well as from Trinidad, West Indies, may need confirmation (Tanha Maafi et al., 2007).  The record from Trinidad


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Morphology and Anatomy:

General Characteristics of the Sacchari Group of Heterodera

Cysts lemon-shaped
Bullae present, finger-like
Underbridge well-developed
J2 incisures 3
Monocotyledonous hosts

(Handoo and Subbotin, 2018; Subbotin et al., 2010,  Tanha Maafi et al., 2005, 2007)


Mature Females: White, lemon-shaped.

Lip region truncated, with 2 offset retrorse annules; weak cephalic sclerotization.

Weak stylet, 23-25 µm long, with small, rounded knobs.

Opening of the dorsal esophageal gland situated 4-5 µm behind the spear; procorpus cylindrical.

Heavy median bulb with strong valve; short isthmus.

Basal glandular part of the esophagus nearly spherical.

Nerve ring encircling the isthmus.

Excretory pore situated 165-175 µm from the anterior extremity.

Cysts: Mature cysts brown to dark brown, lemon-shaped with prominent vulval cone and neck of medium size.

Cuticle with a lace-like pattern; numerous perforations evenly but irregularly distributed on the inner layer. Young cysts with a thick subcrystalline layer.

Cyst cone ambifenestrate: fenestrae rather obscurely demarcated.

45-55 µm long and 35-45 µm wide.

Vulval slit 50-52 µm long, as long as vulval bridge; underbridge strongly developed with finger-like bullae: length = 100-150 µm; width 50-70 µm; depth = 22-28 µm (Mulvey, 1972); few peripheral bullae.

Males: Rare. Body from nearly straight to C-shaped when relaxed by heat, twisted at the posterior end, cylindrical, slightly tapering anteriorly, terminus rounded.

Cuticle distinctly annulated; annules 2.5 µm in the middle of body; lateral field faintly marked by 3 longitudinal lines (=incisures) irregularly crossed by annules, occupying 1/5 of the corresponding body width.

Lip region dome-shaped, with 4-5 (exceptionally 6) annules, often anastomosed, without longitudinal striations; cephalic framework heavily sclerotized, with outer margin conspicuously marked; cephalids difficult to see, situated 2 and 6-7 annules behind labial

Spear stong, anterior and posterior parts of the same length; knobs rounded posteriorly and with flat, sloping anterior surface.

Dorsal esophageal gland opening situated 4 µm behind the spear.

Median esophageal bulb ovoid.

Dorsal gland short and wide with a large nucleus, anterior to the subventral glands which are elongated and narrow, and each with a small nucleus.

Nerve ring strongly marked, encircling the esophagus just behind the median bulb.

Excretory pore situated 137-150 µm from the anterior end. Hemizonid flat, 7-15 µm anterior to the excretory pore and extending over one annule. Hemizonion not seen.

Testis single; spermatozoa 4-5 µm in diameter.

Spicules curved, notched at the tip; specimens with abnormal atrophied spicules frequent (up to 20% of specimens); exceptionally, three spicules were observed. Spicular sheath present. Lamellate gubernaculum.

Tail 1/4 to 1/3 of the cloacal body diameter. Phasmids not observed.

Second-stage Juveniles: Body straight or slightly ventrally curved when heat-relaxed, slightly tapering at the anterior end, more attenuated at the posterior end.

Cuticle annulated; annules 1.7 µm wide in the middle of body; lateral field composed of 3 longitudinal lines, not crossed by annules except in the fore-part and at the level of the phasmids.

Lip region dome-shaped, with 3 annules; cephalic framework heavily sclerotized.

Stylet stong, anterior part slightly shorter than the posterior; knobs well developed, rounded posteriorly, concave anteriorly.

Dorsal esophageal gland opening situated 5-8 µm behind the spear

Median esophageal bulb ovoid, with strong valve.

Dorsal gland broad,anterior to the subventrals, with a large nucleus. Subventral glands not completely filling the body cavity, each with a small nucleus.

Nerve ring well defined, situated immediately behind the median bulb.

Excretory pore104-130 µm from the anterior end. Hemizonid immediately in from of the excretory pore, lenticular, extending over 2 annules.

Genital primordium with 2 nuclei, located at the mid-body.

Tail elongated, conical, with pointed terminus. Phasmids pore-like, situated 45-47 µm from the posterior end.

Reported median body size for this species (Length mm; width micrometers; weight micrograms) - Click:

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The sugarcane cyst nematode was originally reported from sugarcane, Saccharum officinale L., in Congo-Brazzaville (Luc & Merny, 1963) and was later found parasitising roots of rice, Oryza sativa L.  With these two plants as the primary hosts, H. sacchari is usually found in warmer regions of the world.

Several West African countries including Congo-Brazzaville, Nigeria, Cote d’Ivoire, Liberia, Benin, Burkina Faso, Senegal, Cameroon, Gambia, Ghana, Guinea, Togo (Luc & Merny, 1963; Jerath, 1968; Merny, 1970; Luc, 1986; Nobbs et al., 1992; Coyne et al., 1996; Evans & Rowe, 1998; Tanha Maafi et al,., 2007).

Also, India (Swarup et al., 1964) and Pakistan. (Maqbool, 1981). It has also been reported from Trinidad (Singh, 1974) but that report has not been confirmed by molecular studies (Tanha Maafi et al., 2007).  The record from Trinidad (Singh, 1974) appears to have been incorrectly reported as being from Jamaica by Luc (1974).


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Economic Importance:

Considered a potentially important pest of sugarcane and rice and, accordingly, is on the quarantine list of several countries.

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Feeding site establishment and development typical of genus.

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Hosts of the Sacchari group of Heterodera spp. are generally moncotyledons.. Sugarcane (Saccharum officinale L.), Rice (Oryza sativa L.).

 Also, several wild grasses sour grass (Paspalum conjugatum), bermudagrass (Cynodon dactylon), carpet grass (Axonopus compressus), Mariscus umbellatus, Brachiaria brizantha and Elesune indica, were considered to be hosts by Odihirin (1975) in Southern Nigeria, namely However, information on the morphology of the populations isolated from the grasses is lacking and the host status of these plants to H. sacchari needs to be confirmed (Tanha Maafi et al., 2007).


For an extensive host range list for this species, click



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Life Cycle:

Ecophysiological Parameters:

For Ecophysiological Parameters for this species, click If species level data are not available, click for genus level parameters

Parthenogenetic reproduction. Males rare, many have degenerate copulatory apparatus. 3n=27, triploid.

Juveniles hatch readily in water.

In Nigeria, Jerath (1968) observed that H. sacchari was more abundant in sandy soils than in clay soils, and that cysts were more numerous in the soil during the dry season.

On pluronic gel medium at 25C, the life cycle was completed in 7-9 weeks on rice (Oryza sativa, cv Nipponbare). After infection, juveniles developed and reached the reproducing adult female stage in 25 days. All females had produced eggs by 35 days.  Female bodies began to turn into dark brown cysts at 49 days.  Hatching of huveniles from eggs could be stimukated by  3mM ZnCl2 but not by rice root exudate (Pokhare et al., 2019).



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Sugarcane plants become stunted, measuring 1-1.5 m in height and remaining thin as compared with healthy plants which are 3-4 m high.

Secondary roots are fewer in damaged plants (Jerath, 1968).

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Little work has been done on control.

Jerath (1968) indicated that molasses applied to sugarcane fields decreased the number of cysts recovered to one-fourth the number found in untreated soils.

Host Plant Resistance, Non-hosts and Crop Rotation alternatives:

For plants reported to have some level of resistance to this species, click


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BABATOLA, J.O. (1983). Pathogenicity of Heterodera sacchari on rice. Nematologia Mediterranea 11, 21-25.

BRIDGE, J., LUC, M. & PLOWRIGHT, R.A. (1990). Nematode parasites of rice. In: Luc, M., Sikora, R.A. & Bridge, J.(Eds). Plant parasitic nematodes in tropical and subtropical agriculture. Wallingford, UK, CABI Publishing, pp. 69-108.

COYNE, D.I., PLOWRIGHT, R.A. & FOFANA, I. (1996). Preliminary investigations of nematodes associated with rice in Guinea, Benin and Togo. Afro-Asian Journal of Nematology 6, 70-73.

Pokhare, S.S., Samer S. Habash, John T. Jones, Abdelnaser Elashry and Florian M.W. Grundler. 2019. In vitro life cycle of Heterodera sacchari on Pluronic gel. Nematology 21:573-579.

DI EDWARDO, A.A. & PERRY, V.G. (1964). Heterodera leuceilyma n. sp. (Nemata: Heteroderidae), a severe pathogen of St Augustine grass in Florida. Agricultural Experiment Stations, University of Florida, Gainesville, USA, Bulletin 687, 35 pp.

HANDOO, Z.A. & IBRAHIM, I.K.A. (2002). Description and SEM observations of a new species of cyst nematode Heterodera goldeni (Nematoda: Heteroderidae) attacking Panicum coloratum in Egypt. Journal of Nematology 34, 312-318.

Handoo, Z.A., Subbotin, S.A. 2018. Taxonomy, identification and principal species. Chapter 15 in Perry, R.N. Moens, M., and Jones, J.T.(eds)  Cyst Nematodes. CAB International.

JERATH, M.L. (1968). Heterodera sacchari, a cyst nematode pest of sugar cane new to Nigeria. Plant Disease Reporter 52, 237.

LUC, M. (1974). Heterodera sacchari. CIH Descriptions of plant-parasitic nematodes, Set 4, No. 48. St Albans, UK, Commonwealth Institute of Helminthology, 3 pp.

LUC, M. (1986). Cyst nematodes in equatorial and hot tropical regions. In: Lamberti, F. & Taylor, C.E. (Eds). Cyst nematodes. New York & London, Plenum Press, pp. 355-372. 496 NematologyMorphological and molecular studies on Heterodera

LUC, M. & MERNY, G. (1963). Heterodera sacchari n. sp. (Nematoda: Tylenchoidea) parasite de la canne à sucre au Congo-Brazzaville. Nematologica 9, 31-37.

 MULVEY, R.H. (1972). Identification of Heterodera cysts by terminal and cone top structures. Canadian Journal of Zoology 50, 1277-1292.

NETSCHER, C. (1969). L’ovogénèse et la reproduction chez Heterodera oryzae et H. sacchari (Nematoda: Heteroderidae). Nematologica 15, 10-14.

NETSCHER, C., LUC, M. & MERNY, G. (1969). Description du mâle d’Heterodera sacchari Luc & Merny, 1963. Nematologica 15, 156-157.

NOBBS, J.M., IBRAHIM, S.K. & ROWE, J. (1992). A morphological and biochemical comparison of the four cyst nematodes species, Heterodera elachista, H. oryzicola, H. oryzae, and H. sacchari (Nematoda: Heteroderidae) known to attack rice. Fundamental and Applied Nematology 15, 551-562.

ODIHIRIN, R.A. (1975). Occurrence of Heterodera cysts nematode (Nematoda: Heteroderidae) on wild grasses in southern Nigeria. Occasional Publications of Nigerian Society of Plant Protection 1, 24.

SHAHINA, F. & MAQBOOL, M.A. (1995). Cyst nematodes of Pakistan (Heteroderidae). Karachi, Pakistan, National Nematological Research Centre, University of Karachi, 155 pp.

SINGH, N.D. (1974). Nematode pests of sugarcane new to Trinidad. Plant Disease Reporter 58, 122.

STONE, A.R. (1975). Head morphology of second stage juveniles of some Heteroderinae (Nematoda: Tylenchoidea). Nematologica 21, 81-88.

STONE, A.R. (1979). Co-evolution of nematodes and plants. In: Hedberg, I. (Ed.). Parasites as plant taxonomists. Uppsala, Sweden, Acta Universitatis Upsaliensis, Symbolae Botanicae Upsaliensis 22, part 4, pp. 46-61.

STURHAN, D. (1998). Notes on the taxonomy and phylogeny of Heteroderidae parasitising Gramineae. Nematologica  44, 585-586.

Subbotin, S,A., Mundo-Ocampo, M., Baldwin, J.G. 2010. Systematics of Cyst Nematodes (Nematode: Heteroderinae). Nematology Monographs and Perspectives Volume 8B, D.J. Hunt and R.N. Perry (eds) Brill, Leiden, The Netherlands 512p

SUBBOTIN, S.A., WAEYENBERGE, L. & MOENS, M. (2000).Identification of cyst forming nematodes of the genus Heterodera (Nematoda: Heteroderidae) based on the ribosomal DNA-RFLP. Nematology 2, 153-164.

SWARUP, G., PRASAD, S.K. & RASKI, D.J. (1964). Some Heterodera species from India. Plant Disease Reporter 48, 235.

TANHA MAAFI, Z. (2002). Morphological and molecular identification of cyst-forming nematodes from Iran. Ph.D. Thesis, Department of Biology, Faculty of Sciences, University of Gent, Belgium, 237 pp.

TANHA MAAFI, Z, SUBBOTIN, S.A. & MOENS, M. (2003). Molecular identification of cyst-forming nematodes (Heteroderidae) from Iran and a phylogeny based on ITS-rDNA sequences. Nematology 5

TANHA MAAFI, Z.T., STURHAN, D., HANDOO, Z., MOR, M., MOENS, M. & SUBBOTIN, S.A. (2005). Morphological and molecular characterisation of Heterodera sacchari Luc & Merny, 1963 and H. goldeni Handoo & Ibrahim, 2002 (Nematoda: Heteroderidae). Russian Journal of Nematology 13, 152-153., 99-111.

TANHA MAAFI, Z., STURHAN, D., HANDOO, Z., MOR, M., MOENS , M. SUBBOTIN, S.A.. 2007. Morphological and molecular studies on Heterodera sacchari, H. goldeni and H. leuceilyma (Nematoda: Heteroderidae). Nematology 9: 483-497.

VOVLAS, N., LAMBERTI, F. & TUOPAY, D.K. (1986). Observations on the morphology and histopathology of Heterodera sacchari attacking rice in Liberia. FAO Plant Protection Bulletin 34, 153-156.

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Copyright © 1999 by Howard Ferris.
Revised: October 20, 2023.